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NOTES ON ERIOGONUM - VI

A REVISION OF THE

ERIOGONUM MICROTHECUM COMPLEX

( POLYGONACEAE )

by

James L. Reveal

BIOLOGICAL SERIES— VOLUME XIII, NUMBER 1

APRIL 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany,

Brigham Young University, Prove, Utah

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

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Ernest L. Olson, Chairman, University Publications

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Brigham Young University
Science Bulletin

NOTES ON ERIOGONUM - VI

A REVISION OF THE

ERIOGONUM MICROTHECUM COMPLEX

(POLYGONACEAE)

by

James L. Reveal

BIOLOGICAL SERIES— VOLUME XIII, NUMBER 1

APRIL 1971

TABLE OF CONTENTS

ABSTRACT 1

INTRODUCTION 1

HISTORICAL STUDIES 1

TAXONOMY 4

Key to the Species of Coryiiibosa 6

Eriogonum leptophylluiii 8

Eriogonuni clavellatum 9

Eriogonum bicolor 10

Eriogonum ripleyi 12

Eriogonuni ericifolium 13

Key to the Varieties of E. ericifolium 14

Var. pulchrum 14

Var. ericifolium 15

Eriogonum microthecum 15

Key to the Varieties of E. microthecum 16

Var. laxiflorum 17

Var. toliosum 22

Var. panamintense 25

Var. corymbosoides 25

Var. johnstonii 28

Var. lapidicola 28

Var. alpiniun ^ 1

Var. microthecum 32

Var. ambiguum ^^

Eriogonum effusum 36

Key to the Varieties of E. effusum 38

Var. effusum 38

Var. rosmarinoides 39

SUMMARY 42

ACKNOWLEDGMENTS 42

LITERATURE CITED 44

NOTES ON ERIOGONUM - VI
A REVISION OF THE ERIOGONUM MICROTHECUM COMPLEX ( POLYGON ACEAE)

by

James L. Reveal'

ABSTRACT

This revision discusses seven closely related species
of Eriogonum (Polygonaceae) found in the western
L'nited States of North America. One of the two
major species, E. micratliccwn. is composed of nine
varieties which range over a wide area of the Far
West. The other major species, E. effimim, is found
over a more restrictive geographical range mainly east
of the Continental Divide. The remaining five species
are primarily restricted to the Colorado-Green river
drainage basin of Utah, Colorado, New Mexico, and
northern Arizona. Together these entities form the
core of a group of plants known as Section Corym-

bosa. The new varieties proposed are E. micwthcciim
var. coryinbosoides and M-dr.johnstonii from southern
California; E. micnitheciiin var. lapidicola of eastern
California, southern Nevada, and perhaps adjacent
Utah; and, E. mi era the cum var. alpimim from the
Sierra Nevada of east-central California. The follow-
ing new combinations are proposed: E. ericifolium
var. piihiirum and E. microtheciim var. foliosiim.
Detailed distribution maps are provided for each
taxonomic element recognized, and the variation in
E. microtheciim and E. effitsiim is illustrated.

INTRODUCTION

The Eriogonum microthecum complex, as here de-
fined, includes those species of Eriogamim which are
woody perennial shrubs or subshrubs with small lin-
ear-lanceolate to narrowly elliptic leaves with acute
apices mainly restricted to the lower half of the her-
baceous stems, and with glabrous, white to yellow,
small fiowers with rounded t)r more often tnmcate to
slightly cordate outer tepals. The complex is related
to E. corymbosum Benth. in DC. which differs in hav-
ing large lanceolate to elliptic or orbicular leaves and
slightly larger flowers with essentially round bases on
the outer tepals. This latter complex was treated as
Part V of this series of papers (Reveal, 1968). These
two species groups form a distinct subsection within
the larger section called Corymbosa which was named
by Bentham in deCandolle's Prodromus (1856) and
later typified with E. microthecum Nutt. (Reveal,
196%).

Eriogomim microthecum and the species most
closely related to it occur througliout much of the
western United States. They are found in numerous
habitats ranging from the high alpine reaches of the

Sierra Nevada to the hot desert floors of Nevada and
Arizima. The two species discussed in detail in this
paper, /:". microtlicciini and E. effusum Nutt., are not
narrowly endemic in their distribution, but other en-
tities associated with them tend to be restricted in
their geographical range. Consequently, other species,
now unknown and undiscovered, are likely to be
found in the future.

The Section Corymbosa is not a simple group. The
species referred to it are exceedingly variable in most
diagnostic features, and a lack of familiarity with the
plants in the field will put one at an immediate disad-
vantage in using keys to the species. Likewise, the
extreme variation that is characteristic of some taxa
does not reduce all of the problems. The polymorphic
characteristics of Eriogonum microthecum var. foUo-
siim is matched by the variation that exists in E. co-
rynihosum var. corymhosum. and even the relatively
restricted E. microthecum var. lapidicola is now de-
fined with such broad latitude that most certainly at
least two more variants will be described when addi-
tional material is available.

HISTORICAL STUDIES

The first species to be described in the complex occupation by the British and American gtwern-

were Eriogonum microthecum and E. effusum. ments) in 1834 with Nathaniel J. Wyeth and actively

Thomas Nuttall, the famous English botanist, had collected along the Oregon Trail (McKelvey, 1955:

traveled to the western half of what is now the Graustein, 1967). Although he described many of his

United States (then the Oregon Country held in joint newly discovered plants fiom 18.^7 to 1842, a few

'Department of Botany, University of Maryland, College Park, Maryland 20742; U.S. National Herbarium.
Smithsonian Institution, Washington, D.C. 20560.

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

remained to be described, and names for these were
not formally proposed until 1848 (Reveal and
Spevak. 1967). Among the new species proposed
were these two members of Hriogonum. Niittall sug-
gested herbarium names for some additional plants
which niiglit be worthy of publication but neglected
to provide them with the official endorsement of put-
ting them in print. Consequently, he left these to be
evaluated by others. In 1854, while discussing the
plant collections of Karl Geyer, William J. Hooker
proposed E. microtheciim var. laxijlontm. citing in
synonymy one of NuttalTs herbarium names. In 1856
Benthan wrote in de Candollc's famous world mono-
graphic series. Prodromus , and proposed the same
name. At the same time Benthani added E. simpsonii
from northern New Mexico, an area just acquired by
the United Slates from Me.xico; E. coitferiijlonim
from what is now northern California, altht)Ugh at the
time the collection was made the area was part of
Mexico; and, E. effusum var. rosmarinoides which
was designated as being from "California" but actual-
ly came from Kansas, an error pointed out by Torrey
and Gray in 1870 but not adopted into the California
literature until much later (Reveal and Munz, 1968).

In 1870 John Torrey of New York and Asa Gray
of Harvard University revised the genus Eriogonum
and discussed the E. microtheciim complex at some
length. Gray, who was the major author of the revi-
sion,^ took an exceedingly conservative view of Ben-
tham's Section Corymbosa which contained both E.
corynihosiim and E. microthecuin. One new species
was described (E. ericifolium). and E. corymhosum
was retained as a valid species. However, within E.
microtheciim. considerable "lumping" was done. In
the revision, Torrey and Gray described the flowers of
E. microtheciim as being "... albis nunc roseis raro
luteis . . ." for the overall concept of the species, and
from this action was introduced the misapplication of
the concept of var. microtheciim to what is now
called var. laxiflorum Hook., an error that continued
in the literature until the recent works by Hitchcock
(1964) and Reveal and Munz (1968). The var. laxi-
florum was only questionable recognized by Torrey
and Gray (their var. a) and included in their discus-
sion elements now referred to as var. alpiiuim in the
present paper.

Following their linear sequence, Torrey and Gray
recognized Benthani's vd\ . feihllcrianiaii (now a valid
species, viz. Reveal, 1968), but included in this entity
specimens referable to var. laxiflorum and var. amhig-
uum (M. E. Jones) Reveal in Munz. They proposed a
new combination, reducing Eriogonum confcrtiflo-
rum to a variety under E. microthecum and placing
under this name elements of both var. microtliecum
and var. laxiflorum. They next transferred E. effusum
var. leptophyllum {o E. microthecum. commenting

^(See Gray's footnote to Torrey's delayed publication of
Exploring Expedition which was finally released in 1874 -here

that Bentham's E. confertiflorum var. stansburyi was
an intermediate stage between var. confertiflorum
and var. leptopliylliim. With only fragments of var.
stansburyi and var. leptophyllum at hand in 1870,
this transfer is not at all surprising. However, now it is
possible to refer var. stansburyi to /;'. hrevicaule
Nutt., and var. leptoplnilum is considered a valid spe-
cies by most authors.

In their treatment of var. leptophyllum, Torrey
and Gray put a diverse series of entities under this
name. Eriogonum leptophyllum. elevated to the spe-
cies rank by Woott)n and Standley in 1913. was con-
fused by Torrey and Gray with specimens of what is
here called E. microthecum var. foliosum. Although
they did correctly associate var. foliosum (in name
only) with Bentham's E. simpsonii. they failed to see
the distinct differences which exist between E. lepto-
phyllum and E. microthecum var. foliosum.

The reduction of Eriogonum effusum to the vari-
etal rank under E. microthecum was critical and is
followed today by some {viz. Hitchcock, 1964;
Porter, 1968). This is both unfortunate and unneces-
sary. Porter (personal communication of 6 February
l'-)67) claims that the two "...seem to intergrade
badly and . . . have almost continuous (at least contig-
uous) ranges . . . ." From my data no such intergrades
have been detected. However, even if this were to
happen, the two species are distinct, both morpholog-
ically and geographically, and should not be con-
fused. As will be seen, the zone of possible overlap is
small (only in southern Colorado and northern New
Mexico) and no major taxonomic change would be
warranted.

The last variety Torrey and Gray discussed in their
revision of Eriogonum microthecum was var. leptocla-
don. In 1857 they proposed this entity as a valid spe-
cies, but were forced to reduce it to the varietal rank
to conform with the rest of their revision. In 1966 I
suggested that E. leptocladon be retained as a species
and that Eastwood's E. ramosissimum be placed as a
variety within it. Since then, fieldwork has shown
that the var. ramosissimum (Eastw.) Reveal tends to
come close to E. kearneyi Tidestr. in Kane County,
Utah on the western flank of the range of vdt.ramo-
sissimutn. while on the eastern flank it is approached
by the New Mexican phase of E. effusum wdi. effusum.

The complex remained neglected for some years
after the Torrey and Gray revision. Watson (1877)
did not alter their concepts to any great degree
although he transferred Eriogonumm ericifolium to
E. fasciculaium Benth. and placed E. microthecum
var. fendlerianum in synonymy under E. corym-
bosum. Charles C. Parry described E. mearnsii in
188'). Marcus E. Jones described E. bicolor from east-
ern Utah in 1893, added a few varieties in 1895, and
proposed E. friscanum in 1903, based on specimens

the plants collected by the 1838-1842 Wilkes' linited States
Gray reveals the role he played in the revision)

BIOLOGICAL SERIKS. VOL. I J, NO. 1 NOTES ON ERIOGONUM - VI

from southwestern Utah. Small added E. clavellatuin
in 1898 tVom a L-nllection gathered by Alice East-
wood in liS'-)5. Ill 1906 Michel Gandoger, Europe's
famous "splitter,"" described several new species and
varieties, all of which are now placed in synonymy!
In 1936 Susan G. Stokes published her monograph
of Ehogonitm and presented a bewildering arrange-
ment of the various taxa associated with the £". micro-
theciim complex. Her typical subspecies appears to be
the same as my var. inicrothecum, h\.i\ the placement
of £". effiisiim var. rosntariiioiclcs as a synonym under
it is difficult to explain. Within ssp. typicum. she rec-
ognized a series of varieties that actually belong to
vastly different forms. Her other subspecies are all
synonyms except for var. pananiiiitense (see Table 1 ).

Name in Stokes (1936)

Present Name or Concept

Ssp. typicum S. Stokes
Var. macdougalii

(Gand.) S. Stokes
Var. paiiamimeitse

S. Stokes
Var. friscanuin

(M.E. Jones) S. Stokes
Var. spatbulare

(Gand.) S. Stokes
Var. idahoense

(Rydb.) S. Stokes
Ssp. laxifloniin (Hook.)

S. Stokes
Ssp. confenijloruin

(Benth. in DC.) S. Stokes
Ssp. rigidiim (Eastw.)

S. Stokes
Ssp. ericifoiium

(Torr. & Gray) S. Stokes
Ssp. intermedium

S. Stokes
Ssp. mearnsii

(Parry in Britt.) S. Stokes
Ssp. bicolor (M.E. Jones)

S. Stokes
Ssp. pulclirum (Eastw.)
S. Stokes

Ssp. aureuin (M. E. Jones)
S. Stokes

Var. crispum (L. Will.)
S. Stokes

Var. expanmin S. Stokes

Var. microthecum
Var. laxijlorum Hook.

Var. paiiaininieme S. Stokes

Var. foliosum

(torr, & Gray) Reveal
Var. laxitloruiii Hook.

Var. microthecum

Var. kxiflorum Hook.

Var. laxijlorum Hook.

Var. foUosu}n

(Torr. & Gray) Reveal
E. ericifoiium Torr. &

Gray var. ericifoiium
\:li. foliosum (Torr. &

Gray) Reveal
E. ericifoiium Torr. &

Gray var. ericifoiium
£". bicolor M. E. Jones

E. ericifoiium Torr &
Gray var. pulcbrum
(Eastw.) Reveal

E. corymbusuiu Benth. in DC.

v'di. glutiiiosuin (M.E. Jones)

M. E. Jones
E. corymhosum Benth. in DC.

var. glutiiiosum (M. E. Jones)

M. E. Jones
Var. ambiguum (M. E. Jones)

Reveal in Mun?.

Table I. The disposition of ta.xa recognized by Stokes
(1936) in Eriogonum microthecum. unless otherwise indi-
cated, all varieties are referable to E. microthecum.

As can be seen in Table I . Stokes includes four dif-
ferent elements within her definition of .ssp. typicum.
Only var. idahoense is actually referable to var. inicro-
ihecitm. while both var. niacdinigalii and var. spathii-
lare are placed in synonymy under var. laxijlorum in
the present treatment of the group. The inclusion of
var. panainintense is difficult to explain as is her
placement of var. friscanuin - a typical form of var.
foliosum. or what she called ssp. rigiJum. Her con-

cept of var. laxiflorum (her ssp. laxijlorum) is correct,
but I cannot maintain ssp. confcrtijlorum at any
rank. When one compares the type of Eriogomtin
confertijlorwn with Stokes' description of the entity,
they do not match. From the description, I would
judge that her intention was to include the tall forms
ot var. foliosum. but as no specimens were ever anno-
tated with this name, one can only speculate.

Stokes proposed ssp. rigidum for what I am calling
var. foliosum, but, as Torrey and Gray's Eriogonum
effusum var. foliosum and Eastwood's E. microthe-
cum var. rigidum represent the same kind of plant,
Stokes should have adopted the earlier var. foliosum
as the basionym of her name. The ssp. intemwdlum.
according to Stokes, is an internrediate form between
E. microthecum and E. fasciculatum. a point I cannot
accept. If anything, the type of her new subspecies
appears to be an intermediate between var. laxiflorum
and vix. foliosum of E. microthecum.

The reiriaining subspecies referred to Eriogoinim
microthecum should be retained as distinct species or
associated with other species. None of these except
her var. expansum should be associated with E.
inicri)tlu'cum.

Name in Stokes (1936)

Present Name or Concept

Ssp. typicum S. Stokes
Ssp. lielichrysoides

(Gand.) S. Stokes
Ssp. feudleriainim

(Benth. in DC.) S. Stokes
Ssp. ainslcl (Woot. &

Standi.) S. Stokes
Ssp. salicimnn (Greene)

S. Stokes
Ssp. orbiculaluiii S. Stokes

Ssp. corvnibosuin (Benth.

in Di:.) S. Stokes
Ssp. divaricatum (Torr. &

Gray) S. Stokes
Ssp. durum S. Stokes

Ssp. sabinim (A. Nels.)

S. Stokes
Ssp. pallidum (Small)

S. Stokes

Var. shaudsii S. Stokes

Ssp. leptocladon (Torr. &

Gray) S. Stokes
Ssp. simpsonii (Benth. in

DC.) S. Stokes

Ssp. nelsomi (L. Will.)
S. Stokes

Ssp. coHtorium (Small e\
Rydb.) S. Stokes

Var. effusum

Var. rosmariuoides Benth.

in DC.
£". fendlerianutu (Benth. in DC.)

Small
E. fendlerianum (Benth. in

DC.) Small
E. lonchophyllum Torr. & Gray

E. corymbosum Benth. in DC.
var. orbiculalum
(S. Stokes) Reveal
and Brotherson
E. corymbosum Benth. in DC.

var. corymbosum
E. corymbosum Benth. in DC.

var. corymbosuui
E. corymbosum Benth. in DC.

var. corymbosum
E. corymbosum Benth. in DC.

var. corymbosum
E. leptocladon Torr. &
Gray var. ramosissimum
(Eastw.) Reveal
E. leptocladon Torr. & Gray

var. leptocladon
E. leptocladon Torr. & Gray

var. leptocladon
E. microthecum Nutt. var.
foliosum (Torr. &
Gray) Reveal
E. microthecum Nutt. var.
foliosum (Torr. &
Gray) Reveal
E. contortum Small ex
Rvdb.

Table 2. The disposition of taxa recognized by Stokes
( 1936) in Eriogonum effusum: unless otherwise indicated, all
varieties arc referable to £'. effusum. The nomenclature o{ E.
corymbosum follows Reveal (1968) while the reinaining taxa
are based on Reveal ( 1969a).

BRIC.HAM YOUNG UNIVIIKSIIY SC11:NCE BULLEIIN

Under I-'riogonum efjhsiim. Stokes treated the
reinaining members of the t. micwthccwn complex.
Her concept of typical E. effuswn (as ssp. typicum) is
correct, but she failed to realize that var. rosmari-
noides (which she placed under F.. micwtheaim ssp.
typicum as a synonym) and tiie new combination.
ssp. helichrysoides (Gand.) S. Stokes, were the same
element. Most of the remaining subspecies under H.
effiisiiiii belong to tiie E. corymhosuin complex as
summarized in Table 2.

In the list of synonyms found under some ot the
subspecies recognized by Stokes were some odd ele-
ments. In addition to Eriogoinini effusiiiii var. ros-
luarinoides which she referred to E. microthecwn ssp.
typicum, she also placed E. confertiflorum var. staiis-
huryi under her ssp. confertiflorum: E. aureum M. E.
Jones var. amhiguum M. E. Jones was put in synon-
ymy under her ssp. aureum: yet. at the same time, she
described the exact same plant as var. expaiisum.
Under the subspecies placed with E. effusum she
referred /•.'. iniriaitum Gand. under ssp. typicum
when the name should have been placed under /;'.
microtliecum var. laxiflorum. The puz/ling species, E.
sarolhriforme Gand., was placed under the ssp.
corvmliosum. when in reality the name sht)uld be
referred to either /;". hrevicaule. or as 1 have ventured
to suggest (Reveal,, 196^3). to /;". lonchopliyllum
Torr. & Gray - both species quite distinct and differ-
ent from /;'. curymhosum.

In her monograph, Stokes maintained Eriogonum
leptophyllum and /■.'. clavcllatum as distinct species.

Work on the Eriogonum microtliecum complex
since 19,>6 has been minimal. Stokes described /;'.
efflisum var. limln/lum in 1 441, bui this is identical

to her /:'. microtliecum var. punaminlcnsc named five
years before. Kearney and Peebles, unaware of the
nature of /;'. ericifolium. made the combination E.
mearnsii var. pulchrum in 1939. and I proposed the
new combination, E. microtliecum var. amhiguum in
1968.

!n l9t)X 1 published a revision o\ \W isiogonum
corymbosum complex. Since then, one additional
species has been found. Eriogonum revealianum
Welsh ( 1970), was known to me in 1968 and included
in my concept of £". corymbosum var. corymbosum. I
believe that this new name should be reduced to
synonymy under var. corymbosum.

Within Eriogonum microtliecum, a number of new
entities are proposed at this time. Two of the new
varieties, var. alpimim and var. jolmsionii. have been
suspected as being undescribed for a U)ng time but
never formally proposed until now. The var. corym-
hosoiUcs has been collected for several years in the
mountains of southern California, but its unique na-
ture generally has gone unnoticed. The var. lapidicola
has been known to collectors since the late 1890s.
but it was not until the fine collections of Dr. Janice
C. Beatley on the Nevada Test Site in southern
Nevada that the distinctiveness of this entity became
obvious.

The population called vdw fuliosum by me in this
paper has been outlined in various manuals and tloras
under a variety of names, but the proper combination
has not been made until now.

One undescribed species is known to me that is
not included in this paper. This species is known from
a single plant gathered in Delta County, Colorado,
some years ago, but until it can be rediscovered and
studied in detail, it seems unwise to name it now.

TAXONOMY

In my doctoral dissertation (Reveal, I'-Hi'^hi). I pro- 1903. Species lectotypus: /;. microtliecum Nutt., se-
posed that the Section Corymbosa be divided into lected by Reveal in Gunckel, Curr. Topics PI. Sci.
three Subsections. One was designed to include Eriog-
onum deserticola S. Wats, of extreme southern Cali-
fornia, while a second (also undescribed) was outlined
to include E. leptocladon Torr. & Gray, /;'. kearneyi
Tidestr., the recently described E. ammopliilum Re-
veal, and the rare western Utah endemic, E. iium-
mularc M. E. Jones. A pt)rtion of the typical subsec-
tion is the subject of this paper and may be described
and discussed as follows:*

Eriogonum Sect. Corymbosa

Eriogonum Michx. Sect. Corymbosa Benth. in
DC., Prodr. 14:17. 1856. Sect. Corymharium
Kuntze in Post & Kuntze, Lexicon Gen. Phan. 204.

236. 1969.

Low to liifih. spreading to erect, piiiviniite to compact or
open siilishrubs to sliriilis, lierliaceoiis or \vooil\ . (1,02-1.5 iii
liigli, 0.1-2.5 m across, the lower stems reddish-brown to
tirown or somewhat grayish, woody at least as much as hall'
the height of plants or less, leafless or nearly so, the upper
stems herliacoous; leaves solitary or in fascicles at the tips of
short dwarf shoots, variously scattered or even rather con-
gested along the lower portions of the herliaceous branches,
occasionally on tlie upper portion of the woody stems, the
leaf-blades variable, linear to orbicular, densely tomenlose on
both surfaces, less so to glabrous above, or totally glabrous
on both surfaces, the margins plane to revolute. on short to
long petioles; iiijloresceiices cyniose, compact or congested,
mfrequently open and expanded, the internodcs short, glali-
rous to densely tomenlose. the involucres arranged in the
forks of the branches or at the tips of the ultimate branches,
not racemosely arranged at the tips; involucres solitary, tur-
binate to broadly campanulatc, glabrous to tomenlose with-

"Unlil the oilier subsections are proposed, the typical subsection of Section C'orymbo.sa does not c\isl. Consequently, the
description of the "section" given below is actually based on the subsection as 1 have defined it (Reveal, 1969a) but not called
such for nomenclaliiral reasons.

BIOLOGICAL SERIES. VOL. I 3, NO. I NOIKS ON ERIOGONUM

VI

5

out. mostly 5-tootlied, the tcctli usualh' not dccph dividing
the tube; /7ou'('rs white to pink or yellow, with greenish, red-
dish, or reddish-brown midribs and bases, essentially glabrous
or (in E. effiisiim) rarely sparsely pubescent at the base and
along the midribs without, the tepals similar or dissimilar, the
outer whorl of tepals oblong, obovate, spathulate, or elliptic
to fan-shaped, or nearly to quite orbicular, the inner whorl of
tepals narrower, mostly oblanceolatc to spathulate or elliptic;
stamens slightly to long e\serted, the filaments glabrous to
pilose basally, the anthers variable in color, mostly oblong;
achenes light brown to brown, glabrous, not winged.

Dislribiition.

In several habitat types from the low, warm desert
floors to exposed mountain ridges, from (1500)
3000-10,500 feet elevation, througliout much of the
western United States, from Washington to California
eastward to the Great Plains, from Nebraska and
South Dakota southward to central New Mexico with
a predominance of species in the Green and Colorado
rivers drainage basin, especially in eastern Utah.

The subsection is a difficult group. It is large and
complex, containing some sixteen species and perhaps
twenty varieties; yet, the taxonomy of the group is
now tairly well understood and documented. The
taxon is divided into two major species complexes,
one centering around Eriogoiniin corymbosiim and
the other around E. microtheciim. The foriner com-
plex was the subject of my fifth part of this series,
and the latter complex is the group discussed in this,
the sixth part, of the series. Within these two groups
are a great many forms. Some, like E. lancifolium Re-
veal and Brotherson, E. saitriniim Reveal, and E.
smithii Reveal, are narrowly endemic to a small spe-
cific site. This seems to be a characteristic of the E.
corymbosiim complex, as the taxa in this group tend
to be more restrictive in their distribution than the
taxa associated with E. microthecum. For example,
both E. microthecum var. la.xiflonim and E. cfftisiim
var. effusiim occur in a number of contrasting ecolog-
ical niches. Even the various variants within E. mi-
crothecum occupy a multitude of sites except the re-
stricted var. johnstonii. The only species in the E.
microthecum complex which are even somewhat re-
stricted in their distribution are E. clavellatum in
southern Utah, E. ripleyi J. T. Howell and E. ericifol-
ium of northern Arizona, and the undescribed species
apparently restricted to clay slopes near Hotchkiss,
Delta County, Colorado. Some additional forms oi E.
microthecum (as yet undescribed) may prove to be
narrowly distributed.

Some hybridization is known to occur in this part
of the Section Corymbosa. Eriogonum microthecum
var. laxijlorum and E. corymbosiim vars. corymbo-
siim and erectum Reveal and Brotherson will hybrid-
ize with E. brevicaule, a herbaceous species of
another section in the genus. It is suggested here that
the entity known as E. nehraskense is the probable
result of hybridization between E. effitsum var. ef
fusum and E. pauciflorum Pursh, but until this can be

demonstrated, formal recognition of this as a hybrid
species will have to wait. It is likely that other hy-
brids will be found in the future.

Some entities show amazing variability. Eriogo-
num microlliecum \\\\ . foliosiim probably is the most
variable plant in the genus over its entire range. Other
species in this species complex are extremely stable,
but these tend to be the endemics restricted to a
small geographical range. One exception to this is E.
bicolor which is consistent throughout its rather ex-
tensive range in eastern Utah and western Colorado.
Variation is the source of new kinds within a popula-
tion, and it should be expected that the more widely
distributed species should be the most variable. How-
ever, E. microthecum var. corymbosoides, in its small
geographical range in the mountains of southern Cali-
fornia is exceedingly variable in its pubescent charac-
ters, much more so than most of the more widespread
entities. This may result, over a long period of time,
in the further evolution of new kinds within this area.

The reader is forewarned that the exact determina-
tion of some populations within this group will be
difficult. This section has been studied extensively in
the field, and I liave called upon this tleld experience
in arriving at the taxonomic conclusion presented
herein. Consequently, mere fragments or poorly pre-
served herbarium material may be impossible to de-
termine, especially at the infraspecific rank. Geo-
graphical and edaphic isolation has seemingly played
a major role in the evolution of this group, and the
exact location of an undetermined collection will
prove helpful in arriving at its proper name. The vari-
eties within Eriogomim microthecum are easily recog-
nized in the field or on well prepared herbarium
sheets once the entity is known in the field, but when
only the upper branches are gathered, the determina-
tion may prove fruitless. The key to the various spe-
cies in the Section Corymbosa (as discussed here) at-
tempts to take in all exceptions and provide the user
with several possible choices. As the key is based
mainly on leaf features, it is to be understood that
the general trend of all leaves is to be considered
rather than the one leaf that differs from the rest.
However, I have attempted to provide a lead for most
of the commonly encountered exceptions even if
these exist on only a few herbarium specimens.

Note.

The following key is to the entire Corymbosa
taxon. It includes those species treated in this paper,
part V of this species, and other species as suggested
in my overall revision of the genus (Reveal, 1969a). It
is hoped that an inclusive key will be useful to those
faced with the necessity of determining a member of
the group and not knowing whether it is more closely
related to Eriogomim microiliccum or E. corym-
bosum.

HRICll AM YOUNG UNIVF.USITY SCIliNCK BULLETIN

Key to the Species of Corymbosa

A. Leaves narrow, linear or narrowly elliptic or narrowly
lanceolate, the apices sharply acute or nearly so, or if the
leaves broader, then plants from the mountains of east-
ern and southern California or of the plains of central
Washington

B. Leaves tightly revolnte. narrow
C. Leaves 2-6 cm long

D. Inflorescences densely cyniose, 0.2-1.2
(1.5) dm long, glabrous and bright green;
plants forming round compact shrubs 2-6
(8) dm high, 3-10 (12) dm across: involu-
cres 2-3 mm long, 1.1-1.7 (2) mm wide,
glabrous; clay hills and slopes, Sandoval
County, New Mexico, westward across
northwestern New Mexico into Apache,
Navajo, and extreme eastern Coconino
counties, Arizona, northward to the lour
Corners area of Utah and Colorado

1. /■.'. leptoph villi m.

DD. Intlorescences less dense, or if so. then the

stems not glabrous; plants forming erect to

spreading shrubs and subslirubs; involucres

tloccose to lanate without

E. Inflorescences (L5) 2-6 cm long,

white-tomentose or lanate, rarely

glabrate; leaves 0.3-3.5 (4) cm long,

mostly rigid, thickish in some; clay

hills and slopes, sandy desert washes

and rocky outcrops, San Juan

County, New Mexico, and adjacent

southern Colorado westward across

northern Arizona, southern Utah, and

southern Nevada into extreme eastern

California (the var. joliosum phase of

£". microllieciim)

6. £'. microilieaiiit.
Lh. Inllorescences 5-15 cm long, tloccose
(o glabrate; leaves 3.5-6 cm long,
twisted in some, thin and linear; clay
hills and washes, west-central Kansas
(the var. rosmarninoides phase of E.
ef I'll. •ill in)

7. /■,'. ctfusuiii
CC. Leaves 0.5-2 cm long

D. Plants subshrubs. 1-2 dm high; involucres
sessile

E. Involucres (3.5) 4-4.5 mm long, glab-
rous; leaves 0.8-1.7 (2) mm wide,
glabrous above; stems green and glab-
rous, sparsely branched; low round
compact subshrubs; clay slopes and
washes, rare, San .luan County, Utah
2. E. clavellaliiin.
V.V.. Involucres 2-2.5 mm long, pubescent
or rarely glabrate; leaves 2-4 mm
wide, sparsely pubescent above; stems
white-tomentose, rarely glabrate in
New Mexico and Arizona, cymoscly
branched; low decumbent, prostrate,
or erect subshrubs; granite or lime-
stone outcrops and slopes in the high
mountains of eastern California (var.
alpimim) and Nevada (low forms of
var. laxiflonim), or on rolling clay
hills and slopes at lower elevations in
western Colorado and adjacent Utah
into northern Arizona and southern
and central Nevada (low forms of var.
Joliosum and var. lapidkola)

6. E. microthniiin.

DD. Plants matted, woody, less than 1 dm
high, or if higher, then lacking bracts
immediately below the invohicres; involu-
cres sessile or on short peduncles
E. Flowers 2.5-4.5 mm long; achenes
3-3.5 mm long; west-central Colo-
rado, eastern Utah, and northwestern
Arizona

F. Tepals similar or nearly so,
oblong, united at least 'A the
length of the flower; leaves thin-
ly tloccose to glabrate; inflores-
cences cymose; involucres nar-
rowly turbinate, 3-3.5 mm long,
1-1.5 min wide; Delta County,
Colorado.

Eriogomim species (undescribed).
FF. Tepals distinctly dissimilar, the
outer whorl of tepals broadly
obovate to orbicular, united
about 'A the length of the tlower;
inflore.scences umbellate-cymose;
involucres turbinate to campanu-
late

G. Leaves densely tomentose
on both surfaces, 5-12 (15)
mm long, 1-2 (3) mm wide;
involucres turbinate-cam-
panulate, 5-lobed; tlowers
2.5-4 mm long, the outer
whorl of tepals (2) 2.5-3
mm wide; involucres pedun-
cled, the bracts 1.5-3 (4)
mm below the base of the
involucral tube; clay fiats
and rolling hills. Mesa Coun-
ty, Colorado, and adjacent
eastern Utah from Grand
County south to northern
San Juan County and Wayne
Countv'.

3. E. hi color.
GG. Leaves densely tomentose
below, thinly tloccose to vil-
lous and greenish above, 2-6
mm long. 0.5-1 mm wide;
involucres campanulate, 3-5
mm long, 3-5 lobed; tlowers
3.5-4.5 mm long, the outer
whorl of tepals 3.5-4 mm
wide; involucres on top of
bractlcss peduncles; sandy
clay slopes and washes in
south w e s t e r n Coconino
County and northern Yava-
pai County, Arizona

4. E. ripleyi.

EE. Mowers 2-2.5 mm long; achenes 2-2.5
1. inm long; leaf-blades 5-8 mm long,

0.8-L5 (1.9) mm wide; involucres
2.5-3 mm long, 1.5-2 mm wide; rocky
places on flats and slopes, western
Mohave County, southern and central
Navajo and Coconino counties, and
Yavapai Count\ . Arizona

5. E. cricifoliiiin.
HH. 1 eaves tlat. not revolutc or with the margins rolkxl,
i)ccasionally with margins thickened
C. Leaves 0.2-4 cm long

D. Leaf-apices sharply acute, the bl.idcs most-
ly narrowly elliptic or narrower, 1-8 (20)
mm wide; plants mostly low subshrubs
E. Inflorescences (1.5-6 (12) cm long.

BIOLOGICAL SERIES. VOL. I 3, NO. I NOTES ON ERIOGONUM - VI

compact and rather congested, the
branches uhite-tonientose to lanate
or more commonly tloccose to sub-
glabrous, rarely glabrous, green: leaves
0.3-3.5 (4) cm long, 1-8 (20) mm
wide; widespread on a variety of soil
types west of the Continental Divide,
from Washington eastward to Mon-
tana, hence south to southern Califor-
nia, northern ."Arizona, and New Mex-
ico

6. E. luicrolliccmn.

EE. Inflorescences 6-25 cm long, open,
the branches floccose, white but
often drying blackish; leaves linear-
oblanceolate or oblanceolate to ob-
long or obovate, (I) 2-4 cm long,
mostly 3-7 (10) mm wide, the blades
plane or in the mountains of central
Colorado, some plants with blades
rolled; gravelly to sandy hills, slopes
and flats, east of the Continental
Divide from South Dakota, Wyoming,
and adjacent Nebraska soutliward
along and in the Front Range and on
the western edge of the Great Plains
in Colorado to central and northwest-
ern (west of the Divide) New .Mexico
7. E. effiisum.
DD. Leaf-apices slightly acute to rounded, the

blades oblanceolate to elliptic. (0.5) 1-3

cm wide; plants decidedly woody shrubs;

Utah and Colorado south to New Mexico

and Arizona

8. E. coirmhosinn.
CC. Leaves 3-8 cm long

D. Leaf-apices mostly rounded, the blades

oblanceolate to elliptic, 1-3 (5) cm long;

southern Utah and northern Arizona (the

var. ghitiiiosiiin phase off. corymhosum)
8. E. corymhosum.

DD. Leaf-apices acute, usually sharply so, the
blades mostly lanceolate, usually more
than 3 cm long

E. Branches subglabrous to tomentose;
involucres tomentose without
L. Involucres 2.5-3 mm long; IntTo-
rescences with several short
branches; flowers white, 3-3.5
mm long, the tepals slightly dis-
similar; leaves 3-5 cm long, the
petioles 3-6 mm long, deciduous
on the lower portion of the stem;
Mancos Shale hills east and south
of Wellington, Carbon Countv.
Utah

9. E. lancifoliiim.
FF. Involucres 3-4 mm long; inflores-
cences open with few long
branches; basal stem leaves usual-
ly persistent; Duchesne County.
Utah, and adjacent areas
G. Involucres 3.5-4 mm long;
flowers white, 3.5-4.5 mm
long, the tepals slightly dis-
similar; leaves 3.5-7 cm long.
the petioles 5-10 (18) mm
long; Bad Land Cliffs,
Duchesne County, Utah

10. E. hylophilum.

GG. Involucres (2.5) 3-3.5 mm
long

H. Flowers yellow, 2.5-3
mm long, the tepals es-
sentially similar; leaves
2.5-4 cm long, the peti-
oles 4-7 mm long;
Indian Creek Canyon,
Duchesne County, Utah
\\. E. duchcsnese.
HH. 1- lowers white, or if yel-
lowish, then plants
from southern LItah
and northern Arizona,
or if from Duchesne
County, then tlowers
pale-yellow and leaf-
blades less than 3 cm
long; leaves mostly less
than 3 cm long, but an
occasional blade up to
4 cm long (various
forms of £". corymho-
sum)

8. £ corymhosum.
EE. Branches glabrous or floccose; involu-
cres glabrous without
F. Leaves tomentose below; stems,
among the leaves, floccose to
tomentose

G. Involucres 2-3 mm long;
flowers 2-3 mm long, cream
to pale yellowish-white;
leaves 3-6 cm long; erect
shrubs 3-5 dm high; Mowry
Shale, Dinosaur National
Monument. Uintah County,
Utah

12. E. sauriimm.

GG. Involucres 2.5-3.5 (4) mm

long; tlowers white. 2.5-3.5

(4) mm long; leaves 1.5-4

(5) cm long; southern Colo-
rado and adjacent northern
New Mexico eastwiud into
northern Texas

13. E. fciidleriaiium.
FF. Leaves as well as the entire plant
totally glabrous throughout (ex-
cept for the cottony-tomentose
leaf-buds in the axils of the
leaves), bright green and shining;
flowers bright yellow. 3-4 mm
long; red blow sand, San Rafael
Desert, Emery County, Utah

14. £■. smithii.

AA. Leaves broad, oblanceolate to lanceolate or elliptic,
cordate to nearly orbicular, the apices round or nearly so
B. Leaf-blades oblanceolate to elliptic or nearly orbicu-
lar. 1-4(5) cm long; widespread
C. Inflorescences 6-25 cm long, highly branched,
stems floccose, white but often drying black-
ish; leaves narrowly oblanceolate to oblanceo-
late, 2-4 cm long; gravelly to sandy hills, slopes
and flats east of the Continental Divide, South
Dakota, eastern Wyoming and western Nebras-
ka southward through eastern Colorado to
northern and central New Mexico

7. cffusiim.
CC. Inflorescences 1-5 (10) cm long, sparsely

HKK'.HAM YOUNG UNIVKRSITY SCIKNCK BULLETIN

liranchcd, or if with several branches present,
tlien divaricately arranged and blades nearly or
quite orbicular, stems mostly white-, silvery-,
brownish", or reddish-brown tomentose: leaves
oblanceolate to lanceolate or elliptic to nearly
orbjcidar. 1-3 (5) cm long; clay or gravelly
slopes, hills and Hats, southwestern Wyoming
and western Colorado southward into eastern
and southern Utah, and into northern Arizona
and New Mexico, with an isolated series of
popidalioTi in central New Mexico

8. E. coryiiihiisiini.
BB. Leaf-blades cordate, (1.5) 2-2.5 cm long, 1-2 (2.5)
cm wide, densely white-tomcntosc below, Hoccosc
and greenish- or brownish-white above; stems
brownish-white tomentose; infrequent in sandstone,
limestone, or other rocky areas of northern Arizona
from extreme western Navajo County westward
across central Coconino County to extreme easl-
central Mohave County

15. £■. joiicsii.

1. Eriogoiiimi leptophylluni (Torr. in .Sitgr.) Woot. &
Standi.

Eriogoinim Icptuphvlliiin (Tun. in Sitgr.) Wool. &
Standi., Contr. U. S. Natl. Herb. 16: 1 18. 1^)1.^^, based
on E. effusum Nutt. var. leptophyUiiin Torr. in Sitgr..
Rept. Exped. ZuTTi and Colorado Riv. 168. 1854. -
Hriognmtm miirothcciim Null. var. lepiopliylliiDi
(Torr. In Sitgr.) Torr. & Gray, Proc. Amer. Acad. Arts
8:171. 1870. - Near Zuni, McKinley County, New
Me.xico, 24 September 1851, Woodliousc s.ii. Holo-
type, NY! Isotype, GH!

Largo, rounded, heavily-branched shrubs 2-6 (S) dm high.
3-1(1 ( 1 2) dm across, the lower stems reddish-brown to brown
or more frequently gray to grayish-brown, woody, the bark
often exfoliating in large platelike segments, mostly leatless,
the upper branches herbaceous, slender to stout, thinly pub-
escent and green when young but often totally glabrous, es-
pecially at maturity; leaves solitary or in fascicles on short
dwarf shoots, rather closely grouped along the lower 1/2 to
2/3 of the herbaceous stems, the leaf-blades thin, linear to
linear-oblanceolate. (1.5) 2-6 cm long. (O.S) 1-2.5 (3) mm
wide, densely to thinly white-tomcntose below, the midveins
obvious, thinly pubescent above when young or glabrous,
glabrous and green above by anthesis in all, the margins en-
tire, tightly revolule, the bases and apices sharply acute, the
leaves persistent, but soon deciduous in post-anthesis. the
petioles short, 0.4-1.1 mm long, membranaceous and light
yellowish-brown, glabrous, the petiole-bases triangular to del-
toid, 1-1.5 mm long. 1-2 mm wide, slightly tomentose to
glabrous without, light to dark brown, tomentose within,
more or less clasping the stems; //ou'cto;^ stems slender. l-S
cm long, green and glabrous, the area below among the leaves
remaining tomentose; inflorescences cymose, dense and
broomlike, congested with numerous shorl-internode
branches, 2-12 (15) cm long, 4-15 (30) cm wide, trichoto-
mously branched throughout, green and glabrous; hracls
.scalelike, ternate, 1-4 mm long, 0.4-1 mm wide, linear to
elongate-triangular, glabrous within and without, connate at
the bMi:\ peduncles lacking; /»i'o/((cn'.s solitary, narrowly tur-
binate, 2-3 mm long, 1.1-1.7 (2) mm wide, glabrous and
green within and without, the 5 acute teeth 0.3-0.7 mm long,
the bractlets linear, 1.5-3 mm long, minutely fringed with
short capitate gland-tipped cells, pale yellowish, the pedicels
2.5-5 mm long, glabrous; /7ouw.? while with greenish-brown
midribs and bases, 2.5-4 mm long, glabrous witliin and with-
out except for a few microscopic glands at the base of the

united tepals, the tepals essentially similar, oblong to narrow-
ly obovate, the outer whorl of tepals 1.4-1.7 mm wide, the
inner whorl of tepals 1.1-1.4 mm wide, united only at the
base of the tlower; stamens long exserted, 3-6 mm long, the
filaments subglabrous to sparingly puberulent basally, the
anthers reddish or light pink, 0.5-0.7 mm long, oblong;
achenes brown, 3.5-4 inm long, the globose base tapering to a
long. 3-angled beak. - Representative collections: Arsene
IMIO (G)', 19433 (P, US); Barnehy 129M (CAS, NY);
Biande^ee 12128 (MO, \iQ^\Castcltcr s.n. (UNM);£'a.v/ivooJ
and Howell 6W9 (CAS, Gil, P()M. IIC. US); Gooddini; 46 72.
-/6.V.5 (ARIZ);yo»<'.v 493S(\i\\. BR, POM, VS)\Mielwels 890
(ASU); Reveal and Davidse 924. 929. 942 (BRY. LL. NY,
U\C)\Standley 7342 (GH. MO. UC. US); Weber 5223 (CAS,
COLO. (ill. ISC, OKL, UC, US); /;/<A- s.n. (CAS, MO. NY.
US).

Distribution.

Dry clay Hats, washes and slopes in pliiyon-juiiiper
woodlands, in Bernarlillo, Santa Fe, and Taos coun-
ties. New Mexico, westward into Apache, Navajo, and
eastern Coconino counties, Arizona, and northward
and just entering Colorado in Montezuma County
near Four Corners and thus likely in extreme south-
eastern San Juan Comity, Utah, from 4500-6000 feet
elevation. Flowering frtim July to October. Figure 1 .

The type of the Broom-head Buckwheat, Erlogo-
iiiiiu leptophylluni. was collected by Dr. Samuel W.
Woodhouse, the naturalist assigned to Captain
Lorenzo Sit greaves on the latter's reconnaissance west
ot' the Indian village ZuTii across northern Arizona to
the Colorado River. The collection was made on 24
September 1851 , and according to Sitgreaves' Report
(185.>). the expedition was near Zimi, and Wood-
house ;ipparently obtained his collection in low
moimtaiiis east ot the village. In Torrey's report of
the pkint collections made by the naturalist, he
placed this buckwheat under F.. cfj'iisuiii. but the il-
lustration accompanying the description was of E.
hrerlcaiilc and not his var. Icpinpliylliini. Torrey
(185.^) stated that the original specimens were found
both at Zimi and on the San Francisco Mountains in
northern Arizona; however, no specimens of this spe-
cies from Arizona have been fomid among the Wood-
house material or among the other specimens in the
Torrey Herbarium collected prior to I85.v The \;ni-
ety was based on fragmentary material.

The var. leptophylhim was transferred to Eriogo-
tiiiiu iiiicrnlhcciim by Torrey and Gray in IS70. but
was elevated to the species r;mk b\- Wooton and
Standlev' ( \^>\}) while they were working on the flora
of New Mexico. Subsec|uent tlorislic works have con-
tinued to maintain the specific r;mk fur this pLinl.

EriiigoniiDi leptophylhim may be looked upon as
the most primitive extant member of an evolutionary
line that extends through /:'. chivclhittim to E. hicolor,
E. riplcyi, and perhaps to /;'. cncifoliiim. The origin of
/;'. Icpiophylluin is to be sought among the various
forms of ancient entities associated with the develop-
ment of £. mkrothecitiit. and most likely those forms
;issociated with var. foliosuin. This concept is sup-

BIOLOGICAL SERIES. VOL. I 3. NO. 1 NOTES ON I'RIOCION UM - VI

-^ ^

_J

I

Fig. 1 Distiibulioii map o( Urioguiiiim leptophyllum.

ported in pari by the appearance ot^ tlie young speci-
mens of botii buckwheats which tend to resemble
each other in their long leaves, narrow involucres, and
rather compact inflorescences. The demarcation be-
tween E. Icpiophylliiui and E. clavellatiim is sharp,
but allowing for evolutionary divergency, it is pos-
sible to speculate such an origin. The linear arrange-
ment of the species in the Section Corymbosa could
be reversed with E. leptophylhun following E.
microtheciim. but as those species following E. mi-
crothecwn (i.e.. E. effuswn and the E. corymbosuin
comple.\) exhibit a stronger tendency for a more di-
rect relationship, it simply proved more convenient to
arrange the species as seen in this revision.

In the iK\i,Eriog<miuu leptophyllum is distinctive
and not likely to be confused with any other species.
The plants are rather tall and roundish, with only the
upper portions bearing herbaceous stems. The inflor-
escences are compact and somewhat broomlike in
that they are composed of numerous erect branches
bearing short, flower-bearing nodes and internodes.
The result is a shrub, which in anthesis, becomes a
mass of white flowers. This produces a colorful sight
on the red or gray gumbo clay hills in northwestern
New Mexico and adjacent Arizona as the plants stand
out in brilliant contrast against the color of the soil.
It would seem that the horticulturalists have over-
looked a beautiful plant that might make a unique
addition to the garden.

2. Eriogonum clavellatuni Small

EriogDimin ehirellaliim Small, Bull. Torrey Bot. Club
25:48. 1898. - "Barton's Range," likely between
Comb Wash and Lime Creek, San Juan County, Utah,
13 July 1895, Eastwood 132. Holotype, NY! Iso-

types, CAS, GH, MO, NY, UC, US!

Low-rounded, lieavity-branched pofygamo-dioecioiis sub-
sfirubs 1-2 dm liigti, 3-8 dm wide, the tower stems reddisli-
browii. woody, tiie baric exfoliating in long, loose strips,
often feafy, the upper branches herbaceous, slender, thinly
tloccose to glabrous: leaves solitary or in fascicles on short
dwarf shoots, scattered along the lower .V4 of the herbaceous
stems, somewhat scattered and evenly-spaced but in fascicles
on the woody portion of the plant, the leaf-blades oblancco-
late, 5-12 (15) mm long, 0.8-1.7 (2) mm wide, densely
white-tomcntose below, the midveins totally obscured by the
tomentum, thinly pubescent and green above when young,
becoming glabrous by anthesis, the margins entire, tightly
revolute and completely enclosing the lower surface, the
apices rounded to obtuse, the bases acute, the leaves persis-
tent, the petioles short, 0.5-1.5 mm long, light greenisli-
brown and thinly pubescent when young, becoming brown
and glabrous at maturity, the petiole-bases triangular, (1.8-1.2
mm long, 1-1..^ mm wide, slightly tomentose to glabrous
without, tomentose within, not clasping the slem^; flowering
steins slender, 0.6-2 cm long, slightly pubescent when young
but becoming glabrous at maturity, the area among the leaves
below becoming sparsely tomentose; inflorescenees
nmbellate-cymose, more or less compact and congested,
0.5-1.5 cm long, 1-2 cm wide, trichotomous, the rays 2-5 mm
long, green and essentially glabrous: /tracts scalelike, ternate.
linear, 1.5-2.5 (3) mm long, 0.3-0.8 mm wide, glabrous with-
out, sparsely tomentose within, connate at the base: i>ed-
iinelcs slender. 1.5-4 mm long, green and glabrous, erect: »i-

10

hku;ham younc. univi-rsity scikncI:; bulletin

volucres solitary, turbinate-cainpamilate, (3.5) 4-4.5 mm
long, 2.5-4.5 mm wide, green ami tllaliroiis within and with-
out, the 5 acute teeth 0.6-0.9 mm long, mcmbranaicmis
along the margins, the bractlcts linear-oblanceolate, hyaline.
2-4 mm long, fringed with short eapitate gland-tipped celK.
the pedicels .1.5-7 mm long, glabrous; flowers white with
greenish-brown to reddish-brown midribs and base.s, 3-3.5
mm long, glabrous within and without except for scattered
microscopic glands along the midribs within, the tepals dis-
similar, the outer whorl of tepals broadly obovate to nearly
fan-shaped. 2-2.5 mm wide, the apices rounded to retuse. the
inner whorl of tepals oblanceolate to spathulatc, 0.9-1.5 mm
wide, slightly shorter than the outer whorl, united about 1/4
the length of the ({own: slameiis long exscrted, 3-fi mm long,
the filaments sparsely pilose basally, the anthers reddish (o
light pink, 0.5-0.6 mm long, oblong; achcnes light brown.
3-3.5 mm long, the globose base tapering to a long, 3-angled
beak. Representative collections: Harrison cl al. IU359
(US); Mamrc .^'.S'j.i' ( B 1< Y , GH, NY, \JC, UTC, \\TV):Reri'al
et al. S40 (ARIZ, BRY, (AS, CJH, MO, NY. OKL, KM, RSA,
UC, US, UTC, WIL!).

Distribution.

Li)vv-rolliiig clay liills and slopes along dry washes
in and art)iind Comb Wash west to Lime Creek, south-
west oF Bluff, San Jti;ni County. Utah, from
4,500-5,200 feet elevation. Flowering from April to
June. Figure 2.

I'lg. 2. Disli ibulmii ni.ip nt /ic/c.i,""""" cliircHiilliiii.

The type of the CItih-lcafed Buckwheat. F.riogi^-
imm clarc'llaliiiii. was collected between Comb Wash
and Lime Creek-or at least it would appear that is
the case. Alice Eastwotid (1896) staled that her "/:'.
mcanisli" (as she called the collection) was gathered
■"on ;t rocky hill on Barton's Range, between Epsom
and Willow creeks," but it is diftlctilt to determine
what her Epsom and Willow creeks were. In retracing
her travels, it would seem that after leaving Btitler
Wash she traveled up onto the mesa east of Comb
Wash and then down into Comb Wash itself. From
here, she traveled with Alfred Wetherill acrtjss Lime
Ridge, reaching Lime Creek, which, it is thought, she
called Willow Creek. As E. clavcllaliim occurs on the
western rim of Comb Wash and thus on the eastern
edge of Lime Ridge, it is proposed that this is the
type locality. Nevertheless, the species is rather rare.

both in the field and herbaria and is known t)nly from
the collections cited above.

The species is the largest ;ind most sliiubby of
those laxa having elongated peduncles, tight rcvolule
leaves, and dissimilar whorls of tepals. It is I'elt that
Eriogonuni clavellatiim evolved froin E. leptophyllum
(or some form related to that entity), and the other
members of this group which follow can be looked
upon as isolated modifications of the ancestoral type
which gave rise to, and evolved tVom, /:'. chiveUatuin.

In Ihe field, ihis species is quickly recognized by
its bright green appearance, whitish flowers, and low
stature. These morphological features, plus the re-
stricted distribution and the early tlowering of this
species, easily distinguishes Eriogonuni chivclhuuni
from all other species in the genus. The low-roimded
subshrubs are easily seen on the reddish soils of Comb
Wash, and in the spring of the year seem like small
white boulders on the hillsides when viewed from a
distance. The Comb Wash area is a pleasant place to
collect, and richly rewards the more diligent botanist
who cares to venture into the area at all seasons.

3. Eriogonum bicolor M. E. Jones

Eriogonuni bicolor M. E. Jones, Zoe 4:281. 1893. —
Eriogonuni niicrolliecuni Nutt. ssp. bicolor (M. E.
Jones) S. Stokes, Gen. Eriog. 75. 1936. - Thompson
Springs [now Thompson], Grand County, Utah, 7
May 1891, A/. E. Jones s.n. Holotype, POM! Isotypes.
A,MO,NY,PH, UCUS!

Low-rounded, heavily-branched pulvmate, polygamo-
dioecious subshrubs, 2-6 cm high, 5-20 (30) em across, the
lower stems reddish-brown to light brown, woody, the bark
exfoliating in long loose strips, leafy. Ihe upper branches her-
baceous, slender, tomentose; leaves solitary or in fascicles on
short dwarf shoots, scattered along the lower 1/2 to 3/4 of
the herbaceous stems, somewhat closely placed and con-
gested, the fascicles restricted to the tips of stemless caudices
or on the woody portions of the plants, the leaf-blades
liiicir-oblanceolate to narrowly elliptic, 5-12 (15) mm long,
1-2 (3) mm wide, densely white-tomentose below, the mid-
veins totally obscured by the tomentum. slightl\' less tomen-
tose and white to whitish-green above, the margins entire,
revolute and often completely enclosing the lower surface,
the apices and bases acute, the leaves persistent, the petioles
short. 1-1.5 mm long, light greenish- lo yellowish-brown and
thinly pubescent when young, becoming subglabrous at ma-
turity, the petiole-bases triangular, 0.5-1 mm long and wide,
slightly tomentose to glabrous without and light brown and
densely tomentose within, not clasping the stems; //oiecn/;.?
slenis slender, 3-22 mm long, densely tomentose, remaining
tomentose among the leaves below as well; inflorescences
umbellate-cymose. more or less compact and congested, 5-10
mm long, 5-15 mm wide, dichotomous or trichotomous,
rarely reduced to a single ray, the rays 1-5 mm long, tomen-
tose; liracis scalelike, ternate, 0.7-1.3 nun long. 0.2-0.6 mm
wide, linear, glabrous to sparsely pubescent without, tomen-
tose within, connate at the base; pcrfK^c/cv slender, 1.5-3 (4)
mm long, tomentose when young, becoming suliglabrous al
matiuity; involucres solitary, turbinate-campanulate. 2-4 mm
long, 1.5-3 mm wide, tomentose to subglabrous or glabrous
without, glabrous within, the 5 acute teeth 0.4-0.7 mm long,
membranous along the margins, the bractlets oblanceolate.
1.5-3.5 mm Ions;, hirsutulous wilh short acute cells scattered

BIOLOGICAL SKRIKS. V(_)L. I J. NO. I NOIKS ON I-KIOGONUM - VI

11

among gland-tipped (.ells. li\almo. Ilie pedicels 3-6 nun long,
glabrous: flowers white with gieciiisli-lirovvn to reddish-
brown midribs and bases, 2.5-4 mm long, glabrous within and
without except for scattered glands along the midribs within,
the tepals dissimilar, the outer whorl of tepals broadly obo-
vate to nearly orbicidar, (2) 2.5-3 mm wide, the apices
rounded to emarginate, the inner whorl oblanceolatc to nar-
rowly elliptic, 1-1.5 mm wide, slightly shorter than the outer
whorl, united about 1/4 the length of the flower: stameiu
long e.xserted, 3-5 mm long, the filaments glabrous to pilose
basally, the anthers reddish, 0.5-0.6 mm long, oblong;
achenes light brown, 3-3.5 mm long, the narrowly globose
base tapering to a long, 3-angled beak. - Representative col-
lections: AtwooJ 1340 (BRY): Bariiebv 13134 (CAS, NY):
Cronqiiisl 9(MI (MICH. NY, RSA, UC. UTC, VVS, VVTU):
Harrison 11149. 11409. 1 1515. U 705 {BRy . NY): Higgiiis
and Reveal 12S2. 1295 (BRY): Jones s.n.. on S May 1914
(CAS. GH, MICH, NY, POM, US): on 28 May 1914 (CAS,
MICH. POM, UC. US): Magiiire 1S241 (GH, NY, UC, UTC,
WTU): Ripley and Barnehv 8635 (CAS, NY. inC):Siokes
s.n. (NY, UC, US): Welsh 3976. 7072 (BRY).

Distiibiitioii.

Dry sandy clay to giiinbd clay flats and low rolling
hills in eastern Utah and adjacent western Colorado,
from Castle Valley and the San Rafael Swells of Car-
bon and Emery counties into the Grand River Valley
of Grand County, Utah, and Mesa County, Colorado,
southward into Utah to Capitol Reef National Monu-
ment and the Green River Desert of Wayne County,
the Waterpocket Fold area near Bullfrog, the foothills
of the Henry Mountains of Garfield County, and en-
tering San Juan County along the Colorado River in
Canyonlands National Park. Flowering from April to
June. Figure 3.

The type collection of the Bicolored Buckwheat.
Eriogomim bicolur. was possibly distributed to her-
baria on two different occasions by Jones; once with
the label data as "Thompson Springs" and again as
"Cisco." The first is cited in the original publication
(Jones, 1893), and the handwritten labels bearing
Thompson Springs are found in several herbaria (such
as Arnold Arboretum, Gray Herbarium. The New
York Botanical Garden, and Tlie Academy of Natural
Sciences). The second label is printed and has the

words "N. Sp. Type" found below the scientific
name. .According to Jones (19(i5). he was at West-
water in eastern Grand County, Utah, on 6 May
1891. ;ind at Thompson Springs [now Thompson on
road maps] on 7 May. No mention is made of Cisco,
but if Jones was traveling by train or if he was going
by buckboard, he had to pass through Cisco, the
largest town in the area. It is considered at this time
that the printed labels were prepared later and the
plants distributed to other herbaria (such as The Mis-
souri Bi)tanical Garden, the University of California
at Berkeley, and the United States National Museum)
sometime after the otliers had been sent out. Never-
theless, they are still part of the original type collec-
tion gathered near Thompson, and Jones was simply
identifying the location as being near the largest town
in the area at that time.

Eriogomim bicolor is a low, pulvinate species
which flowers in the spring of the year, being at the

\

\
/

4

^J

Fig. 3 Distribution map of Eriogoni/ni bicolor.

12

HKICIIA.M YOUNG UNIVliKSlTY SCIKNCF. BULLEIIN

peak of antliesis in May. It is rather closely related to
E. clavellatiim. differing essentially only in the degree
of pubescence and stature. Both occur on clay soils,
but so far as is known, their respective ranges do not
overlap.

During the course of tiiis study, a single collection
was discovered which seems to represent a new spe-
cies for the Hriogomim mkrotheciiin complex. This is
a low, pulvinate subshrub also related to E. ckivclla-
iiiDi as it possess the same type of leaf construction
and. to a great degree, a similar type of pubescence as
well. The distinctive features of this buckwheat are
the narrowly turbinate involucres, the flowers with
the essentially similar tepals, and the large globose
flower bases. The plants are more pulvinate and not
as woody as E. chiveUaiuiu. nor do they have the
same type of inflorescence. In most of these features,
this seemingly new species also differs from E. hi-
color. At present, this new entity is known from a
single collection made by Harold Gentry from near
Hotchkiss, Delta County, Colorado. However, recent
attempts to rediscover the population have failed. Dr.
Larry C. Higgins has discovered a new species of
CryptiDitha in the area while searching for EriogoiUDii
and thus this part of Colorado seems to need addi-
tional collecting. Hopefully, additional material will
come to light, whicli may form the basis of a future
publication.

Of all the species in this complex, the Bicolored
Buckwheat is the only one that has been studied suf-
ficiently to allow some comment on the polygamo-
dioecious condition. The female plants are male ster-
ile, that is. the anthers do not form or function and
the filaments tail to elongate: however, vestigial re-
mains of the anthers and filaments are clearly visible.
The male plants, on the other hand, produce both
function stamens and ovaries. Collections at the Brig-
ham Young University Herbarium which are only fe-
male (at least in part) are Cottam 2000: llarrhnii
11149. 1 1206. 11409: lliggim ami Reveal 1295. and
ItWiV; 7072. Otlen the two conditions occur on
plants growing tc)gether, but at least on one occasion,
plants found on the .San Rafael Desert were only fe-
male with no herntaphiditic plants discovered. Unfor-
tunately it was too early in the year to determine if
such plants would have viable seeds; still, it could be
that some populations of Eriogonum hivolor are
apomictic, although it is too early to make such sug-
gestions. Nevertheless, this evolutionary condition
seems to point to the fact that this species represents
an end point, or at least a highly specialized cimdilion
in the genus, even if it occurs in only a few members
of the population.

4. Eriogonum ripieyi J- T. Howell

Eriogdiiiim riplcyi J. T. llovvcll, Leall. W. Bot. 4:5.
1444. On the edge of sandstone mesas 13 miles
southwest of Frasier's Wells. Coconino County,

Arizona, 13 .May 1^43, Riplev & Baniebv 5226.
Holotype.CAS!

Low-depressed, heavily branclied, possibly polygamo-dioe-
cious subshrubs 0.5-1.5 (2) dm high, 0.5-3 (5) dm across, the
lower stems reddish-brown or grayish, woody, the bark exfo-
liating in long loose strips of plalelike segments, leafy nearly
throughout, the upper branches woody, slender to stoutish,
suhglabrous to glabrous, the bark grayish; leaves solitary or
much more commonly in fascicles on short dwarf shoots,
these alternately scattered along the upper 3/4 to 7/8 of the
upper woody steins, closely placed and congested, the Icaf-
bladcs narrowly oblaneeolate, 2-6 mm long, 0.5-1 mm wide,
densely vvhite-tomentose below, thinly lloccose to villous and
greenish above, the midveins totally obscured by the toinen-
tum, the margins entire, strongly revolute and completely
enclosing the lower surface, the apices and bases acute, the
leaves persistent, the petioles short. (0.5) 1-1.5 mm long,
light greenish- to yellowish-brown and thinly pubescent when
young, becoming subglabrous at maturity, the petiole-bases
triangular, 0.5-1 mm long and wide, slightly tonientose to
glabrous without, light brown and membranous, densely
tonientose within, not clasping the stems; flowering stems
indistinguishable from the other branches, woody, grayish;
infloreseenees reduced cymose-umbellate, consisting almost
entirely of a single involucre arising from the apex of each
dwarf shoot; bracts lacking; peduncles slender, thinly tloc-
cose, 1-10 mm long, erect, terminating the dwarf shoot; /»ro-
lucres solitary, campanulate, 3-3.5 mm long and wide, thinly
floccose or villous to suliglabrous without, glabrous within,
the 3-5 unequal acute teeth 0.7-1 mm long, more or less
membranous along the margins, the bractlets linear, 2-3 mm
long, hirsutulous with short acute cells scattered among the
gland-tipped cells, hyaline, the pedicels 3-5 mm long, glab-
rous or sparseh' pilose at the base; flowers white with red-
dish-brown midribs and bases, 3.5-4.5 mm long, glabrous
within and without, the tepals dissimilar, the outer whorl of
tepals suborbicular, 3-3.5 mm long, 3.5-4 mm wide, the
apices rounded to emarginato, the inner whorl of tepals
broadly obovate. 2.8-3.2 mm long. 2-2.5 mm wide, united
about 1/5 the length of the tlower; stamens exserted. the fila-
ments villous to densely pilose basally. tlie anthers reddish.
0.5-0.6 mm long, oblong; aclienes light brown to brown.
2-2.5 mm long, the narrow base tapering gradually to a
short. 3-angled beak. - Representative collections: Harris s.n.
( ASLl. BRY, n\\: Ripley and Barneby S445 (CAS).

Distribution.

Sandy clay soil on the edge of sandstone mesas
associated with pinyon, about thirteen miles southwest
of Frasier's Wells, at 6,000 feet elevation, Coconino
County, and at the north end of the Horseshoe Dam
area on calcareous soil, Yavapai County, Arizona.
Flowering from April to June. Figure 4.

The most remarkable characteristic of the Ripley
Buckwheat is the bractless flowering stems and the
resulting inflorescence which is essentially composed
of a single terminal involucre. This trend is first seen
in Ericgomim clavellaimn and carried to this extreme
in /:'. ripieyi. The ebracteated condition is likely due
to the shortening of a line fiowering stem, leaving
only the peduncle and the uppermost whorl of leaves
of the fascicle may be true bracts. However, based on
gross morphology, it is impossible to determine at
this time whether or not the leaves are all "leaves" or
the upper whorl is "foliaceous bracts." As noted by
Howell (I ''44), this situation is similar to the condi-

BIOLOGICAL SERIES. VOL. 1 3. NO. I NOTES ON ERIOGONUM - VI

13

E-ig. 4 Distribution map o( Eriogoniiin ripleyi. Coconino and
Yavapai counties, .Arizona.

tion found in E. caespitosum Nutt., but tills is only
another example of parallel reduction of similar struc-
tures in unrelated species groups in the genus.

Of the remaining vegetative features, the arrange-
ment of the leaf fascicles with the involucres extend-
ing out from these on slender peduncles represents
another extreme in the reduction of the inflores-
cences. It is possible to imagine that at one time short
branches radiated out along the main woody branch-
es, each containing a small-branched inflorescence.
However, with the reduction of the shorter branches,
the leaves were reduced to mere fascicles, and the
inflorescence reduced to a single involucre. Tl'is trend
is not seen in other species in the Hriogimiim micro-
ihecum complex although it is hinted at in /.". hicolor.

In 1969 a second location of this species was dis-
covered in Yavapai County, Arizona. Until then, Erio-
gonuiu ripleyi was known only from the sandstone
mesas southwest of Frasier's Well. The plants from
Yavapai County are larger and more robust than
those from the type location, and the two differ in
some minor ways as well. Whether or not the difter-
ences are important will have to remain until the nec-
essary field work can be done and the variation
studied in detail.

5. Eriogonum ericifolium Torr. & Gray

Low spreading pulvinate. proliably polygamo-dioecious,
subshrubs 0.5-0.9 (1.2) dm high, 0.8-2 (3.5) dm across, the
lower stems reddish-brown or gray, woody, the bark extoMat-
ing in long loose plates, essentially leafless, the upper branch-
es herbaceous, slender, tloccose: leaves solitary or in fascicles
on short dwarf shoots, scattered along the lower 3/4 to 7/8
of the herbaceous stems, rather closely crowded, the fascicles
restricted to the tips of the stemless caudices, the leaf-blades
oblanceolate to narrowly elliptic, 5-8 mm long, O.S-1.5 (1.9)
mm wide, densely white-tomentose below, the midveins evi-
dent and less pubescent, glabrous and green to tloccose and
whitish-green above, (he margins entire, revolute or at least
with thickened margins, fully or only partially obscuring the
undcrsurface of the blade, the apices and bases acute, the
leaves persistent, the petioles short. 1.5-2 mm long, tloccose,
the petiole-bases elongate-triangular, 1-2 mm long, 0.8-1.2
mm wide, light tan to greenish-brown, glabrous or thinly
pubescent, light tan and densely tomentose within, not clasp-
ing the stems; flowering stems slender, 0.3-2 cm long, tloc-
cose to slightly tomentose, the area below among the leaves
remaining tloccose to tomentose or rarely glabrate; in flares-
censes umbellate-cymose, compact and congested, 0.5-1 cm
long, 0.5-1.5 cm wide, spar.sely tomentose to tloccose, di-
cliotomous, the rays up to 7 mm long; brads ternate, scale-
like, 1-1.5 mm long, 0.4-0.6 mm wide, linear, tloccose with-
out, tomentose within, connate at the bdf.e\ peduncles lack-
ing; involucres solitary, turbinate, 2.5-3 mm long, 1.5-2 mm
wide, slightly pubescent without, glabrous within, the 5 acute
teeth 0.4-1 mm long, not membranous along the margin, the
bractlets linear, 2-3 mm long, hyaline, minutely fringed with
capitate gland-tipped cells, the pedicels 2.5-4.5 mm long,
glabrous; flowers white with reddish-brown midribs, becom-
ing tinged with pink or red to rose at maturity, 2-2.5 mm
long, gFabrous within and without except for a few scattered
microscopic glands along the midribs within, the tepals dis-
similar, the outer whorl to tepals broadly obovate to nearly
orbicular, more or less claw-shaped in some, 2-2.5 mm wide,
the apices rounded to retuse or emarginate, the bases obtuse
to truncate or cordate, the inner whorl of tepals oblanceolate
to oblong, 0.8-1.2 mm wide, about as long as the outer
whorl, united about 1/5 the length of the tlower and forming
a subglobose base; stamens slightly exserted, 2-3 mm long,
the filaments pilose basally, the anthers reddish, 0.5-0. b mm
long, oblong; achenes liglit brown, 2-2.5 mm long, the nar-
rowly globose base tapering to a papillate, 3-angled beak.

Distribution.

Dry gravelly to rocky places mainly in pinyon-
juniper woodlands from 3,000-6,600 feet elevation,
extreme western Mohave County eastward across
Coconino County and southern Navajo County, and
in Yavapai County, Arizona. Flowering from August
to October. Figure 5.

Unfortunately the Heath-leaf Buckwheat, Eriogo-
intiii ericifolium. has not been investigated in the
field. Specimens are infrequently encountered in
herbaria, and I liave yet to be in northern Arizona at
the right time of the year to study the species in the
field. The typical phase is known only from two
collections, both made in the last century. The more
widespread variant, var. piikhntm. is not abundant,
and preliminary studies have done little more than
indicate its relationship to other members in this
species complex. Hopefully, this plant can be fully
studied in the field, but based on information now at
hand, the following treatment seems correct.

14

bkigham young univi:ksh y science bulletin

Fig. 5. Distribution map of Eriogonum ericifolhiiit. with var. p((/(7;n/»;f solid circles) and var. ericifntiinn (open
circles). Nortlicrn .■\rizona.

Key to tlie Varieties of t'riogonuin ericifoliuin

A

llppcr surface of tlie leaves tloccose; outer tepals nearly
orbicular, claw-shaped; infrequent, extreme western
Mohave County eastward to southern Navajo County,
Arizona

5a. var. piiUhrtim.
AA. Upper surface of the leaves glabrous: outer tepals obo-
vate, now claw-shaped; rare, Yavapai County. Arizona

5b, var. cricifolium,

5a. Eriogonum ericifoliiim var. pulcliruin (Eastw.)
Reveal

Eriogonum ericifoliiim Torr. & Gray var, pulclmim
(Eastw.) Reveal, cumb. iiov., based on E. pulciinim
Eastw., Proc. Calif. Acad, Sci, IV, 20: LW, 1931.
Eriogonum microthecum Nutt. ssp, pulchrum
(Eastw.) S, Stokes, Gen, Eriog, 76, 1936, - Eriog-
onum mearnsii Parry in Britt, var. pulchrum (Eastw.)
Keain. & Peebl., J, Wash. Acad, Sci, 29:474. 1939, -
Near the Meteor Crater, near Canyon Diablo, Coco-
nino County, Arizona, 21 October 1928, Eastwood
15746. Holotype, CAS! Isotypes, GH, NY, POM, US!

Low-spreading suhshruhs 0.8-1.2 dm high, woody at the
base; leaves 5-8 mm long, lloccose and whitish-green above,
slightly revolute or with thickened margins;/7oum- 2-2.5 mm
long, the outer whorl of tepals nearly orbicular, more or less
claw-shaped, 2-2.5 mm wide. - Representative collections:
Baniehy 12964 iCAS): Bolirer 1097b. 11J2. 1133. (ARIZ);
Cottam 13S36 (CAS): Danow 3292 (ARIZ, CAS): Eastwood
ami Howell h9l)4 (CAS); Jaeger s.ii. (MNA); Lehto 3462
( ASU), jVy<V ( ASU, W\}): Michaels 814 (CAS).

Distribution.

Dry gravelly and rocky places t'roni Toroweap
Point, Mohave County, souliieastwardly into east-
central Coconino County in the Painted Desert region
northwest and west of Winslow, and continuing
southeastward in Navajo County to near the Mogol-
lon Rim in ihe vicinity of Snowflake and Heber,
.'i, 400-0, (>00 feet elevation, northwestern Arizona,
Elowering from August to October, Figure 5.

The var, pulchrum is poorly known to me and is in
need of much critical field and herbarium study. It is
seemingly endemic to northern Arizona but some spe-

BIOLOGICAL SERIES. VOL. 1 3. NO. 1 NO lES ON ICRIOCiONUM - VI

15

cimens of Hriogoniim inicn)thcciim do appear to be
similar, especially some found in southern Utah. The
variety seems to bridge the gap between E. leptophyl-
Iwn and E. micwthecum througli the latter's var.foli-
osiini. However, this point is higiiiy speculative. Tiie
VAX . pulchnim also grossly resembles E. hicolor. but in
this case, the similarities are likely due to parallel evo-
lution and not because of any direct association.

The relationship between var. pidchmm and var.
ericifolium seems clear, with the rare var. erlcifoliinn
being a slightly more depauperate, glabrous phase
occupying an area disjunct from var. pitlchrum. Based
on existing herbarium material, the placement of var.
piilclinim under this species seems valid, but addition-
al studies will be needed to confirm this belief.
5b. Eriogonum ericifolium var. ericifolium
Eriogomun ericifolium Torr. & Gray, Proc. Amer.
Acad. Arts 8:170. 1870, as ericaefoliuin. - Eriogo-
num fasciculatuin Benth. var. ericifolium (Torr. &
Gray) M. E. Jones, Contr. W. Bot. 13:14. 1903. -
Eriogonum microthecum Nutt. ssp. ericifolium (Torr.
& Gray) S. Stokes. Gen. Eriog. 75. 1936. — Near Fort
Whipple [now Whipple], Yavapai County, Arizona.
25 September 1865, Coues & Palmer 581. Holotype,
GH! Isotype,MO!

Eriogonum mearnsii Parry in Britt., Trans. New
York Acad. Sci. 8:72. 1889. - Eriogonum microthe-
cum Nutt. ssp. mearnsii (Parry in Britt.) S. Stokes,
Gen. Eriog. 75. 1936. — Near Fort Verde [now Camp
Verde], Yavapai County, Arizona, 1 1 October 1887,
Mearns 1 79. Holotype, NY! Isotypes. NY!

Low-spreading siibshrubs 0.5-0.8 dm high, woody at the
base; leaves 5-6 mm long, glabrous and green above, tightly
revoliite; flowers 2 mm long, the outer whorl of tepals obo-
vate. not claw-shaped, 2 mm wide. - Representative collec-
tions: Known only from the two cited type collections.

Distribution.

Probably in gravelly or rocky places on slopes,
known only from near Prescott and Camp Verde,
Yavapai County, Arizona, from 3,000-5,200 feet ele-
vation. Probably flowering from August to October.
Figure 5.

The type of the Heath-leaf Buckwheat, Eriogonum
ericifolium, was collected by Elliott Coues and
Edward Palmer near Fort Whipple in 1865. The mate-
rial is exceedingly fragmentary and consists of only
the upper herbaceous branches, but based on this,
Torrey and Gray (1870) described the species. A sec-
ond collection was made by IVIearns near Fort Verde,
and this was described by Parry in Britton's ( 1889)
paper on the Mearns collections gathered in Arizona
from 1884 to 1888. In 1903 Jones proposed the vari-
etal combination E. fasciculatum var. ericifolium per-
haps following Watson's (1877) suggestion that E.
ericifolium was a synonym of E. fasciculatum. I have
been unable to understand this conclusion, but since
Coues and Palmer did collect E. fasciculatum var.

polifolium (Benth. in DC.) Torr. & Gray at Fort
Whipple, Watson (and perhaps Jones) mistook this
collection to represent the type of E. ericifolium.

In the literature on the genus, the name Eriogo-
num ericifolium has been generally ignored. The vari-
ous treatments of the genus for Arizona in the past
have not attempted to place the species, or, as is the
more frequent case, the authors have simply not men-
tioned the name.

6. Eriogonum microthecum Nutt.

Low to tall, spreading to erect, open to compact, sparsely
to diffusely branclied herbaceous subshrubs to woody shrubs,
0.5-15 dm high, 0.6-1.3 (1.8) m across, the lower stems red-
dish-brown, light brown, yellowish-brown, to grayish, woody,
the bark exfoliating in loose platelike strips or long loose
strips, essentially leafless or leaves restricted to fascicles, the
upper brandies herbaceous, mostly slender, glabrous to floc-
cose or densely tomentose to lanate when young, remaining
so or becoming less densely pubescent as the plant matures,
often greenish at maturity; leaves solitary or in fascicles on
short dwarf shoots, scattered along the upper part of the
woody stems in some, becoming alternate and variously
spaced along the lower 1/3 to 3/4 (7/8) of the herbaceous
brandies, the leaf-blades various, linear to linear-oblanceolate
or narrowly elliptic to elliptic-obovate, 0.3-3.5 (4) cm long,
1-20 mm wide, variously white-tomentose below, the mid-
veins usually obvious and less pubescent and greenish, as
pubescent to slightly less so or nearly to quite glabrous
above, the tomentum white to brownish or reddish over the
greenish surface, the margins entire, variously revolute or
plane, usually with at least the margins thickened, the lower
surface completely enclosed in some, the apices mostly acute
but occasionally rounded or at least obtuse, the bases acute
or infrequently rounded, the leaves persistent, the petioles
short, 0.5-5 mm long, light greenish- to yellowish- or tannish-
brown and thinly pubescent to nearly or quite glabrous when
young, becoming usually less pubescent and more brownish
in age, the petiole-bases triangular, deltoid, or elongate-trian-
gular, 0.5-1.5 mm long and wide, slightly tomentose to glab-
rous without, various shades of brown but u.sually a less in-
tensive hue than the petioles without, cottony tomentose
within, not clasping the stem; flowering stems slender to
more or less stout, I-IO cm long, densely lanate to tloccose
when young, remaining so or becoming subglabrous or glab-
rous at maturity, the area among the leaves usually remaining
tomentose; inflorescences cymose, rather congested and com-
pact, more or less flat-topped, 0.5-6 (12) cm long, 1-10(13)
cm wide, trichotomous throughout or at the lower nodes
only with the upper branches dichotomously branched, the
internodes variously shortened above, tomentose to glabrous,
the involucres in the forks of the branches or at the tips of
the ultimate bracteated branches; bracts scalelike, ternate,
1-5 mm long, 0.5-2 mm wide, linear to triangular, tomentose
to glabrous without, tomentose within at least at the connate
base; peduncles, when present, slender and up to 1.5 cm long,
erect, variously pubescent or glabrous; involucres solitary,
turbinate, (1.5) 2-3.5 (4) mm long, 1.3-2.5 (3) mm wide, vari-
ously tomentose to subglabrous when young, becoming less
pubescent or more frequently merely floccose between the
angled ridges or even glabrous without, glabrous within, the 5
rounded to triangular teeth (0.3) 0.5-1 (1.7) mm long, often
with membranous margins, the bractlets linear to oblanceo-
late, 1-4 mm long, variously pubescent from only sparsely
fringed with scattered acute or gland-tipped cells to rather
hirsutulous with long white marginal cells, the pedicels 2-4.5
mm long. glabrous;/7oH'«'S white with green, greenish-brown,
pink, red, rose, or reddish-brown midribs and bases, becom-

16

BKKIHAM YdUNG UNIVIRSUY SCIKNCF, BULLETIN

ing tinged wilh, or fully colored with, pink to rose or even
orange in ago in some, or yellowish to liriglil yellow with
greenish-yellow midribs and bases, 1.5-3 (4) mm long, glab-
rous within and without except for scattered microscopic
glands along the midribs within, the tepal similar or slightly
dissimilar, the outer whorl of tepals oblong to obovate, 1.2-2
mm wide, the apices rounded to obtuse, the bases rounded to
more or less cordate, the inner whorl of tepals oblanceolate,
elliptic, or oblong, 0.8-1.5 nun wide, as long to slightly longer
than the outer whorl, rarely shorter, united about 1/5 to 2/5
the length of the tlower; stamens slightly to long exserted,
2.5-4 mm long, the filaments sparsely to densely pilose basal-
ly, rarely subglabrous. the anthers white, pink, rose, red. or
infrequently purplish, 0.35-0.5 mm long, oblong or nearly so;
achenes light brown to brown, 1.5-3 mm long, the narrowly
globose base tapering to a long, 3-angled beak.

Dislrilnitioii.

Widespiead in a variety of iiabitats from eastern
Washington southward to tiie mountains of soutiiern
California, hence eastward in the deserts, higii moun-
tain valleys, and mountain ranges to the Rocky
Mountains from western Montana southward througii
Nevada, Utah, western Wyoming, and western Colo-
rado into northern Arizona and New Mexico, from
(1,500) 2,200-10,500 feet elevation. Flowering from
June to October.

The Great Basin ^i\.ck\\\\ed{, Hriogonwn nikrothc-
ciim, as outlined and defined in this revision, is com-
posed of nine closely related and somewhat overlap-
ping varieties. The species is exceedingly complex,
and althougli it has been under intensive study for the
past five years, some entities as yet unnamed may
prove to be worthy of laxonomic cimsideration in the
future.

Hybridization within lindgoniiiii micr<>!hcciii)i is
limited, and is believed to occur only between var.
laxijloruin and vdt.fnliosian. This is thought to occur
in the zone of their overlapping geographical ranges
across southern Nevada and in northern Arizona
where var. laxiflonim from the north gradually
merges with vat.fdliosuni of the south. Hybridization
between /;'. microtheciim var. la.xijlonini and t'. hrevi-
caiilc var. hrcvicaiile is believed to occur in northeast-
ern Utah and adjacent southwestern Wyoming where
the two grow together. Supposed accounts of hybridi-
zation or introgression between E. inivrollurinu and
E. cffiisuni are unfounded.

Almost all of the infraspecific elements in Eriogo-
nitin microtheciim are difficult to consistently recog-
nize, especially in all of their various phases. The vast
majority of specimens can be distinguished and
placed, but as some forms tend to blend into each
other, these intermediates are often difficult to place.
Thus, plants from central Nevada, soutiiern Utah, and
northern Arizona are often impossible to place in
either var. laxiflonim or wdr.foliosum. Some fragmen-
tary specimens may prove difficult to place.

Some populations from totally different habitats
are morphologically somewhat similar. Plants from
the higli alpine reaches of the Sierra Nevada (called

var. alpimiiu) will approach var. lapidicola which
grows on volcanic mesas in southern Nevada. Forms
of var. microtheciim in the low mountains of north-
ern Oregon approach some plants from the foothills
of the Sierra Nevada in Inyo County. California,
which are recognized here as var. amhigiiiim. At the
other extreme, some populations which are thought
to represent a single variety may vastly differ from
one site to the next. In northern and eastern Utah
and portions of southern Colorado, var. foliosum is
only a few centimeters higli. However, through a
series of intermediate populations in southeastern
Utah and northern Arizona, this variety reaches
heiglits of a meter or more.

In summary, the varieties oi' Eriogoiiiim microthe-
ciim are interesting. They represent an example of the
kind of variabilily found in the genus and in plants of
the western Linited States in general.

Key to the Varieties o( Eriogoniim microtheciim

A. Flowers white, not yellow or yellowish

B. Tomentum whitish (see also var. alpiiniiii of the
high Sierra Nevada); widespread and common from
Washington and Montana southward to southern
California, northern fXiizona, southern Colorado,
and northwestern New Mexico
C. Leaves plane, infrequently revolute; stems and
inflorescences tloccose to glabrous; northern
populations of the species, from Washington
and western Montana southward to California,
extreme northern Arizona, and western Colo-
rado 6a. var. laxiflonim.
CC. Leaves revolute. rarely plane; stems and inflo-
rescences densely lanate to tomentose, or if
glabrous, then plants not in the range of the
above variant; southern populations of the
species, from southeastern California across
southern and central Nevada into southern and
eastern Utah, western and southern Colorado,
northern .Arizona, and northwestern and cen-
tral New Mexico 6b. var. foliosum.
BB. Tomentum brownish to reddish; scattered popula-
tions in southern California, central Sierra Nevada,
and from eastern California across Nevada to ex-
treme western Utah
C. Plants shrubby, 3-6 dm high

D. Stems and inflorescences tomentose when
young, becoming floccose at maturity;
flowers 1.5-2 (2.5) mm long; achenes
1.8-2.1 mm long; Panamint Range and the
Inyo Mountains, Inyo County, California
6c. var. panamiiitcnse.
|)D. Stems and inflorescences lanate to tomen-
tose even at maturity; flowers 2-2.5 (3)
mm long; achenes 2.5-3 mm long; San
Bernardino and San Gabriel mountains.
San Bernardino and Los Angeles counties,
(California 6d. var. corymlmsoides.

Ci'. Plants subshrubs. 0.5-1.5 dm liigli

D. Leaves elliptic, not revolute; involucres (2)
2.5-3.5 mm long; flowers (1.5) 2-3.5 (4)
mm long; southern and southeastern Cali-
fornia eastward across central Nevada to
extreme western Utah
E. Leaves elliptic to ovate, 5-10 mm
long, (2) 3-5 (6) mm wide; involucres

BIOLOGICAL SERIES. VOL. 1.1. NO. I NOTES ON ERIOGONUM - VI

17

(2) 2.5-3 mm long: tlowers (2.5) 3-3.5
(4) mm long; San Gabriel Mountains,
southern California

5e. var. johnstonii.
EE. Leaves elliptic, 3-7 mm long, 1-4 mm
wide; involucres (2.5) 3-3.5 mm long;
flowers (1.5) 2-3 mm long; Inyo
Mountains, Inyo County, California
across southern Nevada to extreme
western Utah 6f. var. lapidicola.

DD. Leaves linear-ohlanceolate to narrowly
elliptic, often revolute; involucres (1.5)
2-2.5 mm long; flowers 1.5-2.3 mm long;
central Sierra Nevada of Alpine, Tuolum-
ne, and Mono counties, California

6g. var. alpimim.
\.\. Flowers yellowish to yellow, not white

B. Plants tloccose to glabrous; leaves (3) 4-9 (12) mm
wide; involucres 2.5-3 mm long; achenes 2-2.5 mm
long; central Oregon eastward to the Idaho line, and
southward to northern Humboldt County, Nevada,
and Modoc and Lassen counties. California

6h. var. microthecum.
BB. Plants tomentose to floccose; leaves (2) 3-6 (8) mm
wide; involucres 2-2.5 mm long; achenes 1.5-2 mm
long; extreme west-central Nevada and adjacent
eastern California southward to Inyo County, Cali-
fornia, and Esmeralda County, Nevada

6i. var. ambiguum.

6a. Eriogonum microthecum var. laxiflorum Hook.

Eriogoiniin microtliecuin Nutt. var. laxiflorum Hook.,
Hooker's J. Bot. Kew Card. Misc. 5:264. 1853. - Eri-
ogonum microthecum Nutt. ssp. laxiflorum (Hook.)
S. Stokes, Gen. Eriog. 74. 1936. - "Rocky Moun-
tains of the Columbia in Oregon," actually from west-
ern Idaho or perhaps northeastern Oregon, probably
collected in August 1834. Nuttall s.n. Holotype, K!
Isotypes, BM, K!

Eriogonum confertiflorum Benth. in DCProdr.
14:17. 1856. - Eriogonum microthecum Nutt. var.
confertiflorum (Benth. in DC.) Torr. & Gray, Proc.
Amer. Acad. Arts 8:171. 1870. — Eriogonum micro-
thecum Nutt. ssp. confertiflorum (Benth. in DC.) S.
Stokes, Gen. Eriog. 75. 1936. — Along the Shasta
River, Siskiyou County, California, 1-4 October
1841. Brackenridge 1570. Holotype, NY! Isotype,
GH!

Eriogonum macdougalii Gand., Bull. Soc. Roy.
Bot. Belgique 42:191. 1906. - Eriogonum microthe-
cum Nutt. var. macdougalii (Gand.) S. Stokes, Gen.
Eriog. 74. 1936. - About the Grand Canyon, Coco-
nino (probably not Mohave) County, Arizona, at
7.000 feet elevation, 28 June 1898, MacDougal 176.
Holotype, LY! Isotype, GH, NY, UC, US!

Eriogonum spathulare Gand., Bull. Soc. Roy.
Bot. Belgique 42:191. 1906. — Eriogonum microthe-
cum Nutt. var. spathulare (Gand.) S. Stokes. Gen.
Eriog. 74. 1936. - On sterile white clay hillsides,
Maurey's Mountain, Crook County, Oregon, 25 July
1901, Cusick 2698. Holotype, LY! Isotypes, GH, K,
NY, ORE, POM, UC, US!

Eriogonum intricalum Gand., Bull. Soc. Roy.
Bot. Belgique 42:191. 1906, non Benth., 1844.'-
Near Townsend, Broadwater County, Montana. 13

August 1899, Blankingship s.n. Holotype, LY! Iso-
types, MONT, NY!

Eriogonum tenellum Torr. var. grandiflorum
Gand., Bull. Soc. Roy. Bot. Belgique 42:197. 1906.-
From an unknown location in Rich County, Utah,
August \W1 Jjnford s.n. Holotype, LY!

Eriogornmi tenellum Torr. var. sessiliflorum
Gand., Bull. Soc. Roy. Bot. Belgique 42: 198. 1906. -
Near Reno, Washoe County, Nevada, September
\894, Hillman s.n. Holotype, LY! Isotype, RENO!

