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Science Bulletin

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THE MYOLOGY OF SCELOPORUS C. CLARKI
BAIRD AND GIRARD (REPTILIA: IGUANIDAE)

by

D. M. Secoy

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 1

JUNE 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

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Brigham Young University
Science Bulletin

THE MYOLOGY OF SCELOPORUS C. CLARKI
BAIRD AND GIRARD (REPTILIA: IGUANIDAE)

by
D. M. Secoy

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 1

JUNE 1971

TABLE OF CONTENTS

MATERIALS AND METHODS 1

INTRODUCTION 2

MYOLOGY 3

DISCUSSION 10

CONCLUSIONS 21

LITERATURE CITED 21

THE MYOLOGY OF

SCELOPORUS C. CLARKI BAIRD AND GIRARD

(REPTILIA: IGUANIDAE)

by

D. M. Secoy'

ABSTRACT

Examination of the complete musculatm-e of
Sceloporus clarki clarki Baird and Girard and
seven other species of Scelojwrns revealed varia-
ation among species of Sceloporus and between
Sceloporus and other iguanid lizards. The
muscles in which the greatest variation was
found were the intermandibularis group, the con-
strictor colli, the epistemo-cleido-mastoideus, the

epistemohyoideus, the coracoid head of the tri-
ceps, the costocoracoid and the flexor tibialis
extemus. This study indicates: (1) Sceloporus
is more closely allied to Crotaphyius than to the
ground-dwelling iguanines;(2) the possibility of
the basal stock of Sceloporus being arboreal,
and, (3) Sceloporus is a genus in the process of
rapid differentiation.

INTRODUCTION

Even a cursory glance at the literature deal-
ing with the morphology of the suborder Lacer-
tiJia reveals the scarcity and scattered nature of
studies concerned with the soft anatomy.

The myology of the entire body has been
described in varjdng detail in eight genera of
the suborder Lacertilia: Iguana ( Mivart, 1867),
Chaniaeleon (Mivart, 1870) ,Platydactt/lus (San-
ders, 1870), Liolepis (Sanders, 1872), Plmjno-
soma (Sanders, 1874), Pseudopus (Humphry,
1872), Chlamtjdosaurm (de Vis, 1883), and Vro-
mastix (George, 1948) have been examined and
described. Laboratory dissection guides showing
the antomy of Crotaphijtus (Davis, 1934) and
Aganm (Harris, 1963) are brief.

A search of the literature for descriptions of
portions of the anatomy of a genus, or compara-
tive works examining representatives of different
genera or families is much more rewarding. The
former includes the recent work by Oelrich
(1956) on the head of Ctenosaura pectinata,
Avery and Tanner's paper ( 1964 ) on the head
and thorax of Sauromalus, Robison and Tanner's
paper (1962) on the anterior region of Crota-
plnjtus and Gambelui. and Jenkins and Tanner's
paper (1968) on Phnjnosoma.

Comparative studies have been divided into
three main groups: those concerned with the
muscles of the jaw articulation; those which
have examined the variation in the muscles con-

nected with the hyoid apparatus; and those
head of reptiles, as is the case with other verte-
brate groups, has been studied most extensively.
The works on the jaw muscles include those of
Versluys (1904), Bradley (1903), Adams (1919),
Lakjer (1926), and Haas (1960). Some of the
more important works on the hyoid apparatus
are those by Zavattari (1908), Gandolfi (1908),
and Gnanamuthu ( 1936 ) . Edgeworth's treatise
(1935) on cranial muscles includes tiU head
musculature and discusses the homologies of
structures extensively, although Brock (1938)
disagrees with his ideas of muscle origins. Camp
( 1923 ) has used the numerous generic descrip-
tions of the hyoid and its associated musculatiu-e
in the construction of his lizard classification.

The other area of intensive interest and work
hiis been in limb musculature. Romer (1922,
1923, 1942) has estabUshed homologies in the
pelvic limb muscles among the several reptilian
groups based on his research and that of Gadow
(1882). Haines (1934, 1935) and Appleton
( 1928 ) have also discussed the homologies of
the pelvic and thigh muscles. The anterior limb
and pectoral girdle have been studied by Fiir-
bringer (1876), Romer (1924, 1944), McMurrich
(1903, 1903a), and Howell (1938).

Since its description by Wiegmann in 1828
Sceloporus has been one of the most intensively
studied of the New World iguanid lizards. The

'Department of Biology, University of Saskatchewan, Regina, Saskatchewan. Canada.

Brigham Young University Science Bulletin

large size of the genus and its broad distribution
concerned with the muscles of tlie lijnbs. The
have made it ideal for studies on taxonomy and
distribution. The same two factors have made it
unwieldy and a deterrent to those who wish to
study the entire genus.

Early studies of the genus include those of
Dumeril and Bibron (1837), Boulenger (1897),
and Cope (1900). The definitive work on the
genus is that of Hobart Smith ( 1936, 1937, 1937a,
1938, 1939). These studies have been based on
external characteristics and the only mention of
skeletal or soft anatomy is at the generic level.

The osteology of the genus has been de-
scribed piecemeal, usually in conjunction with
that of other reptiles. Cope ( 1892 ) gives a com-
plete description of the skull and skeleton but
gives no figures. Camp (1923), in his work on
lizard classification, refers to Sceloponis in the
discussion of many of the skeletal characters.
Stokely (1950) mentions the possibility of the
occurrence of an intermedium in SceJoporus.
Hotton (1955) discusses the dentition of Sce-
lopoTus graciosus, S. magister and S. undulatus in
relation to their prey choice. Romer (1956)
illustrates various skeletal elements in his treatise
on reptilian osteology. Lundelius ( 1957 ) made
a statistical analysis of the skeletal adaptations
of Sceloporus oUvacetis and S. undulatus hi/a-
cinthinus in relation to their environments. Jollie
(1960) mcluded Sceloporus undulatus in his
analysis of the lizard skull. Avery and Tanner
( 1964 ) illustrated and discussed the variation in
the wrist bones of several genera of iguanids,
including Sceloporus. Etheridge ( 1964, 1965,
1967) discussed and illustrated several skeletal
elements of Sceloporus in his studies of skeletal

variation within the Iguanidae. Miller (1966)
discussed Sceloporus in his work on the cochlear
duct in the Lacertilia.

The myology has been less well studied.
Camp ( 1923 ) mentioas the genus in his discus-
sion of the rectus abdominis and the mandibulo-
hyoideus (his genio-hyoideus ) . Avery and Tan-
ner ( 1964 ) use a specimen of Sceloporus magis-
ter for comparison with Sauroinalus. Snyder
( 1962 ) illustrated the hind leg of Sceloporus in
his discussion of muscle grouping and develop-
ment in relation to lizard bipedahsm. In his
paper on the anatomical ratios of lizard limbs,
Snyder ( 1954 ) used Sceloporus as an example
of persistent quadrupedality.

The paucity of information on the muscular
anatomy of one of the largest genera of New
World lizards was the impetus for this study.
The primary concern of the study was the de-
scription of the myology of the entire body of
Sceloporus clarki. Comparative dissections of
other species of Sceloporus were made. Com-
parisons were also made with descriptions of the
myology of other iguanids.

I wish to thank Dr. Hugo Rodeck, director,
and Dr. T. Paul Maslin, curator of zoology, of
the Museum of the University of Colorado for
permission to dissect specimens from the herpe-
tological collections. Funds for the collection of
specimens were provided by the Kathv Lichty
Fund and Grant #GB 2362, National' Science
Foundation made to Dr. T. Paul Maslin. The
figures were drawn by Miss Susan George.

This manuscript has been reviewed by Drs.
W. W. Tanner, H. M. Smith, and D. F. Avery.
Their comments and suggestions are greatly
appreciated.

MATERIALS AND METHODS

The species used for the basic dissections
was Sceloporus c. clarki Baird and Girard. Suf-
ficient specimens were dissected to note varia-
tion found within a species. Sceloporus magister,
a closely related species (Lowe, Cole and Patton,
1967), was selected for comparative purposes.
The phylogenetic tree postulated by Smith
( 1939 ) was used to select several other species
of Sceloporus for comparative examination.
These species were used to detennine at what
level of consanguinity, anatomical differences,
if any, occurred.

Speciment of Sceloporus c. clarki examined
were UCM 13289, 134095, .34099, .34183, .34184,
34185, 34202, 34215, 34216, ^4217, 34218, .34157.

341.58, 34159, .35160, 34179, 34180. Five unregis-
tered specimens were also dissected.

Specimens of other species of Sceloporus
used for comparative purposes were Sceloporus
magister Hallowell UCM ;3017S; Sceloporus poin-
setti Baird and Girard UCM 34132; Sceloporus
V. variabilis Wiegmann UCM 28778, 28782; Sce-
loporus nuilachiticus smargadinus Boucourt
UCM 24523, 24525; Sceloporus undulatus ery-
throcheilus M;islin UCM 17414, 17442; Scelo-
porus grammicus dlspariUs Stejneger UCM
20034; Sceloporus chrtfsostictus Cope UCM
16511, 16548.

All specimens had been fixed in 10% formalin
and stored in 70% ethanol.

BiOLOOicAL Series, Vol. 14, No. 1 Myology of Sceloporu.s

Myology

The basic description following the muscle
name is that of the condition found in Scelopo-
rus clarki. Any deviation from this condition
which was found in other species of Sceloporus
dissected is then noted. Comparisons were then
made with the published descriptions and illus-
trations of other iguanid lizards. The genera
used for comparison were Iguana ( Mivart, 1867;
Romer, 1922; Haines, 1934, 1936; Howell, 1936,
1938; and Evans, 1939), Phnjnosonui (Sanders,
1874; Jenkins and Tanner, 1968), Ctenosaura
(Oelrich, 19.56), Sauromahts ( Avery and Tanner,
1964) and Crotaphytus (Robison and Tanner,
1962). Only those muscles which varied within
the genus Sceloporus or between Sceloporus and
other iguanids are included in the present
account.

Muscle nomenclature used here is based pri-
marily on that of Romer for the body muscula-
ture and Edgeworth for the musculatm-e of the
head.

Throat and Hyok Musculature

Depressor palpebrae inferioris

Origin: from the floor of the orbit.
Insertion: into the whole of the lower eyelid.
This is an extremely thin and diffuse set of
fibers found in the lower eyelid. No levator
palpebrae superioris was found. Of the other
studies, this muscle was noted only in Cteno-
saura.

Intermandibularis

This is a thin sheet of muscle which lies on
the ventral aspect of the lower jaw over the
hyoid musculature. A varying number of fibers
insert into the skin of the gular region. The
muscle is divided into an anterior and posterior
portion. The ;mterior portion is designated the
intermandibularis anterior profundus, since it
appears to be homologous to the deep portion
of the intermandibularis anterior in those genera
in which the intermandibularis has three slips.
Tlie fibers of this muscle He perpendicular to
the long axis of the body.

Intermandibularis anterior profundus

Origin: from the medial side of each dentary.
Insertion: into the midventral fascia.
In this genus the only differentiation be-
tween the anterior and posterior slips of the
intermandibularis is the unmasked origin of the
anterior profundus from the ramus. Its origin
lies between that of the mandibulohyoideus and
the genioglossus.

The intermandibularis anterior superficiaUs
is not present in any species of Sceloporus ex-
amined. A narrow band of this muscle is present
in Crotaphijtus, Sauromahts and Ctenosaura. In
Phrijnosorjui, the muscle appears to be fairly
broadly developed.

Intermandibularis posterior

Origin: from the la tero- ventral to lateral sur-
face of the posterior half of the mandible.

Insertion: into the midventral fascia.

The anterior portion of the muscle comes to
the surface of the throat by a varying number
(3-7, usually 5-6) of slips which interdigitate
with the mandibidohyoideus I. The number of
interdigitations is usually asymmetrical for an
individual. The posterior portion comes from
the lateral surface of the jaw, from the fascia
covering the pterygomandibularis and also di-
rectly by a tendon from the lateral surface of the
ramus between the pterygomandibularis and the
insertion of the depressor mandibularis. In many
specimens the muscle is composed only of fascia
in the center so that the body of the hyoid and
the base of the hngual process are visible
through the fascia. This window of fascia occurs
at the posterior boundary of the intermandibu-
laris posterior. The fibers of the constrictor colli
go to the midline just posterior to the window.
In cases where this window was not present, it
was very difficult to differentiate this muscle
from the constrictor colli.

In the specimen of magister examined in the
course of the dissection the intermandibularis
posterior and constrictor colli appeared as one
sheet of muscle. There also was no great divi-
sion between the two portions of the intermandi-
bularis. Avery and Tanner ( 1964, p. 15 ) state
that the specimen of magister which they ex-
amined had a broad separation between the two
portions of the intermandibularis. In variabilis
the intennandibularis posterior was very thin
and composed of fascia over the entire ventral
surface of the jaw while the anterior portion was
very well developed and several fibers thick.
In gramniicus there was a window of fascia over
the hyoid body. In poinsetti the posterior was
thinlv developed while the anterior was heavily
developed. In chnj.sostictus there was no dif-
ferentiation between the posterior and the con-
strictor colli. This is also the case in undulatus.

In Crotaphtjtu,y it was stated (Robison and
Tanner, 1962) that the anterior profundis and
posterior portions were not easily separated ex-
cept by origin as in Sceloporus. Rut, the con-
strictor colli was separated from the posterior
margin of the intermandibularis posterior by a

Bhicham Young University Science Bulletin

band of fascia. There are three or four interdigi-
tations of the posterior section with the mandi-
bulohyoideus I. In Sauwnialns the posterior is
a narrow band of muscle which is separated
from both the anterior section of the muscle and
the constrictor colli by bands of fascia. In
Ctenosoiira the posterior section is broad and in
contact with the constrictor colli. In Iguana the
intermandibularis is heavily developed in con-
junction with the dewlap. In Phnjnosonm the
intermandibularis group is well developed, but
Sanders mentions no interdigitations of the pos-
terior with the mandibulohyoideus I.

Constrictor colli

Origin: from the cervical portion of the
dorsal aponeurosis.

Insertion: into the midventral fascia.

This is the thin sheet of muscle which covers
the side of the neck and the posterior portion
of the throat and hyoid. It is superficial to the
depressor mandibulae group, the omohyoid,
epistemo-cleido-mastoid, epistemohyoid and the
deep neck muscles. The muscle is usually so
broadly developed as to cover part of the tem-
poral fossa. Some clarki specimens had fibers
inserting into the surface fascia of the epistemo-
hyoid superficialis.

In poirusetti the constrictor colli was broadly
developed but liecame tendinous before reach-
ing the midline. In undulatus the constrictor
colli was connected to the epistemohyoideus but
was not as broad as in clarki. In variabilis, nuila-
chiticus, and graminicus, it was present only as
a narrow band across the throat, widely separat-
ed from the intennandibularis posterior, and
covering only a portion of the omohyoid and
epistemohyoideus superficialis.

The constrictor colli is present as a narrow
band also in Phnjnosoma, Crotaphtjttis, Satiro-
nuiJtis, and Ctenosaura. In Iguaiui it is more
heavily developed in conjunction with the throat
fan and Ls continuous with the intermandibularis
posterior.

Mandibulohyoideus I

Origin: on the medial surface of the posterior
half of the ramus.

Insertion: to almost the entire length of the
first ceratobranchial.

Tliis muscle lies immediately deep to the
intermandibularis posterior, lateral to the hyo-
glossus, medial to the pterygomandibularis and
superficial to the branchiohyoideus. The inter-
mandibularis rises to the surface through inter-
digitations with this muscle.

Mandibulohyoideus II

Origin: by a thin, broad band of fascia from
the dentary symphysis and the adjacent bone.

Insertion: on the imterior portion of the first
ceratobranchial near the body of the hyoid.

The insertion may extend down the anterior
half of the first ceratobranchial on the ventral
and ventrolateral surfaces.

H yogi OSS us

Origin: from ceratobranchial I.

Insertion: to the lingual process of the hyoid
and the fleshy base of the tongue.

This muscle is deep to the posterior portion
of the genioglossus and superficial to the brachi-
ohyoideus. It occupies the space between the
ceratohyal and ceratobranchial I.

Genioglossus

Origin: from the anterior third of the mandi-
ble anterior to the origin of mandibulohyoideus
I.

Insertion: on the first ceratobranchial near
the body of the hyoid and also by fascia into the
covering fascia of the dorsal surface of mandi-
bulohyoideus I.

The origin is deep to that of mandibulohyo-
ideus II and the insertion is superficial to the
hvoglossus.

Branchiohyoideus

Origin: from the center of the ceratohyal.

Insertion: to the posterior portion of cerato-
branchial I.

This small muscle fills almost the entire
space between the hyoid bones except for the
most medial portion.

In CrotapJu/tits and Sauromalus this is a
much smaller muscle which is confined to the
lateral portion of the space between the hyoid
bones. It is fairly broad in Ctenosaura but does
not fill the entire space.

Constrictor laryngi

Origin: from the midventral line of the crico-
thyroid cartilage.

Insertion: to the middorsal line of the epi-
glotic cartilage.

Dilator laryngi

Origin: from the base of the cricothyroid
cartilage.

Insertion: to the dorso-lateral surface of the
posterior process of the cricothyroid cartilage.

Epistcmo-cleido-mastoideus

Origin: from the ventral siu'face of the clavi-
cle, from the ventral surface of the stemum by

Biological Series, Vol. 14, No. I Myology of Sceloporus

a heavy, broad band of fascia which does not
develop muscle fibers until it passes over the
clavicle and, usually, a slip of varying size which
rises from the fascia of the epistemohyoideus
superficialis.

Insertion: to back of quadrate and lateral
surface of exoccipitals.

The origin is superficial to that of the omo-
hyoid. The muscle lies deep to the constrictor
colli and the depressor mandibularis group. In
the other species of Sceloporus, except in the
specimen of grammicus which was examined,
there was also a slip from the surface of the
epistemohyoideus superficialis. However, the
variability of the development of this muscle in
clarki would suggest that the condition in the
single specimen of grammicus might not always
be the case.

Crotaphtjtus may have a sHp from the stemo-
hyoideus superficialis, depending on the species.
Ctenosaura has a doubleheaded epistemo-cleido-
mastoideus. Sauwmalus has no carry-over of
muscle fibers from the stemohyoideus super-
ficialis. Avery and Tanner reported a single
head in the specimen of Sceloporus magister
they dissected.

Omohyoideus

Origin: from the scapula and the lateral part
of the clavicle.

Insertion: into the body of the hyoid and
the whole of ceratobranchial I.

This is a straplike muscle, triangular in cross-
section, which is deep to the intemiandibularis
posterior and constrictor colli, medial to mandi-
bulohyoideus I and superficial to the insertion
of the epistemohyoideus superficialis. The ori-
gin is covered by that of the epistemo-cleido-
mastoideus. In one case the omohyoid took part
of its origin from the midhne fascia superficial
and lateral to the epistemohyoideus superficialis.
In another case this muscle was markedly asym-
metrical in its development. In some cases a
raphe was present in the center of the muscle
at the level of the raphe marking the insertion of
the epistemo-hyoideus profundus into the fibers
of the epistemohyoideus superficialis, which lies
medial of the omohyoid.

This muscle was well developed in most of
the other species of Sceloporus. In undulatus
and chrysostictus it was thin, straplike and rec-
tangular in cross-section. In chrysostictus the
muscle was elongated, as were the epistemo-
hyoideus and epistemo-cleido-mastoideus, due to
the lengthening of the neck and hyoid apparatus
in this species.

The investigators of Crotaphytus and Sauro-
nuilus did not find the omohyoid to be distin-
guishable from the epistemohyoideus. Sanders
illustrates a well-developed omohyoid in Phryno-
sonui. In Ctenosaura the muscle was present and
rectangular in cross-section.

Epistemohyoideus superficialis

Origin: by a thin, broad fascia from the
interclavicle and the fascia covering the pec-
toralis.

Insertion: onto almost the entire length of
ceratobranchial I.

The origin of tliis muscle is superficial to the
pectoralis and deep only to the skin. The inser-
tion is immediately deep to the omohyoid. The
body of the muscle nms parallel to the cerato-
branchial II and is attached to it by a fascia but
no muscle fibers. A varying number of lateral
fibers are incorporated into the epistemo-cleido-
mastoideus. In most specimens of clarki only
one slip of this muscle is present.

Epistemohyoideus profundus

Origin: from the midline fascia and inter-
clavicle.

Insertion: into the dorsal surface of the epi-
stemohyoideus superficialis by a discernible
raphe.

This muscle is either only occasionally pres-
ent or very difficult to divide from the super-
ficialis.

In magister, undulatus, and malachiticus,
only the superficialis was present. In one speci-
men of variabilis the muscle was single. In the
other, the superficialis inserted onto the proxi-
mal portion of ceratobranchial I and completely
enveloped ceratobranchial II. In grammicus the
muscle had a single origin but a double inser-
tion. One insertion was to the body and proxi-
mal portion of ceratobranchial I, while the other
slip was to the most distal portion of cerato-
branchial I. In poinsetti the profundus extends
to the dorsal surface of ceratobranchial I. The
superficialis extends to the ventral surface of the
proximal portion of ceratobranchial I with ad-
ditional fibers to the medial and lateral surfaces.
In chn/sostictus it was single with a heavier con-
centration of fibers to the proximal section of the
first ceratobranchial. In this species the elonga-
tion of the neck has caused the ceratobranchial I
to lie parallel to the ramus of the mandible for a
longer distance than is found in other species of
Sceloporus. This elongation of the cerato-
branchial has caused the epistemohyoideus to
assume the shape of a parallelogram rather than
the fan-shape found in other species.

Bricham Young University Science Bulletin

In Crotaj)htjtus and Sauromalus this muscle
was found to have a inidventral fold which
causes it to have the appearance of two muscles.
Sanders illustrates ( 1874, fig. 1 ) a narrow, band-
like epistemohyoideus.

Jaw Musculature

Levator anguli oris

Origin: from the squamosal, postorbital and
dorsal portion of the tympanic crest.

Insertion: into the skin of the comer of the
jaw and the mundplatte.

This muscle lies directly below the heavy
infratemporal fascia. It is superficial to the ad-
ductor mandibularis extemus. The mundplatte
into which this muscle inserts, is a heavy sheet
of fascia which arises on the postorbital bar and
inserts onto the articulare. The mundplatte is
the only fascial sheet in Sceloporus which is
underlain by a fold of skin.

In Ctenosaura and Crotaplit/tus the levator
also takes origin from the medial surface of the
infratemporal fascia.

Adductor mandibularis extemus

Origin: from the posterior portion of the
postorbital and squamosal.

Insertion: onto the lateral surface of the pos-
terior third of the ramus of the mandible.

This is the large muscle at the angle of the
jaw. It is dorsal to the pterygomandibularis,
anterior to the depressor mandibularis group
and deep to the levator anguli oris. It is covered
by the temporal fascia and the levator anguli
oris so that it is not visible directly. In Scelopo-
rus it is not easily divided into the component
elements and only two portions were found.
Three were reported from Ctenosaura, Crota-
phijtus and Sauromalus.

Depressor mandibularis

Origin: from the dorsal aponeurosis.

Insertion: into the back of the mandible.

The group of depressor mandibularis muscles
appears on the lateral surface of the head and
neck. They lie deep to the constrictor colli, su-
perficial to the epistemo-cleido-mastoideus and
axial muscles and anterior to the trapezius. In
Sceloporus three muscle slips are separable. The
anterior two are usually divided from the pos-
teriormost and are termed the depressor mandi-
bularis. The posterior section is usually called
the cervicomandibularis in anatomical studies
but could be called the depressor mandibularis
posterior.

Depressor mandibularis anterior

Origin : from the back of the parietal and the
very anterior portion of the dorsal aponeurosis.

Insertion: on the articulare.

If heavily developed, this muscle will mask
the back half of the temporal opening. It ex-
tends posteriorly for a short distance from its
origin then turns ventrad and lies along the
posterior margin of the external auditory meatus.
The muscle fibers are replaced by a tendon ap-
proximately at the middle of the auditory meatus
which then passes below the tendon of insertion
of the depressor mimdibularis lateralis.

Depressor mandibularis lateralis

Origin: from the cervical portion of the dor-
sal aponeurosis.

Insertion: on the articulare.

The lateralis is much broader and thicker
than the anterior slip. It extends ventrad from
its origin and then turns anterior to its insertion.
Tlie insertion and ventral portion of the belly
cover the posterior portion of the anterior slip.

Cervicomandibularis

Origin: from the dorsal aponeurosis at the
level of the more posterior cervical vertebrae.

Insertion: in the fascia of the intermandi-
bularis posterior and on the back of the ramus.

From the broad origin the fibers of the
muscle extend ventrally and anteriorly to con-
verge at the insertion. The breadth of origin and
thickness of development are variable, but there
did not seem to be any correlation between
development and either size or sex. The de-
velopment of the muscle ranged from a broad,
straplike muscle to a thin sheet of fascia with
a few muscle fibers scattered through it. In one
individual it was not discernible. Its variability,
posterior origin, and weak insertion would indi-
cate that it is the weakest of the jaw depressors.

This muscle was similar in the other species
of Sceloporus, except that in one specimen of
ttmliilatus the fascia of insertion was into the
ventral midline at the level of the hyoid body.

Three section of this muscle were present in
Sauronuilus. Ctenosaura, Phn/nosoma. Iguana
and Crotaphijtus uislizeni and reticulatus. Cro-
taphytus collaris is reported as having only two
})undles which seem to be the anterior and lateral
slips. According to Sanders the cervicomandibu-
laris (his neuro-mandibularis ) was a well-
developed band which inserted onto the back of
the mandible. Camp (1923, Fig. 45) indicates
in Phripwsonw that the insertion was into the
fascia of the central part of the intemiandibularis
posterior.

Biological Series, Vol. 14, No. 1 Myology of Sceloporus

Shoulder Giedle Musculature

Trapezius

Origin: from the dorsal aponeurosis at the
level of the posterior cervical and anterior dorsal
vertebrae.

Insertion: onto the scapula by the overlying
fascia.

The anterior portion of this muscle Ues deep
to the cervicomandibularis. The posterior por-
tion is superficial to the latissimus dorsi and the
;ixial musculature. In two specimens some fibers
inserted into the lateral section of the clavicle
next to the origin of the epistemo-cleido-niastoi-
deus.

Crotaphijtus was also noted to have a partial
insertion onto the clavicle. Sanders (1874) re-
ported that the specimen of Phnjnosoma, which
he examined, had no trapezius. Jenkins and
Tanner ( 1968 ) found a narrow trapezius in
Phrtjnosoma phitiirlunos and douglassi.

Latissimus dorsi

Origin: from the dorsal aponeurosis of most
of the dorsal vertebrae.

Insertion: on the processus latissimus dorsi
just below the head of the humerus between the
scapular and coracoid heads of the triceps.

This is the largest of the superficial dorsal
muscles. The posterior border is sometimes dif-
ficult to determine as fibers running at the same
angle are present in the fascia to the level of the
sacrum. Its borders are further disguised by a
heavy concentration of melanophores which oc-
cur in the surface fascia of the entire body.

In poinsetti the latissimus dorsi was only one
fiber thick so that the iliocostalis and tlie tendons
of the spinalis and semispinalis were visible.
The insertion fibers appeared to be as heavy,
however.

In Phnjiwsoma, Sanders (1874) found the
latissinius to be only a narrow band arising from
the level of the spines of the third and fourth
dorsal vertebrae. Jenkins and Tanner ( 1968 )
reported a broad latissimus dorsi originating
from the superficial dorsal fascia.

Serratus dorsalis

Origin: by three slips from the cervical ribs.

Insertion : on the medial surface of the scapu-
la dorsal to the insertions of the serratus ven-
tralis and subscapularis.

These muscles, which form part of the sling
by which the anterior limb is attached to the
body, are visible when the lattissimus dorsi and
trapezius are cut and the scapular elements are
pulled away from the body.

In Plinjnosoiiui Sanders (1874) found only
two slips to tlie dorsal section of the serratus.
Jenkins and Tanner ( 1968 ) reported three slips
of this muscle.

Serratus ventralis

Origin: from the middle of the first thoracic
ribs.

Insertion: on the posterior margin of the
scapula posterior to the insertions of the serratus
dorsalis and subscapularis.

In Crotaphijtus the serratus ventralis had two
slips rather than one large muscle. In this case
they also took origin from the cervical ribs.

Subscapularis

Origin : from the medial surface of the supra-
scapula and scapula ventral to the insertion of
the serratus ventralis.

Insertion: onto the proximal, medial surface
of the humerus.

In Crotaphijtus, two slips of this muscle were
reported.

Biceps brachii

Origin: originates by two broad, flat tendons
from the ventroposterior margin of the coracoid.

Insertion: by a single tendon into the tendon
of insertion of the humeroantebrachialis.

This is the largest muscle of the anterior sur-
face of the humerus. The anterior tendon de-
velops a small flat belly of muscle which runs
almost to the shoulder joint which it crosses as
a broad tendon just posterior to the insertion of
the pectoralis. This becomes the lateral belly of
the biceps. The tendon of the medial belly re-
mains tendinous from its origin, over the shoul-
der joint, luitil the belly develops approximately
one-third of the way down the humerus. The
two bellies join in the distal third of the brachi-
um and a single tendon moves over the elbow
to its insertion.

Robison and Tanner (1962, p. 16) consider
this unit to be composed of two muscles, after
Mivart's (1867) work on Iguana. Tlie anterior
belly is considered to be a separate muscle—
the brachialls inferior. Romer ( 1924 ) considers
the muscle to be single but doubleheaded. San-
ders considers the muscle in Phnjnosoma to con-
sist of the biceps and the brachialis anterior.
The inclusion of muscle fibers in the anterior
head as it passes over the coracoid is not men-
tioned elsewhere.

Coracobrachialis brevis

Origin: from the majority of the medial por-
tion of the coracoid.

Bkiciiam Young University Science Bulletin

Insertion: to the proximal two-thirds of the
medial surface of the humerus.

The origin is deep to the tendons of origin
of the biceps brachii. In Iguami Howell (1936)
found two slips of this muscle. In Phnjnosuvui
the insertion is to nearly the entire length of the
humerus.

Triceps brachii

This is the large muscle on the posterior sur-
face of the brachium. The origins of the various
slips are well separated, but the insertion is by a
common tendon which passes over the elbow to
insert onto the olecranon of the ulna and which
contains the sesamoid ulnar patella.

Lateral humeral head of the triceps

Origin: from the lateral surface of the proxi-
mal half of the humerus.

Insertion : into the common tendon.

This head of the triceps lies lateral to the
scapular head and joins witli it halfway down
the brachium.

Medial humeral head of the triceps

Origin: from the medial surface of the hu-
merus about halfway down the shaft.

Insertion: into the common tendon.

This head of the triceps lies medial to the
scapular head. It joins the other heads of the
triceps just tiefore the tendon crosses the elbow
joint.

Scapular head of the triceps

Origin: by a heavy tendon from the postero-
ventral comer of the external surface of the
scapula.

Insertion: into the common tendon.

This head of the triceps joins the lateral
humeral head at approximately the middle of the
brachium. It is joined to the coracoid head by a
sling tendon in all examined species of Sce-
loporiis. This tendon was not noted in Crota-
phijtus and Sauwmalus but was noted to be
shongly developed in Iguana by Howell ( 1936 ) .

Coracoid head of the triceps

Origin: by a strong tendon either directly
from the posterior portion of the medial surface
of the coracoid or from the stemoscapular liga-
ment.

Insertion: into the common tendon.

This is the most variable head of the triceps.
In clarki the head varied from strongly devel-
oped to completely degenerated. In the case
where the belly was completely degenerated the
tendon was still present. If the belly is present.

it develops at the level of the insertion of the
latissinms dorsi. The shp hes medial to the
scapular head and superficial to the medial
humeral head. The sling tendon was well de-
veloped in nui<iister, chnjsostictUf; and unduJatiis.
It was rather weakly developed in grammiais
and malachiticxis.

In PImjnosonui and Mivart's (1867) Iguana
account, the inner (coracoid) head of the triceps
receives a tendon from the latissimus dorsi, very
close to the latter's insertion. From the illustra-
tion, this appears to be the sling tendon.

In Sceloporus the sling tendon is connected
with a subscapular ligament and together they
fonn a complicated set of tendons. The sub-
scapular ligament usually has three points of
connection. The most lateral point is that of
a strong but narrow tendon which arises from
the posterior comer of the coracoid. The tendon
of origin of tlie coracoid head of the triceps
arises from this tendon a few millimeters from
its origin. The tendon mns medially where an
anterior tendon extends forward to the fascia of
insertion of the levator scapula profundus. The
original tendon then continues medially to the
anterior portion of the costocoracoid muscle.
The three points of connection then are the pos-
terior comer of the coracoid, the back of the
levator scapulae profimdus, and the sternum, by
the fascia of the costocoracoid.

Variations of this basic pattern were speci-
mens of clarki in which the coracoid head arose
directly from the coracoid in conjunction with
the subscapular ligament, and specimens of un-
dulatus hi which an additional slip of muscle
from the center of the first abdominal rib enters
the costocoracoid where that muscle becomes
tendinous and enters the subscapular ligament.
In grammicus the ligament proceeds directly
from the stemum to the coracoid without send-
ing an extension to the levator scapulae profun-
dus. This species also has the extra slip of the
costocoracoid.

Howell (1936, p. 200), in his dissection of
Iguana, notes a similar, but not identical, situa-

te
tion:

"Thus, in Iguana, the scapulo- and coraco-
triceps arise from a continuous sling or loop,
suspended laterally between scapula and hu-
merus and mediallv between stemum, coracoid
and scapula."

In the dissections of Crotaphytus and Sauro-
mahis, no mention is made of the sling tendon
between the scapular and coracoid heads of the
triceps but in their discussion of the costocora-
coid, Robison and Tanner (p. 20) note that "the
costocoracoid . . . inserts, just dorsal to the mid-

BionxjicAL Series, Vol. 14, No. 1 Myology of Scelopohus

region of the first sub.scapularis, on a ligament
which extends between tlie inner surface of the
sternum . . . and the anterior border of the
scapuhi."

Costocoracoid

Origin: from the inedial portion of the first
abdominal rib and from the lateral margin of the
posterior half of the sternum.

Insertion: to the back of the coracoid by the
stemoscapular ligament.

This muscle consists of fibers which arise
from the first rib and the sternum posterior to
the point from which the ligament extends to
the back of the coracoid and a broad, flat sheet
of fascia anterior to the point of attachment of
the ligament.

In grammiciis and utuJtilatus there is also a
slip from the center of the first abdominal rib
into the costocoracoid just before it becomes
tendinous.

Axial Musculature

Rectus abdominis superficialis

Origin: on the anterior margin of the pubis.

Insertion: on the last abdominal rib posterior
to the origin of the pectoralis.

This is the large superficial muscle of the
belly. The identity of the lateral boundaries of
of the rectus is lost in the middle of the belly as
fibers from the rectus are incorporated into the
obliquus extemus. In Sceloporus there are three
inscriptions in the body of the muscle.

Rectus abdominis intemus

Origin: from the symphyseal area of the
pubis.

Insertion: into the midventral line or, in
some cases, onto the last abdominal rib.

In clarki the rectus intemus inserts into the
midventral line to the level of the xiphisternal
ribs, resulting in a triangular muscle. In other
species the muscle inserts into only the posterior
section of the midventral line. In g^rammiciis
and one specimen of chnjsostictus the insertion
was directly onto the xiphisternal ribs.

Obliquus extemus

Origin: from the middle of the body of the
dorsal ribs.

Insertion: into the midventral line of the
dorsal surface of the rectus abdominis.

The direction of the fibers is ventral and
posterior. Since fibers of the muscle do not arise
from the posterior dorsal vertebrae, there re-

mains a triangular space in front of die hind leg
in which the viscera is covered only by the peri-
toneum, the thin sheets of the obliquus intemus,
and the transversus abdominis.

In some individuals the slips retain individual
identity as they arise from the ribs almost to
their insertion posterior to the point where the
posterior margin of the latissimus dorsi crosses
the level of the insertion of the obliquus in-
temus. This tendency was also noted in
iimgister.

Musculature of Hind Limb

Ambiens

Origin: from the anterior surface of the
femur near the head below the insertion of
pubo-ischio-femoralis intemus.

Insertion: onto the patellar capsule and the
head of the tibia.

This muscle covers the leading edge of the
thigh. In Sceloporus the muscle is single and
simple. The origin is covered by the origin of
the femorotibialis.

Femorotibialis

Origin: broadly from the head of the femur
and the fascia of the pubo-ischio-femoralis in-
temus.

Insertion: broadly to the head of the tibia,
just proximal to the insertion of the flexor
tibialis intemus superficialis.

This muscle is visible on the ventral sur-
face of the thigh posterior to the ambiens and
anterior to the pubo-ischio-tibialis. It extends
diagonally from its origin beneath the pubo-
ischio-tibialis and the insertion of the adductor.

Pubo-ischio-tibialis

Origin: from the puboischiadic ligament
Insertion: onto the Hba by a broad tendon

which it has in common with the flexor tibalis

extemus.

This is the largest muscle of the ventral sur-
face of the thigh. The insertion is on the an-
terior surface of the tibia between the bodies
of the tibialis anterior and the extensor digitorum
communis several millimeters below the patella.
This muscle lies directly below the femoral pores
and bears their imprint in preserved specimens.

In poinsetti the muscle is less heavily devel-
oped and has a definite sHp between the main
portion and the flexor tibialis extemus. It in-
serts into the broad tendon of the main part of
the muscle by a thin band which remains visible
in the fascia across to the tibia. In malachiticus

10

Bricham Young UNrvERsiTv Science Bulletin

the common tendon enters the tibia just below
the knee joint.

Flexor tibialis extemus

Origin: from the tubercle of the ischium and

the ihoischiadic hgament.

Insertion: into the body of the pubo-ischio-
tibialis before it bec-omes tendinous.

This muscle occupies the posterior border
of the thigh. In all species examined the flexor
enters into the pubo-Lschio-tibialis before that
muscle crosses the knee, although there is some
variation in the distance above the knee the
junction occurs. In some specimens of clarki
some of the more posterior fibers of the flexor
enter the tendon rather than the muscle. In
iindulatiis the more posterior fibers give rise to
a small, independent tendon which crosses the
knee posterior to the larger tendon.

In Plmjnosonm and Igxiana the flexor tibialis
extemus crosses the knee independently.

Flexor tibialis intemus

Origin: from the iUo-ischiadic hgament deep
to the origin of the flexor tibialis extemus.

Insertion: by three tendons on the head of
the tibia.

The flexor tibialis intemus spUts into tliree
bellies at the level of the head of the femur.
The largest belly inserts broadly to the head of
the tibia immediately deep to the common ten-
don of the pubo-ischio-tibialis and the femoro-
tibialis. Of the two smaller bellies, one belly
develops a long, narrow tendon two-thirds of the
way down the thigh which inserts into the
posterior surface of the head of the tibia. This
is the deepest slip. The third slip is slightly
posterior to the other tvvo and develops a tendon
of insertion just proximal to the knee joint.

Popliteus

Origin: from the postero-lateral surface of
the head of the fibula.

Insertion: into the tendon of the flexor hallu-
cis longus.

In Sceloporus the popliteus is an extremely
short muscle which arises directly from the bone.
The fibers wrap around the tendon of the flexor
hallucis longus. This is almost the mammahan
condition rather than the normal reptilian con-
dition of a well-developed popliteus.

Gastrocnemius

Origin: the lateral head from the head of
the fibula, the medial from the head of the tibia.

Insertion: by a common tendon into the plan-
tar aponeiu-osis.

In Sceloporus this muscle is two-headed and
two-bellied but has a single insertion and will
be considered as a single muscle. In some cases
the muscle has been considered as two mus-
cles—the soleus and the gastrocnemius. There
has been little agreement over which is which,
but most recently George ( 1948 ) called the fibu-
lar head; the soleus.
Flexor digitoriuni brevus

Origin: from the tarsals and the plantar apo-
neurosis.

Insertion: to the ventral surfaces of the
phalanges.

This muscle is superficial to the flexor digi-
torum longus. The palmar aponeurosis is present
but is not as heavily developed as has been re-
ported from other lizards. Also, there is no sesa-
moid bone found, although Cope (1892, p. 196)
states that "In all the nomial Lacertilia the ten-
dons of the flexors of the digits are combined on
the palm and the point of junction is occupied by
a larire. flat sesamoid bone."

DISCUSSION

Within the genus Sceloporus, or at least in
the species examined, it can be seen that the
myology is generally very constant. Those mus-
cles which are constant within the genus are
essentially those which are constant within the
members of the family which have been exam-
ined.

Etheridge (1964) has indicated that Srelo-
ponts is an advanced iguaiiid in comparison to
other iguanid genera by virtue of ostcologieal
characters. Only a limited amount of myological
comparison can be made because of the small

number of myological studies available within
the family.

The muscles in which the greatest variability
occurred were the intennandibularis group, the
constrictor colli, the epistemo-cleido-mastoideus,
the epistemohyoideus, the coracoid head of the
triceps, the costocoracoid, and the flexor tibialis
extemus. It will be noted that most of this varia-
tion is in the neck region.

The variation in the intermandibularis was
used by Camp ( 192.3 ) in the construction of his
lizard classification. The results of the present

Biological Series, Vol. 14, \o. 1 Myology of Sceloporus

11

study are not always in agreement with his con-
clusions. According to Camp, Sceloporus is one
of the genera in which separation of the pro-
fundus and superficialis bmidles of the inter-
mandibularis anterior has occurred, and he states
(p. 370) that "in the iguanids, Callisaunts, Unia
and Uolhrookia, alone, is the superficial layer
absent." It was not noted in any of the species
of Sceloporus dissected. According to Camp, the
number of interdigitations of the intermandibu-
laris posterior with the mandibulohyoideus I
can be used for purposes of classification be-
cause they are regular and equal. In clarki alone,
the number of interdigitations ranged from three
to seven and usually were not the same in num-
br for the two sides. Therfore, the use of this
muscle for classification purposes in Sceloporus
does not seem warranted. Camp's criterion of
the primiti\'e iguanid throat musculature is the
concUtion in which there is a higher number of
interdigitations of the posterior slip and the lack
of or slight separation of the two slips of the
anterior slip of the intennandibularis. In this re-
spect, Sceloporus would maintain its place as one
of the primitive iguanids.

The variable development of the constrictor
colli is understandable if its broad development
is associated with the throat fan of Iguana, while
the absence of this throat flap in Sauromalus and
the other ground iguanids is accompanied by a
narrow constrictor colli. The variability of the
muscle is less easUy understood in the genus
Sceloporus. It is broadly developed in the tree-
living clarki and the rock-living nuigister. It oc-
curs as a narrow band in nmlachiticus and vari-
abilis which are quite far removed from each
other according to Smith (1939). It apparently
has no ecological and phylogenetic basis al-
though more data are necessary to draw any
conclusions. The behavior of the various species
of Sceloporus is not well enough known to at-
tempt any correlations.

The comparative development of the inter-
mandibularis posterior and the constrictor colli
seems to follow no patterns. In Sceloporus, the
development of both muscles is variable with
no relation between the two. In Iguana they are
both heavily constructed. In Sauromalus both are
narrow bands. In Crotapht/tus and Ctenosaura
the intennandibularis is broad and the constric-
tor colli is a narrow band.

The variation of the epistenio-cieido-mastoid-
eus seems to be without correlation. All species
of Sceloporus examined, except grammicus, had
a double-headed muscle. The variation in the
size of the head from the epistemohyoideus
superficialis was negligible.

The muscles connecting the pectoral girdle
and the hyoid apparatus show a great deal of
variation. There is controversy among anatomists
as to the nature and identification of these mus-
cles. Camp illustrates (fig. 42-47, 53-65) the
muscles he calls omohyoideus, stemohyoideus
and stemothyreoideus and shows the great di-
versity of this group of muscles. The three are
not always present simultaneously. Of the iguan-
ids studied Camp indicates that all three were
present in Prijnosoma. Avery and Tanner ( 1964 )
in Sauromalus and Robison and Tanner (1962)
in Crotaphijtus reported that there was a single
sheet of muscle folded back on itself at the
midventral line. Camp illustrates this condition
only in Gerrhosaurus (Fig. 63, 66). This folded
condition has been reported for no other iguan-
ids. If the muscle deep to the stemohyoideus
superficialis is the epistemothyreoideus, then the
litter muscle is present in poinsetti and spo-
radic in clarki.

There is a graduation of conditions in Scelo-
porus of the epistemohyoideus supericialis. In
niagister, undulatus, malachiticus, and most
specimens of clarki, the muscle was single and
inserted unifonnly on the entire length of the
first ceratobranchial. In chnjsostictus there was
a tendency for increase of fiber number to the
proximal portion of ceratobranchial I. In one
specimen of variabilis the muscle fibers were
confined to the proximal portion of ceratobran-
chial I and had shifted part of their insertion to
ceratobranchial II. Thus, concentration of fibers
to the medial portion of the hyoid occiu^s in one
of Smith's basic divisions of Sceloporus while
the species with insertion into the whole of
ceratobranchial I occur in the other branch.
The tendency to develop the deeper slip is
noted only for clarki and poinsetti which are
fairly closely related, according to Smith, and
would be an independent evolutionary trend.

Perhaps the most interesting of the muscle
variations found in this investigation is the varia-
bility and the relationships of the subscapular
ligament and the related muscles— the coracoid
head of the triceps, the costocoracoid and the
levator scapulae profundus. In the simplest con-
dition the ligament seems to be the tendon of
insertion of the costocoracoid in conjunction
with a heavy fascia from the anterior edge of the
sternum ( Robison and Tanner, 1962 ) . In Iguana
( Howell, 1936 ) , the tendon of the coracoid head
of the triceps has entered into the ligament.
Howell states that the ligament is presently in
Cijclura but is less strongly defined, and does
not elaborate. Sceloporus has added the further
refinement of the inclusion of the levator scapu-

12

Bricham Young University Science Bulletin

lac profundus. At this stage of investigation the
only attempt at correlation can be to say that
the ligament seems to be more strongly and com-
plexly developed iii aboreal iguanids than in the
ground dwellers.

The heavy development of the flexor tibialis
extemus in Scelopoms and its insertion on the
body of the pubo-Lschio-tibialLs chffer from other
iguanids and other lizards. The heavy tibial
flexors, which also act as femoral extensors, are
used by Snyder (1954) in his analysis of the
locomotor components in lizards. He indicates
that Scelopoms is a natural quadruped, with no
possibility of achieving the liipedality of other
iguanids. The flexor tibialis extemus was the
most highly developed in clarki; less developed
in magister and poinsetti; moderately developed
in undulatiis and nialachiticus; and rather weakly
developed in variabilis. Thus, the decidedly ar-
boreal clarki has the most heavily developed
flexor while it is rather weakly developed in
the ground-dwelling variabilis. This muscle
would be useful in a half-contracted position in
the maintenance of posture on a tree trunk in a
highly arboreal animal.

Avery and Tamier ( 1964, p. 28 ) have con-
structed a natural group of the genera Sauro-
malus, Ctenosaura, and Dipsosaurus which
share the characteristics of ( 1 ) an intermandi-
bularis anterior and posterior, (2) a single-head-
ed epistemo-cleido-mastoideus, (.3) a depressor
mandibularis group with three muscles and, (4)
an herbivorous diet. They state that, in their
opinion, Scchjwrus differs onlv in that its spe-
cies are omnivorous or carnivorous. Further work
by Avery has indic;\ted that Scelopoms is not
very close to the iguanines (pers. comm. ).

From the investigation herein described it
would seem that the relationship of Scelopoms
to these genera is not as close as is indicated.

1. Scelopoms lacks an intermandibularis an-
terior supcrficialis which is present in these
genera and the posterior and anterior slips of the
muscle are in contact rather than divided.

2. The epistemo-cleido-mastoideus usually
possesses a slip from the surface of the epi-
stemohyoideus supcrficialis and cannot be con-
sidered single-headed.

3. The third head of the depressor mandibu-
laris group, or cervicomandibularis, is usually
weakly developed and is sometimes absent.

The folded nature of the epistemohyoideus in
Sauromahis would seem to be a basic difference,
as is the simple nature of the subscapular liga-
ment.

These muscle differences would indicate that
Srclo])onis is not too closely related to Sauro-
iii'ilus and Ctenosaura. This is in agreement with
Etheridge's findings (1964).

Crotaplu/tus is considered by Etheridge to
be fairly closely related to the sceloporine lizards.
The myology is more similar to that of Scelopoms
than is that of Sauronuilus. but there are still

TABLE I
List of abbreviations used in the figures.

a - ambiens

add - adductor

anif - adductor mandiljularis extemus

anri - ring muscle

l)li - branchiohyoideus

c - cloaca

cb I - ceratobranchial I

cb II - ceratobranchial II

cc - constrictor colli

ch - ceratohyal

cm - cervicomandibularis

CO - costocoracoid

dcm - caudac dorsalis

ecd - e.xtensor communis digitonim
ecm - epistemo-cleido-mastoideus
chs - epistemohyoideus supcrficialis
CO - obli(jiuis extemus

fdc - flexor digitonmi communis
ft - fcmorotibialis
fte - flexor tibialis extemus
fti - flexor tibialis intemus

g - gastrocnemius
gg - genioglossus

hb

hyoid body
hyoglossus

iap - intermandibularis anterior profundus

ifb - ischiofibularis

ip - intermandibularis posterior

it - ischiotibialis

Ip - lingual process

m.ind - mandible
mil 1 - mandibulohvoideus I
mh II - mandibulohvoideus II
mil II - mandibulohvoideus III

oh - omohyoideus

pb - peroneus brevis

pect - pectoralLs

pife - pubo-ischio-fenioralis

pi - peroneus longus

pt - pubo-ischio-tibialis

pterv - pter\gomandibularis

ra - rectus abdominis

si) - subscapular ligament
si - stemum

ta - tibialis anterior

tlsp - tendon to levator scapulae profundus

toe - tendon to origin of cor.icoid head of triceps

tp - transversus perinei

Biological Series, Vol. 14, No. 1 Myology of Sc.

ELOPORUS

13

PECT

Fig. 1. Ventral view of throat, superficial mu.seulature.

14

Bricham Young University Science Bulletin

MHE

PECT

Fig. 2. Ventral view of throat, second layer.

Biological Series, \'ol. 14, No. 1 Myology of Sceloporus

15

MAND

ANN

PECT

Fig. 3. Ventral view of throat, third layer.

16

Brigham Young University Science Bulletin

— MAND

Fig. 4. Ventral view of throat, deepest musculature.

Rioi.ocicAL Series, Vol. 14. No. 1 Myology of ScELOPOiiU.s

17

Fig. .5. Subscapular ligament and a.ssociated muscles.

18

Bricham Young University Science Bulletin

DCM

Fig. 6. Dorsal view of superficial musculature of leg.

Biological Series, Vol. 14. Ncx I Myology ok Sceloi'ohus

19

///fiuim

Fic. 7. Ventral view of superficial musculature of leg.

20

Bricham Youno UNrvERSiTY Science Bulletin

Fic. 8. Ventral view of leg, pubo-ischio-femoralis and flexor tibialis extemiis removed.

Biological Series, Vol. 14, No. I Myology of Sceloporus

21

some differences. The intermandibularis anterior
and posterior are not separated by fascia as in
Sauromalus, but thev differ from Sceloporus in
liaving a definite superficial slip of the anterior.
Oiilv tvvo slips of the depressor mandibularis are
present in some species of Crotii])htjtUfi, but three
are present in the other species, as in Scclojwnis.
More important are the folded nature of the
episteniohyoid muscle and die simple nature of
the subscapular ligament. The nature of the
flexor tibialis extemus is not known.

Etheridge (1964) considers Phrynosoma to
be very closely related to the sceloporine lizards.
It differs, according to Camp (1923), by having
a verv well-developed intennandibularis anterior
superficialis. According to the Sanders (1874)
the body musculature differs by the absence of
the trapezius, an extremely narrow latissimus
dorsi and a flexor tibialis extemus which crosses
the knee independent of the pubo-ischio- tibialis.

Jenkins and Tanner ( 1968 ) consider that the
myological peculiarities of Plirijnosovia consist
of "The divided nature of the M. stemohyoideus,
M. subscapularis II, and M. epistemocleidomas-
toideus, and the reduced condition of the M.
serratus, M. trapezius, and M. obliquus abdomni-
is extemus, and the expanded nature of the M.
branchiohyoideus . . . " . Therefore, the body
musculature is decidedly different.

It has been shown that there are discernible
differences between the species of Sceloporus.
Little pattern can be seen in the differences and
one cannot be projected until further work is
done.

Even between the closely related clarki and
magister differences can be seen. Magister pos-
sesses a constrictor coUi which is merged with
the intennandibularis posterior; the episterno-
hyoideus profundus is not present; and the ilio-
costalis is much larger than that found in clarki.

CONCLUSIONS

1. Sceloporus is not closely related to the
ground-dwelling Sauroiruilus, Ctenosaura and
Dipsosaurus.

2. Crotaphiftus is more closely allied to Scelo-
porus than to the ground-dwelling iguanines.

.3. The musculature of Phrynosoma is aber-
rant in several respects, while other aspects in-
dicate that its relationship is not as close to
Sceloporus as is that of Crotaphytus.

4. The high degree of development of the
subscapular ligament in all species of Sceloporus

and its associated development in Iguana, an ar-
boreal lizard, would indicate the possibility of
the basic stock of Sceloporus being arboreal.
This is born by the nature of the flexor tibialis
extemus.

5. The small differences in myology between
the species of Sceloporus would indicate the
presence of a rapidly evolving group with, as yet,
little morphological variation.

6. Differences in the myology of reptiles can
and do occur at the species group and genus
group levels.

LITERATURE CITED

.\n.\.\i, L..\. 1919. .A memior on tht- phylogeny of the

jam muscles in recent and fossil \ertebrates. Ann.

N.Y. Acad. Sci. 28:51-166.
Appleton, A.B. 1928. The muscles and nenes of the

post-;L\ial region of the tetrapod thigh. 1 and II. J.

Anat., London 62( .3) :.364-438.
.\vEnY, D.F. ."VND T.\NNER, W.W. 1964. The osteology

and myology of the head and thorax regions of the

obesus group of the genas Sauromalus Dumeril

(Iguanidae). Brig. Young Univ. Sci. Bull., Biol.

Ser. 5(3): 1-30.
BouLENGER. G.A. 1897. A revision of the genus Sce-

loponis. Proc. Zool. Soc. London 1897:474-522.
Bn.\i)LEY, O.C. 1903. The muscles of mastication and

the movements of the skull in Lacertilia. Zool.

Jahrb., Abth. fur Anat. u. Ont. 18:475-486.
Brock, G.T. 1938. The cranial muscles of the gecko-

A general account with a comparison of the muscles

of other gnathostomes. Proc. Zool. Soc. London,

Ser. B 1938:735-761.
Camp, C.L. 1923. Classification of the lizards. Bull.

Amer. Mus. Nat. Hist. 48:289-481.
Cope, E.D. 1892. The osteology of the L:icertilia.

Proc. Amer. Phil. Soc. 30:185-221.
1900. The crocodilians, lizards and snakes

of North America. Ann. Rep. U.S. Nat. Mus. 1898

(1900): 151-1294.
Davis, D.D. 1934. The collared lizard, a lalx)ratory

guide. Macmillan, New York. 57 pp.
DE Vis, C.W. 1883. Myology of Chlamtjdosaurus

kingi. Proc-. Linn. Soc. N.S. Wales 8:300-320.
Dumeril. A.M.C. and G. Bibron 1837. Erpetologie

general ( Histoire naturelle complete des Reptiles ) .

iV. Fain, Paris.
Edgeworth, F.H. 1935. The cranial muscles of the

vertebrates. University Press, Cambridge. 493 pp.

Bhicham Young University Science Bulletin

Etheridge, R. 1964. The skeletal morphology ;iikI
systematic relationships of sceloporine lizards. Co-
p'cia 1964(4):61()-631.

196.5. The alKlominal skeleton of lizards in

the family Iguanitlae. Heqjetologiea 21(3) .161-168.

1967. Lizard caudal vertebrae. Copeia 1967

(4):699-721.

Evans, F'.G. 1939. Tlie moqihology and functional

evolution of the atlas-a.xis complex from fish to

mammals. Ann. N.Y. Acad. Sci. 39(2):29-UW.
FuRisRiNciim, M. 1876. Zur vergleichenden Anatomic

der .Schultennuskeln hei Saurier und Crocodile.

Gegen. Moq)h. Jahrh. 1:636-816.
Cadow, H. 1882. Beitrage zur Myolog)' der hinteren

E.xtremitaten der Reptilien. Gegen. Morph. [ahrb.

7:329-466.
Gandolfi, H. 1908. Der Zunge des Agamidae und

Iguanidae. Zool. Anz. 32:569-580.
George, |.C. 1948. The muscular system of Uromas-

tix hardwickii Gray. J. Univ. Bombay 17(3):-123.
Gnanamuthu, C.P. 19.37. Comparative study of the

h)oid and tongue of some typical genera of reptiles.

Proc. Zool. Soc. London, Ser. B 1937:1-63.
Haas, G. 1960. On the trigeminus muscles of the

lizards Xctiosauriis grandis and Shinisuunm crocidi-

liints. Amer. Mus. Novitates 2017:1-54.
Haines, R.W. 1934. The homologies of tlie flexor and

adductor muscles of the thigh. (. Morph. .56(1):21-

49.

1935. Some muscular clianges in the tail and

thigh of reptiles and mammals. J. Morph. 58(2):
355-383.

Harris, V.A. 1963. The anatomy of the rainbow
lizard. Hutchinson and Co., Ltd. London. 104 pp.

HoTTON, N. 195.5. A survey of adaptive relationships
of dentition to diet in the North American Iguani-
dae. Evol. 53(1):88-114.

Howell, A.B. 1936. Morphogenesis of the shoulder
architecture. IV. Reptilia. Quart. Rev. Biol. 11:183-
208.

1938. Phylogeny of the distal musculature of

the pectoral appendage. J. Morph. 60( 1 ) :287-315.

Humphry, CM. 1872. Notes on the mu,scles of the
glass-snake (Pseudajms Pdlhisii ) . J. Anat. Physiol.
6:287-292.

