The Great Basin Naturalist

VOLUME XXVII, 1967

Editor: Vasco M. Tanner
Associate Editor: Stephen L. Wood

Published at Provo, Utah, by
Brigham Young University

TABLE OF CONTENTS
Volume XXVII

Number 1 - April 22, 1967

On Two Little-Known Species of the Earthworm Genus

Diplocardia. G. E. Gates 1

Spiders of the Nevada Test Site. Illustrated. Dorald M.

Allred and D Elden Beck _ 11

Problems of Sampling Desert Arthropods Before and After
a Thermonuclear Cratering Test. Illustrated. Clayton
S. Gist 26

New Species of Bark Beetles (Coleoptera: Scolytidae),
Mostly Mexican. Part VII. Illustrated. Stephen L.
Wood 36

Number 2 - September 5, 1967

A Comparative Study of the Mountain Brush Vegetation in

Utah. Illustrated. Elray S. Nixon 59

Predaceous-Scavenger Ants in Utah. D Elden Beck, Dorald

M. Allred. and William J. Despain 67

New Records and Species of Neotropical Bark Beetles

(Scolytidae: Coleoptera). Stephen L. Wood 79

A New Species of Spinibdella from Utah (Bdellidae: Aca-

rina). Illustrated. Clive D. Jorgensen 98

Notes on Eriogonum - III on the Status of Eriogonum pauci-

florum Pursh. Illustrated. James L. Reveal 102

Number 3 - November 30, 1967

New Records and Species of Neotropical Bark Beetles

(Scolytidae: Coleoptera) Part II. Stephen L. Wood 119

On the Earthworm Fauna of the Great American Desert

and Adjacent Areas. G. E. Gates 142

A New Name for a Utah Lepidium. Illustrated. James

L. Reveal 177

Number 4 - December 30, 1967

Notes on Eriogonum - V. A Revision of the Eriogonum

Corymhosum Complex. Illustrated. James L. Reveal 183

D Elden Beck (1906-1967). Illustrated. Vasco M. Tanner 230

Index 241

II

The

Volume XXVII, No. 1
April 22, 1967

MUS. COMP. ZOOL.
LIBRARY

MAY 3 1 1967

HARVARD
UNIN^RSITY

Great Basin

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: "Vasco M. Tanner, Department of Zoology and Entomology
Brigham Young University, Provo, Utan

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:
J. V, Beck, Bacteriology
C. Lynn Hayward, Zoology
W. Derby Laws, Agronomy
Howard C. Stutz, Botany

WiLMER W. Tanner, Zoology, Chairman of the Board
Stanley Welsh, Botany

Ex officio Members

RUDGER H.

cultur
Ernest L. Olson, Chairman, University Phiblications

RuDGER H. Walker, Dean, College of Biological and Agri-
cultural Sciences

^^S. COMP. Z

The Great Basin Naturalist

Published at Provo, Utah by ^' ^ ^ '^t

Brigham Young University

HARVARD

Volume XXVII April 22, 1967 W6^J'yERSITV

ON TWO LITTLE-KNOWN SPECIES OF THE
EARTHWORM GENUS DIPLOCARDIA'

G. E. Gates-
Few zoologists realize how little is known about the earthworms
of North America. An erroneous assumption that their distributions
in the United States are adequately characterized certainly seems to
have been involved in some recent zoogeographical discussions.

During the last 16 or so years, Professor D Elden Beck of the
Department of Zoology and Entomology, Brigham Young Univer-
sity. Provo. Utah, has been collecting earthworms on his travels
through various states from Maine to California. Those collections
eventually will provide information about some of the many areas
with unknown faunas. A series from a single site provides the basis
for much of this first report on the "Beck" material.

That series is of interest for two reasons: (1) It provides the first
record of an earthworm from the state of New Mexico. (2) The
species is native, not exotic as is each of the three hitherto reported
from the adjacent state of Arizona.

A single specimen of the same species from Wisconsin, under
different circumstances, could have been of equal interest. However,
it is noteworthy that only two megadrile taxa had been reported
from Wisconsin hitherto. Alltolobophora iurgida Eisen, 1874, re-
corded by Ude in 1885, could have been any one or more of three dif-
ferent species. Another. Sparganophilus eiseni Smith. 1885, was
recorded by Hague from the state as long ago as 1923.

No single species was added to the short list of Arizona earth-
worms since 1900. Fortunately, Professor Beck has interested Pro-
fessor T. W. Barrett of Arizona State University. Tempe, in adding
to our limited knowledge of the area. As a result of IVofessor Bar-
rett's early collecting, it is now possible, for the first time, to record
a North American native from Arizona.

Diplocardia verrucosa Ude

1895. Diplocardia verrucosa Ude, Zoologischer Anzeiger, 18:339. (Type locality,
Omaha. Nebraska. Types in the Zool. Mus., Hamburg, Germany.)

1 . I'rom research financed by the National Science Foundation.
1. University of Maine, Orono.

The Great Basin Naturalist
2 G. E. GATES Vol. XXVII. No. 1

1962. Diplocardia verrucosa Murchie, Ohio Journal of Science, 62:185. (Two
subspecies recognized after examination of a '"syntype" and other speci-
mens. )

Habitat. — New Mexico (Hidalgo County). Rodeo, six inches
below the surface in sticky, black, gumbo-like soil under a rank
growth of Russian thistle, February 27, 1966, 1 (+ 16 mostly juve-
nile and 10 fragments?) -0-30. D E. Beck.

Wisconsin (Dane County). Between Arena and Mazomanie,
damp, willow, swamp loam. September 22, 1963, 0-0-1. Christina
Hobby.

External Characteristics. — Size, 80 by 4 mm. (Wisconsin),
to 107 by 3 mm. (New Mexico, but softened). Undisturbed speci-
mens in the soil, according to Professor Beck, reached a length of
six inches. Segments, 118 (Wisconsin). Color, white, clitellum yel-
lowish brown. Prostomium, epilobous; tongue narrowing to a point,
or open, with a transverse furrow slightly behind anterior margin
of the peristomium. Secondary annulation, one presetal and one
postsetal secondary furrow present per segment beginning with v or
vi. Setae, small, retracted; follicles not visible in coelomic cavities,
AB ca. = CD, little difference in sizes of BC and A A or BC <AA
(Wisconsin), DD ca. = V2 C. a,b/xix, xxi penial, a,^/xviii lacking.
Nephropores, inconspicuous whenever distinguishable at or slightly
above D and so in a single rank on each side of the body. First dorsal
pore, at 9/10 (1 specimen). 10/11 (9, including Wisconsin),
?11/12 (2). 11/12 (3).

Spermathecal pores, transverse slits (New Mexico), small, but ob-
viously larger than the other genital apertures, superficial, at or
close to A, just behind intersegmental furrows 7/8 and 8/9. Centered
around each spermathecal pore (New Mexico) is a small circular
area of slight tumescence. Female pores, paired, anteromedian to a,
in some New Mexico specimens within a single small field lacking
clitellar coloration. Seminal grooves, broad and shallow, nearly
straight, in AB between eq/xix and eq/xxi. Male pores, each on a
tiny spheroidal protuberance from bottom of seminal groove, at or
immedately behind probable level of 19/20. Two follicle apertures
are present at each end of a seminal groove, sometimes definitely
not within the grooves. The body wall seems to be slightly thick-
ened ventrally in xix-xxi, but aside from the grooves there is no
specially characterized male field (New Mexico). The ventral
region between grooves of the Wisconsin worm is slightly timies-
cent and after staining seemed to contain three pairs of slight cir-
cular tumescences. Clitellimi, saddle-shaped (all), extending down
to B (Wisconsin), nearly to mV (all New Mexico), reaching into xiii
and xix (Wisconsin), in xii-xix (1), xiii-xix (28),xiii-xx (1), lack-
ing ventrally in xix or xix-xx.

Genital markings, lacking (all New Mexico specimens). Each
marking of the Wisconsin worm has a circular area of translucence
surrounded by a broad rim of slight epidermal tumescence so

April 22. 1967 earthworm genus diplocardia 3

shaped that the marking has a circular to shortly and transversely
elliptical shape. Markings, usually paired but not median when
single, usually in the region of AB or centered there. Locations,
only approximate because of invisibility of intersegmental furrows
in some areas. 8/9, postsetal in ix, x. (in x on left side only and
centered at A), 10/11 just lateral to B, presetal in xviii, xix, post-
setal in xxi, xxii, xxiii, perhaps also in xxiv but there tumescence
slight and no translucence is recognizable.

