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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

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MUS. CO MP. ZOOU
LIBRARY

JAMS t973

HARVARD
JUNlVERSiTY,

Brigham Young University
Science Bulletin

THE HYOBRANCHIUM

AND THROAT MYOLOGY OF THE

ADULT AMBYSTOMIDAE

OF THE UNITED STATES

AND NORTHERN MEXICO

by

John E. Krogh

and

Wllmer W. Tanner

BIOLOGICAL SERIES — VOLUME XVI, NUMBER 1

MAY 1972

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.

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^i^ i

Brigham Young University
Science Bulletin

THE HYOBRANCHIUM

AND THROAT MYOLOGY OF THE

ADULT AMBYSTOMIDAE

OF THE UNITED STATES

AND NORTHERN MEXICO

by

John E. Krogh

and

Wilmer W. Tanner

BIOLOGICAL SERIES — VOLUME XVI, NUMBER 1

MAY 1972

TABLE OF CONTENTS

INTRODUCTION 1

MATKRIALS AND METHODS 4

CENERAL FEATURES OF THE HYOBRANCHIAL SKELETON 4

GENERAL FEATURES OF THE MUSCULATURE 23

SPECIES DESCRIPTIONS 27

P;UMfic Ciant Sahimander. Dicav\ptodon ensalus ( Eschscholtz ) 27

OKmpic Salum.iiulfr. Rhijcicotriton oli/inpicus (Gaige) 30

Ringed SalaniaiHler. Aiiiln/.stiniui cninuhiliuii Cope 33

Rt'ticulated SalaniaiultT. Aiuhijstoiiia c'nt'^uldluin Cope 35

Xoilliwestern Saianiaiuler. Ainhf/.ttoiiui '^rac'dc (Baird) 38

lefferson's Salamander. Amhi/.stonui jcfjcisoixiantivt ( (ireen ) 40

Malx-e's Salamander. Aiuhi/.stiiiiia iiuilwci Bishop 42

Long-toed Salamander. Amhi/.slDitui iiiucnHlticti/hiin Baird 44

Spotted Salamander. Anil)i/.sliiiiiii maculatuni ( Sliaw ) 47

Mole Salamander. Anthi/^itonui talpoidcuin (Holhrook) 49

Marbled Salamander. Anilii/'iloniu d/Kici/m ( (^ra\cnhorst ) 51

Texa.s Salamander. Ainhi/stdnia trxmiinn ( Matthes ) 53

Chihuahua Salamander. Aiuhi/.tloina riisacriiiii Ta\lor 55

Tiger Salamander. Aniln/stoniii ti^iimiiii Creen 36

Ea.stern Tiger Salamander. Amhi/itoina tii:,riitum (ig/iin/Hi (Creen) 56

Northwestern Tiger Salamantler. .\»i/'i/v/i)/ii(( liiiniiinii tiichinofitictiiDi (Baird) 57

Clouded Tiger S:ilaniander. Aiuhi/stonia ti^rinuiii nchiilositiii Hallowell 57

DISCUSSION 59

SUMMARY 66

ack\o\vledc;ments 68

LITERATURE CITED 69

THE HYOBRANCHIUM AND THROAT MYOLOGY

OF THE ADULT AMBYSTOMIDAE OF THE UNITED STATES

AND NORTHERN MEXICO

by

John E. Krogh'

and

Wilmer W. Tanner-

ABSTRACT

The 16 species and subspecies of the Ambys-
tomidae of the United States are placed into
six groups according to their throat morphology
as follows: The Dicamptodon group, the Rhy-
acotriton group, the A. Annulatum-A. cingulatum
group, the A. gracile-A. jeffersoniamim-A. ma-
beei-A. macrodactijlum-A. inacttlatiim group, the
A. taljwideum group, the A. opacum-A. tcxanum
group, and the A. rosaccum-A. t. tigrinuin-A. t.
melanostictum-A. t. nchtdostim group. Dicamp-
todon appears to be the most primitive group
and Rhijacotriton the most specialized. Based

on the morphology of the throat region it is
suggested that A. rosaccum be placed in the
tiger salamander group as a subspecies of A.
ti<'rmum.

The results of this investigation support the
hypothesis that Ambystomidae is primitive to
Salamandridae and to Plethodontidae. The find-
ings which have led to this conclusion are based
upon the protrusibility of the tongue, and upon
the structure and presence of the epibranchial,
otoglossal, second radial, and second basi-
branchial cartilages of the three families.

INTRODUCTION

The ambystomid salamanders are strictly
North American in distribution, ranging from
southern .Alaska and extreme southern Labrador
in the north to the southern parts of the central
Mexican Plateau. Most salamander taxonomists
recognize five living ii^cnera—Diccimptodon.
Rhijacotriton, Rhijacosiredon, Ambijstonui, and
Bathijsiredon. Two of these, Rhijacosiredon and
Bathijsircdon, are found onlv in Mexico. Di-
campiodon and Rhijacotriton are found along the
Pacific Coast of the United States, and the
genus Amhiistoma is essentiallv coextensive with
the entire family range. This study deals with
the ambystomids of the United States, plus one
Mexican species Amhijstoimi rosaceurn. There
are about .30 species in the entire family, and of
this number 13 are found within the boundaries
of the United States.

Most workers believe that Dicamptodon and
Rhyacotriton are related and differ from Amhys-

'University of UUh. Salt Lake City. Utah 84112

^Department nf 7/inIogy. Rrigham YnnnK University, Provo, Utah 84<)01 .

toma in certain features. Tihen (1958) sub-
divides the genus Ambijstonm as follows: (An
asterisk following the name indicates that it is
found in the United States. )

Subgenus: Ambystonia

The mexicanum group
Arnhtjstuinu hansense
Amitijstoma Icrmacnsis
Ari\hiistoma mexicanum

The tigrinum group

Atithi/\toin(i (imhlt/ccplwlum

Anihi/stonui Itotnliypclhiin

Amhijstoma jluvinatum

Amhi/stoniu p^ranulosuin

A m h t/sto ma hihhanli

Arnhijsloma lacustri.s

Aml)tistoma ordincirium

Amhi/stoma rosaccum

Ainhijstinna suhsdlsum

Amhystoma tigrinuvi'

Bricham Young Universitv Science Bulletin

Subge

Subgenus ;

The opacum group
Ambijstoma opacum"
Ambystoma ialpoidciim"

The maculatiim group
Ambysloma gracile"
Ambijstoma jeffcrsoiiiaiium '
Ambystoma laterale'
Ambys-toma macrodactylum °
Ambystoma jiiaculatum'

Linguaelapsus

Ambystoma antiiilatiim'

Ambystoma cin^tilatum °

Ambystoma mabcei'

Ambystoma schmidti

Ambystoma tcxantim'

Bathysiredon

BathysiredoH dumcrili

Tihen (1958) has suggested that the family
Ambystomidae had its origin in the late Meso-
zoic and was probably derived from Asiatic
hynobiids. He further suggests that Dicamptodott
and Rlu/acotrituii were established in the west-
em part of the continent at this early date, and
that the remainder of the family had its begin-
ning at a later date in the northeastern part of
the continent.

There is a lack of agreement among taxono-
mists as to the relationships of the various fami-
lies of salamanders. Some investigators (Dunn.
1926; Noble, 1927) consider the Ambystomidae
to be primitive in relation to the Salamandridae
and seem to be convinced that the Plethodonti-
dae can be derived from the Salamandridae,
whereas others (Regal, 1966; Bishop, 1943)
would derive them from the Ambystomidae.
The primary objective of this study has not
been to help resolve this c|uestion, since our
studies are directed primarily at the generic
and species levels. However, the data do sug-
gest certain conclusions pertaining to inter-
familial relationships, and these are discussed
later in the paper.

Inasmuch as this paper is concerned only
with the hyobranchial apparatus and the myol-
ogy of the throat, it contains some inherent
limitations, and the conclusions which will be
drawn are subject to modification ba.sed on
evidence from other sources.

One of the first workers to studv the anato-
my of the throat region in ambvstomid sala-
manders was Cope (1^^7). The findings of
several subse(juent workers were later sum-
marized by Piatt ( 1940). With respect to cer-
tain structures, our oliservations do not agree
with those made bv these earlier workers, as
will be pointed out hereinafter.

Other nonanatomical approaches to the study
of amphibian evolution have been made in the
past. Some investigators (Boulenger, 1910;
Sampson, 1900; Brandes, 1901; Kerr, 1919; Bar-
bour, 1926) have used developmental patterns
in the study of amphibian evolution. Noble
( 1927 ) contends that, while habits are change-
able among the higher vertebrates, they are the
direct result of inherited organization in the
lower vertebrates, and he suggests that these
habits may even be more stable than many
structural features in these lower forms. Accord-
ingly, he has used courtship behavior extensively,
based on the premise that the more specialized
the behavioral pattern the greater the probabil-
ity that it will be found only in closely related
forms.

One of the obvious manifestations of the
process of evolution is the series of anatomical
changes in a group of organisms which accom-
pany and reflect the changes that occur in the
environment.

The throat region is a fruitful area of anato-
my on which to study salamander evolution
since it is involved in both feeding and breath-
ing mechanisms. Feeding structures associated
with the afjuatic habitat are notably different
trom those found in terrestrial fonns. Terrestrial
fonns arc unable to utilize water for the capture,
manipulation, and softening of food items, but
they have evolved instead a tongue to accom-
plish these purposes in a terrestrial environment.

Hegal ( 1966) has used tongue structure in
the classification of terrestrial salamanders. He
describes the aquatic tongue as consisting of a
mere thickening of the mouth fl(x:)r around the
branchial elements. He groups terrestrial
tongues into three types based on their freedom
from the floor of the mouth, and the ability to
extend the tongue out of the mouth by means of
the hyoid elements. The ainbystomids have a
terrestrial-type tongue that is free at the lateral
and posterior margins but is not as profrusible
as the tongue found in the Salamandridae and
the Plethodontidae. As a result of this structure,
the amb\stomids are restricted to larger food
items. Salamanders with the more advanced
tongues are able to flip the tongue out of the
mouth to catch small, ('lusi\e insects, and, in
addition, thev are able to use the more conser-
vati\e methods for the capture of larger food
items.

Some of the other structures which are also
associated with feeding and breathing mecha-
nisms that have been studied bv various workers
include teeth, skull bones, and \ertebrae (Tihen,
1958); the direction of tooth replacement

Biological Series, Vol. 16, No. 1 Hyobranchium and Throat Myology

(Regal, 1966); and the hyobranchial apparatus
( Cope, 1887; Smith, 1920; Francis, 1934; Tanner,
1952; Wake and Ozeti, 1969).

The present study is based exclusively on the
dissection of adult salamanders. Larval forms
liavc been avoided owing to the great variability
that exists in their development. Brandon
(1961) points to the fact that earlier workers
had concentrated their descriptions on either
newly hatched or fully grown larvae but that
nothing was known for many of the intermediate
sizes. He prepared a table based on balancers,
limbs (buds and toes), and skin pigmentation
to be used to identifv all sizes of larvae.

Dunn ( 1940), in his paper on the races of
Ambt/stovui tigrinum, makes the statement that
larvae and recentlv transformed voung are usual-
Iv useless in racial discrimination. Furthemiore,
the anatomical changes that occur at metamor-
phosis are rather extensive, and the adult stage
of development, therefore, appears to be the
most reliable ontogenetic level at which an
anatomical investigation could be undertaken.
For a cjuantitative treatment of the changes in
weights and in linear measurements of the body
and organs that occur at metamorphosis, the
reader is referred to Latimer and Roofe (1964).

Inasmuch as we have considered in this
paper only those ambvstomids that normallv
occur in the United States (with the exception
noted above), the (juestion of neoteny does not
concern us. On the other hand, a complete
treatment of the familv would include specimens
from populations in which there is complete or
partial neotenv, and an investigator undertaking
such a studv would be forced to compare gilled
animals with nongilled forms. Complete neo-
tenv is found in populations of the tigrinum,
mexicanuin. and Bathtisircdon groups of Tihcn
( 1958), and partial neoteny is found throughout
the family.

We have been able to confirm the platelike
otoglossal cartilage of Dicamptoclon as reported
bv Cope (1887), but our findings concerning
this structure in Amhijsioma gracHe and A. texa-
num do not agree with his observations. The
otoglossal cartilage in these latter two species
is not platelike as in Dicamptoclon but is an-
nular in shape.

We have found the otoglossal cartilage to be
of special importance in the process of feeding.
The vomerine teeth are located on the roof of
the mouth directlv opposite the otoglossal in the
Ambystomidae, and thLs cartilage appears to
function as a rod with which to impinge food
against these teeth. The otoglossal is either
absent or drastically reduced in the Salaman-

dridae and Plethodontidae, and the vomerine
teeth in these two families do not form a trans-
verse series opposite the position of the otoglos-
sal as in the Ambystomidae, but tend rather to
be scattered throughout the mouth. The sala-
mandrids and plethodontids use the entire
tongue to press food up against their teeth,
whereas the ambystomids with their rather im-
movable tongues rely heavily on the otoglossal
cartilage for this purpose.

We have found, contrary to the statements
of earlier workers (Tihen, 1958, for example),
that the majority of the hyobranchium in Di-
camptodon is not solidly ossified, but rather is
mostly cartilaginous. The only structures which
are solidly ossified in Dicamptodon are the tips
of the ceratohyals and the epibranchials. The
second basibranchial cartilage is partially ossi-
fied in most of the specimens studied, but in a
few of them it was completely cartilaginous.
An epibranchial cartilage, the presence of which
is considered to be primitive (Ozeti and Wake,
1969), was observed in all of the specimens
we have studied. The first basibranchial carti-
lage and also the ceratohyal are primitively os-
sified, and thev were at least partly ossified in
most of the species observed in this study.

For the most part, the muscle terminology
we have used is in agreement with that sug-
gested bv Driiner ( 1901), Francis ( 1934), Edge-
worth (1935), and Piatt (1939). We have used
the name interhijoideus posterior for the most
posterior of the ventral hyoid constrictors, and
we agree with Piatt ( 1940) that the terms gularis
and quadratopectoralis should be used only in
the Plethodontidae. Piatt has also stated that
the subhyoideus is a special anterior derivative
of the intcrhyoideus, and that no such muscle
exists in the Ambystomidae. In all of the speci-
mens we have observed, however, this muscle
is present, extending forward from the posterior
tip of the ceratohyal to join the fibers of the
intcrossa<juadrata near the midline of the throat.
In Rhyacotriton, the subhyoideus inserts even
further forward along the posterior edge of the
mandible.

Previous studies on the hyobranchial appa-
ratus and throat myology of the ambystomid
salamanders have been sketchy and incomplete.
The families Salamandridae and Plethodontidae
have been studied relativclv recently (Piatt,
1935; Hilton, l<M7a; Hilton, 1947b; Tanner,
1952; Ozeti and Wake, 1969) and rather com-
pletely, but the same cannot be said for the
Ambystomidae. It has been this need for fur-
ther study which has provided the major stimu-
lus for this investigation.

Bricham Young University Science Bulletin

MATERIAL AND METHODS

The materials which have been used in this
study were made available for dissection by the
Brigham Young University Natural History Mu-
seum, California Academy, Museum of Natural
History, U.S. Natural History Museum, and from
museum collections of other universities through-
out the United States. Numerous specimens were
also obtained from the field collections of private
individuals. Specimens of the Ambystomidae
which we dissected are: Dicamptodon ensatus,
Rhtjacotriton ohjmpicus, Anibijstoma rosaceiim,
A. tigrinum, A. opacuni, A. talpoicleum, A. grac-
ile, A. jeffersonianum, A. nmcrodactylum, A.
maculatum, A. anntdatum, A. cingulatum, A.
inaheei, and A. texunttm. We have also dissected
the subspecies A. t. tigrinum, A. t. nehulosum,
and A. (. melanostictum of the tiger salamander.
For comparison we also dissected one specimen
of Sahniandra salamandra.

In the case of some species there were as few
as two specimens available for dissection, and
in the case of the Long-toed Salamander, Ambijs-
tonia macrodactijlum, only one was available.
We have therefore made no attempt to ascertain
the degree of individual variation within each
species. In making some conclusions concerning
phylogeny, we have attempted to use morpho-
logical structures which tend to have low vari-
ability between individuals of the same species.
All of the dissections were perfonned with the
aid of a stereoscopic microscope at magnifica-
tions varying from 10 to .30 diameters. All mea-
surements were made with calipers to one-tenth
of a millimeter.

The origin, insertion, size, and shape of thir-
teen muscles are described in detail for each

specimen, and the hyobranchium is, likewise,
described for each. The percentages of ossifica-
tion and/or chondrification for the various ele-
ments of the hyoid apparatus are mentioned and
their significance in determining phylogeny is
discussed.

Since most of the muscles are bilaterally sym-
metrical, two depths are shown on some illustra-
tions in order to reduce the number of drawings.
All of the descriptions and illustrations have ref-
erence to adult specimens. Eosin dye was used
only occasionally to distinguish muscular from
connective tissue on very small specimens. A few
specimens were also cleared according to the
Schultz-Potash technique as modified by Han-
sen and Tanner ( 195S ) and then stained with
alizarin red and stored in glycerin. The major-
ity of the specimens have been preserved in 60
percent alcohol and have been accessioned in the
collections of the Museum of Natural History
at Brigham Young University in Provo, Utah.

We have selected 48 morphological charac-
ters, and have used them in the deduction of a
phylogeny for the genus Ainbtjstoiim according
to the method of Wishart (1969). The morphol-
ogy which is included in this paper includes
information on feeding and breathing structures,
and does not attempt to include any of the nu-
merous other body systems or information con-
cerned with life history. The reader is therefore
reminded as to the limitations of this work, and
he is advised that the findings discussed herein
be used in conjunction with information from
other sources in the final decisions concerning
relationships of the Ambystomidae.

GENERAL FEATURES OF THE HYOBRANCHIAL SKELETON

There has been considerable controversy
concerning the tenninology of the hyobranchial
skeleton, as evidenced by the large amount of
synonomy which exists in the literature at the
present time. We have followed Wiedershetm
(1877), Noble (1931), Piatt (1935, 1938), Tan-
ner (1952), Wake (1963, 1966), and Ozeti and
Wake ( 1969 ) as far as hyobranchial skeleton
terminology is concerned. We use the terms
ceratobranchial rather than hypobranchial, epi-
branchial rather than ceratobranchial, first basi-
branchial rather than copula or basihyal, second
basibranchial rather than os triangulare, and ra-
dial cartilages rather than coniua.

In larval forms, the hyoid arch is present as
well as portions of four branchial arches. Fol-

lowing metamorphosis, however, there is a re-
duction of both the hyoid and branchial arch
components, leaving only portions of the hyoid
arch and the first two branchial arches in the
adult. The majority of these stnictiu-es is usually
cartilaginous with centers of ossification most
commonly occurring in the center of the first
basibranchial, the tips of the epibranchiais, the
posterior tips of the ceratohyals, and the medial
portion of the second basibranchial.

The first basibranchial cartilage is the main
supporting element inasmuch as it provides a
base of attachment for most of the other com-
ponents. It is located in a midventral position,
and the centra! portion is usually ossified, ;is
mentioned above. Ozeti and Wake (1969) indi-

Biological Series, Vol. 16, No. 1

HvoBR.-kNCHIUM AND TnHOAT MYOLOGY

cated that the shape of the first basibranchial
was characteristic of each genus in the Sala-
mandridae, and it appears that the same trend
can be found in the genera of ambystomids pre-
sented in this paper. In Dicatnptodon this struc-
ture is truncate, in Rhtjacotriton it is shaped like
an elongated pear, and in Ambystonui it is either
pear-shaped or shield-shaped (Fig. 1-16).

In adults, the hyoid arch consists of two ele-
ments on each side — the first radial cartilage
and the ceratohyal. In the larval stage, these two
structures articulate with each other, but in the
adult condition they are often separated or at-
tached by way of a ligament. The first radial
cartilage attaches to the anterior edge of the
ceratohyal. There are no muscles which attach
to these first radial cartilages. The anterior end
of the ceratohyal is flat and spatulate in shape
while the posterior end becomes rounded in
cross section and curves dorsally to attach to the
quadrate by way of the hyoquadrate ligament.
The anterior end serves as an attachment point
for the subarcualis rectus I, and the posterior end
is the point of origin for the subhyoideus. In the
majority of specimens observed, the posterior tip
of the ceratohyal was ossified. Ozeti and Wake
( 1969) report that from one-half to two-thirds of
the ceratohyal is ossified in salamandrids, and
Tanner ( 1952) reported a complete lack of bone
in these structures in the plethodontids which he
studied.

The two second radial cartilages arise from
the sides of the first basibranchial and also curve
anterolaterally while tapering gradually to a
point. They are usually longer than the first ra-
dials and are typically curved dorsally to artic-
ulate with the otoglossal cartilage. In contrast
to the first radials, the second radials do have
muscles attached to them. This second pair was
shown by Driiner ( 1901 ) to arise at metamor-
phosis. He showed in Salamandra that the sec-
ond pair of radials forms while the first pair is
still present. Later, while investigating Triturus,
Bogoljubsky ( 1924 ) also found a second pair of
radials forming while the first pair was still in
place. The first pair was later lost, and only the
second pair was found to be present in the fully
transformed adult. Ozeti and Wake (1969)
found several genera of salamandrids Chioglossa,
Salaniandrirui, Taricha, Triturus, Notoplithal-
rntis, Cynops, and Paramesotriton) to have only
one pair of radials in the adults, and, moreover,
there were muscles attached to this pair. In
those salamanders which have two pairs of ra-
dials, it seems likely that the second pair is ho-
mologous with the single pair of other genera.

Ozeti and Wake ( 1969) report an "interradial
airtilage" extending between the two second ra-

dials, and a structure by the same name had
been reported earlier by others (Driiner, 1901;
Kallius, 1901; StadtmuUer, 1936). Oppel
(1900) found a ligamentous plate, the "sehnen-
platte," in the substance of the tongue in Sala-
nuuulra maculosa which serves for the attach-
ment of the hyoglossus muscle and also for the
insertion of the abdominohyoideus (rectus cer-
vicis profundus ) . One of the morphological char-
acters that has been used to distinguish primi-
tive salamanders is the otoglossal cartilage
(Dunn, 1926). Cope (1887) figured the otoglos-
sal cartilage in his drawings of the hyobranchial
skeleton of ambystomids and shows this struc-
ture forming an arc between the tips of the sec-
ond radials. We likewise have found this struc-
ture in the specimens which were dissected. It is
a semicircular ring of cartilage, the ends of
which are attached to the tips of the second ra-
dials with the arch directed dorsad into the sub-
stance of the tongue. In some species, it may be
jointed at the midline. It serves as an attachment
for the rectus cervicis profundus and genioglos-
sus muscles. It is hinged at its junction with the
2nd radial and apparently rotates forward with
the radial cartilages as the tongue is extended in
feeding. Tanner ( 1952 ) indicated that the
"lingual cartilage" was missing in all of the
Plethodontidae which he studied, and Dunn
( 1926 ) states that it is missing in all Plethodon-
tidae, although Piatt ( 1935 ) reported finding it
in several plethodontids. We believe that the in-
terradial cartilage referred to above is in reality
the otoglossal cartilage and that the "sehnen-
platte" or "lingual cartilage" is a vestige of this
structure as it exists in salamanders in which the
tongue has attained a greater degree of freedom
from the floor of the mouth than that which is
seen in the Ambystomidae. Its position dorsal to
the first basibranchial, its attachment to the
second radial cartilages, and the fact that the
rectus cervicus profundus also inserts on this
structure in some of the Salamandridae seem to
confirm such a belief.

The first branchial arch is formed by a nar-
row, curved bar of cartilage consisting of the
first ceratobranchial and the epibranchial. The
proximal end of the first ceratobranchial attaches
to the posterolateral edge of the first basibran-
chial and extends posterolaterally and also some-
what dorsally, such that the posterior portion lies
parallel with the ceratohyal. It is a flat oval in
cross section and is wholly cartilaginous in the
ambystomids observed, with the exception of
one specimen in which a small ossification cen-
ter was found (Table 1). Others (Hilton, 1947b;
Ozeti and Wake, 1969), in working with the
salamandrids, have found this element to be well

Brigham Young Univehsitv Science Bulletin

Fig. 1. Dicmnptodon etisutus. Upper left: Supcrtuial throat mu.sciil.iliirc. Upper right: Intermethate muscles of
the throat. Lower left: Deep museles of the tliroat. Lower right; Hvohraiielii:il skeleton.

al, aponeurosis lingualis; hi, first hasibranchial; hll, seeond hasihranehial; el, first ceratobranehial: cll, .second
ceratobranehial; eh, eeratoiiyal; eh, epihranehial; gd. dorsal geiiioglossus; gh, genioh\()ideus; gv. ventral geni-
oglossus; hh, helx)steoypsiloideus; hy hyoglossus; ih, interhvoideus [X)sterior; ip, intemiandihularis posterior; iq,
intero.ssa(iuadrat:i; la, linea alba: m, mandible; og, otoglossal; ogp, otoglossal pl;ite; oh, omohyoideiLs ; p, pec-
toralis; pe, pericardium; rl, first radial; rll, sect)nd radial; rp, rectus eervicis profundus: rs, rectus cervicis
superficialis; s, sternum; sh, subhyoideu.s; sr, subarcualis rectus I; t leirdon

Biological Series, Vol. 16, No. 1 Hvobh.\nchium .-^nd Tiikoai Myologv

Fig. 2. Rhijacotriton olijmpicus. Upper left: Superficial throat musculature. Upper right: Intermediate mu.scles
of the throat. Lower left: Deep muscles of the throat. Lower right: Hyobranchial skeleton.

Bkicham Young Univehsitv Scienc:e Bulletin

Fig. 3. Ambystoma (iriiiuUitiirn. Upper left: Superficial throat miiscniatiire. Upper right: Intermediate muscles of
the throat. Lower left: Deep mu.scles of the tliroal. Lower right: HyobraiKhial skeleton.

Biological Series. Vol. 16, No. 1 Hyobhanchium and Throat Myology

Fig. 4. Amhijstoma cingulatum. Upper left: Superfici.il throat musculature. Upper right: Intermediate mu.scles
the throat. Lower left: Deep muscles of the throat. Lower rigiit: Hyobranchial skeleton.

10

Bhicham Young University Science Bulletin

Fig. 5. Ambtjstoma gracile. Upper left: Superficial throat musculature. Upper right: Intermediate muscles of
the throat. Lower left: Deep muscles of the throat. Lower left; Deep muscles of the throat. Lower right: Hyo-
branchial skeleton.

Biological Series. Vol. 16, No. 1 Hyobr.\nchium and Thhoat .Myology

11

Fig. 6. Ambystoma jefjcrsonianum. Upper left; Superficial throat musculature. Upper right: Intermediate
muscles of the throat. Lower left: Deep muscles of the throat. Lower right: Hyobranchial skeleton.

12

Bhicham VouNt; IInivkrsitv S(:ienc:k Bulletin

Fig. 7. Amhtjstnma mahcci. Upper U-ft: Superficial throat niusculatiirc. Upper right: Intemuihate muscles ol the
throat. Lower left: Deep muscles of llie throat. Lower right: llvol>rani Iii.i! skeleton.

Biological Series, Vol. 16, No. 1 Hv(jiiranchium and Thhoat Myology

13

Fig. 8. Ambystoma macrndactijlum . Upper left: Superficial throat musculature. Upper right: Intermediate
mu.scle.s of the throat. Lower left: Deep muscles of the throat. Lower right: Hyobranchial skeleton.

14

Bhigham ^'ouno Univkhsity Science Bulletin

Fig. 9. Amhijstoma muculatum. Upper left: Supcrfitial throat inusciilaturc. Upper riglit: liitemiedi.ite muscles of
till' throat. Lower left: Deep muscles of the throat. Lower right: Hyohrauehial skeleton.

Biological Sehies. Vol. 16, No. 1 Hyobk.\nchium .\nd Thbo.\t Myology

15

Fig. 10. Ambxjstoma tuljHUdcum. Upper left: Superficial throat musculature. Upper right: Intermediate mu.scles
of the throat. Lower left: Deep mu.scle.s of the throat. Lower right: Hyobrancliial skeleton.

16

Bkicham Young University Science Bulletin

Fig. 11. Ainhi/stonui opaciini. Upper left; Superficial tliroat miiseuhitiire. Upper right: Interinediate inu.scles of the
throat. Lower left: Deep miisc-les of tlie throat. Lower right: Hyohraiichial .ski4eti)n.

BioLor.K:AL Series, Vol. 16, No. 1 Hv()1)h.\N{:iiium a.nd Thro.m Mvoi.ocy

17

Fig. 12, Ambijstoma tcxanum. Upper left: Superficial throat mu.sculature. Upper right: Intermediate muscles of
the throat. Lower left: Deep muscles of the throat. Lower right: Hyohranchial skeleton.

18

Brigham Voung Univehsity Science Bulletin

Fig. 13. Amhijatoma rosaccum. Upper loft: Supcrlici.il throat nuisculaturc. Upper right: Intermediate muselcs of
the tiiroat. Lower left: Deep muscles of the throat. Lower right: Hyohranchial skeleton.

Biological Series. Vol. 16, No. 1 Hvohh.wchium and Thhoat Myology

19

Fig. 14. Anthtjstonm tigrimim. Upper left: Superficial throat mu.sculature. Upper right: Intermediate mu.scle.s of the
throat. Lower left: Deep mu.scle.s of the throat. Lower right: Hyobranchial skeleton.

20

Bhicham Young University Science Bulletin

Fig. 15. Amhi/stonui (i>nrii/m mcltirwsticttwt. Upper left: Superficial throat miisciilatiire. I'ppcr right: Interme-
diate rmi.s(.le.s of tlio throat. Lower left: Deep muscles of the throat. Lower right: Hyohranchial skeleton.

BioLOGiCAi, Series. Vol. 16, No. 1 Hyohh.vnciiium .\nu Thho.\t Myology

21

Fig. 16. Amhystoma tigrinum nehulosum. Upper left: Superficial tliroat musculature. Upper right: Intermediate
muscles of the tliroat. Lower left: Deep muscles of the throat. Lower right: Hyobranchial skeleton.

22

Brigham Young Univehsity Science Bulletin

%#-'

■ig. 17. Representative liyobranchial skeletons of Amin stoniid le, Salainaiulridae, and I'lethodontidae. Upper left:
/:)i(vjm/)^i(/()ri cm-Hfu.v, Upper ri,i;lit: Amhi/stavKi taUioidcum. Middle left: Siihimandra suUimamlra . Middle
right: Taricha toro.sd. Lower left: Ko/idig/oswi rufcsicii.s. Lower right: Thorius ;i«ri.S()t«/i,v. Middle left and
middle right from Ozeti mid W;ike (1969), Lower left and lower right from Tiinner (1952).

Biological Sehies, Vol. 16, No. 1

HvOIinANCIIIUM AND TlIROAT MVOLOCV

23

ossified. The distal end of the first ceratobran-
chial attaches to the proximal end of the epibran-
chial, the articulation of which is usually per-
ceptible only as a thin line separating these two
elements. There is no second epibranchial. The
posterior tip of the epibranchial is ossified in nine
of the fourteen species which we have dissected
and serves as an attachment for the subarciialis
rectus I muscle. Ozeti and Wake ( 1969 ) report
that the epibranchial is usually almost entirely

Table 1. .\verage percent ossification of hyobranchial
elements.

Species

bl bll cl cll rl rll ch eb

Dicamptodon

cnsatus
Rliiiacolritoii

olympicus
Amhijstoimi

(inmilatum
Ainhi/stonni

cinguldltini
Anihtjstoma

grdcilc
Amhi/stoma

jcffcrsonuinum
Ambi/stomu

mahcci
Ambijstoma

macrodactijlum
Andit/stoniii

macnUitum
Amht/stonni

t(dpoidcuin
Ambiistoma

opacum
Amhi/stoma

tcxamn7i
Aiid>ijstonii\

rosaccttm
Arrd)t)stomti

tigrinurti
Ambystomii tigrinum

mclanostictum .37. .5 71 0.0 0

Arnbi/slonui tifirinum

nebulosurn 59.0 .59 0.0 0

0.0 0 0.0 0

0.0 0 0.0 0

77.0 43 0.0 0

73.0 62 2.2 0

0.0 78 0.0 0

.37.4 80 0.0 0

37.0 63 0.0 0

34.0 50 0.0 0

25.0 73 0.0 0

32.6 76 0.0 0

41.0 0 0.0 0

.38.0 65 0.0 0

28.0 69 0.0 0

40.0 72 0.0 0

0 0 15.0 56.5

0 0 0.0 0.0

0 0 0.0 12.0

0 0 5.0 15.0

0 0 0.0 0,0

0 0 6.4 39.7

0 0 6.8 .58.0

0 0 5.0 0.0

0 0 10.6 .32.0

0 0 3.6 0.0

0 0 2.7 11.0

0 0 3.3 0.0

0 0 4.8 .36.0

0 0 5.0 17.0

0 0 0.0 21.0

0 0 8.0 60.0

ossified in the Salamandridae but that some gen-
era have lost this element or have experienced a
fusion of the latter with the ceratobranchial.
These investigators also report that the epibran-
chial is longer than the ceratobranchial, but we
have foimd that the ambystomid epibranchial is
consistently shorter than the first ceratobranchial
with which it is articulated (Table 13).

The second branchial arch consists of a pair
of slender, curved bars of cartilage which articu-
late proximally with the posterior end of the
first basibranchial and curve posterolaterally to
articulate distally with the first branchial arch
at a point about two-thirds along the latter's
length. It is entirely cartilaginous in all speci-
mens which we have observed. The space be-
tween these two arches is covered by the inter-
cartilaginous membrane (Driiner, 1901) except
for the mesial portion through which the rec-
tus cervicis profundus passes.

The second basibranchial is a small element
located in the tendonous inscription which sep-
arates the geniohyoideus from the rectus cervi-
cis superficialis. It usually assumes the shape of
a three-pointed star in which one arm is directed
anteriorly with the other two arms pointing pos-
terolaterally. In most of the specimens studied in
this paper, the second basibranchial is cartilag-
inous on the tips of the lateral arms, the re-
mainder being ossified. In a few specimens, the
entire element is cartilage. In the larval state,
this structure is connected with the first basi-
branchial by a slender median rod which disap-
pears at metamorphosis, leaving the two basi-
branchials separated from each other. In most
genera of the Salamandridae, the second basi-
branchial is lost (Ozeti and Wake, 1969), and
in the Plethodontidae it is reported by some
authorities as completely missing (Dunn, 1926;
Tanner, 1952; Wake, 1966), although Piatt
(1935) recorded its presence in several pletho-
dontid genera.

GENERAL FEATURES OF THE MUSCULATURE

The following paragraphs constitute some
general statements as to position and size of the
throat muscles. More detailed descriptions will
be found in the following section which deals
with each species.

Intermandibularis Posterior

This muscle is found just under the skin of
the throat and covers about two-thirds of the area
between the two rami of the mandible. There

are two intennandibularis muscles in the larva,
a small anterior portion, and a larger posterior
portion. It is the posterior portion that persists
in the adult, and the anterior portion, the inter-
mandibularis anterior, is lost at the time of trans-
formation, or if it persists it is so completely
fused with the posterior muscle that no separa-
tion can be made in adults. The origin of the
intermandibularis posterior is along the dorso-
medial surface of the mandible. In most speci-
mens there is a small open area at the angle of

24

Bbiciiam Young University Science Bulletin

the chin wliere there are no fibers and the ven-
tral portion of the genioglossus can be seen upon
removal of the skin. The insertion is at the mid-
line fascia of the throat— the linea alba. The
width of this fascia is variable among the several
species studied. In Rhyacotriton and A. gracile
the fibers of the anterior portion extend with-
out interruption across the midventral line. Also,
in A. clngulatum the anterior portion of the in-
termandibularis posterior is noticeably separate
from the large posterior portion of the muscle.
The anterior portion of the interossa(|uadrata is
covered ventrally by the posterior fibers of this
muscle.

This is one of the least variable muscles of
the throat area. Contraction results in the raising
of the floor of the mouth, which is a component
of the breathing mechanism. This action also
helps to force food items up against the vomer-
ine teeth in feeding.

Table 2. Numeric Characters'

Inlerhyoideus Posterior

It has been suggested (Piatt, 1940) that this
muscle in the Plethodontidae gives rise to two
adult muscles, the gularis and the quadratopec-
toralis. In the ambystomids we have seen no
tendency toward subdivision of the intcrhyoideus
posterior, and it would therefore appear that this
muscle is undifferentiated as noted by Piatt
(1940). The origin is in fascia which attaches
to the tip of the mandible, the quadrate, the
s(|uamosal, and the otic capsule and is continu-
ous with the fascia cephalodorsalis. The fibers
extend ventromesial to insert into the skin of the
gular fold and the fascia along the midventral
Vme.

This muscle would appear to function in
swallowing food and in breathing, inasmuch as
it is in a position to constrict the pharynx. It
might also act in head flexion, since the posterior
fibers are attached to the skin of the gular fold.

Species

Museum
Number

Attribute Numbers

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

18

17

18

19

20

21

22

Dicamptodon

32467

0.50

0.75

1.29

0.52

0.21

0.55

0.27

0.47

2.09

0.52

0.93

1.10

0.53

0.55

0.35

0.72

0.36

litiir

00.0

00.0

16,0

60.0

Dicampiudon

3i465

0,49

0,71

1.09

0.55

0,21

0.45

0,25

0,40

1,90

0.55

1.21

0.91

050

0,53

0.29

0.71

0.39

00.0

00.0

00,0

140

.530

Hh

tjacotritoii

1248U

0.40

0,&4

1,03

0,80

0,27

0.50

0.29

0,27

1,07

0,54

1.15

1.17

0,63

0,35

0.18

091

037

00.0

00 0

00,0

000

0.00

Kh

tjactttritini

12479

0.35

0,62

0,97

0,89

0,33

0,38

0,21

0,33

120

0,50

1,26

0.85

0,47

0,29

031

1.16

0.31

00,0

(X).(l

00,0

0,00

000

A.

annulatum

32521

0.38

0,67

1,02

0,83

0,23

0,46

0,27

0,43

0,62

0,60

1,48

1.15

0,69

0,39

0.29

0.73

0.39

730

48,(1

00(1

00,0

20.0

A.

»nnulatlttn

.TIS'.I

0.37

0,67

1,06

0,83

0,24

0,45

0,26

0.58

0,71

0.58

1,39

1.08

0.63

0,40

0 40

072

0-38

78()

.54,0

00(1

00,0

07.0

A.

annulatum

32523

0.36

0,66

1,10

0,91

0,21

0,48

0,25

0,55

0,55

0.53

1,14

1.33

0,71

041

0 43

0,67

0.37

84,0

42,0

00,0

00,0

07.0

A.

annulatum

32524

0.39

0,67

1,09

0.97

021

041

0,26

071

081

0.63

1,70

1.25

0,79

0,41

0.24

0-71

0.39

75,0

32,0

ooo

00,0

10.0

A.

annuLilum

32525

0.39

0,63

1,03

1.05

0,24

0,49

0,27

0.59

0,77

O.M

1,18

1.22

0,66

0,37

0,35

071

0 35

78,0

42,0

000

00,0

16.0

A.

cingulatum

33307

0.42

0,71

1,11

0.75

0,25

0,50

0,29

0,72

0,22

0.57

1,33

1.17

0,67

0,46

0,30

(1,77

0,32

60,0

6O0

00,0

09,0

18,0

A.

cingulatum

33309

044

0,75

1,13

0,71

0,23

0,43

0,26

0,93

0.33

0.61

1,59

1.20

(1,74

0,45

0,48

082

034

72,0

52,0

000

000

28,0

A.

ciuf^ulatum

33310

0.39

0,71

0,89

0,77

0,21

0.43

0,26

0,80

0,33

061

1,59

1-09

0,67

0,48

041

0.71

0.33

90 0

63,0

00,0

08,0

00,0

A.

cin^uUituvt

33312

0,47

0,69

1.14

0,86

0 21

0.45

0.31

0,73

0.27

0.67

2,06

1,12

0,76

0,.5O

0,41

0-77

035

78(1

71,0

00,(1

000

00,0

A.

cin^ulatuin

33308

0.40

0.67

1.03

0.82

0.21

0.46

0.28

0,69

0.38

0.60

1.52

1,17

0,71

0,48

0.58

066

0.37

67.0

67,0

11,0

08.0

27.0

A-

gracile

32503

0.39

0.67

0.94

0.46

0,23

0.49

0.27

0,34

0,92

0,55

1.24

II 1

0,61

0,57

0.35

0.67

0,35

00.0

82,0

00,0

ooo

00.0

A.

gracUf

.32504

038

0.69

0,99

0.55

0,24

0.55

0.27

0,46

0,86

0,.50

1.00

1,14

0.57

0,49

0.45

0 78

0.32

00,0

74,0

(X),0

000

00.0

A.

jefjcTsonitmum

32487

0.46

0,64

1,12

0.42

0,26

0.62

0,32

05I

0,63

052

1,09

1 18

0,61

0,49

0.30

0.64

0.32

360

9l,n

000

08,5

27,5

A.

jejfersonianum

32489

0.44

0,68

1,00

0.44

0,25

061

0,31

0,46

0,66

0.5(1

1,00

1,12

0.56

0,52

0,55

0,69

0 26

345

85.0

00,0

06,1

32,0

A.

ji-j^CTstinwnum

32488

0.47

0,64

0,93

0.44

0,24

0,55

0,31

0,49

0,46

0.56

1,26

1,12

0.63

0,53

0,43

0,63

033

,39 4

80.0

00,0

06,1

48,0

A.

jcficTsonumum

32508

0.49

0,70

1,12

0.42

0,25

0,57

0,31

0,41

0,68

0,55

1,20

1,12

061

0.53

0,45

0,69

0.28

39,(1

82.0

00,0

05,9

45,0

A.

jefjersonUinum

32.509

0.45

0,62

0,89

0,49

0,24

11,59

0,3 1

0,55

0 82

0.52

1,08

1,19

0.61

0.52

0,67

0.71

028

38,5

76,0

00,(1

05 4

36,0

A.

maheei

33313

0.40

0,68

1,18

0,71

031

0,56

0,32

0.56

1,00

056

1,28

1,22

0.68

0.37

0,30

0.92

0.34

.39,0

70,0

(KIO

06 0

45,0

A.

ntaticci

33314

0.41

0,67

126

0 75

028

0,57

031

0 48

0,86

0.54

1,19

1,03

(1.56

0,44

0,36

0.75

0.36

38,0

a5,o

00 0

07,0

73,0

A.

nuiha-i

33318

0.37

0,65

1,12

0,88

0,30

0,46

0,24

0.67

1,33

0.53

1,11

1,10

0.58

0,33

0,40

1,02

0.31

39,0

65,0

(K),0

08,0

64 0

A.

nmbcci

33317

0,39

0,64

1,13

0,6«

0,26

0,50

0,25

0.63

0,95

0,51

1,03

1,08

0„55

0,42

046

0,85

0.38

37,0

63,0

00.0

08,0

55 0

A.

maheci

33315

0.40

0,65

1,19

0,63

0,25

0,49

0,25

0.56

1,17

0,52

1,09

III

0„58

0.47

0,44

081

0,35

45,0

56,0

00,0

05,0

56,0

A.

maciotlacttjlum

13735

036

0,67

1.01

0,56

0,24

0,67

0,36

0.36

0,91

0.54

1,18

1,15

0,62

0.53

0,54

0.62

0,32

34,0

50,0

00,0

05,0

00,0

A-

nuiculaium

23311

0.40

0,71

100

0,6!

0,26

0,51

0,29

0.49

0.85

0,57

1.33

1,08

0.62

0.52

0,19

0.69

0,32

31.0

74,0

00,0

155

36.0

A.

nuicutatunt

32512

0.43

0,64

1,07

0,53

0,30

0.60

0,33

0.56

0,68

056

1.27

1,26

0 71

043

0,26

0.67

0,23

25.0

72,0

00,0

11.0

28.0

A.

maculatum

.32515

0.41

0,66

1,09

0,55

0,28

0.57

0,29

0.45

0,58

051

1.05

1,18

060

0,47

0,30

0,75

0,32

260

67,0

00,0

08.6

24.0

A.

maculatmn

.32516

0.40

0.62

1,07

0,68

0,28

0.57

0,30

0.49

0,63

0.53

l.ll

1,13

0.59

0,42

0,29

0,77

0,34

18,0

78,0

(X),0

08.6

38.0

A.

tjuiculatum

32517

0..38

0,64

1,07

0,60

0,29

0,61

0,31

051

0,50

0.51

1.05

1,26

(1.64

0.45

0,36

071

0.32

26,0

74,0

00,0

09,5

35.0

A.

talpoidfum

1421.35

0.46

0,61

0,92

0,58

028

0,.57

0,30

0.52

0,85

0.52

1.09

1,14

0.59

0.56

0,25

0,70

0.36

.36.5

80,0

00,0

11,0

00.0

A,

talpoidcunt

142136

0.45

0,62

0,89

0,60

026

0,57

0,28

0,48

0,64

0.50

1.00

0,98

0.49

0.51

0,.54

076

0.31

250

77,0

(KIO

ooo

OO.O

A.

talpoitleum

142137

0.42

0,65

0,78

0,73

0,.38

0,71

0.34

0,44

0,78

0.48

0.93

1,18

0.57

0.39

0.58

1,00

0.37

36.5

72,0

(10,0

00,0

OOO

A.

vpacum

23310

0,33

0,72

0,92

0,5()

0.34

0,72

0.37

0.50

0,60

0.52

1.08

107

0..56

0.48

0.20

075

0..36

.53.5

(K),(l

OOO

00,0

oo.o

A.

opacum

.325 U

0,38

0,62

1,10

0,69

0.41

0,88

0,43

0,51

0,68

0,48

0.93

1,24

0.60

039

0.27

0,88

0.28

37,0

(H),0

(100

05,8

(Hid

A.

opactitn

32510

0,43

0,6«

1,16

0,58

030

0..57

0.32

0,62

0,81

0.56

1.28

1,04

0,59

0.42

O.M

0,70

0.37

51,0

(KID

00,(1

00,0

00,0

A.

opacum

32313

0.48

0,67

0,96

0,66

031

0.59

0.34

0.54

0,73

0.57

1.33

1,18

0,68

0.46

0.31

0,86

0.28

650

00,0

oo.o

05,3

45,0

A.

texanum

13098

0,43

0,65

1,04

0,06

1)26

0..57

0,30

0,70

1.20

0.53

1.13

1,12

0,.59

0,44

018

0,69

0..33

36,5

69.0

00,0

00,0

00,0

A.

tcianum

13099

0,39

0.63

099

0,74

0.31

0.6«

0,36

0,43

1.05

053

1.15

1,26

0.67

0,44

0..32

0,69

0.51

35,0

59.0

00,0

00,0

00,0

A.

tcuinum

13104

0,40

0,67

1,06

0,61

11,25

0.51

0,29

0,74

0,95

0.57

1..32

1,16

071

0,48

0.45

0,71

040

.39.0

53,0

00,0

04,3

00,0

A.

texanum

13101

0,39

0,67

am

0,76

0,29

0.62

0,35

0.78

1,00

0.57

1.32

1.24

0.71

0,40

0.40

0,85

038

36.0

79,0

00,0

07,0

00,0

A.

texanum

13102

0,36

0.63

0,99

0,69

022

0..50

0,28

0,81

1 (10

0..56

1.28

1.09

OOl

0,45

0,30

0.70

0,29

43,0

67,0

000

05,5

000

A.

TOiaceum

32507

0.45

0.65

1,11

0.37

0,25

0.59

0,32

0,42

1,13

«..54

1.17

1.17

(1.63

0,67

0,44

0.63

0,27

32,0

61,0

00,0

08,5

64,0

A.

rosaceum

,)2.5(X)

0,40

0.70

1,07

0.40

0,27

0..54

0.30

0,41

0,82

0.55

1.23

1.17

0.65

0,.55

0,47

0.67

0,27

28,0

71,0

00,0

06,4

44.0

A.

Tosaceum

32505

0,46

0,69

0.97

0.51

0,26

0.61

031

0,46

0.92

0.52

1.07

1,18

0.61

(1.55

0.39

0.73

0,24

24.0

74,0

(K1,0

00,0

00,0

A.

1. tigrinum

31462

0,42

0.66

0,92

0,52

0,27

0.61

0..34

049

0.80

0.56

1.27

1,05

0.59

0,64

0,50

0.61

0.30

43.0

69,0

ooo

050

19,0

A

t. tigrinum

32513

0,44

0.68

0,97

0,49

0,28

0.61

0.32

0,46

074

0.53

1.11

1,01

0,5.3

0.53

031

0.65

036

39,0

73,0

000

06,6

200

A

t- tif^rinum

.32514

045

0.65

1,03

0,39

0,29

0.63

0,34

0 40

062

0,a5

1,23

1,01

0.56

0.54

0,35

0.70

0.33

38,0

74,0

ooo

0.3,6

140

A.

t. mclanoslictunt

703

0,44

0.59

0,88

0.40

0.23

0..50

0,26

0,54

1.07

0.52

1 06

I 00

0,55

0.58

0,25

0.85

0.28

40,0

70,0

(KIO

00,0

19,5

A-

t. mcLinostictum

717

0,46

0.67

0,93

0,41

025

0.01

0.30

0,55

0,65

(1,49

0,97

1,03

0,51

0.61

0,39

0.81

0.23

.35.0

72,0

00,0

ooo

22,0

A.

t. ncbulostim

15103

0,39

0.49

0,94

0,47

0.27

0.62

0.34

0.52

0,70

0,.53

1,11

1,07

0.56

0,.58

0,46

0-79

0.33

63.0

70,5

00,0

08,0

55,0

A.

t. nchutosum

32494

0,46

0.65

0,92

045

024

0,52

0,27

0 43

0,85

0.52

1,10

1.02

0..54

0.64

0,46

0-78

030

70.0

63,0

(K),0

06,7

630

A

t. ncbulosvm

32493

0,40

0..59

0,S7

0,47

0.32

061

0,32

0.39

0,72

0.52

1,09

1.00

0.52

0.53

0,32

0.83

0.32

.53.0

59.0

000

10,5

56.0

A.

t. nehulosum

15105

0.39

0.57

0,93

0.46

0.25

0,53

0,27

0.61

0,88

0.51

1.05

1.02

0.52

0.54

0.44

0.81

0.35

55.0

52.0

00,0

08.0

48.0

A

t. iiebulasum

32S20

0.45

0.62

0.94

0.44

0.23

0.56

0.27

0.52

0.96

0.49

0.94

l.OS

0.53

0.63

0.38

0.72

0.35

57.0

52.0

00.0

06.7

69.0

'Sec test for explonatiaii.

BiOLooiCAL Sehies. Vol. 16, No. 1

HvoliKANCHIUM .AND ThROAT MYOLOGY

25

Table 3. Binary Characters'

Character Numbers

Species

Dicamplodon

Rfujacotritim

A. (itmuUiltim

.\ cin^uUiUifii

A. ^racile

A jefffrsoniamtin

A. nuihcci

A. tnuirotiuctl/luii

A nuuiilatuiii

A Uitiioidiitni

A. iijmiutn

A. trxiiituin

A. riisiKCttni

23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 ^ 56 57 58 59 60 61 62 63

0 0 1 0 0 i i 0 1 1 i I 0 10 0 i i o oo loooooi iooioof)ioooio

110 10 0 1
I 0 I) n ) 0 II

10 0 0101 OOIOOI 1 001 1010101(1

I II 1 0 I 1 0

0 I) 0 10 0 1 II

n (I (I I II n 1 II

I) 1 I 1 II I)

0 10 1

0 1 I) 1 0 1

t 1 II 0 0 I

I 0 0 II II II 1 II

1 II I) II I) 0 I n

I II O II I) II I 0

I 1 II 0 I I (I ] II 1 I 1 o 1 O (1 0

I 0 II ] I It

1 0 1 (1 0 1 1 II 0 II 1 0 I 0 0 (1 0 I)

1 II 11 1 I 0 1 II 1 0 I) ! 0 I) 0
1 II 0 I [ 0 I (I I I) 0 I (I 1) 0

0 1 0 II 1 I) 0 II

I 1 n 1 1 0

1 i (1 I 0 n II

I n 0 <i 0 1

1 II 1 1 II I

I II I II 1 O II ] 0 0 0 I)

10 0 1 II II 0 1 0 II 0 I II 0 (I

A. (ig
A. I. mt-

hinostictum
'htilosu fji

0 0 1 (1 n 0 1 0 0 0 10 10 1

0 0 1 0 0 0 0
II 0 I 0 II II 0

I (1 0 II 1 0 ] II I) II 0 0 I n

I II 0 II 1 0 1 0 II 0 0 (I I (I

Olio I)

I 10 0 II II 0

0 0 1 1 II [ O I 0 I 10 0 0 0

0 II 1 1 II

0 II 0 0
0 II II I)

0 I 1 II 0 O O 1 1 0 0 II II II 1 II I 0 1 1 0 II 1 1 0 1 0 0 0 0

O II 0 0

'See text for cxplanatioii-

Depressor Mandibulae

The depressor mandibulae originates on the
squamosal, the otic capsule, and the fascia ceph-
alodorsalis and inserts on the posterior tip of the
mandible. It is the major jaw-lowering muscle
in salamanders.

Levator Mandibulae

This muscle consists of four portions, the
levator mandibulae extemus, the levator mandib-
ulae posterior, and the levator mandibulae an-
terior, which is further divided into a superfi-
cial portion and a deep portion. The fibers in-
termingle considerably near the origins, and
separation is rather difficult in some specimens.

The levator mandibulae extemus lies lateral
to the mandibular branch of the fifth cranial
nerve and completelv covers the deeper portion
posteriorly. The fibers originate from the lateral
margin of the squamosal and from the otic cap-
sule, and they insert on the dentary and coro-
noid process of the prearticular. The levator
mandibulae posterior is a short, triangular mus-
cle which lies mesial and posterior to the man-
dibular nerve, originating on the squamosal,
quadrate, and pterygoid and insertijig on the ar-
ticular and prearticular. Its distal fibers blend in
with the tendon of the levator mandibulae an-
terior. The levator mandibulae anterior lies me-
sial to the mandibular branch of the fifth cranial
nerve. The superficial portion, or temporalis, is
a long, pinnate muscle arising by way of a ten-
don from the neural crest of the cervical verte-
bra and from fascia which is attached to the
lateral margin of the parietal bone. The fibers
extend in an anterior direction over the dorsal
surface of the parietal and then turn sharply
ventrad in front of the ear capsule to insert on
the coronoid process of the prearticular. The deep
portion of the levator mandibulae anterior orig-
inates on the lateral border of the frontal and
parietal bones. The fibers of this deep portion
blend in with those of the superficial portion,
and together they insert on the dorsal margin of

the prearticular. The fibers of this short, fan-
shaped muscle extend in an almost vertical direc-
tion.

With respect to origins and insertions, the
levator mandibulae is one of the most uniform
muscles of those that are considered in this
study. The size of this muscle varies consider-
ably within the family as seen in Dicamptodon
and Rhijacotriton, in which it is hypertrophied
to the extent that part of it can be seen from a
ventral view of the throat. Its action is to close
the mouth by raising the mandible.

Interossaquadrata

According to Piatt ( 1940 ) , this muscle, along
with the subhyoideus, is derived from the inter-
hyoideus. He further states that the subhyoideus
and interossaquadrata are not found in the
Ambystomidae; but, as mentioned in the intro-
duction, we have found both muscles in each
specimen dissected. The interossaquadrata is
thick and narrow at its origin but spreads into a
thin, fan-shaped muscle at its insertion. It origi-
nates on the ventral surface of the quadrate or
on the hyoquadrate ligament which attaches the
tip of the ceratohyal to the ((uadrate. The most
posterior fibers may also attach on the tip of the
mandible and lie ventral to the other fibers.
These ventral fibers run caudo-mesiad, whereas
the dorsal fibers extend anteriomesial. The fi-
bers may meet at the midline in some specimens,
or thev may insert into the linea alba as the lat-
ter continues posteriorly from the intermandibu-
laris posterior. The medial ends of the anterior
fibers lie dorsal to the intermandibularis poste-
rior.

Contraction of this muscle results in con-
striction of the hyobranchial skeleton and
posterior mouth cavity.

Subhyoideus

The subhyoideus originates along the pos-
terior tip of the ceratohyal. In Rhijacotriton this

26

BmciiAM Young University Science Bulletin

line of origin extends a considerable distance an-
teriorly along the lateral margin of the cera-
tohyal. The insertion is also atypical in this
species, being attached to the posterior edge of
the mandible just lateral to the symphysis. In
other ambystomids that are included in this
study, the subhyoideus inserts along the dorsal
surface of the linea alba and into fascia over
the anterior end of the ccratohyal. The fibers
fan out slightly so that the insertion is broader
than the origin.

This muscle is apparently able to assist in
bringing the hyobranchial apparatus forward in
feeding.

Geniohyoideus

Originating on the posteroventral surface of
the mandible just lateral to the symphysis, this
muscle passes caudad as a rather narrow but
stout band to insert onto the second basibran-
chial and onto the tendonous inscription that ex-
tends laterally from the base of the second basi-
branchial. The geniohyoideus can be seen in
some specimens beneath the transparent linea al-
ba. In many specimens, the anterior ends of these
two muscles are separated, and the ventral por-
tion of the genioglossus can be seen between
them at the angle of the chin. The lateral edge
of the genioglossus partially overlaps the origin
of the geniohyoideus ventrally. In Dicamptodon,
the two geniohyoideus muscles are not separated
anteriorly.

This muscle acts to depress the lower jaw or
the entire head. It also advances the second
basibranchial cartilage.

Genioglossus

There are two parts to this muscle. The ven-
tral portion fills the angle of the chin and orig-
inates on the posteroventral edge of the mandi-
ble, beginning at the symphvsis and extending
laterally for a millimeter or two. Tlie fibers ex-
tend posteromesially to fonn a small triangle-
shaped muscle at the point of the chin. These fi-
bers insert into the aponeurosis lingualis which
connects the root of the tongue with the first
basibranchial cartilage. A few mesial fibers may
also continue caudad for a short distance to at-
tach to the anterodorsal edge of the first basi-
branchial and onto the otoglossal.

The dorsal portion of the genioglossus arises
from the posterior edge of the mandible im-
rnediatelv dorsal to the origin of the ventral por-
tion. The geniohyoideus must be removed in or-
der to see the entire muscle since the latter over-
laps the dorsal portion. If one were to proceed
dorsad from the lateral edge of the ventral por-

tion of this muscle, he would encounter the ven-
tral portion of the genioglossus, the geniohyoid-
eus, and the dorsal portion of the genioglossus.
The dorsal portion spreads out as a fan over the
floor of the mouth. The medial fibers often pass
posteriorly to insert onto the tip of the second
radial, while the lateral fibers continue in a
posterolateral direction to insert into the sides of
the base of the tongue.

Contraction of this muscle helps to pull the
tongue out of the mouth. It may also act to com-
press the glandular tissue of the tongue prepara-
tory to opening the mouth and extending the
tongue in feeding.

Subarcualis Rectus I

This muscle originates on the tip of the epi-
branchial, around which the fibers wrap, form-
ing a muscular sheath. It then extends antero-
medially, parallel with the ceratohyal to insert
on the expanded anterior end of the latter and,
therefore, loosely binds the epibranchial and the
ceratohyal together. Along with the subhyoideus,
this muscle acts to bring the hyobranchial ap-
paratus forward in feeding.

Rectus Cervicis Superficialis

This muscle originates on the dorsal surface
of the sternum and extends dorsal to the anterior
end of the sternum, and to the coracoid carti-
lages. It continues forward as a thin, broad sheet
ventral to the pericardium to attach to the second
basibranchial and the tendonous inscription from
which the geniohvoideus arises. It then continues
forward as a thick, narrou' band on either side
of the midline to insert onto the posterior edge
of the medial end of the ceratobranchial and
onto the posterior end of the first basibranchial.
The portion between the sternum and the second
basibranchial has three transverse tendonous in-
scriptions in all of the specimens that are in-
cluded in this paper, except for Rhtjacotriton
and A. taljioideum. in which there are only two.
The omohyoideus is a muscle associated with the
rectus cervicus superficialis. It arises from the
ventral surface of the scapula and extends to the
lateral edge of the rectus cervicis superficialis,
attaching to the latter along the margin between
the second basibranchial cartilage and the first
or second tendonous inscription posterior to the
second basibranchial. The deep fibers of the
two rectus cervicis muscles intermingle consider-
ablv dorsal to the insertion of the omohyoideus.
The omohyoideus is not included in the descrip-
tions of each specimen. It is the only muscle
of the throat region that links the hyobranchial

Biological Series. \\)l. 16, No. 1

HvoBR.\Nc:niuM .^nd Tubo.^t Myology

27

apparatu.s with the cartilages of the pectoral
girdle. The omohvoideus serves to brace the rec-
tus cervicis superficialis and to adduct the
shoulder girdle (Tanner, 1952: 596).

The mesial fibers of the rectus cervicis super-
ficialis contract to stabilize the second basi-
branchial, thereby providing a stable attach-
ment for the insertion of the geniohyoideus. The
lateral fibers assist the rectus cervicis profundus
in retracting the tongue.

Rectus Cervicis Profundus

This muscle is actually a direct forward con-
tinuation of the rectus abdoniinus muscle which
originates on the puboischium. Tlie portion un-
der consideration in this paper begins at the level
of the sternum and continues forward in a posi-
tion dorsolateral to the rectus cervicis superfi-
cialis. It lies alongside the pericardium and lat-
eral to the second basibranchial. It then be-
comes thicker and narrower and passes through
the space between the ceratobranchial cartilages.
In all of the animals included in this paper, it
proceeds for^vard to attach onto the posterior
edge of the otoglossal cartilage. Since Dicampto-
don does not have an annular otoglossal carti-
lage, this muscle is not attached in the same
manner as in the other ambystomids. In this par-
ticular case, the forward portion of the rectus cer-
vicis profundus attaches to the posterodorsal
surface of the otoglossal plate, the latter forming
a continuous structure with the second radial
cartilages.

Myological studies of the Salamandridae
( Francis, 1934; Ozeti and Wake, 1969) indicate
that the rectus cervicis profundus lies ventral to
the hyoglossus muscle, but it is definitely dorsal
to the hvoglossus in the Ambystomidae. In most
specimens examined, the rectus cervicis muscle
inserts on the otoglossal at a point that is an-
terior as well as dorsal to the hyoglossus.

When this muscle contracts, the entire hyo-

l^ranchial apparatus is pulled posteriorly as in the
retraction of the tongue. It may also function in
flexion of the head.

Hebosteoypsiloideus

This muscle is a narrow band that takes its
origin on the dorsal surface of one of the myo-
septa of the rectus cervicis profundus. From its
origin, it proceeds anteriorly dorsal to the rectus
muscles. It turns mediad at the level of the sec-
ond basibranchial to insert on its dorsal surface.
The function is to assist the rectus cervicis pro-
fundus in stabilizing and in retracting the second
basibranchial.

Hyoglossus

This tiny muscle originates on the dorsal sur-
face of the anterior end of the first basibranchial
cartilage. There may be a few of the anterior-
most fibers that also attach to the aponeurosis
lingualis. The fibers are directed caudolaterally
to insert on the lateral edge of the second radial
cartilage. In some specimens there may be a few
fibers which continue laterally for a short dis-
tance into the substance of the tongue.

Tliese latter fibers may be homologous with
the muscle referred to by Driiner ( 1901 ) as the
basiradialis. In the ambystomids, however, we
were unable to detect a separation along the
course of the hyoglossus in any of the specimens
observed.

As indicated above, this muscle lies ventral to
the rectus cervicis profundus. This is in sharp
contrast to the report by Francis (1934) for
Salanumdra sakiiruindra in which he states that
the rectus cervicis profundus lies ventral to the
hyoglossus and that the former must be removed
in order to expose the latter.

The contraction of the hyoglossus causes the
second radial cartilage to swing in an antero-
dorsal direction, thereby elevating and protrud-
ing the pad of the tongue.

SPECIES DESCRIPTIONS

Pacific Giant Salamander
Dicamptodon ensatus (Eschscholtz)

Triton eii.satus Eschscholtz, Zool. Atlas, Pt. 5,
1833, p. 6, pi. 22. Dicamptodon, Strauch, Mem.
Acad. Sci. St. Petersb. Ser. 7, Vol. 16, 1870.

The range of this .species is along the humid
coastal regions of the Northwest. It extends from
Santa Cruz County in California, north to the
southern coastal region of British Columbia, and

into northwestern Idaho and the Rocky Moun-
tains of Montana. The ground color of the skin is
brown, mottled or marbled with darker brown or
black. This is the largest species of the Am-
bystomidae, transfonned adults attaining oc-
casionally a length of 300 mm. The body is stout,
and the limbs are large and strong. The head is
proportionately not as broad in this salamander
as in many of the other ambystomids. There are
12 costal grooves, but they are poorly defined.

28

BiiiCHAM VouNC Uniseksitv Science Bulletin

Vomerine teeth form a transverse series behind
the internal nares and are interrupted by a nar-
row break at the midhne where the series bends
anteriorly. The teeth extend to the lateral mar-
gin of the internal nares.

Hyobranchial Skeleton (Fig. 1)

The first basibranchial is not separated from
the radials by definite lines of articulation. It
is truncate in shape and uniformly, though
slightly, mineralized. In fact, the hyobrancfiium
as a whole appears to be of a consistency inter-
mediate between bone and cartilage. The first
basibranchial is located in the center of the area
circumscribed by the mandible.

This cartilage is shorter in relation to the
total length of the hyobranchial skeleton than in
any of the other ambystomids (Table 4). This
is probably because the radials are especially
long in this species. As seen in Table 5, the ratio
of the first basibranchial length to first cerato-
branchial length in Dicamptodon is the shortest
of all the species being considered, with the ex-
ception of A. annulatuin, A. cingulatttm, and
Rhyacotriton.

Table 4. Average ratios of length of first basibranchial
to total length of hyobranchiiim.

Species 0

1

2

3

4

5

Dicamptodon

I'tlSUtUS

Hhifucotriton

oUfinpicus
Atuhijstotna

(innulaturn
Ainhi/stoiua

cinguhitum
Amhijstojua

gracile
AiiihtfstoTnu

jeffcTsoniaiiuni
Amhystoma

maheei
Amhystoma

Ttiacrodacttfhtm
Amhystoma

macuhituin
Am})ystoma

talpoidcum
Amhystoma

opacum
Andiystoma

tcxamwi
Amhystoma

rosaceum
Amhystoma

tigrimim
Amhystoma tiorimtm

meUmostictam
Amhystoma tigrmum

nchuhsuvi

The first radial cartilages are long, curved,
and jointed a little beyond their midpoint. Their
distal ends articulate with the ceratohyals. They
contain no distinct centers of ossification. The
ratio of length of the first radial to the length
of the first basibranchial is much greater in this
species than in the other members of this family
which we have studied (Table 6).

The second radials join the first basibranchial
at their bases without a visible line of articula-
tion, as mentioned above. These structures also
unite with the otoglossal plate without any visi-
ble joint line. The otoglossal plate extends trans-
versely between the two second radials and is
tilted such that its anterior edge is more dorsal
than its posterior edge. These structures appear
to be composed of heavily calcified cartilage. In
cutting into this tissue, one is impressed with the
fact that it is denser than ordinary cartilage but
less dense than bone.

Tlie first ceratobranchial contains no distinct
center of ossification and articulates, as in all of
the species being studied in this paper, with the
first basibranchial, the second ceratobranchial,
and the epibranchial. As mentioned above, the
first ceratobranchial is larger in relation to the
first basibranchial than in all of the other species
except A. annulatuin, A. cingulatum, and Wiy-
acotriton.

No definite centers of ossification are found
in the second ceratobranchials. The distal ends
of these structures articulate with the first bran-
chial arch at a point posterior to the articulation
between the first ceratobranchial and the epi-
branchial. The ratio of the second ceratobran-
chial length to first ceratobranchial length is
equal to that seen in Rhyacotriton and smaller
than in the other ambystomids (Table 8).

A large center of ossification exists in the dis-
tal ends of the ceratohyals and is the largest per-
centage of ossification for this cartilage seen in
the family (Table 1). The expanded anterior end
is mostly cartilage, with a very slight amount of
mineralization imifomilv present throughout it.

The lateral arms of the second basibranchial
are much larger than the anterior ann. No dis-
tinct center of ossification exists in this element.

The ratio of mandible length to width is
greatest in this species (Table 9).

MUSCULATXJBE (Fig. 1)

Intermandihularis posterior originates along
the dorsomedial edge of approximately the an-
terior three-cjuarters of the mandible. After ex-

Biological Series. Vol. 16, No. 1

HYOBn.\NCHiuM .\Nn Tniio.\T Myology

29

tending in ;i transverse direction across the
throat, the fibers insert into the linea alba. The
posterior portion of this muscle overlaps the
interossatjuadrata ventrally. The ratio of the an-
teroposterior length of this muscle to the dis-
tance between the mental symphysis and the
pectoralis muscle is greater in this species than
in any of the others (Table 10).

Interhijoideus posterior originates on the
ventrolateral surface of the quadrate, the lateral
surface of the tip of the mandible, and the dis-
tal end of the scjuamosal. The fibers run trans-
versely and slightly posteriorly to insert into the
midventral line of fascia and into the skin and
fascia of the gular fold. The anterior fibers form
an almost indistinguishable margin with the pos-
terior fibers of the interossaquadrata. The pos-
terior edge of this muscle (the gular fold) is
more posteriorlv situated in this species than in
other species (Table 11).

Depressor mandibiihe originates in this spe-
cies along the dorsal aspect of the squamosal
and on the posteroventral surface of the otic
capsule. It also attaches to the fascia cephalo-
dorsalis. The fibers converge ventrally onto the
posterior tip of the mandible.

Levator nwndihuiae is basically the same as
that given above in the general statement of
throat musculature of the Ambystomidae. ex-
cept that the levator mandibulae posterior is not
covered largely bv the levator mandibulae ex-
temus. The majoritv of the latter muscle lies pos-
terior to the levator mandibulae posterior. The
tendon of insertion is tougher and larger than
that seen in the other species. The overall size of
the muscle, in fact, is hypertrophied as compared
to other ambvstomids, with perhaps the excep-
tion of Rhyacotriton, in which this particular
muscle also appears to be enlarged.

Interossaquadrata originates on the medial
aspect of the posterior tip of the mandible and
the ventral surface of the quadrate immediately
posterior to the mandibuloquadrate articulation.
It inserts into the linea alba as the latter contin-
ues caudad from the intermandibularis posterior.
Suhhijoidcus originates on the posterior tip of
the ceratohval and wraps around the latter be-
fore proceeding forward parallel to the subar-
cualis rectus I. This is a thicker muscle than in
the other species included in this investigation.
It inserts into the dorsal fascia of the interman-
dibularis posterior. The interossaquadrata blends
in with this muscle and overlaps it ventrally
along the posterior one-third of its length.

Geniohijoideus takes its origin on the poste-
rior edge of the mandible extending from the

Tiible 5. Average ratios of length of first basifirancfiial
to length of first ceratobranchial.

Species

)

1

2

3

4

5 .6 .7

Dicamptodon

I'Tisattis
Rhyacotriton

nlympicus
Ambystoma

uutiulatum
Ambystoma

cingultittim
Ambystoma

gracilv
Ambystoma

jeffersoniatium
Am!)ystoma

1
1

iTwbeei
Ambystoma

macrodactylum
Ambystoma

maculatum
Ambystoma

talpoideum
Ambystoma

opacum
Ambystoma

texamim
Ambystoma

rosaceum
Aminjstoma

tigriuum
Ambystoma tigrinum

meliitwstictuTu
Ambystoma tigrinum

ncbtilosi^tm

1

symphysis laterally for a distance of about 3 mm.
The insertion is onto the anterior edge of the
second basibranchial and laterally into the ten-
donous inscription that separates the geniohyoid-
eus from the rectus cervicis superficialis.

Genioslossus is divided into ventromedial
and dorsolateral portions as in the other species
of this familv. However, it appears to be a stout-
er muscle than in the others. It originates along
the posterior edge of the mandible dorsal to the
origin of the geniohyoideus. The medial fibers
attach to the aponeurosis lingualis, and then pro-
ceed caudad to insert onto the anteroventral sur-
face of the otoglossal plate. The lateral fibers
overlap the tip of the second radial cartilage
ventrallv, and then insert into the adjacent fas-
cia.

Srilyarrualis rectus I wraps around the poste-
rior end of the epibranchial and forms a
sheath which encloses the epibranchial. As the
fibers pass for\vard, thev follow the ceratohyal
and spread out over the hitter's expanded ante-
rior end. The insertion is along the anterior edge
of the ceratohyal.

Rectus cervicis superficialis originates on the

30

Bricham Vounc University Science Bulletin

Table 6. Average ratios of length of first radial tii
length of first basibranuhial.

Species 0 0.5 1.0 1.5 2.

0

Dicamptodou

t'lisatus
Hhyacotriton

ohfmpicus
Amltystomu

unnulatum
ATrd>ystom(i

cingulatxtm
Amhystoma

gracilc
Amhystoma

jeffcrsoniamun
Amhystoma

nwheei
Amhysti>Tuu

mucrodactyhm
At7ihystoma

maculattim
Amhystomu

talpoideum
Atnhystoma

opacum

Amhystoma

texanum
Amhystoma

rosaccum
Am}}ystoma

Amhystoma tighmim
meUitiostictum

Am})ystoma tignnum
iichulosiim

dorsal surface of the sternum and extends for-
ward dorsal to the sternum, dorsal to the cora-
coid cartilages, and ventral to the pericardium.
It attaches to the dorsal surface of the second
basibranchial and laterally to the tendonous in-
scription from which the geniohyoideus arises.
The superficial fibers of the portion which ex-
tends forward from this point insert into the
posterior edge of the first ceratobranchial lateral
to, but not inserting onto, the posterior tip of
the first basibranchial. The deeper fibers con-
tinue forward to insert onto the posterior sur-
face of the otoglossal plate ventral to the inser-
tion of the rectus cervicis profundus.

Rectus cervicis profundus represents a direct
anterior continuation of the rectus abdominus.
From the level of the sternum, it passes forward
along the dorsolateral edge of the rectus cervicis
superficialis. It passes ventrally over the second
ceratobranchial and dorsally over the first cera-
tobranchial. It inserts onto the posterodorsal sur-
face of the otoglossal plate dorsal to the attach-
ment of the rectiis cervicis superficialis.

Hehosteoifpsiloideu.s originates from the dor-
sal surface of the second myoseptum posterior

to the second basibranchial. The fibers extend
forward dorsal to the rectus cervicis profundus
and then medially to insert onto the dorsal sur-
face of the second basibranchial. The distal fi-
bers in this species are strongly intermingled
with those of the rectus cervicis profundus.

Ht/oglossiis. We have been unable to find a
hyoglossus muscle in this species. The muscle
which most nearly occupies the position of the
hyoglossus is the posterior portion of the genio-
glossus, whose fibers extend in an anteroposte-
rior direction for the most part. The lateral fibers
of the genioglossus, however, do extend in a
posterolateral direction as is seen in the hyoglos-
sus of other salamanders.

Olympic Salamander.

Rhyacotriton ohjmpicus (Gaige)

Ranodon ohjmpicus Gaige, Occ. Papers Mus.
Zool. Univ. Mich., No. 40, May 30, 1917, p. 2
pi. 1.

Rhyacotriton Dunn, Proc. N. Engl. Zool. Club,
7:56, 1920.

The Olympic Salamander is found in western
Oregon and Washington. It has been collected in
small springs and streams and among stones
along the banks. This is a small ambystomid,
occasionally attaining the size of 130 mm. It is
characterized by its large, protuberant eyes, the
horizontal diameter of which is equal to or
greater than the length of the snout. The ground
color is seal-brown with small white flecks dis-
tributed over the skin. The belly is yellow or
orange in color. The tail is short and the gular
fold is strongly developed. Costal grooves num-
ber 14 when the one in the axilla and the one in
the groin are included. The vomerine teeth are
formed into two strongly arched series that are
separated at the midline. In the male, these se-
ries consist of fewer and larger teeth than in the
female, and they extend in the male to a some-
what more lateral position in relation to the in-
ternal nares than in the female. The shape of
the vent Ls different in the two sexes, the male
being fonned into squarish lobes which can
readily be seen from either a dorsal or a ventral
view, whereas in the female these lobes are con-
siderably reduced in size.

Hyobranchial Skeleton (Fig. 2)

There is no ossification in the various com-
ponents of the hyobranchium in this species
(Table 1).

The first basibranchial is pear-shaped, but
narrow. The width-to-length ratio is smaller, in

RtoLooiCAi. Sebies, Vol. 16, No. 1

HYObH.-\NCHlUM AND ThHOAT MvOLOCV

31

fact, than in any of the other ambystomids in-
cluded in this study (Table 15). The distance
posterior from the mental symphysis to the an-
terior edt^e of this structure in relation to the
mandible length is exceeded only by A. annula-
tum (Table 12). The length of the first basi-
branchial is also small when compared to the
total length of the first branchial arch (first
ceratobranchial plus epibranchial) (Table 20).

The first radials attach to the anterior tips
of the ceratohyals by a short ligament. These
radials are delicate, cartilaginous rods project-
ing anterolaterad from the pointed anterior tip
of the first basibranchial. The shortness of the
first basibranchial is reflected in the high ratio
of radial length to length of first basibranchial.
This ratio is exceeded only by Dicamptodon
(Table 6).

The second radials are approximately the
same length as the first radials, but the former
are curved dorsally to articulate with the oto-
glossal.

The first ceratobranchial is not uniformly
wide along its length, being narrower at the
anterior end and then becoming wider about
midway along its length. It is flattened dorso-
ventrally.

The moderately long epibranchial forms a
smooth continuation of the first ceratobranchial
and is also flattened dorsoventrally. The second
ceratobranchial is a narrow, smoothly curved rod
of cartilage that forms a link between the pos-
terior tip of the first basibranchial and the an-
terior end of the epibranchial. In relation to the
first ceratobranchial, the second ceratobranchial
is short when compared with the other species
(Table 8).

The hyobranchial skeleton is shortened in an
anteroposterior direction in this .species. This is
evident in the high ratio of ceratohyal length to
total length of hyobranchium (Table 19).

The second basibranchial cartilage has a
verv small anterior projection. The ratio of an-
teroposterior length to side-to-side length of this
element is smaller than in other species (Table
18).

Musculature (Fig. 2)

Intermandibularis posterior originates on
the dorsomedial surface of the mandible. The fi-
bers lie transversely for the most part, except for
the posterior ones which are directed slightly
caudad. The anterior portion appears almost as
a separate muscle because it is a narrow band
of continuous fibers extending across from one
side to the other. Posterior to this band of mus-
cle the linea alba, into which the remainder of

the fibers insert, is broad at first and then tapers
to a point at the level of the posterior edge of
this muscle. The geniohyoideus can be clearly
seen through the broad portion of the linea alba.

Intcrhijoklcxis posterior is . a thin ribbon-
shaped muscle. The origin is broader than that
seen in other species. The anterior fibers arise
from the posterior tip of the mandible, while
the remainder of the muscle originates from the
fascia cephalodorsalis, which covers the lateral
aspect of the depressor mandibulae, and from
the lateral surface of the quadrate and the squa-
mosal. It inserts into the gular fold and the mid-
ventral fascia. It appears to be a weaker mus-
cle than the interhyoideus posterior of other
species of the family. It especially appears small
in contrast to the hypertrophied levator mandi-
bulae of this species.

Depressor mandibulae originates from the
fascia cephalodorsalis (posterior fibers), the up-
per posterolateral surface of the squamosal and
from the posterolateral surface of the otic cap-
sule. It inserts onto the posterior tip of the man-
dible.

Table 7. Average ratios of length of first cerato-
branchial to total length of first branchial arch.

Species

.5 .6 .7

Dicamptodon

cnsatus
Rhijacotriton

olympicus
Anihijstoma

antiulafum
AjJibifstoma

cingulatum
Arnbijstoma

f^racilc
Amhystomu

jeffcrsonianum
Amhystoma

maheei
Aml)ystotna

macrodactylum
Amhystoma

maculatum
Ajnhystoma

talpoideum
Amhijstoma

opacum
Amhystoma

tcxanum
Amhystoma

Tosaceuni
Amhystoma

tigrinum
Amhystoma tigrinum

melanofitictum
Amhystoma tigrinum

nchulosum

32

BiuGiiAM V()UN<; IIni\ek.sity .Scienc:e Bulletin

Levator mandihulae has its origin and inser-
tions essentially the same as that given in the
section on general musculature of the family.
The size, however, is a departure from the con-
dition seen in the other species. This muscle is
hypertrophied as noted by the fact that it can
be seen as a bulge behind the eye when viewed
from the ventral surface. It can also be seen in
Dicamptodon from a ventral view, but not to the
same extent in the other species included in this
study. The depressor mandibulae, which opens
the mouth, is not enlarged in RJitjacotriton and
perhaps closing the jaw with a powerful stroke
is relatively more important in this species than
opening the mouth, whereas this may not be the
case in other species.

Interossaquadrata originates from the poste-
rior surface of the (juadrate, and ventral sur-
face of the articulation between the ceratohyal
and the (juadrate. The middle fibers are at-
tached, in addition, to the posterior tip of the
mandible, and are hypertrophied to form a band
of muscle extending directly mesiad. This mus-
cle fans out over the venter of the throat and in-
serts into the fascia of the linea alba. The fibers
from each side do not meet at the midline. It ap-
pears to be better developed and more easily
separated from the subhyoideus than in other
species of Ambystomidae.

Subhijoideiis originates along the lateral mar-
gin of the ceratohyal beginning at a point about
1.0 mm from the posterior tip of this cartilage
and extending forward about thrce-cjuarters of
the distance to the anterior tip. It is a heavy
muscle which is about as thick as the geniohyoi-
deus, lying lateral and nearly parallel with the
latter. The insertion is at a point on the posterior
surface of the mandible immediately lateral to
the origin of the geniohyoideus. This appears to
be better developed than the subhyoideus in
other species of the family.

Geniolnjoideus is attached to the posterior
surface of the mandible along a line which be-
gins at the symphysis and extends laterally about
1.0 mm. The lateral fibers proceed directly cau-
dad while the medial fibers are directed shghtly
mediad before turning in a caudal direction.
This is a strong band of muscle, one on each side
of the midline and overlapped ventrally by the
intemiandibularis posterior. It inserts onto the
second hasibranchial and in the tendonous in-
scription which separates this muscle from the
rectus cervicis superficialis.

Genio<iIossus has two portions, a ventral part
whicfi is much the same as in other species, and
a dorsal portion. Both of these appear to be
slightly longer than in other species.

The ventral portion originates on the ventral
and posterior surfaces of the mandible, begin-
ning about 0.5 mm lateral to the symphysis and
extending along this line of origin on the mandi-
ble for about 1.0 mm. The fibers converge to in-
sert along the lateral edge of the ventral sur-
face of the anterior end of the first hasibran-
chial. It also sends a few fibers onto the otoglos-
sal. This muscle is longer in this species than in
others.

The dorsal portion originates from the same
position as the ventral portion just described,
except that it is immediately dorsal to the origin
of the ventral portion. The lateral fibers pass
caudad and slightly laterally to insert into the
lateral part of the floor of the mouth dorsal to
the subhyoideus belly. The medial fibers insert
onto the ventral surface of the first hasibranchial
and the anterior edge of the first ceratobran-
chial. This muscle overlaps the radial cartilages
ventrally.

Suharcualis rectus I attaches at its origin to
the dorsal side of the posterior tip of the epi-
branchial, wraps around it, and encloses it in a
sheath. The fibers continue obliquely in an an-
teromesial direction and fan out over the ex-
panded anterior end of the ceratohyal. The in-
sertion of these fibers is along the anterior mar-
gin of the ceratohyal. The overall position of
this muscle appears to be more posterior than in
the other species, the anterior tip not being very
far anterior to the posterior tip of the mandible.
In addition, these two muscles lie in a position
in which the anterior ends tend to point strongly
toward each other.

Rectus cervicli superficialis originates me-
dially from the dorsal surface of the sternum
and laterally along the tendonous inscription
separating this miLscle from tlie rectus abdom-
iiuis. There are two transverse inscriptions along
its length between the sternum and the second
hasibranchial. A small section of the pericardium
is visible between these two muscles at the mid-
line. The fibers that extend forward from the
second basil^ranchial insert along the posterior
edge of the medial end of the first ceratobran-
chial and onto the posterior tip of the first hasi-
branchial. The sternum in this species is reduced
to a small triangle with one angle facing anteri-
orly.

Rectus cervicis profundus originates techni-
cally on the puboischium. It is a broad sheet of
muscle extending along the ventral body wall
and continuing forward at the level of the ster-
num as a flat band of muscle that is overlapped
partially on its medioventral surface by the rec-
tus cervicis superficialis. It tunis mediad and

Bioi.oGK'Ai, Series, Vol. 16, No. 1

Hyohranchium and Tumoat Myology

33

passes through the arch formed by the two cera-
tobranchials, and then forward dorsal to the first
basibranchial to attach to the posterior edge of
the otoglossal.

Hebosteoypsiioidcus originates from the dor-
sal surface of the rectus cervicis muscles at the
most anterior tendonous inscription of those in-
scriptions occurring between the sternum and
the second basibranchial. It is thin and narrow
and extends forward dorsal to the rectus cervi-
cis profundus for a short distance, where it turns
mediad across the rectus cervicis superficialis to-
ward the second basibranchial, upon whose dor-
sal surface it inserts.

Hijoglos.ms originates on the dorsal surface
of the anterior end of the first basibranchial and
along the lateral margin of the dorsal surface of
this cartilage. This small muscle is situated dor-
sal to the first basibranchial and ventral to the
insertion end of the rectus cervicis profundus.
The fibers extend laterally to insert onto the tip
of the second radial and into the fascia at the
sides of the base of the tongue.

Ringed Salamander.
Ambystonm annulatum Cope

Amblystoma annulatum Cope, Proc. Amer.
Philos. Soc, Vol. 23, 1886, p. 525.

This is a strikingly marked salamander with
yellow spots and cross bars on a dark brown to
black ground color. The cross bars on the trunk
extend ventrad to the level of the legs, whereas
those on the tail reach the ventral edge. The
head is small and the tail long and slender. To-
tal length rarely exceeds 203 mm. There are 15
costal grooves, counting one each in the axilla
and groin. The vomerine teeth consist of two
short series that lie obliquely forward between
the internal nares to fonn an inverted V. Each
consists of three rows of 7 to 11 small, blunt
teeth. This species is abundant in parts of Ar-
kansas and Missouri and appears to be a bur-
rower, spending most of the year beneath the
surface of the ground..

Hyobranchial Skeleton ( Fig. 3 )

\ definite center of ossification exists in the
first basibranchial. This center is located in an
anteromedial position, and constitutes a greater
percentage of the total volume of the first basi-
branchial than in any of the other species ( Table
1). There are definite lines of articulation be-
t^veen this cartilage and the first radials. These
lines do not exist, however, between the first

Table 8. Average ratios of length of second cerato-
hranchial to length of first ceratobranchial.

Species

0 0.2 0.4 0.6 0.8 1.0 1

2

Dicamptoilon

ni.satus
Hhijaiotriton

ohjmfmu.s
Ainhifstiwia

anmilntum
Atnhifstoma

ciiif^uUitum
Amin/stoinii

gmcilc

jcffcr.soniamim
ArithijstoTua

mubeei
Amhystomu

iiKiiroihicttjlum
Anihy.stonia

tnaculiitum
Anihi/stoma

talpoidcuin
Avihtj.stonia

Aiuhi/.'ttoinii
texunum

Arnhysionui
rosaccum

AinhifstiniKi

Atn}>ystomu tigrinum

Anihystoina (igrhnim

basibranchial and the second radials. The first
basibranchial is shaped like a slightly elongated
shield. This structure is situated in a more pos-
terior position in relation to the position of the
mandible than in other species (Table 12).

There is no ossification in the first radial car-
tilages. These structures are thin and taper to a
fine point. They do not attach to the anterior
end of the ceratohyal and they are not segment-
ed.

The second radials also are entirely carti-
laginous. They articulate with the otoglossal
cartilage at their distal tips. The otoglossal is a
thin bar of cartilage fomiing an arch which is
directed dorsally into the tongue.

The first ceratobranchial is devoid of bone.
In relation to the length of the first basibran-
chial, this structure is exceeded in length only
in RJufacotriton and A. cin<ntlatum (Table 5).

The epibranchial contains an ossified tip con-
stituting about 12 percent of its total volume
(Table 1), the remainder being cartilage. The
first ceratobranchial is long in relation to the
epibranchial, this ratio being exceeded only in
A. cinguhitum (Table 13). The articulation be-

34

Brigham Young Univehsitv Science Bulletin

tween the epibranchial and the first cerato-
branchial is almost indistinguishable.

The second ceratobranchial is longer in rela-
tion to the length of the first ceratobranchial in
this species than in the others (Table 8) and
contains no bone.

The ceratohyal shows no ossification. The
width of this structure in relation to its length is
exceeded only in Dicamptodon and A. texanutn
(Table 14). '

The second basibranchial has a shape not
seen in most otlier ambystomids. A small, posteri-
orly directed ami exists in addition to the other
three, and the lateral arms are slightly expanded
at their tips. More than half of the structure is
cartilage, the lateral anns being almost entirely
chondrified (Table 1).

The ratio of mandible length to width is ex-
ceeded only liy Dicamptodon and A. mabeei, al-
though it is equalled by A. nmcidattim and A.
cingulatum (Table 9).

Musculature

InternuimUhidaris posterior originates along
the dorsomedial surface of the mandible. The fi-

Table 9. Average ratios of mandible length to width.

Species

0 0

2 0.4 0.6 0.8 1.0 1

o

Dicamptodon

ensatus
Hhifacotritoii

oh/mpicus
Atnhy.stoiJUi

annidatuTti
Arnhifstoma

t higiilatiitn
Amhy.'ytoma

gracilc
Amhystoma

jeffcrsontontiiit
Anihif.stiiinn

rtuihcei
AmhystoUKi

macrodactylum
Amhystotna

nuiculatuttt
Atid>ii\l(>mu

ta{poideu7)t
Ai}d}y\tiniui

opacum
Amhystoma

texumwt

Anthystotna

wsaccum

Amhyatonui

tigrinutn

Amhystouui ti^ritutm

melunostutujii
Amhystoma tifi^rinum
nehulosttm

1

bers lie transversely and insert into the broad
linea alba at the midline of the throat. The linea
alba is wider than in the other species studied.
The gcnioglossus is visible at the point of the
chin, as in most other species.

Intcrhtfoidctis posterior originates from the
fascia which is attached to the lateral edge of
the posterior tip of the mandible, the quadrate,
the squamosal, and the otic capsule. The poste-
rior fibers arise in the fascia cephalodorsalis. The
insertion is into the skin and fascia of the gular
fold and into the linea alba at the midline. The
anterior fibers blend in with the posterior fibers
of the intcrossa(|uadrata.

Depressor nuindibidae has its origin along
the dorsal end of the squamosal, the ventral and
posterior surfaces of the otic capsule, and the
fascia cephalodorsalis. The fibers converge into
a short, stout tendon which inserts onto the pos-
terior tip of the mandible.

Levator vuindibulae corresponds very closely
to the description given in the section on general
musculature.

Interossaquadrata originates from the ventral
surface of the hyocjuadrate ligament and from
the quadrate bone immediately anterior to this
ligament. The fibers fan out over the ventral sur-
face of the throat, with the anterior fibers ex-
tending anteromedially over the ventral surface
of the subarcualis rectus I, and the posterior fi-
bers extending posteromedially to parallel the
fibers of the interhyoideus posterior. The inser-
tion is into the linea alba.

Stibhyoideus originates on the ventrolateral
margin of the posterior tip of the ceratohyal. It
is thin and delicate, especially at its insertion.
The posterior fibers are difficult to distinguish
from those of the interossaquadrata. It lies over
the ventral surface of the subarcualis rectus I
and inserts into the dorsal surface of the fascia
of the intermandibularis posterior.

Genioliyoidcus originates along a line on the
posterior edge of the mandible beginning at a
point just lateral to the symphysis and extending
laterallv about 1.5 to 2.0 mm. This end is over-
lapped ventrally by the ventral portion of the
gcnioglossus. The fibers extend directly caudad
to insert on the second basibranchial cartilage.

Geniof^lossiis originates along a Une on the
ventral surface of the mandible beginning at the
symphysis and extending laterally over the ven-
tral surface of the origin of the geniohyoideus.
The fibers converge medially to form a strong
band which contributes to the aponeurosis lin-
gualis. Some of these fibers, in turn, continue
distally to attach to the anterior end of the first
basibranchial and to the otoglossal. Contraction

BioLOGicAi. Series, \'ol. 16, No. 1

Hyobranchium and TunoAT Myology

35

Table 10, Average ratios of distance between mental
symphysis and posterior edge of intermandibularis
posterior to distance between mental symphysis and
pectoralis.

Species

0

,1

Dicamptodon

ensattm
Hlufiicotriton

otympicus
Amhystoina

cintwlatum
Amhystoma

cinguldtitm
Amhystomu

gracilc
Amhystoma

Jefferson tatium
Amhystoma

mabeei
Amhystoma

i

macrodactylum
Avihystoma

maculatum
Amhystormi

talpoideuTti
Amhystoma

opacum
Amhystoma

texanum
Amhystoma

rusacettni
Amhystoma

tigrinum
Amhustoma tigrinum

melarwstictum
Ambustoma tignnum

nebuhsitm

of this muscle therefore serves to advance both
of these cartilages. The ventral portion of the
genioglossus is large in this species, as compared
to the same muscle in most of the other species
investigated,

Tlie dorsal portion originates along the man-
dible just dorsal to the origin of the ventral por-
tion. This muscle spreads out in a fan shape over
the floor of the mouth. The medial fibers extend
across the dorsal surface of the tips of the first
radials and continue distally to insert along the
posterolateral edge of the second radials. The
lateral fibers insert into the fascia of the floor of
the mouth.

Subarcualis rectus I originates along the dor-
sal edge of the tip of the epibranchial. The fi-
bers radiate around this cartilage and enclose it
in a sheath. They then continue forward in a
inedialantcrior direction. When they reach the
junction between the epibranchial and the sec-
ond ceratobranchial, they lose their sheath shape
and come to lie in a ventral position over the
first ceratobranchial and the ceratohyal. Anteri-
orly, they spread out over the expanded an-

terior end of the ceratohyal and insert along the
latters anterior edge.

Rectus cervicis superfickilis originates from
the dorsal surface of the sternum. The medial fi-
bers attach to the second basibranchial and the
lateral fibers form a thicker, narrower band
which runs forward to insert along the posterior
edge of the medial end of the first ceratobran-
chial. There are three transverse tendonous in-
scriptions in this muscle between the sternum
and the second basibranchial.

Rectus cervicis profundus is actually an an-
terior continuation of the rectus abdominus. At
the level of the sternum, this muscle extends an-
teriorly alongside of, and dorsal to, the rectus
cervicis superficialis. Near its middle, the hebo-
steoypsiloideus can be seen along the border be-
tween the two rectus muscles. The rectus cervi-
cis profundus becomes tendonous as it passes
through the opening formed by the two cerato-
branchials. This thin tendon then continues for-
ward, dorsal to the first basibranchial, to attach
to the medial section of the otoglossal. This ten-
don was seen only in this species and in A. cin-
gulatum. The other species have fleshy inser-
tions of this muscle on the otoglossal.

Hebosteoypsiloideus originates as a nar-
row band of muscle from the dorsal surface
of the third inscription of the rectus cervicis pro-
fundus posterior to the second basibranchial. Its
origin can be seen between the two rectus mus-
cles. From here it extends forward dorsal to the
medial edge of the rectus cervicis profundus and
then turns medially to insert onto the dorsal sur-
face of the central portion and anterior arm of
the second basibranchial.

Hyoglossus originates on the dorsal surface
of the anterior end of the first basibranchial and
the adjacent fascia. Its fibers extend laterally and
slightly posteriorly to attach along the posterior
margin of the tip of the second radial. This is a
small muscle.

Reticulated Salamander.

Ambtjstoina cingulatum Cope

Amhh/stonui cingidatum Cope, Proc. Ac. Nat.
Sci, Phil,, 1867, p. 205.

This is a medium-sized species, less than 102
mm in total length. The ground color is black
and has many small, gray spots which form a
reticulated pattern on the dorsal surfaces of the
head, trunk, and tail. Along the sides, these spots
are concentrated into narrow vertical lines be-
tween the 13-14 costal grooves. Vomerine teeth
are found in tvvo short series, one on either side
of the midline, and each contains about seven

36

BiuGiiAM VoUNc University Science Bulletin

teeth in two or three rows. This species is found
from South Carolina to nortliern Florida and
Alabama. Little is known about its habitat pref-
erence.

Hyobranchial Skeleton (Fig. 4)

The average ossification of the first basibran-
chial in the five specimens studied was 73 per-
cent (Table 1 ). This ossification is centrally lo-
cated, with a narrow margin of cartilage existing
around the structure. No definite articulation
lines were visible between the radials and the
first basibranchial. This shield-shaped structure
is located in a posterior position as compared to
most of the other species. Only in Rlit/ucotriton
and A. anmdatuin is the first basibranchial in a
more posterior position (Tabic 12). It is shorter
in relation to the total length of the hyobran-
chium than in all of the other species except in
Dicamptodon and is wider in relation to its
length than in other species (Tables 4 and 15).

The first radial cartilages are short, earlike
projections without ossification. They are greatly
reduced as compared to the other species ( Table

Table 11. Average ratios of distance between mental
symphysis and giilar fold to distance between
mental symphysis and pectoralis.

Species

0 .1 .2 ..3

.8

Dicamptodon

etu^utus
Rhijacotriton

ohfmpicus
Anihystotiw

(intutlatinn
Aniliystottui

cin^uliituni
Ambystuma

gracile
Ambttstoma

jejjt'Tsonianum
Ambystoma

mabeei

Ambystoma

macwilactyhim
Ambystoma

muculatum
Ambystoma

talpoideum
Ambystoma

opacuni
Ambysti)ma

texanum

Amifystouia

rosacfum
Ambystoma

til^rinum
Ambystoma tig^rinum

jfwhinostictum
Ambystoma tij^rinum
nehttlostim

_

6). They do not articulate with the anterior end
of the ceratohyal.

The second radials are short and delicate and
wholly cartilaginous. Their tips articulate with
the otoglossal, the outline of which is pear-
shaped rather than the usual annular shape.