Low to erect, spreading to sparsely branched subshrubs,
or infrequently shrubs, (1) 2-4 (5) dm high, 2-8 dm across;
leaves mostly elliptic, (0.5) 1-2 (2.5) cm long, (1.5) 2.5-6 (8)
mm wide, densely to sparsely white-tomentose below, less so
to sparsely floccose and whitish-green above, the lomentum
whitish, the margins plane or with thickened edges, infre-
quently revolute, the apices acute; flowering stems slender,
2-6 (8) cm long, tloccose to sparsely tomentose when young,
remaining at least floccose in some in the northern part of
the variant's range, otherwise frequently becoming green and
glabrous or subglabrous, the tonientum whitish; inflores-
cences (1) 2-4 (8) cm long, floccose to glabrous when young,
usually becoming subglabrous or glabrous at maturity ;oti'o/;(-
cres 2-3 (3.5) mm long, subglabrous to glabrous or merely
tloccose between the angled ridges ;/7oH'fl'S white with green-
ish- or reddish-brown midribs and bases, becoming tinged
with pink or wholly pinkish in most especially at maturity,
2-3 mm long, the tepals slightly dissimilar, the outer whorl of
tepals obovate, the bases truncate to nearly cordate, the inner
whorl of tepals mrrowei; achenes 2-3 mm long. - Represen-
tative collections: Archer 6979 (NA, RSA, V/S): Baker A'6J5.
S745 (WS), 9444. 9529. 9984 (VJTV): Beach 86 7 { ARM.
BM, COLO, IDS, NY, OKL, SD. US, UTC, WIS, US, WTU);
Christ 5813. 5824. 6551. 8491. 8894. 8964. 9029. 9799.
11452. 12238. 15514. 15540. 16104 (NY); Cottam 1599S
(RSA. UT. WIS, WTU); Cronqiiist 1961 (IDS, MO, NY),
3056 (IDS, MO, ND), 7523 (GH, NY, UC, UTC, WS, WTU),
7766 (CAS, NY, RSA, UTC, WTU); Detling 3155 (ORE,
UC); Eastwood and Howell 7035 (CAS, GH, \}C): Ferris and
Lorraine 10995 (CAS. IDS, GH, RSA, VC.WTV): Goodding
1 742 (COLO, GH, MO, NY. UC, VS):Heller 9511 (MO, NY,
RENO, US), 9979 (A, GH, RENO); C L. Hitchcock 15587
(COLO, NY, RSA, WS. WTU), 15657. 16703 (NY, RSA, UC,
WS, WTU); Hitchcock and Martin 5579 (ISC, NY, UC, WS,
WTU). 5658 (MO, NY, OKL, POM, UC, US, UTC, WS,
WTU); Leiberg 489 (BM, BR, GH, NY, OKL, UC, US);
Magiiire and Holmgren 22193 (GH, MO, NY, UC, UTC,
WTU), 26729 (GH, NY, UC, US, \}lC):Nelson 8116 (ARIZ,
GH, MO, NY, POM, RENO, US); Peck 9715 (GH, NY,
WILLU); Remy s.n. (P); Reveal and Holmgren 1905 (NTS,
NY, UTC); Roos and Roos 5931 (CAS, RSA): Sandherg and
Leiberg 383 (BM, BR, CAS, GH, MO, NY, OKL, POM, UC,
US, WS); Torrev 449 (GH, MO, NY, US); Watson 1024 (NY,
US); Welsh and Moore 5148 (BRY); Whited 85 (A, MO, ND,
NY).

Distribution.

Widespread and common from southeastern Wash-
ington southward into north-central California, other-
wise east of the Sierra Nevada as far south as extreme
northeastern San Bernardino County, California, then
eastward in the high valleys, foothills, and mountain
ranges througli central and southern Idalu), all except
southernmost Nevada, Utah, and northern Arizona
(mainly on the Kaibab Plateau) into western Mon-

18

BKI(;HAM YOUNC. UNIVERSITY SCIENCE BULLETIN

tana, and west of the Continental Divide in western liabuats. fioni (1.500) 5.000-10.500 feet elevation.
Wyoming and Colorado, found in a wide variety of Howering from June to October. Figures 6 and 7.

l-'ig. 6. Disirilnilion map oi l-jiugoimiii microlhecuin var. laxijloniiii.

BIOLOGICAL SERIES. VOL. 1 3, NO. I NOTES ON ERIOGONUM - VI

19

Fig. 7. Illustration oi Eriogoinim microthecum var. laxijloiinn showing variants from throughout the taxon's range.

20

HKU.HAM YOUNC. UNIVKKSl lY SCIKNCF HULLI-.TIN

The history of the discovery and naming of var.
laxiflonim is long and complex. Thomas Nuttal! trav-
eled westward with Nalhaniel J. Wyeth, a Bostt)n ad-
venturer, in 1834. Accompanying Niitlall was John
K. Townsend, an ornithologist who kept an excellent
journal (Townsend, 1839) which was used by McKel-
vey (1955), Graustein (1967) and Reveal and Hafen
(1970) to trace Nuttall's round-about route from .St.
Louis, Missouri, to Fort Vancouver near the mouth of
the Columbia River. It has been possible to essentially
pin-point the type location of var. mkrothecum
(which see), but the exact type locality of var. laxiflo-
nim remains a mystery. The label datum on the col-
lection at the British Museum (Natural History) is as
cited above, but the exact intended meaning of the
decriptive phase "Rocky Mountains of the Columbia
in Oregon" is difficult to understand. Comparing Nut-
tall's original collections with others made along the
Oregon Trail where he traveled, it would seem that
the type came from Idaho or, and less likely, from
adjacent eastern Oregon. The leaves are plane (that is,
not revolute) and densely tomentose below; the intlo-
rescences are open, glabrous, and the white (lowers
have outer tepals with cordate bases. Nuttall made
several collections of this variety while in the Oregon
Country, no doubt dming the different years he was
in the area ( 1834 and 1835), and assigned a series of
herbarimn names to each. None has exact location
data and thus their exact origin can only be prcsmiip-
tive.

The name, var. laxijloriim. was first proposed by
Hooker (1853) although three years later, Bentham
proposed the same name, based on the same type.
Both men cited Ehogoiniw kixijlorum Nutt. in syn-
onymy, but this herbarium name actually was never
published by Nuttall himself. In as much as Hooker
was referring to the Nuttall name, the type is the
Nuttall specimen in Hooker's Herbarium now depos-
ited at Kew even though he mentioned a Geyer col-
lection. The fact that Hooker was the first author of
this name was not noted until recently (Reveal and
Munz, 1968). However, it is difficult to explain
Bentham's actions in redescribing the variety alter his
friend and colleague, Hooker, did so. I believe that
Bentham actually suggested the name to Hooker and
the latter included it in his article on Geyer's plant
collection without giving Bentham credit for the
name. Later, Bentham knowing he was responsible
for the name, simply credited himself. However, this
is speculation which is not admissible and Ihus, we
must accept Hooker as the author of var. laxifloniiu
even though Bentham may have been the original
authority.

Over the years, var. laxifloniiu has been lolally
misunderstood. Stokes (1936) recognized which ele-
ment represented the type of Eriogimum microthe-
cum (that is, the \ar. micnilheciim), but she failed to
apply the information to the taxonomy of the species

even as she understood il. and thus hci name, ssp.
typiciim and the concept of the taxon which went
with it, was applied to what is here called var. laxiflo-
niiiK In all lloras and manuals up to 1964 the concept
of typical microtlicciim was associated with var. laxi-
Jlonim, and var. idalioense was considered the valid
name for what is now referred to as var. inicroihe-
nim. The first author to reverse this oversight was C.
Leo Hitchcock (1964) who settled the problem at
least for the Pacific Northwest flora. Thus, in most
lloras, the distribution and description of the species
is based on a variety of the species (var. laxiflonim)
aiul nut on the typical form at all.

It IS herein propcised ihal all subsequent varieties
arose from this phase of ilie species. Four rather dis-
tinct lines evolved independently from var. laxiflonim
and the linear arrangement is simply for convenience
as one could have started with any one of the other
lines and proceeded to the others. The \-dr. foliosum
is the closest of the extant variants and is treated
next. It is ditTicidt to separate the two varieties at
times and thus they seem to be still in the process of
evolving into two discrete entities. The next line is
that which extends from var. paiiainiiitense to var.
jolmstoiiii via var. corvmbosoiUes. This group is re-
stricted to the mountains of southern California, and,
although somewhat isolated from each other geo-
graphically, they are still very similar in some basic
morphological characteristics. The vars. lapidicola and
alpiiuim are two extremes, both highly reduced. The
first is at lower elevations in the deserts from eastern
California to (perhaps) western Utah, while the sec-
ond is more restricted, being found on the Sierra
Nevada of east-central California. The last line of evo-
lution is that formd in vars. microilieciim and ambig-
iiiim. Botli have yellow fiowers and are difficult to
distinguish in the herbariimi. although in the field the
two are distinctive enough to merit formal recogni-
tion. The presence of the yellow fiower color in
otherwise white tlowered species is relatively frequent
in the genus, but the taxonomic significance of such a
distinction can only be determined on the merits of
each individual case. Here, the two yellow flowered
entities have oilier nwrphological differences and
seemingly occupy, at least in part, a unique geograph-
ical range, and thus, they are given formal status.

As now defined, the var. laxiflonim is still variable
and contains many micropopulations that may or
may not be worthy to taxonomic recognition in the
future (see below). What is considered to have
evolved from tins eiiliiy and consequently described
as taxonomic units at this time are those phases of
the overall species that represent major lines or modes
of development. Some of the additional elements
within var. laxiflonim and most likely \m. foli(>siim.
may still be distinguislied in time.

Certain individual populations denn)nstrate the
taxonomic dilTiculties associated with var. laxiflonim.

BIOLOGICAL SERIES. VOL. I .1. NO. 1 NOTES ON ERIOGONUM - VI

21

In soullieiii Wasliington, tlie plants arc readily distin-
guished, but as one proceeds into Oregon, and espe-
cially eastern Oregon, the features of the broad, plane
leaves, open inflorescences and tall stature give way
to the lower, more scraggly forms so typical of the
plants found in the Great Basin of Nevada and Utah.
However, in north-central and central Oregon, the
robust feature persists and the yellow flowered vari-
ant, var. micwthecum. becomes common. In parts of
the John Day Valley, both variants occur, but var.
lii.xiflonim is very much in the minority. It is likely
the two merge in various parts of their range in Ore-
gon as they certainly do in Lassen County, California.

The Idaho plants fall well within the morphologi-
cal delimitations of var. laxijlonini. In this area, and
especially in the mountains of central Idaho, the invo-
lucres are often long peduncled and the flowering
stems thinl>' floccose. The leaves are narrowly oblanc-
eolate and nearly glabrous above, but on the Snake
River Plains, the leaves are wider and more densely
pubescent although the stature of the plants are not
too greatly reduced except when the plants occur on
harsh edaphic sites.

On the whole, the plants in Montana of this vari-
ety are small, being rarely more than 2.5 dm high.
Correspondingly, the leaves are small, narrow and
more densely pubescent (especially above) and thus
similar to those individuals found in the Great Basin.
In the mountainous part of the state, the plants tend
to be similar to those in adjacent Idaho, as would be
expected.

In the Intermountain West the var. lii.\ijli)riiiu
occupies sites (as in southwestern Wyoming) that are
of a similar nature. The plants tend to be reduced in
stature, more spreading, woody, and less leafy. Like-
wise the density of the pubescence increases. How-
ever, these features are associated with two factors:
one, the southward extension of the variety, and sec-
ond, the upward advance of populations onto tops of
mountain ranges often well above ^),000 feet eleva-
tion. The compaction of forms here is unlike that
found in eastern Utah where forms of var. laxijlonim
(and var. foliosum) occur on clay slopes. In the Great
Basin, the plants spread outwardly from a gnarled
root crown, with elongated caudex branches armed
with oblanceolate leaf-blades and small but open
inflorescences. Those plants at the lower elevations
tend to be more pubescent than those at higlier eleva-
tions. In eastern Utah and adjacent Colorado the
plants are pulvinate, and Reveal 683 from Emery
County, Utah, and Reveal and Davidse 856 from Rio
Blanco County, Colorado, are so reduced they tend
to resemble Eriogomim eontortwn Small ex Rydb.
These plants would tend to fall into var. foliosum and
may be better placed there along with other reduced
forms of var. foliosum in eastern Utah and adjacent
western Colorado. However, these elements do not fit
within any established taxonomic group in the species

as now defined, and perhaps they should be given
formal recognition. The elevational gradients in Utah
are not as pronounced as it is in Nevada except on the
Aquarius Plateau and Henry Mountains where plants
also lend to be more pulvinate than spreading as is
the case on the Great Basin ranges of central and east-
ern Nevada.

In the field, var. laxijlorum occupies a multitude
of habitats and comes, therefore, in a wide variety of
shapes and forms. In addition to the general aspect of
the plants as noted above, the variety ditTers in vari-
ous ecological sites. For example, when the plants
occur in scattered stands of Artemisia (sagebrush) as
in southern Idaho, parts of Oregon and Nevada, the
plants are often large, rounded, and rather robust at
lower elevations. When plants occur on steep road
banks, and especially road cuts, the plants are often
extra large. Yet, as one proceeds to higher elevations,
the plants become more prostrate, with their long
stems spreading along the ground usually arising from
root crowns situated in open places between individ-
ual plants of Artemisia. In protected areas at these
higher elevations, the plants may be more leafy, but
they are still prostrate and appear depauperate.

The ability to consistently determine var. laxiflo-
rum horn var. foliosum is impaired by a band of
intermediate populations extending nearly the entire
length of their contact zone in southern Nevada and
Utah, and northern Arizona. For the most part the
two may be readily and simply determined, but in
some areas, the characteristics completely break
down. For example, on the foothills of the Toquima
and Monitor ranges in central Nevada, these two vari-
ants come together and cannot be distinguished; the
same is true in the Grand Canyon area of northern
Arizona. Variation may be seen in a single series of
collections made by a single collector, no doubt to
show the variation he has seen in the field. Unfortu-
nately, these collections are broken up into herbar-
ium sheets which are then distributed to herbaria far
and wide. Once reassembled, the variation is difficult
to fully understand as the growth liabit of the plant is
gone, the aspect of the ecological niche is not pre-
served, and it is impossible to know exactly what all
the phases originally represented. This is especially
true of the large collections of Ira W. Clokey's from
the Spring (Charleston) Mountains of Clark County,
Nevada. In this area both elements are present, and
the entire sample seems to represent both var. folio-
sum and plants which approach (but never really
reach) var. laxiflorum.

There is no solid line of demarcation between
these two varieties, and as the taxonomic rank of
"variety" is used here, this zone of morphological
overlap is to be expected. If the differentiations were
more consistent and rigid, one would be compelled to
use a higher taxonomic rank. Likewise, as the two
entities are distinct throughout much of their respec-

22

BKIC.HAM YOUNG UNIVKKSll Y SCIKNCE BULLETIN

live ranges, it winild be equally unrealistic to reduce
these to a single taxon.

6b. Eriogoniim niicrothecum var. foliosum (Torr. &
Gray) Reveal

Eriogoiiuui niicrothecuni Nutt. VM.foUosunt (Torr. &
Gray) Reveal, comb, nov., based on E. effiisitm Nutt.
var. foliosum Torr. & Gray, Rapt. Expior. Surv.
Ascert. Pract. Econ. Route Railroad Miss. River to
Pacific Ocean 2:129. 1857. - San Luis Valley, Ala-
mosa or Saguache counties, Colorado, July 1853,
Cruetzfeldt s.n. Holotype, NY! Isotype, GH!

Eriogomim simpsonii Benth. in DC, Prodr. 14:18.
1856. - Eriogonum effiisuni Nutt. ssp. simpsonii
(Benth. in DC.) S. Stokes, Gen. Eriog. 81. 1936. -
Sierra de Tunecha, northwestern New Mexico, Sep-
tember 1849, Simpson s.n. Holotype, NY!

Eriogonum mircothecum Nutt. var. rigiclum
Eastw., Zoe 4:11. 1893. - Eriogonum microlhevum
Nutt. ssp. rigiclum (Eastw.) S. Stokes, Gen. Eriog. 75.
1936. — On mesas near Diuango, La Plata County.
Colorado, August 1892, Eastwood s.n. Holotype,
CAS! Isotypes, GH. MO, UC, US!

Eriogonum frisainum M. E. Jones, Contr. W. Bot.
11:14. 1903. — Eriogonum microthecwn Nutl. var.
friscanum (M. E. Jones) S. Stokes, Gen. Eriog. 74.
1936. - Frisco, Beaver County, Utah, 24 June 1880.
M. E. Jones s.n. Lectotype, POM!

Eriogonum nelsonii L. Will., Bull. Torrey Bot.
Club 59:428. 1932. — Eriogonum effusum Nutt. ssp.
neLionii (L. Will.) S. Stokes, Gen. Eriog. 81. 1936. -
Geyser Basin, San Juan County, Utah, 30 July 1912,
Walker 368. Holotype, RM!

Eriogonum microtliecum Nutt. ssp. intermedium
S. Stokes, Gen. Eriog. 75. 1936. - Rocky slopes near
Ely, White Pine County, Nevada, 24 August 1931,/
T. Howell 795f). Holo'type. CAS! Isotypes, GH, US!

Low and spreading to tail and creel sulishriibs and shrulis
(I) 4-15 dm liigli and (1) 4-16 dm across; leaves mostly nar-
rowly elliptic, 0.5-1. « (2.5) cm long, (0.5) 1-2 (2.5) mm
wide, densely white-tomentose below, lloccose and whitish-
green above, rarely snbglabrons or glabrous and green above,
the tonientum white, tightly revolute in most or at least with
rolled, thickened margins, the apkcs Mute; Jluwerini; sleius
slender to ± stout, 2-7 cm long, densely lanale to tonientose
throughout, rarely tloccose at maturity, or if subglabrous and
greenish, then in the southeastern part of the variety's range,
the tomentum \vhH\^h; inflorescences (1.5) 2-4 (6) cm long,
tonientose to tloccose. rarely suliglabrous at maturity in New
Mexico and northeastern Arizona; involucres 2-3 mm long,
tonientose to tloccose or suliglalirous, the tomentum usually
dense between the angled ridges ;y/ovt'en' white with greenisli-
to reddish-brown midribs and bases, becoming pinkish in
fruit in some, 2-3 mm long, the tepals essentially similar to
slightly dissimilar, the outer whorl of tepals narrowly obovale
to obovate, the bases rounded to truncate or t truncate-cord-
ate to cordate; ffc/;c/icj' 2-3 mm long. - Representative collec-
tions: Appleqiiisl s.n. iMNA):. Arsene and Benedict l()6l() (P,
US); Beal 562 (ARIZ, \\\\ WW): Clokev and Clokev 7071
(ARIZ, BRY, CAS, US, Gil, MO, ND, NY, OKL, RM, UC,
US, UTC, \VS, WTU); Culler 2783 (GH, MNA. NY, WIS);
Deaver 3765 (ARIZ, MNA); Eastwood and Howell 6531.

6555. 6986 (CAS, US), 7316 (CAS. GH, POM), 7332 (CAS,
GH, NY, US); Flowers 6377 (UT); G'a/wav 8243 (15RY, US);
Goodman and Parson 3260 (UC, WTU); Howell and True
44714. 44835. 44878 (CAS): Jones J 795 (BM, BR, CAS, G,
GH, NY, POM, US, UTC); Kearnev and Peebles 12820
(ARIZ, NY, US); Magidre 17659 (GH, NY, UlC); Munz
12855 (A, POM, UC); Parson 613 (COLO, GH, RM, WTU);
Popenoe sn. (A, KSC): Purpus 6280 (UC, US). 6296 (CAS,
NY, VOUy.Ramalev 14370. 15241. 15817 (COLO): Reveal
683 (ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, OKL, RM.
RSA, UC, US, UT, UTC); Reveal and Beattev 1691 (BRY,
NTS, NY, UTC); Reveal and Davidse 933 (BRY, CAS, GH,
LL, NY, OKL, RSA, SMU. TUX, UC, UTC); Reveal and
Holmgren 1813 (BRY, N IS, NY, VTC): Riplev and Harnehv
4005. 8684 (CAS); Rollins 1532 (GH, NY), 1934 (NU. NY,
WTU); Rusbv 815 (CAS, NY, US); Rrdhcrg and Garrett
8442 (GH, NY. WIS); Weber 3868. 7827 (COLO): Welsh and
Moore 1838 (HRY. ISC, WIS), 1993. 2221. 2348 (BRY, NY);
Wetherill s.n. (MNA); Wooton s.n. (US).

Distribution.

Widespread and common from southeastern Cali-
fornia in eastern San Bernardino and Inyo counties
eastward across southern and central Nevada into
northern Arizona and southern Utah, northward on
the Colorado Plateau to Emery and Grand counties,
and in widely scattered locations as far north as the
Wyoiriing state line, entering western Colorado in
Mesa County and proceeding southward and across
southern Colorado to San Luis Valley and across
northern New Mexico to the Sangre de Cristo Moun-
tains, hence southward to central New Mexico, in a
wide variety of ecological niches and communities,
from 4,500-7.500 feet elevation. Flowering from
June to October. Figures 8 and 9.

The type of var. foliosum was discovered by Fred-
erick Creuzefeldt in San Luis Valley of south-central
Colorado while with the Gunnison Expedition in
1853. The entity was described by Torrey and Gray
in 1857. However, the variety was first found by
Lieutenant James H. Simpson while on a Navajo raid
into northwestern New Mexico lead by Lieutenant
Colonel John M. Washington, the military governor of
New Mexico (Goetzman, 1959). The Simpson collec-
tion, made in 1849, consisted of a small stem which
was used by Bentham ( 1856) as the basis for his new
species. As can be seen in the above list of synonyms,
several specimens representing var. foliosum were
described. Eastwood (1893) described the var. rigi-
diim from material obtained only a few miles west of
the type locality of var. foliosum, but then it was
only a short distance north of the type area of Erio-
gonum simpsonii] Jones (1903) added E. friscanum
to the list by naming this species from western Utah,
but L. O. Williams (1932) brought the two "centers"
of type distribution together by describing /■.'. iwlsonii
from southeastern Utah.

The treatment by Stokes (1936) of what I have
called var. foliosum requires close examination. She
ni;maged to recognize all of the above synonyms, ex-
cept var. foliosum, under Eriogonum microtliecum or

BIOLOGICAL SERIES. VOL. 1.1. NO. 1 NOTES ON ERIOGONUM - VI

23

E. effusion. It lias been Impossible to discover the
combinatitm of characters she used to place the vari-
ous tbrnis into one or the other of these species.
Under E. microlhecuin she placed var. rigidiiiu (and
included var. follosum as a synonym, in part - but
there is no indication where the other "part" should
have been applied). E. friscanum, and described ssp.
intermedium - one each tVom Colorado. Utah, and
Nevada respectively. Under E. effusum she placed E.
simpsonii and E. nelsonii - one each from New Mex-
ico and Utah.

The var. folioswn is now defined to include those
populations of Eriogonum microthecum from the
southern part of the species' range. It is easily recog-

nized by the lightly revolute leaves with densely Ian-
ate to tomentose stems and inflorescences. Still var.
folidsum is exceedingly variable even as now defined.
The largest forms of the species are found within this
variety as some shrubs are over a meter high in north-
ern New Mexico, northeastern Arizona, and southern
Nevada; yet, within this same variety, 1 have been
forced to place some highly reduced populations
found on gumbo clay hills in eastern Utah and central
Nevada. These latter populations are thought to rep-
resent an extreme in the variation, and except for the
isolated plants in central Nevada which are provision-
ally placed here, all of these depauperate entities can
be referred to the var. foliosum in the strictest sense.

k • • • • •

Fig. 8. Distribution map of Eriogonum microthecum var. foliosum. Stars refer to populations intermediate between var. laxi-
florum and var. foliosum.

24

BKIGHAM YOUNC; UNIVKKSIIY SCIENCE BULLETIN

M

"^^^k^ , ^"^y^

$ y /

T "5V-' ■'x''^ '.''■i %<3^''H'^i:V~J.v ^y'

\

Fig. 9. Illustration of Eriogomim inurolhccum \m. jolUtsum showing varianis Ironi thiouahoiu Ihc t;i\on\ r;im;i.' ami botli
the large and small extremes in height.

BIOLOGICAL -SERIES. VOL. 1.1, NO. I NOTES ON ERlOCiONUM - VI

25

The variation in tiie amount of puLiescenco may be
worthy of more investigation. In tiie tali erect plants,
the stems may be very densely tomentose to lanate or
glabrous. Those of the first group are common
throughout the southern part of the variety's range,
extending from southeastern California to New Mex-
ico. However, the glabrous (or nearly so) plants are
found in northwestern New Mexico and adjacent
northeastern .Arizona. Again, at the extremes, there
does seem some value in attempting to distinguish
between them, but as numerous intermediates are
seen, the value of such a taxonomic decision becomes
less obvious, and thus no new entities are proposed at
this time.

As noted under var. kixitlonini. it and the present
variety are often difficult to separate where their
ranges overlap.

6c. Eriogonum microthecum var. panamintense S.
Stokes

Eriogonum nilcrotlicciini Nutt. var. panaiuinteiise S.
Stokes. Gen. Eriog. 74. 1936. - Wild Rose Canyon.
Panamint Range, Inyo County, California, at 7,800
feet elevation, 29 September 1931, Hiiffmaiiu s.ii.
Holotype, CAS.

Eriogonum effusum Nutt. var. limhatum S. Stokes,
Leaf!. W. Bot. 3:15. 1941. - Pinon Mesa, Panamint
Range, Inyo County, California, at 6.200 feet eleva-
tion. 28 September 1939, Oilman 3954. Holotype,
CAS.

Large, rounded to flat-topped slirubs 3-6 dm liigti and (4)
5-12 (IS) dm across; leaves mostly broadly elliptical, 0.6-1.8
cm long, 3-8 mm wide, tomentose below, tloccose to snbglab-
rous above, the tomentum whitish-brown to brown, the mar-
gins plane, not revolute, the apices acute to obtuse;/7ovv(T»;^p
stems slender, 5-10 (15) cm long, tloccose, the tomentum
reddish-brown in most; involucres 2-2.5 mm long, subglab-
rous to glabrous without: /7t>ivw.? whitish-brown with large
reddish-brown midribs and bases, becoming reddish-brown in
fruit, 1.5-2 (2.5) mm long, the tepals dissimilar, the outer
whorl of tepals obovate, the bases truncate to cordate, the
inner whorl nmowei'.achenes 1,8-2.1 mm long. - Represent-
ative collections: Gilman 2023 (US), 27U0 (CAS), 2701
{VOW: Hoffmann 431 {CAS); Reveal and Holmgren 1779
(NTS. NY, VTO.Roos 7J(P0M).

Distribution.

Restricted to the Panamint Range and the Inyo
Mountains, Inyo County, California, in gravelly to
rocky soils on slopes and steep hillsides in sagebrush
scrub and pinyon-juniper woodlands, from
6,000-9,000 feet elevation. Flowering from July to
October. Figures 10 and 1 1 .

The type of var. panamintense was collected by
Ralph Hoffmann in 1931, and named by Stokes as a
variety of Eriogonum microthecum in 1936. In 1941
she redescribed the same plant as a variety under E.
effusum.

The affinity of vars. panamintense and iaxiflorum
cannot be denied. The two are very similar, ditfering

Fig. 10. Distribution map of Eriogonum microtliecum var.
panamintense. Inyo Co., California.

in the color of the tomentum, the broader leaves (as
compared with the var. Iaxiflorum in the same area),
and the brownish tinge to the flower color. As an iso-
lated population, adaptive radiation has likely played
a major role in the evolution of var. panamintense,
but the degree of isolation is not as strong as it is in
the following two variants.

In the field var. panamintense is easily distin-
guished by the large brownish shrubs which occupy
the lower elevations on the west side of the Panamint
Range and the Inyo Mountains. At the higlier eleva-
tions, the plants tend to be shorter and more com-
pact, but still the plants are relatively large when
compared with the low scraggly forms of var. Iaxiflo-
rum at these same higher elevations. Of all the plants
in this species, this variety and the next would make
fine additions to the garden.

6d. Eriogonum microthecum var. corymbosoides
Reveal

Eriogonum microthecum Nutt. var. corymbosoides
Reveal, var. nov. — Johnston Grade, 0.8 miles below
the summit northeast of Baldwin Lake along Califor-
nia Highway 18, in granitic soils among pinyon and
mountain mahogony, San Bernardino Mountains, San
Bernardino County, California, 10 September 1968,
Reveal 2090. Holotype, US! Isotypes, 30 duplicates
to be distributed to various herbaria from the United
States National Museum. - A var. panamintensi foliis
longioribus, (0.8) 1-2 (2.5) cm longis, caulibus lanatis,
involucris 2-3 mm longis, floccosis, fioribus 2-2,5 (3)
mm longis, acheniis 2.5-3 mm longis differt: a var.
laxiflora tomentoso ferrugineo et a var. johnstonii
statura maiore differt.

Large rounded to spreading shrubs 3-6 dm high and 6-12
(15) dm across; leaves elliptic to obovate, the leaf-blades
(0.8) 1-2 (2.5) cm long, (4) 6-10 mm wide, densely tomen-
tose below, floccose to subglabrous above, the tomentum
whitish-brown, the margins plane or merely rolled, not revo-
lute, the apices acute to oh\.uss. flowering stems slender to ±
stout, 5-13 (15) cm long, lanate to tomentose, rarely thinly
tloccose at maturity, the tomentum tannish- to reddish-
brown: inflorescences densely cymose, 1-4 cm long, lanate to
tomentose, the tomentum tannish- to rcddisli-hrown: //;ro/»-

26

liRIGHAM YOUNG UNIVEKSI lY SCIENCK BULLETIN

cres 2-3 mm limg. Iloccosc when yoiing. bfcoming levs pubes-
cent to •iiibglubrous at maturity in some; flowers whitish-
brown with large reddish-brown midribs and bases, becoming
reddish-brown in fruit, 2-2.5 mm long, the tepals essentially
similar to slightly dissimilar, the outer whorl of tepals merely
rounded at the base in most; aclieiies 2.5-3 mm long. Rep-
resentative collections: Balls 20193 (BM, CAS. RSA, UT);
Ewan H332 (GH. NO, VC): Johnston s.n. (POM); Knox s.n.
(UT);A/»/;z 7661 IGII. NY, VC). 10784 (POM. Vn. 12706
(A, HM, MO, POM, VO-.Pfirson 4003. 51.^0 (KSAI; Whcchr
I2SS['H\). R.SA.WIU).

Distribution.

Loose gravelly to rocky granitic or limestone soils
in the San Bernardino Mountains, San Bernardino
County, from 5,800-4,500 feet elevation, and t)n the
north slope of the San Gabriel Mountains, Los
Angeles County, from 7,000-7,500 feet elevation,
southern California. Flowering from July to Septem-
ber. Figures 12 and 13.

The name "corymbosoides" is derived the Greek
koryiuhos. corymbose, and coides, likeness, alluding
to the similarities between the new variety and lirio-
gonum coryinbositin Beiith. in DC.

Ehogonum microtliecuni has long been known
from the mountains of southern California, but iniiil

:&::.

I ig. 1 1. Illustration o( L'riogoniim microlliecuni \.\i. piimtniinlcnse.

BIOLOGICAL SERIES. VOL. 1 J, NO. 1 NOTES ON ERIOGON UM - VI

21

t...

- — icl 1? )

Fig. 12. Distribution map of Eriogonum inicrorheciim var
corymhosiodes. San Bernadino and Los .'Vngcles counties
California.

Fig. 13. Illustration of Erfi>f;i)iniiii microrlieciini var. cnrynibosoides.

28

liRKlHAM YOUNG UNIVKRSI lY SCIENCE BULLETIN

now, the various elements restricted to this area have
not been distinguished. The var. corymbosoides is
well isolated From var. panaminteiise which is about
135 miles northward, as well as from the var. /aA^y/o-
nim which just enters the northeastern corner of San
Bernardino County in the Kingston Mountains. How-
ever, within the immediate area of var. corym-
bosoides is the var. plinstonli. another variety but
restricted to the higher ridges in the San Gabriel
Mountains.

The var. corymbosoides resembles many specimens
of Eriogomim corymhosum var. corymliosum, an
entity primarily of Utah and Colorado. The broad
leaves are distinctive as is the dense, essentially lanate,
tomeiitum on the stems and inflorescences. In its
color, the tomentum approaches that found in var.
corymliosum from eastern Utah and the southern
populations of £. corymbosum vdt. glii I inosiim (M. E.
Jones) M. E. Jones. In the denseness of the tomen-
tum, the new variety is approached only by the var.
foliosiim, but the color is different, and the habit of
these two forms o( E. microiheciim is most distinct.
In the field, specimens of var. corymbosoides vary in
the degree of the tomentum, much as in other phases
oi E. microtheciim. The plants normally in the shade
of pinyon and mountain mahogony are less densely
pubescent than those plants found on flats among
sagebrush. In the area around Cactus Flat, a complete
trend may be seen from the sandy flat to the adjacent
slopes with regards to this feature of the tomentum.

Some specimens from the Sugarloaf region of the
San Bernardino Mtnmtains approach var. jolmstonii
(Balls 20193). but in other high elevation places, the
plants clearly are var. corymbosoides.

Three collections require special comment. The
Parish Brothers, M. E. Jones, and Anstruther David-
son collected an odd form of Eriogonum microthe-
ciim in Bear Valley of the San Bernardino Mountains:
Parish and Parish 1512 in August of 1882 (BM, US),
Jones s.n. on 19 July 1900 (POM), and Davidson
2284 in July of an unrecorded year (US). These speci-
mens are totally glabrous except for the lower leaf
surface, although the upper surface of the leaves and
the stem among the leaves may be thinly floccose at
times. The leaves are oblanceolate, 1-1.5 cm long and
3-5 mm wide with revolute margins. The involucres
are 3-4.5 mm long and 2-2.5 mm wide. The plants are
less than 2 dm tall and appear to be very distinct.
Unfortunately, additional specimens of this form
have not been discovered in recent years, and imtil
this entity can be more thoroughly studied, its proper
placement in the species is impossible. Dr. Philip A.
Munz recently visited this pi)pular resort area, and
reports the area is so built up that it may be difficult
to ever find the plants again.

6e. Eriogonum niicrothecum var. johnstonii Reveal
Eriogonum microiliecum Null, var.johnsionii Reveal,

var. nov. - West spur of Mt. San Antonio [then, Old
Baldyl , San Gabriel Mountains, on the ridge in loose
broken granite soil at 9,000 feet elevatitm, Los
Angeles County, California, 16 September 19|7,
Johnston 1726. Holotype, UC! Isotypes. GH, POM!
— A var. corymbosoides foliis minoribus, 5-10 mm
longis et 3-5 mm latis, caulibus floccosis vel subglab-
ris, involucris (2) 2.5-3 mm longis. floccosis vel glab-
ris, florlbus (2.5) 3-3.5 (4) mm longis dilTert.

Low decumbent spreading siil)shriilis 0.6-1.3 dm liigli and
2-5 dm across; leaves elliptic to ovate, the leat'-blades 5-10
mm long, (2) 3-5 (6) mm wide, densely lomentose below,
tloccose to subglabrous above, the tomentum whitish-brown,
the margins plane or merely rolled, not revolute, the apices
acute; y7oH'tTO;jif stems slender, 3-6 cm long, tomentose when
>oung or more commonly tloccose to subglabrous especially
at maturity, the tomentum whitish- to reddish-brown; inflo-
rescences cymose, 0.5-3 cm long, tloccose to subglabrous, the
tomentum whitish- to reddish-brown; involucres (2) 2.5-3
mm long, tloccose when young, becoming glabrous at matu-
rity;//oive/'S whitish-brown with large reddish-lirown midribs
and bases, becoming reddish-brown in fruit, (2.5) 3-3.5 (4)
mm long, the tepals essentially similar to slightly dissimilar,
the outer whorl of tepals merely rounded at the base in most;
achenes 2.5-3 mm long. - Representative collections: John-
ston 1530 (GH, POM. \}C)\Mwu 16S7, 6U9iS I.POM):Peirson
JI99tRSA).

Distribution.

Loose granitic soil in the San Gabriel Mountains
near Cucamonga Peak and Mt. San Antonio (Old
Baldy), San Bernardino and Los Angeles counties,
California, from 8,500-9.500 feet elevation. Flower-
ing from July to September. Figures 14 and 1 5.

The name is selected to honor Ivan Miura\' John-
ston (1898-1960), professor of botany at Harvard,
the authority on Boraginaceae, and a collector in
southern California during a period from the late
1910s and early 1920s. He was the first to note the
distinctiveness of this variety (in 1923), but never for-
mally proposed a n;ime for it.

The var. johnstonii is obviously related to var.
corymbosoides. The two are similar in leaf shape,
pubescence color, and flower color. They differ In the
amount and degree of the tomentum, especially on
the stems and inflorescences, and in statrn-e and dis-
tribution. The vat. johnstonii is similar to ww.lapidi-
cola In a generalized manner and points to the kind of
parallel evolution that is so commonly found in this
large and complex genus of flowering plants.
6f. Eriogonum microtheciim var. lapidicola Reveal
Eriogonum microiliecum UuU.Vdv. lapidicola Reveal,
var. nov. — North end of Rainier Mesa at the south-
ern end of the Belted Range at the head of The Aque-
duct, 0.5 mile north of Rainier Mesa Road near the
jimction of Old Rainier (or Back) Mesa Road on shal-
low soil assiiciated with sagebrush and pinyon-jimiper
woodlands on flat-rock outcrops of a volcanic origin.
Nevada Test Site, Nye County, Nevada, at 7.400 feel
elevation, 25 August 1968, Reveal and Holmgren

BIOLOGICAL SERIES. VOL. \ ^. NO. 1 NOTES ON EKIOGONUM - VI

nistnbulion.

Rocky slopes and Hats in thin shallow soils often
on rocky outcrops or on sandstone ledges, from the
Inyo Mountains, Inyo County, California, eastward
into Esmeralda County, Nevada, on Magruder Moun-
tain, and in the low mountains of central Nye and
western Lincoln counties, Nevada, and (perhaps)
westward into extreme western Utah, from
(•1.000-8.500 feet elevation. Flowering from July to
September. Figures 16 and 17.

The name lapidicola is derived from the Latin
lapis, rock, and -cola, dweller, as to the rocky habitat
where this variety occurs on the Nevada Test Site, the
type location.

The typical phase of var. lapidicola is a densely
branched form, low and spreading, and generally red-
dish-brown in color. Of the various varieties investi-
gated so far. this is the only polygamo-dioecious one
noted althougli I have reasons to suspect this condi-
tion may occur sporadically elsewhere in the species.
On the Nevada Test Site, where the variety has been
extensively studied, the plants are confined to thin
soils usually on top of tlat volcanic rocks which make
up the mesas. The rooting system is superficial, being
less than a centimeter or two below the surface, but
covering a wide area. This is similar to the rooting
system of Astragalus beatleyae Barneby, a recently
described local endemic which also grows in the same

1 ig. 14. Distribution map of Enogoiiuiu microthecuin var.
johimonii. San Bernadino and Los Angeles counties, Cali-
fornia.

1926. Holotype, UTC! Isotypes, 35 duplicated will be
distributed to various herbaria from the Intermoun-
tain Flerbarium. L'tah State University, Logan. - A
var. paiuimiiitensi statura niinore differt et similis a
var. jolmstonii sed foliis 3-7 mm longis et 1-4 mm
latis. floribus (1 .5) 2-3 mm longis.

Low dwarfed polygamo-dioecious (at least in some) sub-
shrubs 0.5-1.5 dm high and 0.8-2.5 dm across; /eai'es elliptic,
the leaf-blades 3-7 mm long. 1-4 mm wide, densely tomen-
tose below, tomentose to floccose or glabrous above, the
tomentum reddish-brown, the margins plane or merely rolled,
not revohite, the apices acute to obtuse;/7ou'(.TO!^ steins slen-
der. 2-6 cm long, tomentose when young, becoming tloccose
at maturity, rarely glabrous, the tomentum reddish-brown;
inflorescences cymose, 2-6 cm long, tomentose to floccose,
the tomentum mostly reddish-tiro wn; involucres (2.5) 3-3.5
mm long, floccose to subglabrous; flowers whitish-red with
red midribs and bases, becoming pink to rose or even orange
in fruit. (1.5) 2-3 mm long, the tepals slightly dissimilar, the
outer whorl of tepals subcordate at the base; achenes 2.5-3
mm long. - Representative collections: Alexander and Kel-
logg 3060 (OKL, VO.Beatlev 599. 604. :.i26. 390S. 4S08
(NTS). 3IO.i (DS, NTS, RSA), 50J2 (MARY. NTS): Jaeger
s.n. (POM): Reveal 1528. 1717. 1729, 1945. 2022, 2047
(BRY, NIS, NY, UTC); Reveal and Holmgren 1819 (BRY,
NTS, NY, UTC); Wells s.n. (CAS).

>jM y

Fig. 15. Illustration of Friogonum microthecum \ax. johnstonii.

30

BKIGHAM YOUNG UNIVKRSI lY SCIENCE BULLETIN

type of habitat on the Test Site. Only one collection
(Reveal 1972) has been found on sandstone. The
flower color changes from a whitish tinge to a pink or
deep rose or even an orange color in fruit, giving tlie
plants a unique color not seen elsewhere ni the spe-
cies.

In general the var. lupliliaila grows with sagebrush
under pinyon-juniper. hi eastern California the plants
grow on more exposed ridges than on the sites in
Nevada, and thus the plants tend to be somewhat less
spreading in the Inyo Mountains.

The variety is clearly related to var. kixijlcruin
although it is believed to have evolved from an ances-

toral group similar to vin. pafwiniiitense. it Is with the
latter variant that var. lupiJicola seems to merge
based on a small sample of specimens from eastern
California; however, the significance of this remains
to be studied in the field.

Two discordant elements related to, and likely
associated with, var. hipidicola can be mentioned
here. One phase is a highly depauperate population
found on sandy calcareous clay soils in Lander and
Eureka cos., Nevada {Ripley & Barneby 9330). These
plants are similar to var. foliosum populations in
northeastern Utah in size and stature, but are distinct
in a number of minor moiphological characteristics.

Fig. 16. Distritnitiim map o( L'riogoiiuni microthccum var. lapidicola.

BIOLOGICAL SERIES. VOL. I .!, NO. I NO lES ON ERIOGONUM - VI

31

^m

Fig. 17. Illuslration o( Eriogontini nucrorht'cuni vur. lapidicola.

Unfoitunately 1 liave not been able to study these
plants in the tleld, and it is possible that they might
represent another distinct taxon. The tonientum is
whitish, and thus different from var. lapidicola in this
respect, but whether or not this is a consistent feature
of the central Nevada plants cannot be determined on
the basis of a small number of collections. The second
phase is represented by Pitrpiis ^24^. This too is a
depauperate variant oi Eriogomiui fiiicrolhevum and
is questionably referred to var. lapidicola. In this very
distinct phase the leaves are glabrous above, tiglitly
revolute with thin, nearly glabrous, tlowering stems
and short, compact inflorescences. At first glance this
population reminds one of E. ehcifolium of northern
Arizona, but that species is far to the south, and the
flowers are different. One major problem with the
Purpus collection is the location-it is given as Jimiper
Mountain, a place unknown to me. However, I sus-
pect the site may be a part of the House Range in
Beaver Co.. Utah, but until this area can be visited
and the population rediscovered, the fate of the pop-
ulation as a valid entity within var. lapidicola must
await its time.

6g. Eriogonum microthecuni var. alpinum Reveal

Eriogontini microthecuni Nutt. var. alpinum Reveal.
var. nov. — Loose dry soil. Sonora Pass, Tuolumne

and Mono counties, California, at 9,300 feet eleva-
tion, 16 July 1863, Brewer 1888. Holotype, US! Iso-
types, GH, MO, UC! - A var. laxiflora statura
minore, 0.4-1 dm alta, foliis anguste ellipticis, 3-7 C?)
mm longis et 1-2.5 (3) mm latis, involucris (1.5) 2-2.5
mm longis, floribus 1.5-2.3 mm longis differt; a var.
lapidicola et var. jolmstouii foliis anguste ellipticis et
floribus 1.5-2.3 mm longis differt.

Low decumbent spreading subshrubs 0.4-1 dm high and
0.5-2.5 dm across; leaves linear-oblanceolate to narrowly
elliptic, the leaf-blades 3-7 (9) mm long, 1-2.5 (3) mm wide,
densely tomentose below, floccose to subglabrous above, the
tomentum whitish- to reddish-brown, the margins rolled to
revolute, infrequently plane, the apices acute; flowering
stems thin to slender, 1.5-4 (5) cm long, floccose to subglab-
rous, the tomentum whitish- to reddish-brown; />(y7o/'t's<;'e/!ce.s
cymose. 0.5-2 (3) cm long, floccose to subglabrous; />n'o/i;-
cres (1.5) 2-2.5 mm long, floccose when young, becoming
thinly floccose to glabrous at maturity ;//oiyf7-i' white to red-
dish- or brownish-white with reddish midribs and bases, be-
coming rose in fruit in some, 1.5-2.3 mm long, the tepals
essentially similar, the outer whorl of tepals merely rounded
at the base; at7)('/;<;'s 1.5-2 mm long. - Representative collec-
tions: Alexander and Kellogg 4020 (VVTU); Cantelow s.n.
(CAS); Eastwood 597 (CAS, GH, MO, US); Hendrix 332
(OKL); Peirson 11647 (A, CAS, COLO. RSA); Ripley and
Barneby 9909 (CAS); Roos and Roos 5985 (CAS. RSA);
Thorne and Henrickson 33116 (RSA); Wiggins 9268 (DS,
MICH, POM, UC, UTC); Wiggins and Rollins 565 (CAS, DS,
c;il. RSA, WTU).

32

URIGHAM YOUNG UNIVERSITY SCIENCT. BULLETIN

Distribution.

Dry sandy to graveii\' takis slopes und ridges In the
Sierra Nevada IVoni Alpine County south to central
Mono County and southern Tuolumne County, and
in the Sweetwater Mountains, Mono County, Calil'or-
nia, from 8,500-10,500 teet elevation. Flowering
from July to September. Figures 18 and 1''.

Fig. IS. Distribution map of Erio^oniiin iiiicrotliLCiini var.
alpliniin. Alpine. Mono, and Tuolumne counties. Calilornia.

I-'ig. 19. Illustration of Eriogonuin niicroiliccuni var. al-
piniuni.

The name alpinum is derived from the Lulin alpiii-
iis, alpine, alluding to the habitat of the new variety.

The var. alpinum was first characterized by Torrey
and Gray (1870), but they did not give the entity a
name. They drew up their discussion from Brewer's
collection and another depauperate collection from
the Ruby Mountains t)f northeastern Nevada (a form
of var. hixijhmimj. I am adopting their concept of
the variety, excltiding the Nevadan element, and
restricting the entity to the Sierra Nevada and the
closely associated Sweetwater Mountains of Califor-
nia. It is a higli alpine form growing at or above tim-
berline in the Sonora Pass region and elsewhere^on
the Sweetwater Mountains and in Alpine County, the
plants are often below tiinberline. For the most part
the var. alpinum may be looked tipon as a high alti-
tude ecotype of var. laxiflonim -much as Torrey and
Gray did— possessing a series of distinctive morpho-
logical features which allow It to be formally recog-
nized at a ta.xonomic rank. Within the small geograph-
ical range of the variety, the morphological variation
is not too extensive, but it does appear that the vari-
ety may gradually grade into var. laxijlonim in the
Tioga Pass area where the southern limit of var. alpin-
um Is encountered.

One unnamed, and seemingly related series of pop-
ulations, occurs in the higli mountains of central
Nevada. I have been luiable to place the plants from
the Toiyabe Dome area of the Toiyabe Mountains;
however, I strongly suspect these populations repre-
sent alpine forms of var. laxiflonim such as I have
seen elsewhere in Nevada. The plants are depauperate,
with long scraggly branches, scattered leaves, and
small compact inflorescences of wiiite flowers. Some
representative collections are Hitchcock and Martin
5616 (OKL. UC, UTC, WTU); Linsdale and Linsdale
970 (CAS); and Maguirc and Holmgren 25996 (GH,
NY, UC, UTC).

6h. Eriogonum microthecum var. inicrothecuin
Eriogomiin microthecum Nutt., Proc. Acad. Nat. Sci.
Philadelphia 4:15. 1848. - Eriogonum microthecum
Nutt. ssp. typicum S. Stokes, Gen. Eriog. 74. 1936. —
"Hills In Oregon, east of Walla-walla [sic]," possibly
near Huntington, Baker County, Oregon, August
1834, Nultafi s.n. Hoiotypo. BM!' Isotypes. BM, GH!

Eriogonum idahoense R\db.. Bull. Torrey Bot.
Club 3'^):307. 1912. - Eriogonum microthecum Nutt.
var. idahoense (Rydb.) S. Stokes, Gen. Eriog. 74.
1936. - Weiser, Washington County. Idaho, 7 July
1899, M. E. Jones 6511. Holotype, NY! Lsolypes,
BM,MO, POM, US!

Erect and spreading shrubs (2.5) .1-5 dm high and 3-7 dm
acrcss; /eares oblanceolate to elliptic, the leaf-blades (0.8) 1-2
(2.7) cm long, {i) 4-9 (12) mm wide, densely tomentose
below, floccose to essentially glabrous above, the tomentuin
whitish, the margins plane or with thickened margins, not
revolute, the apices acute and even apiculate in somQ: jlower-
iiig stems slender to ± stout, 3-7 cm long, tloccose to glab-
rous; inflorescences dense to open cymose. 3-10 cm long.

BIOLOGICAL SERIES. VOL. 1 3, NO. 1 NOTES ON ERIOGONUM - VI

33

tloccose to glabrous; ;/M'o/i«TP.s- narrowly turbinate, 2.5-3 mm
long, tloccose to glabrous;ffoi«'n' yellow, 2-2.5 mm long, the
midribs and bases brownish-\ellow, the tepals similar, the
outer whorl of tepals tapering at the bi\s.e:acl!eiies 2-2.5 mm
long. - Representative collections: Cronquist 6127 (CAS,
COLO, GH, MICH, NY, RSA, TE.X, UC, UTC, WS, WTU),
7743 (CAS, NY, RSA, WS, WTU). 77-^7 (CAS, NY, UC, WS,
WTU);fi/7/o/i 935 (GH, NY, UC, WS, V/TV); Henderson 5431
(CAS, GH, MO): Hobngi-cn and Reveal 1346. 76^7 (ARIZ,
BRY, CAS, DS, GH, KSC, MO, NY, OKL, RS, RSA, VC. US,
UTC, WTU): J. T. Howell 1209,S (A, CAS, GH, NY):A7. E.
Jones 2SS59 (BM, CAS, MO, POM, UC); Magtiire and Holm-
gren 26730 (CAS, IDS, GH, MO, NY, POM, US, WS, WTU);
£. / Palmer 37990 (A, NY, US, WTU); Peck IS63S (NY,
WILLU); Thompson 11901 (A, CAS, DS. GH, NY, POM, \}Q.
US, WTU).

Distiibiitioii.

Dry rocky slopes anii hills on clay to sandy-loam
soils of central Oregon from southeastern Wasco
County, and southern Gillman County southeast into
the John Day Valley of Wlieeler and Grant cotmties
and northern Crook County; disjunct in southeastern
Baker County and adjacent northern Malheur Cotm-
ty, Oregon and adjacent Washington County, Idaho;
in isolated and very scattered populations in northern
Hiunboldt County, Nevada; and in eastern Modoc and
Lassen counties, northeastern California, from
2,200-5,500 feet elevation. Flowering from July to
September. Figures 20 and 21 .

The type of Eriogomim microthecum was collect-
ed by Thomas Nuttall in August of 1834. The exact
location cannot be determined althougli comparison
of modern specimens with that of NuttalFs is rather
revealing. The route followed by Nuttall is clearly
reported by Wyeth (189Q) and Townsend (1839).
After following the Boise River to the Snake River in
western Idaho, the party crossed the Snake River on
23 August, and entered the present-day state of Ore-
gon. Wyeth guided the party northward overland to
the Malheur River which they then descended back to
ihe Snake. On the 26th, they arrived at the Burnt
River and followed it northward until the 28th. This
is the only area where Nuttall could have found this
species as determined from modern-day distribution.
The type compares favorably with plants collected
more recently from this area and is found to match
almost identically with M. E. Jones 651 1, the type of
E. idahoense. It is suggested that the type of E.
microthecum was collected on the rocky slopes above
the Snake River below Huntington, Baker Co., Ore-
gon, on or about the 26 August 1834.

No sooner had Nuttall named the species than
authors in America and England were applying the
concept of var. microthecum to specimens of var. lax-
ifhrum—an error that has continued in tloras, man-
ual, and revisions of the genus up to 1964.

The var. microthecum, as here defined, consists of
two rather easily distinguished groups. The one, rep-
resented by the type, has tloccose flowering stems, in-

florescences and involucres, oblanceolate leaves, and
densely compact inflorescences. The other phase,
restricted to the John Day Valley region of central
Oregon, has subglabrous to glabrous stems, involu-
cres, and a more open, glabrous inflorescence. The
leaves are generally more elliptical than oblanceolate.
The populations in northern Nevada and California,
while similar to each other, tend to bridge the mor-
phological gap (although not completely so) between
the two Oregon types.

In the field, these two groups attributed to var.
microthecum, ditTer in minor ways. For e.xaiuple, the
plants associated with the type are usually found
associated with species of Artemisia on open slopes or
the lower foothills. If these plants are associated with
Pimis, the species is normally pinyon. The John Day
Valley phase is also associated with Artemisia, but
less directly, and the Eriogomim is more ol'ten on
open slopes under taller species of conifers than pin-
yon. The soils are different too. That of the typical
phase tends to be more clayey than that found in
John Day Valley wliich is distinctly a loam soil. The
plants in northwestern Nevada are almost always on
heavy clay soils, and the same is likely true of the
plants in northeastern California. If further work
should demonstrate the distinction between these
two groups of var. microthecum, the definition of the
variety will become even more restricted.

The var. microthecum clearly evolved from var.
laxiflorum. The two variants are very similar and non-
flowering plants can be difficult to place, especially
when they come from northwestern Nevada or north-
eastern California. The presence of yellow flowers is a
rather dubious feature, but when added to the other
minor morphological differences and the geographical
range is considered, a varietal distinction seems justi-
fied. The distinction between var. microthecum and
var. ambiguum is one that is largely based on ecologi-
cal and geographical differences. To be sure, morpho-
logical differences exist, but these are somewhat over-
lapping in nature. However, in the field, the two seem
quite different and one never finds var. microthecum
in the ecological habitats where the vast majority of
specimens of var. ambiguum occur.
6i. Eriogonum microthecum var. ambiguum (M. E.
Jones) Reveal in Munz

Eriogomim microthecum Nutt. vnr. ambiguum (M. E.
Jones) Reveal in Munz, Suppl. A Calif. Flora 61.
1968, based on E. auretim M. E. Jones var. ambiguum
M. E. Jones, Proc. Calif. Acad. Sci. 11,5:719. 1895.-^
Eriogonum fruticosum A. Nels. var. ambiguum (M. E.
Jones) A. Nels.. Bot. Gaz. 34:23. 1902. — Eriogonum
corymhosum. Benth. in DC. var. ambiguum (M. E.
Jones) M. E. Jones, Contr. W. Bot. 11:14. 1903. -
Along the Hockett Trail in the valley of Little Cot-
tonwood Creek on the east slope of the Sierra Nevada
near Lone Pine, Inyo County, California, 24 August
1891, Covillc and Funston 1688. Holotype, US! Iso-

34

bRlGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

Kig. 20. Distribution map of /:

nogomw, micmthecum var. micmthecum. Portions of Oregon, Idaho. California, and Nevada

BIOLOGICAL SERIES. VOL. 13, NO. 1 NOTES ON ERIOGONUM - VI

35

1 i\~~iy /tZ^ -i*"

Fig. 21. liluslration o( Eriogomnn micmthecuin var. micro rlit'cinii.

36

imiGllAM YOUNG UNIVERSITY SCIENCE BULLEriN

type, NY!

Eriogonuin icnelliini Toir. var. criuiuliuni Gand.,
Bull. Soc. Roy. Bot. Belgique 42:198. 1906. -
Truckee Meadows, Nevada County, ralitornia, 1892,
llillman s.n. Holotype, LY! Isotypes, RLNO. UC!

Eriogonum mkrothecum Nutt. var. expansum S.
Stokes, Gen. Eiiog. 76. 1936. - Rocky places in the
Wliite Mountains, probably Inyo County, California,
September 1898, Pwpus 6464. Lectotype, UC! Iso-
lectotypes, SD, US!

Ercci or spreaJiiig siilishrubs or shrubs. (I..S-5 dm liigli and
1-8 dm across; leaves linear-oblaiucolatc to elliptic. 0.8-2.5
cm long, (2) 3-6 (8) mm wide, densely lomentose below,
floccose (rarely subglabrous) above, llie tomentum wliilish or
reddish-brown, the margins plane or with thinkened margins,
occasionally revolute, the apices acute to rounded, rarely
apiculate; /7">vfn»,? sleins stoutish, 2-10 cm long, tomentose
when young, becoming tloccose or infrequently subglabrous
at maturity; injlorcscences open cymose, 1-5 (12) cm long,
tomentose to tloccose, rarely glabrous, the tomentum whitish
or reddish-brown; involucres 2-2.5 mm long, turbinate,
tomentose to tloccose, glabrous along the angled ridges;/7ow-
ers yellow, (1.5) 2-2.5 (3) mm long, the midribs and bases
occasionally brownish-yellow in fruit, the tepals essentially
similar, the outer whorl of tepals rounded at the base;
aclieiies 1.5-2 mm long. - Representative collections: /t/ev-
aiider and Kellogg 2542 (GH. NY, VC, WS, WW); Archer
7018 (ARIZ, NA, NY, VC): Balls and Everett IH024 (CAS
NY, RSA); Duran 31U (ARIZ. BM, BR, CAS, DS, GH,
MICH. MO, NY, POM. RSA, SD, UC, US, UTC, WIS, WTU);
Ferris 6754. h7(,4 (US. POM); Graham 65, 75. 235 (UC);
Hall II8HI (GH, VC); Heller IU220 (CAS, GH, MO, US);/
T. Howell 14324. 22841. 24155. 26294. 40223. 40987
(CAS); Mwiz 21165 (CAS. NY. RSA); Reveal 392, 414
(CAS, UTC. WTU): Roos and Roos 5931 (CAS, RSA);
Shocklev 544 (ND-G, OKL, UC), 66rt (ND-G, UC); Stokes
s.n. (SD. UT); Train 43IS (NA, RSA, WIU); Twisselmann
5636. 5833 (CAS).

Distribution.

Dry rocky places from southern Washoe County.
Nevada and adjacent Nevada County, California,
southward along the eastern tlank of the Sierra
Nevada and adjacent desert ranges to Mineral and
Esmeralda counties. Nevada, and Mono and Inyo
counties, California, from (5,000) 6„500-10,500 feet
elevation. Floweiing from July to September. Figures
22 and 23.

The var. iimbigiium has suffered from numerous
interpretations since it was described by Jones in
1895. First it was referred to Eriogonum aiireum, a
synonym of E. coryiuhosuin var. gliiiinosuin. then
transferred to E. fruticoswn by Aven Nelson as part
of an error discussed under E. corymbo.sum (Reveal.
1968), and finally placed in E. corymbosum by
Jones-all of this done in a period of eight years!
However, it was not put into its proper taxonomic
position until some sixty years later when 1 called
attention to this forgotten entity (Reveal & Munz,
1968). In the intervening year Gandoger (1906) and
Stokes (1936) rediscovered and renamed the variety.

I"ig. 22. Distribution map of Eriogonum micrnihccnm var.
ambigmim. Portions of California and Nevada.

first under E. teneUum and finally under E. microtbe-
ciim. Even so the situation with this entity is not
st;ible. As one may see from the key, the separation
t)f var. ainhiguum from var. microlltecwn is a matter
of some difficulty. However, as the two differ in sev-
eral minor ways which seem to be consistent, they are
given formal taxonomic recognition as distinct vari-
ants.

In the field, var. ambigmim is a most interesting
subject. In much of its range it occurs with var. laxi-
llonim ;ind it may be easily looked upon as a yellow-
fiowered pluise of var. laxijlonim. The two grow
together in the lower desert foothills in Mono and
Inyo counties, California, but in the higher elevations,
especially on the Sierra Nevada, the var. iimbigiium
can be found separated from var. laxijlonim.
7. Eriogonum effusum Nutt.

DitruscK liranched spreading shrubs 1.5-7 dm high and
3-12 (15) dm across, the lower stems grayish to reddish-
brown, woody, the bark shreddy or exfoliating in platelike
segments, lealless or leafy on the upper portions only, the
upper branches herbaceous, slender to ± stout, scraggly and
fragile, tloccose to glabrous, the stems dark green; /('orcv soli-
tary and scattered along the lower 1/4 of the herbaceous
stems, ± decurrent, the leaf-blades thin, linear to linear-
oblanceolatc or oblanceolate to oblong, (1) 1.5-5 (6) cm
long, 1-7 mm wide, densely white-tomenlose below, the mid-
veins usually distinct and not densely tomentose, white-floc-
cose and green above, the tomentimi and leaf-color drying
blackish in most, the margins entire or slightly undulate in
some, plane or revolute, the apices acute to subacute or
obtuse, the bases cuneate, the leaves soon deciduous, the
plants often leafless in late anthesis, the petioles slender, 2-7
mm long, tloccose below, subglabrous to glabrate above, the
petiole-bases elongated-triangular. 1.5-4.5 mm long, 1-2 mm

BIOLOGICAL SERIES. VOL. I 3, NO. I NOTES ON ERIOGONUM - VI

37

Fig. 23. Illustration of Enogoinim miciotheciim var. uinhiguuin sliowing extremes in variation.

38

BRir.HAM YOUNG UNIVIiRSITY SCIENCE BULLETIN

wide, tomcnlose to tloccosc without, tomentose within, not
clasping tlie stems; /?ovi'OT«^ sleitis slender to ± stout, lew to
numerous per phmts, (2) 3-8 (10) em long, tloccose to glab-
rous, the tomenlum blackish at maturity in most, often more
heavily pubescent among the basal leaves and below to the
woody stems; injlorcsccnccs densely cymose, diffuse and ±
congested with numerous trichotomous and dichotomous
branches, (0.5) 1-3 (4) dm long, 1-4 dm across, white-lloc-
cose to green and glabrous, the tomentum and branches often
becoming blackish; brads scalelike, ternate, mostly triangu-
lar, 0.5-2 (5) mm long, 0.5-1.5 mm wide, tloccosc to subglab-
rous or glabrous, usually dark brown to black without, cot-
tony tomentose within, connate at the b-dse; peduncles, when
present, slender and up to 2.C cm long, mostly floccose,
restricted to the lower nodes; involiieres solitary, turbinate,
1.5-3 mm long, 1-2 mm wide, sparsely tloccose without, glab-
rous within, the 5 minute acute to triangular lobes 0.3-0.6
mm long, the bractlets linear to oblanccolate, 1.5-3 mm long,
fringed with several short capitate gland-tipped cells, the ped-
icels 1.5-4.5 mm long, glabrous; /7(nvf/-.v white with greenish
or reddish midribs and bases, 2-4 mm long, glabrous within
and without except for scattered pilose to strigose hairs with-
out in some and for a few microscopic glands within along
the midribs and lower part of the tlower tube, the tepals
nearly similar to slightly dissimilar, the outer whorl of tepals
elliptic or more frequently obovate, 1.3-I.H mm wide, the
apices rounded to emarginated, the bases rounded to nearly
cordate, the inner whorl of tepals narrowly oblong lo oblong,
0.6-1 mm wide, the apices mostly acute, united about ',4 the
length of the llovver; slainens mostly exserted, 2-4.5 mm
long, the filaments sparsely pubsecent to short pilose basally,
the anthers reddish to purplish-red, 0.5-0.7 mm long, oblong;
achenes brown, 2-2.5 mm long, the large globose base taper-
ing abruptly to a long 3-angled beak.

Distribiitiiiii.

Dry rocky slopes to sandy plains and flats or infre-
quently on heavy clay slopes, chalky bluffs, or gyp-
sum outcrops, in the mountains and on the Great
Plains frotn southwestern South Dakota .southward to
eastern Wyoming and extreme western Nebraska
across eastern Colorado in the Front Ranges and on
the plains into north-central, ceiiti;il, and northeast-
ern New Mexico, and in west-central Kansas, from
3,000-7,500 feet elevation. Flowering from June to
Septetuber.

The Great Plains Buckwheat, t'riogdiiuin cjjiisiiin.
has been placed under /•,'. micrnlhecum as a variety by
some authors. This treatment is most unrealistic as
the two are well separated on morphological and geo-
graphical bases. Both species occur together in south-
ern Colorado and northwestern New Mexico, but in
this area the two are very distinct and no introgres-
sion or hybridization is seen in the field nor in the
numerous herbarium specimens examined.

Within the species as now recognized, only two
varieties are recognized.

Key to the varieties of Eriogomim effiisiim

A. Leaves oblanccolate to oblong or obovate, (1) 1.5-3 cm
long, (2) 3-7 mm wide, not rcvolute; South Dakota and
Wyoming southward to New Mexico

8a. var. effustim.

AA. Leaves linear to linear-oblanceolate, (2) 3-6 cm long,

1-2.5 (3) mm wide, usually revolute; west-central Kansas

8b. var. rosmarlnoidcs.

7a. Eriogonuin effusuni var. effiisum

Hriogonwu ejjusum Nutt., Proc. Acad. Nat. Sci.
Philadelphia 4:15. 1848. — Eriogomim microthecum
Nutt. var. effimim (Nutt.) Torr. & Gray, Proc. Amer.
Acad. Arts 8:172. 1870. — Eriogomim effustim Nutt.
ssp. typicum S. Stokes, Gen. Eriog. 78. 1936. -
"Rocky Mountains of the Platte," likely along the
Platte River in extreme western Nebraska or adjacent
Wyoming, late May or early June 1834, Niinall s.ii.
Ilolotype. BM! Isotypes, GH, PH!

Eriogomim myriantlnim Gand., Bull. Soc. Roy.
Bot. Belgique 42:191. 1906. - Fort Collins, Larimer
County, Colorado, 1 September 1898, Crandall s.n.
Lectotype, LY! Isoieclotype. US!

Eriogomim lu'iviiskense Rydb., Flora Rocky Mts.
lOhl. (917. - Eriogomim imilticeps Nees in Wied-
Neuw. ssp. nehraskensc (Rydb.) S. Stokes, Gen.
Eriog. 94. 1936. - Eriogomim paiiciflonim Pursh var.
nebraskeiise (Rydb.) Reveal, Great Basin Naturalist
27:1 13. 1967. - Prairies in Kimball County, Kimball
Co., Nebraska, 12 August 1891, Rydberg 336. Holo-
type, NY! Isotypes, KSC, NY, US!

Diffusely branched shrubs (1.5) 2-5 (7) dm high and up to
15 dm across; leaves oblanccolate to oblong, (1) 1.5-3 cm
long, (2) 3-7 mm wide, densely tomentose below, white-tloc-
cose to glabrate or glabrous and green above, the margins
plane, not revolute, or if so, then the plants from the moun-
tains of central Colorado; /Tou'cri/i^ steins slender to ± stout,
3-8 cm long, floccose to glabrous; inflorescences 1-3 (4) dm
long, tloccose to glabrate or subglabrous; mi'o//(crfs 1.5-2.5
(3) mm long; //oivCTW white, 2-4 mm \on%\ aclienes 2-3 mm
long. - Representative collections: Arsene and Benedict
15418 (P, US): Bacigahiin M2. 912 (GH, VC): Clements and
Clements 21 (ISC, NY, US); Cloker 2948 (ARIZ, CAS, GH,
NY, UC, US); Dodds 1957. 2041, 2069 {COLO): Eastwood
123 (CAS, COLO, GH, UC. VS): Elilers 7827 {ARM. MICH.
ND); Engelmann s.n. (GH, MO, NY); Ewan U691 (COLO,
ND. NO, OKL); Fendler 768 (GH, MO); Fremont s.n. (NY);
Goodman 2008 (ISC, MO, NY, OKL): Hall and Harbour 502
(BM, GH, MO, NY, US): Heller 14307. 14324 (MO, WTU);
litis and litis I86.'i8 {\\\S): Johnston 403 (GH. MICH, MO,
VS): Jones 544 (BM. BR, NY, POM, VTC):Mulford s.n. (GH,
MO, NY, VS): Nelson 329 (MO. NY. POM, RM), 1138 (GH,
RM, US, WIS). 7631 (ARIZ, COLO, GH. ISC. MO. NY. POM.
RKNO. RM, US); Parrv 32! (GH, ISC, MO, NY):Ramalev
863 (CAS, COLO, RSA, UC, WTV): Reveal and Davidse 865.
869. 870 (BRY, CAS, GH, LL, NY, OkL, RSA, SMU, TEX.
UC, US, UTC); Robhins 900 (COLO, NY, VC):Rollins 1072
(G, GH, ND, RM, VC. US, WTU): Rvdherg 185, 335 (NY);
Sheldon 572 (LIVU. UC. VS): Standlev 6941. 7107 (US);
Stokes 210 (ARIZ, BM, CAS, DS, GH. MO. NY. POM. RSA,
UC, US, UTC); Waterfall 12041 (OKI . RSA, US): Williams
2449 (G, MO, ND, UC, US, WTU).

Distribution.

Rocky slopes of the mountains and ranges onto
the sandy soils of the plains, from Pennington
County, South Dakota, southward into Converse and
southern Niobrana counties, Wyoming, southward in
southeastern Wyoming and adjacent extreme western
Nebraska to central and east-central Colorado in the
Front Ranges and on the Great Plains lo northern

BIOLOGICAL SERIES. VOL. I 3, NO. : NOTES ON ERIOGONUM

VI

39

New Mexico from eastern San Juan County eastward
to Union County and with an outlying population in
Socorro County, New Mexico. Flowering from June
to September. Figures 24 and 25.

Fig. 24. Distribution map o( Enogonuin effusiiiu var. eJiusion.

The type of Eriogoniim effuswn was collected in a
premature state by Thomas Nuttall while on his trans-
continental trip with Nathaniel J. Wyeth The e.xact

location cannot be determmed, but on the basis of
known distribution of the species, and his route, the
type area can be estimated fairly accurately. On 28
May, the party reached Scotts Bluff where E. effusum
is known to occur. That Nuttall collected here is
attested by Townsend (1839) who commented:

These mounds (at Scotts Bluff] were of hard yellow
c ay, without a particle of rock of any kind, and
along their bases, and in the narrow passages tlowers
ot every hue were growing. It was a most enchanting
sight: even the men noticed it. and more than one of
our mattcr-ot-tact people exclaimed, beautiful beau-
tiful Mr. N|uttall| was here in his glory. He rode on
ahead ol the company, and cleared the passages with
a trembling and eager hand, looking anxiously back
at the approaching party, as though he feared it
would come ere he had finished, and tread his lovely
prizes under tool.

The species is equally common on the Laramie Hills,
and it is possible that Nuttall gathered his specimens
on 2 June when they crossed these mountains in
southeastern Wyoming.

The variety, as now defined, includes Eriogoniim
nebraskense. An understanding of this taxon has
plagued botanists for years, and even its placement in
synonymy here may not be a final answer. The distin-
guishing features are the pubescent flowers and
densely tomentose leaves, both characters that may
have come about as a result of int regression or
hybridization with E. paiiciflomm Pursh (Reveal,
1967). Its placement with E. paiidflonim was based
on an over emphasis of the pubescent flowers, a point
of view that is now considered to be in error. How-
ever, whether or not the species should be recognized
as a hybrid species remains to be demonstrated by
cytological studies.

In the field, var, effusum is a spectacular shrub. It
torms large spreading masses of intricately branched
inflorescences capped with numerous whitish to pink-
ish flowers. In some areas, as in central Colorado, the
shrubs are massive, often being up to 1.5 m across,
but in parts of northern New Mexico, they are low
scraggly bushes with only a few branches. This variety
miglit make an excellent garden shrub were it not for
the fact that when it is not flowering, the dead and
dried branches are rather bare and forlorn looking,
and even in the field the plants look unbecoming.

The suggestion that var. effusum forms hybrids
with Eriogumim microlhecuin appears to be un-
founded. In this investigation, the distinction be-
tween these two taxa has been easily made, and not a
single specimen has presented a problem. Conse-
quently, E. effusum is maintained as a distinct spe-
cies.

7b. Eriogonum effusum var. rosmarinoides Benth in
DC.

Eriogonum effusum Nutt. var. rosmarinoides Benth.
in DC, Prodr. 14:18. 1856. - Along Smoky Hill
River, Gove or Trego counties, Kansas, 21 July 1845,
Fremont 181. Lectotype, NY! Isolectotype GH k'
MO!

40

BRIUIIAM YOUNG UNIVERSITY SCIENCE BULLETIN

M-^M

Fig. 25. Illustration o( l-'riogoniiin effusimi var. eflKsuin.

Lriogonum lielichrysoidcs Gaud.. Bull. Soc. Rny. Rydb., Biiiuniia 1:87. 1*^)31. Hriogoiium cjjiistiiii
Bot. Belgique 42:192. 1906. - t'riogoniim micro- Nutt. ssp. helichrysolJes (Gand.) S. Stokes, Gen.
//lecMW Null. var. //eZ/c'/imo/c/ei (attrib. lo Gaud, by) Eriog. 78. 1936. - In canyons of Gove County,

lilOLOGICAL SERIES. VOL. 13, NO. I NOTES ON ERIOGONUM - VI

41

Kansas, 21 July 1895, .4. S. Hllclicock 44S. Holo-

type, LY! Isotypes, GH, KSC, MO, NY, US!

Low diffusely branclied shrubs 1.5-3.5 (4) dm high and up
to 8 dm across; leases hnear to linear-oblanccolale, (2) 3-6
cm long, 1-2.5 (3) mm wide, densely tomentose below, Hoc-
cose above, often revolute: flowering stems slender, 2-4 cm
long, tloccose: inflorescences 0.5-1.3 (1.6) dm long, tloccose
to nearly glabrous; ;>n'o/!«rex 2.5-3 mm long: flowers white,
2-2.5 (3) mm long; ac'/;e/it'.s 2-2.5 mm long. - Representative
collections: Agrelius and Stephens s.n. (WIS); Gates lh676
(KSC, US); Goodman 220t) (ISC, MO. NY, OKL); Gordon
s.n. (MO); Hitchcock 55 7 (GH); Horr and Horr 4154 ( KANU.
U\): McGregor 12471. IJh52. IJh44. IJhh6 i.KANV): Rich
1297 (KSC); Rvdbcr^ and Imler IU34 (COLO, KANU, KSC,
MO, NY). 1132 (KANU, KSC. NY); Stephens 8996. 9770
(KANU). 7 WcSS (KANU, UC); Weber 142 (KSC).

Distiibuiioii

Clay slopes and chalky limestone outcrops in
Logan. Gove. Trego. Scott, and Lane counties. Kan-
sas, from 3.000-3.500 feet elevation. Flowering from
July to September. Figiues 26 and 27.

The type of var. rosmarinoides was collected by
John C. Fre'mont on the last leg of his western trip
which lasted from 1843 to 1845. However, when it
was described by Bentham (1856), the location date
was given as "California." In 1870, Torrey and Gray
called attention to this error, but the name remained
associated with the California flora until this fact was
pointed out again by Reveal and Munz ( l'-'68).

The Gandoger name has been the subject of some
nomenclatural confusion. Even thougli Gandoger spe-
cifically stated his name, Eriogomim helichrysoides,
to be a new species, Rydberg considered the name to

Fig. 27. Illustration of Eriogomim cffusuin var. rosmarinoides.

Fig. 26. Distribution map o'i Eriogunum effusum var. rosmarinoides. Logan, Gove, Trego, Scott, and Lane counties, Kansas.

42

DKIGHAM YOUNG UNIVEKSITY SCIENCE BULLETIN

be merely a variety of E. microtheaim based on
Gandoger's past usage of this rank in other taxonomic
treatments. Nevertheless, as Gandoger was specific in
his application of the rank he used in tiiis paper, it
cannot be questioned.

In tiie Held, var. rosmarinoides occurs in exposed

areas on steep chalky bluffs in west-central Kansas.
So far as known it is endemic to this area. The low
shrubs are rather woody, tend to be spreading, and
have numerous branches. The difTerences between
var. rosmarinoides and var. effimim are subtle, but
their geographical ranges do not overlap.

SUMMARY

This revision discusses seven species of Eriogonuni
(Polygonaceae) found in the western United States of
North America. One of the two major species. E.
microthecum Nutt., is composed of nine varieties
which range over a wide area of the Far West. The
other major species, /•.'. effitsum Nutt., is found in a
smaller geographical area mainly east of the Conti-
nental Divide. The five remaining species are pri-
marily restricted to the Colorado-Green rivers drain-
age basin of Utah, Colorado, New Mexico, and Ari-
zona. Together these entities form the core of a group
of plants known as Section Corymbosa Benth. in DC.
The remaining species of the section (not discussed
here) are typified by E. desenicola S. Wats., E. lepto-
cladon Torr. & Gray, and E. corymbositm Benth. in
DC. Tlie latter group of species was reviewed in Part

V of this series. Notes on Eriogonuni. The new en-
tities proposed in the present paper are E. micro-
theaim var. corymhosoides and var. jolmstonii from
southern California; £. microthecum var. lapidicola of
eastern California, southern Nevada, and perhaps
western Utah; and, E. microthecum var. alpinum
from the Sierra Nevada of east-central California. The
proper characterization of E. ericifolium Torr. &
Gray is proposed and E. mearnsii Parry in Britt. is
reduced to synonymy. Two new combinations are
suggested: E. microthecum vm. foliosum and E. erici-
folium var. pulchrum. Keys and descriptions for each
entity are provided as are maps showing the distri-
bution of each taxon. Illustrations of the variation
within E. microthecum and E. effusum are provided.

ACKNOWLEDGMENTS

I extend my sincere thanks to Dr. Stanley L. Welsh
of Brigliam Young University, Provo, Utah, for sug-
gestions on various aspects of this study. Dr. Wallace
R. Ernest and Mr. Conrad V. Morton of the Depart-
ment of Botany, United States National Herbarium,
Smithsonian Institution have kindly read the manu-
script and made many helpful suggestions. Fieldwork
since 1965 has been supported by both private
financing and a National Science Foundation grant to
Dr. Arthur Cronquist of the New York Botanical
Garden for the Intermountain Flora Project, a coop-
erative program between the New York Botanical
Garden and Utah State University. Summer fieldwork
conducted in 1968 was aided by Contract No.
AT(04-1) Gen- 12 between the University of Cali-
fornia at Los Angeles and the Division of Biology and
Medicine, United States Atomic Energy Commission
while collecting plants on the Nevada Test Site. Cri-
tical herbarium material has been obtained through
support furnished by Brigliam Young University.
Visits to herbaria have been largely supported by
Utah State University, New York Botanical Garden,
the Smithsonian Research Foundation, the Texas
Research Foundation, Brigham Young University,
and the Atomic Energy Commission. The final phases
were partially supported by a National Science Foun-
dation Grant (GB-22645) for studies on Eriogonuin.
The illustrations were prepared by Mrs. Twila Davis
Bird.

The several herbaria which have been consulted
are noted below and abbreviated as suggested by
Lanjouw and Stafieu (1964). I am grateful to the
numerous curators who have so kindly provided me
with material for this study.

A Arnold Arboretum, Harvard University Her-

baria, Cambridge. Massachusetts
ARIZ University of Arizona. Tucson, Arizona
ASC Northern Arizona University, Flagstaff,

Arizona
ASU Arizona State University, Tempe, Arizona
BM British Museum (Natural History). London,

England
BR Jardin Botanique de I'Etat, Bruxelles, Bel-

gium
BRY Brigham Young University, Provo, Utah
CAS California Academy of Sciences. San Fran-
cisco, California
COLO Museum, University of Colorado, Boulder,

Colorado
DS Dudley Herbarium. Stanford University,

Stanford. California
G Conservatoire et Jardin bdlaniques. Geneve.

Switzerland
GH Gray Herbarium. Harvard University Her-

baria. Cambridge. Massachusetts
IDS Idaho State University, Pocatello, Idaho

ISC Iowa State University, Ames, Iowa

JEPS Jepson Herbarium. University of California.

BIOLOGICAL SERIFS. VOL. I .!, NO. 1 NO lES ON ERIOGONUM

VI

43

Berkeley, California

K Royal Botanic Garden, Kew, England

KANU University of Kansas, Lawrence, Kansas

KSC Kansas State University, Manhattan, Kansas

LIVLI The Hurley Botanical Laboratories, Liver-
pool, England

LL Liuidell Herbarimn, Texas Research Founda-

tion, Renner, Te.\as

L\ Herbiers de la Faculte des Sciences de Lyon,

France

MARY University of Maryland, College Park, Mary-
land

MICH University Herbarimn, University of Michi-
gan, Ann Arbor, Michigan

NINA Museum of Northern Arizona, Flagstaff,
Arizona

MO Missouri Botanical Garden, St. Louis, Mis-

souri

MONT Montana State University, Bozeman, Mon-
tana

NA United States National Arboretum, Washing-

ton, D.C.

ND University of Notre Dame, Notre Dame,

Indiana

ND-G Greene Herbarimn, University of Notre
Dame, Notre Dame, Indiana

NO Tulane University, New Orleans, Louisiana

NTS Nevada Test Site Herbarium, Mercury,
Nevada

NY The New York Botanical Garden, Bronx

Park, Bronx, New York

OKL Bebb Herbarium, University of Oklahoma,
Norman, Oklahoma

ORE University of Oregon, Eugene, Oregon

OSC Oregon State University, Corvallis, Oregon

P Museum National d'Histoire Naturelle,

Laboratoire de Phanerogamic, Paris, France

PH Academy of Natural Sciences, Philadelphia,

Pennsylvania

POM Pomona College Herbarium, Rancho Santa
Ana Botanic Garden, Claremont, California

RENO University of Nevada, Reno, Nevada

RM Rocky Mountain Herbarimn, University of

Wyoming, Laramie, Wyoming

RSA Rancho Santa Botanic Garden, Claremont,
California

SD San Diego Museum of Natural History, San

Diego, California

SMU Southern Methodist University, Dallas,
Texas

TEX University of Texas, Austin, Texas

UC University of California, Berkeley, California

UNM University of New Mexico, Albuquerque,
New Mexico

US United States National Museum, Smith-

sonian Institution, Washington, D.C,

USES L'nited States Forest Service Herbarium,
Boulder, Colorado

UT University of Utah, Salt Lake City, Utah

UTC Intermountain Herbarium, Utah State Uni-
versity, Logan, Utah

WILLU Peck Herbarium, Willamette University,
Salem, Oregon

WIS University of Wisconsin, Madison, Wisconsin

WS Washington State University, Pullman, Wash-

ington

WTU LIniversity of Washington, Seattle, Washing-
ton

44

BRIGHAM YOUNC. UNIVERSl lY SCIINCK BULLETIN

LITERATURE CITED

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Eastwood, A. 1893. Notes on some Colorado plants, /oe.
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1896. Report on a c-ollection of plants from San

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1931. New species of plants from western North

.America. I'roceedings of tlic California Academy of
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Gandoger, M. 1906. Le genere Eriogonum (Polygonaceae).
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Goet7.mann, W. H. 1959. Army Exploration in the Amer-
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Graustein, J. \. 1967. Tliomas Nuttall. Naturalist. Explora-
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Hooker, J. 1853. Catalogue of Mr. Geyer's collection of
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Howell, J. 1. 1944. ,\ new fruticulose Eriogonum. Z.ca//eM
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1895. Contributions to western botany. VII. Pro-
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1903. Erioaonum. Contributions to Western Bot-
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Nuttall, T. 1848. Descriptions of plants collected by Mr.
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Parry, C. C. 1889. "Eriogonum mcarnsii." fn: Britton, N.
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Porter, C. L. 1968. A flora of Wyoming. Part VI. /?csea/-c/!
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Reveal. J. L. 1966. Notes on three Utah Eriogonums. Pro-
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1967. Notes on Eriogonum - III. On the status of

Eriogonum paucitlorum Pursh. Great Basin Naturalist.
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1968. Notes on E^riogonum - V. .\ revision of the

Eriogonum corymbosum complex. Great Basin Natural-
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1969a. A Revision of the Genus Eriogonum (Poly-
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1969b. "The subgeneric concept In Eriogonum

(Polygonaceae)." //;: Gunckel, J. Current Topics in
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Reveal, J. L., and P. A. Munz. 1968. "Eriogonum." In:
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Reveal. J. L.. and V. S. Spevak. 1967. Publication dates and
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^-0J<^ -Ki^rov;o3

■i

Brigham Young University
Science Bulletin

..If

^ TAXONOMIC STUDY OF CROTAPHYTUS COLLARIS
ETWEEN THE RIO GRANDE AND COLORADO RIVERS

by

William Ingram III

and
Wilmer W. Tanner

BIOLOGICAL SERIES— VOLUME XIII, NUMBER 2
April 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany,

Brigham Young University, Provo, Utah

Members of the Editorial Board:

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

Ex officio Members:

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

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

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

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

Brigham Young University
Science Bulletin

A TAXONOMIC STUDY OF CROTAPHYTUS COLLARIS
IeTWEEN the RIO GRANDE AND COLORADO RIVERS

by

William Ingram III

and
Wilmer W. Tanner

BIOLOGICAL SERIES— VOLUME XIII, NUMBER 2
April 1971

TABLE OF CONTENTS

INTRODUCTION 1

REVIEW OF LITERATURE 1

MATERIALS AND METHODS 3

Selection and Gathering of Material 3

Statistical Methods 4

Taxonomic Characters 5

Color-pattern Combinations 5

RESULTS 6

Cluster Analysis 6

Analysis of Variance 7

Coloration and Pattern 11

Discriminant Analysis II

DISCUSSION 13

Zoological Discussion 13

Statistical Discussion 24

SUMMARY AND CONCLUSIONS 27

ACKNOWLEDGMENTS 28

LITERATURE CITED 29

A TAXONOMIC STUDY OF CROTAPHYTUS COLLARIS
BETWEEN THE RIO GRANDE AND COLORADO RIVERS

by

William Ingram III' and Wilmer W. Tanner^

INTRODUCTION

The Western Collared Lizard, Crotaphytiis coUaris
baileyi, Stejneger. is an attractive lizard of marked
variability. Its range, as now recognized, includes the
area from the Great Basin south to central Baja Cali-
fornia and Sonora; the desert slopes of mountains in
southern California, east to the Continental Divide in
southwestern Colorado and western New Mexico, and
south into Chihuahua, Durango, Coahuila, Nuevo
Leon, and San Luis Potosi (Stebbins, 1966; Smith
and Taylor, 1950).

The range of C. c. baileyi is divided by several
natural geographic barriers which have been shown to
mark boundary lines for subspecies of other saurian
forms. Stebbins (1954) describes patternal differences
of C. coUahs which are both consistent and specific
for certain geographic areas. Fitch and Tanner (1951)
proposed that at least one geographic area within the
range of C. c. baileyi. the Upper Colorado River

Basin, contains a population of Collared Lizards that
is distinguishable on the subspecitlc level from C. c.
baileyi. These studies suggest that C. c. baileyi may in
reality be a heterogeneous taxonomic unit requiring
more study.

The first step in the comprehensive study of the
populations presently contained in C. c. baileyi re-
quires a detailed study of the population represented
by the type material. An analysis of the total Western
Collared Lizard problem also includes a study of the
Collared Lizard populations located in the Chihua-
huan Desert and the Upper Colorado River Basin.
This includes the three populations" interactions with
each other. The analysis of the remaining populations
of Collared Lizards now presently contained within
C. c. baileyi and occurring in areas located primarily
west of the Colorado River and Baja, California is the
subject of another study currently in progress.

REVIEW OF LITERATURE

In 1890, Dr. Leonhard Stejneger described as new,
Crotaphytiis baileyi. The type locality was listed as
"Painted Desert, Little Colorado River, Arizona," and
the type specimen designated as U.S. National
Museum No. 15281. Other specimens included in the
type series were USNM 15282-15287.

Stejneger (1890) distinguished C. baileyi from C.
collaris on the basis of four characters: ( 1 ) two rows
of interorbitals, (2) smaller supraocular scales, (3)
narrower head, and (4) longer snout. At the time of
the description, Stejneger indicated that further in-
vestigation probably would show C. baileyi to inter-
grade with C. collaris. However, he described it as a
new species until conclusive evidence of intergrada-
tion was provided. The geographic range of C baileyi
was set forth as western New Mexico, Arizona,
Nevada, and northern Mexico.

E. D. Cope (1900), in his monumental work on
North American reptiles, recognized the differences
listed by Stejneger. However, he stated that "transi-
tions (of characters between geographic areas) are so
numerous that a distinct subspecific name is of

doubtful utility." Cope failed to support his conclu-
sions with a clear presentation of data, but merely
compared the number of specimens on hand (80) as
to the condition of their interorbitals. He failed to
recognize C. baileyi as either a species or a subspecies
because his series contained a large number of speci-
mens intermediate to C. collaris or C. baileyi with
respect to interorbital scalation. However, he states
that most of these intermediate specimens came from
the central portion of New Mexico and western
Texas, the area proposed by Stejneger as the probable
region of intergradation. Thus, one wonders why
Cope failed to recognize C. baileyi and place it as a
subspecies of C. collaris.

Stone and Rehn (1910) was the first to recognize
C. baileyi as a subspecies of C. collaris. This was based
on a series of eleven specimens from Pecos, Texas, in
which specimens with characteristics of both C.
baileyi and C. collaris were in the series.

In 1917 Stejneger and Barbour listed C. c. collaris
and C. c. baileyi in their check list using central New
Mexico as the dividing line between the two sub-

I, 2

Department of Zoology and Entomology, Brigham Young University, Provo, Utah 84601

BRICHAM YOUNG UNIVERSITY SCIENCE BULLETIN

species.

Van Denburgli and Slevin (1921) described as new,
Croiaphytus insiilaris from Angel de la Guardia Is-
land, Baja California, Mexico. Crotaphytus insularis
was separated from C. c. baileyi by three charac-
ters: (1) longer snout, (2) narrower head, and (3)
single incomplete collar (Van Denburgh 1922). Burt
(1928) showed definite relationships between the
Collared Lizards of Angel de la Guardia Island and
those of the mainland of Baja California and southern
California. The lizards of this area are also in need of
further study as a pari of those populations occurring
west of the Colorado River.

Crotaphytus Jickersoiiac, was described by
Schmidt (1922) from Tiburon Island, Sonora, Mex-
ico. It was said to be distinguishable from C. c. baileyi
by having "hindleg considerably longer than the
body, a longer more distinctly compressed tail and
slightly enlarged scales on the middorsal line of the
tail." Burt (1928) expressed doubts as to the validity
of this species, thus necessitating a further analysis of
C. dickersonae.

Burt (1928) provided tlie only complete review of
the species to date. He presented a detailed review of
the literature and an excellent grasp of the problem
of Collared Li/ard taxonomy. He recognized the need
for further study of the populations of Mexico and
southern California and saw trends of variation within
the Collared Lizards, but his investigations were not
detailed.

Burt's analysis of the data was based on three
assumptions, which are not substantiated by our data.
He examined a large series (1,252) of lizards for the
presence of ten characters, one meristic and nine
proportional. His first assumption concerns the
method used to assign lizards to either a western or
an eastern group. He divided the Collared Lizards into
two groups along state lines. The eastern group in-
cluded Oklahoma, Kansas and Texas; and the western
group, New Mexico, Arizona, Utah, Idaho, Oregon,
California and Mexico. As was pointed out by Fitch
and Tanner (1951), this completely disregards the
possibility of the two subspecies' ranges not con-
forming to state lines. This is especially perplexing
since all of the published ranges of C c. baileyi and C.
c. cnllaris designate the area of intergradation to be in
central New Mexico and western Texas (Stejneger,
1890;Stejneger and Barbour, 1917).

-Secondly, within the area Burt assigned to the
Western Collared Lizard are the ranges of two forms
of Collared Lizards, one form centered in the Great
Basin and the other in Baja California and southern
California.*

Thirdly, in the comparisons of body size and
proportions, il is obvious from Burt's data (79-345

*.'\n analysis of morphological similarity of Collared
Lizards demonstrated the difference between forms.

mm. tolal lenglh for the eastern group) that there was
no attempt to group specimens by age class. It is well
known that body propt)rlions, as well as length,
change significantly from hatchling to adult (Mayr,
1969), The use of different age classes in the analysis
of leiiglh and proportion characters may bias results
proportionally to the percent of the total sample
represented by each age class.

Fitch and Tanner (1951) separated the Yellow-
headed Collared Lizard, C. c. aiiriceps, from C. c.
baileyi. It was described as a population from the
Upper Colorado River Basin, type locality: three and
one-half miles north northeast of Dewey Bridge,
Grand Co., Utah. Crotaphytus c. auriccps was sepa-
rated primarily on the characters of coloration and
supralabial scalation, thus introducing the factors of
coloration and pattern as characters for distinguishing
Collared Lizard populations.

