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SMITHSONIAN
MISCELLANEOUS COLLECTIONS

VOL. 116

ZaXtSON ay

FEB 18 1952
LIBRAR Ls

“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—JAMES SMITHSON

(Pustication 4061)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
il" 1952

The Lord Baltimore Press

BALTIMORE, MD., U. S» As

ADVERTISEMENT

The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge,
all the publications issued directly by the Institution except the An-
nual Report and occasional publications of a special nature. As the
name of the series implies, its scope is not limited, and the volumes
thus far issued relate to nearly every branch of science. Papers in
the fields of biology, geology, anthropology, and astrophysics have
predominated.

A. WETMORE,
Secretary, Smithsonian Institution.

(ii)

TYAM ARP CRS

Sih adeie wiatiior wits: erorrsnll,2 wosaia iM pnts
viata tints al ganintivide 2 asitontier at) Docket aie
SrA uli hye vourting. ot plage ious enothe ae
ut @/. ality 1 wore A 4 creuhys Terman’ Darel ; i
memersitind iLh lGtey Satie? TO ee eS sai eon exits odf {9-3

mp xenpitl ‘riage bo he ond vise bed an oie facut)
pial sere riag. Lice enolonprtsltian, phew nes to

mow TAL of
gosiul*) ree nn? uM

CONTENTS

. Swoperass, R. E. Comparative studies on the jaws of mandibu-
late arthropods. 85 pp., 25 figs. Nov. 16, 1950. (Publ. 4018.)
. Gazin, C. Lewis, and Cotiins, R. Lee. Remains of land mam-
mals from the Miocene of the Chesapeake Bay region. 21 pp.,
7 figs. Oct. 12, 1950. (Publ. 4019.)

. THALBITZER, WILLIAM. Two runic stones, from Greenland and
Minnesota. 71 pp., 7 figs. Aug. 30, 1951. (Publ. 4021.)

. Assot,C.G. Precipitation and temperature in Washington, D. C.,
for 1950 and 1951. O pp., 2 figs. Mar. 1, 1951. (Publ. 4045.)
. RaAsett1, Franco. Middle Cambrian stratigraphy and faunas of
the Canadian Rocky Mountains. 277 pp., 34 pls., 5 figs.
Sept. 18, 1951. (Publ. 4046.)

. Mutr-Woop, HELEN M., and Cooper, G. ARTHUR. A new species
of the Jurassic brachiopod genus Septirhynchia. 6 pp., 2 pls.
June 5, 1951. (Publ. 4047.)

. CrarK, Austin H., and Crarx, Leita F. The butterflies of
Virginia. 239 pp., 31 pls., 1 fig. Dec. 30, 1951. (Publ. 4050.)

(v)

pt SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 1

}| COMPARATIVE STUDIES ON THE JAWS
|| OF MANDIBULATE ARTHROPODS

BY
R. E. SNODGRASS

Collaborator, Bureau of Entomology and Plant Quarantine
: U. S. Department of Agriculture

i

he
sania HET >}
Xs" . hs
NOV IL? tos

(Pustication 4018)

eC Crew OR WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
Hider - NOVEMBER 16, 1950

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 1

COMPARATIVE STUDIES ON THE JAWS
OF MANDIBULATE ARTHROPODS

BY
R. E. SNODGRASS

Collaborator, Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture

(Pustication 4018)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 16, 1950

The Lord Galtimore Press

BALTIMORE, MD., U. 8. A.

COMPARATIVE STUDIES, ON THE JAWS ‘OF

CONTENTS

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INTRODUCTION

MANDIBULATE ARTHROPODS
By R. E. SNODGRASS

Collaborator, Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture

The organs of feeding associated with the mouth that may be called
the jaws of an arthropod are usually mandibles, but not always, since
in some species one pair or both pairs of the maxillae may take over
the biting if not the chewing function. The mandibles, moreover,

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 1

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

are sometimes highly developed for purposes other than feeding, or,
on the other hand, they may be so reduced as to have little use of
any kind, and finally they may be suppressed entirely. However, the
principal subject of the following discussion will be the mandibles.

It may be taken for granted that the arthropod mandibles have been
evolved from a pair of legs, since all the postoral appendages of the
trilobites were fully developed as ambulatory limbs. There is some
difference of opinion, however, as to what part or parts of a general-
ized limb the mandible represents, though a reasonable answer should
be obtained by comparing a mandible of primitive type with an ordi-
nary ambulatory appendage of modern arthropods.

A typical arthropod leg (fig. 1A) consists of a basal segment
known as the coxa, or coxopodite (Cx), and of a segmented distal
shaft called the telopodite (Tipd). The usual movement of a locomo-
tor appendage on the body, whether for walking or swimming, is
anterior and posterior. Generally the coxa is specifically articulated
dorsally and ventrally on the body, or if not articulated, it is so at-
tached that the axis of rotation is essentially dorsoventral, though
actually it may be oblique at various angles. The dorsal articulation
(a), when present, is on the tergum of the body segment, or on a
laterodorsal plate termed the pleuron (P/) ; the primary ventral articu-
lation (b) is on the sternum. Departures from this type of structure
are clearly secondary and need not be considered here.

The body musculature of an appendage is appropriate to the move-
ments of the appendage on the body. If the limb turns anteriorly and
posteriorly, or approximately so, it is provided with promotor and
remotor muscles. The legs of most arthropods have both dorsal and
ventral muscles, though some have only dorsal muscles, and others
only ventral muscles. The dorsal muscles arise on the tergum of the
body segment; the ventral muscles usually have their origins on an
endosternal support of some kind, but since such structures are sec-
ondary formations it is reasonable to suppose that the ventral limb
muscles were first attached on the sternal surface of the segment. The
number of individual muscles for each appendage is variable, but
when dorsal and ventral muscles are both present, the functional
groups of fibers are four. In their action on the limb as a whole they
were probably in the first place dorsal and ventral promotors (fig. 1 A,
dpm, vpm), and dorsal and ventral remotors (drm, vrm) ; in their
action specifically on the coxa, they are anterior and posterior rotators.
With changes in the coxal articulation, or in the points of origin of
the muscles relative to the coxa, however, the same muscles may take
on quite different functions.

WO: LT JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 3

An arthropod mandible that most closely resembles the coxa of a
leg both in structure and musculature may be regarded as a generalized
mandible. Such a mandible occurs in many of the entomostracan and
in some of the malacostracan Crustacea, and in the Machilidae among
the insects. A mandible of this type (fig. 1 B, Md) is suspended ap-
proximately vertically from a single dorsal point of articulation (a)
on the tergum of its segment or on the head ; its lower end is produced
into a strong, usually toothed endite, or gnathal lobe (gnL); the
telopodite may be represented by a palpus (P/p), or it may be sup-
pressed. Inasmuch as a mandible of this kind, in order to be a func-
tional jaw, must be able to swing toward its fellow, the pendent mandi-
ble has no ventral point of articulation. Furthermore, since the mouth
(Mth) for practical purposes lies between the jaws, the mandibu-
lar sternum itself has been obliterated or reduced and displaced
posteriorly.

The musculature of a pair of primitive mandibles includes indi-
vidual anterior and posterior dorsal muscles for each jaw (fig. 1 B),
but in Crustacea with this type of mandible all the ventral fibers from
each jaw are attached medially on a transverse sheet or cylinder of
fibrous tissue suspended between the mandibles and forming a com-
mon ligament uniting the fibers from the opposite jaws. Since the
mandibular musculature becomes thus reduced to three functional
groups of fibers, it will be convenient to designate the groups simply
A, P, and V. The A and P muscles very clearly are the dorsal pro-
motor and dorsal remotor of the leg (A), but in their action on a
pendent mandible they become an anterior rotator (B, A) and a pos-
terior rotator (P). The single large mass of ventral fibers of the
mandible (V), representing the combined ventral muscles of the leg
coxa (A), constitute a particularly effective ventral adductor. The
mandibles being suspended on single dorsal points of articulation (a),
the adductor muscles, pulling against each other on the median liga-
ment, bring the gnathal lobes of the jaws strongly together beneath
the mouth. In mandibles of this type there is no apparent muscular
mechanism of abduction, the opening of the jaws evidently depends
on the elasticity of their basal connections.

With the further evolution of the mandibles the fibers of the three
primary muscles may become dissociated into distinct secondary
muscles, with diversified functions correlated with changes in the
mandibular mechanism. Functional names for the muscles, therefore,
cannot be consistently carried over from one type of mandible to
another. In a species of Collembola as many as 17 distinct muscles

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Fic. 1—Diagrams of the various types of arthropod mandibles, and their
apparent derivation from the coxae of a pair of leglike appendages.

A, section of a body segment bearing a pair of legs with generalized dorsal
and ventral musculature. B, a pair of mandibles of generalized structure,
pendent from single dorsal articulations, adducted by the united ventral muscles.
C, a generalized decapod mandible, with gnathal lobe in line with the length
of the jaw, doubly articulated, and rotating on a horizontal axis. D, the
astacuran mandible, same as the last but with a lateral apodemal lobe (Ap).
E, the anomuran-brachyuran type of mandible, the apodeme extended in line
with the body of the jaw. F, a protractile mandible. G, a doubly articulated
mandible with horizontal axis of rotation and gnathal lobe perpendicular to
axis (Isopoda, Amphipoda, Lepismatidae, most Pterygota). H, the diplopod-
symphylan type of mandible, gnathal lobe freely movable on the mandibular
base, and independently musculated.

SS ) e

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 5

with varied functions have been described attached on a single mandi-
ble, yet they can all be referred to the three original fiber groups.

It is, of course, not literally correct to say that a muscle changes its
function, since the only physical activity of muscle tissue is that of
contraction. What the muscle accomplishes usually depends on the
mechanism of its skeletal connections ; so, speaking of the mandibles,
the idea would be better expressed if we might say that the arthropods
have shown a great versatility in adapting their mandibles by mechani-
cal alterations to different kinds of movement operated by the same
muscles. Yet also, shifts in the muscle attachments may bring about
radical changes in the action of the muscles on the mandibles.

The intermandibular ligament on which the adductor muscles of
the jaws are attached is not a structure limited to Crustacea. In the
Diplopoda a large group of the adductor fibers from each mandible
merges into a common, transverse, cylindrical ligament (fig. 20 F),
and in the thysanuran insect Machilis, groups of fibers from opposite
jaws are similarly connected by a ligament through the base of the
hypopharynx (fig. 22 A, rV). The intermandibular ligament of Crus-
tacea is usually connected with a similar though smaller ligament
between the first maxillae and another between the second maxillae,
or the three ligaments may be united in a single sheet of tissue. A
composite structure of this kind is strongly developed in the gnathal
region of Anaspides (fig. 5 F), and is supported from the dorsum by
three pairs of suspensory branches (sl). An even more complex
structure of the intergnathal ligament is shown by Manton (1934,
fig. 17) to be present in Nebalia (E) and several other crustaceans.
On the other hand, in the copepod Calanus the large bundles of ventral
fibers of the mandibles (fig. 4 F, V) and also the maxillary muscles
are attached on an extremely slender median ligament running length-
wise over the nerve cord, and dividing anteriorly into a pair of finely
branched suspensory ligaments.

The term “ligament,” or “tendon,” seems hardly appropriate for
the intergnathal muscle-supporting structure when the latter takes the
form of a broad, elaborately developed, composite plate, which very
much resembles the so-called “endosternum” of Limulus and the
arachnids, from which the ventral muscles of the prosomatic ap-
pendages arise. Both structures are composed of a nonchitinous cellu-
lar and fibrillated tissue, and the fibrillae appear to be directly con-
tinued into those of the striated muscle fibers (figs. 2H, 5 A, 6H).
The suspensory branches in Crustacea (figs. 2 E, H; 8 D, E, sl) are
attached either directly on the dorsal body wall, or by groups of short
muscle fibers, as are those of Limulus.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

It has been shown by Manton (1928, 1934) that in the development
of Hemimysis and Nebalia a transverse ligament is formed from the
ectoderm of each ventral intersegmental fold throughout the length
of the trunk. On each fold there first appears a median ingrowth
between the nerve cords, and at each side a lateral ingrowth. The
lateral rudiments then extend mesally and unite with the median
rudiment, thus forming an arch or bar over the nerve cords. The
lateral connections with the ectoderm are later severed, and the bar
sinks backward into the segment behind. The antenno-mandibular
bar comes into contact with the mandibular mesoderm, which grows
along it from each side and becomes differentiated into the fibers of
the adductor muscles, while the bar itself transforms into the sup-
porting tendon. In the same manner are formed the maxillulary and
maxillary tendons, and, according to Manton, from the last is devel-
oped in Nebalia the tendon of the adductor muscle of the shell. The
mandibular, maxillulary, and maxillary tendons finally become inter-
connected, resulting in the formation of the complex intergnathal
ligament of the adult (fig. 5 E).

Embryonic phenomena can seldom be translated literally into evo-
lutionary history. According to the above account by Manton, the
embryonic adductor muscles of the gnathal appendages wait for the
formation of the ligament before they become functional; in evolu-
tionary development the muscles must have been functional from the
beginning. We may suppose, therefore, that the ventral muscles of
the appendages were first attached on the ventral body wall laterad
of the nerve cords, and that the ectoderm then formed a bridge over —
the nerve cords in the manner described, carrying the muscles with it.
In the Arachnida, according to Purcell (1909), the segmental groups
of embryonic muscle cells are at first attached on the ventral inter-
segmental folds, but the areas of contact soon become marked by the
appearance of the intermuscular tendons that will form the endosterna
(“entochondrites”). The tendons, however, are said by Purcell to
be a product of the fusion and metamorphosis of the muscle cells
themselves where the latter come into contact with one another and
with the epidermis. Purcell thus agrees with Schimkewitsch (1895),
who traces the development of the endosternum in spiders from trans-
formed muscle tissue.

Finally, it may be noted that in the Scutigeromorpha among the
chilopods the adductor muscles of the mandibles and both pairs of
maxillae arise from a plate of tissue (fig. 18D) much like that of
the intergnathal ligament of Crustacea and the endosternum of Limu-
lus and Arachnida, which Fahlander (1938) claims to be a non-

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 7

chitinous endoskeletal substance derived from the inner surface of
the epidermis. Furthermore, in each body segment of the Scutigero-
morpha the ventral muscles are attached on a cross bar in the posterior
part of the segment, which structures are suggestive of the segmental
endosternal plates in the opisthosoma of Limulus.

All these “ligamentous” or “endosternal,” nonchitinous muscle-
bearing tissues of the arthropods appear to be related or analogous
formations, but their origins and finer structure should be more ex-
actly studied, and their chemical composition determined by modern
technique. For the present we can simply accept them as anatomical
facts.

In the higher Crustacea the intergnathal ligaments become sup-
ported on ventral cuticular apodemes, and are reduced to thin, fascia-
like membranes, or they practically disappear, while the muscles are
taken over directly by the apodemes. The same transfer of the ventral
muscles to apodemes is seen in the chilopods; in Scutigera the liga-
mentous muscle-bearing plate of the head is supported on a pair of
apodemes, in other groups the ligament is reduced or absent and the
muscles in part or entirety go over to the apodemes. In the diplopods
and the insect Machilis the ligament persists between one group of
mandibular adductor fibers, but the other fibers take their origins
directly from head apodemes, and in the rest of the insects all the
ventral muscles of the gnathal appendages are attached on the apo-
demal tentorium of the head. That the transfer of the muscles to
cuticular apodemes is secondary is shown by the fact that in the dif-
ferent arthropod groups the apodemes may have quite different ori-
gins, and are certainly not homologous structures. It is only among
the holometabolous insects that the original ventral muscles of the
mandibles are themselves suppressed.

The body of a generalized mandible (corpus mandibulae) is broadly
attached by its mesal surface to the membranous lateral wall of the
mandibular segment or the head, and the gnathal lobe projects freely
from its distal end (fig. 1 B). Inasmuch as the basal muscles of the
mandible evidently correspond with the coxal muscles of a leg (A),
it is most reasonable to assume that the body of the mandible repre-
sents the basal segment of an ordinary limb, which is that commonly
called the coxa, or coxopodite, and that the gnathal lobe is a coxal
endite. That the basipodite of the mandibular appendage is the first
segment of the palpus is shown in crustaceans having a biramous
palpus, in which the two rami are carried by the basal segment of the
palpus (figs. 2D, 4F). Some writers contend, however, that the
primitive arthropod limb had a “precoxal,” or “subcoxal,” segment

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

proximal to the coxa, and from this idea the mandible has been inter-
preted as being either the “precoxa,” or the “precoxa and coxa com-
bined.” The evidence of a subcoxal limb segment is based principally
on the occasional presence of small sclerites at the root of the limb,
or on ringlike thickenings at the base of the coxa. Stormer (1944),
for example, interprets a short ring supporting the coxa in the leg of
a trilobite as a “precoxa,” but it is difficult to believe that such a
structure observed in a fossil can be regarded with any assurance as
a limb segment. There is no specific evidence in any case that a
so-called ‘“‘precoxa” or ‘‘subcoxa” was ever an individually musculated
and independently movable part of the appendage.

Discussions concerning the nature of the arthropod jaw have cen-
tered largely around the mandible of the copepod Calanus. It happens
that the mandibular palpus of Calanus is distinctly biramous (fig.
4 F), so that the segment supporting the two rami can be identified as
the basipodite (Bspd). The jaw part of the appendage is transverse
and ends with a broad, toothed gnathal lobe (gnL). The basipodite
is attached to the jaw segment by a small ring (bspd). Some writers,
therefore, as Borradaile (1917) and Hansen (1925), have regarded
the intercalated ring as the coxa, and interpret the basal segment as a
“precoxa.” If this interpretation is true for Calanus it would have to
be carried over to all the other arthropods. An examination of the
mandible of Calanus, however, gives no support to the idea that the
ring supporting the basipodite is a true segment; no muscles arise
within it, one small muscle is attached by a tendon on its base, and the
other muscles traverse the ring to be attached on the basipodite. More
definite evidence as to the nature of the ring may be deduced from
the study of Campbell (1934) on the development of the mandible in
Calanus tonsus Brady ; from her figures it appears that the basipodite
ring is not present in the appendage until the first copepodid stage.
From this fact, therefore, Heegaard (1947, p. 197) convincingly
argues that the alleged “coxa” of the Calanus mandible “is merely a
later sclerite ring separated from the basis, so as to give the mandibular
palp a greater mobility,’ and cannot be regarded as a primary seg-
ment. In the following descriptions it will be assumed that the arthro-
pod mandible is in all cases the coxa of the mandibular appendage,
since on it are attached the muscles that clearly correspond with the
coxal muscles of a leg.

The gnathal lobe of the mandible, often called the “gnathobase,”
being the functional part of the jaw, takes on various forms accord-
ing to the nature of the food or the manner of feeding of the animal.
Very commonly the lobe is differentiated into a toothed incisor process,

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 9

and a proximal molar process or masticatory surface (figs. 5 B, C, D;
8B, I; 17F; 22B). Crampton (1921) has followed the relative
development of these processes in the various mandibulate groups of
arthropods, but he has probably attributed too much phylogenetic sig-
nificance to them, since the structure of the gnathal lobe may be quite
different in related forms with different feeding habits. On the incisor
process of the mandible there may be present a small group of loose
teeth, or a small, flexibly attached dentate plate. This structure occurs
among the peracaridan Crustacea, in the Symphyla, the Diplopoda, and
in some insects ; it is known as the “lacinia mobilis” (figs. 6 C; 17 G;
20 G, Im). The name might be appropriate if the term “‘lacinia” is
taken in its literal meaning of a “fringe,” but the structure cannot be
supposed to have any relation to the Jacimia of an insect maxilla, as
Crampton (1921) has sufficiently emphasized, the maxillary lacinia
being itself a musculated endite equivalent to the entire gnathal lobe
of the mandible.

The most leglike mandibular appendage to be found among the
mandibulate arthropods occurs in the ostracod family Cypridinidae.
The mandible of Philomedes, for example (fig. 2 F, G), has the form
of a simple, biramous limb consisting of a basal coxopodite (C#) and
a 3-segmented telopodite, the exopodite being represented by a small
external lobe (F, Expd) of the basipodite. A gnathal lobe is usually
absent, but in some species of the family, as in the male of Philomedes
globosus (G), the coxa bears distally on its mesal surface a small,
weak, bidentate process (guL) that evidently represents the gnathal
lobe of other forms, though certainly it can have little function as a
feeding organ. Though the leglike mandible of Philomedes is not to
be regarded as a primitive mandibular appendage, but rather as a
simplified jaw, which, armed with strong apical spines, has been trans-
formed into a grasping organ for securing food particles, it does,
however, give a clear suggestion of how a simple limb might be con-
verted into a jaw by the development of a gnathal endite on the coxa,
and the reduction of the telopodite. The functional jaws of the
cypridinids are the first maxillae, which in most forms are armed with
strong spines for tearing the food.

That the mandibles are appendages of the same segment in all the
mandibulate arthropods is generally unquestioned. Silvestri (1933)
alone has contended that the jaws of the chilopods, diplopods, and
insects represent the first maxillae of Crustacea, and that the crus-
tacean mandibles are the appendages of a segment that corresponds
with the so-called intercalary, or premandibular, segment in the em-
bryo of the other forms, in which this segment lacks appendages in

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the adult stage. The segment in question, however, is commonly re-
garded as representing the second antennal segment of Crustacea, the
antennae of the chilopods, diplopods, and insects being identified with
the antennules of the Crustacea. The essential likeness in the struc-
ture and musculature of the mandibles in all the mandibulate arthro-
pods, and their innervation from corresponding ganglia make it dif-
ficult to believe that the arthropod mandible is not a common in-
heritance from a common ancestor; differences in the mandibles are
easily seen to be structural modifications correlated with changes in
the jaw mechanism.

The principal structural changes of the mandibles and the func-
tional changes of the muscles that take place in the higher arthropods
result from the acquisition by the mandible of a second articulation
with the head on the anterior margin of its base (fig. 1 C, c) at the
end opposite from that bearing the primary dorsal articulation (a).
This secondary articulation is ventral, dorsal, mesal, or anterior in
relation to the primary articulation, according to the position assumed
by the mandible, and it is not always with the same part of the head,
but it gives the jaw a fixed axis of rotation (a-c) between the two
articular points. The distal articulation does not represent the primary
sternal articulation of the appendage; it is never on a true sternal part
of the head, and it lies anterior to the ventral muscles. The newly
established axis (a-c) thus runs close along the anterior, or outer,
side of the mandible, but just within the attachment of the anterior
dorsal muscle (4). The doubly articulated mandible, therefore, is
closely hinged to the head by its anterior margin, and its movements
resemble those of a door on its hinges; but the mandible differs from
a door in that the motor power is applied on both sides of the axis.
According to Schmidt (1915), Berkeley (1928), and Cochran (1935),
the doubly articulated mandible can no longer “rotate.” Actually,
however, it is to be seen that its movements are the rotary motions of
a pendent jaw with one articulation, but on a doubly articulated axis;
the rotary movements are now called abduction and adduction.

With the altered mechanism of the mandible resulting from the
articular innovation, the anterior dorsal muscle (fig. 1C, A), if it
retains its origin dorsal to the mandible, becomes an abductor, and
the directly opposed posterior dorsal muscle (P) becomes an adduc-
tor. The ventral muscles (/) are still adductors as in a singly articu-
lated mandible. Schmidt (1915) and Berkeley (1928) contend that
it is impossible to determine the homologies of the muscles of a doubly
articulated mandible with those of an ambulatory limb, but Cochran

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS Wa

(1935) has shown that the relation is very simple, and her explanation
is that given here.

If the gnathal lobe of the doubly articulated mandible retains its
primitive position in line with the axis of the mandibular base (fig.
1 C, gnL), as it does in most of the malacostracan Crustacea and in
the Chilopoda, the opposing lobes cannot now swing toward each other,
they simply work in the manner of a pair of valves opening and closing
from below with the rotation of the mandibles. Mandibles having
this kind of mechanism are not efficient biting and chewing organs,
and must depend on having the food passed to them by some of the
following appendages, which are particularly modified to serve as
accessory feeding organs.

An improvement in the mechanism of the jaws for increasing the
adductor power of the gnathal lobes, however, has been evolved in
the decapod Crustacea. By the development of an apodemal lobe on
the marginal part of the mandible (fig. 1D, Ap) that carries the
anterior muscle, which here consists usually of two or three bundles of
fibers (A), the insertion of the muscles is brought above the hinge
line (a-c), and is accompanied by a lowering of the points of origin
of the muscles on the carapace, so that these primarily abductor fibers
(C, A) now become adductors (D). Opposed to them is a differen-
tiated group of the ventral fibers (D, 2V’) attached on the inner face
of the apodeme above the hinge line ; these fibers thus lose their origi-
nal adductor action and become a ventral abductor muscle. This struc-
ture and mechanism of the mandible is characteristic of the natantian
and astacuran decapods. The Anomura and Brachyura have still fur-
ther improved on it by carrying the apodeme out proximally as an
arm projecting in line with the body of the jaw beyond the lateral
articulation (E, Ap). The apodeme, with its opposing muscles, by
this alteration becomes an efficient lever for operating the gnathal lobe.
This type of mechanism reaches its highest development in the crabs.
A mandible rotating on its long axis, however, is still not the most
effective kind of jaw, since it has little power of grasping and in
general serves only as a masticatory organ for food passed on to it by
the following appendages.

The mandibles of the chilopods and the entognathous apterygote
insects resemble those of the decapods in that they lie horizontally
against the under surface of the head, and the gnathal lobes project
in line with the mandibular axes. In these terrestrial groups the
mandibles may also be doubly articulated for rotary movement, but
the articulations are not fixed points of attachment, and the jaws are
more or less protractile. The protractor muscle is generally a differ-

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

entiated group of the ventral fibers (fig. 1 F, 2V) attached on an
apodemal support.

By a simple modification of form the doubly articulated mandible
has acquired its greatest efficiency as a biting and chewing jaw. The
change involves merely a shift of the gnathal lobe from a position in
line with the mandibular axis (fig. IC) to one approximately per-
pendicular to the axis (G). The mandible thus again swings trans-
versely, but now on a firmly hinged axis (a-c) instead of on a single
point of articulation as in the primitive mandible (B), so that the two
jaws are able to close strongly against each other. Mandibles of this
type are largely relieved of dependence on other appendages for the
securing of food; by elongation they may become fangs for grasp-
ing living prey, and they are particularly amenable to modifications
by which they become piercing organs. The anterior, or outer, dorsal
muscle of the mandible (fig. 1 G, A) retains the abductor function,
the posterior dorsal (P) becomes the chief or only adductor; the
ventral muscles (V) lose their importance as adductors, and are
reduced or eliminated. Mandibles of this kind have been evolved,
apparently independently, in the amphipods and isopods among the
Crustacea, and in the Lepismatidae and Pterygota among the insects.
The winged insects have the most efficient jaws of all the arthropods
for direct mandibular feeding.

Finally, we encounter the curious condition in which the gnathal
lobe, ordinarily a solid outgrowth of the mandibular base, becomes
either flexible or movably articulated on the base, and independently
musculated. The first condition occurs in the Chilopoda, the second
(fig. 1 H) is characteristic of the Diplopoda and Symphyla. In the
diplopods and symphylids the gnathal lobe (gnL), which is the func-
tional jaw of the animal, is supported on a large basal plate (mdB)
on the side of the head (fig. 20 A, B). That this plate, though rela-
tively immovable, is the true base of the mandible is shown by the
fact that most of the usual mandibular muscles are inserted on it
(fig. 1H). Attached on the gnathal lobe, however, is a huge cranial
flexor of the lobe (14), and a smaller muscle (/) arising within the
basal plate. The cranial muscle, since it is attached on the margin of
the lobe and goes anterior to the ventral muscles (1V, 2V), may be
regarded as an anterior dorsal muscle. More difficult to explain is
the presence of the intramandibular muscle (J). It can hardly be
doubted that the gnathal lobe of the diplopod and symphylid mandibles
is the homologue of the immovable lobe in other arthropods, and that
it does not represent a segment of the telopodite. It may be supposed
to be derived from a flexible lobe with a similar musculature such as

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 1

that of most of the chilopods. The movable gnathal lobe of the mandi-
ble is identical in its essential structure and its musculature with the
lacinial lobe of an insect maxilla, and it is to be noted that endites of
the maxillary appendages in general are movable and independently
musculated from the limb segment on which they arise.

The distribution of the types of mandibular structure among the
arthropods does not show any evolution of the types from one major
group to another. Among the Crustacea, for example, are found all
the different kinds of mandibles having an immovable gnathal lobe,
including mandibles with a single point of articulation, doubly articu-
lated mandibles, some with a horizontal valvelike action, others with a
transverse swinging movement, and also piercing mandibles. In the
Chilopoda the mandibles are of the horizontal valve type, rocking on a
lengthwise axis, though without fixed articulations. The mandibles
of the entognathous apterygote insects somewhat resemble the chilo-
pod jaws, but they may be modified for piercing. Among the other
insects, mandibles of the generalized type with a single point of articu-
lation recur in the thysanuran Machilidae, and in modified form in
larval Ephemeroptera. The characteristic insect mandible, however,
is a doubly articulated jaw with a free transverse movement, though
the piercing type is of frequent recurrence among the Pterygota.
Mandibles with a movable, independently musculated gnathal lobe are
characteristic of the Symphyla and Diplopoda, but the jaws of the
crustacean Branchiura and Cirripedia, 7f they are mandibles, are to
be included in the same category. There is good reason for believing
that the pendent, singly articulated mandible represents the primitive
arthropod jaw, because it shows the least departure from the coxa
of a leg, but it is evident that the other types of mandibular structure
and mechanism have been independently evolved in the various arthro-
pod groups.

E. CRUSTACEA

The principal types of mandibular structure that occur in the Crus-
tacea have been sufficiently outlined in the Introduction. A review of
the subject, therefore, need not be repeated here, and the following
descriptions will simply give examples of the jaw structure and
mechanism developed in the various crustacean groups.

Branchiopoda and Ostracoda.—The jaws of the branchiopods
well illustrate the structure of the pendent type of mandible with a
single dorsal point of articulation (fig. 1B). In the Anostraca (fig.
2A, B) the mandibles (Md) are articulated on the tergum of the
mandibular segment (JJ), which is a small but distinct plate between

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

_ Endpd Expd |

Fic. 2.—Crustacea—Branchiopoda and Ostracoda.

A, Eubranchipus vernalis Hay, female (Branchiopoda: Anostraca), head and
anterior part of body, lateral. B, same, mandibles and muscles, anterior. C,
Apus longicaudatus Leconte (Branchiopoda: Notostraca), mandibles and mus-
cles, anterior. D, Cypris testudinaria Sharpe (Ostracoda), right mandible,
mesal. E, Daphnia pulex Degeer (Branchiopoda: Cladocera), mandibles and
muscles, anterior. F, Philomedes globosa (Lillj.) (Ostracoda), left mandible,
lateral. G, same, left mandible, anterior. H, Estheria clarkii Packard
(Branchiopoda: Conchostraca), mandibles and intergnathal muscles, anterior.

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 15

the protocephalic head (A, Prtc) and the large tergal plate of the
maxillary segments (//J, JV). In the notostracan Apus (C) the
mandibles are suspended from the under lamellae of the lateral folds
of the mandibular part of the shell. The mandibles in each case have
large gnathal lobes, but the lobes are not differentiated into incisor and
molar processes. Palpi are absent. The same type of mandible is
seen in the conchostracan Estheria (H), and in the cladoceran
Daphnia (E). Each mandible is equipped with strong anterior and
posterior dorsal muscles (B, C, E, A, P), and the whole inner cavity
of the jaw is occupied by the spreading fibers of the ventral adductor
muscle (V), the convergent ends of which are united in a thick median
ligament with those from the opposite mandible. The strong de-
velopment of the dorsal muscles leaves no doubt that these muscles
are functionally important ; probably they give a rotary motion to the
jaws, or perhaps some degree of anterior and posterior movement,
but, acting together, they might also be adductors. A muscular mech-
anism of abduction, however, is not evident.

The apparent strength of the mandibles and their musculature in
some of these small or minute crustaceans is surprising considering
the nature of the food, which, for the most part, consists of organic
detritus or micro-organisms filtered from the water, only a few species
being predaceous. Elaborate studies have been made by Cannon and
others on the feeding mechanism that brings the food to the mouth,
but little is said about the specific action of the jaws.

The Branchiopoda in general, except Notostraca, as described by
Cannon (1928; 1933b), obtain their food from water currents driven
forward to the mouth in a median channel of the ventral body wall by
movements of the trunk limbs. The water enters the food channel
through the interlimb spaces, and the contained particles are either
filtered off on setal fringes of the basal endites of the limbs, or are
carried directly in the forward current to the mouth region. The
particles lodged on the filters are scraped off, as the latter move for-
ward and backward, by combs of setae on the walls of the food chan-
nel, and are then caught in the water current. On reaching the mouth
region the accumulated food may be introduced at once between the
mandibles by the maxillules, or in some species it is first agglutinated
into a mass by a secretion of glands in the labrum. A special descrip-
tion of the labral glands of cladocerans is given by Cannon (1922).
The Notostraca lack a median food channel, and with them there is no
perceptible forward-flowing water stream. Food particles entering
between the limbs are caught on the spiny basal endites, and, with the
forward and backward movement of the limbs, are successively

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

scraped off upon the preceding endites, and so eventually reach the
mouth. Both Apus and Lepidurus, Cannon says, feed also on large
food masses grasped with the anterior trunk limbs and held against
the mouth.

Among the Ostracoda the mandibles take on various forms, but
they are more generalized than those of the branchiopods in the reten-
tion of a segmented palpus, which is biramous. In most species, as in
Cypris (fig. 2D), the long basal part of the mandible has the struc-
ture typical of the branchiopod jaw with a simple, strongly toothed
gnathal lobe. In certain species, however, the lobe is armed with
strong spines, and in some of the Cytheridae it is produced into a
piercing stylet. The simplified, leglike mandible of Philomedes (F,
G) has been noted in the Introduction.

The Ostracoda feed in various ways. According to Schmitt (1931)
fresh-water forms, so far as observed, seem to be omnivorous, but
marine species feed largely on diatoms and other plants of the ocean.
Some, however, feed on copepods, which they ensnare with a sticky
secretion spread over the prey. Species with piercing mouth parts
suck the juices of marine plants, and a few are predaceous on other
animals. Among the marine ostracods, Asterope and Cytherella are
said by Cannon (1933a) to be purely filtratory feeders. By the
activity of the maxillary epipodites currents of water are drawn
through the chambers within the shell valves. The filters of Asterope
are combs of long setae on the maxillules, the particles lodged on them
are scraped off by setae of the maxillae and spinous lobes of the
mandibles, and passed to long setae on the maxillularly endites, which
deliver the food mass to the mouth, from which finally it is introduced
into the esophagus by curved processes of the mandibles. Members
of the Cypridininae, Cannon says, may feed on detritus from currents
driven through the shell chamber by the same mechanism as in Aste-
rope, but they are not true filter feeders, and some or raptatory. The
food particles from water currents are collected on setae of the basal
parts of the maxillules, the maxillae, and the first trunk limbs, and the
food is entangled in a secretion from glands in the large labrum.
Cypridina feeds on large food masses, which, Cannon suggests, are
held by the mandibular palps directly under the mouth and here torn
to pieces by the strongly armed maxillulae. Philomedes, on the other
hand, feeds on small particles dislodged by means of its spiny mandib-
ular palps (fig. 2 F, G) from the mud over which it swims, and it
has only a relatively weak maxillulary armature. Gigantocypris,
Cannon notes, “must be an efficient hunter of living prey,” since its

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS W7,

stomach was found to be full of large copepods; Cypridina castanea
“feeds on comparatively large Crustacea.”

Branchiura.—The functional jaws of adult branchiurans are small,
toothed plates or hooks mostly or entirely concealed within a preoral
“buccal cavity” in the end of a proboscis. In Argulus the proboscis
is an elongate tubular organ (fig. 3 A) projecting posteriorly on the
under side of the head; in Dolops it is a mere hexagonal mound (1)
between the bases of the maxillulary hooks. The preoral cavity of
the proboscis is enclosed between an anterior upper lip (Lm) and a
posterior lower lip (Mst) ; the jaws, commonly called “the mandibles,”
project mesally from the lateral walls of the cavity. Rodlike thicken-
ings of the proboscis wall strengthen the latter and serve as supports
for the jaws.

In a young stage of the embryo the apparent mandibles are fully
exposed appendages on the under side of the head; as shown by
Martin (1932) in Argulus “viridis” (fig. 3C), each mandible of a
26-day embryo consists of a basal segment (mdB) bearing a large,
lateral, 3-segmented palpus (P/p), and a small, toothed gnathal lobe
(gnL) projecting from its mesal end. At a later embryonic stage
(D), according to Martin, the basal segments of the mandibles have
increased greatly in size and appear to have formed the principal
part of the proboscis, their distal parts uniting with lobes of the head
that become the upper and lower lips of the preoral cavity. The apical
lobes of the mandibles now appear as a pair of hooks (D, gnL) in the
end of the proboscis. The same structure found by Martin in the
35-day embryo of Argulus “viridis” (D) is carried over into the first
larval stage, as shown here in Argulus americanus (E), except that
the proboscis spine (Spi) is now present. At this stage the mandibular
palps (mdPlp) still arise from the base of the proboscis, and are
widely separated from the hooklike gnathal lobes (gn) in the end
of the proboscis. Martin’s statement, therefore, that the principal
part of the proboscis is derived from the mandibular bases seems to
be well substantiated. If so, the mouth hooks are not themselves “the
mandibles,’ but are the displaced gnathal lobes. The labrum and
paragnaths, Martin believes, are represented by three small processes
that form a filter apparatus at the mouth entrance within the preoral
cavity. In no other crustacean, however, do these structures occur in
any such place; it would seem much more probable that the upper lip
is the labrum (A, B, I. Lm), and that the lower lip (A, I, Mst) isa
metastomal lobe formed of the paragnaths. These elements would be
readily available in any crustacean for the construction of a proboscis.
The mandibular palps, Martin says, are lost at the first larval moult.

18

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Z:-
4.
in
ba
ie
Z
bs
A:
Se
ZS:
Ze.
Zz
Ae
&:
ee
E
Rt

Fic. 3.—Crustacea—Branchiura.

A, Argulus americanus Wilson, end of proboscis, ventral. B, same, labrum
and jaws, ventral. C, Argulus viridis Nettovitch (foliaceus L.), mandibles
of 26-day embryo (from Martin, 1932). D, same, proboscis of 35-day embryo
(from Martin, 1932). E, Argulus americanus Wilson, proboscis of newly hatched
larva. F, Argulus viridis Nettovitch (foliaceus L.), gnathal lobe of mandible
(from Martin, 1932). G, Argulus laticauda Smith, gnathal lobe of mandible.
H, Argulus pugettensis Dana, gnathal lobes of mandibles. I, Dolops doradis
Cornalia, proboscis, ventral. J, same, gnathal lobe of mandible. K, same,

gnathal lobe of mandible, and supporting plates in wall of proboscis.

a a

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 19

The adult jaws of Argulus vary in shape and dentition in different
species, as shown at B, F, G, and H of figure 3. The jaws of Dolops
doradis are strong serrated hooks (J, K) projecting mesally from the
lateral walls of the moundlike proboscis, and are partly exposed in
the preoral cavity between the labrum and the metastome (I). Each
hook arises from the posterior end of an elongate base (J, K), which
is merely a local sclerotization in the otherwise membranous lateral
wall of the proboscis. At about its middle the hook base is pivoted
on the tapering end of a slender transverse sclerite (J, K, w) in the
ventral wall of the proboscis, the outer end of which is held in a
notch of a second more lateral sclerite (K, v). On each side of the
fulcral point muscles are attached on the hook base that evidently
rock the latter and thus produce adduction and abduction of the hooks,
the jaw hooks being movable by reason of the flexibility of the pro-
boscis integument in which their bases are implanted.

Inasmuch as the observations above cited, if true, seem to show
that the functional jaws of adult branchiurans represent the apical
hooks of the embryonic mandibles, and thus evidently correspond
with the immovable gnathal lobes of the mandibles of most other
Crustacea, it is surprising that the structures in question are indi-
vidually movable and independently musculated. Though the muscula-
ture of the branchiuran proboscis needs to be more carefully studied,
there is no question that the jaw muscles arise within the proboscis
itself ; but this condition is one characteristic of maxillary endites,
and becomes so pronounced in the case of the first pair of jaws in the
Cirripedia that the latter have been interpreted as maxillulary endites,
and not as mandibles. The musculature of the branchiuran jaw lobes
might be justified if we could suppose that the lobe of the embryonic
mandible (fig. 3 C, gnL) represents the endopodite of the appendage
and that the palp (Plp) is the exopodite, but there is little in the struc-
ture of the organ to support such an interpretation. The interpretation
of the branchiuran jaws, as given in figure 3, therefore, must be held
subject to further investigation, but the same anomalous condition
seems to be even more pronounced in the Cirripedia.

Cirripedia.—The mouth parts of the ordinary nonparasitic bar-
nacles, or Thoracica, are so closely associated with one another
around the mouth that together they form a thick, proboscislike lobe
with a somewhat constricted base (fig. 4A) projecting from the
ventral side of the head. The large, swollen, strongly sclerotized
anterior part of the lobe is the labrum (Lim). Closely adnate on each
side of the labrum is the wide base of an appendage that supports
ventrally an elongate, hairy palpus (P/p) projecting forward beneath

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Gn n ff IN B
wi gs in awe i

Fic. 4.—Crustacea—Cirripedia (Lepas) and Copepoda (Calanus).

A, Lepas anserifera L., mouth parts, left side. B, same, mouth parts and
mouth, ventral. C, same, labrum and first gnathal appendages, posterior,
showing first pair of jaws (1Gn). D, same, second jaw of right side, mesal.
E, same, second and third gnathal appendages, posterior, showing position of
nephropores. F, Calanus cristatus Kréyer, left mandible of fifth copepodid
stage, ventral.

Pom.

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 21

the labrum, and bears behind the palpus a large, free, flat, strongly
toothed jaw lobe, or gnathos (zG7), which is independently movable
on the base. The jaw is turned mesally behind the labrum (B, C,
1Gn), so that only its outer edge is visible in side view (A). Above
and behind this first lateral appendage on each side is the base of a
second, smaller appendage (A, 1M), which bears a second platelike
jaw (2Gn) similar to the first though smaller and not so strongly
toothed (D). Finally, projecting ventrally behind the other mouth
parts is a pair of large, thick, soft, rounded, hairy lobes (A, B, 2M~x),
with a deep groove between them that runs forward to the mouth.

The cirriped mouth parts were first well described by Darwin (1851,
1854), who regarded the first pair of appendages and their jaw lobes
as the mandibles, the second pair as the first (“inner”) maxillae, and
the two postoral lobes as the second (“outer”) maxillae. Darwin’s
interpretation and nomenclature have been followed by most subse-
quent students of the cirripeds.

An examination of the so-called mandible will show at once that its
structure is quite unlike that of any ordinary crustacean mandible.
The effective jaw lobe of Lepas anserifera (fig. 4C, 1Gn) is freely
articulated on a small sclerite of the posterior edge of the base of
the appendage, and has a strong individual musculature consisting of
abductor and adductor muscles arising in the base. The base itself is
immovably attached to the side of the labrum, and its outer wall is
divided by a groove into an upper and a lower part (A) suggestive
of a segmentation, but the division appears to be merely a surface
differentiation. The entire body of the appendage is filled with muscle
fibers inserted on the palpus and the jaw lobe; the lower part con-
tains an external layer of longitudinal fibers. From their structure,
these appendages might well pass for maxillae with a highly developed
biting endite. In fact, it has already been said by Hansen (1925,
p. 51) that the appendages of the cirripeds called “mandibles” differ
“so strongly from the mandibles in other Arthropoda while agree-
ing much more with the maxillulae or maxillae, that I prefer to name
them maxillulae; consequently mandibles are absent.” It should be
noted, however, that the mandibles of the Diplopoda and the Symphyla
have independently movable and individually musculated gnathal
lobes quite comparable to the jaw lobes of cirripeds. Borradaile (1917,
1926) suggests that the jaw lobes of the cirriped mandibles may be
endites of the second segments of the appendages and not those of
the first, but it is not clear how this interpretation makes the matter
any easier to understand. In the cypris stage of cirriped ontogeny
the second antennae and the mandibles of the nauplius are suppressed ;

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Darwin (1851) says the mouth parts of the adult are all present in
the cypris stage in an undeveloped condition, but apparently it has
not been shown that the first pair of mouth appendages of the adult
are derived from the naupliar mandibles.

The second pair of mouth-part appendages of Lepas (fig. 4 D) are
much simpler than the first. The basal part of each is membranous
(A, 1Mz) ; the jaw lobes are smaller and simpler than those of the
first pair, and lie behind the latter (B, 2G”). From the base of each
second jaw projects a large apodemal arm (D, Ap), but the relation
of the muscles to the apodeme was not determined. If the first ap-
pendages are the mandibles, the second are the first maxillae.

The postoral lobes of the cirriped group of mouth parts (fig. 4 A,
B, E, 2Mx), as Hansen (1925) notes, in no way suggest by their
form or position that they represent the second maxillae; the deep
groove between them runs forward to the muoth. In short, these
lobes have the position and character of a pair of thick paragnaths.
Hansen remarks that only a single circumstance makes it doubtful
that the postoral lobes are the paragnaths, which is that the excretory
glands are said to open on their bases. In Lepas anserifera the nephro-
pores (E, npr) are not on the bases of the lobes, but lie behind them
plainly exposed in the membrane between the second and third ap-
pendages. The apertures are shown in the same position’ by Darwin,
who regarded them as “olfactory organs.” On the other hand, in the
lepadid Conchoderma Dephner (1910) plainly shows the gland ducts
opening on the bases of the postoral lobes, as he says they do also
in Balanus, and Batham (1945) shows the nephropores on the bases
of the lobes in Pollicipes. The position of the gland openings must
be given priority over all other considerations as evidence that the post-
oral lobes of the cirripeds are the second maxillae, for, as Borradaile
(1926) says, the assumption that the gland apertures have migrated
from the maxillae to the paragnaths “will probably not commend
itself to carcinologists.” The usual interpretation of the mouth parts
of Lepas is implied in the lettering given here on the figures, but the
homology of the parts of the mandibles with those of the jaws of other
crustaceans must be left undetermined. The cirriped “mandibles,”
however, appear to have something in common with the branchiuran
“mandibles.”

Copepoda.—The copepod mandibles show an extreme degree of
variation from the generalized calanoid type of jaw to that of parasitic
forms in which the mandible takes the form of a long arm or slender
stylet armed with teeth on its distal part. The mandible of Calanus
cristatus (fig. 4 F) has been sufficiently discussed in the Introduc-

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 23

tion, but we may note again that its basal segment is produced mesally
in a large, flat, toothed gnathal lobe (gnL), and bears a biramous
palp. Between the basal segment, or coxopodite (Cxpd), and the
basipodite (Bspd), which supports the two rami, is an intermediate
ring (bspd) that has been interpreted by Borradaile (1917) and by
Hansen (1925) as the “coxa” of the appendage, making the basal
segment a “‘precoxa.” There is nothing in the nature of the ring, how-
ever, to give it the status of a true segment ; in appearance it is merely
a secondary subdivision of the basipodite. Many examples might be
cited among the arthropods of subdivision of primary limb segments,
producing parts that may have the appearance externally of seg-
ments, but which have no musculature of their own; the ring sup-
porting the basipodite in the Calanus mandible is evidently a struc-
ture of this kind. Before any limb section devoid of muscles can be
held to be a true segment it must be shown by evidence from some
source that it was once an individually musculated and therefore inde-
pendently movable part of the appendage. The basal segment of the
Calanus mandible certainly is the coxopodite of an ordinary limb ;
its ventral muscles (/) consist of eight or nine slender bundles of
fibers, and those from the opposite jaws come together medially,
where they are attached on a narrow intergnathal ligament above the
nerve cord, which expands anteriorly and branches into a pair of
suspensory ligaments attached on the back.

Numerous examples of the elongate or styletlike mandibles of
parasitic copepods have been well illustrated by Heegaard (1947),
who shows that while some are simple rods, most of them are divided
by joints into several segmentlike parts. From comparative studies
on different forms Heegaard argues that the divisions represent true
segments in course of fusion. From this interpretation it would ap-
pear that the styletlike mandible of the copepods is formed of the
main shaft of the appendage. The usual mandibular stylet of other
arthropods, however, is the elongated gnathal lobe, the telopodite being
absent.

Leptostraca.—The mandibles of Nebalia (fig. 5 A) are of the
suspended type of structure; they hang from dorsal articulations
(a) on the head, with the large, 3-segmented palpi projecting from
their lower ends. The long gnathal lobes turn mesally at right angles
from the bases of the jaws, and come together behind the edge of the
labrum. From the base of each lobe there arises anteriorly a small,
thin, bidentate incisor process (C, inc). The main part of the lobe,
therefore, constitutes a molar process (mol). The incisor process of
the mandible, first met with in the Leptostraca, as seen in Nebalia

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Fic. 5.—Crustacea—Leptostraca (Nebalia) and Anaspidacea

(Anaspides and Paranaspides).

A, Nebalia bipes (Fabr.), mandibles and muscles, posterior. B, Anaspides
tasmaniae Thomson, mandibles and muscles, posterior. C, Nebalia bipes
(Fabr.), right mandible, anterior. D, Paranaspides lacustris (Smith), right
mandible, anterior (from Smith, 1908). E, Nebalia bipes (Fabr.), intergnathal
ligament (from Manton, 1934). F, Anaspides tasmaniae Thomson, intergnathal
ligament and suspensory branches, dorsal, flattened under cover glass.

NOs JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 25

appears to be a secondary outgrowth of the gnathal lobe rather than
a result of the division of the lobe into two parts; but, on the other
hand, if Nebalia, as Cannon (1927) contends, has been evolved from
a mysidlike malacostracan, its small incisors may be remnants of once
large and functional processes. The musculature of the Nebalia
mandibles includes two relatively weak dorsal muscles for each jaw
(A, A, P), and large ventral adductors (V) attached medially on the
mandibular expansions of an elaborate intergnathal ligament (E).

Nebalia lives in shallow water along the shore and in soft mud
where there is much organic decay, and feeds largely on particles
filtered out of water currents produced by movements of the trunk
appendages. The filter chamber, as described by Cannon (1927), is
an enclosure beneath the thorax shut in laterally by the appendages,
below by a mat of setae on the ends of the endopodites, and closed
behind by setae on the last pair of limbs. The chamber is thus open
anteriorly, and the water currents enter im front and flow posteriorly.
The contained particles are filtered out on fringes of long setae on
the endopodites of the trunk limbs, which overlap mesally from be-
hind forward. By the forward and backward movement of the
appendages, therefore, the particles lodged on the setae are auto-
matically transferred to those of the limb in front, and so on until
they reach the maxillary region. Here they are taken over first by
the maxillae and transferred to the maxillules, which give the food
a preliminary mastication, and then finally pass it forward between
the paragnaths to the molar processes of the mandibles. The small,
widely separated incisor processes, Cannon notes, do not act as biting
parts. Particles that are too large to pass the filters are stopped in the
maxillary region and utilized directly.

Anaspidacea.—Though the anaspidaceans, or syncarids, are true
Malacostraca, they still retain the primitive type of mandible (fig.
5B) supported by a single point of articulation (@) on the tergum
of the mandibular segment. The large gnathal lobes are differentiated
into broad molar processes (B, D, mo/) and toothed incisor processes
(inc). The mandibular palps of Anaspides (B) are 3-segmented,
but in Paranaspides (D) the palps of the adult are shown by Smith
(1908) to have four apparent segments, and the first segment bears
a small lateral lobe suggestive of being a palpal exopodite. In young
specimens, however, Smith (1909) finds that the palp may have only
three segments and that the basal lobe is less conspicuous, from which
fact he doubts that the palp of Paranaspides shows a true biramous
structure.

The musculature of the Anaspides mandibles (fig. 5 B) is the same

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 116

as that of the branchiopod mandibles (fig. 2B, C) in that each jaw
has an anterior and a posterior dorsal muscle (A, P) and a large
bundle of ventral adductor fibers (V). In Anaspides, however, the
posterior dorsal muscle (P) is attached, as in the Malacostraca gen-
erally, by a thick tendon on the posterior mandibular margin close to
the base of the gnathal lobe. In the higher Malacostraca having a
doubly articulated jaw, this muscle becomes a powerful dorsal adduc-
tor (fig.9 D, E, P). The ventral adductor fibers of Anaspides (fig. 5
B, V) all arise on the surface and margins of the mandibular part of a
large, strongly developed intergnathal ligament (F) but in the natural
condition only a small median area of the ligament is exposed (B, Lg).

Concerning the feeding habits of Anaspides tasmaniae, Smith
(1909) says these fresh-water crustaceans “appear to be omnivorous,
as they will feed upon the dead bodies of insect larvae or even upon
one another, but their chief food is the algal slime covering the rocks
among which they live, and they also browse upon the submerged
shoots of mosses and liverworts.” Manton (1930) notes that in addi-
tion to feeding on algae and detritus covering weeds and stones, the
adults of Anaspides eat also tadpoles and worms.

The large size of the jaws of Anaspides, their strong musculature,
and their well-developed incisor and molar processes indicate that
these crustaceans are well equipped for direct feeding on the bodies of
dead animals, and Cannon and Manton (1929) observe that “an ex-
ternal view of the mouth parts suggests a raptatory function rather
than a filtratory type of feeding mechanism.’ However, these writers
show that Anaspides and Paranaspides are both filtratory and rapta-
tory feeders, though the related Koonunga, judging from its mouth
parts, apparently is entirely raptatory. (By “raptatory” is meant
grasping or scraping, but not necessarily predatory habits.) The filter
apparatus of Anaspides and Paranaspides as described by Cannon and
Manton is formed of the broad maxillae and their dense fringes of
marginal setae, which lie close against the maxillules. The maxillary
filter plates enclose between them a filter chamber through which
water is driven from behind by the action of the trunk limbs. Food
particles in the water caught on the setal fringes of the maxillae are
scraped off by the basal endites of the maxillulae and passed forward
to the mouth. The animals are thus equipped for feeding on detritus
dislodged from submerged plants and stones.

Euphausiacea, Cumacea, Mysidacea.—In these three malacostra-
can groups the mandibles begin to take on the structure and mechanism
that become characteristically developed in the doubly articulated jaws
of the decapods, resulting first from a close association of the mandi-

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 27

bles with lateral expansions of the epistome. The calcification of the
epistomal region spreads to the sides between the bases of the second
antennae and the mandibles, usually uniting with the inner lamellae
of the carapace folds, and thus establishes a firm support for the
anterior margins of the mandibles, which take a more or less horizontal
position. The movement of the mandibles is thereby limited to a
lengthwise rocking or rotary motion on the epistomal wings, and
abduction and adduction now consist of a valvelike opening and clos-
ing of the gnathal lobes from below. In the stomatopods and decapods
the mandibles further acquire specific articulations on the epistome
at the sides of the labrum. Characteristic of mandibles of this type
is the attachment of the posterior dorsal muscle by a strong tendon
on the mandibular margin at the base of the gnathal lobe (fig. 6 A,
D, F, P or Pt), a condition already noted in Anaspides (fig. 5 B),
so that this muscle becomes an effective dorsal adductor.

The mandibles of Thysanopoda tricuspidata (fig. 6D), taken as
an example of the Euphausiaceae, are strongly developed, with broad
gnathal lobes differentiated into thick molar processes and toothed
incisor processes. The molar process of the left jaw is a blunt projec-
tion, that of the right is cup-shaped; when the jaws are closed the
left molar fits into the cavity of the right as a pestle into a mortar,
and the incisor teeth interlock with each other. The mandibles are
closely attached by their anterior margins on the weakly calcified post-
antennal expansions of the epistome, but they have no specific episto-
mal articulations or hinges. The fibers of the ventral adductor muscles
arise from a broad, membranous median ligament (Lg), which is
supported by two pairs of suspensory branches (s/) attached directly
on the dorsal epidermis of the carapace by their expanded but non-
muscular ends.

The elongate mandibles of Cumacea (fig. 6 FE, F) lie almost in a
horizontal plane, directed convergently forward from their articula-
tions on the carapace, with the gnathal lobes lying against the labrum.
The well-sclerotized epistome (G, Epst) has broad lateral extensions
reflected back along the sides of the labrum, and the mandibles are
hinged (h) on the epistomal margins, so that their movement is
definitely restricted to a lengthwise rotation on their dorsal anterior
margins. The mandibles of Diastylis glabra (fig. 6E) are more
slender than those of D. rathkei (F) figured by Hansen (1930),
but in both species the molar lobes and the incisor lobes are widely
divergent, giving the jaw a branched appearance. The mandibular
musculature (E) includes the usual anterior and posterior dorsal
muscles (A, P) and large ventral muscles. The anterior dorsal

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

ig
( Wy
, ig
\WY;

a
i)

Fic. 6.—Crustacea—Mysidacea (Neomysis), Euphausiacea (Thysanopoda), and
Cumacea (Diastylis).

A, Neomysis franciscorum Holmes, mandibles and intergnathal muscles,
posterior. B, same, right mandible and muscles, mesal. C, same, gnathal
lobe of left mandible, posterior. D, Thysanopoda tricuspidata M. Edw., man-
dibles and intergnathal muscles, posterior. E. Diastylis glabra (Zimmer),
mandibles and muscles, anterior. FF, Diastylis rathkei Kr., mandibles and
muscles, posterior (from Hansen, 1930). G, Diastylis glabra (Zimmer),
Sole epistome and labrum. H, same, intergnathal ligament and its mandibular
muscles.

=

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 29

(A) is differentiated into several fiber groups, but they all appear to
be abductors opposed to the posterior adductor (P). The ventral
adductors are composed mostly of bundles of striated fibers (H, Mcl)
within the cavities of the mandibles that are continuous from non-
striated fibers radiating from the median ligament (Lg). Some of
the posterior muscle fibers, however, as shown by Hansen (F), run
without interruption from one mandible to the other, though they
are attached on the ligament. The intermandibular ligament is sup-
ported by a pair of thick suspensory branches (FE, F, H, sl) with
muscular attachments on the carapace.

The cumaceans are said to feed on detritus from the mud in which
they bury themselves, and the dense setal fringes on the incisor
processes of the mandibles (fig. 6E, F), which cover the mouth
entrance, appear to make a retaining sieve when the jaws are closed.
The strong incisor and molar lobes and the elaborate musculature of
the jaws, however, suggest that these crustaceans are capable of
handling tougher kinds of food.

In the Mysidacea the mandibles (fig. 6 A, B) in their structure and
mechanism closely approach those of the decapods. They are hinged
by their dorsal anterior margins on lateral wings of the epistome,
though there are no specific points of articulation. The anterior
dorsal muscles include three groups of fibers (B, 1d, 2A, 3A). The
first two (1A, 2A), being attached on a flange mesad of the hinge
line (1), appear to have thus become dorsal adductors, as are the
corresponding muscles in the decapods, but the large third muscle
(3A) evidently retains its original abductor function. The posterior
muscle (P), attached by a strong tendon (A, Pt) near the base of
the gnathal lobe, is a dorsal adductor. The fibers of the ventral mus-
cles (V) occupy the entire cavities of the mandibles (A, B), and
were not observed to be differentiated into abductor and adductor
groups as in the decapods. The mysid mandible gives a good example
of the so-called “lacinia mobilis,” a small toothed plate (C, Jm) flexibly
attached on the incisor area of the gnathal lobe.

It is shown by Manton (1928) that in the development of Hemimy-
sis lamornae the ligament of the ventral adductor muscles of the
mandibles becomes attached on the apex of a V-shaped apodemal
plate arising from sternal invaginations in the maxillary region. This
condition foreshadows that in the decapods, in which the ventral
adductor fibers of the mandibles arise directly on the cuticular “head
apodeme” formed of sternal invaginations between the first and
second maxillary segments.

Hemimysis is said by Cannon and Manton (1927) to exhibit “two

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

distinct types of feeding, one on large food masses and the other on
minute particles filtered from a water current.” The filtratory mecha-
nism is that common to all the less-specialized Malacostraca, the
maxillae and the adjacent appendages forming the filter apparatus ;
the thoracic limbs furnish the motor power for producing currents of
water. In Hemimysis, according to Cannon and Manton, centripetal
streams of water are generated by the rotary movements of the
exopodites of the thoracic limbs, which currents pass between the
limb bases into the food channel below the body, where the water is
drawn forward principally by vibrations of the maxillae. In feeding
on large food masses, these writers say, the food is held by the thoracic
endopodites and consumed as the mysid swims. Such food includes
particles too large for filter feeding, and small animals, even Sagitta
worms longer than the mysid itself. The food is held beneath the
mouth parts by the third to eighth endopodites, properly oriented by
the long mandibular palps and the first and second endopodites, and
is bitten into by the incisor processes of the mandibles and the spi-
nous distal endites of the first maxillae.

Stomatopoda.—The stomatopod is a taxonomic misfit ; it is intro-
duced here because its mandibles are doubly articulated, though in
other respects they do not much resemble the doubly articulated jaws
of the decapods.

The mandibles of Squilla (fig. 7D, Md) hang vertically behind the
epistome and labrum from weak dorsal articulations (A, a) on the
doublure of the carapace, but they have strong ventral articulations
(c) with small condyles on the posterior margins of the long lateral
wings of the epistome (D, Epst) that embrace the labrum (Lm).
The lower end of each mandible is produced downward into a large,
tapering, toothed incisor process (A, B, imc), and from the base of
the latter, just below the epistomal articulation (c), there extends for-
ward a long molar process (mol) with a double row of teeth. In the
natural position of the jaws the ends of the incisor processes come
together with their teeth interlocked behind the mouth, and the molar
processes project straight forward into the large mouth cavity above
the labrum.

The mandibular musculature of Squilla consists of two anterior
muscles (fig. 7B, rAd, 2A), a single posterior dorsal muscle (P),
and a large ventral muscle (C, V’), the spreading fibers of which fill
the cavity of the mandible. The fibers of the ventral muscles of the
jaws do not arise from a supporting ligament, but are attached sepa-
rately on large lateral expansions of an apodemal arch arising from
premandibular invaginations between the epistome and the doublure

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 31

of the carapace (D, inv). A corresponding apodeme has not been
observed in any other crustacean.

Manipulation of the mandibles of a preserved specimen shows that
the only movement the jaws can make is a partial rotation on their
vertical epistomal hinges between the two points of articulation (fig.
7B, a,c). Since the molar processes project forward at right angles

Fic. 7.—Crustacea—Stomatopoda.

A, Squilla afinis Berthold, left mandible, lateral. B, Squilla panamensis
Bigelow, right mandible, mesal. C, Squilla affinis Berthold, right mandible
alee mesal. D, same, anterior part of body, showing left mandible
in place.

to the axes of the mandibles, a rotary movement that corresponds with
adduction in a doubly articulated horizontal mandible separates the
molar processes, and the opposite movement brings them together.
The movements have little effect on the incisor processes other than
that of rotation. The stomatopods are predaceous, and are said to
feed on small crustaceans, mollusks, and worms.
Decapoda—Stenopidea.—The mandibles of Stenopus hispidus
(fig. 8E), as those of Penaeus, hang from dorsal articulations (a)
on the carapace, and are articulated ventrally on the lateral wings of

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the epistome. The two jaws are connected by the large ventral adduc-
tors (V) united on a median ligament which is supported anteriorly
by a pair of suspensory branches (s/) attached on the dorsum of the
carapace (Cp). From the adductor ligament there extends posteriorly
a broad sheet of membranous tissue on which are attached the ventral
muscles of the maxillae, and which is itself supported by two pairs of
suspensory ligaments. This maxillary “fascia” is common to other
natantian decapods. On the anterior margin of the mandible are
inserted three dorsal muscles (C, A), all of which appear to be abduc-
tors. The posterior adductor (P) arises by a broad base on the cara-
pace (E) and is attached by a long tendon on the posterior margin
of the jaw at the base of the gnathal lobe. The fibers of the ventral
muscles (V) are apparently all adductors, since they show no evi-
dent differentiation into adductor and abductor groups.

Decapoda—Penaeidea.—Inasmuch as the mandibles of the deca-
pods differ very much in the several suborders, they will be described
separately in each group. There is no characteristic difference, how-
ever, in the structure and mechanism of the jaws as between those
forms classed as Natantia and those included in the Reptantia.

The mandibles of Penaeus (fig. 8A) lie almost in a transverse
vertical plane, but they are slanted somewhat forward, and hang
obliquely inward, so that the gnathal lobes come together below the
mouth. Each jaw is articulated dorsally (a) on the base of the inner
lamella of the carapace fold, and ventrally by a small process on its
anterior margin (A, B, c) with the narrow postantennal wing of
the epistome. The Penaeus mandibles are thus doubly articulated,
and rock lengthwise on the axes between the two articular points.
The base of each mandible is broadly oval and deeply concave (B) ;
the gnathal lobe is split into a flat, toothed incisor process (inc) and
a thick molar process (mol), both of which are turned mesally from
the end of the mandibular base (A). On the anterior margin of the
base of the jaw are attached three anterior dorsal muscles (B, IA,
2A, 3A), which, since they arise on the carapace dorsal to the man-
dible, and are inserted laterad of the mandibular axis, appear to be
all abductors. The opposed dorsal adductor (P) arises dorsomedially
on the carapace (A). The ventral muscles of the two mandibles
(A, V) are united by a strong median ligament, and most of their
fibers (z/’) are adductors. An anterior group of the ventral fibers
(2V), however, evidently functions as a ventral abductor, since it
is attached on the mandibular margin above the line of rotation.
This differentiation of the primarily adductor ventral fibers of the
mandible into adductor fibers and abductor fibers is an essential

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 33

feature of the jaw mechanism in the Reptantia, and is seen also in
such natantian forms as Lebbeus (D) and Macrobrachium (F), in
which the opposed anterior dorsal muscles (1A, 24) take on an adduc-
tor action.

Decapoda—Caridea.—The mandibles and the mandibular muscu-
lature of the caridean Pandalus danae Stimpson have been described
by Berkeley (1928), who shows that the mandibles and their mecha-
nism in this representative of the natantian decapods are the same
as in the Reptantia, and she follows the muscle nomenclature used
by Schmidt (1915) for Astacus. In the Caridea the mandibles take
a more horizontal position than in Penaeus and Stenopus, and in their
structure and mechanism they more closely resemble the jaws of the
Astacura.

The mandible of Macrobrachium latimanus (fig. 81) is a large,
strongly calcified boat-shaped structure, doubly articulated (a-c)
between the carapace and the epistome. The gnathal lobe is divided
into a broad, thick, bluntly toothed incisor process (inc), and a long,
expanding molar process (mol) strongly toothed on its truncate end.
The anterior margin of the base of the mandible, proximal to the
epistomal articulation (c), is somewhat elevated and bears a low,
flangelike apodemal ridge (Ap).

The musculature of the Macrobrachium mandible (fig. 8 F) in-
cludes four anterior muscles separated into a distal pair (14, 2A) and
a proximal pair (34). The two distal muscles are inserted on the
apodemal flange of the mandibular margin (I, Ap), laterad of the
hinge line of the jaw, and take their origins somewhat ventrally (F)
on the side of the carapace.. These muscles in Macrobrachium, there-
fore, are anterior adductors of the mandible, as they are in Pandalus
and in the Astacura (fig. 9 D, E, 1A, 2A), though they belong to the
original group of dorsal abductors. The double proximal anterior
muscle (fig. 8 F, 34) arises dorsally on the carapace and is an anterior
abductor; it is not given by Berkeley (1928) in Pandalus, but is
represented in Astacura (fig. 9D, E, 34). The posterior dorsal
muscle (fig. 8 F, P), attached distally on the mandible by a long thick
tendon, is the usual posterior adductor. The ventral muscles from the
opposite jaws are united by a thick, cylindrical median ligament (H,
Lg). On each side the ligament gives off dorsally three short branches
from which spreading groups of muscle fibers go to the carapace
(Cp). The ligament itself has a horizontally lamellate structure. The
lamellae, which are separated by interlamellar spaces, are distinctly
fibrillated and break up at their ends into bundles of fibers that soon
become striated muscle tissue. In Pandalus Berkeley (1928) notes

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

ANY
\\\

\\

SS ZZ

Fic. 8.—Crustacea—Decapoda: Natantia.

A, Penaeus duorarum Burkenroad, section of carapace, and suspended man-
dibles, anterior. B, same, right mandible and marginal muscles, mesal. C,
Stenopus hispidus (Olivier), left mandible and muscles, anteromesal. D,
Lebbeus groenlandicus (Fabr.), mandibles and muscles, anterior. E, Stenopus
hispidus (Olivier), section of carapace, and suspended mandibles, posterior.
F, Macrobrachium latimanus von Martens, left mandible and muscles, lateral
(anterior). G, Sclerocrangon boreas (Phipps), left mandible, ventral. H,
Macrobrachium latimanus yon Martens, right half of intergnathal ligament of
mandibles, with its suspensory muscles and mandibular adductor (V). I, same,
left mandible, dorsolateral.

a, a

NO; I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 35

that the ligament, or “fascia,” is supported on sternal apodemes. Most
of the ventral muscle fibers are attached within the cavities of the
mandibles and have an adductor function. On each side, however, a
broad sheet of dorsal fibers (F, 2’) is inserted on the marginal
flange of the mandible opposite the lateral adductors (1A, 2A), and
these fibers, therefore, constitute a ventral abductor muscle of the
mandible, as in Pandalus and Astacura, opposed to the lateral
adductors.

The mandibular musculature of Macrobrachium illustrates very
clearly how the primitive three muscles of the crustacean jaw (4,
P,V) may become differentiated into five groups of fibers with diver-
sified functions; only the posterior dorsal muscle (P) preserves its
integrity and its original function. It will later be seen that these
same muscles persist in the reptantian decapods and become adapted
to further changes in the mandibular mechanism.

The mandibles of Lebbeus (fig. 8D) resemble those of Macro-
brachium except in that the incisor processes (imc) are relatively
small and slender. The large molar processes (mol), as in Macro-
brachium, are turned toward each other at right angles to the axes of
the mandibles, so that with the rocking of the jaws on their axes the
opposed surfaces of the molars work upon each other, and, it should
be noted, they are turned outward by the adductor movement of the
mandible, and inward with the abductor movement. Each mandible
has at least two anterior dorsal muscles (D, 2A, 3A) ; it is probable
that a first muscle of the series was lost in dissection, since a well-
differentiated abductor set of ventral fibers (2V) is inserted on the
inner surface of an elevated part of the mandibular margin that in
Macrobrachium (F) gives insertion to two lateral adductor muscles
(1A, 2A). The relatively slender ligament of the ventral adductors in
Lebbeus is supported by a pair of dorsal branches attached directly
on the carapace.

In the caridean family Crangonidae the mandibles take on a highly
aberrant form, and evidently function in a manner quite different
from that of the ordinary decapod jaw. The mandible of Sclero-
crangon boreas (fig. 8G) is attached by its basal part in a wide mem-
branous area between the mouth and the carapace, but it has no direct
connection with either the carapace or the epistome. Its anterior end
is turned abruptly mesally as a long, slender gnathal lobe that ends
with a toothed expansion. The lobes of the opposing mandibles project
directly toward each other into the sides of a capacious preoral cavity
enclosed between the huge, 3-lobed labrum in front, and the long,
divergent paragnaths behind, which arise from a thick, semicircular

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

metastomal base (see Snodgrass, in press, fig. 11 F). Mandibles such
as these evidently can have no biting or chewing function; they must
be used for jabbing into a mouthful of food. The musculature of the
Sclerocrangon mandibles could not be determined from the material
studied.

Decapoda—Astacura.—The astacuran mandibles lie transversely
between the carapace and the mouth (fig. 9 A, Md), but they are
somewhat inclined downward and strongly slanted forward, so that
the gnathal lobes come together beneath and behind the labrum with
their toothed margins in apposition. Laterally each mandible is articu-
lated by a condyle (a) on the base of the inner lamella (Dbl) of the
carapace fold in the angle between the latter and the broad pleural
bridge of the maxillary segment (mxB) that limits the branchial
chamber anteriorly (brC). Mesally the mandible is articulated in a
socket (c) of the epistome at the side of the labrum, and furthermore,
it has a strong linear hinge (1) on the epistomal margin laterad of
the articular socket.

The mandible of Cambarus, or of other cambarine species, is some-
what quadrate in shape as seen from below (fig. 9B). Projecting
beyond the palpus as a direct continuation from the body of the man-
dible is the broad gnathal lobe (gn), and in front of the palpus rises
a large process that bears the epistomal articulation (c). On the
lateral angle of the base of the jaw is a cup-shaped knob (a) that ar-
ticulates with the carapace. The axis of rotation (a-c), therefore, is
strongly oblique between the two articular points; the long epistomal
hinge (/) falls in the plane of the axis. The movements of the man-
dible are thus strictly limited to movements of rotation, but the obliq-
uity of the axis line gives the gnathal lobe (guL) a wide swing on the
perpendicular (d) from the axis (a-c) ; in adduction the toothed mar-
gins of the opposing lobes come directly together.

The operation of the astacuran mandible depends largely on the
fact that the anterior margin of the jaw is produced into a wide,
triangular apodemal lobe (fig. 9 B, C, Ap) between the epistomal
hinge (/) and the lateral articulation (a). The apodeme is smaller
in Homarus (D) than in Cambarus or Astacus, but it is charac-
teristically more developed in the Astacura than in the natantian
decapods, in which, as in Macrobrachium (fig. 81, Ap) it is repre-
sented at best only by a slight elevation of the mandibular margin.
The apodeme attains its highest development in the Anomura and
Brachyura.

The musculature of the astacuran mandibles presents the same
functional fiber groups seen in Macrobrachium, but in the reptantian

JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 37

NO. I
decapods the ventral fibers no longer form an intermandibular muscle,
since they have become attached on the intermaxillary apodemes of
the ventral endoskeleton. The mandibular muscles of Astacus have

(VV Halll) S)-2Ant
‘Y Sy 1D)

Vy YICCSTT 3S
fg >

'
'
!
'
' \
U

Md Mth Mad

R

Fic. 9.—Crustacea—Decapoda: Astacura.

A, Cambarus longulus Girard, ventral surface of anterior part of body, with
C, same,

left mandible removed. B, Cambarus sp., right mandible, lateral.
D, Homarus americanus M. Edw., right mandible

right mandible, dorsomesal. ;
E, Cambarus sp., right mandible and muscles, mesal.

and muscles, mesal.
F, sectional diagram of astacuran mandibular mechanism.

been fully described by Schmidt (1915); those of Cambarus and
related genera (fig. 9 E) are the same as in Astacus, but in Homarus

(D) the relative size of some of the muscles is different.
The first two anterior dorsal muscles of the mandible in both

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Homarus and Cambarus (fig. 9D, E, 1A, 2A), as in Astacus, are
attached on the outer face of the mandibular apodeme (Ap) and pull
laterally from their origins on the carapace. These two muscles to-
gether, therefore, are termed by Schmidt the adductor lateralis of the
mandible. The third muscle of the anterior series (34), inserted
laterad of the apodeme, arises dorsally on the carapace and preserves
its original abductor function. It is the abductor minor of Schmidt,
and is also very small in Cambarus (E), but in Homarus it is a muscle
of large size (D, 3A). The adductor posterior (P) is a huge, conical
muscle arising by a broad base on the dorsum of the carapace anterior
to the “cervical” groove, with its fibers converging to a long, thick
tendon attached on the posterior margin of the mandible at the base
of the gnathal lobe (C, Pt). The principal group of ventral fibers in
Astacus and Cambarus (FE, 1V) spreads into the entire cavity of the
mandible, but in Homarus (D) the fibers form a relatively small
muscle in the anterior part of the mandibular cavity. These fibers
retain the primitive adductor function of the ventral muscles, and
constitute the adductor anterior of Schmidt. A second and quite dis-
tinct group of ventral fibers (D, E, 2V’) is attached on the inner face
of the mandibular apodeme, and thus forms a muscle antagonistic to
the lateral adductors (7A, 2A) attached on the outer face of the
apodeme. This muscle (2V) is termed by Schmidt in Astacus the
abductor maior, but in Homarus it is exceded in size by the “abduc-
tor minor” (D, 3A).

The mechanism of the astacuran mandibles will be readily under-
stood from the diagram, figure 9 F. Each mandible is hinged on the
lateral wing of the epistome at h. The mandibular apodeme (Ap)
is inflected above the line of the hinge, so that the muscles (1A, 2V’)
inserted on its opposite surfaces become respectively adductors and
abductors. The principal adductor power of the mandibles, how-
ever, must reside in the great posterior dorsal muscles (P), and in
the ventral muscles (1V) attached within each mandible below the
hinge line.

Decapoda—Palinura.—The mandibles in this group (fig. 10) are
variable in their form and structure, and at one extreme they take on
special features of the anomuran and brachyuran mandibles. The
palinuran mandibles, however, have one distinctive character, which
is the presence of a long, tapering articular process arising mesad of
the palpus, which fits into a deep notch between the labrum and the
epistome (see Snodgrass, in press, fig. 15 C). On the outer edge of
the articular process is the epistomal hinge of the mandible (fig. 10 A,
D, h), and at its apex the epistomal articulation (c). The gnathal lobe

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 39

is simple, its margin may be even (A) or toothed (D), but there is
no differentiated molar process. The mandibular musculature is the
same as in Astacura.

The mandible of Panulirus (fig. 10 A) has the general features of
the astacuran mandible, but the apodemal lobe (Ap) is greatly en-
larged and produced proximally so that it not only rises far above
the axis of the mandible (a-c), but projects somewhat beyond the
lateral articulation (a). The apodeme thus becomes a very effective

Fic. 10.—Crustacea—Decapoda: Palinura.

A, Panulirus argus (Latr.), right mandible, dorsal. B, Scyllarus americanus
(Smith), mandibles, ventral. C, same, right mandible, dorsal. D, Polycheles
tanneri Faxon, right mandible, dorsal.

lever for the abductor and adductor muscles attached on its opposite
sides.

In Polycheles (fig. 10 D) the mandible is more slender and elon-
gate ; its apodeme (Ap) is a small knob near the proximal end of the
anterior margin of the mandible, but the articular process (a) is turned
posteriorly to give leverage to the short apodeme.

The mandibles of Scyllarus (fig. 10 B, C) are still more slender
than those of Polycheles. The apodeme (Ap) is a large, expanded
lobe extended into the body cavity so far beyond the lateral articula-
tion (a) that it appears to be the outer end of the mandible itself,
with the articulation transposed to the posterior margin. The true

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

relation of the parts, however, is evident on referring back to Poly-
cheles (D), Panulirus (A), and the Astacura (fig. 9B). On the
other hand it will be seen that the scyllarid type of mandibles is still
more elaborated in the Anomura and the Brachyura.

Decapoda—Anomura and Brachyura.—The structure and mech-
anism of the mandibles are so much alike in the anomuran and brachy-
uran decapods that the mandibles of the two groups may be treated
together. Before taking up the mandibles, however, attention must
be given to the ventral skeleton on which the jaws are supported. In
the Astacura, as already noted, the mandibles are articulated laterally
on the inner walls of the carapace folds (fig. 9 A, a), and behind
them are the broad, horizontal pleural bridges (mxB) that connect
the sternal region of the second maxillary segment with the carapace.
In the Anomura, as seen in Aegla and Galathea (fig. 11), the maxil-
lary bridges are reduced to narrow bars (m+xB), and the mandibles
(Md), instead of being articulated on the carapace, as in Astacura
and most other Crustacea, have their lateral articulations (a) on the
mesal ends of the pleural bars close to the second maxillary foramina
(2Mx). As a consequence, the mandibles are much shortened. In
Petrolisthes eriomerus (fig. 12 A, B) the sclerotic bridges themselves
(mxb) do not reach to the carapace and appear as small lateral exten-
sions from the rims of the maxillary foramina carrying the mandibu-
lar articulations (a). In the Brachyura the maxillary bridges are
still more reduced; in Callinectes (fig. 14) for example, the only
sclerotic connection of the rim of the second maxillary foramen
(2M) with the carapace is a slender, tapering rod on the anterior
margin of the branchial chamber (brC), and the mandibles have
become articulated laterally on the anterior rims of the maxillary
foramina (fig. 14; fig. 15 D, H, a). Between the mandible and the
carapace there is now a wide space (fig. 14) occupied by a thin, semi-
membranous extension of the doublure of the carapace that extends
forward to the epistome (Epst) and forms the dorsal wall of the
pump chamber of the respiratory passage. In both the Anomura and
the Brachyura the mandibles retain the usual epistomal connections
(figs. 12 A; 15 C, F, G) including the mesal articulations (c) and
the marginal hinges (/).

Another character apparently peculiar to the anomurans and brachy-
urans is the presence of a pair of arms arising from the metastomal
plate of the ventral skeleton (figs. 11 A, B; 16D, H, ¢) that extend
forward along the folds at the sides of the mouth and support the
mandibles mesally at the base of the gnathal lobe (fig. 16H). In
Uca pugilator these mandibular props are wide sclerites at the sides

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 4I

Stl ——

I
/

\
1Mxpd 2Mxpd_ IIS

Fic. 11.—Crustacea—Decapoda: Anomura

|
lddltttttzz#7e

; = a SO,

Rois >

A, Aegla prado Schmitt, ventral skeleton of anterior body region, with
mandibles in place, exposed by removal of ventral folds of carapace. B, Galathea
californiensis Benedict, ventral surface of protocephalon and skeleton of gnathal
region, ventral folds of carapace cut off at 2.

Note in each species mandibles articulated at a on mesal ends of narrow
maxillary pleural bridges (m+rB).

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Fic. 12.—Crustacea—Decapoda: Anomura.

A, Petrolisthes eriomerus Stimpson, epistomal region and mandibles, dorsal
(interior) view, showing mandibles articulated at a@ on remnants of maxillary
bridges (mxb). B, same, right mandible, ventral. C, Galathea californiensis
Benedict, muscles of mandibular apodeme arising on carapace. D, same, right
mandible, dorsal. E, Pagurus pollicaris Say, left mandible in position of adduc-
tion, ventral. F, same, same mandible in position of abduction. G, same, distal
part of left mandible, showing articulation on epistome. H, Galathea californi-
ensis Benedict, right mandible and its skeletal supports, ventral.

NOs Er JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 43

of the mouth folds (fig. 16 E, t), each of which at its distal end
divides into a mesal branch that goes into the paragnath (Pgn), and
a lateral branch that supports the mandible by means of a small inter-
vening sclerite (E, H, e¢). The connection on the mandible is with
a process behind the base of the gnathal lobe (figs. 12D; 15C, D,
F, G; 16 A, B, G, H, d), a feature characteristic of anomuran and
brachyuran mandibles; in most cases the metastomal arm makes a
direct contact with the mandibular process. Inasmuch as the mandibu-
lar attachments on the metastomal arms are not in line with the man-
dibular axes of rotation, they cannot be regarded as true articulations ;
the arms merely furnish an extra support for the jaws, and evidently
they must be flexible in order to permit the normal movement of the
latter.

The most important modification of the mandible in the Anomura
and Brachyura is the progressive elongation of the mandibular apo-
deme in line with the body of the jaw, by which the apodeme becomes
an increasingly efficient lever for abduction and adduction of the
gnathal lobe.

Among the Anomura, the mandibles of Petrolisthes (fig. 12 A, B)
and of Pagurus (E) resemble those of the palinuran Polycheles (fig.
10 D) in that the apodeme (AP) arises from the anterior margin of
the mandible a short distance mesad of the lateral articulation (a).
The apodeme carries loosely attached to its apex a large thin plate
(fig. 12 E, mp), on the opposite sides of which are inserted the first
and second anterior muscles of the mandible (C, rd, 2A), which
arise laterally on the carapace (Cp). Antagonistic to these muscles
is a large group of ventral fibers attached posteriorly on the apodeme
itself, as shown in the brachyuran Callinectes (fig. 16 A, C, 2V). The
opposing sets of muscles, therefore, alternately pulling in opposite
directions on the apodeme, rotate a mandible such as that of Pagurus
(fig. 12 E, F) on its lengthwise axis (a-c), the mandible being articu-
lated mesally on the epistome (G).

By a change in the position of the apodeme relative to the lateral
articulation of the mandible, the anomuran mandible may take on the
type of structure and mechanism that is particularly developed in the
Brachyura. In Galathea, for example (fig. 12 D, H), the mandibular
apodeme (Ap) is produced laterally far beyond the articulation (a)
so that it appears to be a proximal extension of the body of the
mandible itself. The apodeme thus takes a position more nearly
perpendicular to the oblique rotation axis of the jaw (H, a-c), and
hence acquires a mechanical advantage in being almost directly op-
posed to the gnathai lobe.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Again, among the Anomura, mandibles occur that show a retrogres-
sive type of structure. The weak, fragile mandibles of Callianassa
major, for example (fig. 13 A), are well developed at their mesal
ends, where they are strongly articulated and hinged on the epistome,
but proximally they taper out into slender arms that do not quite
make contact with the ventral skeleton. The mandibular apodeme is
merely a small, angulated rod (B, Ap) arising near the end of the
slender shaft. In the sand crab, Emerita talpoida, the mandibles (D)
somewhat resemble those of Callianassa, but they are of minute size

Fic. 13.—Crustacea—Decapoda: Anomura.

A, Callianassa major Say, mandibles, labrum and epistome, ventral. B, same,
proximal end of mandible, with small apodeme. C, Emerita talpoida (Say),
mouth region and mandibles, ventral (labrum normally turned forward). D,
same, right mandible, dorsal.

and are immovably fixed on the epistome (C). The small, thin
gnathal lobes of the opposite jaws are widely separated, and apparently
only the palpi can be functional organs. Since the food of Emerita
consists of particles gathered from the water on the large, feathery
antennae, this highly specialized inhabitant of sand beaches evidently
has no use for functional jaws.

Among the Brachyura there are forms, such as Dromidia (fig.
15 B), in which the mandibular apodeme (Ap) projects as a short
arm from the anterior margin of the mandible, but it is almost per-
pendicularly opposed to the axis of rotation. In most of the brachy-
urans, however, the apodeme is extended beyond the lateral articula-
tion directly in line with the body of the jaw (figs. 15 D-H; 16A,
B, G, H, Ap). The lateral point of articulation (a) thus comes to lie

NO. I JAWS OF MANDIBULATE ARTHROPODS—-SNODGRASS 45

behind the base of the apodeme, and the axis of rotation becomes
increasingly oblique (fig. 15 F, G, a-c) instead of longitudinal. As
if to bring the axis as near as possible to a transverse position between
the apodemal lever and the gnathal lobe, the articular process becomes
elongate and extended mesally (G), and finally it is supplemented by
a small articular sclerite, as in Cancer (fig. 15 H), Callinectes (fig.
16 A), and Uca (fig. 16 H). The evolution of the brachyuran jaw,

YN»

Fic. 14.—Crustacea—Decapoda: Brachyura. Callinectes sapidus Rathbun.

Ventral view of anterior part of body with mesal lobes of carapace cut away
before bases of chelipeds to expose the ventral skeleton of the maxillary and
maxilliped region, after removal of the appendages, showing the mandibles
articulated laterally on the anterior rims of the second maxillary foramina (see
also fig. 15D, H).

therefore, is toward a more efficient leverage action of the apodeme
on the gnathal lobe. The axis of the jaw, however, can never become
completely transverse in position because of the intervention of the
first maxilla between the mouth and the articular point of the mandible
on the ventral skeleton (figs. 15 H; 16 H, rM-x).

The muscles of the brachyuran mandibles have been described by
Borradaile (1922) in Carcinus maenas, and by Cochran (1935) in
Callinectes sapidus, but each of these writers overlooked a small group
of fibers representing the ventral adductor of Astacura (fig. 9 D, E,
1V). The musculature and mechanism of the crab mandible is well
shown in Callinectes (fig. 16A, C) and Cancer (B). The great

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Fic. 15.—Crustacea—Decapoda: Brachyura.

A, Dromidia antillensis Stimpson, epistome and labrum, ventral. B, same,
mandibles, ventral. C, same, right mandible, mesal. D, Calappa flaminea
(Herbst), right mandible, mesal. E, Callinectes sapidus Rathbun, mandibles,
anterior. F, Ovalipes ocellatus (Herbst), right mandible, mesal. G, Ocypoda
albicans Bosc, right mandible, mesal. H, Cancer borealis Stimpson, left man-
dible, ventral.

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 47

Fic. 16.—Crustacea—Decapoda: Brachyura.

A, Callinectes sapidus Rathbun, right mandible, mesal. B, Cancer borealis
Stimpson, right mandible, mesal. C, diagram of mandibles and their muscles
in Callinectes sapidus Rathbun, ventral. D, Libinia emarginata Leach, mouth
region, ventral. E, Uca pugilator (Bosc), mouth region, ventral. F, Libinia
emarginata Leach, metastomal prop of left mandible, ventral. G, Uca pugilator
(Bosc), right mandible, mesal. H, same, right mandible and its skeletal con-
nections, dorsal (internal view).

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

elongation of the mandibular apodeme (Ap) in these forms has car-
ried the first and second anterior dorsal muscles (B, C, 14, 2A)
attached on the apodeme far beyond the lateral mandibular articula-
tion (a). By the same process, the third muscles of the anterior series
(C, 34) comes to be attached on the mandible (by a long tendon)
below the base of the apodeme (A, B, 3A4t). The two apodemal
muscles are regarded by Borradaile and by Cochran as a single muscle,
as indeed they are functionally, but they clearly represent the two
lateral muscles of the apodeme in Astacura (fig. 9 D, E, 1A, 2A).
The third anterior muscle of the crab (fig. 16 C, 34) is erroneously
grouped by Cochran with the posterior adductor of the jaw. The
ventral muscles of the crab mandible arise on the endosternal skele-
ton, and consist of two groups of fibers. A small mesal group (fig.
16 A, B, C, rV), attached within the cavity of the mandible, repre-
sents the principal mass of ventral fibers in other forms, but in
the crab these fibers form only an unimportant ventral adductor of
the jaw. The other, much larger group of ventral fibers (2V) is
attached mesally on the upper margin of the mandibular apodeme.
These fibers, therefore, constitute the principal abductor of the gnathal
lobe, and are directly opposed by the lateral muscles (1A, 2A) at-
tached on the apex of the apodeme, which are the principal adductors.
In brachyurans such as Ovalipes (fig. 15 F), Callinectes (fig. 16 A),
and Cancer (B), in which the axis of the jaw (a-c) becomes strongly
oblique across the body of the mandible, the decapod type of mandible
attains its most efficient mechanism for abduction and adduction of
the gnathal lobe.

The crab, in feeding, Borradaile (1922) says, cuts or tears its
food into small pieces, which it swallows without chewing. The food
is seized by the chelae, and by them placed between the jaws, perhaps
assisted by one or more of the legs. If the food is soft, pieces are
bitten off by the mandibles ; otherwise it is securely held in the grip
of the mandibles, while the chelae or the third maxillipeds pull on it
until a fragment comes off in the jaws, which is then swallowed with-
out chewing. The “molar” surfaces of the mandibles were not ob-
served to have any grinding action on the food. The mechanics of
the crab mandible would indicate that the jaws are merely a pair of
very efficient pincers, but, as in the decapods generally, they have
little capacity for grasping food directly.

Tanaidacea, Isopoda, Amphipoda.—In these three groups of
malacostracan Crustacea we find a new mechanical principle devel-
oped in connection with the mandibles, which converts the latter
again into a pair of jaws swinging transversely against each other,

NO: L JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 49

but now on a doubly articulated axis instead of on a single dorsal
point of suspension. The jaws thus acquire a strong grasping and
biting action.

The change in the mandibular mechanism involves three correlated
structural modifications. The mandible itself is altered in shape by a
shift of the gnathal lobe from a position in line with the length of
the jaw, as in the decapods, to one perpendicular to the axis (fig. 17 B,
E, G, Md). This change in the form of the mandible, to be effec-
tive, must be accompanied by a change in the position of the mandible
on the head. The mandible still keeps its cranial and epistomal articu-
lations (a, c), but, in order to swing transversely, the axis between the
two articular points, instead of lying crosswise on the under side of
the head, has taken a longitudinal position on the lateral margin of
the cranium. Finally, to accommodate the anterior position of the
epistomal articulation, it is necessary that the epistome itself should
have a frontal position on the head (A, Epst). The whole cranial
structure in the Tannaidacea, Isopoda, and Amphipoda, therefore, is
quite different from that in any other crustaceans, but it is duplicated
among the insects with doubly articulated jaws working transversely.

In the Tanaidacea (fig. 17 C) the mandible is hinged on the side
of the cranium between the usual articular points (a, c), but the line
of the hinge is strongly declivous from behind forward, and the
gnathal lobe, therefore, is but little deflected from the body of the
mandible. Each lobe consists of a long, toothed incisor process, and a
large molar process arising at the base of the incisor. Palpi are present
in some genera, but are absent in Tanais.

In Isopoda and Amphipoda with typical biting and chewing jaws
(fig. 17 A, Md) the mandibular axis (B, a-c) becomes horizontal
along the edge of the cranium, and the gnathal lobe (gnL) projects
downward from the anterior part of the mandible at right angles to
the axis. The lobes of the opposing jaws, therefore, open and close
directly against each other (D) in the space below the mouth between
the labrum in front and the paragnaths behind.

The musculature of the amphipod mandible as seen in Gammarus
(fig. 17 H) has been described by Borner (1909), whose chief inter-
est was in showing that it is carried over into the orthopteroid insects.
On the other hand, the mandibular musculature of both the amphipods
(E) and the isopods (F) is directly comparable with that of the
Branchiopoda, Nebalia, and Anaspides. The anterior rotator of the
primitive mandible becomes a dorsal abductor (E, F, G, A), the
posterior rotator a large dorsal adductor (P) ; the ventral muscle (V)
retains its original adductor function, but the fiber bundles from the

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Fic. 17.—Crustacea—Tanaidacea (Tanais), Amphipoda (Orchestoidea,
Talorchestia, Gammarus), and Isopoda (Ligyda).

A, Orchestoidea californiana (Brandt), head, lateral. B, Talorchestia longi-
cornis (Say), left mandible and adjacent parts of head, lateral. C, Tanais
carolint M. Edw., head, lateral. D, Talorchestia longicornis (Say), mouth
region, ventral, with mandibles in abduction. E, same, right mandible
and muscles, mesal. F, Ligyda exotica Roux, left mandible and muscles,
anterior. G, Orchestoidea californiana (Brandt), lower part of inner wall of
right side of cranium, with head apodeme of same side, and right mandible
in place. H, Gammarus locusta (L.), outline of head with mandibles, posterior,
showing head apodemes united in a transverse bar (Ap).

a

INO: E JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 51

two jaws are separately attached on head apodemes, and not united
by a median ligament. In the isopods the head apodemes arise from
the intermaxillary sternal brachia (see Snodgrass, in press), and thus
have the same position morphologically as the first sternal apodemes
of the decapods, though they take on very different forms.

The head apodemes of the amphipods arise from grooves (fig. 17 A,
inv) between the lower posterior parts of the cranium and the plate
(VS) that carries the first maxillipeds. Since this plate can hardly
be interpreted otherwise than as the sternum of the first maxilliped
segment (or possibly as the base of the maxillipeds themselves), the
apodemes appear to be intersegmental inflections between the second
maxillary segment and the first maxilliped segment. Further evi-
dence that the amphipod head apodemes (G, AP) are postmaxillary
and not intermaxillary is seen in the fact that the postmaxillary arms
of the head sternum (pmB) are directly continuous with the bases of
the apodemes. The apodemes of the amphipod head, therefore, rep-
resent the second pair of potential intersegmental apodemes. In the
decapod Cambarus there are no apodemes between the maxillary and
first maxilliped segments, the second developed pair of sternal apo-
demes being between the first and second maxilliped segments, and
the first pleural apodemes between the second and third maxilliped
segments.

The postmaxillary apodemes of the amphipod Orchestoidea (fig.
17 G, Ap) have broad bases from each of which a slender arm curves
forward and downward, and gives attachment to the ventral adduc-
tor muscle (V) of the corresponding mandible. In Gammarus (H)
the two apodemes are united in a thick bar (Ap) that extends
through the back of the head, and the mandibular adductors (V) are
attached on anterior branches of the bar. This structure in Gammarus
suggests the tentorial bridge of the insect head, and is termed the
“tentorium” by Borner (1909), who notes the attachment of the
mandibular muscles on it, but does not discuss the nature of the
apodeme, except to say that it is invaginated on each side behind the
maxillary bases. The posterior bridge of the insect tentorium cer-
tainly does not arise behind the second maxillae; it represents a pair
of intermaxillary apodemes. The amphipods, therefore, furnish an-
other example of the apparent indifference of the ventral mandibular
adductors as to what apodemal structures they become attached after
severing connections with each other. We have thus far noted their
attachment on premandibular apodemes in the stomatopods, on inter-
maxillary apodemes in the decapods and isopods, and on postmaxil-

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

lary apodemes in the amphipods; other examples of inconsistency will
be seen in the myriapods and insects.

I]. CHIEOPODA

The mandibles of the chilopods seem to initiate a new line of jaw
evolution, since their structure cannot be matched anywhere among
the Crustacea, and, on the other hand, it appears to lead into the
type of mandible characteristic of the Diplopoda and Symphyla,
while some of its lesser features are repeated in the entognathous
apterygote hexapods.

The mandibles of a centipede (fig. 18 A) are elongate, widened
anteriorly, narrowed posteriorly, and lie horizontally against the under
side of the head with their axes convergent toward the mouth lying
above the approximated gnathal lobes, which latter are strongly
toothed on their opposed margins. In the normal condition the anterior
parts of the mandibles are mostly concealed above the edge of the
labrum, and are covered below by the broad palps of the first maxil-
lae. The tapering posterior end of the jaws are deeply sunken into
pouches of the membranous ventral wall of the head invaginated at
the sides of the first maxillae, and extended above the basis of the
second maxillae.

Each mandible has an anterior and a posterior point of articulation.
The anterior articulation (fig. 18 A, c) is by means of a knob or
hook on the lateral surface of the jaw, some distance back from the
anterior end, which is loosely held in the notch (F, g) between the
epipharyngeal arm (f) and the hypopharyngeal arm of the corre-
sponding premandibular sternal sclerite of the head (Flt) that sup-
ports the hypopharynx. The posterior articulation (A, a) is at the
rear extremity of the mandible, where the latter is attached to the
end of a slender rod (mdr) in the wall of the enclosing pouch, which
extends posteriorly and mesally from the cranial margin ; in Lithobius
the rod arises from a small marginal plate of the cranium (m). The
posterior attachment of the mandible (a) on the rod is evidently the
primary dorsal articulation of a primitive mandible on the mandibular
tergum, which in the chilopods is intermediated by the articular rod
as a result of the invagination of the mandible. The anterior articu-
lation, on the other hand, is clearly secondary; it is merely one of
contact serving to hold the mandible against the head, and has no
relation to the anterior articulation of a doubly articulated crustacean
or insect jaw. The different positions of the two articular points rela-
tive to the axis of the chilopod mandible brings the hinge line of the

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 53

Ephy

al \p z

Mth Hphy

Fic. 18.—Chilopoda.

A, Lithobius sp., mandibles and muscles, fultural sclerites with cranial con-
nections, and head apodemes, ventral. B, Scutigera coleoptrata (L.), right
mandible, dorsal. C, Lithobius sp., right mandible and muscles, dorsal. D,
Scutigera coleoptrata (L.), intergnathal ligament supported on ends of head
apodemes, dorsal. E, same, right mandible and muscles, mesal. F, Lithobius
sp., hypopharynx, fultural sclerites, and head apodemes, ventral. G, a geophilid,
hypopharynx and mandibles, ventral.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

jaw (A, a-c) almost parallel with the midline of the head. In action,
therefore, the mandibles rotate lengthwise, and the toothed edges of
the gnathal lobes separate ventrally in abduction and come together
in adduction.

In Scutigeromorpha, Lithobiomorpha, and Scolopendromorpha the
broad anterior part of the mandible bearing the articular process
projects forward and mesally as a free gnathal lobe (fig. 18 A, B),
armed on its distal margin with strong teeth, and fringed with brushes
of hairs. On the dorsal surface of the mandible (C) the sclerotized
wall of the lobe is separated from the basal part of the jaw by an
oblique line of flexibility proximal to the articular process (c). At-
tached mesally on the base of the lobe are two large muscles (C, E),
one (J), arising in the base of the mandible, the other (1d) by
widely spreading fibers on the dorsal wall of the cranium. The gnathal
lobe of the jaw in these three chilopod groups, therefore, would
appear to have an independent movement of flexion on the base of
the organ; the huge cranial muscle (14) otherwise would be merely
an abductor or retractor of the mandible as a whole. A live specimen,
however, refuses to demonstrate any action whatever of its jaws.
The cranial muscle of the gnathal lobe may be supposed to be derived
from the anterior dorsal musculature (A) of a generalized mandible ;
the intramandibular muscle (/) is comparable to the stipital flexor
of the lacinea of an insect maxilla, but it appears to have no antecedent
in the mandible. This same musculature of the gnathal lobe, however,
is carried over into the Diplopoda and Symphyla, in which the jaw
lobe becomes freely movable by a definite articulation on the base of
the mandible.

In the Geophilomorpha the mandibles are extremely small and
weak, relative to the length of the body, but they have the general
shape of the jaws in the other groups. The mandibular base is a
slender, curved bar, from the end of which the gnathal lobe expands
mesally against the side of the hypopharynx (fig. 18G, Md). In
unidentified specimens examined by the writer, however, there is no
line of flexibility separating the lobe of the mandible from the base,
and no cranial muscle of the lobe could be discovered, such as that
so highly developed in the rest of the chilopods, and which is charac-
teristic also of the symphylids and diplopods. On the other hand
there is a striking similarity between the geophilid mandible and the
mandible of the pauropods (fig. 19 A, Md).

The basal musculature of the mandible of Lithobius (fig. 18 C)
includes a dorsal muscle (24), perhaps a rotator, attached on the
posterior end of the mandible, and three groups of ventral fibers (V).

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 55

The fibers of the first ventral group (zV) arise on the head apodeme
of the same side (A, hAp), those of the second group (2/) appear
to be continuous from one jaw to the other, but they are attached on
a ligamentous bridge (A, Lg) between the apodemes; these two sets
of fibers are adductors of the mandible. The third group of ventral
fibers (3) arises on the base of the apodeme, and is attached on the
posterior end of the mandible; these fibers constitute a protractor
muscle, the cranial muscle of the gnathal lobe (zA) being evidently
a retractor of the mandible as well as a flexor of the lobe.

In the Scutigeromorpha the mandible has the same musculature
(fig. 18 E) as in Lithobius, but all the adductor fibers of the ventral
muscles are attached on a broad sheet of ligamentous tissue (D, Lg)
supported from below on the posterior ends of the head apodemes
(hAp), and suspended from the dorsal wall of the cranium by suspen-
sory ligaments (s/). The intergnathal ligament of Thereuonema tuber-
culata (Wood) has been described in detail by Fahlander (1938),
who shows by its muscle connections that it pertains to the three
gnathal segments of the head, and is correspondingly divided into
three parts by median foramina. The ligament is a nonchitinous tis-
sue which Fahlander says has a fibrillar structure and contains cell
remnants ; this and other similar tissues of the chilopods he contends
are formed of an “endoskeletal substance” generated from the inner
surface of the epidermis. A comparison of the intergnathal ligament
of the scutigeromorph chilopods (fig. 18 D) with the similar ligament
in lower Crustacea (fig. 5 E, F) can scarcely leave any doubt of the
identity of the two structures. As in the higher Crustacea, the liga-
ment in the chilopods has become supported from below on ventral
head apodemes. In Lithobiomorpha the ligament is reduced to a
bridge between the apodemes (fig. 18 A, F, Lg) and some of the
ventral fibers of the jaws have become attached on the apodemes
themselves ; in Scolopendromorpha the bridge is still narrower; in
Geophilomorpha it is eliminated, all the ventral muscles of the man-
dibles (G, V) are attached directly on the head apodemes (HAP).

The head apodemes of the chilopods are cuticular ingrowths from
the mesal ends of a pair of premandibular sternal sclerites of the
head (fig. 18 A, F, Fit) that extend transversely from the margins
of the cranium to the sides of the hypopharynx (F, Hphy). They
are the “Kommandibulares Geriist’”’ of German writers, so named
because the mandibles loosely articulate on their mesal parts, but
since the sclerites appear to support the hypopharynx, the writer (in
press) has termed them the hypopharyngeal fulturae. It is clear that
the head apodemes of the Chilopoda, being premandibular in origin,

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

can have no homology with the postmandibular head apodemes of
Crustacea, though the same muscles come to be attached on them in
both groups, showing that the apodemal support of the intergnathal
ligament, and finally of the muscles, is secondary. On the other hand,
head apodemes clearly homologous with those of chilopods are present
in the diplopods, pauropods, and symphylans, and possibly in the
insects. Fahlander (1938) calls the interapodemal ligament of the
chilopod head the “tentorialkérper,” but, being a nonchitinous struc-
ture, the ligament can have no homology with the central plate of
the insect tentorium, since the latter is entirely a cuticular structure.

III. PAUROPODA

In the general structure of the head, the presence of a single mouth-
part appendage, the gnathochilarium, behind the mandibles, and in the
anterior position of the genital outlets, the pauropods appear to be
closely related to the diplopods. In common with the chilopods and
some of the diplopods, the pauropods have premandibular sclerites
of the ventral head wall (fig. 19 A, Flt) attached laterally on the
cranium and mesally on the hypopharynx. These sclerites are de-
scribed in Pauropus silvaticus by Tiegs (1947) as suspensorial scle-
rites of the head apodemes, or hypopharyngeal apophyses (hAp),
which arise from their inner ends. The head apodemes of the pauro-
pods, as shown by Silvestri (1902) and by Tiegs (1947), are long
slender arms extending posteriorly through the head into the first
body segment, each arm giving off a lateral branch attached to the
occipital margin of the cranium. The pauropods thus show unques-
tionably that they belong to the “myriapod” series of arthropods.
The pauropod mandibles resemble the chilopod mandibles in that
they have a longitudinal position against the under side of the head,
and the long tapering bases are invaginated into the head (A, Md) ;
the gnathal lobes are direct extensions from the bases of the ap-
pendages. As noted by Hansen (1930), therefore, the pauropod
mandibles differ strongly from the mandibles of both the Diplopoda
and the Symphyla, in which the gnathal lobes are independently
movable on the mandibular bases.

The mandibles of Allopauropus brevisetus, as shown by Silvestri
(1902), are simple elongate organs (fig. 19 B, Md), the wide taper-
ing bases of which are sunken into the head, and the free distal parts
armed mesally with rows of slender teeth. The mandibles of Pauropus
silvaticus (A, Md), as illustrated by Tiegs (1947), have a striking
resemblance to the mandibles of a geophilid centipede (fig. 18G).

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 57

Each mandible of Pauropus, Tiegs says, consists “of an unsegmented
piece of chitin, whose base is prolonged into an apodeme, a long
curved blade of chitin, which extends far back into the cavity of
the head.” The mandibular “apodeme” is said to be attached medially
by a fibrous ligament to the head apodeme, and laterally by a similar
ligament (not shown in the figure) “to the wall of the head just to
the rear of the pseudoculus.” As already noted, the cranial rod sup-
porting the mandible is characteristic of the invaginated jaws of the
chilopods ; it will be met with again in the Collembola and Protura.
In these forms the “ligament” is a sclerotic rod in the membranous
wall of the containing pouch, and it seems probable that the mandible

Fic. 19.—Pauropoda.

A, Pauropus silvaticus Tiegs, outline of head, ventral, showing right mandible
and head apodemes (adapted from Tiegs, 1947). B, Allopauropus brevisetus
Silvestri, head and first body segment, ventral (from Silvestri, 1902).

of Pauropus is likewise simply invaginated and connected in the same
way with the cranium. However, it is most surprising to find that
the pauropod mandible is so entirely different from the diplopod
mandible, and so nearly duplicates the mandible of a geophilid centi-
pede. What significance, if any, this fact may have as to the taxonomic
position of the Pauropoda the writer leaves to the phylogenists.

The mandibular musculature of Pauropus as described by Tiegs
consists of four groups of fibers, two of which are dorsal in origin,
and two ventral. The two dorsal muscles arising on the roof of the
head are attached anteriorly and posteriorly on the base of the man-
dible. A group of obliquely transverse adductor fibers arises on the
corresponding head apodeme. The fourth muscle, a ventral protractor,
has its origin on the suspensory plate (fultura) of the apodeme, and
is inserted on the posterior end of the mandible. It is noteworthy

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

that there are no special muscles of the gnathal lobe, such as are
present in those myriapods that have a flexible or articulated gnathal
lobe, and which appear to be absent also in the geophilids.

IV. SYMPHYLA AND DIPLOPODA

The symphylids and the diplopods are two groups of progoneate
myriapods that have little likeness in their body structure, but in
the structure of the head and particularly in that of the mandible
they have features in common that set them off from all the other
arthropods, and would seem to link them with each other. The man-
dible in each group consists of a basal plate implanted on the side of
the head (fig. 20 A, B, mdB), where it is but little movable, and of a
gnathal lobe (gnL) articulated on the base, and freely movable by
its own muscles. The musculature of the lobe is the same as that of
the flexible gnathal lobe of a chilopod mandible. It might be supposed,
therefore, that the symphylid-diplopod jaw has been evolved from the
more simple jaw of the chilopods; but, aside from the musculature,
the mandibles of the two groups are widely different. The gnathal lobe
of the symphylid-diplopod mandible and its musculature have a curi-
ous resemblance to the lacinial lobe and musculature of an insect
maxilla, but since the symphylids have both first and second maxillae
in addition to the mandibles, there can be no question of the identity
of their jaws with the mandibles of other arthropods.

The mandibles of Symphyla are somewhat simpler than those of
the diplopods. The elongate basal plates (fig. 20 B, mdB) lie longi-
tudinally on the sides of the head, separated from the deeply angulated
cranial margins above them by wide membranous spaces containing
the head spiracles (Sp). The flattened gnathal lobes (G, gnL) project
forward, and are flexible in a horizontal plane. Each lobe is articu-
lated by the outer angle of its base on the end of the supporting plate.
On the dorsal side is a small process (c) that bears against a sclerotic
ridge of the epipharyngeal surface above it, but does not form a true
articulation, and is suggestive of the similar structure in the chilopods.
The biting edge of the gnathal lobe is strongly toothed, and bears a
small dentate plate (Jm), or so-called “lacinia mobilis,” flexibly in-
serted between the fixed teeth on both sides of it. Attached on the
mesal angle of the gnathal lobe is the tendon of the flexor, or adductor,
muscle, a huge bundle of convergent fibers (14) from the posterior
part of the cranium. The musculature of the basal plate includes a
second dorsal (primitively anterior) muscle (24), arising on the
cranium, and two ventral adductors (zV, 2V), both of which arise on

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 59

Fic. 20.—Symphyla (Scutigerella) and Diplopoda (Fontaria, Thyropygus).

A, Fontaria virginiensis (Drury), head, lateral. B, Scutigerella immaculata
(Newp.), head, lateral. C, Fontaria virginiensis (Drury), left mandible, with
cranial flexor muscle of gnathal lobe, ventral. D, Thyropygus sp., right mandible
and muscles, dorsal. E, Fontaria virginiensis (Drury), head apodeme (hypo-
pharyngeal apophysis) of left side, and attached muscles, ventral. F, same,
intergnathal ligament and its muscles, dorsal. G, Scutigerella immaculata
(Newp.), outline of right half of head, and right mandible with muscles, dorsal.

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

the corresponding head apodeme (iAP), there being no intermandibu-
lar fibers in the symphylids or any connecting ligament between the
apodemes. The head apodemes, or hypopharyngeal apophyses, are
invaginated at the side of the hypopharynx, but supporting plates, such
as the premandibular fultural sclerites of the chilopods, pauropods,
and some diplopods are absent in the Symphyla. Since the symphylids
are exceptional among the myriapodous groups in lacking fultural
sclerites, and have no intermandibular muscle fibers, they are probably
to be regarded as specialized by the loss of these features, though in
other respects they are fairly generalized.

In Scutigerella immaculata (fig. 20G) the anterior fibers of the -
adductor musculature of the mandible form a large, fan-shaped muscle
(zV) arising on a small lateral lobe of the base of the apodeme and
spreading to most of the length of the basal plate of the mandible.
The fibers of the second muscle (2V) arise by a wide base on the
apodeme, and converge to a narrow insertion on the posterior part
of the mandibular plate. The same musculature is given by Tiegs
(1940) for the mandible of Hanseniella agilis.

It is in the Diplopoda that the arthropod mandible attains its most
remarkable development. The base of the organ in this group is a
large plate on the side of the head (fig. 20 A, mdB) occupying the
entire space between the emarginate edge of the cranium and the
enathochilarium, and is usually divided by a groove into a proximal
and a distal part. The plate, in fact, is virtually a part of the head
wall, but the attachment on it of the usual mandibular muscles shows
that it is truly the base of the mandible. The basal plate of a dead
specimen appears to be but little movable, and a live specimen, when
handled, keeps its jaws so tightly closed that no evidence of their
action can be obtained. The great size of the muscles inserted on
the mandibular plates (fig. 20 D), however, attests that these muscles
must have some important function. The large gnathal lobe of the
mandible, being freely movable on the basal plate and strongly muscu-
lated (C, D, gnL), becomes the functional jaw of the animal. It is
armed at the apex with large teeth flexible at their bases, and on the
mesal side is a variously developed molar surface. The principal
muscle of the lobe is a huge cranial flexor (14) composed of spread-
ing fibers arising on the dorsal wall of the head and inserted on the
mesal angle of the base of the gnathal lobe by a thick tendon. A
smaller intramandibular flexor, present at least in some forms (D, J),
has its origin within the basal plate of the mandible.

The musculature of the basal plate, as above noted, includes the
usual dorsal and ventral muscles of a typical mandible. Inserted on

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 61

the dorsal (anterior) margin is a thick muscle (fig. 20D, 24) from
the dorsal wall of the head, corresponding with the anterior rotator of
a primitive mandible; a ventral (posterior) rotator, however, appears
to be absent in the Diplopoda, as also in the Chilopoda and Symphyla.
The true ventral muscles include an anterior adductor (r1V) arising
on a thick intergnathal ligament (Lg), and a posterior group of ad-
ductor fibers (2V) arising on the head apodeme. In Fontaria vir-
giniensis the intergnathal ligament (F, Lg) gives off on each side a
large conical mass of fibers (zV) into the anterior part of the man-
dibular base, and branches posteriorly into two slender arms from
which arise smaller groups of fibers (3V) inserted in the posterior
parts of the mandibular bases, while a pair of short anterior branches
support small fiber groups (wv) inserted within the gnathal lobes.
From the bases of the anterior branches groups of slender fibers
(1) go to the under surface of the stomodaeum. The second large
bundle of adductor fiber in Fontaria (E, 2V) arises on the head apo-
deme (HAP) and spreads into the middle part of the basal mandibular
plate. The intergnathal ligament of the diplopods is free from the head
apodemes below it, but the muscle fibers arising from it are evidently
the first groups of adductor fibers in Symphyla (G, rl’), which have
become attached on the apodemes.

Fully developed premandibular ventral sclerites attached laterally
on the cranial margins are present in some diplopods as in the chilo-
pods and pauropods, but in most forms the sclerites are reduced to
small plate at the sides of the hypopharynx, from which the head
apodemes arise. The apodemes of Fontaria (fig. 20E, hAp) taper
posteriorly and dorsally in the head, and their apices are attached
to the central discs of the organs of T6mdsvary (A, OT). Two small
muscles from the dorsal head wall are inserted on each apodeme
(E, i, 7), so that the apodemes appear to act as levers possibly effect-
ing an elevation of the hypopharynx.

V. THE ENTOGNATHOUS APTERYGOTE HEXAPODS

The wingless, 6-legged arthropods included under this heading are
the Protura, the Collembola, and the Diplura (or Dicellura). They
are usually classed as insects along with the Thysanura and Pterygota
but are here treated as a separate group of hexapods because, in addi-
tion to being entognathous, they have head characters that set them
apart from all the other insects, while their mandibles have no counter-
parts among the Thysanura or Pterygota, either in structure or mecha-
nism, and show certain features suggestive of the mandibles of the

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

chilopods. The mandibles together with the maxillae lie horizontally
on the under side of the head, where their basal parts are enclosed in
gnathal pouches formed by union of the labium beneath them with
the lateral edges of the cranium. Because of the horizontal position
of the mouth parts, the primitive anterior surfaces become dorsal, the
posterior surfaces ventral.

A characteristic feature of the entognathous apterygotes, not found
in any of the other insects, is the presence of a pair of rodlike scleroti-
zations extending posteriorly from the hypopharynx in the inner walls
of the gnathal pouches between the bases of the maxillae and the
labium (fig. 21 D, mB), which diverge posteriorly and support the
maxillary cardines. As seen in specimens cleared and mounted whole,
these rods appear to be apodemes, and have been regarded as “ten-
torial arms,” but it has been shown by Folsom (1900) that, in their
origin in Collembola, they are linear sclerotizations of the ventral head
wall. In their position between the maxillae and the labium, or second
maxillae, the rods are quite comparable to the intermaxillary sternal
brachia of Crustacea, which in the isopods and amphipods similarly
support the first maxillae (see Snodgrass, in press). In the Protura the
anterior parts of the rods are united with each other forming a
Y-shaped structure. In the Diplura the two rods are connected inside
the head by an arched ligamentous bridge (fig. 21 D, Lg), on which
are attached muscles of the mandibles and maxillae; in Collembola
the bridge is elaborated into a highly developed platform for muscle
attachments, which is known as a “‘tentorium,” though it can have
no relation to the tentorium of Thysanura and Pterygota.

The mandibles of the entognathous hexapods are elongate, with
the gnathal lobes extended in line with the bases of the appendages,
sometimes drawn out into piercing stylets, but never flexible or inde-
pendently movable as in the myriapods. In Protura and Collembola
each mandible is connected posteriorly with the cranial margin, as in
Chilopoda, by a slender rod in the lateral wall of the gnathal pouch.
According to Hoffmann (1908) the mandibles of Collembola have
also an anterior articulation on the head. In Diplura there is neither
an anterior nor a posterior articulation. Regardless of the type of
articulation, however, the mandibles are protractile and retractile, and
in addition apparently have a strong rotary movement on their long
axes. The mandibular musculature is similar in the three entognathous
groups ; it becomes unusually complex in the Collembola, probably no
other arthropod having so many muscles attached on its jaw, but
the musculature is merely an elaboration of the usual anterior and

| ae

Ee eee a

--
\
(op)

---74f--

/ “S
3 ve lbrmel

Stom V (adrnd)

Fic. 21—Hexapoda—Entagnathous apterygotes: Diplura, Collembola,
and Protura.

A, Heterojapyx gallardi Tillyard, left mandible and muscles, dorsal. B,
Orchesella cincta L., left mandible and muscles, dorsal (from Folsom, 1899).
C, Tomocerus plumbeus L., right mandible and muscles, dorsal (adapted from
Hoffmann, 1908). D, Heterojapyx gallardi Tillyard, cross section of head.
E, Campodea sp., mandibles and adductor muscles. F, Tomocerus plumbeus L.,
right mandible, ventral (from Hoffmann, 1908). G, Entomobrya lanugenosa
(Nicolet), mandible (from von Stummer-Traunfels, 1891). H, Anurida
maritima Guér., left mandible, dorsal (from Folsom, 1900). I, Eosentomon
germanicum Prell, right mandible, ventral (from Prell, 1913). J, Acerentomon
doderoi Silvestri, left mandible and muscles, dorsal (from Berlese, 1910, part
ie fig. 121, pl. 12). P

3

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

posterior dorsal muscles and the ventral muscles of a more primitive
mandible.

Diplura (Dicellura).—The simple, elongate mandible in this en-
tognathous order (fig. 21 A) has a long mesal attachment on the
membranous inner wall of the gnathal pouch (D, Md), but it has no
articulation with the cranium either anteriorly or posteriorly. The
elongate, toothed gnathal lobe is blunt in Japygidae (A) ; in Cam-
podea (E) it is more acute and bears mesally a small setose appendage.
The proximal end of the mandible is extended into a tapering process
(A) that lies free in the containing gnathal pouch (gnP).

The mandibular musculature, as seen in Heterojapyx gallardi (fig.
21 A), consists of six dorsal muscles (Z-6) arising on the cranium,
and two ventral muscles (7, 8) attached on the ligamentous bridge
(Lg) that connects the two intermaxillary sternal brachia (D, Lg)
of the inner walls of the gnathal pouches. The cranial muscles at-
tached dorsally on the mandible (representing the A fibers of a more
generalized mandible) include two distal lateral rotators (A, 1, 2)
and two proximal retractors (3, 4) ; those attached ventrally (the P
fibers) are two large mesal rotators (5, 6) from the median ridge of
the cranium. The ventral muscles include a wide sheet of transverse
adductor fibers (7), and a smaller band of oblique fibers (8), which
is evidently a protractor. All the ventral fibers of the mandibles, as
above noted, arise from the ligamentous bridge between the inter-
maxillary sternal brachia (D, Lg). Superficially the fibers from the
opposite mandibles united on the bridge give the appearance of an
intermandibular muscle, as seen in Campodea (E), and were formerly
described as such by the writer (1928). The association of the
mandibular muscles with the sternal brachia (D, Lg), however, is
certainly secondary, since these rods support the first maxillae and
can have no relation to the mandibular segment. The mandibular
adductors of the Diplura represent the ventral muscles of the man-
dibles that are attached on an intergnathal ligament in more general-
ized arthropods.

Within the mandible of Heterojapyx there arises posteriorly a
slender muscle (fig. 21 A, 9) that anteriorly joins the tendon of a
long labral muscle (/brmcl) arising posteriorly on the cranium, and
inserted by its tendon on the base of the under wall of the labrum.
No such muscle has been observed in any other insect.

Collembola.—The collembolan mandibles in general resemble the
mandibles of Diplura, but in vegetable-feeding species the gnathal lobe
is more or less differentiated into an incisor process and a molar sur-
face (fig. 21 B, C, F, G); in Anurida, however, which is said by

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 65

Folsom (1900) to feed on the soft tissues of the mollusk Littorina,
the slender mandible (H) simply expands distally in a sharply toothed
lobe.

The mandibles of Collembola, being protractile and retractile within
the gnathal pouches, have no fixed articulations, but each mandible is
supported posteriorly on the end of a slender, rodlike thickening of
the lateral pouch wall (fig. 21 B, mdr), which is attached anteriorly
on the margin of the cranium (H). This suspensory structure of the
collembolan mandible, found also in Protura (I), exactly duplicates
the attachment mechanism of the jaw in the Chilopoda (fig. 18 A,
mdr) and Pauropoda. The mandibular rod is called the “stirrup” by
Folsom (1899) ; Denis (1928) refers to it as a “ligament.”’ Folsom
believed that in protraction the mandible left the socketlike end of
the “stirrup,” but Denis points out that the rod, or “ligament,” is
attached to the mandible and is itself a part of the wall of the gnathal
pouch, so that it simply bends forward or backward with the move-
ment of the jaw. According to Hoffmann (1908) the collembolan
mandible has an anterior process (fig. 21 F, c) that bears against an
articular surface of the head at the base of the “anterior tentorial
arm.” Evidently a disarticulation must take place here when the
mandible is protracted. Other writers have not recorded the existence
of an anterior articulation in Collembola.

Descriptions of the collembolan mandibles have been given by sev-
eral writers, including von Stummer-Traunfels (1891), Folsom
(1899, 1900), Hoffmann (1908), Denis (1928), Hansen (1930).
Folsom, Hoffmann, and Denis include also a full account of the
mandibular musculature. The number of muscles attached on each
mandible differs in different genera, but the fiber bundles all fall into
the usual three major groups attached on a generalized mandible,
namely, anterior dorsal muscles and posterior dorsal muscles arising
on the cranial wall, and ventral muscles, which in the Collembola
arise on the “tentorium.” Because of the horizontal position of the
collembolan jaws, the primitively anterior muscles become dorsal,
and the posterior ventral. By diversification of their points of origin
on the head, the dorsal muscles are differentiated functionally into
rotators and retractors of the jaw, and similarly the fibers of the
primary ventral adductor are differentiated into adductors and
protractors.

Folsom (1899) finds in Orchesella cincta 10 muscles attached on
each mandible (fig. 21 B), 7 arising on the cranial wall, and 3 on the
“tentorium.” Of the cranial muscles, four are attached dorsally on
the mandible and three ventrally. As named and numbered by Folsom,

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the dorsal muscles include a lateral rotator (1), an abductor (2), a
retractor and rotator (3), and a retractor (4) ; the ventral muscles
include two long rotators (7, 6) and another not shown in the figure
lying beneath 7. The “tentorial” fibers are differentiated into a broad
transverse adductor (9), and into two protractors (5, 6) extending
from the posterior end of the mandible to the anterior arm of the
“tentorium.”

It will be observed that the musculature of the mandible in Orche-
sella (fig. 21 B) is essentially identical with that of the dipluran
Heterojapyx (A). A much simpler mandibular musculature is found
by Denis (1928) in Anurida maritima, consisting of three rotators
arising on the head wall, adductors from the body of the “tentorium,”
and protractors arising at the base of the lingua. In Onychiurus
fimentarius Denis shows the musculature to be more complex than
that of Anurida, since it includes the same muscles and some others
in addition. In Tomocerus catalanus he finds an elaborate mandibular
musculature essentially the same as that described by Hoffmann for
Tomocerus plumbeus.

The mandibles of Tomocerus plumbeus have the greatest number
of muscles attached on them yet recorded for any collembolan. As
given by Hoffmann (1908) there are 17 distinct fiber bundles inserted
on each mandible of this species (fig. 21C). To eight of these
muscles (2-6, 9, 10) arising dorsally on the head wall, Hoffmann
ascribes a rotary function, while two others (rz, 72) with posterior
origins are retractors. Opposed to the last are two muscles (7, 8)
arising anteriorly on the “‘tentorial” arm, which are protractors. The
rest of the muscles, five in number (13-17), are adductors with their
origins on the body of the “tentorium” (14, being ventral, is not seen
in the figure). Clearly, the mandible of this collembolan is equipped
for hard work.

It should be noted here that the so-called “tentorium” of the Col-
lembola appears to be a structure of the nature of the intergnathal
ligament of the crustaceans and myriapods, supported on the inter-
maxillary sternal brachia, and clearly has no homology with the cuticu-
lar tentorium of Thysanura and Pterygota (see Snodgrass, in press).

Protura.—The mouth parts of Protura have been described by
Berlese (1910), Prell (1913), Hansen (1930), and Tuxen (1931).
The mandibles are slender and sharp-pointed (fig. 211), in some
forms drawn out into long, tapering stylets (J, Md). The base of
each mandible is attached mesally to the membranous inner wall of
the gnathal pouch and opens by an elongate foramen into the head
cavity (1); the narrowed posterior end is shown by Prell to be con-

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 67

nected with the cranium by a slendor rod (I, mdr). Tuxen interpreted
the mandibular rod as a “pleural apodeme” of the head, but there
can be little doubt that it is a mere thickening of the wall of the
gnathal pouch, as is the similar rod in Collembola and Chilopoda.
The mandibular musculature in the Protura is simplified because
its chief function is that of protraction and retraction. The muscula-
ture of the mandible of Acerentomon doderoi (fig. 21 J), as described
and figured by Berlese (1910), consists of anterior and posterior
fiber bundles disposed in the form of an X, so that the two opposing
sets appear to be protractors and retractors. The lateral anterior mus-
cles (z) and the mesal posterior muscles (2), however, both arising
on the cranium, are diagonal with respect to the axis of the man-
dible and are attached on its upper margin. These muscles, there-
fore, must have a rotary action on the mandible accompanying pro-
traction and retraction. The other muscles (3, 4) are approximately
parallel with the mandibular axis, and undoubtedly are protractors
and a retractor. The single retractor (3) arises posteriorly on the
cranium ; the three protractors (4) take their origins anteriorly on the
anterior arm of the Y-shaped sternal sclerite of the head (the so-
called “tentorium” of Berlese and Prell). These ventral protractors
of the proturan mandible, as noted by Tuxen, are derivatives of the
primitive ventral adductors of the appendage; as are the corresponding
protractors of the mandible in Collembola, Diplura, and Chilopoda ;
functionally adductor fibers appear to be absent in Protura.

VI. THYSANURA

There can be little question that the ectognathous thysanuran and
pterygote insects are more closely related to each other than either
group is to the entognathous apterygotes, and yet their likenesses and
differences are so inconsistently distributed that it is difficult to dis-
cover the nature of their interrelationships. The mandibles of the
Thysanura differ conspicuously between the two principal families of
the order, the Machilidae and the Lepismatidae, but not in a way
that would preclude the evolution of the lepismatid type of jaw from
that of the machilid. The machilid mandible is a pendent jaw with
a single dorsal articulation, and is quite comparable in its form,
musculature, and mechanism to the mandibles of lower Crustacea.
The lepismatid mandible has a horizontally elongate base, is doubly
articulated on the head, and the gnathal lobe projects downward from
the axis of rotation, in which features it closely resembles, on the
one hand, the mandibles of the isopods and amphipods among the

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Crustacea, and, on the other, the biting and chewing type of jaw of
the Pterygota, except larval Ephemeroptera. The ephemeropterid
mandible more resembles the mandible of Machilis than that of the
other pterygotes. The machilid jaw is simply a mandible of general-
ized type from which the doubly articulated jaw has been evolved
independently in the Crustacea, Thysanura, and Pterygota.

The mandibles of Machilidae (fig. 22 A, B) are elongate and hang
downward from the cranial margins behind the antennae on single
points of articulation (a). The long oval base of each jaw is attached
to the membranous lateral wall of the head below the cranial support,
so that the jaw swings freely in a transverse plane; the free gnathal
lobe is divided into an elongate, apical incisor process, and a thick
molar process directed mesally. The musculature of the machilid
mandible is typical of a jaw of its kind. Each mandible has an anterior
and a posterior dorsal muscle (A, A, P), apparently rotary in func-
tion, and a large mass of ventral adductor fibers. The adductor fibers,
however, are differentiated into a distal adductor (1) and a proxi-
mal adductor (2V). The fibers of the first spread into the cavity of
the mandible (A, B, 1V) and converge to a median ligament (Lg)
by which they are united with the corresponding fibers of the opposite
mandible. The ligament passes through the base of the hypopharynx
behind the roots of the anterior tentorial arms (A, AT). The other
ventral fibers form a wide, flat muscle (A, 2V) for each mandible
arising on the tentorial arm and inserted on the posterior edge of the
mandibular base. This muscle would appear to have a rotary as well
as an adductor action on the jaw. The ventral musculature of the
machilid mandible is thus seen to be the same as that of Lithobius
(fig. 18 A) or a diplopod (fig. 20 D) in that the adductor fibers of
each jaw are separated into a distal group united with the correspond-
ing fibers from the opposite jaw, and into a posterior group attached
individually on a head apodeme. The entire musculature of the machi-
lid mandible is carried over into the Lepismatidae and the Pterygota,
but in these groups the anterior fibers of the ventral adductors from
each jaw become separately attached on the base of the hypopharynx.

The mandibles of Lepismatidae (fig. 22 D, Md) are attached to the
lower margins of the head by their elongate bases, with anterior and
posterior articulations, so that the gnathal lobes swing transversely
on horizontal axes of rotation. The posterior articulation (a) is in a
notch of the cranial margin. The anterior articulation (c) is not
with the clypeus, as it is in the pterygote insects, but with a small
condyle (F, c) on the ventrally inflected anterior angle of the gena
(Ge) behind the clypeus, just outside the invagination (at) of the

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 69

My

TAN
fl il j

|

n

Vy

/

Pe
Pr
YI

Fic. 22.—Hexapoda—Thysanura.

A, Nesomachilis maoricus Tillyard, vertical cross section through back of
head, with suspended mandibles and their muscles, posterior. B, Machilis sp.,
mandibles, with intergnathal adductor and posterior cranial muscles, posterior.
C, Ctenolepisma urbana Slabaugh, left mandible and muscles, with part of
tentorium, dorsal. D, same, head, lateral. E, same, left mandible and muscles,
lateral. F, same, horizontal section of head below tentorium, ventral. G, same,
“oa of head and mandibles, showing distribution of mandibular muscles,
ventral.

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

anterior arm of the tentorium (AT). It is evident that this anterior
articulation of the mandible has been independently acquired in the
Lepismatidae, since it is not the same as that in the Pterygota. The
musculature of the lepismatid mandible corresponds with that of
the machilid mandible in that it consists of anterior and posterior
dorsal muscles and two ventral muscles (fig. 22 C, E) ; but, because
of the horizontal axis of rotation of the lepismatid jaw (E, a-c), the
anterior muscles, of which there are two in Ctenolepisma (1A, 2A),
become lateral abductors, and the single, greatly enlarged posterior
muscle (P) a mesal cranial adductor. Of the two ventral muscles
(C, E, 1V, 2V), both of which evidently are adductors, the second
(2l”) is much the larger and is attached on the anterior arm of the
tentorium (C, F, G); the first (zV) is a relatively small muscle
clearly corresponding with the intermandibular muscle of Machilis
(A, B, 1V), but the fibers from each mandible are attached separately
on the suspensory arms of the hypopharynx, just as they are in the
lower pterygote insects (fig. 24 B, C, 1V). It is of interest to note
again here that the lepismatid hypopharynx, as the writer has else-
where shown (in press), has the structure typical of the orthopteroid
hypopharynx, and in no way resembles the primitive 3-lobed hypo-
pharynx of Machilidae and larval Ephemeroptera.

Vil. PrERYGOTA

The winged insects in their fundamental adult structure are well
standardized, probably because the function of flight does not permit
any great degree of diversification in body form. The flightless young,
however, being free from the restrictions placed on the adults, have
had liberty to adapt themselves to various special environments, and
many of them have taken advantage of their freedom to the extent of
assuming bodily forms that in most cases have no relation to the
ancestral adult form, though, for the most part, the larval feeding
organs have retained a fairly generalized structure. On the other
hand, since the manner of feeding has little effect on the function of
flight, adult insects have been free to develop highly specialized kinds
of mouth parts. Hence, in a study of the mandibles, the jaws of the
larva are likely to be more representative of the primitive mandibles
of the species than are those of the adult.

Among the winged insects the Ephemeroptera stand apart from
the other orders by various features that give them a relatively primi-
tive status, as the wing venation and the paired genital openings of
the adult, but other primitive characters, less commonly considered,

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 71

are found in the head and the mouth parts of the larva. The larval
tentorium, for example, has its anterior roots in the ventrally inflected
lower margins of the genae mesad of the mandibles (fig. 23 C, at)
between the base of the clypeus and the posterior articulations of the
mandibles, while in all other pterygote insects the anterior tentorial
invaginations are Jaterad of the mandibles on the sides or the facial
aspect of the head. The mayfly larva has a well-developed 3-lobed

Fic. 23—Hexapoda—Pterygota: Ephemeroptera.

A, Hexagenia sp., adult, ventral view of head and mouth parts. B, same, left
mandible, ventral. C, Leptophlebia sp., larva, head, lateral. D, same, right
mandible and muscles, mesal. FE, same, left mandible, dorsolateral.

hypopharynx, such as occurs in Machilidae, the entognathous aptery-
gote hexapods, the symphylids, and various crustaceans, but is found
in no other pterygote insects, nor even in the Lepismatidae among
the apterygotes. The large jaws of the ephemeropterid larva (fig.
23 C, Md) resemble those of other pterygote insects in general ap-
pearance and in their position on the head, but most surprising it is
to find that they have no anterior articulations and no connection what-
ever with the clypeus. Each mandible has a posterior articulation, and
is membranously attached to the subgenal margin of the cranium by
only its posterior part, leaving a long, concave space on the ventrally

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

inflected area of the gena between it and the clypeus, in which the
anterior part of the jaw rests and turns. Though the ephemerid larval
jaw resembles in form and position the mandible of an orthopteroid
insect (fig. 24 A), its actual structure is that of the singly articulated,
pendent mandible of the Machilidae. Its musculature (fig. 23 D) is
that typical of Lepismatidae and the lower Pterygota, there being a
huge dorsal adductor inserted on a broad tendon (Pt), a slender dorsal
abductor attached on a smaller tendon (At), and two ventral muscles,
one of the latter (rV) being attached on the base of the hypopharynx,
the other (2/) on the tentorium. Murphy (1922) erroneously
ascribes all the muscle fibers inserted within the mandible, termed
“flexors,” to the tentorium.

In the adult mayfly (fig. 23 A) the mouth parts are greatly reduced.
Sternfeld (1907) says the mandibles have vanished entirely, but in
the species of Hexagenia here figured mandibles (A, Md) are dis-
tinctly present, though each mandible (B) is merely a small, soft,
somewhat quadrate, flattened lobe with a long vermiform flagellum
arising from its anterior mesal angle. Mandibles of similar form are
shown by Murphy (1922) in the subimago of Hexagemia recurvata.
Sternfeld notes that the reduction of the mayfly mouth parts begins
in the nymphal stage, but that the alimentary canal, on the contrary,
is not reduced at any stage. In the imago, Sternfeld says the ali-
mentary canal is filled with air and serves for facilitating flight and
particularly the act of hovering; the digestive tract thus gives up its
original function, but it takes on a new one. From Sternfeld’s sec-
tional view of the anterior part of an adult mayfly it would appear that
the pharyngeal muscles and the sucking apparatus are well preserved.

The typical pterygote mandible, as seen in the cockroach (fig. 24 A),
is a doubly articulated jaw with an approximately horizontal axis of
movement (a-c) on the lower, lateral margin of the cranium, so that
it swings transversely toward its opponent. The mandibles close
between the clypeus and labrum in front and the hypopharynx behind
(D); in the cockroach the left mandible overlaps the right. The
mandible is membranously attached to the head and the articular
surfaces are external points of contact with the head wall (A). The
primary articulation (a) is posterior on the subgenal margin of the
cranium, the secondary articulation (c) is anterior on the base of the
clypeus (Clp), which latter in the cockroach is not separated from
the frons (Fr). If the insect is prognathous, the axis of the jaw
becomes vertical instead of horizontal, but the relation of the mandible
to the head is not changed. In biting-and-chewing insects a molar
surface of the mandible is differentiated from a toothed incisor

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 73

if

Fic. 24.—Hexapoda—Miscellaneous Pterygota.

A, Blatta orientalis L. (Blattidae), left mandible and adjoining parts of head,
lateral. B, same, right mandible and muscles, anterior. C, same, section of
head showing mandibles with muscles, hypopharynx, and tentorium, anterior.
D, Periplaneta americana (L.) (Blattidae), section of mouth parts showing
relative positions. E, Popillia japonica Newm. (Scarabaeidae), left maxilla,
dorsal. F, same, mandibles, ventral. G, Cotinis nitida (L.) (Scarabaeidae),
mandibles, ventral. H, Berosus pugnax LeConte (Hydrophilidae), left mandible,
ventral. I, Mycetobia sp. (Mycetophilidae), left mandible of larva, lateral.
J, same, incisor lobe of mandible, mesal. K, Popillia japonica Newm. (Scara-
baeidae), left and right mandibles turned dorsally, showing masticatory surfaces
on mesal sides.

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

process. The ordinary mandible of the winged insects is practically
identical in its structure and mechanism with the mandibles of the
isopods and amphipods among the Crustacea, but since the mandibles
of the apterygote insects and the ephemeropterid larva are not of this
type, it is clear that the doubly articulated mandible having the gnathal
lobe perpendicular to the axis of rotation has been independently
developed in both the crustaceans and the insects.

The mandibular musculature of the cockroach and of most other
orthopteroid insects except Acrididae is the same as that of the lepis-
matid mandibles (fig. 22 E) in that each jaw has four distinct muscles
(fig. 24 B), though the corresponding muscles differ in relative size
in the two groups. The principal motors of the pterygote mandible,
however, are the dorsal muscles (C), including a slender lateral abduc-
tor (A) and a huge mesal adductor (P). The ventral muscles of the
cockroach mandible (B, C, V) are inserted within the cavity of the
jaw and consist of two distinct fiber groups, one being a fan-shaped
muscle (zV) inserted laterally in the mandible, with its convergent
fibers attached on a small arm (1) of the hypopharyngeal suspen-
sorium (HS), the other (2V) a short, thick muscle inserted poste-
riorly in the mandible and attached on the anterior arm of the ten-
torium (C, Tut). The musculature of the pterygote mandible is thus
seen to be fundamentally the same as in the Thysanura and other
arthropods. In most of the holometabolous insects, however, the
ventral muscles have been lost, and the dorsal muscles become the only
muscles of the jaw (F), though in some of the mandibulate Diptera,
as in Tabanidae, a small tentorial muscle of the mandible may be
retained.

In no other group of arthropods do the mandibles undergo such
diverse and numerous modifications in adaptation to different ways of
feeding as they do in the insects. Though the typical biting and chew-
ing mandible has a well-differentiated toothed incisor process and a
grinding molar surface, either part may be reduced or eliminated.
Among the beetles the mandibles of species that practice extra-oral
digestion of the food have no molar surfaces; in the leaf-feeding
Scarabaeidae on the other hand, in which strongly toothed lobes of
the maxillae serve as jaws for biting off the leaf surface (fig. 24 E),
the mandibles have become specialized as masticatory organs by a
great development of the molar surfaces and a reduction of the incisor
processes (F, G, K). In various orders the mandibles are drawn
out into piercing stylets, in the worker bees they are modified for pur-
poses other than that of feeding, and finally the mandibles may be
greatly reduced in size or entirely suppressed. Weber (1939) has

NO. I JAWS OF MANDIBULATE ARTHROPODS—-SNODGRASS 75

given a classified account of various atypical kinds of insect mandibles,
among which may be mentioned the mandibles of certain leaf-mining
lepidopterous larvae in which the molar lobes have been converted
into saws, and others in which the usual action of the jaws and the
1elative size of the muscles have been reversed, so that the jaws work
outward instead of inward, as in the elephant louse, Haematomyzus,
and in the larvae of the tenthredinid Phyllotoma aceris.

The mandibles of some of the Hydrophilidae, as in Berosus (fig.
24H), are armed each with a pair of large dentate processes (/m),
flexibly attached on the inner margin of the jaw, which have been
cited by Hansen (1930) as examples of the occurrence of a “lacinia
mobilis” among the insects; similar processes he says are present in
Hydrobius, but not in Hydrophilus. A small, flexible appendage bear-
ing a brush of hairs is seen also on the larval mandibles of Ephemerop-
tera (fig. 23 D, E), and is termed a “lacinia mobilis” by Murphy
(1922). Though such structures are widely distributed among the
arthropods, as already pointed out, there is no reasonable basis for
regarding them as homologues of the lacinia of an insect maxilla.

In the larvae of lower nematocerous Diptera the entire incisor
process of the mandible may be separated by membrane from the
sclerotized base of the jaw, and thus becomes freely flexible on the
latter. An example is here given in a species of Mycetobia (fig.
241, J). The mandibles of such species are so articulated on the head
that they work in longitudinal planes parallel with each other and
not in opposition. In a comparative study of families having this type
of larval jaw Anthon (1943a, 1943b) finds that the distal lobe, or
“segment,” of the mandible is most freely movable in the Sylvicolidae
(Rhyphidae), in which it is doubly articulated on the base. In the
Melusinidae (Trichoceridae) the lobe is united with the base by a
sclerotic bridge on the dorsal side. Among the Psychodidae the lobe
in some species is freely movable, while in others it has an extensive
union with the base. In the Liriopeidae (Ptychopteridae) the incisor
lobe is apparently immovable, since it is set off from the base only by
a groove. The apical lobes of the jaws in Sylvicolidae (Rhyphidae)
bear large brushes of long hairs, which Anthon says serve as brooms
for sweeping into the mouth particles of detritus, algae, and diatoms
on which the larva feeds. Other forms, in which the lobes are armed
with strong teeth and are not so freely movable, feed on vegetable
fragments less easily removed from their source. The purely aquatic
rhyphid larvae, Anthon observes, use the mandibles also for locomo-
tion, creeping by means of the jaws on the bottom, while the rear end
of the body with the spiracles is held at the surface of the water.

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Insect mandibles of the type just described have a certain re-
semblance to the diplopod jaw, but the flexible incisor lobe of the
dipterous larval mandible is entirely devoid of muscles, and is mov-
able merely by a desclerotization of the mandibular wall around its
base. Considering the evidence that the entire arthropod jaw repre-
sents only the coxa of the mandibular appendage, and that the true
distal segments, when present, form a palpus, it is impossible to
accept Anthon’s conclusion that the divided mandible of the sylvicolid
(rhyphid) larval type is a primitive, segmented mandible, from which
the solid type of jaw has been derived by union of the two “segments.”
The dipterous larva certainly does not represent a primitive arthropod.

The most specialized mandibular mechanism among the insects is
that of the muscularly protractile and retractile styletlike or bristlelike
mandibles of Hemiptera. In the Homoptera the enlarged base of
each mandibular stylet (fig. 25 E), where it is attached to the wall
of the containing pouch, is produced into a slender retractor arm (ra)
and a broad protractor arm (pa). The retractor muscles (F, A, P)
are attached on the retractor arm and in the notch between the bases
of the two arms; these muscles evidently represent the dorsal muscles
of a generalized mandible. The protractor muscles (1V), inserted
on the upper end of the protractor arm, arise on the lower end of a
plate of the head wall (A, Lor), inserted between the clypeus and the
maxillary lobe, termed the lorum by taxonomists. The identity of
the protractor muscles of the mandibles, therefore, depends on the
homology of the loral plates.

By some students of Homoptera the loral plates of the head have
been thought to be detached parts of the clypeus, while others have
regarded them as extensions of the genae. The writer (1938) sug-
gested that the lora belong to the hypopharynx, since their lower ends
are confluent with the hypopharyngeal floor of the sucking pump
(fig. 25 C). The independence of the loral plates from the clypeus,
and their direct continuity with the sides of the hypopharynx in the
cicada has been fully demonstrated by Butt (1943, fig. 2), and ina
sectional view of the head of Typhlocyba ulmi, Willis (1949, fig. 4)
shows clearly that the inflected lateral edges of the clypeus are con-
tinued into the epipharyngeal roof of the sucking pump.

A horizontal section of the head of a cicada through the lower part
of the postclypeus (fig. 25 B) shows that the contiguous edges of the
postclypeus (Pclp) and the lora (Lor) are inflected individually
clear through the head. The clypeal lamellae become the epipharyngeal
wall (Ephy) of the sucking pump (CbP) with the plunger on which
the dilator muscles (dicb) are attached ; the loral lamellae (n) become

,

No. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 7/

Fic. 25.—Hexapoda—Pterygota: Homoptera.

A, Cephisus siccifolia Walker, head, anterior. B, Magicicada septemdecim
(L.), horizontal section of head through lower part of postclypeus. C, same,
anterior surfaces of loral plates connected with hypopharyngeal trough (Sit)
of sucking pump. D, Tibicen sp., posterior view of lower part of head after
removal of labium, lower ends of maxillary lobes and stylets, showing posterior
median lobe of hypopharynx (p) membranously connected (o) with loral plates
(Lor). E, Magicicada septemdecim (L.), basal part of right mandible, posterior.
F, same, basal part of left mandible with muscles, anterior; anterior branch of
protractor muscle (z/) attached below r on C, posterior branch at s.

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the hypopharyngeal floor of the pump with the deep, median, trough-
like sitophore (Sit). Likewise, the posterior edges of the loral plates
are inflected, but become thin membranes (0) continuous medially
with the ventral wall of the posterior lobe (p) of the hypopharynx.
The hypopharynx and the lora, therefore, constitute an anatomically
integral structure; the loral plates are merely the exposed outer sur-
faces of lateral expansions of the hypopharynx (Hphy). The same
relations can be demonstrated by dissection. On lifting or removing
the anteclypeus (A, Aclp), it is seen that the lower ends of the loral
plates (C, Lor) converge behind the anteclypeus and that their broad,
sclerotized anterior lamellae are united with the sitophore trough (Sit)
of the hypopharynx. Working from the back of the head (D), by
removal of the labium and cutting off the lower parts of the maxil-
lary lobes (ML), the median posterior lobe of the hypopharynx (/)
is exposed, and is seen to be continuous with the membranous areas
(0) on each side inflected from the posterior margins of the loral
plates (Lor). .

From these facts it is clear that the loral plates of Homoptera are
literally parts of a complex hypopharyngeal structure. It then fol-
lows that the protractor muscles of the mandibles arising on the lora
can be no other than the hypopharyngeal muscles of the orthopteroid
mandibles (fig. 24 B, 1V), the altered relative position of which re-
sults from the retraction of the mandibles and the lateral expansion
of the loral wings of the hypopharynx. In some Fulgoridae the
upper ends of the loral plates appear to be united with the clypeus.
In the Heteroptera (Hemiptera) it is shown by Butt (1943) that the
sclerites corresponding with the lora of Homoptera are concealed
within the head.

In the cicada, either Tibicen or Magicicada, there are two large
branches of the protractor muscle inserted on the upper end of the
protractor arm of each mandible (fig. 25 F, 1V), one lying anterior
to the other. The anterior branch arises ventrally on the inner sur-
face of the lower part of the inflected dorsal lamella of the lorum
behind the point r on C of the figure; the posterior branch arises
on the same loral lamella at the base of the sitophore (s). The pro-
tractor arm of the mandible is not an apodeme; it is a sclerotic exten-
sion of the mandibular base in the membranous anterior wall of the
gnathal pouch. The muscle-bearing haemocoelic surface, therefore,
corresponds with the inner lateral wall of the orthopteroid mandible
on which the hypopharyngeal muscle of the jaw is inserted. The
homopterous head, the mandibles and their muscles can thus be inter-

No. I

JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 79

preted in terms of orthopteroid structure; they show merely the ex-
treme degree to which modifications of a basic anatomical complex
may be carried in the evolution of a new mechanism.

EXPLANATION OF LETTERING ON THE FIGURES

ABBREVIATIONS

A, anterior dorsal muscle of a primi-
tive mandible (14, 2A,
3A, subdivisions of same).

ablvr, abductor lever.

Aclp, anteclypeus.

admd, adductor muscle of mandible.

rtAnt, first antenna (antennule), or its
head foramen.

2Ant, second antenna, or its head fora-
men.

Ap, apodeme.

at, anterior tentorial invagination.

At, tendon of muscle A.

AT, anterior tentorial arm.

brC, branchial chamber.
bspd, subdivision of basipodite.
Bspd, basipodite.

CbP, cibarial sucking pump.

Chpd, cheliped, or its foramen.

Cp, carapace.

CT, central plate of tentorium (corpo-
tentorium).

Cvux, neck, cervix.

Cx, Cxpd, coxa, or coxopodite.

Dbl, doublure (inner wall) of carapace
fold.

dpm, dorsal promotor muscle of coxa.

drm, dorsal remotor muscle of coxa.

DT, dorsal tentorial arm.

Endst, endosternum.

Ephy, epipharyngeal surface.
Epst, epistome.

es, epistomal sulcus.

Expd, exopodite.

Fit, fultura (premandibular sternal
sclerite supporting hypo-
pharynx).

Fr, frons.

Ga, galea.

Ge, gena.

Gn, jaw (gnathal endite).
Gnch, gnathochilarium.

gnL, gnathal lobe of mandible.
gnP, gnathal pouch.

H, head.

hAp, head apodeme, “hypopharyngeal
apophysis” of myriapods.

Hphy, hypopharynx.

HS, hypopharyngeal suspensorium.

I, intramandibular muscle of gnathal
lobe.

IT, III, IV, etc., somatic segments (J,
segments of second anten-
nae).

imB, intermaxillary sternal brachium.

inc, incisor process of mandible.

inv, site of an endoskeletal invagina-
tion.

Lb, labium.

Le, lacinia.

Lg, intergnathal ligament.
Im, “lacinia mobilis.”

Lm, labrum.

Lor, lorum.

lvr, leverlike sclerite.

mel, striated muscle tissue.

Md, mandible.

mdB, base of mandible (corpus man-
dibulae).

mdF, mandibular foramen.

mdmcls, mandibular muscles.

mdr, articular rod of mandible.

mol, molar process or surface of man-
dible.

mp, muscle plate.

Mst, metastome.

80

Mth, mouth.

1Mx, first maxilla (maxillula), or its
foramen.

2M, second maxilla, or its foramen.

mb, remnant of maxillary bridge.

mxB, maxillary bridge (pleuron of
maxillary segment).
2MxF, foramen of second maxilla.

MxL, maxillary lobe.

Mexpd, maxilliped (1Maxpd, 2M xpd,
3M xpd, first, second, and
third maxillipeds, or their
foramina).

npr, nephropore.
Oe, oesophagus.

P, posterior dorsal muscle of a primi-
tive mandible.

pa, protractor arm of mandible.

Pclp, postclypeus.

Pgn, paragnath.

Pl, pleuron.

Plp, palpus (telopodite or a part of it).

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116

pmB, postmaxillary sternal brachium.
Prtc, protocephalon.
Pt, tendon of muscle P.

ra, retractor arm of mandible.

S, sternum.

Sit, sitophore (troughlike floor of ciba-
rial pump).

sl, suspensory ligament.

smcl, suspensory muscle.

Sp, spiracle.

Stom, stomodaeum.

Sty, mandibular stylet.

T, tergum.
Tipd, telopodite.
Tnt, tentorium.

V, ventral muscle, or muscles (1V, 2V,
3V), of mandible.

vpm, ventral promotor muscle of coxa.

vrm, ventral remotor muscle of coxa.

VS, sternum of first maxilliped seg-
ment.

ALPHABETICAL LETTERING

a, primary dorsal articulation of coxa or mandible.
a-c, axis of rotation of doubly articulated mandible.

b, primary ventral articulation of coxa.

c, secondary ventral or anterior articulation of mandible.
d, process of mandible supported on metastomal prop (t).

e, small sclerite between d and t.
f, epipharyngeal arm of fultura.

g, angle of mandibular articulation on fultura.

h, epistomal hinge of mandible.
i, j, cranial muscles of head apodeme.

k, lateral, postantennal expansion of epistome.

I, stomodaeal fibers of intergnathal ligament.

m, marginal sclerite of cranium in Lithobius.

n, sclerotic anterior wall of loral lobe of hypopharynx.

o, membranous posterior wall of loral lobe of hypopharynx.

p, median ventral lobe of hypopharynx.
q, apical lobe of hypopharynx.

r
Ss
t,
u
v

, origin on lorum of anterior protractor of mandible.
, origin on lorum of posterior protractor of mandible.
arm of metastomal plate supporting mandible.
, branch of intergnathal ligament to gnathal lobe of mandible.
, w, sclerites of proboscis wall in Dolops.

iN

NO. I JAWS OF MANDIBULATE ARTHROPODS—SNODGRASS 81

a, loral arm of hypopharyngeal suspensorium.
y, oral arm of hypopharyngeal suspensorium.
g, cut edge of carapace.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 2

Charles D. and Mary Waux Walcott
Research Fund

REMAINS OF LAND MAMMALS FROM
fia MIOCENE OF THE CHESAPEAKE
BAY REGION

By
C. LEWIS GAZIN
U. S. National Museum
AND
Rh. LEE COLLINS

University of Tennessee

OWE INCRE”
G cool OA ones Sa
BAe SCN

(PusicaTion 4019)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 12, 1950

The Lord Baltimore Press

BALTIMORE, MD., U. & 4.

Charles DB. and Mary Waux Walcott Research Fund

REMAINS OF LAND MAMMALS FROM THE
MIOCENE OF THE CHESAPEAKE
BAY REGION

By C. LEWIS GAZIN
U. S. National Museum
AND
R LEE. COELINS

University of Tennessee

The occasional discovery of the remains of various land mammals
in the Miocene marine deposits, principally along the cliffs of the
Chesapeake Bay in Maryland, has, in the course of a number of
years, resulted in a rather significant representation of these forms.
Heretofore no systematic attempt has been made to bring together
the various previous reports of land mammals from these beds and to
place on record the undescribed remains known to exist. Much of
the material extant is preserved in the U. S. National Museum. It
is comprised of an accumulation of gifts from private collectors,
Johns Hopkins University, U. S. Geological Survey, and U. S. Na-
tional Museum personnel and, in particular, represents the results
of searches by the junior author. The drawings depicting the speci-
mens were prepared by William D. Crockett, of the U. S. National
Museum.

The materials described in this paper are for the most part from
the Calvert formation, associated with a wealth of invertebrate fossils,
and with unquestionably the most abundant representation known,
both in numbers and diversity, of Miocene marine mammals. In some
instances the fossils have weathered out and were found loose on the
strand at the foot of the cliffs, and in cases where only Calvert is
reported exposed in the immediate section their derivation from this
formation is logically inferred. However, where Choptank formation,
also Miocene in age, is exposed in the upper portion of the cliff
section some doubt attaches to the origin, unless the specimen was
uncovered in place.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 2

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

PREVIOUS REPORTS

The occurrence of remains of Mammalia in the Chesapeake Mio-
cene is the basis of some of the very early reports on vertebrate fossils
in this country. The cast of a mastodon tooth that Leidy* included
in his description of Mastodon obscurus was considered to have been
made from the tooth mentioned by Harlan * in 1842 as coming from
the Miocene of Maryland, and by Lyell in the Proceedings of the
Geological Society of London for 1843 to 1845 (p. 38). Further
discussion is found in Warren’s * work on “Mastodon giganteus.”

Manatee and porpoise were also described from the Maryland
Miocene at about this time,* with subsequent papers by Leidy and
others on occasional finds. However, the first extensive reports on
the marine mammals of these beds were Cope’s studies of 1867-1869.
Included in Cope’s 1867 * paper was the description of Cynorca pro-
terva, the second type of land mammal to be described from these
‘beds, first thought to be a squalodont but later recognized as a peccary.

In the general review of the fauna from the Maryland Miocene in
the Maryland Geological Survey report of 1904, E. C. Case, in his
treatment of the mammals, overlooked the records of land types.

GEOLOGIC RELATIONS

The Miocene series in Maryland, known as the Chesapeake group,
includes three formations, the Calvert, Choptank, and St. Marys, in
ascending order. These consist essentially of sands, clays, marls, and
diatomaceous sediments, for the most part highly fossiliferous. The
beds are exposed in southern and eastern Maryland, to the southeast
of Baltimore and Washington, where they unconformably overlie the
greensand marls of the Eocene. For the most part they dip gently
to the southeast, preserving the higher formations of the group in
the more southerly portions of the State. The Chesapeake group is,
in turn, unconformably overlain by Pleistocene and possibly Pliocene
detritus over much of its extent, exposure of the Miocene resulting

1 Leidy, Joseph, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 7, pp. 244,
306, pl. 27, fig. 13, 1869.

2 Harlan, Richard, Amer. Journ. Sci., vol. 43, p. 143, 1842.

8’ Warren, John C., The Mastodon giganteus of North America, pp. 78-85,
175-180, 1852.

# Harlan, Richard, Medical and physical researches, p. 385, 1835; Bull. Proc.
Nat. Inst. No. 2, p. 195, 1842. Also, Dekay, J. E., Zoology of New York, pt. 1,
pp. 123, 136, 1842.

5 Cope E. D., Proc. Acad. Nat. Sci. Philadelphia, 1867, pp. 151-152.

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 3

largely from dissection of the older surfaces and terraces. Its de-
velopment is most impressive in the cliffs along the Chesapeake Bay.

The Calvert formation, with the superadjacent portions of the
Choptank, is the principal mammalian fossil-bearing formation of
the group. It is exposed from near Washington to below the mouth
of the Wicomico River along the Potomac, associated with Eocene
northward and with the Choptank southeastward. It extends south
into Virginia, and as a belt northeastward across eastern Maryland
and Delaware, and probably has for its equivalent the Kirkwood in
New Jersey.

AGE AND FAUNAL RELATIONSHIPS

The age of the Chesapeake series has been regarded by Dall® as
most nearly equivalent to the Helvetian of the European sequence,
which in turn is currently regarded as about middle Miocene. In the
committee report’ on the “Nomenclature and Correlation of the
North American Continental Tertiary,” the Calvert is regarded as
more nearly equivalent to the Tortonian of Europe, a stage higher
than Helvetian, and, hence, approximately upper middle Miocene
or upper Hemingfordian in the North American continental sequence.
It is compared to the “lower Sheep Creek” of the Nebraska Miocene,
“or, possibly, a slightly earlier age.”

From a critical examination of the fauna, the composition of which
was not fully known to the committee, we are inclined to regard the
stage of development as slightly later. This is based to a large
extent on the development reached in the Equidae and to some
extent on certain of the other forms. The horizon represented is
probably a little later than the lower Sheep Creek, or than the Florida
Hawthorn (as determined by the vertebrate localities in the northern
part of the State), although the extent to which the fauna is ad-
vanced over the faunas of those horizons is not great. The most
nearly comparable fauna is to be seen in the Merychippus zone of
the Temblor formation in California. The stage represented, if not
the latest Hemingfordian, is early Barstovian, earlier than the typical
Barstow fauna, and might be regarded as early Sarmatian.

The land mammals thus far recognized from the Calvert formation
are as follows:

Carnivora:
Tomarctus marylandica Berry

6 Dall, W. H., The Maryland Geological Survey, Miocene, p. cxliii, 1904.
7 Wood, H. E., 2d, et al. Bull. Geol. Soc. Amer., vol. 52, pl. 1 and p. 16, 1941.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Proboscidea :
Gomphotherium calvertensts, n. sp.
Perissodactyla :
Tapiravus cf. validus (Marsh)
Aphelops?, sp.
Archaeohippus, sp.
Merychippus, sp.
Artiodactyla :
Cf. Cynorca proterva (Cope)
Hesperhys (Desmathyus) ?,8 sp.

In a consideration of the age assignment, the Carnivora and
Artiodactyla listed above are not significant, although Hesperhys
would suggest a relatively earlier age than some of the other forms.
Its listing, however, cannot be regarded as in any way indicative,
inasmuch as Hesperhys has not been identified with any degree of
assurance. The presence of “Desmathyus” in the base of the Kirk-
wood is open to various interpretations, but for whatever it is worth
- attention should be called to the fact that Dall ® regarded the southern
New Jersey Miocene as containing reworked fossils from older
sediments.

Gomphotherium is significant in either of two ways. If the presence
of mastodon is to be regarded as placing an early limit on the
possible age assignment, then a lower Barstovian age would likely be
indicated. On the other hand, there is always the possibility that the
Calvert represents the first appearance of mastodons in North Amer-
ica, a conclusion not out of keeping with its geographic remoteness
from the western occurrences. Early appearances of mastodons in
other regions of North America include their presence in the Pawnee
Creek, Deep River, Mascall, Skull Spring, Sucker Creek, and Temblor
faunas. So far no mastodon has been recorded from the lower Sheep
Creek beds of Nebraska, from the Hawthorn of Florida, or from
the Phillips Ranch locality in California.

Among the Perissodactyla, the tapir gives little evidence regarding
the age assignment which may be made, other than to indicate a
relationship with the Shiloh marl of the Kirkwood formation in
New Jersey (from which Marsh described the species, Tapiravus
validus), inasmuch as tapir remains are exceedingly scarce in this
portion of the geologic column. Also, the rhinoceros present is of
no great help because the remains are too fragmentary for certain
generic reference, and Aphelops has been recorded from Hawthorn
to Pliocene times.

8 So far recorded only in zones 17 and 19, which are allocated to the Choptank.
® Dall, W. H., ibid., p. cxliv, 1904.

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 5

The horses, however, although only the lower teeth have been found,
give rather good indication of the portion of the Miocene represented.
The Merychippus teeth appear advanced over related types from the
Hawthorn or from the Sheep Creek beds, but, as noted under the
systematic treatment of this portion of the fauna, the stage of
development reached is nearly comparable to that in Merychippus
californicus of the Temblor. The Temblor mammalian fauna is repre-
sented, as is the Calvert, by materials derived from a marine sequence
and includes a strikingly similar assemblage, suggesting an ecologic
relationship in addition to one in time.

SYSTEMATIC DESCRIPTION OF THE MAMMALIAN REMAINS

Order CARNIVORA
Family CANIDAE
TOMARCTUS MARYLANDICA Berry

A second lower molar and the greater part of the lower carnassial of
a dog (U.S.N.M. No. 15561) were found by Charles T. Berry in
zone 10 of the Calvert formation, about 14 miles south of Plumpoint
Wharf, Calvert County, Md. These were described and given the
name Tomarctus marylandica by Berry in 1938. He noted a marked
resemblance of his form to that described by Merriam *° as Tephro-
cyon kelloggi from the Miocene Virgin Valley beds of northwestern
Nevada. The Calvert material appears inadequate for good specific
diagnosis. Nevertheless, the geographic remoteness of its occurrence
suggests a probable distinctness from previously described forms,
a matter for future collecting to demonstrate.

Order PROBOSCIDEA
Family GOMPHOTHERIIDEA

Considerable confusion has existed, and still exists, in mastodont
nomenclature, particularly as applied to the mastodon known to occur
in the Maryland Miocene. Details of its complexity in this instance
were not clarified by Osborn in his monograph of 1936.** In order
to illustrate the taxonomic procedure that the present authors have
followed, it is necessary to review briefly the origin of the name
“Mastodon” obscurus Leidy and the history of the use to which it
has been put.

10 Merriam, J. C., Univ. California Publ., Bull. Dept. Geol., vol. 6, pp. 235-238,
IQIl.
11 Osborn, H. F., Proboscidea, vol. 1, pp. 285-287, figs. 232 and 233, 1036.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Leidy,?? in his work on the “Extinct Mammalian Fauna of Dakota
and Nebraska,” described several mastodon teeth and figured three,
which were considered to be from various localities in the eastern
States. One of these, the first of the series discussed, was a cast of
a tooth (see fig. 3), the original of which was lost, but demonstrated
with some degree of probability by Leidy to have come from near
Greensboro, Caroline County, Md. As shown by the Maryland
Geological Survey *® this area is underlain by Choptank Miocene. A
second tooth of confused history included in the discussion was pur-
chased in London and purported to be American. Two incomplete
teeth from Tarboro, N. C., a fragment of a molar from an unknown
locality referred to as M. chapmani, and a tooth portion from Darien,
Ga., were also described.

In a later part of the paper Leidy ** proposed the name Mastodon
obscurus, citing the description beginning on an earlier page and the
_three figured teeth. However, he did not designate a type. Beneath
the synonymy the following statement is made:

Apparently a species distinct from the preceding (M. mirificus), indicated by
specimens from North Carolina and Georgia. Other specimens from unknown
localities supposed, however, to be American probably belong to the same. One
of the latter was referred to a separate species by Dr. Hays under the name of
M. chapmani, but Dr. Hays expresses the opinion to me that this is distinct
from the former. Under the circumstances I propose to distinguish the species
represented by the undoubted American specimens by the name heading the
article. The species has been suspected to be of Miocene age.... .

This discussion, constituting the designation of the typical and
referred materials, does not include Maryland specimens as indicative
of the species, but regards them as of dubious origin; hence the cast
of the lost tooth, believed to have come from Caroline County, is
eliminated from consideration as the type.

In keeping with the above limitations, Merrill** in 1907 selected
as type, by lectotype, one of the Tarboro, N. C., specimens, U.S.N.M.
No. 426. At this point the record is clarified, but in 1936 Osborn *°
stated that Merrill’s designation of the type was an error and that
the cast of the lost tooth is the proper type. He further indicated that
the Tarboro tooth really represents the species, Ocalientinus (Ser.)

12 Leidy, Joseph, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 7, pp. 244-
240, pl. 27, figs. 13, 15-16, 1860.

18 Maryland Geological Survey, Miocene, map facing p. xxiii, 1904.

14 Leidy, Joseph, ibid., p. 396, 1860.

15 Merrill, G. P., U. S. Nat. Mus. Bull. 53, pt. 2, p. 45, 1907.

16 Osborn, H. F., ibid., p. 286, 1936.

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS Ff

obliquidens, of somewhat later age. The taxonomic arrangement
made by Osborn is untenable, and we are forced to the following
conclusions: The Tarboro specimen is the type of “Mastodon”
obscurus, and Serridentinus obliquidens is a synonym of Serridentinus
obscurus (Leidy). “M.” chapmani Hays must be ignored, as there
is a strong probability that this specimen is of foreign origin. The
various mastodon teeth now known from the Miocene of Maryland
are without a specific designation. For these we propose the following
name:

GOMPHOTHERIUM CALVERTENSIS, new species

Type.—Left upper second molar, U.S.N.M. No. 13764.

Horizon and locality—Zone 13 or 14, Calvert formation ; in green
sandy clay about 4 feet above beach, 1.1 miles north of Governor
Run Road, Calvert County, Md.

Specific characters—Size of teeth approximating those in the
genotype, G. angustidens (Cuvier), from the Miocene of France.
Teeth low-crowned. Bunodont lophs of upper molar arranged trans-
versely, with well-developed trefoils. Anterior and posterior lobes
of trefoils partially distinct or cuspate. Lophs of lower molars more
arcuate and forward sloping, and with posterior lobe of trefoils better
developed than anterior, and with more noticeable separation. Slight
tendency to the formation of trefoil form on inner part of lower
teeth and outer part of upper teeth. Teeth noticeably resemble those
of G. (Serridentinus) obscurum from North Carolina, but decidedly
smaller and lower-crowned.

Description.—In addition to the type upper molar (fig. 1), which
was collected by M. H. White about 1931 and given to the National
Museum in 1935, there is a complete unworn last lower molar, also
from near Governor Run, collected by A. Hecklinger in 1931 and
deposited in collections of the Maryland Academy of Sciences. A
cast of this tooth (fig. 2) is in the collections of the National Museum,
U.S.N.M. No. 12134. The controversial lost tooth that Leidy re-
garded as coming from Caroline County, Md., and which Osborn
considered as the type of G. obscurum, is represented also in the
national collections by a duplicate (fig. 3) of the cast in the Academy
of Natural Sciences of Philadelphia.

The second upper molar of G. calvertensis lacks only the front half
of the anterior loph and the root portions. It is relatively unworn and
shows clearly the details of the cusp development. In occlusal view the
sides of the tooth are nearly parallel and the lophs almost at right

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

angles to the long diameter. The crests of the lophs are characterized
in the unworn condition by a series of rounded cusps. The trefoil plan
is markedly developed with the anterior and posterior lobes, between
the lophs, about equally developed and tending to be separated from

>>
W/ r B\ \)

Q\

“
a”
9 5)

Fic. 1—Gomphotherium calvertensis, new species: Left upper second molar
(U.S.N.M. No. 13764) type specimen, lateral and occlusal views, X §. Calvert
Miocene, Maryland.

the lophs at their summits. A slight, or subdued, trefoil arrangement
is also seen on the outer half of the lophs on the opposite side of
the clearly defined median crease or fold extending the length of
the tooth. In end view the inner and outer portions of the lophs are
of about equal height, except for the first loph where wear has

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 9

somewhat lowered the lingual half. The slopes of the inner and outer
columns are nearly equal, although the inner wall appears to be less
erect. No cingular crest is developed on the buccal wall and only
between the lophs lingually. The character of the forward margin of
the tooth cannot be determined, but the posterior margin exhibits

fe

A

i

)

i

Fic. 2—Gomphotherium calvertensis, new species: Drawing from cast
(U.S.N.M. No. 12134) of right last lower molar belonging to Maryland
Academy of Sciences, lateral and occlusal views, X 3. Calvert(?) Miocene,
Maryland.

a markedly cuspate crest across the midportion and about halfway up
the posterior loph, rising toward the inner extremity.

The last lower molar, belonging to the Maryland Academy of
Sciences, appears somewhat smaller than would likely have been
associated with the type upper molar but may well be within the
range of this species. It is also a little shorter and narrower than
the cast of the Caroline County specimen. The tooth is unworn and
complete except for roots. Resemblance to the corresponding tooth
of G. angustidens is striking, although the Maryland Academy tooth

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

is noticeably smaller than U.S.N.M. No. 150 from the Miocene of
France. These teeth, including the cast of the Caroline County
specimens, are alike in structural pattern with arcuate lophs in which
the outer trefoil portion is offset or directed backward, and the poste-
rior buttress or lobe of the trefoil is very well developed. This lobe
is even more prominent and shows better separation directly behind

SSS.
WW
SSN \ \\ NaH

\’

= Ss
U,
s

Zi hg
ef). ji,
BZ

Z: N e
yr # NV

Hits, an: \ | i,
oy é wil \ HIN

Fic. 3—Cf. Gomphotherium calvertensis, new species: Drawing from cast
(duplicate of Acad. Nat. Sci. Philadelphia No. 13278), lateral and occlusal views,
> 3. Original purported to be from Choptank Miocene, Maryland.

the first loph. It is essentially single in the Maryland Academy speci-
men but may be bilobed in G. angustidens. The posterior lobe of
the trefoil is less distinctly separate from the second loph but there
is a suggestion of bilobing near the base in the Maryland form.
Slight basal cuspules or weak and irregularly developed trefoiling
is seen inward of the median longitudinal cleft.

In critically examining the type and other materials of Serridentinus
productus and Serridentinus floridanus the structural arrangement is
scarcely different from that in typical Gomphotherium. Except for

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS II

size and height of crown, the molar teeth of Pliocene Serridentinus
in this country differ but little from Miocene Gomphotherium molars
at hand. We suspect that but one phyletic group is represented.

Measurements (in millimeters) of cheek teeth of
Gomphotherium calvertensis

Transverse diameter of M?, U.S.N.M. No. 13764 (type)...............- 70.5

Length of Ms, U.S.N.M. No. 12134 (cast of Maryland Acad. Sci. speci-
ENS END Mee eer We el eo Yevec cen wey Pave Shar EOS shaban oe eG Bie race Bai ctcia sie ov eishatals 140.

Gransverse diameter of Ms, U:S:NIM. No. T2134... 2... c....2ce secs cece 61.

Transverse diameter of Ms, duplicate of cast in Acad. Nat. Sci. Philadelphia. 60.

Order PERISSODACTYLA
Family TAPIRIDAE
TAPIRAVUS cf. VALIDUS (Marsh)

A right maxilla, U.S.N.M. No. 18372 (fig. 4), with P? and M?
preserved, but exhibiting the root portions or alveoli for the re-
maining cheek teeth, was found by J. E. Smedley, of the U. S.
Geological Survey, about 2 miles south of the Chesapeake Beach
resort. The specimen was discovered weathered-out on the beach, but
at this point only the Calvert formation is reported in the adjacent cliff
section so that there seems little doubt of its origin.

The teeth resemble to a marked degree those seen in the modern
tapir, but are very much smaller and appear to be relatively lower-
crowned. Their size is rather close to the dimensions given by
Marsh ** for Tapiravus validus from the “Miocene marl of Cumber-
land Co., New Jersey.” The Calvert specimen is tentatively referred
to the New Jersey species.

Schlaikjer,1® in his work on the tapir from the lower Miocene of
Wyoming, considered that Marsh’s form represented uppermost Mio-
cene or lower Pliocene time. However, from a consideration of
Tertiary stratigraphy in southern New Jersey it seems probable that
the type is from the marl of the Kirkwood formation, which is
considered to be very close in age to the Calvert formation in
Maryland. Except for clay lenses in the Cohansey formation, there
does not appear to be any beds of later date that could properly be
referred to as marl. Since we have some assurance that the Maryland
specimen is from the Calvert we may also reason that Marsh’s type

may well have come from the Kirkwood.

17 Marsh, O. C., Proc. Acad. Nat. Sci. Philadelphia, vol. 23, pp. 9-10, 1871.
18 Schlaikjer, E. M., Bull. Mus. Comp. Zool., vol. 70, No. 4, p. 249, 1937.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Comparison of the Calvert specimen with modern Tapirus shows
that its maxilla, though smaller, is relatively a little deeper, when the
lengths of the cheek tooth series are considered. The infraorbital
foramen opens anteriorly above the posterior part of P?, much as in re-
cent specimens ; also the notch between the zygomatic wing of the max-
illa and the maxilla proper is about opposite M*, and the anterior mar-
gin of the orbit is located above about the middle of P*. However, the

CFO >
NEN

:
y

Fic. 4.—Tapiravus, cf. validus (Marsh): Right maxilla with P® and M’*
(U.S.N.M. No. 18372), lateral and ventral views, 4%. Calvert Miocene,
Maryland.

outward face of the lachrymal is less modified, though actually nearly
as long, and the lachrymal processes are not nearly so prominent. The
lachrymal foramen is divided posteriorly, and the two openings are
close together and face posteriorly. In modern Tapirus skulls the lower
of these foramina, when separate, is often observed to face outward
beneath the posterior lachrymal process. The sulcus on the lateral
face of the ascending portion of the maxilla is shallow and confined
to the maxilla and probably the frontal, whereas in Tapirus this sulcus
is much more deeply impressed, involves the lachrymal bone, and

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 13

may extend as far down as the infraorbital foramen. A small segment
of the frontal bone in the Calvert specimen is seen wedged between
the maxilla and lachrymal, much as in the modern tapir, so that we
may infer frontal and nasal modification for a proboscis, although
probably less strikingly developed. Modification of this portion of
the skull was probably not greatly different from that shown by
Schlaikjer ?® for the Harrison tapir.

Measurements in millimeters

U.S.N.M. No. 18372
Ss — T. validus type
P4 or M?

Measurement ps M?
Anteroposterior diameter ......... 13.4 15.0 14.8 *
Mransverse diameter, .4......20% 05 16.4 16.0 17.5\*

*7 and 8} lines, respectively.

Family RHINOCEROTIDAE
APHELOPS?, sp.

Three isolated lower molars, representing two individuals of rhi-
noceros, were collected by Collins from a locality about 0.8 mile south
of Randle Cliff Beach, Calvert County, Md. Two of these, unworn
right and left (fig. 5a) molars, U.S.N.M. No. 18427, were found to-
gether on the beach, but with matrix attached which suggested their
derivation from a weathered zone high on the cliff, probably zone 14
or 15 of the Calvert. The third tooth, partially worn molar in jaw frag-
ment, U.S.N.M. No. 18428 (fig. 5b), was found in the same general
locality at a later date, and presumably represents a different individ-
ual. The worn tooth is considered to be a first molar and the unworn
teeth may well be second molars.

The teeth are of moderate size, broad, and appear fairly high-
crowned, particularly when viewed from the outer side. The tooth
which Marsh *° described as Rhinoceros matutinus from Squankum,
in Monmouth County, N. J., revised by Wood * as Diceratherium
matutinum, may not be significantly different in size from the Calvert
teeth, as indicated by Marsh’s illustrations, but appears appreciably
lower-crowned and exhibits a strong external cingulum not seen in
the Maryland material. The only other Miocene rhinoceros material

19 Schlaikjer, E. M., ibid., figs. 1 and 2, 1937.

20 Marsh, O. C., Proc. Acad. Nat. Sci. Philadelphia, vol. 23, p. 3, 1870. See
also Amer. Journ. Sci., vol. 46, pp. 411-412, pl. 10, fig. 4, 1893.

21 Wood, H. E., 2d, Bull. Geol. Soc. Amer., vol. 50, pp. 1968-1969, 1939
(abstract).

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

from eastern United States is that described by Colbert *? from the
Hawthorn formation in Florida. The material which the Florida
Geological Survey lent to Dr. Colbert for study included five lower
teeth, apparently of one individual, which he referred to Aphelops.
Again, these teeth do not appear to be significantly different in size
from those of the Calvert form but are much more worn, and are
described as having a strong internal cingulum, closing the inner valley
of the premolars. No premolars are represented in the Calvert material

|

\

Oa
a |
fi

NG
——~ Hiri
e "I

Fic. 5.—Aphelops?, sp.: a, Unworn left M2? (U.S.N.M. No. 18427) ; b, left
Mi (U.S.N.M. No. 18428). Lateral and occlusal views, & %. Calvert Miocene,
Maryland.

but the molars do not have the cingulum developed either internally
or externally.

Reference of the Maryland material to Aphelops is highly tentative.
The specimens furnish little or no information which would serve
to distinguish the form represented from Aphelops, and at the same
time the material is too incomplete to warrant certain generic reference.

Measurements in millimeters

My? (U.S.N.M. No. 18428) Mo? (U.S.N.M. No. 18427)
Anteroposterior Transverse Anteroposterior Transverse
diameter diameter diameter diameter
40.0 28.6 47.0 28.5

22 Colbert, E. H., Florida State Geol. Surv. Bull. 10, pp. 55-58, fig. 30, 1932.

—

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS I5

Family EQUIDAE
ARCHAEOHIPPUS, sp.

A slightly worn lower cheek tooth in a jaw fragment, U.S.N.M.
No. 18431 (fig. 6a), is the only specimen of the small brachyodont
horse Archacohippus known from these beds. It was found by
F, Stearns MacNeil, of the U. S. Geological Survey, a little less than
a mile south of Plumpoint, Calvert County, Md. The Calvert forma-
tion is the only Tertiary reported in the cliff section in this vicinity.

Fic. 6.—a, Archaeohippus sp., portion of right ramus of mandible with M:
(U.S.N.M. No. 18431); b, Merychippus sp., left lower premolar (U.S.N.M.
No. 12032); c, Merychippus sp., left lower premolar (U.S.N.M. No. 18430).
Lateral and occlusal view, X 1. Calvert Miocene, Maryland.

The tooth of No. 18431, apparently Mz, is rather simple in con-
struction, decidedly brachyodont, without cement, and with very
little evidence at its present stage of wear for any separation of the
metaconid and metastylid. A cingulum is developed only on the
anterior and posterior walls, where it rises internally to the parasty-
lid and hypoconulid, respectively. Its size is distinctly less than this
tooth in the paratype of Archaecohippus mourningi (Merriam) *
from the Barstow beds, but only a little less, and it is somewhat more
slender than in A. mourningi from the Temblor ** formation in Cali-
fornia. Size difference is not significant in comparison with isolated

23 Merriam, J. C., Univ. California Publ., Bull. Dept. Geol., vol. 7, pp. 427-434,

1913.
24 See Bode, F. D., Carnegie Inst. Washington Publ. 440, pp. 48-50, 1033.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

lower molars of Archaeohippus ultimus Cope*® from the Mascall
formation of Oregon. The tooth is longer and narrower than lower
cheek teeth in the type of Archaeohippus penultimus Matthew *
from the Sheep Creek horizon but appreciably smaller than in
referred specimens. The Maryland tooth is also, presumably, lower-
crowned.

The only Archaeohippus species known from the eastern region
is A. nanus Simpson ?* from the Thomas Farm fauna of the Haw-
thorn formation in Florida. The Maryland tooth is very close in
size to M2 in A. nanus. However, there is no assurance that the form
represented is this species, as the specimen is much too incomplete
to warrant specific diagnosis.

Measurements (in millimeters) of M2, U.S.N.M. No. 18431

Anteroposterior, diametet, <j)... 2.44 + e+e css se ih3
Transverse? diameter? cevitews ios cieisiueyse ssa eeiecias 7.6

MERYCHIPPUS, sp.

Two left lower premolars of the horse Merychippus were found
on the beach between the Chesapeake Beach resort and Randle Cliff.
One of these, an unworn tooth, U.S.N.M. No. 12032 (fig. 6b), was
discovered by John J. O’Brien, of the Soldiers’ Home, and the other,
U.S.N.M. No. 18430 (fig. 6c), worn for probably half of its original
length, was collected by Charles Welter, of New York City. Between
the two beach resorts only Calvert of the Miocene sequence is
represented in the cliff section, as shown by the columnar sections in
the Maryland Geological Survey’s report on the Miocene.

Two phalanges, collected by W. E. Salter, of the U. S. Geological
Survey, from the vicinity of Scientists Cliffs, a short distance north
of Governor Run, may also represent Merychippus. Scientists Cliffs
includes the lower part of the Choptank formation as well as the upper
part of the Calvert, so that although a Miocene age is assumed for
the toe bones they are not certainly Calvert.

Unfortunately, no upper teeth have been found in the Calvert, and.
lower teeth are less satisfactory for comparative purposes. Never-
theless, the developmental stage reached in the lower teeth affords
strong evidence toward conclusions as to the stage of the Miocene
represented.

25 See Bode, F. D., ibid., pp. 51-52, pl. 3, figs. 1-3.
26 Matthew, W. D., Bull. Amer. Mus. Nat. Hist., vol. 50, pp. 158-159, 1924.
27 Simpson, G. G., Florida State Geol. Surv. Bull. 10, pp. 28-31, 1932.

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 17

The two lower teeth have very nearly the same cross section and
must surely represent the same species. Together they demonstrate
the unworn or total length of the tooth and the enamel pattern at
about one-half the tooth length. Their construction does not represent
a very early stage of Merychippus, but demonstrates pronounced
hypsodonty and well-developed enamel folding. Both teeth show
cement-filled folds on the lingual side and discontinuous or a scattering
of cement deposition on the outer walls and in the median fold. The
teeth are of small size, but distinctly more advanced than Merychippus
primus from the Sheep Creek horizon. They more nearly approach
in pattern and proportions those of Merychippus californicus Mer-
riam 7° from the Merychippus zone of the Temblor. They are dis-
tinctly less advanced than material of this genus from the Barstow.

It would seem that the stage of evolution reached in the Calvert
teeth is approximately that of Merychippus californicus, but with a
simpler pattern than those illustrated. They may not be far removed
in time from the Merychippus horizons represented in the Mascall
and Virgin Valley beds of Oregon and Nevada but appear advanced
over the stage indicated by Merychippus gunteri and Merychippus
westont from the Hawthorn beds of Florida.

Measurements (in millimeters) of Merychippus sp. cheek teeth

U.S.N.M. No. 12032 U.S.N.M. No. 18430
Antero- Antero-
posterior Transverse posterior Transverse
21.0 LO 19.0 12.0

* About midsection of tooth length.

Order ARTIODACTYLA
Family TAYASSUIDAE
Cf. CYNORCA PROTERVA (Cope)

The presence of a distinctly small peccary in the Maryland Miocene
has been a matter of record since 1868, but since that time very few
references have been made to the occurrence. In the preceding year
Cope ?® described what he regarded as a new type of squalodont on
the basis of material found by James T. Thomas near his residence
in the eastern part of Charles County, Md., not far from the Patuxent

28 Bode, F. D., Carnegie Inst. Washington Publ. 453, pp. 39-63, figs. 1-6,
pls. 1-2, 1934.
29 Cope, E. D., Proc. Acad. Nat. Sci. Philadelphia, vol. 19, pp. 151-152, 1867.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

River—not the Ashley River, S. C., as Palmer *° indicated. Cope’s
description was based on characters of an upper canine of a peccary,
but included in the material mentioned were two squalodont cheek
teeth. In 1868 Cope ** announced that Leidy had called his attention
to the fact that the canine belonged to a small peccary, whereupon he
attempted to retain this name for the other teeth. The correction
was not noticed by Case,** but Kellogg ** later demonstrated that since
the distinctive characteristics of Cynorca proterva were based on a
peccary canine, obviously the type, the name was not available for
use as a squalodont, and in this he was supported by Hay.**

Leidy,®® on the other hand, followed Cope in retaining the name
for a squalodont and referred the peccary canine to his species,
Dicotyles lenis, the type of which is from the Ashley River deposits,
and also included additional peccary material from Charles County,
Md., which Cope *® regarded as Pleistocene and referred to the
modern species under the name Dicotyles torquatus (Pecari angulatus ).
We have not examined these specimens so are unable to verify their
age assignment. The Chesapeake Bay collections of the National
Museum include peccary material from both the Miocene and Pleisto-
cene, the latter apparently being referable to Tayassu lenis (Leidy).

The peccary material at hand from various localities in Calvert
County, along the cliffs adjacent to Chesapeake Bay, indicates a
noticeably small form which is distinct from species of Miocene pec-
caries known from elsewhere. Since Cope described Cynorca proterva
as coming from the Miocene of the same general area, and Leidy
characterized it as distinctly smaller than modern forms, it seems
probable, although no other canines are at hand, that our material
belongs to this species.

DescriptionAmong the materials here referred to Cynorca pro-
terva are some six isolated upper and lower cheek teeth and a lower jaw
portion, U.S.N.M. No. 18429 (fig. 7), including M, and M;. More-
over, some incomplete limb and foot bones may belong to the same
species. The lower jaw was found on the beach 0.7 mile south of

30 Palmer, T. S., Index generum mammalium, p. 213, 1904.

31 Cope, E. D., ibid., vol. 20, pp. 185-186, 1868.

32 Case, E. C., Maryland Geological Survey, Miocene, pp. 7-8, 1904.

33 Kellogg, R., Proc. U. S. Nat. Mus., vol. 62, art. 16, pp. 28-29, 1923.

34 Hay, O. P., Second bibliography and catalogue of the fossil vertebrates of
North America, vol. 2, p. 770, 1930.

35 Leidy, Joseph, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 7, pp. 384-
385, 1860.

36 Cope, E. D., ibid., vol. 19, p. 155, 1867.

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 19

Randle Cliff Beach, where the Maryland Geological Survey indicates
only Calvert.

Two premolars, possibly P2 and Ps, but not of the same individual,
are simple and smooth with the elevated crest of the trigonid and the
lower talonid divided but free of accessory cuspules. The paraconid
is weak on the smaller of the two teeth, but prominent, though lower,
on the other. These premolars, if properly referred to the same
species as the molars, are relatively large.

Fic. 7.—Cf. Cynorca proterva (Cope): Right ramus of mandible with Ma
and Ms (U.S.N.M. No. 18429), lateral and occlusal views, X 1. Calvert Miocene,
Maryland.

Two last upper molars are strikingly smaller and relatively nar-
rower than in the living collared peccary, the cusps are more nearly
conical, the cingulum is much less developed, and there are fewer
accessory cuspules.

The lower jaw, No. 18424, is relatively shallow and exhibits rather
smooth, narrow molars, particularly Ms, with cusps, as in M*, more
nearly conical than in modern peccaries. It should be noted, however,
in comparison with modern forms that the large white-lipped peccary,
Tayassu pecari, exhibits teeth having less tendency toward the for-
mation of transverse crests, lower trigonid portions in the premolars,
and more nearly conical cusps in the molars than the smaller Pecart

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

angulatus. The absence of cingular cusps between the external cusps
of the lower molars, noted by Leidy for T. lenis, is also characteristic
of T. pecari, but not P. angulatus.

The dental characteristics noted for the form Cynorca are much
more prophetic of the modern peccaries than are characters of the
better-known Pliocene species of Prosthennops. Although it is pos-
sible that Prosthennops is a synonym of Cynorca it seems more
likely that the two are distinct and that Cynorca is actually in the
line of descent for Tayassu and Pecari. As noted by Colbert,37
Prosthennops does not fall in this line and Platygonus has become too
specialized in its dental characters. On the other hand, “Desmathyus”
and the closely related form, Floridachoerus, are known only from
large species, having teeth so modified that it seems unlikely they
in turn could have given rise to Cynorca.

Measurements in millimeters of lower jaw

(Cf. Cynorca proterva, U.S.N.M. No. 18429)

Depth of jaw beneath point between M2 and Ms................- 24.0
Anteroposterior diameter: transverse diameter, Mo............. rae: 0.95
Anteroposterior diameter: transverse diameter, Ms............. 1.47: 0.86

HESPERHYS (DESMATHYUS) ? sp.