In one of the five specimens which was dis-
sected, the first ceratobranchial on each side
contained a small ossification center on its me-
dial end. The other four specimens were devoid
of bone in this structure. In relation to the total
length of the first branchial arch, the first cera-
tobranchial length is exceeded by no other spe-
cies (Table 7); and, when compared to the
length of the first basibranchial, it is only ex-
ceeded by Wiyacotriton (Table 5).

The joint between the first ceratobranchial
and the cpibranchial is difficult to locate, the
two structures appearing to be well fused. Two
of the five specimens contained no bone in the
cpibranchial, whereas the other three had an
average of 15 percent ossification, the center of
ossification in each case being located in the tip.
The fraction of the total length of the first bran-
chial arch that is contributed by the cpibran-
chial is smaller than in other species (Table 16).

The second ceratobranchial contains no carti-
lage. When this structure (which is actually the
second branchial arch) is compared in length to
the first branchial arch ( first ceratobranchial
plus cpibranchial), the ratio is found to be
greater in this species than in others (Table 17).

The ceratohyal contains a long, narrow center
of ossification running lengthwise at its posterior
end. The bone in this stnicturc amounts to an
average of approximately 5 percent of the total
volume. Two of the five specimens, however,
had no ossification in the ceratohyal.

The second basibranchial has a small fourth
ann projecting posteriorly. The cartilage portion
of this structure is located on the lateral arms
and is about 38 percent of the total volume. The
anteroposterior length of the second basibran-
chial is almost half as long as the side-to-side
length (Table IS).

The ratio of mandible length to mandible
widtli is high for this species, being etjualled by
A. nwculatum and A. annuhitttm and exceeded
only by Dicamptodon and A. iiud)cei (Table 9).

Mu.scuLA'ruHE (Fig. 4)

Internumddndaris posterior has the major
portion originating along the dorsomedial sur-
face of the mandible and inserting medially into
the linea alba. Tlie most anterior portion forms
a stout band extending from the ventromedial
surface of the mandible to the midline. The in-

Biological Series. Vol. 16, No. 1

Hyodh.vnciiium and Tiihoat Myology

37

sertion of this anterior portion is more medial
than the insertion of the remainder of the mus-
cle. The genioglossus and the geniohyoideus can
be seen passing dorsal to the forward edge of
the anterior portion.

Interhtjoidcus posterior originates from a fas-
cial sheet that is attached to the lateral aspect
of the mandible tip, and from the quadrate,
squamosal, and otic capsule. As in most of the
other species, the posterior fibers attach to the
fascia cephalodorsalLs. The fibers extend postero-
mediallv to form a prominent muscle whose in-
sertion is along the skin of the gular fold and the
midventral line of the throat. The anterior fibers
blend in with those of the interossaquadrata.

Depressor mandibulae arises from the fascia
cephalodorsalis, the dorsal and posterior surfaces
of the squamosal, and from the ventral and pos-
terior surfaces of the otic capsule. It inserts onto
the posterior tip of the mandible.

Levator immdihulae is essentially the same as
the description given earlier for the levator man-
dibulae in the general section on musculature.

Interossaquadrata originates on the lateral
surface of the quadrate just anterior to the hyo-
quadrate ligament, from the lateral surface of
the hyoquadrate ligament, and from the dorsal
and lateral edges of the tip of the ceratohyal.
The fibers wrap around the tip of the ceratohyal
completely covering its lateral surface. They
pass ventral to the subarcualis rectus I and fan
out over the ventral surface of the throat. No
noticeable boundary exists between the anterior
fibers and those of the subhyoideus. The inser-
tion is into the fascia along the midventral line
of the throat and the dorsal surface of the fascia
into which the intermandibularis posterior in-
serts.

Subhyoideus is a delicate muscle closely as-
sociated with the interossaquadrata. It originates
along the lateral edge of the posterior end of the
ceratohyal and extends anteromedially to insert
into the dorsal surface of the fascia of the linea
alba. The fibers at the origin lie dorsal to those
of the interossaquadrata, but both muscles
spread out over the throat; they form a broad
fan-shaped structure, the anterior one-fifth of
which is made up of fibers of the subhyoideus,
whereas the remaining posterior portion is the
interossaquadrata.

Geniohyoideus originates on the ventromedial
surface of the mandible along a line beginning
about 1.0 mm lateral to the symphysis and ex-
tending laterally about 1.5 mm. It is not over-
lapped ventrally by the genioglossus but is
crossed ventrally by the anterior band of the in-
termandibularis posterior. The medial fibers ex-

tend in a somewhat medial direction as they pro-
ceed caudad. The insertion is onto the second
basibranchial cartilage and into the tendonous
inscription which extends laterally from this
cartilage.

Genioglossus is delicate when compared to
the same structure in the other species investi-
gated. It originates on the ventromedial surface
of the mandible beginning at the symphysis and
extending about 1.0 mm laterally. The fibers ex-
tend caudad to attach to the aponeurosis lin-
gualis but continue on as a narrow band to in-
sert onto the anterior edge of the first basibran-
chial and onto the otoglossal.

The dorsal portion originates along the man-
dible from a position immediately dorsal to the
origin of the geniohyoideus. The fibers insert
along the base of the anterior margin and the
sides of the tongue. The medial fibers extend
directly caudad to insert onto the second radial
cartilage.

Subarcualis rectus I originates on the dorsal
surface of the posterior tip of the epibranchial
and then wraps around and encloses the epi-

Table 12. Average ratios of distance between mental
symphysis and anterior edge of first basibranchial
to mandible length.

Species 0 .1 .2 .3 .4 .5 .6 .7 .8 .9 1.0

Dicamptocion

ftifiatlis
Rhi/ucotriton

oltirupictts
Ajulnjstomu

amntlatum
Anthjfstojiui

cin<^ulutUJii
Amhtj.'iloma

Anihl/stointi

jcffcr:ioniiimiin
Aminjfttonici

maheci
Ainhti.-itoruu

nuuTodacttjhim
Anthij^tuTiici

muculatitm
Aininj^toina

talpoidcum
Aininjstoitut

OptlCUfU

Amhystoina

texanum
Ambystoma

rasaccurji
Amhi/slonia

tigrinuni
Amlnjstonia tigriutim

lUL'lttiwstictuvi
AnthijstoTiia tigrinuni

nvhulo:iunt

38

BniGH.\M Young UNrvER.siTY Science Bulletin

Table 13. Average ratio.s of length of first cerato-
branchial to length of epibranchial.

Species

0 0.5 1.0 1

5

Dicamptodon

ensatus

Rfujacotriton

ohjmpicus
Amhystuma

annulatum
Amhystoma

cinguUititni
Amhystoma

grati/c
Amhystoma

jeffcrsonunium
Amiyystomii

maheei
Amhystotiia

macrodactyhim
Ai7d>tfstoma

nuiculaturji
Amhystoma

tal)>oidi'um
Aruhystoma

opacum
Am})ystoma

texanum
Amhystoma

rosaccum
Amhystoma

tigrimim
Amfnjsiovia tigrmutn

melaiwstictum
Amhystoma tigrinum

nehulosum

branchial in a muscular .sheath. This sheath con-
tinues anteromedially to the level of the cerato-
hyal where it leaves the first ceratobranchial and
spreads out over the ventral surface of the cera-
tohyal. It inserts along the forward margin of
the expanded anterior end of the ceratohyal.

Rectus cervicis superjiciaJis originates on the
dorsal surface of the sternum and extends for-
ward ventral to the pericardium fonning a thin,
broad sheet. Between the .sternum and the sec-
ond basibranchial there are three transverse in-
scriptions. The portion which continues forward
from the second basibranchial is narrower and
thicker than the posterior portion. The anterior
portion attaches to the anterior edge of the lat-
eral tip of the second basibranchial and the ad-
jacent tendonous inscription, and the fibers pro-
ceed anteromediallv to insert along the posterior
edge of the medial end of the first ceratobran-
chial and along the lateral edge of the posterior
tip of the first basibranchial.

Rectus cervicis profundtis lies dorsolateral to
the rectus cervicis superlicialis, and its shape is
essentially the same as the latter. It is actually a
continuation of the rectus abdominus, and so the

origin is, technically, the puboischium. The por-
tion extending anteriorly from the second basi-
branchial is also narrower and thicker than the
posterior portion. This anterior portion passes be-
tween the ceratobranchials and dorsal to the
first basibranchial. At this point, the fibers con-
verge into a thin, narrow tendon which inserts
onto the medial section of the otoglossal. The
tendon is 1.5 to 2.0 mm in length.

Hehosteoijpsiloideus originates as a thin, nar-
row muscle from the dorsal surface of an in-
cription on the rectus cervicis profundus. Tlie
particular inscription is the second one posterior
to the second basibranchial. The fibers extend
directly anterior at first, and then turn medially,
dorsal to the rectus cervicis superficialis, and in-
sert onto the dorsal surface of the second basi-
branchial.

Htjoglossus is small, being slightly less than
1.0 mm in length. It originates from the dorsal
surface of the anterior end of the first basibran-
chial and extends laterally to insert on the tip of
the second radial.

NORTHWESTEHN SALAMANDER.

Ambtjstoma gracile (Baird)

Siredon gracilis Baird, Pacif. R. R. Rep., Vol. 10,
Williamsons Route, Pt. 4, No. 4, 1859, p. 13, pi.
44, fig. 2.

The range of this species is from northern
California to British Columbia. It is found in
damp habitats and has been collected rather fre-
(juently under logs and bark, except during the
breeding season, at which time it resorts to
ponds and slow streams. It is less than 195 mm
in total length. Probably the most distinguishing
characteristic is the large, raised, kidney-shaped
parotid glands behind the eyes. There is also a
glandular region along the dorsal ridge of the
tail. The color is dark brown; however, the head
and tail may be slightly lighter. Some may be
marked dorsally with yellow spots or flecks.
There are 11 costal grooves, counting one each
in the axilla and groin. The legs are large and
strong. In contrast to most Ambi/stoma the ven-
tral edge of the tail tends to be knifelike, while
the dorsal edge is rounded. Vomerine teeth are
in a transverse line which may be broken into
three or four series.

Hyobranchial Skeleton (Fig. 5)

The first basibranchial is a rod-shaped struc-
ture Iving in the center of the area outlined by
the mandible. It contains no bone and has defi-
nite lines of articulation with the radials.

BioLt)ciCAL Series, \'ol. 16, No. 1

Hyobhanchium and Thuoat Myology

39

The first radials are moderately long and at-
tached to the anterior edge of the ceratohyals
by a hganient. Thev contain no bone.

The second radials are thin, cartilaginous
rods that curve dorsally to attach to the annu-
lar-shaped otoglossal. Neither the second radial
nor the otoglossal cartilages contains bone.

There is also a lack of bone in the first cera-
tobranchial, the second ceratobranchial, the epi-
hranchial. and the ceratohyal. The only element
which is ossified is the second basibranchial,
which a\'erages 78 percent bone for the two
specimen studied ( Table 1 ) .

The ceratohyals are wide, being exceeded in
a width-to-length ratio only bv Dicamptodon, A.
anmilatum, A. inaheei, and A. texanum (Table
14).

The second basibranchial, as mentioned
above, is the only ossified portion of the hyo-
branchium, the cartilage being located on the
tips of the two lateral arms. This structure is
stronglv triradiate, the anterior arm being al-
most as long as the two lateral ones.

Only in three cases have we found a smaller
ratio of mandible length to mandible width than
in A. gracilc. Those three are A. talpoideiim, A.
t. nehulosiim. and .A. f. melanostictum, the short-
est being found in A. talpoideiim (Table 9).

Masculature (Fig. 5)

Intermandibularis posterior has its origin
along the dorsomedial surface of the rami of the
mandible. The fibers extend transversely to in-
sert into the fascia of the linea alba. A small
area at the point of the chin is open so that the
ventral portion of the genioglossus is visible.
The posterior fibers overlap the anterior end of
the interossaquadrata. The anterior fibers of this
muscle are continuous across the midventral line.
Thev constitute a band that is 2.0 mm wide at
the midline.

Interlu/oideus posterior originates in fascia
that is attached to the posterior tip of the mandi-
ble, quadrate, squamosal, and the otic capsule.
The posterior fibers arise from the fascia cepha-
lodorsalis. The fibers spread out as they pass
obliquelv caudomediallv to insert along the skin
of the gular fold and the midventral line of fas-
cia.

Depressor mandibulae takes its origin from
the sfjuamosal bone, attaching along its entire
posterolateral surface except for the ventral 1-2
mm. It also arises from the lateral surface of the
otic capsule and from the fascia cephalodorsalLs.
The fibers converge onto the posterior tip of the
mandible.

Levator mandibulae corresponds to the de-
scription given earlier in the general section on
musculature except that the fibers of the levator
mandibulae extenius and the levator mandibu-
lae posterior intenningle considerably, making it
difficult to separate these two portions.

Interossaquadrata originates from the ventral
surface of the quadrate just anterior to the hyo-
cjuadrate ligament and from the ventral surface
of the hyoquadrate ligament itself. A few fibers
are also attached to the ventral aspect of the
posterior tip of the ceratohyal. The anterior fi-
bers extend anteromedially over the subarcualis
rectus I, the medial fibers extend directly me-
dially, and the posterior fibers extend caudo-
mediad. These posterior fibers lie parallel with
the anterior fibers of the interhyoideus posterior.
It inserts into the median raphe of the throat.

Subhijoideus attaches along the ventrolateral
margin of the posterior 2-3 mm of the ceratohyal.
It becomes a thin muscle as it fans out over the
anterior end of the subarcualis rectus I and is
easily removed from the intennandibularis pos-
terior as the deeper layers are exposed. It is
readily distinguished from the interossaquadrata

Table 14. Average width to length ratios of ceratohyal.
Species 0 .1 .2 .3 .4

Dicamptodon

eiisatus
Hhyacotritou

olympicus
Amhystoma

annulotum
Amhystovia

cinguhtum
Amhystomu

gracile
Ambustoma

jeffersonianum
Ambystoma

mabeei

Ambystoma

macrodactylum
Amhystoma

maculatum
Amhystoma

talpoideiim
Ambystoma

opacum
Atnbystoma

texanum
Amhystoma

rosacea m
Am}>y\t(>ma

ti^rinutit
Amhystoma tigrinum

mclarwsiictum
Ambysioma tigrinum

Twffulositm

40

Bricham Young University Science Bulletin

at its origin but blends in with the latter near
the insertion. The insertion is by way of: the dor-
sal surface of the fascia of the intermandibularis
posterior.

Geniolnjoideus originates from the posterior
edge of the mandible along a line which begins
about 1.5 mm lateral to the symphysis and ex-
tends laterally for approximately 1.0 mm. The
anterior end of this muscle is overlapped along
its medial edge by the ventral portion of the
genioglossus. The lateral fibers extend directly
caudad, whereas the medial fibers lie in a medio-
caudal direction for a short distance before con-
tinuing directly caudad. Insertion is onto the sec-
ond basibranchial cartilage, with the lateral fi-
bers attaching into the tendonous inscription of
the rectus cervicis superficialis.

Genioglossus originates in two parts, the ven-
tral portion from the ventral and posterior sur-
faces of the mandible beginning at the symphy-
sis and extending laterally about 2.5-3.0 mm. The
fibers extend caudomedially for a short distance,
overlapping the anteromedial end of the genio-
hyoideus as indicated above. Insertion is into the
aponeurosis lingualis. A few fibers also continue
on to the first basibranchial and the otoglossal.
This latter portion is undivided.

The dorsal portion of the genioglossus origi-
nates along the posterior surface of the mandible
from the symphysis to a point about 3-4 mm lat-
eral from the symphysis. The lateral fibers pro-
ceed dorsal to the tip of the first radial and at-
tach into the fascia that forms the floor of the
mouth. The medial fibers insert onto the poste-
rior edge of the lateral half of the second radial.

Subarcualis rectus I originates along the dor-
sal edge of the tip of the epibranchial. These fi-
bers fonn a strong sheath around the epibran-
chial and then extend forward over the ventral
surface of the ceratohyal. The insertion is along
the anterior margin of the expanded end of the
ceratohyal.

Rectus cervicis snperficiuUs originates along
the lateral edge of the dorsal surface of the
sternum and from the tendonous inscription
which separates this nuiscle from the rectus ab-
dominus. It inserts onto the dorsal surface of the
second basibranchial and then extends forward
onto the posterior end of the first basibranchial
and the posterior edge of the medial end of the
first eeratobranchial.

Rectua cervicis profundus is actually a contin-
uation of the rectus abdoininus. It continues for-
ward at the level of the sternum as a flat band
and is situated dorsolati'rally with respect to the
rectus cervicis superficialis. The fibers turn
slightly medially at the lev(>l of the second basi-

branchial and then pass through the opening be-
tween the ceratobranchials, where they extend
dorsal to the first basibranchial and the hyoglos-
sus and finally inserts along the posterior edge of
the otoglossal.

Hchostcoijpsiloideus consists of a narrow
band of fibers arising from the dorsal surface of
the second myoseptum of the rectus cervicis pro-
fundus posterior to the second basibranchial. It
lies along the dorsomedial edge of the rectus
cervicis profundus and extends forward and me-
dially to insert along the lateral edge of the dor-
sal surface of the anterior arm of the second
basibranchial cartilage. Like A. taljwideum it
makes no insertion onto the lateral anns of the
second basibranchial.

Hyoglosstis originates from the dorsal surface
of the anterior end of the first basibranchial and
along the lateral edge of this cartilage to the
base of the second radial. The anterior fibers ex-
tend laterally and posteriorly to insert onto the
posterolateral edge of the second radial. The
posterior fibers extend more directly lateral and
insert into the fascia adjoining the second radial.

Jefferson's Salamander.

Ambystonui jeffersonianum (Green)

Sahunandra jeffersoniana Green, Contr. Maclur.
Lye, Vol. 1, No. 1, Jan. 1827, p. 4. pi. 1, fig. 1.

Tliis is a rather slender species which may at-
tain a length of 185 mm. Adults have faint blu-
ish markings on the sides. The ground color is
dark brown. There are 12 costal grooves and a
prominent gular fold. Toes are very long and
slender. The vomerine teeth form a transverse
line that is interrupted back of the internal nares
and occasionally at the midline. The range ex-
tends from Hudson Bay south to Virginia. It is
found westward to Wisconsin, Illinois, and Ar-
kansas.

HVOBRANCHIAL SKELETON (Fig. 6)

The first basibranchial is shaped like an
elongated pear when viewed from a ventrodorsal
direction and is a flat oval when viewed in cross
section. It contains a central mass of bone
amounting to about 37 percent of the total vol-
ume (Table 1). Definite lines of articulation
exist between this stnicture and the radials.
There are only four other species in which the
first basibranchial is more anteriorly located
(Table 12).

The first radial cartilages contain no bone
and are attached to the anterior tips of the cera-
tohvals bv small ligaments. These radials are

Biological Series Vol. 16, No. 1 Hyobranchium and Tuboat Myology

41

short in relation to the length of the first basi-
branchial. This ratio is the same in A. macula-
turn and smaller only in A. cingulaturyi (Table
6).

The second radials are broad at their bases
and taper quickly to fine tips which articulate
with the tips of the otoglossal. The otoglossal is
annular in shape, and neither the second radials
nor the otoglossal contains bone. There are no
centers of ossification located on the first cera-
tobranchials.

The epibranchials are long and contain bone
only in their tips. It is because of their length
that the length ratio of the first ceratobranchial
to epibranchial is so small (Table 13). Only in
Dicamptodon, A. taljtoideum, A. t. nebulosum,
and A. t. melanostictum is this ratio smaller. An
average of more than one-third of the volume
of the epibranchial is ossified (Table 1).

No bone is found in the second ceratobran-
chial. These cartilages are moderately long as in-
dicated bv the fact that the length ratio of the
second ceratobranchial to the first ceratobran-
chial is greater only in four other species— A.
anmdatum, A. maculatum, A. rosaceum, and A.
texanum (Table S).

There is a center of ossification in the pos-
terior tip of each ceratohyal. The average per-
cent of ossification is 6.4 (Table 1). These
cartilages are rather narrow when compared to
the ceratohyal width-to-length ratios of the
other species (Table 14).

Only the tips of the lateral arms of the
second basibranchial are chondrified. The
structure is distinctly triradiate in shape, and
averages 80 percent bone.

Musculature (Fig. 6)

Intermandibidaris posterior originates along
the dorsomedial surface of the rami of the man-
dible. A small area at the point of the chin is
open through which the genioglossus can be
seen. The fibers extend medially to insert into
the fascia of the linea alba. The posterior fibers
also extend slightly caudad to overlap the an-
terior end of the interossaquadrata.

Interhijoideua ])osterior originates in fascia,
which is attached to the posterior tip of the
mandible, the quadrate, the scjuamosal, and the
otic capsule. Posterior fibers also originate in
the fascia cephalodorsalis. The fibers fan out
over the throat to insert into the skin of the gular
fold and the midventral fascia. The anterior
fibers blend in with those of the interossaquad-
rata.

Depressor niandihuhw originates along the
posterolateral surface of the dorsal half of the

squamosal, the posterolateral surface of the otic
capsule, and from the fascia cephalodorsalis.
The insertion is onto the posterior tip of the
mandible.

Levator nmndihulae appears to have no sig-
nificant differences between the configuration of
this muscle and the description given in the
section on general musculature.

Interossaquadrata originates along the pos-
terior surface of the quadrate, and at this end
of the muscle these fibers lie ventral to the
origin of the subhyoideus. The insertion is into
the fascia of the linea alba at the midventral
line. The insertion end is very delicate and is
easily pulled loose with the overlying inter-
mandibularis posterior. As mentioned above, the
posterior fibers blend in with those of the inter-
hyoideus posterior so that a borderline between
these muscles is located only by tracing the
course of individual fibers back to their origin.

Subhyoideus originates along the ventro-
lateral margin of the posterior tip of the cera-
tohyal. This muscle passes forward and medially
to blend in with the anterior edge of the inter-
ossaquadrata and inserts into the dorsal side of

Table 15. Average width to length ratio of first basi-
branchial.

Species

0 .1

.3 .4 .5 .6 .7

Dicamptodou

CTisatus
Rhyacotritott

olyjJipicus
Anihystoma

annulatum
Anihystoma

cinguUitum
Amhystoma

^racile
Amhystoma

jefjiTsoniamim
Amhystoma

maheei
Amhystoma

macrodactylum
Amhystoma

maculatum
Amhystoma

talpoideum
Amhystoma

opacum
Amhystoma

texanum
Amhystoma

rosaceum
Amhystoma

tigrinum
Amhystoma tignnum

mehinosiictum
Amhystoma tigriruim

nchulostim

42

BmciiAM YoUNC Univeiisitv SriEXt k Bulletin

the tasciii of the intermandibularis posterior.
This muscle is also very dehcate and is easily
removed inadvertently with the intermandib-
ularis posterior.

Geniohyoideus takes its origin from a short
line along the posterior surface of the mandible,
the line beguming about 1.5 mm lateral to the
symphysis and extending laterally for approxi-
mately 2. .5 to 3.0 mm. The lateral fibers coming
from this line extend directly caudad, whereas
the medial fibers extend medially for a short
distance and then proceed directly caudad. This
muscle appears as a band proceeding between
the chin and the second basibranchial into which
it inserts. A few fibers also insert laterally into
the tendonous inscription alongside of the second
basibranchial.

Genioglossus originates in two parts, the ven-
tral along a line on the posterior surface of the
mandible. This line begins about 2.0 mm lateral
to the symphysis and extends laterally along the
mandible for about another 2.0 mm. This origin
is immediatelv ventral to the medial half of the
origin of the geniohyoideus. It inserts into the
aponem'osis lingualis, with a few fibers continu-
ing on to the first basibranchial and the oto-
glossal.

The origin of the dorsal portion of the genio-
glossus is along a line on the posterior surface
of the mandible just dorsal to the origin of the
geniohyoideus. The fibers extend laterally as well
as caudallv to insert into the floor of the mouth.
The medial fibers extend caudally to the apo-
neurosis lingualis and then oblicjuely in a caudo-
lateral direction to attach to the tip of the second
radial.

SuharcualK rectus I originates on the pos-
terior tip of the epibranchial and forms a sheath
around the cartilage. It also extends out onto the
ventral surface of the ceratohyal where it inserts
along the anterior margin. The two ceratohyals
almost touch each other at their anterior ends.
They overlap the lateral edge of the anterior
half of the first basibranchial, and these muscles,
therefore, are also almost touching at that point.

Rectus cervicis superfickilis originates on
the posterior edge of the dorsal surface of the
sternum and from the tendonous inscription
which separates this muscle from the rectus
abdominus. These fibers extend forward over
the pericardium attaching to the second basi-
branchial and then continuing forward to insert
onto the posterior tip of the first basibranchial
and the posterior edge of the medial end of the
first ceratobranchial cartilage.

Rectus cervicis- jjrofuiulus extends forward
from the sternum as a stout band of muscle

situated alongside of the rectus cervicis super-
ficialis. It partially overlaps the latter dorsally
as the two muscles move forward to the level of
the second basibranchial. The archway between
the two ceratobranchials serves as a passageway
for this muscle as it continues forward to finally
attach to the posterior edge of the otoglossal.

Ilebosteoiij)siIaideus originates from one of
the myosepta of the rectus cervicis profundus.
The particular myoseptum is the second one
posterior to the second basibranchial, from whose
dorsal surface it arises as a narrow band and
proceeds directly anteriorly. After a short dis-
tance, it turns medially to insert onto the dorsal
surface of the second basibranchial.

Hyofdossus is a tiny muscle originating on
the dorsal surface and lateral margin of the an-
terior end of the first basibranchial. It extends
laterally to insert on the tip of the second radial
cartilage and into the adjoining fascia.

Mabee's Salamander.
Ambystoma mabeei (Bishop)

Ambystoma nuibeei Bishop, Joum. Elisha
Mitchell Sci. Soc, Vol. 43, No. .3-4, 1928, p. 157.

Tliis species inhabits rotted logs and bark in
moist areas of the Carolinas and Georgia. The
color above is deep brown, and the sides are
covered with white or grayish flecks which be-
come indistinct as they approach the dorsal sur-
face, where they appear to be overlain with
darker pigment. This salamander resembles A.
texanum in size, color, and shape of the head.
It reaches 102 mm in total length. The vomerine
teeth form a transverse line that is broken back
of the internal nares. There are 13 costal
grooves, counting two that come together in the
groin and one in the axilla.

Hyobranchial Skeleton ( Fig. 7 )

The first basibranchial cartilage in this spe-
cies is pear-shaped when viewed from its ven-
tral surface. It is a flat oval in cross section. Tlie
core is bony and is surrounded on all sides by a
layer of cartilage. The average percent of ossifi-
cation is 37. The first basibranchial is situated in
a rather posterior position as compared to its
position in other species, being exceeded in this
respect onlv by A. anrutlatum, A. cingulatum,
and Rhyacotriton (Table 12). The length of the
first basibranchial as compared to the total
length of the hyobranchium is greater in this
species than in others except A. TaJpoidcum, A.
opiicuin. and Rln/cicotriton (Table 4).

The first radials are entirclv cartilaginous and
arc relatively longer than in the other species ex-

Biological Sehies, \'ol. 16, No. 1 HvoniiANCiiiuM and Thuoat Myology

43

Table 16. Average ratios of epibranchial length to
length of first branchial arch.

Specie.s 0 .1 .2 .3 .4 .5

Dkaniptodon

crisatus
Wiyacutriton

olympicus
Amhystonw

tiutiuiatiini
Aminjfitomu

ciu^ulatum
Ambystoma

gracile
Atnbifstoma

jeffcrsoniatiuui
Amijystoma

rnabeei
Anihyfitoma

macwdactylum
Amhystoma

maculatum
Amhystoma

talpoideum
Aml}ystoma

opacum
Ambystoma

tcxanum
Ambystoma

Tosaceum
Ambystoma

tigrinum
Amhystoma tigrinum

meUiTWstictum
Amhysioma tigrinum
riehulosum

cept Dicamptodon and Rliijacotriton (Table 6).
Their tips are attached to the anterior tips of the
ceratohvals by a short hganient, and a definite
hne exists at their base, indicating the point of
articulation with the first basibranchial.

The second radials attach to the sides of the
first basibranchial and then taper to a fine point
in an anterodorsal direction. These distal tips
articulate with the otoglossal, which is an annu-
lar-shaped rod of cartilage arched forward and
dorsad into the tongue.

There is no bone in the first ceratobranchial.
It is flattened dorsoventrallv and forms a smooth
arch which is continued distallv bv the epibran-
chial. The epibranchial averages 58 percent os-
sification, the bony portion constituting the dis-
tal end (Table 1)'.

The second ceratobranchial is entirely carti-
lage. Its distal end articulates with the medial
side of the epibranchial at a point well posterior
to the first ceratobranchial-epibranchial articu-
lation.

The ceratohyal is longer in relation to the
total length of the hyobranchium in this species

than in other ambystomids, except for Rhyacotri-
t07i ( Table 19 ) . The average percent ossification
for this structure is 6.S (Table 1).

Only the extreme tips of the lateral arms of
the second basibranchial are cartilage. The ma-
jority of the volume of this structure (63 per-
cent) is bone. The anterior arm is much shorter
than either of the lateral arms.

The ratio of the mandible length to mandible
width is exceeded only by Dicamptodon (Table
9).

Musculature ( Fig. 7 )

Intermandibtdaris posterior originates on the
dorsomedial surface of the mandible, and the
fibers extend medially to the linea alba where
they insert. The ventral portion of the genioglos-
sus is visible at the point of the chin. The in-
terossaquadrata and subhyoideus muscles are
partly overlapped ventrally by this muscle. In
contrast to most of the other species, the poste-
rior fibers in this salamander exend almost di-
rectly transversely with only a slight deviation
caudad.

Interhyoideus posterior is broad in this spe-
cies, the anterior fibers extending directly me-
sially to the midline. The origin is by way of
fascia which attaches to the posterior tip of the
mandible, the squamosal, the fjuadrate, and the
otic capsule and is also continuous with the fas-
cia cephalodorsalis. The fibers pass over the
mandible tip, cross the middle section of the
subarcualis rectus I and insert into the skin and
fascia of the gular fold and into the linea alba
midventrally.

Depressor mandihulae fibers arise from the
fascia cephalodorsalis, dorsal end of the squamo-
sal and the posterior and ventral surfaces of the
otic capsule. They converge to a short, tough
tendon which inserts onto the posterior tip of the
mandible.

Levator itwndihttlac consists of four portions
as outlined in the general section on mascula-
ture.

Jntcrossuiptadrata originates on the posterior
surface of the cjuadrate and the ventral surface
of the ceratohyal-tjuadrate articulation. The fi-
bers lie in an anteromedial direction. They in-
sert into the fascia of the linea alba as it pro-
ceeds posteriorly from the intermandibularis pos-
terior. This muscle is narrower in this species
than in the others since there are no fibers ex-
tending in a posteromedial direction.

Sublujoideus originates from the ventral sur-
face of the posterior tip of the ceratohyal. The
fibers at the origin are dorsal to the interossa-
(juadrata, but they proceed in a more anterior

44

Bhigiiam Vounc IInisehsity Science Bulletin

direction than those ot the interossatiuadrata, re-
sulting in a side-bv-side location of their distal
ends. From a point about midway along the
length of these muscles, they appear as a single
muscle since no definite line of separation is visi-
ble between them. The subhyoideus inserts into
the dorsal surface of the fascia of the interman-
dibularis posterior.

Geniohyoidcus originates from the posterior
surface of the mandible along a line that begins
at the symphysis and extends about 1.5 mm lat-
erally. The medial fibers pass around the genio-
glossus at the point where the fibers of the latter
converge and pass dorsad. From this point pos-
teriorly, the fibers proceed directly caudad to in-
sert onto the anteroventral surface of the second
basibranchial and into the inscription that is
continuous with the latter's lateral arm.

GeniogJossus originates in two parts. The ven-
tral portion arises along the ventral surface of
the mandible, beginning at the symphysis and
extending laterally about 1.5 mm. The origin is
ventral to the origin of the geniohyoidcus. The
fibers converge toward the midline where they
attach to the aponeurosis lingualis. From this
point, they extend directly caudad and, together
with the medial fibers of the dorsal genioglossus,
form a broad sheet of muscle contributing sub-
stantially to the anterior half of the floor of the
mouth. The insertion is onto the anterior edge
of the first basibranchial and onto the otoglossal.

The origin of the dorsal genioglossus is im-
mediately dorsal to the origin of the ventral por-
tion. The fibers tend to fan out as thev proceed
caudad and in so doing form the broad sheet of
muscle mentioned above. The medial fibers at-
tach to the tip of the second radial. The lateral
fibers insert into the fascia of the mouth floor
at the base of the tongue.

SuharcualLs rectus I originates from the dor-
sal edge of the epibranchial cartilage. The fibers
form a sheath of muscle which completely en-
closes the epibranchial and then moves anterior-
iv parallel with the ceratohyal. At the level of
the articulation between the epibranchial and
the second ceratobranchial, the sheath opens up
medially to pennit the entrance of the second
ceratobranchial into the sheath. From this point
anteriorly, the muscle gradually loses its sheath-
shap(> and comes to lie on the ventral surface of
the ceratohyal. It inserts onto the anterior edge
of this cartilage.

Rectus cervicis superjicialis originates from
the lateral half of the dorsal surface of the ster-
num and from the tendonous inscription that ex-
tends laterally Iroin the steniuuL There are three
iMVOsepta lietween the sternum and the second
basibranchial. The oinohyoideus arises along the

lateral margin of this muscle between the first
two of these mvosepta. The rectus cervicis super-
ficialis lies dorsal to the coracoids and ventral to
the pericardium. A portion of the pericardium
can be seen along the midventral line between
these two muscles. The medial two-thirds of the
libers attach to the second basibranchial, but
only the lateral half of these continues forward
together with the remainder of the fibers to in-
sert onto the posterior tip of the first basibran-
chial and the posterior edge of the medial end of
the first ceratobranchial.

Rectus cervicis profundus is actually a for-
ward continuation of the rectus abdominus and
its origin is, therefore, from the puboischium of
the pelvic girdle. There are three myosepta along
the course of this muscle between the sternum
and the insertion. It is situated in a position dor-
sal and lateral with respect to the rectus cervicis
superficialis. The profundus is basically a thin,
broad muscle, but the fibers become compacted
into a thicker and narrower muscle belly at the
level of the hyobranchium. The medial half of
the opening between the ceratobranchials serves
as a passagewav through which the fibers pass.
From this point, they proceed forward, dorsal to
the first basibranchial and the second radial.
They finally insert onto the otoglossal at a point
well anterior to the first basibranchial.

Hebosteoypsiloideus is a narrow muscle
which originates from the second tendonous in-
scription of the rectus cervicis profundus, pos-
terior to the second basibranchial. It extends for-
ward dorsal to the rectus cervicis profundus and
then crosses over medially to insert onto the dor-
sal surface of the second basibranchial.

Hijo<yIosstis is a small muscle seemingly hy-
pertrophied in this species, as compared to the
same muscle in other species. It originates on
the dorsal surface of the anterior end of the first
basibranchial and from the dorsal surface of the
articulation between the first radial and the first
basibranchial. The fibers converge slightly so
that the insertion is narrower than the origin.
Thev wrap around the ventral aspect of the tip
of the second radial and insert along the latter's
posterior edge.

Long-toed Salamander.

Amhystoma macrodactylum Baird

Amhijsioma macrodactyla Baird, Joum. Ac. Nat.
Sci. Phila., Ser. 2, Vol. 1, 1849, p. 292.

This species has been found in breeding
ponds, beneath rocks that are just above the
waterline of these ponds, and Ix'neath the loose
bark of fallen trees. It is not unusual to find

BroLOGiCAL Sebies, Vol. 16, No. 1

HvoiiH.\N(:illUM .\N1) THIU).\T .\IvOLO(;V

45

them in the breeding ponds while these ponds
are still icebound in the spring. The geographi-
cal range is along the Pacilic Coast from Califor-
nia to British Columbia and Alberta, and then
eastvvard into Idaho and Montana. It has also
been reported in parts of Iowa. This is a slender
animal whose total length may reach 12S mm.
The chocolate brown to black ground color is
topped by a median dorsal stripe that is dull
tan to bright yellow in color. In some individuals,
this stripe is greenish yellow. It may occur along
the entire length of the animal's body, including
head, trunk, and tail, or it may begin just behind
the head. The usual number of costal grooves,
including one each in the axilla and groin, is 12.
The vomerine teeth form a broken line across the
roof of the mouth. There are four sections to this
line, the two middle ones being the longest, con-
taining 9 to 10 teeth each, and lying wholly be-
tween the internal nares. The lateral sections
contain 3 to 5 teeth each and are directed pos-
terolaterallv behind the inner nares.

Hyobraxchial Skeleton (Fig. 8)

The first basibranchial cartilage is a pear-
shaped structure with a central core of bone
surrounded by a layer of cartilage. The bony
portion constitutes .34 percent of the total volume
( Table 1 ) . There are definite lines of articula-
tion with the radials. The first basibranchial is
long when compared to the length of the first
ceratobranchial. The ratio of lengths of first basi-
branchial to first ceratobranchial is exceeded
only by A. opacuin (Table 5). This structure is
also rather narrow. The ratio of width to length
is smaller only in Rhijacotriton (Table 15).

The first radials are long in relation to the
first basibranchial length (Table 6). They are
entirely cartilaginous and are connected to the
ceratohyal tip by a short ligament.

The second radials are likewise entirely carti-
laginous. They articulate with the tips of the
otoglossal, the latter structure forming a carti-
lage ring which arches dorsad into the substance
of the tongue.

The first ceratobranchial forms a smoothly
arched bar of cartilage completely devoid of os-
sification. The epibranchial is also completely de-
void of bone. It continues the smooth arch be-
gun by the first ceratobranchial. The second
ceratobranchial, like the first branchial arch, con-
tains no bone.

A small tip of bone exists in the posterior end
of the ceratohyal in this species and constitutes
5 percent of the total volume of this element
(Table 1).

The triradiate second basibranchial was

found to be 50 percent cartilage and was shorter
from side-to-side in comparison to its anteropos-
terior length than the same structure in other
ambystomids (Table 18).

Although the mandible length appears quite
short, the ratio of length to width is actually
smaller in several other species (Table 9).

Musculature (Fig. 8)

Intermandibularis posterior originates along
the dorsomedial edge of the mandible. There
are fibers lacking at the point of the chin, and
consequently the genioglossus is visible there.
The fibers lie mostly in a medial direction where
they attach into the linea alba of the throat. The
fibers from each side do not quite meet at the
midline. The ratio of distance bctvveen the men-
tal symphysis and the posterior edge of this mus-
cle to the distance between the mental sym-
physis and the pcctoralis muscle is smaller in
this species than in any of the others (Table 10).

Interhi/oideus posterior has a complex origin
with the lateral fibers arising from the fascia
cephalodorsalis, and the middle fibers attaching

Table 17. Average ratios of length of second cerato-
branchial to total length of first branchial arch.

Species

0 .1

.6 .7 .8

Dicamptodon

CHsatus
Hlufacothtou

ohfmpicus
Amhtfstoma

anuulatuni
Amhtfstonui

cinouldtuni
Ambifstojuu

Amht/stonui

jcfjeraoniamim
Ajtihijstoma

mahcci
Aiuhijstoniu

nmcrodacttjluni
Amhystoma

maculatum
Ajtihystonia

talpoidcimi
Amhystoma

opacum
Amhy.stoJna

tcxanu7u
Amhystotnii

rosaceum
Amhystoma

tifihnum
Amhystoma tignnum

mcUinosiictum
Amhystoma ti^riuum

ruhulosiim

46

to the posterior tip of the mandible just anterior
to the insertion of the depressor mandibulae. By
way of fascia, this muscle also attaches to the
lateral surface of the mandible tip, the lateral
surface of the mandibulo-tjuadrate articulation,
and the lateral surface of the ventral end of the
sfjuamosal. The fibers fan out over the posterior
throat region to insert along the skin of the gu-
lar fold and into the midvcntral line of fascia,
the linea alba.

Depressor nuindibulae originates from the
posterolateral edge of the squamosal except for
the ventral 1-2 mm, which attaches to the fascia
cephalodorsalis and the lateral surface of the otic
capsule. It inserts onto the posterior tip of the
mandible.

Levator numdihulue is essentially the same as
that description given in the general musculatiu-e
section of this paper.

Interossiujuadrata originates on the posterior
surface of the ([uadrate, the ventral surface of
the articulation of the ceratohyal and the quad-
rate, and from the posterior tip of the mandible.
The fibers spread out over the throat with the

Table 18. Average ratios of anterior to posterior length
to second basibranchial to width of second basi-
branchial.

Species

0

.1

.3

.5

enmtMs

oUfmpicus

tmnuUitum
Atuhi/.stoma

fin^ulotum
Amhystoma

gracilv
Amhtjstoriui

jcffcr.soutaiium
Amhystoma

maheei
Amhystoma

macwdactyhim
Amhystoma

morulatum
AmhystoTua

tatpoideum
Amhystoma

opacum
Amhystoma

texouum
Amhystoma

romrtum
Amhystoma

tifi^rimun
Amhystoma ti^nmnn

mehnwstutum
Amhm'toma tigrinum

nehulosum

,

Bbigham VouNt; University Science Bulletin

anterior fibers directed anteromedially and the
posterior fibers in a posteromedial direction. The
insertion is into the fascia of the linea alba.

Sublitjoideus originates along the anteroven-
tral edge of the posterior tip of the ceratohyal.
It fans out as it proceeds forward over the throat
and inserts into the dorsal fascia of the interman-
dibularis posterior. Tlie posterior fibers blend in
with those of the interossacjuadrata so that no
liorder is distinguishable between these two mus-
cles.

Genioliijoideus originates along a line on the
posterior surface of the mandible which begins
about 1.0 mm lateral to the symphysis and ex-
tends laterally about 1.0 additional mm. The fi-
bers pass caudad and mediad, and then directly
caudad to fomi a stout band on either side of the
midline. It inserts onto the second basibranchial
and in the tendonous inscription that extends
laterally from the arms of the second basibran-
chial.

Genioglosstis has a ventral portion which orig-
inates on the ventral and posterior surfaces of
the mandible, beginning at the symphysis and
extending laterally approximately 1.0 mm. The
fibers overlap the anterior end of the genio-
hyoideus and converge medially to attach them-
selves into the aponeurosis lingualis before con-
tinuing to attach to the anterior tip of the first
basibranchial and the otoglossal.

The lateral fibers are directed laterally as
well as caudad and form a thin sheet of muscle
which inserts into the fascia forming the floor
of the mouth. These fibers lie in a dorsolateral
position with respect to the anterior end of the
ceratohyal. The medial fibers proceed directly
caudad into the aponeurosis lingualis and then
to the tip of the second radial.

SuburctuiJis rectus I originates on the tip of
the epibranchial cartilage, from which its fibers
wrap around and enclose the latter in a muscular
sheath and then continue forward over the ven-
tral surface of the ceratohyal. The insertion is
along the ventral siuface of the anterior margin
of the ceratohyal.

Rectus cervicis s-uperficialls originates along
the dorsal surface of the posterior end of the
steminn and the tendonous inscription which
separates this muscle from the rectus abdominus.
The fibers extend forward ventral to the peri-
cardium. In this species there is a large section
of the pericardium which is exposed at the mid-
line between the two rectus cervicis superficialis
muscles. The second basibranchial is attached to
this muscle and serves as a landmark between
the thin, broad portion of the muscle and the
anterior portion which becomes thick and nar-

Biological Series. Vol. 16, No. 1

Hyoiiii.vnciiium .\ni) Tiiiio.\t Mvoloov

47

row. This anterior portion inserts onto the pos-
terior tip of the tirst basibranchial and tlic pos-
terior ed£;e of the medial end of the ceratobran-
chial.

Rectus cervicis profundus at the level of the
sternum extends forward alongside of and deep
to the rectus cervicis superficialis. After passing
dorsal to the omohyoideus, it turns slightly me-
diad and proceeds through the opening between
the ceratobranehials to insert onto the posterior
edge of the otoglossal.

Hebostcoi/psiloideus originates as a narrow
muscle on the dorsal surface of the rectus cervi-
cis profundus. It is separated from the latter only
bv careful dissection since the fibers of these
two muscles lie parallel with each other. It is
from the second myoseptum posterior to the sec-
ond basibranchial that this muscle arises. It
crosses the rectus cervicis superficialis and then
inserts onto the dorsal surface of the second basi-
branchial.

Ihiof^lossus is a small muscle situated be-
tween the dorsal surface of the first basibran-
chial and the ventral surface of the rectus cervi-
cis profundus. It originates along the dorsolateral

Table 19. Average ratios of length of ceratohyal to
total length of hvobranchium.

Species 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0

Dicamptodon

cnsatus
Hfujucotrituit

olympicus
Ambystoma

anmUatum
Ambystoma

cingulatum
Aml)ystoma

gracile
Ambystoma

jeffcrsoiiUinujn
Ambystoma

mabeei
Ambystoma

nwcrodactylum
Ambystoma

muciilatum
Ambystoma

talpoideum
Ambystoma

opacum
Ambystoma

texanum
Ambystoma

rosaceum
Ambystoma

tigrinum
Ambystoma tigrinum

mcUttu>\tiituTji
Ambysioma tigrinum

nehutosum

edge of the first basibranchial between the an-
terior tip and the base of the second radial. The
anterior fibers extend obli([uely in a posterolat-
eral direction whereas the posterior fibers extend
directly laterad and converge with the former to
insert onto the posterolateral edge of the second
radial as it curves dorsad into the tongue.

Spotted Salamander.

Ambystoma maculatum (Shaw)

Lacerta nmculata Shaw, Gen. Zool., Vol. 3, Pt. 1,
1802, p. 304.

The spotted salamander is distinguished by
a row of large, round, yellow or orange spots oc-
curring on either side of the middorsal line. The
ground color is a deep bluish black. Males are
usually more slender than females, and the for-
mer are readily recognized in the breeding sea-
son by the swollen vent. Total length reaches
206 mm. There are usually 12 costal grooves, a
prominent gular fold, and, in this species, larger
legs. The vomerine teeth extend across the
roof of the mouth in a smoothly curved line im-
mediately b?hind the internal nares. This line is
sometimes interrupted at the midline and in
back of the internal nares. The geographical
range is from Nova Scotia to Wisconsin and
southward to Florida, Louisiana, and Texas.
They are found in ponds, slow moving streams,
or temporary pools which offer .spring breeding
sites.

Hyobranchial Skeleton (Fig. 9)

The first basibranchial is a pear-shaped
structure when viewed ventrally and has a cen-
tral core of bone. In cross section it is flat oval
in shape. The bone constitutes an average of
about 25 percent of its volume ( Table 1 ) . There
are lines of articulation visible between the first
basibranchial and the radials.

The first radials are short in relation to the
length of the first basibranchial. The only other
species with a smaller ratio of length of first ra-
dial to length of first basibranchial is A. cingula-
tum, (Table 6). No bone is found in these struc-
tures.

The second radials articulate with the oto-
glossal at their distal tips. The otoglossal forms
a thin arch of cartilage extending dorsad into
the tongue.

The first ceratobranehials are nearly straight,
the only prominent curve in the first branchial
arch occurring in the anterior half of the epi-
branchial. There is no ossification in either cera-
tobranchal.

18

HmciiAM VouNO Univehsii Y Science Bulletin

The epibranchial is ossified at its tip, the os-
sification making up an average of 32 percent of
the total vokime (Table 1). The width of this
element is unifonn along most of its length ex-
cept near the tip where it tapers abruptly to a
point.

The ratio ot the lengths ot the second cerato-
branchial to the first ceratobranchial is high in
this species, being exceeded only by A. annula-
tiim (Table S). This element is unossified.

The posterior tip of the ceratohyal is ossified
and constitutes an average of 10.6 percent of the
total volume (Table 1). The anterior tips over-
lap slightly at the midline.

The lateral arms of the second basibranchial
are much longer than the anterior arm, and it is
on these lateral tips that the cartilage is found.
This cartilage constitutes 27 percent of the total
(Table 1)."

The mandible is longer than it is wide, the
ratio of length to width averaging 1.06 for the
five specimens of A. maculatum used in this
study (Table 9).

Mu.SCULATURE (Fig. 9)

Intenmiiulihularis posterior originates along
the dorsomedial surfaces of the rami of the man-
dible. There is a small opening at the angle of
the chin through which the genioglossus can be
seen. This muscle overlaps the anterior end of
the interossa(|uadrata and the anterior portion
of the subhyoideus. The fibers pass, basically, in
a transverse direction to the midventral line
where they insert. There is no broad fascial sheet
in this species occupying the midventral line as
is seen in most other species.

In one of five specimens the anterior fibers of
the intermandibniaris posterior were continuous
across the midline. This band was 0.7 mm wide
at the midventral line.

Interhyoidcus posterior originates on the fas-
cia cephalodorsalis and on the posterior tip of
the mandible immediately medial and anterior
to the insertion of the depressor mandibulae.
The majority of this muscle attaches by way of
a fascial sheet to the lateral surface of the tip of
the mandible, the lateral surface of the articula-
tion between the mandible and the quadrate,
and the lateral surface of the ventral end of the
s(|ua!nosal. This muscle inserts into the gular
fold and midventral fascia of the throat.

Depressor mandibulae originates from the
fascia cephalodorsalis, ihe lateral surface of the
otic capsuk', and the dorsal half of the lateral
surface of the stjuamosa. The fibers converge to
insert onto the posterior tip of the mandible.

Levator mundihulae has its origin and inser-
tion essentially as in the description given in the
section on general musculature.

JnterossMpiadrata originates from the poste-
rior surface of the (|uadrate and the ventral sur-
face of the articulation between the quadrate
and the ceratohyal. The most posterior fibers al-
so attach to the posterior tip of the mandible and
lie ventral to the other fibers of this muscle.
These ventral fibers extend caudomesially
whereas the dorsal fibers extend anteromesially.
The insertion is into the linea alba as it con-
tinues caudad from the intennandibularis poste-
rior.

Sublnjoidctts originates from the dorsum of
the posterior or distal tip of the ceratohyal and
inserts into the dorsal fascia of the intennandib-
ularis posterior.

Geniohijoideus originates along the posterior
surface of the mandible along a line beginning
just lateral to the symphysis and extends laterally
approximately 3.0 mm, deep to the ventral por-
tion of the genioglossus. This muscle is narrower
anteriorly than posteriorly, and consequently the
genioglossus is exposed along the midline in this
species. There are some deep fibers which at-
tach to the aponeurosis lingualis and extend
posteriorly with the remainder of the muscle.
The muscle inserts onto the second basibranchial
and into the tendonous inscription which sepa-
rates this muscle from the rectus cervicis super-
ficialis.

Genioglossus has a ventral portion which orig-
inates on the ventral and posterior surfaces of
the mandible beginning at the svinphysis and ex-
tending laterally for about 3.0 mm. Tlie fibers ex-
tend in a eaudomedial direction to form a small
triangular-shaped muscle. This portion of the
genioglossus overlaps the anterior end of the
gcniohyoideus. The insertion is into the aponeu-
rosis lingualis which continues caudad to attach
to the anterior end of the first basibranchial
c-artilage. A few fibers coiitinui- along in associa-
tion with the aponeurosis lingualis and insert on-
to the otoglossal.

The dorsal portion of this muscle attaches
along the posterior edge of the mandible begin-
ning at a point on the mandibular symphysis
and extending laterally for about 2.0 mm. The
fibers fan out as a thin sheet which contributes
to the anterior floor of the mouth. The medial
fibers extend caudad to insert onto the posterior
edge of the second radial tip, while the lateral
fibers continue caudad and laterally to attach in-
to the fascia lateral to the base of the tongue.
The aponeurosis lingualis, in this as well as most
other species of this family, sends a slip onto the

Bioloc:k:al Series. \'ol. 16, No. 1 Hv()BR.\nchium .\Nn Thro.\t Myology

49

Table 20. Average ratios of length of first basibranchial
to total length of first branchial arch.

Species

0

1

2

3

.4

Dicamptodon

f}L\atu.s
Rhijacutriton

oli/niiiU'its
Anihi/slonia

annitlatuin
Amhi/stotiui

cintiulutuni
Aml)tjstoinu

gracile
Ambt/stoma

jeffersonianum
Amhystonia

mabeei
Ambtjstomu

nmcwdactylum
Avtbystoma

maculatum
Arnbijstitvia

talpoidcum
Andnjstoutti

opacum
Ambystoma

texanum
Ambysloma

rosaceum
Amby.stonia

tigrinuru
Ambysionia tigrtnum

incl/inostictum
Aniliystdtna tigrintiiu

iiclnilo^tim

first ceratobranchial wliere tlie latter articulates
with the first basibranchial. It also sends fibers
onto the dorsal surface of the base of the first
ceratobranchial and the lateral portion of the
dorsal surface of the posterior end of the first
basibranchial.

Rectus cervicis pwfumltis is extended for-
ward alongside and dorsal to the rectus cervicis
superficialis. At the level of the second basibran-
chial, it begins to turn medially and become
thicker and narrower. It passes through the arch-
way fonned by the two ceratobranchials and
continues forward dorsal to the first basibran-
chial to its insertion on the posterior edge of
the otoglossal cartilage.

Hehosteoijpsiloideus originates from the dor-
sal surface of the most posterior inscription on
the rectus cervicis profundus. It extends forward
dorsal to the latter until, at the level of the omo-
hyoideus, it turns medially to cross the rectus
cervicis superficialis dorsally and insert onto the
dorsal surface of the second basibranchial.

Hijoglossus is a small triangular-shaped mus-
cle which originates on the dorsal surface of the
anterior end of the first basibranchial and along
the lateral edge of the dorsal surface of this car-
tilage to the base of the second radial. The fi-
bers converge to insert on the tip of the second
radial where the latter turns dorsad to join the
otoglossal. A few fibers, in addition, continue
laterally to attach to the fascia of the floor of
the mouth.

tip of the first radial cartilage and continues on
to attach the first radial to the anterior tip of the
ceratohyal.

Suharcualis rectus I originates on the tip of
the epibranchial cartilage. It wraps around this
tip and proceeds forward, medial to the angle
of the jaw where it then begins to fan out into
a thin sheet over the ventral surface of the cera-
tohyal. It inserts along the anterior margin of
the ceratohyal.

Rectus cervicis supcrjicialvi originates from
the dorsal surface of the sternum and the ten-
donous inscription which separates this muscle
from the abdominal muscles posteriorly. It
passes forward ventral to the pericardium. The
medial fibers attach to the second basibranchial,
which is actuallv partlv imbedded in this muscle.
The lateral fibers end on the tendonous inscrip-
tion which extends outward from the lateral arms
of the second basibranchial. .^t this point, the
nmscle continues forward again to become a
thick band which inserts onto the hyobranchial
skeleton. It inserts onto the posterior tip of the
first basibranchial and the posterior edge of the

Mole Salamander.

Ambystoma talpoideum (Holbrook)

Salamandra talpoiclea Holbrook, N. Amer. Herp.,
Ed. 1, Vol. 3, 18.38, p. 117, pi. 29.

This is a small species with a broad, de-
pressed head, a rounded body, and a short tail.
Total length ranges up to 97 mm. It has 10 cos-
tal grooves, counting one each in the axilla and
groin. Vomerine teeth fonn three distinct series,
the middle one being situated some distance
posterior to the internal nares. The lateral series
begin a short distance behind the internal nares
and extend a short distance laterally. The ground
color is dark brown dorsally, becoming bluish
gray along the lower sides. Small bluish white
flecks are scattered over the surface of the skin,
becoming numerous enough on the lower sides
to fonn large patches. The range of this species
is along the Southeast and Gulf Coasts of the
United States from North Carolina to Louisiana
and northward to Illinois. It has also been found
in Oklahoma, Arkansas, and Texas.

50

Bricham YouNn UN'ivmsiTY Science Bulletin

HvOBRANCHtAL SkFXETON (Fig. 10)

The ossification center which makes up tlie
core of the first basibranchial amounts to an
average of 32.6 percent of tlie total voKime. This
structure is pear-sluiped when viewed from tile
ventral surface, and the cross sectional area is
a flat oval. The lines of articulation with the
radials are distinct. The first basibranchial is
long compared to the total length of the entire
hyobranchium and as compared to the length ot
the first ceratobranchial (Tables 4 and 5).

There is no bone in the radials. The first pair
is free from the ceratohyals and unjointed. The
second pair attaches to the tips of the otoglossal.
The otoglossal cartilage is a smoothly curved rod
of cartilage which arches into the tongue from
below.

The first ceratobranchial is a slightly curved,
unossified, rod of cartilage. It is flattened dorso-
ventrally. The major bend in the first branchial
arch occurs in the anterior half of the epibran-
chial. The latter is also unossified. The epibran-
chial length in relation to the length of the en-
tire first branchial arch and in relation to the
length of the first ceratobranchial is greater than
in anv other species (Tables 13 and 16).

The second ceratobranchial is an unossified,
slightly curved rod of cartilage articulating at
its proximal end with the tip of the first basi-
branchial and with the anterior end of the epi-
branchial at its distal end.

The ceratohyal was ossified in one specimen
and unossified in the other two. In the specimen
with ossification, this amounted to 11 percent
of the total volume (Table 1). Only in A.
mabeei. A. t. iiwhinostictum, and Rhi/acotnton
is the ratio of ceratohyal length to total hyo-
branchial skeleton length greater than in this
species (Table 19).

The lateral arms of the second basibranchial
are longer than the anterior arm, and the former
contain cartilage on their tips. The average ossi-
fication is 76 percent.

The ratio of mandible length to mandible
width is much smaller than in other species
(Table 14), indicating a short mandible.

Musculature (Fig. 10)

Internumdihtilam posterior originates along
the dorsomedial margin of each mandibular ra-
mus. The fibers extend transversely and slightlv
posteriorly to attach to the fascia of the linea al-
ba. It covers more than three-tjuarters of the
area between the rami of the mandible. The an-
terior portion at the point oi the chin is missing,
and the genioglossus can be seen in that posi-
tion.

Intcrlitioidcm posterior originates from fascia
which attaches to the posterior tip of the mandi-
ble, tlie ([uadrate, the squamosal, and the otic
tapsuie. The posterior fillers originate in the fas-
cia cephalodorsalis. The origin overlaps the in-
sertion of the depressor mandibulae on the tip
of the mandible. The insertion is into the mid-
ventral raphe. The anterior fibers blend in with
those of the interossacjuadrata.

Depressor nuindibulae originates along the
dorsal end of the squamosal, the ventral and pos-
terior surfaces of the otic capsule, and along the
fascia cephalodorsalis, the fibers of this muscle
converge onto the posterior tip of the mandible
where they insert as a short, stout tendon. There
are also a few short fibers which extend from
the ventral surface of the ((uadrate to the poste-
rior tip of the mandible.

Levator inandihulae corresponds closely to
the description that is given in the section on
general musculature. There is considerable inter-
mingling of fibers between the levator mandibu-
lae posterior and the levator mandibulae exter-
nus.

Interossaquadrata is a fan-shaped muscle
which originates from the ventral surface of the
(juadrate anterior to the hyoquadrate ligament,
from the articulation of the ceratohyal and the
c{uadrate, and from the ventral surface of the
posterior tip of the ceratohyal. The anterior fi-
llers, arising from the quadrate, lie dorsal to the
posterior fibers and are directed in an antero-
medial direction. The posterior fibers, which
arise from the ceratohyal tip, extend transversely
and slightly posteriorly. The fibers form a deli-
cate fan which adheres to the dorsal surface of
the intermandibularis posterior and blends in
well with the subhyoidens. The insertion is into
the fascia of the dorsal surface of the interman-
dibularis posterior.

Subhtioidcus originates from the ventral sur-
face of the posterior tip of the ceratohyal im-
mediately medial to the origin of the interossa-
cjuadrata and the insertion blends in with the
latter before inserting into the dorsal fascia of
the intemiandibuiaris posterior.

Geniohtjoideus originates from a line on the
posterior edge of the mandible beginning at the
symphysis and extending laterally about 1.5 mm.
The fibers extend directly caudad to form two
bands of muscle Iving side bv side along the
venter of the throat. The thin fascia of the linea
alba overlaps this muscle ventrally. The inser-
tion is onto the second basibranchial cartilage
and tlu> tendonous inscription that extends lat-
erally from till' amis of the second basibranchial.

C.enio<!lossus has two parts. Tlie ventral por-
tion originates along a line on the posterior edge

Biological Series, \'ol. 16, No. 1

Hyobranchium anu Throat Myology

51

of the mandible. The Hne in this case Hes im-
mediately ventral to the origin of the geniohyoid-
eus and extends from the symphysis to a point
about 2.0 mm lateral to the symphysis. The fi-
bers extend eaudomesially to insert into the
aponeurosis lingualis and onto the otoglossal.
This is a large muscle in this species and is
thicker and broader than in the other species in-
vestigated.

The origin of the dorsal portion of the genio-
glossus is from a line along the mandible im-
mediately dorsal to the origin of the geniohyoi-
deus. The fibers spread out as they pass distally
over the throat and become almost indistinguish-
able from the fascia of the floor of the mouth. It
is into this fascia that these fibers finally insert
at a position lateral to the second radial carti-
lage. The tip of the first radial is overlapped by
the fibers of this muscle, and the medial fibers
also attach to the posterior edge of the tip of
the second radial cartilage as the latter turns
dorsad to articulate with the end of the otoglos-
sal.

Suharcualis rectus I attaches at its origin to
the dorsal edge of the posterior tip of the epi-
branchial. The fibers wrap around and enclose
the epibranchial and the lateral portion of the
ceratobranchial in a sheath. There is an opening
in the sheath which faces medially and into
which the second ceratobranchial passes on its
way to articulate with the epibranchial. The fi-
bers lie parallel with the ceratohyal and spread
out over the latter's anterior expanded end. The
insertion is along the anterior margin of the cera-
tohyal.

Rectus cervicis superficialis originates along
the lateral margin of the dorsal surface of the
sternum and along the transverse tendonous in-
scription which separates it from the rectus ab-
dominus. It passes forward as a thin, broad band
to the level of the second basibranchial, which
is partially imbedded in this muscle. The portion
of this muscle up to this point has only two ten-
donous inscriptions crossing it. The portion run-
ning anterior from the second basibranchial is
thicker and narrower than the posterior portion.
This anterior portion inserts onto the posterior
edge of the medial end of the first ceratobran-
chial.

Rectus cervicis profundus is a direct forward
continuation of the rectus abdominus, arising at
the level of the sternum and passing for\vard
alongside of the pericardium and both dorsal
and lateral to the rectus cervicis superficialis. At
the level of the second basibranchial, it is direct-
ly dorsal to the rectus cervicis superficialis. It
continues forward from this point to lie in a posi-

tion dorsal to the first basibranchial and the
hyoglossus. The two rectus cervicis profundus
muscles together are about as wide as the first
basibranchial. This muscle inserts on the poste-
rior edge of the otoglossal.

H ehosteotjpsiloideus is a narrow muscle
which arises from the dorsal surface of one of
the myosepta of the rectus cervicis profundus.
In Ambtjstoma talpoideum. this muscle origi-
nates from the second myoseptum posterior to the
second basibranchial. The fibers extend forward
dorsal to the rectus cervicis profundus and then
cross the rectus cervicis superficialis to attach to
the dorsal surface of the anterior arm of the
second basibranchial.

Ht/oglossus is a thick, well-developed muscle
which originates from the dorsal surface of the
base of the first radial and the adjacent surface
of the first basibranchial. The insertion is laterad
along the tip of the second radial.

Marbled SALAMA>a)ER.

Ambtjstoma opacum ( Gravenhorst )

Salamandra opaca Gravenhorst, Vergl. Uebers.
Zool. Syst, 1807, p. 421.

The geographical range extends along the At-
lantic Coast of the United States from Massa-
chusetts to Florida and westward to Texas, Ark-
ansas, Missouri, Indiana, Illinois, and Wisconsin.
This species is found in drier habitats than most
of the other species of the genus. It is often col-
lected in sandy areas bordering ponds and slow-
moving streams. The females are slightly larger
than the males, the total length ranging up to
120 mm. The ground color is black and is inter-
rupted by light markings in the form of trans-
verse bands. These bands are narrow dorsally
and widen as they approach the upper sides of
the body, where thev may unite with the ends
of adjacent bands to enclose a series of regular
black spots along the midline of the back. Males
have bright white transverse bands, while those
of the females are grayish in color. The body is
thick, short, and cylinder shaped. The gular
fold is prominent, and there are 12 costal
grooves if the one immediately above the fore-
leg is counted. The vomerine teeth form a trans-
verse line which is interrupted both at the mid-
line and at a point just back of the internal
nares. Most species of this genus deposit their
eggs in the .springtime in ponds or slow-moving
streams. The Marbled Salamander, however, de-
posits its eggs in the fall under logs, pieces of
bark, or other materials on the surface of the
ground. These laying sites are subsequently
flooded by the last summer or fall rains.

52

BnionAM Young Univehsity Scienc:e Bulletin

Hyobranchial Skeleton ( Fig. 1 1 )

The average amount of bone in the first basi-
branchial for the four specimens was 41 per-
cent. The basibranchial-radial articulations are
seen as definite lines. When the length of the
first basibranchial is compared with the total
length of the hyobranchial apparatus or with the
length of the first ceratobranchial, it is seen to be
greater than in any of the other species (Tables
4 and 5).

The first radials do not connect with the
anterior tip of the ceratohyals. The radials are
wholly cartilaginous and are intermediate in
length between the extremes seen in other mem-
bers of the family.

The second radials are directed in a dorso-
lateral direction while gradually tapering to a
fine point which articulates with the otoglossal.
The otoglossal is arched dorsally into the tongue
and is entirely cartilaginous.

The boneless first ceratobranchial forms a
smooth arch that continues with the epibranchial
to fomi a (juarter circle of cartilage.

The second ceratobranchial is a thin carti-
laginous rod only slightly curved and articulat-
ing on the epibranchial well posterior to the
first ceratobranchial-epibranchial articulation.

The ceratohyal is unossified except in two
specimens, in which a small ossification center
is found in the posterior tip of each of these
structures.

The second basibranchial is unossified ex-
cept in one specimen, and is shaped as an in-
verted V.

Musculature (Fig. 11)

Internumdibularis posterior originates along
the dorsomedial edge of the rami of the mandi-
ble and passes transversely over the venter of
the throat to insert into the linea alba. There is
no subdivision of this muscle into parts. The
ventral portion of the genioglossus can be seen
in the angle of the chin inasmuch as the inter-
mandibularis posterior does not completely
cover the intennandibular space at that point.
The posterior fillers extend slightly caudad and
overlap the interossacjuadrata.

IntcTlujoidcus fuisterior originates from the
fascia which is attached to the lateral aspect of
the tip of the mandible, the quadrate, the squa-
mosal, the otic capsule, and from fibers which
attach to the fascia cephalodorsalis. Tlie fibers
spread out as thev move distally and medially
until they fonn a thin sheet which inserts into
the skin of the gular fold and the midline fas-
cia. The anteriontiost tifjers are blended with

those of the interossa<|uadrata near their inser-
tion.

Depressor inandihulac originates from the
scjuamosal hone of the skull, attaching along
its entire posterolateral surface except for the
ventral 1 to 2 mm. It also arises from the fascia
cephalodorsalis and the lateral surface of the
otic capsule. It inserts onto the posterior tip of
the mandible.

Levator mandibuhie is essentially the same
in its origins and insertions as that description
which has been given previously in the section
on general musculature.

Interossuqmidrata originates on the posterior
surface of the cjuadrate and from the ventral sur-
face of the articulation between the cjuadrate
and the tip of the ceratohyal. A few fibers also
arise from the posterior tip of the mandible. The
anterior fibers extend anteromedially, while the
posterior ones are in a posteromedial direction.
It inserts into the midventral raphe of the throat.
This is a thin, fan-shaped muscle.

Sublitioidcus originates along the anteroven-
tral edge of the posterior end of the ceratohyal
and then fans out slightly as it proceeds forward
over the throat. It lies ventral to the subarcualis
rectus I and inserts into the dorsal fascia of the
intemiandibularis posterior. The latter covers
the anterior portion of this muscle ventrally. The
insertion end is displaced slightly lateral from
the subarcualis rectus I.

Geniolttjoidetis originates from the posterior
edge of the mandible along a line which begins
about 1.5 mm lateral to the symphysis and ex-
tends laterallv about 1.0 mm. The anteromedial
end of this muscle is overlapped ventrally by the
ventral portion of the genioglossus. The lateral
fibers extend directly caudad, while the medial
fibers are in a mediocaudal direction for a short
distance before continuing directly caudad. The
insertion is onto the second basibranchial carti-
lage, with the lateral fibers attaching into the
tendonous inscription of the rectus cervicis su-
perficialis.

Genio<sIossiis has a ventral portion which ori-
ginates from the ventral and posterior surfaces
of the mandible beginning at the symphysis and
extending laterallv about 2 to 3 mm. From this
area of origin, the fibers extend caudomediallv
to fomi a small triangle which overlaps the an-
teromedial end of the geniohyoideus. This ven-
tral portion is divided into two portions by a
separation in the fibers. It inserts into the apo-
neurosis lingualis. A few fibers continue along
in this ligament to attach to the first basibran-
chial and to the otoglossal.

The dorsal portion of the genioglossus origi-
nates along the posterior edge of the mandible

Biological Sehiks, Vol. 16, No. 1 Hvouhanchium and Tiihoa r Mvoi,oc:y

53

from the symphysis to a point about 1.5 mm
lateral from the symphysis. The medial fibers
insert onto the posterior edge of the lateral half
of the second radial, while the lateral fibers ex-
tend beyond this point and attach into the fas-
cia, fonning the floor of the mouth.

Suharcualis rectus I originates on the dorsal
edge of the tip of the epibranchial. The fibers
wrap around and enclose this end of the epi-
branchial in a sheath and extend forward paral-
lel with the ceratohyal. After spreading out over
the ceratohyal's expanded anterior end, these
fibers insert along the ventral edge of the ante-
rior margin of this cartilage.

Rectus cervicis superficialis originates from
the dorsal surface of the lateral half of the ster-
num. Lateral fibers originate on the tendonous
inscription which separates this muscle from the
rectus abdominus. After passing forward from
the second basibranchial, the medial fibers in-
sert onto the posterior end of the first basibran-
chial, whereas the lateral fibers insert onto the
posterior surface of the medial end of the first
ceratobranchial.

Rectus cervicis projundus constitutes a direct
forward continuation of the rectus abdominus,
arising at the level of the sternum. It is situa-
ted in a dorsolateral position with respect to the
rectus cervicis superficialis. The fibers pass ven-
tral to the second ceratobranchial and dorsal to
the first ceratobranchial. It continues forward
dorsal to the first basibranchial and the hyo-
glossus to insert onto the posterior edge of the
otoglossal.

Hehostcotipsiloidcus is a narrow muscle
which originates from a more posterior position
than in most of the other species. It arises from
the third tendonous inscription posterior to the
second basibranchial and extends anteriorly and
then medially to insert onto the dorsal aspect of
the second basibranchial.

Htjoglossus originates from the aponeurosis
lingualis and the lateral edge of the dorsal sur-
face of the anterior end of the first basibran-
chial. The fibers converge toward the insertion
to fonn a small, triangular-shaped muscle that
attaches laterally to the posterolateral edge of
the second radial cartilage as the latter curves
dorsad to join the end of the otoglossal. There
are also a few fibers which extend laterally to
attach to the fascia at the base of the tongue.

Texas Salamander.

Amhijstoma texanum (Matthes)

Salamandra texana Matthes, Allg. deutsche
naturh, Zeitschr., N. S. Vol. 1, 1855, p. 266.

The Texas Salamander is dark brown in color
with grayish, irregular, lichenlike blotches. Total
length may reach 153 mm. A slender head and
small mouth are characteristic of this species.
The vomerine teeth are very slender and are ar-
ranged in a transverse line that is entirely be-
tween the internal nares and is interrupted at
the midline. There arc usually 14 costal grooves.
It is found in Iowa, Ohio, Indiana, and Illinois
in the north and westward to Nebraska and Kan-
sas. It extends southward to Texas and Louisiana
and has also been reported from North and
South Carolina, Tennessee, Kentucky, and West
Virginia.

Hyobbanchial Skeleton (Fig. 12)

The first basibranchial is a shield-shaped
structure with a large core of bone averaging
38 percent of the total volume (Table 1). The
lines of articulation with the radials are dis-
tinct. The first basibranchial is situated poste-
riorly as compared to its position in most of the
other species. Only in A. anmdatum, A. cingula-
tum, A. maheei, and Rhyacotriton is the first
basibranchial in a more posterior position (Ta-
ble 12). This structure is wider in this species
in relation to its length than in any of the
others, except A. cingulatuin (Table 15).

The first radials are long and curved, and
they are attached to the tips of the ceratohyals
by a short ligament. Their length in relation to
the first basibranchial is exceeded only by Rhya-
cotriton, A. niaheei and Dicamptodon (Table 6).

The bases of the second radials overlap the
medial end of the first ceratobranchial ventrally.
These radials are also without ossification. They
articulate with the tips of the otoglossal, a dor-
sally curved rod of cartilage.

The first ceratobranchial is cartilage. It is a
gradually curved rod whose medial end is fused
to the first basibranchial.

The epibranchial is devoid of bone and mod-
erate in length (Table 16).

The second ceratobranchial is longer than the
first ceratobranchial and, in fact, the ratio of the
length of the second ceratobranchial (the second
branchial arch) to the length of the first bran-
chial arch ( the first ceratobranchial plus the epi-
branchial) is greater than in the others except
A. anmdatum and A. cingulatum (Table 17).
The second ceratobranchial is entirely cartila-
ginous.

In three of the five specimens, the ceratohy-
als contained ossification centers in the posterior
tips. Tlie other specimens were unossified in the
ceratohyals. This lack of consistency in ossifica-
tion between these five specimens cannot be at-

.54

Bricham '\'ounc Univeiisitv Science Bulletin

tributed to size differences since they were all
approximately equal in this respect. There may
have been an age difference between the five
specimens, although they all were collected on
the same day and in the same location, and all
five were completely transformed adults.

The second basibranchial is shaped like an
inverted V and contains cartilage on the two
tips. The average ossification is 65.1 percent.

Musculature (Fig. 12)

Internuindibularis posterior originates along
the dorsomedial edge of the mandible. The fi-
bers extend transversely to insert into a broad
midline of fascia, the linea alba of the throat. A
small area is opened at the point of the chin
through which the ventral portion of the genio-
glossus can be seen. The posterior portion of the
intermandibularis posterior overlaps the anterior
end of the interossaquadrata ventrally.

Interliijoidetis jwsterior is attached at its ori-
gin to several bonv and fascial surfaces, arising
from the posterior tip of the mandible, the later-
al surface of the quadrate, the lateral surface of
the squamosal, and from the fascia cephalodor-
salis. It inserts into the gular fold and into the
median ventral raphe which is a posterior con-
tinuation of the fascial material into which the
intemiandibularis posterior and the interossa-
quadrata insert.

Depressor mandibulae originates along the
upper squamosal, the ventral and posterior sur-
faces of the otic capsule, and posteriorly from
the fascia cephalodorsalis. The fibers of this
muscle insert onto the posterior tip of the man-
dible where they function to depress or lower
the mandible.

Levator mandibulae in Amlnistonui tcxanuni
is essentially the same as the description given
previously in the section on general musculature.

Interossaijuadrata originates along the ven-
tral surface of the articulation between the cera-
tohyal and the c|uadrate. The insertion is into
the dorsal side of the median fascia into which
the intermandibularis posterior inserts. The fi-
bers i'xtend medially to a point somewhat short
of the midline and, consequently, this muscle
overlaps ventrally the lateral half of the genio-
hyoideus.

Stibhiioideus originates along the anteroven-
tral edge of the posterior end of the ceratohyal.
It extends dorsal to the interossacjuadrata at its
origin end. The fibers are directed more medial-
ly in this species than in most of the others, and
consecjuently the insertion end overlaps the ven-
ter of the subarcualis rectus I medially rather

than laterally. It fans out to insert into the dor-
sal fascia of the intermandibularis posterior.

GenioJujoidcus originates on the posterior sur-
face of the mandible. More precisely, the ori-
gin is along a line extending laterally from the
mandibular symphysis tor a distance of 1.5 to
2.0 mm. The fibers run caudad to insert onto
the second basibranchial and into the tendon-
ous inscription of the rectus cervicis superfi-
cialis that extends laterally at that point.

Genioglossus has a ventral portion which orig-
inates along the posterior edge of the mandible
immediately ventral to the medial portion of the
origin of the geniohyoideus. The fibers extend
caudoinediallv and overlap the medial anterior
end of the geniohyoideus. They insert into the
aponeurosis lingualis at about the level of the
anterior edges of the ceratohyals, and a few fi-
bers continue in the aponeurosis lingualis to the
first basibranchial and to the otoglossal.

The dorsal portion originates along the same
line on the mandible as the ventral portion, but
the fibers extend caudolaterally. The medial fi-
bers are dorsal to the geniohyoideus, and they
insert onto the anterior edge of the second radial
near its tip. The lateral fibers insert into the fas-
cia of the tongue at its base and contribute to
the floor of the mouth.

SubarctiaUs rectus I originates on the dorsal
side of the posterior tip of the epibranchial, from
which the fibers proceed forward, wrapping
around and enclosing the epibranchial in a
sheath of muscle. These fibers continue forward
parallel with the ceratohyal to insert onto the
hitter's expanded anterior end along the anterior
margin.

Rectus cervicis supcrficialis originates on the
dorsal surface of the posterior end of the ster-
num. The fibers extend dorsal to the anterior
part of the .scapula and dorsal to the coracoids.
It lies on the ventral surface of the pericardium.
These fibers insert onto the medial end of the
first ceratobranchial just before the latter artic-
ulates with the first basibranchial. It also in-
serts onto the point of articulation of these two
cartilages and onto the posterior tip of the first
basibranchial. The medial three-fourths of this
muscle also attaches to the second basibranchial.

Rectus cervicis profundus passes forward
alongside the pericardium and is both dorsal and
iatt'ral to the ri'ctus cervicis supcrficialis. It
continues forward through the opening formed
by the two ceratobranchial cartilages and finally
inserts onto the posterior edge of the otoglossal.

1 1 ebosicotipsiloidcus originates from the dor-
sal surface of a myoseptum of the rectus cervi-
cis profundus. The particular myoseptum in this
case is the second one posterior to the second

Biological Series. Vol. 16, No. 1

HYOim.\NCIIIUM .AND TlIHO.\T MYOLOGY

55

basibranchial. Tlii.s narrow muscle extends for-
ward dorsal to the rectus cervicis profundus. At
the level of the second basibranchial, it tiuns
medially and crosses the rectus cervicis superfi-
cialis to attach along the dorsal surface of the
second basibranchial.

Hyogloss-us is a small muscle situated be-
tween the first basibranchial ventrally and the
rectus cervicis profundus dorsally. It originates
from the dorsal surface of the anterior end of
the first basibranchial and along the lateral mar-
gin of the dorsal surface of this cartilage. From
this point, the fibers extend caudolaterally to
wrap around and insert onto the posterolateral
edge of the tip of the second radial cartilage.
There are also a few fibers which continue for
a short distance into the sides of the base of the
tongue.

Chihuahua Salamander.

Ambijstovm rosaceum Taylor

Amhijstoma rosaceum Taylor, Copeia 1941 (3):
143-144, figs, la and lb. '

This species has a uniform dark brown
ground color with a lighter brown ventrally.
Along the chin and tail there are light, indistinct
spots which may also continue along the sides
of the body in a somewhat linear pattern. There
are usually 11 costal grooves, counting one each
in the axilla and groin. Vomerine teeth are di-
vided into two groups of 26 each, separated by
a midline diastema. The tail is compressed lat-
erally and has a small dorsal tail fin. There are
glands scattered over the body, becoming prom-
inent along the proximal third of the dorsal ridge
of the tail and also forming a parotid gland.
These glandular areas arc similar to but not as
well developed as in A. gracUe. Total length may
reach 165 mm. These salamanders are found in
the mountains of the Sierra Madre Occidental
from El Salto, Durango, to northern Chihuahua
and northeastern Sonora.

Hvobranchial Skeleton (Fig. 13)

The first basibranchial is shield-shaped when
viewed from the ventral aspect. In cross section,
it is a flat oval. The core of this structure is
bone, constituting an average of 28 percent of
the total volume (Table 1). The articulation
lines separating this structure from the radials
are distinct. The first basibranchial is situated
anteriorly with respect to the position of the
mandible. Only A. t. nebulosuin has a more an-
teriorly situated first basibranchial (Table 12).

The first radials are sharply curved laterally
and entirely cartilaginous. They are long, as in-

dicated by the fact that the ratio— radial I length
to basibranchial I length— is greater in this spe-
cies than in the others, with the exception of A.
mahcei, A. texanum, Rhi/utotriton, and Dicamp-
todon ( Table 6 ) .

The second radials are bent in a slightly pos-
terior direction when in a resting position. They
are wholly cartilaginous and attach to the ends
of the annular otoglossal.

No bone is found in the first ceratobranchial,
and they are rather straight, the major curve in
the first branchial arch occurring at the anterior
end of the epibranchial.

In one of the three available specimens, the
epibranchials and ceratohyals were devoid of
bone in their posterior tips. The other two speci-
mens had well-developed ossification centers in
these structures. The average ossification for the
three specimens was 36 percent for the epi-
branchials and 4.8 percent for the ceratohyals.

The ratio of ceratohyal width to ceratohyal
length is equalled in A. t. melanosticttim and ex-
ceeded in all of the others (Table 14).

The totally cartilaginous second ceratobran-
chial is long when analyzed by the ratio of
length of second ceratobranchial to length of
first ceratobranchial (Table 8).

The lateral arms of the second basibranchial
contain cartilage on their tips, the cartilage con-
stituting an average of 31 percent of the total
volume (Table 1).

The mandible length compared to the mandi-
ble width is a rather high ratio in this species,
being exceeded only by four others— Dicampto-
(ion, A. annulatttm, A. nuibeei, A. inaculatum,
and A. cingulattnn (Table 9).

Musculature (Fig. 13)

Intermandibularis posterior originates on the
dorsomedial surface of the mandible and ex-
tends medially to the linea alba. It is seen im-
mediately under the skin of the throat and over-
laps the more dorsally situated interossaquadrata
and subhyoideus muscles. The posterior fibers
are directed oblifjuely caudomediad, while the
majority of the filjers are transverse.

Intcrluioideus posterior originates from fas-
cia which is in turn attached at several points,
the lateral aspect of the posterior tip of the
mandible, the quadrate, the squamosal, the otic
capsule and the fascia cephalodorsalis. The fi-
bers spread out as they extend in a caudomesial
direction to insert into the gular fold and the
inidventral fascia.

Depressor mamlihulae originates on the dor-
sal end of the scjuamosal, the ventral and poste-
rior surfaces of the ear capsule and along the

56

Bhigham Young Univkrsity Science Bulletin

fascia cephalodorsalis. It inserts onto the poste-
rior tip of the mandible.

Levator iiumdibulac is essentially the same as
the description which is given in the earlier sec-
tion on general musculature.

Interossacjuadrata originates on the ventral
surface of the cjuadrate, anterior to the hyoquad-
rate ligament and from the hyoquadrate liga-
ment itself. Tliis is a delicate muscle which is
easily lifted along with the intennandibularis
posterior. It is fan-shaped, the anterior fibers ly-
ing in an antcromesial direction and the poste-
rior fibers in a posteromesial position. It inserts
into the median raphe into which the intennan-
dibularis posterior and the interhyoideus poste-
rior also insert.

SubJiyoideus originates from the ventrolateral
margin of the posterior end of the ceratohyal.
Tlie posterior fibers of this thin muscle blend in
with those of the interossaquadrata, making sep-
aration of these two muscles difficult at their in-
sertion ends. This is also a delicate muscle that
is easilv disrupted when the intennandibularis
posterior is lifted. The insertion end overlaps
laterally part of the ventral surface of the sub-
arcualis rectus I. It inserts into the dorsal sur-
face of the fascia of the intermandibularis pos-
terior.

Geniohijokleiis originates from a short line
on the posterior surface of the mandible. This
line runs laterally for about 3.0 mm from a point
about 1.5 mm lateral to the symphysis. The fi-
bers extend caudad to form a band of muscle
which inserts onto the second basibranchial and
onto the tendonous inscription lateral to the sec-
ond basibranchial. This muscle appears to be
rather narrow in this species.

Genioglossus originates along a line begin-
ning at the mandibular symphysis and extends
laterally about 2.0 mm along the posterior sur-
face of the mandible. Tlie fibers extend caudo-
mesiad to form a small triangle. The insertion is
into the aponeurosis lingualis, and a few fibers
continue on to the anterior edge of the first basi-
branchial. A few fibers also attach to the oto-
glossal cartilage.

The broader lateral portion originates im-
mediately dorsal to the origin of the ventral por-
tion but extends laterally from the symphysis for
a distance of 3 to 4 mm. The fibers fan out and
extend posterolaterally to insert into the fascia
of the floor of the mouth at about the level of
the second radial tip.

Subarciudls rectus I originates along the dor-
sal edge of the posterior end of the epibranchial,
wraps around this cartilage, and extends forward
over the ventral surface of the ceratohyal. The

fibers fan out and insert along the anterior mar-
gin of this cartilage.

Rectus cervici.s supcrficialis originates on the
dorsal surface of the sternum and the tendonous
inscription extending laterally from the sternum.
It extends dorsal to the coracoid cartilages and
ventral to the pericardium. The second basi-
branchial is partially enveloped by this muscle
as it continues anteriorly. It then becomes nar-
rower and thicker and turns medially to insert
onto the posterior tip of the first basibranchial
and onto the posterior edge of the medial end
of the first ceratobranchial.

Rectus cervicis profundus appears at the level
of the sternum as a flat band extending forward
alongside and dorsal to the rectus cervicis su-
pcrficialis. At the level of the omohyoideus, it
begins to turn medially. Forward from the sec-
ond basibranchial, it is a thick, narrow band
which lies dorsal to the rectus cervicis supcrfi-
cialis. The fibers pass ventral to the second cera-
tobranchial and dorsal to the first ceratobran-
chial. From this point forward, it leaves the rec-
tus cervicis supcrficialis and inserts onto the
posterior edge of the otoglossal.

Hebosteoijpsiloideus originates from one of
the myosepta of the rectus cervicis profundus.
The particular myoseptum is the second one
posterior to the second basibranchial. It is a nar-
row muscle which at first extends directly for-
ward dorsal to the rectus cervicis profundus.
After a short distance, it turns across the dorsal
surface of the rectus cervicis supcrficialis and
inserts onto tlic dorsum oi tlie second basibran-
chial.

Hi/oglossus is a small muscle extending from
the first basibranchial to the tip of the second
radial. It is triangular in shape, with the base of
the triangle situated along the dorsal surface of
the anterior end of the first basibranchial and
along the lateral margiii of this cartilage to the
base of the second radial. The point of the tri-
angle is the insertion end which attaches onto
the tip of the second radial and out into the ad-
joining fascia.

Tiger Salamander.
Ambystoma trigrinum Green

^(davuiudra tigrina Green, 1825, Jour. Acad. Nat.
Sci. Phil., (1), 5:116.

Eastern Tiger Salamander.

Ambijstonui tigrinum tigrinum (Green)

Salamundra tigrina Green, Journ. Ac. Nat. Sci.
Phila., Vol. 5, 182.5, p. 116.

Hioi.iuacAi. Sehies. \'()l. 16. No, 1

HVODHANCIIIUM AND TllliOAl \Iv<>l-(><:^

57

This subspecies ranges extensively through-
out most of the United States, from New York to
northern Florida and westward to Texas in the
south and Minnesota and parts of Ontario in
the north. It is a burrower and spends most of
the vear underground. Its burrowing habit is
abandoned onlv during the breeding season,
during which time it can be found in ponds and
slow streams. This is the largest species of the
genus Ambystoma, attaining a length of 254 mm
in some individuals. It is a stout-bodied salaman-
der with 12 costal grooves when those in the
axilla and groin are counted. The gular fold is
prominent, and the vomerine teeth are usually in
a continuous series, although there is occasional-
Iv a break in this series at the midline. Males
are usuallv larger than females, and the tail and
hind legs are also usually longer in male speci-
mens. Ground color is dark brown or dull black.
The venter is olive yellow in color. Brownish
oli\e or brownish yellow blotches are scattered
over the dark ground color. These blotches may
fuse somewhat between the costal grooves to
form short bands along the sides of the trunk.

This subspecies of tiger salamander is char-
acterized by having black spots on a dark
ground color of olive green or dark gray. It has
a well developed gular fold and usually 13 cos-
tal grooves when the one in the axilla and the
two that run together in the groin are counted.
The line formed by the vomerine teeth tends to
arch forward between the internal nares and
may or may not form a continuous series. When
not continuous, the tooth line will be interrupted
just medial to the inner margin of the naris and
also at the midline. Males tend to be slightly
larger than females. Total length may reach 230
mm. During the breeding season the swollen
vent is characteristic of the male. These sala-
manders are found commonlv in the mountain
lakes and ponds and also occasionally beneath
logs and stones. Their range extends over west-
ern Colorado, northwestern New Mexico, north-
em Arizona, the inner basin of Utah, and the
Colorado Plateau in Utah.

The anatomical descriptions which follow are
based upon a study of the above three subspe-
cies of Tiger Salamander.

Northwestern Tiger Salamander.

Ambystoma tigrinum melanostictum (Baird)

Sircdon lichenoides melanostictum Baird, 1860
Expl. Surv. R. R. Miss. Pacific, 12, pt. 2, p. 306.

This subspecies is characterized by extreme
development of the light colored areas of the
skin. These areas converge greatly sucii that the
darker ground color persists as irregular patches.
The light areas are pale yellow and the ground
color is brown to black. There are usually 13
costal grooves when one in the axilla and two
that converge in the groin are counted. There
may be three series of vomerine teeth which
form a forward arching line in front of the in-
ternal nares; however, the central series is oc-
casionally interrupted at the midline. Adults may
reach a total length of 219 mm. Adult specimens
are sometimes found in damp situations under
logs or stones; whereas, during the breeding sea-
son, they may be abundant in the mountain
lakes and ponds throughout their range. They
are found in British Columbia, Alberta, Wash-
ington. Oregon, Idaho, Montana, Wvoming,
North Dakota, South Dakota, and Nebraska.

Clouded Tiger Salamander.

Ambystoma. tigrinum nebulo.mm Hallowell

Ambystoimi nebnlosum Hallowell, Proc. Acad.
Nat. "Sci. Phila., 6, p. 209.

Hyobranchial Skeleton (Fig. 14-16)

The first basibranchial i.s shield shaped and
has a central core of bone. The percentages of
ossification of the first basibranchial are similar
for A. t. melanostictum and A. t. tigrinum
(Table 1).

The relative position of the first basibranchial
along the anteroposterior axis of the throat is
verv similar for these three subspecies and, as a
group, their first basibranchials are more ante-
rior than thev are in the other species (Table
12).

The first radials attach to the tips of the cera-
tohyals by a short ligament. These radials are
moderate in length and unossificd.

The second radials also contain no bone and
are curved dorsally where their distal ends artic-
ulate with the ends of the otoglossal. The oto-
irlossal is unossificd and ;irched dorsallv into the
tongue.

The first ceratobranchials are entirelv carti-
laginous and moderate in length. They are uni-
fonn in width and thickness throughout their
length and are flattened dorsoventrally.

The epibranchials were ossified in all 10 of
the specimens examined. The amount of ossifi-
cation was notably different, however, in A. t.
ncbulosum from that found in the other two sub-
species (Table 1). This, however, is based on a
limited number of specimens.

The average first ceratobranehial length,
when compared to the length of the epibran-

58

Bhigham Y(>iiN(; University Science Bulletin

chial, was shorter for A. t. melanostictum and A.
t. nebulo.nun than all of the others except A.
talpoideum (Table 13).

The second ceratobranchial compared to the
first ceratobranchial has a smaller ratio in the
f!>rtnu;«-groiip than in other species except for
Dicamptoclon and Rlujacotriton (Table 8).

Two of the subspecies were found to have
ossification centers in the ceratohyal tips. The
third, A. t. melanostictum, has no bone in the
ceratohyals. The average percent ossification of
the ceratohyals in A. t. nebulosum and A. t.
ti^rimim was 8 and 5 respectively (Table 1).
The ceratohyal length in relation to the total
length of the hyobranchium is shorter in A. t.
tigrinum than in the other two tiger salamander
subspecies (Table 19). This ratio is moderate
for the tigrintim group as a whole.

The second basibranchial in the tigrimim
group is triradiate in shape, with the anterior
ann being considerably shorter than the lateral
anns. The anteroposterior length of this element
is greater in A. t. tigrimim in relation to the side-
to-side length than in the other two subspecies
(Table 18). This ratio is also moderate com-
pared with all the other species.

As a group, the tigrimim subspecies have a
broad head compared with the other species.
The ratio— mandible length to mandible width-
is smaller in A. tigrimim than in the others ex-
cept for the mole salamander, A. talpoideum
(Table 9).

MUSCUL.A.TUHE (Fig. 14-16)

Intermamlibiilari.s posterior originates on the
dorsomedial surface of the mandible. The direc-
tion of the fibers is transverse for the most part,
except for the posterior fibers which are directed
caudomedially. It inserts into the linea alba of
the throat.

Interhyoideus ])osterior originates from fas-
cia which is attached to the lateral part of the
posterior tip of the mandible, the (juadrate, the
s(|uamosal, and the otic capsule, with some pos-
terior fibers originating from the fascia cephalo-
dorsalis. It inserts into the skin and fascia of the
gular fold and into the midventral line. The an-
terior fibers blend imperceptibly with those of
the interossaquadrata. The posterior edge of this
muscle (the gular fold) is more anteriorly situ-
ated in A. t. mdndosum than in anv of the other
specimens included in this study (Table 11).

Depressor numdibulae attaches along the dor-
sal end of the s([uamosal, the ventral and pos-
terior surfaces of the otic capsule, and in the
fascia cephalodorsalis. The insertion is onto the
posterior tip of the mandible.

Levator mandibtdae is essentially the same as
the description given in the earlier section on
general musculature.

Interossaquadrata originates from the ventral
surface of the (juadrate anterior to the hyoquad-
rate ligament and from the ligament itself. The
anterior fibers extend anteromediad and ventral
to the subarcualis rectus I, whereas the posterior
fibers lie parallel with those of the interhyoideus
posterior. The insertion is in the median raphe
into which the intemiandibularis posterior and
interhyoideus posterior also insert.

Subhyoideus originates on the ventrolateral
margin of the posterior 2 to 3 mm of the cerato-
hyal. It is a thin muscle, the posterior fibers of
which blend in closely with those of the interos-
sa(juadiata. The fibers lie parallel with the sub-
arcualis rectus I; and, in fact, if the intemiandib-
ularis posterior is removed carefully, the sub-
hyoideus can be seen lying against the ventral
surface of the subarcualis rectus I. The insertion
end overlaps the venter of the subarcualis rectus
I laterally. This muscle inserts into the dorsal
fascia of the intemiandibularis posterior.

Geniolnjoideus originates from a short line
along the posterior surface of the mandible. This
line begins at a point about 1..5 mm lateral to
the symphysis and extends laterally about 3.0
mm. The fibers pass caudad and slightly mediad
to insert onto the second basibranchial. A few of
the lateral fibers also insert onto the tendonous
inscription of the rectus cervicis superficialis that
extends laterally from the amis of the second
basibranchial.

Genioglossiis has a ventral portion which ori-
ginates along a line beginning at the mandibular
symphysis and extending laterally about 2.0 mm.
The fibers extend caudomedially and form a
small triangle that is seen between the anterior
ends of the geniohyoideus. The insertion is into
the aponeurosis lingualis which connects the
root of the tongue with the first basibranchial
and with the otoglossal.

The lateral portion is a broader muscle which
originates along a line immediatelv dorsal to the
origin of the ventral portion but extends laterally
Irom the symphysis for a distance of 3 to 4 mm.
These fibers extend posterolaterally to insert in-
to the base of the tongue. The medial fibers at-
tach to the tips of the second radials.

Snbarctialis rectus I originates from the dor-
sal side of the posterior end of the epibranchial
cartilage. The fibers radiate around and enclose
this end of the epibranchial, forming a sheath at
the posterior end of the muscle with the opening
of the sheath facing mediallv and into which the
ceratobranchial passes. The fibers lie parallel
with the cartilage and insert on the ventroante-

Bioi.o(:ic:ai. Skhies. \'in,. 1(>. No. 1

IlvoHliAN'ClllUM AND TllHOAl Mvi)l.(K;V

59

rior border of the expanded anterior end ot the
ceratohval.

Rectus cervicis sujicrficialis originates from
the posterior two-thirds of the dorsal surface of
the sternum. The fibers of this muscle extend
fonvard dorsal to the sternum, dorsal to the cor-
aeoids, and ventral to the pericardium to insert
onto the dorsal surface of the second basibran-
chial and into the tcndonous inscription from
which the geniohyoideus arises. The part ante-
rior to the second basibranchial inserts onto the
posterior tip of the first basibranchial and the
posterior edge of the medial end of the first
ceratobranchial cartilage.

Rectus cervicis profundus is a direct forward
continuation of the rectus abdominus. .\t the
level of the sternum, it passes forward along the
side of the pericardium and both dorsal and lat-
eral to the rectus cervicis superficialis. It is ven-
tral to the second ceratobranchial and dorsal to
the first ceratobranchial and inserts finallv on

the posterior edge ot the otoglossal cartilage.

Hehosteotjpsiloideus is a narrow muscle orig-
inating from the dorsal surface of one of the
nivosepta of the rectus cervicis profundus. In
this species it arises from the second myosep-
tum posterior to the second basibranchial, ex-
tends dorsal to the rectus cervicis profundus,
turns medially for a short distance dorsal to the
rectus cervicis superficialis, and inserts on the
dorsal surface of the second basibranchial.

Uijopjossus is a small muscle which lies ven-
tral to the rectus cervicis profundus and origi-
nates from the dorsal surface of the anterior end
of the first basibranchial cartilage and along the
lateral margin of the dorsal surface of this carti-
lage. Tlie fibers extend caudolaterally to wrap
around the tip of the second radial cartilage.
The point of insertion is on the posterolateral
edge of the second radial, although a few of the
posterior fibers extend laterally to insert into
the sides of the base of the tongue.

DISCUSSION

The foregoing descriptions of comparative
morphology have been condensed to 48 charac-
ters which we have used in a cluster analysis us-
ing the method of Wishart (1969). We believe
that the use of this method, together with the
methods of descriptive morphology, provides for
an analysis of the data which is more precise
than is possible by use of descriptive morphol-
ogy alone.