The taxonomic history of C. c. baileyi and other
Collared Lizards, along with their present status, is
summarized in the following synonymies.

Crotaphytus collaris baileyi Stejneger

Crotaphytus baileyi Stejneger, 1890, N. Amer. Fauna,

3:103 [Type locality: Painted Desert, Desert of

the Little Colorado R., Arizona; U.S. Nat. Mus.]

Crotaphytus collaris bailevi Stone and Rehn, 1903,

Proc." Acad. Nat. Sci. Phila., 55:30.

Crotaphytus insularis Van Denburgh and Slevui

Crotaphytus Insularis Van Denburgh and Slevin,
1921,' Proc. Calif. Acad. Sci., ser. 4, vol. 11:96
[Type locality: Angel de la Guardia Island. Baja
California, Mexico; Calif. Acad. Sci.) .

Crotaphytus collaris baileyi Stone and Rehn. 1903,
Proc. Acad. Nat. Sci. Phila., 55:30.

Croiaphytus dickersonae Sclmiidt

Crotaphytus dickersonae Schmidt, 1922, Bull. Amer.

Mus. Nat. Hist., 46:638 [Type locality: Tiburon

Island, Sonora, Mexico; U.S. Nat. Mus.] .
Crotaphytus collaris dickersonae Allen. 1933, Occ.

Pap. Mus. Zool. Univ. Mich., 259:7.
Crotaphytus dickersonae Smith and Taylor, 1950,

Bull. U.S. Nat. Mus., 199:93
Crotaphytus collaris baileyi Stone and Rehn, 1903,

Proc' Acad. Nat. Sci. Phila.. 55:30.

Crotaphytus collaris auriccps Fitch and Tanner

Crotaphytus collaris auriccps Filch and Tanner, 195 I ,
Trans. Kans. Acad. Sci., 54(4):553 [Type locality:
three and a half miles north northeast of Dewey
Bridge, Grand Co., Utah: Kans. Mus. Nat. Hist.].

Crotaphytus collaris baileyi Stone and Rehn, 1903,
Proc. Acad. Nat. Sci. Phila., 55:30.

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAPHYTUS COLL.^RIS

MATERIALS AND METHODS

Selection and Gathering of Material

Specimens of C. collahs utilized in this study were
I'rom three sources: (1) specimens in the Brigham
Young University Herpetological Museum. (2) speci-
mens borrowed from several museums, and (3) live
specimens collected in the field. Specimens from the
first two sources were used for the measurement of
meristic and proportional characters. Those collected
in the tleld were used for pattern and coloration
determination.

The museums with specimens on loan are listed
below, followed by the abbreviation to be used
througliout the remainder of this paper: Brigliam
Young University, BYU; California Academy of
Science, CAS; California State College at Long Beach,
CSCLB; University of Colorado Museum, CUM; Uni-
versity of Kansas. KU; Los Angeles County Museum,
LACM; San Diego Society of Natural History,
SDSNH; University of California, L!C; University of
Illinois Natural History Museum, UIMNH; United
States National Museum, USNM; University of Texas
at El Paso, UTEP; University of Utah, UU.

Four collecting trips were made to the area en-
compassed by this study. In May 1969, one short trip
was made to southeastern Utah and a longer trip (two
weeks in June) covered western New Mexico to the
Nexican border and most of Arizona. During these
trips specimens were collected from the Upper Colo-
rado River Basin of Utah and Arizona, and the Ciii-
huahuan Desert of New Mexico. Another extended
trip to New Mexico and Arizona in May 1970 re-
sulted in the collection of living specimens from the
Upper Colorado River Basin of New Mexico; central
New Mexico; Arizona, south of the Mogollon Rim,
and additional specimens from the Upper Colorado
River Basin of Utah and Arizona. During a final short
trip in 1970 to southeastern Utah, we secured speci-
mens from the northernmost extension of the range
in Grand County. Utah. All living specimens ex-
amined during the course of this study will be depos-
ited in the Brigliam Young University Herpetological
Museum.

Only those specimens whose snout-vent length was
longer than 80 mm. were used in this study for the
determination of both meristic-proportional and
coloration-patternal characters. This was done to
reduce the amount of bias caused by the mixing of
age classes on character determination. The cutoff
point was reached by determining the lower limit of a
907c confidence interval on the mean of the snout-
vent length of adult C. c. collahs, using Fitch's data
(Fitch, 1956). His data was used rather than data on
adult C. c. baileyi. because a realistic diagnosis for C.
c. baileyi has not been determined. Also, Fitch did
the only ecological study on the Collared Lizards and

is perhaps the most reliable source of adult snout-vent
lengths.

The I'oUowing lizards, listed by county within each
state, were examined in the course of this study:

ARIZONA - Apaclie Co.: BYU 497; LACM 16895: UIMNH
7524: USNM 29184. 38056, 45035. 58610. Cochise Co.:
CAS 35128-35135, 48615-48617 80748: USNM 8463, 8466,
8467. 14748, 19704-19706, 24462. Coconino Co.: BYU
506. 11388. 32110, 32109: UIMNH 6543. 35945,
74786-74790: USNM 15821, 15822, 60110-60113, 60115,
60117-60121. Gila Co.: UIMNH 34336, 74797, 74798.
Graham Co.: UIMNH 24507, 82348-82353: USNM 5153,
51737. 51739, 54599. 54606. Maricopa Co.: CAS 80681,
80682. Mohave Co.: BYU 32116: UIMNH 74778,
74781-74784. Navaio Co.: BYU 13574: LACM 16894:
UIMNH 74794-74796. Pima Co.: LACM 3983: SDSNH
15214: UIMNH 5899. Pinal Co.: UIMNH 74800: USNM
22129, 44681, 44708. Santa Cruz Co.: BYU 32106: LACM
26833: UIMNH 5900; USNM 16807, 17183. Yavapai Co.:
BYU 33322: UIMNH 43208, 74767-74777, 82354; USNM
11860, 14814, 15689, 15690.14710,15892,22206,59750.

COLORADO - Baca Co.: CUM 9678-9680, 11340, 11343,
13666, 21727, 32278-32280. Bent Co.: CUM 19652, 19653.
Las Animas Co.: CUM 1292, 2939, 7560-7562. 9675, 9681,
10030-10034, 11345, 32276. Mesa Co.: BYU 11342. I 1344.
Montezuma Co.: BYU 1577. 32108. Otero Co.: CUM 19654,
19655. Pueblo Co.: CUM 2622. San Miguel Co.: CUM 1333,
4448, 4450, 4451, 4453, 4456, 4458.

KANSAS - Anderson Co.: BYU 898. Montgomery Co.: BYU
22167. Wilson Co.: KU 41. 45. 46, 48-50, 54.

NEW MEXICO - Bernalillo Co.: USNM 58604. Chaves Co.:
LACM 3974-3976. Dona Ana Co.: LACM 3971: USNM
22268, 25423: UTEP 54. Eddy Co.: LACM 3973,
16981-16983: UIMNH 8690: USNM 93034. Guadalupe Co.:
LACM 16984, 16985. 16987; USNM 32862. Hildago Co.:
BYU 32107: LACM 3977. Lea Co.: USNM 94360. Lincoln
Co.: LACM 16990. Luna Co.: BYU 31940, 31942, 31944,
32120, 32121; USNM 44955, 80072. McKinley Co.: USNM
27738. Otero Co.: LACM 16975, 16988. Quay Co.: USNM
44940. Rio Arriba Co.: UU 3724-3732. Santa l-e Co.: CUM
7007: LACM 16907. 16908, USNM 8408, 8471. Sierra Co.:
LACM 3981, 16992. Socorro Co.: LACM 3979. 3980,
16909. 16910, 16918, 16919, 16923, 16924, 16927-16929,
16931, 16932. 16934, 16935, 16940, 16942, 16944, 16945,
16947-16953. 16957-16962, 16966-16972, 16976, 16977,
16979; USNM 44573. Taos Co.: CUM 7006.

OKLAHOMA - Carter Co.: BYU 500, 1574.

TE.\AS ~ Bexar Co.: BYU 13047, 13050, 13051. Brewster
Co.: USNM 32852, 103663. Clay Co.: USNM 32857, El Paso
Co.: USNM 59351, 59352; UTEP 52. 55-57; UU 493, Garza
Co.: CUM 32277. Llano Co.: USNM 42309. Randall Co.:
CUM 13554-13556. Roberts Co.: USNM 32866. Stephans
Co.: BYU 13117. ValverdeCo.: USNM 32850.

UTAH - Grand Co.: BYU 1625, 1626, 10338, 12854,
12855, 31949, 31950, 31981. San Juan Co.: BYU 1461,
1464, 12619, I3006-I3008, 16484. 16801. 16802,
18333-18340, 21706, 31951, 31982, 32088, 32112-32117;
UU 1461, 2427.

MEXICO - Chihuahua: BYU 13383-13386. 13410. 13411.
13736, 14211. 14212, 15184, 15I86-I5188, 15305, 15325,
16969-16976, 17010. 17014; KU3378, 33789. 44127: UC
70704; USNM 14242. Coahuila: UC 24721. Nuevo Leon:
USNM 2728. Sonora: CSCLB 2752-2755, 2757, 2759-2764;
LACM 8798, 8799, 52882. 52886; UC 10163; USNM 2694.

BRIGHAM YOUNC; UNIVERSITY SCIENCE BULLETIN

Statistical Methods

Upon initial examination of both living and pre-
served specimens, three distinct populations (groups)
were postulated: (1) Upper Colorado River Basin. (2)
central Arizona. (3) Chihuahuan Desert. An initial
analysis consisting of six steps was performed to test
the null hypothesis of no difference between groups.
Multivariate techniques of data analysis were used
extensively in this study for reasons to be discussed
later.*

Step ( I ).

There were 66 meristic and proportional char-
acters chosen to represent all observable areas of
phenotypic variation. Tliese characters included scale
counts, proportions of body parts, and those pat-
ternal aspects that remain even after long periods of
preservation. Body parts were measured using a
Golgau Vernier Caliper. All scale counts of paired
structures (e.g. supralabials) were done on the right
side only.
Step (2).

A random sample of ten males and ten females
from each proposed group was selected from the pre-
served material on hand. Ostle's random number table
and method for entering the table randomly were
used to select the random sample (Ostle. 1963). To
enable the use of multivariate analysis, only those
specimens that possessed some state of each of the 66
characters were used. For example, specimens that
were damaged in some aspect were omitted from this
part of the study.
Step (3).

The characters were measured on the sample and
correlation analysis was performed. This eliminated
those characters highly correlated with each other
(hence measuring the same source of variation). Wlien
two or more characters were found to be higlily cor-
related (0.75 or greater) the character with the
greatest variation between groups was selected to
represent all the correlates, and the other characters
were dropped from the analysis.
Step (4).

A data organizing technique developed and pro-
grammed by Wishart (1968) was used to group the
lizards in clusters of highest morphological similarity.
This technique, known as Ward's Minimum Variance
Cluster Analysis, arranges individuals in hierarchal
minimum variance clusters, thus grouping together
those lizards that are most alike, as defined by the
characters measured. Chosen to represent the range of
geographic locations available, 80 individuals were

*AII statistical techniques which are relatively new or
unfamiliar to workers in herpetological taxonomy will be
treated in the section, "Statistical Discussion." A brief non-
technical description of each method will be presented.

used as input (see Fig. I for areas represented). The
program was halted when four clusters had been
formed and the members of each of the clusters were
recorded as to which of the proposed groups they
represented. A chi-square contingency table was
formed (see Table I ). This tested the null hypothesis
that the grouping originally proposed was completely
independent of a grouping formed by clustering those
lizards of closest morphological similarity.
Step (5).

A two-way multivariate analysis of variance fol-
lowing the methods of Anderson (1958) and
Morrison (1967) was performed on the data. The
following model was used:

Y,^ = U^A,^B.^^e.^

where:

U
A

B:

m

J

ijk

a vector of measurements on an individual
= a vector of effects on Y due to the mean
= a vector of effects on Y due to location
= a vector of effects on Y due to sex

experimental error

The U-statistic and Mahalanobis D-square (An-
derson, 1958) were used to test the null hypothesis of
no difference between groups.
Step (6).

Using Mayr's coefficient of difference as an indi-
cator. 24 characters were chosen that maximized the
variation between groups (Mayr. 1969). These char-
acters are listed in a separate section (entitled Taxon-
omic Characters) which immediately follows this
section.

This step was done for three reasons: (1) to elim-
inate those characters that are relatively invariant
from group to group, (2) to reduce the time involved
in measuring characters on each lizard, and (3) to
keep the matrices used in the analysis within the
limits imposed by computer storage space.

Stepwise multiple discriminant analysis was then
applied to the data to select the set of functions to be
used in placing additional specimens into their proper
group (Dixon, 1968).

Living specimens were examined for coloration
and patternal characters. All specimens so examined
were warmed under 200 watt liglit bulbs for 15-30
minutes before being analyzed. This was done to
approximate the warmth and liglit the animal receives
in nature and to reduce the variability in coloration
and pattern that is the result of internal temperature
variations of these animals.

Color and pattern characters, chosen to represent
all of the observable differences, were combined into

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAPHYTUS COLLARIS

groups, thus expressing a lizard's color and pattern as
a single variable. Aspects of color or pattern that were
invariant or so variable as to present no recognizable
pattern were discarded from the analysis. (The final
list of color-pattern combinations follows the list of
proportional and meristic characters.) The combina-
tions were then tested for independence when com-
pared with geographic locations. Wliere significant,
the color-pattern combinations were used to supple-
ment the discriminant functions in identification.

The remaining specimens were then identified to
determine the extent of the ranges and intergrading
areas of the groups. The probabilities for each individ-
ual to be identified as a member of each of the groups
were calculated. These probabilities, using a modifica-
tion of Rao (1952), were used in outlining the areas
of intergradation.

Taxonomic Characters

The following is a list of the 24 characters finally
selected for measurement on preserved specimens (see
Figs. 1 and 2) as well as the pattern and coloration
characters selected from live specimens (see Fig. 3).
Terms are taken from Smith (1946).
Body measurements.

Snout-vent length, length of second collar, tail
length, and hindleg length were measured to the near-
est tenth of a millimeter. Hindleg length was mea-
sured from the midline to the tip of the fourth toe.
The second collar was measured from the insertion, in
a straight line, to either its dorsal end or to the dorsal
midline if the collar was not disjunct medially. Pro-
portions were then formed from these measurements
and used as the actual characters. The proportions
were tail/snout-vent, tail/hindleg, and the second
collar/snout-vent.
Internasals.

These are the number of scales in a straiglit line
between the middle of each nasal.
Enlarged internasals.

These are the number of scales in the internasal
series which were noticeably larger than the rest.
These scales invariably formed a median row. If the
row began at the anterior or posterior end of the
internasal series, it was also recorded.
Fused interorbitals.

These are the number of interorbitals belonging to
both supraorbital semicircles.
Frontoparietals.

These are scales in the midline anterior to the
interparietal extending anteriorly to the meeting of
supraorbital semicircles.
Head dorsal scales.

These are the number of scales lying in the mid-
dorsal line between the rostral and interparietal
scales.
Loreal-lorilabial series.

These are the number of scales along a straight line
perpendicular to the supralabials running through the
loreals and lorilabials to the junction of the canthals
and suboculars.
Supralabials.

These are the number of scales between the
rostral, but not including it, and the point where the
scales' shape change from rectangular to pentagonal,
with the apex of the pentagon pointing ventrally.
Post mentals in contact with infralabials. These are
recorded as 1 or 0 to correspond with yes or no.
Gulars.

These are the number of scales along a transverse
line connecting the last infralabial on each side.
Dorsal scales.

Three characters were determined wdthin the
dorsals: (1) between the interpariental and the
anterior-most projection of the first collar, (2)
between the anterior-most projection of the first
collar and the posterior border of the second collar,
and (3) total dorsals.
Scales between the collar separations.

Two characters were determined from the collar's
dorsal separation and one character from the pattern
of the first collar: (1 ) the number of scales along a
line connecting the lateral boundaries of the first
collar's separation, (2) the number of scales along a
line connecting the lateral boundaries of the second
collar's separation, and (3) the number of spots, com-
pletely isolated from the main portion of the collar,
witliin the first collar's separation.
Ventrals.

These are the number of scales along a midventral
line connecting the mental and the anterior edge of
the anus.
Subdigital lamellae.

These are the number of lamellae of the second,
fourth and fifth toes on the right hind foot. The
lamellae were considered to begin with the first scale
that was obviously a member of the subdigital lamella
series.
Femoral pores.

These are the number of pores in a straight line on
the right hindleg.

Color-pattern Combinations

Five combinations of color were chosen. In all
cases the color refers to ground color, and variations
in hue around the basic color were considered equal.
The color combinations, now known as color-pattern
types, are (1) body dorsum green, head yellow to sec-
ond collar, gular patch green, area between infralabi-
als and gular patch yellow, (2) body dorsum green,
head yellow to second collar, gular patch green, area
between infralabials and gular patch white, (3) body
dorsum green, head yellow not past eyes, posterior
portion head pale in color, gular patch green, area

between infralabials and gulai patch white, (4) center
body dorsum brown, sides body dorsum green, head
less than half yellow, head posterior cream, gular
patch green, area between infralabials and gular patch

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

white. (5) body dorsum brown, head white or cream,
gular patch brown, black or intermediate, area
between infralabials and gular patch white.

ROSTRAL

INTERNASALS
NASAL

SUPRAORBITAL SEMICIRCLE
(INTERORBITALS)

SUPRAOCULARS
FRONTOPARIETALS

INTERPARIETAL
NUCHALS

r-'ig. 1. A dorsal view of head scalation typical for the populations studied. (Drawn from BYU 21705)

RESULTS

Cluster Analysis

The four groupings formed by this method (see
Fig. 4) were tested to see if they would support the
groupings originally proposed in this study. All indi-
viduals from southeastern Utah and southwestern
Colorado were considered to be C. c. aiiriceps, Ari-
zonan specimens were labeled as C. c. baileyi. south-
ern New Mexican and Mexican specimens were
assigned to the Chihuahuan Desert population. Other
specimens from Colorado. New Mexico, Texas, and
Oklahoma were put in the C. C. collahs category.*

A contingency table was prepared comparing the
groupings as proposed initially by the study (Upper
Colorado River Basin, Central Arizona Plateau. Chi-

huahuan Desert, and Great Plains) with the four
groupings formed by the cluster analysis (see Table
I). The null hypothesis (the two classifications, one
by closest morphological resemblance and the other
by geographical location, are completely independent
of each other) was tested by a chi-square of nine de-
grees of freedom. The lest statistic is significant at the
0.001 level.

*Twenty-five specimens from western Utah, Idaho.
Nevada. California, and Baja California were also used in this
cluster analysis. These specimens clustered together with no
exceptions and remained separate until coefficient of approx-
imately .^0.0 was reached. This is taken as evidence that the
Collared Lizards found west of the Colorado River are very
dilTercnt from those to the cast.

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAPHYTUS COLLARIS

ROSTRAL

MENTAL

POSTMENTAL

CANTHALS

SUPRALABIALS

INFRALABIALS

GULARS
SUBOCULARS

EAR

Fig. 2. A lateral view of head scalation typical for the population.-; studied. (Drawn from BYU 21705)

X-^(l-a.9) = 80.768
X~ (0.999 ,9) =29.7

U-statistic (Anderson, 1958). The U-statistic was
determined to be U,^^ :, -75. = 0.0209. Since most
U-statistic tables only go up to p=10, Paul Sampson's

Table 1. A contingency table testing the indepen-
dence of Ward's clustering method and the proposed
grt)ups.

Therefore X" (l-a,9) ^ '(^"'(0.999.9) 3"*^ the null
hypothesis is rejected. An analysis of the dependence
pattern is as follows: The morphological relationships
of the lizards examined form essentially the same
groupings as those proposed at the onset of this
study. (Ostle, 1963).

Analysis of Variance

One of the multivariate generalizations of the anal-
ysis of variance tests its hypothesis by means of the

Proposed groups

Clusters

Upper

Central

Chihuahuan

Great

Colorado

Arizona

Desert

Plains

1

11

4

1

4

2

5

13

0

3

3

1

2

17

1

4

3

1

2

12

BKICHAM YOUNG UNIVERSITY SCIENCE BULLETIN

>^^ YI LLOW ^5^^ (iRHEN

UROWN

WHITE

Fig. 3. Color-pattern characters: (A) type one, (B» type two, (O type tliree, (D) type five, and (E) type tour.

BIOLOGICAL SERIES. VOL. 1 J, NO. 2 CROTAPHYTUS COLLARIS

10

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

MOHAVLCO, ARIZ

COCONINO CO.. \KI/

PIM\ CO. ARIZ

1 W \l'\l CO . ARIZ.

PIMXCO. ARIZ

N W AJOCO . ARIZ

SAN JUAN CO.. Ul AM
C;R\NI) to . UTAH
MOHAVtCO. ARIZ
YAVAPM CO. \KIZ
COCONINO CO . \RIZ
Ml SA CO. COLO
COCONINO CO . \RIZ
CRAHAM CO . ARIZ
CH WLSCO . NIAV Ml \
WILSON CO . KANSAS
SAN Jl.lAN CO.. IITAH
LUNA CO . NLW MtX
CHAVLSCO. NtWMHX
(JUAY CO , NLW MLX

COCONINO CO. ARIZ
COCONINO CO . ARIZ
SAN J I) AN CO.. UTAH
GARZA CO . TLXAS
LINC OLN CO . NLW MLX
RANDALL CO. TEXAS
COCONINO CO. ARIZ
SAN MICULLCO . COLO
SAN MIC.ULLCO . COLO
CJRAHAM CO . ARIZ
SAN JUAN CO. UTAH
SAN JUAN CO. UTAH
SAN JUAN CO . UTAH
APACHL CO . ARIZ
SAN JUAN CO. UTAH
GRAND CO^UTAH
SAN MIGUEL CO. COLO
SAN JUAN CO. UTAH
SAN MIGULLC O.COLO
SA1\JUAN CO.. UTAH
SAN JUAN CO.. UTAH

APACHE CO. ARIZ
GRAND CO.. UTAH
SAN MIGUEL CO .COLO
DONA AN A CO . NEW MIX
CHIHU \HliA. MEXICO
CHIHILAHUA. MEXICO
CHIHUAHUA. MEXICO
LAS ANIMAS CO COLO
EL PASO CO TEXAS
CHIHUAHUA. MEXICO
EL PASO CO . TEXAS
CHIHUAHUA. MEXICO
CHIHUAHUA. MEXK O
EL PASO CO . TEXAS
LUNACO . NEW MEXICO
SIERRA CO . NEW MLX
CHIHUAHUA. MEXICO
LUNA CO . NEW MLXKO
CHIHUAHUA. MEXICO
DONA \NA (O . NEW Ml X
EL PASOC C) . lEXAS

NAVAJO CO. ARIZ
MOHAVE CO . ARIZ
SAN JUAN CO . Ul AH
C\RU R CO OKLAHOM \
MONICOMI RV CO , KAN
GRAHAM CO . ARIZ
WIISON C O , KANSAS
I MIHI MIL \ Ml \l( C)
CIIIIU MIUA. MEXICO
WIISON CO. KANSAS
ROm RISCO. TEXAS
RAM) \LLCO . TLXAS
WILSON CO. KANSAS
Sll PHANSCO . EEXAS
CAREER CO. OKLAHOM A
LAS ANIMAS CO. COLO
RANDALL CO . TEX \S
LAS ANIMAS CO.COIC)

Fig. 4. Results c)l' Ward's clustering metliod: (I) auric

cps, (11) haileyi. (Ill) Chilmaliua, (IV) coltaris.

BIOLOGICAL SERIES. VOL. 13, NO. 2 CROTAPHY lUS COLLARIS

11

approximate F-value (Di.xon, 1968) was used instead
of the U-statistic. The approximate F-value is

'(la,72.266.84)

= 80.2955. The tabular F-value is

F(0.999,50,I20) ~ --01 (Ostle. 1963). Therefore.
the approximate F-value is signitlcant at tiie 0.001
level and tiie null iiypothesis of equal group means is
rejected.

The Maiialanobis D-square statistic, which is
approximated by the chi-square distribution, also
tests the hypothesis of equal group means. This statis-
tic has degrees of freedom equal to the number of
variables being measured times the number of groups
minus one. The value of the D-square statistic is
D-(l-a,198) = 571.160. The tabular chi-square value
is X-(0.999.100) = 153.2 (Ostle, 1963). The D-square
statistic is also significant at the 0.00 1 level. These
statistics give two of the appropriate multivariate
methods for determining if the proposed groups are
the same or different and therefore distinguishable
from each other.

Coloration and Pattern

ico; and Ciiihuahua, Mexico, and (4) the Great Plains
population, C. c. collahs (N = 12), represented by
those specimens found in Kansas and Oklahoma.*
Only specimens from geographic areas not thought to
contain any intergrading populations were used to
form the discriminant functions. Ail other specimens
were lumped into an unclassified group to be evaluat-
ed by the functions. This is known as model I.

The originally proposed groups were used as data
to examine the effect that the change in groups
would have upon the program's ability to identify the
individuals correctly. The original groups were de-
fined as follows: Upper Colorado River Basin popula-

Table 2. A contingency table testing the indepen-
dence of coloration-pattern types and geographical
locations.

Coloration-pattern types

Locations

It was observed that the coloration and pattern
groupings did not quite reproduce the groupings first
proposed. Therefore, a contingency table, testing for
independence of coloration and pattern with regard
to geographic location was constructed. The test sta-
tistic was found to be significant at the 0.001 level.

X-(l-a.20)=(Oij-Eij)2/Eij

X- (1-Q,20)= 148.136

7

'^"'(.999,20)

47.5

Therefore X"^ x"( ggq iq) ^'""^ the null hypothesis is
rejected. It was concluded that there is a definite
association between the color-pattern and geographic
location (see Table 2).

Discriminate Analysis

The above results were used in altering the pro-
posed groups sliglitly to see if the population in ques-
tion could be more closely defined. The UCLA
BM0D7 stepwise discriminant analysis program
(Dixon, 1968) was run using the following groups as
data: ( I ) Upper Colorado River Basin population, C.
c. aitriceps (N = 10), consisting of those individuals
found north of the union of the Colorado and Green
rivers at the approximate location of Moab, Grand
Co.. Utah, (2) the central Arizona plateau population.
C. c. haileyi (N = 30), containing those individuals
found in Coconino and Yavapai counties, Arizona,
(3) the Chihuahuan Desert population (N = 45), con-
sisting of those individuals found in El Paso Co.,
Texas; Luana, Sierra and Hildago counties. New Mex-

Upper Colorado River Basin
of Utah (north of the
union of the Green and
Colorado rivers) 7 0 0 0 0

Upper Colorado River Basin
of Utah (south of the
union of the Green and
Colorado rivers) 0 2 0 0 0

Upper Colorado River Basin

of New Mexico 14 0 0 0

Upper Colorado River Basin
of Arizona ( east and
north of the Little
Colorado River) 0 13 0 0

Central Arizona (west of
the Little Colorado
River and north of the
MogollonRim) 0 0 17 0 0

Southern Arizona (south

of the Mogollon Rim) 0 0 2 3 0

Chihuahuan Desert of New
Mexico (southern Luna
County) 0 0 0 0 7

*The members of Group (4) represent specimens from a
close proximity of the type locality for C. c. collaris. They
were included to demonstrate the presence of a difference
between the type population of C. c. collaris and the popula-
tions of real concern in this study.

12

HRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

tion, C. c. auriceps; those individuals found in the
Colorado River drainage of Colorado and Utah. All
other groups remained the same. This is known as
model II.

The classification formed by the first discriminant
analysis was chosen as the one that best represented
the actual populations of lizards. The determination
was made by selecting the model that made fewer
wrong classifications of lizards previously defined as
C. c. auriceps. Model I identified 9 out of 10 cor-
rectly or 90%. Model II identified 24 out of 32 cor-
rectly or 75%.

The analysis of the output of the stepwise discrim-
inant program reveals two interesting statistics, the
U-statistic and the F-statistic. The U-statistic tests
equality of means between groups using the variables
included in the discriminant function. The U-statistic
is 0.06346 with 24,3,93 degrees of freedom. The
F-value approximation to the U-statistic was used
because of the availability of F-tables. The approxi-
mate F-value is 4.42269 with 72,210.06 degrees of
freedom. This statistical test determines whether the
groups are, or are not statistically separable when
given a set of taxonomic characters to be used for
classificatory purposes. The tabular F-value is
F(. 999, 100, 120) = 1.82. Therefore F(1-q;,72,2 10.06)
^f^(.999j00,120) 3nd the null hypothesis is rejected.

The F-statistic is used to test the difference
between each pair of groups, thus making it possible
to determine if all groups are separate from each
other. It is measured with 24.70 degrees of freedom,
and all groups are separated at the 0.001 level. The
F-statistic is summarized in Table 3, and the discrimi-
nant functions formed are listed in Table 4.

The taxonomic characters that correspond with
the coefficients of the discriminant functions are
listed as follows: (1) tail length/hindleg length, (2)
tail length/snout-vent length, (3) snout-vent length,
(4) internasal scales, (5) number of fused interorbitai
scales, (6) frontoparietal scales, (7) scales from tiie
union of the posterior canthal and subocular to the
supralabial, (8) infralabial contact with postmetal, (9)
supralabial scales, (10) gular scale rows, (11) number
of enlarged internasals, (12) scales from rostral to
interparietal, (13) dorsal scales from interparietal to

Table 3. A summary of the F-statistics which show
differences between individual groups.

Table 4. A listing of the coefficients of the discrimi-
nant functions.

Groups

Groups

auriceps
bailcvi

Chiluialuia

haileyi

2.99625

2.29124

2.65349

Groups

Variable

auriceps

Chihuahua

bailey i

1

51.39813

45.22476

47.94501

2

2.14172

8.31362

5.60604

3

1.27552

1.34056

1.21061

4

10.33790

10.90252

11.58937

5

15.42395

16.15672

15.77870

6

-13.55824

-13.54851

-13.42835

7

7.06088

4.44547

5.51994

8

9.06605

9.79720

9.35053

9

0.56730

-0.59190

0.83744[

10

1.28302

1.24001

1.18661

11

-5.04772

-3.78518

-4.35537

12

1.2393

0.81388

0.85222

13

0.85929

0.99943

0.73786

14

1.63115

1.57809

1.54830

15

0.41538

0.39279

0.56490

16

0.80368

0.81625

0.71332

17

0.70005

0.44894

0.49637

18

0.01570

-0.01466

0.04405

19

1.38450

1.95044

1.19760

20

269.06104

264.62378

294.33545

21

2.11617

2.84886

3.45656

22

1.17804

1.35396

1.12000

23

0.96154

0.98708

1.14926

24

2.53244

2.28217

2.30347

constant

-526.26660

-523.28274

-527.89990

the anterior edge of the first collar, (14) dorsal scales
from the anterior edge of the first collar to the pos-
terior edge of the second collar, (15) total dorsal
scales, (16) total ventral scales, (17) dorsal separation
of the first collar, (18) dorsal separation of the sec-
ond collar, (19) number of spots within the dorsal
separation of the first collar, (20) second collar
length/snout-vent length, (21) subdigital lamellae of
the riglit iiind foot, second toe, (22) fourth toe sub-
digital lamellae, (23) fifth toe subdigital lamellae,
(24) femoral pores.

The measurements on each lizard were put into all
three functions and a numerical value for that lizard,
as evaluated by each of the functions, was obtained.
Identification was made by placing the lizard into the
group whose function resulted in the largest numeri-
cal value. Along with placing each individual into a
group, the probability thai it belonged in that group
as well as the probability of it belonging to each of
the other groups was calculated. This probability indi-
cated the assurance with which each individual was
classified.

Two aspects of the discriminant analysis" identifi-
cation were considered. First, the degree of reliability
of the identification was examined. Lizards that were

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAFHYTUS COLLAKIS

13

from locations known to be within the ranges of one
of the groups, were examined for their reaction to
being identified by the discriminant functions. Areas
considered definitely to belong to one of the groups
are as follows: the Upper Colorado River Basin popu-
lation, C. c. aiiriceps, Grand Co., Utah and Mesa Co.,
Colorado; the Central Arizona Plateau population, C.
c. baileyi, Yavapai and Mohave counties, Arizona; the
Chihuahuan Desert population, Hildago, Luna, and
Dona Ana counties of New Mexico and Chihuahua,
Mexico. In Yavapai and Mohave counties, only those
Collared Lizards not resembling the Western form of
Collared Lizard were considered. The percentage of
individuals classified correctly was determined by
dividing the number correctly classified by the total
number in the geographic area considered (Ns.corr./
Nj , ). This was summed over all groups to get the
total percentage of correctly classified individuals in
the sample (see Table 5). Thus, approximately 80% of
the sample was identified correctly; wliich is well
within the bounds set by Mayr and others in the 75%
rule(Mayr, 1969).

Table 5. Percentage of sample identified correctly
using tiie discriminant analysis.

Group

Sample

Ncorr.

Percentage

size

Ntot.

auriceps

10

9/10

90.00

bailcvi

42

34/42

80.95

Chihuahua

93

75/93

80.64

Total

145

118/145

81.38

Another aspect of the discriminant analysis dealt
with using the discriminant functions' identification
and the probability for group membersliip to investi-
gate intergradation between populations. This has

always been a problem that made identification of
Collared Lizards difficult (Burt, 1928).

The individual, when identified by the discrimi-
nant function program as belonging to a group, is
labeled with the a posteriori probability for its mem-
bership in all of the groups. The a posteriori probabil-
ity is the probability of an individual belonging to a
group once the group has been defined. This set of
probabilities always sums to unity. The probabilities
for membership in a group have three options: (1)
There will be one large probability and the rest small
(e.g., 0.982 and 0.044 and 0.044). In this situation,
the individual is placed in the group with the largest
probability of membership. Any probability that
exceeds 0.70 is considered a large probability of
membership. (2) There will be two approximately
equal probabilities and the rest small (e.g., 0.5602
and 0.4498). This indicates that the individual is not
distinct enougli to fit with much assurance into either
group. A specimen of this type is considered to repre-
sent an intergrade between the groups given the larg-
est probabilities. (3) All the probabilities will be
approximately equal. In this case, the individual is
assumed to be unidentifiable. This aspect of discrimi-
nant analysis is an adaption of Rao's three population
discriminant analysis procedure (Rao, 1952). Identifi-
cation of each specimen used in the study was exam-
ined. The percentage of the total sample placed in
each group was recorded. This percentage was
arranged by geograpliic area in a north-south line. The
results of this analysis are summarized in Figs. 5 and
6.

The characters themselves were examined to estab-
lish their effect on the discrimination between
groups. This is summarized in Table 6. Table 7 lists
the means and standard deviations for the 24 charac-
ters of each group. Figure 7 also shows the means and
one standard deviation for the characters that give the
best individual discrimination between groups.

DISCUSSION

Zoological Discussion

The initial question proposed by a taxonomic
study at the subspecific level deals with identifica-
tion. Specifically, are there any populations that are
geographically continuous and also identifiable, fol-
lowing the 75% rule, in respect to other populations?

The multivariate analysis of variance shows that
the populations proposed in this study are distin-
guishable at a high confidence level. This analysis was
done using external morphological characters exclu-
sively. The groupings of the cluster analysis, along
with its test of independence, lends support to the
actual existence of the proposed groups.

Once assured of the existence of these popula-

tions, the problem becomes one of identification.
Using external morphological characters exclusively,
an examination of the means and standard deviations
reveals that although their means were different (the
statistical tests of population difference showed this),
their overlap was such that no one character could be
used to identify an unknown specimen with complete
accuracy. Discriminant analysis computes a new char-
acter, "Z," which is the value of a set of functions or
equations. These functions are constructed from
linear combinations of the original characters in such
a way that as many members as possible from each
population have higli values for the function that cor-
responds to their population. In a sense, this is a new
taxonomic character that identifies members of each

14 hrk;ham young univeksity science bulletin

A key to the abbn;vialions used in I'igs. 5 and 6.

atiriceps " The Collared Lizard population ot the Upper Collarado River Basin, north of the union of the
Colorado and (Ireen Rivers.

baiteyi - The Collared Lizard population of central Arizona.

AxB - The intergrade popidation o( aiiriccps and haileyi.

chihuahua ■ Lhe Collared Lizard population of (he Chihuahuan Desert.

B\C' - The inlergradc population of /)u//t'i7' and Chihuahua.

LInident ■ Individuals not assignable to any of the above groups.

Upper Colorado River Basin
(north of the union of the Green and Colorado rivers)

Percent of
sample

100-
80-
60-
40-
20-

Auriceps AxB Baileyi BxC Chihuahua Unident.

Fig. 5. A comparison of Collared Lizard populations along a line connecting Grand Co., Utah and northern Sonora. Mexico.

Upper Colorado River Basin of Utah and Colorado

(between the union of the Green and Colorado rivers

and Monticello. Utah)

Percent of
sample

Auriceps AxB Baileyi BxC Chihuahua Unident.

I'ig. 5. (continued)

BIOLOGICAL SERIES. VOL. 1 J. NO. 2 CROTAFHYTUS COLLARIS

Upper Colorado River Basin of Utah and Colorado
(south of Moiiticello, tlie San Juan River drainage)

15

100-
80-
Percent of ^0-
saniple

Auriceps AxB Bailey i

BxC Chihuahua Unident.

Upper Colorado River Basin of Arizona
(east of the Little Colorado River)

Percent of
sample

100-
80-
60-
40-
20-
0

■ 1 ■

Auriceps AxB Baileyi

BxC Chihuahua Uindent.

Central Arizona
(between the Little Colorado River and the Mogollon Rim)

Percent of
sample

100-

80-

■

60-

1

40-

1

20-

1

0

^

■

—

Auriceps

AxB Baileyi

Fig. 5 continued

BxC

Chihuahua Uin

16

BKIC.HAM YOUNG UNIVERSITY SCIENCE BULLETIN

Gila County, Arizona
(the base of the Mogollon Rim)

100-

80-

Percent of
sample

60-
40-

1

20-
0

I 1

1

Auriceps AxB Baileyi BxC Chihuahua Unident.

Pinal and Graham Counties, Arizona
(mountains south of the Mogollon Rim)

Percent of
sample

100-
80-
60-
40-
20-
0

J

Auriceps AxB Baileyi BxC Chihuahua Unident.

Percent of
sample

Sonoran Desert of Arizona and Mexico
(Cochise, Pima, and Santa Cruz counties and Sonora, Mexico)

100-

80-
60-
40-
20-
0

■ ■

Auriceps AxB Baileyi BxC Chihuahua Unident.

fig. 5 conlimicd

BIOLOGICAL SERIES. VOL. 1 3. NO. 2 CROTAPHYTUS COLLARIS

17

(Upper Colorado River Basin of New Mexico
(the San Juan River drainage of Rio Arriba County)

Percent of
sample

100-
80-
60-
40-
20-
0

Auriceps

AxB Baileyi

BxC

Chihuahua

Unident.

Percent of
sample

Upper Colorado River Basin of New Mexico

(the southern portion located in

McKineley and Santa Fe Counties)

00-

80-

60-

40-

■

20-
0

■

1

1

1

Auriceps

AxB

Baileyi

BxC

Chihuahua

Unident.

Percent of
sample

Bernallilo and Torrance counties. New Mexico
(mountains south of the edge of the Upper Colorado Plateau)

100-
80-
60-
40-
20-
0

1

Auriceps

AxB Baileyi

BxC Chihuahua

Unident.

Fig. 6. A comparison of Collared Lizard populations along a line connecting Rio .Arriba County, New Mexico and Chiliuahua,
Me.xico.

18 I5RIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

Central Rio Grande River Valley
(Lincoln and Socorro counties. New Mexico)

100-

80-

Perccnt of

, 60-

saniple

40-

20-

0

1

Auriceps AxB Baileyi BxC Chihuahua Unident.

Southern portion of New Mexico's Rio Grande River Valley
(Sierra and Otero counties)

100-
80-

Percent of ,^
sample

40-

20-

0

l_L

Chihuahuan Desert of New Mexico
(Dona Ana, Hildago and Luna counties. New Mexico)

100-
80-
Percent of ^n

60-
sample

40-
20-

0

Auriceps AxB Baileyi BxC Chihuahua Unident.

Auriceps AxB Baileyi BxC Chihuahua Unident.

lig. 6. (continued)

BIOLOGICAL SERIES. VOL. 13. NO. 2 CROTAPHYTUS COLL.^RIS

Percent of
sample

100-

80-
60-
40-
20-
0

Chihuahuan Desert of Mexico
(Chihuahua, Coahuila and Nuevo Leon, Mexico)

1

Auriceps

AxB Baileyi

BxC

Chihuahua

Unident.

group with a higli degree of reliability. This rehabihty
is expressed in the form of a probability for joining
each group (Sokal and Rolilf, 1969). The discrimi-
nant functions have demonstrated their ability to
identify the three populations accurately.

Coloration was studied separately from the mor-
phological characters. This aspect of a lizard's pheno-
type consistently demonstrated differences between
populations. There may be some questions as to the
validity of coloration when used as a taxonomic char-
acter due to a possible relationship between colora-
tion and diet. However, the similarity of classifica-
tions of Collared Lizards by color patterns and by
morphological characters tends to dispute this idea.
In an animal such as the Collared Lizard, that is
active, diurnal, higlily territorial, and uses sight to rec-
ognize both territorial intruders and potential mates
(Fitch, 1956), it is difficult to imagine a character
more important than color pattern. Since coloration
fades rapidly on preserved specimens, tliis method of
identification works on live specimens only. The pop-
ulation of C. c. auriceps is characterized by colora-
tion-patternal type \\C.c. baileyi. by type 3; inter-
grades between the two, by type 2; the Chihuahuan
Desert population, by type 5; and intergrades
between the Chihuahuan Desert population and C. c.
baileyi. by type 4.

By combining the information obtained from both
coloration-patternal and morphological characters, it
is possible to explain the structure of the Collared
Lizard populations between the Rio Grande and
Colorado Rivers (see Fig. 8). Located in the northern
extension of the Upper Colorado River Basin is C. c.
auriceps. its range should now be restricted on the
south, to the region of the union of the Green and
Colorado rivers in the vicinity of Moab, Grand
County, Utah. Northward, they extend at least to the
Book Cliffs area and possibly further. South of the

Table 6. A comparison of the percent of correctly
identified individuals within groups as new variables
are added to the discriminant functions.

Variable

Groups

Step

added

auriceps

baileyi

Chihuahua

coUaris

1

5

0

90

16

75

■)

8

40

43

47

92

3

7

80

50

60

83

4

21

80

60

69

92

5

10

80

63

69

92

6

14

80

67

73

92

7

4

80

60

73

100

8

17

80

70

73

100

t)

18

80

67

71

100

10

3

80

73

73

100

12

19

80

73

78

100

13

20

80

80

84

100

14

13

80

77

89

92

15

15

80

70

88

100

16

12

80

70

88

100

17

-)T

80

73

88

100

18

-> 1

80

77

89

92

19

U

90

77

89

92

20

~t

90

77

92

92

24

h

90

77

92

100

union of the Green and Colorado rivers, an increasing
number of individuals is identified by the discrimi-
nant functions as C. c. auriceps X C. c. baileyi inter-
grades. This corresponds with a decreasing number
identified as C. c. auriceps. which is also supported by
the color patterns observed. In this region, numerous
lizards are found of color type 2, wliich is intermedi-
ate to color type 1, C. c. auriceps: and color type 3.
C. c. baileyi. The area of intergradation is limited to
the region somewhat south of Moab, Utah, and
extending to the vicinity of the Little Colorado River
of Arizona.

Crotaphytus collaris baileyi is now restricted to
the region south of the Painted Desert, across the
center of Arizona. It is also expected to occur in the

20

BRIGHAM YOUNC; UNIVKRSITY SCIENCE BULLETIN

o.n 0.25 -SO 0.75 II) I :.5 i so its :u : 25 2 so

B
CH

1.(1 'IS 10. h 114 122 l.iU l.l.S M.(i IS.-) 16.2 17.0
lllll

?n 7 5 so S5 'M) 9 5 10 0

B
CH

15 0 1 5 (, I (. 2

1 (. N 17 4

l-"ig. 7. Means and standard deviations plotted tor the lour
characters that contribute the most to population discrimina-
tion: (I) fused interorbitals, (li) supralabials, (Hi) loreal-
lorilabial series. (IV) subdigital lamellae of second toe on
hind foot, (("O) Great Plains population, (CI I) Chiluiahuan
Desert population, (B) central Arizona population. (A) Upper
Colorado River Basin population.

western central mountains of New Mexico; however,
specimens from this area were too few to state this
with assurance. C. c. hailcyi seems to be centered in
Mohave, Yavapai, and southern Coconino counties of
Arizona and follows the Mogollon Rim and adjacent
mountains to the east. In the south, the picture

becomes more confused. Isolated populations of C. c.
baileyi are found on the mountain tops, and, in the
lower elevations, intergrades between C. c. baileyi and
the Chihuahuan population are found. In general, this
area is populated by C. c. hailcyi X Chiluiahuan inter-
grades; however, the exact relalit)nships are in need of
further study.

The exact type locality of C c. baileyi is unknown.
Stejneger describes it as the Painted Desert of the
Little Colorado River. In his original description of
baileyi, two facts become apparent. First, the red
spots described on the neck of the type (which is a
male), and the time of collection (late August, when
generally only juveniles and hatchlings are active)
indicate the type to be a juvenile. Secondly, the type
locality is somewhere between Cameron and Wupatki
National Monument in Coconino County, Arizona.
Stejneger describes the locality for collection of the
type as in the vicinity of the Little Colorado River.
The type was collected on the second excursion to
the desert, which took him north from Flagstaff to
Tuba City on a route that is followed by U.S. High-
way 89 (Stejneger, 1890). Locating the type locality
as south and west of the Little Colorado River places
it within the range of C. c. baileyi as determined by
this study.

The population centered in Chihuahua should now
be recognized as a subspecies and separate from C. c.
baileyi and C. c. auriceps. This is done on the basis of
its overall morphological distinctiveness as expressed
by the discriminant functions and also by its strik-
ingly different coloration. The brown of its doral
coloration is easily distinguished from the green of
either C. c. baileyi or C. c. auriceps. The new popula-
tion, centered in Chihuahua, Mexico, extends north-
ward to Socorro County, New Mexico, dispersing up
the lowlands of the Rio Grande River Valley. The
eastern and southern extents of its range are in need
of further study. To the west is an area where isolated
populations of C. c. baileyi occur on the higher
mountain ranges (e.g., Tanciue Verde Mountains near
Tucson). This provides for intergradation to occur
and for the coloration-pattern type 4 to become com-
mon. The overall ranges are shown for the three pop-
ulations in Fig. 8.

Twelve individuals from the area of the type local-
ity of C. c. collaris were included in the discriminant
analysis. This was done to obtain an idea as to the
relatitniship of C. c. collaris to the pi>pulalions
studied. Also included in the analysis were a number
of individuals from eastern New Mexico, Texas, and
eastern Colorado. All are within the presently deter-
mined range of C. c. collaris. Two facts resulted from
this analysis: ( 1 ) the lizards from the type locality
were identified with 100% assurance, and (2) no
meaningful pattern could be discerned for the remain-
ing supposed C. c. collaris. This suggests that the sub-
species, C. c. collaris, is also a heterogeneous grouping

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAPHYTUS COLLARIS

21

Table 7. A listing of the means and standard deviations for the 24 characters measured on the individuals used
in forming the discriminant functions.

Groups

auriceps

baileyi

Chihuahua

collaris

Character

Mean

Stand, dev.

Mean

Stand, dcv. Mean

Stand, dev.

Mean

Stand, dev,

1

2.2

0.3

2.2

0.2

2.2

0.2

2.1

0.1

2

1.8

0.2

1.9

0.1

1.9

0.2

1.8

0.1

3

94.3

7.2

93.1

6.2

95.3

7.8

91.8

7.8

4

5.8

0.8

6.0

0.7

6.0

0.7

4.8

0.8

5

0.1

0.3

0.1

0.3

0.4

0.7

1.8

0.8

6

2.3

0.8

2.0

0.7

2.0

0.9

1.2

0.4

7

8.6

1.5

7.3

0.9

6.7

0.9

7.4

1.2

8

12.5

0.7

13.2

1.6

13.9

1.6

10.5

0.8

9

0.9

0.3

1.0

0.2

0.9

0.2

0.8

0.4

10

66.1

4.5

61.0

6.2

62.8

6.0

52.8

6.1

11

0.1

0.3

0.5

0.7

0.6

0.7

0.1

0.3

12

15.7

1.2

14.7

1.7

14.4

2.7

14.7

1.0

13

24.5

6.1

25.4

7.3

28.6

3.9

26.2

2.6

14

33.7

6.5

33.1

5.8

28.4

4.3

24.5

3.9

15

158.1

7.9

163.5

11.7

155.8

10.4

142.1

9.5

16

195.4

9.8

187.5

11.5

187.7

10.9

170.2

6.2

17

28.9

4.7

24.2

7.8

23.3

5.8

24.2

10.3

18

1.4

1.8

4.3

3.8

5.4

4.3

7.8

6.1

19

1.9

0.3

1.7

0.6

1.9

0.4

1.0

0.3

20

0.2

0.0

0.2

0.0

0.2

0.0

0.1

0.0

21

17.3

1.3

19.5

1.5

18.7

1.4

16.4

1.2

22

33.0

3.9

34.7

3.0

34.4

3.0

28.8

2.1

23

15.0

1.7

15.5

1.5

15.0

1.8

13.9

1.5

24

18.6

1.6

17.8

1.4

18.2

1.9

17.6

1.3

and should be studied. The ease of separation of the
specimens from the area of the type for C. c. collaris
supports the present separation of three western pop-
ulations from an eastern Great Plains group now des-
ignated as C. c. collaris. This was also supported by
the grouping of the cluster analysis. Following Rao's
technique (1952), a graph of the tlrst two canonical
variables was plotted (see Statistical Discussion). This
provides a two-dimensional representation of the
interrelationship of the populations (see Fig. 9).

A diagnosis of the three populations' characteris-
tics and their comparison with the material represent-
ing the type population of C. c. collaris is as follows:

Crotaphytiis collaris auriceps is separated, as are
the other groups, primarily on coloration and pattern.
It has a light green body with a briglit yellow head.
The yellow on the head extends posteriorly to or just
past the second collar and ventrally onto the throat.
In males, the yellow on the throat meets the green of
the gular patch. Morphologically C. c. auriceps is sep-
arated from C. c. collaris by a fewer number of speci-
mens with fused interorbitals. It is separated from C.
c. baileyi primarily by a smaller number of supralabi-
als, as indicated by Fitch and Tanner (1951), and a
greater number in the loreal-lorilabial series. C. c.
auriceps is also distinguishable from the Chihuahuan

Desert population by the above characters and is fur-
ther separated by possessing fewer subdigital lamellae
on the second toe of the hind foot.

Crotaphytus collaris baileyi has a dark green body,
and, if yellow is present on the head, it does not
extend posteriorly beyond a line drawn between the
rear of the supraorbital semicircles. Yellow is never
found on the throat. Using morphological characters
exclusively, C. c. baileyi is separated from C. c. coll-
aris by a smaller number of specimens with fused
interoribitals, a greater number of supralabials, and a
greater number in the loreal-lorilabial series. C. c.
baileyi was not separated with much assurance from
the Chihuahuan Desert population until 14 characters
had been added to the discriminant function. This
suggests the differences between the populations are
expressed as a function of many variables (a sum of
many small differences) rather than just one.

The Chihuahuan Desert population differs from C.
c. collaris morphologically mainly by having fewer
individuals with fused interorbitals and having more
supralabials. Color and patterns of C. c. collaris were
not analyzed. The Chihuahuan Desert population has
been demonstrated to be sufficiently different from
all presently recognized populations to merit designa-
tion at the subspecific level. It is therefore named as

22

A key lo the sumbols used in lig. 8.

Range of the Upper Colorado River
oj Basin population. C c. auriccps.

I -}- I Range of the centriil Arizona
I 1 ij piipuljticin, C. c. hailvvi

HRIC.HAM YOUNG UNIVERSITY SCIENCE BULLETIN

Vl" "xl Range of C. c. baileyi x C. c. auriceps

I o4- ." I intergrades.

Pi •" )> I Range of the Chihuahua Desert

.'■"' ' "* f population, C. c. fuscus.

VF^ "^J Range of C. c. fuscus x C. c. baileyi

I -H-^^ 1 intergrades.

1 ig. 8. A range map for the populations studied.

BIOLOGICAL SERIES. VOL. 1 3. NO. 2 CROTAPHYTUS COLL.'XRIS

23

A kcv to tlie symbols used

A - The upper Colorado River Basin population.
B - The central Arizona population.
F - The Chihuahuan Desert population.
C - The Great Plains population.

■ Misidentified A.

■ Misidentified B.
Misidentified F.

A A

A BB

BB

*
C C

AB
B
B BB

^\

F B B B^B

B $

F F & F

0 F FF
F 0 *F F F

F F F F

F F F

FF F

F F F F

F F ff 0

Fig. 9. A two-dimensional representation of the interpopulation morphological relationships formed from the canonical
variates.

Crotaphytus vollaris fusciis subsp. nov.

Type. - Adult male, Brighain Young University No.
I(i^»70, collected 6.5 mi. N. and 1.5 mi. W. of Chihua-
hua City, Chihuahua, Mexico, by Wilmer W. Tanner
on July 21, 1960.

Paratypes. — Chihuahua: topotypes, BYU 14211,
14212, 15305, 15325-15331. 15817-15822, 16%9,
16971-16977, 17010; Chihuahua City: UC 70704;
Colonia Juarez: BYU 3736, 15185-15188; Hechi-
chero: KU 33789; Nuevo Casas Grandes: BYU
15184; palomas: BYU 17014; Ricardo Magon: BYU
13382-13386, 13410, 1341 1 ; Victoria: KU 33788.
Diagnosis. - It differs from C. c. baileyi. C. c. aiiri-
ceps and C. c. collahs in having a brown dorsal color
with no trace of green and a liglit to cream colored
head with no trace of yellow. The morphological dil-
ferences, no one of which is conclusive, are many and
add up to a general difference from the recognized
populations that is best expressed by the previously
mentioned discriminant functions.
Description of the type. ~ Head and body length 106
mm. tail length 222 mm, width of head at angle of

jaw 28 mm, hindleg length from midline to tip of
fourth toe 89 mm, femoral pores 20-20, supralabials
14-14, infralabials 12-11, fused interorbitals 0, inter-
nasals 6, frontoparietals 2, loreal-lorilabial series 7,
postmentals in contact with infralabials, gular scale
rows at angle of jaw 69, scales from rostral to inter-
parietal 15, scales from interparietal to anterior edge
of first collar 25, scales from anterior edge of first
collar to posterior edge of second collar 28, total dor-
sals 140, total ventrals 181, scales within dorsal sepa-
ration of first collar 26, scales within dorsal separa-
tion of second collar 5, number of spots within dorsal
separation of first collar 2, subdigital lamellae of sec-
ond toe of right hind foot 17, subdigital lamellae of
fourth toe 31 , subdigital lamellae of fifth toe 16.

Type described from preserved specimen and nat-
ural coloration not apparent. General pattern and col-
oration as described from living specimens. Head pale
(white or cream) with a few small dark spots distrib-
uted randomly across back of head, gular patch black,
bluish black or dark brown, never green, reticulation
present on lateral edges of gular patch, reticulations
to infralabials, first collar widely disjunct dorsally

24

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

with two small spots in disjunction, second collar nar-
rowly disjunct dorsally, second collar on forearm,
body dorsum of varying shades of brown, never
green; small white spots, rarely any yellow, scattered
profusely; front legs same color as body dorsum but
patternless; hindlegs same as body dorsum with spot-
ting of body continued to thigli; feet pale; body
venter white or cream; body dorsum ground color,
fades into venter in region of midbody; no dark axil-
lary or groin patches present.

The populations represented by C. c. collaris, C. c.
baileyi, C. c. auriceps, and C. c. fuscus should be con-
sidered as an evolutionary group because of their
greater morphological and patternal similarity when
compared with the Collared Lizards west of the
Colorado River. This group will be referred to as the
co//an.v -complex.

The following phylogeny is based on morphologi-
cal and patternal evidence. The population ancestral
to the co//ans-complex originated in east central Mex-
ico, probably Chihuahua or Coahuila, from which
they established themselves in the border states of
Mexico and the United States. With improving cli-
matic conditions, following the Pleistocene, they
advanced northward. The population followed three
corridors of dispersal: ( 1 ) along the low mountains of
southwestern New Mexico and southeastern Arizona,
(2) up the Rio Grande River Valley, and (3) east of
the central mountains of New Mexico.

The first corridor led to the high plateaus of cen-
tral Arizona and the Upper Colorado River Basin.
Upon reaching the elevation barrier presented by the
southern edge of the plateaus, a segment of the ances-
tral population, which invaded the higlier elevations,
was isolated and continued to disperse along the
drainages of the Colorado River and its tributaries.
This population, C. c. baileyi, moved northeast into
Utah and Colorado following the mountain ranges
that skirt the relatively uninhabitable Painted Desert
and Monument Valley region of Arizona. This
resulted in a large population centered in the Upper
Colorado River Basin of Utah and Colorado, loosely
associated with the main population of central
Arizona. The length of the connection, coupled with
the spotty distribution in northeastern Arizona,
reduced the amount of genetic exchange possible
between the two main populations. This allowed a
distinct population, C. c. auriceps, to form at the
northern boundary of the Upper Colorado River Basin.

Crolaphytus collaris auriceps is probably the
youngest population of the co//am -complex. Three
facts support this idea: (1) C. c. auriceps is located in
the area most recently open to expansion by reptilian
forms, the central portion of the Upper Colorado
Eliver Basin. (2) C. c. auriceps presents the smallest
change in color and pattern from C. c. baileyi, thus
suggesting recent evolution from that form, and the
greatest change from the population closest to ances-

tral stock (the Chihuahuan Desert population, C. c.
fuscus). (3) C. c. auriceps still possesses a wide inter-
grade zone with C. c. baileyi, suggesting that they
have had little time to separate.

Crotaphytus collaris fuscus has probably changed
little from the ancestral form, as it occupies essential-
ly the same range. As conditions improved, following
the Pleistocene, C. c. fuscus dispersed northward fol-
lowing the Rio Grande River along the low mountain
ranges and river basins. It appears to be contained by
the higlier elevations encountered to the north and
west. The range of C. c. fuscus may be outlined by
constructing a line following the 5,000 foot elevation
level in southern Arizona and New Mexico.

The population that dispersed eastward presently
inhabits the Great Plains region. It became isolated
from the western segment of the aj/tom-complex by
the Rocky Mountains and differentiated into a dis-
tinct population now considered to be C. c. collaris
(see Fig. 10).

Statistical Discussion

One of the major difficulties in using statistics in
the zoological sciences is the lack of descriptions of
the techniques phrased in the language of a zoologist.
The purpose of this section is twofold: ( 1 ) to explain
the necessity of using multivariate statistics in the
taxonomy of subspecies, and (2) to explain in under-
standable terms the statistical methods used in this
paper.

One of the more important discoveries of this
study has been in the realm of methodology. Univari-
ate methods are those techniques of data analysis and
statistical decision-making where only one variable is
measured on each experimental unit. Examples of
this type of analysis are T-test of means, Chi-square,
and Analysis of Variance. In recent years, a new sta-
tistical technique has grown out of classical univariate
analysis. This form of statistics, the multivariate or
generalized analysis, should be recognized as the
proper form to use in taxonomic studies where more
than one character is being analyzed. Multivariate
analysis is designed to perform analyses that are anal-
ogous to those of univariate methods for cases where
more than one measurement, or variate, is being
determined on each experimental unit or specimen
(Anderson, 1958).

There are definite hazards to using univariate
methods where multiple measurements are being
made on a single experiinental unit. These hazards are
centered in the inability of knowing the exact alpha-
level of a statistical test, unless the assumptions of
that test are complied with. The alpha-level of a test
is the probability that the difference observed in the
data is due to chance.

In most herpetological taxonomic studies, more
than one variate or taxonomic character is measured

BIOLOGICAL SERIES. VOL. 1 3, NO. 2 CROTAPHYTUS COLLARIS
AURICEPS

25

Fig. 10. Proposed phylogenetic relationship of the populations of the c'o//ora-comple\.

on each individual. These variates usually are then
analyzed separately with a univariate method, and the
results are combined to support a conclusion.

If such multiple measuretnents are analyzed on the
basis of separate univariate treatments of the vari-
ables, the combination of the results of the univariate
tests and the assignment of a level of confidence to
any inference drawn from these tests present a prob-
lem. If all the variables are perfectly correlated, the
same conclusion is drawn from each variable, and uni-
variate methods are acceptable. Perfect correlation
means that all the variables change values in the same
relationship to each other (e.g.. for each five scale in-
crease of the dorsals, the femoral pores increased
one). However, variables that are perfectly correlated
are each measuring the same source of variation. To
prevent heavy weighting of that source of variation.
only one of the correlates should be measured. If the
variables are completely independent and significance
at the 0.10 level is claimed (when at least one variable
shows significance), the true level of significance is
l-{.90n) with ^ equal to the number of variables
measured (Steel. 1955). Suppose four different vari-
ables are measured and tested at an alpha-level of
0.10. Using the above formula, it can be seen that the
true alpha-level of any conclusion drawn from the
combination of the four tests is actually l-(.90 ) =
l-{.66) = 0.34. This differs greatly from the alpha-
level of 0.10 that would normally be assuined. If the
rule would be to claim significance when all the vari-
ables show significance, the alpha-level would become
(O.IO)", with " equal to the number of variables

measured. Using the previous example, the true
alpha-level becomes 0.0001. This makes it practically
impossible ever to detect a difference. Often indepen-
dence of variables is assutiied without proof. For a
variable to be considered completely independent, its
value must not be influenced by the value of any
other of the measured variables. In taxonomy, which
deals with characters controlled by an unknown
arrangeirient of the genotype, the assumption of in-
dependence of variables without prior verification
seems untenable.

If either complete dependence or independence of
variables were known to be the case, rules could be
formulated to allow inferences from a combination of
univariate analyses of the data. However, the true sit-
uation invariably lies somewhere between the two
extremes. Thus, one would not know the true level of
significance of the inferences on the combined results
of univariate analyses. By using the multivariate gen-
eralizations of univariate methods, this problem of
indeterniinant alpha-level is controlled (Steel. 1955).

Mayr (1969) has advocated the use of multivariate
methods wherever multiple measurements are used.
He states also that often the calculations (e.g., the
determinant of a 100 X 100 matrix) are prohibitive.
With the advent of fast digital computers and pack-
aged programs, this is no longer true.

Another more compelling reason for using multi-
variate analysis of data concerns what is actually
being analyzed. Taxonomists are classifying whole
organisms, not any one scale count (Mayr, 1969;
Sokal and Sneath, 1963). Univariate methods con-

26

liKlCHAM YOUNC; UNIVI.KSIIY SCIENCE BULLETIN

sider only one variable at a time as completely unre-
lated to all other variables. Multivariate methods con-
sider groups of characters, as a unit, and their rela-
tionships with each other. This is a better approxima-
tion of the organisms with which taxonomists are
concerned. The following is a description of the mul-
tivariate techniques used:

Multivariate analysis of variance. A method lo lest
the difference of group means for those cases where
more than one variable is recorded for each indi-
vidual. This is the multivariate extension of the famil-
iar analysis of variance and F-Test. It is appropriate
for testing hypothesis concerning differences between
populations.

Cluster analysis. When a taxonomic study is made
taking two measurements on each individual, the
specimens studied could be represented as points on a
two-dimensional space. The resulting graph would
illustrate the phenotypic interrelations of the indi-
viduals. Expanding this to 90 measurements on each
individual, the specimens could be represented as
points in a hypothetical 90 (or p)-diniensional hyper-
space. The representation of individuals on a
90-dimensional graph is best grasped by visualizing
many points in space grouped in clusters of varying
size. The number of dimensions in the hyperspace is
equal to the number of variables measured. This
concept of individuals being represented as points in a
p-dimensional space is essential to cluster and discrim-
inant analyses.

Ward's method of cluster analysis forms spherical
clusters of individuals in the hyperspace. New clusters
are formed by measuring the distance from each
individual in the original cluster to the center of the
cluster, called the centroid. These distances are sum-
med to form the error sum of squares for the cluster.
The individuals to be added to the cluster are con-
ditionally added, and the new centroid formed. An
error sum of squares for the newly formed cluster is
calculated. This procedure is done for all possible
entries to the original cluster (possible entries include
other clusters as well as individuals). The entry that
causes the least increase in the error sum of squares is
joined to the original cluster. Each new cluster is
formed by joining those individuals that move the
centroid the smallest distance. In other words, each
cluster is composed of those individuals located
closest to each other in the hyperspace. Thus, it is
seen that this method unites individuals of the highest
morphological similarity first (Wishart, 1969). The
main assumption that must be valid for this proce-
dure to give meaningful results is that the characters
chosen represent the phenotype of the animal as well
as possible.

Canonical analysis. This method allows the examina-
tion of the relationship of two sets of variables. The
two sets used in this study were (1) groups and (2)

variables measured on the individuals. This resulted in
two variables, evaluated for each individual, formed
from a linear function of all the variables measured.
These two new variables maximized the correlation
between groups and originally measured variables.
When plotted on an .v and i' axis, the variables t'orm a
two-dimensional graph of the relationships of the
groups to each t)ther (Dixon, 1^)68; Rao, 1952).
Discriminate analysis. This technique theoretically
constructs p-dimensional planes in the hyperspace,
which separate the clusters of individuals. In practice,
it builds a single variable from all the variables mea-
sured and maximizes the difference between groups
(Anderson, I958;Sokal and Rohlf, 1969).

Rao (1952) describes a "gray" area located be-
tween two clusters, in which a few individuals may
occur. In this area the individual's probability of
belonging to either cluster is not great enough to
grant membership with assurance. Rao states the
possible conclusions in a system consisting of three
clusters. The individuals either belong to (1) one of
the three clusters, (2) one of two clusters, or (.>) all
three clusters and no conclusion may be drawn.

The procedure for using discriminant analysis in a
subspecitlc problem, as developed in this study, par-
allels Rao's concept. The major distinction is because
of the ability of members of iwo distinct clusters
(subspecitlc population) to interbreed and produce
individuals with characteristics intermediate to either
of the clusters. Thus, Rao's "gray" area becomes a
region occupied by intergrades between the two
clusters. As in Rao, an individual with an equal proba-
bility of joining all of the groups is considered
unidentifiable (see Fig. 1 1 ). This procedure, like so
many procedures in taxonomy, is based on certain
subjective decisions. Therefore, the validity of its
results is dependent upon the validity of the assump-
tion made. The assumptions are as follows below:

( 1 ) Among the individuals to be classified, at least
two distinct populations must be represented. Prior
to using discrinunant analysis, an appropriate method
must be employed to determine the number of popu-
lations present. A test of the population difference is
also advisable.

(2) The most crucial assumption concerns the
selection of members used in forming the discrim-
inant functions. In order to identify intergrade popu-
lations correctly, the individuals used to form the
functions must be selected so that only "pure stock"
of the populations being investigated is represented.
The sample used to form the discriminant functions
define, as far as the analysis is concerned, the param-
eters of that population. Thus, as more intergrades
are included in tlic sample, a less precise definition of
the population and its parameters results; and the
identificatit)!! of individuals by the discriminant
analysis declines in reliability.

BIOLOGICAL SERIES. VOL. I 3. NO. 2 CROT.-XPHYTUS COLLARIS

27

SUBSPECIES

A

Probability to Join

A - High

B- Low

C- Low

A \ B INTERGRADES

Probability to Join

A = B

C - Low

UNIDENTIFIABLE

SPECIMENS

Probability to Join

A = B = C

A X C INTERGRADES

Probability to Join

A = C

B - Low

SUBSPECIES B

Probability to Join

A - Low

B - High

C - Low

B.xC

INTERGRADES

Probability to Join

A - Low

B = C

SUBSPECIES C

Probability to Join

A - Low

B - Low

C - High

Fig. 1 1. The subspecifif identification problem as viewed by discriminant analysis (modified after Rao. 1952).

(3) Any one individual to be identified, must be
considered to have an equal prior probability of
belonging to any of the populations. This can be
assured by placing an equal number in each of the
groups used to form the discriminant functions. It is
acceptable to use unequal numbers if specimen avail-
ability or some other factor necessitates. In this
study, availability of pure C. c. auriceps and C. c.
coUaris limited those samples and a decision was
made to sacrifice equality of a priori probabilities to
gain a better definition of the population parameters
of the other two subspecies.

(4) Discriininant analysis is a statistical method,
and, as such, its reliability hinges on the amount of
information put into it. As the number of individuals

and variables used in forming the functions increases,
so does the reliability of the results. This is also true
of the reliability of identification of new specimens.
The results of statistics are always phrased in prob-
abilities and the higlier the probability, the more sure
the conclusion. It is possible, especially in a subspe-
cific problem, to have any one individual wrongly
identified as belonging to a population. This becomes
more likely, the closer the populations resemble each
other. Therefore, conclusions about which population
is present in a certain area should be made on a basis
of the population most frequently identified from
that area. One specimen (or perhaps a few) is not
enough to make a valid conclusion about the
population structure of an entire area.

SUMMARY AND CONCLUSIONS

Little work has been done previously on the tax-
onomy of Crotaphytus collaris baileyi. This and the
demonstration by Fitch and Tanner that C. c. baileyi
is a heterogeneous grouping prompted the present
study. Only the populations of the type material, the
Upper Colorado River Basin, and the Chihuahuan
Desert were studied. Multivariate, variance, canonical
and discriminant analyses of external characters were
performed, and pattern-coloration characters were
examined on living specimens.

The results of the analyses show significance be-
tween all three populations. The discriminant func-
tions distinguished between the populations with 80%
reliability and patternal characters were discrimin-
atory with near 100% reliability. Therefore, a new

subspecific name, C. c fiisciis, was applied to the
Chihuahuan population.

The co//ffra-complex was shown to consist of at
least four subspecies: C. c. auriceps. C. c. baileyi, C. c.
collaris and the new subspecies, C. c. fuscits from the
Chihuahuan Desert. C. c. auriceps' range was re-
stricted to the area near Moab, Utah, and north of the
union of the" Green and Colorado rivers. A broad
intergrade zone south into the Painted Desert was
established between C. c. auriceps and C. c. baileyi.
The range of C. c. baileyi was established as central
Arizona. Southern and central New Mexico and most
of Mexico east of central Sonora were established as
the range for C. c. fuscus. Further study of the popu-
lations presently recognized as C. c. collaris was ad-

28

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

vised.

While all the populations were separable on the
basis of morphology, the best characters tor identifi-
cation were color and pattern. C. c. auriceps has a
light green body and the yellow of the head extends
onto the side of the throat. C. c. baileyi has a darker
green body with reduced yellow on the head. The
area of the throat between the infralabials and the
gular patch is always white. C. c. fusciis has a brown

bt)dy and a white to cream head.

Important discoveries were also made in method-
ology. The necessity for using multivariate statistics
in taxonomic studies which investigate more than one
character was demonstrated. The use of a posteriori
probabilities was presented as a new technique for
investigation of taxonomic problems involving inter-
gradation.

ACKNOWLEDGMENTS

We wish to extend our gratitude to Dr. Melvin W.
Carter and Mr. Vernon Crandall for their suggestions
and aid in the preparation of the statistical material
and to Dr. Clive D. Jorgensen for introducing to us
Cluster analysis.

The manuscript was read by Drs. Ferron L.
Anderson, August W. Jaussi, H. Duane Smith and
Stephen L. Wood of Brigham Young University and
by Dr. Henry S. Fitch of Kansas University. We ap-
preciate their helpful suggestions.

Materials were supplied by the following institu-
tions. Brigliam Young University (BYU), California
Academy of Science (CAS), California State College
at Long Beach (CSCLB), Los Angeles County
Museum (LACM), Museum of Vertebrate Zoology

(MVZ), United States National Museum (USNM),
University of Colorado (UC), University of Illinois
Natural History Museum (UIMNH). University of
Kansas (KU), and University of Texas at El Paso
(UTEP). For the loan of specimens and courtesies
shown us by the personnel of these institutions, we
are most grateful.

We also express our appreciation to Mr. Nathan M.
Smith and Mr. J. Robert McMorris for their help with
specimens in the laboratory and in the field. Mr.
Michael Andelin prepared the art work, and Mrs.
Janice Ingram assisted in editing and typing the rough
draft of this manuscript. We appreciate their assis-
tance and the assistance of all those involved in this
study.

BIOLOGICAL SERIES. VOL. 13, NO. 2 CROTAPHYTUS COLLARIS

29

LITERATURE CITED

Allen. M. J. 1933. Report on a collection of amphibians and
reptiles from Sonora, Mexico, with description of a new
lizard. Occ. Pap. Miis. Zool. Univ. Mich., 259:1-15.

Anderson. T. W. 1958. An introduction to multivariate
statistics. John Wilev and Sons, Inc., London, p.
126-307.

Burt, C. E. 1928. The synonymy, variation and distribution
of the Collared Lizard, Crotaphvtus coHaris, (Say). Occ.
Pap. Mas. ZooL Univ. Mich., 196:1-19.

Cope, E. D. 1900. The crocadilians, lizards, and snakes of
North America. Ann. Rep. U.S. Nat. Mus. for 1898,
p.153-1270.

Di.xon, W. J. 1968. Biomedical computer programs. Univ. of
Calif. Press, Los Angeles, p. 214a-215.

Fitch, H. S. and W. W. Tanner. 1951. Remarks concerning
the systematics of the Collared Lizard {.Crotaphvtus
collaris). Trans Kans. Acad. Sci., 54(4):548-559.

Fitch, H. S. 1956. An ecological study of the Collared Liz-
ard {Crotaphytus collaris). Univ. Kans. Mus. Nat. Hist.
Publ., 8:213-274.

Mavr, E. 1969. Principles of systematic zoology. McGraw-
Hill Book Co., New York. p. 181-197.

Morrison, D. F. 1967. Multivariate statistical methods.
McGraw-Hill Book Co., New York. p. 159-207.

Ostle, B. 1966. Statistics in research. Iowa State Univ. Press,
Ames, Iowa. p. 129, 512-564.

Pack, L. C. and W. W. Tanner. 1970. A taxonomic compar-
ison of Uta stansburiana of the Great Basin and the
Upper Colorado River Basm in Utah, with a description
of new subspecies. Gt. Basin Nat., 30(2) :7 1-90.

Rao, C. R. 1952. Advanced statistical methods in biometric
research. John Wilev and Sons, Inc., New York. p.
273-370.

Schmidt. K. P. 1922. The amphibians and reptiles of Lower
California and the neighboring islands. Bull. Amer. Mus.
Nat. Hist., 46:607-707.

Smith. H. M. 1946. Handbook of lizards: lizards of the
United States and Canada. Comstock Publishing Co.,
Ithaca. New York. p. 166-172.

Smith, H. M. and E. H. Taylor. 1950. An annotated check-

list and key to the reptiles of Mexico, exclusive of the
snakes. Bull U.S. Nat. Mus. p. 199.

Sokal. R. R. and P. H. A. Sneath. 1963. Principles of numer-
ical taxonomy. W. H. Freeman and Co., San Francisco,
p. 1-357.

Sokal, R. R. and F. J. Rohlf. 1969. Biometry: the principles
and practices of statistics in biological research. W. H.
Freeman and Co., San Francisco, p. 1-776.

Stebbins, R. C. 1954. Reptiles and amphibians of western
North America. Univ. of Calif. Press, Berkeley, Cali-
fornia, p. 227-229.

Stebbins, R. C. 1966. A field guide to the western reptiles
and amphibians. Houghton Mifflin Co., Boston, p.
99-100.

Steel, R. G. D. 1955. An analysis of perennial crop data.
Biometrics, 1 1(2):201-212.

Stejneger, L. 1890. An annotated list of reptiles and ba-
trachians collected by Dr. C. Hart Merriam and Vernon
Bailey on the San Francisco Mountain Plateau and
Desert of the Little Colorado River, Arizona, with de-
scriptions of new species. N. Amer. Fauna. 3:103-118.

Stejneger, L. and T. Barbour. 1917. A checklist of North
American amphibians and reptiles. Harvard Univ. Press,
Cambridge, Massachusetts, p. 45.

Stone, W. and A. G. Rehn. 1910. On the terrestrial verte-
brates of portions of southern New Mexico and western
Texas. Proc. Acad. Nat. Sci. Phila., 40:16-34.

Van Denburgh, J. and J. R. Slevin. 1921. Preliminary diag-
nosis of new species of reptiles from islands in the Gulf
of California, Mexico. Proc. Calif. Acad. Sci. Ser. 4,
11:95-98.

Van Denburgh. J. 1922. The reptiles of western North
.'\merica. Vol. 1, "Lizards", Occ. Pap. Calif. Acad. Sci.,
10:95-98.

Wishart, D. 1968. A fortran II programme (CLUSTAN) for
numerical classification. C^omputing Laboratory, St.
Andrews, Fife. Scotland, p. 1-50.

Wishart, D. 1969. Mode analysis: a generalisation of nearest
neighbour which reduces chaining effects, pp. 282-308.
In A. J. Cole (ed.) Numerical taxonomy. Academic
Press, London.

c^-CV)^^-^"^^^^

Brigham Young University makv

UNivizr

Science Bulletin

A TAXONOMIC REVISION OF THE

WEEVIL GENUS TYCHIUS GERMAR

IN AMERICA NORTH OF MEXICO

(COLEOPTERA: CURCULIONIDAE)

by

Wayne E. Clark

BIOLOGICAL SERIES — VOLUME XIII, NUMBER 3

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, Utaii. All remittances should be made payable to Brigliam
Young University.

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

Brigham Young University
Science Bulletin

A TAXONOMIC REVISION OF THE

WEEVIL GENUS TYCHIUS GERMAR

IN AMERICA NORTH OF MEXICO

(COLEOPTERA: CURCULIONIDAE)

by

Wayne E. Clark

BIOLOGICAL SERIES — VOLUME XIII, NUMBER 3

APRIL 1971

TABLE OF CONTENTS

Page

INTRODUCTION 1

HISTORY 1

MATERIALS AND METHODS 2

BIOLOGY 3

INTRASPECIFIC VARIATION 4

TAXONOMIC CHARACTERS 4

PHYLOGENY 5

SYSTEMATIC SECTION 6

Genus Tychius Germar 6

Key to North American species of Tychius 8

Tychius stcphcnsi Schoenherr 10

Tychius sordidus LeConte 14

Tychius caesius, new name 15

Tychius lincelhts LeConte 16

Tychius liljcbladi Blatchley 19

Tychius tcctus LeConte 20

Tychius semisqucimosus LeConte 23

Tychius lamellosus Casey 24

Tychius badius, n.sp 25

Tychius proUxus Casey 26

Tychius soltaui Casey 28

Tychius montanus, n.sp 29

Tychius hirsutus, new name 30

Tychius phalarus, n.sp ,31

Tycliius arutus Say 33

.\CKNO\VLEDGME\TS .' 36

LITERATURE CITED 38

A TAXONOMIC REVISION OF THE WEEVIL GENUS

TYCHIUS GERMAR IN AMERICA NORTH OF MEXICO

(COLEOPTERA: CURCULIONIDAE)'

by
Wayne E. Clark-

ABSTRACT

A study of morphological characters of 4,000
adult weevils used in preparing a key and
descriptions indicates there are fifteen North
American species in the genus Tyclihts Gennar.
Adults occur on plants in the genera Astragalus,
Baptisia, Hcdi/sarum. Lotus, Lupinus, and Oxij-
tropis.

The genus is divided into two species groups.
The T. sordidiis group appears to have repre-
sentatives in the Old World fauna, but the T.
seiniscjuamosus group is probablv native to
North America.

New names, Tijchius caesius, and T. hirsutus
are proposed for T. arniatus Green, 1920 (not
Tournier, 1873), and T. hirtellus LeConte, 1876
( not Tournier, 1873 ) respectively. Three species,
T. badius, T. montanus, and T. pJialarus, are
described as new. A neotype is designated for
T. aratus Say. Lectotypes are designated for T.
tomcnto.ms (Herbst), 1785, T. stephensi Schoen-
herr, 1836, T. lineeUus LeConte, 1876, T. semis-
ijuoinosus LeConte, 1876, and T. hlrtelhis Le-
Conte, 1876. The name Puratychus Casey, 1910,
is newly placed in synonymy with Ty chins Ger-
mar, 1813.

INTRODUCTION

Weevil species assigned to the genus Tychius
Gennar, 1817, have been described from North
and South America, Europe, Africa, Asia, and
Australia. The majority of approximately 266
species occur in the Mediterranean region
( Klima, 1934 ) . Fourteen native and one in-
troduced species are known to occur in North
America.

So far as is known, all species of Tychius in-
fest the pods of leguminous plants. Several
species are of economic importance in the Old
U'orld ( Muka, 1955 ) . One of these, T. stephensi
Schoenherr, 1836, has been introduced into North
.America and is a pest of cultivated clover. The
native North American species have been re-
corded from plant species in the genera Astra-

galus, Hedysarum, Oxytropis, Lotus, Lupinus
and Baptisia. Some Tychius species may play a
part in the natural control of these plants
(Marcovitch, 1916), some of which are poisonous
to livestock (Hulbert and Oehme, 1961).

To date the most complete treatments of the
genus are the works of LeConte ( 1876), and
Casey (1892, 1910). These papers provide keys
and descriptions of some species, but are of
limited use in identifying specimens. The objec-
tives of this revision have been to provide accu-
rate descriptions and keys for the identification
of new and known species, and to contribute to
the knowledge of the biology and phylogeny of
the group.

HISTORY

The genus Tychius was established by Ger- stus Fabricius, and ])icirostris Fabricius. By sub-
mar (1817:.'}4), who used the name in associa- sequent designation, Schoenherr (1825:583) des-
tion with the previously validated specific names, ignated Curculio (juinquejninctatus Linnaeus as
cpiinquepunctatus L. (cited .5-punctatus), venu- the type-species. In the same work Schoenherr

'Thesis subniilted in partial fulfillment nf requirements for the MS. degree at Brigliani Young University.

-Department of Zoology, Brigham Young University, Provo. Utah: now at the Department of Entomology. Texas A&M University. (College
Sution, Texas 77801.

BniGHAM Young University Science Bulletin

erected Miccotrogus as a svibgemis of Ti/rliuis.
Later (1826:245-247) he characterized Tyclnm
as having seven and the subgenus Miccotrogus
as having six antennal funicular segments.
Stephens (1839:229) elevated Miccotrogus to
the rank of genus.

Tiicliius stephensi was described by Schoen-
herr (18.36:412), evidently prior to its introduc-
tion into North America. Thomas Say ( 1831 )
described Ti/chius (initus and T. amoenit.s. Ty-
chitis amucnus was transferred to Pachijtycliius
Jekel by LeConte (1876:168, 216), and was in-
cluded in Smicront/x Schoenherr bv Anderson
(1962:264-266). Clyllenhal (18.36:414-415) ap-
plied the name Tychius arator to a specimen re-
ceived from Thomas Say, identified by Say as T.
(iratm. LeConte (1876:216-218) described as
new, Tychius lincclhts, T. sordidit.s, T. tcctus, T.
semiscjuainoms, T. hirtclhts, and T. setosus.

Casey (1892:411-425) divided the North
American species of Tychius into four subgenera;
I and II possessing seven and III and IV possess-
ing six antennal funicular segments. Subgenus
I was characterized as having ". . . the elytral
intervals entirely devoid of recurved setae," and
subgenus II as having ". . . elytral intervals with
recurved semi-erect setae." Subgenus III was de-
fined as ". . . with recurved setae, the entire facies
almost as in group II . . . ," and subgenus IV as
". . . the species generaly minute, with or without
erect setae. . . . ' Subgenus I contained T. lineel-
hts LeConte, T. sordidus LeConte, T. tcctus Le-
Conte, and T. arator Gyllenhal. Subgenus II con-
tained T. liirtelhis LeConte, T. semi.squamosus
LeConte, T. aratus Say, and two species de-
scribed as new, T. soUaui and T. lameUosiis. In
subgenus III Casey placed a single species de-
scribed as new, T. proxihis. Subgenus IV con-
tained T. setosus LeConte and six others de-
scribed as new, T. varicgatus, T. simplex. T.
sihinoidcs. T. mica, T. subfasciatus, and T.
hispichis. CJasey did not recognize Miccotrogus
in this work because a specimen sent to him by

Desbrochers, identified as M. j)icirostris, had
seven instead of six antennal funicular segments,
and because his own North American species
with six funicular segments, T. proxilus, agreed
closely in other respects with the other species
of Tychius. I have examined a specimen in the
Casey collection identified as M. picirostris and
found it to be a Tychius stephensi Schoenherr.
Casey (1897:664-666) described three species
which he assigned to subgenus IV; T. sidtcatu-
lus, T. inermis, and T. transversus. Another
North American worker, Schaeffer (1908:217-
219) described T. griseus, T. suturalis, T. palli-
dus, and T. ulhidus. In a subsecjuent note (191.5:
197) he stated that T. griseus was a synonym
of a species he called Tychius (Miccotrogus)
picirostris ( Fabricius ) .

Casey (1910:1.32-142) established the sub-
genus Paratychiiis for T. proxilus which he had
formerly assigned to subgenus III and T. imhri-
catus which he described as new. In the same
publication he erected the subgenus Micro-
tt/chius to include the species fonnerly as-
signed to subgenus IV, as well as thirteen species
described as new; T. erraticiis, T. j)ucUus, T.
atomus, T. cchimis, T. vcrnilis, T. fatuus, T.
fratercuhis, T. gryjuis, T. dtdcis, T. imheUis, T.
jwrcatus, T. curtipemiis and T. errans. He also
described nine species belonging to subgenus I,
T. tacitus, T. hesperis, T. radians, T. dilectus, T.
probus, T. tcxanus, T. carolinae, and T. lan-
guidus. Five species occurring in the eastern
United States, including one described as new,
7'. liljclihidi, were treated !)v Blatcliley and Leng
(1916:24.5-247). Leng ( 192d:.321) listed all of the
species described by Schaeffer (1908:217-219),
except T. griseus, under Microtychius Casey.

Kissinger ( 1964:57-.5S) transferred Paraty-
chius and Microtychius to tiie genus Sibiniu
Gemiar and suggested further study to deter-
mine the true relationship of these groups to
Tychius. I liave followed his classification, but I
include Parati/chius in Tychius instead of
Sibinia.

MATERIALS AND METHODS

Most ol approximately 4,(X)0 specimens ex-
amined in this study were borrowed from col-
lections of institutions in the United States and
("anada. I collected about 1,5(X) specimens.

The following abbre\iations are used to indi-
cate the collections in which the specimens
examined are deposited: AMNH, American Mu-
seum of Natural History, New York; UA. Uni-

versit\' of Arizona, Tucson; BYU, Brigham
Young University, Provo; CAS, California Acad-
emy of Sciences, San Francisco; CIS, California
Insect Survey, University of California, Berkeley;
CNC, Canadian National Clollection, Ottawa;
CU, Cornell University, Ithaca; FMNH, Field
Museum of Natural Histor)-, Chicago; INHS,
Illinois Natural Historv Survey, Urbana; ISU,

Bioi.oGicAi. Series, Vol. 13, No. 3 Revision ok the Weevil Genus Tvchius

Iowa State Uiii\er,sity, Ames; UK, University oi
Kansas, Lawrence; LA, Los Angeles County
Museum. Los Angeles; MCZ, Museum of Com-
parati\e Zoology, Harvard University, Cam-
liridge; CW'O, collection of Charles \V.' O'Brien,
Texas Tech Uni\ersit)', Lubbock; OSC, Ohio
State University, Coliunbus; OSU, Oregon State
I'niversity, Corvallis; PANS, Academy of Natural
Sciences of Philadelphia; TANL Texas A. & M.
I'niversity, College Station; USNNL United
States National Museum, Washington, D.C.;
USU, Utah State University, Logan;"^ WEC, col-
ii'ction of the autlior.

I ii;i\e exiunined the types of all known North
American species and their synonyms except
the types of some alleged synonyms of Ti/chiiis
stcphcnsi Schoenherr and the type of T. aratus
S;n' which is presumably destroyed (LeConte,
lS59:vi).

Ail measurements were made using a cali-
brated eyepiece reticule with a dissecting micro-
scope at magnifications up to 80 times. Total
lenirth and width were measured from the dor-
sal aspect, length from the dorsal margin of the
eyes to the elytral apices, width at the widest
point across the ehtra. Length of the rostrum
was measured from the lateral aspect from the

apex to the anteroventral margin of the eye.
Length of the pronotum was measured from the
lateral aspect from the anterior margin to the
base. Other measurements recjuire no further
explanation.

Male external sjenitalia were removed for
study. Specimens were taken directly from al-
cohol, or if previously mounted, soaked in warm
water until soft. Holding the specimen between
the thumb and forefinger, the abdomen was
forced down with a pin exposing the tergum.
The tergimi was torn with a pin and the pin
inserted beneath the median lobe to lift it into
view. The structure was then removed with a
pair of jeweller's forceps. Genitalia were placed
in 10 percent KOH to remove muscle tissues,
washed in 90 percent alcohol, then stored with
glycerin in polyethylene microvials attached to
the pin with the specimen.

Line drawings were made with grid paper
and an eyepiece reticule in a dissecting micro-
scope. Genitalia were drawn immersed in glyc-
erin. Definitions of terms used, except those
describing genitalia, may be found in Torre-
Bueno (1962). Terms used in reference to geni-
talia are those of Sharp and Muir ( 1912).

BIOLOGY

Biology of the red clover seed weevil Tijchius
stcphcnsi Schoenherr, has been studied by Muka
(1955). According to him larvae feed on de-
veloping seeds of red clover while adults feed
on reproductive portions of flowers of the same
plant. Adults overwinter in soil around the host
plants, ;md emerge in the spring and commence
a migration flight. Females ovijjosit in the florets,
laving one egg per floret on the ovarv inside the
corolla tube. In New York state there are two
generations per year on red clover.

.\dults ot Tijchhis linecUus were observed on
lAijiiiuis Icucoiihijlhis at the mouth of Hobble
Creek Canyon, Utah County, Utah, on May 3,
before the plants were in bloom. Copulating
pairs were seen on florets; females with their
rostra piercing the corolla of partially opened
flowers. Apparently females feed on pollen
grains. This was indicated by examination of gut
contents and fecal material which were similar
in color and texture to pollen of the Lupines.

Before ovipositing, the female makes a hole
in the calyx and deposits one or two eggs on

the side of the ovary. Larvae feed on seeds in
the developing pods. When the larvae are mature
they evidently chew a hole in the side of the
pod and drop to the ground to pupate. Although
no larvae or pupae were actu;dly found in the
soil, holes were observed in the sides of mature
pods which showed signs of infestation. Muka
(1955) described similar habits in T. stephensi.
According to Mitchell and Pierce (1911), larvae
of T. sordidiis "emerge " from Baptisia pods and
pupate in the ground.

On May 19, pods of Astra^aJtis titaliensis
(Torr. ) T. and G., the host of Tycluus prolixus,
were collected at Provo, Utah. Several larvae
were taken from the pods at that time, and on
July 28, four adult weevils were taken from
the bag containing the pods. These pods \yere
subse(juently dissected and out of 266 pods, 18
showed signs of infestation including several
containing dead larvae. White cocoons about
3.5 mm in length were found in two of the pods,
and holes about 1.3 mm in diameter were ob-
served in the sides of the pods in the portion
covered by the cocoons. This does not provide

Bhk.ham Young Univf.hsitv Sciknce Bulletin

condiisivc evidence that the weevils normally On August 1, a few live adults were sifted from
pupate in pods, since Muka (1955) states that T. soil taken beneath A. utahetisis indicating that
stepheiisi can be "forced" to pupate in the pods. the weevils may on envinter as adults.