A peccary of considerable size is represented in the collections by a
humerus, the distal portion of a radius, several carpals and phalanges,
and some vertebrae and ribs, all belonging to one individual. These
were found by Dr. W. F. Foshag, of the U. S. National Museum, in
1940 in fallen blocks from zone 17 of the Choptank Miocene immedi-
ately above the Calvert, at Calvert Beach, Md. Three isolated meta-
podials from along the Chesapeake are of a size comparable to the
foregoing. Two of these are not documented as to horizon, but the
third was collected by Collins from zone 19 of the Choptank.

There is no certainty as to the genus represented, but it is thought
that the form might possibly be “Desmathyus,’ as a peccary from
the base of the Miocene Kirkwood formation in New Jersey, which
Marsh *® described as Perchoerus (Dicotyles) antiquus, has since
been referred by Wood * to “Desmathyus.” Moreover, “Desmathyus”

87 Colbert, E. H., Nebraska State Mus. Bull., vol. 1, No. 44, p. 421, 1935.

88 Marsh, O. C., Amer. Journ. Sci., vol. 46, p. 411, 1893.

39 Wood, H. E., 2d, Bull. Geol. Soc. Amer., vol. 50, pp. 1968-1969, 1039
(abstract).

NO. 2 MIOCENE LAND MAMMALS—GAZIN AND COLLINS 21

has been described from the Merychippus primus zone of the Snake
Creek beds as well as from the Upper Rosebud and Harrison.

The humerus and other elements of this peccary are as large and
massive as in the Pleistocene form, Platygonus cumberlandensis. No
limb material of true Hesperhys was available for direct comparison
and these seem undescribed in the literature.

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Fa Naa
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 3

tWO RUNIC STONES, FROM
GREENLAND AND MINNESOTA

BY
WILLIAM THALBITZER

(Pusrication 4021)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 30, 1951

iW,

The Lord Baltimore Press

BALTIMORE, MD., U. 8 A.

TWO RUNIC STONES, FROM GREENLAND
AND MINNESOTA *

By WILLIAM THALBITZER
Copenhagen, Denmark

CONTENTS

Page
er its COL Cd OT cee ves eae ae es 01G Os SENG ok ous in sece Sys ETL HEU PAC Tey aT kW oo eusears 2

I. The Kingigtorssuaq stone from the neighborhood of Upernavik,
Moreawest “Greenland So. 2. cn eee See es Pa eed era aten 6
Mine he wicensinetom StOLeiasieia cs .tdosctant <r meatal ae Poles eit els GaeieIane 14
Hustoricalubackerounid) >< kevin verte ten si erasous atae serestoe piiene 14
The Kensington runes and the numeral signs.................-2. 18
PAleCOPTADMICHSIPNG saci secre cintec high hiticte otal kote eee Te 23
he Walecarhian runes and the El-rane. ....-co+css se tee cee = Caer 26
MER IiTISTIStIC ALOLIM ed oicteva ce arse one cre ee eee ie enone eke Oe oetee 28
OldeSwedish: Bit) 2k Sheek cece. elses Ate eben ees 29
PNG IPUNSITIST Oy 5, cca tacyoickehe Seah se seretns seen ae WarsPiss ero Gacnci need cleus fake Meus ebb exch 32
INGO OSES ITS way egos. & ayannys &, 7)s.5s5 wi ogni ahs, AS Se oy oeoyanntovensie isl ese aa ars aetereneatoeres 35
SIWIAG ERS EVIE vestte nc yaya ere sani ede evar? crete siclioerepanetonets sts, ore + enehsvayaguya mus etaetehe arctan 39
Barthes piilolopicaly ampnressionsw scree cee oe ceieiare as ete rele 42
PAPEL Ie LOS O) Maa ereaie votes tchasurets, eee etoee ONS chee wie Heiale STAelcal ate SFM eeaoieEs 49
Mitee, MiStOniCis GOCUIMENES as ch. das wraciaye Seveporew ioe o oeietdiolelepeliertus ovestatwethe 49
Final comments on the Danish treatise (1946-1947) ..........eeeeeeeee 51
GRE > ea Re eee they ee cere A gs i eS ek I Si 54
1. The Greenland runes as carved on stone or wood in Greenland.. 54
2. The secret runes on the Kingigtorssuag stone...............-. 54
3. Singular runes on the Kensington stone, K and f .......... 56
Gy eh a sacl eeteyvicsens uncraes Maaaeriant veucias Susi shmne See oners 57
eA ROISCLIN oa ne cat eie Gariee ake this otek iat ere eee aoe 57
G HAE Xe shior or 2) (Ene. sear, “CHih, POCK)) <biinececncien se 58
7. rise = Modern Sw. resa, “journey, (warlike) expedition”..... 59
8. at se adptir, “to look after, guard, superintend (the ships)”.... 61
G) CAG ie 85SEC." Kday 7S JOULNEY): «[tavarfoysters stave se cele sholalo\wters le «tyes epesateus 61
TO Wm Heat SEG. eich cals ciate. chaicrevcyu lo cians rere uaneisnors Gig, tists avi via ch cree eee has 62
11. Medieval Swedish-Norwegian mixed language................ 63
125 Olds Swedish as) aenypotetical pPrODleMl. ah ~ tos are wieie es ate 64
13. A final comparison between the two runic inscriptions........ 65
PADS liy he Peery ett. hereto tas sta adeteabste ste anh ete eet epoiapern bl nut ly. « prvpsidye Welaket 67

* This paper, by a distinguished Danish authority, is published by the Smithso-
nian Institution as an interesting contribution to a discussion of a subject that, so
far as it relates to the Kensington stone, has aroused much controversy.—EbITor.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 3

AUG 3 0 195)

INTRODUCTION

nysta ek niér,

nam ek upp runar

cepande nam;

fell ek aptr badan.
(From Hovamegl.)

Of the two most interesting and puzzling runic stones known to
me, one is from the small island of Kingigtorssuaq, “towering moun-
tain,” situated among the skerries about 10 miles north of the colony
of Upernavik in northern West Greenland, in latitude 72°58’ N.,
longitude 56°14’ W.; the other is from Minnesota near the village of
Kensington, situated about latitude 47° N., longitude 96° W., west of
the Great Lakes. Both stones tell of expeditions of discovery, one
starting from the Norse settlements in Greenland and reaching far
north, the other starting from Scandinavia and traveling by way of
Greenland to Vinland, and from there west.

The first stone was found in 1824 by Pelimut (i.e., Philemon), a
Greenlander, and the same year was sent to the museum at Copen-
hagen. It was very small, only 10 cm. in length. All runologists ac-
knowledge the authenticity and old age of the inscription; it is esti-
mated to date from about A.D. 1300.

The other stone, the American one from Minnesota, was dug up
in 1898 by a Swedish farmer named Olof Ohman when grubbing
trees on a hill, formerly an island, on his land. He found the stone
clasped by two large roots of an aspen, where, to judge from the age
of the tree, it had been buried for many years. It is a longish stone
(78.7 by 40.6 by 15.2 cm.) weighing nearly 200 pounds. The inscrip-
tion is long, one of the longest runic inscriptions known, and by a
kind of “runic figures,” i.e., pentathic numeral figures (used accord-
ing to our Arabic numeral system), it dates itself in the year 1362.
The 221 (222) large, clear runic signs are arranged in nine horizontal
lines on the face and three vertical lines on the left edge of the stone.
Geologists who have inspected the stone state that the runes bear the
stamp of a very long period of weathering.

But in the contents of the inscription there were certain peculiari-
ties that went contrary to this impression. Hence the authenticity of
the inscription was seriously questioned by two American runologists,
Prof. O. J. Breda, of the University of Minnesota, and Prof. G. O.
Curme, of Evanston, Ill., who 50 years ago first saw the stone or
photographs of it. The inscription appeared to contain English words
such as from, of, ded, illy, and several other peculiarities such as

2

NO. 3 TWO RUNIC STONES—THALBITZER 3

mans, “men,” rise, “journey, voyage,” etc., which seemed to them
absolutely inexplicable on a runic stone. Certain statements made
locally unfortunately led to suspicion being directed to a Swedish im-
migrant, a friend of the farmer who had found the stone. He was a
schoolmaster wandering in certain parts of Minnesota, originally an
unfrocked clergyman from the province of Sodermanland, “who had
perhaps half forgotten his mother tongue.” He was even said to have
possessed a Swedish textbook in which a runic alphabet was printed.
The man had died a few years before the stone was found.

Since the first critical treatment, no thorough philological investi-
gation has been made to this day, perhaps with one exception, which
will be mentioned presently. It is true that the stone was sent to ex-
hibitions in France and Norway, where it occasioned animated dis-
cussions in the newspapers; but the real experts of Scandinavia one
and all brushed aside the view that the stone could be genuine. This
was in the years immediately after 1908, when the Norwegian-
American Hjalmar R. Holand, living in Wisconsin, had obtained the
stone from Ohman and taken up the study of runes and related sub-
jects. He first wrote about the find in Skandinaven (January 17,
1908), one of the leading newspapers of Minnesota. He was of the
opinion that the philologists had judged the inscription on erroneous
supposition that it belonged to the old Vinland period and was written
in Old Icelandic. That it was found hidden in the earth so far from
the Atlantic coast was a further reason for rejection. The last point
was, indeed, difficult to account for. But Holand was able to offer a
surprising explanation, which justified both the year 1362 and the re-
mote place of discovery (see p. 16 f.). It seems, however, that the
verdict by the professors that the stone was a “clumsy fraud” had
made an indelible impression. The weighty scientific condemnation
and the distortion of the facts in the papers were like a dead weight on
the proper investigation of the problem. Charges of that kind are
hard to silence.

The suspicious stone was first placed in a bank in Alexandria,
Minn., where it remained for a number of years, still tabooed by the
runologists ; but in 1948 it was moved to the United States National
Museum under the direction of the Smithsonian Institution, in
Washington, D. C. Meanwhile the indefatigable Holand had con-
tinued his private studies on Scandinavian runes, medieval dialects,
and Scandinavian archeology, and in the course of 40 years he has

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

written three very thorough books on the Kensington stone and the
Newport tower, besides some shorter articles.*

For a long time I, too, had considered the Kensington stone a
fraud, and the late Prof. Finnur Jonsson and other Scandinavian
runologists confirmed my view. However, from time to time certain
fresh facts bearing on the matter have come to light, in archeology,
runology, and philology, especially Prof. Axel Kock’s later studies on
medieval Swedish dialects. As new light is gradually being thrown
on this amazing find from the West, I cannot but waver in my doubt
and am forced to see the question from a new viewpoint. Not only
Holand’s books but my own investigations as well have set me think-
ing along new lines.? I now maintain that this matter in its entirety is
worthy of restudy; it seems to me that, after all, the inscription may
be authentic.

I shall not dwell on the great number of discoveries of presum-
ably medieval antiquities made near or in the general neighborhood
of the locality where the Kensington stone was found, in Minnesota
and elsewhere (described in Holand’s books) ; this must be left to
the archeologists. I shall keep to the inner linguistic and historical
criteria of the inscription and try to state the evidence that seems to
me to speak for its linguistic credibility. This, perhaps, will provide
encouragement to others more competent than I to give their attention
to the problem once again.

Acknowledgments.—I am greatly indebted to Gunnar Knudsen,
M. A., editor of the Danske Studier, for many helpful comments
during the preparation of the manuscript of my paper (in Danske
Studier, 1946-47). During the preparation of the English translation
and the appendix to the present work, I collaborated with Vilhelm
Marstrand, Secretary to our new Academy of Technical Sciences,
who, I found, possessed an intimate knowledge of everything written
about the runes. He has himself examined a great many runic in-
scriptions and interpreted them in a very independent manner. He
has given numeric magic particular attention. The results he put

1 Holand, Hj. R., The Kensington stone, a study in pre-Columbian Ameri-
can history, pp. 75, 78 ff., 1932 (2d ed. 1940) ; Westward from Vinland, 1940;
America 1355-1364, 1946.

2 One competent observer who has not taken a hypercritical attitude like most
other Scandinavian philologists is the late Prof. Hjalmar Lindroth, of Goteborg.
In a letter to Prof. Richard Hennig (author of Terrae Incognitae, vols. 1 to 3,
1938), published in Petermann’s Geogr. Mitteil., pp. 89-90, 1938, Lindroth
weighs the evidence pro and con in an unbiased manner, admitting that hitherto
the investigations have been superficial. “The last word has not been spoken,”
writes Lindroth.

{i i

ae WY Even
it ae
MA Stak ff

yee BE, - ,
Pula Qterreléie

Fic. 1—The Kingigtorssuaq stone from northwest Greenland. (From
Grgnlands Historiske Mindesmeerker, vol. 3, pl. 9, K@benhavn, 1845.)

Moen Ch
Taek Leng
EAE i!
Y AARC i

Fic. 2—The Kensington stone from Minnesota. Left, front view; right, side
view. (From Hjalmar R. Holand, 1932, figs. 1-2.)

NO. 3 TWO RUNIC STONES—THALBITZER 5

before me, a continuation of the studies of Magnus Olsen and others
of numeric ‘‘magic” in runology (cf. p. 54 f.), impressed me so
much that I could not disregard the possibility that the runes of the
Kensington stone might present a similar secret numeric system. Mr.
Marstrand, it is true, had at first been skeptical concerning this
possibility. The precarious conditions under which this inscription was
carved made it seem improbable that it should have been adapted to
the usual system of numeric magic. However, on my suggestion he
made a fresh investigation of the number of runes and points of the
inscription, and arrived at a positive result. I wanted Mr. Marstrand’s
account to be included above his own signature in the appendix to this
paper, but this did not prove feasible. If his theory is correct his result
will present a cogent criterion of the genuineness of the inscription.

I. THE. KINGIGTORSSUAQ. STONE FROM THE
NEIGHBORHOOD OF UPERNAVIK,
NORTHWEST GREENLAND

The Kingigtorssuaq stone is a blackish-gray or dark-green quartz
slate, with a smooth surface for the inscription, found on stony ground
beside a very old ruined cairn in which the stone was probably once
placed. Close by, at the top of the same island, two somewhat smaller
cairns are found, so that the three cairns form the corners of an
equilateral triangle.

These cairns may have given the bearings of the permanent resi-
dence of the people who had erected them as being on another island
or on the mainland. The same year it was found, the small runic
stone—undoubtedly one of the smallest in the world, but at the same
time the most significant of all runic inscriptions yet found in Green-
land—was sent down to Copenhagen by W. A. Graah on his first
voyage to Greenland (1823-24).

Rasmus Rask interpreted the inscription, and on November 2, 1824,
wrote an account of it—brief, but for the time expert and penetrat-
ing—which was published by Prof. Finn Magnusen in the Antiq-
variske Annaler.* Rask’s interpretation has stood its ground, as it is
repeated practically unaltered by the later interpreters of the inscrip-
tion, Finnur Jonsson (1914) and Magnus Olsen (1932), except that
they hesitate to accept the interpretation of the last words and the
secret runes. (The last word is followed by six unusual and unex-
plained runic signs, probably some magic letters. )*

Nobody has ever doubted the authenticity of this runic inscrip-
tion, although Finnur Jonsson has considered the possibility of its
being of later origin because at the time of the discovery there was
an Icelander in north Greenland who was for some time employed
as manager at Godhayn on Disko Island and as such was accountant-

’ Rask, in the Antiqvariske Annaler, vol. 4, pp. 309-342 and 367-378, 1827,
quoted by Finn Magnusen, who also later wrote about the inscription in
Grgnlands Historiske Mindesmerker, vol. 3, p. 843, 1845. Cf. Rafn, in the
Antiquitates Americanae, pp. 347-355, 1837, in both places with drawings of the
stone (pls. 8 and 9). Cf. also Pingel, in Nord. Tidsskr. f. Oldkyndighed, vol. 1,
p. 108, 1832, and Liljegren, Run-lara, p. 157, note, and p. 211, 1832.

4 Jonsson, Finnur, Runestenen fra Kingigtorssuaq, Grgnlandske Selskab
Aarsskrift, 1914 (cf. ibid., 1916 and 1922). Olsen, Magnus, Kingigtérsoak-
stenen og sproget i de grénlandske runeinnskrifter, Norsk Tidsskr. f. Sprog-
videnskap, vol. 5, 1932.

6

NO. 3 TWO RUNIC STONES—THALBITZER 13

manager of Upernavik. This Icelander, however, did not get so far
north until 1825, the year after the runic stone had been discovered.
He was a member of the Royal Scandinavian Ancient Text Society
(Det Kongelige Oldskrift Selskab). It may be added that the first
missionary at Upernavik, sent there in 1779, was also an Icelander.°
The fact that the strange cairns there had long ago aroused attention
among the Greenlanders and Danes in north Greenland appears from
statements in Paul Egede’s Greenlandic Journal (printed in 1780),
about which I have written elsewhere.®
Rask’s interpretation runs as follows:
Ellingr Sigvab sonr ok Bjanne Torterson

ok Endridi Osson laugarSaginn fyrir gangdag
hl6Su varda te... ok ruddu MCXXXV

Erling Sighvatsson and Bjarne Thordarson and Endrithi Oddson Saturday
before “gangdag” raised the(se) cairns ...and “rydu”......

“T attach no importance at all to the explanation of the date of
year,” says Rask, and Finn Magnusen queries it.’

In Finnur Jonsson’s rendering the inscription shows the following
form and interpretation:

(a) ellikr: sikvabs: son: r. ok. baanne: tortarson:
(b) ok: enribi: osson: laukardak. in: fyrir. gakndag

(ce) hlobu. wardate. okrydu: ... (7)...
“Erlingr Sigvatssonr ok Bjarne borSarson ok Enridi Oddson laugardaginn
fyrir gagndag (—gangdag) hlodu vardu Se... ok rydu?...”

And he renders it in Danish thus:

Erling Sigvatsson og Bjarne Tordsson og Enride Oddson opfgrte Igrdagen
f¢dr gangdag denne (disse) varde(r) og...

As appears from the inscription (fig. 1) there is an initial sign in
the lines (a) and (b) that has not been considered in the interpreta-
tions of Rask, Finn Magnusen, and Finnur Jonsson, all having con-
sidered these “initial signs” as ornamental strokes without any
meaning.

5 Egede, Paul, Efterretninger om Grgnland, p. 249, 1788, and Jonsson, Finnur,
IGG, (Cit, p. 321:

6 Thalbitzer, W., Nordboerne ved Upernavik, Grgnlandske Selskab Aarsskrift,
1945; Jonsson, Finnur, Grdénlandske runestenen, ibid., 1916.

7 Antiqvariske Annaler, vol. 4, pp. 311 and 318, 1827. Notoriously the ancient
knowledge of runic and numeral magic was quite lost in the days of F. Mag-
nusen and R. Rask, and their interpretation of the secret signs (“runes”) of
this inscription is guesswork.

The first ones to search the old lore about these matters were the Swede
F. Laffler and the Norwegian M. Olsen. (Cf. pp. 9-10 and 55.)

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Finnur Jonsson describes their forms and adds: “These signs are
only ornaments without any literal meaning” (i.e., they are not
letters).

The two first-mentioned interpreters termed them “initial orna-
ments.” In the case of the final signs, the six secret runes in (c)
(fig. 1; pp. 7 and 54 f.), both Rask and Finn Magnusen, although with
some doubts, would read an indication of a year as if they were Latin
letters (MCXXXV), an otherwise unknown phenomenon on our
early rune stones. On the other hand, this feature, a date in connec-
tion with a communication, might very well be expected on a stone
like this, which does not belong with the commonest kind of runic
stones such as tombstones, but may rather be compared to the calendar
statements, diplomas, and other dated documents of the time, where
a year is generally given at the top, more rarely at the end.® As this
inscription, like most diplomas, states a day—the Saturday before
“sangdag’’—there would be reason to expect a year, too. Without this
the dating would be rather absurd. Finn Magnusen gives his opinion
as to the exact meaning of the statement of the day: According to
the old calendars and other Catholic sources it refers to the 25th of
April,® a time of year when the sea between the skerries in northern
Greenland is always frozen over. Thus any possibility for the three
men wintering there to escape home in their vessel must have been
excluded. They were obviously badly in need of help and eagerly
awaiting the breaking up of the ice, which never takes place until the
end of June or the beginning of July. The miserable men then
must have erected the cairns, partly as signals to draw the attention of
their countrymen in the south to their whereabouts and partly as an
account of their expedition to be left behind if they were to perish.
The cairn with the runic inscription on the small stone, then, was
erected by the men as a possibly posthumous bit of information for
those who missed them and perhaps would search for them.

Rask and Finn Magnusen were unable to offer any convincing in-
terpretation of the concluding secret runes of the inscription. The
interpretation of these as denoting a year was based on guesswork,
although they tried to connect their conjectures with the information
they thought it possible to adduce from ancient manuscripts and
calendars. Finnur Jonsson gave up the interpretation of the mystical
signs.

8 Diplomatarium Norvegicum, Kristiania, vol. 1, 1847, wherein No. 237 ends
like this: “Septimo ydus Martii, anno domini M°CCC° XXXVJ°” (i.e., March
9, 1336), and No. 252: “m®°ccc® XXXqvarto idus Julii” (i.e, July 12, 1334).

® Magnusen, Finn, im Antiqvariske Annaler, vol. 4, pp. 318-320, 1827.

NO. 3 TWO RUNIC STONES—THALBITZER 9g

But Magnus Olsen, the great Norwegian runologist of our day,
undertakes to explain the signs as a disguise of a magical word that
might fit into the preceding context.° In his opinion these secret
runes are intended to complete the meaning of the last runic words
proper, okrydu, i.e., ok ryddu, which, having been written in haste,
have a punctuation mark missing. Magnus Olsen reads this as ok
ryndu “and runed,” i.e., “and wrote runes”—with the same y-rune
as in fyrir, “for,” and with a substitution of md for t, which has sev-
eral analogies in runology (cf. also here ellingr for ellikr), and which
seems to me a fine and ingenious reading. Then he interprets the
secret runes as vel or vit, thus reading it all as “(and runed) well,
or widely.”

I am sorry that I must consider the latter part of this solution—
his interpretation of the secret runes—improbable; it seems to me
to be too artificial and the resulting word too insignificant, its mean-
ing too unsatisfactory. In the first place one wonders why such
an adverbial addition as the innocent little word “well” or “widely”
should be expressed by magical runes. Further, Olsen’s interpreta-
tion of the individual runes seems to be difficult and far-fetched and
is surely not correct with reference to the last sign, which cannot
be J.

There is a former interpretation by Fritz Laffler that seems more
probable to me.*4 He, too, had studied the secret runes of the
Kingigtorssuaq stone and points out (as also mentioned by Magnus
Olsen) a striking correspondence of these with the secret runes on
the Norum font from Bohuslan, a similarity that cannot be disre-
garded. He reads the secret runes of the Kingigtorssuaq stone as
is, “ice,” interpreting the preceding rydw (in agreement with Rask
and Finn Magnusen) as ruddu, the whole thus reading: “they cleared
away ice.” In my opinion this would be an acceptable meaning, but
it is linguistically unwarrantable to read ruddu when the stone actu-
ally had rydu (or rytu). I decidedly prefer Magnus Olsen’s reading
ryndu, which would give a better sense than that adduced above; for
there would be a clear purpose in the suggestion that the men ryndu

10 Olsen, Magnus, Kingigtérsoak-stenen (etc.) im Norsk Tidsskr. f. Sprog-
videnskap, vol. 5, Oslo, 1932.

Later, Olsen came to another conclusion. Cf. Appendix, p. 54.

11 Laffler, Fritz, at the annual meeting of the Swedish Antiquarian Associa-
tion, March 1906 (Fornvannen, Stockholm, 1906, p. 52). Cf. Olsen, Magnus,
Norsk Tidsskr. £. Sprogvidenskap, vol. 5, pp. 194-198 and 200, 1932, and Nansen,
Fridtjof, Nordpa i Takeheimen, p. 227, with reference to Svenska Dagbladet,
March 14, 1906.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

is(?), “runed (for) the ice,” i.e., conjured it away with runes to
make the water open so that they might escape in their icebound boat
from the long involuntary winter stay and go south to the home settle-
ments. Anyone who has traveled in that region and has been ice-
bound, even on a summer’s day, at the mouth of the large ice fjord
north of Upernavik, will easily realize how hopeless the situation
must have appeared to the three icebound men after their confine-
ment during the long winter. In all directions the sea must still have
been frozen up to the end of April, while it was light during most
of the 24 hours of the day as in south Greenland in summer. The
three men must have been longing for deliverance. Also the Eskimo
knew the use of magic in making the ice break up and go away.

The Norsemen had the same custom as the Eskimos, namely, not
to mention by its true name the object to which one applies the magic,
or the animal one wants to catch at sea—a kind of name taboo. When
hunting at sea the Eskimos use substitute or circumlocutory words to
denote the game in order not to scare it away by betraying their own
intentions. Thus also when the ice was to be attacked ; they must not
mention the word “ice” or spell the word with the ordinary runes. Here
magic signs were called for to attack the hostile ice: ok ryndu is (is
to be written with magic signs). Linguistically the last-mentioned
form may be open to criticism, viz, someone might maintain that
rynda (from ryna, a verb derived from the stem of riinar, “runic
letters”) is an intransitive verb; if so, it would be expected that és
would be put in the dative, ist. But ryma (imperfect ryndi) was a
rare word, and, after all, might it not be used in an intransitive sense?
Otherwise, ryna isi, or eptir ist.

All interpreters consider the language of this inscription to be Ice-
landic or even (Magnus Olsen) Greenlandic-Icelandic, a proper
Norsemen’s language. Rasmus Rask only wrote that it “is pure and
proper Icelandic, but written carelessly according to the pronuncia-
tion of an unlearned man.” Rask rightly regarded most of the forms
of the personal names as being in accordance with modern Icelandic
pronunciation. He considers ellikr (of the inscription) to stand for
Erlingr ; sikvaps for Sigvats ; baanne (bjanne) for Bjarne; tértarson
for borparson; enridi for E(i)ndridi; osson for Oddson. Thus he
constructs the old Icelandic forms on the basis of those of the stone,
which really look quite modern. Ellingr, Osson, Bjanne “are very
close to the modern Icelandic pronunciation.” 1 He further writes
that “te for be are undoubtedly the initials of the word Penna,’ the

12 Cf. also Finn Magnusen in Antiqvariske Annaler, vol. 4, pp. 314-316, 1827.

a

NO. 3 TWO RUNIC STONES—THALBITZER II

sense of the final line being: “erected this cairn” (etc.). Finnur
Jonsson and Magnus Olsen accept without reservation Rask’s read-
ing of the three men’s names but prefer to interpret varda (varda)
as in the plural, since, as we know now, there were three cairns on
the top of the same islet. Hence they read pe as pessa, “these
(cairns).”

On the other hand, Magnus Olsen looks on the linguistic forms of
the inscription and the rune carver’s relation to the language some-
what differently from Rask. We now, with Olsen, feel that the in-
scription testifies to a thorough insight into the use of the runes; it
must have been an extremely skilled rune master who carved it—in
spite of a few peculiar slips or mistakes, such as the separation of r
from the stem of son:r, the omission of the punctuation mark in
okrytu (for ok rytu), and t for d in the last-mentioned word, but ¢
for ) in Sigvap, te for pe in tessa (pessa), etc.

The runes, according to Magnus Olsen, belong to the late rune
alphabet, “which made the correspondence with the Latin alphabet
Complete .. .” 1

And at the end of the same treatise he says (p. 230): “The
Kingigtorssuaq inscription was written by a man with a considerable
training, and it is imaginable that this training was acquired not only
by carving of runes, but also by occupation with texts in the mother
tongue written with Latin characters.” So much here for Magnus
Olsen’s interpretation.

A feature that has not been pointed out by the interpreters of this
inscription, but that seems important to me, is this: the rune carver
mentions the three men in the third person, none of them as identical
with himself.

12a Norsk Tidsskr. f. Sprogvidenskap, vol. 5, p. 250. Here he continues as fol-
lows: “Unfortunately we have no criteria to decide in which settlement and in
which milieu the man belonged who carved the runes on the Kingigtorssuaq stone.
It would seem that he was a prominent man who not only distinguished himself
by courage and enterprise, but also readily devoted himself to the spiritual pur-
suits to which he might have access under the restricted conditions in a remote
and isolated country. It seems to me indubitable that he had his home in the
East Settlement (Eystri-bygd), the southernmost settlement on the west coast
of Greenland, and I should think that he belonged to or was in touch with the
leading circles there. This agrees with the whole character of the inscription,
its literary stamp and, further, the runic art with which the inscription was made
from first to last. ... But a written Greenlandic literature seems to be out
of the question. Even in a runic inscription written so excellently as that on
the Kingigtorssuaq stone, living speech greatly asserts itself.”

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

A most remarkable feature is the first rune, which has been inter-
preted as an e (ellikr-ellingr), this e here having been made into a
dotted rune, i.e., provided with two internal clearly separated dots, @
(see fig. 1), but not repeated later in the inscription. Might it not be
possible that the expert, perhaps learned, rune carver by this means
wanted to denote a special variant of e, perhaps 6 or a diphthong ey
or oy? 38

Olsen makes much of the two dots, being of the opinion that
the rune carver wanted in this way to characterize the initial letter
as a “golden number” in the calendar. In the year 1333 the golden
number of Easter Day belonged to the 4th of April; therefore the
gangdag would be Sunday the 25th of April. The “Saturday before
gangdag” then would correspond to the 24th of April. Magnus Olsen
concludes that the cairn was built and the inscription made on April
24, 1333. If so, this is a “historic inscription.” (Olsen, loc. cit.,
p. 226.)

Thus there would still be a year indicated on the stone, an extremely
rare occurrence on a runic stone from a medieval period, the begin-
ning of the fourteenth century. The rune carver obviously was not
quite so illiterate as thought by Rask. Nor can it be denied that the
historical experience and the accurate calendar date of the inscrip-
tion almost certainly must presuppose the statement of a year. If it
had been written on vellum, one might have expected the application
of Latin monastic letters, which presumably were well known to the
rune carver from annals and similar documents stating years. In-
deed, this occurs here and there in late medieval script in Scandinavia,
at any rate in annals, calendars, and diplomas.

But let us once more look at the silent initial signs of the first
lines.**

The two initial secret runes or unknown “stave signs” A, |§
apparently are comparable with Ogham, twig runes, nail script, etc.

13 A similar name is found in a Norwegian diploma. See Unger and Lange,
Diplomatarium Norvegicum, Kristiania, vol. 1, No. 163, p. 142, 1847; cf. facsimile
No. 2, p. 4: Ollingi-Ellingi. An Eylingr perhaps might also be imagined in
Greenland (a patronymic?).

14 The intial sign in line 1 resembles a corresponding one in a late Dalecarlian
runic inscription; it is seen on a “food bowl made of curly-grained wood.” Cf.
Erixon, Sigurd, Runeskrift fran Dalarna, Fataburen, p. 151, fig. 4, 1915. For
that matter, it also resembles the old -rune known from the Kallerup, Snoldelev,
and Helnzs stones and might here be an archaically used sign (Bréndum-
Nielsen, Danske runeindskrifter, Nordisk Kultur, vol. 6, p. 119, 1933).

15 Cf. Pedersen, Holger, in Runernes oprindelse, Aarb. Nord. Oldk. og Hist.,
1923, pp. 80-81; Olsen, Magnus, De norrgne runeinnskrifter, in Runerne
(runorna), Nordisk Kultur, vol. 6, pp. 102, 108, 157, 1933.

NO. 3 TWO RUNIC STONES—THALBITZER 13

If these unknown signs with which this inscription is started are not
mere ornament or fancy sample, might they not after all be hidden
numeral figures? One might feel tempted to adjust them so as to
agree with our decimal system and to be interpreted as 1314, but
probably this would be too rash an interpretation—particularly in
the face of Magnus Olsen’s ingenious hypothesis, which perhaps after
all is justifiable and which leads to a later decade. At any rate, it
would be remarkable if there were not a vestige of a preposed A.D.
(Anno Domini) or the word “year.” But what liberties cannot a
Greenlandic rune carver take when on a dangerous voyage of dis-
covery to the “Ultima Thule”?

II. THE KENSINGTON STONE

HisroricAL BACKGROUND
(The documents here mentioned will be found reproduced in the Appendix.)

Let us (with Holand) suppose that the rune carver was an expert
on runes and literature, of Swedish stock, a member of Powell Knuts-
son’s expedition starting from Sweden-Norway, which, according to
a document '® dated at Bergen, Monday the 3d of November, 1354,
was to leave Norway for Greenland by order of King Magnus Eriks-
son (sometimes nicknamed Smek). At that time King Magnus
still reigned over parts of Sweden and the whole of Norway; in 1355
his son Hakon took over the government of Norway.

Information on what happened on the expedition is very scarce.
It no doubt started out, although probably not until the spring of
1356. The document “Koning Magni Befalingsbref Powell Knuts-
son paa Anarm Gifvet, at Sejle til Grgnland,” has been printed from
a late copy among the diplomas in volume 3 of Grgnlands Historiske
Mindesmerker (pp. 120-123, 1845), as the original does not exist.
Anarm must be a mistake in copying for Onarheim. A distinguished
man, formerly an official in the service of Duchess Ingeborg, in 1347-
48 law-speaker of Gulathing, Powell Knutsson ‘* was appointed to be
“commandant” of the knorr (a type of ship) to take the men to
Greenland. He is commissioned to select among the King’s body-
guard the “housecarls or others” whose relations to the King are
such that he can command them to take part in the expedition. Thus
it is prominent Norwegian and Swedish men with their “retainers”
who are mentioned, fit to take part in an expedition to Greenland.
This is in keeping with King Magnus’ previous missionary and com-
mercial policy toward Russia, a policy that, however, had been rather
unsuccessful and had impoverished him and the realm. But in 1346
the knorr (the royal “brig” or a ship of similar size) had returned
with riches from the southernmost settlement of Greenland, the
Eastern Settlement, an extremely fortunate event, as the realm was

16 The document reads: “Mandagen efter Simonis oc Judae Dag” (the Mon-
day after —). As Simon’s and Judas’ Day (28th of October) in 1354 fell on
a Tuesday, the document thus originated from the 3d of November.

17 Munch, P. A., Det norske folks historie, Unionsperioden, first part, p. 420,
Kristiania, 1862. Here the name is spelled “Paal Knutss¢gn.”

14

————_—————

NO. 3 TWO RUNIC STONES—THALBITZER 15

in need of money. It was further known that the Skrelings (old
Icelandic name for the Eskimos) had attacked the Norsemen from
the north, as may be read in the Icelandic annals: “They are now
in possession of the Western Settlement.” Already in 1341 this settle-
ment had been found deserted. (Cf. what is said in a much later
document from Skalholt, that the people of Greenland in 1342 were
apostates from “the true faith and the religion of the Christians and
had turned to the people of America.”)*® This was the reason why
“the Christians began to abstain from the voyages to Greenland.”

In other words, there is no small probability that most of the
Norsemen of the Western Settlement after the first attack of the
Skreelings, about 1341, preferred to emigrate to the other side of the
sea, where they constantly had connections with Helluland, Mark-
land, and Vinland.

King Magnus, in short, had in mind a kind of small-scale crusade
to support the declining Christianity in Greenland, either to set it on
its feet again or to support the emigrants wherever they might be.

This strange expedition from Scandinavia to Greenland is not
frequently mentioned in the literature about the medieval voyages to
Greenland and Vinland. Little is known about it, and it is not known
to have been of any importance to the early history of Greenland.
Only upon arriving at Greenland could Powell Knutsson learn where
the apostates were to be found.1® However, no information on the
return of the explorers is extant. The expedition was not profitable

18 Sixteenth-century copy of an earlier document, printed in Grgnlands
Historiske Mindesmerker, vol. 3, pp. 459-460 (cf. p. 887), 1845. The year 1347,
given in the work mentioned above the passage on page 459, is a misprint; it
should be 1342. The passage in question is found in Bishop Gisle Oddson’s copy
made before the year 1637 from an old document in the archives of the bishopric.
(See the Latin rendering in the Appendix, p. 51.) Munch, P. A., Det norske
folks historie, Unionsperioden, first part, pp. 313-315, 1862.

19 Besides the above-mentioned pages in Grénlands Historiske Mindesmerker,
refer to Munch, P. A., Det norske folks historie, Unionsperioden, first part, pp.
633-634, 1862 (cf. pp. 413-415, 429). Storm, G., Studier over Vinlandsreiserne,
pp. 73-74, 1888. Holand, Hj. R., The Kensington stone: a study in pre-
Columbian American history, pp. 75, 78 ff., 1932, 1940; Westward from Vinland,
1940; America 1355-1364, 1946. Négrlund, P., Nordbobygderne, p. 128, 1034.
Hennig, R., Terrae Incognitae, vol. 3, pp. 268-281. Thalbitzer, W., Powell
Knutssons rejse, en forsvunden ferd til Grgnland og Markland, Grgnlandske
Selskab Aarsskrift, pp. 54-60, 1948.

On the Swedish King’s politics and lack of money see also Munch, P. A,,
op cit., pp. 599-603; Hildebrand, E., Sveriges Historia, vol. 2, Medeltiden, pp.
207, 214, 228-235, 250 ff., 1905; and @verland, O. A., Norges Historie, vol. 3,
pp. 814 and 816 ff., Kristiania, 1888.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

to the king. Powell Knutsson is not mentioned in any known docu-
ment after his departure in 1356.

Credit is due Hjalmar Holand for having pointed out the possi-
bility of a connection between this early Scandinavian Greenland
expedition and the runes of the Kensington stone. In this inscrip-
tion in a Scandinavian language it seems that a lost expedition has

OBS YOTER AAS FESRARRY 44:84:

8 otervok 2A NOT MER PA

2. ‘ABPXTHPHE PX RB: VRa:
opPpagelse Earp ya
3. VIthX tbe: Vtntevi:

weutarwp of wert wi

4. EXPEERKYERIV ED FSG ERR +t:
Rape ee ep Weep x skjiaren
5S PRSGERV GT SAR RP RA bth Ht 4:
pags NESLR WOAK Pra pena sten
YER AT: PENA td: PRY ERB TIER
we war ok Pidbhke en pagh AafRtinr
PEAVEY PR EPEYX FRG Pt:
We ROM haw hor 10 man VS pe
SK LM bd 4: bt bs AVM"
af blop og pep AVM
9. PRX FPGESXP: ING:
frietoe af illu (or illu)
10, AK REPKE PEARY PERT AE:
ae 10 mand we hawet at ase

MBP RPA REGAL BPE P ERI GE:

Actin wore shife sH pogh wees
OVC aak nee

am pengohkh ahr 14362

Fic. 3.—Transcription and translation of the Kensington inscription. (From
Holand, 1932, p. 5.)

erected a monument to itself somewhere inland about “14 days’ jour-
ney from the coast” nearly on the watershed between the sources of
the Mississippi and the prairie rivers going north or northwest.
Powell Knutsson and his men, then, are supposed to have sailed
first to Greenland on one or two ships at their disposal, from there
southwest toward Vinland, and, as the people searched for were not
found there, then “west,” i.e., according to Holand (as Vinland was
regarded as part of a large island), first north, back to the Hudson
Strait, and then west through this along the coast of the supposed

NO. 3 TWO RUNIC STONES—THALBITZER 17
large island (i.e., Labrador) ; from there either across Hudson Bay
or along the coast, until a favorable point at a broad estuary was
reached. Here then was a place where an armed group might go
inland in smaller boats, while the ship (or ships) was left behind
near the coast, guarded by a crew of Io men.

The inscription records a sanguinary catastrophe and the danger-
ous position of the men left behind “‘on this island”; but the lake in
which the island once was located later dried up; now there is only
a hill.

The men mentioned, 8 Goths 18 and 22 Norwegians, at the time
when the stone was erected, would have been away from their home-
lands in Scandinavia for six or seven years on a warlike expedition.

The inscription on the Kensington stone

. 8: géter :

As it probably is to be read
(with all its ‘errors’’)

ok : 22 : norrmen :

po:

As it ought to be, according to
the medieval (14th-century) Swedish
written language as it is known
to us

8 géter ok 22 northmen pa

2. : opdagelsefard : fra *updagelseferdum fra

3. winland : of : west : wi winlandi *of west. wi

4. hade : leger : wed : 2 skjar : hafSum legher web twem skizrum en
en:

5. dags : rise: norr : fra: beno: daghs *rese nor fra bessom steni.
sten :
: : dum :

6. wi: war: ok: fiske: en: dagh: wi warum ok fiska en dagh. eptir
eptir :

7. wi: kom: hem: fan: 10: man wi komum hem funnum Io men rgda
réde

8. af: blod : og : ded’: AVM: af blodi ok ddéSa. A.V.M.

0. freelse : af: illy: frelse af illu.

Io. har : 10: mans : we: hawet: (wi ?)** hawum Io men wep hafinu

dat cSe) :

at sea

II. zeptir : wore: skip: 14: dagh: zptir warum skipum 14 dagh *resor
rise :
12. fram : beno: gh: ahr : 1362: fra(m) bessi g. ar A.D. MCCCLXII

* Before unusual forms or words.
** or: tak (hawer?).

19a Goths = Ggtar in Modern Swedish will here be used for the inhabitants of
the Swedish provinces East- and West-Ggtaland and of the neighboring provinces
to the south.

20 Cf, Hildebrand, E., Svenska skriftprov (Andra haftet), 1900; Noreen, Ad.,
Altschwedisches Lesebuch, 1892-1894; Klemming, G. E., Svenska Medeltidens
Rimkr¢gnikor ; and Swedish homilies from the Middle Ages: Medeltids-Postillor.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Translation

8 Goths (Swedes) and 22 Norwegians on exploration journey from Vinland
westward. We had camp by two skerries one day’s journey north from this
stone. We were and fish(ed) one day. After we came home (we) found 10
(of our) men red with blood and dead. A.V.M. (Ave Virgo Maria) save (us)
from evil.

(We) have 10 men by the sea to look after our ship(s) 14 days’ journey
from this island. Year 1362.

The inscription is easy to read; the runes are large and distinct.
Most of them are unambiguous (cf. fig. 6). Ambiguous (uncertain)
are the runes:

4=0,4a = 9, oh f=Jor el (or le)
b=), éord Y=7,.0 (a)

We may believe that they reached the then unknown coasts of
Hudson Bay although it was indeed little short of a miracle (cf. Jens
Munk’s voyage, 1619-1620). The possibility of the same ships being
able to return home from there would be just as amazing. Probably
this first Scandinavian “group of emigrants” never returned, but
scattered and. perished. On this question Holand has his own
opinion.* The return is not mentioned in any extant documents.
Powell Knutsson and his men have disappeared in the mists of
antiquity. The expedition seems to have been entirely forgotten in
Scandinavia, like so many other things.

THE KENSINGTON RUNES AND THE NUMERAL SIGNS

One of the most surprising features that meets the eye in this in-
scription is the use of the old-fashioned runic numerals, “‘five-twig
signs” known from antiquated calendars (‘“‘runestave,” “primstave”’)
and other documents, discussed in Worm’s work dating from 1643 on
the runes and the “golden figures” (“gyldental’’) and in later works
written by Swedish experts.*? To find these numeral signs in a

21 Holand, Hj. R., The Kensington stone: a study in pre-Columbian Ameri-
can history, 1932 (2d ed., 1940). The author (with G. Storm) here seems to
believe in the return of the ship in 1363 or 1364 (pp. 88-95) although with a
great number of casualties, among them Powell Knutsson. Cf. Westward from
Vinland, pp. 146 ff., 1940; America 1355-1364, p. 14 (note), 1946. But in a
later chapter he follows up the fate of the group of the emigrated Norsemen in
America and their supposed merging in the peculiar culture of the blond Mandan
Indians, which appeared so remarkable to later explorers.

22Worm, Ole, Fasti Danici, Hafnia (Copenhagen), 1643. Martin P: son
Nilsson, N. Beckman, N. Lithberg, and N. Lid: Tidsregning, Tidsrakningen,
im Nordisk Kultur, vol. 21, 1934. Tuneld, J., in K. Humanistiska Vetenskaps-
samfundet i Lund, vol. 18. Brate, E., Sveriges runinskrifter, pp. 95-102, 1922.

NO. 3 TWO RUNIC STONES—THALBITZER 19

medieval runic inscription on stone is quite incredible and hitherto
unheard of in the history of runology!

The question arises: how old is this custom? The very first line
of the Kensington stone contains such “pentathic numeral signs,”
and the inscription ends by stating the year 1362 written with the
same kind of signs. On closer inspection, however, the numeral signs

=

rt
<

cai

PPPPLTT TELE ey
MEEPEE Te eerrereer re
PPP PEP EPPP PEL

errr errery
peers

Fic. 4—A page with calendar and “golden” figures, from Ole Worm’s Fasti
Danici (1643), Hafnia (= K¢benhavn).

of the stone seem somewhat different in form as well as in applica-
tion from the types given by Ole Worm. The Kensington rune carver,
e.g., wrote the number 14 with two digits, I and 4, as we do, instead
of with the single 4-twigged stave, as in Worm (see fig. 4), and the
signs for 22 and 1362 likewise follow the method of division of the
Arabic system. For to (?) the Kensington stone has a circular form
( ? ) instead of the cross-shaped sign in Worm.?* On the stone the

23 Holand identifies this sign, which looks like the Dalecarlian rune for the
letter O and the runic numeral for 109, with 10, probably through conjecture
(twice 5 written with runic numerals, nearly like two P’s close together: 9P ).
According to the Fasti Danici this sign (a circle on a stave) might correspond
to our 19.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

uppermost twig is always drawn from the top of the stave; in Ole
Worm, somewhat below the top. Holand shows some very similar
numeral signs in a vellum almanac from the fifteenth (?) century
found in the museum of Oslo (see fig. 5).

These numeral signs were already in use at the time in medieval
Scandinavia along with the Roman letters. The fact that the carver
of the Kensington inscription used the runic numerals of the calendars

Fic. 5.—Extract from a calendar. The pentathic figures (the calendar
gyllental, “golden numbers,” 19 in all) are seen on the lowermost line. The
Arabic figures in the lowermost line but one have been added later and refer
to the Latin letters on the middle line above them. These letters represent the
weekdays of the month of November, the week being reckoned as Abcdefg.
The small illustrations in the top section are connected with certain of the
weekdays, indicating seven festivals or calendar days of the month (from left
to right): All Saints, All Souls, Martinmas, The Virgin (Maria), St. Clemens,
St. Catherine, St. Andrews. (From Holand, 1932, p. 129.)

with twig digits in this way shows a detached modern attitude. It
occurred to him that they would be easier to carve on the stone. No
doubt he was even accustomed to writing them on vellum. But it is
true that they have not been found on any other runic stone in the
world. The case is unique—just as unusual as the placing of the date
at the end of the inscription. The man was obviously used to writing
both ordinary figures and dates according to the same system, i.e.,
Arabic numerals.

Anyone who has read P. A. Munch’s treatise on the Algorismus
(the Latin work in the Hauksbok), now a hundred years old, need
not wonder at the case. Writing with Arabic numerals in the four-
teenth century was a new art in the Scandinavian countries, but an
old one abroad. “After Hauk’s time, in the beginning of the four-
teenth century, the Arabic numerals are frequently found in Nor-
wegian and Icelandic codexes, partly as chapter numbers in the
margin, partly also as dates and sums.” ** Holand has mentioned this

24 Munch, P. A., Algorismus, eller Anviisning til at kjende og anvende de

sakaldte arabiske Tal, efter Erlendson’s Codex, Ann. Nord. Oldkyndighed, vol. 2,
Pp. 354, 1887-1888.

NO. 3 TWO RUNIC STONES—THALBITZER 2I

in his books and refers to works by O. Friesen, Bugge, and Gjessing.”®

The custom is so old that a rune carver about the middle of the
fourteenth century might very well be familiar with the use of these
numeral signs, the twig signs, then so modern. He must have been
a trained scribe and rune carver.

If we imagine a forger as author of the inscription, working far
west in Minnesota after the middle of the last century, but before
1895, where would he have obtained his knowledge of the runic
numerals? And if this incredible thing should be so, how foolish to
use them if he wanted to make an imitation of a runic stone! A forger
would at most have ventured to put Roman numerals on his runic
fabrication, and even then he must have been fairly skilled in the
late runic custom.”®

Indeed, much more probable is the explanation that the rune carver
was an Old-Swedish scribe or expert on writing who was accustomed
to using these signs, well known at the time, and who, perhaps be-
cause he was in a hurry, found it more practical to carve them in
stone. Indeed, he was on an expedition of discovery, had traveled
much, and for many years had been out of touch with conditions at
home.

More than half of the Kensington runes look correct and genuine,
as they agree with the 20 runes of the Codex runicus of the Scanian
Law. (MS. extant from about 1300.) The following nine deviate
more or less (see fig. 6) :

abdknt—u (=w) @6

25 Holand, Hj. R., 1932, pp. 127 and 256 (with illustration from Algorismus) ;
1940, pp. 180-182. For later works about this question, see here footnote 22,
p. 18.

26 Runic stones with Latin dates are known particularly from Gotland, the
earliest one, from Vallstena, dated 1326; a Latin inscription with black-letter
writing occurs on the edge of the stone. Ona chancel painting in St. Radakirke
in Varmland the year 1323 is rendered with Roman numerals (Brate, E.,
Sveriges runinskrifter, pp. 92-93, 1922). In the calendars and annals of the
time there are numerous examples of the use of Roman numerals. Otherwise
a circumscription was used, such as, “fourteen hundred years and one year less
than fem tigi (i.e., fifty),” as seen on two stones in the parish of Lye, Gotland.
But as shown by Holand there are in the documents of the Middle Ages, in
the Swedish diplomataria and in the Icelandic annals (Vetustissimi down to,
thus before, 1314, that of Skalholt down to 1362, and Rymbegla from about
1300) numerous examples of a knowledge of decimal notation. The Kensington
rune carver thus may have belonged to the Old Swedish period. On the other
hand, a later rune forger cannot have found the twig signs in Liljegren’s Run-
lara from 1832, where only the real runes are mentioned (p. 194 ff., cf. pp.
211-213).

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Selected specimens of characters (majuscules and

E : s
I Si a g g minuscules), chiefly from L. F. aaf, “Bokstafs-
£ H 5 G} ee former under medeltiden enligt Sverges offentliga
So aay =| af bon handlingar” (between 1164 and 1513) *
“= _ s
25 aU.s Bo eia0
oh 060 SOS SES
i n aS 1250-1300 1300-1500

WAS aoa

x
|
p
+
ie
q
|
f
‘
i
We
+
4
p
R
A
he
i‘
4
©

* Table in vol. 15 of K. Vitterhets-, Historie- och Antikvitets-Akademien Handlingar,
Stockholm, 1838.

Fic. 6.—Runes and old-fashioned characters.

NO. 3 TWO RUNIC STONES—THALBITZER 23

Further, the Kensington stone has three signs not found in the
Codex runicus, viz, the signs for p, j, and y (=ii). Among the above-
mentioned signs, however, the »- and ¢-runes are old forms, common
on most of the early runic stones, also a j-rune was known in the
older rune alphabet. But the others are rare or unique, e.g., B used
for the sound p. When B in the “fourth runic period’ was to be
used for P, the rune was provided with two dots (a “dotted” rune
B, at any rate on the Danish stones).*7 The rune carver here did
not use this. In a single place he used a b, viz, in blod (line 8).
The rune of the Kensington stone for that sound, b, looks com-
pletely wrong, but the rune may not have been finished (because of
the necessity of haste on the part of the rune carver?) and perhaps
should have been a complete B or b (cf. Appendix, p. 54).

The other peculiarities perhaps are worse, but we shall see pres-
ently that ) used for d and 0 in dagh, blod, Vinland, etc., is not sur-
prising (cf. the sword pommel from Ikast with the Latin word
bominum=dominum, and several similar instances).?* Indeed, the
inscription on the Kensington stone is from a period when new forms
were introduced by imitation of the Latin letters.*®

PALEOGRAPHIC SIGNS
(Cf. fig. 6)

The time of the runes in Scandinavia was ebbing away. The Latin
writing was imitated on stone. It began with the first dotted runes
in the eleventh century, which attained a certain similarity to black-
letter writing when a dot or a small upright twig was placed in the
u-rune, the inverted U, in order that it might be used to denote the
Scandinavian sound of y (ii), sometimes even two dots or points in-
side the U *° or a small horizontal pointlike twig on the J-stave, which
was to be turned into a sign for the vowel e (pp. 54, 58). The Latin
script came to central and southern Sweden via Norway, brought by
monks from England.*t They often used a manuscript y-sign provided

27 Jacobsen, Lis, and Moltke, E., Danmarks runeindskrifter, texts 947 and
966-967, 1942.

28 Brdndum-Nielsen, J., Danske runeindskrifter, Aarb. Nord. Oldkynd. og
Hist., pp. 205-206, 1917. Holand, 1932, pp. 238-239.

29 From the tenth century the Latin writing had forced the rune masters to
invent different signs for w and y, 7 and e, k and g, etc.

30 Jacobsen, Lis, and Moltke, E., op. cit., texts 980-981.

31 Friesen, O.v., Var Alsta handskrift pa fornsvenska, K. Humanistiska
Vetenskaps-samfundet i Lund, vol. 4, p. 3, 1904. “Vastergdtland got her Anglo-
Saxon script via Norway.” Christianity was preached in these regions by

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

with a dot above (4, insular point script). Later, from the fourteenth
century, the same feature appears in German texts as sign for an in-
verted wu (originally with a small e placed above the uw): ii. In the
same way the signs 6 and 6 arose in German script (originally with
a small e placed above an o).*”

The peculiar 6- and y-runes on the Kensington stone thus are easily
explained when interpreted as applied black-letter writing. They are
not runes but transferred (conventionalized) minuscules or majus-
cules. The rune carver has constructed them from the medieval mo-
nastic script. He may similarly in a hurry have constructed some
of the other unusual signs, because on his long travels covering so
many years he had forgotten some of the runes he was in need of.
Or the man may actually have known such signs from his monastic
school at home—more or less locally used runic signs resembling the
kind we find in the Dalecarlian runes of later times. But also the
Dalecarlian runes originate from the Middle Ages.**

The k-rune is here only an inverted k, i.e., conventionalized black-
letter writing in which the lateral twigs of the k are turned forward
instead of backward.

The j-rune (used in skjar, line 4), which is otherwise unknown,
is very much like a written black-letter i or j, stiff and conventional-
ized, really transformed like an inverted rune.

The v- or w-rune occurs in several places in the inscription; here
a consonantal v was needed, whereas the u-vowel accidentally and
strangely enough does not occur at all. The rune carver has resorted
to the expedient of using the Latin minuscule w (double wv or 1)
placed on a stave. In order not to mistake this sign for the m-rune,

Englishmen (cf. the list of the bishops of Vastergétland: “a large number of
these were English”). It thus appears in the fact that the earliest book script
in Swedish was with Anglo-Saxon types: y, @, p, and J, but Latin signs were
used for v, f, r. b supersedes the voiced fricative 0, which is only regarded as
a variant of d. “The fixed dot over the o-sign to denote 6 in the thirteenth
century is detached from the o-element (the oval), the beginning of our two-
dotted 6.”

82 Cf. Brgndum-Nielsen, J., Paleografi, Nordisk Kultur, vol. 28, A, pp. 14-15
and 23. “The two dots above uw in modern German ii are remnants of a dissolved
e.” ... “g in German words is designated by oe or 6 or 6.” Cf. examples in
Jansson, Sam, Svensk paleografi, p. 127: “vilia binder aff niio biiggia bi.”
Translation: (if) peasants will build a village of new.

83 Liljegren, J. T., Run-lara, p. 40 and pl. 1, 1832. Noreen, Ad., together
with Boéthius and Levander, Dalska runinskrifter fran nyare tid, I-III, Forn-
vannen, 1906 (see especially the table between pp. 90 and 91). Cf. Holand,
Hj. R., 1032, pp. 120-121, 123.

NO. 3 TWO RUNIC STONES—THALBITZER 25

he put a dot in the left pocket, the rune thus becoming a new
“dotted rune.” **

His y-rune with two dots over (in ily, line 9) is also easily recog-
nized from the y of the medieval manuscripts, but is nicely conven-
tionalized. This rune might easily be mistaken for a g-rune if it had
not been provided with an additional dot and also with a transverse
line through the stave as in the case of the j-rune.

At first glance the a-rune is no less singular. Its peculiarity, how-
ever, is chiefly due to the small hook on the upper right branch. As
in the Dalecarlian runes we here find that the original rune + gradu-
aly develops into a sloping position in the direction of an X. It might
be supposed that the carver of the Kensington inscription was a
Swede from somewhere in central or southern Sweden, who had
been accustomed to finding, perhaps even to using, a similar hook
in the written sign for a, a form that (as shown by Holand) is not
rare in ordinary black-letter writing in various Swedish documents
(cf. fig. 6).8° The rune carver probably transferred his own written
form to the runic sign. This type of an a-rune has never before been
found in any runic inscription and thus may be considered a fresh
late medieval ‘“‘runic sign’”—a discovery for runology!

The man also seems to have been accustomed to adding two dots
above his a-rune to denote an @; in the rune he contents himself with
adding a single dot before the hook, so that it looks almost like two
dots. This sign also is otherwise unknown, but it fits in with the fact
that in the Dalecarlian runes an d-sign is formed from the o-rune: 0.

These new runic signs (a, 6, ii) are distinctly and consistently
carried through in the Kensington inscription,** e.g., @ in lager, aptir,
perhaps also in frdelse. An d-sign, however, would seem to be super-
fluous in the last-mentioned word. As the dot (in the figure) even
seems to be missing in the rune in question, the @-sound, if so, is in
this word denoted in the Latin way by @ [from ae]: frelse (a verbal
optative form).*”

34 Holand (1932, 1946) regards the w-rune of the Kensington stone as a
further development of the corresponding letter in a document drawn up by
Bishop Nikulas of Oslo in the thirteenth century (Diplomatarium Norvegicum,
vol. 1, No. 7).

35 See, e.g., Hildebrand, E., Svenska skriftprof. Medeltiden No. 30 (specimen
from Viastgétalagen dating from 1280), No. 33 (Saint Birgitta’s notes from
about 1360), No. 34 (from 1387), Stockholm, 1894.

36 Holand, 1932, pp. 120-121 and 123.

37 If the word is to be read as frelse, @ is presumably intended to denote a
long vowel, with a reminiscence of the origin: frje-.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

O with an inside cross has rightly aroused surprise among the
runologists. Holand is of the opinion that the d-rune of the Kensing-
ton inscription in its shape combines the two (equivalent) signs for
6 and g. The latter (¢) is well known from the necrologies, calendars,
and similar works of the Middle Ages (cf. here fig. 6). Holand
refers to the Nestved Obituary, in which an initial majuscule has
been used, which is formed as an O with an oblique line through it
and provided with a pointlike hook above; and in a Norwegian letter
from 1321 (Diplomatarium Norvegicum, vol. 1) Ellingi or Ollingi
occurs, beginning with a sign, which very much resembles the rune
of the Kensington inscription with an inscribed small e-rune.*®

And both the latter and the name itself resemble the first name and
first rune on the Kingigtorssuaq stone from Upernavik! Indeed, the
first name (Ellingr) on the Greenlandic runic stone even begins with
a circular e-rune with two dots in it on each side of a vertical streak
(chip: 12).

Thus the ii-sign (minuscule of y) is the first to have loose dots on
top; but the d-sign had them at nearly as early a date.

The Kensington rune master, who was active between the years
1350 and 1362, was familiar with dots placed above the letters or
within them from Swedish medieval black-letter writing (fig. 6).

THE DALECARLIAN RUNES AND THE H-RUNE

One of the critics of the Kensington inscription (G. T. Flom)
thought he could prove imitation of the modern “Dalecarlian runes,”
thus branding the inscription as a forgery. Some of the signs just
mentioned (a, 6) and the runes for d, f, h, i, and m, then, were
simply to be explained from a modern knowledge of the late Dale-
carlian runes.*® But the rest of the runes of the inscription deviate
from the Dalecarlian ones! (Cf. fig. 6 and pp. 21, 24-25.)

88 Holand, 1932, pp. 112-118, and 1940, pp. 166-170. Also he refers to Axel
Kock, Svensk Ljudhistoria, vol. 2, pp. 1-2, 1909: “When rendering OSw.
manuscripts in print the editors generally have not considered the different
forms of the g-sign, but have printed either 6 or g.” Kock mentions, e.g., in
one of the earliest Swedish prints from 1495, “o with a dot above it.” The
Kensington inscription, however, was dated more than a century before, and
we still lack other examples of a distinctly 2-dotted 6 dating from its period
(the fourteenth century). Holand, however, has found a single instance (1932,
p. 117): “In Onarheims Gildeskraa (MS. of 1397) the g has a single dot (6),
but the w has a double dot (aw) ... In Den Norske Landslov (MS. of 1325)
the ¢ usually is written with one dot: 6, but once it is written with two dots:
6 (Paleografisk Atlas, ed. Kaalund, No. 22a, p. 11, 1903).”

39 Noreen, Ad., Dalska runinskrifter . . ., 1906.

NO. 3 TWO RUNIC STONES—THALBITZER 27

Most of the Kensington runes agree with the runic forms in the
Scanian Law or those on the runic stones from the “Fourth Period.”
The rune carver obviously followed an old tradition of writing inde-
pendent of both the Dalecarlian runes and the Scanian Law. Every-
thing suggests that he was a contemporary of some latinized runic
system—in a monastic school—about the middle of the fourteenth
century. At any rate, it is neither the early nor the late Dalecarlian
runes he used for his work. Nor is it the dialect of Dalarne he used.
This, too, has been pointed out by Holand.*°

The h-rune was used initially in hem, hawet, har, hade, in final posi-
tion in dagh in order to emphasize that the g was aspirated,** and
after the vowels in 6h and ahr to indicate vowel length. This last
use, i. as a sign of long vowel, is a most unusual feature, almost with-
out parallel in Swedish or Danish spelling in the Middle Ages, where
it was a custom to denote vowel length by doubling (aa, yy, etc.) or
by putting a small stroke above the letter.*

In contrast to this genuine h-rune, which in final position may be
used as a kind of diacritical mark (on which more below, see pp. 44-45),
the other signs here discussed, those denoting k, 7, u (w), y, are not
at all runes proper but black-letter minuscules or majuscules as these
were written in the fourteenth century, conventionalized for the pur-
pose of being carved on stone. For adaptation to the hard stone they
had to be changed a little and provided with diacritical marks.

The rune carver may have known these special forms. If so, it
would be valuable if somebody could find out in which Swedish
province these singular runic forms belonged, which might give a
clue as to whence this rune carver came and which “dialect” he spoke.
Or the man may have formed them on the spur of the moment, from
the image in his memory of the usual signs of the vellum
manuscripts.**

40 Holand, 1932, p. 124.

41 Wimmer, L. F., De danske Runemindesmerker, manual ed. by Lis Jacob-
sen, 1914, p. 28: “The h-rune: ... Only in very late inscriptions A is further
used to denote g: dah = dag.” In the Kensington inscription there is in a way
a double denotation (gh) of the spirant g. This fact may have given rise to the
use of h to denote vowel length (e.g., -a from -agh).

42 Holand, 1932, p. 254, may, however, refer to such a spelling as ahr efter
gudz fdédelse in Klemming’s edition of Svenska Medeltidens Rimkrgniker.

43 Cf. fig. 6, adduced from Raaf, L. F., Bokstafsformer under Medeltiden
enligt Sverges offentliga Handlingar (specimens of writing from Latin and
Swedish documents, between 1164 and 1513). Table to vol. 15 of K. Vitterhets-,
Historie- och Antikvitets-Akademien Handlingar, Stockholm, 1838.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

So far the script of the stone need not create any doubts.**

More peculiar and suspicious are (1) the 2-dotted 6 and (2) h
used after a vowel to denote length of the vowel. Both of these
features of the inscription look like anachronisms. Otherwise all
the signs, the runic and the paleographic ones, as well as the three
Latin majuscules, AVM line 8, are easily explained and warrantable
as seen in the light of the authentic year of the stone, 1362.

The runologists of our day have shown the occurrence of so many
new, in part unique, runic variants from the late Middle Ages, on
church walls, vellum manuscripts, and numerous other objects, that
the peculiarities of the Kensington inscription, apart from the excep-
tions mentioned, need no longer alarm us. The boldness of the vari-
ants should rather, if anything, inspire us with confidence in the
genuineness of the inscription. A modern forger with the knowledge
of runes and the Old Swedish language to which the inscription testi-
fies would not have been so foolish as to introduce the “anachronisms”
mentioned. In other words, we may accept the anomalies as evidence
of a medieval style of writing not hitherto met with. The rune carver
was a scribe or clerk with a character of his own.

Tue Lincuistic Form

The style looks strange, and here and there nearly modern—quite
incredible, say the critics, for the time to which the inscription pre-
tends to belong. The language does not look quite like any known
Scandinavian form of speech from the fourteenth century; it re-
minds one a little of Danish, but more of Swedish. But what about
such forms as wi har, vi hade, vi var, kom, fan (without the ending
-um for the first person plural) ? Quite improbable to the sensitive
ears of a philologist accustomed to the medieval scholar. Unheard
of, unseen in the old vellum manuscripts from the fourteenth cen-
tury! And further, what of such expressions as pa opdagelsefard, vi
var ok fiske, 10 man, 10 mans ve havet, 14 dagrise fram peno gh?
These and other things sound absolutely spurious, say the doubters.
No small effort is needed for a Scandinavian philologist, who knows
the medieval language from old diplomas and laws, to tackle these

44 Holand (1932, p. 108) may be right in his view that the new formation
may be due to a failing memory. The men of the expedition would hardly have
carried any “rune book” or list of the current runes with them. The rune car-
ver had to rely on his memory, after being away from home and having traveled
through foreign parts. He could not, perhaps, remember any correct runes for
k, w (uw), @, 9, y (2), and hence resorted to the Latin letters.

NO. 3 TWO RUNIC STONES—THALBITZER 29

monstrous linguistic morsels and test their contents. I must confess
that I have myself had some qualms.

But the inscription is long and undoubtedly also contains some old-
fashioned forms, indeed some obviously quite archaic forms (of west,
aptir, fram, beno, etc.). Consequently it must be examined. Analyze,
isolate, test, consider the individual words in the following investiga-
tion, and then sum up the whole!

After all, what is linguistic style? Something fixed and immovable?
What remarkable speech do we not hear every day, in lectures, on
the radio, in the street, in children’s speech and mixed language. The
written language was never the highest norm for the living language.
Those earlier times, indeed, also had both class language and living
dialects, of which the texts that have come down to us—on runic
stones or vellum—have left us extremely sparse evidence, which per-
mits only a fragmentary knowledge. Undoubtedly we do not know
the spoken language of the time.

And here we have to do with quite abnormal conditions. This
American runic inscription is more homeless and rootless than any
other inscription known to us—more than the Kingigtorssuaq in-
scription from North Greenland. On the other hand, it is dated, and
is Swedishlike in its linguistic form. Its contents tell us that the
rune carver must have been surrounded by Norwegian and Swedish
men, and their speech seems to have been mixed with both foreign
and Old Swedish words and word forms.

The language of the inscription is a kind of simplified basic Old
Swedish engrafted with new tendencies; and, as on so many other
runic stones, there are plenty of grammatical and orthographical
errors and inconsistencies. But, as we shall see, the inscription still
testifies to an astonishing knowledge of Swedish pronunciation in
the fourteenth century.

OLp SweEpIsH

A great many of the word forms of the inscription decidedly agree
with old-fashioned Swedish. Old Swedish (OSw.) meets the eye in
the very first line.

Line 1. In Modern Swedish (ModSw.) the plural of the word
denoting an inhabitant of the Swedish provinces of Ostergotland and
Vastergétland is Gétar; but the Goter (-@r) of the inscription is typi-
cally OSw. as in the manuscripts from the fourteenth century (in

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

the Swedish laws, diplomas, etc.) : ok e gleddus vestgoter af honum
meden hans lifdagher waru.**

Line 1. po or pa for early pd, with the rune 1=0 or dad. Com-
mon in OSw.*®

Line 2. op, with the vowel o (close) is OSw., but ModSw. up,
upp.?

Line 4. veb corresponding to Old Norse (ON.) wip, vid; the
original 7 became e before the consonant ) from about 1300.** Thus
it is a typical OSw. form for ModSw. wid.

Line 5. fro or fra peno sten, cf. line 12, frdm peno oh. Here
fram is an archaic form of what later became ModSw. fran. Origi-
nally (in Gothic) the form was fram, from which such intermediate
forms as fran and fra are frequently found in OSw. and also in
stressed position with the form fram some time after (e.g., in
St. Birgitta’s Uppenbarelser, MS. from about 1360, and Lydikinus,
Vestgotalagen, MS. from 1300).*° Fran from fram gradually carried
the day; but in an OSw. dialect the m may have been preserved for
a long time and a transition fram>frdm is possible: the Sw. a (long)
before a labial consonant was apt to develop into a or o in the four-
teenth century, and the transition, according to Axel Kock,®® was
completed about 1400. Hence ModSw. fran.

The Kensington inscription has twice fra and once *frém, OSw.
forms. Whatever may be the reason for this vacillation, the occur-

45 From a letter of about 1325 written by a clergyman in Vastgotland (see
Noreen, Ad., Altschwedisches Lesebuch, 2d ed., p. 143, 1904); @ denotes an
e-sound; hence we find Gdter or Géter. On @ > e, cf. Rydquist, Svenska
sprakets lagar, vol. 4, pp. 16-19, 93-96; Kock, Axel, Fornsvensk Ljudlara, vol.
I, pp. 116, 121, and 144, 1882. In Magnus Eriksson’s Konungsbalkr of about 1350
(Noreen, loc. cit., pp. 32-33) we find a plural form kununger (ModSw.
konungar).

46 Kock, Axel, Svensk Ljudhistoria, vol. 1, § 430 (431), p. 353, and vol. 2,
§ 801, p. 201, 1906-1911, “with labial consonants: o <a,” e.g., suo, swo < swa;
hwo (hoo) < hwia; two < twa.

47 Kock, Axel, Svensk Ljudhistoria, vol. 2, § 801, pp. 200-201, 1906-11, u>o in
OSw. in compounds, e.g., opbera (op with close o and comparatively unstressed
as in opd@). On fro see line 5.

48 Kock, Axel, ibid., vol. 1, §§ 30 and 47, pp. 27 and 41.

#9 Cf. some more examples in Holand, 1932, pp. 257-258, which in part are
due to information from Professor Séderwall.

50 Kock, Axel, Svensk Ljudhistoria, vol. 1, § 364, p. 299: “about the year
1400 the transition to a real d-sound was completed (early OSw. ga > late OSw.
ga, mostly written gaa or ga)”; cf. about a > 6, d, pp. 348-354, especially § 431:
“from about 1400 we find examples of o instead of a (a), such as wor “var,”
gordh (< gardh < garb) “gard,” fron “fran” ... in diplomas from Ostergot-
land from about 1400.”

NO. 3 TWO RUNIC STONES—THALBITZER 31

rence of these Swedish variants on the stone, even though fram has
not hitherto been found in other inscriptions or manuscripts, is
strong evidence that the language of the Kensington inscription as a
whole belongs in the fourteenth century.

Lines 5 and 12. eno is an OSw. dative corresponding to ModSw.
denna. The form pheno (also benom) is one that occurs extremely
rarely in OSw.,** and hence it is remarkable that it is found twice in
this inscription. The accusative masculine stem thenna, penna is
somewhat more frequent. But generally the stem Passe is used in
OSw. both in the singular and the plural, and this probably also
applies to Old Norwegian and Danish. Still the stem of Penna is
old and is found also on old runic stones (accusative Panna, dative
panni) alternating with the more frequent forms passi, p@ssi
(paimsi, beimsi). Peno in the Kensington inscription in both cases
covers a dative, apparently in the neuter. A stone (sten) was
masculine, an island (dh), feminine; but the demonstrative pronoun
at the time, in Holand’s opinion, had probably fossilized in a certain
insusceptibility to gender, so that we may find 7 thesso helgho script
(neuter dative in o although script is feminine) .°*

Sten and 6(h) in the ancient language would have had diphthongs
(stain, ey), but in many East Swedish provinces the vowels were
monophthongized as early as the thirteenth century, just as the earlier
(=ON.) ein became een.®? The use of h as a “‘diacritical” mark for
long vowel, as stated above, is rare in medieval writings, though not
entirely unknown.** (See, further, pp. 44-45.)

Lines 6 and 8. og=ok. Kock points out that the transition of
k to g had already begun in the fourteenth century; for instance,
jak=jag, taka=tagha, etc., in alternating use often by the same
writers. ok and og are OSw. forms; NSw. ock or och.*°

Lines 6 and 11. dptir is only found in OSw. In OSw. runic lan-
guage generally eftir (or aftir, uftir) ; ModSw. efter.

51 Sdderwall, A., Ordbok 6fver svenska spraket i medeltiden, Lund, 1925, sub
verbo pdnne (thenne, thetne). He mentions thenno as the dative neuter oc-
curring “a single time”; Noreen, Ad., Altschwedische Grammatik, 1904, men-
tions the form in § 500, note 8, as rarely occurring in writing, e.g., in Diploma-
tarium Swecanum (ed. Hildebrand, 1878) from the years 1348, 1405, etc.

52 Holand, 1932, pp. 242-243, but ON. and OSw. script might also be the neuter,
so the example is not worth much; more examples on p. 253.

53 Kock, Axel, Svensk Ljudhistoria, vol. 2, § 481.

54 Jacobsen, Lis, and Moltke, E., Danmarks runeindskrifter, text, p. 957, note
4, 1942.

55 Kock, Axel, Studier 6fver Fornsvensk Ljudlara, vol. 1, pp. 35-36, 1882-86;
Holand, 1942, pp. 290-291.

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Line 10. hawet (not hawit) is OSw. (cf. p. 38).

Line 11. skip with distinct 7 in OSw. in contrast to ModSw. skepp.
Kock has shown that the originally short 7 in certain forms developed
into e in OSw.** He writes: “The forms with i (gifva, kid, skipa)
and the forms with e (geffua, ked, skepa) originally belonged to
different dialectal shades, and a preceding palatal consonant thus has
only in certain districts been able to preserve 7. The above, pp. 454-
456, quoted skeneben, skepp, etc., like skepelse, skepnad, with e after
palatal, originally belonged to a region where this was not the case.”
To which Swedish province the rune carver belonged may perhaps
be suggested by means of such criteria as those mentioned here.

wore, “our,’ with close o in OSw. (as in swo)*?; ModSw. with
open d.

Line 12. ahr, “year,” with long a is an OSw. form; ModSw. ar.*®

ARCHAISMS

Most of the particularly remarkable words and word forms also
bear an old-fashioned stamp.

Line 1, norr(men), line 5, norr (adverb). The former of these
words in ON. is generally spelled nordmen (plural), but the form
here is spelled according to the pronunciation in OSw., with assimi-
lated rd>rr as in Norwegian dialects; cf. Aasen, Ivar, Norsk Ord-
bok, 1873, s.v. Nordmann, “inhabitant of Norway: in this meaning
always pronounced norrmann, without any dialectal difference, Old
Norw. nordmadr.” Cf. norrgn (particularly about the wind) <
nordrgn. As an adverb the word for “north” is otherwise generally
spelled nor (Aasen).°® Cf. Rydquist, Svenska sprakets lagar (1850-
1883), vol. 2, p. 451, where we find nor in OSw. But why not write
norr as we often spell off for of ? This is the principle followed by the
rune carver. He wrote norr phonetically according to the Norwegian
and Swedish pronunciation of the fourteenth century.

Line 3. of west. If the inscription had been Old Icelandic or Old
Norwegian, there would be nothing surprising in this expression.

56 Kock, Axel, ibid., pp. 459-460; cf. p. 456.

57 Kock, Axel, Svensk Ljudhistoria, vol. 1, §§ 430-431, pp. 353-354.

58 Kock, Axel, 1882-86, vol. 2, p. 407, and 1906, vol. 1. § 420, p. 352. On the
spelling with h, see here pp. 44-45.

59 Cf. ModSw. adverb ner neder, “comparative form,” Rydquist, vol. 5, p. 125:
“a few stems ... originally: hvar (ubi), ber (par), her, nor, etc.”

6° Fritzner, Johan, Ordbog, vol. 2, p. 867, 1886, s.v. of “over,” when referring
to movement over something which is thus passed: sudr of fjall (southwards
over the fell) ; of lopt ok wm log (through air and over sea). Holand (1940,

p. 204) renders it by “(from Vinland) round about the West.” Cf. in Ynglin-
gatal: of austr, “eastwards.”

rr —

NO. 3 TWO RUNIC STONES—THALBITZER 33

It would then presumably have to be translated “westward” (from
Vinland), and this is probably the meaning of it in this place, even
though the language is Swedish and we do not (unfortunately)
happen to find the same old expression in the extant literature from
the fourteenth century. No doubt it might very well be OSw., but
it is hardly common usage in ModSw. in any part of the country.
For that matter it might also, in accordance with usage in the old
language, be identical with what would have been written as ofuest,
offuest, “very far west,” an adverbial phrase that is hardly ever found
in OSw. literature either but might very well have been preserved in
some dialect through the fourteenth century—in that case found re-
corded only on the Kensington stone in Swedish. This is one of the
points of conjectural criticism of the inscription. But at any rate the
phrase is old-fashioned.

Line 4. skjar (plural) is also a form belonging to the Middle Ages,
as the word otherwise both in ON. and ModSw. is sker, skair. We
have here presumably a late “broken form” (by analogy) as it sounded
in a Swedish dialect, corresponding to contemporary words from the
fourteenth century such as siax (gra sak), “six,” sial (German Seele)
in Swedish manuscripts; ® in the Jutlandish Law skial (wm marke
skial), “boundary.” ®? The tongue or ear of the rune carver led him
to render the unusual broken form by the rare j-rune instead of 1,
perhaps in order to indicate a “hard” pronunciation of the preceding
k: s-kj-ar (j pronounced as English y). Was the pronunciation
foreign to his ear? (Was he a foreigner ?)

Lines 1, 7, and 10. men (e.g., in norrmen, line 1) is the regular
plural of man.®* The forms show that in OSw. it was possible to use
the word collectively in the singular as it still is in Danish: alle mand
pa plads! to mand frem! ti (stykker) til mands. Besides we find in
OSw., preserved from the early language, combination of a numeral
with the following noun in the genitive plural, e.g., “thirty men’:
XXX manna (partitive genitive, manna is plural), a mixing well
known in all history of language (parataxis, contamination) ; a mixed
construction as still used in Icelandic: 200 manns, annan hundrad
manns (manns is genitive singular).°* A corresponding way of ex-

61 Noreen, Ad., Altschwedisches Lesebuch, 203, 3125, 3325. In ODan. sial,
siael (Bertelsen, H., Dansk Sproghistorisk Lzsebog, 1905).

62 Kaalund, Kr., Palzeografisk Atlas, No. 23, 1903.

83 See Noreen, Ad., Altschwedisches Lesebuch, 1894, Glossary, s.v. maper,
porsi.

64 Holand (1932, p. 260 ff.) has collected a good number of instances of this
use of the genitive singular (mans in a collective function) from Snorre Stur-

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

pression is not met with in the OSw. literature. Some day, however,
we may by chance find in some old document that it was once possible
to use such a construction in the spoken language, at any rate here and
there in Sweden ; then the ro mans of the Kensington inscription (line
10) will establish another example: meaning “a crew of Io,” “ a Io-
man party.” The strange genitive in line 10 thus is to be interpreted
as a collective and elliptical one. The collective use of a man(s) is no
more peculiar in OSw. than in Modern Danish.

Line g. illy, “evil.” This somewhat strange form in -ly of *illa
(adjective) should be a dative governed by the preposition af,
“from’’; but it sounds strange and wrong, and the y may be due to
an error by carving an extra dot in the otherwise unknown, but quite
correct, u-rune used here. If we read without the dot, illu, this would
be a normal dative of *7la in OSw.

ily or illu can have been intended only as an adjective in the neuter
(dative), “the evil,’ thus an abstract and nominal notion, but the
word is not found as such in the large Swedish dictionaries, where
only the adverb illa, “in an evil way,” occurs. The same applies to
the large Danish dictionary, but differently in Norwegian and Old
Norse (Icelandic). Fritzner, in his ON. dictionary, 1886 (West
Norse), quotes ilr as an adjective, “evil” (about a person or thing) :
sja vid illo, “beware of what is evil” (Sigrdrifa 37) ; gjalt bu eigi illo
alt (Gammel Norsk Homiliubok) ; laustu (=leys pu) oss fré illo,
“deliver us from evil’ (ibid.), the last example thus the very form
from the Lord’s Prayer as on the stone. But the first printed large
Swedish translation of the Bible from 1541 had formed the prayer
as it is now spoken everywhere in Scandinavia: frels oss ifraé ondo
(Matthew 6:9).

Here we have thus again an instance of the Kensington inscrip-
tion showing us a phrase kept in medieval Swedish popular speech
but not otherwise recorded in the extant literature (like of west, etc.).
In Norwegian, too, we find illa recorded not only as an adverb but
also as the neuter and as an adjective (see Heggstad and Torp, Gam-
melnorsk Ordbog, 1930: illr (ilt), eg., hin illi, “the evil one, the
devil” ; alt er illum at vera, “it is evil to be evil’). Still more impres-
sive in this connection is the old Norwegian rhyme (fragment) known

lason’s Konungasogur, and from the Flateyarbok: fj¢lda mannz. An old usage
analogous to mans is here folks: bar war talid 300 folkz eda meir (Oddveira
Annall sub 1341 in Storm, G., Islandske Annaler, p. 480, 1888) ; but more fre-
quently with genitive plural: letuzs XXX manna, “30 men perished,” ibid., p.
403 (the years 1346-1349). By a similar development we have in Danish the
word slags (< slag), “kind, sort”: ti slags, mange slags.

NO. 3 TWO RUNIC STONES—THALBITZER 35

to Holand through the Norwegian peasants in Minnesota, in which
illi ends a line in an adjectival function.®

In the Lord’s Prayer in Swedish, in the form from about 1300, it
says: frelse os af illu. “God save us from evil.”

The phrase of the Kensington inscription thus is correct OSw.

NEOLOGISMS

The following words are still more surprising, as they look too
modern for the language of the fourteenth century as known from
medieval literature:

Line 2. opdagelse (ferd)—not the ending -else, which makes its
appearance and spreads as a productive suffix as early as the four-
teenth century,®’ but the word opdage. Who in Scandinavia ever
used this word before the rune carver of Kensington? It is not found
in recorded language of the Middle Ages. A word like opdagelse in
OSw. looks like an utterly strange word, as if it here turns up long
before its proper lease of life. Still, opdage is undoubtedly, as sug-
gested by Holand, a word of genuine Scandinavian origin,®* but
spreading to both East Frisian as uppdagen and Dutch as opdagen,
“come to light, appear,” transitively “bring to light, reveal, or dis-
cover.” It has not been found recorded in early Norwegian, Swed-
ish, or Danish. From West Norse the word landalcita is known with
the same meaning, but it became obsolete at an early date. When
was it replaced by opdage? Holand’s reference to German influence
on the Swedish vocabulary as early as about 1350 is particularly
instructive.

An extensive immigration from Germany to Sweden took place
about 1350 (or earlier). The immigrants obtained rights and privi-

65 Holand, 1932, pp. 268-270; 1942, pp. 308-300.

66 Holand, 1932, p. 268, quoting Ahlquist, The history of the Swedish Bible,
in Scandinavian Studies, vol. 9, p. 93.

67 According to Ad. Noreen’s Altschwedisches Lesebuch, 1892-94, there are
many words ending in -else in OSw. literature, 1356-1400, such words as
liknelse, dppilsi, wighilsi (p. 28), styrkilse, kynnilse (p. 43), styrilse (p. 166),
the first-mentioned of them at any rate 50 years before 1400. Correspondingly in
Old Danish; cf. Bertelsen, H., Dansk Sproghistorisk Lzsebog, vol. 1, 1905:
tilliggelse, “surroundings” (p. 109), bestandelse, aminnelse, begengelse, etc.

68 Holand (1932, p. 229) calls attention to dagar uppi in Alvismal (verse 35,
ed. Finnur Jonsson, 1888) and Grettissaga, p. 152, with reference to Fritzner’s
explanation in Ordbog (s.v. daga). Cf. Falk and Torp’s definition, Etymol.
Worterbuch, vol. 1, s.v. dag, p. 134, and Ordbog 6fver Svenska Spraket, vol. 1,
p. 97, and vol. 2, p. 33, s.v. dag. In Sw. dialect uppedaga, ON. uppidagadr,
about a troll who has been surprised by daylight.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

leges on an equal footing with the country’s own citizens.®® Falk and
Torp are of the opinion that the word mentioned above was influ-
enced by German entdecken, properly “uncover” (a rendering of
French découvrir), which is also reflected in Sw. uppticka, Early
Danish optckke, “unveil, bring to light”; cf. German aufdecken, “re-
veal, lay bare.” The German influence has left its traces in a great
many Swedish words, both in prefixes and endings of an exactly
corresponding kind as in opdagelse, to be found in Sw. manuscripts
as early as the fourteenth century, indeed earlier, e.g., in Vestgota-
lagen from 1285; opstandelse, opfarelse, optagelse, in an increasing
number through the influence of “Den Heliga Birgitta’s Uppen-
barelse,’ which dates from 1360.

The word opdagelse has not otherwise been recorded from OSw.,
but among the above-mentioned words from the literature of the
fourteenth century it would not look remarkable, and it may be quite
by accident that it has not been recorded. It can already have been
current in living speech at that time.

As to the vocalic quality of the initial o (in op-) (see p. 30), the
stress was once minimal on the first syllable of this composite, but
strong on the second one: upddgelse, opddgelse; now 6pdagelse.

A composite like opdagelsefard (“voyage of discovery’) is highly
surprising for the language of the fourteenth century. But we find
early examples of composites in ON., e.g., heimskringla, konungs-
skuggsja, sonatorrek-, and a tendency to even longer compositions in
medieval Swedish and Danish: herabhgfhingi, heresshgfping, em-
betzman, umboszman, hofoplot, midhlungaman, tamlangadagher,
loghordagh, kirkiustefna, scriftemol, tienasta-quinna, pianostoman,
gudzs¢ffwalaghspield, wor herres fgdelses-aar.”°

Lines 5 and 11. The same (a seeming anachronism) applies to
rise, which, of course, is a rendering of German reise (Old High
German reisa, Middle High German reise) and is resa in ModSw.
It is surprising to find this “modern” word so early in the Swedish
language, but the context was created on a long voyage, an expedi-
tion of young and old warriors and clerks traveling together, who-
ever they were, presumably each of them originating from his own
cultural center in Scandinavia. There is every reason to suppose that
the runic scribe was a Swede, but it is possible that one of the partici-

69 Munch, P. A., Det norske folks historie, Unionsperioden, first part, vol. 1,
pp. 596-598, 1862.

70 Noreen, A., Altschwedische Grammatik, 1904; Kock, Axel, Svensk Ljudhis-
toria, vol. 1, 1906.

NO. 3 TWO RUNIC STONES—THALBITZER a7

pants in the expedition was a foreigner (German, Frisian, Dutch, or
English). A form like rise with a long pure 7 does not agree very
well with OSw., the loan from a foreign language perhaps is just
reflected in the pure 7-rune (| ), not (t ), which has been maintained
both times when the word appears in the inscription and is possibly
meant to render a diphthong (e).™* And, be it noted, this word,
which was new and rare at the time, has been used only as a technical
term, viz, in the compound dag-rise “‘day’s journey,” a term that may
have been first used here to replace the maritime unit of the long
voyage, the common measure of the ship’s speed. At sea they prob-
ably used dggr or a corresponding circumlocution in Swedish with
which the distance at sea was measured in early times (before the
invention of the nautical mile). The distances between Iceland and
Norway or Greenland are stated in dggr, “24 hours.” “Day’s jour-
ney” (that is, night and day) was a new fixed unit of length for use
on land, perhaps used the very first time on this expedition. It ap-
pears that the real distance from the seacoast approximately agrees
with the measure of the distance as stated on the stone, when this is
interpreted as “14 day’s journeys” (1 day’s journey=75 nautical
miles, in all 1,050 miles),’* which thus means that the searching
travelers were at least 14 days on the road from the coast of Lake
Superior or the mouth of the Nelson River (?), but, in fact, most
probably a greater number of days. May we rely upon their having
counted the days exactly? They may, of course, have carried a
wooden calendar along with them for the control of time.

Line 8. og pep (dep), “dead.” As the rune carver has found no

’

71 The root vowel in this OSw. form rise, “voyage, journey,’ must have been
long: imitation of the vowel of Middle High German reise, “departure,” “war-
like expedition.” Its transition into e in late Swedish is analogous to the transi-
tion in steg < stigh (Icelandic stigr) ; cf. ModSw. led, sed, fred (beside frid).
See Kock, Axel, Fornsvensk Ljudlara, pp. 457-458, 1882. Conditions are differ-
ent in OSw. risi (rese), “giant,’ with short i (Kock, loc. cit., p. 260).

72JTn connection with the voyages to Vinland, dggr and similar conventional
terms denoting distance have been treated in detail in the literature about those
voyages. Holand has collected the results with reference to the present case
(referring to works by F. Nansen, W. Hovgaard, Fossum, Gathorne-Hardy,
etc.; cf. Holand, The Kensington Stone, 2d ed., pp. 135-139, 1940). In the old
directions for setting a course toward Greenland I find degra sigling, degra
haf, “a night-and-day’s sailing” (Grénlands Historiske Mindesmzrker, vol. 3,
p. 210 ff.), Fritzner’s dictionary has the expression dagrédr, “as far as one
can row in a day,” mikill d., “a long day’s rowing.” On land, rest: Thor’s
hammer lay hidden “eight rosts” down in the interior of the earth. Other terms
were also used: dags fdr, viku fdr, “week’s journey.” See Petersen, N. M.,
Haandbog i den Gammelnordiske Geografi, pp. 132-135, 1834.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

difficulty in writing 6 in other words (lines 1, 7, 12), there must
be a special reason for his spelling this word with an e. Perhaps his
own dialect has just made this spelling natural to him. Thus dggr
and degr alternates in the Flateyjarbok (about 1389).7% Both A.
Noreen and Axel Kock adduce many examples of corresponding
conditions (delabialization) ; g>e is a frequent transition in early
Swedish. Kock adds: “in certain tracts,” eg., svefn for sugfn
(“sleep”), bredr for brgdr (“brothers”), fetr for fotr (“feet”), bret
for brgt (“bread”), hera for hgra (“hear”), ofert for ofgri (“a
mess’’) ; 74 in the Erik’s Chronicle inversely (labialization) gptir for
eptir (“after”), ggnom for genom (“through”), thom for them
(“them”). This shows that this alternation of e and g was fairly
common.”® Rydquist also gives a great many examples: kemr for kgmr
(“comes’’), red for rgd (“red”), ex for gx (yx, “ox’’), etc.7® Holand
has found the same spelling as in the Kensington inscription, ded, in
a Sw. letter dating from 1390: effther the henne hosbonde her Jens
Herne ded er (“as her husband Mr. Jens Herne is dead’’).7

Line 10. (we) havet (“at the sea’) is surprising because it has
the definite article -et instead of -it, and particularly because the noun
is uninflected after we(d) where we should have expected a dative
with the ending -u."* The endings -in, -it, -ir had become -en, -et, -er
in the manuscripts of the fifteenth century.’® Axel Kock has shown
that the Bjorkoratten uses final e after @ in the penultimate (this
occurs frequently also in Vastgotalagen: Jate, present subjunctive of
the verb “to let”: gielde, gielde, present subjunctive “pay”; origi-
nally gialde; ware, gange; bape fgter, “both feet”; and that the
scribe was probably a West Goth. The old laws, as Vastgétalagen,
Bjorkoratten, were drawn up for their respective provinces, and it
is natural to assume that their copyists originated there. The West
Goth must have written under the influence of the dialect change
taking place in Vastergotland.*°

73 Cf. Holand, 1932, pp. 265-266.

74 Kock, Svensk Ljudhistoria, vol. 1, p. 127, and vol. 2, pp. 38-42 (§ 623).

7 Cf. Holand, 1932, pp. 266-267.

76 Rydquist, Svenska sprakets lagar, vol. 4, pp. 98-101.

77 Holand, 1932, p. 267: “in a letter of 1890 from Lédelse, West Gothland”
in Diplomatarium Norvegicum, vol. 4. No. 586.

78 The examples adduced by Holand (1932, p. 251) from about 1350-1400
(j rédha hafuit, Svenska Medeltids-Postillar, Kleming’s ed., pt. 1, p. 7, and in ¢
riket, Konungsbalker, Noreen, Altschwedisches Lesebuch, p. 53) are irrelevant
as they are accusative forms used after verbs of movement.

79 Kock, Axel, Fornsvensk Ljudlara, p. 146, 1882 (from Rydquist).

80 Kock, loc. cit., pp. 145-146 and 156-159.

NO. 3 TWO RUNIC STONES—THALBITZER 35

The omission of final } (in wep, “with,” line ro, cf. line 4, on the
stone) is not unknown but is not frequent in OSw.*! On the vowel e
for ModSw. 7 see page 30, under line 4.

THE STYLE

We have still to consider the total impression of the style. The
runes show the way: only half of them were genuine runes; the rest
were the Middle Ages in disguise, monastic or school script trans-
ferred to stone.

The Kensington inscription resembles no medieval text handed
down, whether on stone or on vellum. As compared with other runic
inscriptions it is more communicative than most and offers special
topical details about something unusual. It is peculiarly personal and
emotional, epic-dramatic in contents.

The rune carver feels himself to be surrounded by unknown
enemies and now wants to leave a memorial behind. He was under
the necessity of using runes on a stone. He must have known some
local individual runic system. But the inscription might just as well
have been written on vellum or on a whitewashed wall, and it is
without any mysticism.

The language is about half OSw. dialect, unraveled word by word;
but as far as we can see it teems with anomalies and grammatical
errors and resembles a medley of archaic and modern forms and,
furthermore, contains an alien element. The style, perhaps, as sug-
gested by Holand, may be interpreted as an approach to the “col-
loquial” language of the fourteenth century. But in my opinion this
holds good only in part; rather this abridged style developed on the
long journey on which many Goths and Norwegians from various
cultural centers were together. Their different dialects and schooling
may have given rise to discussion about “language.” A mutual influ-
ence took place, more or less consciously. The result was a mixture
of a somewhat artificial language and a careless one. Here one can
picture great change and looseness in the words of the inscription.
They are, so to speak, full of foreboding unrest. They originate
from a very unusual situation, which perhaps has only one analogy
in the history of the runic stones—that of the three men from
Kingigtorssuaq north of Upernavik in Greenland.*?

81 Holand, 1932, sect. 58, p. 250, refers to Noreen, Altschwedische Grammatik,
§ 308, 1904, and Vastgétalagen (1285), ed. 1827 by Collin-Schlyter, p. vii.

82 Nearest in situation and style probably comes the Hénen stone of Ringerike
(see Neckel, Die Erste Entdeckung Amerikas, pp. 83-84). We may think also
of Jérgen Brgnlund’s death diary, the lines of which are carved on the Mylius-
Erichsen stone near the Langelinje promenade in Copenhagen.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

On the island near Kensington there must have been a most seri-
ous periculum in mora for the 10 men of the crew who were left be-
hind since the large party of men had ventured on the dangerous
exploration inland. Hence the inscription was made in a hurry, per-
haps in fear and horror, but was copious enough to serve as an
explanation to the men left behind “by the sea’ or whoever else
might search for those missing and remaining.

The rune carver was a learned clerk who was an expert on runes
rather than a warrior. And he must have belonged to the younger
generation among the travelers. He belonged to a party of brisk and
bold adventurers who for six years had been traveling by sea and
land and had gotten far away from the “school” habits of their
homes. They must have been accustomed to hearing one another’s
dialects. The rune carver seems to have disregarded petty academic
rules of spelling. His language seems to have been disintegrating.
Perhaps the words were dictated to him, pithily and hurriedly. We
do not know the origin of his linguistic usage and habits of writing;
they may testify to the instruction of a teacher with a character of
his own, or to discussions taking place in this small traveling com-
munity. Perhaps some individual dialectical habits come through in
his very personal and epically formed style. Hence it has a strange
and novel effect. The secret perhaps may be found through a study
of the style including the dialect.**

To me the colloquial language of the rune carver strikes through
from first to last, not least in the “incantation” in lines 8-9: freelse
(verb in the optative, “save (us)”). Here he reveals his connection
with the church. Of course, in a catastrophe like this, where the routed
remaining members of the expedition were in danger of their lives,
one must invoke the Virgo Maria.**

The young man spoke a different dialect from that of the man who
dictated to him. He did not speak like the older men. Their aftir
varum skipum became in his speech dptir vore skip; réda became

83 Lis Jacobsen in Nye Runeforskninger, p. 5, says: “But further no secret
should be made of the fact that so great real and methodic advances have taken
place within philological and historical research that we are able to understand
the language of the past in a way, it seems to me, essentially different from
what was formerly the case. We have learned to interpret the documents with
an intimate, concrete solidarity such as is extremely rarely found among pre-
vious investigators. This living contact with the sources characterizes the efforts
of our time within the study of antiquity.” Cf. also Beksted, A., Vore yngste
runeindskrifter, Danske Studier, vol. 36, 1939, particularly pp. 111-114.

84 Holand, 1932, on invocation of the Virgin Mary, pp. 247-248, and 1940,
pp. 268-270, 301-309. Cf. pp. 35 and 46.

NOW 3 TWO RUNIC STONES—THALBITZER AI

réde; dopa became dep; the latter form looks like a neuter (singu-
lar), perhaps induced under the influence of the preceding man (in-
stead of men in plural), which means the part of the “crew” that was
dead (dep, cf. pp. 37-38).

He heard the German reise, and perhaps during the travel had
gotten into the habit of pronouncing it with a long e or 1: rese or rise
(rijse). To his ear the foreign diphthong ei, MLG. ei, may have
seemed a monophthong. As a foreign word he left it uninflected in
the inscription.

The short forms (wi) hade, har, war, kom, and fan jar on the
philologist’s ear as deviating from the typical literary style, but they
may very well be explained from the possible (viz, exceptionally oc-
curring) forms in the early language of the middle of the fourteenth
century.®° All this may be considered from the same view as that
taken above—what Holand terms colloquial usage, the forms of the
natural conversational speech of the younger generation, which would
have come through more or less completely in this inscription.*®
After all, they might perhaps sound like this (see pp. 28-29) in the
speech of the younger generation about the middle of the century,
particularly by a somewhat singular individual who was somehow quite
detached from the school language, or was influenced by a dialect
other than his own. It has become a vulgar language in sharp con-
trast to the conservative forms of the literature of school and church.

The fellowship of the old Scandinavians on their travels through
those many years must have had not only a leveling effect on their
speech but must also positively have evoked old-fashioned words from

the Norwegian and Swedish dialects of the Middle Ages. This in part

85] shall not here repeat all the examples of such OSw. and ODan. verbal
forms (first person plural) as those above, mentioned by Holand (1932, pp. 233-
236) and already previously adduced by better-known linguists. The linguists
quoted by Holand on that occasion are Falk and Torp, V. Dahlerup, H. Bertel-
sen, Bréndum-Nielsen, and G. Indrebg ; and among works containing such forms
are Lucidarius (from about 1350); Svensk Diplomatarium; Diplomatarium
Norwegicum; Queen Margrete’s letter 1393 (Molbech and Petersen, Diplomer
og Breve, No. 22); Klemming’s edition of Medeltidens Rimkrgniker; Codex
Bureanus, etc.: zi gaa til bordh, the svenske kom forst; wi gawe, wi unne, wi
bide, we henghe, etc. in the language of about 1350-1400. G. Indreb¢ gives these
examples from the fourteenth-century language: gefuir wer (we give), setti mer,
setti mer (we set, trans.), hefuer ber (you have), ete.

86 Cf. Brdéndum-Nielsen, Gammeldansk Grammatik, vol. 1, p. 24: “The fact
well known in the present day, that there are many degrees of spoken language,
colloquial speech in town and country, language of lectures, of the theater and
church, etc., can be traced far back in time.” Cf. Trygve Knudsen in the Finnur
Jonsson Festskrift, p. 448; Seip, Norsk sprakhistorie, pp. 340-343.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

explains the occurrence of, respectively, the modern and the archaic
forms in this remote inscription.

Could there have been a conscious tendency in this vagary of
speech among the travelers—a linguistic smartness or jargon? Or
was it rather a topical common speech ground down by the daily
frictions of the dialects—a consciously intended “short language,”
a purist “lingua franca’?

It might seem appropriate to call to mind Otto Jespersen’s brilliant
studies on neologisms, mixed languages, hybridism, and pidgin Eng-
lish—all realities. Read in his Language (1922) the chapters about
linguistic play and word creation, about isolated children’s (twin’s)
construction of quite new languages, about the history of language,
sometimes by way of sudden alterations; and about Horatio Hale’s
theory of linguistic development.**

Near Kensington in Minnesota we have, on the other hand,
found a specimen of language apparently from the Middle Ages and
originating from a small isolated Scandinavian community of grown
men roaming beyond Greenland to Vinland, a mixed group with
different dialects and new views. Result: a cocktail of medieval
Scandinavian dialects!

I surmise that in the dramatic hours of the carving of the inscrip-
tion the dictator (perhaps a Norwegian) represented the conserva-
tive tendency (illu, of west, etc.), whereas the scribe or carver (per~
haps a Swede, a Goth from southern Sweden) represented the modern
tone. It was he who in the urgency of the moment kept to the short-
est possible forms, aiming at a kind of old-fashioned telegraphese,
left out a word like “we” and once or twice a verb, dotted the vowels
or forgot the dots. The result was a kind of runic shorthand with
abbreviations, etc.—up-to-the-minute in the 1360’s among these first
emigrants !

FuRTHER PHILOLOGICAL IMPRESSIONS

This matter can and ought to be investigated deeper than has
hitherto been done. The present paper is only to be compared to the
impression after the first turning of the spade. Under the roots of
the tree the stone lay hidden with its philological riddles. It has not
yet been completely brought to light, even though the light falls more
clearly on certain points than it did 50 years ago when the stone was

87 Cf. Otto Jespersen’s Language, p. 181 ff. and 213 f., London, 1922, and vari-
ous Danish works: Nutidssprog hos bgrn og voxne (1916), Bérnesprog (1923),
and Sproget (1941), with more Danish details.

NO. 3 TWO RUNIC STONES—THALBITZER 43

found by the Swedish farmer. Certainly a find with undreamt-of
and far-reaching consequences, if the runes are genuine!

Axel Kock with Lyngby (versus Rydquist) emphasized in his
books that about the middle of the fourteenth century “greater” (i.e.,
increasing) linguistic differences had developed, which resulted in
the Middle Swedish dialects we know from the time of the provincial
laws. Thus the dialects already flourished before 1350, “really differ-
ent dialects,” so that there is a connection between the language of
the old provincial laws and the language of Skane, the Old Skanish
dialect, which again had Skanish-Danish character. But Kock did
not make a secret of the fact that our knowledge of these problems
was still (in 1886) very defective:

Most of our provincial dialects are still too imperfectly investigated, and
among the early monuments which are most likely to give information of the
early form of our dialects, the runic inscriptions, the place-names, and the dip-
lomas are for the most part either unreliably or incompletely published or not
published at all.88 . . . But in this connection it should further be noted on the
one hand that, as remarked by Lyngby, many a minor difference might be found
in pronunciation which the comparatively imperfect writing was unable to ren-
der, and on the other hand that the dialects in more remote districts might differ
materially from the language in the provinces for which we have provincial laws
left.89 . . . And also when a dialectal peculiarity in the provincial laws does
not occur in the present dialect of the province, this is no sure proof that this
dialectal peculiarity did not exist in the province five or six hundred years
ago. This appears clearly from the vowel harmony of the Vestgota writings.°°

It is evident that Kock formed a picture of a number of OSw. dia-
lects, now in part disappeared, a far more variegated picture than
that which the scanty literary samples of those times now reveal to us.

Dialect forms may be found even in runic inscriptions: Funenish
in Master Jakob Réd’s rédlyt for rédlyt, “reddish”; Jutlandish in
gore for gorde, “made,” mek for mik, “me,” on the chest from
Pjedsted (Vejle) ; in the church of Lgnborg (Ringk¢bing) skol for
skule; the Kragelund stone (Viborg, between 1225-1250) bad,
“bade,” rist,’ “carved.” ** Peculiarly truncated forms are found on
Claudius Claussén Swart’s map of the North from the beginning of
the fourteenth century:

haw’ for hauer (present indicative).
driw’ for driver (“the sand drifts from the north’).

88 Kock, Axel, Studier 6fver Fornsvensk Ljudlara, vols. 1 and 2, 1882-1886,
the section “Till fragan om fornsy. rikssprak och fornsvenska dialekter,” p. 490.

89 Thid., pp. 494-495.

90 Thid., p. 500.

91 Jacobsen, Lis, and Moltke, E., Danmarks runeindskrifter, No. 217 (1275-
1300, the censer from Stenstrup, Svendborg), Nos. 221, 222, 231.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 116

If the curly mark after haw and driw in the legends of the old
map are intended to render a dialectal pronunciation (Claudius Clavus
was a native of Funen), these forms after all are just as advanced
as har, war, kom, etc., in the Kensington inscription. But the ques-
tion is, what is this sign (’) intended to denote? Maybe it is only
a shorthand trick to indicate shortening of the full forms haver and
driwer and, if so, is without any importance to our problem.

Mixed language is known also from runic stones, e.g., the Karlevi
stone (Oland) from about the year 1000, with Danish and Norwegian
used indiscriminately (according to Wimmer).®? Strange or wrong
spellings are not rare, such as ura:suti for gresund ** on a stone from
the twelfth century; hir for her (“here”), on a Greenlandic runic
stone (Igaliko) ;°* ston for stin or stein (“stone”), on a Swedish
ones"

Errors in carving were frequent; but even if we keep to the forms
in the Kensington inscription as they look according to the value of
the runes, they may perhaps have been used and heard in a Swedish
Gotaland province about 1300-1350.

We do not, however, know any dialect that gives vi fan for vi
funnom (or funno).®®> Nor do we know the two-dotted vowels G, 6,
even though current script letters may be indicated by the unusual
runic forms of the inscription X®“4. And what is one to think about
h in oh (“island”) and ahr (“year”)? Vowel length denoted by h
was exceedingly rare about that time in Scandinavia. See, however,
an example in Bréndum-Nielsen: gooh pronounced with a long o.°
And Ivar Baardssgn’s description of Greenland from about 1350
(MS. copy from the sixteenth century)? begins as follows:

Saa sigger vis[s] mend som fgd[dle ehre udj Grénland; and farther down:
daa stoer strém ehr (“where strong current is”), with eh = é@.

92 Thid., No. 245; cf. Nos. 263-266.

93 Dresund, The Sound between Skane and Sjelland, ibid., No. 47.

94 See Bruun, Dan., The Icelandic colonization of Greenland, Meddelelser om
Grgnland, vol. 57, p. 121, 1918: Vigdis M’s datter hwilir hir.

94a Dybeck, R., Sverikes runurkunder, vol. 1, p. 37, No. 255, Uppland, 1860.

®5 If “we” covers an “J” the first person singular form becomes more in-
telligible, being analogous to wan (uan) from winna. May the rune carver have
written “we” where the leader (dictator) said “I”?

86 Brgndum-Nielsen, Gammeldansk Grammatik, vol. 1, § 24, p. 59: “h after
vowel o to denote length (rarely): gooh.” As to the dots over the vowels (4,
6, ti), see here also p. 24.

87 Grgnlands Historiske Mindesmerker, vol. 3, p. 248; Meddelelser om
Grgnland, vol. 20, pp. 278 ff. and 322, 1899. Finnur Jonsson, Det gamle
Grgnlands beskrivelse af Ivar BarSarson, 1930.

NO. 3 TWO RUNIC STONES—THALBITZER 45

Is there possibly a foreign influence in this case?

Is there not a possibility that this spelling habit was connected with
the German influence in Central and West Sweden (mentioned above,
pp. 35-36), which even before the middle of the fourteenth century be-
gan to appear in the Swedish language? Might not these spellings with
ah, eh, oh be some early predecessors of the spelling of the German
official (Kanzlei-) style, which gradually gave rise to ah, eh, etc., for
the earlier dG, é, etc., as in modern German Stahl, Mahl, Jahr, Ehre,
Sohn, etc. ? %8

The rune carver must have been an expert on writing, familiar
with medieval OSw. dialect and spelling, comparatively expert on
runes considering the times, but completely unscrupulous as regards
the runology and orthography of the time. Again, is there perhaps
a foreign influence? Was he perhaps more or less a stranger who in
this Nordic language mistook the gender in peno 6h; mistook a for a
in two words, fard for fiard, skjar for skidr; left out endings in order
to make the carving easier; and even skimped his work in his hurry?

A forger living about 1895 able to collect in his product all the
rare traits here mentioned must indeed have been a most cunning and
sophisticated scholar and something of a genius. He would have to
have known Ole Worm’s rare work on the runes with the “golden
figures” and P. A. Munch’s Algorismus; also Liljegren’s and
Dybeck’s works on the runes from the 1830’s and 1860’s, and Axel
Kock’s Studier (1886) and Undersokningar (1887). But what about
several later works (after 1900) on Swedish dialects, runes, and
paleography? Over there in Minnesota! Was he perhaps in collusion
with learning in Sweden? Did he as late as 1895 live at a seat of
learning, near a Swedish library? Who was he, this great X ? Learned
and ahead of his time he must have been—if he ever existed.

A large number (one-third) of the words of the inscription un-
doubtedly are genuine Old Swedish (see p. 29 ff.). The same may
also apply to some forms that look like old-fashioned expressions
(aptir used as a conjunction, of west; illu; beno, etc., pp. 32-35). The
form peno is of particularly rare occurrence; but is there not a pos-
sibility that it was much commoner in OSw. than suggested by our
old manuscripts? After all, it was the victorious form (denna)! [It
is a puzzling fact that this very form occurs repeatedly in the South

98 Behagel, Otto, Geschichte der deutschen Sprache, in Paul’s Grundriss der
Germ. Philologie, vol. 1, p. 676 (cf. p. 671, sect. 17), 1901 (“Schon um
BAGO es « )s

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Greenlandic runic inscriptions of the fourteenth century (cf. Ap-
pendix, p. 53) ].

Finally, it should be kept in mind that some (one-sixth) of the
words of the inscription are neutral: they would in OSw. be written
and pronounced nearly as they are now and therefore look like the
language of our day: ok, wi, Vinland, lager, en dags, nor(r), hem,
af, at, frelse (friilse). They count as genuine OSw. of the fourteenth
century.

The prayer (invocation) corresponds to the genuine form from
the fourteenth century (see pp. 34-35). An Ave Maria is known also
from the late Swedish runic inscriptions. The AVM of the Ken-
sington stone presumably means Ave Virgo Maria. It occurs here
as a fixed formula of invocation, a charm: “Hail, Virgin Mary, save
(us) from evil.” °° The author of the Kensington inscription is prob-
ably here thinking of ila as “misfortune” rather than sin.” The
form illy perhaps may be retained as an imitation of an early Nor-
wegian dialectal form corresponding to ili (dative singular).

The occurrence of this old West Norse word in such a dative form
as lly perhaps more than any of the other peculiarities has brought
the inscription into discredit.°° A dot too much may have been carved
in the rune, in which case the rune is an error for the u-rune (a script
rune of an otherwise unknown form, imitation of the monastic
script), so that the reading is ilu. But after Holand’s account of
this word in ON. and early Norwegian I have little hesitation in
accepting the view that this word seemed quite natural to the Nor-
wegians but perhaps sounded a little strange to the Swedish Goth—
to say nothing of the evidence of the old version of the Bible as re-
gards the use of that identical word. These are weighty arguments.’

The invocation of the Virgin Mary on the Kensington stone is an

99 Holand, 1932, pp. 247-248; cf. above, p. 40, footnote 84. On Christian
formulas on runic stones, etc., see particularly Beksted, A., Runerne, pp. 98-
112, 1943; Jacobsen, Lis, and Molkte, E., Danmarks runeindskrifter, 1942, with
texts, pp. 841-842 (the Ave Maria formula) ; see particularly the Sdborg stone
(vol. 1, p. 308); Friesen, O. von, Runorna, De svenska runinskrifterna, p.
232i ., 1633:

100 The assertion that ily looks like an English word is, of course, sheer non-
sense. It is true that the form in -ly is unknown from Old Norwegian or Old
Swedish literature; but y [i] may be an error for u or a misinterpretation of
a final 7 of a possible illi which may have been dictated to the rune carver.
This latter explanation has not occurred to any other interpreter. But a stroke
of luck brought the advocate of the stone on the right track. See Holand, 1932,
p. 269; 1940, p. 308 (cf. above, pp. 34-35 and 40).

101 Holand’s account (see above) bears the character of being a scientific
investigation and has been very instructive to me. Doubt and criticism are
found in many passages, in Holand always held in a genuinely constructive

NO. 3 TWO RUNIC STONES—THALBITZER 47

impressive argument in favor of the authenticity of the inscription.
No modern forger could in the last century know how deeply this
prayer as here formulated is rooted in the medieval Christian ritual
language. With reference to the sources just quoted (p. 46, ftnt. 99),
I can further note the Lord’s Prayer as recorded in recent times on
Shetland Islands, a survival of the Old Norse (or Norn) of the
islands :

Fai vor o er i chimert “Father our, who art in Himmerig”
Halaght vara nam det “Holy be name thine”

La conungdum din cumma “Let kingdom thine come”

But delivra wus fro adiu idlu “But deliver us from all ill.” 1°?

In none of the other lines does the author of the inscription show
his relation to the powers that be, either God or a leader. No name
is mentioned in this inscription. The man feels only that he is a mem-
ber of the party, one of the survivors. Only the two ethnic names
have been preserved, and in OSw. form: Goter and Norrmen.

The word “we” is otherwise his only distinctive mark, the wi

_ DE Ee ee ee ee ae
spirit, as he has confidence in the presuppositions of the find; but in few others
dominated by a negative tendency, e.g., in Erik Noreen’s Svensk stilparodi
(1944), which practically tries only to ridicule Holand. The same holds good
of Prof. R. B. Anderson’s “Another View of the Kensington Runestone” (Wis-
consin Mag. Hist., vol. 3, 1920) ; here it is a serious misunderstanding to refer
to Fryxell as a runologist, as he never wrote anything about runes. Skeptical
views are also advanced by Milo M. Quaife, “The Myth of the Kensington
Rune Stone” (New England Quarterly, December 1934) ; Halldor Hermanson,
The problem of Wineland, 1936; Wolfgang Krause, Runen in Amerika, Ger-
manien, 1937; A. W. Brogger, Vinlandsferdene, p. 153, 1937; etc.

Among others the following scholars have written with sympathy, more or
less positively in favor of the genuineness of the stone: Profs. Hj. Lindroth,
K. F. Séderwall, P. P. Iverslie, G. Indreb6 (cf. Holand, 1932, pp. 230, 287,
103) ; and in America, O. E. Hagen (South Dakota), Francis Betten (Mar-
quette University), F. S. Cawley (Harvard), Stefan Einarsson (Johns Hop-
kins), A. Fossum, author of The Norse Discovery of America (1918), A. Strom-
berg (University of Minnesota), G. Woodbine (Yale University); also R.
Hennig (see above, pp. 4, 14), and Philip Ainsworth Means (Newport Tower,
1942). Cf. Holand, 1942, pp. 318-334.

A brief but noteworthy controversy took place in Denmark between Prof.
Aug. Krogh (the Nobel Prize winner), who in Flensborg Avis, November
1941 and May 26, 1942, maintained the authenticity of the Kensington inscrip-
tion, and the philologist Dr. Gudmund Schiitte, who for linguistic reasons re-
jected it (ibid. March 14, 1942). Krogh was strongly supported by Vilhelm
Marstrand, the historian and civil engineer ; but none of these showed any inti-
mate contact with linguistic method. Ten years earlier W. Thalbitzer had in a
Danish newspaper (Berlingske Tidende, January 21, 1933) called attention to
the problem with reference to W. Hovgaard’s review of Holand’s book on the
Kensington stone, just published.

102 Cited from Brdégger, A. W., Gamle Emigranter, p. 28.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

(we)*° which implies a solidarity for years between these first emi-
grants from the Scandinavian countries to Greenland and Vinland,
during their wanderings through the huge unknown island country
(as they presumed) west of Vinland. This wi is omitted in the
first line, appears in the third line, and is repeated twice; line 6,
wi war ok fiske, the last word being a careless pronunciation form as
in a hurried report,’°* and line 7, wi kom hem. In line 10, wi is
omitted. The rune carver is nervous and disregards the strict de-
mands of grammar for correspondence (congruity), is doubtful of his
spelling, omits both words and endings. The man lets himself go,
unceremoniously, to give the information needed.

So long an inscription—i12 lines—could hardly have been carved
in less than two days (or by two men in one day). Night may have
intervened in the performance and caused special disturbances farther
down in the text.

The general impression is contrary to the style of the literary
sources on record, in which no parallel is known. But is it not possi-
ble that all this lack of form is to be psychologically explained from
the seven years of travel, the 30 Scandinavians’ confusion of tongues,
the situation as it appears from the contents of the inscription, and
other conditions unknown to us or that may only be guessed at: a
scribe with a character of his own, perhaps a conscious “purism,”
perhaps a foreign element in the party, dictation at the carving and
the dictator’s or the carver’s individual reactions?

The cautious philologist will find that the signification of the Ken-
sington stone has not yet been established. It must be decided whether
it has not been condemned too quickly and whether there are not new
facts available forcing us to consider the find from a fresh angle, per-
haps to begin to come to terms with prejudices of any kind and grad-
ually simply try to take cognizance of the contents. In the develop-
ment of runology and philology in the time since the stone was found
a few years before the end of the last century, so many new facts and
views have appeared that it now seems possible to maintain that this
peculiar inscription—the runes as well as the contents—in spite of
everything may be genuine.

But, false or genuine, a solution is wanted, if possible a proof.

103 a with long i [77] in OSw. as the 7 in illu; see Kock, Fornsvensk Ljudlara,
vol. 1, p. 420; Seip, D. A., Norsk sprakhistorie til omkring 1370, pp. 321-322.

104 Perhaps a paratactic anacoluthia of wi war ok fiskade (verb) and wi war
at fiski (sb.). Many analogous types of sentences are discussed in Otto Jesper-
sen, Tanker og studier (1932), the article “Sammenfaldet og—at,” e.g., behag
at tryk pa dgren; versgo og sid ned! se nu og bliv ferdig! Sw. ga ut a jaga
(p. 186) han holl pa att skratta= han héll pa och skrattade (p. 188) ; de hafde
weret henne och fiske 1 aaen (p. 203).

APPENDIX (1950)

THREE HISTORIC DOCUMENTS
(Cf. above, p. 15 ff.)

I

Translation of the document dated at Bergen 1354 illustrated in
facsimile in figure 7. From the Royal Library of Copenhagen, Old
Collection, 4°, No. 2432.7

King Magni letter of command given to Powell Knutsson at Anarm [prob-
ably Onarheim] to sail to Greenland.

Magnus, by the Grace of God, King of Norway, Sweden, and Skone, sends
to all men who see or hear this letter good health and happiness in God.

We desire to make known to you that you are to take all the men who shall
go in the knorr 2 whether they be named or not named, from my bodyguard or
other men’s attendants or of other men whom you may induce to go with you,
and that Powell Knutsson, who is to be commandant on the knorr, shall have
full authority to name the men whom he thinks are best, both as officers and men.
We ask that you accept this our command with a right good will for the cause,
as we do it for the honor of God and for the sake of our soul and our predeces-
sors, who have introduced Christianity in Greenland and maintained it to this
day, and we will not let it perish [nederfalle| in our days. Let it be known
that whoever breaks this our command shall feel our displeasure and pay us in
full for the offense.

Executed in Bergen on the Monday after Simoni and Judae Day [i.e., Oct. 28]
in the 36th year of our rule [i.e., 1354] Herr Ormer Ostinsson, our Lord High
Constable, set the seal.

King Magnus had come up against the great crisis of his life. “In
1350” (say the annals) “King Magnus and Queen Blanche arrived
at Bergen and then he gave Sweden to his son Erik, but Norway to
Hakon, with solemn promise, and set them on the royal thrones and
ordained a bodyguard for them; but he reserved for himself Haloga-
land, Iceland, the Faroes, and Hjaltland.” Nothing is said about
Greenland, but presumably this time as usual it followed Iceland. A
great decision had been made, and the plan was to be realized within
5 years. It meant the separation of the realm, the dissolution of the
Swedish-Norwegian union.

1 First time printed and commented on in Grgnlands Historiske Mindesmerker,
vol. 3, pp. 120-123, K¢gbenhayn, 1845.

2 knorr originally meant a special type of ship, but in the Icelandic-Norwegian
sources of the fourteenth century it is the king’s ship in the possession of the
crown of Norway, which was regularly used for the sailing between Bergen
and Greenland (cf. p. 14).

49

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

But there was a hope, a rehabilitation in a plan for a crusade to
the remote Greenland. Out there was the golden straw at which the
king was clutching. In four years the plan ripened, and the royal order
was issued, as we have seen.

II

In the Historia Norwegiae, written about 1250 or earlier, we hear
for the first time about skirmishes with the Skrelings in the Nor-
wegian settlements on the west coast of Greenland.? A hundred years
later the Skrzelings had closed in on the northern settlers and already
seem to have expelled the northernmost of them. The story of this
has been told by the Norwegian clergyman Ivar Bardsson (Bdrdarson,
Latinized [varus Barderi, sometimes abbreviated Bertt, Bere), who
(born in Greenland himself) from 1341 for more than 25 years acted
as deputy for the bishop at the Cathedral of Gardar in Greenland
during a long interregnum over there. His words have been rendered
and translated in a short work, a Latin description of Greenland from
the fourteenth century, beginning as follows: “So say wise men who
were born in Greenland,” etc., a script that has often been commented
on.* We shall here quote only a passage toward the end:

Likewise all this, as said before, was told us by Iffver Bardsen Greenlander,
who was principal in the bishop’s residence in Gardar in Greenland for many
years, that he had seen all this, and that he was one of those who was appointed
by the law-speaker to go to the Western Settlement against the Skrellings to
drive them out of the Western Settlement, and when they arrived there, they
found no man, neither Christian nor heathen, nothing but some wild cattle and
sheep, and they fed on the wild cattle, and took as much as the ships could
carry and then returned home [i.e., to the Eastern Settlement] and the men-
tioned Iffver was with them.

Ivar Bardsson thus found the Western Settlement deserted; only
some stray cattle. This was presumably in the summer of 1342.

III

The above-mentioned would be in agreement with another Icelandic
report, which refers to an event in the Western Settlement in the
same year; but it is only extant in a much later copy from 1637 (in

2 See Storm, Monumenta historica Norwegicae, Christiania, 1888; Thalbitzer,
Historical data about the East Eskimo, Meddelelser om Gr¢gnland, vol. 31, p. 26,
1904.

8 Grgnlands Historiske Mindesmeerker, vol. 3, pp. 248-264 (especially p. 259),
886-889; and Finnur Jonsson, Det gamle Grgnlands beskrivelse af Ivar BarSar-
son (Ivar Bardsson), ed efter handskrifterne, Kgbenhavn, 1930.

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which the words ad Americe populos may be a later translation of
an original ad Vinlandie populos) :

1342. Groenlandiz incole a vera fide et religione Christiana sponte sua
defecerunt et repudiatis omnibus honestis moribus et veris virtutibus ad Americe
populos se converterunt; existimant enim quidam Groenlandiam adeo vicinam
esse occidentalibus orbis regionibus. Ac inde factum quod Christiani a Groen-
landicis navigationibus abstinerent .. .

Translation

1342. The inhabitants of Greenland fell voluntarily from the true faith and
the religion of the Christians, and having abandoned all good manners and true
virtues, they turned to the peoples of America. Some are of opinion that Green-
land is quite close to the western regions of the world. This was the reason
why the Christians began to refrain from the Greenlandic navigation . . .4

Here it is thus said with regret (in 1637) that the Icelandic settlers
of the Western Settlement left the community and Christendom in
Greenland ; that about 300 years before 1637 they had emigrated to
the heathen people on the other side of the ocean. One would think
of the savages in Markland or Vinland, the Skrzlings (Icelandic,
skrelingjar), i.e., the American Indians and Eskimos.

FINAL COMMENTS ON THE DANISH TREATISE (1947)

In 1949 I mentioned the Kensington stone for the second time in
a short account of all the runic inscriptions hitherto discovered in
Greenland.® Most of these date from the time shortly before or after
1300. Many of them are fragments only, partly unintelligible. The
longest of them is the one found farthest north, on the small island
of Kingigtorssuaq about 10 miles north of the Danish colony Uper-
navik (lat. 72° 58’ N.). It has already been examined here (p. 6 f.;
er. fig. 1).

As to the Kensington stone in Minnesota I emphasized again intro-
ductorily in 1949 the great number of features suggesting that it is a
forgery. The “wrong” features undeniably make a strong effect by
their number and kind. Thus it has been objected that a few of the
runic signs fall outside ‘‘the short runic alphabet,” the futhark, which
was universal about the year 1300. The same, however, also applies

4Grénlands Historiske Mindesmerker, vol. 3, pp. 459 f., 887. Also mentioned
by P. A. Munch, Det norske folks historie, Unionsperioden, pp. 313-314, 1862,
and with some criticism by G. Storm, Om Gisle Oddson’s Annaler, Archiv f.
Nord. Philoiogi, vol. 6, Lund, 1890. Cf. also Holand (1932) pp. 82-84.

5 Thalbitzer, W., Runeindskrifter i Grgnland, Grgnlandske Selskab Aars-
skrift 1949, pp. 85-92.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

to the “Dalecarlian runes,” which held out in the Swedish province
of Dalarne right down to the last century, but which, indeed, are
genuine enough in their own way and originate continuously from
the Middle Ages. The anachronisms of the Kensington runes must
be viewed with this background (cf. the following notes). They form
a group apart developed from the runes of the past (in Sweden), but
mixed with neologisms or ‘“‘modernisms.”

The Kensington inscription belongs to a period of decay in the
Swedish (in part also the Norwegian) community, language, and
literature. The vocables and grammatical forms of the old language
(classicism) were no longer taken seriously in the Scandinavian coun-
tries but were increasingly influenced by new “modes,” the German
manner, pouring in from the south. The Hanse immigrants made a
strong impression. German and French books obtained access to the
literature ; black-letter writing became dominant in the vellum manu-
scripts. There was plenty of individual neologisms. If we search
sufficiently long in the letters and documents handed down from the
Middle Ages (about 1350), we shall gradually find examples of
“subversive” deviations from the classical court language which for-
merly reigned supreme. We must also use our imagination: we can
imagine considerable lost material. Read again the above-mentioned
paternoster in Norn from the Shetlands! Study not only the old
Swedish chronicles, diplomas, sigils, and homilies from the fourteenth
century but also the letters and other casual writings handed down
to our time, including Axel Kock’s Swedish dialect-studies (1882-
1929).

In Greenland a particular dialect was spoken, and the runic system
used here had its peculiarities. During the first centuries after Eric
the Red’s colonization of part of West Greenland (shortly before the
year 1000) there was a lively communication between Greenland and
the recently discovered countries in the south, on the east coast of
North America. According to the old Icelandic monastic annals the
bishop of the Norse Greenlanders in 1121 went to Vinland: Euzrikr
Uppst Gronlendinga byskup leitadi Vinlands, or according to another
coeval annal: fdr at leita Vinlands “started (from Greenland) in order
to visit Vinland.” As late as 1347 the Icelandic annals mention a ship
“which had gone from Greenland to Markland with 18 men on board
and now with a lost anchor entered a West Icelandic fiord.” The
following year the crew went on to Bergen. Everywhere they ad-
vanced the names of Markland and Vinland were revived and became
topical. But nothing is stated about the great lakes or any stone with
runes or other things they may have observed over there.

NO. 3 TWO RUNIC STONES—-THALBITZER 53

The Kensington stone may be a forgery made later. But “sign
stands against sign,” | repeated in 1949. There isa conflict of evidence.
The language of the Kensington inscription has an old-fashioned
ring in spite of everything, e.g., the datival form eno, “this,” governed
by a preposition: frd(m) peno sten, peno 6h. These expressions,
however, are “bad grammar” from the point of view of that time.
The vocalism of the Old Swedish is more convincing, the vocalic
qualities demonstrated by Axel Kock, so characteristic of the vowels
of the Gotish dialects of the Middle Ages. The words of the
Kensington stone show the same nuances of vowels as the Swedish
medieval language. The frequently mentioned odd runic letters may
be derived from the black-letter writing of the fourteenth century ;
the rune carver must have been familiar with them and transferred
them to the stone in a slightly modified form.

How did this come about? For seven years he had been traveling
in company in which different dialects from Sweden and Norway met
and were mixed into a kind of insular language. The small party
was itself a traveling enclave that was forming a mixed or in part
artificial language in order to facilitate mutual understanding. The
result was a lingua franca, ground-down words without inflexional
endings, phrases and forms of bad grammar, like eptir wi kom hem;
[iak or wi] fan 10 man rode af blod, [tak or wi] har 10 mans we
hawet—all, however, or nearly all, forms found sporadically in Swed-
ish and Danish literature (letters, laws, rhymes) from the fourteenth
century. This may be indicative of the spoken language already then
to a great extent having abandoned the use of the Old Norse endings.
We can use a good number of these sporadic finds to explain the
anachronisms and irregularities in this peculiar inscription.

This is just what I have done in my article in Danske Studier °
(printed in translated form above), in which our runic stones—those
of the whole of Scandinavia—have often been discussed. The Ken-
sington runes are not difficult to interpret and apparently contain no
obscurities. The mystery consists in the geographical and historical
place of the stone and the fatal intrinsic complexity of the inscrip-
tion. The strange runes and word forms in my opinion may be partly
explained from the peculiar circumstances attaching to Powell Knuts-
son’s “expedition of discovery.”* After all, the anomalies of this
inscription are no more remarkable than those of the Kingigtorssuaq
stone from Greenland, which is genuine!

6 Kgbenhavn, 1946-47 (pp. I-40).
7 Thalbitzer, W., Powell Knutssons rejse, Grénlandske Selskab Aarsskrift,
1948.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

In other words, a close analysis of the phonetic and paleographic
details supports the adoption of a new view of this remotely isolated
runic inscription. Still there are a few difficult knots left as long as
we have not yet succeeded in finding, in the literature about 1350, the
use of an 6 with two dots over it and the use of an enclitic h to denote
vowel length, as, e.g., in the final line: fram peno oh ahr 1362 instead
of 6, “island,” and ar, “year.” To this day it must be admitted that
the authenticity of the inscription is uncertain. But we may hope that
we shall succeed in finding an explanation or, still better, a proof on
these points, too.

The last word has not yet been spoken.

NOTES

I. THE GREENLAND RUNES AS CARVED ON STONE OR WOOD IN GREEN-
LAND, WITH CORRESPONDING ICELANDIC LETTERS (TO THE
RIGHT )

Mariela B oie Bist)
HanduGe Cinoaiy
Nanna Cer:
Nh u,v * h,g,gh)
I. (yo Peet

lanes D ane ie
As MA peer iat
ATU P (p)

Bb 5p.) sleet
d4 d,d,p,t brit 2)

2. THE SECRET RUNES ON THE KINGIGTORSSUAQ STONE

Magnus Olsen has later altered part of his interpretation from 1932
and found out that the six mysterious signs indicate the letters FU,

NO. 3 TWO RUNIC STONES—THALBITZER 55

ie., the beginning of the runic alphabet (the fupark), pars pro toto
probably being used for exorcism against some threatening peril,
maybe the stubborn sea ice or the Eskimos. The two letters, how-
ever, are placed in inverted order (UF) in accordance with an old
custom: “To sorcery belongs all that is done backwards, e.g., read-
ing the Lord’s Prayer backwards or moving in the wrong direction,
to the left, counterclockwise, etc.” ®

Further, I shall here mention Magnus Olsen as a keen advocate of
the theory of numeral magic being hidden in most medieval runic
inscriptions. Already in his first papers of 1932 (cf. also above pp. 9
and 13) he proved that the Greenland Kingigtorssuaq inscription,
like several of the runic inscriptions on the Orkneys, is pervaded by
an artificial numeric system, i.e., the number of runes and points
(. or :) within each line and each section and for the inscription as
a whole is accommodated according to certain desirable “lucky num-
bers” (e.g., 9, 18, 24, 48, etc.). Therefore the manner in which
the words are spelled and the points arranged is often influenced by
this consideration. To read the runes is called to radu, the same word
as Gothic rapjan, “to reckon, to count.” The word “rune” (rin) in
itself means “secret.” The question of numeric magic is not men-
tioned frequently in the old literature, maybe only indicated once.®
But it may be understood that it was practiced in the later runic period
from the fact that well-known runologists such as Finnur Jonsson,
Magnus Olsen, and others vouch for the existence of it in the vari-
ous inscriptions they have examined for the purpose of counting their
secret “lucky numbers” and systems of numbers.’°

Our comparatively recent discovery of this custom of “counting
the runes and signs,’ observed alike in the wording and reading of
(perhaps) most of the old inscriptions, will come in very useful for
our explanation of the anomalies or apparent mistakes in them. From
now on we should be allowed to consider the irregularities not as
the mistakes of a carver or scribe, not as accidental slips, but as tech-
nicalities aimed at, a sort of trick. That is, for instance, how Mag-
nus Olsen interprets the singular distribution of the points of punctu-

8 Olsen, Magnus, Sigtuna-amuletten, 1940 (p. 23) ; En futhark-innskrift i Lom
kirke, 1943 (pp. 91-92). Cf. his treatises from 1932.

9 Magnus Olsen cites the Busluben; cf. his Sigtuna-amuletten, p. 29, where
he also mentions papers by N. E. Hammarstedt and A. Nordén. Cf. also
Feilberg, H. F., Tallene i folkets brug og tro, Dania, vol. 2, Kgbenhavn, 1892-94.

10 The first Scandinavian scholar to view the study of the runes in a broader
light of Byzantine, Semitic, and Oriental comparisons (from about 1927 on)
was the Swede Per S. Agrell.

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

ation and the faulty spellings and some of the secret signs of the
Kingigtorssuaq inscription.

If this theory of numeric magic, which was not advanced till some
decades into this century—thus long after the discovery of the in-
scription discussed here—holds some truth, all runic inscriptions ought
to be considered in the light of it. We shall revert to this question
below.

On the basis of that theory, all fairly long inscriptions would have
required laborious preparation. They could not possibly have been
improvised. They may be supposed to have been worked out on the
basis of a kind of draft (written on wax tablets or wood or hide,
etc.) before being carved into the stone.

3. SINGULAR RUNES ON THE KENSINGTON STONE, X anp F

If a cleric or priest in 1362 really wrote these runes as a kind of
draft (on parchment or wood or the like) in order either to carve
them himself or to have somebody else carve them on the stone, he
must—and this is the most probable—have obtained his education in
a monastic school where such runes or quasi-runic signs were known
in the fourteenth century.

A fairly great number of the runes agree with those known to us
from the Skanish runic manuscript (Codex runicus) now found in
the Arnemagnaean Collection in the University Library of Copen-
hagen (AM 28, 8°) and considered to have been written about 1300.
Besides the well-known runes, there are on the Kensington stone a
small number of extraordinary forms which seem to resemble the
majuscules and minuscules used in the Middle Ages. Similarly the
Skanish runic inscription is supposed to have been transferred into
runes—a conscious anachronism—from an original with ordinary
alphabetical writing.

On the Kensington stone particularly the two runes for a X
and 7 * give some trouble. I have above (p. 25) suggested that the
a-rune might originate from a provincial form like the Dalecarlian
rune ( X), which it somewhat resembles, and that the so-called 7-rune
(if, indeed, it is 7) might be an imitation of a manuscript 7 or 7
(ordinary form of the letter 7) in the earliest documents, used alike
for the vowel and the semivowel (cf. fig. 6, p. 22).

The sign X—with an extra dot in the a rune, used in this inscrip-

11 Hanninger, N., Fornskansk ljudutveckling, Lund, 1917 (pp. 4-5) ; Nielsen,
Lauritz, Danmarks middelalderlige haandskrifter, Kg¢benhavn, 1937 (pp. 116-
117).

NO. 3 TWO RUNIC STONES—THALBITZER 57

tion to denote the sound of @—seems to be a secondary formation.
The a X of this inscription perhaps imitates the black-letter of the
manuscripts (cf. fig. 6). In the a and @ of the minuscules we find
the same small hook as found again in the runic form appearing here.
The rune carver may have been familiar with these letters from his
school, or may have brought along a calendarian runic tablet with
the alphabet (perhaps “golden figures”) on which the runic a was
given the same form as we later find in the Dalecarlian runes—the
form which we thus find already on the Kensington stone.

4. 6 =OE
The relation between some of the runelike pairs of vowels of the
Kensington stone appears from what follows:

XX a-—a
YA u—ii.

The dots are taken from the hand-written letter writing. In the
lower pair the first “rune” has a dot between the left twig and the
main stave in order not to be mistaken for the normal m-rune of the
Kensington inscription. This is the starting point of the pair of
rounded labial vowels in the cleric’s alphabet. The last rune has an
extra dot as often the letter denoting y (ii) in the fourteenth century
(see fig. 6). At the same time it has been modified somewhat, viz,
halved and the stave provided with a transverse stroke. Here we
see a vague connection with the Greenlandic runes (see p. 54), as
the y- (i-)rune in Greenland about the year 1300 may have one of
these forms:

Yorenrs
the latter with downward-directed twigs as in the runes of Norway.
Cf. the special Icelandic form for y: A.1* The carver of the Kensing-
ton runes has preferred the upward-directed rune and individualized
it with the dots. He must have cultivated this principle in his runes as
he must have done in his personal writing.

5. OPDAGELSEF&RD (see pp. 35-360)

This word, extraordinarily long for that time, perhaps should be
interpreted in the light of line 10: “(we) have ten men at the sea
to look after our ships.” For whom was this information intended?
Was there anybody who awaited the men’s arrival? Who was to be

12 Beksted, A., Islands runeindskrifter, p. 40, 1942; Jonsson, Finnur, Interpre-
tation of the runic inscriptions from Herjolfsness, pp. 289-290.

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

informed of the presence of the ship(s) ? And was there among the
men participating in the expedition any definite expectation as to
whom they were to “opdage,” i.e., search for?

Opdage in the old Icelandic- Norwegian language particularly meant
“surprise trolls at daybreak.’ In Greenland the Skrelings were called
trolls,’* and as the name of Skreling originally had been used about
the natives of Markland and Vinland, it was just here (in America)
that people in the Middle Ages could surprise trolls, “daga upp troll.”
From this meaning the word may very easily in the fourteenth cen-
tury have been generalized to an approximation of the modern mean-
ing of the word updaga: “discover.” And P. Knutsson’s men no
doubt knew for whom they were searching.

6. HAPXR =skiar or ? (ENGL. scar, “CLIFF, ROCK”

The rune f is otherwise unknown. It is either to be taken as a
retroverted + (d or 0) or as a ligature (“binderune”) of le or el,
or to be read as j, or to be a clerical error for the e rune ¢.

Reading the rune as 7 (as Holand does) is a makeshift, since
the form skjar is unknown from the OSw. or ON. of the Middle
Ages. Moreover, the 7-rune which belonged to the primary runic
alphabet (the futhark of 24 letters) had been dropped in the later
alphabet of 16 letters. The word, however, would remind of ON.
sker or sker, Engl. scar or skerry, and thus make sense in the con-
text ; either it would be interpreted as a big stone (rock, boulder) on
land, or as a rock in the water, standing out like a small island
(skerry).

In my Danish article (p. 33) I have taken the word skjar to be
an artificial “broken” form (brydning) from the fourteenth century
(although the phenomenon of breaking had otherwise been abandoned
long ago), viewing it as a poetic word to be construed on the analogy
of similar inherited forms. The rune then might be considered a j,
but a special rune for 7 was not extant in the shorter futhark of that
period. If it is really a j, the rune carver must have been a learned
cleric who was familiar with many Swedish runic inscriptions where
a j-rune might easily have been found, as it was extant in the old
futhark (24 signs). The rune used in the Kensington inscription is
of an otherwise unknown form, maybe constructed on the pattern of
the black-letter script, even a retroverted 7 (cf. fig. 6). But in the
Middle Ages they did not distinguish between i and 7. So it is

18 See Grgnlands Historiske Mindesmerker, vol. 3, pp. 438-439 and 460.
Cf. Thalbitzer, W., in Meddelelser om Grégnland, vol. 31, pp. 28-29, 1904.

NO. 3 TWO RUNIC STONES—THALBITZER 59

nothing less than a riddle how the carver came across the idea of
constructing such a special sign for 7. Hence, I am more inclined to
consider the rune as misrepresented and mistaken: the carver had
intended to make a simple | ; the rest of the sign was mere blunder-
ing. If so, we have here a spelling with ia simply to denote the
letter @: skiar to be read as sker. This manner of spelling with ia
instead of ae (the Latin way), according to Axel Kock, is found in
a great many cases in the documents from the fourteenth century.
The diphthongization of the sound @ is not uncommon in OSw.
dialects, as in siel (or sial) alongside of sel, “soul”; i hel for i hjal,
“killed, dead”; in runic inscriptions mierki or miarki for merke, “a
mark.” Kock further writes: “In certain OSw. documents a not
rarely occurs instead of an expected @ . . ., e.g., kannir for kennir
(now kdnner, “knows’’) ; 7 malli, emallan, mallen (ModSw. mellan,
“between”), map for mep, “with”; gare or giare for gera (now
gora, “do, make”).” Such forms may be clerical errors, thinks Kock,
though he is not sure.** Other examples are OSw. aft alongside of
eft; aftir or eptir, “after”; qvar or quer “clam”; qwarn or qvern,
“grinding mill”; hwar or hver, ON. hverr, ModSw. hvar, “every.
Forms like miarki, sial, giare would seem to corroborate our im-
pression of skiar as an OSw. form used instead of sker or skar (cf.
Engl. scar). With some reservation it would also be comparable to
the latter part of the name of Torbiarn, which is found in OSW.
alternating with Torbian, later Torban (cf. the Icelandic form Banne
for the early Biarne).1° If we are to read skiar, we might characterize
the form as imitative of the old forms called ‘broken’ forms, an
antiquated form.’’ But if we have to take ia as denoting ae (reading
sker), the word would be quite regular and the mystery solved.

7. ris€=MODERN SW. resa, “JOURNEY, (WARLIKE) EXPEDITION”
Peco. 40.1.)

Thus used in the Swedish “Rimkronike,” the continuation about
King Magnus Eriksson, from about 1450. The original of this sec-
tion of the rhyming chronicle was probably composed and written

14 Kock, Axel, Svensk Ljudhistoria, vol. 1, pt. 1, pp. 256-258, 266-267, 1906.

15 Kock, Axel, loc. cit., pp. 237-250.

16 Kock, Axel, loc. cit., vol. 2, p. 404, 1909-11. Banne is the form we found
on the Kingigtorssuaq stone from Greenland (cf. p. 10).

17 It would remind of an old-fashioned form such as our Danish skjald along-
side of Swedish skald, English scald, ON. skdld. Cf. in OSw. the name of
Spjellebod, in which spjalli-, “associate” is a “broken” form of spell, Goth.
spill, OE, spell.

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116
down soon after the king’s death, which took place in 1374 (Decem-
ber 1),'* but as it has been lost and the copy dates from at least 50
years later, we cannot be sure that its language is in every respect
exactly as it was written in the preceding century. However, there is
nothing surprising in the fact that a German word should have been
adopted in the Swedish language already at the time of the original
owing to the German immigration into Sweden about 1350, let alone
the possibility that one of the members of the Powell Knutsson expe-
dition may have been of German extraction, even though Swedish-
speaking. Already in the beginning of the eleventh century a Ger-
man named Tyrker participated in Thorfin Karlsefne’s expedition.

Here follow some lines from the above-mentioned section of
Sveriges Rimkronike in the linguistic form in which it has been
preserved.’® Note the word resa:

Verse

114 konungen hade skane fangith
127 han ville ena resa till rytza-
landh fara
775 ‘Tha rytza komma j
behaldh
344 huar hade en binzel om sin hals
ok sagde, vy aro her komne
aa jdre nade
Offuer vara lyiff ok lemmer
magen j rade
thz vy jdher loffuade kunnom
vy ey holde
thy giffua vy os jdher j volla

thera

The King had got Skane

He would go on an expedition to
Russia

When the Russians came in their poor
state

Each had a chain round his neck

And said, We have come here by your
grace

You may rule over our lives and limbs

What we promised you we cannot
fulfill

Therefore we place ourselves in your
power

Note the absence of inflexional endings after prepositions (om sin
hals ; ffuer vara lyiff) as on the Kensington stone (aptir) wore skip;
and in the last line vy (wi) giffua instead of giffuom. (More ex-
amples on p. 41.)

The anomalies on the Kensington stone become less astonishing by
such comparisons. They have not been far from the spoken language
which was developing and was to be realized soon after. A purist or
champion of Modern Norwegian might, about the middle of the
fourteenth century, anticipate those kinds of forms in his draft for
the inscription. Particularly if he was a foreigner (a German) who
was to write down the language of a (Swedish) Goth or Norwegian.

Such anomalies may occur at any time, sometimes in an astonish-

18 Mverland, O. A., Norges Historie, vol. 3, p. 867, Kristiania, 1888.
19 Klemming, G. E., Svenska Medeltidens Rimkrgnikor (in 3 vols., 1865-68),
vol. 1, p. 175 fff.

NO. 3 TWO RUNIC STONES—THALBITZER 61

ing accumulation in a narrow space. We have seen a similar occur-
rence on the Greenland runic stone from Kingigtorssuaq (cf.
pp. 7-13).

I want to repeat: what cannot a Greenland rune carver permit him-
self on a dangerous expedition of discovery to the Ultima Thule!

8. at se Gptir, “TO LOOK AFTER, GUARD, SUPERINTEND (THE SHIPS)”
(see lines 10-11 of inscription)

This expression is known from Old Icelandic and may as well have
belonged to the Old Swedish dialect as the above-mentioned other
phrases of an old-fashioned character (of west, aptir wi kom hem,
etc.). Cf. Olcel. sya eptir einu in Fritzner, Ordbog over det gamle
norske sprog, Kristiania, 1886, e.g., ctlu vér at eigi muni adrir meirr
eiga eptir sinum. hlut at sjd in the saga on Vigastyr ok Heidarvigum
(and many other examples).

g. dagh rise, “DAY’s JOURNEY” (see line 11 of inscription)

14 dagh rise, “day’s journeys,” from the “coast of the ocean” is
a somewhat elastic concept, particularly considering the times in
question—vise was a new word. The travelers had no telemeters ;
the regions through which they were traveling were quite unknown
to them. They may have carried a calendar with them and been able
to check the days and dates. Note that, in sharp contrast to the
Kingigtorssuaq stone, the Kensington stone does not give any calen-
dar reckoned date.

Fourteen days gives only an approximation, just as we say “a
fortnight.’ It must have meant from the nearest seashore.

“The ocean” need not be a scientific term, either. “Our ships by
the sea” may have been intended to refer to the nearest shore of Lake
Superior or Lake Winnipeg, or perhaps to the mouth of the Nelson
River in Hudson Bay. If the men had passed the Niagara on their
expedition they would have had to build new ships (or boats) above
the falls. It would have been easy for them to build ships. Perhaps
the “hafsvelg’ of the old geographers is the “falls” where “the ocean
washes the threshold of the earth,” to use another medieval term.*°

20 Cf. Franciscus Irenicus, Germaniae exegeseos volumina, p. 200, Hagenau,
1518; Jos. Fischer, Die Entdeckungen der Normannen in Amerika, Freiburg,
1902; Grgnlands Historiske Mindesmerker, vol. 3.

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

I0. THE STYLE

Explaining and Explaining Away
Scandinavia in the Middle Ages

Pattern and correctness in language, it is true, are firmly anchored
in the classical well-balanced society near court surrounded by the
pursuits of peace until the penetration of the new fashions, due to
the immigration of the foreigners, begins to change the mold of
language.

On this opdagelsefird the very ligaments of language seem to have
broken. Here we trace a decomposition, a confusion, the penetration
of foreign elements coupled with a counteraction, a puristic tendency
and intention. I picture to myself an Old Swedish cleric recently
arrived in Greenland, with tendencies toward new forms aroused in
him, perhaps under the influence of a custom of language and writing
found in the Norsemen’s settlements in Greenland which was strange
to him. The expedition must have passed the Norse settlements in
Greenland and perhaps the men there learned something new. The
Swedish cleric, among other things, must have learned to know the
Greenlandic runic custom and have had a fresh recollection of his
new knowledge in inland Minnesota. Powell Knutsson’s expe-
dition may have been accompanied by a man from Greenland ac-
quainted with the localities, who was to act as pilot on the voyage to
Vinland, and a Norse-Greenlandic cleric. Another stimulating imple-
ment may have been a runic calendar made in Greenland. Little by
little we are removed from the Norwegian-Swedish patterns.

My severe critic, the Swedish docent Sven B. F. Jansson, finds
proofs of a forgery in the great number of grammatical errors in the
inscription. Of course he is right: this mess from Minnesota denotes
a revolt against the style of the fourteenth century in writing and
language. It so greatly challenges the suspicion of all upright phi-
lologists that only a heretic can contradict them. Moreover, the
learned historian Gustav Storm, in the last century, about a decade
before the stone was dug up, had aroused all historians’ interest in
the old voyages to Vinland. About 1890 there was a romantic tune
in the air—also in America as far as the lake district—a great possi-
bility that this thrilling tune might be caught by a monomaniacal Vin-
land fanatic in Minnesota: “Let me arrange a practical joke in honour
and to the delight of the good Swedish farmer Mr. Ohman, who shall
find his own visiting-card between the lines of this runic stone which

21 Jansson, Sven B. F., “Runstenen” fran Kensington, Nordisk Tidskrift
(Letterstedtska), Stockholm, 1949.

NO. 3 TWO RUNIC STONES—THALBITZER 63

I secretly bury on the outskirts of his fields, under the ground among
the roots of the tree!” dh ahr 1362, left anonymously.

Rogues have existed at all times. From the fifteenth century we
know the Dane Claudius Clavus (from Funen, born 1388), who put
fictional names at all rivers and headlands on his Ptolemaic map of
the North, even along the coasts of Greenland, where he had never
been.*? The Kensington rogue was a century ahead of the Dane.
The stone had stood and had tumbled into the soil of Minnesota 500
years before this last rogue was exposed by a Swede!

II. MEDIEVAL SWEDISH-NORWEGIAN MIXED LANGUAGE

On the decay of the language in Norway-Sweden about 1330-
1350, P.A. Munch wrote at two places in Det norske folks historie,
Unionsperioden, II Anden Hovedafdeling, 1862. The following is
an extract from that work (pp. 362-364).”°

But directly Queen Euphemia 24 had begun to spurn the Norwegian transla-
tion and instead encouraged French translations of the fashionable literature
of the time, e.g., the Norman-French adaptations of the legends of King Arthur
and the Knights of the Round Table, this kind of literature seems particularly
to have become the fashion, an end to which the union with Sweden also con-
tributed. From this time we may take it for granted that Swedish or a mixture
of Swedish and Norwegian became the language of the court, which again had
an unfavorable effect on the purity of the language in general, of which much
evidence is extant in many public papers of the time (e.g., bondomen for
bondunum, lagomen for Iggunum)?> ... Among the books which in 1340,
according to an inventory, were found in the Castle of Bagahus [now Bahus],
there were, besides a good number of Swedish books, also some German books,
the Swedish translation of Iwan and Gawian arranged for by Queen Euphemia,
a German Bible and lawbook . . . With the sinking interest in the old national
literature and in the occupation with standard native language the latter lost
its purity ...and now was written more slovenly . .. the fixed rules were
disregarded. The dialects asserted themselves . . . we find in letters and litera-
ture the popular speech (“Almuesprog’’) itself instead of cultivated and well-
arranged written language, although with some exceptions in the official docu-
ments proper.

22 Claudius Clavus (Claus Claussén Swart) [the first cartographer of Scandi-
navia]. A monograph by A. Bjgrnbo and Carl S. Petersen, Kgl. Danske
Videnskabernes Selskab, ser. 6, vol. 6, 1904, K@benhavn.

23 Cf. also Holand, Hjalmar R., The Kensington Stone, pp. 98-103, 1932;
Westward from Vinland, pp. 152-158, 1940.

24 Fuphemia, the Queen of Hakon V (1299-1319).

25 Many more examples of similar “wrong” dative-plural forms are found in
Swedish Medeltids-Postillor (Samlingar ed. by Sv. Fornskrift Sallsk.), vol. 5,
1909-10, e.g., p. 248: fordldroman, dnglachoroman, kenneswenomen, scriptama-
loman. There is also another example of the dative ending in y-: illy instead of
alli (illu).

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Again, later in his work, P. A. Munch has some pages (596-599)
about the mixing and decay of standards. His presentation really
ought to be translated and added here because it so marvelously falls
in with the amazing character of the idiom on the Kensington stone.
It would explain a good many of the anomalies of its grammar. Add
to this that no other has been more familiar with the documents and
letters extant from that period in Norway and Sweden than that great
Norwegian historian and philologist.

I2. OLD SWEDISH AS A HYPOTHETICAL PROBLEM

Prof. Axel Kock (whose works of 1882-1886 and 1906-1929 have
been used in what precedes) was a specialist on Swedish dialects,
both those of the Middle Ages and those still current. Ad. Noreen
stood beside him in these studies (cf. here p. 26, Noreen’s Dalska
runinskrifter, 1906). Kock uncovered a number of medieval dialects
in the provinces of the Goths and the Svear and could demonstrate the
occurrence of dialects now extinct.

In Noreen’s work on the Dalecarlian runes these are considered a
survival of a number of medieval popular runic systems once oc-
curring also in other Swedish provinces (Harjedalen, etc.), but most
of them now extinct; only the system of Dalarne being preserved.
Both the runes and the language of the Kensington inscription are to
a certain degree different from all this. Hence it is excluded that a
forger, if the inscription is a fabrication, should have been a man
from the province of Dalarne. The same is true of his dialect (cf.
here p. 27).

A faker from before 1898 might, but only through Axel Kock’s
works, have obtained a detailed knowledge of the vocalism of the
Old Swedish words in the inscription as analyzed here (pp. 29-32; cf.
pp. 43 and 53). But he could not possibly have known Kock’s manu-
scripts of the works then not published. And it is not very likely that
he should have had information sent to Minnesota from the author’s
(Kock’s) own intimate circle of scholars.

It is evident that the idiom of the Kensington inscription is not at
all a modern one but old-fashioned. It contains twice the old dative
form peno, which a faker would not have invented; likewise the old-
fashioned illu from the nominative ila. And how should the scribe
have hit the right nuances of the Old-Swedish vowels without having
been quite familiar with these forms from his own natural language:
Géter instead of Gotar; op (not up), ve(d) (not vid), skip (not
skepp), ar (not dr). Also, a form like his aptir instead of New
Swedish efter is markedly OSw.

NO. 3 TWO RUNIC STONES—THALBITZER 65

Conclusion: the spelling of the inscription, perhaps even the word-
ing of it, must be due to a man who himself spoke an Old Swedish
dialect as his mother tongue, probably an extinct and to us hitherto
unknown dialect.

13. A FINAL COMPARISON BETWEEN THE TWO RUNIC INSCRIPTIONS

As already pointed out, there are certain points of resemblance
between the two inscriptions.

Finnur Jonsson in the annual of Det Grgnlandske Selskab, 1914
(pp. 94-97), raised the question with reference to the Kingigtorssuaq
stone, whether (or not) it is a fraud (cf. above, pp. 6-7). He came to
the same conclusion as do all other interpreters: the stone is genuine—
there is no cogent reason to doubt the authenticity of this (the Green-
landic) inscription.

As a suspicious circumstance it is mentioned that C. A. Stephen-
sen, who was a manager of the northern Upernavik district, where
a native Greenlander found the stone on the ground beside the cairn,
was himself an Icelander by birth and that the names of the Icelandic
men mentioned on the stone as members of the expedition are ren-
dered (spelled) in conformity with modern Icelandic pronounciation
(cf. above, p. 10), a remarkable feature in an inscription dating from
the beginning of the fourteenth century. In order to exclude the
suspicion, similar reasons must be adduced as in the case of the
Kensington inscription: how could a modern forger, who would
appear to have a good knowledge of the old language and the runes,
make a fool of himself to such a degree, if he really wanted to gain
credence? Finnur Jonsson took great pains to prove that Stephensen,
the Icelander, would have been unable to play such a sophisticated
joke: that of carving quite modern names in an inscription which
otherwise closely resembled the ancient language. “This suspicion
must be flatly rejected.”

There are some few graphic points of resemblance between the
two runic inscriptions.

An 0-like rune combining o and e is found on both (pp. 12 and 26;
cf. p. 44).

The strange b-rune (|) of the Kensington stone approaches the
Greenlandic b ( b) occurring four times in different places in the
Western Settlement. Here, e.g., it reads in runes: Ave Maria grazia
bina (i.e., plena), misspelling with b (the -rune).?° And there are

26 See Moltke, E., Greenland runic inscriptions, Meddelelser om Grgnland,
vol. 88, p. 225, 1936, Sandnes inscription 2, notes 1 and 2.

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

two other runes which the Minnesota and Greenland stones have in
common, the reversed k ** and the B used for p. These agreements
might indicate that the carver of the Kensington runes has been influ-
enced by some peculiar Greenlandic forms of style. Therefore it is
thinkable, in fact credible, that he has passed via Greenland to
Minnesota.*®

As regards the contents, the comparison is also interesting: two
or three of the runic inscriptions of Greenland show similar magic
spells connected with the holy name of the Virgin Mary. And both
inscriptions have a definite dating: the Greenlandic stone both year
and day, the Kensington stone at any rate a definite year.

The differences, however, are apparent: (1) The Minnesota stone
is without any magic sign—even though the Latin letters dV M look
like a kind of magic (a spell) in the middle of the realistic report.
The Kingigtorssuaq stone is full of secret runes (also “ligatures,” or
“knitted” ones, i.e., two runic letters united on one staff), and it
contains plenty of mystical exorcising, numerical magic (cf. Magnus
Olsen). (2) The former bears a personal stamp by its repeated
“we,” the pointing out of a lesser number of Goths and another but
major of Norwegians—but without giving one personal name. The
latter mentions three persons by their full names, but in the third
person, impersonally.

Both stones are connected with a dangerous expedition of dis-
covery via South Greenland to Ultima Thule, or to Vinland, with the
explorers in a fateful situation. Both are among the most dramatic
runic inscriptions ever known.

27 Thid., p. 228, Sandnes 9, note 2.
28 Cf. my Runeindskrifter i Grénland, 1949; also above, pp. 16 and 62 (cf. pp.
57-58).

BIBLIOGRAPHY
AASEN, IVAR.
1873. Norsk Ordbok, Christiania.
AGRELL, PER SicurpD (1881-1937).
1927. Der ursprung der runenschrift und die magie. Arkiv f. Nord. Filologi.
1931. Senantik mysteriereligion och nordisk runmagi.
1932. Die spatantike alphabethmystik und die runenreihe.
1934. Lapptrummor och runmagi.
1938. Die Herkunft der Runenschrift.
B2KSTED, A.
1939. Vore yngste runeindskrifter. Danske Studier, vol. 36. Kgbenhavn.
1942. Islands runeindskrifter. Bibliotheca Arnamagnzana, ed. Jon Hel-
gason, vol. 2. Kgbenhayn.
1943. Runerne, deres historie og brug. Kgbenhavn.
BrateE, E. (See also Séderberg and Brate.)
1911. Oster-Gotlands runinskrifter. Sveriges Runinskrifter, vol. 2. Stock-
holm.
1922. Sveriges runinskrifter.
1924-32. Sddermanlands runinskrifter. Stockholm.
Brix, Hans.
1928-29. Studier i nordisk runemagi, I-II. Kgbenhavn.
1932. Systematiske beregninger i de danske runeindskrifter. Kgbenhavn.
BréccEr, A. W.
1928. Gamle Emigranter. Oslo.
BréNDUM-NIELSEN, J.
1917. Danske runeindskrifter. Aarbgger f. Nordisk Oldkyndighed og Hist.
K¢gbenhavyn.
1933. Danske runeindskrifter. Nordisk Kultur, vol. 6. Stockholm, Oslo,
K¢benhavn.
1928-1935. Gammeldansk Grammatik, vols. 1-3. Kgbenhavn.
1943. Paleografi. Nordisk Kultur, vol. 28.
Dyseck, R.
1855-76. Sverikes runurkunder, vols. 1-2. Stockholm.
FaLk, Hyatmar, and Torp, ALF.
1900. Dansk-Norskens Syntax. Christiania (Oslo).
1910. Norwegisches-Danisches Etymologisches Worterbuch. Heidelberg.
1913. Upplands runstenar. Uppsala.
FRIESEN, O. von. (See VON FRIESEN.)
FRITZNER, JOHAN.
1886. Ordbog over det gamle norske sprog, vols 1-3. Christiania. (Oslo.)
Hacen, S. N.
1950. The Kensington runic inscription. Speculum, vol. 25. Cambridge,
Mass.
HILpEBRAND, EMIL.
1894. Svenska skriftprof (medeltiden). Stockholm.
HILpEBRAND, HANs (editor).
1900-1935. Sveriges runinskrifter, vols. 1-7. Stockholm.
67

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

HyJELMSLEV, Louts (editor).
1932-35. Rasmus Rask, 1832-1932. (Introduction by Holger Pedersen.)
Udvalgte Afhandlinger. Kgbenhavn.
HoLanp, HJALMAR R.
1932. The Kensington stone: a study in pre-Columbian American history.
Ephraim, Wis. (2d ed. 1940.)
1940. Westward from Vinland. New York. (2d ed. 1942.)
1946. America 1355-1364. New York.
1950. Kensington-stenens mangfoldighet. Nordisk Tidskrift (Letter-
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INDREBG, GUSTAV.
1924. Litt um burtfallet av fleirtal i verbalbgygjingi i norsk ca. 1360-
1530, in Festskrift f£. Amund B. Larsen.
JacossEn, Lis.
I93Ia. Eggjum-stenen. Kgbenhavn.
1931b. Nye Runeforskninger (Glavndrup, etc.).
JAcosBsEN, Lis, and Motrxe, E.
1942. Danmarks runeindskrifter, vols. 1-2 (Atlas and Text). Kgbenhayn.
JANSSON, SVEN B. F.
1949. “Runstenen” fran Kensington. Nordisk Tidskrift (Letterstedtska).
Stockholm.
JESPERSEN, OTTO.
1922. Language, its nature, development and origin. London.
JONSSON, FINNUR.
1914. Runestenen fra Kingigtorssuak. Grgnlandske Selskab Aarsskrift.
K¢benhayn.
1916. Grgnlandske runestene. Ibid.
1922. Runeindskrifterne fra Herjolfsness. Ibid.
1924. Interpretation of the runic inscriptions from Herjolfsness. Med-
delelser om Gr¢gnland, vol. 67.
1928. Festskrift til Finnur Jonsson, 29 Maj 1928. Kgbenhayn.
1929. Runic inscriptions from Gardar. Medd. om Grgnland, vol. 76.
1930. Det gamle Grgnlands beskrivelse af Ivar Bardarson. Kgbenhayn.
KAALUND, Kt.
1903. Palzografisk Atlas. Kgbenhayn.
KLEeMMING, G. E.
1865-68. Svenska Medeltidens Rimkr@gnikor, vols. 1-3. Stockholm.
Kock, AXEL.
1882-86. Studier 6fver Fornsvensk Ljudlara. Lund.
1906-29. Svensk Ljudhistoria, vols. 1-5. Lund.
KroMANN, E.
1943. Skriftens Historie i Danmark. Kgbenhavn.
LANGE, C. A., and Uncen, C. R.
1947. Diplomatarium Norvegicum (Oldbreve). Christiania.
LEVANDER, L.
1910. Dalska runinskrifter. Fornyannen, p. 165 ff. Stockholm.
LiLjEGREN, J. G.
1832. Run-lara. Stockholm.

NO. 3 TWO RUNIC STONES—THALBITZER 69

Lyncsy, KristTEN.
1861. Den oldnordiske udtale. Tidsskr. f. Philologi og Pedagogik. Kgben-
havn.
1863. Udsagnsordenes b¢jning i Jyske Lov. Kg@benhayn.
MAGNUSEN, FINN.
1827. “Oplysninger,” in Antiqvariske Annaler, vol. 4. Kgbenhavn. (See
also under Rask.)
MaGNUSEN, FINN (editor), RAFN, C. C., et al.
1838-1845. Grgénlands Historiske Mindesmerker, vols. 1-3.
MArRSTRAND, VILHELM.
1922. Arsenalet i Piraeus og Oldtidens Byggeregler. K¢benhayn.
1933. Aabenraa ca, 1020-1523 (see p. 25). Kg@benhayn.
1949. The runic inscription of No Man’s Land. New England Quarterly,
vol. 22, pp. 85-92.
1951. Asmild-stenen (separate from Viborg Stiftstidende). Kgbenhayn og
Viborg.
Mo.tTKE, Ertx. (See also Jacobsen and Moltke.)
1936. Greenland runic inscriptions. Meddelelser om Grgnland, vol. 88,
No. 2, Appendix. Kgbenhavn.
Mo.rKe, E., and ANDERSEN, Harry.
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Muncu, P. A.
1862. Det norske folks historie. Anden Hovedafdeling. Unionsperioden,
first part. Kristiania.
1877-88. Algorismus ... efter Erlendsén’s Codex. Annaler f. Nordisk
Oldkyndighed, vols. 1-2. Kgbenhavn.
NECKEL, GustTaF.
1913. Die erste Entdeckung Amerikas im Jahre 1000 n. Chr. Jn Voigt-
lander’s Quellenbiicher, vol. 43. Leipzig. (2d ed., 1934.)
NIELSEN, LAvuRITZ.
1937. Danmarks middelalderlige haandskrifter. Kgbenhavn.
Noreen, ADOLF.
1892. Altislandische und altnorwegische Grammatik. Halle.
1892-1894. Altschwedisches Lesebuch. Halle. (2d ed. 1904.)
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holm.
Noreen, Erik.
1944. Svensk stilparodi. Stockholm.
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Ibid.
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vol. 6. Stockholm, Oslo, Kgbenhavn.
1940. Sigtuna-amuletten. Det norske Videnskapsakademi, II. Historisk-
filosofisk klasse, No. 3. Oslo.

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

1941. Norges innskrifter med de yngre runer. Norsk Tidsskrift f. Sprog-
videnskap, vol. 5, Oslo.

1943. En futhark-innskrift i Lom kirke. Maal og minne. Oslo.

PEDERSEN, HotceEr.

1923. Runernes oprindelse. Aarbgger f. Nordisk Oldkyndighed og Historie.

K¢benhavn.
PETERSEN, Cart S.

1950. Wimmer og Kensingtonstenen. Nordisk Tidskrift (Letterstedtska).

Stockholm.
RAAr, L. F.

1838. Bokstafsformer under medeltid enlight Sverges offentliga hand-
lingar (mellan 1164-1513). Kgl. Vitterhets-, Historie- och Antik-
vitets-Akademien Handlingar, vol. 15. Stockholm.

Rarn, C. C. (See also Magnusen, Rafn, et al.)

1837. Antiquitates Americane. Kgbenhavn. (See Societas Regia Anti-

quariorum Septentrionalium. )
Rask, RASMUS.

1827. Efterretning om en i Grgnland funden runesten med dens fork-
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nusen. Antiqvariske Annaler. Kjébenhavn.

Rypgulist, J. E.
1850-1883. Svenska sprakets lagar, vols. 1-6. (Ordbok in vols. 2-3.)
Stockholm.
Sep) DAG
1931. Norsk sprakhistorie til omkring. 1370. Oslo.
Socretas ReciA ANTIQUARIORUM SEPTENTRIONALIUM (editor).
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SODERBERG, SVEN, and BratTe, Erik.
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S@DERWALL, A.
1925. Ordbok 6fver svenska spraket i medeltiden. Lund.
SrorM, GUSTAV.

1888a. Islandske Annaler. Christiania.

1888b. Studier over Vinlandsreiserne. Christiania.

1888c. Om Kilderne til Lyschanders “Grgnlandske Chronica.” Aarboger

f. Nord. Oldkyndighed og Historie. Kjgbenhavn.
THALBITZER, W.

1904. Historical data about the East Eskimo. In A Phonetical Study,
Meddelelser om Gr¢gnland, vol. 31. K@benhavn.

1912. Four Skreling words from Markland in the saga of Erik the Red.
Proc. 18th Internat. Congr. Americanists. London.

1945. Nordboerne ved Upernavik. Grgnlandske Selskab Aarsskrift. Kgben-
havn.

1946-47. To fjerne runestene fra Grgénland og Amerika. Danske Studier,
ed. Universitetsjubilaets Danske Samfund. Kgbenhavn.

1948. Powell Knutssons rejse, en forsvunden ferd til Grénland og Mark-
land. Grgnlandske Selskab Aarsskrift. Kg@benhavn.

1949. Runeindskrifter i Grgnland. Ibid.

NO. 3 TWO RUNIC STONES—THALBITZER ria

TuorsEN, P. G., editor.

1877. Codex Runicus of Skanske Lov, with supplement: Om runernes

brug til skrift udenfor det monumentale. K¢gbenhayn.
Torr, Atr. (See FALK, HJALMAR.)
VON FRIESEN, O.

1928. Runorna i Sverige. Uppsala.

1933. De germanska, anglofrisiska och tyska runorna.—De_ svenska
runinskrifterna. Nordisk Kultur, vol. 6. Stockholm, Oslo,
K¢benhayn.

Wimmer, L. F.
1887. Die Runenschrift. Berlin.
1923-1904. De Danske Runemindesmerker, vols. 1-4. Kgbenhavn.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 4

Roebling Fund

PRECIPITATION AND TEMPERATURE
IN WASHINGTON, D. C.,
FOR 1950 AND 1951

BY
C. G. ABBOT

Research Associate, Smithsonian Institution

(Pusiication 4045)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MARCH 1, 1951

The Lord Galtimore Press

BALTIMORE, MD., U. 8 A.

|
|

Roebling Fund

PRECIPITATION AND TEMPERATURE IN
WASHINGTON, D. C., FOR 1950 AND 1951

By C. G. ABBOT
Research Associate, Smithsonian Institution

A. PRECIPITATION AT WASHINGTON

Referring to previous publications,’ I have predicted in January
of each year, for several years, about 155 dates throughout the
year when average daily precipitation in Washington, D. C., should
exceed that quantity for all other dates of the year. In 1950, for
the seventeenth consecutive year, this expectation was verified.

TABLE 1.—Statistics of Washington precipitation, 1950

(Values in inches)

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. Year

Average |) Pid. .... 0.054 0.095 0.151 0.088 0.296 0.013 0.182 0.223 0.399 0.094 0.186 0.142 0.153
er el other. 0.058 0.095 0.113 0.031 0.288 0.175 0.149 0.122 0.071 0.119 0.005 0.091 0.103
Other Merete, <\ 63 0:04. 0.99 1:35 2.80 1.03 0.07 1.22 1.83 5:62 0:79 35:09 1.57 1.49
Lo.) $e Te7d 2.08 (405. 1.87 §:906 3:14 ° 5.1 §.40 6.73 3°33 2.87 3:69 46.47
Normal ppt. ....... Bang) S027) 3°78 “3127 3:70 “Avg 4i7l: 4.05 “si24, 2.84 2.47) 3:aa) Aacx6
ercent of normal.. 49 82 108 57 fOr 76. §IO8! a2) 208) Try $2i. , Lit 110

Tabulations had indicated that the ratio for average daily precipitation,
preferred dates
all other dates

for 17 years is 147 percent, and for 1950 it was 149 percent.
Preferred days of 1950 had a higher average precipitation than all
other days in all months but January, February, June, and October.
In February the ratio was sensibly 1.00. In October, if the heavy
rainfall of October 23, which fell between midnight and early morn-
ing, had occurred 2 hours earlier, the ratio would have exceeded

, should be 142 percent. The average observed ratio

1 Especially Smithsonian Misc. Coll., vol. 104, Nos. 3 and 5, 1944; vol. 111,
No. 17, 1950.

Errata.—In the last-cited publication, table 1, Normal ppt., year, for “42.46”
read “42.16”: and in table 2, under Sept., for “26” read “27”; Oct., for “23”
read “24”; Nov., for “19” read “20”; Dec., for “16” read “17.” In footnote 3,
p. 2, for “949” read “1949.”

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 4

aR 1 ~ 195°

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116
unity. Had it occurred 6 hours earlier, the ratio for October would
have been about 11.5 instead of 0.70.

Table 2 gives the dates for 1951 when the average daily precipita-
tion in Washington is expected to exceed the average daily precip-
itation in this city on all other days. In the first column are given

TABLE 2.—Predicted dates when average daily precipitation should exceed
average daily precipitation for all other dates for the year 1951
in Washington, D. C.

“Preferred”

cycle places Jan. Feb. Mar. Apr. May June
Ti fh segs eet 24 20 19 15 12 8
LAGE pers ne ayy fc 25 21 20 16 13 9
1 Bt roa Recisaearee 26 22 2I 17 14 10
LG Beams cis 27 23 22 18 15 II
VS Toca eck, 1, 28 24 23 19 16 12
D210 (ilies eakna vee 8 4 3, 30 26 23 19
SOUT ee. aan 9 5 4, 31 27, 24 20
ERG ca nnd crc: II 7 6 2, 29 26 22
XSV ANG 8 Sorat hope 13 9 8 4 1, 28 24
XUN ATUT ee pte fete 14 10 9 5 2520) 25
EXONGTH cubensis 18 14 13 9 6 2, 29
XOXO asin reine 22 18 07; 13 10 6
2,8) a as eine 23 19 18 14 II a
“Preferred”
cycle places July Aug. Sept. Oct. Nov. Dec.
Dipiegs sat ctets ce 5 I, 28 24 21 17 14
DI sh eee as 6 2, 20 25 22 18 15
DTT Avreioenese 7 3, 30 26 23 19 16
Tig ise ictaes 8 4, 31 2 24 20 17
Viento ce 9 5 1, 28 25 21 18
DS) 0) RN Sek or 16 12 8 5 I, 28 25
SS TUT Be eh ees 17 13 9 6 2, 20 26
DO MREANG ceoytfonnne 19 15 II 8 4 1, 28
DON OINY Sat a ae te 2I 1) 13 10 6 3, 30
AVAIL eat 22 18 14 II 7 4, 31
d.2.9) Dig a Bae esse 26 22 18 15 II 8
POV IE reser 3, 30 26 22 19 15 12
EXOXGVEIIE FCP ete ei 4, 31 27 23 20 16 13

in Roman numerals the day numbers of the 27-day cycle when
higher precipitation is expected. These values arise from the sta-
tistical study, 1924 to 1941, cited below. The other columns give
the actual days in the 12 months of 1951 when these Roman cycle
dates will occur. In other words, the remaining columns give the
“preferred” dates for 1951. While it is expected that for the entire
year 1951 the “preferred” dates will yield higher average precipita-

NO. 4 PRECIPITATION, TEMPERATURE IN WASHINGTON—ABBOT 3

tion than all others, and even that this will be so for most of the
individual months of 1951, the probability that any individual “pre-
ferred” day will yield precipitation is scarcely above 50-50.

The basic tabulation, on which table 2 rests, began with January
1924 and ended with December 1941. The length deduced for the
precipitation cycle is 27.0074 days. In 366 cycles of this length there
are 9884.7084 days. From January 1, 1924, to December 31, 1950.
there are 9,862 days. Hence the 367th cycle begins January 24, 1951,
as given in Table 2.?

B. TEMPERATURE AT WASHINGTON

In previous papers * I have drawn attention to a regular periodic
variation of 6.6485 days’ length in the output of radiation from the
sun. Though quite regular intervals occur in the solar variation,
terrestrial responses thereto are sometimes I, 2, or rarely 3 days from
their expected dates. This is due to the complexity of the atmospheric
constituents and reactions. All terrestrial responses to solar impulses
are subject to lag. For instance, the warmest part of the day occurs
I to 6 hours after noon at various stations of the earth. The lag
is not constant from day to day at any station. Hence, from analogy,
the irregularity of terrestrial responses to the 6.6485-day solar varia-
tion is not surprising. Nevertheless they are notable in magnitude.
As shown in earlier papers they range from 2° to 20° F. in the
temperature of Washington.

Notwithstanding the differences in lag just referred to, which
cause displacements of the terrestrial responses, it seemed to me
worth while, in January 1948, to predict for the ensuing year the 55
dates when minima of temperature with respect to surrounding days
might be anticipated in Washington. In doing so I recognized that
actual minima would sometimes fall 1, 2, or even rarely 3 days from
the dates predicted. In January 1949 the prediction was compared
with the event with fair verification.

In table 3 I give the dates predicted for minima of Washington
temperatures in 1950, and the dates when minima occurred. I also
give the dates predicted for minima in 1951. In these cases the
predicted dates tabulated are those within a half day of the accurately
computed minima, based on a period of 6.6485 days, with the zero
date January 17.0000, 1946.

2T.e., January 1 +23 days= January 24.
8 Smithsonian Misc. Coll., vol. 107, No. 4, 1947; vol. 111, No. 6, 1949; vol.
111, No. 13, 1949.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

In order to show how definitely the period reveals itself in Wash-
ington temperatures, and recurs on the average on time, I give in
figure 1 the frequency of occurrence of minima within certain inter-
vals of time from the accurately computed minima, covering the
whole year of 1950.

TABLE 3.—Dates in 1950 when minima in Washington temperatures were
predicted and observed, and dates predicted in 1951

January February March April
1950 predicted .. 5 12 19 25 1 8 14 21 28 6 13 20 26 2 8 15 22 28
1950 observed ... 8 12 20 28 31 4 8 14 20 26 4 ‘OTA 19) 23) 26°30 6

9 14 21 25 30)

1951 predicted .. 6 13 19 26 2° 8 15 22 28 7 14 20 27 3 10 17 23 36
May June July August
1950 predicted .. 5 12 18 25 ay Tae 4 II 17 24 30 6 13 19 26
1950 observed ... 8 12 20 25 28 Bi r2) Tee2e2o Gel2o is) 2r.26 5 13, 22.27
1951 predicted .. 7 13 20 27 2 19) 36° 22) 20 651210) 25 r 8 14 20m
September October November December
1950 predicted .. 2 8 15 22 28 ie sachs AS L007 aer sy, 4 10 17 24 30
1950 observed ... 5 9 13 17 22 24 6 14 21 26 62 r8ier 26 6 13° 20 2%
1951 predicted .. 3 10 17 22 29 6 12 19 26 r 8 15 21 28 4 11 18 24 3k

FREQUENCY OF NMIIN/IMA

oO
& eas" a — ef
3 2 z 2 oe. +¥ +3 +2
DAYS
Fic. 1—Relative frequency with which temperature minima occurred in 1950
in Washington within certain days of predicted minima.

ABBOT

NO. 4 PRECIPITATION, TEMPERATURE IN WASHINGTON

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6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Again, to show how the observed minima seesaw around the pre-
dicted dates, I plot in figure 2 the actual daily departures from Wash-
ington mean temperatures, from October 5 to December 31, 1950,
along with accurately computed predicted minima. The first periods,
October 6 to 27, occur too late. Then the minima come several days
too early, from October 27 to November 18. There follows a time
of transition from November 18 to 30. New conditions prevail from
December I to the end of the year. In this time the minima arrive
about 3 days late.

These seesawing transitions lessen the value of the 6.6485-day
period for forecasting purposes, but of its veridity in Washington
temperatures there can be no doubt. Its accuracy is proved by the
agreement of average recurrences of 1950 with the computed times,
notwithstanding a lapse of five years, 1946-50.

ty: PAN

iv
du a

‘MIDDLE CAMBRIAN STRATIGRAPHY

~ ROCKY MOUNTAINS

(Wirn 34 PLATES)

BY

FRANCO RASETTI

The Johns Hopkins University
, Baltimore, Maryland

yt
een | f
"

‘ arn ate, I ri
Ne Saito” sep 181951

ppveciesbaon 4046) >» _ CARvE LIBE ant

On CITY OF WASHINGTON
LISHED BY THE SMITHSONIAN INSTITUTION
- SEPTEMBER 18, 1951

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 5

Charles D. and Mary Waux Walcott
Research Fund

MIDDLE CAMBRIAN STRATIGRAPHY
AND FAUNAS OF THE CANADIAN
ROCKY MOUNTAINS

(WiTH 34 Prates)

BY
FRANCO RASETTI

The Johns Hopkins University
Baltimore, Maryland

ays? re ows
Gung Y ot

eeceese?®

(Pustication 4046)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 18, 1951

The Lord Baltimore Press

BALTIMORE, MD., U. 8. A.

CONTENTS

PART I. STRATIGRAPHY

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WONSATE TEMES Lit TR CEM dts cys hoe iofevs sto kajol ave via = a'w vidas (im oko vale eee eee eee
ETT cine roe ce sata) Lao Ss. Lp gielvla poses aemtatelsle aie ane
Ee RGUMMIOVCTIRII AISI id 18 ects oso Sisto o ocd bin cccek ba Coca ee ome ea oon were
memecaiich > Micking Elotse Mine 2.5... cc0.dscrareveguscssapeeanes
REE OPT ECSS: OGATTY silt be os aus vee ds awe car eekeneete
EPEAT FOU LG UI ets ys iia k = oi ees eine seo Vine Mialdin wis miae ie a SAG ota
SEHR AGESCHIPEOM) a ahye ory csc claloe ole eala a ular t= arcane Cael seam eee

LA Taste ie Clas CB) aac) he tee ee ORES SEO ACR ACER sae CROC OMT CICS ocr
AGITE TH aig GT Se ee a IE RRM ey Sat, Gino
merit (Gilly And Mossil Golly i... 0c. cess. nvecosesnocesseasanes
Semapriati formations: Lower Cambrian ...:....2-.csccescsseccnsccucces
MEALS ATIC SELIG Ns ho ic oc eyeie ee vig. d winve ne nies 00.6 vl wieiais w gin ee ain Mera TREC
Bieger “potitinaty Of fOrMatiOn. <2... acer edercter vs ennenaunneeue
SU LOMISIMESt ONE “TLEMIDEL. yoo vs,» oack ois vies, vir wos .a wn sn ates aimee ee eee
Cambrian formations: Middle Cambrian ............+ecccseecesnseccaces
MEER INES COVIIATION c5 cc cc beecsrenvswenedonanaekevccdus ie yam
MIOCEIRE ac at ok dé od bn Ce ebie ss hevaveswiteansaiuay eeweed
PeOIOeY Aliel PUICKHESS, 6.6 boc cade en ewcndes sauvssvererasssuneas
Monn Whyte—Gathedral contact: . 0... sc.scccsccrscrecvensensess
Pane Agnes Shale lentil. .....0ccccscsveucsesavecvcssesereeuneuns
CMe SIAte SIGH ots cade eee e aval he Ver eked sev nse nase cae komn

iv SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Page
Cathedral formation: siaoe ccicha Ga crete Sesieie ois. ge settee one eo aasieettoterentocneeete 64
Tyme SOCtION s/s,é/<is\e'sisisie aia isvolsheerote svererstevateranscalel Sistas alevejavetore siohsvonstettrere 64
Lithology..and thickness..2...:.): 5. cee ccasceoe a oahie ae ae ao era 65
Cathedral—Steplien contact. 2... Jc). ccn-wvag sive mite ieeni eae eee 69
Ross) ake. shale-meéember. sino eee onsen). Seo ac cena 69
Stephen “fOrmation 5: sgisicinn.s s d.vwe cawcaree eng cine vie wares 2 aera eee 70
Type SOCHON: « <-cysisitisvateatatans die sera erele eave wietatbie 2p os sie arto ee ooo nee 70
Lithology ‘and* thickness). i060 «sz ain csc ss m= «attests ake eee TE
Stephen-Eldon: contact)... sie: s0scs.easd sees > sear sen alee Soe 74
Ogygapaus shale lentil cc.c50.3s¢r cae es oe eae estos on sete 75
Burgess. shale dental... :5..2. viyce So sies ais oiec eee ee ee 77
EVAUIIAS) Sree kd Svcd e che oars ee Ste teres alee wane sie RIOR ene oe eee 79
General istatemiceint-5)c-csatvo ss suctaie soe ois bse ee tala a lero bee eee ee 79
Raina of theebonma-Olenellusezonens sce tee ee ee sie eee 81
Hower viddles Gambian bouncanvereerainiecit eerie ates 84
Fauna of the Wenkchemnia-Stephenaspis Zone .......-+.eeeeeeeeeees 87
Fauna of the Plagwra-Kochaspis ZONe (an... sns 50 tee a eles aes seen go
Fauna ofthe Aibertella zone ic. <2 sea dea oe tes oo ao ke a eee 93
Fauna. of the Glossopleura’ Zone 01.5 04sec ness es ae eee ee 96
Fauna of the Bathyuriscus-Elrathina zone ........:2cccescccscsccees 99
Order of succession of the faunules, (2504) eecd >... 42's eee 108
Species fiot discussed: in this, pater. ove... se +s ssn enlsise eae ene I10
Geographic and vertical distribution of trilobites................0.00- 112
Environment ‘of deposition, : s....44000 04scses oes ee gee oxen eid ene 117
Hndex (Of localities. .iacis sates groves 4-00 re pce ato aseio OS eretoreec mia, eee re caer ea 121
PART II. PALEONTOLOGY
Generallistatemient: ¢orace atweels cies wgiave tins sisoeuierns cere eau EEE Eee 131
TRETINIMOLO SY auais sea ats aie tie arora ve etorel wie'o enerele caraies Sere on Eee eee 131
@lassitication ot. Cambrian trilobites: «le cnie asisieeiac ine clatter 132
Descriptions of genera and species of trilobites... .-...+- «i ccm «eerie ieee 133
Order Apnostida. 3s. Adhd Jc hore bene oe pees cee ae ee 133
Order Eadiscida. sa. ietiicjnriaeuric os vee a Si ies a eae ee 137
Order Opisthoparida) «<..1 sia as cisiace siete witw- 5 8 eae eee oe ae eee 138
Superfamily ‘Burlingioidae « ¢.:0.0.s00s.c0.n aad oawa ao oo 138
Superfamily Corynexochowdae.. ss sips o. ood ws eo see eee 139
Family Zacanthoididae’ dx saalensy get doscalteva tee 140
Family Dolichometopidad ©. <s%<:ccuitn<.neis/ bom de ete ae ae 156
Family ‘Gorynexcelidae” 4.6.0 ¢sc suman ets beaten nee 188
Bamily Dorypy gidae : os. 40510... caves mus hw ciel 188
Family Osygopsididae’ «.3ois.4 oiaavissdbe sae ac ae ee 190
Ramuly Oryetoceplalidae:\. is os scan soe paca ct eee 192
Superfamily “Ptychoparioidae no. eens c:cclen sec cse canes ae ee 198
BRETEEENICES.,,.\ 5 0 )a:nze are « arneraje ave tye bee eaters ove e-evstoo tie oes aus re ote teeter ear tenet 251
Explanation’ of plates. /./2: 045 Vs siete hove ates ate ee re 255

Index

SO SIAN WD H

Leal

Ah wh xm

LIST OF PLATES
(All plates following page 270.)

South face of Mount Temple.

Middle Cambrian strata on Mount Stephen and Park Mountain.
Northwest face of Mount Stephen.

Lower part of the northwest face of Mount Stephen.

Interfingering limestone and dolomite in the Cathedral formation.
Anomalous lithology of the Cathedral formation on Mount Stephen.
East face of Mount Odaray.

Trilobites of the Bonnia-Olenellus and W enkchemnia-Stephenaspis zones.
Trilobites of the Wenkchemnia-Stephenaspis zone.

Trilobites of the Wenkchemnia-Stephenaspis zone.

Trilobites of the Wenkchemnia-Stephenaspis zone.

Trilobites of the Wenkchemnia-Stephenaspis zone.

Trilobites of the Plagiura-Kochaspis zone.

Trilobites of the Plagiura-Kochaspis zone.

Trilobites of the Plagiura-Kochaspis zone.

Trilobites of the Plagiura-Kochaspis zone.

Trilobites of the Albertella zone.

Trilobites of the Albertella zone.

Trilobites of the Albertella zone.

Trilobites of the Albertella zone.

. Trilobites of the Albertella (?) zone.
. Trilobites of the Albertella (?) zone.
. Trilobites of the Glossopleura zone.

Trilobites of the Glossopleura zone.

Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis
shales).

Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis
shales).

Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis
shales).

Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis
shales).

Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis
shales).

. Trilobites of the Bathyuriscus-Elrathina zone (Burgess and Ogygopsis

shales).

. Trilobites of the Bathyuriscus-Elrathina zone.
. Trilobites of the Bathyuriscus-Elrathina zone.
. Trilobites of the Bathyuriscus-Elrathina zone.

Trilobites of the Bathyuriscus-Elrathina zone.

TEXT FIGURES

Map of the area indicating location of sections............eeeee cae eens 10
EU MUNINEE SCCHOI Gs cle.sic chives sinc cntevssvaensenvvchenewshy wenn ene 17

. Contour map of Mount Stephen indicating fossil localities............+- 40

Section in the eastern and western parts of Mount Stephen............. 44

. Map indicating thickness of the Mount Whyte formation............++- 61

Charles D. and lary Waux Walcott Research Fund

MIDDLE CAMBRIAN STRATIGRAPHY AND
FAUNAS OF THE CANADIAN
ROCKY MOUNTAINS

By FRANCO RASETTI
The Johns Hopkins University
Baltimore, Maryland

(WirH 34 PrLateEs)

BAR los SERATIGRSA EEL
INTRODUCTION
THE PROBLEM

The area described in this work is situated on or near the Conti-
nental Divide in the southern Canadian Rocky Mountains. The portion
of the Divide investigated centers around Kicking Horse Pass and
extends northwest to Bow Pass and south to Wenkchemna Pass. The
mountains forming the Divide are known as the Waputik and Bow
Ranges, respectively north and south of Kicking Horse Pass. All
the localities discussed are either in Banff National Park, Alberta,
or in Yoho National Park, British Columbia.

Stratigraphically this study covers the lower portion of the
Middle Cambrian, i.e., the Mount Whyte, Cathedral, and Stephen
formations. These strata average, in the area investigated, a thick-
ness of somewhat over 2,000 feet and represent about one-half of
the total thickness of the Middle Cambrian series. Immediately un-
derlying and overlying formations are occasionally discussed.

The area is famous for the great development, high fossil content,
and splendid exposures of the entire Cambrian system. The simple
fault-block structure of these mountains facilitates the study of the
stratigraphy. Hence the Canadian Rocky Mountains attracted the at-
tention of the foremost student of the Cambrian in North America,
Charles D. Walcott, who devoted more time and work to the Cam-
brian in this area than in any other portion of North America. He
established the general Cambrian section of the region, named the

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 5

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

formations, described a number of sections in detail, and made an
extensive study of the fossils. His discovery and study of a large
fauna of invertebrates preserved in exceptional manner in the Burgess
shale represents one of the greatest paleontologic contributions of all
time.

Walcott fully realized the provisional character of some of his
work, a consequence of the enormous geographic and stratigraphic
range of his studies, the rugged topography, and difficult communica-
tions. He often pointed out the necessity for more detailed investi-
gation and especially more intensive fossil collecting by future stu-
dents.