A character analysis produced bv statistical
methods concerning relationships between the
various species of the Ambystomidae provides
useful information for the deduction of a phylog-
env. It must be kept in mind, however, that the
basis for these relationships, as used in this
studv, is comparative morphology of the hyo-
branchium and throat musculature. It is entirely
possible that these results could be different us-
ing infomiation derived from different sources,
such as the skull or external morphology.

It is not known whether the individual char-
acters are evolving as independent units or
whether they are parts of larger integrated sys-
tems. We suspect that some are related to the
development of the feeding and/or breathing
mechanisms.

The characters used in the cluster analysis
are of tvvo types— numeric and binarv. The for-
mer are recorded as the length of a particular
structure, the ratio of lengths of two different
structures, or as a percentage of ossification of a
structure. Binary characters are expressed as

either the presence or absence of a particular
structure, and are recorded in the data as 0 or 1
for absence and presence, respectively.

The numbered paragraphs which follow con-
tain explanations of each of the 48 characters.
Some of the binary characters contain two or
three different attributes so that the total num-
ber of attributes amounts to 63 ( 22 numeric and
41 binarv). The number of each paragraph cor-
responds to the character and/or attribute num-
ber. (See Tables 2 and .3 for the specific charac-
ters for each individual. )

1. The ratio of the distance between the
mental symphysis and the posterior edge of
the intermandibularis posterior to the dis-
tance between the mental symphysis and the
anterior edge of the pectoralis.

2. The ratio of the distance between the
mental symphysis and the gular fold to the
distance between the mental symphysis and
the anterior edge of the pectoralis.

3. The ratio of mandible length to mandi-
ble width.

4. The ratio of the distance between the
mental symphysis and the anterior edge of
the first basibranchial to the length of the
mandible.

5. This ratio is a comparison of the length
of the first basibranchial with the total length
of the hyobranchial apparatus. The basibran-

60

Biur.iiAM Young Univehsitv Science Bulletin

chial length is taken as the distance between
the anterior edge of the first basibranchial
and a hne drawn between the posterior edges
of the medial ends of the first two cerato-
branchials. The latter point was chosen
rather than the extreme posterior tip of the
first basibrancliial l^ecause the articulation
between the first basibranchial and the sec-
ond ceratobranchial was often indistinct.

6. The ratio of the length of the first basi-
branchial to the length of the first cerato-
branchial.

7. The ratio of the length of the first basi-
branchial to the total length of the first bran-
chial arch.

8. This ratio compares the width of the
first basibranchial to its length.

9. The ratio of the length of the first ra-
dial to the length of the first basibranchial.

10. This character is expressed as the ra-
tio of the length of the first ceratobranchial
to the total length of the first branchial arch.

11. The ratio of the length of the first
ceratobranchial to the length of the epibran-
chial.

12. The ratio of the length of the second
ceratobranchial to the first ceratobranchial.

13. The ratio of the length of the second
ceratobranchial to the length of the first
branchial arch.

14. The ratio ot tlie length of the hyo-
branchium to the distance between the men-
tal symphysis and tlic sternum.

15. The ratio of the anteroposterior length
of tlie second basibranchial to its side-to-side
width.

16. The ratio of the length of the cerato-
hyal to the length of the entire hyobran-
chium.

17. The ratio of the width of the cera-
tohyal to its length.

18-22. These characters are the percent-
ages of ossification of the first basibranchial,
second basibranchial, first ceratobranchial,
ceratohval, and epibranchial. respectively.

23. This binary character is the absence
or presence of a definite line of articulation
between the first radial and the first basi-
branchial.

24. The absence or presence ot a definite
line of articulation between the second radial
and the first basibranchial.

25-27. This is a binary character with
three different attributes. Each first basibran-
chial either is or is not truncate, pear shaped,

or shield shaped. Column 25 on Table 3 indi-
cates whether or not it is truncate; column 26
indicates whether or not it is pear shaped;
and column 27 indicates whether or not it
is shield shaped,

28. The absence or presence of a joint
midway along the first radial cartilage.

29. A binary character based on whether
or not the first radial tip attaches to the cera-
tohyal.

.30-31. This binary character has two at-
tributes—whether the otoglossal is or is not
annular in shape and whether it is or is not
platelike in shape.

32. This character is whether the otoglos-
sal is lightly ossified, as in Dicamptodon, or
completely cartilaginous. The ossified condi-
tion is designated with a 1 on Table 3, and
the cartilaginous condition is designated with
a 0.

.33. This binary character is whether or
not the otoglossal is fused with the second
radials.

34-35. The posterior tip of the second
ceratobranchial articulates with the first bran-
chial arch at a point either directly on the
joint between the first ceratobranchial and
the epibranchial or at a point slightly poste-
rior to that point. The first attribute of this
character is whether or not the point of artic-
ulation is posterior to the joint. The second
attribute is whether or not it is directly on
the joint.

■36-38. This binary character lias three at-
tributes. The second basibranchial either is or
is not triradiate, it either is or is not 4-radi-
ate, and it either is or is not an inverted V.

39. This is a binary character concerned
with hypertrophy or lack of hypertrophy of
the levator mandibulae.

40-42. This character has three attributes
concerned with the insertion of the rectus
cervicis superficialis— whether this muscle
does or does not insert only on tlie first basi-
branchial, whether it does or does not insert
only on the first ceratobranchial, and whether
it does or does not insert on both the first
basibranchial and the first ceratobranchial.

43-45. The hebosteoypsiloideus originates
on one of the three myosepta posterior to the
second basibranchial. These three attributes
are whether or not this muscle originates on
the first, second, or third myoseptum.

46. This character is the absence or pres-
ence of the hyoglossus.

UiOLOciCAL SiJAiEs. Vol. 16, No. 1

HvollH.^^'^HIU^t .xnd Tiihoai Myology

61

47. This character is whether or not there
is an opening in the intennandibularis poste-
rior at the point of the chin.

48. This is a binary character concerned
with the width of the linea alba. If it is a thin
line rnnning down the midline of the throat,
it is designated with a 1. If it is wider than
this, it is designated with a 0.

49-.50. This character deals with the num-
ber of inscriptions on the rectus cervicis su-
perficialis. Each animal has either two in-
scriptions or three inscriptions.

51-52. The insertion of the rectus cervicis
profundus is either fleshy or tendonous. At-
tribute 51 is either fleshy or not fleshy, and
attribute 52 is either tendonous or not ten-
donous.

53-54. This character is whether the joint
between the first ceratobranchial and the epi-
branchial is distinct or indistinct.

55. A binary character concerned with
whether or not the anterior end of the genio-
hvoideus is overlapped ventrally by the ge-
nioglossus.

56. This character is whether or not the
intermandibularis posterior is divided.

57. This is the presence or absence of
parotid glands and a glandular ridge on the
tail.

58-59. This character has two attributes—
either the ventral edge of the tail is rounded
or it is thin.

60. This is whether the genioglossus does
or does not insert on the first ceratobranchial.

61. This character is whether the sub-
hyoideus inserts on the mandible or not.

62-63. In Dicamptodon the majority of the
hyobranchium is uniformly, but lightly, ossi-
fied or mineralized. In all of the other spe-
cies the majority of the elements of this struc-
ture are cartilage. This character is whether
the hyobranchium is like that of Dicampto-
don, or whether it is mostly cartilaginous.

The numeric characters were tested for
homogeneity of variance using Bartlett's Test
(Dixon and Massey, 1969). Unfortunately,
there was for each character in the cluster analy-
sis at least one species in which the variance was
great enough to cause heterogeneitv. For this
reason the numeric characters were of little use
in the deduction of a phylogeny. In other words,
the variation between individuals of a single
species with reference to a given numeric char-
acter was so excessive that the use of this char-
acter to separate species was invalid. There is

considerable individual variation in the lengths
and widths of the separate components of the
hyobranchial skeleton. Other workers have used
a few of these dimensions for studies at the
generic level (Ozeti and Wake, 1969); however,
this study has shown that they are of little use in
separating species or subspecies.

There were 26 binary characters used in the
cluster analysis. The dendrograms for this analy-
sis are shown in Figures 18 and 19. A brief dis-
cussion and summarization of the results of the
binary characters is as follows:

Dicamptodon remains separate from the
others in this analysis. It is the only species with
a segmented radial, a platelike otoglossal, a
fused otoglossal-second radial, no hyoglossus, no
opening in the intermandibularis posterior at the
chin, and a uniformly, but lightly, ossified hyo-
branchium.

Rhifacotriton has a totally cartilaginous hyo-
branchial skeleton. This is probably an advanced
condition associated with terrestrial feeding hab-
its (Regal, 1966). A cartilaginous hyobranchium
is more flexible than a bony one, and in the act
of feeding is a more pliable part of the tongue
apparatus enabling the tongue to be extended
with more speed and precision than could be
done with a hyobranchium of bone. Bony
hvobranchial elements are characteristic of prim-
itive aquatic salamanders.

Another advanced trait found in Rlujacotriton
is the reduced number of myosepta on the rec-
tus cervicis superficialis. The number is two in
this case as compared to the usual three. A trait
peculiar to RJitjacotriton is the long subhyoid-
eus. In all of the other .species investigated this
muscle is attached to the anterior end of the cera-
tohyal, whereas in Rlujacotriton it extends for-
ward beyond that point to reach the mandible.
In most of the others, the subhyoideus is so re-
duced as to be difficult to detect upon dissec-
tion. It has been completely overlooked by some
anatomists, as pointed out alx)ve. The genioglos-
sus is also longer than in other species, extend-
ing as far posterior as the first ceratobranchial.

The affinity between Dicamptodon and
Rhifacotriton which has been suggested by nu-
merous workers is based on data other than
throat morphology. Dunn ( 1940) says that these
two genera are related to each other and differ
from Andji/stoma in their skull features. Tihen
(19.58) suggested, on the other hand, that Di-
camptodon and RJitjacotriton, based on the oste-
ology of their skulls, were derived independently
from the proto-ambystomatid stock and that they
are different from each other and from the re-
mainder of the family. Noble ( 1927), in attempt-
ing to support the idea that Dicamptodon and

62

A. gracile

A. jeffersotiiauurn

A. mabcei

A. mucnulartijhnn

A. inaitiluttiiii

A. lutpoiiU'iiin

A. ojKicinn

A. tcxuiiiiiii

A. rosdccuiii

A. tigrinuDi

A. iniiitiliituiii
A. ciiiiiuldluin

0 .1

BiiiciiAM Yoi'Nx; UNivKitsiTV Science
.7 .8 .9 1.(1 1.2 1..3 1.4 1..T l.fi

Bulletin
.3.fj

Di

-7^

-y^

Fig. 18. Deiulrogram for tlu' (lenus AnihystDinii.

Rlitjcicotriton are related, has pointed to the fact
that these two genera have stream-type larvae
in contrast to the pond-type larvae of Ambys-
toma. We can add to this the fact that the
adults of these two stream-type genera have
hypertrophied levator mandibulae muscles which
are apparently of great value in grasping and
swallowing the elusive types of food items pres-
ent in a fast-moving stream. It is also note-
worthy that in the mouth of one of the Rlujaco-
Iriton specimens was found a winged insect
the size of a small bee. Furthermore, the "eo-
graphical distributions are basically the same.
They are found in the Pacific Northwest and
nowhere else. In addition, the long subhyoideus
in Rhijacotriton is a primitive trait. In spite of
this, however, our studies have verified the fact
that these two genera are different from each
other and from the genus Ambtjstonia. Not only
is Rhijacotriton distinct from Amhijstonui, but it
is also different from Diaimptodon in its wholly
cartilaginous hyobranchium, fewer transverse in-
scriptions in the rectus cervicis muscles, unseg-
mented radials, annular otoglossal, and presence
of the hyoglossus. We believe, therefore, that
there is a preponderance of evidence for an in-
dependent derivation of DUiimptodon and RJnj-
acotriton from the proto-ambystomatid stock.

A. (iniuiJattim and A. ciniitthituiii consistently
remaiiKH.! togetlK>r throughout the analyses. Tiiev
share several characters. They are the only spe-
cies in which the second basibranchial is 4-ra-

diate and they alone have tendonous insertions
of the rectus cervicis profundus. They and Di-
camptodon are the only species in which the
first ceratobranchial-epibranchial point is indis-
tinct. It is interesting to note that C^ope (1887)
considered A. anntthittun and A. leptttrum (cin-
^uhiturn) to be a distinct and separate group
from the other ambvstomids.

As expected, the tigrinum group remained to-
gether, but it is of interest that A. rosaceum is
aligned closely with them. Tliis is a confinnation
of the idea which others have expressed con-
cerning the close relationship between A. rosa-
cctiin and A. tig,rinuin (Anderson, 1961). We
definitely believe that rosaceum separated early
from the ti<irimtm group and became a mountain
type of tiger salamander. With the dispersion
after the recent ice age salamanders of the tigri-
mini group (or their ancestral stock) in the
southern areas, particularlv those of the Rio
Concha and other smaller drainage basins flow-
ing into the Rio Grande, moved up the streams
to become mountain inhabitants. Such a dispers-
al produced mountain islands of isolation with
wide desert flats to insure the separation. Ambys-
tonui rosaceum is also similar to some of the
subspecies of tigrinum in its basic habitus, color
(light spots or with light mottling as seen in
specimens from the border states) and is a bur-
rower. On till' basis of our data this salamander
should perhaps be realigned as a subspecies of
A. tiiirinum.

JilOLOGlCAL SeHIES, VoL. 16, No. 1 HVDBHANCHIUM AND TllHOA T MYOLOGY

63

Dicamptudon

A. i^nicilc

A. jcffcrsonUimim

A. mtihcci

A. macrodactijluTii

A. mucuhitttnt

A. tutjioidctim

A. OfHicum

A. Icxunuin

A. rosaceum

A. tigrinum

A. (innulutinn

A. cinnulutum

Rhijucotriton

Fig. 19. Pos.sible phylogeny for the Ambystomidae of the United States and northern Me.xico.

A. opactim and A. texanum show a closer re-
lationship to each other on the basis of their
throat anatomy than they do to other members
of the family. They differ from each other in
only tvvo characters, and for the one character
( the shape of the second basibranchial ) they are
different from all of the others. The nearest as-
sociation of the opacum-taxanwn group is with
the wsaceiim-tigrimtm group at .991 (Fig. 18).

A. jeffer.wnianum, A. mabeei, and A. nuicro-
dactylum differ in none of the binary characters.
However, A. nuicrodactylum can be separated
from A. jeffersonianum and A. mabeei by the
fact that it contains no ossification in the epi-
branchial tips in fully transformed adults. A. jef-
fersonianum and A. mabeei can be separated by
the shapes of their otoglossal cartilages (Fig. 6
and 7). A. maculatum differs from the others in
the width of the linea alba. A. gracile is associa-
ted with this same cluster, differing from them
in the shape of the first basibranchial and in
having a divided intennandibularis posterior.

A. talpoideum is aligned more closely with
the gracile - jeffersoniantim - mabeei -mac r o-
dacti/Jum-macuIatum group than with the Oj)a-
cum-texamim-tigrintim group. This is based pri-
marily on the shape of the first basibranchial.
The former group, like A. talpoideu7n, all have
pear-shaped first basibranchials, whereas this
structure in the latter group is shield shaped.

When the binary characters concerned with
muscles are masked from the analysis, the align-
ment of the species is unchanged from that de-

scribed above. This then, represents an arrange-
ment based on the hyobranchial skeleton. The
hyobranchial skeleton characters were also mask-
ed, and the analysis produced thereby is one
based on information concerning muscles. In this
case A. opacuin and A. texamtm are more widely
separated from each other than in the former
method, and A. rosaceum becomes more widely
separated from the tigrinum group. Otherwise,
the cluster arrangement is the same. Such con-
firming data derived from t\vo anatomical
sources add considerable weight to a phyloge-
netic conclusion and seems to us to be of utmost
importance.

There is considerable similarity between the
alignment of the species of the genus Ambys-
toma as presented here and that suggested by
Tihen ( 1958 ) . There is agreement in the place-
ment of A. rosaceum in the tigrinum group, and
in the placement of A. gracile, A. jeffersonianuvi,
A. macrodacttjlum, and A. nuiculatum in the 7nac-
idaluin group. He has also aligned A. annula-
tuni and A. cingulatum. We disagree in the
placement of A. ialpoideum, A. mabeei, and A.
texanum (compare Figure 18 with Tihen's
groups in the introduction).

Tihen's classification is based on skull mor-
phology, whereas ours is based on the hyobran-
chial apparatus. Inasmuch as the hyobranchial
apparatus is more directly involved with those
anatomical structures asociated with the proces-
ses of breathing and feeding than is the skull, this
apparatus provides, we believe, a more sensitive

64

BiiiGiiAM VouNc; University Science Bulletin

indicator of the evolution tliat has occurred in
tliese salamanders.

Some interesting evidence with regard to in-
terfamilial relationships has come to light in the
course of this investigation. Most workers are of
the opinion that the ainhystomids gave rise to
the salamandrids and that the salaniandrids in
turn gave rise to the plethodontids. Regal ( 1966),
in considering feeding mechanisms, affirms that
the free-tongue condition seen in the plethodon-
tids is an advanced condition and that the tongue
of the Salamandridae, which is attached and
less protrusible, is a correspondingly less ad-
vanced condition. He further concludes that in
primitive salamanders the tongue, being firmly
attached at its base and along its sides, is not
capable of extension outside of the mouth cavity.
This latter type is an aquatic structure— a "water
tongue"— whereas the free-tongue condition is
found in terrestrial salamanders which feed upon
more elusive prey. For the most part the Ambys-
tomidae have a tongue which is attached in a
primitive manner. The flexible hyobranchium of
Rhijacotriton, however, enables this advanced
ambystomid to utilize terrestrial feeding more so
than it would appear to occur in most of the
other members of the family. The long subhyoid-
eus and genioglossus muscles have assumed an
important role in propelling the tongue forward.

The epibranchial cartilage was found in all
of the ambystomids disected. The absence of this
cartilage in some of the salamandrids would ap-
pear to be evidence for the advanced nature of
the Salamandridae with respect to the Ambys-
tomidae.

Further evidence for this relationship is
found in the fact that there were two pair of
radials present in all of the ambystomid speci-
mens. Some salamandrids have only one pair,
and Bogoljusky (1924) has confinn'ed that the
two-pair condition is primitive. The otoglossal
cartilage was also present in all of the ambysto-
mid specimens, whereas it is lacking in both
Salamandridae and Plethodontidae (Dunn,
1926). Figure 17 illustrates those changes which
have occurred in the otoglossal and the radials.
The primitive ambystomid, Diccnnptodon, has a
platelike otoglossal which is fused to the second
radials. The more advanced condition in Ambvs-
tomidae consists of an annular otoglossal which
articulates with the tips of the second radials.
The salamandrid condition shows complete loss
of the otoglossal and in some genera a loss also
of the second radial. The otoglossal is absent in
Pli'thodontidae as illustrated in Figure 17, and
there is likewise a noticeable reduction in the
radial cartilages. As the tongue has become more
flexible and freed from the floor of the mouth in

advanced salamanders, it would appear that
tiiere has been a tendency toward reduction and
even loss of the otoglossal as well as a rearrange-
ment of teeth over the roof of the mouth. The
rearrangement consists of a shifting of the teeth
postcriorlv to lie direct!\' above the tongue pad
rather than more anteriorly to correspond in posi-
tion above the otoglossal as in the ambystomids.
In the more terrestrial salamandrids and pletho-
dontids, the tongue has become increasingly
freed from the floor of the mouth and functions
more effectively in the capture of insects. Cap-
tured insects are manipulated bv the tongue and
forced up against the numerous teeth located on
the roof of the mouth. Ambystomids, on the
other hand, have a much less flexible tongue,
and have retained the otoglossal cartilage appar-
entlv as an aid in forcing food up against the
vomerine teeth, which fonn a transverse row
directlv opposite the otoglossal.

The epibranchial is longer with respect to the
ceratobranchial in Salamandridae (Ozeti and
Wake, 1969) than in the Ambystomidae studied.
Tanner (1952) illustrated these extremely long
cartilages in the Plethodontidae.

The second basibranchial is present in all of
the ambystomids studied. It is lost in most sala-
mandrids (Ozeti and Wake, 1969) and is prob-
ably completely missing in plethodontids (Dunn,
1926; Tanner, 1952).

All of the foregoing facts support the hypoth-
esis that the ambystomids are primitive to the
salamandrids and that the latter are in turn prim-
itive to the plethodontids. There is reason, how-
ever, to consider another alternative. If ossifica-
tion of the hvobranchium is to be considered a
primitive character, then with respect to this
character the Salamandridae would be primitive
to the Ambvstomidae and the Plethodontidae.
Tile ceratohyal, for example, is from 30 to 50
percent ossified in most salamandrids ( Ozeti and
Wake, 1969), whereas it is much less than this
in Ambvstomidae (Table 1) and in the Pletho-
dontidae (Tanner, 1952; Wake, 1966). The first
ceratobranchial is totallv unossified in the Am-
bystomidae and well ossified in most Salamandri-
dae. However, there are salamandrids with total-
lv cartilaginous hvobranchial skeletons, and,
therefore, since this high le\'el of ossification is
not consistent throughout the family, the use of
percentages of ossification as an index of phylo-
genetic relationships between families seems to
be unjustified. In fact, there is considerabl(> vari-
ation within each familv with respect to per-
centages of ossification, and, moreover, an inter-
esting parallelism seems to have occurred in each
familv. The advanced members of a particular
family seem to be more c;ipable of terrestrial

Biological Sehils. \'ol. 16. No. 1

HV013HANt:HllM AM) TlIllOAl MVOLOCV

65

feeding habits, and in these salamanders there
has been a reduction in the amount of calcifi-
cation in the hvobranchium, thereby making it
more flexible to facilitate the capture of prey
through tongue pad flipping. The highly ad-
vanced free-tongued plethodontids studied by
Tanner (1952) were shown to have no ossifica-
tion whatever in the hyobranchium, whereas the
more primitive plethodontids as reported by
Wake (1966) did have definite centers of bone
in the hyobranchium. The ambystomid which is
most terrestrial in its feeding habits, Rhijacotri-
ton, has a completely bone-free hyobranchium;
and the terrestrial feeders among the Salaman-
dridae (Salanuindra, Cliioglossa, and SaJainan-
drina ) are likewise e(|uipped with more cartilage
in the hyobranchium than the other presumably
more primitive members of the family ( Ozeti
and Wake, 1969). It would appear, therefore,
that the percentage of ossification of the hyo-
branchium may have some value as an index to
phylogeny. but is of limited value if one wishes
to studv interfamilial phvlogenv. Thus, the per-
centage of ossification has importance in deter-
mining the degree of adaptation toward a ter-
restrial habitat.

An investigation of the muscle adaptations is
also impK>rtant and of interest in an assessment
of de\elopment toward the more terrestrial type
of feeding. An examination of the more ad-
vanced plethodontids indicates a greater devel-
opment of the anterior transverse muscles. Ex-
cept for the genus Thorhis those genera of free-
tongue plethodontids studied by Tanner ( 1952 )
possessed a large anterior development of the
intemiandibularis. According to his analysis, this
represents the intermandibularis anterior, a mus-
cle thought by Piatt (1935) to be lost at meta-
morphosis. This must yet be demonstrated by an
cmbryological investigation; however, the fact
still remains that in these plethodontids there
is a relatively large mass of transverse muscle
which ties the anterior part of the mandibular
area together and which is not interrupted by
the linea alba. The development anteriorly is al-
so accompanied bv a reduction in the size and
extent of the transverse muscles in the more f>os-
terior areas of the throat.

If we examine other related but more primi-
tive families ( salamandrids and ambystomids ) .
we do not see the same types of muscle develop-
ment; but we do see in the more terrestrial forms
of these families a development of the more an-
terior transverse muscles. In the Salamandridae,
for example, the genera in which terrestrial feed-
ing is best developed are Sahinmndni. Chioglos-
sa. and Sahimandriiui; and, furthennore, these
three genera tend to have a greater degree of

elaboration of the anterior portions of the throat
musculature than other genera in the family
(Ozeti and Wake, 1969). There is also a con-
comitant reduction in the posterior parts of the
throat musculature in these land feeders. The
three genera of salamandrids noted above all
have experienced a reduction in myosepta of the
rectus cervicis superficialis as compared to the
other members of the family, and a correspond-
ing reduction in rectus cervicis superficialis myo-
septa has occurred in Wujacotriton of the Am-
bijstomidac .

In the plethodontidae there is also a reduc-
tion of the myosepta in the rectus series. Tan-
ner ( 1952) figures this reduction and also indi-
cates a reduction and finally a complete loss of
the omohyoideus muscle, which in the less ad-
vanced species ties the rectus throat muscles to
the pectoral girdle (scapula). There is also a
gradual reduction in the size of the second basi-
branchial (os thyroideum) until it is lost in the
free-tongued plethodontids. These reductions
and deletions have also served to increase the
flexibilitv of the longitudinal throat muscles and
have produced an uninhibited band of muscle
from the pelvis to the hyobranchium providing
a mechanism for rapid tongue retraction.

An analysis of the myology and osteology of
the hyobranchium of the families Ambystomidae,
Salamandridae, and Plethodontidae provides evi-
dence to support the conclusion that there has
been a degree of parallel development leading
toward a terrestrial development of feeding and
perhaps breathing. In all three families, we note
a reduction in the ossification of the skeletal
elements of the hyobranchium depending, we
believe, on the degree of the terrestrial type
feeding achieved by members of a generic or
specific group within these families. The flexi-
bilitv of the hvobranchium achieved by the free-
tongued plethodontids is most remarkable and
has virtually eliminated in some species that
tvpe of feeding in which jaws are used as the
grasping or food-getting device. One is caused
to wonder if the development of the tongue as
an almost exclusive food-getting device is the
ultimate type of development to be achieved in
the process of adaptive radiation in terrestrial
salamanders and perhaps also in lizards ( Chame-
leon idae).

The conversion of the skeletal elements into
a flexible cartilaginous structure and the reduc-
tion in size and deletion of some elements has
])een accompLmied by modifications in the myol-
ogy. As indicated above, there have been basic
modifications in the muscles which have in ef-
fect reduced the heavy musculature of the throat
area. Associated with this reduction, there has

66

BnicHAM Young University Science Bulletin

l)ecn a specialization in both the transverse and
longitudinal muscle series. A specialization of
particular interest to us is the development of
the most anterior transverse muscles in certain
groups of each family in which the muscle fi-
bers extend uninterrupted across the midline.
Tanner ( 1952) referred to these as the inter-
mandibularis anterior and based his conclusions
on the fact that in the plethodontids studied
there was a small tendinous fascia tying each
end of the muscle to the opposing mandible.
A comparison of this muscle in those ambysto-
inids and salamandrids in which fibers extend
across the linea alba suggests that in these fami-
lies those continuous fibers represent the anterior
fibers of the intennandibularis posterior which
have been modified to strengthen the anterior
area of the now changing food-getting mecha-
nism. We have found no indication that the inter-
mandibularis anterior is retained in any members
of the latter two families. This anterior myologi-
cal specialization suggests that there is a definite
convergence leading toward a terrestrial type of
feeding which involves an increasing use of a
more flexible tongue.

If we examine the posterior transverse mus-
cles, we note a reduction in size and mass with
an increase in the width of the central fascia as
the entire structure becomes more flexible. The
longitudinal series has also been effected to the
extent that they have become more elongate (in
some plethodontids the rectus cervicis series are
actually folded. Tanner, 1952, plate 73, Fig. G).
Ozeti and Wake (1969) indicate a reduction in
the numbers of mvosepta in the rectus series of
such genera as Salamandia, Chioglossa, and
Salanumclrina. We partcularly note a similar
change in Rhtjacotriton in which the myosepta
of the rectus series have been reduced to two.
In RJitiarotritan the genioglossus has been ex-
tended beyond the first radii of the first basi-
branchial and reaches posterior to insert on and
in fascia attached to the first ceratobranchial.
This adaptation provides for a longer muscle and
permits a greater length of contraction in pulling
the hyobranchium anteriorlv in the extending of

the tongue. Although we note that different
structures have been altered, the general effect
appears to be the same, namely, that terrestrial
groups in each faniilv have modified the hyo-
branchium to provide for greater flexibility and
have increasingly involved the tongue as an
evermore important food-getting organ.

We have found the musculature of the Am-
bystomidae to be more advanced in some re-
spects than previous workers have reported. Con-
trary to Piatt's findings, the ambystomids do
have a subhyoideus, and therefore this character
cannot be used to separate the families as it has
been in the past. Piatt further reported that the
transverse ventral throat muscles posterior to the
intermandibularies posterior are the interhyoid-
eus posterior in Crytobranchidae, Amphiunii-
dae, and Ambystomidae and that these two mus-
cles remain in the adults in an essentially un-
changed condition. We have foimd the inter-
hvoideus posterior to be the same as the larval
muscle. In the Plcthodontidae, this muscle be-
comes divided into the gularis and the quadra-
topectoralis. The interhyoideus of the larva, how-
ever, becomes divided into subhyoideus and in-
terossaquadrata in the adults of Ambystomidae
and Salamandridae.

This new infonnation on myology does not
answer the ((uestion with regard to the relation-
ship l^etween Ambystomidae and Salamandridae
except that it does make the ambystomids ap-
pear more advanced than they were previously
thought to be. The preponderance of evidence
still seems to support the concept that the Am-
bystomidae are primitive to both the Salamandri-
dae and the Plcthodontidae.

In the past it has been assumed that the or-
der of phylogeny is from the Ambystomidae to
the Salamandridae to the Plcthodontidae. How-
ever, based on the findings of this study there is
a second distinct possibility that the ambysto-
mids mav have given rise independently to both
the salamandrid and plethodontid families.
These two possibilities are presented in Figure
20.

SUMMARY

1. The hyobranchial skeleton is mostly carti-
laginous with a few isolated areas of bone lo-
cated in certain components. Only in Dkainpto-
don is ossification widespreail throughout the
livobranchium, although this ossification is not
heavy. In only a few areas in Dicamptodon, such

as the tips of the ceiatoii\als, is the ossification
heavy.

2. The first basibranchial is shield-shaped,
pear-shaped or rod-shaped and usually has a
central core of bone. Bone is completely absent
in Rhyacotriton and in A. gracile.

Biological Sekies, Vol. 16, No. 1 lIvoiuiANt hium and TiinoAr Mvi>loc;v

PLETHODON riDAE

67

SALAMANDRIDAE

AMBYSTOMIDAE

■f -f ♦ -f -f

SALAMANDRIDAE PLETHODONTIDAE

AMBYSTOMIDAE

Fig. 20. Diagrams depicting the possible phylogenetic relatioiisliips between the .Ambvsfomidae. Salamandridae,
and Plethodontidae.

3. The first radial cartilages are entirely car-
tilaginous. They are usually long, slender rods
that may or may not unite with the tips of the
ceratohyal; although in A. cingitlatiun they are
very short and rounded. In Dicamptodoti there
is a joint midwav along the length of the first
radial.

4. The second radial cartilages are always
unossified and curve dorsally to meet the oto-
glossal. Dicainptodon has a fusion of the second
radials and the otoglossal plate to form a single
platelike structure.

5. There is usually no ossification in the first
ceratobranchial. One specimen of A. ciiiriiilatum
contained a spot of hone in the medial end of
this structure which represented 11 percent of
the total volume of the first ceratobranchial.

6. The second ceratobranchial is always un-
ossified and articulates with the first branchial
arch either at the joint between the first cera-
tobranchial and the epibranchial or immediatelv
posterior to that joint.

7. There is an epibranchial cartilage in all
species included in this study. However, in some
the articulation of this structure with the first
ceratobranchial is indistinct. This is the case in
Dicamptodoti, A. anmdatnm, and A. cingulatiun.

S. The ceratohval is often ossified on its pos-
terior or distal tip.

9. The second basibranchial is present in all
species. Its shape is .3-radiate, 4-radiate, or an
inverted V. Usually, the central portion is ossi-
fied with the tips of the lateral amis remaining
cartilaginous. In Dicomptodon, Rhtjacotriton,

and A. opacum, the entire structure is cartilag-
inous.

10. The linea alba is usually wide at the
level of the intennandibularis posterior. In A.
nuiculatum it is a very thin line.

11. The fibers of the intennandibularis pos-
terior insert into the linea alba of the throat; ;U-
though, in Rhyacotriton, A. cingtdatum, and A.
gracde, a few of the anterior fibers extend all
the way across the midline without being inter-
rupted by the linea alba.

12. The levator mandibulae is hypertrophied
in Rlii/acotriton and Dicamjdodon. This may be
associated with their stream feeding habit, which
rc<(uires a more aggressive behavior than used
by the other members of the family (genus Am-
I)t/.stoitia) in their pond habitats.

1.3. Contrary to the reports of earlier workers,
the subhyoideus is found in all species studied.
It is derived from the larval interhyoideus which
at metamorphosis divides into the subhyoideus
and the interossa(juadrata. It is usually small and
delicate and extends forward only as far as the
anterior edge of the ceratohyal. In Rhijacotriton,
however, it is longer, extending all the way for-
ward to the mandible. In Dicainptodon, this
muscle is not as long as in Rhyacotriton, but it
is thick and heavv when compared with other
species.

14. The genioglossus is divided into a ven-
tral and dorsal portion in all species. Only in
Rlu/acotriton does this muscle extend as far pos-
teriorlv as the first ceratobranchial.

15. A hvoglossus is present in all .species ex-

m

Bricham Youno Univfrsity Scienck Bulletin

cept Dicamptodon. The posterior portion of the
genioglossus in DUainptodon lies in the position
occupied by the hyoglossus in the other members
of the family, and it appears likely that the hyo-
glossus is derived from the genioglossus by spe-
cialization of the posterior part of the muscle.

f6. There are usually three transverse in-
scriptions on the rectus cervicis superficialis. In
Rliijacotriton and A. talpokleum, however, there
are only two.

17. The rectus cervicis profundus usually at-
taches to the otoglossal cartilage by a fleshy
rather than a tendonous insertion. In A. annula-
tum and A. cing^tilatum this insertion is by way
of a thin tendon.

IS. The hebosteoypsiloideus originates on the
second myoseptum of the rectus cervicis profun-
dus posterior to the second liasibranchial in all
species except A. anmikituin and A. opacum, in
which it arises from the first inscription.

19. The variation in ratios of dimension of
hyobranchial skeleton elements that is found
within a given species is excessive beyond that
which would allow this infomiation to be of
value in species differentiation. Whereas these
dimensions have been used successfully by other
workers in the phylogenetic alignment of sala-
manders at the generic level, this study indi-
cates they are not equally usable in studies at
the species level.

20. The 16 species and subspecies are placed
into six groups according to their throat morphol-
ogy as follows: The Dicmnptodon group, the
Rhtjiicoiriton group, the A. annuhittim-A. ciu'^u-
Idtum group, the A. grr/ri/c-A, jcffcrsonwnum- A.
nud)eei-A. nuicrodactiihim-A. nuicitlatum group,
the A. taJpoidcnm group, the A. opacum-A. tex-
anum group, the A. wsaceum- A. t. tigrinum-A.
t. melonostictum- A. t. nehulosum group.

21. Dicmnptodon appears to be the most

primitive group, and Rlu/acotriton, the most
specialized.

22. Based on the morphology of the throat re-
gion, it is suggested that A. rosacettm be placed
in the tiger salamander group as a subspecies of
A. tigrinum.

2.3. The binar\' characters when applied sep-
arately to the hyobranchial skeleton and the hyo-
branchial musculature produce essentially the
same phylogenetic results.

24. In comparing the hyobranchium and
throat myology in different families of salaman-
ders, there appears to have developed a parallel-
ism within each family with respect to adapta-
tions toward terrestrial feeding. These adapta-
tions include the reduction and even complete
loss of bone in the hyobranchium. Tlie elabora-
tion of the anterior tliroat niusculatiuc to pro-
vide a mechanism for rapid tongue pad flipping
has resulted in a reduction in the size of muscles
and in the numl)cr of nivosepta in the posterior
throat nnisculatiire. These adaptive changes
seem to have occurred as the need for sucking
action and jaw movement seen in aquatic sala-
manders are replaced by rapid tongue protru-
sion and retraction in the terrestrial forms.

25. The results of this investigation support
the hypothesis that Ambystomidae is primitive
to Salamandridae and to Plethodontidae. The
findings which have led to this conclusion are
based upon the protrusibility of the tongue and
upon the structure and presence of the epibran-
chial, otoglossal, second radial, and second basi-
branchial cartilages of the three families. The
muscles which have added infonnation concern-
ing these interfamilial relationships are the in-
temiandibularis posterior, levator mandibulae,
subhyoideus, genioglossus, hyoglossus, rectus cer-
vicis profundus, rectus cervicis superficialis, and
hebosteoypsiloideus.

ACKNOWLEDGEMENTS

We express our sincere gratitud<> to Doctors
F.A. Andersen, H.H. Frost, B.F. Harrison,
Glen Moore, and J.R. Murphy for their advice
and suggestions as they reviewed this study. We
also appreciated the opportunity to dissect many
specimens from the Hrigham Young University
.Natural History Museum (BYU). For the use of
specimens Irom other individuals and niuseiuns,
we are grateful. 'I'hese include the following: J.
A. Peters, U.S. National Mu.seum (USNM); F.
M. Creusere, Universitv of Kentucky (UK); J.

H. Bailey, Duke University (DU); J.M. Walker,
University of Arkansas (UA); A.E. Leviton,
(California Academy of Sciences (CAS); R.C.
Snyder, University of Washington (UW); J.D.
.\nderson, Rutgers University ( RU ) ; D.F.
Avery, Southern Connecticut University (SCU);
J.D. Hardy, Chesapeake Biological Laboratory
(CBL); and L.D. Fisher, student at University
of Minnesota; specimens were donated and bear
a BYU number.

Biological Series, \'c)i.. 16, No. 1

HvdHKANCHIUM AND TllHOAl MvoLOCY

69

For advice concerning the statistical methods William Ingram III. We also appreciate the ef-
and computer programs used we are grateful to fort and patience of Mrs. Karen Krogh who did
Dr. Melvin Carter, Mr. Keith Blake, and Mr. the typing and assisted with the illustrations.

LITERATURE CITED

Anherson, J. D. 1961. The life history and sys-
tematica of .Am/;i/sf()mH rosdccum. Copeia, 1961(4):
371-376.

Barbour, T. 1926. Reptiles and amphibians. Their
habits and adaptations. Mifflin Co.. Boston and
New York, pp. 1-129.

Bishop, S. C. 1943. Handbook of salamanders. Com-
stock Publ. Assoc, Ithica, New York, pp. 1-555.

BocoLjuBSKY. S. N. 1924. Zur morphologies des
hvoid-apparatus bei den amphibien ( Salamandni
ciiucasica nnd Triton). Rev. Zool. Russe 4(1-2):
89-114. (In Russian, German abstract.)

BouLENGEH, G. .•\. 1910. Les batraciens et principalc-
ment ceu.x d'Europe. Paris, pp. 1-305.

Brandes, G., and W. Schoenichen. 1901. Die bnit-
pflege der schwanzlosen batrachier. Abh. Nat. Ges.
Halle, 22:1-69.

Brandon, R. A. 1961. A comparison of the ]ar\ae of
five northern species of Ambystoma ( Amphibia,
Caudata ) . Copeia, 196 1 ( 4 ) : 35.5-383.

Cope, E. D. 1887. The hyoid structures in ambystomid
salamanders. Am. Nat.'. 27:87-88.

Di.xoN, W. J., and F. J. Massev. 1969. Introduction
to Statistical AnaKsis. 3rd ed. New York, McGraw-
Hill.

Druner, L. 1901. Studicn zur anatomic der zungeii-
bein-, kiemenbogen-und kehlkopfmuskeln der uro-
delen. I Theil. Zool. Jahrb. Anat. Ont., 15:435-622.

Dunn, E. R. 1926. The salamanders of the family
Plethodontidae. Smith College, Northhampton,
Mass., pp. 1-441.

. 1940. The races of Anihi/stonui tit^rininii.

Copeia. 1940(3 ): 154-162.

EncEWORTH, F. H. 1935. The cranial muscles of
vertebrates. Cambridge Univ. Press, London, pp.
1-493.

Fr-\ncis, E. B. T. 19.34. The anatomy of the sala-
mander. Clarendon Press, O.vford, pp. 1-381.

Hansen, A. M., and W. W. Tanner. 19.58. A com-
parative osteological study of certain species be-
longing to the genus BoUtonlossa (Ampliibia).
Great Basin Nat., 18(3-4 ) :85-100.

Hilton, W. A. 1947a. The hvobranchial ■.keleton of
Plethodontidae. Herpetologica, 13:191-194.

. 1947b. The hvobranchial apparatus of Sala-

mandridae. Jour. Ent. and Zool.. 38(l):68-72.

Kallius, E. 1901. Beitriige zur entwickelung der
zunge. I. Theil, .\mphibien iind Reptilien. Z. Anat.
Ent^vickl. Anat., I6:,531-760.

Kerr, J. G. 1919. Textbook of embryology, 11 Vertc-
brata. London, pp. 1-163.

Latimer, H. B., ani> P. G. Rf)()EE. 1964. Weights and
linear measurements of the bodv and organs of the
tiger salamander, before and after metamorphosis,
compared with the adult. Anatomical Rec, 148(2):
139-147.

Noble. G. K. 1927. The value of life history data
in the study of the evolution of the amphibia.
Ann. N. Y. Acad Sci.. 30:31-128.

. 1931. The biology of the Amphibia. McGraw-
Hill, New York, pp. 1-174.

Oppel, a. 1900. Lehrbuch der vergleichenden mikro-
skopischen anatomic der wirbeltiere. Bd. 3, S. 1-10,
1-1180. Jena.

OzETi, N., AND D. B. Wake. 1969. The morphology
and evolution of the tongue and associated struc-
tures in salamanders and newts ( Family Salaman-
dridae). Copeia, 1969( 1 ) :91-123.

PiATT, J. 1935. \ comparative study of the hvobran-
chial apparatus and throat musculature of the
Plethodontidae. J. Morph., 57 (1):213-251.

. 1938. Morphogenesis of the cranial muscles

of Aml)hjitom(i punctiitiiut. J. Morph., 63(3) :531-
587.

1939. Correct terminology in salamander

myology. I. Intrinsic Ciill Musculature. Ibid., 1939

(4):22b-224.
. 1940. Correct terminology in salamander

myology. II. Transverse Ventral Musculature.

Copeia.' 1940( 1 ) :9-14.
Regal, P. I. 1966. Feeding specializations and the

classification of terrestrial salamanders. Evolution,

20(3):392-407.
Sampson, L. V. 1900. Unusual modes of breeding

and development among Anura. Amer. Nat., 34:

687-715.
Smith, L. 1920. The hvobranchial apparatus of

Spclcrpes hislineatua. J. Morph., 33:.527-.582.
Stadtmiiller, F. 1936. Kranium und visceralskelett

der stegocephalen und amphibien. Hanb. Bergl,

Anat. Wirbelt. 4:. 501 -698.
Tanner, W. W. 1952. A comparative study of the

throat musculature of the Plethodontidae of Mexico

iind Central America. Lfniv. K'ans. Sci. Bull., 34

(10):. 58.3-677.
Tihen, J. A. 1958. Comments on the osteology and

phylogenv of ambystomid salamanders. Bull. Fla.

St.' Mus.,' 3:1-50.
Wake, D. B. 1963. Comparative osteology of the

plethodontid salamander genus Aneidc.s. ]. Morph.,

1I3(1):77-118.
. 1966. Comp:irative osteology and evolution of

the lungless salamanders, family Plethodontidae.

Mem. So. Calif. Acad. Sci., 4:1-111.
Wake, D. B., and N. Ozeti. 1969. Evolutionary re-

laticmships in the family Salamandridae. Copeia,
1969(1 ):124-127.
WiEDERsiiEiM, R. 1877. Das Kopfskelet der urodelen.

Gegen. Morph. Jahrb.. 3:352-.548.
WisnAHT, D. 1969. Clustan lA user manual. Com-
puting Lab., Univ. of St. .\ndrews, Fife. Scotland.

MUS. CO MP. 200L
LIBRARY

JAN 5 1973

<^_|J/^_P^ov.«'> HARVARD

UNIVERSITY.

Brigham Young University
Science Bulletin

UNDERSTORY CHARACTERISTICS

RELATED TO SITE QUALITY

FOR ASPEN IN UTAH

by

James H. Warner

and

K. T. Harper

BIOLOGICAL SERIES — VOLUME XVI, NO. 2
APRIL 1972

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 Brigham
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Brigham Young University
Science Bulletin

UNDERSTORY CHARACTERISTICS

RELATED TO SITE QUALITY

FOR ASPEN IN UTAH

by

James H. Warner

and

K. T. Harper

BIOLOGICAL SERIES — VOLUME XVI, NO. 2
APRIL 1972

TABLE OF CONTENTS

INTRODUCTION 1

LITERATURE REVIEW 2

METHODS 3

Location of Study Areas 3

Tree Samples 5

Herb Layer 5

Soil Studies 5

LABORATORY METHODS 7

Plant Species _ 7

Environmental Factors 7

Data Analysis 7

RESULTS 7

Stand Distribution 7

Prevalent Species 7

Species Associated by Chi-Square 8

Site-Quality Determinations 11

Environment _ 14

DISCUSSION 14

ACKNOWLEGMENTS 18

LITERATURE CITED 19

Brigham Young University Science Bulletin

tion. Undoubtedly, a large part of the neglect is
attributable to the fact that the species was con-
sidered a "weed" tree for many decades by re-
source managers. At the present time, the litera-
ture pertaining to aspen in the West consists of
numerous notes and peripheral studies consider-
ing individual facets of the species' ecology; but
Baker's study (1925) is the sole attempt to in-
terpret the general ecology of this forest-type.
Even Baker's classical study largely ignores char-
acteristics of the understory vegetation and de-
tails of the soil environment. Thus, at the present
time when new and often conflicting demands
are being made on the aspen-type, western re-
source managers find themselves confronted with
a bewildering array of ecological questions for
which there are few, well-documented answers.

This study was designed to provide a better
understanding of the relationship between un-
derstory composition and site-quality for aspen
itself. An attempt is made to devise a scheme for
predicting site-quality from understory vegeta-
tion. Also, some preliminary correlations between
gross characteristics of the understory and
height-over-age relations of aspen are made.

Inasmuch as a major portion of the acreage
of aspen in western America is confined to the
states of Colorado and Utah, it was considered
desirable to conduct the stud\' within that area.
Preliminary work indicated that a broad range in
site-quality and understory characterisitics could
be found in aspen forests of Utah and northern
Arizona. Accordingly, all field work was confined
to that area to minimize travel time.

LITERATURE REVIEW

Baker's paper ( 1925 ) stands as the only in-
tensive consideration of the ecology of quaking
aspen in the West, although several other inves-
tigators have studied individual facets of aspen
ecology. Age and height values for aspen grow-
ing in association with different understory types
in Montana have been reported by Lynch
(1955). Alder (1969) has reported on the age
structure of aspen forests of Utah. A recent pop-
ularized account of aspen ecology in the West
was published by Cottam ( 1963 ) . Langenheim
( 1962), working in western Colorado, and Ream
(1963), in the Wasatch Mountains of Utah and
Idaho, drew general conclusions about the rela-
tive position of aspen in the environmental com-
plex in their respective areas. Heinselman and
Zasada (1955), Strotham and Zasada (1957), and
Powells ( 1965) have attempted to integrate pub-
lished reports of the behavior and use of quaking
aspen throughout North America.

Compositional characteristics of the under-
story in aspen forests have been more thorough-
ly investigated in the West than has the tree it-
self. Numerous papers have contributed to our
knowledge of the understory vegetation associa-
ted with aspen. Lynch's studies (1955) provide a
useful understanding of the understory vegeta-
tion of aspen in northern Montana. The nature
of the understory in Wyoming is set forth by J.
Reed ( 1952), R. Reed ( 1969)^ and Hoff ( 1957).
For Colorado, papers by Costello (1944), Hoff
(1957), Langenheim '(1962), and Morgan
(1969) give useful data for the aspen-type. In
Utah, Allan (1962) and Ream (1963) provide
data on the understory of aspen in the Wasatch
Mountains, and Ellison and Houston ( 1958) dis-

cuss the production of herbaceous material under
the canopy and in treeless openings of aspen
forests of central Utah. Houston (1954) pub-
lished a "condition" guide for aspen ranges in
the Intermountain Region which gives some idea
of the impact livestock have had on the type. In
none of the foregoing papers, however, has
there been an attempt to relate the performance
of individual species or groups of species to
specific characterisitics of the abiotic environ-
ment or to the performance of aspen itself.

There is clear evidence that many aspen for-
ests throughout the West are currently being re-
placed by conifers in the course of natural suc-
cession ( Baker, 1918 and 1925; Daubenmire,
1943; Hall, 1953; Costing, 19.56; Hoff, 1957; and
Cottam, 1963). There are numerous indications
in the literature, however, that the successional
processes are extremely slow or nonexistent in
manv parts of western America ( Fetherolf, 1917;
Reed, 1952; Lvnch, 1955; Marr, 1961; Langen-
heim, 1962; Juiander et al., 1964; Morgan, 1969;
and Reed, 1969). A more thorough knowledge of
the successional process would seem to be need-
ed if the best management procedures possible
are to be implemented.

The nature of aspen forests can also be great-
ly modified by the influence of domestic graz-
ers (Baker, 1918; Sampson, 1919; Houston,
1945; Coles, 1965; Hutchins, 1965; and Lucas,
1969) and big game (Coles, 1965 and Alder,
1969). It seems possible that aspen can actually
be eliminated from a site if animals are permit-
ted to repeatcdlv browse off the trees' reproduc-
tion (Alder, 1969 and Lucas, 1969).

Biological Series, Vol. 16, No. 2 Aspen in Utah

The most conspicuous deficiency in all of the
foregoing studies lies in their failure to study the
abiotic environment in detail. The soil studies of
Lynch ( 1955 ) are not presented in a form suit-
able for interpretation of the perfoniiance of in-
dividual plant species. Likewise, a detailed study
of the influence of topographic position on the
behavior of aspen has never appeared in the lit-
erature. Crovvther and Harper ( 1965 ) have
studied the influence of parent material on as-
pen forest development in Utah, but the study
was confined to a single watershed and a nar-
row range of parent materials. A useful study by
Jones ( 1967a ) considers the influence of various
combinations of climatic, topographic, and eda-
phic characteristics on height growth of aspen
stands in Colorado, New Mexico, and Arizona.
Additional studies of this type are needed in
other parts of the Mountain West.

Disease appears to play a key role in the dy-
namics of aspen forests in the West. Meinecke
(1929), Mieike (1957), Hinds (1962), and Kre-
bill ( 1971 ) discuss the major diseases attacking
western aspen. The influence of ecological con-
ditions on distribution and severity of impor-
tant diseases is still largely unknown, however.

Attempts to utilize understory species as in-
dicators of site potential for either tree growth
or herbage production as advocated by early
ecologists (Korstian, 1917 and Riggs, 1929) have
not usually been successful (Kittredge, 1938 and
Driscoll, 1963). However, a few workers have
used understory species as indicators of site po-
tential for tree growth with success ( Rowe, 1956;
Gadnon and MacArthur, 1959; Hills and Pier-
pont, 1960; Daubenmire, 1961; and Orloci,
1962), but methods employed have not usually
been well quantified.

METHODS

In this study, an attempt is made to develop
a simplified and completely quantitative ap-
proach to the use of understory species as indi-
cators of site-quality. Statistical reliability of in-
dicators will be verified and a straightforward
technique for integrating response of many indi-
cators devised. Once the indicators are identi-
fied, any person with the ability to identify those
species should be able to derive the index for
any given site.

The 49 stands employed in this study were
initially ordered according to site indices derived
from tree height-over-age data using Jones's
( 1967b) natural site index curves. The statistical
relationship between Jones's natural site index
for each of the several stands and the index val-
ues derived by the use of understory species for
the same stands was determined by simple cor-
relation.

Location of Study Areas

Study sites extend along the mountainous
backbone of Utah from Logan Canyon on the
north to Fish Lake on the south. Six additional
stands are located in northern Arizona on the
Kaibab Plateau (see Fig. 1 and Table 1).

All stands selected were approximately 0.04
liectare (0.1 acre) in size and satisfied the fol-
lowing criteria: (1) tree density appeared aver-
age for the particular site ( very sparse and very
dense stands were avoided); (2) aspen contri-
buted 100 percent of the tree canopy; (3) aver-
age tree age was in excess of 30 years; (4) area

sampled was well buffered on all sides by simi-
lar aspen forest; and (5) aspect, soil, topo-

F"igure 1. Distribution of stands.

Brigham Young University Science Bulletin

Table 1. Location of stands and the gross environmental characteristics of each.

Stand
No.

Elev.

(m)

%
Slope

E.\posure°

Township-Range
Section

Natl. Forest

Area Name

1
2
3

2469
2469
2499

12
15
19

20
295

23

T8N R4E S21
T8N R4E S21
T8N R4E S21

Cache

Squaw Flats

4
5
6

2667
2301
2865

47
53
36

190
180
158

T2S R3E S26
T2S R3E S17
T2S R3E S25

Wasatch

Big Cottonwood
Canyon

7
8
9

2435
2408
242.3

6
43
13

115

47

105

T4S R3E S33
T4S R3E S33
T4S R3E S33

Wasatch

Aspen Grove

( Mt. Timpanogos )

10
11
12

2591
2591

2591

11
14
13

0
325
348

T3S R12W S24
T4S R12W S24
T4S R12W S24

Uinta

Mud Creek

( Strawberry Reservoir )

13
14
15

2789
2576
2667

20

29

8

50
100

70

T35N R3E S34
T38N R2E S20
T37N R2E S34

Kailiali

Jacob Lake
Arizona

16
17
18

2515
2499
2545

5

7

29

120

35

150

T33N R2E S4
T33N R2E S4
T32N R3E S5

Grand Canyon
Natl. Park

North Rim

19
20
21

2652
2682
2743

14
10
32

200
113

225

T17S R4E S20
T17S R4E S20
T17S R4E S20

Manti-LaSal

Clreat Basin E.xp. Sta.
( Ephraim )

22
23

2134
2438

7
17

270
260

T8N R4E S20
T8N R4E S20

Cache

Ant Flats

24
25

2377
2,377

27
15

90
280

T4S R3E S33
T4S R3E S33

Wasatch

Aspen Grove

26
27

2057
2073

31
40

315
315

TIN R3E S20
TIN R3E S20

Wasatch

Red Butte

Canyon

28

2316

NA

195

TIS R3E S31

Wasatch

Mill Creek Canyon

29
30
31

2758
2774
2743

8
30
31

10

75

120

T24S R3E S24
T24S R3E S24
T24S R3E S24

Fish Lake

Cold Springs

32

2323

30

210

T2S R3E S26

Wasatch

Big Cottonwood

33
34
35

2435
2591
2438

11
16

48

150

95

150

T14S R6E S34
T14S R6E S26
T14S R6E S26

Manti-LaSal

Scad Valley

36
37
38
39
40

2731
2792
2484
2408
2522

18
21
20
12
9

70

60

347

145

40

T6S R6E S7
T6S R6E S7
T5S R5E S36
T5S R6E S26
T5S R6E S26

Uinta

Wallsburg Road
( Daniels Canyon)

41

2234

6

60

T3S R12W S35

Uinta

Bryants Fork

42
43
44

2408
2385
2469

13
27
48

160
150
180

T13N R3E S7
T13N R3E S7
T13N R3E S8

Cache

Tony Grove

{ Logan Canyon )

45

2347

43

345

T17S R4E S23

Manti-LaSal

Willow Creek

46

2713

8

300

T17S H.5S S28

Manti-LaSal

Tliistlr Flats

47

2286

NA

200

TIS R3E S31

Wasatch

Mill Creek Canyon

48
49

2432
2408

NA
NA

NA
NA

T4S R3E S33
T4S R3E S33

Wasatch

Aspen Grove

' In degrees, clockwise from north.
NA = Dat.T nol nvail.nble.

Biological Series, Vol. 16, No. 2 Aspen in Utah

graphic position, slope, and vegetational pattern
were unitonn throughout the area sampled.
Stands were almost exclusively grazed by deer
and or domestic sheep. The three stands on the
Kaihab National Forest are now grazed by cat-
tle but have had a long history of sheep use un-
til the recent past. All sampling, except for soil
temperature, was completed during the sum-
mers of 1965 and 1966.

Stands were selected on a variety of sites sup-
porting pure aspen forests (i.e., to the extent
possible, serai stands were avoided). An attempt
was made to include stands on the full range of
site conditions within the defined area (see
Fig. 1 ) . For ease of accessibility all stands were
located within national forests and national
parks. Detailed maps were drawn of each stand
and its surroundings so that the exact location
could be easily located on return visits. Stands
^^'erc square-shaped and marked at each corner
with orange surveyor's tape.

For each stand, tlie following data were re-
corded: (1) location, (2) elevation, (3) aspect,
(4) slope, (5) description of landscape, and (6)
nature of any disturbing influence.

Tree Samples

A tree is here interpreted as any woody stem
having a diameter in excess of 10 cm (4 in) at
breast height (dbh). All trees within the borders
of each stand were measured and the following
data recorded: (1) dbh, (2) height (using
an optical range finder), and (3) evidence of
disease, fire, or animal damage. In addition,
cores from the 10 tallest trees were taken at
breast height with an increment borer for sub-
sequent aging in the laboratory. An attempt was
made to locate stand boundaries so that all trees
of the stand were of the same clone. When it
could be detemiined, the sex of the clone was
recorded.

Herb Layer

The herb layer was sampled immediately fol-
lowing establishment of the stand boundary in
order to minimize the effect of trampling. Twen-
ty-five sc|uare quadrats, each 0.2.5 m- in area
(1/16,000 acre), were uniformly distributed
throughout the stand for frequency data. Above
ground, current year production to a height of
1.8 m (6 ft) was taken in 42 stands between July
15 and August 15 of the years of study. Produc-
tion was estimated on .36 cjuadrats (0.89 m- or
9.6 ft") per stand. Accuracy of the estimate was
regularly checked by clipping and weighing.
Field weights were converted to air dry weights

by use of conversion factors obtained by drying
vegetational samples from each study area.

A list of all vascular plants present within the
boundaries of each stand was prepared (War-
ner, 1971). Species were classified into broad
life-form and longevity classes (Table 2). A de-
gree of arbitrariness was involved in the assign-
ment of perennial forbs to height classes. Osmo-
rluza cJuIensi.s and Valeriana occidentalis, for in-
stance, occasionally grow to a height in excess of
.30 cm ( the breaking point between short and
tall forbs ) ; but since their photosynthetic surface
is almost wholly developed close to the ground,
we have classified these species as short forbs.

Soil Studies

In each stand, soil samples were collected
and soil temperature measured. Soil profile data
were obtained from a pit dug to the depth of
1.5 m (60 in) or to bedrock, whichever was
shallower. Samples were taken from each horizon
of the profile with a bulk density sampler of 750
cc capacity. For each horizon, the occurrence of
mottling or gleying and physical properties such
as structure, color (by Munsell soil color chart),
percent by volume of rock over 2 mm diameter,
and the amount of rooting were recorded. Soil
samples from each horizon were placed in plas-
tic bags and returned to the laboratory, where
they were dried at 105° C for at least 24 hours.

Soil temperatures were taken at the center of
each stand at the following depths: 13, 25, and
38 cm (5, 10, and 15 in), using a steel-stem,
dial-type thermometer. Soil temperatures were
taken in all stands during a two-week period in
mid-August 1968.

Following drying, the portion of soil samples
passing a 2 mm sieve was analyzed for the fol-
lowing characteristics: (1) texture, (2) bulk
density, (3) pH, (4) exchangeable calcium, and
(5) "available" phosphorus. Soil extracts for de-
termination of the nutrient elements were pre-
pared by washing 3 g of sieved soil with 60 ml
of 0.2 N acetic acid for one hour. Calcium was
measured by flame photometry (Jackson, 1958)
and phosphorus by spectrophotometry ( Golden-
berg and Fernandez, 1966). The results are ex-
pressed as mg of substance per g of soil.

Soil texture was analyzed by a hydrometric
method as described by Bouyoucos (1928). The
pH was determined by preparation of a slurry
consisting of a soil-water mixture in a ratio of
1:1. Readings were made with a glass electrode
pH meter.

The weight of a standard volume of rock-free
soil was used to compute the bulk density of the
various horizons of the soil profiles. When aver-

Brigham Young University Science Bulletin

age values for whole profiles were calculated for
pH, phosphorus, calcium, and texture, values for
individual horizons were weighted by horizon

depth (reduced by a quantity equal to the per-
cent volume of rock in the horizon) and bulk
density.

Table 2. Prevalent species list. The letters in parentheses following the scientific names designate the "life-form"
class of the species. Life-form abbreviations are as follows: T, tree; S, shrub; TF, tall forb; SF, short forb;
AF, annual forb; and G, graminoid.

Species

Common Name

Pres.

%

Avg.
Freq.

P.xF
Index

1. Stelkiria jamesiana (SF)

2. Brornus polyanthus (G)

3. Lathynis lanszwertii (SF)

4. Gidium bijolium (AF)

5. CoUomUi linearis (AF)

6. Ncmophilci hrcviflora ( AF )

7. Collinsiu purviflora (AF)

8. Ehpnus glaucus (G)

y. Osmorhiza chilensis (SF)

10. Agropyron trachycaulum (G)

11. Polygonum douglasii ( AF )

12. Vicia auiericuna (TF)

13. Descurainia californica (AF)

14. Poa carta (G)

1.5. Thidictrum fcndleri (TF)

16. Viok nuttallii (SF)

17. Taraxacum officinale (SF)

18. Poa pratensis (G)

19. Chcnopodiuni frcmontii (AF)

20. llackelia florihunda (TF)

21. Achillea millefolium (SF)

22. Valeriana occidenlalis (SF)

23. Populus tremuloidcs (T)

24. Symplioricarpos orcophilus (S)

25. Phacelia hcterophylla (SF)

26. Rudhechia occidentalis (TF)

27. Mertensia arizonica (TF)

28. Srnilacina stellata (TF)

29. Hydroplujllum capitatum (SF)

30. Aster engelmannii ( TF )

31. Silcne menziesii (SF)

32. Pachiatima myrsinites ( S )

33. Stipa Columbiana ( G )

34. Berheris repens ( S )

35. Carex occidentalis (G)

36. Agastache urticifolia (TF)

37. Senecio serra (TF)

38. Delphinium occidentale (TF)

39. Senecio uintahensis (SF)

40. Osmorhiza occidentalis (TF)

41. Melica bulhosa (G)

42. Floerkea proserpinacoides (AF)

43. Gayophytum ramosissimum (AF)

44. Ueraclcum lanatum (TF)

45. Ilelenium hoopesii (SF)

46. Polcmonium foliosissimum (TF)

47. Erysimum argillosum (SF)

48. Ligusticum porteri (TF)

49. Castilleja rhcxifolia ( TF )

50. Bromus ciliatus {(.',)

51. Epilobium angustifolia (TK)

52. Aconitinn colundiianum (TF)

53. Scrophuhria bnceolatus (TF)

Tuber Starwort

80

64

5120

Mt. Brome

84

59

4956

Peavine

78

56

4368

Twinleaf Bedstraw

78

55

4290

Collomia

61

59

3599

Nemophila

51

67

3417

Blue-eyed Mary

63

48

3024

Blue Wildrye

69

43

2967

Sweetroot

73

39

2847

Slender Wheatgrass

71

39

2769

Douglas Knotweed

69

38

2622

American Vetch

67

36

2412

California Tansymustard

67

36

2412

Bluegrass

57

37

2109

Fendler's Meadow Rue

78

27

2106

Nuttall Violet

57

36

2052

Dandehon

61

32

1952

Kentucky Bluegrass

51

35

1785

Fremont Goosefoot

74

24

1776

Stickseed

63

26

1638

Yarrow

84

19

1596

Western Valerian

41

36

1476

Aspen

100

14

1400

Mt. Snowberry

78

17

1326

Vari-le:xf Phacelia

49

23

1127

Niggerhead

61

18

1098

Arizona Bluebell

33

33

1089

Starry False Solomon Seal

31

34

1054

Waterleaf

55

18

990

Engelmann Aster

45

21

945

Menzies Silene

37

25

925

Mt. Lover

16

54

864

Columbia Needlegrass

43

17

731

Oregon Grape

29

24

696

Western Sedge

42

16

672

Nettleleaf Gianthy.ssop

55

12

660

Butterweed Groundsel

41

16

656

Western Tall Larkspur

41

16

656

Uintah Groundsel

12

51

612

Sweet Anice

43

12

516

Onion Gra.ss

45

10

450

False Mermaid

14

31

434

Groundsmoke

12

32

384

Cowparsnip

22

17

374

Sneezeweed

14

27

378

Leafy jacobsladder

41

8

328

Wallflower

20

14

280

Porter Ligusticimi

8

32

256

Sphtleaf Painted Cup

6

40

240

I'^ringed Brome

14

17

238

l'"irewced

14

14

196

Monkshood

10

18

180

Lanceleaf Figwort

22

8

176

Biological Series, Vol. 16, No. 2 Aspen in Utah

LABORATORY METHODS

Plant Species

All species of the study were ranked in des-
cending order based on a presence X frequency
(P X F) inde.x (Anderson, 1954; and Curtis,
1959) utilizing percent presence in the several
stands and average frequency in stands of oc-
currence in the 0.25 m- quadrats. It should be
noted that this method of calculation departs
from the method used by the foregoing authors
\\'ho based average percent frequency on all
stands and not on stands of occurrence only.
However, by basing average frequency on stands
of occurrence only, the index becomes a proba-
bility statement of the likelihood of finding a
species in any quadrat placed within the spec-
trum of stands sampled. This advantage was the
motive for the change in manner of calculation.
Species selected for detailed study were those
with the highest P X F indices. A total of 53
species were used in the initial analyses.

The occurrence of positive or negative asso-
ciation among 42 prevalent understory species
was evaluated by chi-square analysis of joint and
independent occurrences of all possible combina-

tions of species in the 1,225 quadrats used to
sample the herb layer of the 49 stands.

Plant nomenclature follows Holmgren and
Reveal (1966).

ENVmONMENTAL FACTORS

The following factors were tested for environ-
mental differences among stands grouped ac-
cording to site-quality of aspen: (1) slope, (2)
aspect, (3) altitude, (4) soil temperature at
three depths, (5) values for pH, calcium, phos-
phorus, sand, silt, clay, and bulk density in the
Ai horizon, and (6) average values for pH, cal-
cium, phosphorus, sand, silt, clay, and bulk den-
sity for whole profiles. These same environmen-
tal factors were correlated with each other in all
posible combinations by simple statistical corre-
lation to determine the interrelatedness of the
several enviromnental variables (Warner, 1971).

Data Analysis

All analysis was made by computer, using
programs designed and tested by Dennis Powell,
University of Utah.

RESULTS

Stand Distribution

The stands are distributed over a range of
more than 5 degrees of latitude ( from 36° 13' at
the North Rim of the Grand Canyon to 41° 45'
in Logan Canyon, Utah). Elevation varies from
2,059 to 2,867 m (6,750 to 9,400 ft) above sea
level. Average elevation for all stands is 2,500
m (8,198 ft). Stands were selected in six national
forests and one national park (Table 1).

Prevalent Species

A list of the 53 most important species in the
study is shown in Table 2; species are listed in
decreasing magnitude of the P X F indices. An
average of 36 vascular plant species was found
per 0.04 ha stand.

The species listed in Table 2 include but a
single tree (aspen) and only three shrubs, of
which snowberry (Stjmphoricarpos oreophilus) is
the most common. Tlie remaining species include
40 forbs (23 percent of which are annuals) and
9 graminoides. Roughly two-thirds of the
perennial forbs grow to a height in excess of 30
cm (1 ft). On the basis of the P X F index,

which may be viewed as a kind of ubiquity in-
dex. Table 3 demonstrates that the flora of the
aspen type is dominated by forbs ( almost 75
percent of the index sum). Shrubs are unimpor-
tant, and grasses and grasslike plants contribute
only about one-fourth as much as forbs to the
sum. The large contribution of annuals appears
to be related to excessive grazing pressure from
domestic animals. Stands grazed only by deer
rarely support an abundance of annuals. The an-

Table 3. Contribution of various life-form group.s to
the sum of the P x F indices of the .53 species listed
in Table 2.

Sum PxF

%of

Group

Indices

Total

Tree

1,400

1.7">

Shrubs

2.886

3.6

Perennial forbs ■

■ tall (>30 cm)

14,580

18.0

Perennial forbs

- short ( <30 cm

) 23,723

29.2

Annual forbs

21,958

27.0

Graminoides

16,677

20.5

Total

81.224

100.0

'The tree layer is dominated by aspen, but this value considers
rpi( urrenrcs of aspen stems in ihe herb-layer quadrats nnly.

8

Bricham Young UiVivERSiTY Science Bulletin

nuiils involved
Mountain West.

xppear to be native to the

Species Association bv Chi-square

A number of coefficients have been devel-
oped to measure the degree of association be-
tween species based on presence-absence data.
Tiiese have been vi'idely used and adequately re-
\iewed by Cole (1949, 1957) and Hurlbert
( 1969 ) . For the purposes of this study, it was
considered desirable to be able to identify spe-
cies which were either positively or negatively
associated to a significant degree; but the degree
of association was not considered essential. Ac-
cordingly, significant associations among all pos-
sible combinations of understory species were
detennined by chi-s(|uare analysis of joint and
independent occurrences of the specified species
pairs in the 1,225 quadrats taken in the herb
layer. The results are reported in Fig. 2. This
figure shows in matrix fomi the relationship be-
tween species, the level of significance of the
chi-square values, and whether a relationship is
positive or negative.

It is possible to graphically summarize the as-
sociation data in many ways. Kershaw (1964)
demonstrates several techniques by which spe-
cies "constellations" based on significant positive
associations only can be constructed. Sokal and
Sneath (1963) discuss numerous ways in which
entities may be "clustered" using all values of
interspecific association. We have drawn associa-
tion "constellations ' as suggested by Kershaw
( 1964) and considered various "clustering" tech-
niques. The pair-group clustering method based
on simple averages (see Sokal and Sneath, 1963)
was finally adopted, since it has the theoretical
advantage of employing all association values
( negative and positive, significant and nonsignif-
icant) in construction of the association dia-
gram. Furthermore, its synthesis of the data
seemed to square with field observations better
than those based on other techni((ues.

As shown in Fig. 3, the 15 .species on the ex-
treme left form a highly associated group of an-
nual and perennial herbs. The group includes the
most common species of the array of stands
sampled and seems to represent those under-
story species that distinguish modal aspen areas
in the region sampled. Many of these .species are
also known (Hutchins, 1965) to be increasers on
heavily grazed aspen areas. The nine species
at the extreme right in the figure are known, on
the basis of our own unpublished work, to be
most abundant in aspen forests that are prone to
rapid displacement by conifers. Species inter-
mediately positioned between these two extreme

groups are apparently characteristic of special-
ized microhabitats of aspen forests. Within that
group, dandelion and Kentucky bluegrass are
perennial exotics that invade moist, disturbed
sites. Yarrow and Oregon grape also occur on
such sites.

Fringed brome and Columbia needlegrass
were positively associated to a significant degree
with each other but with no other species. This
pattern is related to the fact that only those spe-
cies among the 42 considered in Fig. 3 were
abundant in the six Arizona stands.

Figure 2. As.sociiition by chi-square among 42 impor-
tant understory species in aspen forests of Utah and
northern Arizona. The summ.ition chi-s(]uare vakie
for the association test of each species pair is re-
ported in die bo.x formed by the intersection of the
row and cohimn of the two species compared. Neg-
ative as.sociations are designated by a minus sign.
Significance levels are as noted in the figure. Spe-
cies abbreviations appear below.

Species

Achillea millefolium
Agu.stache urticifolia
Agro/)i/ron truchiji.nihim
Aster engchnunnii
Beiheris repens
Bromus pohfanthus
Bromiis ciliatus
Ctirex occidentali.'!
Chenopodium frcmonlii
Collinsia pariiflora
Ci)llomi:i linearis
Delphinium oecidentale
Deseurainin calif arnica
Eh/mus glaueus
(.'((/(i(»i hifolium
Iluehcliii florihundu
IIildrDphiilluni cujiitatum
Ldthi/ru.s lanszwertii
Melica Imlho.sa
Mertennia erizonieti
Nc7nophila hrcvifloru
Osnunh 'za chilen.si.-i
O.-inuirhizj oce'dentali.i
Piichi.'.am I myrsinUci
Phjccli'i hclerophijUa
l'i>a curia
Poa pratenf:i.\
Pohi<^i>nu)n dougla.sii
Riitlheekia occidcntati.i
Seneeit) serra
Senecio uintalti'n.si.t
Silene menzi<:'iii
Smihicina .■itellat i
Stclliria p.nnesianu
Slipa colinnhianii
S}jmphoiic;iry.()s oreophilus
Tariixaeum officin(dc
I halietrum fendleri
V'tlcrian'i occidcntulus
Vici'i amerieiinn
Viohi uutlallii

.'Viibreviation
Ala
Aur
Atr
Aen
Bre
Bca
Bci
Coc
Cal
Cpa
Ch
Doc
Dca
Egl
Gbi
Hfl
Hca
Lla
Mbl
Mar
Nbr
Och
Ooc
Pmv
Phi'
Pen
Ppr
Pdo
Roe
Sse
Sui
Sme
Sst
Sja
Sco
Sor
Tof
Tfe
Voc
Vam
Vnu

Biological Series, Vol. 16, No. 2 Aspen in Utah

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Brigham Young Unwersitv Science Bulletin

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X'- Value

Figure 3. Results of the "cluster" analysis of interspecific association values among 42 species. Species abbrevia-
tions are as in Fig. 2, except for Achillea inillcfuliurii (Ami), Bivmus polyuutlius (Bpo), and Mclicu hiilhosa
(Mbu).

Biological Series, Vol. 16, No. 2 Aspen in Utah

11

As noted earlier, species "constellations"
based on significant positive associations were al-
so drawn utilizing indices of interspecific asso-
ciation obtained by correlation and chi-square
analysis (Snedecor and Cochran, 1967). See
Warner ( 1971 ) for results of these analyses.

SlTE-QUAUTi' DETERMINATIONS

Site-quality was determined for 43 stands, us-
ing the natural site index curves of Jones
(1967b). The 6 stands in Northern Arizona
were not used in this phase of the study because
of basic compositional differences in the under-
story. Almost none of the dominants of those
stands corresponded with dominants of the Utah
stands. An index value was obtained for each of
the stands used by extrapolation from the nat-
ural site index curves which are derived from
height-over-age at breast-height data. Height and
age values used in this study were the averages
for height and age (in years) of the 10 tallest
trees in each of the stands tested (Warner,
1971). Using these values, stands were ordered
according to Jones's site index.

Forest characteristics of the several stands
are shown in Table 4. Excluding the Arizona
stands ( which are not used in site-quality analy-
ses), average tree density is 1,005 per ha (407/
acre); aspen reproduction (individuals having
dbh less than 10 cm) density averages just over
4,200 individuals per ha ( 1,707/acre). Average
stand age (based on the 10 tallest trees per
stand) is 74 years, with a range running from 42
to 111 for individual stands. Average site index
is 15.8 m (52 ft) at 80 years of age.

In order to identify understory species signif-
icantly associated with stands of high or low site-
quality, average quadrat frequency of understory
species of the top 16 stands on the site-quality
gradient, based on Jones's index, was tested
against the average frequency of the same spe-
cies in the last 16 stands of that gradient. Aver-
age (juadrat frequency was calculated for each of
the 5.3 herbaceous species in those two groups of
stands. Of those species, 15 were found to be
significantly (at the .05 level or better) more or
less abundant on the better quality aspen sites
(Table 5).

An index value was calculated for each of
the 15 indicator species. The derivation of this
index was as follows : ( 1 ) the mean difference
between quadrat frequencies of all species on ex-
cellent as opposed to poor site-quality stands was
used to rank the 15 species, (2) all of the posi-
tive differences were divided by the smallest
positive difference, (3) the same procedure was
used for the negative differences, and finally.

(4) all values were adjusted so that there were
no negative values (Table 5).

The aspen stand site indices were then de-
termined for each stand using such indicator spe-
cies as occurred in the stand and the following
equation:

Understory site index =:

2 (species index value x species % frequency)

i; % frequency of indicator species

Table 4. Stiinc! chunicteristics of the 49 forests sampled.

Aspen Understory

Aspen Repro- Produc-

Stand Trees duction tion Avg. Age Avg. lit. Jones

No. Ind/ha Ind/ha kg/ha (years) (m) Index

1

1458

1483

374

47

12.5

52

2

.544

6259

649

86

13.1

43

3

815

12190

489

111

20.7

57

4

1334

4119

763

68

14.3

52

5

840

4945

720

66

14.6

56

6

1112

9392

2.35

60

12.5

50

7

741

1483

2237

62

17.1

66

8

519

4942

1354

83

18.3

59

9

815

2308

16.58

75

21.0

70

10

568

7416

1246

67

14.6

54

11

791

2634

911

63

15.2

58

12

914

1814

657

78

15.5

53

13°

2372

.329

29

70

12.5

44

14'

1483

1317

1.32

75

23.8

70

15°

544

2471

NA

85

16.5

53

16°

1359

329

75

66

15.2

57

17°

1112

11.54

98

67

16.8

60

18°

1063

166

80

86

18.3

57

19

1211

494

500

75

8.5

30

20

865

329

1221

83

16.8

53

21

815

1646

879

42

9.4

48

22

717

1648

331

94

15.2

46

23

593

1979

175

73

12.2

40

24

1581

0

582

78

14.6

48

25

618

3128

659

71

15.8

56

26

1878

329

776

61

12.2

45

27

NA

NA

NA

59

8.8

35

28

NA

NA

NA

65

9.1

34

29

1211

3625

558

80

19.2

62

30

1260

7744

745

78

18.0

60

31

1211

6096

416

96

18.3

54

32

1112

10791

NA

75

11.6

37

33

1236

988

418

100

19.5

58

34

766

4942

823

98

20.7

60

35

1606

9224

525

68

14.0

50

36

840

5105

462

74

18.3

60

37

10.38

5108

409

88

18.6

58

38

.544

3128

379

67

11.6

42

39

1236

.5271

429

66

11.9

43

40

988

4611

188

84

14.3

46

41

1754

1317

1699

52

17.7

75

42

1532

8402

381

69

1,5.5

55

43

964

2965

392

80

17.4

58

44

1137

1.3.344

.321

63

12.2

46

45

939

494

693

76

16.2

55

46

272

3128

399

95

16.5

48

47

840

3951

NA

47

8.5

37

48

NA

NA

NA

67

14.9

54

49

NA

NA

NA

69

18.3

66

•Stands not used in site-quality analyses.
NA = Data not available.

12

BwGHAM Young University Science Bulletin

Table 5. Comparative frequency of iinderstory .species in aspen stands of contra.sting height growth. Only species
of significantly different abundance in the two groups are listed. Derivation of the indicator index is also
demonstrated.

Average

Frequency

Indicator

16 High Site-

16 Low Site-

Indicator Index''

Species

Quality Stands

; Quality Stands

Difference

Inde.x"

(transformed)

Elijmus ^laiiais

46

23

23

4.68

8.3

Lathijrus lanszwertii

54

33

21

4.47

8.1

Viola nuttaUii

28

7

21

4.42

8.0

Poa curt a

27

11

16

3.37

7.0

Merteiisia arizuitka

16

2

14

2.84

6.5

Floe rkca p rose rpinaco ides

11

0

11

2.26

5.9

Thalictrum fendlcri

19

11

8

1.74

5.4

Heracleiiin lanatum

7

0

7

1.53

5.2

Helenium hoo])esii

6

0

6

1.21

4.8

Acouituiii coliimbiumati

5

0

5

1.00

4.6

Gaijoplujlum ramosissimum

0

9

-9

-1.00

2.6

Sijnij)horicarj>os oreophilus

7

16

-9

-1.03

2.6

Aster cni:^leiiiannii

3

16

-13

-1.30

2.3

Parhistiina mijrsinites

4

20

-16

-1.76

1.9

Polygontiin duugUisii

16

40

-24

-2.62

1.0

a Positive differences were divided by the smallest positive difference anJ llie nepative differences by the smallest negative difference,
b The largest negative index was adjusted to a value of +1.00 and all niher indues were adjusted to the new base by adding the factor used
to adjust the largest negative index.

Table 6 illustrates the method of computing the
understorv site-index, using; stand 34 as an exam-
pie.

The degree of correspondence between
Jones's natural site-index values and the under-
story site-index values for the 43 stands was
evaluated by simple correlation. The correlation
coefficient for the test is +0.70. The r-value is
significant at the 1 percent level. The coefficient
of determination ( r- ) for this relationship is 0.49
Thus approximately half of the variation in
height growth can be accounted for by the linear
regression of Jones's natural site index on under-
story site index. Figure 4 illustrates the results.

■The use of simple presence of indicator spe-
cies, rather than frequency values, was also
tested. If site indices could be predicted with
reasonable accuracy using simple occurrence of
indicator species, sampling and associated ex-
penses in time and money could be saved. In
this case, site values were determined using the
following formula:

Stand index =

2 index values for indicators present

i indicator species present

The degree of correspondences between
Jones's natural site-index values and the under-
story site-index values based on indicator pres-
ence only was also evaluated by simple correla-
tion for the 43 stands. The correlation coeffi-
cient for the test is +0.64 and is significant at the

one percent level. The coefficient of determina-
tion (r-) for this relationship is 0.41. The r- for
indices based on fecjuency is thus slightly higher
than that for indices based on presence data
only, but the difference is not great. In view of
the greater simplicity of the latter method, one
may be willing to accept slightly less precision
in estimates in exchange for savings in labor and
time.

The average species composition of low, in-
termediate, and high site-quality stand-groups

Table 6. Calculation of thi'

urulers

torv sit(

■ index for

aspen stand .34.

Index

Species Species

Species

Value

'i Fre<:

|. Index

Eli/mii.s f^liiticus

8..3

80

664.0

Latlii/ni.s liinszitertii

8.1

88

712.8

Viohi nultnllii

S.O

100

800.0

/'(Kj lurtti

7.0

32

224.0

Mcrtcnsia iiriznitica

6. .5

16

104.0

ylocrkea jnoscrpuiaandt s

5.9

(1

0

'rhalictnmi fcntllcri

.5.4

48

2.59.2

Heriiclcuin liiuatum

5.2

0

0

llcUnium luKiprsii

4.8

8

.38.4

Acoiiittmi colmuhkinuni

1.6

0

0

Cui/cphi/tuin luinosix.siinuin

2.6

0

0

Siiiiiphoricarpus oreophilus

2.6

4

10.4

Aster cn<^chnannii

2.3

0

0

I'ucliixtimd mijrsinites

1.9

0

0

Fiiliiilonurn (/owf^/rwii

1.0

0

0

Total

376

2812.8

■ 1 — S(Species index value -\

species %

f"-")'- 7,,«

S % frequency of all indicator species

Biological Series, Vol. 16, No. 2 Aspen in Utah

13

according to Jones's index is shown in Table 7.
It can be seen that several species have similar
average frequency in all site classes. Bromus
j)flhiatUhus, ColJomia linearis. CoUinsia parvi-
flora, Descttrainia californica, Hackclin floiihun-
da. Polemonittm foUosissimum, Senecio serra, and
Stcllaria jamcsiana are representative of such

Table 7. Average understorv' composition of low, in-
termediate, and high site-quality-stand groups.

Average Frequency

Species

1 (> High 11 16 Low

Site- Intermediate Site-

Ouality Site-Qualily Onalitv

Stands Stands Stands

Achillea millefolium
Acotiitum cotumhianurn
Agustuchc urticifulia
Agropt/ron traclu/cauhim
Aster cngelmannii
Berberis repens
Bromus ciliatus
Bromus poh/anthus
Carex occidentulis
C (1st ill c III rh ex i folia
Clienopodiuin fremoiitii
Collomia linearis
Colliiisia piirviflora
Delphinium oeeideutale
Descurainia ealiforiiiea
Eh/mus glaueus
Epilohium august ifoliiirn
Ert/simiim argillosum
Ftoerkca firoserpinacoides
Qaliuui hifolium
Gai/oplu/tum lumosissimum
llachelia floribunda
Helenium hoopesii
lleraeleum hinatum
Ihjdrophiillum capitate m
Latlu/rus lanszwertii
Ligusticum porteri
Melica hulhosa
Mertensia arizonica
Neuuiphila hreiiflora
Osmorhiza ehilcnsis
Osinorhiza oecidcutalis
Paeh ist inui rni/rsin ites
Phaceliu heteroj)tu/Ua
Poa carta
Poa pralensis

Poleuuniiuni foliosissi>7iui)i
Polt/goniun douglusii
Populus tremuloidcs
Rudbeckia occidcntalis
Scrophidaria lanceolata
Senecio serra
Senecio uiutahensis
Silenc menziesii
Smilacina stellata
Stellaria jamesiana
Stij)a Columbiana
Si/mphoricarpos oreophilus
Taraxacum officinale
Thidictrum fcndlcri
Valcrianu occidentulis
Vicia amcricana
Viola nuttallii

11

19

15

5

0

0

6

1

4

23

32

31

3

10

16

6

3

4

0

3

3

58

41

51

5

54

0

0

0

3

21

16

28

36

43

41

33

33

35

11

6

0

23

28

23

46

25

23

3

0

2

1

0

4

11

1

0

.54

53

40

0

0

9

14

18

10

6

7

0

7

1

0

8

23

6

.54

47

33

6

0

0

4

8

2

16

19

2

42

43

29

40

20

24

6

6

4

4

0

20

8

7

13

27

31

11

17

10

12

1

1

1

16

22

40

14

15

11

13

3

8

0

1

1

6

3

5

7

0

11

8

8

12

12

5

8

55

49

51

3

6

3

(

12

16

18

8

22

19

16

11

15

20

6

25

36

23

28

30

7

rather unifomily distributed species. Other spe-
cies appear to prefer stands of intermediate site-
quality. Such species as Achillea millefolium,
Carex occidentalis, Hijdrophijlliim capitatiim,
Valeriana occidentalis, and Vicia amcricana are
representative of this latter group. Species that
are most abundant in stands of high or low site-
(juality have already been identified in Table 5.
Perhaps the most striking thing about under-
story composition in the three site-quality
groups of Table 7 is the floristic uniformity that
exists among groups.

Understory production (current year, above-
ground gro\\'th to a height of 6 ft ) is significantly
correlated with Jones's site-quality index for the
42 stands for which production data are avail-
able (r =: +0.45). Average understory produc-
tion in these stands was 621 kg/ha (554 lbs/acre)
with the range running from 29 to 2,237 kg/ha.
As shown in Table S, the contrast in production
between selected stands of low as opposed to
high site-quality is marked. Also, stands of low
site-quality are relatively more shrubby than tall
stands. On the average, understories of tall
stands tend to be more heavily dominated by
forbs than do those of short stands; but indi-
vidual values were too variable in our sample to
yield statistically significant differences.

Tree and tree reproduction density values re-
ported in Table 8 would seem to indicate that
the greater understory production of tall stands
is not attributable to lower stocking rates and
consequent better lighting of the forest floor.
Tree and reproduction densities do not differ
significantly in the two stand groups.

Table 8. Comparison of average values for various
vegetational parameters from the 7 shortest as op-
posed to the 7 tallest aspen stands for which data
for all parameters are available. Stand numbers of
the short stands are: 2, 13, 19, 23, 26, 38, and 39.
Tall stands used are numbers 7, 9. 29, 30. 34, .36,
and 41.

Parameter

Site Quality Group
Short ■ Tall

Understorv production (kg/ha)
% composition of imderstory
(by wt)

Shrubs

Graminoides

Perennial forbs

Annual forbs
Aspen trees (no. /ha)
Age of 10 tallest trees (years)
Tree site quality (Jones Index)
Aspen reproduction (no. /ha)

419

1169"'

19.7°

4.2

35.5

36.4

44.7

59.2

3.7

2.8

1196

1055

71

74

41

65"

2540

3788

'Valiio is significantly (0.05 level or better) larger than that with
which it is paired

14

Bhigham Young University Science Bulletin

Environment

Mean values were calculated for various en-
vironmental variables in the two stand groups
previously used for the identification of herba-
ceous species associated with high or low site-
(|uality aspen stands (Table 5). As shown in
Table 9, only two environmental variables (soil

reaction and exposure) differ significantly be-
tvveen the two stand groups. Better sites were
consistently more acidic than were sites of low
cjuality for aspen. Exposure was shown to differ
at the 0.1 level of significance only. In general,
the better sites had an easterly exposure, while
poor sites were more common on westerly ex-
posures.

DISCUSSION

The desire to predict site-qualty for aspen
using understory species occurring on the sites
stems from the fact that aspen is a fire-follower.
As such, the species may become dominant on
large acreages previously controlled by another
forest type. If the tree itself is to provide the
estimate of site potential for aspen on such areas,
one must wait from two to three decades before
an accurate estimate is possible ( Graham, Harri-
son, and Westell, 1963). Obviously, better man-
agement alternatives would be possible if a pre-
diction of potential tree production could be
made earlier.

Predicting aspen site-cjuality from under-
story vegetation may also prove easier than us-

ing tree height-over-age techniques as predictors.
Aspen increment cores are normally difficult to
age and require mounting, drying, sanding, and
staining before accurate counts are possible
(Alder, 1969). Prediction based on understory
vegetation would require only field samples of
relative simplicity. Furthermore, predictions
based on understory species can reveal much
about (juality and cjuantity of forage in the forest
understory. Predictions based on understory
parameters could thus have value for multiple-
use decisions.

The test of validitv of the index developed
in this study admittedly includes a strong ele-
ment of circular reasoning. Inasmuch as 32 of

Table 9. Environmental characteristics
on 16 stands in each group.

Characteristics

of stand groups ot contrasting site quality for aspen. Averages are based

Low Site-
Quality Group

High Site-
Quality Group

Significance of
Difference

Altitude (feet)

Slope (%)

Exposure''

PO," (mg/g)

Ca-" (mg/g)

pH^

bulk den.'' (g/cc)

Sand (%)"

Clay (%)"

Silt {%)'•

PO, (A,) (mg/g)

Ca- (A,) (mg/g)

pH (A,)

bulk den. (A,) (cc/g)

Sand {%) (A,)

Clav (%) (A,)

Silt'(f) (A,)

Soil temp. (5" depth °F)

Soil temp. (10" depth °F)

Soil temp. (15" depth °F;

8283

20.5

.72

.250

3.62

6.02

1.31

32

33

35
.026
3.55
6.20
1.10

36

24

40

.50.7

46.S

46.1

8115

20.8
1.39
.208
5.54
6.13
1.32

30

33

37
.038
3.50
5.60
1.05

35

25

40

47.6

46.0

45.6

n.s.
n.s.

o

n.s.
n.s.
n.s.
n.s.
n.s.
n.s.
n.s.
n.s.
n.s.

o O

U.S.

n.s.

n.s.
n.s.
n.s.
n.s.
n.s.

a F.vposure values arc a sine transformation of direction of slope in azimuth dcfiiees dnckwisc froni ninth using the foi nulla of Beers, Dress,
and W'ensel (HHifii, This fornuda pives southwest expd-.iuos a \f\\iw of zero .Tutl those of iioitheasl 2; all others receive interniediule
values,
1> Mean value for entire soil profile: Olitainod by weiehting values fo scp.n.ite hoii/inis Iiv hoii/on ilepth.
■ = Significant at 0,1 level.
•■ = SignifiraiU at (),ni level.

Biological Series, Vol. 16, No. 2 Aspen in Utah
80 1

75 J

15

41

70

•9

65 -

.49
7

60 -
55

■20

43.3,-3^.^33

50 -
45

2

44.^^22^/^

^^^ 6 35
/^l 46

24

26

39 -36

40 -

23

47

35 -

^^^28

32

27

30 -

' 1 ' 1

-^^^^

1 ' ; ' I ' 1

Understory Index

Figure 4. Computed regression line of Jones's Inde.x on the Understory Inde.x. Dots represent individual stand
locations. Equation for the regression line is: Understory Inde.v = 4.62 (Jones's Index) + 25.96.

the 4.'3 stands used in the vahdity test illustrated
in Fig. 4 were previously used to identify indi-
cator species, a strong positive correlation be-
tween Jones's index and the understory index is
to be expected. Ideally, the derived index should
have been tested on an independent sample of
aspen forests from the same region. Unfortu-
nately, such a set of data was not available.

Several different lines of evidence suggest,
however, that the understory index does have
real validity for Utah aspen forests. One such
evidence stems from analysis of the interspecific
association values for understory indicators of
sites of high as opposed to low quality. Ten of
the 15 indicators identified in Table 5 appear in
the matrix of interspecific association values
shown in Fig. 4. Floerkea prose rpinacoides, Hera-
cleum lanattim, Aconitiiin columbianum, and
Gaijopliytum ramosissiinum were omitted from
Fig. 4 because all occurred with low frequency,
even in stands of occurrence. Six of the 10 indi-
cators represented in Fig. 4 are indicators of bet-
ter sites, while the other 4 are indicative of poor
sites.

Inasmuch as the association data (Fig. 2)
are based on presence or absence in individual
quadrats of all 49 stands, those results are, at

least partially, independent of responses reported
for the same indicators in Table 5 which are
based on percent frequency data from only 32
of the stands. Yet, as shown in Table 10, indica-
tors of good sites still tend to hang together as
a group with 80 percent of the possible inter-
specific associations being significantly positive.
The four indicators of poor sites form a less
homogeneous group but still are significantly
positive in their association in one-third of the
possible comparisons. If, however, indicators of
good sites are compared with indicators of poor
sites, it will be noted that only 8 percent of the
comparisons are significantly positive, and 42
percent become significantly negative. Average
chi-square value for all possible comparisons
between species of the two groups drops to +3.2,
a value far smaller than that obtained when all
possible association values within groups are
averaged.

Another bit of evidence supporting the valid-
ity of the indicator species selected is furnished
by the index based on presence onlv of indica-
tors. Indicators were initially selected on the
basis of average frequency in stand groups of
contrasting site-quality. The fact that a highly
significant correlation exists between the Jones

16

Brigham Young University Science Bulletin

Table 10. Summary of interspecific association
Fig. 2.

s anion"

till' 10 indicator species ol Table 5 that also appear in

No. indicators stiidit-d

No. interspecific comparisons made

Avg. chi-square value for the interspecific

comparisons (from Fig. 2)
Yc chi-s(juare values that are

significantly positive
',c chi-s(juare values that are

significantly negative
C^oefficient of variation"

for the ave. chi-square value

for interspecific comparisons (%)

Interrelationships

among Indicators of:

High Low

Quality Sites Quality Sites

Interrelationships

between Indicators

of High vs. Low

Site Quality

6
15

4
6

10

24

+18.3

+ 11.4

+3.2

80

33

8

0

17

42

85

245

452

'Standard deviation \ 100

C.V.

index and the presence index indicates again that
the selected species do have real value as pre-
dictors.

A final evidence for validity of the indica-
tors is presented in Fig. 5. Here the understory
index is plotted against the Jones index for 15
stands. The 15 stands include the 11 intermediate

80 -1
75

70 -
65
60
55

'm
t>

c 50
o

— >

45 -
40-

1)

c

35 -

30

•28

.19

stands of Table 7 (none of which were used in
identification of the indicator species) and the
two tallest and the two shortest stands shown in
Fig. 4. Thus, 73 percent of these stands provide
independent evidence concerning the validity of
the index. The two tall and the two short stands
were added to the analysis to lend length to the

.41

•45

43.3.11

25-

.31

•1 'li

24

.46

•38

UncJerstory Index

Figure 5. Correspondence between tlu' Jones's Index and the Understory Iiide.\ based on 1.5 stantls, 11 of which
were not used in the original selection of indicators.

Biological Series, Vol. 16, No. 2 Aspen in Utah

17

height gradient under te.st, since it is difficult
to demonstrate significant trends when the range
of variation involved is small (range for the If
intcnnediate stands equals f6 units on the Jones
index). Adding the two extreme stands from each
end of the height gradient represented in Fig. 4
is actually less "loaded" than might appear at
first glance. Indicators were selected on the basis
of stand groups, not individual stands. Hence
the contribution of the tallest and the shortest
stands was no more or less than that of other
stands in the t^vo groups. Utilization of the tal-
lest and the shortest stands in the new test of
utility of the indicator approach thus seems no
more objectionable than would the use of an
equal number of randomly selected stands from
the two stand groups.

The results of the foregoing test of correla-
tion between the Jones and understory indices
give an r- value of 0.79 ( highly significant ) . Thus
the understory index for these stands accounts
for roughly 62 percent of the variation in the
Jones index for the same stands. This is an even
stronger correlation than was obtained in the
original test of the understory index ( Fig. 4, r-
value of 0.70), in which almost 7.5 percent of the
stands had been used in the initial selection of
indicators.

These results indicate that one can expect the
understory index as here calculated to consistent-
ly predict 50 percent or more of the variation in
site potential for aspen growi:h in Utah. Further-
more, there is reason to believe that the index
can be refined to pemiit even better predictions.
The aspen stands on which the present study is
based were not initially sampled for the purpose
of clarifying relations between tree height and
understory vegetation. Also the study extended
into at least two somewhat distinct floristic
zones ( the Wasatch Mountains and the southern
plateaus). Accordingly, many species of poten-
tial worth as indicators were rejected as being
statistically nonsignificant because they occurred
in only a few of the stands sampled. By select-
ing numerous stands of markedly different site
potential within a given floristic zone, one should
be able to increase the number and reliability of
indicators and thus produce a more precise in-
dex. Perfection of a technique for stratifying
stands according to the degree of grazing dis-
turbance should also permit more accurate esti-
mates (Harper and Hutchins. 1965).

Other attempts at predicting site-quality for
aspen from understory vegetation have been
made by Kittredge (19.38) and Graham, Harri-
son, and Westell (196-3). Both studies consider
aspen forests of the Great Lakes area. Kittredge's
"natural community plant-indicator" index was

relatively efficient as a predictor of site-quality
for aspen (r ^ 0.76), but its use required a
highly technical knowledge of the ecology of all
the major forest types of northern Minnesota and
Wisconsin. Graham, Harrison, and Westell
(1963) report that in Michigan sweet fern
{Comptonki peregrina) is common on sites of
low quality for aspen, while height growth of
bracken feni {Pteridiurn aquilinum) is positively
correlated with increasing site-quality. Never-
theless, they conclude that prediction of site-
(juality from individual understory species is too
imprecise to be useful. Kittredge (19.38, p. 238)
came to a somewhat similar conclusion for indi-
vidual species, saying, "On the whole, the con-
clusion seems justified that individual species do
not characterize the site index of aspen specifi-
cally enough to be used for prediction with any
degree of confidence" and that "indicator value
of individual species is distinctly less than that of
groups of species." Data from the current study
seem fully compatible with this view.