Jenkins, R.L. and W.VV. Tanner 1968. Osteology
and myology of Phrt/nosoma p. platt/rhiiios Girard
and Phn/no.wmii d. heriwndesi Girard. Brig. Young
Univ. Sei. Bull., Biol. Ser. 9(4): 1-34.

JoLLiE, M. 1960. The head .skeleton of the lizard.
Acta. Zool. 41:1-64.

Lakjer, T. 1926. Studien iiber die trigeminus ver-
sorgte Kaumuskulature der Sauropsiden. C.A. Reit-
zel, Kopenhagen.

Lowe, C.H., C.J. Cole and J.L. Pa rrON 1967. Kary-
otype evolution and speciation in lizards ( genus
Sceloporus) during evolution of the North Ameri-
can de.sert. Syst. Zool. 16(4):296-300.

LuNDELius, E.L. 1957. Skeletal adaptations in two
species of Sceloporus. Evol. 11:6.5-83.

McMuRHicH, J.P. 1903. The phylogeny of the fore-
arm flexors. Amer. J. Anat. 2:177-210.

1903a. The phvlogen\' of the palni:ir nuiscu-

lature. Amer. J. Anat. 2:46.3-500.

Miller, M.R. 1966. The cochlear duct of lizards.
Proc. Gal. Acad. Sci. 4th Ser. .33( 11) =255-359.

MuART, St. G. 1867. On the myology of Iguana
tuberculata. Proc. Zool. Soc. London 1867:766-797.

1870. On the myology of Chaemcleon par-

sonii. Proc. Zool. Soc, London 1870:850-890.

Oei.ricii, T.M. 1956. The anatomy of the head of
Ctciwsaura pectinata (Iguanidae). Mi.sc. Pub. Mus.
Zool., Univ. Mich. 94:1-22.

RonisoN, VV.G. and W.W. Tanner 1962. A compara-
tive study of the species of the genus Crotaphytus
Holbrook (Iguanidae). Brig. Young Univ. Sci. Bull.,
Biol. Ser. 2(1):1-31.

RoMEH, A.S. 1922. The locomotor apparatus of cer-
tain primitive and mammal-like reptiles. Bull. Amer.
Mus. Nat. Hist. 46:517-606.

192.3. Crocodilian pelvic muscles and their

avian and reptihan homologues. Bull. Amer. Mus.
Nat. Hist. 48:.53.3-.552.

1924. Pectoral limb musculature and shoulder

girdle structure in fish and tetrapods. Anat. Rec.
27(2):119-143.

1942. The development of tetrapod limb

musculature-the thigh of Lacerta. J. Morph. 71:
251-298.

1944. The development of tetrapod limb

musculature-the shoulder region of Lacerta. J.
Morph. 74:1-41.

1956. The osteology of reptiles. Univ. Clii-

cago Press, Chicago. 772 pp.

Sanders, A. 1870. Notes on the myology of Plati/-
dactylu.s jiiponicii.<i. Proc. Zool. Soc. London 1870:
413-426.

1872. Notes on the myology of Liolepis belli.

Proc. Zool. Soc. London 1 872: 1.54-183.

1874. Notes on the myology of Phninosoma

Proc. Zool. Soc. London 1874:71-89.
1936. The lizards of the torquatus

coroitatum.
Smith, H.M.

group of the genus Sceloporus Wiegmann, 1828.
Bull. Univ. Kansas (24) : 539-693.

1937. A synopsis of the variabilis group of

the lizard genus Sceloporus, with description of
new subspecies. Occ. Pap., Univ. Mich. Mus. Zool.
3.58:1-14.

1937a. A synopsis of the scalaris group

of the lizard genus Sceloporus, with description
of new species. Occ. Pap., Univ. Mich. Mus. Zool.
361:1-8.

1938. Rem:irks on the st:itus of the

sulxspeeies

of Sceloporus undulatus, with descriptions of new
species and subspecies of the umluUitus group. Occ.
Pap., Univ. Mich. Mus. Zool. ;587:1-17.

1939. The Mexican and Central .American

lizards of the genus Sceloporus. Zool. Ser. Field
Mus. Nat. Hist. 26:.3-.397.

Snvdeh, R.C. 19.54. The anatomy and function of
the pelvic girdle ;ind hind limb in lizard locomotion.
Amer. J. Anat. 95:1-46.

1962. Adaptations for bipedal locomotion of

lizards. Amer. Zool. 2(2 ) :]91-203.

SioKKi.Y, P.S. 1950. The occurrence of an interme-
dium in lizards. Amer. Midi. Nat. 43:179-182.

ViaisLUYs, [. 1904. Ueber Kaumuskeln hei Lacertilia.
Anat. Anz. 24:641-644.

Wiecmann, A.F.A. 1828. Beitrage zur Amphibien
kunde. Isis von Ok.n 1828:369.

ZwAiTARi, E. 1908. Materiali per lo studio dell osso
ioide dei Sauri. Atti Relae Accad. Sci. Toriono 43:
1-10.

Cuf^ ' T LCicv^o3

Li:

Brigham Young University ^ '371

Science Bulletin harvard

UNIVERSITY)

BOTANICAL AND PHYSIOGRAPHIC

RECONNAISSANCE

OF NORTHERN YUKON

by

Stanley L. Welsh

and

J. Keith Rigby

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 2

JULY 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany,

Brigham Young University, Provo, Utah

Members of the Editorial Board:

Vernon J. Tipton, Zoology
Ferron L. Anderson, Zoology
Joseph R. Murdock, Botany
Wilmer W. Tanner, Zoology

Ex officio Members:

A. Lester Allen, Dean, College of Biological and Agricultural Sciences
Ernest L. Olson, Chairman, University Publications

The Brigham Young University Science Bulletin, Biological Series, publishes acceptable
papers, particularly large manuscripts, on all phases of biology.

Separate numbers and back volumes can be purchased from Publication Sales, Brigham
Young University, Provo, Utah. All remittances should be made payable to Brigliam
Young University.

Orders and materials for library exchange should be directed to the Division of Gifts
and Exchange, Brigham Young University Library, Provo, Utah 84601 .

Brigham Young University
Science Bulletin

BOTANICAL AND PHYSIOGRAPHIC

RECONNAISSANCE

OF NORTHERN YUKON

by

Stanley L Welsh

and

J. Keith Rigby

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 2

JULY 1971

Frontispiece. Soiitii<Mst .uioss tiic nduiii .isiim p.nt ot tlie Bam Mountains into Blow Rivor Pass. Linear trails of
caribou show in the trampled tundra in the foreground. Creeks in the foreground and intermediate distance
are tributaries at the head of Fitton Creek at approximately 6«°29' N: 1.38° 1.5' W. The Rieharilson Mountains
.show as faint ghosts beyond the sea fog blanketing Blow River Valle\ .

TABLE OF CONTENTS •

INTRODUCTION 1

Accessibility 2

TOPOGRAPHY 4

Old Crow Flats 4

Bam Mountains 8

British Mountains 9

Arctic Coastal Plain 9

Buckland and Bahbage Hills 12

Driftwood Mountains 12

WEATHER 13

STRATIGRAPHY 13

VEGETATION 15

Streamsides and Bars 16

Lake Margins 19

Tussock Tundra 20

Alpine Tundra 21

Boreal Forest 24

Maritime Vegetation 26

Aquatic Vegetation 26

COLLECTION LOCALITIES 28

CHECKLIST OF VASCULAR PLANTS 38

ACKNOWLEDGMENTS 63

REFERENCES 63

BOTANICAL AND PHYSIOGRAPHIC
RECONNAISSANCE OF NORTHERN YUKON

by
Stanley L. Welsh' and J. Keith Rigby-

ABSTRACT

The area of study is located in the northern
panhandle of the Yukon Territory, and includes
the British and Bam Mountains, Old Crow Flats,
and the coastal plain of the north slope. The
physiography and geology of the region is de-

scribed and the plant communities are enumer-
ated. An annotated list of 279 species, 15 sub-
species, 63 varieties, and 1 fonn of vascular
plants collected during the summer of 1970 is
included.

INTRODUCTION

The area of study is located within the north-
ern panhandle of the Yukon Territory (Fig. 1),
and extends southward 70 to 80 miles from the
shore of the Beaufort Sea across the British and
Bam mountains and Old Crow Flats to the Por-
cupine River, west and southwest of the north-
em Richardson Mountains. The area under
investigation includes the Bam Mountains and
Blow River valley on the east, and the British
Mountains and Old Crow Flats to the Alaska
border on the west (Bostock, 1948). In general
the area lies between latitudes 68° and 69° 39'
north and longitudes 138° and 141° west, in the
Blow River, Herschel Island, Demarcation Point,
Davidson Mountain, Old Crow, and Bell River
quadrangles. The collected area occurs in quad-
rangles 117 A-2-7, 10-15; 117 B-S-9, and 16; 117-
C 1, 8-9; and 117-D-3-5, of the National Topo-
graphic Survey of Canada.

The central part of the area is approximately
130 miles west of Inuvik, Northwest Territories,
and approximately 70 miles north of Old Crow,
an Indian village on the Porcupine River.

The British Mountains continue to the west
in Alaska as the Romanzof Mountains. The more
eastern Barn Mountains are west across the Blow
River valley from the northern end of the Ricii-
ardson Mountains, which are adjacent on the
west to the lower Mackenzie River delta.

We arrived at the landing strip on the gravel
bar in the Porcupine River upstream from Old
Crow ( Fig. 2 ) on June 17 and set up a tempor-

ary camp while awaiting the arrival of air trans-
portation to Sam Lake. Camp was established
June 18 at Sam Lake, in the topographic saddle
between the British Mountains and Bam Moun-
tains at the northeastern edge of Old Crow
Flats (Fig. 3). Bonney (Bonnet) Lake to the
southeast approximately 20 miles. Trout Lake
to the north approximately 45 miles (Fig. 4),
and other lakes in the Old Crow Flats area were
still frozen. Trout Lake opened up approximately
July 1, and Bonney Lake at about the same time,
McNeish Lake, east of the mouth of Firth River,
remained with ice over much of the lake until
mid-July. The small lake in the headwaters of
the Firth River, near the braided ice field of the
Firth River east of the Alaska border, was clear
early in the season on June 20 when we first
flew into the area. Sam Lake, and Firth Lake,
as we will call it here, are both shallow lakes
and apparently wamied more quickly than the
deeper clearwater lakes both north and south of
the British Mountains.

Timber for tent poles, stakes for tents, poles
for a dock, etc., were obtained from a small
grove of white spruce trees on the south flank
of hills approximately 4 miles northwest of Sam
Lake, or from the valley of Dog Creek and
Black Fox Creek, 5 to 10 miles to the southwest,
the northernmost woods in the area. There is no
available timber to the southeast of Sam Lake
for some distance. Trees sufficient for low tents
and dock constmction are available in the Firth

'Department of Botany. Brighani Young University. Proyo, Utah 84'»0I
-Department of Geology. Brigham Young University, J'rovo, Utah 84501

Brigham Young Univebsity Science Bulletin

BE A U

F O

d^

Old

Mf s.

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S E

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<

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Y U K

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Fig. 1. Regional map of collected area in northern Yukon.

River valley, in the vicinity of Firth Lake and
upstream for a short distance. Only willows are
growing in the lower part of the Firth and Mal-
colm river valleys where these rivers emptv onto
the north slope of the British Mountains and the
Arctic coastal plain.

We moved our camp onto a prominent point
on the northwestern side of Sam Lake on June
18-19 and experienced some difficulty with tent
pegs. First, wooden pegs would not function in
the permafrost which was at that time at the
surface, necessitating use of long metal spikes
(10-12 inches) for initial setting up of etjuip-
ment. However, as frost level lowered, the spikes
did not hold in the spongy ground, and longer
stakes were necessary to hold tents in an upright
position.

Accessibility

Access is currently onlv hv aircraft with
floats landing on the few lakes in the mountains

and along the mountain flanks. McNeish Lake,
Trout Lake, and other lakes along the north
flank, and Bonney Lake, Sam Lake, Firth Lake,
and other minor lakes along the south flank of
the British Mountains are usable by planes of
Otter size and smaller. Several lakes in Old
Crow Flats are also deep enough and large
enough for emergency use, but most are so far
from solid camp areas that they are impractical
for anything but transshipment points.

A very short semigraded strip is located at the
head of Fitton Creek near a mine prospect on
the east side of the Bam Mountains, northeast of
Mt. Fitton, but is soft during the summer. A
landing strip capable of handling a DC-3 is
Hearing completion at the village of Old Crow
on the Porcupine River and will give year-round
use. The first planes landed there during late
July of 1970 on the unfinished runway. This is
an improvement over the gravel channel bar in
the middle of the Porcupine River, 10 miles up-

Biological Series, Vol. 14, No. 2 Reconn-\iss.\nce of Northern Yukon

"1

/»V*

r-f

EL-IBI l7mJ

■'■fH

i

Fig. 2. Junction of Old Crow and Porcupine rivers. Second sandbar east ot Old Crow village (A) in Porcupine
River is site of landing strip and temporary camp ( B ) . Air Photo Division — Energy, Mines, and Resources —
Canadian Government air photo A 13140-3.3.

stream from town, where the airstrip was located
in earher years and was used during the early
part of the summer of 1970. It was usable only
during low-water stages of the river and is peri-
odically flooded, even during the summer, by
floods related to heavy rainfall.

Winter seismic trails cross the Blow River
Pass into Old Crow Flats from the northeast
from the Arctic slope east of the Bam Moun-
tains. A winter road was bladed across the Eagle
Plains and into Old Crow village during the
winter of 1969-70, from the Demster Highway

through the Ogilvie Range. It connects to Daw-
son and roads to the south.

Most aircraft of the region are based at
Inuvik, and both wheeled and float-ecjuipped
planes are available for service and charter.
Inuvik is the local center for shipping and con-
nections to regions outside the lower Mackenzie
River valley and delta region. It is the major
supply point for much of the northern Yukon
and adjacent Northwest Territories, and most
items required can be purchased through sup-
pliers there.

Brigham Young University Science Bulletin

Fig. 3. Sam Lake, northeast portion of Old Crow Flats, and Bam Mountains, with Black Fox Creek at the left
and its major trihutar)-. Dog Creek, in the center. Collecting localities: A = 55; B = 59; C = 58; D = 57; E = 60;
F==56. Air Photo Division — Energy, Mines, and Resources — Canadian Government air photo A 13383-172.

TOPOGRAPHY

The area includes the generally mountainou.s
to hilly region of the British Mountains and
Bam Mountains, the western foothills of the
Richardson Mountains, the western part of the
narrow Arctic coastal plain of Canada, and the
flat, marshy, Old Crow Flats (Fig. 5).

Old Crow Flats

Old Crow Flats covers a wide area of low
relief extending south from the British Moun-

tains and west from the Driftwood Mountains to
the Old Crow Range, north of the village of Old
Crow. It is approximately 40 miles across north-
south and 50 miles across east-west. The entire
flat is between SOO and 1,400 feet above sea
level and gently slopes to the south into the
Old Crow River, which drains the lake-studded
flats into the Porcupine River through a gap at
the eastern end of the Old Crow Range.

The flats arc covered with small lakes and
poorly drained marsh, much of the latter show-

Biological Series, Vol. 14, No.

Reconnaiss.\nce of Northern Yukon

Fig. 4. Trout Lake, north of the junction of Canoe (C) and Babbage rivers (B). Arrow marks collecting locah-
ty on Trout Lake. Air Photo Division — Energy, Mines, and Resources — Canadian Government air photo A
13383-164.

ing excellent patterned-ground development. In-
dividual lakes range from small ponds up to lakes
8 to 10 miles across. Most of the lakes are marsh-
bordered, and only a few have beach develop-
ment at all. Limited beaches are common at
the southern end of the lakes, evidence of domi-
nantly north winds.

Several of the major streams such as Black
Fox Creek, Timber Creek, and Old Crow River
which drain across the flats, are entrenched into
the fine-grained fluvial sediments and lake clay
and silt that form the flats. Entrenchment in-
creases toward the south into the major Old

Crow River drainage where streams may be as
much as 50 to 60 feet below the general flat
upper surface. Streams in the interior of the
flats may be entrenched 10 feet, but in the outer
peripheral areas entrenchment of 4 or 5 feet is
typical. Major streams entering from the moun-
tain slopes have produced belts of gravel, spread
by the meandering streams. These are tree-
covered with white spruce and cottonwood trees
in the central and southern part of the area, but
are willow-covered and barren, meander-scarred
plains in the peripheral areas. Old Crow Flats
is an area of easy winter road development, but

Brigham Young Univebsity Science Bulletin

Fig. 5. Map of collected area in northern Yukon.

Biological Series, Vol. 14, No. 2 Reconnaissance of Nohthern Yukon

Fig. 6. Old Crow Flats, along Timber Creek showing angular lakes with about the same directional development

as the ridge at left. Arrow marks collecting site 67. Air Photo Division — Energy, Mines, and Resources —
Canadian Covernment air photo A 1.3470-76.

a bottomless bog area during summer, and for
much of the year, it is a barrier to overland
movement.

Se\eral low "islands" rise above the general
marshv level of the flats (Fig. 6). These are
bedrock exposures of either Lisburne Limestone,
Kayak Shale or of the yoimger lower sandstones
of the Cretaceous sequence. These rocks are
folded and appear to be overlain unconformably
by the much younger lake silts and clays of the
flats. The islands rise as much as 400 to 500 feet
above the general level of the flats. Minor ice-
shoved gravel beaches produce limited dry-

ground ridges 5 to 15 feet high along the shores
of some lakes.

Lakes of the flats occur generally below the
1000-foot contour, where the flats have a slope
of only a few feet per mile. These lakes range
from angular rectangular to rounded and many
show a distinct lineation with a northwest-
southeast trend and an even more pronounced
northeast-southwest one. Such rectangular out-
line is particularly shown well in the south-
eastern and northwestern quarters of the flats,
but is also evident in the northeastern part. The
northwest-southeast trend is parallel to fold

8

Bhigham Young University Science Bulletin

trends in the hills in the southern margin of the
British Mountains and to the folded rocks in the
islands within the flats. One island, on Timber
Creek in the southwestern part of the Blow
River 1:250,000 (juadrangle, shows the pattern
and relationship fairly well. The sides are also
parallel and normal to the long axis of the Old
Crow Flats basin and could be joint controlled
in the fractured permafrost in the continuing
subsidence in the basin.

Sequences of lake-filling and destruction
show in the patterned ground and marshes of the
flats. Some of the shallow lakes have obviously
filled with marsh plants and are now virtually
destroyed. In some, the patterned ground is now

only a vague ghost in the tussock and willow
tundra, particularly around the higher ground of
the periphery of the flats.

There is some suggestion that the Old Crow
Flats basin is still actively undergoing minor
subsidence along the trough, for lakes away from
the trough appear to have been tilted slightly
toward the northwest-southeast axis of the basin,
with abandoned shorelines on the exterior basin
side and encroaching shorelines on the interior
side.

Barn Mountains

The Barn Mountains (Fig. 7) are a relatively
low. north-south trending range in the north-

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Fig. 7. 15ani Mountains, cast side, with wf stern tributaries of Blow Hiver. and with Twin Peaks (T), and Mt.
Fitton (F), near right center. Air Plioto Division — Energy, Mines, and Resources — Canadian Government air
photo A 14406-66.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

9

eastern side of the area of concern, west of the
Blow River and the northern end of the l\ichard-
son Monntains, and nortlieast of Old Crow Flats.
They rise to approximately 3500 feet, 2000 feet
above Old Crow Flats to the southwest, and
above Blow River valley to the east. The Barn
Mountains are bordered on the northwestern and
northern side by low hills that form a foothill
belt with the Arctic coastal plain.

The Bam Mountains are rolling, occasionally
angular, barren hills and low-rounded moun-
tains, carved on older fractured rocks which
break into angular talus and fragmental rubble
that blanket the slopes in fine debris. There
is little tundra development on the generally
steeper rock-strewai slopes. Several streams cut
or nearly transect the range so that the moun-
tains consist of essentially three separate masses:
one south of the headwaters of Boulder Creek,
a central mass between Boulder Creek and the
northern one in the headwaters of Anker Creek.
All three masses rise to approximately the same
exhumed, warped, peneplain surface. Highest
points in the range are generally concordant at
approximately 3000 feet, but one peak 3506 feet
high occurs in the southern part of the central
area, at the head of Boulder Creek.

Slopes drop off moderately fast along the
eastern side of the elongate triangular range,
with valleys developing on softer shale close to
the older more resistant rocks. Slopes of the
western side are less steep, partially because of
an overlapping sequence of Paleozoic limestone
which allows development of strong cuestas on
the alternating softer and harder beds. The softer
pre- and post-Lisbume shales erode to form
strike valleys flanking the inward facing Lis-
burne cuesta ( facing toward the core of the Barn
Mountains ) .

The Bam Mountains are approximately 16
miles long and 10 miles wide at their widest
on the northern part. They taper southward to
only a mile across near their southern tip north
of the major bend of the Blow River.

British Mountains

The British Mountains are an east-west trend-
ing range which is an eastward continuation of
the Romanzov Mountains of the Brooks Range
complex of Alaska. They are flanked by a series
of moderatelv low disconnected foothills on the
north and south, the fonner of which are termed
the Buckland Hills. The British Mountains are
bounded on the south bv Old Crow Flats, with
a series of isolated partially disconnected hills
and ridges of Paleozoic and Mesozoic rocks at
the transition.

British Mountains rise to a summit level of
approximately 3500 feet in the eastern part, but
to more mountainous expression and elevations
of as much as 5429 feet in the central part of
the belt near the Alaskan boundary, in peaks
between the Firth and Malcolm rivers. The in-
ner margin of the Arctic coastal plain on the
north and Old Crow Flats on the south are both
at elevations of approximately 1500 feet, giving
the mountains a relief of approximately 2000 feet.

At the Alaska-Yukon boundary the range is
approximately 50 miles wide, but it gradually
narrows until it is only 25 to 30 miles wide in the
vicinity of the Babbage River, at its eastern end,
including some of the small outlying dissociated
foothill ranges.

The British Mountains are cut into segments
by the Malcolm River, Firth River ( Fig. 8 ) , and,
at the eastern end, by tributaries of the Babbage
River. Broad passes between the headwaters of
Tliomas and Timber Creek from the south, with
the Firth River on the north, functioned as one
of the main north-south accessways in the west-
em part of the range. The broad Babbage River
valley and Dog Creek and Timber Creek passes
also offer relatively low elevation accessways
around the eastern part of the range.

General summit levels of the range plunge
eastward, perhaps as the generalized surface of
an exhumed peneplain at the base of the upper
Triassic. Summit-level erosional surfaces appear
to be at approximately 5500 feet in the western
end, approximately 4()00 feet in the central part,
and approximately 3000 feet in the area of the
Babbage River at the eastem end of the range.

Local topography in the western part of the
mountains is often precipitous, with sharp di-
vides and cliff-strewn slopes. In the area east of
the Firth River, however, slopes are more gentle
and summits are often rounded with a generally
subdued topography. Chert and quartzite ridges
often show sharp divides, but the doniinantly
softer rocks erode to less expressive forms.

Arctic Coastal Plain

The Arctic coastal plain (Fig. 9) stretches
southward from the margin of the Beaufort
Sea to the northern flanks of the Bam and
British mountains and a foothill belt, sometimes
referred to as the Arctic Plateau or the Buck-
land Hills, in the western part of the British
Mountains. The coastal plain is a region of low
relief, sloping from 1000 to 1500 feet along the
mountains to an elevation of less than 500 feet
at the base of the piedmont only 5 to 10 miles
iroin the mountains. The plain surface has even
a much lower gradient in the outer part of the

xo

Brigham Young Univehsity Science Bulletin

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Fic. 8. Firth River (F), along the coastal plain, 10 to 15 miles south of the Beaufort Sea, at its juncture with
Loney Creek. Arrow' indicates collection locality 6 upstream from Lonev Creek (L) and west of McNeish
Lake (M). Air Photo Division — Energy, Mines, and Resources — Canadian Government air photo A 13751-116.

plain. Three divi.sion.s can thus be recognized in
the plain: an inner piedmont, an intennediate
zone where fluvial processes arc still dominant,
and an outer zone where accretionary coastline
processes dominate.

Tlie inner piedmont slopes the steepest and
is a region of pediment development over the
soft Mesozoic shales and sandstones. Locallv, the
surface slopes approximately I(X) feet per mile,
or even steeper, in the immediate foothill belt,
and is less steep in the outer regions. Slopes in
the intermediate zone are usuallv less than 2.5 to
50 feet per mile and are covered pediment slopes

with gravel and sand blanketing the beveled,
slightly folded, softer, younger rocks. The slope
of the outemiost zone is even less vet terminates
in the embayed coastline-sea cliff development
of the shoreline.

The coastal plain is a region where broad
deltas are developing from each of the major
streams, such as the Malcolm, Firth, Babbage,
and Blow rivers. The Firth and Malcolm rivers
are producing deltas typical of steep headlands
with smooth fronts and multidistributarv fanning
channels. The liabbage l^i\er, on the other hand,
is producing a delta which is a miniature of the

Biological Sebies, Vol. 14, No. 2 Reconnaissance of Nohthern Yukon

11

Fig. 9. Arctic coastal plain, Beaufort Sea, and King Point (K). along the lower reache.s of Deep Creek and
Babbage River. Collecting locality 13 (C). Air Photo Division — Energy, Mines, and Resources — Canadian
Government air photo A 1436.3-55.

Mackenzie River to the east, filling an estuary
between moderately High protecting ridges. The
Blow River is emptying onto the flank of the
Mackenzie delta and is producing much the
same general topographic pattern as seen on the
larger structure.

Lakes are a common feature of the eastern
part of the outer, lower coastal plain. In general
these are shallow, muskeg-bordered lakes with
little through-flowing water. Most appear to be
kettles or the result of melting ice, with a few
related to abandoned channels of the major
streams as the latter have shifted their discharge

channels near the mountain front. A few of these
lakes would be serviceable bases for field par-
ties. In the west, McNeish Lake, east of the Firth
River, is deep enough and ice-free long enough
to function as a campsite in the late summer.
Most of the lakes are not usable, however, be-
cause they are surrounded by marsh and soft
ground.

Lakes are not common on the upper slopes
of the plain, but Trout Lake and the small lake
to the north are deep enough and large enough
to be serviceable for float plane-based operations.
These appear to be ice-melt lakes, possibly even

12

plunge-pool lakes in the ancient surface of the
Babbagc drainage.

Entrenchment of streams below the general
tundra surface is typical of all the streams drain-
ing northward from the British Mountains and
Barn Mountains into the coastal plain. These
major streams arc entrenched not only into their
own debris, but into the underlying bedrock as
well. For example, the Babbage River is en-
trenched 60 to 70 feet bi'low the general pedi-
ment surface on Jurassic Shale near Trout Lake.
The amount of entrenchment decreases coast-
ward, so that halfway across the 25-mile wide
plain, it is entrenched only 10 to 15 feet. The
degree of entrenchment varies somewhat in the
outer reaches of the river, but generally is less
than 3 to 5 feet where the dcpositional surface
of the delta begins 4 or 5 miles from the shore.

The Firth River and other more directly
emptying steeper streams show the same general
degree of entrenchment, apparently related to
uplift of the land mass. The Firth River cuts
across vertical slate and quartzite near the coast-
al plain border 20 miles upstream from the shore
and is entrenched 40 to 50 feet below the general
sloping plain level. Its banks step dowoi in a
series of spectacularly developed, cut bedrock
terraces. It is well entrenched for approximately
10 miles of the inner and medial parts of the
plains, but begins to fan and develop an intri-
cate lobate delta in the outer 10 to 15 miles of
its course. It is entrenched only 2 or 3 feet
throughout most of the lower gradient part of its
coastal plain course.

Four distinct terraces are recognizable on
most major streams, with the lower two as the
most prominent and with deepest entrenchment.
The vertical-walled entrenchment of the Mal-
colm, Firth, and Babbage rivers into bedrock
cut terraces is particularly striking at the moun-
tain front-coastal plain boundary.

Deposits of the Firth River terminate in a
rounded, lobate or arcuate deltaic coastline, with
an offshore lagoon and barrier island secjuence, a
suggestion of moderately active regression and
energy dissipation along the shoreline. The Mal-
colm River shows an almost identical pattern and
has even a more spectacularly fanning delta, ter-
minating in the typical lobate to crescentic sea-
ward margin, with a bordering lagoon and an
offshore barrier island secjuence, named Nunatak
Spit.

Abandoned beaches mark the uplands of the
coastal plain, scribing the flanks of the British
Mountains to elevations of 700 to 800 feet. These
appear to be only moderate-energy beaches of

Bhicham Young UxrvEHSfTY Science Bulletin

relatively short still-stand because of the imma-
turity of the beach elastics, as well as shallowness
of indentation and poor development of sea cliffs.
Some of the high levels may be kamelike features
developed where mountain streams banked
against Pleistocene sea ice from offshore or
against a lobe of continental ice from the Mac-
kenzie River.

The coastal plain varies from 6 or 7 miles
wide at its western development, north of the
British Mountains west of the Malcolm River, to
a broad plain 30 to 55 miles wide north of the
Bams Mountains, west of the Blow River valley.
It narrows again to the east around the northern
end of the Richardson Mountains where there is
only a minimum plain between the mountains
and the southwestern margin of the Mackenzie
delta.

BUCKLAND AND BaBBACE HiLLS

The Buckland Hills are on the north flank of
the British Mountains. The tenn Babbage Hills
is applied to the continuation of these mainly
dissociated hills of Late Paleozoic and Cretace-
ous rocks in the vicinity of the headwaters of the
Babbage River.

On the southern side of the British Mountains
the hills are commonly cuestas or unbreached
anticlinal ridges of Lisburne Limestone from
which the softer Jurassic and Triassic beds have
been stripped. Cuestas of Cretaceous rocks in
the structural saddle between the Barn and
British mountains also fomi features of relief that
rise above the softer shale valleys as distinct
linear features. Similar dissociated hills on the
northern side of the British Mountains are on
outliers of older Neruokpuk rocks and, in the
eastern part, on cuestas of Cretaceous sandstone
in the Crow River, Trout Lake, and Sleepy
Mountain region.

DiuFTWooD Mountains

The Driftwood Mountains are an elongate
north-south range along the east side of Old
Crow Flats, south of the Bam Mountains. They
are cored by ragged-weathering Lower Paleozoic
chert and siliceous shale, surrounded by gentle
cuestas of Upper Paleozoic and Mesozoic rocks.
They fonn a series of hills approximately 12
miles wide and 25 miles long which rise to an
ele\ation of slightly over 3000 feet, approximate-
ly 1500 feet above Old Crow Flats on the west
and 2000 feet above Driftwood River on the
southeast. They are the southwestern end of a
series of hills leading southwest from the main
Richardson Mountains east of Bonney Lake.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

WEATHER

13

Weather experienced during the .summer of
1970 can be subdivided into basic patterns or
periods. During tlie month of June, at least from
the time we arrived at Sam Lake area until the
first week in July, the weather was moderately
clear all the time, with only three one-day per-
iods when fieldwork was not possible because of
weatlier. The days were bright and clear, with a
consistent wind from the north or northeast.

Wind exhibited a diurnal variation related
apparently to the differences in temperature be-
tween Old Crow Flats and the north slope and
Arctic coastal plain. During the afternoon, fol-
lowing warming of the Old Crow Flats, air rose
over the flats and pulled cool air from the north,
producing moderatelv strong winds from ap-
proximately noon until 1:00 to 2:00 a.m. During
the period from 2:00 a.m. until late morning, the
air was usuallv moderately calm.

The several short-duration stonns in June
blew in from the northeast with wind-driven
rain, with the exception of one storm which blew
in from the northwest. In general the weather
pattern was from northwest to southeast or from
west to east for the entre summer.

During June the temperatures on the valley
bottoms and around Sam Lake were approxi-
mately 40° to 50° during low sunlight periods
at night and during the windy part of the day,
but wanned to 60° to 70° during the calm part
of the day. A maximum of 80° occurred during a
still period during the period of 24-hour sun in
late June.

With the disappearance of sea ice from near-
shore in the Beaufort Sea in mid-July, the gen-
eral weather pattern changed. For two weeks
during early July the wind was most commonly
from the south, off Old Crow Flats, with several
intense thundershowers visible on most after-
noons, both in the flats to the south and over the
higher part of the British Mountains to the west.
This general pattern held until approximately
mid-July when the major winds again came from

the north and northeast, bringing thick persistent
banks of sea fog onto the northern slope and into
the lower valleys of the mountains. Only at night
did fog reach onto the southern part of the
mountains in areas west of the major drainage
through Blow River valley. Occasionally fog
would accumulate in the Old Crow Flats area,
but commonly the Sam Lake area remained fog-
free even though surrounding areas were blank-
eted. Bonney (Bonnet) Lake was commonly fog-
blanketed, whde areas to the west were mod-
erately clear. Trout Lake and the lower end of
the Babbage River valley were often fog-blanket-
ed during late July and early August.

Temperatures remained in the 60s and 70s
during the davs in the area around Sam Lake,
and only when cold winds blew off the sea to
the north did temperatures in the mountains
drop much below that of the base camp. Temp-
eratures at night remained near .50° during the
early part of July in the base camp area, but
dropped into the 30s during the nights of late
Julv and earlv August. The first frosts occurred
July .30 and 31 during a period of calm following
a cold north wind.

Weather in late July and early August was
decidedly cyclic. Calm wind and bright clear
weather was generally followed, usually in less
than one day, with high cirrus clouds and in-
creasing winds usually from the south or south-
west. Cloud cover increased and winds calmed,
usually followed by a warm, steady rain, some-
times for 2 or 3 days. Wind velocity then in-
creased and swung from south to southwest, then
west. Patches of broken clouds usually followed
with a change of wind from west or northwest
into the north or northeast, with some increase in
clouds and much increase in local fog on the
ridges north of Sam Lake, in Blow Pass and
Babbage River Pass, and over the coastal plain.
This usually was followed by clearing and strong
northeasterly wind, with later calming and begin-
ning of bright, clear weather again. The entire
cycle usually took from 3 to 5 days to pass.

STRATIGRAPHY

Rocks of the region range in age from Late
Precambrian ( ? ) to Upper Cretaceous, with Ter-
tiary ( ? ) and Quaternary sediments over the more
consolidated part of the section. The section
breaks into four broad sequences of rocks: an

older Precambrain to Devonian(?) strongly fold-
ed argillaceous and cherty section included in
the Neruokpuk and equivalent Road River for-
mations ( Martin, 19.59; Norford, 1964; Norris et
(il., 1963; Reed, 1968); an intermediate Upper

14

Brigham Young Univebsity Science Bulletin

Paleozoic section which includes a basal clastic
st'(iuence of Devonian and Mississippian age
overlain by a carbonate secjuence, the Lisbunie
Limestone, of Mississippian and Pennsylvanian
age; and an upper secjuenee of clastic Jurassic
and Cretaceous rocks including the Kingak Shale
and unnamed Cretaceous formations (Jeletzky,

1961; Mountjoy, 1967b). A fourth sequence of
Fennian and Triassic rocks is exposed in the
western part of the area (Mountjoy, 1967a) be-
tween the Lisbumc Limestone and the Jurassic
Kingak Shale. A total thickness ot approximately
40,000 feet of beds is present in the area (Fig.
10).

Cretaceous

Cretaceous rocks undifferentiated

Gray shale and mudstone with moderately thick coarser grained sandstone
units in the lower part. Mudstone and shale are silcy throughout, with
some beds distinctly siliceous in the lower third. Sandstones of the unit
form prominent ridges and cuestas but mudstones and shales form tlie
lowlands of the northernslope and intermediate valleys.

Jurassic —

Triassic

Permian

Pennsylvanian

Mississippian

Devonian

Precambrian to
Silurian

KINGAK SHALE

Dark-gray to black, somber-weathering shale and siltstone that forms
broad valleys and tundra-covered slopes. Pan of the formation is
very siliceous and all of the formation contains ironstone concretions.

SHUBLIK FORMATION

Dark-gray, brown-weathering sandstone, limestone, and shale .
SADLEROCHIT FORMATION

Brownish gray- weathering siliceous limestone and calcareous sand
stone with considerable quartzite and chert.
LISBURNE UMESTONE

Light gray-weathering limestone and dolomite with considerable
dark-gray to black chert. Forms prominent cliffs and ledges, most
extensive exposures are in the western part of the area. Fresh
carbonate rocks are dark-gray to medium-gray and commonly
are abundantly fossiliferous.
KAYAK SHALE

Dark- gray to black shale and siltstone with interbedded siliceous
sandstone, congbmcrate and quartzite in the lower and middle
part witli calcareous shale and argillaceous limestone in the upper
part. Forms broad valleys and sbpes or passes.

KEKIKTUK CONGLOMERATE

NERUOKPLIK FORMATION

Interbedded black, green, and maroon slate with ^ay and ^ay-
grcen chert and quartzite; with local limestone and dolomite
lenses. Locally schistose and cut by monzonitic intrusions.

Fig. 10. Geological section in northern Vnkon.

Biological Series, Vol. 14, No. 2 Reconnais.sance of Northern Yukon

VEGETATION

15

The general aspect of northern Yukon is
dominated by vegetation of low relief, i.e., tun-
dra. The hills and valleys have the look of a
well-tended golf course (Fig. 11). The vegeta-
tion is not unifonn, however, it varies in cover
and in composition. The major vegetative con-
trol appears to be the amount of moisture that is
available. Substrate differences appear to influ-
ence the water supplv directlv, and thus the
plant cover indirectly. The limestone, shale,
sandstone, and siliceous conglomerates, slates,
and schists of the lower, rounded mountains gen-
erally lack the more lush vegetation of the ba-
jada.s, wet meadows, and stream courses of lower
elevations. Rock stripes, boulder patches, and
rock outcrops appear at first glance to be barren
of higher plants. They are vegetated by a sparse
cover of plants widely spaced. Even the lowlands
display a mosaic pattern in the green mantle of
the land. Frost-heaved patches of gravel of ir-
regular shape produce a varied pattern in the
tundra. The swales where water drains slowly
over the surface are bright green compared to
the slopes alongside them. Polygonal patterns ap-
pear here and there in the tundra, apparently
always associated with poorly drained sites. The

lower trenches of most polygons are moist or
have standing water in them and frequently are
more lushly vegetated than the surrounding area.
Tall vegetation is present only along the streams,
\\'here willows and an occasional cottonwood
stand a few feet above the surface. Southward
along the upper reaches of the drainage of the
Old Crow and rarely in the headwaters of the
streams which drain to the Arctic Sea, there oc-
curs a depauperate assemblage of the Boreal
Forest, chiefly along stream courses, and along
some south facing slopes. This forest extends to
about 69° 15' N'^ along the Firth River, at ap-
proximately 140° 25' W. Eastward, however, the
forest does not extend that far north, reacliing
only to about 68° 39' N along Spruce Creek, a
tributary of the Canoe River, at approximately
1.38° 43' W. The trees reach their best develop-
ment in the mountain system on south facing
slopes near the bases of low rounded hills. In the
Old Crow Flats there has developed a parkland
tundra or taiga, the interspaces of which is dom-
inated by a heathland of dwarf birch, willows,
and other shrubs. In some places the woodland
is really quite dense, the trees reaching a height
of up to 25-30 feet and a few inches in diameter.

\' 'I v-^i,,-^ ,.

Fig. 11. Northwestward from apprcximately 58°32' N; 1.38°27' W, along westward dipping cue.stas of Li.sbume
Limestone, at the northwest edge of the Ban) Mountains, in the headwaters of Wood Creek. Low vegetated
valleys are in the Kayak .shale, in part in fault-repeated sections. Tussock and alpine tundra and heathlands.

16

Brigham Young University Science Bulletin

Lakes dot the land in the Old Crow Flats but
are less common elsewhere; in fact, there are
very few lakes of any consecjuence in the moun-
tains. Numerous lakes, most lacking higher plant
life, occur on the coastal plain of the north slope.
Thus, aquatic vegetation is best represented in
the Old Crow drainage system.

Streamseoes and Bars

The effects of streams seldom reach far from
active running water. However, since the streams
tend to be braided, at least in the lower reaches,
the effect is often more broadly felt. Willows
of one to several species dominate the banks
(Fig. 12 and 13). Where meanders develop
there may be considerable development of wil-
loy dominated vegetation (Fig. 14 and 15). The
height of these riparian plants varies from one
foot to ten or fifteen, or less commonly to twenty
feet. The larger plants are the willow, Salix ala-
xensis, with densely hairy branchlets. However,
the Cottonwood, Popiilus bahamijera, is some-
times present ( Fig. 16 ) . Neither of these plants
reach far from the streamside. As soon as the
riparian boundary is exceeded, there is no more
SdUx (ilaxensis. Certainly Salix alaxensis is the

abundant, tall willow species along all of the
drainages, from high elevations down to very
low ones, except, of course, in the Old Crow
River area where other species become import-
ant, but even there Salix alaxensis coi.tinues as
one of the dominant species.

Other species of willow grow along the
drainages and grade in height backward from
the abundant moisture of the river or stream in-
to the drier heath or tussock tundra along the
margins. Here and there the alder, Alnus cri.ipa,
grows tall enough to add to the thicket along the
drainages and in some places such as along the
Firth River, at its junction with Muskeg Creek,
the alder forms a part of the heath vegetation
some distance back from the bank (Fig. 17).
Similar extensive growth of alder occurs along
the lower portions of the Blow River and in the
Old Crow Flats where river bars are formed
which do not receive an annual scouring. Bars
which receive annual scouring do not support
vascular plants, but are barren throughout the
growing season (Fig. 18).

Many plants occur in the openings among
the willows lining banks and bars. Here such
things as sweet vetch, milk vetch, buttercups.

u-M^K

U^*

1 !<■. i-. ^'imiii .ivai>> >llKlll liUnallKtl ruck Willi gl.lM-l IIRMIkU-IS .llul \\llllA\ llKll^lll^ llllUhHll.llf U ^tnlllR.l^t oi

Sam Lake at 68°23' N; 1.38°.38' W. Broad low country of Old Crow Flats forms the background. The fore-
ground is tussock tundra, with alternating heathlands and tussock tundra in the background.

Biological Series, Vol. 14, No. 2 Heconnaissance of Northern Yukon

17

.^k*

'^'

^/■^

^^■%^

?%:;"^^

•-<*.

Fig. 13. Downstream along Spruce Creek from almost the same position as Figure 23, at an elevation of appro.\i-
mately 1000 feet, Jurassic Kingak Shale is exposed in the gully margins.

chickweeds, and other plants grow in profusion
adding pink, yellow, and white to the color of
the gravel bar vegetation. Poorly drained sites
back from the riparian vegetation develop a
distinctive cover of plants. Drt/as, Empetrum,
Ledum, Arctostaplit/los, Betula, Vaccinium, ( vit-
is-idaea, and (//fgino.si/;n) and Salix dominate the
community. But secondary species brighten the
landscape with shades of yellow (Arnica and

Oxytropis), white and lavender (Castilleja),
blue (Mertensia and Lupinus), and pale cream
( Pedicularis capitata ) , and pink ( Pedicularis and
Iledt/sarum). The tiny flowers of the miniature
plants of Tofieldia and Carex add to the diver-
sity of flower types, and here and there are the
bright pink spikes of bistort and the inconspicu-
ous ones are viviparous bistort. This heath type
forms a dark, dull green stripe along the drain-

18

Brigham Young University Science Bulletin

Fic. 14. Jurassic and Cretaceous sand and shale at the junction of Canoe Kiver with tlie Bahbage River south ot
Trout Lake, taken from approximately 68°48' N; 138°45' W. The river terraces are veneered here and there
with a thin blanket of gravel. Tussock tundra (center), heathland (along slopes) and willow dominated bar
(foreground).

BiOL<x;iCAL Series, Vol. 14, No. 2 Reconnaissance of Nohthern Yukon

19

■"iS|B!!iS5WS^""

<;^

Ijt^gf-'

ifyt

..M

Fig. 15. Cretaceous shale and sandstone at approximately 68°55' N; 138°.3r W, along the lower part of the
Babbage River where the river has emerged from the British Mountains onto the Coastal Plain. Low country
in the background is carved on soft Cretaceous sediments. Willow, alder, and dwarf birch heathland in fore-
ground and along stabilized bar.

age systems in the tundra and can be picked out
from some distance by its contrast with the pale
or gray green of the tussock tundra ( Fig. 19).

Lake Margins

In the tundra, the lakes are generally small
and commonly shallow. The margins are fre-

quently occupied by emergent plants. Principal
among there are the sedge, Carex aquatilis. and
the grass, Arctophila fuloa. Tlie wet meadows
at the waters edge are dominated by the cotton
grass, Eriophonim angtistifolium, and back from
the waters edge the vegetation is dominated by
a thicket of low willows (Salix) and birches
(Betula glandulosa) . Interspersed among the

20

Brigham Young University Science Bulletin

willows and extending into the wet meadows
are Andromeda polifola, Cardamine purpurea,
PedicuUiris kanei, P. sudetica, P. lahradurica, and
Ruhus chamaemorus. The extent of the wet
meadow and the willow-birch thicket is con-
trolled by the topography features of the lake
basin. Also, the height of the willows and birches
is apparently a function of topography and snow
depth, with the height of the inner ridge of the
lake basin and the height of the snow bank
fonned adjacent to the ridge limiting shrub
height. Ledtnn decumhcivi and Empetrum nig-
rum fill in between the willows and the birches.
In some lakes there is an accumulation of several
feet of organic debris along the beaches. The
aquatic and seniiacjuatic plants (X'cupy the or-
ganic fill in the same se(juence as in lakes which
have rocky basins forming the beach. The shal-
low lake at collecting locality 72 has two organic
debris littered beaches ( Fig. 20 ) .

Tussock Tundra

Landward from either lakes or streams, there
is developed a hummocky tundra which covers
more land surface than any other vegetative type.
Even here there is much variation in the com-

position of the vegetation. Near streams and
lakes there is a transition from the tallish, lush
vegetation characterizing these sites to the hum-
mocky tundra type (Fig. 21 and 22). Slopes
along streams are characteristically transitional
to tussock tundra also (Fig. 23). The transition
may be completely subtle or it may be abrupt.
Generally, Salix alaxensus gives way abruptly
along the wet sites, but the other species of wil-
low which may grow to a height of a few feet
along the streams continue to dominate the
rounded hummocks of the tundra, but seldom
reach a height much above the hummocks. They
grow there with Vaccinium vitk-klaea, V. uligin-
ostim, Betula glanduJosa, Drija.s integrifoJia, Ar-
ctostaphtflos alpirui, Pedicularis capitata, P. lab-
radorica, P. kanei, Oxijtropis maijdeUiana, Card-
amine digitata. Polygonum hi^orta, and Carex
species.

In higher, better drained ridges and slopes,
the willow hummocks give way to a cotton grass
tussock tundra dominated by Eriopliorum vagin-
atuin and Carex species. Shrubs are not lacking
in the tussock tundra, but their role is subordin-
ate to the cotton grass. In fact the same species
of shrubs outlined for the wetter hummocky
tundra occur in the tussock tundra as well.

jf^.i

Fig. 16. Undercut meander Ixnd along Black Fox Crock in the cast central part of Old Crow Flat at 139°.'50' W;
68°07' N at Locality 68. Elevation of ahout 900 feet. Willow and Cottonwood dominated bar, with taiga and
heathland in background.

Biological Seiues, Vol. 14, No. 2 Reconn.'Vissance of Northern Yukon

21

Fig. 17. Southwest across the Firth River to Firth Rock. Collecting locality 15, at 68°47' N; 140°28' W and at
approximately 1400 feet in elevation is on the gravel bar in the center of the stream. Vegetation in the fore-
ground is on river gravel terraces. Firth Rock is composed mainly of Lisbunie Limestone. Firth Rock rises
appro.ximately 1000 feet above the ri\er valley. Salix, Bctula. AInus heathland and taiga along valley bottom,
river bank, and bar.

Swales and moist flats in the tussock tundra
appear green and lush by comparison to the well-
developed tussock tundra. The green of the
sites is due in a large part to the presence of
the cotton grass Eriophorum angustijolium, and
in some places to the presence of the horsetail,
Equisetum arvense. Where the rehef is low and
drainage is poor, there is developed a topography
made apparent by the presence of polygonal
figures in the tundra (Fig. 24 and 25). Each
polygon is outlined by a trench, and thLs is often
filled with water or is, at any rate, more moist
than the surrounding area. Here grows Erioph-

orum angustijolium and Carex species, often in
a mass of Sphagnum. Other plants in the poly-
gons are PotentiUa pahistrls, Pedicularis species
and Menijanthes trifoUata. In drier polygons the
trenches often support a community of plants
similar to that along a stream channel, i.e., with
Salix, Betula, Empetrum, Ledum, and Vaccinium.

Alpine Tundra

Mountains in the northern Yukon appear pale
green to gray or else they are green only in
small areas. Talus and rockstripes are often bar-

22

Bricham Young University Science Bulletin

ren of plants or are occupied by clmnps or
stripes and patches of Dri/as, Empetrum, Betuhi,
and Carex. Talus slopes are evidentiv in motion,
and vegetation is commonly oriented in stripes
with an up-down slope axis (Fig. 26). Ridge
tops are characterized by widely spaced plants
of PotentilUi uniflora, Draha caesia, D. lactea,
A.strat^alus australis, Oxijtrof)i.<i ni'^rcscens, Saxi-
jraga reflexa, S. tricuspidata, Douglasia arctica,
D. ochotcnsis, Parn/a ntidicauUs, Phlox sihirica,
and Erctrichium naittiDi. Carex, Kobresia, Fes-
luca, and Poa are the principal monocots. Again,
density and composition varies, with the most
dense vegetation in swales and on north facing
slopes. Seeps and springs are densely covered
with Salix, Betuhi, Empetnim, Drijas, and Carex
and numerous herbaceous species. The herbs in
the seepy areas are the same as those in wet
meadows, i.e., species of Pediculari.s, Cardamine,
Anemoiw, Senecio, Carex, Kobresia, Dodocath-
eon, Lagotis, Parnja, Poleinonium, and Mijosotls.
From the specimens collected, it seems that
each geological stratimi exposed in the alpine
tundra supports a distinctive flora. In regions
where vegetative cover is well developed, the
plants are insulated from the geological forma-
tion by organic debris, permanently frozen

ground, or by alluvium. Geological control of
v(>getation is best demonstrated on outcrops of
formations on ridge crests and mountain tops
where conditions for growth are poor at best.
Some species seem to do well on any substrate,
regardless of the nature of the stratum. Others
occur principally on limestone fonnations ( Figs.
27, 2<S, 29 and .'30), e.g., Alijssiim americanum;
Arenaria rossii var. elegans; Brai/a piirptirascens;
Carex petricosa: Festuca braclnipiujlhi; F. ovina;
Lychnis apetala; L. furcata; and Saxifraga caes-
pitosa. Those which occur most often on slate
and schist (Figs. 31 and 32) include: Arenaria
macrocarpa Pursh; A. rubella; Douglasia arctica;
Saxifraga esrhscholtzii; and Senecio fuscatus. On
siliceous conglomerate, granite, and sandstone
(Figs. 33, 34 and 35), the list of apparently re-
stricted species include Loiseleuria procumbens;
Potentilla elegans; and Saussurea angustifolia.
Those which grow primarily on shale (Fig. 30)
include: Saxifraga exilis and S. serpt/IIifolia. Sili-
ceous conglomerates and sandstones are defin-
itely poorer in species than are the limestones.
The shales and schists are intermediate, but the
limestones support the greatest number of spe-
cies in alpine sites.

\M^uj^^

'«»-

Jb ...

0^ -■---;^^

1 i!.. i.^i. M l.n.ility .'56. Uiiniit; tlic nia.\iiiiuiii im Itw.iti i lilack Ko.\ (;H<k luiis imi tlii_' cxpu.snl i;i.i\(.l iii tin'
foreground.

BioL()r;K.AL Series, Vol. 14, No. 2 Reconnaissance of Nohthehn Yukon

23

Fig. 19. Southwest along one of the numerous willow-bordered drainages southeast ot Sam Lake in a tundra
heathland flat which rims the southwestern edge of the Bam Mountains. Photograph is at appro.ximately
68°2.3' N; 1.38°27' W. Some of the marginal small lakes at the edge of Old Crow Flats show in the back-
ground. Helicopter antenna shows tov\ard the left, and an old seismograph trail shows in the foreground.
.Mternating stripes and patches of Salix and Betula (dark) and Eriophorum, Carex, and Salix tundra (light)
with large areas of tussock tundra along the drainage margin.

24

Bbigham Young University Science Bulletin

Boreal Forkst

Tlic Bort-al Forest is only poorly developed
in northern Yukon. Three main types are reeog-
nizable, e.g., white spruce stands on south facing
slopes, open parkland tundra (i.e., taiga), and
riparian woodland. The first type, the spruce
woodland on the southern bases of hills (Fig.
36) and on south facing slopes in valleys (Figs.
37 and 38 ) , is distinctive. This woodland is made
up of isolated stands which vary in size from a
few trees to rather extensive forests. Southward,
in Old Crow Flats, the stands on hills tend to be-

come continuous with the parkland tundra and
riparian wo(k1s (Fig. .39). The species composi-
tion in the spruce stands is the same as that of
the parkland tundra, differing mainly in the fre-
(juency of species encountered. In both the dom-
inant tree species is the white spruce, Picea
f^latica.

The riparian woods ( Figs. 40, 41, 42, and 43)
are distinct even though they grade into both
of the other tvpes of woodland. White spruce is
the dominant conifer, and other tree species are
represented by white l^irch, Betula papyrifera.

/^

f" ,

y /

Fic. 20. Unnamed lakes in Old Crow I'lat about 16 miles soutli of Sam Lake. Collecting locality- 72 is ak)ni; the
south side of the lake with two developed beaches (A), and 73 is atop the prominent ridge (B) between the
two larger lakes.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

25

'?««l.

Fic. 21. SouthwL'stward aloni; liroail open \allf\ lu-.ii the In-. id cil Trail Himi .it approximately 68' 5.5' ,\, l.j!) ."iS'
W. Rocks in the background are principally of Lisburne Limestone. Streaks are reflections from the helicopter
bubble. Tussock tundra.

gMMjlli^:

•<i«kifl^^^

*«f*J««,t!;i

iftfia^^

Fig. 22. Stream teiraLt-. al.aig tin- Caiidc iiisei suutli ol limit Lake at appioMiiiatcK iiH 47 ;\; l.j8 46' W.
Rocks in the foreground are Jurassic shale. Hills in the background are held up by Cretaceous sandstone.
Tussock tundra on patterned ground.

26

Brioham Young University Science Bulletin

Fig. 23. A view looking upstream along Spruce Creek from 68°38' N; 138°38' W
expo.sure.s in the hackgroiuid with the .steeply dipping cuesta in the backg
Salix heathland and tussock tundra.

Jurassic Kingak Shale forms
round of Cretaceous sandstones.

and the cottonwood, Popiihis balsamifera. Betula
glundulosa and B. glandulifera fonn the iinder-
story in the riparian woods, along with Salix spp.,
Rosa, Ledum, Einpettim, and Rhododendron.
Herbaceou.s .streamside components include;
A(iroj)yron nuicrourum, Arahis hirsuta; Arenaria
j)lii/sodes. Astragalus alj)inus, Barbarea orthocer-
as, Castilleja pallida, Hedtjsarum alpinum, H.
boreale, Stellaria longipes, and Taraxacum cer-
atophorum.

M .\IUTIME VegEI ATION

The maritime vegetation is not essentially
different from that of the lower delta vegetative
types along the northern coast. Beach and spit
gravels and sands ( Fig. 44 and 45) support plant
communities dominated by Arenaria peploides,
Alopecurus alpinus, Cochlearia officinalis, Ehj-
mus mollis, Mertensia maritima, and Stellaria
humifusa. Landward from the beaches, spits, and
bars along the deltaic deposits, the same kinds
of plants occur which grow along streams in the
interior, e.g., with Salix alaxcnsis and other spec-
ies of willow dominating along the edge of
streams, and with Empetrum, Ledum, Rhododen-

dron, and other woody species of plants. Mea-
dows in the deltas are dominated by Eriophor-
um. Maritime vegetation gives way abruptly
along sea cliffs to tussock tundra typical of the
coastal plain.

Aquatic Vegetation

Streams in the region ordinarily have scoured
bottoms and do not support a flora of higher
plants. However, along the edge of some streams
the yellow marsh marigold, Caltha palustris,
grows in shallow water of quiet pools. Lakes
support a variety of kinds of af|uatic plants. In
Old Crow Flats some lakes are filled with yellow
pond lily, Nuphar pohjsepala. The pond lilies
grow in a circular pattern some distance out
from the edge of the pond in the deeper water.
The margin of these ponds is occupied by spec-
ies of Carex, Triglochin tnaritima, Utricularia
intermedia, Mentjanthes trifoliata, and Ranuncu-
lus hi/perboretis. Potamogeton alpinus is attached
and submersed in some ponds. Emergent vegeta-
tion in the lakes consists of Utricularia, Mentjan-
thes, and species of Carex. Several species of

Biological Sekies, Vol. 14, No.

Reconnaissance of Northern Yukon

27

Carex, Scirpus caespitosiis, and Pinguicula vul-
garis grow in moist sites in the heath-taiga lands
in Old Crow Fhits.

A pecuhar semiaquatic habitat occurs along
the Babbage and Canoe Rivers and Timber
Creek, where more or less permanent fields of

ice spread out along basins in the braided stream
course (Fig. 46). Willows and other woody
species are embedded in the ice. As the ice melts
the willows flower and grow leaves, but the
growing season is much shortened and existence
of this vegetative type appears tenuous.

^■•^-

X

*>

• it -J

Fic. 24. Patterned ground and poor Cretaceous exposures along the lower part of the Babbage River at appro.xi-
mately .58°.57' N; 139°2r W. Moderately wet tussock tundra on patterned ground (foreground) and stream-
bank and gravel bar vegetation (center and background).

28

Brigham Young University Science Bulletin
COLLECTION LOCALITIES

During the summer of 1970 collections were
made from 79 localities in northern Yukon ( Fig.
47). Collections were made in the period be-
tween June 17 and August 15.