The choice of the particular problem discussed in this paper was
determined by the following considerations: The area was chosen for
reasons of accessibility and because it includes the type sections of
the formations that it appeared desirable to revise. The portion of
the stratigraphic column investigated, the lower half of the Middle
Cambrian, was chosen because of the poor knowledge of the faunas
of this age, and because the southern Canadian Rockies probably con-
tain one of the thickest, most fossiliferous, and better-exposed sec-
tions of this stratigraphic interval. Hence a study of the faunal suc-
cession in this area was expected to present much more than local
interest leading to the establishment of a standard sequence for the
entire Cordilleran province. The results rewarded these expectations,
as the writer believes that the fossils here described give for the first
time a clear idea of the succession of trilobite faunas that followed
the Olenellus fauna and preceded the Albertella fauna.

The Cambrian beds underlying those investigated (the Lower Cam-
brian sandstones) and those immediately overlying (the Eldon dolo-
mite) are but sparsely fossiliferous and are represented by more
fossiliferous equivalents in other areas of the Cordilleran province;
hence they presented less interest than the strata studied in this work.

ACKNOWLEDGMENTS

The present work was made possible by grant No. 509-47/48 from
the Penrose Bequest of the Geological Society of America. The grant
defrayed the writer’s and an assistant’s expenses for two field seasons.
The writer is glad to acknowledge his deep gratitude to the Society.

Thanks are also due to the Department of Mines and Resources,
Ottawa, for permitting the collection of fossils in the Canadian
National Parks; and to the officials of Banff and Yoho Parks for
their helpful cooperation.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 5

The writer is also greatly indebted to Dr. G. Arthur Cooper for
making available the collections and facilities of the United States
National Museum, for the identification of the brachiopods, and for
taking some of the photographs of specimens in the National Museum
collections.

Thanks are also due to Prof. Christian Poulsen, of the University
of Copenhagen, for information concerning the Cambrian faunas of
Greenland.

Finally, the writer wishes to thank F. Bonenfant and C. T. Moore
who assisted him in the field in 1947 and 1948, respectively.

SUMMARY OF PREVIOUS WORK

The work done on the Cambrian of the area prior to 1939 is
summarized very briefly, as excellent reports have been published by
Deiss (1939; 1940). Only the essential results of Walcott and Bur-
ling are mentioned.

Walcott, in a series of papers (Walcott, 1908a ; 1908c ; 1917a; 1927;
1928) laid the foundation of the Cambrian stratigraphy of the re-
gion. In several other publications he described many of the fossils.
The Cambrian formations established by Walcott are the following.

Lower Cambrian.—Coarse Lower Cambrian clastics were found
by Walcott to overlie unconformably late Proterozoic (Beltian) shale.
Walcott divided the Lower Cambrian into four formations. The
first three, in ascending order, are the Fort Mountain formation
(formerly known as Fairview formation), the Lake Louise shale,
and the St. Piran formation. The Fort Mountain formation consists
of sandstone and quartzite and was reported to be 940+ feet thick.
The Lake Louise shale is a siliceous shale 105 feet thick at the type
locality. The St. Piran formation consists mostly of sandstone and
quartzite with some siliceous shale and was reported to be 2,632 feet
thick at the type locality. Fossils, chiefly olenellid trilobites, are known
from several horizons through the St. Piran formation and indicate
an Early Cambrian age.

The uppermost Lower Cambrian unit was named the Mount Whyte
formation from Mount Whyte west of Lake Louise. The formation
consists of interstratified siliceous shale and crystalline or oolitic lime-
stone. Its thickness on Mount Whyte was given as 386 feet.

The fauna of the Mount Whyte formation was described by Wal-
cott (1917c). However, owing partly to the meager and rather poorly
preserved material available at the time, partly to the difficulty of
correlating different horizons among the various measured sections

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

of the formation, the character of the fauna of the Mount Whyte
formation has remained uncertain until the present time. The pres-
ence of Olenellidae in the basal beds of the formation induced Wal-
cott to consider the entire unit of Early Cambrian age.

Middle Cambrian.—Walcott’s original lowermost Middle Cambrian
unit was named the Cathedral formation (Walcott, 1908a) from
the intended type locality on Cathedral Mountain, and described
as “massive, arenaceous and dolomitic limestones.” Later (Walcott,
1928) he changed the type locality to Mount Bosworth, where he
gave a thickness of 1,212 feet. No fossils were reported from the
Cathedral formation.

In 1917 Walcott (1917a) added a new formation, the Ptarmigan,
chiefly in order to provide a name for the shale bearing the Albertella
fauna (the Ross Lake shale), which Walcott had discovered and
which for many years had been known only from drift blocks col-
lected at the base of Mount Bosworth. However, in defining the
Ptarmigan formation, he designated as typical a series of limestones
on Ptarmigan Peak where the Albertella-bearing shale does not oc-
cur. Furthermore, Burling (1916) had already found the Albertella-
bearing shale in place on Mount Bosworth and proved that its
position is within the Cathedral formation, 375 feet above the base.
Hence the Ptarmigan was essentially a name for the lower portion
of the beds previously included in the Cathedral formation.

The next Middle Cambrian unit was named (Walcott, 1908a;
1908c) the Stephen formation from the type locality on Mount
Stephen. The Stephen formation was reported to consist of 712 feet
of siliceous and calcareous shales and dark-gray, thin-bedded lime-
stones, and was stated to include at the top the extremely fossilifer-
ous shale bearing the Ogygopsis fauna. The Burgess shale, discovered
on Mount Field opposite Mount Stephen, was also considered as a
local development within the Stephen formation,

The uppermost Middle Cambrian unit was named the Eldon forma-
tion and the type section defined on Castle Mountain (now Mount
Eisenhower). The formation was described as 2,195 feet of “mas-
sive, arenaceous, dolomitic limestones, with a few bands of purer
bluish gray limestones.” A few fossils collected at various horizons
indicated a Medial Cambrian age.

Upper Cambrian.—Walcott tentatively placed the Middle-Upper
Cambrian boundary at the top of the Eldon formation, although the
next overlying units, the Arctomys and Bosworth formations, are
unfossiliferous and hence of uncertain age. These are succeeded by

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 5

the Paget and Sherbrooke formations. Fossils collected from both
indicate an early Late Cambrian (Dresbachian) age. Walcott re-
ported a total thickness of 3,590 feet for the Upper Cambrian series
on Mount Bosworth. Upper Cambrian deposits are not known to be
present south of Kicking Horse Pass, the summits of the Bow Range
being usually formed by Middle Cambrian rocks.

A paper was especially devoted by Walcott (1927) to a discussion
of the early Paleozoic sedimentary basins in the Canadian Rockies.
He distinguished the Beaverfoot, Goodsir, Bow, Glacier Lake, and
Sawback troughs. He observed that deposits of particular ages were
represented in some of these troughs but were absent from others.
The area studied in the present work is almost entirely located within
the Bow trough, involving only the northeastern marginal area of
the Goodsir trough. Walcott distinguished the Bow and Goodsir
troughs from the facies of the Upper Cambrian sediments, since
Lower and Middle Cambrian formations are not exposed in the area
of the Goodsir trough. It will be shown in this paper that the Middle
Cambrian formations along the line that Walcott considered as the
border between the Bow and Goodsir troughs already show remark-
able changes of facies from the normal sediments of the Bow trough.

No field work was done on the Cambrian of the area between 1917
and 1938. Discussion of Walcott’s stratigraphic and paleontologic
data by various authors (cited in Deiss, 1939) added little to what had
already been published by Walcott concerning the faunal succession
in the Mount Whyte, “Ptarmigan,” and Stephen formations.

Deiss undertook an accurate revision of several of Walcott’s sec-
tions. Unfortunately he did not supplement his two stratigraphic
papers (Deiss, 1939; 1940) with descriptions of the fossils. Deiss
restudied the Castle Mountain, Ptarmigan Peak, Ross Lake, Mount
Bosworth, and Mount Assiniboine sections (the last outside the area
considered in this paper). His conclusions are briefly summarized in
the following paragraphs.

(1) The Lower Cambrian clastic formations were accurately and
completely measured by Deiss (in the Ptarmigan Peak area). He
reported a thickness of 865 feet for the Fort Mountain sandstone
(including 60 feet of Lake Louise shale member at the top) and 815
feet for the St. Piran sandstone, a total of 1,680 feet. According to
Walcott, the aggregate thickness of these formations in the Bow
Range is 3,677+ feet; hence, even allowing for inaccuracies in the
measurements, it is clear that there is considerable lateral variation
in the thickness of the Lower Cambrian within short distances.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

In the Mount Assiniboine area, Deiss united all these Lower Cam-
brian clastics under one name, the Gog formation, reported to be
1,235 feet thick in that area. Deiss expressed the doubt, shared by
the writer, that the Lake Louise shale is more than a localized deposit ;
in the absence of this shale, there is nothing to distinguish the Fort
Mountain from the St. Piran sandstone. In this case the name Gog
formation could be applied everywhere to the entire series of Lower
Cambrian clastics. However, Walcott’s original names are still used
in this paper as the problem has not been sufficiently investigated.

(2) Four sections of the Mount Whyte formation were measured
by Deiss, at Castle Mountain, Ptarmigan Peak, Ross Lake, and Mount
Bosworth, yielding thicknesses of 145, 275, 315, and 285 feet, re-
spectively. Although he did not measure the Mount Whyte section,
he transferred the type section of the Mount Whyte formation to
Ptarmigan Peak (an action that the writer considers unwarranted).
Deiss found that olenellid trilobites are confined to a few basal feet
of the formation instead of ranging through the entire unit as had
been reported by Walcott. The rest of the formation yields faunules
assigned by Deiss to a “Plagiura zone,’ which, for no clearly stated
reasons, Deiss still included in the Lower Cambrian.

In the Mount Assiniboine area, Deiss applied a new name, Naiset
formation, to 475 feet of shale and limestone between the Gog forma-
tion and the Cathedral dolomite.

(3) Deiss recognized the Ptarmigan limestone, in which he in-
cluded 460 feet of strata on Ptarmigan Peak, 105 feet on Castle
Mountain, and tentatively 195 feet on Mount Bosworth. He could
not find any strata assignable to the Ptarmigan in the Ross Lake
section. The Ptarmigan limestone was considered to be a basal Mid-
dle Cambrian deposit chiefly localized in the Slate Mountains area.
In his later paper, Deiss (1940, p. 777) expressed doubts as to the
validity of the Ptarmigan as an independent formation and suggested
that it might be included in the Cathedral. Deiss saw evidence for
the Medial Cambrian age of the Ptarmigan in the presence of a
Kochaspis cecinna fauna, which he considers sharply distinct from the
underlying Plagiura fauna.

(4) The Cathedral formation (styled by Deiss the Cathedral
dolomite) was measured in three sections, yielding thicknesses of 670
feet on Castle Mountain, 620 feet on Ptarmigan Peak, and 1,085+
feet in the type section on Mount Bosworth. Deiss confirmed Bur-
ling’s finding that the Ross Lake shale member, containing the Al-
bertella fauna, occurs within the formation, but did not find this shale
in the Ptarmigan Peak section.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI Ff

(5) The Stephen formation was measured by Deiss on Castle
Mountain, Ptarmigan Peak, and Mount Bosworth, yielding thick-
nesses of 285, 315, and 470 feet, respectively. In all these sections
the Stephen formation yielded a Glossopleura faunule from the basal
beds, and several distinct faunules in the upper portion. However,
nowhere did Deiss find any strata analogous lithologically and faunally
to the Ogygopsis and Burgess shales. No redefinition of the Stephen
formation was attempted because Walcott’s type section on Mount
Stephen was not remeasured.

(6) The Eldon formation (styled by Deiss the Eldon dolomite)
was measured on Castle Mountain, Ptarmigan Peak, and Mount Bos-
worth, the reported thicknesses being 1,015, 1,230, and 1,110 feet,
respectively. Part of the decrease in thickness as compared with
Walcott’s data is due to Deiss’ setting off the upper part of Walcott’s
Eldon formation as a separate unit, the Pika limestone.

(7) The Pika limestone was defined on Ptarmigan Peak and
stated to be 460 feet thick on Ptarmigan Peak, 550 feet on Castle
Mountain, and 845 feet on Mount Bosworth. The formation yielded
“a new basal Upper Cambrian fauna” and was therefore considered
Upper Cambrian.

(8) Deiss also discussed the Upper Cambrian Arctomys and Bos-
worth formations in some of his sections. He affirmed that Walcott
had greatly overestimated the total thickness of the Upper Cambrian
series in the Mount Bosworth section.

METHOD OF WORK

The area investigated is the most easily accessible in the Canadian
Rocky Mountains. The highway Banff—Lake Louise—Kicking Horse
Pass—Field, the Banff—Jasper highway, and the side roads to Mount
Temple Lodge, Moraine Lake, and Yoho Valley allow one to drive
within easy reach of many of the sections. Many other localities are
accessible by trail from points not situated on the roads, but where
accommodation is available, such as Skoki Lodge in the Slate Moun-
tains and Lake O’Hara Lodge in the valley of Cataract Brook. These
facilities enable one to reach within a day’s walk or climb most of
the important sections in the area. Even when the day’s work involved
a climb of 5,000 feet and a walk of 10 or 15 miles it was found
practical and time-saving to do this from a convenient base where
accommodation was available instead of camping near the place to
be studied. Owing to the unstable mountain weather, it was found
that expeditions of the type that had to be planned in advance were

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

usually doomed to failure, whereas great mobility and the freedom
to decide instantly the location of the day’s work according to weather
conditions, made for good utilization of the available time. The
writer’s long experience of mountain climbing in the Alps proved a
valuable asset. Four months were spent in the field.

The mountains are of the fault-block type, dips seldom exceeding
15°. Often each mountain forms a single block separated from
neighboring mountains by normal faults, as valleys and gullies were
prevalently eroded along fault lines. This structure makes it possible
to find localities where the strata to be investigated are exposed at
the most favorable altitudes, between 7,000 and 10,000 feet. Most
of Walcott’s work was done near the valley bottoms, where the strata
are partially covered by glacial drift, talus, soil, and vegetation. The
writer’s sections were measured mostly above timber line. The section
traverse usually followed either a ridge, generally free from talus
cover even on moderately steep slopes, or a gully bottom, which can
be climbed more easily than an equally steep open slope, and where
the strata often offer fine exposures in the cliffs on either side of
the gully.

Most of the sections were measured on slopes between 30° and 45°
with respect to the horizontal, the dip of the beds not exceeding 20°.
Under these circumstances the following method allowed a fast, if
not extremely accurate, measurement of the thickness. Once the
direction of the traverse on the mountain slope or ridge had been
selected, the level of a Brunton compass was adjusted to the inclina-
tion of the strata in that direction. The angle of inclination could be
anywhere between zero (when the traverse followed the strike) and
the angle of dip. A point sighted on the slope is then known to be
approximately higher stratigraphically than the point where the ob-
server is standing by a constant distance (in the writer’s case 5/3”).
The observer then moves to the sighted point and repeats the opera-
tion. If the inclination of the beds is appreciable, the value found
must be corrected by multiplying by the cosine of the angle that the
line of sight forms with the horizontal. This correction never ex-
ceeded 10 percent. The writer was able to measure almost 1,000 feet
of horizontal strata in one hour on a conveniently steep slope. The
measurement obviously must be effected when climbing the mountain.
Reference marks were written on the rocks, permitting a ready place-
ment in the section of the fossils that were searched for and collected
in the descent. In certain cases thin units were measured directly
with a tape.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 9

Sections are not described in excessive detail, although sometimes
finer subdivisions were distinguished when measuring the section
than are reported here. Observation shows that great detail is of little
or no significance. Minor units in a given section traverse that can
be distinguished by slight differences in thickness of the beds or other
lithologic characters are often lenticular, or no longer recognizable
whenever change in slope or other conditions of exposure alter the
appearance of the weathered rock.

Topographic maps of the region are issued by the Surveys and
Engineering Branch of the Department of Mines and Resources, and
the following sheets are available: Lake Louise and Yoho Park
sheets, scale I inch to 1 mile; Kootenay Park sheet, scale 1 inch to
2 miles ; Banff Park sheet, scale 1 inch to 3 miles. The last, not very
accurate map is the only one that covers the localities investigated
in the upper Bow Valley.

DESCRIPTION OF LOCALITIES AND SECTIONS
TERMINOLOGY

Each section was assigned a number. The fossiliferous horizons
in each section are designated by a capital letter indicating the forma-
tion (P=St. Piran, W=Mount Whyte, C=Cathedral, S=Stephen,
E=Eldon) followed by the number of the section. The last symbol
is a lower-case letter indicating a particular horizon within the forma-
tion. Successive letters indicate progressively higher horizons in the
same section, and as far as possible the same letter was used to desig-
nate equivalent horizons in the different sections. For example, the
letters Cm designate the Ross Lake shale member of the Cathedral
formation, bearing the Albertella bosworthi faunule. The symbol
C3m indicates this horizon on Mount Whyte, C4m at Ross Lake,
Cs5m on Mount Bosworth, etc. An exception in the order of the
letters is represented by the horizons designated as Wh, Wi, Wj, and
Wk in the Fossil Gully section on Mount Stephen. When these beds
were first discovered in 1947, their position in the general section
was unknown and they were given these symbols. In 1948 it was
found that the highest of these horizons, Wk, is approximately equiv-
alent to the horizon designated as Wb in the Mount Whyte and other
sections, but too many fossils had been labeled by that time to make
a change to a more rational terminology worth while. It must also
be noted that the horizons designated as Ch, Cj, and Cj’, and Ck in the
equivalent of a portion of the Cathedral formation on Mount Stephen
may be younger and not older than the Ross Lake shale as the sym-

VOL. I16

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI Ea

bols would imply. Finally, the fossil horizons in the Ogygopsis and
Burgess shales were given the symbols Sd, Se, Sf, Sg, indicating
their probable relative positions with respect to each other and to
the other fossiliferous horizons in the Stephen formation, but this
order is inferred from paleontologic evidence and is not absolutely
certain.

All the fossils collected and prepared, over 4,000 specimens, were
labeled according to these locality and horizon symbols. A list of
all the fossil localities, including topographic and stratigraphic po-
sition and lithology, is given at the end of the stratigraphic part of
this paper.

Figure I represents a diagrammatic map of the area, with the
localities indicated by numbers in circles. The localities at Bow Lake,
Hector Creek, and in the Skoki Valley are not comprised in the map,
lying north of the area represented. Mount Stephen is illustrated by
a special large-scale contour map (fig. 3).

In the fossil lists included with the descriptions of the sections, the
relative abundance of the species is indicated by the abbreviations
cc=very common, c=common, r=rare, rr=very rare. An asterisk
preceding the specific name indicates that the locality is typical for
the species.

BOW LAKE

The Cambrian formations are excellently exposed in the eastern
slopes of the Waputik Range on the west side of the Bow Valley,
from Mount Bosworth to Bow Pass. The slopes of the northeastern
spur of Mount Thompson, between Bow Lake and Bow Pass, offer
the most accessible of these sections. Because of the gentle south-
ward dip of the beds, a very thick section is available, extending from
the base of the Cambrian in the cliffs overlooking Peyto Lake through
the entire Lower and Middle Cambrian and well into the Upper
Cambrian.

A section including the upper portion of the St. Piran sandstone
and the entire Mount Whyte and Cathedral formations was measured
in the slope and ridge just south of a glacial cirque facing the Bow
Valley, 1.0 mile northwest of Num-Ti-Jah Lodge on the north shore
of Bow Lake. Walcott (1928) had measured a portion of this section.

BOW LAKE SECTION (NO. 15)
Mipp_E CAMBRIAN

Stephen formation

Note.—The formation is incompletely represented in the fault
block where the section was measured. The lower portion of the

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

Stephen is an argillaceous, dark-gray, thin-bedded limestone, with
Glossopleura pygidia near the base, and forms a sharp contact with
the underlying dolomite of the Cathedral formation.

Cathedral formation

9. Dolomite: crystalline, light gray, tan weathering, in irregular thick
DOUS! iaire a ic: sis'o.0 igs pinio-Sialeig oiaiaratern/ oie ala gete ete olareeiale nal aptereetere eee
8. Limestone: black-gray, subcrystalline, in irregular beds 1-4 inches
thick s/55.0 i.e eS V eR doe aslette ral i acate ales able ee neta eine
7, Dolomites, as: tinit: Oss casa anion'sa'y « aatsie sine bac eeelep ees cL Ree
6. Limestone: subcrystalline, black-gray, with tan clay partings and
flakes, in irregular beds 1-3 inches thick. Fossils well pre-
served and fairly common in the thicker beds (locality Cr5n) .

FossILs:

*Albertelia lunbataRasettt 22 ia dss adie okie cele een Boe c
* istenorNachts) IRASEELL ara celeiers ats cletie Sets nere rete r
HIS OGHING -MACTOPS: RASCH A: ish Sv ee ae sete Bete r
*Ptarmiganoides bowensis Rasetti...........ceeceeeees c

Cathedral formation (Ross Lake shale member)

5. Shale: siliceous, fine-grained, blue-gray, fissile, with one or two
beds of gray, crystalline limestone 1-2 inches thick at the top.
Fossils abundant both in shale and limestone (locality C15m) .

FossILs (in shale) :

Albertcila bosworiht Walcott, . <5 ci. as esis ace site emer Cc

MACTOPS TRASCUUE 0's ead d.dtis's:s,00.00a)satals avers crete ete r

MALSEG TRESSREN S, Newsbyeakaie alata sd tities tena r

Koching americana (Walcott)... sss. 5<~n vcs + aoe rr

Mexsceltia. stator (Wealeott ic ac sane ne aan ae ora ante eeiet c

Ptarmigania rossensis (Walcott) ......0c0ccucsecstece rr

Vanuzemella mortia: Walcott: s2sab sie od Walkie caisson ties cc
Fossits (in limestone) :

PAT ber telladecuois WRASCHA ccic.cc shicsiwje train sca's pes waht see r

WAECEGPS CAS CU oso ed o!e/ hetece (20 aioe tases ial wR IOS r

WEHGTE IRCSSED! Wn ite neaetsns ve nere ohne ter r

Mesxicélla' stator: CW aleott) oc... ost eeeee ened Sones Cc

Vanuxemella noritia NVAlCOtt. 6. sc oa soe sence ee seen cc

Cathedral formation (continued)

4. Limestone: subcrystalline, dark gray, in irregular beds 1-4 inches
ELON, 9 ce isp sein ess) 0:53p-208 Gis © oh ate winnie Torso wie ingle es Peoaea adhe arene
3. Limestone: finely crystalline, dark or medium gray, mostly thick-
bedded, weathering to rough surfaces; partly grading laterally
to crystalline; light-cray dolomite... 2.24 sccciseesedee een
2. Limestone: subcrystalline, dark gray, in irregular beds 1-4 inches
PRICIE «cos ab accaereadecge e's eevee leis iis erajerein Oa bacetelelcicle See
1. Limestone: subcrystalline, medium gray, weathering light gray, in
irregular, mostly thick beds, up to several feet in thickness....

Total thickness of Cathedral formation. .................-

116

Feet
355

65
210

16

40

155
135

260

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 13

Mount Whyte formation

Feet
3. Limestone: subcrystalline, dark gray, with tan clay partings and
flakes, in irregular beds mostly 1-2 inches thick.............. 78
2. Shale and limestone: alternating layers of siliceous, gray shale and
Subenystalline se craywlimestoniesssn osiace certs tere ee ee cree tree 285
1. Shale and sandstone: gray and greenish siliceous shale alternating
with thin and thick, greenish sandstone beds in lower third.... 140
Total thickness of Mount Whyte formation................ 503
Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member: type locality)
4. Limestone: crystalline or oolitic, partly sandy, gray, in thick, ir-
regular beds. Partings poorly developed, giving the limestone
a massive aspect in cliff exposures. Olenellid fragments and
Bonnia common 130 feet above base. A fault cutting this unit
may cause a small error in the measurement of the thickness. . 180
3. Shale and limestone: siliceous, fine-grained, blue-gray shale alter-
nating with thick-bedded, gray, sandy limestone.............. 28
2. Limestone: crystalline or oolitic, gray, partly sandy, in thick, ir-
| STEATLES oc |S EER Me GRAB i SAU i wag Be 30
1. Sandstone: gray, more or less calcareous especially in upper part.. 40
Total thickness of Peyto limestone member...............- 278
St. Piran sandstone
i sandstone: purple, very. thick-beddeéd... .. «25.00 ssawemaded Not measured

Note.—Below about 100 feet of this purple sandstone is a siliceous
shale yielding rare olenellid shields.

HECTOR CREEK

A section of the Mount Whyte formation was measured on the
northeast spur of Mount Hector, in cliffs facing the Bow Valley just
south of Hector Creek. The section traverse is approximately located
0.8 mile northeast of the lower end of Hector Lake. The cliffs were
climbed following the bottom of a gully offering excellent exposures.
The summit of the ridge is formed by the top unit of the Mount
Whyte formation. The strata dip 23° northeast.

The Peyto limestone member of the St. Piran sandstone attains
here almost as great a thickness as at Bow Lake.

HECTOR CREEK SECTION (NO. 19)
MippLtE CAMBRIAN

Note.—The Cathedral formation consists as elsewhere of alternat-
ing and often interfingering limestone and dolomite. The lowermost
portion is a gray, very thick-bedded limestone.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Mount Whyte formation

F

4. Limestone: finely crystalline, dark gray, in irregular beds 1-4 a

inches thick, weathering to rough surfaces. Thickness ap-

proximate because dip slope makes measurement difficult..... 30
3. Limestone: crystalline, dark gray, with tan clay partings and

flakes, in irregular beds 1-6 inches thick................0e00- 125
2. Shale and limestone: mostly alternating thin layers of siliceous

shale and limestone, but with intervals of almost pure, gray

siliceous shale and others of crystalline or oolitic, dark-gray

limestone in beds 1-6 inches thick. Trilobite fragments com-

mon in limestone in middle of unit (locality Wioc).......... 175

FossIts :

Amecephalus sp.

Plagiura cercops (Walcott)
1. Shale and sandstone: siliceous, gray and greenish shale alternating

with beds of greenish sandstone, more abundant in lower part. 150

Total thickness of Mount Whyte formation...............- 480

Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)

1. Limestone: crystalline or oolitic, partly sandy, gray, in thick, irreg-
ular beds; interstratified with a little green-gray siliceous
shale. Olenellid fragments abundant through most of the
limestone, forming a coquina in certain layers. Thickness
estimated Wie 2..s.ah sists rcwleaseratavs Sera o cei eri naieke SRL eer 200
Note.—The Peyto limestone is underlain by purple and gray,
thick-bedded sandstone.

SLATE MOUNTAINS

The Slate Mountains include the classic Ptarmigan Peak section
first measured by Walcott and later revised by Deiss (1939). The
lower portion of this section (including the Mount Whyte and
“Ptarmigan” formations) was examined by the writer but no new
measurements were effected and the fossil collecting yielded poor
results. The conclusions reached by the writer are discussed in the
descriptions of the Mount Whyte and Cathedral formations and
need not be mentioned here.

The study of the lower Middle Cambrian formations was extended
to the northern part of the Slate Mountains, north of Mount Richard-
son and Pika Peak. The Skoki Valley descends from the northern
slopes of these mountains, and its upper cirque is occupied by a lake
lying about 1 mile north of the summit of Pika Peak. The only avail-
able map of this area is so inaccurate that the writer was unable to
decide whether the lake designated as Merlin Lake is this lake, or
another lake farther up in the valley. Below the lake, steep cliffs are

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 15

formed by the massive limestones of the lower Cathedral formation,
and below these the Mount Whyte formation is excellently exposed.
All these strata present much the same thickness and lithology as in
the Ptarmigan Peak section measured by Deiss.

The lower, calcareous portion of the Cathedral formation was not
measured, but a rough estimate indicates a thickness of the order of
400 feet. This is succeeded by dolomite, and a portion of the dolomitic
Cathedral formation was measured in order to locate the position
of a fossiliferous horizon. The section was measured in the slope
and cliffs southeast of the above-mentioned lake, approximately
1.0 mile N. 15° W. of the summit of Pika Peak. The beds dip about
15° south,

SKOKI VALLEY SECTION (NO. 13)
MrippLE CAMBRIAN

Cathedral formation

et
5. Dolomite: crystalline, light gray, tan weathering, thick-bedded. ki
Forms cliffs and extends for several hundred feet to base of
Stephene Olina Ole se aeric oma cio cevrterse wisrmereretin cael Not measured
4. Limestone: fine-grained, black-gray, in beds 2-4 inches thick.
Grades laterally into dolomite. Fossil locality C13r......... 5
FossILs :
*Glossopleura merlinensis Rasetti........cccceeccsccees c
BEM e erate PARAS ORR RIES wo ROUTE ae eee r
* SRORIWHSIS RASCLE” doce cen ceve sen manc es c
SPL E ASE AIDING! Syste armies © siete 5S Wd « ono ator ees atalalw antes) Oi 8 Sawin wie ate 13
2. Limestone: crystalline or fine-grained, black-gray, in beds 1-5
ATIGHES HEIGL yet k crels tale ete ha ate cid Walt, o'sie, clelel batersmerateterel Sen a ore 32
MCIOMNLSS (AS VRE) Gy wie bk otnceavew beeuw seebise ewe ee wndicde Kael 225
Total measured thickness of Cathedral formation.......... 275

MOUNT NIBLOCK

The Mount Whyte formation is well exposed in the cirque be-
tween Mount Niblock and Mount Whyte, above Lake Agnes. This
is obviously the locality where Walcott measured the upper part of
his “Lakes Louise and Agnes section” (Walcott, 1928, p. 302).

The best and most accessible exposure of the Mount Whyte forma-
tion was found to occur on the south slope of the east ridge of Mount
Niblock, and the section measured here is referred to as the Mount
Niblock section. The beds have a slight westward dip. The top of
the Mount Whyte formation is exposed at the approximate altitude
of 8,250 feet.

The lower portion of the Cathedral formation was examined and
found to consist of the usual alternating dolomite and dark-gray

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 116

limestone. The Ross Lake shale occurs approximately 400 feet above
the base of the formation, but appears thinner than in the neighbor-

ing sections and no fossils were observed.

The Cathedral-Mount Whyte contact is better defined than in most
of the other sections, as an increase in thickness of the beds coincides

with the transition from limestone to dolomite.

MOUNT NIBLOCK SECTION (NO. 2)
MippL_e CAMBRIAN

Mount Whyte formation

9. Limestone: crystalline or oolitic, dark gray, in thin, irregular beds,
with some thicker beds in middle of unit....................
8. Shale and limestone: alternating layers of siliceous, gray, tan-
weathering shale and fine-grained, oolitic, or crystalline gray

MteMEStOME ces) ist aisictad ovate terate sia aeeare ones exe tracey wie ate rate Meee teh eter
7, Limestone: crystalline or oolitic, dark gray, in thin, irregular beds
With) tanjclay pattings and Hakes. os ces recone mania
6. Shale and limestone: as unit 8, with shale predominating.........
5. Shale and sandstone: siliceous, gray shale with thin layers of
greenish sandstone, all weathering tan.............--...«..
4. Shale and dolomite: alternating thin layers of fine-grained, greenish
dolomite and siliceous, gray, tan-weathering shale..........

3. Shale: siliceous, fissile, slightly micaceous, greenish or tan, mostly
tan weathering. Lower part of interval forms bench and is
SUA y COM ELEM occre ois 010.510) 010) e\cyeie) afexgistelnhttete meters aot ole aioe eer

Mount Whyte formation (Lake Agnes shale lentil)

2. Shale: siliceous, fissile, very fine-grained, blue-green-gray (locality

W2Dius cere tc aecln nagidaie sae peeled pees Rete bo eee ae Senet
FOssILs :

Amecephalus agnesensis, (Walcott)............... fcke

Poltella: prima: CWaleott) isd since estas. nies « shh tmadele r

Syspacephalus perola (Walcott) ..........eeeseeeeeee c

Wenkchemnia walcotti Rasetti...........cccccecceee r

Mount Whyte formation (continued)

1. Shale and limestone: siliceous, gray shale with interstratified
nodular beds of gray limestone; weathers to pitted surfaces
after’ solution .of: the, Limestone... <0: < cise sce d «yp oo lel ole

Total thickness of Mount Whyte formation..............

Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)

5. Limestone: crystalline, sandy, medium gray, limonitic, in thick,
irregular beds; including some calcareous sandstone in lower
part. Olenellid fragments common throughout.............

Feet

56

45

13
22

29

44

52

15

281

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI i

St. Piran sandstone

Feet

Bersuale: siliceous, fissile,’ preen=Pray #2 oes <.. «Mele dbs RRS ee oak 2

3. Sandstone: calcareous, light gray, tan weathering.............. I
2. Shale and sandstone: micaceous, green-gray shale with thin inter-

Stee SADOSEANE: IDEOS. a:Wiere ain ane nak «sue peuneewee won oF = 10

1. Sandstone: thick-bedded, gray and purple................ Not measured

Total measured thickness of St. Piran sandstone...... 5 Gals 4!

MOUNT WHYTE—PLAIN OF SIX GLACIERS

An excellent section ranging from the uppermost Lower Cambrian
to the lower portion of the Eldon dolomite is exposed and entirely

LOWER MIDDLE CAMBRIAN
CAMBRIAN
ST. PIRAN CATHEDRAL
Smeal =
SS
Se
SS
SS
s
Ny
q 2 NS
DD DvD > a yD
oo — ~ ss
ss § s 2S
Ss 5 s 32
zones S a
ee 2 g5
aes $ ae
> 7s
“
WN ST lero EE eee eee
0 500 1000

FEET

Fic. 2—The Mount Whyte section. This section is typical for the Bow Range.
The position of the faunules is indicated.

accessible on the south ridge of Mount Whyte, from Plain of Six
Glaciers to near the summit of the mountain. The Stephen and
Cathedral formations were measured following the crest of this ridge.
The Mount Whyte formation, because of better exposure, was meas-
ured on the southeast slope of Popes Peak, a few hundred feet west
of the lowermost measured exposure of the Cathedral formation, in
a small gully 0.2 mile west of Plain of Six Glaciers Chalet. The two
exposures are separated by a normal fault, along which the deep valley
separating the slopes of Popes Peak from those of Mount Whyte
was eroded. The strata on the side of Popes Peak (west side) are
downthrown about 250 feet. This discontinuity in the measured sec-

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

tion does not introduce confusion because the Cathedral-Mount Whyte
contact is clearly defined. The entire section is referred to as the
Mount Whyte section. All these strata are approximately horizontal.
The Eldon-Stephen contact is exposed at an approximate altitude of
9,300 feet on the south ridge of Mount Whyte. The base of the
Mount Whyte formation, in the fault block where it was measured,
is exposed at an approximate altitude of 6,800 feet.

The Eldon dolomite becomes gradually thinner-bedded near the
base and interfingers with limestone, thus making the Eldon—Stephen
contact arbitrary within 50-100 feet (see discussion of Stephen—Eldon
contact).

MOUNT WHYTE SECTION (NO. 3)
MippLeE CAMBRIAN

Stephen formation

Feet

4. Limestone: subcrystalline, dark gray, in beds 2-4 inches thick;

in upper portion interfingering with dolomite, thus making the
boundary with the overlying Eldon dolomite highly arbitrary. 80

3. Limestone: subcrystalline, black-gray, in beds 1-3 inches thick;

in lower third thinner-bedded, with tan clay partings and
MAKES s,s BG stele Sain. dial Shera a vorevere aieneie crear tip ess tome reve strators eterna 130

2. Shale: mostly argillaceous, green-gray, with some interstratified

fine-grained, argillaceous, gray, thin-bedded limestone. Near
basevdarker, harder) siliceous shaleta. ccc ciscle sitet es oir 160

1. Limestone: near top fine-grained, banded; underlying beds finely

crystalline, dark gray, 2-4 inches thick; lower third argil-

laceous, fine-grained, in beds 1-3 inches thick. Fossils abun-

dant in finely crystalline and fine-grained argillaceous beds

FO-00) feet above base! (locality S3b)) access eerie cerletrtee 125
FossILs:
Glossopleura mekeet Resser.. 0s... ccc csscesccccccsecs c
a SHENOTHACHES TRASELLT, sive. sivetsieies oie siete i;
Polypleuraspis msignis Rasetths. : oc 0s. .sce0s «02 ween mane Cc
Total thickness of Stephen formation................e.08. 495

Cathedral formation

8. Dolomite: crystalline, light gray, weathering pale tan, thick-

bedded ; in upper part interfingering with limestone.......... 385
7. Limestone: finely crystalline, dark gray, in irregular thin and thick

beds. Girvanellia in some of the beds. ..........00.c000cee00s 145
6. Limestone: subcrystalline, gray, very thick-bedded.............. 63

5. Limestone: subcrystalline, dark gray, thick-bedded except near
top; weathering to coarse surfaces because of less soluble in-
CUSTOMS. “ierctasave sys see orosese hararecore tee tetaiefeutiets ok cre Soe cTCIReTe rere teereitoes 108

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

Cathedral formation (Ross Lake shale member)

4. Shale: siliceous, hard, fissile, fine-grained, dark blue-gray, highly

PeSREMEOE CS! (LOCALITY (Cath). cc ws ces enc ata cme de on ce eae
FossILs:

wiverteiaposwortht Walcott: oo 0. c....c0cearsaceesess cc
MEF ODS WRASETEN, 5 o's'u, cane eae cee eee teem r
PATA UREGSEL” were oat cae tone pene tone ee neat c

Kockina americana (Walcott). .......csccsdecsecees r

HIezIcerr Sturor” CVV AICOEE) sos 5 oS eos eeaweclencnesunte cc

Manuremecia norta. Walcott: 520i olci.s oieace hued sck cc

Cathedral formation (continued)

3. Dolomite: crystalline, light gray, tan weathering, thick-bedded;
interfingering with subcrystalline, dark-gray limestone.......
2. Limestone: subcrystalline, medium gray, very massive; with less
soluble inclusions weathering in relief...................005-
1. Dolomite: crystalline, light gray, tan weathering, thick-bedded;
with lenses of undolomitized dark-gray limestone............

Motalethickness) of Cathedral formation. .o...00. ova. cere

Mount Whyte formation (type section)

9. Limestone: finely crystalline or oolitic, dark gray, with tan clay
partings and flakes; in irregular beds 1-6 inches thick. Con-
tact with overlying Cathedral formation well marked........

8. Shale and limestone: thin alternating layers of gray, subcrystalline

limestone and gray, siliceous shale... i... ccisc cesses cecesene

TRESTIESEOME SMA SATIN Ea Oieyeiaio.s ahcraiesas oxvetatse) oye atts eletetetommieterene ore iotere eietore

Shialevandalimestone saSeutit Gy «10 siecle tists teenie masieiie cimeieiner

5. Limestone: as unit 9. Contains fragments of trilobites (locality
DM eae EN uchiha cdl «Aud cua. 0c ant Saaracd a prela She vi aaeeane ited Rak

FossiILs:
Amecephalus cleora (Walcott)
Plagiura cercops (Walcott)
4. Shale and limestone: as unit 8, mostly shale in lower third.......

Ove

Mount Whyte formation (Lake Agnes shale lentil: type locality)

3. Shale: siliceous, very fine-grained, fissile, green-blue-gray, highly

TeSEueeroUs (locality AWS) s xs'sida nike sdk oer hte bon wa Regs
FossILs:

AEP OLSE oad Soria auld voit aw doh awns ARN Wig Capua ears c

Amecephalus agnesensis (Walcott) ..........0eeee eens c

Ponelia prima CWacnt) « . << Senveinhes even ieee :

Syspacephalus perola (Walcott) .......scceeeeeeeeees c

*Wenkchemnia walcotti Rasetti.......cccsccsceccccecs c

Feet

120
48

205

1,169

45

20
Io
24

I2

73

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Mount Whyte formation (continued)

Feet
2. Shale and limestone: siliceous, gray shale with thin, nodular beds
of fine-grained, gray limestone. A limestone nodule yielded
fragments of AmecepHnalus..: «x ssa vcs.e sisson ac demaneen aeion 6
1. Shale: siliceous, fine-grained near base, coarser in upper part,
STEEN SLAY... Lteiahecsheie, ava-5 seeisie se cielu le Sai etamier vate nal cloeete emt eee 16
Total thickness of Mount Whyte formation............... 211
Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)
1. Calcareous, limonitic sandstone, grading into sandy limestone near
the top; the more calcareous layers filled with Bonnia and
olenellid fragments. Underlain by brown and purple, thick-
bedded "sandstone “iene c ent oarnaceiwieceesaettee wetecie te ome 20

ROSS LAKE

A section of the Mount Whyte and lower Cathedral formations
in the cliffs southeast of Ross Lake was measured and reported by
Deiss (1939, pp. 988-993). The section was not measured by the
writer, but at least the major units of Deiss’ section could be easily
identified in the field. A summary of Deiss’ section is reported in
order to establish the stratigraphic position of several beds from which
fossils were collected by the writer. The lowermost portion of Deiss’
Mount Whyte formation, containing olenellid trilobites and hence
Lower Cambrian, was reassigned to the Peyto limestone member of
the St. Piran sandstone.

ROSS LAKE SECTION (NO. 4)

(After Deiss, 1939. Fossil localities and lists according to the writer.)
MippLE CAMBRIAN

Mount Whyte Formation ie
eet
6. Limestone: finely to medium crystalline, partly oolitic, dark gray,

with partings and flakes of tan clay, mostly thin-bedded....... 26
5. Limestone: finely crystalline, dark gray, thin- and thick-bedded.
Fossil locality W4e near top of unit, locality W4d 5 feet above

Bases esis rotates Coleen at cca ie Stat Ae) ent en ote eter eer 56
Fossits (loc. W4e) :
Amecephalus cleora: (Walcott) ivi... soc os. cows eek ok c
Plagiuracercops-CWealeott)s 6fecen set habe sed bac meenak r
Fossirs (loc. W4d) :
Ameécephatus: cleora: (NValcott) iciaisca. dees eck ee ve des cc
Caborcellassrapian(\WWalcott)sesnen enero tee eee Cc
Plagimra:cercops. (Walcott) 22. 0.200 de sath pee ee Cc

Schistometopus converus Rasetti.........cceccecccoue i

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 21

4. Limestone: oolitic, dark gray, in beds 1-6 inches thick. Fossil lo- casi
cality Wace 3-5 feet above base of unit: .6)..5.2..6.00e a's de 22
FossILs:

Amecephalus cleora (Walcott) wo00s .0.0s000005 odes cc
Cavorcella skapia’ (Walcott). 2.22 ium ooh Oes cend oe ee r
Plagura cercops (Walcott). 00 tee cl i c
Schistometopus convexus Rasetti..........ccccccceees i

no ouale, sandstone, “and limestone: :....01 05 Seco Meee dees eens. 62

MP SME: ARCS MISTIESEONG SS «alate Slash aiote vidas oie Uvigln we ving e eee ev nee 45

MeeESSUIGE UIE MUIMICSTONGs Ya; ts i oiass oss b bas ceeds aeebedece eee aeones 53

Total thickness of Mount Whyte formation................ 264

MOUNT BOSWORTH

A section of the Cathedral and Mount Whyte formations was
measured by Walcott and more recently by Deiss (1940, pp. 739-751)
on the east ridge of Mount Bosworth, on the Continental Divide above
the Stephen station on the Canadian Pacific Railway. Both forma-
tions were remeasured by the writer. The results concerning the
Cathedral formation entirely agree with Deiss’, hence the writer’s
data are not reported. However, the writer found it difficult to
recognize in the field several of the units into which Deiss subdivided
the Mount Whyte formation. This is entirely attributable to the
different aspect that strata may assume depending on weathering con-
ditions. In the present instance, certain portions of the section can
be measured either in vertical cliffs or on gentle slopes, and lithologic
units that may be separated in one type of exposure become indis-
tinguishable elsewhere. The total thickness measured by the writer
closely agrees with Deiss’ data, but the subdivision into units is some-
what different. Hence the writer’s measured section is reported,
chiefly in order to define the position of a fossiliferous horizon. It
should be noted that, according to the writer’s interpretation, the
Mount Whyte formation includes at the top 105 feet of dark-gray
limestone that Deiss tentatively assigned to the Ptarmigan formation
(Deiss gave a thickness of 195 feet for these strata), whereas the
lowermost 49 feet of Deiss’ section, containing Bonnia and Olenellus,
are reassigned to the Peyto limestone member of the St. Piran sand-
stone. With these reassignments, the thickness of the Mount Whyte
formation as measured by Deiss would be 381 feet, in good agree-
ment with the writer’s value of 389 feet.

On the east ridge of Mount Bosworth, the Mount Whyte—Cathedral
contact is better defined than usual, a thick-bedded dolomite of the
Cathedral formation overlying the thin-bedded, dark-gray limestone
that Deiss assigned to the Ptarmigan formation.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

MOUNT BOSWORTH SECTION (NO. 5)
MippLE CAMBRIAN
Mount Whyte formation

6. Limestone: fine-grained or finely crystalline, dark gray, with tan
clay partings and flakes particularly in the lower part; in ir-
regular beds 1-3 inches thick. Some oolitic beds near the base. 105
5. Limestone: oolitic, dark gray, in irregular beds 1-5 inches thick,
with occasional shaly intervals. Fossil locality W5c 50 feet

ADOVEMDASE. < o.c «sew sents aie sora ws Bho eine ae aie hae eee ee 80
FossIts :

Amecephalus cleord (Walcatt) Gk ussite ao ceisipbidels a2 Cc

Cabercellaishapial (Walcott) wicaenieens aaiess seine sie Cc

Piagumarcercops ((NValcott) meri eeietadaa ire ier ariel Cc

Schistometopus convexus Rasetti........5.00-0csaee0- c

4. Shale and limestone: massive beds composed of thin alternating
layers of siliceous, gray, tan-weathering shale and fine-grained,

tay WiMEstOne) cara. scraid Scheie a asbieremmas meme eos oat cee ee 92
3. Shale and limestone: as unit 4, but with lesser percentage of lime-
SUOMEGH reiclay efores cy aracolahe wrsiaiore a etelouskelare nuise igheketiote ust auevetetchevetenerehe toletonerette 52

2. Shale and limestone: massive beds of siliceous, gray, tan-weather-
ing shale with interstratified thin nodular layers of fine-grained,
gray limestone. Upon solution of the limestone the beds

WEATHER tO PItEd SUGLACES) sree taciole } rus Guero sarees Sate leche omaeiens 36

1. Shale and sandstone: micaceous, green shale with interstratified
Sandstone beds: tj.cisains.cris.cis eee ble owiae ols Sigurt. ae oe eee 24
Total thickness of Mount Whyte formation................ 380

Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)

1. Limestone: crystalline, more or less sandy, limonitic, in thick,
irregular beds, with a little interstratified shale. Olenellid
fragments extremely abundant in the purer calcareous por-

PIONS acorns canta siiens aaa tate pate wth eieintest Sate cuata iets Senda era. Siar Sere 24

MOUNT VICTORIA

A section of the Mount Whyte formation was measured on the
south face of Mount Victoria directly above Lake Oesa. The Mount
Whyte formation is here exposed at an approximate altitude of 9,350
feet. The beds dip 10° east.

MOUNT VICTORIA SECTION (NO. 25)
MippLE CAMBRIAN

Mount Whyte formation
Feet

3. Limestone: crystalline or oolitic, dark gray, in beds 1-3 inches
thick. The upper portion interfingers with dolomite, making
the Cathedral-Mount Whyte contact more or less arbitrary. . 25

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 23

2 ade Feet
2. Shale and limestone: siliceous, gray, tan-weathering shale alternat-

ing with layers of fine-grained or crystalline, mostly thin-

bedded limestone. A crystalline limestone layer near top of

Mutt yielded AMecephnalus SP... ccc. cccccecsccscnccccceccscece 32
1. Limestone: finely crystalline, dark gray, weathering to mottled

surfaces owing to dolomite inclusions; in irregular beds up to

BPRCMCOMERIC Ma cain dais cis sis''s ana tied wee wpe oh ere rama e pa ee ee 6

Total thickness of Mount Whyte formation.............. 63

Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)

1. Dolomite: crystalline, sandy, gray, weathering dark tan, with con-
cretions of fine-grained dolomite resembling algal growths.
Lenticular beds of sandy limestone in top 3 feet yielded
SSPERIE MICE REAR INGHIES:. Sere tack. ctuly be dew nea: women a bavi eie es 17

CATHEDRAL MOUNTAIN

A section of the Mount Whyte formation was measured on the
northernmost spur of Cathedral Mountain. Here the Cathedral for-
mation forms inaccessible cliffs, but the Mount Whyte formation is
entirely accessible and well exposed. The strata are approximately
horizontal.

CATHEDRAL MOUNTAIN SECTION (NO. 23)
Mippte CAMBRIAN

Cathedral formation

Fee

1. Dolomite: crystalline, gray, tan weathering, massive excepting the a
lowermost 5 feet which are thin-bedded. Possibly these lower-
most beds should be included in the Mount Whyte forma-

(BIGTE es ors Geko CeCLCACEC IEC GISIGUICIERE RIOR CRE CDA er Not measured

Mount Whyte formation

2. Limestone: finely crystalline, dark gray, with tan clay partings
and flakes especially in upper part. The lower beds yielded
poorly preserved gastropods and trilobite fragments......... 75
1. Shale and limestone: alternating thin layers of siliceous gray, tan-
weathering shale and finely crystalline limestone. Limestone
prevails in the upper portion, forming a gradual transition to
VCCI IIE IG ods co ce gates wee wee dig ctecenenwae ae yéewaWes 80

Total thickness of Mt. Whyte formation..............0+5+ 155

24. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

LowER CAMBRIAN

St. Piran sandstone (Peyto limestone member)

Feet
1. Sandstone and limestone: more or less calcareous, limonitic sand-
stone alternating with sandy, crystalline, gray limestone. Ole-
nellid fragments and Bonnia abundant in the more calcareous

lavenS ase cece aliratees dauces.ame ee eT ee ee 30

POPES PEAK

A section of the Mount Whyte formation was measured in the
valley of a creek, tributary of Cataract Brook, that originates in a
glacier occupying a cirque between Popes Peak and Mount Victoria.
It is difficult to describe accurately the location of this section, but
the position is clearly indicated on the map (fig. 1).

The Mount Whyte formation is exposed and was measured at an
approximate altitude of 8,000 feet on the north side of a waterfall
formed by the brook a little below the tongue of the glacier. The beds
dip 22° north.

POPES PEAK SECTION (NO. 24)
MippLE CAMBRIAN

Cathedral formation
; Feet
1. Dolomite: crystalline, light gray, tan weathering, thick-bedded.

Lowermost 15 feet darker, still thick-bedded, forming a rela-
tively sharp contact with the underlying limestone of the
Mount Whyte: copia ttonlasereisieeiecrssicciecineresicreeineeiere Not measured

Mount Whyte formation

5. Limestone: subcrystalline, dark gray, in beds 1-3 inches thick, with
tan clay partings and flakes; lower part shaly.............00. 50
Limestone: oolitic, dark gray, thick-bedded................s0008: 27
3. Limestone: finely crystalline or oolitic, dark gray, in irregular beds
I-2 inches thick, alternating with some shale. Trilobite frag-
ments: (Amecephalus Sp.) wos anima ceeis se eens oe aes aes 52
2. Shale: siliceous, gray, tan weathering, with some thin limestone
layers: Gas. Je EA OM ses ates wom eee ee cee ee tees lrne are erate 50
1. Shale, sandstone and limestone: alternating thin layers of siliceous,
green-gray shale and green sandstone, with some thin, len-
ticular limestone beds. No fossils were found in this interval,
hence it is uncertain whether it should be placed in the Mount
Whyte formation or in the underlying St. Piran sandstone.... 47

is

Total thickness of Mount Whyte formation................ 226

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 25

Lower CAMBRIAN

St. Piran sandstone (Peyto limestone member)

Feet
1. Shale, sandstone, and limestone: alternating layers of green-gray
shale, gray sandstone, and crystalline limestone. Olenellid frag-

ments in limestone common up to the top of unit............. 22

EIFFEL PEAK

A section of the lower Cathedral and Mount Whyte formations
was measured on the southeast ridge of Eiffel Peak. The Mount
Whyte formation is exposed at an approximate altitude of 8,500 feet
and the strata are horizontal.

EIFFEL PEAK SECTION (NO. 20)
MippLte CAMBRIAN

Cathedral formation

6. Dolomite: crystalline, light gray, tan weathering, massive, forming ee
GRE pase 5s bears Hated aM TRIN cS Aa le oct Soa Not measured
5. Limestone: finely crystalline, very dark gray, thick-bedded,
WeatheringatonMnestlan GluttaCesap ai. aietereeiiats elec serie ioievelelels 80
Cathedral formation (Ross Lake shale member)
4. Shale and limestone: siliceous, gray shale alternating with irreg-
ular, thin layers of gray limestone. Near middle of unit, a few
inches of fissile, siliceous shale are abundantly fossiliferous
USealitvaC cont lees lOve cetell bc tote e sores we eho es keel kas 3
FossILs :
Albericiia Dosworine Walcott .2 5 6. esd ee causes t sake c
walventeiia: nitida WRESSEL .o\5 5 e's i's ot ven ble dle awe Dee r
Kochina americana (Walcott) .......cceeceeccncevees r
Miexicellaistator™( Walcott) <\0c% o% 0. SISTER ee c
Ptarmigania rossensis (Walcott) ......ccseceeeeeesces r
Vanuzemella:nortiasW alcotts ics. ssiic%-sctce delelewne sis slate c
Cathedral formation (continued )
3. Dolomite: crystalline, light gray or cream-colored, weathering
light tan, thick-bedded (thickness estimated) ..............+. 250
2. Limestone: subcrystalline, medium gray in upper part, light gray
in lower part, thick-bedded (thickness estimated)............ 40
1. Dolomite: crystalline, gray, tan weathering, in beds up to 5 inches
LANG Keteree ayfetcte Sate vateta fal ove ror tual ¥lsteieiuyeteraia wai Utes suoket avasniees! Cnebs eis ratpae cic 115

Total measured thickness of Cathedral formation.......... 488

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Mount Whyte formation

1. Limestone and shale: crystalline, dark-gray limestone with tan =
clay partings and blebs, in beds 1-2 inches thick alternating
with siliceous, fine-grained, green-blue-gray, fissile shale.
Mostly shale 16-22 and 28-34 feet above base, limestone in
other parts of unit. Fossils abundant in limestone 22-28 feet
above base, and in shale immediately above and below (locality
W200) © cs.s'xac case tome AAR USSR EEN oc sxe clea k eae 58
FossILs :
Amecephalus cleora (Walcott) .......0..cccccccceece cc
*Ficldaspis bilobata Rasctl wi. /isaaddsio. sven ase eale cc
furcatan Rasch: #2. decd: seas dh PO rr
HK ochaspis ect clenses RASCH; cas «0. dsdtdvoewmadd eee ee 1%
*Kochsella® maacys Rasettt. 60 <i cuits se asa ewes o'sluaiatorne c
Plaguura: ‘cercops CW alcott) <e.cec nase saws s ones ostsien ee Cc
*Schistometopus collaris Rasetti .........ccccccccecuns r
* convents Rasettat ei. eee eS
LowER CAMBRIAN
St. Piran sandstone (Peyto limestone member)
3. Dolomite: sandy, light gray, tan weathering..................05- 13
2. Sandstone: coarse, brown, massive, including round concretions of
fine-grained dolomite, resembling algal growths.............. 4
1. Dolomite: sandy, massive, weathering dark tan...............05- 12

Total thickness of Peyto limestone member of St. Piran
SANASOME 4.555.075) ite «Fis cide ee poy © a Peis oe ee 29
Note.—On the slope of Eiffel Peak, about 4 mile west of
the ridge where the preceding section was measured, the
Mount Whyte formation was found to consist of 62 feet of
fine-grained, crystalline and oolitic limestone in irregular beds
1-3 inches thick, with tan clay partings and blebs and a few
interstratified layers of siliceous shale. The Peyto limestone
member of the St. Piran sandstone is 20 feet thick, the upper
portion including the usual dolomite concretions.

MOUNT TEMPLE

A section of the Stephen, Cathedral, and Mount Whyte formations
was measured on the southwest ridge of Mount Temple from an
approximate altitude of 11,000 feet (Stephen-Eldon contact) to 9,200
feet (base of Mount Whyte formation) above Sentinel Pass. All
these strata are excellently exposed, undisturbed, and horizontal
throughout the mountain.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 27

MOUNT TEMPLE SECTION (NO. 21)
MippLE CAMBRIAN

Eldon dolomite

1. Dolomite: crystalline, light gray, tan weathering; massive in upper a
part, partly thin-bedded and mottled near base. Eldon-Stephen
contace fainly well. defied ¢.¥s.c..52602.c0¢ne covenees tae Not measured
Stephen formation
5. Limestone: very dark gray, finely crystalline, purer and thicker-
bedded (up to 4 inches) in upper part; thin-bedded and with
tan clay partings and flakes in lower part.................0- 168
4. Shale: siliceous, fine-grained, blue-gray, fissile................... 4
3. Limestone: crystalline, dark gray, with tan clay partings and
Haken peds T—2s1TiGhes sEhNIC kaye eeforeleretstersicie soe eleveietercere eiatels 12

2. Shale: siliceous and argillaceous, green- and blue-gray, fissile,

darker and harder near base; with a few interstratified thin

IIMIESTONE MAY CDSs cere ai ostreraiete da oe cia cla days eteieia oie eintetalevelatorss Cee 165
1. Limestone: finely crystalline to fine-grained, mostly dark gray,

partly (near the top) banded light gray; mostly in beds 4-5

inches thick, thin-bedded and more argillaceous near base.

Fossils abundant in subcrystalline, dark-gray limestone 100-

120 feet above base (locality S21b). Argillaceous beds near

base yielded pygidia of Glossopleura..........ceeesccserces 150
FossILs :
*Glossopleura templensts Rasetti.......ccesccccscccesce c
“Poly pleuraspis insigms RASCH. oii esles dtiess's'c a te sl c
Wotalethickness ot Stephen formation cee « sc slelec't steivieliereyols 499

Cathedral formation

g. Limestone: finely crystalline, gray, mottled, thick-bedded, espe-

cially in uppermost part. Contact with overlying Stephen for-

MHA OURVELVESHAL Dice ce scte cereis oe eres/s nile ie slevlcic creek ie cielo nie See 195
8. Limestone and dolomite: limestone as overlying unit in cliffs east

and west of ridge, partly replaced on ridge by crystalline, light-

gray, tan-weathering dolomite. Dolomite-limestone boundary

ET EAPECR CAT. whelyis ais, rare F Ulareib slate eed Sawin eee Wie mw ese crhletatar htc aieiate 200
7. Limestone: finely crystalline, dark gray, with calcite veinlets,

weathering to rough surfaces; mostly in beds up to 4 inches

thick, some portions thicker-bedded............ccccseececces 155
6. Limestone: subcrystalline, lighter gray than overlying unit, mas-
SVG fete sicvessin ote iapeyeretihars.é mrbbere sab ratt hts rerk iotatieie aise ete suntan tebe buat ae 100

SMR MIC ASIAING OF), 2 shy is ie, Suici cme Bie mya aiaie Dele ahpievh ate Gala Rumen 105

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Cathedral formation (Ross Lake shale member)

4. Shale: siliceous, hard, dark gray, with some interstratified nodular
limestone layers.) Locality Catmy. 2... isciasie oe dine « oh te etaeniets
FossIts:
Albertella bosworthi Walcott
Mexicella stator (Walcott)
Ptarmigania rossensis (Walcott)
Vanuxemella nortia Walcott

Cathedral formation (continued)

3. Limestone: subcrystalline, dark gray, partly mottled, massive.....
2. Limestone: subcrystalline, light gray, partly banded, thick-bedded.
1. Dolomite: crystalline, light gray, tan weathering, mostly thick-

bedded but with some thin-bedded portions................--

otal thickness ot Cathedral formation... +. 242.2 + seer

Mount Whyte formation

4. Limestone: crystalline to fine-grained, dark gray, in beds up to 4
inches thick in upper part; thinner-bedded and with tan clay
partings and flakes in lower part. Contact with overlying
Cathedral formation rather arbitrary because of irregular

dolomite-limestone, DOUnGaty sariisenese ieee icisieias cieinielsiie eine

3. Shale: siliceous, blue-gray, tan weathering, fissile................
2. Shale and limestone: siliceous shale with thin, nodular limestone
LAVEHS:S Eb Seis sere es SHES Save clei atere cr ousrovelae Poe tene Svan cron cies te cleeiete

1. Limestone: subcrystalline, dark gray, in beds 1-2 inches thick.....
Total thickness of Mount Whyte formation...............

Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)

2. Sandstone and limestone: gray, calcareous sandstone and crystal-
line, more or less sandy limestone with abundant olenellid
FEAST TIES We varerstayahccepovere Oe tape: ohesasicievehere cishacers ois xelaiers asin ae
1. Sandstone: coarse, brown, with spherical dolomite concretions re-
Semibling ‘algal, @rOWEIS «on -s1ss coi © xis.0o lees. © as caw ee ie

Total thickness of Peyto limestone member of St. Piran
SANGSEOME™ o/s evejeiersisse sisssre aceteneterstersiobere ercters el oreye ete atevecers ened

PINNACLE MOUNTAIN

Feet

195
45

225

1,228

45
53

24

30

A section of the Mount Whyte formation was measured on the
east ridge of Pinnacle Mountain, above Sentinel Pass. Pinnacle
Mountain constitutes a fault block downthrown approximately 500
feet with respect to the Mount Temple fault block, the fault line

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 29

passing through Sentinel Pass. The Mount Whyte formation is ex-
posed at an approximate altitude of 8,700 feet. The beds are hori-

zontal.
PINNACLE MOUNTAIN SECTION (NO. 27)

MippLe CAMBRIAN

Cathedral formation

Feet

1. Dolomite: crystalline, gray, tan weathering, thick-bedded; grad-
ually becoming thinner-bedded near base, forming transition to
Smuer NINES TOMMANON 5 o5056.s nccee bRaneve wink one EEO Not measured

Mount Whyte formation

4. Dolomite and limestone: finely crystalline, light gray, banded dolo-
mite in beds 2-5 inches thick grading laterally into dark-gray
DIPS URMLC® ip Steve ate, Sr he syay week, Aiuvara Siete RUCIR Sate aie are panacea 20

3. Limestone: subcrystalline, dark gray, with tan clay partings and
Hakes Ptilin=DEddedies & ceccre crs’ c cis «1s ereinremresiesdere s cnettiotectins oats 28

2. Shale: siliceous, gray, tan weathering, fissile; with some thin, nodu-
lar limestone layers. Rare fossils in shale (Amecephalus sp.) . 48

1. Limestone: fine-grained, gray, in thin, irregular beds; some shale
ATIC PYRE tgs DAL terse olc tats aterete aio ei oteroraarcve olete staterireiaieralalwinnsreistete ates 12

Total thickness of Mount Whyte formation................ 08
LoweER CAMBRIAN
St. Piran sandstone (Peyto limestone member)

3. Limestone and sandstone: sandy, crystalline, gray limestone and
calcareous sandstone, with a little siliceous shale. Olenellid
fragments abundant throughout the calcareous layers........ 21

2. Sandstone: dark brown, with round concretions of fine-grained,
tan-weathering dolomite resembling algal growths........... 7

1. Sandstone and dolomite: sandy, gray, tan-weathering dolomite and
AO OUIIt ExSATICStONGm: isin idl cecscvciens cious chores rece ee aerate Caer 15

Total thickness of Peyto limestone member of St. Piran
SARUSEONE MoaGiiis scram anid siti x adonig sas AN Cee 43

MOUNT SCHAFFER

Mount Schaffer is cut by a fault into two blocks. The northeastern
fault block includes the summit of the mountain. The southwestern
block is downfaulted several hundred feet, bringing the base of the
Cathedral formation against the upper part of the St. Piran sand-
stone. The following section was measured in the southwestern fault
block. Another fault separates the southwestern part of Mount
Schaffer from Park Mountain, which belongs to a still farther down-

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

thrown fault block, the middle part of the Cathedral formation in
Park Mountain resting against the top of the St. Piran sandstone
in the southwestern block of Mount Schaffer.

The section was measured on the slopes and cliffs overlooking
McArthur pass, and includes the Mount Whyte formation and the
Peyto limestone which here attains considerable thickness and carries
more abundant and better-preserved fossils than most of the other
sections. The beds dip 15-20° southwest.

MOUNT SCHAFFER SECTION (NO. 22)
MIppLE CAMBRIAN

Cathedral formation

et
1. Dolomite: crystalline, gray, tan weathering, massive; lowermost me
portion is massive undolomitized limestone, distinguished from
underlying Mount Whyte formation by the thickness of the
beds, although the change is not very sharp............ Not measured
Mount Whyte formation
5. Limestone: gray, mottled, subcrystalline, in very irregular beds
Up tos miches ticks 6 35.08 Sha a cite ws nye dare asisleninie ale aa aire 25
4. Limestone: gray, shaly, with tan clay partings and flakes........ 20
3. Limestone: finely crystalline, partly sandy, gray, weathering to
mottled surfaces probably owing to dolomite inclusions; mas-
sive, weathering to irregular fragments.............e.eee00- 45
2. Sandstone: coarse, brown, calcareous in upper part.............. 20
1. Shale: siliceous, fine-grained, dark gray, fissile................. 2
Total thickness of Mount Whyte formation................ 112
Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)
5. Sandstone s (coarsey wiehnte Prowler ate enc latetatelare asic enetamrcrchsvel steve tenerolers 30
4. Limestone: crystalline, gray, partly sandy, irregularly and mostly
thick-bedded, up to 5 inches. Olenellid fragments and Bonnia
abundant throughout. Fossils collected near base (locality
P22k) and 10 feet below top (locality P22m). U.S.N.M.
locality ‘61d is clearly within ‘this unite 6.0... se Vea ener 60
Fossits (loc. P22m) :
CAGr alos: UNnC@ NV Alcott. 1: eine aisle celeb sate = ies [aerate iF
Bonmia feldensys: (GWialeott)ianwrenterct eae eit cc
Crasstjumbra sli OWalcote) ieee ets cieetee ee r
OVEN ELIS ESD ine creristnciche bie aT nie Ghee eee oer cc
POCdeUMtas SON Nabe eee dea atthe owes cee ee caeae sire r

ZLacanthopsis cits lewis) CWalcott) eaectscsecesros eee ee r

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 31

Feet
Fossits (loc. P22k) :
erases Unc VVAICOLL. <3. s\s3 ssa aed dhe Oaede aaedoae c
Domus pipidcuse (Walcott) 2... stsecec cs nuse eee enews cc
CFIERENE SPA Son's sats 6 ve's dae So eae akeos Rae a cc

3. Limestone: thick, irregular beds of sandy or dolomitic limestone
with inclusions of purer limestone, assuming mottled aspect on
MVC ERC ITIE Foye to iain esiarcials oym'e 0) em se MS Sia aie RISE ee OU eee 60
2. Sandstone: partly calcareous or dolomitic, brown, with concretions
of fine-grained dolomite or limestone resembling algal

TOMAS oe cis ocieis< én .c crapp ouc'c ovnn vic oes v OL ee Smee Saale « 16
1. Limestone: finely crystalline, gray, weathering to mottled surfaces
probably owing to dolomite inclusions..............eeeeee005 32

Total thickness of Peyto limestone member of St. Piran
SHMCSEONG. f5 cs asc os pds uivie oa ele a mee a erate a wae one 198

MOUNT ODARAY

A section of the Stephen, Cathedral, and Mount Whyte forma-
tions was measured from the top of the south peak of Mount Odaray
down its southeast ridge, then following a broad shoulder that de-
scends in the northeast direction toward Odaray Plateau.

The summit of the south peak of the mountain is formed by the
black limestone of the upper Stephen formation, which therefore is
not completely represented. The formation is not complete even in
the higher north peak.

The beds dip almost uniformly 15-20° southwest across the entire
mountain.

Every bed of the Cathedral formation is well exposed and was
carefully examined, but the writer was unable to find the Ross Lake
shale. It appears exceedingly unlikely that it could have escaped
observation, hence it is assumed that it is not developed at this locality.

MOUNT ODARAY SECTION (NO. 6)
MipptE CAMBRIAN

Stephen formation
Feet

3. Limestone: subcrystalline, black-gray, more or less argillaceous,
in beds 2-4 inches thick, locally grading laterally into dolomite.
The dolomitized beds preserve the original stratification and
are often banded, a cream-colored dolomite alternating with
dark-gray limestone. Thickness of unit not accurate because
of slope of ridge almost paralleling the strata. Fossils pres-
ent at several horizons but abundant only in top 20 feet, just
below the summit of the mountain (locality S6l)........... 280+

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

FossILs:
Alokistocare sinuatum Rasetti...........cccceccscccce r
+ Athabaskia#, parva Wascttljansasasceioeeld. Hoes eee if
Bathyurtscus adacus Waleotts.6...06ds00.t och secasads c
* Chania OCAKAyensts NaSettinsacces ces ce nace eee meee r
*Elrathing margimalis: Rasettl: ). 699s. ob eeb ite cece ope c
Olenoidessspiic Gace ate te tee an r
*Pachyaspis attenuata, Rasetttt: «. di. sae dade desiddaeh e's c
Pagetia:ci. (Pvboates: Walcott) és c.aban eeiseed le bee a: c
Peronopsis columbiensis Rasetti........0000cceceecens r
SOEROPICUP CUE SO. 5d ak cts Se ee cle amemeca een. nee re
Tonkinella stephensts Kobayashi... ......0e0se0scee0es c
Zacanthowdes divergens Rasettt,.<..:0isclecs 0 oa0e0 0s be r
* langipygus: Rasettl ..s)on.wee cine eoends es (
ns subyunticus. Rasettt «iso «do. as eestnciie stan c

2. Limestone and shale: black-gray, thin-bedded, shaly limestone
alternating with dark-gray and tan siliceous shale. Many of
the thin limestone beds are highly fossiliferous, but it is diffi-

cult to collect unweathered fossils (locality S6k)...........
FossIs :
Bathvuriscus: adacus: Walcott dicate deucdtescaa wd taemare cc
Elrathina paralléla: Rasettt-. ¥ o/sisieas ogee occyenees de's Pale cc
* SPINISETO IRASCUED ive warclctercre clase cate ferme cc
KOotenia Spits ces esc hare dole ooo en ee eke eee SOIC 5
Pagetis ci. P.cbootes Waleott: 20g. fos tee O08 see 2 c
*Peronopsts columbiensis Rasettin.... <2. 00<es+=+sas cc
Tonkinella stephensis Kobayashi..........0ceeeeeeees Cc

I. Shale and limestone: siliceous, fissile, green-gray, tan-weathering
shale alternating with crystalline and fine-grained limestone,
mostly in thin, lenticular beds. Fossils collected in crystalline
limestone layers 12 feet above base (locality S6g). Contact

with underlying Cathedral formation very sharp............
FossILs :
Elmaniella burgessensis Rasetti:.......cccccccccyeccs c
Solenopleurelia: spe INO. Bia 2ilosc. cahsmsbaadewiers detente r
Total thickness of Stephen formation...................5.

Cathedral formation

4. Dolomite: crystalline, light gray or cream-colored, weathering pale
fans thiek-bedded. its eee sees vale neers fo ae eee
3. Limestone: fine-grained, light gray, with less soluble layers
weathering in relief forming banded surfaces: very thick-
bedded, in places grading laterally to dolomite.............. ;
2. Dolomite: crystalline, cream-colored, weathering pale tan, thick-
bedded: :.isicasce erick ede Semreaeeteeida tc Re vee ee
1. Dolomite: light gray, crystalline, weathering dark tan, thick-
bedded! aU snasiate ett sheests acaist « heiapestelomine sche Seite wiers

Feet

40

120

440+

1,000

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 33

Mount Whyte formation

Feet
3. Shale and limestone: massive beds composed of alternating layers
of gray limestone and siliceous, gray, tan-weathering shale.
In top 3-5 feet, beds up to 3 inches thick of black-gray finely
crystalline limestone are locally developed and yield fossils
MRP CELLP NVR Fc a: acid 2, 210:'5:0,0hare aint tans sap sinca mai aia a epee aie oe 85
FossILs :
Faeldashiseh: F: bilobata Rasettt. 05.1.0. sews ss vviseiaw oes c
IG CHAS PES CUTeleHsts = RASCLEL aii «ac clejelectald a eeidieei eva ciniere r
2. Limestone: finely crystalline, dark gray, with sandy layers
weathering in relief; very thick-bedded, forming cliff......... 66
1. Shale: siliceous, fine-grained, fissile, dark gray..............ee0- 10
Total thickness of Mount Whyte formation............... 161
Lower CAMBRIAN
St. Piran sandstone (Peyto limestone member)
5. Limestone: coarsely crystalline, medium gray, pure or sandy, in
thick, irregular beds with poorly developed partings. Olenel-
lids ‘and Bonnia common (locality P6k)....0..5080s vet ece ees 81
FossILs :
Bouma feidensis (Walcott) a6. ».+2 edeesied am nice s s'earw ee c
IGTALNS? Beat Bc coc tee COR COT OC Oa Oo nO aoe en soe c
Any Polomite-.sandy,eray, tan weathering. o. . ces. es cee ce «se clare 5
3. Limestone: coarse, sandy, dark tan; with round concretions of fine-
grained limestone or dolomite resembling algal growths...... 8
2. Sandstone: purple or maroon, massive, with concretions as in over-
lyysuayer eravke he {oyn) Zhaval loYounavedla Roun ocoAocbod Tabu DLHnancondEo lo” 20
1. Sandstone and limestone: massive beds of irregularly interstrati-
Red Eray MimmectOne and SANGSEONE. os a0 cscee eee sce spienp sae 30
Doral taiekness. of Peyto limestone. ..<« 4. <'c2%,<.0ceimaseineste oe 144

PARK MOUNTAIN

The northeast face and the north ridge of Park Mountain, over-
looking Lake McArthur, offer continuous and accessible exposures
of the entire Stephen and the lower part of the Eldon formation.
Park Mountain is separated by a normal fault from Mount Biddle
at the east. Almost immediately southwest of the summit of Park
Mountain, a succession of faults brings highly disturbed Upper Cam-
brian strata to the surface, and no outcrops of the Middle Cambrian
are known farther west. Hence Park Mountain, as Mount Odaray
and Mount Stephen, lies at the southwest margin of the Middle
Cambrian outcrop area where the Middle Cambrian sections present
considerable differences from the typical sections of the Bow Range.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

In Park Mountain the anomalous character of the section chiefly
consists in the unusual lithology of the lower portion of the Eldon
formation.

The Eldon formation is not discussed in detail in this paper, but
it was mentioned that in the type section on Castle Mountain ( Deiss,
1939), on Ptarmigan Peak and Mount Bosworth it consists of over
1,000 feet of dolomite with some calcareous beds. On Mount Stephen,
the Eldon includes in the lower portion 150 feet of siliceous shale,
most of the rest being dolomite. Hence it was entirely unexpected
to find that on Park Mountain the Stephen formation is not overlain
by the usual dolomite, but rather by a thick succession of siliceous
shales and argillaceous limestones. Several hundred feet of such
strata were measured, and there is no doubt that beds of this lithology
extend for a considerably greater thickness.

It would be of great interest to observe the lateral change of the
Eldon formation from dolomite to siliceous shale. However, most of
this change probably took place in an area between Mount Biddle
and Park Mountain where the Eldon has been largely eroded away.
The writer did not climb Mount Biddle, where possibly the Eldon
already shows peculiar lithologic character.

The section given below was measured, for the Eldon formation,
in the middle part of the northeast face of the mountain. The Stephen
formation here is mostly covered by snow and talus, hence it was
measured farther northeast, along the broad north ridge of Park
Mountain. This ridge and the nearby slopes offer excellent exposures
and opportunity for good fossil collecting. The dip of the strata
across all the investigated portion of Park Mountain is 20-24° south-
west.

PARK MOUNTAIN SECTION (NO. I0)
MIppL—E CAMBRIAN

Eldon formation

Feet
4. Shale: siliceous, massive, fine-grained, gray, weathering banded
gray and tan; with a few thin limestone layers. Only the basal
part of this unit was examined in place, but strata of this li-
thology certainly extend to the summit of the mountain.
Whiekmess:; esturtated, ci ek Sate eee 800-++
3. Limestone: crystalline, dark gray, in beds 1-3 inches thick. Abun-
dantly fossiliferous in lower 30 feet (locality TOC) Gee cee 70
FossILs:
Athabaskiaispiwit cee nek ei a, CR r
BOtwuris cis) Sp ists ae bet hos sae ere ee ee r
GLSPROSPIS: SP. a5 nav cbe nae ee ee r
Kootenia Sp. sa: o8 ooteu er et ok ate tree oan eee c

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 35

Feet
2. Limestone: fine-grained, argillaceous, dark gray, in beds 1-3 inches

thick, gradually changing to purer limestone of overlying unit.

In places grades laterally to dolomite lenses, the dolomitized

beds preserving the original stratification marked by clay part-

ings. Unit apparently thickens westward; thickness estimated. 360
1. Limestone and shale: limestone as overlying unit, with bands at top

and bottom of varying thickness of siliceous, gray, tan-

weathering shale enclosing lenses of gray, fine-grained, bright-

tan-weathering limestone. Thickness estimated.............. 90

Total estimated thickness of Eldon formation.............. 1,320+

Stephen formation

3. Limestone: fine-grained or subcrystalline, more or less argilla-
ceous, black-gray, in beds up to 5 inches thick. In upper 50 feet
grading into partially dolomitized beds banded light and dark
gray. Fossils abundant 180-200 feet above base (locality Srol)

and 15-30 feet above base (locality S1ok).................. 420
Fossits (loc. Stol) :
*Alokistocare cataractense Rasetti..........secseeeees c
Bathyuriscus adaews Walcott.........c0..ceeeeccevccs c
*Glyphaspis parkensis Rasetti......ccssccccccesscccess c
RO OLENVE- SPs pois aialeces < olaie 0 Sine eyarsiereeeieterayele agile le dhe) sravs 6 ote r
“Parkaspis endecamera Rasetti. ovis. vs sa sre osieadee ss c
Tonkinella stephensis Kobayashi............6seeeee08 ie
*Vuknessaspis. paradoxa Rasetti.....c.cececsvevessece r
*Zacanthoides divergens. Rasetti......0s00es0se0accs conus r
Fossits (loc. Stok) :
Bathyurtscus adaeus Walcott... ........00.0eseeeeees c
Peronopsis columbiensis Rasetti. ......0600.cses000ees c
Tonkinella stephensis Kobayashi..........seeeesseees c
UNI SOM vo chic cia sal Srvc arafecom aw 'si a*eiainlw cyniwie Reo aR sles c

2. Shale and limestone: massive beds of thin alternating layers of
argillaceous, gray, tan-weathering limestone, some purer, fine-
grained, gray limestone, and siliceous, gray, tan-weathering
shale; mostly shale in middle portion of unit............... 100

1. Shale and limestone: argillaceous, gray, tan-weathering shale alter-
nating with coarsely crystalline, gray, massive limestone. Con-
tact with underlying crystalline, massive, tan-weathering dolo-
mite of Cathedral formation very sharp. Several hundred

feet of the Cathedral formation are well exposed........... 28
Total thickness of Stephen formation..............eee0e5 548

MOUNT FIELD: KICKING HORSE MINE

A good section of the Mount Whyte formation is exposed in the
vicinity of the Kicking Horse Mine at the base of Mount Field. The
base of the formation, however, is everywhere covered with talus. The
beds dip eastward.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The section described below was measured immediately east of
the mine. The Cathedral-Mount Whyte boundary must be defined
arbitrarily within certain limits, as the upper limestone beds of the
Mount Whyte grade laterally into dolomite, and there is no abrupt
change in thickness or other characters of the strata. An average
horizon for the limestone-dolomite boundary was assumed in deter-
mining the thickness of the uppermost unit of the Mount Whyte
formation.

KICKING HORSE MINE SECTION (NO. 7)
MIppLeE CAMBRIAN

Mount Whyte formation

4. Limestone: finely crystalline, hard, black-gray, in beds 1-3 inches "7
thick; top portion irregularly dolomitized. Fossils collected
40-50 feet above: base:; (locality, W 7g) 5 «cise « sila viele ele» visiale 110
FOossILs :
*Caborcella are SRASEU oa =ctioe hats cathe oe ie ois aloeamee r
*Figidaspis superba, IRASCttA. «2625 mt verde oo elawlaihs Oe eutiee c
*Onchocephalus depressus Rasetti .........cecccccceee c
* SUDIAETIS™ INAS@tiltw errs ajecrcisisielelele:ctetnetetete rf
3. Limestone: crystalline or oolitic, dark gray, in beds 1-3 inches
thick: + css vitae nedeslnadion Hediste ate Sacea ete erie eer ae remene 5
2. Limestone: crystalline or oolitic, dark gray, with tan clay partings
and flakes, in beds 1-2 inches thick; more shaly in lower por-
tion. Fossils collected 72 feet above base (locality W7f),
10 feet above base (locality W7e), and 1-2 feet above base
(locality W7d). Also from undetermined horizons within unit
(oealities W 7, WHY) oe ee a. ore oe tants me eas acs caieteioe ae 75
FossILs:
Amecephalus cleora (Walcott)\.......+ 5 .0-0<-uesc0ss r
TP CIGGSOES JUPCULE TRASEUCN occas oun. 26.c%5 oe ioyea oferurar ots pale ala cc
Keochaspis: cuif clemsts’ TRASCUER cies ose» antenna ne yeies s Keats r
Kochrella2 ict. Ka mare vey RASetth ere «ie cope a) ais oleeiets r
*Oryctocephalites ressert RaSettic....0 0c. sjesccesiooness r
Schistometopus converus Rasetti..........ceeeeecees r
*Onchocephalus fieldensis Rasetti........eeeececeeeeees ce
* MNALOT, WRASCLt le Secceeycice. aeeticene Aoaeek c
Fossits (locality W7e) :
Gaborcella: skapta OW aleott) 0 cout wnins <tc = cepaee eens Cc
Faeldaspss furcata (Rasettt. . <5 5. toes wae ae ee eronens c
Plagiura-cercops (Walcott) wacas: «0% saiass eee ee r

Fossits (loc. W7d):
Amecephalus cleora (Walcott)
Caborcella skapta (Walcott)
Onchocephalus fieldensis Rasetti

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 37

1. Shale: siliceous, banded gray and tan. Top of interval forms bench
in cliff, crossing portal of Kicking Horse Mine a little above
floor. Base of unit and Mount Whyte-St. Piran contact
covered by talus. Fossils in thin limestone layers in top 3 feet
Oiesitenvar | lotanty- W'7G) icsesh .xneheercseay ehhuade seks 105+
FossILs:
Amecephalus cleora (Walcott)
Caborcella skapta (Walcott)
Fieldaspis furcata Rasetti
Onchocephalus fieldensis Rasetti
Total thickness of Mount Whyte formation................ 295-+

MOUNT FIELD: BURGESS QUARRY

A section of a portion of the Stephen formation was measured on
the west slope of the ridge between Mount Field and Mount Wapta,
from strata somewhat lower than Walcott’s celebrated quarry in the
Burgess shale to near the top of the ridge. In the description of the
Burgess shale, the problem concerning the stratigraphic position of the
beds measured in this section is discussed in detail. The fault block in
which Walcott’s quarry occurs and where the section was measured
is isolated by faults or drift cover from beds of recognizable position
in the general section. Hence the assignment of the Burgess shale
to the Stephen formation rests on paleontologic evidence. However,
the relative ages of three faunules collected from this section rest
upon unquestioned stratigraphic evidence, even though their relation-
ship to faunules from other sections remains uncertain.

Throughout the measured section the strata are approximately
horizontal and continuously exposed except a portion of the lowermost
beds.

BURGESS QUARRY SECTION (NO. II)
MippLeE CAMBRIAN

Eldon (?) dolomite
} : . Feet
1. Dolomite: crystalline, gray, tan weathering, thick-bedded except

near base, seaching’ tO top OF TIGEEL 2. oak cae wees « Not measured

Stephen formation (Burgess shale lentil: type section)

5. Shale and dolomite: siliceous, gray, tan-weathering shale alternat-

ing with thin- and thick-bedded, gray, tan-weathering dolo-

tal) (owe gee) CI MCE AGO nacre me oT ee oe ae 120
4. Shale and limestone: siliceous and calcareous shale, alternating

with thin beds of gray limestone; all beds except some of the

purer limestone layers weather tam........cccseecscccceceee 100

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

3. Shale: in upper part siliceous, argillaceous, and calcareous; in pe
lower part siliceous, hard, fine-grained, dark gray. Fossils
130-140 feet above base (locality St1g)..............ceecce. 210
FossILs :
Ehmaniella burgessensis Rasetti...........0.--eceece cc
POGCHGUSD. - dete ale wi aicrers aeicieieal cies Sain ieee relic eae c
Solenopleureiia sp. NO.*3..7 .ahewerss caches ae eees c

2. Shale: siliceous, fine-grained, hard, dark gray. This is the cele-
brated “phyllopod bed” of Walcott. Locality S11f (=U.S.N.M.
35K) il walscicc canis dente aise weteerte te betas ere eee 20
FossiLs: see list in discussion of faunas.

Stephen formation (continued)

1. Shale: calcareous and siliceous, mostly covered by talus. Fossils
abundant but poorly preserved about 60 feet below top of unit

(locality Sure) Vas .oc se oointis caves cite haviclevsivee dea mane 100+
Fossits:
Paterma cioP; senovia CWaleott).......2.+--seces cee c
Bathyuriscus rotundatus (Rominger).............+4. Cc
Erathinarspeyenestee 2b ath PO et aS r
Kidoteiiat sp: Vii. bats suet ted waidirct ais Sule ose ele oy cee MoE ete ae r
Olenotdes serratus) (Rominger) 2...) 5.0ssccsecessees c
Pageiia Bootes. NVACOLES - ee aca cae Sineaee cole eeas Cc
Total thickness of Stephen formation...........sse0cce0¢ 550+

Approximately 0.3 mile south of Walcott’s quarry and at the
same altitude, beds of argillaceous, gray, tan-weathering, thin-bedded
limestone are exposed in shallow gullies. These strata would appear
to be a lateral continuation of those exposed in Walcott’s quarry,
notwithstanding the different lithology. However, the beds are not
continuously visible between Walcott’s quarry and the above-men-
tioned outcrops, and small faults may be concealed under the drift
cover. Hence the writer was unable to ascertain the exact strati-
graphic position of this argillaceous limestone in the Burgess quarry
section. The beds are abundantly fossiliferous, entire trilobite shields
weathering in large numbers on the surfaces of the limestone slabs
(locality S1te).

FossILs :
I phadetla’ chs fieldensis Ressets. o:d.,:% sje ae te dentacee c
*Alokistocarella fieldensis. Rasetti..........50s00ceee0s c
Bathyuriscus rotundatus (Rominger)..............+0. r
*Blrathina. brevijroms kasettioes es ce cen se eeece ces cies cc
Kootema. burgessensis TReSSCtjta. osc yess ciee ce stisceieeel Cc
Olenoides’ serratus GRominger)) > cc. cae cee elles oes Cc
Pagetia bootes Walectt .5 ake a at as cy wee ie Cc

Solenopieurelia sp. No: Yay. asi itedatebs tte weeks r

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 39

MOUNT STEPHEN
GENERAL DESCRIPTION

No other mountain in the area presents stratigraphic and paleon-
tologic problems comparable to those encountered on Mount Stephen.
Hence an especially intensive study was made, about 4 weeks being
spent on this mountain alone. Almost every accessible exposure below
the Eldon dolomite was examined.

The better to understand the topography, stratigraphy, and struc-
ture of Mount Stephen, two photographs of its northwest face (pls.
3, 4) and a large-scale map (fig. 3) are presented. Furthermore,
to facilitate the description several minor topographic features are
named. The above-mentioned illustrations enable the reader to locate
exactly topographic features, stratigraphic units, and fossil localities.

Mount Stephen is bounded at the east by the deep valley of Mon-
arch Creek, which follows a fault line (Cathedral-Stephen fault).
A broad ridge (henceforth referred to as the “North Ridge”) de-
scends from the summit of the mountain northward to the Monarch
Mine and the tunnel of the Canadian Pacific Railway (indicated in
incorrect position on the Lake Louise sheet, but placed correctly on
the map, fig. 3). Southwest of this ridge extends the precipitous
northwest face of the mountain, overlooking the Kicking Horse River.
The lower and upper parts of this face are accessible only at a few
points, but the middle slope forms a shelf that permits one to traverse
easily the entire face at an approximate altitude of 7,000 feet. Deep
gullies have been eroded in this face, and the three principal ones are
named, in order from northeast to southwest, ‘North Gully,” ‘“Mid-
dle Gully,” and “Fossil Gully.” They are indicated on the above-
mentioned map and photographs (fig. 3 and pls. 3, 4).

The portion of the mountain so far described is essentially one
fault block of almost horizontal strata. Only minor normal faults,
involving displacements probably nowhere exceeding 100 feet, break
the continuity of this block (one such fault follows North Gully,
others are visible near the Monarch Mine).

A far more important fault, here named the “Fossil Gully fault,”
follows, in approximate north-south direction, the lower part of Mid-
dle Gully, the upper part of Fossil Gully, and traverses horizontally
the entire west slope of the mountain at an altitude somewhat exceed-
ing 7,000 feet. The approximate course of this fault is marked in
figure 3. The fault continues across the Kicking Horse River and is
traceable on Mount Field. The portion of Mount Stephen west of

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

the Fossil Gully fault is upfaulted and the strata dip steeply to the
west, approximately paralleling the slope of the mountain. For this

KICKING
HORSE MINE

MT. STEPHEN
Ja

10495

05
MILES
Fic. 3.—Contour map of Mount Stephen indicating fossil localities. West of the
Fossil Gully fault (dotted line) the strata are greatly disturbed.
reason, besides the presence of other faults, folding, and cover by
glacial drift and vegetation, the portion of Mount Stephen west of
the Fossil Gully fault is unfit for stratigraphic work. The celebrated

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 4I

outcrop of the Ogygopsis shale occurs in this disturbed area and
cannot be placed in the section.

The preceding description shows that the strata underlying the
Eldon dolomite can be studied only in the east face (in the valley
of Monarch Creek), in the North Ridge, and in the northwest face
from North Ridge to Fossil Gully.

In the valley of Monarch Creek, the Mount Whyte and lower for-
mations are not exposed, and a good section of the Cathedral for-
mation cannot be obtained because of the considerable horizontal
extent of any accessible section traverse. Hence only a section of
the Stephen formation was measured (section No. 26).

The North Ridge is largely inaccessible, and at the level of the
Stephen formation forms a bench mostly covered by talus. Only the
Mount Whyte formation is accessible and well exposed at the edge of
the talus below the Monarch Mine (section No. 17).

The northwest face offers continuous exposures of the entire Mid-
dle Cambrian and the uppermost portion of the St. Piran sandstone.
A section (No. 16) was measured along North Gully, where the beds
almost up to the top of the Mount Whyte formation are excellently
exposed and easily accessible. The top unit of the Mount Whyte
formation forms an overhanging cliff and the gully cannot be climbed
farther.

Middle Gully is accessible in its lowermost part, but soon vertical
cliffs prevent further progress. From the top of the talus slope
occupying the lower part of Middle Gully, one can easily reach the
steep wooded ridge separating Middle Gully from Fossil Gully and
follow it until it merges with the slope at the foot of the limestone
cliff formed by the top unit of the Mount Whyte formation. Tra-
versing talus slopes almost horizontally in the southwest direction,
one reaches the bottom of Fossil Gully where large dolomite boulders
emerge from the talus slope. From this point, Fossil Gully is acces-
sible to its termination in the broad ridge that separates the northwest
and west faces of Mount Stephen. Furthermore, one can easily
climb out of the gully on either side and traverse either the shelf of
the northwest face to the North Ridge, or the west slope as far as
the outcrop of the Ogygopsis shale at the celebrated “Fossil Bed.”
It is also possible to examine the lower portion of the Cathedral
formation in Middle Gully, where it forms easy slopes between the
vertical cliffs of the harder underlying and overlying units.

The northeast side of Fossil Gully and the slopes and cliffs above
the upper end of Fossil Gully offer a continuous section of approxi-

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

mately horizontal strata (section No. 9) extending from the top
unit of the Mount Whyte formation inclusive to the top of the
Stephen formation and higher beds if desired. This section is con-
tinuous within one fault block with the section measured in North
Gully, hence the two can be united and represent a continuous section
of the entire portion of the Middle Cambrian studied in this work.
In the bottom of Fossil Gully, the beds are somewhat distorted by the
Fossil Gully fault. On the southwest side of the gully, the beds are
upfaulted by 500 feet with respect to the northeast side. For some
distance, before the strata are further disturbed by other faults, the
beds are still horizontal and supply another excellent partial section
of the upper part of the Mount Whyte and the lower part of the
Cathedral. This section is an exact duplicate of the one measured
on the other side of Fossil Gully, and it would not be worth mention-
ing except for the fact that here the Yoho shale member of the Mount
Whyte formation is unusually well exposed for collecting and affords
one of the richest fossil localities in the entire area (locality Wok).

Walcott (1928, pp. 315-319) described a Mount Stephen section,
stating that his traverse extends “from the summit of the mountain
down its northeast and north slopes to the track of the Canadian
Pacific Railway at the tunnel east of Field.” This would indicate
that Walcott’s theoretical section traverse should follow what is here
designated as the North Ridge. As already pointed out, the Mount
Whyte formation is the only one that is well exposed and accessible
along this traverse. The Cathedral formation is mostly exposed in
inaccessible cliffs and it is clear from Walcott’s description that it
was not measured in place, the thickness being inferred from the dif-
ference in elevation between top and bottom. The Stephen forma-
tion on North Ridge is partly exposed in inaccessible cliffs, partly
covered with talus. Apparently Walcott did not measure here the
section of the Stephen formation described as typical, since he affirmed
(Walcott, 1928, p. 247) that the type locality of the Stephen forma-
tion is the “northwest side of Mt. Stephen.” In the description of
the section, he placed the Ogygopsis shale, stated to be 150 feet thick,
in the upper part of the Stephen formation between two units of
bluish-black limestone. However, according to Deiss’ and the author’s
observations, the only available outcrop of this shale is at the famous
“fossil bed” where the shale forms a deeply weathered dip slope and
is separated either by faults or drift cover from any rocks in recog-
nizable stratigraphic position, At the nearest place where a continuous
section of the Stephen formation can be measured, above Fossil

—————————————

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 43

Gully (section No. g) there is no trace of a shale bearing the Ogygop-
sis klotzi faunule anywhere within the explored stratigraphic range
from the top of the St. Piran sandstone to the middle of the Eldon
dolomite. One must conclude that the Ogygopsis shale, at least in its
recognizable fossiliferous form, is a narrowly localized deposit that
cannot be placed in the section by stratigraphic evidence. In the
Fossil Gully section there are, in the undivided Cathedral and Stephen
formations, approximately 1,000 feet of shales similar lithologically
at least to certain portions of the Ogygopsis shale. However, as no
fossils could be found in these beds, it remains problematical what
part, if any, of these shales is equivalent to the Ogygopsis shale.

The writer’s thorough investigation of Mount Stephen revealed
another important fact that had escaped the attention of previous
observers. This is the astonishing lateral change in thickness and
lithology of several units that takes place within the mountain from
northeast to southwest. Along Monarch Creek the Mount Whyte,
Cathedral, and Stephen formations possess normal thickness and li-
thology (section Nos. 17 and 26; also diagram in fig. 4). In the north-
west face from North Ridge to Fossil Gully, within a distance of
1 mile (fig. 3) the character of the section undergoes great changes.
The Mount Whyte formation thickens from 302 feet to 578 feet (sec-
tion No. 9), most of the thickening taking place in shale units in the
middle of the formation.

An even greater change affects the Cathedral formation. Along
Monarch Creek it consists entirely of dolomite. On the northwest
face of Mount Stephen, tongues of shale and thin-bedded limestone
begin to appear and rapidly thicken toward the southwest, while the
interfingering dolomite tongues thin out. Part of this change takes
place in inaccessible cliffs, but good observations of the strata can be
made from the opposite side of the valley. At Fossil Gully (section
No. 9) the dolomite in the Cathedral formation is reduced to two
tongues, 24 and 100 feet thick respectively, while the shale and thin-
bedded limestone units have acquired an aggregate thickness of the
order of 1,000 feet. The shaly character of the upper part of the
Cathedral formation makes it impossible to separate it here from the
overlying Stephen formation. Hence the two formations are left un-
divided in the Fossil Gully section, where their aggregate thickness is
2.254 feet.

Of particular interest are the limestone and shale units 2 and 3 near
the base of the Cathedral formation (section No. 9) that begin to
appear just west of the Monarch Mine and at Fossil Gully have an

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

MONARCH FOSSIL
CREEK GULLY

LZ ATLL L LLL SL LL LLL Ld

LI TIITIT iil
14454 LIIITIILIT IIT TT
LIZTIIIIILIILTL LTT LITT A

i LITIILITITITLTIT ITIL LTT A
554 LILTITILTIIT LIL TLL TT
TA

PLAS SILLS LLLSLL LTA
LIST ITTIITL IT
LEST TTT TTT TST IITA
USLESELLELELE SELLS
LEITLILT ITLL ILL Th
VILLI LILLE
SSS oo i ee

[TTA
(gsqaanasaasgaauauen (2)
TILIILILI LA =)
Yaaaa. SLISILILI LILES
wae, (=)
a
a
a
C
oO aa
uJ (ILI ILIILTLI LLL TA
LILILLILILILIIT LLL LT Ah
Fa CILILT Th fo) =
= SW
<x Tt)
OU © w
oO

iW
—
=
oe
=
—
=
=a
<a
a
a
—
(ep)

Fic. 4.—Section in the eastern and western parts of Mount Stephen. The '
thickness of the Cathedral formation at Monarch Creek is estimated.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 45

aggregate thickness of 191 feet. These beds carry remarkable fau-
nules not known anywhere else in the region. These units also include
gigantic boulders of dolomite or massive limestone, sometimes up to
30 feet in diameter (pl. 6). These boulders indicate that a portion of
Cathedral limestone or dolomite of normal lithology had been sedi-
mented at the northeast prior to deposition of the shales and thin-
bedded limestones at the southwest. Probably the massive carbonates
represent reefs of algal origin (note the abundance of Girvanella in
the undolomitized portions of the Cathedral formation in many sec-
tions). These reefs appear to have formed a steep submarine slope
along which large boulders, possibly incompletely consolidated, could
slide and become embedded in the argillaceous sediments being de-
posited southwest of the margin of the reef. In terms of the sedi-
mentary basins introduced by Walcott (1927) this conclusion may be
stated by saying that at the time when the Cathedral formation was
deposited, pure carbonates were sedimented in the Bow trough while
a considerable amount of fine clastic material was deposited in the
Goodsir trough. The conditions that produced the large conglomerate
boulders persisted for a considerably long time, since such boulders
occur occasionally through a thickness of 650 feet of strata.

MONARCH CREEK

The valley of Monarch Creek separates Cathedral Mountain from
Mount Stephen and was eroded along a fault line, Mount Stephen
being downfaulted with respect to Cathedral Mountain. In the upper
valley of Monarch Creek, a section of the Stephen formation is well
exposed and almost entirely accessible. The overlying Eldon dolomite
forms the vertical cliffs of the east face of the mountain. The Ca-
thedral formation is entirely exposed, but a section of this formation
was not measured because any accessible traverse would involve a
considerable horizontal distance, rendering the measurement inac-
curate. As far as it could be ascertained, the Cathedral in the east
face of Mount Stephen is entirely dolomitic.

The section of the Stephen formation described below was meas-
ured at a locality approximately 0.8 mile due east of the summit of
Mount Stephen. The strata are approximately horizontal, and the
Stephen-Eldon contact is exposed at an estimated altitude of 8,000
feet.

The Stephen-Eldon boundary must be arbitrarily defined within
certain limits, as there is no sharp lithologic change, and furthermore
near the top of the Stephen or the base of the Eldon there is the usual
lateral intergrading of limestone and dolomite.

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

MONARCH CREEK SECTION (NO. 26)
MippLe CAMBRIAN

Stephen formation

8. Limestone: fine-grained, black-gray, in beds 2-5 inches thick. a
Upper portion not accessible. Thickness estimated........... 80
7, Lamestone: datk gray... snassive.....< >. ,niee teens aes 5
G: Limestone’: a5 Unit (835 vic a s.scicre a as's 50 ena ealeiere Be eae ae 110
5. Limestone: gray, in thin, irregular beds with tan clay partings... II
4; ‘Shales"'siliceaus, “oray, “fissile; 22... 2 s/o. so enero e eee reer 5
gif -Limestoties as: unit 8. oe-20lds oo ils. ee bea ee leaner aon 7
2. Shale: in lower fourth siliceous, hard, dark gray, with lenses of
coarse, gray limestone; grading upward into siliceous, argilla-
ceous and calcareous shales, green-gray or tan, with some
thin-pedded limestones sae ciis oe cidoiee ohineecee ee OO eee 152
1, Limestone: gray, irregularly bedded: .\...44 s.2acseuwes esas nee ce 6
Total thickness of Stephen formation... 00.3. 0.00 0000.00. 376

Cathedral formation

1. Dolomite: coarsely crystalline, gray, tan-weathering, thick-bedded.
Contact with overlying Stephen formation very sharp... .Not measured

MONARCH MINE

A good section of the Mount Whyte formation can be studied in
the cliffs immediately below the Monarch Mine on Mount Stephen,
following the edge of the talus slope. If this section were examined
isolatedly, it would be difficult to decide where to place the upper
boundary of the Mount Whyte formation because the dolomitization
of the limestone at this locality extends into lower beds than usual.
Hence if the dolomite-limestone contact were taken as the Cathedral-
Mount Whyte contact, beds whose equivalents in other sections were
included in the Mount Whyte would be assigned to the Cathedral.
The criterion adopted was to trace the beds across the northwest face
of the mountain from the Monarch Mine to Fossil Gully. Then it is
clear that the lowermost 100 feet of dolomite at the Monarch Mine
represent the calcareous unit of the Fossil Gully section which carries
a faunule of the Plagiwra-Kochaspis zone and is everywhere in-
cluded in the Mount Whyte formation. Hence the Cathedral-Mount
Whyte contact was placed at the top of these 100 feet of dolomite, a
little below the level of the portal of the Monarch Mine, although
there is no definite change in lithology at this horizon.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

MONARCH MINE SECTION (NO. 17)
MippLe CAMBRIAN
Mt. Whyte formation

5. Dolomite: crystalline, gray, tan weathering, in beds 2-4 inches
thick. Thickness somewhat arbitrary because of indefinite
UP PEcabOundanvaOtelOnmiatl Ollsaeer esse aacteniiericia seit acai
4. Shale: siliceous, medium-grained, gray, tan weathering..........
3. Limestone and shale: crystalline, dark-gray limestone in beds 1-3
inches thick, alternating with siliceous, green-gray, tan-
weathering shale. Undetermined ptychoparid trilobites were
collected in limestone in top 5 feet (locality W17b).........
2. Shale: siliceous, fissile, green-gray, tan weathering..............
1. Shale and limestone: siliceous, gray shale alternating with dark-
gray, hard, siliceous, irregularly bedded limestone. A cranid-
ium of Amecephalus sp. was collected 18 feet above base
(locality W17a). This unit appears greatly to thicken west-
ward, corresponding to the combined units 1-3 of the Mount
Whyte formation in the North Gully section (combined thick-
ness of these units at North Gully 262 feet).................

Total thickness of Mount Whyte formation...............

LowER CAMBRIAN
St. Piran sandstone (Peyto limestone member)

1. Limestone, shale and sandstone: crystalline, sandy, gray limestone
in beds up to 2 feet thick alternating with sandstone and sili-
ceous shale. Olenellids and Bonnia common in the calcareous

47

Feet

100
35

40

52

75

302

NERVES By sei Sac nace naetontg eheros-a ions orm ue ene ore teas Not measured

Note—More abundant and better-preserved fossils were
collected from the top 6 feet of the Peyto limestone about
4 mile west of the Monarch Mine, directly above the second
of the three sheds of the Canadian Pacific Railway (locality
P18m, probably identical with U.S.N.M. locality 35f). The
identified species include:

Bouma feldensis (Wealcott))o\.1. 0s <nccam assess cc
Olenelisss Domeracctteto ern see Lier isie c cc
Onchocephalus cleon (Walcott) ............e00 r

Tits (CW AICOUE I Aretcoe esa cen os r
PAG PEME SV feta sseresstenctch nal els sa. uetsisrete ci aiaters eetiera cis ig
PASEURY Dike ( WV ANCORL) a2 aces acest pibic cc svacie sexe aise one r
Syspacephalus charops (Walcott) ............000 cc

NORTH GULLY AND FOSSIL GULLY

As explained in the general description of Mount Stephen, a section
extending from the top of the Lower Cambrian through the entire

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Middle Cambrian (and possibly a portion of the Upper Cambrian) is
exposed, within a single fault block, in the northwest face of Mount
Stephen between Fossil Gully and North Gully. For reasons of ac-
cessibility, the upper portion of the section (from the top of the
Stephen to the uppermost unit of the Mount Whyte inclusive) was
measured along the broad ridge that separates the northwest and west
faces of Mount Stephen, then down the northeast side of Fossil Gully.
A portion of the Mount Whyte formation was also measured in
another fault block on the southwest side of Fossil Gully. Finally, the
Mount Whyte formation exclusive of the uppermost unit was meas-
ured in North Gully. The strata, except in the immediate vicinity of
the fault in the bottom of Fossil Gully, are almost horizontal and well
exposed throughout.

FOSSIL GULLY SECTION (NO. 9)
Mipp_e CAMBRIAN

Eldon dolomite

Feet
4. Dolomite: crystalline, gray, tan weathering, massive...... Not measured
3. Limestone: subcrystalline, dark gray, thin-bedded. Thickness
SStitmateds as lors ores ais oars late rel te) eratarcre ete tees ayetetoucteretaiavae were torerene 30
2. Shale: siliceous, fine-grained, fissile, finely banded gray, green,
and tan. Viewed from a distance forms the “dark band” on the
west face of Mount Stephen mentioned by Walcott. Thickness
@SHimmiat dee oe (oy crete, stevie forsi oualaheyescsonssetanevolers sui siee ereisictetan-Varsien a lerene 150
1. Dolomite: crystalline, gray, tan weathering, thick-bedded. Thick-
Mess estimated. < sce = aarswlesa Sorerele sears ols eiae eine wrens ieee eeiternere 400
Total measured thickness of Eldon dolomite.............. 580
Stephen and Cathedral formations (undivided)
16. Limestone: subcrystalline, black-gray, more or less argillaceous,
mostly thin-bedded, partly purer and thicker-bedded (up to
4-5 inches). Upper part grading laterally to dolomite, making
Eldon-Stephen boundary arbitrary within 100 feet or more.
The thickness given is based on an average horizon for the
dolomite-limestone contact. Beds extremely fossiliferous 30-80
feet above base (locality Sok), most of the fossils occurring
on weathered surfaces of platy, argillaceous limestone layers. . 300
Fossits :
*Alokistocare paranotatum Rasetti........eeseeeeeeees r
Bathyuriscus adaeus Walcott.........eceeeceeeeeeees cc
Chancia cf. C. odarayensis Rasetti...........eeeeecees r
*Elrathina parallela Rasetti .......cceeecececececcees cc

Spinifera Rasettios's 0s .s'c sie clels vac eulses wigs c

NO.

15.

14.

5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

QOH N OSLER DIY 3 Cac cid DIOR IIo G AOR OIST AIOE r
PAGE liaiciey Her UO OLES NNidLGOLE are terse secieleroters ota tele Cc
Peronopsis columbiensis Rasetti........5..e.eseeeeees ce
Tonkinella stephensis Kobayashi..............eseeee. c
ZECAMIBOLD ES SDs esi citi are gle aid ores ds ambardha Qaele haved Oe a r

Limestone: subcrystalline, dark gray, with tan clay partings and
flakes, im irregular beds 1-2) inches thick:/J:.....0..-..-cce.-
Shale: siliceous, gray, fissile, tan weathering, with some inter-
stratified thin limestone beds. Also some gray, thin- and
irregular-bedded limestone with tan clay partings and flakes;
and some smooth-bedded, fine-grained, argillaceous limestone.
Unit forms cliff and is partly of difficult access. Thickness

ES EUTTAL CU Pert sreiaie Secicra eee ee Ren ee aIe a ahevererg shanaere
13. Limestone: subcrystalline, gray, thin- and irregular-bedded, with
tanrelay, partines-and! HAKes2cc cas sake dee hain cae eins eres 8
2s Shalesesiliceous miine-cralned se tavacaeaitemeeciiaecicisin eres) cereale
DD eIMEStOME': ASE UNI 03 = araretern overs e/stevere ashe Sie treveve alc, ole el he arena eve fatah ous’ o>
10. Shale: siliceous, gray, fine-grained, chunky..................0+:
g. Limestone: crystalline, hard, dark gray, in regular beds up to 6
inches thick. Forms good horizon marker between overlying
and underlying shales. Yielded rare, poorly preserved speci-
mens of Kootenia, Olenoides, and Oryctocephalus............
Sie Shall ec cash ari tat Operates fe esses aves teyasebnie de boiwheyis ol sr sseyoreys avshausis le sisletolere
7. Limestone: fine-grained, argillaceous, dark gray, in smooth beds
2-4 inches thick. Occasional lenses of dolomite conglomerate
includingelaneenbouldens. qemestetrceiee sete aeraersiebiers aac cree rns
6. Dolomite: crystalline, light gray, tan weathering, very massive.
Top of this unit is at level of great dolomite boulder emerging
PROM tals si DOLtOml OL HOssils Gully ese acess cress ls eels
5. Limestone: subcrystalline, hard, dark gray, mostly in beds 2-4
inches thick, with occasional massive, lenticular beds. Rare
fossils near base: (locality Cok) )o.55 cdicew sscmes sagen canacnes
FossILs :
Kootenia sp.
Zacanthoides sp.
4. Limestone: massive single bed of sandy, gray limestone..........
3. Limestone: subcrystalline, hard, dark gray, mostly in beds 2-6

inches thick, with massive, partly lenticular beds near base.
Large dolomite or limestone boulders are included in this unit
in the slopes of Middle Gully. Fossils collected at top of unit
(locality Coj’) and 35 feet below top (locality Coj).........
Fossixs (locality Coj’) :

Diraphora sp.

Wimanella sp.

Athabaskia sp.
*Chancia bigranulosa Rasetti

Poliella sp. No. 2

49

Feet

335

350

100

145

116

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Feet
Fossits (locality Coj) :
B GEVUSES CHAS GSD aye arcstie) <!ai ch eve oinicle mje eholateteistalete! ofetele(st storey fete r
Chancia bigranulosa Rasctttv. viscera isos ows aeleiad osierer c
Ogygopsis klotzi (Rominger)............2sccee. pueden
PRB. SD cts wakenthn oe Winle so aeinls sia east e fas seo r
Yohoaspis pachycephala Rasetti............- Rattner e r
*Zacanthoides sexdentatus Rasetti..,......eeeeeeeeeees c
2. Limestone and shale: silty, dark-gray limestone in irregular, often
lenticular beds; alternating with dark-gray, siliceous, and cal-
careous shales. Ogygopsis common throughout unit. Fossils
mostly collected 30-70 feet above base (locality Coh)......... 75
FossILs:
AlcrOthelé. Sp nies ails «is s'eis sie's'ss0 siavalareis wicleoyeisie nto eteene r
* GUNG LEG CLG INASELU Ms iors:cleieretoverets. ceelenalsteccnalerersh <r r
I GOTT HS EE IAOOR HOO OO DODO OOO OD CE ODCUOD On OOS r
Ogygopstis Rlots:. CROMIN ER) |e cee ocr sole lolele clel= cake) -1slet c
* SPIMOsa RASCH |. Git acct Racine mene eee T
PAOTCHA ISP ried ctastenios ofoncleracie tote ti ae ee Pht cleats c
Poltella Spc NOs Dosceeween ceed jecidse ee sch eae aes c
*Syspacephalus tardus, Rasetti..........0c.cccvccsasees c
*Yohoaspis pachycephala Rasettis.. 1.2 ..0iceseesn noice 2 r
ZGCanthoidess sp> actacyeianioe atlases Gare heir ee c
1. Dolomite: crystalline, gray, thick-bedded.................2.e000- 24
Total thickness of undivided Stephen and Cathedral forma-
LIONS whe, cyeusisie als aeereeeeainn slaps ah aaflet ahold yetoiseeyeteacte alate 2,214
Mount Whyte formation
11. Limestone: crystalline, dark gray, with tan clay partings and
flakes, in irregular beds 1-4 inches thick. Fossils common at
Variousw levels (localityanVV.OL) cies ciclate venice eater eieeerene 95
FossILs:
Amecephalus cleora (Walcott) .........cscscecccececs r
veld as pis. fur cata Nasctiteis. sta sere isiacieioeeeee eee c
Kochaspis evffclensts: Rasetti.'.ii. 2.3\00 oan 9e eee r
Mochiellacemaxeys, Rasettitva dice sae ee eens r
Piaguira cercops (Walcott) esas. se ecniene ete caters ik
wcomstometopus convenus Rasetti .2..seseceeeeeccenes r

collaris’ Rasett’ icc fon h weer eee ee r

Note.—The section was continued downward in another
fault block on the southwest side of Fossil Gully. The beds
in this block are upfaulted by 500 feet with respect to the
strata on the northeast side of Fossil Gully, hence strati-
graphically lower beds are easily accessible. This portion of
the section is connected to the preceding by means of the
readily recognizable contact of units 10 and 11. The strata
described in this portion of the section were also measured
in North Gully.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 51

e
Mount Whyte formation (continued) cid
LOM Shaler st CeO Uss 1 aly cos Vpryererevetars «ea lersseusteioreic ore, exsieyovehevesreea iotete: sia’ 40
g. Limestone: sandy, dark gray, in thin, irregular beds............. 6
SeeShalecgsiliceouss Ganka Cray tte cares oie ciicicle siete crsieleierajate steve: siavelstarere 32
7. Limestone: sandy, dark gray, thin-bedded in upper third, thick-
bedded sins lower stwo-thirdsijos > <\-elersieisieisieiorsiare scclewys's waves < es 18
Mount Whyte formation (Yoho shale lentil: type locality)
4-6. Shale: siliceous, fine-grained, fissile, dark gray. Faunule prob-
ably ranges throughout unit; fossils abundant 25-80 feet above
base: (locality: Wolk). dvcs'sveidics Aausysigtents coma tin Ae raleediows 270 « 135
FossILs:
PAV OMEN ESAS Ds warelos siekn oaisatassoteicicl seme oisialsietclsisterelsiseieweeiaie = c
SCONCU ESD? is, sclera tad oc Neier sieh ares ietreaie ee elbies cc
ALE OLMELE SPs ras, « Sisusrere ia sueparols) av ahe)steessyalyh shay oles) e)e1sises oils) aveisie r
AGRO OLS Paps aterds aiers sehets ose eis ayoles ser alsralusveleieNe sisi sysy/aouer als iovsis c
DLCLY ONAN SP sis Assavcretecarsteisieysvoahorem rains) olsletetole: sea\ciopeinvese ie r
JET TAPES We BE EE OCOTISR AS te ASO Onn nee SO Tee it
Amecephalus agnesensis (Walcott)..........cseeeees r
tGhancianstenomero pa RASeLtl eee eich teisis srersieieis cieielels rr
Oogygopsis kota (Rominge®) 5.506 ssee cen. deeds, ole rr
Oryctocephalusy sp) Now 2henaseese se oe eet en souls rr
*Poltelladenticulata: Rasettt, 22 22). /seg i eels es c
prima. CWaleott)) ccccki SoU ee acre Stee r
*Stephenaspis: bispinosa. RaAsettas, «0% caja si sieje'vy0)6 sa0's cc
*Syspacephalus gregarius Rasetti ...........seeeceees cc
laewigatus: Rasettt mack sees cae anekices r
perola (Walcott)iav. 26 eds eh we r
*Wenkchemma spinicollis Rasetti.........ccscceeccees cc
Total measured thickness of Mount Whyte formation...... 326

Note.——Lower beds are of difficult access in Fossil Gully.
The section was continued downward in North Gully, where
the highest measured strata are unit 10 of the Mount Whyte
formation. Hence there is a considerable overlap between
the two measured sections, units 4-10 having been measured
at both localities. The unit representing the Yoho shale lentil
in Fossil Gully was numbered 4-6 in order to designate by the
same numbers equivalent units in the Fossil Gully and North
Gully sections. In the latter the stratigraphic interval repre-
senting the Yoho shale is subdivided into three units, num-
bered 6, 5, and 4.

In North Gully the entire Mount Whyte formation is
excellently exposed, and all the beds are accessible except the
uppermost units which form a vertical or overhanging cliff.
The gully was eroded along a normal fault, the strata being
approximately horizontal on either side and the displacement
being 50-100 feet, with the southwest side downthrown, All

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the beds of the Mount Whyte formation, except the lowermost
unit, were measured on the northeast side of the gully, and
all the fossil localities are also located on this side. The slopes
east of the gully offer excellent exposures for collecting.

NORTH GULLY SECTION (NO. 16)
Mipp_E CAMBRIAN

Mount Whyte formation

Feet

11. Limestone: exposed in inaccessible cliff................0.. Not measured

10. Shale: siliceous, green-gray, tan weathering. Partly exposed in
inaccessible cliff. .Dhickness estimatedj2e- «5. ous s ne heres 50

9. Limestone: crystalline, dark gray, thin-bedded.................. a,

S. Shisha’ AS aah WO caress Waverarcty at eraterardve clan oneaccoratare ees as eee 45

7. Limestone: crystalline, hard, dark gray, in beds up to 5 inches
thick se caeae « siya) chat ay in tshan'a ap avon Gatton teabah ores Sahat hoa inet na Che ae eee 15

Mount Whyte formation (Yoho shale lentil)

6,. “Shales asy wiit 10 oiadinvs,< sisereettiers als ooh dior aist Gay ee Ober eee 10

5. Limestone and shale: limestone as unit 7, alternating with some
siliceous! shale? ss) </rsisjerecs acy ayeyec oe telat oh saelel le he Fae oP ae 24

4. Shale: siliceous, fine-grained, dark gray, fissile. Fossils common
about 20 feet above base and in thin limestone lenses 30 feet
above ibasea(locality > WiGls) jamie nae ineiaeee nee eee 70

FossILs :
Poatiella prima. (Walcott)... oreeuals. ata ese cere een r
Stephenaspis bispinosa Rasetti........cccccecccecceces c
Syspacephalus gregarius Rasetti ........ccecceccccees c
perolalCWaleott)igaesseeie see ee Cente r
Wenkchemmia spinicollis Rasetti..........ceceeeeveees cc
Mount Whyte formation (continued)

3. Shale and limestone: siliceous, dark-gray shale alternating with
fine-grained or finely crystalline, silty, dark-gray limestone in
irregular, often lenticular beds up to 3 inches thick, and rare
lenses of purer, crystalline limestone. Fossils common through-
out the interval; Ogygopsis apparently confined to upper 20
feet of interval, other species may range throughout. Locality
W16j in top 20 feet of interval, W16j” 40-50 feet above base,

W16j’ at undetermined horizon within unit.................- 104
Fossits (locality W16j) :
MirvGtfeta. SPoic>.. vitisic cies cke eermtortne a eine ae eee Peimaaseeoe
Ogygopsts klotst. CRominger,):. oc oss cas voces one os Cc
Oryctocephalus sp. NO. 122 ecco caus + acc neee taste oor r
*Prasella tuberculata IASG. ccs os os, spares coerce hence: i
Pohiella ci. P.. denticulata Raseth. ¢. oc... see sees set een Cc

Stephenaspis ci. S. bispinosa Rasetti......cccccccccees F

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 53

Feet
*Syspacephalus crassus Rasetti ......-.cccccccesecsees c
CES ore gaits) RASEUil. civ <liosialogayetenls r
* TOTICEPSPINASCHUM Dy aoreteieleiaie)-talelale) eistafeleiat ial c
Wenkchemnia sulcata Rasetti..........cceccccccecees c
Fossits (locality W16j”) :
Syspacephalus crassus Rasetti
laticeps Rasetti
*Wenkchemnia sulcata Rasetti
2. Shale: siliceous, medium-grained, fissile, dark gray. Trilobite
shields are common but the preservation is extremely poor.
Better-preserved fossils were collected from thin limestone
fenses aU lOcaILy) WUIOE)) ocx ece 2's t's tere Gine de navalenceaieie vase 16
FossILs :
Syspacephalus sp.
Wenkchemnia sulcata Rasetti
1. Shale and sandstone: siliceous, green and gray shale alternating
with thin layers of fine- to medium-grained, green-gray sand-
stone. Some thicker sandstone beds (up to 6 inches) in upper
part of unit. Considerable cross-bedding in lower part. At top
of unit, on northeast side of gully, beds of crystalline, dark-
gray limestone yielded fossils (locality W16h)............... 142
FossILs:
Acrotreta sp.
Syspacephalus laticeps Rasetti
Wenkchemnia sulcata Rasetti
Total measured thickness of Mount Whyte formation...... 488
LoweER CAMBRIAN
St. Piran sandstone (Peyto limestone member)
1. Sandstone and limestone: gray and brown, partly calcareous sand-
stone with lenses of sandy or almost pure, crystalline lime-
stone. Olenellid fragments abundant in the calcareous layers.. 30

CAMBRIAN FORMATIONS: LOWER CAMBRIAN
ST. PIRAN SANDSTONE
UPPER BOUNDARY OF THE FORMATION

It is not the purpose of this paper to revise the Lower Cambrian
formations. However, the upper portion of the St. Piran sandstone
was studied in most of the sections investigated in order to decide
where the St. Piran-Mount Whyte contact should be placed. The
result of this study was the decision to transfer from the Mount
Whyte to the St. Piran the basal limestone unit of the Mount Whyte
formation as hitherto understood. This unit is only 20 feet thick in
the type section of the Mount Whyte formation.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The St. Piran sandstone was reported by Walcott (1928) to be
approximately 2,600 feet thick in the type section, and there is no
doubt that the thickness actually exceeds 2,000 feet in most of the
sections in the Bow Range where the formation is entirely exposed.
In the Slate Mountains Deiss (1939) found the St. Piran to be much
thinner, 815 feet in the measured Ptarmigan Peak section. The St.
Piran chiefly consists of thick-bedded sandstone and quartzite with
some interstratified siliceous shale. Carbonate deposits are totally
absent from all except the top beds of the formation. Near the top,
calcareous material appears ; there is generally a gradual change from
purely siliceous to increasingly calcareous, highly ferruginous sand-
stone, then to more or less sandy limestone, and in many places to
pure, crystalline or sometimes oolitic limestone. Often calcareous sand-
stone or sandy limestone first appears as isolated lenses in the upper
portion of the sandstone. Olenellid fragments and Bonnia are almost
invariably present in the calcareous layers, which in many cases are
a true coquina of olenellid tests. It is clear both from stratigraphic
and faunal evidence that there is no break between the underlying
siliceous sandstone and these more or less calcareous (more rarely
dolomitic) beds, hence a formational contact placed between them (as
it was hitherto done) would be to a large extent arbitrary. On the
other hand, both stratigraphic and paleontologic arguments favor
placing a formational boundary at the top of the Olenellus beds. The
great faunal change that takes place between these beds and those
immediately overlying is discussed in another part of this paper. A
sharp change in lithology also generally occurs, the crystalline, sandy
limestone usually in thick beds being overlain by siliceous shale with
interstratified thin limestone or sandstone layers. In most of the
sections (see descriptions) the St. Piran-Mount Whyte contact as
here defined can be located within a few inches without ambiguity.
Furthermore, the erratically varying thickness of the Mount Whyte
(see discussion of formation) makes it appear likely that this unit was
deposited on a somewhat irregular surface. This fact, coupled with the
sudden faunal and lithologic changes, strongly suggests an uncon-
formity at the top of the Olenellus-bearing beds. Hence all argu-
ments favor placing the St. Piran-Mount Whyte boundary at the top
and not at the base of the Olenellus-bearing limestone or calcareous
sandstone, contrary to current usage. An analogous suggestion was
first made by Burling (1914, 1916).

The St. Piran-Mount Whyte boundary as here defined is at the
same time the Lower-Middle Cambrian boundary according to the

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 55

faunal definition later discussed in this paper. The writer does not
believe that series boundaries must necessarily coincide with uncon-
formities and formational boundaries, but in the present instance, in
the area investigated, this happens to be the case.

PEYTO LIMESTONE MEMBER

The calcareous beds at the top of the St. Piran sandstone are only
10-30 feet thick in most of the sections studied. However, in two
portions of the area investigated, one in the upper Bow Valley, the
other, probably of more limited extent, in the upper valley of Cataract
Brook, the calcareous beds attain considerable thicknesses, in places
exceeding 200 feet. At such localities the unit deserves a name and is
here designated as the Peyto limestone member of the St. Piran sand-
stone.

The type section of the Peyto limestone is chosen in the cliffs
formed by the west spur of Mount Thompson on the west side of the
upper Bow Valley, north of Bow Lake and south of Peyto Lake (see
section No. 15 for a detailed description). At the type locality the
Peyto limestone is 278 feet thick and is mostly crystalline or oolitic,
medium-gray, thick-bedded limestone, with some more or less cal-
careous sandstone and a little interstratified gray siliceous shale. Part-
ings in the limestone are poorly developed, hence when the strata are
exposed in cliffs they appear very massive. Olenellids and Bonmnia are
present at least in the upper portion of the limestone.

At Hector Creek (section No. 19) the Peyto limestone is well
developed, being about 200 feet thick, and presents the same lithology
as in the type section. Olenellid fragments are here extremely abun-
dant in most of the calcareous beds.

The Peyto limestone also has considerable thickness in the upper
valley of Cataract Brook. The thickness on Mount Odaray (section
No. 6) and Mount Schaffer (section No. 22) is 144 and 198 feet
respectively. In all the other measured sections the unit is much thin-
ner: 28 feet on Mount Niblock; 20 feet in the Mount Whyte section ;
24 feet on Mount Bosworth; 17 feet on Mount Victoria; 30 feet on
Mount Cathedral ; 22 feet on Popes Peak; 29 feet on Eiffel Peak ; 30
feet on Mount Temple; 43 feet on Pinnacle Mountain; 30-40 feet
at North Gully on Mount Stephen. Thicknesses at other localities
can be obtained from Deiss’ published sections (Deiss, like Walcott,
included these beds in the Mount Whyte formation). The resulting
values are 35 feet on Ptarmigan Peak, and 51 feet at Ross Lake. In
the Castle Mountain section, Deiss’ description does not suggest the

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

presence of beds assignable to the Peyto limestone. On account of
the gradual change from sandstone to limestone and the frequently
lenticular character of the limestone layers, exact thicknesses have
little significance.

A characteristic bed of the Peyto limestone that is present in many
sections is a massive sandstone, sandy limestone or dolomite, generally
weathering dark tan, including small, spherical or ovoidal concretions
of fine-grained limestone or dolomite approximately an inch in diame-
ter. These concretions resemble algal growths, but are more likely to
be inorganic. This bed was observed in the Ptarmigan Peak, Mount
Victoria, Eiffel Peak, Mount Temple, Pinnacle Mountain, Mount
Schaffer, and Mount Odaray sections. Since no other bed of similar
lithology has ever been observed in the Cambrian of the area, the con-
cretionary layer, when present, is an excellent horizon marker.

Fossils occur sparsely through the siliceous portion of the St. Piran
sandstone. The oldest faunule was discovered by Walcott (1928, p.
300) and collected again by the writer in sandstone near the base of
the formation, approximately 2,000 feet below the top of the Lower
Cambrian, near Vermilion Pass at the base of Storm Mountain. The
fossils are relatively abundant and fairly well preserved considering
the unfavorable lithology. The faunule includes “Wanneria” cf.
gracilis Walcott and a species of Rustella. The Peyto limestone is in
places highly fossiliferous and carries the Bonnia fieldensis faunule.

The name Peyto limestone derives from Peyto Lake, which is
approximately 2 miles north of the typical exposure.

CAMBRIAN FORMATIONS: MIDDLE CAMBRIAN
MOUNT WHYTE FORMATION
TYPE SECTION

The Mount Whyte formation was defined by Walcott (1908a) as
“alternating bands of limestone and siliceous and calcareous shale,”
the type locality being described as “Mt. Whyte, above Lake Agnes,
and eastern slope of Popes Peak, southwest of Mt. St. Piran.” In 1928
(Walcott, 1928, p. 251) the lithology was redescribed as “alternating
bands of gray and bluish-black limestone, siliceous and calcareous
shales, with some interbedded sandstones near the lower part.” The
type locality was stated to be the “north slope of Mt. Whyte and
southwest of Lake Louise.” A section of the formation under the
name of the “Lakes Louise and Agnes section” was only published
in 1928; the Mount Whyte formation described in this section being

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 57

apparently measured on or near Mount Whyte. The thickness of the
formation was given as 458 feet in the description of the section
(Walcott 1928, p. 302) but only 386 feet on page 251 of the same
paper.

Notwithstanding the somewhat ambiguous indication of the type
locality and thickness, it is clear what beds Walcott intended to include
in the Mount Whyte formation, and also that he meant the type sec-
tion to be in the general area of Mount Whyte. Deiss (1939, p. 998)
transferred the type section to Ptarmigan Peak for the following
reasons :

Examination of the Mt. Whyte section indicates that the formation there
includes Middle Cambrian limestones in the upper 100 feet or more and does
not contain as clearly defined or readily recognizable faunal zones as on
Ptarmigan Peak. Walcott not only never specified this section as typical of the
Mt. Whyte formation but nearly always referred to sections in other areas
when he discussed the formation or its fauna.

In the writer’s opinion, every one of the arguments by which Deiss
justified his change of the type section is invalid. The entire forma-
tion, once Walcott’s basal beds are transferred to the St. Piran, is
Middle Cambrian. The Mount Whyte formation in the Mount Whyte
area includes at least two fossiliferous horizons with characteristic
and well-identified faunules. The fact that Walcott gave two type
localities (both in the general area of Mount Whyte and within 23
miles of each other), or that he referred to other sections more often
than to the Mount Whyte section in discussing the formation, do not
appear sufficient reasons for transferring the type section to a rela-
tively distant area. Furthermore, Deiss’ choice of the Ptarmigan Peak
area for his new type section is particularly inappropriate, as on Ptar-
migan Peak the basal portion of the overlying Cathedral formation
(Deiss’ “Ptarmigan limestone’) is undolomitized and partly thin-
bedded, making it unusually difficult to place the upper boundary of
the Mount Whyte formation.

The formation was found to be particularly well exposed and ac-
cessible at two localities in the Mount Whyte area. The first (section
No. 2) is in the cirque above Lake Agnes, between Mount Whyte and
Mount Niblock, and approximately corresponds to Walcott’s first
type locality. The second locality is in the immediate vicinity of Plain
of Six Glaciers, 14 miles southwest of the upper end of Lake Louise
(section No. 3) and approximately corresponds to Walcott’s second
type locality. At Plain of Six Glaciers, the formation is exposed on
both sides of the gully between Popes Peak and Mount Whyte. The

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Mount Whyte formation was measured on the south side of the gully,
on the slope of Popes Peak, because of better exposure, but since
the overlying Cathedral and Stephen formations were measured on
the ridge of Mount Whyte, the section is referred to as the Mount
Whyte section.

For the purpose of reestablishing the type section of the Mount
Whyte formation in Walcott’s typical area, either the writer’s section
No. 2 or section No. 3 could be used. Each corresponds to one of
the type localities designated by Walcott, and both show closely simi-
lar successions of strata. The preference is given to section No. 3
because the formation is here more fossiliferous and more readily
accessible.

LITHOLOGY AND THICKNESS

The formation at the type locality at Plain of Six Glaciers, on the
east slope of Popes Peak, 14 miles southwest of the upper end of
Lake Louise, consists of 211 feet of gray or greenish siliceous shale,
crystalline or oolitic, dark-gray, mostly thin-bedded limestone, and a
little interstratified thin-bedded sandstone. Shale and limestone alter-
nate irregularly, but here as in all other sections shale prevails in the
lower part and limestone in the upper portion. Many of the limestone
beds include blebs and flakes of tan argillaceous material. The lower
part of the formation includes beds of shale with thin interstratified
nodular layers of limestone.

The St. Piran-Mount Whyte contact and the reasons for transfer-
ring Walcott’s basal beds of the Mount Whyte to the St. Piran have
been discussed. In the type section, the St. Piran-Mount Whyte
boundary as here defined is extremely sharp. Furthermore, olenellids
occur up to the very top of the Peyto limestone member of the St.
Piran, while a totally different Middle Cambrian faunule (the Poliella
prima faunule) occurs in the Lake Agnes shale lentil of the Mount
Whyte formation, 22-27 feet above the top of the Olenellus-bearing
beds.

The Mount Whyte formation in Walcott’s “Lakes Louise and
Agnes section” was measured “‘in the vicinity of Lake Agnes” (Wal-
cott, 1928, p. 302) hence the locality closely corresponds to the one
where the writer measured his Mount Niblock section. The writer’s
measured thickness of the Mount Whyte formation on Mount Nib-
lock is 281 feet. Walcott, in the description of the section, gave a
thickness of 458 feet. The excess is due to an obvious error made by
Walcott in reporting a thickness of 187 feet for the Olenellus-bearing
beds (here assigned to the Lower Cambrian Peyto limestone) which

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 59

in reality are only 28 feet thick. The portion of Walcott’s section
which is here assigned to the Mount Whyte formation hence has,
according to Walcott, a thickness of 271 feet in close agreement with
the writer’s measured value of 281 feet.

The thicknesses of the Mount Whyte formation as measured in
different sections are reported in table 1. In the case of the sections
measured by Deiss, the thickness of the formation was reduced
by subtracting the beds assignable to the Peyto limestone. Deiss’
Ptarmigan Peak section required special attention. Here Deiss rec-
ognized the “Ptarmigan limestone” which, in the writer’s opinion,

TABLE I.—Thickness of the Mount Whyte formation in 18 sections

Author
Section Thickness (D = Deiss,
No. Locality Feet R = Rasetti)
Biss a orerotia 8 in ats Mount Niblock 281 R
2 ae rir iey Mount Whyte 211 R
feito ie ane Ross Lake 264 D
ae cls SORS ees Mount Bosworth 380 R
Oy ct rere aes Seon Mount Odaray 161 R
Gal O ace aso: North Gully-Fossil Gully 578 R
Teieiete cole sapere she Ptarmigan Peak 405 D
Seite aie ios) Bow Lake 503 R
LZ arers . Aoa ee ree Monarch Mine 302 R
EO Gants esa ee Hector Creek 480 R
BO on see See cee Eiffel Peak 58 R
Dera or esc reve Saree Mount Temple 108 R
DP Re Ee ete ee Mount Schaffer 112 R
Ze caste ah EET Mount Cathedral 155 R
BAN cents Noe eee Popes Peak 226 R
2 Evetati BS atelier « Mount Victoria 63 R
Drege stots Bevasece, ular Pinnacle Mountain 98 R
Castle Mountain 145 D

must be redistributed between the Mount Whyte and Cathedral for-
mations. After examination of the section on Ptarmigan Peak, the
writer decided to include the basal 165 feet of Deiss’ “Ptarmigan
limestone” in the Mount Whyte, the rest in the Cathedral.

The thicknesses reported in table 1 are also represented on a map
(fig. 5). The formation is obviously thicker in the northern part of
the area investigated, the upper Bow Valley (not comprised in the
map) but elsewhere the thickness varies erratically, reaching a mini-
mum of 58 feet on Eiffel Peak and a maximum of 578 feet in the Fos-
sil Gully section on Mount Stephen. The Fieldaspis bilobata faunule
collected 22-28 feet above the base of the formation on Eiffel Peak is
essentially the same as the Fieldaspis furcata faunule found in the

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

uppermost 95 feet of the 578 feet of strata at Fossil Gully. The paleon-
tologic evidence seems to indicate that the youngest beds of the for-
mation are approximately of the same age everywhere, while older
strata that are present in the thicker sections are absent where the
formation is thinner. There seems to be an axis of minimum thickness
(indicated in fig. 5) running in a northwest-southeast direction from
the bottom of the Yoho Valley to Eiffel Peak, the Mount Whyte
formation thickening on either side of this axis. Northeast of the axis
the Middle Cambrian formations have the typical facies of the Bow
trough. Southwest of the axis at least some of the Middle Cambrian
formations have different facies, and we may include at least part
of this area in Walcott’s Goodsir trough. Southwest of the fault line,
parallel to the above-mentioned axis, indicated in figure 5, no Middle
Cambrian sediments come to the surface. According to the suggested
interpretation, this anticlinal axis developed in late Early Cambrian
time and represents a line of maximum unconformity between the
Lower and Middle Cambrian sediments.

Although the present work does not include the Mount Assiniboine
area, a comment on the unit there defined by Deiss as the Naiset
formation is in order. Deiss used the name for 475 feet of siliceous
and calcareous shales and argillaceous limestones overlying the Gog
formation and underlying the Cathedral dolomite. Walcott had as-
signed these strata to the Mount Whyte formation. Deiss (1940)
used a new name because fossils collected from the upper part of
the Naiset were considered of Medial Cambrian age, in contrast
with the Early Cambrian age of the fauna of the Mount Whyte for-
mation. If the Mount Whyte formation in the Bow Range and the
Naiset formation in the Mount Assiniboine area were proved to be a
continuous lithogenetic unit, there would be no need for separate
formational names even if this unit were found to transgress the
Lower-Middle Cambrian boundary. However, this nomenclatural
problem does not arise because there is no faunal difference between
the Mount Whyte and Naiset formations, both units including ex-
clusively fossils of Medial Cambrian age. An idea of the fauna of
the Naiset formation can be obtained from the fossils collected by
Walcott from his locality 62w and which are preserved in the U. S.
National Museum. The species include:

*Amecephalus cleora Walcott.
*Caborcella skapta (Walcott).
*Olenoides damia (Walcott).
*“Ptychoparia”’ gogensis Walcott.

MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 61

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62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

The first two species are typical of the Plagiwra-Kochaspis fauna
of the Mount Whyte formation. Olenoides was not found by the
writer in the Mount Whyte formation, but is known elsewhere to
range from the upper Lower Cambrian through the Middle Cambrian.
The last species is a generalized ptychoparid of little stratigraphic
significance. Deiss’ fossil list for the upper part of the Naiset con-
firms that the Naiset formation is an equivalent of the Mount Whyte.
Since the lithology is not essentially different, there seems to be no
need for a separate formational name.

MOUNT WHYTE—CATHEDRAL CONTACT

The Mount Whyte-Cathedral contact is to a greater or lesser extent
arbitrary in most of the sections, suggesting that continuous deposition
was taking place. The essential changes are an upward decrease of
the amount of clastic material and the thicker character of the beds.
Where these changes coincide with a transition from limestone to
dolomite, a fairly sharp boundary results (e.g., in the Mount Whyte
section). Where the lowermost portion of the Cathedral is calcareous,
the contact is more arbitrary (e.g., in the Ptarmigan Peak section).
At many localities (e.g., at the Kicking Horse Mine) the boundary
between limestone and dolomite does not follow the bedding planes,
oscillating within a vertical range of 10-50 feet even within short
horizontal distances. At the same time, the thickness of the beds in-
creases and the distinctness of the partings decreases but slowly and
gradually. In such sections, the Mount Whyte-Cathedral contact is
arbitrary within certain limits. A case of dolomitization of the lime-
stone in the upper portion of the Mount Whyte formation occurs
at the Monarch Mine on Mount Stephen. Here the top 100-feet lime-
stone unit of the Mount Whyte is represented by dolomite. The
reason for including these beds in the Mount Whyte rather than the
Cathedral is that the unit can be traced westward across the face of the
mountain, and at Fossil Gully becomes a limestone unit (unit No. 11
of the Fossil Gully section) which carries the Fieldaspis furcata
faunule characteristic of the upper Mount Whyte.

The character of the Mount Whyte-Cathedral contact is described
in the sections.

LAKE AGNES SHALE LENTIL

The name “Lake Agnes shale” was mentioned by Walcott (1917¢c,
p. 93) incidentally in describing a trilobite and never defined or used
again. Notwithstanding the small thickness and the limited areal ex-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 63

tent of the unit thus designated, the term deserves to be revived to
provide a name for the highly fossiliferous shale bearing the Poliella
prima faunule.

As far as known, the Lake Agnes shale is a lentil developed only
in the Mount Whyte area, hence included in the type section of the
Mount Whyte formation. It was observed at Plain of Six Glaciers
(section No. 3) and on Mount Niblock (section No. 2). At both
places, it consists of 5 feet of very fine-grained, green-blue-gray, fis-
sile siliceous shale. Its position is 22-27 feet above the base of the
formation at Plain of Six Glaciers and 15-20 feet above the base on
Mount Niblock. The former locality, where the unit is better exposed,
more readily accessible, and more fossiliferous, is chosen as the type
locality. This is also the type locality for the Mount Whyte formation.

In the other sections, the same position is occupied by coarser
siliceous shales, often with thin interstratified sandstone layers, and
the beds are unfossiliferous.

The name derives from Lake Agnes, which occupies the cirque
between Mount Niblock and Mount Whyte.

YOHO SHALE LENTIL

This is also a local unit, appearing in the middle portion of the
Mount Whyte formation as a result of the westward thickening of
the strata on Mount Stephen. It is known only in the northwest face
of Mount Stephen, from a little west of the Monarch Mine to Fossil
Gully. The only accessible exposures are in North Gully and Fossil
Gully.

At the type locality, in a fault block on the southwest side of Fossil
Gully, the Yoho shale lentil consists of 135 feet of dark-gray, fine-
grained, fissile siliceous shale. At North Gully, it includes a lower
70-feet unit of similar shale, followed by 24 feet of dark-gray, crystal-
line, mostly thin-bedded limestone interstratified with siliceous shale,
in turn overlain by 10 feet of shale; hence possesses a total thickness
of 104 feet.

The Yoho shale is an example of the peculiar character of several
Middle Cambrian formations in the western part of Mount Stephen.
It may be a tongue extending from unexposed strata existing still
farther west, but owing to the very limited exposure it will be desig-
nated as a lentil. It will be noticed that the Yoho shale is exposed at
the boundary between Walcott’s Bow and Goodsir troughs.

The Yoho shale carries the Stephenaspis bispinosa faunule that
seems to be approximately equivalent to the Poliella prima faunule of

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

the Lake Agnes shale. The unit is next only to the Ogygopsis shale
in the abundance of fossils. The fact that such a remarkably fossilif-
erous shale had not been discovered earlier shows the superficial
character of the previous investigation of Mount Stephen. The name
derives from the Yoho River, which flows into the Kicking Horse
River near the base of Mount Stephen.

CATHEDRAL FORMATION
TYPE SECTION

The Cathedral formation was originally defined as “‘massive, arena-
ceous and dolomitic limestone’ (Walcott, 1908a) and its type locality
was given as “Cathedral Mountain and Cathedral Crags, east of
Mount Stephen and southeast of Mount Bosworth.” When the forma-
tion was redefined (Walcott, 1928) the type locality was changed to
“south face of Mount Bosworth between Hector and Stephen on the
Canadian Pacific Railway, also finely exposed in Cathedral Crags
east of Mount Stephen.” Lithologically the formation was rede-
scribed as “more or less arenaceous, thick and thin layers of hard,
gray limestone forming fine cliffs when not broken down.”

The change in type locality was opportune, since the strata are well
exposed but largely inaccessible in Cathedral Mountain and Cathedral
Crags, hence a section was never measured there. Deiss (1939)
adopted the Mount Bosworth section as typical and later (Deiss,
1940) measured it on the east ridge of the mountain. Unfortunately,
at that locality some of the top beds are cut off by a fault, hence Deiss’
measured thickness of 1,085 feet is not complete. The entire forma-
tion is present farther west in the south slope of Mount Bosworth,
but parts of the beds are covered by talus and soil. It will be shown
that the thickness of the Cathedral formation is fairly constant within
a relatively large area and averages 1,200 feet in the sections nearest
to Mount Bosworth, hence the thickness missing in Deiss’ section
is not great.

Walcott (1917a) subsequently named another unit, the Ptarmigan
formation, supposed to occupy an interval above the Mount Whyte
and below the Cathedral. At the type locality, on the south slope of
Ptarmigan Peak in the Slate Mountains, the Ptarmigan formation
was stated to include 516 feet of limestone. When a first detailed
description of the Ptarmigan Peak section was later given (Walcott,
1928, p. 278), the Mount Whyte-Ptarmigan boundary was raised
go feet, thus reducing the thickness of the Ptarmigan to 426 feet.
Deiss (1939) recognized the “Ptarmigan limestone” in the Ptarmigan

ee

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 65

Peak section and gave its thickness as 460 feet. He placed the Ptar-
migan-Cathedral contact at the boundary between the underlying lime-
stone and the overlying dolomite. By this procedure the thickness of
the Cathedral formation in the Ptarmigan Peak section was reduced
to 620 feet. Deiss could find only 105 feet of Ptarmigan limestone on
Castle Mountain, 195 feet doubtfully assigned to the formation on
Mount Bosworth, and none at Ross Lake. The writer does not
recognize the “Ptarmigan limestone” for reasons that are fully ex-
plained later, hence the beds referred to the Ptarmigan by Deiss are
redistributed among other formations. In the Ptarmigan Peak section,
the basal 165 feet of Deiss’ Ptarmigan are reassigned to the Mount
Whyte formation ; the remaining 395 feet are typical of the Cathedral
(see discussion of lithology) and are included in this formation, thus
bringing it to a more normal thickness of 1,015 feet. The 195 feet
of Deiss’ doubtful Ptarmigan on Mount Bosworth were already re-
assigned to the Mount Whyte formation. The writer has not examined
the Castle Mountain section, but Deiss’ description of the lithology
makes it clear that the 105 feet which he assigned to the Ptarmigan
belong to the Cathedral.

LITHOLOGY AND THICKNESS

The Cathedral formation consists almost entirely of carbonates,
mostly in the form of massive limestone or dolomite.

Deiss designated the formation as the “Cathedral dolomite” because
of the prevalently dolomitic character of the rock in his measured
sections. In the type section on the east ridge of Mount Bosworth,
the formation is stated to include 830 feet of dolomite, 247 feet of
limestone, and 8 feet of shale. This, however, does not give a correct
picture of the average lithology, as many dolomite units grade later-
ally into limestone. On the south slope of Mount Bosworth, 1 mile
west of Deiss’ measured section, part of the strata are covered, but all
the rock that can be seen, certainly more than half the thickness of
the formation, is calcareous. On the east ridge itself, a few hundred
feet north of the crest, several intervals that were described by Deiss
as dolomite change to limestone. This lateral intergrading of lime-
stone and dolomite was observed in almost all sections where the
formation was studied (see also Deiss, 1940, pp. 775-776). The dolo-
mite-limestone-boundary may follow the bedding for considerable
distances, as in the south slope of Ptarmigan Peak, where this fact
led Deiss to recognize the “Ptarmigan limestone.” However, the
writer followed the contact around the mountain, in the cliffs of the

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

northwest face and above the lake in the cirque north of Ptarmigan
and Pika Peaks. At that locality, the limestone-dolomite contact
shifts to a lower horizon, suddenly reducing the “Ptarmigan lime-
stone” from 395 to 100 feet. It is obvious that this contact cannot
be used as a formational boundary. At other localities, lenses of lime-
stone in dolomite, lenses of dolomite in limestone, interfingering
tongues of limestone and dolomite, and other irregular forms of the
limestone-dolomite contact can be observed. The contact is sometimes
so sharp that hand specimens were collected where the lateral change
of limestone into dolomite takes place within less than one inch. The
irregular dolomitization of a dark-gray limestone in the upper Ca-
thedral formation on Mount Temple is illustrated by two photographs
on plate 5.

The better to illustrate the lithology and thickness of the Cathedral
formation, data from Deiss’ and the author’s measured sections are
collected in table 2. The formation is naturally divided into “lower
Cathedral” and “upper Cathedral” by the Ross Lake shale. Where
this shale is absent, the formation is left undivided. The table
includes the respective thicknesses of limestone and dolomite sepa-
rately for the lower Cathedral, the upper Cathedral, and the formation
as a whole. Inspection of the table shows: (1) Limestone is present
in considerable amount, in some of the sections prevailing over dolo-
mite. (2) Dolomite is, on the average, just as abundant in the lower
as in the upper part of the formation. These data clearly show the
invalidity of the “Ptarmigan formation” based on the distinction be-
tween limestone and dolomite. The mixed lithology also makes it
advisable to use the term “Cathedral formation” rather than “Cathe-
dral dolomite.”

Table 2 also shows the remarkably constant thickness through most
of the area, where the formation averages 1,200 feet; excepting the
Castle Mountain section where the formation, measured by Deiss, is
considerably thinner. The Ross Lake shale, when present, occurs on
the average somewhat over 400 feet from the base, hence the lower
Cathedral represents from one-third to two-fifths of the formation.
Only in the Bow Lake section, at a considerable distance from the
other measured sections of the formation, the Ross Lake shale divides
the Cathedral into almost equal parts.

The limestones of the Cathedral formation are mostly thick-bedded,
often massive without apparent partings for considerable thicknesses,
medium to dark gray, sometimes mottled, finely crystalline to fine-
grained. Algal concretions (Girvanella) are frequently observed.

MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

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68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Many beds contain stringers or irregular inclusions of less soluble
material weathering in relief. When the limestone grades laterally
into dolomite, two changes are almost invariably apparent: the dark
pigment disappears, producing light-gray or cream-colored dolomites
even when the corresponding limestone is black-gray ; and the texture
becomes much more coarsely crystalline. These crystalline dolomites
almost invariably weather to a light- to dark-tan color and form
rough surfaces under solution. Sometimes limestone is partially
dolomitized producing a mottled appearance. The thin- or thick-
bedded character of the strata is usually unaffected by the lateral
change of limestone into dolomite.

The writer will not attempt to decide the moot question whether
the dolomite resulted, as Deiss (1940, p. 776) is inclined to believe,
from simultaneous deposition of limestone and dolomite, or an original
limestone was secondarily altered to dolomite. In certain cases there
is some evidence for secondary dolomitization, for example, in the
vicinity of the Monarch Mine, situated in the lowermost portion of
the Cathedral formation, where the unusually strong dolomitization
affects even the uppermost portion of the Mount Whyte formation
and suggests some connection with the formation of lead and zinc
deposits (Allan, 1914). At other localities the writer got the impres-
sion that the Cathedral is more apt to be dolomitized near faults or
other structural disturbances. If this fact were ascertained, it would
prove the secondary origin of at least some of the dolomite. However,
the observations are not extensive enough to show that the effect is
real.

The facies of the formation on Mount Stephen deserves special
discussion. As it was mentioned in the description of Mount Stephen,
in the eastern part of the mountain the Cathedral formation is entirely
dolomitic, and although the thickness was not measured, it appears
to be normal. Proceeding westward, units of shale and thin-bedded
limestone appear and rapidly thicken at the expense of the interfinger-
ing dolomite tongues. At Fossil Gully, 1 mile west of the Monarch
Mine, the Cathedral dolomite is reduced to two units, 24 and 100
feet thick respectively, all the rest being replaced by strata of the
new facies. At the same time the boundary between the Cathedral and
the Stephen becomes indefinite, hence 2,214 feet of beds overlying the
Mount Whyte formation and underlying the Eldon dolomite had to
be assigned to the undivided Cathedral and Stephen formations. The
lower part of these 2,214 feet of strata is fossiliferous and yields
faunules unknown elsewhere in the area. It is clear that within Mount

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 69

Stephen we observe a sharp transition between two different facies
of the Middle Cambrian deposits. A great change was known to
affect the Upper Cambrian sediments at nearly the same boundary,
the prevalently calcareous formations of the Bow trough (Walcott,
1927) being replaced by the shaly sediments of the Goodsir trough.
Now for the first time the evidence presented shows that the distinc-
tion between these two sedimentary basins was already developed in
Medial Cambrian time. Unfortunately only an extremely narrow
area of the Middle Cambrian sediments at the margin of the Goodsir
trough is preserved in Mount Stephen. Farther southwest only
younger formations are exposed and structural complications ac-
companied by metamorphism render a study of the stratigraphy
difficult. Figure 4 represents in diagrammatic form the succession of
beds in both the eastern and western parts of Mount Stephen.

CATHEDRAL—STEPHEN CONTACT

The Cathedral-Stephen contact is one of the best-defined forma-
tional boundaries in the area. The thick-bedded limestone or dolomite
of the Cathedral is usually overlain by thin-bedded, argillaceous, platy
limestone of the Stephen and the contact is sharp in most of the
sections. Furthermore, these basal beds of the Stephen are readily
recognized by the presence of the Glossopleura boccar faunule. In
some of the sections (Mount Odaray, Park Mountain, Monarch
Creek) the Glossopleura fauna seems to be missing, the Stephen for-
mation beginning with the shaly unit that usually forms its middle
portion. The Cathedral-Stephen contact is even sharper in these cases.

The time interval between the deposition of the uppermost Cathe-
dral and the lowermost Stephen in the sections where the latter for-
mation has normal lithology does not seem very great, since in the
Skoki Valley section (No. 13) a Glossopleura merlinensis faunule,
collected from the upper Cathedral, is closely related to the Glosso-
pleura boccar faunule of the basal Stephen. An unconformity must
be present between the Cathedral and the Stephen where the latter
does not include beds of the Glossopleura zone.

ROSS LAKE SHALE MEMBER

The shale now bearing this name was known for a long time only
from drift blocks collected by Walcott at the base of Mount Bos-
worth. These blocks yielded the abundant Albertella bosworthi faunule
and were believed by Walcott to belong in the Mount Whyte forma-
tion. Burling (1916) first located the shale in place in the Cathedral

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

formation on Mount Bosworth. Walcott (1917a, p. 4) named and
defined the shale as follows:

The Ross Lake shale member of the Ptarmigan formation (Albertella zone).—
A name proposed for the fine siliceous shale carrying the Albertella fauna in the
Ptarmigan ? formation.

Type locality—tIn cliffs above Ross Lake.

Character—Dark gray, fine siliceous shale with local fillets and thin layers
of gray limestone.

Thickness.—From 7 to 11 feet.

Walcott’s reference of the Ross Lake shale to the “Ptarmigan
formation” needs no further discussion at this point, as the position
of the shale has been explained in describing the Cathedral formation.

The Ross Lake shale is generally a dark blue-gray, fine-grained,
fissile siliceous shale. At some localities (e.g., Eiffel Peak) the in-
terval is represented by alternating thin, irregular layers of limestone
and siliceous shale. In other sections (e.g., Mount Bosworth, Ross
Lake, Mount Temple) the unit is prevalently siliceous, but still in-
cludes nodules, lenses or thin layers of gray, finely crystalline lime-
stone. The Ross Lake shale does not exceed 8 feet in thickness, yet
is surprisingly widespread for such a thin unit. It extends at least
from Mount Temple at the southeast to Bow Lake at the northwest.
Furthermore, the writer found the shale present near Sunwapta Pass,
at the southern end of Jasper Park, 52 miles northwest of Mount
Temple. Whether the unit is continuous throughout this distance or
consists of isolated lenses remains undetermined.

The only sections where the shale was searched for and not found,
although the Cathedral formation is completely exposed, are the
Ptarmigan Peak and Mount Odaray sections. The shale is probably
also absent in the eastern part of Mount Stephen, in the valley of
Monarch Creek.

The position of the Ross Lake shale in the Cathedral formation
and its thickness in various sections are given in table 2.

The Ross Lake shale is highly fossiliferous in almost all outcrops
and carries the Albertella bosworthi faunule described under the fauna
of the Albertella zone.

STEPHEN FORMATION

TYPE SECTION

The Stephen formation was first defined by Walcott in 1908 (Wal-
cott, 1908a, pp. 3-4) and described as follows:

Type locality——Bluish gray and greenish gray limestone and shale band about
2,700 feet up above railroad track on the north and east sides of Mount Stephen.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 71

Thickness—On Mount Stephen, 562 feet, with 150 feet of local development of
Ogygopsis shales at the summit.

The same year Walcott (1908c) published a detailed section of the
formation on Mount Stephen, in which he gave the thickness of the
formation as 712 feet and thus clarified his earlier statement.

In 1928 (Walcott 1928, pp. 315-319) he again discussed the Stephen
formation. The type locality was now stated to be “northwest side
of Mount Stephen, above Field.” The thickness was again stated to
be 712 feet (Walcott, 1928, p. 247) and the lithology described as
follows:

On Mount Stephen, calcareous, siliceous, and finely arenaceous shales with
more or less argillaceous matter, superjacent to a thick band of bluish-black
limestones in thin layers, which in turn is underlain by a series of oolitic and
gray limestones alternating with bands of dolomitic limestone and a few bands
of siliceous shale.

On pages 315-319 of the same paper, in the detailed description of
the Mount Stephen section, 190 feet of “bluish-gray limestone with
bands of dark siliceous shale in lower portion” were transferred from
the Eldon to the Stephen, thus bringing the latter formation to a
thickness of 912 feet.

In describing Mount Stephen, the writer already pointed out that
Walcott’s section on that mountain was not derived from the measure-
ment of a continuous succession of strata at any one locality, but is
instead a composite section, as evidenced by the inclusion of the
Ogygopsis shale in the Stephen formation and the description of the
Cathedral formation as dolomitic. In the western part of the moun-
tain, where the Ogygopsis shale crops out, the Cathedral is not ex-
posed, and at the nearest exposure, above Fossil Gully, is not dolo-
mitic but almost entirely represented by thin-bedded limestone and
siliceous shale.

Furthermore, the anomalous lithology of the Cathedral formation
in the western part of Mount Stephen (see discussion of formation)
makes even a continuous measured section at that locality (the writer’s
Fossil Gully section) unsuitable for defining the Stephen formation,
as the Cathedral-Stephen boundary is not clearly marked either litho-
logically or faunally. The section in the east face of Mount Stephen
(Monarch Creek section) is unsuitable because the Stephen forma-
tion at that locality is unfossiliferous and the top portion is inacces-
sible. Moreover, the succession of strata that might be defined as the
Stephen formation both in the eastern and western parts of Mount
Stephen is not typical of that portion of the stratigraphic column in
the area investigated.

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The conclusion is that the type section of the Stephen formation
must be established elsewhere than on Mount Stephen. There are
several sections that have been accurately measured either by Deiss (on
Castle Mountain, Ptarmigan Peak, Mount Bosworth) or by the writer
(on Mount Odaray, Park Mountain, Mount Whyte, Mount Temple)
and where the formation is excellently exposed. In the Mount Odaray
and Park Mountain sections, the Stephen is not typical for the area,
these localities being situated at the margin of the “Bow trough”
where the facies of the Middle (and Upper) Cambrian sediments
undergoes a rapid lateral change. Among all the other sections which
are free of this objection, the writer prefers to use the Mount Bos-
worth section as typical because it was one of the first measured by
Walcott (1908, pp. 204-207; 1928, pp. 308-314), has been often
cited in the literature, and is nearest to Mount Stephen. The Mount
Bosworth section was accurately measured by Deiss (1940) on the
south slope of the mountain.

LITHOLOGY AND THICKNESS

At the type locality in the south slope of Mount Bosworth, the
Stephen formation as measured by Deiss consists essentially of three
units. In ascending order, these are: (1) 134 feet of dark-gray, fine-
grained to medium-crystalline limestone, more argillaceous and thin-
bedded in the lower part, purer and thicker-bedded in the upper
portion; (2) 185 feet of gray and greenish siliceous and calcareous
shales with some interstratified thin-bedded limestone; (3) 151 feet
of finely crystalline or fine-grained, black-gray, mostly thin-bedded
limestone. Thus the total thickness of the formation on Mount Bos-
worth measured by Deiss is 470 feet.

The same three lithologic units are recognizable in the sections on
Castle Mountain and Ptarmigan Peak (Deiss, 1939) and in those
measured by the writer on Mount Whyte and Mount Temple. The
total reported thickness of the formation is 285 feet on Castle Moun-
tain, 315 feet on Ptarmigan Peak, 495 feet on Mount Whyte, 499
feet on Mount Temple, 440+ feet on Mount Odaray, 548 feet on
Park Mountain, and 376 feet in the eastern part of Mount Stephen
(Monarch Creek). From these data the thickness appears more vari-
able than it really is, because in Deiss’ Castle Mountain and Ptarmigan
Peak sections thin-bedded dolomite beds that were included in the
basal Eldon probably represent the dolomitized equivalent of a por-
tion of the limestone included in the upper Stephen in the other sec-
tions. The writer would transfer from the Eldon to the Stephen 85

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI F/3}

feet of dolomite in the Castle Mountain section and 170 feet in the
Ptarmigan Peak section, thus bringing the total thickness of the for-
mation in those sections to 370 and 485 feet respectively (see discus-
sion of Stephen-Eldon boundary).

On Park Mountain, Mount Odaray, east Mount Stephen (Monarch
Creek) and west Mount Stephen (Fossil Gully) the character of the
formation is anomalous in that the basal limestone unit seems to be
absent, the middle shaly unit immediately overlying the Cathedral
dolomite or limestone (in the Fossil Gully section the upper Cathe-
dral is replaced by shale and the Cathedral-Stephen boundary remains
undefined). Faunal evidence also supports the conclusion that, at
least on Park Mountain and Mount Odaray, sediments corresponding
to the time interval in which the basal limestone was formed elsewhere
were not deposited. All these localities where the Stephen formation
has anomalous character lie at the southwestern margin of Walcott’s
Bow trough (Walcott, 1927).

The faunas are described in another part of this paper; however,
it is opportune to discuss briefly their distribution within the forma-
tion. Wherever the Stephen has a basal limestone unit, it always
yielded both to Deiss and the writer faunules of the Glossopleura
zone. These beds are highly fossiliferous in many of the sections, but
yield little besides specimens of Glossopleura and Polypleuraspis. The
overlying, mostly shaly unit was found by the writer to be almost
barren. On Mount Odaray, where this unit directly overlies the Ca-
thedral dolomite, it yielded near the base a faunule with Ehmaniella
and Solenopleurella, certainly younger than the fauna of the Glosso-
pleura zone. Hence the conclusion that the Stephen at this locality
does not include beds of the Glossopleura zone. The uppermost unit
of the Stephen formation, a black-gray limestone, is barren in some
of the sections, while in others (Park Mountain, Mount Odaray, Fos-
sil Gully) it is extremely fossiliferous, yielding rich faunules of the
Bathyuriscus-Elrathina zone, the most characteristic and widespread
fossils being Bathyuriscus adaeus and Tonkinella stephensis. The
writer finds it difficult to establish a correlation between the faunules
observed in the Stephen formation above the Glossopleura zone and
those reported by Deiss from the same beds in the Castle Mountain
section. Unfortunately the writer had no opportunity to collect from
the Stephen at this locality and thus acquire first-hand evidence on the
faunas mentioned by Deiss. According to Deiss, in the Castle Moun-
tain section the fauna of the Glossopleura zone is succeeded by several
faunules; the lowermost one is characterized by Ehmaniella and

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Solenopleurella, the next by Solenopleurella and Thomsonaspis, the
uppermost by Olenoides, Solenopleurella, and Thomsonaspis. The
writer did not find any trilobite that could be referred to Thomson-
aspis in any of his sections ; as to the other genera mentioned by Deiss,
they suggest analogy with the above-mentioned faunule from the shaly
portion of the Stephen on Mount Odaray, and also with the Ehmani-
ella burgessensis faunule of the Burgess shale. On the other hand,
Deiss evidently did not find the faunules characterized by Bathyuris-
cus adaeus and Tonkinella stephensis in any of his sections, as he
could not have failed to comment upon such striking assemblages.
The writer’s impression is that these assemblages are younger than
any of the faunules reported by Deiss from the Stephen formation,
but the point cannot be settled until collections from the Castle Moun-
tain section have been examined. The faunules of the Ogygopsis and
Burgess shales are mentioned in discussing these units.

STEPHEN-ELDON CONTACT

The boundary between the Stephen formation and the overlying
Eldon dolomite is artificial and must be located more or less arbitrar-
ily. The situation is similar to that presented by the Mount Whyte-
Cathedral boundary, the uncertainty being still greater in the present
case. The limestone of the upper Stephen gradually becomes thicker-
bedded and changes to dolomite. When a change in thickness of the
beds happens to coincide with the limestone-dolomite boundary, ap-
parently the formations can be sharply separated. However, the
distinction is illusory, because if the strata are followed laterally,
sooner or later the limestone-dolomite contact is observed to shift to
a higher or lower horizon, or even not to follow the bedding planes
at all. This state of affairs was observed in the eastern and western
parts of Mount Stephen, on Mount Bosworth, Mount Whyte, Mount
Victoria, Mount Temple, Mount Odaray and elsewhere. Hence the
reported thickness of the upermost, calcareous unit of the Stephen
is to some extent arbitrary. The writer transferred a portion of thin-
bedded dolomite in Deiss’ Castle Mountain and Ptarmigan Peak sec-
tions from the Eldon to the Stephen because the thickness of the
beds is believed to be more significant than the distinction between
calcareous and dolomitic composition of the rock.

A better-defined Stephen-Eldon contact would result if the upper-
most unit of the Stephen were included in the Eldon. Then the
boundary, placed at the contact of an underlying shaly unit and an
overlying limestone, would become fairly sharp and independent of

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 75

the erratic dolomitization of the limestone. The Stephen formation
would be reduced to an almost uniform thickness of 249 feet on Castle
Mountain, 295 feet on Ptarmigan Peak, 319 feet on Mount Bosworth,
285 feet on Mount Whyte, and 331 feet on Mount Temple. In the
“anomalous” sections lacking the basal limestone unit, the thickness
would be 120 feet on Mount Odaray, 128 feet on Park Mountain,
and 170 feet on east Mount Stephen. The suggested change is not
formally adopted here because the Eldon formation has been in-
sufficiently studied.

On Park Mountain, the black limestone here still included in the
upper part of the Stephen is not overlain by the usual Eldon dolomite,
but by a thick succession of argillaceous limestones and siliceous
shales. Apparently no Eldon dolomite was deposited at this place,
which is only a few miles from localities (e.g., Mount Stephen, Mount
Temple, Mount Bosworth) where the Eldon is normally dolomitic.
The anomalous lithology of the Eldon on Park Mountain is analogous
to the replacement of the Cathedral dolomite by shale and argillaceous
limestone in the western part of Mount Stephen (on Park Mountain
the Cathedral is normally dolomitic). Both localities are evidently
near the margin of Walcott’s Bow trough. The boundaries that sepa-
rated the Bow and Goodsir troughs apparently oscillated with time.
When the Cathedral dolomite and limestone were deposited, Park
Mountain was within the limits of normal sedimentation of the Bow
trough, while the western part of Mount Stephen received different
sediments apparently characteristic of the Goodsir trough at that
time. Later the black limestone bearing the abundant Bathyuriscus
adaeus faunule was deposited identically at both localities. Still later,
an almost normal Eldon dolomite (except for 150 feet of interstrati-
fied siliceous shale, not present in the other sections) was deposited in
the whole area of Mount Stephen; while siliceous shales and argil-
laceous limestones that appear to have been characteristic of the Good-
sir trough at that time, were deposited in Park Mountain. Probably the
only locality where this shaly facies of the Eldon is now exposed is
Park Mountain, as the shaly facies of the Cathedral outcrops only
in Mount Stephen. The character of the sediments of the Bow and
Goodsir troughs in earliest Medial Cambrian time was mentioned in
discussing the Mount Whyte formation.

OGYGOPSIS SHALE LENTIL

The name “Ogygopsis shale’ was first used by Walcott (1908c,
p. 210) with the statement: “This term is applied to the local develop-

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ment of arenaceous and calcareous shale at the summit of the Stephen
formation on the northwest slope of Mount Stephen.”

The extremely abundant fossil content of the Ogygopsis shale was
first noted by Klotz and the fauna was described by Rominger (1887),
Matthew (1899), and Walcott (1888, 1908d).

In describing the Mount Stephen section, Walcott (1908c, 1928)
stated that the Ogygopsis shale is 150 feet thick, and that it overlies
an interval of bluish-black limestone and underlies another interval
of similar lithology which he assigned to the basal Eldon in 1908 and
to the uppermost Stephen in 1928. From this description one might
infer that the Ogygopsis shale has been observed in an orderly suc-
cession of beds and that its position at or near the top of the Stephen
formation rests on unquestionable stratigraphic evidence.

Deiss (1940) examined the Ogygopsis shale outcrop and made the
following remarks:

The Ogygopsis fauna occurs in several different kinds of matrix: brown soft
platy calcareous finely arenaceous shale in the upper beds; green tan platy harder
shale in the underlying beds; black-gray calcareous shale beneath the green and
tan interval; and white-buff soft slightly calcareous clay-shale at the base. The
shale strikes N. 10° to 20° W., and dips 35° to 42° SW. and more steeply than
the slope of Mount'Stephen at that place. Consequently, as one ascends the
slope one goes down-section through the Ogygopsis shale. At an elevation of
approximately 7,000 feet the Ogygopsis shale rests upon steel-gray, buff-weather-
ing, slightly mottled, thin-bedded Cathedral dolomites instead of being overlain
by the Eldon dolomite as had been reported. At 7,200 feet in elevation the
Cathedral dolomite is faulted against what appear to be thicker-bedded dolomites
of the basal Eldon formation.

The foregoing facts suggest that on Mount Stephen the Ogygopsis shale forms
the base instead of the upper part of the Stephen formation, that the published
type section is largely inaccurate and possibly inverted, and that beds equivalent
to the Glossopleura zone on Castle Mountain, Ptarmigan Peak and elsewhere
are absent.

The only strata with which the Ogygopsis shale is seen in contact
are the dolomites of uncertain stratigraphic position mentioned by
Deiss, as the outcrop is separated by faults from the orderly succession
of beds farther north and east. The nearest good section of the strata
underlying the Eldon dolomite is exposed and was measured by the
writer in and above Fossil Gully (section No. 9; see also fig. 3). At
this locality, 1 mile north of the Ogygopsis shale outcrop, there occur
1,000 feet of shales (representing portions of the undivided Cathedral
and Stephen formations) that are similar lithologically to portions
of the Ogygopsis shale. However, these shales are almost unfossilif-
erous, hence a possible equivalence of some part of them to the

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI If

Ogygopsis shale cannot be proved. West and south of Fossil Gully,
the entire Eldon dolomite is still exposed and undisturbed, while
exposures of lower strata are either cut off or greatly disturbed by
the Fossil Gully fault. The Eldon dolomite with the locally developed
150 feet of shale (unit 2 of the Eldon dolomite in the Fossil Gully
section) can be followed to within a distance of 4 mile from the
Ogygopsis shale outcrop. Since this shale appears unfossiliferous and
is different lithologically from any of the varieties of Ogygopsis shale,
it is exceedingly unlikely that it may represent its equivalent.

In conclusion, it appears certain that the Ogygopsis shale in its
recognizable fossiliferous form is nowhere exposed in an orderly
succession of strata and its position must be inferred from faunal
evidence. This evidence, discussed in another part of this paper,
makes it plausible that the Ogygopsis shale lies somewhere between
the lower and upper calcareous units of the Stephen formation in its
typical sections.

The writer considered the opportunity of giving the Ogygopsis
shale a proper stratigraphic name derived from a topographic feature
instead of using the faunal designation. This was not done for two
reasons: (1) following a long-established tradition makes the termi-
nology clearer than the introduction of a new name, and (2) the
Ogygopsts shale is so poorly known as to thickness and position in the
section that it affords little basis for naming a stratigraphic unit.

BURGESS SHALE LENTIL

The name Burgess shale was first used by Walcott in 1911 (Wal-
cott, I911b, p. 51, footnote) and the unit was defined as follows:

Burgess shale—This name is proposed as a geographic name for a shale to
which the term of Ogygopsis shale was given in 1908 (Smithsonian Miscel-
laneous Collections, vol. 53, p. 210). It is proposed to call it the Burgess shale
of the Stephen formation.

Type locality—Burgess Pass east of Mount Burgess and on the west slope of
Mount Field and the ridge extending to Wapta Peak. ... The Burgess for-
mation occurs to the southward across the Kicking Horse Canyon in the side of
Mount Stephen.

Thickness——On the west slope of Mount Field, 420 feet; on the northwest
slope of Mount Stephen, about 150 feet.

Stratigraphic position—Above thin bedded, dark gray and bluish-black lime-
stones of the Stephen formation, and beneath a thin bedded limestone below the
massive, arenaceous limestones of the Eldon formation that cap Mount Burgess,
Mount Field, and Mount Stephen.

This original definition included under the name “Burgess shale”
also the Ogygopsis shale on Mount Stephen. However, Walcott him-

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

self in his later publications never employed the term in this broad
sense, limiting the name Burgess shale to the outcrop on the west slope
of the ridge between Mount Field and Mount Wapta. In 1928 (Wal-
cott, 1928, p. 320) he gave a detailed section at this locality, specify-
ing that the Burgess shale has a thickness of 410 feet and that the
celebrated “‘phyllopod bed” occurs 30-40 feet above the base. In this,
as in his other publication, he considered the Burgess shale an exact
equivalent of the Ogygopsis shale on Mount Stephen.

It is well known that Walcott quarried the Burgess shale and re-
covered from it an exceptional fauna and flora including, besides fos-
sils preserved in ordinary manner, an abundance of forms lacking a
hard test and representing invertebrate groups never before or after
collected from Cambrian deposits. These unique fossils are described
in a series of publications (Walcott, Ig11a, 1911b, IgIIc, 1g12a, 1919,
1920, 1931) and their discussion is beyond the scope of this paper.

The writer examined and measured a section at and near Walcott’s
quarry on the slope of the ridge between Mount Field and Mount
Wapta (section No. 11). About 1,100 feet of approximately hori-
zontal and mostly well-exposed strata can be measured within the
fault block that includes the quarry. Immediately north of the quarry,
the strata are cut off by a normal fault that brings Cathedral dolomite
into contact with the Burgess shale. South of the quarry the beds
disappear under drift cover, and when they reappear there is no
certainty that they represent the same horizon, as the drift seems
to conceal a minor fault ; moreover, the lithology is different.

The problem of placing these 1,000 feet of strata in the general
section cannot be solved by stratigraphic evidence alone, as the suc-
cession of beds does not look like any portion of the standard section
in the area. Evidently we have here another case of sharp lateral
change of facies as we approach the margin of the Bow trough, anal-
ogous to the changes that affect the Mount Whyte, Cathedral, and
Stephen formations in the western part of Mount Stephen and the
Eldon formation on Park Mountain.

The succession of beds in the Burgess Quarry fault block, in as-
cending order, is the following. The lowermost observable strata are
poorly exposed siliceous and calcareous shales, yielding abundant
fossils. The faunule is called the Olenoides serratus faunule and is
almost identical with the faunule of the Ogygopsis shale except for
the presence of Pagetia and the absence of Ogygopsis. This interval
is overlain by 450 feet mostly consisting of dark-gray siliceous shale
with some limestone and dolomite in the upper portion. The “phyllo-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 79

pod bed” of Walcott with its faunule, here named the Pagetia bootes
faunule, occurs at the base of this unit, while an Ehmamiella burges-
sensis faunule was collected 150 feet higher. These beds are overlain
by massive Eldon (?) dolomite that forms the summit of the ridge.
We thus have at least 600 feet of almost uninterrupted shaly sedi-
ments (there may be more as the lower part of the shale disappears
under drift cover) that have no counterpart anywhere in the normal
sections. A thick succession of shales occurs in the undivided Cathe-
dral and Stephen formations on Mount Stephen, but there these strata
are almost unfossiliferous.

Hence we must resort to paleontologic evidence in order to deter-
mine the stratigraphic position of the beds in the Burgess Quarry
fault block. This evidence, presented in another part of this paper,
makes it likely that the shales here discussed represent a development
of the middle portion of the Stephen formation of the typical sections.
It is also likely that the lowermost portion of the shales exposed in the
Burgess Quarry fault block is at least an approximate equivalent of
the Ogygopsis shale, hence the latter would be somewhat older than
the “phyllopod bed” of Walcott.

FAUNAS
GENERAL STATEMENT

The fossils of the formations considered in this study belong to
a number of faunules whose relative ages are known in most cases
from unquestionable stratigraphic evidence. The problem of using
these data in order to define Middle Cambrian faunal zones of more
than local validity presents theoretical and practical difficulties, some
of which are briefly discussed.

Within a limited area such as the one here investigated, sharp verti-
cal changes in composition of the faunas may be chiefly attributed to:
(1) unconformities, i.e., time intervals not represented by sediments,
as the one that occurs at the top of the St. Piran sandstone; (2)
barren strata, as much of the Cathedral formation, especially when
represented by dolomite ; (3) changes in environment and correspond-
ing sedimentary facies, which brought assemblages not greatly different
in time, but rather in habitat, to lie over each other in the strati-
graphic column ; (4) invasion of the basin by organisms that had been
gradually evolving elsewhere.

Extensive stratigraphic and paleontologic investigation of a large
area is required in order to determine, even approximately, how much
of the vertical change of the faunal assemblages in a particular section

80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

is due to each of the above-mentioned factors; until this point has
been ascertained, subdivision into faunal zones based on that particu-
lar section or group of sections can only be tentative.

For the lower part of the stratigraphic range studied in this work,
the lowermost portion of the Middle Cambrian, so little is known of
the faunal succession elsewhere that the writer felt justified in estab-
lishing tentative zones based on the narrow investigated area. The
two lowermost Middle Cambrian zones here proposed, the Wenk-
chemnia-Stephenaspis and Plagiura-Kochaspis zones, seem distinct
enough to deserve separate recognition. However, it is possible that
the differences observed are partly due to differences in facies between
the corresponding stratigraphic intervals, and it would not be sur-
prising to find in another area the genera here considered character-
istic of the respective zones occurring together.

The succeeding Albertella and Glossopleura zones seem well char-
acterized and can be recognized in most of the Cordilleran sections
where the corresponding time intervals are represented. The Middle
Cambrian zones above the Glossopleura zone so far proposed seem
to the writer of doubtful standing. In the Cambrian correlation chart
(Howell et al., 1944), the lists of genera considered characteristic of
these zones include mixtures of forms from the Cordilleran and
Atlantic provinces lumped together on the basis of hypothetical cor-
relations. Even if the genera from the Atlantic province are disre-
garded, the remaining genera do not appear to be sufficiently distinc-
tive and short ranging to allow a clear definition of the zones.

Middle Cambrian zones have always been defined in terms of trilo-
bites, and the paleontologic material available does not suggest de-
parture from this practice. The writer collected non-trilobitic fossils
whenever possible. Next to the trilobites, brachiopods and gastropods
(or at least forms usually placed in the latter group) are the most
important elements of the faunas. Hyolithes, Hyolithellus, and sponge
spicules are fairly common in certain beds. These are all the animal
groups represented in the Mount Whyte, Cathedral, and Stephen
formations apart from the extraordinary fossils of the Burgess shale
and other exceptional occurrences of little use for stratigraphic pur-
poses. Besides the trilobites, the other groups that might have value
as index fossils are the brachiopods and the gastropods. The former
are fairly numerous, but seldom well preserved. Nevertheless it is
possible that a paleontologist possessing profound knowledge of the
brachiopods and skill in the difficult techniques of preparation could
obtain stratigraphic information from these fossils. The excellent

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 81

work of Bell (1941) on the Cambrian brachiopods of Montana is an
example of what can be done in this field. The writer must frankly
acknowledge his inadequate competence and list it as one of the
reasons for not making use of the brachiopods. The gastropods seem
less promising, and a general study of the Cambrian forms would be
required before they can acquire stratigraphic value. No new binomial
combinations are proposed for nontrilobitic forms even when the
present generic reference is known to be erroneous.

Only species collected by the writer from strata in measured sec-
tions are used in determining the faunal succession. Exceptions are
made for the Ogygopsis and Burgess shales where no doubt exists as
to the beds from which the fossils were collected.

Deiss published faunal lists, frequently giving only generic identifi-
cations or referring to undescribed genera and species. Hence his
lists are not quoted as they would add but little information.

FAUNA OF THE BONNIA-OLENELLUS ZONE

Although this paper is primarily concerned with Middle Cambrian
stratigraphy and faunas, a brief description of the youngest Lower
Cambrian fauna observed in the area is required in order to discuss
the problem of the Lower-Middle Cambrian boundary.

Faunal zones for the Lower Cambrian of the Appalachian and Cor-
dilleran provinces have been tentatively defined, but an examination
of the basis for the suggested subdivisions shows their highly doubt-
ful standing. The writer believes that the choice of the southern
Appalachians for a standard Early Cambrian time scale (Resser,
1933) should be abandoned. The structure in that area is excessively
complicated, thick and well-exposed sections of Lower Cambrian
strata are lacking, and most of the fossils are known from scattered
outcrops of uncertain stratigraphic position. Resser (1938b) named,
in ascending order, the Obolella, Bonnia, Kootenia, Olenellus, and
Kochiella zones, the last lacking olenellid trilobites. In the Cambrian
correlation chart (Howell et al., 1944) the Lower Cambrian is divided
into the Obolella, Bonnia, Olenellus, and Syspacephalus zones.

The writer does not recognize a post-olenellian Lower Cambrian
zone (Kochiella or Syspacephalus zone for reasons that are fully
discussed in the next section of this paper. As to the other proposed
zones, the following remarks are indicated. (1) The Obolella zone,
supposed to lack trilobites, was based on poorly fossiliferous forma-
tions of the southern Appalachians. There is no certainty that the
absence of olenellids from these beds is not due to unfavorable facies.

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Olenellids range through great thicknesses of beds in the Cordilleran
province. (2) The Bonnia and Kootenia zones were chiefly estab-
lished on the fauna of the reef limestone at Austinville, Va., attributed
to the Shady formation. However, Stose and Jonas (1938) ques-
tioned the stratigraphic position of the limestone, which occurs in an
isolated fault block. It is true that similar faunules occur in orderly
sections in southern Labrador and western Newfoundland, but at

those localities the faunal succession has been insufficiently investi-

gated. (3) The Olenellus zone may represent a shaly facies of the
Lower Cambrian deposits rather than a definite time interval. In
northwestern Vermont, for example, olenellids seem to prevail when
the Lower Cambrian is represented by siliceous shales, while Bonnia
and small, generalized ptychoparid trilobites are dominant in limestone
or dolomite formations, regardless of age.

It is clear that a satisfactory standard time scale for the Early
Cambrian will have to be based on an area where thick, orderly, and
reasonably fossiliferous successions of Lower Cambrian strata are
present. Possibly such sections will be found in western Newfound-
land. However, it is more likely that a standard section will be
ultimately established in the Cordilleran province, possibly in Sonora
(Cooper and Arellano, 1946), or in Nevada, California, or the Ca-
nadian Rockies. The area here described is inappropriate for the
purpose because most of the Lower Cambrian is unfossiliferous.

The youngest Early Cambrian fauna occurring in the area, the
fauna of the Peyto limestone member of the St. Piran sandstone, is
listed here in order to illustrate the great faunal change taking place
at the Lower-Middle Cambrian boundary. As far as the fossils have
been studied, there seems to be little lateral or vertical change in com-
position of the faunules within the Peyto limestone even where this
member attains considerable thickness. Hence these faunules will be
listed as one unit. Only the trilobites collected and identified by the
writer are listed. This assemblage will be designated as the Bonnia
fieldensis faunule. An asterisk preceding the name indicates that the
type locality and horizon of the species are within the Peyto limestone
in the described area.

BONNIA FIELDENSIS FAUNULE

* “Agraulos” unca Walcott.

*Bonnia fieldensis (Walcott) (= columbensis Resser).
*Crassifimbra lux (Walcott).

*Olenellus canadensis Walcott.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 83

*Onchocephalus cleon (Walcott).

* thia (Walcott).
Paedeumias sp.
Pagetia sp.

*Piazella pia (Walcott).
*Syspacephalus charops (Walcott).
Zacanthopsis cf. levis (Walcott).

To give a correct picture of the composition of the fauna, it must
be stated that the Olenellidae are by far the most abundant fossils,
some beds being a coquina of their tests. Unfortunately, owing to
the large size of these trilobites and the thinness of the test, good
specimens are excessively rare. The next in abundance is Bonnia,
whose thick, well-preserved head and tail shields can be collected from
almost any outcrop of the Peyto limestone. The small, generalized
Ptychoparioidae are rare except at two localities, on Mount Stephen
and Mount Schaffer. Pagetia was collected only on Mount Stephen
and is very rare. The non-trilobitic forms are chiefly represented by
the brachiopod Nisusia and the gastropod Scenella.

The fauna of the Peyto limestone cannot be definitely assigned to
any one of the suggested Lower Cambrian zones. The writer does
not see important differences between this fauna and the faunas of the
Forteau formation of Labrador and Newfoundland, the Mallet dolo-
mite of northwestern Vermont, and the Shady (?) limestone reefs at
Austinville, Va. Resser (1938b) included all these faunas in the
Bonmia zone, supposed to be older than the Olenellus zone. If this
interpretation were accepted, the Peyto limestone would be older than
such formations as the Rome of the southern Appalachians and the
_ Parker shale of northwestern Vermont. On the other hand, Lochman
(1947) suggested the name “Antagmus-Onchocephalus zone’ for
“the uppermost Lower Cambrian zone” in the Cordilleran province.
Apparently she had in mind faunas precisely of the type of that occur-
ring in the Peyto limestone. Deiss assigned the fauna of the Peyto
limestone (which he considered as the basal portion of the Mount
Whyte formation) to a “Bonnia-Olenellus zone.’ Since Bonnia and
Olenellidae are by far the most common fossils in the Peyto limestone,
Deiss’ term will be tentatively used. This does not imply that the
Bonma-Olenellus zone of Deiss, which appears synonymous with
the Antagmus-Onchocephalus zone of Lochman, has been recognized
to possess general validity even within the Cordilleran province. Its
relationship with the tentative zones proposed by Resser remains
problematical.

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

LOWER-MIDDLE CAMBRIAN BOUNDARY

The uncertainty existing about the Lower-Middle Cambrian bound-
ary in the Cordilleran province is chiefly due to three factors: in-
sufficient stratigraphic work, poor knowledge of the faunas, and
failure of the stratigraphers, until recent times, to realize clearly the
distinction betwen time-stratigraphic and lithologic units (Ashley
et al., 1933, 1939; Schenk and Muller, 1941; Moore, 1947). Many
authors did not seem to understand the contradiction between the
definition of periods and epochs in terms of time, and their attempts
to locate the boundaries between the corresponding systems and series
by diastrophic or lithologic criteria.

Much of the discussion of the Lower-Middle Cambrian boundary
in the Cordilleran province was based on the area here described, as
it includes one of the most complete, well-exposed, and fossiliferous
records of this portion of the stratigraphic column.

Granted that series are time-stratigraphic units and that faunas
supply the only criterion for time equivalence of deposits in distant
areas, it follows that the Lower-Middle Cambrian boundary, or the
corresponding time boundary between the Waucobian and Albertan
epochs, must be defined in terms of faunas.

For many years the Lower Cambrian in North America has been
synonymous with the strata carrying the Olenellus fauna (Walcott,
1890). This still appears a sound definition, hence the writer con-
siders the Early Cambrian epoch as terminating with the disappear-
ance of the Olenellidae. Obviously there is no certainty that the
Olenellidae, or any other group of organisms, appeared or became
extinct contemporaneously (or at least within a time interval short
enough to be considered negligible on the geologic time scale) even
within a single epicontinental sea, such as the one that occupied the
Cordilleran geosyncline. The best provisional solution of this difficulty
seems to be to define a conventional time equivalence based on a
widespread and well-characterized group of fossils (in the present
instance the Olenellidae) and use it until the study of the whole
faunas in a wide area suggests different correlations. The writer is
fully aware of the weakness of correlations based on a single group
of organisms. For example, if the Medial Cambrian epoch were de-
fined by the appearance of the Agnostida, the correlation between
deposits of the Appalachian and Cordilleran provinces would be very
different from the generally accepted one. In the Appalachians, rare
agnostids already occur in association with the olenellids, whereas in
the Cordilleran province, and especially in the area investigated, there

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 85

is a wide gap, occupied by several faunas, between the extinction of
the olenellids and the appearance of the agnostids. In this instance,
evidence from most of the other fossils favors the correlation based
on the olenellids, but the true time equivalence may lie somewhere
between the extremes resulting from the consideration of each group
alone.

In recent times attempts have been made to raise the Lower-Middle
Cambrian boundary and thus to admit a post-olenellian Lower Cam-
brian zone. Examination of the stratigraphic and paleontologic
grounds for the suggested modification shows that it was chiefly based
on insufficient knowledge of the faunas and misinterpreted strati-
graphic evidence. This discussion refers only to the Appalachian and
Cordilleran provinces of North America. Precise correlation with the
deposits of the Atlantic province and other regions is still uncertain ;
furthermore, as pointed out by Resser (1933), the Cambrian sequence
in North America is far more complete than in other areas and must
be used for establishing a generalized time scale.

Resser (1933) tentatively established a post-olenellian Lower Cam-
brian zone and named it the Kochiella zone, stating (Resser, 1933,

p- 744) that—

In the Rocky Mountains the Mount Whyte formation contains the usual olenel-
lid trilobites in its lower portions, but toward the top the Kochiella faunas appear.
In Manchuria and Greenland similar changes occur, and possibly this portion may
be ultimately placed in the Middle Cambrian, even though there is no change in
lithology at any place.

The Kochiella zone was again used by Resser (1938b) to designate
the upper portion of the Rome formation in the southern Appala-
chians. Resser and Howell (1938) stated that “the uppermost Lower
Cambrian is characterized by the Kochiella fauna which lacks olenel-
lid trilobites.” Howell et al. (1944) recognized a Syspacephalus zone
as the highest post-olenellian Lower Cambrian zone and listed Sys-
pacephalus, Inglefieldia, and Kochiella as its characteristic genera.

Let us now examine the facts on which this post-olenellian Lower
Cambrian zone was established. The genotype of Kochiella and
several other species were described by Poulsen (1927) and stated to
occur in association with Olenellidae in northwest Greenland. Profes-
sor Poulsen confirmed the presence of this association in a personal
communication to the writer. Forms that may doubtfully be assigned
to Kochiella do occur in post-olenellian strata in the Rocky Moun-
tains ; still it seems a questionable procedure to name a post-olenellian
zone after a genus whose genotype and most of the other described

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

species occur in association with Olenellidae. Similar considerations
apply to /nglefieldia, whose described species, according to Poulsen,
occur in association with Olenellidae in the Cape Kent formation of
Greenland. No forms definitely assignable to Jnglefieldia are known
to the writer from the Cordilleran province. Finally, before the pres-
ent work Syspacephalus was known only from the genotype, a species
occurring at a single locality in the Peyto limestone on Mount Stephen,
where it is associated with Olenellus and Bonnia. We conclude that
the totality of the genotypes and most of the described species of the
genera given as characteristic of the uppermost post-olenellian Lower
Cambrian zone occur in association with Olenellidae.

The present work shows from unmistakable stratigraphic evidence
that Plagiura cercops and associated forms, hitherto considered Lower
Cambrian, occur above the shale bearing Amecephalus agnesensis,
Poliella prima, and other species assigned to the Middle Cambrian.
The latter assignment appears to the writer entirely correct, hence the
Plagiura faunule cannot be considered Lower Cambrian. Deiss
(1939) recognized a Plagiura zone and correctly considered these
beds as post-olenellian, but did not give any reasons for his statement
that these are “undoubted Lower Cambrian rocks.”