INTRASPECIFIC VARIATION

The sexes can be distinguished by differences
in the structure of the pygidium. The pygidiuiii
of the male in its normal position is nearly per-
pendicular to the longitudinal a.xis of the body
and is \isible for more than half its length
beyond the elytral apices. A transverse carina
divides it along the line normally attained by the
elvtral apices. The pvgidium of the female in its
normal positon is obliijue rather than perpendic-
ular to the longitudinal axis, nearly covered by
the elvtra, and lacks a transverse carina.

The rostrum of the female is usually longer
and more slender than that ol the male espeeiallv
in T. (imtus (Fig. 3). where the rostrum of the
female is more than half tiie length of the body.
The antennal insertion is normally more distad
and the distal portion less stronglv tapered and
with deeper pits and rugae in the male. The
apical tibial mucrones are smaller in females
of most species. Females average about 0.1 mm
longer than males.

The average difference in length of native
Xorth .\merican species was 32 percent of the
length of the smallest specimen. Environmental
conditions, espeeiallv size and number of seeds
and or lar\ae per pod, probabh' influence the
size attained by individual specimens.

Color of the integument ranges from light
piceous to black. The general color of a spi'cimen
is imparted to it by the color of the scales. In
7". stepheiLii and T. tectiis, scale color ranges
from light gray to tawniy in specimens w ithin a
given series. Muka (1955) observed tli.il newly

emerged specimens of T. stcphcn.si were yellow-
ish brow n and that with age scale color changed
in many specimens to pale gray. Specimens of
the T. seinisijiuimosus species group often ex-
hibit variation in the color of the round or
elongate-oval, nonstrigose scales of the elytra and
|)rothora\'. These vary from white to dark reddish
brown on each specimen. They are usually dark-
est on interspaces five through seven. The long,
narrow, strigose scales on the pronotum and
elvtra also vaiy from very light to dark reddish
brown in these species.

Specimens may also exhibit variation in the
distribution of certain types of scales. In T. tec-
ins and T. liljehladi, white, round scales on the
elvtra may be very dense or sparse. The num-
bi'r and unifonnity of the median rows of long,
narrow, strigose scales on the elvtral interspaces
may vary, especially in T. semiiquamostts and
T. lamellosiis, and to a lesser extent in other
members of the T. scmisqtiatnosus species group.

In some species the rostrum from lateral as-
pect may vary from evenly and prominently
arcuate from the base to the apex, to prominently
arcuate in the basal portion, and nearly straight
to the apex. In other species one extreme or the
other may be consistent.

Geographic variation was noted mainly in the
overall size of specimens and in the shape, color,
and distribution of scales. Where geographic
variation was observed, it is described in greater
detail in the discussion following the description
of the species involved.

TAXONOMIC CHARACTERS

Color, shape and distribution of scales, shape
of the rostrum and structure of the male geni-
talia, provide good characters for distinguishing
species.

Scale color varies from gray as in 7'. sonlidiis
to tawny yellow as in T. tecttis and a combina-
tion of nearly white and reddish brown as in
most species of the T. .ieinis<niaiiio.sus species
group. The presence or absence of rows of erect
or suberect setae on the elytral interspaces is
important in separating species groups. Posses-

sion or absence of fine erect setae on the ab-
domen and metathorax is an important character
in separating species.

The relative length of the rostrum in com-
parison with tile prothorax varies. From the
dorsal aspect, the rostrum may be wide at the
base, becoming acuminate towards the apex as
in T. lamcllostts, or narrow basallv and widening
at the apex as in T. sonlidtis. Sculpture of the
portion distad of the antennal insertion may be
shallow or deep.

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius

The shape of the apical portion of the medi-
an lobe i.s an important character tor .separating
clo.selv related .species. In its simplest fonn the
apex is more or less evenlv ronnded or \\'ith a
slight apical prominence as in 7'. sonliclit.s ( Fig.
it), T. cacsim (Fig. 15), T. liljchlaili (Fig. 1.3),
r. tectus (Fig. 12), T. soltaui (Fig. ll), T.
phiiUims (Fig. 7), and T. proUxus (Fig. 6). In
T. linccUus (Fig. 16), the apical prominence
is greatK' exaggerated. In T. Jnidiiis (Fig. 5), T.
moiitanus (Fig. IS), and T. Iiirstittts (Fig. 14),
the apical portion bears prominent lateral apical
prominences. Weak lateral apical prominences
are present in T. aiatus (Fig. 4). In T. scmi-

s(juaiiio.sus (Fig. 9) and 7'. JmncUo^us (Fig. 8),
the apical portion is asymmetrical. Size and
shape of the median apical membraneous area is
important in distinguishing between the closely
related species 7'. liljchhidi (Fig. 13) and T.
tcctu.s (Fig. 12), and between T. soltaui (Fig.
11) and r. phakirus (Fig. 7). The median struts
may be stout in some species as in T. hadius
(Fig. .5), or very slender as in T. sordidtts (Fig.
17). The tenninal clubs on these structures in
some species such as T. icctn.s ( Fig. 12) may also
be important. Structure of the genitalia of T.
■stcphcn.si (Fig. 10) appears unrelated to any of
the native North American species.

PHYLOGENY

Since most species of Ttjcluus occur in the
Old World and have not been studied, a de-
tailed discussion of their ph\'logenv \\ ill not be
attempted. Some trends are evident howe\er,
among the North American species.

The native North American species are di-
vided into two species groups. These are char-
acterized in the discussion following the descrip-
tion of the genus. The T. sordidtis species group
is probablv the most primitive. The palearctic
fauna contains species which appear to be close-
ly related to this group. In this group T. sordidti.s.
T. caesius, and T. lincellus are relatively large in
size with gray scales. The toothed protibia of T.
lineeUus is unifjue among North American spe-
cies but some European species possess a similar
tooth. Se\eral characters expressed by T. lilje-
hladi and T. tectus suggest intenncdiacy be-
tvveen the two species groups. In addition to the
long, narrow, strigose scales, T. liljebladi possess-
es a few scattered, round, white, nonstrigose
scales on the elytra. Ti/chius tectus usually pos-
sesses definite rows of white, round scales on
interspace one, near the humeri, and on the pro-
notum, giving the insect a striped appearance.
The general body form in these two species is
also intennediate. The body form in T. liljchhidi
is more like the species of the T. sordidtis species
group, whereas T. tectus more closely resembles
species of the T. semistjuamosus species group.
These two species occur on Astra'^(dtis as do spe-
cies of the 7'. scmisquamosus species group. The
other members of the T. sordidtis species group
occur on species of the plant genera Baptisia and
Lupinus.

The species of the T. semis(ptaiiiostis species
group appear to be more distantly related to
the palearctic fauna.

Reduction of the rows of long, narrow scales
on the elvtral interspaces from multiseriate to
unifonn, median, uniseriate rows, and the de-
\elopment of erect, hairlike setae on the ventral
surfaces appear to be important trends within
the T. scmisquoinostis species group. Ttjchitis
semiscitiamostts and T. himcUosus have multi-
seriate rows of long, narrow scales on the elytral
interspaces but lack erect, hairlike setae on the
ventral surfaces. This indicates relationship to
the T. sordidtis species group in which the ely-
tral interspaces are clothed exclusivelv with long,
narrow scales. Tijchitis hadius appears to occupy
a position intermediate between T. lamellosus.
and T. soltaui. This species has a reduced num-
ber of rows of long, narrow scales on the elytral
interspaces and also lacks erect hairlike setae
on the ventral surfaces.

Structural variation in the male genitalia does
not appear to indicate major trends. The assym-
metrical apical portion of the median lobe in T.
scmiscpiamosus (Fig. 9), and T. lamcUosus (Fig.
S) is uni([ue among the North American species.
The apical and lateral prominences on the me-
dian lobes of the genitalia of T. hadius (Fig. 5),
7'. ntontantis (Fig. 18), and T. Iiirstitus (Fig.
14). may function as isolating mechanisms.

Possession of uniseriate rows of long, narrow
scales on the elytral interspaces, the absence of
erect, hairlike setae from the venter, and close
resemblance to T. lameUosus indicates that T.
proxihis, for which Casey (1910) erected the
subgenus Parattjchius, arose in North America
witli the T. scmisquamosus species group. The
difference in number of antennal funicular seg-
ments does not appear to warrant giving this
taxon generic or subgeneric rank.

BiuGiiAM ViniNc: Univehsitv Science Bulletin

ftjcltius phalarus appears similar to 7". soltaui,
but se\cral diaracters of tlie rostrum and vesti-
ture suggest that they are not closely rehited.
ThLs species is associated with the plant genus
Lotus rather than Astraf^ahis.

The trend in the T. senm(iiia>nosti.s species
group toward refinement of the long, narrow

scales on the elytral interspaces is culminated in
T. Iiirsiitu.s-; these scales taking the form of very
long, white, hairlike setae.

Tyclihis (iratus is distinct in many features
from the otlier members of the T. seinis(juomosus
species group. Its relationship to the group is
uncertain.

SYSTEMATIC SECTION

Genus I'ljcliius (Jennar

Ttjchius Gcnii.ir, 1817, Magazin <lti Kntoinologic (Gcr-
mar), 2:.340 ( Type-specie.s, Ctirculio iittiiKjnepunctii-
tus Linnaeus, 1758, by subscijucnt designation,
Schoenherr, 1825:583).

Parahjchius Ca.sey, 1910, Can. Entomol., 42:1.35 (Typc-
spccies, Ti/chiii.s proxilu.t Casev. by original designa-
tion). NEW SYNONYMY.

The genus Tijchius in North America may
have six or seven antennal funicular segments. It
is closelv allied to .\/irr«/r()gi/v Schoenherr, 1S2.7,
one species of which, the introduced M. piciw-
stris (Fabricius, 1787), occurs in North America.
Ttjchius and Miccotw<^us, in the female, ha\'e
the elytral apices conjointly rounded concealing
the pygidium. Four related genera, Itijchiis
Kissinger, 1962, Par(i<io<^es LeC'onte, 1S76, Me-
ajnopijiiu Pierce, 19()S, and ^ihinia Gennar, 1817,
each with si.x antennal funicular segments, oc-
cur in North .America. These genera all have the
elytral apices separately rounded, leaving the
pygidium broadly exposed in both sexes.

Description. Length 2.0-.5..3 mm, female usu-
ally 0.1-0.2 mm longer than male; integument
light reddish brown to black; appendages and
rostrum usually lighter in color than bodv.
V'estitiire of gray, yellowish or reddish brown
and white scales.

Rostrum longer or shorter tiian protliorax; in
dorsal aspect both nearly parallel from base to
apex; apex wider than frons between dorsal
margin of eyes, or finely tapered from basi> to
apex; frons between dorsal margin of eves as
nuich as 2,5 times wider than rostrum ;it tip;
usually glabrous or with a few elongate scales
distad of antennal insertion; antennal insertion
.It middle of rostrum in female, in distal third or
fourth in male.

.Antennae with last funicular segment with
row of ;ilternately long and short sciiJes.

I'ronotum as wide or wider than long; sides
usually e\enly rounded, slightly constricted
apieally. wider at base than at apical constric-

tion. Vestiture of lon<', narrow scales on dorsum,
round or elongate-oval; usually lighter colored
scales on ventral portion of lateral surface, often
with round or elongate-oval scales on dorsum
in median and lateral vittae.

Elytra nearlv parallel sided in basal two
thirds, humeri not prominent; in lateral aspect
either broadlv rounded or nearly flat in basal
iialf, declivity evenly rounded; striae deep,
punctures even, clearly visible, strial setae fine,
hairlike or broad. Vestiture of long, narrow scales
ot uniform size and shape, or round to elongate-
oval, usually broadly imbricated scales with
median ro\\'s of long, narrow scales on each in-
terspace.

Vt'ntral surface with broadly imbricated,
usually white, round to elongate-ov;d scales; su-
ture between sterna t\\'o and three strongly
produced posterolaterally, reaching or passing
suture between sterna three and four (Fig. .3).
Sterna three and four about etjual to sternum
fi\e in length; sternum five usually with deep
median fovea.

Front coxae contiguous, femora usually
swollen in apical two tliirds, usuallv with strong-
ly ile\eloped apical, ventral emargination; often
witii minute tooth or spine on pro.\imal portion
of apical ventral emargination; vestiture of long,
iKurow and elongate-oval scales, or elongate-oval
sciiles alone.

Tibiae inueronate, muero on protibia usually
larger and stouter; apex of tibia with uniform
row of stout, usuall\- light vellowish brown
bristles; \esfiture of long, narrow, and round or
elongate-oval scales, and elongate, veiT fine, hair-
like setae.

Tarsi with p;ids of very fine white setae on
\entral surfaces, dorsal surfaces with long, nar-
row sc;des and fine, hairlike setae; claw with
biisal process about two thirds length of claw.

Mall' genil;dia with median, usually apical,
nuinbnmous area; apex of median lobe rounded,
or asymmetrical, often with apical, lateral
prominences; median struts articulating with

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tvchius

ventral-lateral projections ot median lobe; teg-
nien small, Y-shaped (Fig. 12), not forming
ring.

Discussion. The native North American spe-
cies may be divided into two species groups.
Species of the T. sordidus species group have a
simple vestiture in which all scales on the elytra
arc long, narrow, and strigose, the elytral inter-
spaces lacking discrete rows of setae. The in-
troduced T. stcphensi is most closely related to
this group. The T. semisquamosus species group

has a complex vestiture in which the elytral
interspaces are clothed with round, usually imbri-
cated, light colored, nonstrigose scales, and
multiseriate or uniseriate rows of long, narrow
setae.

Forms included and host records. A list of
the species groups and species of Ttjcliitis in
North America and host plants from which they
have been recorded is given below. Synonyms are
given in parenthesis following the valid name
of each species.

Tychius species

Host Plants

Introduced species

T. stcphensi Schoenherr, 1836 Melilotns spp.

Fragaria spp.
Crataegus spp.
Vicia spp.
Trifolitim pratense L.

T . sordidus species group

T. sordidus LeConte, 1876 Baptisia leucantha Torr. & Gray

(nimius Casey, 1910) B. hructeatu Muhl.

(texatius Casey, 1910) B. cuneata Small.

(carolinac Cas'ey, 1910) B. i illosa (Walt.) Ell.

T. caesius, new name None Cited

r. liiwellus LeConte, 1876 Lupiniis alhifrons Benth.

(tacitus Casey, 1910) L. ammophilus Greene

{hespcris Casey, 1910) L. urgcnteus Pursh

(radians Casey, 1910) L. arltorus Sims.

(dilectus Casey, 1910) L. hiculor Lindl.

iprobus Casey, 1910) L. cauddtus Kell.

L. chainissionis Esch.

L. exeuliitus [ones

L. leucoplu/llus Dougl.

L. sericcus Pursh

T. liljehludi Blatchley, 1916 Astragalus eanadensis L.

r. tectus LeConte, 1876 Astragalus adstirgens Pallas

{languidus Casey, 1910) ssp. rohustior (Hook.) Welsh

A. hisulcattis ( Hook. ) Gray
var. Iwijdenianus ( Gray ) Bameby
A. scopulorum T. C. Porter e.\
Port. & Coult.
A. tcnellus Pursh
Oxt/tropis hessciji ( Rvdb. ) Blank
O. campestris ( L. ) DC.
O. lamhertii Pursh
(). sericca Nutt.
Hcdt/sartiin sp.

7 . scmi.'itiuainosus species group

T. semvKpiamosus LeConte, 1876 .\one Cited

T. lamelhsus Casey, 1892 Astragalus l>ccktiithii T. & G.

A. druniondii Dougl. ex Hook.
A. lentigiuosus Dougl. e.x Hook. var.
paltiivi ( M. E. Jones) M. E. Jones
.4. lonchocarpus Torr.

T. proxihis Casey, 1892 Astragalus amphioxifs Gray

(imbricatus Casey, 1910) A. doughisii Gray

A. utahcn.sis (Torr.) T. & G.
A. lentigenosus Dougl. ex Hook.

BiUGHAM Young University Science Bulletin

T htiilius. n.sp Astragalus scoimlontm T. C. Porter

ex Port. & Coulf.

A. I>isulcatus ( Hook. ) Gray

T. soUaui Casey, 1892 Astragalus flavus Nutt. ex T. & C.

var. flavus (M. E. Jones) Bameby
A. jlexuusus ( Hook. ) Don

7". montanus, n.sp None Cited

7'. /iir«i//u4-, new name Astragalus uuttallianus A. DC.

T. phalanis. n.sp LofiM rigidus ( Benth. ) Greene

T. aratus Sav. 1831 Astragalus crassicarpus Nutt.

(urator Cvllenhal, 1836)

Key to North American species of Tychitis

1 Elytral iiiterspaci^s bearing long, narrow, strigose scales of unifomi size and
shape; round, or oval, white scales, if present, not present on all interspaces;
dorsal profile of elytra broadly rounded sordultis group 2

r Each elvtral interspace bearing two distinct types of scales; round to elongate-oval,
recumbent, usually broadly imbricated, nonstrigose scales and long, narrow,
often fine and setifomi, strigose scales in uniseriate or multiseriate, median
rows; dorsal profile of elytra straight on disc, broadly rounded to apices on
declivity - semiMjuaino.sits group 7

2(1) M.ile with large triangular tooth near middle on ventral margin of protibia; scales
generally grav in color, often alternate elvtral interspaces with bronze colored
sciiles Unccllus

2' Male without large ti-iangular tooth near middle on ventral margin of protibia,
scales gray or yellow in color, not gray and bronze on alternate elytral inter-
spaces 3

3(2') All femora with small tooth or spine on proximal portion of apical, ventral emar-
giiiation; pronotum with sides broadly roundi'd, widest in about middle; all
scides gray or yellowish gray in color caesius

3' Metafemur often with a small tooth or spine on pro.ximal portion of apical \en-
tral emargination but profemur and mesofemur without tooth; pronotum
widest at middle or at base; .scales gray or yellow in color 4

4(3") Rostrum from dorsal aspect wider at apex than irons between dorsal margin of
eyes; pronotum widest at base; scales yellow; vellowLsh gray, or grav, often
lateral margins of individual scales metallic bronze in color; no round white
scales on elytra sordidus

4' Rostrum from dorsal aspect narrower at apex than frons between dorsal margin
of eyes; pronotum wider in middle than at base; scales yellow, several round,
white scales on elytra 5

5(4') Rostrum from lateral aspect prominendy swollen at base; acuminate, smooth, with
ver)' shallow punctures distad of antennal insertion; round, white scales on
elytra sparse, unevenly distributed liljehladi

5' Rostrum from lateral aspect not prominently swollen at base; portion distad of an-
tennal insertion not acuminate, punctures and rugae deep; round white scales
concentrated on interspaces one and ;iround humeri on interspace eight 6

6(5') Round, white scales on elytra limited to interspace one, small, 2.0-2.6 mm in

length stepliensi

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius 9

6' Rouik], white .scale.s on elytra on interspaces one and eight, especially dense on
humeri, a few scattered scales rarely occur on other interspaces; larger, 2.5-
3.8 mm in length tectus

7(1) Antennal funiculus seven-segmented 8

7' Antennal funiculus six-segmented prolixus

8(7) Abdominal sterna each bearing a distinct transverse row of erect, hairlike setae;
rostrum usually with several round, white, nonstrigose scales on lower por-
tion of sides, or rostrum longer than prothorax 9

8' Al)dominal sterna without distinct, transverse rows of erect, hairlike setae; rostrum

witliout round, white, nonstrigose scales on lower portion of sides 13

9(8) Rostrum longer than prothorax, especially in female (Fig. 3); scales unicolorous;

length 4.1-4.4 mm aratus

9' Rostrum shorter than prothorax; insect with white and dark reddish brown scales;

length 2.6-3.9 mm 10

10(9") Rostrum distad of antennal insertion acuminate; white scales on dorsum of pro-
thorax fonning broad median vitta from base to apex of pronotum . 11

10' Rostrum distad of antennal insertion not acuminate, often slightly expanded in dor-
sal aspect at extreme apex; white scales on dorsum of pronotum, limited to
basal median patch 12

11(10) Long, narrow scales on elytral interspaces fine, hairlike, longer than width of
interspace, usually lighter in color than round, nonstrigose scales; rostnmi
distad of antennal insertion finely acuminate; elongate scales on dorsum of
prothorax narrow, integument broadly visible; median lobe of male gentalia
with lateral, apical prominences (Fig. 14) hirsuttis

11' Long narrow scales on elytral interspaces short, stout, shorter than width of inter-

space, usually darker in color than round, nonstrigose scales; rostrum distad
of antennal insertion evenly tapered, not finely acuminate; elongate scales on
dorsum of prothorax broad, integument concealed or only slightly visible;
median lobe of male genitalia without apical, lateral projections (Fig. 11) . . soltaui

12(10) White scales on dorsum of prothorax forming a large median, basal patch (Fig. 2);
iiu'tatliorax and visible abdominal sternum one with fine, erect, hairlike setae;
long, narrow, scales on femur darker than nonstrigose oval, wliite scales; me-
dian lobe of male genitalia without lateral, apical projections (Fig. 7) phalanis

12' White scales on dorsum of prothorax forming small, median basal patch; metatho-
rax and visible abdominal sternum one lacking fine, hairlike setae; long,
narrow scales on femora lighter in color than nonstrigose oval scales; median
lobe of male genitalia with weaklv developed apical, lateral projections ( Fig.
18) montanus

13(8') Long, narrow scales on elytral interspaces in nearly unifonn uniseriate rows; me-
dian lobe of male genitalia with well-developed lateral, apical projections
( Fig. 5) badius

13' Long, narrow scales on elvtral interspaces in confused, multiseriate rows; median
lobe of male genitalia \\ ith apical portion asymmetrical, lacking apical, later-
al projections (Figs. 8, 9) 14

14(13') Round, nonstrigose scales on elytral interspaces dense, imbricated; distal portion

of rostrum finely acuminate; length 2.4-3.4 mm lamcUous

14' Round, nonstrigose scales on elytral interspaces sparse, rarely imbricated; distal

portion of rostrum not finely acuminate; length 2.3-2.7 mm semmjuamosus

10

Brigham Young University Science Bulletin

Tijchitis stephemi Schocnherr
(Figs. 10,20)

Curculio picirostris Fabricius, 1787, Mantisa insectorum;
1:101 (Holotype: "Hafniae Dom. Liind." Copt-n-
hageii Museum, Fabriciu.s collection); Paykull, 1792,
Monographia curculionum Sueciae, p. 63.

Curculio fuscirostris Paykull, 1792, Monographia curcu-
lionum Sueciae, p. 62 (see discussion for infor-
mation on the "type ' ) .

Curculio hwicnlosus Herbst (not Olivier, 1790), In:
Jal>lonsky, 1795, Natursystem allcr bckannten in

vmd ausiaeiidischen insecten kaefcr, 6:278,

Tab. 81, Fig. 7 ( Lectotype here designated: female,
"Deutschhind", Z<K)logisches Museum, Berlin,
54577).

Rhynchacmis picirostris: CJvlUnlial, 1813, Insecta Sue-
cica , 1(3):121.

Tt/chius picirostris: Gcnnar, 1817. .Magazin der Ento-
mologie (Cermar), 2:340.

Tychiiis tomeniosus: Stephens, 1829, Systematic cata-
logue of British insects p. 160.

Tychius stepheni Selux-nherr, 1836, Genera et species
curculionidum. . . . 3:412 (Lectotype here designa-
ted: Female, "Anglia," British Museum Natur. Hist.,
J. F. Stephens coUecticm).

Tychius stcphensi: Stephens. 1839, A manual of British
coleoptera. or l)eetles. . . . , p. 229 (Emendation of
stepheni Schoenherr ) .

Miccotrogus picirostris: Casey, 1892, J. New York Ento-
mol. Soc, 6:411-412.

Tychius brevicollis Rey, 1895, Echange, 11:3 (types
not seen, synonymy from Klima, 1934:25).

Tychius clavipes Bey, 1895, Echange, 11:3 (types not
seen, synonymy from Klima, 19.34:26).

Tychius mixtus Rey. Echange, 11:4 (types not seen,
synonymy from Klima, 1934: 26).

Tychius griseus Schaeffer, 1908, J. New York Entomol.
Soc. 16:217-218 (Holotvpe: male, Itliaca, New
York, USNM, type 42484).

ThLs i.s the red clover seed weevil of North
American economic literature. It was probably
introduced into North America from Europe. It
closely resembles 7'. tcclti.s and T. Uljehladi in
several characters but differs by its smaller size,
by the structure of the male genitalia ( Fig.
10), and by its host preferences. It also closely
resembles Miccotro<nts picirostris { Fabricius )
but can easily be distinguished by the seven-
rather than six-segmented antennel funiculus
and other characters enumerated by Milliron
(1949). Muka (1955) studied the biology of T.
stcphensi and, Takenouchi (1965) described the
chromosomes of T. steplicnsi and M. picirostrk.

Description. Male: Length 2.0-2.5 mm,
width 0.9-1.2 mm; integument black to dark red-
dLsh browii, appendages light reddish brown.
Vestiture on appendages, thorax and elytra of

long, narrow, light yellowish brown scales, ven-
tral surface with white scales.

Rostrum shorter than prothorax, moderately,
evenly arcuate, slightly tapered to apex. Frons
slightly wid(>r between dorsal margin of eyes
than rostrum at apex. Integument distad of an-
tenna! insertion smooth and shining, rugae very
deep, especially laterally; glabrous except for
sparse fine setae on extreme Hp. Scales proximad
of antennal insertion of uniform size, shape and
color, parallel sided, truncate to rounded at
apices.

Antennal funicle seven-segmented, pedicel
longer than next three segments combined.

Prothorax 1.2 times wider than long, sides
broadly, evenly rounded, slightly constricted at
apex, less than two times as broad at base than
at anterior margin. Scales on dorsum of uniform
size, shape, and color, long, narrow, rounded at
apices; sc;iles on lateral surface round to elon-
gate-oval; long, narrow scales of dorsum ending
abruptly aliout one-fourth of the way down sides,
not intermingled with round or elongate-oval
scales on sides.

Elytra with sides broadly rounded; dorsal
profile broadly rounded, not flat in b;isal third.
Scales on dorsum slightly broader than those on
dorsum of prothorax. Interspace one usually with
distinct row of round, white, nonstrigose scales
extending entire length; round, nonstrigose scales
absent from other interspaces. Strial setae nar-
row, light colored on dorsum, broader and darker
in color laterally.

Ventral surface with dense, slightly imbri-
cated, round to oval, white scales, often with
plumose margins; elongate, hairlike setae absent.
Sternum five lacking median fovea.

Femora with ventral apical, emargination
weakly developed, no minute tooth on proximal
portion of emargination. Scales of unifonn size
and shape, similar to scales on elytra and pro-
thorax .Tibiae mucronate, mucro on protibia larg-
est; vestiture of long, narrow scales and fine
hairlike setae, no round, nonstrigose scales.

Male genitalia (Fig. 10), with median lobe
stout, strongly curved in lateral aspect, median
dorsal membranous area large, extending nearly
to proximal portion ot median lobe, with row of
sclerotic inclusions on e;ich side; median stmts
stout, finely tapered.

Female: length 2.0-2.6 mm. Rostrum more
finely tapcri'd, anti'unal insertion slightly distad
of middle.

Hosts. Recorded 1)\' Muka (1955): In Europe
from Melilottis, Fra<'ciria, Cratac"us, and Vicia,
and hmI clover, Trijolium prutense: in North
.\merica from Trijolium pratcnsc.

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius

11

Distribution. (Fig. 20).

Alberta: Edmonton, \'-14-21. H. W. \\'enzil, 1
male, 2 females (O.SC).

Arizona: Globe, III, D. K. Duncan, 1 male, 2
female.s (CU).

Briti.sh Columbia: Chilliwaek, \'I-I5-53, G. ]. Spen-
cer, 1 female (CNC).

Colorado: Bellvue, 13 mi. \\'., Buckhorn Mts.,
8500', VI-22-66, S. G. Wellso, 1 male (TAM).

Connecticut: Canaan, \'l-12-28, L. B. Woodruff. 2
males, 2 females (AMNH); Cornwall. l\\ V, VI, VII-28,
29, 8, 15, 6, 10, 11-20, 21, 22, 24, Chamberlain, 4 males,
3 females (CU), 3 m;des, 2 females (CAS), 1 male
(USNM); Littlefield, V-30-13, L. B. Woodruff, 1 male
(AMNH); New Ha\en, V-23-19. Chamberlain, 1 male,
1 female (CU); Westport. V-28-31, L. Lacey, I female
(BVU).

Illinois: Hebron, VII-29-,52, C. E. White, red clover,
I m;de (INHS); Lombard, VII-29-52, C. E. White,
nii.xed red clover, alfalfa, rat^veed, 1 male (INHS);
Plainfield, \'1 1-30-52, C. E. White, red clover, 1 female
(INHS); YorkviUe, VII-30-52, C. E. White, red clover.
1 female (INHS).

Indiana: Decatur, F. W. Poos, red and white clover,
3 males, 1 female (USNM).

Maine: Cumberkuid Co., VI, VII-I, 26-16, A, Nico-
lav, 3 males ( BVU ); Bridgton, VIII-20-34, M. E. Griffith,
1 'female (UK); Lincoln Co., VIII-20-40, D. J. Borror,
1 female (OSC); Medomak, VII-4-38, 1 male (OSC):
Millinocket, VII-27-30, C. G. Siepmann, 3 males, 3 fe-
males (OSU); Orono, VIII-19-18, H. Osbom. 1 female
(OSC); Weld, VII-2-51, A. Stone, 1 fem;de (USNM).

Marvland: Montgomery Co., Great Falls, VI-2.5-63,
D. C and K. A. Rentz, 1 male (CAS); Raspeburg, IV-
14-43, Schaeffer, red clover, 5 males, 2 females,
(USNM).

Massachusetts: 1 male (BYU); Ashland, VI-18-5I,
C. A. Frost, I male, 1 female (ISU); Fall River, VI-1,
20-19, 34, N. S. Easton, 3 males, 3 females (MCZ);
Harwichport. VIII-33, L. Lacev, 1 male, 2 females
(BVU); Hopkinton, VI-1-13, 1 mde, 1 female (BVU);
Marblehead, VIII-26-30, H. Dietrich, 1 female (CU);
Salisbur\-, VI-1 1-28, H. Dietrich, 1 male, 2 females
(CU); Sherbom, VI-6-2.5. C. A. Frost. I male, 1 female
(BYU); Sherbom, VI-1, C. A. Frost, 2 fem;iles (PANS);
Wilmington, \'I-26-20, C, C, Speery, 1 female (USNM);
Woods Hole, VII-11-19. L. L. Buchanan. 1 female
(CU).

Michigan: Shebovgan, VII-14-4I. H. B. Hungerford,
1 male (UK); Shebovgan, VII-1-42, E. L. Todd,' 1 male
(UK); Shebovgan, Vn-2-51, D. M. Anderson, 1 female
(CIS); VII. 2.3, 27, 29-51, E, P. Marks, 23 males, 22
females (CIS); Ingram Co.. Vn-23-47, 1 male (USNM);
Missaukee Co., VII-14-45, R. R. Dreisbach, 1 female
(UA).

Minnesota: St. Paul, VI-19-48, H. E. Milliron, red
clover, 1 male (USNM).

New Bnmswick: Halcomb. VIII-9, 11, 14-51, E. E.
Gilbert. 2 males, 4 females (CIS).

New H.mipshire: Mt. Washington, VII-6-14, C. A.
Frost. 1 male (INHS); Peabodv River, White Mts„ VII-
11-2.5, A Nicolay, 1 male (USNM); VII-11-25, E. D.
Quirsfeld, 2 males, 3 females ( UA ) ; Valley Meadow,
White Mts., VII-I 1-25, F. R. Mason. 900'. 2 males, 2
females (PANS).

New Jersey: Haddon Hts., IV-29-35. L. J. Botti-
mer, 2 males, 2 females (CNC); Irvington, A. Bischoff,
1 male (USNM), 1 male (AMNH); Montclair, E. D.

(,)uirsfeld, 2 males, 1 female (CAS), Bi.schoff. 1 female
(AMNH); Pahsades, VI-22-39, Malltin, 1 male, 2 fe-
males (FMNH); Phillosburgh, V, VI-12, 20-17, 31, J.
W. Green, 1 male, 1 female ( CAS ) .

Nova Scotia: Digby Co., VI-27-58, C. V. Reichart.
1 male (OSC); Sidney Mines, VI-19-65, W. J. Brown,
1 male, 1 female (CNC).

New York: Austerlitz, VI-2.5-34, H. Dietrich, 1 male
(CU); Bear Lk., VI-2-40, 3 females (FMNH); Bridge-
port, V-20-14, 8 males, 7 females (USNM); Canton,
\'I-19-25, Bablv, 1 female (CU); Crosby Landing, VI-
26-14, L. Keuke, 1 male (CU); Croton Falls, IV-26-40,
1 male. 2 females (FMNH); Crown Pt., VI-26-34, H.
Dietrich, 1 female (CU); Greenport, VII, VIII-63, R.
Latham, 1 female ( CU ) ; Hancock, VI-18-34, H. Diet-
rich, 1 female (CU); Ithaca, Vn-8-07, 1 male (FMNH),
V, VI-18, 2-14, 1.5, 2 males, 1 fem;de (AMNH), VI-2-
15, 5 m;des, 5 females (BYU), V-.30-14, 1 male
(USNM): III, L\-14-20, 6 m;des, 8 females (CU), V.
VII-3, 24-17, 19. H. Dietrich, 2 females (OSU), 1 fe-
male (CU), VIII-31-1.5, C. W. Leng, 2 females (BYU);
Renwich, VI-2-19, 1 male, 1 female (CU); H. Morrison,
VI-1-13, 1 female (TAM); Courtland Co.. Labrador
Lake, VI-4-38, J. C. Bradley. 1 male, 1 female (CU);
Cape Hopafrieng, VI-9-40, 3 females (FMNH); Lan-
caster, VII-25-46, L, D. Beamer, 1 male (UK). McLean,
\TI-2, 3-04. 1 male (CU); Tompkins Co., McLean
Bogs, V-30-19, H. Dietrich, 1 female (CU); Minetto, VI,
VIII-1-52, A. A. Muka, 36 males, 41 females (CU);
6 males, 20 females (OSU); VIII-52, 1 female (USNM);
Oliverea, VI-18-34, H. Dietrich, 2 males, 4 females
(CU); 0,swego, VII-2, 16, 19-1896, 4 males, 5 females
(CU); Paulsmith, VI-19-25, Bably, 2 males (CU); Pel-
ham, VI-7, 2-30, 34, Lacey, 1 male, 2 females (BYU);
Penn Yen, VII-12-25, Bably, 1 female (CU); Perry,
VII-31-19, 1 male. 1 female (CU); Clinton Co., Peru,
VI-10-16, 3 females (CU); Peterburg, VI-25-34, H.
Dietrich, 1 female (CU); Phoenicia, VI-30-35, J. W.
Green, 1 female (CAS); Port Jarvis, VI-6-56, M, Plavter,
alfalfa, 1 female (CU); Pula.ski, VI-20-25, Bably, 2
females (CLI); Rochester, V-14, M. D. Leonard, 1 female
(LA); 2 m;iles, 2 females (CU); Salem, VI-26-34, H.
Dietrich, 2 females (CU); Slaterville, V-27-38. J. C.
Bradlev, 1 female (CU); Sonyea, VI-22, 1 male (CU);
Cavuga Co., Springlake, VII-23-I8, 1 male (CU); Stoney
Ishuid, VII-8-96, 2 females (CU); Staatsburg, VI-23-34,
H. Dietrich. 2 males, 1 female (CU); Ticonderoga, VII-
3, F. R. Mason, 1 male (PANS); Tuxedo, V-26-40, 1
female (FMNH); Van Cortland Park, V, VI-9, 23, 26-
39, 2 males, 3 females (FMNH); West Point, VI-3-12,
W. Robinson, 1 female (CU).

OHIO: Adams Co., VIII-20-67, R. and L. Hamilton,
1 female (OSC); Clinton Co., VI-IO-61, F. J. Moore, 2
males, 2 fem.iles (OSC); Columbus, VI-8-64, H;imilton
and Bhick, 2 females ( OSC ) ; Wayne Co., Daes, V-3-60.
alfalfa and clover, 1 male (OSC); Delaware Co., IV, V
VII, VIII-13, 2, .30, 4, 9, 56, 65, 66, 67, 68, R. and L
Hamilton, 6 males, 9 females (OSC), VII-4-66, E
Sims, 1 male (OSC); Frankhn Co., V-10-67, R and L
Hamilton, 1 female (OSC); Greene Co., VI-2-59, D. J
and J. N. Knull, 1 male (OSC); Highland Co., V, VI-2
18, 3-61, 67. 68, R. and L. Hamilton, 3 males, 1 female
(OSC); Hocking Co., V-30-64, Hamilton and Black, 1
female (OSC); V-4-68. R. and L. Hamilton, beaten
from Prunus virginiana, 1 male. 1 female (OSC); V-2-
.57, D. J. and J. N. Knull, 1 m;ile (O.SC); Clear Fork
Valley, VI-5-66, R. and L. Hamilton, 1 male (OSC);
Licking Co., VII-30-47, Ladino red clover, 5 males, 3
females (OSU): Pike Co., V-12-63. R. E. White, 4

12

Brigham Youno University Science Bulletin

males, 1 female (OSC); StrongsviUe, VI-30-20, W. H.
Larrimer. 1 male, 1 female (USNM); Vinton Co., 1V-1.>
67, R. aiul L. Hamilton, 1 female (O.SC); Wood Co.,
\'II-30-47, red clover, 2 males (OSU); Madison Co.,
V-27-67, R. and L. Hamilton, 1 male, 1 female (OSC).

Ontario: Prince Edward Co., V-14, 23-20, 21, Hrim-
ley, 4 females (UK), 1 male 9 females (CAS); Ottawa,
V-2()-.5() H. F. Howden, 1 male, I female (CNC); VI-
18-16, I male. 1 female (CU); Rideau Lk., VII-17, V.
H. M.ison, 1 male (PANS).

Pennsylvania: Downington, Vn-4-35, L. J. Bottimer,

1 male (CNC); Duncannon, V-8-40, F. W. Poos, 1
female (USU); Easton. VI, VII-.3, 4-30, 26, J. W. Green,

2 females (CAS); Effort, VI-6-31; J. W. Green, 1 fem;de
(CAS); Creentown, Vl-16-20, I). E. Quirsfeld, 1 male,
6 females (UA); Hnmimlstown, \'-20-2.5, J. N. Knull, 1
m;de (PANS); New Hope. V-.30-3.5, L. ]. Bottimer. 3
m;des 1 female (CNC); Nottingham, V-10-.36, I,. |.
Bottimer, 1 male (CNC); Milford Pike Co., V, Vl-3(),
1-41, B. M;dkin, 3 fem;des (FMNH); North East, VI-
11-17, R. H. Cnshman, red clover, 1 female (USNM);
Snyder Co.. I.\-4-41, J. O. Pepper, clover .seed heads. 2
females (USNM); Spring Bridge, V-26-45, 1 male
(USNM); Wilawana, VI-12-39, R. H. Crandall, clover,

2 females (UA); Wind Gap, V, VI-28, 18-31, J, W.
Green. 7 males, 4 females (CAS).

Quebec; Avlmer, V-31-28. W. ]. Brown, 1 m.ile, 1
female (UK); VI-19-36, G. Stacesmith, 3 males, 5 fe-
males (CAS); Chelsea, Vl-20, 2.5-16, 1 male, 1 female
(CU); Covev Mill, VI-27-24, C, E. Fetch, 1 nwle
(CNC); Depanpiet, V-27-44, G. Stacesmith, 1 m;ile, 1
female (CAS); Ga.spe, 25 mi. W., VI-22-54, W. J.
Brown, 1 male (CNC); Georgesville, VI-23, .36, G. S.
Walley. 1 female (CNC); Hull, VM9, 23-16, 1 female
(CU); V-31-.54. W. J. Brown, 2 males (UK); 1 m;ile,

3 females (AMNH); Megantic, VlI-6, 7-16, 1 fem;de
(CU); Laurentian Mts,, Montfort, Vl-30-16, 1 female
(CU); Montreal, V-31-19, E. S. Ros.s, 1 male (AMNFI);
Perkins' Mills, VI-23-36, G. Stacesmith, 1 male, 1 female
(CAS); Sherhrookc. VII-,5-16, 1 male (CU); Ste. Anne's,
VI-12-1.5, Webster, 1 male (USNM); St. Lambert, VII-
4-27, W. J. Brown, 1 female (CNC).

Rhode Island: VI-7-51, red clover, 2 males
(USNM); Arawan Cliffs, VII-3-,50, C. V. Reichart. 1
male (OSC).

Vermont: Chelsea, \'I-16, H. E. Smith. 1 male
(USNM),

Virginia: Arlington, IV-9-37, F. F. Dicke. 1 male
(USNM); 1 female (OSC).

Washington: Bellingham, VI-4-45, M, J, Forsell, red
clover, 1 female (USNM), 3 mi. N., III-3-60, G. G.
Scudder, 2 females (OSU).

Wisc-onsin: Racine Co.. Dover, VII-10-66, alfalfa, 1
male (USNM); Walworth Co., Geneva, VIll-4-66, alfal-
fa, 1 female (USNM); Green Co., Jefferson, VIIl-4-66,
alfalfa, 1 female (USNM); Kanosha Co., Randall, VII-7-
66. alfalfa, 1 female (USNM).

Total specimens examined: 547

Discu.ssion. The nomenclature of two clo.sc-
ly related weevil pe.sts of cultivated clover wa.s
the subject of a paper by Milliron (1949). He
detennined that one of the .species wliicli pos-
esscs seven antenn;il funicular segments belongs
to the genus Tijchim. The correct name of this
species was determined to be T. stephensi
Schoenherr. He stated that the other species
which possesses six antennal funicular segments

belongs to the genus Miccotwgiis. The correct
name for this species was deteniiincd to be M.
piclrostrw (Fabricius), Since tiieii these names
have been in use for the two weevils in the
literature of North American economic ento-
mology.

Nlilliron's determination of the nomenclature
of these species was made without recourse to
the type specimens. During the course of this
revision I haxe examined the types and other
iiiatirial which relate to this problem. These
were borrowed from the European museums
in which they are preserved. The identity of
other type specimens has been ascertained
through correspondence with Dr. l\. T. Thomp-
son of the British Museum (Natural History)
and Per Inga Persson of the Stockholm Museum
of Natural Histor)'. Examination of this material
has revealed that the current ;ipplication of the
two names in question is incorrect.

For convenience of discussion the s\aionymy
revealed by reference to the types is listed be-
low. The names listed under Tt/cJuus conform
to the current concept of T. stephcnsi Schoen-
herr. Those listed under Miccotrogus confomi to
the current concept of M. jiicirostrLs ( Fabricius).

Tijchius

Ciirctilio jiicirosfris Fabricius, 17S7
Ctirciilio fuscirostris Paykull, 1792?
Ctirculio toniento.sus Herbst, 1795
Tt/cliius stejihensi Schoenherr, 1S.36
Tijchius griscus Schaeffer, 1908

Miccotrogus

Cttrciilio cinerascens Marsham, 1802
Tt/chiiis posticus Gyllenhal, 18.36

Dr. Thompson reported that specimens of T.
stepehensi from the British Museuin (Natural
History) and the type of Curctilio j)icirostris
Fabricius in the Copenhagen Museum were com-
pared bv Dr. B. D. \'alentine at Dr. Thompson's
re(|uest and detcmiined to be conspecifie.

According to Persson there are no specimens
in the Paykull collection at the Stockholm Mu-
seum of C. fuscirostris Paykull. Paykull (1800)
lists fuscirostris under C. picirostris. Apparently
Paykull thought that the name was incorrectly
;»pplied and either removed the specimen or
specimens from his collection or placed them
with his specimens of C. picirostris. 1 have ex-
amined a series of five specimens labeled C.
l)icirostris from the Pavkull collection. Tliese all
confomi to the surrent concept of T. stcjilicusi.
1 can iiiul no evidence for linking fuscirostris

Biological Series, Vol. 13. No. 3 Revision of the Weevil Genus Tychius

13

with Miccotrogus uncliT wliicli it is li.stccl bv
Klimu (1934).

The lectotype designatccl ahoxe for T. to-
mentosus Herbst i.s a female, the first specimen
of a series of eight syntypes recei\cd from the
Zoologische Museum der Humboldt-Universitat,
BerUn. This specimen and the second, third,
fifth, seventh, and ei<rlitli conform to the current
concept of T. stephensi Schoenherr. The fourth
is a Ti/chitis which is unfamiHar to me and the
sixth confonns to tlie current concept of A/. j)ici-
rostris { Fabricius). The name tomentosus is in
current use in Europe for the Tijchius species but
Milliron (1949) rejected it because it is a junior
homonym of CunttUo tomcntoms Olivier, 1790.

Schoenherr (1S.36) gave the name T.
stepheni to the species described by Stephens
( 1S31 ) as T. tomentosus. Schoenherr apparently
considered it to be a new species only on the
basis of Stephens" description and had no speci-
mens in his collection. According to Thompson
there are nine specimens identified as T. tomen-
tosus in tlie Stephens' collection. The first of
these which I ha\'e examined bears the label by
the late Sir Guy Marshall: "Type of T. stephensi
Schonh. ( em ) 1836." I have designated this
specimen as lectotvpe of T. stephensi. According
to Thompson ail of the series agree with the cur-
rent concept of T. stephensi except the fifth,
which is an EUeschus bipunctatus (L.), and the
sixth, which agrees with the current concept of
M. picirostris ( Fabr. ) .

Schoenherr's original spelling of the name was
stepheni. This does not ([ualifv as a lapsus cahi-
mus as it is also spelled stepheni in the inde.x to
his 1S36 work. Stephens ( 1S39) was the first to
use the spelling stephensi which is in current
use today.

1 also examined the type of T. griscus
Schaeffer at the U.S. National Museum. There
is no (juestion on its svnonymy with T. stephensi.

The onlv types which were found to agree
with the current concept of Miecotrogus pici-
rostris were those of Curculio cincrascens
Marsham and Tijcliius j)0sticus Gyllcnhal. The
identity of cinerascens was confirmed by
Thompson \\ho states that its tvpe is in the
Stephens collection. I examined the type of T.
posticus from the Stockholm Museum.

Thompson also checked the type of C. villo.ius
Marsham which Klima (1934) lists in synonymy
with T. tomentosus. Tlie tvpe is in the Kirby
collection and is a Sihinia potentillae Germar,
under which species it is also listed by Klima.

The earlv uorkers knew the identity of
Fabricius" C picirostris. I have examined the
specimens described bv I'avkull (1792:253) as

C. picirostris Fabr. These conform to the cur-
rent concept of T. stcplwnsi Schoenherr. Gyllen-
hal ( 1813:121) considered his Rhijnchaenus pici-
rostris to be the same as R. picirostris Fabr. 1801,
Pavkull's C. picirostris, and C. tomentosus
Ili'rbst. Germar (1817:340) cited tomento.sus
Herbst in synonymy with i)icirostris. Stephens
(1829:160) listed Paykulls j)iciro.stri.i as synony-
mous with R. picirostris Gyllenhal and later
(1839:228) listed tomentosus Flerbst, T. steph-
ensi Schoenherr, and Paykulls i)icirostris as
synonyms.

The association of Fabricius" picirostris with
the name Miccotrogus came about as the result
of a mistake made by Schoenherr. Gennar ap-
parently did not consider C. picirostris Fabr. 1787
to be the same as R. picirostris Fabr. 1801. He
(1824:291) associated R. jiicirostris with the
generic name Sihinia and listed R. picirostris
"\'ar. b" Gvllenhal in synonymy. The following
year Schoenherr (1825:583) listed Gyllenhals
j)icirostris under Ti/chius and under his newly
established subgenus Miccotrogus listed Sibinia
picirostris Gennar and R. jiicirostris "var. Gyll"
( presumably referring to the "var. b " ) thus as-
sociating the specific name picirostris with Mic-
cotrogus for the first time, Schoenherr (1836:
411) then correctly associated Paykulls pici-
rostris with Gyllenhals jiicirostris "var. a" and
then listed Gyllenhals picirostris "var. b, " which
lie considered to belong to Miccotrogus, in syn-
onvmy with C. picirostris Fabricius. Later work-
ers and catalogers copied Schoenherrs error thus
establishing the usage of Fabricius" C. picirostris
for the Miccotrogus species instead of the Ty-
chius species to which its type belongs.

Apparently Fabricius' Cursulio picirostris and
his Rhifnchaenus picirostris are not the same
species. Dr. Thompson reports that Dr. Valentine
saw a specimen in the Fabricius collection, la-
beled Rhijnchaenus picirostris. He noted that
this specimen was a tvchiinine, but "much larger
than T . stephensi."

1 have examined Gyllenhals specimens of
R. picirostris including the "var. b" from the
Gyllenhal collection at Uppsala, Sweden. There
are 46 specimens of "var. a,"' all of which con-
form to the current concept of T. stepliensi. Of
the series of ten specimens designated as R.
jiieirostris "var. b" nine are T. .stephensi. Only
one conforms to the current concept of M. pici-
rostris (Fabr.).

According to the synonymy revealed in this
study the name picirostris Fabricius should re-
place stephensi Schoenherr for the Tychius spe-
cies described above. The name cinerascens
should replace picirostris for the Miccotrogus

14

Brigham Young University Science Bulletin

species. 1 lia\e decided to retain the current
usage of the names in (juestion; however, since
I do not consider that the changes indicated
would lie in the interest of stabihty of nomen-
clature. I intend to appeal to the International
Commission on Zoological Nomenclature to use
its pK-nar\' powers to such extent as may be
necessary to pro\ide a valid basis for the con-
tinued use of the names Tijcliius stephensi
Schoenherr, 1836, and Miccotrogus piciwfitri.s
(Fabricius, 1787) as they are currently applied.

Tijcliius sordidus LeConte
(Figs. 17, 20)

Ti/r/iii/s sordidus LcCoiito, 1876. Proc. Amcr. Philos, Soc,
13:217 (Holotvpc: male, Illinois, MCZ type 5232);
Casev, 1892. Ann. New York Acad. S'ci., 6:414;
Sanderson. 1904. Tex. Agr. E.\p. Sta. Bnll., 74:.3-1.3;
Hunter and Hinds. 1904, U.SDA Bur. Entomol. Bull..
51; Mitchell and Pierce, 1911, Proc. Entomol. .Soc.
Wash., 13:45-62; Pierce, 1907, Entomol. News. 18:
362; Pierce, 1907, Stud. Zool. Lab. Univer. Xebr..
p. 273; Casev, 1910, Can. Entomol.. 42:134-135;
Pierce, 1912,' USDA Bur. Entomol. Bull., 100:77;
Blafchley and Leng, 1916, Hhynchopora or weevils
of northeastern America, p. 245; Frost, 1945, J.
New York Entomol. Soc, 53:221.

Ti/chius nimius Casev. 1910, Can. Entomol., 42:134
(Holotype: male,' Iowa, USNM 36751, T. L. Casey
collection ) .

Tychius texmws Casey, 1910, Can. Entomol., 42:134
(Holotvpe: female. Haw Creek, Texas USNM
36752, T. L. Casey collection).

Tijchius carolinae Casey, 1910, Can. lOiitomol., 42:134-
135 (Holotvpe: female, Southern Pines, North
Carolina, IX, A. R. Manee, USN.M 36750, T. L.
Casey collection ) .

Tychius- sordidus carolinae: Blatchle\- and Leng, 1916,
Rhvnchophora or weevils of northeastern America,
p. 245-246.

Miccotrogus sordidus: Klima, 1934, Coleopterorum Cata-
logus, 29(138): 32.

This is the largest North American species.
It can be distinguished from other North Ameri-
can .species by its size, its gray or vellowish
gray color, its obese shape, and the shape of the
pronotum which is wider at the base than at the
apex. It closely resembles T. caesius and T.
lineeUtts. From the former it can be distinguished
by the broad prothorax and the absence of a
minute t(X)th on the pro- and mesofemora;
from the latter by the absence of a triangular
median tooth on the protibia and the absence of
the apieal projection of the median lobe of the
male genitalia (Fig. 17).

Notes on the biology of this species are given
by Sanderson (1904)' Pierce (1907a, 1907b,

1912), Mitchell and Pierce (1911), Blatchley and
Leng (1916), and Frost (1945).

Description. Male: length .3.0-4.9 mm, 1.8
times longer than wide; integument shining
black on dorsum often piceous to black on ven-
tnd surface; appendiiges dark reddish brown.
V'estiture of gray to yellowish gray scales often
with metalic bronze margins.

Rostrum shorter than prothorax; from lateral
;ispeet nearly straight to imtennal insertion then
tapered slightly to apex; in dorsal aspect wider
at apex than frons between eyes; dorsoventrally
flattened distad of aiitennal insertion, without
dorsal depression between scrobal apices, rugae
deep. Vestiture sparse, composed of long, nar-
row, apically truncate scales; apical portion
glabrous except for row of bristles extending
nearly to apex from bene;ith apical portion of
scrobe distad of antennal insertion.

Antennal funiculus seven segmented, pedicel
longer than next two segments combined.

Prothorax 1.2 times wider than long, widest
at base, base more than twice as wide as apex
from dorsal aspect. Scales on dorsum of uniform
size, shape and color, long, narrow with rounded
apices, broader than scales on elytra; scales on
lower half of sides round to elongate-oval.

Elvtra with sides broadly rounded, widest
just before middle; strongly convex in dorsal
profile. Scales on dorsum of same shape and color
as those on pronotiuii; usually denser on inter-
space one but scales of other interspaces of sim-
ilar size and density. Strial scales slightly, if at
all, narrower than scales on interspaces. Inter-
spaces nine and ten with rounded scales similar
to those on venter.

\'entral surface clothed with dense, imbri-
cated, round to elongate-oval, white or light
gray scales.

Femora stout, especially apically; ventral,
apical emargination prominent , usually with
sm;dl tooth on posti'rior portion of emargina-
tion. Se;des of two distinct t\pes, long, narrow,
strigose scales and broad scales with rounded
sides.

Tibiae iiiucion;ite, inucrones on protibia
slightly larger than on mesotibia and metatibia.
\'estiture of long, narrow, strigose scales, and
\eiy fine hairlike setae.

Tarsi with long, narrow scales and fine hair-
like setae on dorsal surface. Claws long, diver-
gent, bas;d processes convergent.

Male genit;dia (Fig. 17) with apical portion
ol median loix' slightly anguhite; apical, dorsal,
median membranous area nearly round, strongly

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius

15

defined posteriorly; median striit.s verv fine, not
clavate.

Female: length 3.7-5.3 mm, ro.strum sliglitly
longer and more slender, especially distad of
antennal insertion, antennal insertion median.

Hosts. Bapt'ma JeucantJui, and B. hracteata
(Blatchle\ and Leng, 1916:24,5; Pierce, 1907-a:
273; Pierce, 1907-b:362; Frost, 1945:221), B. cti-
mata (Mitchell and Pierce, 1911:61-62); B.
villosa. Also recorded from Acerates and Croton.

Distriution. (Fig. 20)

.Arkansas: 1 male (USNM): "southwesteni," Palm,
2 males, 1 female (AMNH).

Illinois: 2 males (USNM); F. Blanchard. 2 fe-
males (MCZ); Liebeck, 1 male (MCZ); Paiia, VII-2()-
38, J. H. Bigger, Acerates, 7 males, 5 females (INHS);
"southern," 1 male (PANS); F. C. Bowditeh. 1 male,
1 female (MCZ).

Iowa: W. G. Dietz, 3 males (MCZ): Horn, 1 male,

1 female (PANS); Burlington, Liebeck, 2 males, 1 fe-
male (MCZ); Ft. Madison, 2 males (UK).

Kansas: 2 males, 3 females (PANS); Douglas Co.,
F. H. Snow, 1 female (UK); Kansas Co., Liebeck, 1
male, 2 females (MCZ); Chautauqua Co., Niotazi, 2
mi. E., VI-3-68, D. R. Harris, 3 males, 1 female (WEC);
Onaga, \'-20-01, F. C. Bowditeh. 1 female (MCZ), VI-
27-03, Crevacoeur, 1 male, 1 female ( UK ) , 1 female
(USNM); Jefferson Co., 8 mi. N. Lawrence, VII-8-6.5,
J. B. Karen, 1 m;ile (CWO).

Louisiana: Logansport, (Pierce, 19()7-a:273; 1907-b:
362), Natchitoches, III-28-07, Cushman and Pierce,
Baptisia leucantlui, 8 males, 9 females (USNM); Natch-
itoches, III-28-07, Pierce, Btiptistii lillusii, 1 male, 1 fe-
male (USNM).

Michigan: Adrian, Liebeck, 1 male, 1 female
(MCZ).

New Jersey: Cape May Co., Woodbine, 1 mi. E..
VI-21-66, D. G. Kissinger, Baptltia, .5 males, 2 females
(WEC).

New York: Bellport, Long Island, VII-I8-14. A.
Nicolay, 2 males (BYU).

North Carolina: Southern Pines ( Blatchlev and
Leng, 1916:246).

Oklahoma: Okfuskee Co.. VI-31-.34, Hinton, 1 m;ile
(BYU); Stillwater, V-3-31, H. Whitaker, 2 males, 3
females (BYU).

Te.vas: 2 males (INHS); F. H. Chittenden, 2 males,

2 females (USNM); Horn, 1 female (PANS); Liebeck,
2 males (MCZ); Brazos Co., College Sta., III-I8-64, J.
C. Schaffner, 9 males, 6 females (TAM), lV-3, ll-7().
on Bapfmu, 62 males, 4.5 females, W. E. Clark (WEC),
I\'-27-,50, H. J. Reinhard, 1 male (TAM); Colorado Co.,
IV-7-22, G. Wilev, 1 male (UK); Eagle Lake, IV-12-
1899, A. M. Wangh. 1 male (USNM );"Edna, III-24-07,
J. D. Mitchell, 1 male, 3 females (USNM); Anderson
Co., Elkhart, 10 mi. S., III-27-67. H. R. Burke, 3 fe-
males (TAM); Grand Saline, 111-25-04, W. D. Hunter.
1 female (USNM); Houston, IV-1-04, G. W. Curtis, 1
female (USNM); Jackson Co., III-2.5-07, J. D. Mitchell,
1 male (USNM); Keechi IV-4-22, 1 male (TAM):
Kirbvville, 111-20-08, E. S. Tucker, Cro(o)i and Bapti.siii.
6 males, 7 females (USNM); Leon Co., IV-IO-48, J. L.
Ward, I male (USNM); Maud. IV-29-41, D. J. and [.
N. Knull, 2 males (OSC); Panola Co., IV-I5-05, J. John-
.son, Baptisia, 2 males, 4 females (USNM); Swiss Alps,

III-24-1899, Hubbard and Schwarz, wild pea, 5 males,
10 females (USNM); Tenaha, III-23-08, E. S. Tucker,
Buptma, 3 males, 1 female (USNM); Timpson, III-25-
08, E. S. Tucker, Baptisia, 2 males, 2 females (USNM);
Victoria, III-25, 29, 30-05, W. E. Hinds :md E. S.
Tucker, Baptisia and Flowers of "Bull \\'eed," 5 males,
5 females (USNM); Whitewright, IV-15-08, J. W.
Henry, 3 females (USNM); Y'oakum, III-27-1899, Roos
Bros., Baptisia hracteata, 2 males, 8 females (USNM);
Favette Co., LaGrange, III-30-70, on Baptisia, W. E.
Clark. 4 males, 5 females (WEC).
Total specimens examined: 307.

Discussion. Specimens of this species from
adjacent locahHes or from the same series may
be entirely gray in color or have several scales
with bronze margins giving a general yellowish
line. Specimens from the eastern and southern
portion of the range average smaller in size than
those from the northern and western portion.
Specimens from New Jersey averaged 3.65 mm
in length, those from Arkansas 3.85 mm, Louisi-
ana 4.30 mm, Te.xas 4.30 mm, Iowa 4.40 mm,
Michigan 4.60 mm, Kansas 4..55 mm, and Illinois
4.70 mm.

Tychius caesius, new name

(Figs. 15, 20)

Tychius armatiis Green (not Touniier, 1873), 1920,
Entomol. News, 31:198 (Holotype: female, Gray-
beard Mountain, North Carolina, CAS).

Sihinia armata: Klima, 1934, Coleopteronim Catalogus,
29(1.38);45.

This species appears to be most closely re-
lated to T. sordichis. It can be distinguished from
other North American species by its gray vesti-
ture; stout, short rostrum which is slightly wid-
ened at the antennal insertion and prominently
tapered from the antennal insertion to the tip;
and the toothed femora.

Description. Female: Length 3.0-3.6 mm,
2.0 times longer than wide; integument shining
black, appendages and antennae dark reddish
brown. Vestiture of bluish or yellowish gray
scales.

Rostrum as long or shorter than prothorax, an-
tennal insertion in apical third, slightly wider at
antennal insertion than frons between dorsal
margin of eyes, from lateral aspect prominently
evenly arcuate; pits and rugae distad of antennal
insertion deep, especially on dorsum between
apices of scrobes. Vestiture proximad of antennal
insertion of long, narrow scales, nearly glabrous,
distad of antennal insertion. Eye nearly round,
gold in color.

•Antennae with seven funicular segments;
pedicel equal in length to next two segments
combined.

16

BiUGii\.\r YoiTNG Univkksity Sciknce Bulletin

Pronotum 1.2 times wider tlian long, from
dorsal aspect nearly parallel sided in basal half,
narrower than elytra at base. Scales on dorsnin
elongate, broader than scales of elytra, usually
apically rounded; sides with elongate-oval, white
scales.

Elvtra 1.1 times longer than wide; nearly
panillel sided in b;is;d two thirds, widest just
l)ehind humeri, rounded broadly to apices; even-
ly, broadly roinided in dorsal profile. Scales on
interspaces long, narrow, denser on interspaces
one, five, and seven. Strial scales narrower than
scales on interspaces.

Ventral surface sparsely covered by noiiiiii-
bricated, oval, plumose margined scales; integu-
ment finelv \isiblc- bi'tween sc;iles; no erect
setae. Sternum live usualK- with dcvp median
fovea.

Femora toothed on posterior portion of ven-
tral apical emargin;ition. tootii large and promi-
nent on metidemur, sm;dl or very minute on
mesofemur and profemur; ventnd apic;d emar-
gination very prominent. Vestiture ot long, nar-
row scales, sometimes with sp;irse oval scales on
pro.ximal portion.

Tibiii mucronate. inucroiics on protiliia larg-
er than on mesotibia and metatibia; scales long,
narrow, verv fine, liairlike, toward ape.x ot tibi;i.

T.irsi clotlied dorsalK with long, narrow
scales and fine hairlike setae; claws short, di-
vergi'nt, basal proc-esses parallel or slightly con-
vergent.

Male: length 2.7-.'1.5 mm; rostrum shorter
than protlior;i.\. stout; anteiin;d insertion in
apical fourth; pits and rug;K' on distal portion
ver)' deep.

Male genitiilia (Fig. 15) with apical portion
of median lobe rounded; ;ipic;ii, dors;d, median
membnmous ;uea elongate-o\-al, sharplv defined
posteriorly; median struts narrow, moderately
clavate.

Host. Unknown.

Distribution. (Fig. 20).

North Girolina: Black Mts., \ 1, \ 11-02, X'.iti D\kc,
52 males, 29 females (CAS), V-1,5-12, Beutenmulkr. I
male (CAS); Mt. Graybeard, V, Vl-26, 9, 19-04, 25,
26 males, 21 female.s (AMNH), 1 female ( Paratvpe
.>»993, U.SNM), V-15-I2, \V. Beuteiimiiller, 8 male.s,
I females (CAS); VI-02, R. C. Van Dvke, 7 males, 5
females (C.\S).

South Carolina: Hoekv Bottom, V-22-,34. J. A Bervl
1 male (USNM).

Total specimens e.vamiiied: 1.5,3.

Discussion. Green states that the "tvpe" is ;i
m;de, but, the type specimen examined is def-
initcK female.

'iijcliius lincclhis LeC'onte
(Figs. 16, 21)

Ti/chius linccllit.s- l.eCoiite, 1876, Proc. .'\mer. Pliilos.
Soc, 15:217 (I.ecototvpe here designated: male,
California, MCZ type 5231); LeConte, 1881, Trans.
Amer. Entomol. Soc, 9:x.vii; Casev, 1892, Ann. New
York Acad. Sci., 6:412-413; Casey, 1910, Can.
F.ntomol., 42:132; Yothers, 1916, Bull. Wash. State
Agr. Exp. Sta.. 124:7, pi. 1, Fig. 8; Bruhn, 1947, Or.
Basin Natural.. 8:3, 18, Fig. 38 a & b (genitalia
described); Kissinger, 196.3, Ann. Entomol. Soc.
.\rner., 67(6):771 ( provi'niiculiis described).

Ti/rhius tm-itus Casey, 1910, Can. Entomol., 42:132
(Holol\pe: female, California "without more defi-
nite statement of locality," USNM 36745. Para-
types: 3 males, USNM 36745, T. L. Casey col-
lection ) .

Tijchitis licsperis Casey. 1910, Can. Entomol., 42:132-
133 (Holofvpe: female, Siski)ou Co., California,
USNM 36746. T. L. Casey collection).

Tychius radians Casey, 1910. Can. Entomol., 42:133
( Holot\pe: leniale, San Diego, Californi:i, USNM
36747, T. L. C.isey collection).

Ti/chius ililcctus Casey, 1910, Can. Entomol., 42:133
(Holotvpe: female, San Francisco Co., California,
USN.M' 36748. Paratype: 1 male. USNM 36748, T.
L, Casey collection).

'I'ljctiius probii.s Casey, 1910, Can, Entomol,, 42:133-
1.34 (Holotvpe: female, "near San Francisco," Cali-
fornia, USNM .36749),

Miccotrogus lineellus: Klima, 1934, Colcuptcwrum cata-
logus. 29(138): 30-31.

The prominent triangular tooth on the mid-
dle of the protibiii of the male and the projection
on the apex of the median lolx- ot the male geni-
talia ( 1' ig. 16), readily distinguish this species
from its North American relatives. The relatively
large size and grav or brownish grav color are
also chanictcristic. This is the onlv North .\meri-
can Ti/iliiu\ known to be associated with Lti-
;)/;i!/s.

Description. Mali': length .3.0-4.5 mm. \\ idth
1.4-2.1 mm; integument piceous to black, ap-
pendages reddish to orangish brown, scales either
entireh' grav in color or with combination of
grav ;uid bronze colori'd. often metallic scales.

Hostrmn shorter th;in prothorax, antennal in-
sertion in apic;il third; slightlv expanded at an-
tenn;d insertion, width ;it ;mtennal insertion equal
to or slightK' less IIkui widtii between dorsal
margin of eyes; ;ipical third dorsovenlndlv flat-
tened; in latend aspect slightlv tiipered from
anteimal insertion to extreme tip, slightly ex-
panded before eve; pits and rugiie verv deep,
especiallv dors;dly between ;intenn;d insertions
where slight depression between elevated lateral
c.irinae is often evident. N'cstiture of lontj, nar-
row, usually sparse scales, no erect setae, usu;dly

Biological Series, Vol. 13. No. 3 Revision of the Wee\il Genus Tychius

17

with .spursf tine hairlike setae around di.stal
portion of .scrobe.

Antennal fiinicle seven-.scgmented; pedicel
longer than segments two and three combined.

Pronotuiii as wide or wider than long; sides
rounded, 1.6-2.3 times \\ider at base than at
apex. Vestiture on dorsum of long, narrow,
apicallv truncate or acuminate scales; usually
with broad median and lateral vittae of slightly
wider scales; either all scales gray in color or
median and lateral vittae with gray and remain-
ing portion with bronze colored scales; integu-
ment usually clearly visible between scales.
Lower portion of sides with elongate-oval gray
scales.

Elytra in dorsal aspect parallel sided or
tapering slightly in basal two thirds, widest at,
or just beyond humeri, broadly rounded to
apices in distal third; dorsal profile usually
prominently convex but sometimes nearly flat in
basal third, broadlv rounded to apex. Vestiture
of long, narrow, apically truncate or acuminate,
recumbent scales; scales usually denser and
lighter in color on interspace one and alternate
interspaces; often alternate interspaces with
bronze colored scales. Strial scales narrower tlKin
scales on interspaces.

\'entral surface with recumbent, oval, often
plumose margined scales; usually with discrete
transverse rows of suberect hairlike setae on
each sternum. Sternum five without median
fovea.

Femora \\'ith prominent, apical, \entral
emarginations, often with minute tooth on basal
portion of emargination of metafemur. Scales
long, narrow, gray in color, usually longer and
pointed on \entral portions especially on pro-
femur.

Tibiae mucronate, mucro on protibia slightly
larger and stouter than meso- ;uid metatibiac;
protibi;i witli prominent median, ventral, trian-
gular tooth. W'stiture of fine setae, especially
fine apically.

Tarsi clothed with \er\' fine hairlike setae,
sparse on segments three and four, tarsal claws
long, divergent, basal processes convergent.

Male genitalia (Fig. 16) with apical portion
of median lobe constricted, fomiing narrowed
apical process; apical, dorsal, median membran-
ous area sharply defined posteriorly; medi;ui
struts stout, not strongly clavate.

Female: length 3.6-4.6 mm; rostrum more
slender and elongate than in male, pits and
rugae distad of antennal insertion shallow; an-
tennal insertion median. Sternum five with deep
median fovea. Tibiae with slightly smaller mu-

crones, protibia lacking median, ventral triangu-
lar tooth.

Hosts. Liipinus all/if rails. L. caiidattis, L. am-
inopltihis, L. arf^^cntctis, L. arJiorus, L. l)icaIor, L.
cliainissionis, L. excuhitus, L. leiicophijlhis, and
L. scriceus. also recorded from Burr Clover and
GiJia.

Distribution. (Fig. 21)

.Alberta: Lfthbridsie, V-30-33, B, M. White, 1 male
(CNC).

Arizona: Williams, IV-6, Barber :uid Schvvarz, 1
male (USNM); Fort Vallev, Coconino Co., Flagstaff,
I'A mi. N.W., VI-7-e4, R. W. Poole, 7350", 1 female
(CU).

British Columbia: Osovoos, V-30-58, H. and A.
Howden. 2 females (CNC);' Vernon, VI-2, .5, 31-21, 28,
K. Hoppint;, 9 m:iles, 4 females, (CAS), V-16-.53, J. E.
H. Martin,' 1200", 1 female (CNC), Venables, 1 female
(USNM).

California: ALAMEDA COUNTY: k'ocbele, 4 males,
4 females (CAS); 11-30-17, E. R. Leach, 1 female
(CAS); Berkeley Hills, N.E. Oakland, IV-8-64, P. Rude,
1400', 1 male (ClS): Oakland, VI-2-46, B. Adelson, 1
male (CIS); Hav"ward, V-21, 1 male, 1 female (CNC);
Havward. V-21-,30, F. E. Blaisdell, 7 m;des, 13 females
( CAS ) : Oakland, IV-8-06, E. C. Van Dyke, 5 males, 3
females ( CAS ) ; Oak Hills, IV-8-06, E. C. Van Dyke. 2
males, 6 females (CAS); BUTTE COUNT\': IV-29-39,
F. W. Nunemacher, 3 males (FMNH); Oroville, IV-30-
27, H. H. Kelfer, Lupinus alhifrotu; 2 males, 1 female
(CAS); Yankee Hill, V-8-28, H. H. Kelfer, 2 females
(CAS): CALAVERAS COUNTY: V-15-.36, 2 males, 1
female (ISU); Murphvs, V-14, 1.5, 18, 19-36, F. E.
Blaisdell, Alt. 2.500'. 15 males, 28 females (CAS), 1
male (PANS); Mokel Hill, V, F. E. Blai.sdell, 1
male (CAS); CONTRA COSTA COUNTY: Koebele,
I male (CAS); Antioch, III-29-56, B. J. Adelson, 1 male,
1 female (CIS), V-22-48, E. Ehrenford, 1 male (CIS),
III-31-33, G. A. Marsh, 2 males, 3 fem:iles (CIS), IV-5-
56. J. Powell, 1 male (CIS), IV-9-49, L. W, Quate, 1
male. 1 female (CIS), 11-26-39, ]. G. Shenafelt, 1 m;ile
(LA): Berkeley, V-33, E. S. Ro.ss, 1 male (CAS);
Orinda, V-4-34,' 1 male, 7 females (LA): EL DORADO
COUNTY: F. W. Nunemacher, 1 male (BYU); Placer-
viUe, V-20-13, 1 male (ISU); 3 males (CIS), F. H.
Wvmore, 1 male (CAS); FRESNO COUNTY: Coolinga,
l\'-8-51. E. G. Lindsay, 3 males, 3 females (CIS);
HUMBOLDT COUNTY: V-2, 3, 7-11, F. W. Nune-
m:icher, 14 males, 12 females (FMNH); Fieldbrook,
V-29-03, H. S. Barber, Lupinus, 4 males, 6 females
(USNM); Korbel, VI-16-16, F. E. Blaisdell. 1 male, 4
females (CAS); INYO COUNTY: Argus Mts., IV-91,
Koebele, 1 female ( CAS ) ; Independence, 2 males
(CAS); IV-19-19, Blaisdell. 2 males, 2 females (CAS),
VI, A. Fenyes, 1 male (CAS), IV, V-27, 19, 2-18, 19,
L. L. Muchmore, 12 males, 7 females (LA); Lone Pine,
V-26-.37, 2 males (LA); KERN COUNTY: GlennviUe,
V-7-31. A. T. McClay, 5 males, 6 females (CIS), 3
males, 2 fem:.les (CAS); Indian Wells. IV-19-62, E.
Lehre, 1 male (CIS), IV-18-62, C. A. Toschi, 3 males,
4 females (CIS); Isabella, IV-4-34, R. P. Allen, 1
female (CAS). R. Hopping, 1 female (CAS); Woody,
1 mi. E.. V-3-64, [. Powell, 1 male (CIS); LASSEN
COUNTY: Dovle, ^'-20-34, E. O. Essig, 7 males, 8
females (CIS)': LOS ANGELES COUNTY: IV, 10
males, 13 females (USNM): III-22-39, K. E. Stager, 1

18

BlUCIl AM YOUNC: Un1\ EHSITY SCIENCE BULLETIN

niiilf (LA); I\'-2-4(), 1 mall- (LA); Antelope Valley,
111-30-35, 1 male (CAS); Azusa, IV. 2 males, 2 females
(CAS); Dr. A. Feynes, 2 males, 2 females (CU); Cole,
Vn, 1 female (CAS); Durate, Wiekliam. 1 male. 1 fe-
male (USNM); Fairmont, lV-15-28, 1 female (CNC);
El Sef;uiulo, IV-27-38. D. Poole, Lupimif.- clwmisxioiiis,

1 female ( LA ) ; Laneiister, V, 1 male ( CAS ) ; Neenach,
\'- 17-28, J. O. .Martin, .5 male.s, 3 females (CAS); Pasa-
dena, I\', 1 female (CAS): Pasadena, C. Sehaeffer, 2
males, 2 females (BVU); Pomona. 1 female (INHS), 1
m;ile. 1 female (LA), 3 males, 2 females (MCZ):
MADFRA COUNTY: Coarsegold, V-26-42, C. Kennett,
Luiniws, 3 males, 1 female (CIS); MARINA COUNTY:
Fairfax, V-9-20, E. P. Van Duzce, 11 males, 8
females (CAS); Mill V;illey, IV-20-24, F. E. Blais-
dell, 3 males. 4 females (CAS), 1 male, 1 female
(CU), lV-21-24, E. P. Van Duzee, 3 males, 2 females
(CAS); Olema, V-25-.52, O. Bryant, 1 male (BYU);
MARIPOSA COUNTY: El Portal, V-18-3S, C. T. Sierr;i.

2 females (LA), J. R, Warren. 2 females (LA); Coidter-
ville, IV-17-55, J. R. Jessen. 1 male (CIS); Mariposa,
\'- 17-59, C, H. Toschi, 1 female (CIS); Yosemite, V-24-
38, J. R. \\';irren, 388()-4000', 1 female (LA); MODOC
COUN TY: Goose Liike, VII-24-22, C. L. Fo.v, 1 m:ile,
1 female (CAS); Lii.ssen Creek, VII-22-22. F. E. Blais-
dell, 1 male (CAS); MONO COUNTY: VI-4-17, F. E.
Blaisdell, 1 male (CAS); MONTEREY COUNTY: Ar-
royo Seco Camp, V-.5-56, 1 female (USU); Brysoii, IV,
V-I9, 20, 26-17, 20. E. P. Van Duzee, 9 males. 11 fe-
males (CAS); Cannel, IV-2-1I, E. C. Van Dvke, 4
males, I female (CAS), IV, V-2, 25, II. 8-29, 23, L. S.
Slevin. I male, 2 females (CAS): Monterey, IV-12-54,
R. P. Allen, 2 females (CIS), VI, A Feynes. I male
(CAS). 3 males (CU), 2 males, 1 female (CNC); Pacific-
Grove, VI, A. Feynes, I female (CAS), VII-16, 18-
1898, Lii/)ini(.v arbonis, I male, I female (USNM);
Pine Canyon, III-I9-20, L. S. Slevin, 1 male (CAS);
Tassajara, V-26-20, L. S. Slevin. LupinuSy 2 males, 4
females (CAS); Carmel, Tnlareitos Ranch, IV-27-.54, 1
m:ile (CIS); NAPA COUNTY: E. C. Van Dvke. 1
female (CAS); ORANGE COUNTY: E. C. Van' Dyke.
1 male (CAS); PLUMAS COUNTY: 3 mi. S. French-
man Res. 6(K)0', W. Gagne and C. \V. O'Brien, Lupimis
cauilatm, 12 males, 7 females (CWO); RIVERSIDE
COUNTY: ARiianga. V-12-29, 1 m;ile (CNC); Beiming.
IV-I3-I898, L. O. Howard, 23()0" (USNM); Ribbon
Wood, San Jacinto Mts., V-30-.39, E. G. Lindsay, 1 male
(CIS); S.\CRAMENTO COUNTY: Fair Oaks.'V-Il-I3,
Smith and Vosler, 7 males, 1 female (LA), 1 male
(CAS); SAN LUIS OBISPO COUNTY: Guvaina Valley,
V1-21-3I, E. P. Van Diizee, I fem;ile (CAS): Pozo, IV-
30-62, C. A. To.schi, I m;ile (CIS); Simmler, 111-20-40,
J. W. Tilden and G. S. Mansfield, 1 female (CAS);
SAN BERNARDINO COUNTY: Gxniillett, I male
(USNM); Cajon. V-14-,52, O. Bryant, 7 males, 8 fe-
males (BYU); Ont;irio. 1II-7-40, 'llopper and Graves,
Burr Closer, 16 males, 19 fem;iles (USNM); SAN
DIEGO COUNTY: 2 males, 1 female (CIS); III-12-I4,

E. P. Van Duzee, I female (CIS); Alvarado Co., IV-
24-.54, J. Powell, 1 male (CIS); Jacumba, X-26-26. Van
Dyke. 1 male (CAS); Mount P.ilomar. Vl-28-63. II. L.
Griffin. I male, 2 fem:iles (CIS); Pow;iv, F. E. Bhiisdell,
I m:ile, 2 fem;iles (CAS); San Diego', 2 males (CU),

F. E. Blaisdell, 1 fem;ile (CIS), E. C. \an Dvke. 1
in;ile (CAS). Wickham, I m;de, 1 female (MCZ)'; SAN
FRANCISCO COUNTY: Cocpnllett. 1 female (USNM);
V.in Dvke, 2 males, 1 female (CAS); San Francisco,
1889. F. !•:. Bkiisdell, 4 m;(les, 6 females (CAS), Van
Dyke. 1 male (CAS); SAN JO.\QUIN COUNTY: Ripon,
III-25-34, M. Crazier, I male (AMNH); Ripon, 111-23-

34. C. H. Scliw.ih. 1 feni.de (LA): Stockton, 111-19-34,
M. Crazier, I male (LA); Traiv, V-4-33, A. E. .Michel-
bnicher. 1 imie (CIS); SANTA CLARA COUNIT: Mt.
ILimilton. IV- 15-47, G. E. Boshart. CilUi. 8 males. 2 fe-
males (CIS); San Antonio Valley, IV-8-47, R. F.
.Smith, Lupinus, I male (CIS); SANTA CRUZ COUN-
TY: Ben Lomond, V-17-3I, E. C. Van Dyke, 1 male
(CAS), VI-I-30, L. Saylor, 1 female (USNM); Santa
Cruz Mts., VI- 11-22, 1 in;ile (CIS): VI-20-I2. Coleman.

1 male, 3 fem;iles (CIS); SIERRA COUNTY: Gold
Lake. VlI-16-21, 1 female (CAS); SISKIYOU COUNTY:
VII. 2 females, (USNM); SOLANO COUNTY: Rio
Vista, V-19-49, E. G. Lindsay, Lupinm; 7 m;des (CIS);
SONOMA COUNTY: Mark West Spgs,, V-10, 11-30, E.
P. V;in Duzee. 8 males, 3 females (CAS); lV-27-30, J.
<). \I:irtin, 2 males, 2 females (CAS); Mt. St. Helena,
l\'-17-32, C. E. Morh.nd, I fem;de (LA); V-6-30, E.
C. V;in Dyke. 3 males (CAS); Sobre Vista, IV-24-I0,
E. G. Van Dvke, 1 fem;de (CAS); TRINm' COUNTY:
Carrville, V-i28-34, 2400-2590', 1 male (FMNH); TU-
LARE COUNTY: Fail-view, 9 mi. So., V-1-64, [. Doyen,
4 males, 5 females (CIS), IV-29-64, P. Rude,' Lupimis
(■xciil>itits. 4 males, 1 fem;ile (CIS); Greenhoni Mts.,
V-7-31, E. C. Van Dyke, 3 males, 3 females (CAS);
White River, V- 17-30, E. C. Van Dyke, 5 males, 3
Icni.iles (CAS); TUOLUMNE COUNTY: North Fork
Tuohimne River, 3 mi. N.E. Tuolumne, V-1-61, R. M.
Brown. 2 males, 6 females (CAS); Stravvberrj-. VIlI-4-
60, G. W. Colliver, 1 male (CIS); YOLO COUNTY:
Rumsev, V-3-36, B. E. White, 2 males, 2 females (CAS);
SANTA ROSA and SANTA CRUZ ISLANDS: Santa
Cruz Is., IV-8-41. G. P. Kanakoff, Lupimis bicolor, 4
males, 7 fein;iles ( L.\ ) .

Colorado: 1 male, (USNM); Palm, 1 female
(AMNH); Boulder. VI-10-61. B. H. Poole, 5500', 1
male (CNC); Denver. VII-7, Hubbard and Schwary,. 1
miile (USNM); Denver, VII-7, F. C. Bowditch, I male
(MCZ); Denver, Adams Sp., VI-15-49, B. L. and J. G.
Rozen, I femide (CIS); Glemvood Springs, VII, VIII. A.
l'V\nes, 2 males. 2 fem;des (CAS); Pueblo, V-20, H.
Soit;iu. 1 female (USNM); Ste;imboat Springs. VII-42.
VIII-45, 21 males, 28 females (BYU); Valmont Butte,
Boulder, VI-2(>, \II-fil, |. R. St.iiiier. 5300'. 1 fenwle
(CNC).

Idaho: Caribou Co., Soda Springs, 1 mi. N.,
VII-9, 10-68, D. R. Harris. Lupimis, 2 males (WEC);
Coeur D'Alene. VI, Wickham, 1 female (USNM); Win-
chester, V-11-24, M. C. Lane, 1 female (USNM).

Montana: Boulder, Jefferson Co., VII-31-68. W. E.
Clark, Lupiuus scriccus, 5 m;iles, 2 females (WEC):
Bozeman. \'ll-25-03. 4800", 1 male (USNM); Bridger
Canyon, \'II- 12-02, 5(X)0'. 1 female (USNM); Big Horn
Co.,' Bushby, 4 mi. W., VI-8-69, W, E. Clark, Lupimis,

2 m;ili-s (WEC); Florence, V-24-13, H. P. Wood. 2

m;iles (USNM), VI-I, 17-12, 1 male

females.

(USNM); Custer Co., Miles City, 17 mi. N.E., VI-8-69,
W. E. Chirk, I male (WEC); Nlissoula, VII-6-68. 1 fe-
male (USNM), Silver Bow Co., Nissler, 5 mi. N., VIII-
6-68, W. E. Cl.iik. Lupiuus scriccus, I male (WEC);
R;ivalli Co., Roaring Lion Canyon. Vl-23-35, W. L. Jelli-
soii, Lupiuus, 9 males, 7 females (USNM); Big Ilorn
Co.. Wvohi, II mi. S., VI-8-69, W. E. Clark, Lupiuus,
1 inale.'2 females (WEC).

Nevada: Horn, 1 male (PANS); G;irson City, \l-2.5,
26-29, R. R. Usinger, 9 m:iles, 10 females (GAS); Wick-
ham, 1 fem;ile (USNM); Ormsby Co., VII. Biker, 2
males (FMNH). I male (USNM)'.

Oregon: Athena. VI- 12-38, K. Gray and J. Scluih,
1 male (USNM): Corvallis, VI-7-32, E. C. Van Dvke,
I female (CAS), V-22-35, K. Gray, 1 female (CAS);

Biological Seiues, Vol. 13. No. .3 Revision of the Weevil Genus Tvchiu.s

19

l':ii;in, \^I-20-22, A. L. Lovott, 1 tcmale (CAS); Hood
Hivcr, Vl-4-17, F. R. Cole, 1 male (USNM); Kamela,
VMO-25, M. C. Lane, 1 male (USNM); No Powder,
VI-8-24, 1 male (USNM); Steen Mts., 4 mi. W. Fish
Lake, VII- 15-53, Roth and Beer, 1 male, 1 female
(OSU); Woods, Vl-13-39, K. M. and L. M. Fender, 1
female (FMNH).

Saskatchewan: Farewell Creek, I male, 2 females
(BYU).

Utah: Avon, V-29-39, G. F. Knowlton, 1 female
(USNM); Beaver Co., VI-14-57, G. F. Knowlton, 2
males, 2 females ( OSC ) ; Bellexue, Sehaeffer. 1 female
(BYU); Blue Si)rin<;s Hills, Box Elder Co., VI-28, V. M.
Tanner, 1 male (bVU); Cache Jet., VI-11-03, 1 female
(BYU); Cove Fort, V-29-37, G. F. Knowlton, 1 male
(USNM); Dixie N,it'l. Forest, VI-1.5, .3.5, G. F. Knowl-
ton, Li(;)/iii/.s, 2 females (USU); Eden, VII-23-37,

1 fem;ile (USU), Enterprise, 8 mi, S., VI-15-35, G.

F. Knowlton, Lufunns. 2 females (I'SU); Utah Co.,
Hobble Creek Canyon, Sprini;ville, 5 mi. E., V, VI, VII,
VIII-3, 24, 17, li, 16-68, 69, W. E. Clark, Liipinus
leiicophylhis, .33 males, 30 females (WEC), 19 mi. E.,
VI-6-(>8, Lupinus scricciis, 14 males, 22 females (WEC);
Huntsville, V. M. Tanner, 1 male (BYU); Leeds, IV-2.5-
35, G. F. Knowlton and C. F. Smith, Lupinus, 2 males,

2 females (USNM); Logan, VI-10-50, John V. Bmce, 1
male (USU); Mantau,' VI-20-61, G. F. Knowlton, 1
male (USU); Mt. Meadows, VI-15-35, G. F. Knowlton,
1 male (USU); Iron Co., Orton. 12 mi. N.W., VII-17-67,
H. R. Burke, 1 female (TAM); Salt L;ike Citv', Big
Cottonwood Canvon, V-22-33, G. F. Knowlton, Lupinus.
1 male, 4 females (USU), 2 fem;iles (USNM), VI-6-.3.5.
1 m;ile (USU); St, George, V-28-35, E. C. Van Dyke, 1
m;ile (CAS); Duche.sne Co., Mtn. Home, 7 mi. N., VII-
13-68, W .E. Clark, Lupinus sericcus. 11 males, 11 fe-
males (WEC); Trout Creek, \'-8-.34. T. O. Thatcher.
Lupinus, I male (USU); Wasatch, VI-27, Hubbard and
Schwarz, 8 males, 8 females (USNM), VI-27, Horn, 2
m;iles (PANS).

Washington: Blewett, V-29-32, |. Wilcox, 1 female
(OSU); Brewster, IV-29-12, 1 female (USNM): Drvden,
V-16-42, E. C. Johnston, 1 female; Kooskooski, V-1-46,

G. Nelson, 1 female (TAM); Walla Walla, VI-9-38. E.
C. \'an D\ke. I male, 1 female (CAS); Sanpoil, Keller,
VII-3-21, 'M. C. Lane, 1 female (USNM); Waw;.wai, 1
male (USNM).

Wyoming: Johnson Co., Buffalo. 8 mi. S.W , \'I-20-
68, W. E, Clark. Lupinus anunophilus, 8 males, .5 fe-
mides (WEC), 5 mi. W. Lupinus cirgcntcus, 1 m;ile,
I female (WEC). VI-7-69, 3 males, I female (WEC);
Campbell Co., Gillette, 22 mi. W., W. E. Clark, Lu-
pinus argenteus, 8 males, 2 females (WEC); Niobrara
Co.. Lusk, 11 mi. S., VI-15-68, W. E. Clark, Lupinus
argenteus, ,3 male, 1 female (WEC); Teton Co., 12 mi.
S. Jackson, VI-23-62, 6000", 1 male, 1 female, C. W.
O'Brien (CWO).

Total specimens ex.imincd: 1011.

Disciis.sion. X'ariation is e\ident in the .size,
shape, and color of the scales. .Specimens from
California exhibit a wide range of variation and
some distinct varieties can be associated with
particular geographic areas. Some specimens
from the Los Angeles area have gray and lironze-
colored scales which are unusiuilK- long and
acuminate. The lectotvpe localitv is probiibly Los
Angeles, as the lectotvpe has this tvpe of vesti-

turc. Specimens from tlie west slope of the
Sierra Nevada Mountains are relatively small
and have gray and metallic bronze areas on the
prothorax and on alternate elytral interspaces.
Specimens from the east slope have no bronze-
colored scales. The scales on these are also denser
and broader. Two populations were sampled
from Hobble Creek Canyon in Utah County,
Utah. Specimens from the mouth of the canyon
taken on Ltiphuis IcucojiliiiUiii exhibit contrast
Ix'tween grav and bronze scales, but specimens
taken a few miles up the canyon from L. sericeus
are nearly unicolorous, as are specimens from
the same host at Mountain Home, Duchesne
Coimtv, Utah.

Tt/cliius liljchladi Blatchley

(Figs. 13, 19)

Tt/chius litjcbladi Blatchley, 1916, In: Bhitchley and
Leng, Rhynchophora or weevils of northeastern
America, p. 246-247 (Holotype: male, Steuben Co.,
Indiana; Purdue).

Ti/chius arator: LeConte, 1876, Proc. Amer. Philos. Soc,
15:216; Casev, 1892. Ann. New York Acad. Sci.
6:415; Blatchley and Leng, 1916, Rhynchophora
or weevils of northeastem America, p. 247; Marco-
vitch, 1916, Rep. State Entomol. Minn., 16:140,

A/iC(;-ofrog!(.s liljehladi: Klima, 1934, Colcoptcrorum Cuta-
logus, 29(1.38): 30.