Internal Anatomy. — Septa, 4/5 membranous, 5/6 muscular
and translucent, 6/7-8/9 (Wisconsin) thickly muscular, thickness
of 9/10-12/13 decreasing posteriorly. 6/7-12/13 (New Mexico)
slightly muscularized. The usual subpharyngeal and subesophageal
horizontal mesenteries are present. Pigment, none visible in sec-
tions through the body wall. A special mid-dorsal, longitudinal
muscle band is present from the region of 10/11 to the hind end.
Pharyngeal glands, confined to region in front of 4/5.

Gizzards, rather weak, in v-vi (11). Calciferous glands and
lamellae, lacking. Esophagus (Wisconsin) not especially wide in
any segment, wdth a ventral typhlosole and numerous, low, closely
crowded nonlamelliform vertical ridges in x-xiii. Intestinal origin,
doubtfully in xvi (Wisconsin and 2, New Mexico), in xviii (8,
New Mexico). Such variation seemed unusual. After determining
intestinal origin in the eight specimens, the other two New Mexico
worms were reexamined. An esophageal valve was unrecognizable.
The gut in xvi-xvii certainly was grossly distended by the ingesta.
GDndition does not permit determination of the nature of the tissues
in the region in question. Typhlosole, beginning in the region of
xx-xxi. relatively fairly high, simply lamelliform, ending abruptly
in the 87th segment (Wisconsin).

Dorsal blood vessel, single from the periproct to region of iii
where it passes under the brain and bifurcates (Wisconsin). The
branches shortly attenuate to invisibility. Ventral trunk, complete.
Supra -esophageal trunk, at least in part empty and extent not
determinable. Extra -esophageal trunks, median to hearts, close to
or on the gut in vii-xiii. Posterior lateroparietal trunks, no trace
recognized though probably present. Suljneural trunk, unrecog-
nizable throughout and probably lacking. Hearts, of vii-ix (perhaps
also of x) lateral, of xi-xii probably latero-esophageal. Last pair of
hearts, in xii (11). A large, looped vessel from the dorsal trunk
passes down to the ventral parietes on each side of the body and
just in front of each septum in the intestinal region. A short, trans-
parent thread sometimes seems to connect such a segmental vessel
to the ventral trunk.

Nephridia. present from ii. wdthout recognizable vesicles, small,
simple, investing peritoneal cells not distended, funnels about at A,
ducts passing into parietes at D gap in the musculature.

Seminal vesicles, medium-sized, acinous, in ix and xii; the
anterior pair sometimes with several fairly large lobes, the posterior

The Great Basin Naturalist
4 G. E. GATES Vol. XXVII. No. 1

pair sometimes with more but smaller lobes. Sperm ducts, without
epididymis, coming into contact in xiii, at least through xviii not
united, in xix just lateral to prostatic ducts, not traceable to the
male pores. Prostates, large, in some or all of xiii-xxiv, once reach-
ing into xxviii, to 15 mm. long and 0.5 mm. thick. Prostatic ducts,
only a little thicker than the slender penisetal follicles, vsdthout
muscular sheen, 2+ mm. long (Wisconsin), l-f- mm. long (New
Mexico). Diagonal muscles in region of the male field (New Mexi-
co) are few and weak. Penisetal follicles, shorter than the prostatic
ducts, do protrude slightly into coelomic cavities.

Spermathecae, small to medium sized, ampullae empty and
strongly contracted (Wisconsin), large and reaching to the dorsal
parietes (New Mexico). Ducts, slender and shorter than the am-
pullae. Diverticulum, approximately discoidal, vertically placed,
with a row of seminal chambers somewhat like the teeth of a
comb, opening into the duct just below the ampulla. As so
described, the diverticulunri looks much the same as in various other
species of Diplocardia. However, in present specimens of verrucosa,
as well as in various congeners, individual organs often deviate
more or less vsddely from the norm described above. Thus, the semi-
nal chambers may not be in a row but arranged in an arborescent
pattern. The organ may be more or less deeply bi-, tri-, or quadri-
lobed or otherwise. One lobe may be much larger than the others
and without iridescence. The bottom of each of several New Mexico
ampullae was occupied by a flat and markedly iridescent disc. That
material is continuous with the contents of diverticular seminal
chambers. Obviously the seminal chambers did not have space
enough to hold all of the seminal fluid.

Follicles of a and b setae of viii-x are just visible in the parietes
and probably do not contain specially enlarged and modified ( copu-
la tory) setae.

Ovaries, fan-shaped, each with several egg-strings, which are
short (New Mexico) or long (Wisconsin). Ovisacs, small, in xiv
(Wisconsin).

Reproduction. — Iridescence on male funnels is brilliant and
indicative of massive production and aggregation of sperm. Irides-
cence also is obvious in spermathecal diverticula, proving that copu-
lation had been completed. Accordingly, and m absence of any
definite evidence to the contrary, reproduction can be assumed to
be amphimictic.

Ingesta. — Very fine-grained, black (Wisconsin) or red (New
Mexico) earth in which few plant fragments were recognizable.
The species appears to be geophagous.

Parasites. — Coelomic cavities of posteriormost segments (Wis-
consin) were filled with spheroidal cysts. Nematodes and other
foreign organisms were not seen.

April 22, 1967 earthworm genus diplocardia 5

Systematics. — The two subspecies are distinguished from each
other by situation of ventral boundary of the clitellum, location of
spermathecal pores, shape of seminal grooves, and patterns of
genital-marking location. The Wisconsin worm has genital markings
as do both subspecies, is more like the Oklahoma subspecies, recta, as
regards clitellar boundaries and spermathecal pore locations. New
Mexico worms also have the recta location of spermathecal pores
but a verrucosa clitellar boundary and are distinguished from both
subspecies by absence of genital markings as well as by a more
posterior intestinal origin.

Remarks. — One fourth of the area of Wisconsin is a major part
of some 15,000 square miles that until recently was called the
Driftless Area. The name indicated a belief that the region had not
been glaciated during the Quaternary. A much better chance of
survival would seem to have been provided earthworms by such
an area than by the possibly bare, mountain-peak nunataks on
which living organisms sometimes are thought to have survived
through Ice Age millennia. Collections eventually procured for
the author from the Driftless Area contained only one specimen of
a native American species. Any importance that individual might
have had was dissipated by geologists who found that all of Wis-
consin had been covered by ice. Although the glaciation of the
region in question is described as light, earthworms are unlikely to
have survived therein.

D. verrucosa is. of course, endemic somewhere in America —
though iust where still is to be determined. No information is
available as to self -migration rate of any diplocardia. There is, how-
ever, no good reason for believing sufficient time has been available
for migration to Wisconsin from well below the southern limit of
the ice sheet. Presence in Wisconsin then is due to transportation
and probably by man since the first European settlement. Else-
where D. verrucosa has been recorded from Lake St. Marys. Ohio
(a single specimen), central Illinois (where Harman. 1960. found
it in 16 counties), Omaha (Nebraska), and two counties in Okla-
homa. The Ohio site, as also all Illinois sites, is within the glaciated
area. Introduction to Ohio and to Nebraska is anticipated. Unfor-
tunately, our knowledge of Illinois earthworms dates only from
1885, but even so. introduction to central Illinois does not seem
impossible or even improbable. Certainly there seems to be no
reason at present for suspecting introduction by man to the two
Oklahoma counties.

The other distributional discontinuity may at first seem sug-
gestive of human introduction to New Mexico. However, the
paucity of information about the earthworms of an area extending
from central Texas to the Pacific Ocean at present contraindicates
any such conclusion. Also, very much more needs to be learned
about distributions within the diplocardian refugium. Texas
straight east to the Atlantic Ocean.

The Great Basin Naturalist
6 G. E. GATES Vol. XXVII, No. 1

Diplocardia texensis Smith

1924. Diplocardia keyesi texensis Smith, Proceedings, United States National Mu-
seum, 66(12):2. (Type locality, Chillicothe, Texas. Types, 2, both sectioned,
in the U. S. Natl. Mus.)