The entire Mount Whyte formation was formerly considered
Lower Cambrian because of the belief that Olenellidae range through
the entire unit. Deiss observed that these trilobites are confined to the
basal beds but failed to draw the logical conclusion that there was no
longer any basis for assigning the entire unit to the Lower Cambrian.
In this work the basal, Olenellus-bearing beds are assigned to the
underlying St. Piran sandstone, and the Mount Whyte formation thus
restricted is considered entirely Middle Cambrian.

The conclusions here adopted are the following:

1. The Early Cambrian, according to tradition, is defined as ter-
minating with the disappearance of the Olenellidae. This definition,
at least provisionally, applies only to the Appalachian and Cordilleran
provinces.

2. In the area investigated, there are no Early-Medial Cambrian
transition faunas, the oldest post-olenellian fauna already having fully
developed Medial Cambrian character.

3. Formations, being defined as lithologic units, may belong to two
epochs. However, in the area studied, the Lower-Middle boundary as
above defined coincides with the probably unconformable contact be-
tween the Peyto limestone member of the St. Piran sandstone and the
Mount Whyte formation. Burling (1914, 1916) and Grabau (1936)

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 87

had already suggested placing the Lower-Middle Cambrian boundary
at the top of the Olenellus zone, i.e., within the Mount Whyte forma-
tion as understood at that time.

4. In other areas of the Cordilleran geosyncline, according to
published evidence, there seems to be a still greater Lower-Middle
Cambrian unconformity, since the Kochaspis liliana fauna has been
reported almost immediately to overlie the Olenellus fauna, thus in-
dicating absence of the intermediate fauna assigned to the Wenkchem-
mia-Stephenaspis zone in this paper.

FAUNA OF THE WENKCHEMNIA-STEPHENASPIS ZONE

The post-olenellian faunas older than the Albertella fauna were
assigned to a Lower Cambrian Kochiella zone by Resser (1938b) ;
to a Lower Cambrian Plagiura zone and a Middle Cambrian Kochas-
pis zone by Deiss (1939) ; to a Lower Cambrian Syspacephalus zone
and a Middle Cambrian Kochaspis liliana zone by Howell et al.
(1944). Lochman (1947) did not recognize the existence of a Middle
Cambrian zone older than the Albertella zone.

The inconsistencies on which the Kochiella or Syspacephalus zone
was based were listed in the discussion of the Lower-Middle Cambrian
boundary. The Plagiura and Kochaspis liliana zones seem to be es-
sentially equivalent, although Deiss considered them distinct enough
to be placed in different epochs. Since more and better-preserved fos-
sils from these zones are described in this paper than in all the pre-
vious literature, a new approach to the problem is indicated, and two
tentative zones are established on the basis of the faunules collected
from the Mount Whyte formation and studied in the present work.

The lower zone includes assemblages that were hitherto unknown
or misunderstood, and is designated as the W enkchemnia-Stephenaspis
zone after two new trilobite genera. The upper zone is designated as
the Plagiura-Kochaspis zone because it includes the Plagiura zone of
Deiss and the essentially equivalent Kochaspis liliana zone of various
authors.

The fauna of the Wenkchemnia-Stephenaspis zone occurs in the
lower or middle portion of the Mount Whyte formation. However,
in several sections the lower part of the Mount Whyte is unfossilifer-
ous; in others (e.g., the Eiffel Peak section) strata of this age appar-
ently were never deposited (see discussion of Mount Whyte forma-
tion). Hence our knowledge of the fauna is entirely based on two
limited areas: (1) the Mount Whyte-Mount Niblock area, where a
faunule of this age occurs in the Lake Agnes shale lentil ; and (2) the

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

western part of Mount Stephen, where the lower and middle portions
of the Mount Whyte formation are unusually thick and fossiliferous,
and yield several faunules of which the youngest is particularly abun-
dant in the Yoho shale lentil.

In the western part of Mount Stephen, in the Fossil Gully and
North Gully sections, four faunules assigned to the Wenkchemmia-
Stephenaspis zone were observed in orderly succession. The oldest
of these faunules occurs 140 feet above the top of the Lower Cam-
brian Bonnia-Olenellus zone in the North Gully section, while the
youngest occurs in strata immediately underlying the Plagiura-Koch-
aspis zone. The oldest faunule of the Wenkchemnia-Stephenaspis
zone was collected in limestone at locality W16h and includes the
species :

Acrotreta sp.

Syspacephalus laticeps Rasetti.
Wenkchemma sulcata Rasetti.

The next faunule occurs both in shale and limestone at locality
W16i, a few feet above locality W16h, and may be identical with the
preceding. The species include:

Syspacephalus sp.
Wenkchemnia sulcata Rasetti.

The third faunule ranges through almost 100 feet of impure lime-
stone and shale in the overlying interval. Collections were made at
several horizons (see lists of species from localities W16j, W16j’,
W16j”) and showed little change in composition of the faunules,
hence these assemblages will be treated as one unit named the Syspa-
cephalus laticeps faunule. However, it must be noticed that Ogygopsis
klotzi was found only in the upper part of this interval. The list of
species is the following (an asterisk indicates that the types of the
species belong to this assemblage) :

SYSPACEPHALUS LATICEPS FAUNULE

Acrotreta sp.
Ogygopsis klotzi (Rominger).
Oryctocephalus sp. No. 1.
*Piazella tuberculata Rasetti.
Poliella cf. P. denticulata Rasetti.
Stephenaspis cf. S. bispinosa Rasetti.
oa ‘yspacephalus laticeps Rasetti.
crassus Rasetti.
cf. S. gregarius Rasetti.
*WVenkchemnia sulcata Rasetti.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 89

The fourth faunule is represented by great numbers of individuals
in the Yoho shale lentil (localities Wok, Wok’, W16k), and is desig-
nated as the Stephenaspis bispinosa faunule from its most striking
trilobite. The species include:

STEPHENASPIS BISPINOSA FAUNULE

Hyolithes sp.
Scenella sp.
Acrothele sp.
Acrotreta sp.
Dictyonina sp.
Paterina sp.
Amecephalus agnesensis (Walcott).
*Chancia stenometopa Rasetti.
Ogygopsis klotzsi (Rominger).
Oryctocephalus sp. No. 2.
*Poliella denticulata Rasetti.
prima (Walcott).
*Stephenaspis bispinosa Rasetti.
*Syspacephalus gregarius Rasetti.
= laevigatus Rasetti.
perola (Walcott).
*W enkchemma spinicollis Rasetti.

In the Mount Whyte area (localities W2b, W3b; U.S.N.M. 35m),
the Lake Agnes shale lentil yields a faunule that is designated by
the trilobite Poliella prima and includes the species:

POLIELLA PRIMA FAUNULE

Hyolithes sp.
*Acrothele clitus Walcott.

Acrotreta sp.
*Amecephalus agnesensis (Walcott).
*Poliella prima (Walcott).
*Syspacephalus perola (Walcott).
*W enkchemnia walcotti Rasetti.

It is clear from the percentage of common species that the Stephen-
aspis bispinosa and Poliella prima faunules are approximately of the
same age. The latter is likely to be the younger because of the abun-
dance of Amecephalus, which appears to be absent from the earlier
faunules of the Wenkchemnia-Stephenaspis zone and becomes com-
mon in the overlying Plagiura-Kochaspis zone.

The identified trilobite genera in the Wenkchemnia-Stephenaspis
zone are: Amecephalus, Ogygopsis, Oryctocephalus, Piazella, Poliella,
Stephenaspis, Syspacephalus, and Wenkchemnia. The fauna, compared

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

with that of the Bonnia-Olenellus zone, is sharply differentiated by
the absence of Bonnia and the Olenellidae, and the appearance of
Amecephalus, Ogygopsis, Oryctocephalus, and several genera of the
Dolichometopidae, Poliella, Stephenaspis, and Wenkchemnia. The
only genera common with the uppermost Lower Cambrian zone are
the generalized ptychoparids Piazella and Syspacephalus, but little
significance should be attributed to the persistence of such forms that
are known to range, without differences justifying generic separation,
through much of the Lower and Middle Cambrian. On the other
hand, the fauna of the Wenkchemnia-Stephenaspis zone is rather
closely related to those of the overlying Plagiura-Kochaspis zone and
still higher Middle Cambrian zones, and presents fully developed Mid-
dle Cambrian characteristics. Hence the writer considers it amply
justifiable to include this zone in the Middle Cambrian, whereas Deiss
would consider even a portion of the still higher Plagiura-K ochaspis
zone as Lower Cambrian. Furthermore, the great faunal change be-
tween the Bonnia-Olenellus and the Wenkchemnia-Stephenaspis
zones suggests an unrepresented time interval, i.e., an unconformity
between the Peyto limestone and the Mount Whyte formation as it
was indicated in discussing these units.

As far as the writer knows, if the fauna of the Wenkchemmia-
Stephenaspis zone exists in other areas of the Cordilleran province
there is no published evidence to indicate it.

FAUNA OF THE PLAGIURA-KOCHASPIS ZONE

The fauna here designated as characteristic of the Plagiwra-Koch-
aspis zone is a clearly defined assemblage of closely related faunules
occurring in the upper part of the Mount Whyte formation. Some of
its species were assigned by Deiss and Resser to the Lower Cambrian
(e.g., Plagiura cercops), others to the Middle Cambrian (e.g., species
of Kochaspis), but additional collecting has shown that these fossils
occur in the same beds. Furthermore, unquestionable stratigraphic
evidence from the Mount Whyte and Fossil Gully sections shows that
the Plagiura-Kochaspis fauna is younger than any of the faunules
assigned to the Wenkchemnia-Stephenaspis zone. Since the fauna of
the W enkchemnia-Stephenaspis zone was assigned to the Middle Cam-
brian, Plagiura cercops and the associated forms cannot be considered
Lower Cambrian.

The upper part of the Mount Whyte formation is generally more
fossiliferous than the underlying strata, hence the Plagiura-K ochaspis
fauna is represented in most of the investigated sections. Several

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI gI

faunules can be distinguished within the zone, although changes are
gradual and the general character of the fauna is preserved through-
out. The oldest faunule occurs in several sections and is designated
as the Plagiura cercops faunule although this trilobite ranges into
somewhat higher beds. This faunule occurs in limestone and is rep-
resented at localities W4c-d, Ross Lake; W5c, Mount Bosworth;
W7c-e, Kicking Horse Mine; and several others of Walcott’s and
the author’s localities. The lists from these localities are all essentially
the same and include the species:

PLAGIURA CERCOPS FAUNULE

Amecephalus cleora (Walcott).
Caborcella skapta (Walcott).
Fieldaspis furcata Rasetti.
Onchocephalus fieldensis Rasetti.
*Plagiura cercops (Walcott).
Schistometopus convexus Rasetti.

Amecephalus cleora and Plagiura cercops are common at all locali-
ties, while the other species are moderately common or rare.

In somewhat higher beds, a few new forms appear and the relative
abundances of the others undergo considerable changes. At localities
W7! (Kicking Horse Mine) and Wof (Fossil Gully), a highly fos-
siliferous limestone yields a faunule named the Fieldaspis furcata
faunule from its most common and striking trilobite. Plagiura cercops
and Amecephalus cleora are rare members of this assemblage. The
species include:

FIELDASPIS FURCATA FAUNULE

Amecephalus cleora (Walcott).
*Fieldaspis furcata Rasetti.
Kochaspis eiffelensis Rasetti.
Kochiella? maxeyi Rasetti.
*Oryctocephalites resseri Rasetti.
*Onchocephalus fieldensis Rasetti.
* maior Rasetti.
Plagiura cercops (Walcott).
Schistometopus convexus Rasetti.

On Eiffel Peak (locality W2od) a similar faunule occurs abun-
dantly both in limestone and siliceous shale. Its composition is so
similar to that of the Fieldaspis furcata faunule that it is doubtful
whether it should be considered a distinct unit. The chief differences
are the great abundance of Amecephalus cleora and Plagiura cercops

g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

and the replacement of Fieldaspis furcata (which is extremely rare)
by another species of the same genus, Fieldaspis bilobata, from which
the faunule is named. The species include:

FIELDASPIS BILOBATA FAUNULE

Amecephalus cleora (Walcott).
*Fieldaspis bilobata Rasetti.
furcata Rasetti.
*Kochaspis eiffelensis Rasetti.
*Kochiella? maxeyi Rasetti.
Plagiura cercops (Walcott).
*Schistometopus collaris Rasetti.
* convexus Rasetti.

The youngest faunule of the Plagiura-K ochaspis zone was collected
at the Kicking Horse Mine, in a limestone 50 feet stratigraphically
above the beds carrying the Fieldaspis furcata faunule (locality
W7g). This faunule is named from the trilobite Fieldaspis superba
and includes the species:

FIELDASPIS SUPERBA FAUNULE

*Caborcella rara Rasetti.
*Fieldaspis superba Rasetti.
*Onchocephalus depressus Rasetti.
* sublaevis Rasetti.

The Plagiura-Kochaspis fauna as a whole includes in the area
investigated the trilobite genera Amecephalus, Caborcella, Field-
aspis, Kochaspis, Onchocephalus, Oryctocephalites, Plagiura, Schisto-
metopus, and a species doubtfully assigned to Kochiella. Fieldaspis
and Plagiura, as far as known, are confined to this zone. The same
holds for Kochaspis and Schistometopus in the area investigated, but
the genotype of Schistometopus occurs in a younger fauna, and Koch-
aspis is generally considered to range through most of the Middle
Cambrian. However, the late Medial Cambrian species assigned to
this genus (e.g., K. unzia and K. upis) are doubtfully congeneric
with the genotype, and it is possible that when Kochaspis is properly
restricted it will be found to be confined to the early Medial Cambrian
beds. Amecephalus first occurs in the Wenkchemnia-Stephenaspis
zone and ranges into higher beds, hence is of little stratigraphic value.
Onchocephalus is based on a Lower Cambrian genotype, but little
significance should be attributed to the generic assignment of these
generalized ptychoparids.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 93

The Plagiura-Kochaspis fauna is clearly demarcated both from the
older W enkchemnia-Stephenaspis fauna and from the younger Alber-
tella fauna. The composition of the fauna in the Canadian Rockies is
such that the name Plagiura fauna (or Plagiura-Fieldaspis fauna)
would be more appropriate, as Kochaspis is a rare fossil. However,
Plagiura and Fieldaspis seem to be unknown elsewhere in the Cordil-
leran province, whereas Kochaspis is abundant in strata attributed to
this zone (e.g., the genotype, Kochaspis liliana, in the Pioche shale
of the Highland Range, Nevada). Hence the mixed designation for
the zone. Very little has been published on the fossils of the Plagiura-
Kochaspis zone elsewhere.

FAUNA OF THE ALBERTELLA ZONE

The Plagiura-Kochaspis fauna is succeeded in the Canadian
Rockies by the Albertella fauna which occurs in the Cathedral for-
mation. This is a well-known assemblage of widespread occurrence
in the Cordilleran province. The Albertella-bearing beds are separated
from the Mount Whyte formation by 400 or more feet of unfossilifer-
ous limestone or dolomite, hence a considerable time interval is un-
represented by fossils and it is not surprising to find that the Albertella
fauna has few elements in common with the Plagiura-Kochaspis
fauna.

Two faunules definitely belonging to the Albertella zone are known
in the area. The older faunule occurs abundantly in the Ross Lake
shale member of the Cathedral formation and is present in most of
the sections where this horizon was investigated. This faunule is
designated as the Albertella bosworthi faunule from one of the most
common species. The fossils are most frequently collected in shale,
but most of the species are known also in limestone. The following
list includes the species reported from Walcott’s localities U.S.N.M.
63j, 63m, and 35c, and from the writer’s localities C3m, Cqam, C15m,
C2om, and C21m. The list of nontrilobitic species is taken from Wal-
cott (1917b) with the modifications given by Resser (1938c).

ALBERTELLA BOSWORTHI FAUNULE

*Coleoloides hectori (Walcott).
Eocystites? sp.

*Hyolithes cecrops Walcott.

*Tholiasterella? hindei Walcott.
*Urotheca parasitum Resser.
*Acrothele walcotti Resser.

94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

*Obolus parvus Walcott.
*Paterina wapta (Walcott).
*Wimanella rossensis Resser.

* qwalcotti Resser.
*Albertella bosworthi Walcott.
- declivis Rasetti.

x microps Rasetti.

* nitida Resser.

Kochina americana (Walcott).
*Mexicella stator (Walcott).
*Ptarmigania rossensis (Walcott).
*Vanuxemella nortia Walcott.

The younger faunule is known only from the Bow Lake section
(locality C15n) where it occurs in black-gray limestone immediately
above the Ross Lake shale. This assemblage is named the Albertella
limbata faunule and includes the species:

ALBERTELLA LIMBATA FAUNULE

*Albertella limbata Rasetti.

* stenorhachis Rasetti.
*Kochina macrops Rasetti.
*Ptarmiganoides bowensis Rasetti.

The Albertella fauna in the Canadian Rockies thus includes the
trilobite genera Albertella, Kochina, Me.xicella, Ptarmigania, Ptarm-
ganoides, and Vanuxemella, All these genera, possibly excepting
Kochina, are readily recognizable and apparently confined to the zone.
Hence the fauna of the Albertella zone is excellently characterized.
Obviously this is partly due, in the Canadian Rockies, to the absence
of immediately older or younger fossils. The limits that should be
assigned to the Albertella zone are not so clearly demarcated in other
sections of the Cordilleran province. In the Langston (?) formation of
Idaho, an assemblage which must be at least partly included in the Al-
bertella fauna, was described by Resser (1939b) as the “Ptarmigania
fauna.” In this assemblage Albertella is exceedingly rare (Resser did
not describe any species of Albertella, but the writer collected speci-
mens undoubtedly belonging to this genus). The most common trilo-
bites belong to the genera Caborcella, Kootenia, Oryctocephalus, Pa-
getia, Ptarmigania, Ptarmiganoides, and Zacanthoides (only genera
collected and identified by the writer are listed, as Resser described a
mixture of forms from different horizons and the writer does not ac-
cept several of his generic assignments). Of these genera, Albertella,

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 95

Ptarmigania, and Ptarmiganoides are definitely indicative of the Alber-
tella zone, while the others are too long-ranging to give any indication
for or against this assignment. Resser united this faunule with a
younger one characterized by Ogygopsis (Resser’s “Taxioura’) and
Pachyaspis, but the writer found that these two assemblages occur
in different beds. Whether the younger assemblage should still be
included in the Albertella fauna cannot be decided at present.

Several faunules of uncertain age which may belong to the Al-
bertella zone are discussed here, but their fossils were not included in
the list of genera from the Albertella zone given above.

The shales and impure limestones representing the lower portion of
the Cathedral formation in the Fossil Gully section on Mount Stephen
yield several faunules that are tentatively assigned to the Albertella
zone but actually include either long-ranging genera or forms unknown
elsewhere, hence cannot be definitely assigned to a faunal zone from
paleontologic evidence. Unquestionable stratigraphic evidence shows
that these faunules are younger than the fauna of the Plagiura-K och-
aspis zone; the lowermost occurring about 50 feet above the top of
the Plagiura-Kochaspis zone. From this stratigraphic position, one
would be tempted to infer that this faunule is older than the Al-
bertella bosworthi faunule, which occurs in the Ross Lake shale of
the Cathedral formation 400 or more feet above the top of the
Plagiura-Kochaspis zone. However, the Ross Lake shale and the as-
sociated Albertella bosworthi faunule were not observed in the west-
ern part of Mount Stephen, and the section at that locality is very dif-
ferent from the normal sections in the Bow Range. Hence there is no
evidence that the strata 50 feet above the top of the Plagiura-K ochaspis
zone at Fossil Gully are older than the strata 400 feet above the top
of the same zone elsewhere; on the contrary, from the paleontologic
evidence the writer inclines to believe that they are younger, i.e., that
in the Fossil Gully section there is a considerable unconformity at the
top of the Mount Whyte formation and that the oldest beds of the
Cathedral formation were deposited there after the Ross Lake shale
had been sedimented in the normal sections.

The next higher well-identifiable faunule in the Fossil Gully section
is the Bathyuriscus adaeus faunule of the upper Stephen formation,
occuring almost 2,000 feet above the strata containing these faunules
of uncertain age.

The lowermost of the faunules tentatively assigned to the Albertella
zone occurs in the Fossil Gully section at locality Coh and is abun-
dantly but poorly preserved in siltstone and impure, dark-gray lime-

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

stone through 50 or more feet of strata. This faunule is named the
Yohoaspis pachycephala faunule and includes the species :

YOHOASPIS PACHYCEPHALA FAUNULE

Acrothele sp.

*Chancia latigena Rasetti.

Kootenia sp.

Ogygopsis klotzi (Rominger).
= spinulosa Rasetti.

Pagetia sp.

Poliella sp. No. 1.
*Syspacephalus tardus Rasetti.
*Yohoaspis pachycephala Rasetti.

Zacanthoides sp.

The next faunule was collected about 100 feet higher in the section,
in lenticular beds of purer limestone interstratified with slightly argil-
laceous or silty limestone (localities Coj, Coj’). The fossils from
these two horizons are listed together and the faunule is named the
Chancia bigranulosa faunule from a trilobite that is common at both
localities.

CHANCIA BIGRANULOSA FAUNULE

Diraphora sp.
Wimanella sp.
Athabaskia sp.
Bathyuriscus sp.

*Chancia bigranulosa Rasetti.
Ogygopsis klotzi (Rominger).
Pagetia sp.

Poliella sp. No. 2.
Yohoaspis pachycephala Rasetti.

*Zacanthoides sexdentatus Rasetti.

FAUNA OF THE GLOSSOPLEURA ZONE

In the Cambrian correlation chart (Howell et al., 1944) the Al-
bertella zone is succeeded by a Zacanthoides-Anoria zone supposed
to be characterized by the trilobite genera Zacanthoides, Anoria, Daw-
sonia, Kootenia, Strotocephalus, and Clavaspidella, Lochman (1948)
does not recognize the identity of a faunal zone characterized by the
restricted occurrence of Zacanthoides and Anoria, and the writer fully
concurs with her criticism. Zacanthoides and Kootenia range from the
uppermost Lower Cambrian through most of the Middle Cambrian
and are obviously extremely poor diagnostic genera. Anoria probably

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 97

has a restricted vertical range, but is a typical fossil of the Glosso-
pleura zone, as shown by the occurrence of the genotype, Anoria
tontoensis, in the Glossopleura zone of the Grand Canyon area. Daw-
somia is an Atlantic province genus which for obscure reasons was
associated with the other genera from the Cordilleran province in
the faunal lists given in the correlation chart; its age relative to the
Cordilleran faunas is entirely problematical. Strotocephalus (which
the writer considers a synonym of Amecephalus) is common, as
shown in the present work, through the Wenkchemnia-Stephenaspis
and Plagiura-Kochaspis zones and extends into younger beds. Clava-
spidella, or rather Athabaskia, ranges at least from the Albertella zone
to the Glossopleura zone.

In the particular area studied in this work, the problem of recog-
nizing a Zacanthoides-Anoria zone does not arise, since in the typical
sections the Albertella zone is followed by a barren interval (the
upper portion of the Cathedral formation) and the next younger
faunule is typical of the Glossopleura zone. The only possible ex-
ception may be represented by the peculiar faunules occurring in the
equivalent of the lower part of the Cathedral formation in the
Fossil Gully section. These faunules were tentatively assigned to the
Albertella zone.

The writer prefers to use the term Glossopleura zone instead of
Glossopleura-Kootenia zone as in the correlation chart, because the
genotype of Kootenia and many other species of the genus occur in
much younger strata; Kootenia is much too long-ranging to add any-
thing to the definition of the fauna under discussion.

Faunules typical of the Glossopleura zone usually occur in the
area in the lowermost portion of the Stephen formation, represented
by pure or argillaceous limestones. In the discussion of that forma-
tion, it was mentioned that in some of the sections, those at the
southwestern margin of the area investigated (the Park Mountain,
Mount Odaray, and Mount Stephen sections) beds assignable to the
Glossopleura zone seem to be absent.

However, the oldest faunule of the Glossopleura zone was col-
lected from black limestone lenses interstratified with the Cathedral
dolomite in the Skoki Valley section (locality C13r). This finding
indicates that if the undolomitized portions of the Cathedral forma-
tion were searched more extensively and carefully, new faunules
might be discovered, representing intervals that are usually barren
in the area. The writer was able to discover only two such new
faunules, the present one and the Albertella limbata faunule previ-
ously discussed.

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

The faunule from locality C13r is designated as the Glossopleura
merlinensis faunule and includes the species:

GLOSSOPLEURA MERLINENSIS FAUNULE

*Glossopleura merlinensis Rasetti.
* skokiensis Rasetti.
sp.

In the lower portion of the Stephen formation, two faunules of the
Glossopleura zone were observed. The older faunule occurs at or
near the base of the formation, and in the sections studied by the
writer appeared to consist of only one species, Glossopleura boccar,
the type of the genus. This trilobite is common at its type locality
on Mount Bosworth (author’s locality S5a). According to Deiss, in
the Castle Mountain and Ptarmigan Peak sections several other
species of trilobites are associated with Glossopleura boccar. Un-
fortunately the writer had no opportunity to examine the Stephen
formation at those localities.

GLOSSOPLEURA BOCCAR FAUNULE

*Glossopleura boccar (Walcott).

In somewhat higher beds, on Mount Temple (locality S21b) and
Mount Whyte (locality S3b), another faunule occurs, chiefly charac-
terized by the abundance of the trilobite Polypleuraspis in association
with various species of Glossopleura. The assemblages collected at
the two above-mentioned localities are united as one faunule, named
from the trilobite Polypleuraspis insignis.

POLYPLEURASPIS INSIGNIS FAUNULE

Glossopleura mckeei Resser.
* stenorhachis Rasetti.
* templensis Rasetti.
*Polypleuraspis insignis Rasetti.

The fauna of the Glossopleura zone is thus represented in the
writer’s collections from the area by only two genera, Glossopleura
and Polypleuraspis, both apparently confined to the zone. It is sur-
prising that Polypleuraspis had so far escaped observation, as it is
a common and striking fossil. The genus was previously known from
northwest Greenland, where it also occurs in association with Glosso-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 99

pleura. The United States National Museum collections contain
unidentified pygidia of Polypleuraspis from the Glossopleura zone
of Utah.

It is obvious that the collections from the area give but an extremely
incomplete idea of the composition of the fauna of the Glossopleura
zone, as numerous trilobite genera of the Cordilleran province are
known to range from the Albertella zone to the Bathyuriscus-El-
rathina zone, hence undoubtedly existed at the time when the Glosso-
pleura beds were deposited. Evidently ecologic conditions favored
associations of great numbers of individuals of very few species.

FAUNA OF THE BATHYURISCUS-ELRATHINA ZONE

As we proceed toward the later portion of the Medial Cambrian,
the characterization of successive faunas becomes more difficult and
the discussion given by the various authors increasingly contradictory
and confusing. Deiss (1939, 1940) recognized, above the Glosso-
pleura zone, an Ehmania zone (known from Montana, Wyoming,
and Utah, but not identified in the Canadian Rockies). This is suc-
ceeded by a zone which was alternatively called the “Agnostus’’-
Bathyuriscus, the Bathyuriscus-Elrathia, and the Bathyuriscus-Elra-
thina zone, and sometimes split into two zones, each characterized
by two of the above-mentioned genera. The next higher fauna was
designated by Deiss either as the Thomsonaspis fauna or as the
Olenoides (Neolenus) fauna. This fauna was stated by Deiss to be
present in the Canadian Rockies in the uppermost portion of the
Stephen formation (in the Castle Mountain and Ptarmigan Peak
sections).

In the Cambrian correlation chart (Howell et al., 1944) the Middle
Cambrian zones younger than the Glossopleura zone are designated,
in ascending order, as the Bolaspis-Glyphaspis zone, Clappaspis sub-
zone, Elrathiella-Triplagnostus subzone, Olenoides-Marjuniua zone,
Palella-Thomsonaspis zone, and Deissella-Centropleura vermontensis
zone. The lists of the genera supposed to characterize these zones
again include mixtures of forms from the Atlantic and Cordilleran
provinces assumed to be of the same age on the basis of hypothetical
correlations between the strata of Vermont and those of the Rocky
Mountains. Even if all the Atlantic province genera are disregarded,
the writer believes that the assemblages of genera from the Cordilleran
province do not constitute clear-cut faunas recognizable over much
of that area.

In the area investigated, it appears that all the faunules occurring
in the Stephen formation and younger than the Glossopleura fauna

100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

have many genera in common and may well be assigned to a single
faunal zone. This zone seems to correspond approximately to the
Elrathiella-Triplagnostus-Clappaspis zone of the correlation chart
and to the Bathyuriscus-Elrathina zone of Deiss. There are several
good reasons for discarding the first name. Elrathiella is doubtfully
distinct from Elrathia, and its genotype is a species occurring in
Greenland in strata of doubtful stratigraphic position. The genotype
of Triplagnostus occurs in the Swedish Cambrian, and the assignment
of the Cordilleran forms to the genus might be open to question.
Clappaspis, in the writer’s opinion, is not sufficiently distinct from
Ehmaniella to warrant generic recognition. On the contrary, Deiss’
designation seems well chosen, as both Bathyuriscus and Elrathina are
abundant in the faunules of the zone, and even if they are not en-
tirely confined to it, at least they do not appear to constitute important
elements of older or younger faunules. Hence the writer adopts the
term Bathyuriscus-Elrathina zone and tentatively includes in the
fauna of the zone all the faunules collected from the Stephen forma-
tion above the Glossopleura zone.

Unfortunately, in most of the sections the upper two-thirds of the
Stephen formation are almost barren of fossils. In other sections
highly fossiliferous beds are present, but usually not more than one
or two faunules could be identified from a single section. Hence,
although several faunules of the Bathyuriscus-Elrathina zone are
known (some of which are represented by an exceptional wealth of
individuals), the order of succession of the faunules presents the
most difficult problem of this kind encountered by the writer in the
area investigated. In the following discussion it will be made clear
how far the order of succession is derived from unquestionable strati-
graphic evidence and how much had to be inferred from the study
of the fossils.

To clarify the discussion, the faunules of the Bathyuriscus-Elra-
thina zone are designated as usual by a characteristic species and as-
signed an order number, believed by the writer to represent their
most probable order of succession. Faunules collected at different
localities and believed to be essentially contemporaneous, although
not identical, are listed separately and designated by the same num-
ber followed by a dash.

In ascending order, the faunules are: (1) the Ogygopsis klotsi
faunule; (2) the Olenoides serratus faunule; (2’) the Alokistocarella
fieldensis faunule; (3) the Pagetia bootes faunule; (4) the Ehmani-
ella burgessensis faunule; (5) the Bathyuriscus adaeus faunule; (6)

|

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI IOI

the Tonkinella stephensis faunule; (6’) the Parkaspis endecamera
faunule. Each of these faunules and its occurrence will be discussed
and the reasons for the assumed relative ages will be presented.

The oldest faunule of the above-mentioned list is believed to be
the faunule of the celebrated Ogygopsis shale on Mount Stephen
(locality S8d=U.S.N.M. locality 14s). In the discussion of the
Ogygopsis shale lentil it was shown that it is not possible to establish
the position of that unit in the section by stratigraphic evidence.
Hence the determination of the age of the Ogygopsis shale faunule
relative to other faunules occurring in the region must entirely depend
on the study of the fossils. The faunule is designated, according to
tradition, by the trilobite Ogygopsis klotzi, although this species has
an exceptionally long vertical range, extending from the Wenk-
chemnia-Stephenaspis zone in the Mount Whyte formation to the
Ogygopsis shale itself. It is remarkable that this genus, common at
five different horizons on Mount Stephen, was never observed else-
where in the Canadian Rockies.

In the following list of species of the Ogygopsis klotzi faunule,
the non-trilobitic forms are given as identified by Walcott, taking into
account Resser’s new generic references. Abundance data are in-
cluded for the trilobites, as the faunule was not listed in the descrip-
tion of the section, but not for the other fossils, as the writer does
not possess sufficient first-hand evidence on this point.

OGYGOPSIS KLOTZI FAUNULE

*Helcionella belliana (Walcott).

* romingert (Walcott).

* wheelerit (Walcott).
*Hyolithellus annulatus (Matthew).
a flagellum (Matthew).

*Hyolithes carinatus Matthew.
*Orthotheca corrugata Matthew.

* maior Walcott.
*Scenella amit (Matthew).
* columbiana (Walcott).

*Acrotreta depressa Walcott.

*[ phidella fieldensis Resser.

*Nisusia alberta (Walcott).

*Obolus mcconnelli (Walcott).

*Anomalocaris canadensis Whiteaves.
*Anomalocaris? acutangula Walcott.

- whiteavesi Walcott.
*Bathyuriscus rotundatus (Rominger)...... cc
*Bonnaspis stephenensis (Walcott)......... r

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

*Burlingia hectori Walcott................. rr
*Chancia pallisert (Walcott)............... r
*Elrathina cordillerae (Rominger)......... cc
Hanburia gloriosa Walcott................ rr
*Klotziella ornata (Walcott)..............- r
*K ootenia dawsoni (Walcott).............. c
*Ogygopsis klotzi (Rominger)............. ce
*Olenoides serratus (Rominger)........... cc
Oryctocephalus matthewi Rasetti .......... r
reynoldst Reed .....-0cc--: r
walkeri Matthew .......... 5
Pag etsaispis cic chines Raise poe he ee 1
*Peronopsis montis (Matthew)............. 5
*Zacanthoides romingerit Resser..........+- cc

The reasons for believing the Ogygopsis klotzi faunule to be the
oldest of the Bathyuriscus-Elrathina zone in the area will be set
forth after discussing the next faunules, which occur at three differ-
ent levels in the Burgess Quarry section.

The oldest faunule in the Burgess Quarry section occurs about
60 feet below the celebrated ‘“‘phyllopod bed” of Walcott. The locality
is indicated as St1d and the fossils are abundant but poorly preserved
in calcareous and siliceous shales. The faunule is designated as the
Olenoides serratus faunule from the most common form, and in-
cludes the species of the following list.

OLENOIDES SERRATUS FAUNULE

Paterina cf. P. zenobia (Walcott).
Nisusia sp.

Bathyuriscus rotundatus (Rominger ).
Elrathina sp.

Kootema sp.

Olenoides serratus (Rominger).
Pagetia bootes Walcott.

A similar, although apparently not identical faunule, was collected
in greater abundance 0.3 mile south of Walcott’s quarry (locality
Site). Owing to small faults covered by drift, it is difficult to as-
certain whether these beds are lower or higher than those of locality
Sr11d, but in any case the difference in age cannot be considerable.
The fossils at locality S11e weather on the surface of slabs of argil-
laceous limestone, and entire carapaces of trilobites are common. The
faunule is distinguished from the preceding and designated as the
Alokistocarella fieldensis faunule.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 103

ALOKISTOCARELLA FIELDENSIS FAUNULE

I phidella cf. I. fieldensis Resser.
* Alokistocarella fieldensis Rasetti.
Bathyuriscus rotundatus (Rominger ).
*Elrathina brevifrons Rasetti.
Kootenia burgessensis Resser.
Olenoides serratus (Rominger).
Pagetia bootes Walcott.
Solenopleurella sp. No. 2.

The next younger faunule collected from the Burgess Quarry
section occurs in the “phyllopod bed” quarried by Walcott (author’s
locality S11f=U.S.N.M. locality 35k). Notwithstanding the in-
tensive study made by Walcott and others of the exceptionally pre-
served fossils from the Burgess shale, little has been published on the
“conventional” fossils from this bed, the only ones that can be used
for the purpose of correlation. Study of the United States National
Museum collections revealed that several of the common species of
trilobites from locality 35k had never been published. The new species
are described in the paleontologic part of this paper.

The following list includes all the brachiopods described by Wal-
cott and by Resser (1938c) and the trilobites identified by the writer.
The exceptionally preserved fossils, unknown from other formations,
are left out of consideration. The faunule is designated by one of
the most common trilobites, Pagetia bootes. Abundance data are
included for the trilobites but not for the other forms for the reasons
mentioned in listing the Ogygopsis klotzi faunule.

PAGETIA BOOTES FAUNULE

*Acrothyra gregaria Walcott.
*Iphidella pulchra Resser.
*Lingulella waptaensis Walcott.
*Micromitra burgessensis Resser.
*Nisusia burgessensis Walcott.
*Paterina senobia (Walcott).

Chancta pallisert (Walcott) ...........000. f
*Ehmaniella burgessensis Rasetti ........... iF
* waptaensis Rasetti ........... r
*Elrathia permulta (Walcott).............. r

Elrathina cordillerae (Rominger)......... r
*Hanburia gloriosa Walcott.............+.. rr
*K ootenia burgessensis Resser.............- c

Olenoides serratus (Rominger)............ Cc

104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

*Oryctocephalus burgessensis Resser........ c
* matthewi Rasetti .......... r
reynoldst Reed ......c0cese0 r
*Pagetia bootes Walcott. (2.0.03 icentes en esne cc
*Parkaspis decamera Rasetti...........0005 tr
*Peronopsis montis (Matthew)...........+- r
Solenopleurella:sps, <ic\ssnlsiaetes swivereaveleloes ove i
*Triplagnostus burgessensis Rasetti........ cc

The youngest faunule from the Burgess Quarry section occurs
in fine-grained, gray siliceous shale 150-160 feet above the base of
the quarry. Walcott did not mention the occurrence of fossils at this
horizon, although certain layers are fairly covered with trilobite
fragments, belonging to three species. The faunule would not pre-
sent much interest but for the fact that it supplies part of the evidence
for determining the stratigraphic position of the Burgess shale lentil.
The faunule is designated by the trilobite Ehmaniella burgessensis
which also occurs, but much less abundantly, in the “phyllopod bed.”

EHMANIELLA BURGESSENSIS FAUNULE

Ehmaniella burgessensis Rasetti.
Pagetia sp.
Solenopleurella sp. No. I.

It is now opportune to discuss the relative ages of the faunules of
the Bathyuriscus-Elrathina zone listed so far, that is of faunules
(1), (2), (2’), (3), and (4). There is unquestionable evidence that
faunules (2), (3), and (4) occur in ascending order. Faunule (2’)
is almost identical with faunule (2) or possibly intermediate between
faunule (2) and (3). Hence the chief problem is to determine the
relative age of the Ogygopsis klotzi faunule (No. 1) with respect to
the faunules occurring in the Burgess Quarry fault block. The writer
believes that there is hardly any doubt that the Ogygopsis klotzi fau-
nule is older than any other of the above-mentioned faunules. It has
many species in common with faunules (2) and (2’), fewer with
faunule (3), and none with faunule (4), although in the last case
the evidence is less convincing, as faunule (4) includes only three
species. Of the species of faunule (2), occurring at the base of the
Burgess Quarry section, Bathyuriscus rotundatus, Elrathina sp., Koo-
tenia sp., and Olenoides serratus are identical with or are very similar
to some of the most common forms of the Ogygopsis shale. The most
striking difference between faunules (1) and (2) is that the latter
lacks Ogygopsis and includes Pagetia, which is extremely rare in the

:

a

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 105

Ogygopsis shale. The fact that Ogygopsis klotzi is already present in
the Mount Whyte and Cathedral formations adds weight to the as-
sumption that the Ogygopsis shale is older than the Burgess shale.

Accepting the above conclusion, there is still the problem of placing
all these faunules, (1) to (4), in the general section, because the
Burgess Quarry fault block is isolated and its succession of strata does
not resemble any portion of the typical sections of the Bow Range.
Hence the age of the faunules (1) to (4) relative to the other fau-
nules described in this paper must be determined from the study of the
fossils.

In the writer’s opinion, the best evidence for solving this problem
is supplied by the close resemblance of the youngest faunule of the
Burgess Quarry fault block (faunule No. 4, or Ehmaniella burges-
sensis faunule) to a faunule collected from limestone lenses in the
shaly basal portion of the Stephen formation on Mount Odaray (lo-
cality S6g). The latter faunule includes Ehmaniella burgessensis
and Solenopleurella sp. No. 1, which also occur in the Ehmamniella
burgessensis faunule in the Burgess Quarry section. Correlation of
these horizons is confirmed by the fact that the poorly fossiliferous
shaly unit that constitutes the middle portion of the Stephen forma-
tion in most of the sections of the Bow Range, at several localities
yielded cranidia of Ehmaniella. Hence it seems highly plausible to
correlate the Burgess shale with some portion of the middle, shaly
unit of the Stephen formation, which occurs in the sections of the
Bow Range above the beds of the Glossopleura zone and below the
limestone carrying younger faunules of the Bathyuriscus-Elrathina
zone (the Bathyuriscus adaeus and Tonkinella stephensis faunules)
still to be discussed. Evidence about the last-mentioned point is sup-
plied by the Mount Odaray section, where the Ehmaniella burgessen-
sis faunule occurs 110-150 feet below the Bathyuriscus adaeus fau-
nule. On the other hand, the Mount Odaray section does not supply
evidence for the relative ages of the Ehmaniella burgessensis faunule
and the Glossopleura fauna, because at that locality, as mentioned in
discussing the Stephen formation, strata assignable to the Glosso-
pleura zone seem to be missing. However, abundant evidence from
other areas in the Cordilleran province, e.g., northwestern Montana,
unquestionably proves that the Bathyuriscus-Elrathina fauna (to
which the Ehmaniella burgessensis faunule belongs) is much younger
than the Glossopleura fauna. Hence in table 3, representing the
general order of succession of the faunules, faunules (1) to (4) were
placed above the youngest faunule of the Glossoplewra zone and be-
low the Bathyuriscus adaeus faunule.

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

There remain to be discussed the faunules (5), (6), and (6’), re-
spectively designated by the trilobites Bathyuriscus adaeus, Tonkinella
stephensis, and Parkaspis endecamera. All these faunules occur in
pure or argillaceous black-gray limestone in the upper portion of the
Stephen formation in the Mount Odaray, Park Mountain, and Mount
Stephen (Fossil Gully) sections. Specimens observed in drift blocks
show the presence of faunules of this group on Mount Biddle and
Mount Field. However, in some of the best-known sections of the
Stephen formation such as the Mount Bosworth, Mount Whyte, and
Mount Temple sections, the corresponding beds, although similar in
lithology, appear to be barren.

The lowermost faunule, designated as the Bathyuriscus adaeus fau-
nule because of the great abundance of this trilobite, is known from
Mount Odaray (locality S6k), Park Mountain (locality Stok), and
Mount Stephen (locality Sok, probably identical with U.S.N.M.
locality 58j). The species include:

BATHYURISCUS ADAEUS FAUNULE

*Alokistocare paranotatum Rasetti.
*Bathyuriscus adaeus Walcott.
*Chancia odarayensis Rasetti.
*Elrathina parallela Rasetti.

* spinifera Rasetti.

Kootema sp.

Pagetia cf. P. bootes Walcott.
*Peronopsis columbiensis Rasetti.
*Tonkinella stephensis Kobayashi.

Zacanthoides sp.

On Mount Odaray, a similar faunule occurs abundantly 300 feet
higher in the section, at locality S6l. This faunule is designated by
the trilobite Tonkinella stephensis, although this species is equally
common in the preceding faunule. The identified forms are the
following:

TONKINELLA STEPHENSIS FAUNULE

*Alokistocare sinuatum Rasetti.

*Athabaskia? parva Rasetti.
Bathyuriscus adaeus Walcott.

*Chancia odarayensis Rasetti.

*Elrathina marginalis Rasetti.
Olenoides sp.

*Pachyaspis attenuata Rasetti.
Pagetia cf. P. bootes Walcott.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 107

Peronopsis columbiensis Rasetti.
Solenopleurella sp.
Tonkinella stephensis Kobayashi.
Zacanthoides divergens Rasetti.
* longipygus Rasetti.
* submuticus Rasetti.

The third faunule of this group is known only from locality Siol
on Park Mountain, where it occurs about 150 feet above the beds
carrying the Bathyuriscus adacus faunule. From this occurrence it
cannot be determined whether the present faunule is older or younger
than the Tonkinella stephensis faunule collected on Mount Odaray,
but both the stratigraphic position and the number of common forms
indicate that there cannot be any considerable difference in age. The
chief distinctive feature of the faunule collected on Park Mountain
is the abundance of the trilobite Parkaspis endecamera, unknown from
other localities, by which the faunule is designated; and also of a
species of Glyphaspis. The identified species are the following:

PARKASPIS ENDECAMERA FAUNULE

*Alokistocare cataractense Rasetti.
Bathyuriscus adaeus Walcott.

*Glyphaspis parkensis Rasetti.
Kootenia sp.

*Parkaspis endecamera Rasetti.
Peronopsis columbiensis Rasetti.
Tonkinella stephensis Kobayashi.

*Vuknessaspis paradoxa Rasetti.

*Zacanthoides divergens Rasetti.

* planifrons Rasetti.

One more faunule, younger than any so far discussed, was collected
on Park Mountain in the lower portion of the calcareous equivalent
of the Eldon formation (locality E1oc). A list of fossils from this
locality is given in the description of the Park Mountain section. From
the assemblage of trilobite genera, it appears that this faunule does not
greatly differ from those occurring in the upper Stephen formation
and still belongs to the Bathyuriscus-Elrathina zone.

The Bathyuriscus-Elrathina fauna as represented in the Stephen
formation includes the trilobite genera Alokistocare, Alokistocarella,
Bathyuriscus, Bonnaspis, Burlingia, Chancia, Ehmaniella, Elrathia,
Elrathina, Hanburia, Klotziella, Kootenia, Ogygopsis, Olenoides,
Oryctocephalus, Pachyaspis, Pagetia, Parkaspis, Peronopsis, Soleno-
pleurella, Triplagnostus, Tonkinella, Yuknessaspis, and Zacanthoides.

108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

The Agnostida first occur in this zone in the Canadian Rockies.
Alokistocare, Alokistocarella, Chancia, Kootenia, Ogygopsis, Ole-
noides, Oryctocephalus, Pagetia, and Zacanthoides are all long-ranging
genera of little stratigraphic value. Bathyuriscus and Tonkinella are
common genera that seem to be confined to the zone; so are the
rarer or more localized forms Bonnaspis, Burlingia, Hanburia, Klotz-
iella, Parkaspis and Yuknessaspis. The generalized ptychoparids
Ehmaniella, Elrathia, Elrathina, Pachyaspis, and Solenopleurella,
whose generic limits are difficult to establish, are not all confined to
the zone. However, at least Elrathina and Ehmaniella are far more
abundant in these beds than in lower or higher strata and possess strati-
graphic value.

The fauna of the Bathyuriscus-Elrathina zone as represented in
the Stephen formation is sharply distinct from the fauna of the
underlying Glossopleura zone. This fact is probably due to an un-
conformity, within the Stephen formation, between the strata of the
Glossopleura zone and those of the Bathyuriscus-Elrathina zone. The
Ehmama or Bolaspis-Glyphaspis fauna that is believed to occupy
this time interval in Montana, Wyoming, and Utah does not seem to
be represented in the Canadian Rockies.

ORDER OF SUCCESSION OF THE FAUNULES

In this section all the faunules discussed are tabulated in chrono-
logic sequence. Each faunule was named from a characteristic species
and is so designated in the list. Evidence for the relative ages of
two faunules may be either direct stratigraphic evidence, the two
faunules being collected from the same section from beds in unques-
tionable order of sequence; or indirect evidence deduced from cor-
relation of different sections or from the study of the fossils. In the
second case the order of succession may be considered probable but
there cannot be absolute certainty. In order to avoid confusing the
results from the two kinds of evidence, in table 3 the first column
includes only the faunules whose order of succession is unquestion-
able. The other columns contain faunules that were collected from
partial sections, and are interpolated in the main succession according
to the best available evidence. Hence the relative ages of faunules
in different columns are only probable. Faunules united in brackets
are essentially of the same age, and possible slight differences are
not necessarily in the order indicated.

109
Middle Cambrian

MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI
TABLE 3.—Succession of faunules

NO.
Lower
Cambrian

West Mount Stephen
Fossil Gully-North
General section Gully

Parkaspis endecamera
Tonkinella stephensis

Bathyuriscus adaeus Bathyuriscus adaeus

Bathyuriscus-
Elrathina
zone

Polypleuraspis insignis
Glossopleura boccar
Glossoplewra merlinensis

zone

Chancia bigranulosa

Albertella Glossopleura

v Yohoaspis pachycephala
8 Albertella limbata
Alberitella bosworth
ed Fieldaspis superba
232 (Fieldaspis furcata Fieldaspis furcata
aS 0 5 .
zs 8 | Fieldaspis bilobata

Plagiura cercops

Poliella prima Stephenaspis bispinosa
Syspacephalus laticeps

Wenkchemnia-
Stephenaspis
zone

zone

Bonnia fieldensis Bonnia fieldensis

Bonnia-
Olenellus

Burgess Quarry “Fossil bed”
Mount Field Mount Stephen

Ehmaniella burgessensis
Pagetia bootes
Alokistocarella fieldensis
Olenoides serratus
Ogygopsis klotzi

IIO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

SPECIES NOT DISCUSSED IN THIS PAPER

Several of the species described by Walcott, Matthew, and Resser
from the Mount Whyte, Cathedral, and Stephen formations were not
collected by the writer, and owing to uncertainty regarding their
exact stratigraphic position, it was deemed best not to use these species
in discussing the composition of the faunas. Here such species are
listed and their probable stratigraphic position is indicated. A few
species are also included which will not be dealt with in the systematic
part of this work because the writer did not have specimens for study.

“Ptychoparia” cuneas Walcott (type locality U.S.N.M. 35f, Mount
Stephen) was undoubtedly collected from the Lower Cambrian Peyto
limestone, as shown by the association of the type specimens with
olenellids.

The following species, listed with their respective type localities,
are almost undoubtedly Middle Cambrian and were collected from
various horizons in the Mount Whyte formation:

Species Type locality

Billingsella marion Walcott..........++++ 58l
Micromitra (Paterina) charon Walcott.... 61c
Nisusia (Jamesella) lowi Walcott......... 58k
Obolus damo Walcott fic ckin eine asic sicinceea se 63g
whympert Walcott .....-.sccccnes 30j
Wimanella catulus Walcott..........2see0: 63a
Alokistocare stephenense Resser........+.. 58k
Fieldaspis celer (Walcott) ...........se00- 58k
Kochaspis carina (Walcott) ...........00. 35m
cecinna (Walcott) <....sa0 creer 63a
“Olenopsis” leuka Walcott....0....0c00cce- 58¢
“Ptychoparia” adina Walcott ........++..+- 57q
clusta Walcott)... conss0ee 58k

COSSUS WAlCOtt 05 cns6 a8eenss 61a

The next four species were collected by Walcott from the “Ptar-
migan” formation on Ptarmigan Peak, hence belong either in the
Mount Whyte or the Cathedral formation. From the lower of the

two reported fossiliferous horizons in the “Ptarmigan,” at locality —

63d, Walcott listed the following two species that certainly must be
assigned to the Mount Whyte formation and more precisely to the
Plagiura-Kochaspis zone (see description of Kochiella? maxeyi in
the systematic part of this work) :

“Ptychoparia” cilles Walcott.
“Crepicephalus” chares Walcott.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI III

From the higher fossiliferous horizon in the “Ptarmigan” (lo-
cality U.S.N.M. 63b) Walcott reported the species:

Olenoides constans (Walcott).
Albertella cimon (Walcott).

If the reference of Zacanthoides cimon Walcott to Albertella, sug-
gested by Resser, is correct (as the writer is inclined to believe, al-
though the species is known only from cranidia and there can be no
certainty in the reference to Albertella without the pygidium), then
the higher fossiliferous horizon in the “Ptarmigan” would be close
to that of the Ross Lake shale which is not developed in the Ptarmigan
Peak section. The writer has already reassigned these beds of the
“Ptarmigan” to the Cathedral formation.

Glossopleura stephenensis Resser is based on a flattened, poorly
preserved, specifically unidentifiable shield of Glossopleura supposedly
collected from the Ogygopsis shale on Mount Stephen. Since the
writer has never seen any representative of the genus in the Ogygopsis
shale, he suspects that the specimen has been mislabeled.

Bathyuriscus pupa Matthew and Dolichometopus occidentalis Mat-
thew, both from the Ogygopsis shale, were considered by Walcott and
Resser as synonymous. The writer has not seen the types, and the
species is not represented either in the United States National Mu-
seum collections or in the writer’s own. Resser (1935, p. 44) states
that “two excellent carapaces have just been discovered in our col-
lections.” Upon inspection, the two specimens (on one slab) were
found to be labeled from locality 35k (Burgess shale) instead of 14s
(Ogygopsis shale) as stated by Resser. Furthermore, the matrix re-
sembles neither the Burgess shale nor the Ogygopsis shale. These
specimens represent Poliella denticulata, described in this paper from
the Mount Whyte formation on Mount Stephen, hence it is virtually
certain that they do not come from the Ogygopsis shale; they were
probably collected from talus at the base of Mount Stephen.

Oryctocephalus walkeri Matthew is another form from the Ogygop-
sis shale whose exact characters the writer was unable to ascertain.
Probably even examination of the types would leave some uncertainty,
as the specimens appear to be poorly preserved from Matthew’s de-
scription. Examination of collections made by the writer and those
of the United States National Museum seems to indicate the presence
of three species of Oryctocephalus in the Ogygopsis shale, two of
which are definitely O. reynoldsi and O. matthewi. The individuals
that do not appear to belong to either species and may represent O.
walkeri are too poor to show clearly the specific characters. The latter

II2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

species may prove to be identical with O. burgessensis Resser from
the Burgess shale, in which case Resser’s name would become a
synonym. In the meantime it seems better to use the name burges-
sensis for the well-represented form from the Burgess shale.

GEOGRAPHIC AND VERTICAL DISTRIBUTION OF TRILOBITES

In this section the writer wishes to discuss the vertical range and
geographic distribution of some of the trilobite species, genera, and
families encountered in the formations described in this paper.

The Olenellidae occur in extreme abundance in the uppermost beds
here assigned to the Lower Cambrian, and suddenly disappear, indi-
cating a probable unconformity as it has been mentioned in discussing
the contact between the St. Piran sandstone and the Mount Whyte
formation, and the Lower-Middle Cambrian boundary.

The earliest observed occurrence of the Agnostida is an unusually
late one, since no agnostid was found in beds older than the middle
portion of the Stephen formation ; probably the oldest one is Peronop-
sis montis from the Ogygopsis shale. Furthermore, the agnostids
show little variety and a small number of species even when they
become abundant in numbers of individuals. In other areas of the
Cordilleran province, e.g., in Idaho, agnostids are known from con-
siderably older beds, such as the Langston (?) formation (Resser,
1939b). In the Appalachian province they occur, though rarely, even
in late Early Cambrian time. It is clear the Canadian portion of the
Cordilleran geosyncline was one of the last seaways of the world to
be invaded by the Agnostida.

The Eodiscida, recognized by the writer as an order, seem to be
represented in the Cordilleran province exclusively by forms with
eyes and free cheeks (Pagetia). The blind Eodiscida, common in the
Atlantic and Appalachian provinces both in the Lower and Middle
Cambrian, are here totally absent. Pagetia appears in the Lower Cam-
brian Peyto limestone; a form collected by the writer from those
strata is closely similar to Pagetia ellsi Rasetti and other Middle Cam-
brian species. The presence of Pagetia in the Lower Cambrian had
already been noticed by the writer in the Appalachian province (Ra-
setti, 1948a). In the area investigated, Pagetia was never observed
in the Mount Whyte and Cathedral formations of normal lithology ;
for example, it was never seen among the thousands of trilobites filling
certain layers of the Ross Lake shale, although it existed both before
and after the time when the shale was deposited. The same remark
applies to many trilobite genera. The next higher occurrence of

-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI IT3

Pagetia is in the peculiar silty limestone that replaces a portion of the
Cathedral dolomite in the western part of Mount Stephen. The fau-
nule in which Pagetia occurs is of somewhat uncertain age, and is here
tentatively considered younger than the Albertella bosworthi faunule.
Pagetia becomes common in the Stephen formation.

The family Dorypygidae is represented in the Lower Cambrian by
the genus Bonnia, common in the Peyto limestone as in most of the
limestones of late Early Cambrian age in the Cordilleran and Appala-
chian provinces. Kootenia and Olenoides appear much later, although
they are already present in the Lower Cambrian of the Appalachian
province. Kootenia was first observed in the anomalous equivalent
of the Cathedral formation on Mount Stephen, and Olenoides does
not seem to be present in strata older than the middle portion of the
Stephen formation. There is no doubt that Kootenia existed in the
Cordilleran province in strata both older and younger than the Ross
Lake shale, yet this is another genus totally absent from that unit.

The family Oryctocephalidae is represented by Oryctocephalus,
Oryctocephalites, and Tonkinella. Oryctocephalus appears in the
Wenkchemnia-Stephenaspis zone and is evidently long-ranging, as it
occurs again in the Stephen formation. As in the preceding cases,
this genus shows a wide gap in its vertical distribution in the area,
being apparently absent from the Cathedral formation. Of Orycto-
cephalites we know an isolated occurrence in the Plagiura-K ochaspis
zone. Tonkinella seems to be strictly confined to the upper Stephen
formation and to be an excellent index fossil.

The families Dolichometopidae and Zacanthoididae may be treated
together as the boundary between them is to a large extent arbitrary.
In the Lower Cambrian Peyto limestone the genus Zacanthopsis was
found; no forms definitely assignable to the Dolichometopidae were
collected. Such forms, however, are known to be associated with
Olenellidae in northwest Greenland (Poulsen, 1927) and in Quebec
(Rasetti, 1948a). Zacanthoides appears in strata equivalent to a por-
tion of the Cathedral formation in the western part of Mount Stephen.
Several dolichometopid genera (Wenkchemmia, Poliella, Stephenaspis,
Fieldaspis) occur abundantly in the Wenkchemnia-Stephenaspis and
Plagwura-Kochaspis zones and seem to be lacking or extremely rare
both in older and younger beds. Albertella, Vanuxemella, Ptarmigania,
and Ptarmiganoides seem to be confined to beds of the Albertella zone.
Bathyuriscus first appears in the Stephen formation and is there rep-
resented by several species and extremely numerous individuals.
Glossopleura and Polypleuraspis are abundant in the Glossopleura
zone and absent from younger beds.

II4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The genus Ogygopsis deserves special discussion. Individuals that
the writer must assign to a single species, since no morphologic dif-
ferences were observed, were collected in strata of five different ages
ranging over an exceptionally long time interval. Ogygopsis klotat
first appears and is very abundant in the Syspacephalus laticeps fau-
nule of the Wenkchemnia-Stephenaspis zone. The species was found
to be extremely rare in the next younger assemblage, the Stephenaspis
bispinosa faunule of the same zone. It was never observed in the
strata of the overlying Plagiura-Kochaspis zone. It then reappears
abundantly as a member of the Yohoaspis pachycephala and Chancia
bigranulosa faunules in the equivalent of a portion of the Cathedral
formation. Ogygopsis is totally absent from the beds of the Glosso-
pleura zone. It is then extremely abundant in the Ogygopsis klotzi
faunule (Ogygopsis shale), which occurs in the Stephen formation
and is younger than the Glossopleura fauna. Ogygopsis klotzi is as-
sociated with five different assemblages in these different occurrences.
All these occurrences are on Mount Stephen; evidently Ogygopsis was
an inhabitant of the Goodsir trough, southwest of the Bow trough, and
only occasionally was carried by the waters to the extreme south-
western margin of the Bow trough. Other forms, unknown in the
typical Middle Cambrian sections of the Bow Range, occur in the
anomalous assemblages of the Yohoaspis pachycephala and Chancia
bigranulosa faunules on Mount Stephen.

The last group of trilobites still to be discussed is the Ptychopari-
oidae. This superfamily is not divided by the writer into families be-
cause of the great difficulties presented by the taxonomy of the group.
The lack of satisfactory family divisions and the arbitrary definition
of most of the ptychoparid genera renders them poor index fossils,
to be avoided for zone definition whenever anything else is available.
Generalized ptychoparids (Piazella, Onchocephalus, Syspacephalus)
appear in the Lower Cambrian Peyto limestone and occur through the
overlying Middle Cambrian formations. Most of the Middle Cambrian
forms are referred to other genera, but whether this is done or not,
little or no significance for correlation should be attributed to general-
ized ptychoparids. There are, however, a few distinctive Middle
Cambrian ptychoparids that do have stratigraphic significance.

The long-brimmed, flat, micropygous forms (here assigned to
Amecephalus and Alokistocare; partly referred in the literature to
Kochiella or Strotocephalus) were first observed in the W enkchemmia-
Stephenaspis zone. In Greenland, however, Poulsen (1927) reports
Kochiella from the Cape Kent formation in association with Olenel-
lidae. These long-brimmed ptychoparids are common in the Stephen

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI II5

formation, but were never observed in the Albertella zone, unless
Mexicella be considered a member of the group.

Two very distinctive ptychoparid genera are Plagiura and Schisto-
metopus, both collected only from the beds of the Plagiwra-Kochaspis
zone.

A difficult problem, discussed in the systematic part of this paper,
is presented by the genera Kochiella and Kochaspis on account of the
uncertain assignment of the pygidia to the cranidia. Until Kociiella
is better understood it is preferable not to use it for zone designation.
Kochaspis is well based on the genotype Kochaspis liliana, whose head
and tail are properly assigned. For this reason, Kochaspis was used
with Plagiura for zone designation. However, some of the other
described species of Kochaspis were undoubtedly based on arbitrary
combinations of ptychoparid heads and bathyuriscid tails.

A general conclusion that can be derived from these remarks on
the occurrence of trilobite families and genera is that often even
highly fossiliferous formations appear entirely to lack fossils that
are known to range through the corresponding stratigraphic intervals
and to occur in beds of similar lithology elsewhere. Hence very little
significance should be attributed to the absence of genera or families
from certain strata. A typical example is that of the Ross Lake shale
member of the Cathedral formation, a siliceous shale with limestone
beds and nodules. No more than seven species of trilobites, represent-
ing five genera, are known from this extremely fossiliferous unit.
Evidently ecologic conditions favored this small assemblage of species
to the exclusion of a great number of other trilobites that lived at the
time. To mention only a few examples, there is no doubt that Koo-
tenia, Olenoides, Oryctocephalus, Pagetia, Zacanthoides, Ogygopsis,
and a host of ptychoparid genera lived in the Cordilleran province at
the time when the Ross Lake shale was deposited. Hence it would
not be surprising to find elsewhere a faunule exactly equivalent in
age to that of the Ross Lake shale and yet not having any species
or even genera in common. The stratigrapher must constantly bear in
mind the fact that while the presence of several common species in
strata of different areas almost certainly indicates a close time equiva-
lence, lack of common forms does not prove difference in time.

Table 4 indicates the presence of the trilobite genera discussed in
this paper in the various zones (Bonnia-Olenellus zone of the Lower
Cambrian and Wenkchemnia-Stephenaspis, Plagiura-Kochaspis, Al-
bertella, Glossopleura, Bathyuriscus-Elrathina zones of the Middle
Cambrian). The data in this table refer exclusively to the area in-
vestigated, hence the presence of genera in the same zones in other
areas is not reported.

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Taste 4.—Range of trilobite genera in the Canadian Rockies

8.2 . &
33 33 Ss 2 vo $ o =3 o
Soe 7 eee) ee 38 Z8 $88
SON RAR = NR =N =A SON
Genus Q S X aN S FQ
Albertella) ... 32s x
Alokistocare ....... x
Alokistocarella ..... x
Amecephalus ....... x x
Athabaskia ........ ? ?
Bathyuriscus ....... x
Bonnaspis osceisese x
BORNAD Sera iNe cose x
BUPlingia .ccmcles on : x
Gaborcella acco « x
Ghancia Bosaececees x ? x
Crassifimbra ....... x
Ehmaniella ........ xX
Er Gthia raster eloteteiax ae
Elrathina .......++. x
Fieldaspis .....0++. x:
Glossopleura ....... x
Hanburia w...0..s6 De
Klotsrellawscy.ictrakele ae 5
TK OCRGSPAS! sieve teva.siache x
Keochtella Ze arieecete x
TSOCHING S cactecyia ese 38
IKOGLENIA® mene nes ee ? x
WIC AL eau apadoe a
OGVGOPSIS 2. cee. : Bi ? 3
Olenelluscdeciaesc x
Olenoideswemce cece x
Onchocephalus ..... be x
Oryctocephalites ... x
Oryctocephalus .... 3% x
Pachyaspis j«.0s0s0s x
Paedeumias ........ x
Pagetia...... eEaeiet x ? x
POrRaspis jerccie» cae x
IPCrOnopsis Soe.ce se 5.2
PAGE CllON eran teres eX x
PU GQUIO ie nrc cto ees x0 x
IRA ANG MAD OOCOORe x
Polypleuraspis ..... x |
Ptarmigania ....... i:
Ptarmiganoides .... x
Schistometopus ..... x
Solenopleurella ..... x

Stephenaspis ....... x

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI TE.

TABLE 4.—Continued

3.2 :
6 eS 2 os S 2 3
Cia Qos S48 net =N Bs ERS

Syspacephalus ...... x x
Triplagnostus ...... x
MOWRNEUIG se slices ox
Vanuxemella ....... D3
Wenkchemnia ...... BG
PV ONOGSPAS | oc cic csi ve ?
Yuknessaspis ...... x
Zacanthoides ....... ? x
Zacanthopsis ....... mS

ENVIRONMENT OF DEPOSITION

This section includes a brief discussion of the ecology of the Mid-
dle Cambrian deposits and the manner of preservation of the fossils.

Fossils occur in the rocks investigated in several kinds of matrix,
which can be divided into three general lithologic types: (1) crystal-
line or oolitic, thin-bedded limestones; (2) argillaceous, platy lime-
stone; (3) fine-grained, siliceous shales. Massive carbonate forma-
tions (e.g., most of the Cathedral formation) appear barren of fossils
(apart from the algal concretions designated in the literature as
Girvanella) even when represented by calcareous limestone where the
fossils would be expected to be well preserved if they had originally
been present.

The thick Lower Cambrian series in the area consists almost en-
tirely of sandstones, quartzites, and siliceous shales, and is almost
barren of fossils, excepting numerous worm trails and burrows. The
shaly intervals were carefully searched at various localities, in view
of the abundant fossil content reported for beds of this age and lithol-
ogy elsewhere, but yielded few identifiable fossils. A trilobite faunule
collected from quartzitic beds in the lower St. Piran sandstone was
mentioned in discussing that unit. In view of the abundant traces
of animal life in the less coarse portions of the Lower Cambrian
clastics, the scarcity of trilobite shields, brachiopod shells, and other
identifiable remains is probably due to the destructive action of
scavenging organisms.

Fossils appear abundantly near the top of the Lower Cambrian
with the first deposition of carbonates. The uppermost St. Piran beds
gradually change to calcareous sandstone, sandy limestone, and finally

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

pure, crystalline limestone ; the calcareous sediments in many instances
first appearing as lenses in the sandstone. This Lower Cambrian
limestone (Peyto limestone member of the St. Piran sandstone) is
often filled with comminuted tests of olenellid trilobites. The smaller,
thick shells of Bonnia and ptychoparid trilobites are usually much
better preserved. The conditions of fossilization point to a very low
rate of sedimentation for these calcareous Lower Cambrian beds.

The lowermost Middle Cambrian unit (Mount Whyte formation)
first consists to a large extent of siliceous shale with thin interstrati-
fied sandstone beds, succeeded by alternating shale and limestone;
the latter coarsely crystalline or oolitic and predominating in the
upper part of the formation. Most of the shale shows worm trails
and burrows and appears barren of identifiable fossils. However,
local deposits of very fine, siliceous shale lack such worm burrows
and yield numerous well-preserved trilobites and other fossils. These
localized deposits, here designated as the Lake Agnes and Yoho shale
lentils, are approximately of the same age.

The Lake Agnes shale is a blue-green shale yielding chiefly trilo-
bites, accompanied by small inarticulate brachiopods and rare Hyo-
lithes. Most of the trilobites are fragmentary, but complete shields
of the four species were recovered. Some of these shields preserve
the free cheeks while others do not. It is generally assumed that in
the first case the living animals, in the second case moults, were en-
tombed. Early larval stages were not observed.

The Yoho shale is far more fossiliferous than the Lake Agnes
shale, and small inarticulate brachiopods, gastropods (Scenella),
and Hyolithes are almost as abundant as the most common trilobites.
The conditions of fossilization of the trilobites are analogous to those
observed in the Lake Agnes shale.

The Cathedral formation yielded fossils at a few localities and
horizons from its calcareous beds, but most of the fossils reported
from the formation were collected from the Ross Lake shale member.
This fine-grained, siliceous shale averages only 5-6 feet in thickness
but covers a vast area and contains trilobites and other fossils in
great abundance; it represents an interval of clastic sedimentation
underlain and overlain by several hundred feet of almost pure car-
bonates. Besides the trilobites, brachiopods and Hyolithes are com-
mon. Articulated trilobite carapaces were collected for all the seven
species known from the shale; however, the percentage of such
shields is probably less than one in a thousand compared to dismem-
bered fragments. Both moults and living animals were entombed.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI II9Q

Larval forms in various stages of growth are associated with the
adult trilobites, but the preservation is not good enough to allow a
profitable study of the smaller individuals.

A remarkable feature of several faunules preserved in shale, and
particularly of the Albertella bosworthi faunule of the Ross Lake
shale, is the small number of species compared to the number of indi-
viduals. Many thousands of trilobite specimens were examined by the
writer on the surface of slabs of the Ross Lake shale, yet no ad-
ditional species were discovered. The collections from limestone beds,
owing to the difficulty of examining and recovering fossils that do not
appear on weathered surfaces, are much smaller, yet often yielded
larger numbers of species than the collections from shaly units, and
probably many more forms would be revealed by further collecting.
Apparently the ecologic conditions obtaining on the sea bottom where
the Ross Lake shale or similar sediments were deposited favored as-
semblages of a few very prolific species. The Ross Lake shale is
remarkable not only for the abundance of a few species but also for
the apparently total absence of many trilobite genera that are known
to have existed in the Cordilleran province at the time when the shale
was deposited and occur in strata of similar age and lithology else-
where.

The basal portion of the Stephen formation carries faunules pre-
served in argillaceous or pure, fine-grained or crystalline limestone.
The trilobite shields, usually representing only the two genera Glosso-
pleura and Polypleuraspis, are generally concentrated in certain layers,
sometimes covering the upper surfaces of the limestone beds. Entire
trilobite shields are rare.

The Ogygopsis shale includes beds of different lithologies, as men-
tioned in discussing that unit; all of them are crowded with fossils,
especially the five most abundant species of trilobites. The gastropod
Scenella and the peculiar crustacean Anomalocaris are next in abun-
dance. There is an exceptionally high percentage of articulated trilobite
carapaces, but most of them lack the free cheeks, hence are assumed
to represent moults. Such conditions of preservation seem to indicate
an exceptionally high rate of sedimentation as the delicate trilobite
moults, and especially Anomalocaris, could not have remained long
unburied on the sea bottom without falling apart. The absence of
scavenging organisms must have been another essential factor that
made possible the preservation of such great numbers of fossils.
Lithologically, the varieties of Ogygopsis shale seem identical with
the barren siliceous shales common in the Middle Cambrian of the
area. Most of the trilobite shields are of medium or large sizes.

120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The Burgess shale and the conditions under which it was deposited
were extensively discussed by Walcott (1912a) and a few remarks
will suffice. Trilobites, like other fossils, are not abundant, but the
percentage of articulated carapaces is higher than in any other sedi-
ment known to the writer. Furthermore, most of the shields preserve
the free cheeks, hence do not appear to represent moults. Most of
the trilobites are from half to full grown, early larval stages being
rare.

The uppermost, calcareous portion of the Stephen formation carries
abundant fossils at several localities, while similar beds elsewhere
seem barren. The fossils occur in a black-gray, fine-grained, more
or less argillaceous limestone. Articulated trilobite shields are not
uncommon and larval stages are frequently represented.

In two instances it was possible to obtain information on the de-
pendence of faunal assemblages upon the lithology of the sediments.
The beds of the Plagiura-Kochaspis zone of the Mount Whyte
formation on Eiffel Peak (locality W20d) consist of interstratified
crystalline limestone and fine-grained, siliceous shale. The Fieldaspis
bilobata faunule of these beds showed little difference in the relative
abundance of the various species between the two types of matrix.

Another case where the composition of a faunule in limestone and
shale could be compared is offered by the Ross Lake shale, which at
several localities (e.g., Ross Lake, Mount Bosworth, Bow Lake)
includes near the middle or at the top beds of pure, crystalline lime-
stone. In the shaly portion, by far the most common fossils are species
of Albertella, Mexicella stator, and Vanuxemella nortia. All these
forms are also common in the limestone, in addition to Ptarmigania
rossensis, which is a rather rare fossil in the shale. Here again the
difference in composition of the faunule is slight. Albertella has been
considered a representative fossil of shaly beds and stated to be rare
in limestone deposits (Lochman, 1947). The writer’s observations
indicate that the only reason why Albertella is usually represented by
shale specimens in the collections is that such specimens are much
more readily noticed in the field and can be collected in great numbers
with far less expense of time and work than are required to obtain
fossils from limestone. The number of fossils per unit volume of
the rock did not appear to be smaller in the limestone than in the shale.

These remarks should not be interpreted to mean that the compo-
sition of a faunule is unaffected by the environment of deposition,
but rather that in the above-mentioned instances there was essentially
a single environment, independent of the deposition of clastic or car-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI I2I

bonate sediments. This is made plausible by the fact that limestone
layers interbedded with the shale are usually lenticular, hence lime-
stone deposition took place in small, shifting patches on a prevalently
argillaceous sea floor. It is not surprising that under such conditions
we do not find different types of organisms confined to each kind of
sediment.

INDEX, OF LOCALITIES

Table 5 includes a list of the author’s localities from which fossils
were described. The first column gives the locality number, the second
column the topographic location (see also localities indicated on maps,
figs. 1 and 3). The third column gives the stratigraphic position and
lithology. The fourth column gives the approximate altitude in feet.

Table 6 shows the correspondence between all the United States
National Museum localities mentioned in the description of the fos-
sils and the equivalent author’s localities.

TABLE 5.—Topographic and stratigraphic position of author’s localities

Approxi-
mate
Stratigraphic position altitude
Locality Topographic position and lithology Feet
litsjregaresn & Northwest slope of Mount St. Piran sandstone (Peyto 4,700
Stephen, about 4 mile limestone member) : top 6
west of the Monarch feet of limestone. Dark-
Mine, directly above the gray, crystalline limestone.
second of the three sheds
of the Canadian Pacific
Railway; Yoho Park,
British Columbia.
Ee 2IG” isicacs . West ridge of Mount Schaf- St. Piran sandstone (Peyto 7,800
fer, above McArthur limestone member ) : near
Pass; 0.4 mile W. 30° S. base of unit 4 of the
of the summit of Mount Mount Schaffer section,
Schaffer; Yoho Park, about 110 feet above base
British Columbia. of member. Dark-gray,
crystalline limestone.
2-771 ee Same as P22k. St. Piran sandstone (Peyto 7,800

limestone member): top
10 feet of unit 4 of Mount
Schaffer section, about
160 feet above base of
member. Dark-gray, crys-
talline limestone.

Wiebe concer South slope of east ridge of Mount Whyte formation: 8,000
Mount Niblock; 0.3 mile unit 2 of Mount Niblock
NW. of the upper end of section, 15-20 feet above
Lake Agnes; Banff Park, base of formation. Blue-
Alberta. green-gray, fine-grained

siliceous shale.

I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

TABLE 5.—Continued

Approxi-
mate
Stratigraphic position altitude
Locality Topographic position and lithology Feet
Webiwecens. Southeast slope of Popes Mount Whyte formation: 6,800
Peak; 0.2 mile west of lower 5 feet of unit 3 of
Plain of Six Glaciers Mount Whyte section,
chalet; Banff Park, Al- 22-27 feet above base of
berta. formation. Blue-green-
gray, fine-grained siliceous
shale.
Wise’ .voeance Same as W3b. Mount Whyte formation: 6,900

unit 5 of Mount Whyte
section, 100-112 feet above
base of formation. Dark-
gray oolitic limestone.
WAC sa kere Slope 0.2 mile southeast of Mount Whyte formation: 6,300
upper end of Ross Lake; 3-5 feet above base of
Yoho Park, British Co- unit 4 of Ross Lake sec-
lumbia. tion, 162-164 feet above
base of formation. Dark-
gray, oolitic limestone.
Wace cn Same as W4c. Mount Whyte formation: 6,300
unit 4 of Ross Lake sec-
tion, 160-182 feet above
base of formation. Dark-
gray, oolitic limestone.
Wiad eesceeen Same as W4c. Mount Whyte formation: 6,300
5 feet above base of unit
5 of Ross Lake section,
187 feet above base of for-
mation. Dark-gray, crys-
talline or oolitic limestone.

Wad" soviet Same as W4c. Samehorizonas W4d. Dark- 6,300
gray, crystalline limestone.
Wiser eenciate Same as W4c. Mount Whyte formation: 6,400

near top of unit 5 of Ross
Lake section, about 235
feet above base of forma-
tion. Dark-gray, crystal-
line limestone.

Wsby 23nae South slope of Mount Bos- Mount Whyte formation: 6,000
worth; 0.4 mile north of unit 2 of Mount Bos-
western end of Sink Lake; worth section, 24-60 feet
Yoho Park, British Co- above base of formation.
lumbia. Outcrop separated by

fault from measured sec-
tion. Light-gray, crystal-
line limestone lenses in
shale.

NO. 5

Locality
VIS Gave aeeicre

MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

TABLE 5.—Continued

Topographic position

East ridge of Mount Bos-

worth; 0.9 mile N. 15°
E. of western end of Sink
Lake ; Yoho Park, British
Columbia.

East slope of south peak of

Mount Odaray; 0.5 mile
east of summit of south
peak; Yoho Park, British
Columbia.

W7c .......Southeast slope of Mount

Field; just east of portal
of Kicking Horse Mine;
Yoho Park, British Co-
lumbia.

W7d).2...2eoame.as Wc.

Same as W7c.

Wiis. <sos. ame as VV 7c:

Same as W7c.

Stratigraphic position
and lithology

Mount Whyte formation:
50 feet above base of unit
5 of Mount Bosworth
section, 254 feet above
base of formation. Dark-
gray, crystalline or oolitic
limestone.

Mount Whyte formation:
top 3 feet of unit 3 of
Mount Odaray section,
158-161 feet above base of
formation. Very dark-
gray, finely crystalline
limestone.

Mount Whyte formation:
top 3 feet of unit 1 of
Kicking Horse Mine sec-
tion, 190-193 feet below
top of formation. Dark-
gray, crystalline or oolitic
limestone.

Mount Whyte formation:
basal 2 feet of unit 2 of
Kicking Horse Mine sec-
tion, 188-190 feet below
top of formation. Dark-
gray, crystalline lime-
stone.

Mount Whyte formation:
10 feet above base of unit
2 of Kicking Horse Mine
section, 180 feet below top
of formation. Dark-gray,
crystalline or oolitic lime-
stone.

Mount Whyte formation:
72 feet above base of unit
2 of Kicking Horse Mine
section; 118 feet below
top of formation. Dark-
gray, crystalline lime-
stone.

Mount Whyte formation:
unit 2 of Kicking Horse
Mine section, more precise
horizon unknown (isolated
outcrop east of measured

123

Approxi-
mate
altitude
Feet

7,000

8,200

4,700

4,700

4,700

4,800

4,700

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

TABLE 5.—Continued

Approxi-
: t e mate
Stratigraphic position altitude
Topographic position and lithology Feet

section traverse). Dark-

unit 2 of North Gully sec-
tion; 142-158 feet above
base of formation. Gray,
siliceous shale and lenses
of gray, crystalline lime-
stone.

gray, crystalline lime-
stone.

Same as W7c. Mount Whyte formation: 4,500

unit 2 of Kicking Horse
Mine section, more pre-
cise horizon unknown
(talus). Dark-gray, crys-
talline limestone.

Same as W7c. Mount Whyte formation: 4,800
40-50 feet above base of
unit 4 of Kicking Horse
Mine section, 60-70 feet
below top of formation.
Very dark-gray, finely
crystalline limestone.

Southwest side of Fossil Mount Whyte formation: 6,000
Gully ; northwest face of 25-80 feet above base of
Mount Stephen, Yoho unit 4 of Fossil Gully
Park, British Columbia. section. Dark-gray, fine-

grained siliceous shale.

About 4 mile west of lo- Same as locality Wok. 6,500
cality Wok. Dark-gray, fine-grained

siliceous shale.

Southwest and northeast Mount Whyte formation: 5,800
sides of Fossil Gully; from 4 above the base and
northwest face of Mount to top of unit 11 of Fossil 6,300
Stephen, Yoho Park, Gully section; o-60 feet
British Columbia. below top of formation.

Dark-gray, crystalline
limestone.

South slope of Ptarmigan Mount Whyte formation: 7,700
Peak, above Ptarmigan 120 feet above base of for-

Pass; Slate Mountains, mation. Dark-gray, oolitic
Banff Park, Alberta. limestone.

Northeast side of North Mount Whyte formation: 5,200
Gully ; northwest face of top 3 feet of unit 1 of
Mount Stephen; Yoho North Gully section, about
Park, British Columbia. 140 feet above base of for-

mation. Dark-gray, crys-
talline limestone.
Same as Wi6h. Mount Whyte formation: 5,200

NO. 5 MIDDLE CAMBRIAN STRATIGRAPH Y—RASETTI 125
TABLE 5.—Continued
Approxi-
mate
Stratigraphic position altitude

Topographic position

Same as W16h.

Same as W16h.

Same as W16h.

Same as W16h.

North shoulder of Mount

Stephen, directly above
the Canadian Pacific Rail-
way tunnel; Yoho Park,
British Columbia.

Same as Wi7a.

West face of northwest spur
of Mount Hector ; 0.8 mile
northeast of the lower end
of Hector Lake; Banff
Park, British Columbia.

and lithology

Mount Whyte formation:
top 20 feet of unit 3 of
North Gully section, 242-
262 feet above base of
formation. Dark-gray sili-
ceous shale and dark-
gray, crystalline or fine-
grained, silty limestone.

Mount Whyte formation:
unit 3 of North Gully sec-
tion (more precise posi-
tion unknown). Dark-
gray, siliceous shale and
fine-grained, dark-gray,
silty limestone.

Mount Whyte formation:
40-50 feet above base of
unit 3 of North Gully sec-
tion; 198-208 feet above
base of formation. Dark-
gray, siliceous shale and
fine-grained, dark-gray,
silty limestone.

Mount Whyte formation:
20-30 feet above base of
unit 4 of North Gully sec-
tion, 282-292 feet above
base of formation. Dark-
gray, fine-grained _ sili-
ceous shale and lenses of
dark-gray, crystalline
limestone.

Mount Whyte formation:
18 feet above base of unit
1 of Monarch Mine sec-
tion. Dark-gray lime-
stone.

Mount Whyte formation:
top 5 feet of unit 3 of
Monarch Mine section,
162-167 feet above base
of formation. Dark-gray
crystalline limestone.

Mount Whyte formation:
near middle of unit 2 of
Hector Creek section;
about 235 feet above base
of formation. Dark-gray,
oolitic limestone.

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS

TABLE 5.—Continued

Locality Topographic position

W200i icin. c Southeast ridge of Eiffel
Peak; 0.3 mile northeast
of the eastern end of
Eiffel Lake; Banff Park,
Alberta.

S3M) Sievsiocarete South ridge of Mount
Whyte; 0.3 mile north of
Plain of Six Glaciers
chalet; Banff Park, Al-
berta.

(Ci pmacot Northwest slope of the
north spur of Popes peak;
0.3 mile southeast of the
upper end of Ross Lake;
Yoho Park, British Co-
lumbia.

Gomis ace East ridge of Mount Bos-
worth; 1.0 mile north of
the western end of Sink
Lake; Yoho Park, British
Columbia.

Cons 4. same Southwest and _ northeast
sides of Fossil Gully;
northwest face of Mount
Stephen; Yoho Park,
British Columbia.

Gr satxei ne Northeast side of Fossil
Gully; northwest face of
Mount Stephen; Yoho
Park, British Columbia.

(20) eee Northeast side of Fossil
Gully; northwest face of
Mount Stephen; Yoho

Park, British Columbia.

Stratigraphic position
and lithology

Mount Whyte formation:
limestone 22-28 feet above
base of formation, and
shale immediately above
and below. Dark-gray,
crystalline limestone and
green-gray, fine-grained
siliceous shale.

Cathedral formation (Ross
Lake shale member):
unit 4 of Mount Whyte
section, 463-468 feet above
base of formation. Dark-
blue-gray, fine-grained
siliceous shale.

Cathedral formation (Ross
Lake shale member):
410-418 feet above base
of formation. Dark-blue-
gray, fine-grained sili-
ceous shale and _inter-
bedded gray limestone.

Cathedral formation (Ross
Lake shale member):
385-304 feet above base
of formation. Dark-blue-
gray, fine-grained _ sili-
ceous shale.

Cathedral formation: 30-
70 feet above base of unit
2 of undivided Cathedral
and Stephen formations
of Fossil Gully section.
Dark-gray, silty lime-
stone and dark-gray, sili-
ceous shale.

Cathedral formation: 35
feet below top of unit 3
of undivided Cathedral
and Stephen formations
of Fossil Gully section.
Dark-gray, subcrystalline
limestone.

Cathedral formation: top
of unit 3 of undivided

Cathedral and Stephen
formations of Fossil
Gully section. Dark-

gray, subcrystalline lime-
stone.

VOL. 116

Approxi-

mate

altitude

Feet

8,500

8,100

6,700

7,400

5,900
and

6,400

6,000

6,000

NO. 5

MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

TABLE 5.—Continued

Locality
Cok

Topographic position

Reeser Northeast side of Fossil
Gully ; northwest face of
Mount Stephen; Yoho
Park, British Columbia.

Wewarls South slope of Ptarmigan
Peak, above Ptarmigan
Pass; Slate Range, Banff
Park, Alberta.

se caste Skoki valley ; 1 mile N. 15°
W. of the summit of Pika
Peak; Banff Park, Al-
berta.

Beal ohek East slope of unnamed
mountain on west side of
Bow Valley, north of
Bow Lake; 1.0 mile
northwest of Num-Ti-Jah
Lodge; Banff Park, Al-
berta.

Lveateaters Same as Cr5m.

ciate teeta Southeast ridge of Eiffel
Peak; 0.4 mile northeast
of the eastern end of
Eiffel Lake; Banff Park,
Alberta.

Stratigraphic position
and lithology

Cathedral formation: base
of unit 5 of undivided
Cathedral and Stephen
formations of Fossil Gully
section. Black-gray, sub-
crystalline limestone.

Cathedral formation: Deiss’
23-feet unit of “Ptarmigan
formation.” According to
reinterpretation of Deiss’
section, this unit is 363-
386 feet above base of Ca-
thedral formation. Fine-
grained, black-gray lime-
stone.

Cathedral formation: 270-
275 feet above base of
dolomitic portion of for-
mation; probably about
650 feet above base of Ca-
thedral formation. Black-
gray, fine-grained lime-
stone.

Cathedral formation (Ross
Lake shale member):
unit 5 of Bow Lake sec-
tion, 590-596 feet above
base of formation. Blue-
gray, fine-grained, sili-
ceous shale and gray,
crystalline limestone.

Cathedral formation: unit
6 of Bow Lake section,
596-612 feet above base
of formation. Black-gray,
fine-grained limestone.

Cathedral formation (Ross
Lake shale member):
unit 4 of Eiffel Peak sec-
tion, approximately 400
feet above base of forma-
tion. Greenish-gray, fine-
grained siliceous shale.

127

Approxi-
mate
altitude
Feet

6,000

8,000

7,700

8,200

8,200

8,900

SMITHSONIAN MISCELLANEOUS COLLECTIONS

TABLE 5.—Continued

128

Locality Topographic position

G2i ties tars Southwest ridge of Mount
Temple; 0.3 mile north-
east of Sentinel Pass;
Banff Park, Alberta.

SSD scsies .--South ridge of Mount
Whyte; 0.6 mile north of
Plain of Six Glaciers
chalet; Banff Park, AlI-
berta.

SISA) avnels sates South slope of Mount Bos-

worth; 0.8 mile N. 30°
W. of western end of
Sink Lake; Yoho Park,
British Columbia.

SG2 oi ac risiac East slope of south peak of
Mount Odaray; 0.4 mile
southeast of summit of
south peak; Yoho Park,
British Columbia.

Slo eee aans Southeast ridge of south
peak of Mount Odaray;
0.3 mile southeast of sum-
mit; Yoho Park, British
Columbia.

elaine ae Summit of south peak of
Mount Odaray; Yoho
Park, British Columbia.

S8die acess Mount Stephen; 1.5 miles
E. 30° S. of Field sta-
tion; Yoho Park, British
Columbia.

SSOleo. cow acehs Northwest face of Mount
Stephen; 0.6 mile north-
west of summit of moun-
tain, on ridge immediately

Stratigraphic position
and lithology

Cathedral formation (Ross
Lake shale member):
unit 4 of Mount Temple
section, 460-468 feet above
base of formation. Dark
blue-gray, fine-grained
siliceous shale.

Stephen formation: 50-60
feet above base of unit I
of Mount Whyte section.
Dark-gray, finely crystal-
line or fine-grained lime-
stone.

Stephen formation: basal
10 feet of lowermost unit.
Fine-grained, dark-gray,
argillaceous limestone.

Stephen formation: 12 feet
above base of unit I of
Mount Odaray section.
Gray, crystalline lime-
stone lenses in siliceous
shale.

Stephen formation: unit 2
of Mount Odaray sec-
tion; 120-160 feet above
base of formation. Very
dark-gray, fine-grained,
argillaceous, platy lime-
stone.

Stephen formation : 260 feet
above base of unit 3 of
Mount Odaray section;
420 feet above base of
formation. Very dark-
gray, fine-grained lime-
stone.

Stephen formation; Ogy-
gopsis shale at old “fossil
bed.” Position in section
unknown.

Stephen formation: 30-80
feet above base of unit 16
of undivided Cathedral
and Stephen formations

VOL. 116

Approxi-

mate

altitude

Feet
9,700

8,800

7,000

9,000

9,100

9,700

6,800

7,700

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI
TABLE 5.—Continued
Stratigraphic position
Locality Topographic position and lithohlogy
south of upper end of of Fossil Gully section.
Fossil Gully ; Yoho Park, Very dark-gray, fine-
British Columbia. grained, more or less ar-
gillaceous limestone.
Site Sen 058 Northeast slope of north Stephen formation: 15-30
ridge of Park Mountain; feet above base of unit 3
0.4 mile north of summit ; of Park Mountain sec-
Yoho Park, British Co- tion; 143-158 feet above
lumbia. base of formation. Very
dark-gray, fine-grained or
subcrystalline limestone.
S10) econ Same as Siok. Stephen formation: 180-200
feet above base of unit 3
of Park Mountain sec-
tion; 308-328 feet above
base of formation. Very
dark-gray, fine-grained or
subcrystalline limestone.
STO aisteiciects Northeast slope of Park Stephen formation: unit 3
Mountain, near shore of of Park Mountain sec-
Lake McArthur; Yoho tion, more precise horizon
Park, British Columbia. unknown (talus). Very
dark-gray, subcrystalline
limestone.
Sidi 75.5 West slope of ridge between Stephen formation: 60 feet
Mount Field and Mount below top of unit 1 of
Wapta; directly below Burgess Quarry section.
Walcott’s quarry; Yoho Not placed in general sec-
Park, British Columbia. tion. Calcareous and sili-
ceous shale.
Sit Gearon .. West slope of ridge between Stephen formation: prob-
Mount Field and Mount ably at approximate hori-
Wapta; about 0.3 mile zon of Walcott’s quarry.
south of Walcott’s quarry ; Gray, fine-grained, argil-
Yoho Park, British Co- laceous, thin-bedded lime-
lumbia. stone.
Sli cheitterstes West slope of ridge between Stephen formation (Bur-
Mount Field and Mount gess shale member):
Wapta; at Walcott’s basal 20 feet of unit 2

quarry ; Yoho Park, Brit-
ish Columbia.

of Burgess quarry sec-
tion (Walcott’s “phyllo-
pod bed”). Fine-grained,
dark-gray, siliceous shale.

Approxi-

mate

altitude

Feet

8,300

7,500

7,600

7,700

7,700

130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16
TABLE 5.—Continued
Approxi-
mate
Stratigraphic position altitude
Locality Topographic position and lithohlogy Feet
STTGe eater Same as Si1f. Stephen formation (Bur- 7,900
gess shale member):
150-160 feet above base
of unit 2 of Burgess
Quarry section. Fine-
grained, dark-gray, sili-
ceous shale.
Seika sane Southwest ridge of Mount Stephen formation: 100- 10,600
Temple; 0.4 mile south- 120 feet above base of
west of summit of moun- unit 1 of Mount Temple
tain; Banff Park, Alberta. section. Very dark-gray,
fine-grained or subcrys-
talline limestone.
TasLe 6.—Equivalence of U.S.N.M. and author’s localities
U.S.N.M. Author’s
locality locality Topographic position
TAS) = aie aeyataisteene = sae aso ae etyieye S8d Mount Stephen
BEC col atavotayelote ares si cis fares avetanss(aaness Cs5m Mount Bosworth
BEG Mie ceye sieistesesetonns ela eietstars nies taidtourere Web Mount Niblock
BSE Kelsie vine aele Sate eee ete rere P18m Mount Stephen
21 ROH His CIDE CLOT re michtic oo eicoron © Sirf Mount Field
DOTTY Le te eee loarolee onto ne aioe siete s- W3b Mount Whyte
BBS yl arsintetsapcc Gave Chee tehi are Oeeia ates ars We2od Eiffel Peak
BD Ss CULL nach cite ere ral tate wiave, sheveaiw diets Ssa Mount Bosworth
TG) 1a cai sre laysh a Saciala GRO aeamae ral stele cate arate Wsc Mount Bosworth
OTE caches, Vexay ase aiicb hora Mena Ske te eral etaie Sok Mount Stephen
Od) gis, anes ee eas Cio etek ane P22k Mount Schaffer
(OY EAS Cher Aa Ory BETO ID CCS Wi2c Ptarmigan Pass
Te SRO ARG Crees aOR SoraG Silene panes C4m Ross Lake
(0%, } 5019 10} } 70 OE Eko R Ceo OOS TUE C5m Mount Bosworth
OB Way sestese okey sas veot eke iat ols elo atayay creda sone ie ets Ci5m Bow Lake

PAR Dw PALEONTOLOGY
GENERAL STATEMENT

The fossils described and figured in this paper were selected from
approximately 4,000 specimens collected by the writer and several
thousand specimens collected by Walcott and preserved in the United
States National Museum.

Only trilobites are described. A conservative attitude was followed
in naming new species; forms apparently not belonging to described
species but poorly represented were left out of consideration unless
they seemed to possess special stratigraphic significance ; in the latter
case they were described and illustrated but left unnamed.

All the types of the species based on material collected by the writer
were deposited in the United States National Museum collections.

All the specimens figured in the plates show the upper surface of
the test unless otherwise stated.

TERMINOLOGY

The terminology employed in this paper for describing the trilobite
shield is essentially the one proposed by Howell et al. (1947), with
a few modifications. Ross (1948) raised several objections against
the terminology of Howell et al. Some of his suggestions have been
adopted.

Ross suggests modification of the meaning of the term “palpebral
lobe” in order to give a significance to this portion of the cephalic
shield even when the palpebral furrow is obsolete. The writer does
not agree with this view, since the palpebral lobe as defined by Ross
would designate an area bounded by an arbitrary line, whereas the
usual procedure of considering the palpebral lobe bounded by the
palpebral furrow makes it a natural unit. When the palpebral furrow
is obsolete, the palpebral lobe remains undefined from the rest of the
fixed cheek, just as the glabellar area becomes undefined when the
dorsal furrow is obsolete.

The writer agrees with Ross’ suggestion that the old term “rim” is
more appropriate than the term “border”? used by Howell et al., and
will use “rim” in this paper. The term “anterior limb’’ to designate
collectively the brim, marginal furrow and rim is also accepted.

Ross suggests that the glabella should include the occipital ring.

131

132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

This proposal has several obvious advantages ; however, to avoid un-
essential changes in terminology the glabella will still designate the
axial portion of the cephalic shield exclusive of the occipital ring.

The writer agrees with Ross that measurements taken in the longi-
tudinal direction along the midline should be generally designated as
lengths, even if the part being described has greater extension in the
transverse direction, such as the occipital ring. However, in certain
cases it is impossible to follow this suggestion without distorting the
usual meaning of the words beyond a reasonable point. For example,
the “width” and “length” of a furrow or spine obviously signify
measurements respectively taken across and parallel to the furrow
or spine, whatever its direction. It would be impractical to invert the
meaning of the two words when the furrow happens to be perpendicu-
lar to the axis of the animal. Furthermore, some furrows (e. g., the
marginal furrow on the cephalon) describe a semicircle, hence in
some portions are parallel, in others perpendicular to the axis of the
body.

In this paper the term “‘midlength” will indicate the extent of the
rim, brim, or entire frontal limb in the longitudinal direction. The
terms “length,” “long,’ and “short” will also be applied to the oc-
cipital ring, to axial and pleural segments of the thorax, and to the
posterior limbs to indicate longitudinal extent. The words “width,”
“wide,” and “narrow” will always indicate transverse extent of the
same parts. The lateral extent of the brim in the transverse direction
in trilobites (of the Bathyuriscus type) that have no brim in front of
the glabella is designated as “width.”

The glabellar furrows are numbered from one to four beginning
from the front.

It should be noted that the “width of the fixed cheeks” includes
the palpebral lobes.

All descriptions refer to the upper surface of the test unless other-
wise indicated.

CLASSIFICATION OF CAMBRIAN TRILOBITES

The writer (Rasetti, 1948a) recently gave a brief discussion of the
classification of Cambrian trilobites. Study of the material discussed
in the present investigation did not bring out any evidence against
the conclusions reached in the above-cited paper; on the contrary,
several points received valuable confirmation; particularly the sharp
distinction of most of the Lower and Middle Cambrian Opisthoparida
of the Appalachian and Cordilleran provinces into the two super-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 133

families Corynexochoidae (formerly Bathyuriscidea) and Ptycho-
parioidae. Further distinguishing characters for this subdivision are
discussed under the superfamily Corynexochoidae.

The classification here adopted is the following. It must be borne
in mind that it includes only those families that are represented in the
material studied in this work, hence it is not a complete classification
for the trilobites, or even for the Cambrian forms alone.

For superfamily names the uniform ending -oidae was adopted, as
suggested by the editors of the Treatise on Invertebrate Paleontology,
now in preparation.

Order AGNOSTIDA
Family Peronopsidae
Family Agnostidae

Order EODISCIDA
Family Pagetiidae

Order OLENELLIDA
Family Olenellidae

Order OPISTHOPARIDA

Superfamily BuRLINGIOIDAE
Family Burlingiidae

Superfamily CoryNEXOCHOIDAE
Family Zacanthoididae
Family Dolichometopidae
Family Corynexochidae
Family Dorypygidae
Family Ogygopsididae
Family Oryctocephalidae

Superfamily PrycHOPARIOIDAE
(Families undefined)

DESCRIPTIONS OF GENERA AND SPECIES
OF TRILOBITES

Order AGNOSTIDA

Family PERONOPSIDAE Westergird
Genus PERONOPSIS Hawle and Corda, 1847

Genotype: Battus integer Beyrich.

The generic assignment of many Middle Cambrian agnostids from
the Cordilleran province presents a problem that will require extensive
study before it can be satisfactorily solved. The classification of the
agnostid genera was recently discussed by Harrington (1938), Kobay-
ashi (1939), and Westergird (1946) and was based chiefly on the

134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Atlantic province forms. All these authors agree in assigning most of
the agnostid genera to either of two groups, the Peronopsidae and the
Agnostidae (considered by Westergard as subfamilies), but were
unable to find objective criteria for determining whether a given
agnostid belongs to the Peronopsidae or the Agnostidae. The two
groups are assumed to represent distinct lineages of descent undergo-
ing parallel development in most of the observable characters. Hence,
one could decide to which family an agnostid belongs only through the
study of its phylogeny.

So little is known of the agnostids of the Cordilleran province that
the above-mentioned procedure is at present impracticable, hence one
must resort to morphologic criteria to decide the generic assignment.
Two forms discussed in this work are assigned to Peronopsis because
they closely agree with the genotype and other Atlantic province species
attributed to the genus, but an extensive study like that carried out by
Westergard for the Scandinavian agnostids might show that we have
a case of parallel development in a different stock.

PERONOPSIS MONTIS (Matthew)
Plate 25, figures II-14

Agnostus montis MATTHEW, Trans. Roy. Soc. Canada, ser. 2, vol. 5, sect. 4, p. 43,
pl. 1, fig. 6, 1899.

Agnostus montis Matthew, Watcort, Canadian Alpine Journ., vol. 1, No. 2,
pl. 3, fig. 7, 1908.

Peronopsis montis (Matthew), Kopayasuyi, Journ. Fac. Sci. Imp. Univ. Tokyo,
sect. 2, vol. 5, p. I15, 1930.

This species is rare both at the type locality on Mount Stephen and
in the Burgess shale, where only half a dozen specimens were found
among the several hundred agnostids from the locality preserved in
the United States National Museum.

Horizon and locality.—Stephen formation (Ogygopsis and Burgess
shale lentils ; Bathyuriscus-Elrathina zone). Type locality, U.S.N.M.
14s, Mount Stephen. Also locality U.S.N.M. 35k, Burgess Quarry,
Mount Field.

Types.—Holotype: Royal Ontario Museum. Plesiotypes: U.S.N.M.
Nos. 116215-6.

PERONOPSIS COLUMBIENSIS Rasetti, new species

Plate 33, figures 1-8

Known from numerous cephala and pygidia and several complete
shields, all preserved in limestone.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 135

Both shields considerably convex and well rounded. Glabella on
the average very slightly tapered, highly elevated posteriorly and low
anteriorly, well defined by the dorsal furrow; occupying somewhat
more than two-thirds of the cephalic length. Anterior glabellar lobe
well rounded, with just a suggestion of a pointed shape; transverse
furrow straight. Posterior lobe culminating in an elongated elevation
at one-third the distance from the posterior margin. Basal lobes
triangular, defined by rather shallow furrows. The basal lobes are
united mesially through a very thin occipital ring. Cheeks strongly
convex, without trace of a preglabellar furrow. Marginal furrow
narrow and deep; rim convex, narrow, of even width all around the
cephalon.

The thoracic segments do not show any specific features.

Pygidium with an elevated rachis defined by a well-impressed dor-
sal furrow. Rachis rapidly tapered from the anterior end to the first
transverse furrow, slightly expanded from this level to a little back
of the second transverse furrow, then tapering in triangular shape to
a rather sharp point. Rachis elevated anteriorly and low posteriorly,
terminating at a distance from the posterior marginal furrow some-
what greater than the width of the rim. First transverse furrow
represented by a pair of lateral impressions directed inward and
somewhat forward; second transverse furrow almost indistinct. An
elevated tubercle just in front of the second furrow. Marginal furrow
and rim considerably wider than in cephalon. Rim of almost even
width, with a pair of small, short marginal spines.

Surface of test smooth. Length of largest entire shield 10 mm.
Separated shields indicate the presence of somewhat larger individuals.

This species is very close to Peronopsis interstricta (White) from
the Wheeler formation of Utah, differing only in minor details in the
proportions of the various parts, chiefly the somewhat shorter pygidial
rachis.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality S6k, Mount Odaray. Also localities S6l, Mount
Odaray ; Sok, Mount Stephen; Siok, Si1ol, Park Mountain.

Types——Holotype: U.S.N.M. No. 116267. Paratypes: U.S.N.M.
Nos. 116268-9.

Family AGNOSTIDAE Jaekel
Genus TRIPLAGNOSTUS Howell, 1935

Genotype: Agnostus gibbus Linnarsson,

136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

TRIPLAGNOSTUS BURGESSENSIS Rasetti, new species
Plate 25, figures 1-5

Known from a large number of shields more or less flattened in
shale.

Both shields approximately semicircular. Glabella narrow and long,
divided by a straight transverse furrow into anterior and posterior
lobes ; the former slightly more than half as long as the latter. Anter-
ior lobe pointed, narrowly rounded in front. Posterior lobe slightly
tapered forward, showing two pairs of short and shallow furrows at
the sides; with a small, low tubercle somewhat back of its midpoint.
Basal lobes short and wide. Median preglabellar furrow shallow but
distinct in all specimens. Marginal furrow narrow and deep; rim
narrow, convex. Posterior angles sharp but not extended into spines.

Thoracic segments of the shape usual in the genus. The rachis
shows in each segment the large lateral tubercles and, in addition, a
small median tubercle close to the anterior margin in the first segment.

Pygidial rachis with two well-impressed transverse furrows. Rachis
tapered from the anterior end to the first furrow, expanding from
the first furrow to one-third the length of the posterior lobe, hence
tapered, narrowly rounded at the posterior end; occupying on the
average one-third of the width and four-fifths of the length of the
pygidium. Anterior transverse furrow convex forward, posterior
furrow straight. Middle lobe with a tubercle reaching the maximum
elevation near the posterior margin of the lobe. Pleural lobes convex,
confluent behind the rachis. Marginal furrow as in cephalon, rim
slightly wider, without marginal spines. In young individuals the
rachis does not expand as in the adult and there seems to be a median
postrachial furrow.

Surface of test smooth.

Length of largest complete shield 8 mm.

As far as the writer knows, no species referable to Triplagnostus
had previously been described from the Cordilleran province, al-
though the presence of the genus in that area had been reported.
T. burgessensis very closely resembles the genotype from the Acado-
Baltic province, differing chiefly in the proportionately narrower gla-
bella, shallower preglabellar furrow, and lack of spines at the pos-
terior angles of the cephalon and on the second thoracic segment.
These differences are certainly not of generic importance, and the
writer does not consider it proper to remove the species from the
genus on purely geographic grounds.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 137

Horizon and locality—Stephen formation (Burgess shale lentil ;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 35k, Burgess
Quarry, Mount Field.

Types.—Holotype: U.S.N.M. No. 116212. Paratypes: U.S.N.M.
No. 116213.

Order EODISCIDA
Family PAGETIIDAE Kobayashi
Genus PAGETIA Walcott, 1916
Genotype: Pagetia bootes Walcott.

PAGETIA BOOTES Walcott
Plate 25, figures 6-10

Pagetia bootes Watcott, Smithsonian Misc. Coll., vol. 64, No. 5, p. 408, pl. 67,
figs. I, 1a-f, 1916.

Some of the specimens figured are flattened in the usual variety of
Burgess shale, others are preserved in a calcareous shale and show
most of the original convexity.

Horizon and locality——Stephen formation (Burgess shale lentil ;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 35k (=author’s
locality S11f) ; Burgess Quarry, Mount Field.

Types.—Syntypes: U.S.N.M. Nos. 62855-61. Plesiotypes: U.S.-
N.M. No. 116214.

PAGETIA cf. P. BOOTES Walcott
Plate 33, figures 9-I1

Specimens of Pagetia are fairly common in the black limestone
that constitutes the uppermost portion of the Stephen formation in
many sections. These fossils easily escape observation because of
their very small size. The writer was unable to establish specific dif-
ferences among the specimens collected ; however, as the material is
not abundant and close comparison with the type of the genus, P.
bootes, is made difficult by the different manner of preservation, the
specific identification is considered uncertain. The figured specimens
preserve their full convexity, and the pygidia show the unbroken
axial spine.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Localities S61, Mount Odaray, and Sok, Fossil Gully, Mount
Stephen.

Figured specimens.—U.S.N.M. Nos. 116270-1.

138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

PAGETIA, species undetermined
Plate 22, figures 13, 14

A species of Pagetia is common in the impure limestones and silt-
stones that immediately overlie the Mount Whyte formation in the
Fossil Gully section. However, the coarse grain of the matrix, the
presence of distortion, and the small size of the fossil, make it difficult
to ascertain its exact features. Hence specific identification is not
attempted.

The cranidium has the usual features found in the typical forms
of the genus. The pygidium has a narrow, highly elevated axis show-
ing four rings and a terminal section, which was extended into the
usual spine, broken off in all available specimens. The pleural lobes
are unfurrowed.

Horizon and locality—Cathedral formation (Albertella (?) zone).
Localities Cgh and Coj, Fossil Gully, Mount Stephen.

Figured specimens.—U.S.N.M. No. 116192.

Order OPISTHOPARIDA

Superfamily BURLINGIOIDAE
Family BURLINGIIDAE Walcott
Genus BURLINGIA Walcott, 1908
Genotype: Burlingia hectori Walcott.
BURLINGIA HECTORI Walcott
Plate 28, figure 1
Burlingia hectori Waxcort, Smithsonian Misc. Coll., vol. 53, No. 2, p. 15, pl. 1,

fig. 8, 1908.

An unretouched photograph of the holotype of this exceedingly
rare species is reproduced.

Horizon and locality.—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 14s, Mount Ste-
phen.

Types.—Lectotype: U.S.N.M. No. 53418. Paratype: U.S.N.M.

No. 53419.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 139

Superfamily CORYNEXOCHOIDAE

Although this group of trilobites reaches its greatest development
in the Appalachian and Cordilleran provinces of North America, the
type genus and family from which the name is derived are based on
forms from the Atlantic province. The writer does not believe in
strict adherence to the priority rule at the family and superfamily
levels, hence the names Bathyuriscidae and Bathyuriscidea, proposed
by R. Richter (1932), might be accepted in place of Dolichometopidae
and Corynexochidea (emended as Corynexochoidae) respectively.
R. and E. Richter (1941) abandoned the name Bathyuriscidea in
favor of Corynexochidea for priority reasons, and the writer, con-
trary to his previous usage (Rasetti, 1948a, 1948b), will follow the
same procedure.

It must be emphasized that the limits assigned by the writer to the
superfamily Corynexochoidae are somewhat different from those de-
fined by Richter (1932) and Kobayashi (1935). The writer would at
least provisionally exclude from the superfamily all the post-Cambrian
families, whose affinity with the Early and Medial Cambrian Cory-
nexochoidae is doubtful because the lineages cannot be followed
through the late Cambrian. On the other hand, it seems impossible to
separate from the Corynexochoidae the family Zacanthoididae, placed
by Richter in a superfamily Zacanthoididea and by Kobayashi in a
suborder Mesonacida. New forms described in the present work
make this separation even more arbitrary, as an almost continuous
series of genera bridging the gap between Zacanthoides and Bathyuris-
cus is now known. Even a family distinction between the two groups
is difficult, and is maintained more in order to avoid the family
Dolichometopidae attaining an unwieldy size than for any other rea-
son. The only difference between the classification adopted in 1948
and the present one is purely nomenclatural, consisting in the adop-
tion of the superfamily name Corynexochoidae instead of Bathyuris-
cidea, and the family name Dolichometopidae (originally subfamily
Dolichometopinae Walcott of the family Corynexochidae) instead of
Bathyuriscidea.

The superfamily Corynexochoidae is here construed to include the
families Corynexochidae, Dolichometopidae, Ogygopsididae, Dory-
pygidae, Zacanthoididae and Oryctocephalidae. This list includes only
the families known to have North American representatives in the
Medial Cambrian.

The Corynexochoidae are primarily distinguished by the cephalic
features, chiefly the long, parallel-sided or expanding glabella occupy-

I40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ing, with the occipital ring, all or most of the cephalic length. The
glabella often shows four pairs of well-impressed furrows. The pal-
pebral lobes are usually of medium to large size, often curving in semi-
circular form and coming close to the glabella at both ends. One
character that appears common to all the Corynexochoidae, probably
the most fundamental that distinguishes this group from the Ptycho-
parioidae, is the type of ventral cephalic sutures. In several genera of
the Corynexochoidae (Bathyuriscus, Kootenia, Ogygopsis, Olenoides,
Oryctocephalus, etc.) study of shields exposed from the ventral side
shows that the hypostoma is directly attached to the anterior margin
of the cranidium without an intervening rostrum; the hypostomas of
all Corynexochoidae where this part of the shield is known present
the same type of structure with strong anterior wings, showing the
same type of attachment. Hence, we may safely conclude that the
cephalon of the Corynexochoidae consists of four pieces, the cranid-
ium, hypostoma, and free cheeks. The doublures of the free cheeks
are widely separated by the hypostoma. In Ptychoparia, according to
Barrande (see also Stubblefield, 1936) there is a rostrum between
the anterior margin of the cranidium and the hypostoma. Although
little is known by direct observation of the ventral cephalic sutures in
other Ptychoparioidae, it is fairly safe to assume that the same type of
sutures as in Ptychoparia is of general occurrence. This conclusion
is suggested by the considerable distance between the anterior end of
the glabella and the anterior margin of the cephalon. The hypo-
stoma in the Ptychoparioidae consequently has a much more posterior
position than in the Corynexochoidae, and since the known hyposto-
mas of the former group possess very small anterior wings, they could
not have been directly attached to the cranidium. The conclusion that
Corynexochoidae and Ptychoparioidae can be separated on the basis
of such a fundamental character as the number of pieces constituting
the cephalic shield is important for the classification of the Opistho-
parida.

Family ZACANTHOIDIDAE Swinnerton
Genus ZACANTHOIDES Walcott, 1888
Genotype: Olenoides spinosus Walcott.
ZACANTHOIDES, species undetermined
Plate 21, figures 5-8

Known from numerous cranidia and an articulated shield, all poorly
preserved in siltstone or impure limestone.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI I4I

Glabella parallel-sided, with three distinct pairs of glabellar furrows
and the first pair almost obsolete. Occipital ring with a small spine.
Frontal limb slightly concave, not differentiated into rim and brim;
midlength about one-fourth the glabellar length. Anterior facial su-
tures widely divergent ; hence the limb extends widely at the sides.
Palpebral lobes typical of the genus, about three-fourths the glabellar
length; maximum width of fixed cheeks sligthly less than glabellar
width. Posterior limbs very slender, expanded distally, with a small
intergenal spine at the end.

Pygidium of the general shape characteristic of the genus, but
showing an unusual degree of reduction of the marginal spines; the
first pair is reduced to extremely small spines, the other pairs are
absent.

Length of largest cranidium 16 mm.

This form is almost identical with several described species as far
as the cranidial features are concerned, but differs from all in the
reduction of the pygidial spines. The material is considered too poorly
preserved to warrant naming the species, especially since the chief
diagnostic features would have to be based on the pygidium of which
only one poorly preserved specimen is known.

Horizon and locality——Cathedral formation (Albertella(?) zone).
Locality Cgh, Fossil Gully, Mount Stephen.

Figured specimens—U.S.N.M. No. 116180.

ZACANTHOIDES SEXDENTATUS Rasetti, new species
Plate 22, figures 22-27

Known from numerous cranidia and pygidia preserved in lime-
stone but considerably distorted.

Glabella fairly convex in both directions, slightly expanded for-
ward, rounded in front. Four pairs of glabellar furrows visible, the
posterior pair fairly deep. Occipital furrow well impressed ; occipital
ring expanded mesially but not extended into a spine. Brim de-
veloped at the sides, but absent mesially or at least undifferentiated
from the rim. Brim attaining considerable width at the sides owing to
the outward course of the anterior branch of the facial suture. Fixed
cheeks slightly convex, on the average horizontal. Palpebral lobes
long, with curvature increasing from the anterior to the posterior end;
the maximum width of the fixed cheeks is attained almost at the level
of the posterior end of the glabella and equals three-fourths the
glabellar width. The posterior end of the palpebral lobe forms a rather
sharp angle. Posterior limbs not preserved, obviously very slender.

142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Anterior branch of facial suture starting extremely close to the dorsal
furrow at the anterior end of the palpebral lobe, directed almost
straight outward and forward to the anterior angle of the cranidium
which is rather sharp.

Pygidium twice wider than long, subtriangular, with widely rounded
anterior angles. Axis very prominent, tapered, extended into a short
postaxial ridge that reaches the margin. Four axial rings and a
terminal section visible in most of the specimens. No spines or nodes
on the axis. Pleural platforms convex, downsloping; rim flat, de-
limited by a distinct marginal furrow. Pleura strongly curved, di-
rected backward distally. Three pairs of pleural furrows well
impressed, separated by narrow and shallow interpleural grooves.
Margin extended into six pairs of flat, regularly spaced spines of
decreasing length.

Surface of cranidium smooth. Surface of elevated portions of
pygidial axis and pleural lobes with elevated, scattered granules.

Length of largest cranidium 12 mm. Length of largest pygidium
6 mm., width 12 mm.

This species is sharply distinct from most of the others previously
described in possessing an expanding glabella and a pygidium with
convex pleural platforms and six pairs of spines of equal length. Both
these characters would rather suggest reference to Prozacanthoides.
However, in the writer’s opinion the differential characters of the lat-
ter genus are rather doubtful, and Resser (1939b) himself assigned
to Prozacanthoides several species (e.g., all the forms he described
from the “Ptarmigania strata” of Idaho) that do not comply with his
generic diagnosis, but are rather typical of Zacanthoides. The geno-
type of Prozacanthoides is a Lower Cambrian form, then there seems
to be a gap in the stratigraphic distribution of these trilobites ; later in
the Medial Cambrian forms appear, some of which, like the present
one, are very much like the Lower Cambrian Prozacanthoides, while
others have the characters of the typical Zacanthoides. Awaiting
further study, the writers prefer to assign all the Medial Cambrian
species to Zacanthoides. The described species which most closely
resembles the present form is Z. serratus Resser, which chiefly differs
in the less expanding glabella and flat pleural lobes of the pygidium.

Horizon and locality—Cathedral formation (Albertella(?) zone).
Locality Cgj, Fossil Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116196. Paratypes: U.S.N.M.
No. 116197.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 143

ZACANTHOIDES ROMINGERI Resser
Plate 27, figures 8-10

Embolimus spinosa RoMINGER, Proc. Acad. Nat. Sci. Philadelphia, 1887, p. 15,
pli i; fig: 3.

Zacanthoides spinosus (Walcott) (part), Watcotr, Canadian Alpine Journ.,
vol. 1, No. 2, pl. 4, fig. 1, 1908.

Zacanthoides romingeri REssER, Smithsonian Misc. Coll., vol. 101, No. 15, p. 56,
1942.

The nomenclatural history of this trilobite was given by Resser
in proposing the new name, Z. romingeri. Rominger and Walcott had
independently given the same specific name to two trilobites, Olen-
cides spinosus and Embolimus spinosa. Both species were subse-
quently referred to Zacanthoides and believed to be identical. Resser
observed that the species from Mount Stephen is different from the
typical Zacanthoides spinosus (Walcott) from Nevada, hence had
to make a new name for the former. He also gave a sufficient re-
description of the species.

Horizon and locality.—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s (=au-
thor’s locality S8d) ; “fossil bed” on Mount Stephen.

Types.—Holotype: Univ. Michigan 4871. Plesiotypes; U.S.N.M.
Nos. 102324, 116230.

ZACANTHOIDES SUBMUTICUS Rasetti, new species
Plate 32, figures 11-14

Known from several cranidia and pygidia rather poorly preserved
and somewhat flattened in black, argillaceous limestone.

Glabella parallel-sided, with four pairs of furrows relatively well
impressed. Occipital furrow well marked; occipital ring extended
into a slender spine of moderate length. Structure of the frontal limb
as in Z. romingeri; limb in front of the glabella reduced essentially
to a convex rim, at the sides delimited from the brim by a distinct
marginal furrow. Palpebral lobes as in Z. romingeri. Anterior facial
sutures as in that species, i.e., not quite within the dorsal furrow at
the anterior end of the palpebral lobe, then diverging at an average
angle of less than 45° ; maximum lateral width of the brim about one-
third the glabellar width. Posterior limbs not preserved.

Pygidium subtriangular, one and one-half times wider than long ;
anterior angles well rounded. Axis prominent, moderately tapered,
sloping down steeply to the posterior margin. Three axial rings and
a terminal section defined by well-impressed furrows. No spines or

144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

nodes on the axial rings. Pleural lobes slightly convex, with a shallow
but distinct marginal furrow and a flat rim. Pleural furrows curving
backward but not quite reaching a longitudinal direction; furrows
and grooves shallow. Margin extended into a pair of small spines at
the anterior angles; two other pairs barely distinguishable as slight
denticulations of the margin.

Surface apparently smooth. Length of largest cranidium 19 mm.
Lenth of pygidium 6.5 mm., width 10 mm.

This species closely resembles Z. romingeri, from which it would
be difficult to distinguish on the basis of the cranidial features. How-
ever, the pygidium differs considerably in the great reduction of the
marginal spines. This character also differentiates the species from all
others so far described.

Since cranidia and pygidia of two species of Zacanthoides, here
described as submuticus and longipygus, occur associated at the same
locality, some doubt subsists about the assignment of the two parts
of the shield. The pygidia being much more distinctive than the
cranidia, pygidia are chosen as holotypes of both species; hence,
should the assignment of the two shields prove to be erroneous, the
valid descriptions of the species designated by the two names will be
those referring to the pygidia.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S61, Mount Odaray.

Types.—Holotype: U.S.N.M. No. 116263. Paratypes: U.S.N.M.
No. 116264.

ZACANTHOIDES LONGIPYGUS Rasetti, new species
Plate 32, figures 3-7

Known from numerous cranidia and pygidia preserved in limestone.

Glabella of low convexity, parallel-sided. Four pairs of glabellar
furrows impressed; third and fourth pairs rather deep, fourth pair
showing a tendency to bifurcate. Occipital furrow well impressed
especially at the sides; occipital ring extended into a slender spine
of moderate length. Frontal limb differentiated into a very short
brim and an upturned, convex rim. The brim has a lateral extent
equal to one-half the glabellar width, and near the anterior angles of
the cranidium the marginal furrow is well impressed. Palpebral lobes
as in Z. romingeri. Posterior limbs not preserved. Anterior branch
of facial suture coming close to the dorsal furrow in front of the
palpebral lobe, then diverging at an angle greater than 45° to form
the wide lateral portions of the frontal limb.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 145

Pygidium rather subrectangular than triangular, slightly wider than
long (not including the spines). Axis prominent, well tapered, show-
ing three rings and a terminal section, apparently without nodes or
spines, reaching the posterior margin. Pleural platforms with very
little convexity ; marginal furrow shallow, rim poorly defined. Pleural
furrows turning backward to assume a longitudinal course ; two pairs
of furrows visible, interpleural grooves indistinct. Posterior margin
extended into three pairs of flat, slender spines of regularly decreasing
length, plus two innermost pairs of exceedingly short, barely visible
spines.

Surface of test smooth except for a faint reticulated ornamentation
of the pygidial pleura.

Length of largest cranidium 20 mm. Length of pygidium 5 mm.,
width 6 mm.

As this form was collected from the same locality as Z. submuticus,
some uncertainty exists about the assignment of cranidia and pygidia,
as discussed in the description of Z. submuticus. A pygidium is
chosen as the holotype.

This species closely resembles Z. romingeri from the Ogygopsis
shale and several forms from the Spence shale, Z. idahoensis Walcott,
Z. abbreviatus Resser, and Z. adjunctus Resser. Z. longipygus differs
from all these species in the greater divergence of the facial sutures
and in the characters of the pygidium. Z. romingeri has a proportion-
ately shorter and wider, more triangular pygidium; the other above-
mentioned species have the innermost pairs of pygidial spines more
strongly developed. Zacanthoides spinosus (Walcott) closely re-
sembles the present species as far as the cranidium is concerned, but
a close comparison of the two forms is impossible because the other
parts of the shield are unknown. Similar considerations apply to the
poorly represented species Z. walapai Resser.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S6l, Mount Odaray.

Types—Holotype: U.S.N.M. No. 116259. Paratypes: U.S.N.M.
No. 116260.

ZACANTHOIDES PLANIFRONS Rasetti, new species
Plate 32, figures I, 2

Known from three entire shields rather poorly preserved in lime-
stone.

Glabella on the average approximately parallel-sided, slightly ex-
panded in the anterior portion. Third and fourth pairs of glabellar

146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

furrows well impressed, first and second pairs shallow but distinct.
Occipital ring extended into a slender occipital spine. Frontal limb
relatively long for the genus, flat or very slightly concave as a whole,
not differentiated into brim and rim. Fixed cheeks two-thirds the
glabellar width at their maximum; palpebral lobes about two-thirds
the glabellar length, with curvature increasing posteriorly. Posterior
limbs expanding distally, with intergenal spines. Anterior facial su-
tures diverging at an angle somewhat greater than 45°, straight for
a considerable length, narrowly curving inward before reaching the
anterior margin. Free cheeks with a poorly differentiated rim and flat,
moderately long genal spines.

Thorax of nine segments of the usual shape. Axial rings too poorly
preserved to determine the presence of nodes or spines.

Pygidium subtriangular. Axis elevated, almost reaching the pos-
terior margin, extended into a postaxial ridge, with three rings sepa-
rated by shallow furrows and terminal section. Pleural lobes flat.
Pleura turned backward and extended into five pairs of flat marginal
spines of decreasing size. Spines of first pair almost as long as the
pygidium, second to fifth pairs gradually shorter and narrower.

Surface of shield apparently smooth except for anastomosing ridges
on the ocular platforms.

Length of largest complete shield 50 mm. Separated fragments
indicate the presence of individuals of larger size.

This form differs from all the described species of the genus in the
long, flat, undifferentiated frontal limb. The thorax and pygidium are
almost identical to those of Z. romingeri and several other known
species.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S10, Park Mountain.

Types.—Holotype: U.S.N.M. No. 116257. Paratypes: U.S.N.M.
No. 116258.

ZACANTHOIDES DIVERGENS Rasetti, new species
Plate 32, figures 8-10

Known from an entire shield and several cranidia, all preserved in
limestone.

Glabella moderately convex, somewhat expanded forward, with
three pairs of rather shallow glabellar furrows. Occipital furrow
well impressed especially at the sides; occipital ring short, extended
into an exceedingly short, slender spine. Frontal limb poorly differ-
entiated into rim and brim. Rim upturned, slightly convex, narrow ;

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 147

brim flat. Width of brim at the sides almost two-thirds the glabellar
width. Fixed cheeks flat, slightly upsloping, about two-thirds the
glabellar width. Palpebral lobes defined by a distinct palpebral fur-
row, increasingly curved posteriorly, starting at the level of the an-
terior third of the glabella and ending somewhat behind the level of
the occipital furrow. Posterior limbs not preserved. Free cheeks
downsloping, with a much better-defined rim than the frontal limb ;
genal spines slender, about as long as the glabella. Anterior branch
of facial suture directed almost straight outward to the margin, pro-
ducing sharp anterior angles of the cranidium.

Thorax of nine segments, of the general shape characteristic of the
genus, rather rapidly tapered. Axis too poorly preserved to determine
the presence of nodes or spines.

Pygidium about one and one-half times wider than long, with a
prominent, tapered axis. Three axial rings and a terminal section
can be distinguished. Pleural lobes slightly convex; two pairs of
pleural furrows start outward and turn backward. Marginal furrow
and rim poorly defined ; margin extended into four pairs of flat spines
of decreasing length.

Surface of test smooth, except for strong anastomosing lines on
the ocular platforms, and a few tubercles on the pygidial pleura.

Length of shield 18 mm., of which 7 mm. belong to the cephalon,
7 mm. to the thorax, and 4 mm. to the pygidium.

Only two described species of Zacanthoides, Z. sampsoni Resser,
and Z. serratus Resser, closely resemble the present form, in possess-
ing widely divergent anterior facial sutures. The cranidia of all these
species are difficult to distinguish, but differential characters can
be found in the pygidia. The pygidium of Z. serratus has spines of
uniform length, while the pygidium of Z. sampsoni has spines of de-
creasing size, like the present species, but more strongly developed.
Z. sampsoni also apparently lacks the tubercles on the pygidial pleura.
There is no doubt that the three species are very closely related.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality Stol, Park Mountain. Also locality S6l, Mount
Odaray.

Types—Holotype: U.S.N.M. No. 116261. Paratypes: U.S.N.M.
No. 116262.

Genus ALBERTELLA Walcott, 1908

Genotype: Albertella helena Walcott.
Confusion exists about the species of this genus occurring in the
Ross Lake shale at Ross Lake and Mount Bosworth. Walcott recog-

148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6
nized two species, A. bosworthi and A. helena. Resser admitted four
species, A. bosworthi and three new ones, similaris, rossensis, and
nitida. Study of all the types, additional material in the United States
National Museum, and specimens preserved both in shale and lime-
stone, collected by the writer, shows that there are three clear-cut

species. However, one of them requires a new name because the four

TABLE 7.—Differential characters of three species of Albertella

A. bosworthi
Glabella slightly expanded
toward the front.

Palpebral lobes strongly
curved, about 4 the
glabellar length.

Maximum width of fixed
cheeks over 4, mini-
mum ;5 of glabellar

width.

Thorax with macropleural
development of fourth
segment.

than

Pygidium longer

wide.

Pygidial axis with 5 to 7
distinct rings plus ter-
minal section. Axial
spines or nodes strong.

Pleural lobes slightly con-
cave, horizontal.

A. nitida
Glabella very
tapered.

Palpebral lobes moder-
ately curved, about 4
the glabellar length.

Maximum width of fixed
cheeks 4, minimum 4
of glabellar width.

Thorax with macro-
pleural development of
third segment.

Pygidium somewhat wid-
er than long.

Pygidial axis with 4
rings plus terminal
section. Axial nodes
weak, or indistinct.

Pleural lobes
down-sloping.

convex,

A. microps

slightly Glabella slightly expand-

ed toward the front.

Palpebral lobes weakly
curved, about 4 the
glabellar length.

Maximum width of fixed
cheeks over 3, mini-
mum 4 of glabellar
width.

Thorax with macropleu-
ral development of
third segment.
Pygidial width § of
length.

Pygidial axis with 3
rings plus terminal
section. Axial nodes
moderately prominent.

Pleural lobes
down-sloping.

convex,

names used by Resser cover only two species. Two other species are
described from other localities.

To facilitate identification of the three species occurring at Ross
Lake and Mount Bosworth, the chief differences are summarized in

Table 7.

It will be noticed that the species of Albertella vary considerably

in several respects, chiefly the extent of the brim in front of the

glabella, the divergence of the anterior facial sutures, the length of —
the palpebral lobes, the number of segments represented in the py-
gidium. On the other hand, the thorax, whenever known, shows a

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 149

constant number of seven segments. The differences in the cephalic
characters are as important as those which, in other cases, were con-
sidered sufficient for generic distinction. However, when the whole
shield is considered, it clearly appears that all the species assigned
to Albertella are closely related, since intermediate forms bridge the
gaps between the more distant species.

ALBERTELLA BOSWORTHI Walcott
Plate 17, figures I-9

Albertella bosworthi Watcorr (part), Smithsonian Misc. Coll., vol. 53, No. 2,
p. 22, pl. 1, figs. 4-6 (not fig. 7), 1908; vol. 67, No. 2, p. 38, pl. 7, figs. 3-3b
(not figs. 2-2b), 1917.

Albertella similaris RESsER, Smithsonian Misc. Coll., vol. 95, No. 4, p. 2, 1036.

Resser separated his new species similaris from bosworthi stating
that it differs “in its longer pleura, wider pygidium, and striated
free cheeks.’”’ Examination of the material shows that the proportions
of the pygidium vary slightly and continuously due to distortion ; the
surface of the free cheeks presents different aspects according to the
manner of preservation. The character of the “length” (in the term-
inology used here the ‘“‘width’’) of the thoracic pleura, if one examines
only the holotypes of the respective species, at first sight appears to
be a valid one. However, when numerous shields are examined, it
is clear that there is a continuous range of variation in this character,
the holotype of A. bosworthi representing an extreme case of the
form with narrow pleura. The three entire shields illustrated in this
paper show different degrees in width of the pleura. The species is
also variable in other characters, e.g., the number of segments visible
on the pygidial axis, which may vary between five and seven besides
the terminal section, and the distinctness of the axial nodes. The
holotype of A. bosworthi seems to be an individual with a few seg-
ments on the pygidial axis, although this part of the shield is only
partly preserved (in Walcott’s illustration, the pygidial axis was
partly restored from other specimens). However, in other individuals,
narrow thoracic pleura are associated with numerous axial segments
in the pygidium. The conclusion is that the variation in these charac-
ters is continuous and uncorrelated, making it impossible to recognize
two clear-cut species. Hence the name similaris Resser is placed in
synonymy.

Cranidia and pygidia preserved in the limestone interstratified with
the Ross Lake shale show some characters that are not clearly ap-
parent from shields flattened in shale. The glabella slightly expands
forward in the anterior half and has a moderate transverse and longi-

150 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

tudinal convexity, the latter mostly concentrated in the anterior por-
tion. Two posterior pairs of glabellar furrows distinct, other two
pairs very shallow. Occipital ring long, flat, with a very small node
at the posterior margin. Undifferentiated anterior limb somewhat
concave, extending in front of the glabella. Fixed cheeks slightly
convex, on the average horizontal; palpebral furrow deep. Surface
of cranidium finely granulated.

Horizon and locality—Cathedral formation (Ross Lake shale
member ; Albertella zone). Type locality U.S.N.M. 35c, Mount Bos-
worth. Other localities U.S.N.M. 63m, Mount Bosworth; U.S.N.M.
63j, Ross Lake; C3m, Mount Whyte; C4m, Ross Lake; C5m, Mount
Bosworth; C15m, Bow Lake; C2om, Eiffel Peak; C21m, Mount
Temple.

Types.—Lectotype (designated by Resser) : U.S.N.M. No. 53416.
Paratypes: U.S.N.M. Nos. 53413, 53415. Plesiotypes: U.S.N.M.
Nos. 63762-4, I16151-3.

ALBERTELLA DECLIVIS Rasetti, new species
Plate 17, figures 10-15

Known from several cranidia and associated pygidia preserved in
limestone.

Glabella parallel-sided, rather strongly convex transversely and flat
longitudinally for the posterior two-thirds ; steeply sloping downward
in the anterior third. Third and fourth pairs of glabellar furrows
moderately deep, other two pairs indistinct. Occipital furrow well im-
pressed at the sides, shallow mesially ; occipital ring with a small node
near the posterior margin. Undifferentiated frontal limb flat, steeply
inclined ; anterior facial sutures directed straight forward for a short
distance in front of the eyes, then converging, thus producing a nasute
extension of the brim in front of the glabella. Fixed cheeks on the
average horizontal, their maximum width five-sixths of the glabellar
width. Palpebral lobes very long, semicircular, set off by a well-
impressed palpebral furrow. Anterior end of palpebral lobe ex-
tended into a short ocular ridge; posterior end distant from the
glabella about two-fifths the glabellar width. Posterior limbs slender,
of undetermined width.

Pygidium considerably longer than wide. Axis very strongly con-
vex, slightly tapered, showing five rings plus a terminal section; a
strong node or spine on each ring. The axis reaches the posterior
rim. Pleural lobes on the average horizontal, slightly concave in the
transverse direction. Several pleural furrows impressed, the first

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI I5I

four pairs very distinct. Rim convex, fairly well differentiated.
Pleural spines starting at the level of the pygidial midpoint ; appear-
ing as an extension of the second pleural segment. Spines of unknown
length, probably not greatly divergent.

Surface of all the prominent parts of cranidium and pygidium
finely granulose.

Length of largest cranidium 15 mm., width between the palpebral
lobes 16 mm. Length of largest pygidium 11 mm., width 8 mm.

This species most resembles A. bosworthi in all the cranidial fea-
tures, except the structure of the anterior limb. A. bosworthi has
divergent anterior facial sutures and a short, concave rim in front
of the glabella instead of the flat, nasute, downsloping limb of the
present species. This character, unique among the known species of
Albertella, also distinguishes it from A. robsonensis Resser. The py-
gidium is almost indistinguishable from that of A. bosworithi.

Horizon and locality—Cathedral formation (Albertella zone).
Type locality C15m, Bow Lake.

Types.—Holotype: U.S.N.M. No. 116154. Paratype: U.S.N.M.
No. 116155.

ALBERTELLA NITIDA Resser
Plate 18, figures 1-7

Albertella helena Watcott (part), Smithsonian Misc. Coll., vol. 53, No. 2, p. 19,
pl. 2, figs. 7-9 (only), 1908; vol. 67, No. 2, pl. 7, figs. 5, 5a (only), 1917.

Albertella rossensis RESSER, Smithsonian Misc. Coll., vol. 95, No. 4, p. 2, 1936.

Albertella nitida REssER, Smithsonian Misc. Coll., vol. 95, No. 4, p. 2, 1936.

The form from the Ross Lake shale is probably specifically dis-
tinct from A. helena from the Gordon shale. Among the differences
is the size, the largest individuals from the Ross Lake shale being only
half as large as those from Montana.

Resser made two species without clearly stating the differential
characters. He based A. rossensis on a pygidium figured by Walcott,
but assigned to it as paratypes several unfigured entire shields. Study
of all the type material, and of additional specimens in limestone,
shows that although there are two species, Resser’s holotypes both
belong to the same species. The name nitida is maintained because its
holotype is a much better specimen.

This species is now well known from specimens in shale showing
the general proportions, and separated fragments in limestone show-
ing the convexity and surface characters. Owing to the previous con-
fusion, a new complete description is given.

152 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Glabella almost parallel-sided, slightly tapered forward ; moderately
convex transversely and longitudinally, the longitudinal convexity
stronger anteriorly. Four pairs of glabellar furrows usually distin-
guishable, decreasing in depth from the posterior to the anterior pair.
Occipital furrow shallow but impressed throughout. Occipital ring
moderately long, rounded, bearing a small node near the posterior
margin. Frontal limb undifferentiated at the sides, somewhat concave ;
a short rim in front of the glabella. Limb at the sides less than one-
third the glabellar width; anterior facial sutures slightly divergent,
anterior angles of cranidium rather well rounded. Palpebral lobes
moderately curved, set off by a well-impressed palpebral furrow;
about half as long as the glabella; ocular ridges distinct. Maximum
width of fixed cheeks somewhat over one-half the glabellar width;
fixed cheeks moderately convex and on the average downsloping.
Posterior limbs not as wide as the occipital ring, parallel-sided,
rounded at the extremity ; reaching the base of the genal spine.

Free cheeks with wide ocular platforms and a narrow, slightly con-
vex rim. Genal spines slender.

Thorax of seven segments. Axial rings with nodes faint but dis-
tinct in some specimens, indistinct in others. Pleura with a broad,
shallow, oblique furrow. Pleura of the third segment longer than
those of any other segment and with stronger and longer spines. There
is also a special development of the spines of the last segment, but
not as strong as in the third segment.

Pygidium somewhat broader than long. Axis strongly prominent,
tapered, occupying most of the pygidial length; showing four rings
plus a terminal section. Axial nodes very faint or absent. Pleural
lobes weakly convex, downsloping. Three or four pairs of pleural
furrows visible; interpleural grooves barely indicated. Marginal
spines appearing as extensions of the first two pleural segments to-
gether ; long, straight, more or less divergent. Rim poorly defined,
usually indistinct in flattened specimens.

The proportions of an undistorted shield are the following: length
of cephalon 8.5 mm.; length of thorax 11 mm.; length of pygidium
5 mm., width 7 mm. Length of largest shield 36 mm.

Entire surface of shield finely and densely granulated.

Horizon and locality —Cathedral formation (Ross Lake shale mem-
ber ; Albertella zone). Type locality U.S.N.M. 63j, Ross Lake. Other
localities U.S.N.M. 35c, Mount Bosworth; C3m, Mount Whyte ; C4m,
Ross Lake; C5m, Mount Bosworth; C15m, Bow Lake; C2om, Eiffel
Peak.

row?

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 153

Types.—Holotype: U.S.N.M. No. 63766. Paratypes: U.S.N.M.
Nos. 53402, 53404, 53405, 53406, 63764, 63765. Holotype of “Al-
bertella rossensis”: U.S.N.M. No. 53403. Plesiotypes: U.S.N.M. No.
1161506-8.

ALBERTELLA MICROPS Rasetti, new species
Plate 19, figures 1-8

Known from entire shields flattened in shale and separate cranidia
and pygidia in limestone.

Although this is the least common of the three species of Albertella
in the Ross Lake shale, it is surprising that it should have escaped
attention. The United States National Museum collections contain
several specimens.

Glabella strongly convex in both directions, almost parallel-sided.
When flattened, it appears to expand in front because of the greater
convexity of the anterior portion. Glabellar furrows and occipital
furrow about as distinct as in A. nitida. Brim at the sides half the
glabellar width. Anterior facial sutures not divergent. Rim differen-
tiated by gradually curving upward rather than by a definite marginal
furrow, extremely reduced mesially, shorter than in A. nitida, where
in turn it is shorter than in A. bosworthi. Ocular ridges distinct ;
palpebral lobes only one-third the glabellar length, weakly curved,
rather elevated, set off by distinct palpebral furrows. Anterior end of
palpebral lobe distant from glabella one-half the glabellar width;
maximum width of fixed cheeks somewhat greater. Posterior limbs
much less slender than in the two preceding species. Posterior facial
suture directed straight outward; posterior margin of cranidium and
accompanying marginal furrow curving forward. Hence the posterior
limbs are tapered distally.

Thorax of seven segments. Axis prominent, apparently with axial
nodes. Pleura distinct from those of the other species in that the
spines do not start from the posterior margin but near the middle of
each pleuron. Spines of the third segment exceptionally long and
strong.

Pygidium five-thirds wider than long, composed of a lesser number
of segments than in the preceding species. Axis strongly prominent,
tapered, almost reaching the margin ; showing three rings and a termi-
nal section. Axial nodes distinct. Pleural lobes convex, sloping
steeply to a distinct rim. Three pleural furrows rather strong; two
pairs of interpleural grooves narrow and shallow. Spines appearing
as extensions of the first two pleural segments together ; straight and

154 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

very long, at least four times the pygidial length; rather strongly
divergent.

Length of the largest shield 30 mm. However, most of the speci-
mens do not exceed two-thirds of this size.

Surface of cranidium with elevated granules, much less numerous
than in A. nitida. Surface characters of thorax not preserved. Py-
gidia in limestone apparently smooth.

Horizon and locality—Cathedral formation (Ross Lake shale
member; Albertella zone). Type locality U.S.N.M. 63m, Mount
Bosworth. Other localities C3m, Mount Whyte; C4m, Ross Lake;
C5m, Mount Bosworth; C15m, Bow Lake.

Types——Holotype: U.S.N.M. No. 116163. Paratypes: U.S.N.M.
Nos. 116164-6.

ALBERTELLA LIMBATA Rasetti, new species
Plate 18, figures 8-17

Known from numerous cranidia and pygidia preserved in limestone.

Glabella almost parallel-sided, slightly expanded in front, fairly
convex transversely; longitudinal convexity strong only anteriorly.
Three pairs of glabellar furrows rather short but well impressed ;
occipital furrow also well impressed. Occipital ring expanded mesi-
ally, triangular, with a rather indistinct node. Frontal limb flat, almost
horizontal, with an extremely shallow furrow at the sides, a direct
extension of the frontal portion of the dorsal furrow; this may rep-
resent the marginal furrow, indicating that the area in front of the
glabella is the rim. Anterior facial sutures very strongly divergent,
starting a short distance from the dorsal furrows at the anterior end
of the palpebral lobes. Hence the widely rounded anterior angles of
the cranidium flare out for a considerable distance. Fixed cheeks hori-
zontal, their maximum width about three-fourths the glabellar width.
Palpebral lobes about two-thirds the glabellar length, strongly curved ;
palpebral furrows well impressed. Distance from posterior end of
palpebral lobe to dorsal furrow about one-fourth the glabellar width.
Posterior limbs extremely slender, expanded distally, somewhat wider
than the occipital ring.

Pygidium about equally wide and long. Axis strongly convex, ta-
pered, showing five axial rings (with nodes) and a terminal section;
extended into a short postaxial ridge that reaches the margin. Pleural
lobes convex, rather strongly downsloping. Pleural furrows and inter-
pleural grooves together forming wide depressions, separated by sharp,
narrow ridges. Three such pairs are well marked, the successive ones
less distinct. Rim narrow, flat, differentiated by a distinct marginal

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI * I55

furrow. Pleural spines directed straight backward, of unknown
length; their outer outline being a straight extension of the lateral
margin of the pygidium.

Surface of both shields apparently smooth.

Length of the largest cranidium 11 mm.; width and length of the
largest pygidium 10 mm.

This species most resembles A. cimon (Walcott) in the cranidial
features, but possesses a much wider frontal limb. The pygidium of
Walcott’s species is unknown. The pygidium is of the general type
of that of A. bosworthi, differing chiefly in the convexity of the
pleural lobes.

Horizon and locality—Cathedral formation (Albertella zone).
Type locality C15n, Bow Lake.

Types——Holotype: U.S.N.M. No. 116159. Paratypes: U.S.N.M.
No. 116160.

ALBERTELLA STENORHACHIS Rasetti, new species
Plate 18, figures 18-21

Known from several pygidia well preserved in limestone.

Pygidium equally long and wide. Axis moderately tapered, ele-
vated, occupying almost the entire pygidial length and one-third of
the width, extended into a short postaxial ridge that reaches the
posterior margin. Six axial rings and a terminal section defined by
distinct furrows. Axial rings each with a transverse tubercle, broken
off in the available pygidia. Pleural lobes on the average horizontal.
Pleural platforms slightly convex, with at least five pairs of wide
furrows impressed. Interpleural grooves narrow, impressed only
proximally. Marginal furrow wide and shallow, rim slightly convex
and moderately wide. The marginal spines appear as extensions of
the first and second pleura together, and are directed backward and
slightly outward ; total length unknown. Surface of posterior part of
pleural lobes with characteristic wrinkles. Length of largest pygidium
10 mm., width 10 mm.

This form is described, notwithstanding the lack of an associated
cranidium, because the pygidium cannot be confused with those of
described species. It is obviously close to A. bosworthi and A. declivis,
differing from both chiefly in the somewhat shorter axis and the
lesser relative width of the axis with respect to the pleural lobes.

Horizon and locality—Cathedral formation (Albertella zone).
Locality C15n, Bow Lake.

Types——Holotype: U.S.N.M. No. 116161. Paratypes: U.S.N.M.
No. 116162.

156 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Family DOLICHOMETOPIDAE Walcott
Genus ATHABASKIA Raymond, 1928
Genotype: Athabaskia ostheimeri Raymond.
ATHABASKIA, species undetermined
Plate 22, figures I1, 12

Known from one cranidium and one pygidium preserved in lime-
stone and doubtfully assigned to the same species.

The pygidium is the part of the shield that can definitely be as-
signed to Athabaskia and will be described first. Entire pygidium
semicircular, almost twice wider than long, with rounded anterior
angles. Axis prominent, slightly tapered, extended into a postaxial
ridge that reaches the margin. Three axial furrows visible. Pleural
platforms convex, downsloping ; marginal furrow wide, setting off a
slightly upturned rim. Pleural furrows and grooves both well im-
pressed ; four pairs of each are visible, and terminate in the marginal
furrow, each furrow being close to the corresponding groove in the
manner characteristic of Athabaskia.

The associated cranidium very doubtfully assigned to the species has
a long, narrow, slightly expanded, convex glabella. Lateral compres-
sion exaggerates the narrowness of the glabella in this specimen. Four
pairs of glabellar furrows and occipital furrow shallow. Occipital
ring expanded into a thick spine of unknown length. Frontal limb
reduced to a short rim; brim at the sides well developed transversely ;
anterior angles of cranidium very sharp. Fixed cheeks slightly convex.
Palpebral lobes less than half the glabellar length. Anterior branch
of facial suture paralleling the dorsal furrow, posterior branch paral-
leling the posterior margin. Posterior limbs deeply furrowed.

Surface of both shields smooth. Length of cranidium 10 mm.
Width of pygidium 7 mm., length 4 mm.

The pygidium of this species is typical of a group of species of
Athabaskia, especially resembling A. anax (Walcott) from the Spence
shale. The cranidium, if it belongs to the genus, has unusually short
palpebral lobes and strong occipital spine.

Horizon and locality.—Cathedral formation (Albertella(?) zone).
Locality Cgj’, Fossil Gully, Mount Stephen.

Figured specimens.—U.S.N.M. Nos. 116190-I.

ATHABASKIA? PARVA Rasetti, new species
Plate 31, figures II, 12

Known from several cranidia preserved in limestone.
Glabella moderately convex, expanding forward especially in the

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—-RASETTI 157

anterior third, defined by a well-impressed dorsal furrow. Glabellar
furrows very faint. Occipital furrow straight and of uniform depth
throughout ; occipital ring rather long, with a rounded posterior mar-
gin. Brim and rim almost wholly obsolete, except for a very narrow
and short lateral strip in front of the palpebral lobes. Fixed cheeks
slightly convex, on the average horizontal, reaching their maximum
width posteriorly, where they are somewhat more than half as wide
as the glabella at the same level. Palpebral lobes narrow, about two-
thirds the glabellar length, first directed outward and backward, then
curving to assume a straight backward course in their posterior por-
tion. Palpebral furrow narrow but well impressed. Posterior limbs
not exposed in the available specimens.

Surface of test smooth. Length of largest cranidium 4 mm.

This species is tentatively assigned to Athabaskia, since generic
references in this group of trilobites cannot be reasonably certain un-
less the pygidium is known. The cranidial features are intermediate
between those of Athabaskia and Glossopleura. This species closely
resembles a form described by Resser from the Middle Cambrian
of the Grand Canyon as Clavaspidella kanabensis, differing chiefly in
the less-expanded glabella. Resser included Athabaskia in Clavas-
pidella.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S61, Mount Odaray.

Types.—Holotype: U.S.N.M. No. 116253. Paratype: U.S.N.M.
No. 116254.

Genus BATHYURISCUS Meek, 1873
Genotype: Bathyuriscus haydeni Meek.
BATHYURISCUS ADAEUS Walcott
Plate 31, figures 1-6

Bathyuriscus adaeus Watcort, Smithsonian Misc. Coll., vol. 64, No. 5, p. 334,
pl. 47, figs. 3, 3a-c, 1916.

Walcott’s brief description of this exceedingly common species ap-
pears sufficient to characterize it.

The surface of the cephalon is finely granulated. The largest py-
gidium observed has a length of 13 mm. and a width of 24 mm. By
comparison with the proportions of smaller complete shields, it ap-
pears that a full-grown individual had a length of 50 mm.

158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality U.S.N.M. 58j, Mount Stephen (probably close
to or identical with the author’s locality Sok). Other localities S6k,
Mount Odaray; S6l, Mount Odaray; Sok, Mount Stephen; Stok,
Park Mountain; S1ol, Park Mountain.

Types.—Syntypes : U.S.N.M. Nos. 62631-4. Plesiotypes: U.S.N.M.
Nos. 116249-50.

BATHYURISCUS ROTUNDATUS (Rominger)
Plate 28, figures 2, 3

Embolimus rotundata RoMINGER, Proc. Acad. Nat. Sci. Philadelphia, 1887, p. 16,
IG Mpa, 2b Ge

Bathyuriscus howelli Walcott (part), WaAtcorr, Amer. Journ. Sci., ser. 3,
vol. 36, p. 165, 1888.

Bathyuriscus howelli Walcott (part), MatrHew, Trans. Roy. Soc. Canada,
ser. 2, vol. 5, sect. 4, p. 50, 1890.

Bathyuriscus rotundatus (Rominger), Watcort, Smithsonian Misc. Coll., vol.
53, No. 2, p. 41, 1908; Canadian Alpine Journ., vol. 1, No. 2, pl. 4, fig. 2,
1908; Smithsonian Misc. Coll., vol. 64, No. 5, p. 346, pl. 47, figs. 2, 2a-b,
1916.

Horizon and locality.—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s, Mount
Stephen.

Types.—Holotype: Acad. Nat. Sci. Philadelphia. Plesiotypes:
U.S.N.M. Nos. 62629-30; 116232.

BATHYURISCUS? species undetermined
Plate 22, figures 15, 16

Two cranidia referable to Bathyuriscus are figured, notwithstand-
ing the poor state of preservation, because of the unusual interest of
the faunules from the equivalent of the Cathedral formation in the
Fossil Gully section.

Glabella long and narrow, strongly convex, expanded forward.
Posterior glabellar furrows well impressed, other three pairs very
shallow. Occipital ring not preserved. Frontal limb reduced to a
short rim; at the sides represented by the undifferentiated brim and
rim, of width not much greater than the length of the frontal rim.
Anterior angles of cranidium rounded, closely following the outline
of the glabella. Fixed cheeks rather narrow, somewhat convex. Pal-
pebral lobes about half the glabellar length, set off by a distinct
palperal furrow, of about uniform curvature. Posterior limbs slender,

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 159

parallel-sided. Surface of cranidium smooth. Length of largest
cranidium 8 mm.

Although the described specimens do not show any differences of
generic importance from typical species of Bathyuriscus, the generic
identification in trilobites of this group cannot be considered certain
in the absence of the pygidium.

Horizon and locality.—Cathedral formation (Albertella (?) zone).
Locality Coj, Fossil Gully, Mount Stephen.

Figured specimens.—U.S.N.M. No. 116193.

FIELDASPIS Rasetti, new genus

Glabella occupying almost the entire cranidial length, expanded in
front. Dorsal furrow well impressed; four pairs of glabellar furrows
and occipital furrow distinct. Brim and rim poorly differentiated
from each other laterally, almost obsolete in front of the glabella.
Palpebral lobes about half the glabellar length, curved, narrow, close to
the glabella at the anterior end. Posterior limbs wide and slender,
situated at a considerably lower level than the palpebral lobes.

Thorax apparently of nine segments in the genotype. Pleura
curving backward, the last ones enveloping the pygidium.

Pygidium with prominent axis almost reaching the margin. Pleural
lobes more or less strongly bilobate, with weak furrows and grooves,
undefined marginal furrow and wide, flat or concave rim.

Genotype.—Fieldaspis furcata Rasetti, new species.

Stratigraphic range-—Middle Cambrian (Plagiura-Kochaspis zone).

Remarks.—This is another dolichometopid genus whose isolated
cranidia cannot be distinguished generically from Athabaskia, Ptar-
migama, Dolichometopsis, Stephenaspis, and possibly several others.
Fieldaspis differs from each of them in the characteristic shape of the
pygidium.

FIELDASPIS FURCATA Rasetti, new species
Plate 15, figures 1-8

Known from an entire shield and numerous fragments excellently
preserved in limestone.

Glabella moderately convex in both directions, somewhat expanded
in the anterior two-thirds. Posterior glabellar furrows oblique, well
impressed ; three other pairs transverse, shorter and shallower but
distinct. Occipital furrow shallow mesially ; occipital ring expanded
mesially, rounded, with a small node at the posterior margin. Rim

160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL...L1G

almost obsolete in front of the glabella. Limb laterally consisting of
narrow strips, somewhat expanded at the anterior angles. Anterior
outline of cranidium moderately and evenly curved. Palpebral lobes
somewhat more than half the glabellar length; slighty elevated above
the rest of the fixed cheeks, set off by a distinct palpebral furrow,
curved, the curvature increasing posteriorly. Anterior end of palpe-
bral lobe very close to the glabella, posterior end at a distance of
one-third the glabellar width. Posterior end of palpebral lobe at the
level of the occipital furrow. Fixed cheeks horizontal. Posterior
limbs very slender, at a much lower level than the palpebral lobes;
slightly expanded distally, wider than the occipital ring.

Hypostoma of the general bathyuriscid type. Anterior wings, rep-
resenting the fused rostrum, stout, curving backward. Posterior lobe
small, set off by a shallow but uninterrupted furrow. Maculae narrow
and long, oblique, elevated. Rim narrow, expanded into a pair of
blunt, backward-directed spines at the level of the maculae.

Thorax apparently of nine segments; there is the possibility that
one or two segments might be concealed in the known articulated
specimen, but this seems unlikely. The axial rings apparently had no
spines, although this part is not well preserved and does not permit
a definitive statement. Pleura curving backward, the posterior ones
beginning to curve very near the axis, extended into long, flat spines.
Pleural furrows wide and rather deep, sharply delimited anteriorly.
Pleura of the last segment enveloping the pygidium.

Pygidium of equal length and width, subquadrate in shape except
for the deep median notch. Axis occupying half the pygidial length,
prominent, tapered, with two pairs of furrows well impressed only at
the sides ; extended into a postaxial ridge that reaches the margin at
the median notch. Pleural lobes slightly convex proximally and con-
cave distally, strongly bilobate, almost straight laterally. Two pairs
of furrows and one pair of interpleural grooves faintly impressed
proximally.

Surface of the shield generally smooth, except for wrinkles on some
portions of the pleural lobes of the pygidium.

Length of largest cranidium 14 mm., of largest pygidium 11 mm.
The proportions are about the same in the smaller articulated shield.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Type locality W7f, Kicking Horse Mine, Mount Field.
Also localities Wof, Fossil Gully, Mount Stephen, and W2od, Eiffel
Peak.

Types—Holotype: U.S.N.M. No. 116136. Paratypes: U.S.N.M.
No. 116137.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 161

FIELDASPIS BILOBATA Rasetti, new species
Plate 16, figures 1-7

Known from numerous cranidia and pygidia preserved in limestone.

Glabella strongly convex transversely and fairly convex longitudi-
nally, moderately expanded in the anterior half. Four pairs of gla-
bellar furrows well impressed. Occipital furrow deep laterally but
very shallow mesially. Occipital ring extended into a moderately long,
slender spine. Frontal limb almost entirely obsolete in front of the
glabella, developed at the side and differentiated into a flat brim and
an upturned rim. Maximum width of limb at the sides about one-
third the glabellar width. Fixed cheeks slightly convex, on the aver-
age horizontal, with maximum width one-half the glabellar width,
showing an elongated tubercle at the posterior end. Palpebral lobes
delimited by a well-impressed furrow, strongly curved posteriorly.
Posterior limbs slender, parallel-sided, wider than the occipital ring.
Anterior facial sutures more divergent than the dorsal furrow; hence
the brim increases in width toward the front.

Pygidium one and one-half times wider than long. Axis strongly
elevated, moderately tapered, showing three rings and a terminal
section. Axial furrows shallow mesially. A short postaxial ridge
reaches the posterior margin. Pleural lobes widely rounded at the
anterior and posterior angles, extended laterally farther back than on
the midline, thus producing a distinct median notch. Pleural platforms
downsloping ; marginal furrow indistinct, rim wide and concave. Three
or four pairs of pleural furrows impressed proximally, not extending
across the rim. Interpleural grooves distinct for a short distance.

Surface of test smooth.

Length of largest cranidium 14 mm. Length of largest pygidium
Io mm., width 15 mm.

This species differs from the genotype, F. furcata, chiefly in the
somewhat shorter and more strongly curved palpebral lobes, presence
of an occipital spine, parallel-sided instead of expanding posterior
limbs, and much lesser backward extension of the pleural lobes of the
pygidium.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Type locality W2od, Eiffel Peak.

Types.—Holotype: U.S.N.M. No. 116146. Paratypes: U.S.N.M.
No. 116147.

162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

FIELDASPIS cf, F. BILOBATA Rasetti
Plate 16, figures 8, 9

Specimens collected in limestone of the Mount Whyte formation
on Mount Odaray slightly differ from specimens of F. bilobata from
the type locality in the deeper axial furrows on the pygidium and
larger size, some cranidia reaching a length of 24 mm. These differ-
ences do not appear specifically significant.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W6d, Mount Odaray.

Figured spectmens.—U.S.N.M. No. 116148.

FIELDASPIS SUPERBA Rasetti, new species
Plate 16, figures 10-18

Known from numerous cranidia and pygidia and a specimen pre-
serving thorax and pygidium, all preserved in limestone.

Glabella similar to that of F. furcata but more strongly expanded
in the anterior portion. Four pairs of glabellar furrows well im-
pressed. Occipital ring long, extended into a short, blunt spine. Rim
in front of the glabella short and flat. Frontal limb at the sides not
differentiated into brim and rim, wider than in F. furcata owing to
the strong divergence of the anterior facial sutures. Fixed cheeks as
in F. furcata but more definitely upsloping. There is an elongated
turbercle between the dorsal furrow and the posterior end of the
palpebral lobe. Because of this elevation there is a considerable drop
between the level of the fixed cheeks and that of the posterior limbs.
Posterior limbs very slender, slightly expanded distally, considerably
wider than the occipital ring.

Hypostoma very strongly tapered backward, truncated posteriorly.
A shallow furrow separates the anterior portion, representing the
fused rostrum and including the anterior wings, from the main body
of the hypostoma. Posterior furrow well impressed and continuous
across the hypostoma, situated very near the posterior end. Marginal
furrow and rim well developed around the posterior portion. Maculae
well developed. A pair of marginal spines at the level of the maculae.

Thorax of eight-plus segments. The axial rings may possess nodes
or spines. Pleura with a triangular, elongated ridge at their proximal
end as in most of the bathyuriscid trilobites. Pleural furrows well
impressed. Pleura turned backward distally, extending into sharp
spines.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 163

Pygidium more than one and one-half times wider than long. Axis
slightly tapered, not strongly elevated, with four rings and a terminal
section delimited by shallow furrows. A low postaxial ridge reaches
the margin. Pleural lobes giving the entire pygidium a subrectangular
or slightly cordiform shape, with widely rounded anterior angles and
a shallow median notch. Four pairs of broad, shallow pleural furrows
impressed ; interpleural grooves distinct proximally. Marginal furrow
broad and essentially limited to a series of deeper impressions in the
pleural furrows. Pleural platforms slightly convex and downsloping.
Rim wide, flat, poorly defined. A pair of marginal spines, appearing
as extensions of the second and third pleura together, straight, di-
rected outward and backward, slightly upsloping, longer than the
entire pygidium.

Surface smooth except for faint wrinkles on the pleural lobes of
the pygidium.

Length of largest cranidium 26 mm. Length of largest pygidium
21 mm., width 36 mm.

This species differs from the others assigned to the genus in possess-
ing long pygidial spines. This character brings the species close to
Stephenaspis. However, the broad-based spines of this species pre-
sent closer analogy with the rounded lobes of F. furcata than with
the spines of Stephenaspis bispinosa. The cranidium is almost identi-
cal with that of F. bilobata which has a bilobate instead of a spinose
pygidium.

A pygidium almost identical with that of the present species (ex-
cept for the lesser relative width, which may be due to distortion) was
described by Walcott as Crepicephalus celer. Resser united with it a
cranidium from the same locality described by Walcott as Ptychoparia
clusia and made of these two fragments a species of Kochaspis. There
is no doubt that Walcott’s pygidium belongs either to the present
species or to a closely related one, hence the ptychoparid head bearing
the name Ptychoparia clusia is entirely unrelated. In the case of
Fieldaspis superba, although no complete specimen is known, the
intimate association of numerous fragments and the absence of any
other large trilobite from the same beds makes the assignment of
pygidium and cranidium to one species virtually certain.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Locality W7g, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116149. Paratypes: U.S.N.M.
No. r16150.

Genus GLOSSOPLEURA Poulsen, 1927
Genotype: Dolichometopus boccar Walcott.

164 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

GLOSSOPLEURA BOCCAR (Walcott)
Plate 24, figures 1-6

Dolichometopus boccar Wa.cortt, Smithsonian Misc. Coll., vol. 64, No. 5, p. 363,
pl. 53, figs. 1, Ia-f, 1916.

Glossopleura boccar (Walcott), Poutsen, Medd. Gr¢gnland, vol. 70, p. 268, 1927.

Glossopleura bosworthensis REssER, Smithsonian Misc. Coll., vol. 93, No. 5, p. 31,

1935.
Glossopleura nitida RESSER, Smithsonian Misc. Coll., vol. 93, No. 5, p. 31, 1935.
?Glossopleura stephenensis RESSER, Smithsonian Misc. Coll., vol. 93, No. 5, p. 31,

1935.

Resser’s species bosworthensis and nitida were based on small
specimens in limestone, while the individuals that he regarded as
typical of G. boccar are large shields flattened in shale. Shaly and
calcareous layers alternate at the type locality on Mount Bosworth, and
examination of Walcott’s and the writer’s collections shows that the
differences observed by Resser may be entirely attributed to the size
of the individuals and the manner of preservation. Hence Resser’s
names are placed in synonymy. Glossopleura stephenensis is based on
a specifically unidentifiable shield of Glossopleura supposed to have
been collected from the Ogygopsis shale. The writer has never seen
Glossopleura in those beds and suspects that the specimen has been
mislabeled.

Horizon and locality—Stephen formation (Glossopleura zone).
Type locality U.S.N.M. 57g (=author’s locality S5a), Mount Bos-
worth. Also locality U.S.N.M. 57u, Mount Bosworth.

Types.—Syntypes: U.S.N.M. Nos. 62703-8. Plesiotypes: U.S.-
N.M. Nos. 116204-5.

GLOSSOPLEURA TEMPLENSIS Rasetti, new species
Plate 24, figures 14-17

Known from numerous cranidia and pygidia and an entire shield,
all preserved in limestone.

This species closely resembles the genotype, G. boccar, hence in-
stead of giving a complete description, only the specific differences
will be listed. The cranidium differs from that of G. boccar in that
the glabella expands slightly in front of the palpebral lobes, and the
brim is not as completely obliterated as in that species. The glabellar
furrows are faintly impressed, at least in young cranidia, whereas
cranidia of G. boccar of the same size show hardly any trace of fur-
rows. This character also separates the present species from G. mckeer
which occurs approximately at the same horizon.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 165

The pygidium of G. templensis differs from that of G. boccar in
the greater relative length (ratio of width to length about 1.60 in
G. boccar, 1.40 in G. templensis). In the present species the doublure
is narrower, the axis relatively longer, and the axial furrows are al-
most entirely obliterated.

The complete shield has a length of 25 mm., but fragments indicate
the presence of individuals of considerably larger size. The largest
pygidium collected has a length of 16 mm. and a width of 22 mm.

Horizon and locality——Stephen formation (Glossopleura zone).
Locality S21b, Mount Temple.

Types.—Holotype: U.S.N.M. No. 116209. Paratypes: U.S.N.M.
No. 116210.

GLOSSOPLEURA MCKEEI Resser
Plate 24, figures 9-12, 18

Dolichometopus productus Watcotr (part), Smithsonian Misc. Coll., vol. 64,
No. 5, p. 369, pl. 53, figs. 4, 4a, 1916.

Dolichometopus tontoensis Watcotr (part), Smithsonian Misc. Coll., vol. 64,
No. 5; p- 373, pl. 51, figs. 1d’, 1d”, th, 1916.

Glossopleura mckeei RessErR, Smithsonian Misc. Coll., vol. 93, No. 5, p. 33.
1935; in McKee and Resser, Carnegie Inst. Washington Publ. 563, p. 196,
pl. 21, figs. 1-5, 1945.

Specimens well preserved in the limestones of the lower Stephen
formation are referred to this species, described from the Bright
Angel shale of the Grand Canyon area.

The material from the type area includes pygidia in sandstone pre-
serving the convexity, and accurate comparison of the proportions
with the specimens collected by the writer failed to disclose any dif-
ferences of specific value. The comparison of the cranidia is less
significant, first because only flattened cranidia from the type locality
are available, secondly because specific differences in Glossopleura are
usually more apparent in the pygidia than in the cranidia.

Horizon and locality—Stephen formation (Glossopleura zone).
Locality S3b, Mount Whyte.

Types.—Holotype (from the Bright Angel shale of Arizona) : U.S.-
N.M. No. 62714. Plesiotypes: U.S.N.M. No. 116208.

GLOSSOPLEURA STENORHACHIS Rasetti, new species
Plate 24, figures 7, 8

Known from numerous pygidia well preserved in limestone.
Pygidium 1.5-1.6 times wider than long, moderately convex. Axis
tapered, reaching three-fourths of the pygidial length, extended into

166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

a short postaxial ridge that does not reach the margin. Width of axis
at the base one-fourth of the pygidial width. Five or six segments,
defined by shallow furrows, visible on the upper surface of the axis.
Pleural lobes slightly convex, moderately downsloping, showing faint
pleural furrows even on the upper surface. Rim slightly concave, flat
marginally. Width of doublure one-fifth the pygidial width.

Surface smooth. Length of largest pygidium collected 13 mm.,
width 21 mm. The cranidium has not been identified ; it is possibly
difficult to distinguish from that of the associated and more common
species G. mckeet.

This species closely resembles G. perryi Deiss, also known only
from pygidia, but can be distinguished by possessing a relatively nar-
rower, less prominent axis and a narrower doublure.

Horizon and locality—Stephen formation (Glossopleura zone).
Locality S3b, Mount Whyte. Specimens in the United States National
Museum show that this form occurs in the Stephen formation on
Castle Mountain in association with G. boccar.

Types.—Holotype: U.S.N.M. No. 116206. Paratypes: U.S.N.M.
No. 116207.

GLOSSOPLEURA SKOKIENSIS Rasetti, new species
Plate 23, figures 6-10

Known from several cranidia and pygidia preserved in limestone.

Glabella moderately convex, strongly expanded especially in front
of the anterior end of the palpebral lobes. Four pairs of glabellar
furrows faintly impressed, the posterior pair oblique and somewhat
deeper than the others. Occipital furrow impressed throughout ; oc-
cipital ring moderately long, with a suggestion of a small node. Brim
reduced to narrow, poorly defined strips at the sides; no brim or rim
in front of the glabella. Palpebral lobes about two-thirds the glabellar
length, set off by rather well-impressed palpebral furrows; posterior
portion of palpebral lobe curving somewhat inward. Maximum width
of fixed cheeks greater than half the glabellar width. Posterior limbs
not preserved, but evidently very slender.

Pygidium with a strongly prominent, cylindrical axis occupying
more than two-thirds of the length. Six rings and a terminal section
defined by extremely shallow furrows. A postaxial ridge extends to
the posterior margin, which is slightly notched mesially. Pleural plat-
forms faintly furrowed, downsloping. Rim moderately wide, concave.
Anterior outline of pygidium slanting backward, anterior angles
widely rounded.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 167

Surface of both shields showing very fine, wavy lines.

Length of largest cranidium 7 mm. Length of largest pygidium
5.5 mm., width 9 mm.

This species is extremely close to G. walcotti Poulsen from north-
west Greenland, differing in minor characters such as the slightly more
expanded glabella and more rounded anterior angles of the pygidium.
G. expansa Poulsen has a glabella that expands more suddenly in its
anterior portion, and a somewhat different pygidium.

Horizon and locality.—Cathedral formation (Glossopleura zone).
Locality Cr3r, Skoki Valley.

Types——Holotype: U.S.N.M. No. 116200. Paratypes: U.S.N.M.
No. 116201.

GLOSSOPLEURA MERLINENSIS Rasetti, new species
Plate 23, figures 1-5

Known from several cranidia and pygidia preserved in limestone.

Glabella rather strongly expanded especially in front of the anterior
end of the palpebral lobes, with its minimum width back of the middle.
Convexity of glabella moderate in both directions; glabellar furrows
almost indistinct. Occipital furrow visible mesially ; occipital ring with
a small node. Brim at the sides reduced to narrow strips, poorly dif-
ferentiated from the glabella because the dorsal furrow is almost
obsolete in front of the palpebral lobes. No differentiated rim in front
of the glabella. Palpebral lobes about two-thirds the glabellar length,
the anterior portion directed obliquely, the posterior part directed
straight backward; maximum width of the fixed cheeks not more
than half the glabellar width. Palpebral furrow well distinct. Pos-
terior limbs very slender, about as wide as the occipital ring, directed
somewhat backward.

Pygidium with a prominent, slightly tapered axis in which six rings
and a terminal section are defined by extremely shallow furrows.
Axis occupying two-thirds of the pygidial length and extended into
a short postaxial ridge that does not reach the margin. Pleural plat-
forms downsloping, very faintly furrowed. Rim very wide, slightly
concave. Anterior outline of pygidium not very oblique, posterior
outline semicircular, median notch almost indistinct. Anterior angles
rather narrowly rounded.

Surface of cranidium and of prominent areas of pygidium covered
with extremely fine, wavy lines.

Length of the largest cranidium 10 mm. Length of the largest
pygidium 20 mm., width 32 mm.

168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

This species differs from the preceding in that the fixed cheeks
are narrower, the glabellar furrows shallower and the rim occupies
a much greater fraction of the pygidial area. The last-mentioned
character distinguishes it from the species described by Poulsen, and
also from G. mckeei. In the latter species, the dorsal furrow is much
shallower.

Formation and locality.—Cathedral formation (Glossopleura zone).
Locality C13r, Skoki Valley.

Types——Holotype: U.S.N.M. No. 116198. Paratypes: U.S.N.M.
No. 116199.

GLOSSOPLEURA, species undetermined
Plate 24, figure 13

Represented by a single cranidium preserved in limestone.

Dorsal furrow exceedingly shallow, delimiting a uniformly convex
glabella. Brim entirely absent both in front and at the sides of the
glabella. Palpebral lobes set off by an almost indistinct palpebral fur-
row, and showing almost uniform curvature. Occipital furrow faintly
impressed mesially ; occipital ring slightly elevated at the posterior
margin.

The cranidium of this species resembles more closely the smoother
forms from the Stephen formation, such as G. boccar and G. mckeet,
rather than the more strongly furrowed species with which it is associ-
ated. It does not appear worth while naming the species as the py-
gidia usually supply the most significant specific characters in this
genus.

Horizon and locality—Cathedral formation (Glossopleura zone).
Locality C13r, Skoki Valley.

Figured specimen.—U.S.N.M. No. 116211.

Genus KLOTZIELLA Raymond, 1928
Genotype: Bathyuriscus ornatus Walcott.
KLOTZIELLA ORNATA (Walcott)

Plate 28, figures 7, 8

Bathyuriscus ornatus Watcott, Smithsonian Misc. Coll., vol. 53, No. 2, p. 39,
pl. 1, figs. 1-3, 1908; Canadian Alpine Journ., vol. 1, No. 2, pl. 3, fig. 3,
1908; Smithsonian Misc. Coll., vol. 64, No. 5, p. 346, pl. 46, figs. 4, 4a-b,
1916.

Klotziella ornata (Walcott), RAyMonp, Amer. Journ. Sci., ser. 5, vol. 15, No. 88,
p. 310, 1928.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 169

Horizon and locality—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 14s (=author’s lo-
cality S8d), Mount Stephen.

Types.—Syntypes: U.S.N.M. Nos. 53420, 53421, 53423. Plesio-
types: U.S.N.M. No. 116235.

PARKASPIS Rasetti, new genus

Dolichometopid trilobites with pygidium much smaller than the
cephalon.

Glabella long and narrow, moderately expanded forward, reaching
the anterior margin of the cranidium except for a very reduced rim.
Occipital furrow and four pairs of glabellar furrows well impressed.
Brim and rim at the sides of the glabella narrow. Fixed cheeks
slightly convex, about hali the glabellar width. Palpebral lobes nar-
row, curved, somewhat less than half the glabellar length. Posterior
limbs wider than the occipital ring, short, parallel-sided, deeply fur-
rowed. Free cheeks convex, wide, with well-defined rim and strong
genal spines.

Thorax of 11 segments in the genotype and 10 segments in another
species. Axis occupying about one-fourth of the width. Pleura
deeply furrowed, with the usual elongated triangular ridge at their
inner ends; extended into short, backward-directed spines.

Pygidium subtriangular, about twice wider than long. Axis promi-
nent, not tapered, composed of three segments and a terminal section ;
not quite reaching the posterior margin. Pleural lobes flat, without
distinct marginal furrow or rim; with three pairs of pleural furrows,
faint interpleural grooves, and a denticulated margin.

Genotype.—Parkaspis endecamera Rasetti, new species.

Stratigraphic range—Middle Cambrian (Bathyuriscus-Elrathina
zone).

Remarks.—The species on which the genus is based closely re-
sembles Bathyuriscus; in fact, the cranidia of Parkaspis endecamera
and Bathyuriscus adaeus are so much alike that difficulty would have
been encountered in assigning the proper shields to each species, if
complete specimens had not been available. However, there are dif-
ferences of generic value in the thorax and pygidium. Parkaspis has
II or 10 thoracic segments whereas all known species properly as-
signed to Bathyuriscus have 9. The pygidium of Parkaspis is pro-
portionately smaller than in Bathyuriscus, has fewer segments both
in the axis and pleural lobes, indistinct interpleural grooves, a denticu-
lated margin, and lacks a rim.

170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Another genus with which Parkaspis may be closely compared is
Poliella. Species assigned to Poliella have from 8 to 11 thoracic seg-
ments, and pygidia of the same relative size as Parkaspis. However,
the cranidia of Poliella have longer palpebral lobes and less deeply im-
pressed dorsal and glabellar furrows. Furthermore, the triangular
ridges on the thoracic pleura are poorly developed in Poliella. Ptarmi-
gama also resembles Parkaspis in the general shape of cranidium and
pygidium, but has fewer (8) thoracic segments and the anterior angles
of the cranidium are widely rounded.

The name is derived from Park Mountain.

PARKASPIS ENDECAMERA Rasetti, new species
Plate 31, figures 7-10

Known from numerous specimens, including complete shields
slightly flattened in argillaceous limestone.

Glabella slightly tapered in the posterior third, expanding in the
anterior third, narrowest at the level of the third pair of glabellar
furrows. Glabella rather strongly convex in both directions, the longi-
tudinal convexity increasing anteriorly. Dorsal furrow deep. Four
pairs of glabellar furrows: anterior pair deep, short, terminating in
the anterior pit at the dorsal furrow; second and third pairs short
and shallow ; fourth pair longer, deep, oblique. Occipital furrow mod-
erately deep, continuous ; occipital ring apparently possessing a spine
or node, broken off in all available specimens. Rim very short in front
of the glabella, longer at the sides where it is separated from the brim
by a distinct marginal furrow. Anterior angles of cranidium narrowly
rounded. Fixed cheeks somewhat convex and upsloping, about half
the glabellar width at their maximum. Palpebral lobes very narrow,
long, curved, set off by a deep palpebral furrow. Distance from pos-
terior end of palpebral lobe to posterior margin about one-third the
length of palpebral lobe. Posterior limbs wider than the occipital ring,
short, parallel-sided, deeply and broadly furrowed, bearing a short,
very slender intergenal spine at the extremity. Free cheeks fairly
convex; marginal furrow well impressed, rim convex, moderately
wide, genal spines reaching the level of the middle of the thorax.

Thorax of I1 segments. Axis occupying about one-fourth of the
width. The axial rings may have possessed spines or nodes, broken
off in all available shields. Pleura with a triangular, elongated ridge
at their inner ends; broadly furrowed, curving backward near the
end into short, sharp spines.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI L7L

Pygidium twice wider than long. Axis prominent, not tapered,
abruptly terminated, not quite reaching the posterior margin. Three
rings and a terminal section defined by moderately impressed fur-
rows; the axial rings apparently possessed nodes, broken off in the
available specimens. Pleural lobes slightly downsloping. Three pairs
of broad, rather shallow pleural furrows impressed, not reaching the
margin; interpleural grooves distinct but very shallow and narrow.
No distinct marginal furrow and rim. Margin with three pairs of
small spines, and two or three more pairs represented by a faint
waviness of the outline.

Surface of the test covered with exceedingly fine granules.

Length of holotype, the largest complete shield, 52 mm., of which
18 mm. belong to the cephalon, 25 mm. to the thorax, and 9 mm. to
the pygidium.

This species cannot be confused with any described trilobite when
complete shields are available. Separate cranidia, however, may easily
be confused with those of the associated Bathyuriscus adaeus. The
chief distinguishing features are the greater lateral development of
the brim, the separation of brim and rim by a marginal furrow, and
especially the great depth of the first pair of glabellar furrows in
Parkaspis, whereas they are almost obsolete in Bathyuriscus adaeus.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality S1ol, Park Mountain.

Types.—Holotype: U.S.N.M. No. 116251. Paratypes: U.S.N.M.
No. 116252.

PARKASPIS DECAMERA Rasetti, new species
Plate 27, figure 11

Known from a single specimen flattened in shale.

Cephalon poorly preserved on account of the flattening and dis-
placement of the free cheeks. The glabella appears to expand less
than in the genotype, and the glabellar furrows seem to be shallower.

Thorax of 10 segments. Axis proportionately wider than in P. en-
decamera. Pleura with a more distinct geniculation, situated nearer
the proximal than the distal end of each pleuron. Pleura tapered dis-
tally into sharp spines, directed almost straight outward instead of
backward as in the genotype.

Pygidium similar to that of P. endecamera, differing in the more
tapered axis, and two instead of three distinct pleural furrows. Inter-
pleural grooves narrow but distinct.

Length of shield 20 mm.

172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Horizon and locality—Stephen formation (Burgess shale lentil;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 35k, Burgess
Quarry, Mount Field.

Type.—Holotype: U.S.N.M. No. 116231.

Genus POLIELLA Walcott, 1916
Genotype: Bathyuriscus (Poliella) anteros Walcott.
POLIELLA PRIMA (Walcott)
Plate 12, figures 10-13

Bornemannia prima Watcott, Smithsonian Misc. Coll., vol. 53, No. 5, p. 213,
1908. (Nomen nudum.)

Bathyuriscus (Poliella) primus Wa.cotr (part), Smithsonian Misc. Coll., vol.
64, No. 5, p. 352, pl. 46, figs. 6, 6a (only), 1916.

Bornemannia prima (Walcott), Vocpes, Trans. San Diego Soc. Nat. Hist.,

vol. 4, pt. 2, p. 92, 1925.
Poliella prima (Walcott, part), RESSER, Smithsonian Misc. Coll., vol. 93, No. 5,

P. 44, 1935.
?Poliella castlensis RESSER, Smithsonian Misc. Coll., vol. 93, No. 5, p. 44, 1935.

A description of the species as now restricted follows.

Glabella apparently weakly convex, defined by a shallow dorsal
furrow, subparallel-sided, slightly expanded in front. Glabellar fur-
rows very shallow, almost indistinct. Rim flat, developed at the sides
and in front of the glabella. Brim developed only at the sides, slightly
downsloping, delimited by somewhat divergent anterior facial sutures.
Palpebral lobes almost half the glabellar length, very narrow, mod-
erately curved, set off by distinct palpebral furrows. Anterior end
of palpebral lobe almost in contact with the glabella, posterior end
somewhat more distant; maximum width of fixed cheeks slightly less
than half the glabellar width. Posterior limbs very slender, slightly
expanded distally.

Thorax of eight segments. Axis slightly less than one-third the
thoracic width. Axial rings apparently without nodes or spines.
Pleura broadly furrowed, the anterior ones almost straight, the pos-
terior ones curving more and more backward; those of the last seg-
ment partly enveloping the sides of the pygidium. Fulcrum not very
pronounced, situated at one-third the distance from the axis in the
anterior segments and about one-fourth in the posterior segments.

Pygidium somewhat less than twice broader than long. Shape of
entire pygidium ovate ; anterior angles widely rounded, posterior mar-
gin entire. Axis tapered, occupying two-thirds of the length, with

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 173

two distinct rings and a terminal section. Pleura curving backward ;
besides the anterior furrow, one or two shallow, almost indistinct
furrows; interpleural grooves indistinct. Marginal furrow and rim
indistinct.

Surface apparently smooth. Length of largest shield 18 mm., of
which 7.0 mm. belong to the head, 7.4 mm. to the thorax, and 3.6 mm.
to the tail.

Horizon and locality—Mount Whyte formation (Lake Agnes and
Yoho shale lentils; Wenkchemnia-Stephenaspis zone). Type locality
U.S.N.M. 35m, southwest of Lake Louise. Also localities W2b,
Mount Whyte; W3b, Plain of Six Glaciers; Wok, Fossil Gully,
Mount Stephen.

Types.—Lectotype (designated by Resser) : U.S.N.M. No. 62624.
Paratype: U.S.N.M. No. 62623. Holotype of Poliella castlensis:
U.S.N.M. No. 62626. Plesiotypes: U.S.N.M. Nos. 116111-2.

POLIELLA DENTICULATA Rasetti, new species
Plate 12, figures 6-9

Known from several shields and numerous cranidia and pygidia
partially flattened in shale.

Cranidium typical of the genus. Glabella expanded forward in the
anterior half, with shallow furrows. Occipital ring of medium length,
apparently with a small node. Rim distinct at the sides, upturned,
greatly reduced in front of the glabella. Brim developed only at the
sides, consisting of strips about one-eighth the width of the glabella.
Palpebral lobes half the glabellar length, straight in the anterior part,
curving backward posteriorly; set off by a distinct palpebral furrow.
Maximum width of the fixed cheeks somewhat more than half the
glabellar width. Posterior limbs very slender, parallel-sided, equaling
in width the occipital ring.

Thorax of 10 segments. Axial rings possessing nodes or spines,
broken off in all available specimens. Anterior pleura almost straight,
curving backward more and more as one proceeds toward the py-
gidium; terminating in spines, short in the anterior segments and
longer posteriorly ; those of the last pleuron enveloping the sides of
the pygidium. Pleural furrows broad, starting from the anterior
angle of each pleuron and running somewhat obliquely outward and
backward.

Pygidium transversely ovate, twice wider than long. Axis tapered,
with three rings and a terminal section, reaching not far from the
posterior margin. Pleura curving backward; three broad pleural fur-

174 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

rows are impressed, and two narrow, short interpleural grooves are
visible. Anterior angles drawn into a pair of very short spines. Pos-
terior margin wavy, indicating two or three pairs of exceedingly
short spines. Margin with a median notch. Marginal furrow and rim
almost indistinct.

Surface apparently smooth.

Length of largest shield 34 mm., of which 12 mm. belong to the
head, 17 mm. to the thorax, and 5 mm. to the pygidium.

This species is similar to P. prima, from which it differs in possess-
ing 10 instead of 8 thoracic segments, wider fixed cheeks, longer pal-
pebral lobes, and denticulated pygidium. These characters also dis-
tinguish it from other described species.

Horizon and locality —Mount Whyte formation (Yoho shale lentil ;
W enkchemnia-Stephenaspis zone). Type locality Wok, Fossil Gully,
Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116109. Paratypes: U.S.N.M.
No. 116110.

POLIELLA cf. P. DENTICULATA Rasetti
Plate 9, figures 7, 8

Known from numerous cranidia preserved in limestone.

The observable characters do not show any specific differences be-
tween these cranidia and those of P. denticulata as far as the different
manner of preservation allows an accurate comparison. In the absence
of a pygidium, specific identification cannot be reasonably certain.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Locality W16j, North Gully, Mount Stephen.

Figured specimens.—U.S.N.M. No. 116082.

POLIELLA, species undetermined No. 1
Plate 22, figures 17-19

Known from numerous cranidia and one pygidium poorly preserved
in silty limestone.

Glabella slightly expanded in the anterior third, with traces of
glabellar furrows. Occipital furrow shallow, occipital ring slightly ex-
panded mesially but not extended into a spine. Rim greatly reduced,
brim absent in front of the glabella. At the sides the limb has only
one-sixth the width of the glabella and does not widen at the anterior
angles, the facial suture being parallel to the dorsal furrow. Palpebral
lobes narrow, about two-thirds the glabellar length, evenly curved.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 175

Maximum width of fixed cheeks about two-thirds the glabellar width.
Distance from posterior end of palpebral lobe to dorsal furrow one-
half the glabellar width. Posterior limbs slender, slightly expanded
distally, equaling in width the occipital ring.

A pygidium tentatively assigned to the species has a prominent,
slightly tapered axis composed of two rings and a terminal section.
Pleural lobes downsloping ; pleura curved backward, with three pairs
of distinct furrows; marginal furrow and rim indistinct. Shape of
entire pygidium transversely ovate, with a slight median notch in
the posterior outline.

Length of largest cranidium 8 mm. Length of pygidium 3.3 mm.,
width 4.7 mm.

Horizon and locality—Cathedral formation (Albertella (?) zone).
Locality Coh, Fossil Gully, Mount Stephen.

Figured specimens.—U.S.N.M. No. 116194.

POLIELLA, species undetermined No. 2
Plate 22, figures 20, 21

Two poorly preserved pygidia are tentatively assigned to Poliella.
Even the generic identification cannot be certain owing to the poorly
preserved material and the lack of the corresponding cranidium.

The pygidium is transversely suboval, almost twice wider than long.
Anterior angles widely rounded. Axis moderately prominent, showing
three rings and a terminal section, not quite reaching the posterior
margin, which has a shallow median notch. Pleura turned backward ;
three pairs of furrows visible, interpleural grooves indistinct. Pleural
platforms slightly convex, almost horizontal; rim narrow, poorly
defined, slightly concave. There is a pair of exceedingly small spines
at the posterior angles. Surface apparently smooth. Length of py-
gidium 5 mm., width 9 mm.

Horizon and locality Cathedral formation (Albertella (?) zone).
Locality Cgj’, Fossil Gully, Mount Stephen.

Figured specimens —vU.S.N.M. No. 116195.

Genus POLYPLEURASPIS Poulsen, 1927

Genotype: Polypleuraspis solitaria Poulsen.

A species of this genus, described by Poulsen on a single pygidium,
occurs abundantly in certain beds of the Glossopleura zone in the
Canadian Rockies. Complete shields of the new form have been re-
covered. A more complete diagnosis of the genus is based on the new
material.

176 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Entire shield with considerable transverse convexity. Cephalon and
pygidium of equal length, thorax somewhat longer than the other two
parts of the dorsal shield. Glabella long and narrow, expanded for-
ward, reaching the anterior margin of the cranidium. Occipital fur-
row and four pairs of shallow glabellar furrows visible. Brim and
rim obsolete. Palpebral lobes narrow, almost half the glabellar length ;
fixed cheeks about one-third the glabellar width. Posterior limbs
small, narrowly triangular. Free cheeks wide, with a narrow rim and
short, slender genal spines.

Thorax of seven segments. Axis strongly convex. Pleura broadly
furrowed, tapering distally to backward-directed spines.

Pygidium of approximately equal length and width. Axis strongly
prominent, moderately tapered, multisegmented, almost reaching the
posterior margin. Pleural lobes downsloping, with numerous furrows
and a narrow, concave rim.

This genus is obviously closely related to Glossopleura, from which
it differs chiefly in the characters of the pygidium.

POLYPLEURASPIS INSIGNIS Rasetti, new species
Plate 23, figures II-15

Known from large numbers of separate cranidia and pygidia and
a few articulated shields, all excellently preserved in limestone.

Glabella long and narrow, moderately expanded forward, reaching
the anterior margin of the cranidium, moderately convex in either
direction. Glabellar furrows shallow; first pair indistinct, directed
inward and forward; second and third pairs better impressed, trans-
verse; fourth pair directed inward and backward. Occipital furrow
moderately deep; occipital ring rather short and simple. Brim and
rim absent or indistinct both in front and at the sides of the glabella,
as in Glossopleura. Fixed cheeks approximately horizontal, one-third
the glabellar width. Palpebral lobes narrow, somewhat less than half
the glabellar length, demarcated by a distinct palpebral furrow ; an-
terior end of palpebral lobe in contact with the dorsal furrow ; anterior
part of lobe straight, oblique, posterior part curving backward and
approximately parallel to the dorsal furrow. Posterior limbs some-
what narrower than the occipital ring, short, triangular, delimited by
an almost straight posterior branch of the facial suture. Free cheeks
wide, slightly convex, with a narrow rim defined by a well-impressed
marginal furrow and short, slender, backward-directed genal spines.

Thorax of seven segments. Axis as wide as the occipital ring in
its anterior half, slightly tapered posteriorly. There is no evidence for

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI T77

axial spines or nodes. Pleura broadly furrowed, curving backward
and tapering to slender, sharply pointed spines. The pleural lobes
are as wide as the axis.