This species resembles T. tecttis LeConte, in
general facies. It can be distinguished from other
North American species bv the shape of the ros-
trum wliicli is prominently swollen jjasally, and
smooth, sliining, and finely acuminate beyond
the antennal insertion; by the light yellowish
brown scales; and by the sparse, scattered,
round, white scales on the elytra.

Description. Male: Length 2.8-'3.4 mm, width
1.5-1.7 mm; integument black on pronotum, usu-
ally dark reddish brown on elytral apices, ap-
pendages light to dark reddish brown. Vestiture
of light vellowish l)ro\\ai scales on dorsum,
scales on ventral surface white.

Rostrum shorter than prothorax; from lateral
;ispect swollen basally, prominently arcuate
from dors;d margin of eyes to basal fourth, then
slightly to moderately arcuate to apex; antennal
insertion in apical third; moderately to strongly
acuminate, smooth, shining, glabrous, with shal-
low punctures distad of antennal insertion. From
dorsal aspect not strongly tapered from base to
apex, slightly expanded at antennal insertion,
scales of uniform color, size and shape, long, nar-
row or wedge shaped; no erect or suberect setae.

.\ntennal funicle seven-segmented, pedicel
shorter than next three segments combined.

20

Brigham Young University Science Bulletin

Pronotum 1.2 times wider thiin long; sides
prominently arcuate, slightly constricted at an-
terior margin, nearly twice as wide at base as at
anterior constriction. Dorsum covered mainly by
broad, usually apically rounded scales. Sides and
often small median basal portion ot dorsum with
round to elongate, white scales.

Elytra moderately convex in dorsal profile;
in dorsal aspect widest just beyond middle,
prominentlv rounded in apical third. Scales on
interspaces of uniform size, shape and color;
long, narrow, often spatulate; sparse, scattered,
white scales mainly on apical third, denser on
sides. Strial scales narrower than scales on in-
terspaces.

Ventral surface with pale yellow to white,
round to elongate-oval recumbent scales; erect
or suberect setae absent. Sternum live with deep
median fovea.

Femur with prominent, apical, ventral emar-
gination, usuallv with luinnte tooth on proximal
portion of emargination. \'estiture of uiiifonn
size and shape, similar to that of elytra but
lighter in color.

Tibiii mucronate, mucro on protibia usually
larger than mucroiies on mesotibia and meta-
tibia; vestiture of long, narrow, scales and fine
setae, setae predominant apically and ventrally.

Tarsi clothed with fine, eloncate, white to
yellow scales; claws short, stout; tooth connate
in basal fourth, not as long as claw.

Male genitalia ( Fig. 1.3) with apical portion
of median lolie brcKidlv rounded; ;ipical, dorsal,
medi;ui membranous area snuill, oval, strongly
defined posteriorly; median lobe constricted
medi;illv; median struts clavate.

Feiiiiile: similar to male but with rostrum
longer and more finely acuminate beyond an-
tennal insertion.

Host. Astragaltts cunadcnsis.

Distribution. (Fig. 19).

Alberta: Cypress Hills, VI-30, F. S. Carr, 1 male. 1
female (UA), 1 male (CNC); Medicine Hat, VI-6, 28-
26, 28, F. S. Carr. 3 males, 1 female (CAS), 1 female
(UK); 1 male (UA), VI-28-26, 2 males (USNM). 1
male, I female (CNC), 1 male, 3 females (BYU).

Colorado; Denver, \'II-7. Huhhard and Seliwar/.. 1
female (USNM).

Illinois: H. Soltau, 1 female (USNM).

Iowa: Ames, VII-26-51. ]. Laffoon, Astragalus cdna-
demis. 2 males. 2 females (ISU), VI-3-32, J. A. Adams.
1 male (ISU), IV. VIII-30, 25-1897, 2 males, 1 female
(ISU); VII-7-34, H. E. Jacques, 1 male, 2 females
(ISU); Iowa Co., VII-30-35, H. E. Jacques, I female
(ISU), VI 1-5-3.5, G. Warren, 1 female (USNM);
Granite, \II-28-16, D. Stoner, 1 m.ale (USNM); Lake
Okoboji. VII, VIII-6, 13, 22-16, 17, L. L. Bueh.inan, D.
Stoner, II males, 11 females (USNM); Ledvard. 2 mi.

S., V-9-26, G. O. Hendriekson, 1 female (ISU); Lemars,
6 mi. N.W., VII-26-28, G. O. Hendriekson, 1 male, 1 fe-
male, (USNM), 2 females (ISU).

Kansas: 1 male (USNM); Douglas Co.. F. H. Snow.
90()'. 1 female (UK); Topeka, VI, VlII-12, Popenoe, 3
males, 4 females (USNM); Leavenworth Co.. 6 mi. W.
Linwood, VI-17-64, j. B. Karen, 1 female (CVVO); Wal-
kiee Co.. F. H. Snow, ,500'. 1 male (UK).

Manitoba: A«eme. IV, VllI-19, 12-30, R. M. White,
Astragalus caruidensis. 1 male, 2 females ( CNC ) .

Michigan: Grand Ledge, Vll-16, Hubbard and
Scliwar/, 2 males (USNM); Monroe, Hubbard and
Schwar/, 1 male. 1 female (USNM).

Minnesota: Cliisago Co., Chisago Lake, VlI-19-21,
F. P. Mctelaf, in seed of Astragalus canadensis, 1 female
(USNM); St. Anthony Park, Astragalus canadensis, 2
males (USNM).

Missouri: C. Schaeffer, 2 males, 1 female (BYU).

Nebraska: Seward, 1 male, 1 female (BYU).

North Dakota: Case Co., VII-15-63, R. Gordon, 1
male (BYU); Fargo, VlI-22-22, R. L. Webster, 1
female (BYU); Wahpeton, 1933, Wickham. 1 male, 1
female (USNM).

South Dakota: 2 males (CU); Volga, 1 male, 2
females (LA); Truman (Wickham Coll.), 1 female
(USNM); Roberts Co., 21 mi. S. Sisseton, Vll-1-64.
L. and C. W. O'Brien, 3 males (CWO).

Texas; Dallas, F. C. Bowditch, 1 female (MCZ).

Washington; Metaline Falls. Vll-20-32, T. Terrell, 1
female (USNM).

Total specimens examined: 101.

Discussion. The tvpe specimen of this spe-
cies was examined and determined to be con-
specific with specimens identified as T. orator
Gyllenhal in collections in North America. Speci-
mens labeled T. orator Gyllenhal have been ex-
amined from the LeConte, Casey ;md Rhitchley
collections ;md found to be T. UJjchladi. The
basal swelling and acuminate apical portion of
the rostnim are not well dc\'eloped in the holo-
type. TijcJiitis arator Gyllenhal is a synonym of
T. aratus Say.

Tychius tectus LeConte

(Figs. 1, 12. 19)

Tychius tectus LeConte, 1876, Proc. Amer. Philos. Soc,
15:217 (Holotype: female. Kansas, MCZ type
.5233); LeConte, 1879, Bull. U.S. Geol. and Geog.
Survev., 5:506; Casey, 1892, Ann. New York Acad.
Sci., '6:414-415; Ca.sey, 1910. Can. Entomol.,
42:135.

Tychius languidus Casey, 1910, Can. Entomol., 42:135
( Holotype; male. Garland, Colo., USNM 36753, T.
L. Casey collection).

Miccotrogus tectus: Klima, 1934, Colcopterorurn Cata-
Ingm, 29(138):32.

This species differs from other members of
the T. sordulus species group bv the vollow or
reddish brown rather th;m grav vestiture. The
body is oblong ( Fig. 1 ) ; the sides of the elytra
nearly p;irallel, elytra with ba.sal portion flat

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius

21

rather than rounded in dorsal profile. It can be
di.stingui.shed from T. liljcbhidi, which it resem-
l)les in general faeies by the .shape of the ros-
trum, which is not tumidus at the base nor
acuminate in the apical portion, and by the
deeply riigulose distal portion.

Description. Male: length 2.5-3.8 mm; integu-
ment piceous to black, appendages light to dark
reddish browii. Vestiture of yellowish or reddish
brown scales, usually with median and lateral
vittae of white scales.

Rostrum shorter than prothorax; in lateral
aspect usually moderately to prominently and
evenly arcuate, but often nearly straight pro.xi-
mad of antennal insertion; antennal insertion in
apical third; in dorsal aspect moderately, eyenly
tapered from base to apex; apex narrower than
frons between dorsal margin of eyes, distal por-
tion oval in cross section, deeply rugulose. Vesti-
ture of elongate-oval or parallel sided scales with
rounded or truncate apices, scales on sides usu-
ally of lighter color; glabrous distad of antennal
insertion except for sparse setae around apical
portion of scrobe.

Antennal funicle seven-segmented, pedicel
usually as long or longer than next three seg-
ments combined; scales on antennae elongate,
clavate.

Pronotum wider than long, usually widest
in front of middle, rounded slightly to base and
strongly to apical constriction (Fig. 1); 1.4-1.7
times wider at base than at apical constriction.
\'estiture of elongate-oval or parallel sided, api-
callv rounded or truncate, light to dark yellow-
ish or occasionally reddish bro\\ii, strigose scales,
usually with median vittae and lateral, patches of
white scales. Lower portion of sides with elon-
gate-oval, nonstrigose, usually white or light
colored scales, some of which may extend to
dorsum, especially basally.

Elytra in dorsal aspect with sides converging
slightly or parallel in basal two-thirds; promi-
nently tapered to apices, usually widest just
distad of humeri; nearly flat or very slightly
rounded in basal half in dorsal profile, declivity
l^roadly rounded. Interspaces with scales similar
to those on prothorax. Interspace one with dense,
oval, white, nonstrigose scales from base to
apex, and with several long, narrow, darker
colored scales intenningled throughout; usually
with broad lateral vittae of white, oval scales.
Scales of strial punctures elongate, narrower
than scales on interspaces.

\'entral surface densely covered by white
or nearly white, broadly imbricated, oval to
elongate-oval scales; no distinct rows of erect or

suberect setae. Sternum five with median fovea,
usually concealed by scales.

Femur with prominent, apical, ventral emar-
gination; no minute tooth on proximal portion of
emargination. Vestiture of dense, broad, elon-
gate, usually p;nallel sided, truncate, or apically
rounded scales, with elongate-oval, nonstrigose
scales on basal portion.

Tibiae mucronate, mucro on protibia largest,
about ecjual in length to tarsal claw. Vestitme
of elongate, broad, strigose scales with very fine
hairlike setae near apex.

Tarsi clothed dorsally with hairlike setae and
broad, strigose scales; claws with basal processes
parallel, nearly half as long as claw.

Male genitalia (Fig. 12) with apical portion
of median lobe rounded; apical, dorsal, median
membranous area round, extending proximad be-
yond middle of median lobe, strongly defined
posteriorly; median struts and tegminal strut
strongly clavate.

Female: rostrum slightly longer and narrow-
er, antennal insertion near middle; mucrones
slightly smaller.

Hosts. Astragalus adsiirgens var. robustior,
A. histdcatits var. Iict/denianus, A. scopulorum, A.
tenellus, Oxijtwpis besseyi, O. campestris var.
gracilis, O. lambertii and O. sericea, Hedijsarum
sp. Also recorded from "vetch."

Distribution. (Fig. 19).

Alaska: Big Delta, VlI-16-48. R. T. Sailer, 3
males, 1 female (USNM).

Alberta: Cardston, 9 mi. S., VIII-6-68, W. E. Clark,
Oxt/tropi.s campcstris var. gracilis, 4 males, 4 females
(WEC); Edmonton, VII-14-20, F. S. Carr, 1 female
(AMNH), VII-14-20, 1 male, 3 females (CAS), 2 males,
2 females (CU), 2 males, 1 female (MCZ), 3 males
(PANS), 1 female (PA). 5 males, .5 females (UA), 3
males, 2 females (UK), 11 males, .5 females (USNM),
4 males, 1 female (OSU), VII-14-20, J. G. Shenafelt, 1
female (LA); Medicine Hat, VII-14-20, A. C. Davis
coll., 1 male, 1 female (CNC).

British Columbia: Naramata, V-28-58, H. and A.
Howden, "on vetch," 1 male. (CNC); Oliver, 2 mi.
W., V-29-.58, H. and A. Howden, "on vetch," 1 male,

1 female (CNC); Penticton, 3 mi. E., VI-1-58, H. and
.\. Howden, "on vetch," 2 males (CNC); Ritcher Pass
Road, 7 mi. W. O.soyoos, VI-2-.58, H. and A. Howden,

2 males, 1 female (CNC).

Colorado: Buena Vista. VI, VII-15, 30, 1, 6-96, H.
F. Wickham, 7900-8000', 5 males, 2 females (MCZ),
VII-1, 6- 1896, 5 males, 2 females (USNM), VIII-5,
Liebeck Coll., 3 males, 1 female (MCZ), III-7, Hubbard
and Schwarz Coll.. 4 males, 3 females (USNM), H. F.
Wickham. 4 males, 1 female (USNM), 6 males, 3 fe-
males, (AMNH), 4 males (CU), 1 male (CAS), 1 male,
1 female (UK), Boulder, VI-9-61, W. R. M. Mason,
.5.500". 1 male (CNC); Colorado Springs, II-IO; 2-4, H.
Soltau Coll.. 3 males. 10 females (USNM): Garland, 24
males, 14 females (USNM), 2 males (UK), VI-30, F.
C. Bowditch, 1 male (MCZ), VI-29, 30, Horn Coll., 2

BmciiANt Young Univhrsity Science Bulletin

fomalfs (PANS), Vl-29, 30, lUibbard and Schwarz. 6
males. 5 females (USNM); Mancos, La Plata Co., 7 mi.
E., V-30-69, \V. E. Clark, Astrw^aliis hisiilcdtus \ar.
Iicijdenianus, 3 males, 2 females (WEC); Montrose,
VIII-8-1885, F. C. Bowclittli Coll., 6000', 2 males, 1
female (MCZ); Arclinleta County, Pagosa Springs, 26
mi, S.E., V-31-69, W. E. Clark,' Astr(if2,alus hisiilcatiis
var. hcijileniaiuis, 15 males, 8 females (WEC); Poucire
Canvon. Larimer Co., Vl-12-68, W. E. Clark, Oxxjtropis
tiimhcrtii and O. scricea, 57 males, 33 females (WEC);
Hidgeway, Ouray Co., 2 mi. S.W., V-31-69, W. E. Clark,
.A.s7r(/grt/n.v liisulcatux \ar. Iwijdciiiantm, 5 males, 5 fe-
males (WEC); Toponas, Routt Co., 3 mi. E, VI-4-69,
W. E. Cliirk. Aslragiiliis tcnclhts, 9 males, .5 females
(WEC). 19 mi. E., VI-5-69, W. E. Clark, Aslmnahn-
scorpiilortim, 7 males, 3 females (WEC).

Manitol)a; Aweme, \'l 1-4-03. \. Criddle, i male
(CNC), VI-4-29, R. M. White, Oxi/lwpis lamhcrlii, 2
males, (CNC); Treebank, 5-18-27, N. Criddle, Astra-
liiilus, 1 miile (CNC), VI-14-27, R. M. White, 2 males,
2 fem;iles (CNC).

Montana: Powder Hi\cr Co.. Ashl.ind, 7 mi. E.,
VI-8-69, W. E. Clark. O.vi/fro/iK, 3 males, 1 f<m;dc
(WEC); Big Horn Co., Biisliby, 4 mi, W., VI-8-G9, W.I-;.
Clark, Oxi/tropis scrUca. 2.5 males, 10 females (WEC);
Kalispell. Vl- 13-20, Wickham. 1 male (U.SNM); Custer
Co., Miles City. 17 mi. N.E.. VI-8-69, W. E. Clark, Oxij-
tropis lumbcrtii, .3 males, 2 females (WEC); Missoula,
Liebeck Coll,, 5 males, 3 females (USNM); Silver Bow-
Co,, Nissler, 5 mi, N„ VI 11-6-68, W. E, Clark, Astni'iii-
lit.i «(/,si/rgfM,s- ssp, rohu.stior, 1 male. 2 females ( WEC ) ;
Ghieier Co,, Piegan, 1 mi, S„ VIIl-6-C>8, W. E, Clark,
Oxi/lropis campestris var. g,racilis, 1 m;de, 2 females
(WEC), 4 mi, S,. 1 femide (WEC).

Nebraska: MeCook. Hul)l);ird ;uid Sehwiirz, 1 fe-
male (USNM), F, C, Bowditeh Coll,, 1 male (MCZ),
Wiekham Coll., 2 males, (USNM), R, Hopping Coll,,
1 male, I female (C.-VS); War Bonnet C;iri\on, 1 m;de
(USNM).

Nevada: Klko Co., east slope Spruce Mlu., \ 1-26-
.56, W, C, Russell, 1 male (CIS),

New Me.xieo: Rio Arril);i Co.. Cliani.i, 17 mi. \.W..
\'-31-69, W. E, Clark, Astni^iiliis hisiilctitux var, licij-
(Icttiumis, 12 males, 8 fem;iles (WEC;); |eme/ Mts,, VI,
VllI, L\-21. 8-21. 27. J, Woodgate. 4 males, 5 females
(CAS).

North Dakota: Colden Valley Co,, Beiich. 12 mi. E.,
\"I-9-69, W. E. Clark, Oxi/lropis'lanihcrlii, 1 m;de, 2 fe-
males (WEC); Dunn Co', Killdeer 1 mi. S., \I-10-69,
W, E, Clark, Oxtjtwpk larnhcrtii, 1 male (WEC); Mc-
Kenzie Co., Newtown, 17 mi, W,, VI-10-69, W, E.
Clark, Astriif^ahis tcneUus, 10 m;iles, 10 females
(WEC); Mountrail Co,, Parshall, 3 mi. N,W„ \'I-10-69,
W, E, Chirk, Oxijtropis, 10 m;iles, 10 fem;iles (WEC);
Theodore Roosevelt National Park, South Unit, Vl-9,
10-69, W. E. Clark, Oxiftropis sericm, 19 males, 4 fe-
males (WEC); Williams Co., Williston, 33 mi, N.,
VI-ll-69, W, E, Cl;irk, Oxijiropis. 2 m;iles (WEC), 31
mi, N., VI- 1 1-69, W, E, Clark, Oxi/tropis, 1 male, 1 fe-
m;ile (WEC),

Oregon; Kamela, VI- 10-25, M, C, Lane, 2 males, 3
females (USNM).

Saskatchewan: Fish Creek, VII-I8-25, K. M. King.
4 males (CNC); Forta a L:iConne, VII-17-2.5, K, M.
King, 1 fem;de (CNC); Moose Jaw, 16 mi. E, VI-12-
69, W, E. Clark. 1 m;ile, 2 females (WEC).

South Dakota: Lawrence Co., Brownsville. 1 mi. S.,
VI- 18-68, W. K. Clark, O. ciimpc.stris \ar. grmiVis, 14
m;iles. 17 fem;iles (WEC); Lawrence Co., Cheyenne

Crossing, 2 mi, E,, \'l-18-68. W. E, CI. irk, O. camjic.stris
var. gracilis, 5 males, 3 females (WEC); Todd Co.,
Mission, 15 mi. S., VI-11-.50, llieks, Shiter, L;iffoon, 1
male, 2 females (ISU); Pennington Co,, Pactola Reser-
voir, Vl-17-68, W, E, Clark, 1 femiile (WEC),

Wyoming: Horn Coll,, 1 male (PANS); Albany
Co,, Albany, 5 mi, N,E„ VI-.5-69, W, E, Clark, Oxijtropis
scriccd, 3 males (WEC); Johnson Co.. Buffalo, 5 mi,
W, VI-7, 14-69, W, E, Chirk, Astragalu.s udsurgcm ssp,
robuitior, 15 males, 4 fcmides (WEC), 8 mi, S.W„ VI-
20-69, W. E, Chirk, Oxijtropis sericca, 19 m;iles, 10
females (WEC); Campbell Co,, Gillette, 22 mi. W„ VI-
20-68, W, E, Chirk, Oxijtrupi.s himl)ertii, 1 male, 1 fe-
male (WEC); Johnson Co,, Kaycee, 1 mi, N, VI-7-69,
W, E, Clark, Oxijtropis bessei/i, 5 males, 2 females
(WEC); FremimtCo,, Lander, 14 mi, S., VI-I4-69. W.
E, Clark, 1 female (WEC); Albany Co,. Laramie, 4
mi. N,W,, VI-6-69, W. E, Clark. Oxijtropis hmbertii, 5
m;iles, 3 females (WEC); Niobrara Co., Lusk, 11 mi. S.,
VI-15-68, W. E. Clark, Oxi/tropis bcssci/i, 4 males. 1 fe-
male (WEC); Carbon Co., Medicine " Bow, 3 mi. N.,
\T-6-69, W. E. Clark, Oxi/tropis scricea, 5 males. 4
t'em:iles (WEC); Shoshoni." Fremont Co,, 11 mi, N„
\'l-21-68, W, E. Clark, Oxijtropis lagopus, 14 males. 11
females (WEC); Hot Springs Co,, Thermopillis, 10 mi,
N„ Vl-21-68, W, E, Clark, Oxi/tropis hiiio/ms, 7 males,
5 fem;iles (WEC); Washakie Co., \\orland, 7 mi. E.,
VI-14-69, W. E. Clark, Oxi/tro/iis hgo/ms, 6 males. 6 fe-
ni;iles (WEC),

Yukon Territory: Ross River, 132°3', 61°56',
3,000 ft,, VI-20-60, Hcdijstirum, J, E. H, M;irtin, 22
in;iles, 23 females (CNC), VI-19-0(). E. W, Rock-
l)urue, 27 males, 26 females (CNC),

Total specimens examined: 816.

Discussion, Geographic \;iri;iti()n i.s ovident
in scale shape, size, and color, as well as in the
average size of specimens. Individual variation
is evident in size, distribution of white scales,
and color of the dense, elongate scales which
iinixirt the general color to the specimens. In ;i
given series, specimens usuallv agree closch' in
the coloration and distribution of scales, but
often range from verv light to ;i few very dark
colored specimens, Usualh ;i number of gra\' or
silvery gra\' specimens can be observed.

Some specimens from southwestern Colorado
;uul northern New Mexico have verv dark \el-
hnvish l)rown scales. These were associated with
.A.v/r«g«/(/.v l)isitlc(itus. Specimens labeled "Jemez
Mts., " New Mexico, have a verv light red integu-
ment. The scides on these are light to verv dark
reddish brown pro\iding ,i ]U;irki'd contr;ist with
the white se;iles.

Specimens from .\l;isk;i ami the Yukon Terri-
tory of ("iuiada have a lighter, yellowish to gray-
ish \'estiture. Individn;il scales are n;uTOw, leav-
ing the integiiiiicnl bn);Kllv exposed.

A single leiiuile specimen Irom iiortiuMsteni
Nevadii w;is ex;uiiiiied which is uiii(|ue in se\iTal
cliaracters. It is small, 2,.") nun in length, with
scales ver\' broad, elong;ile-()\;il, ;uid relatively
sparse, and integument bro;idlv visible. The pedi-

Biological Series, Vol. 13, No. 3 Revision or the Ween-il Genus Tychius

23

Fig. 1. Dorsal \ie\v of Ti/clihis tcctiis.

eel of the antenna is longer tlian tlic next fonr
.segments eombined.

Five specimens examined from Kamela, Ore-
gon, average 3.9 mm in length and have very
elongate, narrow rostra wiiich in both sexes are
silghtly longer than the prothorax. In these the
rostra are finely tapered from the base to the
apex and amnminate distad of the antennal in-
sertion, but the di,stal portions are deeplv rugu-
lose.

The specimens from Nevada, Oregon, Alaska.
and the Yukon Territorv ha\e extralimital distri-
butions (Fig. 19).

T>j(hi\is .scmis(jU(nnosus LeConte

( Figs. 9. 21 )

Tijchiiis semisquamosus LeConte (not Faiise. 1893).
1876, Proe. .Amer. Philos. Soc., 1.5:217-218 ( Leco-
type here designated: female. Fort Tejon, California,
MCZ type .5229; Paralectotvpe, female, .same lo-
cality, .MCZ, type .52292); Casey, 1892, Ann. New
York Acad. .Sci'.. 6:418.

Miccotrugus .v(';(ii.5r/i/«mo.«/.s: Klima. 1934. Coleoptcronmi
Catalogus, 29(1.38): 32.

This species may be distinguished from other
Nortli American representatives of the genus

by the multiple, confused, as opposed to single,
unifomi, median rows of long, narrow, light to
dark reddish brown scales on the elytral inter-
spaces; by the absence of fine, erect setae on the
abdomen; and bv the asvmmetrical apical por-
tion of the median lobe of the male genitalia
(Fig. 9). It is doubtfully distinct from 7'. himeU-
osus Casey but can be distinguished by the fol-
lowing characters: the elongate-oval, white scales
on the elvtral interspaces are very sparse and
rarely imbricated, and the rostrum is not finely
acuminate and the average size is smaller.

Descri]jtion. Male: length 2.5 mm, width
1.1 mm; integument black to piceous, append-
ages light to dark reddish brown; vestiture of
white to dark reddish brown scales.

Rostrum shorter than prothorax, antennal in-
sertion on distal fourth; moderately evenly arcu-
ate in dorsal profile; in dorsal aspect strongly,
evenly tapered from base to apex. Irons 2.9 times
wider between dorsal margin of eyes than ros-
trum at apex; distal portion strongly tapered,
smooth, shining, pits and rugae shallow. Vesti-
ture of elongate, broad, strigose, recumbent,
apically rounded scales of unifomi shape; distal
portion glabrous except for sparse, fine setae
around apical portion of serobe.

Antennal hinicle seven-segmented; pedicel
as long or longer than next three segments com-
bined; setae broad, elongate, apically rounded.

Pronotum wider than long, about 1.5 times
wider at base than at apical constriction; sides
evenly, prominently arcuate. Vestiture complex,
consisting of long, narrow, apically rounded or
pointed, recumbent, dark reddish browni, strigose
scales covering dorsum and extending about half
wav down sides; integument broadly visible on
dorsum, scales on lower portion of sides oval to
elongate-oval, nonstrigose, light reddish brown,
extending dorsallv intermingled with long, nar-
row scales forming broad lateral vittae on dor-
sum; sparse oval scales scattered throughout on
dorsum, also forming small, median, dorsal, basal
patch.

Elvira nearlv parallel-sided in basal two
thirds, broadly rounded to apices; in dorsal pro-
file nearlv flat in basal half, declivity broadly,
evenly rounded. Vestiture on interspaces of
sparse, scattered, round to elongate-oval, some-
times slitrhtlv imbricated, recumbent, nonstrigose
white to very light reddish brown scales, much
denser, darker, and more broadly imbricated on
interspace one. Each interspace with confused,
multiseriate rows of long, narrow, apically
rounded or pointed, dark reddish brown, usually
suberect, strigose scales; scales not denser on
intersals two through four. Strial setae narrow,

24

Bmcii AM VouNc: Uni\ kissitv Scmence Bulletin

about lialf as wido as long, narrow scales on in-
terspaces, usually lighter in color.

Wntral surface with round to elongate-oval,
recumbent, slightly imbricated, nonstrigose,
white to light reddish brown scales; some scales
slightlv narrower and suberect, especially on
sterna four and five; no discrete transverse rows
of elongate, fine, hairlike setae. Stcniiini live
with deep median fovea.

Femur elongate, apical half slightly swollen,
apical ventral eniarginations prominently de-
veloped, metafemur lacking minute l(K)th on
pro.ximal portion of apical ventral emargination.
Vestiture of elongate-oval, recumbent, white to
light reddish brown, nonstrigose scales, and
long, narrow, strigose, white or light reddish
browTi, apically trimcate or rounded, strigose
scales; no fine, erect, liairlike setae.

Tibiae mucronate, mucrones shorter than
tarsal claws, protibia with larger mucro; vesti-
ture of long, narrow, strigose scales, and fine,
erect, light brown setae on apical portion, rare-
Iv with sparse elongate-oval scales.

Tarsi dorsallv with long, narrow scales and
fine hairiike setae; basal process of claw about
two thirds as long as claw.

Male genitalia (Fig. 9) with apical portion
of median lobe strongly asvmmt'trical; apical,
dorsal, median membranous area elongate, weak-
ly defined posteriorly; median struts c!a\ate.

Female: length 2..3-2.7 mm, rostiinn narrower,
slightlv more acuminate in distal half, antenna]
insertion near middle. Tiliial mucrones slightly
smaller.

Host. Unknown.

Distribution. (Fig. 21).

California: T. I,. Cascv, coll., 1 male (USNM);
.\rgus Mts., V-1891, 1 female. (USNM).
Total .specimens examined: 4.

Discussion. The siruclnre of the male I'eni-

o

talia (Fig. 9) is similar to that of T. hnnellosus
(Fig. 8). Examination of more material mav in-
dicate synonymy between these two. The Le-
Conte specimens and the male in the (-asev
collection are small and have verv' sparse elon-
gate-oval scales on the elytra. The female from
the Argus Mountains of California lias denser
elongate-oval scales and is somewhat larger.

Tijclntis lamellosus Casey
( Figs. S, 21 )

Tijchiu.i liwwllosiis C;isev, 1892, .\iin. New York .Ac.ul.

Sci., 6:418-419 (Holotype: male. Utah, U.SNM

.367.1.5, T. L. Cisey collection ) .
Miccotrogus lamellosus: Klima, 1934, Coleonterorum

Calttlugus, 29 (1.38): 30.

■fhis species is distinguislied from other North
American representatives of the genus by the
multiple, confused, as oppos(>d to single, unifonn
median rows of long, narrow scales on the elytra!
interspaces; by the absence of fine, erect setae
on the abdomen; by the finely acuminate ros-
tniiii. and bv tiii- asymmetrical apical portion
of the HK'dian lobe of the male genitalia (Fig. S).
It is doubtfully distinct from T. .seini.s(iiuimosu.s
LeConte but can be distinguished by the char-
acters enumerated in the diagnosis of that
species.

Description. NIale: length 2.4-.'3..3 mm, width
1.2-1..") mm; integument black on pronotum, black
to pieeous on elytra, appendages light to dark
reddish brown; vestiture of white to dark reddish
brown scales.

Rostrum shorter tlian prothorax, antennal in-
sertion on distal fourth; in lateral aspect mod-
erately to slightlv, evenlv, arcuate in dorsal pro-
file; in dorsal aspect strongly tapered from base
to tip, frons 1.8-2.5 times as wide between dorsal
margin of eyes as rostrum at extreme apex; distal
portion finely acuminate, smooth, shining, pits
very shallow or absent. \'estiture of long, nar-
row, recumbent, or suberect, pointed or wedge
sliaped, strigose, light to dark reddish browTi
scales; distal portion glabrous except for sparse,
fine setae aroimd apical portion of scrobe.

Antennal funicle seven-segmented; pedicel
nearly as long as next three segments combined,
setae very fine, hairlike.

Pronotum wider than long, 1..5 times wider
at base than at apical constriction; sides evenly,
prominentlv arcuate. N'estiture complex, con-
sisting of narrow, dorsal, median vitta of long,
narrow, strigose, and round to oval nonstrigose,
white scales, usually extending to anterior mar-
gin but often confined to basal portion; long,
narrow to broad, recumbent, strigose, apically
pointed to truncate, light to ver\' dark reddish
brown scales covering dorsum and dorsal half
ol lateral surface; lower portion of sides with
round to elongate-oval, white to reddisli brown,
nonstrigose scales which extend dorsalK' forming
bro;id, lateral. \ itt.ie in dorsal aspect; usually
se\eral nonstrigose scales intermingled with long,
luurrow scales on dorsum.

[•llvtra parallel sided in basal two liiirds,
broadly rounded to apices, widest just lnhind
humeri; in dorsal profile nearly flat in basal half
to two-thirds, declivity broadly, evenb' rounded.
Vestiture on interspaces of ne;u'l\' uniform bi-
seri;ite to triseriate rows of round to elongate-
oval, slightK- imbriciited, reevunbeiit, nonstrigose,
white to very ii<'ht reddish brown scales, usually

Biological Series, Vol. 13. No. 3 Revi.sion of the Weevil Genus Tvchius

25

more hroadh' iiiiliricatt'd on intcnal.s one and
five through seven, darker in color and den.ser
on inter.space one; each interspace with confii.sed
uniseriate or multiseriate rows of long, narrow,
apicallv tiiincatc or pointed, light to dark red-
dish brown, suherect, strigose scales which are
usually denser on interspaces two through four;
setae arising from strial punctures narrow, light
colored, hairlike.

Wntral surface densely clothed with round
to elongate-o\aI, recumbent, broadly imbri-
cated, nonstrigose white scales, often some scales
slightly narrower and suherect, especially on
sterna four and five; no discrete transverse rows
of elongate, fine, hairlikc setae. Sternum five
with deep median fo\ea.

Femur elongate, narrow, apical half slightly
swollen. Apical, ventral, emargination weaklv
de\eloped; metafemur often with minute tooth
or spine on pro.ximal portion of emargination.
Vestiturc of elongate-oval, recumbent, white
scales and long, narrow, white or ver\' light red-
dish brown apicallv truncate or rounded, stri-
gose scales; no fine, erect, hairlike setae.

Tibiae mucronate, mucrones usually shorter
than tarsal claws, lacking obtuse tooth on dorsal
portion, usuallv largest mucro on protibia. Vesti-
ture of long, narrow, strigose scales, and fine,
erect, light brown setae on apical portion, rarely
with sparse elongate-oval scales.

Tarsi dorsallv with long, narrow scales, and
hairlike white setae, claw with basal process
about two-thirds as long as claw.

Male genitalia (Fig. S) with apical portion
of median lobe as^aninetrical; apical, dorsal,
median mebranous area elongate, strongly de-
fined posteriorly; median stmts weakly clavate.

Female: similar to male except rostrum
longer, more slender, antennal insertion median,
distal portion very smooth, long, finely acumi-
nate; slightly larger, length 2.4-3.4 mm; mu-
crones on tibiae slightly smaller.

Hosts. Astiagahis l)cckuithii, A. (Iruninioiulii,
A. lcnti<^iuosu.s var. jxiJcns, and A. Ioncliuc(ii])iis.

Distribution. (Fig. 21).

British Columhia: 01i\er, 2 mi. W., V-29-.58, H. liikI
A. How clen. "on vetch," 4 males. 3 females ( CNC ) .

Colorado: Boulder, VI-13-61, J. R. Stainer, 1 female
(CNC); Garfield Co., Grand Valley, 8mi. W.. VI-4-69,
W. E. Clark, A. lonchocarpus, 1 male, 3 females
(WEC); San Miguel Co., PlacerviUe, 4 and 6 mi. \.\V.
and 1 mi. N.E.. VI-3-69, W. E. Clark, A. lonchocarpus,
16 males, 29 females (WEC); LaPlata Co.. Bavfield, 16
mi. E., V-31-69, W. E. Clark. A, lonchocarpus, 29 males.
30 females (WEC); Archuleta Co., Pagosa Springs. 21
mi. W., V-31-69. W. E. Clark, A lonchocarpus. 7 males.
11 females (WEC).

New Mexico: Rio Arriba Co.. Chama, 17 mi. X.W .,

V-31-69, W. E. Clark. A. lonchocarpus, 7 males, 19
females (WEC); Rio Arriba Co., Cebolla, 2 mi. S. and
1.5 mi. S.W., V-31-69, W. E. Clark, A. lonchocarpus, 60
males, 65 females (WEC); Rio Arriba Co., Coyote, 8 mi.
W., VI-1-69, W. E. Clark. A. lonchocarpus,'! females
(WEC); Sandoval Co., Jemez Springs, 3 mi. N.E., VI-
1-69, W. E. Clark, A. lonchocarpus. 12 males, 5 females
(WEC); 1923, Edith W. Mark, 4 females (CI); Jemez
Mts., VI-4. and VI-26, |. Woodgate, 6 males, 8 females
(CAS), VI-4, Shoemaker. 2 males (USNM); Ft. Win-
gate, VI-VII, 5 females (USNM).

Utah; Utah Co., Provo, 1 mi. S.E., V-15, 22, 24-
66, 67. 68, W. E. Clark. A. heckwithii, 71 males, 50 fe-
males (WEC); Utah Co., Provo, Mouth Rock Canyon,
V-3, 10, 21, 69, W. E. Clark, A. heckwithii, 113 males,
110 fcnules (WEC); Utah Co.. Lehi, 13 mi. W., V-10-
69, W. E. Clark, A. heckwithii, 3 males, 1 female
(WEC); Utah Co., Provo Canyon, VI-1-68, W. E.
Clark, 1 male (WEC); Arches National Monument,
\'-10-6S, W. E. Clark. A. Icntijj^inosus var. jialcns, 9 males
6 females (WEC).

Wvoining; Fremont Co.. Lander, 14 mi. S., VI-14-
69, W. E. Clark, A. drummondii, 12 males, 10 females
( WEC ) .

Total specimens examined: 677.

Discussion. Specimens examined from Utah
have the antennal insertion very near the tip of
the rostrum as do specimens from British Co-
lumbia. The most noticable variation is in the
color, shape, and density of the long, narrow,
reddish browai scales on the pronotum and elytral
interspaces. In specimens from Provo, Utah, these
scales range from nearly the color of the round
nonstrigose scales to very dark reddish brown
with ;i corresponding darkening of the round,
nonstrigose scales. In all specimens examined
these scales have bluntly pointed apices. Speci-
mens from southwestern Colorado and north-
western New Mexico have the elongate scales
rather light orangish brown and very broad and
rounded at the apices. Specimens taken on A.
(Intmmondii near Lander, Wyoming, are yery
light in color, with the long, narrow scales barely
darker than the round, nonstrigose scales. Speci-
mens from British Columbia are very similar to
the Wyoming specimens. Specimens from Arches
Nation;d Monument, Ut;di, average smaller in
size than Provo specimens and more nearly re-
semble T. .semiscjtiumosxis in distribution of scales
and size.

Color and siiape of the long, narrow scales
\iuy in a dine from British Columbia, where
they are narrow and light in color, to New
Mexico, \\'here they are broad, darker, ;md
oransjish browai.

Ti/chius had ins, n. sp.

(Figs. 5. 21)

This species is probably closely related to
T. himellosiis Casey, but differs by the following

26

Buiciiwr Young I'nivf.hsity Sc:ii;nce Bulletin

c-liiiracters: tlio median lobe of the male geni-
talia has lateral prominences on the apical por-
tion (Fig. 5); the long, narrow, scales on the
elvtral interspaces are nearly always in single,
uniform median rows while in 7". lamcllosiis they
are usually in multiseriate rows; tiie basal por-
tion of the rostrum is wider in lateral aspect; and
the scales on the elytra are broader ;uid more
densely imbricate.

Description. Male: length 3.1 mm, width 1.5
mm; integument black to piceous, appendages
light reddish brown. X'estiturc of crcamN' white
to reddish brown scales.

Hostrum slight Iv shorter than prothorax;
swollen in basal fourth then slightly arcuate in
apical two-thirds; antenna! insertion in apical
third; distad of antcnnal insertion smooth, shin-
ing, with sparse, shallow, lateral impressions;
finch' acuminate to ape.x; in dorsal aspect promi-
nenti\- t;ipcred from base to apc.x, frons 1.7 times
as wide between dorsal margin of eyes as ros-
trum at apex. Scales above antcnnal insertion of
unilomi size and shape, long, narrow, recum-
lieiit, apicallv roundid, no erect hairlike setae.
Scrobc with elongate, sulxuect setae around an-
terior margm; extreme apical portion with sparse.
yery fine setae.

.Antcnnal funiculus seven-segmented; pedicel
nciirlv twice as long as next two segments com-
bined.

Pronotum 1.2 times as wide as long; sides
prominently arcu;itc, 1.6 times wider at base than
at apical constriction. Scales of two distinct
types, long, narrow, pointed, light reddish brown
scales on dorsum and halfway down sides;
median dorsal vitt;i of bro;id, oval, white scales;
lower portion of sides with o\;d, white scales
wlu'ch extend dorsidly and are densely inter-
mingled with the long, narrow scales on dorsum.

Elytra 1.6 times as long as wide, sides paral-
led to apical fourth then evenly rounded to
apices; nearly flat in basal two-thirds in dorsal
profile. Each interspace with double or triple
rows of round to elong;ite-oval, light brown,
densely inibricale. recumbent scales; sc;des on in-
terspace one denser and more bro;idl\' imbri-
cate. Each interspace with single medi;ui row of
long, narrow, recumbent, light rctidish sc;des,
single rows breaking up into irreguhu- clusters
b;is;dly on interspace two, around humeri. ;uid
at scattered places all over elytra. Stri;il setae
narrow, hairlike, white.

\'entral surface with round to oval, white
sc;des; those on median surface of \enter with
plumose m;irgins; some scales on apical portion
of ;ibdomen elongate and suberect; no distinct

Inmsvcrse rows of erect, tint' setae. Sternum five
with shallow median fovea.

Femur with prominent, ;ipical, yentnd cmar-
gin;ition; sometimes with minute tooth on proxi-
mal portion of emargination. \'estiture ot round
to oval, nonstrigose ;uid long, n;irrow, strigose
scales.

Tibiiie mucronate, mucrones shorter than
trasal claws; vestiture of long, narrow scales and
suberect fine setae.

Tarsi clothed with elongate setae; claws with
broiid, conn;itc b;is;d processes.

Male genitalia (Fig. 5) with prominent lat-
end prominences on apical portion of median
lobe; ;ipit;d, dors;d, median membranous area
nearly round, sluirply defined posteriorK', median
struts very stout, not chivate.

Female: slightly longer, rostrum longer, more
finely tapered, 1.9 times as wide at ;ipex as frons
between dorsal margins of eyes, antenn;d inser-
tion median.

Type Locality. COLOR.VDO: I.;iPlata Co..
Miuicos , 7 mi. K.

Type Material. Male h(ilot\ pe, female allo-
t\pe, 15 males ;ind 1 1 fem;des, parat\ pes taken
;it the type loc;dity, Ma\- ;3(), 1969, ■l)y W. E.
(]|;irk, sweeping Astrap^ahis scojudoniui. The
holotxpc and allot\pe ;ue deposited in the U.S.
National Museum, \\'ashiugton, D,(!.; two male
and two female parat\ pes are deposited in the
Brigham Young Universitx' collection, I'rovo,
Utah; the remaining 2.3 parat\pes are retained
in the ;iuthor"s collection.

Distribution. (Fig. 21).

Ci)lcii;ul(>: .\Rlnilfta Co., P.ii;<)sa .Springs, 1.5 mi.
.S K., \-:31-69, W. E. Clark, 2 males, 19 females, Astm-
f,'(//i/,v hi.sulciittis \ar. hcijdenUinus, (WEC); "Colo.."
Wirkhani rollc-c-tion. 1 male (USN'M).

Total specimens examined: .50.

Discussion. There is a conspicuous tlilfercnce
in the size of the Mancos and P;igosa Springs
specimens. Males from Mancos range from 3.0-
■3. 1 mm, and females range from 3.2-3.4 mm in
Icnglh; males from Pagosa Springs range in
liiigth from 2.5-2.8 mm, females from 2.6-3.0
niuL Males from Pagosa Springs average 0.52
mm, and fem;des 0..55 mm shorti-r than speci-
mens from Mancos.

rt/cliiii.s prolixu.s (';Lsey
(F-igs. 6, 21)

iiichiii.t pnilixua Ca.sev, 1892, Aiiil Neu York .\ea(l.
Sei.. 6:419-420 (Holotvpe: male, Nevada. U.SNM
36756, T. L. Casey eollcction). Tanner. 1966, BYU
Sci. Bnll.. Biologieal .series, 8(2) :26. Klima, 1934,
Colcoptcrurum Catalogiis, 29(138):21.

Bioi.or.rcAL Skhies, Vol. 13. No. 3 Revision of the Weevil Genus Tychius

27

Ti/chiiix (Piiwli/chiiis) yirn/i'.vii.v C^iscv. 1910, CaiL Eiito-
niol.. 42:13.5 ( cstahlished pidlixu.s- as l\pc ot suli-
gfiuis Ptirutychius).

Tycliiiis (Piirati/chius) imbricatus Casey, 1910, Can.
Entomol.. 42:135-136 (Holotype: female, San Diego,
California, USNM 36757, T. L. Casey collection).

This .species most closely resembles T. himcl-
lostis but can be easily distinguished by the six
radier than seven antennal funicular segments,
bv the denser, more broadly imliricated scales
on the ehtra, and bv the synimetrical apical
portion of the median lobe of the male genitalia
(Fig. 6).

Description. Male: length 2.6-4.0 mm; integu-
ment piceous to black, rostrum and appendages
light to dark reddish brown; vcstiturc of white
to dark reddish brown scales.

Rostrum nearly as lonsr or slightly lonsjer than
prothorax; antennal insertion in distal third; near-
ly straight distally; from dorsal aspect strongly,
evenlv tapered from base to apex, frons 2.0-2.5
times wider between dorsal margin of eves than
rostrum at extreme apex, distal portion tapered,
not finely acuminate, smooth, shining, pits dense,
shallow. \'estiture of stout, broad, to long, nar-
row, wedge-shaped to apically rounded, decum-
bent, strigose, white to dark reddish brown
scales; no distinct, erect, hairlike setae; distal
portion glabrous except for sparse, small scales
around ;ipical portion of scrobe.

Antennal funiculus six-segmented; pedicel
about equal in length to next three segments
combined; setae long, hairlikc to clavate.

Prontum wider than long, about 1.6-2.0 times
wider at base than at apical construction; sides
slightly arcu;ite in b;is;d third, strongly arcuate
indistal third. W'stitiire complex, dorsal median,
\itta, of long, narrow, strigose, and round to
o\al, nonstrigose white scales; long narrow, re-
cumbent, strigose, apically truncate to rounded,
light to very dark reddish brown; strigose scales
covering dorsum imd upper h;df of sides; lower
portion of sides with round to elongate-oval,
white to light reddish brown, nonstrigose scales
which extend dorsally, forming broiid Literal
vittae from dors;il aspect; usually several non-
strigose scales interniin<rled with lon<?, narrow
scales on dorsum.

Elytra p;irallel sided on basal two-thirds,
broadly rounded to apices; in dorsal profile near-
ly flat in basal half to two-thirds; declivity
broadly, evenly rounded. Vcstiturc on interspaces
of nearly uniform biseri;ite to triseri;ite rows of
round to oval, broadly imbricated, recumbent,
noastrigose, white to reddish brown scales, den-
ser and darker on interspace one, usually darker

in color on interspaces two through foiu'. Each
interspace with imiform median rows of long,
narrow, recumbent, apically truncate or roimded,
light to dark reddish brown, strigose scales, usu-
ally in uniseriate rows on intervals one, three
and fiye to seven; multiseriate rows on inter-
spaces two and four. .Strial setae ninrow, white,
hairlike.

Ventral surface with round to elongate-oval,
recumbent, broadly imbricated, nonstrigose white
scales; often several scales suberect, especially
on sterna four and five; sometimes with discrete
transverse rows of elongate, fine hairlike setae.
Sti'rnum five with deep median fovea.

Femur elongate, apical half moderately swal-
len; apical, ventral cmargination usually promi-
nent; metafemur often with minute tooth or spine
on proximal portion of cmargination. Vcstiturc
of elongate-oval, recumbent, white scales and
long, narrow, strigose, light, reddish brown, api-
cally truncated or rounded, strigose scales; no
fine, erect, hairlike setae.

Tibiae mucronate, mucro usually shorter than
tarsal claw; lacking obtuse tooth on dorsal por-
tion; usually mucro on protibia largest. \'estiture
of long, narrow, strigose, white to light reddish
brown scales, elontrate-oval, white nonstrigose
scales, and fine, erect, light brown setae on api-
cal portions.

Tarsi dorsally with long, narrow scales and
hairlike, white setae; claw with basal process
about two-thirds as long as claw.

Vlale genitalia (Fig. 6) with apical portion
of median lobe rounded; apical dorsal, median
membranous area nearly round, strongly defined
posteriorly; inedi;m struts fine, clavate.

Female: length 2.7-4.1 mm; rostrum longer,
more slender, often straighter; antennal inser-
tion median; tibial mucrones slightly smaller.

Hosts. Astni<!^(ilus (n)ij>hioxijs\ A. dotii^lasii, A.
iitdlwnsi.s. and A. Icntiiiinosiis (Tanner, 1966).

Distribution. Fig. 21).

Arizona: Morrison, Hubbard and Schwarz, 1 male
(USXM); (Sta. Rita, N.F.), M. Chrisman, Jiinipcnis. 1
male (U.SNM); Snovvflake VIll-1-30 Ballantyne, A.stm-
•idliis diphysiis, 1 female (UA), V-27-32, E. F. Rnssell,
"swept from loco weed," 1 female (USNM); St. John,
VI-7-32, E. F. Rus.sell, "reared from loco plants," 2
males (USNM).

California: 2 males (UK): Lielieck. 2 females
(PANS); Aguanga, V-12-29, 1 male (CNC); Antioch,
V-18-36, 1 female (CNC); Chino, VlI-20-08, 1 female
(USL); Elsinore, III, A. Feynes Coll., 2 males, 1 female
(CAS): Hemet Reservoir, San Jacinto Mts., V-22-40. 1
fem.ile (CIS); Jacumba, lV-17-16, J. O. Martin, 2 males,
1 female (CAS): Riverside Co., Keen Camp, Vl-6, 12-
17, E. P. Van Duzce, 1 male (CAS); 4 mi. E. Keen
Camp, Hemet Res. 4500', VII-1-65, C. D. Johnson,

28

Bmicham Vounc; Univebsity Science Bulletin

Aslragiihis douglasii, 7 males, 3 females ( CWO ) ; Lagiina
Mts., VII-6-29. R. H. Beamer, 1 females (UK); San
Diego Co., La Mesa, III-20-2«, H. G. Barber, 3 males
(USNM); Leijec, IV-15-28, A. C. Davis, 2 males
(CNC), 1 male (CNC); Los Angeles Co., Cociuillett
Coll., 5 males. 2 females (USNM), Van Dyke Coll.,
2 males (USNM); San Diego Co., Paeific Beaeh, VI-23-
37, C. Gammon, "re;irecl from loco weed," 1 female
(CAS); Riverside, 111-18-40, C. Bammeres "flowers of
loeo weed," 1 male, 3 females ( LA ) ; 2 males, 2 females
(USNM); San Diego Co., lV-8-2.5, K. C. Van i:)yke, 1
male. 5 females (CAS); F. E. Blaisdell, 1 male, 2 fe-
males (CAS), F. H. Parker Coll., 4 males, 2 females
(UA), D. K. Dnncan, 2 males, 2 females (UA); IV-2,3-
20, E. P. Van Duzee, 1 female (CAS); Riverside Co.,
San Jiieinto Mts., VII-20-29, I male (UK); San Luis
Obispo. III-14-08, I. J. Condit. Aslra<!,(ilus. 2 males, 3
females (USNM); S. Bernardino Co., 2 m;des, 4 females
(USNM); Buena Ventura, Liebeck Coll., 8 males, 4 fe-
males (MCZ); Upland, IV-,5-20, A. Feynes Coll., 2
males, 2 females (CAS); IV, Wiekham Coll. 1 ni;ile,
2 females (USNM), 1 fem;ile (.MCZ); Monteiev Co.,
King Citv, IV-2-60, C. A. Tosehi, one male (CWO).

New Me.\ico: W. G. Dietz, 1 male (MCZ); Sando-
val Co., San Vsidro, 13 mi. N.VV., VI-1-69, W. E,
Clark, Astragalus amphioxi/s, 9 males, 4 females
(WEC).

Texas: Brewster Co.. 17 mi. S. Alpine. lOOO', \'I-6-
70, L. and C. W. O'Brien, 1 m;de, 1 female (CWO);
Davis Co., 10 mi. S. Tovahvale, \-31-70, R. NL Murray,
1 female (WEC).

Utah: Utah Co., Provo, Mouth Rock Canyon, V-19-
69, W. E. Clark, Astragalus titaliciisis, 10 males, 9 fe-
males (WEC); Utah. Co., Provo, V-9-67. \'-27-67, D.
R. Harris, Astragalus utahensis, 7 males, 1 femali'
(WEC).

Total specimens examined: 1.53.

Discussion. Specimens from Utah and New
Mexico have longer and straighter rostra in both
sexes than do specimens from California. The
possession of six rather than seven antennal
funiciihir segments does not justify giving this
taxon the rank of subgenus because of its ap-
parent close reiiUionship to other inemliers of the
T. semisijuamofius species group, espcci;iily T.
lameUosus.

Tijcliitis sollutii (,'asey
(Figs. 11, 20)

Ttjchius soltaui Casey, 1892, Ann. New York Acad. Sci.,
6:416 (Holotype: male, Laramie, Wyoming, USNM
36734, T. L. Casey collection ).

Miccotrogus soltaui: Klima, 19.34, Coleoptcroruni Cata-
logus, 29(I38):32.

This is tile most widely distributed species
in a group of ven- closely related species within
the 7'. .seinis(iu(inios-us species group. This species
can be distinguished from other members of the
T. .vcmi.sf/»r/»io.v(/.v species group by the posses-
sion of erect hairlike setae on the abdominal
sterna. The male genitalia (Fig. 11), which lack
the ;ipical lateral prominences of T. Jnidins ( Fig.

5) and 7'. hirsuttis (Fig. 14) ;ind have the api-
cal median niembnmous ;uc;i sluuplv defined
posteriorlv nither than not sh;irpiy detined ;is in
T. phalarus (Fig. 7), serve to distinguish T. sol-
taui from other members of the group.

Description. \l;ile length, 2.fi-.3.S mm, width
1.2-1.8 mm; integument piceous to l)iack, ap-
pendiiges piceous to light reddish brown. Vesti-
ture complex, of white (o (Uirk reddish lirown
scales.

Rostrum shorter than prothorax, length of
rostrum 20 to 2.5 percent of total bodv length;
;uiti'nnal insertion in apie;d fourth; usually
strongly arcuate in b;is;il luilf, nearlv straight
;i]oieallv, but often eveiiK arcuate or straight
entire length; from dorsal aspect tapered evenlv
from base to apex, frons 1.7-2.5 times wider
between dorsal margin of eyes at extreme apex,
distal jiortion tapered but not strongly acumin-
;ite, smooth ;uid shining, large, shallow pits espe-
cially dense lateralh. N'estiture of long, narrow,
;ipicall\' truncate or rounded, strigose scales, usu-
;dl\' scales on lower portion of sides smaller,
lighter colored, white to lisrht reddish brown,
scales on dorsal portion dark reddish brown
with several lighter colored, narrow, erect or
suberect setae especially dense distallv.

Antennal funiculus seven-segmented; pedicel
as long or longer than next three segments
combined; setae broad, elongate.

Pronotum wider th;m long l.fi-I.S times wider
;it base than at iipical constriction; sides in
dorsal aspect evenlv, bnxidlv rounded. Vesti-
ture of narrow, dors;i]. median vitta of long,
n;irroy\', strigose, and round to oval white scales;
long, narrow, light to very d;nk reddish brown,
strigose scales covering inost of dorsum and up-
per portion of sides; lower portion of sides with
round to elong;ite-ovaI, \\hite or light reddish
i>n)\\n, nonstrigose scales which extend dorsallv
forming broad, lateral vittae from dorsal aspect,
nonstrigose scales usualh' intermingled with
long, narrow sc;des on dorsum.

El\tr;i p;irallel-sided in l)as;d two-thirds,
broadh' rounded to apices; in dorsal profile
ne;u'I\' tl.it on ilisc, declivity bro;idl\' e\(.'ul\'
roiuuled. N'estiture on interspaces of roiuid to
o\<d. br();idl\ imbric;ited, recumbent, nonstrigose,
white to d;uk ri'ddish brown scales, lighter in
color on interspaces five to eight; each inter-
space with unifonT) row of scales simikir to
long, narrow scales of pronotum. Stri;d setae
n;u'rower th:ui sc;tles on inlers[);ices.

X'ciitr.il suihici' wllh round to elong;iti'-oy;d,
white to light reddish brown, recinnbent, imbri-
cated scales; each abdominal sternum with dis-

Biological Skries, Vol. 13. No. 3 Revlsion of the Weevil Genus Tychius

29

Crete traii.s\erse niw of erect, narrow, liairlike
setae; metasternum witli .some elongate, recum-
bent, narrow setae; sternum fi\e witli dee]')
median fovea.

Femora long, narrow to stout, apical portion
swollen, ventral emargination well developed in
specimens with stout apical portion; metafemur
often with minute tooth or spine on pro.ximal
portion of apical, ventral emargination. Vestiture
of elongate-o\al, recumbent, usually light to dark
reddish brown scales, and long, narrow, strigose,
suberect, usuallv white or very light reddish
brown scales.

Tibiae mucronate, mucrones usuallv as long
as tarsal claws, often with obtuse tooth on dorsal
portion, usuallv largest on protibia. Vestiture of
elongate, oval, and lonsr, narrow scales, and fine,
hairlike setae.

Tarsi dorsalh' with lone, narrow scales;
claws with basal process short, about half as
long as claw.

Male genitalia (Fig. 11) with apex of median
lobe obtuseh' rounded; apical, dorsal, median
membranous are a small, transverselv oval,
strongly defined posteriorlv; median struts slight-
ly clavate.

Female: length 2.6-.3.9 mm; rostrum more
finely tapered distad of antennal insertion; mu-
crones on tibiae usuallv smaller.

Hosts. Astragalus flavtis var. flavtis and A.
flextiostis.

Distribution. (Fig. 20).

Arizona: C. V. Riley, 1 male (USNM); Peath
Springs, VIII-2.5, C. W. Leng, 1 female (BYU).

Colorado: Denver, IV-2, H. Soltaii, 1 m.ile
(USNM); Platte Can., X-27-1889. H. Soltau. 1 male
(USNM).

Manitoba: Aweme, VI-26-30, R. M. White, Astra-
galus flexuosus, 1 female (CNC).

Montana: 1 male (INHS), 1 male (USNM), 1 fe-
male (PANS); Helena, Hubbard and Sehwarz, 1 female
(USNM).

Nebraska; Indianola, H. Soltau, 2 males (USNM).

New Mexico: Sandoval Co., San Ysidro, 1.3 mi.
N.W., V-1-69, W. E. Clark, 1 male (WEC).

North Dakota: Tower Citv, VI-3-().5, G. I. Reeves,
1 female (USNM).

Saskatchewan: Last Mtn. Lake, VI-5-33, Wickhani
Coll., 1 male (USNM).

Soudi Dakota; Pennington Co., Pactola Reservoir,
\'I-17-6,S, W. E. Clark, 1 female (WEC).

Texas: Davis Mts., IV-26-24, J. O. Martin, 1 male,

1 female (CAS); Upton Co., Rankin, VI-3-70, C. W.
Neeb, 1 male (TAM).

Utah: Cache Co., Logan, \'-24-.51, H. G. Egoscue,

2 females (BYU); Uintah Co.. Vernal, 14 mi.' S.W..
V-17-69, W. E. Clark, A.rtr«^«/w.v flaviis, 10 males, 8
females (WEC).

Wvoming; Albanv Co., Laramie, 20 mi. N.W., V-
6-69, \\'. E.' Clark, Aslr,i<iulii.<< flatus, 13 males, 8 fe-
males (WEC); Fremont Co., Lander, 14 mi. S., VI-14-

69, W. E. Clark, I male, 1 female (WEC); Carbon Co.,
Medicine Bow, 32 mi. N., VI-6-69, W. E. Clark, 1 male,
1 female (WEC).

Total specimens e.xiimined: 63.

Discussion. The holot\pe is small, .3.0 mm in
length, unusualh' narrow, and dark in color.
Specimens taken in New Mexico, Arizona, and
Colorado are often somewhat larger in size but
agree in characters of the male genitalia. Most
of the specimens at hand are single or at best
pairs of specimens from widely separated areas.

I have examined a female from Montana in
the Casey collection identified as T. aratiis. but
it is actually T. soltaui. This specimen is fairly
large and the erect setae on the intervals are
fini'r th;in those of the type of T. soltaui.

Tijchius inontanus n.sp.

(Figs. 18, 20)

This species appears closely related to T.
soltaui Casey. The most reliable character for
separating the two species is the structure of the
apical portion of the median lobe of the male
genitalia ( Fig. IS ) which possesses weakly de-
veloped lateral, apical, promineiices in T. mon-
tantis but not in T. soltaui. The rostrum is gen-
erally as long or slightly longer than the pro-
thorax, nearly straight or very slightly arcuate,
and usuallv expanded at the extreme apex. The
pits on the distal portion of the rostrum are
slightly deeper than in T. soltaui. The median
dorsal p;itch of white scales does not extend the
entire length of the pronotum, ;is in T. soltaui,
but forms a small, basal patch. The absence of
erect hairlike setae or recumbent, long, narrow,
white scales on the metasternum and the first
visible ;ibdoniinal sterna is also diagnostic.

Description. M;de: length 3.4 mm, width 1.5
mm; integument light reddish brown, darker on
dorsal surfaces. Vestiture complex, of white to
dark reddish l:)rown scales.

Rostrum slightly-shorter than prothorax, ros-
trum length about 28 percent of total body
length; iintennal insertion in apical third, evenly,
sliglitly arcuate from base to apex in dorsal pro-
file; slightlv tapered from base to apex, some-
times slightly expanded at ape.x, frons about 1.5
times wider between dorsal margin of eyes than
rostrum at apex; distal portion stout, not more
strongly tapered than proximal two-thirds; distal
third densely, deeply rugulose. Vestiture of long,
narrow, apically, rounded, strigose scales; scales
on dorsal portion dark reddish brown, several
lighter colored, slightly narrower, suberect scales
especially dense distally.

30

BiuGiiAM Young University Science Bulletin

AntcMinal funide seven-segmented; pedicel
as long as next three segments combined; setae
long, narrow.

Pronotiini about 1.2 times wider than long,
1.6-1.8 times wider at base than at apical con-
striction; sides prominently, evenly arcuate.
Vesfiture complex, dorsum with small basal patch
of round, and long, narrow, white scales; long,
narrow, often suberect, light to veiy dark red-
dLsh brown, apically truncate or pointed, stri-
gose scales covering most of dorsinn and up-
per half of sides; lower portion of sides with
round to elongate-oval, white to light reddisii
brown, nonstrigose scales whicli cxttMid dorsally
forming broad, lateral, liglit colored vittae from
dorsal aspect.

Klvtra parallel sided in basal two-thirds,
broadh- roumled to apices; in dorsal profile near-
Iv flat to \ery slightK- rounded in basal half; de-
clivity broadly, e\eni\- rounded. X'estiture on in-
terspaces of nearlv uniform biseriate or trisei^iate
rows of round to oval, broadlv imbricated, re-
cumbent, nonstrigose, white to dark reddish
brown scales, sliglit!\' darker on intervals two
through four, lighter and denser on interspace
one; each interspace witli imifonn row of long,
n:u"r()w, usuallv apicallv pointed, suberect, light
to dark reddisii brown, strigose scales, each scale
shorter than width of interspace. Strial setae
lighter in color and slightlv narrower than long
narrow scales on intersp;ices.

\'entral surtace with round to eIongate-o\al,
white, recumbent, nonstrigose scales; abdominal
sterna three and five each with discrete trans-
verse row of erect, hairlike setae, which are ab-
sent from metathorax and abdominal segment.s
one and usually two. Sternum fi\c witli deep
median fovea.

Femora long, narrow or swollen apic;illv;
apical, \entral eiiuugination usuallv well de-
xciopetl; metalemnr often with minute tooth or
spine on proximal portion of emargination. Vesti-
ture of elongate-oval, recumbent, usually light
reddish brown, nonstrigose scales and long, nar-
row, strigose, usually suberect, lighter colored,
often white scales.

Tibiae mucronate, nnicro on protibia about
as long as tarsal claw, mucrones on mesofemur
and metafemur smaller, usually mucrones with
obtuse dorsal tooth. Vestiture of elongate-oval,
usually light reddish brown .scales, long, narrow
white scales, and very fine, hairlike suberect setae
especially dense on apical and \entral portions.

Tarsi dorsally with long, narrow scales; claw
with basal process about half as long as claw.

Male genitiilia (Fig. IS) with weaklv devel-
oped Literal apical prominences on median lobe;

apical, dorsal, median membranous area nearly
round, strongl}' defined posteriorly; median
struts very fine, strongly clavate.

Female: rostrum length 27 percent to 29
percent of total bodv length; total body length
.3.f-.3.5 nini; ;intciin;il insertion median.

Type Locality. MONTANA: Helena.

Type Material. Mal(> holot\pe, female allo-
t\pe, one male and one female paratvpe taken
at the t\pe locality by Hubbard and Schwarz;
on 1-5, all but the male paratype are deposited
in USNM; the male paratype is in my personal
collection.

Host. Unknown.

Distribution. (Fig. 20).

In addition to the t\pe material, two speci-
mens from tlie following localities were exam-
ined:

.\lbert.i; Medicine Hat, VI-1-34. J. Cair, 1 female

(HVU).

North D.ikdl.i: Mandan, F. E. Cobb, 1 female
(USXM).

Tot.il speeimeiis ex.imined: 6.

Ti/cliitis liirstitus. new name

(Figs. 14, 20).

Ti/chiu.s lurtctlus LeConte (not Tournier, 1873), 1S76,
Proc. Amer. Philos, See, 15:218 (Lecototype here
designated: female, Texas, MCZ type 52282).

Miccutrogus hirtellus: Klima, 1934, Coleopterorum Ciita-
logus, 29(1 38): 30.

This species can be distinguished from its
North American relatives bv the veiT fine,
elongate, hairlike setae on the interspaces of the
elvtra, rostnim and appendages. It closelv re-
sembles T. soltinii Casey but can be distin-
guished from tliat species bv the apical, lateral,
])rojections of the median lobe of the male geni-
t:ilia (Fig. 14). The rostrum is more finely acu-
minate in the distal portion, the scales on the
pronotum are narrow, leaving the integimient
l)roacl!\- \isii)Ie, and the variation in color be-
tween interspaces two to three and four to seven
;i|5parc'nt in 7'. sollinii is alisent.

Description. Male: k'ngtii 2.6-2.9 mm; integu-
ment piceous to black, appendages piceous to
light reddish brown. Vestiture complex, of white
to dark reddisii brown scales.

Rostrum shorter th:ui jirotliora.x, ;iiiteniial in-
sertion in apical fourth, usualK e\eiil\, slighth
to moderateh' arcuate in dors;d profile but some-
times prominentK ;ucuate in basiil half, nearh'
straight in distal half; in dorsal aspect tapered
evenly from base to apex, Irons 2.0-2.5 times

Biological Series, Vol. 13, No. 3 Revision ok the Weevil Genus Tychius

31

wider hetwi't'n doisal margin ot c\es than ros-
trum at apex; distal portion stronglv tapered,
often fineh' acuminate, pits and rugae shallow.
\'estiture of elongate, verv narrow, apicalh
pointed or fineh' acuminate, light to dark reddish
brown scales on dorsum and upper portion of
sides, usually with smaller lighter colored nar-
row scales and several light colored round to
oval, nonstrigose scales on lower portion of sides;
suberect, elongate, hairlike white setae proximad
and distad of antennal insertion.

Antennal funicle seven-segmented; pedicel
about equal in length to or longer than ne.xt
three segments combined; setae verv fine, elon-
gate.

Pronotum wider than long, 1.6-1.7 times
wider at base than at apical constriction; sides
evenly, broadly rounded, or nearly parallel in
basal half, rounded acutelv in distal portion.
\'estiture complex, of narrow, dorsal, median
vitta of long, narrow, and oval to elongate-oval
white scales; elongate, narrow, often semierect,
light to dark reddish brown, strigose, scales cov-
ering most of dorsum and upper half of sides;
integument visililc between scales; lo\\er portion
of sides with round to elonizate-oval, white to
dark reddish brown nonstrigose scales which ex-
tend dorsallv forming broad, lateral vittae from
dorsal aspect; usuall\- with several oval sc;iles
intermingled \\ ith long, narrow scales on dorsum.

Elvtra usually broadest at humeri, tapering
slightlv to apices; in dorsal profile nearh' flat on
disc; decliv it\ broadh', evenly rounded. \'esti-
ture on interspaces of round to oval, broadly
imbricated, recumbent, nonstrigose, \\hitc to
dark reddish brown scales, slightlv denser, and
often of different color on interspace one, no
obivous color differences on other interspaces;
each interspace with uniform median row of
narrow, elongate, pointed, erect, white to dark
reddish brown, setae.

\'entral surface with round, to elongate-oval
recumbent, imbricated, white to light reddish
brown, usually plumose margined scales; each
abdominal sternum with discrete transverse row
of erect, fine, white, hairlike setae; metasternum
usually with some elongate, recumbent or sub-
erect, narrow setae, but often with erect very
fine setae. Sternum five with deep median fovea.

Femora long, narrow, apical portion not
prominently swollen, apical ventral emargination
weakly developed; metafemur often with minute
tooth on proximal portion of apical ventral emar-
gination. \'estiture of elongate-oval, recumbent,
light to dark reddish brown scales, and elongate,
pointed, hairlike, erect, white setae, no long,
narrow, strigose scales.

Tibiae mucronate, mucro usually as long as
tarsal claw, often with obtuse tooth on dorsal
l^ortion, mucrones usually of uniform size on all
tibiae but often largest on protibia. Vestiture of
clongate-oval, recumbent scales and fine, hair-
like, erect setae.

Tarsi dorsalh with long, narrow, pointed,
scales; claw with basal process about two-thirds
as long as claw.

Male genitalia (Fig. 14) with apical portion of
median lobe bearing prominent lateral projec-
tions; apical, dorsal, median membraneous area
nearly round, strongly defined posteriorly; medi-
an lobe very long in comparison to median
struts, verv heavily sclerotized; median struts
clavatc.

Female; length 2.7-3.0 mm; antennal inser-
tion median, finely acuminate distallv; tibial
mucrones slightlv smaller.

Hosts. Specimens bearing the following host
data have been examined: Astragalus mittalli-
aiuis. beating Qucictis, and Prosojiis juUflora.

Distribution. (Fig. 20).

New Me.xico: Albuquenjue, VI-27-33, Wickham
and Bowditch. 1 male (USNM), 1 female (MCZ).

Te.vas: 2 females (PANS); Belfrage, Hubbard and
Schwarz, 2 males (USNM); C. V. Riley, 1 male, 2
females (USNM); Bastrop Co., Vl-31-58, H, R. Burke,
1 female (TAM); Brazos Co.. VI-20-60, H. R. Burke,
1 female (TAM); Collinsworth Co., V-18.59, 1 female
(TAM; Corpus Christi, III-30-54, D. J. and J. N.
Knull, 1 female (OSC); Dallas, V-ll-.5'0, E. E. Gil-
bert, 2 males, 1 female (CIS); Dallas Co., lV-18-40,
Knutson, 1 female (CAS); Gillespie Co., VI-1-58, S.
Burke, 1 female (TAM); Kerrville, IV, V, VI-4, .5,
18-52, .55, L. |. Bottimer, Astragalus, V-4-52, 3 males, 7
females (CXC), IV-4, 13, 20-,59, Becker and Howden,
lieatins; Qiicrcus, 2 males, 1 female (CNC); Llano, IV-
21-06.' F. C. Pratt, 1 female (USNM); Marfa, VII-11-
12, J. \V. Green, 1 female (USNM): San .-Antonio, V-31-
03, A. C. Morgan, Prosopis juliflora. 1 female (USNM);
Jim Wells Co.', 7 mi. W. Alice, III-29-70, W. E. Clark,
A.stiaguhis nuttallUiiiiis, 5 males, 4 females (WEC).

Total specimens examined: 46.

Discussion. The range of this species and
T. soltaiti overlap in western Texas and New
Mexico. Some specimens examined from the area
have weakly developed lateral apical promi-
nences on the medi;ui lobe of the male genitalia
which suggests possible intergradation between
the two. More study is necessary to detennine
;iccuratelv the relationship between them.

Tt/chitis j)hatanis, n.sp.

(Figs. 2, 7, 20)

This species closely resembles T. soltaui
Casey. The most reliable character for separating
the two is the apical portion of the median lobe

32

BmcMAM Young Univkhsity Science Bulletin

of the male genitalia ( Fig. 7). The apical, medi-
an membranous area extends proximad for the
greater portion of the length and does not have
a distinct posterior limit as in T. aoltaui { Fig. 11).
The white scales on the median portion of tlu>
dorsum of the pronotum are restricted to a prom-
inent basal patch (Fig. 2) instead of fonning
a median vitta the length of the pronotum. In
most specimens the rostrum is slightly expanded
in the extreme distal portion, scales on the ros-
trum are elongate-oval and pointed ratlier than
parallel sided, scales on tlu- pronotum are nar-
row and dark in color. o\al scales on the femur
are white, and the long, narrow scales are dark
in color.

Description. Male: ieiigtli 2.S-'15 mm; in-
tegument black to piceous appendages iiglit to
dark reddish brown. \'estiturc of white to dark
reddish brown scales.

Rostrum slightJN' shorter or al)out same length
as prodiorax, length 20 to 2S percent of total
body length, antennal insertion on apical fourth;
evenlv, prominenth' arcuate from base to apex
in dorsal pioiile. In dorsal aspect prominently,
esenlv tapered from base to apex, frons 1.8-2.2
times wider between dorsal margin of eyes than
rostrum at extreme apex; distad of antennal in-
sertion oblong in cross section, not finely acu-
minate; smooth, shining, lateral pits shallow;
often slightly expanded at antennal insertion and
at extrem(> apex. Vestiture of elongate-oyal,
apically pointed, white, recumbent scales, and
long, narrow, strigose, suberect scales; no fine
erect setae, some round, nonstrigose scales on
lower portion of sides.

.\ntennal funicle seven-segmented; pedicel
as long or longer than next three segments com-
bined.

Pronotum wider than long, base about 1.7
times wider than apex at apical constiiction; sides
evenly, prominently rounded; punctures large,
evenly spaced, broadly visible on disc. Vestiture
complex, large, median, basal, patch of o\a], non-
strigose and long, narrow, strigose, white scales;
remainder of dorsum and upper portion of sides
with long, narrow, pointed, dark reddish brown,
strigose scales; lower portion of sides with oval
to elongate-oyal, white to light reddish brown,
nonstrigose scales, some extending to dorsum.

ICIytra nearly parallel in basal fointh, broadly
rounded to apices; in dorsal profile nearly flat
in basal half, declivity evenlv, broadly rounded.
\'estiture on interspaces of biseriate to triseriate
rows of oval, recumbent, white to light reddish
brown, nonstrigose, broadly imbricated scales;
scales on interspace one slightly denser, more

i)road]\ imbricated. Each interspace with medi-
an, uniseriate row of long, narrow, strigose,
suberect to erect, usually dark reddish brown,
apically pointed scales. Strial scales elongate,
pointed, while.

N'ciitral surface with oval, or elongate-oval, im-
bricated white, plumose margined, ncmstrigose
scales. Metasternum and abdominal sterna with
discrete, transverse rows of suberect to erect,
hairlike setae; sternum five with deep median
fovea.

I'\'min' long, narrow, apical ventral emargina-
tion well de\eloped; often metafemur with mi-
nute spine on proximal portion of emargination.
\estiture of elongate-oval, recumbent, nonstri-
gose scales, and long, narrow, suberect, strigose,
white to light reddish brown scales.

Tibiae mucronate, mucro on mesofemur and
metafemur usually shorter than tarsal claw; mu-
cro on protibia about ecjual in length to tarsal
claw. \'estiture of elongate-oval, nonstrigose and
long, narrow strigose scales, and fine, hairlike,
usually darker colored setae near apex.

llr^iS 'filial

Kig. 2. Dorsal view ol I'ljchiu.s phiilanis.

Biological Sehies, \'ol. 13. No. .3 Revision of the Weevil Genus Tychius

33

Tarsi dorsallv with long, narrow scales; claw
with short basal process, usuallv onlv half as
long as claw.

Male genitalia ( Fig. 7 ) with apical portion
of median lobe broadh' rounded; apical, dorsal,

sharply defined,

absent; median

rostrum usually
narrower, more

median membranous area not
posterior sclerotized margin
struts fine, clavate.

Female: length 2.S-3.6 mm;
equal in length to pronotum,
slender: tibial mucrones generally smaller.

Type Locality. ARIZONA: Organ Pipe Cac-
tus National Monument, Dripping Springs.

Type Material. Male holotvpe, female allo-
t\pe, 16 male and 15 female paratxpes taken at
the t\pe locality on April 5, 1969, by W. E.
Clark, sweeping the host plant. One female para-
t\pe was taken at the t\pe locality on April 24,
1953, b\' A. and H. Dietrich. Deposition of the
t\pe material is a follows: hoIot\'pe and allotype
(USNM), 1 male, 1 female paratype (BYU), one
female paratype (CAS); the remaining 29 para-
t\pes are retained in the authors collection.

Host. Lotus rigidus.

Distribution. (Fig. 20).

One specimen not included in the tvpe ma-
terial that was examined.

California: Po\va\, 1 female (CAS).
Total specimens examined: 35.

Fig. 3 Lateral view of Ti/c/iin.v aratiis. female.

Ti/cliitifi aratits Say
(Figs. 3, 4, 19)

Tychius aratus Say, 1831, Descriptions of North Ameri-
can curciilionides p. 26, (reprinted In:

LeConte, 1859, The complete writings of Thomas

Say 1:294) (Male holotvpe of Tifrliiim ani-

tor Gyllenhal here designated as neotjpe of Ttjchiu.s

Figs. 4-10. Ttjchius spp., median lobe of male genitalia;
4. T. arafu.i: 5, T. haditis; 6, T. prolixus; 7, T. pha-
larus; 8, T. hmellosus; 9, T. semisqiiamosus; 10,
T. stephensi; a - dorsal view, b - lateral view, c -
dorsal viev\' of apex. Line at right of Fig. 10 repre-
sents 1 mm.

aratus Say: Missouri, Naturhistoriska Riksmuseum,
Stockholm); LeConte, 1876, Proc. Amer. Philos. Soc,
15:432; Gemminger and Harold, 1871, Catalogus
Coleopterorum 8:2514 {=arator Gyllenhal).

Tychius arator Gvllenhal, 1836, In: Schoenherr, Genera
et species curculionidum. .... 3(1):414-415
(Holotype: male, Missouri .Naturhistoriska Riksmu-
seum, Stockholm).

Miccotrogus aratus Khma, 1934, Coleopterorum Cata-
logus, 29 (138): 29.

This species differs from other members of
the r. scmisquamosus species group by the uni-
colorous scales and larger size. The long ros-
trum in the female (Fig. 3) is unique among
known North American Tychius.

Description. Male: length 4.0-4.4 mm, width
1.8-2.1 mm; integument dark reddish brown to
black on bodw appendages dark reddish brown;
coyered b\' imicolorous, light, tawn\' scales.

Rostnmi longer than prothorax, slightly,
evenly arcuate or nearly straight in basal two-
thirds in dorsal profile; slightly wider at antennal
insertion in lateral aspect; in dorsal aspect slight-
ly, evenly tapered from base to tip, frons 1.4
times wider between dorsal margin of eyes than
rostrum at apex; antennal insertion in apical
third; evenly tapered to tip in lateral aspect,
lateral portion with very deep rugae, dorsal por-
tion with median shiny, smooth area. Vestiture
of uniform shape, size and color; distad of an-

34

Bhigham VouNt; Univehsiiv Science Bulletin

(t-niial insertion long, narrow, subcrcct setae ex-
tending t\\'o-tl)irds distance to apex, scales above
insertion stouter, no distinct erect hairlike .setae.

Antennal funicle seven-segmented; pedicel
as long as next two segments combined.

Pronotiim 1.0-1.2 times wider than long, sides
prominently arcuate, l.S-2.0 times wider at base
than at apical constriction. Scales of two types:
long, narrow, recumbent, apically pointed scales
on dorsum; sides with broad, oval, recumbent
scales, extending dorsally, intermingled with
elongate scales halfwav up sides, extending to
dorsum fonninii broad lateral vittae from dorsal
aspect.

Figs. 11-18. Ti/chiuii spp., median lohc of nuilc geni-
talia; U, r. soltmii; 12, T. tectus; 13, T. liljchhuli;
14, 7'. hirsutus; 15, T. ccicsiti.i; 16, T. lincclltis; 17,
7'. sordkltis; 18, 7'. rnontdnu.i; a - dorsal \i<-vv. b -
lateral view. Line at right ol Fi<;. 18 represents
1 mm.

Elytra 1.4-1.5 times longer tlian wide; in dor-
sal profile nearly flat in basal fointh to one-half,
broadly rounded to apices. Sides in dorsal a.spect
slightly rounded in basal two-thirds, broadlv
rounded to apices. Each interspace with thre(> or
four rows of broad, oval, sometimes pointed,
broadly imbricated, nonstrigose scales and with
medi;in, uniseriate to muitiseriate rows of long,
narrow, suberect, strigose sc-ales. Strial scales
n;irrou, pointed.

\enlral surface densely clothed with liroad
to elongate-oval densely imbricated scales and
transverse rows of erect setiform scales, these

often absent from visible abdominal stemiun one.
Slerntun five with broad, deep, median fovea.

Femur stout, apical, ventral emargination well
developed, metafcmur with minute tooth on
proximal portion ol emargination. Vestiture of
round, and long, n;u-row, suberect, strigose scales.
Tibi;ie with sliort, stout, iiuicroni^s, mucro
on protibia largest. \'estiturc of sparse round
scales and long, narrow, apically pointed, sub-
erect, strigose scales. Tarsi with line setae and
long, narrow, strigose scales dorsally, tarsal claw
divergent, basal processes parallel.

Male genitalia ( Fig. 4) with apical portion of
median lobe bearing lateral prominences, apex
roimded; apical, dorsal, median mem])ranous area
nearlv round, strongly defined posteriorly; me-
dian struts clavate.

Female: rostrum extiemelv long, narrow,
nearlv half bodv length. Antennal insertion near
middle, apical portion slightlv expanded, near-
1\- glabrous entire length. Tibi;d miicrones slight-
lv smaller than in male.

Host. Astratj,alti.s crassicarjni.s.

Distribution. (Fig. 19).

MiiinesDta: Duluth, Daggett, 1 male (LA).

Montana: Mason 1 female (USNM); Bozeman, VI-
4-.38, D. R. Lindsay, 1 male, (I.SU).

Wyoming: Johnson Co.. Bnffalo. 5 mi. W'., \l-2()-
68, W. E. Clark, .A.s7r(/^'r;/i/,v crassicurpii'i. 2 males
( WKC ) ,

Total specimens examined: 6.

Discussion. The "t\pe of T. ciratus Sav was
iipparentlv destrovcd (LeConte, 1859 :vi). A
specimen from the Gvllenhal collection in the
Stockholm Museum hdieled Tijrliius- arutus Say
was examined. Ciyllenhal (1836:414-415) states
that this specimen was sent to him by Say and
cites Tijchhis auitus S:i\ as a synon\mi of T.
(irator. LeConte ( 1876:216) says of T. orator that
"Sav :ipparentlv confounded this species with
one described bv him as T. arciius; and Major
Cyllenhal suspecting perhaps the existence of
some error has, while (jiioting Sav in svnonymy,
gi\('n ;i different n;ime to the insect received
Irom that author. 1 have examined the specimen
in the LeConl(> collection libeled T. arator and
determine it to be T. liljchlddi BIatchle\'. The
specimen in the Gvllenhal collection sent by Say
to Gvllenhal is prob;iblv the onlv authentic Say
specimen of T. arutus in existence; therefore, it
is here desigiuitcHl as the neot\pe.

The rehitionship ol T. aratus to the other
members of the scmiscjiiamosufi group is not
clear. It :i]')pears I'alhci' isolated in se\eral
leatm-es.

Biological Series, Vol. 13. No. 3 Revision of the Weevil Genus Tychius

35

NORTH AMER

0 500

MILES

Fif^. 19. Map of North America showing distributions of Tychius liljcbladiU, T. tectus 9, and T. arattisO-

36

Biucii AM VouNt; University Science Bulletin

Fig. 20. Map of tlie United States showing the distributions of Tijchius stephcnsi . T. sordklu.s
<'(i(.'.sin.v T , i ■ sultuui^, T. mnntiinusO, T. hirsutusm, and I", phalarus^.

ACKNOWLEDGMENTS

T wish to express my sincere appreciation
tor tlie help which many people havi- rendered
in the course of this study. Special acknowledg-
ment for assistance is due the following: Dr.
Stephen L. \V(K)d, Department of Zoology, Brig-
ham Young University, who served as chaimian
of the advisor)' committee and directed the
research and writing of the thesis. Dr. Vernon
Tipton, Department of Zoology, Rrighaiii Young
University, who took over as chairman of the
advisory commitee when Dr. Wood went on
sabbatical leave. Dr. Howard C. Stutz, Depart-
ment of Botany and Range Science, Brigiiam
Young University, wiio served as a member of
tile advisory committee. Dr. Vasco M. Tanner,
Detpartment of Zoology, Brigham Young Uni-
\ersity, who made specimens in the Brigham
Young University collection available for studv
and contributed advice and help at many times
during the course of the study, including dona-
tion of the services of his illustrator. Miss Janet

Tj. Tompkinson, who prepared Figures 1 and 2.
Dr. Stanlev E. Welsh, Department of Botany and
l-lange Science, Brigham Young University, for
identifying host plants of the weevils I collected.
Rose Ella Warner, Systematic Entomology Lab-
oratory, Entomology I^esearch Division, Agri-
cultural Research Service, USDA, for arranging
tlie loan of specimens from the United States
National Museum, making facilities available at
the museum for my study of the T. L. Casey
colltction, lielping with the literature survey,
and criticising the manuscript. Dr. John F.
Lawrence, Museinn of Comparative Zoology,
Harvard Uni\c-rsit\-, for arranging the loan of
specimens inc'liiding t\pe material and other
specimens from the LeConte collection. Dr. Ross
H. Arnett, Jr., Department of Entomology, I\ir-
due University, lor arranging tiie loan of speci-
mens from the Blatchley collection. Hugh B.
Ixxx-h, C/alifoniia .Academy ot Science, tor ar-
ranging the loan of specimens including the type

Biological Series, Vol. 13, No. 3 Revision of the Weevil Genus Tychius

37

Fig. 21 Map of uestem United States showing the ilistritnitions of Ti/chius lamcllofius Q , T, scmi.squn-
mnstisii, T. Imcellus^, T. badiuiA, and T pruUxiv a.

38

Brigham Young University Science Bulletin

of Ttjchius armatus Green. Per Inga Persson, De-
partment oF Entomology. Swedish Museum of
Natural History, Stockholm, for sending the types
of Ti/c/iiiM a'rator Gyllcnhal and 7'. posticus
Cvllenhal, specimens from the Paykull collection
and for infonnation regarding type material of
T. stephensi Schoenherr, and Curculio fusciro-
stris Pavkull. Dr. F. Heike, Zoologisches Mu-
seum, Berlin, for sending the type series of C.
tomentoms Herbst. Dr. Lars Medstrom, Univer-
sity of Uppsala, Uppsala, Sweden, for sending the
tv'pe material of Rhtpuhaenus picirostris Gyllen-
lial. Dr. H. T. Thompson. British Museum (Nat-
ural History), for sending specimens from the
Stephens c-ollcction and for infonnation on the
tvpes of C. cinerascens Marsham, C. villostts
Marsham, and C. fiicirostris Fabricius. and for
infonnation pertaining to the nomenclature of
Ti/chitis stcjilicnsi Schoenherr.

The following individuals and instritutions

also lent specimens for studv; D. E. Bright, Jr.,
Entomoiogv Research Institute, Canada Depart-
ment of Agriculture; H. 1^. Burke, Texas A. & M.
University; G. W. Byers, University of Kansas;
(. A. Chemsak, University of California, Berke-
ley; M. G. Emsley, Academv of Natural Sciences
of Philadelphia; W. J. Hanson, Utah State Uni-
versity; L. H. Hennan, Jr., American Museum
of Natural History; J. Laffoon, Iowa State Uni-
versity; C. W. OBrien, Texas Tech University;
P. Oman, Oregon State University; L. L. Pechu-
inan, Cornell University; C. A. Triplehom, Ohio
State University; F . S. Truxal, Los Angeles
County Museum; J. D. Unzicker, Illinois Natural
History Survey; R. L. Wenzel, Field Mu.seum of
Natural History and F. G. Werner, University
of Arizon;i.

I am especially grateful to my wife, CJlenora
|ean, who typed all drafts of the manuscript and
helped prepare the species distribution sections.

LITERATURE CITED

.Xndkhson, D. M. 1962 The weevil genus Smicronyx

in .America north of Me.\ico (Coleoptera: Curcu-

Honid.ie). Proc. U.S. Nat. Mus., 11.3; 18.5-.372.
BL.-VTCHLEY, \V. S. 1916. hi: Bhitchley, \V. S., and C.

W. Leng, Rhynchophora or weevils of northeastern

.■\mcrica. .\aturc Pub. Co., Indianapolis, Indiana.

682 p.
Casey, T. L. 1892. Colcopterological notices. I\'. Ann.

New York Acad. Sci., 6:359-712.
. 1897. Colcoptcrological notices. VII. Ann.

New York Acad. Sci., 9:285-684.
. 1910. On some new species of Balanini,

Tychiini and related tribes. Can. Entomol.. 42:114-

144.
Fabricius, J. C. 1787. Mantissa Insectornm. vol. I.

Hafniae, 348 p.
Frost, S. VV. 1945. Insects feeding and lirccding on

indigo, Baptisia. J. New York Entomol. Soc. .53:219-

225.
Cermar, E. F. 1817. Mlscellen and Corresponden-

Nachrichten. Mag. Entomol. 2:,3.39-341.
. 1824. Insectorum species novae . . . ., Halae,

624 p.
Green, J. W. 1920. Notes on American Wivnclioplior.i

(Col.). Entomol. News 31:19.3-201.
Gyllenhal, L. 1813. Insecta Suecica, vol. 3.
. 1836. In: Schoenherr, C. J. Genera et .species

curciilionidum Vol. 3. Paris; 858 p.

HuLUEBT, 1,. C.,ANi) F. W. Oeh.me. 1961. Plants.

poisonous to livestock. Kansas State Univ., Man-
hattan.
Kissinger, D. G. 1964. Curculionidae of America

north of Mexico a key to the genera. Taxonomic

Publications, South Lanc;ister, Mass., 143 p.
Klima, a. 1934. Coleopterorum Cataloeus, Tvchiinae.

29(138). 61 p.
LeConte, J. L. 1859. The complete writings of

Thomas Say on the entomology of North .America.

Bailliere Bros., New York. Vol. I, xxiv -f 412 p.;
Vol. II, iv + 814 p.

. 1876. In: J. L. LeConte and G. H. Horn

The Rh)nichophora of America North of Mexico.
Proc. Amer. Phil. Soc, 15:wi-xvi -f 455 p.

Leng, C. W. 1920. Catalogue of the Coleoptera of
America north of Mexico. New York, 470 p.

Mahco\itch, S. 1916. Insects attacking weeds in
Minnesota. Rep. Minn. State Entomol., 16:1.35-152.

MiLLiRON, H. E. 1949. The identity of two intro-
duced clover seed weevils (Coleoptera: Curcu-
lionidae). J. New York Entomol. Sc. 57:109-116.

Mitchell, J. D., ,\no \V. D. Pierce. 1911. The wee-
vils of Victoria Count\ , Texas. Proc. Entomol. Soc.
Wa,sh., 13:4.5-62.

McKA, A. A. 19.55. The biolog\- of the clo\er head
weevil, Ttjchiiis stephensi Schoenherr, with some
notes on control. Siimmani In: Dissert. Abstr. 15
(1):12-13.

P.\ykull, 1792. Monographi.i cnrculioniim Suecica,
427 p.

. 180(1. Fauna Suecica, Insecta. part 3.