1965. Diplocardia sp., Bhatti, Proceedings, Pennsylvania Academy of Science,
39:8.

Habitat. — Arizona (Gila County). Watershed F at 3 Bar
Watershed, at ca. 3,800 feet, upper six inches of soil underneath a
Turbinella oak bush, March 31. 1966, 5-0-0-2. T. W. Barrett per
D E. Beck. Sierra Ancha Exp. Forest, moist, dark, clay loam (soil
sample #107) from thicket of walnut and oak trees, about one
mile from waterfall at head of Workman Creek, at 7,450 feet,
November 22, 1966, 0-1-0. Floyd Stockton per T. W. Barrett.

Pennsylvania (Delaware County). Swarthmore, bank of Crum
Creek, November, 1962, 0-1-1. H. K. Bhatti.

Texas (Lubbock County). Lubbock, flower bed. August 16,
1955, 0-0-2. J. E. Sublette per W. J. Harman.

(Victoria County), 0-1-0, (U.S. Natl. Mus. No. 57489). Vic-
toria, 5 miles southeast of town, prairie region, October 7. 1941,
0-2-13. J. D. Mitchell. (U.S. Natl. Mus. No. 57463).

External Characteristics. — Size. 44-100 by 2.0-2.5 mm. Seg-
ments, 74, 83, 100 (posterior amputees?). 121 (Swarthmore), 122
( + ?), 129, 130, 131 (2 specimens), 132, 134, 135. 136. 142, 147,
156 (Lubbock). Color, white. Secondary annulation, usually pro-
nounced behind male field except in last 8-20 segments, a pre- and
postsetal secondary furrow per segment, the posterior secondary
annulus usually with a well-marked tertiary furrow and other
tertiary furrows may be present. Such annulation makes counting
of segments very difficult especially as setae often are unrecogniz-
able externally and dorsal pores are not distinguishable. Body, with
a nearly circular cross section posteriorly. Prostomium, epilobous,
tongue closed (Swarthmore and 5 Arizona), open (17). Setae,
present from ii where none are lacking, AB ^ or slightly < CD,
BC much < AA, DD ca. ^1/2^", posteriorly BC may become so
much smaller as to be only slightly larger than AB, a, b follicles of
viii, ix, xviii-xx confined to parietes and in them setae usually seem
to be lacking. Nephropores, when detectable, about at D in iii-vi
or vii (several), posteriorly at or well above D but without regular
alternation in level, and accordingly in a rather irregular single
rank on each side of the body. First dorsal pore, at 7/8? (1 speci-
men), ?8/9 (4). 8/9 (1), ??9/10 (1). ?9/10 (1). 9/10 (3), ??10/11
(2), ?10/11 (1), 10/11 (2), ???11/12 (1), ??11/12 (1).

Spermathecal pores, sometimes obviously larger than other
genital apertures, superficial, each within a very small protuber-
ance, at or close to A and slightly or immediately behind 7/8, 8/9.
Female pores, anteromedian and much closer to a than to each
other, within a distinctly delimited transverse area (occasionally
dumbbell shaped) that covers the anterior half of xiv in A A (14),

April 22. 1967 earthworm genus diplocardia 7

a female field unrecognizable in the other worms. Prostatic pores,
at ends of seminal grooves and hence at eq/xx and eq/xxii, in AB.
Male pores, each on a very small tubercle in a seminal groove and
at or just behind level of 20/21, perhaps about at eq/xxi of one
Texas worm. Seminal grooves, nearly straight, deep and wide. A
postequatorial and preequatorial furrow in xxii, between the A
meridians, of the Swarthmore specimen, at first appear to be
median continuations of 20/21 and 21/22, with the seminal grooves
seemingly confined to xxi. Intersegmental furrows are, however,
obliterated across the male field. The latter is not distinctly demar-
cated, nor is it depressed, but it is distinguished by a greater epider-
mal opacity that extends well into BC between 19/20 and 22/23.
Genital markings, lacking (all).

Clitellum. reddish brown reaching just below A (Swarthmore),
to just below C in xxi. to mBC in xx, in other segments to well below
A or nearly to mV, from eq/xiv to 16/17 (1),) or annular in xiv (1),
xiv-xv (1), dorsal pores occluded, setae retained, intersegmental
furrows obliterated (Swarthmore) or faintly indicated dorsally and
gradually becoming more obvious ventrally (some Texas), in xii-xx
(Swarthmore, 1 Arizona), xiv-xix (12). xiii/n-xix (1), xiii-xx (2).
xii-xxi (Lubbock).

Internal Anatomy. — Septa, 4/5 membranous. 5/6 slightly
thickened, 6/7-10/11 increasing a little thicker, 11/12 muscular and
opaque or translucent, 12/13 slightly muscular and translucent. In-
sertions on parietes laterally of 6/7-10/11 are posterior to levels of
corresponding intersegmental furrows. Special longitudinal muscle
band at mD. too indistinct to be helpful in determination of function
of pore-like markings. Gaps in longitudinal musculature show that
AB obviously < CD. Pigment, unrecognizable in sections of body
wall.

Gizzards, two, in v-vi (14). Calciferous glands and lamellae lack-
ing. Esophagus usually of nearly uniform width throughout, very
slightly vsddened in xii-xiii, xii-xiv. viii, or xiv and then thin-walled,
or even much widened in xviii (2). The inner wall may have at
mV a well-marked and blood-gorged ridge lateral to which there are
low and rounded, red or white, nonlamelliform, vertical or longi-
tudinal ridges, or ridges may be replaced by villiform protuberances.
Intestinal origin, in xix (12). A widening of the gut in xviii of two
worms is believed not to be intestinal but condition did not permit
certainty. Typhlosole. first recognizable in region of xxi-xxiii, low
but thickened lamelliform. height gradually decreasing posteriorly,
unrecognizable behind 50th (1). 80th (1). 63rd (147). or 74th (of
142), segments. The lamella in a posterior amputee of 83 segments
was rudimentarv behind the 44th and was unrecognizable behind the
63rd.

Dorsal trunk, single throughout (5), complete, bifurcating under
brain, the branches uniting (1) to become the ventral trunk which
also is complete. Supra-esophageal, single, adherent to gut reaching

The Great Basin Naturalist
8 G. E. GATES Vol. XXVII. No. 1

into ix and xii. Extra-esophageals, median to hearts, on gut from ix
or X through most of xiii. Posterior lateroparietal trunks recognized
only from xxi or xx, turning up to join the extra-esophageals on the
anterior face of 12/13. Subneural trunk, no trace recognized in any
dissected specimen, probably lacking. Hearts, in v not traceable all
the way to ventral trunk, in vi-ix lateral, in x-xii probably latero-
esophageal though no blood is present in filaments passing to dorsal
trunk, last pair in xii (14). Segmental commissures, behind clitellum
are covered by a layer of chloragogue that often is thicker than the
vessel itself.

Nephridia. small, simple, seemingly avesiculate, on parietes in
AD or from mBC to somewhat lateral to D, ducts passing into pari-
etes about at D (or? entrance often not recognizable because of con-
dition ) .

Seminal vesicles, acinous, nearly filling coelomic cavities
(Swarthmore) but larger in xii. Vesicles of ix were not found in
several Texas specimens and must have been rather small unless
disintegration was involved. Sperm ducts, slender, those of a side
uniting in xx shortly before disappearing into the parietes of xxi.
Prostates, 5-9 mm. long (Swarthmore and some Texas), ca. 2 mm.
long and laterally directed in own segments (Lubbock), or in a U-
shaped loop in own segment, or variously twisted or extended, in
one worm reaching back into xxviii. Ducts, slender, without muscu-
lar sheen, 1-2 mm. long, sinuous or in one or two shortly U-shaped
loops, passing into body wall at B gap. Diagonal muscles, although
a number are present in xx-xxii. appear to be weak, no depression
or elevation of male field was recognizable. Follicles of ventral setae
may protrude slightly into coelomic cavities of xx and xxii but often
seem to have no shafts. When present, setae are straight or nearly
so, unornamented or with several fine teeth scattered near the tip.

Spermathecae, small, medium-sized, or reaching to or slightly
above level of dorsal face of gut. Ducts, shorter than, as long as, or
somewhat longer than the ampullae, narrowest within the parietes
and gradually widening entally but without becoming markedly
thickened. Diverticulum, as usual attached by a mesentery to the
posterior face of the septum in front, along lateral face of duct
entally, rather sausage-shaped, sessile (no stalk), opening into duct
directly through a single aperture in a middle portion, with 2-5
seminal chambers indicated by the nearly discrete balls of sperm.
Copulatory setae were not recognized in ventral follicles of viii-ix.