1. NE Mount Page. In British Mts., ca 4 miles
northeast of Mount Page, in headwaters of
unnamed river draining to C>larence Lagoon,
ca 16 miles south of Beaufort Sea coast, at
69°24' N, 149°50' W, in alpine tundra on
Kayak shale.

2. Malcolm River. In pass between Malcolm
River and unnamed river draining to Clar-
ence Lagoon, ca 14 miles south of Beaufort
Sea coast, at 69°23' N, 140°40' W, at 2500
feet elevation, in alpine tundra on Kayak
shale.

3. Mt. Conybeare, south flank. South flank of
Mt. Conybeare, ca 8 miles south of Koma-
kuk Beach, Beaufort Sea, at 68°28' N, 140°
07' W, at ca 1100 feet elevation, in rich
tundra on limestone.

4. Mt. Conybeare, north flank. North flank of
Mt. Conybeare, ca 7 miles south of Koma-
kuk Beach, Beaufort Sea, at 68°29' N, 140°
07' W, at ca 1100 feet elevation, in rich
tundra on limestone.

5. Loney Creek. Along Loney Creek, a tribut-
ary of Firth River, 6 miles southwest of
confluence, at 69°20' N, 139°50' W, at ca
100 feet elevation, in arctic tundra, on Tri-
iissic liinestone of the Shublik formation.

6. Lower Firth River. Gravel bar and meadow,
along Firth River, ca 13 miles south of the
Beaufort Sea, and 14 miles southwest of
Herschel Island, at 69°22' N, I39°32' W, at
2(X) feet elevation, in willow dominated bar
and Dn/as-Oxijtwpis meadow.

7. Buckland Hills. North slope of Buckland
Hills, on ridge adjacent to a small lake, ca
H mile east of hill 722 and 20 miles south of
Herschel Island, at 69° 13' N, 139°06' W, at
700 feet elevation, in tundra on Neniopuk
slate.

8. Ancient Beach. In Buckland Hills, on an-
cient, polygonally figured, gravelly beach,
ca 2 miles north of Roland Creek and 23
miles south of Herschel Island, at 69° 11'
N, 139°07' W, at 9(X) feet elevation, in
sparse tundra.

9. DC-3 Wreckage. Site of DC-3 wreckage, on
divide between Spring River and Roland
Creek, at 69°07' N, 139° 10' W, at 1500 feet
elevation, in sparse but rich alpine tundra,
on Neruokpuk slates.

10. Three Pyramids section. British Mts., at 69°
02' N, 139°43' W, at 3000 feet elvation, on
Kayak Shale in alpine tundra.

11. Crow-Trail. Ridge top, between Crow and
Trail rivers, ca 24 miles southwest of Kay
Point, at 69°00' N, 138°46' W, at 1000 feet
elevation, in tundra on Neruokpuk slate.

12. Crow River. Gravel bar and braided stream
channel, Crow River, north slope of Buck-
land Hills, at 69°05' N, 138°40' W, at ca 350
feet elevation, in willow community.

13. Coastal plain lake. Lake margin, ca 4 miles
west-southwest of King Point, at 69°06' N,
138°07' W, at ca 200 feet elevation, in hum-
mocky heath-tundra.

14. King Point. Sandy beach, at King Point,
Beaufort Sea coast, at 69°07' N, 137°58' W,
at near sea level, in maritime vegetation.

15. Firth River. Gravel bar, in Firth River at
junction of Muskeg Creek, at 68°47' N, 140°
28' W, at ca 1400 feet elevation, with wil-
lows, dwarf l>irch, and white spruce.

16. Firth Lake. Lake shore and river gravel,
along Firth River, British Mts., at 68°49' N,
140°37' W, at 1.500 feet elevation, in Bettila-
SaJix heath.

17. Firth-Muskeg divide. Ridge top between
Firth River and Muskeg Creek, ca 12 miles
east of Alaska border and 5 miles southeast
of confluence of River and Creek, at 68°48'
N, 140°33' W, at 3200 feet elevation, in
rich tundra on Lisbume limestone.

18. Head Muskeg Creek. Ridge top, near head
of Muskeg Creek, ca 13 miles south of con-
fluence with Firth River, and 13 miles east
of Alaska border, at 68°38' N, 140°25' W,
at 2500 feet elevation, in poor tundra on
siliceous Triassic Shublik formation.

19. Lisbume cliffs. Snow flush, on north side
of small cliffs, in whitish Lisbume limestone
ca 40 miles west of Sam Lake, near head of
Timber Creek, at 68°32' N, 140°08' W, at
2800 feet elevation, in rich alpine tundra.

20. W Timber Creek. Ridge top, west of Tim-
ber Creek, ca 36 miles west of Sam Lake,
at 6S°26' N, 140°05' W, at 2800 feet eleva-
tion, in poor alpine tundra, on siliceous
conglomerate.

21. Ridge 1800. Ridge top #1800, on Lisbume
limestone, in northwestern portion of Old
Crow Flats, at 68°22' N, 140°40' W, at 1800
feet elevation, in taiga.

22. Bear Creek. Along Bear Creek, a tributary
of the Crow River, Buckland Mts., at 68°57'

Bioi.uGK^AL Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

29

Fig 25. View looking toward tlit- .suuthcast aciu.s.s llic piedmont zone between tJie coastal plain and the hills along
the Barn Mountains to the south. Stream in the foreground is at an elevation of approximately 50 feet. South
across the coiistal plain southwest of King Point from approximately 69°05' N; 138°07' W, near collecting
locality 13. W'et tussock tundra on polygonal ground, dominated by Eriophorum, Carex, and Salix species.

30

Bricham Young Univebsity Science Bulletin

N, 139°35' W, at 2(X)0 feet elevation, in al-
pine tundra.

23. Mt. Sedgwick. Northwest edge of Mt. Sedg-
wick, at 68°52' N, 139°2()' W, at 2500 feet
elevation, alpine tundra, in granite wa.sh
felsenmeer.

24. Trout Lake. North edge of Trout Lake, ca
.30 miles north of Sam Lake, in Babbage
River valley, at 68°50' N, i;38°45' W, at
KXM) feet elevation, in shrubby tundra.

25. Habbage River bend. Prominent bend in
Babbage River, at 68°57' N, 1.3S°20' W, at
ca 400 feet elevation, in arctic tundra.

26. Babbage River lower. Gravel bar in promin-
ent meander of Babbage River, on coastal
plain, at 6S°58' N, 1.38°2.5' W, at ca 400 feet
elevation.

27. Deep Creek. Stream gravel, along Deep
Creek, in shale slope, at 68°50' N, 137°45'
W, at 490 feet elevation, with willows.

28. Deep Creek lower. Along Deep Creek, at
68°52' N, 137°43' W, at 400 feet elevation,
in tundra over Cretaceous shale.

29. Blow River, Gravel bar, in prominent mean-
der of Blow River, ca 12 miles south of
Shingle Point, at 68°46' N, 137°20' W, at .300

feet elevation, in willow dominated com-
munity.

30. Blow River delta. Delta of Blow River, ca
2 miles east of Shingle Point, at 68°55' N,
137° 10' W, at ca 2 feet elevation, in Salix-
Eriophorum—Alopecurus community in open
gravelly areas between rows of driftwood
logs.

31. Upper Babbage River. Along Babbage
River, ca 32 miles west-northwest of Sam
Lake, at 68°38' N, 139°45' W, at 1200 feet
elvation, in willow thicket and meadow,
along black Kingak shale slope.

32. Babbage-Cottonwood. Hill top, between
head of Babbage River and Cottonwood
Creek, at 68°40' N, 139°27' W, at 1900 feet
elevation, in tundra on Shublik sandstone
outcrop.

33. Ridge 3261 north. South-facing slope, ca 3
miles north of #3261, ca 25 miles west-
northwest of Sam Lake, at 68°32' N, 139°24'
W, at 2000 feet elevation, on rocks stripes
of siliceous fonnation.

34. Ridge 3261 NNE. Low cliff, on summit of
ridge, ca 5 miles north-northeast of #3261,
ca 21 miles west-northwest of Sam Lake, at

.^^.

^

;^^^

I'lG. 26. Westward into Lisl)iiriu' Liiiustoiic outcrops in the licaiiw.ittrs ot Cr.utl Crcik at approxiniatcK 68°43'
N; 139°43' W. Peaks in the background have an elevation of approximate!)' 3500 feet and a rehef of 1000 to
1500 feet. Alpine tundra, on solifluction and talus slopes.

Biological Series, Vol. 14, \u. 2 Reconnaissance of Northern Yukon

31

^.

--%^.-

^^

'<,;*«**-,,; -^-

■vr'

. i ■

.••^..■\ ~ -.-, •*'-.>4;jt-' --'^^ •^■^^",-^•-

■:^';

■^:k.;«^

-'T..-^^^

^-■J* T *^>'»*^

Fig. 27. Northwest from near collecting locality 49 at approximately 68°2.> \. I39°07' W. Rocks in the fore-
ground are limestone of the older Paleozoic Neruokpuk sequence exposed in the core of a small dome south
of the main British Mountain trend. The darker slopes in the intermediate distance are on the Kayak Shale,
with Lisbume Limestone forming the gentle cuesta and the hill in the intermediate distance. Rich alpine
tundra.

68°32' N, 139° 18' W, at 2500 feet elevation
in rich tundra, on Permian Sadlerochit lime-
stone.

35. S Babbage River. South fork of Babbage
River, ca 18 miles northwest of Sam Lake,
at 68°35' N, 139°07' W, at ca 2000 feet ele-
vation, in tundra on Jurassic shales.

.36. Babbage ice field. Lower ice field, near
head of Babbage River, along bluff above
south bank, at 68°40' N, 139°07' W, at
1000 feet elevation, in mixed heath-tussock

tundra on Kingak shale.
37-38. Babbage River narrows. Narrows along
Babbage River, ca 20 miles north-northwest
of Sam Lake, at 68°43' N, 139°02' W, at ca
900 feet elevation, in tussock tundra along
ridge crest, on Triassic rocks.

39. Canoe River. Head of Canoe River, ca 12
miles northwest of Sam Lake, at 68°34' N,
138°5S' W, at ca 1700 feet elevation, in
alpine tundra on Cretaceous shales.

40. Dog Creek upper. Along upper Dog Creek,

32

Brigham Voung University Science Bulletin

41

42.

ca 9 miles north of Sam Lake, Barn Mts.,

at 68°32' N, 1.38°42' W, at ca 1800 feet ele-
vation, tussock tundra and stream gravels.
Bam Mts. Reddish, slaty ridge top, in Barn
Mts., ca 10 miles northeast of Sam Lake,
at 68°a3' N, 138° 18' W, at ca 2500 feet ele-
vation, in alpine tundra on Neruokpuk for-
mation.

Fitton Creek. Along west fork of Fitton
Creek, a tributary of Blow River, ca 22 miles
east-northeast of Sam Lake, at 68°34' N,
138°04' W, at 1000 feet elevation, in snow
flush along mossy slope, and in willow
thicket.

NE Mt. Fitton. Ridge top, ca 4 miles north-
east of Mt. Fitton, on summit between Fit-
ton Creek and Blow River, at 68°30' N,
1.37°52' W, at 1900 feet elevation, in sparse
alpine tundra, on Neruokpuk formation.
Ridge 3261 south. Ridge top, ca 2 miles
south of #3261, ca 21 miles west-northwest
of Sam Lake, at 68°28' N, 139°25' W, at
.3200 feet elevation, in alpine tundra, in rock
stripes of Ordovician-Silurian limestone.
45. Siliceous conglomerate. Outcrops of siliceous
conglomerate, ca 23 miles west of Sam Lake,

43.

44

at 68°25' N, 139°30' W, at 2500 feet eleva-
tion, in poor alpine timdra.

46. Ridge 3261, 4 miles S. Ridge crest and
saddle, ca 20 miles west of Sam Lake, at
2S°27' N, 139°22' W, at 2800 feet elevation,
in sparse alpine tundra, on slaty schist, in
Neruokpuk formation.

47. Triassic-Lisbume contact. West Bam Mts.,
at Triassic-Lisbume contact, at 68° 24' N,
1.39°20' W, at 2000 feet elevation, in alpine
tundra.

48. Ridge 2651. Ridge top, about 14 miles west
of Sam Lake, at 68°23' N, 139° 10' W, at
2651 feet elevation, in alpine tundra on
Ordovician-Silurian limestone.

49. Ridge 2651, 1 mile E. Ridge crest, ca 13
miles west of Sam Lake, ca 1 mile east
#2651, at 6S°23' N, 139°08' W, at 2500 feet
elevation, in rock stripes, in patchy but rich
tundra on Lisbume limestone.

50. Ridge 2651, 1 mile S. Ridge top, ca 14 miles
west of Sam Lake, at 68°22' N, 139° 10' W,
at 2500 feet elevation, in tundra-taiga tran-
sition at contact zone between Kayak shale
and Lisbume limestone.

51. Ridge 2651, 1.5 miles S. Ridge top, ca 14

Fig. 28. East across outcrops of massive Lisbume Limestone at locality 19 at approximately 2800 feet in the
.soutliem part of the British Mountains at 68°32' N; 140°08' W. Lakes in Old Crow Flat'can be seen in the
background to the southeast. Rich alpine tundra.

Biological Sehies, Vol. 14, No. 2 Reconnaissance of Northern Yukon

33

Fig. 29. Northweshvard acros.s Li.sburne Limestone outcrops at Locality 19 on the ruigr
British Mountains in the same areas as Figure 10. Alpine tundra, rich in species.

miles west of Sam Lake, ca 1.5 miles south
of #2651, at 68°21' N, 1.39° 10' W, at 2100
feet elevation in rock stripes and boulder
patches with poor alpine tundra, on siliceous
conglomerate and Kayak shale.
Small lake. Small lake in Old Crow Flats, ca
22 miles southwest of Sam Lake, at 68° 16'

N, 1.39°22' W, at ca 1000 feet elevation,
emergent vegetation and surromiding Be-
ttila-Rhododendron heath and spruce woods.
53. Caribou trap. Historic caribou trap (late
19th century?), ca 11 miles west of Sam
Lake, at 68°24' N, 239°04' W, at 2000 feet
elevation, in parkland tundra.

34

54. Dog Creek north. Along Dog Creek, ca 6
miles northwest of Sam Lake, at 68'>28' N,
138°45' W, at ca 2(XX) feet elevation, in a
wet meadow with Salix, Vaccinium, Arcto-
staphijlos. Drtjas, Carex, and Arnica.

55. Kayak sliale. Quartzitic member of Kayak
shale, ca 6 miles northwest of Sam Lake,
in Bam Mts., at 68°26' N, 138°48' W, at

Brigham Young Univer,sity Science Bulletin

2000 feet elevation, in poor alpine tundra.

56. Black Fox Creek. Cravel bar, in Black Fox
Creek, ca 5 miles west of Sam Lake, at
68°24' N, 138°49' W. at 1400 feet elevation,
with willows and tussock tundra.

57. Sedge meadow. Summit of ridge, in Lis-
burne limestone, south end of Bam Mts.,
ca 3 miles northwest of Sam Lake, at 68°26'

" T

\ 'Ma,

tf*'^

<"»

■■•^^^

«S»~— • J' ^ -^rv I. i>

Fic. 30. Kayak Shale and overlying Lishiirne Limestone along the west bank of the headwaters of Canoe River
above the elf icefield at approximately (>8°.'}7' N; 138 °43' \V" at an elevation of approximately 900 feet. Inhos-
pitable barren.s, heath, and patchv tundra on solifluction slope and talus.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

35

':^S5c/W

Fic. 31. B

31. Broad tundra-covered .slope blanketing strongly folded Nemokpuk beds near collecting locality 46 at
68°27' N; 139°22' W at approximately 2700 feet. Small platy angular debris is of siliceous schistose beds in the
Nemokpuk Formation. Sparse alpine tundra, with patches of Dnjas dominating.

N, 138°40' W, at 2500 feet elevation, in 60.
sedge meadow, alpine tundra.
58. Lisbume limestone. Rocky ridge top, ca 4
miles northwest of Sam Lake, at 68°27' N,
138°40' W, at 2578 feet elevation, in alpine 61.
tundra, on Lisbume limestone.
Dog Creek. Gravel bar, along Dog Creek,
ca 4 miles northwest of Sam Lake, at 68°26' 62.
N, 138°45' W, at ca 1400 feet elevation, in
willow community.

59

Spruce woods. Grove of spruce woods, on
south-facing slope, ca 3 miles northwest of
Sam Lake, at 68°26' N, 1,38° 40' W, at 1550
feet elevation, on Lisbume limestone.
NW Sam Lake. Wet, polygonal tundra, ca
1 mile northwest of Sam Lake, at 68"25' N,
138°.38' W, at ca 1600 feet elevation.
Sam Lake. Around margin of north side of
Sam Lake, at 68°25' N, 1,38°37' W, at 1500
feet elevation in Salix-Carex heath.

36

63.

64.

Sam Lake west. Around west margin of 65.
Sam Lake, at 68°25' N, 138°3S' W, at 1500
feet elevation, in Salix-Carex heath.
Dog Creek head. Along head of Dog Creek,
ca 6 miles north of Sam Lake, at 68°28' N, 66.
138°35' W, at 2200 feet elevation, in riparian
vegetation, heath, snow flush, and alpine
tundra, on sandstone. 67.

Brigham Young UNiYERsrrv Science Bulletin

Sandstone. Rock .stripes, in Cretaceous sand-
stone, ca 4 miles northeast of Sam Lake,
Bam Mt., at 68°27' N, 138°31' W, at ca
ca 2000 feet elevation, in poor alpine tundra.
W Barn Mts. Triassic-Lisburne contact, west
Bam Mts., at 68°24' N, 138 °20' W, at 2000
feet elevation, in tundra.
Timber C^reck. Gravel bar, along Timber

•*»...

*-*^-

\

.■5,.-'

*^.>-- *^-

;«il\

'>^<^aM%

32. Northwestward from Mt. Fittoii to Twin Peaks. The rocks on Mt. Fitton in the foreground are slightly
weathered siliceous granitic rocks. Slopes of Twin Peaks in the background are carved in large part in strongly
folded slates of the Neniokpuk se<iuence overlain hy coarse conglomerates of the late Paleozoic sequence.
The Twin Peaks rise to an elevation of approximatelv 2600 feet. Rocks of Mt. Fittoii in the foreground are
at an elevation of appro.ximately 1,5(K) feet. Alpine tundra, with boultler patches, solifluction slopes, aiul sparsely
covered slopes and rock outcrops.

Biological Series, Vol. 14, No. 2 Reconnaissance of Nohthehn Yukon

37

Fig. 33. Southwest from Mt. Fitton into the headwaters of Boulder Creek and the main high ridges of the Bam
Mountains. Rocks in the foreground are the Mt. Fitton granodiorite. and those in the background are strongly
folded Neniokpuk Formation. The high peak along the skyline in the distance is at approximately 3500 feet.
Mt. Fitton exposures in the foreground are at approximately 1500 feet. Alpine tundra.

68

69.

Creek, a tributary of Old Crow River, ca
3.3 miles southwest of Sam Lake, at 68°11'
N, 139 °47' W, at ca 900 feet elevation, in
white spruce-white birch woods and willow
dominated bar.

Lower Black Fox. Gravel bar, along Black
Fox Creek, at 68°07' N, 139°28' W, at ca
900 feet elevation, in spruce-willow-poplar
taiga.

Rose Lake. Gravelly north shore of Rose
Lake, Old Crow Flats, at 68°06' N, 139°23'
W, at 900 feet elevation.
70. Taiga pond. Pond and adjacent taiga-heath-
land, ca 22 miles south-southwest of Sam
Lake, at 68°08' N, 139°05' VV, at ca 1000
feet elevation.

Black Fox Creek lower. Along Black Fox
Creek, ca 13 miles southwest of Sam Lake,
at &Sn.5' N, 1.38 °50' W, at 1100 feet ele-
vation on gravel bar, with cottonwood and
willows.

Unnamed lake. Small unnamed lake, in
Old Crow Flats, ca 16 miles south of Sam
Lake, at 68°12' N, 138°49' W, at ca 1200
feet elevation, on humus beach and lake

73

71

72,

basin margin, with acjuatic plants and heath.
Ridge 1578. Small hill, #1578, in north-
eastern portion of Old Crow Flats, at 68° 12'
N, 138°45' W, at 1578 feet elevation, in
spruce taiga.
74. Neruokpuk limestone. West side of Blow
River Valley, at 68° 17' N, 137°50' W, at
2200 feet elevation, in alpine tundra, on
Neruokpuk limestone.

Cretaceous mudstone. Head of Blow River,
along a creek margin ca 27 miles southeast
of Sam Lake, at 68°17' N, 137°17' W, at
1500 feet elevation, in tundra.
Old Crow. Along gravel bar and first ter-
race above Porcupine River at Old Crow,
at 67°35' N, 139°50' W, at ca 900 feet ele-
vation, in white spruce woods and riparian
vegetation, on alluvium.
Old Crow landing strip. Landing strip on
river bar, ca 8 miles east of Old Crow, at
67°.34' N, 139°32 W, at 900 feet elevation,
in willow community.

Porcupine River bluff. Bluff west of bend
in Porcupine River, ca 31 miles east of Old

75

76

77.

78

38

Brigham Young UNrvEBSiTV Science Bulletin

Crow, at 67°36' N, 138°38' W, at 100 feet
elevation, in spruce woods.
79. Conglomerate hill. On siliceous conglom-
erate hill, ca .39 miles east of Old Crow, at
67°32' N, 138°23' VV, at 2000 feet clevaHon,
in heathlands and tundra.

Checklist of Vasculah Plants

The following list of plants collected during
1970 is arranged in phylogenetic secjuence to the
subdivision level and is alphabetically arranged
thereafter. Collectors names are abbreviated with
WR for S. L. Welsh and J. K. Rigby, and Rigby
for J. K. Rigby. All specimens are deposited at
BRY. A summary of taxa collected is presented
at the end of the checklist (Fig. 48). Distribu-
tion records were checked against maps pub-
lished by Hulten (1968) and Porsild (1964).
Identifications were made by comparison of
specimens collected with keys provided by Hul-
ten and Porsild and with an unpublished manu-
script of "Anderson's Flora of Alaska and ad-
jacent Canada" (Welsh, 1971).

LYCOPSIDA

Lycopodiaceae Clubmoss Family

Li/copo(Uutn selago L.

Locality #41, WR 10291, 8 July; #43, WR
10720, 15 July. Alpine tundra, ridgetops, on
slate.

Selaginellaceae Spikemoss Family

SelagineUa silnrica (Milde) Hieron

Locality #9, WR 10139a, 1 July; #41, WR
10297, 8 July. Alpine tundra, ridgetops, on slate.

SPHENOPSIDA

Equisetaceae Horsetail Family

Equi.setum arvense L.

Locality #15, WR 70578a, 12 July; #16,
Rigby 34, '23 June; #59, WR 10051, 30 June;
#77, Rigby 11, 18 June. Gravel bars, and form-
ing vast green patches along moist upland sites.

■ . ii:r--.'<'^ ■

f!5i'^-.,>-'r-^

I. '■^1^

:^'Mf:^^ -T^-^'^*

I'll. >l Si'iiili .nIl>^^ llic south (lank ot the British Moiiutaiiis into Old Oow Flats from approxiniatclv 68°24' .\;
1.39°16' W near locality 45. Rocks in the foreground are siliceous clierty conglomerate underlying the Kayak
Shale, the soft unit which forms the broad valley in the intermediate distance. The light colored ridges beyond
the valley and along the margin of Old Crow Flats are in Lisbunie Limestone. Old Oow I'lats to the south
is an area where broad marshes and lakes are extensively developed on Tertiary and Quaternary fill. Alpine
tundra dominated by Betulti glandulosa, Dri/as, Loiseleurui, and Etnpctmm.

Biological Series, Vol. 14, No. 2 Reconnais.sance of Nohthebn Yukon

39

.','V*^' /.

^

•>l*f

.^-..-r-.-

&■'
'-*'^.

-•ir^

:=r >.

,■•<«' ■

■-- /ivX-.-ir'v

vA-.-i-

• r .V .''..• '^;,-— ^^ <>r. ■•■^.«S' .,-—•-'.■ .U;v-}-; -, ;'•'.■,

-'■'■!^W ■■'*-

Fig. 35. Siliceous iron-rich .silt.stoncs and shale in the Permian Sadlerochit Formation as seen from the head of
Mu.skeg Creek at 68°.31' N; 140°.32' W near collecting locality 18, looking toward the north to the main part
of the IJritish Mountains. The hroad open flats in the background are at an elevation of 2000 feet. Rocks in
the foreground are at an elevation of 2500-2600 feet. Sparse alpine tundra, dominated by Dnjas, Beiula, and
Empetrum.

Equisetum scirpoides Michx.

Locality it 26, WR 10699, 15 July. Gravel
l>ar.

PTEROPSIDA-FILICINEAE

Polypodiaceae Fern Fiunily
Cijstopterls jragilis (L. ) Bemh.

105,50, 12 July; #32, WR 10209, 3 July. On low
cliffs and talus slopes, in alpine tundra, on
sandstone and limestone. This is the first re-
port of C. jragilis for northern Yukon.

Dnjopteris jragrans (L). Schott.

Locality #33, WR 10.389, 9 July. Siliceous
conglomerate, in alpine tundra. ThLs is the first

Locality #17, WR 10557a, 12 July; #19, WR record of D. fragram for northern Yukon.

40

Bricham Young University Science Bulletin

PTEROPSIDA-GYMNOSPERMAE

Pinaceae Pine Family

Picea glauca ( Moench ) Voss

Locality #16, Rigby 25, 23 June; #60, WR
10046, 30 June; #67, WR 10096, 1 July; #76,
WR 10432, 10 July. On terraces, gravel bars, lake
shores, and mountain slopes, in taiga and boreal
forest. A dominant species.

Picea mariana (Mill.) Britt, Stems., Pogg.

Locality #50, WR 10509, 11 July. Tundra-
taiga transition, an Kayak shale. This specimen
has only a few hairs along the leaf base and
lacks cones. It is tentatively assigned to P. nmri-
ami, but might represent a hybrid with P. glauca.

PTEROPSIDA-ANGIOSPERMAE-DICOLYLEDONEAE

Betulace;\e Birch Family

Alnus crispa ( Ait. ) Pursh

var. cris^m

Locality #30, WR 10323, 8 July, #.36, WR
10194, 3 July; #72, 10654, 13 July. Stream bank
and heathland component, widespread.

Betula glandulifera ( Reg. ) Buder

Lociility #67, WR 10097, 1 July; #79, Rig-
by 130b, 26 July. These are shrubs with ch;u--
acteristias intennediate between B. glwuhtlosa
and B. j)a])t/rifera. They grow along river banks
and bluffs iii Old Crow Flat.

Betula glandulosa Michx.

var. glandulosa

Locality #8, WR 10147, 1 Julv; #16, Rigby
27, 40, 23 June; #24, WR 10126', 1 July; #52,
WR 10479, 11 July; #65, WR 10301, 8 July;
#73, Rigby 54, 24 June. Dwarf birch is abundant
in most ph;xses of alpine and arctic tundra and
heathlands. In favorable locations it is erect,
reaching a height of 2 to 4 feet. In less favorable
sites, it is a prostrate spreading shrub.

Betula papyrifera Marsh

var. neoala.skaim ( Sarg. ) Raup

Locality #67, WR 10095, 1 July; #79, Rig-
by 130, 134, 26 July. White birch Ls a component
of riparian woods and occurs on bluffs along
stream channels in the lower reaches of Old
Crow Flat.

Fig. 36. - '■!. , '- !■ ■ nil^ iiMi,. v-. J l.- ,i- ^ mi I iL.- .^,<u^^ \\,r nnrth si.lr ot Old Crnw Flats.

The woods are at ap|)roximatcly ti8°2.5' N; 138°4()' VV. (^ollfcting locality W) is in the point of woods in the
immediate right center, at the edge of the tundra, at an ele\ation of approximately 1550 feet. Mixed heath-
land and tussock tundra are in the foreground.

Biological Sehies, Vol. 14, No. 2 Reconnaissance of Northern Yukon

41

Fig. .37. East up Skidoo Creek at 68°.51' N; 140°13' W, east of c-oUecting locality 15. AH the rocks immediately
visible in the foreground are in the Lisbume Limestone. The valley bottom in the foreground is at an eleva-
tion of appro.ximately 1500 feet, with peaks along the skyhne at about 2000 feet. White spruce woods, near
northern limit of trees along Firth River valley.

42

Bhicham Young University Science Bulletin

Boraginaceae Borage Family

Eretrichium nanum (Vill. ) Schrad.

var. aretioides (Cham.) Herder

Locality #1, WR 10262, 5 July; #2, WR
10247, 5 July; #48, WR 10518, 11 July. Alpine
tundra and ])arTen.s.

var. chami.ssonis ( DC. ) Herder

Lociility #3, WR 10176, 1 July; #4, WR
10226, 4 July; #23, Rigby 96, 2.5 June. Alpine
tundra and barreas.

Mertensia maritima ( L. ) S. F. Gray

Loaility #14, WR 10033, 30 June; do, WR
10686, 15 July. Gravelly beaches and spits along
the sea coast.

Mertensia paniculata ( Ait. ) G. Don

ssp. paniculata

Locality #5, WR 10603, 13 July; #31, WR
10405, 9 July; #54, WR 10276, 6 July; #59,
10062, 30 June. Gravel bars, meadows, and al-
pine and arctic tundra.

Myosotii- stjlvatica Hoffm.

Locality #3, WR 10178, 1 July; #5, WR
10610, 13 July; #25, WR 10624, 13 July; #27,

Rigby 141, 28 July; #40, WR 10506, 11 July;
#59, WR 10065, 30 June. Streams gravels, tus-
sock tundra, alpine tundra, and barreas.

Caryophyllace;ie Fink Family

Arenaria arctica Stev

Loaility #1, WR 10261, 5 July; #3 WR
10180, 1 July; #4, WR 10218, 3 July; #8, WR
10152, 1 July; #9, WR 101.33, 1 July; #16, Rig-
by 71, 24 June; #17, WR ia558, 12 July; #32,
WR 10211, 3 July; #42, WR 10293a, 8 July;
#43, WR 10714, 15 July; #59, WR 10059, 30
June. Ridge tops, alpine slopes, tussock tundra,
and stream gravels, common to abundant.

Arenaria kiricifolia L.

var. htiltenii Welsh

Locality #46, WR 10378, 9 July; #55, WR
10068a, 30 June; #79, Rigby 133, 26 July. Sili-
ceous outcrops iuid on limestone, in alpine tun-
dra.

var. laricifolia

Locality #29, WR 10355, 9 July; #45, WR
10539, 12 Julv; #.50, WR 10.5^, 11 July; #51,
WR 10514, 11 July; #51, WR 10.362, 9 July;
#66, Rigby 107, 22 July. Siliceous conglomerates

Fig. 38. CJravel bar at Kirtli Rock and tollecting locality 1.5 in the foreground. Most of the plants were collected
to the right beyond the picture on the frciniently covered crest of the gravel bar. White spruce, willow, alder,
and dwarf birch dominate the talus slopes.

Biological Sebies, Vol. 14, No, 2 Rec;onnaissance of Northern Yukon

43

and quartzites and less commonly on limestones,
in alpine tundra.

Arenaria macrocarpa Pursh

Locality #29, WR 10744, 15 July (from a
rounded, hemispheric clump 3 feet in diameter);
#40, WR 10510, 11 July; #41, WR 10293, 8
July; #43, WR 10716, 15 July. Stream gravels
and slaty ridge tops.

Arenaria peploides L.

var. peploides

Locality #14, WR 10032, 30 June; do, WR
10681, 15 July. Sea beach and spit gravels.

Arenaria phijsodes Fisch.

Locality #67, WR 10110,
gravels and bars.

1 July. Stream

Arenaria rossii R. Br.

var. elegans ( C. & S. ) Welsh

Locality #34, WR 10397, 9 July; #46, WR
10369, 9 July. Alpine tundra, on limestone.

Arenaria rubella ( Wahl. ) Smith

fma. exilis ( Fern. ) Polunin

Locality #9, WR 10141, 1 July. Alpine tun-
dra on Neruokpuk fomiation.

'9

'^.

H-

.A

N -^

S- -^n^

4i vi"^

■^'^■:

i:.

Fig. 39. Northwest along Pleistocene beach ridges. The prominent light scar in left center is a graveled beach at
appro.ximately 68°12' N; 139°02' W at the eastern shore of a broad lake which occupied Old Crow Flats. Black
Fox Creek in the middle distance is lined by spruce. British Mountains rise along the skyline beyond Old Crow
Flats. Parkland tundra (taiga) and heathland.

44

Bhicham Young University Science Bulletin

var. rubella

Locality #11, WR 10187, 1 July; #59, WR
10050, .30 June; do, WR 10465, 10 July. Gravel
bars, and alpine tundra on Neruokpuk formation.

Arenaria sajanensis Willd.

Locality #64, WR 10642, 14 July. Shrubby,
alpine tundra, in a snow flush.

Cerastium beeringianum Cham. & Schlecht.

var. beeringiannm

Locality # 12, WR 1045.3, 10 July; #44, WH
10367, 9 July; #.59, WR 100.53. Alpine tundra
on limestone and on gravel bars and braided
stream channels.

Cerastium maximum L.

Locality #76, WR 10434, 10 July. River ter-
race alluvium in white spruce woods.

Lijchnis apetala L.

Locality #44, WR 10365, 9 July; #49, WR
10356, 9 July. Alpine tundra, on limestone.

Lychnis jurcata ( Raf . ) Feni

Locality #34, WR 10391, 9 July. Alpine tun-
dra, on limestone.

Lychnis triflora R. Br.

var. dawsonii Robins.

Locality #67, WR 10113, 1 July. Gravel bar,
in lower Old Crow Flats.

Silene acaulis L.

var. exscapa (All.) DC.

Locality #2, WR 10257, 5 July; #4 WR
10217, 3 July; do, WR 10241, 4 July; #5, WR
10615, 13 July (peduncles filifonn, to 1.5 cm
long); #6, WR 10169, 1 July; #37, WR 10092,
30 June. Alpine and arctic tundra, on ridge tops,
slopes, and gravel bars, common to abundant.

Silene repens Pers.

Locality #31, WR 10401, 9 July; #66, Rigby
109, 22 July. Gravel bars and tussock tundra.

Stellaria huniifusa Rottb.

Locality #14, WR 10682, 15 July. Sea beach
and spit gravels.

Stellaria longijolia Muhl.

Locality #70, WR 10662, 15 July. Pond mar-
gin, in taiga.

Stellaria longipes Goldie

var. altocaulis (Ilulten) C.L. Ilitchc.

Locality #12, WR 10452, 10 July. Gravel
bars, in braided arctic stream channel.

var. edwardsii (R. Rr. ) Gray

Locality #56, WR 10590, 12 July. Gravel
bar, in stream channel.

var. laeta ( Richards. ) Wats.

Locality #14, WR 10674, 15 July; #50, WR
10530, 11 July; #59, WR 10060, 30 June. Sea
beach and spit, alpine tundra, and gravel bars.

var. longipes

Locality #5, WR 10612, 13 July; #29, WR
107.39, 15 July; #67, Rigby 58, 24 June; do, WR
10104, 1 July.' Gravel bars.

Compositae Composite Family

Achillea millejolium L.

ssp. boreale ( Bong. ) Breitung
Localitv #26, WR^^ 10706, 15 July; #29, WR
10736, 15 July; #68, Rigby 1.59, 20 Julv; #76,
WR 10420, 10 July; #78, Rigby 127, 26 July.
River bluffs, terraces, and gravel bars.

Antennaria alpina ( L. ) Gaertn.

var. compactu (Malte) Welsh

Locality #11, WR 10192, 1 July; #1S, WR
10554, 12 July. Alpine tundra, in siliceous sand-
stones and slaty shales.

Antennaria monocephala DC.

Locality #8, WR 10151, 1 Julv; #9, WR
10146a, 1 July; #40, WR 10641, 14 July. Alpine
tundra and heathlands.

Arnica alpina ( L. ) Olin

ViiT. angustifolia (Vahl) Feni.

Locality #1, WR 10263a, 5 July; #4, WR
102.32, 4 July; #5, WR 10601, 13 July; #50, WR
105.32, 11 July. Alpine and arctic tundra.

ssp. attenuata (Greene) Maguire

var. attenuata

Locality #12, WR 10447, 10 Julv; 3i.3.3, WR
10.379, 9 July; #25, WR 10621, 1.3 July; =54.
WR 10273, 6 July. Stream banks, meadows, and
bars.

Arnica lousiana Farr.

var. frigida ( C. A. Mev. ) Welsh

Locality #1, WR 10263, 5 July; #15, WR
10.563, 12 July; #.32, WR 10205, 3 July. Alpine
tundra.

Artemisia alaskana Rydb.

Locality #26, WR 107{X), 15 July. Gravel
bar.

Artemisia glomerata Ledeb.

Locality #9, WR 10138, 1 July; #10, Rigby
75, 91, 25 June. Alpine tundra, on Neruokpuk
formation and Kavak shale.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

45

Artemisia norvegica Fries

var. coiiuita ( Rydb. ) Welsh

Locality #12, WR 10446, 10 July; #43, WR
10725, 15 July; #64, WR 10639, 14 'July. Gravel
bars, and alpine tundra.

Artemisia tilesii Ledeb.

var. iinalaschcensis Besser

Locality #14, WR 10675, 15 July; #26, WR
10701, 15 July; #29, WR 10739, 15 July; #59,
WR 10468, 10 July; #67, WR 10106, 1 July;
#68, Rigby 157, 20 July; #78, Rigby 126, 26
July. Gravelly sea beaches and spits, and on
gravel bars, terraces, and river blufls.

Aster sihiricus L.

Locality #68, Rigby 156, 20 July; #76, WR
10426, 10 July; #78, Rigby 125, 26 July. River
bars, terraces, and bluffs, in Old Crow Flats.

Crepis nana Richards.

Locality #12, WR 10444, 10 July; #15, WR
10570a, 12 July; i±16, Rigby 69, 24 June; #19,
WR 10552, 12 July; #25^ WR 10622a, 13 July;
#49, WR 10352, '9 July; #59, WR 10462, 10
July; #67, WR 10105, 1 July. Bars and stream
banks and on talus and ridge tops in tundra.

Erigeron acris L.

var. kamtscliaticus (DC.) Herder

Locality #68, Rigby 1.54, 20 July; #71, WR

10496, 11 July. Gravel bars, in Old Crow Flats.

Erigeron hiimilus Grab.

Locality #50, WR 10536, 11 July; #51, WR
10515a, 11 July. Kayak shale, in alpine tundra.

Erigeron lonchophijlhts Hook.

Locality #22, WR 0713, 15 July. Tussock
tundra.

Erigeron purpuratus Greene

Locality #59, WR 10459, 10 July; #71, WR
10500, 1 July. Gravel bars, along streams in Old
Crow Flats.

Erigeron ijukonensis Rydb.

Locality #.34, WR 10390, 9 July; #50, WR
10527, 11 July; #51, WR 10508, 11 July; #66,
Rigby 106, 22 July. On Kayak shale, and contact
between Shublik and Lisbume limestones in al-
pine timdra. The specimens do not fit the pub-
lished description of E. ijukonensis exactly. In
fact, they closely resemble Aster alpinus var.
vierliappeii, but have bright pink purple rays.

|1i*«^'W*''

p-.-

F'lc. 40. Southwcstward across lakes along the west bank of Old Crow River, toward Shaffer Creek drainage.
Lakes at the right arc at 67°55'N; 139°44' W. The relatively well-drained bank of Old Crow River is to-
ward the east (Ifft). Elevation is approximately 950 feet. Riparian woods of Picia glauca, Salix, iind Poptilus,
and heathlands of Betula glandulosa and SuUx.

46

Brioham Young Univebsity Science Bulletin

Fio. 41. Parkland tundra near the uppermost limit of trees along Timber Creek at approximately 68°26' N;
139°45' W. Abandoned meanders show along the gravel margin. E.xposures of Lisbume Limestone .show as
the light colored barren area in the upper left. Salix and Picea glauca fringed stream and meander.

Matricaria matricarioides (Les. ) Porter

Locality #76, WR 10435, 10 July. Weed of
disturbed sites, at Old Crow.

Petasites frigidus ( L. ) Fries

var. frigidus

Locality #24, WR 10124a, 1 July; #42, WR
10332, 8 July; #60, WR 10036, 30 June. Mea-
dows, willow thickets, and heathlands.

Saussurea angustifolia ( Willd. ) DC

var. angustifolia

Locality #33, WR 10386, 9 July; #51, WR,
10515, 11 July; #56, WR 10591, 12 July. Siliceous
conglomerates in alpine tundra, heathlands, and
on gravel bars.

Senecio atropurpureu.s (Ledeb. ) Fcdtsch.

VAT. ulriKri (Steffen) Porsild

Locality #4, WR 10239, 4 July; #7, WR
10159, 1 July; #16, Rigbv 68, 24 June; #34, WR
10204, 3 July; #40, WR 10644, 14 July; #57,
WR 10078, .30 June; #60, WR 10044, .30 June.
A dominant species in tundra generally. Both
ligulate and nonligulate phases are represented.

Senecio congestus ( R. Hr. ) DC.

Locality #14, WR 10673, 15 July; #62, WR

10549a, 13 July; #72, WR 10649, 14 July. Gra-
velly seashores and spits, and lakes shores.

Senecio fuscatus ( Jord. & Fourr. ) Hayek

Locality #9, WR 10142, 1 July; #43, WR
10721, 15 July. Alpine sparse tundra, on Ner-
uokpuk formation.

Senecio lugens Hook

Locality #5, WR 10605, 13 July; #25, WR
10622, 13 Julv; #27, Rigbv 143, 28 July; #29,
WR 10737, 15 July; #50, 'WR 10529, 11 July;
#.54, WR 10284, 6 July; #.59, WR 10473, 10
July; #67, WR 10108, 1 July; #71, WR 10495,
1 1 July. Gravel bars and meadows.

Senecio resed if alius Less.

Locality #1, WR 10264, 5 July; #9, WR
10143, 1 July; #44, WR 10364, 9 July; #48, WR
10.348, 9 July; #74, Rigby 1.37, 26 July. Alpine
tundra, on limestone, slates, and shales.

Senecio ijukonensls Porsild

Locality #40, WR 10644a, 14 July. Shrubby
tundra.

Solidago multiradiatu Ait.

Locality #5, WR 10606, 13 July; #15, WR
10.570, 12 July, (iravel bars and stream banks.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

47

Tanacetum bipinnatum (L. ) Schulz-bip

ssp. huroncnse (Nutt. ) Welsh

Locality #76, WR 10424, 10 July; #78, Rig-
by 12.3, 26 July. Gravel bars and river terraces,
in Old Crow Flats.

Taraxacum ce rat ophorum (Ledeb. ) DC.

Locality #59, WR 10458, 10 July; #67, WR
10111, 1 July. Gravel bars and river terraces.

Taraxacum hjratum (Ledeb.) DC.

Locality #2, WR 10245, 5 July; #64, WR
10640, 14 July; #58, WR 10476, 11 July. Alpine
tundra, on Lisbume limestone, Kavak shale, and
Jurassic shales.

Crassulaceae Stonecrop Family

Scdum rosextm (L. ) Scop.

var. integrijolium (Raf. ) Berger

Locality #4, WR 10215, 3 July; #10, Rigby
81, 25 June. Alpine tundra.

Cruciferae Mustard Family

Ahjssum americanum Greene

Locality #48, WR 10344, 9 Jidy; #50, WR
10526a, 11 July; #51, 10513, 11 July. Alpine tun-
dra, on limestone outcrops. These collections
represent a northward extension of the range of
the species in the Yukon.

Arabis hirsuta Lange

var. pijcnocarpa ( Hopkins ) Rollins
Locality #67, WR 10101, 1 July; do, Rigby
60, 24 June. River bar. These collections repre-
sent the northernmost known limits of this entity
in the Yukon.

Barbarea orthoccras Ledeb.

Locality #67, WR 10102, 1 July; #69, Rigby
150, 20 July. Lake shores and gravel bars, in Old
Crow Flats.

Braija Immilus (C.A. May.) Robins

ssp. arctica ( Bucher. ) Rollins

Locality #15, WR 10565, 12 July. Gravel bar,
in Firth River. This specimen is apparently the
first record of B. humihts for northern Yukon.

Braija purpurascens (R.Br.) Bunge

Locality #50, WR 10345, 9 July. Alpine tun-
dra, on limestone. This specimen is apparently
the first record of B. purpurascens for the Yukon.

Cardamine bellidifoUa L.

Locality #42, WR 10322a, 8 July. Snow flush
and willow thicket.

Cardamine digitata Richards.

Locality #11, WR 10191, 1 July; #13, WR
10692, 15 July; #14, WR 10687, 15 July; #34,
WR 10202, 3 July; #57, WR 10076, 30 June;
#64, WR 10641a,' 14 July. Tussock and shrubby
tundra.

Cardamine microphijUu Adams

Locality #1, WR 10260, 5 July; #2, WR
10249, 5 July. Alpine tundra, on Kayak shale.
This is apparently a new record for the Yukon.

Cardamine pratensis L.

Locality #62, WR 10223, 4 July. Lake shore.

Cochearia officinalis L.

var. arctica ( Schlecht. ) Gel.

Locality #14, WR 10688, 15 July. Sea
beaches and spits.

Descurainia sophioides ( Fisch. ) Schulz

Locality #14, WR 10670, 15 July; #76, WR
10425, 10 July. Gravel bars and terraces along
rivers, and near an abandoned dwelling along
the sea coast.

Draba alpitia L.

Locality #16, Rigby 72, 24 June. Gravelly
river bank.

Draba caesia Adams

Locality #3, WR 10182, 1 July; #4, WR
10216, 3 July; #6, WR 10166a, 1 July; #9, WR
101.35, 1 July; #10, Rigby 79, 25 Jmie; #48, WR
10.341, 9 July. Alpine tundra, in limestone, shales,
and slates, and on gravel bars. Common.

Draba cinerea Adams

Locality #5, WR 10618a, 13 July; #21, Rig-
by 115, 25 July; #34, WR 10395, 9 July; #73,
Rigby 49, 24 June. Alpine tundra, on limestone
and sandstone.

Draba glabella Pursh

Locality #10, Rigby 77, 25 June; #14, WR
10685, 15 July; #31, WR 10408, 9 July; #59,
WR 10061, .30 June; #64, WR 10643, 14 July.
Gravel bars along streams, and sea beaches and
spits.

Draba lactea Adams

Locality #2, WR 0254, 5 July; #4, WR
102.38, 4 July; #17, WR 10562, 12 July; #19,
WR 10544, 12 July; #32, WR 10210, 3 July;
#44, WR 10363, 9 July; #64, WR 10636, 14
July. Sandstone, limestone, shales, and slates, in
alpine tundra. The specimens cited herein are
evidently the first records of this species for
northern Yukon.

48

Bhicham Young University Science Bulletin

Draba longipes Raup

Locality #50, WR 105.38, 11 July. In tiindra-
taiga transition at contact between Kayak shale
and Lisbume limestone.

Erysimum chieranthoides L.

Locality #68, Rigby 1.58, 20 July; #78, Rig-
by 129, 26 July. Gravel bars and stream terraces,
in Old Crow Flats.

Erysimum inconspicuum (Wats.) MacM.

Locality #25, WR 10623, 1.3 July; #26, WR
10698, 15 July. Gravel bars, along arctic streams.
These specimens are apparently the first record
of this species for northern Yukon.

Eryismum paUasii (Pursh) Fern.

Locality #10, Rigby 74, 25 June; #12, WR
10443, 10 June; #31, WR 10402, 9 July; #50,
WR 10353, 9 July; #59, WR 10049, 30 June; do,
WR 10464, 10 July. Gravel bars, and alpine
tundra.

Lesquerella arctica ( Wormskj. ) Wats.

Locality #16, Rigby 64, 24 June; #37, WR
10081, 30 June. Alpine tundra, on Triassic rocks,
and on gravelly river bank.

Parrya nudicaulis ( L. ) Reg.

Locality #.3, WR 10171, 1 July; #4, WR
10233, 4 July; #10, Rigby 80, 25 June; #1.3, WR
10693a, 15 July; #16, Rigby 21, 30, 35, 23 June;
do, Rigby 66, 24 June; #.37, WR 10090a, 30
June. A dominant species in tussock and alpine
timdra. Common.

Rorippa islatulica (Oed. ) Borbas

var. his}>ida (Desv. ) Butters & Abbe
Locality #70, WR 10663, 15 July; #72, WR
106.5.5, 14 July; #76, WR 10430, 10 July. Lake
shores, gravel bars, and river terraces.

Smelowskia borealis ( Greene ) Drury & Rollins

Locality #17, WR 10556a, 12 July. Talus
slope, on hmestone. This specimen represents a
range extension eastward and northward from
previous records.

Smelowskia cahjcina ( Steph. ) C. A. Mey.

var. media Drury & Rollins

LocaUty #2, WR 10243, 5 July; #4, WR
10213, 3 July; #6, WR 10167, 1 July; #7, WR
10154, 1 July; #8, WR 10149, 1 July; #9, WR
10140, 1 July; #10, Rigby 90, 25 June; #16,
Rigby 62, 24 June; #24, WR 10125, IJuly; #41,

Fio. 42. Black Fox Creek at the margin of Old Crow I'lat looking toward the southwest from approximately
68°15' N; 138°76' \V. Picca glmica woods, margined with Salix. Bctulti papt/rifcra. and Populus.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

49

WR 10292, S Julv: #4.3, WR 10722, 15 July;
#46, WR 105S2a,' 12 July. A dominant on senii-
barren ridgetops and abandoned beaches, and
less commonlv in stream gravels. This is evidently
the first report of S. cahicina from the Yukon.

Elaeagnaceae Oleaster Family

Shephcrdia canadensis (L. ) Nutt.

Locality #59, WR 10052, .30 June; #68,
Rigby 100, 20 July. Gravel bars, along stream
courses.

Empetraceae Crowberry Family

Empetrum nigrum L.

var. Iiennaphroditicum (Lge. ) Sor.

Locality #36, WR 10199, 3 July; #65, WR
10300, S July. A dominant in tussock tundra and
heathlands. This is evidently the first report of
£. nigrutn for northern Yukon.

Rhododendron Iapj)onictiin (L. ) Wahl

Locality #5, WR 10604, 13 July; #6, WR
10166, 1 July; #16, Rigby 15, 21, 29, 23 June;
#37, WR 10083, .30 June; #60, WR 10042, 30
June; #73, Rigby 52, 24 June. Tundra, heath-
land, and taiga, widespread and common.

Vaccinium uliginosum L.

Locality #7, WR 10160, 1 July; #36, WR
10201, 3 July; #62, WR 10024, 30 June. A dom-
inant species in tussock tundra, heathlands, and
taiga.

Vaccinium vitis-idaea L.

Locality #63, WR 10306, 8 July. Shrubby
tundra and taiga.

Gentianaceae Gentian Family

Gentianella propinqua ( Richards. ) Gillette

Locality #26, WR 10704, 15 July. Gravel bar.
Evidently rare.

Ericaceae Heath Family

Andromeda polifolia L.

Locality #60, WR 10045, 30 June; #62, WR
10031a, 30 June. Lake shores, heathlands, and
taiga.

Arctostaphylos alpina ( L. ) Spreng

Localitv #5, WR 10618, 13 July; #7, WR
10161, 1 July; #15, WR 10564, 12 July; #25,
WR 10628a, 13 July; #73, Rigby 47, 24 June.
A dominant species in tussock tundra and on
ridges and slopes in alpine timdra, and along
stream courses.

Cassiope tetragona (L. ) D. Don

var. tetragona

Locality #1, WR 10269, 5 July; #10, Rigby
82, 2.5 June; #16, Rigby 24, 23 June; #42, WR
10331, 8 July; #55, WR 10070, .30 June; #60,
WR 10041, 30 June; #73, Rigby 53, 24 June. Al-
pine tundra, heathland, and taiga. A dominant
common species.

Ledum decumbens ( Ait. ) Lodd.

Locality #30, WR 10319, 8 July; #55, WR
10071, 30 June; #60, WR 10044b, 30 June; #73,
Rigby 57, 24 June. Woodlands, lake shores,
stream banks, and deltas.

Loiseleuria procumbent (L. ) Desv.

Locality #.39, WR 10129, 1 July; #47. WR
10359, 9 July; #73, Rigby 45, 24 June. Poor tun-
dra on siliceous conglomerates, sandstone, and
shale.

Haloragaceae Watermilfoil Family

Hippurus vulgaris L.

Locality #62, WR 10313, 8 July. Emergent
in lake margin.

Leguminosae Legume Family

Astragalus alpinus L.

Locality #12, WR 10449a, 10 July; #16,
Rigby 14, 23 June; #29, WR 10732a, 15 July;
#36, WR 10203, 3 July; #59, WR 10461, 10
July; do, WR 10047, 30 June; #67, Rigby 61, 24
June. Tussock tundra and gravel bars. Wide-
spread and locally common.

Astragalus ausiralis ( L. ) Lam.

Locality #3, WR 10175, 1 July; #12, WR
10440, 10 July; #15, WR 10580, 12 July; #32,
WR 10207, 3 July; #.37, WR 10080, 30 June.
Ridge tops, on limestone or sandstone, and on
river gravels on the north slope.

Astragalus bodinii Sheld.

Locality #59, WR 10047a, 30 June; do, WR
460, 10 July. Gravel bar, along Dog Greek. This
report is apparently the first record of A. bodinii
from Northern Yukon.

Astragalus umbellatus Bunge

Locality #4, WR 102.34, 4 July; #37, WR
10091, .30 June; #59, WR 10057, .30 June. Alpine
tundra and stream gravels.

Hedijsarum alpinum L.

Locality #1, WR 10270, 5 July; #5, WR

50

Bhicham Young Univehsity Science Bulletin

10613, 13 July; #34, WR 10388; #48, WR 10520,
11 July; #51, WR 10516, 11 July; #54, WR
10277, 6 July; #67, Righy 59, 24 June; do, WR
10116, 1 July; #76, WR 10418, 10 July; #78,
Rigby 128, 26 July. Alpine tundra, on shale and
limestone, and on gravel bars, stream banks and
terraces, and meadows. Widespread and com-
mon. Both the dwarf alpine phase and the tall
woodland phases are represented and also the
intennediates between them.

Hedtjsarum horeale Nutt.

.ssp. nuickenzii ( Richiu-ds. ) Welsh
Locality #1, WR 1027a, 5 July; #12, WR
10439, 10 July; #18, Rigby 120, 25 July; #27,
WR 10712, 15 July; #32, WR 10207, 3 July;
#37, WR 10082, 30 June; #48, WR 10346, 9
July; #50, WR 1052.3, 11 July; #51, WR 10507,
11 July; #59, WR 10048, .30 June; do, WR 10467,
10 July; #67, WR 10099, 10115, 1 July. Alpine
tundra on shale, limestone, and sandstone, and

':(-

'■)i^

)f^i

1

» '^

e:-jaruz\j:<'

Fic. 43. Southwest to the tri-i'-fiin:,! u I ili I l-l.n L : . i ii .t i. loss tlu- tiiii.!!.. iiiui „; jiioxiinatcK 68"I6' N;
138°52' W. Trail in tlic foreground is a winter seismograpli trail. Old Crow Mountain shows vcr\' faintly
along the skyline near the left hordt^r. Black Ko.x Creek in the foreground is at an elevation of 1100 to 1200
feet. Parkland tundra and heathland (foreground) and riparian w<K)ds in background.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

51

on gravel bars and terraces. Widespread and
locally common. There is a dwarf alpine phase in
this species also, and it is likewise connected by
a series of intennediates to the more robust
streamside phase.

Litpinus arctictis Wats.

Locality #2, WR 10256, 5 Julv; #8, WR
10148, 1 July; #11, WR 10184, 1 July; #12,
WR 10445, 10 July; #15, WR 10693, 15 Julv;
#16, Rigbv 43, 23 June; #24, WR 10119, 1 July;
#29, WR' 10740, 15 July; #59, WR 10063, .30
June; #59, WR 10066, 30 June; #62, WR 10026,
30 June; #65, WR 10303, 8 July; #77, WR 6,
18 June. Alpine tundra and heathlands on sand-
stone, shale, and limestone, and on stream gra-
vels and tussock tundra. Widespread.

Oxt/troj)is arctica R. Br.

Locality #12, WR 10448, 8 July; #35, WR
10221, 3 July; #48, WR 10340, 9 July. The latter
two specimens are more or less depauperate al-
pine phases from limestone ridge tops, the first
listed is a more typical plant from river gravels.
These specimens apparently represent the first
report of O. arctica from the Yukon since its ini-
tial collection in 1906 on Herschel Island by
Lindstrom.

Oxytropis campestris ( L. ) DC

var. jordalii (Porsild) Welsh

Locality #15, WR 10572, 12 July; #59, WR
10455, 10 July. Gravel bars. Both specimens were
collected from polychrome-flowered populations.
The latter were pink in the field and have faded
bright blue-purple as the specimens dried. The
combination of small flowers, few flowered ra-
cemes, and few leaflets (17 or less) per leaf
serve to unite these specimens with var. jordalii.
This is the second know report of var. jordalii
from the Yukon.

var. varians (Rydb.) Bameby

Locality #.50, WR 10526, 11 July; #68, Rig-
by 160, 20 July; #71, WR 10494, 11 July. The
first specimen cited is a dwarf alpine phase, the
latter two are typical of the stream phase of
var. varians.

Oxytropis deflexa ( Pallas ) DC.

var. foIioJosa ( Hook. ) Bameby

Locality #59, WR 10058, 30 June; do, WR
10454, 10 July. Gravel bars. This is the first
knov\Ti report of O. deflexa for northern Yukon.

Oxytropis maydelliana Trautv.

Locality #1, WR 10266, 5 July; #32, WR
10206, 3 July; #34, WR 10494, 9 July; #36,
WR 10200, 3 July; #37, WR 10089, 30 June;

#46, WR 10585, 12 July; #51, WR 10513a, 11
July; #,54, WR 102,82, 6 July; #59, WR 10457,
10 July. Alpine tmidra, on shale, slate, schist, and
limestone, and gravel bars, tussock tundra and
heathlands. Widespread and locally abundant.