Pygidium of about equal length and width, subtriangular. Axis
strongly elevated above the pleural lobes, moderately tapered, almost
occupying the whole pygidial length and less than one-third of the
width. About 8 to 10 axial segments are distinguishable, separated by
shallow furrows becoming indistinct posteriorly. Approximately the
same number of furrows distinguishable on the pleural lobes ; furrows
broad, separated by narrow ridges. Pleural lobes steeply downsloping.
Rim narrow and concave; doublure rather narrow, its margin clearly
visible as an impression on the dorsal side of the shield.

Surface of shield smooth.

Length of complete shield 23 mm. Length of largest cranidium
Io mm.; length of largest pygidium 12 mm., width 14 mm.

This species differs from the genotype, P. solitaria Poulsen, known
from a single pygidium, in the greater proportionate length of this
portion of the shield and the more distinct concave rim. Poulsen’s
species also occurs in association with Glossopleura.

Horizon and locality—Stephen formation (Glossopleura zone).
Type locality S2tb, Mount Temple. Also locality S3b, Mount Whyte.

Types.—Holotype: U.S.N.M. No. 116202. Paratypes: U.S.N.M.
No. 116203.

Genus PTARMIGANIA Raymond, 1928
Genotype: Bathyuriscus rossensis Walcott.
PTARMIGANIA ROSSENSIS (Walcott)
Plate 19, figures 9-16

Bathyuriscus rossensis Wa.cott, Smithsonian Misc. Coll., vol. 67, No. 2, p. 46,
pl. 5, figs. 5, 5a-d, 1917.

Bathyuriscus cf. B. rossensis WaAtcott (part), Smithsonian Misc. Coll., vol. 67,
No. 2, p. 49, pl. 5, figs. 6a, 6a’ (only), 1917.

Ptarmigania rossensis (Walcott), RAyMonp, Amer. Journ. Sci., ser. 5, vol. 15,
No. 88, p. 310, 1928.

The species has been adequately described by Walcott, and is based
on specimens in shale. Walcott also described specimens collected
from limestone interstratified with the Ross Lake shale as Bathyuris-
cus cf. rossensis. Resser made a new species, Ptarmigania longula, for
the large cranidium in limestone illustrated by Walcott. Examination
of the type material appears to show that this cranidium is specifically

178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

different from P. rossensis, hence Resser’s species may at least pro-
visionally be considered as valid. However, the numerous other speci-
mens in limestone in the United States National Museum collections,
including the pygidium figured by Walcott, are entirely typical of
P. rossensis, hence the name longula is restricted to the type cranidium.

Horizon and locality.—Cathedral formation (Ross Lake shale mem-
ber ; Albertella zone). Type locality U.S.N.M. 63j, Ross Lake. Other
localities C4m, Ross Lake ; C5m, Mount Bosworth ; C15m, Bow Lake;
U.S.N.M. 35c, 63m, Mount Bosworth.

Types.—Lectotype (hereby designated): U.S.N.M. No. 63733.
Paratypes: U.S.N.M. Nos. 63729-32. Specimen described as “Bathy-
uriscus cf. rossensis’: U.S.N.M. No. 63735. Plesiotype: U.S.N.M.
No. 116167.

PTARMIGANOIDES Rasetti, new genus

Dolichometopid trilobites closely related to Ptarmigania and dis-
tinguishable from the latter genus only by the pygidial features.

Glabella long and narrow, somewhat expanded forward, reaching
the anterior margin of the cephalon except for an exceedingly short
rim. Glabellar furrows generally well developed; four pairs visible
in some of the species. Occipital ring usually extended into a spine.
Brim and rim at the sides of the glabella not sharply differentiated
from each other. Fixed cheeks approximately horizontal, somewhat
convex ; maximum width equaling half the glabellar width. Palpebral
lobes curved, narrow, long, set off by a deep palpebral furrow. Pos-
terior limbs slender.

Thorax probably of seven segments. Axis strongly convex.

Pygidium large, with a very prominent axis occupying most of the
length. Pleural platforms downsloping; marginal furrow shallow,
often reduced to a series of pits. Margin extended into at least four
pairs of strong, cylindrical spines. A strong upright spine on the first
axial ring.

Genotype.—Ptarmuganoides bowensis Rasetti, new species.

Stratigraphic range-——Middle Cambrian (Albertella zone).

Remarks.—Poulsen (1927) based his genus Dolichometopsis on the
genotype, D. resseri, and other species from the Lower Cambrian
Cape Kent formation of northwest Greenland. Poulsen tentatively
assigned to the genotype a small, nonspinose pygidium. Later Resser
(1939b) described a series of species from the Albertella zone (“Ptar-
mugania strata”) of Idaho. These species present cranidia, as Resser
himself states, generically indistinguishable from Ptarmigania associ-
ated with strongly spinose, Kootenia-like pygidia. Resser concluded

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 179

that Poulsen had probably assigned a wrong pygidium to Dolicho-
metopsis and that the latter genus is characterized by spinose pygidia,
hence assigned all the species in question to Dolichometopsis and
modified the diagnosis of the genus accordingly.

Resser’s procedure has already been questioned by Kobayashi
(1942) and the writer (Rasetti, 1948a). The writer does not doubt
that Resser’s assignment of the spinose pygidia to his cranidia is
correct, but sees no reason to assume that the original species of
Dolichometopsis from the Lower Cambrian of Greenland possess
similar pygidia. It is a common occurrence among dolichometopid
trilobites to find almost identical cranidia associated with a great va-
riety of pygidia, and in such cases generic characters must be based
on the features of the entire shield and not of the cranidium alone.
Furthermore, the great difference in age between the two groups of
species favors placing them in different genera. Hence the writer pro-
poses the new name Ptarmiganoides for the forms with a Ptarmi-
gania-like cephalon and a Kootenia-like pygidium that Resser as-
signed to Dolichometopsis. As genotype the writer chooses a new
species from the Albertella zone of the Canadian Rockies, as for this
form, notwithstanding the lack of complete specimens, the association
of the fragments of the shield leaves no doubt as to the assignment
of cranidium and pygidium to one species.

Ptarmiganoides, as the name indicates, is undoubtedly a close rela-
tive of the contemporaneous Ptarmigania from which it differs in the
structure of the pygidium.

Besides the genotype, the following species described by Resser
should be included in Ptarmiganoides: Dolichometopsis alia, D.
comis, D. communis, D. gravis, D. gregalis, D. lepida, D. mansfieldi,
D. media, D. potens, D. poulsem, D. propinqua, D. stella. Possibly
some of these species are synonymous, but further study and additional
material would be required to discuss this question properly.

PTARMIGANOIDES BOWENSIS Rasetti, new species
Plate 20, figures 1-9

Known from numerous cranidia, free cheeks, and pygidia excel-
lently preserved in limestone.

Glabella long, slightly expanded forward, well rounded in front,
showing considerable convexity in both directions, delimited by a deep
dorsal furrow. First pair of glabellar furrows faint but visible, second
pair indistinct, third pair about as deep as the first, fourth pair longer
and deeper than the others, oblique. Occipital furrow well impressed
across the glabella; occipital ring extended into a short, blunt spine.

180 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Rim in front of the glabella exceedingly short; rim and brim at the
sides fairly extended, poorly differentiated by a shallow marginal
furrow. Anterior angles of cranidium well rounded. Fixed cheeks
somewhat upsloping from the dorsal furrow, slightly convex, about
half the glabellar width. Palpebral furrow very deep; palpebral lobes
narrow, straight in the anterior portion, regularly curved posteriorly,
slightly over half the glabellar length. Posterior limbs very slender,
not entirely preserved in any specimen. Anterior facial sutures some-
what divergent in front of the eyes, turning inward with a wide curve
after crossing the marginal furrow; posterior branch directed straight
outward. Free cheeks with convex ocular platforms, a wide, flat rim,
and genal spines of unknown length.

Pygidium with approximately semicircular outline. Axis strongly
prominent, subcylindrical, rounded posteriorly, almost reaching the
posterior margin, composed of four segments and a terminal section.
There is an upright spine of unknown length on the first segment and
a small node on the second. Pleural platforms convex and down-
sloping, showing three distinct furrows and rather indistinct grooves.
Marginal furrow wide and fairly well impressed, deepening into four
pairs of broad pits in correspondence with the pleural furrows. Rim
convex, extended into six pairs of spines. Spines of the first four pairs
about equally strong, approximately half the length of the pygidium.
Spines of the fifth pair much thinner and shorter; spines of the sixth
pair very short and blunt.

Surface of cranidium with granulations particularly marked on
the posterior portion of the glabella and the fixed cheeks. Surface of
pygidium with irregular, elevated, wavy lines.

Length of the largest cranidium 20 mm.; length of the largest
complete pygidium (inclusive of the spines) I0 mm., width 18 mm.
Fragments indicate the presence of individuals of larger sizes.

This species differs from any of those described by Resser under
the genus Dolichometopsis in possessing six instead of four marginal
pygidial spines.

Horizon and locality—Cathedral formation (Albertella zone).
Locality C15n, Bow Lake.

Types—Holotype: U.S.N.M. No. 116172. Paratypes: U.S.N.M.
No. 116173.

STEPHENASPIS Rasetti, new genus

Glabella occupying almost the entire cranidial length, moderately
expanded forward, well delimited by the dorsal furrow. Four pairs
of distinct glabellar furrows. Brim and rim developed at the sides;

NOI 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 181

only an exceedingly short rim in front of the glabella. Fixed cheeks
less than half the glabellar width. Palpebral lobes about half the
glabellar length, curved, close to the glabella at both ends. Posterior
limbs slender, parallel-sided.

Thorax of nine segments. Axis occupying less than one-third of the
width. Pleura curved backward and extended into long, cylindrical
spines. Each pleuron carries an elongated triangular ridge at its inner
end and is broadly furrowed. The pleura of the last segment envelop
the pygidium.

Pygidium subrectangular. Axis prominent, almost reaching the
posterior margin, with several distinct segments. Pleura flat, with fur-
row and grooves curving backward; straight sides extended into a
pair of spines; rim poorly defined.

Genotype.—Stephenaspis bispinosa Rasetti, new species.

Stratigraphic range.—Middle Cambrian (Wenkchemnia-Stephen-
aspis zone).

Remarks.—The genotype and only known species has many char-
acters that place it in an intermediate position between Zacanthoides,
Albertella on one side and Bathyuriscus, Poliella, Fieldaspis and re-
lated genera on the other. However, when the combination of all
characters is considered, it appears that this form cannot be assigned
to any of the above-mentioned genera.

Stephenaspis differs from Zacanthoides chiefly in the reduction of
the frontal limb and different pygidium. Albertella has a somewhat
similar pygidium, but a thorax with seven segments and characteristic
macropleural development of certain segments. The cephalic charac-
ters do not greatly differ from those of Poliella and Ptarmigania, but
the thorax, with its long spines, has a different aspect, and the pygidia
differ considerably. Probably the most closely related genus is Field-
aspis, which differs chiefly in the greater distance of the posterior end
of the palpebral lobe from the dorsal furrow and the shape of the
pygidium.

STEPHENASPIS BISPINOSA Rasetti, new species
Plate 10, figures 1-6

Known from a large number of entire shields partially flattened in
shale.

Glabella prominent, expanded forward in the anterior half, de-
limited by a deep dorsal furrow. Posterior glabellar furrows fairly
deep, oblique, not quite meeting mesially. Next pair shorter and shal-
lower, transverse. Second pair impressed like the preceding, directed
slightly forward. First pair shortest, shallow. Occipital furrow

182 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

straight, well impressed ; occipital ring long, extended into a slender
occipital spine. Anterior outline of cranidium fairly straight. Brim de-
veloped only at the sides ; rim short, upturned, greatly reduced in front
of the glabella. Anterior facial sutures rather divergent; anterior
angles of cranidium narrowly rounded. Palpebral lobes half the
glabellar length, narrow, set off by a distinct palpebral furrow. An-
terior end of palpebral lobe almost in contact with the dorsal furrow;
palpebral lobe straight in its anterior portion, curving posteriorly.
Posterior limbs slender, parallel-sided, about as wide as the occipital
ring, without intergenal spines. Free cheeks wide, with a distinct rim
and genal spines of moderate length.

Thorax of nine segments with a strongly prominent axis. Each
ring carried a spine, which is broken off in all the available specimens.
Pleura first directed straight transversely, then rather suddenly curv-
ing backward. Pleural spines increasing in length from the first to
the last segment. The spines of the last segment are almost parallel
to the sides of the pygidium.

Pygidium 14 times wider than long. Axis slightly tapered, strongly
prominent, showing three or four rings and a terminal section, with-
out axial spines ; extended into a short postaxial ridge that reaches the
margin. Pleura flat, bent backward as the thoracic pleura. First
pleuron curving straight backward and extended into a pair of straight
spines about two-thirds the pygidial length. Three pairs of rather
shallow pleural furrows and interpleural grooves, ending in a shallow,
wide marginal furrow. Rim flat, poorly defined. Posterior outline
of pygidium with distinct mesial notch.

Free cheeks with transverse, anastomosing ridges. Rest of surface
apparently smooth.

Length of adult shield 45 mm., of which 14 mm. belong to the
head, 20 mm. to the thorax, and 11 mm. to the tail.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
W enkchemmia-Stephenaspis zone). Type locality Wok, Fossil Gully,
Mount Stephen. Other locality W16k, North Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116091. Paratypes: U.S.N.M.
No. 116092.

STEPHENASPIS cf. S. BISPINOSA Rasetti
Plate 10, figures 7-9

Known from several pygidia well preserved in limestone.

The available material does not show definite differences between
this form and the typical S. bispinosa from the Yoho shale, apart from
those attributable to the different manner of preservation. Since the

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 183

cranidium is not known, definite identification is not indicated. The
pygidium has almost horizontal pleural platforms and an upturned
flat rim, making the pleural lobes slightly concave as a whole. The
flattening of the shields in the Yoho shale makes it difficult to as-
certain whether the pygidium of S. bispinosa had the same relief.
The axis may be slightly longer in the present form.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Locality W16j, North Gully, Mount Stephen.

Figured specimens—U.S.N.M. No. 116093.

Genus VANUXEMELLA Walcott, 1916
Genotype: Vanuxemella contracta Walcott.
VANUXEMELLA NORTIA Walcott
Plate 20, figures 10-13

Vanuxemella nortia Watcott, Smithsonian Misc. Coll., vol. 64, No. 3, p. 222,
pl. 36, fig. 5, 1916; vol. 67, No. 2, p. 37, pl. 7, fig. 7, 1917.

Vistoia prisca Watcott, Smithsonian Misc. Coll., vol. 75, No. 3, p. 122, pl. 17,
fig. 14, 1925.

Walcott’s generic description illustrates most of the characters of
this species. The writer collected and illustrates here excellently pre-
served cranidia and pygidia in limestone, showing certain details better
than the shields flattened in shale.

The pygidium has the two anterior pairs of pleura terminating in
short, faleate ends beyond the general outline of the shield. There
is a pair of small, sharp spines at the posterolateral angles. Axial
and pleural furrows are extremely shallow on the outer surface, but
quite distinct on the interior cast.

Horizon and locality—Cathedral formation (Ross Lake shale
member ; Albertella zone). Type locality U.S.N.M. 35c, Mount Bos-
worth. Also localities C5m, Mount Bosworth; C4m, Ross Lake;
C2om, Eiffel Peak; C15m, Bow Lake; C21m, Mount Temple; U.S.-
N.M. 63j, Ross Lake; U.S.N.M. 63w, Bow Lake.

Types.—Holotype and paratypes: U.S.N.M. No. 61728. Plesio-
types: U.S.N.M. Nos. 116174-5.

WENKCHEMNIA Rasetti, new genus

Glabella expanded forward in the anterior part, defined by a shal-
low dorsal furrow; glabellar furrows shallow. Frontal limb poorly
differentiated into rim and brim, extremely reduced in front of the
glabella. Palpebral lobes narrow, oblique, about one-third the glabel-

184 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

lar length ; their posterior end distant from the dorsal furrow by more
than half the glabellar width. Posterior limbs wider than the occipital
ring, slightly tapered, rounded distally ; distance from posterior end
of palpebral lobe to posterior margin equal to the length of palpebral
lobe.

Thorax of 9 segments. Pleura straight, broadly furrowed, not ex-
tended into long spines.

Pygidium twice wider than long; subtriangular. Axis prominent,
furrowed, occupying most of the length. Pleural furrows almost
straight, wide; interpleural grooves almost indistinct. Marginal fur-
row and rim narrow, poorly defined.

Genotype.—W enkchemnia walcotti Rasetti, new species.

Stratigraphic range—Middle Cambrian (Wenkchemnia-Stephen-
aspis zone).

Remarks.—The species assigned to this genus show close affinity
with Bathyuriscus and especially Poliella. Wenkchemnia differs from
Bathyuriscus in the shallower dorsal furrow, small, transverse py-
gidium, and less distinct interpleural grooves. In Poliella the palpe-
bral lobes are longer, the posterior limbs are slender and do not taper
but rather expand distally, and the last thoracic segments curve back-
ward enveloping the pygidium.

The name derives from Wenkchemna Pass in the Bow Range.

WENKCHEMNIA WALCOTTI Rasetti, new species
Plate 11, figures 1-3

Bathyuriscus (Poliella) primus Watcott (part), Smithsonian Misc. Coll., vol.
64, No. 5, p. 352, pl. 46, fig. 6b (only), 1916.

In describing Bathyuriscus (Poliella) primus, Walcott remarked
both in the text and in the legend of the plate that there were speci-
mens of two types, differing in the number of thoracic segments and
the shape of the posterior limbs. Resser’s choice of a lectotype re-
stricts the name Poliella prima to the form with long palpebral lobes,
slender and expanding posterior limbs, and eight thoracic segments,
which is a good species of Poliella. The form with tapered posterior
limbs, short palpebral lobes, nine thoracic segments, and wide, short
pygidium presents sufficient differences to induce the writer to make
it the type of a new genus.

Known from numerous entire shields flattened in shale. Glabella
moderately expanded forward in the anterior half, delimited by a shal-
low dorsal furrow. Three or four pairs of glabellar furrows visible
but very shallow. Occipital furrow better impressed; occipital ring

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 185

moderately long, with a very short spine. Brim developed only at
the sides; rim slightly upturned, greatly reduced in front of the gla-
bella. Anterior facial suture parallel to the dorsal furrow, distant
from it one-fifth the glabellar width. Palpebral lobes defined by a
shallow palpebral furrow, narrow, moderately curved, obliquely
placed, about one-third the glabellar length, their posterior end dis-
tant from the glabella slightly more than one-half the glabellar width,
and situated on the transverse line through the posterior fourth of
the glabella. Posterior limbs tapered, slightly wider than the occipital
ring ; about half of the width comprised within the palpebral lobes.
Free cheeks rather narrow.

Thorax of nine segments. Axis slightly tapered, occupying some-
what less than one-third of the width. Pleura bent downward at
geniculation, which is situated at half the length of each pleuron;
furrowed for the entire length, directed transversely.

Pygidium twice wider than long, widely subtriangular. Anterior
margin almost straight. Axis prominent, somewhat tapered, with
three distinct segments and a terminal section, occupying most of the
pygidial length. Pleural lobes slightly convex. Pleural furrows shal-
low, three or four being visible; interpleural grooves indistinct. Mar-
ginal furrow shallow but distinct ; rim narrow, convex.

Surface apparently smooth.

Length of largest shield observed about 30 mm., of which 10 mm.
belong to the head, 15 mm. to the thorax, and 5 mm. to the tail.

Horizon and locality—Mount Whyte formation (Lake Agnes shale
lentil; Wenkchemmia-Stephenaspis zone). Type locality W3b, Plain
of Six Glaciers. Also localities U.S.N.M. 35m, southwest of Lake
Louise, and U.S.N.M. 35e, Mount Whyte.

Types.—Holotype: U.S.N.M. No. 116098. Paratypes: U.S.N.M.
Nos. 62625, 116099-100.

WENKCHEMNIA SPINICOLLIS Rasetti, new species
Plate 11, figures 4-8

Known from numerous cranidia, pygidia, and entire shields partly
flattened in shale.

Glabella of moderate convexity, defined by a shallow dorsal fur-
row, rather strongly expanded in the anterior third. Glabellar fur-
rows very shallow. Occipital furrow shallow; occipital ring extended
into a slender spine about one-fourth the glabellar length. Brim and
rim poorly differentiated from each other at the sides ; rim flat, greatly
reduced in front of the glabella. Palpebral lobes about one-third the

186 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

glabellar length, narrow, delimited by a shallow palpebral furrow;
maximum width of the fixed cheeks near the posterior end of the pal-
pebral lobes and equal to two-thirds of the glabellar width. Posterior
limbs slightly tapered, rounded distally, somewhat wider than the
occipital ring. Distance from posterior end of palpebral lobe to pos-
terior margin about equal to length of palpebral lobe. Anterior facial
sutures divergent, approximately parallel to the dorsal furrow.

Thorax of nine segments. Axial segments each with a spine or
node, broken off in all available specimens. Pleura straight, widely
furrowed, truncated at the extremity, the posterior end produced into
a sharp but very short spine.

Pygidium twice wider than long. Anterior outline almost straight,
anterior angles narrowly rounded. Axis strongly prominent, moder-
ately tapered, showing three rings and a terminal section defined by
shallow furrows, not quite reaching the posterior margin. Pleural lobes
slightly downsloping. Four pairs of rather wide, shallow pleural fur-
rows and very faint, narrow interpleural grooves. Marginal furrow
very shallow, rim slightly convex and poorly defined. Rim of first
pleural segment extended into a pair of short, blunt spines.

Surface apparently smooth.

Proportions of an entire shield are: Length of cephalon 7 mm., of
thorax 8 mm., of pygidium 3.3 mm. Length of largest shield ob-
served 40 mm.

This species is very similar to the genotype, from which it differs
chiefly in the occipital spine, axial spines on the thorax, and small
spines at the anterior angles of the pygidium.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
W enkchemma-Stephenaspis zone). Type locality Wok, Fossil Gully,
Mount Stephen. Also locality W16k, North Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116101. Paratypes: U.S.N.M.
No. 116102.

WENKCHEMNIA SULCATA Rasetti, new species
Plate 11, figures 9-15

Known from numerous cranidia, a few pygidia, and entire shields
preserved in limestone and shale.

The species is so similar in all parts of the carapace to W. spinicol-
lis that it is sufficient to describe it by listing the differences. The fact
that all specimens of W. spinicollis are more or less flattened in shale
while those of the new species preserve the full convexity makes an
exact comparison difficult. Nevertheless the following differences do
not appear to be due to the different manner of preservation. In the

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 187

new species the dorsal furrow is somewhat deeper, making the glabella
better defined, and the glabellar furrows are also deeper. The occipi-
tal ring carries only a short, blunt spine. The thorax presents no
differences. The pygidium has the small marginal spines at the an-
terior angles as in W. spinicollis, but the marginal furrow and rim
are narrower and less well defined.

Length of a complete shield 18 mm. Length of the largest cra-
nidium 13 mm.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Type locality W16j”, North Gully, Mount Ste-
phen. Other localities W16j, W16i, W16h, North Gully.

Types.—Holotype: U.S.N.M. No. 116103. Paratypes: U.S.N.M.
Nos. 116104-6.

Undetermined pygidium No. 1
Plate 15, figures 10, 11

Several specimens of this pygidium were collected from the Plagi-
ura-Kochaspis zone at various localities It has been impossible to as-
sign it with certainty to any of the associated cranidia, hence the
systematic position of this form remains obscure. It appears likely
that the pygidium belongs to a dolichometopid trilobite.

Entire pygidium subcircular, about as wide as long. Axis large,
occupying two-thirds of the length and more than half of the width;
strongly prominent, relatively short and wide, cylindrical; composed
of two rings and a terminal section, very poorly defined. A steep
postaxial slope reaches the posterior margin. Pleural lobes narrow,
concave in the transverse direction, without distinct rim, showing
but traces of segmentation. Anterior angles well rounded, posterior
outline uniformly curved.

Surface smooth. Length of largest pygidium 6 mm., width 6.5 mm.

The writer does not know of any described pygidium resembling
the present one.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Localities W4c-d, Ross Lake; W2od, Eiffel Peak.

Figured specimens.—U.S.N.M. No. 116141.

Undetermined pygidium No. 2
Plate 10, figure 10

Known from a single specimen in limestone.
This pygidium is subtriangular, truncated posteriorly. Axis moder-
ately tapered, strongly prominent, showing three rings and a terminal

188 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

section, extended into a short post-axial ridge that reaches the margin.
Pleural lobes approximately horizontal, flat, with five distinct, broad
furrows and very shallow interpleural grooves. Anterior angles well
rounded. Rim flat, narrow, differentiated by upturning rather than by
a marginal furrow. Length of pygidium 8 mm., width 12 mm.

This pygidium resembles that of Stephenaspis except that it lacks
marginal spines. It almost certainly belongs to a dolichometopid tri-
lobite. It is described because so little is known of the fauna of the
W enkchemnia-Stephenaspis zone.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Locality W16j, North Gully, Mount Stephen.

Figured specimen.—U.S.N.M. No. 116094.

Family CORYNEXOCHIDAE Angelin
Genus BONNASPIS Resser, 1936
Genotype: Karlia stephenensis Walcott.
BONNASPIS STEPHENENSIS (Walcott)
Plate 28, figures 4-6

Menocephalus salteri RoMINGER (not Devine), Proc. Acad. Nat. Sci. Philadel-
phia, 1887, p. 16, pl. 1, fig. 6.

Karlia stephenensis Watcott, Proc. U. S. Nat. Mus., vol. 11, p. 445, 1889; Ca-
nadian Alpine Journ., vol. 1, No. 2, pl. 3, fig. 4, 1908; Smithsonian Mise.
Coll., vol. 64, No. 3, p. 224, pl. 36, fig. 8, 1916.

Corynexochus romingeri MATTHEW, Trans. Roy. Soc. Canada, ser. 2, vol. 5,

SCCh 4, Dar 47a Dla es loses) eTSOO:

Corynexochus stephenensis (Walcott), Watcott, Smithsonian Misc. Coll., vol.
64, No. 5, p. 324, pl. 55, figs. 5-5¢, 1916.

Bonnaspis stephenensis (Walcott), REssER, Smithsonian Misc. Coll., vol. 95,
No. 4, p. 5, 1936.

Horizon and locality Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 14s (=author’s lo-
cality S8d), Mount Stephen.

Types.—Holotype: U.S.N.M. No. 61731. Paratypes: U.S.N.M.
Nos. 62717, 62718. Plesiotype: U.S.N.M. No. 116233.

Family DORYPYGIDAE Kobayashi
Genus KOOTENIA Walcott, 1889

Genotype: Bathyuriscus (Kootenia) dawsoni Walcott.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 189

KOOTENIA DAWSONI (Walcott)
Plate 27, figures 4-7

Bathyuriscus (Kootenia) dawsoni Watcott, Proc. U. S. Nat. Mus., vol. 11,
p. 446, 1880.

Dorypyge dawsoni (Walcott), MatrHew, Trans. Roy. Soc. Canada, ser. 2,
vol. 5, sect. 4, p. 56, pl. 3, fig. 1, 1899.

Dorypyge (Kootenia) dawsoni (Walcott), Watcorr, Canadian Alpine Journ.
vol. 1, No. 2, pl. 3, fig. 9, 1908.

Kootenia dawsoni (Walcott), WaAtcott, Smithsonian Misc. Coll., vol. 67, No. 4,
p. 131, 1918.

Horizon and locality—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s, Mount
Stephen.

Types.—Syntypes: U.S.N.M. No. 108495. Plesiotype: U.S.N.M.
No. 116229.

KOOTENIA BURGESSENSIS Resser
Plate 28, figures 9-11

Kootenia dawsom Watcott (part), Smithsonian Misc. Coll. vol. 67, No. 4,
p. 131, pl. 14, figs. 2, 3, 1918.

Kootenia burgessensis REssER, Smithsonian Misc. Coll., vol. 101, No. 15, p. 27,
1942.

Horizon and locahity.—Stephen formation (Burgess shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 35k, Burgess
Quarry, Mount Field. Also locality St1e, Mount Field.

Types.—Holotype: U.S.N.M. No. 65511. Paratypes: U.S.N.M.
Nos. 65512, 65533. Plesiotypes: U.S.N.M. No. 116234.

Genus OLENOIDES Meek, 1877
Genotype: Paradoxides nevadensis Meek.
OLENOIDES SERRATUS (Rominger)
Plate 27, figures 1-3

Ogygia serrata RomincerR, Proc. Acad. Nat. Sci. Philadelphia, p. 13, pl. 1,
figs. 2, 2a, 1887.

Olenoides nevadensis (Walcott, not Meek), Watcotr, Amer. Journ. Sci., ser. 3,
vol. 36, p. 165, 1888.

Neolenus serratus (Rominger), MAtrHEw, Trans. Roy. Soc. Canada, ser. 2,
vol. 5, sect. 4, p. 53, 1890.

Neolenus granulatus MATTHEW, Trans. Roy. Soc. Canada, ser. 2, vol. 5, sect. 4,
p. 55, pl. 2, figs. 1a-c, 1890.

190 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Neolenus serratus (Rominger), WAtcott, Canadian Alpine Journ., vol. 1, No. 2,
pl. 4, fig. 1, 1908; Smithsonian Misc. Coll., vol. 67, No. 4, p. 126, pls. 14-23,
1918.

Olenoides serratus (Rominger), KopayAsHI, Journ. Fac. Sci. Imp. Univ. Tokyo,
sect. 2, vol. 4, pt. 2, p. 153, 1035.

Horizon and locality —Stephen formation (Ogygopsis and Burgess
shale lentils). Type locality U.S.N.M. 14s, Mount Stephen. Also
locality U.S.N.M. 35k, Burgess Quarry, Mount Field.

Types.—Holotype: Acad. Nat. Sci. Philadelphia. Plesiotypes fig-
ured by Walcott: U.S.N.M. Nos. 57656-7, 58588, 58590, 65510,
65513-15, 65519-21. Plesiotypes figured by the writer: U.S.N.M.
Nos. 116227-8.

OGYGOPSIDIDAE Rasetti, new family
Genus OGYGOPSIS Walcott, 1889

Genotype: Ogygia klotzi Rominger.

The genus Ogygopsis was assigned to the Ordovician family Asa-
phidae, on the basis of superficial resemblance of the entire shield and
especially the macropygous development. However, this resemblance
is not substantiated by the really significant diagnostic features. In
the Asaphidae, the free cheeks meet along a median ventral suture.
Warburg (1925) remarked that the course of the dorsal sutures in
Ogygopsis does not suggest a structure of the ventral sutures of the
type observed in the Asaphidae, hence the placement of the genus in
the family is questionable. Study of the abundant material of Ogy-
gopsis klotzi from the Ogygopsis shale fully supports Warburg’s as-
sumption. Shield, exposed from the ventral side show that the free
cheeks are widely separated by the anterior portion of the hypostoma,
i.e., there is a pair of ventral sutures instead of a median suture as
in the Asaphidae. The hypostoma is fused with the rostrum. This
type of cephalic sutures is observed in the trilobites here assigned
to the superfamily Corynexochoidae. Ogygopsis entirely agrees with
this group not only in the course of the cephalic sutures but also in
the general features of the shield. In many respects Ogygopsis is
intermediate between the Dorypygidae and the Dolichometopidae ; the
cephalon is like Kootenia, the pygidium resembles those of Orria
and other macropygous Dolichometopidae. The hypostoma, however,
is relatively wider posteriorly than in any of the genera of these two
families and the anterior wings are less developed than in the Dory-
pygidae and Dolichometopidae. These characters suggest the assign-
ment of Ogygopsis to a new family Ogygopsididae. The characters

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI IQI

of the family are those of the type genus since no other described
genera appear to belong in the Ogygopsididae.

Ogygopsis was known only from the type species, Ogygopsis klotzt,
until Resser (1939b) described a new genus, Taxioura, based on a
new species, Tavrioura typicalis, from the “Ptarmigamia strata” of
Idaho. The only significant difference that the writer could discover
between Tavrioura typicalis and Ogygopsis klotzi is the lack of inter-
pleural furrows in the former species. This is not regarded as a
charaeter of generic importance, hence the writer considers Taxioura
as a subjective synonym of Ogygopsis.

On Mount Stephen, Ogygopsis ranges through an exceptional thick-
ness of strata and occurs in five different faunules, apparently rep-
resented by the same species. The earliest occurrence of Ogygopsis
is in the Syspacephalus laticeps faunule of the Wenkchemnia-Stephen-
aspis zone. The next occurrence is in the Stephenaspis bispinosa fau-
nule of the same zone. The next higher occurrences are in the Yoho-
aspis pachycephala and Chancia bigranulosa faunules, tentatively
assigned to the Albertella zone, collected from the shaly and calcareous
equivalent of the lower Cathedral formation on Mount Stephen. The
latest occurrence is in the Ogygopsis shale, whose faunule belongs to
the Bathyuriscus-Elrathina zone.

OGYGOPSIS KLOTZI (Rominger)
Plate 12, figures 1-5; Plate 21, figures 1-3; Plate 20, figures 6-8

Ogygia klotsi RomincEr, Proc. Acad. Nat. Sci. Philadelphia, 1887, p. 12, pl. 1,
fig. I.

Ogygopsis klotzi (Rominger), Watcott, Proc. U. S. Nat. Mus. vol. 11, p. 446,
1889; Canadian Alpine Journ., vol. 1, No. 2, pl. 4, fig. 4, 1908; Smithsonian
Misc. Coll., vol. 64, No. 5, p. 377, pl. 66, figs. 1, 1a-b, 1916.

Ogygia (Ogygopsis) klotzi Rominger, MATTHEW, Trans. Roy. Soc. Canada,
ser. 2, vol. 5, sect. 4, p. 58, 1890.

This species was collected at five different horizons ranging from
the Wenkchemnia-Stephenaspis zone in the Mount Whyte formation
to the Ogygopsis shale of the Stephen formation which is the typical
horizon. Entire shields are known from four of these horizons, and
show no difference in the thorax, which has eight segments in all
specimens. The cranidia and pygidia were compared as accurately
as permitted by the imperfect state of preservation. All specimens
from the Ogygopsis shale are flattened to a greater or lesser extent
according to size. Specimens from the Cathedral formation (localities
Coh, Cgj) are either flattened or not according to the shaly or cal-
careous composition of the layer in which they occur, but in either

I92 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

case are poorly preserved in the coarse matrix. Identical conditions
of preservation obtain at locality W16j in the Wenkchemnia-Stephen-
aspis zone of the Mount Whyte formation. Distortion is present at
all localities and horizons. Notwithstanding these circumstances, a
relatively accurate comparison of the material from different beds
is favored by the large size of this trilobite and the possibility of com-
paring specimens preserved in the same type of matrix. This study
did not reveal differential characters in the cranidium or pygidium.
Hence Ogygopsis klotzi seems to possess a vertical range unprece-
dented among Cambrian trilobites. Specimens from different horizons
are figured.

Horizon and locality—(1) Stephen formation (Ogygopsis shale
lentil; Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s,
Mount Stephen.

(2) Cathedral formation (Albertella (?) zone). Localities Coj,
Coh, Fossil Gully, Mount Stephen.

(3) Mount Whyte formation (Yoho shale lentil; Wenkchemnia-
Stephenaspis zone). Locality Wok, Fossil Gully, Mount Stephen.

(4) Mount Whyte formation (Wenkchemnia-Stephenaspis zone).
Locality W16j, North Gully, Mount Stephen.

Types—Holotype: Acad. Nat. Sci. Philadelphia. Plesiotypes:
U.S.N.M. Nos. 62846-8, 116107-8, 116178, 116239, 24040.

OGYGOPSIS SPINULOSA Rasetti, new species
Plate 21, figure 4

Known from a single pygidium preserved in impure limestone and
not flattened.

The pygidium resembles the associated O. klotzi in all respects
except the lack of interpleural furrows, somewhat shorter axis, and
especially the denticulated margin. Each pleural segment is extended
into a minute spine, nine pairs being distinguishable, the last two
represented by a faint waviness of the margin. Length 22 mm., width
30 mm.

Horizon and locality.—Cathedral formation (Albertella (?) zone).
Locality Coh, Fossil Gully, Mount Stephen.

Type.—Holotype: U.S.N.M. No. 116179.

Family ORYCTOCEPHALIDAE Beecher

This family represents a very homogeneous, closely interrelated
group of genera when spurious forms that at one time or other were
included in the Oryctocephalidae are eliminated. The writer considers

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 193

the family to include the genera Oryctocara, Oryctocephalites, Oryc-
tocephalus, and Tonkinella.

Genus ORYCTOCEPHALUS Walcott, 1886

Genotype: Oryctocephalus primus Walcott.

ORYCTOCEPHALUS, species undetermined No. 1
Plate o, figure 24

A single cranidium of a species of Oryctocephalus is figured in
order to present what probably is the earliest known occurrence of the
genus.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Locality W16j, North Gully, Mount Stephen.

Figured specimen.—U.S.N.M. No. 116090.

ORYCTOCEPHALUS, species undetermined No. 2
Plate 9, figure 23

A species of Oryctocephalus is exceedingly rare in the Yoho shale.
Specific identification is not attempted as this form is represented only
by flattened cranidia.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
Wenkchemma-Stephenaspis zone). Locality Wok, Fossil Gully,
Mount Stephen.

Figured specimen.—U.S.N.M. No. 116089.

ORYCTOCEPHALUS REYNOLDSI Reed
Plate 29, figures 4, 5

Oryctocephalus reynoldsi RrEep, Geol. Mag., new ser., dec. IV, vol. 6, No. 8,

p. 350, text fig., 1899.
Oryctocephalus reynoldsi Reed, Watcott, Canadian Alpine Journ., vol. 1, No. 2,

pl. 3, fig. 1, 1908.

This rare species, described from the Ogygopsis shale, also occurs
in the Burgess shale. An excellently preserved shield from the latter
locality is figured, as well as a specimen from the type locality.

Horizon and locality—Stephen formation (Ogygopsis and Bur-
gess shale lentils ; Bathyuriscus-Elrathina zone). Type locality U.S.-
N.M. 14s, Mount Stephen. Also locality U.S.N.M. 35k, Burgess
Quarry, Mount Field.

Types.—Holotype: Woodwardian Museum. Plesiotypes: U.S.N.M.
Nos. 24078, 116237-8.

194 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ORYCTOCEPHALUS BURGESSENSIS Resser
Plate 26, figures 1-3

Oryctocephalus primus Kopayasuti (not Walcott), Journ. Fac. Sci. Imp. Univ.
Tokyo, sect. 2, vol. 4, pt. 2, p. 147, pl. 15, fig. 1, 1935.

Oryctocephalus burgessensis RESSER, Smithsonian Misc. Coll., vol. 97, No. 10,
Pp. 37, 1938.

Resser’s brief description of the species can be supplemented,
thanks to the abundant material available.

Glabella moderately expanded forward in the posterior half, some-
what pear-shaped, almost reaching the anterior margin of the head.
First pair of glabellar furrows indistinct, second and third pairs rep-
resented by fairly deep lateral pits, fourth pair consisting of lateral
pits connected by a shallow transverse furrow; occipital furrow deep
but not reaching the dorsal furrow. Occipital ring short and simple.
Fixed cheeks somewhat narrower than the glabella. Ocular ridges
straight, parallel to the anterior margin of the cephalon. Palpebral
lobes slightly oblique, moderately curved ; distance from posterior end
of palpebral lobe to posterior margin of cephalon somewhat less than
length of palpebral lobe.

Thoracic axis without nodes or spines. Pleural furrows starting
near the anterior end of each pleuron, first directed obliquely outward
and backward, then parallel and close to the posterior margin of the
pleuron. Pleura terminating in slender, straight spines increasing in
length from the first to the seventh segment; those of the seventh
segment exceeding in length the remaining portion of the pleuron.

Pygidial axis tapered, occupying about two-thirds of the pygidial
length, with five segments and a short terminal section, without nodes
or spines. Six pairs of pleural furrows all well impressed, the furrows
of the last pairs directed longitudinally and close to each other. First
to third pairs of interpleural grooves almost indistinct, fourth and
fifth pairs well impressed, slightly concave outward while the cor-
responding pleural furrows are convex outward. Six pairs of long,
slender marginal spines. Spines of the first to third pairs about
equally long and strong, spines of fourth pair strongest and longest,
still directed outward, spines of fifth and sixth pairs slenderer,
straight, directed slightly inward. Average length of spines somewhat
less than pygidial length.

Surface of test smooth. Length of adult shield, exclusive of spines,
16 mm.

This species is in almost all characters intermediate between the
two other forms occurring in the Burgess shale, O. reynoldsi and

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 195

O. matthewi. It may be synonymous with O. walkeri Matthew, but
the question cannot be decided without studying the types of the latter
species.

Horizon and locality—Stephen formation (Burgess shale lentil;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 35k, Burgess
Quarry, Mount Field.

Types.—Holotype: U.S.N.M. No. 96487. Plesiotypes: U.S.N.M.
No. 116220.

ORYCTOCEPHALUS MATTHEWI Rasetti, new species
Plate 26, figures 4, 5

Known from several complete shields flattened in shale.

The cranidium does not require a complete description, as it is
almost identical with that of O. burgessensis, possibly excepting
slightly shallower glabellar furrows and shorter palpebral lobes.

Thoracic axis without spines or nodes. Pleural furrows wider and
shallower than in O. burgessensis. Pleural spines short, rapidly
tapered, even the longest, those of the seventh segment, not equaling
half the length of the pleuron.

Pygidial axis tapered, without nodes or spines, showing five rings
and a terminal section, occupying two-thirds of the pygidial length.
Pleural lobes with six pairs of furrows, the last two shallower ; fur-
rows of the last pair parallel to each other. First three pairs of inter-
pleural grooves indistinct, last two pairs shallow but distinct, dividing
the spaces between pleural furrows more evenly than in O. burges-
sensis. Six pairs of marginal spines, all tapered, relatively short, of
about equal strength.

Surface of test smooth. Length of largest shield 16 mm.

This species differs from the associated O. burgessensis chiefly in
the shortness of the marginal thoracic and pygidial spines.

Horizon and locality—Stephen formation (Burgess and Ogygopsis
shale lentils ; Bathyuriscus-Elrathina zone). Type locality U.S.N.M.
35k, Burgess Quarry, Mount Field. Also locality U.S.N.M. 14s,
Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116221. Paratypes: U.S.N.M.
No. 116222.

Genus ORYCTOCEPHALITES Resser, 1939
Genotype: Oryctocephalites typicalis Resser.

196 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

ORYCTOCEPHALITES RESSERI Rasetti, new species
Plate 15, figure 9

Known from two cranidia preserved in limestone.

Glabella expanded forward, well rounded in front, moderately con-
vex transversely and longitudinally, defined by a narrow but rather
well-impressed dorsal furrow. Glabellar furrows represented by three
pairs of round pits; the posterior and median ones connected across
the glabella by shallow furrows. Occipital furrow consisting of deep
pits at the sides and shallow median portion; not reaching the dorsal
furrow. Anterior pits well developed, indenting the sides of the gla-
bella immediately in front of the ocular ridges. Anterior outline of cra-
nidium strongly oblique at each side. Rim short, convex. Brim rep-
resented by short bands at the sides between the border and the ocular
ridges. Ocular ridges well developed, parallel to the anterior margin.
Palpebral lobes narrow, somewhat elevated, about two-fifths the gla-
bellar length. Fixed cheeks downsloping, slightly concave; anterior
width about one-third the glabellar width, posterior width almost
equal the glabellar width. Posterior limbs not entirely preserved,
but evidently not extending far beyond the palpebral lobes, strongly
bent downward distally. Posterior marginal furrow deep.

Surface covered with exceedingly fine granules. Length of largest
cranidium 6.5 mm.

The species is assigned to Oryctocephalites rather than Oryctocepha-
lus because of the expanding glabella and the strongly curved an-
terior outline of the cranidium. It differs from O. typicalis Resser
in the strong development of the anterior pits and the concavity of the
fixed cheeks.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W7f, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116139. Paratype: U.S.N.M.
No. 116140.

Genus TONKINELLA Mansuy, 1916
Genotype: Tonkinella flabelliformis Mansuy.

TONKINELLA STEPHENSIS Kobayashi
Plate 31, figures 13-18

Tonkinella stephensis KopayAsut, Journ. Fac. Sci. Imp. Univ. Tokyo, sect. 2,
vol. 4, pt. 2, p. 149, pl. 15, figs. 2-5, 1935.

This species was adequately described by Kobayashi, except for his
statement that the thorax has six segments. The only entire shield

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 197

that was available to him is very poorly preserved, and the error is
explicable. Better-preserved shields show unmistakably that there
are only five thoracic segments even in the largest individuals.

A well-preserved cranidium among the material illustrated by
Kobayashi is chosen as lectotype. Among the specimens collected by
the writer is an unusually large shield 25 mm. long.

This species is a common, readily recognizable diagnostic fossil
of the black limestones that constitute the upper portion of the Ste-
phen formation.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality U.S.N.M. 58j, Mount Stephen, probably close
to or identical with the author’s locality Sgk. Other localities S6k,
S6l, Mount Odaray; Stok, Siol, Park Mountain; Sgk, Mount Ste-
phen.

Types.—Lectotype and paratypes: U.S.N.M. No. 108497. Plesio-
types: U.S.N.M. Nos. 116255-6.

Family undetermined
Genus HANBURIA Walcott, 1916
Genotype: Hanburia gloriosa Walcott.
HANBURIA GLORIOSA Walcott
Plate 25, figure 15

Hanburia gloriosa Watcott, Smithsonian Misc. Coll., vol. 64, No. 3, p. 226,
pl. 36, figs. 2, 3, 1916.

Hanburia gloriosa Walcott, Kopayasui, Journ. Fac. Sci. Imp. Univ. Toyko,
sect, 2, vol. 4; pt:.2, p. 143, pl. 14, fig. 15, 1935.

Besides the shield figured by Walcott and a few other poor speci-
mens from the the type locality, there is available an impression of
a shield collected from the Ogygopsis shale by Resser.

All the specimens are flattened and the characters of the cephalon
are somewhat obscure. Walcott questioned the presence of dorsal
facial sutures. Kobayashi stated that “In examining the type speci-
mens under the high magnification with crossed light, they are seen to
have the same kind of facial suture and eyes as in Ogygopsis.”

The writer disagrees with this conclusion and believes that the
species has no dorsal cephalic sutures. Examination of casts of the
new specimen supports this conclusion. The lines that Kobayashi
assumed to be sutures seem to be cracks formed in the flattening of
the shield, as they appear on one side only in each specimen, and in

198 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

different positions in the various specimens; furthermore, the writer
was unable to detect any trace of palpebral lobes. Apparently Han-
buria was a blind trilobite, with free cheeks confined to the doublure
and a marginal cephalic suture as in the Conocoryphidae. However,
the expanding glabella and the general aspect of the shield indicate
a blind relative of the Dolichometopidae or other families of the
Corynexochoidae, and no relationship with the Conocoryphidae which
are obviously an aberrant branch of the Ptychoparioidae. Until more
becomes known of this trilobite, it seems better to leave its more
precise systematic position undetermined.

Horizon and locality Stephen formation (Burgess and Ogygopsis
shale lentils; Bathyuriscus-Elrathina zone). Type locality U.S.N.M.
35k, Mount Field. Also locality U.S.N.M. 14s, Mount Stephen.

Types.—Syntypes: U.S.N.M. Nos. 61724-5.

Superfamily PTYCHOPARIOIDAE

The superfamily name Ptychoparioidae (emendation of Ptycho-
pariidea R. Richter, 1932) is used instead of the new name Conoco-
ryphidea (Conocoryphoidae) suggested by R. and E. Richter (1941)
for priority reasons.

The writer does not believe that the priority rule should be fol-
lowed at the family, superfamily, and order levels whenever its strict
application would require one of these groups to be named after a
genus either poorly known, or nontypical in morphology, age, or
geographical distribution; any of these reasons may induce future
taxonomists to remove the genus from the group that it is supposed to
represent, thus causing instability and confusion in nomenclature. In
the present case Ptychoparia and the Ptychopariidae are perfectly
typical examples of the large group of trilobites assembled under the
superfamily name, whereas Conocoryphe represents an aberrant stock
whose close relationship with the Ptychopariidae has even been ques-
tioned (Poulsen, 1927).

The great difficulty of classifying the Ptychoparioidae is well known
to every student of Cambrian faunas. We find an immense number
of forms intergrading in all observable characters and thus making
many of the suggested generic subdivisions appear artificial and of
questionable practical application. The status of the proposed family
arrangements is even more doubtful.

Students of the Cambrian up to the first decade of this century
solved the problem by referring almost all these trilobites to Cono-
cephalites or Ptychoparia. However, with the discovery of new forms

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 199

it became apparent that the genus Ptychoparia was attaining an un-
wieldy size, and many new genera were established. Today the genera
made to receive forms that earlier workers placed in Ptychoparia
number over 75 for the Lower and Middle Cambrian forms of North
America alone (exclusive of the Atlantic province) ; to which an even
greater number of genera based on European, Asiatic, and Australian
forms could be added. Many more genera were established for Up-
per Cambrian species.

Yet everyone who undertakes the study of a Cambrian fauna soon
meets ptychoparid forms that do not possess the exact combination
of characters of any described genus, according to present standards
of close discrimination. Short of falling back on the old practice
(with which the writer at times feels strongly inclined to sympathize),
of using Ptychoparia s. 1., the only course left is to make further new
genera.

The taxonomy of this group, beset with intrinsic difficulties, has
been made more confused by the careless work of Cambrian paleon-
tologists. The most objectionable practice responsible for unnecessary
confusion has been basing new genera on poorly represented geno-
types. For example, some of the important Lower Cambrian genera,
such as Periomma, Antagmus, Bullingsaspis, Onchocephalus, etc.,
were based on species known from one or a few poorly preserved,
often fragmentary cranidia. These forms are difficult to discriminate
from each other even when complete cranidia ideally preserved in
limestone are available; the problem becomes almost hopeless when
generic names are tied to weathered and distorted specimens. Several
genera were based on such poor genotypes, when excellently preserved
material both from the Appalachian and the Cordilleran provinces was
readily available.

Another source of difficulties, which cannot be laid to the door of
Cambrian paleontologists, is that some of the genera were based ex-
clusively on the characters of the cranidium ; this is the almost general
rule for species preserved in limestone, where all the parts of the
shield occur separately, and the simple, small ptychoparid pygidia and
thoracic segments escape observation, or cannot be assigned to definite
cranidia. However, when genera were founded on genotypes known
from specimens in shale, often generic characters were taken from
features of the entire shield, whereas the cranidial characters could be
less accurately ascertained because of the flattening and poor preserva-
tion of the test. It is sometimes exceedingly difficult to compare
genera based on specimens preserved in the two different manners.

200 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

It appears that few of the characters used to distinguish ptycho-
parid genera are of great significance; in fact, in other groups of tri-
lobites similar characters would not be considered of generic im-
portance. Some of these characters and the general evolutionary
trends of the ptychoparids deserve a brief discussion.

The Ptychoparioidae, in their more primitive forms appearing in the
Early Cambrian, are typically micropygous (Rasetti, 1948a). These
generalized ptychoparids are usually characterized by well-developed
dorsal, glabellar, and marginal furrows; the conical glabella occupies
a moderate fraction of the cephalic area, the brim and rim are well
developed, the anterior facial sutures slightly diverge toward the
front, the frontal portion of the suture is marginal. The thorax, when
known, usually includes 12 to 20 simple segments. The pygidium is
small, transverse, often appearing composed of one or two segments
only. Distinguishing generic characters can usually be found only in
the cranidia. Forms that have been assigned to Antagmus, Piazella,
Ptychoparella, Onchocephalus, Proliostracus, Syspacephalus, etc., typ-
ify this group of primitive, generalized ptychoparids. Ptychoparia
itself, although of Medial Cambrian age, has not greatly evolved from
these older forms.

In Medial Cambrian times we note two distinct evolutionary tend-
encies. One is toward an extension and flattening of the brim and
rim, widening of the doublure, and reduction of the proportionate
size of the glabella; genera like Alokistocare and Amecephalus ex-
emplify this tendency. Often groups evolved in the opposite direction,
leading to forms with very convex cranidia, large glabellae, strong,
convex rims and narrow doublures. These trilobites, as generally all
the very convex forms, developed a thicker test than in the preceding
group. Solenopleura is the typical example. However, generalized
ptychoparids as described under the first group remained abundant
throughout the Medial Cambrian epoch.

In the later part of the Medial Cambrian, certain ptychoparids
became macropygous, with corresponding reduction of the number
of thoracic segments ; in certain cases, the tail became as large as the
head. Asaphiscus and Glyphaspis are outstanding examples.

A somewhat anomalous group of ptychoparids, beginning in early
Medial Cambrian time, developed spinose pygidia. Kochaspis is the
typical genus of this group, and possibly the Upper Cambrian Crepi-
cephalus is its descendant.

There are thus at least two groups of ptychoparids where the
characters of the pygidium are important for generic distinction.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 201

However, in most cases only cranidia are available, and it may be very
difficult to separate such forms as the micropygous Alokistocare from
the macropygous Glyphaspis, or to distinguish cranidia of Kochaspis
from those of several genera with simple pygidia.

It is likely that at least some of these various modifications of the
ptychoparids developed independently in different stocks, so that
genera based on these characters may not represent phylogenetic
groups. However, since more significant characters for the subdi-
vision of the ptychoparids, if they existed in the living animal, are
unknown to us, there is little justification for separating apparently
similar forms on the mere suspicion that they may not be closely
related. A similar procedure was adopted by R. and E. Richter in
uniting all the blind ptychoparids under the family Conocoryphidae,
in spite of the strong suspicion that the genera of this family derived
from different stocks of oculate ptychoparids.

A tendency that is observed in several groups in Medial Cambrian
time is the reduction of the furrows, culminating in smooth forms
that are the most difficult to classify. Their similarity is of little
significance, being based on the absence rather than the presence of
common diagnostic features.

A recent attempt to classify a group of Lower Cambrian ptycho-
parid genera was made by Lochman (1947). She discussed the genera
Antagmus, Litocodia, Onchocephalus, Periomma, Periommella, Plagi-
ura, Plagiurella, Proliostracus, Ptychoparella, and Syspacephalus, and
established two new genera, Crassifimbra and Piazella. Two of these
genera (Plagiura-Plagiurella and Periommella) are very distinctive
and require no comments. The others all belong to the generalized,
most difficult group of ptychoparids. Lochman intended to make the
generic distinctions in this group less subjective by establishing a list
of “diagnostic generic features.” She believes that “‘a definite, recogni-
zable variation in any one of these features constitutes a valid generic
distinction.”

The features mentioned are chiefly proportions of the various
cranidial parts and angles between surfaces. As an example, she
separates two genera chiefly on the basis of the width of the fixed
cheeks being three-fourths and two-thirds of the glabellar width re-
spectively. The writer, having measured that ratio for a considerable
number of Lower and early Middle Cambrian ptychoparids, found,
as expected, values ranging anywhere between 0.5 and 1.0 without
any preference for the simple fractions mentioned by Lochman.
Whatever limits are set to the generic characters, there will always

202 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

be intermediate cases. Furthermore, the writer does not consider
some of the proportions used by Lochman as important. For example,
the relative width of the fixed cheeks is known to vary to a great
extent even within some of the genera of trilobites that represent the
best-established phylogenetic units.

A set of diagnostic generic features may be used, as done by Loch-
man, as a temporary expedient to put a certain degree of order in a
difficult group such as the Lower Cambrian ptychoparids, but it should
be understood that its adoption is apt to lead to an excessively arti-
ficial grouping of forms. It must be remembered that many of the
best-established phylogenetic groups of extinct organisms cannot be
defined in terms of the characters preserved in the fossils, yet we are
able to recognize such groups through their morphologic and chrono-
logic continuity.

An example that illustrates the above-mentioned principle is the
following. The Olenidae of the Atlantic province are one of the
best-established trilobite groups, their intergrading genera showing an
obvious affinity. Yet the writer believes that it is impossible to formu-
late a set of diagnostic features that will give an objective description
of the olenid shield and enable one to decide whether a certain trilobite
belongs to the Olenidae or not. It is likely than an objective definition
of the Olenidae would be possible if the soft parts were preserved ;
as many groups of living animals could no longer be defined if we
were to use only the characters that would be preserved in a fossil.

The genera of the Ptychoparioidae are not assigned to families
and are arranged in alphabetical order.

Genus ALOKISTOCARE Lorenz, 1906

Genotype: Conocephalites subcoronatus Hall and Whitfield.

Several generic names are available for Middle Cambrian ptycho-
parid trilobites with flat cranidium, long, on the average concave
frontal limb, wide fixed cheeks, numerous thoracic segments and a
small pygidium. The forms of this type are numerous and intergrade
in most of their characters, hence it is difficult to establish generic
limits. The generic names Alokistocare Lorenz, Amecephalus Wal-
cott, Chancia Walcott, Amecephalina Poulsen, and Strotocephalus
Resser have been proposed. Resser did not recognize Amecephalus
which he considered a synonym of Alokistocare.

The genotype of Alokistocare is known only from cranidia. If
forms known from complete shields and described by Resser as
Alokistocare belong to the genus, they appear to differ from Ame-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 203

cephalus piochensis in possessing somewhat narrower fixed cheeks and
in the different form of the thoracic pleura, which extend into falcate
terminations in Amecephalus piochensis, while they remain almost
straight in the forms assigned to Alokistocare. Chancia (genotype:
Chancia ebdome Walcott) differs from the other above-mentioned
genera in the more convex fixed cheeks and better-defined rim.
Amecephalina is based on a cranidium from Greenland (Amecepha-
lina mirabilis Poulsen) to which Poulsen tentatively assigned a fairly
large, multisegmented pygidium. If this part of the shield belongs to
the species the genus deserves recognition, otherwise it is doubtful
whether the cephalic characters would justify generic separation from
Amecephalus or Alokistocare. Strotocephalus (genotype: Stroto-
cephalus gordonensis Resser) is known only from cranidia which,
as far as the writer can see, do not differ generically from Amece-
phalus or Alokistocare. Finally it should be mentioned that Kochiella
cannot be easily distinguished from Amecephalus or Alokistocare as
far as the cranidia are concerned, while the characters of other parts
of the shield are uncertain (see discussion of genus).

The writer, at least provisionally, recognizes Amecephalus and
Alokistocare as distinct on the basis of the above-mentioned charac-
ters; however, it must be borne in mind that the diagnostic features
assigned to Alokistocare are uncertain because they are in part de-
rived from species other than the genotype. Amecephalus, as here
construed, occurs on the average in older strata than Alokistocare. Of
the forms described in this paper, those assigned to Amecephalus
occur in the Wenkchemmnia-Stephenaspis and Plagiura-Kochaspis
zones, while the species assigned to Alokistocare occur in the Bathy-
uriscus-Elrathina zone. In other areas of the Cordilleran province,
species assignable to these genera frequently occur also in the Al-
bertella and Glossopleura zones.

ALOKISTOCARE PARANOTATUM Rasetti, new species
Plate 33, figures 17, 18

Known from several cranidia preserved in limestone.

Glabella tapered, moderately convex, somewhat truncated in front.
Glabellar furrows very shallow. Occipital furrow deep at the sides,
shallow mesially ; occipital ring expanded backward mesially, bearing
a small node. Frontal limb two-thirds as long as the glabella, on the
average concave mesially ; brim convex and downsloping at the sides.
Rim flat, upturned with respect to the brim; midlengths of rim and
brim about equal. Marginal furrow deeper laterally than mesially.

204. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Fixed cheeks slightly convex and upsloping, almost as wide as the
glabella at the maximum. Palpebral lobes almost half the glabellar
length, moderately wide, set off by well-impressed palpebral furrows.
Ocular ridges thick and prominent. Posterior limbs deeply furrowed.
Anterior facial sutures somewhat divergent, almost straight to the
anterior margin; anterior angles of cranidium narrowly rounded.
Posterior branch forming slender posterior limbs, wider than the oc-
cipital ring. Distance from posterior end of palpebral lobe to
posterior margin somewhat less than length of palpebral lobe.

Surface of test densely covered with fine granules.

Length of largest cranidium 7.5 mm.

This form very closely resembles several species described by Res-
ser from the Lakeview limestone of Idaho, especially A. natale and
A. notatum. The new species chiefly differs from A. natale in the
deeper dorsal furrow and shallower glabellar furrows; from A. no-
tatum in the greater relative midlength of the brim.

Horizon and locality —Stephen formation (Bathyuriscus-Elrathina
zone). Locality Sgk, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116277. Paratypes: U.S.N.M.
No. 116278.

ALOKISTOCARE SINUATUM Rasetti, new species
Plate 34, figures I, 2

Known from several cranidia preserved in limestone.

Glabella weakly convex, strongly tapered, moderately truncated
in front, with somewhat concave sides. Fourth pair of glabellar fur-
rows turning backward and almost isolating the basal glabellar lobes ;
third pair shorter and very shallow, first and second pairs indistinct.
Occipital furrow impressed throughout ; occipital ring short, bearing
at most a small node. Frontal limb somewhat more than two-thirds
the length of the glabella, with little relief. Brim convex laterally, on
the average flat mesially ; rim of moderate thickness, convex, set off
by a distinct marginal furrow. There is a median swelling of the
anterior portion of the brim that extends backward mesially for
about one-third of the midlength of the brim. In such cases, it is
difficult to decide whether the marginal furrow is the one in front
of this swelling (as assumed in the foregoing description) or the
one back of it (i.e., the swelling belongs to the rim rather than the
brim). Fixed cheeks somewhat upsloping and convex. Ocular ridges
straight, moderately prominent. Palpebral lobes not preserved; ap-
parently of size and position usual in the genus. Anterior facial

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 205

sutures considerably divergent ; anterior angles of cranidium widely
rounded. Posterior branch almost parallel to the posterior margin,
defining slender, deeply furrowed posterior limbs.

Length of largest cranidium 18 mm.

Surface of test covered with small granules, among which are
scattered granules of a much larger size; brim with the usual longi-
tudinal anastomosing lines.

This species differs from all the described forms of the genus in
the shape of the glabella and glabellar lobation.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S6l, Mount Odaray.

Types—Holotype: U.S.N.M. No. 116281. Paratypes: U.S.N.M.
No. 116282.

ALOKISTOCARE CATARACTENSE Rasetti, new species
Plate 34, figures 3, 4

Known from numerous cranidia preserved in limestone but more or
less flattened and distorted.

Glabella weakly convex, relatively wide and short, tapered at
average rate, straight-sided, truncated in front. Glabellar furrows
very shallow, visible chiefly as smooth areas on the otherwise strongly
ornamented surface. Occipital furrow impressed throughout; occip-
ital ring short and simple. Frontal limb as long as the glabella, on
the average slightly concave mesially. Rim upturned, poorly defined
by a very broad marginal furrow. Brim with a strong median swell-
ing, more posteriorly situated than in A. sinuatum. A faint, sinuous
line defines the posterior margin of the swelling, and should possibly
be interpreted as the marginal furrow. Fixed cheeks slightly convex
and upsloping. Ocular ridges straight, fairly prominent; palpebral
lobes not preserved. Anterior facial sutures considerably divergent ;
anterior angles of cranidium well rounded. Posterior branch almost
parallel to the posterior margin, defining slender posterior limbs.

A probable incomplete thorax shows 22 segments.

Surface of test covered with very fine granules, among which are
scattered granules of a much larger size. On the brim and free
cheeks the granules tend to arrange themselves along the usual anasto-
mosing lines.

Length of largest cranidium 20 mm. Length of incomplete shield
with cephalon and 22 thoracic segments, 58 mm.

This species resembles several forms described by Resser from the
Spence shale ; it apparently differs from all of them in the considerable
swelling of the brim and in the surface ornamentation.

206 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality Srol, Park Mountain.

Types.—Holotype: U.S.N.M. No. 116283. Paratypes: U.S.N.M.
No. 116284.

Genus ALOKISTOCARELLA Resser, 1938
Genotype: Alokistocarella typicalis Resser.
ALOKISTOCARELLA FIELDENSIS Rasetti, new species
Plate 30, figures 3-5

Known from numerous shields somewhat flattened in argillaceous
limestone.

Glabella moderately tapered, rounded in front, of average con-
vexity. Glabellar furrows indistinct; occipital furrow straight, mod-
erately impressed; occipital ring short and simple. Frontal limb
somewhat shorter than half the glabellar length, on the average con-
cave. Rim occupying more than half the midlength of the limb, de-
limited by a shallow marginal furrow, slightly concave and upturned.
Fixed cheeks somewhat more than half the glabellar width. Palpebral
lobes small, narrow, situated slightly in advance of the glabellar mid-
point. Ocular ridges faint. Distance from posterior end of palpebral
lobe to posterior margin about three times length of palpebral lobe.
Posterior limbs deeply furrowed. Anterior branch of facial suture
slightly divergent in front of the eye, curving across the rim. Pos-
terior branch first directed outward and somewhat backward, then
curving backward. Width of posterior limbs greater than width of
occipital ring. Free cheeks with poorly defined rim and short, slender
genal spines.

Thorax of 17 segments. Axial rings simple. Pleura considerably
wider than axis; geniculation near halfway in anterior pleura, shift-
ing nearer axis in posterior pleura. Pleural furrows rather narrow
and deep, close to anterior margin of pleura.

Pygidium small, three times wider than long, subelliptical. Axis
faintly elevated, occupying about four-fifths of the length and more
than one-third of the width, rounded at the end. Axial furrows shal-
low. Two pairs of shallow pleural furrows, almost paralleling the
margin, hence curving backward and then slightly inward. Rim
indistinct.

Surface of test smooth, as far as can be ascertained from the
moderately well-preserved specimens.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 207

Length of largest cranidium 10 mm. Length of holotype specimen
13.8 mm., of which 4.9 mm. belong to the cephalon, 8 mm. to the
thorax, and 0.9 mm. to the pygidium,

This species appears referable to the genus, insofar as its cranidial
characters agree with those of the genotype, known from cranidia
alone. The species could not be referred to Alokistocare on account
of the larger proportions of the glabella with respect to the other
cranidial parts, and the writer did not find any other described genus
combining the characters of concave frontal limb, numerous thoracic
segments and very small pygidium. Compared with A. typicalis, the
present species has shorter and less-elevated palpebral lobes and a
somewhat different structure of the frontal limb.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S1tre, Mount Field.

Types——Holotype: U.S.N.M. No. 116241. Paratypes: U.S.N.M.
No. 116242.

Genus AMECEPHALUS Walcott, 1924
Genotype: Ptychoparia piochensis Walcott (part).

AMECEPHALUS AGNESENSIS (Walcott)
Plate 10, figures II-15

Olenopsis? agnesensis Watcott, Smithsonian Misc. Coll., vol. 57, No. 8, p. 242,
pl. 36, fig. 2, 1912; idem (part), vol. 67, No. 3, pl. 13, fig. 5, 1917.

Kochiella agnesensis (Walcott), Watcott, Smithsonian Misc. Coll., vol. 75,
No. 5, p. 302, 1928.

Alokistocare agnesensis (Walcott), REssEr, Smithsonian Misc. Coll., vol. 93,
No. 5, p. 9, 1935.

This species is common in the Lake Agnes shale, where it is rep-
resented by complete shields. These, however, are flattened, hence
the convexity cannot be ascertained, and close comparison with forms
preserved in limestone is difficult. A new description follows.

Entire shield subelliptical, the posterior portion of the thorax taper-
ing rapidly to a small pygidium. Cranidium of the general shape
typical of Amecephalus. The convexity cannot be ascertained, all
known specimens being flattened in shale, but apparently was not
great. Dorsal furrow apparently very shallow. Glabella narrow,
moderately tapered, somewhat truncated in front. Glabellar furrows
almost indistinct, occipital furrow shallow. Length of glabella (in-
clusive of occipital ring) about three-fifths of cranidial length. Brim
and rim poorly differentiated by a narrow, shallow marginal furrow,

208 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

closer to the anterior margin than to the glabella. The rim appears to
have been concave. Ocular ridges starting near the anterior end of
the glabella and directed slightly backward. Palpebral lobes about
half the glabellar length, set off by a shallow palpebral furrow. Dis-
tance from posterior end of palpebral lobe to posterior margin some-
what more than half the length of palpebral lobe. Posterior limbs
extending beyond the palpebral lobes for about two-thirds the distance
from the dorsal furrow to the palpebral lobe. Free cheeks with rim
about as wide as the ocular platform, extending into moderately long
genal spines.

Thorax of 16 segments. Axis occupying on the average one-fifth
of the width. Pleura with a broad furrow, whose anterior margin is
well marked, while posteriorly the furrow merges into the flat portion
of the pleuron. Anterior pleura wholly straight and bluntly terminated,
while the posterior ones have increasingly long falcate terminations ;
the last ones to some extent envelop the pygidium.

Pygidium triangular, about twice wider than long. Axis fairly
prominent, showing at least three segments, reaching the posterior
margin. Pleura with three pairs of shallow furrows; apparently con-
vex, without differentiated rim.

Surface smooth. Length of largest shield 43 mm.

This species is similar to Amecephalus piochensis (Walcott), from
which it differs chiefly in the somewhat longer palpebral lobes, lesser
number of thoracic segments, and proportionately larger pygidium.
It differs from A. cleora in the different relative midlengths of the
brim and rim.

Horizon and locality —Mount Whyte formation (Lake Agnes and
Yoho shale lentils; Wenkchemnia-Stephenaspis zone). Type locality
U.S.N.M. 35m, southwest of Lake Louise. Other localities W3b,
Plain of Six Glaciers, and Wok, Fossil Gully, Mount Stephen.

Types Holotype: U.S.N.M. No. 58363. Plesiotypes: U.S.N.M.
Nos. 116095-7.

AMECEPHALUS CLEORA (Walcott)
Plate 15, figures 12-20

Olenopsis cleora Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 74, pl. 13,
figs. 3, 3a, 1917.

Alokistocare cleora (Walcott), RESsER, Smithsonian Misc. Coll., vol. 93, No. 5,
P- 9, 1935.

?Olenopsis crito Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 75, pl. 11,
figs. 6, 6b, 1917.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 209

?Kochiella crito (Walcott), REssER, Smithsonian Misc. Coll., vol. 93, No. 5,
Pp. 39, 1935.

?Inglefieldia birdsalli Howrett, Wagner Free Inst. Sci. Bull., vol. 11, No. 4,
pl. 1, hg. 2, 1036:

This species, described by Walcott from the Mount Assiniboine
district, is exceedingly common and widespread in the Plagiura-K och-
aspis zone. Scores of excellent cranidia in limestone and a few
articulated shields in shale were collected. Olenopsis crito is based
on a few cranidia poorly preserved in sandstone, and as far as can be
ascertained from the existing material is identical with Olenopsis
cleora. The new material available allows a more complete description
of the species.

Cranidium as a whole with little convexity. Glabella tapered, trun-
cated in front, delimited by a shallow dorsal furrow. Glabellar fur-
rows exceedingly shallow. Occipital furrow shallow, occipital ring
short and simple. Frontal limb long, divided into rim and brim by
a very shallow, sometimes indistinct marginal furrow. The characters
of the rim vary to some extent with the size of the individual. In
small cranidia, of 3-5 mm. length, the marginal furrow is better de-
fined, and the rim itself is marked by a shallow transverse depression ;
the rim is more or less upturned with respect to the brim. In large in-
dividuals, the rim has a tendency to become more definitely concave
and the marginal furrow to become obliterated, the whole frontal limb
possessing a concave longitudinal profile in which no sharp change in
slope between brim and rim is appreciable. In every case, the mid-
length of the rim is somewhat greater than that of the brim. Fixed
cheeks flat, slightly upsloping, their maximum width equaling the
glabellar width. Ocular ridges very low. Palpebral lobes narrow,
somewhat elevated, about half the glabellar length, with strongest
curvature near the middle. Distance from posterior end of palpebral
lobe to posterior margin little more than half the length of palpebral
lobe. Posterior limbs almost parallel-sided, wider than the occipital
ring. Anterior branch of facial suture considerably divergent in front
of the eye, gently curving inward across the marginal furrow and rim,
the frontal portion marginal for a considerable distance. Posterior
branch almost straight outward, narrowly curving backward near its
distal end. Free cheeks flat, with genal spines of moderate length.

Thorax rapidly tapered in the posterior third, giving the entire
shield a subelliptic shape. Pleura broadly furrowed, straight in the
proximal portion; terminating in long, flat spines that in the poste-
rior segments are directed straight backward. The only shield in which

210 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the thorax appears complete has 17 segments. Pygidium small, poorly
preserved in the only specimen where it is present.

Surface smooth. Length of a complete shield 46 mm. Fragmentary
cranidia indicate that the species attains a somewhat larger size.

This species closely resembles A. agnesensis, from which it chiefly
differs in the relative midlengths of brim and rim and in possessing
17 instead of 16 thoracic segments.

Inglefieldia? birdsalli Howell was described from a single poorly
preserved shield collected from the talus at Sentinel Pass, between
Pinnacle Mountain and Mount Temple. The specimen is certainly an
Amecephalus but does not allow specific identification. Since Ameceph-
alus cleora is common in limestone and shale of the Plagiura-Koch-
aspis zone in that area, whereas strata carrying Amecephalus agnes-
ensis are not known to be present, the specimen is tentatively referred
to the former species.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Type locality U.S.N.M. 62w, Mount Assiniboine dis-
trict. Other localities U.S.N.M. 60e, Ptarmigan Peak; W3c, Plain of
Six Glaciers ; W4c-e, Ross Lake; W5c, Mount Bosworth ; W7c-d and
W7i, Kicking Horse Mine; Wof, Fossil Gully ; W2od, Eiffel Peak.

Types.—Holotype: U.S.N.M. No. 64396. Holotype and paratype
of Olenopsis crito: U.S.N.M. Nos. 64371-2. Plesiotypes: U.S.N.M.
Nos. 116142-3.

Genus CABORCELLA Lochman, 1948
Genotype: Caborcella arrojosensis Lochman.

CABORCELLA SKAPTA (Walcott)
Plate 13, figures 1-4

Ptychoparia skapta Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 95,

pl. 12, figs. 9, 9a, I9QI7.
Antagmus skapta (Walcott), Resser, Smithsonian Misc. Coll., vol. 95, No. 22,

Pp. 2, 1937.

This species was based on an incomplete cranidium, collected in the
Mount Assiniboine district. Specimens collected by the writer in
the limestones of the Mount Whyte formation at several localities
are identical with the type in all the characters that can be ascertained ;
hence they are assigned to the species. This procedure is supported
by the fact that the fauna at the type locality (U.S.N.M. loc. 62w)
includes other forms of the Plagiura zone, proving at least approxi-
mate equivalence of the respective horizons at Mount Assiniboine and

Ross Lake.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 2IiI

Walcott gave a satisfactory description of the type cranidium as
far as it is preserved. Some additional remarks are based on the new,
more complete cranidia. The fixed cheeks are rather strongly convex
and on the average slightly downsloping, thus placing the palpebral
lobes slightly below the level of the dorsal furrow. Palpebral lobes
about one-fourth the glabellar length, rather elevated. Distance from
posterior end of palpebral lobe to posterior margin somewhat greater
than length of palpebral lobe. Posterior limbs somewhat wider than
occipital ring. Posterior branch of facial suture directed almost
straight outward behind the eye, sharply turning backward in crossing
the broad distal portion of the marginal furrow. Anterior branch
appearing directed straight forward when seen from above, sharply
turning inward in crossing the marginal furrow, marginal for a long
distance in the median portion. Entire surface covered with fine
granules of one size.

This species seems referable to Caborcella rather than to any other
described genus. It closely agrees with the genotype in all essential
features, differing chiefly in the better-defined border and less-tapered
glabella. Another species that the writer would refer to Caborcella is
Poulsenia granosa Resser (1939b, p. 59, pl. 13, figs. 19-30), whose
cranidium differs from that of C. skapta in the straighter ocular
ridges, less convex fixed cheeks, coarser granules, and larger size.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Type locality U.S.N.M. 62w, Mount Assiniboine district.
Other localities W4c-d, Ross Lake; W5c, Mount Bosworth; W7c-e,
Kicking Horse Mine.

Types —Holotype: U.S.N.M. No. 64392. Plesiotypes: U.S.N.M.
No. 116113.

CABORCELLA RARA Rasetti, new species
Plate 15, figures 21-23

Known from two cranidia preserved in limestone.

Entire cranidium strongly convex. Glabella of moderate convex-
ity, strongly tapered in the posterior two-thirds, almost parallel-sided
in the anterior third, rounded in front. Three pairs of shallow glabel-
lar furrows; occipital furrow well impressed at the sides. Occipital
ring expanded mesially, imperfectly preserved in both specimens.
Brim convex and downsloping ; marginal furrow deep; rim convex
and strongly upturned, arched transversely. Fixed cheeks convex and
downsloping, about two-thirds the glabellar width at its midpoint.
Palpebral lobes small, elevated, at the level of the glabellar midpoint.

212 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Ocular ridges wide, not strongly prominent. Posterior limbs some-
what narrower than the occipital ring, deeply furrowed. Distance
from posterior end of palpebral lobe to posterior margin almost twice
the length of palpebral lobe. Anterior facial sutures slightly convex
outward, on the average slightly convergent from the eyes to the
marginal furrow. Posterior branch with curvature increasing from
the eye to the posterior margin.

Entire surface of cranidium densely covered with very fine gran-
ules; in addition, large scattered tubercles on the elevated portions.

Length of largest cranidium 7 mm.

This form seems referable to Caborcella, although it differs from
the genotype in the narrowness and more definitely downsloping
position of the fixed cheeks.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W7g, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116144. Paratype: U.S.N.M.
No. 116145.

Genus CHANCIA Walcott, 1924

Genotype: Chancia ebdome Walcott.

Walcott gave a good description of the generic characters of Chan-
cia. Forms agreeing with the genotype are common at several horizons
in the Middle Cambrian; however, some of the described species
that should be assigned to Chancia were placed by Resser in Elrathia,
Ehmaniella, or other genera with which they do not agree owing to
the great width of the fixed cheeks and the large number of thoracic
segments.

CHANCIA LATIGENA Rasetti, new species
Plate 21, figures 15-17

Known from several cranidia poorly preserved in shale or impure
limestone, and a specimen preserving a portion of the thorax.

Glabella rather strongly convex transversely, less convex longi-
tudinally, moderately tapered, fairly rounded in front. Third and
fourth pairs of glabellar furrows well impressed, second pair short
and faint. Occipital furrow deep at the sides, shallow mesially ; occipital
ring bearing a node. Brim slightly convex, downsloping. Rim some-
what more than half as long as the brim, slightly convex, tapered
laterally, set off by an almost straight marginal furrow. Fixed cheeks
moderately convex, on the average somewhat downsloping, slightly
wider than the glabella; palpebral lobes small, situated at the level
of the glabellar midpoint. Ocular ridges moderately elevated. Pos-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 213

terior limbs considerably wider than the occipital ring. Distance from
posterior end of palpebral lobe to posterior margin more than twice
length of palpebral lobe. Anterior facial sutures not divergent even
immediately in front of the eyes, then curving inward and converging,
producing widely rounded anterior angles of the cranidium. Posterior
branch directed outward and slightly backward behind the eye.

Surface of test densely covered with fine granules.

Length of the largest cranidium 14 mm., width between extremities
of the posterior limbs 32 mm.

This species is readily distinguished from the genotype, C. ebdome,
by the greater width of the fixed cheeks and the different proportions
of frontal brim and rim.

Horizon and locality.—Cathedral formation (Albertella( ?) zone
Locality Cgh, Fossil Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116184. Paratypes: U.S.N.M.
No. 116185.

CHANCIA BIGRANULOSA Rasetti, new species
Plate 22, figures 1-6

Known from numerous cranidia preserved in limestone but con-
siderably distorted.

Glabella moderately tapered, fairly convex in both directions, some-
what truncated in front. Glabellar furrows very shallow ; however,
in some specimens four pairs are distinguishable under proper light-
ing. Occipital furrow well impressed throughout ; occipital ring rather
short, bearing a node. Frontal limb consisting of brim and rim; in
most specimens two transverse furrows are visible on the limb. This
raises the question, which one should be considered as the marginal
furrow. The writer believes that in similar cases (e.g., in Kochiella;
see discussion of that genus) the furrow nearer the glabella is the
marginal furrow, the other being a furrow across the rim proper. In
the present species the brim is slightly convex and downsloping ; the
rim is convex near the cranidial margin, but on the whole has a some-
what concave profile because of the transverse impression. The mid-
length of the rim is greater than that of the brim. The rim tapers
rapidly at the sides. Fixed cheeks convex, on the average slightly down-
sloping, almost as wide as the glabella. Palpebral lobes small, elevated,
situated at the level of the glabellar midpoint; ocular ridges strong.
Posterior limbs considerably wider than the occipital ring. Distance
from posterior end of palpebral lobe to posterior margin about one
and a half times the length of palpebral lobe. Anterior branch of facial

214 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

suture almost straight from the eye to the marginal furrow, directed
slightly outward; anterior angles of cranidium narrowly rounded.
Posterior branch reaching the posterior margin with almost uniform
curvature.

Surface of entire cranidium densely covered with small granules,
among which are scattered tubercles of a much larger size.

Length of largest cranidium 13 mm.

This species differs from all others assigned to the genus in the
character of the frontal limb and the type of surface ornamentation.
In several features this form is intermediate between Piazella and
Chancia, but seems better placed in the latter genus.

Horizon and locality——Cathedral formation (Albertella(?) zone).
Type locality Coj’, Fossil Gully, Mount Stephen: also locality Cgj,
Fossil Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116186. Paratypes: U.S.N.M.
No. 116187.

CHANCIA PALLISERI (Walcott)
Plate 29, figures 1-3

Ptychoparia palliseri Watcott, Canadian Alpine Journ., vol. 1, No. 2, p. 244,
pl. 3, fig. 6, 1908.

Elrathia palliseri (Walcott), REssEr, Smithsonian Misc. Coll., vol. 95, No. 22,
p. 10, 1937.

No description of this form has ever been given. Hence it is
considered opportune to give a complete description accompanied by a
discussion of the generic position.

Known from numerous shields flattened in shale.

Glabella moderately tapered, rounded in front. Two pairs of gla-
bellar furrows rather well impressed. Convexity of glabella cannot
be determined from the flattened specimens. Frontal limb about two-
thirds as long as the glabella, consisting mostly of brim; rim short,
apparently somewhat upturned. Fixed cheeks appearing slightly nar-
rower than the glabella in flattened specimens ; however, if the gla-
bella had preserved its convexity, it would probably not appear wider
than the fixed cheeks. Ocular ridges moderately elevated ; palpebral
lobes small, situated slightly in advance of the glabellar midpoint.
Distance between posterior end of palpebral lobe and posterior margin
about twice length of palpebral lobe. Posterior limbs wider than
occipital ring. Anterior facial sutures fairly divergent, straight from
the eye toa short distance from the marginal furrow, then curving in-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 215

ward; anterior angles of cranidium not widely rounded. Posterior
branch straight for a considerable portion, then curving backward.

Thorax long, with a moderately tapered axis. Pleura straight for
a considerable distance, curving backward into short spines distally ;
these spinose endings are more pronounced in the median portion of
the thorax. Pleural furrow well impressed over the entire length of
each pleuron.

The number of thoracic segments appears variable in this species,
as in some other cases among multisegmented ptychoparids (see de-
scription of Elrathina cordillerae). Walcott figured the holotype as
possessing 23 thoracic segments. Of the three complete specimens
from the type locality (U.S.N.M. locality 14s, the Ogygopsis shale
bed on Mount Stephen) preserved in the United States National Mu-
seum, two (including the holotype) have 23 segments and one has
only 20 segments. Several complete specimens from the Burgess shale
(locality 35k) have 20 segments. The writer collected a good im-
pression of a complete specimen from the type locality, and this shows
21 segments. These various specimens do not appreciably differ in
size, hence the differences in the number of thoracic segments are
certainly not due to growth stages. As in the case of Elrathina cordil-
lerae, it must be concluded that the number of thoracic segments in
certain species is not absolutely constant. No such case was ever
observed by the writer among bathyuriscid trilobites, where the
number of thoracic segments does not exceed a dozen.

Pygidium small, transverse. Axis occupying less than one-third
of the width and most of the length, distinctly segmented. Pleural
lobes with three or four distinct pairs of furrows and three pairs of
interpleural grooves.

Surface apparently smooth except for the usual longitudinal anas-
tomosing lines on the brim.

The proportions of a complete shield with 20 thoracic segments are
the following: Total length 35 mm.; length of cephalon 11 mm.;
length of thorax 21 mm.; length of pygidium 3 mm.

This species appears to agree in all essential generic features with
Chancia, differing from the genotype chiefly in the different propor-
tions of brim and rim. It disagrees with Elrathia in the great width
of the fixed cheeks, smaller palpebral lobes, great number of thoracic
segments, and small pygidium.

Horizon and locality—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s, Mount
Stephen, identical with the writer’s locality S8d. Also Burgess shale
lentil ; locality U.S.N.M. 35k, Mount Field.

216 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Types.—Holotype: U.S.N.M. No. 94344. Plesiotypes: U.S.N.M.
No. 116236.

CHANCIA ODARAYENSIS Rasetti, new species
Plate 33, figures 15, 16

Known from a few cranidia and two specimens with most of the
thorax, all poorly preserved in limestone.

Glabella moderately tapered, fairly convex in both directions, not
sharply truncated in front. Glabellar furrows shallow. Occipital
ring apparently bearing a node. Brim convex, downsloping. Rim
separated from the brim by a shallow marginal furrow, on the average
horizontal, slightly concave. Midlength of rim somewhat greater
than midlength of brim. Fixed cheeks convex, on the average hori-
zontal, as wide as the glabella. Ocular ridges fairly prominent,
straight, directed slightly backward. Palpebral lobes small, semicircu-
lar, not greatly elevated. Posterior limbs considerably wider than the
occipital ring, deeply furrowed. Distance from posterior end of pal-
pebral lobe to posterior margin about twice length of palpebral lobe.
Anterior facial sutures directed forward for a short distance in front
of the eyes, then turning inward with a wide curve; anterior angles
of cranidium well rounded. Posterior branch almost straight for most
of the length, narrowly rounding off the ends of the posterior limbs.

A possibly complete thorax shows 22 segments. Pygidium small,
triangular, transverse, with a prominent axis reaching the posterior
margin and strongly furrowed pleural lobes.

Surface of test finely granulated.

Length of the largest cranidium 7 mm., of cephalon and thorax
together 25 mm.

This species differs from the others described chiefly in the propor-
tions of the fixed cheeks and palpebral lobes and in the structure
of the frontal limb.

Horizon and locality—Stephen formation (Bathyuriscus-Elra-
thina zone). Type locality S61, Mount Odaray. Specimens collected
at locality Sok, Mount Stephen, doubtfully belong to the species.

Types——Holotype: U.S.N.M. No. 116274. Paratypes: U.S.N.M.
Nos. 116275-6.

CHANCIA STENOMETOPA Rasetti, new species

Plate 8, figure 20

Known from several cranidia slightly flattened in shale, and a
specimen preserving a portion of the thorax.

EE ———E—

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 217

Glabella proportionately long and narrow, moderately tapered,
strongly convex transversely, with well-defined anterior angles,
rounded in front. Glabellar furrows well impressed, decreasing in
depth from the fourth to the second pair; first pair indistinct. Oc-
cipital furrow deep; occipital ring short, apparently simple. Brim about
half the glabellar length, convex, downsloping ; rim short, convex, set
off by a wide, fairly deep marginal furrow. Fixed cheeks as wide as
the glabella, rather flat, probably horizontal. Ocular ridges curved,
well marked. Palpebral lobes small, semicircular, elevated. Posterior
limbs considerably wider than the occipital ring. Distance from
posterior end of palpebral lobe to posterior margin about twice length
of palpebral lobe. Anterior branch of facial suture convex outward,
on the average directed forward from the eye to the marginal furrow,
then curving inward rather sharply across the rim. Posterior branch
rather straight for most of the distance.

An incomplete thorax shows approximately 15 segments. Pleura
deeply furrowed.

Surface of cranidium densely covered with very fine granules, be-
sides a much smaller number of larger granules; the frontal brim
shows, in addition, faint longitudinal ridges.

Length of the largest cranidium 12 mm.

This species agrees in the essential features with the generic diagno-
sis of Chancia, although the narrowness of the glabella gives the
cranidium an unusual aspect. It is also to be noted that it is a much
earlier form than all others assigned to the genus.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
Wenkchemnia-Stephenaspis zone). Locality Wok, Fossil Gully,
Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116078. Paratypes: U.S.N.M.
No. 116079.

Genus EHMANIELLA Resser, 1937

Genotype: Crepicephalus (Loganellus) quadrans Hall and Whit-
field.

EHMANIELLA BURGESSENSIS Rasetti, new species
Plate 30, figures 9-16

Ptychoparia permulta Watcotr (part), Smithsonian Misc. Coll., vol. 67, No. 4,

p. 145, pl. 21, fig. 1 (only), 1918.
Elrathia permulta (Walcott), REsser, Smithsonian Misc. Coll., vol. 95, No. 22,

p. 10, 1937.

218 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II16

Known from several complete shields and numerous cranidia and
pygidia, preserved in siliceous shale and more or less flattened ; besides
a few cranidia and pygidia preserved in limestone.

Glabella moderately tapered, straight-sided, fairly rounded in front,
convex in both directions, slightly keeled. First pair of glabellar
furrows represented by short, shallow pits; following three pairs
longer and rather well impressed. Occipital furrow straight, well
impressed throughout. Occipital ring somewhat expanded mesially,
bearing a node. Frontal limb slightly convex, downsloping, divided
by the marginal furrow into brim and rim; midlength of brim about
twice midlength of rim. Marginal furrow well impressed, and fairly
straight, rim convex, tapered at the sides. Fixed cheeks convex,
slightly downsloping, about two-thirds the glabellar width. Ocular
ridges strong, fairly straight, moderately oblique. Palpebral lobes
about one-fourth the glabellar length, somewhat elevated, narrow ;
proportionately longer in young individuals. Distance from posterior
end of palpebral lobe to posterior margin about twice length of pal-
pebral lobe. Anterior branch of facial suture divergent in front of the
eye, almost straight to the anterior margin; anterior angles of cranid-
ium narrowly rounded. Posterior branch directed outward and
backward without much curvature. Free cheeks with short genal
spines.

Thorax of 13 segments. Axis slightly tapered. Pleura almost
straight in dorsal view, deeply furrowed, with poorly defined genicu-
lation; terminating in fairly sharp but short spines.

Pygidium twice wider than long, subtriangular. Axis moderately
tapered, elevated, showing four or five rings and a terminal section ;
occupying about five-sixths of the pygidial length. Pleural lobes
downsloping ; first pleuron less fused than the others, delimited by a
well-impressed interpleural groove; extended at the sides into a pair
of short, sharp, backward-directed marginal spines. Other inter-
pleural grooves less distinct ; five pairs of pleural furrows impressed.
Marginal furrow distinct, rim narrow.

Surface of cranidium covered with very fine granules. Brim and
free cheeks with the usual anastomosing lines.

Length of largest complete shield 27.5 mm., of which 9 mm. belong
to the cephalon, 15 mm. to the thorax, and 3.5 mm. to the pygidium.

This species closely resembles the genotype, E. quadrans (Hall and
Whitfield). An accurate comparison with the types of quadrans is
impossible because the latter are very poorly preserved. Better-pre-
served material from Utah in the United States National Museum

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 219

collections may be assumed to represent the genotype species, and
affords a more significant comparison. E. burgessensis appears to
differ from quadrans chiefly in the presence of small marginal pygidial
spines, slightly shorter palpebral lobes, and lesser number of distinct
interpleural grooves in the pygidium.

E. burgessensis also closely resembles £. basilica Resser from the
Grand Canyon, differing mainly in the straighter anterior marginal
furrow, straighter ocular ridges, and presence of marginal pygidial
spines.

Horizon and locality.—Stephen formation (Burgess Shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 35k, Burgess
Quarry, Mount Field. The species is much more common at locality
Siig, Mount Field. Several specimens were collected from limestone
at locality S6g, Mount Odaray.

Types—Holotype: U.S.N.M. No. 116245. Paratypes: U.S.N.M.
No. 65516; Nos. 116246-8.

EHMANIELLA WAPTAENSIS Rasetti, new species
Plate 30, figures 6-8

Known from numerous shields partly flattened in shale. Entire
shield ovate.

Glabella tapered at the usual rate, straight-sided, fairly rounded in
front. Three pairs of shallow glabellar furrows. Occipital furrow
deep and straight; occipital ring rather short, possibly bearing a
small node. Brim downsloping; rim convex, almost unifomly wide
around the cephalon, defined by a deep marginal furrow ; midlengths of
brim and rim equal. Fixed cheeks somewhat less than two-thirds the
glabellar width. Ocular ridges fairly prominent. Palpebral lobes about
one-fourth the glabellar length; distance from posterior end of pal-
pebral lobe to posterior margin slightly greater than length of palpe-
bral lobe. Facial sutures as in other species of the genus. Free cheeks
with very short genal spines.

Thorax of 13 segments. Axis somewhat tapered. Pleura deeply
furrowed ; geniculation somewhat nearer to the dorsal furrow than to
the end of the pleura. Distal end of pleura rounded.

Pygidium relatively small, more than twice wider than long. Axis
tapered, not greatly elevated, with two or three very shallow furrows.
Pleural lobes with one or two pairs of pleural furrows and inter-
pleural grooves distinct.

220 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Surface of test covered with granules, larger on the glabella than
on the other portions of the shield.

Length of the largest cranidium 4.5 mm. Length of an entire shield
9.0 mm., of which 3.4 mm. belong to the cephalon, 4.6 mm. to the
thorax, and 1.0 mm. to the pygidium.

This species differs from the genotype and other forms hitherto as-
signed to Ehmamiella in the longer palpebral lobes and lesser relative
midlength of the brim. It is more similar, as far as the cranidial fea-
tures are concerned, to species assigned to Clappaspis, a genus that the
writer would include in Ehmaniella. The present species appears to
differ from the forms described by Deiss and Resser as Clappaspis in
the different surface ornamentation and longer palpebral lobes.

Horizon and locality——Stephen formation (Burgess shale lentil;
Bathyuriscus-Elrathina zone). Locality U.S.N.M. 35k, Burgess
Quarry, Mount Field.

Types.—Holotype: U.S.N.M. No. 116243. Paratypes: U.S.N.M.
No. 116244.

Genus ELRATHIA Walcott, 1924
Genotype: Conocoryphe (Conocephalites) kingii Meek.

ELRATHIA PERMULTA (Walcott)
Plate 30, figures 1, 2

Ptychoparia permulta Watcotr (part), Smithsonian Misc. Coll., vol. 67, No. 4,
p. 145, pl. 21, fig. 2 (only), 1918.
Elrathia permulta (Walcott), REssEr, Smithsonian Misc. Coll., vol. 95, No. 22,
p. 10, 1937.
Elrathia dubia RessEr, Smithsonian Misc. Coll., vol. 95, No. 22, p. 11, 1937.
Resser correctly recognized that the two individuals illustrated by
Walcott as Ptychoparia permulta belong to distinct species. How-
ever, in making a new species, Elrathia dubia, he designated as holo-
type the same specimen that Walcott had unmistakably indicated as
the holotype of permulta. Hence dubia becomes an objective synonym
of permulta, and the other form lacks a name. The latter species is
a typical Ehmaniella and is described in this paper as E. burgessensis.
Horizon and locality—Stephen formation (Burgess shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 35k (=au-
thor’s locality Stif); Burgess Quarry, Mount Field.
Types——Holotype: U.S.N.M. No. 65517. Plesiotypes: U.S.N.M.
No. 116240.

—

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 2

bo
_

Genus ELRATHINA Resser, 1937

Genotype: Conocephalites cordillerae Rominger.

Forms belonging to this genus are abundant in most of the strata
of the Bathyuriscus-Elrathina zone. The writer found considerable
difficulty in recognizing clear-cut species within the available material.
The specific characters, if any, consist in small differences in the pro-
portions of the cranidial parts, and are often masked by individual
variability or by flattening, distortion, and other effects of imperfect
preservation. It is especially difficult to make an accurate comparison
between specimens preserved in shale and limestone. Hence the
validity of the species described below is presented as somewhat
doubtful.

One fact that emerged from a close study of species of Elrathina
is that the number of thoracic segments within a single species appears
to be variable within narrow limits, instead of being absolutely con-
stant as assumed by most modern authors. Apparently a variable
number of thoracic segments is observed only among multisegmented,
micropygous ptychoparids.

ELRATHINA CORDILLERAE (Rominger)
Plate 26, figures 7-9

Conocephalites cordillerae RoMINGER, Proc. Acad. Nat. Sci. Philadelphia, 1887,
Dt7, pl. 1h. 7:

Ptychoparia cordillerae (Rominger), Watcott, Amer. Journ. Sci., ser. 3, vol.
36, p. 165, 1888; Canadian Alpine Journ., vol. 1, No. 2, pl. 3, fig. 5, 1908;
Smithsonian Misc. Coll., vol. 67, No. 4, p. 144, pl. 21, fig. 4 (3 and 5?), 1918.

Ptychoparia cordillerae (Rominger), MatrHew, Trans. Roy. Soc. Canada,
ser. 2, vol. 5, sect. 4, p. 44, pl. 1, fig. 7, 1800.

Elrathina cordillerae (Rominger), ReEsser, Smithsonian Misc. Coll., vol. 95,
No..22;..6.. II, 1937:

The number of thoracic segments in this species varies between 17
and 19, 18 being the most common number and 19 the rarest. These
differences are certainly not due to growth stages as they occur in
individuals of the same size; they are present at both localities where
the species was collected.

Part of the difference in the number of thoracic segments is due
to the last thoracic segment being sometimes included in the pygidium,
which then appears to consist of two segments instead of one. How-
ever, individuals with the smaller pygidium still may have 18 or 19
thoracic segments, while individuals with the larger pygidium may
have 17 or 18 thoracic segments. This shows that the variability in

222 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the number of thoracic segments is not entirely due to anchylosis of
a normally free thoracic segment with the pygidium.

Horizon and locality—Stephen formation (Ogygopsis shale lentil ;
Bathyuriscus-Elrathina zone). Type locality U.S.N.M. 14s (=au-
thor’s locality S8d), Mount Stephen. Also locality U.S.N.M. 35k,
Burgess Quarry, Mount Field.

Types.—Syntypes: Acad. Nat. Sci. Philadelphia. Plesiotypes:
U.S.N.M. Nos. 65518, 57658, 17831, 116225-6.

ELRATHINA PARALLELA Rasetti, new species
Plate 33, figures 19-22

Known from numerous cranidia and complete specimens, all pre-
served in limestone.

Glabella almost parallel-sided, narrow, rounded in front. Glabellar
furrows indistinct. Occipital furrow shallow; occipital ring short,
bearing a small node. Frontal limb as a whole downsloping, about
equally divided into brim and rim mesially ; marginal furrow shallow ;
slopes of rim and brim almost equal. Fixed cheeks convex, on the
average downsloping; their maximum width approximately equaling
the glabellar width. Palpebral lobes narrow, somewhat less than one-
third the glabellar length ; their centers situated somewhat in advance
of the glabellar midpoint but not quite on a transverse line through
the anterior third of the glabella. Ocular ridges very faint, slightly
curved. Distance from posterior end of palpebral lobe to posterior
margin about twice length of palpebral lobe. Posterior limbs consider-
ably wider than the occipital ring, well rounded distally.

Thorax of 17 or 18 segments in the specimens where it is sufficiently
well preserved for the segments to be counted. Pleura straight in
dorsal view, moderately bent down at geniculation which occurs on
the average somewhat beyond the midpoint of each pleuron. Ex-
tremity of pleura rounded.

Pygidium small, transverse, with one distinct pair of pleural fur-
rows.

Surface of test smooth.

Length of an apparently full-grown shield 21.2 mm., of which 7
mm. belong to the head, 13 to the thorax, and 1.2 to the tail.

This species is very similar to the genotype, E. cordillerae, and a
very accurate comparison between the two forms is difficult owing to
the different manner of preservation. The new species differs in
having the geniculation of the thoracic pleura closer to the middle than
to the distal end of each pleuron, and in possessing a less triangular
pygidium with a relatively narrower axis. Other possible differences

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 223

are the proportionately narrower and longer glabella and the slightly
longer and more posteriorly situated palpebral lobes.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality Sgk, Mount Stephen. Also locality S6k, Mount
Odaray.

Types.—Holotype: U.S.N.M. No. 116279. Paratypes: U.S.N.M.
No. 116280.

ELRATHINA BREVIFRONS Rasetti, new species
Plate 26, figure 6

Known from numerous shields partly flattened in argillaceous lime-
stone.

Owing to the close resemblance of this species to the genotype,
E. cordillerae, it will be sufficient to point out the differential charac-
ters instead of giving a complete description.

The cranidium differs from that of E. cordillerae in the reduction of
the midlength of the brim; the dorsal and marginal furrows almost
meet mesially.

The geniculation of the thoracic segments is situated near the mid-
dle of each pleuron instead of near the distal end. The pleura termi-
nate in short spines instead of being rounded as in E. cordillerae. Most
of the specimens have 18 thoracic segments; some have 19. The py-
gidium is subelliptical instead of subtriangular as in E. cordillerae.

The present species also closely resembles E. fecunda Deiss, differ-
ing chiefly in the lesser midlength of the brim. Another closely related
species is Elrathia spencei Resser, which is fully typical of Elrathina
in all characters except the one, which does not appear of generic im-
portance, of having genal spines, whereas in £. cordillerae and the
present species the genal angle is rounded. EF. spencei further has a
more convex cranidium and a proportionately narrower and longer
pygidium.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S11e, Mount Field.

Types.—Holotype: U.S.N.M. No. 116223. Paratypes: U.S.N.M.
No. 116224.

ELRATHINA SPINIFERA Rasetti, new species

Plate 34, figures 11-14

Known from numerous cranidia preserved in limestone.
Cranidium closely similar in all its proportions to that of E. cordil-
lerae, possibly excepting a somewhat greater midlength of the rim.

224. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 26

The chief distinctive character is the presence of a short, backward-
directed occipital spine.

Surface of test smooth. Length of largest cranidium 7 mm.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Type locality S6k, Mount Odaray. Also locality Sok, Mount
Stephen.

Types—Holotype: U.S.N.M. No. 116289. Paratypes: U.S.N.M.
Nos. 116290-1.

ELRATHINA MARGINALIS Rasetti, new species
Plate 34, figures 8-10, 15

Known from numerous cranidia and a specimen with part of the
thorax, all preserved in limestone.

The cranidium has the same general appearance as in E. cordillerae,
hence is described by pointing out the differential characters, consist-
ing in the greater general convexity, strong elevation of the glabella,
and greater depth of the marginal furrow, which defines a more con-
vex and elevated rim than in the other species of the genus here dis-
cussed. The occipital ring bears a node. The geniculation of the
thoracic pleura occurs at about two-thirds the distance from the proxi-
mal end. The pleura do not terminate into spines.

Surface of test smooth. Length of largest cranidium 6 mm.

Horizon and locality —Stephen formation (Bathyuriscus-Elrathina
zone). Locality S61, Mount Odaray.

Types —Holotype: U.S.N.M. No. 116287. Paratypes: U.S.N.M.
No. 116288.

Genus GLYPHASPIS Poulsen, 1927
Genotype: Asaphiscus capella Walcott.

GLYPHASPIS PARKENSIS Rasetti, new species
Plate 34, figures 5-7

Known from numerous cranidia and pygidia and an entire shield,
all preserved in limestone but more or less flattened and distorted.

Glabella weakly convex, tapered, truncated in front. Glabellar
furrows very faint; occipital furrow shallow but impressed through-
out, occipital ring short, bearing a small node. Frontal limb concave,
poorly separated into a brim and rim of about equal midlengths by
a wide, shallow marginal furrow. Total midlength of frontal limb
almost equal to glabellar length. Fixed cheeks about two-thirds the

ee

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 225

glabellar width. Palpebral lobes more than half as long as the glabella,
moderately curved, narrow, set off by a well-impressed palpebral
furrow. Ocular ridges strong. Anterior facial sutures considerably
divergent and almost straight for a considerable distance; anterior
angles of cranidium narrowly rounded. Posterior branch almost par-
allel to posterior margin, defining slender, deeply furrowed posterior
limbs.

Thorax of Io segments. Axis not tapered. Pleural furrows in-
creasing in depth from the proximal end of the pleura to the genicula-
tion.

Pygidium somewhat less than twice wider than long, as wide as the
cephalon. Axis reaching about three-fourths the length of the py-
gidium, tapered, showing approximately six rings and a terminal
section; no axial nodes or spines. Pleural platforms convex, rim
wide, concave. Six or seven pairs of increasingly shallow pleural
furrows, separated by shallower and less distinct interpleural grooves.

Surface of test smooth except for anastomosing lines on the brim
and free cheeks.

Length of the largest cranidium 20 mm. Cephalon, thorax, and
pygidium each of approximately equal length.

This species resembles two forms described by Deiss from Mon-
tana. Glyphaspis paucisulcata Deiss, known from the pygidium alone,
differs from the present species in the much more widely rounded
anterior angles. The other species, G. storeyi, is extremely similar
to G. parkensis in the pygidial features, but the cranidium of G.
storeyi appears to differ in the longer palpebral lobes and narrower
fixed cheeks. The two forms are doubtless very closely related.

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Locality Stol, Park Mountain.

Types.—Holotype: U.S.N.M. No. 116285. Paratypes: U.S.N.M.
No. 116286.

Genus KOCHASPIS Resser, 1935

Genotype: Crepicephalus liliana Walcott.

It appears reasonably certain that Crepicephalus liliana was based
on the correct combination of cranidium and pygidium. Slabs of
sandstone from the Pioche shale (Comet shale of Deiss) of the
Highland Range, Nevada, preserved in the United States National
Museum, are covered with cranidia and pygidia of this species, the
only associated species being Oryctocephalus primus. Of the three
species from the Mount Whyte formation assigned to Kochaspis, K.
celer and K. chares were apparently based on illegitimate combinations

226 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

of a ptychoparid head and a bathyuriscid tail (see descriptions of
Fieldaspis superba and Kochiella? maxeyi). The writer inclines to
believe that the third species, K. cecinna, is a good species of Koch-
aspis founded on correct association of the two shields. The writer
was unable to collect K. cecinna, but found a new species of the
genus at several localities in the Plagiura-Kochaspis zone. Owing to
the rarity of this trilobite, there is no certainty that the assignment
of cranidium and pygidium to one species is correct, but it appears
at least probable.

KOCHASPIS EIFFELENSIS Rasetti, new species
Plate 14, figures 4-10

Known from numerous cranidia and a few pygidia preserved in
limestone.

Glabella rather strongly convex transversely, moderately convex
longitudinally, moderately tapered, straight-sided, rather truncated
in front. Four pairs of glabellar furrows impressed ; first pair very
short, second and third pairs longer, fourth pair still longer and show-
ing a tendency to bifurcate at the inner end. Occipital furrow well
impressed, shallower in the middle. Occipital ring short, bearing a
node. Frontal brim slightly convex, downsloping. Rim almost flat,
differentiated from the brim by its upturned position; in young indi-
viduals the rim is appreciably convex. Midlengths of brim and rim
almost equal; midlength of entire frontal limb about half the glabellar
length. Fixed cheeks flat, very slightly downsloping ; maximum width
almost equaling the glabellar width. Ocular ridges well developed,
straight, strongly oblique. Palpebral lobes curved, slightly elevated,
set off by a shallow palpebral furrow; their length about one-third the
glabellar length. Posterior limbs incompletely preserved in available
specimens.

Pygidium subrectangular, about twice wider than long, with pleural
lobes extended into a pair of strong, long spines. Axis cylindrical,
strongly elevated, composed of two rings and a terminal section, not
quite reaching the posterior margin. Pleural lobes convex, down-
sloping. Two pairs of pleural furrows impressed ; these become deeper
in crossing the marginal furrow (which is otherwise obsolete) and
extend for some distance on the base of the spines. Spines strong,
directed backward and somewhat outward, of unknown total length
as they are broken off in the few available pygidia.

Surface of cranidium and pygidium densely covered with fine gran-
ules of one size.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 227

Length of largest cranidium 14 mm. Length of pygidium 5 mm.,
width Io mm.

This species closely resembles K. cecinna (Walcott) from which
it differs chiefly in the flatter and wider anterior rim, lesser number
of pleural furrows in the pygidium, and lack of large granules on the
surface of the test.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Type locality W2od (=U.S.N.M. 38k), Eiffel Peak.
Other localities W6d, Mount Odaray; W7f, Kicking Horse Mine;
Wofi, Fossil Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116125. Paratypes: U.S.N.M.
Nos. 116126-8. The holotype is a cranidium, hence the name will
remain attached to the cranidium if the assignment of cranidium and
pygidium to one species should prove to be erroneous.

Genus KOCHIELLA Poulsen, 1927

Genotype: Kochiella tuberculata Poulsen.

The genotype and several other species were described by Poulsen
from the Lower Cambrian Cape Kent formation of northwest Green-
land. Poulsen did not assign a pygidium to any of his species, but
described from the same beds a spinose pygidium as Crepicephalus
cf. cecinna Walcott. Resser (1935, p. 39) assigned this pygidium to
K. tuberculata. Following this assignment, Resser described several
Middle Cambrian species of Kochiella as combinations of an Ameceph-
alus-like cephalon and a spinose pygidium. At least in certain in-
stances, e.g., Kochiella mansfieldi (Resser 1939b, p. 57), the associa-
tion appears warranted by the existing material. However, there is
no certainty that the assignment of the pygidium of K. tuberculata
is correct, hence the generic characters of the pygidium remain un-
determined, and it is difficult to decide what forms should be included
in Kochiella.

A cranidium occurring at several localities in the Plagiwra-Koch-
aspis zone agrees generically with Kochiella tuberculata, but since the
pygidia of either form are unknown, the generic assignment of the
species from the Mount Whyte formation is doubtful. A spinose py-
gidium occurs rarely in the same beds, and the cranidia to which it
could be assigned are either the one being discussed or that of Koch-
aspis eiffelensis. The second solution appears the most likely to be
correct and was adopted. There remains no pygidium assignable to
the new form, which makes it plausible that this part of the shield was
small and featureless as in Amecephalus. If this were proved to be

228 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL; 516

the case, possibly Amecephalus would be the proper generic assign-
ment for the form under discussion.

It should be mentioned in this connection that whenever Resser de-
scribed species of Kochiella or similar cranidia he repeatedly referred
to a transverse line across the brim being the “impression of doublure
on upper surface” (Resser, 1935, pp. 8, 38; 1939b, pp. 51, 57). Not
only does this line appear in specimens in limestone where there is no
trace of flattening, but Resser described this feature in cranidia lack-
ing the free cheeks, where obviously the doublure could not be im-
pressed on the dorsal shield because it was not present when the speci-
men was buried in the sediment. The line is simply the marginal
furrow, and its closeness to the glabella in certain cases indicates a
considerable midlength of the rim.

KOCHIELLA? MAXEYI Rasetti, new species
Plate 13, figures 5-8

Known from numerous cranidia preserved in limestone. Glabella
fairly convex, its cross section being of inverted V-shape; strongly
tapered, with slightly concave sides, somewhat truncated in front.
Second, third, and fourth pairs of glabellar furrows well impressed
at the sides; first pair very short but distinct. Occipital ring set off
by a well-impressed furrow, short, bearing a node. Frontal limb divided
by a narrow and shallow, but distinct, marginal furrow into a slightly
convex brim and a concave rim of equal midlengths ; total midlength
of limb almost equaling the glabellar length. Fixed cheeks flat, up-
sloping. Ocular ridges slightly curved, strongly elevated. Palpebral
lobes narrow, somewhat elevated, sharply curved in the middle, almost
half the glabellar length ; width of fixed cheeks about equal to glabellar
width. Posterior limbs wider than the occipital ring, short, parallel-
sided. Anterior sutures somewhat divergent in front of the eyes,
curving inward across the rim. Posterior branch directed almost
straight outward.

Surface of cranidium densely covered with very fine granules,
among which are scattered granules of a much larger size; brim with
irregular, longitudinal ridges.

Length of largest cranidium 18 mm.

The cranidium of this species is very similar to that described by
Walcott as Crepicephalus chares. However, the pygidium assigned
by Walcott to the species is, in the writer’s opinion, the pygidium of
a bathyuriscid trilobite of the genus Fieldaspis. No pygidia assignable
to the cranidium here described were collected by the writer. Hence,

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 229

pending decision as to whether the name chares should apply to the
cranidium or the pygidium (no holotype was designated either by
Walcott or by subsequent authors) the writer believes that confusion
will be avoided by describing this form under a new name rather
than tentatively identifying it with the cranidium of Crepicephalus
chares. The reasons for the tentative assignment of the species to
Kochiella were mentioned in the discussion of the genus.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Type locality W2od, Eiffel Peak. Other locality Wof,
Fossil Gully, Mount Stephen.

Types——Holotype: U.S.N.M. No. 116114. Paratypes: U.S.N.M.
Nos. 116115-6.

KOCHIELLA? cf. K. MAXEYI Rasetti
Plate 13, figure 9

A cranidium collected from the Mount Whyte formation on Mount
Field differs from the types of the species in the greater relative mid-
length of the anterior rim. Such a small difference does not warrant
naming a new species unless it should prove a constant feature of the
specimens at that horizon and locality.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W7f, Kicking Horse Mine, Mount Field.

Figured specimen—vU.S.N.M. No. 116117.

Genus KOCHINA Resser, 1935
Genotype: Olenopsis americanus Walcott.

KOCHINA AMERICANA (Walcott)
Plate 19, figures 20-23

Olenopsis americanus Watcott, Smithsonian Misc. Coll., vol. 57, No. 8, p. 243,
pl. 36, figs. 8-11, 1912.

Olenopsis cf. americanus Watcortt, Smithsonian Misc. Coll., vol. 67, No. 2, p. 37,
pl. 6, figs. 8-8b, 1917.

Kochina americana (Walcott), RESSER, Smithsonian Misc. Coll., vol. 93, No. 5,
Pp. 40, 1935.

Kochina bosworthensis RessEr, Smithsonian Misc. Coll., vol. 93, No. 5, p. 40,
1935.

The types of this species were described from the Gordon shale
of Montana. Later Walcott stated that specimens from the Ross
Lake shale were apparently identical with those from the type locality.

230 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Resser separated the form from the Ross Lake shale under a new
name, Kochina bosworthensis, without stating the characters that
differentiate the new species. Study of the types shows that, as far
as can be ascertained from the existing material, the two forms are
identical.

It is possible to supplement Walcott’s description of the species,
thanks to the discovery of additional material. Cranidia preserved in
the limestone interstratified with the Ross Lake shale show the con-
vexity, which has been lost to a large extent in the specimens in shale.
The convex fixed cheeks rise from the dorsal furrow almost to the
level of the glabella. The frontal limb is turned down rather strongly
at the sides, and the rim shows a moderate convexity. The reproduced
photographs show these characters clearly.

The impression of a complete shield in the Ross Lake shale, dis-
covered by the writer, shows for the first time the size and shape of
the pygidium. The thorax, apparently complete although some of the
segments are slightly displaced, shows 17 segments. The pygidium
is approximately semicircular, more than twice wider than long. The
axis occupies somewhat more than half the length, and about one-
fourth of the width; it is slightly tapered and consists of two distinct
segments plus a terminal section. The pleural lobes show three or
four pairs of shallow furrows and fainter interpleural grooves. The
proportions of the parts of the shield are given by the following
measurements. Cranidium: length Io mm., width 28 mm.; thorax:
length 22 mm.; pygidium: length 5 mm., width 11 mm,

Surface of cranidium densely covered with small granules.

Resser compared the genus with Kochiella. lf the pygidium of
Kochiella is large and spinose like that of Kochaspis (as believed by
Resser), then Kochina with its small, simple pygidium is not close
to the above-mentioned genera. Even the cranidia show only a
moderate resemblance.

Horizon and locality—The types are from the Gordon shale of
Montana. In the area the species occurs in the Cathedral formation
(Ross Lake shale member ; Albertella zone). Localities C3m, Mount
Whyte; C4m and U.S.N.M. 63j, Ross Lake; C5m and U.S.N.M. 35¢,
Mount Bosworth; C15m, Bow Lake; C2om, Eiffel Peak; U.S.N.M.
63m’, Mount Bosworth.

Types.—Syntypes (from the Gordon shale): U.S.N.M. Nos.
58368-71. Lectotype and paratype of Kochina bosworthensis Resser:
U.S.N.M. Nos. 63749-51. Plesiotypes: U.S.N.M. Nos. 116170-1.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 231

KOCHINA MACROPS Rasetti, new species
Plate 19, figures 17-19

Known from a few cranidia preserved in limestone.

Glabella rather strongly tapered, rounded in front, moderately
convex. Posterior glabellar furrows wide and shallow; two other
pairs almost indistinct. Occipital furrow well-impressed at the sides,
shallow mesially ; occipital ring short. Brim slightly convex, strongly
downsloping laterally ; midlength of brim equal to midlength of rim.
Rim tapered laterally, flat, slightly arched transversely, defined by an
almost straight marginal furrow. Fixed cheeks flat, upsloping, al-
most as wide as the glabella. Ocular ridges straight. Palpebral lobes
fairly wide, curved, defined by shallow palpebral furrows, about half
the glabellar length, continuing the upward slope of the fixed cheeks.
Distance from posterior end of palpebral lobe to posterior margin
less than half length of palpebral lobe. Anterior facial sutures on the
average parallel, slightly convex outward, widely curving across mar-
ginal furrow and rim. Posterior branch parallel to posterior margin ;
posterior limbs wide and short.

Elevated parts of cranidium covered with granules of different
sizes.

Length of holotype cranidium 8 mm. Fragments indicate the pres-
ence of individuals of larger size.

This species differs from K. americana in the proportionately wider
and shorter glabella, different proportions of brim and rim, flat in-
stead of convex fixed cheeks, straight ocular ridges, and much larger
palpebral lobes.

Horizon and locality—Cathedral formation (Albertella zone). Lo-
cality C15n, Bow Lake.

Types.—Holotype: U.S.N.M. No. 116168. Paratypes: U.S.N.M.
No. 116169.

Genus MEXICELLA Lochman, 1948
Genotype: Mexicella mexicana Lochman.
MEXICELLA STATOR (Walcott)
Plate 20, figures 14-19
Agraulos stator Waucott, Smithsonian Misc. Coll., vol. 64, No. 3, p. 173, pl. 36,
fig. 6, 1916; vol. 67, No. 2, p. 28, pl. 6, fig. 6, 1917:

Alokistocare stator (Walcott), RESsER, Smithsonian Misc. Coll., vol. 93, No. 5,

Pp. 9, 1935.
Mexicella stator (Walcott), LocHMAN, Journ. Paleontol., vol. 22, p. 457, 1948.

232 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Walcott’s diagnosis adequately describes this species. Cranidia in
limestone are figured in order to show the convexity of the various
parts which cannot be well ascertained from specimens in shale.

Horizon and locality—Cathedral formation (Ross Lake shale mem-
ber ; Albertella zone). Type locality U.S.N.M. 35c, Mount Bosworth.
Other localities U.S.N.M. 63j, Ross Lake; C3m, Mount Whyte ; C4m,
Ross Lake; C5m, Mount Bosworth; C15m, Bow Lake; C2om, Eiffel
Peak; C21m, Mount Temple; U.S.N.M. 63m’, Mount Bosworth.

Types.—Holotype: U.S.N.M. No. 61729. Paratype: U.S.N.M.
No. 61729a. Plesiotypes: U.S.N.M. Nos. 116176-7.

Genus ONCHOCEPHALUS Resser, 1937
Genotype: Ptychoparia thia Walcott.

ONCHOCEPHALUS THIA (Walcott)
Plate 8, figures I, 2
Ptychoparia thia Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 96, pl. 12,
fig. 6, 1917.
Onchocephalus thia (Walcott), REssErR, Smithsonian Misc. Coll., vol. 95, No. 22,
p. 20, 1937.
Onchocephalus thia (Walcott), LocHMAN, Journ. Paleontol., vol. 21, p. 63, 1947.
This species has been discussed by Lochman in her revision of
several Lower Cambrian ptychoparid genera. The writer finds it
exceedingly difficult to establish sharp lines of division between the
present genus and other genera of generalized ptychoparids. Besides
the genotype, which is a Lower Cambrian form, several species from
the early Medial Cambrian beds are assignable to Onchocephalus.
A cranidium collected by the writer from the type horizon is figured.
Horizon and locality—St. Piran formation (Peyto limestone mem-
ber; Bonnia-Olenellus zone). Type locality U.S.N.M. 35h, Mount
Bosworth. Also locality P18m, Mount Stephen.
Types——Holotype: U.S.N.M. No. 64388. Plesiotype: U.S.N.M.
No. 116070.

ONCHOCEPHALUS FIELDENSIS Rasetti, new species
Plate 14, figures 11-14

Known from numerous cranidia excellently preserved in limestone.
Glabella moderately tapered, fairly convex transversely and longi-
tudinally, rounded in front, defined by a dorsal furrow which is fairly
deep posteriorly but very shallow in front. Traces of three pairs of

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 233

glabellar furrows are visible on the upper surface. Occipital furrow
fairly deep laterally, shallow mesially. Occipital ring expanded mesi-
ally, bearing a small node. Brim convex, downsloping. Rim weakly
convex, expanded mesially where it attains the same length as the
brim; marginal furrow wide and shallow. Ocular ridges very low,
curving backward. Palpebral lobes at the level of the glabellar mid-
point, one-third the glabellar length, narrow, set off by a distinct pal-
pebral furrow. Fixed cheeks downsloping, their maximum width
two-thirds the glabellar width. Distance from posterior end of pal-
pebral lobes to posterior margin equal to length of palpebral lobe.
Posterior limb as wide as the occipital ring. Anterior branch of facial
suture straight forward from palpebral lobe to marginal furrow. Pos-
terior branch first directed outward, then curving backward to meet
posterior margin at a right angle.

Surface covered with very fine and not greatly elevated granules.

Length of largest cranidium 5 mm.

This species is very similar to the genotype, O. thia, differing chiefly
in the more definitely downsloping fixed cheeks, proportionately nar-
rower and longer glabella, and lesser midlength of the rim.

Horizon and locality—Mount Whyte formation (Plagiura-K ochas-
pis zone). Type locality W7{, Kicking Horse Mine, Mount Field.
Also locality W7c, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116129. Paratypes: U.S.N.M.
No. 116130.

ONCHOCEPHALUS DEPRESSUS Rasetti, new species
Plate 14, figures 15-17

Known from numerous cranidia well preserved in limestone.

This form is so similar to the preceding species that it will be suf-
ficient to list the differential features instead of repeating a complete
description of the cranidium. Compared with O. fieldensis, the mid-
length of the rim is, on the average, somewhat greater in proportion
to the midlength of the brim. The ocular ridges are considerably more
conspicuous, while the palpebral lobes are a little shorter. The general
convexity of the glabella and the whole cranidium, as well as the
surface ornamentation, are as in the preceding species.

Length of the largest cranidium 3.5 mm.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W7g, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116131. Paratypes: U.S.N.M.
No. 116132.

234 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ONCHOCEPHALUS MAIOR Rasetti, new species
Plate 14, figures 19-23

Known from numerous cranidia well preserved in limestone.

Glabella moderately convex in both directions, with slightly concave
sides, moderately tapered, not sharply truncated in front. Glabellar
furrows shallow; three pairs visible in most specimens. Occipital
furrow deep at the sides, shallow mesially ; occipital ring somewhat
elevated, bearing a small node. Brim convex, downsloping ; rim con-
vex, set off by a well-impressed marginal furrow, slightly swollen
mesially ; midlength of rim somewhat greater than midlength of brim.
Fixed cheeks slightly convex, on the average downsloping, about
two-thirds the glabellar width. Ocular ridges prominent, somewhat
curved, definitely oblique. Palpebral lobes small, elevated, situated
at the level of the glabellar midpoint. Posterior limbs about as wide
as the occipital ring, deeply furrowed. Distance from posterior end of
palpebral lobe to posterior margin somewhat greater than length of
palpebral lobe. Anterior facial sutures slightly divergent in front of
the eyes, curving inward after crossing the marginal furrow. Pos-
terior branch almost straight outward and backward for some distance,
then curving backward to posterior margin.

Surface of cranidium covered with granules of average size.

Length of largest cranidium 9 mm.

This species differs from the associated O. fieldensis chiefly in the
slightly concave sides of the glabella, greater relative midlength of rim,
deeper dorsal furrow, more prominent ocular ridges, and greater size.
Compared with the genotype, O. thia, the entire cranidium, and the gla-
bella in particular, are proportionately longer and narrower in the
present species.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Locality W7f, Kicking Horse Mine, Mount Field.

Types.—Holotype: U.S.N.M. No. 116134. Paratypes: U.S.N.M.
No. 116135.

ONCHOCEPHALUS SUBLAEVIS Rasetti, new species
Plate 14, figure 18

Known from a few cranidia preserved in limestone.

This species is so similar to O. maior that it will be sufficient to
mention the differential characters. The only significant differences
that the writer was able to ascertain are the much finer surface granu-
lation and the absence of an occipital node. The palpebral lobes are

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 235

broken off in the available cranidia, but apparently had the same
position and size as in O. maior.

Length of largest cranidium 7 mm.

Horizon and locality—Mount Whyte formation ( Plagiura-Koch-
aspis zone). Locality W7g, Kicking Horse Mine, Mount Field.

Type.—Holotype: U.S.N.M. No. 116133.

Genus PACHYASPIS Resser, 1939
Genotype: Pachyaspis typicalis Resser.

PACHYASPIS ATTENUATA Rasetti, new species
Plate 33, figures 12-14

Known from numerous cranidia preserved in limestone.

Glabella relatively long and narrow, strongly convex, moderately
tapered, rounded in front. Two pairs of shallow glabellar furrows.
Occipital furrow well impressed, occipital ring short, bearing a small
node. Brim convex, downsloping, with a pair of wide, shallow
depressions running forward and outward from the anterior corners
of the glabella. Rim upturned, well defined by the marginal furrow ;
midlength of brim about three times midlength of rim. Fixed cheeks
convex, on the average slightly downsloping, almost equaling the
glabella in width. Ocular ridges strong, starting from the anterior
angles of the glabella and running rather obliquely outward and back-
ward. Palpebral lobes well defined by the palpebral furrows, mod-
erately wide, about one-fourth the length of the glabella. Distance
from posterior end of palpebral lobe to posterior margin about twice
length of palpebral lobe. Posterior limbs deeply and widely furrowed.
Anterior sutures almost straight to anterior margin and slightly con-
vergent ; posterior branch running outward and backward to posterior
margin without much curvature.

Surface of test densely covered with fine granules and showing a
few scattered larger granules. Brim with the usual anastomosing
lines, more apparent on the impression of the lower surface.

Length of largest cranidium I0 mm.

This species closely resembles the genotype, P. typicalis, in the
essential cranidial features ; the other parts of the shield are unknown
in both forms. The present species chiefly differs in the definitely
convergent anterior facial sutures, less strongly downsloping fixed
cheeks, and more marked longitudinal lines on the brim. The same
features distinguish the new species from P. moorei Resser.

236 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Horizon and locality.—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S6l, Mount Odaray.

Types—Holotype: U.S.N.M. No. 116272. Paratypes: U.S.N.M.
No. 116273.

Genus PIAZELLA Lochman, 1947
Genotype: Ptychoparia pia Walcott.
PIAZELLA PIA (Walcott)
Plate 8, figures 3-5

Ptychoparia pia Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 93, pl. 12,
fig. 8, 1917.

Antagmus pia (Walcott), RESsER, Smithsonian Misc. Coll., vol. 95, No. 22, p. 1,
1937.

Piazella pia (Walcott), LocuMan, Journ. Paleontol., vol. 21, p. 69, 1947.

This Lower Cambrian species has been sufficiently described by
Walcott and by Lochman. However, all the published figures are
poor, hence a good specimen collected by the writer from the typical
horizon and locality is illustrated.

Horizon and locality.—St. Piran sandstone (Peyto limestone mem-
ber; Bonmia-Olenellus zone). Type locality U.S.N.M. 35f, Mount
Stephen. Also author’s locality P18m, Mount Stephen, probably
identical with the preceding.

Types.—Holotype: U.S.N.M. No. 64391. Plesiotype: U.S.N.M.
No. 116071.

PIAZELLA TUBERCULATA Rasetti, new species
Plate 8, figures 6-10

Known from several cranidia preserved in limestone.

Glabella moderately convex, tapered, rather sharply truncated in
front, with slightly concave sides. Three pairs of shallow glabellar
furrows; first pair indistinct. Occipital furrow deep at the sides, shal-
low mesially. Occipital ring bearing a small node. Brim convex and
downsloping at the sides, reduced to a short length mesially. Rim
thick, convex, demarcated by a deep marginal furrow, expanded back-
ward mesially. Midlength of rim somewhat greater than midlength
of brim. Fixed cheeks slightly convex and downsloping, almost equal-
ing the glabella in width. Ocular ridges wide and rather prominent.
Palpebral lobes elevated, somewhat less than one-third the glabellar
length. Distance from posterior end of palpebral lobe to posterior
margin greater than length of palpebral lobe. Posterior limbs con-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 237

siderably wider than occipital ring, deeply furrowed. Anterior facial
sutures divergent in front of the eyes, slightly convex outward to the
marginal furrow, then definitely curving inward. Posterior branch
reaching the posterior margin with a wide curve.

Surface of test covered with small granules, among which are
scattered larger tubercles.

Length of largest cranidium 13 mm.

This species appears to agree with the genotype, P. pia, in all the
essential features, particularly the great relative width of the fixed
cheeks. It differs from that species chiefly in the surface ornamen-
tation.

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Locality W16j, Mount Stephen.

Types——Holotype: U.S.N.M. No. 116072. Paratypes: U.S.N.M.
No. 116073.

Genus PLAGIURA Resser, 1935
Genotype: Ptychoparia? cercops Walcott.

PLAGIURA CERCOPS (Walcott)
Plate 13, figures 10-16

Ptychoparia? cercops Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 81,
pl. 12, figs. 1, 1a-d, 1917.

Plagiura cercops (Walcott), REssEr, Smithsonian Misc. Coll., vol. 93, No. 5,
P. 43, 1935.

Plagiura cercops (Walcott), LocHMAN, Journ. Paleontol., vol. 21, p. 66, 1947.

Ptychoparia? cleadas Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 83,
pl. 12, fig. 2, 1917.

Plagiurella cleadas (Walcott), RESSER, Smithsonian Misc. Coll., vol. 95, No. 22,
Pp. 23, 1937.

Plagiura cleadas (Walcott), LocuMan, Journ. Paleontol., vol. 21, p. 66, 1947.

Lochman showed that there is no reason for the generic separation
of the two species described by Walcott. Further material collected
by the writer shows continuous growth series between the small
cranidia on which Ptychoparia cleadas was based, and the large
cranidia representing the typical Ptychoparia cercops. Hence the
former name is placed in synonymy.

The chief features that differentiate the cranidia of the young from
the adults are the presence of shallow glabellar furrows, the deeper
dorsal and occipital furrows, the distinct ocular ridges, and the pres-
ence of a well-differentiated, convex anterior rim. In mature indi-

238 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

viduals the rim becomes flat and less differentiated from the brim, yet
the marginal furrow does not become entirely obsolete.

Complete articulated shields are unknown ; however, the association
of cranidium and pygidium was observed in so many instances as to
make the assignment virtually certain. The writer collected a shield
in shale preserving the pygidium and 15 thoracic segments, possibly
representing the complex thorax. The thoracic axis is slightly tapered.
The pleura show a fairly sharp geniculation at about one-third the
length from the the proximal end. The pleura terminate in sharp, flat,
slightly falcate spines. Pleural furrows well impressed at the genicu-
lation, shallow proximally and distally. Length of thorax and py-
gidium 19 mm., of pygidium 2 mm.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Type locality U.S.N.M. 63c, Ptarmigan Pass. Type
locality for Ptychoparia cleadas U.S.N.M. 57s, Mount Bosworth.
Other localities W4c, W4d, Ross Lake; W5c, Mount Bosworth ; Wof,
Mount Stephen; Wigc, Hector Creek; W2o0d, Eiffel Peak.

Types.—Holotype: U.S.N.M. No. 64378. Paratypes: U.S.N.M.
Nos. 64377, 64379-81. Holotype of Ptychoparia cleadas: U.S.N.M.
No. 64382. Plesiotypes: U.S.N.M. Nos. 116118-9.

Genus SCHISTOMETOPUS Resser, 1938

Genotype: Schistometopus typicalis Resser.

The genus was described from a single cranidium. One of the
characters of the genus, as stated by Resser, is the elevation of the
central portion of the rim while the lateral portions are depressed.
Although the only known specimen of S. typicalis is badly flattened
in shale, careful examination seems to prove that this feature was
really present in the animal and is not due to an accident of preser-
vation.

A species from the Mount Whyte formation does not possess an
elevated median portion of the rim but agrees so well with Schisto-
metopus typicalis in all other features that a separate genus does not
seem indicated. Resser himself had referred this form to Schisto-
metopus in labels attached to specimens in the United States National
Museum.

SCHISTOMETOPUS CONVEXUS Rasetti, new species
Plate 13, figures 17-22

Known from numerous cranidia preserved in limestone.
Glabella moderately tapered, straight-sided, rather truncated in
front, strongly convex in both directions. Glabellar furrows moder-

=.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 239

ately impressed. Posterior pair oblique, showing a tendency to bi-
furcate near their inner ends. Third pair also directed inward and
backward, shorter than the posterior pair. Second pair shorter, shal-
lower, and directed inward and slightly forward. There is a bare trace
of the first pair. Occipital furrow deep, straight; occipital ring ex-
panded mesially, bearing a small node. Dorsal furrow deep. Brim ab-
sent mesially, convex and downsloping laterally. Rim convex, almost
straight ; marginal furrow somewhat turned backward mesially and
confluent with the dorsal furrow in front of the glabella. Fixed
cheeks convex, on the average horizontal ; maximum width somewhat
more than half the glabellar width. Ocular ridges strongly oblique ;
palpebral lobes about one-third the glabellar length, moderately wide,
set off by well-impressed palpebral furrows; their midpoint situated
somewhat back of the glabellar midpoint. Total width of posterior
limbs slightly less than width of occipital ring. Distance from posterior
end of palpebral lobe to posterior margin less than length of palpebral
lobe. Posterior marginal furrow deep. Anterior facial sutures slightly
convex outward and somewhat divergent; posterior branch directed
straight outward behind palpebral lobe, then curving backward.

Entire surface of cranidium covered with small granules, among
which are scattered a few much larger granules; the latter form a
somewhat regular pattern.

Length of the largest cranidium 13 mm.

This species chiefly differs from the genotype, S. typicalis Resser,
in lacking the elevated median portion of the anterior rim, and having
only three pairs of distinct glabellar furrows.

Horizon and locality—Mount Whyte formation (Plagiura-Koch-
aspis zone). Type locality W2od, Eiffel Peak. Other localities W4c-d,
Ross Lake ; W5c, Mount Bosworth ; W7{, Kicking Horse Mine; Wof,
Fossil Gully.

Types.—Holotype: U.S.N.M. No. 116120. Paratypes: U.S.N.M.
Nos. 116121-2.

SCHISTOMETOPUS COLLARIS Rasetti, new species
Plate 14, figures 1-3

Known from several cranidia preserved in limestone. All the avail-
able specimens are exfoliated, hence the description is based on the
appearance of the internal cast.

Glabella moderately tapered, rounded in front, strongly convex in
both directions. Three pairs of glabellar furrows fairly well impressed.
Occipital ring extended into a slender, short spine. Brim at the sides

240 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

of glabella convex, downsloping; in front of the glabella reduced
to an exceedingly short length but not entirely eliminated as in S.
convexus. Marginal furrow straighter than in S. convexus; rim
convex, thicker mesially and tapered laterally. Fixed cheeks consider-
ably convex, on the average downsloping, between half and two-thirds
the glabellar width. Ocular ridges less prominent and less oblique
than in S. convexus. Palpebral lobes smaller than in S. convexus.
Posterior limbs subtriangular; distance between posterior end of
palpebral lobe and posterior margin about twice the length of palpe-
bral lobe. Anterior facial sutures slightly divergent for a short distance
in front of the eyes, then curving inward and becoming convergent.

Characters of surface unknown. Length of largest cranidium
13 mm.

This species is undoubtedly a close relative of S. convexus, with
which it is associated in the Plagiura-Kochaspis zone. Most of the
differences from the preceding species, mentioned in the description,
bring it closer to the average ptychoparid.

Horizon and locality—Mount Whyte formation (Plagiura-K och-
aspis zone). Locality W2od, Eiffel Peak.

Types—Holotype: U.S.N.M. No. 116123. Paratypes: U.S.N.M.
No. 116124.

Genus SOLENOPLEURELLA Poulsen, 1927

Genotype: Solenopleurella ulrichi Poulsen.

The genotype of Solenopleurella is known from a single cranidium
collected in the Cape Wood formation of northwest Greenland. Forms
that have been assigned to the genus are common in some of the
Middle Cambrian formations of the Appalachian and Cordilleran
provinces, and some of the species are represented by complete shields.
These forms, provided they are congeneric with the genotype, may be
used to supplement the generic diagnosis by stating that the thorax
has 14-16 segments with simple pleura rounded at the extremity, and
the pygidium is small, transverse, showing about two segments on the
axis and pleural lobes.

Resser (1945) assigned to a species of Solenopleurella a large py-
gidium that certainly does not belong to the genus.

Several other genera of ptychoparids are very close to Solenopleur-
ella and intergrade in such a way that further study may lead to
uniting all these forms in one genus. These genera are Crusoia Wal-
cott, 1924 (genotype: Crusoia cebes Walcott) ; Spencia Resser, 1939
(genotype: Spencia typicalis Resser) ; Stauroholcus Resser, 1939
(genotype: Stauroholcus typicalis Resser). The last two genera are

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 241

obviously synonymous, the only differential character being the un-
important presence of an occipital spine in Stauroholcus. The forms
of this group show all intermediate stages between a small occipital
node and a strong spine.

SOLENOPLEURELLA, species undetermined No. 1
Plate 25, figures 17, 18

Apparently the same form of Solenopleurella occurs in the Burgess
shale and in supposedly equivalent beds on Mount Odaray, where the
fossils were collected from limestone lenses in shale. The different
manner of preservation makes an accurate comparison difficult.

This form closely resembles S. diligens Resser from the Grand
Canyon. Owing to the insufficient material, specific identification is
not attempted.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Localities S6g, Mount Odaray, and Siig, above Burgess
Quarry, Mount Field.

Figured specimens.—U.S.N.M. Nos. 116218-9.

SOLENOPLEURELLA, species undetermined No. 2
Plate 25, figure 16

A second species of Solenopleurella is represented by a few weath-
ered shields, one being almost complete.

The cranidium appears to differ from that of the preceding form
in possessing a less-thickened rim. The elevated portions of the gla-
bella were weathered away in the available cranidia.

The thorax consists of 14 segments. The pleura have the genicu-
lation much closer to the proximal than to the distal end, and are
rounded at the extremity. The pleural furrows decrease in depth
toward the pygidium. The pygidium is small, more than twice wider
than long, and shows barely a trace of two pairs of pleural furrows.
Length of cranidium 5 mm., of thorax 8.4 mm., of pygidium 1.3 mm.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S11e, Mount Field.

Figured specimen.—U.S.N.M. No. 116217.

Genus SYSPACEPHALUS Resser, 1936

Genotype: Agraulos charops Walcott.
This genus, recently discussed by Lochman (1947), is somewhat
better characterized than most of the other genera of Early Cambrian

242 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

and early Medial Cambrian generalized ptychoparids, chiefly because
of the unusual anterior position of the palpebral lobes and the con-
vergence of the anterior facial sutures.

The type species is from the Lower Cambrian Peyto limestone.
Nevertheless, it is figured here because it is important that the geno-
types in this difficult group of trilobites be illustrated as well as
possible.

Several Middle Cambrian species are assigned to the genus, as they
wholly agree with the generic characters of the Lower Cambrian
genotype. Some of the new forms are known from complete shields,
hence the generic diagnosis may be completed by stating that the
thorax consists of 13 to 15 segments and the pygidium is small, trans-
verse, with only two distinct segments. Thorax and pygidium do not
supply any characters than can be used to discriminate this genus from
similar generalized ptychoparids.

SYSPACEPHALUS CHAROPS (Walcott)
Plate 8, figures I1-13

Agraulos charops Watcort, Smithsonian Misc. Coll., vol. 67, No. 3, p. 67, pl. 13,
figs. 2, 2a, I917.

Syspacephalus charops (Walcott), RrEssEr, Smithsonian Misc. Coll., vol. 95,
No. 4, p. 28, 1936.

Syspacephalus charops (Walcott), LocoMAn, Journ. Paleontol., vol. 21, p. 64,
1947.

A specimen from the typical horizon and at least approximately the
type locality is figured.

Horizon and locality—Peyto limestone member of the St. Piran
sandstone (Bonnia-Olenellus zone). Type locality U.S.N.M. 35f,
Mount Stephen. Plesiotypes from locality P18m, Mount Stephen.

Types.—Holotype and paratypes: U.S.N.M. No. 64395. Plesio-
type: U.S.N.M. No. 116074.

SYSPACEPHALUS GREGARIUS Rasetti, new species
Plate 8, figures 14-19

Known from a large number of cranidia and several complete
shields in shale, and several cranidia in limestone preserving the full
convexity.

Glabella slightly tapered, truncated in front, rather strongly convex,
defined by a dorsal furrow that is deep at the sides and shallow in
the frontal portion. Three pairs of glabellar furrows well impressed ;
fourth furrow directed backward, third transverse, second directed

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 243

slightly forward. Occipital furrow deep; occipital ring elevated, bear-
ing anode. Brim and fixed cheeks convex ; fixed cheeks on the average
slightly downsloping. Rim convex, expanded mesially, its midlength
about two-thirds of the brim midlength. Marginal furrow almost obso-
lete mesially. Fixed cheeks almost as wide as the glabella. Palpebral
lobes about one-third the glabellar length, somewhat elevated, situated
somewhat in advance of the glabellar midpoint. Ocular ridges narrow,
transverse, slightly curved. Anterior facial suture directed slightly
inward, curving across the border; posterior branch almost straight,
narrowly curving near the distal end. Distance from posterior end of
palpebral lobe to posterior margin of cranidium somewhat greater than
length of palpebral lobe. Posterior limbs about as wide as occipital
ring. Free cheeks small and rounded at the genal angle.

Thorax of 14 segments. Pleura deeply furrowed, rounded at the
distal end.

Pygidium small, elliptical, about three times wider than long. Axis
with two visible segments. Pleural lobes showing one distinct pair
of furrows and traces of another.

Surface characterized by small scattered tubercles.

Length of largest cranidium 4.8 mm. Most of the specimens are
somewhat smaller. The length of an average complete shield is
12 mm,

This species resembles in all characters of generic significance the
genotype, S. charops. Chief distinctive features are the greater con-
vexity of the fixed cheeks, deeper glabellar furrows, more elevated
palpebral lobes, and different course of the ocular ridges.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
Wenkchemmia-Stephenaspis zone). Type locality Wok, Fossil Gully,
Mount Stephen. Other locality W1i6k, North Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116075. Paratypes: U.S.N.M.
Nos. 116076-7.

SYSPACEPHALUS LATICEPS Rasetti, new species
Plate 9, figures 1-6

Known from numerous cranidia preserved in limestone.

Glabella slightly tapered, straight-sided, truncated in front, most
elevated posteriorly, with the third and fourth glabellar furrows fairly
well impressed and traces of one or two other pairs. Occipital furrow
deep at the sides; occipital ring expanded mesially, bearing a small
node. Brim and fixed cheeks convex; fixed cheeks on the average
downsloping. Rim flat, slightly upturned with respect to the brim ; rim

244 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II16

midlength about half of the brim midlength. Fixed cheeks almost as
wide as the glabella. Palpebral lobes slightly elevated, about one-
fourth the glabellar length, situated in advance of a line through the
anterior third of the glabella. Ocular ridges weak, starting forward
from the dorsal furrow, distally curving backward, on the average
transverse. Anterior facial suture directed somewhat inward in front
of the eyes, curving farther inward after crossing the marginal fur-
row, marginal for a short distance. Posterior branch directed almost
straight outward and backward; distance from posterior end of pal-
pebral lobe to posterior margin of cranidium more than twice the
length of palpebral lobe. Posterior limbs considerably wider than the
occipital ring.

Surface smooth except for faint longitudinal lines on the brim.

Length of largest cranidium 6.5 mm.

This species closely resembles the genotype, S. charops, in all fea-
tures of generic significance. It is readily distinguished by the better-
impressed glabellar furrows, wider posterior limbs, and lack of a
median boss on the frontal limb.

Horizon and locality—Mount Whyte formation (Wenkchemmnia-
Stephenaspis zone). Type locality W16j, North Gully, Mount Ste-
phen. Other localities W16j” and W16h, North Gully, Mount
Stephen.

Types.—Holotype: U.S.N.M. No. 116080. Paratypes: U.S.N.M.
No. 116081.

SYSPACEPHALUS PEROLA (Walcott)
Plate 9, figures 17-22

Ptychoparia perola Watcott, Smithsonian Misc. Coll., vol. 67, No. 3, p. 91,
pl. 12, figs. 7, 7a, 1917.

Inglefieldia perola (Walcott), REssER, Smithsonian Misc. Coll., vol. 95, No. 22,
p. 14, 1937.

Antagmus perola (Walcott), LocHMAN, Journ. Paleontol., vol. 21, p. 62, 1947.

Known from a large number of cranidia and several articulated
shields preserved in shale.

All the characters can be ascertained except the exact degree of
convexity because of the partial flattening of the shields. The position
of the palpebral lobes and the convergence of the facial sutures in
front of the eyes exclude reference of the species to Jnglefieldia or
Antagmus, whereas all characters of generic significance agree with
the genotype of Syspacephalus.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 245

Walcott gave a sufficient description of this species, except for the
statement that the free cheeks have genal spines. Study of the type
material in the United States National Museum shows that the speci-
men on which he based this observation is a free cheek of the associ-
ated species Amecephalus agnesensis. The writer collected several
individuals preserving the free cheeks and these show that the genal
angle is rounded.

The present species differs from S. charops chiefly in the deeper
glabellar furrows, reduced marginal portion of the facial suture, and
the course of the ocular ridges.

Horizon and locality—Mount Whyte formation (Lake Agnes and
Yoho shale lentils; Wenkchemmia-Stephenaspis zone). Type locality
U.S.N.M. 35m, southwest of Lake Louise. Other localities W2b,
Mount Niblock; W3b, Plain of Six Glaciers; Wok, Fossil Gully,
Mount Stephen.

Types.—Lectotype (hereby designated): U.S.N.M. No. 643809.
Paratype: U.S.N.M. No. 64390. Plesiotypes: U.S.N.M. Nos.
116087-8.

SYSPACEPHALUS LAEVIGATUS Rasetti, new species
Plate 9, figures 9-11

Known from several cranidia and a complete shield, all preserved
in shale and slightly flattened.

Glabella tapered, unfurrowed, not sharply truncated in front. Oc-
cipital ring expanded, bearing an indistinct node. Frontal limb and
fixed cheeks convex. Limb faintly divided into brim and rim by a
marginal furrow that is distinct only at the sides; brim longer than
rim. Fixed cheeks about two-thirds the glabellar width. Palpebral
lobes narrow, slightly elevated, situated on a transverse line through
the anterior third of the glabella. Ocular ridges indistinct, faintly
curved, directed transversely. Anterior branch of facial suture di-
rected slightly inward in front of the eyes, gently curving across the
rim; marginal portion of moderate extent. Posterior branch directed
outward and backward behind the eye, then curving backward, Dis-
tance from posterior end of palpebral lobe to posterior margin at
least twice length of palpebral lobe. Width of posterior limbs approxi-
mately equal to width of occipital ring.

Thorax of 13 segments. Pleura broadly furrowed, rounded at the
distal end.

Pygidium small, transverse, about three times wider than long. Axis
almost reaching the posterior margin, pleural lobes about as wide as

246 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

the axis. Both axis and pleural lobes show one furrow and traces of
a second pair.

Surface smooth.

Length of largest cranidium 7 mm. Length of a complete shield
14 mm.

Horizon and locality—Mount Whyte formation (Yoho shale lentil ;
W enkchemnia-Stephenaspis zone). Type locality Wok, Fossil Gully,
Mount Stephen.

Types—Holotype: U.S.N.M. No. 116083. Paratypes: U.S.N.M.
No. 116084.

SYSPACEPHALUS CRASSUS Rasetti, new species
Plate 9, figures 12-16

Known from numerous cranidia preserved in limestone and an en-
tire shield in shale.

Glabella slightly tapered, strongly convex, rounded in front, con-
siderably elevated above the fixed cheeks. Two pairs of short glabellar
furrows of medium strength. Occipital furrow well impressed ; oc-
cipital ring expanded, bearing a strong node. Brim and fixed cheeks
convex. Rim about half the length of the brim on midline, slightly
convex; marginal furrow indistinct mesially. Fixed cheeks on the
average slightly downsloping, about two-thirds the glabellar width.
Ocular ridges transverse, slightly curved. Palpebral lobes rather
elevated, about one-third the glabellar length, their centers situated
somewhat in advance of the glabellar midpoint. Anterior facial su-
ture directed somewhat inward in front of the eye, curving across the
rim, marginal for a considerable distance. Posterior branch directed
outward and backward, curving backward near the distal end. Dis-
tance from posterior end of palpebral lobe to posterior margin some-
what greater than length of palpebral lobe. Width of posterior limbs
equal to width of occipital ring.

The thorax has 14 segments and the pygidium is small, transverse
as in S. gregarius.

Surface covered with scattered granules. Length of largest cranid-
ium 4 mm.

This species differs from the genotype, S. charops, in several fea-
tures, chiefly the greater proportional size and elevation of the gla-
bella, distinctness of the glabellar furrows, elevation of the palpebral
lobes, and surface ornamentation. The first two characters also
separate it from S. gregarius which it resembles in most of the other
features.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 247

Horizon and locality—Mount Whyte formation (Wenkchemnia-
Stephenaspis zone). Type locality W16j, North Gully, Mount Ste-
phen. Other locality W16j”, North Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116085. Paratypes: U.S.N.M.
No. 116086.

SYSPACEPHALUS TARDUS Rasetti, new species
Plate 22, figures 7-10

Known from numerous cranidia moderately well preserved in silty
limestone.

Glabella of average convexity, slightly tapered, rounded in front.
Glabellar furrows indistinct. Occipital furrow shallow ; occipital ring
not greatly expanded, possibly bearing a node. Brim convex, down-
sloping ; marginal furrow rather straight; rim on the average hori-
zontal, slightly convex, with a swelling on the median line visible in
most of the specimens. Midlength of rim equal to midlength of brim.
Fixed cheeks convex, on the average slightly downsloping, somewhat
more than two-thirds the glabellar width. Ocular ridges faint. Pal-
pebral lobes small, elevated, situated on a transverse line through the
anterior third of the glabella. Distance from posterior end of palpe-
bral lobe to posterior margin more than twice length of palpebral lobe.
Posterior limbs slightly wider than occipital ring. Anterior facial
sutures slightly convergent in front of the eyes, curving inward across
the rim. Posterior branch directed almost straight outward behind
the eye, then rather sharply turning backward.

Surface of test apparently smooth. However, the state of pres-
ervation would not allow the detection of very small granules.

Length of largest cranidium 6 mm.

This species, although much younger than the genotype, appears
entirely to agree with the generic characters of Syspacephalus and
it does not seem appropriate to remove it from the genus on account
of its age. Compared with the genotype, S. charops, the present
species differs chiefly in the greater convexity of the anterior portion
of the glabella and the horizontal rather than downsloping position
of the rim.

Horizon and locality—Cathedral formation (Albertella(?) zone).
Locality Cgh, Fossil Gully, Mount Stephen.

Type.—Holotype: U.S.N.M. No. 116188. Paratypes: U.S.N.M.
No. 116189.

248 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

YOHOASPIS Rasetti, new genus

Ptychoparid trilobites known from the cranidium.

Cranidium as a whole with considerable relief. Glabella strongly
convex transversely, moderately tapered, rounded in front, defined by
a deep dorsal furrow; glabellar furrows indistinct, occipital furrow
shallow. Frontal limb consisting of rim only, as the marginal furrow
runs into the dorsal furrow in front of the glabella. Rim long, down-
sloping ; marginal furrow almost straight. Fixed cheeks strongly con-
vex, not rising to the level of the glabella, almost as wide as the
glabella. Palpebral lobes narrow, moderately long, rather straight,
set off by a distinct palpebral furrow, situated at the level of the gla-
bellar midpoint. Ocular ridges poorly defined. Posterior limbs rather
slender, tapered, deeply furrowed. Anterior facial sutures somewhat
divergent in front of the eyes, curving inward across the rim, prob-
ably marginal on the median line. Posterior branch directed almost
straight outward and a little backward.

Genotype.—Y ohoaspis pachycephala Rasetti, new species.

Stratigraphic range-——Middle Cambrian (Albertella(?) zone).

Remarks.—Y ohoaspis is a rather large ptychoparid trilobite that
can be compared with other genera only as far as the cranidial fea-
tures are concerned, since the other parts of the shield are unknown.
Its chief diagnostic features are the strong convexity of the fixed
cheeks and especially the structure of the frontal limb, consisting of
a thick rim only. The confluence of the dorsal and marginal furrows
in front of the glabella occurs to a greater or lesser degree in several
genera of Lower and Middle Cambrian ptychoparids, but seldom to
such a complete extent as in the present instance. The genera present-
ing this feature differ from Yohoaspis in the proportions and con-
vexity of other cranidial parts.

The name is derived from the Yoho River.

YOHOASPIS PACHYCEPHALA Rasetti, new species
Plate 21, figures 9-14

Known from several cranidia preserved in shale or impure lime-
stone, the latter preserving the convexity.

Glabella moderately tapered, rounded in front, strongly convex in
both directions, defined by a deep dorsal furrow. Glabellar furrows
very faint. Occipital furrow deep laterally, shallow mesially; oc-
cipital ring apparently short and simple. Brim developed only at the
sides, convex. Rim occupying the entire frontal limb mesially because
of the confluence of the dorsal and marginal furrows; rim down-

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 249

sloping, slightly concave as a whole, convex marginally ; marginal
furrow almost straight. Fixed cheeks well convex, on the average
downsloping. Palpebral lobes small, elevated, situated on the trans-
verse line through the glabellar midpoint. Ocular ridges faint. Pos-
terior limbs deeply furrowed, wider than the occipital ring. Distance
from posterior end of palpebral lobe to posterior margin about twice
length of palpebral lobe. Anterior facial sutures divergent in front of
the eyes, turning inward with a wide curve across the rim. Posterior
branch fairly straight for a distance behind the eye, then gradually
curving backward.

Surface of test densely covered with very fine granules.

Length of largest cranidium 20 mm.

Horizon and locality——Cathedral formation (Albertella(?) zone).
Type locality Coh, Fossil Gully, Mount Stephen. Also locality Cgj,
Fossil Gully, Mount Stephen.

Types.—Holotype: U.S.N.M. No. 116181. Paratypes: U.S.N.M.
Nos. 116182-3.

YUKNESSASPIS Rasetti, new genus

Ptychoparid trilobites with multisegmented thorax and a small py-
gidium.

Glabella moderately convex, tapered, straight-sided, sharply trun-
cated in front. Dorsal furrow unusually deep at the sides. Frontal limb
as long as the glabella, chiefly consisting of a convex brim, with a
short rim. Fixed cheeks about equaling the glabellar width, upsloping.
Ocular ridges starting from the anterior angles of the glabella, un-
usually strong and prominent. Palpebral lobes short and narrow. Pos-
terior limbs wider than the occipital ring, widely furrowed. Distance
from posterior end of palpebral lobe to posterior margin greater than
length of palpebral lobe.

Thorax of at least 14 segments in the genotype. Axis moderately
tapered. Pleura straight, widely furrowed. Pygidium unknown. From
the rate of tapering of the thorax it is certain that this part of the
shield was small.

Genotype.—Yuknessaspis paradoxa Rasetti, new species.

Stratigraphic range-——Middle Cambrian (Bathyuriscus-Elrathina
zone).

Remarks.—The genus cannot be confused with any described pty-
choparid, chiefly on account of the great depth of the dorsal furrow,
strength of the ocular ridges, and long, convex brim that almost rises
to the level of the glabella.

The name is derived from Mount Yukness.

250 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

YUKNESSASPIS PARADOXA Rasetti, new species
Plate 32, figures 15-18

Known from two cranidia and a specimen with attached thorax,
all preserved in limestone.

Glabella moderately tapered, with slightly concave sides, sharply
truncated in front, strongly convex transversely but moderately con-
vex longitudinally. Two or three pairs of oblique glabellar furrows
are barely indicated. Occipital furrow consisting of a pair of narrow,
oblique, rather deep impressions at the sides connected by a shallow
median furrow. Occipital ring extended into a spine. Dorsal furrow
wide and deep at the sides, narrow and moderately deep in front of
the glabella. At the sides of the anterior portion of the glabella the
dorsal furrow forms a pair of deep pits, from which the ocular ridges
rise with unusual prominence. Frontal limb equaling the glabella in
length. Brim strongly convex, almost rising to the level of the gla-
bella; marginal furrow narrow but well impressed, rim convex, its
midlength about one-eighth of the midlength of the brim. Fixed cheeks
flat, upsloping, but not reaching the level of the glabella ; approximately
equaling the glabella in width. Ocular ridges originating at the level
of the anterior angles of the glabella, first directed outward, then
slightly curving backward. Palpebral lobes upsloping, about one-
fourth the glabellar length, set off by a narrow palpebral furrow.
Posterior limbs triangular, considerably wider than the occipital ring.
Distance from posterior end of palpebral lobe to posterior margin
greater than length of palpebral lobe. Marginal furrow on posterior
limbs wide and deep. Anterior facial sutures divergent in front of
the eyes, then describing a wide curve and crossing the rim obliquely.
Posterior branch directed fairly straight outward and backward.

Thorax of 14 segments in the only specimen where this portion
of the shield is preserved. From the rate of tapering of the last seg-
ments it is clear that this thorax is either complete or almost complete,
and that the pygidium was small. Pleural lobes with a fairly sharp
geniculation but no great convexity. Pleura straight, furrowed, not
extended into spines.

Surface of glabella smooth. Surface of fixed cheeks with fine
reticulated ornamentation. Surface of brim with coarse, longitudinal
anastomosing lines.

Length of holotype cranidium 15 mm.; length of another specimen
preserving cranidium and thorax 26 mm.

Horizon and locality—Stephen formation (Bathyuriscus-Elrathina
zone). Locality S1ol, Park Mountain.

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 251

Types.—Holotype: U.S.N.M. No. 116265. Paratype: U.S.N.M.
No. 116266.

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254

1912a.
1912b.
19I6a.
1916b.
I9I7a.
IQI7b.

19I7¢c.

1918.
1919.
1920.
1925.
1927.
1928.

1931.

WARBURG,

1925.

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EXPLANATION OF PLATES

PLATE I. South face of Mount Temple (11,636 feet)

Showing over 4,000 feet of horizontal Lower and Middle Cambrian strata.
P= St Piran;) W= Mount Whyte; C= Cathedral; S= Stephen; E=
Eldon.

Prate 2. Middle Cambrian strata on Mount Stephen and Park Mountain

Upper: Northeast shoulder of Mount Stephen. The portal of the Monarch
Mine is visible in the upper left corner. P=St. Piran; W = Mount
Whyte; C= Cathedral. Note the westward thickening of the lowermost
portion of the Mount Whyte formation.

Lower: East face of Park Mountain. C= Cathedral; S=Stephen; E=
Eldon. The Eldon is here represented by an unusual shaly and calcareous
facies.

PLATE 3. Northwest face of Mount Stephen

Photograph taken from the summit of Mount Field. P=St. Piran; W=
Mount Whyte; SC = Cathedral + Stephen (undivided) ; E= Eldon. NG=
North Gully; MG= Middle Gully; FG=Fossil Gully. Dotted line
represents Fossil Gully fault. The dark band above the letter E in the
Eldon is a siliceous shale.

PLATE 4. Lower part of the northwest face of Mount Stephen

Photographed from the south slope of Mount Field. P=St. Piran; W=
Mount Whyte; C= Cathedral. NG= North Gully; MG = Middle Gully;
FG= Fossil Gully. Dotted line represents Fossil Gully fault.

PLATE 5. Interfingering limestone and dolomite in the Cathedral formation

Dark-gray limestone and light-tan-weathering dolomite interfingering in the
Upper Cathedral formation on Mount Temple.

Piate 6. Anomalous lithology of the Cathedral formation on Mount Stephen

The lower portion of the Cathedral formation is represented by shale and
thin-bedded limestone in the western part of Mount Stephen. A gigantic
dolomite boulder in the thin-bedded sediments in Middle Gully, Mount
Stephen.

PLATE 7. East face of Mount Odaray

Photographed from the west shoulder of Mount Schaffer. P= St. Piran;
W = Mount Whyte; C=Cathedral; S= Stephen. The cliff at the top
of the St. Piran is formed by the Peyto limestone member. The section
was measured on the slope of the south peak (left).

235

256 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Pirate 8. Trilobites of the Bonnia-Olenellus and Wenkchemnia-Stephenaspis
zones

(All the figured specimens are preserved in limestone unless otherwise indicated.)

Page
Bigs: 1; 2iOnchocephalus.thia (Walcott) yo. iaincistcinniap ces 2.3 cero ci miei ae 232
Dorsal and frontal views of cranidium, 6; U.S.N.M. No. 116070,
plesiotype (loc. P18m).

Bigs. 3-5) “rasellasprar (Walcoth) 2 soso vtec state ae atte re nde, 3 are 236
Dorsal, lateral, and frontal views of cranidium, & 3; U.S.N.M. No.
116071, plesiotype (loc. P18m).

Figs. 6-10. Piazella tuberculata Rasetti, new species..............s++seee 236
6-8, Dorsal, frontal, and lateral views of cranidium, X 2; U.S.N.M.
No. 116072, holotype. 9, 10, Two cranidia, 2; U.S.N.M. No.
116073a-b, paratypes (loc. W16j).

Bigs. 11-13.. Syspacephalus charops* (Walcott)... 2... Sees. eee. oe cee toe 242
Dorsal, frontal, and lateral views of cranidium, X 4; U.S.N.M. No.
116074a, plesiotype (loc. P18m).

Figs. 14-19. Syspacephalus gregarius Rasetti, new species..............-. 242
14-16, Lateral, frontal, and dorsal views of cranidium, X 4; U.S.N.M.
No. 116077a, paratype (loc. W16k). 17, 18, Two cranidia in sili-
ceous shale, &X 4; U.S.N.M. No. 116076a-b, paratypes (loc. Wok).
19, Cast from the external mold in siliceous shale, X 4; U.S.N.M.
No. 116075, holotype (loc. Wok).
Fig. 20. Chancia stenometopa Rasetti, new SPeCieS.........-e eee eeeeeeees 216

Cranidium in siliceous shale, &* 3; U.S.N.M. No. 116078, holotype (loc.
Wok).

PiateE 9. Trilobites of the Wenkchemnia-Stephenaspis zone

Figs. 1-6. Syspacephalus laticeps Rasetti, new species.............eeeeeeee 243
1-3, Dorsal, frontal, and lateral views of cranidium in limestone, X 4;
U.S.N.M. No. 116080, holotype. 4, 5, Dorsal and frontal views of
cranidium in limestone, X 4; U.S.N.M. No. 116081a, paratype. 6,
cranidium in limestone, & 4; U.S.N.M. No. 116081b, paratype (loc.

W16j).
Bigs 7 Gameoliclia Civic Aen tGUlal@masettlsaienyssreticeteriaicisteiieil tetera: 174
Cranidia in limestone, & 3; U.S.N.M. No. 116082a-b (loc. W16j).
Figs. 9-11. Syspacephalus laevigatus Rasetti, new species............-.00 245

9, Articulated shield in shale, & 3; U.S.N.M. No. 116083, holotype. 10,
11, Cranidia in shale, * 3; U.S.N.M. No. 116084a-b, paratypes (loc.
Wok).
Figs. 12-16. Syspacephalus crassus Rasetti, mew species.............eee0. 246
12-14 Lateral, frontal, and dorsal views of cranidium in limestone, X 6;
U.S.N.M. No. 116085, holotype. 15, 16, Cranidia in limestone, X 6;
U.S.N.M. No. 116086a-b, paratypes (loc. W16j).

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 257

Bigs. 17-22.) Syspacepnalus perolal (NW alcott) ca siclersiers wieieie ferscay eee) onl cistolore «6 244
17, Cast from mold in shale, X 2; U.S.N.M. No. 64389, holotype (loc.
U.S.N.M. 35m). 18, Cranidium and thorax in shale, K 2; U.S.N.M.
No. 116087, plesiotype (loc. Wok). 19, 21, Specimens in shale, * 2;
U.S.N.M. No. 116088a-b, plesiotypes (loc. W3b). 20, Cranidium in
shale, X 2; U.S.N.M. No. 64390, paratype (loc. U.S.N.M. 35m).
22, Cranidium in shale, X 4; U.S.N.M. No. 116087b, paratype (loc.

Wok).
Fig. 23. Oryctocephalus, species undetermined No. 2..........2eeeeeeeee 193
Cranidium in shale, X 2; U.S.N.M. No. 116089a (loc. Wok).
Fig. 24. Oryctocephalus, species undetermined No. 1...........0cceeeeees 193

Cranidium in limestone, * 3; U.S.N.M. No. 116090 (loc. W16j).

Pirate 10. Trilobites of the Wenkchemnia-Stephenaspis zone

Figs. 1-6. Stephenaspis bispinosa Rasetti, new genus, new species.......... 181
1, Entire shield in shale, & 1.2; U.S.N.M. No. 116092a, paratype. 2,
Shield in shale, X 1.5; U.S.N.M. No. 116091, holotype. 3, 4, Cranidia
in shale, X 2; U.S.N.M. No. 116092b-c, paratypes. 5, 6, Pygidia in
shale, X 2; U.S.N.M. No. 116092d-e, paratypes (loc. Wok).
PIES. -7-0. SHEPNENASPUS! CL...S. (OIUSPUIOSG: KRASEUN ciciarin det aie sine 1 a eeiatd ses aoa 182
Three pygidia, preserved in limestone, & 3; U.S.N.M. No. 116003
(loc. W16j).

Enea TOs WOndeterimineds py eid N One scene scree eereisintesietsre aie oreiocineter ete 187
Pygidium in limestone, X 3; U.S.N.M. No. 116094 (loc. W16j).
Figs, 11-15. Amecephalus agnesensis (Walcott) ...........0..ceseesseeee 207

11, Shield flattened in shale, X 2; U.S.N.M. No. 58363, holotype (loc.
U.S.N.M. 35m). 12, Shield in shale with well-preserved thorax and
pygidium, *1; U.S.N.M. No. 116095, plesiotype (loc. W3b). 13,
Cephalon and thorax of small individual incompletely flattened in
shale, X 3; U.S.N.M. No. 116096, plesiotype (loc. U.S.N.M. 35m).
14, Cephalon and thorax flattened in shale, 1.5; U.S.N.M. No.
116097a, plesiotype (loc. Wok). 15, Pygidium in shale, with a small
cranidium of Syspacephalus gregarius, X 3; U.S.N.M. No. 116097b,
plesiotype (loc. Wok).

PuatTe 11. Trilobites of the Wenkchemnia-Stephenaspis zone

Figs. 1-3. Wenkchemnia walcotti Rasetti, new genus, new species......... 184
1, Shield partly flattened in shale, X 2; U.S.N.M. No. 116099a, paratype
(loc. U.S.N.M. 35m). 2, Cast from the external mold of a shield
in shale, 2; U.S.N.M. No. 116098, holotype (loc. W3b). 3,
Cranidium with left free cheek in shale, X 2; U.S.N.M. No. 116100,
paratype (loc. W3b).
Figs. 4-8. Wenkchemnia spinicollis Rasetti, new species...............00.- 185
4, Cast from the external mold of a shield in shale, * 3; U.S.N.M.
No. 116101, holotype( loc. Wok). 5, Shield in shale, & 1.5; U.S.N.M.
No. 116102a, paratype (loc. Wok). 6, 7, Cranidia in shale, X 1.5;
U.S.N.M. No. 116102b-c, paratypes (loc. Wok). 8, Pygidium in
shale, X 2; U.S.N.M. No. 116102d, paratype (loc. Wok).

258 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Figs. 9-15. Wenkchemnia sulcata Rasetti, new specieS..........0.cceccees
9, Shield in siliceous limestone, X 3; U.S.N.M. No. 116103, holotype
(loc. W16j”). 10, 11, Cranidia in limestone, X 3; U.S.N.M. No.
116104a-b, paratypes (loc. W16h). 12, 13, Cranidia in limestone,

X< 3; U.S.N.M. No. 116105a-b, paratypes (loc. W116i). 14, Pygidium

in limestone, X 3; U.S.N.M. No. 116105c, paratype. 15, Pygidium

in limestone, X 3; U.S.N.M. No. 116106a, paratype (loc. W16j”).

Pirate 12. Trilobites of the Wenkchemnia-Stephenaspis zone

Pigs. 1-5... Ogygopsis kota. Rominger)): inal: cltgeied ace cheat ee eee
I, Cast from the mold of cranidium and portion of thorax, 1;
U.S.N.M. No. 116107a, plesiotype (loc. W16j). 2-4, Pygidia and
portion of thorax, somewhat flattened in calcareous shale, XI;
U.S.N.M. No. 116107b-d, plesiotypes (loc. W16j). 5, Pygidium flat-
tened in siliceous shale, X 1; U.S.N.M. No. 116108, plesiotype (loc.
Wok). (See also pls. 21 and 20.)
Figs. 6-9. Poliella denticulata Rasetti, new SpecieS..........eeeccecccseees
6, Shield flattened in shale, X 1.5; U.S.N.M. No. 116109, holotype (loc.
Wok). 7, 8, Cranidia flattened in shale and somewhat compressed in
the transverse and longitudinal directions respectively, X 3; U.S.N.M.
No. 116110a-b, paratypes. 9, Thorax and pygidium in shale, X 3;
U.S.N.M. No. 116110c, paratype (loc. Wok).
Fags. 10-13: -Poleciia prima Walcott )).< saw ste.ciiuce oie anna eseioeie eee
10, Cranidium partly flattened in shale, X 3; U.S.N.M. No. 116111,
plesiotype (loc. U.S.N.M. 35m). 11, Shield flattened in shale and
slightly compressed longitudinally, X 3; U.S.N.M. No. 62623, para-
type (loc. U.S.N.M. 35m). 12, Shield flattened in shale, x 3;
U.S.N.M. No. 62624, lectotype (loc. U.S.N.M. No. 35m). 13, Shield
flattened in shale, 3; U.S.N.M. No. 116112, plesiotype (loc.
Wok).

PuiaTeE 13. Trilobites of the Plagiura-Kochaspis zone
(All the figured specimens are preserved in limestone.)

Figs. 1-4. (Cavorcella. skapia (Walcott)... :oa nescence ones on eee meee
1, Cranidium, 5; U.S.N.M. No. 116113a, plesiotype. 2-4, Dorsal,
lateral, and frontal views of cranidium, * 5; U.S.N.M. No. 116113b,
plesiotype (loc. W4c).
Figs. 5-8. Kochiella? maxeyi Rasetti, new specieS..........ceccccccceeecs
5-7, Dorsal, lateral, and frontal views of cranidium, <2; U.S.N.M.
No. 116114, holotype. 8, Cast from the external mold of cranidium,
X 2; U.S.N.M. No. 116115a, paratype (loc. W20d).
Pip..o.. Kochella? cf. Kamareyy Rasetticas, go. osteo bees eee ee pate tee
Cranidium, X 2.5; U.S.N.M. No. 116117 (loc. W7f).
Bigs, 10-16. Plagtura.cercops '(Walcott)/22 46. aoe cree ae nies nee ee oe see
10, 11, Frontal and dorsal views of cranidium, * 4; U.S.N.M. No.
116118a, plesiotype (loc. W4d). 12, Cranidium, * 5; U.S.N.M. No.
116118b, plesiotype (loc. W4d). 13, Cranidium, X 3; U.S.N.M. No.

IQI

172

210

228

229

237

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 259

Page
64378, holotype (loc. U.S.N.M. 63c). 14, Exfoliated cranidium, x 1;
U.S.N.M. No. 64377, paratype (loc. U.S.N.M. 63c). 15, Pygidium,
xX 2; U.S.N.M. No. 116119a, plesiotype (loc. W20d). 16, Cranidium,
x8; U.S.N.M. No. 64382, holotype of Ptychoparia cleadas (loc.
U.S.N.M. 57s).
Figs. 17-22. Schistometopus convexus Rasetti, new species.............06- 238
17, Cranidium, X 4; U.S.N.M. No. 116121a, paratype (loc. W7f). 18,
Cranidium, X 5; U.S.N.M. No. 11612Ib, paratype (loc. W7f). 10,
Cranidium, X 2; U.S.N.M. No. 116122a, paratype (loc. W20d). 20-
22, Dorsal, lateral, and frontal views of cranidium, & 2; U.S.N.M.
No. 116120, holotype (loc. W2od).

PiaTeE 14. Trilobites of the Plagiura-Kochaspis zone
(All the figured specimens are preserved in limestone.)

Figs. 1-3. Schistometopus collaris Rasetti, new species............eeeeeees 239
Dorsal, frontal, and lateral views of exfoliated cranidium, X 2;
U.S.N.M. No. 116123, holotype (loc. W2o0d).
Figs. 4-10. Kochaspis eiffelensis Rasetti, new species...........seeeeeeeeee 226
4-6, Dorsal, lateral, and frontal views of cranidium, & 2; U.S.N.M.
No. 116125, holotype (loc. W20d). 7, Cranidium, X2; U.S.N.M. No.
116126a, paratype (loc. W20d). 8, Cranidium, * 4; U.S.N.M. No.
116127a, paratype (loc. W7f). 9, Pygidium, x 3; U.S.N.M. No.
116128, paratype (loc. U.S.N.M. 38k). 10, Pygidium, X 2; U.S.N.M.
No. 116126b, paratype (loc. W20d).
Figs. 11-14. Onchocephalus fieldensis Rasetti, new species................ 232
11-13, Dorsal, lateral, and frontal views of cranidium, * 5; U.S.N.M.
No. 116129, holotype. 14, Cranidium, 5; U.S.N.M. No. 1161302,
paratype (loc. W7f).
Figs. 15-17. Onchocephalus depressus Rasetti, new species.............05 233
15, 16, Dorsal and frontal views of cranidium, 6; U.S.N.M. No.
116131, holotype. 17, Cranidium, X 6; U.S.N.M. No. 116132, para-
type (loc. W7g).

Fig. 18. Onchocephalus sublaevis Rasetti, new species..........eeeeeeeeee 234
Cranidium, X 4; U.S.N.M. No. 116133, holotype (loc. W7g).
Figs. 19-23. Onchocephalus maior Rasetti, new species..........e..eeeeeee 234

19-21, Lateral, dorsal, and frontal views of cranidium, * 2.5; U.S.N.M.
No. 116134, holotype. 22, 23, Cranidia, 2.5; U.S.N.M. No.
116135a-b, paratypes (loc. W7f).

Puate 15. Trilobites of the Plagiwra-Kochaspis zone
(All the figured specimens are preserved in limestone unless otherwise indicated.)

Figs. 1-8. Fieldaspis furcata Rasetti, new genus, new species.............- 159
1, Articulated shield, X 2; U.S.N.M. No. 116136, holotype. 2, 3,
Dorsal and lateral views of cranidium, * 2; U.S.N.M. No. 1161372,
paratype. 4, Cranidium, x2; U.S.N.M. No. 116137b, paratype.
5, Pygidium, X 2; U.S.N.M. No. 116137c, paratype. 6, 7, Dorsal
and lateral views of pygidium, X 2; U.S.N.M. No. 116137d, paratype.
8, Hypostoma, & 2; U.S.N.M. No. 116137e, paratype (loc. W7f).

260 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Page
Fig. 9. Oryctocephalites resseri Rasetti, new species...........--e.eeceece 196
Cranidium, < 4; U.S.N.M. No. 116139, holotype (loc. W7f).
Figs..10,.11..ndetermined pysidiuny Noid scstonn «05 eee eee 187
Two specimens, X 2.5; U.S.N.M. No. 116141a-b (loc. W4d).
Figs. 12-20. Amecephalus cleora (Walcott)........2sesacevcvecddcwacksece 208

12, Shield flattened in siliceous shale, 1; U.S.N.M. No. 1161422,
plesiotype (loc. W2od). 13, 14, Dorsolateral and dorsal views of
cranidium, X 4; U.S.N.M. No. 116143a, plesiotype (loc. W4c). 15,
16, Dorsal and lateral views of cranidium, 4; U.S.N.M. No.
116143b, plesiotype (loc. W4d). 17, Cranidium, & 4; U.S.N.M. No.
116143c, plesiotype (loc. W4d). 18, Exfoliated cranidium, 1;
U.S.N.M. No. 116142b, plesiotype (loc. W20d). 19, 20, Cranidia,
x 2.5; U.S.N.M. No. 116143d-e, plesiotypes (loc. W4c’).

Bigs. 21-23. Caborcella rara Rasetti, mew speciés...sc'< «: 412 elena a Sone 211

21, 22, Dorsal and frontal views, & 3, and 23, lateral view, XX 2.5, of
cranidium; U.S.N.M. No. 116144, holotype (loc. W7g).

PLATE 16. Trilobites of the Plagiura-Kochaspis zone
(All the figured specimens are preserved in limestone.)

Figs. 1-7. Fieldaspis bilobata Rasetti, new specieS...........-..sceeceeeee 161
1, Cranidium, X 2; U.S.N.M. No. 116147a, paratype. 2, 3, Dorsal and
lateral views of exfoliated cranidium, & 2; U.S.N.M. No. 116147b,
paratype. 4, 5, Pygidia, X 2; U.S.N.M. No. 116147c-d, paratypes.
6, 7, Lateral and dorsal views of pygidium, & 2; U.S.N.M. No.
116146, holotype (loc. W2o0d).
Bigse80. ieldaspis cia OtloCatamasettine ei aaeio cei ic eter erate 162
8, Exfoliated pygidium, 1.5. 9, Cranidium, 1.5; U.S.N.M. No.
116148a-b (loc. Wé6d).
Figs. 10-18. Fieldaspis superba Rasetti, new specieS.........ecceececceces 162
10-12, Dorsal, frontal, and lateral views of cranidium, * 1; U.S.N.M.
No. 116150a, paratype. 13, 14, Dorsal and dorsolateral views of
cranidium, X 2; U.S.N.M. No. 116150b, paratype. 15, Cranidium,
<2; U.S.N.M. No. 116150c, paratype. 16, Thorax and pygidium,
xX 2; U.S.N.M. No. 116149, holotype. 17, Pygidium and hypostoma,
x 1.5; U.S.N.M. No. 116150d, paratype. 18, Cast of the external
mold of pygidium, * 1; U.S.N.M. No. 116150e, paratype (loc. W7g).

Puate 17. Trilobites of the Albertella zone

Pigs. 1-0; edlbertella® boscwontht. Walcott eeeeer. see eee acenee eee eee 149
1, Shield partly flattened in shale, X 2; U.S.N.M. No. 53416, lecto-
type (loc. U.S.N.M. 35c). 2, Shield partly flattened in shale, X 1.5;
U.S.N.M. No. 63762, plesiotype of Albertella bosworthi, holotype of
Albertella similaris (loc. U.S.N.M. 63j). 3, Shield partly flattened
in shale, X 1.5; U.S.N.M. No. 116151, plesiotype (loc. U.S.N.M.
63m). 4, Cast from the external mold of a shield in shale, X 2;
U.S.N.M. No. 116152, plesiotype (loc. C5m). 5, Cranidium flattened
in shale, X 2; U.S.N.M. No. 116153a, plesiotype (loc. C4m). 6, 7,
Dorsal and lateral views of cranidium in limestone, X 2; U.S.N.M.
No. 116153b, plesiotype (loc. C4m). 8, Cranidium in limestone,

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 261

Page
x2; U.S.N.M. No. 116153c, plesiotype (loc. C4m). 9, Pygidium
in limestone, X 2; U.S.N.M. No. 116153d, plesiotype (loc. C4m).
Figs. 10-15. Albertella declivis Rasetti, new species..............00eeeees 150
10-12, Dorsal, frontal, and lateral views of cranidium, * 2; U.S.N.M.
No. 116154, holotype. 13-15, Lateral, posterior, and dorsal views of
pygidium, X 2; U.S.N.M. No. 116155a, paratype (loc. C15n).

Puiate 18. Trilobites of the Albertella zone

gcse si-7., eAlbertella matidas IRESSEL: ota fleets cose scycie cisions emcioe teeters I5I
I, Shield flattened in shale, X 3; U.S.N.M. No. 116156, plesiotype
(loc. U.S.N.M. 35c). 2, Cranidium flattened in shale, X 2; U.S.N.M.
No. 116157, plesiotype (loc. C4m). 3, Shield flattened in shale,
X 3; U.S.N.M. No. 63766, holotype (loc. U.S.N.M. 63j). 4, 5,
Dorsal and lateral views of pygidium in limestone, X 3; U.S.N.M.
No. 116158a, plesiotype (loc. C15m). 6, 7, Dorsal and lateral views
of cranidium in limestone, X 3; U.S.N.M. No. 116158b, plesiotype
(loc. C15m).
Figs. 8-17. Albertella limbata Rasetti, new species..................000 154
8-10 Dorsal, frontal, and lateral views of cranidium, & 3; U.S.N.M.
No. 116159, holotype. 11-13, Dorsal, lateral, and posterior views
of pygidium, X 3; U.S.N.M. No. 116160a, paratype. 14-16, Three
cranidia, X 3; U.S.N.M. No. 116160b-d, paratypes. 17, Pygidium,
X 3; U.S.N.M. No. 116160e, paratype (loc. Cr5n). All specimens
preserved in limestone.
Figs. 18-21. Albertella stenorhachis Rasetti, new species...............00- 155
18-20, Dorsal, posterior, and dorsolateral views of pygidium, XX 3;
U.S.N.M. No. 116161, holotype. 21, Pygidium, *K 3; U.S.N.M. No.
116162a, paratype (loc. C15n). All specimens preserved in limestone.

PLaTE 19. Trilobites of the Albertella zone

Figs. 1-8. Alberiella microps Rasetti, new SP€Ci€S..........cecseeseccccces 153
1, Shield partly flattened in shale, X 3; U.S.N.M. No. 116163, holo-
type (loc. U.S.N.M. 63m). 2, Cranidium flattened in shale, x 3;
U.S.N.M. No. 116164, paratype (loc. C5m). 3, Pygidium in shale,
<3; U.S.N.M. No. 116165, paratype (loc. C4m). 4, Cranidium in
limestone, X 3; U.S.N.M. No. 116166a, paratype (loc. C15m).
5, 6, Lateral and dorsal views of cranidium in limestone, xX 3;
U.S.N.M. No. 116166b, paratype (loc. C15m). 7, 8, Dorsal and
lateral views of pygidium in limestone, & 3; U.S.N.M. No. 116166c,
paratype (loc. C15m).
PissuO-10. Plarmiganioa: rossensts (Walcott) mc ies 6 cuess venwcsawcad tears as 177
9, Shield flattened in shale, X 1; U.S.N.M. No. 63733, lectotype (loc.
U.S.N.M. 63j). 10, Cranidium flattened in shale, * 1.5; U.S.N.M.
No. 63729, paratype (loc. U.S.N.M. 63j). 11, Pygidium in shale,
<2; U.S.N.M. No. 63732, paratype (loc. U.S.N.M. 63j). 12, 13,
Top and side views of pygidium in limestone, K 2; U.S.N.M. No.
63735, described by Walcott as Bathyuriscus cf. rossensis (loc.
U.S.N.M. 63m’). 14-16, Dorsal, frontal, and lateral views of cranid-
ium in limestone, 2; U.S.N.M. No. 116167, plesiotype (loc. C4m).

262 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Figs. 17-19. Kochina macrops Rasetti, new SPeCieS..........cecececcceses
Dorsal, dorsolateral, and frontal views of cranidium in limestone, < 3;
U.S.N.M. No. 116168, holotype (loc, C15n).
Pigsi20-23) Kochina americana’ (Walcott) sees sees case coe ciee eee ste ion eee
20, Cranidium flattened in shale, & 1; U.S.N.M. No. 63749, lectotype
of Kochina bosworthensis (loc. U.S.N.M. 35c). 21, 22, Frontal and
dorsal views of cranidium in limestone, X 2; U.S.N.M. No. 116170,
plesiotype (loc. U.S.N.M. 63m’). 23, Cast from the mold of a
shield flattened in shale, 1; U.S.N.M. No. 116171, plesiotype
(loc. C4m).

PLATE 20. Trilobites of the Albertella zone

Figs. 1-9. Ptarmiganoides bowensis Rasetti, new genus, new species.......
1-3, Dorsal, lateral, and frontal views of cranidium, X 2; U.S.N.M.
No. 116173a, paratype. 4, Cranidium, X 2; U.S.N.M. No. 116173b,
paratype. 5-7, Dorsal, lateral, and posterior views of pygidium, X 3;
U.S.N.M. No. 116172, holotype. 8, 9, Pygidia, * 3; U.S.N.M. No.
116173c-d, paratypes. All specimens in limestone (loc. Cr5n).
Bigs. 10-13) VWanumemella norita Waleott..+ acces aes coe cee cee cee ee
10, Shield partly flattened in shale, X 3; U.S.N.M. No. 61728, holo-
type (loc. U.S.N.M. 35c). 11, Cranidium in limestone, X 3; U.S.N.M.
No. 116174a, plesiotype (loc. C15m). 12, Pygidium in limestone,
<3; U.S.N.M. No. 116174b, plesiotype (loc. C15m). 13, Shield
partly flattened in shale, & 3; U.S.N.M. No. 116175, plesiotype (loc.
U.S.N.M. 63w).
Bigsu4-10; Vescellarstacor @Walcott) saeco see cece eeereee
14, 15, Lateral and dorsal views of cranidium in limestone, X 3;
U.S.N.M. No. 116176a, plesiotype (loc. U.S.N.M. 63m’). 16, Ex-
foliated cranidium in limestone, X 2; U.S.N.M. No. 116176b, plesio-
type (loc. U.S.N.M. 63m’). 17, Cast from the mold of shield
flattened in shale, X 2; U.S.N.M. No. 116177, plesiotype (loc. C5m).
18, Shield flattened in shale, X 2; U.S.N.M. No. 61720a, paratype
(loc. U.S.N.M. 35c). 10, Shield flattened in shale and lacking the
pygidium, X 1; U.S.N.M. No. 61729, holotype (loc. U.S.N.M. 35c).

PuateE 21. Trilobites of the Albertella (?) zone

Figs, 1-3. (Ogygopsis felores Ciomingper yes) abe lee cetes «see tee. eee
1, Weathered shield, 1; U.S.N.M. No. 116178a, plesiotype. 2,
Pygidium lacking the test, X 1; U.S.N.M. No. 116178b, plesiotype.
3, Cranidium, <1; U.S.N.M. No. 116178c, plesiotype. All specimens
partly flattened and distorted in silty limestone (loc. Coh). (See
also pls. 12 and 29.)
Fig. 4. Ogygopsis spinulosa Rasetti, new species...........0..cccceuseee
4, Pygidium lacking the test, 1; U.S.N.M. No. 116179, holotype.
Preserved in silty limestone (loc. Coh).
Figs. 5-8. Zacanthoides, species undetermined.............0ccccceccescce
5-7, Cranidia, X 2; U.S.N.M. No. 116180a-c. 8, Poorly preserved articu-
lated shield, 2; U.S.N.M. No. 116180d. All specimens partly
flattened and distorted in silty limestone (loc. Coh).

220

179

183

231

IQI

192

140

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

Figs. 9-14. Yohoaspis pachycephala Rasetti, new genus, new species.......
9, Cranidium flattened and weathered in siliceous shale, * 1; U.S.N.M.
No. 116182a, paratype (loc. Coh). 10-12, Dorsal, lateral, and
frontal views of cranidium in limestone, possibly somewhat com-
pressed transversely, X 2; U.S.N.M. No. 116181, holotype (loc. Coh).
13, 14, Dorsal and frontal views of distorted cranidium in limestone,
X 2; U.S.N.M. No. 116183, paratype (loc. Coj).
Figs. 15-17. Chancia latigena Rasetti, new species..........ecceececcecves
15, Cranidium in limestone, & 2; U.S.N.M. No. 116184, holotype. 16,
Cranidium lacking the test, flattened in shale, «1.5; U.S.N.M.
No. 116185a, paratype. 17, Cranidium in limestone, X 2; U.S.N.M.
No. 116185b, paratype (loc. Coh).

Pate 22. Trilobites of the Albertella (?) zone

Figs. 1-6. Chancia bigranulosa Rasetti, new SpecieS........cccccceccecess
1-3, Dorsal, lateral, and frontal views of cranidium, distorted in lime-
stone, X 2; U.S.N.M. No. 116186, holotype (loc. Coj’). 4, Cranidium
in limestone, X 2; U.S.N.M. No. 116187a, paratype (loc. Coj).
5, 6, Dorsal and lateral views of cranidium in limestone, X 2;
U.S.N.M. No. 116187b, paratype (loc. Coj).
Figs. 7-10. Syspacephalus tardus Rasetti, new specieS.............eceeeees
7, Cranidium, X 3; U.S.N.M. No. 116188, holotype. 8, Cranidium, * 3;
U.S.N.M. No. 116189a, paratype. 9, 10, Lateral and dorsal views of
cranidium, X 3; U.S.N.M. No. 116189b, paratype. All specimens pre-
served in silty limestone (loc. Coh).
Higse lin ie tnabaskia) species undetenminedss..s47 ese se cee ae eee: ec
11, Cranidium doubtfully assigned to the genus, * 3; U.S.N.M. No.
116190. 12, Pygidium, X 3; U.S.N.M. No. 116191. Specimens pre-
served in limestone (loc. Coj’).
Rigs. 13. 14; agenda, Species GndeterMiINeEd. 2... ile. slewdec sees cee es cmreses
13, Cranidium, 8; U.S.N.M. No. 116192a. 14, Pygidium, x 8;
U.S.N.M. No. 116192b. Specimens preserved in limestone (loc. Coh).
Figs. 15, 16. Bathyuriscus? species undetermined..........0..ccecccceees
Cranidia poorly preserved in limestone, X 3; U.S.N.M. No. 116193a-b
(loc. Cgj).
fags. 17-10. Poltelia, species. tadetermuined + NO,) 15 <0 e e ays oe sence
17, Pygidium lacking the test, * 3. 18, 19, Cranidia lacking the test,
<3; U.S.N.M. No. 116194a-c. Specimens preserved in silty lime-
stone (loc. Coh).
Figs. 20, 21. Poliella, species undetermined No. 2...........c0eesseeecees
Pygidia poorly preserved in limestone, XK 3; U.S.N.M. No, 116195a-b
(lac; Coy").
Figs. 22-27. Zacanthoides sexdentatus Rasetti, new species.............06-
22-24, Cranidia, * 3; U.S.N.M. No. 116197a-c, paratypes. 25, Pygid-
ium, X 3; U.S.N.M. No. 116196, holotype. 26, 27, Pygidia, x 3;
U.S.N.M. No. 116197d-e, paratypes. All specimens preserved in lime-
stone and distorted in various directions (loc. Coj).