Ovaries, fan-shaped, with several short or long (and then with
as many as 20 ova) egg strings. One egg string of a Texas specimen
passes well down into the oviduct of its own side. Oviducts, narrowed
behind 13/14. (Ovisacs, recorded by Smith, were not recognized.)

Reproduction. — Spermatozoal iridescence is brilliant on male
funnels of dissected clitellate specimens with ovarian ova in discrete
egg strings. Iridescence in spermathecae was confined to the diverti-

April 22. 1967 earthworm genus diplocardia 9

cult, and as usual in worms of many other species, was not recogniz-
able in the main axis. Iridescence was lacking in Texas individuals
with ovaries still not separated into discrete egg strings even though
clitella seemed to be well developed. Copulatory transfer of sperm
having been demonstrated by the spermathecal iridescence, reproduc-
ing yellowish or reddish fragments.

Distribution. — A self-acquired range from Pennsylvania to Ari-
zona would be very much larger than that of any other diplocardia.
Such is not to be expected of a sj)ecies that seems, in several ways,
to be highly specialized. Absence of records for texensis between the
Delaware and Mississippi Rivers is of no significance at present be-
cause of the paucity of information about earthworms of that area.

Several diplocardias probably have been carried around the
United States by man who may have been responsible for presence
of texensis in the vicinity of Philadelphia as well perhaps as also in
Arizona.

Systematics. — Relationships suggested by the male terminalia
are wdth Diplocardia keyesi Eisen, 1896, known only from descrip-
tions of a single sectioned worm from Baja. California. Differences
from worms herein referred to texeruis now seem too great to be
attributable to individual or even to intraspecific geographic varia-
tion. An intestinal origin in xv. in Diplocardia, certainly seems to be
primitive. Origin in xix, recorded for each of thirteen specimens of
texensis. now seems to represent greatest esophageal elongation in
the genus. However, much more information about variation in each
of the two taxa is needed.

Parasites. — Two small nematodes were found in coelomic cavi-
ties of a Texas worm.

Remarks. — Intestinal contents show the worms are geophagous.
Soil particles were mostly quartz grains along wdth bits of mica and
black mineral particles of similar size (Pennsylvania) or (Lubbock
worms) rounded quartz grains with occasional black bits and numer-
ous yellowish or reddish fragments.

Unusual thickness of the clitellum of one Texas worm is not
associated wdth presence of mature sperm but with spermathecae
and ovaries that seemingly are juvenile. The clitellar tumescence
may then sometimes be a preservation artifact rather than an indica-
tor of height of breeding activity.

I'sually seminal grooves or dij)locardias are recognizable before
the epidermis of clitellar segments begins to become tumescent and
also after that condition has completely disappeared. As types were
said to be clitellate, supposed absence of seminal grooves needs an
explanation, especially as there seems to be no reason to suspect
that parthenogenesis (which would allow their elimination) is in-
volved.

The Great Basin Naturalist
10 G. E. GATES Vol. XXVII. No. 1

References

Hague, F. S. 1923. Studies on Sparganophilus eiseni Smith. Transactions Ameri-
can Microscopical Society, 42.

H.ARM.AN, W. J. 1960. Studies on the taxonomy and musculature of the earth-
worms of central Illinois. Thesis, University of Illinois.

MuRCHiE, W. R. 1962. A new subspecies of diplocardian earthworm from Okla-
homa. Ohio Journal of Science. 62:185-190.

Ude, H. 1885. Ueber die Riickenporen der terricolen Oligochaeten, nebst Beit-
ragen zur Histologie der Leibesschlauches und zur Systematik der Lumbrici-
den. Zeitschrift fiir wissenschaftliche Zoologie, 43:84-143.

SPIDERS OF THE NEVADA TEST SITE*

Dorald M. Allred and D Elden Beck'

Introduction

Since 1959, the Department of Zoology and Entomology of Brig-
ham Young University, under contract with the U.S. Atomic Energy
Commission, has conducted ecological studies at the Nevada Test
Site. Primary objectives are to establish baseline data as a prerequi-
site for determining the effects of nuclear testing on native animals.
Allred, Beck, and Jorgensen (1963) discussed the biotic communities
of the test site, and summarized the initial collection data on some
predominant species, including the spiders identified to that date.

No effort was made to specifically collect spiders in all of their
habitats. However, those reported here were taken in connection with
other studies, and reflect observations made on a daily basis for the
collection methods applied. Principal methods of collecting were can
pit-traps and berlese funnels {ibid.: 8-9; Allred and Beck. 1964),
and a few were taken by hand. We feel the present listing is a good
representation of the spiders which may be considered principally
terrestrial in habit. Specialized collecting applied to all types of
habitats would likely reveal many additional species. For example,
the whole range of flower-inhabiting spiders was not investigated,
nor the collection of specimens by net-sweeping of plants. A number
of new species were taken, but these will be described in a subsequent
paper by Dr. Willis J. Gertsch.

We are grateful to Dr. Willis J. Gertsch, American Museum of
Natural History, New York, N.Y.. for the identifications of the speci-
mens and for checking the names used in this manuscript. Collection
and identification of specimens, and analysis of data (in part) were
accomplished under AEC research grants AT(ll-l)-786. AT(ll-l)-
1326. AT(11-1)-1336. and AT(11-1)-1355.

In the following discussion a phylogenetic presentation has not
been followed. Families and the species assigned to them are listed
alphabetically for convenience. The total numbers of spiders col-
lected are indicated, and the months in which they were taken are
listed. Their relative abundance in each plant community is based
on the number collected in relationship to the number of collecting
attempts which varied between different communities (Table 1).
Where sex is indicated, the specimens are adults. Sex determination
was not made for the immature specimens in most cases.

Agelenidae
Agelenopsis aperta Gertsch. Fhree females were taken in July,

•B.Y.U. AEC. Publication No. C00-13J»)-5.

1 . Department of Zoologj- and Entomology, Rriglism Younp University. Prove, Utah.

11

The Great Basin Naturfdist
12 ALLRED AND BECK Vol. XXVII, No. 1

October, and November, and four immatures from May through
July. Two spiders were found adjacent to the pond at Cane Springs.

Calilena restricta Chamberhn and Ivie. Thirty-five males were
taken from August through December and in February, mostly in
October. Eighteen females were taken in April, August, and from
October through February, mostly in November. Fifty-three im-
matures were taken from January through November except in
February, and mostly during June, July, and August.

Circurina utahana Chamberlin. Four males were taken in No-
vember and December, two females in December and January, and
one immature in August.

Argiopidae

Apollophanes texana Banks. One male was taken in May. and
four females in June and July. One spider was taken adjacent to the
pond at Cane Springs.

Metepeira gosga Chamberlin and Ivie. One male was taken in
August. 15 females in June and August (mainly August), and 17
immatures in June, July and August. Five specimens were taken
adjacent to the pond at Cane Springs.

Tetragnatha laboriosa Hentz. One immature was taken in March.

Caponiidae

Orthonops getrschi Chamberlin. Fifteen males were collected
from April through December, except in August. Twenty-seven
females were taken from March through September, mostly in May.
Six immatures were collected in July, August, and October.

Tarsonops sp. One male was taken in September, and six females
in July, August, and October.

Clubionidae

Anyphaena sp. One immature was taken in October.

Castianeira sp. One immature was collected in October.

Corinna bicalcarata Simon. Three males were taken in Septem-
ber and October, and three females in May, July, and August.

Micaria gosiuta Gertsch. Eleven males were taken in August and
September (mostly in September), and one female in September.

N eoanagraphis chamberlini Gertsch. Seventy-six males were
taken in August, September, and October (mostly in September);
12 females from February through October, except in May; and 66
immatures from April through October, mostly during June, July,
and August. One spider was taken adjacent to the Cane Springs pond.

Neoanagraphis pearcei Gertsch. Forty-seven males were taken
in September and October (mostly in October), one female in July,
and nine immatures in June, August, September, and October.

April 22. 1967 spiders of Nevada test site 13

Phrurotirnpus sp. One male was taken in August.

Piahunn nanna Chamberlin and Ivie. One female was taken in
September.