Oxytropis nigrescens ( Pallas ) DC.

var. nigrescens

Locality #3, WR 10177, 1 July; #7, WR
10153, 1 July; #9, WR 101.39, 1 July; #16, Rig-
by 16, 24 June; #23, Rigby 94, 25 June; #37,
WR 10084, ,30 June; #43, WR 10719, 15 July;
#44, WR 10.366, 9 July; #59, WR 10456, 10
July. Ridge tops, on limestone, sandstone, granite,
shales and slates, and on stream gravels.

Oxytropis viscida Nutt.

Locality #3, WR 10173, 1 July; #6, WR
10165, 1 July; #12, WR 10441, 10 July. Gravel
bars, along arctic streams, and on limestone
ridge tops. This is the first report of a viscid
oxytrope for northern Yukon.

Lentibulariaceae Bladderwort Family

Pinguicula vulgaris L.

Locality #15, WR 10569, 12 July; #52, WR
10483, 11 July. Wet pond margin and river
bank. This is the first apparent report of P. vul-
garis for northern Yukon.

Utricularia intermedia Hayne

Locality #52, WR 10482, 11 July. Emergent,
in shallow pond. This is the first report of U.
intermedia for northern Yukon.

Menyanthaceae Buckbean Family

Menyanthes trifoliata L.

Locality #52, WR 10486, 11 July. Emergent,
in shallow ponds in Old Crow Flats.

Nymphaeaceae Waterlily Family

Nuphar polysepalum Engelm.

Locality #52, WR 10477, 11 July. Lakes, in
Old Crow Flats.

Onagraceae Evening-primrose Family

Epilohium angustifolium L.

Locality #63, Rigby 113, 24 July; #69, Rig-
by 148, 20 July; #70, WR 10666, 15 July; #76,
WR 10429, 10 July. Gravel bars, stream terraces,
and lake and pond margin.

Epilohium latifolium L.

Locality #5, WR 10608, 13 July; #12, WR
10442, 10 July; #25, WR 10626, 13 July; #26,

52

Bricham Vounc: University Science Bulletin

10705, 15 July; #31, WR 10400, 9 July; #68,
Rigby 153, 20 July. Stream ;ind river gravels.

EpHohium palustre L.

var. lapponicutn Walil.

Locality #70, WR 10668, 15 July; #10658,
14 July. Lake margins.

Orobanchaceae Broomrape Family

Boschnkikia rossica ( CI & S. ) Fedtscli.

Locality #28, Rigby 114, 24 July. In shmbby
tundra. This is the most northern record of B.
rossica for the Yukon.

Papaveraceae Poppy Family

Papaver radicatwn Rottb.

Localit)' #2, WR 19242, 5 July; #4, WR
10227, 10228, 4 July; #5, WR 10614, 13 July;
#18, WR 10556, 12 July; #21, Rigby 118, 2.5
July; #22, WR 0708, 15 July; #50, WR 10533,
11 July; #53, Rigby 112, 22 July. Ridge tops
and talus slopes, on limestone, shale, and slate,
and less commonly in tussock tundra.

Plantaginaceae Plaintain Family

Plantago canescens Adams

Locality #76, WR 10419, 10 July; #77, Rig-
by 12, 18 June. River terraces and gravel bars.

Polemoniaceae Phlox Family

Phlox sibirica L.

var. borealis (Wherry) Welsh

Locality #1, WR 10259, 5 July; #3, WR
10181, 1 July; #9, WR 101.34, 1 July; #10, Rig-
by 88, 89, 25 June; #22, WR 10710a, 15 July;
#35, WR 10219, 3 July; #37, WR 10090, 30
June; #48, WR 10342, '9 July. Ridge tops, on
limestone, shale, slate, and sandstone, in tundra.
This is the first report of P. sibirica for northem
Yukon.

Polemonium boreale Adams

Locality #2, WR 10250. 5 July; #3, WR
10172, 1 July; #4, WR 10237, 4 July; #16, Rig-
by 13, 23 June; #2.5, WR 1062.5, 1.3 July; #26,
WR 10703, 15 July; #27, Rigby 142. 28 July. Al-
pine tundra, on shale, slate, and limestone, and
on river bars.

Polemonium caeruleum L.

var. villosuiu ( Rud. ) Brand

Locality #5, WR 10600, 13 July; #14, WR
10669, 15 July; #29, WR 10729, 1.5 July; #3.3,
WR 10.385, 9 July; #42, WR 10.3:38, '8 July.

Heathlands, willow thicket, and meadows, on
gravel bars, beaches, and spits.

Polygonaceae Buckwater Family

Polygonum al])inum All.

ssp. alaskanum (Small) \\'elsh
Locality it 29, WR 107.34, 15 July; #69, Rig-
by 151, 20 July; #76, WR 10428, 10 July; #79,
Rigby 1.32, 26 July. Gravel bars, terraces, and
river bluffs and lake shores, in Old Crow Flat,
and northward along the Blow River almost, or
quite, to the coast. These are the first records
of P. alpinum from northern Yukon.

Polygonum historta L.

ssp. plumosum (Small) Hulten
Locality #5. WR 10620, 13 July; #31, WR
10410, 9 July; #.39, WR 10127, 1 July; #40. WR
10.502, 11 July; #42, WR 10.326, 8 July; #50,
WR 10.528a, 11 July; #56. WR 10,592, 12 July;
#.57, WR 10072a, '30 June; #60, WR 100.34.
Meadows, tussock tundra, stream gravels, and
taiga. Broadly distributed, common.

Rumex arcticus Trautv.

Locality #42. WR 10.3.37, 8 July; #62 WR
10596, 13 July; #70, WR 10667, 15 July. Stream
banks, and lake and pond margins. Uncommon.

Primulaceae Primrose Family

Androsace chamaejasme Host.

ssp. lehmanniana (Spreng. ) Hulten
Localitv' #10. Rigby 84, 25 June; #16, Rig-
by 38, 23 June. River gravels and alpine tundra.
Uncommon.

Dodocatheon frigidum Cham. & Schlecht.

Locality #4, WR 102.30. 4 July; #.39, WR
10128, 1 July; #50, WR 105:^, 11 July; #.54,
WR 10280, 6 July; #57, WR 10079, 30 June.
Meadows, stream banks, and snow flushes.

Douglasia arctica Hook.

Locality #29, WR 10742, 15 July; #41, WR
10295, 8 July; #43, WR 10718. Slaty ridge tops,
in Neruokpuk fonnation and on gravel bars.
Specimens on gravel bars grow in rounded, hemi-
spheric clumps, those on ridge in depressed-
pulvinate mats.

Douglasia ochotensis ( Willd. ) Hulten

Locality #4, WR 10240, 4 July; #9, WR
101.37, 1 July; #17, WR 10559, 12 July. Slaty
ridge tops in Neruokpuk fonnation, and on Lis-
bume limeston(>. This is the first report apparent
of D. ochotensis in northem Yukon.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

53

Pyrolaceae Wintergreen Family

Pi/rola grandiflora Radiu.s

Locality #2-5, WR 10629a, 13 July; #56, WR
10593a, 12'july; #60, WR 10042a, 30 June; #63,
WR 10307, 8 July. Gravel bars and lake margins,
in heath, thickets, and taiga.

Ranunculaceae Buttercup Family

Aconitum dclphinifolium DC.

van delpliiiiifolium

Locality #5, WR 10602, 13 July; #27, Rig-
hy 140, 2S July. Stream gravels.

Anemone driimmondn Wats.

Locality #2, WR 10246, 5 July; #4, WR
1021Sa, 3 July; #6, WR 10168a, 1 July; #10,
Righy 87, 25 June; #48, WR 10347, 9 July. Al-
pine tundra, on limestone, slate, and shale, and
on gravel bars along arctic streams.

Anemone parviflora Michx.

Locality' #16, WR 16, 26, 37. 23 June; #40,
WR 10504, 11 July; ^57, WR 10075, 30 June;
#59, W R10054, 30 June. Alpine tundra in mea-
dows, and on stream gravels. Common.

Anemone patens L.

Locality #5, WR 10607, 13 July; #37, WR
10085, 30 June; #44, WR 10389, 9 July; #73,
Rigby 40, 24 June. Alpine tundra, on sandstone,
shale and limestone outcrops.

Anemone richardsonii Hook.

Locality #3, WR 10411, 9 July; #42, WR
10334, 8 July. Thickets and meadows, along
streams.

Caltha palustris L.

var. arctica ( R. Br. ) Huth.

Locality #6, WR 10168, 1 July; ii62, WR
1002Sa, 30 June. Pond and stream margins.
Uncommon.

Dclplunium ^lanciim Wats.

Locality #27, Rigby 139, 28 July; #71, WR
10497, 11 July. River bars and stream banks. Un-
common.

Ranunculus hyperboreus Rottb.

Locality #14, WR 10672, 15 July; #72, WR
10652, 14 July. Pond and lake margins and
muddy shores.

Ranunculus hipponicus L.

Locality #30, WR 10321, 8 July. Moist wil-
low-grass-cottongrass community. Uncommon.

Ranunculus nivalis L.

Locality #19, WR 10545, 12 July; #42, WR
103.33, 8 July; #64, WR 10637, 14 July. Snow
flushes, in alpine tundra, hcathlands, and willow
thickets. Locally common.

Ranunculus pallasii Schlecht.

Locality 62, WR 10594, 13 July. Boggy area,
adjacent to Sam Lake. Uncommon.

Ranunculus pijgmaeus Wahl.

Locality #42, WR 10329, 8 July; #50, WR
10535, 11 July; #64, WR 10638, 14 July. Snow
flushes, in alpine tundra, heathlands, and wil-
low thickets. Locally common.

Ranunculus turneri Greene

Locality #40, WR 10505, 11 July; #59, WR
10055, .30 June; do, WR 10463, 10 July; #64,
WR 10633, 14 July. Gravel bars and thickets.
Locally common.

Rubiaceae Madder Family

Galium trifidum L.

var. trifidum

Locality #72, WR 10653, 14 July. Lake shore.
Uncommon.

Rosaceae Rose Family

Dnjas integrifolia Vahl

var. integrifolia

Locahty #6, WR 10164, 1 July; #16, Rigby
18, 36, 39, 23 June; do, Rigby 63, 24 June; #57,
WR 10076a, .30 June; #60, WR 10044a, 30 June.
Tundra, heathlands, and taiga. A dominant
species.

Drijas octopetala L.

var. kamtschatica (Juz. ) Hulten

Locality #5, WR 10616, 13 July; #7, WR
10159, 1 July; #33, WR 10.383, 9 July; #55, WR
10067a, 30 June; #57, WR 10077, 30 June. Al-
pine tundra and meadows, on limestone, shale,
siliceous conglomerates, and slate. A dominant
species.

var. octopetala

Locality #10, Rigby 85, 25 June; #73, Rig-
by 46, 24 June. Alpine tundra. These records
are apparently the first for D. octopetala from
northern Yukon.

Getim glaciale Adams

Locality #2, WR 10244, 5 July; #4, WR
10229, 4 July; #10, Rigby 83, 25 June. Alpine
tundra, on slate and shale.

54

Bhigiiam Young Univehsitv Science Bulletin

i

Fig. 44. West along the shoreline from King Point at approximately 69°07' N; 138°00' W. The seacliff is in soft
sediments with considerable permafrost ice showing in some of the deeper gullies. The cliff rises appro.ximately
1.50 feet above the ice-covered Beaufort Sea to the right. Arctic tundra, heathland, and maritime habitat.

Potentilla anserina L. 10551, 12 July; #46. WR 10584, 12 July; #48,

var. amerina WR 10519, 11 July; #53, Rigby 111, 22 July;

Locality #76, VVU 10.527, 10 July. River bars #^4, Rigby 13.5, 26 July. Alpine and arctic tun-

and terraces. This is the first evident report of ^ra, on limestone and slate.

P. anserina for northern Yukon. r, , .n i ^i o r. 1 1 i .

FotenttUa elegans Cham. & Schlecht.

Potentilla biflora Willd. Locality #10, Rigby 78, 25 June; #18, WR

Locality #5, WR 10617, 13 July; #19, WR 10555, 12 July; #20, WR 10543, 12 July. Alpine

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

55

tundra, on siliceou.s conglomerate and sandstone.
This is the first report of P. elcgans for northern
Yukon.

Potentilhi fniticosa L.

Locality #5, WR 10619, 13 July; #15, WR
10566, 12 July; #42, WR 10711, 15 July; #48,
WR 10521, 11 July; #50, WR 10531, 11 July;
#52, WR 10481, 11 July. Taiga, heathland, and
tundra. Widespread, and locally common.

Potentilhi hookeriana Lehm.

Locality #5. WR 0611, 13 July; #25, WR
10627, 13 'July; #33, WR 10381, 9 July; #50,
WR 10525, 11 July. Alpine tundra, and along
streams on gravel.

Potentilhi norvegica L.

Locality #69, Rigby 149, 20 July. Lake shore
This is the first report of P. norvegica from
northern Yukon.

PotentiUa pahistris L.

Locality #61, WR 10289, 7 July; #70, WR
10659, 15 July; #72, WR 10656, 14 July. Lake
shores, pond margins, and wet heathlands.

PotentiUa unijlora Ledeb.

Locality #1, WR 19272, 5 July; #9, WR
1013, 1 July; #10, Rigby 86, 25 June; #48, WR
10343, 9 July; #55, WR 10068, 30 June; #73,
Rigby 44, 24 June. Alpine tundra, on sandstone,
shale, limestone, and slate. Locally abundant.

Rosa acicularis Lindl.

yar. botiigeauiana Crepin

Locality #67, WR 10094, 1 July; #68, Rig-
by 155, 20 July. Grayel bars, terraces, and bluffs.

Rubtis chamaemorus L.

Locality #40, WR 1050.5a, 11 July; #60, WR
10039, 30 June. Heathland and taiga.

Spiraea beauverdiana Schneid.

Locality #29, WR 10735, 15 July; #79, Rig-
by 131, 26 July. Grayel bars and grayelly slopes.
This is the first report of S. beauverdiana for
northern Yukon.

Salicaceae Willow Family

Populus balsamifera L.

Locality #27, Rigby 147, 28 July; #68, Rig-
by 98, 20 July; #71, WR 10499, 11 July. Grayel
bars and terraces. The specimen from locality
#27 is the northernmost record of this species in
the Yukon.

Salix alaxensis (Anderss. ) Coy.

yar. alaxensis

Locality #6, WR 10162, 1 July; #24, WR
10118, 1 July; #36, WR 10193, 3 July; 68, Rig-
by 99, 20 July; #77, Rigby 21, 18 June. Riyer
and stream bars and terraces and lake shores.
A dominant species in the plant community along
drainages throughout the region; the tree willow
of the arctic.

var. longistijlis ( Rydb. ) Schneid.

LocaUty #77, Rigby 3, 18 June. River bar.

Salix arctica Pallas

Locality #57, WR 10075, 30 June. Meadow,
in alpine tundra.

Salix brachijcarpa Nutt.

ssp. niphoclada ( Rydb. ) Argus
Locality #6, WR 10163, 1 July; #11, WR
10188, 1 July; #13, WR 10694, 15 July; #31,
WR 10409, 9 July; #36, WR 10196, 3 July; #50,
WR 10357, 9 July; #59, WR 10064, ,30 June. Al-
pine tundra on hmestone, and arctic tundra on
slate, and in tussock tundra, and on gravel bars
along streams and lake shores. This entity is a
dominant shrub in all major arctic plant com-
munities.

Salix Candida Flugge

Locality #52, WR 10478, 11 July. Lake mar-
gin, in wet taiga. This is the first record of S.
Candida for northern Yukon.

Salix chainissonis Anderss.

Locality #42, WR 10339, S July. Snow flush
and meadow adjacent to willow thicket. Appar-
ently this is the first record of S. chamissonis
for the Yukon.

Salix fuscescens Anderss.

Locality #62, WR 10028, 30 June. Stony
pavement beach of Sam Lake.

Salix glauca L.

var. acutifolia (Hook.) Schneid.

Locality #1, WR 10271, 5 July; #5, WR
10510, 11 July; #15, WR 10581, 12 July; #30,
WR 10324, 8 July; #36, WR 10196a, 3 July;
#52, WR 10664, 15 July; #60, WR 10038, 30
June; #67, WR 10107, 1 July. Lake margins,
stream banks, terraces and gravel bars in timdra,
heathland, and taiga. Widely distributed, com-
mon, and dominant in several plant communities.

Salix hastata L.

Locality #6, WR 10163a, 1 July; #15, WR
10567, 12 July; #25, WR 10629, 13 July; #37,
WR 10404, 9 July; #54, WR 10275, 6 July; #67,

56

Bricham Young University Science Bulletin

WR 10109, 1 July. Gravel bars, stream banks,
and terraces. This is the first report of S. novae-
angliae for nortlieni Yukon.

Salix lanata L.

ssp. richanlsonii (Hook.) Skvortsov
Locality #6, WR 10162a, 1 July. Gravel bar.

Salix pJilchophijUa .Anderss.

Locality :;?S, WR 10150, 1 July; #9, WR
10145, 1 July; #25, WR 10631, 13 July; #40,

WR 10501a, 11 July; #55, WR 10067, 30 June.
Alpine tundra, on shale, ancient beaches, and
alluvium. Widespread, and locally abundant.

Salix planifolia Pursh

ssp. j)ulchra (Cham.) Argus var. pulchra
Locality #11, WR 10189, 1 July; #24, WR
10121, 1 July; #30, WR 10.322, 8 July; #36, WR
10195, 3 July; #62, WR 1002.3, 10027, 30 June;
#64, WR 106.35, 14 July. Lake shores, drainages,
deltas, and shrubby tundra. A dominant species.

StflUtoH^

-j-^i

Fic. 45. Southeast alon- iln ,| ii a Kihl, I hi ilnnj; th li m. -I the Beaufort Sea at approximately 69°07' N;
137°58' W. The Rithanlson Mouiit.iiiis torni tlie laiiit hills .ilong the skyhne. Major large logs in the fore-
ground are principally conifers brought down the Mackenzie River from forests in the Northwest Territories
and British Columbia to the south. Collecting locality 14.

Biological Series, Vol. 14, No.

Reconnaissance of Northern Yukon

57

Salix reticulata L.

Locality #16. Rigby 28, 23 June. River bank,
in gravel, and in alpine tiindra.

Saxifragaceae Saxifrage Family

Bot/kinia richardsonii Hook.

Localitv #1, WR 10258, 5 July; #4, WR
10231, 4 July; #5, WR 10.597, 13 July; #31, WR
10399, 9 July; #46, WR 10582, 12 July. Slopes
and ridges, in alpine tundra, on slate, shale, and
limestone.

Clirijsosplenium tclrandrum (Lund) Fries

Locality #72, WR 10657, 14 July. Lake shore.

Chrijsosplenium nrightii Franch. & Sav.

Locality #2, WR 10255, 5 July. Alpine tun-
dra, on Kayak shale.

Parnassia kotzebuei Cham.

Locality #67, WR 10114, 1 July. Gravel bar.

Parnassia palustris L.

var. neogoea Fern.

LocaUty #15, WR 10571, 12 July; #21, Rig-
by 117, 25 July; #53, Rigby 110, 22 July. On
limestone in alpine tundra, and with willows on
gravel bars.

Saxifraga hronchialis L.

var. purptireomaciilata Hulten

Locality #43, WR 10724, 15 July. Alpine
tundra, on Neruokpuk formation. Evidently un-
common.

Saxifraga cerniia L.

Locality #14, WR 10671, 15 July; #62, WR
10224, 4 July. Lake shores, and gravels along sea
shores.

Saxifraga caespitosa L.

Locality #17, WR 10557, 12 July; #19, WR
10548, 12 July. Snow flushes, cliffs, and talus
slopes, on Lisbunie limestone. This is the first
record of S. caespitosa for northern Yukon.

Saxifraga davurica Willd.

var. grandipetala (Engler& Imischer) Welsh
Locality #2, WR 102.51, 5 July; # 10236, 4

July. Alpine tundra, on shale and slate.

Saxifraga eschscholtzii Stcmb.

Locality #4, WR 10213, 3 July. Alpine tun-
dra, on Neruokpuk slate. This is the first record
of S. eschscholtzii for the Yukon.

Saxifraga exilis Steph.

Locality #31, WR 10406, 9 July. Steep slope,
with willows, on Jurassic Kingak shale.

Saxifraga flagellaris Willd.

var. flagellaris

Locality #9, WR 101,36, 1 July; #47, Rigby
108, 22 July. Alpine tundra, on slate, shale, and
limestone. This is the first record of S. flagellaris
for northern Yukon.

Saxifraga hirculus L.

Locality #22, WR 10707, 15 July. Meadow,
along Bear Creek. Evidently uncommon.

Saxifraga oppositifolia L.

Locality #23, Rigby 95, 2.5 June; #44, WR
10376, 9 July. Alpine tundra, on granite and
limestone.

Saxifraga punctata L.

var. nel-soni (D. Don) Macoun

Locality #4, WR 10238a, 4 July; #11, WR
10185, 1 July; #14, WR 10669a, 15 July; #31,
WR 10407, 9 July; #.36, WR 10197, 3 July; #37,
WR 10086, ,30 June; #40, WR 10,503, 11 July;
#42, WR 10.336, 8 July. A dominant species in
tussock and shrubby tundra over a series of geo-
logical formations.

Saxifraga reflexa Hook.

Locality #1, WR 10265, 5 July; #2, WR
10252, 5 July; #3, WR 10179, 1 July; #9, WR
10144, 1 July; #16, Rigby 70, 24 June; #17, WR
10561, 12 July; #19, WR 10549, 12 July; #37,
WR 0093, ,30 June; #43, WR 10717, 15 July.
Alpine tundra, on slate, shale, and limestone, and
on stream banks.

Saxifraga rivularis L.

var. rivularis

Locality #42, WR 10,328, 8 July. Snow flush,
along a mossy bank.

Saxifraga serpyllifolia Pursh

Locality #2, WR 10253, 5 July. Alpine tun-
dra, on Kayak shale.

Saxifraga tricuspidata Rottb.

Locality #1, WR 10268, 5 July; #9, WR
10132, 1 July; #21, Rigby 116, 25 July; #35, WR
10220, 3 July; #42, WR 1032.5, 8 July; #48, WR
10,3.58, 9 July; #51, WR 10512, ll' July; #,55,
WR 10069, ,30 June; #65, WR 10298, i040b, 8
July; #73, Rigby 48, 24 June; #74, Rigby 136,
26 July; #7.5, Rigby 146, 26 July. A dominant
species in alpine tundra, in several communities.

Scrophulariaceae Figwort Family

Castilleja elegans Malte

Locality #78, Rigby 122, 26 July. Gravelly
bluff.

58

Bricham Young University Science Bulletin

Castillepa Injperborea Peiiiu'll

Locality #22, WR 10710, 15 July; #37, WR
10()88a. 30 June; #43, WR 10723, 15 July; #44,
WR 10373, 9 July; #46, WR 10583, 12 July; #47,
Righy 105, 22 July; #58, WR 10 147, 1 1 July. Al-
pine tundra ridge tops, on limestone, sandstone,
slate, and shale. Loeally common.

Castilleja pallida ( L. ) Spreng.
ssp. caudata Pennell
Locality il3, WR 10695, 15 July; #15, WR
10573, 12 July; #31, WR 10403, 9 July; #54,
WR 10274, 6 July; it 59, WR 10056, .30 June;
do, WR 10466, lOJuly; #67, WR 10100, 1 July.
Grayel bars, terraces, and bluffs, and less com-
monly in tundra.

Castilleja raupii Pennell

Locality #70, WR 10665, 15 July. Pond mar-
gin, in heath-taiga vegetation. Apparently, this
is a new record for northern Yukon.

Lagotis glauca Gaerto.

Locality #3, WR 10170, 1 July; #4, WR
10235, 4 July; #10, Rigby 93, 25 June; #11, WR
10183, 1 July; #24, WR 10124, 1 July; #37, WR
0088, 30 June; #57, WR 10073, 30 June; #62,

WR 10029, 30 June. Meadows, lake margins, tus-
sock tundra, and ridge tops, on shale, slate, and
limestone.

Pedicularis cupilata Adams

Locality #1, WR 10267, 5 July; #3, WR
10174, 1 July; #7, WR 10155, 1 July; #11, WR
10186, 1 July; #13, WR 10691, 15 July; #30,
WR 10320, 8 July; #37, WR 10087, 30 June;
#.50, WR 10,528, 11 July; #.57, WR 10281, 6
July; #62, WR 100.30a, .30 June; #65, WR 10299,
8 July. Tussock tundra, meadows, and deltas,
on sandstone, shale, limestone, slate, and alluv-
ium. Common. A dominant species.

Pedicularis kanei Durand

Locality #1, WR 10272a, 5 July; #16, Rig-
by 17, .32, '23 June; do, Rigby 67, 24 June; #44,
WR 10372, 9 July; #57, WR 10072, .30 June. Al-
pine tundra and with willows along stream
banks.

Pedicularis labradorica Wirsing

Locality #33, WR 10.382, 9 July; #47, WR
10360, 9 July; #48, WR 10520a, 11 July; #52,
WR 10480, 11 July; #60, WR 10040, 30 June;
#61, WR 10288, 7' July; #65, WR 10302, 8 July.

Fic. 46. Elf icefield alciiii; tlie hc-.uiu;itiTs ui Canoe River. K.ivak Shale tonus the low dark e.xposuu's .iloiig the
river and i.s capped by light colored Lisburne Limestone. Cretaceous rocks form the steeply dipping euesta.s
along the skyline in the background. As ice field melts, willows encased in it grow leaves, produce flowers, and
fruit. Tus.sock tundra and heathlands.

Biological Series, Vol. 14, No, 2 Reconnaissance of Northern Yukon

Tussock tundra, heatlilands, and taiga, and alpine
tundra on ridge tops and rock stripes, on sili-
ceous conglomerate, limestone, and sandstone.

Pedicularis langsdorfii Fisch.

Locality #13, WR 10690, 15 July; #14, WR
10684, 15 July; ii24, WR 10125, 1 July; #30,
WR 10316, 8 July; #57, WR 10283, 6 Julv; #62,
WR 1002.5, 30 June; -63, WR 10309, 8 Julv;
#64, WR 106.34, 14 July; #6.5, WR 10.304, ' 8
July. Lake shores, heathlands, wet meadows,
tussock tundra, stream banks, deltas, and spits.

Pedicularis oedcri \'ahl.

Locality #10, Rigby 76, 25 June. Alpine tun-
dra.

Pedicularis sudetica Willd.

var. hicolor Walpers

Locality- #14, WR 10683, 15 Julv; #.30, WR
10317, 8 Julv; :=:60, WR 10043, ;30' June; #62,
WR 10222, i0222ii, 4 July; #63, WR 10.308, 5
July. Lake shores, stream banks, and deltas, in
heathland, tundra, and taiga.

var. gijmnocephala Trautv.

Locality #52, WR 10488, 11 July. Wet heath-
land, in taiga. This is the first report of these ts\'0
varieties for northern Yukon.

Pedicularis verticillata L.

Loailit\' #15, WR 10568, 12 July. Gravel
bar, with willow and dwarf birch.

Umbelliferae Carrot Family

Bupleurum triradiatum Arams

ssp. arcticum (Regel) Hulten
Locality #.33, Wr" 10,387, 9 July; #,34, WR
10.398, 9 July; #43, WR 10715, 15 July; #44,
WR 10374, 9 Julv; #48, WR 10.349, 9 Julv; do,
WR 10519a, 11 July; #66, Rigby 104, 22 July.
Alpine tundra, on siliceous conglomerate, lime-
stone, slate, and shale.

Conioselinum cnidifolium (Turcz. ) Porsild

Localit)' #76, WR 10433, 10 July. River ter-
race, in white spruce woods.

Valerianaceae Valerian Family

Valeriana capitata Pallas

Locality #5, WR 10.599, 13 July; #10, WR
10190, 1 Julv; ii 14, WR 10689, 15 July; #25,
WR 10628, 13 July; #33, WR 10384, 9 July;
#51, WR 10.507, 11 July; #60, WR 10035, .30
June. Tundra, heathland, and taiga.

PTEROPSIDA-ANGIOSPERMAE-
MONOCOTYLEDONAE

Cyperaceae Sedge Family

Carex aquatilis Wahl.
ssp. aquatilis
Locality #15, WR 10575, 12 Julv; #52, WR
10484, 11 July; #62, WR 10031, ,30 June; #70,
WR 10661, 15 July. Emergent in shallow ponds
and lakes, and on lake shores, gravel bars, and
stream banks. This is the first report of ssp.
aquatilis from northern Yukon.

Carex bigelovii Torr.

Locality #14, WR 10680, 15 July; #31, WR
10412, 9 July; #36, WR 10198, 3 July; #40, WR
10506a, 11 July; #42, WR 10327, 8 July; #56,
WR 10.588, 12' July; #57, WR 10077a, ,30 June;
#60, WR 100,37, ,30 June; #61, WR 10285, 7
July; #65. WR 10,30,3a, 8 July. A dominant
species in tussock tundra, and on rockstripes
and outcrops of sandstone and limestone, and on
gravel bars and spits.

Carex capillaris L.

Locality #15, WR 10,574, 12 July. Gravel
bar, with willow and dwarf birch. This is the
first report of C. capillaris for northern Yukon.

Carex chordorrhiza Ehrh.

Locality #52, WR 10489, 11 July. Emergent
in shallow ponds, in wet heath-taiga.

Carex concinna R. Br.

Locality #54, WR 10278, 6 July. Wet mea-
dow.

Carex diandra Schrank

Locality #52, WR 10485, 11 July. Emergent
in margin of shallow lake. This is the first record
of C diandra from northern Yukon.

Carex limosa L.

Locality #52, WR 10491, 11 July. Emergent,
in shallow water, in wet heath-taiga. This is
the first record of C. limosa for northern Yukon.

Carex macloviana d' Urville

Locality #40, WR 10,504a, 11 July. Gravel
bar. The specimen is fragmentary but apparently
belongs to C. macloviana, and is the first record
of the species for northern Yukon.

Carex maritima Gunn.

Locality #61, WR 10287, 7 July. Shallow
pond, in polygonal tundra.

Carex misandra R. Br.

Locality #19, WR 10.546a, 12 July; #44, WR

60

Brigham Young Universiti' Science Bulletin

10371, 9 July. Limestone outcrops, in alpine tun-
dra. This is the first record of C. misandm for
northern Yukon.

Carex nardina Fries

Loc-alitv #11, WR lOlSSa, 1 July; #17, WR
10560, 12 July; #44, WR 10375, 9 July. Lime-
stone and slate ridge tops, in alpine tundra.

Carex petricosa Dewey

Locality #1.5, WR 10576, 12 July; #19, WR
10546, 12 July. Limestone, in alpine tundra, and
on gravel tar in stream course.

Carex podocarpa R. Br.

Locality #7, WR 10156, 1 Julv; #9, WR
10146, 1 July; #20, WR 10542, 12 July; #24,
WR 10122, l' July; #41, WR 10290, 8 July; #43.
WR 10726, 15 July; #64, WR 1063a, 10645, 14
July. On slate, conglomerate, and sandstone; in
alpine tundra, and in tussock tundra, heathlands,
lake shores, stream banks, and grayel bars. A
dominant species.

Carex rotundata Wahl.

Locality #61, WR 10286, 7 July. Emergent
in shallow pond, in polygonal tundra. This is
the first report of C. rotundata for northern
Yukon.

Carex rupestris All.

Locality #34, WR 10394, 9 July; #46, WR
10587, 12 July. Limestone and slate ridgetops,
in alpine tundra.

Carex scirpoidea Michx.

Locality #15, WR 10578, 12 July; #44, WR
10.368, 9 July; #46, WR 10586, 12 July; #52,
WR 10493, 11 July. Lake shores and river bars,
in heath and taiga, and on limestone and slate
in alpine tundra.

Carex siipina Willd.

ssp. spaniocarpa (Steud.) Hulten
Locality #47, WR 10.361, 9 July. Siliceous
conglomerate rock stripes in alpine tundra. This
is the first report of C. supina for northern Yuk-
on.

Eriophorum aneiustifolhim L.

Locality #30, WR 10314, 8 July; #,52, WR
10490, 11 July; #62, WR 10311, 8 Julv; do, WR
100.30, .30 June; #72, WR 10650, 14 July. Lake
and pond margins.

Eriophorum chamissonis C. A. May.

var. alhidtim (Myl. ) Fern.

Locality #.30. WR 10318, 8 July. Delta of
Blow River, This is the first report of E. chamis-
soni for northern Yukon.

Eriophorum scheuchzeri Hoppe

Locality #62, WR 10225, 4 July; #72, WR
646, 14 July. Lake shores. This is the first report
of E. scheuchzeri for northern Yukon.

Eriophorum vaginatum L.

Locality #24, WR 10120, 1 July; #&3, WR
10310, 8 July; #64, WR 10632, 14 July. A dom-
inant species in the tussock tundra. Bv mid-
July, the tundra appeared to be covered by hoar-
frost due to the abundant inflorescences of E.
vaginatum. This is the first record of E. vagina-
tum for northern Yukon.

Kobresia myosuroides ( Ville. ) Fiori & Pavl.

Locality #11, WR 10187a, 1 July; #34, WR
10.392, 9 July. Alpine tundra, on slate and lime-
stone.

Scirpus caespitosus L.

Locality #52, WR 10492, 11 July. Lake shore.
This is the first report of S. caespitosus for north-
em Yukon.

Gramineae Grass Family

Agropijron caninum (L. ) Beauv.

var. latighime ( Scribn. & Sm. Pease & Moore
Locality #12, WR 10451, 10 July; #59, WR

10471, 10 July; #76, WR 10431, 10 July. Stream

gravels and river terraces.

Agropijron macrourum (Turcz. ) Drobov

Locality #26, WR 10697, 15 July; #31, WR
10413, 9 July; #.59, WR 10472, 10 July; #67,
WR 10098, ■ 10103, 10112, 1 July; #76, WR
10421, 10 July. Terraces and gravel bar.

Alopecurus alpinus Sm.

Locality #14, WR 10677, 15 July; #.30, WR
10315, 8 July. Spits, beaches, and deltas along
the Beaufort Sea. This is the first record for
northern Yukon.

Arctagrostis latifolia ( R. Br. ) Griseb.

Locality #15, WR 10577a, 12 July; #29, WR
10733, 15 July; #.56, WR 10.589, 12 July; #72,
WR 10648. 14 Julv. Gravel bars and lake shores.

Arctophila fulva (Trin. ) Anderss.

Locality #62, WR 10312, 8 July; do, WR
10.595, 13 July; #72, WR 10651, 14 July. Emer-
gent in shallow lakes and ponds.

Bromus pumpellianus Scribn,

Locality #12, WR 104.50, 10 July; #26, WR
10702, 15 July; #.32, WR 10212, 3 July; #71,
WR 10498, li July; #76, WR 10434, 10 July.

Biological Series, Vol. 14, No. 2 Reconnaissance of Northern Yukon

61

Fig. 47. Map of collection localities in northern Yukon.

62

Bric.ham Young University Science Bulletin

Stream terraces and gravel bars, and less com-
monly on sandstone in alpine tundra.

Calariwg^rostis canadensis ( L. ) Beanv.

Locality #68, Rigby 152, 20 July. Gravel
bar. This is the first r(>p()rt of C. canadensis for
northern Yukon.

Calanuigrostis inexpansa (Iray

Locality #15, WR 10577, 12 July; #29, VVR
10745, 15 July. Gravel bars. This is the first
record of C. inexpansa for northern Yukon.

CalamagrostLs purpttrascens R. Br.

L(K-ality #21. Rigby 121, 25 Julv; ii:.34, WR
10396, 9 July; #48.^ WR 10351, 9 July; #68,
Rigby 102, 20 July. Gravel bars, and on rocky
limestone outcrops in alpine tundra. This is the
first report of C'. purj)uresccns for the northern
Yukon.

Deschampsia caespitosa ( L. ) Beauv.

Locality #29, WR 10732, 15 July; #31, WR
10416, 9 July. Gravel bars.

Ehpntis innovatiis Real

Locality #2, WR 10248, 5 July; #15, WR
10579, 12 July. Gravel bars, and on shaly slopes
in alpine tundra.

Elipnns nwUis Trin.

Locality #14. WR 10676, 15 July. Sea beach
and spit.

Festuca altaica Trin.

Locality #31, WR 10417. 9 July. Stream
bank.

Festuca haffincnsis Polunin

Locality #29, WR 10731, 10743, 15 July.
Gravel bar. This is the first report of F. hafjincn-
sis for northern Yukon.

Festuca hrachijhpijUa Schult.

Locality #17, WR 10558a, 12 July; dr44, WR
10370, 9 July. Limestone outcrops, in alpine tun-
dra.

Festuca rubra L.

Locality #12, WR 10449, 10 July; #25, WR
10630, 13 July; #26, WR 10696. 1.5 July; #56,
WR 10593, 12 July; #59, WR l()47.3a, 10 Julv;
#68, WR 101, 20 'July. Gravel bars. This is the
first record of F. rubra for northern Yukon.

HierochJoe aJpina ( S\v. ) R. & S,

Locality #7, WR 101.57. 1 July; #18, WR
10553, 12 July; #29, WR 10747, 15 July; #42,
WR 10.330, 8 July. A dominant species in alpine

and dry tussock tundra, on slate, sandstone, and
alluvium.

Hordeum jubatum L.

Locahty #76, WR 10422, 10 July. River ter-
race, in open white spruce woods. This is the
first record ol //. jubaluni tor iiortiieni Yukon.

Poa alpigena ( Fries ) Lindm.

Locahty #76, WR 104.38, 10 July. River bar,
with other grasses and forbs.

Poa alpina L.

Locality ±H), Rigby 92, 25 July. Alpine tun-
dra, on Kayak shale.

Poa arctica R. Br.

Locality #29, WR 10730, 15 July; #34, WR
10.396a, 9 July; #45, WR 10,540, 12 July; #59,
WR 10469, 10474a, 10 July; #65, WR i0305, 8
July. Gravel bars, and in alpine tundra on sand-
stone, siliceous conglomerate and limestone.

Poa glauca Vahl

Locality #9, WR 101.30, 1 July; if29, 10741,
15 July; #42, WR 10296, 8 July; ii43, WR
10728, 15 July; #44, WR 10377, 9 July; #59, WR
10470, 10 July; #74, Rigby 138, 26 July. Alpine
tundra, on limestone and shale, and on gravel
bars.

Poa lanafa Scribn. & Merr.

Locality #29, WR 10730a, 15 July; #72, WR
10647, 14 July. Gravel bars and lake shores. This
is the first record for northern Yukon.

PuccineUia borealis Swallen

Locality #14, WR 10678, 10678a, 10679, 15
July; #76, WR 104.37, 10 July. River bars and
terraces, and sea beaches and spits.

Trisetum spicatuni (L. ) Richt.

var. inollc (Michx. ) Real

Locality #19, WR 10547, 12 July; #29, WR
10746, 15 July; #31, WR 10415, 9 July; #4.3,
WR 10727, 15 July; ii:59, WR 10474. 10 July.
Gravel bars, and in alpine tundra on slate and
limestone.

Juncaginaceae .\rrowgrass Fainih"

Triglochin nuiritinia L.

Locality #52, WR 10487, 11 July. Wet heath-
taiga around a small lake.

Juncaceae Rush Famih'

Junctts arcticus Willd.

Locality #31, WR 10414, 9 July. Gra\el bar.

Biological Series, \'ol. 14, No. 2 Reconnaissance of Northern Yukon

63

This is the first report of /. arctictis for northern
Yukon.

Jtincus trighimis L.

Locahty #52, WR 10490, 11 July. Wet heath-
land, surrounding a small lake. This is the first
record of /. trighimis for northern Yukon.

Luzula confitsa Lindeb.

Locality #41, WR 10294, 8 July; #42, WR
10335, 8 Julv; ti45. WR 10541, 12 July; #65,
WR 10304a, 8 July. Alpine tundra.

Liliaceae Lily Family

Allium scIiocnoj)r(istim L.

var. sibirictim (L. ) Hartm.

Locality #5, WR 10598, 13 July; #77, Rig-
by 1, 6, 10, 18 June; #78, Rigby 124, 26 July.
Gravel bars and terraces. This is the first report
for northern Yukon, and the Loney Creek local-
ity is apparently the only one known for the
species on the north slope.

TofieJcIia pusilla (Michx. ) Pers.

Locality #50, WR 10537, 11 July; #54, WR
10279, 6 July. Meadows and alpine tundra, on
gravel and on shale and limestone.

Zygadenus elegans Pursh

Locality #5, WR 10609, 13 July; #15, WR

10571a, 12 July; #21, Rigby 119, 25; #22, WR
10709, 15 July; #50, WR 10522, 11 July; #51,
WR 10511, 11 July; #71, WR 10497a, 11 July.
Gravel bars, and in alpine tundra on shale and
limestone.

Potamogetonaceae Pondweed Family

Poiamogeton alpimis Balbis

var. tenuifoliiis ( Raf . ) Ogden

Locality #70, WR 10660, 15 July. Shallow
pond. This is the first report of P. alj)inus for
northern Yukon.

<v

rt

d

4)

^

Oh

C8

Oi

lyi

o

0)

ri

a,

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q

C

y

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o

E

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>>

hJ

C/D

u<

O

0)
a!
(U
C
O

o
o

nJ
§

o

8 ^

§

O

Families

2

1

1

1

31

6

42

Genera

2

1

2

2

98

25

125

Species

2

2

2

2

203

68

279

Subspecies

15

15

Varieties

63

63

Formae

1

1

Fig. 48. Summary of ta.xa collected in northern Yukon.

ACKNOWLEDGMENTS

Collections were made as an outgrowth of a
geologic investigation of the British Mountains-
Bam Mountains region by Union Oil Company
of Canada and we express our thanks to officers
of the companv for providing a base. We are
especially grateful for support and interest of
Gerald Salisbury, Homer Johnston, Roland de
Caen, and others of Union Oil Company in Cal-
gary. In addition to collections by the writers,
some specimens were also added by J. Keith
Rigby, Jr., E. Blair Maxfield, C. Fredrick Lohren-

gel n, Lehi F. Hintze, Jim Nixon, and John
Habbishaw. John King and Jim Keene were our
efficient helicopter pilots, and with Ray Fessen-
den, engineer, and Ian Wright, mechanic, all of
Northern Mountain Airlines, Prince George,
B. C, greatly aided our collecting program.
Douglas Cridland was our excellent cook and
was assisted by Lee Whitehead and Tom Bul-
lock. Fixed wing logistical support was by Great
Northern Airlines, Inuvik, NWT.

REFERENCES

BosTOCK, H. S. 1948. Physiography of the Canadian
Cordillera, with special reference to the area north
of the fifty-fifth parallel; Geol. Survey Canada
Mem. 247. 106 p.

Gabhielse, H. 1957. Geological reconnaissance in
the Northern Richardson Mountains, Yukon and
Northwest Territories: Geol. Survey Canada Paper
.56-6, 11 p.

HuLTEN, E. 1968. Flora of Alaska and neighboring
territories. Stanford, California; Stanford Univer-
sity Press, 1008 p.

Jeletzky, y. A. 1961. Upper Jurassic and Cretaceous
rocks, west flank of Richardson Mountains between
the headwaters of Blow River and Bell River,
Yukon Territory, IIP and 117A (parts of); Geol.
Survey Canada Paper 61-9, 42 p.

Martin L. J. 1959. Stratigraphy and depositional
tectonics of the North Yukon-Lower Mackenzie area.
Amer. A,ssoc. Petrol. Geol. Bull., v. 43, no. 10,
p. 2399-2455. , ^

MOUNTJOY E. W. 1967a. Triassic stratigraphy ot
northern Yukon Territory: Geol. .Survey Canada
Paper 66-19, 44 p.

. 1967b. Upper Cretaceous and Tertiary strati-
graphy of northern Yukon Territory and northwest-
em District of Mackenzie; Geol. Survey Canada
Paper 66-17, 70 p. r , ^. i ■

NonFOHD, B. S. 1964. Reconnaissance ot the Urdovi-
cian ' and Silurian rocks of northern Yukon Terri-

tory: Geol. Survey Canada Paper 63-39, 139 p.

Nonnis, D. K., Price, R. A., and Mountjoy, E. W
1963. Geology of northern Yukon Territory and
northwestern District of Mackenzie: Geol. Survey
Canada Map 10-1963.

PoRsiLi), A. E. 1964. Illustrated flora of the C;mad-
ian' Arctic Archipelago. National Mus. Canad. Bulle-
tin No. 146, 218 p.

Reed, B. L. 1968. Geology of the Lake Peters area,
nortlieastem Brooks Range, Ala.ska: U. S. Geol.
Survey Bull. 1236, 132 p.

Welsh, S. L. 1971. Anderson's flora of Ala.ska and
adjacent Canada. Unpuhlished M.S. 1100 p.

rpt

QjO^Jd}

Brigham Young University
Science Bulletin

LIBRARY;

OCT 29 1971

HARVARD
UNlVERSlBCi

A FLORA FROM THE DAKOTA SANDSTONE

FORMATION (CENOMANIAN)
NEAR WESTWATER, GRAND COUNTY, UTAH

by
Samuel R. Rushforth

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 3
SEPTEMBER 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department ot Botany,

Brigham Young University, Provo, Utah

Members of the Editorial Board:

Vernon J. Tipton, Zoology
Ferron L. Anderson, Zoology
Joseph R. Murdock, Botany
Wilmer W. Tanner, Zoology

Ex officio Members:

A. Lester Allen, Dean, College of Biological and Agricultural Sciences
Ernest L. Olson, Chairman, University Publications

The Brigham Young University Science Bulletin, Biological Series, publishes acceptable
papers, particularly large manuscripts, on all phases of biology.

Separate numbers and back volumes can be purchased from Publication Sales, Brigham
Young University, Provo, Utah. All remittances should be made payable to Brigham
Young University.

Orders and materials for library exchange should be directed to the Division of Gifts
and Exchange, Brigham Young University Library, Provo, Utah 84601.

Brigham Young University
Science Bulletin

A FLORA FROM THE DAKOTA SANDSTONE

FORMATION (CENOMANIAN)
NEAR WESTWATER, GRAND COUNTY, UTAH

by
Samuel R. Rushforth

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 3
SEPTEMBER 1971

TABLE OF CONTENTS

Page

INTRODUCTION 1

GEOLOGICAL BACKGROUND 1

Litliological Cliaracteristics 1

Geology 4

THE AGE OF THE DAKOTA SANDSTONE 4

A CONCEPT OF A DAKOTA FLORA 7

PALEOECOLOGY 9

PALEOCLIMATE 12

SYSTEMATICS 13

Genus Equisctum 13

Genus Asplenium 14

Genus Conioplem 18

Genus Ildusmannia 18

Genus Gleichcnin 20

Genus Matouidium 27

Genus Astrulopteris 34

Genus Cliulophlehis 34

Genus Ilex 38

Genus Maniioliu 38

Genus Ficus 38

Genus Eucali/ptus 40

Genus Platanus 42

Genus Salix 42

ACKNOWLEDGEMENTS 42

REFERENCES CITED 43

A FLORA FROM THE DAKOTA SANDSTONE

FORMATION (CENOMANIAN)

NEAR WESTWATER, GRAND COUNTY, UTAH

by

Samuel R. Rushforth'

ABSTRACT

A Cretaceous (Cenomanian) flora from the
Dakota Sandstone Fomiation near Westwater,
Grand County, Utah contains an admixture of
ferns and angiospenns. The ferns of this flora
are representative of an older Jurassic-Wealden
\egetational type, whereas the angiosperms are
t\pical of the modem vegetational type. Species
of GleichenUi and Matonidhim and Astralopteris

coloradica represent the dominant forms in this
flora.

The Westwater flora contains fourteen genera
including nineteen species and one variety. New
species described from this flora include Asplen-
ium dakotensis, Coniopteris westwaterensis and
Ilex serrata.

INTRODUCTION

A fern-angiospenm flora from the Cretaceous
(Cenomanian) Dakota Sandstone Formation has
been under studv for some time. This flora was
collected from an ash seam and sandstone in
the Dakota Sandstone Fonnation from Rabbit
\'alle\', Grand Countv, Utah, and along the
road and surrounding areas between U.S. High-
wav 50 and Westwater, Grand County, Utah.

This Dakota flora is significant for three
reasons. First, it contains several new species
of fossil plants. Second, it extends the known

distribution patterns of manv previously de-
scribed species. The Westwater flora is one of
few paleofloras which illustrates an admixture
of an older Jurassic-Wealden floristic tvpe with
a modern angiospermous floral tvpe. Third,
this flora provides new information on a time
of the earth's history when angiosperms were
expanding from a position of little floristic im-
portance to a position of dominance in Cre-
taceous and later floras.

GEOLOGICAL BACKGROUND

LiTHOLOGic.\L Characteristics.— Two early
workers, Marcou (1864) and Capellini (Capel-
iini and Hecr, 1867), considered the Dakota
Group of Meek and Hayden (1856, 1861) to
have been deposited in fresh water. This deter-
mination was based primarily upon the included
leaf flora. However, Hayden (1867) stated that
together with F.B. Meek, he had colleceted well-
preserved marine invertebrate remains mingled
with the leaves of the Dakota Group. Hayden
concluded that the Dakota Group was marine
in origin. This conclusion was adhered to by
Lescjuereux (1874, and others) and by subse-
quent workers.

Lesquereux (1874) presented an excellent
discussion on the probable origin of the Da-
kota Group. Based upon studies of recent de-
positional facies and comparing them to the
Dakota, Lesquereux stated: "They are beach
formations, like those in progress at the present
time along the shore of the North Sea, in Hol-
land and Belgium, where the widelv extended
mudd\' shores are formed of a soft substance
of the red color." Lescjuereux further mention-
ed that the presenth' forming North Sea beach-
es are characteristic and similar to Dakota beds
by being composed of sands borne by the sea
intermixed with muds borne by the sea inter-

^Departjiient of Botany and Range .Stienie, Brighani Vowig Unl^eI^i^y

Bhicham Young University Science Bulletin

mixed with muds borne by nearby rivers. In
mentioning that the leases of some species often
exhibit a cUimped rather than a random distri-
bution in Dakota strata, Lesquereux (I.e.) postu-
lated:

A distribution of this kind can result only from
the proximity of the trees from which the
leaves liave been derived, and confirms the
opinion that the formation of the Dakota
group is the result of muddy flats whose sur-
face, raised perhaps in hillocks above water-
limits, and already solid j^round, was cut like
an immense swamp, here and there inter-
spersed by rare groups of trees and bushes.

In support of Lesquereuxs hypothesis, it
should be mentioned that within the Dakota
Sandstone Formation there are deposits indica-
tive of fresh and brackish water. Evidence for
this comes from hgnite seams, fossil assem-
blages, stratigraphy and lithology. These sedi-
ments could have been deposited in embay-
ments, estuaries, and on flood plains (Chaney,
1954; Repenning and Page, 1956.). However, be-
cause of frequent lignite beds and carbona-
ceous shales, and the great number of leaf im-
pressions, which, due to their excellent state of
preservation, do not appear to have been trans-
ported for any great distance, these deposits
appear to have been laid down in paludal en-
vironments close to the sea, similar to those
described bv Lesquereux (1874).

In discussing the nomenclature used in con-
nection with the Dakota Sandstone (as used bv
Meek and Hayden, 1856, 1861), Tester (1931)
preferred the useage of the term Dakota Stage.
His reasoning for the application of this term
sheds some light on the nature of the beds re-
ferred to the Dakota Sandstone. Tester stated
in part:

Geologic events which ;ire of considerable
magnitude, and which have some effect over
a large area, or which constitute a normal
progression of rocks, are considered responsi-
ble for the deposition of rocks comprising a
stage. The widespread marine advance, with
its shoreline variations due to minor retreats
and advances of the waters and to the lands
being built out into the oceans, or the migra-
tion of faunas and shifting of ocean currents,
all have tlieir effect on the character of the
rocks. The rocks deposited under such condi-
tions on an extensive scale, as they were dur-
ing Dakota time, are classed as a stage. It
miglit be said tliat a stage is indicative of a
.set of conditions of rock deposition rather than
of a distinct lithological or paleontological di-
vision.

From Tester's discussion, several important
concepts arc apparent. First, the geologic event

responsible for the deposition of the Dakota
Sandstone was of considerable magnitude and
had effect over a large area. Second, this geo-
logical event was the extensive marine advance
during Dakota times. Third, the Dakota Sand-
stone Formation indicates a set depositional
characteristics rather than conforming to a dis-
tinct lithological or paleontological division.

From the foregoing discussion, a working
definition of the Dakota Sandstone Formation
mav be given. This formation is a Cretaceous
time transgressive sequence of marine, fresh or
brackish water, clastic sediments of various
colors (with red, \ellow, and white being com-
mon) deposited under fluvial and paludal con-
ditions, often with interbedded lignite, shale
and ash secjuences. This formation can not be
defined on the basis of paleontological similari-
ties, although included fossil plants may be cor-
related to some extent. Strata comprising the
Dakota Sandstone Formation are related
throughout their geographical extent in that
thev were similarlv deposited as the result of
the invading Cretaceous sea. The Dakota Sand-
stone Fonnation exhibits continuity throughout
much of the western and midwestern United
States, authough sediments of this formation
from different regions need not be of the same
age due to the deposition of Dakota strata at
the edge of a coastline which changed with
time.

Tester (1931) pointed out that a wide variety
of names had been used for discussing Dakota
strata including Dakota Group (originally used
by Meek and Harden in 1861 in referring to
their Formation No. 1 of 1856), Dakota Series,
Dakota Stage, Dakota Formation, Dakota Sand-
stone, and Dakota without an\' additional term.
Tester (1931) preferred the use of Dakota Stage
which he used svnonvmously with the term
group, since he restricted the usage of group
for the rocks of an entire era.

The term group or stage usage is useful in
delimiting the rocks of more than one closely
related formation when the distinction between
them is either difficult or unnecessar\'. How-
ever, later workers (Jcnney, 1899; Darton, 1905;
Stanton, 1905) showed that the Dakota Group
of the Black Hills and Rock\- Mountain regions
could be divided into Lower and Upper Cre-
taceous formations; these workers restricted the
Dakota Sandstone to formational status. Fur-
thermore, the original Dakota Group from the
midwestern United States is similar strati-
graphicalK' and floristicalh' (although some
variation mav be noted in floras from different
regions) to the restricted Dakota Sandstone

Biological Series, Vol. 14, No. .3 D.\kut.'\ Sandstone Flora

UTAH

ARIZONA

Fic. 1. Index map of collecting localities.

HiuciiAM VoLNC University Science Bulletin

Formation of the Black Hills and Rock\' Monn-
tain regions. In light of this, the term Dakota
Sandstone Formation will be used by the author
for discussion of Dakota strata.

Geolcx;y. -Stanton (1905) in working with the
Jurassic Formation and its relationship with the
Comanche Series and the Dakota Formation in
Southern Colorado, New Mexico, and Okla-
homa, demonstrated that the Dakota Sandstone
of this region, as originally defined, contains
both Lower and Upper Cretaceous strata. Thus,
Stanton pointed out that the true Dakota Sand-
stone Fonnation of these widely separated re-
gions overlies Lower Cretaceous deposits which
overlie the Morrison Formation.

Similar conditions have been observed in
several other regions of the western United
States. In Montrose County, Colorado, the Da-
kota Sandstone Formation occupies a position
between the Jurassic Morrison Formation (Mc-
Elmo Formation of Coffin, 1921) and Upper
Cretaceous Mancos Shale Formation. Careful
examination of these strata (Coffin, 1921) in-
dicated that the Jurassic Morrison Fonnation
is overlain 1)\ a Lower Cretaceous sequence.
This sequence was given formational status bv
Coffin (1921) who named it the Post-McElmo
Formation. Stokes (1948) changed the name of
this Lower Cretaceous setjuence to the Burro
Canyon Formation.

Plants of this Lower Cretaceous setiuence in
Colorado were studied by Brown (1950). Brown
concluded that floristic evidence agreed with
the findings of Coffin (1921). A Lower Cre-
taceous flora was described bv Brown from

the Burro Cam on Formation, and a somewhat
at\pical Dakota flora was described from the
Dakota Sandstone Formation. This Dakota flora
is at\pical in that the incidence of ferns with
angiosperms is high.

Jurassic and Cretaceous stratigraphy near
Westwater, Grand County, Utah, is similar to
that of Montrose Countv, Colorado. The Jur-
assic Morrison Formation from this region of
Utah is overlain b\ the Lower Cretaceous
Cedar Mountain Fonnation. This formation is
conelated to the east with the Buno Canyon
Formation. However, no plants have been col-
lected from the Cedar Mountain Formation of
Grand County. This Lower Cretaceous for-
mation is overlain by the Dakota Sandstone
Formation (Plate 3.) This formation from this
region is composed of three lithological units.
The basal unit is a massive, buff-colored sand-
stone of approximately thirty feet in thickness.
This unit is overlain bv a shale-coal-sand se-
quence which is approximately thirt\' feet thick.
Fossil plants collected near Westwater are ob-
tained from this unit. This shale-coal-sand unit
is in turn overlain by a massive buff-colored
sandstone unit. This is the uppermost unit of
this formation near Westwater, and it is ap-
proximately thirty to forty feet in thickness.
This formation is overlain b\- the Upper Cre-
taceous Mances Shale Formation. Most leaf
compressions collected from the Dakota of this
region are obtained from an ash seam approxi-
mately forty to forty-five feet beneath the Grtj-
phiiea ncubernji zone in the overlving Mancos
Shale.