Jones (1967a), working in Colorado, Wyo-
ming, and New Mexico, has attempted to predict
site-quality for aspen using temperature and
moisture. His results show that the multiple re-
gression of average height of aspen at age 80 on
these two ordinates accounted for only slightly
more than .'30 percent of the total variation in the
site index. Working in the nomnountainous ter-
rain of northern Minnesota, Kittredge (1938)
was able to account for over 60 percent of the
total variation in site index using soil-profile
groups as the independent variable.

It seems apparent that methods utilizmg en-
vironmental parameters as predictors of site-
ffuality are not likely to be any more efficient
than indices based on understory vegetation. In
fact, the environmental data from this study
(Table 9) and those from Jones (1967a) would
seem to warn that predictive indices based on
environmental parameters will be very difficult
to perfect in topographically complex areas such
as characterize aspen forests in the Mountain
West. Certainly indices based on enviromnent
will be more time consuming and expensive than
indices based on understory vegetation. Under-
story indices tell one nothing, however, about
causes of variation in site-quality. For that rea-
son, there must be continuing attempts to iden-
tify potent environmental predictors of site-qual-
ity.

As Jones (1967a) and numerous others (see
Baker, 1921; Pauley, 1949; Cottam, 1954; Marr,
1961; Egeberg, 1963; Garret, 1964; and Strain,
1964) point out, marked genetic variation oc-
curs among aspen populations throughout North
.America. Such variation coupled with the clonal

18

Brigham Young UNrvERSixv Science Bulletin

habit is likely to place a rather low upper limit
on the amount of variation in height growth that
one can account for with any kind of predictive
technicjue. In Utah, it is not uncommon to find
large height differences between contiguous as-
pen clones of identical age in what appears to be
an essentially uniform environment. Obviously
such phenomena could inject considerable varia-
tion into analyses in which understory or envi-
ronmental indices are correlated with site poten-
tial for trees. Thus, in the absence of control of
tree genetics, one could never expect perfect
correlations between predictive indices and tree
height.

As Daubenmire (1952, p. .305, .32,3) has
pointed out, frequency values for understory
species are more likely to be altered by disturb-
ing influences such as grazing, logging, or fire
than is presence. Thus he uses presence in pref-
erence to frequency data in identification of
habitat-types (or site-quality classes) which are
intended to reflect basically different abiotic en-
vironments. In theory, species frequency is so
sensitive to disturbing influences that its use
could confuse rather than clarify understanding
of the abiotic system. Results of this study indi-
cate, however, that frequency data carry more
information content concerning site-quality (as
indicated by the coefficient of determination)
than does presence alone.

Data in Table 10 reveal an interesting char-
acteristic of the two groups of indicators em-
ployed in calculation of the understory index.
None of the indicators of good sites are signifi-
cantly negatively associated (only 1 of 15 pos-
sible comparisons is negative). On the other
hand, 17 percent of the possible comparisons
among indicators of poor sites are significantly
negative. Of the 6 possible comparisons, 3 bear
a minus sign. These data seem to suggest that
environmental conditions correlated with good
height growth of aspen are relatively uniform but
that dissimilar kinds of environments can cause
poor growth of aspen. How else can one account

for species being significantly correlated with
sites of poor growth for aspen, while being
significantly disassociated from each other? This
explanation seems easily defensible on theoret-
ical grounds.

A puzzling aspect of the interrelationships
among the indicator species reported in Table 10
is the fact that tvvo indicators of good sites (i.e.,
LatJnjrus lanszwertii and Poa pratensis) are
significantly positive in their association (Fig. 4)
with Polygonum dougkisii, an indicator of poor
sites. One is tempted to reject all three as indi-
cators under such circumstances, but the fact
that both L. lanszwertii and Poa pratensis are
significantly positive in their association with
four of the other five indicators of good sites in-
dicates that there is no a priori reason for doing
so. However, the indicator value of these species
should be viewed with caution until such time as
their status is clarified by independent data.

Both Poa carta and Polygonum douglasii are
known to be strong increasers on overgrazed as-
pen ranges of northern Utah (Hutchins, 1965).
Hutchins also shows that L. lansztcertii declines
slowly with moderate grazing, but more rapidly
as grazing pressure becomes more abusive. The
anomalous response described for these species
in the preceding paragraph may be caused by
differential grazing pressure among the aspen
forests sampled. One could postulate a system in
which Pohjgomtni doiighisii is largely confined to
sites of low quality when grazing pressure is
minimal. With abusive grazing, the species may
become abundant on sites of better quality. It is
possible that Poa pratensis, another increaser
with grazing, increases on both good and poor
sites under grazing but is best adapted to good
sites. L. lanszwertii, being moderately resistant
to grazing, could associate with Poa pratensis
and Pohjgonum douglasii on good and poor sites
alike but be most abundant on sites of good
quality. Clearly, a stratification of stands accord-
ing to grazing intensity would help clarify these
possibilities.

ACKNOWLEDGMENTS

This work was partiallv sujiported hv Public
Health Service Grant AP-00fi3I and the Inter-
mountain Forest and Uang(> Experiment Station
of the U.S. Forest Service. Wisconsin State
University paid the senior author's salary for one
academic vear while he was on education leave

from that institution preparing the disseration
on which this paper is based. We also thank
Keith Hutchins, Michael .\lder, Wes Revnolds,
Dennis Powell, and Fran Harner who assisted
in various wavs in collection of the data.

Biological Series, Vol. 16, No. 2 Aspen in Utah

LITERATURE CITED

19

Alder, Guy Michael. 1969. Age-profiles of aspen

forests in Utah and northern Arizona. Master's

Thesis, University of Utah, Salt Lake City. 31 p.
All.\n, ]ohn S. 1962. The plant communities of the

Big Cottonwood drainage. Master's Thesis, Univer-

sit\ of Utah, Salt Lake City. 108 p.
.\nderson, O. 1964. The phytosociology of dry lime

prairies of Wisconsin. Ph.D. Thesis, University of

Wisconsin.
B.\ker, F. S. 1918. Aspen as a temporary forest type.

J. Forestry 16:294-303.
. 1921. Two races of a.spen. J. Forestry 19:

412-413.

1925. Aspen in the Central Rocky Mountain

region. U.S. Dept. Agr. Bull. 1291. 47 p.

Beers, T. W., P. E. Dress, and L. C. Wensel. 1966.
Aspect transformation in site productivity research.
J. Forestry 64:691-692.

Bouyoucos. G. 1928. The hydrometer method for
making a very detailed mechanical analysis of soils.
Soil Sci. 26:2.33-238.

Cole, L. C. 1949. The measurement of interspecific as-
sociation. Ecol. .30:411-424.

. 1957. The measurement of partial interspe-
cific association. Ecol. 38:226-233.

Coles, F. H. 1965. The effects of big game and cat-
tle grazing on aspen regeneration. Master's Thesis,
Brigham Young University, Provo. 72 p.

Costello, D.'iviD F. 1944. Important species of the
major forage types in Colorado and Wyoming. Ecol.
Monogr. 14:107-134.

CoTTAM, Walter P. 1954. Prevernal leafing of aspen
in Utah mountains. J. Arnold Arboretum 35:239-
248.

. 1963. Quaking aspen {Popuhis tremuloides

Michau.x). Naturalist 14:3-15.

Crowther, Evan C, and K. T. Harper. 1965. Vege-
tational and edaphic characteristics associated with
aspen "strips" in Big Cottonwood Canyon. LUah
Acad. Sci. Arts and Lett. 42 ( ll):222-230.

Curtis, John T. 19.59. The vegetation of Wisconsin.
University of Wisconsin Press, Madison, Wisconsin.
657 p.

Daubenmibe, R. F. 1943. Vegetation zonation in the
Rocky Mountains. Bot. Rev. 9:325-393.

. 1952. Forest vegetation of northern Idaho

and adjacent Washington and its bearing on con-
cepts of vegetation classification. Ecol. Monogr.
22:.301-330.

. 1961. \'egetative indicators of rate of height

growth in ponderosa pine. Forest Sci. 7:24-34.

Driscoll, R. S. 1963. Production and floristic compo-
sition of vegetation as measures of site potential.
In Range Research Methods. U.S. Dept. Agr. Misc.
Publ. 940. 172 p.

Egebero, R. 1963. Inherent variation in the response
of aspen to frost damage. Ecol. 44:1.53-156.

Ellison, L., and W. R. Houston, 1958. Production
of herbaceous vegetation in openings and under
canopies of western aspen. Ecol. 39:337-345.

Fetherolf, J. M. 1917. Aspen as a permanent forest

type. J. Forestry 15:757-760.
Fowells, H. a, 1965. Quaking aspen {Popuhis
tremuloides Michaux), p. 523-534. In Silvics of
Forest Trees of the United States. U.S. Dept. Agr.,
Agr. Handb. No. 271. 762 p.

Gadnon, J. D., AND J. D. MacArthur. 1959. Ground
vegetation as an index of site quality in white
spruce planatations. Canada Dept. North. Affairs
and Nat. Res., Forestry Branch, Forest Res. Div.
Tech. Note, 70. 12 p.

Garret, Peter W. 1964. Clonal variation in sucker-
ing of aspen obscures effect of various clearcutting
treatments. J. Forestry 62:749-750.

GOLDENBERG, HaRRY', AND A. FERNANDEZ. 1966.

Simplified method for the estimation of inorganic
phosphate in body fluids. Clinical Chemistry 12:
871-882.

Graham, Samuel A., R. P. Harrison, Jr., and C. E.
Westell, Jr. 1963. Aspens, Phoenix trees of the
Great Lakes Region. University of Michigan Press,
Ann Arbor. 272 p.

Hall, H. H. 1953. The impact of man on the Upper
Valley Allotment, Garfield Co., Utah. Master's
Thesis, L'niversity of L'tah, Salt Lake City.

Harper, K. T., and S. K. Hutchins 1965. Evaluation
of grazing disturbance in Utah aspen forests. Bull.
Ecol. Soc. of Amer. 46:47 (abstract).

Heinselman, M. L., and Z. A. Zasada. 1955. A re-
view of literature relating to (juaking aspen sites.
Lake States For. Exp. Sta., U.S. Dept. Agr., Sta.
Paper No. 32. 61 p.

Hills, G. A., and G. Piermont. 1960. Forest site
evaluation in Ontario. Ontario Dept. Lands and
Forests. Res. Branch Tech. Ser. Res. Rpt. 42. 64 p.

Hinds, T. E. 1962. Inoculations with the sooty-bark
canker fungus of aspen. Plant Disease Reporter
46:57-58.

HoFF, C. C. 1957. A comparison of soil, climate, and
biota of conifer and aspen communities in the cen-
tral Rocky Mountains. Amer. Midi. Nat. .58:115-
140.

Holmgren, Arthur H., and J.\mes J. Reveal. 1966.
Checklist of the vascular plants of the Intermoun-
tain Region. U.S. For. Ser. Res. Pap. Int. 32.
Intermountain Forest and Range Exp. Sta., For.
Ser., USDA, Ogden, Utah. 160 p.

Houston, Walter R. 1954. A condition guide for aspen
ranges of Utah, Nevada, southern Idaho, and west-
ern Wyoming. Intermoimtain Forest and Range
Exp. Sta. Res. Paper 32. 25 p.

Hurlbert, S. H. 1969. A coefficient of interspecific
association. Ecol. .50:1-9.

Hutchins, S. Keith. 1965. An index of grazing dis-
turbance in the aspen type of the \Vasatch Range,
Utah. Master's Thesis, University of Utah, Salt
Lake City. 46 p.

Jackson, M. L. 1958. Soil Chemical Analysis. Pren-
tice Hall, Inc., Englewood Cliffs, N.V. 498 p.

Jones, John R. 1967a. Environmental coordinates and
their relation to aspen height growth in the south-
em Rocky Mountains. Ph.D. Thesis. Colorado
State University, Fort Collins. 198 p.

. 19671). Aspen site index in the Rocky Moun-
tains. J. Forestry 65:820-821.

JuLANDER, C, M. E. Lewis, and O. C. Olso.n. 1964.
Aspen communities of the Intermountain Region
and problems of integrating game and livestock
grazing in multiple use management. LInpublished
report of Intermountain Forest and Range Exp. Sta.

Kershaw, Kenneth A. 1964. Quantitative and Dy-
namic Ecology. Edward Arnold Publishers (Ltd.),
London. England. 183 p.

20

Brigham Young University Science Bulletin

KiTTREDCE, Joseph, Jr. 1938. The interrelations of
habitat, growtli rate and associated vegetation in
the aspen commnnity of Minnesota and Wisconsin.
Ecol. Monogr. 8:151-246.

KoHSTiAN, C. F. 1917. The indicator significance of
native vegetation in the determination of forest sites.
Plant World 20:267-287.

Kn.\jiN.\, Vl.\dimir |. 1963. Ecology of the forests
of the Paicfic Northwest — 1962 progress report.
Department of Biology and Botany. University of
British Columbia, Vancouver, B.C. 105 p.

Khebill, Ricii.^Ri) G. 1971. Diseases of aspen on an
elk winter range in the Teton National Forest.
Unpublished report in files of Intermountain For.
and Range E.\p. Sta., USDA, Logan, Utah. 4 p.

Lamb, Fred M. 1967. Aspen wood characteristics,
properties and uses. U.S. Forest Service Res. Pap.
NC-13. 15 p.

L.ANCENHEiM, Jean H. 1962. Vegetation and environ-
mental patterns in the Crested Butte Area, Gunni-
son County, Colorado. Ecol. Monogr. 32:249-285,

Lucas, Paul A. 1969. The effect of cattle, sheep,
and other factors on aspen (Populus tremuloides)
reproduction after clear-cut logging in southern
Utah. Master's Thesis, Utah State University, Logan.
70 p.

Lynch, Bro. Daniel. 1955. Ecology of the aspen
groveland in Glacier County, Montana. Ecol.
Monogr. 25:321-344.

Mark, J. W. 1961. Ecosvstems of the East Slope of
the Front Range in Colorado. University of Colo-
rado Studies, Series in Biology No. 8, L'niversity
of Colorado Press, Boulder. 1.34 p.

Meinecke, E. D. 1929. Quaking aspen — a study in
applied forest pathology, U,S. Dept, Agr. Tech,
Bull, 155. 34 p,

MiELKE, James L, 19.57. Aspen leaf iiliglit in the
Intermountain Region. Interm. For. and Range E,\p.
Sta., USDA. Res. Note 42. 5 p,

Morgan, M. D. 1969. Ecology of aspen in Gunnison
County. Colorado. Amer. Midi. Nat. 82:204-228.

OosTiNG, H. J. 1956. The study of plant communities.
W. H. P'reeman and Co., San Francisco. 440 p.

Oiu.oci, L. 1962. Indicator plants in the coastal west-
em hemlock zone, p. 25-34. In V.J. Krajina (ed.)
Ecology of the forests of the Pacific Northwest.
Dept. of Biol, and Botany, University of British
Columbia, Vancouver, B.C.

Pauley, Scott S. 1949. Forest-tree genetics research:
Populus L. Eton, Bot, 3:299-330.

Re,\m, Robert R. 1963. The vegetation of the Wa-
satch Mountains, Utali and Idalio. Ph.D. Thesis,
University of Wisconsin, Madison. 177 p.

Rkko, I, F, 1952. The vegetation of the Jackson Hole
Wildlife Park, Teton County, Wyoming. Amer.
Midi, Nat, 48:700-729,

Reed, Robert MAnsH,'\LL, 1969, A study of forest
vegetation in the Wind Riyer Mountains, Wyoming.
Ph.D, Thesis, Washington State University, Pullman,
77 p.

RiGcs, G, B, 1929. Using the vegetation cover as an
aid in studying logged-off lands as forest sites. J.
Forestr)' 27:539-545.

RowE, J. S. 1956. Use of understory plant species in
forestry. Ecol. 37:461-472.

Sampson, A. W, 1919, Effects of grazing upon aspen
reproduction. U.S. Dept. Agr. Bull, 741, 29 p,

Snedecor, G. W., and W. G, Cochran, 1967, Sta-
tistical methods. 6th ed. Iowa State University
Press, Ames. 593 p.

Sokal, Robert R., and P. H. A. Sne.'^th, 1963, Prin-
ciples of numerical ta.\onom\ , W. H. f'reeman and
Co., San Francisco, 359 p.

Strain, B. R. 1964. Physiological variabilitv of local
quaking aspen clones. Bull, of Ecol, Soc, .America
45(3):86, (AKstract),

Sthotham, R, O,, and Z, A, Zasada, 1957, Silvical
characteristics of quaking aspen (Populus tremu-
loides). Lake States For, Exp, Sta,, Sta, Pap, No.
49, 26 p,

Warner, James H, 1971, LInderstory characteristics
related to site quality for aspen in Utah, Ph,D,
Thesis, University of Utah, Salt Lake City, 77 p.

S-ajA- ^<^^o^^)

Brigham Young University ^'e/^/\o^°^c

Science Bulletin FSB p ;q~

THE BORAGINACEAE OF UTAH

by
Larry C. Higgins

BIOLOGICAL SERIES — VOLUME XVI, NUMBER 3

JULY 1972

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 Brigham
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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

THE BORAGINACEAE OF UTAH

by
Larry C. Higgins

BIOLOGICAL SERIES — VOLUME XVI, NUMBER 3

JULY 1972

I'Vontispiccc. Erilrwhitun niiiuim ( \'ill. ) Sclirad \i\r. floni^iiturn (lUdh. ) (:roni|.

TABLE OF CONTENTS

Page
INTHODUCTIOX 1

TAXONOMY 1

Boraginaccae - 1

Antsimkia '-'

Ancliusii 4

Aspcrtigo •^

Borcigo ■^

Coldenia ^ 6

Crijplinillui 8

Echiuin 54

Erilrichitini 54

Hucki'liu 55

Heliotropium 58

Ltipindii "O

Lithiisjicrrnxirn "2

Mt'rtetma 66

Miio.mti.i '5

Pcct(icurt/u '3

Plagiohothrys - ' '

ack\o\\t.eix;me\ts 8i

LITERATURE CITED 81

THE BORAGINACEAE OF UTAH

by
Larry C. Higgins*

ABSTRACT

This paper presents a revision of the family
Boraginaceae for the state of Utah. Ninety-three
species in seventeen genera are recognized. The
treatment includes keys, synonymy, descriptions,
and citations of representative specimens found

in the various herbaria in the state of Utah. The
new combination Lappula occidentalis (Wats.)
Greene var. ciipulata (Gray) Higgins is pre-
sented.

INTRODUCTION

For the last six years I have been engaged in
research leading to a revision of the Utah Bo-
raginaceae. This family of flowering plants has
long been a stumbling block for beginning stu-
dents in plant taxonomy and anyone interested
in plant identification. For many years the liter-
ature encompassing the Utah borages has been

inadequate or nonexistent. Investigators have
had to resort to many different publications to
possibly identify the plants at hand. At any rate,
no single work has been available which could
be used to identify the native borages of Utah.
This work is an attempt to provide keys and de-
scriptions for all the Boraginaceae found in Utah.

TAXONOMY

Boraginaceae Lindl.

Boraginaceae Lindl., Nat. Syst., ed. 2:274. 1836.

Plants herbaceous, shrubby or sometimes
tropical trees, usually bristly; leaves simple, al-
ternate, or sometimes opposite or whorled, entire
and pubescent, hispid or setose; flowers perfect,
regular, solitary or cymose; cymes glomerate-
racemose or spicate, fretjuently unilateral and
coiled (scorpioid), usually with bracts between,
to one side of, or opposite the flowers; calyx usu-
allv 5-lobed or 5-parted, usually persis-
tent, the lobes valvate; corolla 5-lobed,
sometimes crested or appendaged in the tliroat;
stamens 5, bonie on the corolla-tube alternate
with the lobes; ovary superior, bicarpellate, usu-
aliv 4-ovulate, entire or lobed, becoming tough
or bony at maturity; fruit commonlv breaking up
into 4 single-seeded lobes (nutlets); style sim-
ple or 2-cleft, seated in the pericarp at the apex
of the fruit or borne between the fruit-lobes
( nutlets ) on the receptacle, or on an upward

^Department of Biology, West Texas State University. (Canyon. Texas.

prolongation thereof (gynobase); endosperm
absent or scarce; embryo straight or curved.

The classification of this family is based pri-
marily upon the structure of the fruit. In many
cases it is difficult to recognize the genus and al-
most impossible to obtain a precise identification
of the species if the specimens lack mature
fruiting stnictures.

The Boraginaceae are of little or no economic
value, but the family has numerous species that
are cultivated as ornamentals, principally in the
genera Heliotropiiim (heliotrope), Aiichusa,
Echium, and Myosotis (forget me not).

Style 2-cleft; stigmas 2, distinct; flowers .solitary or

clustered in the stem forks 5. Coldenia

Stvlf simple; stigmas united (2).

.Stvle arising from the pericarp at the apex of the
fruit, falling away with the nutlets; stigma annular-
peltate, surmounted by a conical or cylindrical, sim-
ple or lobed appendage 11. Ilcliotropium

Style borne between tlie lobes of the fruit (nutlets).

Bbigham Young University Science Bulletin

and attached to the receptacle or gynobase; stigma
capitate, unappendaged (3).

3. Nutlets with uncinate, glochidiate or barbed prickles
on the back, margins or the apex (4).

3. Nutlets without hooked or barbed prickles (7).

4. Nutlets subglobose, with dorsal surface rather uni-
formly covered with barbed prickles, no definite
margins present 7. Cytioglossum

4. Nutlets with a definite margin, the prickles con-
fined to this (back may be muricate or tubercu-
late) (5).

5. Nutlets stellately spreading, attached at the apical
(radicle) end, armed with hooked appendages.
Small slender annuals 16. Pectocanju

5. Nutlets erect, incurved or weakly divergent, at-
tached at or below the middle, i.e., toward the
cotyledon end (6).

6. Plants annual; pedicels erect or nearly so; styles
surpassing the nutlets; subulate gynobase about as
long as the nutlets 12. Lappulti

6. Plants perennial or biennial; pedicels reflexed in
fruit; styles usually shorter than the nutlets; pyra-
midal gynobase about half as long a.s the nut-
lets 10. Hackelia

7. Corolla irregular, the upper lobes usually longer
than the lower ones; stamens not all equal in
length 8. Ecliium

7. Corolla regular or nearly so (8).

8. Calyx in fniit much enlarged, becoming conspicu-
ously veiny, folded and flattened; stems procum-
bent, angled, with stiff retrorse bristles on the
angles 3. Aspenigo

8. Calvx in fruit little if any enlarged, not becoming
veiny, folded and flattened; stems various but not
as above (9).

9. Nutlet attachment surrounded by a swollen ring,
leaving a distinct pit on the gynobase; plants of
fields and waste places (10).

9. Nutlet attachment not surrounded by a rim nor
leaving a pit ( 11 ).

10. Stamens appendaged dorsally, closely crowded
around the style; corolla rotate 4. Borago

10. Stamens unappendaged, included within the tubu-
lar corolla 2. Anchusa

11. Corolla normally blue (aberrant white-flowered
plants occasionally are found ) , or reddish in the
bud stage ( 12 ) .

1 1 . Corolla white, greenish-white. \ ellow or orange

(14).

12. Nutlets with ;ui oblifiue dorsal face encircled by an
upturned flange or rim, this often irregularlv
toothed; depressed-pulvinate plants seldom over
7 cm tall, of alpine areas in Utah . 9. Eritrichiuvi

12. Dorsal f;K(' of nutlet (if present) not encircled by
an upturned flange or rim; pl;ints not depressed-
pulvinate, usually over 7 cm tall, most species
growing below alpine areas in Ut;ih (13).

13. Corolla lobes convolute in the bud; nutlets basally
attached to a flat gynobase; corolhi salverform
15. Mijosotit

13. Corolla lobes imbricate in the bud; nutlets obhquely
attached to a convex gynobase; corolla with a tube
and usually a campanulate throat, not salverform
14. Mertetisia

14. Nutlets attached above the base along a usually
open and generally basallv forked ventral groove or
slit, or by a triangular opening in the pericarp
6. Cryptantha

14. Nutlets lacking a distinct ventral groove or open-
ing in the pericarp, this usually replaced by an
elevated ventral keel (15),

15. Plants perennial; nutlets attached by a broad,
rounded, ((uite basal noncaruncular ;ittachment,
nutlets ovoid, smooth and shiny; corolla usually
yellow or orange 13. Lithospermum

15. Plants annual; nutlets attached by a caruncular
scar Ixjme upon or at the basal end of the ventral
keel, the attachment usuallv lateral or suprabasal;
nutlets usually rough ( 16 ) .

16. Corolla white; cotvledons entire .. 17. Plagiohothrys
16. Corolla orange or yellow, the tube definitely longer

than the calyx; cotyledons 2-lobed .... 1. AmsincKia

1. Amsinckia Lehm.

Anisinckia Lehm., Delect. Sem. Hort. Hamburg
7. 1831; Johnston, Contr. Gray Herb. 70:44.
1924.

Annual, pungent-bristly, herbaceous plants;
stems erect or with spreading branches, leafy;
leaves alternate, linear to ovate, usually vein-
less; racemes usually bractless; calyx cut to base
into erect lanceolate or oblong lobes; corolla
tubular or salverform, heterostyled, yellow or
orange, tube cvlindrical, glabrous, unappend-
aged; lobes spreading, rounded, imbricate; sta-
mens included, affixed in the tube, filaments
very short, anthers oblong; obtuse style filiform,
included; stigma capitate, emarginate; ovules 4;
cotyledons 2-parted; nutlets 4, erect, angulate-
ovoid, smooth or rough, unmargined, strongly
keeled ventrally; gynobase pyramidal, frustrate,
about half the height of the nutlet.

A genus of considerable difficulty which is
found mainly in western North America.

Type species: Ainsinckia hjcopsoidcs Lehm.

1. Corolla-tube 20-nerved below attachment of sta-
mens; calyx-lobes unequal in width and reduced in
numliers (2. 3, or 4) by fusion; nutlets tessel-
late 3. A. tessellata

1. Corolla-tube 10-nerved below insertion of stamens;
calyx-lobes 5, distinct (2).

2. Corolla orange-vellow, 7-20 mm long, well exserted
bcvond the calvx; plants usuiillv green; stems hir-
sute-bristlv, but with little or no fine-apprcssed
h;iirs 1. A. intermedia

2. Corolla pale-vellow, 4-7 mm long, little or not at
all exserted beyond the calyx-lobes; leaves pubes-
cent with appressed or ascending hairs .. 2. A. retrarsa

Biological Series, Vol. 16, No. 3 The Boracinaceae of Utah

1. Amsinckia intermedia Fisch. & Mey. Ind.
Sem. Hort. Petrop. 2:2 and 26. 1836.
Amsinckia campestris Greene, Man. Bay Reg.

263. 1894.
Amsinckia velens Macbride, Contr. Gray

Herb. No. 49:14. 1917.
Amsinckia intactilis Macbride, op. cit. 13.
Ainsinckia arvensis Suksd., Werdenda 1:32.

1927.

Stems simple or much branched, erect to
widely spreading, 3-9 dm tall, sparcely bristly
otherwise usually glabrous except for a tomen-
tose pubescence near the base of the spikes;
basal and lower cauline leaves linear or linear-
lanceolate, the upper lanceolate to nearly ovate,
usually clasping at base and acute at apex, thin-
Iv hirsute on both sides with spreading, often
pustulate hairs; spikes short or usually elongat-
ing in fruit, usually leafy-bracted at base; calyx-
lobes linear attenuate, about half as long as the
corolla, rufous-hispid on the back, densely white-
hirsute on the margins; corolla orange-yellow,
8-10 mm long, the limb 3-6 mm wide; nutlets
2.5-3 mm long, incurved, grayish, narrowly
keeled on the back and sharply rugose with the
surface between papUlate or muricate. A com-
mon field and roadside weed; Washington to
Idaho, and south to Arizona and northern Baja
California. Type localitv: near Bodega Bay, Cali-
fornia. March-June. Fig. 1.

A. intermedia is a highly variable species of
wide geographical range. Suksdorf ( Werdenda
1:48-113. 1931) in a study of the genus proposed
over one hundred new species that fall within
this species. In Utah A. intermedia is located al-
most entirely in Washington County, with a few
scattered records in the western tier of counties.

Millard Co., Wah Wah Mountains, Cottam 5659
IVT): Washington Co., St. George, Law s.n. (UT);
Terry's Ranch, J.B. Karen 18 (BRY); Beaverdam
Nfountains. L.C. Higgins 365, 402. 1222 (BRY); Dixie
State Park, L.C. Higgins 869 (BRY).

2. Amsinckia retrorsa Suksd., Deutsch. Bot.

Monatss. 18:1.34. 1900.
Amsinckia parviflora Heller, Muhlenbergia

2:313. 1907 not Bemh. 18.33.
Amsinckia rufipsa Rvdb., Fl. Rockv Mts. 729,

1066. 1917.
Amsinckia hcUeri Brand, Rep. Spec. Nov.

25:212. 1928.

Stems strictly erect, 3-8 dm tall, usually sim-
ple below the inflorescence, bristly-hirsute and
often more or less cinereous with fine appressed
hairs; leaves linear or the upper linear-lanceo-
late, hirsute on both sides with ascending or

UTAH

County Lines and
river drainages

i-

Fig. 1 . Amsinckia intermedia

appressed hairs; inflorescence of 1 or few,
strict, erect or ascending racemes, bractless;
calyx-lobes 5, distinct, 7-13 mm long, linear or
linear-lanceolate; corolla light-yellow, 5-7 mm
long, the tube included or only slightly exserted
beyond the calyx-lobes; style 2.5-3 mm long;
nutlets 2-3 mm long, broadly ovoid, densely tu-
bcrciilate all over, with scattered larger tubercles
intenntxed, the latter on the central and lateral
ridges when these are present. Moist slopes and
fields or sometimes dry ground. Washington to
Idaho and south to Utah and southern Califor-
nia. Type locality: near Bingen, Klickitat Coun-
ty, Washington. April-August. Fig. 2.

Cache Co., 2 miles southwest of Hyrum, T. Jensen
.589 (UTC): Pine Canyon, Welkville Range, B. Ma-
guire 3111 (UTC): Foot of Sardine Canyon, B. Ma-
guire 13788 (UTC); South of Hardware Ranch and
Randolph Road Jet, W.P. Cottam 16001B (UT); Davis
Co., Centerville, S. Flowers 1192 (UT); Farmington,
B. Maguire 12990 (UTC); Salt Lake Co., Salt Lake
Citv, I.E. Diehl. s.n. (BRY); Salt Lake City. K. Briz-
zee 7780 (UT); Salt Lake City, A.O. Carrett .3640C
(UT); City Creek Canyon. K. Brizzee 7845 (UT); Sum-
mit Co.. Canyon near Gorgoza, A.O. Garrett 8646 (UT);
Tooele Co.. Stansbury Range, South Willow Creek, B.
Maguire 21807 (UTC); Tooele, just northwest of city
limits L.C. Higgins 4092 (BRY); Weber Co., Hunts-
ville, C.P. Smith 1944 (UTC).

3. Amsinckia tesseUata A. Gray, Proc. Amer.
Acad. 10:.54. 1874.

BniciiAM Young University Science Bulletin

Counry Lines and
river drainages

i-

Fig. 2. Amisinckia retrnrsa

Amsinckia coUina Greene, Man. Bay Reg. 263.
1894.

Amsinckia pustulata Heller, Muhlenbergia
2:243. 1906.

Amsiiwkia conica Suksd., Werdenda 1:104.
1931. (In the publication fifteen other
species referable to tesseUata were des-
cribed. )

Stems stout, branched throughout or some-
times simple below, 3-6 dm high, hispid with
spreading bristles; leaves linear-lanceolate, 2-7
cm long, rather thinly hispid, the hairs pustulate
at base, sessile except the narrowly oblanceolate
basal ones; spikes elongating with age, often
.5-12 mm long; calyx lobes 3 or 4, when 4 one
broader and notched or 2-lobed at apex, when
3 a little broader and notched at apex, hispid
and on the margins densely white hirsute, 8-13
mm long; corolla orange, tube 5-10 mm long,
limb 2.5-5 mm wide; nutlets 3-3.5 mm long,
ovoid, the back low usually with a median line,
densely tessellate or papillate, and often trans-
versely rugose. Dry, usually sandy or nK-ky soils.
Eastern Washington to Utah, Arizona ;md north-
em Baja California. Type locality: near Mount
Diablo, California. March-June. Fig. 3.

Davi.s Co., near Centerville, S. Flowers s.n. (UT);
Washington Co., St. George, F.W. CkjuIcI 1479 (BHY);
Beaverdam Wash, L.K. Shumway 2.3 (BRY); Beaver-

dam Wash. W.P. Cottam 5093 (UT); St. George, M.E.
Jones 1640 (UT); Beaverdam Slope, Fisk 3 (UT);
West slope of the Beaverdam Mountains, D. Nish 69
(UTC); 4 miles northwest of Welcome Springs, B.
Maguire 20495 (UTC); 1 mile east of Hurricane, B.
Maguire & Blood 1537 (UTC).

2. Anchusa L.

Anchusa L., Sp. PI. 133. 1753.

Annual, biennial or perennial herbs with blue
or purple flowers in panicled, scorpioid racemes;
calyx divided into narrow lobes; corolla trumpet-
shaped, the tube straight, the throat closed by
scales, the limb with widely spreading lobes;
stamens included, style slender, ovary 4-parted,
nutlets 4, their attachment surrounded by an an-
nular ring leaving a pit on the low gynobase.

1. Anchusa officitwlis L., Sp. PI. 133. 1753.

Plants perennial, from a taproot; stems erect,
branched from near the base, 30-100 cm high;
coarsely hirsute, the hairs often pustulate at
base; basal leaves 8-20 cm long, oblanceolate,
stem leaves lanceolate; calyx 5-8 mm long, the
lobes lanceolate to narrowly triangular, about
as long as the tube; corolla about 10 mm long,
dark blue; nutlets 2-3 mm long, rugose or granu-
late; inserted by their bases on a flat gynobase.
Roadsides and waste places. Native to Eurasia

UTAH

Fig. .3. .\m\inckia tcxscllata

Biological Series, Vol. 16, No, 3

The Boraginaceae of Utah

and introduced into the eastern United States as
far west as Utah. June-August. Fig. 4.

UTAH

County Lines and
river drainages

i-

Fig. 4. Aik7iii.su officinal.

Salt Lake Co., Salt Lake City, L. Arnow 327 (UT);
Utah Co., Rock Canyon, C. Sanders s.n. (UT); Rock
Canyon, Cottam s.n. (UT); Rock Canyon, A.O. Garrett
278.3 (UT); BYU campus, K. Shaw 29 (BRY); Rock
Canyon, P.A. Replogle 85 (BRY); Rock Canyon, L.
Hartman 126 (BRY); Rock Canyon, B.F. Harrison 8380
( BRY ) ; Provo, L. Woodbury 33 ( BRY ) ; Hobble Creek,
L.C. Higgins .3512 (BRY. WTSU).

3. Asperugo L.

Aspemgo L., Sp. PI. 138. 1753.

Rough-hispid, annual, procumbent plants,
with stiff bristly hairs; leaves alternate, or the
upper sometimes opposite, entire; calyx campan-
ulate, unequally 5-cleft, much enlarged and re-
ticulate-veiny in fruit, lobes incised-dentate, the
teeth often appearing as extra lobes in the
sinuses; corolla tubular-campanulate, 5-lobed,
1 to 3 together on short, recurved pedicels in the
upper leaf axis; stamens 5, inserted on the corol-
la tube, included; filaments very short; ovary
4-lobed; style short; stigma capitate; nutlets 4,
ovoid, erect, granular-tuberculate, attached lat-
erally above the middle to the elongate-conic
receptacle, the scar not leaving a pit.

1. Asj)erugo procumhens L., Sp. PI. 138. 1753.

Stems 2-6 dm long; diffusely branched, slen-
der and procumbent or ascending, retrorsely
short-hispid; leaves 1-4 cm long, obovate to
oblanceolate, scabrous, obtuse to acutish at apex;
fruiting calyx 8-15 mm wide; corolla small, 2-3
mm long, blue, purple or purplish-red; nutlets
obliquely ovoid, about 4 mm long, granulate-
tuberculate. Waste places. Introduced from
Europe in various parts of Canada and the
United States. May-August. Fig. 5.

County Lines and
river drainages

i-

Fig. 5. Asperugo procumhens

Cache Co., Logan, in garden, N.H. Holmgren 39
(UTC); USU Campus, B. Maguire 3697 (UTC); 1
mile below Dry Canyon, B. Maguire 13787 (UTC); 1
mile west of Benson, B. Maguire 2417 (UTC); Green
Canyon, Melvin Burke 3696 (UTC); Davis Co., Barton
Creek, mouth of Holbrook Canyon, B. Anderson 51
(UTC); Bountiful, S. Flowers s.n. (UT); Salt Lake Co.,
Salt Lake City, Maguire & Blood 1538 (UTC); Salt
Lake City, A. Bleak 20 (UT); Red Butte Canyon, G.
Swanson 103 (UT); Salt Lake City, A.O. Garrett 8697
(UT); Salt Lake City, W.J. Stubbs s.n. (BRY); Tooele
Co., Tooele, just northwest of the city hmits, L.C. Hig-
gins 4091 (BRY.WTSU); Weber Co., Weber State
College Campus, A. Collotzi 76 (UTC); Foothills east
of Ogden, E.L. Miner s.n. (UT).

4. Borago (Toum. ) L.

Borago (Toum.) L., Sp. PI. 137. 1753.

Hirsute or hispid annual or biennial herbs
with alternate, entire leaves and blue flowers in

Brigham Young Universitv Science Bulletin

terminal leafy racemes; calyx deeply 5-cleft or
5-parted; corolla rotate, the tube very short,
throat closed by scales, limb 5-lobed, the lobes
imbricated, acute; stamens 5, inserted on the
corolla tube; filaments dilated below, narrowed
above to a slender appendage; anthers linear,
erect, and connivent with a beak-like cone; ova-
rv' 4-divided; style filiform; nutlets 4, ovoid,
erect, attached by their bases to the flat recepta-
cle; scar of attachment large, concave.

1. Borago officinalis L., Sp. PI. 137. 1753.

Stems erect, 5-8 dm tall, with ascending or
spreading branches; leaves oblong to obovate,
5-11 cm long, rounded to acute at apex, the up-
per ones clasping, lower narrowed to a winged
petiole; pedicels spreading or recurving, 2-5 cm
long; calyx-lobes linear-lanceolate, 7-10 mm long;
corolla 15-20 mm broad, bright blue; anther-
beak dark purple, about 6-7 mm long; nutlets 4
mm long. An escape from gardens and sparing-
ly naturalized in the western United States, a
native of Europe. June-August. Fig. 6.

Cache Co., Logan J. Thieret 209 (UTC).

5. Coldenia L.

Coldenia L., Sp. PI. 1:125. 1753, and Gen. PI. 61.

1754.

UTAH

County Lines and
river drainages

Is-

Fig. 6. Borago ojfkinalis

Herbaceous or suffruticose plants with slen-
der, forking, usually prostrate or widely spread-
ing stems; leaves small, entire, usually strongly
veined, subsessile or petiolate; flowers small,
generally white, generally extra-axillary, along
leafy twigs or at the forks of the branches, some-
times glomerate; commonly opening in late after-
noon; corolla with a short, cylindrical or ampli-
ate tube and spreading lobes, throat naked or
sometimes appendaged; stamens 4-5, included,
their filaments adnate to the corolla-tube; style
terminal on the ovary, short to long, bUobed or
biparted; stigmas 2, not much differentiated
from the style-branch; ovary 2-celled or some-
times 4-celled by the septum-like placentae, en-
tire or 4-lobed; fruit dry, pyramidal or hemi-
spheric, divided into usually 4 single-seeded nut-
lets; nutlets more or less broadly united ventral-
]y or joined to a central prolongation of the re-
ceptacle.

Except for a single old world species, entirely
confined to arid regions of America. Type spe-
cies : Coldenia procumbens L.

1. Fniit nearly globose, unlobed, breaking apart at
maturity into (|uarter sections, each (|iiarter forming
maturity into (juarter sections, each (juarter forming
a nutlet; leaves ovate to elhptic. white-tomentose,
obscurely veined 1. C. canescens

1. Fniit deeply 4-lobed, the lobes joined only by their
inner angle, each lobe forming a nutlet, leaves not
tomentose (2).

2. Plants perennial; leaves not evidently nerved, lan-
ceolate to linear, usually very pimgently setose;
base of petiole expanded, indurate, usually villous;
flowers solitary in the leaf axils; nutlets finely warty,
ovate 2. C. hispidissima

2. Plants annual; leaves with evident impressed nerves,
ovate or obovate to nearly orbicular; base of petiole
not expanded, or indurate or villoiLs; flowers in
dense clusters at the forks of the stem; nutlets
smooth or granulate 3. C nuttaUii

1. Coldenia canescens DC, Prodr. 9:559. 1845.

Type from between Santander (ie., Jimenez)

and Victoria, Tamaulipas, Mexico, Berlandier

2256.

Stegnocarpus canescens ( DC. ) Torr., in Torr.
& Gray Pacif. R. R. Reports, ed. 2, Vol.
2:170, t;ib. 7. 1,857.

Stegnocarpus leiocanja Torr., Pacif. R. R. Re-
ports, ed. 1, 320. 1855.

Coldenia caiwscens var. stdmuda Johnst.,
Proc. Calif. Acad. Sci. IV. 12:1137. 1924.

Plant suffrutescent, perenni;il, often forming
mats 2-6 dm in diameter; stems several to nu-
merous, mostly prostrate but sometimes ascend-
ing, furcatelv branched; old stems de-
corticating, rough, dark-colored; leafy stems and

Biological Series, Vol. 16, No. 3 The Boracinaceae of Utah

branches pallid, tomentose; leaves numerous,
white tomentose; petiole slender, 2-7 mm long,
in extreme maturity usually breaking off near
the middle leaving a stub attached to the stem;
leaf blade ovate to elliptic-lanceolate, obtuse to
broadly acute at the ends, 7-10 (15) mm long,
2-7 (9) mm wide, thickish, the margins some-
what revolute tomentose; flowers usually solitary
in the axils of the leaves and along the main
stem and branches; calyx sessile, persistent, at
anthesis 3-4 mm long, lobes about as long as
the corolla tube, unequal, united above the base;
lobes of the fruiting calyx lanceolate, usually
long-attenuate, 4-7 mm long; corolla 5-6 (8)
mm in total length, pink, rose or rarely white
the lobes broad and rounded, 1.8-3 (4.5) mm
wide, 1.5-2 (3.5) mm long, usually villous in
the buds, margins frequently erose; filaments all
differing slightly in length and height of attach-
ment; ovary 4-ovulate, at anthesis subglobose,
about 0.8 mm high, marked with 4 longitudinal
grooves; fruit at maturity glabrous or hairy,
ovoid-globose, 2.5-3 mm in diameter, 2-2.5 mm
high, not lobed; nutlets bony, back convex,
densely and minutely tuberculate; style seated in
the pericarp at the apex of the fruit, persistent
until the fruit breaks up into 4 nutlets 1.5-2.5
mm long. Rocky ridges and benches. South-
western Utah, east to Texas, and south to Baja
California and northern Mexico. In Utah C.
canescens Ls limited and very rare in Washing-
ton County. April-June. Fig. 7.

Washint^on Co., West slope of the Beaverdam
Mountains. D. Nish 4.3 (UTC).

2. Coldenia hispidissinui (Torr. & Gray) Gray,
Proc. Amer. Acad. 5:340. 1862; Synop. Fl.
2:pt. 1, 182. 1878.
Eddi/a liispidissima Torr. & Gray, Pacif. R. R.

Reports, ed. 2, 2:170, PI. 9. 1857.
Ehretia Jilspida Torr., Pacif. R. R. Reports,

ed. 1, 320. 1855; without description.

Type is C. Wright 485.

Plants prostrate, forming mats 2-6 dm in di-
ameter; stems several to numerous, dicotomously
branched, spreading from a woody taproot;
younger branchlets villulose-hLspidulous; leaves
clustered, borne mostly on very short branch-
lets along the main stem; petioles 1-2 mm long,
broadest (1-1.5) nun at the base, becoming in-
durate, usually pallid, margin hispid-ciliate and
the back usually glabrous or nearly so; leaf
blades usually linear revolute (rarely ovate or
elliptic), 4-10 mm long, 0.6-1 mm wide, usually
narrower than the indurate petiole-base; flowers
borne among the leaves; calyx sessile, broadly

County Lines and
river drainages

L I I

Coldenia ciinc.sceus

and pennanently attached in the leaf axil, at
anthesis about 3 mm long, lanceolate, united at
the base, villulose-ciliate below the middle, fre-
(juently terminated with a stiff bristle; corolla
usually pink, about 7.5 mm long; corolla lobes
rounded, spreading, about 2 mm long and 3 mm
broad; stamens 5, unequal in length and position
within the corolla tube; style 1.5-2.2 mm long,
somewhat flattened, apex bilobed; nutlets ovoid,
usually only 1 or 2 maturing, 1.1-1.4 mm long,
0.8-1 mm broad, papillate, back convex, ventral
rounded or somewhat flattened; scar open, 1.5-2
mm wide at base, nearly as long as the nutlet,
surrounded by a narrow band of smooth non-
papillate pericarp, traversed down the center by
a lineate ridge. Nevada, southern Utah, northern
Arizona, and east to Texas and south to north-
em Mexico. Usually growing on sand dunes and
dry slopes. April-September. Fig. 8.

Emery Co., San Raphael Desert, Brvan & Redd
12-8 (UTC); 6 miles north of Hanksville! B. Maguire
19.386 (UTC); Garfield Co.. Clay Point, J.C. Pederson
49 (BRY); 2 miles dovvn.stream from Hite, B.F. Harri-
son 11737 (BRY); Hite, W.P. Cottam 14769 (UT);
Grand Co.. Castle Valley, S.L. Welsh & D. Atwood
99.54 (BRY); 1 mile north of Moab, B. Maguire 5722
( UTC ) ; 6 miles up river from Moab, Bryan & Moab
School 6-8 (UTC); Kane Co., Bullfrog Canyon, D.W.
Lindsay 93 (UT); San Juan Co.. West of Bluff, W.P.
Cottam 2.560 ( BRY ) ; along the San Juan River, Rydberg
& Garrett 9942 (UT); Middle Blue Notch Canyon, L.
Wilson 273 (UTC); 9 miles southwest of Bluff, B.

Bricham Young Univehsity Science Bulletin

County Lines and
river drainages
1 1 I

Fig. 8. CukU-nia hisimlissima

Maguire 5719 (UTC); 35 miles southwest of Bluff, B.
Maguire 5722 (UTC); Washington Co., Beaverdam
Mountains, L.C. Higgins 838 (BRY); Wayne Co.,
HanksvQle, R.V. Hardy 41 (BRY); 5 miles north of
HanksviUe, S.L. Welsh'& G. Moore 3611 (BRY); de.sert
between HanksviUe and mountains, W.P. Cottam 5126
(BRY).

3. Coldenia nuttallii Hooker, Kew Joum. Bot.
3:296. 1851. Type: "Rocky Mountains," col-
lected by Nuttall.
Tiquilia parviflora Nutt. ex Hook., loc. cit. as

a synonym.
Tiquila brevifolia Nutt. ex Torr.., Bot. Mex.
Bound. 136. 1859.

Prostrate annual with slender, somewhat brit-
tle, dicotomously branched stems forming mats
1-3.5 dm broad, finely strigose; leaves ovate, to
nearly sulx)rbicular, 4-8 mm long, narrowly revo-
lute, and often hispid on the margins, witli 2-3
pairs of distinct veins on the back, thinly strigose
on the upper surface with somewhat stiff hairs,
hirsute on the lower surface; petioles slender,
usually as long or longer than the blade; flowers
in compact c-lusters in the forks and at the ends
of the branches; calyx lobes linear-subulate, 4-5
mm long, villous or set\ilose on the back, sparse-
ly but conspicuously hispid on the margins; co-
rolla pink or nearly white, little exceeding the
calyx, the limb 2-2.5 mm broad, the tube with
5 triangular scales near the base; nutlets oblong-

ovoid, smooth and shining. Sandy or alkaline
soil, on plaias and hillsides. Washington, east
tlirough Idaho to Wyoming and south to Utah,
Arizona, and southern California. May-August.
Fig. 9.

Beaver Co., Pine Valley, W.P. Cottam 5701 (UT);
Box Elder Co., Copper Mountain, W.P. Cottam 4551
(BRY, UT); Iron Co., Hamblin Valley, W.P. Cottam
3335 (BRY, UT); .sand dune areas, W.D. Stanton s.n.
(UT); Millard Co., desert area, W.P. Cottam 5701
( UT ) ; 6 mUes north of Candy, Maguire & Richards
2752 (UTC); 2 miles .south of Desert Range Experi-
ment Station Headquarters, B. Maguire 20913 (UTC);
3 miles south of Carrison, B. Maguire 20854 (UTC);
Uintah Co. Willow Creek, 16 miles south of Ouray, R.C.
Rollins 1724 (UTC); Washington Co., Beaverdam
Mountains, L.C. Higgins 649 (BRY).

6. Cnjptantha Lehm.

Cryptantha Lehm., Del. Sem. Hort. Hamburg 4.
1833, and Fisch. & Mey., Ind. Sem. Hort.
Petrop. 2:35. 1836; Johnston, Contr. Gray
Herb. 74:1-114. 1925, and Contr. Gray Herb.
78:30. 1927.

Knjnitzkia Fisch. & Mey., Ind. Sem. Hort.
Petrop. 7:52. 1841.

Annual or perennial, herbaceous or frutic-
ulose plants, usually with coarse stiff pubes-
cence; leaves opposite at base, or alternate
throughout, finn, veinless; flowers white or rare-

UTAH

County Lines and
river drainages

l'"ii;. 9. (U)l(lcnui nuttallii

Biological Series, Vol. 16, No. 3

The Bohaginaceae of Utah

9

ly yellow, in bractless or bracted spikes or ra-
cemes; calyx divided to the base, the lobes erect
or connivent, linear or oblong; mature calyx in-
vesting the nutlets and falling away entire, or
the calyx persistent and the nutlets falling away
separately; corolla wdth a short to somewhat
elongate cylindrical tube with or without scales
at the base of the tube, the throat with intruded
appendages; corolla lobes imbricate, rounded,
spreading; style slender, short or long, included;
stigma capitate; ovules 2-4; nutlets 1-4, erect,
ovate to triangular, roughened or smooth,
winged, margined or marginless, affixed laterally
through a medial ventral and commonly basally
forked groove to a usually columnar, subulate or
pvramidal gvnobase.

This is one of the most perplexing genera in
the entire family. It is exclusively American,
mostly in the western United States, but com-
mon in the deserts of South America as well.
Several species are reported to have some value
as a forage for sheep.

Type species: Cryptantha glomerata Lehm.

1. Plants annual; stems slender; Subgenus Krynitzkia
(2).

1. Plants biennial or perennial; steins coarse; Subgenus
Oreocarya (31).

2. Nutlets with the margins decidedly winged or
knifelike (3).

2. Nutlets with the margins rounded or angled, never
with a marginal wing or knifelike edge (6).

3. Pedicels usually evident, slender, 1-4 mm long; nut-
lets heteromorphous 18. C. mcemosa

3. Pedicels obscure or none, less than 1 mm long ( 4 ) .

4. Nutlets heteromorphous, the odd nutlet abaxial
12. C. inaequata

4. Nutlets homomorphous or if slightlv heteromor-
phous the odd nutlet axial (5).

5. Nutlets solitary or rarely 2; calyx obliquely conical
at base; corolla conspicuous 22. C. Utahensis

5. Nutlets 4; calyx symmetrical; corolla inconspicuous.
17. C. ptcTocanja

6. Nutlets all smooth ( 7 ) .

6. Nutlets all rough or at least some of them so ( 14 ) .

7. Hairs on the calvx uncinate or decidedly arcuate.
.'. 10. C. flaccida

7. Hairs on the calvx straight (8).

8. Nutlets with an excentric groove; flowers in biserial
naked spikes 1. C ajjinis

8. Nutlets with a centrally placed groove ( 9 ) .

9. Nudets broadly ovate (10).

9. Nutlets oblong-ovate to lanceolate (12).

10. Spikes usu;illv geminate; inflorescence projected
above the leafy mass of the plant, well defined
21. C. torreyana

10. Spikes usually solitary, not sharply differentiated
from the leafy peduncular stems (11).

11. Nutlets homomorphous 2. C. ambigua

11. Nutlets slightly heteromorphous .-.16. C. pattersonii

12. Style reaching }* to % the height of the nutlets;
calyx densely appressed hispid-villous, commonly

lacking conspicuous spreading bristles

11. C. gracilis

12. Style almost reaching the nutlet-tips or surpassing
them (13).

13. Margin of nutlets acute, at least above the middle.
23. C. watsoni

13. Margin of nutlets rounded or obtuse.-.9. C. fendleri

14. Nutlets decidedly heteromorphous (15).

14. Nutlets homomorphous (22).

15. Mature calyces strongly appressed to the flattened
rachis, decidedly gibbous on the axial side, per-
sistent 8. C. dumetoTum

15. Mature calyces somewhat spreading, not at all gib-
bous (16).

16. Odd nutlet abaxial, surpassed by style ( 17 ) .

16. Odd nutlet axial; style surpassed or occasionally
reaching to the nutlet-tips (20).

17. Spikes braeteate throughout; calyx persistent
14. C. micrantha

17. Spikes naked or nearly so; calyx deciduous (18).

18. Pedicels slender, 1-4 mm long 18. C. racemosa

18. Pedicels stout and obscure, less than 1 mm long

(19).

19. Nutlets 1.3-1.7 mm long; calyx 2-3 mm long
12. C. inaequata

19. Nutlets about 1 mm long; calyx 3-4 mm long
3. C. angustifolia

20. Odd nutlet spinular-muricate; calyx-lobes conspicu-
ouslv thickened 6. C. crassisepala

20. Odd nutlet more or less granulate; calyx lobes
moderately thickened (21).

21. Nutlets ovate, smoothish or sparsely tuberculate,
odd one about 1.9 mm long 16. C. pattersonii

21. Nutlets lanceolate or narrowly ovate, coarsely tu-
berculate. odd one 2-2.6 mm long ... IS.C.kelseyarm

22. Calyx circumscis.sle 5. C. circttmscissa

22. Calyx not circumscissle (23).

23. Style surpassing the nutlets 14. C. micrantha

23. Style not surpassing the nutlets or about reaching
to' the Hps of the nutlets ( 24 ) .

24. Corolla conspicuous, 2-5 mm broad (25).

24. Corolla inconspicuous, 0.5-2 mm broad (26).

25. Abaxial nutlet developing; gynobase reaching '3 to !i
the height of the nutlet 7. C. decipiens

25. Axial nutlet developing; gynobase about S height
of nutlet 22. C. utahensis

26. Ovules 2; nutlet and calyx bent .... 19. C. recurvata

26. Ovules 4; nutlets and calyx straight (27).

27. Nutlets usually solitary, abaxial; gynobase reaching
K to S the height of the nutlet 7. C. decipiens

10

Bricham Young University Science Bulletin

27. Nutlets usually 4 (28).

28. Nutlets decidedly ovate, with low rounded tuber-
culations 2. C. ambigua

28. Nudets more or less lanceolate (29).

29. Stems spreading-hirsute 4. C. barbigera

29. Stems strigose (30).

30. Nutlets verrucose or verrucose-muricate

15. C. nevadensis

30. Nutlets spinular-muricate 20. C. scoparia

31. Corolla tube elongate, distinctly surpassing the
calv.x; flowers usually heterostyled (32).

31. Corolla tube short, scarcely if at all surpassing the
calyx; flowers not heterostyled (46).

32. Nutlets smooth and shining (33).

32. Nudets more or less roughened ( 38 ) .

33. Corolla yellow (34).

33. Corolla white (3.5).

34. Inflorescence an elongate, cylindrical thyrse; nut-
lets lanceolate with acute margins, usually only 1
developing 33. C. jlava

34. Inflorescence consisting of a large terminal cluster
with I or more remote, at maturity frequently
stalked much smaller lateral clusters; nutlets broadly
ovate, with winged margins, all 4 usually matur-
ing 31. C. confcrtiflora

35. Inflorescence capitate, 0.1-0.4 dm long; corolla
limb 6-8 mm broad, the tube litde surpassing the
calyx; nutlets lanceolate 29. C. oapitata

35. Inflorescence elongate, 0.4-4.4 dm long; corolla
limb 8-17 mm broad, the tube distinctly surpassing
the calyx except in C. barnebtji; nutlets ovate (36).

36. Ventral surface of the leaves glabrous

48. C. semighbra

36. Ventral surface of the leaves strigo.se or setose-his-
pid (37).

37. Corolla limb 13-17 mm broad, crests at base of
tube absent; nutlets 3-3.5 mm long; native to San
Raphael Swell 39. C. johnstonii

37. Corolla limb 8-11 mm broad, crests at base of tube

conspicuous; nutlets 3.5-4.5 mm long

26. C. hornehxji

38. Nutlets uniformly muricatc or papillose, or oc-
casionally in C. joncsknui also with some incon-
spicuous ridges (39).

38. Nutlets more or less rugose or tuberculate, or
sometimes with a few inconspicuous inurications
(40).

39. Leaves oblanceolate, strigose with pustuliite hairs
small or lacking; corolla 7-10 mm long, fomices
elongate 35. C. jiilcoc-anescens

39. Leaves spatulate, hispid with pustulate bristles;
corolla 10-15 mm long, fomices low ;ind broad
40. C. joncsiana

40. Ventnil or inner surface of the nutlets smooth or
nearly so 46. C. rollinsii

40. Ventral surface of the nutlets distinctlv roughened
(41).

41. Leaves conspicuously pustulate yentrally; corolla

tube 12-16 mm long; calyx segments 7-10 mm long
in anthesis 41. C. longiflora

41. Leaves sparsely if at all pustulate ventrally; corolla
tube 5.5-12 mm long; calyx segments 3.5-7 mm
long in anthesis (42).

42. Inflorescence 0.1-0.4 dm long; corolla tube 10-12
mm long; margin of nutlets not in contact; plants
less than 1.5 dm tall 45. C. pawdoxa

42. Inflorescence 0.5-3 dm long; corolla tube 5-10 mm
long; margin of nutlets in contact or nearly so;
plants usually over 1.5 dm tall (43).

43. Scar of nutlets surrounded by an elevated margin
but tightly closed; style 1-2 mm long; calyx 3.5-4
mm long in anthesis 25. C. bakeri

43. Scar of nutlets conspicuously open; style 3-8 mm
long; calyx 4.5-7 mm long in anthesis (44).

44. Scar of nutlets conspicuously open and surrounded
by a definite elevated margin .... 34. C. fkwoculata

44. Scar of nutlets slightly open and with only an in-
conspicuous elevated margin if any (4.5).

45. Leaves linear-spatulate; nutlets sharply and deeply
rugose; corolla tube 5.5-7 mm long; fomices low
and broad 52. C. tenuis

45. Leaves obovate or broadly oblanceolate; nutlets
with rounded ridges and tubercles; corolla tube

7-10 mm long, fomices long papillose

54. C. wetherillii

46. Nutlets smooth on their dorsal surface, not mgose,
muricate or tuberculate ( 47 ) .

46. Nutlets more or less roughened, muricate, rugose
or tuberculate at least on the dorsal surface ( 48 ) .

47. Fruit depressed globular, nutlets not in contact by
their margins; style exceeding mature fruit 1-3 mm;
corolla tube 2.5-3 mm long 38. C. jamesii

47. Fruit conical, ovoid or lanceolate, nutlets in con-
tact by their margins or nearly .so; style exceeding
mature fruit 5-6 mm; corolla tube 5-7 mm long....
26. C. barnebtji

48. Ventral surface of nutlets smooth or nearly so (49).

48. Ventral surface of nutlets mgose or variously
wrinkled ( 52 ) .

49. Nutlets bordered by a conspicuous wing; robust
plants 5-10 dm tall, with long ebracteate spikes ....

50. C. setosissima

49. Nutlets never conspicuously winged, sometimes
with an acute margin; plants usually lower and
caespitose (50).

50. Corolla tube 7-9 mm long; calyx 6-9 mm long in
anthesis 46. C. rollinsii

50. Corolla tube 2-6 mm long; calyx 2.5-6 mm long in
anthesis ( 51 ) .

51. Nutlets scarcely or not at all muricate between the
nigae; strictly erect, conspicuously hispid perennials
from iiDrtheastem I'tah 51. C. strict<i

51. Nutlets distinctly muricate or tuberculate between
the rugae and near the margins; western Utah. ...
..._ 47. C. nigulosii

52. Nutlets conspicuously muricate, or in (.'. hurnilis
also with a few irregular ridges (53).

52. Nutlets not exclusiyely muricate, but rugose or
tuberculate (55).

Biological Series, Vol. 16, No, 3 The Boracinaceae of Utah

11

53.

53.
.54.

54.

55.
55.

56.
56.

57.

Pubescence of the leaves silky-strigose or strigillose

but not subtomentose or tomentose

27. C. breviflora

Leaves distinctly subtomentose or tomentose, also
setose in C. humilis (54).

Plants 0.3-1 dm tall; leaves 0.5-2.5 cm long; caly.\
2-2.5 mm long

65. Corolla tube 2-2.5 mm long; style exceeding nut-
lets by 1 mm or less; endemic to Garfield Coun-
ty 43. C. ochwleuca

65. Corolla tube 3.5 mm long or longer; style exceed-
ing nutlets by more than 1 mm; northeastern
Utah 49. C. sericea

in anthesis; corolla tube 1.8-2.2 mm 66.

long; Millard County', Utah 30. C. compacta

Plants 0.4-2.5 dm tall; leaves 2.5 cm long or long-
er; calv.x 3-.5 mm long in anthesis; corolla tube 3-5
mm long; plants widespread 37. C. humilis

Scar of nutlets open .some distance above the base
(56).

Scar of nutlets closed or nearlv so, without a con-
spicuous triangular opening toward the base ( 63 ) .

Scar somewhat constricted some distance below the
middle of the open portion ( .57 ) .
Scar triangular and not constricted below the mid-
dle (58).

Elevated margin of the scar definitely limited;

pustules present on both leaf surfaces

42. C. mensana

■57. Elevated margin indefinitely limited; pustules pres-
ent only on dorsal surface of the leaves

44. C. osterhoutii

58. Some tendency to an elevated margin evident
around the scar (59).

58. No tendency to an elevated margin around the
.scar (60).

59. CNTTiules elongating and so the inflorescence
broad; biennial or short-lived perennials; nutlets

usuallv witli an evident dorsal ridge

53. C. virgincnsis

59. Cymules shorter and the inflorescence narrow;
long-lived caespitose perennials; nutlets with only a
slight dorsal ridge if anv 24. C. abaUi

60. Stvle not exceeding the mature nutlets by more
than 0.5 mm; plants usuallv less than 1.3 dm tall
(61).

60. Stvle exceeding the mature nutlets 1.6 mm or

more; plants usually taller than 1.3 dm

^ 37. C. humilis

61. Corolla tube 2-2.6 mm long; nutlets 2.3-3 mm
long 43. C. ochroleucu

61. Corolla tube .3-4 mm long; nutlets .3-3.5 mm long
(62).

62. Ventral surface of nutlets deeply rugose and tuber-
culate 24. C. abata

62. Ventral surface of nutlets indefinitely muricate;
near the Utah border in Summit and Daggett
counties 28. C. caespitosa

63. Upper surface of the leaves uniformly :ippressed
strigose and without pustulate hairs ( 64 ) .

63. Upper surface of the leaves with two distinct kinds
of hairs; pustulate at base (66).

64. Nutlets sharply rugose and tuberculate. scar closed,
and surrounded bv an elevated margin

' 25. C, hakcri

64. Nutlets not so sharply rugose or tuberculate; scar
not surrounded by an elevated margin (65).

66.

67.

Mature calyx exceeding the nutlets by 2-4 mm; in-
florescence broad-topped; Grand County

32. C. elata

Mature calyx exceeding the nutlets by 4-8 mm
(67).

Nutlets tuberculate, scarcely if at all rugose; mostly

in the Uintah Basin 36. C. grahamii

Nutlets more or less rugose; widespread but mostly
in the western tier of counties 37. C. humilis

1. Cryptantha affinis (Gray) Greene, Pittonia
1:119. 1887.

Krt/nitzkia affinis A. Gray, Proc. Amer. Acad.
"20:270. 1885. Type: east side of Cascades.
1860. Lyall s.n.
Crifptantha geminata Greene, Pittonia 1:119.

" 1887.
Cniptantha confusa Rydb. Bull. Torr. Bot.

Club 46:679. 1909.
Cnjptantha eastwoodiae St. John, Fl. S.E.
Washington 342. 1937.

Usually sparsely branched herb 1-2 (4) dm
tall; branches commonly few and ascending but
plant occasionally much branched from the base,
hispid or short-hirsute throughout; leaves nar-
rowly to broadly oblanceolate, 1-4 (5) cm long,
2.5-6 (8) mm broad, few, short-hirsute, usually
minutely pustulate, obtuse or rounded at tip,
lowest pair clearly opposite; spikes geminate or
solitary, usually 2 to 8 becoming 15 mm long,
slender, remotely flowered, commonly with a
very few large leafy bracts below; corolla in-
conspicuous, 1-2 mm long, limb about 1.5 mm
broad; fruiting calyx 2.5-4 mm long, usually
about as broad as long, laterally compressed, as-
cending; pedicels 0.5-1 mm long; mature calyx-
lobes lanceolate, somewhat cormivent, not great-
ly surpassing the nutlets, midrib weakly thick-
ened on the abaxial lobe, sparsely hirsute, mar-
gins appressed hispid; nutlets 4, homomorphous,
smooth or very finely granulate, shiny, brovimish
to greenish, frequently mottled, 1.8-2.5 mm long,
ovate, obliquely compressed, back low-convex,
margins rounded; groove evidently excentric,
closed, simple or shortly and unequally forked
at the base; gynobase .short, stout, about 1/2
height of nutlets; style evidently surpassed by
nutlets or rarely equalling them. Usually in
sandy to rocky soils. Washington and western
Montana, south to Wyoming, Utah, and southern
California. June-September. Fig. 10.

12

Brioham Young Univebsitv Science Bulletin

County Lines and
river drainages
I 1

30 b"

Fit;. 10. Cryptmttha uffiiii.s

The most obvious characteristics of this spe-
cies are the obhquely compressed nutlets which
result in the excentric position of the groove.

Cache Co., 10 miles up Smithfield Canyon, Hobson
& Piranian 13775 (UTC); Inten'ale, Spring Hollow, B,
.\Iaguire 1.38.39 (UTC); White Pine Lake, B. Maguire
16256 (UTC); Logan Canyon, Spring Hollow, B. Ma-
guire 13778 (UTC); Daggett Co., Flaming Gorge, L.
William.s 473 (UTC); Duchesne Co., 10 miles north of
Altonah, J. Brotherson 646 (BRY); Juab Co., Granite
Canyon, Deep Creek Mountains, Maguire & Becraft
2765 (UTC); Utah Co., Spring Dell.' Provo Canyon,
A.O. Garrett 2778 (UT); Weber Co.. Summit of Little
Bear River Canyon, B. Maguire 12976 (UTC).

Cryptantha ambigua ( Grav ) Greene, Pittonia

1:113. 1887.

Eritrichium muricuhitum var. amhiguum

Gray, Synop. Fl. No. Am. 2:194. 1878.

Type: probably in the Walla Walla region

of southeastern Washington, collected by

the Wilkes expedition.
KrynitzkUi ambigua (iray, Proc. Am. Acad.

20:273. 1885.
Cryptantha polycarpa Greene, Pittonia 1:114.

1887.
Cryptantha inuUicaulis A. Nels. Bot. Gaz.

30:194. 1900.
Cryptantha ambigua var. robustior Brand,

Pflanzenreich 4, 252:69. 1931.

Stems usually loosely branched from the
base, ascending, 1-2.5 dm tall, hirsute and some-
what strigose; leaves linear to narrowly lanceo-
late, 2-3 (5) cm long, 1-4 (5) mm broad, obtuse
to acutish, usually somewhat appressed hispid-
hirsute, the hairs commonly pustulate at base;
spikes often solitary, 5-15 cm long, bractless, or
with the lowennost flowers bracteate, commonly
not projecting clear of the leafy mass of the
plant and not clearly differentiated from the
leafy branches; corolla 1-2 mm broad, incon-
spicuous; fruiting calyces ovate-oblong, 4-7 mm
long, crowded or distant, the tube rounded-ob-
conic at base, lobes linear or linear-lanceolate,
more or less connivent, midrib thickened, tawny-
hirsute, margins strigose-hirsute; pedicels 0.5-
0.9 mm long; nutlets 4, broadly ovoid, 1.6-2 mm
long, granulate and coarsely tuberculate or rare-
ly tending to be smooth toward the base, back
low-convex, sides obtuse and rounded, groove
closed or somewhat open at the always broadly
forked base; gvnobase narrow, 1-1.2 mm long,
2/3 height of nutlets; style reaching 4/5 to 5/5
height of nutlets. Dry slopes and ridges, open
pine forests and sagebrush flats. Southern Wash-
ington to southern Montana and then south to
northern Colorado, western Nevada, and north-
eastern California. June-August. Fig. 11.

C. ambigua is nearly always distinguishable
bv its habit of growth, but in a few cases is dif-

UTAH

County Lines and
river drainages

A-

Kil^. 11. Crifpttnttlui uinhi<i\ia

Bk)1.o(;k AL SEHIE.S, \()i.. 16, No. 3

The Boraginaceae of Utah

13

ficult to distinguish from C. kehetjana and C.
pattersonii.

Beaver Co., 11.5 mile.s east of Milford at the Gran-
ite Nfountain Pass, B. Mapiire 20995 (UTC); Bo.\ Elder
Co.. R;ift River Mountain's, CM. Alder 132 (UT); Juab
Co.. Deep Creek Range, Granite Greek, Maguire &
Holmgren 21904 (UTC); Utah Co., Thistle, M.E.
Jones 5370 (UC).

3. Crijptantha angustifolia (Torr. ) Greene,
Pittonia 1:112. 1887.

Erithcliitim angustijolium Torr. Pacif. R. R.
Reports 5:363. 1858. Type: Yuma, Ari-
zona, Thomas s.n.
Krtinitzkki angustifolia A. Gray, Proc. Amer.
"Acad. 20: 272. 1855.

Stems diffusely branched from the base, 0.5-
3 dm tall; canescent, villous-hirsute, with light
ashy gray hairs, the lowest branches decumbent
or ascending; leaves narrowly linear, 1.5-4 cm
long, 1-2 mm wide, hispid or strigose, somewhat
pustulate; spikes usually geminate, about 5 cm
long, rather dense, bractless or with 1-2 bracts
near the base; corolla minute, the tube 1-2 mm
long, limb 1-2.5 mm broad; fruiting calyces
ovate-oblong, 3-4 mm long, stiffly ascending,
stronglv biseriate, slightly asymmetrical; pedi-
cels less than 0.5 mm long; mature calyx-lobes
linear-lanceolate, slightly connivent, hispid on
the thickened midrib, short villous on the mar-
gins, abaxial lobe longest and most hirsute; nut-
lets usually 4, heteromorphous, ovate-oblong,
brown or plumbeous with pale tuberculations or
rarely murications, back convex, face flattish,
margins somewhat angular; odd nutlet next to
the abaxial calyx-lobe, a little larger and more
persistent than the similarly colored and shaped
consimilar nutlets which are about 1 mm long;
groove slightly open above, broadening at the
base; gynobase columnar, equalled by consimilar
nutlets but shorter than odd nutlet; style usually
surpassing even the odd nutlet. Sandy or gravel-
Iv washes. Lower Sonoran Zone. Southern Cali-
fornia to Southern Utah and south to Baja Cali-
fornia, Sonora, and western Texas. March-June.
Fig. 12.

I have not seen any specimens of C. angusti-
folia from Utah, but the species has been col-
lected within a few miles of the Utah border in
both Arizona and Nevada. This species would
most likely be found in the Bcaverdam Wash or
the valleys of the Virgin River.

4. Cnjptantha harhigera (Gray) Greene, Pit-
tonia 1:114. 1887.

Eritrichium harhigerum A. Gray, Synop. Fl.
No. Amer. 2:194. 1878. Type: Utah

(Parry 171) as designated by I. M. Johns-
ton, Contr. Gray Herb. 74:66. 1925.

Krtjnitzkia harbigera A. Gray, Proc. Amer.
Acad. 20:273. 1885.

Knjnitzkiu mixta M. E. Jones, Contr. West.
Bot. 13:6. 1910.

Stems erect, 1-4 dm high, solitary or several
from the base, branches strictly ascending or
spreading, very bristly and sparsely if at all
strigose except in the inflorescence; leaves ob-
long to lance-linear, obtuse, 1-5 (7) cm long,
3-7 ( 13 ) mm broad, hirsute, more or less pustu-
late; spikes geminate or rarely solitary or ter-
nate, naked, as much as 15 cm long; corolla in-
conspicuous, limb 1-2 mm broad; fruiting calyx
ascending, 5-10 mm long, narrowly ovoid-ob-
long to oblong-lanceolate, symmetrical, decidu-
ous; pedicels 0.3-0.7 mm long, villous; mature
calyx-lobes lanceolate to linear-lanceolate, coimi-
vent above with recurving tips, margin con-
spicuously long white-villous, midrib thickened
and hirsute, abaxial lobe slightly the longest;
nutlets 1-4, homomorphous, lance-ovoid, 1.5-2.5
mm long, strongly verrucose, usually brownish,
back convex, edges obscurely angled or rounded,
groove opened or closed but base gradually di-
lated to form a triangular areola; gynobase nar-
row, 2/3 to 3/4 height of nutlets; style reaching
to or slightly beyond the nutlet tips. Sandy des-
ert washes or slopes in the Lower Sonoran Zone.

UTAH

County Lines and
river drainages

i-

Kig. 12. Crijptaniha (in^ustifolia

14

Brigham Ydung University Science Bulletin

Southern Utah south to southern California, Baja
California, Arizona, southern New Mexico, and
northern Sonora. February-June. Fig. 13.

S;m Juan Co., 2 miles west of Bluff, B. Maguire
16279 (UTC); Washington Co., St. George, F. W.
Gould 1478 (DI.X, UTC); 2 miles west of Rockville,
B. Maguire 4978 (UTC); 4 miles northwest of Welcome
Springs, B. Maguire 20527 (UTC); 40 miles west of St.
George. B. Maguire 20506 (UTC); 10 miles west of
Hockville, C.L. Hitchcock 3003 (UTC); St. George,
M.E. Jones 1621 (UT); St. George, F.W. Gould 1539
(UT); Black Hill west of St. George, DE. Beck s.n.
(DIX); Red hill north of St. George, S. Wilson s.n.
(DIX); Beaverdam Mountains at junction of hwy 91
and the Jackson road, L.C, Higgins 447 (BRY); Di.\ie
State Park, L.C. Higgins 930 (BRY); 3 miles south of
Touquerville, B.R. Stahmann & J.L. Jacobs 21 (BRY);
St. George, south slope of Black Hill, F.W. Gould 1478
(BRY): about 5 miles west of hwy 91 along the Jack-
son road, L.C. Higgins 500 (BRY); St. George, D.H.
Galway 8485 (BRY).

5. Cniptantha circumscissa (H. & A.) Johnston,
Contr. Gray Herb. 68:55. 1923. Type: Snake
Fort, Snake River, Idaho.
LHl)o.s])ermiim circumscisstnn H. & A. Hot.

Beechey 370. 1840.
Piptocah/x circumscissus Torr. in S. Wats.

Hot. King Expl. 240. 1871.
Eritrichium circtmiscissum Gray, Proc. Amer.

Acad. 10:58. 1874.
Kn/nitzkia circwnscissa Gray, Proc. Amer.

"Acad. 20:275. 1885.
WheeJereUa circuriiscissa Grant. Bull. Soc.

Calif. Acad. Sci. 5:28. 1906.
Crt/ptantha depressa A. Nels. Bot. Gaz. 34:29.

" 1902.
Greenocharis circumscissa Rydb. Bull. Torrey

Club 36:677. 1909.

Stems few to many from the base, strigose,
more or less branched above, often forming a
dense hemispherical mass 2-10 cm high, the
outer ones often decumbent; leaves oblanceo-
late, .3-15 mm long, 1-2 mm broad, obtuse, sur-
face siliceous especially toward the pale base,
strigose or short-hispid, obscurely pustulate;
flowers in the axils of foliaceous bracts in short
somewhat indefinite racelike clusters; corolla in-
conspicuous, 1-2 (3) mm broad; fruiting calyx
2.5-4 mm long, oblong-ovoid, united to near the
middle, at maturitv the upper half falling away
l>v ;i circumscission just below the sinuses, basal
part persistent, cupulate, appressed-hispid; ma-
tiu-e calyx lobes narrowly linear-lanceolate, firm-
herbaceous, scarcely ribbed, more or less hispid;
p<'dicels obscure, about 0.5 mm long; nutlets 4,
homomorphous, or with the abaxial one slightly
longer, smooth or inconspicuously muricate, tri-
angular-ovoid or oblong-lanceolate, 1.2-1.7 mm
long, back flattened, especially near the apex.

County Lines and
river drainages

i-

Kig. 13. Cn/iitanlha luirhigcra

margins angled, groove closed and forked at
base, gynobase about 2/3 height of nudets; style
equalling or barely exceeded by nutlets. Sandy
to gravelly soils, Sonoran to arid Transition
Zone. Southern British Columbia and Idaho,
south to Arizona and Baja California. April-
August. Fig. 14.

Beaver Co., 8.5 miles east of Milford, B. Maguire
21020 (UTC); Garfield Co., Egnog Spring, Bullfrog
Creek west of Henry Mountains, Welsh 3984 (BRY);
Iron Co., 16 miles south of Lund, L. Anderson 711
(UTC); Juab Co., 3 miles from Trout Creek, Maguire
& Becraft 2769 (UTC); Kane Co,, 6 miles south of
(;rosvenos Arch, |. L. Reveal 809 (UTC); .57 miles east
of Kan;ib, A. Cronquist 10163 (BRY, UTC); % mile
west of Paria River Bridge along hwy 89, Welsh & At-
wood 9747 (BRY); 25 miles southeast of Escalante,
Holmgren, Reveal & LaFrance 2068 (BRY); John.son
Caiivon northeast of Kanab, D. Atwood 1791A (BRY);
Millard Co., 60 miles west of Delta, Maguire & Becraft
2768 (UTC); 20 miles west of Hincklev, B. Maguire
20761 (UTC); Desert Range Exp. Station. M, Barlow
47 (BRY); Desert Range Exp. Station, S. Brewster 22
(BRY); San [nan Co., 8 miles cast of Hite, A. Cron-
(luist 9041 (UTC); Tooele Co., Little Granite Moun-
t;iin, S. Flowers 185 (UT); Washington Co., 20 miles
sonthe;ist of Hurricane, A. Cronquist' 1(M)96 (UTC); St.
George. M.E. Jones s.n. (UTC); Anderson's Ranch, 32
miles south of Cedar City, B. Maguire 13,561 (UTC);
west slope of the Be;iverdam Mountains, D. Nish 12
(UTC); II miles south of Hurricane, F.W. Gould 1685
(ITT); St. George, M.E. Jones 16.52 (UT); St. George,
Red Hill north of toNvn, D. Hall s.n. (BRY, DIX); 1
mile southeast of Ivins Reservoir along hwv 91, L.C.
Higgins 4102 (BRY, WTSU); Santa Clara, W.P. Cot-

Biological Series, \'ol. 16, No. 3

The Bohaginaceae of Utah

15

UTAH

County Lines and
river drainages
I I I

O 8» fc»

Fii;. 14. Cn/ptanthci circiiinscissi

tarn 1164 (BRY); St. George, D. Hall 6283 (DRY);
Belveau Flats, W.P. Cottam 4674 (BRY); Snows Can-
von, D.H. Galway s.n. (BRY); 4 miles east of Harris-
burg, Welsh & Moore 6842 (BRY); near Utah-Arizona
border west of Colorado Cit> , L.C. Higgins 1380 (BRY).

6. Cryptantha crassisepala (Torr. & Gray)
Greene var. elachantha Johnst. Wrightia 2:20.
1959. Type; North end of Quitman Moun-
tains, 8 miles west of Sierra Blanca, Hud-
speth County, Texas, R. McVaugh 8040.

Erect or widely spreading herb 5-15 cm high,
stems normally numerous, loosely ascending,
branched, hirsute to hispid; leaves oblanceolate,
2-3 (6) cm long, 3-4 (6) mm wide, rounded or
obtuse, thickish, hirsute, pustulate, the upper
scarcely reduced; spikes solitary or rarely gemi-
nate, naked or few brac-ted below, 5-8 (15) cm
long, frecjuently produced from the lowest axils;
corolla inconspicuous, 1-1.5 mm wide; fruiting
calyces 6-7 (10) mm long, oblong-ovoid, some-
what asymmetrical, becoming distant below;
inahire calyx lobes linear-lanceolate, connivent
above, midrib strongly thickened and indurate,
hispid, the margins short hLspid; pedicels short,
0.5-1.2 mm long; nutlets 4 ( 1 or 2 rarely
aborted), distinctly heteromorphous; odd nutlet
next to the ;ixial calyx lolx', persistent, 2-2.5 (3)
mm long, brownish, ovoid, acute, finely granu-
late and spinular-muricate; consimilar nutlets

readily deciduous, 1.2-1.5 (2) mm long, oblong-
ovoid, thickish, coarsely tuberculate, very ob-
scurely if at all granulate, groove usually dilated
and commonly excavated to form an areola oc-
cupying much of the ventral face of the nutlet;
gynobase narrowly oblong, usually about 2/3
height of consimilar nutlets; style equalling or a
trifle exceeding the consimilar nutlets, surpassed
by odd nutlet. Sandy to gravelly places. South-
em Utah and Colorado, southward to Arizona,
Texas and northern Mexico. March-July. Fig. 15.

Carbon Co., east of Wellington, W.P. Cottam
2067 (BRY); Emery Co., 20 miles north of Green
River, B. Magiiire s.n. (UTC); San Raphael Swell, B.
Magiiire 18297 (UTC); Robbers Roost, W.A. Shands
s.n. (UT); Buckhom Wash. 22 miles southeast of
Lawerence, Pyrah. Pitts & Bamett 1.5 (BRY); 5 miles
southwest of Temple Mountain, L.C. Higgins 1327
(BRY); 13 miles east of Buckhom Wash, Welsh & At-
wood 9852 (BRY); Garfield Co., Upper Cottonwood
Wash near Cannonville, Reveal. Gentry & Davidse 757
( BRY, UTC ) ; 8 miles southeast of Escalante, Holmgren,
Reveal & LaFrance 2038 (BRY. UTC); 25 miles south
of HanksviUe. Welsh, At%vood & Higgins 8961 (BRY,
UTC); east side of Mount Ellen, W.P. Cottam 5288
(UT); Sandy Ranch, B.F. Harrison 11476 (BRY);
Willow Tank, 45 miles south of Escalante, Beck & Tan-
ner s.n. (BRY); Grand Co., K mile from Dead Horse
Point, A. Carter 1564 (UTC); northwest of Moab, A.H,
Holmgren 3.309 (UTC); entrance to Arches Natl. Monu-
ment, R. Vickery 735 (UT); 7 miles east of Cresent
Junction, FIov Canyon. W.A. Shands (UT); Arches
Natl. Monument, Pitts, Pvrah & Bamett 64 (BRY);
Arches Natl. Monument, B.F. Harrison 11127 (BRY);

UTAH

County Lines and
river drainages

A 30 io

Fit;. 15. Crifpfanthii crassisepala

16

BiUGHAM Young University Science Bulletin

Gunnison Butte, W.P. Cottam 2094 (BRY); 5 miles
west of tlie numth of Seven Mile Canyon, B.F. Harri-
son 12026 (BRY); 22 miles south of Crescent junction,
D. Atwood 2428 (BRY); Kane Co., 0.5 miles east of
Kodachrome Flat road, J.L. Reveal 802 (UTC); 57
miles east of Kanab, A. Cronquist 10157 (UTC); Glen
Canvon road across Paria River, J.R. Murdock 324
(BRY); Escalante Desert near Willow Tank, B.F, Harri-
son 9025 (BRY); 50 Mile Spring south of Escalante,
J.R. Murdock 351 (BRY); )- mile west of Paria River
bridge along hwv 89, Welsh & Atwood 9748 (BRY);
just east of Kanab, D. Atwood 1797 (BRY); San Juan
Co., 17 miles west of Blanding, B. Maguire & Redd
2061 (UTC); 7 miles northwest of Mexican Hat, B.
Maguire 16293 (UTC); west of Bluff. S. Flowers s.n.
(UT); between Moab and Monticello. W.C. Twiss s.n.
(UT); 5.5 miles north of Bluff along hwv 47, C.A.
Han.son 80 (BRY); 3 miles west of Bluff, B.F. Harrison
10.331 (BRY); vicinity of Dead Horse Point, G.L.
Richards s.n. (BRY); Island in the Skv area, G. Moore
233 (BRY); Bknding, Coft;im & Hutchings 2.504 (BRY);
Sqviaw Canvon, 1 mile west of Cave Spring, Welsh,
Moore & Canter 2834 (BRY); Canvonlands Natl. Park.
Virginia Park, G. Moore 318 (BRY); 2 miles west of
Dugout Rock, Welsh, At%vood & Higgins 8886 (BRY);
7.5 miles west of Bluff, D. Atwood 2470 (BRY); 21
miles southwest of Bluff, D. Atwood 2493 (BRY); 15
miles south of Blanding, D. Atwood 2440 (BRY);
W.ivne Co., Fremont Canvon west of Fniita, B. Ma-
guire 18116 (UTC).

7. CniptantJia decipiens (M. E. Jones) Heller,
Muhlenl:.ergia 8:48. 1912. Type: Yucca,
Arizona.

Kn/nitzkia decipiens M. E. Jones, Contr.
West. Bot. 13:6. 1910.

Stems 1-4 dm high, slender, loosely branched,
strigose and frequently short-hispid; leaves few,
linear, obtuse, 1-3 an long, 1-3 (4) mm wide,
strigose and sometimes hispid, minutely pustu-
late; spikes slender, geminate or rarely solitary or
ternate, bractless, usually becoming loosely
flowered, 4-10 cm long; corolla inconspicuous in
ours, 0.8-1 mm broad; fruiting calyx ovoid to
ovoid-oblong, strictly ascending, asymmetrical,
2.5-7 (9) mm long, deciduous, sessile; mature
calyx lobes lance-linear, decidedly connivent
above with the tips frequently spreading or even
recurving, midrib thickened and usually evident-
ly hirsute, the margins strigose, abaxial lobe evi-
dently the longest and most hirsute; nutlets 1 or
rarely 2, next the abaxial calyx lobe, ovoid-
lanceolate, 1.5-2.4 mm long, muriculate-granu-
late to tuberculate, usually brownish, back con-
vex, sides rounded, groove open or closed but
always dilated below to form a definite areola;
gvnobase half to a third as high as nutlet; style
much suipassed bv nutlet, half to two-thirds the
height of nutlet. S;uidy to gravelly slopes and
ridges. Lower Sonoran Zone. Southwestern Utah,
southern Nevada, western Arizona and southern
California. March-May. Fig. 16.

Wiishington Co., 4 miles northwest of Welcome
Springs, B. Maguire 20499 (UTC); 1 mile southwest of
St. George, south slope of Black Hill, D.H. Galway
8528 (BRY).

8. Cnjptantha duiiKtorum Greene, Pittonia
1:112. 1887. Type: California, half climbing
among bushes at Tehachapi Pass, 1884, Cur-
ran.

Krtjnitzkia duvxetorum Greene in Gray, Proc.
Amer. Acad. 20:272. 1885.

Laxly branched closely strigose herb; stems at
first erect but later commonly much elongated
and sprawling or climbing among bushes; leaves
lanceolate, thickish, 2-4 cm long, 2-4 (8) mm
wide, sparsely appressed hirsute-villous, closely
pustulate below and finely so above; spikes soli-
tary or geminate, usually remotely flowered,
5-15 cm long, occasionally with foliaceous bracts
toward base, rachis brittle and tortuously flat-
tened; corolla inconspicuous, about 1 mm broad;
fruiting calyx closely appressed to rachis, 2-3
mm long, very asymmetrical, not deciduous,
base very oblique and downwardly gibbous on
axial side; mature calyx lobes connivent and
reaching about ecjual height; 3 abaxial lobes
lanceolate, somewhat strigose, with the thickened
midribs deflexed hirsute; 2 axial lobes partly
united, hirsute only on outer margins; nutlets 4,
heteromorphous, granulate and muricate, odd

UTAH

l'"ig. 16. Criiiilanthn dccipifus

Biological Series, Vol. 16, No. .3

The Bohacin.\ce.\e ok Utah

17

mitlet persistent, axil, broadly lanceolate, 2-3
mm long, base much developed and distorting
the calyx, groove open and broad, c-onsimilar
nutlets 1.5-2 mm long, deciduous, lanceolate,
groove closed or very narrow; gynobase narrow,
style shortly surpassed by nutlets or reaching to
their tips. Sandy to gravelly deserts. Southwest-
em Utah to southern California. March-May.
Fig. 17.

W.ishington Co., Black sandy .soil just west of Ivins
Reservoir, L.C. Higgins 4130 (BRY, WTSU).

UTAH

County Lines and
river drainages

Fig. 17. Cn/ptantha dimietorum

9. Cniptantlia femUeri (Gray) Greene, Pittonia
1:120. 1887. Type: Probably New Mexico,
Fendler s.n. 1847.
Krtjnitzkui femlleri Gray, Proc. Amer. Acad.

20:268. 1885.
Eritrklmtm his^nduin var. leiocarpum Kuntze,

Rev. Gen. PI. 2:4.37. 1891.
('■ri/fitdutlui nnnulosissinui A. Nels. Ervthea

7:68. 1899.
Cryptaritha wijomingensis Gandoger, Bull.

Soc. Bot. France 65:62. 1918.

Stems erect, usually evident throughout and
bearing lateral branches mostly above the mid-
dle, soinetimes rather bushy-branched from near
the base, 1-5 dm tall, more or less densely hispid
and frecjuently appressedly so; leaves narrowly
oblanceolate to linear, 2-5 cm long, 2-4 mm

broad, appressed-hirsute, often pustulate on the
lower surface; spikes solitary or geminate, 2-12
cm long, loosely flowered, bractless or rarely
bracted below; corolla inconspicuous, about 1
mm broad; fruiting calyces ovate-oblong, 4-5
(7) mm long, ascending, slightly asymmetrical,
obscurely biserial; pedicels alxnit 0.5 mm long;
mature calyx lobes linear to lance-linear, usually
loosely connivent with the tips somewhat
spreading, midrib hirsute, thickened, margins
strigose; nutlets 4, homomorphous, or sometimes
reduced to 1-3, smooth, somewhat shiny, lanceo-
late, acuminate, 1.5-2 mm long, convex on dorsal
face, sides rounded or somewhat obtuse; groove
closed above, but at base opening into a definite
deltoid areola; gynobase subulate, twice the
length of the style, at least two-thirds height of
nutlets; style eijualling or barely surpassing the
nutlets. Sagebrush plains or Pinyon-Juniper asso-
ciation. Southern Alberta and Saskatchewan to
eastern Nebraska, northern New Mexico and
Arizona to eastern Oregon and Washington.
June-August. Fig. 18.

UTAH

County Lines and
river drainages
1 1 1

O JO v>

Fig. 18. Cn/ptaiillia fcndlcri

Duchesne Co., 5 miles soutli of Moon Lake, Ma-
guirc tk Piranian 12524 (UTC); Garfield Co., Juniper-
Hirivon belt, Holmgren & Nielsen 7747 (UTC); Iron
Co., .5 miles west Of Cedar City, B. Maguire 12979
(UTC); Kane Co., 10 miles north of Kanab, B. Ma-
guire 18771 (UTC); 6 miles north of Kiuiab, Holmgren
& Nielsen 7188 (UTC); 3 miles south of Mount Car-
inel junction, B. Maguire 18894 (UTC); 3 miles south

18

Bhigham Young Unmnehsitv Science Bulletin

of Mount Cannel junction, D. Atwood 1382 (BRY);
1 mile west of Mount Cannel junction, L.C. Higgins
3375 (BRY); OrderviUe. Cottam & Hutchings 2725
(BRY); Rich Co., south end of Bear Lake, B. Maguirc
227 (UTC); Wiishington Co.. vicinity of Pine Valley,
NIagiiire & Richards 12978 (UTC).

10. Cryptantha flaccida (Dougl. ) Greene, Pit-
tonia 1:116. 1887. Type: Probably in eastern
Oregon or Washington, collected by Douglas.
Mtjosotis flaccida Dougl. in Lehm. Pugil.

2:22. 1830.
Eritrichium oxycarytnn Gray, Proc. Amer.

Acad. n. 58. 1874.
Krynitzkui (yxycanm Gray, Proc. Amer. Acad.

20:269. 1885. '
Cryptantha mtilticaule Howell, Fl. N. W.

Amer. 1:487. 1901.
Cryptantha Howellii Nels. Bot. Gaz. 34:30.

1902.

Subsimple or ascendingly branched pallid
strigose herb 1.5-4.5 dm high; stems sparsely
closely strigose with short pallid encrusted hairs;
leaves oblanceo-linear to nearly linear of filLfonn,
2-6 cm long, 1-2 mm wide, closely strigose, strict
or ascending firm, basal portion of lower leaves
somewhat persistent; spikes quinate to solitary,
naked, usually stiffish, 4-8 (16) cm long; corolla
inconspicuous or medium sized, 1-4 mm broad;
fruiting calyces oblong-ovate, 2-4 mm long, evi-
dently asymmetrical, usually strict and closely
hugging the stem, cominonlv firm and stiff, ses-
sile or subsessUe, base broadly conic; mature
calyx lobes lance-linear, connivent above with
the tips commonly spreading, margins ciliate or
strigose, midrib thickened and armed with pale
spreading coarse encrusted arcuate or uncinate
bristles, abaxial lobe longest and most hirsute;
nutlet 1, lance-ovate, rostrate-acuminate, sub-
terete or only slightlv compressed, smooth or
very finely granulate; groove closed; style Vs to
% as high as nutlet. Washington ;ind Idaho to
southern California and east to Utah (?). April-
June. Fig. 19.

Washington Co., Black Lava Hill west of St. Ceorge,
C.W. Cottam s.n. (BRY, DIX). This collection fits the
dcscTiption of C. jUiccidu hut mav he only an immature
C. decipierus.

11, Cry])tanthn gracilis Osterh. Bull. Torrey
Club .30:236. 190,3. Type: Glenwood Springs,
Colorado, Osterliout s.n.

Cryptantha hiUmtinnii Nels. & Kenn. Proc.

Biol. S(x-. Wash. 19:157. 1906.
C-ryplantlia <ira(ilLs- var. hilhtumnii Munz &

Johnston, Bull. Torrey Club 49:39. 1922.

Stems slender, 1-2 dm high, erectly branched,
usually solitary, densely spreading short-hirsute;
leaves linear or narrowly oblanceolate, the
lower 1.5-3 cm long, 1-3 mm broad, the upper
usually much reduced, obtuse or rounded, as-
cendingly short-hirsute, minutely pustulate;
spikes solitary or geminate, usually dense, 1-2
dm long, naked; corolla minute, the limb 0.6-1
mm broad; fruiting calyx ovate, divaricate, 2-2.8
mm long, early deciduous, base evidently conical,
sessile; mature calyx lobes lanceolate, densely
appressed tawny hispid-villous, tips erect, mid-
rib sightly thickened and inconspicuously short-
hispid; nutlets 1 or rarely 2-3 and unequally de-
veloped, lanceolate, 1.5-2 mm long, 0.8-1 mm

UTAH

County Lines and
river drainages

I'ig. 19. Cnjptinil}ia jlaccidii

broad, smooth and shiny, acute, back nearly flat,
sides rounded at least near the apex, groove
mostly opened to above the middle and scarcely
forked below; style reaching to /:i-/4 height of
nutlet. Dry usually brushy slopes and ridges.
Southern Idaho and ('olorado to northern Ari-
zona and southern California. April-July. Fig. 20.

Beaver Co., Wah Wah Mountains, Pine Canyon
Pass, B. Maguire 20960 (UTC); Daggett Co., Green
River at Hideout Forest Camp, S. Flowers s.n. (UT);
Juah Co.. just nortli of the Sanpete-Juah Countv line,
I'L, Jensen .503 (UTC); Kane Co., 25 airline miles
southeast of Fscalante, Holmgren, Reveal & LaFrance
2065 (BRY, UTC); Kanah Fairgrounds, McClain s.n.
(UTC); Clay-loam ridge, B.F. Harrison 9070A (BRY);

liioLocicAL Series, Vol. 16, No. 3 The Boh.aginaceae of Utah

19

County Lines and
river drainages

i-

Fig. 20. Cn/planthii gracilis

northeast .slope of Kaiparawits south of Willow Tank,
B.F. Harrison 9()70 (BRY); Johnson Canyon, northeast
of Kanab. D. At%vood 1789D (BRY); Millard Co., 43
miles north of Beaver, Maguire & Holmgren 25076
(UTC); Black Rock Volcano west of Kanosh, C. Mc-
Millan 14.53 (UT); 10 miles west of Fillmore, W.P.
Cottam9568 (UT); San Juan Co.. Fry Canyon Mesa, L.
Wilson 23 (UTC); Canyonlands Natl. Park, Scjuaw
Flat Campground, G. Moore 279 (BRY); Sanpete Co.,
Ephraim Canyon. B. Crane s.n. (UTC); Utah Co., be-
low Timp Cave, s.n. (UT); Uintah Co., Dinosaur Natl.
Monument, ]. Brotherson 804 (BRY); Hill Creek, 12
miles south of Ouray, J. Brotherson 1063 (BRY); 3.6
miles north of Brush Creek along the Manila hwy,
L.C. Higgins 1872 (BRY); .35 miles south of Ouray
along the Hill Creek road, L.C. Higgins 1879 (BRY);
Washington Co., 10 miles east of Hurricane, W.P. Cot-
tam 8469 (UT); north of Enterpri.se, D. Hall EN35
( UTC ); Black Hill west of St. George, J.W. Harrison s.n.
(BRY); north of St. George. D.H. Galway s.n. (BRY);
Terry's Ranch on the Beaverdam Wash, E. Matthews
86 (BRY).