Syspira electica Chamberlin. A total of 113 males was taken
from April through July and in October, mostly during May and
June. Eighty-two females were collected from April through Septem-
ber, mostly in June and July. A total of 390 immatures was taken
from April through October and in December, mostly during July
and August. Two spiders were taken adjacent to the Cane Springs
pond.

DiCTYNIDAE

Dictyna calcarata Banks. One male and seven females were taken
in July, and seven immatures in July and August. One specimen
was taken adjacent to the ])ond at Cane Springs.

Dictyna personata Gertsch and Mulaik. One male was collected
in November, and two females in May and June.

Dictyna reticulata Gertsch and Ivie. Six males. 12 females, and
seven immatures were taken in June and July, and one immature in
October. Twenty-four of these were taken adjacent to the pond at
Cane Springs.

Dictyna tucsona Chamberlin. Two immatures were taken in
June and July.

Mallos mians Chamberlin. One male was collected in October.

Mallos paUidus Banks. Four males, three females, and eight im-
matures were taken in July, all adjacent to the pond at Cane Springs.

Dig

UETIDAE

Diguetia canities McCook. Three females were taken in May.
July, and August, and one immature in June. One spider was taken
adjacent to the pond at Cane Springs.

Diguetia signata Gertsch. One male was collected in June, and
one female in May.

FiLISTATIDAE

Filistatt utahana Chamberlin and Ivie. Six males were taken
in June and July, one female in September, and two immatures in
June.

Gnaphosidae

Cesonia classica Chamberlin. Nineteen males were taken in
June. July, and August (mostly July). 13 females from June through
September, and 14 immatures in A lay, June. July, and October. One
spider was taken adjacent to the Cane Springs pond.

Drassodes celes Chamberlin. One male was collected in April.

The Great Basin Naturalist
14 ALLRED AND BECK Vol. XXVII, No. 1

Drassyllus irritans Chamberlin. Thirty-one males were taken
from February through June and in November (mostly in April),
47 females from April through August (mostly in June and July),
and 47 immatures from February through December (except in
April and June), mostly in August.

Drassyllus moronius Chamberlin. Four males were collected in
May and November, four females in May, and four immatures in
March, May, September, and October.

Gnaphosa californica Banks. One male was collected in November,
nine females in June and November, and 1 1 immatures in July,
August, and November.

Gnaphosa hirsutipes Banks. Twenty-four males were taken from
March through July and in November, 1 7 females in February, May,
and from July through November, and 34 immatures every month
except April.

Hapuodrassus eunis Chamberlin. A total of 149 males was taken
from October through May. mostly from February through April.
Eighteen females were collected from January through June. A total
of 204 immatures was collected, and every month was represented
except May. Greatest numbers were found from October through
January.

Herpyllus hesperolus Chamberlin. Sixteen males were taken
during March, April. May, and December (mostly in April). Ten
females were collected in April, June, and July. Forty-five immatures
were taken representing every month except A{)ril. One specimen
was taken adjacent to the Cane Springs pond.

Megamyrmecion naturalisticum Chamberlin. Nine males were
collected from April through August, six females in June. July,
August, and October, and six immatures from August through No-
vember.

Nodocion utus Chamberlin. A female was taken in July, and two
immatures in July and September.

Zelotes monachus Chamberlin. Thirty-eight males were collected
from April through October (except July), mostly during April,
May. and June. Thirteen females were taken from March through
July and in October. Twenty-six immatures were taken from March
through October.

Zelotes nannodes Chamberlin. Two males and four females were
collected in June.

Zelotes puritanus Chamberlin. One female was taken in May
and an immature in July.

Heteropodidae

Olios fasciculatus Simon. One female was collected in September,
and an immature in July.

April 22. 1967 spiders of Nevada test site 15

HOMALONYCHIDAE

Honialonychus theologus Chamberlin. Four males were collected
in October, one female in April, and four immatures in April, June,
and October.

LiNYPHIIDAE

Ceratinopsis sp. One male was collected in January.

Erigone dentosa Cambridge. Three inales were collected in June
and October, four females in January, March, and September, and
three inmiatures in October. Nine of the specimens were taken
adjacent to the Cane Springs pond.

Meioneta fillmorana Chamberlin. One male and one immature
were taken in November.

Meioneta formica Emerton. Two females were taken in January
and April.

Spirembolus s\). One female was taken in February.

Tapinocyha sp. One male was taken in November.

Lycosidae

Alopecosa kochi Keyserling. Two males were taken in March
and April, and one female in May.

Geolycosa rafaelana Chamberlin. Three males were collected in
June and July, and one immature in August.

Pardosa ramulosa McCook. Six males, two females, and five im-
matures were taken in March, all adjacent to the Cane Springs pond,
and one male was collected in November.

Schizocosa sp. One immature was taken in June.

Tarentula kochi Keyserling. Eighty-seven males were taken from
October through May. mostly from January through March. Forty -
five females were taken e-, ery month except August, mostly from
November through March. A total of 111 immatures was taken,
representing all months except February.

Mimetidae

Mimetus euiypus Chamberlin and Ivie. One immature was taken
in October adjacent to the Cane Springs pond.

OXYOPIDAE

Oxyopes tridens Brady. Fifty-two males were taken from May
through August, 1 1 females from June through September, and 20
immatures from June through November. Greatest numbers of all
stages were taken during July.

Pholcidae

Physocyclus tanneri Chamberlin. Three males were collected in
August and November, and six females and five immatures in June,

The Great Basin Naturalist
16 ALLRED AND BECK Vol. XXVII, No. 1

July, and November. Seven spiders were taken adjacent to the Cane
Springs pond.

Psilochorus papago Gertsch. Twelve males were taken from June
through December, except in September. Seven females were taken
in June, July, and October, and two immatures in June. One spider
was taken adjacent to the pond at Cane Springs.

Psilochorus utahensis Chamberlin. A total of 450 males was
taken representing every month of the year except February. They
were most common during June, July, and August. A total of 499
females was collected over all the months of the year except March.
They were most common also during June, July, and August. The
total immatures collected was 1500. and they were found during
every month of the year, mostly during the same months as the
adults, as well as in September. Two specimens were taken adjacent
to the pond at Cane Springs.

Plectreuridae

Kibramoa paiuta Gertsch. One male was collected in May.
Plectreurys tristis Simon. One male was taken in May, and seven
immatures in January, April, June, July, and September.

Salticidae

Metacyrba arizonensis Barnes. Thirteen males were collected
during May and June, seven females in May, July, and November,
and 16 immatures from April through October. Two specimens were
taken adjacent to the Cane Springs pond.

Metacyrba taeniola Hentz. One male was taken in June, and
two immatures in September.

Pellenes brunneus Peckham. One male was collected in October.

Pellenes hirsutus Peckham and Peckham. One female was taken
during July.

Pellenes limatus Peckham. One male was collected in June.

Pellenes oregonensis Peckham and Peckham. Three males were
taken during July and August, and one female in July.

Phidippus apacheanus Chamberlin and Gertsch. One male was
collected in September.

Phidippus formosus Peckham and Peckham. Three males were
taken in March and June, one female in February, and seven im-
matures in July, August, and September. Two spiders were taken
adjacent to the pond at Cane Springs.

Phidippus opifex McCook. One male was found in August.

Phidippus workmanni Peckham and Peckham. Twenty-seven im-
matures were taken during June. July, and August. Two were taken
near the pond at Cane Springs.

April 22. 1967 spiders of Nevada test site 17

SCYTODIDAE

Loxosceles unicolor Keyserling. Twenty-one males were taken
from April through August and in October. Twelve females were
collected in July. August, September, and November. Twenty-
seven immatures were taken from May through October. Four speci-
mens were taken adjacent to the pond at (]ane Springs.

Theraphosidae

Aphonopelma sp. Twenty-four males were taken from August
through November, and two females in June.

Theridiidae

Enoplognatha joshua Chamberlin and Ivie. Twenty-nine males
were collected in February and March; 21 females from February
through May; and 15 immatures in January. February, March, and
October. One specimen was taken near the pond at Cane Springs.

Euryopis scriptipes Banks. One female was found in June.

Euryopis spinigera Cambridge. Two females were taken in April
and June.

Latrodectus niactans Fabricius. Four males were taken in April,
May. July, and December; 10 females in April, June. July, and
December; and 28 immatures from July through January.