Pierce, W. D. 1907. On the biologies of the Rhyn-
chophora of North America. Studies Zool. Lab.
Univ. Nebraska, p. 249-319.

. 1907b. Contributions to the knowledge of

Rhynchophora. Entomol. News, 13:356-363.

. 1912. The insect enemies of the cotton boll

weevil. USDA Bur. Entomol. Bull., 100:1-99.

Sanderson, E. D. 1904. Insects mist;iken for the
mexican cotton Ixill wee\il. Tex.as Agr. E.xp. Sta.
Bull.. 74:3-13.

Say, T. 1831. Description of new species of Curcu-
lionides of North .America. New Harmonv, Indiana,
30 p. Reprinted In: LeConte. J. L. 1859.' The com-
plete writings of Thomas Sav on the entomology of
North America. Bailliere Bros., New York, \'ol. I,
.xxiv + 413 p.; Vol. II, iv -|- 814 p.

Bidi.ooicAL Series, Vol. 13, No. ,3 Revision of the Weevil Genus Tvchius

39

.Sc:ii.\effeh, C. F. A. 1908. New Rln iichophora. 111.

J. New York Enfomol. Soc, 16:2i3-222.
. 1915. Mi.scelhmeous notes. J. New York Eii-

tomol. Soc, 23:197.
.Schoenhehh, C. J. 1825. Tabula synoptica faniiliae

curculionidiim. Isis von Oken. 5:581-588.
. 1826. Curculionitlum dispositio methodica

cum genera characteribus Li]5siae, 338 p.

. 1836. Genera et species Curculionidiim. . . . ,

\\)1. 3, 858 p.
. 1843. Genera et species Curculionidiim

\'ol. 7, pt. 2, 461 p.
SiiMir, D.. ano F. Mum, 1912. The c(imp:irati\e

aiiiitomy of tlie male genital tube in Colcoptera.

Trans. Entomol. Soc. Loud., 3:477-642.
Stei'iie.vs. }. F. 1829. A systematic catalogue of

British insects.
388 p

London: B:ddwin and Cradock,

1839. A manual of British Coleoptera or
beetles. London: Longman. Oniie Brown, Green,
and Longmans, 443 p.

Takenouchi, Y. 1965. Chromosome survey in thirty-
four species of bise.\ual and parthenogenetic wee-
vils of Canada. Can, J. Genet, and Cytol., 8:663-
687.

Tanneh, v. M. 1966. Rhvnchophora beetles of the
Nev:ida test site. Brigham 'i'oiing Univ. Sci Bull.,
Biol. Series, 8(2): 1-35.

ToHMK-BuENO, J. R. 1962. A glossary of entomology
and supplement A, George S. Tullouch ed. Third
Printing. BrookUn Entomol. Soc, BrookKn. N. Y.,
336 -t-'^ 36 p., 9 pi.

S-WA

'?.

C(-

Brigham Young University
Science Bulletin

A REVISION OF CRYPTANTHA
SUBGENUS OREOCARYA

by

Larry C. Higgins

BIOLOGICAL SERIES — VOLUME XIII, NUMBER 4
MARCH 1971

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany,

Brigham Young University, Provo, Utah

Members of the Editorial Board:

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

Ex officio Members:

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

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

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

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

^^

li^.

■ \ ■

^7 I- ii: s ..^

^7 I- |>^ / ,.^

■ r

I ronlispiccc. Ciypiaiillia flava Nutl. Drawn Iroiii Wi-lsh and Alwooil 9833. (2/3 aflual si/c).

Brigham Young University
Science Bulletin

A REVISION OF CRYPTANTHA
SUBGENUS OREOCARYA

by

Larry C. Higgins

BIOLOGICAL SERIES — VOLUME XIII, NUMBER 4
MARCH 1971

TABLE OF CONTENTS

INTRODUCTION 1

Materials and Methods

History of the Subgenus Oreocarya

General Morphology

Distribution and Ecology

Generic Relations and Phylogeny

Cytology

TAXONOMY 10

ACKNOWLEDGMENTS 59

APPENDIX 60

UST OF REFERENCES 62

A REVISION OF CRYPTANTHA SUBGENUS OREOCARYA

by
Larry C. Higgins'

ABSTRACT

Tills study is a taxonomic revision of Crypiuntliu
subgenus Oreocarya (Boraginaceae). togetiier witii
descriptions of species, synonyms, distribution maps,
and discussions. The present treatment recognizes
llt'ty-seven species and fourteen varieties, all separated
by means of a comprehensive key, A history of the
group, along with a discussion of distribution, ecol-
ogy, and phylogeny is presented. As a basis for the

comparison of taxa much of the research has been
concentrated on morphological variation in her-
barium specimens. Particular attention has been
directed to a study of pubescence types, branching
patterns, llowers, calyces, and fruit. The herbarium
research has been correlated with extensive field
observations and collections made throughout most
of western North America.

INTRODUCTION

The genus Cryptantha subgenus Oreocarya consists
o'i perennial or biennial herbs centered in western
North America and belongs to the tribe Eritrichieae
of the subfamily Boraginoideae of the family Bora-
ginaceae. The subgenus has long been recognized for
its taxonomic difficulty. This is due to the variability
o'i the species and the lack of distinctive vegetative
characters. Experience has shown that characters of
the fruiting and floral structure provide the only satis-
factory basis for precise specific differentation. The
complexit>- of this subgenus plus the large number of

species described since Payson's monograph (1927),
have made a thorough study of the group necessary,
it is hoped that the present revision of the North
American species of this subgenus will partially meet
this need.

The author first became interested in Oreocarya
while doing fieldwork preliminary to the preparation
of a thesis on the flora of the Beaverdam Mountains.
Because of the difTiculty encountered in classifying
the plants of this group and at the suggestion of Dr.
Stanley L. Welsh, the present study was undertaken.

Materials and Methods

The materials used in this study are largely dried
and pressed specimens from a nimiber of institutional
herbaria in the western hemisphere, together with my
collections from sites within the western United
States. Because of the large number of specimens
examined during the course of this investigation, cita-
tion of all specimens has been omitted, and only a
few representative ones for each taxon are included.
Type specimens of most taxa were received on loan
and were subsequently photographed and all photo-
graphs were deposited in the Brigham Young Univer-
sity Herbariiun.

Collectiiins were made during the spring and
summer months of 1967 in Utah and northern Ari-
zona. Additional collections were made in Wyoming.
Colorado. Montana. Washington, Oregon, Idaho,

Nevada, California, Texas, New Mexico, Mexico, Ari-
zona, and Utah during the spring and summer months
of 1968, U)6^),and 1970.

The measurements of a 15 cm ruler were used to
measure leaves, stems, and inflorescences. Plant
heiglit was measured from the base of the stem to the
tip of the inflorescence. Stem length was measured
from the base of the stem to the first flower. The
measurements of calyces, flowers, and nutlets were
facilitated by the use of an ocular micrometer fitted
to a bmocular microscope.

The taxonomic presentation in this revision fol-
lows a conventional pattern. The number in Arabic
numerals following the description of each species or
variety denotes the total number of collections
studied in preparation of the text. The figure in small
Roman numerals immediately following in paren-
theses denotes the number of these collections made

Department of Biology, West Texas State University. Canyon. Texas 79015

ijkk;ha\i younc; univlkshi' sen nci. bi'lliiin

by nio.

Herbaria t'rom which specimens have been seen.
with the standard abbreviations by wiiicii they are
referred to in the text (Lanjow and Stanicu, l')64)
are as follows:

ARIZ University of Arizona Herbarium,
Tuscon, Arizona.

ASC Arizona State College, Flagstaff,
Arizona.

BRY Brigliani Young University, Provo,
Utah.

CAS California Academy of Sciences,
San Francisco, California.

COLO University of Colorado, Bouldci.
Colorado.

CS Colorado State University. Fort

Collins, Colorado.

1)1 \ Dixie Junior College, St. George,
Utah.

(jll Gray Herbarium. Harvard Univer-

sity, Cambridge, Mass.

IDS Idaho State College Herbarium.
Pocatello, Idaho.

LA University of Califnnna. Los An-

geles. California.

LL Lundell Herbarium, Reniier. Texas,

MNA Museum of Northern Arizona. Flag-
staff. Arizona.

MONT Montana State LIniversity. Boze-
man. Montana.

ND-O Greene Herbarium, University o{'
Notre Dame, Indiana.

ND University of Notre Dame, Notre

Dame, Indiana.

NY New York Botanical Garden. New

York, New York.

ORF University of Oregon Herbarium.
Fugene, Oregon.

I'll Philadelphia Academy of Naliiral

Sciences, Plilla., Penii.

POM Pomona College Heibarium. Clare-
moni, California.

RM Rocky Mountain Herbarium, Lara-

mie. Wyoming.

RSA Rancho Santa .Ana Botanic Garden.
Claremont, California.

TTC Texas Tech. University, Lubbock.
Texas.

UC University of Calilornia, Berkeley.

California.

US United States National Museum,

Washington, D.C.

UT University of Utah Herbarium, Salt

Lake City, Utah.

UTC Intermountain Herbarium, Logan.
Utah.

WTU University of Washington Herbar-
ium, Seattle, Washington.

WTSU West Texas State University, Can-
yon, Texas.

History of the Subgenus Oreocarya

The species belonging to this group to be named
first was Cynogldssinii gloDicranim Nutt. in Fras.,
nom. nud. (Crypianiha cclosiaides). the type of the
subgenus. This was collected by Bradbury in "Upper
Louisiaiui" in 1810, and was described by Pursh
(1814). at which time he cited the Fraser Brothers
Catalogue as the source of his name. The citation
should thus read: C. glonunituni Null, ex Pursh. Fl.
Am. Sept. 2:729. 1814. It was transferred to A/v(;so;/.v
by Nuttall (1818). Torrey ( 1828). described a second
species, Myosolis siiflnaicosa [C. jamesii). A third
species was described (M. kucophaea). and also as-
signed to Myosolis by Douglas in Lehmann (1830).

With the appearance of the tenth volume of de
Candolle's Frodroinus (1846), C. gloincralum and A/.
leiicophuca were placed in the genus Ehtrivhiuiii and
this treatment stood until 1885.

.Asa Gray (1875). published the results of his llrsl
study of the eritrichioid borages of western America.
Following de Candolle he referred all the species to
tirithchium section Pseudo-myosotis. Four species
and three varieties were recognized in Gray's revision.

Bentham and Hooker (1876). published that part
of their Gciwra Plautamm treating the Boraginaceae.
These authors accepted the interpretations of de
Candolle and Gray and added still mure diverse ele-
ments to the already overburdened genus h'riiricliiiim.
The species of Ciyptaniha were placed under that
genus and considered generically indistinguishable
from plants now classified under /'liiglohollirvs and
Ij-iiricliiiiiii.

The genus Ehtrkliiuiu became so heterogenous
and varied that its breakup was inevitable. This began
with Gray's notable paper (1885). .4 Revision of
some BoragincoHs Genera, iii which the species of
Crypianiha were transferred to the section Pseudo-
krynit/kia ol Krviiilzkia with the exception of K
selosissiina which was placed in the section Ptery-
giunr. Fight species were recognized by Gray that
now are placed in the subgenus Oreocarya.

In a series of papers published by Greene ( 1887),
the American representatives of de Candolle's h'rilh-
eliiiiin were segregated in detail. First, the genus /l/fo-
earya was formed to include the species Gray had
treated as Kryin'lzkia section Myosr)tidea. Then Pipro-
ealyx was reinstated to include the species with cir-
cumscissile calyces; also two new genera. Eremocarya
and Oreocarya. were lormetl. The first, Eremocarya.
was based upon the plain of southwestern United
Stales described as E. niieranihiini by Torrey ( 1859).
and the second. Oreocarya. upon nine species of /:>;'-
irieliiiiiii section I'seiido-kr\ nitzkia and pari ot Kry-
iiilzkia section Pterygiimi. desciibed by (ira\ (1885).

BIOLOGICAL SLRIKS VOL I,!, NO. 4 A RKVLSION Ol CRYITANTHA Sll BGENUS ORKOCAR Y A

Greene (iS'^'O). described eight new species and rede-
scribed several iild ones. Later Greene (18'^')). de-
scribed two new species and still later two intire
(1^)01).

In the period from IS^-'d to U'lb some 35 species
were proposed as new, mainly by Marcus Jones
(1891, 1895, 1910), Alice Eastwood (1903. 1913).
and Per Axel Rydberg (1905, 1906, 1909, 1913).

Macbride (|9I6). wrote the first comprehensive
revision ol' the subgenus Oreocarya which included 45
species. A dichotomous key was provided, specimens
were cited, and a short discussion of each species was
included.

Ivan M. Johnston ( I'^I24). questioned the status of
the genus Oreocarya. and was ot' the opniion that
both Oreocarya and Cryptaiitha slundd be combined
under Cryptantha. Payson (1927). transferred all
species previously known within Oreocarya to
Cryptantha. His treatment included 45 species and
contained keys to the species and a list of synonymy
tor each species plus descriptions, specimen citations,
and discussions.

Brand ( 1927) proposed the new genus Hemisphae-
rocarya. which included all those species of the sub-
genus Oreocarya in which the fruit is hemispherical.
Cryptantha jamesii and four of its close relatives
formed the basis for this genus.

Since 1927 most of the new species which have
appeared in the literature have been proposed by
Johnston (1932. 1937. 193'^), 1940), and Higgins
( 1968. 1969).

General Morphology

The plant consists of a taproot, varying in degree
of branching, and a simple or branched caude.x. the
apex of which bears a rosette of leaves and gives rise
to the flowering stem. The stem pattern may differ
considerably between species, but fundamentally it
consists of an axis, usually foliate (or bracteate), with
branches bearing several helicoid cymes.

The majority of species live for several years and
are classified as perennials. In C. setusissima and C.
rirgata. however, the plants are biennial with one
prominent rosette giving rise to a central flowering
stem.
Stems:

The steins are herbaceous, usually arising from a
much branched woody caudex. However, in some
species such as C. rirgata and C. setosissima the stems
are simple and erect. Most species are very similar
with respect to the character of the stem, and so
stems can be used only to a limited extent in the dif
ferentation of species.
Leaves:

The leaves are very similar in all species, varying
only in length and width. They range from linear in
C. jamesii var. laxa and C. shackletleana. to spatulate.

or as is most common, oblanceolate. In all species the
margins are entire, and the apices acute, obtuse, or
subretuse. The blade is most often gradually nar-
rowed into a long, slender, ciliate-margined petiole.
Inflorescence:

The inflorescence is an open, rounded leafy-
bracteate thyrsus or a helicoid cyme. The individual
cymules are prevailingly scorpioid, but sometiiries
glomerate or loosely racemose. In some species the
inflorescence is characteristically narrow, while in
others as C fhyrsiflora. it is very broad and open. In
one species. C. rirgata. the floral bracts are extremely
long and greatly exceed the individual glomerate
cymules.
Sepals:

The calyx is five-parted to the base, with the lobes
all equal and linear, lanceolate, or ovate. In the more
primitive species of this subgenus the lobes are only
slightly accrescent in fruit, and in the advanced spe-
cies greatly enlarged. The pedicel of the mature calyx
may be nearly sessile or very short, or up to 10 mm
long in the case of C. fulrocanescens var. echinoides.
Trichomes:

The hairs of Cryptantha are all simple, unicellular,
and more or less silicious. The more silicious hairs are
either smooth and somewhat transparent, or more or
less roughened by encrustations and then somewhat
opaque. The bristles vary notably in attitude (whe-
ther appressed, retrorse, or ascending), length, and
rigidity. Some species are characteristically setose or
hispid and others conspicuously silky-strigose. but
most of the different forms are so much alike in
pubescence that the differences are unnoticed by one
unfamiliar with the group.

Associated with the bristlelike trichomes on the
stem and leaves are the pale, blistery structures called
pustules. These are composed of a circle of slightly
elevated silicified, opaque, tessellately arranged epi-
dermal cells surrounding the base of the trichome.
They show much variety in frequency and size, vary-
ing from totally absent to decidedly abundant, and
up to a diameter of 4 mm.

The terms used in botanical literature to describe
the many diverse types of pubescence are employed
by varitjus authors in different contexts. The follow-
ing list of terms is presented to allow precision in
interpretation of pubescence types.

Hirsute: with long, moderately

stiff hairs.
Hispid: with long, very stiff

hairs.
Setose: with short, rather stiff

hairs.
Strigose: with short, appressed

hairs usually in one

direction.
Tomentose: hairs medium to short,

curled and interwoven.

BKIGHAM YOUNC; UNIVI.KSII V SCll.NCh. UULLEIIN

Ci)rolla:

The corolla linib is nearly always while and roiate-
salverrorm or campanuiate, but is distinctly yellow in
two species. However, in many of the white-flowered
plants the tube is light-yellow. The relative length of
tiie corolla tube to the calyx lobes is an important
diagnostic character. The corollas vary from 2-20 mm
in length, but are quite constant for any given species.
The tube bears at its ape.x five rounded or emarginate
fornices, 0.5-1.5 mm long, that are conspicuously
papillose to nearly glabrous. Near the base of the tube
a ring of crests is usually present, but may be lacking.
especially in the long-tlowered species.
Style:

The length of the style as well as the position of
the stamens in the corolla tube is of little specific
value in most of the long-tlowered species, since most
of them are distinctly dimorphic in these characters.
However, in C. oblata. a long-nowered species, the
stamens are always located at the middle of the tube,
and the style exceeds the mature fruit by 3 to 5 mm.
In the short-flowered species the distance by which
the style exceeds the fruit is of some value in distin-
guishing species.
Fruit and nutlets:

The fruit of Crypiuitlhu consists of four or fewer
elongate, ventrally grt)ovcd nutlets affixed to a us-
ually elongate gynobase. In shape the nutlets vary
t^om broadly ovate or triangular-ovate to narrowly
lanceolate. The dorsal surface occasionally is some-
what flattened, but is commonly more or less convex,
although in C. virgincnsis it is somewhat obtuse with
a meilial dorsal ridge. The sides of the nutlets in most
species are acute, but can be quite obtusely angled, oi
as in C. selosissima the margin is drawn out into a
broad, greatly developed wing. Surface characteristics
of the nutlets are also of diagnostic value. The surface
of the nutlet may be smooth and shiny as in C. con-
fertijlora. C. Jlava, C. bamehyi, C. semiglahra, or may
be variously roughened. In species such as C brcvi-
flora. C. caiui. and C. fiilvocanescens the roughenings
consist of small, numerous, low, rounded, or sharp
projections, a condition here described as muricate.
Other species, such as C. stricta. C. Jhvoaihiici. and C.
celosioidcs have the surface covered with long or
short, irregular or transverse ridges called rugae.

Although four nutlets arc commonly produced.
one or more of them may fail to develop. This abor-
tion can prevail throughout a plan! or be restricted
largely to either its younger or older parts. This sug-
gests that abortions might be partially connected to
the nutrition of the plant as intluenced by seasonal
conditions.

The nutlets are all homomorphic in contrast to
those in the annual species of the subgenus Krynit/-
kia. which contains many species possessing hetero-
morphic nutlets. The ventral groove or scar of the
nutlet can be open or closed and can he simple or

forked below. The margin surrounding the scar is
entirely lacking or is greatly elevated as in C. Jlavo-
culalii and ('. hakcri.

L^istnbutinn and l-.cology

Oreocarya, as circumscribed in tlie present tieat-
ment. is principally western North American in distri-
bution. Only one taxon (C gnaphalioiik's). or pos-
sibly two others, namely C. argciiica and C. amplexi-
aiiilis. are found in South America (Chile), while all
others occur in western North America. Payson
(1927) stated.

It wt)uld seem reasonable to suppose that
the genus originated in the southwestern
part of North America and in its perennial
primitive form spread lo the desert regions
of Chile and Argentina. In that region it be-
came differentiated into three main groups.
The most primitive of these invaded North
America as Cryptantha in the limited sense
of recent American floras.

With this opinion the present author cannot en-
tirely agree. A more reasonable hypothesis is that the
group had its origin in southwestern North America.
In its primitive form (probably similar to C. jainesii
var. imdlicuulis). it spread to the desert regions of
Chile and Argentina. In that region it became ditTer-
entiaied into Geocarya. an advanced group with
highly specialized, cleistogamous flowers. The niaior-
ity of the group that remained in western North
America became specialized and gave rise to the sub-
genus Krynitzkia. The subgenus Krynitzkia in turn
spread to South America and the specialized subgenus
('ry|itantha developed from it.

The center of distribution in North America is
eastern Utah and western Colorado, with the greatest
concentration of species extending from south-
western Wyoming and northwestern Colorado to
snutheaslein California.

Members of the subgenus Oreocarya occupy lor
the most part xerophytic habitats at middle eleva-
tions. The species of Oreocarya grow in widely dif-
leient ecological situations and in a wide elevational
range. Few species are able to tolerate direct compe-
tition with other herbs or endure the shade of over-
hanging trees or shrubs. The prairie species C. cana
and C. cdcspilosu. appear unable lo tolerate the .sod-
grass climax vegetation, but flourish only on the shaly
ridges or gravelly outcrops which here and iheie
break through the prairie sod. Some species are lound
growing on sandy deserts, e.g. C. jamesii var. disticha,
on alpine talus slopes, e.g. C. wchcri, C. cryniopliila,
C. fhonipsonll, and some on clay knolls or desert
playa. e.g. C. virginensis, C. ntgiilosa. and C. semi-
glahni which are associated with .1//7/)/<'.y. Apparently

HIOLOGIC'AL SERIKS VOL I 3, NO. 4 A RKVISION Ol IK> I'l AN I IIA SUBGKNUS OKl-OCAR Y A

no species is able to tolerate a moist, uiidrained soil.
I he ability to grow and reproduce in difficLilt envi-
loniiieiits has enabled members of tiie group to col-
onize a wide variety of raw and newly exposed, often
uiipriimising habitats. It is perhaps this ability, toge-
ther with the selective action of the environment
upon pioneer populations, that has produced the
large number of species and at the same time the pat-
terns of narrow endemism which are a feature of the
subgenus.

The species are commonly found on almost any
barren hillside. Many are restricted to soils that are so
strongly impregnated with mineral salts that few
other plants are able to compete with them. They are
particularly at home on loose hillsides, talus slopes,
shale outcrops, and heavy, detrital, clay soils.

Over 50 percent of the species are narrowly re-
stricted in some specific ecological niche or edaphic
situation. Some of them are limited to a single hill-
side, others to a particular mountain range, and still
others are restricted to some isloated basin.

The greatest number of endemics are found in the
cold desert regions of Utah and Colorado. C. grahamii
and C. hamchyi are restricted to the Green River
Shales of the lower Uintah Basin, being the most con-
spicuous plants on the shales. C. stricta is a very nar-
row endemic, also of the Uintah Basin, but it is lim-
ited to heavy clay habitats on the north rim of the
basin. Other endemics such as C. data. C. aperta. C.
brcviflora. C. wethcnllli, C. loiigiflora. and C. nigii-
losa grow on higlily saline soils. Gypsiferous soils of
western Colorado and eastern Utah are the home of
C. paradoxa. while in this same general region C.
Wniiia and C. osrerlunitii are almost always confined
to sandy soil.

Endemic species also occur in mesic environments.
For example, the Wasatch Formation of western
Garfield County. Utah, is the habitat of the narrowly
endemic C. ochrolcuca. The volcanic talus slopes
along the Salmon River in central Idaho is the habitat
of C. salmonensis. Montane and warm desert regions
include the habitats of other endemics.

Dispersal mechanisms within the subgenus are
almost entirely unknown. One species, C. setosissiina.
has broadly winged nutlets, but the size of the wing
in relation to nutlet size is such that it may have little
or no value in dispersal. It seems probable that long
distance dispersal by means of some animal vector is
the most reasonable answer to dispersal of propagules
in this group. However, the species of Cryptantha are
not known to be grazed or eaten by animals, and thus
the potential of animals in their dispersal is unknown.
It is apparent that much additional information must
be obtained to account for the broad distribution of
members of this group of plants.

Seasonal differences in the Howering period are
highly variable in the subgenus and have probably
served in the isolation of species within the group.

For example. C. joncsiana is closely related to C. fiil-
rocanesccns but fiowers much earlier. By early May
C. juncsiana has completed Howering and produced
fruit, while C. fiilvocancscens has only started to
tlower. Seasonal isolation between other closely re-
lated species is known (e.g. C. virglnensls and C. hoff-
niannli. C. spiculifera and C. intcrnipta, and between
C. Iiumilis and C. propria).

Heterostyly, (regularly associated with outbreed-
ing populations), has been developed in several groups
in the evolution of the subgenus. Some of the widely
distributed species such as C. flavoculata and C. fiilvo-
cancscens are distinctly heterostyled. On the other
hand some of the very narrow endemics such as C.
oclirolciica. C. compacta. and C. joncsiana are hoiiu)-
styled and possibly self-pollinated. Outbreeding in the
subgenus is probably also facilitated by various in-
sects. Butterfiies and beetles have been noted visiting
several of the species and undoubtedly serve as pollin-
ating agents.

Generic Relations and Phylogeny

The genus Cryptantha is a member of the tribe
Eritrichieae and is apparently derived from the Litho-
spermeae through some form similar to the North
American species of .Antiphytum. Johnston (1925).

The primitive fruit was probably similar to that of
the Hydrtiphyllaceae, being two or imperfectly four-
loculed. capsular, and terminated by a lobed style.
The lobed style consistently occurs in those subfam-
ilies of tlie Boraginaceae which are considered primi-
tive, i.e. the Heliotropioideae, Ehretioideae, and
Cordioideae. Within the subfamily Boraginoideae, the
style is lobed or bears geminate stigmas only in the
tribe Lithospermeae.

The nutlets have resulted from a pinching in of the
pericarp walls to form lobes of the fruit each contain-
ing one ovule. The stages of this development may be
appreciated by a comparative study of the fruit of
Heliotropium or better still, of ColJcnia. Coldenia
cancscens has an unlobed fruit bearing a decidedly
terminal style: Coldenia nutiallii has the lobing evi-
dent and the style attached to the pericarp between
and below the apices of the nutlets. In Coldenia litor-
alis the lobing is almost complete and the style is
affixed almost upon the receptacle. Evidently the
development of nutlets has proceeded by the deepen-
ing downward of the lobing from the apex and in-
ward from the sides by the pinching in of the pericarp
between the carpels. This finally results in an appar-
ent basal attachment of the nutlets and a gradual low-
ering of the style base between the nutlets until it is
at last directly and firmly affixed upon the recep-
tacle. Johnston (1925).

The nutlets of the annual as well as the perennial
species of Cryptantha are characterized by the pres-
ence of a medial ventral groove, which is caused by

HKICIIAM YOUNG UNIVl-.RSH Y SClENCt BULLETIN

the nont'usion of tlie pericarp walls. This development
appears to have been brought about by the gradual
encroachment of the pericarp over the surface of tlic
sliarply cut triangular attachment scar such as those
in the Lithospermeae. This encroachment gradually
narrows the attachment surface of the nutlet and
forms a groove which is usually somewhat forked at
the base. In Cryptantha the groove may be narrow
but not completely closed. In Anisinckla. Plagio-
bothrys. etc. the groove is entirely closed and its loca-
tion is marked by a ridge of fused pericarpel tissue
which bears the scar, Johnston (1925).

According to Johnston (1925). it seems quite
probable that the subgenus Oreocarya has been de-
rived from some form t)f Aniiphytum. a genus of
Lithospermeae evidently derived from some ancestral
plant similar to Lithospennum. The subgenus Oreo-
carya is the most primitive in the genus, probably
having given rise to the other subgenera of Krynit/-
kia, Cryptantha, and Geocarya, of which the latter
two possess cleistogamous (lowers. Oreocarya seems
also to have given rise to Flagiohothrys. a genus which
appears then to have evolved Amsinckia. The prin-
cipal derivative of Oreocarya, however, appears to be
llcnkclia. for Oreocarya appears to be connected with
Cynoglossum through llackclia.

According to Johnston ( 1925) and Payson ( 1 927).
the subgenus Krynitzkia is a specialized ofl'sliuot of
Oreocarya. Oreocarya consists of rather coarse peren-
nials or sometimes biennials with usually persistcni
calyces and hoinomorphic nutlets. However, ni ihe
subgenus Krynii/kia, composed mostly of annuals,
the calyces are deciduous and the nutlets somewhat
heteromorphic. In the present group the question of
primitive or of advanced condition is of much inter-
est. In the present work the species have been
grouped together mainly on the basis of similar mor-
phological characters. After reviewing the works by
Johnston (1924, 1925) and Payson (1927), plus this
current research a phylogenetic summary is presented
below. These conclusions aie not final and may be
subject to change after atldilmiuil leseaich.
Nutlets:

in the hypothetical, primitive plant the luillcis
evidently were smooth on both surtaces and ovale in
outline. The scar of the nutlet was straight and nar-
row without an elevated margin. Nutlets also were
attached at a very low position on the gynobase. As
specialization tt)ok place the nutlets were roughened
first on the dorsal and then on the ventral surface.
The nutlets became more lanceolate, with the scar
developing an elevated margin and becoming open
and contorted. The nutlet of the supposedly advanced
plant also became attached at a higher point on the
gynobasic style.
Corolla;

The primitive plant had corollas which were shoit
and never exceeded the lobes of the calyx. This prim-

itive corolla had low, rounded fornices and crests at
the base of the tube. As specialization took place the
corolla became longer until it greatly exceeded the
lobes of the calyx. This long tube apparently devel-
oped along several different lines in the evolution of
the group. Styles of various lengths (heterostyly), are
regularly associated with long tubes. This is also an
advanced characteristic, styles of uniform length
being primitive.
Calyx segments:

Broadly ovate or lanceolate sepals that were
slightly accrescent were primitive, while those which
are narrowly lanceolate or linear and conspicuously
accrescent in iVuit are advanced.
Intlorescence;

The primitive inflorescence was confined to the
upper part o\' the stem and was composed of a few,
conspicuously elongated, scorpioid cymules. As devel-
opment proceeded the cymules became shorter and
the inflorescence tended to cover more and more of
the stem.
.Stem:

The question of the primitive nature of the peren-
nial or annual habit is of much interest. Morpholo-
gists. in tieneral, seem to be ot' the opinion that the
perennial habit is more primitive than the annual
hahil oi biennial habit. The autln)r is inclined to agree
with this generalization. The primitive species in this
group were perennial with slender unbranched stems,
while the less enduring ones with very short branched
steins are specialized.
Leaves and pubescence:

The linear oblanceolate leal' is more piiniili\e lliaii
the broader ovale oi spalulate i\pes. An exlremcK
setose indumenl on the leaf is considered to be a
mark ol speciahzalion. and in like manner a conspic-
uously sericeous and uniform covering is a specializa-
tion in another direction.
Phylogenetic relationships:

The North American species of ihe subgenus Oieo-
car\a are thought lo he inonophylelic. but showing
several maior lines ot development. In the present
treatment the species groups are not given any taxon-
oiiik i,iiik, bill .lie of the rank of sections. .As a sub-
stitute Ini the section the term group is employed.
The peicnnial species placed in the subgenus Oreo-
carya are evidently the most primitive in the genus.
The most primitive of these is C. janu'sii var. nutlli-
caiilis. The vaiietv lUuUicaulis has so nuins piimitixe
characteristics thai it may be used lo visualize ihe
ancestor from uliicli the other siibgeneia developed.
"Ihe authoi ichlIs lo agiee wilh Payson (1''2^) in his
postLilation thai the piimilue ()ieocar\a species
evolved in loin dilleieni directions. The species of
the main section Oieocarva remained perennial and
developed a tendency toward larger flowers and
shorter cymes. This larger flowered gioup is probably
monophyletic in the setise that a single piimiiive

HIOLOCKAL SKRIKS VOL. 13, NO. 4 A RIVISION Ol CRVI'I ANTHA SUBGENUS OREOCARYA

ancestor gave rise to several basic t\ pes and liicse in
turn produced species as we know them. Deveiop-
inenl in another direction produced annuals very
eaii\ ni the history ol" the genus. This group is recog-
nized as the subgenus Krynitzkia. with its reduced
llowers, heteroniorphic nutlets, and elongate cymes.
This subgenus is probably polyphyletic, the species in
Krynitzkia apparently having arisen independently
Irom several perennial ancestors. The other two sub-
genera. Geocarya and Cryptantha had primitive per-
ennial ancestors and probabK' developed along the
same lines as Krynitzkia. since the species in these
subgenera are similar in appearance to Krynitzkia.
Geocarya is probably monophyletic. Evidence for this
is indicated in the peculiar morphology of the cleisto-
gamous llowers. The subgenus Cryptantha. may have
developed along several hues from Krynitzkia.

The groups that occur within the subgenus Oretv
carya are characterized, discussed, and arranged in a
phviogenetic scheme (Fig. 1 ).
flavoculata

elata

humilis

nubigena

virgata

setosissima

I ij:. 1. A proposed pin logcnctic urrangcnicnt ol the groups
wiltiin the siibiicnus Orcocurv u.

The jamesii group: This group is compt)sed o( live
species and seven varieties mostly confined to the
eastern and southeastern section of the range of the
subgenus (Fig. 2). It is characterized by the hemi-
spherical fruit, with the nutlets smooth on all sur-
faces, or rugose and tuberculate, but not at all
muricate on the dorsal surface. Ventral surface of ihe
nutlet quite or nearly smooth, with the scar narrow,
straight, and closed, and without an elevated margin.
The species within the jamesii group form a natural

unit. Cryptuiulhi jamesii has more primitive char-
acters (smooth hemispherical nutlets, short corolla
tube with crests at the base, low fornices. oblance-
ohite leaves, strigose pubescence, and elongate cymes)
than any other species in the subgenus and is prob-
ably similar to the hypothetical, ancestral form. C.
palmcri. although very closely related to C. jamesii,
shows the advanced characteristics of crests lacking at
the base of the tube, accrescent sepals, and longer
style. C. crassipes is to be considered more advanced
than C. palmcri on the basis of the capitate in-
tlorescence. rougliened nutlets, and the longer style,
which are all considered as specialized characters.
It seems probable that long corolla tubes are derived
from shorter i)nes and that heterostyled tlowers are
derived from uniform llowers. On this basis C oblata
is more specialized than C. jamesii. C. paysonii prob-
ably very recently has evolved from C. oblata. but it
is more specialized because of the strongly hetero-
styled flowers and the more capitate inllorescence.

paysonii

crassi

palmer

oblata

lig. 2. A proposed phylogenetie arrangement of Ihe spcejes
wuliin the janicsii group.

The flava group; The flava group is composed of
seven species, most of which are confined to Utah
and northern Arizona (Fig. 3). The basic syndrome of
characters includes the compressed nutlets which are
smooth on both surfaces and the closed scar which
lacks an elevated margin. This group is recognized as
being closely related to the jamesii group primarily on
the basis of the smooth nutlets. The species within
this group, however, become more difficult to place
because lines of development are more obscure. C.
salmanensis is probably the most primitive in this
group because of its short corolla and unit\)rm style
length. All other species within the group possess
corollas that exceed the lobes of the calyx. C. caii-
feriithmi is a widely distributed species of the hot
desert regions of Utah, Arizona, Nevada, and Cal-
ifornia and may have, through selective forces of the
environment, given rise to most of the other species.

BKIC'.HAM YOUNC; UNIVKRSITY SCIlNfl. liULLl 1 IN

many of which are narrow endemics, confined to ;i
particular kind of habitat. C. semiglahra. C. capitata.
and C. johinlonii are closely related to C. confcr-
tijlora, but probably because of their ability to
inhabit peculiar soil types, have been selected out and
isolated from that species. C. leucophaea. a species of
eastern Washington, which inhabits sand dunes along
the Columbia River, was for a time included within
the concept of C. confcrtijlora by Parish. It is be-
lieved, however, to be more advanced because of the
longer intloresccnce, conspicuous foliar bracts, and
narrowly lineai leaves. The aulluir, at one time, was
inclined to combine C. jlava and C con fen i flora
because of a few intermediates where the ranges of
the two species overlap. However, with additional
research the number ol' intermediates were I'ound to
be so few that the two taxa are best treated as
separate species.

flava

stricta

barnebyi

capitata

johnstonjj

leucophaea

glabra

confertiflora

lig. y. A proposed phylogetictii.- ;irrangemeiit of the species
within the tlava group.

The stricta group: The stricta group contains two
species. C. srriciu and C. barnebyi. both restricted to
the Uintah Basin of Utah (Fig. 4). The stricta group
probably had its origin trom the tTava group. The very
setose or hispid indument. stout stems, and the
smooth ventral surface of the nutlets separate this
group from that of llava. C. stricta is considered more
advanced than C. barnebyi because of the roughened
dorsal surface of the nutlet.

The nubigena group: This group is composed of four
species, all of higli alpine or montane areas of Cali-
fornia, Oregon, and Idaho (Fig. 5). The basic charac-
teristics of the group involve the lanceolate nutlets.
which are smooth on the ventral surface and rinigh-
ened dorsally. This group probably had its origin
from the flava group through some form similar to C.
confertiflora. The three species, C. nulngena, C.

I ig. 4. A pioposetl phylogenetic urraiigemeiil ol the species
wilhin the stricta group.

crymophila

hypsophil

subretusa

I ig. 5. A proposed phylogenetic urrangenieiit ol the species
vvithni the nuliigena group.

.subretusa and C liypsophila. are so closely related
that it is difficult to atTord tlicm complete specific
rank: but because of the wide geographical separation
and the slight differences in morphology, they are
treated as distinct species. C. crymophila is pinhably
more advanced than the other three species because
of the hiigei iiuilots. more accrescent sepals, and the
longer foliai bracts. It was apparently derived trom
some lorm similar to C. nubigena.
The abata group: This group is mainly confined to
the southwestern part of the range of the subgenus. It
contains five species which are characterized by
rugose or tuberculatc nutlets, with the ventral scar
open and triangular and usually surrounded by a
slightly elevated margin (Fig. 6). These characters are
all much more advanced than those displayed by the
jamesii group; so it is probably not closely related to

mOLOClR AL SERIES VOL. I 3. NO. 4 ,^ KIVISKJN Ol (.KV I' I \N I H.-\ SLIBGENUS OREOCARYA

that gmiip. All the species in this giotip oeeiii in the
Soniiian Desert except lor C. ahata. C. iKiffnuinnii is
probably a recent derivative from C. rirgliwnsis. The
two species appear to be distinct even thongii they
are quite similar in general appearance. C. Ihiff'iu/iinll
tlowcrs much later and has nontragrunt (lowers. The
relationship of the other three species is not so ob-
vious; however, they appear to have been derived
tioni some common ancestor similar to abata.

hoffmannii

virginensis

tumulosa

insolita

f'Ig. 6. \ proposed phylogenetic arrangement of the species
within the abata group.

The caespitosa group: This group contains two
species, both of which are apparently derived from
the abata or possibly the humilis group. The distribu-
tion of this group is bicentric. C. caespitosa is a very
hummock fortiiing species found throughout south-
ern Wyomitig growing on clay hillsides, while C.
ochruleuca is a narrow endemic growing on gypsifer-
ous soil in western Garfield County, Utah. The two
species probably had a common ancestor similar to C.
caespitosa.

ochroleuca

caespitosa

The Inimilis group: The humihs group is composed
of eiglit species and five varieties (Fig. 8). The basic
syndrome of characters in this group includes the
conspicuously muricate nutlets and a scar with a
tendency to be open and therefore triangular at the
base. Two basic lines of development can be seen
within this group; one in which the species possess
two kinds of hairs and nutlets which have short rugae
between the murications; the other line has leaves
which are usually silky-strigose and only murications
on the nutlets. C. caiia and C. brerijlora are members
of this second group. C. jonesiana and C. fulvocan-
escens are considered advanced because of the long
corolla tubes and more elongated nutlets.

fulvocanescens

propria

Fig. 7. A proposed phylogenetic arrangement of the species
within the caespitosa group.

I ig. S. .-\ proposed phylogenetic arrangement of the species
w ithm the humilis group.

The elata group: The elata group is composed of thir-
teen species, covering a wide geographical and altitud-
inal range (Fig. ^^). The basic syndrome of characters
includes nutlets which are always roughened dorsally
and distinctly rugose or tuberculate or both, and
often muricate also; ventral surface smooth or vari-
ously roughened; scar straight, closed or nearly so,
margins not elevated; corolla tubes never longer than
the calyx lobes, except in C roUinsii. The species in
this group are so similar in overall appearance and
characteristics that lines of development are ob-
scured. C. aperta and C. thyrsiflora seem to form a
natural unit which possess broad inflorescences and
ovate tuberculate nutlets. C. internipta. C. spiciilifera.
C. shackletteana. and alsti C. ntgiilosa seem to form a
natural unit and are probably very closely related to
each other. The lanceolate nutlets which have similar
markings tend to substantiate this hypothesis. C. celo-
sioidcs. a widespread and heteromorphic species,
apparently gave rise to C. sobolifera and C. thoinp-
sonii. The other species in this group are more ob-
scure in their evolutionary history; probably because
they are very narrow endemics which have specialized
out on peculiar soil types.

10

BRIGHAM YOUNG UNIVERSITY SCIF.NCF. BULLI TIN

Ihompsonii

8hackl«tteana

splcullfera

celosioides

thyrsiflora

I'ig. 9. A proposed plnlogcnetic arr;iiigL'nicnl ol llic spci.'n.'s
within the elala group.

The flavoculata group: The tlavoculala group is
composed of eiglit species, most ol" vvliich are con-
fined to eastern Utali and western Colorado (Fig. 10).
The basic syndrome of characters inckides the deeply
and conspicuously rugose and tuberculate. sometimes
muricatc or foveolate nutlets. Scar, except in C.
bakeri. at least slightly open and then showing a tend-
ency to be constricted above the base; margin usually
elevated. The species of this highly developed group
form so natural a unit thai there is little doubt that
tliey had a common origin. C. oslerhmirli and C.
tenuis are two very closely related species, but the
longer corolla tube and style of C. tenuis evidently
make it more advanced. C. Jhvoculata and the closely
related species C. paradoxa. C. Iiakcri. and C. nicn-
sana are the most advanced in the subgenus and un-
doubtedly are very closely related. C. wctlichllii and
C. longijlora are somewhat intermediate between this
group and the elata group, but seem to be more
closely related to the flavoculata group on the basis
of the nutlet characteristics.

The virgata group: This moni>typic group is so differ-
ent from the other basic groups that it is separated
from them in this treatment. The characteristics
which distinguish this distinctive group are the stout,
strict, usually simple, fistulose stems and the long

I ig. in. A proposed phyiogeiielic arrangement ot" the species
within the navocidata group.

spicate inilorescence with the greatly elongated foliar
bracts.

The setosissima group: This monotypic group is re-
stricted to montane areas throughout southern Utah,
Arizona, and southern Nevada. The basic syndrome
of characters includes the simple erect stems, broadly
winged nutlets, and ovate caly.K segments.

Cytology

Very little cytological work has been done on the
subgenus Oreocarya. Delbert Wiens at the University
of Utah began a study of the morphology of the
group; but because of the difficulty he had in being
able to get seeds to germinate, the stud\ was aban-
doned. He did. however, make chromosome counts
on several of the species in this subgenus. The number
in all species for which counts were made was n equals
twelve. These species were C. virgata. C. jamesii. C.
t/iyrsijlura. C. Jlara. C. Inikcri. and C. Julmcanesccns.
Taylor and Brt)ckman counted the chromosomes in
r. cclosiDiJcs (C. macounii) and determined the
numbei to be ii equals nine. There are some discrep-
ancies in the basic number ol this subgenus, which
points out the lacl ihal nuich additional study is
needed on the c\ toli)g\ ol this group.

TAXONOMY

Cryptantha Lehm. subgenus Oreocarya (Greene) llig-
gins

Perennial or biennial herbs with a conspicuous
setose, hirsute, or strigose indumeni (except in C.

jiiincsii var. pusiulosu and C. scniiglal'ia). Leaves en-
tire, oblanceolate, spatulate or linear. Stems solitary
Irom the root or caespitose, commonly unbranched
below the inflorescence, 0.2-1 2 dm tall. Inilorescence
a coniimious or glomerate cluster of elong;iting or

HIOLdC'.IC \l SI Kll S VOL. 1 .1, NO. 4 ..\ Kl VISION Ol (.R'l I'i .XN I H.\ SUBGHNUS ORI'OCAKY.\

11

redLicei.1, simple or biandicd. bnicteato or nearly
ebracleate, two-ranked, unilateral, scorpioid cymes.
Calyx segments distinct, usually conspicuously accres-
cent. Corollas white or yellow, salverform or campan-
ulate; limb 4-1 7 mm broad; tube equalling or exceed-
ing the calyx. Stamens included in the tube, anthers
sessile oi nearly so. Style shorter, equalling or much
exceeding the mature truit; stigma entire. Nutlets
from nearly circular in outline to narrowly lanceolate,
margined or winged, smooth or variously roughened
on the different smfaces, attached to the gynobase at
a pomt I J> to 4,'5 of the distance from the base to
the apex of the nutlet. Scar of nutlets various, open
or closed, and margin elevated or plane. Subgeneric
type: C. cclosiaidcs (Eastw.) Payson.

Key to the Species of Cr_\ piantha Sidigenus Ore^)carya

1. Corolla tube elongate, distinctly surpassing the

calyx; flowers usually heterostyled (2).
I. Corolla tube short, scarcely if at all siupassing the
calyx; tlowers not heterostyled (20).
2. Nutlets smooth and shiny (3).
2. Nutlets more or less rougliened or wrmkled at
least on the dorsal surface (9).
.V Corolla yellow (4).
3. Corolla white (5).

4. Intlorescence an elongate, cylindrical tliyrse;
nutlets lanceolate, with acute margins, usually

only one developing 7 C Jhra

4. inflorescence consisting of a large terminal
cluster with one or more remote, at maturity
frequently stalked, much smaller lateral clust-
ers; nutlets briiadly ovate, with winged margins,
all four usually maturing ... .6. C. confvnlllnni
5. Inflorescence capitate. 0.1-0.4 dm long; corolla
limb 6-8 mm broad, the tube little surpassmg the
calyx; nutlets lanceolate; native lo northern Ari-
zona and southern Utah 8. C. capitata

5. Inllorescence elongate. 0.4-4 dm long; coriilla limb

8-17 mm broad, the tube distinctly surpassing the

calyx except in C barncbyi: nutlets ovate (6).

6. Ventral surface of the leaves glabrous; native to

north-central Arizona and southwestern Utah . .

1 0. C scmlglabni

6. Ventral surface of the leaves strigose or setose-
hispid (7).
7. Corolla limb l.vl7 mm broad, crests at base of
tube absent; nutlets .i-,i.5 mm long; native to San

Rafael Swell 9. C. joluisionii

1 . Corolla limb 8-1 I mm broad, crests at base of tube
conspicuous; nutlets 3.5-4.5 mm long (8).
8. Corolla tube 5-7 mm long, the limb campanu-
late; leaves broadly oblanceolate, setose-hispid;
stems stout; endemic to the Uintah Basin, Utah

I I . C hanu'hyi

8. Corolla tube 8-10 mm long, the limb spreading;
leaves linear or narrowly lanceolate, strigose

with few or no pustulate hairs; stems slender;

native to Washington. 12. C. leitcophaea

■■). Nutlets uniformly muricate or papillose, or some-
times in C. jonesiana also with some inconspicuous
ridges (10).
9. Nutlets more or less rugose oi luberculale, or
sometimes with a few inconspicuous murications
(II).
10, Leaves oblanceolate, strigose with pustulate
hairs small or lacking; corolla 7-10 mm long,
fornices elongate; native to southeastern
Utah, northwestern New Mexico, western

Colorado, and northeastern Arizona

34. C. fiilrdccmcscciis

10. Leaves spatulate, hispid with pustulate
bristles; corolla 10-15 mm long, fornices low
and broad; native to the San Rafael Swell,

Utah 35. C jonesiana

I 1 . Ventral or inner surface of the nutlets smooth or

nearly so (12).
II. Ventral surface of the luitlets distinctly rough-
ened (15).

1 2. Plants biennial; corolla campanulaie. crests at
base of tube evident; nutlets lanceolate; na-
tive to the Uintah Basin, Lltah

49. C. roUinsii

12. Plants perennial; corolla salverform, crests at

base of the tube lacking; plants of Texas and

New Mexico (13).

13. innoresence capitate; nutlets 3.3-3.8 mm long;

leaves densely white strigose or subtomentose;

native to Brewster county, Texas . .5. C. cnissipes

13. Inflorescence elongate; nutlets 2.5-3.2 mm long;

leaves strigose and setose-hispid (14).

14. Corolla tube 7-10 mm long; calyx lobes 5-7

mm long in anthesis; plants not heterostyled;

nutlets conspicuously tuberculate and short

rugose; native to Texas ?■■ C. oblata

14. Corolla tube 12-14 mm long; calyx lobes 7-9
mm long in anthesis; plants strongly hetero-
styled; nutlets finely tuberculate or rugose;
native to New Mexico and western Texas . . .
4. C paysonii

15. Leaves conspicuously pustulate ventrally; corolla
tube 12-16 mm long; calyx segments 7-10 mm
long in anthesis; native to west-central Colorado

and east-central Utah 51 . C longifhra

15. Leaves sparsely if at all pustulate ventrally ; cor-
olla tube 5.5-12 mm long; calyx segments 3-5.7
mm long in anthesis (16).

16. Inflorescence 0.1-0.4 dm long; corolla tube
1 O-I 2 mm long; margin of nutlets not in con-
tact; plants less than 1.2 dm tall; eastern
Lltah and western Colorado. .54. C. parado.xa
16. Inflorescence 0.5-3 dm long; corolla tube
5-10 nmi long; margin of nutlets in contact
or ncarlv so; plants usually over 1.2 dm tall
(17).

12

IfHK.llAM YOUNC; UNIVI-RSIIV SC'IKNCK liULl I-IIN

17. Scar of nutlets surrounded by ;in elevated margin 27.
but lightly closed;style 1-2 mm long: calyx 3.5-4
mm long in anthcsis; native to southern Utah,
southwestern Colorado, and northeastern Arizona 27.

55. C bakeri

17. Scar of nutlets conspicuously open; style 3-8 mm
long; calyx 4.5-7 mm long in anlhesis (18).
18. Scar of nutlets conspicuously open and sur-
rounded by a definite elevated margin; wide-
spread ^1-C. flavocidala

18. Scar of nutlets slightly open and with only

an inconspicuous elevated margin if any ( 1^)).

14. Leaves linear-spatulate; nutlets sharply and

deeply rugose; corolla tube 5.5-7 mm long, for-

nices low and broad; native to southeastern Utah

52. C. tenuis

19. Leaves obovate or broadly oblanceolate; nutlets 29.
with rounded ridges and tubercles; corolla tube
7-10 mm long, fornices long papillose; native to 29.

eastern Utah 50. C wt'lhcnllii

20. Nutlets smooth on their dorsal surface, not

rugose, muricate or tuberculate (21).
20. Nutlets more or less roughened, muricate,
rugose or tuberculate at least on the dorsal
surface (26).
21. Fruit depressed globular, nutlets not in contact

by their margins (22).
21. Fruit conical, ovoid or lanceolate, nutlets in con- 31
tact by their margins or nearly so (23).
22. Crests at base of corolla tube conspicuous;
calyx not conspicuously accrescent; wide- :; |

spread I ■ C jamesii

22. Crests at base of coriilla lube obsolete; calyx
conspicuously accrescent; southwestern New
Mexico, western Texas and northern Mexico

1. C. palmcri

23. Stout, strictly erect plants with many elongated 33
and conspicuous bracts in the inllorescence;
southeastern Wyoming and central Colorado ....

20. C. virgala },},

23. Smaller, usually caespitose plants with few or in-
conspicuous bracts in the inllorescence; native to
west of the continental divide (24).
24. Corolla tube ^-7 mm long; style 5-6 mm
long; calyx 5-7 mm long; cntlemic to the

Uintah Basin, Utah I 1 • C banwbyi

24. Corolla tube 2-4 mm long; style 0.5-2 mm
long; calyx 2.5-4 mm long (25).
25. Nutlets ovoid; corolla tube 3-4 mm long, limb

7-10 mm wide; plants of central Idaho 35

I 3. C siilnioiicnsis

25. Nutlets narrowly lanceolate; corolla tube 2-2.5

mm long, limb 3.5-5 mm wide; native to the high 35
sierras in southern California ... 15. C mihigcna
26. Ventral surface of the nutlets smooth or

nearly so (27).
26. Vemral surface of the nutlets rugose or vari-
ously wrinkled (40).

Nutlets bordered by a conspicuous wing; robust
plants 5-10 dm tall, with long ebracteate spikes . .

19. C. setosissima

Nutlets never conspicuously winged, sometimes
with an acute margin simulating a narrow wing;
plants usually lower and caespitose; inllorescence
bracteate (28).

28. Inllorescence a virgate spikelike thyrsus with
all but the uppermost tloral bracts much
longer than the short cymes; nutlets broadly
ovate, sparcely rugose or smooth; native to

Wyoming and Colorado 20. C. virgala

28. Inflorescence various but seldom if ever so

spikelike, and at least the upper lloral bracts

reduced to short bracts which slightly if at all

exceed the cymes or racemose branches (29).

Corolla tube 7-9 mm long; calyx 6-9 mm long in

anthesis(30).

Corolla tube 2-6 nnii long; calyx 2.5-6 mm long
in anthesis (31 ).

30. Inflorescence capitate, 0.2-0.8 dm long; nut-
lets ovate; leaves densely white strigose or
sublomentose; native to the Big Bend Region

of Texas 5.C. crassipcs

30. InlliMescence elongate, 0.6-2 dm long; nut-
lets lanceolate; leaves setose-hispid; native to

Utah 49. C. rollinsii

Inllorescence very broad and rounded in outline;
native to the eastern slope of the Rocky Moun-
tains 36. C tliyrsijldiv

Inllorescence narrower; plants west of the conti-
nental divide (32).
32. Fruiting calyx ''-14 mm long; nutlets 4-6 mm

long (33).
32. Fruiting calyx 4-9 mm long; nutlets 2.5-3.8

mm long (34).
Nutlets lanceolate, the scar narrowly subulate but
open at the base; native to Alpine and Tuolumne

Counties, California 18, C. aymopliiia

Nutlets more ovate, the scar broader and cuneate
at the base; plants of the high mountains in Kit-
titas and Chelan Counties, Washington

46. C. ihonipsonii

34. Nutlets 2-2.3 mm long, the scar cuneate or
narrowly triangular; plants of Saguache and
Hinsdale Counties, Colorado . . 40. C wvlu'ri
34. Nutlets 2.6-3.7 mm long, scar closed or nar-
rowly linear and open only at the forked
base (35).
. Plants conspicuously setose-hispid; nutlets trans-
versely rugose and tuberculate; plants of Color-
ado, Utah, and Nevada (36).
. Plants strigose or setose, but not as above; nutlets
usually tuberculate or short rugulo.se; plants of
the high moLUitains in California. Oregon, Idaho,
and Montana (37).

36. Nutlets scarcely or not at all muricate be-
tween the rugae; strictly erect, conspicuously

BIOLOC;i(AI SI KIIS VOL. I 3. NO. 4 .A Kl VLSIONOI CK ^ I' LAN IH.-X SLIBGl'.N US ORF.OC^R Y .-^

37.

37.

3*^).

39.

41.
41.

43

49.

49.

45.
45.

liispKl poicniiials t'riMii iKirthwcslcm Colo-
rado and iioitheastein Utah . . .14. C. srricta

36. Nutlets distinctly muricate or tuberculatc
between the rugae and near the inargiiis;
erect perennials tVom western Lltali and east-
ern Nevada 41 . C riigulosa

Nutlets with tubercles but no conspicuous irans- 47

verse ridges, or sometimes nearly smooth; native

to California 1 5. C. mhigena 47

Nutlets with evident ridges on the dorsal surface;

plants of northern California. Oregon, Idaho, and

Montana (38).

38. Style 1.8-2.5 mm long; st)boliferous peren-
nials from western Montana

47 . C sobolifera

38. Style 0.5-1.5 mm long; plants of Idaho and
Oregon (39).

Leaves oblanceolate, acute or obtuse, spreading

bristly setose; style 1.2-1.5 mm long; central Idaho
1 7. C liypsapliila

Leaves spatulate, subretuse or obtuse, subtomen-

tose or strigose; style 0.5-1 mm long; Oregon.

northern Cantomia. and northwestern Nevada . .
1 6. C. siihrclusa

40. Nutlets conspicuously muricate. or in C.
humilis also with a few irregular ridges (41 ).

40. Nutlets not exclusively muricate, but rugose
or tuberculate, also with a few muncatioiis
between the ridges (44).

Leaves distinctly subtomentose or tomentose,

also setose in C. humilis (42).

Pubescence of the leaves silky-strigose or strigil-

lose but not subtomentose or tomentose (43). 5 1

42. Plants 0.3-1 dm tall; leaves 0.5-2.5 cm long;
caly.x 2-2.5 mm long in anthesis; corolla tube 51
1.8-2.2 mm long; native to Millard County.
Utah 30. C. couipacta

42. Plants 0.4-2.5 dm tall; leaves 2.5 cm or
longer; calyx 3-5 mm long in anthesis; cc)ri)lla
tube 3-5 mm long; plants widespread. Colo-
rado to California 28. C huniilis

Plants densely caespitose, caudex multicipital;

leaves linear oblanceolate; native tt) eastern

Wyoming and adjacent Nebraska and Colorado . .
31 . C cciiia

Plants scarcely t)r only moderately caespitose;

leaves broadly oblanceolate or spatulate; native

to the Uintah Basin in northeastern Utah

32. C. brevijlora

44. Scar of nutlets open some distance above the
base (45).

44. Scar of nutlets closed or nearly so. without a

conspicuous triangular opening toward the 55
base (58).

Scar somewhat constricted some distance below

the middle of the open portion (46).

Scar triangular and not constricted below the 55

middle (47).

53,
53

46. Elevated margin of the scar definitely lim-
ited; pustules present on both leaf surfaces;

central Utah 56. C. mensana

46. Elevated margin indefinitely limited; pustules
present only on dorsal surface of the leaves;

southeastern Utah 53. C. osurliotitii

Some tendency to an elevated margin evident
around the scar (48).

No tendency to an elevated margin aroiuid the
scar (52).

48. Cymules elongating and so the inllorescence
broad; biennial or short-lived perennials; nut-
lets usually with an evident dorsal ridge (49).
48. Cymules shorter and the intlorescence nar-
row; long-lived perennials; nutlets with only
a slight dorsal ridge if any. (51 ).
Surface of the leaves with inconspicuous ap-
pressed bristles; inflorescence open, with only a
few elongate cymules, 7-14 cm long terminating
the stem; endemic to near Las Vegas, Nevada. . . .

22. C insolita

Surface of the leaves conspicuously setose-hispid
with spreading bristles; inllorescence open, at
least at maturity (50).

50. Calyx lobes 7-12 mm long in fruit; nutlets
3-4.5 mm long; prominently carinate on the
dorsal side; southwestern Utah to southern

California 23. C. virgineiisis

50. Calyx lobes 5-7 mm long in fruit; nutlets
2.5-3 mm long, with only an indistinct cen-
tral ridge toward the apex; eastern Calilornia

and western Nevada 24. C hofjiitannii

Nutlets indefinitely tuberculate and rugose; Cali-
fornia and southern Nevada .... 21. C liiimilosa
Nutlets definitely tuberculate o\ rugose; native to

Utah and eastern Nevada 25. C ahata

52. Style not exceeding the matiue nutlets by
more than 0.5 mm; plants usually less than
1.3 mm tall (53).
52. Style exceeding the mature nutlets by 1 mm
or more; plants usually taller than 1.3 dm
(56).
Corolla tube 3-4 mm long; nutlets 3-3.5 mm long
(54).

Corolla tube 2-2.6 mm long; nutlets 2.3-3 mm
long (55).

54. Ventral surface of nutlets deeply rugose and
tuberculate; native to southern LItah and

eastern Nevada 25. C abata

54. Ventral surface of nutlets mdefinitely muri-
cate; native to southern Wyoming

26. C. caespi!()sa

Infiorescence less than 2 cm long; calyx segments
3-4 mm long in fruit; plants 0.1-0.3 dm tall; en-
demic to Inyo County. California

29. C. roosidrunt

inflorescence 2.7 cm long or longer; calyx seg-
ments 4-6 mm long in fruit; plants 0.2-1.3 dm

14

HKIl.HAM Y(;UN(; UNIVKKSIIY SCIKNC'K [iULLKIlN

57.

57.

tall: endemic to Garfield County. Utah

27. C oclirolciica

56. Leaves setose-pustulate and tonientosc; nut-
lets muricate or with a few sht)rt rugae

28. C luiniilis

56. Leaves finely strigose and appressed setulose;

pustulate hairs lacking or inconspicut)us on

the ventral surface of the leaves; densely

caespitose perennials (57).

Leaves linear to narrowly ohianccolate; corolla

tube 3-3. .> mm long: nutlets inconspicuously

tuberculate and rugulose: native to Alaska

44. C shacklaicaiui

Leaves ohianccolate to spatulate; corolla tube
3.5-4.5 mm long: nutlets muricate and irregular
rugose: native to Oregon and ucslcm Idaho ....

V V C. prnpria

58. Upper surface of the leaves uniformly ap-
pressed strigose and uithonl pustulate hairs
(59).
58. Upper surface of the leaves with two distinct
kinds of hairs: pustulate at base (63).
59. Nutlets sharply rugose and tuberculate. scar sur-
rounded by an elevated margin ... .55. C bakeri
59. Nutlets ntit so sharply rugose or tuberculate: scar
not surrounded by an elevated margin (60).
60. Leaves linear or narrowly oblanceolate. 2-13
cm long. 0.1-0.5 cm wide: native to Alaska . .

44. C shacklencaiui

60. Leaves shorter and broader; plants from tar-
ther south (61).
61. Corolla tube 2-2.5 nnn long: style exceeding nut-
lets hv I mm or less: endemic to Garfield

County, Utah 27. C. ochroleitca

61 . Corolla tube 3.5 mm long or longer: style exceed-
ing nutlets by more tluin 1 nim(62).
62. Densely caespitose perennial from a multi-
cipital caudcx: native to eastern Oregon and

western Idaho 3,3. C. prupha

62. Less evident or not at all caespitose: native to

Utah. Colorado, and Wyoming

38. C scricea

63. Mature calyx exceeding the nutlets by 2-4 mm:
infiorescence broadtopped: wcslerii Colorado and

eastern Utah 37. C. data

63. Mature calvx exceeding the nutlets bv 4-8 mm
(64).
64. Nutlets tuberculate, scarcely if at all luuose

(65).
64. Nutlets more or less rugose (69),
65. Ventral surface of the nutlets smooth or nearly
so: native to high mountains in western Montana

47. C. SDholifciv

65. Ventral surface of the nutlets distinctly rough-
ened (66).
66. Plant 1-2 dm tall: native to western Colorado

and eastern Utah (67).
66. Plants 2 or more dm tall (68).

67. Corolla tube 2.6-3 mm long: calyx segments 2.8-3
mm long in anthesis: nutlets 2-2.6 mm long:

endennc to Mesa County, Colorado

39. C. apcrta

67. Corolla tube 3.5-5 mm long: calyx segments 5-7
mm long in anthesis: nutlets 3-3.8 mm long:
native to Uintah County, Utah . .48. C. gruhauiii
68. Nutlets broadly lanceolate: murications lack-
ing or indefinite: infiorescence sliglitly open
to very broad; widespread, from North

Dakota to Washington and Oregon

45. C. cclosioidcs

68. Nutlets narrowly lanceiilate: muiications or
tuberculations very definite: LIko County,

Nevada 42. C. inlcrnipia

69. Scar of the nutlets sumewhat open at the base

28. r. hitmilis

69. Scar of the nutlets closed or nearly so (70).

70. Infiorescence very broad and open; plants
native on the eastern slope of the Rocky
Mountains, from southern Wyoming to

northern Texas 36. C. thynifloni

70. Inflorescence narrower; plants more nor-
therly in range (71 ).
71. Leaves narrowly oblancenlale, strongly setose-
ciliate on the margins: stems slender; native to

eastern Washington and Oregon, and Idaho

43. C spiculifera

71 . Leaves usually broader, oblanceolate to spatulate,
the margins not strongly setose-ciliate; stems
more robust ( 72).
72. Inner surface of the nutlets conspicuously

rugose or tubeiculate; widespread

45 . C celosioides

11. Inner sin lace of the nutlets smooth or nearly
so (73).
73. Leaves soboliferous, oblanceolate to spatulate,
setose; native to high mountains of western Mon-
tana 47. C sobolifcra

13. Leaves not soboliferous, spatulate, subretuse,
subtomentose; native to northern California and
Oregon 1 6. C. subrctusa

I . Crvpiaiillui Hiinvsii (Torr.) Payson

Perennials. 1-6 dm tall; stems onc-man\. 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 inconspicuous; infiorescence open,
cy mules usually elongating, tomentose to .setose-
hirsute, fioral bracts inconspicuous to very conspicu-
ous; calyx segments ovate-lanceolate, acute, in anthe-
sis 3-4 mm long, in fruit 5-7 mm long, subtomentose
to setose-hirsute, (tir sometimes nearly glabrous):
pedicels 1-3 mm long: corolla white, the tube 2.5-3
mm long, crests at base of tube conspicuous, fornices

BIOLOGICAL SERIKS VOL. I .i, NO. 4 .X RHVISION Ol' CR V I' 1 .XN rH.<\ SL'BCILNUS ORICOC'AR Y ,\

15

lighi-yellow. eniaiginate, 0.5-1 mm long, limb 5-8 mm
bioad; style exceeding mature tVuit 1-3 mm; tVuit
oblate-ovoid. 1-4 nutlet.s maiming, ovate-lanceolate,
margins acute. 2-2.5 mm long. 1.5-2 mm wide, the
maigms not m coiilacl. both surfaces smooth and
glossy, scar straight, closed, extending from the base
to near the apex, elevated margin lacking.

Key to the varieties of C. jamcsil

I , N'entral surt'ace of the leaves glabrous, the petioles

not ciliate-margined, nor tufted at the base

Ig. var. piisiiildsu

I . Ventral surface of the leaves strigose or setose, the
petioles ciliate-margined; leaves tufted at the base
(2).

2. Stems simple, not branched above the base (3).
2. Stems branched from the base as well as above
(5).
.V Stems 1-4.4 dm long, usually twice as long as the

basal tuft of leaves la. var. imilticaiilis

3. Stems 0.2-0.9 dm long, usually not exceeding the
basal tuft of leaves (4).
4. Floral bracts exceeding the cymules; stems low,

decumbent; Nevada and California

Id. var. ahoriiva

4. Floral bracts not exceeding the cymules; stems

erect or nearly so Ic. var. setosa

5. Stems decumbent; plants of the Great Plains

If. var. jamesii

5. Stems erect; plants west of the continental divide
(6).

6. Leaves linear; cymules 8 cm long or longer, very
lax; native to southern New Mexico. Texas, and

Mexico lb. var. laxa

6. Leaves oblanceolate; cymules usually much

shorter than 8 cm long, and more congested . . .

le. var. disiiclia

la. var. imilticaulis (Torr.) Payson

conspicuous, the petioles conspicuously ciliate on the
margins; intlorescence open, 0.5-1.5 dm long, bracts
inconspicuous. Collections: 387 (x); representative;
Jones 4007 (ARIZ, GH, US, UTC); J. M. Tucker
2771 (GH.ORE);R. C. Rollins 2429 (GH, US, UTC);
B. Maguire I 1975 (ARIZ, UTC); 0. B. Metcalle 70
(ARIZ, GH, ND-G, US); L. C. Higgins 3136, 3169,
3595 (BRY,WTSU).

Holotype: Fendler 636, collected in New Mexico
near Santa Fe, 1847, NY. Isotypes at GH, US.

Distribution: Southern Colorado and eastern New
Mexico, south to western Oklahoma and Texas into
northern Mexico, north through central Arizona to
southern Utah. Growing on a wide variety of soils,
4,500 to 8,000 feet. Map No. la. April to September.

Map No. la. Parts of southwestern United States. Range of
C. jamesii i.lorx.) Payson var. mutticaidis (Torr.) Payson.

Crypiaiilha iainesii (Tiirr.) Payson var. miilticaulis
(Torr.) Payson. Ann. Mo. Bol. Gard. 14:244.
1927.

Eriirichium miilticaule Torr. ni Mare\. 1 xplorallon Red

River. 262. 1854.
Oreocarya mullicaulis (Torr.) Greene. Pitt. 3:114. 1896.
Oreocarva suffniiicosa (Torr.) Payson var. inullicaiitis

(Torr.) Payson. Univ. Wyo. Publ. Bot. 1:171. 1926.
Ih'iiusphairocarya suffniiicosa (Torr.) Brand var. iiui/li-

caiilis (Torr.) Brand, l-'cdde. 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, oblanceolate, 5-15
cm long, 0.4-1 cm wide, dorsal surface strigose and
approssed setose, or sometimes setose-hirsute, pustu-
late, ventral surface uniformly strigose or subtomen-
tose, without pustules, or the pustules small and in-

The original description of /:". nntlricaiilc Torr.,
was based on a collection from near Santa Fe with
setose-hirsute pubescence. This same bristly form also
occurs in the White Moimtains and southward to the
Santa Catalina Mountains of Arizona. A strigose or
subtomentose form occurs about Flagstaff, also into
southwestern New Mexico, and north into southern
Lllah. In northern Arizona var. nmlticaiilis may be
confused with var. setosa. It can be separated from
var. sclosa by the longer stems, becoming twice the
length of the basal tuft of leaves, and the individual
cymes which are longer and more perfectly devel-
oped. On its eastern boundary it may be confused
with var. jamesii, but differs in the simple stem which
is never branched above the base, more perfectly
developed cymes, and the inoic dense strigose pubes-
cence.

16

BKIC.H AM YOUNG UNIVKKSITY SCIKNCK BULLKTIN

lb. var. laxa (Macbr.) Payson

Crypiantha lanu'sii (Tun.) Payson vai. lu.xu (Macbr.)

Payson, Ann. Mo. Bot.Gaid. 14:246. 1427.

Oreocana mutticaiilis var. laxa Macbr. Contr. Clray llcrli
48:35. 1916.

Hcmhphaciocarva laxa (Macbr.) Brand. Icdde. Rep
Spec. Nov. 24:60. 1927.

Perennial, 2.5-4.5 dm tall: stems branched from
the base and upward, stout, l-.^ dm long, strigose and
spreading setose: leaves linear, 4-12 cm long, 0.2-0.5
cm wide, dorsal surface coarsely strigose and pustu-
late, ventral surface finely strigt)se, and with a few
inct)nspicuous pustulate hairs, the petioles long ciliate
margined; inlltMescence very broad and open, lax,
0.7-2.5 dm long, the foliar bracts inconspicuous. Col-
lections: 13 (0); representative: L. C. Hinckley 3480
(GH); V. Harvard s.n. (US): E. L. Reed 3450 (US): E.
O.Wo..ton401 (NY)

liololype: Pringle 776. collected in Cliiluiaulia.
Mexico, on sand hills near Paso Del Norle. 20 Sep-
tember 1886. GH. Isotypes LL. ND-G.

Distribution: Southern New Mexico, western
Texas, and northern Mexico in the state ol' Chihua-
hua. Limited to sand dune areas. Map No. lb. June to
November.

This narrow-leaved variety is apparently confined
to the sandy, dune areas of Mexico and southern New
Mexico. It may be confused with vdi.jamesli in west-
ern Texas and southern New Mexico, but usually can
be separated from that variety by the stout, erect
stems, the narrower leaves, and the longer, more per-
fectly developed cymes. From variety mulliaiiilis il
differs in the steins, which are branched above the

base, and the very narrow leaves scattered along the
stems.

Ic. var. sclosa (Jones) Jnhiist. ex Tidestr.

CryphiiiiUa jiiiiusii (Ion.) Payson var. sclosa (Jones)
Johnst. ex Tidestr. Pioc. Biol. Soc. Wash. 48:42.
1^35.
Orcocarya cinerea Greene, Pitt. 3:113 1S96. (Type:

Southern Colorado, on the plains near Pueblo. 1S73.

(ireenc s.n.).
Krvnitzkia miillicaulis var. sclosa Jones, Conlr. West. Bot.

■ 13:4. 1910.

Orcocarva Icinmoni laslw. Bull, lorres Bot. Club
30:239. 1903. (Type: ,Ari/on:i, without definite local-
ity. 1884, Leninion.l

Orcocarva miillicaulis var. ciiicrca (Greene) Macbr Proc.
Am. Acad. ,'^1:54. 1916.

Orcocarva suflniiizosa \:u . cinerea (Greene) Pavson, Univ.
Wyo. Publ. Bot. 1:171. 1926.

Hcmisphacrocarva siiffriilicosa var. sclosa (.lones) lir:uid.
1 edde, Rep.'Spcc. Nov. 24:60. 1927.

Ilciiiisphacrocarva ciiicrca ((irccne) Brand. I edde. Rep.
Spec. Nov. 24:61. 1927.

Crvplaniha jomcsii (Torr.) Pavson var. cincrca (Greene)

■ Payson. Ann. Mo Hot. Clard. 14:246. 1927.

Perennial. 1-3 dm tall, branched from the base,
simple above; stems slender, 0.2-0.9 dm long, strigose
and weakly setose; leaves mostly basal, oblaiiceolate,
obtuse, 3.5-13 cm long, 0.4-1.5 cm wide, dorsal sur-
face finely strigose, usually conspicuously pustulate,
venlial surface uniformly and densely strigose, the
petioles conspicuously ciliate on the margins: inflor-
escence open, 0.4-2 dm long, bracts evident especially
near the base of the inflorescence. Collections: 163
(vii): representative: T. S. Brandegee B3I (NY, US):
E. L. Greene s.n. (ND-G): Rvdberg and Vreeland
5702 (NY. RM); D. T. Macdougal 204 (ARIZ, US):C.

Map No. lb. Parts of soiilhvveslcrn United States. Range ol
C jamcsii (Torr.) Payson: (Circles), var. selosa (Jones)
Johnst. e\ Tidestr.; (Triangles), var. laxa (Macbr.) Payson.

Map No. Ic. Parts of western L niled States Range of C
jamcsii (Torr.) Payson: (Circles), var. ahoniva (Greene)
l';i\son: (Triangles), var. pusliilosa (R\db.) ll;irnngl

BIOLOGICAL SFRUS VOL. I 3, NO. 4 A RLVISION Ol CKI I'l AN IIIA SUBGKNUS OKKOCARYA

17

F. B;ika 455 (GH. RM. I'S); L. C. Higgins lOOQ.
144;^(BR^ ).

Holotypc; M. E. Jones s.ii., collected near Fort
Cove. Utah, growing under jimipers. 27 June 1^)01.
RSA. Photograph at BRY.

Distribution: South-central Colorado, northern
New Mexico, northern Arizona, eastern Nevada, and
southern Utah. Usually I'oiuid on heavy clay soils.
Map No. lb. Late May to early September.

In southern Utah variety setosa reaches its best
development. It is characterized by the short stems
which never exceed the basal tuft of leaves. In the
outlying areas on the margins ot" its range, it freely
intergrades with variety miillici/iilis on the south,
aboriiva on the west, and variety jamesii on the east-
ern side. Oreocarya lemmoni was separated on the
basis of a more setose indument which occurs on a
population of this variety about Prescott, Arizona;
however, I do not believe that it is worthy of any
taxonomic rank. This variety has been recognized in
the past as variety cincrea. but the older name oi
sciosii must be used, at least at the variety level.

Id. \jL\.ab(>riirii (Greene) Payson

Cryptantha jamesii (Torr.) Payson var. ahoniva

(Greene) Payson. Ann. Mo. Bot. Gard. 14:250.

l')27.

Oreocarya ahoniva Greene, Pitt. 3:114. 1896.
Krvnilzkia nuilticaiilis var. ahoniva (Greene) .lones.

"Gontr. West. Bot. 13:5. 1910.
Orcocarva siiffruricosa var. ahoniva (Greene) Muebr.

Proc. Am. .Acad. 51:547. 1916.
Heinispliacrocarva ahoniva (Greene) Brand. I eddc. Rep.

Spec. Nov. 24:61. 1927.

Prostrate, caespitose perennials, branched from the
base. 0.7-2 dm tall; stems slender, weak, decumbent,
0.3-0.7 dm long, strigose, and with some weak spread-
ing setose hairs; leaves basal, as well as scattered along
the stem, linear to narrowly oblanceolate, 1.5-9 cm
long, 0.3-0.9 cm wide, dorsal surface finely strigose
and setose pustulate, ventral surface finely strigose
and without pustules, the petioles ciliate margined;
inflorescence open, 0.2-1 .3 dm long, floral bracts very
evident, usually exceeding the cymules. Collections:
44 (i); representative: Clokey 7280 (BRY. ND, NY,
ORE. UTC); Maguire and Holmgren 26119 (NY,
UTC); Parish 1480 (US); Purpus 6068 (US).

Lectotype: S. B. Parish 3694, collected in the San
Bernardino Moimtains, Bear Valley, 6,500 feet. 16-20
June 1895, ND-G. Isolectotypes at GH. UC.

Distribution: Central and southern Nevada, west
to the San Bernardino Mountains, north through Inyo
and Mono Ci)unties, California. Growing in sandy
soils from 6,000 to 10,500 feet. Map No. Ic. Late
May to October.

Variety ahortiva is fairly well defined geograph-
ically, and is only to be confused on its eastern
boundary with var. setosa. It may be separated from
that variety by the long tbliar bracts which exceed

the individual cymules and the prt)strate stems.

1 have designated the plant in the Notre Dame
Herbarium with Greene's handwritten notation Oreo-
carya abortiva to be the type specimen, as it agrees
with the plant and the maturity of the fruit which he
discusses in the original description.

1 e. var. dislivlia ( Eastw.) Payson

Cryptaiillia jamesii (Torr.) Payson var. Jisticlia
(Eastw.) Pavson. Ann. Mo. Bot. Gard. 14:248.
1927.

Orcocarva disticlia l-astw . Bull. Torre\ Bol. Club 30:238.
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 cili-
ate-margined; inflorescence open, 0.5-2 dm long, the
foliar bracts not conspicuous. Collections: 60 (iv);
representative: A. H. Holmgren 3243 (ARIZ. BRY,
UC, US. UTC); B. F. Hariison 10370 (BRY, UC); B.
Maguire 18298 (UC, UTC); Eastwood and Howell
6674 (UTC); L.C. Higgins 1004 (BRY).

Holotype: A. Eastwood 90. collected in San Juan
County. Utah, on Bartons Range. 13 July 1895, CAS.
Isotypes at UC, GH, US.

Distribution: Wayne and Emery Counties. Utah,
southeast through southwestern Colorado, north-
western New Mexico, northeastern Arizona in
Apache, Navajo, and Coconino Counties, north to
Garfield County, Utah. Usually found growing on
sand dunes or sandy slopes and ridges, 4.000 to 7,500
feet. Map No. Id. Late April to September.

The variety Jisticlia seems to be intermediate
between variety luulticaiilis and vdw jamesii. It can be
separated from the tbrmer by the more woody stems
which are branched above the base, from var. jamesii
by the erect stems, the leaves which are subglabres-
cent, and the more perfectly developed cymules. The
character of a single nutlet is of no value; as one to
four nutlets may be found on the same plant.

1 f. var. jamesii

Cryptantlia jamesii (Torr.) Payson vat. ju/ncsii.

Myosolis suffnaicosa Torr. Ann. Lye. N.Y. 2:225. 1827,

not Crvplanl/ia siiffruricosa Piper, Proe. Biol. Soc.

Wash. 32:42. 1919.
Eritrichiiim jamesii Torr. In Marey. E\pl. Red River ^6''

1854.
Krviiitzkia jamesii (Torr.) Grav. Proe. .Am, .Aeud. 20:278.

1885 in part.
Oreocarya siiffruricosa (Torr.) Greene, Pitt. 1:57. 1887.
Ilemispliaerocarva sufTrutlcosa (Torr.) Brand, I'edde,

Rep. Spec. Nov. 24:60. 1927.
Hemisphaerocarva siiffruricosa var. rvpica Brand. I edde.

Rep. Spec. Nov. 24:60. 1927.

Decumbent perennials, 1.6-4 dm tall, branched
from the base as well as above; stems decumbent.

18

HKK.IIAM VOllNC; HNIVl KSIIY SCIKNCK BUl.LKTIN

0.6-2 dm long, slrigose and weakly setose: leaves
linear to oblanceolate, 2.5-8 cm long, 0.3-0.9 cm
wide, the dorsal surface coarsely strigose and ap-
pressed setose pustulate, ventral surface uniformly
strigose and without pustules, tiie petioles ciliale-
margined; inflorescence open, 0.4-1.4 dm long, flora!
bracts evident but not conspicuous. Collections: 217
(iii); representative: P. A. Rydbeig 1514 (Gil, ND-G,
NY, US): A. Nelson 477 (GH, ND-G. NY, US);C. L.
Porter ,^951 (BRY. GH. RM): .1. H. Christ ')54 (CS,
GH):L. C.Higgins 1527(HRM.

llolotype: James s.n.. barren deserts high upon the
Platte, NY.

Distribution: Wyoming and .South D;ikol;i, south
through Nebraska, Kansas, and western Oklahoma,
west through northern Texas and western New
Mexico, and nortii througli Colorado east of the
Continental Divide. Growing on sandy to clay soils.
Map No. Id. May to late August.

Variety /«'"«■// is confined mainly to the area east
of the Continental Divide, but with some overlap in
central New Me.\ico and southern Colorado. In east-
ern New Mexico it may be confused with var. nndri-
caulis. and very often it is quite difficult to sepai;iie
the two. In south-central and southern Colorado var.
jamcsii may be confused with var. sclosa. but can us-
uallv be separated from it by the shorter leaves which
are scattered along the stem, rathci than in a basal
tuft.

Ig. var. ptisiidosii ('R\db.) Haningt.

CryptauiUa jamcsii (Ton.) l';iysoii vai. I'listiilosa

(Rydb.) Harnngt. Man. I'l. Colo. 4(i(i. (i41. 1^54.
Orcocarva i>iistulosu Rvdii. Bull. Tones Hot. (luli

40:4K(). 1913.
Crvptaniha piistulom (K\dli I P;ivson. Ann. .\1o Hot

■ Card. 14:252. 1927.
H cmispluicrocarva siilfniticosa var. puiHilosa lR\dli I

Brand, ioddc, Rcp.'Spcc. Nov. 24:60. 1927.

Erect perennials, 2-5. S dm t:ill. branched fioni the
base, simple above: stems slender, 1-3. 9 dm long,
glabrous or finely strigose: leaves linear to broadly
oblanceolate. 2-9 cm long. 0.4-1.5 cm wide, the
dorsal surface apprcssed setose-pustulate, ventral sur-
face glabrous, the petioles not ciliate margined: inflor-
escence open. 0.4-2 dm long, floral bracts inconspic-
uous. Collections: 22 (ii): representative: A. II.
Holmgren and S. Hansen 3489 (BRY, NY, UTC): 1'.
A. Rydberg and A. O. Garrett 9569 (NY, RM, UT);
A. Cronquisl and N. Holmgren 9372 (NY. UTC): J.
Reveal and G. Davidse 9:(i ( mi\). Welsh. Iliggms ;md
Atwood 8933(BR>').

llolotype: Rydberg and Garrett 9320, collected in
San Juan County. Utah, on the l-lk Mountains m
Hammond Canyon, 31 July 1911. NY

Distribution: Southeastern Utah in Ckirficld, Kane,
and San Juan Counties. Southwestern Coloiado,
northwestern New Mexico, and noiiheastern .-Xri/ona.

Cirowing HI ;i wide variety of soils, 4,500 to 8,500
feet. Map No. Ic. Late May to late August.

This variety is quite dilTerent in general appear-
ance 1'rom any of the other varieties in this species
comjilex. This is due to the slender weak sleius, and
the lack ot any pubescence on the ventral surface of
the leaves. The original description characterizes the
leaves as being glabrous beneath, sparingly hairy
above. However, with the specimens at hand it is the
upper and not the lower surface that is glabrous. The
var. disficha in some cases is nearly glabrous, biu
always has a few hairs on the ventral surface and is
probably a connecting link between var. serosa and
the present species.

2. Crypiautha pal/iicn (Gray) Payson

Crvpiantha palineh (Gray) Payson. Ann. Mo. Bol.
Card. 14:253. 1927.
Krynilzkia palnteri Gidy. Proc. Am. Acad. 20:27S. 1885.
Orcocarva palnnri (Gray) Grccni;. Pitt. 1:57. 1S87.
Ilciiiisphacrocarva palmeri (Grav) Brand. I lhUIc, Rep.

Spec. Nov. 24:61. 1927.
Crvplaiitlia ciiryi .lohnsl. .lourn, .Arn. .\xb. 2(1:396. 1939.

Plants biennial or short-lived perennials. 1 .7-4 dm
tall: stems I -several, 0.7-3.5 dm long, spreading setose
hirsute: leaves linear-lanceolate, acute. 3-16 cm long.
0.4-1 cm wide, strigose and subtomentose. pustulate
hairs conspicutuis on the dorsal surtace. few and not
evident on the ventral surface: inflorescence broad-
topped due to the elongation of the cymules in age,
0.3-2.7 dm long, setose, the floral biacts inconspic-
uous: calyx segments lanceolate, in anthesis 4-6 mm
long, in fruit becoming 7-10 mm long, setose or
weakly hispid: corolla white, the tube 4-6 nun long,
crests at base o\ tube lacking, fornices yellow,
rounded, papillose. 0.5- i mm long, limb 7-9 mni
wide: style exceeding mature fruit by 2-3.5 mm: nut-
lets ovate. 2.5-2.S mm long. 2-2.7 mm wide, the
margins not in contact, acute, both surfaces of the
nutlet smooth and glossy, scar tightly closed and
without an elevated margin. Collections; 1 10 (xii):
representative; D. S. Correll and I. M. Johnston
21243 (GH. LL); V. L. Cory 31517 (GH I: D. S. Cor-
rell 16333 (GH. LL); J. Reverchon 21 20 (GH. ND-G);
M. E.Jones 18514 (ND): E. J. I'almei 34009 (c;H); L.
C. lliggins 3097 (BRY).

llolotype; Palmer 895. collected in Coahuila.
Mexico. 40 miles south of Saltillo. March 1880. GH,

Distribution; Lower Sonoran life zone in western
Texas and adjacent Mexico. Growing on limestiuie or
gravelly to rocky hillsides. 1.000 to 4.000 leet. Map
No. 2. April to Late Jul\ .

The type of C. palmeri is very imniaUiie. and be-
c:iiise of this imnuiliiiiix some contusion luis come
about as lo which pkiiit should be;ii the mime ol
palmeri. In observing the type specimen on loan Irom
Gray Herbarium and the original description, which
characterizes the nutlets as follows: "■luiculis op;icis

BIOLOGICAL Sl-RIKS VOL. I J, NO. 4 .-X KI:V1S10N OL e'R Y I' L.XN IHA SUBGENUS ORKOCARYA

H

Map No. Id. Parts of southwestern United States. Range of
C. jamcsii (Torr.) Payson: (Circles), var. jainesii: (Triangles),
var. disticlia (I-astw.) Payson.

rugosiiisciilis."" the nutlets are subnigose only becuuse
the\' are iniiiiatiire. For this reason C. coryi is placed
in synonymy. Immature specimens in all the smooth-
fruited species have a tendency for the nutlets to
appear subrugose until they are fully matured, which
is no e.xceptiim in the present species.

This species may be separated from its nearest rela-
tive C. jamcsii var. nndlicaulis. by its lack of crests at
the base of the corolla tube, the accressent sepals, and
the longer style.

3. Crypttinilia uhlata (Jones) Payson

Crypiantlui olilaiu (Jones) Pavson, Ann. Mo. Bot.

Gaid. 14:254. 1M27.

Krvnitzkia ohiala M. E. Jones, Contr. West. Bot. 1.^:4.

1910.
Orcocarra luspidissima Wooton and Standlev, Contr. L!.

S. Natl. Herb. 19:545. 1915, not O liispiJissima

(Torr.) Rydb.
Oreocarva ohiala (.lones) Macbr Proc Am. Acad. 51 :54N.

1916.
Ih'inispliaeiDcarra ohiala (Jones) Brand, I edde. Rep.

Spec. Nov. 24:61. 1927.

Caespitose perennial, 1-3.5 dm tall; steins several,
0.4-1.5 dm long, retrorsely setose and spreading hir-
sute; leaves oblanceolale. acute, 3-10 cm long, 0.4-1 .4
cm wide, coarsely slrigose and appressed setose dor-
sally, pustules conspicuous, ventral surface weakly
strigose-setose, and with fewer pustulate hairs, the
petioles ciliate margined; inflorescence somewhat
open, especially in age. 0.3-2 dm long, setose-hirsute;
calyx segments linear-lanceolate. 5-7 mm long in
anthesis, becoming S-10 mm long in fruii. densely
setose; corolla white, tube 7-10 mm long, crests at
base of tube lacking, fornices yellow, broad, papil-
lose, limb S-12 mm wide; style 3-5 mm longer than

mature fruit; nutlets ovoid, usually all four maturing,
the margins narrowly separated, acute, 2.5-3 mm
long, 2-2.5 mm wide, dorsal surface rugose-tubercu-
late, ventral surface smooth or slightly uneven, scar
closed, straight, and without an elevated margin.
Collections: 76 (vi); representative; D. S. Correll and
I. M. Johnston 22036 (LL); C. L. and A. Lundell
14309 (GH, LL); G. R. Vasey s.n. (ND-G, US); W. P.
Cottam 1 0228 ( BRY. UT); G. C. Nealley 1 67 (ND-G,
US); E. O. Wooton s.n. (ARIZ. RM. US).

Holotype: M. E. Jones 3759, collected at El Paso,
Texas, 23 April 1884, POM. Photograph at BRY. Iso-
typesal RM, US.

Distribution: Southcentral New Mexico, western
Texas and northern Mexico. Growing on sandy or
gravelly limestone soil, 1,000 to 5,000 feet. Map No.
3, Late March to September.

This species is confined mainly to the western half
of transpecos Texas and southern New Mexico. It
may be distinguished from its nearest relatives, C.
palmcri and C. paysonii on the basis of the tloral and
nutlet characters. From the former it may be distin-
guished by the nutlets which are roughened on the
dorsal side and the corolla tube which definitely
exceeds the calyx segments. It differs from the latter
by the flowers which are not heterostyled. and the
stamens which are always attached at the middle of
the corolla tube, the fornices which are nearly gla-
brous, and the more open loosely thyrsoid inflor-
escence.

4. Cryptantha paysiuiii (Macbr.) Johnst.

Oypniiirlui paysonii (Macbr.) Johnst. Wrightia 2:160.

1961.

Oreocarva pavsonii Macbr. Contr. Grav Herb. 4.S:36.

1916.
Hcmisphacrocarva pavsonii (Macbr.) Brand. I edde. Rep.

Spec. Nov. 24:61. 1927.

Caespitose perennials. 1.6-2.9 dm tall; stems erect,
stout, 0.8-1.6 dm long, strigiise and more or less
spreading setose-hirsute; leaves oblanceolate, obtuse
to acute, 3-9 cm long, 0.5-1.5 cm broad, dorsal sur-
face finely strigose or subtomentose, also setose with
pustulate hairs, ventral surface similar but with less
conspicuous pustulate hairs; inflorescence subcap-
itate, consisting of four to six compact cymules,
0.5-1 .2 dm long, setose; calyx segments linear-lanceo-
late, in anthesis 7-9 nim long, in fruit becoming 9-10
mm long, setose-hirsute; corolla white, the tube 12-14
mm long, crests at base of tube lacking, fornices
yellow, rounded, densely papillose, 0.5-1 mm long,
limb 1(.)-13 mm wide; plants heterostyled; nutlets
ovate. 2.7-3 mm long, 2-2.5 mm wide, usually all four
nutlets maturing, margins nariowly winged, in con-
tact, both surfaces finely rugulose or finely tubercu-
late. scar closed, straight, and lacking an elevated
margin. Collections: 6 (iii); representative: O. B.
Metcalfe 1576 (POM. US); D. S. Correll and I. M.