THE AGE OF THE DAKOTA SANDSTONE

Much of the earliest work concerning the
Dakota Sandstone Fonnation was concerned ex-
tensively with the age of these rocks. Briefly,
the Dakota was the first considered as a Cre-
taceous group (Meek and Hayden, 1S.56, 1858).
This interpretation was refuted b)- Hawn (1858)
who supposed that Dakota strata belonged in
the Triassic. Heer (1859, 1861) proposed that
the Dakota was of Tertiar\ age, and Marcou
(1858) stated that the original Dakota was com-
posed of rocks of both Tertiarx' and Jurassic
age. Later work (Marcou, 1864; Cajiellini and
Heer, 1867) demonstrated that Dakota rocks
indeed belong to the (Cretaceous, as originalh'
postulated bv Meek and Hayden. This age de-
termination has been adhered to b\- ail follow-

ing workers, and the only subsequent contro-
versy has been concerning the proper Creta-
ceous epoch to which Dakota strata should be
assigned.

An accurate ascertainment of the age of the
Dakota Sandstone Fonnation (whether early or
late Cretaceous) has been complicated by two
|)r<)l)lems. The first concerns a poor usage of
the term. "Dakota flora." This term has been
used loosely bv some workers, and as pointed
out by Berry (1920), "Any Cretaceous formation
containing dicot\ledonous leaves and known or
thought to be older than the Benton . . ." was
said to contain a Dakota flora. In other words,
Dakota flora became an adjectival term rather
tliaii dcliiniting a flora which had been ob-

BioLooicAi. Series, \'()l. 14, No. 3 Dakota Sandstone Flora

m

»'■ IT ■

-.iJ9.

.,^

'.r^

/--•

^*^-,

._. -w

..:^

V>.>.^~,*1

within the formation or from evolutionary
changes in plant communities over a period of
time svnchronou.s with deposition.

An alternative to the proposal of Berry
(1920) is that in delimiting a true Dakota flora,
only plants from the Dakota Sandstone Forma-
tion be considered. That is, the onlv valid ap-
plication of the term Dakota flora can be in the
discussion of plants from the Dakota Sandstone
sensu stricto. It is neither valid nor prudent to
delimit a Dakota flora, merclv in the sense of
a Cretaceous dicotyledonous flora, from anv
other geological entity.

This suggestion, however, leads to a dis-
cussion of the second problem which is some-
what less easily treated. It is that the Dakota
Sandstone is a time transgrcssive formation and

Fig. 2. .K. Southeast view down canyon towards West-
water, Grand County, Utah. The Dakota Sandstone
Formation forms the hills and slopes in the fore-
ground and is exposed along the roadcut. Original
collections from the Dakota Sandstone of this region
were made in the roadcut ( A ) . The Mancos Shale
Formation o\erlies the Dakota in this region.
B. Close-up of collecting site (A) in Fig. 1. This
site \ielded the best colections of fossil plants from
this area.

tained from the Dakota Sandstone. This prob-
lem became so acute that Berrv (1920) further
stated, "It has become increasingh' clear of late
years that Dakota flora was not a unit and had
no precise stratigraphic value." Berrv (1920) pro-
posed that in order to eliminate this problem,
a true Dakota flora mav be defined as "mean-
ing thereb\ the equivalent of that of the Wood-
bine Formation of Texas, and those of corre-
sponding age elsewhere. . . ." This interpreta-
tion, houever, has one inherent difficulty. That
is, it does not allow for the possibility that
within the same formation, fossil assemblages
ma\ differ sufficienth' to render their com-
|iarison with the flora from another formation
difficult. This ma\' result either from a non-
random geographical distribution of fossils

»i^fb»n

Fk;. .3. A. Overview of Dakota Sandstone Formation
showing massive upper sandstone unit resting upon
sliale-coal-sand imit. The arrow points to a collect-
ing excavation. B. Close-up of excavation in Fig.
1 illustrating lithology of shale-coal-sand unit. The
upper coal (B) and unfossiliferous ashes (A and C)
have been removed in this excavation exposing the
fossiliferous ash upon which the worker is kneeling.

6

Bni(;nAM VouNc; University' Science Bulletin

was deposited over a range of time that wit-
nessed a vast floristic change in the dominant
vegetation of the earth. That is, although all
formations are time transgressive, many are
deposited within a period of time wherein little
change in the contemporaneous flora or fauna
occurred, and therefore the included fossils of
that formation would be expected to exhibit
homogenitv. However, the Dakota Sandstone
Formation was deposited over a period of time
when the predominant vegetational type of the
earth changed from a fcrn-g\'mnosperm alliance
typical of the older Mesozoic to an angiospemi
dominated flora typical of the Late Cretaceous,
Tertiary, and present times. Therefore it is
possible to collect within this formation, fossil
assemblages which appear to be indicative of
different ages. This is particularly the case
when the Dakota Sandstone Formation from
west and east of the Rockv Mountains are com-
pared, and the floras from the western portion
of this formaton appear to be older than their
counterparts from the east. Further discussion
on both microfloral and megafloral evidence
bearing on this situation will be considered
later.

Earliest detailed estimates (Lesquereux,
1874, 1883, 1892) concerning the age of the
Dakota Sandstone Formation placed it as Ceno-
manian. This concept has been generally ad-
hered to since that time, although some geol-
gists have disagreed with it based chiefly upon
stratigraphy. In connection with this, Twen-
hofel (1920) assigned the Dakota to a Lower
Cretaceous age. Tester (1931) in studying the
type locality of the Dakota Sandstone consid-
ered it to be older than Ccnomanian and placed
Washita-Kiowa-Mentor-Dakota rocks of Kansas
at the base of the Cretaceous, with the Da-
kota being at least as old as the Mentor Forma-
tion of this region (midAlbian). However, the
conclusions of Twenhofel and Tester were
drawn based upon stratigraphic evidence alone
and are not substantiated by floristic evidence.

Cobban and Heeside (1952) assigned the
Dakota (Jroup to various ages ranging from
early Aptian to middle Cenomanian. However,
the lower members of this group (Lakota Sand-
stone and Fuson Shale of Darton, 1905) were
assigned to ages earlier than late Albian, and
the restricted Dakota Sandstone Formation was
considered by these workers to range from late
Albian through middle Cenomanian.

Berry (1920) was opposed to the detenni-
nation of Twenhofel (1920) tliat the Dakota
Sandstone Formation was of Lower Cretaceous
age. In this paper Berry cited floristic evidence

T'^<^arJ--

--^9

■ "7
I

KiG. 4. A. Roadcut exposure of Dakota Sandstone
Formation. The upper massive sandstone unit (A)
of this formation from this region rests upon the
middle shale-eoal-sand unit ( B ) . C dehmits an
unfossiliferous ash, and D represents the chief
fossiliferous ash seam. B. Contact Ix'tween Dakota
Sandstone and Cedar Mountain-Morrison Formation.
The Dakota forms tlie slope and ledge ( A ) which
rests upon the slope of tlie Cedar Mountain-Mor-
rison Formation ( B ) .

that the Dakota was of Upper Cretaceous (Ceno-
manian) age and could be correlated with the
Woodbine Formation of Texas. However, Berr\'
(1922) later considered the Woodbine Forma-
tion to be of Turonian age, although still to be
correlated with the Dakota Sandstone Forma-
tion. This determination was based upon a de-
tailed anahsis of the flora of the Woodbine
i'ormation from Lamar Count\', Texas, where-
in it was noted bv Berry that this flora had
three species in common with Turonian floras
of Europe.

MacNeal (1956) restudied the Woodbine
flora from collections made in Denton County,
Texas. Based upon examination of more and
better material. MacNeal disagreed with the
age determination of Berry, and a.ssigned the

HioLOGiCAL Series, Vol. 14, No. 3 Dakota Sandstone Flora

Woodbine flora to a Cenomanian age. MacNeal
did, however, agree with Berr\- (1920) that the
Woodbine flora is closeh- related to the flora
of the Dakota Sandstone.

A Cenomanian age for the Woodbine was
also agreed upon b\' Stephenson (1952) based
upon a stud\' of the invertibrate fauna from this
formation.

Brown (1952) agreed with Berry (1920) that
the Woodbine flora is verv similar to the flora
of the Dakota Sandstone of Kansas. One appar-
ent difference, however, was the conspicuous
absence of g\mnospermous species in the
Woodbine flora. This difference was elimin-
ated by MacNeal (1958) in his Woodbine studies
and recenth' b\' Hedlund (1966) in his palv-
nological studies on the Red Branch Member
of the Woodbine.

Hedlund (1966) also confirmed the Ceno-
manian age of the Woodbine Formation. In
this paper, Hedlund treated manv species of
fern and g\innosperm palvnomorphs indicating
that this segment of the flora, even though
poorlv represented by macrofossils, nonetheless,
represents an important floristic component of
the Woodbine.

Pierce (1961), Hall (196.3), and Agasie (1967)
in reporting on microflora! assemblages from
the Dakota Sandstone Formation of Minnesota,
Iowa, and Arizona, respectively, assigned this
formation to a Cenomanian age. The present
author agrees with this Cenomanian assignment
of the Dakota Sandstone Formation, although
in part this formation mav be of late Albian
age (Cobban and Reeside, 1952).

A CONCEPT OF A DAKOTA FLORA

It was recognized from earlier plant col-
lections from the Dakota Sandstone Formation
that plants from this formation were rather
closely related to plants growing on the earth
at the present time. Fossils from these collec-
tions were representatives of dicot\'ledonous
families and genera most of which are still
represented in the earth's flora. As Meek and
Hayden (1859) commented, Dakota fossils "be-
long to a higher and more modern tvpe of
dicotyledonous trees. . . ."

As more collections were made from this
formation throughout the midwestern United
States from such diverse localities as Kansas,
Nebraska, North Dakota, South Dakota, Wyo-
ming, New Mexico, etc., a floristic picture of
the Dakota began to evolve. This picture was
essentially that of Meek and Hayden (1859), and
later of Lesr^uereux (1874, 1883), that the flora
of the Dakota was essentallv a modern tvpe of
flora without many fossils representing ante-
cedent floristic types. As mentioned by Les-
quereux (1874) only one specimen collected to
that time could be referred to the Cycadophyta
{PterojiJiijUtim (?) lunjdenii), and even this ref-
erence was doubtful. Conifers were rare from
these early studies, both in number of species
and number of specimens, and represented a
very limited portion of the flora. Likewise, ferns
were few in number and appeared to make up
a ver\- limited part of the total Dakota vegeta-
tion.

On the other hand, even from earliest
studies, it was apparent that plant fossils refer-

able to the modern angiospermous vegetational
type were exceedinglv common and comprised
by far the dominant part of Dakota vegetation.
Indeed, Lesquereux (1874) noted that dicot
species represented in the Dakota floras could
be referred to "genera to which belong most
of the living arborescent plants of this country
(North America) and of our present climate."

When these facts were considered bv Les-
quereux (1874), he wrote a section dealing with
the disconnection of Dakota floras with ante-
cedent types. In this chapter, Lescjuereux wrote:

... it is evident that the flora of the Dakota
Group is as widely dLsconnected from that of
the Jurassic, even of the Lower Cretaceous,
or as distinctly original, as are the flora of
the Carboniferous compared to that of the
Devonian, or the Permian types compared to
those of the Cretaceous.

Later studies (Lestjuereux, 1883, 1892) indi-
cated that although ferns and gymnosperms
were more common than indicated bv earliest
studies, these floral types were still present in
limited numbers. Thus, Lesquereux (1892) in
his final work on the flora of the Dakota Group
summarized the relative importance of the dif-
ferent components of this flora. Of a total of
460 species identified from the Dakota Sand-
stone flora, Lescjuereux recognized six species
of ferns, representing 1.3 percent of the total
flora; twelve cycadean species representing 2.6
percent; fifteen species of conifers represent-
ing 3.5 percent; and 437 species of angiosperms
representing 92.6 percent of the total flora.

8

BmciiAM Voi'Nc: Univkhsitv Science Bulletin

In the liglit ot the above discussion, it may
be summarized that the prevailing early con-
cept of the flora of the Dakota Sandstone For-
mation was that it was essentially a modern
flora composed principalK of dicotyledonous
leaves related to modern arborescent genera.
The apparent paucity of antecedent floral t\pes
(Jurassic-VVealden ferns and gyinnosperms )
common in earlier Mesozoic floras is note-
worth\', although somewhat enigmatic to some
earh' paleobotanists. It must be remembered,
however, that this concept grew out of studies
by Lesquereux and others of macrofossils from
the midwestern portion of North America and
was drawn without the aid of data from two
sources to be added later. These two sources
are microfloral anahsis of the Dakota, and
megafloral analysis of this formation from many
new localities, especially those further west
than original collection sites.

As more data accumulate concerning the
Dakota Sandstone Fonnation, particularly data
dealing with regions west of the Rocky Moun-
tains, it becomes clear that a valid concept
of a Dakota flora must be modified some-
what from early views. While this earlv con-
cept is essentialh' for megafloral assemblages
from that portion of this fonnation east of the
Rocky Mountains, it does not fit well for the
formation as a whole. In this respect, it is sig-
nificant that the incidence of species repre-
sentative of the antecedent Jurassic-Wealden
vegetational type, particularh' Middle Meso-
zoic ferns, is much higher in megafloral assem-
blages from several localities in the Dakota
Sandstone Formation west of the Rocky Moun-
tains. That is not to say that dicotyledonous
species are not present at these localities. The\-
are not only present, but they are directly re-
lated to the dicot\ledonous species from this
formation further to the east. Indeed, at some
of these collection sites, angiospemious fossils
comprise the dominant floristic component of
the fossil assemblage. H()we\er, from man\'
localities in the Dakota Sandstone Formation
in Utah, Colorado, and Arizona, fossils repre-
sentative of the older antecedent vegetational
type are relativel\- more important than their
counterparts east of the Rockies, both in num-
ber of species and number of specimens pre-
sent. Indeed, at some western localtities, repre-
sentatives of this ancient floral l\pe dominate
the floral assemblage.

The Dakota flora from Montrose County,
Colorado, represents an admixtur(> of repre-
sentati\'es of the Jurassic-Wealden vegetational
type with modern angios|)ermous forms. Farl\-

work by Cockerell (1916) and Berr%- (1919a) in-
dicate the presence of matoniaceous ferns from
this region. Ferns referal^le to this famih' are
representative of an earlier flora! t\pe, as they
are common in the European Wealden and
other Middle Mesozoic strata. Matoniaceous
ferns are uncommon from rocks younger than
Earh' Cenomanian, and are unknown from
Tertiarx' strata. Brown (1950) made further
studies on tlu' plants from several localities in
the Dakota of Montrose County. This author
also treated matoniaceous ferns, as well as ferns
of the famih' Gleicheniaceae, which are also
indicative of a preangiospermous floral type.
Several other ferns from the Dakota of this
region were also chscussed b\' Brown. The
author has had occasion to study the Dakota
collections from Montrose County reported by
Brown. It is apparent that the fern families
Matoniaceae and Gleicheniaceae were very well
developed in this region during Dakota times.
Indeed, ferns of these two families are so com-
mon that they make up the predominant part
of the fossil vegetation, although they are col-
lected in association with angiospemious forms
which, as mentioned bv Brown (1950),

. . . include chiefly species that occur in the
large flora described by Les<]uereu.\, New-
berry and others from the brownish Dakota
•Sandstone of Kansas and Nebraska, in the
Woodliine Formation of Texas, the Dakota of
the Black Hills, South Dakota, and the Upper
Cretaceous rocks of Greenland.

Therefore, during Dakota times in southwest-
ern Colorado, the vegetation was composed of
typical Dakota angiospemious species, but with
a large proportion of plants representative of
a more ancient floristic type.

A somewhat similar situation existed in
Grand County, Utah, during Dakota times. Sim-
ilar to the fossils from Colaroda, the Dakota
flora from this part of Utah is comparable to
Dakota floras from other localities. However,
it is both significant and apparent that the
flora from the Grand Count\ Dakota Sand-
stone Formation is ver\' much dominated b\'
ferns representative of the older Jurassic-
W'ealden vegetational t\pe. Indeed, it is al-
most impossible at some localities in Grand
C'ounty to crack open a rock without exposing
at least one specimen of a fern representative
of the families Matoniaceae or Gleicheniaceae.
Dicf)tyledonous species are reduced both in
number of species and number of specimens
present. Gxmnosperms represent a rather minor
part ot the flora from this region as far as
number of species present, although quanti-
tatively they are represented b\- large amoimts

Biological Series, Vol. 14, No. 3 D.\kot.\ S.\Nn.sTONE Floh.\

of .silicified wood. Therefore, the Grand Count)
Dakota flora mav be characterized a.s a fern-
dominated, fern-angiosperin-g\inno,sperm alh-
ance, .similar to what could be expected during
a time which directly preceded the much dis-
cussed "population explosion" of angiosperms
during Cenomanian times ( Seward, 1927 ) .

This same vegetational aspect has been ob-
served b\' the author from several other Da-
kota localities of the western United States.
Floras from Longhouse \'allev and Kayenta,
Na\ajo Countv, Arizona, both exhibit character-
istics discussed above in connection with Colo-
rado and Utah Dakota floras. That is, ferns re-
presentative of the old Jurassic-Wealden vege-
tational t\pe dominated the Dakota vegetation
of these regions, and angiosperms represented
a rather limited vegetational type.

The Dakota Sandstone Formation from Coal
Can\on, Coconio Countv, Arizona, as discussed
bv Agasie (1967) exhibits similar floristic char-
acteristics. Microfloral analysis from this region
indicates that Dakota vegetation was composed
of a fern-angiosperm alliance with a minor
g\ mnospermous component. Fern spores repre-
sent the dominant microfloral component, es-
pecialh' fomis related to the famlv Schizaceae.
This fern famil\- is often represented in pre-
angiospermous floras (Harris, 1961), and pre-
]iminar\' collection of megafossils from the area
also indicates the presence of manv ferns re-
lated to an old Jurassic-Wealden vegetational
tvpe.

In addition to the data presented above, it
has been shown by Pierce (1961) and Hall (1963)

from microfloral anahsis of two localities in
the Dakota from Minnesota and Iowa respec-
tivelv, that ferns and gsmnosperms represented
more important components of the eastern Da-
kota flora than indicated by the megafossil re-
cord. With respect to this. Pierce reported 24
species of pahnomorphs representative of an-
giospermous species, 36 species of gvmnosperm-
ous pahnomorphs, and approximately 20
species of fern pahnomorphs. Based on this,
Pierce postulated that angiospermous species
are not as important in this formation as indi-
cated bv the megafossil record. However, a
more precise conclusion would be that rather
than the angiosperms representing a less im-
])ortant floral component, the gvmnosperm-fern
floral component represents a more important
part of the flora than previously thought.

From the above discussion, a more accurate
concept of the flora of the Dakota Sandstone
Formation as a unit may be formed. In western
America, the Dakota flora is a fern-angiosperm
alliance with a relativeh' small gvmnospermous
component. At several localties west of the
Rockies, ferns dominate the megaflora as well
as the microflora. To the east, this flora
changes character somewhat, becoming an an-
giospenn-dominated flora with a fern and gym-
nosperm component. This fern-gymnosperm
component was originalh' thought to be of very
minor importance in the eastern Dakota, but
recent microfloral studies (Pierce, 1961) have
demonstrated that this component of Dakota
vegetation was more important than had been
previousK' supposed.

PALEOECOLOGY

The plants from the Dakota Sandstone For-
mation near Westwater are preserved in a light
tan ash la\er from five to ten inches in thick-
ness. This laver was deposited directly upon a
coal seam and is overlain b\' another. These
plants are extremely well preserved with manv
of the leaf compressions illustrating cuticle,
vascular tissue, and reproductive structures. In
addition, manv of these fossils are disposed in
the strata in such a manner as to indicate that
the plants were preserved in growth position,
suggesting that deposition of the ash was rapid.

The ash la\er is uniform in color and com-
position throughout, with the exception of a
one- to two-inch portion directK- beneath the
upper coal seam. Lateralh throughout the ash,
distribution of the plants is differentential.
Thus, within a local area, most of the plants are

of the same species, and laterally the species
change abruptlv within a distance of a very
few feet. In certain outcroppings of the ash,
dense mats of leaves are found within a rela-
tiveh' small area. These mats are normally
found in the lower portion of the ash, and if
one traces them lateralh', the\' appear to follow
definite channels indicating possible stream
channels.

Peels of the foliage of the channel areas
were prepared to aid in identif\ing the angio-
sperms. On one of these peels a well-preserved
diatom and several spores of either algal or
fungal origin were found, indicating the likeli-
hood that the depositional environment of
matted leaves in these areas was aijuatic.

More detailed study of the fossihferous ash
brought to light several other important factors.

10

BmcnAM VoiNx; Univeksity Science Bulletin

In all areas lu-ar Westwalcr wIhtc the plant-
beariiii; asli lias hi-en exainiiu'd, tht" contact be-
tween the ash and the nnclerl\ ing coal seam is
very sharp and easily distinguished (Fig. 5).
In addition, the lowemiost portion of the ash
contains large nnmbers of root and rhizoid-
like structures, often in such profusion as to
form dense mats of these structures (Fig. 6).
Leaves are scarce in the lower portion of the
ash with the exception of poorly preserved,
often broken specimens. One of the more im-
portant identifiable fossils common in this region
of approximately one inch in thickness is the
rhizome of Equmtum lijeUi Mantel with at-
tached tubers.

Immediatelv above this rhizome region is a
horizon containing numerous well-preserved
leaf impressions. This region contains foliage
belonging almost exclusivelv to the three fern
genera Matotiklitiin, Astnilo])tcris, and Gleich-
enia. This zone is approximately five to eight
inches thick and is literally full of beautifully
preserved specimens.

At the upper portion of this dense foliage
region, the color and composition of the ash
changes somewhat (Fig. 5). The ash becomes
noticeabh darker in color and somewhat silty-
carbonaceous in composition. Many of the fos-
sils found in this region differ taxonomically
from those found lower in the ash. While the
three genera Matonklitim, Astraloi)t('ri.'i and
Gleichenia are still present as leaf impressions
in this region, thev are not as dominant as they
were in the lower portions of the ash.

Plant fossils in this upper region represent
several different genera. The greatest taxonomic
diversity in the Westwater flora occurs within
this zone. Significantly, Equisetuin rhizomes
reoccur in this upper region after being noticc-
ablv rare in collections from the middle portion
of the ash.

The contact between tlu' ash and the upper
coal seam is not as sharp as the lower contact
(Fig. 5). From the point where the ash be-
comes carbonaceous in nature and diversitv in
the flora appears, the ash laver graduallv be-
comes more carbonaceous until within a dis-
tance of one to two inches it grades into a well-
defined lignitic coal.

Several explanations for the above described
phenomena have been considered and rejected.
The sharp contact between the lower coal and
ash indicates that the ash fell directlv upon
a coal swamp and preserved the jilants that
were growing at the time. Further evidence in
supjiort of this is noted when it is recalled that
at the base of the ash a region rich in rootlike

Fig. 5. A. Collectins; .site e.xhibitiiig characteri.stic lith-
ology of Dakota Sandstone near We.stwater. The
upper massive sandstone unit forms the chff, and
tlie weathered shale-eoal-sand secjuence forms the
tahis slope in the foreground. B. Close-up of
fossiliferous a.sh seam illustrating rhizome region
( 1 ) , leaf region ( 2 ) , and di\'ersit\' region ( 3 ) .

structures, rhizomes, partially decomposed
leaves, stream channels, etc., occurs. Above this
is a region extremely rich in matoniaceous,
gleicheniaceous, and astralopteroid fossils which
appear to be in growth position. This evidence
would tend to support the hypothesis that the
ash was deposited rapidly upon a coal swamp
which was composed chiefly of ferns of the
three genera Matonidium, Gleichenia and Astni-
lopteris. If this indeed occurred, the rhizome
region would contain mostly rhizomes and roots
of these three genera.

The upper portion of tlie ash where the
diversitv appears contains plants which may rep-
resent forms that began to grow upon the
previously deposited ash. That is, the plants of
the upper part of the ash represent forms that
repopulated the swamp following ash deposi-
tion. Tlie likelv reason for the diversity in taxon-
omy of these plants is that they are forms

BioLOGicAi. Skbies, Vol. 14, No. 3 Dakoiw S.^ndsionk Floh.^

n

SAND

REGION OF
DIVERSITY

RHIZOME
REGION

Fk.. 0. I.^ Representative forms foiiiul in rt-f^ioii of diversity: Conioptcris (a); Asplcniutn (b); Equisctum (c);
and Ctmlophlchts (d). 2. Single specimen collected from leaf region illustrating Gfetc/icnifl and A/rttomV/iHni.
3. Rhizome and rootlike structures collected from rhizome region.

12

Bhiciiam Vouno University Science Bulletin

from serai communities in secondary succes-
sion. It is Iikel\- that the climax vegetation
growing in the coal swamp was a Matonidium-
Glcicheniu-Afitraloptchs association. Further evi-
dence in support of this conclusion was obtain-
ed b\' the author from stud\' of mascerations of
the lower and upper coals. Even though paly-
nomorphs from these coals were poorly pre-
served, it was evident that matoniaceous fern
spores were the dominant microfossils from
both, indicating that species of Matonklhim
were prevalent both prior to and subse(|iient to
ash deposition.

To test the above hypothesis, thin sections
of the ash were made from the three discussed
regions (rhizome region, leaf region, and diver-
sity region). The ash of the rhizome and leaf

regions shows no difference when examined
microscopicalK-. However, the ash of the region
of diversity is much more silty and carbo-
naceous than the ash of the two lower regions.
This evidence would tend to support the theory
that the jilants of the rhizome and leaf regions
were pri'siTved in one ash tall. The silty carbo-
naceous region (region of diversity) represents
a region of increased silt and carbon deposition
due to the initial deposition of enough soil to
support plant growth, population of this soil
b\' new plants (secondar\' succession), and sub-
sequent carbon deposition as these plants died
and were deposited. These plants probably rep-
resent stages in serai succession toward the
climax coal swamp vegetation of Matonidium,
Gleiclienia and Astmloptcris.

PALEOCLIMATE

It is sometimes possible, particularly when
working with late Mesozoic and Tertiary paleo-
floras, to reconstruct the presumed climate un-
der which these fossil floras existed. Two basic
metliods used in determination of paleoclimates
are the comparison of fossil genera to their
nearest living relatives existing under known
climatic conditions (extrapolation through uni-
formitarianisin), and the detennination of dicot-
yledonous leaf characteristics and comparison
of these characteristics with those found in
species growing in known climatic conditions.

It was first recognized b\' Bailey and Sin-
nott (1915) that leaves of arborescent dicot\le-
donous species growing in tropical regions ex-
hibit a higher incidence of entire margins tlian
do their counterparts growing under non-
tropical conditions. These authors pointed out
that there is a correlation between the number
of entire margins of arborescent dicot species
to climatic conditions under which these species
exist. Thus, from sevent\' to ninetx' percent of
tropical wood\' dicot species exhibit entire leaf
margins, whereas only twenty to forty percent
of wood\- dicots of cold temperature regions ex-
hibit entire margins. Further studies (Sinnott
and Baile\-, 1915; Chane\- and Sanborn, 19.33)
pointed out that large leaves, thick leaves, com-
pound leaves, and leaves with attenuated apices
are more abundant in tropical regions, decreas-
ing proportionately into temperate regions.

Chane\- (1954) applied these characteristics
in analw.ing 200 species of leaves from the
Dakota flora in an effort to determine the paleo-
climate of Dakota times. He noted that 73 per-
cent of the leaves from this flora exhibit entire

margins, 44 percent are over 10 cm long, and
74 percent are thick in texture. From these
data. Chancy concluded that the Dakota flora
lived under conditions "more subtropical than
temperate." Chancy further pointed out that
72 percent of Dakota leaves exhibit campto-
drome venation patterns which, according to
Chancy, also indicates that the climate during
Dakota times was tropical or subtropical in
nature. (>hanev suggested that this angiosperm
flora existed in a lowland forest, and that some
highland regions were to be found nearb\'.

Pierce (1961) discussed the Cretaceous cli-
mate of Minnesota based upon a fossil pol-
len assemblage from this region. He concluded
that the flora could be characterized as a coni-
ferous rain forest growing under warm tem-
perate climatic conditions similar to conditions
found ]iresently in the Pacific Northwest of the
United States or the coastal regions of eastern
Asia. This was based upon an unusualK- high
tre([uenc\' of g\mnospermous pollen encounter-
ed which. Pierce argued, could not have come
from nearby upland regions. Based upon this
information. Pierce agreed with Les(iuereux
(1S74) who suggested that the Dakota flora was
similar to the modern flora growing between
30° and 45° north latitude. However, Lesquer-
cux never assumed that the Dakota flora was
a coniferous forest, or that it even contained
a large number of g\mnosperms. Indeed, just
the opposite was the case as Lesquereux (1874)
discussed the noteworthy absence of g\mno-
spermous and fern species from the Dakota
flora. The work of Pierce, however, does point
out that both the g\ mnospennous and the fern

Biological Sehies, XOi.. 14, No. .3 Dakota SANDsroNK Flora

13

component of the Dakota flora are larger than
indicated h\- nicgafloral anahsis.

Hedlund (1966), in considering the pollen
flora from the Red Branch Member of the
Woodbine Formation which is closeh' related
to the flora from the Dakota Sandstone, com-
pared component members with their modern
counterparts and concluded that the climate of
this region during \\'oodbine times was warm
temperate to tropical. His determination was
based upon the high incidence of palynomorphs
related to modern plants which are presently
distributed chiefly in wami temperate-sub-
tropical regions.

A Dakota pollen flora from Coal Canvon,
Arizona, indicates subtropical to tropical cli-
matic conditions for this region during Dakota
times (Agasie, 1967). This is based upon rela-
tivel\' high frequenc\' of pahnomorphs from
genera related to extant taxa growing under
wet tropical or subtropical conditions. The rela-
tiveh' high frequenc\' of g\mnospermous pollen
present in this flora indicates that well-drained,
low upland regions existed near the lowland
swamp\' depositional basins.

Several species and hundreds of specimens
of Matonidium are present in the Westwater
flora. These ferns are probablv the best cli-
matic indicators present in this flora. These fos-
sils are related to the fern famih- Matoniaceae
which is presently a monot\pic (Bower, 1923)
or ditypic (Holtman, 1947) family containing
the genus Matonio. This genus is extremely
limited geographically and climatically in the
present world's flora and grows only in the
humid tropical upland regions of the Malayan
Peninsula. As mentioned by Mahabale ( 19.54 )
matoniaceous ferns are among a select group of
ferns that are accurate climatic indicators.

Gleicheniaceous ferns are well represented in
the Westwater flora, although these fossils are
somewhat less yaluable than matoniaceous ferns
as climatic indicators. Of 80 extant species the
majority are of tropical distribution, although
some species extend well into southerK- lati-
tudes (Bower, 1923).

The fern famih' Dipteridaceae is also repre-

sented in the Grand Cx)unt\- flora. This family
is monotypic containing the genus Di])teris,
which is presently restricted to fiye species all
of which are found in the Indio-Malayan re-
gion of the world (Bower, 1923). Thus, this
famih' also may be cited as an excellent ex-
ample of a tropical or subtropical climatic in-
dicator.

In addition to the aboye evidence, most ob-
served angiospermous fossils exhibit entire leaf
margins and two species show long attenuated
apices (drip points) which are thought to func-
tion in facilitating the run off of excess mois-
ture. As mentioned previously, these character-
istics are found more frequently in tropical re-
gions, and thus, the Dakota angiosperms from
Westwater substantiate the conclusion that the
Westwater Dakota flora grew under subtrop-
ical to tropical conditions.

Topography at the time of deposition of the
shalc-coal-sand unit of the Dakota Sandstone
Formation of this region was likely a broad
swampy mudflat near the shore of the Ceno-
manian Sea. Pockets of dense vegetation com-
posed basically of ferns and Equisetum de-
veloped in some of these regions, often accumu-
lating enough volume, under proper conditions,
to comprise a large component of the coal de-
posits which occurred between sandy or shale
sequences. Well-drained highland areas existed
nearby with gvmnosperms and angiosperms
present. Streams ran from the upland regions
into the lowland swampy areas carrying leaves
and other debris in various stages of decom-
position. The vegetative deposits from these
streams (gymnosperm and angiosperm leaves,
and likely wood) probably formed a large por-
tion of the coals, with the remainder being
added from the ferns and associated plants of
the fern swamps.

Following a period of time when deposi-
tion occurred as outlined above, the shoreline
of the sea changed, and the shale-coal-sand se-
quence in the Dakota of the Westwater region
was overlan bv a massive sand sequence. Fol-
lowing this, the entire formation was overlain
by deposits from the Mancos Sea.

SYSTEMATICS

Division ARTHROPHYTA

Genus EQUISETUM L.

Harris (1961) discussed the usage of Equi-
setum and Equisetitcs for fossils exhibiting sim-
ilarities to the extant Equisetum. Harris elected

to use Equisetum since "No morphological dif-
ference has ever been proved between Equi-
setum ami Equiselites. . . ." This statement is
completely accurate, and furthermore, the fos-
sil and living forms exhibit striking morpho-
logical similarit). In view of this, the author

14

has tollowed Harris (J961) and others in using
Equisetiim for placement of fossils wliich ex-
hibit characteristics similar to those of the ex-
tant genus.

Equisetum hjelli Mantel

Figs. 7-3, 7-5; 15-5. 15-9.

1889 Equisetum manjhmdicum Fontainc-U.S.
Geol. Surv. Mon. 15, p". 65, Pi. 170, Fig. 8.

1893 Equisetum hjeUi Mantel, Dawson-Trans.
Roy. Soc. Can. v. 10, pt. 4, p. 83, text-Fig. 1.

1956 Equisetites lijeUi (Mantel) Ungcr, Bcll-
Geol. Surv. Can. Mem. 285, p. 76. PI. 28, Figs.
5-8.

DESCRiPTiON.-Rhizome horizontal, unbranched,
2-4 mm in diameter, smooth to slighth ribbed,
bearing short tuber-bearing branches at inter-
vals as close as 8 mm; tubers ovoid to globose,
commonh- wrinkled, subtended b\- leaflike
bracts, 13 mm long by 6 mm wide, borne in
pairs; upright stems unknown.

Occurrence.— Dakota Sandstone Formation near
Westwater, Grand Count)-, Utah.

Repository. -Brigham Young University, BYU
1807-1810.

Discussion.— Many specimens of Equisetum
have been collected from the Westwater flora.
These specimens may be easily identified by
their attached tubers, although positively identi-
fied stem material is absent. Specimens obtain-
ed from this flora are collected for the most
part from the lowermost and uppermost por-
tions of the ash.

It is rather difficult to assign this plant to
anv known species since no aerial stems have
been observed. However, rhizomes of Equise-
tum are not uncommon in the fossil record, and
the Westwater material rather closely resembles
some described specimens.

Fontaine (1889) described Equisetum vir-
ginicum based upon both aerial stem and rhi-
zome material. The aerial stem was highly
branched, and tubers (called imperfect buds by
Fontaine) were borne individually along the
rhizome. This specific name was used again
later b\- Fontaine (1899) for placement of a
specimen from the Black Hills Lower Cre-
taceous. Bell (1956) noted that in overall mor-
phologv, E. virginicum is very similar to E.
hjeUi Mantel.

Equisetum marijlaiulicum was described in
1889 bv Fontaine. The aerial stem of this

Bhic;h.'vm Vouno University Science Bulletin

species was highly branched, although a rhi-
zome (juestionably attributed to this species
(op. cit., PI. 179, Fig. 8) is very much similar
to rhizomes collected from Westwater.

Bell (1956) illustrated tuber-bearing rhi-
zomes of Equisetites hjelli (Mantel) Unger
(Equisetum hjelli Mantel). These rhizomes dif-
fer from the Westwater specimens by being
branched and exhibiting strongh' ribbed inter-
nodes. Other differences are the whorled dis-
position of tubers and the apparent lack of
bracts subtending tubers on the Canadian
specimens. Howe\er, the dimensions of the
Canadian material compare \sell with those
noted for Westwater specimens, although the
tubers are commonh' somewhat larger in the
latter.

Other citations of E. hjelli are incomparable
to the Utah specimens since onl\' aerial stems
are described. Thus, Schenk (1871) described
aerial stems of £. hjelli as being much branch-
ed, and with larger dimensions than would be
expected for aerial stems of the Westwater
Equisetum, and Fontaine (1889) noted that
characteristics of aerial stems of E. hjelli from
Virginia agreed closely with those of this species
from Europe as noted by Schenk (1871).

Division FILICOPHYTA
Genus ASPLENIUM L.

This genus is rather large and is well repre-
sented both in the modern and fossil flora.
Bower (1923) reported Asplenium to contain
429 extant species, and Jongmans (1957) listed
96 fossil species of this genus with three vari-
eties and listed ten as Asplenium sp. In addi-
tion, Asiilenites has been used for placement
of asplenioid ferns resembling the modem As-
plenittm; this genus contains several additional
species.

Most of the fossil species of Asplenium are
based upon sterile foliage, and undoubtedly
man\ arc s\nonvrns. This problem is further
complicated bv the fact that sterile foliage of
this genus rather closeh resembles foliage of
several other fern genera such as Anemia, Omj-
chiopsis. Sphenojiteris. AcrostirliO])leris. Coni-
opteris, and Dicksonia. There is no adeciuate
wa\' of determining between some species of
these genera and between species of As}>len-
ium, based upon sterile foliage alone. Clarifi-
cation of resulting taxonomic problems awaits
furthcT collection and study of existing speci-
mens.

Biological Skhies, \oi.. 14, \(

Dakota Sandstone Floha

15

Fic. 7. 1. Glcicheniu dcliciilutu Ht-t-r. Portions of pinnae illustrating pirniuk- shapt- and disposition. "(X12:5).
BVU 1829. 2. GleinchenUi dclicatula Heer. Portions of pinnae illustrating pinnule shape and disposition.
(X.3). BYU 1829. .3. Equisftum lijclli Mantel. Portion of rhizome illustrating shape and disposition of tub-
ers. (X.3). BYU 1809. 4. Gleicheniu dclicatula Heer. Pinna fragment illustrating characteristic branching.
(X3). BVU 18.30. .5. Equlsetum Itjclli Mantel. Portion of rhizome illustrating tubers with subtending bracts.
(X3). BYU 1810.

16

Bhk;ii\m ^()^ \(; Univkhsitv Sciknce Bui.lktin

Aspleiiitiiit (licksoiiianutu Ilet-r
Fi^s. 8-4; 11-3; 12-2; 15-4.

1874 As])Ieniuin dicksoniamim Hcht— Flor. Foss.
Arct., V. 3, pt. 2, p. 31, PI. 1, Fig.s. 1 (exd. b, c),
laa, 2-4, 5 (excl. a, b).

1899 As])leniuin (licksonininmi Ilccr, VVard-
U.S. Geol. Surv. 19tli .\iiii. Kept., pt. 2, p. 704,
PI. 170, Fig. 1.

1950 As])lcniiiin .sp. brown— U.S. Cool. Surv.
Prof. Paper 22 1 -D, p. 48, PI. 10, Fig. 4.

Descrh'tion.— ICntiri' leaf unknown, at lea.st bi-
pinnate; ultimate pinnae alternate, up to 90 mm
or more long bv 50 mm wide; pinnule.s lanceo-
late, entire to deeply cut, altcrnati', up to 40
mm long bv 9 mm wide; attachment ba.sal to
.single point; midvein .strong at point of origin,
not extending to pinnule apex, dividing; .second-
ary vein.s repeatedh' divided, free throughout;
fertile .specimen.s unknown.

OccuHRENCE.— Knowni from the Dakota Sand-
stone Formation near Hot Spring.s, South Da-
kota, near Naturita, Montrose C'ountv, Colo-
rado, and near Westwater, Grand County, Utah.

Repository.— Brigham Young University, BYU
1811-1814.

Discussion.— The author has elected to follow
Ward (1899) who collected an asplenioid fern
from the Dakota Sandstone of the Black Hills
region and pliiced it in A. dicksonianuiu. How-
ever, Brown ( 1950), in describing ;i simihu' fern
collected 1)V Stokes from the Dakota Sandstone
of Montrose (bounty, Colorado, noted that both
the Colorado and the South Dakota specimens
appeared to be somewhat smaller and more
delicate than typical specimens of A. dickson-
ianum. However, b;ised upon material in his
possession. Brown (1950) was hesitant to name
a new species of this genus and so ascribed the
material to AsjileiUuin sp.

The observ;itions of Brown were accurate
for some specimens ol Asj)lciututi from Colo-
rado. However, with the smaller forms men-
tioned bv Brown (1950) are inanv which are
larger and resemble rather closely A. dirkson-
kmuin. Westwater specimens of Asplenitini are
also like A. dicksonianuin. although positive
identity of this feni awaits collection o( speci-
mens illustrating soral and spor;ingiaI character-
istics.

Asplenitini dnkolcnsi.'i Hushfortli, sp. nov.

Kij^s. 11 -.5; 12-(i.

Descuii'tion. —Entire leaf unknown, bipinnate;
petiole smooth, 1-2 mm in di;imeter, bifurcat-

ing to give rise to two r;uhi; piniKie alternate,
35 mm or more long In 11 mm wide, lance
olate; pinnules ovoid, up to 9 mm long by 3.5
mm wide, alternate, attachment basal with
rounded sinuses, to nearly single point, apices
obtuse to rounded, margins entire; midvein
strong at point of origin, not extending to pin-
nule apex, dixiding; secondarx veins mosth' ob-
scure, free, extending to pinnule margin; fer-
tile specimens imknown.

Occurrence.— Known from the Dakota Sand-
stone Formation near Westwater, Grand f'oun-
t\', Utah.

Repository.— Brigham Young University, Iiolo-
t\pe; BYU 1806.'

Dificvssio\. —Asjilcnium dakotensis is proposed
for a small, rather delic;ite Asiilcniuni from the
Westwater flora. It differs from A. dick.wnian-
tini Heer from the s;ime locality by being much
smaller, both in pinnule and pinna size, and by
pinnules being entire and rather strap-shaped,
rather than dentate or deeph cut and lance-
olate as are pinnules of A. dicksonwnum (Table
1). A. dakotensis Rushforth is also much small-
er than most other species of Asjilcniuni and
differs from m;uiv by Ii;iving pinnules with en-
tire margins. It is also rather unique in its bi-
furcated petiole which gives rise to two rachi.
This last characteristic coupled with ]iinnule
shape and margination led the author to con-
sider this fern closely related to Knowltonella
Berr\' (1911). However, examination of the type
specimens of KnowltoncUa indicates that the
branching habit of the two differs, as does
pinnule shape. Fertile specimens of neither
Knowltonella nor A. dakotensis have been dis-
covered, and for the present, the author prefers
to assign this fern to the Aspleniaceae ;is As-
j)Icniutn dakotensis, rather than to the Maton-
iaceae imder KnoulfonclJa.

T;il)!e 1

Comparative M
I leer am

orplioloijN ol Asplcnutru die

1 AKplfiiitini ilitkotciisis Hiishfi

A. clickiimidnurii A.

ksDHUinum

irtli

Jiikolcnsin

Lent^th of

pinna

90 mm

3^ mm

Wiilth of

pinna

riO nmi

1 1 nun

I.<-ii<;lli of

Fininilc

■10 nun

9 nun

Widtli of
Pinnule

9 mm

3.5 mm

Pinnule shape

Lanceolate

Strap-.shi

iped

Pinnule margin

Entire to
deeply cut

Kntire

Bioi.ocicAi. Sehiks. \()1.. 14, No. 3 Dakota Sands ione Fi.oha

17

-r

L

4 ^^'^^ ^

^^^^ ^■,.^-

•r^S:!^'

4»"

Kir.

8 1 C.U-ichcnia drlicatulu Hver. Pinna fragment illustrating clmracteristic hranchuig (M.-). BVU 18-7 -
Ltoni^un hrownii var „,««>„■,„•„„,./»», Rushforth. Pinna fragment illustratn,g long, hnear pnmnk^. (XI 2
H Z- 3. Cleichcuia' dclicutula Heer. Pinna fragment illustrating pinna and p.nnule cbspos, k n

\1 ' ) BVU 1828 4 Asplcninm ,licksouu.num Heer. Pinna illnstrating nltmiate pnmae drspos.t.on. X .2
BVU 81 1 5 ChnlophkL pana Fontaine em, Berry. Pinna fragment illustrating pinnule chsposition. (X
BVU 184V 6 GlcichcnU, comptonuwjoliu (Deb. and Ett.) Heer. Pinna fragment . lustratmg eharaeter.sfc
l,ranel'ing:'(X1.2) BVU 1817. 7. Asplcmum dick.somanum Heer. Pinna fragment dlustratmg disposition of
ultimate pinnae. (X1.2). BVU 1812.

18

BiutaiAM VouNc Univehsity Science Bulletin

Family 1:)K:KS0NIACEAE

Genus CONIOPTERIS Brongniart

Coniopleris is probably the best known fos-
sil genus of the fern family Dicksoniaceae. As
listed bv Jongmans (1959) this genus contains
approximateh- 45 species and three varieties.
However, as discussed by Harris (1961) this
genus is in need of revision, and fewer than 45
species are likely to be valid.

Conioptcris uestwaterensis Rushforth, sp. nov.

Fig.s. 10-2; 11-4; 13-5.

Description.— Entire leaf unknown, at least bi-
pinnate, some divided into a sterile portion be-
low and fertile portion above; sterile pinnae
lanceoiale, up to 20 mm wide by greater than
60 mm long; pinnules suboppositc to alternate,
up to 12 mm long b\' 3 mm wide, lanceolate,
margins entire near pinna apex to strongly den-
tate further down the pinna, apices generally
acute but range to somewhat obtuse, attach-
ment basal to constricted basal, sinuses round-
ed (pinnules connected along rachis); midvein
prominent; secondary veins mosth' obscure; fer-
tile foliage consists of sterile pinnae below and
fertile pinnae above; fertile portion of frond up
to 65 mm long or longer above highest sterile
pinna; fertile pinnules highK' reduced, alter-
nate, connected along the rachis; venation ob-
scure; sori, sporangia and spores not observed.

OccuHHENCE.— Known from the Dakota Sand-
stone Formation near Westwater, Grand Coun-
ty, Utah.

Repository.— Brigham Young Universitv, holo-
type: BYU 180,3.

Discussion.— This fern is known from several
specimens collected from the Westwater local-
tv. A few of these are rather complete, illus-
trating fertile as well as sterile foliage. How-
ever, even though fertile specimens have been
collected none illustrate sori or sporangia.

C. westwaterensis is similar in some respects
to several other species of Coniojiteris. It per-
haps approaches most closely C. I>ttiejen.sis
(Zalcssk\) Seward which is a common species
from the Mid Jurassic of Asia and has also
been collected from Alaska and Europe. C.
westwaterensis differs from C. burejcnsis (Zales-
skv) Scuard in having less crowded pinnae and
bv having the fertile portion of the foliage
separated from the sterile portion, often oc-
curring as much reduced pinnae apicalK' to
sterile pinnae.

C. bella Harris, which is similar to C. bure-
jensis (Zalessky) Seward also is similar to C.
westwaterensis. C. bella is only separated from
C. burejensis based on pinnule margination
and fine soral details, and C uestwaterensis
differs from this fern as from C. Imrejensis. C.
westwaterensis also differs from both of these
ferns in that it exhibits secondary pinnae which
are consistenth' larger, being up to 20 mm wide,
whereas the pinnae of C. burejensis and C. bella
are typically 10 mm wide.

Family DIPTERIDACEAE

Genus HAUSMANNIA Dunker

Dunker (1846) proposed this genus for ster-
ile leaf fragments collected from the Wealden
of Germany. These fragments were divided by
nearlv ecjual ditchotomies to form strap-shaped
leaf segments. Subsequent to this time, several
other specimens of this genus have been col-
lected, and a rather adeejuate generic diagnosis
has evolved (Harris, 1961).

Ilatisuiannia rii^ida Newberry

¥\gs. 9-2, 9-3; 13-4.

1895 llausmunnia rigida Newberrv— U.S. Geol.
Surv. Mon. 27, p. .35, PI. 1, Figs. 2, .3', 5.

1910 Newberri/ana rigida Berry— Jour. Geol., v.

18, p. 254.

1911 Neivbern/anu ri<^ida Hcri-y— Geol. Surv.
New Jersey Bull. 3, p. 220.

Desciui'tion.— Entire leaf unknown, divided to
form tongue-shaped pinnules; pinnules typical-
l\- 9 mm long b\- 3 mm \\k\v. apices rounded,
margins entire, attachment basal with rounded
sinuses; midrib prominent, apparently not ex-
tending to pinnule apex; secondary veins ob-
scure if present; fertile specimens unknown.

OcciiRHENCE.— Known from near Westwater,
Ciriuul (bounty, Utah.

Repository.- Brigham Young Universitv, BYU
1815.

Discussion.— The botanical affinity of llausnuin-
nid luis been in (juestion in the past. Some early
workers considered this plant to be ;i hepatic.
Newberry (1885) said in this connection:

1 li;i\c bffii It'll to think it possible it was a
liiUhtT kiiul of hepiitic-, a Marchanliu. lor cx-
amplf, lifted from its cr<-i-piiig (.oiulitioii into
an iiulfpt-nilent and erect phint, trained and
disciplined into syininetr\ by the occult in-

Biol<k;i(:al Skhies, \'<)i.. 14, No. 3 Dakota Sandmone Floha

19

'!•:

Ay;

y

i' ^ -^^^^

Fic. 9. 1. Astradopteris colurudicu (Brown) Reveal, Tidwell, and Ruslifortli. Near apical pinna fragment. (X1.2).
BYU 1839. 2. Ilansmannia rigidci Newbern,'. Near apical pinna fragment illustrating tongue-sliaped pin-
nules. (X1.2). BYU 1815. .3. llausmunniu rigidu Newberry. Enlargement of Fig. 2. (X3). BYU 181.5. 4.
Gleichcnia delictitula Ilccr. Piima fragment illustrating disposition of ultimate pinnae. (X1.2). BYU 1826.
5. Cladopldchis crenuta Fontaine. Pinnule fragment illustrating venation. (X3). BYU 1841. 6. Gleichcnia
comptoniucfolia (Deb. and Ett. ) Heer. Pinna fragment illustrating disposition of ultimate pinnae. (XI. 2).
BYU 1816. 7. Astmlnptcris roloriidica (Brown) Rexcal, ■I'iducll, and Rushforth. Pinnae fragment. (X1.2).
BYU 1840.

20

BmcuAM YoiNc; University Science Bulletin

fluencc uhicli lias given siuli grace and ex-
actness to the foliage of fenis, lycopods, and
some conifers.

Later studies have shown tluit Ihtusiiumnia
is ;i fern of the family Dipteridaceae ( Richter,
1906), although fertile speeiiuens demonstrat-
in<4 soral characteristics are still unknown. The
Westwater llausmannki doc>s not shed any light
on this problem since fertile specimens are un-
known.

llmismannui was first treated in America by
Newberry (1S85) who described //. rigida. This
species was proposed for several plant fragments
collected from the Amboy Clays of New Jersey.

Berry ( I9I0) later transferred llausmannia
rigida to the new genus Newbern/ana without
description, discussion, or illustration, and
hence, his reasons for this move were unknown.
However, Berry (1911, p. 220-221) further
discussed Haiisni'innia rigida and stated that
Newberry had referred this plant to the Hep-
aticae, and due to this:

It is, obviously, not related to the genus llaiis-
mitnnia, Dunker, which has been definitelv
proven to be a fern genus of the familv Dip-
teridaceae, so that I have selected to propose
a new generic name. . . .

However, Berry gave no reasons why this plant
should not be considered to be related to Haus-
nuinnia, and further went on to say that indeed
it is probably a fern. It is the opinion of the
present author that Newberry was justified in
placing his material in llausmannia as a new
species since this material agrees rather well
with the generic diagnosis of JIausiminniu. Fur-
ther collections and reevaluation of Newberry's
material should, however, be undertaken.

//. rigida Newberry ( 1895, p. 35, PI. 1, Figs.
2, 3, 5.) is probably most closely related to the
Westwater llausmannia. This species lacks di-
cholomous leal ilivisions, and more regular divi-
sions of the frond-forming tongue-shaped seg-
ments is also found in the Westwater species.
As discussed by Nevvberr\- (1895), the midrib
in //. rigida gives rise to numerous fine secon-
dary veins, although his illustrations do not dem-
onstrate this feature. These secondarv veins
cannot be observed in the s[)ecimen from West-
water.

//. dicliotonui Dunker is a common species
of this genus from the Jurassic and Lower Cre-
taceous of Europe. This species usually has
been well described and, as mentioned by Neic-
berry (1895), is closely related to llausmannia
rigida. II. rigida differs from //. dichotoma in

tiiat tiie divisions of the leaf are not dichoto-
mous, and are more regular in the former.

Family GLEICHENIACEAE
Ceims GLEICHENIA Smith

Berry (1924) discussed the usage of Glei-
cJienites and Glciclicnia. In this paper, Berry
rather stronglv supported the use of Glcichcnia
rather than CAcichenitcs and stated in this re-
gard, "It is surelv nothing but a mental illusion
to imagine that the iLse of an objectionable
tenn like Clciclicnitcs indicates a conservation
of judgment." However, Seward ( 1927 ) dis-
agreed with Berry and used the name Gleichen-
itcs for placement of several Greenland species.
Seward stated that in his opinion it is a sound
practice to use an ite.s ending for a generic
name when "either our ignorance or the occur-
rence of some character in which a fossil species
differs from anv existing type" may apply to an
extinct taxon.

Harris (1961) pointed out that the practice
of using genera with an ites ending is being
dropped bv most paleobotanists in recent times.
Harris, however, rightfully suggested that when
a question exists as to whether a fossil taxon may
with ecjual validity be placed with more than
one living taxa, a different generic name for
the fossil must be used.

It is my opinion that it is sound to use the
ites ending under certain circumstances when
evidence of the sameness of a fossil and living
genus is suggested but not definite. However,
the use of this ending merely because of the
older age of a fossil is not valid. When there
is little or no (juestion that a fossill may be
placed in an extant genus, it is both acceptable
and desirable to use the modem generic name
for its placement.

The fossil section of the genus Glcichcnia
is in urgent need of revision and reevaluation.
Many species are undoubtedlv svnonyms and
could be more correctly placed. In addition,
this genus afford.s an excellent opportunitv to
follow the cNolution and migration of a tern
genus from Paleozoic to recent times, and its
study could contribute greatly to botanical
knowledge.

Glcichcnia coniptoniaefolia ( Deb. and
Ett.) Heer

Fig.s. 8-6; lO-I, 10-6; I2-I, 12-3; 13-1.

13-3; 14-3; 20;i.
1859 Didt/tnosarus coniptoniaefolia Deb. and

Bi()L()(:ic:al Skuies, Vol. 14, No. 3 Oakoia Sandsionk I'i.oha

21

•^^"'x..

'*^%if5'''

Fig. 10. 1. Gleichenia comptoniuefoliu (Del), and Ett. ) Heer. Pinna fragment illustratiiif; disposition of ultimate
pinnae. (XI. 2). BYU 1821. 2. Conioptehs wcstwatercnsis Rushfortli, sp. nov. Pinna fragment illustrating dis-
|X>siti()n of fertile pinnae. (X1.2). Paratope BYU 1804. .3. Matonklium hrownii Hirsliforth. Pinna fragment il-
lustrating pinnule disposition. Tiiis fragment is .somewhat atypieal and resembles some speeies of Sclenocar-
/)i/.s- Selienk. (XI. 2). BYU 1833. 4. Gleiclwnki dcUcntula Ileer. Pinna fragment illustrating ultimate pinnae
ilis|«>sition and pinnule shape. (X1.6). BYU 1829. .5. Miitonklium hrownii Hushforth. Pinna fragments and
fiddle head. (X1.2). BYU 1834. 6. Glciclwnia coviptomacjolm (Deb. and Ett.) Heer. Pinna fragment illus-
trating ultimate pinnae disposition and pinnule shape. (XI. 2). BYU 1822.

Bhicham Vounc Univehsity Science Bulletin

Ett.-Denksch. K. Akad. Wiss. Wicn., 17, PI. 1,
Figs. 1-5.

1868 Gleichenia gieseckiarui Heer, pro partt-—
Flor. Foss. Arct., v. 1, p. 78, Pis. 43, Figs, la,
2a, 3a; 44, Figs. 2-3.

1868 Pecopteris borealis Brong., Heer, pro parte
-Flor. Foss. Arct., v. 1, p. 81, PI. 44, Fig. 5.

1868 Gleichenia zippei (Corda) Heer-Flor.
Foss. Arct., V. 1, p. 79, PI. 43, Fig. 4.

1868 Pecopteris arctica Heer, pro parte— Flor.
Foss. Arct., V. 1, p. 80, PI. 43, Fig. 5.

1868 Pecopteris rinkiiina Heer— Flor. Foss.
Arct., V. 1, p. 81, PI. 44, Fig. 4.

1874 Gleichenia longipennis Heer, pro parte—
Flor. Foss. Arct., v. 3, pt. 2, p. 46, PI. 6, Figs.
4-6.

1874 Gleichenia tliulensls Heer— Flor. Foss.
Arct., V. 3. pt. 2, p. 4S, Pis. 5, Fig. 9b; 10,
Fig. 18.

1874 Gleichenia comptoniaejolia ( Deb. and Ett. )
Heer, pro parte— Flor. Foss. Arct., v. 3, pt. 2,
p. 49, PI. 11, Figs. 1-2.

1874 Gleichenia gracilis, Heer, pro parte— Flor.
Foss. Arct., V. 3, pt. 2, p. 52, PI. 10, Fig. 5.

1874 Gleichenia gieseckiana Heer— Flor. Foss.
Arct., V. 3, pt. 2, p. 43, Pis. .3, Id, 8; 7, Fig. 1.

1874 Gleichenia nauckhoffi Heer— Flor. Foss.
Arct., V. 3, pt. 2, PI. 25. Fig. 4.

1874 Gleichenia zippei (Corda) Heer, pro
parte-Flor. Foss. Arct., v. 3, pt. 2, p. 44, 90, 97,
Pis. 4-7; 25, Figs. 1-3.

1874 Pecopteris hijperborea Heer— Flor. Foss.
Arct., V. 3., pt. 2, p. 41.

1874 Gleichenia kurriana Heer, Lescjuereux-
U.S. Geol. Surv. Terr. Kept., v. 6, p. 47, PI. 1,
Figs. 5, 5b, .5c.

1882 Gleichenia geiseckiana Heer— Flor. Foss.
Arct., V. 6, pt. 2, p. 6. PI. 2, Fig. 9.

1882 Gleichenia zijipei ( Corda ) Heer- Flor.
Foss. Arct., V. 6. pt. 2, p. 7, PI. 3, Fig. 2.

1882 Gleichenia longipennis Heer— Flor. Foss.
Arct., V. 6, pt. 2, p. 7, PI. 2, Fig. 5.

1882 Gleichenia comjHoniaefolia (Deb. and
Ett.) Heer-Flor. Foss. Arct., v. 6, pt. 2, p. 8,
36, PI. 46, Fig. 25.

1883 Gleichenia norclenskioldi Heer, Lesquereux
-U.S. Geol. Surv. of Terr. Repts., v. 8, p. 26,
PI. 1, Figs. 1, la.

1889 Gleichenia nordenskioldi Heer, Fontaine

-U.S. Geol. Surv. Mon. 15, p. 119, PI. 21, Fig.
11.