12. C rtjptantha inaequata Johnston, Univ. Calif.
Publ. Bot. 7:444. 1922. Type: Among rocks.
Pleasant Canyon, Panamint Mountains, Hall
& Chandler 692.5.

JolmstotieUu inaequata Brand, Rep. Spec.
Nov. 21:250. 1925.

Stems erect or ascending, 3-4 dm high,
branched throughout, or sometimes the basal
branches elongated and simple or nearly so, his-
pid and strigose or hirsute toward the base;
leaves linear to narrowly-oblanceolate, 2-4 cm

long, acute, often becoming more or less convo-
lute, more or less hispid, pustulate, especially be-
neath; spikes geminate or solitary, 4-12 cm long,
;it times sparsely bracted below; corolla incon-
spicuous, the tube shorter than the calyx, limb
1-2.5 mm broad; fruiting calyx ovoid-oblong, 2.5-
.3 mm long, ascending; pedicels less than 0.5 mm
long; mature calyx-lobes lanceolate, midrib
thickened and hirsute, villous-ciliate on the mar-
gins; nudets 4, heteromorphous, triangular-ovate,
dark with small pale tuberculations, margins
acute, groove closed above but below gradually
enlarging into a shallow triangular areola, odd
nutlet about 1.7 mm long, somewhat persistent,
slightly lighter than the others, next the abaxial
calyx-lobe; consimflar nutlets about 1.3 mm long;
gynobase equalling consimilar nutlets but sur-
passed by odd nutlet; style much surpassing the
nutlets. Among rocks and shrubs. Southwestern
Utah to northwestern Arizona and southeastern
California. March-May. Fig. 21.

UTAH

County Lines and
river drainages

l"ig. 21. CrtipUiniha incieijtuita

Washington Co., Black Lava Hill west of St.
George, C.W. Cottam s.n. (DIX); Black Hill, St.
George, Field Biology Class s.n. (UT); clay hills north-
west of new golf course southwest of St. George, N.D.
Atwood 1707 (BRY).

13. Crijptantha kehet/ana Greene, Pittonia 2:232
1892. Type: ElUston, Montana, August 1889,
Greene s.n.

20

BnlCHAM VoUNC: 1'm\ KHMI Y SCIENCE BULLETIN

Spreading or ascending hirsute herb 5-25 cm
high; stems 1-several, hirsute and also hispid-
strigose; leaves linear or narrowly oblanceolate,
1.5-3 (4) cm long, 2-4 mm wide, rounded or ob-
tuse, thickish, hirsute, pustulate, the upper ones
scarcely reduced; spikes usually solitary, 4-9 cm
long, naked or with a few bracts near the base;
corolla minute, 1-2 mm broad; fruiting calyces
4-6 mm long, ovate-oblong, spreading, loose or
dense, somewhat asymmetrical; pedicels short
but definite, about 0.8 mm long; mature calyx-
lobes linear, weakly connivent above, midrib
thickened, hirsute to hispid, margins inconspicu-
ously villous-strigose; nutlets 4, heteromorphous;
odd nutlet next to the axial calyx-lobe, broadly
lance-ovoid, 2-2.6 mm long, smoothish or granu-
late or muriculate-granulate or rarely somewhat
tuberculate, standing off slightly from the gyno-
base, consimilar nutlets lance or oblong-ovoid,
1.8-2.3 mm long, coarsely tuberculate and usual-
ly granulate darker than the odd nutlet, sides
rounded, groove narrow or closed, near base
abruptly dilated to form a small triangular ar-
eola; gynobase subulate, a little longer than the
style, V2 to % height of consimilar nutlets; style
surpassed by odd nutlet and just surpassing or
even exceeded by consimilar ones. Sandy plains
to rocky hillsides. Saskatchewan and Montana
southward through Wyoming to northern Colo-
rado and Utah. May-July. Fig. 22.

County Lines and
river drainages

Fig. 22. Cn/ptuitthi kcl.sci/<nui

Grand Co., 1.3 miles due southwest of Moab, Har-
rison, Marshal & Nielson 10285 (BRY); Porcupine Ridge,
La Sal Mountain.s, J. Pederson 114 (BRY); Tooele
Co., 10 miles southeast of Cline, W.H. Hirst 36
(UTC); Uintah Co.. 13 miles west of Vernal, R.S.
Ferris 11384 (UTC); 8 miles west of Bitter Creek on
Ourav-Bonanza Road, N.D. Atwood 1556A (BRY); 20
miles south of Bonanza, L.C. Higgins 1079 (BRY).

14. Crtjj)tontlui micrantlia (Torr. ) Johnst. Contr.
Gray Herb. 68:56. 1923. Type: "Sand hills,
Fontera, Texas, and in other places along
the Rio Grande."

Eritrichuim micranthum Torr. Bot. Mex.
Bound. 141. 1859.

Kn/nitzkia micrantlia Gray, Proc. Amer.

Acad. 20:275. 1885.
Eremocartja micrantlia Greene, Pittonia 1:59

1887.
Eremocanja mtiricata Rydb. Bull. Torrey

Club 36:677. 1909.

Slender, dichotomously branched herb 5-15
cm high, drying brownish; root and lower parts
of the stem stained with a purple dye; leaves ob-
long-oblanceolate, 3-7 mm long, 0.8-1.4 mm
wide, whitish-strigose or short hirsute, rounded
at apex, uppennost scarcely reduced and extend-
ing through the inflorescence; spikes numerous,
solitary or geminate, densely flowered and
strongly unilateral, leafy-bracted throughout, 1-4
cm long; corolla inconspicuous to medium-sized,
limb ()..5-2.5 mm broad; fruiting calvx ovoid-
oblong, 1.8-2.5 mm long, slightly asymmetrical,
decidedly biseriate, base broadly conic;il; pedi-
cels 0.5-0.8 mm long; mature calyx lobes oblong-
lanceolate, broad, erect, hirsute, midrib not evi-
dently thickened; nutlets 4, 1-1.3 mm long,
smooth or tuberculate, 1 nutlet sometimes a little
longer and more persistent than the others,
groove extending full length of nutlet, narrow,
scarcelv broadened at base; gynobase subulate,
nearly as long as the calyx, much surpassing the
nutlets and bearing at its summit the sessile stig-
ma. Dry sandv soils in the Lower Sonoran Zone.
Southeastern Oregon to Utah, southward to Baja
California, Arizona, New Mexico, and Texas.
March- June. Fig. 23.

(iarfield Co.. Egnog Spring. Bullfrog Creek, west
of Heiirv Mountains, S.L. Welsh .3983 (BRY); Kane
Co., 23 miles south of Alton, A. Cron(|uist 10192 (BRY,
UTC); sand dunes nortli of Kanal), W.P. Cottam 4304
( BRY ) ; )■: mile west of P;iria River bridge along hwy
89, Wclsli *( Atwood 9746 (BRY); Washington Co., 12
miles northeast of St. George, A. Cron(|uist 10103 (BRY,
tITC); Be;iverdam Mountains, D. Nish 64 (UTC); 8
miles southwest of St. George, Holmgren & Thieret
8229 (UTC); !4 mile north of To<iuerville, B. Maguire
2047.5 (UTC); 32 miles south of Cedar Citv. B. Ma-
guire 13.562 (UTC); St. George, M.E. Jones s.n.

Biological Series. Vol. 16, No. 3

The Bohaginaceae ok Utah

21

County Lines and
river drainages

i-

3« ^

Fig. 23. Cn/ptuntha 7iiicrantha

(UTC); Diamond Valley, W.P. Cottam 4034B (BRY);
St. George, D.H. Galway 6284 (BRY); 5.7 miles .south
of Gunlock, F.W. Gould 1594 (BRY); Terry's Ranch on
the Beaverdam Wash, L.C. Higgiiis 406 (BRY); Bea-
verdam Mountains near the Utah-Arizona border, L.C.
Higgins .344 (BRY); DLxie State Park, 5 miles north-
west of Ivins, L.C. Higgins 857 (BRY); 4 miles east of
Harrisburg, south of the Virgin River, Welsh & Moore
6841 (BRY); 1 mile southeast of Ivins Reservoir along
hwv 91, L.C. Higgins 4103 (BRY); about 4 miles west
of Hurricane, L.C. Higgins 4214 (BRY).

15. Cnjptantha nevadensis Nels. & Kenn. Proc.
Biol. Soc. Wash. 19:157. 1906. Type: Reno,
W;i.shoe Co., Nevada.
Kryiiitzkiu harhigera var. inops Brandg. Zee

5:228. 1906.
Cry})tantlui barbigera var. inops Macbr.

Proc. Amer. Acad. 51:548. 1916.
CniptantJui arenicola Heller, Muhlenbergia

'2:242. 1906.
Cnjptantha IcptophijUa Rydb. Bull. Torrey

Club 36:678. 1909.

Stem.s .slender, 1-5 dm high, 1 to several, erect
or usually flexuous, appressed short-strigose,
mostlv hixlv ])ranched; leaves linear-oblanceolate
to linear, acute or obtuse, 1-4 cm long, 1-5 (7)
mm broad, not numerous, appresse^i short-his-
pid, more or less pustulate; spikes geminate or
temate, tenninal, also scattered along the stem
on short, slender, axillary branches, occasionally

bracted toward the base, congested and some-
what glomerate or elongate and becoming 15
cm long; corolla inconspicuous, limb 1-2 mm
broad; fruiting calyx oblong-ovoid to lanceolate,
5-12 mm long, ascending, slightly asymmetrical;
pedicels about 0.5 mm long; mature calyx-lobes
lanceolate or hnear, connivent above with the
slender tips usually recurving, margins more or
less villous and hispid on the somewhat thicken-
ed midrib; nutlets 4, homomorphous, verrucose
or muriculate toward the tip, lanceolate to lance-
ovoid, 2-2.9 mm long, back convex, margins
somewhat angled, groove open or closed, dilated
below into a small areola; gynobase narrow;
style reaching to or almost to the tips of the
nutlets. Sandy to gravelly slopes and washes in
the Lower Sonoran Zone. Deserts of Utah and
Nevada to Arizona and Baja California. March-
June. Fig. 24.

UTAH

County Lines and
river drainages

Fig. 24. Crijptantha neviulcnsi.s

San Juan Co., 10 miles east of Hite, A. Cronquist
9036 ( UTC ) ; vicinity of Mexican Hat, B. Maguire
16277 (UTC); WiLshington Co., west slope of the Bea-
verdam Mountains, D. Nish 32 (UTC); 2 miles west of
Rockvillc, B. Maguire 4979 (UTC); 1 mile ea.st of Hur-
ricane, Maguire & Blood 1529 (UTC); north of St.
George. D.H. Galway s.n. (DIX); Beaverdam Moun-
t;iins near the Utah-Arizona border along hwy 91, L.C.
Higgins 333 (BRY); Black Hill west of St. George, D.H.
G;ilway s.n. (BRY); Terry's Ranch on the Beaverdam
W:ish,' L.C. Higgins 303 (BRY); Dixie State Park, 5
miles northwest of Ivins, l,.C. Higgins 852A (BRY).

Bbioham Young University Science Bulletin

16. Cnjptantha patlersonii (Gray) Greene, Pit-
tonia 1:120. 1887. Type: Without locality,
1875 Patterson, but probably came from the
Rocky Mountains near Golden, Colorado.
Krynitzkia pattersonii Gray, Proc. Amer.
Acad. 20:268. 1885.

Stems usually several 1-1.5 dm high, loosely
branched, hirsute and somewhat strigose; leaves
oblanceolate, 1-3 cm long, 2-4 mm broad, rather
firm, obtuse, hirsute with somewhat pustulate
bases, upper ones little reduced; spikes soUtary
or geminate, naked, 2-5 (7) cm long; corolla
inconspicuous, 1-1.5 mm broad; fruiting calyx ob-
long-ovoid, 4-5 mm long, spreading, slightly a-
symmetrical, evidently biseriate, lower-most be-
coming 2-6 mm distant; pedicels about 0.5 mm
long; mature calyx-lobes linear-lanceolate, tips
more or less connivent, the midrib hirsute and
thickened, margins appressed hispid; nudets 4,
heteromorphous, odd nudet next to the axial ca-
lyx-lobe, slightly the largest, about 1.9 mm long,
ovate, acute, smooth or obscurely rugulose or
sparsely tuberculate, somewhat persistent, stand-
ing off slightly from the gynobase; consimilar
nutlets oblong-ovoid, about 1.6 mm long, decidu-
ous, closely appressed to gynobase, smooth, back
convex, sides rounded or obtuse, groove opened
or closed and abruptly broadened below into a
small triangular areola; gynobase narrow, reach-
ing to about 73 height of consimilar nutlets; style
exceeded by odd nutlet, equalling or a little
shorter than consimilar nutlets. Dry, usually
somewhat sandy or gravelly soil, mostly in the
mountains. Wyoming, northern Colorado, and
northeastern Utah. May-July. Fig. 25.

Duche.sne Co., 10 miles north of Altonali, J.
Brotherson 651 (BRY).

C. pattersoni is rarely collected. It is also very
closely related to C. kehetjana and C. ambigua
and intergrades with both. In habit it is similar
to C. amhigxia and immature specimens ane
nearly impossible to distinguish.

17. CnjptantlHi pterocanja (Torr. ) (Ireene, Pit-
tonia 1:120. 1887.

Stems erect, branched throughout with as-
cending branches, 1-5 dm high, short hirsute
with either appressed or ascending slender stri-
gose hairs; leaves linear or the reduced upper-
most ones lanceolate or oblong, strigose, 1-2.5
(4) cm long, 1-3 (5) mm broad, obtuse, con-
spicuously pustulate below but usually finely
so above; spikes geminate or rarely temate or
solitary, naked or bracted below, 2-6 (12) cm
long, becoming loosely flowered, corolla incon-

spicuous, 0.5-1 (2) mm broad; fruiting calyces
distinctly accrescent, (2) 3-5 mm long, sym-
metrical, ascending on short pedicels 0.5-1 mm
long; mature calyx-lobes ovate to lanceolate,
somewhat connivent, thin, margins more or less
tawny, appressed hispid, midrib slightly thicken-
ed and sparsely hispid; nutlets 4, homomorphous
and all winged, or heteromorphous with axial
nutlet wingless; body of nutlet oblong-lance-
olate or lanceolate, 2-2.5 (3) mm long, muricate
or verrucose, wing-margin of nutlet broad or nar-
row, entire or crenate or lobed, extending com-
pletely around the nutlet or only down the sides,
groove open or closed ( even in the same plant )
and dilated below into an open excavated areola;
gvnohase slender, about % height of nutlets;
style subulate, slightly surpassing or somewhat
surpassed by the wing-margin of the nutlets but
always exceeding the body proper.

1. Nutlets heteromorphous. a.xial one wingless

17A. var. ptewcarya

1. Nutlets homomorphous, all winged

17B. var. cijcloptera

17.\. CrtiptiiHlJia pterocanja ( Torr.) Greene var.
pterocanja. Type: Walla Walla, Washing-
ton, as designated by I. M. Johnston,
Contr. Gray Herb. 74:53. 1925.
Eritrichium pterocanpnn Torr. Bot. Mex.
Bound. 142. 1859. "

County Lines and
river drainages

Is-

Fig. 25. Crijpuinthd )><ittcr.to»ii

BroLOCiCAL Series, Vol. 16, No. 3 The Boracinaceae of Utah

23

Eritrichium pterocanjum var. pectiimtum
Gray, Proc. Amer. Acad. 10:61. 1874.

Krtjnitzkici ptcwcanjum var. pectiniita
Gray, op. cit. 20:276. 1885.

KrynitzkUi pterocanja Gray, Proc. Amer.
Acad. 20:276. 1885.

Nutlet.s licteromorphous, axial one wingless.
Sandy or gravelly places, Upper and Lower
Sonoran Zones. Southern Washington, southern
Idaho, Utah, Arizona, and northern Sonora.
April-June. Fig. 26.

UTAH

County Lines and
river drainages

Cnjptantha pterocan/a wir. ptcrocuiijii

Beaver Co., Warm Point Ridge, Stahmann &
Hutchiiigs 67 (UT); Garfield Co., 4 iiiile.s northeast of
Hite, A. Cronquist 9053 (UTC); Grand Co., west of
Moab, E.L. Miner 1 (UT); confluence of Delores &
Colorado Rivers, Vickery 287 (UT); Juab Co., 7 miles
east of Trout Creek, B. 'Vlaguire 2777 (UTC); 39 miles
west of junction of Utah hwy 50-6 and Jericho road
along the Callao Road, K.D. Kaneko .50 (BRY); Kane
Co.. 25 airhne miles southeast of Escalante, Holmgren. Re-
veal & LaFrance 2064 (UTC); 57 miles east of Kanab,
A. Cronquist 101.55 (BRY, UTC); 23 miles south of
.\lton, A. Cronquist 10193 (BRY. UTC); Johnson Can-
yon, northeast of Kanab, N.D. Atwood 1789A (BRY);
Millard Co., Black Rock Volcano west of Kanosh, C.
McMillan 1484 (UT); 43 miles north of Beaver, Ma-
guire & Holmgren 2.5075 (UTC); Dessert Range Exper.
Station, Maguire & Holmgren 25109 (UTC); White
Sage Vallev. B.F. Harrison 6377 (BRY); east side of
Pine Vallev, B.F. Harrison 6.393 (BRY); Warm Point
Ridge, Stahmann & Hutchings 50 (BRY); San Juan
Co., 2 miles west of Bluff, B. Maguire 1.3525 (UTC);
Comb Wash, L. Amow 231 (UT); 35 miles west of

Bluff, B.F. Harrison 11947 (BRY); 1 mile east of Moab
bridge, Deming 1-24 (BRY); 15 miles south of Bland-
ing, N.D. Atwood 2439 (BRY); Sevier Co., 2 miles
north of Elsinore, Reveal, Gentry & Davidse 735 ( BRY,
UTC); Washington Co., St. George, D.H. Galway s.n.
(UT); Di;imond V:Uley, F.W. Gould 1705 (BRY, UT);
St. George, M.E. Jones 2034 (UT); Zion Natl. Park, B.
Maguire 13575 (UTC); 1 mile east of Hurricane, Ma-
guire & Blood 1534 (UTC); 5 miles west of Springdale,

A. Cronquist s.n. (UTC); 32 miles south of Cedar City,

B. Maguire 16280 (UTC); Price Bench, R. Hardy s.n.
(DIX); St. George, C.W. Cottam s.n. (BRY, DIX);
Diamond Vidley, D.H. Galway s.n. (BRY); Diamond
Valley, W.P. Cottam 4044 (BRY); 4 miles east of Har-
risburg, Welsh & Moore 6843A (BRY); 15 miles south
of Enterprise, L.C. Higgius 3480 (BRY); Dixie State
Park, L.C. Higgins 934 ( BRY ) ; Beaverdam Mountains,
near the Utah-Arizona border along hwy 91, L.C. Hig-
gins 3.34 ( BRY ) ; about 5 miles west of hwy 91 along
the Jackson road, L.C. Higgins 501 (BRY); 4 miles west
of Hurricane. L.C. Higgins 4218 (BRY); Diamond Val-
lev. L.C. Higgins 4199 (BRY, WTSU).

17B. Cnjptantha pterocanja var. cycloptera
(Greene) Macbr. Contr. Gray Herb. 48:44.
1916. Type: Near Tucson, Arizona. 1884
Pringle.
Knjnitzkia cycloptera Greene, Bull. Calif.

Acad. Sci. 1:20. 1884.
Cryptantha cycloptera Greene, Pittonia
1:120. 1887."

Nutlets homomorphous, all winged, other-
wise like the typical material. Sandy or gravelly
deserts. Southern California east to southern
Utah, Arizona, New Mexico, and southwestern
Texas. April-June. Fig. 27.

Kane Co., Kaiparowits Plateau, about 25 airline
miles southeast of Escalante. Holmgren, Reveal & La-
France 2064 (BRY. UTC); red ledges, north of St.
George, D.H. Galway s.n. (BRY).

18. Cryptantha racemosa (Wats.) Greene, Pit-
tonia 1:115. 1887. Type: Ehrenberg, Ari-
zona.
Krynitzkia racemosa Greene, Bull. Calif.

Acad. Sci. 1:208. 1885.
Eritrichium racemosum Wats, in Gray, Proc.

Amer. Acad. 17:226. 1882.
Krynitzkia ramosissima Gray, Proc. Amer.

" Acad. 20:277. 1885.
Cryptantha suffniticosa Piper, Proc. Biol.

Soc. Wash. 32:42. 1919.
Cryptantha racemosa var. lignosa Johnst.

Univ. Calif. Pub. Bot. 7:445. 1922.
Johnstonella racemosa Brand, Rep. Spec.

Nov. 21:249. 1925.
Johnstonella racemosa var. lignosa Brand,

Rep. Spec. Nov. 21:249. 1925.

Long-lived annual, often decidedly suffruti-
cose near the base, 1-10 dm high; stems single

24

Bric.ham Young University Science Bulletin

County Lines and
river drainages

L 1 I

o j5 *•

Fig. 27. Cryptantha pterocanja var. cycloptera

with numerous ascending branches or many and
diffusely branched younger parts, green, strigose
and commonly hirsute, epidermis at length ex-
foliating leaving the older woody stems glabrous
and brown; leaves oblanceolate, acute, hirsute,
pustulate, the early ones 3-6 cm long, 6-12 mm
broad; racemes apparently forked and panicu-
latelv disposed, minutely bracteate, 3-15 cm
long; corolla very inconspicuous, limb about 1
mm wide; fruiting calyces oblong-ovoid, 2-4 mm
long, slightly asymmetrical, ascending, tardily
deciduous; pedicels usually well developed, 1-4
mm long, slender, frequently nodding; mature
calyx-lobes lance-linear, somewhat strigose, hir-
sute along the thickened midrib; nutlets 4, het-
eromorphous, triangular-ovate, the acute tips
slightly outcurved, groove open or closed above
but below broadening out into a shallow broad-
ly triangular areola; odd nutlet next to the abax-
ial calyx-lobe, 1-2 mm long, somewhat persistent,
finely muricate or tuberculate, light or dark,
consimilar nutlets 0.8-1.5 mm long, acute, tend-
ing to be very narrowly wing-margined, dark
with pallid tuberculations; gynobase subulate,
% length of odd nutlet and about ecjualling con-
similar nutlets; style much surpassing the nut-
lets. Sandy flats and rcK-ky ridges of the Lower
Sonoran Zone. Southwestern Utah, southern
Nevada, western Arizona, and southwest to Baja
California. March-June. Fig. 28.

Washington Co., Black Lava Hill west of St.
George, J. W. Harrison s.n. (DIX).

19. Cryptantha rectirvata Coville, Contr. U.S.
Natl. Herb. 4:165. 1893. Type: Surprise
Canyon, Panamint Mountains, Coville &
Funston 713.

Stems branched from the base, slender, as-
cending or decumbent at base, 1-3 dm high, stri-
gose, root often dye-stained, basal leaves ob-
lanceolate or spatulate, 1.5-2 cm long, those of
the stem remote, linear or lanceolate, 5-10 mm
long, rounded or obtuse, rather finely appressed-
hispid and minutely pustulate; spikes bractless,
slender, loose, 2-10 cm long, solitary or gemi-
nate; corolla inconspicuous, about 2 mm long,
subtubular, not exserted, lobes short; fruiting
calyx slender, asymmetrical, bent and recurved,
■3-4 mm long, tardily deciduous, sessile; mature
calyx-lobes linear, midrib somewhat thickened
and hirsute, rarely merely strigose, axial lobe
longest, thickest and most hirsute; ovules 2, nut-
let 1, subpersistent, oblong-lanceolate, curved
inwardly, dull brownish, granulate-muricate,
next to the axial calyx-lobe, edges obtusish,
groove somewhat oblique, narrow or closed,
opening into a small basal areola; gynobase
slender, about J2 the length of the matured nut-
let and about as long as the abortive ones; style
commonly much surpassed l^y nutlet. Sandy or

County Lines and
river drainages

Fig. 28. Cii/pliiiillia iiicciii<is<i

Biological Series, Vol. 16, No. 3 The Boraginaceae of Utah

25

UTAH

County Lines and
river drainages

Cnjptanthd rcctii vata

gravelly slopes and wash bottoms. Southeastern
Oregon to Utah, Nevada, and southern Califor-
nia. April-June. Fig. 29.

Beaver Co., 26 miles north of Milford, B. Maguire
21034 (UTC); Juab Co., 13 miles east of Trout Creek,
Maguire & Becraft 2771 (UTC); north of Fumarole,
W.P. Cottam 9647 (UT); Emery Co., Temple Moun-
tain, \'.l'. AUman s.n. (BRV); Millard Co.. 20 miles
west of Hinckley, B. Maguire 20760 (UTC); 60 miles
west of Delta, Maguire & Becraft 2774 (UTC); Desert
Range E\per. .Station, R.C. Holmgren 461 (BRY); Kane
Co., 6 miles east along road to Warm Creek, from Glen
Canyon City. Welsh & Atwood 9811 (BRY); N.E. .slope
of Kaiparawits, south of Willow Tank, B.F. Harrison
9070B (BRY); San Juan Co., 18 miles northwest of
Blanding. A. Cronquist 9016 (UTC); 10 miles west of
Bluff, B. Maguire 1.3.541 (UTC); Cottonwood Canyon,
Waylands Ranch, B. Maguire 16271 (UTC); 16 miles
southwest of Bluff on hwy 163, N.D. Atwood 2474
(BRY); Comb Wash, 7..5 niiles west of Bluff, N.D. At-
wood 2469 (BRY); along road just west of Monument
Valley Hospital, N.D. Atwood 2.512 (BRY); 15 mUes
south of Bhmding on road to Bluff, N.D. Atwood 24.38
(BRY); Washington Co., 12 miles northeast of St.
George, A. Cronquist 10116 (BRY, UTC); north of
St. George, D.H. Galwav s.n. (DIX); Black Hill west
of St. George, D. Hal! s.n. (BRY); 4 miles east of
Harrisburg, Welsh & Moore 6843 (BRY); 4 miles west
of Hurricane. L.C. Higgins 4215 (BRY, WTSU); 1
mile southeast of Ivins Reservoir ;dong hwy 91 L.C.
Higgins 4126 (BRY, WTSU).

Cniptantha murictilata var. montana Nels.
Erythea 7:69. 1899.

Stems with several to many stiffly erect
branches 1-3.5 dm high; stems closely short-stri-
gose and frequently also sparsely hispid; leaves
linear to lance-linear, 2-4 cm long, obtuse, 1-3
mm broad, strictly ascending, strigose or ap-
pressed hirsute, finely pustulate, spikes stiff,
bractless, solitary or geminate, 2-10 cm long;
corolla inconspicuous, about 1 mm wide, tube
about etjualling calyx; fruiting calyx ovate-ob-
long, 5-6 mm long, strictly ascending subsessile,
slightly asymmetrical, becoming rather obscurely
biserial at maturity and 5-10 mm distant; ma-
ture calyx-lobes linear-lanceolate, rather stiff,
somewhat connivent above with the tips slightly
spreading, midrib thickened and hirsute, mar-
gins ascending-hispidulous, axial lobe slightly
the longest; nutlets 4, homomorphous, lanceolate
to broadly lanceolate, 1.8-2.2 mm long, antrorse-
ly spinulose-muriculate, especially toward the
apex, margins and base rounded, groove narrow
and forked below where occasionally open to
form a small triangular areola; gynobase subu-
late, about y^ height of nutlets; style reaching to
tips of nutlets. Dry sagebrush plains, usually in
sandy soils. Washington and eastern Oregon, al-
so in Idaho, Wyoming, and northern Utah. May-
July, Fig. 30.

Weber Co., Ogden Hot Springs. Stokes s.n. (UC).

UTAH

County Lines and
river drainages

i-

20. Cnjptantfia scoparw \. Nels.
.54:144. 1912. Type: Minidoka,
Nelson & Macbride 1311.

Bot. Gaz.

Wyoming,

Fig. .30. ('n/j)l(inthii scopariii

26

BniGHAM Young Univehsitv Science Bulletin

21. Cryptantha torreyana (Gray) Greene, Pit-
tonia 1:118. 1887. Type: Yosemite Valley,
Torrey 337.
Krynitzkia torreyaiui A. Gray, Proc. Amer.

Acad. 20:271. 1885.
Krynitzkia torreyaiui var. calycosa Gray, loc.

cit.
Cnjptantha torreyana var. calycosa Greene,

Pittonia 1:119. 1887.
Cryptantha calycosa Rydb. Mem. N. Y. Bot.

Gard. 1:3,31. 1900.
Cryptantha affinis var. flexuosa Nels. Bot.

Gaz. 30:195. 1900.
Cryptantha flexuosa Nels. New Man. Bot.

Cent. Rocky Mts. 416. 1909.
Cryptantha torreyana var. calistogae Johnst.

Contr. Gray Herb. 74:80. 1925.

Stems erect, 1-4 dm high, solitary or several
with erect or more often spreading branches,
finely strigose and sparsely hirsutulous; leaves
oblanceolate to linear, strict or ascending, 2-5
( 7 ) cm long, 3-6 ( 8 ) mm wide, obtuse or round-
ed, hispid, inconspicuously pustulate if at all so;
spikes commonly geminate, bractless, 4-8 (15)
cm long, more or less projected from the leafy
mass of the plant, very elongate and loosely
flowered or congested and glomerate; corolla in-
conspicuous, about 1 mm broad; fruiting calyx
ovoid or oblong-ovoid, 2-7 mm long, ascending,
asymmetrical, base rounded or broadly conic;
pedicels about 0.5 mm long; mature calyx-lobes
lanceolate to linear-lanceolate, connivent above
with tips usually spreading, midrib slightly thick-
ened and hispid-hirsute, margins hispid-strigose;
nutlets 4, occasionallv 1 or more aborted, usually
broadly ovate, 1.5-2.2 (2.5) mm long, 0.8-1.3 mm
broad, smooth and polished, usually mottled,
rarely finely granulate, back low and convex,
sides rounded or obtuse, groove broadly forked
below and closed throughout; gynobase about li
height of nutlets, about 1 mm tall; style reaching
to % height of nutlets or rarely even to their tips.
Open slopes or sometimes growing in partial
shade, mainly transition zones. Extreme western
Wyoming and northern Utah to northern Cali-
fornia and northward to British Columbia and
Alaska. May- August. Fig. 31.

Box Elder Co.. Ruft River Mountains, Dunn Can-
yon, Maguire & Holmgren 222.3.3 (UTC); Cache Co.,
tony Crove Lake, B. Maguire 2166.5 (UTC); 12 miles
southwest of l.aketown, Harrison & l.arsen 7963 (BRY);
Intervale in Providence Canyon, B. .Vlaguire 138.38
(UTC); Logan Canyon, H.B. Pa.ssey 108 (UTC);
Blacksmith Fork Canyon, B. Maguire .3701 (UTC);
Wellsville Range, Pine Canyon. B. Maguire 3110
(UTC): Davis Co., Lagoon, s.'n. (UT); Salt Lake Co.,
north slope of Mount Olympus, Vickery 8.32 (UT); Red
Butte, A.O. Garrett 1869 (UT); Red Butte Canyon, L.

.\mow 1534 (UT, BRY); Sanpete Co., vicinity of
Ephraim, R. Olsen s.n. (UTC); Sevier Co., 5 miles
southeast of Sigurd, B. Maguire 18097 (UTC); Tooele
Co., South Willow Creek, B. Maguire 21757 (UTC);
Utah Co., nortli fork of Provo River, S. Flowers 24
(UT); Aspen Grove, A.O. Garrett 3448 (UT); Mount
Timpanogos, A.O. Garrett .5551 (UT); 2 miles below
Aspen Grove, B.F. Harrison 9392 (BRY); Mount Tim-
panogos loop road, E. Larsen 7 16.5 ( BRY ) ; Aspen
Grove. B.F. Harrison 6776 (BRY); American Fork Can-
yon, LE. Diehl D3 (BRY); hills around Castilla, LE.
biehl D4 (BRY); Provo Canyon near Deer Creek, D.H.
Galway s.n. (BRY); Aspen Grove, J.D. Walker s.n.
(BRY); head of Rock Canyon, B.F. Harrison 9403
(BRY); Washington Co., Pine Valley, W.P. Cottam
8916 (UT); Brouse Area Ranger Station, R.K. Gierisch
.332 (UTC); Weber Co., near Ogden, A.O. Garrett
6278 (UT).

29

Cryptantha utahensis (Gray) Greene, Pit-
tonia 1:120. 1887. Type: St. George, Palmer
.352.
Krynitzkia utahensis Gray, Syn. Fl. N. Amer.

2:427. 1886.
Eritrichium holopterum var. suhmolle Gray,

Proc. Amer. Acad. 13:374. 1878.
Cryptantha suhmollis Coville, Contr. U.S.

Natl. Herb. 4:166. 1893.

Plant usually with a main erect stem with a
few scattered ascending or erect branches, 1-3
dm high, strigose or appressed short-hispid;
leaves not numerous, strongly reduced above.

County Lines and
river drainages

ir-

I'ig. 31. Cnfptdiitltii torritfdii'i

Biological Series, Vol. 16, No. 3 The Boracinaceae of Utah

27

linear to oblanceolate-linear, 1-5 (7) cm long,
1-4 mm wide, rounded at apex, commonly pus-
tulate and .short-hirsutv' especiallv heneatli; spikes
usually geminate, commonly 1-2.5 (5) cm long,
dense, bractless; corolla rather conspicuous, 2-3
mm broad; fruiting calyx ovoid or ovoid-oblong,
2-3 (4) mm long, subsessile, spreading or some-
what recurved, deciduous, usually densely ap-
pressed hirsute and silky; mature calyx-lobes
lanceolate, connivent, midrib thick and usually
brownish, and frequentlv bearing spreading or
recurved hairs; ovules 4. nutlets 1 or rarely 2,
next to the abaxial calyx-lobe, 1.7-2.5 mm long,
1-1.5 mm broad, pale, broadly lanceolate, granu-
late, muricate-papillate or rarely spinulose, back
low-convex or flat, margins sharply angled or
with a verv narrow knifelike margin, groove
open, narrow, opening into a small areola below;
gynobase subulate, about % height of nutlet, not
markedly differentiated from style; style usually
a trifle shorter than the nutlet. Desert washes
and ridges, sandy to rocky soils. Southern Utah
and western Arizona, then westward into south-
em Nevada and the desert regions of California.
March-May. Fig. 32.

Washington Co.. St. George, Palmer 352 (G); 1
mile east of Hurricane, Maguire'& Blood 1532 (UTC); 2
miles north of St. George, E. Cox s.n. (DIX); Terry's
Ranch on the Beaverdam Wash, A.H. Bamum 988
(DIX); Terry's Ranch, L.C. Higgins 1225 (BRY);
Castle Cliffs, along hwy 91. L.C. Higgins 4168 (BRY.

UTAH

County Lines and
river drainages

i-

Fig. 32. Cnjptiinthii utahcnsis

WTSU); Terry's Ranch on the Beaverdam Wash, L.C.
Higgins 292 (BRY).

2.3. Cryptantha watsoni (Gr Ay) Greene, Pittonia
1:120. 1887. Type: Utah, Wasatch Moun-
tains, Watson 858.
Krynitzkia watsoni Gray, Proc. Amer. Acad.

20:271. 1885.
Cryptantha vinctens Nels. & Macbr. Bot.

Gaz. 62:143. 1916.

Slender strictly branched hispid herb 1-3 dm
high; stems solitary, sparsely or loosely branched,
spreading short-hispid; leaves linear to oblanceo-
late, 1-4 (5) cm long, 1-4 (5) mm broad, obtuse
or rounded, ascending hispid, and merely pustu-
late; spikes solitary or in pairs, 1-4 (6) cm long,
bractless or rarely bracted below; corolla incon-
spicuous, about 1 mm broad; fruiting calyx ovoid
to oblong-ovoid, 2-3.5 (4) mm long, subsessile,
rounded at base, early deciduous, oldest ones
becoming distant; mature calyx-lobes lanceolate,
tips usually connivent, hirsute with ascending
hairs, the midrib also with a few spreading
bristles and scarcely thickened; nutlets 4, homo-
morphous or practically so, lanceolate, 1.5-2 mm
long, about 0.8 mm broad, smooth, shiny or at
times dulled by minute granulations, back nearly
flat, margins definitely angled, groove closed
or nearly so and forked at base; gynobase subu-
late, about % height of nutlets; style equalling
nutlets or a trifle surpassed by them. Sandy or
rocky slopes and plains of the arid Transition
Zone. Eastern Washington, south through Ore-
gon and Nevada to California, and east to Mon-
tana, Wvoming, and Colorado. May-September.
Fig. .3.3. ■

Beaver Co., 11.5 miles east of Milford, B. Maguire
21013 (UTC); Juab Co., Deep Creek Mountains, Ma-
guire & Becraft 2775 (UTC); Tooele Co., near the
Great Salt L;ike, A.O. Garrett 2864a (UT); Utah Co.,
Rock Canyon. A.O. Garrett .3316 (UT).

24. Cryptantha abata Johnst. Joum. Am. Arb.
241240. 1948. Type: Arum, Nevada, Jones
6692.
Krynitzkia depressa Jones, Contr. West. Bot.

13:5. 1910. not C. depressa A. Nels.
Oreocarya depressa Macbr. Contr. Gray

Herb. 48:, 32. 1916.
Cryptantha rnodesta Payson, Ann. Mo. Bot.

Card. 14:278. 1927. Not C. rnodesta

Brand.

Plants perennial, arising from a strong, woody
taproot, 0.5-1.8 dm high; stems many, 0.2-1.5
dm long, strigose and weakly setose; leaves ob-
lanceolate to spatulate, obtuse, strigose, setose

BnicnAM Young IKivehsitv Science Bulletin

UTAH

Cn/jitaullui udtsoiii

and subtomentose, the petioles ciliate margined;
inflorescence narrow, 0.2-0.8 dm long; calyx seg-
ments lanceolate to ovate, 2.5-4 mm long in
anthesis, in l^ruit becoming 5-8 mm long, setose;
corolla white, the tube 3-4 mm long, crests at
base of tube conspicuous, fomices yellow, round-
ed, papillose, about 0.5 mm long, limb 7-8 mm
wide; style exceeding mature fruit 0.5-1 mm;
nutlets ov;ite, 3-3.5 mm long, 2-2.5 mm wide,
usually all 4 maturing, margins in contact, ob-
tuse to acute, dorsal surface carinate, tubercu-
late, muricate and sometimes with low inconspic-
uous ridges, ventral surface deeply and irregu-
larly rugose, scar open, triangular, surrounded
by a slightly elevated margin. Growing on sandy
or gra\elly soil usually in the Transition Zone.
South central Utah, northwestern Arizona, and
eastern Nevada. April-July. Fig. .34.

Garfield Co.. Brvce Canyon, K.E. Weight B-32
(UT); 25 miles north'of Panguitch. B.F. Harrison 9009
(BRY); about 800 yards northwest of Sunset Point,
W.S. Bovie 1117 (Bin', UTC); 17 miles southwest of
Escalante, A. Crontjuist 91.52 (UTC); 2.4 miles east
of Red Canyon Campground, D. Atwood 1888 (BRY);
Iron Co., 1() miles northeast of Pinto, P. Plummer 7410
(UT); Piute Co., Marysvale. M.K. Jones 6679 (POM);
Sevier Co., about 28..3 miles north of Fremont, Welsh,
Ativood & Higgins 897.3 (BRY); Wayne Co., 10 miles
south of Fish Lake, L.C, Higgins 101.5 (BRY); about 5
miles south of Torrev, 1..C, Higgins 1016 (BRY); 5
miles north of Fremont. B,F. Harrison 7.364 (BRY);
about 8 miles west of Loa, L.C, Higgins 1.346 (BRY).

25. Cnjptantha bakeri Payson, Ann. Mo. Bot.
Card. 14:331. 1927. Type: Mancos River
sage plains, southern Colorado, Baker, Earle
& Tracy 827.
Orcocanja bakeri Greene, Pittonia 4:92.

1899.
Oreocarya eulopJm.s Rvdb. Bull. Torrey
Club 31:637. 1904.

Biennial or short-lived perennials, 1-4 dm
tall; stems 1-4 (6), 0.5-1.5 dm long, spreading
setose-hirsute; leaves oblanceolate, obtuse, most-
ly basal, 3-6 (8) cm long, 0.5-1.2 (2) cm wide,
dorsal surface strigose and spreading setose,
pustulate, ventral surface unifonnlv strigose and
with few or no pustulate hairs; inflorescence
narrow, 0.6-2.5 (3) dm long, setose-hirsute, fob'-
ar bracts evident, slightly surpassing the indi-
vidual cymes; calyx segments broadly lanceolate
or ovoid, in anthesis 3.5-4 mm long, in fruit be-
coming 6-8 mm long, conspicuously setose; corol-
la white, the tube 4-6 mm long, crests at base of
tube lacking, fomices vellow, emarginate, 1-1.5
mm long, limb 6-8 mm broad; nutlets ovate-
lanceolate, 2.5-3 mm long, 1.5-2 mm wide, 3 to
4 usually maturing, margins obtuse, nearly in
contact, dorsal surface deeply and sharply ru-
gose, scar closed, surroimded by a definitely ele-
vated margin; style exceeding mature fruit 1-2
mm. Growing on sandy or clay soil in the Pin-

UTAH

County Lines and
river drainages
u I -I

Kig. 34. Cnfpianthd <ihit(i

BioL(x;icAL Sekiks, Vol. 16, No. 3 The Bokacinaceae of Utah

29

yon-Juniper community. Southea.stem Utah,
.southwestern Colorado, northwestern New Mex-
ico, and northeastern Arizona. May-August. Fig.
35.

1 I

County Lines and
river drainages

i a5 io

Cn/ptantha bakcii

Garfield Co., Bryce Canyon Natl. Park, B. Maguire
19101 (UTC); about 17 miles southwest of Escalante,
.■\. Cron(iuist 91.53 (UTC); Br>ce Canyon Natl. Park,
K. Weight s.n. (BRY); Kane Co., headwaters of the
Virgin River, 1.5 miles north of Orderville, B. Maguire
188.55 (UTC); San Juan Co., La Sal Ranger Station, B.
Maguire 2059 (UTC); 5 miles west of Blanding, B.
Maguire 5102 (UTC); 8 miles north of Blanding, A.H.
Holmgren 3.374 (BRY. UTC); 18 miles southwest of
Blanding, A. Cronquist 9013 (UTC); 12 miles north of
junction 47-261, D. Atwood 1.5.39a (BRY); Natural
Bridges Natl. Monument, Welsh, Atwood & Higgins
8949 (BRY); about 15 miles south of junction of Utah
hwv 95-261, Welsh & Atwood 9979 (BRY); 10 miles
south of Monticello, L.C. Higgins 3558 (BRY); 8 miles
south of La Sal, L.C. Higgins 3.548 (BRY); about 16
miles west of Blanding, Welsh. Atwood & Higgins 8927
(BRY).

26. Cnjptantha harnehiji Johnst. Joum. Am.
Arb. 29:240. 194S. type: Uintah County, 30
miles south of Otirav. Riplev and Bameby

8748.

Perennial, 1.5-3.5 dm tall; stems stout, erect,
several, 0.8-1.2 dm long, conspicuously yellow-
hispid; leaves oblanceolate, thick, acute, 5-9 cm
long, 0.5-1.4 cm wide, coarsely appressed hispid-
pustulate on both sides, and with some finer

hairs beneath, the petioles conspicuously ciliate;
inflorescence narrow, 1-1.5 dm long, densely
vellowish-hispid, foliar bracts evident to con-
spicuous; calyx segments lanceolate, in anthesis
5-7 mm long, in fruit becoming 8-13 mm long,
yellowish-hirsute; corolla white or light yellow,
the tube 5-7 mm long, crests at base of tube
very conspicuous, fornices yellow, emarginate,
distinctly papillose, 0.5 mm long, limb 8-11 mm
wide; style exceeding mature fruit 5-6 mm; nut-
lets ovate, 3.5-4 mm long, 2.5 i.. mm wide, all 4
usually maturing, margins of nutlets in contact,
acute, smooth and glossy on both surfaces, scar
closed, straight, and without an elevated mar-
gin. Growing on white barren shale knolls. En-
demic to the lower part of the Uintah Basin in
Uintah County, Utah. May-June. Fig. 36.

Fig. 36. Cniptantlia barnclnji

Uintah Co., 8 miles west of Bitter Creek along the
Watson-Ouray road. L.C. Higgins 1887 (BRY); about
27 miles south of Ouray along the Hill Creek road, L.C.
Higgins 1877 (BRY);' along the Watson-Ouray road
at Buck Canvon, L.C. Higgins 1884 (BRY); 8 miles
west of Bitter Creek, L.C. Higgins 1.587 (BRY); about
6.5 miles west of Bitter Creek, L.C. Higgins 1586
(BRY); ;d)out 2 miles south of the Knools, L.C. Higgins
1.584 (BRY).

27. CnifiUintha breviflora (Osterh. ) Payson,
Ann. Mo. Bot. Card. 14:318. 1927. Type:
6.5 miles north of Jensen, Uintah County,
Utah, G. E. Osterhout 6414.

30

Bhicham Vciunc; Univebsitv Science Bulletin

Oreocarija breviflora Osterh. Univ. Wyom.
Publ. Bot. 1:169. 1926.

Long-lived perennials, 1.6-3 dm tall; stems
several, slender, 0.7-1.7 dm long, densely wliite
setose at the base, strigose above; leaves ob-
lanceolate to .spatulate, 2.5-9 cm long, 0.4-1.4
cm wide, clustered at the ends of the branched
caudices the apices obtuse, dorsal surface
densely and uniformly silky-strigose with many
very small pustules, ventral surface similar but
with fewer pustules; inflorescence in flower nar-
row but becoming broad and open at maturity,
0.6-2.7 dm long, setose; calyx segments linear-
lanceolate, 4.5-6 mm long in anthesis, in fruit be-
coming 7-9 mm long, setose; corolla white, .3.5-4.5
mm long, crests at base of tube evident, fomices
vellow, rounded, about 0.5 mm long, limb 8-12
mm wide; nutlets lanceolate, 3.4-4 mm long,
2-2.5 mm wide, less than 4 nutlets maturing,
margins in contact, knifelike, dorsal surface uni-
fonnly muricate or tuberculate, ventral surface
similar, scar open, narrowly triangular, margin
not elevated; style exceeding mature fruit by
2 mm or less. Growing on heavy clay soils.
Northeastern Utah in Duchesne and Uintah
counties. May-July. Fig. 37.

Duchesne Co., Red Creek, 3 mile.s ea.st of Fruit-
hind. L.C. Higgins 1044 (BRV); Bluebell, L.C. Hig-
gins 1060 (BRY); 5 miles east of Red Creek, S.L.
Welsh 1774 (BRY); about 2 miles east of Fniitland,

UTAH

County Lines and
river drainages

JL 1 1

Kig. 37. Cri/pUintlut hrcvifloni

Higgins & Welsh 1018 (BRY); Uintah Co., 2 miles
north of Bnish Creek sheep pens, D. Atwood 1611
(BRY); 3.5 miles north of Brush Creek sheep pens,
Higgins & Atwood 1868 (BRY); 8 miles south of hwy
40 on the road to Bonanza, L.C. Higgins 1084 (BRY);
Dinosaur Natl. Monument, S.L. Welsh 193 (BRY);
Red Wash Oil Field, S.L. Welsh .3901 (BRY); Dino-
saur Natl. Monument, J. Brotherson 806 (BRY); Dino-
saur Natl. Monument, S.L. Welsh 119, 148 (BRY);
Split Mountain Corge, J. Brotherson 993 (BRY); 14
miles west of Vernal. R.C. Rollins 17.36 (UTC).

28. CrifpiantJui caespitosa (A. Nels.) Payson,
Ann. Mo. Bot. Card. 14:281. 1927. Type:
Point of Rocks, Sweetwater County, Wyo-
ming, Nelson 4749.

Oreocanja caespitosa A. Nels. Erythea 7:65.
1899.

Densely caespitose or mat forming peren-
nials, 0.5-1.5 dm tall; stems 1-many, arising from
a much-branched woody caudex, 0.2-0.9 dm
long, weakly setose and appressed strigose;
leaves oblanceolate to spatulate, 1-3 cm long,
0.. 3-0.7 cm wide, pubescence of two kinds, stri-
gose and appressed setose, becoming tomentu-
lose toward the petiole; inflorescence narrow,
0.3-1 dm long, foliar bracts inconspicuous; calyx
segments lanceolate, in anthesis 3-4 mm long,
in fruit becoming 5-8 mm long, strigose and
weakly setose, also somewhat tomentulose; cor-
olla white, the tube 3-4 mm long, crests at base
of tube conspicuous, fomices yellow, rounded,
about 0.5 mm long, limb 4-7 mm wide; nutlets
lanceolate, 3-3.5 mm long, 2-2.5 mm wide, the
margins acute, in contact, dorsal surface with
low rounded rugae, also tuberculate, and with
numerous murications between the ridges, ven-
tnil surface muricate, scar open, narrowly tri-
;mgular, margin of scar not elevated; style
equalling or 0.5 mm longer than mature fruit.
Crowing on heavy clav soils. Southern Wyo-
ming and probablv northeastern Utah. May-
July. Fig. .38.

Crijptantha caespitosa occurs within a few
miles of the Utah border near Mckinnon and
Lonetree. With additional collections, this spe-
cies will without (juestion be found in Utah in
either Summit or Daggett counties or maybe
both.

29. C.njptantha capiiata (Eastw. ) Johnst. Joum.
Am. Arb. 21:66. 1940. Type: Hermit Trail
on the south rim of the Grand Canyon, A.
Ivistwood 5969.

Oreocanja capitata Eastw. Leaflets West.
Bot 1:9. 1937.

Perennial, 1.5-2.7 dm tall; stems weak, 1-sev-
eral. 1.2-2.4 dm long, appressed setose; leaves

UioLOCK Ai, StiuEs, \'oL. 16, No. 3 The Bouaginaceae of Utah

31

UTAH

County Lines and

river drainages

Cn/ptunthu caespitom

linear or very narowly oblanceolate, 3-8 cm long,
0.3-0.5 (0.8) cm wide, dorsal .surface appressed
setose-pustulate, ventral surface uniformly stri-
gose and without pustules; inflorescence capi-
tate, or with 1 or 2 glomerules below the termi-
nal cluster, 0.1-0.4 dm long, spreading white se-
tose; calyx segments linear-lanceolate, 7-9 mm
long in anthesis, in fruit becoming 11-16 mm
long, conspicuously setose-pustulate; corolla
white, the tube 9-12 mm long, crests at base of
tube conspicuous, fomices yellow, emarginate,
about 1 mm long, papillose, limb 6-8 mm wide;
nutlets lanceolate, 4-5 mm long, 2-3 mm wide.
2-4 usually maturing, the margins in contact,
knifelike, both surfaces glossy-smooth, scar
closed, straight, and without an elevated margin;
style exceeding mature fruit 4-5 mm. Growing in
sandv to sandy-loam soil in the transition zone.
South central Utah and north central Arizona.
.\pril-July. Fig. .39.

Garfield Co., Bull Creek, a little .southwest of Bull
Mountain. Henrv Mountains, 20 miles south of Hanks-
ville. 8,000 feet' Cronciuist & HolmRren 929f) (UTC);
\Va\ne Co.. with scattered Ponderosa Pine on the north-
east slope of A(|uarius Plateau southeast of Torrev,
Cronquist & Holmgren 9,36.5 (UTC).

30. ('niptantha compacta Higgins, Great Basin
Naturalist 28:196. 1968. Type: 8 miles we.st
of the Desert Range Experiment Station

Headquarters along hwy 21, L.C. Higgins
1613.

Densely caespitose perennials, 0.3-1 dm tall;
stems numerous, arising from a woody root,
0.1-0.4 dm long, tomentose below, weakly stri-
gose above, leaves oblanceolate to spatulate, ob-
tuse, 0.5-1.5 (2) cm long, 0.2-0.4 cm wide, dor-
sal surface with appressed setose-pustulate bris-
tles, also densely strigose or subtomentose, ven-
tral surface similar but with fewer pustulate
hairs, the petioles tomentose; inflorescence nar-
row, nearly capitate, 1-5 cm long; foliar bracts
evident but not conspicuous; calyx segments
lanceolate, 2-2.5 mm long in anthesis, in fruit
l>ecoming 3.5-4.5 mm long, densely white setose
and tomentose; corolla white, the tube 1.8-2.2
mm long, crests at base of tube evident, fom-
ices yellow, rounded, papillose, about 0.5 mm
long, limb 4.5-5.5 mm wide; nutlets lance-ovate,
acute, 2.5-3 mm long, 1.5-1.8 mm wide, only 1

UTAH

County Lines and
river drainages

i-

•'ig. .39. Cn/plaiithu capil.iUi

or 2 maturing, dorsal surface muricate or weakly
tuberculate-nigulose. ventral surface muricate,
scar open, subulate to narrowly triangular, ele-
vated margin lacking; style equalling or shorter
than mature fruit. Open slopes and ridges, grow-
ing on gravelly loam soil. Known only from
southwestern Millard Gounty, Utah. May-July.
Fig. 40.

32

Bric.ham Vounc Univehsitv Science Bulletin

UTAH

County Lines and
river drainages

i fc i.

Cn/ptaiitha i ompacia

Millard Co., about 8 miles west of the Desert Range
Experiment Station Headtiuarters along hwy 21, L.C.
Higgins 1462, 1613 (BRY); north slope of Pine Valley,
H.C. Holmgren 521 (BRY).

about 1 mm long, crest.s at base of tube evident
or sometimes lacking, limb 8-10 mm wide; nut-
lets triangular or ovate, 3.5-4 mm long, 2.5-3 mm
wide, usually all 4 maturing, margins narrowly
winged, in contact, surfaces smooth and glossy,
scar straight, closed, and lacking an elevated
margin; plants distinctly heterostyled. Growing
in a wide variety of soil types. Western Utah,
northern Arizona, southern Nevada, and west to
southern and western California. April-July. Fig.
41.

Garfield Co., about 8 miles southeast of Escalante,
Holmgren, Reveal & LaFrance 20.36 (BRY); Brvce Can-
yon Natl. Park, B. Maguire 19100 (UTC); near Granite
Ranch, W.P. Cottam 5.543 (BRY); Tropic Canyon, H.
Buchanan 122 (UT); 5 miles south of Boulder, L.C.
Higgins 1013 (BRY); |uab Co., between Callao and
Trout Creek, Maguire & Holmgren 21857 (UTC); 15
miles east of Trout Creek, B. Maguire 2753 (UTC); 18
miles east of Trout Creek, B. Maguire 27.54 (UTC);
Kane Co., Kanab Dunes, J.R. Murdock 101 (BRY);
Kanab, Anderson 16 (UTC); Cottonwood Canyon, 20
miles south of Cannonville, L.C. Higgins 1007 (BRY);
6 miles west from head of Collets Wash, D. Atwood 1873
(BRY); Milhird Co., north of Paddocks, P. Plummer 131
(UT); Desert Range Experiment Station, M. G. Bar-
low 84 (BRY); White Sage Valley, B.F. Harrison 6365
(BRY); Sanpete Co., Manti Canyon, 1 mile east of
Manti, A.H. Bamum 1291 (DIX);' 1 mile northwest of
Mavfield, R.S. Bjerregaard 39 (BRY); Sevier Co., west
of Richfield, B.F. Harrison 260 (BRY); Washington Co.,
entrance to Zion Natl. Park, B. Maguire 13557 (UTC);
0.5 miles north of Andersons Ranch, B. Maguire 16289
(UTC); Zion Natl. Park, trail to Angels Landing, B.

31. (-n/pt(intlui confertiflora (Greene) Payson,
Ann. Mo. Bot. Gard. 14:2.56. 1927. Type:
Cushenberry Springs on the north side of
the San Bernardino Mountains, S.B. Parish
1316.
Oreocarya confertiflora Greene, Pittonia

3:112. 1896.
Oreocartja lutea Greene, Muhlenbergia

2:240. 1906.
Oreocartja alata (Jones) Nels. Coult. & Nels.

Man. Cent. Rocky Mts. 417. 1909.

Perennial herbs, 1.7-4.3 dm tall; stems 1-7,
slender, 1.5-2.5 dm long, tomentose at the base,
strigose and setose upward; leaves linear to ob-
lanceolate, 3-12 cm long, 0.2-1.6 cm wide, acute,
dorsal surface densely strigose and appressed
setose with pustulate bases, ventral surface uni-
fonnlv strigose, and with few or no pustules;
inflorescence subcapitate, 0.3-2 dm long, strigose
and with twisted setose hairs, bracts inconspicoi-
ous; calvx segments linear-lanceolate, in anthesis
6-8 mm long, in fruit becoming 10-14 mm long,
strigose and spreading setose; corolla yellow, the
tube 9-13 nnn long, fornices broad, emarginate,

County Lines and
river drainages

Ir-

Cnjptanthu cuiifcrtifloru

Bioi.ocicAi. Sehies, Vol. 16, \o. 3

The Boracinaceae of Utah

33

Manure 16287 (UTC); Beaverdam Mountains, B. Ma-
miirc 20511 (UTC); 5 miles west of Virgin, C.L. Hitch-
cock 3015 (UTC); Pine Valley, M. Cox s.n. (DIX);
hills north of St. George, V. Worthen 42 (BRY); Santa
Clara, W.P. Cottam 1163 (BRY); between Hurricane
and Kanal). |.W. Harrison s.n. (DIX); Diamond Valley,
F.VV. Could' 1550 (BRY, UT, UTC); Diamond Valley.
L.C. Higgins 42(X> (BRY, WTSU); Zion Natl. Park, B.
[ohnson s.n. (DIX); 7 miles north of St. (Jeorge, E.
H.iwkins s.n. (DIX); Zion Natl. Park, A.M. Woodbury
s.n. (DIX); Zion Natl. P;u-k, W.P. Cottam 5149 (UT);
Diamond Valley, W.P. Cottam 4038 (BRY); Zion Natl.
P;irk, W.P. Cottam 4754 (BRY); 5 miles south of Veyo.
B.F. Harrison 10207 (BRY); 9 miles east of Hurricane,
L.C. Higgins 4224 (BRY. WTSU).

32. Cnjptantha ehita (Eastw. ) Payson, Ann.
Mo. Bot. Card. 14:285. 1927. Type: on the
road to the coal mines. Mesa County, Colo-
rado, Eastwood.

Oreocanja elata Eastw. Bull. Torrey Club
30:241. 1903.

Biennial or short-lived perennials, 3-5 dm
tall; stems 1-6, erect, stout, weakly setose with
spreading white hairs, 0.9-1.5 dm long; leaves
oblanceolate to spatulate, 2-5 cm long, 0.4-1.3
cm wide, apices acute to obtuse, the blade
abruptly tapering to the narrow petiole, dorsal
surface strigose and appressed setose, ventral
surface strigose, lx>th surfaces pustulate; inflo-
rescx'nce spreading or open in age, 1.5-3.5 dm
long, setose, foliar bracts inconspicuous; calyx
segments lanceolate, in anthesis 3-4.5 mm long,
in fruit Incoming 7-8 mm long, hirsute; corolla
white, the tube 3.5-5 mm long, fomices yellow,
rounded, papillose, about 1 mm long, crests at
base of tube well developed, limb 6-8 mm wide;
nutlets lanceolate-ovate, 4-4.5 mm long, 2-2.5
mm wide, usually all 4 maturing, margins in
contract, doreal surface densely tuberculate and
somewhat rugulose, the surface also covered with
dense minute papillae, ventral surface similar
but the roughenings less prominent, scar closed,
or narrowly open at the base, and without an
elevated margin; style exceeding mature fruit
0.5-2 mm. Growing on heavy clay soils. Western
Colorado and eastern Utah in Grand County.
May-June. Fig. 42.

Grand Co., about 3 miles south of U.S. hwy .50-6
along hwv 128, S.L. Welsh 69.52 (BRY); near milepost
39 along hwv 128, L.C. Higgins 1479 (BRY).

33. Crtiptantha flava (A. Nels. ) Payson, Ann.
Mo. Bot. Card. 14:259. 1927. Type: Point of
Hocks, Sweetwater County, Wyoming, Nel-
son .•3074.

Oreocanja flava Nelson, Bull. Torrey Club
2.5:202. 1898.

Oreocanja lutescens Greene, Pittonia 4:93.

1899.
Cnjptantha confertiflora var. flava Brand,

Pflanzenreich, (Heft 97) 4, Fam.

252:90. 1931.
Cnj}>tantha confertiflora var. lutescens

Brand, Pflanzenreich, (Heft 97) 4, Fam.

252:90. 1931.

Perennial, 1.3-4 dm tall; stems many from a
multiple caudex, 0.8-2.6 dm long, densely setose,
white hairy at the base, becoming setose and
strigose upward; leaves narrowly oblanceolate to
nearly linear, acute, 2-9 cm long, 0.3-0.8 cm
wide, dorsal surface strigose and appressed se-
tose with pustulate hairs, ventral surface almost
uniformly strigose, and with the pustulates less
conspicuous; inflorescence narrow to somewhat
open, 0.5-2.5 dm long, conspicuously yellow se-
tose, the floral bracts inconspicuous; calyx seg-
ments linear, in anthesis 8-10 mm long, in fruit
becoming 9-12 mm long, densely setose, with
vellowish hairs, pedicels 3-5 mm long in fruit;
corolla yellow, the tube 9-12 mm long, crests
at the base of the tube absent or nearly so,
fornices yellow, truncate, emarginate, 1-1.5 mm
long, limb 8-10 mm wide; nutlets lanceolate,
3.4-4 mm long, 1.9-2.2 mm wide, 1 or 2 usually
maturing, margins acute, in contact when more
than 1 nutlet matures, both surfaces of nutlet

UTAH

County Lines and
river drainages

o 80

Kig. 42. Crifptantha data

34

BninnANf YouNC I'nmversitv Scifncf Bulletin

smooth and glossy, scar straight, closed, elevated
margin lacking; style exceeding mature fruit 3-7
mm (heterostyled). Usually growing in sandy
soils. Southern Wyoming, south through western
C>olorado and eastern Utah to northern New
Mexico and Arizona. April-August. Fig. 43.

UTAH

County Lines and
river drainages

A JO io

C.rijptiintlui jln i

Carbon Co., 2 miles east of roadside geyser, R.K.
V'ickery 721 (UT); Ducfiesne Co., 2 miles west of
Duehesne, A. Collotzi 54.5 (UTC); 8 miles north of
Duehesne, L.C. Higgins 10.57 (BRV); 1 mile west of
Duehesne, L.C. Higgins 1051 (BRY); 12 miles north
of Duehesne. J. Brotherson 925 (BRY); Emery Co..
alxiut 15 miles west of hwv 50-6 along the road to
Huntington, L.C. Higgins I.'J05 (BRY); about 5 miles
west of Temple Mountain, L.C. Higgins 1.32.5a (BRY);
50 miles south of Green River, A. Cronciuist 9086
(UTC); Goblin V;illev, S.L. Welsh 3925 (BRY); San
Raph.»el Suell. B. Maguire 18302 (UTC); 20 miles east
of Hank-sville, B. Maguire 18211 (UTC); along hwy 24,
10 miles south of San Raph;iel River bridge, LB. Bar-
nett 22 (BRY, UTC); Temple Mountain ' V. P. ,\llman
s.n. (BRY); San Raphael River. B.F. Harrison 8058
(BRY); 18 miles east of Wellington, L.C. Higgins 998
(BRY); C;reen River, W.P. Cottam s.n. (BRY); east of
Wellington, W.P. C:<)ttam 2050 (BRY); .50 miles south
of San Raphael River, B.F. Harrison 9627 (BRY); 20
miles e;ist of Price, Welsh & Moore 2791 (BRY); C^ar-
field Co., 25 miles southeast of Hanksville (;. Parrv
s.n. (UTC); 50 miles southwest of San Raphael B.F.
Harrison 7441 (BRY); ;il)Out 7 miles south of Trarhvtc
Ranch, S.L. Welsh 3940 (BRY); Sandv Ranch, B.F.
Harrison 11489 (BRY, UTC); Calf Creek on the Es-
calante River, McArthur & Beck 195 (BRY); 23 miles
up North Wash from Hite, B.F. Harrison 11.5.50 (BRY);

5 miles south of Boulder, G.E. Bohart .s.n. (UTC);
cast Henry Mountains, near Granite Ranch. B.F. Harri-
son 5544 (BRY); 14 miles south of the Kane County
line on Hole in Rock road, J.B. Karen 89 (BRY); Es-
calante near town. ME. Jones 4408 (BRY); Grand Co.,
Arches Natl. Monument, L.C. Anderson 48 (UTC);
De;ul Horse Point, A. Carter 1.560 (UTC); Arches Natl.
Monument, B.F. Harrison 11129 (BRY); Arches Natl.
Monument. L.B. Bamett 44, .57 (BRY. UTC); Dead
Horse Point, B.F. Harrison 10293 (BRY); Kane Co.,
Willow Tank, about 17 miles south of Garfield County
line, D.H. White 95 (BRY); Glen Canyon area, F.
Wright s.n. (ARIZ); about 3 miles north of U.S. hwv 89
in Buckskin Gulch. S.L. Welsh 5319 (BRY); Cockscomb
Ridge, about 40 miles east of Kanab, S.L. Welsh 5342
(BRY); Plateau near head of Rock Creek, B.F. Harrison
9033 (BRY); Escalante Desert near Willow Tank, B.F.
Harrison 9033 (BRY); San Juan Co., White Canvon. L.
O. Wilson 51 ( UTC ) ; 5 miles northwest of Mexican Hat,
B. Maguire 16290 (UTC); 16 miles east of Hite, A.
Cron<|uist 9023 (UTC); Hatch Rock, S. Flowers 2071
(UT); Rainbow Bridge Trail. J.T. Howell 24617
(ARIZ); ;ibout 10 miles east of H;ills Crossing. L.C.
Higgins .527 (BRY); Natural Bridges Natl. Monument,
G. Moore 140 (BRY); Island in the Sky area, G. Moore
190 (BRY); La S;il Range. W.P. Cottam 2197 (BRY);
4 miles above Cow Canyon. L.K. Shnmwav 64 (BRY);

6 miles south of Bluff. D.E. Bright 114 (BRY); 2 miles
northwest of Bluff. B. Maguire 13.531 (UTC); 18 miles
southwest of Blanding, A. Cronquist 89.56 (UTC); 7
miles northeast of Mexican Hat, A. Cronquist 9133
(UTC); Cow Canvon, AH. Holmgren .3212 (UTC);
Robbers Roost, L.A'. Stoddart s.n. (UTC); Rock House,
Monument Valley. A.H. Holmgren .3220 (UTC); 19
miles south of Blanding, B. Maguire 5104 (UTC); 14
miles south of Bluff, b'. Maguire 5103 (UTC): Natural
Bridges Natl. Monument, B. Maguire 20.58 (UTC);
S(|naw Canyon, 1 mile west of Cane Spring. Welsh,
Moore & Canter 2864 (BRY); about 5 miles north of
Bluff S.L. Welsh 1493 (BRY); about 5.5 miles north
of Bluff, C.A. Hanson 81 (BRY); west rim of Browns
Canvon, L.K. Shumwav 1.33 (BRY); ledges north of
San Juan River just east of Johns Canvon. B F. Harrison
119.54 (BRY. UTC); 10 miles west of Bluff, B.F. Harri-
son 10.363 (BRY); Uintah Co.. 14 miles west of Venial,
RC. Rollins 1734 (UTC): Dinosaur Natl. Monument,
T. Brotherson 1100 (BRY); desert west of Vem;il.
L.O. Williams 619 (I'TC): about 5 miles west of Ver-
n:il L.C Higgins 1069 (BRY): 7 miles south of Fort
Duchesne. J. Brotherson 1105 (BRY): .ibout 8 miles
south of hw\' 40 :ilong the road to Bonanza. L.C. Hig-
gins 1082 (BRY); about 3 miles northwest of Vernal,
LC. Hiuiiins 1073 (BRY): Dinosaur Natl. Monument,
S.L. Welsh 176 (BRY); W.ivne Co. Fnnt;i Arch Can-
von. B. Maguire 181.34 (UTC): 17 miles south of
ILuik'.ville. B Macuire 18190 (UTC); 13 miles west of
Hinksville B Maguire 18149 (ITTC); Fniita. DE. Beck
sn (BRY).

34. Cniptnnfhd flnvnrulnfn (A. Nels.) Pavson.
Ann. Mo. Rot. Card. 14:.a34. 1927. Tvpe:
Piedmont. Wyoming, A. Nelson 4572.
Oreocania flavoctilata A. Nels. Ervthea 7:67.

1899.
Orroraryo fhivnruliitd spattilafii A. Nels.

Erv+he;i 7:f}7. 1899.
Oreoamja rr'nsrtata Eastw. Bull. Torrev Club

.30:244. 190.3.

BioLooicAi. Serifs. Vol. 16, No. 3 The Bon acinaceaf. of Utah

35

Oreocarija .fhockletji Eastw. Bull. Torrey

Club 30:245. 1903.
Oreocarya eastwoodae Nels. & Kenn. Muh-

lenbergia 3:141. 1908.

Caespitosc perennial, 1-3.7 dm tall; stems
I-several, slender, 0.5-2 dm long, strigose and
spreading setose with slender bristles; leaves
linear-oblanceolate to spatulate, obtuse or some-
times acute, 3-11 em long, 0.3-1.5 cm wide, dense-
ly strigose and weakly setose, dorsal surface con-
spicuously pustulate, ventral surface with few
pustules or sometimes silky-strigose; inflores-
cence narrow or sometimes slightly open and
lax, 0.5-3 dm long, foliar bracts evident but not
conspicuous; calyx segments in anthesis linear-
lanceolate, 5-6 mm long, in fruit becoming 8-10
mm long and becoming broadly lanceolate to
ovate; corolla white or pale-vellow, the tube 7-10
mm long, crests at base of tube lacking, fomices
yellow, minutely papillose, 1-2 mm long, limb
S-12 mm wide; nutlets lanceolate to lance-ovate,
2.5-3.5 mm long, 1.8-2 mm wide, usually all 4
maturing, margins obtuse, in contact or slightly
separated, dorsal surface muricate, tuberculate,
and with conspicuous ridges, sometimes nearly
foveolate, ventral surface tuberculate, rarely with
ridges, scar open, constricted near the middle
and surrounded by a highly elevated margin;
style exceeding mature fruit 4-8 mm (hetero-
styled). Open slopes and ridges mostly in the
Pinvon-Juniper community. Growing on a wide
\ariety of soils. Southern Wyoming, western
Colorado, Utah, Nevada, and southeastern Cali-
fornia, south to Arizona and New Mexico. April-
July. Fig. 44.

Beaver Co.. about 5 mile.s north of Cove Fort, L.C.
Higgin.s 903 (BRY); Wall Wah Mountains. 4.5 mile.s
up' Pine Canvon, B. Maguire 20941 (UTC); Wah Wah
Mountains, Pine Grove, W.P. Cottam 8031 (UT); Car-
bon Co., 5 miles east of Wellington, L.C. Higgins 997
(BRY); near Carbon-Emery countv line along hwv
.50-6, L.C. Higgins 3.324 (BRY); Daggett Co., 2 miles
east of Sheep Creek Junction with hwv 44, D. Atwood
1.586 (BRY); 15 miles south of Manila. R.C. Rollins
2280 (UTC); Carter Creek. E.E. Jensen s.n. (UTC);
Duchesne Co., Indian Canyon, 3.5 miles south of
Duchesne. Holmgren, Reveal' & Lafrance 1760 (BRY);
27 miles soutli of Mvton, A. Cronquist & Holmgren
92.38 (UTC); about 6 miles south of Tabiona, L.C.
Higgins 1046 (BRY); about 20 miles south of Myton,
L.C. Higgins 1065 (BRY); about 2 miles east of Fniit-
land, Higgins & Welsh 1020 (BRY); north rim of Nine
Mile Creek. Holmgren. Reveal & LaFrance 19.50 (BRY,
UTC); ;ibout 5 miles west of Fniitland, L.C. Higgins
1086 (BRY); about 6 miles west of Duchesne, L.C. Hig-
gins 1049 (BRY); ;ibout 2 miles e;ist of Fruitland, L.C.
Higgins 1019 (BRY); Red Creek 5 miles north of
Fniitland, J. Brotherson 1128 (BRY); Rock Creek, J.
Brotherson ' 965 (BRY); Bluebell, L.C. Higgins 1061
(BRY); Red Creek 3 miles east of Fniitland, L.C. Hig-
gins 1045 (BRY); Emerv Co., 10 miles up Hiintingtoii

Canvon, B.F. Harrison 8167 (BRY); :ibout 9 miles south
of L;iwrence, L..\I. Pitts 2.3 (BRY); Grand Co., Arches
N;itl. Monument, L.B. Bamett 41 (BRY); Iron Co.,
Kanarraville, W.P. Cottam .5009 (UT); Juab Co., .south
base of Topaz Mountain, near Thomas Pass, Welsh &
Atwood 9667 (BRY); foothills east of Mona, D. Hatch
109 (UTC); Deep Creek Mountains, moutli of Thomas
Creek, D.W. Lindsav 206 (UT); Millard Co., Fillmore,
D.M. Leidig 237 (UTC); Hou.se Range, 44 miles west
of Delta, \iaguire & Becraft 27.55 (UTC); Warm Point,
Desert Range Experiment Station, V.B. Matthews I
(BRY); Wami Point Ridge. F. Coles 6 (BRY); Desert
Range Experiment Station, Tunnel Springs, W.P. Cot-
tam' 8.523 (UT); Pruess Lake, V.B. Matthews 25
(BRY); Rich Co., St;inley Cessell s.n. (UTC); San
|uan Co., 10 miles north of Blanding, B. M;iguire 13505
(UTC); Natural Bridges Natl. Monument, L.C. Higgins
522 (BRY); head of Gravel Canyon, L.O. Wilson 235
(UTC); 13 miles south of Moab. A. Cronquist 9001
(UTC); 8 miles north of Monticello, B.F. Harrison 5891
(BRY); about 5 miles .southwest of Blanding, L.C. Hig-
gins 517 (BRY); Sanpete Co., 0.5 miles north of upper
power plant in Ephraim Canyon, T. Jensen s.n. (DIX);
Ephraim Canyon, K. Peterson s.n. (DIX); 5 miles east
of Ephraim, AH. Bamum 1259 (DIX); benveen Eph-
raim and Great Basin Experiment Station, A.H. Holm-
gren 76.55 (UTC); ;ibout 8 miles southeast of Mt. Pleas-
ant, L.C. Higgins 112 (BRY); Sevier Co., Siilina Can-
von, R. Stevens 33 (BRY); 40 miles north of Fremont,
B.F. Harrison 7341 (BRY); 5 miles southeast of Sigurd,
B. Maguire 18098 (UTC); Summit Co., 1 mile west of
Upton' along hwy 133, Welsh & Murdock 6264 (BRY);
Tooele Co., Deep Creek Mountains, Johnson Canyon 10
miles southeast of Ibapah, W.P. Cottam 7218 (UT);
Stansburv Range, up Willow Creek, B. Maguire 21824
(UTC); Uintah Co., 5 miles northwest of Whiterocks,
I. Brotherson 1027 (BRY); behveen Two Waters Creek

County Lines and

river drainages

I I 1

6 55 fco

I'ig. 44. Crt/ptantha fluvoculnUi

36

HnK.llAM YoUNC l'Nl\tHSITV SciliNCK BULLETIN

and Willow Creek, Holmgren & Reveal 1857 ( BHY,
UTC); Seep Canyon. Holmgren & Reveal 1862 (UTC);
about 8 miles south of hwv 40 along the road to Bonan-
za, L.C. Higgins 1083 (BRY); Dinosaur Natl. Monu-
ment. S.L. Welsh 62, 92, 155, 272 (BRY); Merkeley
Park, Welsh & .Moore 1955 (BRY); about 3 miles north-
west ol \'ernal. L.C. Higgins 1072 (BRV); Big Spring,
V.A, Murphv 19 (UTC); Utah Co,, about 1 mile north
of Divide along hwy 50-6, W,R. Buss 61 (BRY); just
east of Eureka, J.B, Karen 71a (BRY); Soldier Summit,
Holmgren & Boyle 7057 (UTC); W;ishington Co,.
Beaverdam Mountains, near summit. L,C, Higgins 557
(BRY); summit of Beaverdam .Mountains near television
relay tower, L,C, Higgins .3403 (BRY); Wavne Co.,
Fruita Areh Canvon,' B, Maguire 18136 (UTC); 17
miles south of Hanksville, B. Maguire 18170 (UTC).

35, Cn/ptantha juJvocanescens (Wats.) Payson,
Ann. Mo. Bot. Card. 14:319. 1927.

Densely caespitose perennials from a strongly
lignified taproot, 0.8-3 dm tall; stems many from
a multiple caudex, 0.5-1,3 dm long, white hairy
at the base, setose-hispid upward; leaves ob-
lanceolate to spatulate, acute to obtuse, 1.5-7 cm
long, 0.4-1.2 cm wide, uniformly strigose, pus-
tules mainly confined to the dorsal surface; in-
florescence narrow or somewhat open at matur-
ity. 0.3-1.9 dm long, vyhite or yellowish setose,
foliar bracts inconspicuous; calyx segments lin-
ear, 4-6 mm long in anthesis, in fruit becoming
9-13 mm long, densely white or yellowish setose,
pedicels 2-10 mm long; corolla white, the tube
7-11 mm long, crests at base of tube eyident or
lacking, fornices yellow, emarginate or rounded,
0.7-1.3 mm long, limb 7-9 mm broad, nutlets
lance-oyate, 3.5-4.5 mm long, 2-3 mm broad, 1
or 2 usually maturing, margins acute to obtuse,
in contact when more than 1 nutlet matures,
both surfaces densely and uniformly muricate,
scar open or nearly closed, elevated margin lack-
ing; style exceeding mature fruit 3-7 mm.

1. Muriciitions on the nutlet rounded; corolki 9-13
mm long; inflorescence narrow, white setose at
miituritv 3.5A, var, fulvocanescens

1, ,Murie;iti<)ns on the nutlet with 1 or 2 setose pro-
jections; corolhi 7-9 mm long; inflorescence bro;ider

and usually yellowish .setose at maturity

35B, var. crhinoidc.s

35A. Crijj)t(tntlui fulvocanescens (Wats.) Pay-
son, var. fuh'Ocaiwscetis Type: near Santa
Fe, New Mexico, Fendler 632.
Eritrichium glome mtum var. fulvocanes-
cens Wats. Bot. King Exp. 243. 1871.
Eritrictiitivi fulvocanescens Cray, Proc,

Amer. Acad. 10:61. 1875,
Kn/nitzkia fulvocanescens Cray, Proc.

Amer. Acad. 20:280. 1885.
Oreocart/a fulvocanescens Creene, Pittonia
1:58. 1887.

Oreocanfa nltkla Creene,
1901.

PI. Baker. 3:21

Densely caespitose perennial, 1-3 dm tall;
inflorescence narrow, white setose; pedicels 2-3
mm long; corolla white, the tube 9-13 mm long,
crests at base of tube evident or lacking, fornices
yellow, rounded to acute; nutlets lanceolate, 3.5-
4 mm long, 2-2.5 mm wide, the dorsal surface
with rounded murications, scar straight, closed
or slightly open. Sandy to sandy clay-loam soils,
on open slopes and ridges. Western Colorado,
northwestern New Mexico, northeastern Arizona,
and southern Utah. April-August. Fig. 45.

County Lines and
river drainages

I'ig, 45. Ciiiptanllht juit ociinesceiks v.ir, jiiii)i>Ciiiicscivs

Kmery Co., Mouth of Huntington Canyon, B.F",
H.irrison 8136 (BRY); about 15 miles west of 'hwv 50-6

1307
L.C.
Har-
Har-
G.L.

ak)ng the road to Huntington, L.C, Higgins
(BRY); Grand Co., .ibout 7 miles south of Moab,
Higgins 999 (BRY); 5 miles south of Moab, B.F.
rison 10.306 (BRY); Arches Natl, Monument, B,F.
rison 11135 (BRY); ,Arches Natl, Monument,
Pvrah 48 (BRY); 16 miles northwest of Moab, A.
CroiKiuist 9064 (UTC); Arches Natl. Moiuiment, L.C.
Anderson 49 (UTC); S;m Juan Co,, 16 miles east of
Whit.' Canyon on hwv 95, D,K. Bright 75 (BRY);
Srpuiw C.mvon. 1 mile east of C;ive Spring. Welsh &
Moore 2849 (BRY); 30 miles south of Mexic;m Hat.
|.R, Murdock 308 (BRY); 2 mih-s west of Bluff. B.
Maguire 1.3.530 (UTC); 18 miles southwest of Bland-
ing! A. CnnKpiist 8949 (UTC); Monument V;illey, AH,
Holmgren 3225 (UTC): Wayne Co.. 28 miles southeast
of Hanksville, A, Crompiist 9181 (UTC); .32 miles east
of Fniil;!. B, \1;iguirc- 18156 (UTC).

Biological Series, Vol. 16, No. .3 The Roh.vcin.^ce.^e of Ut.\h

.37

35B. Cniptcmtlia fttlvocanescens (Wats.) Puyson
var. cchinoicles (Jones) Higgins, Great
Basin Naturalist 29:30. 1969. Type: Pahria
Canvon, Kane County, Utah, Jones 5297p.
KrynLtzkia ecliiiwicles Jones, Proc. Calif.

Acad. Sci. II. 5:709. 1895.
Orcocanja cchinoidea Macbr. Contr. Gray

Herb. 48:31. 1916.
Cnjptantha ecltinoides Payson, Ann. Mo.

Bot. Card. 14:321. 1927.

Caespitose perennials, 0.8-3.6 dm tall; inllo-
rescence narrow to somewhat open at maturity,
yellowish setose; pedicels 3-10 mm long; corolla
white, the tube 7-9 mm long, crests at base of
tube lacking, or sometimes evident, fornices yel-
low, emarginate; nutlets lance-ovoid, 4-4.7 mm
long, 2.5-3 mm broad, the dorsal surface with
1 or 2 setose projections temiinating each muri-
cation, scar asymetrical, and without an elevated
margin. Usually growing on heavy clay soils
charged with alkali or salts. South central Utah
and north central Arizona. April-July. Fig. 46.

UTAH

Kis;. 46. Cn/ptnntha fuli<)C(ine\cen.'! wir. ccliinoidc :

Kmcrv Co., near Cedar Mountain, about 39 miles
west ot Crccn liivcr along road to Castlf Dale, A. Cron-
(|uist 9096 (UTC); .50 miles south of San Raphael River,
li.F. Harrison 9626 (BRV): C:arfiekl Co., west side of
Henr>- Mountains, B.F. Harrison 11695 (BRY); Kane
Co., .5 miles from Mount Carmel, Kastwood & Howell
926,5 (UTC); Cottonwood Canyon. 20 miles south of
CaiinniiMlle. L.C. His^gins 1008 (BRY); 20 miles east

of Kanab, D. At\vood 1792 (BRY); 3.7 miles east of
Skutumpah-Alton Junction at head of Johnson Canyon,
D. Atwood 1802 (BRV); Washington Co., west en-
trance to Zion Natl. Park, H.A. Canning 2288 (ARIZ);
Wavne Co., 1 mile east of Bicknell, A. Cronquist 9171
(UTC); 9 miles east of Torrey, B. Maguire 18111
( UTC ) ; about 2 miles west of River Bridge, at junction
of Buckhoni Wash and San Raphael River, Welsh & At-
wood 9904 (BRY); about 5 miles east of Torrey, L.C.
Higgins 1344 (BRY).

36. Cnjptantha grahainii Johnst. Journ. Am.
Arb. 20:391. 1939. Type: Bench west of
Green River north of mouth of Sand Wash,
Uintah County, Utah, Graham 7924.

Long-lived perennial from a thick woody
taproot, 1.5-2.5 dm tall; stems several, 0.4-1.2 dm
long, weakly spreading setose; leaves spatulate to
oblanceolate, 2-4.5 cm long, 0.4-1 cm wide, con-
spicuously setose pustulate on both surfaces,
with some finer pubescence beneath; inflores-
cence narrow, 0.4-1 dm long setose, foliar bracts
evident but not conspicuous; calyx segments
lanceolate, in anthesis 5-7 mm long, in fruit be-
coming 7-9 mm long, very setose; corolla white,
the tube 3.5-5 mm long, constricted at the mid-
dle, crests at base of tube evident, fornices yel-
low, emarginate, papillose, 0.5-1 mm long, limb
11-15 mm wide; nutlets lanceolate, 3-3.8 mm
long, 1.7-2 mm wide, 2-4 maturing, margins in
contact, acute, both surfaces of nut with incon-
spicuous, small, low-rounded tubercles, or some
of these confluent into short irregular ridges,
scar straight, open, narrowly linear, the margin
not elevated; style coarse, exceeding mature fruit
1.8-2.2 mm. Found growing only on Green River
Shale. Endemic to the Uintah Basin, Uintah
County, Utah. May-June. Fig. 47.

Uintah Co., Sunday School Canyon of Willow
Creek drainage, Holmgren & Reveal 1879 (BRY, UTC);
Willow Creek, R.C. Rollins 1707 (UTC); 30 miles south
of Ouray along the Hill Creek Road, Higgins & Atwood
1878 (BRY); Buck Canyon along the Watson-Ouray
Road, Higgins & Atwood 1885 (BRY); along Willow
Creek 5 miles north of the mouth of Agency Draw, L.C.
Higgins 1608 (BRY); 35 miles south of Ouray along
the Hill Creek Road, Higgins & Atwood 1881 (BRY)';
about 18 miles south of Ouray along the Hill Creek
Road, Higgins & At\vood 1876 (BRY).

.37. Crt/ptantha hwniJis (Gray) Payson, Ann.
Mo. Bot. Card. 14:278. 1927.

Perennials, more or less densely caespitose,
0.5-3 dm tall; stems many, arising from the ends
of the branched caudex, 0.2-1.5 dm long, strigose
to spreading seto.se; leaves oblanceolate to spatu-
late, 1-6 cm long, 0.2-1.2 cm wide, strigose, se-
tose or subtomentose, pustulate on both surfaces;
inflorescence narrowly cylindrical to open and
lax, 0.2-1.8 dm long, tomentose to conspicuously

38

Bnir.HAM Young University Science Bulletin

UTAH

Fig. 47. Cii/j)lanlli(i •^luluniiii

setose; calyx segments linear-lanceolate, in anthe-
sis 2.5-4.5 mm long, in fruit becoming 6-13 mm
long, setose or tomentose; corolla white, the
tube 2.5-4.5 mm long, crests at base of tube
conspicuous to nearly obsolete, fomices yellow,
more or less papillose, rounded, about 0.5 mm
long, limb 7-10 mm wide; nutlets lanceolate to
ovate-lanceolate, 3-4.5 mm long, 1.8-3.2 mm
wide, 1 to 4 of them maturing, margins in con-
tact, acute to somewhat obtuse, dorsal surface
inuricate, tuberculate, or somewhat rugulose,
ventral surface indistinctly muricate or tuber-
culate, scar open, triangular, margin not ele-
vated; style from shorter than to longer than the
mature fruit by 2.5 mm.

1. Lfavcs strigosf and .setose but not conspicuousl)'
tomentose; calyx evidently setose (2).

1. Leaves densely strigose as well as tomentose; caly.x
setose and suhtomentosc ( 3 ) .

2. Style e.\ceeding the mature nutlets 1-1. .5 mm; the
inflorescence open and broad; plants loosely
tufted 37A. var. comriiixtu

2. Style not or only slightly surpassing the nutlets;
inflorescence congested, even in fniit; plants densely
caespito.se 378. var. naua

3. Style scarcely exceeding the mature nutlets; inflo-
rescence somewhat open at maturity

.37C. var. shwitzii

3. Style exceeding the mature fniit 0.5-1.5 mm; in-
florescence cylindrical and congested in fruit
37D. var. ovijui

37A. Cnjptanthii humilis (Gray) Payson, var.
commixta (Macbr. ) Higgins, Brigham
Young Univ. Sci. Bull. Vol. 13, No. 4:36.
1971 ' Type: Juab, Juab County, Utah
Goodding 1074.
Oreocarya commixta Macbr. Contr. Gray

Herb. 48:33. 1916.
Cniptantha nana var. commixta Payson,

Ann. Mo. Bot. Card. 14:312. 1927.

Caespitose perennials, 1-2.7 dm tall; stems
1-several, arising from the ends of the branched
caudex, 0.3-1 dm long, weakly strigose and
spreading setose; leaves spatulate to broadly ob-
lanceolate, 2.5-6 cm long, 0.5-1.2 cm wide, stri-
gose and spreading setose; inflorescence open,
0.8-1.8 dm long, foliar bracts evident on lower
part of stem; calyx segments linear-lanceolate,
in anthesis 3.5-4.5 mm long, in fruit becoming
7-10 mm long, seto.se; nutlets lance-ovate, 3.5-4
mm long, muricate, tuberculate, or sometimes
with the murications joined to form short irregu-
lar ridges, scar subulate or nearly closed; style
exceeding mature fruit 0.7-1.6 mm. Usually
growing on gravelly soil or talus slopes in the
Pinyon-Juniper community. Central Utah to
eastern Nevada. May-July. Fig. 48.

Beaver Co., Summit of Wah Wah Mountains along
bwv 21, L.C. Higgins 1617. 1468 ( BRY); Juab Co., Juab,
Coodding 1074 (RM); Millard Co.. Ca. 70 miles west

UTAH

I'ig. 48. (.'rt/jiliiulhii htnnili.\ \,ir. itmniiixUi

Biological Series, Vol. 16, No. .3 The BoR.\r.iNACEAE ok Utah

39

of Dflta, L.C. Higgiiis 1612 (BRV); I'avant Butte, 17
miles northwest ot Fillmore, Cottam & McMillan 9592
(UT); Piute Co., Marysvale, M.E. Jones .5388 (UT);
6 miles east of Kingston along liwv 62, L.C. Higgins
1618 (BRY); Sevier Co., near the Big Rock Candy
Mountain, L.C. Higgins 1620 (BRY); Tooele Co.,
Stansbury Lsland, G. Saccomanno 7680 (L'T).

37B. Cryptantha hwiiilk (Gray) Payson, var.
luina (Eastw. ) Higgins, Brigham Young
Univ. Sci. Bull. ^ Vol. 13,' No. 4:37.
1971. Type: near Grand Junction, Colora-
do, Ea.stwood s.n. 1S92.
Oreocari/a luina Eastw. Bull. Torrey Club

30:243. 1903.
Cryptantha nana (Eastw.) Payson, var.

tijpica, Ann. Mo. Bot. Card. 14:315.

1927.

Caespitose perennial, 0.5-1..5 dm tall; stems
several, 0.2-0.7 dm long, setose; leaves oblanceo-
late to spatulate, 0.5-4 cm long, 0.2-0.6 cm wide,
strigose to subtomentose, spreading setose; in-
florescence narrow, cylindriail, 0.2-1 dm long,
setose; calyx segments linear-lanceolate, in an-
thesis 2.5-.3.5 mm long, in fniit becoming 6-8
mm long, setose; corolla white, the tube 2.5-3.5
mm long, crests at base of tube evident; nutlets
muricate or sometimes tuberculate or rugulose;
stvle shorter to slightly longer than mature fruit.
Growing mostly in heavy clav soils but occasion-
ally on sandy-loam soil. Western Colorado and
eastern Utah. April-July. Fig. 49.

Duchesne Co., west of Duchesne, Ripley & Bameby
4675 (C.\S); north of Duchesne, G.E. Osterhout 6200
(RM); Summit Co., Echo Canyon, G.E. Osterhout 6231
(RM); San Juan Co., 13 miles south of Moab, A. Cron-
(|uist 9108 (NY); Uintah Co., 1 mile west of Red Pine,
Book Cliffs, L.C. Higgins 1.598 (BRY).

37C. Cryptantha humills (Gray) Payson, var.
shantzii (Tidest. ) Higgins, Brigham Young
University Sci. Bull. Vol. 13, No. 4. pp.
37 1971 Type: Grant's station south of the
Great Salt' Lake, Kearney & Shantz 3098.
Oreocarya sliantzii Tidestr. Proc. Biol. Soc.

Wash. 26:122. 1913.
Oreocarya dolosa Macbr. Contr. Gray Herb.

48:32. 1916.
Crijptantha naiui var. shantzii Payson,
Ann. Mo. Bot. Card. 14:313. 1927.

Caespitose perennial, 1-2 dm tall; stems 1-
manv from the ends of the much-branched cau-
dex, 0.3-1 dm long, we;ikly setose and strigose;
leaves spatulate to oblanceolate, obtuse, 2-5 cm
long, 0.3-0.7 cm wide, tomentose and appressed
setose, with slender more or less appressed
bristles; inflorescence usually narrow, but the

County Lines and
river drainages

i-

Kig. 49. Cniptantlta liiiiuili.s var. naiid

cvinules slightly elongating, lower foliar bracts
rather conspicuous in the young inflorescence;
calyx segments densely setose and subtomentose;
stvle scarcelv exceeding the nature, muricate
nutlets. Open, s;mdy to gravelly slopes and
ridges. Southwestern Montana, eastern Idaho,
and northern Utah. April-July. Fig. 50.

Bo.\ Elder Co.. Kelton Pa.s.s, Maguire & Holmgren
26247 (UTC); Raft River Mountains, S.J. Preece 889
Cache Co., Providence Canyon, B. Maguire
(UTC); College Bench, B. Maguire 3700
; College Bench, L.C. Higgins 1087 (BRY);
River Bench, W.C. Muenscher 2415 (UTC);
Co.. Mueller Park, K. Brizzee 7876 (UT); Salt
City Creek Canyon, s.n. (UT); Bonneville
Flowers 444 (UT); City Creek, M.E. Jones
; Dry Fork Canyon, P.C. Famsworth s.n.
(UT); Utah Co., west side of Mount Timpanogos, R.
Gourley 8086 (UT).

(UT);
129.52
( UTC )
Logan
Davis
Lake Co.,
Terrace, S.
s.n. (RM)

.37D. Cryptantha humilis (Gray) Payson, var.
ovina (Payson) Higgins, Brigham Young
University Sci. Bull. Vol. 13, No. 4:37.
1971. Type: Vicinity of Current, Nye
County, Nevada, G. H. Bentley s.n.
Cryptantha nana (Eashv.) Payson, var.
ovina Payson, Ann. Mo. Bot. Card.
14:314. 1927.

Densely caespitose long-lived perermials, 0.5-
1.5 dm tall; stems several, 0.2-0.7 dm long;
knives spatulate to oblanceolate, obtuse, 2-4 cm

40

BmcnAM YouNc llNivKHsn v Science Bulletin

Cn/i>kinllui liuinilix var. sluiiitzii

long, tomentose and appressed setose with
rather weak bristles; inflorescence narrow, cylin-
drical, lower foliar bracts inconspicuous; calyx
segments linear-lanceolate, densely setose and
tomentose; style exceeding the mature fruit 0.5-
1 mm. Growing on gravelly loam to clay soils, in
the Pinyon-Juniper community. Southwestern
Utah, southern Nevada, and southeastern Cali-
fornia. April-July. Fig. 51.

Only a few of the several hundred specimens
of the various varieties of C. Jiutnilis have been
cited in this paper for two reasons: (1) most
collections are so immature that it is almost im-
possible to assign them to a particular variety;
(2) in order to conserve space in the paper.
Future collections of Cryptantha would be much
more adec|uate if fruiting plants rather than
flowering specimens were collected, particularly
with members of this species.

Washington Co.. Bt'avcrclam Mountains, L.C. Hig-
gins 12.34 (BRY); summit of the Beaverdam Mountains,
L.C. Higgins 1409 (BRY).

.38. Cryptantha jamesii (Torr. ) Payson, Ann.
Mo. Bot. Card. 14:242. 1927.

Perennials, 1-6 dm tall; stems 1-many, 0.4-4
dm long, glabrous to conspicuously hirsute;
leaves linear to broadly oblanceolate, obtuse to
acute, 2-15 cm long, 0.2-1.5 cm wide, glabrous

to hirsute, usually pustulate dorsally, ventral
surface lacking pustules or the pustules very in-
conspicuous; inflorescence open, cymules usually
elongating, tomentose to setose-hirsute, floral
bracts inconspicuous to very conspicuous; calyx
segments ovate-lanceolate, acute, in anthesis 3-4
nun long, in fruit 5-7 mm long, subtomentose to
setose-hirsute; pedicels 1-3 mm long; corolla
white, the tulx' 2.5-3 mm long, crests at base of
tube conspicuous, fomices light yellow, emar-
ginate, 0.5-1 mm long, limb 5-8 mm broad; fruit
(>blate= ovoid, 1-4 nutk'ts maturing, ovate-lan-
ceolate, margins acute, 2-2.5 mm long, 1.5-2 mm
wide, the margins not in contact, botli surfaces
smooth and glossy, scar straight, closed, extend-
ing from the base to near the apex; style exceed-
ing mature fruit 1-3 mm.

1. Ventral surface of the leaves glabrous, the petioles
not ciliate-margined, nor the leaves tufted at the
base 38A. var. pustulosa

1. Ventral surface of the leaves strigose or setose, the
petioles ciliate-margined; leaves tufted at the base
(2).

2. .Stems simple, not branched aljove the base (3).

2. Stems branched from the base as well as
;ibc>ve 388. var. disticha

3. Stems 1-4.4 dm long, usually twice as long as the
basal tuft of leaves 38C. var. multicaulis

3. Stems 0.2-0.9 dm long, usually not exceeding the
bas;d tuft of le;ives 38D. var. setosa

UTAH

County Lines and
river drainages

ir

l''ig. .51. Cnipltiiithd liiiinilix \.ir. oviiid

Biological Series, Vol. 16, No. 3 The BonAGiN.\CE.\E of Ut.\h

41

38A. Cnjptantlui jamesii (Torr. ) Payson, var.
])u.stulosa (Rydb. ) Harringt. Man. P. Colo.
466, 641. 1954. Type: San Juan County,
Utah.
Oreocanja jmsttdosa Rydb. Bull. Torrey

Club 40:480. 1913.
Cnjptantha jmsttdosa (Rydb.) Payson,

Ann. Mo. Bot. Card. 14:252. 1927.