Steatoda juhm Keyserling. Twelve males were found from May
through September; five females in February. August, and Septem-
ber; and nine immatures from May through September.

Steatoda medialis (Banks). One female was taken in July, and
one immature in June.

Steatoda washona Gertsch. Two females were found during
August.

Theridion sjx One immature was taken in July.

Thomisidae

Ebo dispar Schick. Three males were taken in October and De-
cember, and five females during October. November, and December.

Ebo merkeli Schick. One female and two immatures were taken
in July.

Ebo mexicanus Banks. One female was taken in May.

Misumenops desertus Schick. One male was taken in August,
and six immatures in August and October. One spider was taken near
the Cane Springs pond.

Misumenops rot hi Schick. Eight males were collected in June
and July. 13 females in May, June, and July, and 16 immatures in
June. Jul}', and October. Ten spiders were taken near the Cane
Springs pond.

The Great Basin Naturalist
18 ALLRED AND BECK Vol. XXVII. No. 1

Philodromus infuscatus Keyserling. One immature was found in
July.

Rhysodromus clarus Keyserling. Fifteen immatures were taken
in July.

Thanatus texanus Banks. Fourteen males were found from April
through August, eight females in May and from July through No-
vember, and eight immatures from June through November, except
during September.

Xysticus californicus Keyserling. One male was taken in May.

Xysticus iviei Schick. One female was found in June.

Xysticus lassanus Chamberlin. Eight males were collected in
January, March, April, May, and September. Nine females were
taken in February. April. May. June, September, and December.
Twenty-seven immatures were taken from May through November.

Uloboridae
Uloborus diversus Marx. Three females were taken in August.

Summary

During the years 1959-1965, more than 5600 spiders were col-
lected in connection with other studies at the Nevada Test Site.
These represent 94 species of 65 genera in 22 families, not counting
approximately 17 new species not reported here. The greatest num-
bers of species were found during June and July (Fig. 1 ) , and popula-
tions of spiders were highest from June through September (Fig. 2).
The Coleogyne and Mixed communities supported the greatest num-
bers of species, and the Coleogyne and Salsola conmiunities possessed
the highest populations (Table 2). Fewest species were found in the

40

y^

32

,/■■■ '"v^

.^\

24

^

\

y^

"7 '~"""\,.^ ^

16

^ /

^^4^

^__\

'

_/,

y^J^^

<<1

/

^ -^. \ J>.

8

Mq 1 G S

Fema les

-w-^

1 1

Immatures —

1 1 1

1 1 1 1

Jan Feb Mar Apr May June July Aug Sept Oct Mov Dec

Fig. 1. Seasonal representation showing number of species present at the
Nevada Test Site, 1959-1965.

April 22. 1967

SPIDERS OF NEVADA TEST SITE

19

o8

U

ill

♦ « « «

vn ♦

« •

« • •

CO

CS) ♦

♦ ^O ♦ CSI ro «

t^ « (N -^

♦ Tf- • ro rn

CQ

^ CM T}- CO

in » "^ •

« t^ •^ CO

» fO PO

♦ *-<^(Ovri »t^ •

• ro -^ CSI C<J

« csj ■^ u^

T- G

•a ^ S

41 ? ? "^

.o
a o

^;.

<

-a o
■a

o ^

Sd S

ft -^

k. ^

§«l

o

^ ?

re

-o

6c '^

C5. Ci.

o o

O o

2 c.
fc£ kl

3

~ ~ !> •»-

<3

^

« o

■^ -ii

« .'^

?. ;^ ^ S: q: c^' = Q Q

u

20

ALLRED AND BECK

The Great Basin Naturalist

Vol. XXVII, No. 1

o

♦ ♦

'sD*»c<ivnc<i ♦<©

csi ♦ tJ- csi

CSl

* CO ♦xn»*<-DTj-oo fO fO

04 t}- tn CO vn 04 »

CO ^o o

T-l « » ■<— I ■,— I •,— (

CO ♦ CO vn tj-

QCl

^ tie S «
'^ -is

^ g 3 « C

■° c: ° 5-ii: ^

^ ^ f* C<; "** k.

Q..O

!/5 /T»

3

<o 3

!5

3

c/2 to

^

ta

CO

o

^NNN

April 22. 1967

SPIDERS OF NEVADA TEST SITE

21

U

Ah
u O

o

u

0)
C/2

c

* « vo

OQ

04

CO

3

<u

CO

"c :h

3

"a
<>^

to 2 (-
>^-2 5 6
o

3

^ ci.

. o

1-1 1

So

•- ^ c ;~^ t,

2* ^" ~ ^ "^ ^ C

■ ~ -3?' "^ ~ o c c

(/5

- o

52 <3

^ ^ r^ rR ^ ^ ^ At!'
— O <0 kj < < r§ 00 t^

c

2 o ? >^

o

h-1

3

a,

3

«ij

22

The Great Basin Naturalist
ALLRED AND BECK Vol. XXVII, No. 1

♦ » «

» • ♦

■^ xn

^ Tj-

Cs)

CS) CSl CSJ lO c^

^ CT) ^ rj-

^ ^ 'JZ>

« CSI CS)

c^j» ♦ *-^(Mt-hcn

PO ro

o

3 i5'5

w '^- o

•^ G, a, c^

"o

0.

_2-r

■ 2 *>.

X

1'

<u o a

-sil

•E g 12

0)

3<i ^

CO

0) •— ~-

""O

(V

CO

K

C/D

3 3

^2

2

i 6 H

*J >; rs\

C/j

^ ~u;

^ T3

0) ^
CO ^

£^Ci,=i,

o

o

ry: r^ r-

April 22, 1967

SPIDERS OF NEVADA TEST SITE

23

6C
4)

o

Oh

«8 ?

ih o

°;3

• (N ♦

rj- •

♦ CI

♦ ♦ -^ CSI PO

♦ C^ CSl OO

-^ -"J-

rn

•^ ^

(N

rO -^ y-i

PO

<2

3

5

o "^ S
k; kj '-J c/D

o ^

§ ^

•J '^ -a

c ^ c

g 3 3

C5.
O

s; ^ -h ^
^ ^- ^ -s: rc

c
3'S §

gkjt*iki<<ci,ft:K><><>< o^

o

3

o
x>
o

-31
II I

E -^ = «
"-' :- s c

2 C Eb

on f ICO

^ ■»; L- 4)

■^ c c

E S'^ *

3 '^ 2(j

= P " *-
c I a "

II 9 = S

o. N, 5 a

I i^l

I ^xjll
'-' t. .= ti

° ^ i>

to

O; O -y -^

1 11°

I P3^

a P c

s "all

C a ^ ;u

§ |<8
1 |ll g

c

1-^1.

3

•-

• S-S S

i: 2 ^

c

•^it •-,

~

ll'g-S

"x

(oa.

c

S"

■■tin

c
O

^£

24

ALLRED AND BECK

The Great Basin Naturalist

Vol. XXVII, No. 1

t^i 5;

Oh

O

X

X

CN

X

Csl

i.E c

.5^^-

3 ^

April 22, 1967 spiders of Nevada test site

25

600

/
/

\

500

/
/
/

\
\
\
\

400

M a

es

/

Fe ma

es

_ _ _

_

_

/

\

Immatures

_

/

\
\

300

/

/

\

\
\

200

^

/

\

\ \

100

_-- - ~

X
1

1

'

1

1 1

1 I 1 J

.^^

Jan Feb Mar Apr" May June July Aug Sept Oct Nov Dec

Fig. 2. Seasonal abundance of spiders (all species) at the Nevada Test Site,
based on total numbers of individuals collected, 1959-1965.

Pinyon-Juniper community, and lowest populations in the Grayia-
Lycium (Table 2).

Eight species were found in each of the seven plant communities,
and were widely distributed at the test site. These are Calilena re-
stricta, Drassyllus irrltans, Gnaphosa hirsutipes, Haplodrassus eunis,
Neoanagraphis chamberlini, Psilochorus utahensis, Syspira eclectica,
and Tarentula kochi. Psilochorus utahensis was by far the most
abundant in numbers of individuals, followed by S. eclectica, H.
eunis, and T. kochi. Haplocrassus eunis, P. utahensis, and T. kochi
were the most abundant in numbers of individuals during all 12
months of the year.