212

213

247

156

138

158

174

175

141

264 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Page
PLATE 23. Trilobites of the Glossopleura zone
(All the figured specimens are preserved in limestone.)
Figs. 1-5. Glossopleura merlinensis Rasetti, new species..............44- 167

I, 2, Dorsal and lateral views of cranidium, X 3; U.S.N.M. No. 116199a,
paratype. 3, Cranidium, * 3; U.S.N.M. No. 116199b, paratype. 4, 5,
Dorsal and posterior views of pygidium, & 3; U.S.N.M. No. 116198,
holotype (loc. C13r).

Figs. 6-10. Glossopleura skokiensis Rasetti, new species............++eeee0% 166

6, 7, Lateral and dorsal views of cranidium, 3; U.S.N.M. No.
116201a, paratype. 8, Free cheek, K 3; U.S.N.M. No. 116201b, para-
type. 9, 10, Posterior and dorsal views of pygidium, X 3; U.S.N.M.
No. 116200, holotype (loc. Cr3r).

Figs. 11-15. Polypleuraspis insignis Rasetti, new species..............++-- 176

11, Pygidium, X 2; U.S.N.M. No. 116203a, paratype. 12, Cranidium,

xX 3; U.S.N.M. No. 116203b, paratype. 13, Slab of limestone with
numerous cranidia and pygidia, & 2; U.S.N.M. No. 116203c, paratypes.
14, 15, Lateral and dorsal views of cast from the external mold of a
perfect shield, X 3; U.S.N.M. No. 116202, holotype (loc. S2r1b).

Puiate 24. Trilobites of the Glossopleura zone
(All the figured specimens are preserved in limestone.)

Figs. 1-6. Glossopleura boccar (Walcott)... ....0csceeesececsnenenaanee 164
1, Partly exfoliated cranidium, < 2; U.S.N.M. No. 62704, syntype, holo-
type of Glossopleura bosworthensis Resser (loc. U.S.N.M. 57g). 2,
Partly exfoliated pygidium, X 2; U.S.N.M. No. 62708, syntype, holo-
type of Glossopleura nitida Resser (loc. U.S.N.M. 57g). 3, Cranid-
ium, X 2; U.S.N.M. No. 116204a, plesiotype (loc. U.S.N.M. 57u).
4, 5, Dorsal and posterior views of pygidium, X 2; U.S.N.M. No.
116204b, plesiotype (loc. U.S.N.M. 57u). 6, Partly exfoliated pygid-
ium, X 2; U.S.N.M. No. 116205, plesiotype (loc. S5a).
Figs. 7, 8. Glossopleura stenorhachis Rasetti, new specieS...........eeeeee 165
7, Pygidium, X 2; U.S.N.M. No. 116206, holotype. 8, Pygidium, X 2;
U.S.N.M. No. 116207a, paratype (loc. S3b).
Figs. 0-12, 18-1 Glossopleura mckéet RESSere. <oi.<ns siete sie © serge eran 165
0, 10, Cranidiay x2. 10, (12;" Pygidia,)°X</2. 1/18, Free eneck see,
U.S.N.M. No. 116208a-e, plesiotypes (loc. S3b).

Fig. 13. Glossopleura, species undetermined, ... 02.0.2... . tse 0s ee cae 168
Cranidium, X 3; U.S.N.M. No. 116211 (loc. C1r3r).
Figs. 14-17. Glossopleura templensis Rasetti, new species................ 164

14, Cranidium, X 3; U.S.N.M. No. 116210a, paratype. 15, 16, Pygidia,
xX 2; U.S.N.M. No. 116210b-c, paratypes. 17, Exfoliated shield,
xX 2; U.S.N.M. No. 116209, holotype (loc. S21b).

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI

Piate 25. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

Figs. 1-5. Triplagnostus burgessensis Rasetti, new species........ ...ee0
1, Shield partly flattened in siliceous shale, & 5; U.S.N.M. No. 116212,
holotype. 2-5, Shields partly flattened in siliceous shale, X 5;
U.S.N.M. No. 116213a-d, paratypes (loc. U.S.N.M. 35k).
ies. G-101, Paget Oogles” Walcottim: « a. ceuek ven «hues ate seek wens
6, 9, Shields in calcareous shale preserving the convexity, <5. 7, 8,
Shields partly flattened in siliceous shale, 5. 10, Shield in cal-
careous shale preserving the convexity, X 6; U.S.N.M. No. 116214
a-e, plesiotypes (loc. U.S.N.M. 35k).
mies! 11-14, Peronopss montis (Matthew) ive. c2 ei eco. ccce cs ed eee ees oes
11, Shield partly flattened in siliceous shale, X 5; U.S.N.M. No. 116215,
plesiotype (loc. U.S.N.M. 35k). 12-14, Shields flattened in siliceous
shale, X 5; U.S.N.M. No. 116216a-c, plesiotypes (loc. U.S.N.M. 14s).
Bape 15. Lianouria’ Gloriosa” Walcott. (out), sok. co treme sae eres eee sists eres
Shield flattened in siliceous shale, X 3; U.S.N.M. No. 61724, syntype
(loc. U.S.N.M. 35k).
Fig. 16. Solenopleurella, species undetermined No. 2..........-.2s0eee0e:
Weathered shield in argillaceous limestone, 3; U.S.N.M. No.
116217a (loc. Site).
Figs. 17, 18. Solenopleurella, species undetermined No. 1...........--+-00:
17, Cranidium in siliceous shale, * 3; U.S.N.M. No. 116218a (loc.
Siig). 18, Cranidium in limestone, X 5; U.S.N.M. No. 116219a (loc.
S6g).

PLATE 26. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

Bigs. 1-3. Oryctocephalus burgessensts ReSSEr.. ... 2.60000 .e cess cccceness
1, Shield flattened in siliceous shale, * 5; U.S.N.M. No. 1162202,
plesiotype. 2, Shield flattened in siliceous shale, & 5; U.S.N.M. No.
96487, holotype. 3, Shield flattened in siliceous shale, 3; U.S.N.M.
No. 116220b, plesiotype (loc. U.S.N.M. 35k).
Figs. 4, 5. Oryctocephalus matthewi Rasetti, new species.............200-
4, Shield flattened in siliceous shale, X 5; U.S.N.M. No. 116222a, para-
type. 5, Cast from external mold of shield flattened in siliceous shale,
<5; U.S.N.M. No. 116221, holotype (loc. U.S.N.M. 35k).
Pig. 6. Elrathina brevifrons Rasetti, new SPeCieS...........scecsseccenees
Shield partly flattened in argillaceous limestone, * 3; U.S.N.M. No.
116223, holotype (loc. Stte).
Figs. 7-9. Elrathina cordillerae (Rominger) ..........cccccsscccscseccces
7, Cast from the external mold of shield partly flattened in siliceous
shale, X 3; U.S.N.M. No. 116225, plesiotype (loc. U.S.N.M. 35k).
8, Shield partly flattened in siliceous shale, * 3; U.S.N.M. No.
17831, plesiotype (loc. U.S.N.M. 14s). 9, Shield partly flattened in
siliceous shale, X 3; U.S.N.M. No. 116226, plesiotype (loc. U.S.N.M.
14s).

137

134

197

241

241

104

105

223

221

266 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

PLATE 27. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

(All the figured specimens are preserved in siliceous shale.)

Figs. .1=3.' Olenoides serratus .(Rominger) (i205 .ii5.c%ai see's oe bee peewee oF 189
1, Cast from the external mold of partly flattened shield, & 3; U.S.N.M.
No. 116227, plesiotype (loc. U.S.N.M. 14s). 2, Cast from the ex-
ternal mold of partly flattened shield, X 3; U.S.N.M. No. 116228a,
plesiotype (loc. U.S.N.M. 35k). 3, Cast from the external mold
of partly flattened pygidium, « 2; U.S.N.M. No. 116228b, plesiotype
(loc. U.S.N.M. 35k).
Bigs: 4-7: Kootenta: dawson (Walcott) -. 6. 34 a fe. veces Snaeaiene toe 189
4, Partly flattened, exfoliated shield, X 3; U.S.N.M. No. 116229, plesio-
type (loc. U.S.N.M. 14s). 5, Flattened, exfoliated shield, 1;
U.S.N.M. No. 108495, syntype (loc. U.S.N.M. 14s). 6, 7, Lateral ~
and dorsal views of cranidium preserving the convexity, XX 2;
U.S.N.M. No. 108495, syntype (loc. U.S.N.M. 14s).
Figs. 8-10. Zacanthoides romingeri Resser........0cccccceccccescececvce 143
8, Flattened, exfoliated shield, & 1.5; U.S.N.M. No. 102324, plesiotype.
9, Partly flattened cranidium, <2; U.S.N.M. No. 116230, plesio-
type. 10, Partly flattened, exfoliated pygidium, 2; U.S.N.M. No.
102324, plesiotype (loc. U.S.N.M. 14s).
Fig. 11. Parkaspis decamera Rasetti, new species..........0esceseeeeeees 171
Poorly preserved, flattened shield, 2.5; U.S.N.M. No. 116231, holo-
type (loc. U.S.N.M. 35k).

PLATE 28. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

(All the figured specimens are preserved in siliceous shale.)

Fig. 1. Burlingsa hectors Walcott... sossk cca eas ase chee spend. ane 138
Shield, X 8; U.S.N.M. No. 53418, lectotype (loc. U.S.N.M. 14s).
Figs. 2, 3. Bathyuriscus rotundatus CRominger) ........0.+..s.+-0ceeeees 158

2, Partly flattened shield, X 2. 3, Partly flattened small shield, X 3;
U.S.N.M. No. 116232a-b, plesiotypes (loc. S8d).
Pigs. 4-6. Bonnaspis stephenensts iC Waleott) cack oatseee 12+ sce sews eee ee 188
~ 4, Cast from the external mold of partly flattened shield, X 3; U.S.N.M.
No. 61731, holotype (loc. U.S.N.M. 14s). 5, Exfoliated cranidium
with five thoracic segments, X 3; U.S.N.M. No. 116233a, plesiotype
(loc. S8d). 6, Partly weathered shield preserving most of the test
and the original convexity, X 3; U.S.N.M. No. 116233b, plesiotype
(loc. S8d).
Bisse7 8. Klotetellaconnatan GW Alcott actita clic clelsl alesteretere eitereiie eens 168
Casts from the external molds of two shields partly flattened in shale,

X 3; U.S.N.M. No. 116235a-b, plesiotypes (loc. U.S.N.M. 14s).
Bigssto-11. Kodtentaburgessensis: IRESSerom eens cele tes ae ete ae oeinie ete 189
9, Exfoliated pygidium, X 2; U.S.N.M. No. 116234a, plesiotype. 10,
Cast of the interior of pygidium, X 3; U.S.N.M. No. 116234b, plesio-
type. 11, Partly flattened, exfoliated shield, X 1.5; U.S.N.M. No.

65511, holotype (loc. U.S.N.M. 35k).

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 267

PLATE 29. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

(All the figured specimens are preserved in siliceous shale.)

Bigs 1=3 (Ghanciarpalissert CW alcott) i... Sh Pee ate ees cee ccal 214
Casts from the molds of 3 shields flattened in shale, * 1.5; the two
complete individuals possess 20 thoracic segments; U.S.N.M. No.
116236a-c, plesiotypes (loc. U.S.N.M. 35k).

Bigs A, 5. OFVCEOCePAAIUS TEYNGIGSt REEG visi. ¢ o2ciele tiscd bv v dae's Hem weiner 193
4, Flattened shield, «2.5; U.S.N.M. No. 116237, plesiotype (loc.
U.S.N.M. 14s). 5, Flattened shield, 2.5; U.S.N.M. No. 116238,
plesiotype (loc. U.S.N.M. 35k).

Pigs16-0. JOgvaopsts kiaist (Rominger))..o. 2... Bales o8b ae nda Sake sae onic IQI
6, Flattened hypostoma, 2; U.S.N.M. No. 116239a, plesiotype. 7,
Small, flattened shield, 2; U.S.N.M. No. 24040, plesiotype. 8,
Flattened shield, X11; U.S.N.M. No. 116239b, plesiotype (loc.
U.S.N.M. 14s). (See also pls. 12 and 21).

PLATE 30. Trilobites of the Bathyuriscus-Elrathina zone
(Burgess and Ogygopsis shales)

(All the figured specimens are preserved in siliceous shale unless otherwise indicated.)

Hiss, 2. irate permulta (Walcott). s.ksdes dante da sneonde sds ailion. 220
Casts from the external molds of two partly flattened shields, >< 2;
U.S.N.M. No. 116240a-b, plesiotypes (loc. U.S.N.M. 35k).

Figs. 3-5. Alokistocarella fieldensis Rasetti, new species........-.++eeeee 206
3, 5, Partly flattened shields in argillaceous limestone, X 3; U.S.N.M.
No. 116242a-b, paratypes. 4, Partly flattened shield in argillaceous
limestone, X 3; U.S.N.M. No. 116241, holotype (loc. Site).

Figs. 6-8. Ehmaniella waptaensis Rasetti, new species..........-.e00eeeee 219
6, Cast from the external mold of small, partly flattened shield, X 5;
U.S.N.M. No. 116244a, paratype. 7, Partly flattened shield, < 5;
U.S.N.M. No. 116244b, paratype. 8, Partly flattened cephalon with
several thoracic segments, <5; U.S.N.M. No. 116243, holotype

(loc. U.S.N.M. 35k).

Figs. 9-16. Ehmaniella burgessensis Rasetti, new sSpecieS..........seeeeees 217

9, Partly exfoliated, flattened shield, X 2; U.S.N.M. No. 116245, holo-
type (loc. U.S.N.M. 35k). 10, Partly exfoliated, flattened shield,
X< 3; U.S.N.M. No. 116246a, paratype (loc. U.S.N.M. 35k). 11, 14,
Pygidia, X 2; U.S.N.M. No. 116247a-b, paratypes (loc. Siig). 12,
13, Cranidia, X 2; U.S.N.M. No. 116247c-d, paratypes (loc. Siig).
15, Exfoliated pygidium in limestone, X 3; U.S.N.M. No. 116248a,
paratype (loc. S6g). 16, Exfoliated cranidium in limestone, X 2;
U.S.N.M. No. 116248b, paratype (loc. S6g).

268 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

PLATE 31. Trilobites of the Bathyuriscus-Elrathina zone
(All the figured specimens are preserved in limestone.)

Figs: 1-6. Bathyuriscus adacus Walcott) occ s a. elniet ttetlnt's © melseants
1, Cast from the external mold of shield, & 2.5; U.S.N.M. No. 116249,
plesiotype (loc. S10). 2, 3, Dorsal and lateral views of cranidium pre-
serving the full convexity, X 3; U.S.N.M. No. 116250a, plesiotype
(loc. Sok). 4, Small, unflattened pygidium, x 3; U.S.N.M. No.
116250b, plesiotype (loc. Sok). 5, Pygidium, somewhat flattened and
weathered, X 1; U.S.N.M. No. 116250c, plesiotype (loc. Sok). 6,
Flattened and slightly weathered shield, X 2; U.S.N.M. No. 116250d,
plesiotype (loc. Sok).
Figs. 7-10. Parkaspis endecamera Rasetti, new genus, new species.........
7, Partly flattened and slightly distorted shield, 1; U.S.N.M. No.
116251, holotype. 8, Pygidium of the same specimen, X 2. 9, 10,
Partly flattened, distorted cranidia, K 2; U.S.N.M. No. 116252a-b,
paratypes (loc. Srol).
Figs. 11, 12. Athabaskia? parva Rasetti, new species. .:.........05-.+ser ve
11, Cranidium, * 6; U.S.N.M. No. 116253, holotype. 12, Cranidium,
<3; U.S.N.M. No. 116254, paratype (loc. Sél).
Bigs. 13-195 ehonkinellarstephensis Iobayashiw. saree ne 2 « eile eee
13, Cranidium, X 3; U.S.N.M. No. 116255a, plesiotype (loc. Sok).
14, Cast from the weathered impression of large shield, & 2; U.S.N.M.
No. 116256a, plesiotype (loc. S61). 15, Pygidium, X 2; U.S.N.M.
No. 116256b, plesiotype (loc. S61). 16-18, frontal, lateral, and
dorsal views of cast of external mold of cranidium, X 4; U.S.N.M.
No. 108497, lectotype (loc. U.S.N.M. 58j).

PLATE 32. Trilobites of the Bathyuriscus-Elrathina zone
(All the figured specimens are preserved in limestone.)

Figs. 1, 2. Zacanthoides planifrons Rasetti, new species............2eeeee.
1, Shield, X 1; U.S.N.M. No. 116258a, paratype. 2, Shield, X 1.5;
U.S.N.M. No. 116257, holotype (loc. S1o).
Figs. 3-7. Zacanthoides longipygus Rasetti, new species............00eeeee
3, Exfoliated cranidium, X 1; U.S.N.M. No. 116260a, paratype. 4, Ex-
foliated cranidium, X 2; U.S.N.M. No. 116260b, paratype. 5, 6,
Pygidia, 3; U.S.N.M. No. 116260c-d, paratypes. 7, Slightly
weathered pygidium, <3; U.S.N.M. No. 116259, holotype (loc.
S6l).
Figs. 8-10. Zacanthoides divergens Rasetti, new species...............--.
8, Entire shield, X 3; U.S.N.M. No. 116261, holotype (loc. S1ol). 9,
10, Cranidia, X 3; U.S.N.M. No. 116262a-b, paratypes (loc. S6l).
Figs. 11-14. Zacanthoides submuticus Rasetti, new species............+04
11, Small pygidium preserving the convexity, 6; U.S.N.M. No.
116263, holotype. 12, partly flattened cranidium, K 2; U.S.N.M. No.
116264a, paratype. 13, 14, Partly flattened pygidia, X 2; U.S.N.M.
No. 116264b-c, paratypes (loc. S6l).

170

156

196

144

146

143

NO. 5 MIDDLE CAMBRIAN STRATIGRAPHY—RASETTI 269

Figs. 15-18. Yuknessaspis paradoxa Rasetti, new genus, new species........

15, Cranidium and thorax, * 1; U.S.N.M. No. 116266, paratype. 16-18,

Dorsal, frontal, and lateral views of cranidium, * 2; U.S.N.M. No.
116265, holotype (loc. Srol).

PLATE 33. Trilobites of the Bathyuriscus-Elrathina zone
(All the figured specimens are preserved in limestone.)

Figs. 1-8. Peronopsis columbiensis Rasetti, new specieS..........0+.e eens
1, Shield slightly compressed transversely, & 4; U.S.N.M. No. 116267,
holotype. 2, Shield slightly compressed longitudinally, & 4; U.S.N.M.
No. 116268a, paratype. 3, 4, Dorsal and lateral views of pygidium,
x 4; U.S.N.M. No. 116268b, paratype. 5, 6, Lateral and dorsal views
of cephalon, X 4; U.S.N.M. No. 116268c, paratype. 7, 8, Pygidia,
<4; U.S.N.M. No. 116268d-e, paratypes (loc. S6k).
moseo-rn., Pagena ct. Ps, bootes’ Walcott sic ace ceicys vices ctere.n ate sce stetveeis ve
9, Cranidium, < 8; U.S.N.M. No. 116270a (loc. S61). 10, Pygidium,
<8; U.S.N.M. No. 116270b (loc. S61). 11, Pygidium, xX 8;
U.S.N.M. No. 116271a (loc. Sok).
Figs. 12-14. Pachyaspis attenuata Rasetti, new species.............0.00%
12, 13, Lateral and dorsal views of cranidium, X 2; U.S.N.M. No.
116272, holotype. 14, Cranidium, X 2; U.S.N.M. No. 116273a, para-
type (loc. Sél).
Figs. 15, 16. Chancia odarayensis Rasetti, new species..........eeeeeeees
15, Cranidium doubtfully assigned to the species, * 3; U.S.N.M. No.
116276a (loc. Sok). 16, Weathered shield lacking the pygidium,
X 2; U.S.N.M. No. 116274, holotype (loc. S61).
Figs. 17, 18. Alokistocare paranotatum Rasetti, new species..........+..-
17, Cranidium, X 3; U.S.N.M. No. 116278a, paratype. 18, Cranidium,
X 3; U.S.N.M. No. 116277, holotype (loc. Sok).
Figs. 19-22. Elrathina parallela Rasetti, new species...........eeeeeeeees
19, Several specimens, 1; U.S.N.M. No. 116280a, paratypes. 20,
Slightly weathered shield, «3; U.S.N.M. No. 116279, holotype.
21, 22, Lateral and dorsal views of cranidium, X 3; U.S.N.M. No.
116280b, paratype (loc. Sok).

Pirate 34. Trilobites of the Bathyurtscus-Elrathina zone
(All the figured specimens are preserved in limestone.)

Figs. 1, 2. Alokistocare sinuatum Rasetti, new species...........eeeeeeee
1, Cast of the external mold of cranidium, X 2; U.S.N.M. No. 116282a,
paratype. 2, Cranidium, * 2; U.S.N.M. No. 116281, holotype (loc.
Sél).
Figs. 3, 4. Alokistocare cataractense Rasetti, new species..............00:
3, Cranidium, 2; U.S.N.M. No. 116283, holotype. 4, Cranidium,
X 2; U.S.N.M. No. 116284a, paratype (loc. Srol).

134

137

235

216

203

222

204

205

270 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Figs. 5-7. Glyphaspis parkensis Rasetti, new specieS...........s2eeeeeeee 224
5, Cranidia, * 1.5; U.S.N.M. No. 116286a, paratypes. 6, Somewhat
flattened shield, & 1; U.S.N.M. No. 116285, holotype. 7, Exfoliated
pygidium somewhat compressed transversely, * 1.5; U.S.N.M. No.
116286b, paratype (loc. S1ol).
Figs. 8-10, 15. Elrathina marginalis Rasetti, new species.............-0e- 224
8, Cranidium, * 3; U.S.N.M. No. 116288a, paratype. 9, 10, Lateral and
dorsal views of cranidium, X 3; U.S.N.M. No. 116287, holotype.
15, Cranidium with the glabella broken off and portion of thorax,
x 1.5; U.S.N.M. No. 116288b, paratype (loc. S6l).
Figs. 11-14. Elrathina spinifera Rasetti, new SPeCi€S.........sssccevecease 223
11, 12, Lateral and dorsal views of cranidium, X 3; U.S.N.M. No.
116290a, paratype (loc. S6k). 13, Cranidium, * 3; U.S.N.M. No.
116289, holotype (loc. S6k). 14, Distorted cranidium, * 3; U.S.N.M.
No. 116291a, paratype (loc. Sok).

PLATES

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 126, INO. 5, Pio

SOUTH FACE OF MOUNT TEMPLE (11,636 FEET)

(See explanation of plates at end of text.)

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MIDDLE CAMBRIAN STRATA ON MOUNT STEPHEN AND PARK MOUNTAIN

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TRILOBITES OF THE BATHYURISCUS-ELRATHINA ZONE
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ELRATHINA ZONE

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TRILOBITES OF THE BATHYURISCUS-ELRATHINA ZONE

(BURGESS AND OGYGOPSIS SHALES )

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VOL. 116, NO. 5, PL. 3

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TRILOBITES OF THE BATHYURISCUS-ELRATHINA ZONE
(See explanation of plates at end of text.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 5, PL. 32

TRILOBITES OF THE BATHYURISCUS-ELRATHINA ZONE
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TRILOBITES OF THE BATHYURISCUS-ELRATHINA ZONE
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INDEX

(Figures in boldface indicate detailed descriptions.)

Acrothele, 50, 51, 89, 96 Amecephalina, 202, 203

clitus, 89 mirabilis, 203

walcotti, 93 Amecephalus, 14, 20, 23, 24, 20, 47,
Acrothyra gregaria, 103 89, 92, 114, 116, 200, 202, 207, 227,
Acrotreta, 19, 51, 52, 53, 88, 80 228

depressa, 101
Agnostus gibbus, 135

montis, 134
“Agnostus’-Bathyuriscus zone, 90
Agraulos charops, 241, 242

stator, 231

unca, 30, 31, 82
Albertella, 94, 113, 116, 120, 147, 181

agnesensis, 16, 19, 51, 86, 80, 207,
210, 245; pl. 10

cleora, 19, 20, 21, 22, 26, 36, 37,
50, 60, OI, 92, 2083 pl. 15

piochensis, 203, 208

Anomalocaris, 119

canadensis, 101%

Anomalocaris? acutangula, 101%

whiteavesi, 101

bosworthi, 12, 19, 25, 27, 94, 148, Anoria, 96, 97

149, I51, 153, 155; pl. 17

tontoensis, 97

bosworthi faunule, 69, 70, 93, 95, Antagmus, 199, 200, 201

109, II9
cimon, III, 155
declivis, 12, 94, 150, 155; pl. 17

perola, 244
pia, 236
skapta, 210

helena, 147, 148, 151 Arctomys formation, 4, 7
limbata, 12, 94, 1543 pl. 18 Asaphidae, 191
limbata faunule, 94, 97, 109 Asaphiscus, 200

microps, 12, 19, 94, 148, 1533 pl. 19

capella, 224

nitida, 12, 19, 25, 94, 148, 151,153, Ashley, G. H., 84

154; pl. 18
robsonensis, 151 Bathyuriscidae, 139
rossensis, 148, 151 Bathyuriscidea, 133, 139
similaris, 148, 149 Bathyuriscus, 34, 100, 107, 108, 116,
stenorhachis, 12, 94, 1553 pl. 18 130, 140, 157, 169, 181, 184

Albertella zone, 4, 6, 80, 93, 97, 113,
II5, IQI, 192
Allan, J. A., 68
Alokistocare, 107, 108, 114, 116, 200,
201, 202, 203
agnesensis, 207
cataractense, 35, 107, 205; pl. 34
cleora, 208
natale, 204
notatum, 204
paranotatum, 48, 106, 293; pl. 33
sinuatum, 32, 706, 204, 205; pl. 34
stator, 231

adaeus, 32, 35, 48, 73, 106, 107,
157, 160, 171; pl. 31

adaeus faunule, 74, 75, 96, 100, 105,
106, 107, 109

heydent, 157

howelli, 158

ornatus, 168

pupa, 111

rossensis, 177

cf. rossensis, 177

rotundatus, 38, 101, 102, 103, 1583
pl. 28

sp. undet., 50, 96, 1583 pl. 22

stephenense, 110 Bathyuriscus (Poliella) anteros, 172
Alokistocarella, 107, 108, 116, 206 primus, 172, 184

fieldensis, 38, 103, 206; pl. 30 Bathyuriscus (Kootenia) dawsoni, 188,

fieldensis faunule, 100, 103, 109 189

typicalis, 206, 207 Bathyuriscus-Elrathia zone, 99

271

272

Bathyuriscus-Elrathina zone, 73, 99,
115, IOI
Battus integer, 133
Bell, W. C., 81
Billingsaspis, 199
Billingsella marion, 110
Bolaspis-Glyphaspis zone, 99, 108
Bonnaspis, 107, 108, 117, 188
stephenensis, 101, 188; pl. 28
Bonnia, 13, 20, 24, 47, 54, 55, 86, 90,
D13) 116, 12S
columbensis, 82
fieldensis, 30, 31, 33, 47, 82
fieldensis faunule, 56, 82, 109
Bonnia zone, 81, 82
Bonnia-Olenellus zone, 81, 88, 90, 115
Bosworth formation, 4, 7
Bow Lake, 11
Bow trough, 5, 45, 60, 72, 73, 75
Burgess quarry, 37, 78, 102, 103
Burgess shale, 37, 97, 103, 120
Burling, L. D., 4, 54, 60, 86
Burlingia, 107, 108, 116, 138
hectori, 102, 138; pl. 28
Burlingiidae, 133, 138
Burlingioidae, 133, 138

Caborcella, 92, 94, 116, 210
arrojosensis, 210
rara, 36, 92, 2113 pl. 15
skapta, 20, 21, 22, 36, 37, 60, 91,
210; pl. 13
Cathedral formation, 4, 6, 12, 15, 18,
23, 24, 25, 27, 20, 30, 32, 43, 46, 48,
64, 93, 94, 95, 97, 118
Cathedral Mountain, 23
Chancia, 107, 108, 116, 203, 212
bigranulosa, 49, 50, 96, 2133 pl. 22
bigranulosa faunule, 96, 109, 114,
IQI
ebdome, 203, 212, 213
latigena, 50, 96, 2123 pl. 21
odarayensis, 32 48, 106, 216; pl.
33
palliseri, 102, 103, 2143 pl. 29
stenometopa, 61, 89, 216; pl. 8
Clappaspis, 100, 220
Clappaspis subzone, 99
Classification of trilobites, 132
Clavaspidella, 96, 157
kanabensis, 157
Coleoloides hectori, 93
Conocephalites, 198
cordillerae, 221
subcoronatus, 202
Conocoryphe, 1098
(Conocephalites) kingti, 220

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116

Conocoryphidae, 201
Conocoryphidea, 198
Cooper, G. A., and Arellano, A. R. V.,
82

Corynexochidae, 133, 139, 188
Corynexochidea, 139
Corynexochoidae, 133, 139
Corynexochus romingeri, 188

stephenensis, 188
Crassifimbra, 116, 201

lux, 30, 82
Crepicephalus, 200

celer, 163

chares, 110, 228, 229

liliana, 93, 115

(Loganellus) quadrans, 217
Crusoia, 240

cebes, 240

Dawsonia, 96
Deiss, C., 5, 6, 14, 20, 21, 34, 54, 57,
60, 64, 65, 68, 72, 76, 86, 87, 99
Deissella-Centropleura vermontensis
zone, 99
Dictyonina, 51, 890
Diraphora, 49, 96
Dolichometopidae, 113, 133, 139, 156,
190
Dolichometopinae, 139
Dolichometopsis, 159, 178, 179, 180
alia, 179
comis, 179
communis, 179
gravis, 179
gregalis, 179
lepida, 179
mansfieldi, 179
media, 179
potens, 179
poulseni, 179
propinqua, 179
ressert, 178
stella, 179
Dolichometopus boccar, 163, 164
occidentalis, 111
productus, 165
tontoensis, 165
Dorypyge dawsoni, 189
(Kootenia) dawsoni, 189

Dorypygidae, 113, 133, 139, 188, 190

Ehmania zone, 99, 108
Ehmaniella, 73, 100, 105, 107, 108, 116,
212, 217, 220
basilica, 219
burgessensis, 38,
217, 220; pl. 30

103, 104, 105,

NO. 5

Ehmaniella burgessensis faunule, 74,
79, 100, 104, 105, 109
quadrans, 218
waptaensis, 103, 2193 pl. 30
Eiffel Peak, 25, 120
Eldon dolomite, 4, 7, 27, 34, 37, 48,
74, 107
Elrathia, 100, 107, 108, 116, 212, 220
dubia, 220
palliseri, 214
permulta, 103, 217, 2203 pl. 30
spencei, 223
Elrathiella, 100
Elrathiella-Triplagnostus subzone, 99
Elrathina, 35, 38, 100, 102, 107, 108,
116, 221
brevifrons, 38, 103, 2233 pl. 26
cordillerae, 102, 103, 215, 2213
pl. 26
fecunda, 223
marginalis, 32, 106, 224; pl. 34
parallela, 32, 48, 106, 222; pl. 33
Spinifera, 32, 48, 106, 2233 pl. 34
Embolimus rotundata, 158
spinosa, 143
Environment of deposition, 117
Eocystites, 93
Eodiscida, 133, 137

Fieldaspis, 92, 93, 113, 116, 159, 181,
228
bilobata, 26, 92, 161, 162, 163;
pl. 16
bilobata faunule, 59, 92, 100, 120
cf. bilobata, 33, 1623 pl. 16
celer, 110
furcata, 26, 36, 37, 50, 91, 92, 159,
161, 162; pl. 15
furcata faunule, 50, 62, 91, 109
superba, 36, 92, 162, 226; pl. 16
superba faunule, 92, 109
Fort Mountain sandstone, 3, 5
Fossil Gully, 39, 47, 50, 51, 60, 63, 68,
73, 77; 95

Geographic and vertical distribution
of trilobites, 112
Girvanella, 45, 66, 117
Glossopleura, 73, 98, 113, 116, 119, 157,
163, 176
boccar, 98, 164, 165, 168; pl. 24
boccar faunule, 60, 98, 109
bosworthensis, 164
expansa, 167
mckeei, 18, 98, 165, 166, 168;
pl. 24

INDEX

273

Glossopleura merlinensis, 15, 98, 167;
pl. 23

merlinensis faunule, 69, 98, 109
nitida, 164
perryi, 166
skokiensis, 15, 98, 1663 pl. 23
sp. undet., 15, 98, 168; pl. 24
stenorhachis, 18, 98, 1653 pl. 24
stephenensis, 111, 164
templensis, 27, 98, 164; pl. 24
walcotti, 167

Glossopleura zone, 69 73, 80, 96, 105,

108, 113, 114, I15

Glyphaspis, 34, 116, 200, 201, 224
parkensis, 35, 107, 224; pl. 34
paucisulcata, 225
storeyt, 225

Gog formation, 6

Goodsir trough, 5, 45, 60. 75

Grabau, A. W. R., 86

Hanburia, 107, 108, 116, 197
gloriosa, 102, 103, 197; pl. 25
Harrington, H. J., 133
Hector Creek, 13
Helcyonella belliana, 101
romingeri, IOI
wheeleri, 101
Howell, B. F., 80, 81, 85, 87, 96, 99, 131
Hyolithellus, 80
annulatus, IOI
flagelllum, 101
Hyolithes, 51, 80, 89, 118
carinatus, IOI
cecrops, 93

Inglefieldia, 85, 86
perola, 244
Inglefieldia? birdsalli, 209, 210
Iphidella fieldensis, 38, 101, 103
pulchra, 103

Karlia stephenensis, 188
Kicking Horse Mine, 35
Klotziella, 107, 108, 116, 168

ornata, 102, 168; pl. 28
Kobayashi, T., 133, 139, 179
Kochaspis, 90, 92, 93, 115, 116, 163,

200, 201, 225, 230

carina, 110

cecinna, 6, 110, 226, 227

celer, 225

chares, 225

eiffelensis, 26, 33, 36, 50, 9I, 92,

226, 227; pl. 14

274. SMITHSONIAN MISCELLANEOUS COLLECTIONS

Kochaspis liliana, 93, 115
liliana zone, 87
Kochiella, 85, 114, 115, 116, 227, 230
agnesensis, 207
crito, 209
mansfieldi, 227
maxeyi, 26, 50, 91, 92, I10, 226,
228eup)lu0s
cf. maxeyi, 36, 229; pl. 13
tuberculata, 227
Kochiella zone, 81, 85, 87
Kochina, 94, 116, 229
americana, 12, 19, 25, 94, 229, 231;
pl. 19
bosworthensis, 229
macrops, 12, 2313 pl. 19
Kootenia, 32, 34, 35, 38, 49, 50, 94, 96,
97, 102, 106, 107, 108, I13, 115, 116,
140, 178, 188, 190
burgessensis, 38, 103, 1893 pl. 28
dawsoni, 102, 1893 pl. 27
Kootenia zone, 81, 82

Lake Agnes shale, 16, 19, 62, 118
Lake Louise shale, 3

Lingulella waptaensis, 103

Litocodia, 201

Localities, index of, 121-130
Lochman, C., 83, 87, 96, 120, 201
Lower-Middle Cambrian boundary, 84

Matthew, G. F., 76
Menocephalus salteri, 188
Mesonacida, 139
Method of work, 7
Mesxicella, 94, 115, 116, 23%
mexicana, 231
Stator, 12, 10,25, 28,04, 120, 23%
pl. 20
Micromitra burgessensis, 103
(Paterina) charon, 110
Middle Gully, 39
Monarch Creek, 45, 72, 73
Monarch Mine, 46
Moore, R. C., 84
Mount Assiniboine, 60
Mount Bosworth, 21
Mount Field, 35, 37, 78
Mount Niblock, 15
Mount Odaray, 31; pl. 7
Mount Schaffer, 29
Mount Stephen, 39, 63, 68, 71, 73, 76,
96; pls. 2, 3, 4, 6
Mount Temple, 26; pls. 1, 5
Mount Victoria, 22
Mount Whyte, 17

VOL. I16

Mount Whyte formation, 3, 6, 13, 14,
16, 19, 20, 22, 23, 24, 26, 28, 20, 30,
33, 36, 43, 47, 50, 52, 56, 84, 87, 88,
89, 90, 91, 92, 118, 120

Naiset formation, 6, 60
Neolenus granulatus, 189
serratus, 189
Nisusia, 102
alberta, 101
burgessensis, 103
(Jamesella) lowi, 110
North Gully, 39, 47, 51, 52, 63

Obolella zone, 81
Obolus damo, 110
mceconnelli, 10%
parvus, 94
whymperi, 110
Ogygia klotzi, 191
Ogygopsididae, 133, 139, 190
Ogygopsis, 78, 80, 95, 104, 114, 115,
116, 140, 190
klotzi, 50, 51, 52, 88, 89, 96, 102,
107, 108, 114, 1913 pls. 12, 21, 29
klotzi faunule, 100, I0I, 109, 114
spinulosa, 50, 96, 1923 pl. 21
Ogygopsis shale, 42, 64, 71, 75, IOI,
104, III, I19, IQI
Olenellida, 133
Olenellidae, 133
Olenellus, 30, 31, 33, 47, 54, 58, 86, 116
canadensis, 82
Olenellus zone, 81, 82, 84, 86
Olenidae, 202
Olenoides, 32, 62, 74, 106, 107, 108,
I10, 113, 115, 116, 140, 189
constans, III
damia, 60
nevadensis, 189
serratus, 38, 102, 103, 189; pl. 27
serratus faunule, 78, 100, 102, 109
Spinosus, 140, 143
Olenoides-Marjumia zone, 99
Olenoides (Neolenus) zone, 99
Olenopsis agnesensis, 207
americanus, 229
cleora, 208, 209
crito, 208, 209
leuka, 110
Onchocephalus, 92, 114, 116, 199, 201,
232
cleon, 47
depressus, 36, 92, 2333 pl. 14
fieldensis, 36, 37, 91, 232, 233,
234; pl. 14

a

NO. 5

Onchocephalus maior, 36, 92, 234; pl.
14
‘sublaevis, 36, 92, 2343 pl. 14
thie, 47, 232, 233, 234; pl..8
Opisthoparida, 133, 138
Orria, 191
Orthotheca corrugata, 10%
maior, 10l
Oryctocephalidae, 113, 133, 130, 192
Oryctocephalites, 92, 113, 116, 195
ressert, 36, 91, 1963 pl. 15
typicalis, 195, 196
Oryctocephalus, 48, 89, 94, 107, 108,
Tig, 115, 116, T40) ros
burgessensis, 104, 112, 194, 195;
pl. 26
matthewi, 102, 104, III, 1953 pl.
26
primus, 193, 194, 225
reynoldsi, 102, 104, 111, 1933 pl. 20
sp. undet. No. 1, 52, 88, 1933 pl. 9
sp. undet. No. 2, 51, 89, 1933 pl. 9
walkeri, 102, 111, 195

Pachyaspis, 95, 107, 108, 116, 235
attenuata, 32, 106, 2353 pl. 33
moorei, 235
typicalis, 235

Paedeumias, 30, 116

Paget formation, 5

Pagetia, 38, 47, 78, 94, 102, 104, 107,

TOO, 112. 113, 11S, 116,-137
bootes, 38, 102, 103, 104, 1373
pl. 25
bootes faunule, 79, 100, 103, 109
cf. bootes, 32, 49, 106, 1373 pl. 33
ellsi, 112
sp. undet., 50, 96, 138; pl. 22

Pagetiidae, 133, 137

Palella-Thomsonaspis zone, 99

Park Mountain, 33, 75; pl. 2

Parkaspis, 107, 108, 116, 169
decamera, 104, 1713 pl. 27
endecamera, 35, 107, 160, 170, 171;

pl. 31
endecamera faunule, 101, 105, 106,
109

Paterina, 51, 89
wapta, 94
zenobia, 38, 102, 103

Periomma, 199, 201

Periommella, 201

Peronopsidae, 133

Peronopsis, 107, 108, 116, 133
columbiensis, 32, 35, 49, 106, 107,

1343 pl. 33

INDEX 275

Peronopsis interstricta, 135
montis, 102, 104, 112, 1343 pl. 25
Peyto limestone, 13, 14, 16, 20, 22, 23,
24, 25, 26, 28, 20, 30, 33, 47, 53, 55;
82, 118
Piazella, 89, 114, 116, 201, 213, 236
pia, 47, 236, 237; pl. 8
tuberculata, 52, 88, 236; pl. 8
Pika limestone, 7
Pinnacle Mountain, 28
Plagiura, 6, 92, 93, 115, 116, 201, 237
cercops, 14, 19, 20, 21, 22, 26, 36,
50, 86, 90, 91, 92, 2373 pl. 13
cercops faunule, 91, 109
cleadas, 237
Plagiura zone, 86, 87, 93
Plagiura-Kochaspis zone, 46, 62, 80,
88, 90, 93, 95, II0, 113, 114, II5, 120,
126, 127
Plagiurella, 201
cleadas, 237
Plain of Six Glaciers, 17
Poliella, 81, 84, 89, 113, 116, 170, 172
castlensis, 172
denticulata, 51, 89, III, 173, 1743
pl. 12
cf. denticulata, 52, 88, 1743 pl. 9
prima, 16, 19, 51, 52, 89, £72, 174,
184, 186; pl. 12
prima faunule, 63, 80, 109
sp. undet. No. 1, 50, 96, 1743 pl. 22
sp. undet. No. 2, 49, 96, 1753 pl. 22
Polypleuraspis, 73, 98, 113, 116, 1109,
175
insignis, 18, 27, 1763 pl. 23
insignis faunule, 98, 109
solitaria, 175, 177
Popes Peak, 24
Poulsen, C., 85, 113, 114, 178, 198
Poulsenia granosa, 211
Proliostracus, 200, 201
Prozacanthoides, 142
Ptarmigan formation, 4, 6, 64, 65, 66,
III
Ptarmigania, 94, 95, 113, 116, 159, 170,
177, 178, 181
longula, 177, 178
rossensis, 12, 25, 28, 94, 120,
177; pl. 19
Ptarmigania strata, 142, 178, 1901
Ptarmiganoides, 94, 95, 113, 116, 178
bowensis, 12, 94, 178, 1793 pl. 20
Ptychoparella, 200, 201
Ptychoparia, 140, 198, 199
adina, 110
cilles, 110

276

Ptychoparia clusia, 110, 163
cordillerae, 221, 222, 223, 224
cossus, 110
gogensis, 60
palliseri, 214
permulta, 217, 220
perola, 244
pia, 236
piochensis, 207
skapta, 210
thia, 232

Ptychoparia? cercops, 237
cleadas, 237

Ptychopariidae, 198

Piychoparioidae, 114, 133, 140, 198

Range of trilobite genera, 116

Rasett) Hs Lies 113) 132.8130) 170

Resser, C. FE), 81,63, 85, 87, 93, 94;
LOS) hits ln TAS aT 7 Ome OT a e27.
228, 240

Richter, R., 139

Richter, R., and Richter, E., 130, 108

Rominger, C., 76

IMO RS, Rae oy agin

Ross Lake, 20

Ross Lake shale, 4, 6, 12, 19, 25, 28,
66, 69, 93, 115, 118, 119, 120

Rustella, 56

Stibitane foninationy a) )5s0ls, 275) 5S:
117
Scenella, 51, 89, 118, 119
amit, 101
columbiana, 101
Schenk, H. G., and Muller, S. W., 84
Schistometopus, 92, 115, 116, 238
collaris, 26, 50, 92, 2393 pl. 14
convexus, 20, 21, 22, 26, 36, 50,
OI, 92, 238, 239; pl. 13
typicalis, 238, 239
Sherbrooke formation, 5
Skoki Valley, 15
Slate Mountains, 14
Solenopleura, 200
Solenopleurella, 32, 74, 104, 107, 108,
116, 240
diligens, 241
sp. undet. No. 1, 38, 104, 2413
pl. 25
sp. aindet. No, 2,. 38, 103, 2413
0) ed
Spencia, 240
Stauroholcus, 240
typicalis, 240

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116

Stephen formation, 4. 7, 11, 18, 27,
31, 35, 37, 46, 48, 70, 98, 99, 119
Stephenaspis, 89, 113, 116, 159, 163,
180, 188
bispinosa, 51, 52, 89, 163, 181, 182,
183; pl. 10
bispinosa faunule, 63, 89, 114, 191
cf. bispinosa, 52, 88, 182; pl. 10
Stose, G. W., and Jonas, A. I., 82
Strotocephalus, 96, 97, 114, 202, 203
gordonensis, 203
Stubblefield, C. J., 140
Syspacephalus, 53, 85, 86, 88, 89, 114,
£17,201, 240
charops, 47, 242, 243, 244, 246,
247; pl. 8
crassus, 53, 88, 246; pl. 9
gregarius, 51, 52, 53, 88, 89, 242,
246; pl. 8
laevigatus, 51, 890, 2453 pl. 9
laticeps, 53, 88, 2433 pl. 9
laticeps faunule, 88, 109, 114, 101
perola, 16, 19, 51, 52, 80, 2445
pl. 9
tardus, 50, 96, 2473 pl. 22
Syspacephalus zone, 81, 85, 87

Taxioura, 95, 101
typicalis, 191
Terminology, paleontologic, 131
stratigraphic, 9
Tholiasterella? hindei, 93
Thomsonaspis, 74
Thomsonaspis zone, 99
Tonkinella, 107, 108, 113, 117, 196
flabelliformis, 106
stephensis, 32, 35, 49, 73, 106, 107,
196; pl. 31
stephensis faunule, 74, I01, 105,
106, 107, I09
Triplagnostus, 100, 107, 108, I17, 135
burgessensis, 104, 1363 pl. 25

Undetermined pygidium No. 1, 1873
pers

Undetermined pygidium No. 2, 187;
pl. Io

Urotheca parasitum, 93

Vanuxemella, 04, 113, 117, 183
contracta, 183
nortia, 12, 19, 25, 28, 94, 120, 1833
pl. 20
Vistoia prisca, 183

NO. 5

Walcott, C. D., 3, 4, 5, 11, 15, 42, 45,
54, 56, 57, 58, 62, 64, 69, 70, 71, 72,
73, 75, 76, 77, 78, 84, 93, 102, I10, 120

“Wanneria”’ gracilis, 56

Warburg, E., 190

Wenkchemma, 89, 113, 117, 183

Spinicollis, 51, 52, 80, 185, 186,
187; pl. 11

sulcata, 53, 88, 1863 pl. 11

walcotti, 16, 19, 89, 1843 pl. 11

Wenkchemnia-Stephenaspis zone, 8o,
87, 113, I14, 115, 191, 192

Westergard, A. H., 133

Wimanella, 49, 96

catulus, 110
rossensis, 04
walcotti, 94

Yoho shale, 51, 52, 63, 118
Yohoaspis, 117, 248
pachycephala, 50, 96, 2483 pl. 21
pachycephala faunule, 96, 109, 114,
IQI
Yuknessaspis, 107, 108, 117, 249
paradoxa, 35, 107, 2503; pl. 32

INDEX 277

Zacanthoides, 34, 49, 94, 96, 106, 107,

TOG, Lae 105) LiOs) 117s 130, 540,
181

abbreviatus, 145

adjunctus, 145

cimon, III

divergens, 32, 35, 107, 1463 pl. 32

idahoensis, 145

longipygus, 32, 107, 1443 pl. 32

planifrons, 107, 1453 pl. 32

romingeri, 102, 143, 144, 145, 146;

Hlee27

sampsoni, 147

serratus, 142, 147

sexdentatus, 50, 96, 1413 pl. 22

sp. undet., 50, 96, 1403 pl. 21

spinosus, 143, 145

submuticus, 32, 107,

pl. 32

walapai, 145
Zacanthoides-Anoria zone, 96, 97
Zacanthoididae, 113, 133, 139, 140
Zacanthoididea, 139
Zacanthopsis, 113, 116

cf. levis, 30

143, 145;

Oo ee

AEE ea:
DAD oot

j 3 fA

Me ‘SMITHSONIAN MISCELLANEOUS COLLECTIONS
| VOLUME 116, NUMBER 6

Charles D. and Mary Waux Walcott
Research Fund

te

oe NEW SPECIES OF THE JURASSIC
~ BRACHIOPOD GENUS SEPTIRHYNCHIA

Wire Two Prates)

. ‘+ BY
ay ee HELEN M. MUIR-WOOD |
{ British Museum (Natural History), London

Lee AND
G. ARTHUR COOPER
U, S. National Muceuni, Washington, D. C.

S82ece0ce*

(Pustication 4047)

raters CITY OF WASHINGTON
"PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 5, 1951

Sa SG LES SOY Joa eee
per me * eae : : 3
oe cs ic

on
Soy

FEA
te

Sages
at:

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 116, NUMBER 6

Charles D. and Flary Waux Galcott
Research Fund

Pane Wo SseB@iEs OF THE: JURASSIC
BRACHIOPOD GENUS SEPTIRHYNCHIA

(WituH Two PLrateEs)

BY
HELEN M. MUIR-WOOD
British Museum (Natural History), London
AND
G. ARTHUR COOPER

U. S. National Museum, Washington, D. C.

(PUBLICATION 4047)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 5, 1951

I 4
j

rat i yar :
i i y ry 7 oT ; ‘ | a.
Ba ys Oars ps5 6 fi

xX

The Lord Baltimore Drees

BALTIMORE, MD., U. 8. A.

Charles D. and Mary Waux Walcott Research Fund

im NEW SPECIES, OF THE JURASSIC BRACHIO-
POD: GENUS SEPTIREAY NCHIA
By HELEN M. MUIR-WOOD
British Museum (Natural History), London

AND

G. ARTHUR COOPER
U. S. National Museum, Washington, D. C.

(With Two PLatEs)

The specimens described herein were collected by Dr. Barnum
Brown while on the Dudley Expedition of the Anglo-American
Oil Co., which was organized in 1920 to examine part of Harrar
Province, Abyssinia. Some of the brachiopod material collected by
this expedition is silicified and yielded well-preserved interiors of
species of Pseudoglossothyris and Somalirhynchia, as well as of the
rare Septirhynchia. The attention of collectors in this portion of
Africa is drawn to fossils having this type of preservation.

The preparation of the specimens was made by dissolving away
the matrix in dilute hydrochloric acid. This yielded two fragmentary
pedicle valves of Septirhynchia from Ab-Uqua-Kurtcha and a com-
plete specimen from Ego-Gambo with both valves in contact repre-
senting a new species. The latter produced a nearly perfect interior,
which was photographed with the valves intact. After the valves were
photographed, they were pried apart to facilitate further study and
illustration. Present information on the interior of Septirhynchia
was obtained by sectioning the pedicle beak. The interior details of
the brachial valve were hitherto little known.

The genus Septirhynchia was originally described by Muir-Wood
(1935, p. 106, pl. 9, figs. 3 a-c, 4; text fig. 11) with Rhynchonella
azaist Cottreau as type species. This species was described in 1924
by Cottreau from Lower Kimmeridgian rocks of Harrar, Abyssinia.
Basse (1930, p. 117) recorded this species from the Kimmeridgian
of Cabenaoua, south of Harrar. The silicified specimen figured in
Weir’s earlier paper (1925, p. 83, pl. 12, fig. 26) as Stolmorhynchia

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 6

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

cf. S. nobilis (Sowerby) from the base of the Jurassic ( ?Kim-
meridgian) of Hamud, western British Somaliland, corresponds with
Cottreau’s figures of R. agaist. Weir’s specimen is said to be 53 mm.
wide and 60 mm. thick and to have 12 to 16 costae.

Septirhynchia azaisi is found in a grayish limestone weathered to
a reddish brown. The shell is usually preserved in beekite. This
limestone bed can be traced from Harrar, Abyssinia, through Hamud,
western Somaliland, to Ida Kabeita and Ambal, central Somaliland,
and thence to Anole Issa, Italian Somaliland. It is possible that this
limestone is of Callovian and not Kimmeridgian age, according to
Dr. Migliorini (personal communication to Muir-Wood). The age
of the other three species originally assigned to the genus is: Rhyn-
chonella budulcaensis Stefanini, Kimmeridgian, Italian Somaliland;
Septirhynchia mogharaensis Muir-Wood (=hynchonella decorata
Cossman, 1925), Callovian of Sinai Peninsula ; and S. madashonensis
Muir-Wood, Callovian of eastern British Somaliland.

The original diagnosis of this genus given by Muir-Wood is—

Shell large, pentameroid in contour; median fold low, median sinus shallow;
ventral umbo long, produced and incurved, concealing foramen and deltidial
plates in adult; hypothyrid, deltidial plates conjunct. Dental lamellae strong,
uniting with base of ventral median septum posteriorly. Ventral septum ex-
tending for two-thirds of length of pedicle valve. Dorsal median septum low.
Two slightly divergent septal plates extend across posterior part of dorsal

umbonal cavity. Interareas large and flattened. Pedicle collar developed. Shell
thick, ornamented with coarse, prominent, subangular costae.

A study of the silicified material has shown that this diagnosis
requires to be emended, since the “divergent septal plates” are ac-
tually the long, curved radulifer crura. Comparison of our specimens
with descriptions of described forms shows that the silicified complete
specimen is a new species.

SEPTIRHYNCHIA PULCHRA, new species
Plate 1, figs. 1-5, 11, 12; plate 2, figs. 1-6

Diagnosis of species ——Species small for the genus: length 36.5 mm.
(incomplete), brachial length 31 mm., width 30 mm., thickness 32 mm.,
longer than wide, with the pedicle valve slightly deeper than the bra-
chial valve; fold and sulcus poorly defined ; anterior commissure not
preserved ; costae continuous from umbo to anterior margin, broadly
rounded, and numbering about 13. Costae crowded and narrow pos-
teriorly, but increasing in width anteriorly and lacking on the umbonal
slopes. Beak of pedicle valve strongly incurved, overhanging and en-

No. 6 NEW JURASSIC BRACHIOPOD—M UIR-WOOD AND COOPER 3

closing the incurved umbo of the brachial valve. Areolae broad and
smooth.

Description of interior—The interior of the pedicle valve is charac-
terized by a median septum extending just beyond the half length
of the shell and dividing the delthyrial cavity. Dental lamellae de-
veloped as thin vertical plates on either side of the septum and ap-
proaching it posteriorly and equal to its length anteriorly. The portion
of the dental plates near the teeth is bowed in a median direction.
These dental plates decrease in height anteriorly and support long,
slender teeth or articulating processes, which project posteriorly above
the shell margin in the separated valves and are separated from
the margin by a narrow furrow. The foramen is small and hypothyrid
and passes inwardly into a short pedicle collar. The deltidial plates
are conjunct posteriorly but form a henidium anteriorly. Muscle
scars were not distinguishable.

The crura are curved (radulifer type, i.e., of uniform width and
without terminal hooks), 4.5 mm. long, projecting anteriorly with
their ends curved strongly toward the pedicle valve. These are di-
rectly continuous with and are given off from the anterior ends of
divided hinge-plates, which are broad umbonally but decrease rapidly
in width. Crura are not well demarcated from the inner socket-ridges,
which bound narrow, deeply inserted hinge-sockets. The septalium
is partially concealed by the incurvature of the umbo. The septalial
plates are thin and delicate and unite with the bladelike median septum,
which extends more than half the shell length and is most elevated at
its center. The cardinal process is small and knoblike, terminating in
a short spike, and, because of the curvature of the umbo, projects
anteriorly. Muscle marks are obliterated by silicification.

Holotype—U.S.N.M. No. 103961.

Locality and horizon.—?Callovian of Ego Gambo, Harrar Provy-
ince, Abyssinia.

Discussion—Septirhynchia pulchra differs from S. agaisi (Cot-
treau) by its smaller dimensions, narrower shell, and fewer costae,
13 as compared with 16 in Cottreau’s species. The internal characters
of S. pulchra also differ from those of two pedicle valves described
below, which are here referred to S. azaisi.

The dimensions of S. pulchra appear to be nearer to those of
S. budulcaensis (Stefanini) from the Middle Oolites of Bur Budulca,
southern Somaliland. There are, however, 24 to 28 costae in S. budul-
caensis, while in S. pulchra there are only 13 in each valve, with 4
on the depressed median fold and 3 in the sulcus of the pedicle valve.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Stefanini assigned Weir’s Stolmorhynchia ? azaisi var. from Jubaland
(Weir, 1929, p. 36, pl. 4, fig. 5) to his species R. budulcaensis.

Septirhynchia mogharaensis Muir-Wood, the type of which was
figured by Cossman as Rhynchonella decorata (1925, p. 326, pl. 7,
fig. 9 a-c) from the Callovian of Darb-el-Cheikh, Moghara Massif,
Isthmus of Suez, is larger than S. pulchra and has a length of 45 mm.,
width 40 mm., and thickness 45 mm., with a well-defined median fold
and sulcus. From the figures the type specimen appears to have about
17 promiment, rounded costae. The anterior portion of the pedicle
valve tapers and forms a short linguliform extension.

The small specimen figured by Stefanini (1932, pl. 6, fig. 9 a, b)
as R. azaisi from the Middle Oolites of Anole Issa, southern Somali-
land, corresponds in dimensions and general shell outline with those
of S. pulchra, but the costae appear to be slightly coarser and less
numerous (II costae) in the Somaliland specimen. It should, how-
ever, be identified as S. pulchra var. rather than S. azaisi. The dimen-
sions of this specimen from the figures are: length 40 mm., width
34 mm., and thickness 31.5 mm. A suggestion is here made to col-
lectors in Africa that other rhynchonellid genera resemble Septi-
rhynchia in the strongly curved umbo and costae but differ markedly
internally. A silicified fragmentary specimen from Jigjiga (U.S.N.M.
No. 107067) at 7,150 feet, shows both valves and has similar dimen-
sions and beak characters to those of S. pulchra, but the median
fold is scarcely defined and the costae are more numerous (19). The
septum of the pedicle valve is developed only as a low ridge. Other-
wise the internal characters, teeth and dental lamellae, are similar. A
cardinal process occurs in the brachial valve. The crura are not
preserved and the hinge-plates are imperfect.

A fragmentary specimen (U.S.N.M. No. 107068) from Dire Daoua
(from base of 2- to 10-foot beds above the conglomeratic limestone
that overlies the sandstone) with the two valves in contact and show-
ing internal structure appears to be distinct. It lacks the cardinal
process and septum of the pedicle valve so characteristic of S$. pulchra.
The interior has 20 costae, which are less coarse, and flattened areolae
are not developed on the flanks. The foramen is larger than that of
S. pulchra, and the deltidial plates as preserved are not conjunct.

SEPTIRHYNCHIA AZAISI (Cottreau)
Plate 1, figs. 6-10
For full synonymy see Muir-Wood, 1935.

Two fragmentary pedicle valves from Ab-uqua-Kurtcha show the
septum and dental lamellae uniting about 5 mm. anterior to the umbo.

No. 6 NEW J URASSIC BRACHIOPOD—-M UIR-WOOD AND COOPER 5

The posterior part of the umbonal cavity is occupied by a tubular
pedicle collar about 4 mm. long in specimen (U.S.N.M. No. 107066a).

The median septum is a knifelike ridge which increases gradually
in height anteriorly and extends for a length of 28 mm. The dental
lamellae are also knifelike plates 11 mm. in length in both specimens
and unlike those of S. pulchra considerably shorter than the median
septum of the pedicle valve. The lateral cavities between the dental
lamellae and the inner shell wall are very narrow. The deltidial plates
are conjunct with the line of junction traceable interiorly. They are
continued farther anteriorly as a single plate (henidium of Cloud,
1942, p. 12), which is externally concave and depressed below the
margins of the delthyrium. The teeth are not preserved.

The external shell characters, ornament, and outline agree well with
those of S. agaisi (Cottreau). The original description of R. azaisi
(Cottreau, 1924, p. 581, pl. 17, figs. I-4) gives the dimensions as
length 64 mm., width 56 mm., thickness 56 mm. There are, as a rule,
16 subangular costae, some 6 mm. wide. A low median fold and
shallow sulcus are developed. The internal characters are not men-
tioned by Cottreau, but his longitudinal section (fig. 4 b) appears to
show one radulifer crus and part of the median septum.

The relationships of this remarkable genus are still unknown. The
development of a median septum in the pedicle valve as well as a car-
dinal process in the brachial valve are exceptional characters in Juras-
sic rhynchonellids. A new family Septirhynchiidae is therefore pro-
posed for this genus. Its distribution, as at present known, is restricted
to the Ethiopian province of the Upper Jurassic.

REFERENCES
Bassg&, E.
1930. Contribution a l’étude du Jurassique supérieur (facies corallien) en
Ethiopie et Arabie méridionale. Mém. Soc. Géol. France, n.s., vol.
6, pts. 3-4, pp. 105-148, pls. 20, 21.
Croup, BE Jr:
1942. Terebratuloid Brachiopoda of the Silurian and Devonian. Geol. Soc.
Amer., Spec. Pap. 38.
CossmAn, M.
1925. Description des espéces. Jn Douvillé, H., Le Callovien dans le massif
de Moghara. Bull. Soc. Géol. France, ser. 4, vol. 25, pp. 305-328,
pls. 5-8. (Type of R. mogharaensis.)
CorrrEAu, J.
1924. Invertébrés Jurassiques de la région de Harrar (Abyssinia). Bull.
Soc. Géol. France, ser. 4, vol. 24, pp. 579-591, pls. 17, 18.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

DovuvILLeé, H.

1925. Le Callovien dans le massif de Moghara: Avec description des fos-
siles par M. Cossman. Bull. Soc. Géol. France, ser. 4, vol. 25,
PP. 303-328.

Mutrr-Woop, HELEN M.

1935. The Mesozoic palaeontology of British Somaliland: Pt. 7, Jurassic

Brachiopoda, pp. 75-147, pls. 8-13. London.
STEFANINI, G.

1932. Echinodermi, Vermi, Briozoi e Brachiopodi del Guiralias della
Somalia. Palaeontographia Italica, vol. 32 (n.s. vol. 2), pp. 81-141,
pls. 4-8. (R. budulcaensis.)

WET, J.

1925. Brachiopoda, Lamellibranchiata, Gastropoda and Belemnites in the
collection from Somaliland. Mon. Geol. Dep. Hunterian Mus.,
Glasgow Univ., vol. 1, pt. 6, pp. 79-110, pls. 11-14.

1929. Jurassic fossils from Jubaland, East Africa, collected by V. G. Glen-
day, and the Jurassic geology of Somaliland. Ibid., vol. 3, pp. 1-63.
pls. 1-5.

EXPLANATION OF PLATES
PLATE I

Figs. 1-5, 11, 12, Septirhynchia pulchra, new species, Upper Jurassic (?Cal-
lovian), Ego-Gambo, Harrar Province, Abyssinia: 1, 2, 4, 5, Brachial,
posterior, lateral, and pedicle views, respectively, of the complete specimen,
X 1, holotype (U.S.N.M. No. 103961) ; 3, anterior view of the same speci-
men showing anterior profile and interior, X 1; 11, 12, brachial valve of
holotype looking directly into the interior and showing the median septum,
hinge-plates, crura, and septalium.

Figs. 6-10, Septirhynchia azaisi (Cottreau), Upper Jurassic (?Callovian) of
Ab-Uqua-Kurtcha, Harrar Province, Abyssinia: 6, Pedicle valve showing
interior from side, hypotype (U.S.N.M. No. 107066a) ; 7, beak of preceding,
showing small foramen and henidium, * 2; 8, interior of delthyrial cavity
of same hypotype showing dental plates, pedicle collar, and henidium, * 2; 9,
interior of pedicle valve showing long septum and short dental plates, « 1
(U.S.N.M. No. 107066b) ; 10, beak of the same specimen showing conjunct
deltidial plates, henidium, and small foramen, * 2.

PLATE 2

Figs. 1-6, Septirhynchia pulchra, new species, Upper Jurassic (?Callovian),
Ego-Gambo, Harrar Province, Abyssinia: 1, 2, Pedicle valve of holotype
(U.S.N.M. No. 103961) showing interior, and tilted to show dental plates,
septum, and teeth, 2; 3-5, views of the brachial interior of the holotype
showing septalium, septum, and details of the crura and hinge-plates, & 2;
6, holotype with both valves in contact showing deep insertion of brachial
umbo into the delthyrial cavity of the pedicle valve, * 2.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 6, PL. I

SEPTIRHYNCHIA

(SEE EXPLANATION OF PLATES AT END OF TEXT.)

Z@ “Id ‘9 “ON ‘91L “1OA SNOILDA1109 SNOANVYTISDSIN NVINOSHLIWS

“SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 116, NUMBER 7
(End of Volume)

i jeiae BUTTERFLIES OF
VIRGINIA

~ (Wir 31 Pratzs)

Se ER eS

iS ——-s ie Wy Ae.
a. a 2 in = fA S
c x a hoe oo Pye

BY
AUSTIN H, CLARK
AND

LEILA F. CLARK
Smithsonian Institution ty AS ON A

DEC 29 12

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PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 20, 1951

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, FRONTISPIECE

10

BUTTERFLIES OF VIRGINIA

(From photograph by Frederick M. Bayer. For explanation, see page 195.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 116, NUMBER 7
(End of Volume)

tia tet) Tee lES: OF
VIRGINIA

(WitH 31 PLATES)

BY
AUSTIN H. CLARK
AND
LEILA F. CLARK

Smithsonian Institution

AFR AE-INCp,
Z OS
By sense Avo On. (

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fe
ee

(PUBLICATION 4050)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 20, 1951

The Lord Baltimore Press

BALTIMORE, MD., U. 8. A.

PREFACE

SINCE 1933 we have devoted practically all our leisure time to an
intensive study of the butterflies of Virginia. We have regularly spent
our annual leave in the State, stopping at various places from which
each day we drove out into the surrounding country. In addition to
prolonged visits of 2 weeks or more to various towns and cities, we
spent many week ends in particularly interesting localities. We have
visited all the 100 counties in the State at least twice, most of them
many times, and our personal records are from more than 800 locali-
ties. We have paid special attention to the Coastal Plain, particularly
the great swamps in Nansemond, Norfolk, and Princess Anne
Counties, and to the western mountains.

Virginia is so large and so diversified that it would have been im-
possible for us, without assistance, to have made more than a super-
ficial and unsatisfactory study of the local butterflies. We have been
so fortunate as to have had invaluable assistance from many friends
within the State, some of whom have sent us hundreds of records and
many specimens from their home territory.

Prof. Carroll M. Williams, of Harvard, provided us with records
from Richmond and localities farther east, extending over several
years. Dr. Carroll E. Wood, Jr., and Dr. Carl W. Gottschalk collected
records and specimens for us at Salem and in Roanoke County for
more than 7 years. Frank W. Trainer sent us detailed records and
many specimens, covering a period of several years, from Farmville,
and later from Charlottesville. Warren P. Stoutamire sent us his
records from Gala, and Lloyd G. Carr sent us records and specimens
from Mountain Lake.

But this is by no means all the assistance we have had. Prof. Ellison
A. Smyth, Jr., was so very kind as to send us his records for more
than 50 years’ collecting, chiefly from Blacksburg and Poverty Hol-
low, Montgomery County, and also from Salem. These were supple-
mented by records from Poverty Hollow sent us by Herman J. Erb.
Dr. Frank Morton Jones most generously sent us his records from
eastern Virginia, especially Accomack, Northampton, Princess Anne,
and Nansemond Counties. Jackson H., John, and Alexander Boyd
have sent us many interesting records from Woodberry Forest and the
Dismal Swamp region. John M. Burns has given us numerous records
from Mountain Lake.

ili

iv SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Otto Buchholz has been so good as to send us detailed records of
all the species he collected over a period of several years in eastern
Virginia, several of which were not found by anyone else. Dr. War-
ren Herbert Wagner, Jr., made many trips to different parts of Vir-
ginia to get records for us, and we owe most of the information re-
garding the distribution of Poanes aaront to him. He and Dr. George
W. Rawson paid special attention to the Second Swamp near New
Bohemia and also to the North Landing River swamp, providing us
with many records from those interesting areas.

On our visits to various localities in Virginia we have from time to
time been accompanied by friends who have assisted us in gathering
specimens and records. Mr. and Mrs. Ernest L. Bell have been with
us in eastern Virginia, and also in Frederick County. Prof. and Mrs.
Charles T. Brues and Dr. Alice Brues joined the Bells and ourselves
on a visit to the Dismal Swamp region. Dr. and Mrs. William M.
Mann accompanied us on a 2 weeks’ trip to southern and southwestern
Virginia, and also on other trips to the Dismal Swamp and the North-
ern Neck. Jackson H., John, and Alexander Boyd joined us in a
visit to the Dismal Swamp, and Dr. and Mrs. Harald A. Rehder have
also been with us in the same area. Mr. and Mrs. William D. Field
and later Mr. and Mrs. R. Tucker Abbott accompanied us on expedi-
tions to Middle Mountain in western Highland County.

Others who have been with us on collecting trips are Dr. and Mrs.
Adam G. Boving, Dr. and Mrs. Grover C. Pitts, Dr. Carl W. Gott-
schalk, Dr. Laurence Ilsley Hewes, Dr. William T. M. Forbes, Dr.
Walter S. Hough, Mr. and Mrs. John F. Gates Clarke, Mr. and Mrs.
David Hall, Lt. Col. and Mrs. Edvald L. Rasmussen, Dr. and Mrs.
George S. Myers, Dr. and Mrs. Matthew W. Stirling, Dr. Herbert
Friedmann, Mr. and Mrs. John E. Graf, Prof. John W. Bailey, John
B. Lewis, Mr. and Mrs. Gerrit S. Miller, Dr. and Mrs. William F.
Foshag, Mr. and Mrs. Stephen J. McDonough, Dr. Richard P. Dow,
Miss Grace Sandhouse, Dr. Ashley B. Gurney, Dr. Norman B. Tin-
dale of Adelaide, South Australia, Mr. and Mrs. Herbert G. Deignan,
Dr. Ellis LeG. Traughton of Sydney, New South Wales, Dr. Austin
Roberts and Dr. E. Percy Phillips of South Africa, Dr. H. Boschma
and Dr. L. P. Holthuis of Leiden, The Netherlands, Prof. and Mrs.
Torsten Gislén of Lund, Sweden, Mrs. G. S. Pobst of Tazewell, and
Vilhelm Lauritzen and V. Lauritsen of Copenhagen, Denmark.

During the course of our work in Virginia we have been the re-
cipients of generous hospitality in many places. Among our hosts and
hostesses we are especially indebted to Miss Lilian E. Smith of Mount

NO. 7 PREFACE Vv

Solon ; Miss Willie T. Weathers of Aylett; Miss Florence Walker of
Bayford ; Dr. and Mrs. William T. Sanger, Dr. and Mrs. Wortley F.
Rudd, and Dr. and Mrs. Sidney S. Negus of Richmond; Lt. Col. and
Mrs. Edvald L. Rasmussen and Mr. and Mrs. Richard H. Rule of
Fairfax; Mr. and Mrs. John A. Blakemore of Abingdon; Dr. and
Mrs. George W. Jeffers of Farmville; Mr. and Mrs. Gus Welsh of
Apple Orchard Mountain; Theodor Mussaeus of Limeton; John B.
Lewis of Amelia Court House; and L. Parker Hill of Suffolk.

While this memoir was in press we received many additional records
from Petersburg sent us by Bryant Mather of Jackson, Miss.; from
the Dismal Swamp region sent us by Kilian Roever of Jackson, Tenn. ;
and from Boyd’s Tavern, which we owe to the kindness of Miss
Leila A. Henry of Arlington, Va. Also, we enjoyed the hospitality
of Mr. and Mrs. Douglas D. Withers of Mount Solon, and accom-
panied Mr. and Mrs. William J. Faymonville, Jr., of Louisville, Ky.,
on a visit to Reddish Knob.

William D. Field, associate curator of insects, United States Na-
tional Museum, has been so good as to read over the entire manu-
script for us, and we are greatly obliged to him for his critical com-
ments.

Cyril F. dos Passos most kindly checked all the scientific names,
bringing them into conformity with current usage.

We realize that this report represents only an introduction to the
study of the butterflies of Virginia. A vast amount of work still re-
mains to be done before they can be considered as adequately known.
Many areas in the western mountains and on the Coastal Plain have
not been explored, and far more information is necessary for the ac-
curate determination of the seasons in different parts of the State.

We hope later to publish our detailed records, and also our notes
on the habits and other attributes of the various species.

For the illustrations we are indebted to the late Gurney I. High-
tower and to Floyd B. Kestner, Smithsonian Institution photog-
raphers. All the specimens shown are in the United States National
Museum. ,

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CONTENTS

Page
CURE Te CSS Sen Gey i MRR sp Ogi ee, A ar or ed iii
Uy SLR SEES GEaTVe Ta Val Ag er et lg PRR eo eR on LE ar a a I
BIStribUtton OL DUtteniies) Ine VALeinidee + eriniersetee ccieereieeiciclarenisielacinia cetera 2
SHER IMAP etacistc cate ues cise: « Sielefeheh eles Rio = ELIE ets So noiore wlavelevele inves cieyelee 8
evs tothe butteciites of Virginia. ©... 2c cesiiab oeeie «ec omaie's oats ee ald epee 9
1. Key to the families and subfamilies of butterflies represented in Vir-
Ciara), settee cites ENS REED POIS AI OS oiIe Gb 0 SMSO Or5 CEI 10
2uaiiey, toytie species: Of Methiinae’y.jsiclericiseis ce oe= us sis sie) a steiets Si-seia’ aioe’ ere II
tov ney 0: te SPecles/Of SAby Hide 4 fy siaty, 5c ais ays eletwia, wave, ph iaydle, ovo 0 er sve oye II
4. Key to the species of Apaturidae, Nymphalidae, Argynnidae, and
Wanaidaer er onyvonre noe cae alee ice ic mee T aee aek ee eee ce
me awey-to the species of Riodimdae sss. 0. . cfg 6 Se. he rei. doe veles 15
Gquixey-torthe speciessor Ibycacnidae vasnisces sce ote fede le eoaaee ess 15
Peer tO. be, SHECIES OL MrICEIGAE. bas saltigues) neeis ered swish be. Soe Ddlns anid’ « 18
fa ikey tothe species: of Papiliontdae. ..)sscid<ac oye 2 o> suemn oo wae sc eee 20
Opeweyztouthe Species: OL EP ycoitiae) oa <acisiereinrs cine. chose sissies. scone = a ctindi die wore dre 21
10. Key to the males of the species of Hesperiinae.................... 24
paanotired list ox butterilies'ef Virginiass ig. 7.. s.r. obs ik Pee OLS A. 30
Pea LIEN IAG AGH yaaa ia Os nisiales Ok COUN b So attache te Cae as 30
Maem RYE ALTE IG EIS 52 oe ioc 5 poe oust pba eens: Ay ckeisis, Potten whee Scie oa 39
Mar saM AMMO TESST IS AChE accor ca Sota brers Wiech tas semis ct qu hep Danke aie Poe hake SEE 40
LESTER Gols Canny 101 7) RG maa CS 2 ea SP poe ee 49
PRATITILY RD AMAT AG Heys eee) en ic eta ates gee re Tee eee 65
er MTA PARI AICTE 5. oie tins he SaaS Mae ire Sie a a ae. Be ee ee 68
a MEA Gene aN EAA LRAT OS oh crane) TE 9 ahs 5 Sox w 58 RPSL ale oR fate od to iol Se a 69
ecm yar ley Gaeii Ga men: eae seit seiner ei ks eect a tere Rays © ont een ame 70
LAIN y RCRA AG Merten et epee. orto sae aralcverhrs sicicictaress ciaia ote serie iste ee tin anerclt 86
pee LVES AD TOMMOAGT cw oa tees Sete cars 2 Vitis sy oerds Rene meres waded! 118
amily Plesneniidaew Lees cee. tate et kOe Sele). cee ola Sem ee 147
PURER CHUSCECOGEN SI ks PA ha ree t an: atts jsin cle oe ae ac copa axis Woes ea athas 186
UNM AIT WESC Soe eT, oc a he ace Daan cS oa ole bwn Caer d ja/bie se Seisitie ee 187
IED 2 teh er Seas SG EL Range CR bala, AER is viel gies, Aa 197
ERENT Sei ce aeete. ers ee PT ME, ooke | Se wera eis

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THE BUTTERFLIES: OF : VIRGINIA
By AUSTIN H. CLARK
AND
LEILA F. CLARK
Smithsonian Institution

(WiTH 31 PLATES)

INTRODUCTION

THE number of species and subspecies of butterflies for which we
have definite records for Virginia is 154. Two of these, Pieris vir-
giniensis and Phoebis philea, are accidental visitors. Several others,
as Agraulis vanillae, Strymon ontario, Ascia phileta, and Urbanus
proteus, are irregularly recurrent visitors. Several butterflies of regu-
lar occurrence in the State are summer visitors only, dying out com-
pletely during winter. This is certainly true of Phoebis sennae and
Calpodes ethlius and probably of four or five more. Vanessa carduti is
usually present and occasionally common, though in some years absent.

Two of the most abundant butterflies in the State are introductions.
Pieris rapae, an immigrant from Europe, first appeared in 1870-71,
and Colias chrysotheme eurytheme, an intruder from the west and
south, was first noted by Dr. Frank Morton Jones in 1923. If Augi-
ades sylvanus really occurs at Richmond, it represents a second intro-
duction from Europe.

On the other hand, Pieris virginiensis undoubtedly occurred in the
State before the deforestation of the western mountains. <Atrytone
arogos has not been found for nearly 50 years, but this small species
is easily overlooked. Speyeria diana and Glaucopsyche lygdamus now
appear to be restricted to only a portion of their former range. It is
probable that those species like Speyeria diana, Atrytone conspicua,
and Poanes hobomok that occur in the western mountains and in the
Dismal Swamp region formerly ranged widely over the State.

Boloria selene marilandica and Poanes massassoit hughi occur just
across the Potomac River in Maryland, and both probably will be
discovered in Virginia when the counties along the south bank of the
Potomac have been more thoroughly explored. Indeed, we suspect

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 116, NO. 7

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

that the record for Boloria selene from Bayford is based upon B. s.
marilandica; we have not seen the specimen.

Three species that occur in nearby West Virginia and one in Ken-
tucky will probably be found in Virginia, while a fifth, which occurs in
Pennsylvania and North Carolina, we have been unable to find in
Virginia. This last may have been extirpated by deforestation.

The number of butterflies known from North Carolina, a State
more than one-third again as large as Virginia, is 147; but in North
Carolina the localities where collecting has been done are relatively
few, especially in the eastern and presumably richer half. The number
known from New York, which is about one-fourth again as large as
Virginia, is 132. Being farther north, New York would be expected
to have fewer species than Virginia; but there are some obvious gaps
in the New York list, and the State has not been covered nearly so
intensively as Virginia. From England, with an area slightly greater
than that of New York but less than that of North Carolina, accord-
ing to Dr. E. B. Ford, there are 68 butterflies recorded, several of
which are rare casuals. Nine of these 68 occur in Virginia—N ymph-
alis antiopa, Vanessa atalanta, V. cardui, Boloria selene, Danaus
plexippus, D. p. megalippe, Lycaena phlaeas, Cyamris argiolus, and
Pieris rapae. But Nymphalis antiopa, Boloria selene, Lycaena phlaeas,
and Cyaniris argiolus occur in slightly different forms.

There are recognized in North America north of Mexico 700 spe-
cies, the number of species and subspecies together being 1,110.

DISTRIBUTION OF BUTTERFLIES IN VIRGINIA

The distribution of the butterflies of any highly diversified region
such as Virginia is dependent upon a number of different factors, the
most important of which are the distribution of the food plant or
plants and the conditions of humidity and temperature, especially the
daily and seasonal oscillations, under which these plants are available
for food, or under which the pupal stage must be passed.

Although the caterpillars of most butterflies will feed on several or
many different kinds of related or unrelated plants, the majority show
decided preferences, and these preferences may be uniform through-
out their range or may differ in various regions. In some cases our
native butterflies show marked preference for introduced over native
plants. Many species are confined to a single kind of plant with per-
haps a few of its close relatives. A few are confined to a single or a
few plants in their early stages, later becoming general feeders.
Several feed on plants that are widely different botanically, but
similar in their chemical composition.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 3

Since most of the plants that serve as food for butterflies in Virginia
are generally distributed over the State, the food factor is of relatively
slight significance in determining the distribution of the local species.
A few, however, are limited in their distribution by the localization of
their food plants, or by the plants with which they are invariably asso-
ciated and which for this reason are assumed to be their food plants.
Such plants are the mistletoe (Phoradendron flavescens), the large
passionflower (Passiflora incarnata), the red bay (Persea borbonia),
the canes (Arundinaria gigantea and A. tecta), the Carolina vetch
(Vicia caroliniana), and certain marsh sedges and grasses, particularly
those confined to salt or brackish marshes (species of Scirpus and
Carex, Spartina alterniflora var. glabra, and Zizania palustris).

Many species, .the food plants of which are generally distributed
over the State, are confined to regions in which, during their active or
inactive early stages, the humidity is subject to the least variation.
Most of these occur only in the mountains, particularly the higher and
wetter mountains of the southwest. At the lower altitudes in the
mountains they are found, if at all, only in permanently damp valleys
near streams, though on or near the mountain tops they become dis-
tributed generally over the frequently cloud-enshrouded open pastures.
A few, however, live in the coastal swamps or wet pine barrens, as
here near the ground where the caterpillars feed it is always damp,
though much warmer than it is in the mountains. A few species occur
both in the cool, damp mountain valleys and in the coastal swamps but
are absent from the intervening area.

An interesting illustration of the independent distribution of related
species all feeding on the same generally distributed food plants is
afforded by the several local fritillaries belonging to the genera Spey-
eria and Boloria. All the species of Speyeria and Boloria living in
Virginia feed on violets, which are abundant everywhere from the
higher mountain tops to the eastern swamps. Speyeria cybele occurs
at all points within the State and is almost everywhere common. Spey-
eria diana lives both in the mountains in damp, wooded ravines and
valleys near cold streams and on the Coastal Plain near cold springs
or oozes in the woods and swamps, but not on the Piedmont. Speyeria
idalia is found in wet, open, grassy meadows on the higher western
Piedmont, in the broader mountain valleys, and in the high mountain
pastures. Speyeria aphrodite occurs throughout the wooded mountain
valleys and on mountain pastures. Boloria toddi frequents mountain
bogs and the higher pastures. Boloria selene lives in wet open mead-
ows at more or less high altitudes, always in association with the much
more generally distributed B. toddt.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

In considering the present distribution of the butterflies of Virginia
it is important to bear in mind that it is by no means a natural one,
for local conditions have been greatly altered since the coming of the
Europeans. The lowlands have long been under intensive agricultural
development, and the endemic flora has been greatly restricted in area
and profoundly modified both by the changed conditions and by the
introduction of many European and other weeds. The mountains have
been almost wholly deforested, the original forests being now replaced
by second growth more or less extensively invaded by foreign weeds
from the lowlands or from the north. Swamps, bogs, and marshes
have been partly or completely drained, damp mountain valleys have
become dry valleys, and in general there is greater uniformity over
the State than was the case in the past.

This present relative uniformity in conditions is reflected in the
distribution of the butterflies. About 100 species, or two-thirds of the
butterflies known from Virginia, are found in all sections of the State,
from the western mountains to the coast. These are all hardy and
adaptable species, able to maintain themselves, and often to increase,
under changed conditions.

These are almost the only butterflies that occur on the Piedmont
except locally in the north, and over the cultivated areas on the Coastal
Plain. To the west they become somewhat less numerous, but they
occur in all the valleys and cleared areas, some of them also in the
woods. This faunal complex, in origin a secondary association of
vigorous, hardy, aggressive types chiefly of local origin but with im-
migrant elements from the south and west and from Europe, is dis-
tantly suggestive of the floral associations characteristic of abandoned
fields in the same area.

Superposed upon this originally secondary but now primary faunal
complex of recent origin are certain fairly distinct faunal divisions
corresponding to the well-known life zones. In Virginia these life
zones are indicated by only a small minority of the species, which for
the most part are rather strictly localized.

Canadian Zone.—In Virginia the Canadian Zone, so far as the but-
terflies are concerned, is represented only by a few isolated areas in
the west and southwest above about 4,000 feet in Augusta, Highland,
Giles, Montgomery, Grayson, and Washington Counties.

The characteristic butterfly of the Canadian Zone in Virginia is
Polygonia faunus smythi, which is everywhere common; Speyeria at-
lantis is common in the Canadian Zone on Middle Mountain in High-
land County, where Colias interior also occurs. These may occur else-
where in the Canadian Zone, but little collecting has been done there

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 5

at the time they are on the wing, and both are easily overlooked be-
cause of their resemblance to the common Speyerta aphrodite and
Colias philodice. Three additional Canadian Zone species occur in
West Virginia within sight of the Virginia border, and probably
originally these were found in Virginia.

Transition Zone.—In Virginia the Transition Zone includes the en-
tire western portion of the State from the eastern base of the Blue
Ridge westward, except for the Shenandoah Valley from Augusta
County northward and a small area in Lee County. In the north its
lower limit is at about 1,500 feet, in the south at about 2,000 feet. In
the north the eastern border of this zone becomes somewhat indefinite,
characteristic Transition Zone species occurring locally or casually
eastward, especially in the Potomac Valley, to northeastern Fairfax
County. The characteristic Transition Zone butterflies in Virginia
are: Enodia portlandia anthedon, Euphydryas phaéton, Phyciodes
batesu, Boloria toddi, B. selene myrina, Speyeria aphrodite, Polygonia
progne, Calephelis borealis, Glaucopsyche lygdamus, Incisalia irus,
I. augustinus, I. polios, Erora laeta, Pieris virginiensis, Pyrgus cen-
taureae, Erynnis persius, E. lucilius, Hesperia sassacus, Atrytone con-
spicua (also in the Dismal Swamp), Aérytone bimacula, Poanes
hobomok (also in the Dismal Swamp), and Polites mystic. The most
convenient index species of the Transition Zone in Virginia are Spey-
erta aphrodite and Polygomia progne, both of which are common and
generally distributed. The other species confined to this zone are
nearly all of local occurrence. It should be noted that species confined
to localities within the Transition Zone in Virginia are not necessarily
confined to this zone elsewhere. None of the species characteristic of
the Transition Zone in Virginia occur throughout the zone beyond the
limits of the State, and very few are found even throughout the Al-
leghanian or eastern portion. Notwithstanding these irregularities, the
Transition Zone as it is found in Virginia is a distinct and fairly well
marked zoogeographic province. Of the species characteristic of
the Transition Zone in Virginia slightly more than 80 percent are,
like the species of the Canadian Zone, single-brooded, half flying in
spring only, the other half for a more or less extended period in
summer.

Upper Austral Zone—The area herein regarded as representing the
Upper Austral Zone is bounded on the west by the Blue Ridge; on
the north it is represented by a long, narrow tongue running down the
Shenandoah Valley southwestward to Augusta County, and by irregu-
lar interdigitations with the Transition Zone in Loudoun, Prince
William, and Fairfax Counties; in the southwest an Upper Austral

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

tongue enters Lee County from Tennessee; on the western side of
Chesapeake Bay the eastern boundary of the Upper Austral may be
considered as a more or less indefinite line from the head of Mobjack
Bay to south-central Greensville County, bending westward to include
a considerable portion of Prince George County. Along this line the
Upper and Lower Austral are inextricably interdigitated. The river
bottoms and extensive swamps as far as the Second Swamp on the
Blackwater River are Lower Austral, inhabited by more or fewer of
the Lower Austral species. But the higher ground between the river
bottoms lacks these Lower Austral species and may therefore be con-
sidered as representing irregular fingers of the Upper Austral. As
far as the butterflies are concerned, the Eastern Shore (Accomack and
Northampton Counties) is Upper Austral.

In Virginia the area corresponding to the Upper Austral elsewhere
is in reality nothing more than a featureless faunal no-man’s land.
It has been wholly deforested, and the greater part has long been
under intensive cultivation. Such patches of woodland as are found
are various second-growth associations.

There are no butterflies that are confined to the Upper Austral
Zone in Virginia, all the species found within that zone being either
of very general distribution or else occurring also in the Lower
Austral Zone, more rarely also in the Transition Zone. The Upper
Austral Zone can therefore be defined only in negative terms as a
region between the Transition and Lower Austral Zones from which
the species characteristic of these two zones are absent.

The sporadic occurrence in widely separated localities within the
Upper Austral Zones of such primarily Transition Zone species as
Strymon edwardsti, Hesperia metea, H. leonardus, Atrytonopsis
hianna, and Amblyscirtes hegon, and until recently the general occur-
rence of Colias philodice, suggest that originally most of this area
belonged to the Transition Zone and that the clearing-off of the
forests and other plant cover, and consequent exposure of the land
surface to the drying effect of the hot sun in summer, have rendered
the region unsuitable for the original Transition Zone species, and
at the same time have led to the extensive immigration of hardy
Lower Austral types.

Indeed, we have seen this very process taking place within the
past 20 years. With the invasion from the south and west of Colias
eurytheme the original population of C. philodice has now almost
completely disappeared by replacement and apparently also through
extensive hybridization, though the two subspecies still remain dis-
tinct for the most part in the Transition Zone. Over this area, too,

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 7

the yellow swallowtail (Papilio glaucus) is so excessively variable
that typical examples of the northern forms may be found early in
spring, and typical examples of the southern form in summer, with
all possible intermediates. This suggests that originally the region
was inhabited by the northern form, but the clearing of the land was
accompanied by infiltration of the southern form with resultant
hybridization, so that now the entire Upper Austral is inhabited by
a hybrid population, although the northern 1-brooded forms still
persist in the mountains and the southern 2-brooded form is domi-
nant in the Lower Austral.

The occurrence throughout the so-called Upper Austral in Vir-
ginia of hybrids between the two species of Colias, and of presumed
hybrids between northern and southern forms of Papilio glaucus,
emphasizes the fact that in Virginia this zone is a secondary faunal
complex resulting from the clearing of the land over an area orig-
inally divided among three life zones, the Transition and the Upper
and Lower Austral. These unnatural conditions characteristic of
the Upper Austral in Virginia occur generally from southern New
Jersey southward along the Piedmont to North Carolina.

Lower Austral Zone-—The Lower Austral Zone is represented
at the present time in Virginia by more or less restricted and some-
times isolated areas—river bottoms, extensive swamps, marshes, and
pine barrens—which are surrounded by a greater area of intensively
cultivated higher country supporting the easterly extension of the
generalized and featureless Upper Austral fauna. These Lower
Austral localities are confined to an area that includes the outer
Coastal Plain as far as Cape Henry, and the western shore of Chesa-
peake Bay as far as Mobjack Bay. Its western limit may be ap-
proximately indicated by an irregular line from the head of Mobjack
Bay southwestward to south-central Greensville County, in its cen-
tral portion bending westward to include the Second Swamp on the
Blackwater River near New Bohemia. This area as outlined may be
defined as an area for the most part Upper Austral within which,
though not beyond its borders, Lower Austral species occur in suit-
able localities. There is no really definite boundary between the
Upper and Lower Austral such as those between the Upper Austral
and Transition or between the Transition and the Canadian. It is
possible that the western side of the southern tip of the Eastern
Shore peninsula should be included in the Lower Austral, for Dr.
Frank Morton Jones has found here a colony of Papilio palamedes.
Farther north in the Dahl Swamp in Accomack County we found
Atrytone alabamae common; this is presumably a Lower Austral

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

species, although it ranges north to the vicinity of Lakewood, N. J.

The characteristic Lower Austral species in Virginia are: Eupty-
chia areolatus areolatus, E. gemma, E. sosybius, Lethe portlandia
portlandia, L. creola, Agraulis vanillae, Calephelis virginiensis, Atlides
halesus, Papilio palamedes, Atrytone dukesi, A. dion, A. palatka,
Poanes yehl, Amblyscirtes textor, A. carolina, and A. alternata.
The index species, common and of general distribution throughout
the zone, are Papilio palamedes, Lethe portlandia portlandia, L.
creola, Poanes yehl, Amblyscirtes textor, and A. carolina. The other
species characteristic of this zone are mostly of local occurrence.
Although they are primarily characteristic of this zone, Euptychia
areolatus septentrionalis, E. gemma, E. sosybius, and Lerodea eufala
occur sparingly beyond its borders.

In Virginia it is possible to distinguish three divisions of the Lower
Austral. Along the coast just behind the sand dunes there are ex-
tensive sedge marshes with scattered willow and other shrubs. Here
Minois pegala pegala and Limenitis archippus floridensis are the
characteristic butterflies. Farther west in the great gum (Nyssa
aquatica) swamps along the North Landing River occur Airytone
dukesi, A. palatka, and, on the west side of the Dismal Swamp,
Euptychia areolatus areolatus and Amblyscirtes alternata. These two
subdivisions of the Lower Austral as a whole are characterized by
southern species that are restricted to the extreme southeastern part
of Virginia.

Not properly included within any of the faunal regions are the
two salt-marsh butterflies, Panoquina panoquin and Poanes aaron,
which occur in all suitable localities along the coast, and also Poanes
viator, found abundantly in all brackish marshes with an abundance
of wildrice.

THE MAP

The accompanying map (fig. 1) of the faunal zones of Virginia
has been prepared wholly from our personal records. The Canadian
Zone begins at an altitude of about 4,000 feet in Highland County,
and at an altitude of about 4,500 feet near the southern border
of the State.

The lower border of the Transition Zone is at an altitude of about
1,500 feet in the northern portion of Virginia, rising to about 2,000
feet in the south; its lower border is higher on the eastern face of
the mountains than on the western.

The Lower Austral is wholly on the Coastal Plain.

VIRGINIA

ore ae oe cameramen —

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Os ae in Ls iY
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Paes ee

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: we La ‘
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"Upper Austral.-«----2*=-

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st rvoeuat SURVEY

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lower We.......c=

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ic i

VIRGINIA

Fic. 1—Faunal zones of Virginia

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NO. 7

BUTTERFLIES OF VIRGINIA—-CLARK 9

The counties are numbered and may be identified by reference to
the following list.

. Accomack.

. Albemarle.

. Alleghany.

. Amelia.

. Amherst.

. Appomattox.
. Arlington.

. Augusta.

. Bath.

. Bedford.

. Bland.

. Botetourt.

. Brunswick.
. Buchanan.

. Buckingham.
. Campbell.

. Caroline.

. Carroll.

. Charles City.
. Charlotte.

. Chesterfield.
. Clarke.

. Craig.

. Culpeper.
25. Cumberland.
26. Dickenson.
27. Dinwiddie.
28. Elizabeth City.
29. Essex.

30. Fairfax.

31. Fauquier.
32. Floyd.

33. Fluvanna.

al
ODO ON An AWN 4

eS oe
dO FR

|
OO ON AM Bh W

bo bw HH
&whHD

THE COUNTIES OF VIRGINIA

. Franklin.

. Frederick.

. Giles.

. Gloucester.

. Goochland.

. Grayson,

. Greene.

. Greensville.
Petlalitas:

. Hanover.

. Henrico.

. Henry.

. Highland.

. Isle of Wight.
. James City.

. King and Queen.
. King George.
. King William.
. Lancaster.

. Lee.

. Loudoun.

. Louisa.

. Lunenburg.

. Madison.

. Mathews.

. Mecklenburg.
. Middlesex.

. Montgomery.
. Nansemond.

. Nelson.

. New Kent.

. Norfolk

. Northampton.
. Northumberland.

68.
69.
70.
aT
72.
73:
74.
75:
76.
77+
78.
79.

Nottoway.
Orange.

Page.

Patrick.
Pittsylvania.
Powhatan.
Prince Edward.
Prince George.
Princess Anne.
Prince William.
Pulaski.
Rappahannock.
. Richmond.

. Roanoke.

. Rockbridge.

. Rockingham.

. Russell.

. Scott.

. Shenandoah.

. Smyth.

. Southampton.
. Spotsylvania.

. Stafford.

. Surry.

. Sussex.

. Tazewell.

. Warren.

. Warwick.

. Washington.

. Westmoreland.
. Wise.

. Wythe.

York.

KEYS TO THE BUTTERFLIES OF VIRGINIA

The following keys to the butterflies found in Virginia are in-
tended for use primarily by those who are not specialists in the
Rhopalocera. They are therefore based for the most part on the
color and color pattern, by which most of the local butterflies are
readily distinguished. In the family Hesperiidae it has not been
practicable to include the females in the keys. These may be identi-

Io

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

fied in the field through association with the males. Although often
differing more or less widely on the upper surface, most females
may be identified by a comparison of the under side of the hind wings
with that of the males. But the identification of many of the smaller
species of Hesperiidae is by no means easy and whenever possible
should be checked by comparison with authentic specimens or by
someone familiar with the group.

Id.

Ib.

2a.

2b.
3a.

3b.
4a.
4b.
5a.
5b.
6a.
6b.
7a.
70.

8a.

8b.

. Key To THE FAMILIES AND SUBFAMILIES OF BUTTERFLIES REPRESENTED

IN VIRGINIA

Head of moderate width or narrow, the front less than twice as broad
as high; antennae separated by about the width of their bases or
LEGS OVEV CLASHES |S 0:5 leg occ jors od ale, makgganeretenapeds eas siete leleis tien Sete rere 2

Head very broad, the front twice as broad as high; antennae sepa-
rated by 2 to 4 times the width of their bases; a strong but slender
tuft of lashes in front of the eyes (Hesperiidae)............eee0e> 12

Fore legs in the males or in both sexes imperfect, the foot without
normalsterminaleclawSes <r calc ete emcee aise aeravonicicle eh aeieneeaee 3

Fore legs perfect and used in walking by both sexes................ II

Head large, nearly or quite as broad as the thorax; bases of the
antennae not notching the eyes; fore legs in the males, and usually
also in the females, greatly reduced and brushlike, not used in

Wallet ey 5, SEAS athectotorsth ce arsieS hala breve eceketate opens et stereo cleo chai e eatey tetera 4
Head small and narrow, narrower than the thorax; bases of the
antennae notching the eyes: size smalls. fcc. «ee oe esse oles 8

Palpi not elongated; fore legs of females as well as males defective... 5
Palpi greatly elongated, more than half as long as the antennae; fore

legs of males greatly reduced and brushlike, of females perfect.....

aietalerafeualsraloweieiercie a8 5b gy le veiay aes Walden haan atic re SUB Vectsnerers lsvenelmtevenave ress LIBYTHEIDAE
Antennae clothed with scales; body not spotted with white.......... 6
Antennae without scales; body spotted with white; size large......

a piesa uad scclalia A/a ani > nolo 0oc (ate tad elie haneedes Rim coer crete ane) sie DANAIDAE (key 4)
One or more of the veins of fore wings enlarged or swollen (Sa-

PV LIMAE) biased « vk Sia tensceth/ouc oes eras Ok tegen eis aac t ead cae eae Ee ae i
None of the veins in the fore wings enlarged or swollen.............

MAMET COTE oe APATURIDAE, NYMPHALIDAE, and ARGYNNIDAE (key 4)
Costal vein of the fore wings enlarged and stout, gradually tapering

distally. ..sex Sai icoreuiene eae beac ela eo ats elena LETHIINAE (key 2)
All the veins of the fore wings abruptly swollen at the base..........

Be AAC ROE Pat eer ae Sh wir hetin nce: SATYRINAE (key 3)

Fore legs of males greatly reduced and brushlike, of females perfect;
no white ring about the eyes; antennae not ringed with white; wings
held horizontally: >whemnl atreSt.42).6 1. «evisisiele lets oteret> RIODINIDAE (key 5)
Fore legs of males with defective terminal claw only; a conspicuous
white ring about the eyes; antennae ringed with white; wings held
vertically when jt rest Guycaenigae) .<cc% ca os esas ces sine emielobee 9

. Antennae short, less than half the length of the fore wings, the seg-

ments not much longer than broad. ;Vc2 0. -- es te es SPALGINAE (key 6)

NO.

ob.
10a.
IoD.

Ilda.

11b.

12a.

12b.

Ia.
Ib.

2d.

2b.

Ia.

1b.
2a.

2b.
3a.

Fe BUTTERFLIES OF VIRGINIA—CLARK II

Antennae at least half as long as the fore wings, the segments twice as
FotigtaswbtaniunG tancen eens guts Sales cele aide ese Ose eles ie 10
Hind wings rounded at the anal angle, or at least without a conspicu-
ous lobe, tailless or with a single tail.............. LYCAENINAE (key 6)
Hind wings with a conspicuous lobe at the anal angle, or with two
PeLALS 2 acts ASE es atte OT 5 NS LS Sista ahelehs be alowed ale THECLINAE (key 6)
Fore legs with the tibia (outermost long joint or segment) bearing a
leaflike structure, the epiphysis, on the inner side; hind wings with
conspicuous tails; wings chiefly dark brown or black, if yellow or
white, then with transverse dark stripes......... PAPILIONIDAE (key 8)
No epiphysis on the tibia of the fore legs; hind wings without tails;
wings white, yellow, or orange, with or without a dark border, but
Never with) trans Verse StCIPES...<iiciejarciedcicseve cre-seieeeioess PIERIDAE (key 7)
Distal recurved portion of the antennal club as long as, or nearly as
long as, the proximal part, the club being bent in its proximal part;
abdomen shorter than the hind wings; males often with a recurved
fold on the costa of the fore wings containing androconia..........
ae aiercca ie nictetnits Metetoe Siar atte malais slaw cigtees suideteaiss EP VRGINAE(key:'9)
Distal recurved portion of the antennal club much shorter than the
proximal part or absent, the club being bent beyond its thickest
part; abdomen as long as, or longer than, the hind wings; males
usually with an elongate black velvety patch on the fore wings con-
Rai PE ANGLOCOMIA .). 2)! wes VS Ab Ak ee eee ks HESPERIINAE (key 10)

2. KEY TO THE SPECIES OF LETHIINAE

Farid pwings with ai slight: angle atevelt 4s si. sine de cllacewadeleele sees ce 2
Hind wings with the border evenly rounded (pl. 1, 7, j)..........
BGC OD at De oto Aap oT Dia) aca Da mac cS aD rmeh acted te Lethe eurydice (p. 31)
Under side of fore wings with the postmedian line produced in a single
sharp point on vein 4; sexes essentially alike (pl. 1, e-h)..........
RSE ES CAS SCE Ee eer te ttc eRe Gere reer T Lethe portlandia (p. 30)
Under side of fore wings with the postmedian line produced outward
opposite the cell, with an angle on vein 4 and another on vein 6,
and a more or less straight line between them; males with the fore
wings more pointed than those of the females and with broad patches
of androconia in the inner portions of the interspaces on the fore
wings, running out to a point along the veins (pl. 1, a-d)........
to PEAS CON EIOD ADEE AA DOE aCMAP ene nic Lethe creola (p. 30)

3. Key TO THE SPECIES OF SATYRINAE

Fore wings with a broad yellow band in the outer half including two
large eye spots (rarely one) (pls. 2, f-i; 8, e)....Minois pegala (p. 32)

Ten wellow,s Dalit. On tie, LOLS! WAHL Sisvre0,6 5:4 <0 w Rainn ou npndcelnuniw sieleip nie wistele & 2

Upper surface with two conspicuous eye spots near the apex and lower
angles of the fore wings, and two similar eye spots near the upper
and anal angles of the hind wings, or a single eye spot near the anal

PaO ES ie Tl oe 3's hs 9 vwiiaalaalon tamil Euptychia cymela (p. 38)
Upper surface without eye spots, plain brownish or grayish.......... a
Wnder surtace with conspictious Eye Spots.......««cs.scacteevcce ses 4

3b.

4a.

4b.

4b.

5a.

50.

6a.

6b.

7a.

7b.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Under surface without eye spots; upper surface of hind wings with
three black marginal spots; hind wings below with marginal silver
matings i(pltig.ig2h)!. tnevs'. doth chee eee Euptychia gemma (p. 36)

Under surface of fore wings with a conspicuous circular eye spot near
the apex, followed by several less conspicuous ones; under surface
of hind wings with a submarginal row of 5 circular eye spots of
which the one near the outer angle and the second from the anal
angle are black-centered and conspicuous (pl. 3, 7).......-..eeeees
Tip Halaiciotelele lobe lo vs slate saliieelen et aE ee Euptychia sosybius (p. 38)

Under surface of hind wings with a row of large oval or elongate eye
spots, the three central the largest, the whole series surrounded by
a irregularly .oval. reddish ring (pliig,, a=d ign inc . otic ee
fe Seatostetatctel ae talon (ave. wiieicin ove ve nip ets intone Mi Euptychia areolatus (p. 36)

4. Key To THE SPECIES OF APATURIDAE, NyMPHALIDAE, ARGYNNIDAE,
AND DANAIDAE

. Under side of hind wings with numerous conspicuous silver spots... 2
. Under side of hind wings without numerous conspicuous silver spots... 6
. Fore wings much less than twice as long as broad..........eseeeeees
. Fore wings very long, more than twice as long as broad; dull reddish

with black markings; cell of fore wings with three black spots with
mihite centers Col <Fie, Pic. scgumce ek sceee eet Agraulis vanillae (p. 64)

. Hore wings more than 30 mim: longest cvtera'oc cause ose cee aeiee sees 4

Fore wings less than 25 mm. long; wings above dull orange with a
black border and fine black markings (pl. 9, a-c)..Boloria selene (p. 62)

. Fore wings above reddish orange with delicate black markings; hind

wings above blue-black with two rows of spots, both white (females)
or the outer orange and the inner white (males) (pl. 7, i, 7)...... :
DAS TH actasne wie alee tie lean Sao eee Bie Speyeria idalia (p. 54)
Both fore and hind wings above similar, brownish golden with dark
Dprongal tenting, rs. aras'aie sin ca tials emane Gee oe mas aed ee eee 5
Under side of hind wings with the light-yellow submarginal band
broad, occupying the entire space between the two outer rows of
silver spots, and at least twice as broad as the silver spots on either
Sule: Gols: 2216F Wg WD aiendk Seas Sas Speyeria cybele (p. 58)
Under side of hind wings with the light-yellow submarginal band
narrow, not so broad as the largest silver spots on its inner side.... 26
Both hind and fore wings strongly angulated; under side of wings
vermiculated light and dark, barklike in appearance, without any
definite’ color pattentic. sia. cate cre aiR eee ore eae a
Hind wings not angulated; under side of wings not vermiculated and
barklike; under side of fore wings, and often of hind wings, with
the same color pattefn' as. the upper sidey. 22.02.22 0c. ee eee ee II
Above dark maroon with a broad yellow border followed by a row of
small metallic blue spots; large, fore wings about 40 mm. long
Cpl 2b ORE Poe Se see 2 ee eee ..+.Nymphalis antiopa (p. 42)
Upper surface of wings reddish, the fore wings bordered with dark
brown, the hind wings more or less extensively dark brown, the
light areas on both wings with large dark-brown spots............ 8

NO.

8b.

Qa.
ob.

10a.

100.

IIa.

11D.
12a.

12b.
13a.

13).

I4a.

14D.

15a.

7 BUTTERFLIES OF VIRGINIA—CLARK 13

Larger, the fore wings 30-35 mm. long; basal quarter of costal border
of fore wings beneath conspicuously mottled with pale yellow and
brown; wings narrowly edged with pale violet; center of under
surface of hind wings with a silver semicolon (pl. 16, fig. f).......
Be Ae ee ee Oe ROE rr er eee Polygonia interrogationis (p. 40)
Smaller, the fore wings about 25-27 mm. long; no mottling on costal
border of the fore wings beneath; silver mark in center of under side

efile witieeENUGGs <5 250 Sas kceceles dnd Saleh ae lalee © pee Seeds oe 9
Silver boomerang-shaped mark in center of hind wings beneath abruptly

broadened tateeach end 5 o.2.cicts is. Siscke ak ako emiaceie teclee ote eae eee 10
Silver boomerang-shaped mark in center of hind wings beneath

pointed atweachvend iG@pliys. 62d) syne erica Polygomia progne (p. 41)

Under side of wings not very dark; fore wings with the border below
the subapical angle, and the hind wings, with the border on either
side of the tail, but slightly irregular at the veins; tails of hind
wings triangular with converging sides; no submarginal chevrons
on under side of wings (pl. 5, ¢, f)........06. Polygonia comma (p. 40)

Under side of wings very dark, mostly blackish; borders of wings very
jagged, with a process at each vein; tails of hind wings twice as long
as broad, with parallel sides; a row of dark green submarginal
chevron-shaped marks on the under side; dark spots above large
ROMS: war Deron hal. vate seancweeek wee Polygonia faunus (p. 41)

Above dark brown or blackish; fore wings with a bright red or orange
band from the inner third of the costal border to the lower angle,
and several subapical white spots; hind wings broadly bordered with
bright red or red-orange; fore wings angulated, 25-30 mm. long
MLROAT ieee Rises Oo fe laesk OE Vanessa atalanta (p. 43)

Noibnighttred orordnge abovess 6... ose cde ae adiiewigelns clement es 12

Hind wings above with two large and conspicuous eye spots; fore
wings with a large and conspicuous eye spot in a broad whitish
band; fore wings angulated, 25-30 mm. long (pls. 4, h-j; 5, g, h)

SC OD OCTO RODD COO. COCO ODOT ICC OC OR ORDO tn ore Junonia evarete (p. 45)
Hind wings above without two large and conspicuous eye spots....... 13
Wings above uniform dull orange to brownish orange or dull mahogany

with black veins and a broad black border including small white

Sy EL eis Se et it Re Eee RA ae eee Cen peices Ay Feo dae 14
Wings above not dark orange with black veins and a black border
Ie Hiei. Stall WEEE: SHOES sx wiv devs ie asd wwe iad wer teeta oleracea 6 15

Larger, the fore wings about 50 mm. long; black border of wings with
a double row of small white spots; no black line across hind wings;
thorax spotted with white (pl. 10, a-b)........ Danaus plexippus (p. 65)
Smaller, the fore wings 35-43 mm. long; black border of wings with
a single row of white spots; hind wings crossed by a narrow black
line parallel to the margin in the outer third (pl. 6, d-f)...........
earn beh ee ca vice caves fame esas oy eens Limenitis archippus (p. 52)
Under side of hind wings uniform brownish yellow or steel blue, lighter
in the outer third, the two sections separated by a thin silver line;
a row of thin silver submarginal dashes; sometimes a silver spot in
the cell and another at the inner third of the costal border; above,
inner two-thirds of wings black, outer third orange (males), or steel

14

150).

16a.

16D.

17a.

17).

18a.

18b.

19a.

19).

20d.

20b.

21a.

2ib.

22a.

22b.

23a.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

blue with light-blue and whitish spots in the outer third (females) ;

fore wings 43-55 mm. long (pl. 7, a-d).......... Speyeria diana (p. 55)
Under side of hind wings not uniformly colored—variegated, or with

several spots in the cell, or with a row of submarginal spots or dots;

HO “Sil Ver -: si .25 ca SAE a hw) vd Rietaiavniess ew’ phorst pata eter hei eat ae 16
Wings above blackish, becoming metallic bluish or greenish on outer

third of hind wings; under side with a few large spots in the cell,

and hind wings with a submarginal row of orange spots; rarely with

a white band across the fore wings, or across both wings; fore wings

35-45. mm, lone (pls. 2,106,619, 0) adeeeen Limenitis arthemis (p. 49)
Wings not uniform blackish unmarked except in outer portion; no
metallic’ sreenCOr \DIMek isis, clerisvs uo ieesvale Gisle tiske eee ete ee eee 17

Black with an interrupted brick-red border and numerous whitish
spots in the outer third of the wings; below with the whitish spots
continued inward to base of wings; fore wings 25-30 mm. long
(0) Ge PY NS Aires nr SORES ae RY Hon ey Euphydryas phaéton (p. 47)

Ground color of wings above not black—yellow, orange, reddish, gray-
ish, or brown; no brick-red border or uniformly distributed small
Wihitishr Spots! soe thes Sab Sole Be ee steht ee aoc le eee 18

Wings above grayish, reddish, or orange brown, the fore wings with
dark lines and numerous light spots, the hind wings with a sub-
marginal row of blackish spots, which may be concealed by the dark

CLouild (COLO MIE Esaki ews Silo a cide eS ae eels Sele Mela ae ee eels 19
Wings above with the ground color light, yellow, orange or reddish,
withedani= linestandeother amarkingsin: sees see aoeilo se eemela steerer 20

Grayish brown; fore wings with a rounded black spot in the outer
portion of the interspace between the two lowest veins; fore wings
25-30 tam done iplicaire) eso oes Ueto cee Asterocampa celtis (p. 39)

Reddish or orange brown; no rounded black spot on fore wings; hind
wings may be uniformly dark; fore wings 20-32 mm. long (pl. 4,

pb) he Bad Pas tne. a les Sah ae eae Asterocampa clyton (p. 39)
Under side of hind wings with conspicuous submarginal eye spots,
the inner portion grayish with an intricate reticulated pattern....... 21
No eye spots on under side of hind wings...............0ceesenccces 22
Under side of hind wings with two large and conspicuous eye spots
CDA VODA KR eeM Aes Ke MANE RRE UR ORSRNRERRE Vanessa virginiensis (p. 44)
Under side of hind wings with a series of five or six submarginal
eye spots; fore wings 30-35 mm. long (pl. 4, d, €).......-..ceeeeee
PASE SRST RI Sabeeted ORS Stole te a oe ete Vanessa cardui (p. 44)

Under side of hind wings variegated purplish; wings above with an
intricate pattern of black lines and a row of small submarginal
black spots; fore wings with an obtuse angle below the apex, about

es min long (ploy dye) aoa oe eee eee Boloria toddi (p. 61)
Under side of hind wings yellowish or whitish, usually variegated with
prow sso Je OP SES ind s Gee ees d Mee Wacod Saat a et oan mae tere 23

Size medium, the fore wings usually 30-35 mm. long; brownish yellow,
above with a submarginal row of rounded black spots, each in a
large pale spot, on both wings; bordering this interiorly is a line of
large dark-bordered pale spots; cell of fore wings with a large
pale dark-bordered spot at the end and a smaller one near the
middle (pl. G, GO) sadoxs walocenae stems oe mer Euptoieta claudia (p. 63)

NO.

23b.

24a.

24b.

25a.

25).

26a.

26D.

Ta.

Ib.

Id.

Ib.

2b.

3a.

3b.

7 BUTTERFLIES OF VIRGINIA—-CLARK 15

Smaller, the fore wings not over 25 mm. long; no submarginal rounded
ACH SIE GI, OTE: AUER chaps cis sptalic Syosa;e\a,oseh'c, ay erpobyasnrheoy ath dl acne debates 24
Under side of hind wings clear bright yellow, almost unmarked ; above
very dark, the blackish markings exceeding the yellow in area, at
least on the fore wings; fore wings 17-18 mm. long (pl. 11, f-i).....
AOC DAIATISE 6 AO APO RO SERS ADO Runs CHC OcE DEmrTe Phyciodes batesii (p. 49)
Under side of hind wings with more or less extensive dark-brown
lines or other markings; a marginal pearly crescent between veins
3 and 4; on the upper surface the yellowish ground color exceeds in

areas theudarisrmar kine specspattss isin Vetaicibic’« ajalarardtelaale askngeehe Sees 25
Submarginal spots on hind wings above uniform, all solid black, fore
wings 15-20 mm. long (pl. 11, j-m)........... Phyciodes tharos (p. 48)

Submarginal spot on hind wings between veins 3 and 4 in the form of
a ring and larger than the others; fore wings 18-25 mm. long (pl. 9,
PR aN PATO So lS ora. a eins cle acted’ oad alaveeane cd Melitaea nycteis (p. 47)
Border of wings marked with a narrow black submarginal line, the
veins from this to the margin more or less broadly black; fore

wings 35-43 mm. long (pl. 8, a, b).......... Speyeria aphrodite (p. 59)
Border of fore wings solid black, of hind wings mostly black; smaller,
fore wings 30-33 mm. long (pl. 8, c,d)......... Speyeria atlantis (p. 61)

5. Key To THE SPECIES OF RIODINIDAE

Fringes of fore wings uniform brown, at most with an indefinite light
spot at the apex; ground color above uniform brownish red; small,
the fore wings 9-12 mm. long (pl. 12, 7)...Calephelis virginiensis (p. 69)

Fringes of fore wings with three conspicuous white spots, one at the
apex, one in the middle, and one at the lower angle; ground color
above dark purplish brown with a broad vaguely defined darker
band across the middle of each wing; larger, the fore wings 15-16
thie MORO C AOE) Sac cvs «cob Seemies ©e vinea ae ¢ Calephelis borealis (p. 69)

6. Key To THE SPECIES OF LYCAENIDAE

Fore wings bright coppery red with a brown border and a few black
spots; hind wings above dark grayish brown with a red submarginal
band's fore wines about 15, mm. lone (ply 15, [=%)j.cssese cesses
Pete ctevecnie iis (6 shove tel & apaisiat sJe'a/s10 aidudsate.cionei cts sien eet etary eis Lycaena phlaeas (p. 71)

HOLE wills not DESH COPPELL, LEGsr cece sire cise aiise ele leinis eter citeysieeieee 2

Wings beneath blackish brown, the hind wings with a group of large
metallic green or coppery spots near the anal angle, the fore wings
sometimes (males) with a metallic green streak; above brilliant
metallic blue-green with a broad brown border; hind wings with two
tails ; fore wings about 20 mm. long (pl. 12, c,d) ..Atlides halesus (p. 77)

Under side of wings not blackish brown with conspicuous metallic
SROs GAS ii NS eee acer Sop eet totam Gch ote dace 3

Under side of wings dark slaty or brownish with a submarginal row
of large circular black spots conspicuously ringed with white; above
light silky blue (pls. 14, g; 15, a-c)..... Glaucopsyche lygdamus (p. 72)

Under side of wings without a submarginal row of large circular black
SEES RIAO, SUITS WHILE: iu ale sia we vip SON nin ais Wale wines Salemi lates @anewley 4

16

6a.

6b.

7a.
7b.
8a.

8b.
Oa.

ob.
10a.

10b.

Iida.

I1b.

I2d.

12b.

13a.

130.

14a.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Under side of hind wings plain light blue-gray or brown with black
OES. aicienie Sb caik favceateters cre oeee tam gna oaaeiel ond Oe Te ee ee ee 5

. Under side of hind wings brown or gray or green ornamented with

fine lines, never with dots’ 2cwid clea ala cake one eae Fi

. Under side of hind wings gray-brown or brown with a submarginal

row of large orange-red spots; above plain gray-brown or brown
(OIE; {G5 Weck SRE OR ae ae Strymon titus (p. 79)

. Under side of hind wings light blue-gray without submarginal orange-

Peds spots? ais .ov AS ak eit APS wins, see ee eee 6
Hind wings rounded, without tails; above clear blue or (summer
females) blue with the costal and outer borders of the fore wings
broadly brown (pl. 14, a-f, h).........ceee0e. Cyaniris argiolus (p. 73)
Hind wings with a hairlike tail at the base of which is an orange spot;
above blue or brown, if brown usually with some blue near the
HodyGplietAay a tf) sac, aeasea se lead aa ete BRAS Everes comyntas (p. 75)
Hind ‘wings with two. thim hairlike:tailsi... i200. sk ve ee ee 8
Hind ‘wings withott tailsi). sce sak esas Sc eee eee eee 15
Above brilliant metallic blue with a brown border (pl. 12, e, f)......
siateiatess otebetancts a etatavotamtreeieioiclend ioe, Gols eect nee Eupsyche m-album (p. 78)
Above brown, sometimes tinged with dull blue................ceeeeee 9
Under side of fore and hind wings with a broad bright-red stripe
outwardly bordered with white just beyond the middle; more or
less dull blue on the inner portion of the wings above (pl. 12, h)...
ssa sole tach covet cat taeestate cote chepterarces Ne niin 1h ol PA Ang ee ae Sirymon cecrops (p. 78)
No redband: on wits below. 265.1. «ca: cee donee ae ree 10
Under side of hind wings mostly bright green, with two very irregular
conspicuous white lines; a conspicuous white streak on the outer
border of the fore wings parallel to the margin; above golden brown

with a dark-brown border (pl. 12, g)......... Mitoura gryneus (p. 84)
Under side of hind wings brown or gray; an orange spot between the
pases GP thectails is ..:5:5.2.0052e aden dea eyed oan eee ee ee ee II

Under side of wings gray with a fine line of white, interiorly bordered
with black and brownish red in the outer third, and a large orange
spot above the interval between the tails; above gray with an orange
crescent at the base of the tails (pl. 11, , 0)...Strymon melinus (p. 81)

Wings brown or blackish brown, beneath with a conspicuous blue patch
between the orange spot and the anal angle...............eceeeeee 12

Under side of wings with four evenly spaced irregular fine white lines
in the outer two-thirds and a complete row of small submarginal
Orange or maroon) spots, (pl. 13,1é)). nee eiecles = Strymon liparops (p. 81)

Under side of wings with a narrow submarginal white line or series
of dashes, and a single or double continuous or interrupted white

lide in the outer thicd.c, <0 scarsocnacictepeiiee ecole aie eee 13
A broad dark bar narrowly bordered with white at the end of the cell
Of each wing’ on) the wader Sides «. J.c.neacssiot cess, eee ee serosa 14

Under side with no markings in the cell; the two white lines in the
outer third of the lower surface of the wings continuous and con-
spicuous; fore wings above usually with a dull-orange patch, or at
least some dull-orange scales (pl. 13, f)....... Strymon ontario (p. 79)

Inner white line continuous or nearly so, forming the outer border

NO.

14).

15a.
I5b.
16a.

16b.

17a.

20b.

¥ BUTTERFLIES OF VIRGINIA—-CLARK i7
of a dark band; sometimes a finer white line along the inner border
OGLchisypandy Cpl: D3,16))isetars sereyacisieteree eis « ceareare tee Strymon falacer (p. 80)
Inner white line double, equally strong on each side of the dark band,
and interrupted, forming a series of disconnected white-ringed spots,
especially on the hind wings (pl. 13, d)...... Strymon edwardsti (p. 80)
Eindswinesi without a lobevat the atiallangles.ctee-rcleel-/elee este ictete + 01= 16
Hind wings with a rounded lobe at the anal angle................... 17
Under side light grayish green with an irregular bright-red band in the

outer third; wings above brown, the hind wings more or less exten-
sively blue in the posterior portion; antennae half as long as fore
wings, with elongated segments (pl. 12, a, b)....... Erora laeta (p. 85)
Under side of hind wings coarsely checkered brownish yellow; above
black with a large irregular fulvous patch in the center of the fore
wings and a large fulvous patch in the outer portion of the hind
wings, these patches sometimes occupying most of the wings; an-
tennae less than half as long as fore wings, with short segments
CAT a Defias, oS <veea abe atcvahc) SEE eae a Feniseca tarquinius (p. 70)
Under side of hind wings with the inner half similar to the outer,
both crossed by irregular or jagged very dark lines, the central one
narrowly bordered with white outwardly, giving an irregularly check-
ered effect; border of hind wings conspicuously scalloped; ground
color of under side with a purplish cast (pls. 13, b; 30, h,7).......
ay ieeyas sles ahi cNe ls Sy.sirary va: 6 niaisioje\s tel aisaun MA SEE ange reEe Incisalia niphon (p. 82)

. Under side of hind wings with the inner half uniformly colored and

darker than the outer half, from which it is separated by a very ir-
POOMAT MEGe doyirs'c asc. 04 ura v «40d Sloteleie c Sante Mielec RCRA eLearn 18

. Hind wings with an even and unscalloped border, beneath plain brown,

lighter in the outer half (pl. 16, e)......... Incisalia augustinus (p. 83)

. Hind wings with a scalloped border, beneath with an overwash of whit-

ish scales in the lower two-thirds of the outer half................ 19

. Under side of fore wings with a narrow light-gray line along the

OuULeL DOLGEr CD13, @)) s.0:s = ceisler sieve tiasicter spies Incisalia polios (p. 83)

. No narrow light-gray line along the outer border of the fore wings

16d bo Anis ces eae eigen an atric Sanco ues core ae 20

. Hind wings beneath only slightly darker in the outer half; outer half

of lower two-thirds thickly flecked with whitish scales; more or less
evident white scaling in a broad band along the inner margin; band
across middle of fore wings below scalloped and very irregular;
male with a conspicuous stigma on the fore wings (pl. 16, c).......
aie a aro tata ee slele oes cae es aha site aus alee Aorta Incisalia irus (p. 84)
Hind wings beneath with the inner half very dark brown, contrasting
strongly with the much lighter outer half, uniform in color or with
a large indefinite area of olive or lighter-brown scales near the
anterior border; hoary scaling confined to a broad definite or indefi-
nite band in the hinder two-thirds of the wing which seldom extends
beyond the row of spots or crescents in the light outer portion;
band across under side of fore wings in middle of outer half not
scalloped, parallel to the outer border, usually with the lower half
abruptly nearer the outer border than the upper; male without a
stistay GolviG; @) oii vinta ive wee de cate ween ee Incisalia henrici (p. 84)

18

Ia.
1b.
2a.

2b.
3a.

3b.

4a.

4b.
5a.

5b.

6a.

6b.
70.

7b.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

7. KEY TO THE SPECIES OF PIERIDAE

Ground icolor- above, whites... / rer oirs aero aioe ele teeter bie 2
Groundicolor above notswihiteseparnactesis se eeieler nee biekiie een eer 9
Small, the fore wings 15-20 mm. long; hind wings below marbled or
reticulated with green or erayishverecn: se. ../ecce cele cles viele 3
Larger, the fore winestmoremthan! 20 “mine longa cliente sites ebienele 4
Fore wings with the outer part of the costal margin curved downward

to a sharply pointed apex; a small black spot at the end of the cell;

hind wings below mottled gray-green and white; males with the

apex of the fore wings broadly orange (pl. 0, 1, 7)......-c.seeeees

Pea eratadieee ceehre ahamhicte a ata e tet etele cig ave wage chee Anthocharis genutia (p. 86)
Fore wings of normal shape, the apex rounded; apical portion of

fore wings broadly dusky; a black bar across the end of the cell;

hind wings below coarsely reticulated with apple-green and basally

finved with pink (plir6fab)* 123..80e, oP ee Euchloé olympia (p. 87)
A black spot or bar at the end of the cell of the fore wings........... 5
No black spot or bar at the end of the cell of the fore wings........ 6

A broad rectangular black bar across the end of the cell of the fore
wings; a large black spot on the lower margin of the fore wings
in the outer third; fore wings otherwise plain or nearly so, or with
a border about 5 mm. wide of broadly black veins and white inter-
spaces; no orange or yellow spot in center of hind wings above; hind
wings below pure white, or more or less marked with yellowish, or
(in spring) white with broad dark-gray veins (pl. 18, c-h)........
G-aie horaexe. ste aes tape feiatats Loe retaaateleuatsictaieheinieistt tise teak tetihs Pieris protodice (p. 92)
A small black spot at the end of the cell of the fore wings; no black
spot near the lower margin; border of fore wings broadly black
with included white spots; hind wings above with a conspicuous
orange or yellow spot in the center; hind wings below uniform whit-
ish or yellowish, usually more or less speckled with dark scales
and with a central circular pearly spot ringed with brown, and a
smaller supplementary spot behind it (pls. 10, e; 17, e, f)......++.-
Erctahar Se eseeIeR: white females of Colias eurytheme or philodice (pp. 93, 97)
Fore wings with the apex black, a rounded black spot halfway be-
tween the lower end of the cell and the outer border and another
below it between veins 1 and 2, the latter sometimes absent; hind
wings with a small black spot on the outer third of the costal
bordetuGpls To.0@) pes tis anita lat cetaieeree women mae eee Pieris rapae (p. 88)

Fore wings without conspicuous rounded black spots..............+- 7

Fore wings with black triangles at the outer ends of the veins, longest
at the apex and disappearing after vein 2 (males), or with the
apex broadly blackish, the black decreasing in width but persist-
ing to the lower angle, and the costal margin blackish, giving off
a narrow black line following the anterior border of the cell and
curving about its outer end; hind wings plain white (males) or with
black triangles at the ends of the veins (females) ; fore wings about
30mm. long (pli8,9=1))s sew aatiowel eyaner aes Ascia phileta (p. 93)
Above plain white, unmarked; fore wings 20-25 mm. long........... 8

NO.

8a.

8b.

Oa.

ob.
10a.

rob.
IIa.
11b.

12a.

12b.

13a.

13.
14a.
14D.

15a.

15).

7 BUTTERFLIES OF VIRGINIA—-CLARK 19

Apex of fore wings faintly grayish; under side of hind wings plain
white or faintly yellowish, usually with scattered dark scales (pl.
IS: Was « SUMO eed veRdeeE bee ss Pieris rapae (early spring) (p. 88)
Under side of hind wings white, the veins broadly bordered with light
grayish; no dusky scaling at apex of fore wings above (pl. 18, b)

Biadik le Veh Supiralatdi afeisidte, dhain wees akan «gpk Pieris virginiensis (p. 90)
Ground.color above dark gray: (pl. '8,.g5/A) ics sc%s susleasdsecisaw oedanes

stich hth a: thaberahrasd my 955) ian ars Sexes exdbeicass Dark females of Ascia phileta (p. 93)
Ground color above yellow or orange.....:...cccccecccacceccsvcces 10
Fore wings without a broad black or brown border; large, the fore

OIE] Shes Hal OTT) BON is syeu azole eiare apetstecietacycroicls suctarstecotere oisievese « attewiet ss II
Fore wings with a broad black or dark-brown border; smaller, the

tore wings seldom: so) mttch as) 30 mim: lONG. 6c... 0.0000 + cle avis ce « 12
Bright clear yellow, unmarked (pl. 10, c)........ Phoebis sennae (p. 112)
Bright yellow; fore wings with a large orange patch at the end of

the cell, hind wings broadly bordered with orange (pl. 10, d)......

BioSeta it tidal Mintle sw aGi ces wie eee sas aati Phoebis philea (p. 112)

Hind wings below with a central circular pearly spot ringed with
brown or red, above with an orange or yellow spot at the end of

the cell; fore wings with a black spot at the end of the cell........ 13
No central circular pearly spot in center of hind wings below, or orange
or yellow spot at the end of the cell of the hind wings above....... 14

Fore wings with the apex sharply pointed and a very broad black
border with a sharp angle extending inward toward the middle of
the cell and a truncated angle extending inward between veins I
and 2; a large black patch including most of the cell, reaching the
costal border anteriorly and extending inward and downward to the
base of the wings; yellow (pl. 2, d)........... Zerene caesonia (p. 112)

Fore wings with the apex rounded, the black border, broad or nar-
row, with an even inner edge, plain or including yellow spots; yel-

lows on orange «(temales sometimes! witite) ine. ce eiertoele serie sie e' wei 17
Lower border of fore wings with a broad dark-brown band.......... 15
No dark markings except for the dark border and a small spot at the

enusotthe celltob thettore wists s ees cote ste ce cates strates etre 16

Dark band on lower portion of fore wings broadest outwardly, taper-
inging to a point at the wing base; a narrow dark border on inner
two-thirds of the costal border of the hind wings; apex of fore
wings broadly blackish, the blackish portion bordered by a straight
line running from about the middle of the costal border to the end
of vein 3; under side yellow; small, the fore wings 12-15 mm. long
MIPRRE ly FEN oes Waia'.d bau bas At wih eae siteew eee Nathalis iole (p. 115)

Dark band on lower portion of fore wings sharply defined with nearly
parallel borders and a rounded outer end, separated from the lower
border of the wing by a narrow brownish-orange line; no dark line
on costal border of hind wings; inner border of black apical por-
tion of fore wings a curved line running from the middle of the
costal border to the lower angle; under surface of hind wings silky
white; larger, the fore wings about 17 mm. long (pl. 16, 7,7).......
SOOT OTe RETO Re ar sche rice ne Drier Eurema jucunda (p. 117)

20

16a.

16D.

17a.

17).

Id.

Ib.

2a.
2b.

3a.

3b.

4a.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Larger, the fore wings 23-25 mm. long; blackish border on fore
wings narrowest in the middle, on hind wings with an angle extend-
ing inward toward the cell; bright orange (males rarely bright
pellow) (pk 2%) Voie eis See eee ee Eurema nicippe (p. 115)

Smaller, the fore wings 16-20 mm. long; black border of fore wings
with the inner edge an even curve from just beyond the middle of
the costal border to the lower angle; black border of hind wings
narrow, tapering to a point at the anal angle; bright yellow, the
females sometimes whitish, rarely pure white (pl. 9, h)...........
Rlataralaneid spa cin ls Siva ore lates ree eis alee et taele eRe ee Eurema lisa (p. 116)

Under side of fore wings with a row of submarginal black dots
approximately under the inner margin of the black border on the
upper side; spot at the end of the cell of the fore wings conspicu-
ous, more than half as high as the width of the cell; pearly spot in
center of under side of hind wings ringed with red-brown and with
a small supplementary spot on its anterior border; fringes dull,
more or less pinkish; color yellow or orange (pls. 10, e; 17, a, b,

CO; FRCS GR Oe Colias eurytheme or philodice (pp. 93, 97)

No submarginal black dots on under side of fore wings; spot at
the end of the cell of the fore wings small and obsolescent, not half
so high as the width of the cell; pearly circular spot in center
of under side of fore wings ringed with red and without a supple-
mentary spot; fringes bright rosy red; color light yellow (pl. 17,
CROs WM crareita Ralole We iehe RST ah ee Colias interior (p. 111)

8. Key TO THE SPECIES OF PAPILIONIDAE

White or greenish white with broad dark-brown bands and a scarlet
spot at the anal angle; tails long and slender, half as long as the
hind wings or longer, with straight and slowly converging sides

De TOG ina toeiclae cea eeeals Meera mae Graphium marcellus (p. 145)
Color not white and dark brown; tails shorter, much less than half

asvlone: asp the hind Switig's accu teiein aiersicsnicreiclor recor eieienelatornic ta intense terete 2
Under surface of the wings chiefly yellow...........ceeesecccccceces 3
Under surface of the wings black or blackish brown...............+- 4

Upper surface of wings yellow with a broad black border including
yellow submarginal lunules or broad dashes; fore wings with four
black bands extending downward from the costal margin, only
the innermost reaching the lower margin; hind wings with a long
narrow tapering black stripe and the inner margin narrowly black
(pis 10) jo 200s 21. Wt). tcken aieeice ene see Papilio glaucus (p. 124)

Upper surface of wings brown; fore wings with a broad band of yel-
low from the apex to the inner third of the lower border, and a
curved row of large submarginal spots; hind wings with a row of
large yellow spots in the outer third and a broad transverse yellow
band across the basal portion; a large yellow spot on the tails
PLEO. 12) wteiain «els whtaishekiae waeeiehs REIN ee Papilio cresphontes (p. 120)

Under surface of the wings with a single submarginal row of orange
or yellow or orange and: yellow spots, ois <\i0 cic s.a/s6ddeis as d.sjeis wists 5

NO.
4b.

5a.

5b.

6a.

6b.

7a.

7b.

Id.

Ib.
2a.

2b.

7 BUTTERFLIES OF VIRGINIA—CLARK 21

Under surface of the wings with two rows of conspicuous orange or
orange and yellow spots parallel with the margin.................. 6
Outer half of hind wings below metallic steel-blue with a curved
submarginal row of large rounded spots set in black, each with a
small silver patch on the upper side; wings above brown or black-
ish becoming metallic steel-blue or green on the hind wings; hind
wings and sometimes (females) the lower half of the fore wings
with a submarginal row of small light spots (pl. 10, a-d)........
Meee AS ee Che ea cet ade b cee irre ee eo erate ae Battus philenor (p. 118)
Outer third of hind wings below darker than the proximal two-thirds
from which it is marked off by a narrow black line bordered out-
wardly by a narrow interrupted metallic-blue line; submarginal spots
consisting of stout orange and yellow crescents near the margin;
hind wings above with scattered blue scales which become much
more dense beyond a conspicuous narrow black line along the
outer third; a conspicuous orange spot at the outer angle (pls. 10, f;
DTMEAG ee a a tae Pelneetonen s dark females of Papilio glaucus (p. 124)
Upper surface of wings with two rows of yellow spots, the large spots
of the inner row usually coalesced into a yellow band on the hind
vans CE! ME HOSEN 2 GR Re Ns Pa IY 7
Upper surface of the wings with a single submarginal row of green-
ish spots; hind wings above crossed in the middle by a broad vaguely
defined band of grayish green (males) or with diffuse metallic-
blue scaling (females) ; a conspicuous orange spot on outer third
of costal border of hind wings (pl. 22, a, b)..... Papilio troilus (p. 122
Inner row of large yellow spots on the fore wings in a straight line
parallel to the smaller spots of the submarginal row; yellow band on
hind wings crossed by black veins or (females) represented by a
row of spots; orange spot at the anal angle with a conspicuous black
center; no yellow line on inner portion of hind wings below (pl. 22,
Baa aoa ce ORD te kk cele oie Hed Hae ee ae ee Papilio polyxenes (p. 144)
Inner row of large yellow spots on the fore wings in an irregular
line, the three uppermost and the lowest displaced outwardly, the
second lowest displaced inwardly; yellow band on hind wings con-
tinuous, not crossed by black veins; inner portion of under side
of hind wings with a narrow yellow line from the inner portion of
the costal border to just above the spot at the anal angle (pl. 22,
CRO) RRC IP GS eee Nees lonen es Papilio palamedes (p. 121)

9. KEY TO THE SPECIES OF PyYRGINAE’

Hind wings with the anal angle produced into a long tail more than
half as long as the wing; fore wings with a row of angular yel-
lowish spots from the middle of the costal border to the lower angle,
and a few smaller spots toward the apex; fore wings 20-23 mm.

MOT SGD ASSIGN) ech er evsm: ole icavoete. Sata als Velavela Stele Urbanus proteus (p. 148)
tae matin cu ihiotth: AAlSii¢t viebde dx «ae sin «le duclenes aided aeena tae ete’ <iers 2
Ground color brown or blackish; no conspicuous white spots on the

Htindh Garis sal: Wehcted vet si gad «et.al daw 1 kiss Seen apa A 3

Wings above gray, both fore and hind wings with conspicuous white
gio bo uch eee SRO E eee S TORIC Ae OAD OCH baal Oboe nUnOTa on. 19

22

3a.

3b.

4a.

4b.

5a.
5b.
6a.
6b.
7a.

7b.
8a.

8b.
ga.

ob.

10a.

rob.

Ila.
IIb.
12a.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Fore wings with a conspicuous broad yellow or brownish-yellow band
or row of large angular spots running from the middle of the costal
border to the lower angle; size large, the fore wings more than
22 mm. long; dark brown, the hind wings above unmarked........ 4
Fore wings without a conspicuous broad yellow or brownish-yellow
band or row of large spots; smaller, the fore wings not over 20 mm.

TONG wie onde: slate. aids leolandiaie ats oahbie'nletichaphebie bosentan iclawty docks Sane ieeteae 6

Band on fore wings of irregular width, brownish yellow; hind wings
below: withiavconspicuotis! whiter patchinss sass cer ac iclels eielerserorelelateeielete 5

Band on fore wings with almost parallel sides, 3-4 mm. broad, lustrous
golden; hind wings below dark brown with a few scattered light
spots near the outer border; fore wings about 25 mm. long (pl. 23,
Dab) sawisre seae leone niereieia baw pean ot ad Meenas Rhabdoides cellus (p. 149)

Hinds wings below with a large irregular central patch of bright
silvery ; fore wings 25-30 mm. long (pl. 23, a)..Proteides clarus (p. 147)

Hind wings below with the outer third chalky white; smaller, the
fore wings 22-25 mm. long (pl. 23, e)...... Achalarus lyciades (p. 148)
Fore wings above uniform brown or blackish, the fore wings with
tisually) «small white spotsesh deideres aoe eeinels eee etoile ere eeaerere 7
Fore wings above with numerous grayish scales or mottled black-
ish or brownishvand plighten saat om scciatsve told ele shale cos sl etetel ol ehetel ae eeteneteta II
Fore wings 18-22 mm. long, with a few to many angular white spots.. 8
Smaller, the fore wings less than 15 mm. long............seecceeese 10
Fore wings crossed by a conspicuous narrow interrupted line of largely
contiguous white angular spots running from the middle of the
costal border to the lower angle (pls. 23, g, h; 25, €)....-.seeeee
svotand ba: Zitipts dia s.ctatatete ume eee ralieveelanareie acorns wletemtes Thorybes bathyllus (p. 149)
Fore wings with the white spots small and widely separated.......... 9

Spots on the fore wings irregularly arranged, but present from the
costal border to the lower angle; under side of hind wings without
numerous whitish scales (pl. 25, a, b)........ Thorybes pylades (p. 150)

Spots on the fore wings in a straight line from the middle of the
costal border to the lower angle; outer portion of hind wings more
or less ‘dusted’ with ‘whitislr ‘scales “(pl!’ 25. c,d). 22: ese es eee oeee
JAA eto Coen tonic cer ICIS rica vce, 6 Ok Thorybes confusis (p. 150)

Wings of normal shape, uniform silky blackish, the fore wings with
more or less numerous minute circular white spots (pl. 26 7)......
SOAP CHICIO oN TON ERE HE SILOS Chicas Pholisora catullus (p. 153)

Hind wings with a coarsely crenulate border; wings clouded darker
and lighter; fore wings with two minute white spots near the apex
and another just below the middle (pl. 15, t, 7) .........ssscsseess
eriehadh ehegovers ve (1c ablinvaper Ne atthe (evetovaletaiees ote etarate Pholisora hayhurstti (p. 153)

Fore wings with no subapical: glassy Spots’. ssi22: wiese ale as ee eens s 2) ie

Fore wings with subapical glassy spots... <s'. deeweea: sna as eee 13

Smaller, the fore wings rarely more than 17 mm. long; a distinct hoary
patch between the upper halves of the two bands crossing the upper
surface of the fore wings; males with a pencil of hair on the hind
tibiae; costal fold of males with flagellate tapering scales (pl. 24,

Oy. Ve Disc aetna gee SERRE aes eee Erynnis icelus (p. 154)

|

NO.

12).

13a.

13b.

14a.

14).

15a.

150.

16a.

16b.

17a.

17d.
18a.

18b.

Iga.

19b.

Ze BUTTERFLIES OF VIRGINIA—CLARK 23

Larger, the fore wings rarely less than 17 mm. long; no distinct hoary
patch between the upper halves of the two bands crossing the upper
surface of the fore wings; males without a long pencil of hair on
the hind tibiae; costal fold of males with twisted ribbonlike scales
CS eI cite wei tee eae sae ween ees Erynnis brizo (p. 155)

Larger, the fore wings seldom less than 17 mm. long; costal fold of
males with no rodlike androconia, the androconia being at least twice
asescole apically as basally. si). %.< lee seo sha Mare ee areata ete ieee eyes 14

Smaller, the fore wings rarely more than 17 mm. long; costal fold
of males with many rodlike subequal androconia, sometimes two-
pronged, apically not twice as stout as basally..............seeeee- 16

Upper surface of fore wings flecked with whitish scales throughout
so as to be quite different in general tint from the hind wings; hind
wings below with numerous light spots in the outer third and usually
with one or two small circular light spots near the outer third of
the costal border (pls. 24, 0, p; 25, f)......-. Erynnis juvenalis (p. 157)

Upper surface of fore wings flecked with whitish scales only in small
patches or not at all; hind wings below with the spots in the outer
third obsolescent or absent, and no small circular spots near the
GOStAIMDOLGEE Pree se ceic osins Gre-storeta Sinlote cyohite aeteie ate siererersve eiehcle cfete's? oye) state 15

Glassy spots on the fore wings conspicuous; under side of hind wings
with faintly marked light spots toward the margin (pl. 25, j, k)...

+ dil Es SEER PEI NC a aa eR ree Aes ACE tee Erynnis horatius (p. 158)

Glassy spots on the fore wings small and inconspicuous; under side
of hind wings almost or quite plain blackish (pl. 25, g, h).........
ep aS Beenie © bak w Gud oo on ul eke Coon cows tae Erynnis sarucco (p. 158)

Dark spots on upper surface of wings well defined and in sharp con-
trast to the ground color, giving a checkered appearance; costal
fold of males with many “apple-seed” androconia (pl. 24, 1)......

4 BOO COS CCI Ce HERR rain eRe aA ao cn AH eRe ror Erynnis martialis (p. 156)

Dark spots on upper surface of wings poorly defined and not sharply
contrasted with the ground color; costal fold of males without

RAMIG-SCCK ANGLOCONIA 2. s.0s vi hs e.g uss ween eden tc bese eee aan 17
Inner portion of wings with numerous long light grayish hairs; spring
Only MQple aang UA sh ele ctorcis te crtovaleale steithettrere ake Erynnis persius (p. 155)
Few or no light hairs on inner portion of wings; spring and summer.. 18
Markings of fore and hind wings fairly well defined (pl. 24, k).....
PELE ie slotha le OY or roe 6 hia eels a5 aie ard a ara wk eth aia Erynnis lucilius (p. 155)
Darker, the markings on fore and hind wings obscured and poorly
STC CDA TIN) oreo alata seat ne Cie ee mest Erynnis baptisiae (p. 156)

Fore wings with a broad white band formed of a series of spots
usually at least twice as broad as high and frequently much broader,
separated only by the dark veins, beyond which is a submarginal
row of white spots; fore wings about 15 mm. long (pl. 24, b, c, e, f).
sto6.c0 oC RECORDED EEO aT CIO DOME EADIE rte Pyrgus communis (p. 152)

Fore wings with an irregular series of discontinuous spots which are not
broader than long in the outer third, and no submarginal spots;
hind wings with a row of submarginal spots and a pair of elongate
spots beyond the end of the cell; fore wings 13-14 mm. long (pl. 24,
MEMO a a tr atele esis alk ois sale, tava ete erecine Gretta Pyrgus centaureae (p. 151)

24

Ia.
1b.
2a.
2b.

3a.

3b.

4a.

4b.

5a.

50.
6a.

6b.

7.

8b.

Qa.
ob.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

10. Key To THE MALES OF THE SPECIES OF HESPERIINAE

Under side of hind wings plain yellow, unmarked.................... 2
Under side of hind wings not plain yellow, unmarked............... 5
Small) the fore wings 12-14. mm longer ony eleca inne re eee 3
Larger, the fore wings..15-18, mim. Jogi. io ooo: <mai-ntanimpsin'cha silva apiece 4

Fore wings about 12 mm. long; fore wings above infuscated yellow
with a broad brown outer border; hind wings above clear yellow with
a brown border; fore wings below black with the costal and outer
margin reddish-fulvous (pl. 3, k)....... Ancyloxypha nwmitor (p. 159)
Fore wings 13-14 mm. long; both wings above clear yellow with a
brown border 1.5-2 mm. broad; fore wings below pale yellow, the
lower border clouded with brown (pl. 11, a, b)..Atrytone arogos (p. 172)
Fore wings above clear yellow with an even brown outer border
1.5 mm. broad, a dark streak along the inner half of the lower
border of the cell, and a narrow black line running from the outer
third of the costal border parallel to the outer margin to the lower
part of the outer end of the cell; no stigma; hind wings with a very
narrow brown border; fore wings below with the lower and inner
half of the baseiblack \( pli 27; (6-e))msjee eure Atrytone logan (p. 173)
Fore wings about 18 mm. long, bright yellow with a broad brown
even outer border 4 mm. wide and a narrow black stigma; hind
wings darker and duller yellow with a very broad brown costal

and outer border’ (pls 20, 0-1). aie «aisetnernnne es Atrytone palatka (p. 173)
Under side of hind wings yellow with black or brown spots or blotches ;

NON GATK WHORE Eis stare 's cele eteusi ee Gree CiterR eG eile a ci ae eI eee 6
Under side of hind wings not yellow with dark spots or blotches...... 9
Under side of hind wings evenly dotted with small black or brown

SDOUSI va iaj(c..0) shapcle caslolonecousvavsnans ea cps cspeusfeuin tet nKe cuss iris SS ichel acer oie keenest aie en een 7.
Under side of hind wings with small and large spots of red-brown
irregularly distributed ; fore wings 13-14 mm. long...............+- 8

Hind wings below spotted with black; fore wings 14 mm. long; above
bright yellow with a series of contiguous marginal triangles ex-
tending inward, a large black stigma, and a curved black line from
the stigma to the apex; hind wings with a narrow brown border
widening at the anal angle (pl. 28, c)....... Hylephila phyleus (p. 161)

Hind wings below with dark brown spots; fore wings 12 mm. long;
wings above dark brown, the fore wings with about 6 small yellow
SHOTS H(DIS..25, (PF 2O40=6) Addnetdeae cies ee Amblyscirtes carolina (p. 180)

. Wings above golden yellow with a brown border having an angular

inner edge; no stigma; hind wings below with a large brown spot

at the base sending off a branch to the inner third of the costal

margin, a narrow irregular interrupted brown border, and usually

a few small spots about the middle of the inner half (pl. 27, a, b)..

LHR ete HS Ae ta 'ah Seatecmpcis 4 Sic Brcta Whale NOOR ic Oey Ee Poanes zabulon (p. 169)
Above yellow, marked as in H. phyleus (7a); hind wings below with

two large brown spots near the outer margin, one near the inner

margin, and one or two small spots above the latter (pls. 27, f;

2 AE Vis, cicksive! cratptal-i vik wre minelaye eieid inte MiSeTeis cn teeta Polites vibex (p. 167)
Upper side of wings plain dark brown or black, unmarked............ 10
Upper side of wings not plain brown unmarked..............0.200005 13

Idd.

15a.

15D.

16a.

16b.

17a.
17d.
18a.

4 9 BUTTERFLIES OF VIRGINIA—CLARK 25

Larger, the fore wings 17 mm. long; hind wings below with two
long light streaks diverging from the base to the outer margin

GDI 20s Ged) ht ds eis GR See eas das Atrytone dukest (p. 175)
Smaller, the fore wings not more than 14 mm. long; no streaks on

the unden suntace of the: himd) wines da ei nls ernie) -/lertlale sleisjots cleo eyes II

LH ROLELWIN Sea boute 4 vintitsl 11s farts) aticleye deja helaele oaks anaes edi beieie cannes 12
Fore wings about 12 mm. long; wings unmarked above and below

(GINIEAG: HOD ak Seeiztes azaeto pieces eee Lerodea l’herminieri (p. 181)
Hind wings below plain brown, unmarked (pls. 11, e; 26, ¢)........

B Hyadh Sta balsas ceal SAe ahaha) Ste ori Soua ihe. Sivas NR wrod Atrytone ruricola (p. 177)

Hind wings below with a chevron-shaped row of contiguous bright-
yellow spots in the outer third and a yellow line running from
the base of the wing through the angle of the chevron (pl. 26,
ela ac a an OOOO ACE ORD oc Hoe Poanes massassoit (p. 172)

. Under side of hind wings without spots or sharply contrasting mark-

ings, plain, with one or two long ill-defined light streaks, with
whitish veins, or with light veins and a sharply defined light streak.. 14
Under side of hind wings with spots or sharply contrasting markings.. 21

. Under side of hind wings with one or two long ill-defined light streaks

LUNN PatEOmNtoespase tonthenmag eit sey. es, sco sre sees neeracen s+. 15

. Under side of hind wings without ill-defined light streaks running

FLOM Hes MasentOthey May mrbecperveerae aa aiee swayed she o waists reread a as 17
Both wings above bright yellow with a uniform brown border about
2 mm. wide; fore wings without a stigma, 13-17 mm. long; hind
wings below with a single median light streak (pl. 28, m-o, r)......
SS ORIG 20 CE ITO ICYD ERT EIRS PR ICTR GED Cee Recent Poanes aaroni (p. 170)
Wings above dark brown, the stigma bordered outwardly by a con-
spicuous yellow band; a narrower yellow band just within the
costal border from about the middle of the cell to beyond the end of
the stigma; hind wings with a broad yellow line from about the
middle of the cell to near the outer margin; hind wings below with
two diverging ill-defined light streaks from the base to the outer
OGG trom rete teverctceneyes Hareiato: cosy oats: avo eis cater chad harass oles cue chee SePseaeere 16
Yellow band bordering the stigma outwardly about 2 mm. wide, nearly
as wide as the brown border between it and the margin, and broadly
united with the yellow stripe near the costal border; hind wings
above with the yellow band about 5 mm. long; fore wings 16-17 mm.
Longa Gple2o we) Ae anette. Sos aeereou aes sews Atrytone dion (p. 173)
Slightly smaller and darker; yellow band bordering the stigma out-
wardly about I mm. wide, much narrower than the brown border
between it and the margin, not connected with the yellow sub-
costal line; yellow band on hind wings about 2-3 mm. long, or

more or less obsolete (pl. 20, f)..........+ 02 4trytone alabamae (p. 174)
Hinde wings below splaiti brown: W4s« siys.w Varlcncle vigswad Ge rene oeaees 18
Find wings below net plaitt- Brow? «coc ¢sa% sal duu snes Hecate 19

Larger, the fore wings 17-18 mm. long; fore wings above with a
conspicuous white spot concave outwardly below the end of the
cell, a small one beyond its upper end, one below the large one near
the lower border, and one or two small ones near the apex (pl. 26,

PAYER ctetehass eAis leparsys lors Uiausrecape LP orate, Woot. eigrents Panoquina ocola (p. 186)

18D.

10a.
19b.

20a.

20b.

21a.

2ib.

22a.

22b.
23d.

23D.
24a.

24b.

25a.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Smaller, the fore wings 13-14 mm. long; above more grayish, the
fore wings with two small white spots below the end of the cell and

three very small ones in a row near the apex (pl. 24, q)..........
eR aR Bis Use GME O OE OOS OU ee nae Lerodea eufala (p. 182)
Hind wings below grayish olive with white or yellowish veins....... 20

Hind wings below with an ill-defined dark long-triangular patch ex-
tending inward from the outer third of the costal margin to beyond
the middle; fore wings above with a few small white spots in the
subapicalireeion (ply 27) oF) 7) eee cetie ects sees Lerema accius (p. 178)

Hind wings below with a broad white streak from the outer part of
the cell to the margin; above grayish brown, the fore wings with
numerous whitish spots in the outer half, the hind wings with a
whitish streak from the outer part of the cell to the margin (pl. 26, /)

Si RR Ne aerate hasetiahslanatahelatanatas atoattctoratihs Panoquina panoquin (p. 185)

Hinds wings below with white veins but no broad white streak; fore
wings below with the lower half of the basal portion as far as the
middle black, the upper portion of the black area bordered with two
white spots, the lower the larger; fore wings above with the stigma
bordered outwardly with dull reddish (pl. 27, 7, R)..........0000s
Stat sahatatNaledatirclaratctetateyelalatclitelenianc eure ce armioe amas Atrytone bimacula (p. 176)

Hind wings with a transverse row of three conspicuous spots both
above and below; fore wings with about 6 or 7 white spots, a large
one at the end of the cell, one about half as large about the middle
of the lower border, the others smaller and subapical; wings above
brown, lighter toward the body; large, the fore wings about 25 mm.

Lone AC PLS SIF) ic tehasnre "sta 0%e sararecatars 'o Gta trates tose Calpodes ethlius (p. 183)
Hind wings without a row of three conspicuous white spots on both
SUBTACES Es Scb AEE Oe et de eis Meche Aes PO eee aalae emt 22

Under side of hind wings black, the inner two-thirds with an intri-
cate white reticulation; fore wings with an angled line of small
white spots on the outer third; above black with an angled row of
white spots on the outer third-of the fore wings; small, the fore
wings 13-15 mm. long (pl. 3, 7)........... Amblyscirtes textor (p. 180)

Under side of hind wings not black with a reticulate white pattern.... 23

Fore wings above with the stigma in two well-separated parts; above
brown, the fore wings with an elongate yellow spot extending out-
ward from the anterior portion of the stigma and a smaller spot near
the apex; hind wings below yellow, or purplish brown, with a short
row of obscure light spots in the outer third (pl. 26, m-o).........
Bee Mes Me orc lsteichacs tancheterr ee elettvowioteiaters ortiele oe Wallengrenia otho (p. 167)

Fore wings above with the stigma continuous or absent............. 24

No yellow, above or below; markings confined to isolated spots, never
in ‘bands ‘or Stréalesty sc sale rato oo ocra hei reteetole: state stotevereiee clecctes erartererate 25

Fore wings above marked with yellow, broadly yellow or with a yel-
low band or row of large spots bordering the stigma on its outer
SIS ei cick iche Fete ceiclaele Sieve tie eee Catotere States cus Siete TeNe ore etait cvehate coceken eaetatane 28

Larger, the fore wings 15-17 mm. long; above brown with a few white
spots in the outer third of the fore wings; hind wings below brown
becoming hoary gray in the outer half; a small but conspicuous
white circular spot near the base, and usually another about half-

25.

26a.

26b.

274.

28a.
28b.
204.

29b.

30a.

30b.

31a.

3Ib.

32a.

a7, BUTTERFLIES OF VIRGINIA—-CLARK 27

way to the outer angle; spring only (pl. 26, q)....c.cccccwecesces

Steere acdc: Salat die amen ee Mean ebe we Atrytonopsis hianna (p. 177)
Smaller, the fore wings 10-14 mm. long; above blackish with small

white spots in the outer third of the fore wings.................4. 26

Hind wings below pale gray with a semicircular band of white spots,
a small spot at the end of the cell, and another at about the middle

@b thercostals marein “springs (pls25; t= 275 bis strcie stelle eemiealeiere «
OR eid Se ees aS oseiciale ole starts Seats eRe Amblyscirtes hegon (p. 179)
Hind wings below blackish; spring and summer.................0000: 27

No spot in the middle of the costal border of the fore wings, and hind
wings immaculate; fore wings with 2 or 3 (rarely more) minute
white spots in the outer fourth of the costal margin; hind wings
below blackish, slightly mottled, clouded with pale-bluish scales,
especially in the outer third (pl. 27, n, g)....Amblyscirtes vialis (p. 179)

. A small white spot near the costal border of the fore wings at about

the middle; three minute white spots near the apex, and an irregu-
lar row of small white spots in the outer third parallel to the margin;
hind wings with a small white spot above the end of the cell, and

often additional spots (pl. 25, J, m)..... Amblyscirtes alternata (p. 179)
Hind wings’ abave plain brown, ‘unmarked. i eicien ositecgs oder ae ticles as 29
indiwines marked! -withiyellow aboverat..-cciccas declee ee cscawees ce 31

Hind wings below olive-gray with a curved row of obscure spots in
the outer portion; fore wings above dull golden as far as the minute
subapical spots and the stigma; an elongate spot at the anterior
end of the stigma and commonly other spots along its outer side.. 30

Hind wings below dark purplish brown with a curved row of paler
spots; fore wings above with three minute subapical spots and a row
of whitish-yellow spots along the outer side of the stigma, the upper
the longest and the lowest somewhat isolated, forming an interrupted
band about one-third as broad as the distance from the stigma to
the margin; fore wings 14 mm. long (pl. 28, d)...Polites verna (p. 164)

Fore wings 12-13 mm. long; stigma broad, short, and sinuous (pl. 28, f)

SESE, LES AE NE Se eae ere Polites themistocles (p. 165)
Fore wings usually 14-15 mm. long; stigma longer and narrower, al-
most a straight line (pls. 15, d; 28, e)..... Polites manataaqua (p. 164)

Hind wings below with a narrow irregular white band or row of
white spots forming a right angle, with the apex at about the middle
of the outer border, and a short white oblique line in the middle
of the wing; above dull, with a broad dingy-yellow band bordering
the stigma outwardly, an obscure angular line of dull-yellow spots
on the hind wings, and some minute dull-whitish spots in the apex
of the fore wings; fore wings 13 mm. long; spring (pl. 28, a, b)...
“HAE Ad 0275 ACE AREER OL EPL eee Hesperia metea (p. 160)
Hind wings not marked with white below; yellow above bright, in
Bite COMiLaSE FOSEME: DEGWOS wec-a njca0 so 0 ip dr weeks cinaveckytae oud’ 32
Fore wings yellow with a broad brown outer border and a very large
black oval stigma; hind wings brown broadly streaked with yellow;
under side of hind wings yellow with a few large obscure darker
Repos. tote wings 14 fom. lone (pl. 26, 7-4). .<se<cancse gedaee cine
SORES EME sae e wiee cay eh sedan eee bs Atalopedes campestris (p. 163)

34d.

34D.

35a.

35D.

36a.

36D.

374.

370.
38a.

38b.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Stigma ‘narrowly. linéarjor absent... <j. «eames eee tains eee 33
Under side of hind wings with a series of small clearly defined sub-
mareinal spots,afd.ese.in the, cells ph sisi pms iso's Haims here 34
Under side of hind wings without small clearly defined spots; no small
Spot, dir. the Gels vacherged a sik ire oe ssh Mab wlhale em ea nels panes eens 36
Hind wings below with an angular row of five to seven clearly defined
and conspicuous, SpOtSs;:.. > 2/20.) «mbiesie a abit biter © ee 35

Hind wings below pale cinnamon with four circular yellowish spots in
the outer portion and another in the cell toward the base; wings
above yellow with brown borders, on the fore wings anteriorly ex-
tending inward to the end of the cell; stigma and streaks in the
cell brown, the stigma sometimes with a yellow center ; three minute
yellow spots in the broad brown apical portion (very variable in the
extent of the yellow markings above) (pls. 28, q; 30, d, f, g)......-
eRe Sats BANS Ne itch te SRLS shee PE shel Seibel Aa Poanes yehl (p. 171)

Hind wings below bright cinnamon with an angular row of seven
clearly defined submarginal light spots and another in the cell; fore
wings above orange-yellow with a broad brown border, the apical
portion extending inward to nearly the middle of the costal margin,
and more or less dusky as far as the stigma; hind wings with the
central portion orange-yellow, which may be restricted to a band
in the middle of the outer third (pl. 26, a, b)..Hesperia leonardus (p. 160)

Hind wings below dark brown (or yellow) with an angular row of
usually five well-separated whitish circular spots, another in the cell,
and another above it near the costal border; fore wings above brown,
the stigma broadly bordered outwardly with yellow, and some yel-
low subapical spots; hind wings above with the angular line of small
spots belowsrepeated (pll 2754) 22). Soames ote Hesperia attalus (p. 161)

Hind wings above uniform yellow or orange-yellow with a broad
sharply defined border, or with the central portion occupied by a very

laree’ yellow tspOte Vise. vects tte cue cole Are eee See oe re oer eee 37
Hind wings above not uniform yellow with a sharply defined brown
ORO ire eeesaiactas a ahdgcie eke cake ad tere eciseet e eat Oe ON eae Ee rte a Pe 30

Fore wings above golden yellow with a brown outer border 3 mm.
wide including two small elongate yellow spots beyond the end of
the cell; no stigma; under side of hind wings yellow with a broad
brown border, essentially as above; fore wings 16-17 mm. long
UpleZG ete) adhe cists ter cach ne cg Sea heehee Ae nt Poanes hobomok (p. 169)

Fore wings above mostly brown, or with a conspicuous black stigma.. 38

Larger, the fore wings about 20 mm. long; hind wings below dark
gray with a broad more or less obscure light band from the base to
the outer margin and some light spots near its outer end; fore wings
above brown with a row of orange-yellow spots in the outer third
increasing in size downward, a large spot in the cell, and some small
Spots near therapex: (pll2o; 7) ances eee Poanes viator (p. 168)

Smaller, the fore wings 13 mm. long; hind wings below dull yellow,
the central portion with a large blotchy lighter patch; fore wings
above yellow with a brown outer border 2-3 mm. wide and a long
narrow black stigma (pl. 27, m, p).......... Hesperia sassacus (p. 161)

NO.

30a.

30b.
40a.

4ob.

7/ BUTTERFLIES OF VIRGINIA—CLARK 29

Hind wings below brown with a small group of elongate indistinct
light spots beyond the cell; upper surface of hind wings with similar
but better-defined yellow spots; fore wings above yellow with a
broad brown border 3 mm. broad, a dusky costal border, and a
large black stigma surrounded by yellow at its upper end; fore wings
about 15 mm, long) (pli 27 igs W)e sna.ce 4-2 Atrytone conspicua (p. 176)
Hind wings below with large sharply contrasting light areas.......... 40
Hind wings below brown with a broad and irregular submarginal
crescent of strongly contrasting bright yellow or orange-yellow, a
large spot in the cell, and another at the base of the wing, the mark-
ings often confluent; fore wings above as far as the stigma, and a
few spots beyond, yellow; hind wings above with a group of 4 or 5
elongate yellow spots alternating large and small in the outer por-
tion; fore wings 12-13 mm. long (pl. 28, g, h)..Polites peckius (p. 165)
Hind wings below with a broad band of straw yellow on the outer
half and a patch of darker yellow on the inner portion; outer por-
tion of fore wings below with a broad triangular patch of straw
yellow, the inner portion darker yellow; fore wings above yellow
with a brown outer border 2-3 mm. wide and a black stigma joined
to the apical portion of the border by two diverging lines; hind
wings above with the central portion yellow, sharply defined out-
wardly but inwardly becoming dusky; fore wings 14 mm. long
(Es ene ae Ta) oh le a alee ae ee a tae orth sin i Beeb Polites mystic (p. 166)

ANNOTATED LIST OF BUTTERFLIES OF VIRGINIA
Family SATYRIDAE

Subfamily LeTHImnaE
Genus LETHE Hiibner
LETHE CREOLA (Skinner)
Plate 3, a; bs co

Range—From southern Princess Anne County westward and
northwestward to Greensville, Dinwiddie, and Prince George Coun-
ties; our records are from Dinwiddie, Greensville, Isle of Wight,
Nansemond, Prince George, and Princess Anne Counties.

Occurrence.—Found in wet woods with an abundant undergrowth
of Arundinaria tecta; very local; common in certain localities along
the western border of the Dismal Swamp and near Petersburg and
New Bohemia, elsewhere not numerous. This species usually, but
not always, occurs with L. portlandia portlandia. It is easily over-
looked as it is crepuscular, seldom flying during the day but becoming
very active just before dark. It is readily secured by beating it out
of the cane in which it rests during the daylight hours.

Seasons.—Two broods. This species first appears in the second
week in June, flying until early in July. The second brood is on the
wing shortly after the middle of August and flies until the end of the
second week in September.

LETHE PORTLANDIA (Fabricius)

There are two subspecies in Virginia.

LETHE PORTLANDIA PORTLANDIA (Fabricius)
Plate a;1¢) 7

Diagnostic features—Row of ocelli on under side of fore wings
curved; second and third ocelli on under side of hind wings with
elongate pupils, the fourth usually without a pupil; under side of
fore wings with the row of eye spots edged with white interiorly and
more or less completely exteriorly ; a white band runs from the costal
margin to the region of the second ocellus, and beyond this a narrower
white band runs from the costal margin to the upper part of the first
ocellus ; discocellular vein closing the cell of the hind wings angled.

30

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 31

Range.—tThis subspecies is found on the southeastern Coastal Plain
from the sea westward to Prince George, Dinwiddie, and Greensville
Counties. Our records are from Dinwiddie, Greensville, Isle of
Wight, Nansemond, Norfolk, Prince George, Princess Anne, Surry,
and Southampton Counties.

Occurrence—Found in damp and wet woods and along streams
and roadside ditches wherever cane (Arundinaria gigantea) is abun-
dant; generally distributed and locally common—in wet seasons
abundant—from the coast to the western border of the Dismal
Swamp, becoming more local and less numerous farther west.
Though flying by day, especially on cloudy days, this species is most
active in early morning and at dusk.

Seasons—Three broods. This butterfly is on the wing almost con-
tinuously from the end of May until the end of September or early
October, becoming progressively more numerous as the season ad-
vances, and often abundant in September. The first brood appears
at the very end of May and flies until nearly the middle of July. The
second brood is on the wing shortly after the end of the third week in
July, and the third brood appears late in August; freshly emerged
individuals are found up to the end of September.

LETHE PORTLANDIA ANTHEDON A. H. Clark
Plate 1, g, h

Diagnostic features—Row of ocelli on under side of fore wings
straight ; all the ocelli on the under side of the hind wings with similar
circular pupils; no clear white on wings beneath; discocellular vein
closing the cell of the hind wings evenly curved.

Range.—Found in the Blue Ridge and in the Alleghanies, chiefly
at high altitudes. Our records are from Bedford, Botetourt, Giles,
and Madison Counties.

Occurrence——This subspecies inhabits damp wooded ravines and
steep damp wooded mountain sides with a growth of bottle-brush
grass (Hystrix patula). It is very local and not numerous, the widely
scattered colonies occupying limited areas and including few indi-
viduals. It is active during the day, though more or less crepuscular.

Season.—One brood. Our records are all from early July to early
August.

LETHE EURYDICE (Johansson)

The subspecies represented in Virginia is—

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

LETHE EURYDICE APPALACHIA R. L. Chermock
Plate 1, 2, 7

Range.—Found locally throughout the State, though chiefly in the
north and west. Our records are from Augusta, Fairfax, Frederick,
Giles, Grayson, Greensville, Henrico, Montgomery, New Kent,
Prince Edward, Prince George, Rockbridge, Stafford, Tazewell, and
Wythe Counties.

Occurrence.—In the mountains and on the Piedmont this butterfly
is found in permanently wet sedgy areas, almost invariably with
alders, in and near woods; it is very local and seldom common. On
the Coastal Plain it lives in wet woods and swamps with abundant
Arundinaria, always with Lethe portlandia portlandia; here it is very
local and, so far as we have seen, decidedly rare.

Seasons.—Probably one brood, with a few individuals emerging
late in the season possibly representing a partial second brood. Our
earliest date is June 17, and most of our records are in July. In
August the numbers decrease, but we have found it as late as Septem-
ber 19. At Mountain Lake, in Giles County, the earliest record is
July 4, and here the species is more common in the last half of August
than it appears to be elsewhere.

Subfamily SATYRINAE
Genus MINOIS Hiibner
MINOIS PEGALA (Fabricius)

There are four intergrading subspecies in Virginia.

MINOIS PEGALA MARITIMA (W. H. Edwards)
Plate 8, e

Diagnostic features—Fore wings of the females 28-30 mm. long,
of the males 23-27 mm.; three anterior eye spots under side of hind
wing circular ; color dark, the females usually as dark as the males.

Range.—From the vicinity of the District of Columbia locally
southward on the Coastal Plain to Dinwiddie, Sussex, Southampton,
Nansemond, and Norfolk Counties, where it is typically developed
and distinctive, though intergrading to some extent with alope; also
from Fauquier County westward at the higher altitudes in the moun-
tains, and southwestward to the southern border of the State, here
intergrading extensively with alope and seldom so typically developed
as on the Coastal Plain.

Variation—This small dark form is most typically developed in

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 33

the pine barrens of the inner Coastal Plain. Here the individuals are
very dark and are quite the same as others from similar situations in
New Jersey. As in New Jersey, males with the lower eye spot on the
fore wings greatly reduced or wholly absent are fairly common; such
males (pl. 8, e¢) also occur in the District of Columbia and in eastern
Massachusetts.

In the mountains this subspecies is somewhat less typical than it is
on the Coastal Plain, with the yellow band on the fore wings usually
somewhat lighter. In this area it grades insensibly into alope, so that
it is impossible to draw any definite line between them. In many
places specimens may equally well be referred to maritima or alope,
and over large areas on the Piedmont the first individuals to emerge
are small and dark and resemble maritima, while those emerging later
are larger and lighter and would unhesitatingly be referred to alope.

In a meadow just south of Monterey, Highland County, the indi-
viduals are very variable in size, in the size of the eye spots on the
fore wings, and in the color of the yellow band on the fore wings
which in fresh specimens varies from reddish to almost pure white,
though white-banded individuals are rare. Many of the smaller indi-
viduals in this meadow would be unhesitatingly identified as mari-
tima, others are equally typical alope. Such diversity in any one lo-
cality is unusual, though we have found cases in wet pastures in
West Virginia.

In Frederick County, and in the northern mountains, the males oc-
casionally have the yellow band on the fore wings somewhat narrowed
and constricted in the middle, the central portion of both the outer
and inner borders being concave. Some of these are referable to
maritima, others to alope.

In a small male from Short Mountain taken by Dr. Warren H.
Wagner, Jr., on August 3, 1940, the band on the fore wings is much
darkened, being about halfway between the normal clear yellow and
the general ground color of the wings. A similar male with the fore
wings 24 mm. long was taken by Dr. Wagner at Panorama on July
19, 1936. These two individuals, both of which were flying with
many typical maritima, approach the northern subspecies nephele,
and resemble some specimens determined as nephele from Beaver
County, Pa.

Occurrence.—On the Coastal Plain maritima is found in open and
more or less grassy pine woods, especially along their borders; it is
very local, though common wherever it occurs. In the mountains it
lives along the borders of spruce woods, about the edges of mountain
bogs and the brushy borders of mountain pastures, and also in open

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

bogs and wet pastures ; here it is rather generally distributed and com-
mon wherever it is found.

Season.—One brood. This subspecies appears usually shortly after
the first of July, sometimes as early as the last week in June, becomes
common the second week in July, and flies in gradually diminishing
numbers until early in September. The males emerge a week or more
before the first females, and disappear usually by the middle of Au-
gust when the females are still common.

MINOIS PEGALA ALOPE (Fabricius)
Plate: 2,ah,.g

Diagnostic features—Fore wings of females 31-34 mm. long, of
males 27-30 mm.; three anterior eye spots on under side of hind
wings circular; color usually somewhat lighter than in maritima,
with the females commonly lighter and more grayish than the males.

Range.—Throughout the State, except in the sedge marshes south
of Virginia Beach and in the highest mountain regions. In the moun-
_tains it usually occurs with maritima, into which it intergrades through
an unbroken series of intermediates. On the Coastal Plain it often,
though not always, occurs with maritima. West of the coastal
marshes, north of Virginia Beach, and on the outer coast of the
Eastern Shore it occurs with pegala, though the latter is the dominant
form; a short distance inland from the sea alope becomes dominant,
and intergrades between pegala and alope are common. On the
Eastern Shore alope occurs alone in the Dahl Swamp and probably in
the interior generally.

The range of this subspecies has the appearance of being largely
unnatural. Of the three subspecies of pegala this is the only one that
is thoroughly at home on cultivated land. It is probable that the ex-
tensive overlapping by this subspecies of the territory in which mari-
tima and pegala occur has been due, in part at least, to the clearing of
the land which has resulted in the infiltration by the most adaptable
subspecies, alope, of the territory normally occupied by the two others.

Occurrence.—F ound along the borders of woodlands, about groups
of bushes in open fields, and along streams through pastures; it also
occurs in open deciduous woods with an abundance of coarse grass,
and in patches of pines, especially scrub pines, avoiding thick woods
and open grasslands. Rather local, but in suitable localities usually
common, though seldom abundant.

Season.—One brood. This subspecies first appears between the end
of the third week in June and the first of July, the females coming

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 35

out about a week later than the first males, and reaches the peak of
its abundance in the last half of July and the first half of August.
Later in August the numbers decline, the males disappearing in the
last half of the month and the females flying until late in September.

MINOIS PEGALA CAROLINA R. W. Chermock

Diagnostic features—Resembling M. p. alope, but with the band
on the fore wings cream white.

Range.—The white-banded individuals from a wet meadow just
south of Monterey, Highland County, mentioned on page 33 probably
should be referred to this recently described form. The senior au-
thor has also found it, flying with M. p. alope, in Greenbrier County,
West Virginia.

MINOIS PEGALA PEGALA (Fabricius)

Plate 2, h, i

Diagnostic features—Fore wings of females 35-41 mm. long, of
males 29-32 mm.; three anterior eye spots on under side of hind
wings, or at least the two lower, oval. Throughout most of the range
in the south the lower eye spot on the fore wings is lacking in both
sexes ; in North Carolina most of the females have this eye spot more
or less developed, in northern North Carolina and Virginia all the
females have two eye spots, and the males begin to acquire the lower
eye spot, north of Virginia Beach both sexes having both eye spots
equally developed.

Range.—Coastal region of eastern Virginia from the North Caro-
lina to the Maryland borders. In the sedge marshes from Knotts
Island to Virginia Beach this is the only subspecies present. Just
west of the sedge marshes and along the coast north of Virginia Beach
pegala is the dominant form, though alope also occurs. Farther inland
it reaches the western border of the Dismal Swamp, where alope is
the dominant form, as it is away from the coast on the Eastern Shore.
Where the ranges of pegala and alope overlap there is complete inter-
gradation between them.

Variation.—In the sedge marshes south of Virginia Beach, where
pegala occurs alone, the females always have both eye spots on the
fore wings fully developed; in the males the lower eye spot may be
present, more or less reduced, or absent. North of Virginia Beach the
size decreases slightly, and the males usually have both eye spots on the
fore wings fully developed. In the sand-dune area between Virginia
Beach and Cape Henry very pale more or less grayish individuals

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

occur which appear to agree perfectly with certain specimens of the
form texana.

Occurrence.—In the sedge marshes south of Virginia Beach this
butterfly lives largely in the open, though always in the more or less
immediate vicinity of the scattered bushes in which it takes refuge
when alarmed. Farther north it keeps close to the seacoast, being
replaced by alope a short distance inland. Here alope also is found
along the coast, but in lesser numbers.

Season.—One brood. The males first appear in the last week in
June, becoming common early in July, when the first females appear.
Both sexes are common through July and the first half of August,
the males preponderating in July and the females becoming relatively
more numerous in August. The males disappear toward the end of
August but the females persist until the end of September.

Genus EUPTYCHIA Hiibner
EUPTYCHIA GEMMA (Hiibner)
Plate 3, g,h

Range-——From Princess Anne to Hanover and southern Cumber-
land Counties and southward; also southern Scott County. We have
records from Hanover, Henrico, Isle of Wight, Lunenburg, Nanse-
mond, Norfolk, Prince Edward, Prince George, Princess Anne, and
Scott Counties.

Occurrence.—Found on the Coastal Plain in wet pine woods; local
and usually infrequent, though sometimes common. On higher land
it occurs along streams and in damp valleys and ravines.

Seasons.—Three broods. Our records indicate that this species
first appears shortly after the end of the third week in April and flies
until about the second week in May. The second brood appears about
the middle of June and flies into early July. The third brood, in which
the individuals are more numerous than in the two preceding, appears
early in August, becomes common after the middle of the month,
and remains on the wing until early October.

EUPTYCHIA AREOLATUS (J. E. Smith)

Two subspecies occur in Virginia.

EUPTYCHIA AREOLATUS AREOLATUS (J. E. Smith)
Plate 3, b, d
Diagnostic features—Dark mouse gray, lighter below; eye spots

on under side of hind wings elongated, usually at least twice as long
as broad, often much longer.

INO: 37 BUTTERFLIES OF VIRGINIA—CLARK 37

Range.—Known only from two localities on the western border of
the Dismal Swamp and from Holland, all in Nansemond County.
We found it common about 8 miles south of Suffolk, and our friend
Otto Buchholtz took it at another locality on the edge of the Dismal
Swamp, and also at Holland.

V ariation.—At the locality about 8 miles south of Suffolk areolatus
occurred with septentrionalis. Most of the individuals were inter-
grades between the two, but some were typical areolatus agreeing
with specimens from South Carolina and Georgia, while others were
equally typical septentrionalis agreeing with specimens from New
Jersey.

Occurrence.—Found in grassy open areas in pine woods and along
the grassy borders of pine woods, especially along roads and railway
tracks.

Seasons—Two broods. Mr. Buchholtz found it on May 15, and
we took it on May 30. Mr. Buchholtz also took it on August 8 and
20.

EUPTYCHIA AREOLATUS SEPTENTRIONALIS Davis
Plate 3, a, ¢

Diagnostic features—Dark warm brown; eye spots on under side
of hind wings short and broad, from scarcely longer than broad to
about twice as long as broad.

Range.—Southeastern Virginia from Lunenburg, Nottoway, and
Dinwiddie Counties eastward to Nansemond County. Our records
are from Dinwiddie, Greensville, Lunenburg, Nansemond, Nottoway,
and Southampton Counties.

Variation.—Specimens from Virginia are usually slightly larger
than others from Lakehurst, N. J., with the eye spots on the under
side of the hind wings usually slightly longer, but many Virginia
specimens agree perfectly with others from New Jersey. As already
noted, in the locality about 8 miles south of Suffolk this intergrades
with the typical southern subspecies.

Occurrence-—This subspecies occurs in the same situations as E.
a. areolatus.

Seasons.—Two broods. We have found it in the last half of April
and on May 30; Mr. Buchholtz took it on May g. It also flies from
August 8 (Buchholtz) into September.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

EUPTYCHIA SOSYBIUS (Fabricius)
Plate 3, 7

Range.—Southeastern Virginia westward and northward to Meck-
lenburg, Prince Edward, Henrico, Caroline, King William, New
Kent, Gloucester, and Mathews Counties ; also southwestern Virginia
eastward and northeastward to Buchanan, Russell, Montgomery, and
Roanoke Counties. We have records from Buchanan, Caroline,
Gloucester, Greensville, Henrico, Isle of Wight, King William, Lee,
Mathews, Mecklenburg, Montgomery, Nansemond, New Kent, Nor-
folk, Nottoway, Prince Edward, Prince George, Princess Anne, Ro-
anoke, Russell, Scott, Southampton, Surry, and Sussex Counties.

Occurrence.—On the Coastal Plain this species is found every-
where in pine and mixed woods, least commonly in the very wet areas
preferred by E. gemma. It is common to abundant in all suitable
localities from Princess Anne to Prince George and Greensville Coun-
ties, becoming more local and less numerous farther north and west.
In the southwestern mountains it lives in wooded river bottoms and
in the lower portions of damp valleys; here it is local and not very
common.

Seasons.—Three broods. This species appears shortly after the
end of the third week in April and becomes numerous in the last half
of May, the numbers falling off in the first half of June. The second
brood appears at about the end of the third week in June, reaches its
maximum in the last half of July, and falls off in numbers in the first
half of August. The third brood, of which in most localities the indi-
viduals are most numerous, appears in the last half of August and
flies until the first week in October. Dr. Carroll M. Williams writes
us that he has found this species around Richmond only in spring.
Later in the season it disappears and is replaced by E. cymela, which
then becomes common. Farther east on the Coastal Plain E. sosybius
is the dominant species throughout the summer.

EUPTYCHIA CYMELA (Cramer)
Plate 3, e, f

Range.—Throughout the State.

V ariation.—Specimens from the Coastal Plain east of the Dismal
Swamp usually have larger eye spots than those from farther west.

Occurrence.—Found everywhere in open deciduous woods; gener-
ally distributed and usually fairly common locally, and in some years
common generally.

Season.—One brood. This species usually first appears, in very

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 39

small numbers, about the middle of May, very rarely as early as the
last week in April, and in the last week in May suddenly becomes
common. Increasing in numbers through June, it reaches a maximum
late in June and in the first half of July on the Coastal Plain, in the
last half of July in the mountains. It usually disappears in the first
half of August, though stray individuals are sometimes found as late
as the end of the third week in August.

Family APATURIDAE

Subfamily APATURINAE
Genus ASTEROCAMPA Rober
ASTEROCAMPA CELTIS (Boisduval and LeConte)
Plate 4, c

Range.—Northern and western parts of the State southward to
Essex, Henrico, Nelson, and Amherst Counties, and in the mountains
southwestward to the southern border. Our records are from Am-
herst, Arlington, Augusta, Botetourt, Essex, Fairfax, Fauquier, Fred-
erick, Henrico, Loudoun, Madison, Nelson, Page, Roanoke, Rock-
bridge, Shenandoah, Warren, Washington, Westmoreland, and Wise
Counties.

Occurrence.—Found in woodland glades, about the edges of woods,
and in groves including hackberry trees; very local and irregular in
its appearance, but usually common when and where it occurs; more
generally distributed and more numerous in dry than in wet summers.

Seasons——Two broods. The hackberry butterfly first appears in
the last week in May, becomes common in the last half of June, and
flies until about the middle of July. It reappears in the last half of
July and flies until the end of the season in September or early Oc-
tober. It is most numerous in the last half of the summer.

ASTEROCAMPA CLYTON (Boisduval and LeConte)
Plate 4, a, b

Range.—Northern and western part of the State southward to
Northampton, Lancaster, and Henrico Counties, and in the mountains
southwestward to the southern border. Our records are from Bote-
tourt, Giles, Henrico, Lancaster, Montgomery, Northampton, Page,
Shenandoah, Warren, and Wythe Counties.

Occurrence.—Found along woodland roads and streams, in glades
in the woods, and along tree-lined roadsides, always in the vicinity of
hackberry trees ; very locally distributed and erratic in its occurrence,

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

but usually rather common when and where it is found. This species
is frequently associated with the more generally distributed and
more numerous A. celtis, though by preference rather more of a
woodland species.

Seasons—Two broods. This species appears to have the same
seasons as A. celtis, but it is not in evidence until about the middle of
June. The second brood is on the wing toward the end of July and
flies until about the middle of September.

Family NYMPHALIDAE

Subfamily NyMPHALINAE
Genus POLYGONIA Hiibner
POLYGONIA INTERROGATIONIS (Fabricius)
Plate 16, f

Range.—Throughout the State.

Occurrence-—Found in open deciduous woods, in brushy regions,
about willows in damp meadows and along roadsides, and sometimes
in damp open fields; generally distributed, everywhere frequent and
locally common.

Seasons.—Three broods. This species, popularly called the ques-
tion mark, emerges from hibernation in the last half of March and
flies through May and early June. In the latter part of May, some-
times as early as the end of the third week, a fresh brood appears, and
the insect is on the wing continuously until the end of the season in
October or November. In July a second brood appears, and toward
the end of August a third brood, the individuals of which live through
the winter.

Note.—The dark summer form (wmbrosa) occasionally hibernates,
and the light winter form (fabricti) is sometimes found in summer.

POLYGONIA COMMA (Harris)
Plate 5, e, f

Range.—Throughout the State. We have records from Albemarle,
Alleghany, Amelia, Arlington, Augusta, Bedford, Clarke, Fairfax,
Fauquier, Giles, Grayson, Hanover, Henrico, Madison, Montgomery,
Nansemond, New Kent, Northampton, Page, Prince Edward, Prince
George, Prince William, Princess Anne, Pulaski, Roanoke, Shenan-
doah, Smyth, and Warren Counties.

Occurrence.—Found in open deciduous woods, especially in glades
and along the roads; local and usually not very common ; most gener-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 4I

ally distributed and most numerous in the mountains, especially at
the higher altitudes, becoming very local on the outer Coastal Plain.

Seasons.—Three broods. The comma, sometimes called the hop
merchant, appears in the first warm days of spring, in the lower
areas usually in the last half of March, in the mountains usually in
April. Indeed, it is occasionally noticed on warm days throughout
the winter flying along sheltered woodland roads. These early indi-
viduals, which have overwintered, fly until about the middle of May,
sometimes until the end of the month. In the last week in May the
first brood appears, flying through June and until about the middle
of July. After the third week in July individuals of the second brood
appear. In late August, September, and October the third brood is
on the wing, flying until the end of the season in October. It is
possible that in the northern part of the mountain region the third
brood is only partial. This species usually passes the winter in the
adult stage. A perfectly fresh individual taken by John Boyd at
Woodberry Forest on April 23, and others noticed by us in Augusta
County on May 12, suggest that the winter may sometimes be passed
in the pupa.

POLYGONIA PROGNE (Cramer)
Plate 5, c,d

Range.—Confined to the Transition Zone in the western part of
the State, ranging eastward to Warren, Page, Madison, Bedford, and
Grayson Counties, casually farther eastward. Our records are from
Albemarle, Bath, Bedford, Bland, Giles, Grayson, Highland, Madison,
Montgomery, Page, Rockbridge, Rockingham, Shenandoah, Smyth,
Tazewell, Warren, and Washington Counties.

Occurrence.—Found in and near open deciduous woods, especially
along woodland roads and in brushy areas; generally distributed
throughout its range, frequent and locally often rather common.

Season.—Two broods. This species, the gray comma, appears
early in May, rarely as early as the middle of April, and flies through
May and into June. The first brood is on the wing by the end of the
first week in July, and the butterfly becomes common in the last half
of the month, flying until about the middle of August. The second
brood appears toward the middle of August and flies until the end of
the season in September, when the individuals go into hibernation.

POLYGONIA FAUNUS (W. H. Edwards)

The subspecies found in Virginia is—

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

POLYGONIA FAUNUS SMYTHI A. H. Clark
Frontispiece, figs. 3, 6; plate 5, a, b

Range.——Confined to the Canadian Zone in the higher mountains in
the western part of the State in Highland, Augusta, Giles, Mont-
gomery, Washington, Smyth, and Grayson Counties.

Occurrence.—Found chiefly in the spruce or “lashhorn” forests,
especially in clearings, along forest roads, and along the borders of
the forests; in some places occurring in deciduous woods that have
replaced the original spruce forest; locally distributed, but common
wherever it is found, least numerous in deciduous woods.

Season.—One brood. Smyth’s anglewing first appears at about the
end of the first week in July, becomes common in the last half of the
month, and continues to emerge until at least the middle of August,
the individuals going into hibernation at the end of the season. Our
only spring record is Augusta County, May 12, 1947 (Dr. G. C.
Pitts).

Genus NYMPHALIS Kluk
NYMPHALIS ANTIOPA (Linné)
Plate 2, b

Range.—Throughout the State. Our records are from Albemarle,
Amelia, Augusta, Bedford, Buckingham, Chesterfield, Clarke, Cul-
peper, Fairfax, Fauquier, Frederick, Giles, Grayson, Hanover, Hen-
rico, Highland, Isle of Wight, Madison, Mecklenburg, Montgomery,
Nansemond, New Kent, Northampton, Page, Prince Edward, Prince
William, Princess Anne, Roanoke, Rockingham, Shenandoah, Smyth,
Warren, Washington, and Westmoreland Counties.

Occurrence-—Found in open deciduous woods and groves, and
about willows, poplars, and elms in open country; generally dis-
tributed, more numerous in the higher Piedmont and in the mountains
than elsewhere, though nowhere very common.

Seasons.—Two or three broods. The mourning cloak, the hardiest
of all the butterflies occurring in Virginia and the one active at the
lowest temperatures, may be seen flying about in sheltered places in
the woods at any time during the winter if there happens to be a suc-
cession of warm sunny days. It is always the first butterfly to appear
in spring, flying in some numbers on the first warm days, usually in
the first half, or shortly after the middle, of March and becoming
common in April. In May the numbers decrease, but a new brood
appears at the end of the month or early in June, and a second brood
appears in August, flying into September. Fresh individuals appear-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 43

ing in late September and October presumably represent a partial
third brood. We have no definite proof, but we believe that on the
Coastal Plain there are three full or nearly full broods, in the moun-
tains two broods and a partial third.

Genus VANESSA Fabricius
VANESSA ATALANTA (Linné)
Plate 8, f

Range.—Throughout the State.

Variation.—Throughout its wide range this butterfly is singularly
uniform, without definite local races. In Virginia it varies between
two extremes. At one extreme the individuals are dark golden-brown
above with the orange band across the fore wings broad; the size is
rather small, the fore wings being 27-30 mm. long. At the other
extreme the individuals are larger, with the fore wings up to 35 mm.
long, and black with the band across the fore wings more reddish and
narrow, often interrupted, and commonly including a small white
circular spot ; the under side is very dark.