1889 Gleichenia zippei ( Corda ) Heer, Fontaine
-in Ward, U.S. Geol. Surv. 19th Ann. Kept., p.
664, PI. 162, Fig. 9.

1927 Gleichenites gieseckiana (Heer) Seward—
Phil. Trans. Rov. Soc. London B, v. 215, p. 69,
PI. 5, Figs. 1-4,' 6-14, 16, 17; PI. 10, Fig. 96; PI.
12, Fig. 118; text-Fig. 2.

1950 Gleichenia kurriana Heer, Brown-U.S.
Geol. Surv. Prot. Paper 221-D, p. 48, PI. 10,
Figs. 5-6.

1956 Gleichenia gieseckiaiui (Heer) Seward,
Bell-Geol. Surv. Can. Mem. 28>5, p. 61, PI. 17,
Figs. 1, 2; PI. 18, Fig. 5.

Deschiption. —Entire leal unknown, at least bi-
pinnate; rachis stout, slightly greater than 1 nun
in diameter, smooth, appearing dichotomously
branched (although not truly dichotomizing
[Bower, 1926] ), with a bud in the axil of dichot-
omy; ultimate pinnae alternate, occasionally sub-
opposite, nearly linear, approximately 80 mm
long by 10 mm wide; pinnules arising obtusely
from rachis, narrow eliptic to deltoid near pinna
apex, oriented slightly toward pinna apex, rang-
ing from 1 mm wide by 2 mm long at apex of
pinna to 3 mm wide by 6 mm long near pinna
base, margins entire, apices obtuse; major vein
strong at point of origin, not extending to pin-
nule apex; secondary veins numerous, dividing,
extending to pinnule margins; sori commonly
three per pinnule, round; sporangia and spores
unknown.

OccuRKENCE.— Near Westwater, Grand County,
Utah, and Rabbit Valley, Grand County, Utah.
Rei'ositoky.— Brigham Young University, BYU
1816-1825, 1853a^

Discussion.— As may be deduced from the syn-
onomv for G. comptoniac folia, Heer proposed
many species of Gleichenia in the late ISOOs
based upon material collected from several
Greenland localities. Heer treated any fonn that
illustrated any variation whatsoever as a new
species. Seward (1927) reviewed the work of
Heer and studied many of his original collec-
tions. Based upon his studies, Seward concluded
that many of the species proposed by Heer mere-
ly represent variations of a few species.

In discussing the synonomy tor C;;. giesecki-
ana, into which Seward ( 1927 ) lumped many of
Heer's ( I86S, 1S74, 1872) species, Seward men-
tions two names which predate G. gieseckiana.
The first of these Pecopteris borealis Brongniart,
was applied by Brongniart (1828). Seward ex-

Biological Series, Vol. 14, No. 3 Dakot.\ Samwionk Floha

23

Fig. U. 1. Cladophlebis constricta Fontaine em. Bern'. Pinnules illustrating deeply lobed margins. (X3). BYU
1941. 2. Clmlophlehis puna Fontaine em. Berr\'. Pinna fragment illustrating pinnule shape and venation.
(X3). BYU 1842. 3. Asplenium dichsnnkmum Heer. Pinna fragment illustrating shape and venation of pin-
nules. (X3). BYU 1814. 4. Coniopteris iLcstwtiteremis Rushforth, sp. nov. Portion of pinna illustrating
pinnule shape and margination. (X3). Paratype; BYU 1805. 5. Aiplcniurn Jakotcnsis Rushforth, sp. nov.
Portion of frond illustration pinnae and pinnule disposition. (X3). Holotype: BYU 1806.

Bni(;nAM ^ OuNo Univehsitv Science Bulletin

aniined the type specimen of this fern and con-
cluded that it was not possible to detennine if it
is synononioiis with G. <iies('<ki(ina. Later collec-
tions from the probable locality where P.horealis
WAS collected have turned up specimens which
iire identical with C. gieseckiana. However,
since the original type specimen is not identifi-
able, Seward chose to retain Heer's specific epi-
Didtjmosarus comptoniaejolius Deb. and Ett.
that.

( 1859) is also an older name which has been ap-
plied to C. gk'seckiuno Heer. Heer recombined
D. comptonkiefoliiis Deb. and Ett. as GlwcJienia
gieseckkimi since the fonner is undoubtedly
gieicheniaceous. Seward (1927) placed C comp-
toniaefoUa ( Deb. and Ett. ) Heer in synonomy
with G. gieseckkimi Heer. Seward did not use the
oldest specific epithat l>ecause "evidence of iden-
tity is not convincing." However, Seward m the
same pajx^r noted that some of the feni.s Heer
had placed in G. coniptonwefoUu were.without
question, referable to G. gwseckkma. Further-
more, the illustrations of Debey and Ettinghau-
sen ( 18^59) indicate that it is very likely that the
ferns referred by them to Didtjmosarus comp-
toniaejolius are identical to G. gieseckkina. In
view of this, it is preferable to use the oldest
specific epithat, and therefore the most accept-
able binomial applied to G. gieseckiana of Heer,
is G. comptoniaefolki ( Deb. and Ett. ) Heer.

Lesquereux (1874) figured a gleichenious
fern from the Dakota Group of Kansas. These
fenis agree in ever\' detail with Glek'henia
comptoniaefolki ( Deb. and Ett. ) Heer, but were
placed in Glewhenki kurrkiiui Heer by Les(|uer-
eux. However, it is important to note that Les-
(juereux considered the Dakota ferns closely al-
lied to Duit/mososorus comptoniifoJius Deb. and
Ett., and in fact noted that in pinnule shape,
disposition, and venation, "our American species
or variety would be more closely related to
Dklymososonis comptoniifoUus Deb. and Ett. . ."
Les(|uereux ( 188.3) again treated gliechenious
ferns in his work on the Cretaceous and Tertiary
of the Western Territories. Pinna fragments of
Gleichenia from the Dakota Croup were placed
by Les(|uereux in G. nordcn.skwldi. However, as
noted bv Seward ( 1927 ) the feni examined by
Lesquereux agrees in all respects with G. giescck-
iana (G. comptoniacjolia) .

Fontaine (1889) discussed Glewhemu nord-
enskioJdl Heer as known from the Fotamac
Croup. However, this fern agrees with G. comp-
tonkicfolki closely, and should be placed in this
species.

In discussing Lower Cretaceous plants from
the Black Hills region, Fontaine (1899) again

discussed gieicheniaceous ferns. He placed a
specimen of Glek'henia in G. zippei (Corda)
Heer. This feni likewise is verv similar to G.
eomptoniaefoJia, and can be correctly considered
in this species.

Velenovskv (1888) described and illustrated
several fossil ferns with gleichenious affinities
collected from the European Cretaceous. Two of
these which are very similar to G. comptoniae-
folici are G. zippei ( Corda ) Heer and G. imdtin-
ervosa Velen.

Brown ( 1950) reported on gleicheniou.s fos-
sils from the Dakota Sandstone of southwestern
Colorado. He placed these ferns in Gleichenia
kurriana following Lescjuereux (1874). Recent
examination of these ferns by the present author
indicates that they are indeed very similar to
those figured by Les(|uereux (1874) as G. kur-
rianu which are correctly placed with G. comp-
toiiidcfolia.

Bell (1956) described Gleichenites gieseck-
ianus Heer em. Seward from several localities in
western Canada. These ferns agree with the
Westwatcr Gleichenia in all respects except that
the pinnules on the Canadian specimens are often
somewhat reflexed. Again, these ferns are correct-
Iv classified as Gleichenia cornptoniaefolia ( Deb.
and Ett.) Heer.

Seward ( 1927 ) discussed Gleichenites nor-
detiskioldi Heer, and retained it as a valid species
wliich differs from G. comptoniacjolia by having
long pinna of uniform breadth and pinnules with
somewhat more frecjuent veins. Soriation in the
two species is similar, but Seward noted that G.
nordenskioldi has more sori per pinnule. All of
these features are extremely variable, and speci-
mens from Westwater may be found intermediate
to all of these species-differentiating character-
istics. However, the author has elected not to
make any changes in classical usage of G. nor-
denskioldi and G. comptoniacjolia until further
review.

Gleicheneous fossils from the Westvvater lo-
cality are similar to those figured by Heer ( 1868,
1874, 1882), Seward (1927)"; and others. Similar
problems of classification have been encoimtered
with these ferns in that they exhibit rather wide
morphological variation. However, the present
author has elected to be more conservative than
Heer and Seward, and many fossils which exhibit
some degree of variation are nonetheless placed
in G. comptoniacjolia (Deb. and Ett.) Heer.

This species of fem is the most common ele-
ment from the Westwater area and it is present
in such profusion that at certain outcroppings
one can scarceh' break open a rock without ex-
posing at least tme beautiful specimen. It is in-

Biological Series, Vol. 14, No. 3 Dakoia Sandstone Flora

25

f-:-:-, •

2

v»«i •

Sf

'if*-

Fig. 12. 1. Gleicheniu compUmuwfnlia (Deb. and Ett.) Heer. Pinna illustrating ultimate pinnae and pinnules.
(X1.2). BVU 1818. 2. A.splenium dicksimianum Heer. Pinna fragment illustrating disposition of ultimate pin-
nae. (X1.2). BYU 1812. .3. Gleicheniu complnniaefolia (Deb. and Ett.) Heer. Pinna illustrating ultimate pin-
nae and pinnules. (X1.2). BYU 1819. 4. Sniix newhern/ana Hollick. Leaf demonstrating characteristic shape
and margination. (X1.2). BYU 18.51. 5. Ilex .serrtilus Rushforth, sp. nov. Leaf illustrating shape and margin-
ation. (X1.2). Paratype; BYU 1802. 6. Aspletiium dakotensis Rushforth, sp. nov. Pinna illustrating shape and
disposition of ultimate pinnae and pinnules. (X1.2). Holotype: BYU 1806.

26

Bhic.ham Vou.nc Univeksitv Science Bulletin

Fic. 13. 1. Cleichcnm comptoniacfolut (Deb. and Ett. ) Heer. Pinna frajrment iUustratint; soral disposition. (.X.3).
BYU 1823. 2. Clciclicniii comptonuwfolia { Del), and Ett. ) Hccr. I'innae frapnents u ith pinnnlcs exhibiting
venation. (X3). BVU 1824. 3. Cleichcnm compltonUicfoliu ( Deb. and Ett.) Heer. Pinna fragment illnstrating
characteristic branching. (X3). BVU 182.5. 4. Hausrrmnnia rigida Newberry. Pinnae fragment demonstrating
tongue-shaped pinnules. (X3). BYU 181.5.

Biological Series, Vol. 14, No. .3 D.\kot.\ Sandstone Flora

27

teresting to note that gleicheniaceous ferns are
also most common from the Greenland Cre-
taceous flora. Indeed, as mentioned by Seward,
". . . part of the country was occupied b\' a
vertiable Gleichenietum."

Gleichenia delicatula

Figs. 7-1, 7-2, 7-4; 8-1, 8-3; 9-4; 10-4.

1874 Gleichenia delicatula Heer— Flor. Foss.
Arct., V. 3, pt. 2, p. .54, PI, 9, Figs, lie, llf; PI.
10, Figs, 16, 17,

1882 Gleichenia delicatula Heer— Flor. Foss.
Arct., V, 6, pt, 2, p. 9.

1888 Gleichenia delicatula Heer, Velenovsky—
Abh. K. Bohm, Ges. der Wiss., Math.-Natw.,
ser. 7, V. 1, p. 7, PI. 3, Figs. 12-14.

1910 Gleichenites delicatula (Heer) Seward—
Fossil Plants: v. 2, p. 3.54, Fig. 262b.

1919 Gleichenia delicatula Heer, Berry— U.S.
Geol. Sury. Prof, Pap. 112, p, 54.

Description.— Entire leaf unknown, at least bi-
pinnate; petiole less than 1 mm in diameter,
smooth, appearing dichotomously branched with
a bud in the axis of dichotomy; ultimate pinnae
alternate, linear, up to 15 mm long by 2 mm
wide; pinnules tending toward oyate but with
truncated edge toward pinna apex, oriented
slighth' tovyard pinna apex, approximately 1 mm
long by 1.2 mm wide, attachment basal with
rounded sinuses, margins entire, apices acute to
obtuse, directed apically; primary yein diyiding
immediately upon arising from rachis; secondary
yeins dividing, giving rise to three to five ulti-
mate veinlets which extend to pinnule margin;
fertile specimens unknown.

Occurrence,— Near Westwater, Grand County,
Utah, and Rabbit Valley, Grand County, Utah

Repository.— Brigham Young University, BYU
1826-17.30.

Discussion.- G/eic/ierii'a delicatula was proposed
by Heer in 1874 for placement of several pinna
fragments collected from the Greenland Cre-
taceous. This fern was originally described by
Heer (1874) as follows:

Gl. fronde gracillima, (lichotoma, bipinnata,
racchi tennuissima, pinnis approximatis, pat-
entibus. linearibiis, pinnvilis miiiiitissinms, ro-
tundatus.

This description, and the illustrations of
Heer agree rather well with the Westwater
specimens. One readily apparent difference is

the pinnule shape, which as described by Heer
is round in the Greenland specimens, but tends
toward ovate with an angular apex in the West-
water material. In the oblong pinnule shape,
the Utah material resembles Gleichenia micro-
mera Heer, a contemporaneous species with G.
delicatula. However, the westwater material is
dissimilar to G. micromera in that the pinnules
in the latter are not attached to the rachis with
their entire base, whereas, they are broadly at-
tached and with rounded sinuses in the Utah
Gleichenia.

Velenocsky ( 1888 ) discussed and figured
two specimens of Gleichenia delicatula. These
specimens agree in all details with Heer's
(1874) specimens, and differ from the West-
water specimens only in pinnule shape. Pinna
morphology and size of Velenovsky's specimens
is identical to that of the Utah Gleichenia.

Seward (1910) used the generic name Glei-
chenites for the placement of Gleichenia deli-
catula. He made no changes in description, and
figured a specimen of Gleichenia delicatula
which appears very close to those of Heer
(1874), Velenovsky (1883), and the material
from Utah. Again, the pinnule shape varies be-
tween Seward's specimens and those from Utah,

Berry (1919) reported on Gleichenia deli-
catula collected from the Tuscaloosa Formation
of Fayette County, Alabama, This fern was
placed with G. delicatula based upon the pin-
nules being united to the rachis along their en-
tire base. Berry further mentioned that Fon-
taine (1895) had wrongly placed pinna frag-
ments collected from the Raritan Fomiation in
G. niicroniera. Berry placed these in G. deli-
catula based upon their completely basal at-
tachment. Again, this fern is similar in size to
the Westwater fern, but pimiule shape differs
somewhat.

Even though in pinnule shape the West-
water Gleichenia differs somewhat from other
small pinnuled species of this genus, for the
present time the author has elected to place it
with G.delicatula Heer, This placement is based
upon similar pinna morphology and pinnule size
and disposition.

Family MATONIACEAE

Genus MATOIIDIUM Schenk

Schenk (1871) was the first to use the gen-
eric name Matonidium for placement of fossil
ferns exhibiting similar characteristics as the ex-
tant Matonia pectinata \\. Br. Fossil ferns placed
in this genus are morphologically very similar

28

l$Hi(;nAM ^ouNG Univebsitv Science Bulletin

..'**

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r

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v-*^

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4.

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s:i ■^:^»

7 \

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it^

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V

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Fk;. 11. 1. Mdloiiidiiiiii hrouiiii Hiishtditli. I'iMiia iraKmi'iil illu>tiating \t'ii.iti(>ii .iiid soriatioii <>l piniuik'S. (.\1.2).
BVl' I5fi3. 2. Mdtdiiidium hrownii Busliiortli. Kiihirncineiit of Fijj. 1. (.\2.7). ti\'V 156;5. 3. Clcichcnia
comptunmefolia (Del), and Ktt.) Heer. I'iniia fraf^moiil illiistiatini^ ultimati- pinnae disijosition. (XI. 2). BYU
1820. 4. Astrdloptcris coloraclUu Heveal, Tidwfll, and Huslifortli. Pinna fragment illustrating pinnule attaeh-
ment and shape. (\1.2). BVU 1838. 5. Coiiiopteris urstwiitcrcn.'iK Bushforth, sp. nov. .Sterile and fertile
pinnae. (.\1.2). Holotvpe: BWJ 1803. (i. .\ngiosperni leaf base possihK helduging to h'icus sp. (X1.2). BYU
1847. ...

Biological Series, Vol. 14, Ni). 3 Dakota Sandstone Flora

29

• tin*

*•# . ^ 'If.- . '-•

SsV-'l>

N^

Fic. 15. 1. Miitanidiuni brouuii Rusliforth. Pinna apt'x dfmunstratini^ disposition of pinnules. ( -XI. 2). BYU 1832
2. Matonidiuni hroiinii Hiishforth. Pinna fragment demonstrating venation and soriation. (X2.8). Para-
type: BYU 15.59. 3. Malonidium (?) lanicipinmiliim linshforth. Pinna transfer illustrating pinnule shape
and disposition. (X1..5). Holotype: BVl' 1569. 4. Asiileiiiunt dicksonianunt Heer. Pinna transfer illustrat-
ing pinnule disposition and venation. (X2.4). BYU 1813. 5. Eqiii.\ctitin li/clli Mantel. Portion of rhizome
with two tuhers attaehed. (X1.2).B^'U 1807. 6. Mutimidium hrouitii var. magniiniiiiuluni Rushforth. Pinna
fragment showing long, linear pinnules. (X1.2). BYU 18.36. 7. Satix netcherr\janu Hollick. Leaf demonstrating
characteristic shape and size. (XI. 2). BYU 18.50. 8. Eucahjptus dakotenxis Lesquereu.x. Leaf of character-
istic size and shape. (X1.2). BYU 1848. 9. Equisctum Itjclli Mantel. Rhizome transfer with attached tubers.
(X1.2). BYU 1808.

30

HniciiAM VouNc. University Science Bulletin

and often essentiallv identical to modem maton-
iaceous ferns.

Matonidium is often difficult to differentiate
from other members of the Matoniaceae, par-
ticularly PhU'l)Oj)teris. However, Matonidium is
separated from PliIcl)Oj)trri.s liased upon the pre-
sence of an indusium in the loniier. This differ-
ence is often very difficult to determine even in
fertile specimens, and is impossible to ascertain
in sterile material. Fiirtliermore. to compound
this problem, Matonidium and Phlebojiteris are
often very similar in venation. Further work
dealing with this family will likely show that the
distinction I)etween these two genera is unnat-
ural and that ferns variouslv placed in the two
should be classified under a single genus.

Matonidium americanum Berry em. Rushforth

Kij;s. 18-:5. 18-4; 19-.5.

1916 Matonidium althausii (Dunker) Ward,
Cockerell-J. Wash. Acad. Sci., v. 6, p. Ill, Fig.
2.

1916 Cijcadospadix sp. Cockerell— J. Wash. Acad.
Sci., V. '6, p. 111. Fig. 1.

1918 Matonidium americanum Berrv— Bull. Tor-
rey Bot. Club, v. 46, p. 287, Fig. 2. JPls. 12, Figs.
1-12; 13, .3-6.

1970 Matonidium americanum Berrv, Rushforth
-BYU Geol. Stud., v. 16, pt. .3, p. 25, PI. 8, Fig.
I; PI. 13, Fig. 2.

Description.— Fronds pedate, petiole stout, to
approximately 1 cm in diameter, furrowed (?),
dividing to form a collar from which pinnae are
bom; pinnae lanceolate to linear, 20 cm to 25
cm long, up to 39 per frond; pinnules inserted
near the upper margin of the rachis, up to 9 mm
long by 3 mm wide, coriaceous, more or less
falcate; apices rounded; margins entire or revo-
lute; midvein prominent to near the pinnule
apex; secondary veins prominent, bifurcate ap-
proximately one-half to two-tliirds of the way to
margin, remain tree; sori biseriate, round (ex-
cept where crowded), decrease in size toward
pinnule apex, indusiate; indusium peltate; spor-
angia and spores unknown.

OccuRRENCK.— Near Cutthroat Clulch and Hoven-
weep Canyon west of Dolores, Montrose Coun-
ty, Colorado, and Rabbit Vallev, Grand County,
Utah.

Repository.— Brigham Young University, BYU
1572, 1831.

Discu.ssioM.-As mentioned by Berry (1919) the
pinnules of M. anwricanum are coriaceous. This

feature is easily discerned in the specimens ex-
amined by the present author and may be the
reason that venation in this species is poorly pre-
served. Berry also described the pinnules of this
species as having revolute margins. While this
is possible, it is likely that this appearance is
due to the coriaceous nature of the pinnules.
This feature is (juite unicjue within the Maton-
iaceae.

This species also differs from other species
of Matonidium in the large number of pinnae
bome on a single frond. In addition, the open
venation pattern is somewhat uncommon al-
though not unprecedented within the Maton-

Matonidium brownii Rushforth

FiK.s. 10-3, 10-5; 14-1, 14-2; 1.5-1; 17; 18-2; 18-1,
18-3. 18-6.

1950 Matonidium americanum Berry, Brown—
U.S. Geol. Surv. Prof. Paper 221-D. p. 48, PI. 10.
Figs. 7-8.

1970 Matonidium hrounii Rishforth-BYU Geol.
Stud. 16(3), p. 9, PI. 1, Fig. 2; Pis. 2-5; PI. 6,
Fig. 2; PI. 8. Fig. 2; PI. 11, Fig. 1.

Description.- Petiole stout, between 5 mm ;ind
15 mm in diameter, divided apicallv into
two short arms; pinnae four to ten or more
per frond, lanceolate, up to 14 cm in length,
piniKi rachis between 0.5 mm and 2..5 mm in
di;uneter; pinnules opposite to alternate, basal in
attachment, decurrent but free from each other,
up to 30 mm in length by 9 mm in width, arise
nearly perpendicular to the rachis but become
oriented toward pinna apex; margins entire;
;ipices rounded; midrib prominent to pinnule
apex; secondary veins arise from midrib, divide
to form costal ;ieroles; tertiary veins arise from
costal areole and other secondary veins and
anastomose in tlic pinnule lamiuii; costal areoles
absent in fertile specimens; secondarv veins
fomi an arch above placenta; tertiarv veins arise
from this arch ;md extend to the pLiccnta; upper
epi(lcrm;il cells more or less isodi;mietric, aver-
age 45u in diameter; lower epidemial cells un-
known; sori round, biseriate 1.7 mm to 2.9 mm
in diameter, with a single ring of from ten to
fourteen sponuigia; indusi;i present, peltate, per-
sistent; sp()r;ingi;i cuneatt- sessile or nearly so;
iinnuli obli(|ue; spores trilete, laesurae three
(jUiirters of the distance to the efjuator, rounded-
tri;ingular, 57u in diameter, with a weak margo.

Occurrence.— Known from ne;ir N;iturita, Mon-
trose ('ounty, Colorado, and nc;ir W'estwatcr,
CJnmd (>ountv, Utah.

BioLocK.AL Skries, \()l. 14. No, 3 Dakota Sandstone Flora

31

sILJh T'^1 '"^ (M-2) BVU 1843 ^ Ilex

Sm ^"p.^"'*^-^'', ""^'- AnRuIar-ovate leaf with typical serrate to spinose Lr^in. (XI 2) Holot pe"' BYU

1801. 3 Ficus daphno<,cno,dc., (Heer) Berry. Cuneate leaf hase exhibitini- cuneate base XI f^^^' 1845

5 Ji'X "'''"-g-."'"'" («-■■), Berry. Leaf e.xhibiting drip point, sbape and mar.ination. X ) BYU IH^

neer. rortion ot leal exhibiting characteristic- venation. (.\]..5). BVU 1849.

32

Bhicham Young University Science Bulletin

Hki'ositohy.— Brigham Young University, holo-
tvpe: BYU 1557; paratypw; BYU 1559, 156.3;
BYU 1832-18^5.

Discussion.— Af. brotvnii was first tk'scribed by
Brown (1950) from specimens collected from
the Dakota Sandstone of southwestern Colorado.
However, Brown placed this feni in ^^ . amcri-
canum Berr\'. The present author examined the
specimens collected from this locality and con-
cluded that these specimens should not be placed
with M. (imericamtm. Due to this. M. hrounii
Hushforth ( 1970) was proposed.

M. I)ruunii differs from M. americamtm by
being much larger; by having fewer piniuie per
frond; bv having anastomosing rather than open
venation; and in soriation (Hushforth, 1970).

M. hroumi differs from a related species, M.
uicstwri Krasser, by having anastomosing rather
than open venation; by being larger in overall
size; and bv having more numerous sori. M.
(iltlmusii (bunker) Ward differs from M.
hrotinii in piniuile shape and habit, ;uid in ven-
ation. Soriation of M. althausii also differs from
M. hrotinii in that the fomier exhibits sori bom
to the extreme pinnule apex decreasing in size
as the apex is approached.

In venation M. hrounii is similar to Phleh-
0})teris ( Schenk ) Schenk.

Matonidiurn hroicnii var. nuignipinnulum
Rushforth

Kigs. 1.5-6; 18-2; 2Ib.

1970 Matonidiurn hroicnii var. magnipinmdum
Uushforth-BYU Geol. Stud., v. 16, pt. 3, p. 11,
Pi. 6, Fig. 1; PI. 7, Fig. 2; PI. 14, Fig. 2.

Dk.scriptiox.— Entire leaf unknown, pedate; pin-
nae observed up to 30 cm long, longer in
growth, lanceolate; rachis up to 1.5 mm in di-
ameter; pinnules opposite to alternate, basal in
attachment, often decurrent, free, up to 60 mm
in length bv 6 mm in width, arise at ^X) but be-
come oriented toward pinna apex; margins en-
tire; apices rounded; midvein prominent to pin-
nule apex; secondary veins prominent, forming
costal areoles in sterile pinnules, progressing di-
rectly to phicenta in fertile tonus; tertiary veins
often anastomose except in pinnule tips; lower
epidermal cells large (up to 95u by 45u), lobed;
fertile specimens not observed.

Occurrence.— Known from the Dakota Sandstone
Fomiation ne;ir Westwater, (irand C-ounty, Utah.
Reposiiohy.— Ikigham Young Universitv, holo-
type; BYU 1566; BYU 18.36. '

Dlscussion.— M. hrounii var. m<i<ini])innuhim
was described by Rushforth (1970) for maton-
iaceous frond fragments from the Westvvater
flora. In pinnule habit and venation M. hrownii
var. in'ii!_nipinnuhim is very similar to M. hroionii
var. hrounii. However, the pinnules of var.
nii<^ni]iinnuhiin are consistently longer and nar-
rower than those of var. hrownii. In addition,
the disposition of the pinnules on the rachis Ls
farther ;ipart in var. nuiiinijiinntilum.

As mentioned by RusMorth ( 1970 ) , this fern
inav be distinguished from other matoniaceous
ferns due to its large pinna and pinnule size.
However, long, linear pinnules are not unknown
within the Nlatoniaceae. Bell (1956) reported
on Phk'hopteris (?) elongata for sterile pinna
fragments from the Cretaceous of Canada, and
P. imiensteri also exhibits long linear pinnules.

Miitonichuin (?) lanci])innuhini Rushforth

Figs. 1.5-:3; 18-1; 19-2.

1970 Matonidiurn (?) Iancii>innuhini Rushforth
BYU C;col. Studies, v. 16 pt. 3, p. 24, Pi. 7, Fig.
1; PI. 9; PI. 10; PI. 13, Fig. I.

Description.— Entire leaf unknown; pinnules
sub-opposite, linear-lanceolate, 60 mm long by
12 mm wide, essentiallv perpendicular to rachis
except at extreme pinnule apex, attachment ba-
sal with rounded sinuses; midvein prominent to
pinnule apex; secondary veins prominent, form-
ing costal areoles, anastomosing above c-ostal
areoles except near laminar edge, composed of
tracheids with annular, helical and reticulate
wall thickenings; guard cells 4(hi long by 15u
wide, leaving stomata lOu wide; fertile speci-
mens unknown.

Occurrence.— Dakota Sandstone Fonnation near
Westwater, Grand ("ounty, Ut;di.

Repository.— Brigham Young University, holo-
type: BYU 1569; paratype: BYU 1579. "

DiscussioN.-Rushforth ( 1070 ) phiced this fern
in the Nlatoniaceae based upon pinnule venation
and shape. However, the assignment of this fern
to this familv must remain provisional until fer-
tile specimens are collected.

One interesting fact concerning this feni is
that it seems to be somewluit intermediate in
venation pattern betwi'cn wiil-di'tiiu-d members
of the Matoniaceae and another WesU\'ater feni,
.\.stralo))t('ri.s ( Polypodiaceae ) . This latter genus
illustrates \cnation which would tend to align
it with the \hitoniacc;ie, but sond characteristics
align it witli the Polypodiaceae. Further at-

BioLocicAL Sehiks, \'()i.. 14, No. 3 Dakoia Sandstone Floha

33

Fig. 17. Mahmidium hroicnii Rushforth. Portion of a frond illustratinsr pinnae and pinnule di.sposition (.\12)
Holotype: BYU 1557. f i

34

Bhigham Young Univehsitv Science Bulletin

tempts to locate more specimens of M. l(mci])in-
ntilum are presently under way.

FamUy POLYPODIACEAE

Genus ASTRALOPTERIS Reveal, Tidwell
and Rushforth

Astralopteris coloradica ( Brown ) Reveal,
Tidwell and Rushforth

Fig.s. 9-1, 9-7; 14-4; 19-4; 20.

1950 Bohtis coloradica Brown-U.S. Geol. Surv.
Prof. Paper 221-D, p. 49, PI. 12, Figs. 6-7.

1967 Astralopteris coloradica ( Brovra ) Reveal,
Tidwell and Rushforth, Tidwell et o/.-BYU
Geol. Studies, V. 14, p. 2.39, PI. 2, Fig. 1; Pis. 3-6.

1968 Astralo])lcris coloradica ( Brown ) Reveal,
Tidwell and Rushforth, Rushforth and Tidwell-
BYU Geol. Studies, v. 15, p. 109, Pis. 1-3.
Description. —Entire frond unknown; pinnae in-
complete, large, ovate, tapering to rather acute
apices; pinnules coriaceous, pinnate, alternate to
opposite on rather stout rachises; lower pinnules
linear-lanceolate, about 7 to 11 times as long as
broad, sessile to short stalked attaclunent; mar-
gins entire; apices acute to rounded; upper pin-
nules similar to lower ones, 4 to 5 times as long
as broad, basal attachment with rounded sinuses
in the uppennost pinnules; midvein prominent
to the pinnule apex or nearly so, arising decur-
rently from rachis; lateral veins numerous aris-
ing acutely and then becoming nearly at right
angles to the midvein (or divaricate) for nearly
their entire length, simple for one-half to three-
quarters or more of their length then usually bi-
furcating two or three times (or rarely trifur-
cating at the first fork); ultimate veins finer,
anastomosing, often obscure; sori round, biseri-
ate between the main lateral veins, about one-
fourth of the distance from the midrib to the
margin, fed from a costal areole and other sec-
ondary veins which extend to the placenta; spor-
angia 12 to 15, arranged in a ring around a cen-
tral placenta; annuli vertical; spores unknown.

OccunRENCF..— Known from the Dakota Sandstone
Formation near Westwater, Grand C;ounty,
Utah.

Rei'Ositohy.— Brigham Young Universitv, BYU
1415-1422; 1837-1840.

Discussion.— A. coloradica was originally de-
scribed as Bolhiti.s coloradica by Brown (19.50).
This placement was based upon sterile speci-
mens collected from Naturita, Montrose Gounty,
Golorado. Brown encountered difficulty in plac-
ing this fern and consefjuently he conferred

with extant feni workers from the Smithsonian
Institute. These workers suggested that in over-
all gross morphology (pinnule shape, size, ven-
ation, etc. ) the Naturita fossils are similar to
fenis of the extant genus Bolbitis.

Tidwell et al. ( 1967 ) reported fertile speci-
mens of this fern and illustrated that this fern
eould not be placed in the extant Bolbitis. Fur-
thennore, since no living or fossil genus allowed
for the placement of this fern the new genus
Astralojiteri.s- was proposed.

A. coloradica is one of the most common ele-
ments of the Westwater flora. In some collect-
ing sites it is collected almost exclusively, al-
though it is most commonly collected in associ-
ation with Matonidium and Gleichenia. GoUec-
tioas of fertile specimens of this fern are rare at
the Westwater locality but, from recent exami-
nations bv the author, appear to be somewhat
more common from the Dakota Sandstone of
southwestern Golorado and Arizona.

UNGLASSIFIED FERNS

Genus GLADOPHLEBIS Brongniart

This generic n.mie was originally proposed
by Brongniart (1849) but Saporta (1873) first
defined (Cladophlebis) as follows:

Frond pinnatelv cli\'idecl; pinnules separate
from one another, or slightlv miited, attached
In the entire base.

Schimper (1874) emended this diagnosis to:

Fronds pinnately divided; pinnae spreading;
lolies or pinnules attached by the entire liase,
sometimes confluent, rarely slightly auriculate,
accuniinate or obtuse. occa.sional!)' dentate,
especialK- at the ape.\, not rarely subfalcately
curved upwards; midner\es pretty strong; sec-
ondary nerves departing at a more or less
acute angle, dichotomous; slender to very
slender.

Fontaine ( 1889) modified this diagnosis only in
regard to venation by saying, ". . . midnerve
strong at base, and towards the summit dissolv-
ing into branches."

Fontaine ( 1889 ) discussed the usage of CAad-
ophlebifi and concluded that it is a useful genus
for placement of sterile foliage which may be
described as above. This is indeed the case, as
foliage assigned to Cladopldebis could fit well
into ;i luimber of different genera. Therefore,
Cladophlebis has become a useful, though un-
niitunil genus which is very characteristic of the
Mesozoie, particularly of Jurassic times. Thus
Berrv ( 1912) mentioned that Cladophlebis is
essentially a fomi genus and includes species

Biou)ck;al Skhies, \'ol. 14, No. 3 Dakoi a Sandstone Flora

35

^^f^"^^

J

k

\

2

/

,4

'J

>. t

*''-^ii^

'^J

' '■/,

'*• : h?

'^nr^-

* ^

Fig. 18. 1. MatonidiiiTii (?) hmcipinnulum Rusliforth. I^iriiiu fragment illustratini; pinnule dispositidn. (X1.2).
Paratype: BYU 1570. 2. Matoniditim hrvunii Rusliforth. Trilete spore illii.strating weakly defined margo.
(X1200). Paratvpe: BVU 1.561. 3. Mutonidium amcricanum Berry em. Rusliforth. Two pinna fragments.
(X1.2). BVU 1.572. 4. Mutonidium aTiiericaninn Berry em. Rushforth. Portions of two pinnae illustrating
venation. (X.3.6). BVU 1572.

36

Bricham Young University Science Bulletin

which when fertile may be placed into several
different genera and even families.

Fontaine (18S9) treated 23 species and sev-
eral varieties of Cladoplilchis from the American
Potomac Group. However, Berry (1912) revised
this genus, as known from the Potomac, and rec-
ognizezd eight species. As mentioned by Berry,
Fontaine recorded thi.s high number of species:

. . . altogether losing .sight of variation and
changes due to age or to position of the fos-
sils with regard to the frond as a whole, as
well as changes due to the direct action of
the environment.

Seward (1894) in treating Wealden fossils
in the Britich Museum of Natural History ac-
cepted the generic diagnosis of Schimper ( 1874)
as emended by Fontaine (1889). Berry (1912)
also followed this same generic diagnosis, and
most workers subsequent to Berry have done
likewise.

The literature dealing with Cladophlethis is
voluminous and is in need of reviewing. Many
species are recognized, although this is to be
expected in anv unnatural genus as reproductive
morphology may not be compared. However,
many species appear to be identical, and the
genus should be monographed.

Cladophlebi.s cotutricta Fontaine em. Berry

Figs. 9-5; 11- 1.

1889 Cladophlebis constricta Fontaine— U.S.
Geol. Surv. Mon. 15, p. 68, PI. 2. Fig. 11; PI. 3,
Fig. 2; PI. 6, Figs. 5, 6, 8-14; PI. 21,>igs. 9, 13;
PI. 169, Fig. 2.

1888 Chidophlehis latifolia Fontaine-U.S. Geol.
Surv. Mon. 15, p. 68, PI. .3, Fig. 1; PI. 6, Fig. 4.

1904 Cladophlebis constricta Fontaine, in Ward
-U.S. Geol. Surv. Mon. 48, p. 197, PI. 71, Fig. 26.

1905 Cladophlebis virginensl's Fontaine, pro
parte, in VVard-U.S. Geol. Surv. Mon. 48, p. 512,
PI. Ill, Fig. 7.

91 Cladophlebis constricta l-'ontaine. Berry—
Proc. U.S. Nat. Mus., v. 41, p. 314.

Desciuition'.— Entire leaf unknown; pinnules
typically greater than 27 mm long by 10 mm
wide, margins deeply lobed up to one-half of the
distance to the rnidvein, lobe margins entire,
lobe apices obtuse; midvein very strong; second-
ary veins repeatedly forked, open to the margin;
fertile specimens unknown.

Occurrence.— Near Westwater, Grand (^ountv,
Utah.

Hiii'osiTOHY.— Brigham Young University, BYU
1841.

Discussion.— This feni is known from the West-
water flora only from a single specimen. This
sp(>cimen illustrates fragments of two pinnules
each of which is rather well preserved, exhibit-
ing venation well.

This specimen agrees well with some speci-
mens placed by Fontaine (1889) in Cladophle-
bis constricta, especially PI. 2. Fig. 11a. How-
ever, due to the paucity of fossil material from
Westwater, it is not known if the Utah speci-
mens agree in all details with the specimens
from Virginia. Even though this is the case,
venation, overall size, and pinnule lobing, are all
in agreement with C. constricta Fontaine em.
Berry, and the author believes these similarities
are sufficient to warrant the placement of the
Westwater fern in this species.

Cladophlebis parta Fontaine em. Berry

Figs. 8-5; 11-2.

1889 Cladophlebis parva Fontaine-U.S. Geol.
Surv. Mon. 15, p. 73, PI. 4, Fig. 7; PI. 6, Figs. 1-3.

1899 Cladophlebis parva Fontaine, in Ward—
U.S. Geol. Surv. 19th Ann. Rept., pt. 2, p. 657,
PI. 160, Fig. IS.

1904 Cladophlebis parva Fontaine, in Ward—
U.S. Geol. Surv. Mon. 48, p. 125, Pl. 65. Figs.
5-8.

1911 Cladophlebis ])arva Fontaine, Berry— Mary-
land Geol. Surv., Low. Cret., p. 250, PI. 27, Figs.
1-2; Pis. 39-31.

I'dl'lCladophlebls parva Fontaine, Berry— Proc.
U.S. Nat. Mus., V. 41, p. 316.

1936 Cladophlebis ]>arva Fontaine, Bell— Geol.
Curv. Can. Mem. 285, p. 56, PI. 11, Figs. 3-5; PI.
12, Fig. 3; PI. 13, Figs. 1-2; PI. 14, Fig. 3; PI. 14,

Fig. 3":

Description.— Entire leaf unknown; pinnae prob-
ably lanceolate; rachis about 1 mm in diameter;
pinnules alternate, linear-lanceolate, up to 26
mm long bv 6 mm wide, lobed approximateh'
one-half of the way to the midrib, margins of
lobes sliglitlv undulate, apices acute, attach-
ment basal to somewhat constricted basal; mid-
vein prominent, undi\ided to pinnule ;ipex; sec-
ondary veins divide from one to thn-e times, re-
main free; fertile specimens unknown.

Occurrence. —Near Westwater, Grand County,
Utah.

Bi()L(k:ic:al Series, Vol. 14. No. 3 D.akot.\ S.^niwtone Fi.oh a

37

Fig. 19. 1. Matonidium brounii Rushfortli. Pinna fninrriL-nt illustrating soriation and venation. (X3). Paratype:
BYU 1563. 2. Malonidium (?) laiuijnniitilum Riishforth. Pinna fragment illii.strating pinnule disposition
and venation. Holotvpe; BYU 1.569. 3. Matonidium hroivnii Rushforth. Pinnule exhibiting venation. (.\3).
Paratype: BYU 1559. 4. Astralopteris cnlnradertsis Reveal, Tidwell, and Rushforth. Pinnule e.vhihiting
venation. (X3). BYU 1837. 5. Matonidium americanum Berry em. Rushforth. Pinna fragment e.vhihiting
venation. (X3). BYU 1831. 6. Matonidium brownii Rushforth. Pinna apex illustrating pinnule disposition.
(X3). BYU 183.5.

38

Brigham Young Univebsity Sciencf Bulletin

Repository.— Brigham Young University, BYU
1842.

Discussion.— Cladophlehi.s parva Fontaine em.
Berry is known only from one specimen in the
VVestwater flora. This specimen illustrates sev-
eral pinnules on a pinna Iragment, a few of
which are rather complete. This fern from the
VVestsvater locality has been assigned to Clcid-
ophlebis parva based upon venation, pinnule
shape and habit, and size. However, it is al-
ways difficult to place fragments of Cladophle-
hus accurately, and this feni resembles several
other species of CAadophlehis.

Division

ANTHOPHYTA ( MAGNOLIOPHYTA )

Family AQUIFOLIACEAE

Genus ILEX L.

Uex serrate Rushforth, sp. nov.

Figs. 12-5; 16-2.

Description.— Leaves vary from fairly short and
wide (3.5 cm long by L7 cm wide) to longer
and thinner (4 cm long by 1 cm wide), some-
what coriaceous, angular-ovate; apex acute to
accuminate; base narrowing to the petiole; mar-
gins serrate to somewhat spinose; petiole short;
midvein weak, continuing to apex; secondary
veins mostly obscured, borne at acute angles,
progressing to margin.

Occurrence.— Dakota Sandstone Formation near
Westwater, Grand County, Utah.

Repository.— Brigham Young University, holo-
type: BYU 1801; paratype BYU 1892.

Dlscu.ssion.— Leaves assigned to this species
from near Westwater are somewhat similar to
Ilex dakotensis Les(juereux in shape and size.
However, the leaf originally placed in l.dako-
tetm.'i dtx-s not exhibit serrate margins, and ven-
ation differs somewhat from Utah specimens. In
overall shape, the Westwater material agrees
rather closely with Ilex stenoplit/lla Unger from
the Tertiary of Europe, as does Ilex dakotett^is.
However, Les(|uereux (1892) pointed out that
these two species differ in size and somewhat in
shape. In addition, /. stenoplujlla is not serrate
to spinose as is the Utah Ilex.

Ilex serrata differs from other species of Ilex
in its small size coupled with spinose margins.
In margination, /. serrata is similar to /. armata
Lesquereux, although the latter is much larger
and of different shape than /. serrata.

Family MAGNOLIACEAE

Genus .MAGNOLIA L.
Magnolia houlaijarui Lescjuereux

FiR. 16-1.

1792 Maglunia hotddt/anu Lesquereux— U.S.
Geol .Surv. Mon. 17. p.' 202, PI. 60. Fig. 2.

1894 Ma<inolia ^lauroides Holliek— Bull. Torrey
Got. Club, V. 21, p. 60, PI. 175, Fig. 1-7.

1895 Magnolia glaueoides Holliek, Newberry-
U.S. Geol. Surv. Mon. 26, p. 74, PI. 47, Figs. 1-4.

1995 Magnolia glaueoides Hollick-U.S. Geol.
Surv. Mon. 50, p. 67. PI. 19, Fig. 6; PI. 20, Fig. 6.

1909 Magnolia hoidai/aiui Les<juereux, Berry-
Bull. Torrey Bot. Club, v. 36, p. 254.

1911 Magnolia boulatjana Lesquereux, Berry-
New Jersey Geol. Surv. Bull. 3, p. 131, PI. 14,
Fig. 2.

Description.— Leaf greater than 7.5 cm long by
4 cm wide, narrowly elliptic in outline, coria-
ceous; base rounded to bluntly rounded; margins
entire; midvein strong, bearing somewhat in-
fre(juent secondaries; secondary veias pinnate,
parallel, progressing to near margin and then
beinding to unite with vein above to fomi
rounded areoles near the margin; tertiary veins
connecting the secondaries.

Occurrence.— Near Westwater, Grand County,
Utah.

Repository.- Brigham Young Universitv, BYU

184,3.

Discussion.— Berry (1911) noted that leaves of
Magnolia boulai/ana Lescjuereux are remarkably
constant in size and shape. This is particularly
noteworthv when it is recalled that this con-
stancy extends over wide geographical areas and
for a good deal of time.

Berry (1911) noted that leaves assigned to
Magnolia glaueoides Holliek by Holliek and
Newberry are identical to leaves of M. houlaij-
ana. In view of this. Berry placed M. glaueoides
in synonomy under M. houtihii/ana.

Placement of the Utah Magnolia in Magnolia
houllayana Lesquereux may be made without
hesitation, as size, shape, and venation are all
identical between the two.

Family MORACEAE

Genus FICUS L.

This genus in regard to the Mesozoic has be-
come more of a form genus than a natural

Biological Series, Vol. 14, No. 3 D.^ikota Sandstone Flora

39

^4»^'J3

' '■.A?i^

Fic. 20. Astralopterui coloradica Reveal, Tidwell. and Rushforth. Pinna exhibiting pinnule disposition. (X1.2).
BVU 1415.

40

Brigham Young University Science Bulletin

unit. Into this genus are placed leaves which
are somewhat similar in that they exhibit laii-
raceous characteristics. However, Cretaceous
leaves placed in this gcmis often could be placed
with ecjual logic in various other genera such as
Salix or Laurus.

Ficus daplaiugenoidcs (Heer) Berry

Figs. 16-3, 16-4.

1867 Proteoide.1 daphnogenoides Heer, in Ca-
pellini and Heer— Denksch. Allgem. Schweiz.
Ges. Naturwiss., v. 22, p. 17, PI. 4, Figs. 9-10.

1874 Proteoides daphnogenoides Heer, Lesquer-
eux— U.S. Geo!. Surv. Terr. Kept., v. 6, p. 85,
PI. 15, Figs. 1-2.

1905 Ficu.i daphnogenoides (Heer) Berry-
Bull. Torrey Bot. Club. v. 32, p.329, PI. 21.

1921 Ficus daphnogenoides (Heer) Berry— U.S.
Geol. Surv. Prof. Paper 129, p. 163, PI. 39, Fig. 1.

1959 Ficus daphnogenoides (Heer) Berry,
BrowTi-U.S. Geol. Surv. Prof. Paper221-D, p.
50, PI. 11, Figs. 1, 2, 4.

Description.— Leaves typically greater than 12
cm long bv 3 cm wide, more or less oblanceo-
late; apex accuminate, fonning a rather long
drip pohit; base cuneate; margins entire; mid-
vein strong at point of origin, becoming weaker
towards leaf apex; secondary veins pinnate, a-
rising acutely, bending to parallel margin; terti-
ary veins obscured.

Occurrence.— Near Westwater, Grand County,
Utah.

Repository.- Brigham Young University, BYU
1844-7846.

Discussion.— Lesiiuereiix (1874) described Pro-
teoides daphnogenoides Heer (Ficus daphno-
genoides) as follows:

Lfiivcs oviitc-laiiccolatf near the base, j^radii-
ally tapering uiiuard to a long, aeiite, seythe-
shaped point, entire, smooth, and coriaceous;
medial ner\e narrow; secondary veins obsolete,
few, ascending under a verv acute angle from
the medial nerse and following the borders.

This description fits the Westwater Ficus in all
respects, and the identity of the two may bo
fairly certain.

Berry ( 1905) transferred Proteoides daphno-
genoides Heer to the genus Ficus. This trans-
ferral was valid, ;dtliongh Brown (1950) re-
marked:

These entire-margined leaves li.ivc been re-
ferred to manv different genera with little

satisfaction, as might l)e expected for entire
leaves, the internal senation strucure of which
tends in m;iny towanl similarity. 1 regartl this
reference to Fici/.v its makeshift. Many similar
leaves from the Dakota Sandstone have been
referred with little evidence to Andromeda.
All are perhaps examples of lauraceous species
that are difficult to separate.

Family MYRTACEAE

Genus EUCALYPTUS L. Herit
Eucahjptus dakoten.'iis Les(juereux

Fig. 1.5-8.

1892 Eucahjptus dakotensis Lesquereux— U.S.
Geol. Surv. Mon. 17, p. 1.37, PI. .37, Figs. 14-19.

1895 Eucahfptus angustifolia Newberry— U.S.
Geol. Surv. Mon. 26, p. Ill, PI. .32, Figs. 1, 6, 7.

1895 Eucahjptus (?) nervosa Newberry— U.S.
Geol. Surv. Mon. 26, p. Ill, PI. .32. Figs. .3-5, 8.

Descriition.— Leaf linear, 4.5 cm long by 1.1 cm
wide, somewhat coriaceous; base cuneate; mar-
gins entire; midvein strong; secondary veins pin-
nate, arising obtusely, extending to leaf margin.

Occurrence.— Near Westwater, Grand County,
Utah.

Repository.— Brigham Young University, BYU
1848.

Discussion.— Lesquereux ( 1892) described Eu-
cahjptus dakotensis as follows:

Leaves coriaceous, linear, or gradually narrow-
ed from an obtuse apex to the base, deeur-
ring into short, alate petiole; borders recur\ed,
medi;m nerve strong; secondaries thin, obli(|ue.
proximate, parrallel. camptodrome.

Lesquereux further pointed out that the margin
of some specimens was flat rather than recurved.

Eucahjptus angustifolia Newberry and £. (?)
nervosa Newberry appear to be specifically
identical both with each other and with E. dako-
tensis Lescjuereux. The only apparent difference
is that the petiole of E. angustifolia is naked
rather than a];ite.

The Westwater Eucahjptus also agree well
with the description of Les(|uereux (1892), ex-
cept that the petiole of the Utah sjjecimen is
not winged. However, this difference does not
warrant specific separation, and E. dakotensis
is used for placement of the Westwater speci-
men.

As mentioned by Les((uereux ( 1892) E. dako-
tensis differs from an ;ipp;ireiitlv closely rei;ited
fonii, £. geinitzi in that the tonner is snuiller
and linear in shape as opposed to larger and
broadly spatulate in shape.

Biological Sehies, \'ol. 14, No. 3 Dakota Sands ione Flora

41

t

^

1* *

a
'W:-*^!

s ■

■} "V -.,^

X

x

^*^v

; .^.i* «??v

•^.'

a

{"

^ 'V-'

■*?,

'^^>

<^'

\Ul»'

4 ■" ;^, ^^.

■'^, ^A,^ w ^,

Fic. 21. A. Gk'ichenia conipfoniticfolia (Deb. and Ett.) Heer. Pinna fragments e.xhibiting pinnule dispo.sition.
(XI. 5). BVU 1853a. B. Mutoiiidium hroicnii var. magnipinnuluni Hiishforth. Pinna fragments. (XI. 5). BYU
18.53b.

42

Bbigham Young University Science Bulletin

Family PLATANACEAE

Genus PLATANUS L.

Leaves of the genus Platanus from the Cre-
taceous are well known and rather common.
Manv of these leaves are essentially indistin-
guishable from those of extant species (Seward,
1827).

Platanus newbernjana Heer

Fif;. 16-6.

1867 Platanus newbernjana Heer, in Capellini
and Heer— Denksch. Allgem. Schweiz. Ges. Na-
turwiss., V. 22, p. 16, PI. 1, Fig. 4.

1868 Platanus neioberrtjana Heer. Lesquereux—
Amer. Jour. Sci., v. 46, p. 97.

1874 Platanus netLhern/aiui Heer, Lesquereux—
U.S. Geo!. Surv. Terr. Rept., v. 6, p. 72, PI. 8,
Figs. 2-3; PI. 9, Fig. 3.

1874 Platanus affinls Lesquereux— U.S. Geol.
Surv. Terr. Rept., v. 6, p. 71, PI. 4, Fig. 4.

1873 ProtophtjUum minus Lesquereux— U.S.
Geol. Surv. Terr. Rept., v. 6, p. 194, PI. 27 Fig. 1.

1874 Protojiht/lluin ncbrascense Lesquereux—
U.S. Geol. Surv. Terr. Rept., v. 6, p. 103, Pi. 27,
Fig. 3.

1882 Platanus affinis Lesquereux, Heer— Flor.
Foss. Arct., V. 6, pt. 2, p. 73, PI. 18, Figs. 16-17.

1883 Platanus newbern/ana Heer, Lesquereux—
U.S. Geol. Surv. Terr. Rept., v. 8, p. 28, PI. 59,
Figs. 1-6; PI. 60, Fig. 1.

1883 Cissites affinis Lesquereux— U.S. Geol.
Surv. Terr. Rept., v. 8, p. 67.

1927 Platanus neiiberrtjana Heer, Seward— Phil.
Trans. Rov. Soc-. London B, v. 215, p. 128, PI.
11, Fig. 116; Text- Fig. ,30.

Description.— Leaf greater than 5 cm long by 3
cm wide, coriaceous; base rounded; midvein
strong bearing numerous secondary veins at
acute angles; secondary veins apparently extend
to margin ( caspcd(xlrome ) ; tertiary veins nu-
merous, borne at nearly 90° angles to secondary
veins, connecting secondaries.

Occurrence.— Dakota Sandstone Formation near
Rabbit Valley, Grand County, Utah.

Rkpository.— Brigham Young University, BYU
1849.

Discussion.— This leaf agrees in all details with
Plantanus neuherryana Heer, and is very similar
to leaves figured under this species by Lesquer-
eux and Seward. In addition, it is very similar
to leaves described as Platanus affinis, Pro-
tophijllum minus, and ProtophtjUum nebrascense
by Lesquereux (1874, 1882). The Westwater
leaf exhibits a rounded to angular base, which
agrees with the description of Platanus new-
bernjana by Lescjuereux ( 1874 ) , although in
this respect it differs somewhat from specimens
placed in this species by Seward ( 1927 ) which
exhibited very rounded bases.

Seward (1927) noted that leaves of Platanus
newbernjana "agree very closely with the recent
Platanus mexicana Moric."

FamUy SALICACEAE

Genus SALIX L.
Salix newbernjana Hollick

Figs. 12-4; 15-7.

1895 Salix newbernjana Hollick, in Newberry—
U.S. Geol. Surv. Mon. 26, p. 68, PI. 14, Figs. 1-7.
1911 Salix newbernjana Hollick, Berry— New
Jersey Geol. Surv. Bull. 3, p. 113, PI. 11,' Fig. 2.

Description.— Leaves greater than 6 cm long by
up to 2.5 cm wide, lanceolate; base rounded;
apex accuminate to fonn a rather well defined
drip point; margins finely crenate to dentate;
midvein fairly strong; other veins obscure.

Occurrence.— Dakota Sandstone Fonnation near
Westwater, Grand County, Utah.

Repository.— Brigham Young University, BYU
1850-1851.

Discussion.— This leaf from Westwater is essen-
tially identical to those described by Hollick (in
Newberry, 1895) and Berry (1911). It may be
placed with confidence in Salix twwbernjana
Hollick.

ACKNOWLEDGMENTS

The author wishes to express his appreci- also due to several people who aided the author

ation for the many helpful suggestions of Dr. in completing field work and technical aspects

W. D. Tidwell and Dr. J, R. Bushman during of this manuscript.
the preparation of this manuscript. Thanks are

Biological Sehies, Vol. 14, No. 3 D.\kot.\ S.\ndstone F"lora

REFERENCES CITED

43

.■\c;a.sie, ]. M. 1967. Upper Cretaceous palynomor-
phs from Coal Can von, Coconio County, Arizona:
Unpub. M.S. ThesLs, Univ. of Arizona, 1(M p.

Bailev, E. W. and I. W. SiNNOTT. 1915. A botanical
inde.\ of Cretaceous and Tertiary climates; Science,
V. 41, p. 831-834.

Bell, W. A. 1956. Lower Cretaceous flonis of west-
em Canada: Geol. Surv. Can. Mem. 285, 331 p.

Berry, E. W. 1905. Additions to the fossil flora
from Cliff wood. New Jersev: Bull. Torrey Bot.
Club, v. 32, pp. 43-48.

. 1910. The evidence of the flora regarding

the age of the Baritan Formation: Jour. Geol., v.
18, pp. 252-2.58.

. 1911a. Svstematic palentology of the Lower

Cretaceous deposits of Maryland. Pteridophyta:
Marvland Geol. Surv., Lower Cretaceous, pp. 285-
294.'

. 1911b. The flora of the Raritan Formation:

New Jersey Geol. Sur\-. BuU. 3, 233 p., 29 Pis,

. 1912. A revision of the fossil fenis from the

Potomac Group which have been referred to the
the genera Cladophlehis and Thnjsopteris: Proc.
U.S. Nat. Mus.. V. 41, pp. 307-332.

. 1919a. A new Matonidiuin from Colorado,

with remarks on the distribution of the Maton-
iaceae: Bull. Torrey Bot. Club. v. 46, pp. 285-294.

. 1919b. Upper Cretaceous floras of the east-
em Gulf region in Tennessee, Mississippi, Alabama,
and Georgia: L'.S. Geol. Surv. Prof. Paper 112,
175 p.

. 1920. The age of the Dakota flora. Amer.

Jour. Sci. and Arts,"v. .50, pp. 387-390.