20

BKKJHAM YOUNG UNIVERSITY SCIKNC'K BULl 1 UN

Map No. 2. Western Texas and northern Mexico. Range of C
palmcri (Gray) I'ayson,

Johnston ::003 (LL); L. C. Higgins 3151 (BRY,
WTSU).

Holotype: O. B. Mctciilfe I 576. collccled in Sierin
County. New MckIl-o, on linie.stone hills at Betendo
Creek. 12 May 1%5.GH. isotypesat POM. US.

Distribution: New Mexico in Debaca. Otero, and
Sierra Counties, south into Culberson County, Texas.
Growing on limestone soil. 4,000 to 7,500 feet. Map
No. 4. April to June.

This species has been contused in the past with C.
ohlata. but differs from it in the larger corollas which
are strongly helerostyled. the more compact inflor-
escence, and the nutlets which are only finely rugu-
lose or tuberculate.

More collections of this laxon arc badly needed in
order to determine its exact geographical range.

5. Cryphintha crassipcs Johnst.

Cryptantha crassipcs Johnsi. Joum. Am. Arb.

20:.107. 143^).

Plants perennial. 1.5-2.4 dm tali; stems 1 -several.
1.2-1.'-.' dm long, setose; leaves linear-lanceolate,
obtuse to acute, mostly basal, reduced upward. 3-6
cm long, 0.2-0.6 cm wide, densely while strigose. also
setose-pustulate on the dorsal surface: inflorescence
capitate. 0.2-0.8 dm long, white strigose. floral bracts
inconspicuous; calyx segments linear-lanceolate, in
anthesis 7-9 mm long, in fruit becoming 9-1 1 mm
long, setose: corolla white, the tube 8-9 mm long,
crests at base of lube lacking, fornices yellow.

Map No. 3. Parts of Texas and New Me.xico. Range of C.
ohlala (Jones) Payson.

rounded, about 1 mm long, limb 9-1 1 min wide: style
exceeding mature fruit 4-7 mm; nutlets ovate or tri-
angular-ovate. 3.3-3.8 mm long. 2.5-3 mm wide,
dorsal surface finely rugulose. ventral surface smooth
or only slightly uneven, margin acute or narrowly
winged, scar closed, and without an elevated margin.
Collections: 15 (iii): representative: V. L. Cory 18613
(GH): D. S. Correll and I. M. Johnston 21934 (GH.
LL); D. S. Correll and R. C. Rollins 23604 (LL); V.
L. Cory 31 585 (GH); L. C. Higgins 2767. 2940 (BRY.
WTSU).

Holotype: V. L. Cory 18613. collected in Brewster
County. Texas. 6.5 miles east of Agua Fria Springs.
13 April 1936. GH. Photograph at BRY.

DistribiUion: Brewster Coimty. Texas, m the Big
Bend region. Growing on white limestone which is
shaley or clayey. 1.500 to 4.500 feet. Map No. 5.
Lale March to early June.

This distinctive plant is closely related to C. pal-
mcri and C. paysonii. From the former it can be dis-
tinguished by the compact capitate inflorescence, the
thicker more woody caudex. and the shorter corolla
lube. From C. paysonii. it differs in the shorter cor-
olla tube, the monomorphic flowers, the nutlets
which are more compressed or flattened, and the very
woody caudcx.

6. Cryphinilhi coiifcnijlora (Greene) Payson

Cryptantha ccnlcrti flora (Greene) Payson. Ann. Mo.
Bot.Gard. 14:256. 1927.

IJIOLOC.ICAL SKRiES VOL. I 3. NO. 4 A REVI.SION Ol CRVITAN IHA SUBGENUS ORtXJCARYA

21

Krvniizkia leucophaca var. alala Jones. Proc. Calif. .'\cad.
'Sfi. II. 5:710. 1S95. (Type; Silver Reef. Utah, on

sandstone clil'fs. 4.500 feet. 3 Ma\ 1894. M. i:. .lones

5144.)
Orcocarya coiijcrlijlorii Greene. Pitt. 3:112. 1896.
Oreocarva lulea Greene. Muhlenberttia 2:240. 1906.

Name only. 1-cddc. Rep. Spec. Nov. 19:72. 1923.

Description. (Tvpe: California. Invo Coimt\. While

Mountanis, 9 May 1906. Heller 821 i.)
Oreocarva alala (Jones) A. Nels. Coulter and Nelson. Man.

Cent. Roekv Mts. 417. 1909; Rvdb. 11. Roekv Mts.

725. 1917.

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 oblanceolate, 3-12 cm
long 0.2-1.6 cm wide, acute, dorsal surface densely
strigose and appressed setose with pustulate bases,
ventral surface unitormly strigose and with tew or no
pustules; intlorescence subcapitate. 0.3-2 dtn long,
strigose and with twisted setose hairs, bracts incon-
spicuous; calyx segments linear-lanceolate, in anthesis
6-8 mm long, in fruit becoming 10-14 mtn long, stri-
gose and spreading setose; corolla yellow, the tube
9-13 mm long, fornices broad, emarginate, about I
mm long, crests at base of tube evident or sometimes
lacking, limb 8-10 mm wide; plants distinctly hetero-
styled; nutlets triangular or ovate, 3.5-4 mm long.
2.5-3 mm wide, ustially all four maturing, margins
narrowly winged, in contact, surfaces smooth and
glossy, scar straight, closed, and lacking an elevated
margin. Collections: 117 (vi); representative: I. W.
Clokev 7659 (ARIZ, LL, ND, ORE, UTC); S. B. and
W. F. Parish 1316 (ND-G, US); F. W. Gould 1550
(BRY, VJ. UTC); P. Train s.n. (ARIZ, ORE); L. C.

Higgins 1365, 1475. 1771 (BRY).

Holotype: S. B. Parish 1316, collected in San
Bernardino County. California, at Cushenberry
Springs on the north side of the San Bernaidino
Mountains, 1882. ND-G. Photograph at BRY. Isotype
at US.

Distribtition: Southwestern Utah, northern Ari-
zona, southern Nevada, and west to southern and
western California. Growing in a wide variety of soils,
2,000 to 8,500 feet. Map No. 6. April to July.

This distinctive, yellow-tlowered species is one of
the largest of the perennial species of Cryplantha. and
is not common, bitt covers a wide distributional
range. This species may be distinguished from its
closest relative, C. flava. by the broader more ovate
nutlets and the longer stems with a subcapitate inflor-
escence.

Brand legally published the nomen nudum Oreo-
carva lulea Greene. He did this t)n the basis that the
stamens were in a different position in the coiolla
tube. However, this taxonomic criteria is of no value
in the present species because the corolla is very
dimorphic in regards to this character.

Jones described a narrow-leaved form from south-
ern Utah as a new variety, but it is not worthy of any
subspecific rank.

7. Cryptaiiilia Jhra (A. Nels.) Payson

Crvptantha Jlava (A. Nels.) Payson, Ann. Mo. Bot.
Gard. 14:259. 1927.

®

, 1

Map No. 4. Northwestern Texas and southern New Mexico.
Range of (' paysoiiii (Machride) JolinsiiMi.

Map No. 5. Brewster Co., Texas. Range of C. crassipes
Johnston.

-I-)

liRK.HAM YOIJNC. UNIVKKSITY SCIF.NCK BULLKTIN

Oreocarya Jlara A. Ncls. Bull. Torrcv Hot Club 25:202.

1898.
Orcocarva liilcsceiis Greene, Pitt. 4:9.^. 1899. (Type: On

hills about Aztee. New Mexico, 25 April 1899, C. 1 .

Baker.)
Crvpianlha confertiflora var. tlava Brand. Ptlan^enreich

(Hel't. 97) 4. fam. 252:90. 19.'!1.
Crvpianlha confertiflora var. hilesccns Brand. Hlkin/en-
■ reidi(Helt.97)4, lam. 252:90. 19.M.

Perennial, 1.3-4 dm tall; stems many, tVoni a mul-
tiple caude.x, 0.8-2.6 dm long, densely long white-
hairy at the base, becoming setose and strigose up-
ward; leaves narrowly oblanceolate to nearly linear,
acute,- 2-9 cm long, 0.3-0.8 cm wide, dorsal surface
strigose and appressed setose with pustulate hairs.
ventral surface almost uiiilorinly strigose, and with
the pustules less conspicuous; nitlorescencc narrow lo
siimcwluit open, 0.5-2..^ dm long, conspicuously
yellow setose, the tloral bracts inconspicuous; calyx
segments linear, in anthesis 8-10 mm long, in Iruit
becoming ^-1 2 mm long, densely setose, with yellow-
ish hairs, pedicels 3-5 mm long in fruit; corolla
yellow, the tube 9-1 2 mm long, crests at base of lube
absent or nearly so. Cornices yellow, truncate, emar-
ginate, 1-1.5 mm long, limb 8-10 mm broad; style
exceeding mature fruit 3-7 mm (heterostyled); nutlets
lanceolate. 3.4-4 mm long. 1.9-2.2 mm wide, 1-2
usually maturing, margins acute, in contact when
more than I nutlet matures, both surfaces of nutlet
smooth and glossy, scar straight, closed, elevated
margin lacking. Collections: 193 (xj); representative:

C. V. Baker 562 (ND-G); A. and R. Nelson 785
(ARIZ, ORE); W. A. Weber 383S (ARIZ. COLO); W.
P. Cottam 2050 (BR^); B. Maguire 18302 (UTC); L.
C. Higgins 527, 998, 1057, 1073. 1082. 1 105 (BRY).

Holotype: A. Nelson 3074. collected In Sweet-
water County, Wyoming at Point of Rocks. 1 .lune
1897, RM.

Dislribulion: Sinilhein Wyoming, south through
western Colorado and eastern Utah to northern New
Mexico and Arizona. Usually found growing in sandy
soil. 4,000 to 7.500 feet. Map No. 7. April to August.

This yellow-tlowered species is closely related to
C. confcnifhira. but may be separated from that
species by the narrowly lanceolate nutlets and ilic
longer thyrsoid intlorescence.

8. Cryptaiuha aipitata ( tastw.) Johnst.

Cryptantlia capitula (Eastw.) Johnst. Journ. .\\n.

Arb. 21:66. 1940.

Oreocarya capitala lastw. Leaflets West. Bol. 1:9. 1937.

Perennial, 1.5-2.7 dm tall; stems weak, 1 -several,
1.2-2.4 dm long, appressed setose; leaves linear, or
very narrowly oblanceolate, 3-8 cm long, 0.3-0.5 cm
wide, dorsal surface appressed setose-pustulate, ven-
tral surface uniformly strigose and without pustules;
inllorescence capitate, or with one or two glomerules
below the terminal cluster. 0.1-0.4 dm long, spreading
white-setose; calyx segments linear-lanceolate, 7-9

Map No. 6. Paris of western United .Slates. Raiiiie of C
conferlijlora (Greene) Payson

Map No. 7. Parts of western L'nited States. Rani;e ol'C ilava
(A. Nels.) Payson.

BIOLOGICAL SERltS VOL. I J, NO. 4 A REVISION Ol CK Y I' 1 ANTHA SU BGENUS OREOCAR Y A

23

mm long in anthesis, in fruit becoming 11-16 mm
long, conspicuously setose-pustulate; corolla white,
the tube *)-! 2 mm long, crests at base of tube conspic-
uous, fornices yellow, eniarginate, about I mm long,
papillose, limb d-cS nim wide; style exceeding mature
I'ruit 4-5 mm; nutlets lanceolate. 4-5 mm long. 2-3
mm wide, two to four usually maturing, the margins
in contact, knifelike, both surfaces glossy-smooth,
scar closed, straight, and without an elevated margin.
Collections: 1 1 (0); representative: A. Eastwood and
J. T. Howell 1005 (CAS, GH); A. Eastwood 5S.^2.
596^) (CAS, GH); L. White s.n. (MNA); R. E. Collom
ki20(ASC);F. W. Pennell 21575 (ARIZ).

Lectotype: A. Eastwood 5'560. collected in Coco-
nino County, Arizona, from Hermit Trail on the
south rim of the Grand Canyon, 4 April I'll?, CAS.
Pholograpli at BRY. Isolectotype at GH.

Distribution: Southcentral Utah, and northcentral
Arizona in the Colorado River drainage basin. Grow-
ing in sandy soil, 6,500 to 8,500 teet. Map No. 8.
April to July.

C. aipitata is probably most closely related to C.
confertijicni. but differs from that species in the
more capitate inflorescence, the narrower leaves,
white flowers, with crests at the base of the tube, and
usually smaller size.

Two collections by Cronquist and N. Holmgren
9299 and 9365. appear to be the same species, but
the leaves are broader. These two collections came
from the Henrv Mountains and Aquarius Plateau in
Ltah.

9. Cryptauilui iohnsionil Higgins

Oyptaitllia johnstotiil Higgins. Great Basin Naturalist

28:195. 1968.

Caespitose perennial 1-2.5 dm tall; stems several,
arising from the branched caude.x, 0.6-1.3 dm long,
very weakly strigose; leaves oblanceolate, the apices
obtuse to acute. 2-6.5 cm long. 0.4-1 cm wide, dorsal
surface strigose with conspicuous pustulate hairs;
inflorescence somewhat open, 0.5-2 dm long; foliar
bracts evident but not conspicuous, 1-2 cm long;
calyx segments 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; co-
rolla white, the tube 12-15 mm long, flaring in the
throat, crests at base of tube lacking, fornices yellow,
1-1 .5 mm long, emarginate, papillose, limb 13-17 mm
broad; style exceeding mature fruit 3-8 mm (hetero-
styled); nutlets ovate, 3-3.5 mm long. 2.3-2.7 mm
wide, usually all four maturing, the margins acute or
knifelike, in contact, both surfaces smooth and
glossy, scar straight, closed, elevated margin lacking.
Collections: 5 (iii); representative: L. C. Higgins 1310
(BRY); B. F. Harrison 5628 (BRY).

Holotype: L. C. Higgins 1310, collected in Emery
County, Llfah, on low rolling hills about 15 miles
west of hwy. 50-6 along the road from Woodside to
Castle Dale'. 25 May 1968. BRY. isotypes at CAS,
GH.NY. POM. RM.'US, UTC.

Distribution: Known only from the type locality
15 miles west of hwy. 50-6 on the San Rafael Swell,

Map No. 8. Southern Utah and northern Arizona. Range of Map. No. 9. Emery (ountv, L tah. Range of C. /oliitstonii
C capitala (I:ast\v.) Johnston. Higgins.

24

BRIC.HAM YnUNG UNIVKRSITY SCIENCE BULLEl'lN

Emery County, Utah. Growing itn clayey lo sandy
soils, 5.000 to 5,500 teei. Map No. 9. May and .lime.
Cryptantha johiistoiiii is most closely related lo C
confcnijlora known from western Utah, northern
Arizona, Nevada, and southwestern Calitornia. It can
be distinguished from that species by its smaller size,
longer and more open intlorescence, white llower
color, larger corolla with longer fornices and no basal
crest.

10. Crypiaiillia scntiglahni Barncby

Cryptaiuha semiglahra Barneby. Leatlels West, Bot.

3:1^7. 1943.

Erect perennials, 2-3 dm tall: stems 1 -several.
0.9-1.8 dm long, retrorsely strigose and weakly
spreading setose; leaves oblanceolate, acute, 3-7 cm
long, 0.3-0.6 cm wide, dorsal surface appressed
setose-pustulate, ventral surface glabrous, the old leaf
bases long white-hairy; intlorescence narrow, or some-
what open, 0.4-1.3 dm long, foliar bracts slightly
surpassing the cymes, 1.5-2 cm long; calyx segments
lanceolate, in anthesis 5-X mm long, in fruit becoming
10-13 mm long, setose; pedicels 1-2 mm long; corolla
white, the tube 10-1 2 mm long, crests at base of lube
conspicuous, fornices yellow, rounded, 1-1.2 mm
long, obscurely papillose, limb 8-10 mm wide; style
surpassing the mature fruit 5-7 nun; nutlets ovate,
3.5-4 mm long. 2-2.5 mm wide, usually all four
matming, margins acute, in contact, both surfaces
smooth and glossy, scar closed, elevated margin lack-
ing. Collections; 5 (ii); representative; J. W. Harrison
s.n. (DIX); H. D. Ripley and R. C. Barneby 8519
(UTC); D. Atwood 1525 (BRY); L. C. Higgins 1357,
I364(BRY).

Lectotype: Ripley and Barneby 8429. collected in
Coconino County, Arizona, on detrital clay hills
about 2 miles east of l-'rcdoma, 4.900 feet. 5 .lune
1942, CAS.

Distribution; Apparently confined lo near llie
type locality and north just across the stale line into
Washington and perhaps Kane County, Uiah. Grow-
ing in clay soils, 4,500 to 5.000 feet. Map No. 10.
Early May to July.

Cryptaiuha scinlglabra is closely related to C
capitata and C. confertijlora. From the tormci it
differs in several important aspects, ihc most immed-
iate difference being the distribution of pubescence.
The upper part of the caude.x branches are densely
clothed with long white hairs, the lower surface of
the leaves are beset with stout appressed setose hairs
with pustular bases, while the upper surface is gla-
brous. The inllorescence of a long narrow thyrsus also
distinguishes it from C. capitata. From the latter ii
may be distinguished by the white flowers, the gla-
brous ventral surface of the leaves, crests at the base
of the corolla tube, and the longer foliar bracts.

I 1 . Crypidiiilki hanichvi .lohnsl.

Crypianilta hunichvi .loliiisl. .Iiunn. .'\rn. Arb.

29;240. 1948.

Perennial. 1.5-3.5 dm tall; stems stout, erect, sev-
eral. 0.8-1 .2 dm long, conspicuously yellowish hispid;
leaves oblanceolate. thick, acute. 5-9 cm long. 0.5-1 .4
cm wide, coarsely appressed hispid pustulate on both
surfaces, and with some flner hairs beneath, the peti-
oles conspicuously ciliate; inflorescence narrow. I-1 .5
dm long, densely yellowish hispid, foliar bracts evi-
dent to conspicuous; caly.x 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 8-1 I lum wide; style
exceeding mature fruit 5-6 mm; nutlets ovate. 3.5-4
mm long. 2.5-3 mm wide, all Inur maiming, margins
of nutlets in contact, acute, smooth and glossy on
both surfaces, scar closed, straight, and without an
elevated margin. Collections; 10 (viii); representative;
Riplev and Barneby 8748 (GH); D. Atwood 1562
(BRY); L. C. Higgins 1584, 1587, 1599, 1601 (BRY).

Holotype; Ripley and Barneby 8748, collected in
Uintah County, Utah, 30 miles south of Ouray on
white shale knolls, 5,500 feel. 17 June 1947. GH.
Photograph at BRY.

Distribution: Confined lo the lower part of the
Uintah Basin, Uintah County, Utah. Growing on
white barren shale knolls, 5,000 to 6,000 feel. Map
No. 1 1 . May to June.

This endemic species is confined to white shaley
knolls, and is the only conspicuous plant on them. In
the western part of its range it is t'ound growing with
C. grahaiuii, bul still limited to while shale.

In the original description of this species ihc
corolla was described as being long tubed and pro-
truding beyond Ihe sepals. In observing this plant in
the field the corollas do not exceed the calyx, or if
they do. it is by less than 1 mm.

C. hanichyi is a very distinctive species, and is
probably most closely related to C. confertijlora or C.
jolinstoiiii. but may be distinguished from both of
Ihem by the thick stout stems. camp;iiiiihiic coiolla.
harsher hispid pubescence, and I he more woody
caudex.

I 2. Crypiaiului Iciicoi'liaca ( Dongl.) Payson

Cryptiinllia Icucophaca (Dougl.) Payson, .■\im. Mo.

Bot. Gard. 14:262. 1927.

MyoKoiis Icinopliaca Doug). In l.chm. Pug. 2:22. 1830.

Eniriihinm Ictaophacuin (Doiml.) .\. DC. Prod. 10:129.

1846,
Krvnitzkia Iciicoplwea (Dons;!.) CJrav. Proc. .Am. Acjil.

'20:280. 188.S.
Orcocarya Icucopliacci (Dougl. I tlrci'iic. Pitt. 1:58. 1SS7.

Long-lived perennials. 1.5-3.7 dm tall: stems

Bl

OLOCMCAL SKRIKS VOL. 1 3. NO. 4 .\ RIVKSION OF C KYIT.XN IHA SUBGENUS OREOCAKYA

25

.Map No. 10.
C. semiglahra

Southern
Barneby.

Utah and northern Arizona. Range of

Map. No. I I .
.lohnston.

Uintah County. Utah. Range of C. banicbx

slendei', 1 -several fiom a tmilliple caiidex. 1.2-2.3 dm
long, strigose, and appicssed setose, leaves linear to
narrowly oblanceolate, acute, 3-9 cm long, 0.4-0.7 cm
wide, dorsal surface densely strigose, and with
appressed setose hairs, pustulate, ventral surface
uniformly strigose and with few or no pustulate hairs,
petioles white-ciliate; intlorescencc narrow. 0.<S-I.7
dm long, conspicuously white setose, foliar bracts
evident but not conspicuous; calyx segments linear, in
anthesis 6-8 mm long, in fruit becoming 10-15 mm
long, setose; corolla white, the tube 8-10 mm long,
crests at base of tube evident, fornices yellow, emar-
ginate, 0.5-1 mm long, limb 8-10 mm wide; style
exceeding mature fruit 2-8 mm (heterostyled); nutlets
ovate, 3.5-4.5 mm long, 2.5-3 mm wide, usually less
than four maturing, margins acute, in contact, both
surfaces smooth and glossy, scar stiaight, closed, ele-
vated margin lacking. Collections: 16 (0); repiesenta-
tive: J. H. Sandberg and J. B. Leibeig 373 (RM. US);
J. W. Thompson I 1453 (US. WTU); T. S. Brandegee
W7 (US); L. Hitchcock 20952 (RM); A. Elmer 1056
(US);J. S.Cotton 1027 (US).

Type: Douglas s.n.. collected in Washington State,
on arid barrens of the Columbia River. Not seen.

Distribution: Upper Sonoraii Zone in south-central
Washington along the Columbia River and its north-
ern and southern tributaries. Growing in sandy soil.
Map No. 1 2. Late April to early July.

This species of south-central Washington is most
closely related to C. confertijlom. but differs from
this species by the white corolla, the open and more
elongated intlorescence, and the evident crests at the
base of the corolla tube.

13. CiyptaiUha sahnonensis (Nels. & Macbr.) Payson

Cryptaiitha sahnonensis (Nels. & Macbr.) Payson.
Ann. Mo. Bot. Card. 14:263. 1927.
Oreocarva saliuoiiciisis Nels. & Maebr. Bol. Gaz 61 43
1916.

Moderately caespitose perennials, 1.5-3.5 dm tall;
stems erect, 0.7-1.5 dm long, strigose, and with some
while spreading setose haiis; leaves oblanceolate to
spatulate, 2-9 cm long, 0.5-1.5 cm wide, strigose and
spreading setose, also somewhat tomentose, pustulate
on both of the surfaces; inflorescence narrow, 1-1.8
dm long, foliar bracts evident but not conspicuous;
calyx segments lanceolate. 3-4 mm long in anthesis, in
fruit becoming 6-8 mm long, setose; corolla white,
the tube 3-4 mm long, fornices yellow, rounded,
papillose, crests at base of tube well developed, limb
7-10 mm wide; nutlets lanceolate, 3-4 mm long, 1 .5-2
mm wide, all four usually maturing, acute or nar-
rowly winged-margined, the margins in contact,
smooth and glossy on both surfaces, scar straight,
closed, and without an elevated margin. Collections:
10 (v); representative: Macbride and Payson 3348
(RM); Hitchcock and Muhlick 8950 (UTC); A.
Cionquist 3812 (UTC); E. B. Payson 1880 (RM); L.

26

liKK.llAM YOUNC, UNIVI RSIIV SCIENCI- BULLKMN

Map No. 12. Central and soutlK-rn Wasliington. Range ol C
leuiophaea (Dougl.) Pay son.

C. liiggiiis 1710, 1711, 1713. 1714. 1715 (BRY).

Holotype: Kirtley s.n.. collected in Lemhi County.
Idaho, a'l Salmon, June 1890. RM. Photograph at

BR^.

Distribution: Along the .Salmon River in Lemhi
and Custer Counties, Idaho. Growing on loose talus
slopes of volcanic origin. 4.500 to 7.500 feet. Map
No. 13. June to August.

Crvptcmiha sahnonensis is appaienily confined to
the Salmon River drainage of south-central Idaho. It
IS perhaps most closely related to C. lencopluicu. but
differs from that taxon by the short corollas, more
lanceolate nutlets, shorter style, and shorter intlor-
escence.

14. Crypiuniha siricta (Osterh.) Payson

Boi.

CrvptaiUha stricta (Osterh.) Payson, Ann. M'
'Card. 14:264. 1027.

Orcocarva slricla Osterli. liull, Iorre\ Bol. ( lub 50;21/.
192.V

Strict perennial. 1-3.7 dm talk stems 1 -several.
0.4-2 dm long, strigose and conspicuously setose-
hirsute; leaves mostly basal, oblanceolate to spatulate.
acute. 2-7 cm long. 0.4-0.9 cm wide, retrorsely stri-
gose and spreading setose-hirsute, pustulate: inllor-
escence narrow, interrupted below the terminal

Map No. 1 3- Central Idaho. Range of C sahnonensis (Ncls. &
Maehr.) Pawon.

cluster. 0.5-2 dm long. setose4iirsute. tloral bracts
inconspicuous; calyx segments lanceolate. 4-h miu
long in anthesis. in fruit becoming 7-9 mm long,
setose-hirsute; corolla white, the tube 3-4 mm long,
crests at base of tube conspicuous, fornices yellow.
rounded, papillose, limb 7-10 mm wide: style exceed-
mu mature fruit 1-1.5 mm: nutlets lanceolate to el-
liptic. 3-3.5 mm long. 1.5-2 mm wide, usually all four
mauinng. margins in contact, knifelike. dorsal surlace
with definite transverse ridges, also somewhat tuber-
culate. ventral surface smooth or nearly so. scar open.
very narrowly linear, elevated margin lacking. Collec-
tions: 17 (hi): representative: E. H. Ciraham 8163
(GH): L. Williams 489 (GH); R. C. Bameby 9145
(GH): Welsh and Moore 6714 (BRY): W. J. MacLeod
10a (COLO. CS); G. E. Osteihout 6.^91 (RM): L. C.
Higgins 1869. 1874 (BRY. TTC. WTSU).

Holotype: G. E. Osterhout 61 95. collected in
Moffat County. Colorado, some distance south of the
Yampa or Bear River along the Victory hwy.. 21 June
1922, GH. Photograph at BRY. Isotype at RM.

Distribution: Southwestern Wyoming in Carbon
County, south into Moftat County. Colorado, and
west to I'lah in I'intah. Daggett, and Summit
Counties. Growing on clay soils. 5,000 lo S.500 teel.
Map No. 14. June to September.

Crvpuintlhi siricia is an endemic species, confined
lo the thiec corners area o\ Colorado. Wyoming, and

lilOlOf.UAL SI KIIS VOL. I -i, N(1. 4 A KI'VISION OL IK\ I' I AN IHA SUBGENUS ORKOCARYA

27

Utah. In genei;il iippearaiice il sumcvviuit resembles C.
celosioitlcs. but is probably not very closely related to
that plant, it may be distinguished from other species
in the I'intah Basin by the strict stems with harsh
setose liairs, the nutlets which are smooth on the
ventral surface, and the conspicuous transverse ridges
on the dorsal surface.

15. Cryptaiiilhi luihigcna (Greene) Payson

Crvpiantlia iiuhigcnu (Greene) Payson. Ann. Mo. Bot.

Card. 14:265. 1927.

Orcocarya mibigena Greene. Pitt. 3; 1 1 2. 1 896.
Crvpiaiilha clcniciisac Pav.son. ."^nn. Mo. Bot. Gard.
■ 14:267. 1927. (Tvpe: Glenn's Pass. California. Mrs.
.loscph Clemens, 22 July 1910. RM.)

Short-lived perennials. O.H-2.5 dm tall; stems sev-
eral from a slender taproot. 0.4-1 dm long, setose;
leaves narrowly oblanceolate, tlaccid, obtuse to acute.
2-5 cm long, 0.3-0.7 cm wide, strigose and spreading
setose, dorsal surface pustidate. ventral surface with
tew or no pustules; intlorescence narrow, cylindrical
or nearly capitate. 0.4-2 dm long, foliar bracts incon-
spicuous; calyx segments linear-lanceolate, in anthesis
2.5-3 mm long, in fnut becoming 4-6 mm long,
setose; corolla white, the tube 2-2.5 mm long, crests
at base of tube evident, fornices light-yellow to nearly
white, rounded, about 0.5 mm long, slightly papil-
lose, limb 3.5-5 mm wide; style exceeding mature
fruit 0.5-1 mm; nutlets narri)wly lanceolate, papery,
2.8-3.2 mm long, 1.3-1.5 mm wide, margin narrowly
winged, in contact, dorsal surface weakly tuberculate
to nearly smooth, ventral surface smooth, scar open,
narrowly linear, margin of scar not elevated. Collec-
tions: 32 (0); representative: Chestnut and Drew s.n.
(ND-G); G. T. Robbins 339'-) (RM. WTU); P. A. Mun/
12547 (POM. WTL!); J. T. Howell 25933 (POM.
UTC); F. W. Pierson 14030 (POM); Alexander and
Kellogg 455 2 (UTC).

Lectotype: Chestnut & Drew s.n.. collected in
Mariposa County, California, on Clouds Rest,
Yosemite Natl. Park. 10 .luly 1889. ND-G. Photo-
graph at BRY. Isolectotype at CALIF.

Distribution: East-central California in Inyo,
Tulare, Fresno, Mono, and Mariposa Counties. Grow-
ing in gravelly soil or talus slopes of volcanic origin.
9,000 to 13,000 teet. Map No. 15. July to Sep-
tember.

This species is endemic to the high Sierras, chielh
between 9.000 and 12,500 feet. The specimens of
this species available to past monographers has been
very poor and scanty. Payson saw a poor isotype of
the species and mistakenly identified it with some
plants of Oregon and Idaho. The reasimably good
specimens of this plant t>om the southern Sierras
Payson described as a new species C. clenicnsac. This
latter name however falls in synonymy of C inihigcim
and the plants of Oregon and Idaho mistakenly called
nuhigcna were described as new (Jolinston 1939),

N
J ®

* 1 /

1 — ■ — ^

1

/ * /

Map. No. 14. Northeastern L'tah and ailjoining Colorado and
Wvoniine. Raniie ofC slricta (Osterhoutl Pa\son.

Map No. 15. Parts of easlcenlral California. Range of C.
mibigcna (Greene) Payson,

HKIC.llAM YOUNCi UNIVIKSIIV SCIKNCK lUH.I 1 UN

Tlic Oregon plant is now known as C. subretiisa. and
ihe Idaho plant as C hypsopliila. C. mibigeiia may be
separated from the latter two by being a weaker.
more slender, more bristly plant with less firm, green,
basal leaves, much smaller, smoother nutlets, and a
more interrupted inllorescencc with a capitate ter-
minal cluster, and scattered smaller lateral ones
below.

16. Crypiaiiilui suhrcima Johnsl.

Crypianllui subretiisa Johnsl. Joum. Am. Aib.

20:3^)3. 193^.

Cryi>laiillia amliiia John.sl. e\ M. I. I'eck. Man. PI. Ore.

601. 1941. (Without latin diagnosis or type.)
OreocaiYa siihreliisa (Jolinst.) Abrams. 111. I'l. Pac. St.

.^:599. 1951.

Caespitose perennial. 1-2 dm i:ill; stems several.
0.5-1.5 dm long, setose; leaves spatulate. subretuse or
obtuse at ape.x, congested at the base, reduced
upward, 1-4 cm long, 0.4-1 cm wide, tomentose and
weakly setose, pustules conspicuous on the dorsal
surface, fewer and less evident on the ventral surface;
inflorescence compact, cylindric, 0.2-1 dm long,
spreading setose, floral bracts inconspicuous; calyx
segments lanceolate, 3-4 mm long in anthesis. in liuil
becoming 6-9 tnm long, setose and subtomentose;
corolla white, the tube 3-4 mm long, crests at base of
tube conspicuous, fornices yellow, rounded, papil-
lose, about 0.5 mm long, limb 4-6 mm wide; si\ie
exceeding mature fruit 0.5-1 mm; nutlets lanceolate.
3-4 mm long, 2-2.5 mm wide, 2-4 maturing, the
margins in contact, knifclike or narrowly winged,
dorsal surface inconspicuously tuberculale, and with
low short ridges, ventral surface similar, but the mark-
ings much less evident, or smooth, scar open, subulate
or narrowly linear, and without an elevated margin.
Collections: 25 (0); representative: J. W. Thomspon
12206 (Gil, POM, WTU); R. L. Rogers 87 (ORH);W.
H. Baker 6282 (WTU); W. C. Cusick 2028 (ND-G),
RM); C. G, Hansen 534 (GH, ORE); G. Mason 7502
(ORi:);M. H.Peck 19480 (GH).

Iloloiypc: J. W. Thompson 12206, collected in
Klamath County, Oregon, on pumice rim of Crater
Lake, 7,000 feet, 20 July 1935. GH. Photograph at
BRY. Isotypes at POM . WTU.

Distribution: Northeastern Oregon in Wallowa
County, south to Harney County and to Humboli
County, Nevada. West to northern California and
north into Klamath County, Oregon. Growing on
pumice or talus slopes of volcanic origin, 6,500 to
10,000 feel. Map. No. 16. June to August.

Generally C. siihrelusa may be distinguished from
the other plants or Oregon by its elongate nutlets,
tt)mentulose, thickish. obtuse, truncate or subretuse
basal leaves, and the small corollas.

The accmnit of C. niibigena in Payson's mono-
graph applies almost entirely to this species. His
description, as well as the nutlets belong to C. siihre-
lusa.

17. Crypiantlui hypsopliila Johnsl.

Crvptaiitha hvpsupliila Johnst. Joum. Am. Arb.

20:395. 1939.

Caespitose perennials. 1-2 dm lall; stems several,
0.5-1.4 dm long, setose-hispid; leaves oblanceolale-
spatulate, obtuse, 2-6 cm long, 0.3-0.7 cm wide,
strigose and spreading setose, dorsal surface evidently
pustulate, ventral surface with fewer pustulae; inflor-
escence narrow, 0.2-0.7 dm long, foliar bracts incon-
spicuous; calyx segments lanceolate, 3-4 mm long in
anthesis. in fruit becoming 5-7 mm long, setose-
hirsule, corolla white, the lube 3-4 mm long, crests at
base of tube well developed, fornices yellow,
rounded, limb 4-5 mm wide; style exceeding mature
fruit 1 .2-1 .5 mm; nutlets oblong-lanceolate, 3-3.7 mm
long, 1.5-1.8 mm wide, margins acute, in contact.
di)rsal surface tuberculate or some of these connected
to form short low ridges, ventral surface smooth or
nearly so, scar narrowly linear, the margin not ele-
vated. Collections: II (0); representative: J. W.
Thompson 14129 (GH, WTU); A. Cronquist 2894
(GH, IDS); C. L. Hitchcock and C. V. Muhlick 10676
(WTU); Macbride and Payson 3771 (POM).

Holotype: J. W. Thompson 14129. collected in
Blaine County, Idaho, on crest of high barren ridge at
head of Boulder Creek, Sawtooth Mountains. I 1,000
feet, 6 August 1937, GH. Photograph at BRY. Iso-
typeal WTU.

Disiriliulion: Endemic to central Idaho in Blaine
and Custer Counties. Growing on talus slopes and
soils of volcanic origin. 8,000 to 11,500 feel. Map
No. 17. July and August.

Payson identified this plant as part of C. niihlgciia.
Its atTmities. however, are not with thai plant but
with C. siibrcliisa of Oregon. The Idaho plant can be
distinguished by its spreading biistly hairs, narrower
nutlets, and narrower less firm leaves that are obtuse
or acute at the apex.

This species may also be confused with C. spicii-
lifcra. but ditTers in being a smaller and more caespi-
tose plant, with more elongate nutlets, and shorter
basal leaves. It is also separated from this plant in
elevalimi and flowering time.

18. Crypiantlui ciyinopliilii Johnsl.

Crypiaiillia crriiioplilla Johnsl. Joum. Am. Arb.

21:65. 1940.

Oicocarva cryinoplula (Johnsl.) Jeiis. & lloo\or, in
Jepson I-l. Calif. 3:.'!28. 1943.

Perennial, 1.5-3 dm tall; stems 1 -several, 0.9-1 .3

dm long, erect, hirsute; leaves oblanceolate, 4-10 cm

long, 0.5-1.3 cm wide, finely setose and appressed

hirsute, the dorsal setae pustulate at base, the ventral

with fewer puslules; inflorescence narrow, 1.9-2.3 dm

long, selose; calyx segments lanceolate. 4-5 mm long

in anihesis. in fruii becoming 10-14 mm long, hirsute;

corolla white, the lube 3-5 mm long, crests al base of

BIOLOGICAL SKRIES VOL. I 3, NO. 4 A Kl VISION Ol CRVl'IAN Il'A SUIiGKNUS OHIlOCARYA

29

Map No.
Johnston.

16. Western Linited States. Range of C siihirriisa

/'\

@

/ "- \/ 1

'*\--l

*•

; -.__L_.

Map No.
Range of

17. Central Idaho. Blaine
C. hypsophila Johnston.

and Custer Counties.

Iiibe evident, tomices yellow, rounded, papillose, 0.5
mill long, limb 4-7 mm wide; style exceeding mature
tViiil 1-2 mm; nutlets ovoid, usually four maturing,
5-(i mm king, 3-3.5 mm wide, the margins in contact,
winged, dorsal surface with low ridges, also incon-
spicuously niuricate, to nearly smooth, ventral sur-
face smooth, scar open, linear, and without an ele-
vated margin. Collections: 4(ii); representative: R.
F. Hoover 4193 (GH); C. B. Hardham 65 10 (POM);
L.C.Higgins 1766, 1767 (BRY).

Holotype: R. F. Hoover 4193, collected in Alpine
County, California, on Red Peak. 28 July 1939, GH.
Photograph at BRY.

Distribtition: Alpine ridges between the Clark
Fork and the Middle Fork of the Stanislaus River in
Alpine and Tuolumne Counties, California. Growing
in loose rocks of volcanic origin. '^).000 to 10.000
feet. Map No. 18. Late June to September.

C. crymophila is probably closely related to C.
mibigcna of the southern Sierras. However, it ditt'ers
from C. mibigena by being much taller and more
robust, with longer leaves, much larger tYuiting
calyces, and the larger more rugose nutlets which are
definitely wiiiged-mar.ained.

I'). Cryptaniha setdsissiniu (Gray) Payson

Oyptciiifha sc'iosissinm (Gray) Payson. Ann. Mo. Bot.

Gard. 14:268. 1927.

Erilrichium sctosissima Gra\, Proc. .\m. Aead. 12:80.

1877.
Krvnilzkia scloslssiina (Gray) Grav. Proc. Am. Aead.

20:276. 1885.
Oreocarya selosissiina (Gray) Greene. Pitt. 1 :58. 1887.

Biennial or short-lived perennials. 3-10 dm tall.

stems tisually 1-3 erect. 1.5-5 dm long, hirsute; leaves

clustered at the base, reduced upward, oblanceolate.

the apices obtuse to acute, 3-13 cm long. 0.5-1.5 cm

wide, setose, with some finer twisted pubescence

beneath, pustulate hairs numerous on both surfaces;

inflorescence broad-topped due to the elongation of

the scorpioid racemes, 1-5 dm long; caly.x segments

broadly lanceolate, 4-6 mm long in anthesis, in fruit

becoming ''-I I mm long, setose, and strigose; corolla

white, the tube 3-5 mm long, constricted above the

ovary by the conspicuous ring of crests, fornices

yellow, emarginate, 0.5 mm long, limb 7-'-) mm wide;

style exceeding mature fruit 1-2 mm; nutlets ovate,

5-6 mm long, 3.5-4.5 mm wide, papery, with a

broad-winged margin, dorsal sinface niuricate. and

inconspicuously rugose or tubercidale. ventral surface

smooth or nearly so, scar straight, narrow, slightly

open, elevated margin lacking. Collections: 58 (viii);

representative: Maguire and Holmgren 25583 (BRY,

ORE. RM, UTC); L. F. Ward 646 (UC); R. H. Peebles

12566 (ARIZ);D. T. MacDougal 165 (ARIZ. RM);C.

F. Deaver 6306 (ASC); W. D. Stanton 516 (UT); E.

Palmer 591 (US); L. C. Higgins 1125. ll!7. 1440,

1775, 1795 (BRY).

30

HKK.H AM VOUNC. UNI VI'.RSITY SCIKNCF BULIKIIN

N

/

®

?

^^ \

^ ■' \

\ /f ^V

\ ^1 > \

I -' ^ \

l^—~—'- J • \ \

1 ,,'-->----,'#./ \

\ ^' / ' \

\^ / ^ \

\ / ^^

\ /

\ ' 1

^ / V

\ / ^.

/'-.

\ ) -' \

\^' ■' \

.-^\

Map No. 18. t'ast central Caliloinia. Alpine and Tuolumne
Counties, and adjoining Nevada. Range of C cryuuipliihi
Johnston.

Ht)li)type: L. F. Ward (-i46. collected in Sevier
Counly, Utah, at Fish Lake, 25 August 1875. GH.
Photograph at BRY. Isotypcs at UC. MO, PH. US.

Distribution: Central LUali. siuitli through most of
Arizona in the mountainous areas, west \.o Nye
County, Nevada. Growing in gravelly to sandy soils.
7,000 to 10,000 ft. Map No. 10. .lane to September.

This is one of the most distinctive species in the
entire genus, it may be separated from all other
species by the stout, strict, solitary stems, and the
broadly winged ovate nutlets.

20. Ciyptamha yinj,uia (i'ortei ) Payson

CrvphiniUa vir^aui (i'orter) Pavson. .Ann. Mo. Bot.
Gard. 14:270. 1427,

liritrichiiim virgalum I'orter, ll,i\den R. pi. 479. IS7U.
lirilrichhim f^lomcraniin var. yirjialiiin I'orler, \n I'orter &

Coulter, Syn. II. Colo. 102. 1874.
Krviiilzkia virs;a!ci (I'orter) Cira\. Proc. Am. .-Neail

20:279. 1885.
Oreocarya viifiulu (Porter) Creene, Pitt. I :58, 1887.
Oreocaiva spicala Rvdli. Bull, lorrev Hot. Cluli .i(i:(i78.

1909, (Tvpc: Artists Glen, I'ikes Peak, Colorado, I

August 1901, Clements 102.)

Oreocarya vir^ata I'orma spicalu (R\dlil -NLiebr. Proi. .
Ain, Acad, 51:546. 1916.

Strict biennial, arising frotii a sti)ut taproot, 2.5-8

(10) dm tall: sletns usually solitary, but sometimes

several from the base, stout, 0.5-2 dm lotm, setose or

Map. No. 19. Utah, .Arizona, and southe.islcrn Nevada.
Range of C sctosissima (Gray) Payson.

hirsute-hispid; leaves narrow!) oblaiiceolate, obtuse,
.^-20 cm long, 0.4-1,5 cm wide, setose-hirsute, with
pustuhite hairs on both surfaces: innorescence cylin-
drical. 1.5-7 C")) dm long, with conspicuous, liiiear-
oblanceolate foliar bracts that much exceed the
cymules: calyx segments lanceolate, in anthesis ,v5-4
mm long, in fruit becoming 10-12 mm long, hirsute;
corolla white, the tube 3.5-4 mm long, crests at base
of tube very conspicuous, fornices yellow, emar-
ginate. papillose, about 0.5 mm long, limb 8-i 1 mm
wide; style surpassing the mature fruit 1.6-2 mm;
nutlets ovate. 2.7-3.5 mm long. 2.4-3 mm wide. u,s-
ually all four maturing, the margins iti cont;ict. acute,
dorsal surface usually with conspicuous low ridges,
and a few tubei'cles. or sometimes ncaiK smooth,
ventral surface smooth or with a few indistinct
tubercles, scar narrowly open, linear, and without an
elevated margin. Collections: 62 (vi); repiescntative:
B. Maguire 16292 (UTC); A. Nelson 1437 (ND-G,
RM); G. T. Robbins 3358 (ARIZ); H. B. Payson 4253
(RM); J. Fwan I4')47 (COLO); A. Nelson 1267 (UC.
RM);L.C.Higgins 14m. 1501. 1543 (BRY),

Type: B. H. Smith s.n,. collected near Denver,
Colorado. Colorado Territory. 1864. Not seen.

Dist I ihutioii : Southeasterti Wyoming, south
through central Colorado, on foothills on the eastern
side of the Rocky Mountains. Growing on gravelly
soils. 5.000 to 4.500 feet. Map No. 20. Farly May to

DIOl OCllCAL SFRIF.S VOL. I 3. NO. 4 A Rl VISION Ol CRYl'T ANTKA SUBGENUS OREOCARYA

31

Seplombor.

C. virgata is one of the most conspicuous her-
baceous piiiiits on liie eastern tootiiills ol" the Roci<y
Mountains. Tlie stems of this plant are usually soli-
tary, with long leafy foliar bracts that greatly exceed
the individual cymules. For a few years this plant was
treated as a variety of C cclosioides. but there is no
reason to believe that it is even remotely related to
that species.

Rydberg described Oreocarya spicata as a new
species on the basis of the smooth nutlets. Later it
was transferred as a form of O. virgutu. Macbride
contended that O. spiaUa was not deserving, even of
varietal rank. In obseiving specimens from the type
locality about Pikes Peak, it is even more evident that
sptaiia is only a form of virgata.

21 . Cryptaiitha unuulnsa (Payson) Payson

Crvpitinilhi nmmldsii (Payson) Passon, Ann. Mo. Bot.
Card. 14:276. l'-)27.

Oreocarva ninnilosa Pavson, Univ. VVvo. Piilil. Bot. 1 :164.
1926.

Long-lived perennials. 0.8-2.^' dm tall; stems
1-several from a woody taproot. 0.4-1.4 dm long,
spreading setose; leaves oblanceolate. obtuse, 3-5 cm
long, 0.4-0.4 cm wide, strigose. setose and tomentose.
pustulate on both surfaces, but more conspicuous on
the dorsal side; intlorescence narrow, cylindric. unin-
terrupted. 0.4-2 dm long, tloral bracts not evident;
calyx segments linear-lanceolate. 4-5 mm long in
anthesis. in fruit becoming 7-10 mm long, yellowish
setose; corolla white, the tube 3.5-4.5 mm long, crests
at base of tube evident, fornices yellow, acute, papil-
lose. 0.5-1 mm long, limb 6-8 mm wide; style exceed-
ing mature truit 0.5-1 mm; nutlets ovate. 3-4 nnn
long. 2.5-3 mm wide, one to three usually matming.
the margins in contact when more than one matures,
acute, dorsal surface with a low inconspicuous crest,
tuberculate. and with some low ridges, ventral surface
similar, scar open, triangular, margin of scar slightly
elevated. Collections; 40 (0); representative; I. W.
Clokev 7667 (ARIZ. UC. LL. ORE. RM, UTC); T. S.
Brandegee s.n. (UC); P. A. Munz 14787 (BRY, GH);
R. S. Ferris 11265 (RM); E. K. Ball 1*^)346 (POM);
Alexander and Kellogg 1463 (UC).

Holotype; T. S. Brandegee s.n.. collected in .San
Bernardino County. California, on the Providence
Mountains. May 1902. UC. Photograph at BRY.

Distribution; Clark County. Nevada, m the
Charleston Mountains, southwest to the Providence
Mountains. California, north to Inyo County. Grow-
ing on gravelly to clayey soils. 5,000 to 10.200 feet.
Map No. 2 1 . March to July.

This species has been confused in the past with C
liuniilis and C. nuhigena. The nutlets are quite dif-
ferent from either of those species. Its nearest rela-
tives are probably C. virginensis, C. insolita and C.

1 1

1 1

1 1

N

1 1 1
1 ' 1

j

1 1 ,
1 1 1

@

1

1 • -• ..

V

'*■ •

■-•-^•l-n

Map No. 20. Soiitlieastern Wyoming and central Colorado.
Range of C virgata (Porter! Payson.

Map No. 21. Southern Nevada and adjoining Calil'ornia.
Range of C. tiimtilosa (Pay.son) Payson.

32

HKIGHAM YOUNG IJNIVIKSI lY Sl'lKNtK BULLETIN

ahaia. From C. ahala it differs in liie lack of a con-
spicuous elevated margin around the scar and by the
indelniite roughenings on tiie dorsal surface. It differs
from C. virgiiieiisis and C. iiisolita in its definite
perennial habit, narrow congested intloresccnce. more
lomentosc indument, and the smoother nutlets with
only a slightly elevated margin around the scar.

22. Crypfaiiflia iiisoUta (Macbr.) Payson

Crvptaiulia iiisolira (Macbr.) Payson, Ann. Mi). Bot.

'Card. 14:273. 1927.
Oreocarya iiisolila Macbr. Contr. Gray Herb. 48:28. 1916.

Biennial or shoit-livcd perennial from a slender
taproot. 3-4 dm tall; stems 1-sevcral. 1-3 dm long,
strigose and conspicuously setose: leaves spaliilatc.
mostly basal, obtuse, 3-5 cm long, 0.5-1.4 cm wide.
dorsal surface subtomcntose and sparsely appressed
setose pustulate, ventral surface similar but the setae
smaller and fewer, pustules few and small, petioles
long-hairy at the base: intlorescence open, cymes few,
much elongating, 0.7-1.4 dm long, weakly setose,
foliar bracts inconspicuous: calyx segments linear-
lanceolate, in anthesis 3.5-4.5 mm long, in fruit
becoming 7-9 mm long, densely hirsute: corolla
white, the tube 3-4 mm long, crests at base of lube
well developed, fornices yellow, slightly emarginate.
papillose. 0.5-1 nun long, limb (i-<S mm wide: style
exceeding matiue fruit i-1.5 mm: nutlets ovate to
lanceolate. 3.7-4 mm long, one to four maturing, the
margins acute, in contact or nearly so, dorsal surface
carinate, luberculate. granulo-muricate and some-
times slightly rugulose. ventral surface tuberculatc
and somewhat rugulose. scar narrow but open, the
margin showing some tendency to become elevated.
Collections: 2 (0): representative: L. N. Gooddiiig
2286(GH.RM).

Hok)type: L. N. Goodding 2286. collecied m
Clark County. Nevada, at L.as Vegas, 4 May 1905.
GH. Photograph at BRY. Isotype at RM.

Distribution: Known only from the region about
Las Vegas. Growing in white alkaline soil. 1 .900 to
2,500 feet. Map No. 22. April to June.

This species somewhat resembles C data ot east-
ern Utah and western Colorado, but it is doubtful if
the two are even remotely related. However, this
plant is probably closely related toC vlri;iiiciisis, but
differs in the length of the tloral bracts, number of
cymules, and the shape of the nutlets.

23. Crypnuilha rliwliicnsis (Jones) Payson

Crvpuniiha virgincnsis (Jones) Payson, .^nn. Mo. Bot.

Card. 14:274. 1927.

Krvnilzkia ^lonicrala var. viixincmis Jones, (onlr. West

Bol. \i:i. 1910.
Oreocarva vir,^iiu'iisis (.loiies) Macbr. Proc. Am. Acad.

51:547. 1916.

Biennial, 1.5-3.5 (4) dm tall: stems 1-several, aris-

ing from a stout taproot, 0.3-0.{i dm long, setose-
hirsute; leaves oblanceolate to spatidate. obtuse. 3-10
( 12) cm long, 0.5-1 .5 cm wide, dorsal surface sparsely
setose, pustulate, also with some fine tangled hair
beneath, ventral surface subtomentose and weakly
appressed setose, with only a few pustulate hairs;
inllorescence a broad thyrsus with the individual
cymes much elongating. 0.5-3 dm long, foliar bracts
conspicuous; calyx segments linear-lanceolate, in
anthesis 3-4 mm long, in fruit becoming 7-1 1 mm
long, hirsute: corolla white, the tube 3-4 mm long,
crests at the base of tube conspicuous, fornices
yellow, emarginate. papillose, about 1 mm long, limb
7-9 mm broad; style exceeding matiue fruit 1-1.5
mm: nutlets ovate, 3.3-4.5 mm long, 2.4-2.6 iimi
wide, usually only one to two nutlets maturing, mar-
gins in contact, acute, dorsal surface with a distinct
ridge, the surface tubercuhile and usually rugulose,
ventral surface very uneven with indeterminate rugae
and tubercles, scar open, and triangular, with an
elevated margin. Collections: 64 (ii); representative: I.
W. Clokey 5820 (ARIZ, BRY, ORE, RM, UTC): B.
Maguire 4470 (RM. LUC); I-. W.Gould 1580 (ARIZ,
BRY. DIX. RM): Maguire and Holmgren 25404
(ARIZ. BRY, UTC): Alexander and Kellogg 3019
(RM. UTC); J. Beailey 4275 (BR^. LA): L. C.
Higgins 1243 (BRY).

Holotype: M. E. Jones 5195a. collected in Wash-
ington County. Utah, at Lavcikin. 8 May 1894, POM.
Isotypesat UC, RM, MO, US.

Distribution: Southwestern Utah, southern
Nevada, northwestern Arizona, and southeastern
California. Growing on gravelly to clay soils. 2.000 to
8,000 feet. Map No. 23. March to July.

Cryptiiiului yirgiiu'iisis has its closest relatives with
C tiiiuulosa and C. Iiofjiiuiiiiiii. From the former it
differs in the biennial liabii. more open intlorescence,
more setose indument. and the nutlets which are
more conspicuously roughened.

There can be no di)ubl that C. rirgiiicnsis and C.
liolJ)iiannii are very closely related; and, because of
the variation encountered in each, it is difHcult to
find characters of a high order which are consistently
differential. Variation as now known in C. virgincnsls
is rather great, especially marked in size of nutlets,
length of the calyx, and the markings on the nutlets.
However, where they approach each other in range,
they occupy different life zones, and C. rirglncnsis
conies into tlower a month or more earlier. The
former also has fragrant tlowers while C lioflnniniiil
does not. .■\dditional collections ot' this complex are
badly needed fiom weslein Nevad;i and c:islern
California.

24. CrypiaiuliLi UdfjDianiui Johnst.

Crypianilui liDjI'niaiiiiii Johnst. Contr. Am. Arb.
3:90. 1932.

BIOLOGICAL SERIES VOL. 1 3, NO. 4 .'\ REVISION Ol' CKYl'TAN I KA SUBOENUS OREOCARY A

33

Map. No. 22. Southern Nevada. Range of C insolila (Macbr.)
Payson.

Map No. 23. Soulliern Nevada and parts of adjoining state
Range of C virgineiisis (M. I',. Jones) Payson.

Oreocarva hoffmannii (.lohnst.) .Abranis. Abrams. III. II.
Pae.St. 3:600. 1951.

Plants bientiial. 1.7-3.4 dm tall; stems l-several,
0.2-1. 6 dm long, conspicuously hirsute; leaves spatu-
late. crowded at the base, reduced upward. 2-5 cm
long. 0.5-1 .2 cm wide, spreading setose-hirsute, pustu-
late on both leaf surfaces, but more conspicuous
dorsally; inflorescence broad-topped, interrupted,
1-2.8 dm long, floral bracts evident but not conspic-
uous; calyx segments lanceolate, in anthesis 3-5 mm
long, in fruit becoming 5-8 mm long, hirsute-hispid;
corolla white, the tube 3-4 mm long, crests at base of
tube evident, fornices yellow, rounded, 0.5 mm long,
papillose, limb 5-7 mm wide; style e.xceeding mature
fruit 0.2-0.8 mm; nutlets ovate, 3-3.5 mm long, 2-2.5
mm wide, two to four nutlets maturing, the margins
in contact, acute, both surfaces irregularly low rugose
and minutely tuberculate. the dorsal with a low
inconspicuous crest, scar open, triangular, without an
elevated margin. Collections: 10 (i); representative:
R. Hoffmann 78 (GH); Alexander and Kellogg 2503
(ARIZ, POM, RM); F. W. Pierson 7544 (GH. POM);
P. Train 3977 (UTC); J. Roos 5849 (POM).

Holotype: R. Hoffmann 78. collected in Inyo
County, California, on rocky open slopes of Westgard
Pass. 7,300 feet, II July 1930, GH. Photograph at
BRY.

Distribution: Western Nevada and southeastern
California. Growing on gravelly soil in the pinyon-
juniper community. 7.000 to 9.000 feet. Map No. 24.
June and July.

Johnston, in the original description, said that this
species was most closely related to C. insolita. how-
ever; I believe that its closest relative is C. virgiiiensis.
as discussed imder that taxon.

25. Cryptantha tilniiti Johnst.

Crvptautha ahala Johnst. Journ. Am. .4rb. 24:240.

'l948.
Krvnitzkia depressa .lones. Contr. West. Bol. 13:5. 1910,

' not C depressa A. Nels. Bot. Gaz. 34:29. 1902.
Oreocarva depressa (Jones) Maebr. Contr. (Iras Herb.

48:32. 1916.
Crvptautha modesta Pavson. .Ann. Mo. Bot. Gard.
14:278. 1927. not C modesta Brand, 1 edde. Rep.
Spec. Nov. 24:48. 1924.
Plants perennial, arising from a strong woody
taproot. 0.5-1.8 dm tall; stems many, 0.2-1.5 dm
long, strigose and weakly setose, leaves oblanceolate
to spatulate, obtuse, strigose, setose, and subtomen-
tose. the petioles ciliate margined; inflorescence
narrow, 0.2-0.8 dm long; calyx segments lanceolate to
ovate. 2.5-4 mm long in anthesis. in fruit becoming
5-8 mm long, setose; corolla white, the tube 3-4 mm
long, crests at base of tube conspicuous, fornices yel-
low, rounded, papillose, about 0.5 mm long, limb 7-8
mm wide; style exceeding mature fruit 0.5-1 mm;
nutlets ovate, 3-3.5 mm long. 2-2.5 mm wide, usually
all four maturing, margins in contact, obtuse to acute,
dorsal surface carinate, tuberculate, nunicate. and

34

HKKillAM VOIINC; LINI Vi: KSl TV SCIKNCt BULLETIN

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Map No.
Range of

24. Inyo Con
C. Iiojfinaniui

ntv
J oh

California, anil adjoining Nevada.
nslon.

soinolimcs with low liicunspicLunis ridges, ventral
surface deeply and irregularly rugose, scar open, tri-
angular, surrounded by a slightly elevated margin.
Collections: 28 (v); representative: M. E. Jones 6692
(POM, UTC): B. F. Harrison 0000 (BRY);W. S. Boyle
1 1 17 (BRY. UTC): L. N. Gooddmg W(-,(P()M, RM);
Eastwood and Howell 651 (CAS); L. C. Higgins 1015.
1016 (BRY).

Lectotype: M. H. Jones 66'-)2. collected at Auriuii.
Nevada. 7 JOO feet, 20 June I.S');^, POM. Photograph
at BRY. Isotypeat L'S.

Distribution: South-central (itah. northwestern
Arizoira. and eastern Nevada. Ciriiwing on sandy or
gravelly soil. 4.000 to O.OOO feet. Map No. 25. April
to early July.

The name I'ayson applied to this species of south-
ern Utah and eastern Nevada was invalid because of
an earlier honionyni. Johnston. luiting this, named
the plant t'. ahalii.

Cryptaiiilia ahahi is a modest, densely tufted
Crvptaiithu. thai is not greatly different in general
appearance fioni some of the varieties of C. huniilis
or of C lunnilosa. It differs from these species by the
short intlorescence. the deeply rugose nutlets on the
ventral surlace. and the spatulate leaves.

This plant is not often collected because of the
early flowering time.

26. Crvpiaiulia cacspiiosa (A. Nels.) Payson

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Map. No. 2
of C. abala

5. Soutliwoslcrn lltali and adjolnlni; slates. Range
Johnston.

Crvptiinrlu/ nicspitosu (A, Nels.) Pas'son. .Ann. Mo.
Bot.Gaid. 14:281. 1427.

Oreocarva cacspiiosa A. NeK. I r>lhej 7:(i5. 1X94.

Densely caespitose pereiinials. 0.5-1.5 dm tall;
stems 1-niany. arising finiii :i much-branched woody
caude.K. 0.2-0.0 dm long, weakly setose, and ap-
pressed strigose; leaves oblanceolate to spatulate, 1-3
cm long, 0.3-0.7 cm wide, pubescence of two kinds,
strigose and appressed setose, becoming tomentulose
toward the petiole; intlorescence narrow, 0.3-1 dm
long, toliur bracts inconspicuous; caly.x segments
lanceolate, in anthesis 3-4 mm long, in tVuit becoming
5-cS irim long, strigose and weakly setose, also some-
what tomentulose; corolla white, the tube 3-4 mm
long, crests at base of tube conspicuous, fornices
yellow, rounded, about 0.5 mm long, limb 4-7 mm
witle: style equallying or 0.5 mm longei than mature
tiuit; nutlets lanceolate. 3-3.5 mm long. 2-2.5 mm
wide, the margins acute, m contact, dorsal surlace
with low iiuinded rugae, also tubeiculate. and with
numerous muncations between ihe ndges. ventral
surlace nuiric;itc. scai open, iiarriiwly triangular,
nuirgin ot' scai not elevated. Collections: 40 (vi);
representative: .'\. Nelson 4671 (CS. RM): b. Nelson
4407 (RM);C,. I-. Osierhout 6248 (RM I: I.. B. Payson
4240 (RM); K ( Rollins 1085 (RM. UTC); D.
Atwood 156.S (BR^). L. C. Higgins 1557. 1562.
1563. 1566. 1586, 1570(BR^ ).

BIOIOC.KAI SIKIFS VOL. 13. NO. 4 .X RFVISION OI CR Y l> TAN TFA SU HGKNIIS OREOCAR Y A

l.cctolxpc: A. Nelson 474^^ collected in Svveot-
vvalei Ceuinly. Wyoming, at Point of Rocks, 15 June
1 848, RM. Piiotogiiiph at BRY. Isotype at US.

Distribution: Southern Wyoming, but to be
expected in northern Colorado and L'tah, and perhaps
eastern Idaho. Growing on heavy clay soils, 5,000 to
7,500 feet. Map No. 2(^. Early May to late July.

From the original description it is evident that
Nelson also included in this species the plants that he
later described as C. vana. In C. caiui the leaves are
silky-strigose, the intlorescence more capitate, and
the millets are shai pl\ muricate.

27. Cryptiiniha oclirolaica Higgins

Crvptantlia ochrdlciiai Uiggins, Great Basin Natur-
alist 28:1^7. |4{i8.

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 oblanceolate. the apices
acute or sometimes obtuse, 1-2.5 cm long, 0.1-0.,> cm
wide, basal leaves uniformly and densely strigose,
sparsely setose, the petiole 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 {?•) mm long in
anthesis. in fruit 4-6 mm long, setose; corolla pale-
yellow, the tube 2-2.5 mm long, crests at base of tube
conspicuous, tornices yellow, rounded, about 0..^ mm
long, limb 4-5 mm wide; style scarcely surpassing
mature fruit; nutlets lanceolate, 2.5-3 mm long.
1.4-1.0 mm wide, usually only one maturing, margin
acute, dorsal surface irregularly rugose with low,
rounded ridges; ventral surface only slightly uneven,
scar open, narrowly triangular, extending U the
length of nutlet, no elevated margin. Collections: 5
(i); representative: L. C. Higgins 1 788 (BRY); Reveal
and Reveal 1031 (BRY); D. Atwood 1891 (BRY,
WTSU).

Holotype: L. C. Higgins 1788. collected in Gar-
field County. Utah, on outcrop 100 meters south of
Red Canyon Campground along hwy. 12. 6.500 feet,
21 July 1968. BRY. Isotypes at GH. NY, US.

Distribution: Limited to the red Wasatch Forma-
tion near Red Canyon Campground in southwestern
Garfield County. Utah. 6.500 to 7.000 leet. Map No.
27. May to August.

Crvptaiuha dclirolciica is apparently most closely
related to C. cacspitosa of southwestern Wyoming,
but also has some affinities with C. Iiumilis. it difters
from C. cacspitosa by its less caespitose habit, the
slender, less woody taproot, shorter calyx, shorter,
pale yellow instead of white corolla, and the smaller
nutlets which are more rugose. From C. huniilis. C.
ochralcuca differs in the shorter calyx, pale yellow
corolla, and the rugose nutlets.

This local species is apparently confined to the red
Wasatch Formation in southwestern Garfield County,
Utah.

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Map No. 26. Soutliorn Wyoming.
Ncls.) Payson.

Range of C cacspitosa (A.

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Map No. 27.
Higgins.

Garfield County, LUah. Range of C, ocltrolcuca

36

BRIGIIAM YOUNG UNIVKRSITY SCIENCK BULLETIN

28. Cryptantlia humilis (A. Gray) Payson

Perennials, more or less densely caespitose. 0.5-3
dm tail: stems many, arising from the ends of the
branciied caudex, 0.2-1.5 dm long, strigose to spread-
ing setose; leaves oblanceolate to spatulate, 1-6 cm
long, 0.2-1.2 cm wide, strigose, setose or sublo-
mentose, pustulate on both surfaces; innorescciice
narrowly cylindrical to open and lax, 0.2-1. 8 dm
long, tomentose to conspicuously setose; calyx
segments linear-lanceolate, in anthesis 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,
fornices yellow, more or less papillose, rounded,
about 0.5 mm long, limb 7-10 nun wide; the style
shorter than to exceeding the mature fruit by 2.5
mm; nutlets lanceolate to ovate-lanceolate, 3-4.5 mm
long, 1 .8-3.2 mm wide, one to four of them maturing,
margins in contact, acute to obtuse, dorsal surface
muricate. tuberculate. or somewhat rugulose. ventral
surface indistinctly muricate or tuberculate, scar
open, triangular, margin not elevated.

Key to the varieties of C. hiiinilis

1. Leaves strigose and setose but not conspicuously

tomentose; calyx conspicuously setose (2).
1 . Leaves densely strigose as well as tomentose; calyx
setose and subtomentose (4).
2. Nutlets rugulose as well as muricate; style

1 .5-2.5 nun longer than mature fruit

28a. var. Iiuniilis

2. Nutlets nuiricatc or tuberculate; style not

exceeding the fruit by more than 1.5 nun (3).

3. Style exceeding the mature nutlets 1-1.5 nun; the

inflorescence open and broad; plants loosely tufted

28b. var. comini.xni

3. Style not or only slightly surpassing the nutlets;
inflorescence congested, even in fruil; plants

densely caespitose 28e. var. naiiu

4. Style scarcely exceeding the mature nutlets;
inflorescence somewhat open at maturity;
north-central Utah and southeastern Idaho ....

28c. var. sluinrzii

4. Style exceeding the mature nutlets 0.5-1 .5 mm;

inflorescence cylindrical and congested in tVuit;

southwestern Utah to southeastern California . .

28d. var. orina

28a. var. Iniiii/lis

Crypiuuilui humilis (A. Gray) Paysi.n var. humilis
hhirichium nlomcralum var 'hiimile A. Cray, Proc Am
Acad. 10:61. 1875.
OrcDcarya humilis Greene. Pitt. .1:1 12. 1896.
Oieocarya liispiila Nels. & Kennedy, Proe. Biol. Soc Wasli
19:156. 1906. (Type: Ormsliy Counlv. Nevada, in
Carson Valley, 24 April 1904. G. II True 865.)
Orcocarya ccliinoiJcs Maebride. C'ontr. Gray Herb. 48:31
1916, not Kryitilzl<ia ccliiiioiJcs M. l;. Jones.

Oreocarva iiiachridii Brand. I edde. Rep. Spec. Nov.

19:73. 1923. (Type: Mt. Jarbidue. Nevada. 6 July

1912, Nelson and Maebride 1960.)
Crvplanllia Iniinilis (Greene) Pavson. .^nn. .\lo. Bol. Gard.
'14:278. 1927.

Caespitose perennial, 0.5-3 dm tall; stems 1-many,
0.4-1.5 dm long, erect, strigose and sparsely setose;
leaves spatulate to oblanceolate. 1.5-7 cm long. 0.2-1
cm wide, weakly setose, strigose, and subtomentose,
the petioles ciliate-margined, bt)th siutaces pustulate;
intlorescence narrow to somewhat open, 0.8-1.7 dm
long, the floral bracts inconspicuous; calyx segments
linear-lanceolate, in anthesis 4-5 mm long, in fruit
becoming 7-13 mm long, setose; corolla white, the
tube 3.4-5 mm long, crests at base of tube evident to
nearly obsolete, fornices yellow, rounded, papillose,
about 0.5 mm long, limb 8-10 mm wide; style exceed-
ing mature fruit 1.5-2.5 mm; nutlets ovate-lanceolate.
3-4.5 mm long, 1.8-2 mm wide, dorsal surface muri-
cate, tuberculate, and usually rugulose, scar open at
the base or nearly closed. Collections: 62 (vii); repre-
sentative: Alexander and Kellogg 4443 (UC, US,
UTC); Maguire and Holmgren 25^)38 (ARIZ, UC,
ORE, UTC); A. Cronquist 8312 (ORE, UTC, WTU);
P. A. Munz 21036 (CAS); M. E. Jones 5163 (US);
Eastwood and Howell 8451 (POM); L. C. Higgins
1745, 1747, 1757, 1761 (BRY).

Lectotype: Bolander s.n., collected at Summit
Station (Donner Pass), Nevada County, California,
1871. Not seen.

Distribution: Southeastern Oregon and soinh-
western Idaho, south throughout Nevada and into
eastern California, mainly in mountainous regions.
Usually growing on gravelly slopes and ridges, 4,500
to 12,000 feet. Map No. 28a. April to August.

In eastern Nevada and western Utah there is a
mixing of two populations formerly referred to as C.
humilis and C. nana var. annmixta (Macbr.) Payson.
The specific limits between C. humilis of A. Gray and
that of C. nana are uncertain, and too many inter-
mediates are present to hold them apart. For this
reason the two species have been combined under
humilis. The varieties within this complex are also
very difficult to separate and can be done only tenta-
tively in certain areas. For example, in central Utah
the varieties shanizii. commixia. :md avina are ex-
tremely difficult to separate.

Brand's O. macbridii. iVom .larbidge, Nevada, was
described as new on the basis of the nutlets which
lend to have rugae between the muricalions. The
present author has found this same nutlet tirnamcn-
tation in all the varieties of C. humilis which is just
another reason for coiribining C. iiuniilis and ('. nana.

28b. var. coniniixta (Macbr.) Higgins stat. nov.

Crypianiha humilis (A. Gray) Payson var. commixia
(Macbr.) Higgins

Oreocarva coiiiiiiixia M:icbr. (ontr. Grav Herb. 48:33.
1916.

B10L0C;iCAL SFRIFS VOL. I .1, NO. 4 A Kl VISION Ol t'R Y 1' IAN Tl',\ SUBC ;i;N US ( )Ki;oC,AR Y A

37

Crvptanlha iianu (luistu.) I'a\sonvar. coininixiu (Macbr.)
■ Payson. Ann. Mo. Bol. Gard. 14:312. 1927.

Ciiespilosc perennials. 1-2.7 dm tall; stems 1 -sev-
eral, arising from the ends of the branched caudex,
0..^-l dm long, weakly strigose and spreading setose;
leaves spatulale to broadly oblanceolate, 2.5-6 cm
long, 0.5-1.2 cm wide, strigose and spreading setose;
intlorescence open. 0.8-1.8 dm long, foliar bracts
evident on lower part of the inflorescence; calyx
segments linear-lanceolate, in anthesis 3.5-4.5 mm
long, in fruit becoming 7-10 mm long, setose; style
exceeding nuitme fruit 0.7-1.6 mm; nutlets lance
-ovate. 3.5-4 mm long, muricate. tuberculate. or
sometimes wilh the niurications joined to form short
irregular ridges, scar subulate or nearly closed. Collec-
tions: 18 (iv); representative: L. N. Goodding 1074
(GH. RM); M. H. Jones 5388L (US); B. F. Harrison
10148 (BRY); R. C. Holmgren 266 (BRY); L. C.
Higgins 1468. 1617 (BRY).

Holotype: L. N. goodding 1074. collected in Juab
County. Utah, on sandy slides at Juab. '^' June 1'^'02.
GH. Photograph at BRY. Isotype at RM.

Distribution: Central Utah to eastern Nevada.
Growing on gravelly soil oi talus slopes. 4.500 to
7.500 feet. Map No. 28b. May to July.

This variety may be distinguished by its open
inflorescence, setose or coarsely strigose leaves, and
the length of the style.

The variation within coiiimixia is also noticeable,
and a population along the Sevier River in Sevier and
Piute Counties may prove to be another variety or
species. The nutlets of this population along the
Sevier River are more tuberculate than muricate. the
scar is closed or nearly so. and the leaves are greener
than in typical commixta.

28c. var. shantzii (Tidcstr.) Higgins stat. nov.

Crypiiiiuha liiiiiiilis (A. Gray) Payson var. sliantzii
(Tidestr.) Higgins.

Orcocarva shaiuzii J\A';\U. Proc. Biol. Soc. Wash. 26:122-

1913.
Oreocarva dolosa Macbr. Conlr. Grav Herb. 48:32. 1916.

(Type: College Beneh. Logan, Utah. 4 lune 1909.

Smith 1605.)
Crypiantha nana (l.aslw.) Payson var. slianlzii ( I'ldeslr.)

Payson. Ann. Mo. Bol. Gard. 14:313. 1927.

Caespitose perennial. 1-2 dm tall; stems I -many
from the ends of the much branched caudex. 0.3-1
dm long, weakly setose and strigose; leaves spatulale
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 cymules slightly elongating, lower
foliar bracts rather conspicuous in the young inflor-
escence; calyx segments densely setose and subto-
mentose; style scarcely exceeding the mature nutlets;
nutlets muricate. Collections: 40 (v); representative:
B. Maguire 12952 (RM); C. P. Smith 1573 (RM); C.
P. Smith 1605 (RM); Kearney and .Shantz 3098 (US);

R. Gourley 8068 (UT); R. J. Davis 988 (IDS); A. A.
Beetle 5792 (ND); H. Palmer 42 (RM); L. C. Higgins
1087 (BRY).

Holotype: Kearney and Shantz 3098, collected in
dry saline soil at Grants Station south of the Great
Salt Lake, Utah, 6 August 1912, GH.

Distribution: Southwestern Montana, eastern
Idaho, and northern Utah. Growing on a wide variety
of soils, 4,500 to 8,000 feet. Map No. 28b. April to
July.

This variety may be distinguished by the short
style, evident foliar bracts, and the basal leaves, which
are silvery strigose and setose.

Payson designated the specimen collected by C. !'.
Smith 1605. as the type of var. shaiuzii. However, the
plant collected by Kearney and Shantz 3098, should
have been used since it was designated as the type in
the original description.

28d. var. orina (Payson) Higgins stat. nov.

Cryptautlui humilis (A. Gray) Payson var. orina

(Payson) Higgins.

Crvpranllw nana (Eastw.) Payson var. or/zjo Payson, Ann.
■ Mo. Bot. Gard. 14:314. 1927.

Densely caespitose long4ived perennials, 0.5-1.5
dm tall; stems several, 0.2-0.7 dm long; leaves spatu-
lale to oblanceolate, obtuse, 2-4 cm long, tomentose
and appressed setose with rather weak bristles; inflor-
escence narrow, cylindrical, lower foliar bracts
inconspicuous; calyx segments linear-lanceolate,
densely setose and tomentose; style exceeding the
mature fruit 0.5-1 mm; nutlets muricate or tubercu-
late. Collections: 21 (v); representative: P. A. Munz
21036 (UT); Eastwood and Howell 9377 (CAS);
Ripley and Barneby 3485 (CAS); G. H. Bentley s.n.
(RM); J. L. Reveal 1414 (BRY, LA); S. L. Welsh
5226 (BRY); L. C. Higgins 1234, 1409, 1449, 1455
(BRY).

Holotype: Georgia H. Bentley s.n., collected in
Nye CoLMity, Nevada, in the vicinity of Currant, June
1916, RM.

Distribution: Southwestern Utah, southern Ne-
vada, and southeastern California. Growing on
gravelly loam or clayey soils, mainly in the pinyon-
juniper belt. 3,500 to 7.000 leet. Map No. 28b. April
to July.

Variety orina has its closest relalive in var.
slnintzii. It may be separated from that variety only
tentatively by the more tomentose leaves and calyces
and the longer style. The foliar bracts are also less
evident in this variety.

28e. var. nana (Easw.) Higgins stat. nov.

Crypianlha Ininiilis (A. Gray) Payson var. nana
(Easlw.) Higgins.

Orcocarva nana lastw. Bull. Torrey Bol. Club 30:243.
1903.

liKU.llAM YOUNCI LINIVKKSI I Y SCIKNCE BULLEIIN

Map No. 28a. Parts of western United States. Range of C.
Iiiimilis (Gray) Payson: (Cireles), var. humilis: (Tria
var. nana (l-astw.) Higcins.

oi c. Map No. 2Sh. Parts of western United Stales. Range of C

ngles) humilis (Ciray) Payson: (circles), var. oviiw (Pa\sonl Higgins;

(triangles) var. s/iantzii (Tidestr.) Higgins; niset, (circles), var.

(triangle

coiniiuxla (Macbr.) Higgins.

Crvptantha nana (lastw.) Pa\'son var. ivpica. Ann. Mo.

Bot.CIard. 14:315. 1427.

Caespitose perennial. 0.5-I..S dm tall; stems .sev-
eral, 0.2-0.7 dm long, selnse; leaves oblanceolate to
spatulate, 0..^-4 cm long. 0.2-0.(i cm wide, strigose to
subtomentose, spreading setose; inllorescence narrow,
cylindrical. 0.2-1 dm long, setose: calyx segments
linear-lanceolale. in anihesis 2.5-3.5 mm long, in fruit
becoming 6-8 mm long, setose; corolla white, the
tube 2. 5-.?. 5 mm long, crests at base of tube evident;
style shorter to slightly longer than mature liuit;
nutlets nuiricate or sometimes tuberculate or rugu-
lose. Collections: 18 (v); representative: Ripley and
Barneby 4675 (CAS); W. A. Weber I 1258 (COLO);
A. Eastwood s.n. (CAS):G. E. Osteihout 4484 (RM).
D. Wiens 3066 (COLO): L. C. Higgins 1066. 15^)8
(BRY).

Holotype: A. Eastwood s.n., collected m Mesa
County, Colorado, near Grand Junction on the mesa
above the Gunnison River. 17 May 1842, CAS.
Photograph at BRY. Isotypes at RM. VC. GH.

Distribution: Western Colorado and eastern Utah.
Growing on sandy or clay soils, 4,500 to 7.000 feet.
Map No. 28a. April to July.

Variety nana can usually be separated from the
other varieties of this complex by the setose leaves
which are only rarely tonientose, the very short style
which usually does not exceed the fruit, and the
compact intJorescence.

The C. Iiuniilis complex is distinguished maiiiK by
the short corollas and the uniformly muricale nutlets.
These are also characteristics shared by C. cana and C.
brcvijlnni: however, these species have an indument
which is silky-strigose and with few or no pustulate
hairs especially on the ventral surface.

24. Cryptautha nnisun-wu Mun/

Cryptiuiiliu roasionim Munz. El Aliso 3:124. 1455.
Densely caespitose, long-lived perennial. 0.1-0.3
dm tall: stems many from the end of the branched
caudex. raiher slender. 0.1-0.2 dm long: leaves spatu-
late lo oblanceolate, acute. 0.5-1.2 cm long, densely
strigose and appressed setose, appearing almost
tomentose, scarcely pustulate: inllorescence compact,
about I cm long, foliar bracts inconspicuous: calyx
segments linear. In aiithesis 2.8-3.1 mm long, in fruit
becoming 3.8-4.3 mm long, strigose. and more or less
setose: corolla white, the tube 2.5-3 iinn long, crests
at base of tube well developed, fornices low, rounded,
yellow, about 0.5 mm long, limb 4.5-5.5 mm wide:
style slightly surpassing the nutlets: nutlets 2.3-2.5
mm long, lance-ovate, acutish, dorsal surface rugu-
lose. with low, rounded, irregular ridges, also some-
what nuiricate, ventral surface similar but the mark-
ings less evident, scar narrowly triangular, and lacking
;in elevated margin. Collections: 1 (0); representative:
J. C. and A. R. Roos 6015 (RSA).

lilOL(X;iCAL SKRIKS VOL. I J. NO. 4 A RKVISION 0\ CR Y I' I ..\N rH.\ .SUBGENUS ORKOCARY A

39

Hololype: J. C. and A. R. Roos 601 5, collected in
Inyo Cnunty, California, along crest of Inyo Moun-
tanis three miles east of Badger Flat, at 10,600 feet,
l.> August l^^'.^.v RSA. Photograph at BRY. Isotypes
alCA.S,GH.

Distribution: Apparently endemic to the type
locality. Inyo County. California. Growing on open
rocky slopes, 10,600 feet. Map No. 2'-). July and
August.

Cryptantha roosioruiii is not very well known, and
maii\' more collections of this species are badly
needed. It is perhaps most closely related to C.
Ininiilis. but differs in the moie caespitose habit,
smaller leaves, calyx, and corolla. The inflorescence is
also more compact, and the whole plant is much
smaller.

30. Cryptaiitlia cioiipacla lliggins

Crypianilia voiiipacla Higgins, Great Basin Naturalist

:X;|'-)(i. 1968.

Densely caespitose perennial, 0.3-1 dm tall; stems
numerous, arising from a woody root, 0.1-0.4 dm
long, tomentose below, weakly strigose above; leaves
oblanceolate to spatulate, obtuse. 0.5-1.5 (2) cm
long. 0.2-0.4 cm wide, dorsal surface with appressed
setose-pustulate bristles, also densely strigose or
subtomentose. ventral surface similar but with fewer
pustulate hairs, the petioles tomentose; inflorescence
narrow, nearly capitate. 1 -5 cm long; foliar bracts
evident but not conspicuous; calyx segments lanceo-
late. 2-2.5 mm long in antliesis. in fruit becoming
3.5-4.5 (5) mm long, densely white setose and tomen-
tose; corolla white, the tube 1.8-2.2 mm long, crests
at base of tube evident, fornices yellow, rounded,
papillose, about 0.5 nnn long, limb 4.5-5.5 mm wide;
style equalling or shorter than mature fruit; nutlets
lance-ovate, acute. 2.5-3 mm long. 1.5-1.8 mm wide,
only one to two maturing, dorsal surface muricate or
weakly tubercuiate-rugulose. ventral surface muricate.
scar open, subulate to narrowly triangular, elevated
margin lacking. Collections; 6 (ii); representative: R.
C. Holmgren 521 (BRY); B F. Harrison 6371 (BRY);
L. C. Higgins 1462 (BR>).

Holotype: L. C. Higgins 1613, collected in Millard
County, Utah, about 8 miles west of Desert Range
E.xperiment Station Headquarters along Hwy. 21 . 100
m west of pass at the north end of Needle Range. 18
Jime 1968. BRY. Isotypes at CAS. GH. NY. POM.
US. UTC.

Distribution: Known only from southwestern
Millard County. Utah, but to be expected from north-
ern Beaver County. Utah, and perhaps in eastern
Nevada. Growing on gravelly loam soil. 4.500 to
6.000 feet. Map No. 30. May to July.

Cryptantha compacta is most closely related to C.
hiauilis but differs in its more compact and caespitose
habit, smaller leaves, shorter caly.x segments, and

Map. No. 29.
onim Munz.

Inyo County. California. I^angc of C roosi-

Map No. 30. Western Utah and adjoining Nevada. Range of
C compacta Higgins.

40

BRIC.HAM YOUNCi UNIVI.KSIIY SCII-NCl- liULLKTIN

smaller corolla. This plant has been known for over
30 years, but has been placed with C. Iiuinilis. prob-
ably due to the immaturity of the specimens. In
observing this species in the field it becomes even
more apparent of its right to specific distinction, due
to its dense caespitose habit that more resembles C.
caespiiosa than C. Iiumilis. At tiie type locality it is
the most common plant, growing on shallow stony
loam.

31 . Ciyiuaiuha aiiui (A. Nels.) Payson

Civpiaiilha cana (A. Nels.) Payson. Ann. Mo. Bot.

Card. 14:316. 1927.

Oreocarya cam .\. Nels. Bol. Gaz. 34:30. 1902.

Caespitose perennials. 0.5-2 dm tall; stems many,
arising from a multiple caudex. 0.2-0.5 dm long.
weakly setose; leaves narrowly oblanceolate. acute.
very dense at the ends of the caudices. 2-6 cm long,
0.3-1 cm wide, uniformly silky-strigose, also with
small inconspicuous pustulate hairs on both leal
surfaces; inflorescence narrow, setose, foliar bracts
inconspicuous; calyx segments linear-lanceolate. 3-4
mm long in anthesis, in fruit becoming 6-7 mm long,
weakly setose; corolla white, the tube 3-4 mm long,
crests at base of tube evident, fornices yellow,
rounded, papillose, about 0.5 mm long, limb 6-9 mm
wide; nutlets lance-ovoid, 2.5-3 mm long, 1.4-l.cS mm
wide, usually only one maturing, margins acute,
dorsal surface muricate with elongated papillae, or
sometimes tuberculate, ventral surface similar but less
roughened, scar narrowly triangular, and without an
elevated margin; style shorter than the mature fruit.
Collections: 31 (iv); representative: E. J. Palmer
37423 (GH); C. L. Porter 5723 (GH. RM); Ripley and
Barneby 10547 (GH); A. Nelson 2876 (ND-G); J.
Ewan l'2770 (GH); L. C. Higgins 1534. 1542. 1537
(BRY).

llolotype: A. Nelson 8309, collected in Cioshen
County, Wyoming, at Fort Laramie, on gravelly hill-
top, 29 June 1901, RM. Photograph at BRY. Isolype
at GH.

Distribution: Western Nebraska, northeastern
Colorado, and southeastern Wyoming. Growing on
gravelly loam soils, 4,000 to 6,000 feet. Map No. 31 .
May to early September.

Crypiaiuha cana is similar to C. caespiiosa in
habit, but can be distinguished from C caespiiosa by
the silky-strigose pubescence, sharply muricate nut-
lets, and the different range.

32. Cryptaniha Itrcvijlora (Osicrh.) Payson

Crvptanilia hrcviilDia (Osterh.) Payson. Ann. Mo.
Bot. Card. 14:318. 1927.

Orcovarva hicviflora Ostcrli. Univ. Wyo. Piibl. Bol. 1 : lfi9
1926.

Long-lived perennials. 1.6-3 dm tall; stems several.

slender. 0.7-1.7 dm long, densely white setose at the

base, strigose above; leaves oblanceolate 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 small pustulae. ventral surface similar but with
fewer pustules; inflorescence narrow in fiower. but
becoming broad and open at maturity. 0.6-2.7 dm
long, setose; caly.x segments linear-lanceolate. 4.5-6
mm long in anthesis, in fruit becoming 7-9 mni long,
setose; corolla white, 3.5-4.5 mm long, crests at base
of tube evident, fornices yellow, rounded, about 0.5
mm long, limb 8-1 2 mm wide; style exceeding mature
fruit by 2 mm or less; nutlets lanceolate, 3.4-4 mm
long, 2-2.5 mm wide, less than four nutlets maturing,
margins in contact, knifelike, dorsal surface uni-
formly muricate or tuberculate. ventral surface siirii-
lar, scar open, narrowly triangular, margin not ele-
vated. Collections: 25 (vii); representative: R. C.
Rollins 1736 (UTC); .S. L. Welsh 466 (COLO); J.
Brotherson 806 (BRY); W. A. Weber 5310 (COLO);
Higgins and Welsh 1018 (BRY) L. C. Higgins 1084
(BRY).

Holotype: G. E. Osterhout 6414. collected in
Uintah County, Utah. 6': miles north of Jensen. 19
June 1925. RM. Photograph at BRY.

Distribution: Northeastern Utah in Duchesne and
Uintah counties. Growing on heavy clay soils, 4,500
to 7,000 feet. Map No. 32. May to July.

Crypiaiuha hrcrijlora is apparently endemic to the
Uintah Basin. It differs from its closest relative, C.
fulvocanesceiis. by the short corolla, low fornices,
different pubescence, and shorter style. The range of
the two is also consistently different, C. fulvo-
canesceiis being more southerly and never in the
L'iiUah Basin.

33. Ciypianiha propria (Nels. & Macbr.) Payson

Crvptanilia propria (Nels. & Macbr.) Payson, Ann.

Mo. Bot.Gard. 14:317. 1927.

Kivnitzkia fuhocaiwscens var. idahocitsis Jones. Conn.

West. Bot. 13:6. 1910. (Tvpc: .loncs 6474. near

Wciscr, Idaho. 28 April 1910.)
Oreocarya propria Nels & Mactir. Bol. Cj?. 62:14.^. 1916.

Caespitose perennials from a strongly lignified
taproot. 1-2.3 dm tall; stems several. 0.7-1 .2 dm long,
finely strigose and setose; leaves oblanceolate to
spatulate. obtuse. 3-9 cm long. 0.4-1.2 cm wide,
dorsal surface finely strigose and scattered appressed
setose, ventral surface finely and uniformly strigose,
without pustules; inflorescence narrow, 0.5-1.2 dm
k>ng, foliar bracts inconspicuous; calyx segments
linear-lanceolate, in anthesis 4-6 mm long, in fruit
becoming 7-13 mm long, densely setose-hirsute;
corolla white, the tube 3.5-4.5 mm long, crests at
base of tube conspicuous, fornices yellow, rounded,
papillose, about 0.5-1 mm long, limb 6-8 mm wide;
style exceeding mature fruit 1.5-2 mm; nutlets ovate-
lanceolate, 3-4 mm long, 2.5-3 mm wide, margins in

BIOLOGIf.Al SI RII S VOL. 1 .1. NO. 4 .'\ RKVISION Ol I K"! I' I .\N 11 :,\ SUBGENUS OKI- OC AKY A

41

®

•

•

1

i

N

1

» [
••

•
•

•

' 1

. ^

1
1

J

H -|

' 1

1 ' 1

1 "n J

>""' 1 Ia i'

Map. No
Dakota,
Range of

. 31. Southeastern Wyoming, southwestern South
western Nebraska, and north eentral Colorado.
C caim (A. Nels.) Payson.

Map. No. 32. Northeastern LItah, Duehesne and Uintah
Counties. Range of C. brciifloia (Osterh.) Payson.

coii1;ilH, acute, dorsal surface densely and irregularly
rugulose, with narrow high ridges, inuricate between
the ridges, ventral surface tuberculate, or with some
of the tuberculations joined to form short ridges, scar
open, linear, and without an elevated margin. Collec-
tions: (i (0); representative: J. B. Leiberg 2049 (UC,
GH, POM, US); J. B. Leiberg 2223 (UC, GH, US); R.
J. Davis 4496 (GH); H. M. Tucker 1022 (GH. IDS);
M. E.Jones 4674 (POM).

Holotype: J. B. Leiberg 2049, collected in Malheur
County, Oregon, at Vale, 14 May 1896, GH. Photo-
graph at BRY. Isotypes at UC, GH, POM, US.

Distribution: Southwestern and western Idaho and
eastern Oregon in Malheur County. Growing in clay
soils, 2,000 to 4,000 feet. Map No. 3.^. April to June.