In Virginia most individuals are intermediate between these two
extremes—very dark brown above with the band of the fore wings of
moderate width. They agree with typical atalanta of Europe. In the
spring many of the individuals tend to approach the small light ex-
treme, and large dark individuals do not occur.

In the summer the individuals average somewhat larger and darker
than in spring, and in boggy areas on the Piedmont and on the Coastal
Plain there occur large and very dark individuals which are less active
than those usually seen and which range northward locally along the
coast to Long Island. Southward they become progressively more
numerous and more generally distributed, and finally predominate.
In Virginia this is a “wet” form, confined to boggy areas, although
farther south it represents the species almost or quite to the exclusion
of the other forms.

Occurrence-—Found in open fields, especially boggy fields with
buttonbush, and in open deciduous woods; generally distributed and
usually rather common, occasionally locally or generally abundant. It
is sometimes noticed in numbers in patches of pine woods along the
seacoast, where it seeks shelter from the wind.

Seasons.—Three broods. The red admiral first appears at the end
of March or early in April, both worn and fresh individuals flying
together. The first fresh individuals to appear are small males which
are followed in about a week by larger and darker males and females.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The first brood reaches its maximum in the last half of May and al-
most or quite disappears in the first week in June. About the middle
of June individuals of the second brood appear, these reaching a maxi-
mum in the last half of July, after which the numbers fall off. After
about the middle of August the third brood is on the wing, reaching a
maximum in the first half of September and flying into October or
even as late as the last week of November.

The red admiral hibernates both as an adult and in the pupal stage.
It would appear that in the coastal regions hibernation is mainly in
the pupa, in the colder and drier regions chiefly in the adult stage.

VANESSA CARDUI (Linné)
Plate 4, d, e

Range.—Throughout the State.

Occurrence.—Found in open country everywhere, and when abun-
dant also in open woods; very erratic in occurrence, usually infre-
quent to frequent, occasionally everywhere abundant, and in some
years apparently entirely absent; more regularly present and more
numerous on the Coastal Plain and in the southern half of the western
mountains than elsewhere.

Seasons.—Three broods. Our records are insufficient to enable us
to make any definite statement regarding the broods of this species in
Virginia further than to say that these appear to agree with the broods
of the two related species Vanessa atalanta and V. virginiensis. No
individual of the overwintering brood has ever been found in Virginia,
although we have found a few on the District of Columbia side of the
Potomac. The painted lady therefore is almost or quite killed out each
winter, the summer individuals being the young either of a few that
have overwintered in protected situations and escaped observation or
of wandering females that have entered the State from outside. The
second brood appears at the very end of May and slowly increases in
numbers, reaching a maximum in the last half of July. In the last half
of August brilliant fresh individuals indicate the appearance of the
third brood, which is on the wing until the end of the season, usually
in early, sometimes in late, October.

VANESSA VIRGINIENSIS (Drury)
Plates 4, f, g; 30, 7, k (aberration)
Range.—Throughout the State.

Occurrence.—Found in open country everywhere, generally com-
mon, most numerous on the Coastal Plain. This species, in contrast

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 45

to Vanessa atalanta and V. cardui, varies but little in numbers from
year to year.

Seasons —Three broods. This species, the American painted lady,
appears on the Coastal Plain toward the end of March or early in
April, in the interior somewhat later, both worn and fresh individuals
flying together. About the first of June, later in the higher country,
the first individuals of the second brood are on the wing and the butter-
fly increases in numbers, reaching a maximum in the last half of July
and the first half of August. About the middle of August while the in-
dividuals of the second brood are still numerous the first individuals of
the third brood appear, and the insect flies until the end of the season,
usually early in October. Fresh individuals sometimes noted in No-
vember and even as late as the middle of December are probably pre-
mature emergences of the next spring brood.

This species hibernates both as an adult and in the pupa, near the
coast chiefly in the pupa, elsewhere largely or mainly as an adult.

Genus JUNONIA Hiibner
JUNONIA EVARETE (Cramer)

The subspecies occurring in Virginia is—

JUNONIA EVARETE COENIA (Hiibner)
Plates 4, h, 4,9; 5,9; 4

Range.—Throughout the State.

Variation—tIn Virginia this butterfly has two seasonal forms, a
spring-and-autumn form and a summer form, and in addition a very
distinctive form confined to localized wet areas.

In the spring-and-autumn form (pl. 4, 1), which may also be found
in dry areas during the summer, the fore wings measure about 27 mm.
long in the females and about 24 mm. in the males. The ground color
above is brown. On the under side the ground color of the hind wings
and of the apex of the fore wings is light gray, usually slightly tinged
with buff, and there are two conspicuous oval black spots narrowly
ringed with buff about as broad as, or narrower than, an interspace,
not far from the outer border of the hind wings. In this form the
wings are dry and brittle, and nearly all the individuals captured are
nicked or more extensively damaged.

The summer form (pl. 4. 7, 7) is very similar but averages slightly
larger and somewhat darker. It is characterized especially by having
on the under side of the hind wings a more or less developed irregular
pinkish or reddish band with indefinite edges crossing the wings from

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

the outer third of the anterior border to near the anal angle, passing
just in front of the two black spots. This is by far the commonest and
most widely spread form in summer.

In the form confined to localized wet areas (pl. 5, g, 1) the fore
wings are about 30 mm. long in the females and about 28 mm. long
in the males. On the upper surface the ground color is dark and the
brown of the fore wings and of the outer half of the hind wings some-
times shows dark green metallic reflections. On the under side the
apical portion of the fore wings and the whole of the hind wings are
dull pinkish red, often quite uniform, but usually darkest in a long
irregular narrow triangle bordered by irregular crenulate lines running
from a base near the outer angle of the hind wings to an apex near the
anal angle. This triangle often includes two small oval blue spots nar-
rowly ringed with lighter, or some trace of such spots. Occasionally
the under side of the hind wings is uniform bronze instead of red.
This form has curiously soft wings and always feels as if it had re-
cently emerged from the pupa. Though the wings may be rubbed, they
are rarely broken or torn. With this form there are always to be found
more or fewer of the usual summer form that drift into its restricted
areas from the surrounding country, and in some places all intergrades
between this and the summer form may be found.

Occurrence.—Found in open country, especially in dry and more or
less barren areas and along roadsides, but also in wet fields, when
abundant penetrating into open woods, especially on the Coastal Plain.

Seasons—Three broods. The buckeye appears at the end of May
and is on the wing continuously in increasing numbers until the end of
the season, toward the middle of November or later. The second brood
appears after the middle of July when many individuals of the first
brood are still on the wing and flies until at least the end of August.
The third brood appears late in August and flies until the end of the
season. The butterflies of this brood may be seen on warm days
throughout the first half of the winter, but they have never been
found in spring in Virginia.

Our records indicate that this butterfly is merely a summer visitor
to Virginia, dying out completely in the winter, for we have no record
of any overwintered individual. Our earliest records are all for the
last days of May and represent the earlier individuals of the first
brood. These records are all from the outer Coastal Plain—Princess
Anne, Norfolk, Nansemond, and Accomack Counties. During June
the butterfly becomes distributed all over the Coastal Plain and the
adjacent Piedmont, and also in the Shenandoah Valley as far north-
east as Warren and Fauquier Counties. By mid-July it has spread

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 47

over the entire State, and in August it is found everywhere, most
abundantly on the Coastal Plain, though it is common or even abun-
dant on the Piedmont, and frequent even in the higher mountain
valleys. It varies more or less in abundance from year to year, oc-
casionally being scarce on the Coastal Plain and almost completely
absent elsewhere.

Although we know of no instance of this butterfly’s surviving the
winter in Virginia, we have seen ragged overwintered individuals
in March, April, and early May in Washington, D. C.

Subfamily MELITAEINAE
Genus EUPHYDRYAS Scudder
EUPHYDRYAS PHAETON (Drury)
Plates 3, 1; 30, e (aberration)

Range—From Fairfax County westward and in the mountains
southwestward to the southern border of the State. Our records are
from Alleghany, Amherst, Arlington, Bedford, Botetourt, Giles,
Madison, Montgomery, Page, Roanoke, and Rockbridge Counties.

Occurrence.—Found, always in association with balmony, or turtle-
head (Chelone glabra), in bogs in or near woods and in wet meadows
with bushes or trees, more rarely in damp ravines near mountain
streams; very local, but often abundant in the restricted areas in
which it occurs.

Season.—One brood. The balmony, or turtlehead, butterfly is on
the wing from early in June until about the end of the third week in
July; at Mountain Lake, Giles County, where for most species the
seasons are later than elsewhere in the State, it has even been taken
in August. For a large colony that formerly inhabited a meadow at
Cabin John, Md., just across the Potomac River from Fairfax
County, the dates were from May 30 to July 11, and presumably the
individuals of the colony that formerly was found at the southern
end of the Highway Bridge, on the site of the old Washington Air-
port, had the same flight period. At Forks of Cacapon in Hampshire
County, W. Va., a short distance west of the Frederick County line,
we have taken this species as early as May 29.

Genus MELITAEA Fabricius
MELITAEA NYCTEIS (Doubleday and Hewitson)
Plate 9, f, g

Range.—Found in the mountainous western portion of the State,
ranging eastward to Warren, Fauquier, Page, Madison, Augusta,

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Rockbridge, Bedford, Prince Edward, and Lunenburg Counties.
Our records are from Alleghany, Amherst, Augusta, Bedford, Bu-
chanan, Fairfax, Fauquier, Giles, Highland, Lunenburg, Madison,
Montgomery, Page, Prince Edward, Pulaski, Roanoke, Rockingham,
Russell, Scott, Shenandoah, Smyth, Warren, Washington, Wise, and
Wythe Counties. We have not found it in Frederick County, though
it is common in Hampshire County, W. Va., a few miles farther west.
It has recently been taken in some numbers by T. B. Blivens at Cabin
John, Md., just across the Potomac River from Fairfax County, and
by Dr. Laurence I. Hewes in Fairfax County.

Occurrence-—Found in open woods with brushy undergrowth in
hilly or mountainous country, especially along the brushy borders of
roads through woods and in brushy clearings; generally distributed
and rather common in the southwest, elsewhere local and not
numerous.

Seasons.—Two broods. This species first appears early in May and
flies until about the middle of June. It reappears in the last week in
June, becomes most numerous in the last half of July and the first
half of August, and flies until the first week in September. The early
records, which were kindly given us by Dr. Warren H. Wagner, Jr.,
and others, are from Hampshire County, W. Va., not far from the
Frederick County line, where this species is common, and from Cabin
John, Md.

MELITAEA ISMERIA Boisduval and LeConte
Frontispiece, fig. 9

This species was recorded from Virginia by Boisduval and LeConte
in 1833 and by Strecker in 1878, possibly quoting Boisduval and
LeConte. It was again recorded from Virginia by Hall in 1930 (Bull.
Hill Mus., vol. 2, Supplement, p. 34), but this may refer to West
Virginia.

Genus PHYCIODES Hiibner
PHYCIODES THAROS (Drury)
Plate 11, 7, k, 1, m

Range.—Throughout the State.

Occurrence.—Found everywhere in open country, in scrubby land,
and in the more open woods—in fact, everywhere but in the thickest
woods ; common throughout, though most numerous in weedy fields
with asters and along weedy roadsides. This species and Everes
comyntas are the most generally distributed and the most numerous
butterflies in Virginia.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 49

Seasons.—Three broods. The pearl crescent usually first appears
shortly after the middle of April (rarely early in April or even late
in March), becoming common by the end of the month and flying
continuously until the end of the season, usually the first or second
week in October, though sometimes as late as almost the middle of
November. It is least common in the first half of June, in the interval
between the first two broods, but from the middle of June onward
it is abundant and fresh individuals are always obtainable, though
they become relatively scarce in the first half of August. The second
brood appears shortly before the middle of June (along the coast as
early as the end of the first week). The third brood appears in the
latter half of August, when worn individuals of the second brood
still are numerous, and flies until the end of the season.

PHYCIODES BATESII (Reakirt)
Plate 11, f, g, h, 4

Range.—Definitely known only from Bedford, Frederick, and
Giles Counties. In Bedford County a pair was taken by Dr. Carroll
E. Wood, Jr., in Sunset Field, Apple Orchard Mountain, on June 1,
1938 ; these are now in the United States National Museum. In Giles
County one was taken on Beanfield Mountain, Pembroke Road, July
I, 1940, and another at Mountain Lake on the same day, both by
Lloyd G. Carr. Bates’s pearl crescent was described in 1865 from
specimens from Winchester, Va., and Gloucester, N. J. Samuel H.
Scudder recorded it from Virginia without definite locality in 1872.

Occurrence——Found along the borders of woods, in glades in
woods, and on bushy hillsides. It is probably more common than
these few records would indicate, as its close resemblance to P. tharos
would cause it to be overlooked. But throughout its range it is very
local and seldom common where it occurs.

Season.—One brood. The dates are June 1 and July 1. Dr. W. T.
M. Forbes has pointed out that in any given locality this species flies
between the first two broods of P. tharos.

Family ARGYNNIDAE

Subfamily LiMENITINAE
Genus LIMENITIS Fabricius
LIMENITIS ARTHEMIS (Drury)

The subspecies found in Virginia is—

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

LIMENITIS ARTHEMIS ASTYANAX (Fabricius)
Plates! aia (OMe gf

Range.—Throughout the State.

Variation —Limenitis arthemis is divisible into three subspecies,
which when typically developed are quite distinctive. The northern
subspecies, rubrofasciata Barnes and McDunnough, most nearly ap-
proaches the Old World species and probably should be regarded as
the parent stock in America. The other two subspecies are astyanax
Fabricius in the Southeast and arizonensis W. H. Edwards in the
Southwest. Over a broad area extending from Nova Scotia south-
ward through New England and New York to Pennsylvania and
westward to Minnesota rubrofasciata passes over into the south-
eastern astyanax through a great number of intermediate forms, of
which nearly 20 may be recognized and others also occur. Many of
these forms have received names. Whether these forms represent
simply transitional types between the stable rubrofasciata and the
equally stable astyanax, or whether they are hybrids, is debatable. A
few of these forms reappear in areas so far south that there can be no
question of hybridization. White-banded individuals of astyanax in
southwestern Virginia (pls. 2, a; 6, g), for instance, are undoubtedly
of local origin. They cannot be directly related to white-banded indi-
viduals living farther north from which they are separated by a broad
area in which no white-banded individuals occur.

A parallel case occurring in Australia has been studied in detail by
Dr. G. A. Waterhouse. The satyrid Tisiphone abeona Donovan is
divisible into seven subspecies or local races. One of these, T. a.
joanna Butler, apparently confined to the watershed of the Hastings
River, is so variable that no two specimens are alike. Some speci-
mens from Port Macquarie are somewhat similar to the subspecies
morrist Waterhouse; others are similar to the quite different sub-
species aurelia Waterhouse. Twenty-two varieties of joanna are fig-
ured by Waterhouse. He found great variation in butterflies raised
from eggs all laid by the same female, the variation being equally
great in both sexes.

Twenty miles north of Port Macquarie the subspecies morrisi
occurs, and about 25 miles south the subspecies aurelia. The sub-
species joanna, therefore, is parallel to the intermediate forms of
Limenitis arthemis connecting rubrofasciata on the north with asty-
anax on the south, though the latter cover a much greater extent of
territory.

Dr. Waterhouse has successfully paired the subspecies abeona with

NOS 7 BUTTERFLIES OF VIRGINIA—CLARK 5I

the subspecies morrisi and rawnsleyi Miskin and carried the progeny
through two and three generations. This would suggest that joanna
is a natural hybrid between morrisi and aurelia, and by inference that
the intergrades between Limenitis arthemis rubrofasciata and L, a.
astyanax may be also fertile hybrids between these two forms.

It is most unfortunate that Drury’s type specimen of arthemis was
a specimen almost exactly halfway between rubrofasciata and asty-
anax. The name arthemis cannot logically be used for any of the
subspecies; though applicable to the species as a whole it is based
upon a single individual in the midst of a long series of unstable in-
tergrades. Exact duplicates of Drury’s type specimen are rare,
though approximate duplicates are not uncommon, flying with various
other types of intergrades.

A large female agreeing closely with others from the Adirondacks
of New York and with Drury’s figure of arthemis was taken at
Charlottesville, Albemarle County, by Frank W. Trainer on October
5, 1941. It is very worn and appears to have come from a considera-
ble distance, possibly from West Virginia or the mountains of Penn-
sylvania. Thanks to the generosity of Mr. Trainer this specimen is
now in the United States National Museum.

A fresh and perfect female referable to the form albofasciata
Newcomb (pl. 2, a) was taken by Col. Wirt Robinson, U. S. Army,
in Nelson County on July 23, 1919. In the extent of the white band
and the blue on the hind wings it agrees minutely with specimens
from Sharon and Deerfield, Mass., from Long Island, the type lo-
cality of albofasciata, and with Newcomb’s figure. It is, however,
brighter and more metallic than northern specimens, and the under
side is darker. Except for the broad white bands it agrees with speci-
mens of astyanax from the same region. This appears to be simply
an individual from the local astyanax stock in which, for some reason,
the broad white bands have reappeared.

A small male from Nelson County (pl. 6, g), also taken by Colonel
Robinson, is essentially a small example of astyanax with a narrow
white band on the fore wings. It is intermediate between the forms
albofasciata and proserpina W. H. Edwards. Both this and the pre-
ceding specimen are now in the United States National Museum.

Occasional individuals from Mountain Lake, Giles County, show
more or less strongly developed indications of a white band on the
under side of the fore wings, much more rarely on the upper side.
Except for this they agree completely with specimens of astyanax
from the same region. These resemble proserpina, but are probably
of local origin, unconnected with proserpina from farther north.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Similar individuals occur rarely in Augusta County, and probably
elsewhere in the mountains.

Occurrence.—Found in open deciduous woods, especially along the
roads, about groves of trees in open country, in orchards, and oc-
casionally about willows in boggy fields ; generally distributed, usually
infrequent to frequent, locally rather common. It is rather more
numerous and more generally distributed in the western mountains
than on the Piedmont or on the Coastal Plain.

Seasons —Three broods. In Virginia the black admiral first ap-
pears in May, usually about the second week, rarely as early as the
sixth, and slowly increases to a maximum at the end of May and the
first week in June. The numbers then decrease and the insect becomes
rare toward the end of June. Early in July it reappears, reaching a
maximum in the last half of the month and the first half of August.
In the last half of August it again becomes scarce, but it reappears
again at the end of the month and reaches a third maximum in the
first week in September. After the middle of September the numbers
decrease, but it is sometimes found until nearly the middle of October.

LIMENITIS ARCHIPPUS (Cramer)

Two intergrading subspecies of the viceroy occur in Virginia.

LIMENITIS ARCHIPPUS ARCHIPPUS (Cramer)
Plate 6, f

Diagnostic features—Color above light, pale dull orange, some-
times darker and more brownish on the fore wings; under side of
wings with no trace of bands in the cell.

Range.—Throughout the State ; but in the region east of the Dismal
Swamp interdigitating and intergrading with L. a. floridensis, which
finally becomes dominant near the coast south of Virginia Beach.

Variation.—This subspecies is fairly uniform throughout its wide
range, and most of the specimens from Virginia agree perfectly with
others from farther north (New York) or west (Illinois). In the
northern and western portions of the State very pale females are
sometimes found that resemble the lightest from the Middle West
(pl. 6, f), though in the same areas occasional males will be nearly
as dark as examples of floridensis. On the outer Piedmont and on the
Coastal Plain the average color becomes darker, and dark males are
more frequent. In the Dismal Swamp region the butterfly is more or
less intermediate between archippus and floridensis. Here, so far as
we have seen, there are no very light individuals such as occur in the

NOS 7. BUTTERFLIES OF VIRGINIA—CLARK 53

north and west, but individuals darker than the average, sometimes
much darker, are frequent. Normally colored individuals resembling
the average of those from New York or the Middle West occur east-
ward to the coast, though only as local colonies or individuals in a
population composed chiefly of the dark floridensis.

It is possible that the draining of the eastern part of the Dismal
Swamp and the Green Sea by the digging of the Dismal Swamp
canal, and the draining of the northern portion east of Suffolk, has
resulted in the intrusion of archippus into territory normally inhabited
by floridensis, for under the original conditions in this area there was
a broad belt of territory of a nature unsuited to either subspecies.

Occurrence.—Found about willows, especially along streams and
the borders of ponds in low open country and in the broader and more
open valleys, occasionally on hillsides about poplars and along road-
sides; rather local, though usually frequent to common where it
occurs. This is more of a lowland and open-country butterfly than
L. arthemis astyanax and is usually more numerous in the areas in
which it is found.

Seasons.—Three broods. The viceroy first appears about the mid-
dle of May and is on the wing continuously until the end of the sea-
son, the first or second week in October. As a result of the varied
growth stages in which the caterpillars spend the winter the emer-
gence period of the first brood is unusually long and the broods are
broadly overlapping, so that the fluctuations in numbers during the
season are relatively slight. The butterfly is most abundant between
the middle of July and the middle of August. Fresh individuals rep-
resenting the second brood appear early in July when worn individ-
uals of the first brood are still numerous. Late in August the third
brood puts in its appearance, flying until the end of the season,

LIMENITIS ARCHIPPUS FLORIDENSIS Strecker
Plate 6, d, e

Diagnostic features——Color above darker, orange-brown to chest-
nut or dark mahogany, usually darker on the fore wings; under side
of wings usually with the cell bands characteristic of L. arthemis
astyanax faintly indicated in pale blue.

Variation—The dark individuals forming the great majority of
those occurring east of the Dismal Swamp appear to be inseparable
from the subspecies floridensis, though in the region just east of the
Dismal Swamp there are some colonies composed of individuals of
average coloration. On the other hand, many are darker than some

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

specimens from Florida, and Dr. Warren H. Wagner, Jr., took a
specimen at Wallaceton on July 5, 1941, that was almost as dark as
L. a. eros. Just west and north of the Dismal Swamp very dark indi-
viduals, especially males, are rather frequent, and they occur, with
rapidly diminishing frequency, westward throughout the State,
though rarely in the western and northern portion.

In the revised edition of “The Butterfly Book” (1931), Dr. W. J.
Holland figures (pl. 73, figs. 1, 2) the type specimen of W. H. Ed-
wards’s eros under the name floridensis. In floridensis the hind wings
above are usually slightly lighter than the fore wings; the under sur-
face of the hind wings is usually considerably lighter than in eros,
and there are no white spots along the inner side of the black band.
The subspecies floridensis more nearly resembles his figure of L. a.
archippus (pl. 7, fig. 4), but it is darker, especially on the fore wings.
In southern Florida there is complete intergradation between floriden-
sis and eros, just as in Virginia there is complete intergradation be-
tween floridensis and archippus. Strecker followed his original de-
scription of floridensis with a description of what he called “ab. b.
male Nig.” This form, considered as an aberration by Strecker, was
later described as eros by W. H. Edwards.

Occurrence.—Found about willows in the marshes and along the
roadsides ; rather local, but usually common where it occurs, especially
in the sedge marshes near the coast. This form is more generally
distributed and, near the sea, usually more common than L. a. archip-
pus farther west.

Seasons.—Three broods. Our records are too few to give a very
definite picture of the broods, but there is no evidence that these differ
from the broods of L. a. archippus.

Subfamily ArGYNNINAE

Genus SPEYERIA Scudder

SPEYERIA IDALIA (Drury)
Plate 7, 1,7

Range.—Throughout northern and western Virginia south and
east to Fairfax, Prince William, Stafford, Spotsylvania, Orange, Al-
bemarle, Nelson, Rockbridge, Alleghany, Roanoke, Floyd, and Pat-
rick Counties. We have records from Albemarle, Alleghany, Au-
gusta, Buchanan, Fairfax, Fauquier, Frederick, Grayson, Greene,
Highland, Loudoun, Madison, Montgomery, Nelson, Orange, Pat-
rick, Prince William, Pulaski, Rappahannock, Roanoke, Russell,
Shenandoah, Smyth, Spotsylvania, Stafford, Tazewell, Warren,
Washington, and Wythe Counties.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 55

Variation—In the northern Piedmont region the regal fritillary
reaches a large size, the males having the fore wings 39-42 mm. long,
the fore wings of the females being 46-50 mm. In the higher moun-
tain regions the size is appreciably less, especially in the females.
Mountain males have the fore wings 37-41 mm. long, the fore wings
in mountain females being 42-46 mm. The males taken at high alti-
tudes are less richly colored than the larger lowland males, and in
both sexes the black markings on the fore wings tend to be slightly
heavier, and the dark color near the inner border of the hind wings
below more extensive.

Occurrence.—Found in pastures with boggy or marshy areas, in
damp open grasslands, in extensive grassy bogs, and at high altitudes
in dry pastures; generally distributed in the northern part of the
State as far south as Spotsylvania, Orange, and Albemarle Counties,
farther south found only in the mountains and more localized ; usu-
ally frequent to common in the areas in which it occurs.

Season.—One brood. The males of this species appear about the
middle of June, rarely as early as the end of May, and are followed
about 2 weeks later by the females. The males continue to emerge
until about the middle of July, and the females until after the middle
of August. Both sexes fly until the end of the season in September,
but after about the first of August the males are all worn and shabby,
and their numbers decrease rapidly. By the end of August males have
become rare, though females still are common.

SPEYERIA DIANA (Cramer)
Plate; a,b} c. a

Range.—From Chesterfield to James City County and southeast-
ward to the Dismal Swamp; from Highland County southwestward
in the mountains, eastward to Rockbridge, Bedford, Roanoke, Mont-
gomery, Floyd, and Patrick Counties. Our records are from Alle-
ghany, Bath, Bedford, Bland, Botetourt, Buchanan, Charles City,
Chesterfield, Dickenson, Floyd, Giles, Grayson, Highland, Isle of
Wight, James City, Montgomery, Nansemond, Northampton, Pat-
rick, Prince George, Pulaski, Roanoke, Rockbridge, Smyth, Surry,
Tazewell, Washington, Wise, and Wythe Counties.

The present range of this fritillary, covering only the outer Coastal
Plain and the mountains, is quite abnormal; furthermore, it is de-
cidedly rare except in a very few widely scattered localities. We have
only a single record for no less than 14 counties. These facts suggest
that it has been largely extirpated from a much more extensive for-

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

mer range by the clearing of the land and that its disappearance from
most of the State is only a matter of time. The records for Nor-
thampton and Chesterfield Counties are old; it has not been seen in
either county for at least 20 years.

Variation.—The males have the fore wings usually between 43 and
47 mm. in length. The largest males we have seen were from south-
ern Illinois, with the fore wings 52 mm. long. The first males to
appear are smaller than those that emerge later, and the males from
Highland County and from the higher altitudes in the mountains
farther south are always small, resembling the earliest males from
other regions.

On the Coastal Plain the males occasionally have on the under side
of the hind wings at the end of the cell a conspicuous silver spot
bordered inwardly and outwardly by black lines, corresponding to the
silver spot in the same position in S. cybele though smaller, seldom
reaching more than halfway across the cell, and the silver markings
on the outer portion of the hind wings may be enlarged. Occasionally
on the upper surface of the fore wings there are broad light dashes
in the black ground color beyond the cell in the interspaces between
veins 3 and 4, 4 and 5, 5 and 6, corresponding to the light dashes
on the under side though with indefinite borders. So far as we have
seen, these features are confined to males from the Virginia Coastal
Plain. The majority of the males from the Coastal Plain, however,
do not differ in any way from others from the mountains.

In perfectly fresh males, at least in those from the Coastal Plain,
the black on the fore wings is overlaid with a beautiful blue reflection
recalling that in certain species of Euploea, though not so intense.
This soon disappears.

Dr. Henry Skinner has pointed out that the males differ materially
in the number and size of the black spots on the upper surface of the
hind wings. We have seen males, both from the Coastal Plain and
the mountains, with the light border of the hind wings almost immac-
ulate. There is some variation in the size of the black spots on the
fore wings, and veins 2, 3, and 4 may be narrowly or rather broadly
infuscated. The contrast between the dark basal and light outer por-
tions of the under surface of the hind wings is sometimes accentu-
ated. These variations seem to bear no relation to locality.

The females have the fore wings 50 to 55 mm. long. Dr. Skinner
said that females from eastern Tennessee, western North Carolina,
and southern Illinois are larger than those found in Virginia ; but the
females from the lower altitudes in southwestern Virginia appear to
be quite as large as any from farther south. Dr. Skinner pointed out

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 57

that the females vary a great deal in the degree of silver beneath and
also in the band of large bluish or greenish spots on the hind wings
above. In some specimens these spots are large, and in others they
are confined to a small area around the black spots. The number and
size of the cream-colored or white spots on the upper surface of the
fore wings is also quite variable.

A female from Spring Grove, Surry County, captured on Septem-
ber 4, 1940, has the fore wings with blue and the hind wings with
blue-green reflections. The band on the hind wings is blue, broader
and more continuous than in any others we have seen from elsewhere,
crossed by very narrow dark lines, and separated from the submar-
ginal dashes, which have white centers, by a relatively narrow black
line. Another female taken at the same time and place is similar, but
the hind wings have blue reflections like the fore wings, and the band
is blue-green.

Prof. Ellison A. Smyth, Jr., writes that he is inclined to believe
that perfect freshly emerged females always have the under side of
the hind wings and of the apex of the fore wings dark blue-black and
that as the insect ages this speedily assumes a rusty-brown color, even
before the upper surface shows any wear or dimness, All the fresh
specimens in his collection, as well as his recollection and notes of
other captures, indicate the correctness of this conclusion. Some
slightly worn females show in direct light the brown color, but held
at a slight angle, and particularly in artificial light, the blue-black
can still be seen. He has one female, nearly fresh, in which the outer
third of the under side of the hind wings, which usually retains the
blue-black, is also rusty brown with blue-black angular dashes run-
ning to the outer margin. He has specimens that are green on the
upper surface instead of blue, and also specimens showing various
shades of blue. The blue area on the upper surface of the hind wings
varies from an almost complete band to isolated blue angular dashes.
Professor Smyth’s specimens were from Montgomery County.

Occurrence.—Found on the Coastal Plain and outer edge of the
Piedmont in and near old pine woods with running streams or ex-
tensive cold seepages. In the mountains it occurs chiefly at moderate
or high, though not the highest, altitudes in deep, damp, well-wooded
valleys or ravines, or on damp, wooded mountain sides, always more
or less near cold streams, and always where the undergrowth, es-
pecially rhododendrons or alders, is thick. It is very local, and as a
rule rare to infrequent where found. In a number of localities we
have found it but once, although we have visited them several or
many times. We have searched carefully for it in many likely-looking

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

localities in the southwestern counties without finding any trace of it.

We have found this species most numerous in Surry County, from
Surry to Cabin Point, especially about Spring Grove. We first no-
ticed it in this region on June 15, 1938, when we captured four males
along the main highway (Route 10) in about 20 minutes and saw
others. On July 5 we found two males and a female in the same
place, and Mrs. Barnes, who lives nearby, told us that sometimes
there would be as many as 25 at one time about her butterflybush.
Dr. Warren H. Wagner, Jr., who visited this locality on June 25,
1940, reported seeing about 12 males and a female in a couple of
hours. Otto Buchholtz and J. W. Obenderfer, Jr., have also taken
specimens here. It is fairly common in Poverty Hollow, Montgomery
County, where Professor Smyth obtained his specimens, in certain
valleys near Hot Springs, Bath County, and near Mountain Lake,
Giles County.

Season—One brood. The males of this species first appear shortly
before the middle of June (June 10) and soon become common. The
females appear in the third week in June (June 25) and increase
slowly in numbers until August, when they are at their height, though
all the males are more or less worn. Both sexes fly until early Sep-
tember, the males becoming very scarce toward the end of the season.

SPEYERIA CYBELE (Fabricius)
Plates! 659, 0n

Range.—Throughout the State.

Variation.—The ground color of the females may be more or less
dark yellowish or buffy. The inner portion of the wings is often
heavily infuscated. The black markings, particularly those of the fore
wings, may be enlarged and confluent, especially on the under side.
Quite commonly the second and third black bars across the cell of the
fore wings are merged into a large black spot, and the spot in the
expanded portion of the outer black bar is solid black. The spots of
ground color enclosed by the marginal chevrons on the fore wings
may be whitened from the apex downward, this whitening usually
affecting only the uppermost three or four spots, rarely more. The
color of the under side of the hind wings varies from a uniform
rather light yellow-brown to uniform cinnamon, chocolate, or purplish
brown, the last two being very rare. It is usually yellow-brown more
or less extensively washed with cinnamon or light chocolate.

In males from the lowlands of Virginia the fore wings are usually
38-40 mm. long, those of the females 47-50 mm. long, though smaller
individuals occur. In the mountains the size is somewhat less.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 59

Occurrence-—Found in open woods, especially in rolling, hilly, or
mountainous country, and in open fields and bogs more or less near
woods; generally distributed and in most places frequent to common ;
infrequent on the outer Coastal Plain and on the Eastern Shore
(Accomack and Northampton Counties).

Season.—One brood. The males of the cybele first appear about
the middle of May and soon become common. The females appear
about the first of June and slowly increase in numbers. After about
the middle of August all the males are worn and ragged, but fresh
females continue to appear throughout August and even in early
September when the males have become very scarce. We do not un-
derstand Dr. W. J. Holland’s statement that this species “seems to be
single-brooded in the north and double-brooded in Virginia, the Caro-
linas, and regions westward in the same latitude.” Although fresh
females appear throughout the summer, we know of no records of
fresh males after about the first of August.

SPEYERIA APHRODITE (Fabricius)
Plate 8, a, b

Range.—Confined to the Transition Zone in the western moun-
tains, occurring eastward to Frederick, Warren, Rappahannock, Mad-
ison, Rockingham, Albemarle, Nelson, Rockbridge, Bedford, Floyd,
and Patrick Counties; occasional at Cabin John, Md., across the
Potomac River from Fairfax County. We have records from Albe-
marle, Alleghany, Amherst, Augusta, Bath, Bedford, Bland, Bu-
chanan, Carroll, Dickenson, Fauquier, Floyd, Frederick, Giles,
Grayson, Highland, Lee, Madison, Montgomery, Nelson, Page, Pat-
rick, Pulaski, Rappahannock, Roanoke, Rockbridge, Rockingham,
Russell, Scott, Shenandoah, Smyth, Tazewell, Warren, Washington,
Wise, and Wythe Counties.

Variation—In specimens from Virginia and the Carolinas the
light spots in the outer half of the anterior portion of the under side
of the fore wings are duller and more brownish than in those from
farther north, and the pink flush at the base is usually less pro-
nounced, these southern individuals showing an approach to the
form alcestis.

In Virginia this species reaches a large size, the fore wings in the
males being about 37 mm. long and in the females 38-44 mm. The
earliest individuals and those of the high mountain “balds” or open
grassy areas are somewhat smaller than those appearing in summer
in more favorable localities.

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

The earliest individuals to appear early in summer are bright and
clear, and in both sexes the ground color of the under side of the
hind wings is uniform light yellowish brown. Very soon females are
noted in which the under side of the hind wings is more or less ex-
tensively and heavily shaded or blotched with deeper cinnamon-brown
in the interspaces between the silver spots. Later these females are
supplemented by others in which the ground color of the under side
of the hind wings is uniform deep cinnamon-brown, this color usually
more or less completely obliterating the submarginal yellow band,
which is always prominent in the lighter females. In summer and
autumn all three types of females, with all possible intergrades, occur,
those of the intermediate type being the most numerous and those of
the dark type common. The female forms of this species are parallel
to those of S. cybele, but the extreme dark form, very rare in cybele,
is common.

Like S. cybele this fritillary is subject to frequent and occasionally
extensive aberrations. The black spots on both surfaces may become
enlarged, elongating within the interspaces, and more or less exten-
sively coalesced, while the ground color is heavily obscured with dark
scales above and below. We have seen more or less extreme females
of this type from Syria, from Long Mountain on Route 60, and from
White Sulphur Springs, W. Va. In the males this infuscation takes a
somewhat different form, the individuals becoming above velvety
brownish black with a more or less heavily infuscated fulvous mar-
gin about 5 mm. wide which on the fore wings is crossed by long,
broad triangles at the veins and on the hind wings includes a par-
tially double row of black bars in the interspaces, and just within
these, more or less fused with the dark interior portion of the wings,
a row of black spots. On the under side the wings are almost entirely
blackish, the hind wings showing the usual silver spots only on the
inner third (ab. bakeri). We have not seen this aberration from Vir-
ginia, but we have seen four somewhat diverse examples from
Waynesburg, Ohio.

Among the minor variations the following five are more or less fre-
quently noted. The submarginal spots on the fore wings may be
greatly enlarged. The area between the second and third bars across
the cell of the fore wings may be entirely black, a variation more
common in S. cybele and also occurring in Euptoieta claudia. In the
females the broad dark band between veins 2 and 3 of the fore wings
may be continued inward to the cell, forming a conspicuous black
trapezoid. In the males the black dashes forming the irregular post-
median line on the fore wings may all be extended outward to a

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 61

straight line parallel to the outer border of the wings (ab. hughi). In
the males also the submarginal chevrons may be extended inward,
becoming completely coalesced with the row of submarginal spots,
with which they form an elongated spot with a deeply concave outer
end.

Occurrence.—Found in bogs, glades, open woods, and fields, es-
pecially boggy fields, in mountainous country, and also in high moun-
tain pastures ; generally distributed and common to abundant where-
ever found, especially at the higher altitudes.

Season.—One brood. The males are on the wing in small numbers
early in June, occasionally at the end of May, the females appearing
toward the end of June, both sexes becoming common early in July.
Both sexes remain on the wing until early in October, though after
the end of August the proportion of the now worn and ragged males
rapidly diminishes. Fresh females, however, may be found until the
end of the season.

SPEYERIA ATLANTIS (W. H. Edwards)
Plate 8, c, d

Range.—Known only from Highland County. On June 19, 1948,
when in company with Mr. and Mrs. William D. Field, we found
this species frequent in a meadow on the crest of Middle Mountain
near the mission school and obtained several specimens. Flying with
it were Boloria toddi ammuralis and B. selene myrina.

Several years ago our friend the late John Boyd told us that he
had seen an individual in Bath County but failed to catch it.

Occurrence.—Found in high mountain bogs and damp pastures.

Genus BOLORIA Moore
BOLORIA TODDI (Holland)

The subspecies occurring in Virginia is—

BOLORIA TODDI AMMIRALIS (Hemming)
Plate 9, d, e

Range.——Confined to the Transition Zone in the western part of
the State, occurring eastward to the higher altitudes in Page, Madi-
son, Rockingham, Augusta, Nelson, Bath, Montgomery, Floyd, and
Patrick Counties. Our records are from Augusta, Bath, Carroll,
Floyd, Giles, Grayson, Highland, Madison, Montgomery, Nelson,
Page, Patrick, Pulaski, Rappahannock, Roanoke, Rockingham,
Smyth, Tazewell, Washington, and Wythe Counties.

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Variation Individuals of the first brood are often rather small
and light-colored with a washed-out appearance below and with the
dark markings above restricted and more or less disconnected. Sum-
mer individuals as a rule are very uniform. Occasional specimens
have the dark markings above reduced in size and disconnected, while
in others they are enlarged and confluent; but these aberrations are
as arule rare. In certain localities, however, as in a wet pasture near
Independence, Grayson County, many, or even most, individuals in
summer have the dark markings above enlarged and confluent, in
extreme cases the inner half of the wings being almost wholly black.

Occurrence.—Found in high, dry mountain pastures, in wet or
boggy pastures at somewhat lower altitudes, and in mountain bogs;
locally distributed, though common where it occurs.

Seasons.—Three broods. This fritillary first appears in the last
half of May, flying nearly through June. It reappears shortly after
the first of July, and the third brood is on the wing in the last half
of August, broadly overlapping the second brood and flying until
early September.

BOLORIA SELENE (Schiffermiiller)

One subspecies is known definitely from Virginia, and another
occurs in Maryland just across the Potomac River and should be
found in suitable localities on the Virginia side.

BOLORIA SELENE MYRINA (Cramer)
Plate 9, a

Diagnostic features Small, the fore wings 22-23 mm. long; color
above dull yellowish orange, the black markings narrow and delicate.

Range.—Known only from Highland, Bath, Nelson, and Giles
Counties.

Variation—Specimens from Virginia do not differ appreciably
from others from farther north, although the average size is slightly
larger, the fore wings being 22-23 mm. long, and the color averages
slightly more reddish than in individuals from Massachusetts. Occa-
sional summer individuals have the dark markings above enlarged and
confluent, like corresponding individuals from the north.

Occurrence—F ound in wet pastures or in extensive grassy moun-
tain bogs; very local, but frequent to common wherever it occurs;
most generally distributed in Highland County and the northwestern
part of Giles County. Wherever this subspecies occurs B. toddi am-
miralis is also found, but the latter is much more generally distributed

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 63

and lives under more diverse conditions. But B. toddi ammiralis does
not occur with the following subspecies.

Seasons.—Three broods. This little fritillary first appears in the
first week in May (records from Hampshire County, W. Va.) and
flies until the middle of June. The second brood is on the wing after
the middle of July and flies until the second half of August. Pre-
sumably there is in Virginia, as elsewhere, a third brood in the last
half of August and September, although we have no records for that
period.

BOLORIA SELENE MARILANDICA (A. H. Clark)

Frontispiece, figs. 1, 2; plate 9, b, c

Diagnostic features—Larger than B. s. myrina, the fore wings
about 25 mm. long ; color above more reddish with the black markings
broad and heavy and more or less confluent.

Range.—?Northampton County. An old record from Northamp-
ton County (Bayford) may refer to this subspecies.

This butterfly was formerly rather common in a bog at Beltsville,
Md., near the railway station, but it has not been found there since
1929. In 1941 it was rediscovered by Dr. Warren H. Wagner, Jr.,
in a bog near Largo, Md., but has now disappeared from that locality
also. No locality where it occurs is known at present.

Occurrence.—Found in open, wet, grassy bogs.

Season.—One brood. In contrast to the 3-brooded B. s. myrina,
this subspecies flies only from the very end of June until near the end

cf July.
Genus EUPTOIETA Doubleday
EUPTOIETA CLAUDIA (Cramer)
Plate 6, a, b

Range.—Throughout the State.

Variation —The earliest individuals appearing in spring in Virginia
are always very small with the fore wings 22-25 mm. long. and are
easily mistaken for a species of Boloria. Later individuals have
the fore wings 28-35 mm. long, the females being larger than the
males.

There is more or less variation in the intensity and clearness of the
dark markings, especially in the females. In some of these the ground
color of the inner half of the wings is more or less heavily infuscated
and the dark markings broadened and blurred. Rarely the light spot
near the costal margin in the basal third of the fore wings may be

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

black, or the three light spots in the three lowest interspaces just be-
yond the irregular median line on the fore wings may be black.

Occurrence——Found chiefly in dry open fields and along roads
through fields; in late summer usually generally common and locally
abundant on the outer Coastal Plain, frequent to common on the
Piedmont, and infrequent on the higher mountain pastures. This
butterfly varies considerably in abundance in different years. It was
especially numerous in 1935 and 1936, but we did not meet with it
at all in 1939, our only record for that year being a single individual
noted by Dr. Carl W. Gottschalk at Salem, Roanoke County, on
September 7.

Our records indicate that this butterfly in Virginia overwinters
locally in small numbers, at least in certain years, for we have records
of the dwarf spring form in April and May from Henrico, Albemarle,
and Warren Counties, and from Hampshire County, W. Va. For
June we have records only from Princess Anne, Accomack, Henrico,
Fauquier, Albemarle, and Roanoke Counties, most of them from the
Coastal Plain. By the end of July the butterfly has become generally
distributed, and by the end of the season it is usually one of the
common butterflies in most parts of the State, and is abundant on the
Coastal Plain wherever the common passionflower or maypop (Passi-
flora incarnata) thrives in the corn and cotton fields and along the
roadsides. Our spring records are to be regarded merely as showing
the very spotty nature of the occurrence of this species in spring, not
as giving a true picture of the localities in which it may be expected to
occur, as these localities vary more or less from year to year.

Seasons.—Three broods. The variegated fritillary appears on the
Coastal Plain early in April, or even toward the end of March, some-
what later inland, and flies into June. Toward the end of June fresh
individuals of the second brood begin to appear, and the insect be-
comes more and more common as the second brood develops. About
the middle of August the first individuals of the third brood appear,
and the butterfly soon extends to every section of the State. At the
end of September and in October fresh individuals may represent a
partial fourth brood. This species is on the wing until some time in
October or even November, occasionally into early December.

Subfamily HELICcONIINAE
Genus AGRAULIS Boisduval and LeConte
AGRAULIS VANILLAE (Linné)

The subspecies occurring in Virginia is—

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 65

AGRAULIS VANILLAE NIGRIOR Michener
Plate 7, e, f

Range.—Known to us only from Princess Anne and Norfolk
Counties on the outer Coastal Plain.

Occurrence-—Found in open country generally, especially in waste
land and along roadsides where maypops (Passiflora incarnata) are
abundant; an irregular summer visitor from farther south, usually
rare or wholly absent, occasionally locally frequent. The senior au-
thor found it rather common about Norfolk in 1929, and Dr. George
W. Rawson tells us that he has seen it common there.

Season.—Our only records are Norfolk, August 3, 1929, and
Princess Anne County, August 19, 1936 (Warren H. Wagner, Jr.).

Family DANAIDAE

Subfamily DANAINAE
Genus DANAUS Kluk
DANAUS PLEXIPPUS (Linné)

Two subspecies occur in Virginia.

DANAUS PLEXIPPUS PLEXIPPUS (Linné)

Plate 10, a

Diagnostic features——Preapical spots on fore wings light dull
orange; apical portion of fore wings, beyond the preapical spots,
more or less extensively dull orange; two complete rows of small
white spots in the black border of the hind wings.

Range.—Throughout the State.

Variation—We have noticed no appreciable variation in this but-
terfly in Virginia. An individual with the ground color mouse gray
instead of dull orange (ab. fumosus) was captured at Farmville in
October 1941 by Miss Marie Bricker, who very kindly presented it
to the United States National Museum.

Occurrence-—Generally distributed in open country, preferring
damp or wet pastures with an abundance of flowers, unkept and
weedy fields, particularly along streams and about ponds, and weedy
roadsides ; it is also quite at home in the extensive cattail marshes
along the southeastern coast and occasional in the more open pine
woods of the Coastal Plain, though avoiding thick woods and woods
with undergrowth. This butterfly is most abundant in the higher
areas in the north and west, especially in the mountains, less common
on the Piedmont, and least numerous on the Coastal Plain, though

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

usually common in the great marshes along the southeastern coast.
Its numbers show considerable fluctuation from year to year.

Migratory flights in Virginia—While driving from Washington
to Alexandria along the main boulevard on September 22, 1934, we
saw between 200 and 300 monarchs in the adjoining fields or crossing
the road, flying singly or more rarely in twos or threes, occasionally
in larger groups, from 6 to 15 or 20 feet above the ground. All were
flying directly west. The light wind, as determined from weather
vanes in Alexandria, was east or slightly north of east. Continuing
from Alexandria to Accotink, the same phenomenon was noticed ; be-
tween 200 and 300 were seen, all flying west. On September 25, 1934,
on the road from Accotink to Alexandria, between 5 and 5 :30 o’clock
in the afternoon, a few butterflies were noticed, flying west as before.

Dr. Carroll M. Williams writes us that in the autumn of 1936 he
noticed a southerly movement of this species at Richmond. “The
steady, yet completely scattered, migration in a general southerly
direction was quite apparent. No hint of mass migration was visible,
however.”

At Bear Trap Farm near Mount Solon, Augusta County, in Sep-
tember 1944, we noticed that all the individuals seen were flying
southwest, parallel to Narrowback Mountain. They were usually
4 to 6 feet above ground, but one was at a height of about 50 feet.
Only once were two in sight at the same time, a male and a female
feeding on the same group of flowers a few inches apart. This was
evidently a scattered migration like that described by Dr. Williams.

Dr. Williams quotes from a letter from Mrs. Walton, of Clifton
Forge, Alleghany County, as follows: “During the first of October
1935, a great mass of monarch butterflies passed through the valley
coming from the northeast and following the valley river (the James)
and railroad. They were about 30 yards in width and 6 feet in depth.
They were flying just high enough to miss the house tops and so on
down until you could touch them. They were flying at a moderate
speed, and some of them would pause a second to rest; and when
night came they would all rest just where darkness caught them.
They were so thick in places that they seemed like a cloud passing.
Some of them would dart in and out between the houses. In their
flight they followed the mountain valley.”

This unusual flight was the subject of much discussion locally at
the time.

Dr. Frank Morton Jones, of Wilmington, Del., writes us that at
Virginia Beach, Princess Anne County, from April 18 to 30, 1906,
throughout most days from one to three or four of these butterflies

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 67

were usually in sight flying toward the north or northeast, some ob-
viously seeking the larval food plant. There were many dead ones
along the shore, washed up by the waves.

This note is particularly interesting and significant in view of the
fact that later in the season, in summer and autumn, this is not a
common insect about Virginia Beach. It also suggests an explanation
of the occurrence of the tropical form (megalippe) in southeastern
Virginia. Individuals of this form, which so far as is known is nor-
mally nonmigratory, may join the flocks of plexippus in the far south
where both occur and be, so to speak, swept north with it.

On April 28, 1931, Prof. and Mrs. Torsten Gislén of Lund, Swe-
den, reported a single individual flying westward along the Virginia
shore of the Potomac River rather high in the air, and on May 9,
1931, when on the Maryland side of Great Falls, in company with
Miss Dorothea M. A. Bate of the British Museum and Drs. Doris
M. Cochran and Herbert Friedmann of the United States National
Museum, the senior author saw one flying slowly up the river before
the wind about 20 feet above the cliffs, just as it flies in the same
region in autumn.

Seasons.—Four to six broods. The monarch first appears shortly
after the middle of April, on the Piedmont and in the west late in
April or early in May, the dull and worn individuals seen at this time
having come from hibernation. So far as is known, all the individuals
on the wing at this time are females. Toward the end of May fresh
individuals of both sexes appear, and the butterfly is continually on
the wing in increasing numbers until toward the end of October.
There are four to six broods during the summer, but fresh individ-
uals are to be found at all times as each female deposits her eggs over
an unusually long period. In the last half of the summer if the sea-
son be dry fresh individuals will be seen in numbers after a heavy
rain, a phenomenon from which this butterfly’s popular name “storm
fritillary” is derived. This species does not survive the winter in
Virginia, coming in each year from the south.

DANAUS PLEXIPPUS MEGALIPPE (Hiibner)
Plate 10, b

Diagnostic features—Preapical spots on fore wings white; apical
portion of fore wings, beyond the preapical spots, heavily infuscated ;
small white spots in the black border of the hind wings obsolete or
wholly absent in the central portion.

In the revised edition of Dr. W. J. Holland’s “The Butterfly Book”
(1931), the figure of Danais plexippus (pl. 7, fig. 1) is not plexippus

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

but megalippe. The specimen from which this figure was made came
from South America, probably from Surinam.

Range.—Our records are from Essex, Norfolk, Princess Anne,
Shenandoah, and Surry Counties, and Alexandria.

Occurrence-—An uncommon casual visitor to Virginia from tropi-
cal America, found chiefly on the outer Coastal Plain.

Seasons.—Our dates are June 15, July 4, 5, 6, 23, and Septem-
ber 2.

Note.—In addition to those taken in Virginia, we have examined
specimens of this form from St. Tammany Parish, La., near New
Orleans (Percy Viosca, Jr.), Key West (Lucien Harris, Jr.), the
northern end of Currituck Sound, N. C. (captured by us), and Long
Island, N. Y.; also, one from Decatur, Ill. Except for the one from
Decatur, Ill., and the one from Shenandoah County, Va., all these
specimens are from the vicinity of seaports (Norfolk, New Orleans,
Key West, and New York) at which steamers are constantly ar-
riving from tropical American ports. We are inclined to believe that
these individuals were brought to this country on these steamers and
that, though a rather frequent visitor, this form is not properly an
element of our fauna.

This form, as well as typical D. p. plexippus, has been captured
in England. Of Danaus plexippus 157 individuals have been reported
as seen in England, of which 62 were captured. Dr. E. B. Ford
writes that of these only 22 have been determined as to subspecies,
and 1 of these, caught in Cornwall in 1885, is said to be of the
Central American type. On August 30, 1941, he was so fortunate
as to capture a fine example at Kynance Cove, Cornwall, of which
he published a colored figure. His figure also appears to represent
the Central American form and is most closely matched by one from
Costa Rica in the National Museum collection.

It has been suggested that our Virginia specimens are in reality
aberrations of D. p. plexippus; but if that were the case they should
be found in the west, where the species is much commoner than it
is on the Coastal Plain, and not almost exclusively in the general
vicinity of Norfolk.

Family LIBYTHEIDAE
Genus LIBYTHEANA Michener
LIBYTHEANA BACHMANII (Kirtland)
Plate 23, 1

Range.—Throughout the State. Our records are from Accomack,
Albemarle, Amelia, Arlington, Augusta, Botetourt, Culpeper, Fair-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 69

fax, Fauquier, Frederick, Giles, Goochland, Henrico, Isle of Wight,
Lee, Madison, Montgomery, Nansemond, Northampton, Page,
Prince Edward, Prince William, Princess Anne, Rappahannock,
Roanoke, Rockbridge, Rockingham, Shenandoah, Surry, Sussex, War-
ren, and Westmoreland Counties.

Occurrence.—Found in open fields near woods, in brushy fields,
along tree-lined roadsides, and about hackberry trees; very local
and very erratic in its occurrence, though usually frequent to abun-
dant when and where found.

Seasons.—Three or possibly four broods. The beaked butterfly
first appears, coming out of hibernation, in the first half of April and
flies into early May. The first brood is on the wing in the last half of
June, the second in the last half of July and especially in early
August, and the third in late August and September. It is probable
that some of the September butterflies represent a fourth brood.
According to W. H. Edwards the period from the laying of the eggs
to the emergence of the butterfly is from 15 to 17 days.

Family RIODINIDAE
Genus CALEPHELIS Grote and Robinson
CALEPHELIS VIRGINIENSIS (Guérin)

Plate 12, 7

Range-——Known only from Virginia Beach and Lake Tecomseh
in Princess Anne County, and from Brinkley and the western border
of the Dismal Swamp in Nansemond County.

Occurrence-—Found along the grassy and weedy borders of pine
woods; very local, but common where it occurs. It has a curious
way of making itself conspicuous by resting in the center of the
mass of white flowers of the common yarrow or milfoil (Achillea
millefolium), with the wings extended horizontally.

Seasons.—Three broods. The first brood flies late in April and
in May, the second in July, and the third from early in September
to early in October.

CALEPHELIS BOREALIS (Grote and Robinson)

Plate 12, i

Range.—Known only from Hot Springs in Bath County, Gala and
Tinker Mountain in Botetourt County, Blacksburg, Palmers Hill,
and Poverty Hollow in Montgomery County, and Fort Lewis Moun-
tain in Roanoke County.

70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Occurrence.—Found in open woods, about the borders of woods,
and in clearings in damp valleys in limestone regions; very local, but
common where it occurs.

We have never been so fortunate as to meet with this butterfly,
the northern metalmark, ourselves. Regarding its occurrence in
Montgomery County Herman J. Erb wrote us: “It was the early
part of July. I went alone on top of Palmer’s Hill, about 2,500
feet above sea level, and found a gully running toward the Roanoke
River. There is a small brook in the center of the gully almost dry
in the summer. . . . Farther down I came to a little open spot in
the woods and saw a few small butterflies. They were easy to catch,
but new to me. I showed them to Professor Smyth, and he told
me they were C. borealis. He told me that he was 12 years in Blacks-
burg, and all he ever got was one. Since then I got my C. borealis
from the same locality every time I went to Virginia. They are on
the wing from about the 25th of June until the latter part of July.
Three years ago last summer (i1.e., in 1933) .. . I discovered a new
locality for C. borealis. I crossed Brush Mountain about Io miles
west of Blacksburg and over 3,000 feet above sea level, and went up
Poverty Hollow. It was tough going, but I was amply paid. I struck
a paradise all along the path. In the little open spots I took about
forty C. borealis. All together I took about 100 C. borealis in 1933.”

Although we have never found this species in Frederick County,
Dr. Warren H. Wagner, Jr., found it common in nearby West Vir-
ginia, at Ice Mountain in Hampshire County and at Caudy Castle
in Morgan County.

Season.—One brood. This species is on the wing from the middle
of June to the first week in August. It is most numerous during
the first 3 weeks of July.

Family LYCAENIDAE
Subfamily SPALGINAE
Genus FENISECA Grote
FENISECA TARQUINIUS (Fabricius)
Plate 15, e, k

Range.—Northern and western portions of the State eastward
to Fairfax, Henrico, Prince Edward, and Halifax Counties. Our
records are from Albemarle, Amherst, Augusta, Fairfax, Giles, Gray-
son, Halifax, Henrico, Montgomery, Prince Edward, Roanoke, Rock-
bridge, and Smyth Counties. It is probably more generally distrib-
uted than these records would indicate.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 71

Variation.—Although this species varies between unusually wide
extremes, the fore wings being in normal individuals 11-20 mm.
long, our series from Virginia is very uniform in size, the fore wings
being usually 15-17 mm. long, and also in color. We have found that
bred individuals of this species are so very variable as to give quite
a false idea of this butterfly as it exists in nature in the same region.

Seasons.—One to eight broods, usually four or five. The alder
butterfly appears shortly after, or sometimes before, the middle of
April and is on the wing continually until early October. The cater-
pillar develops very rapidly, molting only three times, but the pupal
stage is of very variable length, usually from 8 to 11 days, though
sometimes as long as 11 months in pupae from the same lot. The
butterflies seen at the end of the season are probably for the most
part of the fourth or fifth brood, but a few may represent only a
second brood and others a sixth, seventh, or even eighth brood.

Subfamily LycaENINAE
Genus LYCAENA Fabricius
LYCAENA PHLAEAS (Linné)

The subspecies occurring in Virginia is—

LYCAENA PHLAEAS AMERICANA (Harris)
Plate 15, 1, m, n

Range.—Throughout the State.

Variation —Early-spring individuals are lighter and less reddish
on the upper surface of the fore wings than individuals of the sum-
mer brood, and the spots of the outer row, together with the inner
cell spot, are smaller, sometimes much smaller. The dark border
is also narrower. On the under side the hind wings are darker and
the submarginal scarlet line is often more or less obscured. The hind
wings are rounded, or with a broadly obtuse angle at the end of vein
2;in the summer form the angle at the end of vein 2 is more promi-
nent and may even be produced into a short tooth.

This species is subject to considerable variation. Occasionally the
red above and below is replaced by straw yellow (ab. fulliolus). A
fine example of this aberration was taken by the senior author at
Rocky Run, Fairfax County, on May 2, 1937.

Occurrence——Found in dry fields and waste lands wherever the
sheep sorrel (Rumex acetosella) grows; common in the northern
portion of the State and southwestward on the higher elevations

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

in the mountains, and also among the sand dunes south of Virginia
Beach ; elsewhere local and seldom common.

Seasons.—Three broods. The little copper usually first appears
shortly after the middle of April, rarely in the first half of the month,
and flies through May and into early June. The second brood appears
in the first half of June and flies until just after the middle of
August. The third brood is on the wing in the last week in August
and flies until the end of the season early in October.

Note.—The large copper, Lycaena thoé (pl. 15, f, g, h), has been
taken in the District of Columbia and in adjacent Maryland and
probably will be found in Virginia.

Genus GLAUCOPSYCHE Scudder
GLAUCOPSYCHE LYGDAMUS (Doubleday)
Plate 15, a, b

The subspecies represented in Virginia is—

GLAUCOPSYCHE LYGDAMUS NITTANYENSIS F. H. Chermock
Frontispiece, figs. 4, 5; plates 14, g; 15, c

Range.—Confined to the mountains in the western part of the
State. Our records are from Augusta, Bath, Botetourt, Buchanan,
Frederick, Highland, Montgomery, Page, Roanoke, Rockbridge, and
Warren Counties. Our few records from Frederick County were
possibly strays from Hampshire County, W. Va., where this butter-
fly is common. The records from Highland and Augusta Counties
are from the west and east sides of Shenandoah Mountain, the
boundary line between these two counties. Most of our records are
from Roanoke and Montgomery Counties. This form is probably
of local occurrence throughout the mountain area.

Occurrence.—Found in hilly or mountainous country in damp,
wooded valleys and in rich woods always in association with the food
plant, the Carolina vetch (Vicia carolimana), rarely straying into open
fields or visiting gardens; very local, but frequent to common where-
ever it occurs.

Season.—One brood. Our dates are from March 30 to May 19;
it is most numerous in the last half of April.

Note.—In October 1948 (Proc. Ent. Soc. Washington, vol. 50,
No. 7, pp. 176-178) the senior author described this subspecies under
the name of Glaucopsyche lygdamus bovydi, listing all the localities

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 73

known to him in Pennsylvania, West Virginia, Virginia, North Caro-
lina, and Arkansas, with the dates of capture. The type locality was
given as Ice Mountain, Hampshire County, W. Va. Cyril F. dos
Passos called his attention to F. H. Chermock’s G. 1. nittanyensis, the
description of which, from State College, Pa., had appeared a short
time before. Dr. Ralph L. Chermock was so very kind as to send
him a series of five specimens of nitianyensis, including two paratypes.
These left no doubt that boydi is a synonym of nittanyensis. William
D. Field agrees with this conclusion.

Genus CYANIRIS Dalman

CYANIRIS ARGIOLUS (Linné)

The subspecies occurring in Virginia is—

CYANIRIS ARGIOLUS PSEUDARGIOLUS (Boisduval and LeConte)
Plate 14, a, b, c, d, e, f, h

Range.—Throughout the State.

Variation.—The early-spring form of this species in Virginia is
pseudargiolus (=violacea). This form is variable, especially in the
mountains. The upper surface may be either clear blue or somewhat
tinged with violet. The under surface varies from grayish white to
brownish gray, and the markings vary from light to rather heavy.
Occasionally individuals are taken (for instance in Frederick County,
April 24, 1938) in which the marginal spots and accompanying chev-
rons are more or less completely fused into a broad brown marginal
border. Such examples are properly referable to marginata.

We have taken the form Jucia only in western Frederick County
in Virginia, but we have a specimen from Cabin John, Md., just
across the Potomac River from Fairfax County, taken on April 19,
1926. Strecker’s record of lucia from Virginia may refer either to
Virginia or to West Virginia.

The brown male (nigra) we have found only in western Frederick
County, where it is not common. We took one brown female (inter-
media) in Frederick County west of Cross Junction on the West
Virginia line on April 24, 1938. Strecker described this form from
Virginia, which may have referred to West Virginia.

Toward the end of the emergence period of the first brood, from
about May 1 until nearly the middle of June according to the locality
and year—usually in the last half of May—the form neglecta makes
its appearance. In these early individuals of the form neglecta the
males are somewhat paler than those occurring later in the summer
and have a slight violet tinge ; the hind wings are indistinctly marked,

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

with the light areas usually reduced and heavily dusted with blue
scales. The females are dull and the light areas on both wings, which
are not sharply defined, are rather heavily dusted with blue or light
brownish scales. This form, in which the fore wings are usually
14-15 mm. long, resemble summer individuals from the vicinity of
Boston, Mass. It appears to occur all over the State, but in the
mountains from Page and Highland Counties southwestward it is
largely replaced by a larger form (meglecta-major) with the fore
wings 15-18 mm. long in which the colors, especially in the females,
are clearer and better defined above, and the under surface is whitish
with the spots and other markings greatly reduced. We have not
seen this form except in the mountains.

In contrast to the individuals of the spring brood, those of the
summer broods are everywhere similar and show little variation.
They represent typical neglecta. The fore wings are 12-15 (usually
about 14) mm. long. The males are clear blue, with the hind wings
white crossed by blue veins and somewhat dusted with blue scales,
and with a blue border; occasionally there is a large vague and ill-
defined triangular patch of whitish heavily dusted with blue scales on
the lower portion of the fore wings. The females are white with the
fore wings metallic blue in the basal portion, with the costal and
outer borders broadly brown and a brown streak at the end of cell;
the hind wings have the anterior border and a row of small marginal
spots brown.

In the autumn occasional individuals of the spring form pseudargi-
olus are to be found (Nansemond escarpment, September 2, 1935),
in association with much more numerous examples of neglecta.

Dr. Carl W. Gottschalk has a specimen of the summer form
neglecta in which the left side is female and the right side is male.
It was captured at Salem, Roanoke County.

Occurrence.—Found in rich woods, especially along the roads and
along their borders, in moist brushy fields, brushy bogs, and swamps ;
everywhere frequent, and in some localities, particularly in the moun-
tains, common or even abundant, especially in spring.

Seasons.—One complete brood, followed by two incomplete broods.
The common blue appears in the last half of March or early in April
and soon becomes common, flying through May and into June. At
about the middle of June, or somewhat earlier, a new brood appears
which is on the wing until about the middle of August. Toward the
end of August a third brood appears which flies until the end of the
season, usually about the end of the first week in September.

This butterfly is most abundant in late April and May when large

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 75

numbers are frequently seen about dogwood, wild-cherry, and other
flowering trees in favorable localities. At this season it is confined
to woods and swamps, where it is rather local, except for stray indi-
viduals. The individuals of the second brood are more generally
distributed than are those of the first, occurring in brushy fields as
well as in the woods, but they are never so numerous in any one
locality. The second brood is not complete, many of the pupae formed
by caterpillars originating from eggs laid by females of the first brood
lasting over until the following spring, especially in the drier areas.
The third brood is composed of only a small number of individuals.

Genus EVERES Hiibner
EVERES COMYNTAS (Godart)
Plate 14, 4,7

Range.—Throughout the State.

Variation—tThis butterfly varies considerably in size, the fore
wings of apparently normal individuals ranging all the way from
5 to 14 mm. There are two fairly well-marked size groups, one with
the fore wings less than 10 mm. long, the other with the fore wings
10-14 mm., long. The average length of the fore wings in both sexes
is about 12.2 mm. Large individuals are most numerous in the moun-
tains. Dwarfs of both sexes with the fore wings 5-6 mm. long are
especially common in spring. The average size is greater in the moun-
tains than in the lower areas, as in the case of the common blue
(Cyaniris argiolus pseudargiolus). It is also greater farther north, in
the vicinity of Boston, Mass., than it is in Virginia.

William D. Field has noted that in Kansas the females in the
summer broods are brown or blackish brown above with two mar-
ginal black spots capped inwardly with orange in the lower portion
of the hind wings between veins 2 and 3 and 3 and 4. The under
side is grayish white, with a marginal series of dark points and a
submarginal series of dark bars along the outer margin of the wings.
The marginal points in the two interspaces beyond the tail are black
with a suffused border of metallic green and silver, and with orange
crescents on the inner side; sometimes there is a little orange above
the marginal spot in the next interspace. There is a submarginal
series of black spots crossing both wings, a bent bar at the end of the
cell in both wings, and three black spots across the base of the
hind wings, all these spots being ringed with white.

The males, according to Mr. Field, are similar to the females on
the under surface but usually have only two orange crescents on

79 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the hind wings. On the upper surface they are violet-blue with a
dark-brown border usually about 1 mm. broad. There is a very faint
series of dark points in the marginal border of the hind wings, that
between veins 2 and 3 being larger and darker than the others and
often bordered or capped interiorly with orange; sometimes the
point in the interspace between veins 3 and 4 is also enlarged and
bordered interiorly with orange. Mr. Field points out that on the
under side the males usually differ slightly from the females by hav-
ing a little less of the marginal orange on the hind wings, there being
usually two of these orange spots in the males and three in the females.

According to Mr. Field the spring brood differs quite consistently
from the summer broods in a number of features. The females are
slightly darker above, often almost black, and are usually more or less
extensively suffused with blue, this suffusion varying from a few blue
scales at the base of the wings to an almost complete suffusion over
both pairs of wings, except for the apex of the fore wings. Most
commonly in Virginia the fore wings are heavily dusted with blue
scales in a rounded triangle extending from the base to the outer
end of the cell and thence to the lower angle, and the hind wings are
thickly dusted with blue scales in their inner half. There is a com-
plete series of black dots along the outer margin of the hind wings,
those in the two interspaces beyond the tail being capped with small
orange crescents, the others encircled with blue, and there is a sub-
marginal series of brownish-black bars. On the under side the ground
color is lighter than in the summer form with the maculation much
less distinct, especially in regard to the marginal and submarginal
markings. There are two instead of three orange crescents in the
posterior region of the hind wings.

The males above are violet-blue with a very narrow brownish mar-
ginal border, which is much less than 1 mm. in width. On the hind
wings the marginal black spots are more distinct than in the summer
form, those in the two interspaces beyond the tail being capped with
small but distinct orange crescents. On the under side they are paler
than the males of the summer form, with less distinct markings.

In Virginia the spring and summer broods differ as described by
Mr. Field for Kansas. Summer individuals at hand from New
England (the vicinity of Boston) resemble most closely spring indi-
viduals from Virginia, though they are to a certain extent interme-
diate. The under side is slightly duller and less uniform than in
summer individuals from Virginia, with the markings more distinct.
The New England males are bluer than the Virginia summer males

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 77

with a narrower dark border, and the females show no blue dusting
on the upper surface.

Occasional individuals from all parts of Virginia entirely lack
the tails on the hind wings, there being merely a small sharp point
at the end of vein 2, and rarely this point is so reduced as scarcely
to be noticeable. Individuals without tails occur in both sexes, but
appear to be more frequent in the females.

The black spots of the submarginal row on the under side of the
hind wings are sometimes enlarged and produced inward in a long
sharp point; rarely they reach the spots of the inner row, forming
long black dashes in the interspaces.

Occurrence—Common to abundant everywhere along weedy road-
sides, in open fields, more especially the drier fields, and in open
woods ; the most uniformly distributed, locally as well as throughout
the summer, and the most numerous butterfly in the State.

Seasons.—Four broods. The tailed blue appears about the first
of April, somewhat earlier on the outer Coastal Plain and somewhat
later in the mountains, soon becomes common, and is on the wing
continuously until the end of the season in the first or second week
in October. The first brood flies from early April until toward the
end of May, the second begins to appear about the end of the third
week in May, the third appears early in July, and the fourth flies
from about the third week in August until the end of the season.
It is possible that at the end of the season some of the fresh indi-
viduals may represent a fifth brood, at least on the outer Coastal
Plain.

Subfamily THECLINAE
Genus ATLIDES Hiibner
ATLIDES HALESUS (Cramer)

Frontispiece, fig. 10; plate 12, c, d

Range.—Regularly present only in the southeastern part of the
State, in Princess Anne, Norfolk, and Nansemond Counties; casual
or accidental in Montgomery County.

Occurrence.—Found wherever the mistletoe (Phoradendron fla-
vescens) grows abundantly; regularly present, though not very
common, in the Dismal Swamp and about its borders; more or less
casual elsewhere. This butterfly as a rule keeps high in the trees
about the mistletoe and is therefore easily overlooked. It is most
frequently seen on white flowers in fields more or less near the swamp.

Seasons.—Three broods. The mistletoe hairstreak first appears at

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the end of March or early in April; the second brood is on the wing
in July; and the third brood flies from the latter part of August until
the end of the season late in October.

Genus EUPSYCHE Scudder
EUPSYCHE M-ALBUM (Boisduval and LeConte)
Plate 12, e, f

Range.—Found in the southern part of the State north to Acco-
mack, Fairfax, Madison, Albemarle, and Augusta Counties. Our
records are from Accomack, Albemarle, Augusta, Botetourt, Fair-
fax, Floyd, Madison, Montgomery, Nansemond, Prince Edward,
Prince George, and Roanoke Counties. Our Accomack County rec-
ord is based upon one specimen we captured on Tangier Island.

Occurrence.—Found in moist fields near woods and along the edges
of woods; apparently a permanent resident of the State as a whole,
but of irregular occurrence in any one locality; usually local and
uncommon, though from time to time locally frequent.

Seasons.—Three broods. The azure hairstreak appears in the
latter half of April and flies until early June. The second brood is
on the wing the last week in June and flies through July and into
August. The third brood appears toward the end of August and
flies until the end of the season early in October.

Genus STRYMON Hiibner
STRYMON CECROPS (Fabricius)
Frontispiece, fig. 7; plate 12, h

Range.—Southern portion of the State north to northern Acco-
mack, Westmoreland, Fairfax, Albemarle, Augusta, and Highland
Counties. Before 1946 it was not known in the eastern part of the
State north of Westmoreland and Caroline Counties. In 1946 it was
taken just south of Alexandria, and in 1947 it was fairly common in
northern Fairfax County and appeared in the District of Columbia.

V ariation.—An interesting form of this butterfly in which the red
on the under side is replaced by light yellow (ab. gottschalki) has
been taken several times in Roanoke and Rockbridge Counties.

Occurrence.—Found chiefly in damp open woods with abundant
undergrowth, and in the mountains along the borders of woods and
in brushy pastures; rather local, but frequent to abundant wher-
ever it occurs. This butterfly varies more or less in numbers locally
from year to year, from time to time becoming very common in
restricted areas.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 79

Seasons.—Three broods. The red-banded hairstreak usually appears
in the latter half of April, on the Coastal Plain sometimes as early
as the first of the month, and flies through May into early June. The
second brood is on the wing shortly after the middle of July, and
the insect continues to fly until the end of the season. The third
brood appears toward the end of August.

STRYMON TITUS (Fabricius)

Two subspecies are found in Virginia.

STRYMON TITUS TITUS (Fabricius)
Plate 13, g

Diagnostic features—Black spots on under side of hind wings
ringed with pale brownish, the rings being inconspicuous against the
ground color.

Range.—Known only from Highland County—Strait Creek, July
25, 1939; Monterey, July 26, 1939; Buckeye, July 27, 1939.

Occurrence-—Found in open fields near woods and in extensive
clearings in woods; rare.

Season.—One brood. The three specimens known from Virginia
are all worn. The butterfly probably flies from early July to early
August.

STRYMON TITUS MOPSUS (Hiibner)
Plate 13, h

Diagnostic features—Black spots on under side of hind wings
conspicuously ringed with clear pure white.

Range.—Apparently confined to the Piedmont and the Shenan-
doah valley. Our records are from Arlington, Fairfax, Fauquier,
Henrico, Montgomery, Prince Edward, and Warren Counties. In
North Carolina it has been reported only from Raleigh and Tryon.

Occurrence.—Found along the brushy borders of woods and in
open fields often at a considerable distance from woods or brush;
very local, and in most places infrequent. Except at Farmville,
where this butterfly appears to be fairly common, all the records repre-
sent only one or two individuals.

Season.—One brood. This butterfly is on the wing from about
the middle of June to about the middle of July. It is most numerous
in the last week in June and the first week in July.

STRYMON ONTARIO (W. H. Edwards)

The subspecies found in Virginia is—

80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

STRYMON ONTARIO ONTARIO (W. H. Edwards)
Plate 13, f

Range.—Known only from Botetourt County, Gala, June 29, 1942
(Warren P. Stoutamire) ; Fairfaix County, Difficult Run, June 29,
1920 (Ernest Shoemaker) ; and Nansemond County, Dismal Swamp,
May 25, 1945 (Otto Buchholtz). We have examined the specimen
captured by Mr. Stoutamire and one of the two taken by Ernest
Shoemaker, which is in the American Museum of Natural History in
New York.

Occurrence——Apparently an infrequent casual.

Season.—One brood. Virginia records are May 25 and June 29.
A specimen from White Oaks, Md., was taken by Dr. Warren H.
Wagner, Jr., on June 17, 1942. In Ohio a specimen was taken in
Washington Township, Jackson County, by Edward S. Thomas on
June II, 1933, and another was taken by Joe Enke at Columbus on
June 18, 1933.

STRYMON EDWARDSII (Saunders)
Plate 13, d

Range.—Probably throughout the State. The records are: Rich-
mond County, Warsaw, June 27, 1937; Prince Edward County, Farm-
ville, June 15, 29, July 20, 1941 (Frank W. Trainer) ; Augusta
County, Sherando, July 5, 1937; and Roanoke County, McAfee Knob,
July 6, 1946 (Carroll E. Wood, Jr.). There is a single record for
North Carolina, Tryon, in July.

Occurrence.—In and near open deciduous woods with oaks; rare.

Season.—One brood. The dates in Virginia run from the middle
of June to the end of July.

STRYMON FALACER (Godart)
Plate 13, c

_ Range—Probably throughout the State. Our records are from
Augusta, Botetourt, Giles, Montgomery, Prince Edward, Rappahan-
nock, Roanoke, Shenandoah, Smyth, Surry, and Warren Counties.

Occurrence.—Found in and near deciduous woods with oaks; not
common. The senior author once found it in some numbers in Rap-
pahannock County just east of Panorama.

Season.—One brood. Our dates in Virginia are from June 7
(Gala, Botetourt County) to August 3 (Hungry Mother, Smyth
County).

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 81

STRYMON LIPAROPS (Boisduval and LeConte)
Plate 13, e

Range.—Probably throughout the State. The records are: Mont-
gomery County, Palmer Hill, June 15, 1896, and Blacksburg, June 24,
1902 (Ellison A. Smyth, Jr.) ; Roanoke County, Fort Lewis Moun-
tain, July 3, 1937 (Carroll E. Wood, Jr.), July 15, 18, 1949 (Carl
W. Gottschalk) ; Tazewell County, Burke’s Garden, August 8, 1940;
Prince Edward County, Farmville, June 30, 1940 (Frank W.
Trainer) ; Nansemond County, Dismal Swamp, June 5, 1944, May
14, 1945 (Otto Buchholtz). There is a single record for North Caro-
lina, at Southern Pines (John Boyd). All the specimens are the
form strigosa.

Occurrence.—Found in and near open deciduous woods with oaks ;
rare.

Season.—One brood. The dates of capture in Virginia run from
May 14 (Dismal Swamp) to August 8 (Burke’s Garden).

STRYMON MELINUS (Hiibner)

Two subspecies occur in Virginia.

STRYMON MELINUS MELINUS (Hiibner)

Plate 11, ”

Diagnostic features—Under side of wings pale gray; anterior
orange spot on under side of hind wings large, extending anteriorly
to vein 4 or 5 and inward so as to interrupt the postmedian line, and
usually fused with the posterior spot.

Range.—FEastern Princess Anne County, south of Virginia Beach.
There is no sharp line of demarcation between this subspecies and
the following. Specimens from Princess Anne County south of Vir-
ginia Beach agree with average S. m. melinus from Florida. Farther
north and west, in northern and western Princess Anne, Norfolk,
Nansemond, Isle of Wight, and Surry Counties, the individuals are
mostly intermediate between melinus and humuli, though many are
more or less typical humuli.

Occurrence.—Found in weedy open fields with more or less abun-
dant Lespedeza.

Seasons.—Apparently four broods, agreeing with those of the
following subspecies.

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

STRYMON MELINUS HUMULI (Harris)

Plate 11, 0

Diagnostic features—Under side of wings darker ; anterior orange
spot on under side of hind wings smaller, scarcely or not at all ex-
tending beyond vein 3, not reaching the postmedian line, and usually
separated from the posterior spot.

Range.—Throughout the State, except in eastern Princess Anne
County south of Virginia Beach where it is replaced by the preceding
subspecies, with which it intergrades.

Occurrence.—Found in open country, especially in weedy unkept
fields and along roadsides with Lespedeza; generally distributed and
everywhere uniformly frequent, though seldom very numerous; most
common on the Coastal Plain, least common in the mountains. We
have found it most numerous on Tangier Island, playing about the
border of the groundsel (Baccharis halmifolia) scrub near the
southern end of the island. On Tangier Island it is one of the
commoner butterflies, occurring in the fields and salt marshes wher-
ever there are flowers. This is the most widely distributed and the
commonest hairstreak in Virginia.

Seasons—Four broods. The gray hairstreak first appears early
in April, or even late in March, later in the higher regions, and from
then on is found in increasing numbers until the end of the season
in late September or early October. There appear to be four broods,
the first appearing late in March or early in April, the second at the
end of May, the third shortly before the middle of July, and the
fourth in the last half of August.

Genus INCISALIA Scudder
INCISALIA NIPHON (Hiibner)

The subspecies found in Virginia is—

INCISALIA NIPHON NIPHON (Hiibner)
Plate 13, D

Range.—Throughout the State. Our records are from Albemarle,
Arlington, Fairfax, Frederick, Henrico, Prince Edward, Prince Wil-
liam, Princess Anne, Roanoke, and Westmoreland Counties.

Occurrence-—Found in and near open pine woods, patches of scrub
pine, and deciduous woods with an abundance of pine; very local and
not common; most numerous in Fairfax and western Frederick
Counties.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 83

Season.—One brood. This species appears in the second week in
April and flies until the end of June. It is most numerous in the last
half of April and the first half of May.

Note.—There is a possibility that the northern Jncisalia niphon
clarki T. N. Freeman (pl. 30, 4, 7) may occur in the Canadian Zone
in the higher mountain regions. We have seen no specimens from
these areas.

INCISALIA AUGUSTINUS (Westwood)

The subspecies found in Virginia is—

INCISALIA AUGUSTINUS CROESIOIDES Scudder
Plate 16, e

Range.—From Fairfax County westward to Frederick and Page
Counties and southwestward in the mountains to Bedford and Mont-
gomery Counties. Our records are from Albemarle, Augusta, Bed-
ford, Botetourt, Clarke, Fairfax, Frederick, Madison, Montgomery,
Page, Rappahannock, and Roanoke Counties.

Variation—Females from Roanoke County frequently show a line
of rusty or brick red across the end of the cell of the fore wings,
the red sometimes being extended for a greater or less distance
along the veins. A specimen from Roanoke County, April 15, 1938,
another unusually large one with the fore wings 15 mm. long from
Castleman’s Ferry, Clarke County, April 20, 1941, and two from
Rocky Run, Fairfax County, have the fringes entirely white.

Occurrence.—Found about blueberry, huckleberry, or laurel (Kal-
mia latifolia) bushes in open woods and along the borders of woods
in hilly or mountainous country ; locally distributed, but common to
abundant wherever found.

Season.—One brood. This butterfly first appears in the third
week in March, or as late as early April, and flies until the second
week in May, or in the higher altitudes until about the last of the
month. It is most numerous in the last half of April.

INCISALIA POLIOS Cook and Watson
Plate 13, a

Range.—The only Virginia specimen was taken at Orphanage
Falls, at the foot of Fort Lewis Mountain, Roanoke County, on
April 5, 1938, by Carroll E. Wood, Jr. It is a fresh specimen and
quite typical, resembling others from New Jersey. Mr. Wood pre-

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

sented it to the United States National Museum where Frank Watson
examined it and confirmed the identification.

Occurrence.—This individual was found in association with large
numbers of J. augustinus.

Season.—One brood. In New Jersey, and presumably also in
Virginia, this butterfly appears with J. augustinus, and from a week
to 10 days earlier than either J. henrici or I. irus. In Virginia it
probably flies from the latter part of March to early May.

INCISALIA IRUS (Godart)
Plate 16, c

Range.—From Fairfax County to Frederick County, and south-
westward in the mountains to Roanoke County. Our records are
from Arlington, Fairfax, Frederick, Madison, Page, Rappahannock,
and Roanoke Counties.

Occurrence.—Found about the borders of woods and along roads
through open woods; very local and not common.

Season.—One brood. This species first appears in the last week in
March and flies until about the middle of May, and in the higher
altitudes as late as the first week in June.

INCISALIA HENRICI (Grote and Robinson)
Plate 16, d

Range.—From Westmoreland and Fairfax to Frederick Counties
and southwestward in the mountains to Montgomery and Giles Coun-
ties. Our records are from Albemarle, Clarke, Fairfax, Frederick,
Giles, Montgomery, Orange, Roanoke, and Westmoreland Counties.

Occurrence.—Found in open woods, along the borders of woods,
and in brushy areas, usually in association with the redbud (Cercis
canadensis) ; very local, but often common where it occurs. Except
for J. augustinus this is the commonest species of Jncisalia in
Virginia.

Season.—One brood. This species appears in the last week in
March and flies until the second week in May; it is most numerous in
the last half of April.

Genus MITOURA Scudder
MITOURA GRYNEUS (Hiibner)
Plate 12, g

Range.—Probably throughout the State. We have records from
Accomack, Albemarle, Amelia, Arlington, Augusta, Fairfax, Giles,

NOD 7 BUTTERFLIES OF VIRGINIA—CLARK 85

Henrico, Madison, Montgomery, Nansemond, Nelson, Northampton,
Orange, Prince Edward, Prince George, Prince William, Roanoke,
Rockbridge, Stafford, Surry, Warren, and Westmoreland Counties.

Occurrence-—Found in the vicinity of red cedars (Juniperus vir-
ginianus), never straying very far from them; very local, and erra-
tic in its appearance, likely to appear suddenly in a locality where
it was previously unknown, persist for one or a few seasons, then
as suddenly disappear ; usually common when and where it is found.

Seasons.——Two broods, but the second brood is incomplete, as
more or fewer of the pupae formed by the young of the first brood
do not give forth the adults until the following spring. The green
hairstreak appears usually about the middle of April, rarely as early
as the first week in April, and flies until the middle of June. The
second brood appears early in July, or even toward the end of June,
and flies until about the middle of August. This butterfly is most
numerous in the last half of April and the first half of May and
common, though less numerous, in the last half of July.

Genus ERORA Scudder
ERORA LAETA (W. H. Edwards)
Frontispiece, fig. 8; plate 12, a, b

Range.—The only Virginia specimen was captured at the Moun-
tain Lake Biological Station, Giles County, on June 23, 1938, by
Prof. Lorus J. Milne. It is a fresh specimen, very recently emerged.
Professor Milne very kindly presented it to the United States Na-
tional Museum.

Occurrence —Mr. Scudder wrote: “As to the haunts of this insect,
all (unless the New Jersey specimen, and perhaps the London, be
exceptions) seem to have been taken in mountainous regions. Mr.
Saunders took his specimen in a wood; Mr. Edwards one of his at
the bottom of a freshly dug post hole near a hop vine. Mine was
taken on a road into a mountain ravine, just before it entered the
woods from partially cleared ground.”

Seasons.—Probably three broods in Virginia, one in the last half
of April and May, another in the last half of June and July, and a
third in August and September.

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Family PIERIDAE

Subfamily PrerINAE
Genus ANTHOCHARIS Boisduval, Rambur, and Graslin
ANTHOCHARIS GENUTIA (Fabricius)
Plate 9, 4, 7

Range.—Throughout the State.

Variation.—The earliest individuals to appear in spring are small,
both sexes having the fore wings about 16 mm. long. The wings
are somewhat narrowed, and the outer border of the fore wings makes
an obtuse angle with the lower border so that the pointed apex is
rather prominent. Gradually the individuals become larger, the fore
wings finally reaching 20 mm. or even more in length, and the wings
become more ample, the hind wings somewhat more broadly rounded
and the fore wings longer in the basal half so that the outer border
makes approximately a right angle with the lower border and the
pointed apex becomes less conspicuous. In size and in wing shape
the earliest individuals agree with others from farther north, and the
latest agree essentially with the subspecies flavida from the coast of
Georgia; but we have seen no specimens from Virginia in which the
orange patch is extended inward as in flavida. Occasional females
have the outer portion of the fore wings lemon yellow, the yellow
covering the area that is orange in the males. More rarely in the
females this area is heavily infuscated.

Occurrence.—Found chiefly in low, open deciduous woods with
large rough-barked trees near streams or swamps; in the mountains
it prefers the damp ravines and valley bottoms, though occurring
sparingly in the higher levels; locally distributed and usually infre-
quent, though in some places common, regularly or in certain years.

Season.—One brood; locally in the mountains from Page County
southwestward an incomplete second brood immediately follows
the first, and occasional individuals representing this second brood
appear along the Potomac in Fairfax County.

On the outer Coastal Plain the orangetip appears soon after the
middle of March, on the Piedmont about the end of March, and in
the mountains shortly after the middle of April. The time of its
first appearance is, however, subject to considerable variation in
different years. In any one locality it is on the wing for from 4 to 5
weeks at the most, sometimes for only 2 or 3. For about the first
week only males are found, and in the last week practically all the
individuals seen are females. The length of the flight period varies

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 87

considerably from year to year. It disappears on the outer Coastal
Plain early in April, but in the mountains and in the northern part
of the State it flies until the second week in May or even later, by
which time, according to Dr. Ellison A. Smyth, Jr., well-grown
larvae can be found on the food plant. In the mountains in Page and
Montgomery Counties and probably elsewhere, and rarely in northern
Fairfax County, a small partial second brood appears shortly after
the end of the third week in May and flies until about the second
week in June. At Blacksburg, Montgomery County, Dr. Smyth
found the second brood on the wing as early as May 26; he noted
that this brood flies for only about a week and is not so numerous as
the first brood.

Dr. W. J. Holland (“The Butterfly Book,” revised edition, 1931,
p. 287) wrote: “The first brood appears in early spring. It is double-
brooded in the western portions of North Carolina, where I have
taken it in the spring, and quite abundantly late in the autumn.” We
do not understand this statement. In his revision of the list of but-
terflies of North Carolina (in C. S. Brimley, “The Insects of North
Carolina,” 1938, p. 257) the senior author was able to include definite
records only from Raleigh, Chapel Hill, and Roanoke Rapids, all in
the north-central part of the State, in March and April.

Genus EUCHLOE Hiibner
EUCHLOE OLYMPIA (W. H. Edwards)
Plate 16, a, b

Range—Known only from northern and western Frederick
County.

Variation—In Virginia and in adjacent West Virginia the early
individuals of this species are always typical olympia, but as the
season progresses the amount of gray on the apex of the fore wings
decreases and the body becomes less shaggy. If during the period
of emergence the weather becomes hot and dry, this tendency is
carried to an extreme and the form rosa appears, quite like specimens
from Texas except for the black spot at the end of the cell of the
fore wings, which is slightly smaller. The form rosa flies with worn
individuals of typical olympia and a more or less complete series of
intergrades. In Virginia and West Virginia it is possible to capture
on the same day typical olympia and typical rosa, though most indi-
viduals will be intermediate between the two forms.

In fresh individuals of this species the markings on the under side
of the hind wings are a beautiful apple green, and the anterior por-

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

tion of the hind wings below is strongly suffused with a delicate pink
that gradually pales toward the outer edge; the costal border of
the fore wings may be pink, above as well as below. The extent and
intensity of this pink color are variable, but it appears to be as well
developed in olympia as in rosa. After death the pink gradually fades
away and the delicate apple green changes to a rather disagreeable
dingy yellow-green, so that preserved specimens give but little idea
of the exquisite coloring of this “apple-blossom” butterfly in life.

Occurrence-—Found in open woods, especially in the higher por-
tions, and along the rocky exposed crests of high ridges; common
along the crest of the ridge forming the western border of Frederick
County, infrequent or casual farther eastward. It is common to
abundant in the higher regions of adjacent Hampshire County, W. Va.

Season.—One brood. Our records indicate that this species first
appears about the middle of April and flies until about the middle
of May.

Genus PIERIS Schrank
PIERIS RAPAE (Linné)
Plate 18, a

Range.—Throughout the State.

Variation—tThe earliest individuals seen in spring are rather
small, with the fore wings about 23 mm. long. The outer border of
the fore wings is more or less strongly convex. At the apex of the
fore wings there is a small, often more or less indefinite, light-gray
patch which in the female does not quite reach the outer edge of the
costal border. The under side of the hind wings is more or less heay-
ily speckled with dark-gray scales, especially in the inner half, most
densely in a large but indefinite patch in the inner half of the wing
just below a more or less conspicuous light stripe running from the
base of the wing to just above the middle of the outer border. The
males are often unmarked above and below, but usually there is a
single dark spot, more distinct and darker below, on the fore wings
just beyond the end of the cell and about halfway between the costal
and the lower border (pl. 18, a), and a similar spot on the anterior
border of the hind wings about two-thirds of the distance from the
base to the outer angle. In the females the spot on the fore wings is
always present and below it in the interspace between veins 1 and 2,
just above the spot on the anterior border of the hind wings, there
is a more or less developed broad dash of dusky scales. In the males
the wing bases, and in the females the basal third of the wings, are

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 89

more or less heavily infuscated. The early spring form in its fullest
development with the males wholly immaculate and the females with
only a small subcentral spot on the fore wings does not occur every
year, and we have never found it on the Coastal Plain.

As the season advances the individuals increase somewhat in size
and pass into the late spring or intermediate form, which in turn
passes into the summer form. Although it is easily possible to get a
complete series of intergrades between the earliest spring form and
the summer form, the early-spring form, the late-spring or inter-
mediate form, and the summer form are to a certain extent distinc-
tive. The first two grade imperceptibly into each other, but inter-
grades between the late-spring and summer forms are rather less
numerous.

In summer individuals the fore wings are usually 25-27 mm. long,
the outer border of the fore wings is more nearly straight, the dark
markings are extended and intensified, and the infuscation of the
under side of the hind wings more or less completely disappears. Sum-
mer males have the dark apical patch on the fore wings considerably
larger than spring males, and also darker, especially the inner por-
tion, where the scales may be quite black. There is commonly a
slender tooth of black running inward along vein 5. The spot on
the fore wing is larger and darker, especially on the inner side, and
there is a gray spot below it near the inner border. The spot on the
anterior border of the hind wings is large and dark. The inner por-
tion of the wings is much less heavily infuscated, and the lower surface
of the hind wings is less extensively stippled with dark scales—indeed
in some examples these may be very few. Occasional summer indi-
viduals occur in which the dark apical patch is reduced, the spot on
the fore wings is small, and the faint spot below it and that on the
anterior border of the hind wings are merely vestigial ; but these have
the other features characteristic of the summer form.

In the summer females the dark apical area of the fore wings is
more extensive than it is in spring individuals, and much darker.
It either extends so as to include the end of vein 5, reaching the
outer margin halfway between veins 5 and 4, in which case vein 3
may be dusky at the tip, or it terminates in the interspace between
veins 5 and 6, in which case a narrow dark triangle extends inward
for some distance along vein 4; this triangle may be quite isolated, or
it may be united basally with the patch above. The two spots on the
fore wings and the spot on the hind wings are much larger and
darker than in the spring form. There is commonly a narrow dusky
line extending inward from beneath the middle of the lower spot

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 116

along the inner border, and rarely the two spots on the fore wings
are connected by a crescent of dark scales thickest in the middle with
the convexity outward. The females are frequently more or less
yellowish, rarely a rather bright pale yellow. k

Occurrence.—Everywhere common in low open country and abun-
dant about farms where cabbages are grown, becoming less numerous,
though universally present, in the mountains; though everywhere
present at all seasons, its numbers are to a considerable extent pro-
portionate to the available supply of its favorite food plant, cabbage.

Introduction and establishment in Virginia—B. W. Jones, writing
to Samuel H. Scudder from Spottsville, said that “it was a general
complaint (in Surry County) as early as 1870-71 among farmers
that they could raise no good cabbage on account of it. In 1872-73 it
infested the gardens about Petersburg in untold numbers.” This
was the first notice of the species in the State. It reached Wash-
ington, D. C., in 1872. By 1874 it had covered the northeastern
half of Virginia, and in the year following it was found throughout
the State.

Seasons.—Three to eight broods, usually four to six. The Euro-
pean cabbage white appears early in the third week in March and,
if the weather be warm, soon becomes common. It continues on the
wing until about the middle of October. Over most of Virginia there
appear to be four to six broods a year, but in the cooler mountain
regions there are probably only three, and in neglected cabbage fields
under proper conditions of temperature and moisture there may be
as many as seven or even eight. In this species the life span of the
individual is subject to great variation. On food plants other than
cabbage and in cool temperatures the caterpillars develop rather
slowly, but when feeding on cabbage the generations may succeed
each other with extraordinary rapidity. The cabbage white hiber-
nates in the pupa. Some of the pupae of each brood do not give forth
the adults until the spring following, so that spring individuals may
be the young of any of the broods of the preceding summer, though
most of them are the young of the fourth, fifth, or sixth broods.
Occasional pupae remain dormant over two winters.

PIERIS VIRGINIENSIS W. H. Edwards
Plate 18, b
Range.—One record, Frederick County northwest of Cross Junc-

tion on the Bloomery Road (Route 698) about 200 feet east of the
West Virginia line, April 24, 1938.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK gi

Variation.—This species varies considerably in the intensity of
the bordering of the veins on the under side of the hind wings.
This bordering is pale in all the individuals from Virginia and ad-
jacent West Virginia that we have seen, some having the under
side of the hind wings almost immaculate white.

Occurrence.—Confined to rich deciduous woods and extremely
local, occurring in limited numbers at usually widely separated locali-
ties. The reduction in the numbers of Pieris virginiensis and its
present occurrence only in widely separated localities are undoubtedly
the result of deforestation, which has greatly reduced the areas in
which it is possible for it to maintain itself. The case of Pieris
virginiensis is similar to that of Speyeria diana.

We agree with Dr. Alexander B. Klots that the habitat of this
species is so very different from that of P. rapae that there can be
no question of any direct competition between them. Pieris rapae is
an open-country butterfly and avoids the woods. The early-spring
form in Virginia frequents the edges of woods but does not enter
them, flying off over the fields when frightened. Pieris virginiensis
keeps strictly to the woods. We have found both species on the same
groups of flowers by the roadside at different times, but this is
unusual,

The individual we captured in Virginia had evidently been carried
over the border from some nearby locality in West Virginia by the
strong west wind that was blowing at the time. Curiosity in regard
to its true home led us to investigate the matter in 1939. On May 8,
following the Bloomery Road (Virginia 698, becoming West Vir-
ginia 45) into West Virginia, we passed the Bloomery Post Office
and soon entered a valley with a wooded hillside on the right just
beyond a wooden bridge over a small stream. Here Pieris virginiensis
was not infrequent, flying in indolent fashion among the trees, and
several were captured both by us and by Mr. and Mrs. Ernest L. Bell
who accompanied us. We noted what we assumed were individuals of
this species in a number of similar localities along the road from
Forks of Cacapon through Largent to Great Cacapon. Though no-
where numerous, this species seems to be generally distributed through-
out this region. We noted it in the same area on May 12, 1941.

Season.—One brood. This species appears shortly after the middle
of April and flies until about the middle of May.

92 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

PIERIS PROTODICE Boisduval and LeConte
Plate 18, c, d, ec, f, g, h

Range—Throughout the State.

Variation.—Early in spring this butterfly appears in a small form
(vernalis) which is dull chalky white above with the hind wings on
the under side heavily marked with dark along the veins, these dark
markings below being similar in kind and in degree in the two sexes
(pl. 18, c, d). This early spring form varies in size and in the extent
and intensity of its markings in different regions, and also in the
same region in different years. The fore wings are commonly 21-23
mm. long.

The early-spring form is soon replaced by a larger intermediate
form in which the markings on the under side of the hind wings are
paler in the females and very faint in the males.

In the summer form the fore wings are usually 23-27 mm. long,
the females being larger than the males. The males have the under
side of the hind wings pure white and unmarked (pl. 18, e, f), while
in the females they are marked with dirty yellowish (pl. 18, g, h).

In autumn individuals occasionally are found that resemble the
summer individuals in size and in the markings of the upper sur-
face, except that the markings of the females are clearer and blacker
than in most summer females. On the under side of the hind wings
the veins are broadly dark, greenish or grayish yellow, as in the early-
spring form. This autumnal form is seldom common and does not
appear every year. It is interesting in approaching very closely the
western subspecies occidentalis, which occurs in the northern Rocky
Mountain region and in Alaska. In fact the only obvious difference
between occidentalis and the eastern autumnal form of typical eastern
protodice seems to be that the dark spot in the outer third of the
interspace between veins I and 2 of the fore wings is uniformly
larger in the autumnal, as in the other, forms of typical protodice.

Occurrence.—Found in open fields, early in spring chiefly near
the borders of woods, later generally distributed; very erratic in its
occurrence, usually local and rather infrequent or even rare, though
in some years common to abundant locally or, more rarely, through-
out. The early-spring form sometimes appears in considerable num-
bers in restricted areas where, later in the season, no individuals of
the summer form are found. This species is the most unpredictable
in its occurrence of all the endemic butterflies in Virginia.

Seasons.—Three broods and a partial fourth. The checkered white
appears with the first warm weather of spring, usually in the last

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 93

half of April, often early in April, occasionally in March, and flies
until about the middle of May. In the last week in May the second
brood appears and is on the wing until about the middle of July,
when the third brood appears, flying through August and into Sep-
tember. In the last half of September, a partial fourth brood appears,
augmenting the numbers of the third brood, which is still flying.

Genus ASCIA Scopoli
ASCIA PHILETA (Fabricius)

The subspecies represented in Virginia is—

ASCIA PHILETA PHILETA (Fabricius)
Plate 8, g, h, i, 7

Range.—One record, Chincoteague Island, Accomack County, July
28, 1928 (Dr. Frank Morton Jones).

Occurrence——An irregular or casual visitor from farther south.
Dr. Jones captured three individuals, one of which was a female
of the dark smoky form, on the flowers of a privet hedge. Boisduval
and LeConte recorded this species from “Virginia” but said that it
was rare.

Note.—William P. Comstock has recorded, under the name A.
monuste, a specimen of this species from Virginia. As this specimen
was in the collection of Henry Edwards it may have come from
West Virginia.

Genus COLIAS Fabricius
COLIAS PHILODICE Godart
Plate 17, a, b, e

Diagnostic features—Ground color above clear light yellow with
no trace of orange.

Range.—Throughout the State.

Variation —The individuals on the wing early in spring are small,
with the fore wings 24-25 mm. long. In the females the dark border
of the hind wings is narrow with a slightly scalloped inner edge,
which shows a slight tendency to continue inward along the veins.
It may terminate just after vein 5, or it may be continued somewhat
farther. In a white female taken near Washington in April it is
rather broad and is continued as a haze of dark scales almost to the
anal angle. The dark border on the fore wings is narrow, the portion
below vein 4 being about as wide as the cell with the included spots

94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

separated from the yellow of the wing only by a fine hazy line, or
confluent with it. In two white females this border is somewhat
broader and the apical portion is extended inward for about two-
thirds of the distance from the apex to the black spot at the end of
the cell, while the included spots are much reduced in size. Yellow
females are occasionally similar. The black spot at the end of the
cell may have a more or less conspicuous yellow center. The early-
spring males have the borders of the wings narrow and more or less
brownish. In both sexes the under side of the hind wings and
apical portion of the fore wings is more or less strongly infuscated,
tinged with olive-green, or with grayish in the white females. In these
early individuals the hind wings are broadly and evenly rounded and
the fore wings are rather short with the outer border always more or
less convex and the apex well rounded. The later-spring individuals
are somewhat larger and resemble more or less closely those of the
brood following.

In the second and following broods the males vary but little. The
fore wings are usually 24-32 (averaging 26.1) mm. long. Most of
the variation is in the shape of the hind wings and concerns the de-
velopment of a rounded angle at the end of vein 7, which may be
wholly wanting, and the relative development of the anal angle, which
occasionally is so very broad as to be almost absent. The dark borders
are broad and black. In the southwestern portion of the State very
large males with the fore wings 30-32 mm. long are occasionally
found. These have the hind wings roundedly subangulate at the
end of vein 3 as well as at the end of vein 7. Similar males occur in
the valleys of West Virginia. The senior author took one on August
3, 1927, at Silver Spring, Md., but we have not noted this variety in
northern or eastern Virginia.

The females are more variable than the males. The average size
is slightly greater, but the maximum size is less. In the summer broods
the fore wings are 24-31 (averaging 27.8) mm. long. Although
as a rule the hind wings of the females are broader and more evenly
rounded than those of the males, they occasionally have the same
form with a well-developed rounded angle at the end of vein 7 and
a well-developed rounded anal angle. The angle between the lower
and outer border of the fore wings is ordinarily obtuse, somewhat
less so than in the males, more rarely nearly a right angle. Rarely
females have the hind wings broadly and evenly rounded and the
fore wings rather strongly convex on the outer margin. The black
border on the fore wings is usually heavy, the black extending in-
ward for two-thirds of the distance from the apex of the wings to

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 95

the black cell spot. The spots within this border are very variable
in size. Occasionally they are wholly absent, the margin of the wing
both in the yellow and in the white form being solid black. Sometimes
the black border is narrow and of uniform width, not broadening
toward the apex, so that it extends less than half the distance from
the apex to the black discal spot. The dark border on the hind wings
is very variable, though usually broad and rather heavy. Its upper
portion may enclose a rather large rounded yellow (or white) spot
and sometimes it may more or less completely enclose another below
it. Usually the first of these is more or less definitely indicated.
Beneath these spots the dark scaling usually runs up the veins in a
short tooth; more rarely the inner border of the dark margin is
smooth. Occasionally the margin is reduced to a series of curved
triangles with their broad bases, which are separated from one an-
other by narrow yellow (or white) lines, situated on the margin of
the wing and their attenuated apices running inward along the veins.
In extreme cases the bases of the triangles may be very narrow and
widely separated and the black may run for a long distance inward
along the veins, especially the two uppermost, as a black line. When
the black on the fore wings is much reduced there may be simply a
light dusting of dark scales on the ends of the four anterior veins. In
some individuals, both yellow and white, the base of the fore wings
as far as a line at right angles to the lower border passing through
the origin of vein 2 is very heavily dusted with deep-olive (in the
yellow individuals) or dark-gray (in the white) scales, and similar
scales are also abundant on the abdominal half of the hind wings,
especially toward the base.

In both sexes there is much variation in the color of the spot at
the end of the cell on the upper surface of the hind wings. It is
commonly straw yellow, sometimes so pale as scarcely to be dis-
tinguished against the ground color of the wings, but it may be more
or less bright orange, or even reddish orange, nearly as bright as in
eurytheme. There is also much variation in the black spot at the end
of the cell in the fore wings. This is usually about twice as long as
broad in the males, and from one-half to one-third again as long as
broad in the females. In both sexes it often bears a prominent tooth
in the middle of the apical side. In the males it is sometimes reduced
to a narrow black line, and in the females rather frequently and in
the males occasionally it has a conspicuous yellow (or white) center.

White females are frequently flushed with sulphur yellow on the
hind wings from the abdominal border as far as the cell and vein 2,

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

where the yellow ends abruptly, and also, less strongly, in the cor-
responding areas on the fore wings. In some cases the yellow is
bright and very conspicuous. In all such individuals that we have seen
the hind wings are shaped like those of Zerene caesoma, with a re-
duced dark border, and the fore wings have the outer margin at right
angles to the lower.

In Virginia white females are relatively as common early in spring
as later in the season, appearing with the earliest yellow females of
the spring brood. In spring, however, they occur in only one type,
a white replica of the corresponding yellow females.

In addition to the common white female there is also a dark female
which is deep, almost chrome, yellow on the fore wings with a pro-
nounced dull-orange flush on the hind wings. This dark female is
not very common. We have it from Massachusetts as well as from
Virginia.

Occurrence.—Found in open country generally, especially in clover
fields; formerly abundant throughout, now common in the higher
mountain pastures and frequent in the southwest, elsewhere infrequent
or even rare, having been almost completely replaced by the western
and southern subspecies eurytheme. Until about 20 years ago this
was the most characteristic and most noticeable butterfly of fields and
pastures, especially in the Piedmont and mountain regions. Though
less common, it was still numerous in the peanut- and cotton-growing
areas in the Southeast. Apparently a somewhat hardier butterfly than
eurytheme and better adapted for life in rugged regions, it is still
predominant in the higher mountain pastures and still frequent in the
valleys among the southwestern mountains.

Seasons.—Three broods, with a partial fourth. The yellow clover
butterfly appears usually about the middle of April, sometimes as
early as the end of March, and flies almost continuously until the
end of the season in October, usually disappearing in the first half of
the month, though sometimes persisting until the end, or even into
November. It is least common, and locally absent, about the end of
May. In the first or second week in June, or somewhat earlier, the
second brood begins to put in its appearance, and in about a week the
butterfly is common. About the second week in August the third
brood is on the wing. Fresh individuals appearing in October probably
represent a partial fourth as well as a delayed third brood.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 97

COLIAS EURYTHEME Boisduval

Plates 10, e; 17, f

Diagnostic features—Ground color orange varying to yellow with
a mere trace of orange on the hinder portion or lower side of the
fore wings.

Range.—Throughout the State.

Variation.—This butterfly is exceedingly variable, much more so
than C. philodice. Before considering this subspecies as it occurs
in Virginia it is of interest to present a composite picture of it based
upon a long series taken by Dr. Warren H. Wagner, Jr., in the Dis-
trict of Columbia, which presumably could be duplicated south of the
Potomac.

The fore wings in the males vary from 18 to 32 mm. in length, in
the females from 18 to 33 mm. Dwarfs are most common early in
spring, though they occur at all seasons. Giants are found only in
the last half of the summer, in low and more or less damp meadows.

The shape of the fore wings is very varied. They may be short
and broad with the outer edge at right angles to the lower edge and
the outer edge rather strongly convex, or longer with the angle be-
tween the outer and lower borders obtuse, the outer border straight,
and the apex pointed. In early spring or winter individuals they may
be markedly elongated and narrow. The extreme type of short wing
and the extreme type of long wing occur most frequently in very
small individuals, and are rare in individuals above medium size. The
pointed wing with the straight outer border is characteristic of all
very large individuals, but also occurs typically developed among the
smallest. The lower border of the fore wings is straight in the
females, in the males either straight or bowed outward, forming a
very broadly rounded obtuse angle approximately in the center. There
is little difference between males and females in the shape of the fore
wings, though in the females they are never so pointed as in the
more extreme males, and the lower border is always straight.

The hind wings vary from evenly rounded with scarcely any trace
of an anal angle to subangulate with a sharply rounded anal angle,
almost a right angle, and another sharply rounded angle at the end
of vein 6. The wings are usually broad, the maximum width in the
females and in many males being 88 percent of the length. In the
larger males with strongly angulated wings the width is 80 percent
of the length. In long-winged early-spring or winter individuals the
maximum width is only 70 percent of the length.

The color varies from light clear citron yellow, sometimes more

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

or less greenish above, with a faint orange flush on the inner por-
tion of the under side of the fore wings (? hybrids with philodice),
to a uniform brilliant orange usually with the costal margin bright
yellow, the males with a more or less intense violet iridescence, the
females with the spots included in the dark border yellow. But the
costal margin in both sexes and the included spots in the dark border
of the females may be orange like the rest of the wings. Rarely the
males are uniformly chrome yellow or, light dull orange (? hybrids).
In the transition from the form wholly yellow above to the deep-
orange form, the orange first appears as a faint flush on the fore
wings between the lower border and vein 1. From this the orange
spreads anteriorly to the cell. Individuals are common that have the
fore wings suffused with orange in a roughly triangular patch ex-
tending from the wing base outward to a line from the end of the
cell to the lower end of the dark margin, the upper and outer sides
of this orange triangle gradually shading into the yellow of the rest
of the wing. More rarely this orange patch has sharply defined
borders, or the veins anterior to the orange patch are broadly bor-
dered with orange, the borders being broadest at the base and tapering
outwardly. Next, the orange suffusion appears on the hind wings,
but here it becomes evident uniformly over the entire wing, except
beyond vein 7 and below vein 1, these areas always remaining yellow.
As the orange spreads over the wings it usually deepens in color,
though this is not always true. Uniform pale-orange, chrome-yellow,
or dull-orange individuals occur, flying with the much more numerous
bright-orange ones.

The dark border of the wings in the males is very variable,
ranging from vestigial to very dark and broad, 7 mm. wide at the
narrowest point in a specimen with the fore wings 30 mm. long. In
a long-winged spring male with the fore wings 22 mm. long taken on
April 12, 1938, the border of the fore wings below vein 4 is repre-
sented merely by a fine diffuse dusting of dark scales forming a series
of narrow crescents with the convexity inward, one in each interspace.
Anterior to vein 4 the dark dusting in the interspaces extends farther
and farther inward so that the whole apex is dusted with dark scales ;
the infuscated area is crossed by broadly yellow veins, and there is a
marginal more or less semicircular spot at the outer end of each
interspace. On the hind wings there is a dusting of dark scales at
the ends of the interspaces between veins 4 and 5, 5 and 6, 6 and 7,
that between veins 4 and 5 being very slight, that between veins 5
and 6 the most extensive, with that between veins 6 and 7 smaller.
The margin of the wing beyond these small patches of dark scales is

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK 99

narrowly yellow. The reduction of the dark border in the males to a
rather faint and very narrow submarginal dusting of dark scales is
rare ; usually the border is continuous and dark, and the dark scales ex-
tend outward to the base of the fringe. On the fore wings the dark
border varies in width in its lower half from scarcely more than
one-third of an interspace to more than twice the width of an inter-
space. Most commonly it is approximately the width of an interspace,
often slightly more or slightly less. When the border of the fore
wings is narrow its inner edge frequently is deeply indented by long
narrow angles running inward along the veins, and the veins may
cross it as narrow yellow lines. Most commonly, however, only the
veins at the apex are yellow, and these do not quite reach the outer
edge. When the dark border is broad the inner edge may be more
or less deeply scalloped, the black of the border extending outward
as long narrow angles along the veins. In most cases the inner edge
of the dark border is simply irregular. In the males the dark border
of the hind wings is developed proportionately to that of the fore
wings. It may reach downward to vein 1, but usually ends at about
vein 2. When the border is narrow it may not pass vein 3, and in
extreme cases it is developed only between veins 6 and 4.

In the females the inner edge of the dark border of the fore wings
is abruptly bent inward in the interspace between veins 3 and 4.
Rarely it is broadly curved inward in the anterior half. The inner
edge of the border is usually very irregular, though occasionally
smooth. The spots in the dark border vary greatly in size, being
largest in the small light-orange individuals. They are usually of
different sizes, that in the interspace between veins 3 and 4 being
much smaller than the others, or absent. Rarely they are large and
subequal, and more or less confluent, forming a partially interrupted
broad yellow band separated from the orange or yellow on the inner
portion of the wing by a narrow dark band of uniform width broadly
and evenly curved in its anterior half. Not infrequently the spots are
entirely absent, the black border then closely resembling that of the
male. In spring individuals the dark border is narrower than it is in
summer individuals, and below vein 4 the inner portion may be nar-
row, vestigial, or even entirely absent, the dark border of the female
| then resembling approximately that of the male except for the in-
| clusion of a curved row of four spots in the apical portion.

On the hind wings in the females the dark border may be narrow,
resembling that of the male though with the inner edge vaguely de-
fined. Sometimes it is as broad as it is on the fore wings with the
inner edge parallel to the edge of the wings and completely enclosing

I0O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

a row of subequal yellow spots, largest anteriorly, one in each inter-
space, much as in the females of C. eogene from Kashmir. Usually it
is broad anteriorly where it completely encloses from one to three
spots, the inner border then becoming obsolescent or represented by
a slight dusting of dark scales in the interspaces. Occasionally the
border is represented merely by long narrow dark triangles with
their bases outward that extend inward along the veins. Rarely it is
wholly absent, there being merely a few dark scales at the outer ends
of the anterior veins.

The spot at the end of the cell of the fore wings is usually well
developed, black, and conspicuous, sometimes with an orange, yellow,
or white center. Rarely it is much enlarged, taking the form of a
black circular ring surrounding a white center. It may be reduced to
a narrow line, or it may even be vestigial, represented simply by a
few dusky scales. Not infrequently it is produced into a more or less
extended angle on the side toward the apex. It may be bright orange
instead of black. In pale-orange early-spring individuals it is com-
monly more or less broadly bordered with orange or mixed with
orange scales, sometimes entirely orange.

The spot at the end of the cell of the hind wings varies from pale
straw yellow to orange-red. In light-orange early-spring individuals
it is sometimes very large, its greatest diameter, parallel with the cell,
being as great as the maximum width of the cell. Usually its diameter
is somewhat less than the width of the interspace between veins 4
and 5. Rarely it is very small, only about one-third the width of this
interspace. There is usually a small supplementary spot on its outer
side just across vein 5, but this may be absent, especially if the spot
be small.

The wing bases above are usually marked with blackish scales.
In the males these may be very dark, and the blackish patch is some-
times extended as a sooty infuscation along the lower border of the
fore wings halfway to the outer margin, and also on the hind wings
in the interspace between veins 1 and 2 and the lower half of the cell
downward almost to the anal angle. In the females the infuscation is
less dense than in the males, but more extensive. On the fore wings
it may be confined to the costal border, though it commonly affects
about the basal third of the fore wings and the area below the cell on
the hind wings, becoming diffuse toward the outer edge. Occasionally
the entire hind wing is infuscated except for a light marginal band or
row of more or less indistinct spots.

In the females the veins of the fore wings for their whole length
and the veins in the outer half of the hind wings may be narrowly

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK IOI

blackish. Rarely in the males all the veins are marked by narrow
black lines. We have not seen an example of this last from Virginia,
but Mr. Wagner has one from Nebraska.

The under side of the hind wings may be clear yellow, yellowish
white, dull white, or grayish blue. They are usually more or less
heavily dusted with dark scales, when the dusting is heavy becoming
dusky olive-yellow and in extreme cases in winter dull green with a
broad indefinite lighter border.

On the under side of the fore wings the antemarginal spots are
usually well developed, with the three lowest the largest. In the forms
with very narrow dark borders above they may be as much as four
times as far from the edge of the wings as the inner edge of the dark
border. In the forms with broad dark borders they may be somewhat
nearer the edge of the wings than the inner edge of the dark border.
Usually they are approximately under the inner edge of the dark
border. In philodice they are slightly farther from the edge of the
wings, beyond the inner edge of the black border. This line of spots
is usually straight and parallel with the outer edge of the wings, but
it may be somewhat curved inward. It is more frequently curved in-
ward in philodice. In males in which the dark border is narrow, the
outer edge of the wing markedly convex, and the spot at the end of
the cell of the hind wings small and that at the end of the cell of the
fore wings vestigial, the spots are greatly reduced and not infrequently
are wholly absent.

The fringes of the wings vary from light dull olive with or without
a pink edging to entirely bright pink. They are usually more or less
dull rosy or pinkish.

In Virginia Colias eurytheme occurs in a considerable number of
more or less distinctive though intergrading forms. The relative fre-
quency of these forms and their seasonal distribution have undergone
marked changes since the establishment of this butterfly as a common
species and at present differ more or less widely in different areas.
The greatest variability appears to occur in the southwestern part of
the State where in the lowlands a long series of variations may be
taken at any time during the summer. It is in this region that C.
philodice is most numerous at the lower altitudes, and also is most
variable. Farther north and east where, except in the higher mountain
pastures, eurytheme predominates almost or quite to the exclusion of
philodice, both are less variable. Here the great majority of the in-
dividuals taken at any one time conform to a single type, and vari-
ants are uncommon, although they do occur.

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Several of the forms of ewrytheme occurring in Virginia have re-
ceived names, but these are deceptive in that they imply a certain
fixity that does not exist. The forms are in reality fluid in varying
degrees and for the most part indefinite, varying in size and inter-
grading completely with others.

Each of the forms commonly recognized—eurytheme (=ariadne),
keewaydin, and amphidusa—is subject to parallel variation in Vir-
ginia. Each may be very pale with scarcely a trace of orange, females
of each may be white, and the characteristic color and wing shape of
winter individuals is superposed upon each of these forms equally.
The chief forms of C. eurytheme occurring in Virginia may be de-
scribed as follows.

A small pale form has the fore wings 20-23 mm. long. It is yellow
with a flush of orange on the inner portion of the lower part of the
fore wings, and sometimes also on the hind wings; the dark borders
of the wings are narrow and more or less brownish. This form occurs
alone in early spring, when the bases of the wings above and the under
side of the hind wings are more or less heavily infuscated. Individuals
of this spring type (eurytheme) are on the wing at Farmville, ac-
cording to Frank W. Trainer, until late in May, broadly overlapping
individuals of the orange form (amphidusa), which are on the wing
from May 5. We have found eurytheme on the wing from the end
of March to May 109, at which time individuals of the larger orange
form (amphidusa) were common. But the small pale form also oc-
curs locally throughout the summer in the drier regions, flying with
the much more numerous individuals of the orange form. Summer
individuals of the small pale form, which are usually somewhat rare,
differ from those flying in spring in lacking the infuscation at the
base of the wings above and on the under side of the hind wings.
Toward autumn in dry seasons the numbers of the small pale form
increase and as the weather becomes cool the under side of the hind
wings becomes increasingly infuscated so that the individuals tend to
approach the spring form. Spring specimens have been called eury-
theme (ariadne), but these grade imperceptibly into others taken in
summer and autumn that differ more or less widely, especially in the
degree of infuscation. In habits the individuals of the small pale form
resemble C. philodice rather than the larger forms of eurytheme. The
flight of the males is lower and less swift than that of the males of the
larger forms, and in a narrower zigzag, made up of shorter lines. It
is noticeable that the males prefer the company of the males of
philodice, and associate with them on muddy spots. On the mud they

NO: 7. BUTTERFLIES OF VIRGINIA—CLARK 103

are quite indistinguishable, but when the group takes wing the slight
flush of orange is at once evident. In summer males of this form are
most easily obtained by searching for them among the companies of
the males of C. philodice resting on mud.

The large pale form resembles the preceding in color, except that
the dark borders of the wings are broader and blacker and the size
is larger, the fore wings being usually 25-27 mm. in length, and
rarely as much as 30 mm. It occurs only in summer, and we have
found it only in the extreme southwestern counties, though it may
occur elsewhere. It is possibly a hybrid between C. eurytheme and
C. philodice.

In an intermediate form the size is large, with the fore wings 25-
27 mm. long as a rule, though occasional individuals may have the
fore wings up to 32 mm. long. The color is yellow faintly and uni-
formly flushed with orange except along the costal border of the
fore wings. It resembles the following forms, but is paler. We have
it only from southwestern Virginia. It may be a hybrid.

The early-summer form is the same size as the preceding, which
it resembles except for its darker color. It is the predominant form
in early summer. It occurs throughout the summer, but in most re-
gions it gradually becomes more or less extensively diluted with the
following form, though always remaining abundant. Before 1931
spring individuals in Virginia were all of this form, differing from
summer individuals only in being smaller and with the bases of the
wings above and the under side of the hind wings more or less in-
fuscated. They were somewhat deeper orange than the usual run of
summer individuals though by no means so dark as, and with less
black than, the dark-orange summer form.

The dark-orange form should probably be regarded as an extreme
development of the preceding, with which it intergrades completely
and abundantly. It resembles the preceding, but is deeper orange with
broader and blacker dark margins. The males, in contrast to those of
the preceding forms, always show a violet iridescence which is often
strong. The large females occasionally have the spots within the dark
border of the fore wings nearly or even quite as deep orange as the
ground color of the wing instead of the usual yellow. White females
of this form are often flushed with salmon-pink on the lower inner
portion of the fore wings and with yellow on the hind wings. Though
the fore wings usually measure 25-27 mm. in length, individuals of
both sexes with the fore wings 30-32 mm. long are not infrequent in
low moist areas. As in the case with large individuals in other forms,

104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

there is usually a rounded angle on the hind wings at the end of vein 3.
This form, which flies in summer with the preceding, reaches its most
intense coloration after the first of August when many of the males
are very richly colored with violet reflections and broad and very
black borders. Richly colored individuals increase in number until
the end of the warm weather. The relative proportion of this and
the preceding form vary from year to year. In 1930 no richly colored
individuals were found in the vicinity of Washington, the darkest
being intermediate between this form and the form preceding. This is
the form to which the name amphidusa was applied by Dr. Boisduval.

A curiously colored chrome-yellow form is probably a variant of
the preceding forms. It is rare and should possibly be regarded as an
aberration.

The females of all the forms are more or less frequently white,
these white females being apparently least common in the small pale
form and most numerous in the dark-orange form. The white females
of the latter in the last half of the summer are frequently flushed with
salmon-pink below the cell of the fore wings, fading to white near
the black border, and with greenish yellow on the hind wings ; in some
individuals the basal third of the costal border of the fore wings, the
hairs on the anterior portion of the dorsal surface of the thorax and
on the head, and on the central part of the thorax below, and the legs
are bright pink.

The winter forms of C. eurytheme are worthy of special mention.
During autumn, if the weather remains warm enough to permit this
butterfly to persist in the adult stage, the individuals become paler
and paler, and finally after the middle of November all the individuals
are very pale. The males are washed with light orange on the hind
wings and on the lower portion of the fore wings, on which it passes
gradually into yellow on the anterior half or two-thirds. A few of
these males are of the small pale type with only a very faint trace of
orange on the hind wings and a scarcely stronger flush on the lower
half of the fore wings, and narrow more or less brownish borders;
the lower edge of the fore wings is straight, and the hind wings are
evenly rounded. Although they are as pale, or nearly as pale, as the
small spring form, most of the males have the dark border of the
wings broad and black, while the lower border of the fore wings is
distinctly convex and the hind wings are subangulate at the end of
vein 3 as in summer individuals. Though most of the males are of
medium size or rather small, a few are as large as average large
summer males. In all the males the base of the fore wings and the

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 105

inner third of the hind wings are more heavily and extensively in-
fuscated than in individuals taken at other seasons, and on the hind
wings the infuscation, becoming less and less intense, runs almost or
quite to the outer border in the vicinity of the anal angle.

The females are as pale as the males. Though the fore wings are
bright and clear, the hind wings are dull yellowish green becoming
abruptly lighter near the dark border, sometimes with a very faint
overwash of orange. The dark border of the fore wings is narrower
than in the summer individuals, and the dark border of the hind wings
is much narrowed, reaching inward only to the line represented by the
outer ends of the more or less completely included light spots in the
summer form, and also much shortened, ending about two-thirds of
the distance from the outer to the anal angle. The fore wings are less
pointed and the hind wings are distinctly narrower and longer than
in the summer females.

On the under side the males are infuscated on the outer half of the
fore wings, and heavily infuscated on the hind wings so that the hind
wings appear strongly washed with olive-green, or sometimes entirely
dull light olive-green. The females are much more heavily infuscated
than the males, the entire under surface of the hind wings and the
outer portion of the fore wings being dull olive-green of the same
tone as in Colias hecla, though not quite so dark.

In the summer forms the spot in the middle of the under surface
of the hind wings consists of a circular red-brown ring surrounding a
pearly circular center, with a conspicuous rounded lobe, also with a
pearly center, on the side toward the dark spot on the costal margin.
The width of the ring is usually about half the diameter of the en-
closed pearly spot. The ring may be evenly red-brown throughout, or
there may be a very narrow dark line immediately surrounding the
pearly spot with traces of another partially outlining its outer border,
the rest of the ring being sparsely scaled; or in extreme cases the
large and small pearly spots may be quite independent of each other,
each being surrounded by an exceedingly narrow deep red-brown
border. Rather rarely there is merely a single circular pearly spot
very narrowly ringed with dark red-brown beyond which is a very
faint and indefinite ring only slightly differentiated from the ground
color.

In the winter individuals the supplementary spot decreases in size
and in some cases disappears entirely, while the large pearly spot tends
to become elliptical with the long axis of the ellipse parallel with the
long axis of the cell. At the same time the brown border of the pearly

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

spot on the side toward the margin of the wing, and to a lesser extent
on the inner side, becomes broadened so that the pearly spot lies in
an oval near the proximal end at about its own diameter away from
the distal end. An exactly similar transformation from the usual
double spot into a more or less elongated spot with a considerable area
of red-brown on the side away from the wing base is characteristic
of a number of Arctic and alpine species that are similarly dark green
on the under surface of the hind wings, as for instance C. hecla and
C. nastes werdandi.

The white females of the winter forms are a more or less dark
dull green on the hind wings above, lighter greenish on the fore wings.

The males of the winter forms, no matter which of the forms they
represent, have a weak, slow, uncertain, desultory and more or less
direct flight which is usually high, 3 or 4 feet or more above the
ground. When frightened they fly away in a straight line and rather
rapidly, though much less rapidly than males of the same size do in
summer. In a prolonged autumn with gradually decreasing tempera-
tures all the forms of this butterfly little by little take on the features
of the winter form, slowly converging toward the common pale color
type and wing shape characteristic of that form. But the fully de-
veloped winter forms occur only in very exceptional years. Since this
butterfly became established in the vicinity of Washington we have
seen it only twice—in 1935, when it was on the wing until December
24, and in the winter of 1946-47, when it was on the wing up to Jan-
uary 27, 1947.

Occurrence——The orange clover butterfly is a recent immigrant
into Virginia from the south and west, now thoroughly established
and abundant everywhere. It is perhaps the most conspicuous and
characteristic butterfly of open fields, especially clover and alfalfa
fields in the Piedmont region. It is common, though less conspicuous,
in the peanut and cotton areas in the southeast. In the western part
of the State it is found in abundance in the valleys but is less numer-
ous or even infrequent in the higher and more exposed mountain pas-
tures where the native philodice still predominates. Over most of
Virginia it is far more abundant than plilodice, but in the southwest,
from Montgomery County southward and westward, it is in many
places scarcely more numerous than philodice, in some, indeed, less
numerous. It is primarily a butterfly of low open country, particularly
farmlands, and does not appear to thrive, as does piilodice, in the
more rugged areas.

Imnugration and establishment.—Although at present this is one

~

SE ———— Eee —E—E———————————— ll

i

NO! 7 BUTTERFLIES OF VIRGINIA—CLARK 107

of the commonest butterflies in Virginia, it is a very recent addition
to the fauna of the State, into which it spread from the south and
southwest.

Prof. Ellison A. Smyth, Jr., writes us that during his residence
in Blacksburg, Montgomery County, up to 1925, he never saw eury-
theme there. Dr. George W. Rawson also writes us that he does
not remember having taken eurytheme anywhere in Virginia in his
collecting up to 1925.

Dr. Frank Morton Jones writes that on coming north by rail on
August 10, 1894, he noted the last eurytheme, seen from the train,
at Salisbury, Rowan County, N. C., and the first philodice at Dan-
ville, Va. Between April 18 and 30, 1906, he took philodice at Vir-
ginia Beach, but he can find no record for eurytheme over that
period.

He did not see eurytheme on the Delaware-Maryland-Virginia
peninsula until July 29, 1920, when he took a fresh male at Ocean
City, Md. In 1923, proceeding southward from Wilmington, Del.,
by motor, Dr. Jones found philodice prevalent until he reached Berlin,
Worcester County, Md., where eurytheme began to appear. Farther
south, at Wachapreague, Accomack County, Va., only eurytheme was
seen. In the last week of July, 1925, eurytheme was among his
captures in the Dismal Swamp. In the spring of 1929 Austin B. J.
Clark found eurytheme abundant as far northwest as Lexington,
Rockbridge County, where it was much more numerous than philo-
dice, though northeast of Lexington philodice was much more numer-
ous than eurytheme.

In considering the spread of eurytheme into Virginia it is of
interest to review the history of its establishment in the adjacent
District of Columbia. On November 11, 1886, Dr. Otto Lugger saw
a “eurytheme, or rather a very pale variety of it, looking like a Pieris,”
flying about some flowers of dandelion in the Department of Agri-
culture grounds. This may have been an example of the pale winter
phase. In the United States National Museum there is a male of
the small pale form (eurytheme) taken in the District on September
6, 1904, by the late Prof. C. R. Ely, and we have seen a male of
the dark-orange form (amphidusa) that was taken in the District
about 1910. These individuals were apparently only casuals. There is
a single broken male of the dark-orange form taken in the District,
but without date, in the Henry F. Schénborn collection. In view of
the fact that Mr. Schénborn kept only small series of perfect speci-
mens, this would indicate that it was an unusual capture.

The next recorded individual was a female taken by Austin B. J.

108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Clark on September 24, 1923. In 1925 eurytheme was occasional
in September in the meadows along the river west of Cabin John,
Md. In 1926 it was much more numerous and was found from August
27 until the end of the season. In 1927 it was abundant from the
middle of July until the end of the season. In the higher and more
exposed areas, as about Silver Spring and Somerset, Md., its num-
bers were equal to more than half the numbers of philodice when
the latter was at its maximum—indeed on some days there seemed to
be no difference in the frequency of the two. In 1928 it was first
taken on June 21 and was very common from the first of July on-
ward. In 1929 it was taken as early as May 12. It disappeared dur-
ing the last week in May, but fresh individuals appeared in the second
week in June and by the end of the month it had become abundant,
its numbers being, in some places at least, equal to the numbers of
philodice. It remained abundant until the end of the season. In 1930
it was first noticed on April 27, and at the end of the summer it
far outnumbered philodice.

It has frequently been stated, following Scudder, that prior to
1889 eurytheme was known east of the Appalachians only from a very
few casuals. Our friend the late Prof. Ellison A. Smyth, Jr., as-
sured us that this is an error and that it was common in his boyhood
days about Charleston, S. C. He wrote us that his earliest recollec-
tion of butterflies, in 1874, and his efforts at his then very youthful
start at forming a collection are vividly concerned with the late-
winter or early-spring form, eurytheme, as common around Charles-
ton. He still has in his collection two specimens dated 1876, and
later he added others of the summer forms caught near Charleston.

In February 1889, when he was adjunct professor of biology at
the University of South Carolina at Columbia, he found eurytheme
quite common. He wrote to William H. Edwards about this, and Mr.
Edwards urged him to try breeding it. This he did for 2 years.
He found that the eggs of eurytheme laid on clover in February
produced the form keewaydin, and eggs from this form gave indi-
viduals that were apparently identical with the bright and large
western summer form (amphidusa). He also sent eggs to Mr.
Edwards.

Professor Smyth never saw p/ilodice around Charleston or Co-
lumbia, S. C. In fact, he never saw it at all until he went to Prince-
ton, N. J., in 1880.

In March 1907 Drs. C. S. Brimley and Franklin Sherman, Jr.,
wrote that eurytheme occurs in North Carolina from the east-central
portion (Raleigh) eastward, while philodice is generally distributed

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 109

and abundant in the mountains. In a recent letter Dr. Brimley tells
us that eurytheme is common all over the State and has been ever
since he first began keeping records in 1900.

When William H. Edwards was living at Coalburg, in the Kana-
wha valley in southern West Virginia, eurytheme was entirely un-
known there. The senior author and Hugh Upham Clark found it
common in the Greenbrier, New River, and Kanawha Valleys in 1929.

The evidence appears to indicate that eurytheme is endemic in
South Carolina and in the Piedmont of North Carolina, or at least
has been in these regions for a very long time, but until recently it
did not occur in Virginia or West Virginia. It was first recorded
from the Delaware-Maryland-Virginia peninsula in the summer of
1920. By 1923 it was common as far north as Wachapreague, where
it had supplanted philodice, and to the northward it occurred with
philodice as far as Berlin, Md. In the same year it appeared in the
vicinity of Washington, D. C., where it had been earlier noted as
a rare casual in autumn. Two years later, in 1925, it was occasional
in autumn. In 1926 it was found as early as August 27 and in autumn
was much more numerous than it had been in the year preceding. In
1927 it appeared in the middle of July and later became abundant.
In 1928 it appeared on June 21, and in 1929 it was taken in May, pre-
sumably, therefore, having overwintered for the first time. In the
western part of Virginia it was abundant in 1929 as far north as
Lexington, farther north occurring in lesser numbers. In that year
it was also common in the valleys of southern West Virginia.

Several interesting features have been noticed in connection with
the acclimatization of eurytheme in Virginia. Prior to 1931 the
spring individuals found in the District of Columbia, and presumably
also in nearby Virginia, were all of the dark-colored amphidusa form,
though of rather small size. The small pale form (eurytheme) did
not appear until the last week in July, from that time on slowly in-
creasing in numbers, though being at no time very numerous. In the
spring of 1931 the butterfly first appeared on April 22, and 2 days
later the males had become abundant and females appeared. For
the first time the spring individuals were almost exclusively of the
small pale form (eurytheme), only two individuals of the dwarfed
amphidusa form being noted, one on April 22 and the other on April
24. Ever since then the early-spring individuals have been of the
eurytheme form, though some of the later spring emergences may
be of the amphidusa type.

Up to 1937 it was noticed that whereas the males of philodice were
to be seen individually or in groups of various sizes on every muddy

Ilo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

spot, the males of euryiheme remained always in the fields, except
that a few males of the eurytheme form might sometimes be found
in a group of males of philodice. In the summer of 1937 in several
different places in western Virginia we noticed for the first time males
of the large brightly colored forms of eurytheme resting on mud
singly or in groups. For instance, near Moscow, Augusta County, on
August 14 we saw about 35 males of eurytheme in a compact group
on mud in a road, together with 3 males of philodice and a few males
of Eurema lisa, these last at some distance from the others.

With the increase in the numbers of eurytheme in Virginia the
numbers of philodice have diminished both relatively and absolutely.
By 1934 philodice had become decidedly uncommon except in the
mountain pastures. At this time at the height of the season several
days might pass without a single philodice being seen among the
hundreds of eurytheme.

In the spring of 1936 Dr. Carroll M. Williams wrote us that
philodice was more common in the vicinity of Richmond than it
had been for several years past. Warren H. Wagner, Jr., indepen-
dently reported an increase in the number of philodice in several
districts in Virginia, and we ourselves noticed it in Fairfax County.
But it disappeared almost completely after the spring brood.

In the spring of 1937 Dr. Williams wrote us that, strange to say,
eurytheme was rare at Richmond, and in Fairfax County we found
it much less numerous than it had been in the years immediately pre-
ceding. Later in the season it became common, and philodice also
remained common. But we are not certain whether these yellow indi-
viduals, most of which seemed to be different from our series of
true philodice, were not hybrids. For the first time we found several
mated pairs in which one individual was orange, the other yellow.

Seasons.—Three, four, or five broods. The orange clover butterfly
usually first appears early in April, sometimes as early as the end
of the third week in March, and flies continuously until the end of
the season, usually about the middle or toward the end of October,
though often into November, occasionally as late as the last week in
December, or even until January 27. The second brood begins to
appear in May, sometimes early in the month, the third about the
first of July, the fourth in August, and the fifth in September. Fresh
individuals on the wing in late autumn or early winter, which often
differ more or less widely in color and in wing shape from those seen
in summer, are probably delayed emergences from the fifth brood.
The prolonged emergence period of the spring brood and the progres-
sively longer emergence periods of the later broods make it quite im-

|

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK EEL

possible to determine the broods from observations in the field, for
after early May worn and fresh individuals are found flying together
at all times. In the higher country this butterfly usually does not
appear until late in April, and there are here probably three broods
and a more or less complete fourth. Up to 1931 this butterfly did
not appear in the vicinity of Washington until toward the end of
April or early in May, but since then it has appeared early in April
or even late in March.

According to V. L. Wildermuth the time from the laying of the
eggs to the appearance of the adults in California and Arizona varies
from 16 to 29 days. It is probably from 3 to 4 weeks or somewhat
longer in Virginia.

Mr. Wildermuth says that the winter is usually passed in the pupa,
although in the Southwest both larvae and adults have been taken
during every month of the winter. It would appear from the rela-
tively late appearance of the butterfly in spring in the early years
of its establishment in Virginia that at that time the winter was
passed as a nearly or quite full-grown caterpillar ; as it now appears
in the spring 3 weeks or more earlier than it did some years ago it
probably passes the winter chiefly in the pupa, from which comes the
small pale form, eurytheme, but also as a caterpillar, the overwinter-
ing caterpillars producing the small spring individuals of the form
amphidusa which fly with the later emergences of eurytheme. This,
however, requires confirmation.

COLIAS INTERIOR Scudder
Plate 17, c, d, g,h

Range.—Known only from a large male with the fore wings 30
mm. long caught by John E. Graf on Middle Mountain, Highland
County, at an altitude of 4,010 feet, on June 14, 1936, and another
similar male taken by William D. Field in the same locality on June
19, 1948.

Occurrence.—During a visit to Middle Mountain in 1936 John E.
Graf was so kind as to collect a number of butterflies for us. These
were examined and recorded as soon as received and then turned
over to a preparator for setting. At the time the specimen was re-
garded as an example of Colias philodice, as we had no idea that
C. interior occurred in Virginia. However, when it was set it was
added to the series of C. interior in the Museum collection, presum-
ably by John F. Gates Clarke, who was then in charge. We have to
thank Dr. William Hovanitz, who found it while studying the

II2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

Museum’s series of C. interior, for calling our attention to it. The
records of this species from Virginia by Hovanitz (1950) and Klots
(1951) are based on these specimens.

It is quite possible that this species occurs regularly in certain
Canadian areas in western Virginia, having been overlooked because
if its superficial similarity to C. philodice.

Genus ZERENE Hiibner
ZERENE CAESONIA (Stoll)
Plate 2, d

Range.—Recorded from Suffolk, Nansemond County, July 27,
1938 (Otto Buchholtz), and from Bayford, Northampton County
(Florence Walker).

Sidney A. Hessel writes us that sometime between October 12 and
20, 1937, when driving from Norfolk to Washington he “recognized a
caesonia flying across in front of the car, and being my first I could
not resist a short stop. I must have seen Io or more in 20 minutes
or less. They were in an overgrown field not more than 2 acres in
area. The general picture I recall of the locality was that the ad-
jacent fields were in truck farming and the vicinity definitely unat-
tractive for collecting. The land along the roadside was in small
parcels, with houses, gas stations, etc., rather frequent. For want
of the exact locality the specimens were labeled ‘30 miles south of
Washington, D. C.’” This would place the locality in the vicinity
of Quantico (Triangle) in Prince William County.

Occurrence.—The dog’s-head clover is an infrequent casual visitor
from the south found in open country.

Genus PHOEBIS Hiibner
PHOEBIS PHILEA (Linné)
Plate 10, d

Range.—Recorded from Gala, Botetourt County, October 11, 1941.
(Donald Stoutamire), and from Salem, Roanoke County, October 15,
1928 (Prof. Ellison A. Smyth, Jr.) ; both specimens are males.

Occurrence.—A rare casual or accidental visitor.

PHOEBIS SENNAE (Linné)
Plate 10, c
Range.—Throughout the State.

V ariation.—According to F. Martin Brown the form eubule, with
the under side of the hind wings and the apical portion of the fore

|

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK II3

_ wings rather deep yellow, contrasting with the lighter greenish yellow
of the inner portion of the under side of the fore wings, and the
under side of the hind wings sparsely spotted, appears in the southern
States in spring and autumn. It ranges as far north as the Coastal
Plain of Virginia, where it occurs in autumn (specimen in the U. S.
National Museum from Suffolk, October 2, 1937) together with
the following form. The form drya, uniformly slightly greenish
yellow with the under side of the hind wings immaculate or nearly
so, occurs throughout the range of the local subspecies and is by far
the commoner form in Virginia, ranging throughout the State.
Farther south, according to Mr. Brown, it appears in autumn.

Occurrence—A summer visitor, appearing usually after the middle
of June, occasionally late in May, rarely late in April, on the outer
Coastal Plain, and moving up the shores of Chesapeake Bay to the
Northern Neck and inland as far as Richmond. On the Eastern
Shore it appears first on the western coast, passing across the northern
part of Accomack County to Chincoteague Island. Soon afterward it
appears in the southwestern portion of the State moving north-
eastward, chiefly along the eastern side of the Blue Ridge, though
also in the Shenandoah Valley. By early August it has usually be-
come very common on the entire Coastal Plain and locally in the
west, and has spread over most of the State, with local foci of
abundance coinciding with centers of abundance of its chief local
food plant (Cassia chamaecrista). In September it is usually abun-
dant all over the Coastal Plain, common in nearly all sections of the
west, reaching even the mountain tops, and frequent everywhere
else except in the extreme north, where it is usually more or less
casual.

This butterfly is irregular in its occurrence. In some years (as in
1939) it may cross the border into the south-central part of the
State as early as the last week in April. In other years (as in 1940)
it may be almost wholly absent, represented only by a few individuals
on the outer Coastal Plain and in the extreme southwest. In still
other years (as in 1941) it is everywhere common to abundant.

A single record from the Dismal Swamp near Suffolk on April 1,
1938, and a female captured at Rocky Run, Fairfax County, May 19,
1935, suggest that this butterfly occasionally overwinters in Virginia.

Migratory flights in Virginia—Although we have kept a special
watch for migratory flights of this butterfly in Virginia, very few
such flights have been noticed. This species is often exceedingly
common on the Coastal Plain and abundant wherever the food plant

II4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

grows, but the individuals dash about irregularly over the fields in
any direction. The spread of the butterfly over the State each year
follows a definite pattern, but this may be due to the distribution
of the food plant on the Coastal Plain and along the western moun-
tains rather than to a directional migration on the part of the
butterfly.

During the course of a drive from Richmond to Burkeville, Not-
toway County, on September 24, 1933, we noticed many individuals
of this species all flying toward the southeast across the road. They
flew as widely separated individuals, far apart, but all in the same
direction.

At Greenville and Lyndhurst, Augusta County, on August Io,
1937, and at Lovingston, Nelson County, 2 days later, occasional
individuals were observed flying toward the northeast. Although the
individuals seen were few, all seen were traveling in the same direc-
tion. At Roseland and Massies Mills, Nelson County, on August 12,
1937, the food plant (Cassia chamaecrista) was extraordinarily
abundant, some old fields being completely filled with it. Here the
butterflies were very common. Most of the individuals seen were
flying irregularly about the weedy fields, but some were observed in
traveling flight toward the northeast. This region would seem to be
an excellent relay point in the seasonal extension of the range toward
the northeast.

At the Agricultural Station north of Alexandria on September 12,
1937, occasional individuals were seen traveling with great speed in
direct flight toward the southeast.

At Hanover Court House, Hanover County, on August 31, 1935,
and again on September 23, 1937, a few individuals were seen flying
directly toward the east.

These few directional flights that we have observed in Virginia
were in no sense mass migrations. The individuals were in all cases
widely separated and seldom in sight of each other. The maintenance
of the same direction by each individual as it came in sight was, how-
ever, striking and significant.

Seasons.—Two broods. Our records indicate an early-spring brood
reaching a peak in the latter half of April and disappearing toward
the end of May. The individuals of this brood in Virginia are prob-
ably all immigrants from the south. In the last half of June the but-
terfly reappears and becomes increasingly common until the first half
of September, after which the numbers fall off.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK II5

Genus NATHALIS Boisduval
NATHALIS IOLE Boisduval
Plate 16, g, h

Range.—Recorded from Roanoke River, South Salem, Roanoke
County, September 26, 1944; and from Katz Hill, Salem, on the same
date (Dr. Carl W. Gottschalk). One of the specimens was presented
by Dr. Gottschalk to the United States National Museum.

Occurrence.—A rare casual; Dr. Gottschalk wrote that the speci-
men from Katz Hill was perfect, and he was sure it had not flown
for but a day or so, and that it must have come from a larva hatched
out in Salem.

Genus EUREMA Hiibner
EUREMA NICIPPE (Cramer)

Plate 2, e

Range.—Throughout the State, but in the mountainous regions
living only in the valleys along the larger streams.

Variation—The males of this species are occasionally clear bril-
liant yellow instead of orange. We have seen none of these, in
Virginia, but the Schonborn collection in the United States National
Museum contains two fine specimens from the District of Columbia.
The females are sometimes very light yellow, rarely white. In the
females the dark border of the wings may be broad and complete, or
it may not extend below vein 2 on the fore wings, or beyond vein
2 on the hind wings; commonly the ground color of the fore wings
extends to the outer margin between veins I and 2.

As in many other species of this group, the fore wings, more
rarely both fore and hind wings, may be more or less heavily
suffused with black scales.

The under side of the hind wings varies greatly in the extent of
the dark markings, which in extreme cases may occupy most of

the central portion; it is sometimes strongly suffused with pink.

Occurrence.—Found in open country, usually near woods, espe-
cially in broad river bottoms, along the sides of streams bordered with
more or less level ground, and near ponds; regularly present and
locally frequent or common in the southwestern portion of the State,
in the Shenandoah Valley, and along the eastern side of the Blue
Ridge, as far northeast as Augusta and Madison Counties; also
regularly present, though less numerous, on the Coastal Plain (except
the Eastern Shore) and on the Chain Bridge flats along the Poto-
mac River opposite Arlington County ; elsewhere of irregular or more

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

or less casual occurrence, though in some years (as in 1941) fre-
quent to common everywhere. Except in the western valleys, on
the Coastal Plain, and on the Chain Bridge flats this butterfly sel-
dom survives the winter in this region.

Seasons —From three to six broods, depending upon local condi-
tions. The nicippe appears on the outer Coastal Plain at the end of
March, later elsewhere, and flies into early October or later, fresh
individuals emerging after a few warm days up to at least the mid-
dle of December. The number of broods during the summer varies
from three to six and probably is usually four or five. In hot weather
in regions where the food plant is abundant the entire life cycle occu-
pies less than a month.

Dr. Warren H. Wagner, Jr., has a male taken March 21, 1938, in
downtown Washington, and a female taken April 27, 1935, at Cabin
John, Md.

EUREMA LISA (Boisduval and LeConte)
Plate 0, h

Range.—Throughout the State.

V ariation—Most individuals have the fore wings 15 to 18 mm.
long, but in some they reach 20 mm. In the males the ground color
of the upper surface is usually light clear yellow, rarely deep yellow,
or with a dull orange tinge. In some individuals the dark borders,
especially on the hind wings, are considerably broadened, while in
others they may be narrowed as in the West Indian and Central
American EF. euterpe. Occasionally the dark border on the hind wings
is reduced to a small dark patch at the outer angle followed by a thin
marginal line (form clappi Maynard), such individuals more or less
resembling females except for the brighter color. Rarely the fore
wings are more or less heavily dusted with black scales.

The females are almost always paler than the males, not infre-
quently cream white, much more rarely chalky white (form alba
Strecker). Of this last, pure white, variety we have seen only one
from Virginia, captured by Dr. Warren H. Wagner, Jr., in West-
moreland County.

Occurrence—Found in open fields and along roadsides, sometimes
entering open woods; in the last half of the summer usually common
to abundant everywhere, though in some years (as in 1940) local
and not common.

So far as we have been able to determine, this little butterfly, al-
though it overwinters locally far to the northward, is only a summer
visitor to Virginia, coming in each year from farther south. Our

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK Ly,

earliest record is for an individual taken near Fredericksburg on
May 29. After the middle of June we have found it fairly common
on the eastern Coastal Plain, along the shores of Chesapeake Bay,
on the Northern Neck, and casually as far north as Prince William
and Fairfax Counties, but we have no June records from elsewhere
in the State. By the end of July it has penetrated in greater or lesser
numbers to every portion of the State, and in August and later it is
usually one of the commonest butterflies everywhere, reaching even
the high mountain pastures. In some years, however, it is absent
from the greater part of the State, occurring in small numbers only
on the Coastal Plain and in the southwest.

Seasons—Usually four broods. The lisa appears toward the
end of May and flies until about the middle of June. It reappears
in the last week in June and from that time onward flies continuously
in increasing numbers until the end of the season, usually the first
week in October, but in some years as late as about the middle of
November. The butterflies found late in May and early in June are
individuals that have strayed from farther south, or the young of
such individuals. The second brood begins to appear usually toward
the end of June, the third after the middle of July, and the fourth in
August. The fresh individuals on the wing at the end of the sea-
son may be in part representatives of a fifth brood. So far as is
known, this common little butterfly does not survive the winter in
Virginia.

EUREMA JUCUNDA (Boisduval and LeConte)
Plate 16, 4, 7

Range.—Recorded only from the southern portion of the State,
in Roanoke, Giles, Bedford, Rockbridge, Prince Edward, Henrico,
and Prince George Counties. Most of the records are from Roanoke
County.

Variation—All the specimens from Virginia are typical jucunda,
with the under side of the hind wings pure white.

Occurrence.—This species is for the most part an irregular visitor
to Virginia, appearing from time to time in various localities east
of the mountains, occasionally in considerable numbers. All but
two of the records are dated after the first week in July, which
would indicate a summer visitor. Unfortunately the single spring
individual is not available for study. It was seen by the late Prof.
Ellison A. Smyth, Jr., who wrote us under date of June 26, 1937:
“Two or three springs ago I saw a jucunda flying down the hillside
about 100 yards from my house, here near Salem. I ran back to

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the house for my net, but could not again locate it when I returned.
The black border on rear of fore wings, and the white under surface
of hind wings were very much in evidence, however, and it was
beyond question jucunda.”

It is possible that this species is established in this section of the
State, though the number of records from Roanoke County may be
due simply to the fact that the records from this area are more com-
plete than from any other.

In Virginia this butterfly has been found only in open country.

Seasons.—There would appear to be four broods in Virginia, one
in spring, the second in June and early July, the third in the last
week in July extending into early August, and the fourth late in
August and September.

Family PAPILIONIDAE

Subfamily PApILIoNINAE
Genus BATTUS Scopoli
BATTUS PHILENOR (Linné)
Plate 10, a, b, c, d

Range.—Throughout the State.

Variation—Lord Rothschild and Dr. Karl Jordan have pointed
out that individuals of the blue swallowtail from the Atlantic States
generally have a more elongate hind wing than the majority of
southern and western specimens ; the submarginal spots of both wings
are often smaller, the spots on the upper side of the fore wings are
more often absent or vestigial in the males, and the outer margin
of this wing is rather more concave; the metallic distal area of the
under side of the hind wings is less often green, and the body is
rarely so shaggy as in some western specimens. But they remark
that these distinctions are by no means reliable, the species being
apparently in a process of separating into an Atlantic, a southern, and
a Pacific coast form.

Early-spring individuals in Virginia (pl. 19, d) are small, with
the fore wing about 40 mm. long. In contrast to the other local swal-
lowtails, except G. marcellus, both sexes appear at the same time
and are of the same size. The hair on the body is very long, and
there is a tuft of long hair on the frons. The fore wings are less
pointed than those of summer individuals, and the outer border is
straight, while the hind wings are more rounded and the tails are
relatively shorter. The whitish spots on the wings are enlarged, these
in spring males resembling those in summer females, and the color

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK I1g

is dull, the males being scarcely brighter than the brightest summer
females.

These early-spring individuals approach some varieties of the form
from northern California known as hirsuta Skinner and _ possibly
might be referred to it. This is another example of a form repre-
senting a geographical race in one area, but only a seasonal variant in
another.

Together with the hirsuta-like form there occurs sporadically in
northern Virginia another small form of the same size with the tails
reduced to mere vestiges, the white spots much larger, and the metal-
lic distal area of the under side of the hind wing more or less reduced

in width, its inner edge, and the white discal dots, being more or

less distant from the cell. In Virginia this form seems to appear
only when the butterfly emerges in warm weather immediately fol-
lowing prolonged and severe cold. We know it in Virginia only from
Leesburg. This is almost, perhaps quite, identical with the form
acauda Oberthiir, which appears as a mutant in various places under
certain conditions. It is recorded from Yucatan and Guadalajara,
México; New Mexico; the Mesa Verde National Park, Colo.;
northern Virginia; near College Park, Md.; and Brooklyn, N. Y.
Late-spring individuals, as in the case of the yellow (Papilio
glaucus) and the zebra (Graphium marcellus) swallowtails, approach
more and more closely the summer form. As with P. glaucus, the
fully developed early-spring form does not appear after mild winters,
the earliest individuals to emerge under these conditions being of
the late-spring form. Also as in P. glaucus, the early-spring form
seems not to occur on the eastern Coastal Plain.
In summer individuals from Virginia (pl. 19, a-c) the fore wings
are 50-55 mm. long in the males, and 50-60 mm. in the females.
Occurrence.—Found in open places in woods, in open country near
woods, and in the farming districts wherever the Dutchman’s-pipe
(Aristolochia macrophylla) is grown as an ornamental; very com-
mon throughout the higher country, particularly in the mountains,
less common and more local on the lower Piedmont, infrequent on
the Coastal Plain, and generally rare or local on the Eastern Shore.
Seasons ——Two broods and a partial third. The blue swallowtail
appears shortly after the middle of April and continues on the wing
through May and into early June. Shortly before the middle of June
the first individuals of the second brood appear, and the butterflies
gradually increase in numbers. The individuals of this brood con-
tinue to emerge through July and August and into September. In
September they are joined by more or fewer individuals of a third

I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

brood that fly until the end of the season, usually early in October,
though sometimes as late as near the middle of November.

Genus PAPILIO Linné
PAPILIO CRESPHONTES Cramer
Plate 10, e

Range.—Throughout the State.

Variation—The giant swallowtail is subject to considerable varia-
tion in size and in the extent of the yellow markings on the upper
surface, particularly the width of the median band on the fore wings
and the band across the base of the hind wings. These variations ap-
pear to be individual and to have no demonstrable relation to locality
and but little to season. Spring individuals as a rule are somewhat
smaller than those flying in summer, spring males having the fore
wings 55-60 mm. long. But one of the largest individuals we have
seen was a female taken on May 29, 1939, near Bloomery, W. Va.,
about a mile and a half west of the Frederick County line. The
largest specimen we have seen is in the Schdnborn collection in the
United States National Museum and was taken in southern Mary-
land; it is a female with the fore wings 73 mm. in length.

Occurrence——Found in open country with scattered trees, par-
ticularly about farmhouses, in brushy land, in open deciduous or pine
woods, and commonly seen in gardens. It is fairly common, some-
times quite common, in the extreme southeastern portion of the State,
particularly on Knotts Island, and is regularly present, though not
in large numbers, northward along the outer Coastal Plain as far as
Gloucester County, and on the Eastern Shore (Northampton and
Accomack Counties). It also appears to be regularly present, though
not very common, in a few localities in the west and north, as about
Narrows in Giles County, between Front Royal and Limeton in War-
ren County, and at Great Falls in Fairfax County. Elsewhere it is
of irregular or casual occurrence.

Dr. Frank Morton Jones writes us that this species breeds regu-
larly on Chincoteague Island, Accomack County, on Zanthoxylum
clavaherculis. On August 27, 1891, he found larvae of various ages
and pupae. Adults were feeding on the flowers of milkweed (Ascle-
pias). On September 28, 1928, larvae collected here earlier in the
month pupated.

The males of this species have the habit of frequenting definite
flyways in open pine or deciduous woods. These flyways are near
and parallel to water and may be a mile or more long, but are nar-

|
|
|

——————— OO CF

= a

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK I2I

row, perhaps a hundred feet in width. Here the males fly back and
forth 5 to 8 feet above the ground. We have noticed such flyways
at Kiptopeake in Northampton County and along the cliffs forming
the east bank of the Shenandoah at Limeton in Warren County.
Dr. Warren H. Wagner, Jr., reported one near Lynnhaven in
Princess Anne County, and on July 17, 1947, David Shappirio found
this butterfly common along the south bank of the Potomac at Great
Falls, Fairfax County. Many years ago a flyway was reported on
the Maryland side of the river at Great Falls.

Seasons.—Two broods. The giant swallowtail appears just before
the middle of May and flies through June and into early July. The
second brood appears after the middle of July and flies until the
end of the season, usually about the end of the first week in Sep-
tember, though sometimes until the end of the month.

PAPILIO PALAMEDES Drury
Plate 22, c,d

Range.—Swamps of Princess Anne, Norfolk, Nansemond, Isle of
Wight, Gloucester, King William, and Northampton Counties ; acci-
dental in Fauquier County and the District of Columbia.

Variation—Individuals from Virginia are usually somewhat
smaller than those from farther south, with the fore wings 50-53
mm. long, those from South Carolina and Florida having the fore
wings usually about 60 mm. long. Late in summer, however, large
individuals become rather common in the Dismal Swamp, where we
have taken several with the fore wings 59 mm. long. In Virginia
individuals of the spring brood average smaller than those of the
summer brood. In Virginia, also, the butterflies on the wing at any
one time are of fairly uniform size. Farther south the variation in
size is greater, small individuals resembling in size those found in Vir-
ginia flying with the larger ones. There appears to be no constant
local difference in color.

When freshly emerged this species is deep sooty black with bril-
liantly contrasting cream-colored markings, but the black soon fades
to brown and the light markings darken. Most museum specimens
give a poor idea of this handsome swallowtail in life.

Occurrence.—Confined to the great swamps and wet woods of
the Coastal Plain and their immediate vicinity, but usually abundant
in the restricted areas in which it is found.

Seasons.—Two broods. This species appears in the last half of
May and by the end of the month has become common. By the

I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

middle of June the individuals are all more or less worn and faded,
and at the end of the month few are to be seen. The earliest indi-
viduals of the second brood are on the wing shortly after the mid-
dle of July, and the butterfly soon becomes abundant. Toward the
end of August the individuals become worn, though a few, in excep-
tional years many, fresh ones are still in evidence, and after early
September they decrease in numbers, disappearing at the end of
the month or early in October.

PAPILIO TROILUS Linné

Plate 22, a, b

Range.—Throughout the State.

Variation.—In southern Florida and as far north as eastern South
Carolina and west to Houston, Tex., a form of this butterfly occurs
in which the submarginal spots of both wings are much enlarged
and light-colored, and on the under side there is frequently a narrow
yellow band on the hind wings corresponding to the yellow band in
Papilio palamedes, sometimes also a subbasal band on the fore wings,
and a streak behind vein SMg.

This is the prevailing form in southern Florida, but the typical
form, with which it intergrades, occurs everywhere with it. Northward
from South Carolina and the Gulf coast this form (tlioneus J. E.
Smith) departs less and less widely from the usual form, and be-
comes less and less frequent. In coastal and also in southwestern
Virginia in summer and early autumn individuals are occasionally
found that resemble the less extreme examples of ilionews from
Florida. One of the most striking of these was captured at Cabin
John, Md., and another almost as extreme in Augusta County. The
enlarged light submarginal spots render these individuals conspic-
uously different from the usual form when in flight. Farther north
the difference between this and the usual form becomes less and less
marked until in eastern Massachusetts it is represented by individuals
with the submarginal spots only slightly enlarged, those on the hind
wing having more or less extensive light centers; the yellow stripe
on the under side of the hind wings is very rare, though it does occur.

The form ilioneus has been commonly treated as a subspecies. But
it occurs throughout the range of the species, at least in the east,
differing widely from the typical form in Florida, but northward
departing less and less from it so that in the northern portion of the
range the two are scarcely to be distinguished. In this it agrees with
the brown males of Papilio glaucus, which, scarcely differing from

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 123

the usual males in Massachusetts, where they are usually overlooked,
reach an extreme development in Florida.

In the males the broad band across the hind wings may be yellow-
green or blue-green or, much more rarely, a delicate grayish blue.
The submarginal lunules on the hind wings are usually pale yellow-
green or blue-green, often more or less extensively yellow in the
center, rarely entirely yellow like those on the fore wings.

The normally orange spot on the anterior border of the hind wings
is occasionally green, like the submarginal lunules. Rarely it is fol-
lowed by from one to three progressively smaller spots in the inter-
spaces below, these additional spots being situated over the cor-
responding spots on the under side of the wing.

Early-spring individuals are small with the fore wings 42-45 mm.
long ; the body is hairy and there is a tuft of long hair on the front
of the head, and the submarginal spots are somewhat enlarged. They
resemble individuals from the northern border of the range. These
small individuals are soon joined by larger ones of intermediate
character. In summer individuals the fore wings are usually 50-55
mm. long.

Occurrence.—Found in open woods and swamps, in scrubby land
with sassafras, in fields more or less near woods, and in gardens;
generally distributed, and everywhere common to abundant; except
on the Eastern Shore and the Northern Neck, and in the coastal
swamps, this in the summer is usually the commonest swallowtail.
In spring it is outnumbered by the yellow swallowtail (Papilio
glaucus), but with the yellow swallowtail the summer brood is only
partial, many of the pupae that might be expected to give forth adults
in the summer remaining dormant until the following spring, which is
not the case, or at least does not occur to anything like the same
extent, with P. troilus.

Seasons.—Two broods, with a partial third. The males of the
spicebush swallowtail appear usually just after the middle of April,
occasionally earlier, sometimes as early as the first of the month on the
Coastal Plain, and the females appear about a week later. The indi-
viduals increase in numbers, and the butterfly becomes common after
the middle of May. It flies until late September or even well into
October, occasionally until almost the middle of November. The sec-
ond brood puts in its appearance about the middle of June, increasing
in numbers through July; the period of greatest abundance is in
August and early September. Fresh individuals found from late Sep-
tember until the end of the season, usually few in numbers though
numerous in some years, are undoubtedly of the third generation.

124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

PAPILIO GLAUCUS Linné

Plates 19, f; 20; 21

Range.—Throughout the State.

Variation—The yellow swallowtail is the most widely distributed
and over the greater part of its range the commonest of the swallow-
tails, at least in spring. The western race (rutulus) shows surpris-
ingly little variation throughout its range, but in the north and east
this is one of the most variable of all our butterflies, and certainly
the most variable of the American swallowtails.

This variation, which is much more marked in the females than
in the males, is correlated in part with locality. Far northern indi-
viduals (pls. 20, a, b; 21, a-c) and the earliest individuals of the
spring brood farther south are much smaller than southern sum-
mer and extreme southern individuals, and they also differ in color
pattern, especially the females. In the north this variation is cor-
related in part with local differences in humidity and other factors.
In sub-Arctic regions where it becomes more or less localized this
butterfly tends to vary from place to place, and to occur in a great
number of usually slightly different local forms somewhat after the
fashion of the species of Boloria, Oeneis, and Erebia living in the
same region. In the central eastern portion of the range, from
southern New England to North Carolina, the variation, other than
seasonal, is for the most part purely individual, several more or
less widely different forms occurring together.

Except in the matter of size there is little definite correlation be-
tween any of the several forms and locality. All far-northern indi-
viduals are small, but the earliest spring individuals from as far
south as the mountains of North Carolina may be equally small and
similar in wing shape and color pattern, though here the later spring
and summer individuals are large. Furthermore, small northern indi-
viduals may be of a type chiefly confined to the north, or they may be
dwarf representatives of a southern form. There are therefore no
grounds for a further subdivision of the species in this region, though
its great variability, partially correlated with locality, should be
recognized.

In the extreme northern portion of the range the fore wings in
both sexes measure from 40 to 45 mm. in length. In males from
Florida the fore wings measure about 65 mm. in length, and in Florida
females the fore wings measure up to 76 mm., these females being
very nearly twice as large as the smaller northern females.

In the East the yellow swallowtail occurs in two widely different

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 125

forms between which every conceivable intergrade occurs. One of
these forms is most highly developed along the southern border of
the range. In this form the male is always yellow (pl. 20, d), varying
from light clear yellow to a fairly deep yellow-brown, and the female
is usually blackish brown (pl. 19, f), if yellow (pl. 20, g) having a
great amount of blue on the dark border of the hind wings. In this
form sexual diversity reaches the maximum, whether the females are
blackish or yellow. The caterpillars usually live high in the trees,
preferring especially different species of ash (Fraxinus), magnolia
(Magnolia), and the tuliptree (Liriodendron), though feeding also
on many other plants, including the introduced camphor (Cinnamo-
mum camphora).

The other form is most highly developed in New England, northern
New York, and the adjacent parts of Quebec and New Brunswick.
In this form the male and female are alike in size, wing shape, color,
and color pattern (pl. 20, c, e, f). The caterpillars feed mainly on
the leaves of bushes and of the lower limbs of trees, preferring wild
cherry, birch, and poplar, though also feeding on many other kinds of
plants.

The males of the second form differ from the males of the first
in having the fore wings somewhat more pointed, the hind wings
narrower and usually more strongly scalloped, the tails more slender,
the black markings, especially the black outer border of the hind
wings, narrower and more sharply defined, and the black abdominal
border of the hind wings usually much broadened. On the under side
the submarginal spots on the fore wings are usually coalesced into a
tapering light band, and on the hind wings the dark border is very
narrow and is heavily washed with light scaling, and the included
submarginal lunules are large and less strongly curved than in the
first form, with a large central orange patch, or in extreme cases
wholly orange.

This form shows a decided approach to the western rutulus in the
usual occurrence of a broad submarginal band instead of a row of
spots on the under side of the fore wings, in the occasional narrowing
of the submarginal spots on the fore wings above, in the straighten-
ing of the lunules, which are sometimes narrowed, on the hind wings,
in the occasional obsolescence or even in extreme cases the complete
disappearance of the orange spot at the anterior angle of the hind
wings, and occasionally in the great reduction of the light center of
the subapical comma-shaped mark on the fore wings below. The
similarity of the sexes, and the similarity of the food plants and habits

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

of the caterpillars, increase its resemblance to rutulus, with which
it has often been confused.

In the extreme south the females are mostly blackish brown (pl.
19, f), but not very far to the northward yellow females (pl. 20, g)
become frequent. These gradually increase in relative abundance,
though in most of Virginia, West Virginia, Kentucky, southern Illi-
nois, and central Nebraska dark females are still more numerous
than yellow ones. North of this line dark females rather rapidly
become less common than yellow ones, then merely occasional, and
in the vicinity of New York City, Buffalo, northern Ohio, southern
Wisconsin, central Iowa, and South Dakota they are rare. As very
rare casuals dark females are known from Newfoundland (pl. 21, c)
and southern Alberta, and even from the Upper Liard River in
British Columbia. In southern Florida the females are always yellow.

In the extreme south the yellow females, as far as we have seen,
are all of a single type, but farther northward, as the proportion of
yellow to dark females increases, the yellow females—and to a certain
extent the dark females also—become diversified, appearing in a
number of more or less distinct types differing in wing shape, in
color, and in color pattern. Some of these types are well marked and
fairly stable, while others are simply inconstant intergrades. Some
are common, while others are rare, or even very rare. Some are
generally distributed, and others are known from a restricted area
only. These different female types seem to be most numerous and
most diversified within the zone wherein dark and yellow females are
about equal in numbers, or at least the former are common, but
some are to be found north of this zone wherever dark females occur,
and one is almost exclusively confined to the area wherein dark
females are never found. "

Within the zone in which the dark females are rare—that is, from
central New York to northern Ohio, southern Illinois, southern Wis-
consin, central Iowa and South Dakota—the size of the individuals
is considerably less than it is in the region where dark females are
abundant, though large individuals, especially females, are not in-
frequent in the second brood.

North of this zone the size of the individuals rather rapidly de-
creases with latitude, and also with altitude. From Alaska to New-
foundland throughout the whole of Canada, except in southern
British Columbia and Ontario, and southward into the mountains of
Montana, central New York, and New Hampshire, the great majority
of the individuals have the fore wings between 40 and 45 mm. in
length, while even smaller ones are not infrequent.

WO7, BUTTERFLIES OF VIRGINIA—CLARK 127

In addition to decreasing in size, the butterfly here exhibits an-
other peculiarity—a greater or less proportion of the females tend
to resemble the males in wing shape and in color pattern. This
assumption by the female of the male wing shape and color shows
curious geographical irregularities. There is the least difference
between the sexes in New England (pl. 20, c, e, f) except in the
southern and southeastern portions, and in central and northern New
York, where the great majority of the females, at least in the spring
brood, are practically indistinguishable from the males. Within this
area the size varies from large in southern New England (pl. 20,
e, f) to very small in the White Mountains of New Hampshire, but
it is everywhere less than the size farther south, and usually much
less. As the form with the sexes alike is especially characteristic of,
and most highly developed in, New England it will be referred to as
the New England form.

In Newfoundland the individuals are always small (pls. 20, a, b;
21, a-c). In the collections we have examined from that island some
females are almost identical with the males in coloration, though
the black markings are usually somewhat broader and less sharply
defined. In both sexes the submarginal spots on the under side of
the fore wings are usually coalesced into a tapering band, and the
dark border of the hind wings beneath is as narrow as in individuals
from New England. Other specimens, however, are quite different,
the males having the submarginal spots on the under side of the
fore wings more or less completely separated from each other (pl.
20, b) and the dark border of the hind wings on both surfaces as
broad as in perhaps the majority of the males from Virginia. With
these males fly females which are very different from the males and,
except for being always of small size, are similar to corresponding
females from southern Maryland. Though they are very rare, black
females of this type occur (pl. 21, c).

Thus in Newfoundland a close approximation to the New England
type is shown by certain individuals of both sexes, but the butterfly
is chiefly represented by individuals which, though dwarfs and usually
somewhat dusky, are obviously of the type found in Maryland and
Virginia, like that type having an alternative black female, and
quite different from the New England form in which black females
are never found. This southern form just reaches southern New
England, and there are a few records of black females from southern
Connecticut. It is somewhat surprising to see it reappear far to the
northeast in Newfoundland where it is presumably a local develop-
ment from endemic stock of the New England type. In both sexes

128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

complete intergradation between these two types occurs in Newfound-
land, just as it occurs in the corresponding, but much larger, indi-
viduals in southern New England and in New York.

Practically all the specimens from Newfoundland that we have
been able to examine have been without definite locality. It is prob-
able that in portions of the island the New England form prevails,
or at least is common, while in other portions the dwarf representa-
tive of the southern type more or less completely replaces it. In a
single small bog in Essex, Mass., on Cape Ann, we have taken during
the course of a week late in July large and small females indistinguish-
able from males, very large deep-yellow females with a large amount
of blue in the dark border of the hind wings, and every possible
intergrade between the two forms. But just west of Boston, about
30 miles away, we have never found any but malelike females.

Only two individuals from the Hudson Bay region have been
available for study. A female from Kettle Rapids on the Nelson
River is quite like many females from New England. The fore wings
are 50 mm. in length. The hind wings above show only a trace of
blue—no more than in some New England males—in the dark bor-
der. A male from the Piquitenay River on the Hudson Bay Railway
has the fore wing 44 mm. long and resembles New England males.

Specimens from southern Ontario resemble others from southern
New York, but are smaller.

In western Ontario, northern Saskatchewan, and along the Atha-
basca River in northern Alberta both sexes are of the New England
type with the females showing relatively little variation. The dark
border of the hind wings is often narrower than in the New England
form, and on the under side more strongly suffused with light scales,
with the included submarginal lunules commonly narrower and more
straightened and bandlike, while the submarginal spots on the fore
wings above may be narrowly linear.

Farther to the northwest in the region of Great Slave Lake in
Mackenzie, the females again become variable and may be duller than
the males with broader and less sharply defined black markings, and
a considerable amount of blue on the black outer margin of the hind
wings above. But the wing shape is only slightly different from that
of the males, more or less suggesting, though by no means closely
approaching, the wing shape of the more extreme females from New-
foundland, and the dark border of the hind wings is less broadened
than in the latter. The submarginal lunules on the hind wings are
straightened and bandlike, and the submarginal spots on the fore
wings are narrowly linear.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK I29

Still farther to the northwest, in eastern and central Alaska, the
females again become very similar to the males in wing shape and in
color, usually having no more blue on the hind wings above than the
males, and never having much more. In this region the females are
more uniformly like the males than they are anywhere else except in
New England. In both sexes the dark border of the hind wings is
commonly narrower than it is in individuals from any other region,
though it is not much narrower than in some from the Athabasca
River and in many from New England, and the amount of yellow
scaling on the black and of black scaling on the yellow just above
the orange anal spot is reduced to a minimum as in the more extreme
New England specimens. On the under side of the fore wings the
light tapering marginal band outwardly narrowly edged with black
is usually about as broad as the dark band along its inner border,
but it may be somewhat narrower, and in extreme specimens is
broader. The comma-shaped subcostal mark in the apical portion
of the fore wing is usually the same as in individuals from other
regions, but it may be more extensively black, and even entirely black
with merely a sprinkling of light scales near the outer border and
along the middle of its extension toward the apex of the wing. On
the hind wings beneath, the submarginal lunules are very large and
much straightened, forming broad slightly curved bands which usually
nearly touch each other and are never more than slightly separated.
Although in many specimens the orange spot is no larger than in
many, or perhaps most, specimens from New England, in those with
the dark border most reduced in width the orange centers are much
broadened, the lunules being wholly orange except for a narrow edg-
ing of yellow at the two ends. The spot in the uppermost interspace
on the hind wings is often, though not invariably, reduced to a narrow
straight line as is frequently the case in New England and often in
north-central Canada, or it may be obsolescent. But if present it is
always orange, or at least mostly orange. The dark border of the
hind wings below is margined interiorly by black lines which are
often broader than in specimens from elsewhere, except British
Columbia, though usually they are no broader than in examples from
New England. The lines in the interspaces as far as the vein running
to the tail meet end to end forming a straight line, as is commonly
the case in New England specimens. Just beyond this black line there
is a very heavy light-blue scaling. When the dark border is very
narrow this blue scaling, which is about as broad as the black line
separating it from the yellow of the wing, is separated from the
lunules by a line of black plentifully sprinkled with blue scales nar-

I30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

rower than the blue scaling itself. Thus in specimens with the dark
border of the hind wings very narrow the broad black regularly
linear inner margin is followed by a line of bright blue, slightly or
vaguely interrupted at the veins, of about its own width or somewhat
broader, which is separated from the large and mostly orange lunules
in each interspace by a narrower line of black mixed with lighter.
In many Alaskan specimens, however, the dark border of the hind
wings beneath does not differ from that in New England examples.

Mr. Scudder long ago wrote that specimens from Alaska agree in
coloration and markings with New England specimens in every par-
ticular, although they are slightly smaller than White Mountain
specimens, and these are generally smaller than the average of speci-
mens from other parts of New England, and are apt to be duskier.
This statement is perfectly true for a large proportion of Alaskan
individuals, but extreme Alaskan specimens cannot be matched by
any specimens from elsewhere.

Certain specimens from Alaska are worthy of special mention. In
one from the Yukon River the black borders on both wings are
unusually narrow. In another from Chatanika, Alaska, the submar-
ginal lunules on the hind wings are rather narrowly linear instead of
broad as usual, and the subcostal comma-shaped mark on the under
side of the fore wings near the apex is almost entirely black, the
resemblance to the subspecies rutulus being very close. But the first
submarginal spot on the upper side of the hind wings is orange. We
have seen a similar specimen from Fairbanks. Another from the
Yukon and one from the Porcupine River show a rather close ap-
proach to rutulus. In the revised edition of “The Butterfly Book”
Dr. W. J. Holland figured a female from Alaska with the wing shape
of a male but slightly duller and with conspicuous blue spots some-
what smaller than the very large lunules in each interspace in the
very narrow black border of the hind wings. He did not say where
this specimen came from, and we have seen none like it from central
or eastern Alaska. In the first edition of “The Butterfly Book” he
said that he had numerous specimens from Sitka in southeastern
Alaska, so that this female is probably from that region.

In males at hand from Arrowhead Lake, British Columbia, the
ground color is rather darker than usual, and the black border of
both fore and hind wings is broad. The inner margin of the dark
border of the hind wings beneath is broadly black, and the black
abdominal border, almost always broader in northern than in southern
individuals, is unusually heavy. There is a submarginal light band
on the fore wings beneath. No females are available from this
locality.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK I31

Speaking generally, the small northern males of glaucus on the
whole resemble males from New England very closely, except only
in size. In some regions, as in the White Mountains of New Hamp-
shire and in Newfoundland, the black markings are often somewhat
broadened and have ill-defined borders; in others, as in British Co-
lumbia, they may be unusually heavy ; and in still others they may be
prevailingly narrow and sharply defined, standing out clearly against
the light-yellow ground color, as in central and eastern Alaska, the
Athabaska River region, and most of New England and New York.
On the fore wings the submarginal spots are almost always some-
what narrowed, and in many specimens from central Canada and
Alaska they are narrowly linear ; in most specimens from Newfound-
land, however, they are scarcely narrower than in others from the
south, and not so narrow as in many from New England. On the
under side the fore wing shows a tapering marginal light band out-
wardly edged with black; but in Newfoundland, in southern Ontario,
and in the southern portion of western Canada this band may be
more or less completely broken up into large spots. Inwardly this
band is bordered by a dark band as wide as, or slightly wider than,
itself, though in Alaska occasionally distinctly narrower. The black
band crossing the cell just beyond the middle is usually continued
downward as a solid band, or at least a broad E-shaped mark, to
vein 2 or slightly beyond, sometimes to vein 1; but frequently in
Newfoundland and occasionally elsewhere the band ends at the lower
border of the cell beyond which there are even less distinct traces of
it than in many specimens from the south. The comma-shaped sub-
costal mark near the apex of the fore wings is sometimes reduced in
size, and in specimens from Alaska it is occasionally wholly black
above and almost wholly black beneath.

On the hind wings above, the dark border is usually only very
slightly broader than the dark border of the fore wings. It is nar-
rowest in specimens from Alaska and central Canada, while in many
specimens from Newfoundland it is as broad as in the average
southern individual. The uppermost submarginal spot is reduced in
size and usually linear, most commonly orange or with an orange
center, rarely entirely yellow or absent altogether. The submarginal
lunules are always only slightly curved and are also of more or less
uniform width so that they are short bands rather than lunules; in
specimens from Alaska they are sometimes narrowly linear. The
black abdominal border is usually very broad, occupying more than
half the area between the edge of the wing and the cell and vein 2.
In western specimens it commonly covers all, or nearly all, this area

132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

and broadly overlaps the base of the cell itself, but in specimens from
Newfoundland it is frequently so narrow as to occupy less than half
the area, as in most southern specimens. On the under side the dark
border is narrow, the width from the inner margin to the inner angle
of the serrations, measured along the veins, being about the same
as the width of the dark border of the fore wings in its widest part.
In some Alaskan specimens it is even narrower, so narrow that the
lower inner corner of the enlarged, broadly bandlike, and only
slightly curved lunules as far as the vein running into the tail are
separated from the blue bands bordering the black lines forming
the inner margin of the dark border by a space less than the width
of the black lines themselves. The black lines forming the inner
margin of the dark border in each interspace are straight and, joined
end to end, form a continuous line as far as the vein running into
the tail; this line is usually rather broad and is especially broad in
specimens from British Columbia and in some from Alaska. In cer-
tain specimens from Newfoundland the dark border of the hind
wings beneath may be twice as broad as the dark border of the fore
wings in its widest part, with the included lunules small and having
pointed instead of truncate ends, and bordered interiorly in each in-
terspace by curved instead of straight lines which are more or less
discontinuous. Such specimens resemble those from Maryland and
Virginia more closely, except in size, than they resemble others from
New England and the north. In small northern males the much-
broadened and bandlike lunules have a conspicuous orange spot in
the center, and in many specimens from Alaska the lunules above the
tail are wholly orange except for a narrow yellow margin at each end.
Often in northern specimens the outer point of the lunule just below
the tail is prolonged for some distance down the inner side of the
latter.

The typical northern female exactly resembles the male in size, in
wing shape, and in color, and this type of female occurs throughout
New England and everywhere in Canada and in Alaska, in some re-
gions predominating to the almost complete exclusion of any other
type of female, and in others representing only a minority of the
females present. The females are everywhere more variable than
the males. The amount of variation in the females differs greatly in
different regions. There seems to be least variation among females in
New England, except along the southern and eastern coasts, and in
eastern and central Alaska. In Newfoundland they vary to an extent
greater than anywhere else north of southern New York. Exceed-
ingly small females are of sporadic occurrence everywhere, but
dwarfs are rare among the males.

a

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 133

On August 30, 1906, Lord Rothschild and Dr. Karl Jordan
described Papilio glaucus canadensis from 80 males and 2 females,
giving the range as Newfoundland, Anticosti, New Brunswick,
Canada, the northern districts of British Columbia, and Alaska. The
type locality is Newfoundland. The description covers both sexes.
It was said to be a small form, but no measurements were given. On
the upper side, according to the authors, the third band from the
base of the fore wing reaches nearly always down to vein Mz; in the
male and to SM, in the female. The marginal spots are thinner and
longer than in Papilio glaucus glaucus. The black abdominal border
of the hind wings is broader than the yellow interspace between it and
the cell. On the under side of the fore wings the submarginal spots
form a continuous band, only the last one or two spots being sepa-
rated. The abdominal border of the hind wings is as broad as above.
The submarginal spots on the whole are less curved than in P. g.
glaucus, the blue spots are larger, and the black proximal borders to
them are on the whole more straight.

This description fits perfectly well certain individuals of both
sexes found from Newfoundland to central Alaska and southward
into the mountains of Montana, New York, and New England. Al-
though it applies to individuals of both sexes from Newfoundland, it
describes Newfoundland specimens as a whole less accurately than
it does specimens from any other northern region. On the basis of
the differential characters given, many individuals of both sexes from
Newfoundland would, except that they are small, be unhesitatingly
assigned to P. g. glaucus. For instance, in some females from
Newfoundland the marginal spots above are neither thinner nor
longer than in P. g. glaucus; the black abdominal border of the
hind wings is narrow; the submarginal spots on the under side of
the fore wings are not united into a band; the submarginal lunules
are not less curved than in P. g. glaucus; the blue spots are not
larger; and the black inner border of the dark margin of the hind
wings below is scarcely straighter. Some males depart quite as
widely from the description.

The name canadensis as defined by Rothschild and Jordan does
cover a subspecies in the sense of a definite geographical race. It
covers a form highly characteristic of the north with which almost
everywhere other dwarf forms coexist which, intergrading with it,
depart more or less widely in the direction of P. g. glaucus. It is not
in any way separable from the type characteristic of New England
where, although in Massachusetts reaching a size approximately equal
to that of P. g. glaucus from much farther south, it remains less

134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

variable than in Newfoundland. It is only fair to Rothschild and
Jordan to say that when they described P. g. canadensis they had only
two females, and no specimens from New England at all.

In December 1906 Dr. Henry Skinner described Papilio rutulus
arcticus. He said that this subspecies is smaller than P. r. rutulus,
the distance from the center of the thorax to the tip of the fore
wing being 43 mm. The orange spot at the anterior angle of the
hind wings, generally absent in rutulus, is large and distinct. The
marginal lunules of the hind wings are wider and not so elongate as
in rutulus, and the blue bands on the hind wings below are narrower
and more distinct. On the under side of the hind wings the marginal
lunules are orange, and there is an orange wash running to the cell.
This new form was described from six males and one female. Five
males and the female were from Eagle City, Alaska, in the middle of
the Alaska-Yukon border, taken between June 1 and 15, and one male
was from the Athabasca River. Dr. Skinner indicated no differences
between the sexes in this new form.

Dr. Skinner’s arcticus is not a form of the western rutulus but,
as was shown by Barnes and McDunnough in 1916, is a form of the
eastern and northern glaucus. The name arcticus is applicable to the
form represented by the small Alaskan specimens in which the width
of the dark border of the hind wings below is reduced to a minimum,
and the broadly bandlike slightly curved lunules are orange and are
separated from the broadly black inner margin by conspicuous biue
bands narrowly edged outwardly with blackish. As such individuals
usually fly together with a greater or less number of others which are
less extreme and do not differ from individuals found throughout the
north and as far east as Newfoundland, the name arcticus cannot be
considered as covering a true geographical race or subspecies any
more than can canadensis. But just as the name canadensis is use-
ful in designating the form of the species in which the sexes are
alike, although it may occur with and grade imperceptibly into the
southern form, so arcticus is useful in designating those individuals,
confined to the Northwest, that represent the extreme development
of the canadensis type.

In the light of a knowledge of the various forms in which Papilio
glaucus glaucus occurs in different portions of its extensive range,
the different forms which it assumes within a single limited area—
the District of Columbia and adjacent Virginia—take on an increased
interest.

This butterfly makes its first appearance in this region in late
March or early April—usually in April. All the earliest individuals

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 135

are males. They are always small, with the fore wings from 42 to
45 mm. in length, and they keep almost entirely to the more or less
open woods, coursing along in a fairly direct line 4 or 5 feet above
ground, rarely rising higher, with a rather nervous flight.

These early males have the body clothed with long hair, and there
is a conspicuous tuft of long hair on the front of the head. The
shape of the wings is very characteristic. The fore wings have the
angle between the costal margin (measured as a straight line from
the base of the wing to the apex) and the inner margin more acute
(47°) than it is in the summer form (53°), and the distal portion
of the costal border is less strongly recurved. The apex of the wing
is more pointed, and the straight or slightly convex outer margin
(which is slightly concave in the summer form) makes a greater
angle with the longitudinal axis of the body. The hind wings are
distinctly subtriangular, the outer margin not being markedly con-
vex as is usually the case in the summer form. The outer margin
is deeply scalloped, and the tails are narrow with the outer half only
slightly broadened. The lobe at the anal angle is small, but abruptly
developed and conspicuous.

On the fore wings the dark margin is broadest at the apex and
slowly decreases in width posteriorly. On the upper side the in-
cluded yellow dashes are large. On the under side the spots corre-
sponding to these dashes are fused into a yellowish-white band
tapering posteriorly, only the lowest one or two spots being isolated.
In the upper half of the wing the dark band bordering the light sub-
marginal band interiorly is only slightly broader. The three bands
crossing the cell are equally spaced, and the central one is as nar-
row as, or narrower than, the outer. The outermost band commonly
includes an ill-defined light spot near the lower end, or near both
ends. On the hind wings the black border is very narrow, scarcely
broader than the border of the fore wings, the included lunules are
large, and there is a minimum of light scaling on the black, and
black scaling on the yellow, at its lower end. The uppermost sub-
marginal spot is usually subcircular or oval, but it may be much
narrowed or even linear; it is always orange, usually with a few
yellow scales at the upper and lower ends, and often also along the
outer margin. The lunule next below may have more or fewer
scattered orange scales in the middle. The black abdominal border is
usually about half as broad as the space between the edge of the
wing and the cell and vein M,, but it may be much broader, and
sometimes occupies nearly all this space. On the under side the
black lines forming the inner margin of the dark border in each in-

136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

terspace are straight and usually rather broad; they follow each
other end to end, forming a continuous line as far as the vein run-
ning to the tail (R;). Just beyond these black lines is a series of
bright-blue metallic lines that are rather sharply set off from the
very abundant olive-gray scaling that runs outward as far as the
narrow black inner margin of the lunules. The lunules are very
large and broad with broadly truncated ends—slightly curved broad
bands rather than lunules—and the orange centers are broad and
conspicuous, often occupying more than half the area of the lunules.
The outer point of the lunule just below the tail is prolonged and
more or less extended down the inner side of the latter.

These early-spring individuals from the District of Columbia and
from adjacent Virginia, although very different from summer individ-
uals from the same localities, do not differ appreciably from specimens
taken in the far north on the Athabasca River and in Alaska. They
are usually of the same size, with the fore wings measuring 42-45
mm. in length, and the shape and proportions of the wings are
identical. On the fore wings the black bands may be slightly broader,
especially that across the end of the cell, and the broad E-shaped
mark beneath the middle band across the cell characteristic of
northern examples, though in them of variable development, is
as a rule only partially indicated, though occasionally strongly
developed.

The points of agreement between the early-spring individuals from
the District of Columbia and adjacent Virginia and individuals from
Alaska involve the wing shape, all the major features of the color
pattern, and the hairiness, while the points of difference have to do
only with minor variations in the extent of the development of the
black markings, which are more or less variable in all localities.
Although there is an average difference in these black markings, this
difference is slight, and in all their variations early-spring speci-
mens from the District of Columbia and nearby Virginia may be
matched with others from the far north.

Lord Rothschild and Dr. Jordan wrote that southern spring speci-
mens of Papilio glaucus glaucus somewhat resemble the small north-
ern form which they called canadensis, but are easily distinguishable
by the much narrower black abdominal border of the hind wings.
This supposed difference does not hold good. Although in some indi-
viduals this border is narrow, it is often very broad in early-spring
specimens—indeed it is often quite as broad as in most Alaskan
examples. Early-spring individuals from the District of Columbia
and adjacent Virginia are very close to others from Alaska, and

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 137

some from the two regions are quite indistinguishable. In all there
is a very broad overlapping in all their characters.

Early in spring in this region females are very rare. We have
seen only a single specimen, which resembled the males. All indi-
viduals seen on the wing are yellow, so we have no evidence that
dark females occur. Throughout the north wherever the form of
the male corresponding to the early-spring form in this region is
found the sexes are alike or nearly alike, and dark females do not
occur. This is as true in New England as it is farther north and
northwest. Dark females are, however, produced in New York west
and south of New England, by the dwarf representative of the
southern summer form northeast of New England in Newfound-
land, and in the west as far north as northern British Columbia.