References

Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. Biotic communities

of the Nevada Test Site. Brigham Young University Sci. Bull., Biol. Ser.,

2(2): 1-52.
Allred, D. M., and D E. Beck. 1964. Arthropod associates of plants at the

Nevada Test Site, Brigham Young Univ. Sci. Bull., Biol. Sen, 5 (2): 1-1 6.
CoMSTocK, J. H. (revised AND EDITED BY W. J. Gertsch). 1948. The spider

book. Gjmstock Publ. Assoc.. Ithaca.

PROBLEMS OF SAMPLING DESERT ARTHROPODS BEFORE
AND AFTER A THERMONUOLEAR CRATERING TEST

Clayton S. Gist^

Introduction

On July 6, 1962, the U. S. Atomic Energy Commission detonated
a large thermonuclear device at the Nevada Test Site (Project
Sedan). The device was buried 635 feet underground and had a total
yield of 100=bl5 kilotons. The explosion ejected about 7.5 million
yards of alluvium and produced a crater 320 feet deep and 1200
feet in diameter (Fig. 1). The ejected alluvium v\as radioactive, and
gamma intensities of several hundred milliroentgens/hour persisted
for months within a mile of ground zero. After a year, gamma inten-
sities at the edge of the crater were still about 1 5 mr/hour.

Some of the effects of this test on plants and animals in areas
near ground zero have already been reported (Allred et al. 1964,
Jorgensen et al. 1963. Martin 1963, and Turner and Gist 1965). One
feature of the biological studies was an analysis of selected arthro-
pods near ground zero before and after the test. The study was re-
stricted to species which could be sampled readily with sunken can
traps, and which were known to be well represented in the test area.
I sought evidence of possible changes in species composition, or in
relative abundance of species as a result of the test. These factors
might influence the long-term recovery of the close-in area, or in-
directly affect the survival of vertebrates in these areas.

The area selected for the test was 4317 feet above sea level and
located at the north end of Yucca Flat. This desert basin is about
15 miles long and 10 miles wide, in the northern portion of the
Nevada Test Site. Floristically it is part of the broad transition be-
tween the Mojave Desert and Great Basin (Beatley 1962). The area
in the vicinity of ground zero had been used for testing in the past and
much of the native vegetation had been destroyed and replaced by
Russian thistle {Salsola kali) and stick-leaf (Mentzelia veatchiana).
However, to the northeast of ground zero — at least beyond 3000 feet
— the vegetation in 1962 was relatively undisturbed (Fig. 2) and
composed predominantly of shrubs, e.g., hopsage {Grayia spinosa)
and blackbrush (Coleogyne ramosissima) . Studies of arthropods
were centered in this area.

Methods and Results

Two study plots were established along a line running from
ground zero to 10.000 feet on a grid azimuth of 58°. Between 3530
and 3800 feet, a 10 x 10 grid of 100 buried can traps was established

1. Laboratory of Nuclear Medicine and Radiation Biology, UCLA School of Medicine, Los
Angeles, California 90024.

26

April 22. 1967 sampling desert arthropods

27

>

<v ^

CO -,

^ O

The Great Basin Naturalist

28 c. s. GIST Vol. XXVII, No. 1

on June 21 (Fig. 3). The traps were 30 feet apart and the grid was
centered on the surveyed line. A similar grid was set up between 8730
and 9000 feet on June 23. The area of such grids was about 72,900
square feet (1.7 acres) in extent. The cans were 6.9 inches deep and
6.3 inches in diameter. All traps were covered with a square foot of
masonite wdth one-inch legs at each corner. Except for June 27,
when a small device was exploded on Yucca Flat, the traps in both
grids were examined daily between June 22 and July 5. Live arthro-
pods were removed from the traps, recorded, and released beneath the
nearest bush so as to prevent unnatural losses due to predation. Dead
animals were recorded and discarded. Traps were sealed on the
afternoon of July 5. and remained closed until radiation intensities
in the study area had declined to a safe level. Sampling was resumed
on July 30. 24 days after the test, and continued for two weeks
during August.

In June 1963 a new grid of 100 can traps was established be-
tween 4730 and 5000 feet from ground zero on the line described
above. This new grid of traps, along wdth the two grids at 3800 and
9000 feet, was sampled daily between June 10 and June 28.

The arthropods recorded were common species (or species
groups) which could be distinguished in the field. If errors in iden-
tification occurred they were probably in attempting to discriminate
between species of young scorpions and between Eleodes hispilabris
and other beetles of this genus.

The results of the sampling are given in Table 1.

Discussion

The interpretation of the numbers in Table 1 is uncertain for the
following reasons. Small differences in the numbers of individuals
captured do not necessarily imply real differences in abundance.
Thus, when small numbers are involved, even relatively "large"
differences are probably insignificant. Without repeated trapping
in the same area, preferably by means of replicated grids in identical
habitats, there is no way to assess the significance of such data except
intuitively. The problem is first to identify any real differences
suggested by the data, and then to determine whether these are
related to the Sedan test.

The tenebrionid beetle. Pelecyphorus pantex, can be eliminated
from further consideration. Its absence from traps during June, and
its presence during August, is a seasonal activity effect unrelated
to the Sedan test (see Allred et al. 1963). Secondly, the area at 9000
feet was not comparable to that at 3800 feet. Not only did the vegeta-
tion differ (see Jorgensen et al. 1963). but also the substratvun at
9000 feet was desert pavement, while at 3800 feet it was sandy. The
niunber of arthropods captured at 9000 feet was invariably less than
that taken at 3800 feet.

The data based on samples taken at 9000 feet do not suggest any
changes associated with the test. In fact, probably the only significant

April 22. 1967 sampling desert arthropods

29

^

'^v?

Li^' ^

1

.^^,

•.V.

.^jf^.

1 *

;-*■''"

" -.f.

•1

1

tlH

3

0)

>^

'o

be

C

c

o

t

T3

o

a

s

g

4;

1

o

Z

O

o

vn

oi

tfl

be

I.

E

The Great Basin Naturalist

30 c. s. GIST Vol. XXVII. No. 1

Table 1. Captures of arthropods in grids of 100 traps before and after the
Sedan test of July 6, 1962.

3800 feet 5000 feet 9000 feet

^

a-,

■^

CO

Species

0)

3
1—1

3

3
<

0*

d

3

3

1-5

C
3

>-5

1

3
<

4)

c

3
>->

Scorpions

Vejovis confusus Stahnke

28

106

40

60

19

22

10

Hadrurus hirsutus Wood

11

43

2

14

5

14

14

Orthopterans

Arenivaga

apacha (Saussure)

2

16

0

8

2

4

1

Stenopelmatus

fuse us Thomas

76

19

97

225

3

0

0

Ceuthophilus spp.

16

6

2

4

5

6

0

Tenebrionid beetles

Trogloderus

costatus LeGonte

20

91

122

144

6

1

5

Eleodes hispilabris (Say)

11

31

6

5

8

8

3

Pelecyphorus pantex Casey 0

532

0

0

0

6

0

Eleodes spp. (mostly

E. armata LeConte)

40

20

0

2

18

9

0

'Number of days traps were examined.

difference between June of 1962 and 1963 was the failure to capture
beetles of the genus Eleodes (except E. hispilabris) in 1963. The
numbers of these beetles apparently declined at 3800 feet also but
I do not believe the Sedan test was responsible. Other investigations
have indicated that, except for the deposition of radioactive dust,
there were no demonstrable effects of the Sedan test at 9000 feet
(Martin 1963, Jorgensen et al. 1963. Turner and Gist 1965).

The remaining comparisons involve samples taken at 3800 feet at
different times, and samples taken at 3800 and 5000 feet in June
1963. The 1962 samples at 3800 feet showed considerable differences
in the numbers of arthropods captured. I believe it reasonable to
assert that changes in numbers of certain species, or changes in
susceptibility to capture, occurred between June and August. The
increase in scorpions was due to the appearance of young animals
in the traps, and is what would have been expected on the basis of
earlier work by AUred et al. ( 1 963 ) . Arenivaga apacha, Trogloderus
costatus, and E. hispilabris were captured in what were probably
significantly larger numbers, but the causes for this increase are
not known. On the other hand, the number of Stenopelmatus fuscus

April 22. 1967 sampling desert arthropods

31

0)

I

<c

r

. C

ky

[*,'JL.A

^

M

The Great Basin Naturalist

32 c. s. GIST Vol. XXVIL No. 1

captured decreased sharply, and there may have been a significant
drop in the number of camel crickets {Ceuthophilus spp.) captured.
However, I discern no patterns in these events implicating the
Sedan test.