Erect perennials, 2-5.8 dm tall, branched
from the base, simple above; stems slender, 1-3.9
dm long, glabrous or finely strigose; leaves linear
to broadlv oblanceolate, 2-9 cm long, 0.4-1.5 cm
wide, the dorsal surface appressed setose-pustu-
late, ventral surface glabrous, the petioles not
ciliate-margined; inflorescence open, 0.4-2 dm
long, floral bracts inconspicuous. Sandy to grav-
elly soils, mainly in the Pinyon-Juniper com-
inunitv. Southwestern Colorado, northwestern
New Mexico, northeastern Arizona, and south-
eastern Utah. May-August. Fig. 52.

UTAH

County Lines and
river drainages

i-

Fig. .52, Cnjptantha jamesii \ar. pustulosa

Garfield Co., King Ranch, Boulder, Utah, Beck &
Tanner 8204 (BRY); about 4 miles south of Boulder,
D. Atwood 1864 (BRY); Kane Co., Mokie Tanks, DE.
Beck s.n. (BRY); San Juan Co., east slope of Elk Ridge,
Magiiire & Redd 206.5 (UTC); between Blanding and
Kigalia Ranger Station, A.H. Holmgren 3489 (BRY);
10 miles west of Blanding, D. Henriques H-9 (UT);
Hammond Canvon, Elk Mountains, Rydberg & Gar-
rett 9569 (UT)': Arch Canyon, 18 miles' west of Bland-
ing, Cronciuist & Holmgren 9372 (UTC); bottom of

Comb Wash about 20 miles west of Blanding along hwy
95, Welsh, Atwood & Higgins 8933 (BRY).

38B. Cnjptantha jamesii (Torr.) Payson, var.
(Usticha (Eastw. ) Payson, Ann. Mo. Bot.
Card. 14:248. 1927. Type: Bartons Range,
San Juan County, Utah, Alice Eastwood
90.

Oreocanja disticha Eastw. Bull. Torrey
Club 30:2.38. 1903.

Erect perennials, branched from the base as
well as above, 2.5-4.2 dm tall; stems somewhat
woody near the base, 1.2-2.9 dm long, strigose
and weakly setose; leaves narrowly oblanceolate,
3-12 cm long, dorsal surface setose-pustulate
and strigose, ventral surface strigose to setose
or silky-strigose, without pustulate hairs or the
pustules inconspicuous, the petioles ciliate-mar-
gined; inflorescence open, 0.5-2 dm long, fo!iar
bracts not very conspicuous. Southwestern Colo-
rado, northeastern Arizona, northwestern New
Mexico, and southeastern Utah. Usually found
growing on sandy soils. April-September. Fig.
53.

Emery Co., 4 miles up Calf Spring Wash, San
Raphael Swell, B. Maguire 18446 (UTC); first fork of
Calf Spring Canyon, San Raphael Swell. B. Maguire
18298 (UTC); Buckhorn Wash, B.F. Harrison 9643
( BRY ) ; junction of Buckhom Wash and San Raphael
River, Welsh & Atwood 9829 (BRY); Garfield Co., 5
miles south of Boulder, L.C. Higgins 1011 (BRY); 4

County Lines and
river drainages

^ so *«

Kig. .53. Cri/plantlut jamesii var. disticha

42

BllK^HAM VdUNG UnIVEKSII Y SCIENCE BULLETIN

.nlles south of HenrieviUe, L.C. Higgins 1010 (BRY);
Hi miles west of Bryce Canyon Natl. Park, B.C. Rollins
2449 (UTC); 5 miles east of Escalante, A.H. Holmgren
7716 (UTC); 10 miles east of Escalante, A.H. Holmgren
7748 (UTC); Kane Co., 3.7 miles east of Skutumpah-
Alton junction at head of Jolmson Canyon, D. Atvvood
1804 ( BRV ) ; about 7 miles east of Skutumpah-Alton
junction, D. Atwood 1807 (BRY); one h;ilf mile west
of Paria Ri\er bridge along hwy 89, Welsh & Atwood
9748 (BRY); 47 miles east of' Kanab, B.F. Harrison
12092 (BRY); San Juan Co., bottom of Comb Wash
along hwy 47 west of Bluff, Welsh & Ahvood 9970
(BRY); Monument Valley, Eastwood & Howell 6674
(UTC); near summit of Navajo Mt., A.H. Holmgren
10661 (UTC); Mexican Hat, B. Maguire 16291 (UTC);
Monument Valley, A.H. Holmgren 3243 (BRY, UTC);
10 miles west of Bluff, C.A. Hanson 87 (BRY); mesa
north of Bluff. B. Magnire .5101 (UTC); head of Cow
Canyon. 2 miles north of Bluff, AH. Holmgren 3213
(UTC); Wayne Co., junction of road to Ekkers Ranch,
Welsh & Atwood 9859 (BRY); 3 miles north of Grover,
D. Anvood 1862 (BRY); 3 miles southeast of Torrey,
Hohngren, Reveal & LaFrance 1091 (BRY. UTC);
about 2 miles south of Lyman, L.C. Higgins 1014
( BRY ) ; 5 miles east of Teasdale. Holmgren, Reveal &
LaFrance 2551 (BRY).

38C. Cnjptantha jarnem ( Torr. ) Payson, var.
muhicaulw ( Torr. ) Payson, Aiin. Mo. Bot.
Gard. 14:244. 1927. Type: New Mexico
near Santa Fe, Fendler 636.
Eritrichiiirn multicaule Torr. in Marcy,

Expl. Red River 262. 1S,54.
Oreocanja mitlticaulus ( Torr. ) Greene, Pit-

tonia 3:114. 1896.
Oreocanja suffruticosa ( Torr. ) Pay.son, var.

midticauli.s (Torr.) Payson, Univ.

Wyo. Publ. Bot. 1:171. 1926.
Hemisphaerocarija sitffniticosa ( Torr. )

Brand, var. midticaulls ( Torr. ) Brand,

Rep. Spec. Nov. 24:60. 1927.

Perennial, 2-5.5 dm tall, branched from the
base, simple above; stems slender, 1-4.4 dm long,
weakly strigose-setose; leaves mostly basal, ob-
lanceolate, 5-15 cm long, 0.4-1 cm wide, dorsal
surface strigose and appressed setose, or some-
times seto.se-hirsute, pustulate, ventral surface
unifonnly strigose or subtomentose, without
pustules, or the pustules small and inconspicu-
ous, the petioles conspicuously ciliate on the
margins; inflorescence open, 0.5-1.5 dm long,
bracts inconspicuous. Growing on a wide variety
of soils, but mainly in sands in Utah. Southern
Colorado and eastern New Mexico, south to
western Oklah(«n;i ;ind Texas into northern Mex-
ico, and north through central Arizona to south-
em Utah. April-September. Fig. .54.

Ciiirfield Co., canyon above Tropic, M.E. [ones
5.300 (US); Kane Co., 4 miles north of Kanab, Ripley
& Banieby 4854 (CH); plate;ui near head of Hock-
Creek, B.F. Harrison 9040 (BRY); on slopes southeast
of dunes, Knudsen 44 (BRY); Coral Pink Sand Dunes,

S.L. Welsh .5299 (BRY); 23 miles south of Alton, A.
Cromiuist 10191 (UTC); Johnson Canyon, D. At\vood
1789C (BRY); Washington Co., Zion Canyon Natl. Park,
east tunnel road, W.P. Cottam 4763 (BRY); Zion Natl.
Park, Clear Creek Canvon, Eiistwood & Howell 9219
(CH).

38D.

Crtjjdantha jamesii (Torr.) Payson, var.
setusa (Jones) Johnst. ex Tidest. Proc. Biol.
Soc. Wash. 48:42. 1935. Type: Cove Fort,
Utah, Jones s.n.

Oreocanja
1896.

Knjnitzkui
Contr.

Oreocanja

cinerea Greene, Pittonia 3:113.

setosa Jones,
1910.
Bull. Torrey

multicauli.i var.
West. Bot. 13:4.
iemmoni Eastw.

Club .30:2,39. 1903.
Oreocanja muUicaulis var. cinerea

(Greene) Macbr. Proc. Amer. Acad.

51:.54. 1916.
Oreocanja suffruticosa var. cinerea

(Greene) Payson, Univ. Wyo. Publ.

Bot. 1:171. 1926.
llemisphaerocanja suffruticosa var. setosa

(Jones) Brand, Rep. Spec. Nov. 24:60.

1927.
Hemisphaerocanja cinerea (Greene)

Brand, Rep. Spec. Nov. 24:61. 1927.
Cnjptantha jamesii var. cinerea (Greene)

Payson, Ann. Mo. Bot. Gard. 14:246.

1927.

UTAH

County Lines and
river drainages

i 20 fco

Fili;. 54. ('n/fitunffui ja})u:\ii \,ir. niiilticftuli.s

Biological Series, Vol. 16, No. .3 The Bor.aginaceae of Utah

43

Perennial, 1-3 dm tall, branched from the
liase, simple above; .stems slender, 0.2-0.9 dm
long, strigose and weakly setose; leaves mostly
basal, oblanccolate, obtuse, .3.?-13 cm long, 0.4-
1.5 cm wide, dorsal surface finely strigose, usual-
Iv conspicuously pustulate, ventral surface uni-
fonnly and densely strigose, the petioles con-
spicuously ciliate on the margins; inflorescence
open, 0.4-2 dm long, bracts evident, especially
near the base of the inflorescence. Growing on
a wide variety of soils, especially in the Pinyon-
Juniper community. South central Colorado,
northern New Mexico, northern Arizona, eastern
Nevada, and southern Utah. May-September.
Fig. 55.

County Lines and
river drainages

i-

3e ho

Fig. 55. Cryptantha jatticsii var. setosa

Emery Co., about 10 miles south of the San Raphael
River bridge, San Raphael Swell, L.C. Higgins 1321
(BRY); Garfield Co.. near milepost 41 south of Circle-
ville, L.C. Higgins 13.52 (BRY); head of Red Canyon,
B. Maguire 19053 (UTC); 5 miles south of Panguitch,
B. Maguire 18706 (UTC); Iron Co., Beryl Desert, C.
Lambert 34L (BRY, UT); Kane Co., 20 miles east of
Kanab, D. Atwood 1792A (BRY); 4 miles south of
Cannonville, A. Cronquist 10213 (UTC); 24 miles west
of Collets Wash, D. Atwood 1873A (BRY); about 32
miles southeast of Kanab, D. Atwood 1.535 (BRY); Cot-
tonwood Canyon, 20 miles south of Cannonville, L.C.
Higgins 1009 (BRY); Sevier Co., Fish Creek Canyon,
A.O. Garrett 2.5.57 (UT); Washington Co.. Mountain
Meadows, W.P. Cottam .3351 (BRY); Mountain Mead-
ows, W.P. Cottam 6770 (UT); Wavne Co., 7 miles
southeast of Teasdale, B.F. Harrison 7.529 (BRY); 1

mile west of Bicknell, R.K, Vickery 604 (UT); 1 mile
southeast of Torrey, L.C. Higgins 1345 (BRY).

39. Cnjptantha johnstonii Higgins, Great Basin

Naturalist 28:195. 1968. Type: About 15
miles west of hwy 50-6 along the road from
VVoodside to Huntington, Emery County,
Utah, L.C. Higgins 1310.

Caespitose perennial 1-2.5 dm tall; stems
several, arising from the branched caudex, 0.6-
1.3 dm long, very weakly strigose; leaves ob-
lanccolate, the apices obtuse to acute, 2-6.5 cm
long, 0.4-1 cm wide, dorsal surface strigose with
conspicuous pustulate hairs; inflorescence some-
what open, 0.5-2 dm long; floral bracts evident
but not conspicuous, 1-2 cm long; calyx seg-
ments linear-lanceolate, in anthesis 5-6 mm long,
in fruit becoming 8-10 mm long, strigose and
spreading white-setose; pedicels 0.5-1 mm long;
corolla white, the tube 12-15 mm long, flaring in
the throat, crests at base of tube lacking, fomices
yellow, 1-1.5 mm long, emarginate, papillose,
limb 13-17 mm broad; nutlets ovate, 3-3.5 mm
long, 2.3-2.7 mm wide, usually all 4 maturing,
margins acute and knifelike, in contact, both sur-
faces smooth and glossy, scar straight, closed,
elevated margin lacking; style exceeding mature
fruit 3-8 mm ( hetero.styled ) . Growing in clay or
clay-loam soils. Known only from the type local-
ity near the head of Cottonwood Wash on the
San Raphael Swell. May-June. Fig. 56.

Emery Co., about 15 miles west of hwy .50-6 along
the road from Woodside to Castle Dale, L.C. Higgins
1310 (BRY); San Raphael Swell, B.F. Harrison 5628
(BRY); about 6 miles down Cottonwood Wash. L.C.
Higgins 3522 (BRY); San Raphael Swell, at the north
end of Cottonwood Wash, L.C. Higgins 3.520 (BRY,
WTSU).

40. Criiptantha jonesiana (Payson) Payson,
Ann. Mo. Bot. Card. 14:323. 1927. Type:
San Raphael Swell, Emery County, Utah,
M. E. Jones s.n.

Oreocarija jonesiana Payson, Univ. Wyo.
Publ. Bot. 1:168. 1926.

Coarse caespitose perennials, 0.5-1.5 dm tall;
stems many, arising from a thick woody multi-
ple caudex, 0.2-0.7 dm long, setose; leaves spatu-
l;vte, 1-4 cm long, 0.4-1.3 cm wide, coarsely ap-
pressed setose-pustulate, leaf bases also setose
with dense white hairs; inflorescence narrow,
somewhat capitate, with 1-3 flowers in the axils
of the bnicts below the terminal cluster; calyx
segments lanceolate to nearly linear, in anthesis
5-7 mm long, in fruit becoming 7-10 mm long,
densely setose, with ascending yellowish bristles;
corolla white, the tube 10-15 mm long, campanu-

44

HuioiiAM YouNc. Univehsitv Science Bulletin

UTAH

County Lines and
river drainages

i-

Fig. 56. Cnjptantha joluisfomi

late in the throat, fomices low and broad, papil-
lose, crests at base of tube lacking, limb 9-13
mm wide; nutlets lanceolate, 3.5-4.5 mm long,
densely and uniformly muricate, or with a few
short, low ridges, scar narrow, open, and without
an elevated margin; style exceeding mature fruit
4-6 mm. Barren clay hills. Endemic to the San
Raphael Swell, Emery County, Utah. April-May.
Fig. 57.

Emen,' Co., .San Raphael Swell, W.P. Cottam 5247
(UT); about 10 miles .south of the San Raphael River
bridge, San Raphael Swell, L.C. Higgins 1322 (BRY);
about 15 miles west of hwy 50-6 along the road from
Woodside to Castle Dale, L.C. Higgins 1308 (BRY);
San Raphael Swell, about 1 mile south of river bridge,
Higgins & Reveal 1265 (BRY); San Raphael Swell, D.
At\vood 1301 (BRY).

41. Criiptaniha JongifUmi (A. Nels.) Payson,
Ann. Mo. Bot. Card. 14:326. 1927. Type:
Palisades, Mesa County, Colorado, C. S.
Crandall s.n.
Oreocanja longiflom \. Nels. Erythea 7:67.

1899.
Oreocanja liorrklulu (Jreene, PI. Baker. 3:20.

1901.

Short-lived perennial or possibly biennials,
0.8-3 (5) dm tall; stems l-several, 0.5-1 dm long,
setose antl spreading hirsute; leaves spatulate,
obovate or oblanceolatc. 2-7 cm long, 0.5-1.5

Fig. .57. Cryptantlui joncsiana

cm wide, both surfaces strigose and strongly hir-
sute, pustulate; inflorescence broad and open,
0.7-2.5 dm long, setose, foliar bracts inconspicu-
ous; calyx segments linear-lanceolate, in anthesis
7-10 mm long, in fruit becoming 10-16 mm long,
setose; corolla white, the tube 12-14 mm long,
crests at base of tube lacking, fomices yellow,
emarginate, broad, rounded, papillose, 0.5-1 mm
long, limb 9-11 mm wide; style exceeding mature
fruit 4-9 mm (heterostyled); nutlets lanceolate-
ovate 3-4 mm long, 2.2-2.6 mm wide, 2-4 matur-
ing, both surfaces with tubercles and low round-
ed ridges, scar straight, closed or very narrowly
open, with a slightly elevated margin. Sandy
to clayev open ridges and flats. Western Colora-
do and eastern Utah along the Colorado River
drainage. May-June. Fig. 58.

Crand Co., bluffs along the Colorado River, about
10 miles south of U.S. hwy 50-6 along hwv 24, S.L.
Welsh 6966 (BRY); Morrison Formation along the
Colorado River about .32. .5 miles from Moab, S.L. Welsh
6989 (BRY); near Cisco. L.C. Higgins .3314 (BRY);
near milepost 32 along hwy 128, L.C. Higgins 1478
(BRY).

42. Cn/ptantha rnensana (Jones) Payson, Ann.
Mo. Bot. Card. 14:333. 1927. Type: Emery
County, Utah. M. E. Jones 5445p.

Oreocanja mensarui (Jones) Payson, Univ.
Wyo. Publ. BoL 1:171. 1926.

BioLocicAi, Sehies, Vol. 16, No. 3 The BoiL-iciNACEAE ok Utah

45

UTAH

Fig. 58. Crijptunthfi huij^iflora

Krijnitzkia mcnsana Jones, Contr. West. Bot.
1.3:4. 1910.

Short-lived perennials, 1-1.5 (2) dm tall;
stems 1-several, 0.5-1.2 dm long, setose-hirsute,
with some finer strigose hairs beneath; leaves
oblanceolate to spatiilate, obtuse, 3-8 cm long,
0.5-1.4 cm wide, lower surface setose with pustu-
late hiiirs, also finely strigose, ventral surface
strigose, less setose and with fewer pustules; in-
florescence broad, open, 0.4-1.2 dm long, setose,
foliar bracts well developed; calyx segments
lanceolate, in anthesis 4-5 mm long, in fruit be-
coming 7-8 mm long, setose-hirsute; corol-
la white, the tube 3-4 mm long, crests at
base of tube lacking or nearly so, fomices yel-
low, rounded, slightly papillose, about 0.5 mm
long, limb 5-8 mm wide; nutlets ovoid, 3-3.5 mm
long, 1.6-1.9 mm wide, margins obtuse, not in
contact, dorsal surface rugose, tuberculate and
somewhat muricate, ventral surface conspicuous-
ly tuberculate, scar open, constricted at the mid-
dle and surrounded by a high, elevated margin;
style exceeding mature fruit 1.5-2 mm. Clay to
clay-loam soils. Central and eastern Utah in
Emery, Carbon, and Grand counties. April-July.
Fig. 59.

Carbon Co., 20 miles north of Wellington, B, Ma-
guire 18.536 (UTC); left fork of Minnie-Maud Creek,
B. Maguire 18596 (UTC); West Tavaputs Plateau, B.
Maguire 18503 (UTC); 5 miles ea.st of Wellington, L.C.

Higgins 996 (BRY); 5 miles east of Wellington, Hig-
gins & Welsh 1043, 1041 (BRY); near Carbon-Emery
County line along hwy .50-6, Higgins 3323 (BRY,
WTSU); Emery Co., about 1.5 miles west of hwy 50-6
along the road from Woodside to Castle Dale, L.C.
Higgins 1306 (BRY); about 7 miles south of the San
Raphael River bridge, San Raphael Swell, L.C. Higgins
1318 (BRY); Carbon-Emery County hne along hwy
50-6, Higgins & Welsh 1039 (BRY); San Raphael River,
B.F. Harrison 8060 (BRY); about 9 miles south of
Lawrence, G.L. Pyrah 15 (BRY); Grand Co., vicinity of
Sego, about 5 miles north of Thomp.son, S.L. Welsh
6915 (BRY); Wayne Co., 5 miles east of Buckhom
Wash, Welsh & Atwood 9908 (BRY).

43. Cnjptantha ochroleuca Higgins, Great Ba-
sin Naturalist 28:197. 1968. Type: Garfield
County, Utah, on outcrop 100 meters south
of Red Canyon Campground along hwy 12,
L. C. Higgins 1788.

Low caespitose perennial, 0.2-1.3 dm tall;
stems several, 0.1-0.4 dm long, strigose and
weakly setose; leaves linear-oblanceolate to ob-
lanceolate, the apices acute or sometimes obtuse,
1-2.5 mm long, 0.1-0.3 cm wide, basal leaves uni-
formly and densely strigose, sparsely setose, the
petioles white-hairy, cauline leaves strigose and
with some setose-pustulate bristles; inflorescence
narrow, 0.2-0.7 dm long, weakly setose; calyx
segments linear-lanceolate, 2-2.5 (3) mm long in
anthesis, in fruit becoming 4-6 mm long, setose;
corolla pale-yellow, the tube 2-2.5 mm long,

UTAH

County Lines and
river drainages

Mj^. 59. CTijptantha nicnsand

46

BniCHAM Y(;UNG Univehsitv Science Bulletin

cTi'sts at base of tube conspicuous, fomices yel-
low, rounded, about 0.3 mm long, limb 4-5 mm
wide; nutlets lanceolate, 2.5-3 mm long, 1.4-1.6
mm wide, usually onlv 1 maturing, margin acute,
dorsal surface irregularly rugose with low round-
ed ridges, ventral surface only slightly uneven,
scar open, narrowly triangular, extending % the
length of the nutlet, and lackuig an elevated
margin; style scarcely surpassing the mature
fruit. Limited to the red Wasatch Fomiation
near Red Canyon Campground in southwestern
Garfield County, Utah. May-August. Fig. 60.

UTAH

Cnjptunthu ochrolcuca

Garfield Co.. on outcrop 100 meters south of Red
Canyon Campground, L.C. Higgins 1788 (BRV); Red
Canyon Campground, Reveal & Reveal 10.31 (BRY);
Kocky outcrops north of Red Canyon Campground, D.
Atwood 1891 (BRY); 2.4 miles east of Red Canyon
Campground, D. Atwood 1879 (BRY).

44. Crijpkintba osterhoutii (Payson) Payson,
Ann. Mo. Bot. Card. 14:.329. 1927. Type:
Monument Park, near Grand Junc-tion, Col-
orado, G. E. Osterhout 6138.
Oreocart/a osterhoutii Payson, Univ. Wyoni.
Pub). Bot. 1:167. 1926.

Den.sely caespitose perennials, 0.7-1.2 dm tall;
stems slender, many, arising from the densely
branched multiple caudex, 0.3-0.6 dm long, stri-
gose and spreading setose; leaves spatiilate to
oblanceolate, obtu.se, 1-3 cm long, 0.3-0.8 cm

wide, dorsal surface strigosc and appressed se-
tose, pustulate, ventral surface strigose, not pust-
ulate or the pustules inconspicuous, the petioles
ciliate-margincd; inflorescence open, 0.3-0.8 dm
long, weakly white setose, foliar bracts incon-
spicuous; calyx segments lanceolate, in anthe-
sis 2.5-4 mm long, in fruit becoming 5-6.5 mm
long, strigose and spreading white setose; corolla
white, the tube 2-3 mm long, crest at base of
tube usually evident but poorly developed, fom-
ices yellow, broad, emarginate, papillose, about
0.5 mm long, limli 5-7 mm wide; nutlets lanceo-
late, 2.7-3.2 mm long, 1.8-2.2 mm broad, usually
less than 4 nutlets maturing, margins obtuse, not
in contact, dorsal surface carinate, sharply tuber-
culate and rugose, ventral surface sharply tu-
berculatc, scar open, constricted above the base,
elevated margin evident but not conspicuous;
style exceeding mature fruit 0.2-0.7 mm. Sandy
soil or rocky ledges and slopes of the Pinyon-
Junipcr community. Mesa County, Colorado,
Wayne and San Juan counties, Utah. May-June.
Fig! 61.

UTAH

County Lines and

river drainages

I I 1

o b5 5o

Fig. (>1. Criiptiintlui osterhoutii

Sail Juan Co., Elepliant Hill, Cauv'onlands Natl.
Park, W.'Ish. Atwood & Higgins 887.5 (BRY); 25 mUes
west of Bluff near Johns Canyon, B.F. Harrison 1192.3
(BRV); nine miles north of junction 47-261 at summit
of .lugway. D. Atwood l.'5.38A (BRY); Wayne Co.,
North Point, ahont 8 miles northwest of road to Flint
Trail, above Orange Cliffs, Welsh ti Atwood 9872
(BRV).

Biolocic:al Series, Vol. 16. No. 3

The Bon,\c:iNACE.\E of Ut.\h

47

45. Cn/ptantlui paradoxci (A. Nels.) Payson,
Ann. Mo. Bot. Card. 14:330. 1927. Type:
Paradox Valley, Montrose County, Colorado,
E. P. Walker 91.

Oreocarija paradoxa A. Nels. Bot. Gaz.

56:69. 1913.
Oreocarija gijpsophila Payson, Bot. Gaz.

60:3S0. 1915.

Caespitose perennial, 0.4-1.2 dm tall; stems
1-many, slender, 0.2-0.8 dm long, subtomentose
near the base, weakly setose above; leaves ob-
ianceolate to spatiilate, usually folded, obtuse,
1.5-4 em long, 0.2-0.4 em wide, dorsal surface
with appressed setose-pustulate hairs, ventral
surface uniformly strigose and without pustulate
hairs, the petioles ciliate-margined; inflorescence
subcapitate, 0.1-0.4 dm long, setose, foliar bracts
inconspicuous; calyx segments linear-lanceolate
in anthesis 5-6 mm long, in fruit becoming 6-8
mm long, weakly setose; corolla white, usually
with a vellow tube 10-12 mm long, crests at base
of tube lacking, fomices vellow, broad, emar-
ginate, papillose, about 0.5 mm long, limb 10-12
(15) mm wide; nutlets lanceolate, turgid, 2-3
mm long, 1.3-1.6 mm wide, all 4 usually matur-
ing, margins acute to obtuse, not in contact,
dorsal surface densely tuberculate and conspicu-
ously rugose, ventral surface tuberculate, also
somewhat rugulose, scar open, constricted be-
low the middle, the margin elevated; style ex-
ceeding mature fruit 4-9 mm. Sandy to heavy
clav soils on flats and open ridges. Western
Colorado, northwestern New Mexico, and east-
em Utah. May-June. Fig. 62.

Emerv Co., 5 miles south of the San Raphael River,
IVF. Harrison 9607 (BRV): San Raphael Swell. 50 miles
north of Hanksville, A. Cronquist 9204 (UTC); San
Raphael Swell, B. Maguire 18282 (UTC); .3 miles south
of San Raphael River Bridge, San Raphael Swell, L.C.
Higgins 1314 ( BRY): 10 miles south of the San Raphael
River Bridge, San Raphael Swell, L.C. Higgins 1319
( BRV ) .

46. CrijptantJia rollinsii Johnst. Joum. Am. Arb.
20:391. 1939. Type: on Thomes Ranch near
Willow Creek, 22 miles south of Ouray,
Uintah County, Utah, R. C. Rollins 1715.

Biennial herbs, 1-3.5 dm tall; stems 1-several,
0.2-1 dm long, setose, leaves clustered at the
base, gradually reduced upward, oblanceolate to
spatulate, obtuse to acute, 2-5 cm long, 0.5-1.5
cm wide, setose and hispid, pustulate on both
surfaces; inflorescence narrow to somewhat open
at maturity, cvlindrical or obovoid. racemes in
dense glomemles, 3-6 flowered, hispid, 0.5-2 dm
long; calyx segments linear, in anthesis 7-8 mm
long, in fruit becoming 8-10 mm long, hispid;

UTAH

County Lines and
river drainages

A a5 iio

Fig. 62. Crijptantha fxiradoxa

corolla white, campanulate, the tube 7-9 mm
long, crests at base of tube evident, fomices
vellow, papillose, about 0.5-1 mm long, limb 7-S
mm wide; plants slightly heterostyled; nutlets
lanceolate, 3-4 mm long, 1-1.5 mm wide, ob-
scurely rugulose and tuberculate on the dorsal
surface, ventral surface smooth, scar closed, and
without an elevated margin. Open hills and
ridges, growing on white or red shale. Central
and northeastem Utah in Emery, Uintah, and
Duchesne counties. May-July. Fig. 63.

Duehesne Co., about 10 miles south of Myton, L.C.
Higgins 1067 (BRY); 10 miles south of Duehesne, L.C.
Higgins 1052 ( BRY ) ; Indian Canyon, 10 miles south of
Duchesne, B.F. Harrison 40t)H (BRY); between Myton
and Wellington, 19 miles south of junction with U.S.
hwv 40, Holmgren, Reveal & LaFrance 1946 (BRY); 2
miles south of Bridgeland, L.C. Higgins 1068 (BRY);
20 miles south of Myton, L.C. Higgins 1063 (BRY);
Parley Canyon, tributary to Argyle Canvon, south end
of county, Welsh & Christensen 6622 (BRY); 1 mile
south of Duchesne, L.C. Higgins 10.56 (BRY); about
8 miles north of Duchesne, L.C. Higgins 10.58 (BRY);
Emery Co., near Temple Mountain, L.C. Higgins 1324
(BRY); west of Hidden Splendor Mine, D. Atwood
1846 (BRY); 8 miles north of Goblin Valley tumoff
on Temple Mountain road, D. Atwood 1859' (BRY);
about 5 mill's east of Buckhom Wash, along road north
of San Raphael River. Welsh & Atwood 9843 (BRY);

1 1 miles north of Goblin Valley tumoff on Temple Mt.
road, thence 40 miles west-southwest on mining road,
D. Atwood 1858 (BRY); Uintah Co.. Hill Creek, about

12 miles south of Ouray, J. Brotherson .541 (BRY); Hill
Creek, .33 miles south of Ouray. ]. Brotherson 473

48

Biiir.iivM Young Uni\ihsitv Science Bulletin

UTAH

County Lines and
river drainages

o 30 io

Fig. 63. Cn/plantha rollinsii

( BRY ) ; about .5 miles south of White River bridge, D.
.\t\vood 1617, 1618A (BRY); Dino.saur Natl. Monu-
ment, Split Mountain Corge Campground, J. Brotherson
994 (BRY); 18 miles south of Ouray, L.C. Higgins 1875
(BRY); .3 miles west of Bitter Creek, L.C. Higgins 1581
(BRY).

47,

Cnjptantha rugulosa (Payson) Payson, Ann.
Mo. Bot. Card. 14:295. 1927. Type: Fish
Spring.s, Juab County, Utah, M. E. Jones s.n.
Oreocariju rugulosa Payson, Univ. Wyom,
Puhl. Bot: 1:166. 1926.

Biennial or .short-lived perennial, 1.2-3 dm
tall; stems slender, 1-several, 0.8-1.6 dm long,
spreading scstosc-hispid; leaves oblaneeolate to
spattiiate, obtu.se to acute, strigose and conspicu-
ously setose-hispid, pustulate on both surfaces;
inflorescence 0.2-2 dm long, hispid; foliar bracts
inconspicuous; calyx segments linear-lanceolate,
in iinthesis 4-5 mm long, in fruit becoming 7-9
mm long, strigose ;md spreading hirsute; corolla
white, the tube 3-4 mm long, crests at base of
tube conspicuous, fomices rounded, distinctly
p;ipillose, iibout 0.5 mm long, limb 5-7 mm
broad; nutlets lanceolate, 2.8-3.2 mm long, 1.3-
1.7 mm wide, all 4 usually maturing, margins in
contact, acute, dorsal surface with short low
ridges, also somewhat tubt^rculate, ventral sur-
face smooth or nearlv so, scar open, subulate,
without an elevated margin; stvle exceeding

mature fruit 1-1.5 mm. Upper Sonoran Zone in
gravelly loam to clay soils. Central Utah to
northeastern Nevada. May-July. Fig. 64.

Juab Co., 7 miles east of Trout Creek, Maguire &
Becraft 2761 (UTC); 13 miles east of Trout Creek, Ma-
guire & Becraft 2760 ( UTC ) ; near Topaz Reservoir,
B.K. Harrison 11796 (BRY); Millard Co., Desert Range
Experiment Station, W.P. Cott;im 8.523 (UT); 10 miles
west of Fillmore, Ice Spring Crater, W.P. Cottam 9569
( UT ) ; 26 miles west of Delta, Magnire & Becraft 3943
( UTC ) ; 9 miles north of Desert Range Experiment Sta-
tion Headquarters, R.C. Holmgren 519 (BRY); 1 mile
south of Grandy, B.F. Harrison 116.58 (BRY); Cinder
Cones. W.P. Cottam 8003 (UT); House Range, 37 miles
west of Delta, Maguire & Becraft 2758 (UTC); Cow-
bov Pass, Q. A, Hare I30F (UTC); 20 miles west of
Hinckley, B. Maguire 20759 (UTC); about 15 miles
west of Desert Range Experiment Station, L.C. Higgins
1614. 1463 (BRY); Sevier Co., about 1 mile southeast
of Sigurd, L.C. Higgins 1623 (BRY); 6 miles south of
Big Rock Candv Mountain, D. Atwood 1787 (BRY); 1
mile south of Sevier. L.C. Higgins 1474, 1621 (BRY);
Tooele Co., 4 miles north of Gold Hill, B. Maguire
22021 (UTC); 2 miles west of Gold Hill. Maguire &
Becraft 2762 (UTC); near Ibapah, W.P. Cottam 3159
(BRY); about 3 miles east of Wendover, L.C. Higgins
1720 (BRY).

48. Criiptantha semiglahra Bameby, Leaf. West.
Bot. 3:197. 1943. Type: Arizona, about 2
miles east of Fredonia, Ripley & Bameby
8429.

Erect perennials, 2-3 (4) dm tall; stems 1-
several, 0.9-1.8 (2) dm long, retrorsely strigose

UTAH

County Lines and
river drainages

is-

Kig. 64. Cri/iilinitha nii^iilosii

Biological Series, Vol. 16, No. 3 The Bouaginaceae ok Utah

49

and weaklv spreading .setose; leaves oblanceo-
late, acute, 3-7 cm long, 0.3-0.6 cm wide, dorsal
surface appressed setose-pustulate, ventral sur-
face glabrous, the old leaf bases long vvhite-
Iiairy; inflorescence narrow to somewhat open,
0.4-1.3 dm long, foliar bracts slightly surpassing
the cymes, 1.5-2 cm long; calyx segments lanceo-
late, in anthesis 5-8 mm long, in fruit becoming
10-13 mm long, setose; pedicels 1-2 mm long;
corolla white, the tube 10-12 mm long, crests
at base of tube conspicuous, fomioes yellow,
rounded, 1-1.2 mm long, obscurely papillose,
limb 8-10 mm wide; nutlets ovate, 3.5-4 mm
long, 2-2.5 mm wide, usually all 4 maturing,
margins acute, in contact, both surfaces smooth
and glossy, scar closed, elevated margin lacking,
style surpasing the mature fruit 5-7 mm. Clay
soils. Upper Sonoran Zone. Northern Arizona
and southwestern Utah. May-July. Fig. 65.

Washington Co., Galagers Hill, Hurricane-Kanab
road, J.W. Harrison s.n. (DIX).

49. Crijptantha sericea (Gray) Payson, Ann.
Mo. Bot. Card. 14:286. 1927. Type: Bridger
Pass, Wyoming, H. Engelmann s.n.
Knjnitzkia sericea Gray, Proc. Amer. Acad.

20:279. 1885.
Oreocanja sericea Greene, Pittonia 1:58.
1887.

UTAH

County Lines and
river drainages

i-

Torrey

(Nels.)
14:288.

I'ig. 6.5. Cn/ptimtlui .'iciiiifilahni

Oreocanja afjinis perennis A. Nels. Erythea
7:67. 1899.

Oreocanja argentea Rydb. Bull. Torrey Club
31:637. 1904.

Oreocanja perennis Rydb. Bull. Torrey Club
33:150. 1906.

Oreocanja procera Osterh. Bull.
Club 47:211. 1920.

Cnjptantha sericea var. perennis
Payson, Ann. Mo. Bot. Gard.
1927.
Perennials, 1.5-4.3 (5) dm tall; stems 1-sever-
al, branched from near the base, 0.5-1.2 (3) dm
long, setose with spreading hairs; leaves ob-
lanceolate to spatulate, obtuse, 2.5-10 (15) cm
long, 0.5-2 cm wide, dorsal surface strigose and
weakly appressed to spreading setose, pustulate,
ventral surface silky-strigose, pustules lacking or
very inconspicuous; inflorescence narrow to
somewhat open, 0.5-3.2 dm long, setose-hispid,
foliar bracts 2-5 cm long; calyx segments lanceo-
late, 2.5-4 mm long in anthesis, in fniit becoming
6-8 mm long; pedicels 0.5-1 mm long; corolla
white, the tube 2.5-3.5 mm long, crests at base
of tube conspicuous, fomices yellow, broad, de-
pressed, 0.5-0.6 mm long, limb 7-9 mm wide;
nutlets lanceolate, 2.5-3.5 mm long, 1.5-2 mm
wide, usually all 4 maturing, margins acute or
narrowly winged, in contact, dorsal surface with
low rounded tuberculations, also somewhat
rugulose and muriculate, ventral surface similar
but the markings less evident, scar straight,
closed and without an elevated margin; style
exceeding mature fruit 0.5-1.3 mm. Growing on
heavy clay soils in the Pinyon-Juniper commu-
nity. Southwestern Wyoming, northwestern Colo-
rado, and northeastern Utah. May-August. Fig.
66.

Carbon Co., West Tavaputs Plateau, Welsh &
Christensen 6.572 (BRY); Daggett Co., 7 miles south of
Manila, R.C. Rollins 1772 (UTC); 6 miles south of
Manila, B. Maguire 12.378 (UTC): Duchesne Co., In-
dian Canyon, A.O. Garrett 8330 (UT); Indian Canyon,
R.S. Ferris 11331 (UTC); about 2 miles south of
Duchesne, L.C. Higgins 1055 (BRY); 10 miles east of
Fniitland, L.C. Higgins 1048 (BRY); about 14 miles
west of Duchesne, Higgins & Welsh 1023 (BRY); Rich
Co., about .5 miles west of Evanston just across the
Utah line, L.C. Higgins 1.571 (BRY); Uintah Co..
Bnish Creek, 1 mile east of sheep corrals, D. Atwood
1585 (BRY); I mile north of Brush Creek sheep pens,
D. Atwood 1.589 (BRY); 5 miles west of White River
bridge, D. Ahvood 1620 (BRY); about I mile west of
Rainbow. Holmgren, Reveal & LaFrance 1806 (BRY);
.3 miles north of Brush Creek sheep corrals, L.C. Hig-
gins 1867 (BRY); Book Cliffs, about 8 miles south of
the junc-tion to Cook Ridge, L.C. Higgins 1596 (BRY).

•50. Cn/jUantJia setosissima (Gray) Payson, Ann.
Mo. Bot. Card. 14:268. 1927. Type: Fish
Lake, Sevier County, Utah, L. F. Ward 646.

50

BmcHAM VouNc; INiviiiMTs Science Bulletin

l''ig. 66. Cn/pttintha scricca

Eritrichiiim setosis.sima Gray, Proc. Aiiier.

Acad. 12:80. 1877.
Krijnitzkki setosis^inui Gray, Proc. Amer.

Acad. 20:276. 1885.
Oreocanja setosissima (Gray) Greene, Pit-

tonia 1:58. 1887.

Biennial or .short-li\ed perennials, 3-10 dm
tall; stems usually 1-3, erect, 1.5-5 (6) dm long,
hirsute; leaves clustered at the base, reduced up-
ward, oblanceolate, the apices obtuse to acute,
3-13 cm long, 0.5-1.5 cm wide, setose, and with
some finer twisted pubescence beneath, pustu-
late hairs numerous on both surfaces; inflores-
cence broad-topped due to the elongation of the
scorpioid racemes, 1-5 dm long; calyx segments
broiidiv lanceolate, 4-6 mm long in anthesis, in
fruit liecoming 9-11 mm long, setose, and stri-
gose; corolla white, the tube 3-5 mm long, con-
stricted above the ovary by the conspicuous ring
of crests, fornicvs veilow. emarginate. 0.5 mm
long, limb 7-9 mm briKid; nutlets ovate, 5-6 mm
long, .3..5-4.5 mm wide, papery, with a broad,
winged iiuu-gin. dorsal surface muricate, and in-
conspicuously rugulose or tuberculate, ventral
surface smooth or nearlv so. sc;u' straight, mir-
row, slightly open, elevated margin lacking; style
excveding mature fruit 1-2 mm. Sandy to gravel-
ly soils in the Transition Zone. Central Utah,
south throuirh most of .^ri/oiia in the moimtain-

ous areas, west to Nye County, Nevada. June-
September. Fig. 67.

Giirfield Co., east side ot Mount Ellen, Henry
Mountains, W.D. Stanton .516 (UT); Piute Co., 10 miles
west of junetion along hwy 1.53, L.C. Higgins 1785
iBRY); Sevier Co., Bowery Camp, Fish Lake, B. Ma-
guire 19890 (UTC); Fish 'Lake, Bower\' Creek Camp-
ground, L.C. Higgins 1117 (BUY); Fish Lake, Main
Lodge. L.C. Higgins 1125 (BRY); Washington Co.,
about 6 miles west of the town of Pine Vallev, L.C.
Higgins 1440 (BRY).

51. Cnjpidixthu strict (Osterh.) Payson, Ann.
Mo. Bot. Card. 14:264. 1927. Type: Some
distance south of the Yampa or Bear River
along the Victory highw;iy, Moffatt County,
Colonido, G. E.'Osterhout 6195.
Oreocanja striata Osterh. Bull. Torrey Club
50:217. 1923.

Strict perennial, 1-3.9 dm tall; stems 1-several,
0.4-2 dm long, strigose and conspicuously
setose-hispid; leaves mostly basal, reduced up-
ward, obhmceolate, acute, 2-7 cm long, 0.4-0.9
cm wide, retrorsely strigose and spreading se-
tose-hispid, pustulate; inflorescence n;irrow, in-
terrupted below the terminal cluster, 0.5-2 dm
long, setose-hispid, foliar bracts conspicuous,
especially near the base; calyx segments lanceo-
late. 4-6 mm long in ;uithesis, in fniit becoming
7-9 mm long, setose-hispid; corolla white, the

UTAH

County Lines and
river drainages

I'il^. f^7, Cn/ptdnthd -\ff(>sissitn(i

l$ioLOGif Ai, Sp.HiES, Vol. 16, No. 3

The BoR.\r.iNACE.\E of Utah

51

tube 3-4 mm long, cre.sts at base of tube con-
spicuous, fomices yellow, rounded, papillose,
limb 7-10 mm wide; nudets lanceolate to ellip-
tic, 3-3.5 mm long, 1.5-2 mm wide, usually all
4 maturing, margins in contact, knifelike, dorsal
surface with definite transverse ridges, iilso
somewhat tuberculate or nearly smooth, ventral
surface smooth or nearly so, scar open, very
narrowlv linear, elevated margin lacking; style
exceeding matiu^e fruit 1-1.5 mm. Growing on
clay or shale in the Transition Zone. South-
western Wvoming, northwestern Colorado, and
northeastern Utah. June-August. Fig. 68.

UTAH

County Lines and
river drainages

O JO

Fig. 68. Cryptantha strictci

Daggett Co., Flaming Gorge, L. Williams 489
(RM); Summit Co.. about 10 miles south of Lonetrce,
Wyoming, Welsh & Moore 6714 (BRY); Uintah Co., 2
miles north of Brush Creek on the Manila-Vemal liigh-
way, D. Ahvood 1609 (BRY); .3.6 miles north of Bmsh
Creek along the Manila-Vernal highway, L.C. Higgins
1869 (BRY): 2.5 miles north of Brush Creek, L.C. Hig-
gins 1874 (BRY).

52.

Cryptantha tenuis (Eastw. ) Payson, Ann.

Mo. Bot. Card. 14:327. 1927. Type: Court

House Wash, Grand County, Utah, Alice

Eastwood s.n.

Oreocania tennis Eastw. Bull. Torrey Club
30:244. 1903.
Caespitose perennials, 1.3-2.5 dm tall; stems
ider, 1-many, 0.8-1.2 dm long, strigose and

weakly spreading setose; leaves linear-spatulate,
mostly basal, obtuse, 2-5 cm long, 0.3-0.6 cm
wide, dorsal surface strigose and weakly spread-
ing setose, conspicuously pustulate, ventral sur-
face unifonnly strigose and without pustules; in-
florescence narrow, interrupted, 0.6-1.4 dm long,
weaklv setose, foliar bracts inconspicuous; calyx
segments linear-lanceolate, in anthesis 4.5-6 mm
long, in fruit becoming 7-9 mm long, white
setose; corolla white, the tube 5.5-7 mm long,
crests at base of tube lacking or sometimes evi-
dent, fomices yellow, broad, emarginate, papil-
lose, about 0.5 mm long, limb campanulate, 5-8
mm wide; nutlets lanceolate, 3-4 mm long, 1.8-2
mm wide, all 4 nutlets usually maturing, margin
acute to somewhat obtuse, nearly in contact,
dorsal surface carinate, sharply and deeply ru-
gose, ventral surface i-ugose, scar open, constrict-
ed above the base, and with an elevated mar-
gin; style exceeding mature fruit 3-4 mm. Sandy
or sandy-loam soils in the Upper Sonoran Zone.
Southeastern Utah in Emery, Grand, Wayne, and
San Juan counties. April-July. Fig. 69.

County Lines and
river drainages

i-

slende

Fig. 69. Cryptantha tenuis

Emery Co., about .5 miles southwest of Temple
Mountain,' L.C. Higgins 1326 (BRY); ;ibout 26 miles
southwest of Green River along hwy 24, Higgins & Re-
veal 1281 (BRY); Grand Co., lO' miles northwest of
Moab, B.F. Harrison ,5967 (BRY); Headquarters, Arches
Natl. Monument, Welsh & Moore 2801 (BRY); 1 mile
west of Delicate Arch, B.F. Harrison 112.58 (BRY);
White C:invon. B.F. Harrison 11.576 (BRY); Fisher

52

ISmcHAM Young Umvehsetv SciENCii Bulletin

Towers, Welsh & Atwood 9961 (BRY); Castle Valley,
Welsh & Atwood 9950 (BRY); northeast of Moab, just
l.elow Salt Wash, A. Cronquist 8978 (UTC); 1 mile
east of Moab bridge, Bryan & Moab School s.n. (UTC);
San Juan Co., 14 miles southeast of Moab along power
transmission line, L.C. Higgins 3547 (BRY, WTSU); 2
miles west of Hite, Welsh, Atwood & Higgins 8952
(BRY); Fry Canyon, Welsh, Atwood & Higgins 8951
( BRY ) ; about 2 miles west of Dugout Rock, Welsh, At-
wood & Higgins 8883 (BRY); 1 mile soutlieast of Natu-
ral Bridges Natl. Monument visiting center, D. Atwood
1541 (BRY); north slope of divide into Monument Val-
ley, A.H. Holmgren 3237 (UTC); 10 miles east of Hite.
A. Cronquist 9035 (UTC); about 12 miles .south of
Moab, L.C. Higgins 1000 (BRY); 19 miles west of Fry
Canyon Post Office, L.C. Higgins .537 (BRY); 2 miles
up road to Dead Horse Point, R.K. Vickery 751 (UT);
\Vayne Co., about 1 mile east of Hanksville, L.C. Hig-
gins 1334 (BRY); Three Canyon, along road about 25
miles south of Creen River along county road, Welsh
& At\vood 9895 (BRY).

53. Cryptantha virginensis (Jones) Payson,
Ann. Mo. Bot. Card. 14:274. 1927. Type;
Laverkin, Washington County, Utah, M. E.
Jones 5195A.
Krynitzkia glornerata var. virginensis Jones

Contr. West. Bot 13:5. 1910.
Oreocarya virginensis (Jones) Macbr. Proc.

Amer. Acad. 51:547. 1916.

Biennial, 1.5-3.5 (4) dm tall; stems 1-several,
arising from a stout taproot, 0.3-0.6 dm long,
setose-hirsute; leaves oblanceolate to spatidate,
obtuse, 3-10 (12) cm long, 0.5-1.5 cm wide, dor-
sal surface sparsely setose, pustulate, also with
some fine tangled hair Ixmeath, ventral surface
subtomentose and weakly appressed setose, with
only a few pustulate hairs; inflorescence a broad
thyrsus with the individual cymes much elon-
gating, 0.5-3 dm long, foliar bracts conspicuous,
2-4 cm long; calyx segments linear-lanceolate,
in anthesis 3-4 mm long, in fruit becoming 7-11
mm long, hirsute; corolla white, the tube 3-4
mm long, crests at base of tube conspicuous,
fomices yellow, emarginate, papillose, about 1
mm long, limb 7-9 mm broad; nutlets ovate,
3.3-4.5 mm long, 2.4-2.6 mm wide, usually only
1-2 nutlets maturing, margins in contact, acute,
dorsal surface with a distinct ridge, the surface
tuberculate and usually nigulose, ventral surface
very uneven with indetenninate rugae and tu-
bercles, scar open and triangular with ;in ele-
vated margin; style exceeding mature fruit 1-
1.5 mm. Usually growing on clay or clay-loam
soils in the Lower Sonoran Zone. Southwestern
Utah, southern Nevada, northwestern Arizona,
and soiitheastem (Jidifomia. March- July. Fig. 70.

Washington Co., ;ihout 5 miles southwest of St
George, L.C. Higgins 1243 (BRY); Beavcrdam Moun-
tains, L.C. Higgins 12.32 (BRY); 11 miles e;ist of St.
George. F.W. Could 1.580 fBRY, DIX); north of Gun-

lock, P. Nyberg s.n. ( DIX ) ; 4 miles west of St. George,
A. Terril s.n. (DIX); near the Shivwitt Indian Reserva-
tion, E, Hawkins s.n. (DIX); Beaverdam Mountains,
R.W. Christian 86.3 (UT); 8 miles north of Santa Clara,
A.H. Holmgren 8224 (UTC); Virgin River above Hot
Mineral Springs, Laverkin, Utah, B. Maguirc 4470
(UTC); Jackson W;ish. Beaverdam Mountains, D.H.
Nish .36 (UTC); about 4 miles west of the Shivwitt In-
dian Reservation, L.C. Higgins 4154 (BRY, WTSU).

.54. Cryptantha wetheriUii (Eastw.) Payson,
Ann. Mo. Bot. Card. 14:324. 1927. Type:
Court House Wash near Moab, Grand
Countv, Utah, Alic-e Eastwood s.n.
Krynitzkia glomeratu var. acuta Jones, Zoe

2:250. 1891.
Oreocarija wetheriUii Eastw. Bull. Torrey

Club .30:242. 1903.

Biennial or short-lived perennials, 1-3.5 (4)
dm tall; stems 1-6, 0.5-0.8 dm long, branched
from the base with 1 stout and usually several
low slender ascending stems; leaves clustered
at the base, gradually reduced upward, spatu-
late to broadly oblanceolate, the apices obtuse
to rounded, 2.5-5 cm long, 0.7-1.6 cm wide, stri-
gose and appressed setose, dorsal surface con-
spicuoiLslv pustulate, ventral surface with few
or no pustules; inflorescence becoming broad in
;ige due to the elongation of the cymes, 0.6-3
dm long, calyx segments lanceolate, in anthesis
5-7 mm long, in fruit becoming 7-1.3 mm long.

County Lines and
river drainages

Fig. 70. Cn/iitiiiilhii i irf^/iic/i.viv

Biological Sekies, Vol. 16, No. 3 The Bohaginaceak oi- Utah

53

white setose; corolla white, the tube 7-10 mm
long, crests at base oi tube lacking, fomices
light-yellow, emarginate, papillose, about 1 mm
long, limb 6-13 mm wide; nutlets lanceolate, or
o\'ate-lanceolate, 3.5-4 mm long, 2-2.5 mm wide,
usually all 4 maturing, margins acute, in contact,
dorsal surface distinctly tuberculate and often
rugulose as well, scar open, linear, surrounded
bv a slightly elevated margin; stvle exceeding
mature fruit 3-5 mm. Usually growing on heavy
clay soils associated with Atriplex. East central
Utah in Grand, Carbon, Emery, Wayne, and
Garfield counties. April-June. Fig. 71.

UTAH

County Lines and
river drainages

L, 1 1

#^ s5 io

Fig. 71. Cn/ptantlui ucthcriUii

Carbon Co., Price. W.P. Cottam .s.n. (UT); Wel-
lington, VV.P. Cottam 2031 (BRY); 2 miles south of
Price, B. Maguire 18264 (UTC); Emery Co., 9 miles
north of Green River, B. Maguire 18229 (UTC); Green
River, W.P. Cottam 2073 (BRY); Big Springs Wash,
25 miles south of the Carbon County line. Welsh &
Moore 2786 (BRY); Carbon-Emery County line, Hig-
gins & Welsh 1042 (BRY); 7 miles south of Hunting-
ton, 1..C. Higgins 1017 (BRY); Gunnison Butte, O.S.
Walsh 29 (UT); 6 miles north of Woodside, B.F. Har-
rison 10412 (BRY); along hwy 24, .3-6 miles north of
,San Raphael River bridge. G.L.' Pyrah 21 (BRY. UTC);
10 miles .south of Woodside, J. Brotherson 90 (BRY);
7 miles west of Lawrence, D. ."Ktwood 18-39 (BRY);
Garfield Co.. Henry Mountains, W.D. Stanton s.n.
(BRY); Grand Co., 4 miles south of Crescent Junction,
Welsh & Higgins 8874 (BRY); 10 miles east of Green
River. C.A. Hanson 64 (BRY); 10 miles east of Green
River. D.E. Bright 26 (BRY): about 6 miles east of
Thompson along hwy 50-6. L.C. Higgins 1476 (BRY).

7. Ci/noglosswn (Touni. ) L.

Ctjnoglosstim (Toum.) L., Sp. PI. 134. 1753, and
Gen. PL 65. 1754.

Biennial or perennial or rarely annuals; leaves
alternate, the basal ones long petioled; racemes
elongating, usually without bracts, or rarely
bracted at base; calyx cut to beyond the middle,
somewhat accrescent, segments often spreading
or reflexed in fruit; corolla cylindrical or funnel-
fonn, the tube short, lobes broad, spreading, im-
bricate, throat with trapezifomi oblong or subu-
late appendages; stamens included; filaments
short, anthers oblong or elliptic; ovules 4; nut-
lets 4, equally divergent, depressed ovoid or or-
bicular, glochidiate, back flat or convex, fre-
quently with an elevated margin, attached by
a small or large medial to apical scar to a con-
vex or pyramidal gynobase and frequently with
a free subulate prolongation decurrent on the
short entire style.

Type Species: Cynoglossum officinale L. A
cosmopolitan genus of about 50 species.

1. Cynoglossum officinale L. Sp. PI. 134. 1753.
Type: Europe.

Biennial, villous-tomentose throughout; stems
stout, erect, leafy to the top, 4-5 dm high; lower
leaves oblong to oblong-lanceolate, slender peti-
olate, 15-30 cm long, 2-7 cm vyide; upper leaves
lanceolate, acute or acuminate, sessile or the
upper mostly clasping; racemes several to many,
simple or branched, sparingly bracted or bract-
less; much elongating in fruit; pedicels 5-12 mm
long; calyx segments ovate-lanceolate, obtuse to
acutish, 5-7 mm long in fruit, corolla reddish-
purple, the broad tube 3-5 mm long, the limb
6-8 mm broad; nudets ascending on the pyra-
midal gynobase, about 6 mm high, flattish on
the upper surface and margined, splitting away
from the gynobase at maturity but hanging at-
tached to the subulate style. Native to Europe
and Asia but widely distributed over North
America. May-July. Fig. 72.

Bo.x Elder Co., Perry Canyon, Cottam. Allen &
Rowland 16231 (UT); Blue Creek, W.P. Cottam 2838
(BRY); Cache Co., Twin Creek drainage of Cache
Natl. Forest, CO. Baker 125 (UTC); fields and waste
places. N.H. Holmgren 40 (UTC); Logan, B. Maguire
.3716 (UTC); Paradise-Hyrum road, C.P. Smith 1744
(UTC); Logan, D. Hobson 1,3770 (UTC); 2 miles west
of Logan, B. Maguire 3717 (UTC); Duchesne Co., 18
miles north of Duchesne, J. Brotherson 829 (BRY);
[uab Co.. 8 miles west of sand dune on the road to
Calico. ]. Anderson 842 (BRY); Mount Nebo, Cottam
& Anderson 15105 (UT); Mill.ard Co., 9 miles east of
Fillmore, G. Worthen 348 (UT); Confusion Range,
21.8 miles east of the Utah-Nevada line, J. Gentry 2006
(BRY. UTC); Salt Lake Co., Red Butte Canyon, G.
Swanson 288 (UT); Emigration Canyon, E.B. Robinson

54

BmcHAM Young Uni\ tusixv Science Bulletin

County Lines and
river drainages

i-

l''ig. 72, Cyiioglossuni officinale

s.n. (UT); Sanpete Co., Ephraim Canyon, A.H. Bamum
s.n. (DIX); 4 miles east of Ephraim, R. Stevens 30
(BRV); 5 miles up Manti Canyon, A. CoUotzi 511
(UTC); Tooele Co., south of Vernon, R.H. Foster 22
(BRY); Utah Co., Timp Haven, K. Shaw 95 (BRY);
2 miles east of Thistle, J. Clentry 2001 (UTC); VVa.sateh
Co., 5.7 miles east of Heber, L. Arnow 676 (UT); 5
miles southwest of Heber, L.C. Higgins 1863 (BRY);
Washington Co., no locality, J-W. Harrison s.n. (BRY);
Weber Co., Weber River, W.S.F. 755 (UT); Ogden, on
south Harrison Blvd., A. Collotzi 51 (UTC).

8. Echium L.

Echmm L. Sp. PI. 139. 1753, and Gen. PI. 68.
1754.

Biennial or pos.sibly perennial, hispid, herba-
ceous plants; leaves alternate, entire; flowers
blue to violet-purple, in leafy bracted scorpioid,
spikelike racemes; calyx 5-parted; corolla tiibu-
l;ir-funnclform, irregular, usually 5-lolx;d, the
throat not appendaged; stamens ime(jual, at least
the longer ones exserted on long filaments: ovary
4-lolx'd, these separating in fruit; s-tvle 2-cleft at
api'X; nutlets erect, rugose, attaclied bv their
bases to a flat gynobase, the scar flat or some-
what concave, not leaving a pit.

Type Species: Echium italicum L.

1. Echium vul<iare L. Sp. PI. 139. 1753.

Stems 3-9 dm tall, erect, solitary or occa-
sionally several, finely hispid-villous as well as

shaggy coarse-hirsute; leaves appressed-hispid
and along the margin and midrib .somewhat
hirsute, with a strong midrib but very obscure
or absent veins; lower leaves 8-16 cm long, ob-
lanceolate, broadly stalked, forming a rosette
which withers away at anthesis; cauline leaves
reduced up the stem, the middle ones linear-
lanceolate, 3-9 cm long, contracted to a rounded
sessile base; racemes short, lateral, dispo.sed in
a long narrow thyrse or open panicle; corolla
bright blue, rarely rose or white, pubescent and
sparsely setose, rather finn in texture, 10-15 mm
long, the tube about ecjualing the calyx; stamens
very unecjual, the two pair slightly unequal but
both surpassing the lower corolla lobe, the odd
stamen included; nutlets alx>ut 2 mm long, erect,
rugose. Native of Europe, introduced into east-
em United States where it has spread westward
to the Rocky Mountaias. Jime-August. Fig. 73.

County Lines and
river drainages

i-

I'ig. 7.3. Eriuiim v<il'^aTc

Summit Co., Echo Canyon, C.H. Field 37 (UTC);
Echo C;invi)n, 2 miles above town of Echo on old road,
|.H. Moore s.n. (UTC); V.vhu Canvon, S. Flowers s.n.

(UT).

9. Eri.trichitini Sihnid.

Eritrirhiuin Schrad. in (iaud. Fl. Helv. 2:57.
1828.

Low depressed cushion-Iikt- perennials, with
the short stems densely clothed with small often

Biological Series, Vol. 16, No. .3

The Boracinaceae of Utah

55

imbricate leaves; flowers few in a racemelike
cluster tenninating the slender flowering stem;
calyx-lobes ascending, linear; corolla blue, fun-
nelfomi, with a .short tube; nutlets obliquely at-
tached to the conical gynobase, smooth, the apex
oblicjuely truncate, with a distinct, entire or
toothed margin.

Type Species: Eritricliium nantim Schrad.

1. Eritriclutim iwmiin (Vill.) Schrad. var. elon-
gattnn (Rydb. ) Cronq. Vase. Pis. Pacif. N.
W. 4:20.3. 1959.
Eritrichium urgcntcum Wight, Bull. Torrey

Club 29:411. 1902.
EritricJiutm clongatum Wight, Bull. Torrey

Club 29:408. 1902.
Eritrichium elongatum var. argenteum I. M.

Johnst. Conti-. Gray Herb. 70:53. 1924.
Oreocanja pulvinata A. Nels. Bot. Gaz.

40:63. 1905.

Plants villous often silvery looking, forming
a tuft about 2-4 cm tall (not counting the
flowering branches); leaves closely overlapping,
5-10 mm long, narrowly ovate to oblong or ob-
lanceolate, 1.5-2 mm broad, acute or obtuse,
pilose, especially on the margins and tips, with
long white hairs; flower cluster compact when
sessile among the leaves or sometimes raceme-
like when borne on a leafy flowering branch up
to T cm long; calyx-lobes linear, 1.5-3 mm long;
corolla tube e([ualing the calyx-lobes, limb vari-
able in size (1) 4-5 (7) mm broad, bright blue
(rarely white), crests in the throat pubenilent;
nutlets smooth, with an entire margin to the
trimcated oblique portion, rarely with a few ob-
scured teeth. Rocky ridges in high alpine areas,
10.000 to 13,000 feet. Oregon, ' east to Idaho,
Montana, Wyoming, Utah, and Colorado. July-
September. Fig. 74.

Bo.\ Elder Co., Raft River Range, W.P. Cottani
2849 (BRV, UT); Raft River Mountains. S.J. Preece 640
(UT); Duchesne Co.. Uintah Mountains, l.R. Murdock
.554 (BRY); Summit of Mt. Agazziz, W.P.Cottam 3718
(UT); Summit Co., Upper Henrys Fork, north of Lake
Blanchard. B. Maguire 14346 (UTC); about Henrys
Fork Lake, B. Maguire 14385 (UTC).

10. Hackelia Opiz in Bercht.

Uackelia Opiz in Bercht. Fl. Boehm. 2 pt. 2:146.
1839.

Coarse bienni;il or perennial or rarely annual
Iicrbs; leaves alternate, broad and veiny; flowers
in naked or inconspicuous bracted racemes pan-
iculately disposed; pedicels slender, recurving in
truit; calyx cut to the base into .spreading ovate

to oblong or lanceolate lobes; corolla white or
blue, with a short or elongate tube, lobes round-
ed, imbricate; throat with trapeziform intruded
appendages; stamens included, affixed at middle
of tube; filaments slender, short; anthers oblong
to elliptic; style slender, scarcely if at all surpas-
sing the nutlets; stigma capitate; ovules 4; nut-
lets 4, erect, ovate, affixed ventrally to the pyra-
midal gynobase by a broad medial or submedial
areola, margin with subulate glochidiate append-
ages which are frequently confluent at the
base, back smooth or with glochidiate append-
ages.

Type Species: Hackelia deflexa (Wahl. )
Opiz (Myosotis deflexa Wahl.)

I. Surface of nutlet more or less ridged but without
prickles; branches many flowered 1. H. floribunda

1. Surface of nutlet more or less prickly (may be
ridged ) ; branches fewer and with fewer flowers

^ (-2)

2. Nutlets broadly ovate; basal leaves few, stems
leafy above, not conspicuously reduced in size;
corolla blue 2. H. jessicae

2. Nutlets narrowly ovate; basal leaves many, stem

leaves few and reduced in size; corolla white

3.//. ixilcm-

1. Hackelia floribunda (Lehm. ) Johnst. Contr.
Gray Herb. 68:46. 1923. Type: "Lake Penta-
guishene to the Rocky Mountains, Drum-

County Lines and
river drainages

Fig. 74. Eritrichium nanum

56

Hiii(;iiAM Vdi'nc; U.\i\ i-usii'i Scmk.nck Bulletin

mond" according to Lelmiarui in Hooker

Fl. Bor. Amer. 2:84. 1838.

Echinospermiim floribundum Lelun. Stirp.

Pub. 2:24. 1830.
Lappula floribunda (Lehm.) Greene, Pit-

tonia 2:182. 1891.

Stem erect, .stout, from a short-lived peren-
nial root, 5-12 dm high, the rough pubescence
deflexed, mixed with some spreading hairs;
leaves oblanceolate to linear or oblong, hirsutu-
lous-appnessed, the basal leaves petiolate, with
spreading hairs, the stem leaves sessile above;
racemes of the inflorescence many, rather strict,
densely flowered, pedicels short, 5-7 mm long in
fruit; corolla blue, 4-7 mm broad, appendages
small, obscurelv papillate, not closing the throat;
nutlets 3-5 mm long, the face with a medial
ridge, muriculate, hiisutulous, without short
glochidiate prickles, the marginal spines much
flattened at base, distinct or somewhat confluent,
4 to 6 on each side, mostlv exceeding in width
the face of the nutlet. Brushv slopes and edge of
woods. Mostly western United States and Cana-
da, east to Ontario and Minnesota to western
Texas. June-August. Fig. 75.

Cache Co., Logan, C.P. Smith 1766b (UTC); Lo-
gan Canyon, B. Maguire 12981 (UTC); Tony Grove,
Logan Canyon, F.B.' Wiuin .370,5 (UTC); City Park.
Logan Canyon, B. Maguire 21540 (UTC); West Hodges
pasture, Logan Canyon, H.B. Passev 2 (UTC); Logan,

UTAH

County Lines and
river drainages
i 1 I

O 90 bo

ig. 75. Hackelid florihiimlu

Logan River. Northrop & Crockett s.n. (UTC); Emery
Co^ Huntington Canyon, W.P. Cottam 7437 (UT);
Garfield Co.. Henry Mountain.s, 22.9 miles south of
Hanksville, J. Gentry 1753 (UTC); east slope of Mount
Ellen, B. Maguire 19322 (UTC); 15 miles southwest
of Eseakmte. H. H;dl s.n. (UT); Henry Mountains,
W'.D. Stanton 4888 (BRV); Juab Co.. Thoms Creek,
W.P. Cottimi 3244 (BRV); Salt Lake Co., Little Cotton-
wood Canyon, W.P. Cottam 3543 (BRV); Sanpete Co.,
\ieinit\ of Ephraini, R. Olsen s.n. (IITC); 10 miles up
Ephraini Canyon, j.D. Walker s.n. (BRV); Sevier Co.,
Fish L;ike. Maguire cSc Richards 12982 (UTC); 20 miles
uest of Richfield, F. Coles 53 (BRV); Wasatch Co.,
Cascade Terraces, E.G. Davenport 23 (BRV); Utah Co.,
Mount Timpanogos, B. Maguire 3704 (UTC); Mount
Timpanogos. W.P. Cottam 1265 (BRY).

2. Hackelia jessicae (McGregor) Brand, Pflan-
zenreich 4, 252:132. 1931. Type: Halfmoon
Lake, Eldorado County, California.
La])])iil(i micranthci Eastw. Bull. Torrey

Club 30:497. 1903, not Hackelia mi-

crantha Opiz.
L(i])l)ula jessicae McGregor, Bull. Torrey

Club .37:262. 1910.
Hackelia castuoodiac I. M. Johnst. Contr.

Gray Herb. 68:47. 1923.
La])))ula floribunda var. jessicae Jepson &

Hoover in Jepson Fl. Calif. 3:307. 1943.

Stems erect or ascending from a .stout root,
sparsely to rather densely villous-hirsute; basal
leaves 8-15 cm long, the blades oblanceolate,
15-20 mm wide, narrowed to a winged petiole of
about ecjual length; upper stem leaves sessile,
lanceolate, acute, the reduced ones subtending
the lower racemes often ovate-lanceolate; ra-
cemes several in an open panicle; pedicels slen-
der, at length recurved retlexed, 5-10 mm long;
c-alyx-lobes oblong to oblong-lanceolate, 2-3 mm
long; corolla small, pale blue, 3.5-5 mm broad,
tube often whitish, 1.5-2 mm long, lobes oblong-
ovate, crests yellowish, rounded, pubeiiilent;
nutlets 4-6 mm long, marginal prickles broadly
dilated at base, about 10, distinct, often with a
shorter one in between; dorsal face broadly
ovate, usuallv flattened with a distinct median
ridge, pubeiiilent and in age more or less muri-
culate, usually with 1 or more short barbed
prickles near the center. Usuallv on moist banks
or slopes. British Columbia, Washington, and
Idaho, south to Utah, Nevada, and California.
May-August. Fig. 76.

C;Khe Co.. 5 miles up road to Tony Grove, Gen-
fr\- & Davidse 1729 (UTC); Pine C;uiv()n, B. Maguire
3101 (UTC); Logan Canyon. C.P. Smith 2227 (UTC);
Sardine Canyon, B. Maguire 12985 (UTC); 12 miles up
Smithfield Canyon, B.' Maguire 13777 (UTC); Davis
Co., Centerville Canyon, .\.C. Hull s.n. (UTC); Salt
L;ike Co.. Little Cottonwood C;niyon. R.D. Day 35
'BRV): Summit Co., north slope of Big Mountain, D.H.
Galw;iv 2382 (BRY); Ut;ih Co., divide between Ameri-

Biological Series, Vol. 16, No. .3 The BoK.\ciN.\CE.\ii oi- Utah

57

UTAH

Fig. 76. Hackelia jessicoe

can Fork and Snake Creek canyons, Welsh, Murdock
& Stocks 6348 (BRY); Mount Timpanogos, Little Basin
above falls. B. .\Iaguire 17496 (UTC); Wasatch Co.,
Daniels Canvon, B.F. Harrison 8717 (UTC); Strawberry
Valley, D.H.' Galway 8717 (BRY).

3. Hackelia patens (nutt. ) Johnst. Jouni. Am.
Arb. 16:194. 1935. Type: Near the Flathead
River. Collected by Wyeth.
RocheUa paten's Nutt. Joum. Acad. Phila.
7:44. 1834.

I.up])ula coerulcscens Rydb.
Bot. Card. 1:328. 1900.

Mem. N. Y.

Stems 2 or 3 short, deflexed hirsute, with
some hairs sprtMding, arising from a woody cau-
dex covered with old leaf bases; basal leaves
many, oblanceolate to lanceolate, stem leaves
reduced upward, pubescence rather dense, short,
appressed; branches of the inflorescence 5-10
flowered, pedicels short, elongating somewhat
in fruit; corolla whitish or light blue, usually
(juite large, appendages somewhat broader than
long, obscurely papillose; nutlets small, marginal
glochidiate prickles 3-5 on each side, a few short
prickles interspersed, the longest prickles sur-
passing the body of the nutlet; face of nutlet
with a faint median ridge, muricate and occa-
sionally bearing 1 or 2 prickles. Sandy or gravel-
ly slopes and foothills. Western Montana and
Idaho to Utah and Nevada. May-August. Fig. 77.

Maguire 12988 (UTC)
13771 (UTC); Davis Co..

Box Elder Co., Cold Water Canyon, L. Williams
636 (UTC); Bench east of Honeyville, M. Burke 3102
(UTC); Smithfield junkpile, D. Stocks 12 (BRY); Raft
River .Mountains, J. Berryman 35 (UT); Perry Canyon,
Cottam, Allen, & Rowland 16244 (UT); Beaver Co.,
Pine Canyon Pass, south ridge, B. Maguire 20962a
(UTC); Cache Co., vicinity of Green Canyon, Maguire
& Piranian 12987 (UTC); ii mile east of the fish hatch-
ery, B. Maguire 3708 (UTC); Sardine Canyon, B.
Maguire 12986 (UTC); mouth of Logan Canyon, C.P.
Smith s.n. (UTC); Intervale, Logan Canyon, Muencher
& Maguire 2422 ( UTC ) ; Providence Bench, Logan,
C.P. Smith 1593 (UTC); 4 miles up Logan Canyon, B.
Logan Bench, B. Maguire
mouth of Holbrook Canyon,
R. & B. Anderson s.n. (UTC); Muehler Park, Cottam
s.n. (UT); Garfield Co., Henry Mountains, (UT); Juab
Co., Granite Canvon, Trout Creek, Maguire & Becraft
2779 (UTC); Deep Creek, W.P. Cottam .3231 (BRY,
UT); Iron Co., 12 miles east of Cedar City, B. Ma-
guire s.n. (UTC); Kane Co., vicinity of Glendale, F.B.
Wann 36 (UTC); Millard Co., Confusion Range,
miles east of the Utah-Nevada line, J. Gentry
(UTC); Salt Lake Co., Mill Creek Canyon, L.
37 (BRY); Jordan Narrows, E. Young s.n.' (BRY); Dry
Creek Canyon, J. Ludwig 193 (UT); Citv Creek Can-
von, R.K. Vickerv 1918 (UT); Big Cottonwood Can-
yon, H.K. Harrison 68 (UT); Fort Douglas, R.K. Vick-
ery 563 (UT); Sanpete Co., 8 miles southeast of Mt.
Pleasant, L.C. Higgins 1095 (BRY); 10 miles west of
Moroni, B. Maguire 18657 (UTC); Tooele Co., Inter-
mountain Forest and Range Experiment Station, N.
Frischknecht 25 ( UTC ) ; Benmore Experiment Station,
W.H. Blackburn 39 (BRY); Utah Co., 3 miles south of
Thistle, B. Maguire s.n. (UTC); Rock Canyon, P.A.
Replogle 80 (BRY); Y Mountain. Decker & Eggertson
.347 (BRY): Wasatch Co.. 1 mile east of Heber City,

County Lines and
river drainages

i-

21.8
2003
Eyre

Fig. 77. Hackelia patens

58

BniGHAM YdUNC University Science Bulletin

B.F. Harrison 8709 (BRY); Weber Co., Ben Lomond
Trail, W. Call 17 (UT); North Ogden, A. Collotzi 77

(UTC).

The following citations are for the variety
harrisonii Gentry, which has recently been de-
scribed. Utah Co., near Tucker, Spanish Fork
Canyon, L. Mason 4323 (BRY); hogs back be-
tween right fork of Hobble Creek and Shingle
(Mill) Creek, B.F. Harrison 8323 (BRY); Span-
ish Fork Canyon, 2 miles east of Thistle, J.
Gentry 2000 (UTC); Washington Co., Pine Val-
ley, 1.5 miles up Forsvth Trail, J. Gentry 2002
(UTC).

11. Heliotropium L.

Heliotropium L. Sp. PI. 1:130. 1753, and Gen.
PI. 130. 17.54.

Annual or perennial, herbaceous or more or
less shrubby plants; leaves small to large, sessile
or petiolate; cymes unilateral and usually dis-
tinctly scorpioid, with or without bracts; corolla
white, yellow or purple, variable m fonn, throat
frecjuently pubescent inside; anthers included,
filaments extremely short; style present or ab-
sent; stigma usually frustrumlike or conic, mostly
sterile, receptive only in a band around the base;
fruit dry, at maturity breaking up into 4 single-
seeded or 2 biseminiferous nutlets; seeds with a
thin endospenn. A large genus widely distrib-
uted in the warmer parts of the world, also arid
regions.

Type species: Heliotropium europaeum L.

1. Plant not succulent, hairv. never glaucous; fniit 2-
lobed, each lobe splitting into 2 nutlets; stigma
caped by a tuft of bristles .... 1. H. convohulaceum

1. Plants very succulent, glabrous, usually glaucous;
fruit not lobed, stigma discoid, naked; perennials
(2).

2. Fniit 2.5 mm wide; corolla .5-16 broad, at most

only purplish-tinged at the throat

2a. H. curasscniciim var. ohovatum

2. Fruit 1..5-2 mm wide; corolla ■3-.5 (7) mm broad,
usuallv becoming distinctly purple or purplish ;it
the throat 2b. H. curassavicum var. ocuhitiini

1. Heliotropium convolvulaceum (Nutt. ) Gray,
Mem. Amer. Acad. Sci. 6:403. 1859. Type:
Sandv b;uiks of the .Ark;uis,is, Nutt;ill
Euploca convolvulacea Nutt. Trans. Amer.

Phil. Soc. 2, 5: 190. 18.37.
Batschia iilbiflorci Raf. New FI. No. Amer.

4:19. 18.36. Not H. alhiflorum Engler,

1924.
Euploca (ilhiflora (Raf.) Johnst. Contr.

Gray Herb. 70:53. 192.1.
Euploca <irandiflora Torr. in Emory, Notes

Mil. Reconn. 147. 1848.

Annual, 1-4 dm tall, stems at first usually
simple, but later developing elongate branches,
which are ascending or sprawling; strigose and
cinereous; leaves numerous, .strigose, pustulate,
lanceolate to ovate, 10-14 mm long, 4-15 mm
broad, apex acute, base acute to rounded, petiole
slender, .3-8 mm long, midrib evident; flowers
extra axillary, appearing to be borne along the
elongating leafy branches, bracts leaflike, nu-
merous; calvx 5-lobed, at anthesis 4-6 mm long,
becoming 6-9 mm long at maturity; lobes linear-
lanceolate or linear, unequal; pedicels at anthe-
sis 1-3 mm long, in fruit 3-5 mm long, corolla
white with a yellow throat, fragrant, opening
during the morning and evening, limb widely
funnelform, expanded, 15-22 mm broad, not
lobed pentagonal, plicate in the bud with the
sinus inflexed; tubular portion of corolla 8-11
mm long, strigose on the outside; anthers lanceo-
late, basifixed, 2-2.5 mm long; filaments about
1 mm long; ovary globose, glabrous; style slen-
der, 3-4 mm long; stigmatic head with a prom-
inent stigmatic band 0.5 mm in diameter at the
base, and this surmounted by a truncate append-
age bearing a cluster of hairs; fniit laterally
compressed, hairy, 2-lobed, 3-4 mm long, 2-2.5
mm thick, 2-2.5 mm high; at maturity first divid-
ing transversely and the lobes separating, form-
ing halves, each with a broad flat commisural
face; each half next dividing on the narrow lon-
gitudinal axis to form the asymmetrical single
seeded nutlets. In sandy soils. Nebraska and
Wyoming, south to northern Chihuahua and
Texas, and westward into Arizona and Nevada.
June-September. Fig. 78.

Emerv Co., 20 miles north of Hanksville, B. Ma-
guire 19.397 (UTC); San Raphael Desert. A. Smith s.n.
(UTC); 25 miles southwest of Creen River. W.P. Cot-
tam 17740 (UT); Carfield Co., 30 miles south of Hanks-
ville, Cronquist & Holmgren 9316 (UTC); 8 miles down
Shita Marine Creek, J.C. Pederson 16 (BRY); Hite, D.
Lindsay 2 (UT); Crand Co., .5 miles west of Moab,
Maguire & Redd 2067 (UTC): north of Moab, B.F.
Harrison 12024 (BRY); Juab Co.. near Champlin rail-
road stop, B.F. Harrison 3.52 (BRY); Millard Co., sand
dunes. W.P. Cottam .3792 (UT); San Juan Co.. Bluff
Cemcterv. Holmgren & Hansen .3442 (UTC); Y- mile
north of Chocolate Drop Butte, L.O. Wilson 155
(UTC); 5 miles east of Hite, Holmgren & Maguire
10615 (UTC); 8 miles northeast of ' Hite. Cronrpiist
& Holmgren 9490 (IITC); Bluff. W.P. Cottam 2547
(BRY, UT); Bluff, W.C. Twiss s.n. (UT); Tooele Co.,
Diigway Vallev, S. Flowers 46 (UT); Wavne Co.. .5M
miles north of Hanksville, Cron(|uist &• Holmgren 9424
(UTC): between Hanks\iIIe and Hite, AH. Holmgren
7799 (UTC); 10 miles north of Hanksville. S. Flowers
.57 (UT); W;ishington Co., 2.5 miles west of Toquer-
ville, Maguire &• Piranian 12316 (UTC); 2 miles south
of Anderson's Ranch Maguire & Richards 1.5927 (UTC);
Santa Clara. W.P. Cott.im 1519 (BRY): Leeds, W.P.
Cottam .5391 (UT): Cunlock, W.P. Cott;im .5718 (UT);
Santa Clara Bench, W.P. Cottam 127.52 (UT); Zion
Natl. Park, W.P. Cottam (UT).

RioLOGiCAL Series, Vol. 16, No. .3 The BoH.'Vf;iNACE.'^E ok Utah

59

County Lines and
river drainages
I I J

Hcliotropiuin comDliulacciiiii

2. Heliotwphim curassavicum L. Sp. Pi. 130.
1753.

Annual or short-lived perennial, fleshy, glau-
cous, glabrous throughout, stems diffusely
branched, 1-6 dm long; leaves succulent, vary-
ing from linear to obovate, but commonly spath-
ulate, 1-4 cm long, obtuse, narrow to a thick
petiole; spike mostly in pairs, sometimes 3-5,
often 6-12 cm long; calyx-segments ovate-lanceo-
late, acute, 2-3 mm long; corolla 3-5 mm long,
white with a violet-purple eye on the throat;
stigma glabrous; stamens included, the anthers
subsessile; fruit subglobose, at length separat-
ing into 4 nutlets.

2A. Heliotropium curassavicum var. obovatum
A. DC. Prod. 9:5.38. 1845. Type: Colum-
bia River, probably near the Blue Moun-
tains, Oregon

Ueliotrophnn sjHithuldtum Rydb. Bull. Tor-
rcy Club 30:262. 1903.

Leaves spathulate to obovate; corolla white
or slightly tinged with purple, 6-8 mm long, the
limb about as broad; nutlets 2.5-3 mm long. Al-
kaline or saline places. Ea.stem Washington,
eastern Oresron, and northwestern Nevada, east
to the Rocky Mountains. June-October. Fig. 79.

Bo.\ Elder Co., Bear River Refuge, D. Hobson
14839 (UTC); Kmidsen Marsh, Jen.sen & Dargan s.n.

(UTC); Cache Co., Pehcan Pond, J. Thieret 60 (UTC)
Daggett Co., Linwood Bridge, S. Flowers 127 (UT)
Garfield Co., Bakers Ranch, B. Markham B-9 (UTC)
Millard Co., between Delta and Hinckley, Maguire &
Becraft 3941 (UTC); Salt Lake Co.. Salt Lake City,
W.P. Cottam s.n. (UT); Tooele Co., Smelter Bench,
C.P. Smith 1893 (UTC); Utah Co., near Provo, W.P.
Cottam 3171 (UT); Uintah Co., 8.5 miles north of
Ouray, N. Folks 2.35 (UTC).

2B. Heliotropium curassavicum var. oculatum
(Heller) Johnst. ex Tidestr. Proc. Biol. Soc.
Wash. 48:42. 1935. Type: Sand along the
Russian River near Healdsburg, Sonoma
County, California.
Heliotropium oculatum Heller, Muhlenber-

gia 1:58. 194.
Heliotropium spathulatum subsp. ocula-
tum Ewan, Bull. So. Calif. Acad. 4:56.
1942.

Middle cauline leaves oblanceolate to spath-
ulate, smaller lower and upper ones commonly
oblong and acutish; corolla limb about 4 mm
wide, the lobes white or bluish and the throat
with a violet-purple eye. Southwestern Utah,
south to Baja California, and east to New Mexi-
co and Texas. March-September. Fig. 80.

Washington Co., Arizona Strip, B. McAllister s.n.
(UTC); St. George, F.B. Wann .32 (UTC); Santa Clara,
W.P. Cottam 6900 (UT); 2 miles southeast of Santa
Clara, R. Christian 9.34 (UT); St. George, W.P. Cot-

UTAH

County Lines and
river drainages

Hcliotropiiiin ciirdsKnirutii

ohovatum

60

BlUCIlAM Vol'NC UnIVFRSITV SCIENCE BULLETIN

UTAH

County Lines and
river drainages

o" so

Fig. 80. llcliutrojtiuin cnrussuiicum \ar. oruhituin

tarn 3386 (UT); Beaverdam Wash, K.R. Kelson s.n.
(UT); Terry's Raneh, S.L. Welsh 5270 (BRV); Terry's
Ranch, in the bottom of Beaverdam Wash, L.C. Hig-
gins 513 (BRY).

12. Lappula Moench
Lappula Moench, Metli. 416. 1794.

Annual herbs; leaves alternate, usually nar-
row, firm and veinless; flowers small, blue or
white, on usually erect pedicels or rarely sub-
sessile, in bracted racemes; caly.x 5-partcd into
spreading lanceolate lobes; corolla with a rather
short tube; lobes rounded, ascending imbricate;
throat closed by intruded appendages; stamens
affixed in the tube, included; filaments slender,
short; ;inthers oblong, obtuse; style short, sur-
mounting the subulate-columnar gynobase, com-
monly surpassing the mature nutlets; stigma sub-
capitate; ovules 4; nutlets 4, erect, smooth or
vernicose, narrowly but firmly attached to the
gynoba.sc along the length of the well-developed
ventral keel, back angulate or margined by a
single or double row of prickles which by con-
fluence frequently form a winglike or cupulate
border.

Type species: Mtjosotis lappula L. {Lappula
echinata Gilib. )

1. Nutlets with marginal prickles in at least 2
rows 1. L. echinata

1. Nutlets with the marginal prickles definitely in a

single row ( 2 ) .

2. Marginal prickles distinct to the base or nearly so,

not confluent to form a cupulate structure

2A. L. occidcntalis var. occidentalis

2. Marginal prickles confluent, forming a conspicuous
smooth cupulate structure on the back of some or
all the nutlets .... 23. L. occidentalis var. cupulata

1. Lappula echuuita Cilib. FI. Lith. 1:25. 1781.

Type: Europe.

Mtjosotis lappula L. Sp. PI. 131. 1753.
Lappula imfosotus Moench Meth. 417. 1794.
Lappula lappula Karst. Deutsch. Fl. 979.

1880-83.
Echinafpennum lappula Lehm. Asperif . 121.

1818.

Annual, with erect, simple to freely branched
stems 1.5-8 dm tall, villous-hirsute with upward-
ly more or less appressed hairs; lower leaves
linear to linear-lanceolate or oblong, acute to
obtuse, narrowed to a sessile base, closely as-
cending, 2-5 c-m long, roughly pubescent like
the stem, passing above into the linear or lanceo-
late bracts of the usually numerous racemes;
pedicels 1-3 mm long; calyx-lobes broadly linear,
appressed bristly, in fruit .spreading, 2.5-3 mm
long; corolla bright blue, the limb 2-4 mm
broad, the tube surpassing the calyx; nutlets 3-4
mm long, sharply verrucose or muricate dorsal-
ly, with 2 marginal rows of long slender bristles
not confluent at the base, these sometimes ir-
regularly distributed over the back. Dry plains,
hillsides, and waste places, also cultivated
ground. Native of Eurasia. Widespread as a
weed in the United States and Canada. June-
August. Fig. 81.

Summit Co., John Lambert Ranch, P.B. Lister 12

(UT).

2. Lappula occidentalis (Wats.) Greene, Pit-

tonia 4:97. 1899. Not Lap])ula redowskii of

European authors.

Echinosjicnnum redowskii var. occidentalis

S. Wats. IJot. King Expl. 246. 1871.
Lappula redowskii occidentalis Rydb. Contr.

U. S. Natl. Herb. 3:170. 1895.

.Animal, the stems simple or few branched at
bas(^ ;md erect or sometimes diffiLse, 15-35 cm
tall, herbag(> more or less canescent with a stri-
gose and also a villous pubescence; leaves nar-
rowly linear to narrowly lanceolate or the lower
n;irT(>wly oblanceolate, 1-3 cm long; flowers in
the axils of the small foliaceous bracts forming
open and at length elongated tenninal racemes;
pedicels 1-2 mm long; calyx-segments narrowly

BioLtx:ic;AL Series, \'ol. 16, No. 3

The Bohaginaceae oe Utah

61

County Lines and
river drainages

i JO it"

Fig. 81. Lappula ccliinuta

lanceolate, erect, or but little spreading in fruit,
a little shorter than the corolla-tube, corolla
blue, 3-4 mm long, conspicuously crested on the
throat; nutlets 2-2.5 mm long, bordered by a
single row of barbed prickles, the prickles dis-
tinct at base or joined to fonu a cupulate mar-
gin, the dorsal area of nutlets distinctly tuber-
culate.

2A. Lappula occidentalis (Wats.) Greene, var.
occidentalis

Distinguished by the marginal prickles which
are not united at the base, or do not form a cu-
pulate structure of any kind. Dry hillsides and
vallevs. British Columbia southward to Mexico
and eastward to the Dakotas and Texas. April-
July. Fig. 82.

Bo.x Elder Co., Hardup, G.F. Knowlton 332 (UTC);
Bear River Canyon, C.P. Smith 1678a (UTC); Cotton-
wood CroN-e, M. Burke 3104 (UTC); Raft River Moun-
tains S.J. Preece 7.32 (UT); Beaver Co., Wah Wah
Range, Stahmann & Hatchings 40 (UT); Little Reser-
voir, Beaver Canyon, R.G. Wamock s.n. ( UT ) ; Wah
Wall Mountains, F. Coles 22 (BRY); Cache Co., Pine
Canyon, B. Maguire 3105 (UTC); Logan, Muenscher &
Maguire 2421 (UTC); intervale, Logan Canvon, B.
M;iguire .3713 (UTC); above Pelican Pond, J. Thieret
17 (UTC); Oirbon Co., 4 miles north of Price. B. Ma-
guire 18348 (UTC); Price Canvon, S. Flowers 6422
( UT ) ; .5 miles east of Price, B.F. Harrison 10262 ( BRY ) ;
road to Mounds. D. Atwood 126.5 (BRY); Daggett Co.,

vicinity of Flaming Gorge, L. Williams 478 (UTC);
12 miles .south of Manila, B.F. Harrison 7901 (BRY);
Duchesne Co., Indian Canyon, A.O. Garrett 8329 (UT);
15 miles southwest of Myton, J. Brotherson 1074
(BRY); 5 miles north of Fruitland, ]. Brotherson 503
(BRY); 6 miles south of Duchesne, B.F. Harrison 3981
(BRY); 3.5 miles southwest of Duchesne, N.H. Holm-
gren 1771 (BRY); west of Fruitland, B.F. Harrison
87.50b (BRY); Emery Co., 20 miles north of Green
River, B. Maguire 18234 (UTC); San Raphael Swell,
B. Maguire 18284 (UTC); Huntington Canyon, A.O.
Garrett""7018 (UT); Horse Canyon Junction, J.L. Peder-
son 8 ( BRY ) ; 5 miles south of the San Raphael River,
B.F. Harrison 9613 (BRY); 5 miles southwest of Tem-
ple Mountain, L.C. Higgins 1.328 (BRY); Temple
Mountain, V.P. AUman s.n. (BRY); Garfield Co., 5
miles south of Hatch, B. Maguire 18270 (UTC); Brom-
ide Peak. Henry Mountains. B.F. Harrison 74.50 (UTC);
Brvce Canvon Natl. Park, H. Buchanan 70 (UT); Can-
nonville, ].' Reveal 752 (BRY); .Mt. Ellen, Henry Moun-
tains, W.D. Stanton 4863 (BRY); Grand Co., near
Moab, B.F. Harrison .5957 (BRY); Hill Creek Ranger
Station. Vickery & Wiens 16.34 (UT); 4 miles east of
Green River. |.L. Pederson 10 (BRY); 2 miles north of
Thompson, S.L. Welsh 6898 (BRY); Iron Co., Modena,
L.N. Goodding 1010 (UT); Juab Co., Deep Creek
Mountains, Maguire & Becraft 2783 (UTC); Mt. Nebo
Ranger Station, R.K. Gierisch 274 (UTC); Trout
Creek. Maguire & Becraft 2782 (TC); north of Fuma-
role, Cottam & MacMillian 9646 (UT); Kane Co., vicin-
ity of Glendale, F.B. Wann .37 (UTC); 2 miles south-
east of Kanab, McClain s.n. (UTC); Buckskin Gulch,
S.L. Welsh .5312 (BRY); Millard Co., .37 miles west of
Delta, Maguire & Becraft 2781 (UTC); 5 miles south-
west of Desert Range E\p. Station. B. Maguire 20808
( UTC ) ; 27 miles west of Delta, Maguire & Becraft 2780
(UTC); Desert Range E.\p. StiUion, P. Plummer .30

UTAH

County Lines and
river drainages

A 30 fco

Fig. 82. iMpptiUi occidentalis \ar. nccidcntalis

62

Bjucham Vcunc Uni\eusitv Science Bulletin'

(UT); Confusion Hange. J. Gentry 2033 (BRY); north
end of Pine Valley, S. Brewster 18 (BRY); 33 miles
north of Milfoul, Welsh & Moore 6800 (BRY); Piute

(Greene) Johnst.
3:93. 1932.

Contr. Am. Arb.

C(

Higgins 1158 (BRY); 7
M.l. Morris s.n. (BRY);
D. Atwood 1376 (BRY);
R.K. Vickerv 2360 (UT)

Monroe Mountain, L.C
miles south of Puffer L;ike,
Highway 22 near milepost 3.
.Salt Lake Co., Murdock Peak

City Creek Canyon, K. Brizzee 7783 (UT); San Juan
Co., 2 miles east of .\Ie.\iean Hat, B, Maguire 16258
(UTC); 2 miles northwest of Bluff, B. Maguire 16253
(UTC); 2 miles west of Bluff, B. Maguire 13512
(UTC); 5 miles southwest of Blanding, L.C. Higgins
519 (BRY); .'Vbajo Mountains. Rvdberg & Garrett 9663
(UT); Canvonlands Natl. Park. Virginia Park, G.
Moore 319 (BRY); Chesler Park, Welsh, Moore & Can-
ter 2905 (BRY); Natural Bridges Natl. Monument, G.
Moore 134 (BRY); Sanpete Co., Sanpete-Juab Countv
line, T. Jensen 504 (UTC); Fountain Green, T.R. Stutz
9 (BRY); Tooele Co., Blaek Rock Cimyon, J.L. Re-
ve;il 215 (UTC); Benmore E.xp. Station,' W.H. Black-
hum 40 (BRY); near Stockton, L. Amow 278 (UT);
Ib;ip;ih, W.P. Cottam 3149 (BRY); Uintah Co., Dino-
saur Natl. Monument. S.L. Welsh 1966 (BRY); 5 miles
northwest of Whiterocks, J. Brotherson 559 (BRY); 5
miles south of Ouray, A.D. Youngberg 1106 (BRY);
Two Waters Creek drainage, N.H. Holmgren 1870
(BRY); Utah Co., Sp;inish Fork Canyon, L. Mason s.n.
(BRY); 7 miles north of Elberta, R. Coombs 8 (BRY);
Diamond Fork Canyon, P.A. Replogle 101 (BRY); west
of Utah Lake. W.W. Patton 117 (BRY); Sevier Co.,
Willow Creek. Duane & Jefferv s.n. (UTC); 5 miles
southeast of Sigurd, B. Maguire 18096 (UTC); Fish
Lake, B. Markham s.n. (BRY); Fish Lake, L.C. Higgins
1119 (BRY); Fish Creek Canyon, A.O. Garrett 2591
(UT); 2 miles north of Els'inore, J.L. Reyeal 7.34
(BRY); Summit Co., Burnt Fork Creek, E.E. Jen.sen
s.n. (UTC); Browns Meadow, E.M. Christensen s.n.
(BRY); Wayne Co., 3 miles from Pace Ranch. B. Nfa-
guire 18180' (UTC); Fremont Canyon north of Fniita,
B. Maguire 18114 (UTC); Parker Mountain, D.M.
Bcale 112 (UTC); 6 miles east of Torrey, B. Maguire
IS109 (UTC); Fruita. DE. Beck s.n. (BRY); Bo.x Can-
yon, V. Alexson 47 (BRY); near Torrey, W.P. Cottam
9281 (UT); edge of Bicknell Bottoms, R.K. Vickery
.593 (UT); Washington Co., Pine Valley Mountains,
Gentry & Jen.sen 2206 (UTC); 10 miles east of Hurri-
cane,'Welsh & Moore 6866 (BRY); 5 miles southwest
of St. George, L.C. Higgins 1239 (BRY); Pine Valley
Monnt;iins. W.P. Cottam 8917 (BRY); 2 miles e;ist of
Pinto, P. Plummer 7407 (UT); Wasatch Co., east rim
of Str;uvberrv Valley, I. Brotherson 694 (BRY); Daniels
Canyon, A.O. Garrett' 2843 (UT); Weber Co., Snow
Basin. A. Collotzi 79 (UTC).

2B. Lappula occklentaKs (Wut.s.) Greene var.
ctqmlata (Gray) Higgins, comb. nov.
(ba.scd upon Echinospermum redotvskii
var. cuptihitum Gray, Bot. Calif. 1:5.30.
1876).

Echinospermum redowskii var. cnpnhitum
A. Gray, Bot. Calif. 1:5.30. 1876.

Lappiila nipnlata (Gniv) Rvdb. Ru)]. Tor-
rey Club 28:31. 1901.

Lappula texana var. columhimm (A. Ncls. )
Johnst. Contr. Gray Herb. 70:50. 1924.

Lappula redoivskii var. clesertorutn

Distinguished from the typical species by
JKiving the marginal prickles confluent to about
the middle, forming ;i definite cupulate margin.
.Southeastern Washington, southward east of the
Cascade-Sierra Divide to southern California,
eastward to Montana, Wyoming, ;md New Mexi-
co. .April-August. Fig. 83.

UTAH

County Lines and
river drainages

Fig. 83. Lappula occidcntalis \ar. lupuhitii

Carbon Co., 3 miles north of Price, R. Hardy s.n.
(UT); 1 mile east of Roadside Geyser. R.K. Vickery 718
(UT); Daggett Co., 1 mile south of Manila. B. Maguire
12373 (UTC); Emery Co., 10 miles west of Green
River, B. Maguire 2056 (UTC); Grand Co., 3 miles
north of Dewey, R.K. Vickerv 238 (UT); Kane Co., 47
miles east of Kanab, A. Cron'quist 10203 (BRY. UTC);
Millard Go. Confusion Range, J. Cientry 20.33 (UTC);
W.ishington Co., St. George," W.P. Cottam s.n. (UT).

13. Lithospermunt L.

Lithospcimum L. Sp. PI. 1:132. 1753, and Gen.
PI. 64. 17.54.