— . 1922. The flora of the Woodbine Sand at

.Arthurs Bluff, Te.xas: U.S. Geol. Surv. Prof. Paper
129-G, pp. 153-180.

1924. Mesozoic Gleichenia from Argentina:

Pan-Amer. Geologist, v. 41, pp. 17-24.

Bower, F. O. 1926. The Fems: v. 2, 344 p., Cam-
bridge.

. ' 1928. The Fems: v. 3, 306 p., Cambridge.

Broncniart, a. C. 1849. Tableau des Genres de
Vegetau.x Fossiles Consideres sous le Point de Vue
de Leur Classification Botanique et de Leur Dis-
tribution Geologifjue: 127 p., Paris.

Brown, R. W. 1950, Cretaceous plants from south-
western Colorado: U.S. Geol. Surv. Prof. Paper
221-D, pp. 45-66.

. 1952. in Stephenson. L. W., Larger inverte-
brate fossils of the Woodbine Formation ( Ceno-
manian) of Texas: Lf.S. Geol. Surv. Prof. Paper
242. p. 35.

Capellini, G. and A. Heeh. 1867. Les phyllites Cre-
tacees du Nebraska: Denksch. Allgem. Schweiz.
Ges. Naturwiss., v. 22, p, 1-20, 4 Pis,

Chaney, R. W. 1954. A new pine from the Cre-
taceous of Minnesota and its paleoecological sig-
nificance: Ecology, v. 35, p. 145-151.

Chaney. R. W. and E. 1. Sanborn. 1933. Eocene
Goshen flora of Oregon: Carnegie Inst. Wash. Pub.
439, 342. p.

Cobban, W. A. and J. B. Reeside, Jr. 19.52. Corre-
lation of the Cretaceous formations of the western
interior of the United States: Geol. Soc. Amer.
Bull. 63, p. 1011-1044.

Cockerell, T. D, a. 1916, A Lower Cretaceous
flora in Colorado: Jour. Wash. Acad. Sci., v. 6,
p. 109-112.

Coffin, R. C. 1921. Radium, uranium, and van-
adium deposits of southwestern Colorado: Colorado
Geol. Surv. Bull. 16, 230 p.

Darton, H. 1905. Preliminary report on the geology
and underground water resources of the central
Great Plains: U.S. Geol. Surv. Prof. Paper 32,
435 p.

Dunker, W. 1846. Monographic der Norddeutschen
Wealdenbildung. Ein Beitrag zur Geognoise und
Naturgeschichte der Vorwelt: 83 p., 21 Pis.. Braun-
schweig.

Fontaine, W. M. 1889. The Potomac or younger
Mesozoic flora: U.S. Geol. Surv. Mon, 15, 377 p.,
180 Pis.

. 1899. Notes on the Lower Cretaceous plants

from the Hay Creek Coal Field, Crook County,
Wyoming, in Ward, L. F., The Cretaceous Forma-
tion of the Black Hills as indicated by the fossil
plants: U.S. Geol. Surv. 19th Ann. Rept., p. 645-
702.

Hall, J. W. 1963. Megaspores and other fossils in
the Dakota Formation (Cenomanian) of Iowa:
Pollen et Spores, v. 5, pt. 2. p. 425-443.

Harris, T. M. 1961. The Yorkshire Jurassic Flora, I.
Thallophyta-Pteridophyta : British Mus. (Nat.
Hi.st. ), 212 p., London.

Hawn, F. 1858. The Trias of Kansas: Trans. St.
Louis Acad. Sci., v. 1, p. 171-172.

Hedlund, R. W. 1966. Palynology of the Red Branch
Member of the Woodbine Formation ( Cenoman-
ian ) , Bryan County, Oklahoma : Oklahoma Geol.
Surv. Buil. 112. pp. 1-69.

Heer, O. 1859. Descriptions of fossil plants from No.
1 of the Nebraska section: Proc. Phil. Acad. Nat.
Sci., V. 10, p. 265-266.

. 1861. Prof. Heer's reply to Dr. Newberry on

the age of the Nebraska leaves: Amer. Jour. Sci.
and Arts, v. 31, p. 4.33-440.

. 1868. Die fossile flora der Polarlander: Flor.

Foss. Arct., V. 1, 189 p., .50 Pis.

. 1874. Die Kreideflora der arctischen zone:

Flor. Foss. Arct., v. 3, pt. 2, 1.38 p„ .38 Pis,

, 1882, Den erster theil der fossilen flora Gron-

lands: Flor. Foss. Arct., v. 6, pt. 2, 112 p., 47 Pis.
HoLLiCK, A. 1906. The Cretaceous flora of southern

New York and New England: U.S. Geol, Surv.

Mon. 50, 129 p. 40 Pis.
HoLTTUM, R. E. 1947. A revised classification of

leptosporangiate ferns: Juor. Linn. Soc. (Bot.)

London, v. 53, p. 123-158.
Jenney, W. p. 1899. Field observations in the Hay

Creek Coal Field, in Ward, L. F., The Cretaceous

formation of the Black Hills as indicated by the

fossil plants: U.S. Geol. Sur\. 19th Ann. Aept., p.

568-594.
JoNCMANs, W. J. 1957. Fossilium catalogus. II. Plan-

tae: Filicales. Pteridospermae, Cycadales, pt. 5, p.

247-366,
, 1959. Fossihum catalogus, II. Plantae: Fili-
cales, Pteridospermae, Cycadales, pt. 10, p. 751-848.
Le,squereux, L. 1868. On some Cretaceous fossil

plants from Nebra,ska: Amer. Jour. Sci. and Arts, v.

46. p. 91-10.5.

44

BiUGiiAM Young University Science Bulletin

. 1874. Contributions to the fossil flora of the

Western Territories, [)art I. The Cretaceous flora:
U.S. Ceol. Surv. Terr. Kept., v. 6, 136 p., 39 Pis.

. 1883. Contrihutioiis to the fossil flora of the

Western Territories, part III. The Cretaceoas and
Tertiary floras: U.S. Geol. Surv. Terr. Rept., v. 8,
283 p.,' 56 Pis.

1892. Tlie flora of the Dakota (;roup: U.S.

Geol. Surv. Mon. 17, 409 p.

MacNeal, D. L., 1858. The flora of the Upper Cre-
taceous Woodhine Sand in Denton County, Texas:
Phil. Acad. Nat. Sci. Mon. 10, 1.52 p.

Marcou, J. 1858. Letter on some points of the Cleol-
ogy of Texas, New Mexico, Kansas and Nebraska,
addressed to Messrs, F, B, Meek and F, V. Hayden:
16 p., Zurich.

. 1864. Une reconaissance geologi(jue an Ne-
braska: Soc. C;eol. France Bull. 21, p. 132-146.

Meek, F. B. and F. V. H.ayden. 1856. Description of
new species of gastropods from tlie Cretaceous for-
mations of Nel)raska: Proc. Phil. Acad. Nat .Sci., v.
8, p. 63-69.

. 1958. Heniarks on the Lower Cretaceous beds

of Kansas and Nebraska, togetlicr with descriptions
of some new species of Carboniferous fossils from
the valley of the Kansas River: Proc. Phil. Acad.
Nat. Sci., V. 10, p. 256-260.

. 1859. On the so-called Triassic rocks of Kan-

sa.s and Nebraska: Amer. Jour. Sci. and Arts, v. 27,
p. 31-35.

1861. Descriptions of new Lower Silurian

(Primordial), Jurassic, Cretaceous, and Tertiary
fossils collected in Nebraska Territorv. with some
remarks on the rocks from which they were obtain-
ed: Proc. Phil. Acad. Nat. Sci., v. 13, p. 417-432.

Newbehrv, J. S. 1868. The late extinct floras of North
.•Vmeric:!: .^nn. Neu York Lvceum Nat. Hist., v. 9,
p. 1-76,

. 1895, The flora of the Ambov Clavs: U,S,

Geol, Surv. Mon. 26, 258 p.

1898. The later extinct floras of Nortli

America: U.S. Geol. Surv. Mon. 35, 295 p.

PiEHCE, R. L. 1961. Lower Upper Cretaceous phuit
microfossils from Minnesota: Minnesota Geol. Surv.
Bull. 42, 86 p.

Repenning. C. a. and H. G. Page. 19.56. Late Cre-
taceous stratigraphy of Black Mesa, Navajo and
Hopi Indian Reservations, Arizona: Amer. Assoc.
Petroleum Gm\. Bull. \. 40, No. 2, p. 2.55-294.

RicHTEH, p. B. 1906. Beitrage zur flora der Unteni
Kreide (^uedlinburgs, I. Die gattnng Hau.smuitnid
Dunker unci eiuige seitenere pflanzenreste: 25 p.,
7 pLs., Leipzig,

RusHFORTii, S. R, and W. D. Tidewell. 1968. Notes
on the distribution and morphology of the fern
genus AstruloptiTis: Brigham ^'oung Uni\. Cleoi.
Stud., V. 15, pt. 1, p. 1(W-115.

HisiiioKTn, S. R. 1970. Notes on the fern family
M.itoniaceae from tlie western United States: Brig-
lunn Young Univ. Geol. Stud., v. 16, pt. 3, p. 1-.33.

Sapor ia, (;. de. 1873. Paleontologie Francaise on
Description des Fossiles de la France. Plantes Jur-
assitjues, I. E<|uisetacees. Characees, Fougers: 506
p., Paris.

ScHEHK, A, 187 L Beitrage znr flor.i der vorwclt. I\'.
Die flora der nordwestdentschen W'lMldenformation:
Palaeontographica, v. 19 B, p. 201-276.

ScHiMPKR, W . P. 1874. Traitc de Paleontologie Veg-
etale: v. 3, 896 p., Paris.

Sewari>, a. C. 1894. Catalogue of the Mesozoic
plants in the British Museum. The Wealden flora.
Part I.— Thallophyta-Ptcridophyta: 179 p., 11 Pis.,
London.

. 1910. Fossil Plants: \. 2, 624 p., Cambridge.

. 1927. Tile Cretaceous plant-bearing rocks of

western Greenland: Phil. Trans. Rov. Soc. London
(B), V. 215, p. 57-173.

SiNNOTT, I. W. and E. W. Bailey. 1915. Foliar evi-
dence as to the ancestory and early climatic en-
vironment of the angiosperms: Amer. [our. Bot.. v.
2, no. 1, p. 1-22, 4 Pis.

Stanton, T. W. 1905. The Morrison Formation and
its Relations with the Comanche Series and the Da-
kotii Formation: Jour. Geology, v, 13, p, 657-669,

Stephenson, L. W, 1952, Larger invertebrate fossils
of the Woodbine Formation (Cenomanian) of
Texas: U.S, Geol, Surv. Prof. Paper 242, 226 p..
51 PLs,

Stokes, W. L, 1948. (leologv of the Egnar-Gypsum
V'allev area, San Miguel and Montrose Counties.
Colorado: LI.S. Geol. Surv. Oil and Gas Invest. Pre-
lim. Map 93.

Tester, A. C. 1931. The Dakota Stage of the type
localitv: Iowa Geol. Surv. Ann. Rept. 35, p. 200-
332.

TnnvELL. W, D., S. R. Rushforth and J. L. Reveal.
1967. Astraloptcris, a new Cretaceous feni genus
from Ut:ili :uul Colorado: Brigham Young Univ.
Geol. Stn<l.. s. 14. p. 137-140.

TwENHOFEL, W. H. 1920. The Comanchean and Da-
kota Strata of Kansas: .Amer. |our. Sci. and .Arts., v.
49, p. 281-297.

Velenovsky, |. 1888. Die fame der Bohinischen
Kreitlformation: .Abh. K. Bohm. Ges. Wiss., Math-
Natw., ser. 7. v. 2. p. 1-.32, 6 Pis.

W'ahi), L. F. 1899. The Cretaceous formation of the
Black Hills as indicated bv the fossil plants: U.S.
Geol. Sur\. 19th Ann, Rept,, pt. 2. p. .527-.551,
702-810.

. 1995. Status 111 the Mesozoic Floras of the

United States; U.S. G.'ol. Sur\ . Mon. 48, 616 p„
119 Pis.

J

r?.

Brigham Young University haivm u

.UJMlVERSiTvirj

Science Bulletin

BOTANICAL AND PHYSIOGRAPHIC

RECONNAISSANCE

OF

NORTHERN BRITISH COLUMBIA

by

Stanley L. Welsh

and

J. Keith Rigby

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 4
SEPTEMBER 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany,

Brigham Young University, Provo, Utah

Members of the Editorial Board:

Vernon J. Tipton, Zoology
Ferron L. Anderson, Zoology
Joseph R. Murdock. Botany
Wilmer W. Tanner, Zoology

Ex officio Members:

A. Lester Allen, Dean, College of Biological and Agricultural Sciences
Ernest L. Olson, Chairman, University Publications

The Brigham Young University Science Bulletin, Biological Series, publishes acceptable
papers, particularly large manuscripts, on all phases of biology.

Separate numbers and back volumes can be purchased from Publication Sales, Brigham
Young University, Provo, Utah. All remittances should be made payable to Brigliam
Young University.

Orders and materials for library exchange should be directed to the Division of Gifts
and Exchange, Brigham Young University Library, Provo, Utah 84601.

Frontispiece. Picea glauca woods along the Narrows of the Grand Can\on of the Stikine River, approximately
.35 miles npstrcam from Telegraph Creek, British Colnmhia. In this seetion the .Stikine Ri\er has eut a spec-
tacular gorjje lip to 1000 feet deep through horizontal Tertiary rocks and underlying steeply folded Triassic
and Upper Paleozoic rocks. The riser has an elevation of approximatt'K 1,^00 feet here at approximatelv
.58°08' X; i:}0°20' W.

Brigham Young University
Science Bulletin

BOTANICAL AND PHYSIOGRAPHIC

RECONNAISSANCE

OF

NORTHERN BRITISH COLUMBIA

by

Stanley L. Welsh

and

J. Keith Rigby

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 4
SEPTEMBER 1971

TABLE OF CONTENTS

Page

INTRODUCTION 1

Location 1

Accessibility 1

PHYSICAL FEATURES 6

\ KGKTATION 12

Boreal forest 12

Alpine tundra 14

COLLECTING LOCALITIES 15

ANNOTATED LIST OF SPECIES 19

REFERENCES 49

BOTANICAL AND PHYSIOGRAPHIC RECONNAISSANCE
OF NORTHERN BRITISH COLUMBIA

bv

Stanley L. Welsh' and J. Keith Rigby-

ABSTRACT

The area of study is located in northern and the plant communities are enumerated. An

central British Columbia and includes the Sus- annotated list of 205 species, 11 subspecies, and

tut Basin and surrounding mountains. The phy- 35 varieties collected during the summer of

siography and geology of the region is described 1969 is included.

INTRODUCTION

Location

The Sustut Basin and surrounding mountains,
the primary area of collecting, are located in
north central British Columbia ( Fig. 1 ) in parts
of the McConnell Creek, Spatsizi, Toodoggone,
Dease Lake, and Cry Lake quadrangles. The
collecting area, approximately 150 miles long and
40 to 50 miles wide, extends northward from
approximately .56°39' to 58°30' North latitude,
and westNvard from 126°.30' to 1.30°30' West
longitude. Collections are representative of the
region between Thutade Lake and Sustut Peak,
northwestward to Dease Lake, in a belt north-
east of the Skeena River and Little Klappan
River valleys and southwest of the Omineca and
Thudaka Ranges in the headwaters of the Sti-
kine, Finlay, and Skeena Rivers (Fig. 2).

Prince George is 279 miles southeast of the
southern part of the basin and Hazelton is 120
miles to the south-southwest (Fig. 1). Tele-
graph Creek is 99 miles northwest, and Watson
Lake, Yukon, is 135 miles north of the northern
part of the area.

The Sustut Basin is situated along the Con-
tinental Divide between Arctic and Pacific
drainages. Much of the southern part of the
basin drains through the Finlay River into the
Peace River and the Arctic Mackenzie drainage.
The southwestern part drains into the Skeena
River, and the central and northwestern part
drains into the Stikine River in the Pacific drain-
age. The Kechika and Dease Rivers drain the

north and northeastern area into the Liard River
and Arctic drainage.

Accessibility

The area of primary concern is currently
accessible only by float planes to the several
lakes in the area and by helicopter. Even large
planes can work from Thutade, Tatlatui, Trygve,
Kitchener, Cold Fish, and Dease Lakes (Fig. 2).
Several of the smaller lakes are adequate for
landing by either a Beaver or Otter and could
function as a secondary campsite or base. A
short, graded but unsurfaced, air strip is avail-
able at the southeastern corner of the area in
the vicinity of Sustut Peak and Moose Valley.
The strip is currently maintained by the New
Wellington mine operation and is capable of
handling a Twin Beechcraft, although at certain
times of the year the strip is soft. A second dirt
strip is maintained at Hyland Post, in the central
part of the area, on the Spatsizi River. A third
was contemplated by 1970 at the northern end of
Cold Fish Lake in the northwestern part of the
basin.

Although no roads lead directly into the area,
several roads are constructed into relatively
nearby areas. An all-weather, maintained, grad-
ed road leads from Watson Lake and the Alaska
Highway south to Dease Lake and southwest to
Telegraph Creek. A spur road is currently under
construction which will connect Dease Lake and
Stewart. Tliis road is finished as far south as

'Department of Botany. Briphani Young University, Provo, Utah 84fiOI.
-Department of Geology. Brigham Young University. Provo. Utah 84601.

Bhigham Young University Science Bulletin

Kinaskan Lake and could be utilized by field
vehicles as far as Bob (^uinn Lake in 1969. A
winter caterpillar tractor trail is developed east-
ward from Dease Lake approximately 20 miles in
the vicinity of the east side of Dome Mountain
and could afford access to the northern part of
the area. A mine access road along the north side
of the Stikine River was under construction late

in the summer of 1969. It extends east from
near the Stikine River ferry to the southeastern
margin of the Three Sisters Peaks region, near
the northern edge of the Sustut Basin,

Topographic maps with a scale of 1:250,000
are available for the entire area. Relatively re-
cent geologic maps are available for the eastern
half of the McConnell Creek Quadrangle (Lord,

• Whi I e h o r se

AREA OF COLLECTION

Fort St John

Dawson

^

/

6

A

George

>

V

A

o

U.

o

^

/

o

<^

6 .

s

Vancouver

0

C I — I

100

200

300

400 miles

•"iG. 1. Map of British Columbia showing area of collection.

Biological Series, Vol. 14, No. 4 Reconnaissance of Nohthern British Columiua

KiG. 2. Map of area of roUection. in north central British Columbia.

Brigham Young University Science Bi'lletin

Tl'YA BASALT and overlying glacial debris and alUivium

Sl'STl'T t.ROUP

Well-bedded sandstone, conglomerate, miidstone with minor
interbedded dacite tuff, carbonaceous shale and coal, tiroup
may include some Qetaceous rocks in the lower part. General;
well-bedded and flat lying in plateaus, with ledges and slope*.

BOWSER GROUP

Interbedded gray -green and dark -gray, thin-bedded mudstone
and siltstone, gray- green, thin- to thick-bedded graywacke
sandstone, massive to thick-bedded gra)^^■ackc pebble- to
cobble-conglomerate, with minor thin limestone, coal, and
carbonaceous shale units.

TAKl.A (.ROL'P

\I?.5si\c gray-green, reddish brown, and maroon agglomerate
with -ocal lieds of graywacke and tuff with minor siliceous and
tuffaccous argillitf and basalt. Forms resistant crags and cliffs
in higher ranges of the region.

(:Af;HE CREEK GROUP

Interbedded chert, graywacke, tuff, basalt, and Ume^one in
somewhat cyclic sequence. Limestones arc locally fossiliferous
and cherty; grawackc and tuff units are siliceous. Entire unit
shows some metamorphism and structural complexity. Forms
shoulders below crags of Triassic volcanic section.

ASITKA CftOl'P

Reddish brown, pinkish itay and bright apple green tuff and
argillite with interbedded massive pinkish brown to maroon
rhyolite and andesite flows. Forms craggy xposures near the
\ alley margins.

KiG. .3. (k'dlogical section for Mortliciii Brits

Biological Series, Vol. 14, No. 4 Reconn.^iss.vnce of Nohthehn Biuti.sii Columuia

1948), and Cry Lake Quadrangle (Gabrielse,
1962) and the Dease Lake Qiwdrangle (Ga-
brielse and Souther, 1961). The Spatsizi and
Bow.ser Lake (|uadrangle.s are covered as part of
the older Project Stikine map by the Geological

Survey of Canada (1957). Aerial photographs,
with a scale of approximately 1:30,000 are avail-
able for most of the area of primary concern
through offices of the Geological Survey of Can-
ada in Calgary.

Fic. 4. C.fologiL.il L.iiii[) at Goldfish Laki_. hm Ih.hIi ni.r.il of high .stand of the lake, in a protected fore
collecting locality 6, at approximately .57°39' N; 128°44' W, at an elevation of appro.ximately 3820
Mi.xed woodland dominated hy Vinus contorta, Picea glauca, and Betula ghindiilosu .

:st at
feet.

Bhk.ii AM VoiNt; University .Science Bulletin

PHYSICAL FEATURES

All of the collecting area is included in the
Interior Ranges of liritish Columbia and in the
Interior Plateau section in the Stikine River
region along the east side of the Bowser Basin,
60 miles southwest of the Rocky Mountain
Trench. In broad terms the entire region can be
subdivided into: 1 ) the high rugged topography
of the Sustut belt; 2) the low bordering lowland
river vallevs along the southwestern, northern
and northeastern sides of the plateaus; 3) the
horizontally bedded plateau belt from Thutade
Lake northwestward into the Spatsizi Plateau; 4)
folded Sustut and Bowser beds and the intensely
folded high mountains and glaciated areas east
of the Skeena River Valley; 5) the Cry Lake
hills to the north; 6) the uplands and highlands
to the northeast of the basin; 7) the Eaglenest
Range uplands; and 8) Three Sisters Range.

The Sustut Peak highlands and adjacent Asit-
ka uplands are a region of relatively mountain-

ous terrain that extends from Niven Peak south-
east beyond Sustut Peak in the region of out-
cropping Paleozoic and Triassic to upper Jurassic
volcanic rocks (Lord, 1948). The belt of high-
lands is dominated by Sustut Peak which rises
to an elevation of 8100 feet, 4000 feet above the
valley floor of Moose Valley on the northeast and
Sustut Lake directlv to the east. Savage Peak,
with an elevation of 7600 feet, and Dewar Peak,
with an elevation of 7350 feet, are northern parts
of the highlands. All of the major high peaks
are horns and are connected to one another bv
arete ridges. The entire area has been intensely
glaciated, and active glaciers still persist on the
northeast side of Savage and Dewar Peaks and
around the east and north sides of Sustut Peak.
The t^astem part of the range is carved in large
part from folded Late Paleozoic volcanic and
carbonate rocks that produce a relatively sub-
dued topography in contrast to the high peaks

^^

1M

m\

Flc;. ... 1,.,.,^ ,.. ,i,, ,,wi,,i> ,,i tin (....111 1 i^li l..iki- t.iinii. Liii.iliU (.. ,ii .>, ■>:! \. i_."i 4-1 \\ , ,it .ippiuMiii.iU-ly
3800 feet eIe\ation, Boreal forest of Pinus contorta and Picea glauca, \%ith tents surrounded b\ Su/i.v species
iiiid Betula slandiilosa.

Biological Series, Vol. 14, No. 4 Reconnaissance ok Nohihern British Columbia

Fic. 6. West across Goldfish Lake from the campsite at 57°39' N; 128°44' W. toward the south end of the
Eaglenest Range, with well-developed Boreal F"orest in the foreground and along the low flanks of the
volcanic range in the backgroimd. The camp is at ajipniximately 3800 feet and the range crest at approxi-
matel) 7500 feet. Boreal Forest of ?kca glaucu, Ah'iex lasincarpa, and Pimis conturta. The lake shore is
fringed with Betuki glandulosa and Salix species.

and ragged massive cliffs carved on the thick-
bedded Triassic and Jurassic agglomerate along
the west side of the belt. Toward the northwest
Sustut Peak highlands are separated from the
southeastern plateaus by the broad Niven River
vallev which separates the folded Paleozoic Tri-
assic and Jurassic rocks from the nearly flat-
King Cretaceous and Tertiary Sustut rocks to the
northwest (Fig. 3). Broad Moose Valley and

Asitka River valley on the east separate the
Sustut Peak area from the McConnell Range and
other mountainous areas of the Omineca batho-
lithic areas to the northwest.

Broad flat-topped, but deeply indented, glaci-
ated plateaus have developed along the north-
eastern side of Sustut Basin from near Thutade
Lake, northwestward beyond the Spatsizi Pla-
teau, to the Stikine Valley in the vicinity of the

Q Bhk.ham Voi-Nt; University Science Bulletin

Three Sisters Range, a distance ot approximately River region southeast ot Thutade Lake north-

100 miles. The plateau section is carved on hori- westward across the entire Spatsizi Plateau. The

zontal or very nearly horizontal Sustut beds characteristic topographic development of the

which form relatively open, simple-contoured horizontal beds is perhaps best sho\\Ti in the

topography and extends from Mt. Forest-Niven area between Tudatui and Thutade Lakes in the

\h

\

\

J

%^

■■\>

t

I "10

7. Southwest along Buckinghorsc Creek near its junction with the Spatsizi River. Outcrops of Jurassic
canics are in the foreground with spruce woods on the talus of the overlying Bowser formation at approxim
.57°27' N; 128°35' W. Picea glauca woodland, with Populus tremuloides, lietulti glamlulosa. an '
species in the openings.

vol-
atelv
S«/iA:

Biological Sehies. \'ol. 14, Xo. 4 Rlconnaissance of Northern British Columhla

vicinit)' of Tabletop Mountain and Mt. Jorgen-
sen (Fig. 25) where the alternating resistant
and nonrcsistant beds form almost lake terrace-
like topographic features. The same erosional
pattern can be seen from southeast of Thutade
Lake on Thutade Peak and on Mt. Forest and to
the Stikine River Valley and Idozadelly Moun-
tain and the Spatsizi Plateau in the bend of the
Stikine River. The entire flat upland of the
plateau rises to an elevation of 6.500 to 6800
feet, approximatelv 3000 to 4000 feet above the
vallev floor lowlands. It is a region of broad
open summits and relatively gentlv sloping walls
to broad, flat, river valleys. In general, it is a
region where lake development is not extensive.
The northeastern tip of Thutade, Tatlatui, Kitch-
ener and Laslui Lakes extend into the western
part of the plateaus, but broad valleys of these
lakes and their drainages dissect the plateau
surface.

.\n eastern lowland belt separates the hori-
zontal plateau section, where Tertiary and Cre-
taceous horizontal rocks dominate, from the dip-
ping Triassic and Jurassic rocks along the flanks

of the Omineca and Cassiar batholith belt to the
east and northeast. A nearly continuous, low,
broad valley extends southeast from near Dease
Lake along tributaries and main valley of the
Stikine River into tlie vicinity of Caribou Hide,
and then on the Toodoggone River and continu-
ing southeastward into the headwaters of the
Finlay River and Moose Valley. These broad,
low valleys have an elevation of from 3000 to
4500 and range up to 15 to 20 miles wide. The
various drainages are separated by low, broad
passes such as Matzantan Pass between the Too-
doggone and Stikine rivers; Lawyer's Pass be-
tween the Toodoggone and Chapea rivers, the
latter a tributary near the head of the Finlay
River at Thutade Lake. A broad pass in the
middle of Moose Vallev separates the Finlay
River drainage from that of the Skeena and the
Sustut Rivers and is almost unnoticed when fly-
ing over the region in the southwestern end of
the valley.

A western lowland along the Klappan and the
Little Klappan and the Skeena rivers continues
N .30° W, almost along strike near the south-

^i.

' «*•

Fig. a. Boreal Forest on the north think ot the deep Grand Canyon of the Stikine Hiver at approximatelv .SS^OO'
N; 1.30°2r W, abo\e the junction of the Tanzilla Hi\er with the .Stikine River. Prominent ledges are of
cont;lomeratie sandstone separated bv broad shale belts along the shoulder of the canyon at an elevation of
appro.\imately 2.500 feet. .Mi.xed Picea glauca-Popidus trcmuloides woodland.

10

Bhk;ha.m Vouno UNivtHsnv Science Bulletin

Fic. 9. Northwest to\v;iril the main massif of Savage Peak, elevation 76(X), at 56°39' \; 126°42' \V, eroded from
massive agglomerate of the lower part of the Takla CIroiip. The roeks whieh hold up Savage Peak are
essentially of the same unit as that uhieh holds up Sustut Peak to the southeast. Savage Peak is here proposed
for Douglas Savage who has spent manv years in the area and who was killed in a helicopter crash just east
of the peak in Moose Valley in the sununer of 1969. X'allev boKom witli meadow and Boreal Forest extending
up slope to alpine lundra.

western border ot the area of primary concern.
These valleys are 4 to 5 miles wide, broad, with
steep walls and are intensely glaciated. They
separate the interior of the Bowser Basin from
the folded Bowser beds to the northeast. Both
the Skccna and Klappan valleys have elevations
between 3000 and 4(XM) feet and bridge over a
broad open pass at almost midlcnglh along the
area of concern near Mt. CJunnanoot. The

Skeena River drains to thi' southwest into the
Pacific, and the Klappan and Little Klappan
River drain iiltimateK into the Stikine River near
the norlliwesteni edge of tlie acreage and then
into the Pacific.

West and south of the hori/.ontallv bedded
plateaus of the Sustut Basin is a mountainous
area carved, in large part, in complexh' folded
and faulted Jurassic Bowser (Jroup and (^reta-

Biological Series, \'ol. 14, NO. 4 Reconnaissance oe- \ohtiiehn Bihtish Columbia

11

ceous Tertiary Sustut beds. The boundary be-
tween these folded rocks and the relatively
simple to horizontal Sustut beds of the basin is
along a prominent hogback or a line almost due
northwest from the middle of Thutade Lake,
northwest across the tip of Tatlatui and Kitche-
ner lakes, along the west base of the Brothers
Peak and northwest across Laslui Lake toward
the western Spatsizi Plateau east of Goldfish
Lake. Southwest of this hogback both the Bow-
ser and Sustut rocks are comple.xlv folded in a
series of asymmetric anticlines and synclines.
The folded beds and the relatively high eleva-
tion produce rather spectacular, sharp, serrated
peaks, many of which are now glaciated, char-
acterized by such heights as Kama Peak, with an
elevation of 7105 feet. Chipmunk Peak with an
elevation of approximately 71.35 feet, and Mel-
lanistic Peak, with an elevation of 7110 feet. All
of these rise 4000 to 4500 feet above the general
low valley along the Skeena River to the west

and above the general elevation of 6000 to 6500
feet ot the plateau regions to the northeast.

Three Sisters Range is a complex igneous in-
trusive mass in Triassic and Jurassic volcanic
rocks. It occurs near the northwest border of the
area a few miles southeast of Dease Lake. In
this general area the rock dip regularly from
the intrusive mass near the Three Sisters Peak
into the low country along the southeast bor-
der of Dease Lake and south and eastward
into the Stikine River Valley and its tributaries.
Peaks in the range rise to 7565 feet in the north-
eastern part. Three Sisters Range is nearly cir-
cular, 15 to 20 miles in diameter, and is the south-
westernmost major promontory of igneous activi-
ty associated with the Omineca-Cassiar batholith
sequence. The Three Sisters rises above the
McBride River hills to the east and the Hotailuh
Range to the west. These latter areas are of rela-
tively moderate relief and are in folded Triassic
rocks, in the main.

Fig. 10. Meadow, shmbland, and Boreal Poorest along Spat.sizi V'allev, at lat. 57°34' N. long, 128°33' W, elevation
3700".

12

Bricham Vounc University Science Bulletin

The Eaglenest Range highland is carved in
folded Jurassic (?) volcanic rocks hut is distinc-
tive from less prominent peaks carved in the
folded Jurassic Bowser beds because of the spec-
tacular, sharp, serrated ridges and peaks which
the massive volcanic rocks have produced. The
Eaglenest Range rises above broad Goldfish Val-
ley, which separates it from the Spatsizi Plateau,
to the northwest, and generally above the ranges
of folded Bowser rocks to the west and south-
west. Like other ranges, glaciated valleys are
often connected by low passes through the mar-
gin of the range or near the valley floor of
adjacent lowlands. Peaks, such as Nation Peak,
with an elevation of 7741 feet; Mt. Will, with an
elevation of approximately 7600 feet, and Cart-
mell Mountain, with an elevation of 7135 feet,
rise three to four thousand feet above adjacent
valleys.

Cry Lake Hills, at the northern margin of
the area studied, are carved in metamorphosed
Paleozoic and Mesozoic rocks and fonn rather
subdued to rounded ranges and hills, all of
which have been glaciated and rounded beneath
ice masses (Gabrielse, 1962). The Cry Lake
Hills are relatively low features, with a relief of
appro.xiniatelv 2000 feet in the main, and occur
in front ot the main Stikine Ranges to the north-
east in the belt of Omineca and Cassiar batholith
development. These hills are more or less topo-
graphic continuations of the Mt. Dease and the
McConnell Range uplands which rise onlv mod-
erately above the bordering Stikine River-
Sturdee River-Moose Valley lowlands. The Cry
Lake Hills and the other southeastward continu-
ations show on the topographic maps as general-
ly rounded topography rather than as sharpiv
ridged and jagged topography of adjacent high-
lands to the northeast.

VEGETATION

Boreal Forest

The Boreal Forest, which occupies most of
the area of northern British Columbia, is domi-
nated by three tree species (Figs. 4, 5, and 6).
The most abundant of these is the white spruce,
Picea glatica. Next in importance is the alpine
fir, Abies lasiocarpa. Third in importance nu-
merically is the lodgepole pine, Pinus contorto.
There are two other tree species which make up
a smaller part of the total Boreal Forest. They
are the aspen, Populus treimiloides, and the Cot-
tonwood, P. balsdinifeni. White spruce is abun-
dant on alluvial gravels in valley bottoms, and
on glacial moraine and talus. Alpine fir occurs
as individual trees and in small groups within
the spruce-dominated woods in the lower eleva-
tions. Upwards, the alpine fir is more abundant,
and in timberline situations alpine fir is the
dominant conifer. Lodgepole pine occurs mostiv
on ridges of glacial moraine, which are evidently
better drained than the surrounding regions. In
some sites, such as north of New Wellington
mining camp, lodgepole pine occur in almost
pure stands.

Aspen occurs in parkland areas, usually above
the spruce forest, but also in openings in spruce
woodlands, commonly on slopes ( Figs. 7 and 8 ) .

Major components in the understory of the
spruce-fir-pine w(K)ds include species of willow
{Salix spp. ), and the almost onnnpresent dwarf
birch, Betula glandulosa. Other components of
the understory include:

Linnaea horealis, Vihtirnum echile, Shep-
herdia canadeiviK, Lonkcra iniolucrata, Cornus
canadensis, and Empetrtnn nigrum.

The ground layer in the woodlands consists
of sparse to dense vegetation. In some areas
there are three species of violet growing sym-
patrically, two of which are common ( Viola
adunca, V. glabella) and one uncommon (V.
umfolia). Other plants in the ground layer
include:

Lijcopoditnn annotinum, Lycopodium com-
panatum. Lt/cupodiuin alpiniini. E(piisetum ar-
veiue. Anemone ricliard.wnii. and Listera cor-
data.

During the second week of June 1969, the
spores were released from Lijcopodium annoti-
num in large numbers. Spores covered the en-
tire surface of Thutade Lake. Wind and water
currents concentrated the spores into long yel-
lowish streamers on the lake surface, and finally
onshore breezes stacked windrows of spores
along the lake shore. Some of the windrows
were to one-half an inch in depth.

Parkland meadows are present in valley bot-
toms in the Boreal Forest. Willows and dwarf
birch grow along the drainages in the meadow-
lands (Figs. 9 and 10). The meadows proper
are occupied in wet sites b\- the eottongrass,
Eriophonim iingustifolitim. and the sedge, Carex
aifiiatihs. Drier sites in the meadows are domi-
nated by species of Poa. Festtica altaiea, and
Calamagrostis purpurascens.

BiOLor.icAi. Series, Vol. 14, No. 4 RECoNN.\i.ss.\NrE of Nortiiehn British Columrlv

13

Fig. 11. Steep monoL-line and flat-King portion of the mkklle and upper part of the Sustut Formation 4 miles
northwest of Thutade Lake at approximately 56°53' N; 127°11' W. Resistant beds are conglomerate and
separated by thick units of softer silty mudstone and shale.

Riparian and Palustrine vegetation consists
of essentially the same species which occur in
the Boreal Forest. However, on alluvial gravels
along stream courses the cottonwoods {Populus
})a]samifera) and willows {Salix .spp. ) and alder
( Alnus cri.spa ) become more dominant.

A steep south-facing slope about two miles
southwest of the southern end of Thutade Lake
is worthy of mention. This region is occupied
by up and down slope stripes of trees which
alternate with similarly oriented stripes dominat-
ed bv herbs and shrubs. The herb and shrub
stripes are apparently due to snow slides which
sweep away tree growth and allow herbs and
shrubs to dominate. On June 14 at about (S:()0
p.m., we landed the helicopter on a small out-
crop in one of the slide areas, and in about an
hour we were able to collect some 39 species.
The shrubby \egetation consists of Vil)unuim
edtilc. Ril>es <^landulosiim, and Sambticus riice-
mosa. Important herbaceous species include:

Valeriana sitchensis, Aquilegia fonnosa, Hera-
cleum hinatum, Hackelia jessicae. Geranium eri-
antlnim, Senecio triangularis, Castilleja unalas-
chcensis, Veratrum eschscholtzii, Myosotis syl-
vatica, PecUcularis bracteosa, Lupinus nootkaten-
sis, and Urtica clioica.

Principal grasses on the slope include species
of Poa and Festuca altaica. The habitat and
species list is reminiscent of open slopes in Little
Susitna Canyon in south central Alaska.

The upper elevational limit of the Boreal
Forest varies considerably, but most of the forest
occurs below the 5,000 foot line. However, there
are trees growing to about the 6,000 foot line
in some regions. In these sites, the trees occur
singly, in small clumps, or in long lines trending
upward along drainages or slopes ( Figs. 11 and
12). In other sites the tree line is truncated
along the apparent upper limit of glacier ice of
the Pleistocene glaciation (Figs. 13 and 14).
The forest is best developed on glacial debris

14

BnicHAM Young University Science Bulletin

along the slopes and on moraines and allnvinni
in the valleys. The tree species above about
5000 feet in elevation are limited to spruce and
alpine fir. The trees take on a kniminholz ap-
pearance, spreading out horizontally along the
slopes above the tree phases more typical of the
forest at lower elevation (Figs. 15 and 16). The
krummholz type occurs in patches to several
acres in extent, mostly above areas affected by
glacier ice, and it seems probable that some of
these may have persisted throughout the Pleisto-
cene glaciation and have provided propagules
for reforestation of the valleys following the
retreat of glaciers. The knmimholz interfingers
upwards with alpine timdra ( Figs. 17, 18, 19,
and 20). In its upper limits, the alpine fir
krummholz grows with Plit/Uodoce empetri-
formis, P. glatululiflora, and Cassiope ietragona.

Alpine Tundra

Alpine timdra is well developed in the moun-
tains of northern British Columbia. On alpine
slopes and rounded ridge tops the tundra is

dominated by Dnjas integrifolia, Festuca aJtaica,
Carex sp., Cassiope tetragumi, Salix arctica, S.
glauca, S. reticulata, and S. stolonifeni ( Figs. 21,
22, 23, 24, and 25). On ridges, beghming at
about 6000 feet elevation, and steep slopes up-
wards from that elevation, the tundra consists
of sparse vegetation, dominated bv clumps of
Drijas integrifolia, PotentiJhi diversifolia, P. Inj-
parctica, P. uniflora, Lupinus arcticus, Oxijtropis
campestris, Luzula confusa, L. nivalis, and Carex
sp. (Figs. 26, 27, and 28). In some areas the
bearberry, Arctostaplujios alpina is present near
the upper limits of alpine tundra. Between 6000
and 7000 feet elevation, the open spaces between
plants on rock outcrops and roc-kv ridge crests,
crustose lichens fonn the most abundant plant
cover (Figs. 29, 30, 31). These semibarren
lichen dominated lands are herein called fell
fields. Important species of flowering plants in
fell fields include:

Ranunculus eschscholtzii, PotcntiUa diversi-
folia, P. hijparctica, Sihbaldia procundtens,
Hierochloe alpina, Pedicularis stidetica, Anten-

FiG. 12. Xortlnvest to the iiorflieast spur of Siistiit lV:ik at 56°3T N; 126°.35' W. 'Ilie relatively smooth, even
Crestline of the ridge is in the sandstone seciuenee near the base of the Triassie Takia Group. The deep
snow-filled eol near the right margin is at the approximate l)Oundar\ between Triassie and Permian rocks.
The ridge crest is at an <'le\ation of approxim.itely 7000 feet. .Mpiue tundra grading downward with knmim-

holz and Boreal Forest.

Biological Series. Vol. 14, No. 4 Reconn.\iss.\nce of Northehn Bbitish Columhia

15

naria monocephala, Antennaria (ilpina, and S«/i.v

glauca.

A particularly inhospitable habitat wa,s en-
countered on an outcrop of .silt.stone and coal
beds in a saddle about 5 miles southeast of the
south end of Thutade Lake ( Fig. 32). The sur-

face is in large part barren of vegetation, but
plants, Draha incerta and Carex podocarpa, oc-
cur widely spaced on the soft surface. This is
the only locality wherein AreruirM sajanensis
was collected, growing directly on the carbona-
ceous surface.

COLLECTING LOCALITIES

Plants were collected at localities throughout
the Sustut Basin during much of the growing
season (Fig. 33). Collecting was initiated on

June 7 and the final collections for the season
were taken on August 13. All major plant com-
munities were sampled, but the list of specimens

Fic. 13. Dfwar Peak at .56°43' .\; 126°.50' \V. as seen from the southeast. Dewar Peak is composed of pinkish
gray rhyolite and andesite in the upper part of the Asitka Group which also forms the ragged rocky exposures
on the skyline to the right. Locality 34 is in the prominent saddle on the skyline to the immediate left of
Dewar Peak and is at an elevation of approximateK 6400 feet. Woodland dominated by Picca glauca and
Ahies hmocurpa grading upwards with krummholz of Ahics la.siocarpa and alpine tundra and fell fields.

16

Bricham Vouno University Science Bulletin

Fig. 14. Southwest across Goldfish Lake Valley to the prominent serrated massive volcanic rocks of the Eagle-
nest Range. The snow line is at about the upper level of the Boreal Forest. The snow is from the early
snowstorms which came the 10th to 12th of August and blanketed the uplands. The snow line corresponds
with the appro.ximate upper limit of trees and to near the elevation of ma.vimum elevation of valley glaciers
during major Pleistocene glaciations.

collected is not represented as being exhaustive
for the region. Many additional .species will un-
doubtedly be added as the flora of this magnifi-
cent region becomes better known. The nature
of the Sustut Basin can be seen from the aerial
photographs of selected areas (Figs. 34-42).

1. Ironbridge. South bank of the Tanzilla River, 8
miles south of Dease Lake village, at Ironbridge,
along Dease Lake-Stewart Highway, 58°2r N;
129°52' W, at 3090 feet elevation, on river terrace
gravels.

2. Letain Lake. Asbestos prospect, northeast of Letain
Lake, and ca 7 miles west of King Mt., at 58°20'
N; 128°45' W, at 6000 feet elevation, in Alpine
tundra, on weathered serpentine intrusive.

.3. Kehlechoa River. Ridge west of Kehechoa Ri\er, at
58°12' N; 128°4.5' W. at 5000 feet elevation, in
alpine tundra, on green micaceous scfiists.

4. Cold Fish Lake, north shore. North end of C^oid
Fish Lake, at 57°42' N; 127°50' W, at 3800 feet
elevation, in Boreal Forest, on glacial moraine.

5. Cold Fish Lake, camp ridge. Cold Fish Lake,
south end of camp ridge, at .57°40' N; 128°45' W,
at 3800 feet ele\ation, in open woods on glacial
moraine.

6. Cold Fish Lake. South end of Cold I'ish L.ike, at

57°34' N; 128°44' W, at 3800 feet elevation, in
Boreal Forest, on glacial moraine and beach gravel.

7. Chukachida River. South of the mouth of Chuka-
chida River, at 57°40' N; 127°33' \V, at 5200 feet
elevation, in tundra, on Jurassic volcanic rocks.

8. S.W. Mt. Will. Pass Lakes, ca 6 miles southwest of
Mt, Will, at 57°29' N; 128°53' W, at 6500 feet
elevation, in Alpine tundra on talus and outwash
of Bowser Formation.

9. Griffith Greek. Head of Griffith Creek, on Bowser
Formation, at 7000 feet elevation, at 57°28' N;
128°28' W. Alpine tundra.

10. Caribou Hide. At Caribou Hide, along the Stikine
River at 57°27' N; 127°34' W, at 3700 feet eleva-
tion, in open woods, on river fill.

U. Tuaton Lake. West side of Tuaton Lake, at 57°17'
N; 128°06' W, at 6000 feet elev.ition. in alpine
tundra, on Bowser Formation.

12. Stalk Ridge. Northeast Stalk Ridge, at 57°09' N;
127°37' \\', at 6000 feet elevation, on siliceous
conglomerates, of Bowser Formation, in •■Mpinc
tundra.

13. Stalk Peak. Northwest spur of Stalk Peak, at
.57°08' N; 127°44' W, at 6000 feet ele\atioii, in
alpine tundra, on liarrcn shale slope of Bowser
Formation.

14. Stalk Lakes. Fast shore at .south end of llie north-

Biological Series, \'ol. 14, No. 4 Reconnaissance of Northern British Columhia

17

enimost lake of Stalk Lakes, at 57°07' N; 127°36'
W, at 4500 feet ele\'ation. in meadows, on alhnium.

15. N. Kitchener Lake. Ca 4 miles north of Kitchener
Lake, at 57°07' N; 127°32' W, at 5000 feet eleva-
tion, in alpine tundra on alluvium.

16. Kitchener Crag. Rockv, bedrock ridge crest, im-
mediatelv west of Kitchener Crag, southwest of
Kitchener Lake, at 57°01' N; 127°26' W, at 6500
feet elevation, in alpine tundra on Bowser Forma-
tion.

17. Kitchener Lake. Along south shore of Kitchener
Lake, at 57°11' N; 126°57' W, at 5000 feet eleva-
tion, in alpine tundra on Triassic marble.

19. N. tip Thutade Lake. Northwest shore, north tip
of Thutade Lake, at 57°08' N; 126°54' W, at 3700
feet elevation, in Boreal Forest, on granite bedrock.

20. N.W. Tatlatui Lake. Ca 3 miles northwest of
Tatlatui Lake, at 56°58' N; I27°23' W, at 6000
feet elevation, in alpine tiindra on Bowser Forma-
tion.

21. Skeena River Vallev. Tatlatui Range, ca 6 miles

south and 60° west of Ahna Peak, at 56°40' N;
127°38' W, on slope above Skeena River Valley,
at 6200 feet elevation, ca 17 miles southwest of
Thutade Lake, in Bowser beds or Takla Formation.

22. Head Thutade Creek. Tatlatui Range, in pass be-
tween liead of Thutade Creek and unnamed tribu-
tary of Skeena River, ca 12 miles southwest of
Thutade Lake, at 56°44' N; 127°33' W, at 5600
feet elevation, in a fell field, on Asitka slate.

2,3. Thutade Creek. Near head of Thutade Creek, ca
5 miles southwest of Thutade Lake, at 56°47' N;
127°18' W, at ca 3800 feet elevation, on stream
gravels, in willow heathland.

24. Tatlatui Lake. Tatlatui Lake shore, at 56°54' N;
127°24' W. at 4080 feet elevation, in beach de-
posits and morainic gravels, in mixed woodland.

25. S.W. Thutade Lake. Steep, southeast-facing slope,
ca 2 miles due west of the southwest comer of
Thutade Lake, at 56°47' N; 127°17' W, at 4500
feet elevation, on middle Bowser Formation.

26. Thutade Lake camp. West shore of Thutade Lake,

-I
nil

,msi&

Fig. 15. .\ortii to tiic honndary tank zone at the west edge ni tin' Sustut Basiu exposed m tin- ndgc crest
immediately north of Thutade Lake at 56°50' N; 127° 12' W. Minor affect of differences in lithology of
Sustut Formation and the Bowser group shows in nearly uniform plant patterns. The alternating light-banded
beds on the left are overturned, and sheared middle beds of the Sustut Formation and the more ma.ssive,
nearK horizontal beds exposed on the left are in che Jurassic Bowser Formation, here composed of mudstone
and minor beds of conglomerate. Krummholz of Abk'.i lasiocarpa and open-grass and sedge-covered slopes.

18

Bhigham Young University Science Bulletin

Fig. 16. Northeast .spur of Niven Peak at 56°56' N; 126°52' W, as s, , n I mm tin- east. Tlir three low s.iwtooth-
.shaped ridges along the spur near the eenter of tlie photograph are held up by fossihferous hmestone at the
top of tlie Caehe Creek set|uence and are at an elevation of approximately 7000 feet, 20(M) feet above the
valley floor. The general level spur toward the left is held up by graywaeke sandstone at the base of the
Triassie Takia sequence. The ragged ledge-forming exposures in the center and right are on tlie upper part
of the volcanic Asitka seciuence near Locality 3.3. The steep, apparentis barren, slopes in center background
are clothed with grassy alpine tundra dominated by Festuca altaica.

near south end, in mixed lodgepole pine, white
spnice, alpine fir woodland, at .56°48' N; 127° 12'
W, at 3625 feet elevation, on morainic gravels.

27. Thutade Mt. On mountain behind camp at Thutade
Lake, at 56°49' N; 127°11' \V, at .5000-6000 feet
elevation, in alpine tundra and steep grassy slopes,
on Bowser Formation.

28. .South Pass Peak. Ca L5 miles northwest of South
Pass Peak, ca 7 miles south-southwest of "Thutade
Village," at ,56°42' N; 127°0y' \V, at 4300 feel
elevation, in open woods, on alhi\ium.

29. Coal beds. In saddle of niovuitains. ca .5 miles
southeast of south end of Thutade Lake, at 56°45'
N; 127°05' W, at ca 5000 feet elevation, in alpine
tundra and barrens on Tertiary Sustut coal beds.

30. Mt. Jorgensen. On Mt. [orgensen. ca 12 miles
northeast of •Thutade Village," at 5e°54' N; 126°
56' W, at .5200 feet elevation, in alpine tundra on
Sustut Formation.

31. Firesteel Kiver. Ca 7 miles east of Kitchener Lake,

at 57°03' N; 127°10' W, 4000 feet elevation, in
Boreal Forest on alluvium.

32. Niven Creek. Tributary of Ni\en Creek, ca 3 miles
east-southeast of Dewar Peak, at 56°42' N: 126°52'
W, at 4500 feet eknation. in open willow-meadow
lands surroimded b\ wliitc spruce woods, on allu-
vium.

33. W. and S.W. Dewar Peak. Ridge crests, west and
southwest of Dewar Peak, at ca 56°42' N; 126°50'
W, at ca 5500 feet elevation, in alpine tundra in
I'pper Takla Croup volcanic rocks.

.34. Dewar Peak vicinity. Alpine tundra, at .56°45' N;

126°4.5' \V. at 6000-6500 feet eIe\ation, on lavas

of the Asitka Croup.
35. De«ar Peak NNE. Ca 3 miles NNF Deu.ir Peak,

at 6125 feet elevation on Tertiary Sustut Formation,

at .56°46' N; 126°45' W. Alpine tundra and fell

fields.
.36. 'I'home Lake. Marshy east shore of Thome Lake,

at ,56°50' N; 126°42' \V, at 3900 feet elevation.

Marshy east shore, on allu\iuin.

Hioi.ocicAL Sehies, Vol. 14, No. 4 Re(:onn.\iss.\nc;e of Nohihehn Bkitksh Columdi.v

19

38.

39.

Mt. Savage. Ma.ssif .south of Moo.sevale Creek, ea
3 miles .southeast of Dewar Peak, at 56°40' N;
126°45' VV, at 6000 feet elevation, in krummholz
and alpine tundra, on lavas of the Takla Group.
New Wellington. At New Wellington mining camp,
in Moose \'allev at the mouth of Moosevale Creek,
at 56°44' N; 126°37' W, at ca 4000 feet elevation,
in wet to dry meadow, stream bank, and open
lodgepole pine woods on allmium.
Rognaas Peak. Northeast spur of Rognaas Peak, at

40.

41.

57°09' N; 127°04' W. at 4700 feet elevation, in
open woods, on cherty nihhle at Triassic Takla
Formation.

Trygve Lake. North shore west end, on glacial
moraine, at 56°59' N; 127°30' W, at 4570 feet ele-
vation. Heathland on Pleistocene glacial moraine.
Lawyer Pass. North side of hill, 4 miles east of
Lawyer Pass, at 57°18' N; 127°13' W, at 6000
feet elevation, in deep soils over Jurassic Takla
Group ( ? ) volcanics. in alpine tundra.

ANNOTATED LIST OF SPECIES

The following list of species was collected in
northern British Columbia during the summer
of 1969. Two main collections are included; one
by S. L. Welsh and J. K. Rigby taken during the
first t\\o weeks of June, and a second collected
by J. K. Rigby and G. Cuddy throughout the

growing season of 1969. In order to save space,
the collection localities have been numbered and
described separately from the list of species. The
names of the collectors are abbreviated; with
WR standing for Welsh and Rigby, and RC
for Rigby and Cuddy.

Kic. 17. Northwest along the eastern ba.se of the Niven Peak massif at 56°46' N; 126°5r W. Light colored
tuffaceous and rhyolitic units are a distinctive feature in the central part of the photograph beyond the heli-
copter. Well-bedded rocks in the middle distance are on the southeast spur of Forrest Mountain to the north-
east of Niven River. Peaks along the skyline are in the vicinity of Fredrickson Peak and other peaks to the
northeast to Thutadc Lake and are composed of Takla volcanics cut by Ominica intrusive granodiorite. Thome
Lake, Locality .36, occurs in the vallev between the batholithic belt and Mount Forrest. Locality 38, at New
WelliiigliiM. is in Moose Valley a short distance to the right of the photograpli. Ridge crest dominated by
FotcntilUi hypurctica, Lupinus arcticus, Dnjas integiijolUi, Antennaria monoccphala, and SelagincHti sihirica.

20

Hui(;iiAM ^oi'Nc Univehsitv Science Bulletin

Ku.. 18. Northwest toward tlif northeast spur of Niveii IVak with Cache Creek roeks lorming most ot the central
exposures and Asitka roeks forming the Hght colored to brilliantly colored roeks along the right part of the
photograph at 5(i°47' N; 126°50' W. Collections from Locality 35, Dewar Peak north northeast, were collected
from the saddle and cliff zone separating the Hght and dark-colored rocks at the right center skyline. Sparse
alpine tundra dominated In cnistose lichens in foreground, with krunimholz and tunilra covered slopes in
center and on ridge in background.

Collection localities arc luiiiibcrcd from
northern portion.s of the region southward and
are not in order a.s to date of collection.s.

In the checklist, the subdivision.s are ar-
ranged in phylogenetic setjuence, but families,
genera, species, and infraspecific taxa are in
alphabetical order. A summary of the number
of taxa is presented at the end of the checklist
(Fig. .■3.3). All specimens are deposited in the
herbarium of Brigham Young University (BRY).
Identifications were based on works by Hultcn
(1968) and Welsh (1971).

LYCOPSIDA

Lycopodiaceac Clubmoss Family

LijcojHxIiuin cilpinum L.

Locality #26, VVH 8996, 7 June. Ground

layer in woods, on glacial moraine.

Lt/(0j)0(liu7n annotimim L.

ssp. (itinotinum

Locality i26, WR 8992, 7 June. Ground
layer in woods, on glacial moraine. This species
sporulated during the second week of June and
covered the surface of Thutade Lake with spores.
Onshore breezes concentrated them along the
beaches in windrows to half an inch deep.

Lt/copodium com]>\anatiim L.

Locality #26, WR 8991, 8994, 7 June. Croimd
layer in woods, on glacial moraine.

Lijcopodium selago L.

L(x.>ality #12, RC 1.39, .lO June; #2.5, WR
9041, 9 June, in alpine tundra, on Bowser and
Siistiit Formations.

l?ion>cicAL Series, N'ol. It, No. 4 Reconnaissance of Northern British Columria

21

Selaginellaceae Selaginella Family

Selaginella sibirica (Milde.) Heiron

Locality #24, WR 9037, 9 June. Ridge crest,
on lavas, in alpine t\indra. This is evidently the
first report of S. sihirirn for British Columbia.

SPHENOPSIDA
Equisetaceae Horsetail Family

Ecjuisetttm arvense L.

Locality #26, WR 9074, 14 June. In thicket
along stream bank, on glacial moraine.

Equisettim variegatum Schleich.

Locality #23, WR 9091, 14 June. In moist
site, on gravel bar.

PTEROPSIDA FILICINEAE

Polypodiaceae Fern Family

Cryptogramnui crisjxi ( L. ) R. Br.

Locality #25, WR 9111, 14 June. Rock out-
crop, on steep south-facing, grassy slope, on
Bowser Fonnation.

Cystopteris fragilifi ( L. ) Bemh.

Locality #25, WR 9098, 14 June. Rock out-
crop, on steep south-facing, grassy slope, on
Bowser Fonnation.

PTEROPSIDA GYMNOSPERMAE

Pinaceae Pine Family

Abies lasiocarpa (Hook.) Nutt.

Locality #26, WR 8981, 7 June. A dominant
in the Boreal Forest, and the chief component of
the well-developed krummholz of the region.

juniperus connnunis L.

Locality #26, WR 8989, 7 June. Open
woods, on morainic gravels.

Picea glaiica ( Moench ) Voss

Locidity #6, RC 262, 6 Aug.; 26, WR 8985,
7 June. A dominant species in the Boreal Forest.

Pinus contorta Dougl.

var. latifolki Engelm.