Cryptaiilha propria, a relatively sporadic species
occurring in western Idaho and eastern Oregon, is not
often collected throughout its range. In general
appearance it is not unlike C. Iiumllis var. hitmilis. C.
caiia, or C. brevijlora. The latter two species are far
removed geographically, occurring in Wyoming and
Utah respectively. This species is probably most
closely related to C Inimilis. however, very distinct in
its racemose inflorescence and the uniformly strigose
ventral leaf surface which lacks pustules.

34. Cryptantba fiilvocancsccns (Wats.) Payson

Densely caespitose perennials from a strongly ligni-
fied taproot, 0.8-3 dm tall; stems many from a
multiple caude.x, 0.5-1.3 dm long, white hairy at the
base, setose-htrsute upward; leaves spatulate or oblan-
ceolate, acute to obtuse, 1.5-7 cm long, 0.4-1.2 ctii
wide, uniformly strigose, pustules mainly confined to
the dorsal surface; inflorescence narrow or somewhat
open at maturity, 0.3-1 .9 dm long, white or yellowish
setose, foliar bracts inconspicuous; caly.x segments
linear, 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 evident or lacking,
fornices yellow, emarginate or rounded, 0.7-1.3 mm
long, limb 7-9 mm broad; style e.xceeding mature
frtiit 3-7 mm; nutlets lance-ovate. 3.5-4.5 mm long,
2-3 mm wide, one to two usually maturing, margins
acute to obtuse, in contact when more than one
nutlet matures, both surfaces densely and uniformly
inuricate, scar open or nearly closed, elevated margin
lacking.

Key to the varieties of C. lUlrnccincsccns

I. Miirications on the nutlet rounded; corolla 9-13
mm long; inllorescence narrow, white setose at
maturity 34a. m. fulvocanescens

I. Murications on the nutlet with one or two setose
projections; corolla 7-9 nim long; inflorescence

broader and usually yellowish at maturity

34b. var. echinoides

BKICJHAM YOUNG UNIVERSITY SC1HNCT-: HULLKTIN

Map. No. 33. Harts of western Idaho and eastern Oregon.
Range of C propria (Ncls. & Maebr.) Payson.

Map No. 34. Paris of western United States. Range of /i(/i'o-
canescens: Ovar. fiiliocanescens (Gray) Payson; .% var. echi-
noides (Jones) Iliggins.

34a. var. fulvocancsccns

Cryplanllia fiilrocaiicsccns (Wiits.) I'aysoii vai. ////n»-

cviu'sccns

luilrichiuin f:,k>mera!um var. '.' fiilyocaiicscciis S Wats.

Bot. Kingh\p. 243. 1871.
Kri I rich ill 1)1 fulvocancsccns Clrav. Proe. .\m. .Acad. lOdl.

1875.
Krvnitzkia fulvocancsccns Gray. Proe. Am. Acad. 20:280.

■ 1885.
Orcocarva fulvocanescens (Wats.) Greene. Pitt. 1:58.

1887.
Orcocarva nilida Greene. PI. Baker. 3:21. r)01. (Type:

Deer Run, Colorado. 11 .iiine 1901, C. I-. Baker 95.)
Cryplanllia Julvocancsceus (Wats.) Pavson. .Ann. Mo. Bot.

Gard. 14:319. 1927.

Densely caespitose perennial. 1-.^ dm tall; iiinor-
escence narrow, white setose; pedicels 2-.^ mm long;
corolla white, the tube 9-1 I mm long, crests at base
of lube evident or lacking, t'ornices 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. Collections: 5S
(viii); representative: A. Cronquist 9006 (POM, IITD;
A. H. Holmgren 3225 (US); G. Heller 3517 (ND-G,
US); Fendler 632 (GH);C. F. Baker 561 (ND-G, US);
C. F. Baker 95 (UC, POM. GH. RM. US); L. C.
Higgins W9, 1012. 1307 (BRY).

Holotype: Fendler o32, collected in Santa Fe
County. New Mexico, near Santa Fe, 1847. GH.
Photograph at BRY. Isotypes at PH, US.

Distribution: Western Colorado, northwestern
New Mexico, northeastern Arizona, and eastern Utah.
Growing on sandy soil. 4.000 to 7.500 feet. Map No.
34. April to August.

Cryptantha fulvocanescens var. fulrociincscciis is
most likely to be confused with C. hrcvijlora but
differs in the longer corolla tube, narrower intlor-
escence, longer style, and different pubescence.

34b. var. cchinoiJcs (Junes) Higgins

Ciypuintlki Julrocanesccns (Wats.) Payson var. «■/»'-
iioIlIcs (Jones) Higgins. Great Basm Nattnalisl

29:30. 1969.

Krvniizkia cchinoidcs .tones, Proe. Calif. .Xead. Sei. II.

'5:709. 1895. . .
Orcocarva cchinoidcs (Jones) Maelir. Conlr. Gray Herb.
48:31. 1916, as to synonvmy. not .is to speeimens
cited.
Crvplaiuha cchinoidcs (JonesI I'aNsun. \nn Mo. liol

Gard. 14:321. 1927.
Caespitose perennials. 0.8-3.6 dm tall: iiillor-
escence narrow to somewhat open ai maiurity. yel-
lowish setose; pedicels 3-10 imn long; coiolla white,
the tube 7-9 mm long, crests at base of tube lacking
or sometimes evident, fornices yellow, omarginate;
nutlets lance-ovoid. 4-4.7 mm long. 2.5-3 mm wide.
the dorsal surface with one to two setose projections
lerminating each murication, scar asymmetrical, and
withoui an elevated margin. Collections: 19 (vi);
representative: M. F. Jones 5297p (POM); M. F.
Jones 5312ac (POM); Fastwood and Howell 9265

BIOLOGICAL SFRIKS VOL. I .^. NO. 4 .A Kl- VISION Ol CR'i I'l AN ri'.X SUBOLNUS ORKOCAKYA

43

(L'TO; R. 11. Peebles l4(vS8 (ARIZ. LiS); Reveal.
Gentry and Davidsee 781 (BRY); .4. Croiuiuist lo: 1 2
(BRY, NY);L.C. Higgins 1008, l.U4(BR1).

Lectotype: M. E. Jones 5297p. collected in Kane
County, Utah, in Pahria Canyon. 26 May 18^4, POM.
Photograph at \iR\ .

Distribution: South-cenlral Utah and north-central
Arizona. Usually growing on heavy saline clay soils,
4,000 to 7,500 feet. Map No. 34. April to July.

"Payson cites Fritrichhim gloiiicratum var. C^)
fulvocancsc-cns. Wats. Bot. King Exp. 243 ( 1871 ), as a
synonym of C. Iniinil/s. This 1 believe is incorrect.
Watson took a herbarium name of Gray's and de-
scribed under it an aggregate species. When Gray
finally published his name h'. tiih'ocancsccns he cited
Watsons trinomial as a synonym and appears to have
accepted the limits given it by Watson. Watson, how-
ever, found Gray's name on a specimen from New
Me.xico collected by Fendler (no. 632). He accepted
that specimen as belonging to his variety and indi-
cated it as the source of his botanical name. He erred
in taxonomic judgment, though, in placing with it
certain material from Nevada and possibly Utah that
belongs at least in part to C. Iiiimllis. Payson placed
particular emphasis on tliese latter specimens. How-
ever, since 'Vatson included tlie Fendlerian New
Mexican pla.it in his concept and indicated it as the
source of his name, then, it should be taken as tlie
type of fulvocancscens. both as variety and species.
Consequently the name applies to a very different
plant from that described as Orcovarya hinnilis by
Greene." (Johnston 1932)

The variety ccliinolclcs was entirely misunderstood
by Macbride (1916). From the specimens he cited
from western Nevada and adjacent California, it is
evident that he never saw a specimen of true cclii-
iioulcs. It ditTers from the typical plant in that the
murications are setose, and the nutlets are usually
larger.

35. Crypiantha jouesiuna (Payson) Payson

Crvpiantlia joncsiana (Payson) Payson, Ann. Mo. Bot.

Card. 14:323. 1927.

Oreocarva joncsiana Payson. Univ. Wvo. PubL 15ot. 1 :U->iS.
1926.

Coarse perennials, 0.5-1.5 dm tall; stems many.

arising from a thick multiple caudex, 0.2-0.7 dm long,

setose: leaves spalulate, 1-4 cm long, 0.4-1 .3 cm wide,

coarsely appressed setose-pustulate, leaf bases also

setose with dense white hairs; inflorescence narrow,

somewhat capitate, with one to three flowers in the

axis of the bracts 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, campanulate in the

throat, fornices low and broad, papillose, crests at

base ol' tube lacking, limb 4-13 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. Collections:
1 1 (V); representative; W. P. Cottam 5247 (UT); M. E.
Jones s.n. (POM); D. Atwood 1301 (BRY); Higgins
and Reveal 1265, 1275, 129') (BRY); L. C. Higgins
1322, 1308 (BRY).

Holotype; M. E. Jones s.n., collected in Emery
County. Utah, on the San Rafael .Swell. 15 May 1914,
POM. Photograph at BRY.

Distribution: Endemic to the San Rafael Swell in
Emery County, Utah. Usually growing on barren clay
hills, 4,000 to 7,000 feet. Map No. 35. April to June.

Cryptantha joncsiana is probably most closely
related to C. fulvocancscens. but resembles it not at
all. This handsome plant can be distinguished by its
very large corolla, low broad fornices, spatulate
leaves, and harsh pubescence.

36. Cryptuntlia tliyrsijlora (Greene) Payson

Crvpiantlia llivrsi flora (Greene) Payson, Ann. Mo.

Bot.Gard. 14:283. 1927.

Eiiirkhhim ,^!otncraliiiit var. Iiispidissiiiniiu Torr. Bot.
Me\. Bound. Surv. 140. 1 859. at least m part.

Oreocarva tliyrsijlora Greene, Pitt. 3:111. 1896.

Oreocarva liispidissinia (Torr.) Rydb. Bull. Torrcv Bot.
Club 33:150. 1906.

Oreocarva urticacca Wooton & Standi. Contr. U. S. Natl.
Herb. 16:166. 1913. (Type: Canyoncito, Santa Fe
County, New Mexico, IS .lune 1897, A. A. & E. G.
Heller 3731.)

Oreocarva dura Nels. & Macbr. Bot. Gaz. 62:144. 1916.
(Type: E. L. Johnston 418. 1907. central Colorado.)

Oreocarva monospeniia Osterli. Bull. Torrey Bot. Club
46:55. 1919. (Tvpe: Truudad. Las Animas Co., Colo-
rado, 20 July I9IK, Osterhout 5754.)

Short-lived perennials or sometimes biennial. 1.7-4
dm tall; stems stout, 1-several, arising from the base,
0.5-1.8 dm long, conspicuously spreading setose;
leaves oblanceolate, obtuse, 5-12 cm long, 0.5-1 .4 cm
wide, conspicuously spreading setose, pustulate on
both surfaces, with some finer strigose hairs beneath,
the petioles ciliate-hirsute; intlorescence very broad
and open, cymules much elongating, 1-2 dm long,
0.6-2.5 dm wide, setose, foliar bracts 2-3 cm long, but
not very conspicuous due to the width of the inflor-
escence; calyx segments linear, in anthesis 3-4 mm
long, in fruit becoming 6-9 mm long, setose; corolla
white, the tube 3-4 mm long, crests at base of tube
conspicuous, fornices yellow, emarginate, papillose,
about 0.5 mm long, limb 5-8 mm wide; style exceed-
ing mature fruit 1-1.5 mm; nutlets ovate to ovate-
lanceolate, 2.5-3.5 mm long. 1.5-2 mm wide, usually
two to four maturing, acute, margins in contact,
dorsal surface low rugulose and tuberculate, some-
times with murications between the rugae, ventral
siuface similar but with fewer ridges or sometimes
almost smooth, scar subulate, the margin not ele-
vated. Collections: 110 (xv); representative; A.
Nelson 7306 (RM. COLO, US); M. E. Jones 972 (RM.

44

liKK.HAM YOUNC; UNIVI KSITY SCIl-NCK BULLI- UN

US, UTC); C. F. Baker. F. S. Haiic & Tracy 13
(ND-G. RM. US); W. A. Wcbcr 43cS9 (ARIZ. COLO):
R. C. Rollins 18(i5 (ND. RM): U. T. Walcrfall (COLO,
UTC); M. Ownbey 131 1 (COLO. RM, UTC): L. C.
Higgins 2030, 3290. 3823. 3830. 2042 (RRY,
WTSU ).

Lectotype: E. L. Greene s.n.. collecled in soulli-
eastern Wyoming, at Cheyenne. 6 July 1892, ND-G.
Photograph at BRY.

Distribution: Western Nebraska and southeastern
Wyoming, south ihrougliout eastern Colorado, New
Mexico, and into Oklahoma. Growing in gravelly
loam soil. 4. .500 to 9,600 feet. Map No. 36. Late May
to September.

Harly collection.s of this species were ol'len re-
ferred to C. celosioides. but it is probably not very
closely related to that plant. The broad inflorescence
is the outstanding characteristic of the species, but in
additiiHi the tlowers of /liyrsijlora are much smaller
and the blooming season later than in C. celosioides.
In observing the types of (). urticacca. dura, and
monospenna it is apparent iluit they are e.xact syno-
nyms o f tliyrsijlt >ra.

37. Crrpiiiiiilia ciciiii ( Lasiw.) Payson

Cryptanilia elata (Eastw.) Payson. Ann. Mo. Bot.

'card. 14:285. 1927.

Oreocarva elala I'.aslw. Bull. Torrcv Bol. Club :>0:24l.
1903.

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 tapering abruptly to the narrow petiole, dorsal
surface strigose and appressed setose, ventral surface
strigose. btith surfaces pustulate; inflorescence spread-
ing in age, i .5-3.5 dm long, setose, foliar bracts incon-
spicuous; calyx segments lanceolate, in anthesis 3-4.5
mm long, in fruit becoming 7-8 mm long, hirsute;
corolla white, the tube 3.5-5 mm long, fornices
yellow, rounded, papillose, about 1 mm long, crests
at base of tube well developed, limb 6-.S mm wide:
style exceeding mature fruit 0.5-2 mm: nutlets lance-
olate-ovate, 4-4.5 mm long, 2-2.5 mm wide, usually
all four maturing, margins in contact, dorsal surface
densely luberculate and somewhat rugulose. the sur-
face also covered with dense, minute papillae, ventral
surface similar but the rouglienings less prominent,
scar closed, or narrowly open at the base, and with-
out an elevated margin. Collections: I 1 (iii); repre-
sentative: A. Eastwood s.n. (CAS. Gil): S. L. Welsh
6952 (BRY); G. E. Osterhout 5996 (RM); W. A.
Wcbcr 11294 (BRY. COLO): L. C. Higgins 1479
(BRY).

Lectotype: A. Eastwood s.n.. collected in Mesa
County. Colorado, near Grand Junction on the road
lo the coal mines, growing on bare clay hills charac-

Map No. 35. F.mery County. Utati. Range of C. jonesiana
iPav'son) Parson.

Map No. 36. Parts of western United State
lliyrsiflora (Greene) Payson.

Range of C

Bini (k;ic'ai sikii's vol. i j, nci. 4 a ri;vision oI' cryi'tantim subgenus oreocarya

45

terislic of Ihc region, 25 May 18'-)2. CAS. I'liotograph
at BRY. Isoiypes at GH. NY, US.

Distribution: West central Colorado in Mesa
County, and east-central Utah in Grand County.
Growing on heavy clay soil. 4,500 to 5,500 feet. Map
No. 37. May to June.

This species resembles C. iiisolira from southern
Nevada, but is probably not very closely related to
that laxon. it differs in the shorter spatulate leaves
and larger differently marked nutlets which have the
scar closed. In practice this plant is so distinct that it
is not to be confused with any other species in this
subgenus. Additional collections of this very narrow
endemic are badly needed in order to determine the
exact limits of the species.

38. Cryptantlici scrlcca (Gray) Payson

Crvptaniha sericca (Gray) Pavson, Ann. Mo. Bol.
Gard. 14:286. 1927.

Kiyiiitzkia sericea Gray, Proc. Am. Acad. 20:279. 1SX5.
Oreocarya sericea Greene, Pitt. 1 ;58. 1887.
Oreocarya affiius perennis A. Nels. F.rytliea 7:67. 1899.

(Type: Green River, VVyo^lin^, 31 Mav 1897, A.

Nelson 3035.)
Oreocarva an;ei!lea R\db. Bull. Torrcv Bot. Club 31 :637.

1904. (Type: Ritle, Gart'iekl County. Colorado, 1900.

Osterhout 2122.)
Oreocarva perennis Rvdb. Bull. Torrey Bot. Club 33:150.

1906.
Oreocarya proeera Ostcrh. Bull. Torrey Bot. Club 47:21 1.

1920. (Type: Glenwood Springs. Garfield Countv,

Colorado, 18 June 1 899. Osterhout 1867.)
Cryplantha sericea var. perennis (Nels.) Payson, Ann. Mo.

Bot. Gard. 14:288. 1927.

Perennials, I .5-4.3 dm tall; stems 1-several,
branched from the base, 0.5-1 .2 dm long, setose with
spreading hairs; leaves oblanceolate to spatulate,
obtuse. 2.5-10 (15) cm long, 0.5-2 cm wide, dorsal
surface strigose and slightly appressed to spreading
setose, pustulate, ventral surface silky-slrigose, pus-
tules lacking or very inconspicuous; intlorescence
narrow to somewhat open, 0.5-3.2 dm long, setose-
hispid, foliar bracts 2-5 cm long; calyx segments
lanceolate, 2.5-4 mm long in anthesis, in fruit becom-
ing 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, fornices yellow, depressed, broad,
0.5-0.6 mm long, limb 7-9 mm wide; style exceeding
the mature fruit 0.5-1.3 mm; nutlets lanceolate,
2.5-3.5 mm long, 1.5-2 mm wide, usually all four
maturing, margins acute or narrowly winged, in
contact, dorsal surface with low rounded tubercula-
tions, also somewhat rugulose and muricate, ventral
surface similar but the markings less evident, scar
straiglit. closed, and without an elevated margin.
Collections: 80 (xii); representative: C. L. Porter
4583 (MONT, RM, WTU); R. C. Rollins 1772 (ND,
UTC); B. Magiiire 12378 (UTC); L. Williams 464
(RM);S. L. Welsh and E. M. Christensen 6572 (BRY);
W. A. Weber 61 1 1 (ARIZ. COLO. RM, UTC. CS); G.
E. Osterhout 5119 (RM); L. C. Higgins 1048. 1055
(BRY).

y"~

®

N

i J

7

/

/

Map No. 37. Mesa Co., Colorado and Grand Co., Utah.
of C. elala (Eastw.) Payson.

Range

Map No. 38. Southwestern Wyoming, northeastern Utah,
and adjoining Colorado. Range of C sericea (Gray) Payson.

46

liRK.HAM YtniNC; UNIVF.K.SITY SCIKNCK liLlLl.FlIN

Lectotype: H. tngehiuinn s.ii., collected in
Wyoming at Biidger Pass. 1856, GH. vide Paysoii.

Distribution: Southwestern Wyoming, northwest-
ern Colorado, and northeastern Utah. Growing on
heavy clay soils, 4,200 to 7.000 feet. Map No. .«.
Late May to August.

The name sericea has always been a stumbling
block in the way ofany satisfactory treatment ol this
group of plants. Payson (l')27) was faced with the
problem of selecting a type from the collections avail-
able to Dr. Gray at the lime he described scriccci. The
specimens that were considered to compete loi ihc
type of scricca were as follows:

Sheet 1 contains tour specimens:

a. Bridacr's I'ass, 18,S6,

II. l-ngelniann ecjualsO. aiin'iiwa R\dh,

b. Wasatch .Mts., 1844, Iremonl equals (). Iiiiiiiilis C^reene

c. Clover Mis.. Nevada, 1868.

Watson e(|u,ils ^;' //k/(;///\ Cieene

d. Mountaui Hot Springs.
Vellovvstone Park. 1885.

Tweedy 816 ecpuils C a'losioidcs

Sheet 2 contains three specimens;

a. Montana Terr.. 1867 eijuals C. cc/oivo/rfcs

b. Summit. Calirornia, 1871,

liolander eiiualsC iuihii;ciw GrcL'nt:

c. Grass Valley. Utah. 1875.

Ward 49 equals C. aha/a .lohnsl.

Sheet .^ contains two specimens at the present lime .nui
probalily five I including rragmenls) in Gray's time:
a. Southern Montana. 188(1.

Watson 287 equals (' cclosioiJcs

li. A specimen of celosioides

without data ecpials C celosioiJcs

c. Iragment. Baker (ounlv .

Oregon, 1879, Cusiek equals C siihrcliisu .lohnst.

d. I ragment troin southern

Wyoming equals C. cucspilosa

c. Iragment trom Scolls

Bluff. 1858 equals C. cuiui

The Specimens that were considered to compete
for the type of sericea were then O (iriiciiieu. Iiunulis.
C. celosioides. alvtiti. iiiihigena. sut'iriiisu. aicspirosa.
and cana. By a process of cliiniiKition. a type for
scricca was selected- that from Bridger's Pass, col-
lected by Engelmann-as this was the only specimen
that fit the published descriplioii. the maturity of the
plant Dr. Gray had m mind. :iiid ilic geographical
range.

Cryptunllni scricca is simikii in ;ippearance to C
celosioides but can be recognized ;it once by the
silky-strigose ventral surface of the leaves, which hick
pustulate hairs and the differently marked nutlets.

39. Crypiaiillia apcria ( |-;isl\\ .) Payson

Cryplaiulia apcrta (Lastw.) Payson, Ann. Mo. Bot.
Gard. I4:29.S. 1927.

Orcocarva apcria l^aslw. Bull lorrev Bol. Club '0:241
1903.

Caespitose perennial. 1.2-2 dm tall; steins several.

slender, arising from a woody root. 0..V0.5 dm long.

strigose and conspicuously white setose-hispid; leaves

spalulate to oblanceolate. somewhat lolded, and wiih
Ihc niidiib strongly developed, obtuse. 2-3,5 cm long.
0.3-0.(1 cm wide, both surfaces setose-hispid and
pustulate, with fine appressed hairs beneath the
bristles; intlorescence open, branched from near Ihe
base, with simple or two-t'orked spikes. 1-1.3 dm
long, the individual spikes becoming 4-7 cm long,
foliar bracts inconspicuous; caly.x segments linear-
lanceolate, in anthesis 2.8-3 mm long, in fruit becom-
ing 7-9 mm long, densely setose; corolla white, the
tube 2.6-3 mm long, crests at base of lube conspic-
uous, fornices yellow, iiuncate. distinctly papillose,
about 0.5 mm long, limb 4-6 mm wide; style exceed-
ing mature fruit 1.5-2 mm; nutlets ovate-lanceolate,
2-2.6 mm long. 1.4-1.6 mm wide, usually all four
maturing, margins acute, in contact, dorsal surface
indistinctly carinate. tubcrculate, somewhat rugulose.
and indistinctly muricale, ventral surface indistinctly
roughened, scar closed, and without an elevated
margin. Collections: 1 (0); representative: A.
Eastwood s.n. (CAS),

Holotype: A. Eastwood s.n.. collected in Mesa
County. Colorado, at Grand .lunclion. 27 June 1892.
CAS. Photograph at BRY.

Distribution: Known only from the type locality.
Mesa County. Colorado. Probably growing on clay
soil characteristic of the region, 4,000 to 5,500 feet.
Map No. 39. May to July.

This species still remains obscure because of the
lack of herbarium material. In observing the type
specimen it appears that the plant is quite distinct,
with its briiad inflorescence and the ornairientation
on the nutlet. It is perhaps closely related to C.
thyrsijlora. but is entirely distinct.

40. Crypiantlia wcluri Johiist.

Crypkiiillui wclu'n Johnsi. Journ. Am. .-Xib. 33:72.

1952.

Caespitose perennials. 1-1. S dm tall; stems numer-
ous, erect. 0.2-0,5 dm long, strigose, and weakly
setose; leaves numerous, narrowly oblanceolate. 3-8
cm long, 0.3-0.7 cm wide, densely hispid villous, with
pustules on both surfaces; inflorescence cylindrical,
narrow, 0.4-1 dm long, hispid-villous; foliar bracts
evident on lower part of inflorescence; caly.x seg-
ments lanceolate, in anthesis 3-4 mm long, in IVuit
becoming 5-6 mm long, hispid-villous; corolla white,
the tube 3-3.2 mm long, crests at base of tube con-
spicuous, fornices yellow, emarginate, somewhat
papillose, about 0.5 mm long; style exceeding mature
hull i.5-i.S nmi; nutlets ovate. 2-2.3 mm long.
1.3-1.8 mm wide, all four usually maturing, margins
;icute or narrowly winged, dorsal surface tuberculate,
and with short irregular transverse ridges, ventral
surlace nearly smooth, scar open, triangular or
narrowly cuneate, elevated m;irgiii Licking. Collec-
tions: II (\); representative: \\. ,A. Weber 5778

BIOLOCICAl. SKRIKS VOL. 13, NO. 4 A Ki:VlS10N Ol CK Y I' 1 AN I>:a SUBGKNUS ORKOCARYA

47

N

® r

J

K 1 l_
1 \

/f

■~^1

1 ■%

\ \ \ ®

i \

\ 1 1— 1 (
III )

II ' '

1 1 ' '

1 ' >

If 1 /

1 ' ' 1 /

ill 1 /

^ ^ 1 /

Map No. 39. Mesa
(Eastw.) Payson.

County, Colorado. Range of C apcna

Map No. 40. Central Colorado, Saguache and Hinsdale
Counties. Range of C irc-fctr; Johnston.

(t OLO. Gil, LL); J. H. Langeiiheini 4047 (RM); H.
Gentry 2405 (ARIZ); J. Barrell 92-55 (CS); S, A.
Spongberg 62-55 (CS); W. A. Weber ^41 1 (UT); L. C.
Higgins 2256, 2268. 2269. 3719. 3727 (BRY,
\vf$L).

Holotype: W. A. Weber 5778. collected in
Saguache County. Colorado, along road to Stone
Cellar Ranger Station and Saguache Park, near junc-
tion of main highway. 4 miles west of Cochetopa
Pass, volcanic ash deposit. 9,700 feet, 28 July 1950,
GH. Photograph at BRY. Isotypes at COLO. LL.

Distribution: Saguache and Hinsdale counties.
Colorado. Growing on volcanic ash deposits. 9.000 to
10.500 feet. Map No. 40. July and August.

This delicate little Ciyptantha from the high
mountains of Colorado is one of the most distinct in
the entire subgenus and is not confused with any
other species because of the narrow inflorescence,
pubescence, and the very distinctive nutlets, it keys
out in Payson's monograph to C nigiilosci. but is only
remotely related to that species.

41 . Cryptiiiuha nigiilosa (Payson) Payson

Crvpiaiiiha nigiilosa (Payson) Payson. Ann. Mo. Bot.

Gard. 14:295. 1927.

Oreocarva ntmlosa Payson. L'niv. Wvo. Publ. iiol. 1:166.
1926.

Biennial or short-lived perennial. 1 .2-.^i iliii tall:

stems slender. 1 -several. 0.8-1.6 dm long, spreading
setose-hispid; leaves oblanceolate to spatulate. obtuse
to acute, strigose and conspicuously setose-hispid,
pustulate on both surfaces; inflorescence 0.2-2 dm
long, hispid; foliar bracts inconspicuous; calyx seg-
ments linear-lanceolate, in anthesis 4-5 mm long, in
fruit becoming 7-9 mm long, strigose and spreading
hirsute; corolla white, the tube 3-4 mm long, crests at
base of tube conspicuous, fornices rounded, distinctly
papillose, about 0.5 mm long, limb 5-7 mm wide;
style exceeding mature fruit 1-1.5 mm; nutlets lance-
olate, 2.8-3.2 mm long, 1.3-1.7 mm wide, all four
usually maturing, margins in contact, acute, dorsal
surface with short low ridges, also somewhat tubercu-
late, ventral surface smooth or nearly so, scar open,
subulate, without an elevated margin. Collections: 30
(vii); representative: B. Maguire 22021 (ARIZ, UTC);
Maguire and Becraft 2729 (RM, UTC); W. P. Coltom
9569 (UT); B. F. Harrison 1 1658 (BRY); M. E. Jones
s.n. (POM, RM); L. C. Higgins 1463, 1474. I hi 4,
1621, 1720 (BRY).

Holotype: M. E. Jones s.n.. collected m Juab
County. Utah, at Fish Springs. 4 June 1891. RM.
Photograph at BRY. Isotypes a^t UC, POM.

Distribution: Central Utali to northeastern
Nevada. Growing in clay or gravelly loam soils. 4.500
to 6,500 feel. Map No, 41 , May to July.

Crypiaiuha nigiilosa is closely relaled to C. spicu-

48

HRIC.IIAM YOUNG UNIVKKSl I Y HCIKNCi: liULLIIlN

///era ;irKl C. iiiicmipui. bul differs in llic very setose-
hispid iiidument, differenlly muiked nutlets wiiicli
arc snioolii on the ventral surface, and different
geographical range.

42. Crypiantlia tnicrnipia (Greene) Payson

CrvptantUa intcrrupta (Greene) Payson. Ann. Mo.

Bot.Gard. 14:246. 1927.

Orcocarya iiilerrupla Greene. Pilt. .1:111. 1896.

Long-lived perennial, 1.7-6 dm tall; stems few to
several, slender, 1-3.5 dm long, strigose and weakly
setose with slender white hairs; leaves oblanceolate to
spatulale, obtuse, 1.5-7 cm long. 0.4-1.2 cm wide,
dorsal surface densely strigose, and appressed setose
pustulate, upper surface more finely strigose, setose
hairs less conspicuous, pustules fewer; intlorescence
narrow, interrupted, 1-2.5 dm long, densely setose,
cymes somewhat elongating at the top, foliar bracts
evident on lower pari of stem; calyx segments lance-
olate, 2-3 mm long in anthesis, in fruit becoming 5-(S
mm long, setose; corolla white, the tube 2-2.5 mm
long, crests at base of tube very conspicuous, fornices
light yellow, slightly emarginate, about 0.5 mm long,
limb 5-6 mm wide; style exceeding mature fruit by
less than I mm; nutlets lanceolate, 3.3-3.6 mm long,
1.7-2 mm wide, all four usually maturing, margins in
contact, acute, bolh surfaces tuberculate with scat-
tered, rounded tubercles, or sometimes nearly
smooth, scar slightly open, linear, margin not ele-
vated. Collectit)ns: 10 (iv); representative: A. A.
Heller 9185 (NY, PH, RM, US); Hitchcock 1005, 929
(US); R. C. Rollins 2542 (UTC); Gentry and Davidse
1824 (BRY. NY); L. C. Higgins 1721, 1724 (BRY).

Neotype: L. C. Higgins 1721, collected about 8
miles east of Wells, biko County, Nevada. 13 July
1968, BRY.

Distribution: .Apparently endemic to Elko County,
Nevada. Growing on clay soil, in the pinyon-juniper
community, 5,000 to 7.500 feet. Map No. 42. June
to August.

Dr. Greene stated that he had not seen this plant
in any herbarium, but it was collected by him m ihe
moiuitains of eastern Nevada. He also said, "it
abounds in open woods some miles east of Wells." In
an attempt to locate this collection of Greene's from
east of Wells, no specimen could be found which fit
his description. Payson also stated that he was unable
to locate the specimen. This type specimen is ap-
parently lost or has been destroyed; as a result a
neotype has been selected, Higgins 1 721 .

This distinctive plant is most closely related to C.
spiculifera. bul differs in the shorter style, tubercu-
late nutlets, and the longer stems. In a collection by
Gentry and Davidse 1824, from Harrison Pass the
nutlets were almost smooth; otherwise the plant was
quite the same.

Map No. 41. Western Utah and adjoining Nevada. Range of
C. nigulosa (Payson) Payson.

. —

N

j ®

• ••

1 1

1
1 1

1 \
1 \

i \ •

1 \

N.

Map No. 42. Northeastern Nevada. Range of C. mlcintpta
((ireene) Paxson.

BIOLOC.It'AL SI:RII:S vol . 13. NO. 4 A Kl-VISION Ol- (.'R Yl' lANTHA SU BGKN US ORF.GCARY A

49

4.V Crvpiiiiulni spiciilifera (I'ipoi) I'liyson

Crvphinilhi spividifcra (Piper) Payson, Ann. Mo. Bot.

Gard. 14;:4X. l'):7.

Oieocarva spiculifaa Ilpci. Contr. U.S. Natl. Herb.

11:481. 1906.
Orcocarva cilioliirxtiia Ncls. & Macbr. Bot. (ia/.. 55:.i7K.

1913. (Type: Nelson and Macbride 1799. Minidoka.

Idaho. 23 .lunc 1912.)

Perennial, 1.5-3 dm tall; stems 1-several, 1 .2-2 dm
lung, strigose, and spreading setose; leaves oblanceo-
laie. acute to obtuse, 2-7 cm long, 0.3-0.6 cm wide,
dorsal surface densely strigose and spreading setose,
pustulate, ventral surface similar, petioles conspic-
ui)usly setose ciliate-margined; inflorescence narrow,
0.4-1.5 dm long, foliar bracts inconspicuous; calyx
segments linear-lanceolate, in anthesis 4-5 mm long,
in fruit becoming 6-10 mm long, strigose and spread-
ing setose; corolla white, the tube 3-4 mm long, crests
at base of tube conspicuous, fornices yellow,
rounded, nearly glabrous, about 0.5 mm long, limb
5-9 mm wide; style exceeding mature fruit 1 .5-2 mm;
nutlets lanceolate to ovate-lanceolate. 2.S-4 mm long.
1.5-2.2 mm wide, one to four nutlets maturing,
margins acute, in contact, dorsal surface rugulose, or
tuberculate, and with some inconspicuous murica-
tions, scar subulate, the margin not elevated. Collec-
lions: 65 (ii); representative: A. Cronquist 2396 (GH,
ins, UTC); J. S. Cotton 359 (WTU); R. J. Davis 3803
(GH, IDS); J. F. Macbride 875 (GH,RM,LITC);M. E.
Peck 19918 (GH, ORE); Sandberg and Leiberg 164
(GH); Maguire and Holmgren 26223 (GH, UTC,
WTU); L. C. Higgins 1635. 1636 (BRY).

Holotype: J. H. Sandberg and J. B. Leiberg 164,
collected in Adams County, Washington, at Ritzville,
6 June 1893, US. Photograph at BRY. Isotypes at
UC. GH. WTU.

Distribution: Southeastern Washington to south-
ern Idaho and eastern Oregon. Growing on sandy or
clay soils. 1.500 to 7.000 feet. Map No. 43. May to
July.

Cryptanllia spiciilifera is a fairly well defined spe-
cies characterized by narrow radial leaves with their
ciliate-margined petioles. In various parts of its range,
it may be confused with C. intcmipla. C. humilis. or
C. celosioides. It differs from C. intermpta by the
narrower leaves, rugulose nutlets, and the longer
style. It can be distinguished from C humilis by its
taller habit, rugose nutlets which are not at all muri-
cate, and the ditTerent pubescence. From C. celo-
sioides it ditl'ers in the narrower leaves, a stronger
tendency to a multiple caudex, somewhat ditTerent
nutlets, and a more southerly range.

The plant described as C. cilio-hirsuta by Nelson
and Macbride is almost identical to the type of C.
spiciilifera, and so is placed in synonymy under that
species.

44. Crypuiiulni sliackkilcuna Higgins
Cryptaiiiha sliackkiieana Higgins. Great Basin Natur-

alist 29:28-30. 1969.

Caespitose perennial herb, 1-3 dm tall; stems
slender, weak, 1 -several, 0.7-1 .8 dm long, strigose and
spreading setose with slender weak hairs; leaves linear,
2-13 cm long, 0.1-0.5 cm wide, strigose on both sur-
faces, and with a few inconspicuous pustulate hairs
on the dorsal surface; inflorescence narrow, nearly
capitate. 0.2-0.8 dm long; calyx segments linear or
narrowly lanceolate, in anthesis 3-5 mm long, in fruit
becoming 7-10 mm long, setose, with yellowish
spreading hairs; corolla white, the tube 3-i3 mm
long, crests at base of tube evident, fornices yellow,
emarginate, 0.5 mm long, limb 5-6 mm wide; style
exceeding mature fruit 1.3-1.6 mm; nutlets lance-
olate, 3.3-3.6 mm long, 1.6-2 mm wide, usually all
four maturing, the margins acute, in contact, dorsal
surface muricate and rugulose with low inconspicuous
ridges, the ventral surface similar but the markings
much less evident, scar open, subulate, and without
an elevated margin. Collections: 2 (0); representative:
H. T. Shacklette 6183 (US): Welsh and Moore 8629
(BRY).

Holotype: S. L. Welsh and G. Moore 8629. col-
lected in Alaska, on steep south-facing slope of Eagle
Bluff, about 1 mile northwest of Eagle. Growing with
Artemisia frigida and Agroprron smiiliii. 26 July
1968. BRY.

Cryplautha shackleiieaua is probably closely re-
lated to C. spiciilifera but differs in the longer and
narrower leaves with only inconspicuous pustulate
hairs, the more capitate inflorescence, more elongated
nutlets with less evident markings, and weaker stems.

45. Cryptaiitha celosioides ( Eastw.) Payson

Crvptaiitlia celosioides (Eastw.) Payson. Ann. Mo.

Bot. Gard. 14:299. 1927.

Cynoglossoni gloniei-atiini Nutt. c\ Pursh. I'l. Am. Sept.

2:729. 1814. not Crvplaiuha };lonicra!a Lelini.
Myosotis glomerata Niiit. Gen. Pi. 1:112. 1818.
Roclielia glomerata Torr. Ann. Lye. N.Y. 2:226. 1827.
F.iinichium glomeratum A. DC. Prod. 10:131. 1846.
Krviiilzkia g,lomerala Gray. Proe. Am. Aead. 20:279.

1885.
Orcocarva glomerata Gimne.VM. 1:58. 1887.
Oreocarya affinis Greene. Pitt. 3:1 10. 1896. (Type: Sand

hills near Red Biittes. Wyoming. 5 .luly 1896. Greene.)

not O. affinis (Gray) Greene.
Orcocarva cclosoiodcs Fastw. Bull. Torrey Bot. Club

30:240. 1903. (Type: T. J. Howell, eollceted on the

banks ot the Columbia River, eastern Washincton

Territory. July 1881.)
Krvititzkia piistiilata Blankenslnp, Mont. .Agt. Coll. Stud.

Bot. 1:96. 1905. (Greene. Red Buttes. Wyoming. 5

.luly 1896.) not C. inisliilata (Rydb.) Payson.
Orcocarva sericea sensii Piper. Contr. U.S. Natl. Herb.

1 1 :482. 1906, not C sericea (Gray) Payson.
Orcocarva macounii E^astw. Bull. Torrey Bot. Club

40:480. 1913. (Type: Moose Mt. Creek, Saskatche-
wan. 6 .luly 1880. .lohn Maeoun.)
Oreocarya pcrcnnis Rydb. II. Roeky M(s. 722. 1917. in

part, not (). affinis pcrcnnis A. Nels.
Oreocarya shelcionii Brand, I edde, Rep. Spec. Nov.

19:73. 1923. (Type: Deep Creek. Wallowa County.

Orcjion. 16 June 1897. Sheldon 8315.)

50

HKH.IIAM YUUNC. UNI VKKSI lY SCILNCIl BULLEIIN

Map No. 43.
C spiciilifcra

Parts of northwestern United States. Range of
(Piper) Payson.

Map No. 44. .Alaska. RantteofC. shacklcttcaiia Higjiins

Crvplaitrlia shcldonii (Brand) Pavson, Ann. Mo. Hot.

'Card. 14:301. 1927.
Crvplaiitlia nmcoiinii (l.aslw.) Pavson. .\nn. Mo. Mot.

■ Card. 14:303. 1927.
Cryplanlha bradhuriaim Payson. .Ann. Mo. liol. (Jard.
14:307. 1927.

Biennial or short-lived perennial, with or without a
brancheiJ cau(Jex; stems l-several. l-() dm tall, olien
relatively robtist. strigose. sett)se-hirsiite, and sub-
lomentose with pustulate bristles; leaves oblanceolate
to spatulate, obtuse to acute, 2.5-9 cm long. 0.4-1.5
cm wide, strigose. setose, and subtomentose. pustu-
late on both surfaces; intlorescence narrow to open
and very broad. l-.^.X dm long, setose-hirsute; calyx
segments lanceolate. 3-(i mm long in anlhesis. in fruit
becoming 7-1 2 mm long, setose-hirsute; corolla white,
the tube .V7 mm long, crests at base of tube evident,
fornices yellow, rounded or emarginale. about 0.5
mm long, limb 7-1 2 mm wide; style exceeding mature
liuil 1 .5-6 mm; nutlets lanceolate to ovate-lanceolate.
2.S-5 mm long. 1 .5-2.(i mm wide, two to four matur-
ing, margins acute or narrowly winged, in contact,
dorsal surface tuberculate to deeply and sharply
rugose, sometimes papillose between the markings,
ventral surface similar but the markings less evident,
scar closed or narrowly open at the base, elevated
margin lacking. Collections: 457 (xxiv); represen-
tative: A. Nelson 1956 (ND-G. RM); M. Ownbey
104;. (IDS. RM);C. L. Hitchcock 17955 ( RM. WTU);
W. H. Booth 55110 (MONT. RM);P. A. Rydbeigand
E. A. Bessey 4883 (MONT. RM); C. L. Porter 7752
(RM, WTLl); Hitchcock and Muhlick 12553 (RM,
UTC. WTLl); Maguire and Holmgren 26615 (UC,
CAS, IDS, UTC, WTU); A. R. Kruckeberg 2197 (UC,
CAS. ORE. UTC. WTU); T. J. Howell s.n. (UC. CAS);
.1. W. Thompson 11696 (MONT. US, WTU); H. T.
Rogers 613 (UC. CAS. UTC); L. C. Higgins 1535.
1538. 1558. 1689. 1693. 1697. 1701. 1704 (BRY).

Lectotype: Bradbury s.n.. collected in Upper
Louisiana, supposed to be about the Big Bend of the
Missouri in what is now South Dakota, PH. Photo-
graph at BRY.

Distribution: Eastern Oregon from Grant County
to northern Washington and southern British Colum-
bia, east through the lower parts of northern Idaho to
Montana, thence south and east into North Dakota,
Nebraska, and Colorado. Growing on dry open slopes
and valleys, plains and foothills, occasionally ascend-
ing to moderate elevations in the mountains, 1,500 to
8.500 feel. Map No. 45. May to August.

Cryphnnlui celosioidcs is a very widespread and
polymorphic species with a great amount of variation
throughout its range. The species C. hradhuriana.
shcldonii. luacoitnii, and celosioidcs as recognized by
Payson in his monograph have here been combined.
The basis for this wholesale combining of species is
that no differential characters of high enough magni-
tude exist lo separate out species or even varieties
with an\' consistency. The consistent characters used

BIOLOGICAL SlRll'S VOL. 13. NO. 4 A KL;VISION OLC'R Y P LAN I'L' A SUBGENUS 0RL:0CAF<Y A

51

Map No. 45. Parts of nortli\\estcin L nitcd States and adjuiniiig Canada. Range of C. celosioidcs (Eastw.) Payson.

to distinguish species throughout this subgenus seem
to entirely break down in the present case; so until
more information can be gleened and utilized, this
comple.x is best treated as a single taxon, even though
this is not entirely satisfactory.

This species is distinguished by its setose indu-
ment, large corollas, habit, and characteristic inflor-
escence.

46. Crypiamha tlionipsoiiii Johnst.

Cryptaiitlhi iluniipsitiui Johnst. Contr. Am. Arb.

3:88. 1932.

Orcocarva thomiKonii (Johnst.) Abranis. Ahrams IH. I L
Pae.St. 3:600. 1951.

Caespitose perennials. 1.5-3 dm tall; stems several
from a woody caudex. 1-1.7 dm long, setose-hirsute;
leaves oblanceolaie. acute to obtuse, 4-8 cm long,
0.4-0.8 cm wide, yellowish tomentose and with scat-
tered appressed setose bristles on both surfaces; in-
tlorescence usually narrow. 0.2-1.3 dm long, foliar
bracts evident to conspicuous; calyx segments lance-
olate, in anthesis 3.5-4.5 mm long, in fruit becoming
9-12 mm long, setose; corolla white, the tube 3-4 mm
long, crests at base of tube evident, fornices yellow.

low, rounded or emarginate, papillose, limb 6-8 mm
wide; style exceeding mature fruit 1-2 mm; nutlets
lanceolate, 4-5 mm long, 2-3 mm wide, all four us-
ually maturing, margins narrowly winged or knifelike,
in contact, dorsal surface tuberculate and irregularly
rugose, ventral surface smooth or slightly uneven, scar
open, cuneate, and without an elevated margin. Col-
lections: 25 (ii); representative; J. W. Thompson
8742 (GH, WTU); A. R. Kruckeberg 2750 (ORE,
WTU); W. W. Canby ^^h (UC); L. C. Higgins 166 1,
1667 (BRY).

Holotype: J. W. Thompson 7663, collected in
Kittitas County, Washington, on the crest of Iron
Moimtain, Mount Stuart Region, 1930, GH.

Distribution: Kittitas and Chelan counties in the
Wenatchee Mountains of central Washington. Grow-
ing on sleep talus slopes. 3,000 to 7,000 feet. Map
No. 46. Late May to August.

Cryptantha thompsonii is a loosely caespitose
species arising from a thick, very woody taproot. It is
endemic to the higlT mountains of east-central
Washington. The plant may be distinguished by its
thick leaves, nutlets which are smooth on the ventral
surface, and Ihe scar which is evidently open for most

liKK;HAM YOUNG UNIVtKSI lY SCltNCl. BULLETIN

of its length. A very distinct species not to be con-
fused witii any other in the region wiiere it grows.

47. Cryptaniha sohollfcra Payson

Crvptantha sobolifera Payson. Ann. Mo. Bot. Card.
' 14:305. 1927.

Long-Hved perennials. 1-1. S dm tall; stems I-
several from the branched caiidex. some of them
sterile and prostrate and terminating in soboles,
0.5-0.7 dm long; leaves spatulate to oblanceolate,
obtuse, 1.5-4 cm long, 0.5-O.X cm wide, strigose, and
setose-hirsute, also somewhat tomentose, pustules
conspicuous on both surfaces, petioles ciliate-
margined; inflorescence cylindric, narrow, 0..V0.8 dm
long, setose, foliar bracts inconspicuous; caly.x seg-
ments linear-lanceolate, in anthesis 3-5 mm long, in
fruit becoming 6-7 mm long, strigose and conspicu-
ously setose; corolla white, the tube 3.5-4 mm long,
crests at base of tube well developed, fornices yellow,
emarginate, distinctly papillose, about I mm long,
limb 6-8 mm wide; style exceeding mature fruit
1.8-2.5 mm; nutlets ovate-lanceolate. 2.5-3.5 mm
long, 1.4-1.7 mm wide, dorsal surface with low in-
conspicuous tubercles or ridges, or sometimes almost
smooth, ventral surface smooth or nearly so, scar
straight, closed, the margin not elevated. Collections:
3 (0); representative: M. E. Jones s.n. (POM); Pennell,
Cotner and Schaeffer 23928 (US): Hitchcock and
Muhlick 13021 (RM).

llolotype: M. I:. Jones s.n.. collected in Glacier
County, Montana, at Upper Marias Pass, 10 .Sep-
tember 1909, POM. Photograph a I BRY.

Distribution: Western Montana in the high Rocky
Mountains. Growing on sandy soil or serpentine lalus
slopes, 5,000 to 10,000 teet. Map no. 47. Late June
to September.

This species is not very well known or represented
by herbarium specimens, and more collections of it
are badly needed. It is perhaps most closely related to
C. cclosiokles. but differs in the smooth ventral
surface of the nutlets, the soboliferous leaves, and
different tToweriiig lime.

48. Crvpiuiulki griiliainii Jolmst.

Crypianiha graluiniii Johnsl. Jourii. Arn. Arb.

20:391. 1939.

Long-lived perennial from a thick woiidy taproot.
1.5-2 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, conspicuously setose-
pustulate on both surfaces, with some finer pubes-
cence beneath; inllorescence narrow, 0.4-1 dm long,
setose, foliar bracts evident but not conspicuous;
calyx segments lanceolate, in anthesis 5-7 mm long, in
fruit becoming 7-9 mm long, abundantly setose;

Map No. 46. Central Washington, Kittitas and Chelan
Counties. Range of C Ihompsonii .lohnston.

®

J

J \

--i
I

I I

Map No. 47. Glacier County, Montana. Range of C. sobo-
lifera Pavson.

BIOLUt.R AL SlRIIiS VOL. 1 J, NO. 4 ,.\ RKVISION OI LKYHIAN IKA SUHGENUS OREOCARYA

53

Map No.
Johnston

48. Uintuli Comity. Utah. Range of C. grahaiiiti

corolki white, the tube 3.5-5 tntn long, constricted at
llie tiiiddie. crests at base ot" lube evidetit. i'ornices
yellow, eniarginate, papillose, 0.5-1 mtn long, limb
11-15 mm wide; style coarse, exceeding mature fruit
1.8-2.1 mm; nutlets lanceolate. 3-3.8 mm long, 1.7-2
mm wide, two to four maturing, margins in contact,
acute, both surfaces of nutlet with inconspicuous
small, low rounded tubercles, or some t)f these
confluent into short irregular ridges, scar straight.
open, narrowly linear, the margin not elevated. Col-
lections; 17 (.\); representative: R. C. Rollins 1707
(GH. RM); E. H. Graham 7924 (GH); Holmgren and
Reveal 1879 (BRY. LITC); R. C. Rollins 1716 (GH);
t;. H. Graham 8Mb2 (GH); L. C. Higgins 1602, 1607,
1610, 1876. 1885 (BRY).

Holotype; Edward H. Graham 7^24. collected in
I'iniah County, Utah, on bench west of Green River
north of mouth of Sand Wash, 4,500 feet, 28 May
1933. GH. Photograph at BRY.

Distribution; Endemic to the Uintah Basin, Uintah
County, Utah, along Willow Creek and the Willow
Creep drainage basin. Growing on while shale. 4,300
to 6,000 feet. Map No. 48. May to June.

Cryptantha grahaniii can be distinguished from
other species of the Uintah Basin by its large corolla,
tuberculate nutlets, coarse style, and the thick, black,
woody caudex.

49. CrypUtutha rciUiusii Johnst.

Cryptantha rolliiisii .lohnst. .iourn. Arn. Arb. 20;391.

1939.

Biennial herbs 1-3.5 dm talhstems I -several, 0.2-1
dm long, setose; leaves clustered at the base, gradually
reduced upward, oblanceolate to spatulate, obtuse to
acule, 2-5 cm long, 0.5-1.5 cm wide, setose and
hispid, pustulate on both surfaces; inflorescence
narrow to somewhat open at maturity, cylindric to
obovoid, racemes in dense glomerules, three to six
flowered, hispid, 0.5-2 dm long; calyx segments
linear, in anlhesis 7-8 mm long, in fruit becoming
8-10 turn long, hispid; corolla while, campanulate, the
tube 7-9 mm long, crests at base of tube evident,
fornices yellow, papillose, about 0.5-1 mm long, limb
7-8 mm wide; plants slightly heterostyled; nutlets
lanceolate, 3-4 miu long, 1-1.5 mm wide, obscurely
rugulose and tuberculate on the dorsal surface, ven-
tral surface smooth, scar closed, and without an
elevated margin. Collections; 42 (xii); representative:
R. C. Rollins 3084 (GH, RM); R. C. Rollins 1715
(GH); Ripley and Barneby 7804 (GH); E. H. Graham
7870 (GH); B. F. Hariison 400H (BRY, GH); J.
Brotherson 1049 (BRY); D. Atwood 1617 (BRY);
Welsh and Christensen 6622 (BRY); L. C. Higgins

Map
Jolin

Ihcrn Utah.

' of C roUinsii

54

IJRICHAM YOUNG UNIVKRSIIY SCIKNCK HULLKIIN

1056. 1068. 1324. 1606. I880(BRY).

Hololype: Reed C. Rollins 1715. collccicd in
Uintah County, Utah, shale hillside on Thome's
Ranch near Willow Creek, 22 miles south of Ouray,
5.500 feet. 16 June 1^).^7, GH. Photograph at BRY.

Distribution; Central and northeastern Utah in
Emery, Uintah, and Duchesne counties. Growing on
white or red shale, 4,000 to 6.000 feel. Map No. 4').
May to July.

Cryptanilia mllliisil is very connnon in the Uniijh
Basin, and can be found on most shaley hillsides
throughout the region. An isolated population also
occurs in Emery County on the south end ot the San
Rafael .Swell near Temple Mountain, but is un-
doubtedly the same species.

This distinctive plant may be recognized by iis
campanulate corolla, nearly smooth nutlets, non-
caespitose habit, and single stem.

50. Cryptiiinltci wvilicnllii (Hastw.) Payson

Crvptaiuha wcllwrillii (Eastw.) Payson. Ann. Mo.

Bot.Gard. 14:324. 1^)27.

Krvniizkia gtowcrala var. aaila Jones. Zoc 2:250. 1S41.
'(Type: Ciseo. Utah. 2 May 189(1. Jones.)

Oreocarva welhcrillii llastw. Bull. Torrey Bot. Club
30:242. 1903.

Biennial or short4ived perennials. 1-3.5 dm tall;
stems 1-6. 0.5-0.8 dm long, branched from the base
with one stout stem and usually several low slender
stems; leaves clustered at the base, gradually reduced
upward, spatulate to broadly oblanceolate. the apices
obtuse to rounded, 2.5-5 cm long, 0.7-1.6 cm wide,
strigose and appressed setose, dorsal surface con-
spicuously pustulate, ventral surface with few or no
pustules; inflorescence becoming broad in age due to
the elongation of the racemes, 0.6-3 dm long; calyx
segments lanceolate, in anthesis 5-7 mm long, in fruit
becoming 7-13 mm long, while setose; corolla while,
the tube 7-10 mm long, crests at base of tube lacking,
fornices light-yellow, emarginate, papillose, about 1
mm long, limb 6-13 mm wide; style exceeding mature
fruit 3-5 mm; nutlets lanceolate or ovate-lanceolale.
3.5-4 mm long. 2-2.5 mm wide, usually all four
maturing, margins acute, in contact, dorsal surface
distinctly luberculate and often rugulose as well as
with numerous murications between the larger rough-
enings. ventral surface similar but the markings not as
distinct, scar open, linear, surrounded by a slightly
elevated margin. Collections: 24 (v); representative:
A. Eastwood s.n. (UC. CAS); M. E. Jones 6734 (\.\C .
GH. RM); B. Magmre 18229 (UC. UTC); W. P.
Cottam 2073 (BRY); G. L. Pyrah (BRY, UTC); Wclsli
and Moore 2786 (BRY); L. C. Iliggms 1476 (BRV)

Lectotype: A. Eastwood s.n., collected in Grand
County. Utah, near Moab. Court House Wash. 25 May
1 842. CAS. Photograph at BRY. Isolype at VK\

Distribution: East central Utah in Grand, Carbon.
Emery, Wayne, and Garfield counties. Usually grow-

ing on heavy clay soils. 4.000 to 6.000 feet. Map No.
50. April to June.

Crvpuiiifliii wctlivrlllii is a close relative ot C.
langiJJuiv. but difters in the shorter corolla tube,
tuberculate nutlets, ventral surface of the leaves with-
out or with only a few pustules, and the (lowers
which are not strongly dimorphic.

5 1 . Crypiantha lungijlura (A. Nels.) Payson

Crvptantha longiflora (A. Nels.) Payson, Ann. Mo.

Bol.Gard. 14:326. 1927.

Oreocarva longiflora A. Nels. F.rythca 7:67. 1899.

Oreocarva liorridula (ireene. PI. Baker. 3:20. 1901.

(Type: Deer Run, Colorado, 1 1 June, on dr\ bank. C.

1 Baker 133.)

Short-lived perennial or possibly biennial. 0.8-3 (5)
dm tall; stems 1 -several. 0.5-1 dm long, setose and
spreading hirsute; leaves spatulate. obovate or
oblanceolate, 2-7 cm long, 0.5-1.5 cm wide, both
surfaces strigose and strongly hirsute, pustulate;
inllorescence broad and open. 0.7-2.5 dm long, se-
tose, foliar bracts inconspicuous; calyx segments
linear-lanceolate, in anthesis 7-10 mm long, in fruit
becoming 10-16 nini long, setose; corolla white, the
lube 12-14 mm long, crests at base of tube lacking,
fornices yellow, emarginate. broad, rounded, papil-
lose. 0.5-1 mm long, limb 9-11 mm wide; style ex-
ceeding mature fruit 4-9 mm (heterostyled); nutlets
lanceolate-ovate. 3-4 mm long. 2.2-2.6 mm wide. 2-4
maturing, both surfaces with tubercles and low
rounded ridges, scar straight, closed or very narrowly
open, with a slightly elevated margin. Collections: 36
(vi); representative;' S. L. Welsh 6989, 6966 (BRY);
R. C. Rollins 2181 (RM. UTC); W. A, Weber 3799
(COLO, UTC); D. Wiens .^061 (BRY. COLO); G. E.
Osterhout 5995 (RM); A. H. Barnum 799 (D1X);C.
F. Baker 133 (ND-G); L. C. Higgins 1478. 3314
(BRY).

Hololype: C. S. Crandall s.n.. collected in Mesa
County. Colorado, at Palisades. 14 May 1898. RM.
Photograph at BRY.

Distribution; Western Colorado and eastern Utah
along the Colorado River drainage. Growing on sandy
to clay soils. 3.800 to 6.000 feet. Map No. 51. May
and June.

This species is perhaps most closely rehited to C.
wclhchllii. but differs in several nolablc respects as
discussed under that species.

Ihe type of Oreocarva liorridula. described by
Greene from Deer Run is the same in every respect as
I his taxon. so is placed in synonymy under ii.

52. Cryptanilia ninin (l-asU\ .) P.iyson

Crvp/aiillhi iciniis (l^aslw.) Pavson. .-Xnn. Mo. Bol.
Gaid. 14:327. 1927.

Oreocarva leina's I-astu Hull lorroN Bol Club 311:244.
190,V

Caespitose perennials. 1.3-2.5 dm tall; sicms

BIOLOGICAL SERIES VOL. 13. NO. 4 A REVISION Ol- CRYI'1 ANTPA SUBCiENUS OREOCAR Y A

55

Map No.
Payson.

50. Eastern Utah. Range of C. wetherillii {Edslw.)

.Map No. 51. Western Colorado and eastern Utah. Range of
C. longiflora (A. Nels.) Payson.

slendei , 1 -many, 0.8-1 .2 dm long, sliigose 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 spreading setose, conspicuously
pustulate, ventral surface uniformly strigose and with-
out pustules; inflorescence narrow, interrupted,
0.6-1.4 dm long, weakly setose, foliar bracts incon-
spicuous; 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 U)ng, crests
at base of tube lacking or sometimes evident, fornices
yellow, broad, emarginate. papillose, about 0.5 mm
long, limb campanulate, 5-8 mm wide: style exceed-
ing mature fruit 3-4 mm; nutlets lanceolate, 3-4 mm
long, 1.8-2 mm wide, all four usually maturing,
margins acute, nearly in contact, dorsal surface cari-
nate, sharply and deeply rugose, ventral surface
rugose, scar open, constricted above the base, and
with an elevated margin. Collections: 40 (vii); repre-
sentative: A. H. Holmgren 3237 (US); R. C. Barneby
13075 (CAS); A. Eastwood s.n. (UC, CAS); B. F.
Harrison 11576 (BRY, UC); A. Cronquist 8978
(UTC); S. L. Welsh 6999. 7053. 7061, 7066 (BRY);
D. Atwood 1541 (BRY); L. C. Higgins 1000. 537,
1281, 1326, 1334 (BRY).

Holotype: Alice Eastwood s.n., collected in Grand
County, Utah, near Moab, in Court House Wash. 25
May 1892, CAS. Photograph at BRY.

Distribution: Southeastern Utah in Emery, Grand,
Wayne, and San Juan counties. Growing on sandy to
clay soils, 2,500 to 5,500 feet. Map No. 52. Late
April to July.

Cryptautha tenuis is often confused with C. oster-
lunitii. biU differs in its taller habit, longer leaves,
larger campanulate corolla, and the longer calyx and
style.

53. Cryptantlia nsrerhoutii (Payson) Payson

Crypiiiiulni ostcrlioutii (Payson) Payson, Ann. Mo.

Bot.Gard. 14:329. 1927,'

Orcocarva osterhoutii Payson. Univ. Wyo. Pubf Bot
1:167. 1926.

Densely caespitose perennials, 0.7-1.2 dm tall;
stems slender, many, arising from the densely
branched multiple caudex, 0.3-0.6 mm long, strigose
and spreading setose; leaves spatulate to oblanceolate,
obtuse, 1-3 cm long, 0.3-0.8 cm wide, dorsal surface
strigose and appressed setose, pustulate, ventral sur-
face strigose, not pustulate or the pustulae incon-
spicuous, petioles ciliate margined; intlorescence
open, 0.3-0.8 dm long, weakly white setose, foliar
bracts inconspicuous; calyx segments lanceolate, in
anthesis 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, crests at base of tube
usually evident but poorly developed, fornices yel-
low, broad, emarginate, papillose, about 0.5 mm long,
limb 5-7 mm wide: stvle exceedins; mature fruit

5 b

HKKIIIXM VOUNC. IINIVI KSirV S( UNCI. lUJLLI IIN

Map No
Payson,

52. Soulheastcin L'tah. Range of C /f;;»/x (l-.astw.)

Map No. 53. Western Colorado and soiilhcaslcrn
Range olC. ostcrhoutii (Payson) Payson.

lltah

0.2-0.7 mm; mulcts hmcculaio, 2.1-}. 2 iniii long,
1.8-2.2 mm wide, usiuilly less lluin t'mii inaluring,
margins obtuse, not in contact, dorsal surface cari-
iiate, sharply tuberculatc and rugose, ventral surface
sharply tuberculate, scar open, constricted above the
base, elevated margin evident but not conspicuous.
Collections: 16 (ii); representative: W. A. Weber 6088
(ARIZ. COLO. CS. RM. UTC):G. E. Osterhout 6138
(COLO. RM): D. Alwood I5.^8A(BRY); B. F. Har-
rison 11923 (BRY): Welsh. Moore and Canter 2946
(BRY); G. Moore 399 (BRY): G. Moore 299 (BRY):
S. L.Welsh 7070 (BRY).

ilolotypc: G. L. Osterhout 6138. ct)llecled in
Monument I'ark. near Grand Junction. Mesa County,
Colorado, 3 June 1921, RM. Photograph at BrS'.
Isotypc at COLO.

Distribution: Mesa County. Colorado, and San
Juan (\)unly, Utah. Growing in sandy soil, 2. .^00 to
6.000 feet. Map No. .53. May to June.

This very distinctive specie is not often collected,
but is one of the most distinct in the entire subgenus
because of its small size.

54. Cryptaiitlia pcirado.xu (A. Nels.) Payson

Crvptantha paivdoxa (A. Nels.) Payson. Ann. Mo.

Bot.Gaid. 14:330. 1927.

Orcocarva paradoxa A. Nels. Bot. Cm. 56:69. 1913.

Oreocarya gypsophila Payson. Bot. Claz. 60:380. 1915.
(Type: On dry gvpsiim hill in Paradox Valle\. Colo-
rado. 18 June '1914. Payson 458.)

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 oblanceolate to
spatulatc. usually folded, obtuse. 1.5-4 cm long,
0.2-0.4 cm wide, dorsal surface with appressed se-
tose-pustulate hairs, ventral surface uniformly strigose
and without pustulate hairs, the petioles ciliate-
niargined; 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 yellow tube 10-12 mm long, crests at
base of tube lacking, fornices yellow, broad, sliglitly
emarginate, papillose, about 0.5 mm long, limb 10-12
(15) mm wide; style e.xceeding matiue fruit 4-9 mm;
nutlets lanceolate, turgid. 2-3 mm long. 1.3-1.6 mm
wide, all four usually maturing, margins acute to
obtuse, not in contact, dorsal smi'ace densely tuber-
culate and conspicuously rugose, ventral surface
tuberculate. also somewhat rugulose. scar open,
constricted below the middle, the margin elevated.
Collections: 16 (vii); representative: E. P. Walker 91
(RM); Payson and Payson 4223 (GH, RM): E. B.
Payson 458 (GH. RM); W. A. Weber 4722 (COLO.
RM. UTC); A. Cronquist 9204 (UTC); B. Maguire
18282 (UTC); B. F. Harrison 9607 (BRY): Higgins
and Reveal 1272 (BRY); L. C. Higgins 1314, 1317.
1319, 1901. 3298 (BRY).

BIOLOGICAL SERIFS VOL. 1.1. NO. 4 .A RKVLSION OL CRYPL.XNTIIA SUBGLNUS OREOCARYA

57

N
f

1 ! c'

•V

• •

c.- J • r

vV--

r , /\

_ J

• • '' 1— ^

- r ; \s

1

•

i • •

/

, •

1 •

1
1

1

Map No. 54, Western Colorailo ami eastern LUah, Range of
C. paradoxa (A. Nels.) Payson.

Map No. 55. Southwestern Colorado, southeastern Utah, and
parts of adjoining states. Range of C. bakeri (Greene) Payson.

Holotype: E. P. Walker ^1, collected in Montrose
County, Colorado, on dry gypsum hills in Parado.\
Valley, 17 June 1^12, RM. Photograph at BRY. Iso-
lypesat GH, POM, US.

Distribution: Western and southwestern Colorado
and Emery County, Utah. Growing on clay or sandy
soil, 4,000 to 7,000 feet. Map No. 54. May and June.

Cryptantha gypsophila is the same plant in every
respect as C. paradoxa. Payson noted, "unfortunately
the specimens of paradoxa had not been mounted
and so were not available when gypsophila was de-
scribed. However, there is no doubt that the two
names are completely synonymous.""

This delicate little species may be distinguished by
its densely caespitose habit, short leaves, and the long
dimorphic corollas.

55. Crypiaiulia hakcri (Greene) Payson

Cryptantha hakcri (Greene) Payson. Ann. Mo. Bot.
Gard. 14:331. 1927.
Oreocarya bakeri Greene, Pitt. 4:92. 1899.
Oreocarva eulnplms Rydb. Bull. Torrey Bot. Club
31:6.37. 1904. (Type: Delores, Colorado, 1892, Cran-
dall s.n.)

Biennial or sln)rt-lived perennials, 1-3 dm tall;
stems 1-4(6). 0.5-1 dm long, spreading setose-hirsute;
leaves oblanceolate, obtuse, mostly basal, 3-6 cm
long, 0.5-1.2 cm wide, dorsal surface strigose and
spreading setose, pustulate, ventral surface uniformly
strigose and with few or no pustulate hairs; inflor-
escence narrow, 0.6-2.5 dm long, setose-hirsute, foliar
bracts evident, slightly surpassing the individual

cymes; caly.x segments broadly lanceolate or ovate, in
anthesis 3.5-4 mm long, in fruit becoming 6-8 mm
long, conspicuously setose; corolla white, the tube
4-6 lum long, crests at base of tube lacking, fornices
yellow, emarginate, 1-1.5 mm long, limb 6-8 mm
wide; style e.xceeding mature fruit 1-2 nun; nutlets
ovate-lanceolate, 2.5-3 mm long, 1.5-2 mm wide,
three to four usually maturing, margins obtuse, nearly
in contact, dorsal surface deeply and sharply rugose,
ventral surface tuberculate and short rugose, scar
closed, sinrounded by a definitely elevated white
margin. Collections: 49 (vi); representative: Baker,
Earle and Tracy 827 (ND-G); R. C. Rollins 2223
(RM, UTC); W. A. Weber 8732 (COLO); Eardman 39
(BR>-, COLO); A. H. Holmgren 3374 (BRY, UTC);
H. M. Schmoll 1281 (COLO. RM); A. Nelson 10408
(RM); D. Atwood 1539A (BRY). L. C. Higgins 1903,
1948, 3558 (BRY, WTSU).

Holotype: Baker, Earle and Tracy 827, collected
on the Mancos River sage plains in southern Colo-
rado, 8 July 1898, ND-G. Photograph at BRY.
Isotype at POM.

Distribution: Southwestern Colorado, northeast-
ern Arizona, and southeastern LItah. Growing on
sandy or clay soils, 4,000 to 8,000 feet. Map No. 55.
May to August.

This species is very distinct, however, very closely
related to C. flavoculata and often confused with it.
it can be distinguished by its leaves which lack pus-
tules on the ventral surf'ace, shorter corolla tube, the
shorter style, and the nutlets which have the scar
tightly closed and the luargin elevated.

58

BKIC.HAM YOIJNC; UNIVESITY SCIENCE BULLETIN

56. Crypiantlui mensaiui (Jones) Payson
Ciyptantlia niciisana (Jones) Payson. Ann. Mo. B^

ot.

Card. 14:33

1927.

Krviiitzkia mensana Jones, Contr. West. Bot. 13:4. 1910.

Oreocarva mensana (Jones) Pavson, Univ. Wyo. Piibl. Bot.
1;17I. 1926.

Short-lived perennials, 1-1.5 dm tall; sienis I-
several, 0.5-1.2 dm long, setose-hirsute, with some
finer strigose hairs beneath; leaves oblanceolale to
spatulate, obtuse, 3-8 cm long. 0.5-1.4 cm wide,
lower surface setose with pustulate hairs, also finely
strigose, ventral side strigose, less setose, and with
fewer pustules; inflorescence broad, open, 0.4-1 .2 dm
long, setose, foliar bracts well developed; calyx seg-
ments lanceolate, in anthesis 4-5 mm long, in fruit
becoming 7-8 mm long, setose-hirsute; corolla white,
the tube 3-4 mm long, crests at base of tube lacking
or nearly so, fornices yellow, rounded, slightly papil-
lose, about 0.5 mm long, limb 5-8 mm wide; style
exceeding mature fruit 1 .5-2 mm; nutlets ovoid, 3-3.5
mm long, 1.6-1.9 mm wide, margins obtuse, not in
contact, dorsal surface rugose, tuberculate and some-
what muricate, ventral surface conspicuously tuber-
culate, scar open, constricted at the middle and
surrounded by a high elevated margin. Collections: 20
(vi); representative: B. F. Harrison 5625 (RM);M. E.
Jones 5445 (POM); S. L. Welsh 6915 (BRY); B.
Maguire 18596 (UTC); A. Nelson 5625 (RM); G. L.
Pvrah 15 (BRY); D. Atwood 1270, 1284 (BRY);
Higgins and Reveal 1298 (BRY); Higgins and Welsh
1043 (BRY); L. C. Higgins 996, 1039, 1318, 3323
(BRY).

llolotype: M. E. Jones 5445p. collected in Emery
County, Utah, 16 May 1894. POM. Photograph al
BRY. isotype at US.

Distribution: Central and eastern Utah in Emery,
Carbon, and Grand counties. Growing on clay soils,
4,500 to 6,500 feet. Map No. 56. Late April to July.

Ciyptantha mensana is closely related to C. Jlaro-
ciilata. but the short corolla and the more open
inflorescence serve to distinguish it from thai species.

57. Crypianiha jlavdviilata (k. Nels.) Payson

Cryptanthu /lavocitlara (A. Nels.) Payson. Ann. Mo.

Bot. Card. 14:334. 1927.

Oreocarva flai'ociilala .\. Nels. Erythea 7:66. 1899.

Oreocarva flavociilata sparulata A. Nels. lirythea 7:67.

1899. (Type: from gravelly hilltops near Evanston,

Wyoming, Nelson 2977, 29 May 1897.)
Oreocarva crislala Eastw. Bull Torrey Bot. Cluli ,^(.1:244.

1903. (Type: Grand Junction. Colorado. 17 Ma\'

1892, Eastwood.)
Oreocarva shocklevi Eastw. Bull. Torrev Bol. Club

30:245. 1903. (Type: Miller Mountain, Esmeralda

County. Nevada, elevation 7.500 feet. Slioekley 244.)
Oreocarva castwoodae Nels. & Kennedy. Mulilenbergia

3:141. 1908. (Type: Mormon Mountains, Lincoln

County, Nevada. P. B. Kennedv and L. N. Gooddini;

146.)

Caespitose perennial, 1-3.7 dm tall; stems 1-
several, slender, 0.5-2 dm long, strigose and spreading

Map No. 56. Central and eastern Utah. Range ofC mensana
(M. E. Jones) Payson.

Map No. 57. Parts of western United Stales. Range of C.
flavoculata (A. Nels.) Payson.

BIOLOGICAL SKRIKS VOL. I .(. NO. 4 A RFVISION ()!■ I'K V I' LAN I 1 ! A SU HC;i;NUS OKHOCAR Y A

59

setose vvitli slender bristles; leaves lineai-oblanceolLile
to spatulate, obtuse to sometimes acute, 3-1 I cm
long. 0.3-1.5 cm wide, densely strigose and weakly
setose, dorsal surface conspicuously pustulate, ventral
surface with few pustules or sometimes silky-strigose;
intlorescence narrow, or sometimes slightly open and
lax, 0.5-3 dm long, foliar bracts evident but not con-
spicuous; calyx segments in anthesis linear-lanceolate.
5-(-) mm long, in fruit becoming 8-10 mm long and
becoming broadly lanceolate to ovate; corolla white
or pale yellow, tube 7-10 mm long, crests at base of
tube lacking, fornices yellow, minutely papillose. 1-2
mm long, limb 8-1 2 mm wide; style exceeding mature
fruit 4-8 mm (heterostyled); nutlets lanceolate to
lance-ovate, 2.5-3.5 mm long. 1.8-2 mm wide, usually
all four maturing, margins obtuse, in contact or
slightly separated, dorsal smface muricate, tuber-
culate, and with conspicuously ridges, sometimes
nearly foveolate, ventral surface tuberculate, rarely
with ridges, scar open, constricted near the middle
and surrounded by a high elevated margin. Collec-
tions: 188 (xix); representative: Maguire and Holm-
gren 26064 (ORE. UTC); I. W. Clokey 7668 (ARIZ.
ORE. LL, UTC); B. Maguire 25234 (ARIZ, BRY.
ORE, UTC); J. Beatley^4007 (BRY. LA); B. F.
Harrison 10320 (BRY, UTC); A. Nelson 4572 (RM);
A. Eastwood s.n. (UC); G. E. Osterhout 6006 (GH.
RM. US); Kennedy and Goodding 146 (RM. US);

Shocklcv 244 (UC) L. C. Higgins 557, W7. 1026.
1001. I'l 12 (BRY); L. C. Higgins 3291. 3324. 3403
(BRY.WTSU).

Holotype; A. Nelson 4572. collected at Piedmont,
Wyoming. 7 June 1898. RM.

Distribution: Southern Wyoming, western Colo-
rado, Utah, Nevada, and southeastern California.
Growing in a wide variety soils. 3,000 to 8.500 feet.
Map No. 57. April to July.

This widespread species may be distinguished by
its long corolla tube, very rugose nutlets with the scar
open and the margin elevated, and the only slightly
heterostyled flowers.

This species possesses a number of different forms,
but they seem to be unworthy of named segregation
t'rom the main specific complex. In western Colorado
the author is familiar with two forms, on the basis of
setose-hispid and silky-strigose indument. For a
limited locality it would seem that these two forms
are worthy of some subspecific rank, but on an exam-
ination of a series of specimens it appears the varia-
tion is only local. Oreocarya cristata has very narrow
leaves and so has a slightly different aspect. On the
same basis of leaf width spaiiilata sliockleyi. and
eastwoodac were described. At the present time the
author can see no difference on which to separate
them.

ACKNOWLEDGMENTS

The author is particularly indebted to Dr. Stanley
Welsh for his suggestion of the problem and his
guidance and interest. Thanks is due my wife tor her
active cooperation. Appreciation is likewise extended
to Dr. James Reveal for many valuable suggestions
and constructive criticism; to Drs. Glen Moore.
Stephen Wood, and Dayna Stocks for critical reading
of the manuscripl. Appreciation is also given to the
Society of Sigma Xi. West Texas State University,

The Kilgore Research Center, and Brigham Young
University for their financial help which made it
possible to do the necessary field work. Special
thanks is given to the curators of the many herbaria
for making specimens available for study. Wliat
authenticity this study may possess is due in large
part to the 7,000 sheets they made available for
study, including the vast majority of types.

60

liKK.HAM YOUNG UNIVKRSITY SCll-NCE HULLKTIN

APPENDIX

Synonyms

The names presented in the following list consist
of the synonyms of the species treated in the fore-
going work. The names in the left colmiin are the
synonyms; those in the cokimn on the right are the
names of the species in the present treatment.

CYNOGLOSSVM

CRYPTA.\T/IA

C. gloineranmi Nutt. o\ Puisli C celosioides (liastw.l

Pays. p.

C. suhreiusa Jolinst. p.
C. celosioides (Eastw.)

Pays. p.

C. inihigeiw (Greene)

Pays. p.

C flava (A. NeK.)

Pays. p.

CRYI'I AMI/A

C. aiijina .lolinsl.
C. bradbuiiaiw Pays.

C. clemensae Pays.

C. confertillora var. flava

(.A. Nets.) Brand
C conferli flora var. lutescens C. flava (A. Nets.)

(Greene) lirand Pays. p.

C. com Jolinst. C. pa/men (Gray) Pays.p.

C. ff/imo/c/c'.v (.tones) Pays. C. fulvocanescens vdr.

echiiwides ( Jones)

C. jainesii var. cincrca

(Greene) Pays.
C. macounii ( Kastw.) Pays.

C. modesta Pay son
C. nana var. commixia

(Macbr.) Pays.
C. nana var. ovina

Pays.
C. nana var. shantzii

(Tidestr.) Pays.
C nana var, naita

C. pnstiilosa (R\db.)

Pays.
C sericea var. perennis

(A. Nels.) Pays.
C. sheldonii (Hr:>nd) Pays.

I RITRICIiUM

E. fiihocanescens Gray

h'. gloineraluni A. DC.

/;'. glomeratum var. fulvo-
canescens S. Wats.
li. glonieraliint var. hispid- C. ihyisijlora (Greene)

issimtini lorr. Pays. p.

t'. ^lomeralinn var. Inimile Gray

C tniniilis var. huniitis p.
E. glomeraliini \:\r. virxaliini C. viigata (.Wirier)

Porter Pays. p.

E. jamcsii Torr. C jamesii var. jamesii p.

£'. leucophaeiim (Dougl.) A. IX".

C /('IKOp/lflffl (Dougl.)
Pays. p.

/^ nnillicaiile lorr. C. jamesii var. mulli-

catilis (Torr.) Pays. p.
E. selosissiino Gray C selosissima (Gray)

Pays. p.

/;. i7/-^a/i/»; Porter C. virgala (¥orWr)

Pays. p

4y

28
50
27
22
22

Is

Higgins

P

42

c.

jamesii var. selosa

(Jones) Jolinst.

P

16

c

celosioides ( taslw .

Payson

P

50

c.

ahata Johnsl.

P

33

c.

Inimilis var. commixia

(Macbr.) Higgins

P

37

c.

Inimilis var. ovina

(Pa\s.) Higgins

P

37

c.

Ininiilis var. shuiuzi

(Tidestr.) Higgins

P

37

c.

Inimilis var. nana

1 lastw.) Higgins

P

37

c

jamesii var. piisiu-

losa (R>db.)

P

IS

c

sericea (Ciray) Pays

P

45

c.

celosioides (Eastw.

Pays.

P

50

C. fulvocanescens var.

fulvocanescens p.

C. celosioides (I'.astw.)

Pays. p.

C fulvocanescens var.

fulvocanescens p.

42

49

42

43

36

30
17

24
15
29
30

IIEMISI'IIAEROCARYA

II. cinerea ((iiecne) lirand C.

II laxa (Macbr.) lirand C

//. oblata (Jones) lirand C.

//. /w//;i['n (Gray) Brand C

//. /)(7iso»;/ (Macbr.) Brand C.

II. ahortiva (Greene) Brand C

//. suffrulicosa (Torr.) Brand C

11. suffrulicosa var. C

miillicaiilis (Torr.) Brand

//. suffrulicosa var. jnislulosa C.

(Rydb.) Brand

H. suffrulicosa var. selosa C.

Jones) Brand

H. suffrulicosa var. lypica C.

KRYMl/.KIA

K. dcprcssa Jones C.

K. echinoides Jones C.

K. fulvocanescens Gray C.

K fulvocanescens var. C.

idahoensis Jones

K. iilomcrala Gray C.

K ,i;lomerala var. acuta C.

Jones

K. iilonierala var. viriiiiiensis C.

Jones

A'. /o»!t's// (Torr.) (iray C.

K. leucopliaeo {Doug\.) Grdv C.

K Icucopliaea var. alala JonesC

K. mullicatilis var. ahortiva C.

(Cireene) Jones

K multicatilis vdr.selosa C.

K mcnsana Jones C

K ohlala Jones C.

K. palnieri Gray C.

K piistulala lilankensliip C.

K. sericea Ciray C.

K. selosissima iGrii\ ) C.

Gray

A'. i»%'ufo (Porter) Gra\ C.

.MYo.soris

M iilomcrala Niit(. C

M. teucoplwca Doiigl. C.

in l.elim.

.17. suffrulicosa Torr. C

OREOCARYA

(). ahortiva Greene C.

() affinis Greene C.

(). affinis perennis .\. Nels C.

O. aiata (Jones) A. Nels. C.

O aperta lastvv. C.

jamesii var. selosa
(Jones) Jolinst. p. 16

jamesii var. laxa
(Macbr.) Pays p

oblata (Jones) Pays p
palmeri (Gray) Pays.p
paysonii (Macbr.)
Johnst. p

jamesii var.
abortiva (Cireene) p
jamesii var. jamesii p
jamesii var.
multicaulis (Torr.) p
jamesii var. piis-
nilosa (Rydb.) p

jamesii var. selosa
(Jones) Jobnst. p

jamesii var. jamesii p

abata Jolinst. p. 33

fulvocanescens var.
echinoides (Jones)
Higgins p. 42

fulvocanescens var.
fulvocanescens p. 42

propria (Nels &
Macbr.) Pays. p. 40

celosioides (Eastw.)
Pays. p. 49

welherillii ( Eastw . )
Pays. p. 54

virginensis (Jones)
Pays. p. 32

jamesii \aT. jamesii p. 17
leucophaea (Dougl.)
Pays. p. 24

confeniflora

(Greene) Pays. p. 21

jamesii var.

a/)0/-nia (Greene) p. 17
jamesii var. selosa
(Jones) Johnst. p. 16

mcnsana (Jones)
Pays. p. 58

oblata (Jones) Pays. p. 19
palmeri (Gray) Pays.p. 18
celosioides ( Eastw . )
Pays. p. 49

sericea (Gray) Pa> s. p. 45
selosissima (Gray)
Pays. " p. 29

virgala (Porter)
Pays. p. 30

celosioides ( lastw ,i
Pays. p. 49

leucophaea ( Dougl.
in t.elm.) Pays. p. 24

jamesii \-M. jamesii p. 17

jamesii var. ahortiva

(Greene) Pays. p. 17

celosioides ( Eastw .)

Payson p. 49

sericea (Ciray) Pays. p. 45

confeniflora

(Greene) Pays. p 21

aperla (Eastw.)

Pays. p. 46

BIOLOC.IC'AL SERIKS VOL. 13, NO. 4 ,\ REVISION Ol fR Y I' IAN I 1 : A .SU HCiKN US OKI'OC AR V.'\

(il

0.
0.

argcntea Rytlb.
bakeri Greene

C.
C.

0.

breviflora Ostcrh.

c.

o.

caespilosa A. Nels.

c.

0.

caiia A. Nels

c.

o.

capitala Ea.stw.

c.

o.

celosioides Eastw.

c.

O. cilio-lursura Nels. & Macbr.C

O. ciiiera Greene

O. cominixia Maebr.

O. confer! itlora Greene

O. crislala Ea.stw.

O. crymophila (Jolinsi.)

Jeps. & Hoover
O. depressa (Jones) Macbr. C abala Jolinst.
O Jisliclw Eastw. C.

scricea (Gray) Pays. p. 45

bakeri (Greene)

Pays. p. 57

breviflora (Osterli.)

Pays. p. 40

caespilosa (A. Nels.i

Pays. p. }4

caiia (A. Nels.)

Pays. p. 40

capitala (Eastw .)

.lolinsl. p. 22

celosioides ( Eastw . )

Pa\'s. p. 44

spiciilifcra (Piper)

Pays. p. 49

jamcsii var. setosa

(Jones) Johnst. p. 16

humilis var. commixta

(Maebr.) Higgms p. 30

confertiflora

(Greene) Pays. p. 2 1

Oava (A. Ne'ls.)

Pays. p. 58

crviuoplula Johnst.

p. 2S
11. 33

O. dolosa Maebr. C.

O. dura Nels. & Macbr. C.

O. easrwoodae Nels. & Kennedv

C.

O. echinoides iionQi) Mdcbr. C

O. elala Eastw.
O. euloplius Rydb.

O. flava A. Nels.

O. flavocidata .\. Nels

O. flavocidata spatulata

A. Nels
O. fulvocanescens iS.W'dts.) C.

Greene
O. glomcrata Greene C.

O gypsophila Pays.

C

O hispida Nels. & Kennedy C.
O. hispidissima (Torr.) R\db. C

O. hispidissima Wooton & Standi,

C.

o.

hoffmanuii (Johnst.)
.Abranis.

C.

o.

horridula Greene

C.

().

huinilis Greene

C.

o.

insolita Maebr.

C

o.

internipta Greene

C.

o.

joiiesiana Pays.

C.

o.

lemmoni Eastw.

C.

o.

leucophaea (Dougl.)
Greene

c.

0.

loiigi flora A. Nels

c

o.

lutea Greene

c

0.

lutescens Greene

c

o.

machridii Brand

c

o.

macounii Eastw.

c.

jamesii var. disticha
(Eastw.) Pays. p. 17

humilis var. shaiuzii
(Tidestr.) Higgins p. 37
tlivrsi flora { Greene )
Pa\s. p. 43

flavoculata

(A. Nels.) Pays. p. 58

humilis var. humilis p. 42

Wfl/fl (Eastw.) Pays. p. 44

bakeri (Greene)

Pays. p. 5 7

Flava (A. Nels.)

Pays. p. 22

flavoculata

(A. Nels.) Pays. p. 58

llavoculala

(A. Nels.) Pays. p. 58

flilvocanescens

(S. Wats.) Pays. p. 42

celosioides (Eastw.)

Pays. p. 49

paradoxa (."X. Nels.)

Pays. p. 56

humilis var. humilis p. 36

thyrsi flora (Greene)

I'ays. p. 43

ohiata (Jones) Pays. p. 19
hoffmaunii Johnst. p. 33

longiflora ( A. Nels.)
Pays. p. 54

humilis (Gray) Pays,
var. humilis p. 36

insolita (Macbr.)
Pays. p. 32

intcrrupta (Greene)
Pays. p. 48

Jonesiana (Pays.)
Pays. p. 43

jamcsii var. setosa
(Jones) Johnst. p. 16

leucophaea (Dougl.)
Pays. p. 24

longiflora (A. Nels.)
Pays. p. 54

confertiflora
(Greene) Pays. p. 21

flava (A. Nels.)
Pays. p. 22

humilis var. humilis p. 3f,
celosioides ( Eastw .)
Pays. p. 49

(). nwnsana iioncs) Pays. C.

<). mouosperma Osterh. C.

O. multicaulis iVoir.) C.

(Jreene

(X mullicaulis var. cinerea C.

(Greene) Macbr.

O. mullicaulis var. laxa C.

Macbr.

(9. nana Eastw. C.

() niiida Greene C

(). nuhigena Greene C

O. oblala (Jones) Macbr. C.

O. oslerhouiii Pays. C.

O. palmcri (Gray) Greene C

O. paradoxa A. Nels C.

O. paysonii Macbr. C.

O. percnnis Rydb. C.

O. procera Osterh. C

O. propria Nels. & Macbr. C.

U pustiilosa Rydb. C.

O. rugulosa Pays. C.

O. salmonensis Nels. & C.

Macbr.

O. sericea (Gray) Greene C.

O sericea sensu Piper C.

O. selosissima (Gray) Greene C.

nwnsuna (Jones)

O. sliantzii Tidestr.

O. sheldonii Brand

O. shocklcyi I-'ast\v.

O. spicala Rydb.

O. spieulifera Piper

O. stricla Osterh.

O. subrelusa (Johnst.) Abrams

C.
O. suffrulicosa (Torr.) C.

Greene
O. suffrulicosa var. aboriiva C.

(Greene) Macbr.
O. suffrulicosa var. cinerea C.

(Greene) Pays.
O. suffrulicosa var. C

mullicaulis (Torr.) Pays
O. tenuis Eastw. C.

O thompsonii iiohw^i.) Abrams

C.
O. ihyrsifloraGr'^ene C.

O. lumuiosa Pays. C.

O urlicacea Wooton & Standl.C

O. r;;;i;«/o (Porter) Greene C

O. virgala forma spicala C.

(Rydb.) Macbr.

O. virgincnsis (Jones) Macbr. C.

O. wetherillii liaslw. C.

ROCHELIA

R glomcrala Torr.

Pays.

P-

58

ihvrsiflora (Greene)

Pays. ■

P-

43

jamesii var. mulli-

caulis (Torr.) Pays.

P-

15

jamesii var. setosa

(Jones) Johnst.

P-

16

jamcsii var. laxa

(Macbr.) Pays.

p.

16

humilis var. nana

(I'astw.) Higgins

P-

37

julvocancscens var.

luhocanescens

P-

42

nuhigena (Greene)

Pays.

P-

27

oblala (Jones) Pays.

P-

19

oslerhouiii (Pa\'s. )

Pays.

P-

55

palmcri (Gray) Pays

■P-

18

paradoxa (A. Nels.)

Pays.

P-

56

paysonii (Macbr.)

Johnst.

P-

19

celosioides (Eastw.)

Pays.

P-

45

sericea (Gray) Pays.

P-

45

propria (Nels. &

Macbr.) Pays.

P-

40

jamcsii var. puslulosa

(Rydb.) Harringt.

P-

18

rugulosa (Pays.)

Pays.

P-

47

salmonensis (Nels.

& Macbr.) Pays.

P-

25

sericea (Gray) Pays.

P-

45

celosioides (Eastw.)

Pays.

P-

49

selosissima (Grav)

Pays.

. P-

29

humilis var. shanlzii

Tidestr.) Higgins

P-

37

celosioides (Eastw.)

Pays.

P-

49

flavoculata (A. Nels

.)

Pays.

P-

58

virgala (Porter)

P-

30

spieulifera (Piper)

Pays. ■

P-

49

stricla (Osterh.)

Pays.

P-

26

subrelusa Johnst.

P-

28

jamcsii var. jamesii

P-

17

jamesii var. aboriiva

(Greene) Pays.

P-

17

jamesii var. setosa

(Jones) Johnst.

P-

16

jamesii var. miilti-

eaulis (Torr.) Pays.

P-

15

tennis (Eastw.)

Pays.

P-

54

ihompsonii Johnst. p. 5 1
ihyrsiflora (Greene) p. 43
tunnilosa Pays.
Pays. p. 3 1

ihvrsiflora (Greene)
Pays. p. 43

virgala tPorti:!) p. 30

virgala (Porter)

p. 30
virgincnsis (Jones)
Pays p. 32

welherilli (l-aslw .)
Pays. p. 52

celosioides ( 1 astw .)
Pays. p. 49

62

BRIGHAM YOUNC UNIVl RSIIY SCltNCE BULLKTIN

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II Ij;' ■■

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645