.\nnu;d or perennial, herbaceous or fruticose
plants with alternate leaves; flowers white, yel-
low, or violet, in bracted racemes; calyx usually
undivided; corolla tubular or salverfonn, the
tube cvlindric;il, lobes spreading and imbricate,
the throat with intruded appendages or with
pubescent or glandul;ir areas; stamens affixed in

BuiLoGiCAL Skbies, \ol. 16, No. 3

TUF. BoHACINACIiAK OF I'XAII

63

the tube, included; filaments short, anthers ob-
long, usually with apieulate connectives; style
filiform; stignias geminate; ovules 4; nutlets 4
or rarely fewer, erect, ovoid or angular, smooth
or verrucose, affixed by a broad horizontal or
slightly obli([ue basal areola; gynobase flat or
very broadly pyramidal.

Type Species: Lithospermum officinale L.

1. Annuah flowcr.s white; nutlets den.sely tiibercu-
late and dull 1. L. arvcnsc

1. Perennial; flowers greenish to yellow, nutlets white,

smooth ( 2 ) .

2. Corolla 10 mm long or more, tube definitely ex-
ceeding the caly.v, green or pale yellow, nutlets
4-6 mm long 4. L. ruderale

2. Corolla 10 mm long or more, tube definitely ex-
ceeding the calyx, yellow (3).

3. Styles of all flowers about the same length; stamens
all bonie near the top of corolla tube; corolla bright
yellow, usually over 20 mm long, its lobes toothed
or with a fringe of hairs; later flowers cleistoga-
mous, much smaller, in fruit with recurved pedi-
cels 2. L. incisum

3. Styles of flowers of 2 lengths ( heterostyled ) ; sta-
mens borne either about at the middle or near the
top of the corolla tube; corolla deep yellow or
orange, not over 20 mm long, its lobes entire or
nearly so; smaller cleistogamous flowers absent;

root thick, containing a purple dye

.3. L. mtiltifloriim

1. Lithospermum arven.se L. Sp. Pi. 132. 1753.
Type: Europe.

Annual; stems erect, 2-7 dm tall, one to
several siiuple or sparsely branched above, hoary
strigose; leaves 2-5 cm long, 2-8 mm broad,
closely appressed-hispid, firm, veinless, with
prominent midrib, pale beneath, basal leaves
rosulate, oblanceolate, or spathulate, those of
the stem acute and lanceolate or linear; racemes
bracted, becoming loosely flowered; calyx parted
into linear-subulate lobes, hispid, mature calyx
with the erect or ascending lol>es becoming 8-13
mm long, the very short tube oblique, pale and
chartaceous, pedicels short and stout, about 1
mm long, corolla white or yellowish or even
purplish, 5-7 mm long, tubular-funnelform, the
tube glabrous within and scarcely if at all sur-
passing the calyx; corolla-throat gradually ex-
panded, without protuberances, merely pubes-
cent; corolla-lobes ascending ovate, obtuse; nut-
lehi brown, dull, roughened, tul>erculate, or
rough-wrinkled or pitted, about 3 mm long,
ovate. Grassy hillside and grain fields. Natural-
ized from Europe, widely introduced in the
United States. June-August. Fig. 84.

Cache Co.. 'A mile south of Providence, B. Maguire
216.58 (UTC); just east of Millville, C. Davidse 1026

(UTC); Providence Canyon, B. Maguire 12989 (UTC)
Davis Co., above Bountifid, R.K. Vickerv 1910 (UT)
Tooele Co., south of Tooi-le, A.F. Heaps .3907 (UT)
about 12 miles southeast of Tooele, L.C. Higgins 3345
(BRY); Salt Lake Co., Dr)' Creek Canyon, ]. Ludwig
233 (UT); 5 miles east of Salt Lake City, L. Amow
291 (UT); Fort Douglas, R.K. Vickery 561 (UT); Utah
Co., mouth of Hobble Creek Canyon, B.F. Harrison
12216 (BRY).

2. Lithospcrmum incisum Lehm. Asperif. 2:303.
1818.
Litl>o,','j)ermum angtistifolitim Michx. Fl.

Bor. Amer. 1:1.30. 1903. not Forsk 1775.
Lithospcrmum lineurifoUum Goldie, Edinb.

Phil. Joum. 6:322. 1822.
Baischia longiflora Nutt. in Pursh, Fl. Sept.

Amer. 1:132. 1814.
Lithospcrmum longiflorum (Pursh) Spreng.

Syst. 1:544. 1825.
Batsciua decttmbens Nutt. Gen. 1:114. 1818.
Lithospcrmum decumhens (Nutt.) Torr.

Ann. Lye. N.Y. 2:225. 1826.
Lithospermum crtiptanthiflorum Brand in

Fedde, Report 28:13. 1930.
Lithospermum horealc Brand in Fedde, Re-
port 28:13. 1930.
Lithospermum breviflorum Engelm. & Gray,

Joum. Host. Soc. N.H. 5:252. 1845.

Perennial plants from a thick woody root,
stems 1-5 dm tall, usually several, erect or as-

County Lines and
river drainages

i-

90 ko

Fig. 84. Lithospermum arieihse

64

BiuciiANf Young Uni\ fusitv Scienck Bulletin

ceiiding, strigose to somewhat hirsute; leaves
10-50 mm long, linear to linear-oblong, strigose;
inflorescence of terminal leafy racemes; calyx
6-10 mm long; corolla 10-30 mm long, yellow,
the tube seldom over 2.5 mm wide when
pressed, salverform, limb 9-18 mm wide, lobes
fimbriate to toothed; nutlets 3-4 mm long, white
and shining; cleistogamous flowers present, in
fruit usually with recurved pedicels; flowers
monomorphic. Dry plains and slopes. Ontario to
British Columbia, south to Illinois, Texas, Mexi-
co, and Arizona. April-July. Fig. 85.

Kig. 8.5. Litho.spi-'ninim iitcisiini

Beaver Co., Wah Wah Mountains, 4.5 miles up
Pine Canvon, B. Maguire 20942 (UTC); Pine Grove,
Wah Wail Mountains. W.P. Cottam 80.55 (UT); Bo.x
Elder Co., .3 miles south of Deweyville, A.H. Holmgren
7002 (UTC); Carl)on Co., 10 miles north of Welling-
ton on the Mvton highway, B. Maguire 18.57.3 (UTC);
Price, S. Flowers 911 (UT); road to Mounds, D. At-
wood 1263 (BRY); Daggett Co., 1 mile below Hole in
the Rock Grazing Station. A.H. Holmgren & Tillett
9519 (UTC); north slope of the Uintah Moimtains, V.B.
lUchens s.n. (UTC); Duchesne Co.. north rim of Nine
Mile Creek, N.H. Holmgren 1951 (BRY. UTC); 6 mih-s
uest of Mt. Home, J. Brotherson 875 (BRY); Emer\
Co.. San Raphael Swell. B. Maguire 18430 (UTC)';
hc;id of Cottonwood Wash, D. Atxvood 1.303 (BRY);
lie;Kl of Buckhorn Wash. B.E. Harrison 81.38 (BRY);
G;irfield Co., 17 miles southwest of Esealante, A. Cron-
(|uist 91.58 (UTC); 5 miles northeast of Ruby's Inn,
Holmgren & Nielsen 7756 (UTC); southeast of Ese;d-
;mte. I). Atwood 301 (UTC); Henry Mountains, W.P.
Cottam .5.540 (UT); Brvee Canvon Natl. Park, L. Ar-

novv 4 (UT); south of Circleville, D. Atwood 1378
(BRY); north side of Boulder Mountain, B.F. Harrison
9164 (BRY); west of Mt. Ellen near Kings Ranch, B.F.
Harrison .5540 (BRY); Bryce Canyon Natl. Park, W.S.
Boyle 1112 (BRY); Grand Co., near Thompson, S.L.
Welsh 6922 (BRY); Iron Co., Iron Mountain, P. Plum-
mer 7425 (UT); Cedar Canyon, W.P. Cottam .3931
(BRY); above Orderville, W.P. Cottam 4270 (BRY);
Kane Co., 2.5 miles north of Cottonwood Wash Spring,
Reveal, Gentry & Davidse 814 (BRY. UTC); K mile
north of Mt. Carmel Junction, Maguire & Piranian 1229.3
(UTC); 10 mdes north of Glendale, B. Maguire 18948
(UTC); 15 miles north of Orderville, B. Maguire
18873 (UTC); Juab Co., south base of Topaz Moun-
tain near Thomas, Welsh & Atwood 9675 (BRY); Mil-
lard Co.. Pavant Butte, W.P. Cottam s.n. (UT); San
juan Co., 1 mile west of Monticello, Holmgren & An-
derson s.n. (UTC); Kigalia Ranger Station, D.E. Hen-
riques s.n. (UT); 20 miles west of Blanding. B.F. Har-
rison .5896 (BRY); 10 miles south of Monticello, B.F.
Harrison 11910 (BRY); Natunil Bridges Natl. Monu-
ment, G. Moore 44 (BRY); Salt Canyon, Welsh, Moore
& Canter 2991 (BRY); Canyonlands Natl. Park, Vir-
ginia Park, G. Moore 334 (BRY); 5 miles west of
Blanding. L.C. Higgins 518 (BRY); Salt Lake Co.,
point of mountain, E.M. Christensen s.n. (BRY); San-
pete Co.. 11 miles west of Fayette, T. Jensen 573
(UTC); Sevier Co., Salina Canvon. W.P. Cottam 9194
(UT); 40 miles north of Fremont, B.F. Harrison 7.340
(BRY); Uintah Co., Willow Creek drainage. Holm-
gren & Reveal 18.5.3a (BRY, UTC); Book Cliffs, L.C.
Higgins 1.593 (BRY); Dinosaur Natl. Monument, S.L.
Welsh 86 (BRY); Washington Co., Zion Natl. Park,
Holmgren. Reveal & LaFrance 1985 (BRY, UTC):
Zion Natl. Park, W.P. Cottam 6987 (UT); Beaverdam
Mountains, L.C. Higgins 909 (BRY); Pine Valley. W.P.
Cottam 4053 (BRY)'; Wayne Co., %
Teasdale, G. Davidse 1080 (UTC)
Elatterite Basin, Bryan s.n. (UTC
area. W.P. Cottam 17774 (UT).

mile southwest of

north trail into

; Standinji Rock

3. Lithospermum imiltiflonan Torr. ex Gray,
Proc. Amer. Acad. 10:51. 1874. Type; Prob-
ably New Mexico.

Lithosjiennum co^natum Greene ex Speng-
ler, Oesterr. Bot. Zeit. 68:118. 1919.

Perennial plants from a thick woody root con-
taining a purple dye; stem 3-6 dm tall, more or
less tufted, often virgately branched above, stri-
gose-hispid; leaves 2-6 cm long, linear or linear-
lanceolate, appressed-strigose above, hirsute be-
neath, becoming smaller and bractlike near the
flowers, scarcely if at all longer than the calyx
lobes and simuhiting them; flowers racemose,
short-pcdiccled, often on .several ascending
corvmblikc branches; calyx lobes about 4-6 mm
long; corolla yellow or orange-yellow, tubular
funnelfomi, the tube ;ibout 8-13 mm long, the
lobes short, about 2 mm long, rounded, not fim-
briate; nutlets about 3 mm long, white and shin-
ing; cleistogamous flowers absent; flowers heter-
ostyled and dimorphic. Hills, canyons and
mountain slopes. Wyoming to Mexico. May-July.
Fig. 86.

BiOLOOtcAL Sehies. \'oi.. 16, No. 3

The Bohacinackae ok Utah

65

Fig. 86. Lithospcrmum multiflonim

Beaver Co., 8 miles east of Beaver, B. Maguire
19666 (UTC); Garfield Co.. 15 miles north of Esca-
lante, N.H. Holmgren 24.38 ( BRY, UTC); Bryce Can-
yon Natl Park, Snnrise Point, W.S. Boyle 1114 (BRV);
4 miles east of Widtsoe Junction, Welsh, Isley &
Moore 6496 (BRY); Red Canyon, B. Maguire 19082
( BRY ) ; north of Red Canyon Campground, D. Atwood
1889 (BRY); Brvce Canvon Natl. Park, W.P. Cottam
11454 (UT); 15miles southwest of Escalante. H. Hall
s.n. (UT); Iron Co., 10 miles east of Cedar City, Rethke
& Raadshoven 4628 (UTC); highway 14, 7.8 miles east
of [unction highway 55, L. Amow 101 (UT); Kane
Co.; 10 miles north of Order\ille, B. Maguire 18767
(UTC); 15 miles northwest of Orderville, B. Maguire
18809 (UTC); San Juan Co., Arch Canyon, W.P.
Cottam 2433 (BRY); Church Rock. W.P. Cottam 2316
(BRY); Devils Canvon between Blanding and Monti-
cello, AH. Holmgren .3492 (BRY, UTC); Elk Ridge,
Maguire & Redd 2063 (UTC); 8 miles north of Bland-
ing.' Holmgren & Hansen .3381 (UTC); Abajo Moun-
tains, W.C. Twiss s.n. (UT); Sevier Co., Fish Lake.
D.W. Lindsay s.n. (BRY); Wayne Co., 16 miles south-
east of Torrey, Cronqui.st & Holmgren 9357 (UTC).

4. Lithospciiiunn ntdcidlc Dougl. c.v. Lcliin.
Stirp. Pug. 2:28. 1830. Type: Gravelly banks
of the Columbia and Willamette river.s.
Lithospernunn ])ilosum Nutt. Joum. Acad.

Phila. 7:4.3. 1834.
LitJiospermtnn torrcyi Nutt. op. cit. 44.
LithosjK'rmum laxum Greene, Pittonia 3:263.

1898.
Litlios])crmum rudcralc var. hmceolatum A.

Nels. Bot. Gaz. 52:272. 1911.

Stem.s u.sually several from a large root, erect
or decumbent, rather stout, 2-5 dm high, simple
or branched, hirsute and somewhat hispid to
densely villous; leaves numerous, usually crowd-
ed above, mostly ascending or sometimes re-
flexed, linear-lanceolate to lanceolate, 3-8 cm
long, 2-12 mm wide, softly to rather harshly
pubescent on both sides, scabrous on the mar-
gins, flowers in the axils of the upper leaves;
pedicels stout, 1-3 mm long; calyx lobes in fruit
subulate, 7-10 mm long; corolla pale, often
greenish-yellow, 9-12 mm long, tube broad,
scarcely dilated at the throat, lobes about 3 mm
long; nutlets broadly ovoid, 5-6 mm long, usually
abruptly attenuate at the apex into a stout beak,
whitish, smooth and highly polished. Dry plains
and hillsides. British Columbia, south in the
Pacific States east of the Cascade and Sierra
Ranges, ea.st to Alberta, Montana, Wyoming,
Colorado, and Utah. April-July. Fig. 87.

Counly Lines and
river drainages

i-

Fii;. 87. I AthospcrmMm rmlrnilc

Box Elder Co., Copper Mountain, W.P. Cottam
3120 (UT); Cottonwood C;>nyon, M. Burke 3115
(UTC); south of Stanrod, Raft River Mountains, K.S.
Erdman 15.30 (BRY); Grouse Creek, K.S. Erdman 1629
(BRY); Ciiche Co., Logan, S. M. Lewis 7 (UTC); top
of Sardine Canyon, N.H. Holmgren s.n. (UTC); 3.5
miles south of Hvmm turnoff. R. Eastmond 245 (BRY);
Daggett Co., CIreen Lakes, L. Williams 601 (UTC);
Diivis Co., Mueller Park, K. Brizzee 7771 (UTC); near
Bountiful Reservoir, B.A. Anderson 52 (UTC); Iron
Co., Dixie Natl. Forest. CJ.S. Grew s.n. (UTC); Juab

66

Bkigiiam Younc Univkrsitv Science Bulletin

Co., JoluLsons Canyon, Deep Creek Mountains, W.P.
Cottam 7184 (UT); Mt. Nebo Ranger Station, R.K.
Gierisch 286 (UTC); Robinson, I.E. Diehl Dl (BRY);
Salt Lake Co., Dry Creek Canyon, J. Ludwig 20 (UT);
San Jiian Co., 1 mile west of Montieello, Holmgren &
Hansen 3384 (UTC); 3.5 miles south of Montieello, J.
Christensen s.n. (UTC); Sevier Co., summit east of
Cove Fort, D. Atwood 1.516 (BRY); Summit Co., Peoa
Cemetery, S.L. Welsh .541 (BRY); Tooele Co., Sheep-
rock Mountixins. E.M. Christensen s.n. (BRY); Uintah
Co., 5 miles north of White Rocks, A.D. Youngberg
1017 (BRY); Dino.saur Natl. Monument, S.L. Welsh
489 (BRY); Utah Co., right fork of Hobble Creek Can-
yon, L.C. Higgins 3.504 (BRY); mouth of Hobble Creek
Canvon. Condom s.n. (UT); 3 miles south of Thistle,
B. Maguire 18645 (UTC); Diamond Fork, J.W. Thomas
141 (BRY); east of Provo. B.F. Harrison 6582 (BRY);
southe;ist of Spanish Fork, E. Ni.xon 64 (BRY); east
Tintic Mountains, R. Coombs 132 (BRY); 10 miles east
of Springville, W.M. Tingey 148 (BRY); Wasatch Co..
southeast slope of Bear Canyon, A.C. Blauer 59 ( BRY' ) .

14. Mertensia Roth.

Merteiisia Roth, Cat. Bot. 1:34. 1797.

Glabrous or pubescent caulescent perennial
lierbs with fleshy, fusiform, rhizomelike or corm-
like roots; leaves entire, linear to cordate, ses-
sile or petiolate, alternate; stems 1-niany from
each root, decumbent to erect, usually branched
below the inflorescence, 0.3-17 dm tall; inflores-
cence a lax or congested, ebracteate, unilateral,
modified scorpioid cyme, or with the lowest
flowers often single and subtended by leaves,
often becoming panicled in age; calyx 5-parted,
occasionally campanulate, the expanded limb
exceeding or exceeded bv the tube, with or with-
out foniices in the throat, blue, occasionally
white or pink; filaments attached below the
throat, the anthers exserted or included; style
shorter or longer than the corolla, in some di-
or trimoi-phic; stigma entire or slightly lobcd;
ovary 2-celled, each cell 2-lobed; nudets 4, at-
tached laterally to the gynobase, usually rugose
or pectinately mgose, coriaceous or smooth ;md
shining, utriclelike.

Type species: Mertensia virginica (L. ) Pers.

1. Phuits usually with prominent lateral veins in the
cauline le;ives; stems usually 4 dm or more tall
( 1-17); nomiidly flowering in late spring anil in the
summer; mostly (Hciirring in moist, slKuled situ;i-
tions (2).

1. Plants usually without lateral veins in the cauline
leaves, some specimens of M. oblongijoUa var.
tic't iiilciusis e.vceptcd; stems usu;dlv less than 4 dm
t;ill, normally flowering in early spring, kiter when
growing in the mountains but commonly ;is soon
as the snow and tem])er;itures permit; mostly in
fairly open habits (7).

2. Limb of the eorolhi longer th;ui tlie lube; le:ives
usu;dlv acuminate (.3).

2. Limb ot the corolhi shorter th;ui the tube, or the

two about svibequal; leaves usually not acuminate

(6).

.3. Leaves pubescent at least on one surface ( 4 ) .

3. Leaves glabrous on both surfaces ( 5 ) .

4. Calyx not accrescent, margins densely ciUate, backs
pubescent or glabrous 5. M. frunciscana

4. Caly.\ accrescent, margins not densely ciliate, backs
glabrous IC. M. urizotiicci var. subnuda

5. Calyx campanulate, lobes shorter than the tube ...-
lA. M. arizunica var. urizonica

5. Calyx not campanulate, the lobes longer than the
tube IB. .\/. arizonica var. Iconurdii

6. Leaves pubescent at least on one surface

5. M. franciscana

6. Leaves glabrous on both surfaces, sometimes the
upper surface papillate 4. M. ciliata

7. Filaments attached in the corolla tube, the anthers
not projecting beyond the throat, contained within
the tube 3. M. brevlstijla

7. Filaments attached near the throat of the corolla
tube, anthers projecting beyond the throat, not
contained within the tube (8).

8. Limb of the corolla longer than or subequal to the
tube (8).

8. Limb of the corolla shorter than the tube (11).

9. Leaves pubescent on both surfaces, usually uni-
lateral 8B. M. viridis var. cana

9. Leaves strigose only above or glabrous on both
surfaces (10).

10. Filaments shorter than the anthers; calyx divided
nearly to the base; style usually not reaching the

anthers; anthers straight; plants alpine

8A. M. viridis var. viridis

10. Filaments longer than the anthers; calyx not divid-
ed to near the base; style usually reaching or sur-
passing the anthers; anthers usually cur\ed; plants
usu;dlv not alpine or sub;ilpine .... 6. M. fusiforviis

I 1 . Tube of the mature corolla only slightly longer
than the limb; plants of the high mountains (12).

IL Tube of the mature corolla usually much longer
than the limb; plains and low hills (15).

12. Lea\'es pubescent only above, or glabrous on both
surfaces ( 13).

12. Le;iyes pubescent on both surfaces (14).

13. Leaves glabrous on both surfiices, Uinta Moun-
tains 8C. M. vrridii var. dihilata

1.'5. L<'ayes strigose above - 8A. A/, lini/i.v v.ir. viridis

14. 1. caves usually unilateral, 1.5-3 cm long, stems
.isccnding 8B. A/, viridis var. cana

14. Leaves not unilateral, usually larger than ;d)ove;
stems more erect; Uint;i Mountains . 2. M. bakcri

i .5. Lt'.t\'es gl;d)rous on both surfaces

7C. M. ohlon<^ifoli(i var. ncvndensis

15. Leaves pubescent, ;it least on one surface (16).

16. Leaves pubescent iilxjve, glabrous below

7A. A/, obloiifiifolia var. oblongifolia

16. Leaves pubescent on both surf;ices

7B. A/, oblongifolia v;ir imwciui

Biological Skhies, \'ol. 16. No. .3

The Bor.\cinaceae of Ut.\ii

67

lA. Merteiisia arizonicci Greene, Pittonia 3:197.
1897.

Plants erect or ascending, 3-S cliii tall or
more; stems 1-several from each root stock, basal
leaves narrowly to broadly ovate or oblong-
lanceolate, 7.5-15 cm long, 2-6 cm broad, slight-
ly decurrent on the petiole, petiole as long as
the blade, glabrous but slightly papillate, mar-
gin ciliate; lower cauline leaves spathulate to
elliptical, usually petiolate, the petiole winged,
upper cauline leaves usually sessile, elliptical to
narrowly ovate, acute, 3-12 cm long, 1-5 cm
broad, base attenuate, apex acute; inflorescence
of axillary peduncles with branches elongating in
age; calyx 4-8 mm long, campanulate, glabrous
on the back, hairv within, the lobes }i or less the
entire length of the calyx, 2-4 mm long, acute or
obtuse, ciliate; pedicels 2-30 mm long, glabrous,
papillose or sometimes the papillae developing
short hairs; corolla tube 6-9 mm long, with a
definite ring of hairs at the base within; corolla-
limb 7-11 mm long, always longer than tube,
moderately expanded; anthers 2.5-3.5 mm long,
as long as or shorter and narrower than the fila-
ments, filaments 3-4 mm long, fomices conspicu-
ous, pubescent; style 10-15 mm long, usu;dly
shorter than the corolla; nutlets rugose, shorter
than the calvx. Central to southwestern Utah.
Moist stream banks and shaded areas. May-Au-
gust. Fig. 88.

;

UTAH

(L r^ '^ '

County Lines and

\a %X'

river drainages

i so fco

<"

1

1

1 / /^ ^> 1

-_A_i\ X

,"s 'l

1

/ lAv^—

'y--'

i/

_^ ry k

^

/

.___! -^

/—-^,

0\ /

v:/ , 1

A /

i^r^^^v"""

-^. <^' \

^S^ 5

C-..-^^

Fig. 88. \fcrtcn\Ui iLrizcnicii

var. (inzotucii

Beaver Co.. Big Flat, 5 mile.s .south of Puffer Lake,
K.H. Coles 88 (BRY); Tushar Mountains, vicinity of
Puffer Lake, M.L Morris 147 (BRY); Garfield Co.,
16 miles south of Esealante, N.H. Holmgren 2519
lUTC); Iron Co., Cedar Breaks Nad. Monument, A.
Bruhn 43 (UT); Cedar Breaks Natl. Monument, near
Point Supreme, W.S. Boyle 731 (BRY); Cedar Moun-
tain, Duek Creek, A.M. Woodbury s.n. (BRY); Cedar
Breaks. W.P. Cottam 3960 (BRY)'; Cedar Breaks, C.L.
Hitchcock 4.553. 4.572 (UTC); 10.5 miles east of Cedar
City. B. Maguire 19469 (UTC); Cedar Breaks, Brian
Hekd Peak, "B. Maguire 18995. 17577 (UTC); Kane
Co., south side of Nayajo Lake, R.K. Gierisch 491
(UTC); Piute Co., Tushar Mountains, Bullion Creek,
N.H. Holmgren 2147 (BRY, UTC); Washington Co.,
Pine Valley Campground, L.C. Higgins 1438 (BRY);
Pine Valley Mountains, along middle fork of the Santa
Clara Riyer. J.L. Gentry 2231 (BRY, UTC); Zion Natl.
Park, N.H. Holmgren i994 (BRY, UTC); Pine Valley.
W.P. Cottam 8874 (UT).

IB. Mcrtenski arizonica var. leonardii (Rydb. )
Johnst. Contr. Am. Arb. 3. S3. 1932.
Mertensia leonardii Rydb. Bull. Torrey

Club 36:680. 1909.
Mertensia siiinp'ionii Tidest. Proc. Biol. Soc.

Wash. 26:122. 1913.
Mertensia arizonica var. timbratahs Macbr.

ace. to Macbride Contr. Gray Herb.

U.S. 48:9. 1916.

Plants similar to the species; calyx 4-8 mm
long, divided almost to the base, lobes 3-7 mm
long, lanceolate, acute, ciliate; anthers and
filaments averaging slightly shorter than in
the species. Central and north central Utah and
southwestern Wyoming. Moist slopes and bot-
toms and shaded areas. May-August. Fig. 89.

Beaver Co., Puffer Lake, W.P. Cottam 3490 (BRY,
UT); headwaters of the Beaver River, B. Maguire 19793
(UTC); Cache Co., Spring Hollovv, B. Maguire 12996
(UTC); Logan Canyon, C.P. Smith 2216 (UTC); 1
mile above Tony Grove, B. Maguire 16711 (UTC);
Duchesne Co.. head of Blind Stream Canyon, B.F.
Harrison 88.39 (BRY); Emery Co., Huntington Canyon,
W.P. Cottam 7452 (UT); Garfield Co., Aquarius Pla-
te;iu, N.H. Holmgren 2112 (BRY, UTC); 15 miles north
of Esealante, N.H. Holmgren 2439 (UTC); Posey Lake,
Esealante Road. I. McArthur 173 (BRY); Aquarius Pla-
teau, B. Maguire 19175 (UTC); Iron Co., Cedar Breaks.
A. Eastwood & J.T. Howell 7277 (UTC); Juab Co.,
north of Mt. Nebo, W.P. Cottam 15648 (UT); Millard
Co., 2 miles above Adelaide Park, P. Plummer 178
(UT); Robins Vallev 20 miles west of Salina. F.H.
Coles 42 (BRY); Piute Co.. Tushar Mountains, Bullion
Creek, N.H. Hohngren 2144 (BRY, UTC); Salt Lake
Co., Big Cottonwood Canyon, E.B. Robinson s.n, (UT);
above Alta. L. Amow 4,58 (UT); Parleys Canyon, G.
Davidse 402 (UTC); Emigration Canyon, C.P. Smith
1846 (UTC); Brighton Guard Station, L. Eyre 12
(BRY); Sanpete Co., 15 miles east of Ephraim, R.D.
lorgensen 44 (UT); vicinity of Ephraim, R. Olsen s.n.
(UTC); head of Mayfield Canyon, B. Maguire 19992
(UTC); Manti Canyon, N.H. Holmgren 2.30 (UTC);
Horseshoe Flats, H. Johnson s.n. (BRY); 10 miles up
Ephraim Canvon, J.D. Walker s.n. (BRY); .Sevier Co..

68

Riur.nAM Young Univkhsitv Science Bulletin

UTAH

County Lines and
river drainages

L 1 I

o a5 io

Fig. 89.

Icrti'iisiti iirizoiiica var. Icoinirdii

Tushar Mountains, H.G. Warnock .s.n. (UT); Musunia
Hunger Station, D. Jt'ffery .s.n. (UTC); sunimit 7 miles
east of Cove Fort, L.C. Higgins 1473 (BRY); Summit
Co., Soapstone. Creek border. W.P. Cottam 3754 (BHY,
UT); about 3 miles west of Holiday Park, near head of
Weber River, Welsh, Murdoek & Stocks 6307 (BRY);
nortii slope of Big Moimtain, D.H. Galway 2.381
(BRY); Utah Co., Aspen C;rove. .A..O. Garrett 6187
(UT); Mount Timpanogos, Aspen Grove, H.C. Stutz 5
(BRY); Hobble Creek Canyon, W. Patrick 111 (BRY);
Mount Timpanogos, T.E. Northstrom 112 (BRY); sum-
mit of right fork of Hobble Creek Canvon, R.D. Day
40 (BRY); Aspen Grove. B.F. Harrison 580,5 (BRY)';
Wasatch Co., Provo River near Wolf Creek, W.P. Cot-
tam 9000 (UT); 16 miles southeast of Heber, Cron-
quist & Holmgren 9228 (UTC); 2 miles east of Soldier
Summit, B. Maguire 18.396 (UTC); Alpine, at first
bridge of Timp." Trail, L.D. Williams 32 (BRY); Cas-
cade Springs. S.W. Sanderson xl3 (BRY); Bryants
Fork, Strawberry Vallev, B.F. Harrison 87.39 (BRY);
Strawberry Valley. V.B'. Matthews 76 (BRY); Weber
Co., base of Ben Lomond. W. C ill 18 ( I't I (i nvlcs
southeast of monastery in Huntsville, L. Huffaker .54
(BRY); Ibnitsvillo, |. McNeley 47 (BRY); 5 miles south
of Huntsville AH. Mnir .57 (BRY).

KL McrtcmUi arizonica var. suhnuda (Maclir. )
Williani.s, Ann. Mo. Bot. Card. 24:66. 19.37.
Mertensia toi/<il)en.si.s var. stihnudo Machr.
Contr. Gray Hcrh. 48:7. 1916.

Differs from the typical plant in that iIk'
leav(?.s are .short-.strigo.se on tfie iipjKT surface,
glabrous or sparingly hairv on th<> lower surface;
calyx lohes divided almost to the base, glabrou.s

on the back or sparingly pubescent, ciliate; an-
thers 2-2..5 mm long. Sevier County, Utah, to
White Pine County, Nevada. May-August. Fig.
89.

Sevier Co., head of Niotche Creek, M.I. Morris 1.39
(BRY); Fish Lake Plateau, 8 miles up Goose Creek
road from highway 10, Salina Canyon, N.H. Holmgren
1970 (BRY).

2. Mertensia hakeri Greene, Pittonia 4:90.
1899.
Mertensia panicidata var. nivalis S. Wats.

U. S. Geol. Expl. 4()th Par. ( Bot. Kings

Exped.) 5:239. 1871.
Mertensia nivalis Rydb. Mem. N. Y. Bot.

Card. 1:336. 1900.
Mertensia lateriflora Grt^ne, Pi. Baker. 3:18.

1901.
Mertensia ini/osotifolia (Heller) Rydb. Colo.

Agr. Exp. Sta. Bull. 100:292. 1906.
Mertensia refracta A. Nels. Bot. Gaz. 56:69.

1913.
Mertensia hinceolata var. imjosotifolia

Macbr. Contr. Gray Herb. 48:15. 1916.

Plants with erect or ascending stems, 1-
scveral, 0.8-4 dm tall, simple or rarely branched,
pubescent with soft hairs; basal leaves linear-
lanceolate to ovate elliptic, more or less densely
canescent on both surfaces, 2-11 (4-6) cm long,
0.5-3.5 ( 1-1.5) cm broad, petiole longer or short-
er than the blade; cauline leaves linear-lanceo-
late to ovate, usually more or less densely canes-
cent on both surfaces, some plants from the Uin-
ta Mountains sparsely so, sessile or nearly so,
semiamplexicaule, 1.5-8 cm long (mostly 3-4)
cm, 0.5-2.5 cm broad; inflorescence congested to
loosely panicled; pedicels usually canescent, of-
ten reflexed in fruit, 1.5 cm or less long; calyx
2.5-5 mm long, divided almost to the base,
sparsely to usually densely pubescent on the
back and margins of lobes, lobes linear-lanceo-
late to lanceolate, acute, about 0.5 mm shorter
than tin- entire calyx; flowers di-, possibly tri-
morphic, the tiibe typically longer than the limb;
corolla-tube with more or less definite ring of
hairs toward the base within, 3.5-9 mm long;
c-orolla-limb moderately expanded, 4-6 mm long;
anthers 1.2-2 mm long, shorter and narrower
than the normal phase, longer and broader in
tlimorphic phases; fomices usually prominent,
glabrous, papillose, or pubescent; style reaching
or surpassing the :mthcrs; nutlets rugose, 2.5-3.5
mm long. Monnt;iins of Colorado, and the Uinta
Mountains Ut;ih to northern New Mexico. June-
ScptenilKT. Fig. 90.

Daggett Co.. north slope of the
Hiclinis .52 (UTC); Duchesne Co

Uint;i Mountains,
. Mount Fnimons,

I?i()i,()f;K:Ai, Sebies, Vol. 16, No.

The Boii.'VciNACE.vE or Ut.\h

69

UTAH

Fii;. 90. Mcrtcnsia hakcri

west meadow. J.R. Murdock 600 (BRY); Atwood Lake,
C.L. Hay%vard 26 (BRY); divide above Daggett Lake,
A.O. Spear 145 (BRY); Mount Agassiz, rocky summit,
W.P. Cottam .371.3 (BRY, UT); Bald Mountain, A.H.
Holmgren 703.5 (UTC); Mount Agassiz, B. Maguire
4240 (UTC); .Summit Co., Mount Lofty, B. Maguire
42.38 (UTC); Red Castle Lake, N.H. Holmgren 318
(UTC); 7 miles south of China Lake, N.H. Hohngren
.308 (UTC).

•3. Mertensia brevisiijhi S. Wats. U. S. Geol.

E.xpl. 40th Par. ( Bot. King.s Exped.) 5:239.

1S71. Type: Wasatch Mounbiins, Wat.son.

Mertensia alpirui var. brevKttjIa Jones

Contr. West. Bot. 12:56. 1908.

Plants wath erect or ascending stems, 1-4 dm
tall; 1-many from each fusiform rootstalk, more
or less pubescent; basal leaves broadly lanceo-
late to oblong, acute or obtuse, strigillose above,
glabrous below, 5-13 cm long, 2-4 cm broad,
petioles longer than the blade; cauline leaves ob-
ovate-oblong to narrowly elliptic, obtuse to
acute, densely strigillose above, glabrous below,
2-6 cm long 0.5-3 cm broad; inflorescence con-
gested at first, becoming panicled in age; pedi-
cels strigose, 1-14 mm long; calyx 2-5 mm long,
divided almost to the base, strigose, the lobe^
narrowly triangular to linear, acute, 1.5-4 mm
long, 0.5-1 mm brotid at the base; corolla-tube
2-4 mm long, slightly shorter to a little longer
than the calyx-lobes, with or witliout a ring of

scattered hairs toward the base within; corolla-
limb rotate, 4-6 mm long; anthers 1-1.3 mm
long, longer than the filaments, inserted on the
tube and not exceeding the throat; fomices more
or less prominent; style shorter than the calyx-
lobes; nutlets rugose, 2-3.5 mm long. Southern
Wyoming, west central Colorado, Utah, and
southeastern Idaho. May-July. Fig. 91.

UTAH

County Lines and
river drainages

i B» bo

1 1/ • 1

1

--A*Vh

~"K J 1

IK 1
-' / 1

"s3

'fty^

y

Fig. 91. Mertensia brevisUjhi

Box Elder Co., southeast of Deweyville, G. Piranian
s.n. (UTC); Cache Co., Logan Canyon, H. Richardson
2 (UTC); Sardine Canyon, B. Maguire 21662 (UTC);
K mile west of Mendon, Thomas & Marshall 44 (UT);
Davis Co., head of Chicken Creek, S.L. Clark 517
(UTC); Bountiful Peak, W.P. Cottam 15012 (UT);
Duchesne Co., head of Blindstream Canyon, Harrison
& Nisson 8835 (BRY); Juab Co., north of Mount Nebo
Ranger Station. R.K. Gierisch 267 (UTC); Salt Lake
Co., Parleys Canyon, G. Davidse 40 (UTC); Lambs
Canyon, R.K. Vickerv 19.34 (UT); south fork of Mill
Creek Canyon, W.J. Stubbs 84 (BRY); Summit Co., 3
miles east of Kamas on highway 150, R.J. Eastmond
218 (BRY); Tooele Co.. 12 miles southeast of Tooele,
L.C. Higgins .3334 (BRY); Utah Co., ridge southwest
of Pavson, G. David.se 312.5 (UTC); Pay.son Canyon,
W.P. Cottam s.n. (UT); Rock Canyon, A.O. Garrett
7550 (BRY); 4 miles up Pole Canyon, L.K. Shumway
46A (BRY); Hope Picnic Grounds, Pole Canyon, C.L.
Havward 20 (BRY); Deer Creek Canyon, B.F. Harrison
8311 (BRY); 1 mile east of Soldier .Summit, B.F. Har-
rison 7323 (BRY); Pole Canyon Exclosure, R.J. East-
mond 2.54 (BRY); Hobble Creek Canyon, B.F. Harri-
son 2514 (BRY); Squaw Peak, J.B. Karren 63 (BRY);
Provo Canyon, C.A. Han.son 14 (BRY); Wasatch Co.,
Soldier Summit, B. Maguire 18386 (UTC); Cascade

70

BmcMAM Young University Science Bulletin

Springs, S. Sanderson 11 (BRY); Hailstone junction,
R.J. Eastmond 232 (BRY); 1 mile west of Deer Creek
Reservoir, R.J. Eastmond 2.35 (BRY); Bryants Eork,
Strawberry VaUey, B.E. Harrison 8735 (BRY); Weber
Co., Snow Basin, A. Collotzi 5 (UTC); base of Ben
Lomond Peak, W. CaU 38 (UT).

inilate. Foothills and mountains up to 12,000
feet. Montana, Colorado, and Wyoming to east-
em Oregon and .south to northern New Mexico.
May-July. Fig. 92.

4. Mertensia cilmta (James) G. Don, Gen.
Hist. 4:372. 1838. Type: "Along streams
within the Rocky Mountains," collected by
James.

Mertensia ))ohjphiiUa Greene, Pittonia 4:87.

1899.
Mertensia punctata Greene, loc. cit. 88.
Mertensia ciliata var. longipedunculata A.

Nels. Bull. Torrey Club 29:402. 1902.
Mertensia picta Rydb. Bull. Torrey Club

31:638. 1904.
Mertensia pallida Rydb. Bull. Torrey Club

36:680. 1909.
Mertensia incongruens (Macbr. ) Payson,

Contr. Gray Herb. 49:66. 1917.
Pulmonaria ciliata James, Trans. Amer. Phil.

Soc. II. 2:176. 1825. Name only; Torr.

Ann. Lye. N. Y. 2:224. 1828.

Plants erect or ascending, 1-12 dm tall, usual-
ly with many stems from each rootstock; basal
leaves variable, oblong to ovate, or lance-
olate, subcordate, 4-15 cm long, 3-10 cm
broad, ciliate on the margins, often pa-
pillate on the upper surface, petioles long-
er or shorter than the blades; cauline leaves
lanceolate to ovate, acute, acuminate or obtuse
at apex, attenuate to subcordate at the base, the
lowennost short-petiolate, the uppermost sessile,
ciliate on the margins, often papillate on the up-
per surface, often (juite glaucous, thin in texture;
pedicels 1-10 mm long, glabrous, papillose or
rarely with a few short strigose hairs; inflores-
cence from the axils of leaves, the peduncles
elongated in mature or well-developed plants, in
young plants the flowers aggregated at the top
of the plant, each peduncle terminated in a
modified ebracteate scorpioid cyme, or occa-
sionallv subumbellate; calyx-lobes 1.5-3 mm
long, glabrous on the back, ciliate to papillate
on the margins, more or less strigose within,
obtuse or rarely somewhat acute, divided almost
or quite to the base, rarely enlarged in fruit;
corolla tube 6-8 mm long, glabrous or with
crisf>ed hairs within; corolla-limb 4-10 mm long,
sometimes longer than the tube, moderately ex-
panded; anthers 1-2.5 mm long, as long as or
shorter and narrower than the exp;indcd p;irt ol
the filament; fomices prominent, glabrous, papil-
late or pubescent; style alwut as long as the
corolla or exceeding it; nutlets rugose or mam-

County Lines and
river drainages
I I 1

Fig. 92. Mcrtoisui ciliata

Bo.\ Elder Co., Rosver Canyon, W.P. Cottam 2945
(BRY); R;ift River Mountains, W.P. Cottam 7036
(UT); Cache Co., Tony Grove, B. Maguire 3726
(UTC); near Tony Grove,' E. Jensen 92 (UTC); White
Pine Lake, B. Maguire 14283 (UTC); Daggett Co.,
Carter Creek, 12 miles south of Manila, B.F. Harrison
7885 (BBY, UTC); Carter Creek, B. Maguire 12387
(UTC); Duchesne Co., Roberts Pass, Uinta Mountains,
J.R, Murdock .584 (BRY); Moon L;ike, B.F. Harrison
7741 (BRY); Mount Agassiz, W.P. Cottiim 3714 (BRY,
UT ) ; southeast base of Movmt Emmons, C.L. Hayward
145 (BRY); Mount Emmons, J.R. Murdock 53 (BRY);
west of Mount Agassiz, B. Maguire 4242 (UTC);
Grand Co.. north slope of Mineral Peak, La Sal Moun-
tains, B.F. Harrison 12472 (BRY); north side of Gold
Mountain, B. Maguire 4.52 (UTC); north base of Hay-
stack Mountain, B. Miiguire 5115 (UTC); Rich Co., 10
miles west of Bear Lake, C. Greenhalgh 93 (UT); Salt
Lake Co., Albian Basin, W.P. Cottam 17704 (UT);
south of Brighton, R.K. Vicken, 1199 (UT); 1 mile
southwest of Brighton. B. Maguire 17349 (UTC); San
|uan Co., Mount Peal, La Sal Mountains, Hvdberg &
Garrett 8999 (UT); Canvon e;ist of Mount Tukuhniki-
viitz. A. Cronquisf 9478 (UTC); between Mount Peal
and Mount Tukuhnikivatz, B. Maguire 2116 (UTC);
Summit Co., headwaters of Provo River, Cottam & Bid-
.Inlph .3642 (BUY. UT); Soapstone, W.P. Cottam .3755
(BRY); Bald Mountain, Cottam & Biddulph 3699
(BRY): Stillwater Fork. Bear River, D. Henriques s.n.
(UT): Upper Henrys Fork Basin, B. Maguire 14334
(UTC); above Dolhir Lake, B. Maguire 14.596 (UTC);

Biological Series. X'ol. 16, No. 3 The Bohaginace.'ve of IItah

71

Uintah Co., 29 miles east of Kamas, B. Maguire 12535
(UTC); Utah Co., Hidden Lake, E. Castle 63c (BRY);
.-Vmerican Fork Canyon, I.E. Diehl s.n. (BRY); cirque
at base of Elk Peak, Mount Timpanogos, E. Castle 9767
(BRY); Silver Lake Area, American Fork Canyon, E.
Nixon 25 (BRY); Aspen Grove, B. Decker s.n. (BRY);
Aspen Grove, A.O. Garrett 3960 (UT); Mount Timpan-
ogos. B. Maguire 17461 (UTC); Wasatch Co., near
Lost Lake, about 2 miles from Trial Lake, E.C. Daven-
port 36 (BRY); Wolf Creek Pass, A.O. Garrett 7721
(UT).

5. Mertensia franciscaiui Heller Bull. Torrev
Club 26:549. 1899. Type: Vicinity of Flag-
staff, MacDougal 232.
Mertensia pratensis Heller loc. cit. 550.
Mertensia alba Rydb. Bull. Torrey Club

31:638. 1904.
Mertensia grandis Woot. & Standi. Contr.

U.S. Natl. Herb. 16:165. 1913.
Mertensia pratensis f. alba Macbride, Contr.

Gray Herb. 48:8. 1916.

Plants with erect or ascending stems, 1-10
( 17 ) dm tall, usually with several from each
rootstock; basal leaves oblong-elliptic to elliptic,
6-20 cm long, 5-9 cm broad, base subcordate to
obtuse, apex acuminate, acute or obtuse, upper
surface short strigillose, lower surface glabrous
or with spreading pubescence; petioles longer or
shorter than the blade; cauline leaves elliptical
to narrowly ovate, 4-14 cm long, 1-5 cm broad,
obtuse to acuminate, the lowermost petiolate,
becoming sessile toward the inflorescence, strig-
illose on the upper surface, glabrous to densely
pubescent with spreading hairs below; flowers
of the inflorescence paniculately disposed in an
ebracteate modified scorpioid cyme, the
branches of the inflorescence elongating in age;
pedicels strigose, 1-20 mm long; calyx 2.5-5 mm
long, divided almost to the base, the lobes linear
to lanceolate, 1-2 mm wide at the base, acute,
rarely obtuse, glabrous or pubescent on the back,
strongly ciliate; corolla-tube 5-9 mm long, gla-
brous or pubescent within; corolla limb 4-6 (9)
mm long, subequal to or slightly shorter or
longer than the corolla-tube, moderately ex-
panded; anthers 2.5-3 mm long, longer than the
filaments; filaments 2-2.5 mm long, glabrous or
with spreading hairs; fomices prominent, usual-
ly pubescent; style 9-20 mm long, usually short-
er than the corolla, sometimes exceeding it; nut-
lets rugose and papilliferous. Southern Colorado,
New Mexico, Arizona, southern Utah, eastern
Nevada, and California. June-August. Fig. 93.

Grand Co., Warner Ranger Station, La Sal Moun-
tains, H.K. Sharsmith 4432 (BRY); La Sal Mountains,
S. Flowers s.n. (UT); Oowah Lake, La Sal Mountains,
Weins & Amow 4147 ( UT, UTC); south ba.se of Gold
Hill, B. Maguire 5120 (UTC); north base of Haystack

Mountain, B. Maguire 5119 (UTC); San Juan Co., La
Sal, Aspen, W.P. Cottam 2220 (BRY); La Sal Ranger
Station, Cottam & Hutchings 2192 (BRY); Kigaha,
along creek, W.P. Cottam 2489 (BRY); 10 miles north
of Blanding, B. Maguire 2109 (UTC); Abajo Peak, B.
Maguire 2115 (UTC); La Sal Pass, Medicine Lake,
Cronquist & Holmgren 9469 ( UTC ) ; Kigaha Ranger
Station, D. Henriques 128 (UT); La Sal Mountains,
Rydberg & Garrett 8597 (UT).

6. Mertensia fu^iformis Greene, Pittonia 4:89.
1899. Type: Open glades. Bob Creek, West
La Plata Mountains, Baker, Earle & Tracy
206.
Mertensia congesta Greene, PI. Baker. 3:17.

1901.
Mertensia papillosa fusiformis A. Nels.

Coult. & Nels. Man. Rocky Mt. Bot. 421.

1909.

Plants with erect stems or nearly so, 1-3 dm
tall, glabrous or sparingly pubescent, 1-few from
each root stalk which is usually rather large and
fusiform; basal leaves elliptic to oblong-ovate,
4-12 cm long, 1.5-3 cm broad, usually densely
strigose above, glabrous below, petiole 7-12 cm
long; cauline leaves linear-oblong to ovate-ob-
long, 1.5-10 cm long, 0.4-3 cm broad, sessile or
the lowermost short-petiolate, more or less dense-
ly strigose above, glabrous below, usually quite
obtuse, rarely somewhat acute; inflorescence usu-
ally congested, sometimes slightly panicled; ped-

County Lines and
river drainages

i-

Fig. 93. Mertensia francvicaim

72

Bnir.HAM Young Universitv Science Bulletin

icels 1-15 mm long, densely strigose; calyx 3-6 mm
long, slightly accrescent, the lobes lanceolate to
lanceolate-ovate, 2-5 mm long, acute, ciliate,
usually pubescent on the backs, occasionally
nearly glabrous, not divided to the base; corolla-
tube 4-7 mm long, with a ring of crisp hairs
within at the base; corolla-limb 5-7 mm long,
moderately expanded, usually suljccjual to or
shorter than the limb, but sometimes longer;
anthers 1.5-2.5 mm long; filaments 1-3 mm long
fomices present but usually not conspicuous,
glabrous or nearly so; style usually surpassing
the anthers, sometimes shorter; nutlets rugose,
about 3 mm long. Wyoming, Utah, and Colora-
do in the mountains. June-September. Fig. 94.

UTAH

County Lines and
river drainages

i-

Fijj. 94. Mcrtensui fusifonnis

Daggett Co., summit along the Vernal-Manila road,
W.I'. Cottam 6070 (UT); Duchesne Co.. shore of Moon
Lake, I..M. Hartman .36 (BRY); Rock Creek, J. Broth-
erson 907 (BRV); 4 miles south of Moon Lake, Har-
rison & Larsen 7618 (BRY); Kane Co., 1.5 miles nortli-
west of Ordt-rville, B. Maguire 18802 (UTC); 20 miles
northwest of Orderville, B. Maguire 18756 (UTC); Salt
Lake Co.. Parleys Canyon, R.K. Selander 1.502 (UT);
.San Juan Co., west slope of the La .Sal Mountains
between Horse and Bmmler Creek, Cron()uist & Holm-
gren 9285 (UTC); along La Sal Creek, A. Cronquist
9215 (UTC); Yellow Pine Association, B.F. Harrison,
.5895 (BRY); La Sal, along La Sal C;rc<'k. Cottam &
Hutchings 2270 (BRY); Uintah Co., Diamond Motui-
tain, Lavcock & Conrad 1.37 (BRY); Diamond Moun-
tain, 15 miles north of Vernal, Welsh & Moore 1959
(BRY); Wasatch Co., Wolf Creek Pass W.P. Cottam

9013 (UT); Washington Co., Zion Natl. Park, Horse
Pasture Plateau, Potato Hollow, N.H. Holmgren 1976
(BRY, UTC).

7A. Mertensia oblongijolia { Nutt. ) G. Don,
Gen. Hist. 4:372. 1838. Type: Montana,
probably collected by Wyeth.
Mertensia nutans subsp. subclava Piper,

coiitr. U.S. Natl. Herh. 11:479, 1906.
Mertensm foliosa var. subclava Macbr.

Contr. Gray Herb. 58:18. 1916.
Mertensia nelsonii Macbr. Ic. 19, in part.
Mertensia bakeri var. subglabra Macbr. &

Payson, Contr. Gray Herb. 49:66. 1917.
Mertensia foliosa var. subclava i. macbridii

Johnst. Contr. Am. Arb. 3:84. 1932.
Pulmonaria oblongifolia Nutt. Journ. Acad.

Nat. Sci. Phila. 7:43. 1834.
CerintJiodes obhngifolium Kuntze, Reo.

Gen. PI. pt. 2:436. 1891.

Plantis with erect or ascending stems, 1-3 dm
tall, one to many from each elongated rootstalk;
blade of basal leaves 3-8 cm long, 0.5-2 cm
liroad, oblong or spathulate to narrowly oblong-
ovate, usually obtuse, strigose on the upper sur-
face, glabrous below, petiole longer or shorter
than the blade; cauline leaves sessile or the
lowermost short-petiolate, linear to oblong-ellip-
tical, 2-8 cm long, 0.3-1.5 cm broad, pubescent
as of basal leaves; inflorescence congested, be-
coming panicled with age; pedicels strigose to
essentially ghibrous, 1-10 mm long; calyx 3-7
mm long, divided to within about 1 mm of the
base, the lobes linear to hmceolate-triangular,
acute, ciliate, glabrous doreally or rarely with a
few hairs; corolla-tube 5-12 mm long, usually
(juite glabrous within, occasionally with a few
scattered hairs; corolla-limb 4-7 mm broad; an-
thers 1.2-2 mm long, oblong and straight; fila-
ments 2-4 mm long, usually longer and broader
than the anthers; style exceeding the anthers;
fomices prominent, glabrous or occasionally
sparsely hairy; immature nutlets 3-4 mm long,
mgose. Westem Montana, Wvoming, and Utah
to Washington and northem Califomia. June-
July. Fig. 95.

Salt Lake Co.. Little Cottonwood Canyon, W.P.
Cottam .3.5.32 (BRY); Uintah Co. Split ' Mountain
Corge. S.L. Welsh 244 (BRY); Utah Co., Silver Lake,
American Fork Canyon, LE. Diehl s.n. (BRY).

7B. Mertensia oblongifolia var. amocna (A.
Nels. ) Williams, Ann. Mo. Bot. Card.
24:1.30. 19.37. Tvpe: Monida, Madison Co.,
Nel.son & Nelson .5413.
Mertensia amoena A. Nels. Bot. Gaz.
30:195. 1900.

RiOLOCicAL Series, Vol. 16, No. 3 The Boraginaceae of Utah

73

Mertensui ctisickii Piper, Bull. Torrey Club

29:643. 1902.
Mertensia pubescens Piper, Contr. U.S.

Natl. Herb. 11:479. 1906.
Mertensia bakeri amoena A. Nels. Coult.

& Nels. Man. Rocky Mt. Bot. 422. 1909.
Mertensia joUosa var. fiubesccns Macbr.

Ck)ntr. Gray Herb. 48:19. 1916.
Mertensia obJongifoIia var. nimbata Macbr.

Contr. Gray Herb. 53:18. 1918.
Mertensia cooperae Peck, Torreya 32:151.

1932.
Mertensia foJiosa var. cnnocna Johnst.

Contr. Am. Arb. 3:85. 1932.

Similar to the species, often more robust;
leaves more or less densely pubescent on both
surfaces. Sporadic with the species and variety
nevadensis. Fig. 95.

UTAH

Counry Lines and
river drainages

i 30 *>"

Fig. 95. Mertensia ohlongifolia var. ohlongifolia

Rich Co., among sagebrush, north slope of a small
canyon about 3 miles east of I^aketown, 28 May 193.^.
Williams & Williams 21.52 (NY).

7C. Mertensia oblontgifalia \ar. ncvadensi.'i (A.
Nels.) Williams, Ann. Mo. Bot. Card.
24:125. 1937. Type: 5 miles west of Reno,
Kenned V & True 711.

Mertensia folio.sa A. Nels. Bull. Torrctj
Club 26:243. 1899.

Mertensia tubiflora Rydb. Bull. Torrey

Club 26:544. 1899.
Mertensia intennedia Rydb. Mem. N.Y.

Bot. Card. 1:335. 1900.
Mertensia nutans Howell, Fl. N.W. Amer-

491. 1901.
Mertensia stenoloba Greene, PI. Baker.

3:20. 1901.
Mertensia sijmphtjtoides Greene, Ic. non

Fisch. 1872.
Mertensia coronata A. Nels. Bull. Torrey

Club 29:403. 1902.
Mertensia nevadensis A. Nels. Proc. Biol.

Soc. Wash. 17:96. 1904.
Mertensia praecox Smiley ex Macbr. Contr.

Gray Herb. 48:10. 1916.
Mertensia foliosa var. nevadensis Macbr. Ic.

19.
Mertensia iwhonii Macbr. Ic. in part as to

type specimen.

Similar to the species but often more robust;
cauline leaves 2-10 cm long, 0.8-6 cm broad,
lanceolate-oblong to ovate, glabrous or the upper
surface pustulate, sometimes the pustules toward
the apex of the leaves developing mucros, some
specimens, particularly from northeastern Utah;
with lateral veins in the well-developed cauline
leaves; corolla extremely variable as to size in
different localities. Western Montana, Wyoming,
and Utah west to Washington and northern Cali-
fornia. April-July. Fig. 96.

Box Elder Co., Goose Creek Moimtains. 3 miles
northwest of Divide, K.S. Erdman 1644 (BRY); Ros-
vere, top of mesa, W.P. Cottam 2847 (BRY); Raft
River Mountains, at summit, N.H. Holmgren 2850
(BRY); above Cottonwood Grove, M. Burke 3118
(UTC); Raft River Mountains, C. McMillan 1.503
(UT); Cache Co., Logan Canyon, B. Maguire s.n.
(UTC); Providence Canyon, B. Maguire 3719 (UTC);
Logan Canyon, M. Burke 3725 (UTC); 11 miles up
Smithfield Canvon, B. Maguire 13776 (UTC); Juab Co.,
Deep Creek Range, Maguire & Holmgren 22050 ( UTC ) ;
Sheba Mine, Deep Creek Mountains, W.P. Cottam 3176
(BRY, UT); Deep Creek Mountains, Maguire & Be-
craft 2784 (UTC); Deep Creek Range, A.H. Holmgren
.3746 (UTC); Salt Lake Co., Big Cottonwood Canyon,
R.J. Eastmond 186 (BRY); Big Cottonwood Canyon,
B. Fowler s.n. (UT); Summit Co., Henrys Fork, A.
Collotzi 45 (UTC); Weber Co., Snow Basin, A, Collot-
zi .50 (UTC); Pine View Dam, J. McNeley 40 (BRY);
Tooele Co., 3 miles south of Tooele, L. Arnow 266
(UT); trail to Mount Deseret, B. Maguire 21767
(BRY); Bennion Canyon, N.C. Frischknecht 81 (BRY);
Muir Trail, Oquirrh Mountains, Cottam, Allen & Row-
land 16867 (BRY); Black Bunch, 1 mile north of Miner
Fork Road, M.E. Lewis 1115 (BRY); Utah Co., Spring-
\ille, B. Dougall 97 (UT); Provo Canvon, M.E. Jones
s.n. (UTC); Hobble Creek Canyon, L.C. Pack 7
(BRY); along Timp. Creek, L.K. Shumway 92 (BRY);
Bridal Veil Falls, S.L. Welsh 3053 (BRY); north end of
Lake Mountain, Welsh & Moore 1927 (BRY); Hobble
Creek Canvon, B.F. Harrison 264 (BRY); just east of

74

Bnir.iiAM Young UsivEnsiTV Science Bulletin

Counfy Lines and

river drainages

I I I

«5 so fco

Fig. 96. Merteiiiia ohlongifolia var. iicuadensis

Eureka, J.B. Karren 79 (BRY); Hobble Creek Canyon,
F.J. Camcnzind s.n. (BRY); Hobble Creek Canyon, T.E.
Northstrom 13 (BRY); Hobble Creek Canyon. L.C.
Higgins 3500 ( BRY ) .

.SO, lanceolate to broiidly ovate, 2-7 cm long, 0.7-
2.5 cni bro;id, strigillose above, glabrous below,
lateral veins rarely visible; inflorescence a
crowded, modified, scorpioid cyme; pedicels stri-
gose or glabrous, 1-10 mm long; calyx 2-6 mm
long; the lobes divided almost to the base, linear-
lanceolate to narrowly ovate-lanceolate, acute or
obtuse, glabrous on the backs, ciliate, somewhat
accrescent in fruit; corolla-tube .3-9 mm long,
usually with a ring of crisped hairs near the base
within, occasionally also with scattered hairs;
corolla-limb 4-9 mm long, moderately expanded,
usually a little shorter than the tube in flowers
having a long style and filaments longer than
the anthers, in flowers having a short style and
filaments about the same length as or shorter
than the anthers, the tube may be much shorter
than the limb; anthers 1-2.5 mm long; filaments
1-.3.5 mm long, longer to shorter than the an-
there; fomices conspicuous, glabrous to densely
pubescent; style various, exceeding the anthers
and about as long as the corolla to shorter than
the tube of the corolla; nutlets 2-3 mm long,
rugose. Montana, south through Utah and Colo-
rado. July-September. Fig. 97.

Carfield Co., 25 miles south of Hank.sville, Henry
Mountains, Cronquist & Holmgren 9435 (UTC); south
rim of Bull Creek Basin, Henry Mountains, B. Maguire
19.362 (UTC); Grand Co., north side of Gold Moun-
tain. B. Maguire 5113 (UTC); saddle bet\veen Castle
and Waas mountains, B. Maguire 5112 (UTC); west

8A. Mertensia viridis A. Nels. Bull. Torrey
Club 26:244. 1899. Type: Wyoming, Lara-
mie Peak, Nelson 1608.
Mcrtemki lanceolata var. viridis A. Nels.

First Kept. Fl. Wyo. 158. 1896.
Meftensi<i ovata Rvdb. Bull. Torrey Club

28:32. 1901.
Mertensia lirwariloba Rydb. Ic.
Mertensia parri/i Rydb. Bull. Torrey Club

31:6.39. 1904.
Mertensia perplexa Rydb. Ic.
Mertensia viridula Rydb. Ic.
Mertensia papiUosa Uneariloba A. Nels.

Coult & Nels. Man. Rocky Mt. Bot.

421. 1909.
Mertensia lanceolata var. Uneariloba Macbr.

Contr. Gray Herb. 48:15. 1916.
Mertensia alpina var. perplexa Macbr. Ic.

20.

Stems erect or ascending, 5-3.5 cm tall, 1-
several from each rootstalk; basal leaves lanceo-
late to ovate, 2-10 cm long, 1-4 cm broad, strig-
illose above, glabrous below, lateral veins
sometimes apparent, petioles longer or shorter
than the blade; cauline leaves sessile or nearlv

UTAH

Fig. 97. Mcrtcnaiii viridis var. viridis

Biological Series, \<)l. 16, No. 3

The Bohaginaceae of L'tah

75

slope of Mount Hobbs, B. Maguire 5111 (UTC); San
Jiuin Co., west slope of Mount Peal, B. Maguire 2117
(UTC); saddle south of Mellenthin, B. Maguire 5109
(UTC); La Sal Mountains, Mount Peal, N.H. Holm-
gren 3003 ( BRY ) .

SB. Mertensia viridis var. cana (Rydb. ) Wil-
liams, Ann. Mo. Bot. Card. 24:115. 1937.
Type: Colorado, Grand Co., Berthoud Pass,
Tweedy 5664.
Mertensia canescens Rydb. Bull. Torrey

Club 31:640. 1904. '
Mertensia cana Rydb. Bull. Torrey Club

36:698. 1909. '

Cauline leaves linear to narrowly ovate, more
or less densely canescent on both surfaces; calyx
lobes usually glabrous on the backs but some-
times slightly pubescent. Bald Mountain, Utah.
Fig. 97.

Bald Mountain. 11 August 1911. demons s.n.
(POM).

8C. Mertensia viriditl var. dUatata (A. Nek.)
Williams, Ann. Mo. Bot. Card. 24:113. 1937.
Type: Wyoming, Albany Co., Medicine
Bow Mts., Nelson 7844.
Mertensia coriacea A. Nels. Bull. Torrey

Club 29:402. 1902.
Mertensia coriacea var. dilatata A. Nels. Ic.

403.

Similar to the species but the leaves glabrous
on both sides. Uinta Mountains, Utah. Fig. 97.

Daggett Co.. Uinta Mountains, 10,000 feet, 11 June
1932, L. WiUiams 599 (UTC).

15. Myosotis (Dill.) L.

Myosotis (Dill.) L. Sp. PI. 1:131. 1753, and Gen.
PI. 63. 17,54.

Annual or perennial herbs; leaves alternate;
racemose, bractless or bracted, calyx cut to be-
yond the middle into lanceolate or triangular
lobes; corolla blue, white or rarely rose, with a
short tube; lobes contorted, rounded, spreading;
throat with intruded appendages; stamens af-
fixed on the tube, included or exserted, the fila-
ments filiform; anthers oblong or ovate, obtuse;
ovules 4; nutlets 4, erect, ellipsoid, smooth and
shinv, with a basal areola; gynobase flat or high
convex; style filiform; a stigma disciform.

Type species : Myosotis scorpioides L.

1. Myosotis scorpioi^les L. Sp. PI. 131. 1753.
Myosotis scorpioides var. palustris L. Ic.
Myosotis palustris Lem. Fl. Fr. 2:2a3. 1778.

Perennial, with slender rootstocks or stolons,
herbage appressed-pubescent with straight
pointed hairs; stems slender, 1.5-4 dm long, de-
cumbent or ascending, rooting at the lower
nodes; leaves oblanceolate to oblong-lanceolate,
2.5-8 cm long, 4-12 mm wide, upper stem leaves
sessile, the lower narrowed to a winged petiole;
racemes loosely many flowered; fruiting pedicels
longer than the calyx; calyx with straight ap-
pressed hairs, the lobes equal, ovate triangular,
acute, shorter than the tube, more or less spread-
ing in fruit; corolla blue with a yellow eye, the
limb flat, 6-8 mm broad; nutlets angled and
keeled on the inside. Wet meadows and margins
of streams. Native of Europe and Asia, well es-
tablished in the United States. June-August. Fig.
98.

UTAH

Fig. 98. Myosotis scorpioides

Cache Co., H mile west of Logan, B. Maguire
20100 (UTC); 1 mile west of Logan along canal, B
Maguire 21579 (UTC); Logan, ditch bank, W.S.F. 544

(UT).

16. Pectocarya DC. ex Meisn.
Pectocarya DC. ex Meisn. Gen. 279. 1840.

Low, often spreading annual herbs, with
slender stems and narrowly linear leaves, canes-
cent with a close-appressed pubescence; flowers
scattered along the stems or branches, on short
pedicels, solitary in the axils; calyx 5-parted, the

76

Unir.HAM Young University Science Bulletin

lobes narrow, spreading or reflexed in fruit; co-
rolla white, the tube shorter than the calyx, lobes
broadly oval, the throat nearly closed by prom-
inent crests; stamens included; style very short;
nutlets flattened, thin, widely divergent either
radiately or in pairs, their margins, at least to-
ward the apex, with a row of hooked bristles.
Type species; CA'noglossuni lateriflorum Lan.

1. Nutlets orbicular or nearly so, both the body aiul
the very thin conspicuous wing beset with slender
uncinate brisdes 4. P. setosa

1. Nutlets oblong or linear, the body without uncinate
bristles (2).

2. Nutlets heteromorphic, 1 of each divergent pair
wingless, or merely margined, the other with a
broad somewhat incurved uncinate-toothed wing
1. r. hetewcarpa

2. Nutlets not heteromorphic, all 4 winged-margined
or toothed ( 3 ) .

3. Margin of nutlet conspicuous, the teeth confluent
at base 2. P. platycaqm

3. Margin of the nutlet very narrow or wanting, the
teeth being nearly or (juite distinct, subulate, nut-
lets strongly recurved 3. P. recurvata

1. Pectocarya hetewcarpa Johnst. Joum. Am.
Arb. 20:399. 1939. Type: Com Springs,
Chuckwalla Valley, Riverside County, Cali-
fornia.

Pectocarya penicillata var. heterocarpa
Johnst. Contr. Gray Herb. 70:37. 1924.

Diffusely branched from the base; stems
slender, ascending or spreading, 3-15 cm long,
strigose and canescent throughout; leaves nar-
rowlv linear, 1-3 cm long, 1-2 mm wide, the hairs
on tile basal ones often pustulate at base; corol-
la minute, its limb about 1.5 mm broad; fruit-
ing nutlets widely divergent, dissimilar, 2 nar-
rower and with or without a narrow margin, and
2 prominentlv winged-margined, the wings pec-
tinately bri.stly at the apex, irregular, few
toothed and with or without scattered bristles
on the sides. Sandy or gravelly slopes and plains.
Southwestern Utah, southern Nevada south
through Arizona and southern California to
Sonora, Mexico. January-May. Fig. 99.

Washington Co., 1 mile cast of Hurricane, B. Ma-
guire 1.522 (U'PC); volcanic hills west of Hurricane, B.
Maguire 1.52.5 (UTC); St, George. M.E. Jones s.n.
(UTC); Di.xie State Park, L.C. Higgins 918 (BRY):
Bcaverdam Mountains, L.C. Higgins 284. 3.52 (BRY);
Black Hills west of St. George, C.W. Cottam s.n. (DIX).

2. Pectocarifa platycarpa Munz & Johnst. Contr.
Gray Herb. Sl;81. 1928. Type: Mesa near
Camp Lowell, Arizona.
Pectocarya aracilis var. platycarpa Munz

& Johnst.
1924.

Contr. Gray Herb. 70:36.

Stems slender, diffusely branched from the
base, prostrate or widely ascending, 5-20 cm
long, cinercous-strigillose throughout; leaves nar-
rowlv linear to linear-oblanceolate, 0.5-1.5 mm
wide, 1-3.5 cm long; calyx-lobes nearly as long as
the nutlets; corolla 2 mm long; nutlets divergent
in pairs, sometime*; heteromorphous, linear-ob-
long or spatulate-oblong, 2.5-3 mm long, with a
wide conspicuous stramineous margin bearing
irregular uncinate-tipped teeth, the odd nutlet
when differentiated, with more deeply dissected
wing and with more pubescent body. Dry grav-
elly slopes and benches. Southwestern Utah,
south and west through Arizona, southern Neva-
da and California. Febmary-May. Fig. 100.

Washington Co., Watercress Springs, D.H. Galway
s.n. (DIX); north of St. George, D. Hall s.n. (DIX);
Beaverdam Mountains, L.C. Higgins 350 (BRY); St.
George, D.H. Galwav 6286 (BRY); Black Hill west of
St. George. D. H.all s.n. (BRY).

3. Pectocarya recurvata Johnst. Contr. Am.
Arb. 3:97. 19.32. Type: near Chandler, Mari-
copa County, Arizona.

Stems slender, simple below, \vith 2 to sev-
eral erect or ascending branches above, or some-
times diffusely branched throughout and more

UTAH

County Lines and
river drainages

A to U

Fig. 99. Pectocartfa heterocarpa

Biological Series, Vol. 16, No. 3 The Boracinaceae of Utah

77

UTAH

County Lines and
river drainages

i-

Fig. 100. Pectocaryo platycarpa

spreading. 5-25 cm long; herbage cinereou.s-stri
gose; leaves narrowly linear, acute, 1-3.5 cm
long, 0.5-2 mm broad; calyx-lobes barely 2 mm
long in fruit, acute; nutlets divergent in pairs,
linear, strongly recurved, the wing divided to or
almost to the body into prominent subulate
straw-colored uncinate bristles, at the apex the
wing prolonged into a short scarious tip, unci-
nate bristly on the margin. Sandy or gravelly
slopes and ridges. Lower Sonoran Zone. South-
em Nevada, western Utah, Arizona, and south-
ern California. March-May. Fig. 101 (semi-
circle ) .

I have seen no Utah specimens of P. recur-
vata, but the species has been collected a few
miles from the Utah border in Mohave County,
Arizona.

4. Pectocanja setosa A. Gray, Proc. Amer.
Acad. 12:81. 1S76. Type: desert plains of
the Upper Mohave River.
Pectocanja setosa var. aperta Johnst. Contr.

Gray Herb. 70:38. 1924.
Pectocanja seiosa var. holoptera Johnst. op

cit. 39.
Gruvelia setosa Rydb. Bull. Torrey Club

40:479. 1913.

Stems usually diffusely branched from the
base, ascending, slender to rather stout, 5-20 cm

tall, herbage rather thinly strigose and setose
with spreading bristle-like hairs; leaves linear
to linear-oblanceolate, 5-20 mm long; calyx-
lobes narrowly linear, 3-4 mm long in fruit,
armed with 3 to 6 straight divergent bristles;
nutlets divergent in pairs, broadly obovate to
orbicular, 2 borders all around with a thin scar-
ious wing, 2 wingless, the body of the nutlets
and usually the wing bearing slender uncinate
bristles, the wing usually slightly undulate and
slightly curved upward saucerlike. Usually fine
sandy areas. Eastern Washington and central
Idaho, southward to Utah, Arizona, Nevada,
and Baja California. April-June. Fig. 101.
( circles ) .

Washington Co., Diamond Valley, F.W. Gould
1.557 (DIX); 10 miles north of St. George, F.W. Gould
1557 (UTC); Diamond Valley, L.N. Goodding 815
(UTC); Welcome Springs, Beaverdam Mountains, B.
Maguire 20521 (UTC); Beaverdam Mountains, B. Ma-
guire 4952 (UTC); Diamond Valley, 10 miles north
of St. George, L.C. Higgins 4203 (BRY,WTSU).

17. Plagiobothnjs Fisch. & Mey.

Plagiobothrys Fisch. & Mey. Ind. Sem. Hort.
Petrop. 2:46. 1835.

Slender, glabrate or mostly soft-pubescent,
annual or perennial herbs; leaves mostly linear
or linear-lanceolate, alternate above and either
opposite at base or forming a rosette; flowers

UTAH

Counly Lines and
river drainages

i-

Fig, 101. Pectocarya recurvata

78

IJmcuAM Young University Science Bulletin

in bractless or bracteate spikelike racemes, the
racemes more or less scorpioid and usually
elongated in fruit; corolla white, small, salver-
form, with crests at the mouth of the throat;
nutlets rugose, erect or incurved, attached at or
below the middle to a depressed gynobase
through a caruncular scar, this decurrent on the
lower part of the ventral keel or situated at the
lower end of the keel and sunken below its
crest.

Type species: Plagiobotlinjsrujescens Fisch.
& Mey.

1. Plants glabrous or nearly so; nutlets attached basal-
ly or nearly so 3. P. Icptocladus

1. Plants strigose or bristly hairy; nutlets attached
laterally or obliquely basal (2).

2. Nutlets checkered with broad flattened, contiguous,
pavementlike raised areas; plants erect, hispid, with
terminal bractless scorpioid cymes 2. P. joncsii

2. Nutlets not checkered, the back rugose, the raised
areas scattered or none (3).

3. Leaves charged with purple dye, particularly at
midrib and margins; calyx with a weakened ring
which allows it to break loose, lobes short, strongly
pressed together at maturity 1. P. arizonicus

3. Leaves lacking conspicuous purple dye, green;
calyx without weakened ring (4).

4. Basal leaves crowded into a rosette, none opposite,
plants slender, erect, loosely branched, not produc-
ing flowers near the base; nutlets incurved, con-
tracted at both ends, somewhat cruciform, trans-
verse ridges very broad 5. P. tenellus

4. Basal leaves distinct, at least not in well-developed
rosettes, lower leaves opposite; nutlets not incurved,
but highly variable 4. P. scouleri

1. Plagiobothrys arizonicus (A. Gray) Greene,
Proc. Amer. Acad. 20:284. 1885. Type: Ari-
zona.

Eritricliium canescens var. arizonicum A.
Gray, Proc. Amer. Acad. 17:227. 1882.

Stems slender, several from the base, as-
cending, simple or few branched, 1-4 dm high,
hirsute-hispid with spreading hairs and also
rather sparinglv villous pubescent; leaves hir-
sute-hispid with more or less appressed hairs,
pustulate at base, without shorter pubescence,
the lower linear-lanceolate, 1.5-5 cm long, upper
linear-oblong to lanceolate; roots, lower parts of
stems and veins of the leaves, or sometimes the
whole plant purplish; spikes at length elongated,
remotely flowered and bractless or with a few
foliaceous bracts; calyx about 3 mm long, cleft
to about the middle, lobes narrow-attenuate,
eonnivent, hirsute-hispid, the tube at length usu-
ally circumscissle near the ba.se; corolla 2-2.5
mm broad; nutlets 1-4 commonly 2, ovoid and
abruptly acute at apex, median and lateral keels

often tuberculate, and with connecting trans-
verse ruiiae, the areolae between smooth or
minutely papillate; scar median, seated in a
sunken area at the base of the keel. Sandy to
gravelly slopes and plains. Southwestern Utah,
south to southern California, Sonora, and New
Mexico. March-June. Fig. 102.

County Lines and
river drainages

i-

Fig. 102. PUigiobothrys ahzoniciLi

Washington Co., west slope of the Beaverdam
Mountains, W.P. Cottam 7545 (UT); east of Laverkin,
W.P. Cottam 5165 (UT); west slope of the Beaver-
dam Mountains, D. Nish 68 (UTC); Beaverdam Moun-
tains, B. Maguire 20508 (UTC); St. George, D.H. Gal-
way 8451 (BRY); Black Hill west of St. George, F.W.
Gould 1481 (BRY); Beaverdam Wash at Terry's Ranch,
L.C. Higgins 300 (BRY); Dixie State Park, L.C. Hig-
gins 850 ( BRY ) .

2. Phi'^iohothriis jonesii A. Gray, Svn. Fl. N.
Amer. 2:430. 1886. Type: Needles, Califor-
nia.
Sontiea jonesii Greene, Pittonia 1:23. 1887.

Stems erect, 1 -several from the base, diver-
gently branched, 1-3 dm high, hispid with
spreading bristly hairs pustulate at base, and
also retror,sely-pul)escent; basal leaves linear or
narrowlv oblanceolate, cauline mostly lanceo-
late with pubescence similar to the stem but
thinner; racemes terminating the branches, most-
Iv conspicuously leafy-bra oted at base, 1.5-3 cm
long, the lower leaves of the branches often

Biological Series, Vol. 16, No. .3 The Borac.inaceae ok I'tah

79

bearing one or tew axillary flowers, calyx lobes
.subulate-linear, 6-8 mm long, corolla 1-2
mm broad; nutlets 3 mm long, incurved
and 4-angled bv the dorsal and ventral
keels and the 2 lateral ridges, abruptly pointed
at apex, the keel and lateral angles tuberculate,
the conc;ive surface between densely tessellate;
scar narrow or medial-narrow merging into the
keel alx)ve, and with a diverging lateral ridge
extending to either side. Washes and rocky
desert slopes. Southwestern Utah, south to
southern Califomia. and Arizona to Sonora.
April-May. Fig. 103.

UTAH

m^

County Lines and

river drainages

d 8« fc*

1

1

-.-'--T-'-^-'V

i jyy-^Qy-^

-X-U./

!

TyL^

'y '-- - - -

A

1

__ / -^

u /

v-y I !

<> /

~\ "1 elz;^

7 /

_-_,/i^i__

A r

-T

I w _

-y

Fig. 103. Plagiobothrys jonesii

Washington Co., 5 mile.s west of Leeds, B. Ma-
guire 1.536 (UTC); Black Hill west of St. George, D.H.
Galwav 8.532 (BRY); Beaverdam Mountains, L.C. Hig-
gins 454 (BRY).

Plaoiohotlinis leptocladus (Greene) Johnst.
Contr. Am. Arb. 3:38. 1932. Type: Pine
Creek, Eureka County, Nevada.
Eritricliium californictim var. \s-ubglochidia-

tum A. Gray, Bot. Calif. 1:526. 1876.
AUocanja leptodada Greene, Pittonia 3:109.

1896.
AUocart/a orthocarjm Greene, op. cit. 4:235.

1901.
AUocanja versicolor Brand. Rep. Spec. Nov.

19:71. 1923.

Stems branched from the base, the branches
prostrate, 1-3 dm long, straight, slender, and
somewhat wiry, thinly strigose or glaberate, of-
ten floriferous nearly to the base; leaves narrow-
ly linear, the lower 3-10 cm long, glabrous or
nearly so above, thinly strigose beneath, the
hairs mostly pustulate at base; racemes simple,
Ijecoming loosely flowered; mature calyx-lobes
usually accrescent, 3-8 mm long, about 1 mm
wide, connivent or sometimes spreading, more
or less definitely curved toward one side; corolla
1-2 mm broad; nutlets narrowly to broadly lan-
ceolate, acute; dorsal side keeled only above the
middle, more or less obliquely or transversely
rugose, smooth, granulate or penicillate-hairy;
ventral side keeled down to the basal scar, this
horizontal or slightly oblique, not surrounded by
a ridge, but frequently with a downwardly
directed dorsal flange. In heavy, usually alka-
line soils. Eastern Oregon to southern Idaho
and northern Utah, south to southern Califomia.
March-July. Fig. 104.

Fig. 104. Plagiobothrys leptocladus

Bo.\ Elder Co., 10 miles east of Bear River Refuge,
B. Maguire 13989 (BRY, UTC); alkaline flats west of
Perry, B. Maguire 3727 (UTC); 7 miles west of Brig-
ham City. H. Jensen s.n. (UTC); 9 miles east of Bear
I?iver Refuge, Maguire & Piranian 1.3985 (UTC); 4
miles northwest of Logan, B. Maguire 12966 (UTC); 4
miles west of Smithfield, B. Maguire 1.3782 (UTC); 4.5
miles west of Logan. B. Maguire 1.3781 (UTC); 3 miles
northwest of Logan, B. Maguire I2JJ65 (UTC).

80

Brigham Young Univehsitv Science Bulletin

4. Plagiohotlirtjs scouleri (Hook & Am.)
Johnst. Joum. Am. Arb. 16:192. 1935. Type:
Northwest Coast, Dr. Scouler.

Myasotis scouleri Hook & Am. Bot. Beechey

370. 1840.
AUocanja scouleri Greene, Pittonia 1:18.

1887.
AUocanja cusickii Greene, Pittonia 1:17.

1887.
AUocanja hispidula Greene, op. cit.
Allocanja penicillata Greene, op. cit.
AUocanja nitetis Greene, Pittonia 3:108.

1896.
Allocanja cognata Greene, Pittonia 4:235.

1901.
Plagiobothnjs scopuloruni Johnst. Contr.

Gray Herb. 68:79. 1923.
Plagiobothnjs lu^Isonii Johnst. Contr. Gray

Herb. 68:77. 1923.
Plagiobothnjs nitens Johnst. Contr. Gray

Herb. 68:78. 1923.
Plagiobothn/s hi^ndulus Johnst. Contr. Am.

Arb. 3:71. 19.32.
Plagiobothnjs cusickii Johnst. op. cit. 3:63.

1932.
Plagiobothnjs cognatus Johnst. op. cit. 3:59.

1932.

Plants more or less densely branched from
the base; stems 5-25 cm tall, ascending or

spreading, with stiff appressed hairs; leaves 1-8
cm long, linear to oblanceolate, strigose to some-
what setose; racemes rather lax, the bracts re-
sembling the leaves; calyx 1-1.5 (2) mm long
in flower, in fruit becoming 1.5-2.5 mm long;
corolla about 1.5 mm long, inconspicuous; nut-
lets 1.5-2 mm long, variously roughened, with or
without setose projections. Moist soils in sandy
to clayey areas. Widely scattered over the west-
em United States. May-July. Fig. 105.

Box Elder Co., Raft River Mountains, VV.P. Cot-
tarn 3051 (BRY); Raft River Mountains, S.J. Preece 752
(UT); Cache Co., % mile west of Logan Airport, B.
Maguire 2412 (UTC); Davis Co., Farmington Water-
fowl Refuge, B. Maguire 13905 (UT); Centerville, S.
Flowers 1193 (UT); Garfield Co., I'anguitch Lake, B.
Maguire 12958 (UTC); Aquarius Plateau, 19 miles
north of Escalante, N.H. Holmgren 2442 (BRY); Salt
Lake Co., Big Cottonwood Canyon, A.O. Garrett 1629a
(UT); Sanpete Co., 2 miles south of Indianola, B.F.
Harrison 10422 (BRY); Ephraim, W.P. Cottam 2783
(UT); Sevier Co., Fish Lake, mud flats, B. Maguire
129,57 (UTC); Koosharem Reservoir, W.P. Cottam
9472 (UT); Summit Co., north .slope of Big Mountain,
D.H. Galwav 2375 (BRY); north slope of the Uinta
Mountains, Welsh, Moore & Matthews 9171 (BRY);
Gorgoza, A.O. Garrett 2291 (UT); Snyderville, A.O.
Garrett 7758 (UT); Tooele Co., Pilot Mountain, W.P.
Cottam 1.3812 (UT); Uintah Co., Upper Bear River,
B.F. Harrison 10968 (BRY); Utah Co., east .slope of
Mount Timpanogos, B.F. Harrison 11447 (BRY); Ameri-
can Fork Canyon, A.O. Garrett 3800a (UT); Wasatch
Co.. summit of Daniels Canyon, A.O. Garrett 8050
(UT); Washington Co., Pine Valley Mountains, B. Ma-

UTAH

County Lines and
river drainages

i-

Fig. 105. Phgiobothrys scouleri

Fig. 106. Plagiobothnjs tenellus

Biological Series, Vol. 16, No. 3 The Boracinaceae of Utah

81

guire 12956 (UTC); Rock Spring, Pine Valley Moun-
tains, R. Hardy s.n. (DIX).

5. Phgiobothrys tenellus (Nutt. ) Gray, Proc.
Amer. Acad. 20:283. 1885. Type: Sunny
rocky slopes of the mountains along the
valley of Coeur d'Alene River, Idaho.
Myosotis tenella Nutt. Kew Journ. Bot.

3:295. 1851.
Plagiohothnjs parvalus Greene, Pittonia

3:261. 1898.
Plagiobothnfs asper Greene, op. cit. 262.
Tlagiohothrtjs liumifusus M. E. Jones, Contr.

West. Bot. 13:7. 1910.
Plagiohothnjs tenellus var. parvalus 5ubvar.

liumifusus Brand, Pflanzenreich 4,

2.52:108. 1931.

Stems slender, erect, freely branched from

the base or sometimes simple, 5-25 cm high, soft
villous with spreading and reflexed hairs; leaves
of the basal rosette oblong-lanceolate or oblong-
oblanceolate, obtuse or acutish, villous, 1-2.5 cm
long, cauline distinct, the lower ones linear-ob-
long, the upper becoming lanceolate or ovate-
lanceolate, gradually reduced in size; spikes
elongated in age and loosely flowered, only the
lowest flowers bracteate; calyx densely short-
villous with whitish or more often rufous hairs,
about 3 mm long in age; corolla limb about 2-3
mm broad; nutlets 1.5-2 mm long, thick cruci-
form, light colored, sharply ridged dorsally and
on the margins, the ridges commonly tubercu-
late. Grassy slopes and meadows. British Colimi-
bia and Idaho, south to Arizona, Utah, and
northern lower California. March-June. Fig. 106.

Salt Lake Co., near mouth of Big Cottonwood Can-
yon, A.O. Garrett 2233 (UT).

ACKNOWLEDGMENTS

I wish to extend my sincere thanks to Dr.
Stanley L. Welsh of Brigham Young University
for his encouragement over the last seven years
and for various suggestions in the preparation of
this work; and to Dr. Duane Atwood for his in-
terest and association on many collecting trips
and various trips to the herbaria in Utah and
elsewhere. Field work and publication costs
since 1966 have been supported by private fi-
nancing, I)v Brigham Young University, by Kill-
gore Research Center, and by West Texas State
Universitv. To these institutions I am most
grateful.

The help of the curators of the following her-

baria which have so graciously provided me
with assistance and the use of their facilities is
greatly appreciated.

BRY Brigham Young University, Provo,

Utah

DIX Dixie Junior College, St. George,

Utah

UTC Intermountain Herbarium, Utah

State University, Logan, Utah

UT Garrett Herbarium, University of

Utah, Salt Lake City, Utah

WTSU West Texas State University, Can-
yon, Texas

LITERATURE CITED

Abram.s, L. 19.54. Illustrated Flora of the Pacific States.
Vol, 111: Stanford University, California: Stanford
University Press.

Bahneby, R. 1943. Miscellaneous Diagnoses. Leaflets
West. Bot. 3:197.

Bentham, G. and ]. D. Hooker. 1876. Genera Plant-
arum. Vol. I: London: Reeve and Company.

Bi.ANKENSHip, ].W. 190.5. Supplement to the Flora of
Montana; Additions and Corrections. Mont. Agr.
Coll. Std. 1:96.

BHAniiUBY, J. 1817. Travels in the interior of America,
in the years 1809, 1810, 1811. including a descrip-
tion of Upper Louisiana, Tennessee, with the Illi-
nois and western territories, and containing re-
marks and observations useful to persons immi-
grating to those countries. Liverpool.

Brand, A. 1921. Borraginaceae-Borraginoideae. Cyno-
glosseae and Cryptantheae. Pflanzenr. IV. 252.
Heft 78, 1921 and Heft 97. 1932.

. 1923. Dacas Specierum Novarum Tertia. Fedde,

Rep. Spec. Nov. 19:70-73.

1927. Dacas Specierum Novarum Septima.

Fedde, Rep. Spec. Nov. 24:60-62.
Britton, D. M. 1951. Cytogenetic Studies on the Bora-

ginaceae. Britt. 7:233-266.
Candolle, a. p. de. 1846. Prodromus Systematis Na-

turalis Regi Vegetahilis. Vol. 10. Parisiis: Treutel.
Coulter, J. and A. Nelson. 1909. New Manual of

Botany of the Central Rocky Mountains. American

Book Company; New York.
Davis, R. J. 1952. Flora of Idaho. Dubuque, Iowa:

Wm. C. Brown Company.

82

BniGHAM Young Univkrsitv Science Bulletin

Eastwood, A. 19()3. New Species of OreoccLnju. Bull.
Torrey Club 30:238-246.

. 1913. in Rydberg, Studies on the Rocky Moun-
tain Flora XXIX. Bull. Torrey Club 40:480.

Ewan, J. 1942. A Review of the North American
Weedy Heliotropes. Bull. South. Calif. Acad. Sci.
41:51-57.

Graham, E. H. 1937. Botanical Studies in the Uintah
Basin of Utah and Colorado. Pittsburgh, Pa.:
Carnegie Institute.

Gray, A. 1875. Contributions in North American Bot-
any: Notes on the Boraginaceae. Proc. Amer. Acad.
10:48-62.

CJray, a. 1877. Contributions in North American Bot-
any: Characters of New Species. Proc. Amer. Acad.
12:80.

. 1878. Synoptical Flora of North America. Vol.

2, pt. 1. Cambridge: John Wilson and Sons.

1885. Contributions in North American Bot-

any: A Revision of some Boragineous Genera.
Proc. Amer. Acad. 20:257-286.

Greene, E. L. 1887. West American Asperifoliae. Pitt.
1:55-60.

. 1890. Reprint of Fraser's Catalogue. Pitt. 2:114-

119.

. 1896. New or Noteworthy Species. Pitt. 3:109-

115.

. 1901. Plantae Bakeriane 3:20. By the Author.

. 1902. New Species of Cryptantlw.' Pitt. 5:53-55.

Harrington, H. D. 1954. Manual of the Plants of
Colorado. Denver: Sage Books.

Hayden, F. V. 1870. Annual Report of F. D. Hayden,
in each of Twelve Annual Reports of the U.S.
Geological and Geographical Survey of the Terri-
tories. Washington, D.C.; Government Printing Of-
fice.

Hitchcock, C. L., A. Cronquist, M. Ownbey, and
J. W. Thompson. 1959. Vascular Plants of the
Pacific Northwest. Part 4, University of Washing-
ton, Seattle: University of Washington Press.

HiGciNs, L. C. 1968. New Species of Perennial Cryp-
tantha from Utah. Great Basin Naturalist 28:195-
198.

. 1969. New Combinations and a new species of

perennial Cryptantha. Great Basin Naturalist 29:29-
30.

. 1971. A Revision of Cn/ptantlui Subgenus

Oreocarya. Brigham Young University Science Bul-
letin, Vol. 13, No. 4. 63 pp.

Holmgren, A. H. 1948. Handbook of the Vascular
Plants of the Northern Wasatch. San Francisco,
California: Lithotype Process Company.

Holmgren, A. H. and J. L. Reveal. 1967. Checklist
of the Vascular Plants of the Intermountain Region.
U.S. Forest Service Res. Paper Int-32. Ogden, Utah.

Jackson, B. D. 1881. Guide to the Literature of Botany.
London: Longmans, Greene and Co.

Jackson, B. I). 1895. Inde.x Kewensis. Oxford: The
Clarendon Press.

Jepson, W. L. 1923. Flowering Plants of California.
University of California, Berkeley: University of
California Press.

John,ston, I. M. 1923. Studies in the Boraginaceae. II.
Restoration of the genus H<ickclui. Contr. Gray
Herb. 68:43-55.

. 192.3. Studies in the Boraginaceae. t. A Syn-
opsis and Redefinition of Pta^iohothrys. (^ontr.
Gray Herb. 68:, 57-80.

. 1924. Studies in the Boraginaceae. A synopsis

of the American Native and Immigrant Borages of

the subfamily Boraginoideae. Contr. Gray Herb.
70:1-54.

. 1925. Studies in the Boraginaceae IV: North

American species of Cryptantha. Contr. Gray Herb.
74:1-114.

. 1932. Studies in the Boraginaceae IX: The

AUocarya section of Plagwbothrijs in the western
United States. Contr. Am. Arb, 3:1-102.

. 1937. Studies in the Boraginaceae XII: Novel-
ties and Critical Notes. Joum. Am. Arb. 18:23-24.

. 1939. Studies in the Boraginaceae .XIII: New or

otherwise noteworthy species chiefly from western
United States. Joum'. Am. Arb. 20:390-399.

. 1940. Studies in the Boraginaceae XIV: Mis-
cellaneous species from Asia, Malaysia, and Ameri-
ca. Journ. Am. Arb. 21:48-66.

. 1948. Studies in the Boraginaceae XVII: Spe-
cies chiefly from Mexico and western United States.
Journ. Am. Arb. 29:240-241.

. 1952. Studies in the Boraginaceae XXII: Joum.

Am. Arb. 33:72-73.

1952. Studies in the Boraginaceae XXIII: A

survey of the genus Lithospermum . Joum. Arn.
Arb. 33:299-366.
. 1959. Some noteworthy Borages. Wrightia 2:13-

. 1961. Notes on some Texan Borages. Wrightia

2:156-162.
Jones, M. E. 1891. Contributions to Westem Botany:

New plants from .Arizona, Utah, and Nevada. Zoe

2:250.
. 1895. Contributions to Westem Botany VII:

Proc. Calif. Acad. Sci. II. 5:709-710.
. 1910. Contributions to Westem Botany: Contr.

West. Bot. 13:4-7.
Kearney, T. H. and R. H. Peebles. 1960. Arizona

Flora. Berkeley, Calif omia: University of California

Press.
Lanjow, y. and F. A. Stafleu. 1964. Index Herbar-

iorum. Part 1. 5th ed. Utrecht, Netherlands: Kem-

ink and Zoon.
. 1966. International Code of Botanical Nomen-
clature. Utrecht. Netherlands: Kemink and Zoon.
Lehmann, J. G. C. 1830. Novamm et Minus Cogni-

tarum Stirpium Pvigillus. Vol. 2. Publi.shed by the

author.
Linnaeus, C. 1753. Species Plantarum. Vol. 1. Holmiae:

Laurentii.

. 17.54. Genera Plantamm. Holmiae: Laurentii.

Macbride, J. F. 1916. The tme Mertensias of westem

North America. Contr. Gray Herb. 48:1-20.
. 1916. Revision of the Genus Oreocarya. Contr.

Gray Herb. 48:20-38.
. 1916. Notes on certain Boraginaceae. Contr.

Gray Herb. 48:39-59.
. 1916. Certain Boraginaceae new or transferred.

Proc. Amer. Acad. 51 :.541-.548.
. 1917. Further notes on the Boraginaceae. Contr.

Gray Herb. 48:16-22.
-. 1919. A revision of the North American species

of Anuinckui. Contr. Gray Herb. 49:1-16.
Matthews, V. B. 1968. A ta.vonomic study of Mertensia

(Bluebells) in Utah. Utah Acad. Proc. Vol. 45, pt.

2:. 590-602.
Moore, J. A. 19.36. Morphology of the gynobase in

Merten.sui. Amer. Midi. Natural. 17:749-758.
Munz, p. a. 19.55. California Miscellany. El Aliso

3:124-12,5.
Nelson, .\. 1898. New plants from Wyoming I. Bull.

Torrey Club 25:202-206.

Biological Series, \'ol. 16, No. 3

The Boiiac:ina{:eae of L' tah

83

. 1899. New species in Orcocanja and its allies.

Erythea 7:65-70.
. 1902. Contributions from the Rocky Mountain

Herbarium 111. Hot. Gaz. 34:30.
- — . 1913. Contributions from the Rocky Mountain

Herbarium XIU. Bot. Gaz. 56:69.
. 1934. Rockv Mountain Herbarium Studies II.

Amcr. Journ. Bot. 21:578.
. 1936. Rocky Mountain Herbarium Studies IV.

\mex. Journ. Bot. 23:269-270.

and P. B. Kennedy. 1906. New plants from the

Great Basin. Proc. Biol. Soc. Wash. 19:155-157.

and P. B. Kennedy. 1908. New plants from the

Great Basin. Muhlenbergia 3:141-142.

and J. F. M.\cdride. 1913. Western plant studies

I. Bot. Gaz. 55:378.

and J. F. Macbride. 1916. Western plant studie;

111. Bot. Gaz. 61:43.

and |. F. Macdkuje. 1916. Western plant studies

IV. Bot.' Gaz. 62:144-14.5.
NuTTALL, T. 1818. The genera of North American

Plants. Philadelphia: D. Heartt.
Ornduff, R. 1968. Inde.x to Chromosome Numbers for

1966. Regum Vegetabile. Vol. .55.
Osterhout, G. E. 1919. Additions to the Flora of

Colorado. Bull. Torrey Club 46:53-56.
. 1920. A new Orcocurya from Colorado. Bull.

Torrey Club 47:211.
. 1923. Two new plants from Colorado. Bull.

Torrey Club .50:217-218.

1926. Oreocarya. Univ. Wyoming Publ. Bot.

1:169.
Parish, S. B. 1898. New or little known plants of

southern California I. Ervthea 6:85-92.
. 1899. New or little known plants of southern

California II. Erythea 7:9.5.
Payson, E. B. 1915. New and noteworthy plants from

southwestern Colorado. Bot. Gaz. 60:380.
. 1926. Thalaspi, Oreocarya and Erigeron. Univ.

Wyoming Publ. Bot. 1:164-186.
-. 1927. A monograph of the section Oreocarya

of Cr>ptantha. Ann. Mo. Bot. Card. 14:211-358.'
Peck, M. E. 1941. .\ manual of the higher plants of

Oregon. Portland: Binfords and Mort.
Piper, C. V. 1902. Notes on the biennial and perennial

west American species of Lappula. Bull. Torrev

Club 29:535-549.
. 1906. Flora of Washington. Contr. U.S. Natl.

Herb. 11:9-637.

1920. A study of Allocarya. Contr. U.S. Natl.

Herb. 22: (2) 79-113.
Pritzel, G. A. 1872. Thesaurus Literature Botanicae.
Lip,siae: F. A. Brockhaus.