It is impossible to diagnose the dwarf northern representatives of
Papilio glaucus glaucus in such a way as to exclude typical early-
spring individuals from the District of Columbia and northern
Virginia. If their origin were unknown such individuals would un-
hesitatingly be placed with others from the far north, especially
with specimens from Alaska and from the region of the Athabasca
River. Nomenclatorily they would unhesitatingly be referred to
Skinner’s arcticus, being more or less intermediate between arcticus
and canadensis, though rather nearer the former.

Ten days or two weeks after the first appearance of this butter-
fly in the District of Columbia and northern Virginia it rather sud-
denly becomes much commoner. The individuals are noticeably
larger, their flight is less hurried and more devious and often higher,
and they are seen more often in the open, especially in gardens.
These individuals are of a type which is not the same as that to which
the individuals earlier on the wing belonged, nor do they resemble
the summer individuals. Although as a whole the males form an
unbroken series between the early and the typical form, the great
majority are easily distinguishable from both.

The body is usually more or less conspicuously hairy. In wing
form and in color pattern they resemble individuals from southern
New England and New York and westward within the area in which
the species is partially 2-brooded. Flying with the males there are
two types of females, one yellow and one dark blackish brown.

These late-spring individuals are readily distinguished by the
hairy body and the tuft of hair on the front of the head; the com-
plete isolation of the submarginal yellow spots on the under side of
the fore wings; the relatively large size of the yellow dashes on the
upper surface of the fore wings corresponding to these spots, par-

138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ticularly noticeable in the black females; and the relative narrow-
ness, as compared with the summer form, of the dark border on the
under side of the hind wings, which is rather heavily suffused with
light scales.

The dark females (pl. 21, d, e) are usually markedly smaller than
the summer dark females, with larger, sometimes much larger, sub-
marginal spots on the fore wings and lunules on the hind wings, the
submarginal spots on the fore wings below rarely replaced by a
tapering band, and with the marginal region of the hind wings be-
neath somewhat, occasionally considerably, narrower and more
heavily washed with blue and olive-gray scales so as to appear no-
ticeably lighter and inwardly delimited by a continuous series of
curved black lines, one in each interspace. Except that they are
somewhat larger, these late-spring females closely resemble a dark
female at hand from Newfoundland.

Although as a rule spring and summer dark females are at a
glance distinguishable from each other, complete intergradation oc-
curs when the end of the spring brood overlaps the beginning of the
summer brood. Furthermore, late in summer occasional dark females
are sometimes found which, though usually large, resemble more
or less closely the dark females of the late spring form. This form
flies until the end of the spring brood.

The summer brood, developing from eggs laid by both spring
forms, is composed of still larger individuals with the dark border
of the hind wings broader and, on the under side where it is de-
limited internally by a scalloped line, darker; with the submarginal
lunules on the hind wings-and dashes on the fore wings much reduced
in size; and with the dark border of the fore wings on the under
side including a series of separate spots instead of a continuous light
band as in the individuals found early in spring.

The males are rather variable and, taken collectively, form a con-
tinuous series from the late-spring type to a form with the dark
border of the hind wings much broader. They also vary in color
from a light to a darker and somewhat ochreous-yellow, though most
of them are clear bright yellow. In most years there is an interval
of a week or more between the disappearance of the last ragged
individuals of the spring brood and the appearance of the first indi-
viduals of the summer brood. Occasionally, however, the two broods
broadly overlap, and when this occurs there is perfect intergradation
between the late-spring and the summer forms. But it is only in ex-
ceptional cases that these two forms in the District of Columbia and
in northern Virginia cannot be at once distinguished by the rela-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 139

tive hairiness, especially on the front of the head, and the relative
size of the yellow dashes parallel to and near the outer border of
the fore wings.

In the summer form there are seven or more different and fairly
constant, though intergrading, types of females ranging in color from
creamy white to sooty black and varying in the shape of the hind
wings as well as in color.

The interrelationships of the three seasonal forms of the yellow
swallowtail occurring in northern and western Virginia are shown in
the following key.

Key To THE SEASONAL ForMs OF PAPILIO GLAUCUS GLAUCUS OcCURRING
IN VIRGINIA

a1, Body conspicuously hairy; a conspicuous tuft of long hair on the frons;

submarginal yellow dashes on upper side of fore wings thick and rather

long; dark border of hind wings below narrow, interiorly delimited by a

straight or approximately straight line and abundantly suffused with light
scales.

b1. Submarginal yellow spots on the fore wings below fused into a broad

band that tapers posteriorly, only one or two of the lowest spots being

isolated; submarginal lunules on the hind wings below much enlarged,

with their ends broadly truncated; lunule at base of tail with the outer

horn prolonged and running far down the inner side of the tail (flying

As) MASON Gnd. APEH ie) wa 2 bin nada oats cients tea agheens Early-spring form

b2. Submarginal yellow spots on the fore wings below all isolated; submargi-

nal lunules on the hind wings below smaller, crescentic, with pointed

or at least not truncated ends; lunule at base of tail with the outer

horn not produced into the tail (flying from late April to the middle

EAE) 42°F 5c a scrttetd ais, sn Ma eieinetel Che Sea Gas SIE, does Late-spring form

a”. Body not conspicuously hairy, and hair on the frons not elongated; sub-

marginal dashes on upper side of fore wings thin and short, widely sepa-

rated; dark border of hind wings below broader, interiorly delimited by

a strongly scalloped line, and only moderately sprinkled with light scales

(fying from late! June te: October) Wi.io 0s. felis. SS Ss Soh Summer form

Summing all this up, the eastern and northern yellow swallowtail
(Papilio glaucus glaucus) is represented early in spring in the Dis-
trict of Columbia and in northern and western Virginia by far-
northern forms (arcticus or canadensis, or intergrades between the
two), or by a close approximation to them. These are soon joined
by the form found in the northern portion of the United States, and
not long afterward they disappear. As the spring advances the
individuals become, on the average, more and more like those of
the summer form, though seldom quite like them.

The spring brood disappears in June. Late in June or early in
July the summer brood appears. Though variable, the individuals of

140 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

this brood are almost without exception easily distinguishable from
those of the spring brood.

On the Coastal Plain and on the southern and eastern Piedmont
of Virginia the earliest spring individuals, so far as we have seen,
are larger and somewhat deeper yellow than those from farther
north and west, though the color pattern is the same. These re-
semble others from New England, as those from the northern and
western portions of the State resemble those from farther north.

But everywhere the facies of the early-spring individuals is de-
pendent upon meteorological conditions at the time of its first ap-
pearance. The extreme development of the spring form, as in the
case of Battus philenor, appears to occur if the weather is cold,
rather suddenly becoming warm at the time of emergence. If it
becomes gradually warm, with abundant moisture, the extreme spring
form appears to be omitted.

In the summer form in Virginia the males have the fore wings
50 to 60 mm. in length, most of them having the fore wings about
55 mm. or slightly less. Summer females have the fore wings 55 to
69 mm. long, usually slightly less than 60 mm.; individuals with the
fore wings more than 65 mm. long are uncommon.

The summer males are divisible into the usual clear light yellow
males, which are by far the commonest; ochreous males, which are
rather rare; and short-winged males in which the outer and lower
margins of the fore wings are at right angles to each other and the
hind wings are unusually broad. Farther south the color of the
ochreous males deepens into yellow-brown, but such dark individuals
are not recorded from north of South Carolina. The ochreous type
of male occurs as far north at least as eastern Massachusetts, though
here the difference between males of the ochreous and light clear
yellow types is slight.

The females are highly variable; including the spring forms, 11
main color types may be distinguished, as shown in the following
key:

KEy TO THE ForMS OF THE FEMALE OF PAPILIO GLAUCUS GLAUCUS
OccURRING IN VIRGINIA

a1, Predominantly yellow.
b1. Under side of fore wings with a submarginal yellow or whitish band
tapering to a point near the lower angle; little or no blue scaling in
the dark border of the hind wings above (color pattern of the early-
spring: males yaaht end ie oxteies = GRR are oe cee be eee No. I
b?. Under side of fore wings with a submarginal row of spots; dark outer
border of hind wings above with abundant blue scaling.

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK I4!I

c1, Hair on front of head long, forming a conspicuous tuft; submarginal

yellow dashes on the black border of the fore wings above large;

dark border of the under side of the hind wings narrow and heavily

sittused wath dight.scales *(late spring)y.ie4 ss one eerie walsleisie +e No. 2

c2, Hair on front of head short; submarginal yellow dashes in the black

border of the fore wings above smaller and narrower; dark border

on the under side of the hind wings wide and less heavily suffused
with light scales (summer forms).

d1, Yellow areas of the wings thickly and uniformly speckled with black

SGaLEShy sh aead ale sh dieealn wine Qabeerale AMS stared ents BIN SR on eee ae No. 3

d?. Yellow areas of the wings not thickly and uniformly speckled with

black scales.

e1, Region between the base of the wings, the innermost band of the

fore wings, and the narrow stripe on the hind wings heavily

infuscated with sooty brown or black scales, which are also

speckled over the remaining yellow areas; light areas of hind

wings thickly speckled with blue scales (pl. 21, h)........ No. 4

e2. Yellow areas of the wings clear, without dark scaling.

fi. Yellow of wings ochreous, usually darkest on the fore wings

and on the inner half of the hind wings; tails slender, narrow

throughout, or much broadened in the outer half; scallops of

the hind wings deep; process between tail and anal angle long,

with the sides of the black central portion parallel for some

Gistanice’ seleiwe vedane cial). eta aetaye LOR Tatwre feveles Brs.cjid Beebe evened No. 5

f?. Yellow of wings light and clear, creamy; tails broad; scallops

of the hind wings relatively shallow; process between tail and

anal angle short, the sides of the black central portion con-

verging.
gi. Few or no blue scales on the yellow discal area of the hind
WVIMES HINT chsh. dis Selbale Lis daid <a bets Cad» ha leh eae ee ete No. 6

g?. Yellow of the hind wings thickly speckled with metallic blue
scales, appearing opalescent whitish; blue scales also occur
along the inner side of the lower end of the innermost black
Stripe ofthe fore wanes, (pli! 20;) gw sic Neies sieeve oe wh No. 7

a2, Predominantly deep brown or brownish black.
bi. No yellow or whitish scales on the upper surface of the wings other than
those in the submarginal spots.
cl, Submarginal yellow dashes on the fore wings above large; submarginal
lunules on the hind wings large; dark border of the under side of
the hind wings narrow and heavily suffused with light scales (spring).
d1, Under side of fore wings with a tapering submarginal band, only the
two lowest spots separate; submarginal dashes on fore wings and
lunules on hind wings very large and conspicucus; wings narrow

Geaslya Spreiie) 2s Bais ciwbtvine.evs pets ale wyeiaiate Aotdiclae oi eldtdiay fis ea asia No. 8

d?, Under side of fore wings with a row of well-separated spots; sub-
marginal dashes on fore wings and lunules on hind wings smaller ;

wings broader (late spring) (pl. 21, d, €)......cceescucees No. 9

c?. Submarginal yellow dashes on the fore wings above smaller and nar-
rower; submarginal lunules on the hind wings smaller and narrower,
and more curved; dark border of the under side of the hind wings

I42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

broad and less heavily suffused with light scales (summer) (pl. 19 f)

ahaa Sieh ia otayale ere gic Rates ce fene cha ahe lc ANNE UIC RIE et ir) en at No. 10

b2. Wings beyond the position of the innermost black band on the fore
wings and the narrow black stripe on the hind wings with the black
markings standing out prominently against the lighter background
which is more or less heavily dusted with yellow or whitish scales, these
becoming more numerous toward the lower margin of the fore wings

and the upper margin of the hind wings (pl. 21, f, g)........... No. II

No. 1.—Females with the color pattern of the males and with a submarginal
band instead of a row of spots on the under side of the fore wings occur only in
early spring and appear to be very rare; we have seen only a single individual,
caught in the District of Columbia.

No. 2.—The late-spring females are rather variable and grade completely
into the summer females. Although when typically developed they are quite
distinctive, they represent a transition form.

No. 3.—Yellow females with the yellow areas of the wings uniformly speckled
with black scales are rare. We have seen two from Virginia.

No. 4.—Yellow females with the yellow of the inner portions of the wings
heavily infuscated represent a step toward the dark females. Heavy infuscation
of the inner portions of the wings is found in yellow females of both (a) type 5
and (b) type 6. A different step from the yellow to the dark females is seen
in type 7. We have seen no specimens of type 4 from Virginia, but we have
seen specimens of both varieties a and b from Silver Spring, Md.

No. 5.—This is the commonest of the yellow females, and includes more than
half the yellow females found in Virginia. Farther south, so far as we have
seen, all the yellow females are of this type. We have a fine typical example
taken in a bog in Essex, Mass.

No. 6.—The light clear yellow or cream-colored females are, next to type 5,
the commonest yellow type, and become relatively more numerous farther
north. As in type 5, the three black bars on the fore wings are typically evenly
spaced.

No. 7.—This cream-colored or whitish female (pl. 20, g) shows an initial
step toward the dark form in the abundant blue scaling on the upper surface
of the hind wings and also in the approximation of the two outer black bars
on the fore wings. The delicate opalescence of the upper surface of the hind
wings is very striking. This modification of type 6 (a) is duplicated by a cor-
responding variant of type 5 (b). Neither is common.

No. 8.—This type, with the very large submarginal lunules on the hind wings
and submarginal spots on the fore wings, and the wings narrow, is essentially
a black variant of the arcticus-canadensis form. We know of only one specimen,
which was taken in the District of Columbia.

No. 9.—The dark females appearing in late spring (pl. 21, d, ¢) are usually
quite distinctive. They are smaller than summer dark females, though not so
small as type 8, with larger submarginal spots and lunules, and sometimes have
the submarginal spots on the under side of the fore wings united into a band.
They appear to be quite distinct from type 8, but grade more or less completely
into type 10. They are not common.

No. 10.—This is the common black female everywhere. There are various
minor varieties. In one (a) such light scales as may occur on the upper sur-
face, particularly in the form of a bar across the cell near the outer end, are
whitish; in another (b) they are deep yellow.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 143

No. 11.—This dark female, showing a tendency toward the yellow form in
the more or less abundant yellow scaling on the outer half of the wings, is not
uncommon in the vicinity of Washington. There are two main varieties. In
one (a) the light scales are whitish, in the other (b) they are more or less
deep yellow.

The steps in the passage from the male type of coloration to
abundant extreme form of black female are represented by the fol-
lowing sequences: I, 2, 5, 7@ or 4a or 3, 11d, 100; or I, 2, 6, 7b
or 4b or 3, I1a, 10a. No intermediates between the two types, early
and late, of yellow and black spring females, 1 and 8, and 2 and 9,
have been found. Both yellow and black females have corresponding
light and ochreous varieties.

There are two records of black males, neither from the region
under consideration.

The extreme of diversity in this species is found in an area ex-
tending southward along the Piedmont to North Carolina and
through the mountain valleys of western Virginia and West Vir-
ginia. Of the 11 female types, 5 (Nos. 3, 4, 7, 8, and 11) have been
noted only in this region. Here the conditions in Papilio glaucus sug-
gest a parallel with those in Limenitis arthemis from Nova Scotia
to Pennsylvania.

It is tempting to consider the variations of Papilio glaucus in this
area as parallel with those of Graphium marcellus. But the parallelism
is true only of certain aspects of the seasonal sequence of forms. In
G. marcellus there is a linear series of variants from the small early-
spring individuals to the largest summer ones, and the deviation
from this linear series is negligible. In P. glaucus there is great
diversity at all points along the line between two constant, or prac-
tically constant, extremes, as in Limenitis arthemis.

Some of the forms of P. glaucus suggest hybrids between the
northern type with the sexes alike and the southern type. For in-
stance, female forms 8 and 9, which are quite anomalous, have the
pattern of northern females but the color of southern females. It
may be that in this area originally the New England type, or an
approximation to it, predominated in the mountains and on the
Piedmont and that deforestation induced the intrusion of the southern
type with a resultant large proportion of inconstant hybrids.

Occurrence.—Found in and about deciduous woods, especially along
their borders and in glades and clearings, in orchards and brushy
areas, and about isolated groves. It wanders widely over clover fields
and is a common visitor to gardens. This butterfly is everywhere
common in spring. In summer it is much less numerous over most of

144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

the State, though usually common in the mountain regions, especially
in the southwest. In wet summers it is sometimes common on the
Piedmont and on the Coastal Plain as well as in the mountains.

Seasons.—Two broods. The yellow swallowtail appears in the last
half of March or early in April. From the time of its first appearance
this species is usually on the wing continuously until the end of
the season, though in June its numbers decrease and it may disappear
altogether after the middle of the month. The individuals of the
second brood begin to appear in small numbers toward the end of
June or even as early as the middle of the month, becoming common
in July and continuing to emerge into early August, or in excep-
tionally cold and wet summers even into early September. During
September the numbers decrease and after the middle of the month
the much-worn individuals are few in numbers; they disappear en-
tirely in the first or second week in October.

PAPILIO POLYXENES Fabricius

This wide-ranging species is represented in Virginia by—

PAPILIO POLYXENES ASTERIUS Cramer
Plate 22, e, f, g, h

Range.—Throughout the State.

Variation.—In Virginia the males occur in the same three slightly
different intergrading forms that are found in the District of Co-
lumbia (U.S. Nat. Mus. Bull. 157, pp. 193-196, 1932). These three
male forms are the eastern representatives of three much more
widely different forms, asterius, curvifascia, and ampliata, occurring
in the Southwest and in Central America.

It may be recalled that Battus philenor in Virginia has a form
approaching the western hirsuta though less extreme, and another
approaching the southwestern and Mexican acauda. Phoebis sen-
nae also in Virginia has variants suggesting the different forms in
the West.

Dr. E. P. Meiners has pointed out that spring individuals of
P. p. asterius in the vicinity of St. Louis, Mo., where there are three
broods, are somewhat smaller than those of the summer broods,
though not so markedly so as in the case of some of the other swal-
lowtails. He noted that the spots of the yellow bands are somewhat
restricted in size, and there is usually an absence of the spot in the
cell of the hind wings as in the form curvifascia. He said he had
never seen any specimens of this variety in the summer broods. They
occur, though rarely, in the summer broods in Virginia.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 145

In this species the fore wings of the males are usually 40-48 mm.
long ; those of the female 45-53 mm. Giants are occasionally found,
especially among the females. In the United States National Mu-
seum there is a giant female from Dade City, Fla., with the fore
wings 65 mm. long.

Occurrence.—Found in open fields and especially about farms,
frequently visiting gardens; an open-country butterfly, avoiding
wooded regions ; generally and quite uniformly distributed, and com-
mon everywhere; the commonest swallowtail on the Eastern Shore
and on the Northern Neck.

It is probable that originally this butterfly was much less widely
distributed and more localized than it is at present, as are its rela-
tives farther north and in Europe and Asia. The clearing away of
the forests and the introduction of the wild carrot (Daucus carota),
now its favorite wild food plant, presumably are responsible for its
present abundance throughout the eastern States.

Seasons.—Three broods, the third brood incomplete except on the
outer Coastal Plain. The parsnip swallowtail appears usually during
the last week in April and by the second week in May has become
common. It is found continuously until the end of the season early
in October, sometimes as late as the middle of November, and is most
abundant late in July and in August. Over most of Virginia there are
two broods, the second brood putting in its appearance just before the
end of June or early in July. In September the butterflies become
worn and their numbers decrease, but a few fresh individuals repre-
senting a partial third brood, the number of which varies from year
to year, may usually be found until the end of the season. On the
outer Coastal Plain the third brood is almost or quite complete, the
butterflies appearing in considerable numbers from the first to about
the middle of September and flying into October or November.

Genus GRAPHIUM Scopoli
GRAPHIUM MARCELLUS (Cramer)
Plate 19, g, h

Range.—Throughout the State, except the Eastern Shore (Acco-
mack and Northampton Counties).

Variation —The individuals of both sexes flying in early spring
are small with the fore wings usually 32-35 mm. long; the hair on the
body is long, giving the insect a shaggy appearance, and there is a
tuft of long hair on the front of the head; the black bands on the
wings are reduced in width, the light areas correspondingly extended ;

146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

there is usually no distinct pale band along the abdominal fold of the
hind wings; the red anal spot on the hind wings is large and undi-
vided ; and only the tip of the tail is white.

On the outer Coastal Plain of Virginia the size of the early-spring
form increases considerably, the fore wings being up to 40 mm. in
length, though the color remains the same. Farther southward the
dark bands become broader, and finally as broad as in the summer
form, but the tails have only the tips white, and the red anal spot on
the hind wings is large and undivided. This early-spring form with
the broad dark bands (floridensis Holland) ranges northward, ac-
cording to our friend John Boyd, to eastern North Carolina, where
it intergrades with the normally colored but large early-spring form
found along the coast farther north.

The late-spring and summer forms in Virginia resemble those from
elsewhere. In the summer form the fore wings are about 45 mm.
long.

Early in September 1940 we were surprised to find occasional
individuals in the Dismal Swamp and elsewhere on the Coastal
Plain. Although only the tip of the tails was white and the red anal
spot on the hind wings was large as in the spring form, these late-
summer individuals had the dark bands much broader than others
taken in the same region early in spring. They resembled closely the
southeastern spring form floridensis and, were the origin and date of
capture unknown, would certainly be referred to it. Not infrequently
the late-spring form, telamonides, reappears in autumn in small
numbers.

Occurrence-—Found in open deciduous woods in more or less
hilly country, on scrubby hillsides, and in the swamps and pine bar-
rens of the Coastal Plain, often visiting gardens; generally dis-
tributed and common—locally the commonest swallowtail—in the
southwestern portion of the State as far northeastward as Albe-
marle County, more or less local elsewhere, though usually common
where it occurs. The spring brood is more generally distributed and
in most places much more numerous than the later brood or broods.

Seasons.—From one to four or five broods according to locality.
Over most of Virginia there are two broods in the first half of the
season, followed by two or three vestigial broods in the last half. In
most localities only the first is a complete brood, the second varying
from not quite complete to much reduced, and those following being
represented, if at all, only by very few individuals so that the butter-
fly disappears almost completely after the middle of July or the first

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 147

of August. The numbers in the second and following broods vary
greatly in different localities, and in different years.

In the region from Cabin John to Great Falls on the Maryland
side of the Potomac the zebra swallowtail is usually fairly common
early in spring, later occurring in reduced numbers and becoming
very scarce late in summer, though it may be found into early October.
In 1929 and 1930 it was not to be found anywhere in this region after
the first week in May, in those years being represented only by the
first brood; but in 1931 it was more numerous than it had been for
many years, occurring throughout the summer.

In the Dismal Swamp region it is very common from late March
until the middle of July, at the end of June being usually the com-
monest of the swallowtails. After the middle of July its numbers
rapidly decrease and it disappears almost completely by the middle
of August, our last record in a normal year being August 18. Here
there are two full, or nearly full, broods, followed by one vestigial
one.

In the extreme southwestern part of the State and about as far
northeastward as Bland and Pulaski Counties, though especially in
Lee and Scott Counties, there is little, if any, diminution in the num-
bers of this butterfly throughout the summer, and in many localities
in this region it is the commonest summer swallowtail.

The zebra swallowtail appears toward the end of March or early
in April, both sexes at the same time, and soon becomes common.
The second brood begins to appear early in June or even in the last
week in May and flies until after the appearance of the third brood,
about the middle of July. The succeeding broods are on the wing
about the end of August, and in late September and October.

Family HESPERIIDAE
Subfamily PyrcInaE
Genus PROTEIDES Hiibner
PROTEIDES CLARUS (Cramer)
Plate 23, a

Range.—Throughout the State.

Occurrence.—The most generally distributed butterfly in Virginia,
found everywhere and almost everywhere abundant, or at least com-
mon; it is most numerous in scrubby and brushy regions with an
abundance of black locust (Robinia pseudacacia) and least numer-
ous in deep woods and swamps; in certain localities in hilly or moun-

148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

tainous country with open woods and scrub and an abundance of
its food plant it is the commonest butterfly.

Seasons——Two broods with a third incomplete brood the indi-
viduals of which are more numerous on the outer Coastal Plain than
elsewhere. The silver-spotted skipper first appears shortly after
the end of the third week in April, sometimes as early as the nine-
teenth ; slowly increasing in numbers, it becomes common toward the
end of May and abundant by the end of June. It is most abundant
in the last half of July and the first half of August after which its
numbers rather rapidly decrease, though occasional individuals may
be found as late as the first week in October. The individuals of the
first brood continue to emerge until about the middle of June, when
niost of those seen are worn and faded. A little before the middle of
July the first individuals of the second brood appear ; these gradually
increase in numbers, the worn individuals of the first brood becoming
less and less common. The butterflies of the second brood continue to
emerge during most of August. Toward the end of August and in
September a third brood appears that is often more or less complete on
the outer Coastal Plain, but elsewhere is represented only by occa-
sional individuals.

Genus URBANUS Hiibner
URBANUS PROTEUS (Linné)
Plate 23, d

Range.—Recorded only from the Coastal Plain as far north and
west as Richmond. The records are: Henrico County, Richmond,
beneath the Boulevard Bridge, August 1933 (George I. Allen) ; Prin-
cess Anne County, 2 miles west of Lynnhaven, August 19, 20, 1936
(W. H. Wagner, Jr.) ; Virginia Beach, September 24, 1934; Din-
widdie County, Petersburg, 1944 (B. Mather).

Occurrence——Found in open country, particularly in and near
fields of beans ; an uncommon casual visitor, but in some years locally
frequent on the outer Coastal Plain.

Seasons—In Florida there are three broods of the swallowtailed
skipper, but in Virginia and northward only representatives of the
late-summer brood, rarely also of the midsummer brood, have been
reported.

Genus ACHALARUS Scudder
ACHALARUS LYCIADES (Geyer)
Plate 23, e

Range.—Throughout the State.
Occurrence-—Found especially in open deciduous woods and on

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 149

brushy hillsides, also in weedy pastures and in adjacent open country ;
usually infrequent, though in some places fairly common.

Seasons.—One full brood, a second partial brood, and a vestigial
third brood. The frosted skipper usually first appears during the
last week in May, sometimes shortly before the middle of the month,
and slowly increases in numbers, being most numerous in the last
half of June. During the first half of July the numbers fall off,
but about the middle of the month a new brood appears, which
reaches its maximum in the first half of August and disappears about
the end of the third week of that month. Scattered individuals on
the wing late in August and early in September represent a vestigial
third brood.

Genus RHABDOIDES Scudder
RHABDOIDES CELLUS (Boisduval and LeConte)
Plate 23, b, c

Range—Known from Buckingham, Fairfax, Fauquier, Mont-
gomery, Nelson, Page, Prince Edward, and Roanoke Counties.

Occurrence.—Found in or near damp wooded ravines in hilly or
mountainous regions, and most frequently seen along the sides of
streams or near bogs or ponds; very local, the widely separated
colonies consisting usually of few individuals.

Seasons.—One brood and a partial second. The gold-banded
skipper first appears (at Great Falls, Md.) toward the end of May
and flies through June and into the first week in July. A second
partial brood, much less numerous than the first, is on the wing at
the end of the third week in July and flies until after the middle of
August. In Roanoke County we have records for May 3 and 7, 1938.

Genus THORYBES Scudder
THORYBES BATHYLLUS (J. E. Smith)
Plates'235g; h;25;\¢

Range.—Throughout the State.

Variation.—There is considerable variation in the size of the white
markings on the fore wings, especially in the males. In some males
they are so reduced that these might easily be mistaken for males of
T. pylades were it not for the absence of the costal fold.

Recently emerged individuals of this and the related species are
sooty blackish, but they soon fade to dark golden-brown, when old
becoming a medium or even a rather light brown. Some individuals
captured are so worn as to have lost most of the scales on the upper
surface, making them difficult to identify.

I50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Occurrence-—Found in scrub land, open pine woods, and open
fields ; generally distributed and everywhere frequent to common, in
certain localities abundant.

Seasons.—Two broods with, locally, a partial third. The southern
smoky skipper appears shortly after the end of the third week in
April and is on the wing almost continuously until the first week in
October. The first brood, slowly increasing in numbers, becomes
most numerous in June and disappears at about the end of the first
week in July. Shortly after the first of July the second brood appears,
reaching its maximum in the first half of August and flying until
nearly the middle of September. On the Coastal Plain and in the
southern portions of the State a few fresh individuals appear about
the middle of September, flying until about the first of October. The
second brood at its maximum is about twice as numerous as the first,
but the third is represented by very few individuals.

THORYBES PYLADES (Scudder)
Plate 25, a, b

Range.—Throughout the State.

Occurrence.—Found in open fields and open woods, mainly in
hilly or mountainous regions, and generally distributed; frequent to
common in suitable localities on the Piedmont and in the mountains,
but less numerous on the Coastal Plain.

Seasons—One brood and a partial second. The northern smoky
skipper first appears usually about the first of May (rarely as early as
April 23) and slowly increases in numbers, reaching a maximum in,
the first half of June, after which its numbers decline and the first
brood disappears toward the middle of July. Shortly after the first
of July fresh individuals appear, this second brood reaching its
maximum in the last half of July and disappearing before the first
week of August. The individuals of the second brood at its maxi-
mum number only a small fraction of those of the first brood.

THORYBES CONFUSIS Bell
Plate 25, c, d

Range.—Probably throughout the State, but definite records are
few because this species has been confused with T. pylades. Our
records are from Augusta, Caroline, Fairfax, King William, Mid-
diesex, Prince Edward, Prince George, Princess Anne, Southampton,
Spotsylvania, Surry, and Wythe Counties.

Variation.—The white spots on the fore wings, especially in the

INO. 7, BUTTERFLIES OF VIRGINIA—CLARK I51

males, may be greatly reduced and occasionally are wholly absent.
Individuals are sometimes captured in which nearly all the scales
have been rubbed off the upper surface of the wings.

Occurrence.—Found in open fields together with T. bathyllus and
T. pylades, but apparently more local and occurring chiefly on the
Coastal Plain; frequent to common in the regions in which it occurs.
The males frequent the same playgrounds as the males of the other
two species.

Seasons.—One full and one partial brood, with possibly a vestigial
brood late in the season. Bell’s smoky skipper first appears, as
shown by our records, at the end of the third week in May and
becomes common in June, remaining common until the end of the
month when its numbers fall off and it disappears early in the second
week in July. The second brood, in which the individuals are much
less numerous than in the first, appears shortly after the middle of
July and flies until the latter part of August. It is possible that fresh
individuals on the wing late in August and until the last week in
September (September 23) represent a vestigial third brood.

Genus PYRGUS Hiibner
PYRGUS CENTAUREAE (Rambur)

The subspecies represented in Virginia is—

PYRGUS CENTAUREAE WYANDOT (W. H. Edwards)
Plate 24, a, d

Range.—From northern Fairfax County westward to Frederick
County and southwestward along the Blue Ridge and the Alleghenies
to the southern border of the State. Our records are from Augusta,
Fairfax, Frederick, Giles, Highland, Montgomery, Rappahannock,
and Roanoke Counties.

Occurrence.—Found in clearings and open places in woods, and in
open fields near woods; generally distributed, occurring in all suit-
able localities ; frequent to abundant in Frederick County and in the
mountains, infrequent east of Frederick County. It is most com-
monly noticed resting with the wings partially extended on leaves
near the ground, or on the ground itself, or on the flowers of the
common creeping Potentilla (P. canadensis).

Season—One brood. In southwestern Virginia the grizzled darter
first appears at about the end of March, but in the north not until
after the middle of April, at the higher altitudes at the end of April
or early in May. The date of its first appearance varies considerably

152 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

from year to year. It is common for about 10 days or 2 weeks after
which its numbers rapidly decline and the males disappear, the fe-
males continuing to fly until about the end of the third week.

PYRGUS COMMUNIS (Grote)
Plate 24, b, c, e, f

Range.—Throughout the State.

V ariation.—Speaking of this species as it occurs in Kansas, Wil-
liam D. Field says that the dark form is common in spring and early
in summer and that later in summer almost all individuals are of the
whiter form.

In Virginia the males have the fore wings 13-15 mm. long, the
females averaging slightly larger. The amount of white on the upper
surface is variable. The males usually have more white than the
females, with a more definite and continuous postmedian line. Occa-
sionally in the males the postmedian line is broadened and quite con-
tinuous, not being interrupted at the veins. The females are some-
times very dark with the white markings above much reduced in size.
The ground color above, especially in the males, may be light and
grayish.

The markings on the under side of the hind wings are usually
light to medium yellow-brown, with some dark brown or blackish,
and the spots very distinct. But not infrequently, especially late in
summer, they are faint and more or less indistinct, the under side of
the hind wings presenting a washed-out appearance as in the western
form albescens Ploetz. Specimens caught in the winter in southern
Maryland just across the Potomac from Virginia have the spots on
the under side of the hind wings black shading into dark gray with
no trace of yellowish.

Occurrence——Found in open fields and along roadsides, and es-
pecially in and near farmyards where the common mallow or cheeses
(Malva rotundifolia) grows; generally distributed and about equally
numerous at all points; usually infrequent to frequent, but locally
common.

Seasons.—Three broods, the last incomplete. This darter usually
first appears in the first half of May, rarely as early as April 24,
and increases in numbers until the last half of July and the first
half of August, after which the numbers gradually fall off and the
butterfly disappears early in October, occasional individuals persist-
ing, in some years, until nearly the middle of November, rarely until
the last week in December. The first individuals of the second brood

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 153

appear toward the end of June, and those of the third brood toward
the end of August, broadly overlapping the later individuals of the
second brood.

Genus PHOLISORA Scudder
PHOLISORA CATULLUS (Fabricius)
Plate 26, r

Range.—Throughout the State.

Occurrence.—Found occasionally to frequently in open fields and
along roadsides, and commonly or even abundantly in and about
cultivated areas, particularly truck farms and gardens, and in unkept
town and city lots where its chief food plant (Chenopodium album)
thrives. The caterpillars are common on this plant everywhere and
the presence of the insect often may be determined from these when
no adults are in evidence.

Seasons——Two broods and a partial third. The pigweed skipper
first appears usually toward the end of the first week in May, on the
Coastal Plain toward the end of April, and flies continuously until
about the end of the second or third week in September, reaching its
maximum abundance in the last half of July and the first half of
August. In the first half of June the numbers decline, but toward
the end of the month the second brood appears, and the butterfly
becomes increasingly common, reaching its peak late in July. Late
in August and in September the appearance of numerous fresh indi-
viduals indicates a third brood in which, however, the numbers do not
reach half of those of the midsummer brood.

PHOLISORA HAYHURSTII (W. H. Edwards)
Plate 15, ¢, j

Range.—Confined to the Coastal Plain. Our records are: Caro-
line County, between Port Royal Forks and Windsor, May 29, 1937;
Isle of Wight County, Rescue, September 4, 1940; Nansemond
County, western border of the Dismal Swamp about 2 miles south
of Suffolk, September 4, 1936, Dismal Swamp, May 14, 1945 (Otto
Buchholz), Holland, July 13-26, 1944 (Otto Buchholz) ; Norfolk
County, Wallaceton, July 6, 1941.

Occurrence-—Found in and near more or less wet open pine
woods ; very local and rare.

Seasons.—Our records indicate three broods, one in the last half
of May, one in July, and the third in September.

154 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Genus ERYNNIS Schrank
ERYNNIS ICELUS (Scudder and Burgess)
Plate 24, g, h

Range.—Northern and western portions of the State and locally
eastward and southward to Amelia, Henrico, New Kent, and West-
moreland Counties. Our records are from Albemarle, Amelia, Bed-
ford, Fairfax, Frederick, Henrico, Madison, Nelson, New Kent,
Page, Patrick, Pulaski, Roanoke, Rockbridge, Shenandoah, Smyth,
Tazewell, Warren, and Westmoreland Counties.

V ariation.—The fore wings in this species are usually about 15-16
mm. long. There is considerable variation in the amount of grayish
white on the outer half of the fore wings, the light area in the outer
third being much more marked in some individuals than in others. It
is usually prominent in blackish individuals fresh from the pupa,
becoming more or less inconspicuous in old, worn, faded, brownish
ones.

In Burkes Garden, Tazewell County, on June 4, 1940, John E.
Graf captured a male with the wings rather light bronzy in color, the
fore wings being faintly clouded and almost immaculate.

Occurrence.—Found in hilly and mountainous country in rich open
woods, along the borders of more or less damp woods, along roads
through woods, and widely dispersed over fields near woods; com-
mon to abundant in all suitable localities from Fairfax to Frederick
and Page Counties and southwestward to the southern border of the
State, becoming increasingly local and infrequent farther south and
east. In some places, notably on the Peaks of Otter and in certain
localities in Fairfax, Frederick, and Page Counties, we have found it
in its season the most abundant butterfly.

Season.—One brood; a few individuals taken in Bedford, Patrick,
Pulaski, and Smyth Counties between July 22 and August 4 indicate
the occurrence of a partial second brood in the southwestern moun-
tain region. This butterfly usually appears about the middle of April,
sometimes as early as the gth, later in the higher altitudes, and reaches
the peak of its abundance in the first half of May in the lower alti-
tudes, in the last half of May in the higher. It remains abundant for
about 10 days or 2 weeks, after which its numbers fall off and it
usually disappears in the first week in June, though occasional indi-
viduals may be found up to nearly the end of the month. In the
southwestern mountains it reappears in small numbers in the last
week in July and the first week in August.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 155

ERYNNIS BRIZO (Boisduval and LeConte)
Plate 24, 1

Range.—Throughout the State. We have records from Albemarle,
Amelia, Bedford, Buckingham, Clarke, Frederick, Giles, Henrico,
James City, Loudon, Montgomery, Nansemond, Nelson, Page, Pow-
hatan, Prince Edward, Roanoke, Surry, and Warren Counties.

Variation—tThe fore wings are usually about 18 mm. long. At
Williamsburg on April 16, 1940, we captured a specimen which was
a rather light bronze with the fore wings uniformly colored except
for a few small black spots.

Occurrence-—Found in open woods, especially along the roads and
paths and along the borders of adjacent fields; it is abundant and
generally distributed in the woods of Frederick County, and common
in the mountains as far, at least, as Montgomery County; east of
the mountains it is locally distributed, though common wherever it
occurs. At Williamsburg on April 16, 1940, we found it much the
commonest species of the genus.

Season.—One brood. In the lower sections of Virginia this species
usually appears about the first of April, sometimes late in March,
but at the higher elevations its appearance is delayed until the latter
part of April. It becomes common about a week after its first appear-
ance, or shortly after the first females are on the wing, and remains
common for about 2 weeks, after which the number of the males
declines and later that of the females, until only a few very worn
females are left. It flies until near the middle of May in the low-
lands, and until about the first of June in the highest elevations.

ERYNNIS LUCILIUS (Scudder and Burgess)
Plate 24, k

Range.—Higher mountain regions in the western part of the State;
our records are from Bedford, Highland, Page, Roanoke, and Smyth
Counties.

Occurrence—Found in usually rocky woodlands and in adjacent
fields in hilly or mountainous country ; uncommon.

Seasons——Two broods. The records from April 23 to June 12
seem to represent the first brood, those from July 22 to August 17
the second.

ERYNNIS PERSIUS (Scudder)
Plate 24, 7, 1

Range.—Mountains in the western part of the State and the Poto-
mac valley. Our records are from Bedford, Fairfax, Highland, Roa-

150 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

noke, and Tazewell Counties; most of them are from Highland
County.

Occurrence.—Found in open fields, usually near woods, in boggy
places, and along roads near woods; it appears to be rather common
in Highland County, uncommon elsewhere.

Season.—One brood. This species appears shortly after the end of
the third week in April (April 23) and flies until after the middle of
June (June 17); it is most numerous from the last of May to the
middle of June.

ERYNNIS BAPTISIAE Forbes
Plate 24, m

Range.—Throughout the State. Our records are from Accomack,
Alleghany, Arlington, Augusta, Bedford, Caroline, Fairfax, Giles,
Isle of Wight, King William, Madison, Montgomery, Page, Roanoke,
Rockbridge, Rockingham, Shenandoah, Smyth, Southampton, and
Surry Counties.

Occurrence.—This recently described species, the range of which
is as yet very imperfectly known, is found in open fields more or
less near woods and in very open woods, especially in grassy areas
and along roads. It is the commonest of the smaller species of the
genus with the exception of the less generally distributed E. icelus
with which, in life, it is easily confused.

Seasons.—Two broods. Although we have no records before the
first of May, this species probably first appears shortly after the
end of the third week in April, the first brood flying through May
and into the first week in June. The second brood appears during
the first week in July, reaches its maximum in the last half of the
month, and disappears early in September.

ERYNNIS MARTIALIS (Scudder)
Plate 24, n

Range.—Mountains in the western portion of the State and locally
eastward to Prince Edward, Henrico, and Fairfax Counties. Our
records are from Alleghany, Augusta, Bedford, Fairfax, Giles, Hen-
rico, Highland, Montgomery, Page, Prince Edward, Roanoke, Rus-
sell, Warren, and Wythe Counties.

Occurrence.—Found in roads and clearings in woods and in fields
near woods in hilly or mountainous regions; local and usually infre-
quent, though in some places fairly common.

Seasons.—Two broods. This butterfly first appears usually in the

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 157

last half of April, sometimes as early as April 11, and flies until the
end of the third week in June. The second brood appears early in
July and flies until just beyond the middle of August. Freshly
emerged. individuals, which are very dark, are easily confused in
life with E. baptisiae.

ERYNNIS JUVENALIS (Fabricius)
Plates 24, 0, p; 25, f

Range.—Throughout the State.

Variation —This species is usually easily distinguished by the pres-
ence of two small circular white or whitish spots on the under side
of the hind wings near the outer angle, but one or even both of these
spots may be absent. There is considerable variation in color, some
individuals having the fore wings much grayer than others, with the
dark markings more distinct, and the dark markings vary consider-
ably in size and in extent. There is frequently some difference in
the markings of the fore wings on the two sides. The size of the
hyaline spots varies, and one or more may be lacking. Occasional
females are very light, with the ground color above light sandy, be-
coming dirty whitish on the outer half of the fore wings.

Occurrence.—Found in open woods, especially along the roads and
paths, and in adjacent open fields; generally distributed and every-
where common to abundant.

Seasons.—One full brood, a partial second brood, mainly on the
Coastal Plain and the outer Piedmont, and a sporadic third brood
represented by occasional individuals on the wing late in autumn in
certain years. This species first appears about the first of April, some-
times late in March, and by the middle of April has become abundant
over most of the State. On the Coastal Plain it is commonly abun-
dant by the end of the first week in April, but in the higher moun-
tain regions it does not usually become abundant until toward the
end of April. After the first of May, or the middle of May at the
higher altitudes, the individuals are worn and their numbers decrease,
but a few are still to be found until the end of the third week in June
or even the first week in July. About the middle of July fresh indi-
viduals appear and the butterfly may be found in small numbers until
toward the end of August. Occasional individuals on the wing in
October probably represent a vestigial third brood. In some years we
have seen this species rather frequent in October at Cabin John, Md.,
just across the Potomac from Fairfax County.

158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

ERYNNIS HORATIUS (Scudder and Burgess)
Plate 25, 7, k

Range.—Throughout the State.

V ariation.—Occasional females of this species are light sandy in
color with the outer half of the fore wings dirty whitish.

Occurrence.—Found in open fields and open woods; generally
distributed and frequent to common everywhere.

Seasons.—Two broods and a vestigial third brood. This species
first appears about the first of April and slowly increases in numbers,
reaching a maximum in the first half of May, the second half at the
higher altitudes, the numbers then declining. In July a new brood ap-
pears which becomes abundant in the last half of the month and
remains numerous through the first of August, continuing on the
wing, in diminishing numbers, through September. Fresh individuals
on the wing in late September and October probably represent a
vestigial third brood. In this species the second brood appears to
be more nearly complete than the second brood of E. juvenalis. In
the spring E. horatius, though common, is far outnumbered by £.
juvenalis, but in midsummer and later we have found E. horatius
much more numerous than E. juvenalis. We are not prepared to
say, however, that these relative seasonal proportions would be main-
tained over a long series of years.

ERYNNIS ZARUCCO (Lucas)
Plate 25, g, h

Range.—Throughout the Coastal Plain, and farther inland north
to Prince Edward, Pittsylvania, Roanoke, Montgomery, Giles, and
Augusta Counties. Our records are from Accomack, Augusta, Giles,
Grayson, Isle of Wight, Mathews, Montgomery, Nansemond, Nor-
folk, Northampton, Pittsylvania, Prince Edward, Prince George,
Princess Anne, Pulaski, Roanoke, and Surry Counties.

Occurrence—Found in open country, usually near the borders
of woods.

Seasons.—Two broods. This species first appears in April, reaches
a maximum in the first half of June, and flies until the end of the
month. In July a new brood appears, and the butterfly is most com-
mon in July and the first half of August, after which time the num-
bers decrease and it disappears after the first week in September.

The very few records for April (April 15, Salem) and May (May
19, Farmville; May 30, Dismal Swamp) seem to indicate that Vir-
ginia is the northern limit of the range, and that mortality during the

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 159

winter is very heavy. It is probable that this species is for the most
part an immigrant from farther south entering the State each year,
a few individuals occasionally surviving the winter. However, it may
be more common in spring than the records indicate, for in the field
it is scarcely distinguishable from the abundant EF. juvenalis and the
common E. horatius. In the summer when the numbers of E. juve-
nalis are greatly reduced and all the individuals of the larger species
of Erynnis are captured and examined it is more likely to be detected.

Subfamily HEsPERIINAE
Genus ANCYLOXYPHA Felder
ANCYLOXYPHA NUMITOR (Fabricius)
Plate 3, k

Range.—Throughout the State.

Variation.—In this species the fore wings are usually about 11 mm.
long, but occasional individuals are much larger. At Rescue, Isle of
Wight County, on September 3, 1940, we captured one with the fore
wings 15 mm. long. Not infrequently the fore wings are wholly
black (form longleyi French). We found this form rather common
at Rescue on September 3, 1940, and at Sunken Meadows Beach,
Surry County, on the following day. A specimen from the Dismal
Swamp taken on September 1, 1940, has the fore wings above light
bronzy, slightly dusky over the normally black areas; the under
surface is a strongly opalescent dull straw yellow.

Occurrence.—Found in wet grassy regions and along the grassy
borders of sluggish streams, ponds, and roadside ditches; generally
distributed and common everywhere. In all wet grassy areas not
subject to too extensive spring flooding or to too severe drying late
in summer this little skipper is certain to occur in more or less
abundance.

Seasons—Three broods. The least skipper first appears just after
the middle of May (earliest date May 17) and by the last week in
the month has become common. The first brood is most numerous
in the first half of June, its numbers falling off rapidly in the last
half of the month. Toward the end of June a new brood appears
which reaches its maximum in the last half of July and the first half
of August, when this little butterfly is at the peak of its abundance.
In the last half of August the numbers decline, but toward the end
of August they are reinforced by the appearance of a third brood,
which remains on the wing until shortly after the first of October, or
even until the second week. The second and third broods spread out

160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

over a considerably larger area than the first brood, especially if the
summer be wet. An extraordinary record we cannot explain is
furnished by a specimen captured at Richmond on March 30, 1936, by
Dr. Carroll M. Williams.

Genus HESPERIA Fabricius
HESPERIA METEA (Scudder)
Plate 28, a, b

Range.—Known from Fairfax, Frederick, Henrico, Prince Ed-
ward, Roanoke, and Montgomery Counties; but undoubtedly much
more generally distributed in the western part of the State than these
records would indicate.

V ariation.—Although the distinctness of the light markings on the
under side of the hind wings is fairly constant in the males, it varies
considerably in the females. In some females these light markings
are represented merely by indistinct vestiges, and we have seen one
female from Rocky Run, Fairfax County, in which they appeared to
be wholly absent.

Occurrence.—F ound in open brushy clearings in dry woods, usually
with pines, and in dry open fields near woods, in hilly or mountainous
country ; very local, but frequent to common wherever it occurs.

Season.—One brood. This skipper usually appears in the middle
of April, sometimes in the first week in the month, and exceptionally
as early as March 12, and flies until the last week in May (latest
date May 26). It is usually most numerous in the first half of May,
in some years in the last week in April and the first week in May.

HESPERIA LEONARDUS Harris
Plate 26, a, b

Range—From Arlington and Fairfax Counties westward and
southwestward in the mountains to Giles, Montgomery, and Patrick
Counties. Our records are from Arlington, Fairfax, Giles, Madison,
Montgomery, and Patrick Counties.

Occurrence.—Found in permanently wet meadows and bogs; gen-
erally distributed, though not common, in Arlington and Fairfax
Counties, very local and rather rare in the western part of the State.

Season.—One brood. Our few records run from August 15 (Pov-
erty Hollow, Montgomery County) to September 22 (Rocky Run,
Fairfax County). In the vicinity of Washington Leonard’s skipper
appears about the first of September, reaches its maximum about the
middle of the month, and usually disappears before the first of

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 161

October. In the mountains it appears at least as early as the middle
of August and presumably reaches its peak of abundance in the last
week in August, as it does in eastern Massachusetts.

HESPERIA SASSACUS Harris
Plate 27, m, p

Range.—Confined to the mountains in the western portion of the
State from Frederick, Warren, and Page Counties southwestward.
Our records are from Frederick, Highland, Montgomery, Page,
Roanoke, Tazewell, and Warren Counties.

Variation—Prof. A. W. Lindsey writes (Denison Univ. Bull.,
Journ. Sci. Lab., vol. 37, p. 48, April 1942) that the darkest speci-
men of the form manitoboides Fletcher he has is from Frederick
County, Va. This was one of a series of 34 specimens taken at
Chambersville on May 28, 1939, that we sent him in connection with
his revision of the genus Hesperia.

Occurrence.—Found in open fields in or near woods in hilly or
mountainous country ; generally distributed in Frederick, Warren, and
Page Counties, becoming increasingly localized southwestward. This
species varies greatly in abundance from year to year. In the
northern counties it is usually the commonest butterfly during its
flight period.

Season.—One brood. The Indian skipper appears early in May
(May 6) and flies until the middle of June (June 19). It is abun-
dant in the last week in May and early in June.

HESPERIA ATTALUS (W. H. Edwards)
Plate 27, i

Range.—Two records: Loudoun County, Clark’s Gap, September
22, 1940 (A. H. Clark); Prince Edward County, Farmville, June
9, 1940 (Frank W. Trainer).

Variation.—Both specimens, now in the United States National
Museum, are females. They are quite alike and represent exception-
ally dark examples of the form seminole Scudder. They are the
darkest specimens of the species we have seen.

Genus HYLEPHILA Billberg
HYLEPHILA PHYLEUS (Drury)
Plate 28, c

Range.—Throughout the State.
Variation ——The females of this species occur in three forms: (1)

162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

The usual form, which is by far the commonest; (2) a light form
with the light markings above greatly extended, which is rare; and
(3) a dark form, very dark with the light markings above much
reduced and the under side uniformly dark with an angular band of
light spots and a light dash extending inward from the angle along
the cell toward the base of the hind wings. This very dark form,
which is not uncommon in Virginia, is sometimes puzzling as it
suggests in its color pattern a species of Hesperia. Intergrades occur
between these three forms.

Occurrence-—Found in fields, along roadsides, and very com-
monly in gardens.

Although our records are far too few to be regarded as giving
more than an indication, a fair inference from them is that this
species is only a summer visitor to Virginia. The only captures that
could be referred to a spring brood are one on May 31, 1936, at
Creeds on the North Carolina line in Princess Anne County, and
another on June 11, 1938, in the Dismal Swamp in Nansemond
County. These records would seem to indicate that in certain years
this butterfly may overwinter in the extreme southeastern portion
of the State.

We have no other records before July when, in the first half of the
month, we have this species from Henrico, King and Queen, Nanse-
mond, Norfolk, Northumberland, Princess Anne, and Surry Coun-
ties, and in the last half from Accomack, Giles, Northampton, Prince
Edward, Pulaski, and Roanoke Counties. In the first half of August
we have found it in Prince William and Nelson Counties, and also in
the District of Columbia.

It would appear that, in most years, this species first enters Vir-
ginia early in July, coming north along the Coastal Plain, in the latter
half of the month spreading widely over the Coastal Plain and also
entering the State in the southwestern and south-central sections.
By the end of the season it is usually abundant throughout the
Coastal Plain and common in the southern half of the State, becoming
less numerous in the northern Piedmont and mountain counties.

The numbers of this butterfly fluctuate considerably from year
to year, and not infrequently it is wholly absent from the northern
Piedmont and mountain counties, and scarce elsewhere except on the
outer Coastal Plain where it is always to be found in greater or less
numbers, and is normally, with the possible exception of Atalopedes
campestris, the commonest of the skippers in the last half of the
summer.

Seasons.—Three broods. The first brood, flying in May and early

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 163

June, is usually not represented in Virginia, though we have two
records, for May 31 and June 11, the latter based upon a worn
female. The second brood appears early in July and flies through
July and August. Apparently there is a third brood that first appears
late in August and continues to emerge until the end of the season,
usually about the end of October, sometimes as late as the middle of
November (November 12). The butterfly is on the wing continu-
ously from about the first of July until the end of the season.

Genus ATALOPEDES Scudder
ATALOPEDES CAMPESTRIS (Boisduval)
Plate 26, f, g, h,i

Range.—Throughout the State.

Variation.—In Virginia the fore wings of the males are 14-15 mm.
long ; those of the females 17-18 mm. There appears to be but little
variation. Occasional females are much lighter and more yellowish
than the average, and some are much darker (pl. 26, f).

Occurrence——Found in open fields, along weedy roadsides, and
in gardens. Very scarce in spring, the sachem appears in numbers
early in July, and from the middle of July onward it is everywhere
abundant except in the highest mountain regions, where it is fre-
quent or sometimes common. Late in summer and in autumn it is in
almost all sections the commonest of the skippers.

Virginia is approximately the northern limit of this species as a
permanent resident. Although we have spring records from 13
counties situated in all sections of the State and including Frederick
County in the extreme north, and also from the District of Columbia
and from Montgomery County, Md., across the Potomac from Fair-
fax County, these records are very few, indicating that only a small
number of individuals succeed in surviving the winter. So scarce is
this insect in spring that we believe the large population in mid-
summer and later is mainly the result of immigration from farther
south.

Seasons.—Three broods. This species first appears in the first
half of May (May 5) and increases in numbers until the last week
in the month, after which the numbers slowly fall off and it disappears
shortly before the middle of June. The males appear a week or so
in advance of the females. The second brood appears about the first
of July, and the butterfly attains its maximum abundance in the last
half of this month and the first half of August. During the last half
of August the numbers decrease, but toward the end of the month

164 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

they are augmented by the appearance of a third brood which is on
the wing until late in October or even until nearly the middle of
November (November 12).

Genus POLITES Scudder
POLITES VERNA (W. H. Edwards)
Plate 28, d

Range.—Throughout the State. Our records are from Accomack,
Albemarle, Bland, Fairfax, Fauquier, Frederick, Grayson, Hanover,
Henrico, Highland, Loudoun, Madison, Montgomery, Nansemond,
Prince Edward, Prince William, Princess Anne, Rockingham, Scott,
Shenandoah, Smyth, Tazewell, Washington, and Westmoreland
Counties.

Occurrence.—Found in damp or wet open woods, brushy bogs, and
wet pastures near woods; very local; in the mountains and on the
northern Piedmont occurring in most suitable situations, usually in
small numbers, though in some places common, especially in and near
bogs ; rare on the Coastal Plain.

Seasons.—Two broods. This skipper appears at about the end of
May and slowly increases in numbers, becoming common in the last
half of June and in the first half of July, after which the numbers fall
off and it becomes scarce or disappears toward the end of the month.
At the end of July fresh individuals appear, and the butterfly is
again common in August, flying until about the end of the first week
in September, or even later (September 22).

POLITES MANATAAQUA (Scudder)
Plates 15, d; 28, e

Range.—Throughout the State.

V ariation—Occasional males of this species entirely lack the
fulvous on the upper surface, being plain dark brown with a black
stigma, like the males of Atrytone ruricola. Although we have not
found this variety in Virginia, we have seen one specimen from the
District of Columbia and another from nearby Maryland.

In Stafford County late in the season we captured a male that was
pale yellowish bronze with silky reflections above, the fulvous patches
being scarcely distinguishable from the ground color.

Occurrence-—Found in open fields and along weedy roadsides in
open country; one of the most generally distributed of the skippers,
though nowhere abundant and seldom very common, and in some
years infrequent. On the highest mountain pastures this species is

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 165

especially noticeable, not because it is common, but because it is
often the only skipper to be found.

Seasons—Two broods. This species first appears after the middle
of May (earliest date May 18) and becomes common toward the end
of the month and in early June, disappearing at the end of June. The
second brood is on the wing shortly before the middle of July
(July 10), reaches its maximum in August, and continues on the
wing until the end of the season, toward the end of September. It is
possible that fresh individuals found late in September may repre-
sent a partial third brood.

POLITES THEMISTOCLES (Latreille)
Plate 28, f

Range.—Throughout the State.

Variation—We have not found any appreciable variation in the
males of this species in Virginia. The females may resemble those
of P. manataaqua except in being smaller and with the spots on the

fore wings more strongly yellow, but usually there is more or less

fulvous along the costal border of the fore wings and in the cell,
not infrequently quite as much as in the males. A female from Vir-
ginia Beach taken on September 24, 1934, is very dark, with no ful-
vous along the costal border of the fore wings and with the spots
greatly reduced in size.

Occurrence.—Found in fields and along roadsides, everywhere one
of the commoner skippers.

Seasons——Two broods. This species first appears usually in the
last half of May (rarely as early as May 7), becoming common to-
ward the end of the month and through the first 3 weeks of June,
the numbers then decreasing, though occasional individuals are to
be found as late as the end of the first week in July. About the mid-
dle of July a new brood appears, and the butterfly increases in num-
bers, reaching a maximum in the first half of August and continuing
on the wing in diminishing numbers until the end of the season. It
is possible that fresh individuals taken late in September may repre-
sent a vestigial third brood.

POLITES PECKIUS (Kirby)
Plate 28, g, h
Range.—Northern and western portions of the State, as far east

as Westmoreland, Essex, Prince Edward, and northern Pittsylvania
Counties. Our records are from Albemarle, Alleghany, Amherst,

166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

Arlington, Augusta, Bath, Campbell, Clarke, Essex, Fairfax, Fau-
quier, Floyd, Frederick, Giles, Highland, King George, Loudoun,
Madison, Montgomery, Nelson, Orange, Page, Patrick, Pittsylvania,
Prince William, Pulaski, Rappahannock, Roanoke, Rockbridge, Rock-
ingham, Russell, Scott, Shenandoah, Smyth, Spotsylvania, Stafford,
Tazewell, Warren, Washington, Westmoreland, and Wythe Counties.

Occurrence.—In the western mountains and in the northern coun-
ties this is one of the commonest and most generally distributed
skippers, found in greater or less numbers in every open field, most
abundantly in the valleys and in mountain bogs and damp pastures.
Farther east and south it rapidly becomes increasingly localized, oc-
curring in colonies in low damp areas, or near water.

Seasons —Two broods. Peck’s skipper usually appears about the
end of the third week in May and by the end of the month has be-
come common, remaining common through early June and disap-
pearing about the middle of the month. The second brood appears
in the third week in July, reaches its maximum in August, and flies
in diminishing numbers until the end of the season, about the end of
the third week in September, though sometimes much later. It is
possible that individuals on the wing late in September and early in
October, seen in certain years, represent an occasional partial third
brood. The broods of this species, confined to the cooler regions of
the State, appear to be more definitely marked than those of its more
generally distributed relatives P. verna, P. manataaqua, and P.
themistocles.

POLITES MYSTIC (Scudder)
Plate 26, j, k

Range.—Known only from Highland, Giles, Montgomery, and
Roanoke Counties. The records are: Highland County, Middle
Mountain, June 19, 1948 (common) ; Giles County, Mountain Lake,
July 21, 1940 (2 females, Lloyd G. Carr), Rich Creek, August 1, 1938
(2 females) ; Montgomery County, Old Preston graveyard, Blacks-
burg, June 2, 6, 1899 (22 males and females, Ellison A. Smyth, Jr.) ;
Roanoke County, Ash Bottom, near Salem, June 1, 1937 (Carl W.
Gottschalk, specimens identified by Ellison A. Smyth, Jr.).

Occurrence.—Found in fields near woods. We have met with this
species personally only twice, along the edge of the woods on a hill-
side near Rich Creek, Giles County, on August 1, 1938, and on the
crest of Middle Mountain, Highland County, on June 19, 1948, where
we found it common, flying with Hesperia sassacus.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 167

POLITES VIBEX (Geyer)
Plates 27, f; 28, 7

Range.—One record, Montgomery County, near Blacksburg (2
males taken by Prof. Ellison A. Smyth, Jr., probably about 1900).
Occurrence-—Probably a casual or irregular visitor.

Genus WALLENGRENIA Berg
WALLENGRENIA OTHO (J. E. Smith)

Two subspecies are represented in Virginia.

WALLENGRENIA OTHO OTHO (J. E. Smith)
Plate 26, o

Diagnostic features—The under side of the hind wings is dull
orange or orange-red.

Range.—North to the south-central part of the State and along the
coast to Gloucester and Mathews Counties. The records are: Glouces-
ter County, Bellamy, September 1, 1941, Severn, August 31, 1941;
Mathews County, Foster, August 30, 1941; Middlesex County, Ur-
banna, September 2, 1941; Nansemond County, Dismal Swamp,
1941, June 2, 5, 1944 (Otto Buchholz), May 20, 1945 (Otto Buch-
holz) ; Prince Edward County, Farmville, September 11, 1940 (Frank
W. Trainer).

Occurrence——We have found this form only in swamps or in
more or less wet localities.

There is something of a mystery about the occurrence of this
form in Virginia. There is no record before 1940, when Frank W.
Trainer took it at Farmville. In 1941 we found it common in
Gloucester and Mathews Counties, and received it from the Dismal
Swamp where previously we had taken only the form egeremet, which
is rather common there. Our friend Otto Buchholz found it in the
Dismal Swamp in 1944 and 1945. It does not seem possible that this
well-marked form could have been overlooked in previous years.

WALLENGRENIA OTHO EGEREMET (Scudder)

Plate 26, m,n

Diagnostic features—The under side of the hind wings is dark
purplish brown.

Range.—Throughout the State.

Occurrence.—Found in fields and meadows, along roadsides, and
in gardens, but avoiding the driest localities; everywhere frequent,
though never very numerous.

168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IIO

Seasons.—Two broods. This species first appears toward the end
of May, sometimes as early as the end of the third week, and flies
until about the end of June, worn individuals continuing on the wing
into July. The second brood appears in the second week in July and
flies until the end of the season, usually about the middle of Sep-
tember. The single record for November 12 may represent a third
brood.

Genus POANES Scudder
POANES VIATOR (W. H. Edwards)
Plate 20, j

Range.—Brackish marshes from the Northern Neck to the North
Carolina border. Our records are: Essex County, Tappahannock,
September 2, 1941, marsh north of Tappahannock, September 2,
1941; Gloucester County, Allmondsville, September 1, 1941; Isle of
Wight County, Smithfield, at the bridge over the Pagan River at
Route 10, June 11, 1938, June 10, 1939; James City County, James-
town, June 14, July 5, 1939; King George County, swamp 2 miles
down the road from the Potomac River bridge, August 31, 1941;
King William County, Port Richmond, June 7, 1936; Mathews
County, Foster, August 30, 1941; Middlesex County, Urbanna, Sep-
tember 2, 1941 ; Nansemond County, Chuckatuck, June 15, 1938, east
of Nansemond River to Kings Highway bridge (Otto Buchholz),
northeast of Suffolk, 1944 (Otto Buchholz), Dismal Swamp, June 6,
July 25, 1944 (Otto Buchholz) ; Princess Anne County, Knotts Island,
July 4, 1939, Lake Tecomseh, West Neck Creek at Gum Swamp road
(603) ; Richmond County, east end of the bridge from Tappahan-
nock, June 27, 1937, August 31, 1941 ; Westmoreland County, swamp
along Mattox Creek, Route 205, bridge, August 31, 1941 (W. H.
Wagner, Jr.).

Variation —lIndividuals from the several colonies show slight
differences.

Occurrence.—The broad-winged skipper is confined to marshes
with an abundance of wildrice (Zizania aquatica) ; very local, but
common to incredibly abundant wherever found.

Seasons.—Apparently two broods. The records indicate one brood
from June 6 to July 5, and another from July 25 to September 2.
At Jamestown on July 5 most of the exceedingly numerous individ-
uals were fresh males, indicating that the first brood must fly much
longer than would be inferred from the records. A curious feature
of our records is that, except in Richmond County, we have never
found any indication of more than one brood in any one locality, al-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 169

though we have visited some of the localities, as Smithfield and
the Dismal Swamp, at various times throughout the summer. In Rich-
mond County we found this species in early summer in 1937, and in
late summer in 1941, not at both times in the same year.

POANES HOBOMOK (Harris)
Plate 28, j, k, J

Range.—From Fairfax to Caroline and Prince Edward Counties
and westward, in the mountains southwestward to the North Caro-
lina border. Our records are from Arlington, Bedford, Caroline,
Fairfax, Frederick, Giles, Highland, Montgomery, Page, Prince
Edward, Rappahannock, Roanoke, and Warren Counties. Also the
Dismal Swamp, Nansemond County (Otto Buchholz).

Variation.—Both light and dark females, together with various
intermediates, occur everywhere in Virginia.

Occurrence.—Found in deciduous woods along streams with more
or less grassy banks, and along the borders of woodland bogs, the
males only in the woods, the females also along the edges of the
woods, along roads, and in adjacent fields; generally distributed and
frequent to common in the mountains.

Season.—One brood. This species usually first appears about the
end of the first week in May, rarely as early as the end of April
(April 28), and reaches its maximum in the last week in May and
the first week in June, disappearing at the end of the latter month

(June 30).
POANES ZABULON (Boisduval and LeConte)
Plate 27, a, b

Range.—Throughout the State.

Occurrence.—On the Coastal Plain this species is found in wet
open woodlands, but in the higher country and in the mountains it
lives along the more or less grassy banks of woodland streams, the
females and more rarely the males straying out into adjoining fields;
most generally distributed and most numerous in the higher western
part of the State where it is often frequent or even rather common,
though never abundant ; local and infrequent on the Coastal Plain.

Seasons.—Two broods. This species first appears toward the end
of April, sometimes as early as the end of the third week, reaches a
maximum in the last half of May, and disappears about the middle
of June. The second brood is on the wing shortly after the end of
the third week in July, reaches a maximum in the first half of August,

170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

and flies in decreasing numbers until about the end of the third week
in September, or until the end of the season.

POANES AARONI (Skinner)

Plate 28, m, n, 0, r

Range.—Salt marshes of Chesapeake Bay and the Eastern Shore
(Accomack and Northampton Counties) and offshore islands, and
southward. Our records are as follows: Accomack County, Chinco-
teague Island, August 27, 1891 (F. M. Jones), September 13, 1938
(W. H. Wagner, Jr.), August 21, 1941 (W. H:. Wagner, fee
August 22, 1944 (Otto Buchholz), Assateague Island, May 31, 1929
(F. M. Jones) ; Gloucester County, Gloucester Point, August 31,
1941; Nansemond County, Dismal Swamp, June 5, 1944, May 28,
1945 (Otto Buchholz) ; Richmond County, east end of the bridge
from Tappahannock, August 31, 1941 (W. H. Wagner, Jr.) ; West-
moreland County, Potomac Beach, August 31, 1941 (W. H. Wag-
Net, Wwhice).

Variation.—This species is usually regarded as having two races,
typical aaroni in the north and the larger and darker howardi in
Florida. Specimens from Virginia are practically all of the large
dark Florida type, some females, indeed, having broader dark mar-
gins than any we have seen from Florida. The form aaroni repre-
sents nothing more than a depauperate light and somewhat under-
sized variety of the species which in reality is typified by the Florida
howardi. This depauperate form aaroni occurs occasionally in the
Chesapeake Bay region, especially in the first brood. The relation
between the form aaroni and the normal form howardi seems to us
to be quite the same as that between the small form of Atrytone logan
that occurs in the vicinity of Boston, Mass., and in the mountains
farther south, and the larger typical form.

Dr. Frank Morton Jones and Dr. Warren Herbert Wagner, Jr.,
both independently reached the conclusion that there is no real differ-
ence between aaroni and howardi. Dr. Jones writes us that this species
occurs both in peninsular Maryland and in Sussex County, Del. He
said he doubted any real distinction between aarom and howardi
except average size. Some examples equal in size those from Florida,
while others closely match New Jersey specimens of aaroni. He
said that the 21 specimens in his collection gave the following data,
the expanse being the distance from the apex of the fore wing to
the center of the thorax, doubled: Anglesea, N. J. (2), 32, 34 mm.,
average 33 mm.; Rehobeth, Del. (2), 34, 38 mm., average 36 mm.;

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK GA

Ocean City, Md. (6), 34, 36, 36, 37, 38, 42, average 37 mm.; Shell-
ton, Md. (1), 35 mm.; Royal Palm State Park, Fla. (10), 32, 32,
34, 37, 38, 38, 38, 40, 40, 40, average 37 mm. He noted that the
New Jersey specimens are perhaps a trifle paler than the others of
like freshness. On the characters given for their separation most
peninsula specimens fall to howardi rather than to aaronit. We have
examined a large series of specimens taken by Mr. Wagner at Chin-
coteague Island and at various points in Chesapeake Bay nearly all
of which are referable to howardi. We see no reason for the re-
tention of the name howardi. Dr. W. J. Holland in the revised edi-
tion of “The Butterfly Book,” 1931, wrote under Poanes howardi:
“All of the specimens I possess show a pale longitudinal streak on
the lower side of the hind wing, which does not occur in P. aaront.”
There is really no difference in this respect between the two forms.

Occurrence.—Found in the salt marshes, as noticed by Dr. Wagner
always in association with the salt-marsh grass (Spartina alterniflora
var. glabra), often straying for some distance into adjacent fields;
local, but common to abundant wherever found; abundant along the
causeway from the mainland to Chincoteague Island.

Seasons.—Two broods. Our records, supplemented by others from
Maryland and Delaware kindly given us by Drs. Frank Morton
Jones, George W. Rawson, and Warren Herbert Wagner, Jr., indi-
cate that this species first appears at the end of May and flies almost
or quite through June. The second brood appears at about the end of
the third week in August (August 21) and flies until the end of the
season, about the middle of September (September 13).

During a recent visit to Emory University, Atlanta, Ga., in com-
pany with Lucien Harris, Jr., Mr. Harris showed the senior author
a bound volume of original water colors by John Abbot that he had
recently presented to the University library. The figures were chiefly
of birds, but some of the plates included butterflies also. Among
the butterflies figured was this species, as well as the somewhat
similar Atrytone arogos.

POANES YEHL (Skinner)
Plates 28, q; 30, d, f, g

Range.—Deep swamps in Nansemond, Norfolk, and Prince George
Counties. Our records are: Nansemond County, Dismal Swamp,
near Suffolk, fourth week of July, 1925 (F. M. Jones), September 1,
1935, July 5 to August 20, 1944 (Otto Buchholz), Suffolk, Septem-
ber 4, 1936, July 27, 1938, Holland, July 5 to August 20, 1944

172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

(Otto Buchholz) ; Norfolk County, Green Sea, September 2, 1935,
North Landing River swamp, September 5, 1936; Prince George
County, Second Swamp near New Bohemia, July 19, 1941.

Variation.—This species is unusually variable. There are two
quite distinct forms connected by a complete series of intergrades.
In one of the forms the under side of the hind wings is cinnamon
with the light spots clearly defined and conspicuous (pl. 30, g). In
the other the under side of the hind wings is yellow with the spots
vaguely and indefinitely outlined and scarcely contrasting with the
light background (pl. 30, d). In the males the dark border may be
scarcely half the usual width, the ground color may be light and
rather bright yellow, and the stigma may be yellow with narrow
black edges (pl. 28, q).

Occurrence—Found in thick and very wet deciduous swamps,
occasionally straying to more open and drier localities; we found it
abundant in the North Landing River swamp, and Mr. Buchholz
found it abundant in the Dismal Swamp and at Holland. It is ex-
ceedingly shy and difficult to catch.

Season.—Apparently one brood, flying from July 4 to September 4.

POANES MASSASOIT (Scudder)

The subspecies represented in the vicinity of Virginia is—

POANES MASSASOIT HUGHI A. H. Clark
Frontispiece, fig. 11; plate 26, c, d

Range.—This butterfly has not been taken in Virginia, but it is
abundant in bogs at Hyattsville and Beltsville, Md., a short dis-
tance north of the Potomac, and probably will be found in nearby
Virginia.

Occurrence.—Found in very wet bogs surrounded by woods and
subject to inundation in the spring; very local, but abundant wher-
ever found.

Season.—One brood. This butterfly is on the wing from July 1 to
the last week in July.

Genus ATRYTONE Scudder
ATRYTONE AROGOS (Boisduval and LeConte)
Plate 11, a, b
Range.—Known only from Montgomery County. Prof. Ellison

A. Smyth, Jr., writes us that he has taken this species at Blacks-
burg but that it is not common.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 173

Occurrence.—Found in open fields.
Season.—Apparently one brood. The records for Tryon, N. C., are
from the latter part of June to August.

ATRYTONE LOGAN (W. H. Edwards)
Plate 27, c, d, e

Range.—Throughout the State.

Variation.—This species varies very considerably in size along the
eastern seaboard. In a series from Newtonville, near Boston, Mass.,
the fore wings are 12-13 mm. long. Over most of Virginia the
fore wings are 14-15 mm. long. In southern Princess Anne County
large individuals occur with the fore wings 16-18 mm. long; they
resemble others from Florida. In Isle of Wight County we have
found large individuals resembling those from southern Princess
Anne County, or from southern Florida, flying with more numerous
smaller ones.

Occurrence.—Found in bogs and wet meadows in the higher coun-
try, in marshy areas and near wet woods on the Coastal Plain; very
local; infrequent to frequent; never very common.

Seasons.—Two broods. This species first appears at the end of
May and flies until the second week of July (July 10). The second
brood is on the wing shortly after the middle of July (July 17) and
flies until the end of the season in the first week of September.

ATRYTONE PALATKA (W. H. Edwards)
Plate 20, g, h, 1

Range.—Southern Princess Anne County; Otto Buchholz took
this species near Munden, June 21, 1939; in southern Princess Anne
County on June 16, 1941; and along Route 614 2 miles north of the
North Carolina line on June 16-22, 1944. He was so kind as to
present a pair of Virginia specimens to the United States National
Museum. We have not found it.

ATRYTONE DION (W. H. Edwards)
Plate 20, e

Range.—Bogs and swamps in Nansemond, Norfolk, and Princess
Anne Counties. Our records are: Nansemond County, Dismal Swamp
near Suffolk, June 11, 14, 1937, June 11, 1939, Dismal Swamp,
April 20, 1945 (Otto Buchholz) ; Norfolk County, Green Sea, Sep-
tember 2, 1935, North Landing River swamp, June 12, 1939, June

174 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

19, 1944 (Otto Buchholz), Northwest, May 31, 1937, Wallaceton,
June 12, 1938, July 5, 1941; Princess Anne County, Gum Swamp
road at the bridge over West Neck Creek, September 2, 1940, Lake
Tecomseh, July 3, 1939, North Landing River swamp, July 5, 6, 1941,
Seaside State Park, September 5, 1936.

Occurrence.—Found in wet sedgy hollows in pine barrens and in
sedgy areas in swamps; usually infrequent to frequent, sometimes
common. We have found this species most numerous where the
Gum Swamp road crosses West Neck Creek, 1 mile west of Pungo,
Princess Anne County. It is fairly common in the wet hollows among
the sand dunes west of the road north from Virginia Beach.

Seasons.—Two broods. This species first appears toward the end
of April (April 20) and flies through May and June and into the
first week in July. The second brood probably appears toward the
end of August and flies until the end of the season in September. We
have found this butterfly most numerous early in September.

ATRYTONE ALABAMAE Lindsey
Plate 20, f

Range-——Known only from Accomack and Nansemond Counties.
Our records are: Accomack County, central portion of the Dahl
Swamp three-quarters of a mile southeast of Cashville, July 23, 25,
1935; Nansemond County, Dismal Swamp, May 29, 1945 (Otto
Buchholz).

Discussion.—We do not believe that this form can be conspecific
with dion, for the following reasons: Though it is very similar to
dion, we have seen no intergrades between the two. It is most numer-
ous in the latter part of July when dion is not flying. It is asso-
ciated with a different type of sedge (Carex striata var. brevis),
which is presumably its food plant.

We have before us a series of 27 males from the Dahl Swamp,
and also Professor Lindsey’s type specimen, a male from Mobile,
Ala., for the privilege of examining which we are indebted to Dr.
Hugo Kahl, who personally brought it to us from Pittsburgh, Pa.

On the upper surface some of our males agree perfectly with Lind-
sey’s type specimen, though most of them are somewhat brighter
with the fulvous band on the fore wings bordering the stigma ex-
ternally somewhat broader; one is even darker, with this band nar-
rower. This fulvous band is usually less than half as broad as the
distance between its outer edge and the edge of the wing, whereas in
dion it is nearly or quite as broad as the distance between its outer

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 175

border and the edge of the wing. In alabamae the fulvous band is
relatively short, ending abruptly at the upper end of the stigma, and
is not practically continuous with the subapical spots as it is in dion.
In the type of specimen of alabamae the subapical spots on the fore
wings are reduced to one, small and inconspicuous, with a faint ves-
tige of another above it. They are similarly reduced in some speci-
mens from the Dahl Swamp, though in these there are usually two
present.

In alabamae the fulvous dash on the hind wings is always short,
about 3 mm. long as in the type specimen, and in some individuals it
it absent. In dion it is always present and is 5-6 mm. long.

On the under side the specimens from the Dahl Swamp agree
perfectly with the type specimen, except that in all but one the spots
on the under side of the fore wings are larger. The ground color of
the under side of the hind wings is darker and duller than in dion.

An interesting feature of the type specimen of alabamae is the
narrow stigma, which is not much more than half as broad as the
stigma of the type specimen of dion. In a few of the specimens from
the Dahl Swamp, the stigma is similarly narrow, though usually it is
more as it is in dion,

In the eight females from the Dahl Swamp, which we have com-
pared directly with the female type of dion from Whiting, Ind., the
spots on the fore wings are smaller, especially the lowest, and the
one above this is usually absent. The dash on the hind wings is short,
dark, and inconspicuous.

Occurrence—In the Dahl Swamp confined to the wet, open, sedgy
areas in the central portion; very common in the limited areas in
which it occurs.

Seasons.—Probably two broods. Mr. Buchholz took it on May 29,
and we found it on July 23 and 25.

ATRYTONE DUKESI Lindsey
Plate 290, a, b, c, d

Range—Known only from the deep swamp bordering the North
Landing River and Pocaty Creek in Norfolk and Princess Anne
Counties. Our records are: Norfolk and Princess Anne Counties,
swamps bordering the North Landing River, Pocaty Creek, and the
Chesapeake and Albemarle Canal, June 14, July 3, 1938, June 12, 13,
23, July 2, 1939, July 5, 16, 1941, June 16-26, 1944 (Otto Buchholz).

Occurrence.—Found in the interior and along the borders of very
wet gum (Nyssa aquatica) swamps, seldom venturing out of the deep

176 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

shade of the swamp vegetation so that it is easily overlooked ; com-
mon in the very limited areas in which it occurs.

The most practicable way to obtain this species in quantity is from
a boat on the North Landing River and along the Chesapeake and
Albemarle Canal. It may also be collected in quantity by wading
through the swamp, which must be done with caution because of the
presence of water moccasins (Agkistrodon piscivorus) and, in the
drier areas, of canebrake rattlers (Crotalus horridus).

Variation.—When freshly emerged this species is sooty black, but
it soon becomes more or less brown. Museum specimens are always
brown contrasting strongly with freshly caught individuals.

Season.—One brood, June 12 to July 16.

ATRYTONE CONSPICUA (W. H. Edwards)
Plate 27, g, h

Range.—Known only from Giles and Nansemond Counties. Our
records are: Giles County, Little Meadows, near Mountain Lake,
July 21, 25, 26, 1940, July 1941 (Lloyd G. Carr and Carroll E. Wood,
Jr.), July 21, 1949 (Carl W. Gottschalk) ; Nansemond County, Dis-
mal Swamp, June 3, 1944 (Otto Buchholz). Mr. Wood has kindly
presented specimens taken on July 25, 1940, to the United States
National Museum.

Occurrence-—Found in bogs in, or adjacent to, woods, and along
the sedgy and marshy banks of slow woodland streams in hilly coun-
try; very local, but usually abundant wherever found.

Mr. Wood raised this species on Carex stricta, and presented lar-
vae to the United States National Museum.

Season.—One brood. All the specimens from Giles County were
taken from July 21 to 26. At Beltsville, Md., this species flies from
the last week in June to about the end of July. Apparently it flies
earlier in the Dismal Swamp, as might be expected.

ATRYTONE BIMACULA (Grote and Robinson)
Plate 27, 7, k

Range.—One Virginia record, August County, Mountain Lake
(Shenandoah Acres), July 5, 1937.

Occurrence.—Confined to permanently wet meadows and bogs.
This species is common at Caudy’s Castle in Hampshire County,
W. Va., a few miles west of the Frederick County line, where numer-
ous specimens have been taken by Warren Herbert Wagner, Jr.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 177

Season.—One brood. In this region this butterfly probably flies
from about the middle of June to about the middle of July.

ATRYTONE RURICOLA (Boisduval)

The subspecies represented in Virginia is—

ATRYTONE RURICOLA METACOMET (Harris)

Plates 11, e; 26, e

Range.—Throughout the State.

Variation —Intergrades between the eastern form (metacomet
Harris) and typical western ruricola are occasionally found in eastern
Virginia, and some individuals would certainly be referred to the
western form if their origin were not known. We have seen two males
from the Dismal Swamp in which the stigma was bordered on the
outer side by a continuous narrow whitish stripe. Immaculate females
are occasionally found.

Occurrence.—Found in damp meadows and boggy pastures and
along the grassy borders of swamps and marshes, straying more or
less into drier territory; fairly numerous everywhere, except in the
highest mountain areas, and locally common.

Seasons——Two broods. This species first appears at about the
end of the third week in May (May 21) and becomes common later
in the month and in the first half of June. The numbers then decrease,
though individuals of the first brood are still on the wing when the
second brood appears at about the end of the third week in July. The
second brood is most numerous toward the end of July and in the
first half of August, after which time the numbers decline and the
butterfly disappears in the first week in September.

Genus ATRYTONOPSIS Godman and Salvin
ATRYTONOPSIS HIANNA (Scudder)
Plate 26, q

Range.—From Fairfax to Westmoreland and Henrico Counties, and
westward to Alleghany County; probably also southwestward in the
mountains, as it occurs in central and western North Carolina (Ra-
leigh and Tryon). Our records are from Alleghany, Fairfax, Fau-
quier, Henrico, Prince Edward, Roanoke, and Westmoreland
Counties.

V ariation—Most individuals have two small, but conspicuous, white
spots on the under side of the hind wings near the anterior edge,

178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

one near the base and the other just beyond the middle, but the outer
spot is frequently absent, and both spots may be absent. Occasional
individuals may have more spots, sometimes as many as Six, scat-
tered over the under surface of the hind wings.

Occurrence.—Found in dry open fields and brushy woodland clear-
ings ; very local, but often frequent or even common where it occurs.
It is often found in the same localities as Hesperia metea.

Season.—One brood. This species appears in the first week in May
(rarely in April), becomes numerous in the last half of the month,
and disappears early in June.

Genus LEREMA Scudder
LEREMA ACCIUS (J. E. Smith)

Plate 27, 0, r

Range.—Throughout the Coastal Plain, in the extreme southern
part of the central portion of the State, and in the valley bottoms of
the western mountains as far northeast as Augusta and Madison
Counties. Our records are from Accomack, Arlington, Augusta,
Giles, Isle of Wight, King William, Lancaster, Madison, Mecklen-
burg, Montgomery, Nansemond, Norfolk, Northampton, Patrick,
Prince George, Princess Anne, Roanoke, Southampton, Sussex,
Warwick, and Wythe Counties.

Occurrence.—Found in open fields, particularly along the road-
sides and the borders of woods, along the sides of roads through
woods, and occasionally in open woods, in low and usually more or
less damp regions or valley bottoms; very irregular in its occurrence;
usually common in late summer on the outer Coastal Plain, sometimes
(as in 1935) the commonest skipper, though in some years scarce or
(as in 1941) apparently absent; elsewhere infrequent or casual, or
apparently absent.

Seasons.—Three broods. From our records it would appear that
this species is only a summer visitor to Virginja—at least in most
years. The only Virginia record that can represent the spring brood
is that of a single individual taken on May 12, 1938, in Roanoke
County. This butterfly appears usually in the second week in June, its
numbers reaching a maximum in the first half of July, and becoming
greatly reduced in August. A new brood, much more numerous than
the midsummer brood, appears about the first half of September and
flies until the end of the season early in October.

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK 179

Genus AMBLYSCIRTES Scudder

AMBLYSCIRTES VIALIS (W. H. Edwards)
Plate 27, n, q

Range.—Throughout the State.

Occurrence.—Found in rich open deciduous woods, along roads
through woods, and along the borders of woods; in the north and
in the western mountains generally distributed and often locally com-
mon; more local, though in some places common, on the Piedmont
and on the Coastal Plain; rare in the south-central and southeastern
sections.

Seasons.—Two broods. The roadside skipper appears in the second
week in April (April 11) and flies through May and into the first
week in June (June 6). The second brood appears early in July
(July 6) and flies until just after the middle of August (August 18).

AMBLYSCIRTES HEGON (Scudder)
Plates 25, 1; 27, |

Range.—Known only from Henrico, Warren, Botetourt, Roanoke,
and Montgomery Counties. Our records are: Henrico County, Rich-
mond, May 16, 1937; Warren County, Limeton, May 12, 1941;
Botetourt County, Gala, May 16, 1943; Roanoke County, Ash Bot-
tom, near Salem, April 23, 1938, Dixie Caverns, May 5, 1940, Fort
Lewis Mountain, May 2, 1937, April 23, 1938, Salem, 1927, April 24,
1938; Montgomery County, Blacksburg, June 2, 1899. It is probably
locally distributed in the northern half of the State and throughout
the western mountains.

Occurrence.—Found in open places in woods, along roads through
woods, and about the borders of woods; very local and never very
numerous ; it is common at Berkeley Springs, W. Va.

Season.—One brood. According to our records this species is on
the wing from April 23 to June 2.

AMBLYSCIRTES ALTERNATA (Grote and Robinson)
Plate 25, 1, m

Range.—Nansemond County, taken by Otto Buchholz on June 5
and from July 15 to August 15, 1944, and in the Dismal Swamp on
April 26, 1945. Mr. Buchholz has kindly presented a pair of Virginia
specimens to the United States National Museum,

180 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

AMBLYSCIRTES TEXTOR (Hiibner)
Plate 3, 7

Range.—From Princess Anne westward to Surry, Prince George,
and Greensville Counties. Our records are from Greensville, Nanse-
mond, Norfolk, Prince George, Princess Anne, and Surry Counties.

Occurrence.—Found in thick wet woods, especially in the glades
and along the roads; generally distributed but usually not common,
and met with as occasional individuals resting on leaves ; locally where
conditions are especially favorable and flowers (particularly Prunella
or Elephantopus) are present, as in various localities along the west-
ern border of the Dismal Swamp, it is common to exceedingly abun-
dant. This species varies considerably in numbers in different years,
wet years being most favorable for it.

Seasons ——Two broods. This butterfly first appears in the latter
part of May, becomes abundant in June and early July, and disap-
pears before the first of August. The second brood is on the wing
at the end of August, becomes common in early September, and flies
until the end of the season in late September (September 23). Otto
Buchholz has taken it in the Dismal Swamp as early as April 27, so it
is possible that there is a spring brood we have overlooked.

AMBLYSCIRTES CAROLINA (Skinner)
Plates 28, p; 30, a, b,c

Range.—From Princess Anne County westward to Prince George
and eastern Greensville Counties.

V ariation—This species, always easily recognizable by the spotting
on the upper side of the fore wings (pl. 28, p), varies greatly in the
color of the under side of the hind wings. Most commonly the under
side of the hind wings is light, rather dull yellow with a submarginal
row of cinnamon or rusty-brown spots, and a number of similar spots
are evenly distributed between this row and the wing base (pl. 30, a).
In a large percentage of the individuals the ground color of the hind
wings is a uniform cinnamon or rusty brown with a submarginal row
of small yellow spots and a long or short streak in the cell (form
reversa F. M. Jones, pl. 30, b). In extreme cases these submarginal
spots may be reduced to two or three very small spots, or in worn
individuals they may be wholly obscured so that the under side of the
hind wings is uniform brown varying from a rich chestnut to light
yellow-brown, with a more or less conspicuous narrow or broad streak
in the cell (pl. 30, c).

The usual form in Virginia is carolina, but reversa is found every-

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 181

where with it and is especially common in the Dismal Swamp near
Suffolk, where sometimes more than half the individuals captured are
referable to it. The immaculate form is not common; we have taken
it in the Dismal Swamp near Suffolk, and near Surry.

Occurrence—Found in wet woods with an undergrowth of Arun-
dinaria tecta near swamps or sluggish streams; generally distributed
but not common; most numerous locally along the western border of
the Dismal Swamp and in southern Princess Anne County.

Seasons.—There appear to be three broods. In 1938 we captured
three worn females on April 1 so that the butterfly must have been
on the wing since about the middle of March, and Otto Buchholz has
taken it in the Dismal Swamp on April 26. It reappears at the end
of May, becomes common in June, and flies through July and into
August. The third brood is on the wing toward the end of August,
and the individuals are most numerous in early September.

Genus LERODEA Scudder
LERODEA L’HERMINIERI (Godart)
Plate 11, c, d

Range.—Throughout the State.

Variation.—The males of this species may be uniform brown above,
or the fore wings may have a more or less diffuse large dark brick-
red patch including the inner half of the interspace between veins
2 and 3, the basal portion of the interspace just above, the entire
lower border of the wing below vein 1, and the borders, or almost
the whole of the interspace between veins 1 and 2. We have taken
specimens with this patch extensively developed at Aylett, King Wil-
liam County, and elsewhere. All degrees of development of this patch
are found. In faded specimens this patch becomes dark dull yellowish.

Occurrence-—Found in open fields, more or less damp or with a
thick growth of luxuriant grass or clover, within which it is very
local, confined to limited areas sometimes of only a few hundred
square feet within which it may be common but from which it seldom
strays. Because of its localization, its habit of usually keeping well
down in the grass, its small size, and its inconspicuousness, it is easily
overlooked. ;

Virginia appears to be approximately the northern limit of this
species as a permanent resident, for here it seems to be mostly killed
out during the winter. Our records for May are all from the Coastal
Plain (Norfolk, Nansemond, Westmoreland, and Caroline Counties)
and from the west (Roanoke and Albemarle Counties), and the indi-

182 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. II6

viduals represented by these records are very few. June records add
Prince George, Dinwiddie, and Fairfax Counties, and the District of
Columbia area in the east, Prince Edward County in the center, and
Montgomery County in the west. These additions only confirm the
localization indicated by the May records, except for Prince Edward
County, which might easily be reached by individuals raised in North
Carolina. We are inclined to believe that over most of Virginia this
species is only a summer visitor.

Seasons.—Two broods. This species first appears about the middle
of May (earliest date May 12) and is common in the latter part of
May and in the first half of June, after which its numbers fall off
and it disappears in the first week in July. The second brood is on
the wing shortly after the middle of July (July 18), and the insect is
common from the end of July until about the middle of September,
sometimes continuing on the wing into early October. It is most
numerous in August, especially in the first half of the month,

LERODEA EUFALA (W. H. Edwards)
Plate 24, q

Range.—Found on the outer Coastal Plain as far west as Nanse-
mond County, and in the southwest as far northeast as Roanoke and
Nelson Counties. Our records are: Nansemond County, Brinkley,
September 26, 1937, Chuckatuck, October 1, 1937, Suffolk, Septem-
ber 26, 1937, North Carolina line at Route 53, September 24, 1937;
Nelson County, Lovingston, August 12, 1937; Norfolk County, east
of the Dismal Swamp, September 2, 1935; Princess Anne County,
Virginia Beach, dunes south of the military airport, September 30,
1937 ; Roanoke County, Fort Lewis, September 6, 1937, July 23, 1939,
Salem.

Occurrence.—Found in open fields. So far as is known this species
is only a summer visitor to Virginia, where there is no evidence that
it survives the winter. It has recently been recorded from the Dis-
trict of Columbia. This species is reported as apparently a recent
invader in California, rapidly becoming common at Indio. Leussler
says that whereas it was formerly rare in Nebraska it is now abundant
there during September and early October, and is spreading north-
ward.

Season.—Only the summer brood is known from Virginia, flying
from the last week in July (July 23) until early in October (Octo-
ber I) ; most of the records are in the first week in September. Dr.
Warren Herbert Wagner, Jr., captured a male in the District of
Columbia on September 7, 1935.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 183

Genus CALPODES Hiibner
CALPODES ETHLIUS (Cramer)
Plate 23, f

Range.—Confined to the Coastal Plain, extending westward to
Southampton County, Petersburg, Richmond, Fredericksburg, Alex-
andria, and Washington, D. C. Our records are from Alexandria, and
Accomack (Tangier Island, September 18, 1939), Arlington, Din-
widdie, Gloucester, Henrico, Isle of Wight, James City, King William,
Nansemond, New Kent, Norfolk, Princess Anne, Southampton, Spot-
sylvania, Surry, Sussex, Warwick, Westmoreland, and York Counties.

Occurrence.—Very erratic; commonly, or perhaps usually, present
in small numbers on the outer Coastal Plain, though in some years
wholly absent; occasionally abundant over the entire Coastal Plain,
doing great damage to garden cannas.

All our records, from 52 localities, are for the late summer of 1937,
between September 8 and October 3, except the following: Tangier
Island, September 18, 1939; Richmond, July 1934 (Carroll M. Wil-
liams), September 10, 1938 (Carroll M. Williams) ; and Virginia
Beach, August 21, 1936 (Warren H. Wagner, Jr.).

In 1937 we planned an extensive investigation of the Dismal Swamp
region between September 24 and October 3, but the weather was
rainy and no butterflies were flying. So we devoted ourselves to the
examination of canna beds to determine the presence of the larvae
of this species. Larvae were found throughout the Coastal Plain,
where they had done extensive damage to cannas everywhere.

We have never seen the adult of this species in Virginia although
we have found thousands of the larvae, some of which we raised to
maturity. All our records are for larvae except for a single adult
from Virginia Beach taken on August 21, 1936, by Dr. Warren Her-
bert Wagner, Jr., and one seen at Franklin, Southampton County, by
Dr. Carroll M. Williams on September 8, 1937. In the Department
of Agriculture grounds at Washington, D. C., we saw two females
ovipositing on canna leaves between I and 1:30 p.m. on Septem-
ber 14, 1937.

Dr. Carroll M. Williams, who reared this butterfly from larvae
found on cannas at Richmond in July 1934, made a special study of
it in the great outbreak of 1937. His attention was first drawn to it
in the week of June 20 when, as a result of a telephone call from his
cousin, Gordon Williams, he found several larvae on the identical
plant on which he had found others in 1934. Later he found larvae
in great abundance at all points on the Coastal Plain. But he did not

184 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

see an adult under natural conditions until September 8. He writes
us that on that date, at Franklin, at 2: 45 p.m., he observed a female
ovipositing. Upon being attracted to a group of green cannas by
their tattered appearance, he saw a female dashing madly about, so
fast that the eye could scarcely follow it. At intervals it would return
and come to rest on the canna leaves. After about half a minute or
less it would again dart into flight. Upon examination of the point
where it had been resting, from one to four bluish-green eggs were
found. These were dome-shaped and minutely reticulated. Close
examinations of the plants revealed many eggs deposited on both sur-
faces of the leaves, as well as fragments of eggs from which the
larvae had emerged. Hundreds of larvae in all stages were present,
as well as a few pupae and larvae that were suspended for pupation.

In the Department of Agriculture grounds we noted that the female
dashes up to the cannas, alights on a leaf, and with much deliberation
attaches an egg. When this is done, after a minute or so, she dashes
off at great speed and is soon lost to view.

On August 26, 1927, Dr. Williams made an examination of a canna
bed in a cemetery at Richmond and found much damage to have been
done. A few very young larvae were still present, but most of the
insects had disappeared. A week before, he had seen dozens of full-
grown larvae in this one canna bed, but he now found only three
pupae and one larva that was ready to pupate. He examined what
had formerly been the shelters of the larvae but in practically every
case the folds seemed to have been devoured. He did find a partial
explanation. Of the three pupae he took, only one was in its leaf roll
in accordance with its habits as reported in the literature on the sub-
ject. The two others were on the under side of the leaf and next to
the midrib. No complete fold was made, but rather the leaf was
drawn downward in a sort of inverted trough effect, in the upper
concave of which the pupa was suspended.

On September 12, 1937, we examined a large plat of fancy cannas
at the Agricultural Station north of Alexandria. Larvae in all stages,
and pupae, were abundant. In making the shelters the edge of the
leaf may be broadly folded inward without any cuts, usually over
the upper surface, but sometimes over the lower, or the leaf may be
cut above or below the fold, or cut and more or less eaten in both
places. The pupae are usually found on the higher fully developed
leaves. They usually lie in a broad fold over the upper surface of
the leaf, though they may be in a fold on the lower surface, or they
may lie transversely on the lower surface with the leaf gabled above
them.

NO. 7 BUTTERFLIES OF VIRGINIA—CLARK 185

We have watched a caterpillar fold over the edge of a leaf. Several
very wet threads were spun, making a slight concavity in the leaf
surface. As the threads dried they shortened and the edge of the leaf
was thus drawn in over the upper surface. The actual folding of the
leaf is not done by the caterpillar, but by the drying and shrinking
of the threads.

The upper leaves of the cannas are those usually attacked, and we
have seen many canna beds in which the upper leaves of all the plants
were reduced to little more than the midribs. Dr. Williams tells us
that in some beds in Richmond the leaf infestation seemed to be
nearly 100 percent, what had formerly been leaves being represented
in many cases by only the central thick midrib.

We noticed in the Department of Agriculture grounds at Wash-
ington that larvae were found on September 20, 1937, in all the beds
of cannas with green leaves, but none were found in a bed of red-
leafed cannas.

Season.—Probably one brood in Virginia. The earliest records for
adults that we have are for July 9 and 11, 1937, when two emerged
from pupae sent us from Richmond by Dr. Williams. From this time
on the butterfly is continuously on the wing in increasing numbers
until the end of the season early in October. Caterpillars have been
reported from Richmond about the middle of June. These are from
eggs that have been deposited by females coming from the south, for
this butterfly is not known to survive the winter north of South
Carolina. From June onward caterpillars in all stages may be found,
but from about the third week in August large caterpillars become
scarce and small ones relatively much more numerous. These small
caterpillars are evidently the young of adults raised in Virginia, but
there is no evidence that any young from Virginia-raised parents
reach maturity before the end of the season.

Genus PANOQUINA Hemming
PANOQUINA PANOQUIN (Scudder)
Plate 26, |

Range.—Found in salt marshes along the entire coast. We have
records from Accomack, Gloucester, Mathews, Middlesex, Northamp-
ton, Princess Anne, Richmond, and Westmoreland Counties.

Variation When fresh the under side of the hind wings has a
marked yellowish tinge and the veins are dull yellow, contrasting with
the white streak beyond the end of the cell. In flight freshly emerged

*

186 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

individuals appear brownish yellow. The yellow soon fades to dull
grayish and is never preserved in museum specimens.

Occurrence.—Confined to salt marshes with an abundance of sedge
(Scirpus), the roots of which are covered at high tide, seldom stray-
ing inland much beyond the high-tide mark; common to abundant
wherever found.

Seasons.—Three broods. The first brood appears at the end of May
(May 31) and flies until the first week in July. The second brood
flies abundantly in the last half of July. The third brood appears at
about the end of the third week in August (August 23) and flies
until after the middle of September (September 18).

PANOQUINA OCOLA (W. H. Edwards)
Plate 26, p

Range.—Occurs chiefly on the outer Coastal Plain, ranging west-
ward to Arlington, Fairfax, Prince William, Henrico, Sussex, and
Greensville Counties ; also found in the southwest as far northeast as
Nelson County. Our records are from Arlington, Gloucester, Greens-
ville, Henrico, Isle of Wight, Mathews, Nansemond, Nelson, Nor-
folk, Prince William, Princess Anne, Roanoke, Southampton, Staf-
ford, Sussex, and Wythe Counties.

Occurrence.—This species is an inhabitant of open fields and gar-
dens in low and more or less damp regions. It is very variable in
its occurrence, being usually fairly common late in summer, in some
years (as in 1937) abundant, in others (as in 1941) scarce.

Seasons.—Three broods. Apparently this species does not usually
survive the winter in Virginia but comes into the State each summer
from farther south. We have only two records that can be considered
as representing the spring brood, one in late May (May 28, Suffolk),
and one near the middle of June (June 13, North Landing River).
The midsummer brood usually appears shortly after the first of July
(July 3) and flies until about the first week in September. The third
brood, in which the individuals are much more numerous than they
are in the midsummer brood, appears after the third week in Sep-
tember and flies into early October, sometimes as late as October 26.

ERRONEOUS RECORDS

The following species have been erroneously credited to Virginia
in the literature:

Limenitis arthemis (Drury): The white-banded form credited to
Virginia by Emmons does not occur in the State.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK 187

Lycorea cleobaea (Godart): Recorded from Virginia by error for
Virgin Islands.

Ithomia drymo (Hubner): From Virginia, Minas Gerais, Brazil.

Hemuargus hanno (Stoll) : Credited to Virginia by Boisduval and
LeConte, but there is no evidence that it occurs in the State.

Pieris virginiensis W. H. Edwards: Credited to Virginia by
Strecker in error for West Virginia.

Colias alexandra edwardsii W. H. Edwards: Given in Seitz from
Virginia by error for Virginia City, Nev.

Eurema elathea (Cramer): We have no evidence that this species
is found in the State.

Erora laeta (W. H. Edwards): Listed by Strecker from Vir-
ginia by error for West Virginia.

Erynnis martialis ab. ausonius (Lintner): Given by Strecker from
Virginia: probably E. martialis.

Potanthus mingo (W. H. Edwards) : A Philippine species described
from West Virginia, identified by Evans, who gave the locality as
Virginia.

Ochlodes sylvanus (Esper) : In the revised edition of “The Butter-
fly Book,” 1931, Dr. W. J. Holland wrote: “I have in my possession
a male of this European species taken at Richmond, Virginia. Its
chrysalis may have been brought over in hay, or packing material, and
the butterfly emerged on this side of the Atlantic. It differs in no
respect from specimens from Germany, of which I have many...
It is a stray immigrant.” We wrote to Dr. Walter R. Sweadner,
curator of entomology, Carnegie Museum, Pittsburgh, regarding this
record. He very kindly looked up the matter for us, and replied on
November 3, 1947: “We have no specimen of Augiades [=Ochlodes]
sylvanus from North America. I do not know anything with regard
to the specimens about which Dr. Holland wrote.”

BIBLIOGRAPHY

The following list includes all the books and articles of importance
to the student of the butterflies of Virginia.

The first butterfly recorded from “Virginia” was a yellow female
of Papilio glaucus, a description and figure of which was published
by Thomas Moufet in 1634. The figure was from a painting made
by John White in 1587 in North Carolina, then a part of Virginia.
There are many later references to this description and figure.

A few early records from “Virginia” refer to West Virginia, which
was a part of Virginia until 1863. Rothschild and Jordan record

188 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I16

certain swallowtails collected by Col. Wirt Robinson near his home
in Nelson County as from “W. Va.” instead of western Virginia.
A number of species have been recorded from Virginia by error for
the Virgin Islands; Virginia, Brazil; and Virginia City, Nev.

ANONYMOUS.

1890. [Papilio palamedes in Virginia.] Ent. News, vol. 1, No. 7, p. 110.
BAILEY, JOHN WENDELL.

1939. Biology at the University of Richmond, pp. 1-194. Richmond, Va.
Bete, BE:

1923. Amblyscirtes textor Httbner from Virginia. Bull. Brooklyn Ent.
Soc., vol. 18, No. 1, p. 12, February.

1926. Collecting at Wilmington, North Carolina, and Suffolk, Virginia.
Journ. New York Ent. Soc., vol. 34, No. 4, pp. 351-354, December.

BotspUVAL, JEAN Baptiste ALPHONSE DECHAUFFOUR DE, and LECoNTE, JOHN.

1829-1837. Histoire générale et iconographie des lépidoptéres et des chenilles
de l’Amérique septentrionale, vol. 1, pp. 1-228, pls. 1-78, May 2,
1829, to July 24, 1837. Paris.

BRIMLEY, C. S.

1938. The insects of North Carolina. North Carolina Dept. Agr., pp. 1-560.
Papilionoidea (revised by Austin H. Clark), pp. 255-263. Sup-
plement, pp. I-39, 1942. Butterflies, p. 16.

CaTEesBy, Mark.

1731-1743. The natural history of Carolina, Florida, and the Bahama
Islands; containing the figures of birds, beasts, fishes, serpents,
insects and plants . . . with their description in English and French,
&c. Histoire naturelle de la Caroline, &c. (appendix), 2 vols.,
colored pls. London.

CHERMOCK, RALPH L.

1942. Notes on collecting Argynnis diana. Proc. Pennsylvania Acad. Sci.,

vol. 16, pp. 59-61.
CHITTENDEN, F. H.

1909. Some insects injurious to cabbages, cucumbers and related crops.
Virginia Truck Exp. Stat., Norfolk, Va., Bull. No. 2, pp. 19-48,
figs. 1-17, Sept. 20.

Criark, AusTIN H.

1932. The butterflies of the District of Columbia and vicinity. U. S. Nat.
Mus. Bull. 157, pp. i-ix, 1-337, pls. 1-64, Feb. 13.

1934. Observations on the butterflies of Apple Orchard Mountain, Bedford
County, Virginia. Proc. Biol. Soc. Washingtgn, vol. 47, pp. 177-
180, Oct. 2.

1035. [Hesperia metea and Thorybes confusis from Difficult Run, Virginia.]
Proc. Ent. Soc. Washington, vol. 37, No. 8, p. 169, November.

1936a. Notes on the butterflies of the genus Enodia and description of a new
fritillary from Peru. Proc. U. S. Nat. Mus., vol. 83, pp. 251-259,
pl. 22, Apr. 11.

1936b. The gold-banded skipper (Rhabdoides cellus). Smithsonian Misc.
Coll., vol. 95, No. 7, pp. 1-50, pls. 1-8, figs. A-D (p. 26), E-H
(p. 29), May 6.

NO.

7 BUTTERFLIES OF VIRGINIA—CLARK 189

1936c. Who’s who among the butterflies. Nat. Geogr. Mag., vol. 69, No. 5,
pp. 679-602, May.

1937a. The butterflies of Virginia. Expl. and Field Work of the Smith-
sonian Inst. in 1936, pp. 47-52, figs. 40-45, Apr. 6.

1937b. A new subspecies of the nymphalid butterfly Polygonia faunus. Proc.
U. S. Nat. Mus., vol. 84, pp. 219-222, pl. 10, figs. 1-8, Apr. 9.

1937c. The butterflies of Virginia. (Abstract of a paper read before the
Entomological Society of Washington.) Proc. Ent. Soc. Wash-
ington, vol. 39, No. 5, p. 116, May.

1937d. Surveying the butterflies of Virginia. Sci. Monthly, vol. 43, No. 3,
pp. 256-265, September.

1938a. The butterflies of Virginia. Expl. and Field Work of the Smith-
sonian Inst. in 1937, pp. 77-80, figs. 80-82, Apr. 12.

1938b. [Note on Holland’s figure of Danais plexippus; and Note on the
occurrence of Calpodes ethlius in Virginia in 1937.] Proc. Ent.
Soc. Washington, vol. 40, No. 4, p. 111, April.

1939. The butterflies of Virginia. Expl. and Field Work of the Smith-
sonian Inst. in 1938, pp. 65-68, figs. 64-68, Apr. II.

1940. Butterflies of Virginia. Expl. and Field Work of the Smithsonian
Inst. in 1930, pp. 63-66, figs. 68-73, Apr. 5.

1941a. Butterflies of Virginia. Expl. and Field Work of the Smithsonian
Inst. in 1940, pp. 57-60, figs. 59-62, Apr. 2.

1941b. Notes on North and Middle American danaid butterflies. Proc.
U. S. Nat. Mus., vol. 90, pp. 531-542, pls. 71-74, Nov. 4.

1948a. [Butterflies of Virginia. Presidential address before the Entomo-
logical Society of Washington.] Proc. Ent. Soc. Washington,
vol. 50, No. 3, pp. 74-76, March.

1948b. A new subspecies of Glaucopsyche lygdamus. Proc. Ent. Soc. Wash-
ington, vol. 50, No. 7, pp. 176-178, October.

1950. Butterfly flyways and playgrounds. Lepidopterists’ News, vol. 4,
Nos. 1-2, p. 13, May.

Crark, AusTIN H., and Crark, Letra F.

1936a. Some butterflies from eastern Virginia. Journ, Washington Acad.
Sci., vol. 26, No. 2, pp. 66-70, figs. 1-14, Feb. 15.

1937. Preliminary list of the butterflies of Virginia. Proc. Biol. Soc. Wash-
ington, vol. 50, pp. 87-92, June 22.

1938a. Butterflies from Virginia and the District of Columbia. Proc. Biol.
Soc. Washington, vol. 51, pp. 1-6, Feb. 18.

1938b. Notes on Virginia butterflies. Proc. Biol. Soc. Washington, vol. 51,
pp. 177-182, Nov. 3.

1939a. Butterflies of a wood road at Suffolk, Virginia. Ent. News, vol. 50,
No. 1, pp. 1-5, January.

1939b. Butterflies from Virginia. Proc. Biol. Soc. Washington, vol. 52,
pp. 177-184, Dec. 15.

Crark, AustIN H., and TRAINER, FRANK W.

1941. Butterflies of Farmville, Virginia. Journ. Washington Acad. Sci.,
vol. 31, No. 1, pp. 38-40, Jan. 15.

Ciark, AustIN H., and WitiiaMs, CarroLt M.

1937. Records of Argynnis diana and some other butterflies from Vir-
ginia. Journ. Washington Acad. Sci., vol. 27, No. 5, pp. 209-213,
May I5.

I9gO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Comstock, WILLIAM P.

1943. The genus Ascia in the Antilles (Lepidoptera, Pieridae). Amer.

Mus. Nov., No. 1229, pp. 1-7, May 5.
CRAMER, PIETER.

1775-1782. De uitlandsche Kapellen, &c . . . . Papillons exotiques des trois
parties du Monde, Il’Asie, l’Afrique et l’Amérique, &c. 4 vols.
Amsterdam and Utrecht.

DeEnToN, S. F.

1898. Moths and butterflies of the United States east of the Rocky Moun-

tains. Boston. (Original Virginia records in sections 4, 5, 6, 7, 8.)
Dotiey, Wo. L., Jr.

1916-1917. The rate of locomotion of Vanessa antiopa (Lep.) in different
luminous intensities and its bearing on the “continuous action”
theory of orientation. Anat. Rec., vol. 11, August 1916-January
1917; Proc. Amer. Soc. Zool., No. 6, p. 519, Jan. 20, 1917; Ent.
News, vol. 28, No. 2, p. 83, February 1917.

pos Passos, Cyrit F.

1940. A new subspecies of Erora laeta from Arizona and New Mexico
(Rhopalocera: Lycaenidae). Amer. Mus. Nov., No. 1052, pp. 1, 2,
Mar. 15.

pos Passos, Cyriz F., and Grey, LioneLt PAUL.

1947. Systematic catalogue of Speyeria (Lepidoptera, Nymphalidae) with
designations of types and fixations of type localities. Amer. Mus.
Nov., No. 1370, pp. 1-30, December 12.

Epwarps, W. H.

1864. Description of the female of Argynnis diana. Proc. Ent. Soc. Phila-
delphia, vol. 3, pp. 431-434.

1877. Catalogue of the diurnal Lepidoptera north of Mexico. Trans.
Amer. Ent. Soc., vol. 6, pp. 1-68.

1884. Revised catalogue of the diurnal Lepidoptera of North America
north of Mexico. Trans. Amer. Ent. Soc., vol. 11, pp. 245-337.

1868-1897. The butterflies of North America, Ist. ser., 52 colored pls.
with descriptive letterpress, Philadelphia, 1868-1872; 2d ser., 51
colored pls. with descriptive letterpress, Boston and New York,
1884; 3d ser., 51 colored pls. with letterpress, Boston and New
York, [1887]-1897.

Emmons, EBENEZER.

1854. The natural history of New York..... , vol. 5, pp. 198-256, pls.

35-47-
FALeEs, JoHN H.

1949. Monarch butterfly migrating in Botetourt County, Virginia. Ent.

News, vol. 60, p. 68, March.
Fazzini, LILt1AN DaAvips.
1934. Butterflies and moths of America, pp. 1-96, 64 colored figs. Racine,

Wis.
Frencu, G. H.
1886. The butterflies of the eastern United States, pp. 1-402, figs. Phila-
delphia.

GrIRAULT, A. A.
1912. Fragments on North American insects.—I. Ent. News, vol. 23, No. 9,
Pp. 399-411, November.

NO. 7 BUTTERFLIES OF VIRGINIA—-CLARK IgI

1915a. Fragments on North American insects—VIII. Ent. News, vol. 26,
No. 3, pp. 127-133, March.

1915b. Fragments on North American insects—IX. Ent. News, vol. 26,
No. 5, pp. 219-227, May.

GROSSBECK, JOHN A.

1909. [Notice of a mecting of the Newark Entomological Society on

November 8, 1908.] Ent. News, vol. 20, No. 1, pp. 47-48, January.
Grote, Auc. R., and Roprnson, CoLEMAN T.

1867-1868. Descriptions of American Lepidoptera, No. 1. Trans. Amer.
Ent. Soc., vol. I, pp. 1-30, June 1867; No. 2, pp. 171-192, August
1867; No. 3, pp. 323-360, January 1868.

Harris, LucIEN, JR.

1950. The butterflies of Georgia. [Revised.] Georgia Soc. Naturalists

Bull. No. 5, pp. i-vii, 1-33, August 16.
Haypon, STANSBURY.

1933. The Papilionidae of Maryland. Proc. Nat. Hist. Soc. Maryland,
pp. 1-14, Mar. 1.

1934. The Satyridae of Maryland. Proc. Nat. Hist. Soc. Maryland, pp. 1-10,
Feb. 1.

Hewes, LAuURENCE ILSLEY.

1936. Butterflies—try and get them. Nat. Geogr. Mag., vol. 69, No. 5,

pp. 667-678, May.
Hotranp, W, J.

1898. The butterfly book, pp. i-xx, 1-382, pls. 1-48 (all colored). New
York. [Rev. ed., pp. i-xii, 1-424, pls. 1-77 (1-73 colored). New
York, 1931.]

Hovanitz, WILLIAM.

1950. The biology of Colias butterflies, I: The distribution of the North
American species. Wasmann Journ. Biol., vol. 8, No. 1, pp. 40-75,
8 figs. (maps), Spring number.

Jones, FRANK Morton.

1897. [Note on Pamphila aaroni.] Ent. News, vol. 8, No. 3, p. 60, March.

1909. Thecla damon Cram., and its varieties. Ent. News, vol. 20, No. 9,
p. 397, November.

1926. The rediscovery of “Hesperia bulenta’ Bdl.-Lec., with notes on
other species (Lepid. Hesperiidae). Ent. News, vol. 37, No. 7,
pp. 193-198, pl. 9 facing p. 193, July.

Ktots, ALEXANDER B.

1951. A field guide to the butterflies, pp. i-xvi, 1-349, 40 pls. (14 in color).

Boston.
LinpseEy, A. W.

1942. A preliminary revision of Hesperia. Denison Univ. Bull., Journ. Sci.

Laboratories, vol. 37, pp. 1-50, 6 pls., April.
Linpsey, A. W., Bett, E. L., and WittiaMs, R. C., Jr.
1931. The Hesperioidea of North America. Denison Univ. Bull., Journ.
Sci. Laboratories, vol. 26, pp. 1-142, pls. 1-32, April.
Lutz, Frank E.
1918. Field book of insects, pp. i-ix, 1-562, 101 pls. (24 colored).
Morrts, JoHN G.

1860. Catalogue of the described Lepidoptera of North America. Smith-

sonian Misc. Coll., vol. 3, pt. 2, pp. i-viii, 1-68, May.

192 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116

Pickens, A. L.

1941. Population recovery in two insect species. Meridian States Research,

Paducah [Ky.] Junior College, vol. 4, pp. 118, 119, Autumn number.
PoLLaArD, CHARLES L.

1910. [Notes on some butterflies from Virginia Beach, Va.] Journ. New

York Ent. Soc., vol. 18, No. 2, p. 129, June.
Prrer, Hi. 1b.

1909. Fighting the insect pests and diseases of orchard, field, and garden
crops. Virginia Agr. Exp. Stat. Circ. No. 7, pp. 1-148, figs. 1-65,
March. [Rev. ed., March 1g10.]

REAKIRT, TRYON.

1865. Descriptions of some new Species of Eresia. Proc. Ent. Soc. Phila-

delphia, vol. 5, pp. 224-227, December.
Ritey, C. V., and Howarp, L. O.

1893. Unusual abundance of butterfly larvae. Insect Life, vol. 5, No. 3,
p. 207, January.

RotTHSscHILD, Hon. WALTER, and JorDAN, Kart.

1906. A revision of the American Papilios. Nov. Zool., vol. 13, No. 3,
pp. 411-744, pls. 4-9.

ScHwarz, Mr. [E. A.].
1889. [Note on Pieris rapae.] Insect Life, vol. 2, No. 4, p. 123, October.
ScuppER, SAMUEL HUBBARD.

1889. The butterflies of the eastern United States and Canada, with spe-
cial reference to New England, vol. 1, pp. i-xxiv, 1-766; vol. 2,
pp. 1-x, 767-1774; vol. 3, pp. i-vi, 1775-1958, pls. 1-89, 3 maps.

Seitz, A.

1924. The Macrolepidoptera of the world, vol. 5, The American Rhopalo-

cera, pp. I-1139, 203 pls. Stuttgart.
SHOWALTER, WILLIAM JOSEPH.

1927. Strange habits of familiar moths and butterflies. Nat. Geogr. Mag.,

vol. 52, No. 1, pp. 77-126, 16 color pls. and many halftones, July.
SKINNER, HEnry.

1917. Lycaena lygdamus Doubleday and its races, with a description of
a new one. Ent. News, vol. 28, No. 5, pp. 212-214, May.

1921. Moths collected at Hot Springs, Virginia (Lepid.). Ent. News, vol. 32,
No. 3, pp. 65-71, March.

SmitrH, JAMES EDWARD.

1797. The natural history of the rarer lepidopterous insects of Georgia,
including their systematic characters, the particulars of their sev-
eral metamorphoses, and the plants on which they feed, collected
from the observations of Mr. John Abbot, many years resident of
that country, vol. I, pp. i-xv, I-102, pls. 1-51; vol. 2, pp. 103-214,
pls. 52-104. London.

SmytTH, Ettison A., Jr.

1895. Some entomological notes from Montgomery County, Virginia. Ent.
News, vol. 6, No. 8, pp. 243-245, October.

1896. Calephelis borealis. Psyche, vol. 7, No. 243, p. 403, July.

1900. Anthocharis genutia. Ent. News, vol. 11, No. 5, pp. 465-468, May.

1907. Note on Thecla damon. Ent. News, vol. 18, No. 8, p. 364, October.

1908. Two freaks—Papilio ajax and Eudamus tityrus. Ent. News, vol. 19,
No. 5, pp. 191-192, pl. 10, May.

NO: 7 BUTTERFLIES OF VIRGINIA—CLARK 193

1916. Color phases in Argynnis diana. Ent. News, vol. 27, No. 3, pp. 136-
137, March.
SmytTH, Etttson A., IV.
1938. A preliminary list of the butterflies of Rockbridge County, Virginia.
The Raven, vol. 9, No. 7, pp. 56-58, July.
STRECKER, HERMAN.
1878. Butterflies and moths of North America, pp. i-ii, 1-283.
WaGNER, WARREN HERBERT, JR.
1941. District of Columbia butterfly notes (Lepidoptera: Rhopalocera).
Ent. News, vol. 53, pp. 196-200, July ; pp. 245-249, November.
WEED, CLARENCE M.
1923. Butterflies worth knowing, pp. i-xiii, 1-286, 43 pls. (27 colored).
New York.
Woop, Carrotu E., Jr., and GotrscHALK, CarL W.
1942. The butterflies of Roanoke and Montgomery Counties, Virginia
(Lepid. Rhopalocera). Ent. News, vol. 53, pp. 143-146, 159-164,
IQI-197.
Woop, WILLIAM C.
1916. Argynnis diana. Ent. News, vol. 27, No. 1, p. 35, January.
WRIGHT, WILLIAM GREENWOOD.
1906. The butterflies of the west coast of the United States, pp. 1-250, i-vii,
pls. 1-32. San Bernardino, Calif.

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EXPLANATION OF FRONTISPIECE

(Photograph by Frederick M. Bayer.)

(All figures natural size.)

Boloria selene marilandica (A. H. Clark), under side, near Landover, Md.,
W. H. Wagner, Jr., July 15, 1941.

Boloria selene marilandica (A. H. Clark), upper side, type specimen, Belts-
ville, Md., A. H. Clark, July 6, 1920.

Polygonia faunus smythi A. H. Clark, upper side, Hardscrabble, Highland
County, Va., A. H. Clark, July 26, 1929.

Glaucopsyche lygdamus nittanyensis F. H. Chermock, upper side, Ice Moun-
tain, W. Va., A. H. Clark, April 19, 1938.

Glaucopsyche lygdamus nittanyensis F. H. Chermock, under side, Forks of
Cacapon, W. Va., W. H. Wagner, Jr., April 16, 1941.

Polygonia faunus smythi A. H. Clark, under side, White Top Mountain,
A. H. Clark, August 6, 1940.

Strymon cecrops ab. gottschalki A. H. Clark, under side, type specimen,
Fort Lewis, Roanoke County, Va., Carl W. Gottschalk, August I1, 1937.

Erora laeta (W. H. Edwards), under side, Mountain Lake, Giles County,
Va., Lorus J. and M. J. Milne, June 23, 1938.

Melitaea ismeria Boisduval and LeConte, under side, Old Fort Mountain,
Ga., Lucien Harris, Jr., May 2, 1950.

Atlides halesus (Cramer), under side, Nashville, Tenn. (Barnes Collection.)

Poanes massassoit hughi A. H. Clark, under side, Hyattsville, Md., W. H.
Wagner, Jr., July 2, 1941.

195

PLAT Es

PLATE I
(All figures reduced by one-third)

Lethe creola, male, western edge of the Dismal Swamp about 8 miles south
of Suffolk, September 1, 1935.

Lethe creola, male, under side, Dismal Swamp near Suffolk, September 3,
1936.

. Lethe creola, female, Dismal Swamp near Suffolk, September 3, 1936.
. Lethe creola, female, under side of the specimen shown in figure c.

Lethe portlandia portlandia, female, Princess Anne, September 24, 1934.
Lethe portlandia portlandia, female, under side of the specimen shown in
figure e.

g. Lethe portlandia anthedon, type specimen, Lava, Sullivan County, N. Y.

. Lethe portlandia anthedon, under side of the specimen shown in figure g.

Lethe eurydice, Beltsville, Md., July 17, 1920.
Lethe eurydice, under side of the specimen shown in figure 1.

198

VOL. 116, NO. 7, PL. 1

SMITHSONIAN MISCELLANEOUS COLLECTIONS

BUTTERFLIES OF VIRGINIA

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, PL. 2

BUTTERFLIES OF VIRGINIA

| PLATE 2
|
| (AIL figures reduced by one-third)

a. Limenitis arthemis astyanax, female, white-banded form, Nelson County,
Col. Wirt Robinson, July 23, 1910.

b. Nymphalis antiopa, female, Cabin John, Md., September 1, 1920.

c. Speyeria cybele, male, under side.

d. Zerene caesoma, Cotulla, Tex.

e. Eurema nicippe, male, Kansas.

f. Minois pegala alope, male, Sitka Church, Md., July 4, 1020.

g. Minois pegala alope, female, Beltsville, Md., July 23, 1928.

h. Minois pegala pegala, male, Fort Macon, N. C., August 4, 1920.

i. Minois pegala pegala, female, Fort Macon, N. C., August 4, 1920.

=

199

b.

k.

Euptychia

September 3, 1935.

Euptychia
Euptychia

May 30,
Euptychia
Euptychia
Euptychia
Euptychia
Euptychia
Euptycha

1935.

Ancyloxypha numitor, Newtonville, Mass., August 17, 1923.
Euphydryas phaéton, male, Cabin John, Md., June 20, 1926.

sosybius, male, under side, Florida.
Amblyscirtes textor, under side, Dismal Swamp near Suffolk, September 1,

PLATE 3
(All figures natural size)

areolatus septentrionalis, under side, 8 miles south of Suffolk,

areolatus areolatus, under side, Florida.

areolatus septentrionalis, under side, 8 miles south of Suffolk,
1037.

areolatus areolatus, under side, Texas.

cymela, female, Cabin John, Md., July 18, 1926.

cymela, female, under side of the specimen shown in figure e.
gemma, Florida.

gemma, under side of the specimen shown in figure g.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 116; NOP 7. PE:

BUTTERFLIES OF VIRGINIA

VOL. 116, NO. 7, PL. 4

MISCELLANEOUS COLLECTIONS

SMITHSONIAN

BUTTERFLIES OF VIRGINIA

SS hee SE Sot etna)

PLATE 4
(All figures reduced by one-third)

Asterocampa clyton, female, dark form, Cabin John, Md., September 8, 1926.

Asterocampa clyton, female, light form, Cabin John, Md., September 11, 1926.

Asterocampa celtis, female, Cabin John, Md., September 11, 1926.

Vanessa cardui, female, under side, Essex, Mass., August 4, 1924.

Vanessa cardui, female, Cabin John, Md., June 25, 19260.

Vanessa virginiensis, female, Cabin John, Md., September 11, 1926.

Vanessa virginiensis, female, under side, Cabin John, Md., September 11,
1920.

. Junonia evarete coenia, dry form, under side.

Junonia evarete coenia, intermediate form, under side.
Junonia evarete coenia, intermediate form approaching the wet form, under
side.

201

PLATE 5
(All figures natural size)

Polygomia faunus smythi, White Top Mountain, July 10, 1936.

Polygomia faunus smythi, under side of the specimen shown in figure a.

Polygonia progne, male, Center, N. Y., September 24, 1872.

Polygonia progne, male, under side of the specimen shown in figure c.

Polygoma comma, female, light form, Cabin John, Md., October 21, 19209.

Polygonia comma, female, under side of the specimen shown in figure e.

Junonia evarete coenia, female, wet form, Cabin John, Md., September 27,
1925.

Junonia evarete coenia, female, wet form, under side, Cabin John, Md., Sep-
tember 27, 10925.

202

SMITHSONIAN MISCELLANEOUS COLLECTIONS

BUTTERFLIES OF VIRGINIA

7, PENG

NO.

116,

VOL.

COLLECTIONS

CELLANEOUS

MIS

ONIAN

SMITHS

BUTTERFLIES OF VIRGINIA

PLATE 6
(All figures reduced by one-third)

Euptoteta claudia, female, Silver Spring, Md., August 1, 1927.

Euptoicta claudia, female, under side of the specimen shown in figure a.

Limenitis arthemis astyanax, male, under side, Silver Spring, Md., Sep-
tember 11, 1928 (upper side in figure /1).

Limenitis archippus floridensis, Miakka City, Fla., July 1900.

Limenitis archippus floridensis, under side of the specimen shown in figure d
(faint traces of the pattern of L. a. astyanax are seen in the cells of both
wings).

Limenitis archippus archippus, Decatur, Ill.

g. Limenitis arthemis astyanax, male, white-banded form, Nelson County, Col.

h.

Wirt Robinson.
Limenitis arthemis astyanax, male, Silver Spring, Md., September 11, 1928
(under side of this specimen shown in figure c).

203

PLATE 7
(All figures half natural size)

Speyeria diana, male, Spring Grove, Surry County, June 15, 1938.
Speyeria diana, male, under side of the specimen shown in figure a.
Speyeria diana, female.

Speyeria diana, female, under side of the specimen shown in figure c.
Agraulis vanillae ngrior, Fernandina, Fla., August 7, 1920.

Agraulis vanillae nigrior, under side of the specimen shown in figure e.

. Speyeria cybele, female, Rock Creek Park, Washington, D. C., June 23, 1923.
. Speyeria cybele, female, under side of the specimen shown in figure g.
Speyeria idalia, female, Cabin John, Md., July 17, 1926.

Speveria idalia, male, Silver Spring, Md., June 16, 19209.

204

VOL. 116, NO. 7, PL. 7

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116, NO.

VOL.

ITHSONIAN MISCELLANEOUS COLLECTIONS

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}

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BUTTERFLIES OF VIRGINIA

Saas S

PLATE 8
(All figures reduced by one-third)

Speyeria aphrodite, Buckeye, Highland County, July 27, 1930.

Speveria aphrodite, under side of the specimen shown in figure a.

Spevyeria atlantis, Middle Mountain, Highland County, June 19, 1948.

Speyeria atlantis, under side of the specimen shown in figure c.

Minois pegala maritima, male, Rock Creek Park, Washington, D. C., W. H.
Waener, July 6, 1935.

Vanessa atalanta, female, wet form, Cabin John, Md., August 27, 1926.

Ascia phileta, dark female.

Ascia phileta, dark female, under side of the specimen shown in figure g.

Ascia phileta, male, Cochise County, Ariz., July.

Ascia phileta, male, under side of the specimen shown in figure 1.

to
=)
OV

Q Ae aoe

. Boloria selene myrina, Essex, Mass., August 12, 1925.
. Boloria selene marilandica, Beltsville, Md., July 6, 1920.

. Boloria toddi ammiralis, Essex, Mass., August 30, 1925.

. Melitaea nycteis, under side of the specimen shown in figure f.
. Eurema lisa, male.

PLATE Q

(All figures natural size)

Boloria selene marilandica, under side of the specimen shown in figure b.
Boloria toddi ammuralis, under side of the specimen shown in figure d.

Melitaea nycteis, Lunenburg, September 2, 1936.

Anthocharis genutia, male.
Anthocharis genutia, female, Cabin John, Md., May 2, 1926.

VOL. 116,

SMITHSONIAN MISCELLANEOUS COLLECTIONS

BUTTERFLIES OF VIRGINIA

SMITHSONIAN MISCE

LLANEOUS COLLECTIONS

VOL.

116,

ee)

BUTTERFLIES OF VIRGINIA

PLATE I0
(All figures reduced by one-third)

Danaus plexippus plexippus, female.

. Danaus plexippus megalippe, female, British Guiana.

Phoebis sennae, male.

. Phoebis philea, male.

Colias eurytheme, female, Cabin John, Md., September 14, 1928.

207

PLATE II
(Figures n, 0, X 14, others natural size)

Atrytone arogos, St. Petersburg, Fla.

Atrytone arogos, under side of the specimen shown in figure a.

Lerodea Vherminiert.

Lerodea lV’herminieri, under side of the specimen shown in figure c.

Atrytone ruricola metacomet, male, Jamesburg, N. J., August 4, 1907.

Phyciodes batesti, male, Albany, N. Y., June 6, 1876.

Phyciodes batesii, male, under side of the specimen shown in figure f.

Phyciodes batesti, female, Apple Orchard Mountain, 3,200 feet, Carroll E.
Wood, Jr., June 1, 1938.

Phyciodes batesti, female, under side of the specimen shown in figure h.

Phyciodes tharos, male, Silver Spring, Md., July 7, 1928.

Phyciodes tharos, female, under side, Cabin John, Md., May 10, 1925.

Phyciodes tharos, female, upper side of the specimen shown in figure k.

. Phyciodes tharos, under side.

Strymon melinus melinus, Florida.
Strymon melinus humuli, Apple Orchard Mountain, over 4,000 feet, July 24,
1934.

208

11

re

NO. 7,

116,

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SMITHSONIAN MISCELLANEOUS COLLECTIONS

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BUTTERFLIES OF VIRGINIA

PLATE 12
(Figures c, d, natural size, others « 14)

Erora laeta, Mountain Lake, Giles County, Lorus J. Milne, June 23, 10938.
Erora laeta, under side of the specimen shown in figure a.

Atlides halesus, male, Florida.

Atlides halesus, male, under side of the specimen shown in figure c.
Eupsyche m-album, male, Silver Spring, Md., September 2, 1927.
Eupsyche m-album, male, under side of the specimen shown in figure e.
Mitoura gryneus, male, under side.

Strymon cecrops, under side.

Calephelis borealis, Kerrville, Tex.

Calephelis virginiensis, Miami, Fla.

200

=a

PLATE 13
(All figures & 14)

1. Incisalia polios, under side, Digby, Nova Scotia.

. Incisalia mphon niphon, under side, Washington, D. C., W. H. Wagner,

April 26, 1938.

. Strymon falacer, male, under side, Essex, Mass., July 9, 1925.

. Strymon edwardsii, female, under side, Kerrville, Tex.

Strymon liparops, under side, Essex, Mass., July 7, 1925.

Strymon ontario, female, under side, Kirkwood, Mo., Mary E. Murtfeldt.

. Strymon titus titus, female, under side, New York.

Strymon titus mopsus, female, under side, Washington, D. C., W. H. Wagner,
June 24, 1934.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, PL. 13

BUTTERFLIES OF VIRGINIA

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, PL. 14

BUTTERFLIES OF VIRGINIA

aoc,

PLATE 14
(All figures & 14)

Cyaniris argiolus pseudargiolus, male, early spring form.

Cyaniris argiolus pseudargiolus, female, early spring form.

Cyaniris argiolus pseudargiolus, male, late spring form.

Cyaniris argiolus pseudargiolus, female, late spring form.

Cyaniris argiolus pseudargiolus, male, summer form, Beltsville, Md., July 9,
1928.

Cyaniris argiolus pseudargiolus, female, summer form, Cabin John, Md.,
June 16, 1920.

Glaucopsyche lygdamus mittanyensis, male, Ice Mountain, W. Va., May 7,
1939.

Cyaniris argiolus pseudargiolus, female, under side of the specimen shown in
figure f.

Everes comyntas, male, Cabin John, Md., June 10, 1928.

Everes comyntas, male, under side of the specimen shown in figure 7.

>

PLATE I5
(Figures a, b, c, 1, m,n, X 14, others natural size)

Glaucopsyche lygdamus couperi, under side, Chicago, May 20, 1917.

Glaucopsyche lygdamus lygdamus, female, under side, Georgia.

Glaucopsyche lygdamus mittanyensis, under side, Forks of Cacapon, W. Va.,
May 8, 1030.

Polites manataaqua, left side male, right side female, Washington, D. C.,
W. H. Wagner, August 12, 1935.

Feniseca tarquinius, larva from Weston, Mass., emerged at Washington,
D. C., November 19, 1923.

Lycaena thoé, female, Essex, Mass., September I, 1925.

. Lycaena thoé, male, Detroit, Mich., G. W. Rawson, June 7, 1927.
. Lycaena thoé, male, under side of the specimen shown in figure g.

Pholisora hayhurstu, male, Texas.

Pholisora hayhurstu, male, under side of the specimen shown in figure 1.

Feniseca tarquinius, larva from Weston, Mass., emerged at Washington,
D. C., August 22, 1923.

Lycaena phlaeas americana, male, Beltsville, Md., September 7, 19209.

. Lycaena phlaeas americana, male, under side of the specimen shown in fig-

ure 1.

. Lycaena phlaeas americana, female, Rock Creek Park, Washington, D. C.,

May 6, 1923.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS NOES 116,;) NOle7, Rio

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, PL. 16

BUTTERFLIES OF VIRGINIA

2s Rose

PLATE 16
(Figures c, d, ce, X 14, others natural size)

Euchloé olympia, west of Cross Junction, Frederick County, April 24, 1938.

Euchloé olympia, under side of the specimen shown in figure a.

Incitsalia irus, Washington, D. C., W. H. Wagner, April 26, 1938.

Incisalia henrici, west of Cross Junction, Frederick County, April 24, 1928.

Incisahia augustinus croesioides, under side, Difficult Run, Fairfax County,
April 29, 1931.

Polygonia interrogationis.

Nathalis iole, Brownsville, Tex.

Nathalis iole, under side of the specimen shown in figure g.

Eurema jucunda, Glenwood, Fla.

Eurema jucunda, under side of the specimen shown in figure 7.

Qs

=~
=

PLATE 17
(All figures natural size)

Colias philodice, male, Silver Spring, Md., August 3, 1927.

Colias philodice, male, under side.

Colias interior, male, Middle Mountain, Highland County, John E. Graf,
June 14, 1936.

Colias interior, male, under side of the specimen shown in figure c.

Colias philodice, female, Cabin John, Md., May 9, 1926.

Colias curytheme, female, under side.

Colias interior, female, Nepigon, Ontario.

Colias interior, female, under side of the specimen shown in figure g.

214

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BUTTERFLIES OF VIRGINIA

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7,

BUTTERFLIES OF VIRGINIA

b.

h.

PLATE 18
(All figures natural size)

Pieris rapae, male, spring form, Cabin John, Md., April 8, 1925.

Pieris virginiensis, under side, west of Cross Junction, Frederick County,
April 24, 1938.

Pieris protodice, female, spring form, Paint Branch, Md.

Pieris protodice, female, spring form, under side of the specimen shown in
figure c.

Pieris protodice, male, summer form, Cabin John, Md., July 25, 1926.

Pieris protodice, male, summer form, under side of the specimen shown in
figure e.

Pieris protodice, female, summer form, Cabin John, Md., September 27, 1925.

Pieris protodice, female, summer form, under side of the specimen shown in
figure g.

215

QS RP ee

=

=

PLATE 19
(All figures half natural size)

Battus philenor, male, Cabin John, Md., September 10, 1025.

Battus philenor, male, under side of the specimen shown in figure a.

Battus philenor, female, Cabin John, Md., September 19, 1925.

Battus philenor, male, early spring form.

Papilio cresphontes, Decatur, Il.

Papilio glaucus, female, dark form.

Graphium marcellus, female, spring form, Great Falls, Md., May 2, 1926.
Graphium marcellus, female, summer form, Cabin John, Md., July 20, 1928.

216

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116,

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BUTTERFLIES OF VIRGINIA

VOL. 116, NO. 7, PL. 20

ION

JUS COLLECT

NEC

MISCELLA

)NIAN

SMITHS

BUTTERFLIES OF VIRGINIA

Teawhe aoses

Papilio
Papilio
Papilio
Papilio
Papilio
Papilio
Papilio
Papilio

PLATE 20
(All figures half natural size)

glaucus, male, Newfoundland.

glaucus, male, under side of the specimen shown in figure a.
glaucus, male, Essex, Mass., July 13, 1925.

glaucus, male, Cabin John, Md., August 22, 1926.

glaucus, female, Essex, Mass., July 26, 1925.

glaucus, female, under side of the specimen shown in figure e.

glaucus, female, Silver Spring, Md., August 6, 1927.
glaucus, male, under side of the specimen shown in figure d.

bo

NI

h.

PLATE 21
(All figures half natural size)

Papilio glaucus, female, Newfoundland.

. Papilio glaucus, female, under side of the specimen shown in figure a.

Papilio glaucus, female, dark form, Newfoundland.

Papilio glaucus, female, dark form, spring type, Great Falls, Md., W. H.
Wagner, April 24, 1938.

Papilio glaucus, female, dark form, spring type, under side of the specimen
shown in figure d.

Papilio glaucus, female, dark form with the outer portion of the wings
thickly sprinkled with yellow scales, Washington, D. C.

Papilio glaucus, female, dark form with the outer portion of the wings
thickly sprinkled with yellow scales, under side of the specimen shown in
figure f.

Papilio glaucus, female, with the inner portion of the wings thickly sprinkled
with dark scales, Silver Spring, Md., August 3, 1927.

218

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BUTTERFLIES OF VIRGINIA

PL. 22

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VOL. 116, NO.

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BUTTERFLIES OF VIRGINIA

PLATE 22
(All figures half natural size )

Papilio troilus, male, Manassas, August 18, 1035.

Papilio troilus, female, Silver Spring, Md., August 6, 1927.

Papilio palamedes, male, Virginia Beach, William Schaus.

Papilio palamedes, male, under side of the specimen shown in figure c.

Papilio polyxenes asterius, male, with an unusually broad yellow band, Silver
Spring, Md., August 14, 1927.

Papilio polyxenes asterius, male, with a narrow yellow band, Silver Spring,
Md., July 24, 1927.

Papilio polyxenes asterius, female, Silver Spring, Md., July 21, 1927.

Papilio polyxenes asterius, female, with patches of the male coloration, es-
pecially on the left hind wing, Nelson County, Col. Wirt Robinson.

210

ota!

PLATE 23
(All figures natural size)

Proteides clarus, male, under side, Cabin John, Md., June 1, 1925.

. Rhabdoides cellus, near Great Falls, Md., June 3, 1934.

Rhabdoides cellus, under side of the specimen shown in figure D.
Urbanus proteus, female, Florida.

Achalarus lyciades, female, under side, Cabin John, Md., June 13, 1926.
Calpodes ethlius, Brownsville, Tex.

Thorybes bathyllus, male, Silver Spring, Md., July 3, 1928.

Thorybes bathyllus, female, Cabin John, Md., August 28, 1926.
Libytheana bachmanii, Cotulla, Tex., J. C. Crawford, May 11, 1906.

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VOL. 116, NO. 7,

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 116, NO 7, RES 24

BUTTERFLIES OF VIRGINIA

PLATE 24
(All figures natural size)

Pyrgus centaureae wyandot, Cabin John, Md., April 10, 1926.
Pyrgus communis.
Pyrgus conunumns.

Pyrgus centaureae wyandot, under side of the specimen shown in figure a.

Pyrgus communis, under side of the specimen shown in figure D.
Pyrgus communis, under side of the specimen shown in figure c.
Erynnis icelus, male, Cabin John, Md., May 25, 1930.

Erynnis icelus, female, Cabin John, Md., May 25, 1930.

Erynnis brizo, male, Albany, N. Y., May 19, 1870.

Erynnis persius, male.

Erynnis lucilius, male.

Erynnis persius, female.

Erynnis martialis, female, June 3, 1870.
Erynnis juvenalis, male.

Erynnis juvenalis, female.

Lerodea cufala, Brownsville, Tex.

. Erynnis baptisiae, male, Camp Letts, Md., W. H. Wagner, July 9, 1941.

SOP APN SS

ctrl ea nee

—

PLATE 25
(All figures natural size)

Thorybes pylades, female, Cabin John, Md., June 6, 1926.

Thorybes pylades, female, under side of the specimen shown in figure a.
Thorybes confusis, Difficult Run, Fairfax County, June 16, 1935.
Thorybes confusis, under side of the specimen shown in figure c.
Thorybes bathyllus, female, under side, Cabin John, Md., August 28, 1926.
Erynnis juvenalis, female, under side.

. Erynnts sarucco, male.
. Erynnis zarucco, male, under side of the specimen shown in figure g.

Amblyscirtes hegon.
Erynnis horatius, male, Miami, Fla.

. Erynnis horatius, female.

Amblyscirtes alternata, Suffolk, Otto Buchholz, May 9, 1945.

m. Amblyscirtes alternata, under side of the specimen shown in figure J.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 116, NO. 7, PL. 25

BUTTERFLIES OF VIRGINIA

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MISCELLANEOUS COLLECTIONS

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BUTTERFLIES OF VIRGINIA

maps

PLATE 26
(All figures natural size)

Hesperia leonardus, male, Silver Spring, Md., September 11, 1928.

Hesperia leonardus, female, under side, Silver Spring, Md., September 14,
1928.

Poanes massassoit hughi, male, Beltsville, Md., July 15, 1928.

. Poanes massassoit hughi, female, under side, Beltsville, Md., July 15, 1928.

Atrytone ruricola metacomet, male, under side, Jamesburg, N. J., August 4,
1907.

Atalopedes campestris, female, dark form, under side, Manassas, August 25,
1935.

Atalopedes campestris, male, Silver Spring, Md., August 13, 1927.

Atalopedes campestris, female, Silver Spring, Md., July 28, 1928.

Atalopedes campestris, female, under side of the specimen shown in figure h.

Polites mystic, male, Middle Mountain, Highland County, June 19, 1948.

Polites mystic, male, under side of the specimen shown in figure /.

Panoquina panoquin, Anglesea, N. J., June 1923.

. Wallengrenia otho egeremet, male, Aylett, July 18, 1937.

Wallengrenia otho egeremet, male, under side of the specimen shown in fig-
ure m.

Wallengrenia otho otho, under side, Florida.

Panoquina ocola.

Atrytonopsis hianna, female, Cabin John, Md., June 2, 1920.

Pholisora catullus, female, Onaga, Kans.

PLATE 27
(All figures natural size)

Poanes gabulon, male, High Island, Md.

Poanes zabulon, male, under side of the specimen shown in figure a.
Atrytone logan, male, Newtonville, Mass., August 16, 1923.
Atrytone logan, male, under side of the specimen shown in figure c.
Atrytone logan, female, Beltsville, Md., July 23, 1928.

Polites vibex, male, under side, St. Petersburg, Fla.

. Atrytone conspicua, male, Beltsville, Md., July 15, 1928.
. Atrytone conspicua, male, under side, Newtonville, Mass., July 11, 1923.

Hesperia attalus, male, under side, St. Petersburg, Fla., April 24, 1914.
Atrytone bimacula, male, Albany, N. Y., June 15, 1870.

Atrytone bimacula, male, under side of the specimen shown in figure 7.
Amblyscirtes hegon, under side.

.. Hesperia sassacus, male, Middle Mountain, Highland County, June 19, 1948.
. Amblyscirtes vialis, Wellington, British Columbia, June 27, 1903.

Lerema accius, male, Indian River, Fla.

Hesperia sassacus, male, under side of the specimen shown in figure m1.
Amblyscirtes vialis, under side of the specimen shown in figure n.
Lerema accius, male, under side of the specimen shown in figure o.

224

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Se rere roe

eR Saat oe

PLATE 28
(All figures natural size)

Hesperia metea, male.
Hesperia metea, male, under side of the specimen shown in figure a.
Hylephila phyleus, male, Cabin John, Md., August 29, 1920.

. |Polites verna, male, July 14, 1907.

Polites manataaqua, male, Washington, D. C., August 18, 1883.
Polites themistocles, male, Silver Spring, Md., June 3, 1928.

. Polites peckius, female, Cabin John, Md., August 10, 1927.

Polhites peckius, female, under side of the specimen shown in figure g.
Polites vibex, male, St. Petersburg, Fla.

Poanes hobomok, female, Cabin John, Md., June 2, 1920.

Poanes hobomok, female, New Jersey.

Poanes hobomok, female, under side of the specimen shown in figure k.

. Poanes aaroni, male, New Jersey.
. Poanes aaroni, male, under side of the specimen shown in figure m1.

Poanes aaron, female, Chincoteague Island, W. H. Wagner, August 21, 1941.
Amblyscirtes carolina, Dismal Swamp near Suffolk, April 1, 1938.

Poanes yehl, male, New Bohemia, W. H. Wagner, July 10, 1941.

Poanes aaroni, female, under side of the specimen shown in figure o.

to
to
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PLATE 29
(All figures natural size)

Atrytone dukesi, male, Mobile, Ala., September 10, 1934.

Atrytone dukesi, male, under side of the specimen shown in figure a.

Atrytone dukesi, female, Mobile, Ala., September 10, 1934.

Atrytone dukesi, female, under side of the specimen shown in figure c.

Atrytone dion, male, Cape Henry, September 4, 1936.

Atrytone alabamae, male, Dahl Swamp, Accomack County, July 25, 1935.

Atrytone palatka, male, Princess Anne County, Otto Buchholz, June 16,
1041.

Atrytone palatka, male, under side of the specimen shown in figure g.

Atrytone palatka, female, Miami, Fla.

Poanes viator, female, Newark, N. J.

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Sa

"FQ THO AD

k.

PLATE 30
(Figures h, 1, X 14, others natural size)

Amblyscirtes carolina, under side, Dismal Swamp near Suffolk, April 1,
1938.

Amblyscirtes carolina var. reversa, paratype male, under side, Southern
Pines, N. C., July 28, 1911.

Amblyscirtes carolina, unmarked variety, under side, Suffolk, May 30, 10936.

Poanes yehl, male, under side, New Bohemia, W. H. Wagner, July 10, 1941.

Euphydryas phaéton, aberration, Gala, Warren Stoutamire, July 19, 1941.

Poanes yehl, female, New Bohemia, W. H. Wagner, July 109, 1941.

. Poanes yehl, female, under side, Nashville, Tenn., August 23, 1895.

Incisalia niphon clarkt.

Incisalia niphon clarki, paratype, Constance Bay, Ontario, G. S. Walley,
May 23, 1013.

Vanessa virginiensis, aberration, Gilbert Yobst, Scotts Run, Fairfax County,
May 1036.

Vanessa virginiensis, aberration, under side of the specimen shown in fig-
ure 7.

INDIES

a kT

INDEX

aaroni, Poanes, 8, 25 (in key), 170;
pl. 28, m, n, 0, r
aaroni aaroni, Poanes, 170, I71
aaroni howardi, Poanes, 170, 171
abeona, Tisiphone, 50
abeona aurelia, Tisiphone, 50, 51
abeona joanna, Tisiphone, 50
abeona morrisi, Tisiphone, 50, 51
abeona rawnsleyi, Tisiphone, 51
acauda, Battus philenor, 119, 144
accius, Lerema, 26 (in key), 178; pl.
27, 0,7
Achalarus, 148
lyciades, 22 (in key), 148; pl.
23, €
Agkistrodon piscivorus, 176
Agraulis, 64
vanillae, 1, 8, 12 (in key), 64
vanillae nigrior, 653 pl. 7, ¢, f
alabamae, Atrytone, 7, 25 (in key).
174, 175; pl. 20, f
alba, Eurema lisa form, 116
albescens, Pyrgus communis, 153
albofasciata, Limenitis arthemis, 51
alcestis, Speyeria aphrodite, 50
alexandra edwardsii, Colias, 187
alope, Minois pegala, 32, 33, 34, 35, 30;
pl. 2, f, g
alternata, Amblyscirtes, 8, 27 (in key),
179; pl. 25, 1, m
Amblyscirtes, 179
alternata, 8, 27 (in key), 1793; pl.
25, 1, m
carolina, 8, 24 (in key), 180; pls.
28, p, 30, a, b, c
carolina reyersa, 180
hegon, 6, 27 (in key), 179; pls. 25,
a er ha|
textor, 8, 26 (in key), 180; pl. 3, 7
vialis, 27 (in key), 1793 pl. 27,
n,Q
americana, Lycaena phlaeas, 71; pl.
15, J, m, 1
americana ab. fulliolus, Lycaena
phlaeas, 71
ammiralis, Boloria toddi, 61, 62, 63;
pl. 9, d, e
amphidusa, Colias eurytheme, 102, 104,
107, 108, 109, III

(Numbers in boldface are principal references.)

ampliata, Papilio polyxenes, 144
Ancyloxypha, 159
numitor, 24 (in key), 1593 pl. 3, &
numitor longleyi, 159
anthedon, Enodia portlandia, 5
Lethe portlandia, 5, 31; pl. 1, 9g, h
Anthocharis, 86
genutia, 18 (in key), 86; pl. 9, i, 7
genutia flavida, 86
antiopa, Nymphalis, 2, 12 (in key),
423; pl. 2,b
Apaturidae, 10 (in key), 12 (key to
species), 39
Apaturinae, 39
aphrodite, Speyeria, 3, 5, 15 (in key),
59; pl. 8, a, b
aphrodite alcestis, Speyeria, 59
aphrodite ab. bakeri, Speyeria, 60
aphrodite ab. hughi, Speyeria, 61
archippus, Limenitis, 13 (in key), 52
archippus archippus, Limenitis, 52, 53,
54; pl. 6, f
archippus eros, Limenitis, 54
archippus floridensis, Limenitis, 8, 52,
53, 54; pl. 6, d, e
arcticus, Papilio glaucus, 134, 137, 139,
142
Papilio rutulus, 134
areolatus, Euptychia, 12 (in key), 36
areolatus areolatus, Euptychia, 8, 36;
pl. 3,6, d
areolatus septentrionalis, 8, 37; pl. 3,
a,c
argiolus, Cyaniris, 2, 16 (in key), 73
argiolus pseudargiolus, Cyaniris, 73,
75; pl. 14, a, b,c, d, e, f,h
argiolus pseudargiolus form intermedia,
Cyaniris, 73
argiolus pseudargiolus form lucia, Cy-
aniris, 73
argiolus pseudargiolus form marginata,
Cyaniris, 73
argiolus pseudargiolus form neglecta,
Cyaniris, 73, 74
argiolus pseudargiolus form neglecta-
major, Cyaniris, 74
argiolus pseudargiolus form nigra, Cy-
aniris, 73

229

230

argiolus pseudargiolus form pseudar-
giolus, Cyaniris, 73, 74
argiolus pseudargiolus form violacea,
Cyaniris, 73
Argynnidae, 10 (in key), 12 (key to
species), 49
Argynninae, 54
ariadne, Colias eurytheme, 102
Aristolochia macrophylla, 119
arizonensis, Limenitis arthemis, 50
arogos, Atrytone, 1, 24 (in key), 171,
1729 pl. 13,.a, 2
arthemis, Limenitis, 14 (in key), 49,
50, 51, 143, 186
arthemis albofasciata, Limenitis, 51
arthemis arizonensis, Limenitis, 50
arthemis astyanax, Limenitis, 50, 51,
53; Dis. 2, a, 6,c,9,h
arthemis rubrofasciata, Limenitis, 50,
51
Arundinaria, 32
gigantea, 3, 31
tecta, 3, 30
Ascia, 93
monuste, 93
phileta, 1, 18 (in key), 19 (in
key), 93
phileta phileta, 93; pl. 8, 9, h, i, 7
Asclepias, 120
asterius, Papilio polyxenes, 1443 pl.
22, ¢,f,9,h
Asterocampa, 39
celtis, 14 (in key), 39, 40; pl. 4, c
clyton, 14 (in key), 393 pl. 4, a, b
astyanax, Limenitis arthemis, 50, 51,
53; pls. 2, a, 6, ¢, 9, h
astyanax proserpina, Limenitis, 51
atalanta, Vanessa, 2, 13 (in key), 43,
44, 45; pl. 8, f
Atalopedes, 163
campestris, 27 (in key), 162, 163;
pl. 26, f, g, h, 4
atlantis, Speyeria, 4, 15 (in key), 613
pl. 8, c,d
Atlides, 77
halesus, 8, 15 (in key), 773 fron-
tispiece, fig. 10; pl. 12, c, d
Atrytone, 172
alabamae, 7, 25 (in key), 174,
175; pl. 20, f
arogos, I, 24 (in key), 171, 1723
pl. 11, a, b
bimacula, 5, 26 (in key), 1763 pl.
27,.4,.%
conspicua, I, 5, 29 (in key), 176;
pl. 27, 9, h

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116

Atrytone dion, 8, 25 (in key), 173,
174; pl. 20, e
dukesi, 8, 25 (in key), 1753 pl.
20.00 .6.a
logan, 24 (in key), 170, 1733 pl.
D7AE CHE,
palatka, 8, 24 (in key), 1733 pl.
29, 9; h, 1
ruricola, 25 (in key), 164, 177
ruricola metacomet, 1773 pls. 11,
e, 20, e€
Atrytonopsis hianna, 6, 27 (in key),
177; pl. 26, q
attalus, Hesperia, 28 (in key), 161;
pl. 27,1
Augiades sylvanus, 1, 187
augustinus, Incisalia, 5, 17 (in key),
83, 84
augustinus croesioides, Incisalia, 83;
pl. 16, e
aurelia, Tisiphone abeona, 50, 51
ausonius, Erynnis martialis, 187

Baccharis halmifolia, 82
bachmanii, Libytheana, 68; pl. 23, 7
bakeri, Speyeria aphrodite ab., 60
baptisiae, Erynnis, 23 (in key), 156;
pl. 24, m
batesii, Phyciodes, 5, 15 (in key), 493
pl. 11, f, 9,h,4
bathyllus, Thorybes, 22 (in key), 1493
pls. 23, g, h, 25, e
Battus, 118
philenor, 21 (in key), 118, 140,
144; pl. 10, a, b,c, d
philenor acauda, 119, 144
philenor hirsuta, 119, 144
bimacula, Atrytone, 5, 26 (in key),
176; pl. 27, j, k
Boloria, 3, 61, 124
selene, 2, 3, 12 (in key), 62
selene marilandica, I, 2, 63, fron-
tispiece, figs. I, 2; pl. 9, b, c
selene myrina, 5, 62; pl. 9, a
toddi, 3, 5, 14 (in key), 64
toddi ammiralis, 61, 62, 63; pl.
0, d, e
borealis, Calephelis, 5, 15 (in key),
69, 70; pl. 12, 7
boydi, Glaucopsyche lygdamus, 72, 73
brizo, Erynnis, 23 (in key), 1553 pl.
24, 4

caesonia, Zerene, 19 (in key), 1123
pl. 2, d

NO. 7

Calephelis, 69
borealis, 5, 15 (in key), 69, 70;
pl. 12, ¢
virginiensis, 8, 15 (in key), 69;
pl. 12,7
Calpodes, 183
ethlius, 1, 26 (in key), 1833 pl.
23, f
campestris, Atalopedes, 27 (in key),
162, 163; pl. 26, f, 9, h,1
canadensis, Papilio glaucus, 133, 134,
136, 137, 139, 142
cardui, Vanessa, I, 2, 14 (in key), 44,
45; pl. 4, d, e
Carex, 3
striata var. brevis, 174
stricta, 176
carolina, Amblyscirtes, 8, 24 (in key),
180; pls. 28, p, 30, a, b, c
Minois pegala, 35
carolina reversa, Amblyscirtes, 180
Cassia chamaecrista, 113, 114
catullus, Pholisora, 22 (in key), 1533
pl. 26, r
cecrops, Strymon, 16 (in key), 78,
frontispiece, fig. 7; pl. 12, h
cecrops ab. gottschalki, Strymon, 78,
frontispiece, fig. 7
cellus, Rhabdoides, 22 (in key), 1493
Bias. 0: €
celtis, Asterocampa, 14 (in key), 39,
40; pl. 4, ¢
centaureae, Pyrgus, 5, 23 (in key), 151
centaureae wyandot, Pyrgus, 1513 pl.
24, a, d
Chelone glabra, 47
chrysotheme eurytheme, Colias, 1
Cinnamomum camphora, 125
clappi, Eureme lisa form, 116
clarki, Incisalia niphon, 83; pl. 30, h, 1
clarus, Proteides, 22 (in key), 1473
pl. 23, a
claudia, Euptoieta, 14 (in key), 60,
63; pl. 6, a, b
cleobaea, Lycorea, 187
clyton, Asterocampa, 14 (in key), 39;
pl. 4, a,b
coenia, Junonia evarete, 45; pls. 4, h,
4, a; 5; gY; h
Colias, 7, 93
alexandra edwardsii, 187
chrysotheme eurytheme, I
eogene, 100
eurytheme, 6, 18 (in key), 20 (in
key), 96, 97, 101, 102, 103, 104,
107, 108, 109, I10, III; pls. 10,
€, 17, f

INDEX

231

Colias eurytheme amphidusa, 102, 104,
107, 108, 100, III
eurytheme ariadne, 102
eurytheme eurytheme, 102
eurytheme keewaydin, 102
hecla, 105
interior, 4, 20 (in key), xxx, 112;
Pie Ine 1a. gan
nastes werdandi, 106
philodice, 5, 6, 18 (in key), 20 (in
key), 93, 97, 101, 102, 103, 106,
107, 108, 109, 110, III, 112; pl.
L7s) Qa ORG
comma, Polygonia, 13 (in key), 40;
pl. 5, e, f
communis, Pyrgus, 23 (in key), 152;
pl. 24, b, c, e, f
communis albescens, Pyrgus, 152
comyntas, Everes, 16 (in key), 48, 753
pl. 14, i,7
confusis, Thorybes, 22 (in key), 150;
plese re
conspicua, Atrytone, I, 5, 29 (in key),
176; pl. 27, 9,h
creola, Lethe, 8, 11 (in key), 30; pl.

We eles Coy te!

cresphontes, Papilio, 20 (in key), 120;
pl. 19, e

croesioides, Incisalia augustinus, 83;
pl. 16, e

Crotalus horridus, 176
curvifascia, Papilio polyxenes, 144
Cyaniris, 73
argiolus, 2, 16 (in key), 73
argiolus pseudargiolus, 73, 75; pl.
14, a, b,c, d,e,f,h
argiolus pseudargiolus form inter-
media, 73
argiolus pseudargiolus form lucia,

argiolus pseudargiolus form mar-
ginata, 73

argiolus pseudargiolus form neg-
lecta, 73, 74

argiolus pseudargiolus form neg-
lecta-major, 74

argiolus pseudargiolus form nigra,

argiolus pseudargiolus form pseud-
argiolus, 73, 74
argiolus pseudargiolus form vio-
lacea, 73
cybele, Speyeria, 3, 12 (in key), 56,
§8)'60% pis. 2) ¢;,7;70, &
cymela, Euptychia, 11 (in key), 38;
pl. 3, e, f

232 SMITHSONIAN MISCELLANEOUS COLLECTIONS

Danaidae, 10 (in key), 12 (key to

species), 65
Danainae, 65
Danaus, 65
plexippus, 2, 13 (in key), 65, 67
plexippus ab. fumosus, 65
plexippus megalippe, 2, 67, 68;
pl. 10, b
plexippus plexippus, 65, 67, 68;
pl. 10, a
Daucus carota, 145
diana, Speyeria, I, 3, 14 (in key), 55,
617 pl.7,.aybi cd
dion, Atrytone, 8, 25 (in key), 173,
174, 175; pl. 20, é@
drya, Phoebis sennae, 113
drymo, Ithomia, 187
dukesi, Atrytone, 8, 25 (in key), 1753
pl. 20, a, b, c, d

edwardsii, Colias alexandra, 187
Strymon, 6, 17 (in key), 803; pl.
heyy (a!
egeremet, Wallengrenia otho, 1673; pl.
26, m, n
elathea, Eurema, 187
Elephantopus, 180
Enodia portlandia anthedon, 5
eogene, Colias, 100
Erebia, 124
Erora, 85
laeta, 5, 17 (in key), 85, 187;
frontispiece, fig. 8; pl. 12, a, b
eros, Limenitis archippus, 54
Erynnis, 154, 159
baptisiae, 23 (in key), 156; pl.
24, m
brizo, 23 (in key), 1553 pl. 24,7
horatius, 23 (in key), 158, 159;
pl. 25, J, k
icelus, 22 (in key), 1543 pl. 24,
g,h
juvenalis, 23 (in key), 157, 158,
159; pls. 24, 0, 25, f
lucilius, 5, 23 (in key), 1553 pl.
24, Rk
martialis, 23 (in key), 156, 187;
pl. 24, n
martialis ab. ausonius, 187
persius, 5, 23 (in key), 1553 pl.

24, J, !
zarucco, 23 (in key), 158; pl. 25,
g, h
ethlius, Calpodes, 1, 26 (in key), 1833
pl. 23, f

VOL. 116
Euchloé, 87
olympia, 18 (in key), 87, 88; pl.
16, a, b

olympia rosa, 87, 88
eufala, Lerodea, 8, 26 (in key), 1823
pl. 24, q
Euphydryas, 47
phaéton, 5, 14 (in key), 473 pls.
3, L, 30, e
Euploea, 56
Eupsyche, 78
m-album, 16 (in key), 78; pl. 12,
8
Euptoieta claudia, 14 (in key), 60, 633
ple On G0
Euptychia, 36
areolatus, 12 (in key), 36
areolatus areolatus, 8, 36; pl. 3,
b, d
areolatus septentrionalis, 8, 373
ih eh ta
cymela, 11 (in key), 38; pl. 3,
é, f
gemma, 8, 12 (in key), 36, 38;
pl. 3, 9, h
sosybius, 8, 12 (in key), 38; pl.
3, 4
Eurema, 115
elathea, 187
euterpe, 116
jucunda, 19 (in key), 117, 118;
pl. 16, %,7
lisa, 20 (in key), 110, 1463 pl.
9, h
lisa form alba, 116
lisa form clappi, 116
nicippe, 20 (in key), 115; pl. 2, e
eurydice, Lethe, 11 (in key), 31

eurydice appalachia, Lethe, 32; pl.

I, 1, J
eurytheme, Colias, 6, 18 (in key), 20
(in key), 96, 97, IOI, 102, 103,
104, 107, 108, 100, I10, 111; pls.
ie ae a
Colias chrysotheme, 1
eurytheme amphidusa, Colias, 102, 104,
107, 108, 100, III
eurytheme ariadne, Colias, 102
eurytheme eurytheme, Colias, 102
eurytheme keewaydin, Colias, 102
euterpe, Eurema, 116
evarete, Junonia, 13 (in key), 45
evarete coenia, Junonia, 45; pls. 4, h,
i, J,5,9,h

NO. 7

Everes, 75
comyntas, 16 (in key), 48, 753
pl. 14, 1,7

fabricii, Polygonia interrogationis, 40
falacer, Strymon, 17 (in key), 80;
pl. 13, ¢
faunus, Polygonia, 13 (in key), 41
faunus smythi, Polygonia, 4, 42; fron-
tispiece, figs. 3, 6; pl. 5, a, b
Feniseca, 70
tarquinius, 17 (in key), 703 pl.
15, ¢, k
flavida, Anthocharis genutia, 86
floridensis, Graphium marcellus, 146
Limenitis archippus, 8, 52, 53, 54;
pl. 6, d, e
Fraxinus, 125
fulliolus, Lycaena phlaeas americana
Alok, ZAt
fumosus, Danaus plexippus, 65

gemma, Euptychia, 8, 12 (in key), 36,
38; pl. 3, 9, h
genutia, Anthocharis, 18 (in key), 86;
pl. 9, i, J
genutia flavida, Anthocharis, 86
Glaucopsyche, 72
lygdamus, I, 5, 15 (in key), 72
lygdamus boydi, 72, 73
lygdamus lygdamus, pl. 15, b

lygdamus nittanyensis, 72, 73,
frontispiece, figs. 4, 5; pls. 14,
9, 15, €

glaucus, Papilio, 7, 20 (in key), 21
(ine key) -11G,. 022, 123, 224,
143; pls. 19, f, 20, 21
glaucus arcticus, Papilio,
130, 142
glaucus canadensis, Papilio, 133, 134,
136, 137, 139, 142
glaucus glaucus, Papilio, 133, 134, 136,
137, 139, 140
glaucus rutulus, Papilio, 124, 125, 126,
130, 134
Graphium, 145
marcellus, 20 (in key), 119, 143,
145 pl. 10, 9, h
marcellus floridensis, 146
marcellus telamonides, 146
gryneus, Mitoura, 16 (in key), 84;
Dieieg

134, 137,

halesus, Atlides, 8, 15 (in key), 773
frontispiece, fig. 10; pl. 12, c, d
hanno, Hemiargus, 187

INDEX

233

hayhurstii, Pholisora, 22 (in key),
153; pl. 15, 4, 7
hecla, Colias, 105
hegon, Amblyscirtes, 6, 27 (in key),
£7935 pis, 25, 4,:27, 1
Heliconiinae, 64
Hemiargus hanno, 187
henrici, Incisalia, 17 (in key), 84; pl.
16, d
Hesperia, 160, 162
attalus, 28 (in key), 1613 pl. 27, i
leonardus, 6, 28 (in key), 160;

pl. 26, a, b

metea, 6, 27 (in key), 160, 178;
pl. 28, a, b

sassacus, 5, 28 (in key), 161,

166; pl. 27, m, p
sassacus manitoboides, 161
Hesperiidae, 147
Hesperiinae, 11 (in key), 24 (key to
species), 159
hianna, Atrytonopsis, 6, 27 (in key),
177; pl. 26, g
hirsuta, Battus philenor, 119, 144
hobomok, Poanes, 1, 5, 28 (in key),
169; pl. 28, 7, k, |
horatius, Erynnis, 23 (in key), 158,
159; pl. 25, 7, k
horridus, Crotalus, 176
howardi, Poanes, 171
Poanes aaroni, 170, 171
hughi, Poanes massassoit, I, 172;
frontispiece, fig. 11; pl. 26, c, d
Speyeria aphrodite ab., 61
humuli, Strymon melinus, 81, 82, pl.
II, 0
Hylephila, 161
phyleus, 24 (in key), 161; pl. 28, c
Hystrix patula, 31

icelus, Erynnis, 22 (in key), 154; pl.
24,9,h
idalia, Speyeria, 3, 12 (in key), 54;
oe rN |
ilioneus, Papilio troilus, 122
Incisalia, 82
augustinus, 5, 17 (in key), 83, 84
augustinus croesioides, 83; pl. 16, e
henrici, 17 (in key), 84; pl. 16, d
irus, 5, 17 (in key), 84; pl. 16, c
niphon, 17 (in key), 82
niphon clarki, 83; pl. 30, A, ¢
niphon niphon, 82; pl. 13, 0D
polios, 5, 17 (in key), 83; pl. 13, a
interior, Colias, 4, 20 (in key), 1411,
TI23 Pl. 175 Cy Os 958

234

intermedia, Cyaniris argiolus pseudar-
giolus form, 73

interrogationis, Polygonia, 13 (in key),
40; pl. 16, f

interrogationis fabricii, Polygonia, 40

interrogationis umbrosa, Polygonia, 40

iole, Nathalis, 19 (in key), 1153 pl.

16, 9,h

irus, Incisalia, 5, 17 (in key), 84; pl.
16, ¢

ismeria, Melitaea, 48; frontispiece,
fig. 9

Ithomia, drymo, 187

joanna, Tisiphone abeona, 50
jucunda, Eurema, 19 (in key), 117,
118; pl. 16, 4, 7
Juniperus virginiensis, 85
Junonia, 45
evarete, 13 (in key), 45
evarete coenia, 45; pls. 4, h, 4, 7,
5, 9,4
juvenalis, Erynnis, 23 (in key), 157,
158, 159; pls. 24, 0, p, 25, f

Kalmia latifolia, 83
keewaydin, Colias eurytheme, 102

laeta, Erora, 5, 17 (in key), 85, 187;
frontispiece, fig. 8; pl. 12, a, b
leonardus, Hesperia, 6, 28 (in key),
160; pl. 26, a, b
Lerema, 178
accius, 26 (in key), 178; pl. 27,
0,7
Lerodea, 181
eufala, 8, 26 (in key), 1823 pl.
24, q
lherminieri, 25 (in key), 181;
DI Ure a
Lespedeza, 82
Lethe, 30
creola, 8, 11 (in key), 30; pl. 1,
GionG
eurydice, 11 (in key), 31
eurydice appalachia, 32; pl. 1, i, 7
portlandia, 11 (in key), 30
portlandia anthedon, 5, 3x3 pl. 1,

g,h
portlandia portlandia, 8, 30, 32;
pl. 1,'¢, f
Lethiinae, 10 (in key), 11 (key to
species), 30

Vherminieri, Lerodea, 25
ESQ splo Li, C0

(in key),

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 116

Libytheana, 68
bachmanii, 683 pl. 23, 7
Libytheidae, 10 (in key), 68
Limenitinae, 49
Limenitis, 49
archippus, 13 (in key), 52
archippus archippus, 52, 53, 54;
pl. 6, f
archippus eros, 54
archippus floridensis, 8, 52, 53,

54; pl. 6, d, e
arthemis, 14 (in key), 49, 50, 51,
143, 186

arthemis albofasciata, 51
arthemis arizonensis, 50
arthemis astyanax, 50, 51, 53;
pls) 2, ‘a, 6, cig ok
arthemis proserpina, 51
arthemis rubrofasciata, 50, 51
liparops, Strymon, 16 (in key), 813
pl. 13, e
Liriodendron, 125
lisa, Eurema, 20 (in key), 110, 116;
pl. 9, h
lisa form alba, Eurema, 116
lisa form clappi, Eurema, 116
logan, Atrytone, 24 (in key), 170,
173; pl. 27, c, d, e
longleyi, Ancyloxypha numitor, 159
lucia, Cyaniris argiolus pseudargiolus
form, 73
lucilius, Erynnis, 5, 23 (in key), 1553
pl. 24, k
Lycaena, 71
phlaeas, 2, 15 (in key), 7x
phlaeas americana, 713 pl. 15, J,
ml, 0
phlaeas americana ab. fulliolus, 71
thoé, 72, pl. 15, f, 9, h
Lycaenidae, 10 (in key), 15 (key to
species), 70
Lycaeninae, 11 (in key), 71
lyciades, Achalarus, 22 (in key), 148,
pl. 23, e
Lycorea cleobaea, 187
lygdamus, Glaucopsyche, 1, 5, 15 (in
key), 72
lygdamus boydi, Glaucopsyche, 72, 73
lygdamus lygdamus, Glaucopsyche, pl.
15, b
lygdamus nittanyensis, Glaucopsyche,
72, 73, frontispiece, figs. 4, 5;
pls. 14, g, 15, ¢

Magnolia, 125
m-album, Eupsyche, 16 (in key), 783
Pleo ea,

NO. 7

Malva rotundifolia, 152

manataaqua, Polites, 27 (in key), 164,
166; pls. 15, d, 28, e

manitoboides, Hesperia sassacus, 161

marcellus, Graphium, 20 (in key), 110,

- 143, 1453 pl. 10, g, h

marcellus floridensis, Graphium, 146

marcellus telamonides, Graphium, 146

marginata, Cyaniris argiolus pseudar-
giolus form, 73

marilandica, Boloria selene, I, 2, 633
frontispiece, figs. 1, 2; pl. 9, b, c

maritima, Minois pegala, 32, 33, 343

pl. 8, e
martialis, Erynnis, 23 (in key), 156,
187; pl. 24, 2

martialis ab. ausonius, Erynnis, 187
massassoit, Poanes, 25 (in key), 172
massassoit hughi, Poanes, 1, 1723
frontispiece, fig. 11; pl. 26, c, d
megalippe, Danaus plexippus, 2, 67,

68; pl. 10, b
melinus, Strymon, 16 (in key), 81
melinus humuli, Strymon, 81, 82;
Oe f0

melinus melinus, Strymon, 813 pl. 11, »
Melitaea, 47
ismeria, 48, frontispiece, fig. 9
nycteis, 15 (in key), 473 pl. 9, f, 9
Melitaeinae, 47
metacomet, Atrytone ruricola, 177;
pls. 11, e, 26, e
metea, Hesperia, 6, 27 (in key), 160,
178; pl. 28, a, b
mingo, Potanthus, 187
Minois, 32
pegala, 11 (in key), 32
pegala alope, 32, 33, 34, 35, 36,
pl. 2, ifs g
pegala carolina, 35
pegala maritima,
pl. 8, e
pegala nephele, 33
pegala pegala, 8, 34, 353 pl. 2, 4,1
pegala texana, 36
Mitoura, 84
gryneus, 16 (in key), 84; pl. 12, g
monuste, Ascia, 93
mopsus, Strymon titus, 79; pl. 13, h
morrisi, Tisiphone abeona, 50, 51
myrina, Boloria selene, 5, 623 pl. 9, a
mystic, Polites, 5, 20 (in key), 166;
pl. 26, 7, k

32, 33, 34;

nastes werdandi, Colias, 106
Nathalis, 115
iole, 19 (in key), 1153 pl. 16, 9g, ht

INDEX 235

neglecta, Cyaniris argiolus pseudar-
giolus form, 73, 74

neglecta-major, | Cyaniris
pseudargiolus form, 74

nephele, Minois pegala, 33

nicippe, Eurema, 20 (in key), 1153
pl. 2, e

nigra, Cyaniris argiolus pseudargiolus
form, 73

nigrior, Agraulis vanillae, 65; pl. 7, e, f

niphon, Incisalia, 17 (in key), 82

niphon clarki, Incisalia, 83; pl. 30, A, i

niphon niphon, Incisalia, 82; pl. 13, b

nittanyensis, Glaucopsyche lygdamus,
72, 73; frontispiece, figs. 4, 5;
PISH14) GA1s..c

numitor, Ancyloxypha, 24 (in key),
159; pl. 3, k

numitor longleyi, Ancyloxypha, 159

nycteis, Melitaea, 15 (in key), 473
plo fg

Nymphalidae, 10 (in key), 12 (key to
species), 40

Nymphalinae, 40

Nymphalis, 42

antiopa, 2, 12 (in

pl. 2, b

Nyssa aquatica, 8, 175

argiolus

key), 42's

occidentalis, Pieris protodice, 92

Ochlodes sylvanus, 187

ocola, Panoquina, 25 (in key), 186;
pl. 26, p

Oeneis, 124

olympia, Euchloé, 18 (in key), 87, 88;
pl. 16, a, b

olympia rosa, Euchloé 87, 88

ontario, Strymon, 1, 16 (in key), 79

ontario ontario, Strymon, 80; pl. f

otho, Wallengrenia, 26 (in key), 167

otho egeremet, Wallengrenia, 167;
pl. 26, m, n

otho otho, Wallengrenia, 167; pl. 26, 0

palamedes, Papilio, 7, 8, 21 (in key),
121, 122; pl. 22, c, d
palatka, Atrytone, 8, 24 (in key), 173;
pl. 20, g, h, 4
panoquin, Panoquina, 8, 26 (in key),
185; pl. 26, /
Panoquina, 185
ocola, 25 (in key), 186; pl. 26, p
panoquin, 8, 26 (in key), 185;
pl. 26;.\
Papilio, 120
cresphontes, 25 (in key), 120;
pl. 10, e

236

Papilio glaucus, 7, 20 (in key), 21 (in
key), Wor 122) “T2g4en24. 143):
pls. 19, f, 20, 21
glaucus arcticus, 134, 137, 139, 142
glaucus canadensis, 133, 134, 136,
137, 139, 142
glaucus glaucus, 133, 134, 136, 137,
139, 140
glaucus rutulus, 124, 125, 126, 130,
134
palamedes, 7, 8, 21 (in key), 121,
122) plei2o.c. a
polyxenes, 21 (in key), 144
polyxenes ampliata, 144
polyxenes asterius, 1443 pl. 22,
e,f, 9,h
polyxenes curvifascia, 144
rutulus arcticus, 134
troilus, 21 (in key), 122, 123;
Dl 22 a0
troilus ilioneus, 122
Papilionidae, 11 (in key), 20 (key to
species), 118
Papilioninae, 118
Passiflora incarnata, 2, 64, 65
peckius, Polites, 29 (in key), 1653
pl. 28, g, h
pegala, Minois, 11 (in key), 32
pegala alope, Minois, 32, 33, 34, 35,
36; pl. 2, f, g
pegala carolina, Minois, 35
pegala maritima, Minois, 32, 33, 34;
pl. 8, e
pegala nephele, Minois, 33
pegala pegala, Minois, 8, 24, 35; pl. 2,
ht
pegala texana, Minois, 36
Persea borbonia, 3
persius, Erynnis, 5, 23 (in key), 1553
pl. 24, 7,7
phaéton, Euphydryas, 5, 14 (in key),
47; pis. 3, J, 30, e
philea, Phoebis, I, 19 (in key), 1423
pl. 10, d
philenor, Battus, 21 (in key), 118,
140, 144; pl. 10, a, b, c, d
philenor acauda, Battus, 119, 144
philenor hirsuta, Battus, 119, 144
phileta, Ascia, 1, 18 (in key), 19 (in
key), 93
phileta phileta, Ascia, 93; pl. 8, g, h,

1,

sdiladice: Colias, 5, 6, 18 (in key), 20
(in key), 93, 97, IOI, 102, 103,
106, 107, 108, 100, II10, III, 112;
pl. 17, a, b, e

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I16

phlaeas, Lycaena, 2, 15 (in key), 7x
phlaeas americana, Lycaena, 713 pl.
15, l,m, n
phlaeas americana ab. fulliolus, Ly-
caena, 71
Phoebis, 112
philea, 1, 19 (in key), ux23 pl.
10, d
sennae, I, 19 (in key), 1123 pl.
10, ¢
sennae drya, 113
Pholisora, 153
catullus, 22 (in key), 1533 pl.
20
hayhurstii, 22 (in key), 1533 pl.
15, 4 J
Phoradendron flavescens, 3, 77
Phyciodes, 48
batesii, 5, 15 (in key), 493 pl. 11,
f 9; h, t
tharos, 15 (in key), 48, 49; pl.
11, j,k, l,m
phyleus, Hylephila, 24 (in key), 161;
pl. 28, c
Pieridae, 11 (in key), 18 (key to
species), 86
Pierinae, 86
Pieris, 88, 107
protodice, 18 (in key), 92; pl. 18,
c,d,e,f,g9,h
protodice occidentalis, 92
rapae, I, 2, 18 (in key), 19 (in
key), 88, 91; pl. 18, a
virginiensis, I, 5, 19 (in key),
90, oI, 187; pl. 18, b
piscivorus, Agkistrodon, 176
plexippus, Danaus, 2, 13 (in key), 65,
67
plexippus ab. fumosus, Danaus, 65
plexippus megalippe, Danaus, 2, 67,
68; pl. 10, b
plexippus plexippus, Danaus, 65, 67,
68; pl. 10, a
Poanes, 168
aaroni, 8, 25 (in key), 470; pl.
28, m, n, 0, r
aaroni aaroni, 170, 171
aaroni howardi, 170, I71
hobomok, 1, 5, 28 (in key), 169;
pl. 28, 7, k, 1
howardi, 171
massassoit, 25 (in key), 172
massassoit hughi, 1, 1723 frontis-
piece, fig. 11; pl. 26, c, d
viator, 8, 28 (in key), 168; pl.
20, J

NO. 7

Poanes yehl, 8, 28 (in key), 7713 pls.
28, q, 30, 4, f, g
zabulon, 24 (in key), 1693 pl. 27,
a,
polios, Incisalia, 5, 17 (in key), 833
Beh
Polites, 164
manataaqua, 27 (in key), 164,
166; pls. 15, d, 28, e
mystic, 5, 29 (in key), 1663 pl.
26, j, Rk
peckius, 29 (in key), 1653 pl.
28, g,h
themistocles, 27
166; pl. 28, f
verna, 27 (in key), 164, 166; pl.
28, d
vibex, 24 (in key), 1673 pls. 27,
hy ash oe
Polygonia, 40
comma, 13 (in key), 403 pl. 5,
End
faunus, 13 (in key), 41
faunus smythi, 4, 423; frontispiece,
figs. 3, 6; pl. 5, a, b
interrogationis, 13 (in key), 40;
mbe We i
interrogationis fabricii, 40
interrogationis umbrosa, 40
progne, 5, 13 (in key), 41; pl. 5,
Gra
polyxenes, Papilio, 21 (in key), 144
polyxenes ampliata, Papilio, 144
polyxenes asterius, Papilio, 144; pl.
22, e, f; g; h
polyxenes curvifascia, Papilio, 144
portlandia, Lethe, 11 (in key), 30
portlandia anthedon, Enodia, 5
portlandia anthedon, Lethe, 5, 31; pl.
TQ.
portlandia portlandia, Lethe, 8, 30, 32;
inh CA
Potanthus mingo, 187
Potentilla canadensis, I51
progne, Polygonia, 5, 13 (in key), 413
pl. 5, ¢,d
proserpina, Limenitis astyanax, 51
Proteides, 147
clarus, 22 (in key), 1473; pl. 23, a
proteus, Urbanus, 1, 21 (in key), 148;
pl. 23, d
protodice, Pieris, 18 (in key), 923
Diets GG, Cy 15g), 2
protodice occidentalis, Pieris, 92
Prunella, 180

(in key), 165,

INDEX 237

pseudargiolus, Cyaniris argiolus, 73,
75; pl. 14, a, b,c, d,e, f,h
pseudargiolus form intermedia, Cyani-

ris argiolus, 73
pseudargiolus form lucia, Cyaniris ar-
giolus, 73
pseudargiolus form marginata, Cyaniris
argiolus, 73
pseudargiolus form neglecta, Cyaniris
argiolus, 73, 74
pseudargiolus form neglecta-major, Cy-
aniris argiolus, 74
pseudargiolus form nigra, Cyaniris ar-
giolus, 73
pseudargiolus form pseudargiolus, Cy-
aniris argiolus, 73, 74
pseudargiolus form violacea, Cyaniris
argiolus, 73
pylades, Thorybes, 22 (in key), 149,
150, 151; pl. 25, a, b
Pyrginae, Ir (in key), 21
species), 147
Pyrgus, 151
centaureae, 5, 23 (in key), 151
centaureae wyandot, 151; pl. 24,
a,d
communis, 23 (in key), 152; pl.
DAD Ge ,aT
communis albescens, 152

(key to

rapae, Pieris, 1, 2, 18 (in key), 19 (in
key), 88, 91; pl. 18, a

rawnsleyi, Tisiphone abeona, 51
reversa, Amblyscirtes carolina, 180
Robinia pseudacacia, 147
rosa, Euchloé olympia, 87, 88
Rhabdoides, 449

cellus, 22 (in key), 149; pl. 23,

Bic

Riodinidae, 10 (in key), 15 (key to
species), 69

rubrofasciata, Limenitis arthemis, 50,
51

Rumex acetosella, 71

ruricola, Atrytone, 25 (in key), 164,
177

ruricola metacomet,
pls. 11, e, 26, e

rutulus, Papilio glaucus, 124, 125, 126,
130, 134

rutulus arcticus, Papilio, 134

Atrytone, 1773

sassacus, Hesperia, 5, 28 (in key),
161, 166; pl. 27, m, p
sassacus manitoboides, Hesperia, 161

238

Satyridae, to (in key), 30
Satyrinae, 10 (in key), 11 (key to
species), 32
Scirpus, 3, 186
selene, Boloria, 2, 3, 12 (in key), 62
selene marilandica, Boloria, 1, 2, 633
frontispiece, figs. 1, 2; pl. 9, b, c
selene myrina, Boloria, 5, 62; pl. 9, a
sennae, Phoebis, 1, 19 (in key), 1123
pl. 10, c
sennae drya, Phoebis, 113
septentrionalis, Euptychia areolatus, 8,
373 pl. 3, a, ¢
smythi, Polygonia faunus, 4, 423 fron-
tispiece, figs. 3, 6; pl. 5, a, b
sosybius, Euptychia, 8, 12 (in key),
38; pl. 3,7
Spalginae, 10 (in key), 70
Spartina alterniflora var. glabra, 3, 171
Speyeria, 3, 54
aphrodite, 3, 5, 15 (in key), 59;
pl. 8, a, b
aphrodite alcestis, 59
aphrodite ab. bakeri, 60
aphrodite ab, hughi, 61
atlantis, 4, 15 (in key), 61; pl. 8,
(By a5
cybele, 3, 12 (in key), 56, 58, 60;
pls. 2, ¢, 7, 9, h
diana, 1, 3, 14 (in key), 55, 91;
pl. 7, a, b, c,d
ialia, 3) 12 "(Gan key) ,. S4s.pl. 7,
t, J
Strymon, 78
cecrops, 16 (in key), 783 frontis-
piece, fig. 7; pl. 12,h
cecrops ab. gottschalki, 78; frontis-
piece, fig. 7
edwardsii, 6, 17 (in key), 80; pl.
13, d
falacer, 17 (in key), 80; pl. 13, c
liparops, 16 (in key), 81; pl. 13, e
melinus, 16 (in key), 84
melinus humuli, 81, 82; pl. 11, 0
melinus melinus, 8x; pl. 11,
ontario, I, 16 (in key), 79
ontario ontario, 80; pl. 13, f
titus, 16 (in key), 79
titus mopsus, 79; pl. 13, h
titus titus, 79; pl. 13, g
sylvanus, Augiades, 1, 187
Ochlodes, 187

tarquinius, Feniseca, 17 (in key), 703
pl. 15, e, k

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I16

telamonides, Graphium marcellus, 146
texana, Minois pegala, 36
textor, Amblyscirtes, 8, 26 (in key),
180; pl. 3, j
tharos, Phyciodes, 15 (in key), 48,
49; pl. 11, 7, k, l,m
Theclinae, 11 (in key), 77
themistocles, Polites, 27 (in key), 165,
166; pl. 28, f
thoé, Lycaena, 72; pl. 15, f, g, h
Thorybes, 149
bathyllus, 22 (in key), 1493 pls.
23, 9, h, 25, e
confusis, 22 (in key), 1503 pl. 25,
Bas
pylades, 22 (in key), 140, 150,
T5is pl. 25) /a,.0
Tisiphone abeona, 50
abeona aurelia, 50, 51
abeona joanna, 50
abeona morrisi, 50, 51
abeona rawnsleyi, 51
titus, Strymon, 16 (in key), 79
titus mopsus, Strymon, 79; pl. 13, h
titus titus, Strymon, 79; pl. 13, g
toddi, Boloria, 3, 5, 14 (in key), 61
toddi ammiralis, Boloria, 61, 62, 63;
pl. 0, d, e
troilus, Papilio, 21 (in key), 122, 123;
pl22aee
troilus ilioneus, Papilio, 122

umbrosa, Polygonia interrogationis, 40
Urbanus, 148
proteus, I, 21 (in key), 148; pl.
oad

Vanessa, 43
atalanta, 2, 13 (in key), 43, 44,
45; pl. 8, f
cardui, I, 2, 14 (in key), 44, 45;
pl. 4, d, e
virginiensis, 14 (in key), 443 pls.
4, f, 9, 30, 7, F
vanillae Agraulis, 1, 8, 12 (in key), 64
vanillae nigrior, Agraulis, 65; pl. 7,
ae)
verna, Polites, 27 (in key), 164, 166;
pl. 28, d
vialis, Amblyscirtes, 27 (in key), 1793
pl. 27, 7, q
viator, Poanes, 8, 18 (in key), 168;
pl. 20, 7
vibex, Polites, 24 (in key), 1673 pls.
27), ‘iB 28, t

NO. 7

Vicia caroliniana, 3
violacea, Cyaniris argiolus pseudargio-
lus form, 73
virginiensis, Calephelis, 8, 15 (in key),
69; pl. 12, 7
Pieris, 1, 5, 19 (in key), go, 91,
187; pl. 18, b
Vanessa, 14 (in key), 443 pls. 4,
FyGs 30; 9,

Wallengrenia, 167
otho, 26 (in key), 167
otho egeremet, 1673 pl. 26, m, 1
otho otho, 167; pl. 26, o
werdandi, Colias nastes, 106

INDEX

239

wyandot, Pyrgus centaureae, 151; pl.

24, a, d

yehl, Poanes, 8, 28 (in key), 1713
pls. 28, q, 30, d, f, g

zabulon, Poanes, 24 (in key), 169;
Dl 27a a0.

Zanthoxylum clavaherculis, 120

zarucco, Erynnis, 23 (in key), 1583

pl. 25, g, h
Zerene, 112
caesonia, 19 (in key), 1123 pl.
26
Zizania palustris, 3, 168

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