Localitv #1, RC 275, 13 Aug.; #6, RC 263,
6 Aug.; #26, WR 8990, 7 June. A dominant
species in the Boreal Poorest.

PTEROPSIDA ANGIOSPERMAE
DICOTYLEDONEAE

Betulaceae Birch Family

Alnu,'> crispa (Ait.) Pursh

var. laciniata Hulten

Locality #1, RC 269, 13 Aug. Streamsides.

Alnus incana (L. ) Moench

ssp. nigosa (Dukoi) R. T. Clausen
var. occidcntalis ( Dipp. ) C]. L. Hitchc.
Locality #26, WR 8984, 7 June. Lake shores.

Betiila glandulosa Michx.

var. glandulosa

Localitv #1, RC 266, 13 Aug. 1969; Locality
# 6, RC 264. 6 Aug. 1969; #26, WR 8983, 7
June. Boreal Forest, taiga, and heathlands.

Boraginaceae Borage Family

Hackelia jessicae (McGregor) Brand

Locality #25, WR 9116, 14 June; #26, RC
148, 4 July. A component of the steep meadows
along snow slide tracks on south-facing slopes.

Mertensia panicuhita (Ait.) D.Don

ssp. paniculata

Locality #3, RC 219, 27 July; # 5, RC 170,
22 July; #18, RC 129, 30 June; #26, WR 9062,
13 June; # 30, RC 11, 17 June; #26, RC 47,
19 June. Lake shores, woods, morainic gravels,
meadows, heathlands, and alpine tundra.

Mijosotis sylvatica Hoffm.

Locality #3, RC 209, 216b, 27 July; #7,
RC 257, 5 Aug.; #8, RC 238, 240, 1 Aug.; #16,
RC 23, 31, 19 June; #25, WR 9107, 14 June;
#28, RC 2b, 16 June; #32, WR 9050, 10 June.
Alpine tundra, meadows, and talus.

Campanulaceae Bellflower Family

Campanula lasiocarpa Cham.

Locality #.3. RC 217, 27 July; #7, RC 249,
5 Aug.; #12, RC 137b, 30 June; #13, RC 100,
27 June; #;34, RC 80a, 25 June. Fell fields,
alpine tundra, and talus slopes, on micaceous
schists, sihceous conglomerate, and lavas.

Caprifoliaceae Honeysuckle Family

Linnaea borealis L.

var. longiflora Torr.

Locality #5, RC 171, 22 July; #6, 193, 26
July. Woods and thickets, on glacial moraine.

Lonicera invohicrata ( Richards. ) Banks

Locality #26, WR 9083, 14 June; do RC 71,
92, 25 June; do RC 158, 11 July. Lake shores
and woods, on glacial moraine. These collec-
tions represent a range extension northward in
British Columbia.

22

HiiiniiAM VouNC University Science Bulletin

Viburnum edule ( Michx. ) Raf.

Locality #25, WW 9092, 14 June; #26, WH
9084, 14 June; do RC 88, 25 June. Lake shores,
woods, and slopes, on glacial moraine and Bow-
ser Fonnation.

Caryophyllaceae Pink Family

Arenaria rubella ( Wahl. ) Smith

Locality #2, RC 2a5, 28 July (glabrous
phase); #27, RC 12'3, 29 June. Alpine tundra,
on weathered serpentine intrusive.

Arenaria sajanensis Willd.

Locality #29, WR 91.'>4, 1.5 June. Noted
only on Tertiary' Sustut coal beds.

Cerastium arvense L.

Locality #2, RC 228, 28 July; #25, WR
912.3a, 14 June. Alpine tundra and meadows, on
weathered serpentine-intrusive and on Rowser
Formation.

Cerastium beeringianum Cham. & .Schlecht.

Locality #18, RC 12,Sa, .30 June; #36. RC
48d, 19 June. Alpine tundra and alluvium along

"*.

'V

, A

^

i

■3

Fig. 19. A continiiatioii toward the left of Figure 20 and shows mainly Jurassic volcanic r<xks. Locality 9, tiriftith
Creek, is in the general vicinity at approximately 7()()() feet elevation. Krummholz, alpine tundra, talus .slopes,
and barrens.

HioLOGiCAL Series, \'<)i,. 1}. No. 4 Reconnaissance of Nohtiieiin British Coi.umdia

23

lakes and streams, on Triassic (?) marble and SteUaria longipea CokMe

alluvium.

Sileiie acaulis L.

var. exscapa ( All. ) DC.

Locality #32, \VR 9052, 10 June; #34, WR
8978, 7 June; do WR 9031, 8 June. Alpine tun-
dra and fell fields, on alluvium and lavas.

var. altocaulis (Hulten) C. L. Hitchc.

Locality #16, RC 21a, 28a, 19 June; #36,
RC 48c, 19 June. Alpine tundra, on Bowser
Formation and alluvium.

var. \ongipes

Locality #10, RC 124a, 29 June. River
gravels.

20 Northufst across the headwaters of CIriffith Creek in the east eentral part of the Spatsizi Quadrangle
from .59°29' N; 128°2.5' W. Ragged exposures along the skyline are in the Jurassic Eaglenest volcanic rocks.
The smooth saddle and relatively rounded rocks toward the right are folded Sustut rocks. A well-e.xposed
major fault here separates the Jurassic and Tertiary formations along the west side of the Sustut Basin. Knimm-
holz and alpine tundra on solifliiction slopes and talus.

24

Briciiam Younc; University Science Bulletin

Fic. 21. Lehi F. Hintzo in argillaceous exposure in the upper part of the Bowser Formation at approximately
57°20' N; 28°33' W, on ridges west of Buckinghorse Creek, which is in the background. Grassy slope
dominated by Festuca altaica, with Potentilla diicrsifolia in the foreground.

Chenopodiaceae Goosefoot Family

Cheno])odiii7n capitatiivi (L. ) Asch.

Locality #4, RC 187, 22 July. Glacial mo-
raine, in open woods.

Conipo.sitae Composite Family
Achillea miUcfolhtni L.

.s.sp. borctili.s (Hong.) Hreitimg

Locality #6, RC 200, 26 July; if 7, RC 251c,

5 Aug. Moraines, river gravels,
rock.s in tundra and woods.

uid volcanic

ssp. laiuilosa ( Nutt. ) Piper

Locality #5, RC 16Sa, 22 July; ::;27, RC 119,
29 June; #36, RC 50, 19 June. Moraines, river
gravels, and Bowser Fonnation in mountain
slopes.

Agoseris mirdntiaca (Ho(jk.) Greene

Locality i3, RC 222, 27 July; #6, RC 194,

Biological Series, \'ol. 14, No. 4 Reconnaissance of Northern British Columbia

25

Fig. 22. West across the western edge of the Spatsizi Plateau from 57°42' N; 128°43' W, across Goldfish Lake
V'aUev into the Eaglenest Range during one of the first major snowstorms in early August. The snow line is at
about the timberhne and snow blankets the upper-elevation tundra surfaces developed on volcanic rocks in
the background and on flat-lying siliceous sedimentary rocks in the foreground. Alpine tundra.

26 July; #26, RC 152, 11 July. Ridge tops and
mountain .slopes, in micaceous schists and mo-
raines, in alpine tundra and woods.

Anteniiaria alpina (L. ) Gaertn.

var. stolonifcra (Porsild) Welsh

Locality #22, WR 91.3;3a, 15 June; #36, RC
55a, 19 June. Alpine tundra, heath, and woods,
on slate and alluvium.

Antennaria monocephala DC.

Locality #20, RC 20, 18 June; #21, WR
9089, 14 June; #29, WR 9133, 15 June; #34,
WR 90.3.S, 9 June; #.37, WR 9009, 8 June. Alpine
tundra and heathlands, on Bowser Formation,
Tertiary Sustut coal beds and lavas.

Antennaria neglecta Greene

Loc-ality =26, RC 89, 25 June. Open woods
on glacial moraine.

Antennarui rosea Greene

Locality #25, WR 9112;i, 15 June. Open
wood.s and meadows, on alluvium.

Arnica cordifolia Hook.

Locality #2,5, WR 9110, 14 June; #26, 87,

25 June. Open woods, on alluvium and glacial
moraine.

Arnica latifolia Bong.

Locality #3, RC 215, 27 July; #7, RC 254,
5 Aug.; #26, WR 9076, 14 June; do RC 36, 19
June;" do RC 95, 25 June; do RC 165, 11 July;
#39, RC 1.35, 30 June. Alpine tundra, meadows,
and open woods, on micaceous schist, glacial
moraine, and cherty rubble.

Artemisia campestris (L. ) DC.

ssp. borealis (Pallas) H. & C.

var. borealis

Locality #5, RC 180, 22 July. Glacial mo-
raine, in woods.

Artemisia nowegica Fries

var. saxatilis (Besser) Jeps.

Locality #3, RC 22.3, 27 July; #7, RC 258a,
5 Aug.; #21, WR 909, 14 June;' #26, WR 9078,
14 June; #29, WR 91.32a, 15 June. Open woods,
alpine tundra, and fell field, on micaceous
schists, coal beds, and glacial moraine.

Erigeron JiumiUs Grah.

Locality #8, RC 2.39a, 1 Aug.; #17, RC 21,
19 June; #27, RC 121, 29 June; #29, WR 9128,

26

Bricham Young University Science Bulletin

i«

%:.^ A \ \^^

:»^

Fig. 23. Lehi F. Hintze photographing Monkshood and sedge along the north fhink of tlie northwest spur of
Mt. Will, near Locality 8, on siliceous Jurassic volcanic rocks of the Eaglenest volcanic sefiiience at apjiroxi-
mately 57°.32' N; 128°47' \V, and at an elevation of 6000 feet. Alpine tundra, witli Hicrochloi- alpina, Aconi-
tum delphiiiifoliuni, and Carex.

15 June; #36, RC 48b, 19 June. Alpine tundra

and talus slopes, on Bowser Fonnation, and

alluvium.

Erigeron lonchoplii/llu.s Hook.

Loc-ality #4, KC l.S9h, 22 July. Glacial mo-
raine, in woods.

Eri^cron pcregrinus (Pursh) Greene
ssp. rallimithemus (Greene) Gronq.
Locality #.3, KG 212, 27 July; #7, RC 247.

5 Aug.; #25, WR 9101, 14 June; #26, RC 74,

74a, 93, 25 June. Open woods, meadows, and

alpine tundra, on micaceous schist.

Biological Series, Vol. 14, No. 4 Heconnai,ss.\nce ok NomiiEHN British Columbia

27

Hieracittm gracUe Hook.

Locality #3, RC 220b, 27 July. Alpine tun-
dra, on micaceous schist.

Petasites frigidtts L.

var. frigidtts

Locality #14, RC 60. 25 June; #22, WR
9056, 12 June; #27, RC 144b, 30 June. Alpine
tundra, fell fields, and meadows, on alluvium
and Bowser Formation.

var. palimitus (Ait.) Croncj.
Locality #38, WR 9025, 8 June. Wet mea-
dow, along stream.

Senecio lugens Richards.

Locality #17, RC 97, 27 June; #18, RC 130,
■30 June. Alpine tundra and woods, on Triassic
( ? ) marble and alluvium.

Senecio triangularis Hook.

Locality #3, RC 210, 225, 27 July; #7, RC
258, 5 Aug.; #25, WR 9122, 14 June; #26, RC
166, 11 July. Meadows, open woods, and alpine
tundra, on micaceous schist. Bowser Formation,
and morainic gravels.

Solidago midtiradiata Ait.

Locality #5, RC 169a, 22 July; #6, RC 201,

' '-XjftM^^'- '

f^' '.\^i^-^''*7%

',-^'->'^"V'

r- ." ^'^

-*i0^^

\i^^^^7

•■JS-2

^•::-...^v.>■:.

:-'is-Jr

jV'S.-?*'

• ■ '■-■ ^' ' A^-'lr.

^T''^.

■ r^

■ •.: ii^N'*-

tic. 24. \ lew westward toward ,i tdiigiiimcratf and ci)al-l)c,iriMg sfijiiciicf Jii the lower part ot the Bowser
Group on the northeast side of Stalk Ridge. Conglomerate and sandstone form the prominent ledges with eoal
and earbonaeeous shale forming the distinctive slope zone. Hlant.s in the foreground are growing on siliceous
pebbly conglomerate and are typical of alpine tundra along ridge crests. The foreground is at an elevation
of 6560 feet and at .57°09' N; i27°35' W, a short distance south of collecting locality 12. Plants from Locality
12 were collected on the same beds of sandstone and conglomerate shown abo\e the talus.

28

Bricham Young University Science Bulletin

Kic. 25. Horizontally bedded Sustut rocks on the northwestern part of the Mount Jorgensen massif at the north-
west end of Tafjletop Mountain at 56°57' N; 127°09' VV, at 6500 feet in the Spatsizi Pleateau. Peaks along
the skyline in the distance are part of the folded Bowser sefjuence west of the Sustut Basin. The alternating
terraces are prothiced by interbedded resistant conglomerate and sandstone. Easily eroded shales form vege-
tated stripped surfaces on top of the sandstone ledges. Alpine tundra heathlands and hmited kmmmholz vege-
tation.

201a, 26 July; #7, RC 253, 5 Aug. Open woods,
heathlands, and alpine tundra, on moraine and
volcanic rocks.

Taraxacum ceratophorum (Ledeb. ) DC.

Locality #8, RC 239, 1 Aug.; #13, RC 99b,
27 June; #36, RC 50a, 19 June. Talus slopes
and ridges, in woods, meadows, and alpine tun-
dra, on Bowser Formation and alluvium.

Taraxacum eriophorum Rvdb.

Locality #17, RC 22, 19 June; #20, RC
19b, 18 June. Alpine tundra, fell fields, and
open woods, on allu\ium, and Bowser Fonna-
tion.

Cornaceae Dogwood Family

Conius canaclemis L.

Locality #26, WR 9075, 14 June; do RC 9,
17 June. Woods, on niorainic gravels.

Crassulaceae Stonecrop Family
Seiluni lanceolatum Torr.

Locality #6, RC 191, 26 July; #30, RC 14,
17 Jime. Clacial moraine in open woods, and
alpine tundra on SiLstut Formation.

Cruciferae .Mustard Family
Arahi.s divaricarpa A. Nels.

Locality- #4, RC 186, 26 July. Glacial mo-
raine in open woods.

Arahls drummomlii Gray

Locality #19, RC 114, 28 June. On granite
liedrock. in open woods.

Arabis glabra ( L. ) Bernh.

Locality #25, WR 9103, 14 June; #26, RC
37d, 19 June; ^28, RC 4, 16 June. Alluvium
and moraiiiic gravels in open woods and
meadows.

Biological Skhies, \'ol. 14, No. 4 Mkconnaissanc:!; of Nohihkkn British Columbia

29

Arabis lemmonii Wats.

Locality #;33, WR 9050a, 10 June. Alpine
tundra, on volcanic rocks.

Am his Itjrata L.

Locality #8, RC 237b, 1 Aug.; #14, RC 56,
2.5 June; #28, RC 3, 16 June; #29, WR 9142ii,
15 June. Open woods, heathlands, and alpine
tundra, on alluvium, talus, and coal beds.

Barbarea orthoceras Ledeb.

Locality #20, RC 37f, 19 June; do, RC 79,
25 June. NIorainic gravels in open woods.

Cardamine pratensis L.

Locality #14, RC 60b, 25 June. Alluvium,
in meadows.

Drabii aurca Vahl

Locality #5, RC 178, 22 July; #6, RC 197,

26 July; #40, RC 101, 28 June. Glacial mo-
raines, in open woods and heathlands.

Draba borealis DC.

var. maxima ( Hulten ) Welsh

Locality #7, RC 251, 5 Aug.; #25, WR
9112b, 14 June; #27, RC 144, 30 June.
Meadows, open woods, and alpine tundra; on
volcanic rocks. Bowser Fonnation, and alluvium.

Draba incerta Pay son

Locality #13, RC 100b, 27 June; #16, RC
28b, 19 June; #27, RC 144a, 30 June; #29,
WR 9130, 15 June; #34, WR 9036, 9 June. Al-
pine tundra and fell fields, on Bowser Forma-
tion, coal beds, and volcanic rocks.

Draba nivalis Payson
var. elongata Wats.

;>.y*.-y<v.-^;.^

.*')«'

^^%*>.*^

iH*if ;^?p^:^^«^*S>:JI^'

>^i^''£jv^fC#'!^

-^

Fk;. 26. West from the northea.st spur of Siistut iVak at .56°35' N; 120'=36' W, acro.ss the Sustut f5iver Valley at
■3.5(M) feet to the soiithuest spur of Savage Peak wliieli rises to over 7000 feet. Lower dark green agglomerate
of the Takla Group forms the prominent high .serrated peaks in tlie right center and right of the photograph.
The upper part of tlie Takla group forms tlie somewhat rounded cliffs along the skyline to the left. Prominent
steep valleys are car\ed in softer more distinctly bedded tuffaceous and graywacke sandstone beds inter-
bedded with the agglomerate. Dryus intcgrijolia and crustose lichen fell field in the foreground.

30

Brigham Young University Science Bulletin

*■■•(

'J*-H?ftr-:_

-^sar^*- /-i-Sr^'S

Ik.. 27. CdUrLliiiii lot.ilitv 2, sIkhmi In tlu' anuu, i.s at an ^^.^lK'^Ul^ pmspiLl jicii Ijt-laiii L.ikc, the pioniinent
lake in the left intermediate di.stance. Country rocks here are metamoqiho.sed argillite, argillaceou.s quart-
zite, and volcanic rock.s which have been intnided by serpentine peridotite. Serpentine peridotite form.s the
light-colored exposures in the background along the ridge crest immediately be\ond the collecting locality.
Low-rank mica sciiist foniis the low rounded hills in part of the area. The Localitv '1 collecting site is at 58°20'
N; I28°45' W, at approximately .5400 feet in elevation. Alpine tundra, heathlands, knimmholz and meadows.

Locality #37, \VR 9009a, 8 June. Alpine
tundra, on lava.s.

Draba stcnoloha Trautv.

Locality #28, RC .3a, 16 June. Alluvium, in
heathlands.

Thlaspi arvense L.

Locahty #4, HC 18.5, 22 July. Glacial mo-

raine, in open wood.s.

Elaeagnaceae Oleaster Family
Slieplierdia canadensis (L. ) Nutt.

LcK-ality #1, RC 270, 13 Aug.; #26, WR
9065, 1.3 June. Lake shores and stream banks,
on glacial moraine and stream terrace gravels,
in woods.

Biological Series, Vol. 14, No. 4 RECoNNAi.ssANt:E ok Nohthehn Bhitinh CoLUMniA

31

Empetraceae Crowberry Family

Empetrum nigrum L.

Locality #1, RC 278, 13 Aug.; #26, WR
8982, 7 June. Open woods and heathlands, on
alluvium and moraines.

Ericaceae Heath Family

Arctostaphylos uva-ursi ( L. ) Sprang.

Locality #1, RC 279, 13 Aug.; #26, WR
8997, 7 June; do RC 37, 19 June; do RC 72, 25
June; #30, RC 7, 17 June. Alluvium glacial

Fig. 28. \'ku lonkiiii; nortliwivst fnim Triangulation Station 6.5.39 about four miles ea.st of the junction of the
right and left forks of Kelileehoa Hiver at 59°09' N; 128°37' W. Rocks in the foreground are metamorphosed
phvllite and slate Unit 14 on the Cry Lake quadrangle geologic map of (Jeologic Survey of Canada. Rocks
of Unit 11-B and 12 of the Crv Lake quadrangle can be seen across Kehlechoa River in the background, on
the right of the photograph along the crest of the ridge in the middle distance between the two forks of
Kehlechoa River. Alpine tundra ilominated by species of Carex, Dryas, and Lichen.

32

Bkigham Young Univehshv Science Bulletin

-•<>-;.

^■'.^

'^■'^. >

Fig. 29. View towards the northwest along the northeast face of Stalk Ridge. The Stalk Ridge collection at
Locality 12, 57°09' N; 127°37' W, at appro.vimately 6000 feet, was made in the ledges at the extreme riglit
margin of the photograph. All the rocks seen here are in the Bowser formation. Plants from Localit)' 12 were
collected principally off the siliceous conglomerate and sandstone beds which form the prominent ledges in the
intermediate distance. Alpine tuiulra and fell fields.

moraine and Sustut Fonnation in open woods
and alpine tundra.

Arctostaphtjlos alpina (L. ) Spreng.

var. al])irui

Locality #37, WR 9011, S June 1969. Alpine
tundra, on lavas.

Cassiope niertensiaiui ( Bong. ) D. Don

Locality #.3, RC 215b, 27 July; #15, RC IS,
18 June; #18, RC 134, 30 June.' Alpine tundra
and heathlands, on micaceous schist, alluviiun,
and marble.

Cassiope tetragona (L. ) D. Don

var. saximontana (Small) C. L. Hitchc.
Locality #29, WR 9141, 15 June; #.34. RC

SO. 25 June; #35, WR 9042, 9 June. Alpine
tiHidra and heathlands, on coal beds and lavas.

Ledum <groen\andicum Oeder

Locality #1, RC 277, 13 Aug.; #5, RC 177,
22 July. Glacial moraine and river terrace
gravels, in woods.

PhijUodoce empetrifoniiis (Sm.) D. Don

Localitv #18. RC 133, .30 June; #29 WR
913S, 15 June; #30, RC 10, 17 June. Alpine
tundra and heathlands, on marble, coal beds,
and s;mdstone.

Philllodocc <il(indnHflara ( Hook. ) Coville
Locality #2, RC 231d, 28 July; if? IS,

RC

Biological Series, Vol. 14, No. 4 Reconnaissance of Northern British Columbia

33

132, 30 June; #26, RC 91, 2.5 June; #29, WR
9139, 15 June; #37, WR 9010, 8 June. Alpine tun-
dra, heathlands, and open woods, on weathered
.serpentine intrusive, marble, glacial moraines,
coal beds, and la\a.

Viicciiiitini c(icsj)itosum Michx.

Localitv #25, WR 9099, 14 June; #26, WR
9079, 14 June; do RC 72b, 25 June. Open woods
and meadows on morainic gravels and alluvium.

Vaccinium rnembranaceum Dougl.

Locality #26, WR 9063, 13 June. Open

woods on morainic gravels.

Vaccinium vitis-idaea L.

Locality #1, RC 276, 13 Aug.; #12, RC 141,
30 June. Open woods and heathlands, on allu-
vium and siliceous conglomerate.

Fumariaceae Fumitory Family

Conjdalis seinpervirens ( L. ) Pers.

Locality #1, RC 280, 13 Aug. River terrace
gravels, in open woods.

Fig. .30. .Soiitlieast toward the soutliwi-^t .\ll. Will glacier from .57°32' N; 128°.50' \\ . Uocks along the .skyhne
toward the left on the south spur of Mt. Will are held up by Eaglenest volcanics which are here essentially
flat King. Rocks in the center and the center right which form the high ledges are the topmost beds of the
Eaglenest \olcanic sequence and are overlain by shaly exposures of the Bowser beds at the upper right. Mt.
Will rises to appro.ximately 8()00 feet above valleys at' .5000 to 6000 feet. Alpine barrens and fell fields.

34

Briciiam Young University Science Bulletin

Fig. 31. Omineca and Sustut highlands looking southeast from Sustut Peak towards Svistnt Lake from 56°36' N;
126°32' W. Outcrops immediately hevond the helicopter are of the Asitka and Cache Creek Groups and are
dipping steeply toward the southwest. Asitka Peak, elevation 7055 feet, is the isolateil mountain beyond
Sustut Lake, elevation 4250 feet, and is composed in large part of southwest dipping Takla volcanics. High
peaks along the skyline in the distance are in the Ingenika and Osilinka ranges and are composed in large
part of Takla volcanics intnided by Omineca granodiorite and (juartz diorite. Sparse alpine tundra, with
Lijcopodhim selago, Cassioiw tetragona, Dnjas iiiiegrifotui and S«/i.v rcticuliila.

Gentianaceae Gentian Family
Gentianii filouca Pallas

Locality #3, RC 215a, 27 July; #7, RC
252b, 5 Aug.; #12, RC 136, 30 June; #18, RC
128, 30 June; #27, RC 121a, 29 June. Alpine
tundra and heathiands, on micaceous schist, vol-
canic roc-ks, siliceous conglomerate, and Bowser
Formation.

GentiuneUa ainareUa (L.) Bonier

Lociility #5, RC 173, 27 July; #6, RC 196,
205, 26 July; do, RC 244, 3 Aug. Glacial mo-
raine, in open woods.

Centianella propinqua (Richards.) Gillette

Locality #2, RC 232, 28 July; #4, RC 189a,
22 July; #5, RC 172;i, 22 July. Open woods,
meadows, and alpine tinidra, on weatiiered ser-
pentine intrusive and glacial moraine.

Geraniaceae Geranium Family
Geranium erianthuni DC.

Locality #14, RC 57, 26 June; #25, WR

9119, 14 June; #27, RC 122, 29 June. Open
woods and meadows, on alluvium, glacial mo-
raine, and Bowser Fonnation.

Leguminosae Legume Family

Astragalus alpinus L.

Loc-alitv #26, RC 37c, 19 June; #28, RC 28,
19 June; ;2S, RC 1, 16 June; =r.36, RC 65, 19
June. Open woods, lake shores, and stream
gravels.

Hedijsarum aljiinum L.

Locality #5, RC 175. 22 July; #6, RC 192.
26 Julv. Open woods, on glacial moraine.

Lupinus arcticus Wats.

Locality #5, RC 168, 22 July; #6. RC 204,
26 Julv; #33, WR 904S. 10 June; #.34, WR
8980. 7 June; do WR VKK)1. 9(W2, S June; do WR
9033, 9 June; #36. RC 45, 19 Jime; #37, W'H
9068, 13 June. Open woods, heathiands, and al
pine tundra on moraine and lavas. All speci-
mens collected in alpine sites are considerably

Biological Series, Vol. 14, No. 4 Reconnaissance of Northern British Columbia

35

throughout Alaska and Yukon, but the alpine
pha.ses resemble var. jordalii (Porsild) Welsh in
leaflet size and number. The flower size is
within the range of variation of var. varians and
apparenty the specimens are merely alpine
dwarfs of var. various.

Onagraceae Evening-Primrose Family

Epilobium aJpinum L.

var. ali)imtin

Locality #7, RC 248a, 5 Aug.; #9, RC 237,
1 Aug.; #26, RC M, 19 June. Alpine tundra
and heathlands and open woods, on volcanic
rocks, Bowser Formation, and lavas.

Epilobium angustifoJium L.

Locality #5, RC 182, 22 July; #6, RC 202,
26 July; #7, RC 255, 5 Aug.; #19, RC 112, 28
June. Open woods, heathlands and tundra, on
gravels, moraines, talus, and granite bedrock.

Epilobium latifolium L.

Locality #7, RC 246. 5 Aug.; #8, RC 236,
1 Aug.; #16, RC 96, 27 June. Alpine tundra
stream gravels, and talus, on volcanic rocks and
Bowser Formation.

Fig. 32. Mr. Ciraham Cuddy, who assisted with some
of the plant collection, on barren e.xposures of silt-
stone and coal in the upper part of the Sustut
Formation at the south end of the Sustut Basin and
the Spatsizi Plateau at Locality 29, at appro.ximatelv
56°45' N; 127°05' \V, at approximately 6.500 feet
in elevation. Semibarren alpine tundra, dominated
by Draha incerta and Carcx podocarpa.

more silkv villous than those from woodland
sites at lower elevations.

Lupimis nootkatensis Donn

var. nootkatensis

Localitv #25, WR 9120, 14 June; #26, WR
8993, 7 June; do 9061, 13 June. Grassy slopes
and open woods on glacial moraine. L. nootka-
tensis is common in the Pacific drainages, but
has seldom been reported in the Mackenzie
drainage. Thus, the report of this entity for the
Thutade Lake vicinitv is unique.

Oxtjtro])is eampestris ( L. ) DC.

var. varians ( Rvdb. ) Barn.

Localitv ^6, RC 204b, 26 Julv; #16, RC 25,
43, 19 June; #29, WR 9131, 15 June; #34, RC
81. 25 June; =.37. WR 9004, 8 June. Alpine
tiuidra and heathlands on Bowser Formation,
coal beds, and lavas, and on glacial moraines
and river gravels in open woods. The low ele-
vation phases are similar to specimens from

Papaveraceae Poppy Family

Papaver alboroseum Hulten

Locality #16, RC 44, 19 June; do RC 98,

27 June. Alpine tundra on Bowser Formation.
This is apparently the first report of P. alboro-
seum for British Columbia.

Papaver radicatum Rottb.

Locality #16, RC .30, 19 June; do RC 116,

28 June. Alpine timdra, on Bowser Fonnation.

Polemoniaceae Phlox Family

Polemonium caeruleum L.

ssp. villosuni ( Rud. ) Brand

Locality #2, RC 231a, 28 July; #3, RC 213
(white flowers), 214, 27 July; #5, RC 184, 22
Julv; #6, RC 207, 26 July; #7, RC 257, 5 Aug.;
#14, RC 58, 25 June; #19, RC 111, 28 June;
#31, RC 110, 28 June; #36, RC 49b, 19 June.
Alluvium and moraines along streams and lake
shores in open woods, and in heathlands and
tundra on weathered serpentine intrusive, mica-
ceous schist, and granite.

PolemoniiDn jiideherrimum Hook.

Locality #16, RC 27, 42, 19 June; # 25,
WR 9097, 14 June; #28 RC 2a, 16 June; #31,
RC 108, 28 June; #.3.3, WR 9049. 10 June. Al-
pine tundra, heathlands, meadows, and open

36

Bhigham Young University Science Bulletin

o
c

5

^

1

:3

S'

Fig. 33. Map of colK-Ltiiig localities in northern British Columbi.i.

Biological Series, \'ol. 14, No. 4 Reconnakssance of Northern British Columbia

37

woocLs on glacial moraine, talus, alluvium, Bow- 27 July; #5, RC 177a, 22 July; #", RC 248,

ser Foniiation, and la\as. 5 Aug.; #36, RC 48, 19 June. Open woods,

heathland, and alpine tundra, on alluvium, gla-

Polygonaceae Buckwheat Family cial moraine, weathered serpentine intrusive,

Oxyria digijnia (L.) Hill ^'"^^ micaceous schist.

Locality- =r7, RC 259, 5 Aug.; #22, 9055, 12 r„„^^ ^^^,o,^ L.

June. Alpine tundra, on volcanic rocks and ^ocalitv #3, RC 224, 27 July; #25, WR

^ ■ 9102, 14 June; #28, RC 5. 16 June; #38, WR

Pohj<iomnn vivijximm L. 9016, 8 June. Open woods and meadows, on

Localit\- #2, RC 227, 28 Julv; #3, RC 216a, alluvium and micaceous schist.

Fig. .34. .Siisliit IVak, thi- sharp i;latiate{l horn near the upper center of the photograpli, is one of the striking topo-
graphic features of tlie .Sustut region near tlie southern border of the area investigated. It is held up by mas-
sive volcanic rocks of Triassic age and is flanked bv lower Triassic rocks and Paleozoic rocks in the long ridges
at the northeastern base of the range and bv Jurassic volcanic rocks at the southwestern base. The valleys
are clothed by Boreal Forest which is continuous with knimmholz upwards.

38

Brigiiam Young University Science Bulletin

Fig. 35. Nivt-n Peak, tlie shaipK outliucd pu.ik along tlic ililgi- jii tlir left lliiIii ul tlic pliotograph, is typical of
the topographic expression of Triassic and L'pper Paleozoic rocks of the Sustut Peak region. Locality 35. near
the upper margin, is on siliceous volcanic rocks of the Asitka Group.

Portulacaceae Purslane Family

Monlia sarnwntosa (C. A. Moy. ) Robin.s.

Locality' #2, RC 231e, 28 July; #3, RC 216,
27 July; #8, RC 237a, 1 Aug.; #11, RC 150, 8
July; #41, RC 10.3, 28 Juno. Alpine tundra, on
weathered .serpentine intrusive, niieaceous schist.
Bowser Formation, and on lavas.

Primulaceae Primro.se Family

Thentahs europaea L.

Locality #26, WR 9073, 14 June; do RC:
.37b, 19 June; do RC: 82, 2.5 June; #28, RC 8,
17 June. Claeial moraine and aliuvivmi, in open
\vo<k1s.

Pyrolaceae Wintergreen Family

Pi/rola a.sarifoUa Michx.

Locality #5, RC 172, 22 July; #6, RC 203,
26 July; #26, RC 1.54, 11 July; do RC 84, 25
June. Open woods and meadows, on alluvium
and glacial moraine.

Ranunculaceae Buttercup Family
Aconitum delpltinijolmm D('.

var. delphmijolhim

Locality #.3, RC 221, 27 July; #5, RC 179,
22 July; #6, RC 208, 26 Julv; #7, RC 256, 5
Aug.; RC 127, 30 June; #26, 159, 11 July; #27,
RC] 118, 29 June. Meadows, open woods, lake

Biological Series, Vol. 14, No. 4 Reconnaiss.^nce of Northern Bbiti.sh Columbia

39

shores, river banks, and alpine tundra, on allu-
vium, glacial moraines, talus slopes, micaceous
schist, volcanic rocks, and Bowser Formation.

Anemone multifkhi Poir.

Localitv #5, RC 195, 26 July; #15, RC 18a,
18 June; #33, WR 9051a, 10 June. Alpine tun-
dra, heathlands, and open woods, on volcanic
rocks, alluvium, talus, and glacial moraine.

Ancnwnc narcussiflora L.

var. monantha Schlecht.

Locality #2, RC 233, 28 July; #3, RC 220a,
27 July; #41, RC 104, 28 June' Alpine tundra,

on serpentine intrusives, micaceous schist, and
volcanic rocks.

Anemone parviflora Michx.

Localitv #5, RC 181, 22 July; #16, RC 31a,
19 June; #.34, WR 9003, 9032, 8 June; #37,
WR 9005, 8 June. Alpine tundra and open
woods, on glacial moraine. Bowser Formation,
and lavas.

Anemone richardsonii Hook.

Locality #26, WR 9071, 14 June; #28, RC
la, 16 June. Dense woods, on alluvium.

Fig. .36. Collecting Locality 25 is on the .southern flank of a high ridge of Bowser muclstone and sandstone on
the north side of Thvitade Creek, west of Thutade Lake. It is one of the most prolific locahties collected during
the summer.

40

Brigham Young Universitv Science Bulletin

Fig. 37. West end of Tlmtade Lake is bordered b\ mountains car\eil in mudstone and conglomerate of the Bowser
Group. Locality 26 is at the campsite, on the liikeshore, and Locaiitv 27 is on the higli ridge jjehind camp
to the north.

AqtiHegia fonnosa Fi.sch. ex DC.

Locality *25, WR 9104, 14 June; #26, 146,
4 July. Cnussy .slopes, on talus.

Caltha leptosejyala DC.

Locality #10, RC 124, 29 June; #19, RC
115, 28 June; #26, RC 147, 4 July; #26, RC
95a, 2.5 June; #38, VVR 9020, 8 June. Stream
banks and wet meadows, on alluvium and
granite.

Delplunium <:^latictim Wats.

Locality #5, RC 18.3, 22 July; #6, RC 199,
26 July; #26, RC 160, 11 July. Open woods

and meadows, on glacial moraine and alluvium.

Ramtnntlus eschsclwltzii Schlecht.

Locality ii:8, RC 241, 1 Aug.; #10, RC 125a,
29 June; '#16, RC 44b, 19 June; #25, WR
912:3, 14 June; #26, WR 9000, 7 June; RC 6,
16 June; #29, WR 9126, 15 June; #33, WR
9047, 10 June. Open woods, meadows, alpine
tundra, and lieathlands, on talus, alluvium. Bow-
ser Foniiation, coal beds, and volcanic rocks.

Ranunculus nivalis L.

Locality # 12, RC 138, 30 June. Alpine tun-
dra, on siliceous conglomerate.

Biological Series, Vol, 14, No. 4 Reconnaissance of Northern British Columbia

41

Ranunculus orcidcntalis Nutt.

\;ir. brcvistiflus Greene

Localit)' #8, RC 241a, 1 Aug.; #14, RC 61,
25 June. Open woods, nieadow.s, and alpine
tundra, on Bowser Fonnation, and alluvium.

Ranunculus pijgmaeus Wahl.

Locality #12, RC 137, .'30 June. Alpine tun-
dra, on .siliceous conglomerate.

Ranunculus unicinatus D. Don

Locality #26, RC 163, 11 July. Open woods.

on glacial moraine.

Thalictruin aJpinum L.

Locality #36, RC 51, 19 June. Marshy lake
shore, in woodland.

Thalictruin occidentale Gray

Locality #25, WR 9121, 14 June; #26, WR
9067, 13 June; do RC 39, 19 June; do RC 78, 25
June; #36, RC 49, 19 June. Open woods, grassy
slopes, meadows, and lake shores, on glacial
moraine and alluvium.

I'll.. yH. Ml. Jcirgfiiscn is held iij) by iiuariy liciriziiiital altfriialiiii; beds (it cdiigloim'ralK.- sandstdiic and argillaceous
mudstone. Sandstone beds form ledges and mudstone form slopes in the step-like erosional pattern well ex-
pressed in eirque floors. Locality 30 is on Mt. Jorgenscii.

42

Bhigham Young University Science Bulletin

:^.?-i>W-'.^

y V

y^

Fig. 39 Kittlumi Laki is in the region between tlic lolUnl immiiUuus Id tin- wist and the Spatsizi Phiteau to
the east. Locahty 17 is at the eastern end of the lake near its ontlet. Locahty 16 near Kitchener Crag is
just off the photograph along the sharp ridge at the lower left, and is in Bowser shale and sandstone.

Hosaceae Ro.se Family

Dn/a.s intcfirifolid Valil

var. integrifoUa

Locality #16, RC 41, 19 June; #35, WR
90.39, 9 June; #.37, WR 9()7(), 13 June. Alpine
tundra, on Bowser Fonnation, Su.stut, a domi-
nant .specie.s.

Dnjas octopetala L.

var. kamtschatica (Juz. ) Hulten

Locality #2, RC 231c, 28 July; #13, RC
99, 27 June. Alpine tundra, on .serpentine intru-
sive and Bowser Formation. These specimens

approach D. intc^rifolUi but possess glands simi-
lar to D. octopctcilii.

Fragaria virginiami Duchesne

var. glauca Rydh.

Locality #31,' RC 106, 28 June. Open woods
and meadows, on alluvium.

C.einu nwcroiiliiiUum VVilld.

Locality #26, RC 1,57, II July. Open woods
and meadows, on glacial moraine.

Ltirtkea jwctitiata (Pursh) Kuntze

Locality #2, RC 229, 28 July. Alpine tun-
dra, on serpentine intrusive.

Biological Series, Vol. 14, No. 4 Reconnaissance of Northern British Columbia

43

Potentilhi (UvcrsifoJui Lehm.

LocaliU' #16, RC 32, 44a, 19 June; #25,
WR 9096, '14 June; #26, RC 37e, 19 June; #28,
RC 2, 16 June; #29, WR 9129, 15 June; #31,
RC 107, 28 June; #36, RC 49a. 19 June; #38,
WR 9024, 8 June. Alluvium, Bowser Formation,
and coal beds, in meadows, lake shores, alpine
tundra, and woods.

PotcntiUa fndicosa L.

Loc;ility #5, RC 176, 22 July; #10, RC 125,
29 June; #26, RC 167, 11 July; #36, RC 54,
19 June. Lake shores, open woods, meadows,
and heathlands, on glacial moraine and alluvium.

Potentilla htjparctica Malte

Locality #21, WR 9088a, 14 June; #22, WR
9132, 15 June; #34, WR 9035, 9 June. Alpine
tundra, on slate and lavas.

Potentilla unifloru Ledeb.

Locality #16, RC 29, 19 June; #21, 9088,
14 June; #.33, WR 9051, 10 June; #34, WR
9002, 9030, 8 June; #35, WR 9039a, 9 June.
A dominant in alpine tundra, on Bowser Forma-
tion and lavas.

Rosa acicularis Lindl.

Locality #5, RC 174, 22 July; #6, RC 206,
26 July. Glacial moraine, in open woods.

Fig. 40. Stalk Lakes region collecting localities arc on Jurassic Bowser siliceous conglomerate and sandstone
(locality 12) and on alluvium and lake shore material associated with a landsile (locality 14), at the south-
western margin of the Spatsizi Plateau region.

44

Blllcll AM ^Olno Um\ KHSITV SCIENCE BULLETIN

■.../'•■

\

Fig. 41. Colclfish Lake region between tlie Eaglenest Range, on the lett, and Spatsizi Plateau, on the right. Lo-
cality 6 is at the campsite and is on glacial material, modified by lake margin deposits. The Eaglenest Range
is held up by Jurassic volcanic rocks while the Spatsizi Plateau is held up largely by Tertiary Sustut Forma-
tion.

Ritlms arcticus L.

Locality #26, RC 38, 19 June. Open woods,
on glacial moraine.

Rubus idaeus L.

.ssp. melanoldsius (Dieck.) Focke

Locality #19, RC 1.33, 28 June. Open woods,

on granite bedrock.

Rtihus pedatus J. E. Smith

Locality #26, RC 37a, 19 June; do RC 145,
4 July. Open woods, on glacial moraine.

Rubus stellatus J. E. Smith

Locality #28, RC Ic, 16 June. Ileathland,
on alluvium.

Sangui.sorbd officinalis L.

Locality #2, RC 234, 28 July. Alpine tundra,
on serpentine intrusive. This report represents
a locality intermediate between populations in
the Yukon and those to the south in British
Columbia.

Sanutii.iorbd stipuhita Raf.

Locality #4, RC 189, 22 July; #6, RC 204a,
26 July; #26, RC 161, 11 July. Alluvium and
glacial moraine in open wmxls.

Sibluddia procumbens L.

Locality #29, \VR 9135, 15 June; #36, RC
55, 19 June. Lake shores in open woods and
alpine tundra on coal beds.

Biological Series, Vol. 14, No. 4 Rec;onnaiss.\nce of Northehn Briti.sh Columhia

45

Sorhus scopuliim Greene

Locality it26, VVR 9080, 14 June; do RC
33, 19 June; do RC 72a, 77, 90, 25 June. Open
woods and grassy slopes, on alluvium, glacial
moraine, and talus.

Rubiaceae Madder Family

Galium boreale L.

Locality #5, RC 169, 22 July; #6, RC 190,
26 July, RC 120, 29 June; do RC 156, 11 July;
±t40, RC 102, 25 June. Open woods, meadows,
and grassy slopes, on alluvium, moraine, and
talus.

Salicaceae Willow Family

Popuhis halsamifera L.

Locality #1, RC 273, 13 Aug. River terrace
gravels, in riparian woods.

Popuhis tremuloidcs Michx.

Locality #1, RC 267, 13 Aug.; #6, RC 260,
6 Aug.: #24, WR 9143, 15 June." River terraces,
slopes, and open coniferous woodlands.

Salix (irctica Pallas

Locality #3, RC 218, 27 July; #30, RC 15,
17 June; #37, VVR 9012, 901.3, 8 June; do, WR
9060, 13 June. Alpine tundra, on siliceous schist,
Sustut Fonnation, and lavas.

Sali.x alaxensis Cov.

var. longistijli.'i ( Rydb. ) Schneid.

Locality #1, RC 274, 274a, 13 Aug. River
terrace gravels, in woods and thickets.

Sa/i.v barclayi Anderss.

Locality #26, WR 8986, 8988, 8998, 7 June;
do WR 9082, 14 June; #32, WR 9045, 9046, 10
June; #64, RC 64, 19 June; #.38, WR 9021,
9022, 9023, 8 June. Stream banks, lake shores,
thickets, and meadows, on alluvium and mo-
raine.

Salix harrattiana Hook.

Locality #.32, WA 904.3, 9044, 10 June; #.38,
WR 9026, 8 June. Stream banks and meadows,
on alluvium and moraine.

^alix hraclnjcdrjxi Nutt.

ssp. brachi/carjHi

Locality #.38, WR 9028, 8 June. Stream
banks and meadows, on alluvium.

Salix commutata Bebb.

Locality ::r36, RC 62, 19 June. Lake shore,
in open woods.

Salix drtiinmondiana Barr.

Locality #26, WR 8999, 7 June; do, WR

9086, 14 June; #.36, RC 63, 19 June; #.38, WR
9027, 8 June. Stream banks, lake shores, and
meadows, on alluvium.

S«/;.v glauca L.

Locality #6, RC 26.5, 6 Aug.; #25, WR
9144, 15 June; #26, WR 9064, 13 June; do WR
9081, 9085, 14 June; #29. WR 9140, 15 June.
Lake shores, stream banks, open woods, heath-
lands, and tundra.

Salix vionticola Bebb.

Locality #1, RC 272, 13 Aug. Stream ter-
race gravels.

Salix novae-angliae Anderss.

Locality #38, WR 9017, 8 June. Stream
sides and meadows, on alluvium.

Salix reticulata L.

Locality #16, RC 24, 19 June; #.35, WR
9040, 9 June; #.37, WR 9007, 8 June. Alpine
tundra, on Bowser and Sustut Formations, and
on lavas.

Salix stolonifera Trautv.

Locality #37, WR 9006, 9006a. 8 June. Al-
pine tundra, on lavas.

Santalaceae Sandalwood Family

Geocaulon lividum (Richards.) Fern.

Locality #26, WR 8987, 7 June; do RC 95b,
25 June. Open woods, on alluvium and moraine.

Saxifragaceae Saxifrage Family

Heuchera glabra Willd.

Locality #25, WR 9125, 14 June. Grassy
slope on Bowser Fonnation.

Leptarrhena pijrolijlora (D. Don) R. Br.

Locality #26, RC 94, 95c, 25 June. Grassy
slope, on talus.

Table 1. Summary of taxa collected in northern Brit-
i.sh Columbia.

o

a

<a

a

a

o

o

"3,

■y.

a.

o

_2

s

o

c

o

■T3

s-

_«

c

^^

o

^^

8

j3

,^

S

o

c

rt

a,

u<

6'

Q

TZ

^

Familie.s

2

1

1

1

35

5

45

Cienera

2

I

2

4

89

14

112

Species

.5

2

2

4

167

25

205

Subspecies

0

()

0

0

11

0

11

Varieties

(1

0

0

0

.35

0

35

46

Brigham Young University Science Bulletin

Fig. 42. Mt. Will region at the southern end of the Eaglenest Range. LocaHty 8, near Pass Lake, i.s in a low
saddle carved in lower beds of the Bowser Formation. Mt. Will, between the three glaciers at right center
is in massive Jurassic volcanic rocks.

Parnassia fimbriata Konig.

Locality #2, RC 226, 28 July; #7, RC 2.50,
5 Aug. Alpine tundra, or .serpentine intrusive
and lava.s.

Parnassia pahistris L.

Locality #4, RC 188, 22 July. Glacial mo-
raine, in open wood.s.

Ribes lacustre ( Pers. ) Poir.

Locality #26, WR 9066, 13 June. Glacial
moraine, in open woods.

Rihcs s.la>i(lulosuin Grauer

Localit}' #2.5, WR 9108, 14 June. Gras.sy Bowser Formation.

slope, on Bowser Formation.

Saxifra<s,a jcrni<iinea Grab.

Locality #12, RC 140, 30 June. Alpine tun-
dra, on siliceous conglomerates.

Saxifraga flagellaris Willd.

var. flagellaris

Locality #9, RC 243, 1 Aug. Alpine tundra,
on Bowser Fonnation.

Saxifraga hjallii Engler

Locality #2, RC 2.35a. 28 July; #7, RC 251a,
5 Aug.; #8, RC 2.38b, 1 Aug. Alpine tundra, on
serpentine, intrusive, Jurassic volcanics, and

Biological Series, Vol. 14, No. 4 Reconnaissance of Northern British Columbia

47

Saxifraga nivalis L.

Locality #12, RC 99a, 27 June; #16, RC
26, 19 June. Alpine tundra, on barren shale
slopes of Bowser Formation.

Saxifrage! occidcntalis Wats.

var. rufidida (Small) C. L. Hitchc.

Locality #25, WR 9109, 14 June. Grassy
slope, on Bowser Formation.

Saxifraga oppositifolia L.

Locality #18, RC 131, 30 June; #34, WR
8979, 7 June; #37, WR 9008, 8 June. Alpine
tundra, on marble, volcanic rocks, and lavas.

Saxifraga punctata L.

var. porsildiana (Calder & Savile) Welsh
Locality #12, RC 137a, 30 June. Alpine

tundra, on siliceous conglomerates.

Saxifraga rividaris L.

var. rividaris

Locality #29, WR 9128a, 15 June. Alpine
tundra, in Tertiary Sustut coal beds.

Saxifraga iricuspidata Rottb.

Locality #8, RC 239b, 1 Aug.; #19, RC 12,
13, 17 June; #25, WR 9112, 14 June; #37, WR
9014, S June. Alpine tundra, in lavas and Bow-
ser talus and outwash, and on steep grassy
slopes.

Scrophulariaceae Snapdragon Family

CastiUeja miniata Dougl.

Locality #26, RC 35, 19 June; do RC 76,
25 June; #36, RC 53; 19 June. Open woods,
meadows, and lake shores.

CastiUeja urudaschcensls Hook.

Locality #2, RC 231, 28 July; #3, RC 211,
27 July; #6, RC 198, 26 July; #25, WR 9095,
14 June; #27, RC 117, 29 June. Open woods,
meadows, grassy slopes, and alpine tundra.

Pedictilaris hracteosa Benth, ex Hook.

Locality #25, WR 9117, 14 June; #26, RC
86, 25 June. Steep grassy slopes, on Bowser
Formation. This is the northernmost known
locality for this entity.

Pedicular^ lahradorica Wirsing

Locality #7, RC 252a, 5 Aug.; #26, RC 72c,
86a, 25 June. Open woods, meadows, and alpine
tundra, on volcanic rocks and glacial moraine.

Pedicularis langsdorfii Fisch. ex Steven.

Locality #13, RC 100a, 27 June; #20, RC
19, 18 June. Alpine tundra.

Pedicularis sudetica Willd.

var. gy mnocephala Trautv.

Locality #3, RC 211a, 27 July; #14, RC '9,
25 June; #29, WR 9127, 15 June; #36, RC 46,
19 June. Alpine tundra, open woods, and
meadows, on micaceous schist, coal beds, and
alluvium.

Pensteinon procerus Dougl. •

Locality #10, RC 126a, 29 June; #24, RC
149, 8 July; #31, RC 105, 28 June. Alluvium,
on lake shores, and stream terraces.

Veronica serpyUifoIia L.

var. Iiumifusa (Dickson) Vahl

Locality #2, RC 231b, 28 July; #14, RC
60a, 25 June; #27, RC 121b, 29 June. Alpine
tundra, on serpentine, intrusive, or along streams
and lake shores on alluvium.

Veronica wonnskjoldii

var. wormskjoldii

Locality #7, RC 249a, 5 Aug.; #8, RC
238a, 1 Aug.; #25, WR 9113. 1922a, 14 June.
Alpine tundra and grassy slopes, on volcanic
rocks and Bowser Formation.

Umbelliferae Carrot Family

Angelica hicida L.

Locality #26, WR 9087, 14 June; do RC 73,
25 June. Open woods on glacial moraine.

Heracleum lantuin Michx.

Locality #25, WR 9106, 14 June. Steep,
grassy, south-facing slopes.

Urticaceae Netle Family

Urtica dioica L.

ssp. gracilis (Ait.) Selander

var. /(/rt//i ( Wats. ) C. L. Hitchc.

Locality #25, WR 9124, 14 June. Steep,
grassy, south-facing slope, on Bowser Formation.

Valerianaceae Valerian Family

Valeriana dioica L.

Locality #10, RC 126, 29 June; #31, RC
109, 28 June; #.36, RC 52, 19 June. Wet
meadows, river banks, and lake shores, on allu-
vium.

Valeriana sitchensis Bong.

Locality #3, RC 220, 27 July; #7, RC 251b,
5 Aug.; #25, WR 9093, 14 June; #26, RC 85,
25 June; do RC 164, 11 July. Grassy slopes and
open woods, on alluvium, glacial moraine, and
tiUus.

48

Bhigham Young University Science Bulletin

Violaceae Violet Family
Viola adunca Smith

Locality :ft25, WR 9114, 14 June; #26,
\VR 9054,' 11 Juno; do \\C 40, 19 June; #28,
RC lb, 6b, 16 June; #38, WR 9018, 8 June.
Open woods, and meadow.s, on alluvium and
glacial moraine.

Viola glabella Nutt.

Locality #26, WR 8995, 7 June. Open
wood.s, on glacial moraine. This record repre-
sents an eastern extension of this entity from
the more restricted coastal distribution previous-
ly known.

Viola renifolia Gray

var. hrainerdii (Greene) Fern.

Locality #26, WR 9053, 11 June. Open
woods, on glacial moraine.

PTEROPSIDA-ANGIOSPERMAE-
MONOCOTYLEDONAE

Gyperaceae Sedge Family
Carex aijuatilis Wahl.

Locality #26, RG 26, 25 June. Pond margin,
on alluvium.

Carex deflexa Homem.

Locality #38, WR 9019, 8 June. Dry, open
woods, on alluvium.

Carex hoodii Boott?

Locality #25, WR 9094, 14 June. Steep,
south-facing, grassy slope, on Bowser Forma-
tion. The specimen is somewhat immature and
is difficult to ascribe positively to this entity.

Carex podocarpa R. Br.

Localitv #12, RG 137c, .30 June; #15, RG
17, 18 June; #16, RG 26, 19 June; #20, RG
19a, 18 June; #29, WR 9136, 15 June; #34,
WR 9034, 9 June, .\lpine tundra, in lavas, coal
beds, and siliceous conglomerates.

Carex pruticola Rydb.

Locality #29, WR 91.37, 15 June. Alpine
tundra, on Tertiary Sustut coal beds.

Carex pyrenaica Wahl.

Localitv #18, RG 128b, 30 June; #21, WR
9090a, 14 June; #22, WR 90.58, 9058a, 12 June.
Alpine tundra, on ridge crests, on marble. Bow-
ser Formation, and slate.

C«reA: rossii Boott

Locality #25, WR 9124a, 14 June. Steep,
south-facing, grassy slope, on Bowser P\>nna-
tion.

Carex vaginata Tausch

Locality #36, RG 48a, 19 June. Marshy lake
shore.

Eriophonmi angustifolium Honck.

Locality #26, RG 70, 25 June; do RG 151,
11 July. Pond margin.

Gramineae Grass Family

Calamagrosti.s jnirpiirascens R. Br.

Locality #5, 177b, 27 July. Open woods, on
glacial moraine.

Festuca ultaica Trin.

Locality =:12, RG 14.3, .30 June; #26, RG
66, 67, 25 June. Open woods, meadows, and
grassy slopes, on Bowser Formation.

Hierochloe alpina (Sw. ) R. & S.

Locality #12, RG 142a, .30 June; #29. WR
9142, 15 June. Alpine tundra, on Bowser Forma-
tion and coal beds.

Phletim alpinum L.

Locality it 2, RG 230, 28 July. Alpine tun-
dra, on serpentine intrusive.

Poa laiuita Scribii.

Locality # 12, RG 142, 30 June. Alpine tun-
dra, on siliceous conglomerates of Bowser For-
mation.

Poa Icptocoma Trin.

Localit\' #25, WR 9100, 14 June. Steep,
south-facing, grassy slope, on Bowser Fomia-
tion. This collection represents an eastward
range extension for P. Icptocoma.

Poa paucispictda Scribn. & Men.

Locality #22, WR 90.59, 12 June. Alpine
tundra, on slate.

Trisetuiii spicatiini ( L. ) Richter

Locality #22, WR 9057, 12 June. Alpine
tundra, on slate.

Wihlodca atropurpurca (Wahl.) Fries

Locality #26, RG 68, 25 June. Open woods,
on glacial moraine.

Juncaceae Rush Family

Luzula confusa Lindeb.

Locality =t41, RG 10.3a. 28 June. Alpine
tundra, on alluvium.

Ltizida pariijlora ( Ehrh. ) Desv.

Locality #8, RG 240a, 1 Aug. Alpine lake
shore, on alluvium.

Bkh.oc.icai, Series, Vol. 14. No. 4 Rec;onnai,s,s.\nce of Nohthern Bhu ism Columbia 49

Ltizuhi spicdtu ( L. ) DC. grassy slopes, on Bowser Formation.

Locality #34, WR 8977, 7 June, Alpine fell

field, on lavas. Orichidaceae Orchid Family

Listera cordata ( L. ) R. Br.

Lihaceae Lily Family Locality #26, WR 9072, 14 June. Ground

Streptopus aniplcxifolius ( L. ) DC. '^lyer^ i" spruce wotxls, on glacial moraine.

Locality it 26, WR 9077, 14 June; do RC Uahemma dHataUi (Pursh) Hook.

153, 16/a, 11 July. Open woods, on glacial mo- . ,.^ ,,-_ ,,,„ r.,,o ^^ i ,, r,n nr^

•" - ^ • 6 Locality #25, WR 9118, 14 June; #26, RC

''""^'^" 162, 11 July; do RC 83, 25 June. Marshy areas,

Veratrum eschscholtzii Gray along strea'ms, and lakes, and in open grassy

Locality #25, WR 9105, 14 June; #26, RC slopes, on Bowser Fonnation and glacial mo-

155, 11 July. Meadows and steep, south-facing, raine.

REFERENCES

Gabrielse, H. 1962. Cn,' Lake, British Columbia: Hulten, E. 1968. Flora of Alaska and neighboring

Geol. Survey Canada Map 29-1962. territories. Stanford. California: Stanford University

Gabrielse, H.. and Souther, J. G. 1961. Dease Lake, Press. 1008 p.

British Columbia: Ceol. Survey Canada Map 21- Lord., L. C. 1948. McConnell Creek Map-area, Cas-

1961. siar District, British Columbia: Geol. Survey

Geological Survey Canada. 19.51. Stikine River Memoir 2.51.

Area, Cassiar District, British Columbia: Geol. Sur- Welsh, S. L. 1971. Anderson's Flora of Alaska and

vey Canada Map 9-1957. adjacent Canada. UnpubUshed Ms. 1100 p.

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