PuHSH, F. 1814. Florae Americae Septentrionallis. Vol.

2. London: White, Cochrane, and Co.
Reveal, J. L. 1968. On the names in Fra.ser's 1813

Catalogue. Rhodora 70:32.
Rydberg, p. a. 1905. Studies on the Rocky Mountain

Flora XIII. Bull. Torrey Club 31:637.
. 1906. Studies on the Rocky Mountain Flora

XVI. Bull. Torrey Club 33:150.'
. 1906. Flora of Colorado. Ft. Collins: Colorado

Agriculture College E.xperiment Station.
. 1909. Studies on the Rocky Mountain Flora

XX. Bull. Torrey Club. .36:678.
. 1913. Studies on the Rocky Mountain Flora

XXIX. Bull. Torrey Club 40: 480-481.
-. 1954. Flora of the Rockv Mountains. New

York: Hafner Publishing Co.
SuKSDORF, W. 1931. Untersuchungen in der Gattung

Amsinckia. Werdenda 1:47-113.
Tidestrom, I. 1913. Novitates Florae Utahensis. Proc.

Biol. Soc. Wash. 26:122.
. 1925. Flora of Utah and Nevada. Contr. U.S.

Natl. Herb. 25:1-665.

and KiTTELL. 1941. A Flora of Arizona and New

Mexico. Washington: The Catholic University of

America Press.
Torrey, J. 1827. Some account of a collection of plants

made during a journey to and from the Rocky

Mountains in the summer of 1820, by Edwin P.

(ames, NLD., assistant surgeon U.S. Army. Ann.

Lye. Nat. Hist. N. Y. 2:225.
. 1854. Botany in Marcy, Explorations of the

Red River Washington: U.S. Dept. of Interior.

1859. Botany in Emory, W., Report of U.S.

and Mexican Boundary Survey. Vol. 2. Washington:

U.S. Dept. of Interior.
Watson, S. 1871. Botany in King, C, Report of the

Geological Exploration of the Fortieth Parallel. Vol.

5. Washington: Government Printing Office.
Welsh, S. L., M. Treshow, and G. Moore. 1964.

Guide to the common LItah plants. Provo: Brigham

Young University Press.
Williams, L. O. 1937. A monograph of the genus Mer-

tensia in North America. Ann. Mo. Bot. Gard.

24:17-159.
Wooton, E. O. and P. C. Standley. 1913. Description

of new plants preliminary to a report on the Flora

of New Mexico. Contr. U.S. Natl. Herb. 16:166.
and . 191.5. Flora of New Mexico. Contr.

U.S. Natl. Herb. 19:9-794.
Wright, W. F. 1902. The genus Eritrichium in North

America. Bull. Torrey Club 29:407-414.

S-M''^(JOOq)

MUs.

Brigham Young University
Science Bulletin

'• Co MP.

''ffi 21973

ZOOL.

CONTRIBUTIONS TO THE STUDY
OF THE GENUS ERIASTRUM

II. Notes concerning the type specimens and
descriptions of the species

by
H. Keith Harrison

BIOLOGICAL SERIES — VOLUME 16, NUMBER 4
SEPTEMBER 1972

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsli, 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
Wilnier 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 UniversitN.

Orders and materials for library exchange should be directed to the Division of Gifts
and Exchange, Brigham Young University Library, Provo. Utah 84501 .

Brigham Young University
Science Bulletin

CONTRIBUTIONS TO THE STUDY
OF THE GENUS ERIASTRUM

II. Notes concerning the type specimens and
descriptions of the species

by
H. Keith Harrison

BIOLOGICAL SERIES — VOLUME 16, NUMBER 4
SEPTEMBER 1972

TABLE OF CONTENTS

INTRODUCTION

TAXONOMY

ACKNOWLEDGMENTS 25

LITERATURE CITED 26

CONTRIBUTIONS TO THE STUDY OF THE
GENUS ERIASTRUM

II. Notes concerning the type specimens and
descriptions of the species

by
H. Keith Harrison^

ABSTRACT

The species of the genus Eriastnim are
described. Keys to the species and subspecies
and the geographic ranges are provided. Dis-

cussions with regard to the type specimens or
lectotypes and interspecific relationships are
presented.

INTRODUCTION

The species of the genus Eriastnim are close-
ly related; and the range of variation found in
one species in branching, the lobing of the
leaves, vesture, etc., overlaps that found in other
species to a considerable degree. The species as
thev are now generally known were described bv
Mason ( 1945 ) . At that time, it was noted that
the proportions and size of the corolla and
stamens were important in identification of the
plants. Further study (Harrison 1959, 1968)
introduced new methods, emphasized the im-
portance of the relative size and proportions of
the corolla and androecium, and elucidated the
nature of these characteristics.

During the investigation of the species of
Eriastnim (Harrison 1959), it was found that
the type specimen for one of the species had
not been published and that other problems
concerning types and lectotypes require some
discussion. Inasmuch as nearly three decades
have pa.ssed since the first major paper on the
species of Eriastnim was published (Mason

1945), and as a later work (Harrison 1959) is
not generally available, it seems desirable to
include the information as to the type speci-
mens and synonomy for all the species, together
with the additions and discussions by the author.
It is understood that the first elements to be
found of a new species may represent either the
norm or one extreme in the variation of the
species. At a later time, other plants may be
collected which cause the investigator to create
a name for a subspecies, thereby segregating
certain specimens as being closely related to
(but still distinct from) the type upon which
the species is based. The fonnation of a sub-
specific category for the new element then ne-
cessitates, according to the International Code
of Botanical Nomenclature (Article 25) and
logic, the establishment of another subspecies to
include only those plants most representative of
the type— one could otherwise make no distinc-
tion between the type subspecies and speci-
mens identified only to the species level.

TAXONOMY

Eriastnim Wooton and Standlev

Hugelia Bentham, Bot. Reg. 19: under pi. 1622,
18.33, not Huegelia Reichenbach, Consp. 144.
1828. Wehvitschia Reichenbach, Handb. 194.
1837, not Welwitschia Hooker, Card. Chron.

'WcbiT State College, Ogden, Utah

22:71, nom. cons. Eriastrtim Wooton and
Standley, Contr. U.S. Nat. Herb. 16:160.
1913, Gilia and Navarretia of authors, in part.

Erect, or spreading and erect, annuals or
perennials, virgately, racemosely, corymbosely,

2 Bricham Young UNivEHsiTi- Science Bulletin

diffusely, or paniculately branched; herbage verform, regular or irregular; stamens ecjually

subglabrate to densely floccose; leaves linear, en- inserted at the base of the throat, on the throat

tire or pinnately lobed; bracteate floral heads, or just below the sinuses, equal, subequal, or

few or numerous, few- to many-flowered, rarely distinctly unequal; anthers sagittate, versatile;

solitary, lightly floccose to densely so with capsule ellipsoid, three-loculed, few-seeded;

arachnoid wool; calyx lobes unequal, joined seeds mucilaginous.

along the lower two-thirds of their length by a Type species: E. filifolium (Nutt.) Woot.

hyaline margin; corollas blue, yellow or white, & Standi.

sometimes cream, rarely pink, funnelform to sal- Range: Western North America.

Key to the Species

A. Plants perennial, woody, branching from the base; stamens inserted at the sinus

1. E. densifolium

AA. Plants annual, herbaceous, branching from the base or above; stamens inserted at the sinus
or below.
B. Corolla 15.0 to 23.0 mm long.

C. Stamens inserted at the sinus; corolla salverform, or very narrowly funnelform

— - 2. E. phdriflonim

CC. Stamens inserted at the base of the throat; corolla funnelform.

D. Corolla bilabiate, or irregular to obscurely so; stamens distinctlv unequal, fila-
ments often oriented toward lower lip, and anthers turned toward upper lip,
lavender blue; desert areas 6. E. erernicuni

DD. Corolla regular, or very nearly so; stamens usually equal, filaments not oriented
toward one side of corolla, bright blue; Monterey County 3. E. virgatiim

BB. Corolla 5.0 to 15.0 mm long.

C. Stamens 5.0 to 10.0 mm long, nearly equalling or exceeding tips of the corolla lobes;
anthers (restored)" 2.0 to 3.0 mm long.

D. Corolla bright golden yellow, 8.0 mm long, regular to very slightly irregular;
Monterey, San Luis Obispo counties 4. E. luteitm

DD. Corolla bright sapphire blue, ca. 1.0 to 1.5 cm long, distinctlv irregular to ob-
scurely so; southern California to Baja, 1000 ft to 9000 ft 5. E. sapphirinum

CC. Stamens less than 5.0 mm long, not equalling or exceeding tips of corolla lobes; an-
thers (restored) not more than 2.0 mm long.

D. Stamens equal; corolla lobes 1.0 mm broad, or less; corolla narrowly funnelform;
throat expanding slightly or not at all.

E. Filaments 2.5 to 3.0 mm long; floral heads distinctly "brushy" with tips of
bracts and calyx lobes conspicuous; ovary cylindric, three-times longer than
wide 7. E. filifolium

EE. Filaments 1.5 mm or less; tips of bracts and calyx lobes not conspicuously
prominent; ovary 1 to 1.5 times longer than wide.
F. Corolla 7.0 to 11.0 mm long; plants 15.0 to 35.0 cm high.

G. Leaves mostly 1.0 to 1.5 cm long and entire, or to 3.0 cm and with
one pair of lateral lobes; foliage often remaining somewhat soft; throat
expanding slightly 8. E. sparsiflorum

GO. Leaves mostly 1.5 to 2.5 cm long with one pair of lateral lobes, less
commonly entire; foliage becoming rigid and brittle; throat not ex-
panding.
H. Corolla lobes 3.0 to 3.5 mm long and 1.0 mm bro;id

. . 11. E. hrandegeae

'The addition of a drop of 10 percent liquid detergent restores dried .intliors tn the size .it the time of dehiscence.

Biological Series, Vol. 16, No. 4 The Genus Eriastrum 3

HH. Corolla lobes 2.5 mm long and 1.5 mm broad 12 E. traijci

FF. Corolla 5.0 to 8.0 mm long; plants to 15.0 cm high.

G. Leaves, when present, 2.0 to 4.5 cm long, with one to three pair of
lateral lobes to 1.0 cm or longer; floral heads many- flowered, very
densely floccose, often found under Adenostoma chaparral or on ob-
sidian rubble 13. E. ahramsii

GG. Leaves filiform, 0.5 to 2.5 cm long, entire or with one pair of lateral
lobes 2.0 to 5.0 mm long; floral heads few-flowered, somewhat floc-
cose, occurring in dry stream beds and on alkaline flats

_ 14. E. booveri

DD. Stamens unequal, corolla lobes 1.25 to 2.25 mm broad, corolla funnelform.

E. Plants robust to 30.0 cm high, racemose, corymbose, or branched from the
base; herbage floccose, often densely so; leaves 1.5 to 3.0 cm long, usually

with one or two pair of lateral lobes; floral heads several-flowered

9. E. tuilcoxii

EE. Plants to 15.0 cm high, diffusely branched from the base; stem and leaves
slender; herbage lightly floccose; leaves 1.0 to 2.5 cm, usually with one pair
of lateral lobes; floral heads few-flowered . 10 E. diffiistim

1. Eriastrum densijoUum (Benth. ) H. L. Mason,
Madrono 8:73. 1945.

Perennial, woody subshrub to 75.0 cm high,
branching from the base, erect, or spreading,
then erect; secondary branches erect, appearing
below the dead persistent heads; herbage light
green and hghtly floccose or subglabrate to dull
grey green and grey-canescent; stems leafy with
short inconspicuous intemodes or sparingly
leaved with intemodes to 4.0 cm long; leaves
sessile, 1.0 to 5.0 cm long, pinna tely parted with
one to five pair of lateral lobes or entire; leaf
rachis 1.0 to 3.5 mm wide; axils naked or with a
spur shoot or a canescent bud; heads terminal or
in a compact group; bracts as long as or far ex-
ceeding heads, with one to four pair of lateral
pinnae, less commonly entire; calyces 0.5 to 1.0
cm long, densely floccose to canescent; lobes un-
equal, joined by a hyaline membrane; corolla
funnelform, regular, 1.5 to 3.0 cm long; lobes
blue to lavender blue, elliptic to elliptic-
spatulate, 0.5 to 1.5 cm long and to 5.0 mm
broad; stamens adnate to sinus, equal, 4.0 to 8.0
mm long; filaments 2.5 to 6.0 mm long; anthers
sagittate, versatile, 2.5 to 4.0 mm long; stigma
1.0 to 1.75 mm long; ovary three-loculed, few-
seeded.

Key to the Subspecies

A. Herbage lightly floccose, subglabrous, or
glabrous; leaves light green.

B. Leaves entire or with one to two pair
of lateral lobes; intemodes very short;
spur-shoot in axils often 1.0 to 2.0 cm

long; corolla broadly funnelform; found
at elevations below 1000 ft, San Luis

Obispo to Santa Barbara County

. la. E. densifolium subsp. densijoUum

BB. Leaves with three to four pair of lateral
lobes, rarely entire; intemodes 1.0 to
1.5 cm; axils most commonly naked or
spur-shoot ca. 1.0 cm long; corolla nar-
rowly funnelform; found from 1500 ft
to 8000 ft, Monterey to Baja California,

Inyo County to Baja Califomia lb.

E. densijoUum subsp. austromontanum

AA. Herbage floccose, lanate, or canescent;
leaves grey green.

B. Well-developed spur-shoot often
present in the axils; corolla ca. .30.0
mm long; San Bernardino and

Orange County region

le. E. densijoUum subsp. sanctorum

BB. Short canescent buds often present
in axils; corolla ca. 16.0 to 18.0 mm
long.

C. Plants to 50.0 cm high; leaves 1.5
to 4.0 cm long; leaf rachis to 1.5
mm broad; Monterey to Baja Cali-
fornia, Inyo County to Baja Cali-
fomia

Ic. E. densijoUum subsp. elongatum

CC. Plants 20,0 to 30.0 cm high; leaves
1.0 to 2.0 cm long; leaf rachis 1.5 to
3.5 mm broad; Kem, San Bernardi-
no, Los Angeles County region .. Id.
E. densijoUum subsp. mohavensis

Brigham Young UNrvERsiTv Science Bulletin

Fig. 1. Eria.strum (Icii.sifoliiirn siilisp. dcnsifoliiim. ii. Habit. K. Harrison 250 VC, M141468. X 1.1. b. Habit. K.
Harrison 250 US Ml 41460. X .6.

Biological Series, Vol. 16, No. 4 The Genus Eria.strum

la. Eriastrtim densifolium (Benth. ) H. L. Ma-
.son subsp. dcn.sifoUttm.

Hugelia densifoUa Benth. Bot. Ree;. 19:

under PI. 1622. 1833.
Qilia huegelUi Steud., Norn. ed. 2, 1:683.

1840.
G. densijolUi Benth. in D.C. Prod. 9:311.

1845.
Navarretki densifoJia Kuntze, Rev. Gen.

PI. 2:433. 1891.
N. densifoUa Brand in Engler, Pflanzen-

reich IV, 250:165. 1907.
Welwitscltia densifoUa Tidestroin, Contr.

U.S. Herb. 25:429. 1925.
Hugelia densifoUa Benth. in Jepson, Man.

FI. PI. Calif. 792. 1925 in part.
alia densifoUa var. ft/pica (Benth.)

Craig. Bull. Torrey Club 61:390. 1934.
H. densifoUa Benth. in Jepson, FI. Calif.

3:161-162. 1943 in part.

Perennial, woody subshrub to 50 cm high;
major branching from the base, erect or spread-
ing, then ascending; secondary branches erect,
appearing at various distances below the dead,
persistent heads from the previous year; herbage
subglabrous or glabrate; stems leafy, stems may
be nearly hidden by the close-set leaves; inter-
nodes very short; leaver light green, 2.0 to 4.0
cm long, ascending, commonly entire or with
one or two (rarelv more) pair of short, lateral
pinnae 2.0 to 8.0 mm long, usually set toward
the basal portion of the leaf; the terminal lobe
thus long ( rarely, the outer pair of pinnae may
branch from the midpoint of the leaf), subulate,
awn-tipped; spur-shoots in the axils often well
developed, 1.0 to 2.0 cm long or longer, green;
the floral heads are terminal, or a compact
group of heads may be formed; the terminal
heads themselves are composed of a few smaller
heads in a very compressed inflorescence ap-
pearing as one structure; heads large, 2.0 to 4.0
cm in diameter and 1.5 to 2.5 cm in length, ex-
cluding flowers; in full flower, the tight cluster
of heads may make a mass of flowers 6.0 cm
across; bracts distinct to 2.0 cm long, exceeding
the calyces, with one or two pair of lateral pin-
nae set on the lower portion; the tenninal lobe
long, subulate, awn-tipped, lightly lanate to sub-
glabrous, light green contrasting with the white
wool surrounding the calyces; calyx 1.0 cm long,
covered with a mass of persistent, tangled, white
trichomes, calyx cleft in subecjual lobes; lobes
united two-thirds of their length by a hvaline
membrane, which margins the lobes nearly to
their tips; corolla bright blue; throat and tube

white or yellow; corolla to 2.5 cm long, broadly
funnel fonn; lobes to 1.1 cm long, elliptic to
elliptic-spatulate, to 5 mm broad; throat plus
tube to 1.25 cm long; tube puberulent; stamens
adnate to sinus, 8.0 mm long; filaments 6.0 mm;
anthere versatile sagittate, white, 3.5 mm long;
stigma 1.25 mm long; style 1.5 cm long; ovary
three-loculed; locules approximately six-seeded.

Type: David Douglas in 1833. KEW. Photo-
graph of type POM "188266; probable isotype
UC 163785. "California." Probably San Luis
Obispo County.

Range: San Luis Obispo and Santa Barbara
counties, California.

Eriastrum densifolium subsp. densifolium is
a very distinct subspecies and is limited in its
geographic range to the coastal region, center-
ing about San Luis Obispo. The large, broadly-
spreading, funnelform corolla, the large heads,
and the nearly glabrous, green, closely-set leaves
mark it as distinct. The closest affinities are
with E. densifolium subsp. austromontanum.

lb. Eriastrum densifolium subsp. austromon-
tanum (Craig) H. L. Mason, Madrono 8:74.
1945.

Gilia densifoUa var. austromontana Craig,
Bull. Torrey Club 61:391. 1934.

Huegelia densifoUa subsp. austromontana
Ewan, Bull. Torrey Club 64:520. 1937.

Huegelia densifoUa var. austromontana
Jepson, FI. Calif. 3:162. 1943.

Woody-based perennial to 30 cm high, erect,
or spreading, then erect; secondary branches
erect, axillary branches may be present; herbage
slightly floccose to subglabrous; stems leafy;
intemodes 1.0 to 1.5 cm long, rarely to 2.5 cm;
leaves light green, 1.5 cm long to 3.5 cm, ascend-
ing usually with three to four pair of lateral
pinnae, rarely entire, subulate-tipped; outer two
to three pair of pinnae, sometimes nearly equal-
ly spaced; the terminal lobe thus short; axils
without canescent buds, short spur shoots some-
times present in the axil, or axil naked; floral
heads commonly tenninal, heads small to medi-
um, 1.0 to 1.5 cm long (excluding flowers and
tips of bracts), to 3.0 cm long (including tips),
1.0 to 3.0 mm acro.ss (excluding tips of bracts),
to 4.0 cm across (including tips), moderately
woolly; bracts numerous, well developed, 1.5 to
3.0 cm long, exceeding the calyces by 1.0 to 2.0
cm in length, with one to four pair of lateral
pinnae; pinnae sometimes short or exceeding 1.0

'Abbreviations are in accordance with I.aniouw and Stafleu. Index Hcrbannruni (19o4).

Brigham Young Univehsity Science Bulletin

cm in length; calyx 6.0 to 8.0 mm long, cleft
into .subequal lolx^, hyaline margin joining the
lobes into a tube nearly the length of calyx,
calyx lobes narrow; corolla blue, narrowly fnn-
nelfonn, 16 to IS mm, rarely to 2.3 mm long;
lobes 5.0 to 7.0 mm long, elliptic to elliptic-
spatulate, 2.5 to 3.25 mm broad; tube plus throat
9 to 12 mm long, tube puberulent; stamens
adnate to sinus (or to 0.5 mm below sinus), 5.0
to 6.0 mm long; filaments 3.0 to 4.0 mm long;
anthers versatile, sagittate, white, 3.0 to 3.5
mm long; stigma 1.0 mm long; ovary few-seeded.

Type: P. A. Munz 8,341 POM 48414. Palo-
mar Mountains near Nellie, San Diego County,
California.

Range: Monterey County south to Baja Cali-
fornia and north to Inyo County, California, oc-
curring from 1.500 ft to over 8000 ft, and general-
ly over 4000 ft.

Eriastrum densifolhim subsp. austromon-
tanuin is closely related to the type subspecies.
Similarities may be noted in the branching
habits, the light green, lightly floccose or sub-
glabrate leaves, in the short intemodes, in the
absence of canescent buds in the axils, and in
the size of the heads. It differs in having more
elaborately lobed leaves and bracts and in the
slightly smaller and more narrowlv funnelform
flowers.

Ic. Eriastrum derviijolium subsp. eloiif^atiiin
(Benth.) II. L. Ma.son, Madrono 8:73.
1945.

Hugelia elongata Benth. Bot. Reg. 19:

under PI. 1622. 1833.
Gilia elonoaia Steud., Norn. ed. 2, 1:683.

1840.
NavarretUi iknisijolki subsp. elongata

Brand in Engler, Pflanzenreich IV, 250:

165. 1907.
G. dciViijoUa var. elongata (Benth.) Gray

ex. Brand, loc. cit.
H. denmfoUa Benth. in Jepson, Fl. Calif.

3:161-162. 1943 in part.

Perennial to 50 cm high, branching from the
woody base, erect or spreading, secondary
branches erect; herbage canescent (particularly
in the upper younger region), usually remaining
canescent in age, occasionally subglabrous; inter-
nodes 1.0 to 4.0 cm long; leaves grev green
ascending, 1.5 to 3.0 cm long, rarely to 4.0 cm
long, entire or pinnate, with one pair near the
ba.se of the leaf to four pairs of short lateral
pinnae; pinnae commonly less than 5.0 mm long.

occasionally to 1.0 cm long when three to four
pair of pinnae are present, the outer two to
three pair may be nearly equally spaced, mak-
ing the terminal lobe short and comparable to
the spacing between the outer pinnae; often
many axils show weak development of short
canescent buds; short axillaiy branches with
small heads are often present in the upper por-
tion; the numerous heads may Ix" terminal, or
the terminal branching may be racemose or
corymbose; heads small, 0.75 to 1.5 cm long, 1.0
to 2.0 cm across (excluding tips of bracts),
white-canescent; bract? 0.75 to 1.5 cm long, oc-
casionally to 2.0 cm long, merely equalling to
exceeding the calyces; bracts not always distinct,
as they may be half-hidden by dense, canescent
wool; calyces completely hidden by dense wool,
cleft into subequal lobes, 5.0 to 7.5 mm long;
lobes margined by a hyaline membrane uniting
the lower two-thirds of the calyx, subulate-
tipped; corolla bright blue or lavender blue;
veins may be marked by dark lavender blue,
throat and tube white or yellow, 14 to 18 mm
long, narrowly funnelform to salverform; lobes
to 7.0 mm long, elliptic to clliptic-spatulate, 2.0
to 3.0 mm broad; throat plus tube 8.0 mm to
1.25 cm long, tube puberulent; stamens adnate
to sinus, 5.0 to 7.0 mm long; filaments 3.0 to 4.0
mm long; anthers versatile, sagittate, white, 3.0
to 3.5 mm long; stigma 1.0 mm long, style 1.25
cm long; ovarv three-loculed, locules few-seeded.

Type: David Douglas in 18.33, "California."
KEW. Probably Monterey, San Benito, or San
Luis Obispo County.

Range: San Benito County south to Baja
California and north to Inyo County, California.

Eriastrum densifolium subsp. elongatum dif-
fers from the type subspecies in having weakly
deveIo[X'd white-canescent buds in the axils in-
stead of well-developed spur-shoots, and in hav-
ing lanate-canescent, grey-green herbage, par-
ticularly in the younger parts.

Id. Eriastrum densifolium subsp. mohavensis
(Craig) II. L. Ma.son, Madrono 8:74. 1945.

Gilin densifolia var. mohavensis C>aig,
iUilI. Torrey Club 61:392. 1934.

Hugelia densifolia var. moJmvensis Jep-
.son. Fl. Calif. 3: 162. 1943.

Perennial to 20 or 30 cm high, luncli
branched, dead heads |X'rsisting from the previ-
ous year, new branches arising below; stem in
older portions scaly and light CTeam-tan in color
as outer layers flake off; herbage, except the old

Biological Series, Vol. 16, No. 4 The Genu.s Ehiasthum

•Stem, canescent-lanate; stem somewhat leafy,
intemodes 1.0 cm long; leaves dull grey green,
1.0 to 1.5 cm long, occasionally to 2.0 cm long,
ascending or reflexed; leaf rachis 1.5 to 3.5 mm
broad, with one to four pair of very short lobes
2.0 to 3.0 mm long; lower ;ixils usually with
short canescent buds; heads few to numerous, to
200 in a plant 20.0 cm high, each branch produc-
ing several; heads small, ca. 1.0 cm long and 1.5
cm broad, excluding flowers; Ijracts with one to
three pair of very short spinescent lobes or teeth
or entire, canescent, often only the tip is visible;
calyces hidden by the dense mat of white wool;
calyx cleft into une(]ual lobes; lobes with a hya-
line margin forming a tube and covered with a
dense mat of long trichomes; corolla light or
lavender blue, 16 mm long, narrowly funnelfonn;
lobes to 5.0 mm long, elliptic-spatulate and 2.0
mm broad, throat 2.0 mm long, tube 9.0 mm
long, tulx" puberulent; stamens 4.0 mm long, fila-
ments 2.5 mm long; anthers versatile, sagittate,
2.75 mm long; stigma 1.25 mm long, style 12
mm long, locules few-seeded.

Type: T. Craig 1630. POM 1S212.3. Be-
tween Rosamond and Mojave, Kem County,
California.

Range: Mohave Desert; Kern, San Bernar-
dino, and Los Angeles counties, California.

Eriastrum densifoliuin subsp. mohavensis is
most closely related to E. densifoliuin subsp.
elonr^atum. The relationship may be noted by
the dull, grey-green leaves, the small canescent
heads and the short bracts. The fomier sub-
species differs in its shorter stature, shorter inter-
nodes and in the broader, shorter leaves which
may also be recurved.

le. Emistntm den.sifoUum subsp. sanctorum
(Milliken) H. L. Mason, Madrono 8:75.
1945.

Giliii densifoUa var. sanctora (Milliken)
Univ. Calif. Pub. Bot, 2:.39. 1904.

Hugelia densifolia var. sanctorum (Milli-
ken) Jep.son, Man. Fl. PI. Calif. 792.
192.5.

//. demifolia var. sanctorum (Milliken)
Jep.son, Fl. Calif. 3:162. 1943.

Woodv subshrub to 75 cm high; stems leafy;
herbage canescent, usually remaining so; leaves
dull grey green, ascending, to 5.0 cm long, en-
tire or with one to three pair of lateral pinnae on
th(? lower portion of the leaf; spur-shoots in the
axils well developed and short branches com-
mon; heads terminal or upper branches
racemose-corymbose; heads woolly; bracts to

2.5 cm long equalling or exceeding the heads;
calyces 1.0 cm long, cleft into unequal lobes;
corolla lavender blue; throat and tube yellow;
veins may be marked by deep lavender blue,
to .33.0 mm long, funnelform; lobes to 9.0 mm
long, elliptic to elliptic-spatulate, to 5.0 mm
broad; throat plus tube to 25.0 mm in length,
tube slightly puberulent; stamens adnate to
sinus or to 0.5 mm below, 6.5 to 8.0 mm long;
filaments 4.5 to 5.5 mm long; anthers versatile,
sagittate, white, 4.0 mm long and to 2.0 mm
broad; stigma to 1.75 mm long, style 25 to
.30 mm long, ovary three-loculed, each approxi-
mately five-seeded.

Type: H. M. Hall 683 and 17.3, UC 52454
and UC 2.3.521. Santa Ana River, Spanishtown
Crossing above Riverside, California.

Range: Santa Ana River, San Bernardino
and Orange counties, California.

Eriastrum densifolium subsp. sanctorum may
be recognized as a distinct subspecies by its
long corolla. The large spur-shoots in the axils
and the leafy stems suggest aspects of the type
subspecies. The grey-green, densely woolly herb-
age, the occasional heads with short bracts
and the woolly axillary buds suggest a relation-
ship with E. densifolium subsp. elongdtum.

2. Eriastrum pluriflorum (Heller) H. L. Mason,
Madrono 8:75. 1945.

Annual, 2.0 to 30.0 cm high or more, simple-
to densely-branched; herbage floccose to sub-
glabrous; leaves light green, to 5.0 cm long,
entire or pinnately divided with one to five pair
of lateral lobes; heads bracteate, lanate; bracts
1.0 to 3.0 cm long, pinnately divided with one to
four pair of lobes; calyx 0.5 to 1.0 cm long;
corolla regular, 16.0 to 23.0 mm long, blue,
or throat yellow, or tube and throat yellow;
stamens adnate to the sinus, equal or subequal;
anthers sagittate, versatile; stigma 0.75 to 1.0
mm long, style 11.0 to 16.0 mm long; ovary
three-loculed, locules few-seeded.

Key to the Subspecies

A. Plants 2.0 to 30.0 cm high, lobes of the
leaves to 2.0 cm long, corolla salverform or
very narrowly funnelfonn, lobffi elliptic 2.0
to 2.5 mm or occasionally to 3.0 mm broad;
hills bordering the San Joaquin Valley
2a. E. pluriflorum sub.sp. pluriflorum

AA. Plants 2.0 to 15.0 cm high, lobes of the
leaves to 8.0 mm long, corolla narrowly

BniGHAM Young University Science Bulletin

funnelform, tliroat expanding, corolla
lobes broadly elliptic to 4.0 mm broad;
Antelope Valley, Los Angeles and San

Bernardino counties

. 2b. E. j)Iurif.lomm subsp. sherinan-lioijtac

2a. Eriastrum pluriflorutn (Heller) H. L. Mason
subsp. pluriflorum.

Gilia virgata var. florihunda Gray, Proc.

Amer. Acad. 8:272. 1870. Not G. flori-
hunda Gray.
G. phiriflora Heller. Muhlenbergia 2:113.

1906.
Navarrctia virgata var. florihunda Brand

in Engler, Pflanzcnreich IV, 250:168.

1907. "
G. hrauntanii Jepson and Mason, Fl.

Econ. PI. Calif. 130. 1924.
Hugelia hrauntonii Jepson, Man. Fl. PI.

Calif. 793. 1925.
H. phiriflora Ewan, Bull. Torrey Club

64:520. 1937.
H. phiriflora (Heller) Ewan in Jepson,

Fl. Calif. 3:163. 1943 in part.

Annual, 2.0 to 30.0 cm or more high, and to
50.0 cm in breadth; erect, unbranched, or vir-
gate, racemose, or corymbose; branching above
or from the base; internodes 1.0 to 5.0 cm long;
herbage floccose to subglabrous; leaves light
green, to 5.0 cm long, subulate-tipped, pinnately
divided with three to five pair of lateral lobes,
rarely entire; lateral lobes to 2.0 cm long, spread-
ing; floral heads variable from 1.0 to 4.0 cm
long, broad, lanate, conspicuously bracteate;
bracts 2.0 cm to 3.0 cm long (exceeding the
heads), with three to four pair of lateral lobes,
lobes spreading slender, often to 1.0 cm long or
to 1.5 cm long, subulate-tipped; calyx 0.5 to 1.0
cm long, cleft into unequal lobes, joined bv the
hyaline membrane; flowers numerous; corolla
regular 16.0 to 23.0 mm long, salverfonn or very
narrowly fimnelfonn, far exceeding the heads;
tube long and slender, ca. 1.0 to 1.25 mm in
diameter; corolla bhie to lavender blue, or throat
vellow, or throat and tube yellow; veins of the
lobes often marked bv darker lines, tube pubeni-
lent; lobes 5.0 to 6.0 mm long and 2.0 to 2.5 mm
broad, occasionally to 3.0 mm broad; tube plus
throat 10.0 to 18.0 mm long; stiimens iidnate
just below the sinus, 3.0 to 5.0 lum long, ec|ual
or subecjual, the longest ef)ualling the tip of the
corolla lobe or nearly so; the anther base of the
shortest stamen occasionally may be onlv 0.5
mm above the sinus, filaments 1.5 to 4.0 mm
long; anthers sagittate, versatile, white or vellow
or purple, 2.5 mm long, 1.0 mm broad; stigma
0.75 to 1.0 mm long; s'tvle ca. 11.0 to 16.0 mm

long; ovary three-loculed, approximately three-
seeded.

Lectotype: W. H. Brewer 1212. GH. Corral
Hollow, Alameda ("ounty, California.

Range: West Central Calif oniia, hills bor-
dering the San Joa(juin Valley from Contra
Costa County south to Santa Barbara County,
east to Fresno, Tulare and San Bernardino comi-
ties, California.

The plants described by Gray as Gilia virgata
var. florihunda were based upon California col-
lections bv Fitch, Wallace, and Brewer (Gray
1870). Jepson (1943) felt that the collection by
Fitch, Ix^ing the first cited by Gray, could well
be taken as the tvpe, and so indicated it. In the
Stinoptiral Flora' (1878, 1886), Gray listed the
collection as Wallace, Brewer, and Palmer, omit-
ting the Fitch collection. In the herbarium of the
University of California, there is a collection by
W. H. Brewer 1212, June 3, 1862, taken while
he was engaged on the state survey, from Camp
72, Corral Hollow, Alameda County. California,
that is inscribed by Jepson as the tvpe. In 1905,
A. A. Heller found Gilia virgata var. florihunda
abundant on the dry hillsides near Sunset, Kern
County, California, and listed two collections:
77.34 from Sunset and 7742 from Oil City. He
noted that the varietv was sufficiently distinct
from the species with which it had been as-
sociated to warrant raising it to specific level.
Since Gilia florihunda. Gray was already occu-
pied; he chose another name descriptive of the
numerous flowers, that of Giliu phiriflora. The
Heller collection 7734 was subse(|uently taken to
be the tvpe. and Sunset, Kern County, as the
type locality (Craig 1934). Heller based his
species on Gilia virgata var. floril)iinda: and it is
evident that he did not intend his collection to
be construed as the type, for he was explicit
whenever he referred to a tvpe (Heller 1906,
Ewan 1937).

An isotype of Heller 7734 at UC is adecjuate,
and the Heller 7742 (not mentioned in connec-
tion with the type) is an excellent specimen.
However, in accordance with custom, it is ap-
propriate to refer to the Fitch, Wallace, or
Brewer collection for the type. The Fitch col-
lection has no information ;is to where or when
it was collectic'd other than "C:alif. Fitch." The
specimen is a fragment with very few flowers,
which could be regarded as adec|uate were there
no other selections available. The Wallace col-
lection is much better, with m;mv flowers; and
the locaticm is given as Tejon. The Brewer col-
lection is satisfactory; adc<juate information is
given with respect to location; and specimens of

Biological Series, Vol. 16, No. 4 The Genus Ehi.\strum

the collection are known at Gray, University of
California, and Jepson Herbaria. Of these, the
best is one of those at the Universit)' of Cali-
fornia. This is the specimen inscrilied by Jepson
as being the tvpe and is annotated l)y Craig as
"Gilia phiriflortim Heller, type of Gilki vir<iiit(i
var. jlorihunda Gray"; still. Gray did not cite
this specimen as type. It seems appropriate to
select the Brewer collection 1212 (Gray Her-
barium) from Corral Hollow, Alameda County,
California, and \\'hich is annotated by Gray as
"Gilm (Hugelia) virgata var. jlorihunda," as a
lectotype.

Eria.^lrum phiriflonnn is a very variable spe-
cies with regard to size and branching habit.
Plants inav flower when onlv 2.0 cm high. They
inav "row strictlv vireate to 15.0 or 30.0 cm or
more with one stem only and one terminal head.
Branching mav be virgate. racemose, corymbose,
or paniculate. The plants often po.ssess one stem
below that becomes branched in the upper por-
tion; or branching occurs from the ba.se, produc-
ing a small plant with a cluster of branches or a
verv denselv branched shrubby plant to 50.0
cm or more in diameter.

Probablv the closest connection between the
perennial and the annual species lies between
E. dcnsifolium subsp. austromontanum, and E.
])Jurifloiuiii. Characters which link these entities
are the light green, lightly floccose or sub-
glabrous herbage, the leaves and bracts with
2-5 pair of lobes, the bracteate-lanate heads, the
long, slender corollas and the stamens inserted
at the sinus. The more elaborately lobed leaves
and bracts, the long salverform corolla and the
insertion of the stamens at the sinus, distinguish
E. j)hiriflonim from all of the other annual spe-
ries of Eriastrum: it is perhajis the most easily
recognized of the annual species. Another close
relationship exists between this species and E.
eretnicinn. This relationship will be discussed in
connection with the latter species.

2b. Eriastrum plurijlomm subsp. slierman-
liot/tae (Craig) H. L. Mason, Madrono 8:75.
1945.

Gi7i« slienmin-hotitac Craig, Bull. Torrev

C:lul) 61:41.5. 1934.
Hugelia plurijlora (Heller) Ewan in Jep-

.son, Fl. Calif. 3:163. 1943 in part.

Annual, 3.0 to 15.0 cm high, simple to corym-
boselv branched; herbage floccose to subgla-
brous; leaves green 1.0 to 4.0 cm long, entire or
pinnately divided with one to four pair of lateral
lobes; lobes to 8.0 mm long, often reduced to
teeth; floral heads approximately 1.0 cm broad,

lanate; bracts 1.0 to 2.0, occasionally to 3.0 cm
long, pinnatelv divided with one to three pair
of lobes, exceeding the heads; calyx 5.0 to 7.5
mm long, cleft into unequal lobes; corolla regu-
lar, to 20.0 mm long, narrowly funnelfonn; tube
long and slender; throat expanding, blue or yel-
low, or throat and tulx; yellow, lobes broadly
elliptic to 6.0 mm long and to 4.0 mm broad; sta-
mens adnate just below the sinus or as much
as 0.75 mm below, ecjual or subecjual, 2.0 to
4.0 mm long; filaments 1.0 to 3.0 mm long; an-
thers sagittate, versatile, 1.5 mm long; stigma 1.0
mm long, style ea. 14.0 mm long, ovary three-
loculed.

Type: P. A. Munz and T. Craig 12925. POM
18.5022. Ten miles south of Muroc, Los Angeles
County, California.

Range: Antelope Valley, Los Angeles and
San Bernardino counties, California.

This subspecies differs from the type sub-
species in the short stature of the plants and the
small heads. The corolla lobes are broader and
the corolla more funnelfonn. The stamens are
shorter and are sometimes inserted 0.25 to 0.75
mm below the sinus. The leaves have fewer and
shorter lobes. This subspecies is known from
Antelope Valley near Lancaster and Muroc,
which is near the southern limits of the species.

3. Eriastrum virgatum (Benth.) H. L. Mason,
Madrono 8:84. 1945.

Hugelia virgata Benth. Bot. Reg. under PI.

1622. 1833.
Cilia virgata Steud., Norn. ed. 2, 1:684.

1840.
Navarretia virgata Kuntze, Rev. Gen. PI.

2:4.3.3. 1891.
N. virgata Brand in Engler, Pflanzenreich

IV, 2.50:167. 1907.
N. deihsifolia var. lanata Brand, op. cit.

165.
G. virgata var. ttjpira C;raig, Bull. Torrey

Club 61:.394. 19.34.

Annual, erect, 3 to 40 cm high, occasionally
to .50 cm high, virgate. or racemosely branched
;ibove, or branching from the base, then ascend-
ing virgatelv; herbage light green, usually lanate
to lightlv floccose, sometimes subglabrous, but
remaining woolly on the axillary buds and heads;
leaves linear, filifonn, ascending, often some-
what appressed, 2.0 to 5.0 cm long, entire or
with one pair of lateral pinnae near the base of
the leaf, rarely with two pair; lateral pinnae
slender, to 12 mm long; heads woolly, 1.25 to
2.0 cm long and 1.0 to 3.0 cm broad, including

10

Brigham Young University Scienck Buli.etin

Fig. 2, F.riatrum virgatum, a. Habit. X 1.6. h. Corolla and stamens. X 2.3 c. Calyx, X 2.3. d. Habit, K. Harrison
237 UC Ml 41 467, X .6.

Biological Series, Vol. 16, No. 4 The Genu.s Ehia.strum

11

the bracts; central mass usually 0.75 cm to 2.0
cm broad, few-flowered; bracts 1.0 to 2.5 cm
long exceeding the heads, usually with one pair
of lateral lobes; calyx lobes unequal, to 9.0 mm
long, joined by a hyaline membrane, subulate-
tipped; corolla fuimelform, regular or yery near-
ly so, 15.0 to 22.0 mm long, bright blue with
yellow throat and tube; lobes 6.5 to 9.5 mm long
and to 3.0 mm broad; throat plus tube 8.0 to 13.0
mm long; stamens e(jual or sul>equal, inserted
1.0 to 2.0 mm below the sinuses, rarely less, and
extending to or nearly to the tip of the corolla
lobe, 6.0' to 11.0 mm' long, filaments 5.0 to 9.5
mm long, anthers linear-sagittate, versatile 2.5 to
3.0 mm long and 0.5 to 1.0 mm broad; stigma
0.75 to 1.0 mm long, style approximately 15.0
nmi, oyary three-loculed, few-seeded.

Type: Dayid Douglas in 1833. Photograph
of type POM 188883; probable isotype UC
163784. "California." Probably Monterey County.

Range: Monterey County, California.

The heads are relatively few-flowered; and
the individual flowers, with their bright blue
lol>es and yellow throats, are particularly showy
in E. vir^atuin and in E. sajiphi.rinum. Eriastrum
virgatum is limited in its geographic range and
is far from being the most common of the annual
species. Close relationships are with E. luteum
and E. sapphirinum subsp. clasyontJmm.

4. Eriastrum hiteum (Benth.) H. L. M;ison,
Madrono 8:81. 1945.

Htigelia lutea Benth. Bot. Reg. 19: under

pi. 1622. 1833. Not Gilia lutea Steud.
Cilia lutescem Steud. Nom. ed. 2, 1:684.

1840.
G. floccosa Kuntze, Rev. Cen. PI. 2:4.33.

1891 in part.
Navarretia hitescens Kuntze, loc. cit.
N. lutea Brand, Pflanzenreich IV. 250:

168. 1907.

Annual, to 25.0 cm high, erect, virgate or
racemose or branched from the base; herbage
lightly to densely flcK-cose; stems and lower
leaves often red brown, upper leaves ;ind bracts
light green; intemodes 1.0 to 2.0 cm long; leaves
to 4.0 cm long, ascending, entire or with one pair
of lateral lobes at the base of the leaf, and that
which appears to be a pair of lateral lobes often
is ;i p;irt of the axillary bud; lateral lobes spread-
ing, to 1.0 em long; floral heads to 1.5 cm bro;id.
densely floccose-lan;ite; bracts 0.75 to 1.5 cm
long, rarely more, with one or two pair of lateral
lobes; calyx lobes nearly e(|u;il 5.0 mm long,
joined by a hyaline membrane; corolla reguhir or

very slightly irregular, funnelform, 8.0 mm long,
bright golden yellow; lobes 3.5 mm long, often
obovate, 2.0 to 2.5 mm broad; throat 1.5 mm
long, tube 3.0 mm long; stamens equalling or
exceeding tips of corolla lobes, 5.0 to 5.5 mm
long; filaments 3.5 to 4.5 mm long, anthers 2.0
to 2.25 mm long and 0.75 mm broad; stigma 0.5
mm long, style ca. 7.5 mm long; locules one-to
two-seeded.

Type: David Douglas in 1833. KEW. "Cali-
fornia." Probably Monterey. San Luis Obispo
County.

Range: Santa Lucia Mountains, Monterey
and San Luis Obispo counties, California.

The bright, golden-flowered species, E.
luteum. which is found somewhat infrequently,
seems to be most closely related to E. viraatum.
The corolla form and proportion ;ind stamen
size are approximately the same; the obovate
loljes of the corolla, the branching, plant-size,
leaves and pubescence of E. luteum are found
in E. virgatum and the geographic range over-
laps. Cytological studies will probably yield
further insight into these relationships.

5. Eriastrum sapphirinum (Eastwood) H. L.
Mason, Madroiio 8:79. 1945.

Annual, erect to 40.0 cm high and to 30.0
cm broad; branching open, paniculate, corym-
bose, racemose, or simple, frecjuently branching
from nearly every axil; branches usually slender,
rarely appearing below dead persistent heads
and flowering continuing into a second season;
herbage with minute glandular hairs throughout
or in part, otherwise glabrous to floccose; stems
sparsely leaved; intemodes 1.0 to 5.0 cm long;
leaves 0.5 to 3.0 or to 5.5 cm long; filiform-
linear, entire or with one pair of slender lateral
lobes at leaf base, subulate, or awn-tipped; heads
few to numerous, pedicelled or ses.sile, one- to
many-flowered, to 1.5 cm broad, glabrous,
glandular, to densely floccose; bracts broad and
short, 3.0 to 10.0 mm long, more or less eciual to
the head; lowermost, subtending the head, not
apparent or to 2.0 cm long and exceeding head,
simple or with one to two pair of short and often
broad lateral lobes; hy;iline membrane between
the lobes frequently apparent; calyx lobes un-
equal to nearly ec(ual. 6.0 to 8.0 nmi long, joined
by hyaline membrane; corolla broiidly funnel-
form to narrowly so, to 15.0 mm long, irregular;
three lobes occasionally appearing longer than
the other two due to une(|ual sinuses, especially
in the broad corolhis, less markedly so in nar-
rowly funnelfonn type; lobes broadly to narrow-

12

BniGHAM Young Univehsity Science Bulletin

Fiji. 3. Eriaslrurn lutciiiii. a. Ilahit. I)a\icl Douglas VC 163780, X .6. 7>. Habit, H. F. Hoover 7125 UC 023830,
X .6. c. Habit. Kathcriiif Braii.lcgef 136. 9VI-09. H0858. X .6. ,1. Calyx. G. T. Nordstrom 1353 UC 123876,'
X 3.4. e. Corolla and staineii.s, G. T. Nordstrom 1353 UC 123876. X 3.4. /. Habit. G. T. Nordstrom 1353 UC
123876, X .6. /?. Habit, R. F. Hoover 7192 UC 023829. X .6.

Biological Series, Vol. 16, No. 4 The Genu.s Ehiasthum

13

ly elliptic, half the length of corolla or more, to
9.5 mm long and 2.5 to 5.0 mm broad, bright
blue; throat 1.0 to 3.0 mm long, yellow; tube to
5.5 mm long or 1/5 to 1/3 length of corolla,
blue or yellow; stamens adnate to bottom of
throat, usually equal, to 10.0 mm long, extend-
ing nearly to tip of corolla lobe, occasionally be-
yond, anthers sagittate, versatile, 2.5 to -5.0 mm
long, 1.0 mm broad; filaments to 8.0 mm long;
ovary three-loculed; seeds 2.0 to 2.5 mm long.

Key to the Subspecies

A. Heads single- to few-flowered, 2.5 to 8.0
mm broad, seldom more; stems slender,
bracts subequal to or slightly exceeding the
heads; corolla often broadly funnelform,

lobes broadly elliptic -

... 5a. E. sapphirinum subsp. sapphirinum

A A. Heads several-flowered, 0.75 to 1.5 em
broad or more; stems slender to somewhat
robust; bracts commonly exceeding the
heads, often prominent; corolla mostly nar-
rowly funnelform, lobes narrowly elliptic
5b. E. sapphirinum subsp. dasijanthum

5a. Eriastrum. sapphirinum (Eastwood) H. L.
Mason subsp. sap])hirinum.

Gilia sapphirina Eastwood, Bot. Gaz. 38:

71. 1904.
Navarretia virgata sub.sp. gynmocepJuila

Brand in Engler, Pflanzenreich IV, 250:

168. 1907.
N. virgata subsp. gtjmnocephala var. oli-

gantha Brand, op. cit.
N. virgata subsp. gt/mnocepljalxi var. sap-
phirina Brand, op. cit.
G. floccosa var. ambigua Jones, Contr.

West. Bot. No. 13:2. 1910.
G. virgata var. sapphirina MacBridc,

Contr. Gray Herb. No. 49:58. 1917.
Hugelia virgata var. sapphirina Jepson,

Man. Fl. PI. Calif. 793. 1925.
G. virgata var. ambigua Craig, Bull. Tor-

rey Club 61:412. 1934.
//. virgata var. ambigua (Jones) Jepson

in Jepson, Fl. Calif. 3:165. 1943.
Eriastrum sapphirinum subsp. gijmno-

cephaUim Mason. Madrofio 8:80. 1945.
E. sappliirinum subsp. ainbiguum Mason,

op. cit.

Branches slender; leaves 0.5 to 2.0 cm long,
occasionally to 3.5 cm long, filiform-linear, sub-
terate, entire or with one pair of lateral lobes at
the leaf base; lateral lol>es 2.0 to 5.0 mm long;
heads small, few to a few hundred, pedicelled or

sessile, single- or few-flowered, 2.5 to 8.0 mm
broad, seldom more, glabrous, glandular, to floc-
cose; bracts 3.0 to 10.0 mm long, more or less
equal to the head, the lowennost subtending the
head often not apparent, occasionally exceeding
the head; corolla to 15.0 mm long, broadly fun-
nelform; lobes often broadly elliptic.

Type: Blanche Trask in 1903. CAS 510. San
Jacinto Mountain, Biverside County, California.

Range: Southern California from Los Ange-
les and San Bernardino counties to Baja Cali-
fornia, occurring between 1500 and 8700 feet.

Navarretia virgata subsp. gijmnocephalurn
was described by Brand ( 1907 ) and implied in
the name was the naked head. Its variety oli-
gantha was characterized as having flowers
single or in pairs. The type specimen of E.
sapphirinum has perhaps 1.50 to 175 heads which
are naked or glandular, pedicellate or sessile and
one- to five-flowered. Eriastrum sapphirinum
subsp. gymnocephalum is regarded here as a
synonym of the type subspecies.

The degree of pubescence in the heads of
E. sapphirinum subsp. sapphirinum varies from
being completely naked to lightly floccose. The
type of E. sappliirinum subsp. ambiguum is not
particularly unlike the type subspecies, and the
only character by which it might differ is the
degree of pubescence. Jones described his speci-
men as being "floccose-woollv mostly through-
out" and the type of E. sapphirinum subsp. am-
biguum (Jones 10011 Victor [now Victorville],
California POM 75010) is almost completely gla-
brous. Except for the minute glandular hairs
and the floccose heads, the plant is not at all
floccose. In his descriptions, Jones cited both
10011 from Victorville and 9917 from Bear Val-
ley as being typical. The latter specimen was
reported by Craig ( 19.34 ) as being "not at all
typical," and Mason (1945) referred the speci-
men to E. sapphirinum subsp. dasijanthum. It is
this specimen that is floccose-woolly through-
out. Eriastrum sapphirinum subsp. ambiguum
is regarded here as insufficiently distinct to war-
rant recognition and is regarded as a synonym.

With a large representation of E. sapphiri-
num, a continuous series can be observed. The
size of the heads varies from 2.5 to 15.0 mm or
so with every intergradation. The slender plants
with small heads have a tendency to have broad-
ly spreading and broadly-lobed corollas which
are distinctly irregular; and those plants with
larger heads have a tendency to have more nar-
rowly funnelfonn, less irregular corollas. The
series suggests two entities within the species
with complete interbreeding. The subspecies

14

Brigham Young University Science Bulletin

Fig. 4. Eriastrurn sapphirinum subsp. sapplurintitii. a. Habit, Blaiuhe Trask in 1903. CAS 510 (type). X .6. b.
Two flowers, Katlierine Brandegue 16-VI-()6, VC 1,3 5928. X 2.3, c. Floral head. C. Epling. M. Darsie. r. Knox,
William Robison 20-VI-32, UC 519836. X 2.3.

Biological Series, Vol. 16, No. 4 The Genus Eriastrum

15

overlap almost completely in geographic distri-
bution. The corollas of the tvpe subspecies
have three lobes, which appear longer than the
others due to the unequal sinuses. The lobes
are broadlv elliptic and extend from 1/2 to 2/3
the length of the corolla; the throat is spreading
and prominent; and the tube is approximate-
ly 3.0 mm in length, or 1/5 of the length of the
corolla. Corollas vary from this to the less ir-
regular fonn of E. sa})pliirinuin subsp. dasijan-
thum, in which the slightly expanding throat
and narrowly elliptic lobes are usual. Cytologi-
cal studies are needed to elucidate the nature
of the subspecies.

5b. Erui^lnim sappliirintim subsp. dasijanthtim
(Brand) H. L. Mason, Madroiio 8:S0. 1945.

Nava.rretia virgata var. dasijantha Brand

in Engler, Pflanzenreich IV, 250:168.

1907. ^
Htigelia virgata var. dasyantha Jepson.

Man. Fl. PI. Calif. 793. 1925.
Gilia virgata var. dasyantha Craig, Bull.

Torrey Club 61:395. 1934.

Plants slender to somewhat robust; leaves 1.0
to 3.0 cm long, occasionally to 5.5 cm long, entire
or with one pair of lateral lobes; lateral lobes
2.0 to 15.0 mm long; heads 7.5 to 15.0 mm broad,
several-flowered, denselv floccose; bracts usually
exceeding the heads; bracts subtending the
heads to 2.0 cm long, often prominent; corolla
to 15.0 mm long, more commonly narrowly fun-
nelfomi; the lobes narrowly elliptic.

Type: H. M. Hall 298 6-VIII-96. Swartout
Canvon, San Antonio Mountain, elevation 7000
feet; within the San Gabriel Mountains, Los
Angeles County, California. Present repository
of the specimen is unknown. Also cited in the
original description as representative of the type
are: Parry and Lemmon 249 (represented
POM), S. B. and W. F. Parish 1478 (represent-
ed UC), S. B. Parish 3803 (represented UC).
The specimen, H. M. Hall 237, San Antonio
Mountain (elevation 7000 feet), Los Angeles
County. California (UC), is cited in the H. M.
Hall fieldbook (UC) as being "identical" with
his 298 collection.

Range: Southwestern California from Ven-
tura County, south to Baja California and north-
east to Inyo Countv, California, occurring from
1000 to 7000 feet.

6. Eriastrum eremicum (Jepson) H. L. Mason,
Madrono 8:78. 1945.

Annual, to 30.0 cm high and to 40.0 cm broad,

erect or spreading, branched from the base or
racemose, or virgate; stems floccose to subgla-
brate, sometimes glandular; leaves light green or
grey green, 0.6 to 5.5 cm long, with one to three
pair of lateral lobes, subulate, awn-tipped; heads
one to many, floccose, 0.5 to 1.5 cm broad, ex-
cluding tips of braotis; bracts 1.0 to 2.0 cm long
with one to three pair of lateral lobes; calyx 0.5
to 1.0 cm long; lobes unequal, joined by a hya-
line membrane; corolla bilabiate or irregular to
nearly regular, funnelfonn, 11.0 to 18.0 mm
long; lobes elliptic, 4.0 to 8.0 mm long and 1.75
to 3.5 mm broad, lavender blue to dark blue;
throat 1.0 to 4.0 mm long, sinus to stamen inser-
tion 1.0 to 4.0 mm, tube 4.0 to 8.5 mm long; tube
and throat light lavender, yellow, or blue; sta-
mens adnate to base of throat, unequal, 2.0 to
9.0 mm long; filaments 1.5 to 8.5 mm long; an-
thers sagittate, versatile, 1.0 to 2.2.5 mm long and
1.0 mm broad; stigma ca. 1.0 mm long, style ca.
12.0 mm long, ovary three-loculed.

Key to the Subspecies

A. Corolla bilabiate to obscurely irregular;
lobes narrowly elliptic 1.75 to 2.75 mm
broad, lavender blue; desert regions south-
ern California, Nevada, Utah, and Arizona
6a. E. eremicum subsp. eremicum

AA. Corolla irregular to nearly regular; lobes
broadly elliptic, 3.0 to 3.5 mm broad, blue

to dark blue; Arizona

6b. E. eremicum subsp. yageri

6a. Eriastrum eremicum (Jepson) H. L. Mason
subsp. eremicum.

Nava.rre'tia densifolia var. jacumbarui

Brand, Ann. Conserv. and Jard. Bot.

Geneve 15 and 16:340. 1913.
Hugelia eremica Jepson, Man. Fl. PI.

Calif. 793. 1925.
Gilia eremica Craig, Bull. Torrey Club

61:416. 19.34.
G. eremica var. typica Craig, op. cit. 417.
G. eremica var. zionis Craig, op. cit. 418.

Annual, erect or spreading, much-branched,
to .30.0 cm high and to 40.0 cm across; stems
lightly floccose to nearly glabrate, often wiry or
thick, becoming dark red or becoming very light
with age as epidennal layers flake off; leaves
grey green, 6.0 to .30.0 mm long, pinnatifid with
one to three pair of lateral pinnae, commonly
with two pair; when three pair occur, the lower-
most pinnae are shorter than the adjacent pin-
nae, lightly floccose to glabrate, subulate-Hpped;
pinnae 2.0 to 10.0 mm long; heads small, lanate,

16

Bricham Young Univkhsiiy Science Bulletin

numerous (2-2.50), few-flowered (2-15); bracts
10.0 mm long, with one to two pair lateral lobes,
lanate, recurved, subulate-tipped; calyx 6.0 to
7.0 mm long, sul)efjual; caJyx lobes connected
along two-thirds of the lower margin by a mem-
branous tissue; corolla irregular or obscurely so,
from bilabiate to nearly regular, 11.0 to 18.0 mm
long, the upper lip may consist of either three or
four lobes; lobes 4.0 to 8.0 mm long, differing
in length in a particular flower by 0.50 mm to
1.75 mm due to unequal sinuses, sometimes
asymmetrical with one margin much more con-
vex than the other, 1.75 to 2.75 mm broad; throat
1.0 to 3.5 mm long; sinus to stamen insertion
distance differing in one flower by 0.50 to 2.0
mm, and usually by 1.0 mm or more; tube 4.0
mm to 8.5 mm long, usually less than half the
length of the corolla; inner surface of the tube
puberulent; corolla light lavender to violet,
prominent reddish lines follow veins of lobes;
stamens 2.0 to 9.0 mm long, distinctly unequal in
length, inserted at base of the throat, 1.0 mm to
.3.5 mm below the sinuses; in conjunction with
the corolla shape, the stamen filaments are very
often inclined toward the lowermost lobe with
the anthers turned toward the upper lip; anthers
sagittate, 1.5 mm to 2.5 mm long, filaments 1.5
mm to 8.0 mm long; ovary three-loculed, few-
seeded.

Type: W. L. Jepson .5414. JEPS. Calico
Wash, northeast of Barstow, San Bernardino,
California.

Range: Desert regions. Southern California
to Baja California, southern Nevada and Utah
and northern Arizona.

A few plants of E. eremicum approach the
growth fonns of E. pluriflonnn. The corolla of
the latter is usually regular, and the equal to sub-
equal stamens are adnate throughout the length
of the tube to the sinus. Rarely, a stamen will
become free as much as 0.75 mm below the
sinus. In the irregular to bilabiate flower of
E. eremicum, all the clefts are unecjual and the
lobes may l:>e irregularly disposed. The stamens
are adnate to 1.0 to 3.5 mm below the sinuses.
In many flowers of E. eremicum, the filaments
diverge toward the lowennost lobe and the
anthers turn toward the upper lobes. Within
Eria.strum this characteristic seems to be unicpie
to this species.

6b. Erimtrum eremicum subsp. ijageri (Jones)
H. L. Mason, Madrono 8.78. 1945.

Gilia virgata var.
West. Rot. 13:2.

ijageri
1910.

Jones, Contr.

G. eremica var. iirizonica Craig, Bull.

Torrey Club 61:419. 19.34.
G. eremica var. ii(i<^eri (Jones) Craig, op.

cit. 420, as to name, not as to lectotype.

Annual, 5.0 to 25.0 cm high, branched from
the base, or racemose, or virgate; herbage floc-
cose to subglabrate, often glandular; inteniodes
1.0 to 6.0 cm long; leaves light green, 1.0 to 5.5
cm long, 0.5 to 2.0 mm broad, with one or two
pair of lateral lobes, or entire, subulate, or awn-
tipped; heads 1 to 50, floccose to densely so,
0.5 to 1.5 cm broad, excluding tips of bracts;
bracts 1.0 to 2.0 cm exceeding the heads, 1.0 to
2.5 mm broad at base, with one to three pair
of lateral lobes, lightly floccose; hyaline mem-
brane between lobes sometimes present; calyx
0.5 to 1.0 cm long; lobes une((ual, joined by
hyaline membrane; tips not obscured by tri-
chomes; corolla funnelfomi, to 18.0 mm, irregu-
lar to nearly regular; lobes broadly elliptic, 5.5
to 7.0 mm long and 3.0 to 3.5 mm broad, com-
monly dark blue; throat ca. 2.0 to 4.9 mm long,
broadly spreading, vellow; sinus to stamen in-
sertion 2.0 to 4.0 mm; tulx^ 5.0 to 8.0 mm long,
vellow or blue; stamens adnate to base of throat,
unequal, 5.0 to 9.5 mm long, varying in same
flower by 1.0 to 3.0 mm; filaments 5.0 to 8.5
mm long; anthers sagittate, versatile, 1.0 to 1.5
mm long and 1.0 mm broad; stigma ca. 1.0 mm
long, style ca. 12.0 mm long, ovary three-loculed.

Type: M. E. Jones 102.53 and 10279. POM
74.569 and POM 74570. Wickenburg, Maricopa
County, and Hillside, Yavapai County, Arizona.

Range: Arizona.

Mason (1945) reviewed and corrected the
use of the name "Yageri." It stvms necessary,
however, to include here a history of its usage
in order to treat the lectotype (lectotype only,
not the name) of Gilia eremica var. ijageri (not
E. eremicum subsp. tja^ieri), which is also the
type specimen of Eriaslrum diffusum subsp.
jom'sii, which is here considered aS a synonym
of E. diffusum.

In describing G. virgata var. i/ageri, Jones
( 1910) cited Jones 1025.3 and Jones 10279 as the
type, and included a list of several specimens as
being representative, which, however, included
a few different entities. Presumably, the name
was derived from that of Yager, Arizona, where
one of the specimens had been collected. K.
Rrandegee regarded Jones 10279 ;is too fragmen-
tary for identification, and Craig also regarded
it as inadecjuate. Through use of the lactophcnol
method (Harrison 1968), it w;is possible to ex-
amine a flower of Jones 10279; and it appears

Biological Series, Vol. 16. No. 4 The C^enus Ehiastrum

17

to belong with Jones 10253 a.s designated by
Jones.

Craig evidently thought that because of the
geographic distribution, the name "Arizonica"
would be more appropriate than ijci'^eri and con-
sidered Jones 102.53 (type of G. virgata var.
i/(igcri) as the type for his Gilia ereinica var.
arizonica. Thus, the name "ijageri" was separa-
ted from its type. The name "yageri" was per-
{x>tnated, however, as Craig selected Jones 99.35
from Yager, Arizona, from Jones's list of "repre-
sentative material" (1910) as a lectotype of C.
eremica var. yageri (Craig, 1934).

As discussed by Mason (1945), the adequacy
of Jones 10253 necessitates recognition of Jones's
concept of C virgata var. yageri and the restora-
tion of the name to its type. Gilia eremica var.
arizonica was therefore referred to svnonomy
under Eriastnim eremicum subsp. yageri (Jones)
H. L. Mason.

Jones 99.35 was then selected by Mason as
the type of E. (liffustim subsp. jonesii Mason.
This plant was described as having larger corol-
las, slightlv larger anthers and longer filaments
than the tvpical fonii of the species. The speci-
men is here considered as being insufficiently
distinct to warrant recognition and is considered
by the present author as a synonym of E. dif-
fustnn.

Craig ( 19.34 ) had placed this plant with
G. eremica and had stated that he was uncertain
whether it should be referred to that species or
to G. filifoJiiim var. diffusa, which should be
interpreted as referring to E. diffusum. and not
E. fiJifolium.

Eriastnim eremicum subsp. yageri (Jones)
H. L. Mason is described ( Kearnev & Peebles
1951) as having less irregular corollas and sim-
pler leaves. The leaves of the type are similar
to those of E. eremicum subsp. eremicum, and
some depauperate plants have simple, entire
leaves. Lack of sufficient specimens has pre-
vented a broad examination of this character,
but the leaves appear to have fewer lobes than
those of E. eremicum subsp. eremicum. With
respect to the different depths of the sinuses,
the corollas are often <juite irregular. Corolla
lobes of this subspecies are broader, much dark-
er and brighter blue than the usual pale laven-
der of E. eremicum subsp. eremicum.

7. Eriastrum filifolium (Nutt) Wooton & Stand-
ley, Contr. U.S. Nat. Herb. 16:160. 1913.

Gi7t« filifolia Nutt, |our. Acad. Phila. II.

1:1.56. 1848.
Navarretia fiUfolia Kimt/e. Uev. Cen. PI.

2:4:3.3. 1891.

G. virgata var. fiUfolia Milliken, Univ.

Calif. Pub. Bot. 2:39. 1904.
N. fUifoha subsp. eufilifoHa Brand, Pflan-

zenreich IV, 250:167. 1907.
G. floccosa var. filifolia Nels & Macbride,

Bot. Gaz. 61:35. 1916.
Welwitschia fiUfolia Rydberg, Fl. Rocky

Mtns. 688. 1917.
Hugelia filifolia Jepson , Man. Fl. PI.

Calif. 792. 1925.
G. filifolia var. ti/pica Craig, Bull. Torrev

Club 61:422. 19.34.

Annual, 4.0 to 40.0 cm high, virgate, race-
mose, or diffusely branched; herbage subgla-
brous to lightly floccose; inteniodes 1.0 to 3.0
cm long, occasionally to 4.0 cm long; leaves fili-
fonn, 1.0 to 3.0 cm long, light green becoming
red brown, sometimes appressed, ascending
subulate-, or awn-tipped; first-formed leaves en-
tire; younger leaves with one pair of filiform
lateral lobes to 1.25 cm long, sometimes occur-
ring 5.0 mm above the leaf base, or entire, or
with t\vo pair of lobes from the base of the leaf;
heads few to several, long-tapered, often narrow
and elongate, distinctly brushy; tips of bracts
and calyces not obscured by wool, few-flowered;
bracts 1.25 to 2.0 cm long, exceeding the heads,
with one or two pair of lateral lobes; calyx
slender-cylindric, lobes une(jual to subequal, 6.0
to 8.0 mm long, joined bv a hyaline membrane;
corolla regular, salverfonn, narrowly funnelform,
7.0 to 9.0 mm long; lobes 3.0 mm long and 1.0
mm broad, blue; sinus to stamen insertion 1.0
mm, throat ca. 1.0 mm long; tube 4.0 mm long,
throat and tube yellow; stamens 3.5 mm long,
ecjual; anthers sagittate, versatile, 1.0 mm long;
filaments 2.5 to 3.0 mm long; stigma 0.75 to 1.0
mm long, style 3.0 to 4.0 mm long; ovary long,
cvlindric, three-loculed; capsule 5.0 mm long and
1.5 mm broad, each locule approximately four to
five-seeded.

Type: T. Nuttall. KEW. Santa Barbara, Cali-
fornia.

Range: Southern California from Santa Bar-
bara and Los Angeles counties south to Baja Cali-
fornia.

8. Eriastrum s]>arsiflorum (Eastwood) H. L.
Mason. Madrono 8:86. 1945.

.\nnual. to 30.0 cm high, racemose, corym-
bose, or often branched from the base, more
erect than spreading; herbage not disposed to
becoming brittle, light green, grey green or yel-
low green; lightlv floccose or subglabrate to
conspicuously floccose; internodes 1.0 to 4.0 cm

18

Bhicham VoUiNG Untversitv Science Bueletin

Kig. 5. Eriastrtwt filifoHum. a. Habit, I.. Abiams 3724 POM 156092, X .6. b. Calyx. T. Craig & M. Zornes
1856 POM 186740. X 4.6 c. Corolla and stamens. T. Craig & M. Zornes 1856 POM 186740, X 4.6. d. Habit.
T. Craig & M. Zornes 1856 POM 186740, X .6. e. Habit. Katberiiie Brandegee. V-06. UC 133849. X .6. /.
Habit. T. Craig & M. Zornes 1856 POM 186798. X .6. g. Habit. Nuttall GH (probable isotype), X .6.

Biological Series, Vol. 16, No. 4 The Genus Ehiastrum

19

long; leaves 0.5 to 3.5 cm long, most commonly
1.0 to 1.5 cm long, mostly entire, or with one
pair of lateral lobes; lobes to 7.0 mm long; heads
few to numerous, 0.5 to 1.0 cm broad excluding
tips of bracts, floccose to densely so, few-
flowered; bracts 0.7 to 2.5 cm long exceeding
the heads, entire or with one or two pair of
lateral lobes; calyx lobes unequal, 5.0 to 8.0 mm
long, joined by a hyaline membrane; corolla
regular, salverfonn to narrowly funnclfonn, 6.0
to 10.5 mm long; lobes elliptic, 2.0 to 3.0 mm
long, rarely more, blue, light pink, dull yellow
or cream; throat 0.75 to 1.0 mm long, yellow;
sinus to stamen insertion 0.25 to 0.50 mm, tube
4.0 to 5.5 nmi long; stamens equal, 1.0 to 2.25
mm long, filaments 0.50 to 1.0 mm long; anthers
sagittate, versatile, 0.75 to 1.0 mm long; stigma
0.5 mm long, style 3.5 to 4.5 mm long; ovary
three-loculed, few-seeded.

Key to the Subspecies

A. Leaves light green or light grey green;
corolla 6.5 to 8.5 or to 10.5 mm long, light
blue to light pink or cream; central Oregon,
Nevada, California along the Sierra Nevada,

and south to Baja California

8a. E. sparsiflorum subsp. sparsiflonim

AA. Leaves yellow green; corolla ca. 6.0 to 7.0
mm long, straw yellow; eastern Mohave

desert

8b. E. sparsiflorum subsp. hartcoodii

8a. Eriastrum sparsiflorum (Eastwood) H. L.
Mason subsp. sparsiflorum.

Gilia sparsiflora Eastwood, Proc. Calif.

Acad. Ill, 2:291. 1902.
Navarretia filifolia subsp. sparsiflora

Brand in Engler, Pflanzenreich IV, 250:

167. 1907.
G. filifolia var. sparsiflora Macbride,

Contr. Gray Herb. 49:57. 1917.
Hugelia filifolia var. sparsiflora Jepson,

Man. Fl. Pi. Calif. 792. 192.5.

Annual, erect, 10.0 to 30.0 cm high, occa-
sionally to 35.0 cm high, racemose, or often
much-branched from the base, paniculate, or
corymbose, more erect than spreading; herbage
not disposed to becoming rigid and brittle, light
green or light grey green, lightly floccose to
subglabrate, in part becoming more glabrate
with age; stems sometimes red brown; inter-
nodes 1.0 to 4.0 cm long; leaves few, 0.5 to 3.0
cm long, most commonly 1.0 to 1.5 cm long,
linear or with one pair of lateral lobes, 4.0 to

7.0 mm long; heads few to numerous, small, 0.5
to 1.0 cm broad, floccose, few-flowered; bracts
0.7 to 2.7 cm long, entire or with one pair of
lobes, exceeding the heads by 0.3 to 1.5 cm;
calyx lobes unequal or nearly equal, 5.0 to 8.0
mm long, rarely more, joined by a hyaline mem-
brane; corolla regular, salverform or very nar-
rowly funnelform, 6.5 to 8.5, sometimes to 10.5
mm long; lobes light blue and throat yellow, or
corolla light pink or cream; lobes narrowly el-
liptic 2.0 to 3.0 mm long, less commonly to 4.0
mm; lobes 1.0 mm broad; tube 4.0 to 5.5 mm
long, rarely more; stamens adnate to 0.5 nmi be-
low sinus, equal, 1.50 to 2.25 mm long; filaments
1.0 mm long; anthers sagittate, versatile, 1.0 mm
long; stigma 0.5 mm long, style ca. 4.5 mm long;
ovary three-loculed, few-seeded.

Type: Alice Eastwood. CAS 516. July 1-13,
Bubbs Creek, South Fork of Kings River, Fresno
County, California.

Range: Central Oregon, western Nevada,
Sierra Nevada Mountains from Lassen County,
California, south to Inyo County, to San Ber-
nardino County, west to Ventura County and
south to Baja California.

8b. Eriastrum sparsiflorum (Eastwood) H. L.
Mason subsp. harwooclii (Craig) H. K. Har-
rison, Phytomorphology 18:401. 1968.

Gilia filifolia var. Harwoodii Craig, Bull.

Torrey Club 61:424. 1934.
Hugelia diffusa var. Hardwoodii Jepson,

Fl. Calif. 3:167. 1943.
Eriastrum diffusum subsp. Hardwoodii

(Craig) H. L. Mason, Madroiio 8:77.

1945.

Annual, to 20.0 cm high, racemose or corym-
bose; herbage floccose to conspicuously so, yel-
low green to grey green; internodes 1.0 to 2.0
cm long; leaves 1.0 to 3.5 cm long, entire or
with one pair of lateral lobes, subulate, awn-
tipped; lobes to 7.0 mm long; heads few to
numerous, densely floccose, few-flowered, small,
0.5 to 1.0 cm broad excluding tips of bracts;
bracts 1.0 to 1.5 cm long, exceeding the heads,
with one or two pair of lateral lobes; calyx lobes
une(]ual, .5.0 to 8.0 mm long, joined by a hyaline
membrane; corolla regular, narrowly funnelform,
6.0 to 7.5 mm long, dull straw yellow or cream
vellow; lobes elliptic, 2.25 mm long and 1.0 to
1.5 mm broad; throat 0.75 mm long, sinus to
stamen insertion 0.25 mm, tube 4.0 mm long;
stamens equal, 1.0 mm long; filaments 0.50 to
0.75 mm long; anthers versatile, sagittate-
cordate, 0.75 mm long; stigma 0.50 to 1.0 mm

20

Buir.iiAM V()UN(; Univehsitv Scienck Bulletin

Kij;. (1 Eridslriini sparsijlorni}) suljsp. spiirsitlDruin. ii. Habit. Ali<(r Eastwood 1-H-VIl'W CAS jH) (typo), X .6.
/;. Ilahit, E. C. Twisselmami 2316 CAS .599515, X .6. c. Ihihit. R. Hacigalupi aiul C;. T. Rohhins 5417 CAS
407774. X .6. d. Calvx, H. M. Pollard 18-VI-49 CAS 352063. X 4.6. e. Corolla and stamens, H. M. I'oUard
18-VI-49 CAS 352063, X 4.6.

Biological Sehies, Vol. 16, No. 4 The Genus Ebiastrum

21

long, style ca. 3.5 mm long, ovary three-loculed,
locules approximately tvvo-seeded.

Type: P. A. Munz & R. D. Harwood 3589.
POM 7622. Blythe Junction, Riverside County,
California.

Range: Eastern Mohave Desert in eastern
Riverside and San Bernardino counties, Cali-
fornia.

9. Eriastrum wilcoxii (Nelson) H. L. Mason,
Madroiio 8:85. 1945.

Gilia floccosa Gray, emend. Syn. Fl. N.A.

2:143. 1878. Not type of G. floccosa

Gray, Proc. Anier. Acad. Sci. 8:272.

1870.
G. wilcoxii Nelson, Dot. Gaz. 34:27. 1902.
Welwitschia icilcoxii Rydberg, Fl. Rocky

Mtns. 688. 1917.
Hugelia filifolia var. floccosa Jepson, Fl.

Calif. 3:166. 1943, as to lectotype, not

as to type.

The following in part as to text, not as to
type (See discussion in Mason, 1945):

Gilia floccosa Gray, Proc. Amer. Acad.

Sci. 8:272. 1870.
Navarretia floccosa (Gray) Kuntze, Rev.

Gen. 2:4.33. 1891.
H. floccosa (Gray) Howell, Fl. N.W.

Amer. 458. 1903.
G. virgata var. floccosa (Gray) Milliken,

Univ. Calif. Publ. Bot. 2:40. 1904.
N. virgata subsp. floccosa (Gray) Brand

in Engler, Pflanzenreich IV, 250:168.

1907.
W. floccosa (Gray) Rydberg, Fl. Rocky

Mtns. 688. 1917.
H. virgata var. floccosa (Gray) Jepson,

Man. Fl. PI. Calif. 793. 1925.

Annual, to .30.0 cm high, racemose, coryin-
bose, or branched from the base; herbage floc-
cose, often densely so, gray green, becoming
rigid and brittle, coarse in appearance; inter-
nodes 1.0 to 4.0 cm long; leaves 1.5 to 3.0 cm
long, commonly 2.0 to 2.5 cm long, subulate-
tipped, usually with one pair of lateral lobes or
with two pair or entire; heads several to numer-
ous, 0.70 to 1.75 cm broad, densely floccose,
several-flowered; bracts prominent, 0.7 to 3.0
cm long, equalling or exceeding the heads, with
one or two pair of lateral lobes; calyx lobes un-
equal, 5.5 to 10.0 mm long, joined by a hyaline
membrane; tips of lobes sometimes conspicuous;
corolla regular or slightly irregular, funnelform,

9.0 to 14.0 mm long; lobes blue, tube and throat
yellow; lobes 3.5 to 6.0 mm long, elliptic to
broadly so, 1.50 to 2.25 mm broad; throat 0.5 to
2.0 mm long, sinus to stamen insertion 0.5 to
2.0 mm, tube 5.0 to 7.5 mm long; stamens un-
equal, 1.5 to 4.5 mm long, varying in one flower
by 0.5 mm or 2.5 mm, exserted beyond the sinus
(occasionally on the same plant some stamens
may be nearly equal and shorter; the anthers
barely surpassing the sinus, thus simulating a
characteristic of E. sparsiflorttm) ; filaments 1.5
to 3.5 mm long; anthers sagittate, versatile, 1.0
to 2.0 mm long and to 1.0 mm broad; stigma
0.50 to 0.75 mm long, style ca. 6.0 mm long;
ovary three-loculed, approximately three-seeded.

Type: E. D. Merrill & E. N. Wilcox 752.
RM 30221. Seven miles W St. Anthony, Idaho.

Range: Central Washington, central and
eastern Oregon, south along the Sierra Nevada
to Inyo County, California, southwest to San
Luis Obispo and Santa Barbara counties, south-
em Idaho, Nevada, and western Utah.

Eriastrum sparsiflorum and E. wilcoxii are
closely related. There is a wide range of varia-
tion and much overlapping in the expression of
the characters. Eriastrum wilcoxii plants tend
to be more floccose, more robust and thicker-
stemmed; the leaves tend to have one or two
pair of lateral lobes and to be somewhat rigid.
The heads are usually larger and have more
flowers per head. The flowers are slightly larger
and the filaments are longer and unequal. The
lobes of the corolla are sometimes broader, and
the vascularization of the corolla is less reduced.
The previously reported level of stamen inser-
tion as 2.0 mm below the sinus (Mason 1945)
does not hold, as it varies from 0.5 to 2.0 mm.

The plant body of E. sparsiflorum is a little
less brittle, less rigid, and less floccose than E.
wilcoxii. The plants may be tall and racemose,
or they may have many branches which indi-
vidually have few leaves. Perhaps the best char-
acters for identification of E. sparsiflorum are
the short, equal stamens, which quite consis-
tently become free 0.5 mm below the sinus; the
iinthers, which lie just above the sinus level; the
slightly smaller corolla, apparently always with
narrow elliptic lobes; the smaller heads; and the
greater frequency of short, entire leaves. It
seems that E. wilcoxii is the most nearly inter-
mediate between the large-flowered forms, e.g.,
E. eremicum and the small-flowered forms, e.g.,
E. sparsiflorttm and E. diffusum.

10. Eriastrum diffusum (Gray) H. L. Mason,
Madrono 8:76. 1945.

22

Hhigham Young Univehsity Science Bulletin

Fig. 7. ErUistntm wilcoxii. a. Hahit, I'. A. Miiiiz 11, 077 POM 153278. X .6. b. Habit, Ynez W. WinblaH 26-
VI I -38 CAS 297740, X .6. c. Habit, E. D. Merrill & E. N. Wilcox 9.52 RM 30221 (type). X .6. d. Corolla and
stamens, P. A. Munz 11, 077 POM 153278, X 3.4. c. Calv.v, P. A. Mnnz 11, 077 POM 153278, X 3.4.

Biological Series, Vol. 16, No. 4 The Genus Eriastrum

23

Gilia filifolia var. diffusa Gray, Proc.
Amer. Acad. Sci. 8:272. 1870.

Navarretia filifolia var. diffusa Brand in
En^ler, Pflanzenreich IV, 2.50:167.
1907.

Wehcitschia diffusa Rydberg, Fl. Rocky
Mtns. 688. 1917.

W. filifolia diffusa Tide.strom, Proc. Biol.
Soc. Wasli. 48:42. 19.35.

G. eremica var. tjaf^eri Craig, Bull. Torrey
Club 61:420. 1934, as to lectotypc only,
not G. virgata var. tjageri Jones.

Hugelia diffusa Jepson, Fl. Calif. 3:167.
1943.

Eriastrum diffusum subsp. jonesii H. L.
Mason, Madrono 8:77. 1945. (See dis-
cussion under E. ereinicum subsp.
yageri. )

Annual, 3.0 to 15.0 cm high, commonly S.O
cm high, diffusely branched from the base or
racemose, often spreading to twice the height;
herbage lightly floccose or subglabrous; stems
slender, 0.5 mm to 1.0 mm in diameter, often
becoming red brown; leaves slender, ascending,
light green, small, 1.0 to 2.5 cm long, usually
with one pair of short lateral lobes (2.0 to 7.0),
less commonlv with two pair or entire, subulate,
awn-tipped; inteniodes 1.0 to 2.0 cm long; floral
heads few to several, small, 0.5 to 1.0 cm in
length and breadth excluding tips of bracts,
few-flowered, moderately or lightly floccose;
bracts 0.75 to 1.50 cm long and exceeding the
heads, with one or two pair of lateral lobes;
calyx lobes unequal to nearly equal, 5.0 to 6.0
mm long, joined by a hyaline membrane; corolla
funnelform, 7.0 to 10.0 mm long, regular to
slightlv irregular; lobes light blue to cream, 2.5
to 4.0 mm long and 1.25 to 2.0 mm broad; sinus
to stamen insertion ca. 0.25 to 0.75 mm, throat
ca. 1.0 mm long, tube ca. 3.0 to 4.5 mm long;
stamens adnate nearly to sinus, 2.0 to 3.0 mm
long; anthers sagittate, cordate, versatile, 0.50 to
0.7.5 mm long, occasionally to 1.0 mm long; fila-
ments 1.5 to 2.5 mm long, stigma ca. 0.5 mm
long, style ca. 5.0 mm long, ovary three-loculed.

Lectotype: George Thurber 326. GH. Pa-
chetiju, south of Copper Mines, Grant to Luna
counties, New Mexico, to Arispe, Sonora, Mexico
(cf. Gray 1854. pp. 297-302).

Range: Southern California, Nevada, Utah,
southeast through Arizona, southwestern New
Mexico to Texas.

When Gray described Cilia filifolia var. dif-
fusa (1870), he wrote, "Forms of this approach
the preceding too nearly," by which he meant
Gilia floccosa and not Gilia filifolia. This is evi-

denced by his annotation on an S. Watson 915
collection at the Gray Herbarium to the effect
that forms approached G. floccosa too closely.
However, when the other collections which were
studied by Gray (Thurber 326, Cooper Fort
Mohave [in part] and probably Thurber 172)
are examined together with his description, the
variety to which he applied the name can be
recognized. The sheet bearing the aforemen-
tioned Watson collection has four collections
upon it, and they do not represent the elements
which prompted Gray to name a new variety,
but rather represent the discordant elements
which made him (jualify his original description
as being too similar to Gilia floccosa. As the type
of Eriastrum diffusum has not been indicated,
the selection of a lectotype is in order. In the
original description. Gray (1870) cited no col-
lector but gave the geographic range as "Fort
Mohave and Nevada to New Mexico and the
borders of Texas." Reference to the collections
confirms the Fort Mohave, New Mexico and
Texas border locations.

On the sheet containing the Watson 915 col-
lections, as well as the other collections, the
word "type" unsigned has been printed in pencil
at the bottom. This cannot be accepted as the
type nor as a lectotype, because there is no indi-
cation as to which collection was under consider-
ation; since Gray considered the plants of the
Watson collection as discordant elements and as
being too close to Gilia floccosa {Eriastrtnn
wilcoxii) to which I refer them, they do not
coincide with Gray's intention. The annotation
"type," is therefore ambiguous as to its applica-
tion; it has not been published; and there is no
indication as to whose opinion it represents.
Therefore, I consider the annotation as having
no standing. From the other specimens ex-
amined by Gray, it seems proper to select the
George Thurber collection 326 ( GH ) as a lecto-
type. This collection was made from May to
June 1851 at Pachetiju, south of the "Copper
Mines" (presumably Grand to Luna counties).
New Mexico. In May 1851, Thurber made a
journey from the Copper Mines (now Santa
Rita) and the Mimbres River south, by way of
the Playas Lake area, Animas, Guadalupe Pass,
Black Creek to the border at Agua Prieta, and
south into Mexico to Fronteras, Bacoachi, and
to Arispe (Gray 1854).

The closest affinities of E. diffusum are not
with E. filifolium. and their separation (Jepson
1943) was proper. Eriastrum diffusum plants
are of small stature with slender, fine stems, a
fraction of a millimeter to 1.0 mm in diameter.
The diffuse branching from the base is not the

24

Brigham Young University Science Bulletin

best criterion for recognition, as the branching
may be racemose, and since diffuse branching
occurs in other species. The vegetative portion
has some aspects of E. eremicum, chiefly in the
usual slender and much-branched habit. The
flower of E. difjtisum is regular and like that of
E. wilcoxii, being just a little smaller (although
sometimes somewhat broader), with similar un-
equal stamens with smaller anthers.

11. Eriastnim hrandegeae H. L. Mason, Ma-
droiio8:88. 1945.

Annual, to 25.0 cm high, racemose or virgate;
herbage lightly floccose; intemodes 1.0 to 4.0
cm long; leaves grey green, 1.0 to 2.5 cm long,
subulate, or awn-tipped, with one pair of lateral
lobes, less commonly entire; lobes to 1.0 cm
long; heads one to few, floccose, 0.5 to 1.0 cm
broad excluding tips of bracts; bracts 1.0 to 1.5
cm long, with one to two pair of lateral lobes;
calyx lobes unequal, 5.0 to 8.0 mm long, joined
by a hyaline membrane; tips not obscured by
trichomes; corolla regular to slightly irregular.
8.0 to 11.0 mm long, subsalverform; lobes nar-
rowly elliptic, 3.25 to 5.0 mm long and 1.0 mm
broad, lavender blue to white; throat plus tube
5.0 to 6.0 mm long, sinus to stamen insertion
0.5 to 1.5 mm; stamens 1.5 to 2.0 mm long, fila-
ments 1.0 to 1.25 mm long; anthers versatile,
sagittate, 0.75 to 1.0 mm long; stigma ca. 0.5
mm long, style ca. 3.5 mm long, ovary three-
loculed, each locule one- to two-seeded.

Type: H. L. Mason 12604. UC 693854.
Obsidian Ridge southeast of Borax Lake, Lake
County, California.

Range: Lake County, California.

Vegetatively, E. hrandegeae closely re-
sembles E. traaji. E. tiilcoxii and to a lesser
extent E. sparsiflorum and E. filifolhnn. The
flower, however, suggests a close relationship
with E. traci/i and differs from that of E. s])arsi-
flormn, E. filifolium, and E. wilcoxii, in that the
sides of the opened corolla are more nearly par-
allel. The corolla tube of E. hrandegeae is pro-
portionally broader than that of E. sparsiflorum
or £. filifolium. Eriastrum hrandegeae is further
distinguished from E. sparsiflorum by its larger
leaves, the larger leaf lobes, by a general ab-
sence of entire leaves in a mature plant, and in
having the stamens inserted at a lower level. It
is distinguished from E. filifolium by its shorter
stamens, slightly smaller anthers, by its more
pubescent heads, and by geographic isolation
from the latter. There are few distinctions be-
tween E. hrandegeae and E. traciji.

12. Eriastrum traciji H. L. Mason, Madrofio
8:87. 1945.

Annual, to 22.0 cm high, branching racemose
or virgate; herbage lightly floccose; intemodes
1.0 to 3.0 cm long; leaves grey green, 1.5 to 2.5
cm long, subulate, awn-tipped, with one pair of
lateral lobes or entire; lobes to 7.0 mm; heads
one to few, floccose to densely so, 0.5 to 1.0 cm
broad, excluding tip of bracts; bracts to 1.5 cm
long, exceeding the heads, with one or two pair
of lateral lobes, hyaline membrane between the
lobes sometimes present; calyx lobes unequal
7.0 to 9.0 mm long, joined by a hyaline mem-
brane; corolla regular, subsalverform, 9.0 to
10.0 mm long, light blue to white; lobes broadly
elliptic, 2.5 mm long and 1.25 to 1.50 mm broad;
throat 1.0 mm long, sinus to stamen insertion
0.50 to 0.75 mm, tube 5.5 mm long; stamens
equal, 1.0 to 1.5 mm long; filaments 0.50 to 1.0
mm long; anthers versatile, sagittate, 0.75 to 1.0
mm long; stigma 0.50 to 0.75 mm long; style ca.
4.0 mm long, ovary three-loculed, each locule
having a single seed.

Type: J. P. Tracy 6463. UC 690662. Hayfork
Valley 2600 ft. Trinity County, California.

Range: Trinity County, California.

It is with considerable hesitancy that the
author retains E. traciji as a distinct entity, and
the only reason for so doing is to avoid unneces-
sary synonomy until later collections may sub-
stantiate the distinction. The only characters
bv which the entity may be distinguished from
E. hrandegeae are the corolla proportions and
the exceedingly slight difference in filament
length. The lobes of E. tracyi are minutely
shorter and broader than those of E. hrandegeae.
When described, E. tractji was said (Mason
1945) to be distinguished from E. hrandegeae
and E. filifolium by its branching and by its
very small anthers and the corolla proportions.
Branching is of almost no value in the recogni-
tion of species of Eriastrum. When the anthers
of E. hrandegeae and E. tracyi are restored
( Harrison 1968 ) , they prove to have exactly the
same range in size from 0.75 to 1.0 mm. Each
specimen of E. hrandegeae examined has shown
narrowlv elliptic lobes from 3.0 to 3.5 mm long
and 1.0 to 1.25 mm broiid, those of E. traciji
have measured 2.50 mm long and 1.50 mm
broad.

13. Eriastrum ahramsii (Elmer) H. L. Mason,
Madrono 8:90. 1945.

Navarretia ahramsii Elmer, Bot. Gaz. 41:
314. 1906.

BioL{)CiCAL Series, Vol. 16, No. 4 The Genus Eriasthum

25

Hugelia ahramsii Jepson and Bailey in
Jepson, Fl. Calif. 3:167. 1943.

Annual, to 15.0 cni high, stems solitary or
branched racemosely, corymbosely, or diffusely
from the base; herbage floccose, often densely
so below the floral heads; intemodes 0.5 to 2.5
cm long; leaves when present filiform, light
green, 1.0 to 4.5 cm long or with one to three
pair of lateral lobes, not rigid, subulate, awn-
tipped; heads one to many, densely floccose, 1.0
cm broad, excluding tips of bracts, several-
flowered; bracts to 2.5 cm long with one to three
pair of lateral lobes, exceeding the heads by
1.0 to 1.5 cm, light green; the bases often com-
pletely obscured by dense, white wool; calyx 5.0
to 6.0 mm long; lobes unequal, joined by a hya-
line membrane, embedded in white wool; corol-
la regular, salverform or very narrowly funnel-
form, 5.0 to 8.0 mm long; lobes elliptic, 2.5 mm
long, blue or white; throat 1.2.5 mm long, tube
3.5 mm long, sinus to stamen insertion 1.5 mm;
stamens adnata to base of throat, equal or sub-
equal, 1.5 mm long; filaments 1.0 to 1.25 mm
long; anthers versatile, subsagittate, 0.50 to 0.75
mm long; stigma ca. 0.50 mm long, style 2.0 mm
long, ovary three-loculed, capsule spreading at
dehiscence, locules one-seeded.

Type: A. D. E. Elmer 4586. DS. Black-
Mountain, Santa Clara County, California ( Black
Mountain of the Santa Cruz Mountains, not
Black Mountain of the Mount Hamilton Range,
cf. Sharsmith 1944).

Range: Santa Cruz Mountains and Mount
Hamilton Range in Santa Clara County south to
San Benito County and north in Lake County.

Eriastnim ahrainsii is known from Lake
County and from the Santa Cruz Mountains and
the Mount Hamilton Range, where it is more
common. It is remarkable for the contrast be-
tween the light green of the leaves and bracts
and the very dense white mass of wool, which
obscures the bases of the calyces and the bracts
and continues to ;i short way down the stem.
The plants are of short stature, usually 15.0 cm
or less high; and the stems may be solitary,
branched racemosely, corymbosely, or diffusely
from the base. Flowering may occur when the
plant is a few centimeters high or when the
floral head is just above the ground level. As
the first leaves mav wither, the plant mav be

almost leafless; and a large part of the photo-
synthesis may be carried on by the bracts. The
corollas are white and protrude above the mass
of wool so slightly that they are quite incon-
spicuous. The species was first described as
Navarretia ahrainsii (Elmer 1906) and, as dis-
cussed by Sharsmith (1944) and Mason (1945),
is more closely related with Eriastrum. It oc-
curs as an associate of Adenostonm fasciculatum
chaparrel ( Sharsmith 1944 ) and is found "chiefly
confined to dry gravelly soil immediately bor-
dering thickets of the California Chamiso
{AdenoStome fasciculatum)." (Elmer 1906).

14. Eriastrum hooveri (Jeps. ) H. L. Mason,
Madrono 8:89, 1945.

Hugelia hooveri Jepson, Fl. Calif. 3:167.
1943.

Annual to 15.0 cm high, racemose, corym-
bose, virgate, or branched from the base; her-
bage lightly floccose to subglabrate, intemodes
0.5 to 3.0 cm long; leaves filiform, 0.5 to 2.5 cm
long, entire or with one pair of lateral lobes;
lobes to 5.0 mm long; heads one to few, com-
monly 0.50 to 0.75 mm broad excluding tips of
bracts, two- to eight-flowered, floccose to lightly
so; bracts 0.8 to 1.5 cm long, exceeding the
heads, with one pair of lateral lobes; calyx lobes
imequal, 4.0 to 6.0 mm long, joined by a hyaline
membrane; corolla regular, subsalverform, 5.0 to
6.0 mm long, white; lobes 2.0 to 2.5 mm long and
0.75 to 1.0 mm broad; throat 1.0 mm long, sinus
to stamen insertion 1.0 mm; tube 2.0 to 2. .5 mm
long; stamens 1.0 to 1.2.5 mm long, equal; fila-
ments 0.75 to 1.0 mm long; anthers sagittate-
cordate, versatile, 0.50 mm long; stigmas 0.25 to
0..50 mm long; style ca. 2.0 mm long, ovary
three-loculed, each locule one- to four-seeded.

Type: R. F. Hoover 1846. JEPS. Shafter,
Kern County, California.

Range: Southern San Joaquin Valley, Fi>esno
and Kem counties, California.

Eriastrum hooveri occurs on alkaline areas
and dry stream beds in the San Joaquin Valley.
The slender plants of short stature and heads
of two to five flowers suggest a relationship
with E. diffusum. The flowers of E. hooveri,
in their small size, resemble those of E. ahrainsii.

ACKNOWLEDGMENTS

I wish to thank Mrs. Emily Reid of the Uni-
versity of California, Berkeley, for the drawings

of Eriastrum.

26

Bricham Young U.niveksity Science Bulletin

LITERATURE CITED

Brand, A. 1907. Polemoniaceae, in Engler, Das
Pflanzenreich. IV. 250:1-203. Leipzig.

Craig, T. 1934. A revi.sion of the subgenus Hu^clki
of the genus Gilia (Polemoniaceae). Bull. Torrev
Club 61:385-396.

Elmer, A. D. E. 1906. New and noteworthy western
plants 3. Bot. Gaz. 41:309-326.

Ewan, J. 1937. Annotations upon the California flora.
I. Bull. Torrey Club 64:520-521.

Gray, A. 1854. Plantae novae thurberiauae. Mem.
Amer. Acad. Arts and Sci. N. Ser. 5:298-302.

Cray, A. 1870. Revision of North American Pole-
moniaceae. Proc. Amer. Acad. Sci. 8:247-282.

Cray, A. 1878. Synoptical flora of North America.
2:128-151. New York.

Cray, A. 1886. Synoptical flora of North America.
2nd ed. 2:128-151, 406-412. New York.

Harrison, H. K. 1959. Morphological and taxonomic
studies in the genus Eriastrum. Ph.D. Thesis. Uni-
versity of California, Berkeley.

Harbison, H. K. 1968. Contributions to the study of

the genus Eriastrum. I. The corolla and androecium.
Phytomorphology 18:393-402.

Heller, A. A. 1906. Botanical exploration in Cali-
fornia. Muhlenbergia 2:113.

Jepson, 1943. Flora of California. 3:160-168. ASUC.
Berkeley.

Jones, M. E. 1910. New species and notes. Contr.
West Bot. 13:2.

Kearney', T. H., and R. H. Peeble.s. 1951. Arizona
Flora. University of California Press, Berkeley and
Los Angeles.

Lanjouw, J. et al. 1906. International code of botani-
cal nomenclature. International Bureau for Plant
Taxonomy and Nomenclature. Utrecht.

Lanjouw, J. and F. A. Stafleu. 1964. Index Her-
bariorum. 4th ed. Utrecht.

Mason, H. L. 1945. The genus Eriastrum and the
influence of Bentham & Gray upon the problem of
generic confusion in Polemoniaceae. Madrono
8:65-91.

Sharsmith, H. K. 1944. Notes on Nai'arreti<i ahramsii
of the Polemoniaceae. Amer. Midi. Nat. 32:510-
512.