In June of 1963 there were also indications of change at 3800
feet. The sampling time was about 30% longer in 1963 (18 days vs.
14 days), so some increase might be expected for this reason alone
{e.g., Stenopelmatus, Vejovis). However, the much greater abund-
ance of Trogloderus cannot be explained on this basis. Furthermore,
the samples contained fewer Hadrurus., Ceuthophilus and Eleodes
hispilabris, and no other Eleodes. The apparent decline in numbers
of Eleodes (other than E. hispilabris) has already been discussed. It
is surprising that some arthropods {e.g., Stenopelmatus, Trogloderus,
and Vejovis) persisted so successfully in an area subjected to such
devastation (Fig. 4). One might conjecture that herbivorous species
would be extremely sensitive to the sort of disruption experienced
at 3800 feet. On the other hand, carnivorous and omnivorous species
would, at least for a time, have some available food. The apparent
decline of the large carnivorous Hadrurus hirsutus may in some way
be related to the extensive destruction of surface cover. Both Steno-
pelmatus fuscus and Vejovis confusus are carnivorous. Trogloderus
costatus is probably omnivorous. The plant feeders {Arenivaga
apacha and Ceuthophilus spp.) at least showed no signs of increasing
at 3800 feet.

During June of 1963, more arthropods were captured at 5000
feet than at 3800 feet. Prior to the Sedan test these two areas were
similar in terms of soils and vegetation but I have no pre-test arthro-
pod data from 5000 feet. However, I believe that the greater numbers
of arthropods taken at 5000 feet during June 1963 reflect better sur-
vival in this area. This is not surprising in view of the relative
destruction at 3800 feet (Fig. 4) and at 5000 feet (Fig. 5). In short,
except for the reduction in numbers of arthropods at 3800 feet com-
pared to 5000 feet, I perceive no short-term test-related effects in
the data in Table 1.

There is no clear-cut relationship between apparent changes in
numbers and food habits, although there is a possibility that carni-
vorous or omnivorous forms persist more successfully than herbivor-
ous ones in the areas where the above-ground vegetation is largely
destroyed. If there were close-in effects, it is possible that they could
have occurred because of destruction or modification of habitat, and
not because of ionizing radiation.

Probably the most significant lesson which emerges from the
above experience is that deducing real changes in arthropod popula-
tions on the basis of captures in traps is difficult, and requires a con-
siderable sampling effort in order to avoid ascribing significance to
differences which have occurred simply due to chance. There is also
the difficulty of seasonal variation touched on indirectly in this
paper, and treated in detail for the Nevada Test Site by Allred et al.

April 22. 1967 sampling desert arthropods

33

Ki %

1 ;'ii^'>;*. ' 1

'

^?^

-^

• . > ■

M

■ !|

< ^1^^

>

'

• . ' W

J

•i

\ '

j|

■

•i

4

1 * ' . ■

>

I

A.

■ « . •

■H

,

- , 1

c

o

^

r«0

c/1

CO

hr

^ta

o

J3

c

?

f^

1

C8

e

o

CO

U

ta

o

L.

CO

(1)

34

C. S. GIST

The Great Basin Naturalist

Vol. XXVII, No. 1

'A

1? • ; \ .^---^

•'^^ -.1* - ' iv-! :. ■';•• ^' A

J?^ '^.^

^ CM

ffl be

Oj • —

o

If) ti£

April 22, 1967 sampling desert arthropods 35

(1963). Examining the records of several consecutive years does not
always guarantee the ability to evaluate the observations in some
future year, because no two years are precisely equivalent. Finally,
even if statistics and/or intuition suggest that the sampling reflects
true changes in numbers, the causes for such changes are apt to be
extremely elusive. With so many factors governing the numbers of
arthropods captured, it is almost impossible to pinpoint effects due to
some specific cause, unless there are across-the-board changes ob-
viously attributable to this particular influence.

Acknowledgments. Original reference material was generously
supplied by Dorald Allred of Brigham Young University, and one
subsequent identification was made by Vasco M. Tanner of the
same institution. I thank Frederick iurner of the University of Cali-
fornia, Los Angeles, and Vernon Stern, of the University of Cali-
fornia, Riverside, for useful criticisms and suggestions. I also thank
John Williamson, Civil Effects Test Organization coordinator, for
the many services without which this work could not have been
accomplished. These studies were supported by Contract AT(04-1)
GEN- 12 between the Atomic Energy Commission and the University
of California at Los Angeles.

Literature Cited

Allred. D. M., D Elden Beck, and C. D. Jorgensen. 1963. Biotic communi-
ties of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol Series,
Vol. 2, 52 pp.

. 1964. Close-in effects of an underground nuclear detonation on small

mammals and selected invertebrates. U. S. Atomic Energy Commission Report
PNE-226F, 22 pp.

Be.\tley, J. C. 1962. Vascular plants of the U. S. Atomic Energ>^ Commission's
Nevada Test Site, Nye County, Nevada, U. S. Atomic Energy Commission
Report UCLA 508, 33 pp.

Jorgensen, C. D., D. M. Allred, and D Elden Beck. 1963. Some effects of
an underground nuclear detonation on biotic communities at the Nevada
Test Site. Proc. Utah Acad. Sci.. Arts, and Letters. 40:49-61.

Martin, W. E. 1963. Close-in effects of an underground nuclear detonation on
vegetation. I. Immediate effects of cratering, throw-out, and blast. U. S.
Atomic Energy Commission Report PNE-228P. 45 pp.

Turner, F. B,, and C. S. Gist. 1965. Influence of a thermonuclear cratering
test on close-in populations of lizards. Ecology 46:845-852.

NEW SPECIES OF BARK BEETLES
(COLEOPTERA: SCOLYTIDAE), MOSTLY MEXICAN

PART VIP

Stephen L. Wood-
In order to make names available for other work. 21 species of
Scolytidae are described on the following pages as new to science.
Most of the species are from Mexico, although four species occur
in Honduras; three of these four are common to both areas. The
genera represented include: Hylastes (1), Pseudothysanoes (1),
Cactopinus (1). Pityophthorus (3). Pseudopityophthorus (4),
Gnathotrichus (4), Monarthrum (3). Microcorthylus (1). Amphi-
cranus (2), and Corthylus (1).

Most of the specimens were collected either by Dr. J. B. Thomas.
Canada Department of Forestry, or by myself. The type material is
mostly either in the Canadian National Collection or my collection,
except for a few paratypes in the California Academy of Sciences.

Hylastes mexicanus, n. sp.

This species is more closely allied to nitidus Blackman than to
other known species, but it has the sides of the pronotum much more
strongly arcuate and the pronotal surface much more finely punc-
tured.

Female. — Length 4.5 mm. (paratypes 4.3-4.5 mm.). 2.8 times
as long as wide; color black.

Frons convex, with a weak, transverse impression between eyes
and more strongly impressed just above epistoma; median carina in-
conspicuous, more strongly elevated below, continuing dorsad as a
fine line to interoccular impression; surface smooth and shining
above, rather dull below, very finely, deeply, closely punctured;
vestiture minute, inconspicuous, covering entire surface.

Pronotum 1.1 times as long as wide; sides widest on basal third,
but almost parallel to a point just anterior to middle then broadly,
evenly rounded to anterior margin, appearing much wider than in
nitidus; surface smooth and shining, with punctures fine, deep, close,
separated by distances about equal to their own diameters; glabrous.

Elytra 1.9 times as long as wide. 1.9 times as long as pronotum;
sides straight and subparallel on basal half, then increasingly arcuate
to the rather narrowly rounded posterior margin; striae 1 moder-
ately, others weakly impressed, the punctures small, deep; inter-
striae as wide as striae, feeblv convex, subshining. the punctures
rather coarse, close, deep, confused, their diameters slightly greater

1 . In part, field work for the study in which these species were collected was sponsored by
grants from the National Science Foundation Nos. GB-532 and GB-,S(i78.

2. Department of Zoology and Entomology, Brigham Young University, Provo, Utah, Scolytoidea
Contribution No ^0.

36

April 22. 1967 new species of bark beetles 37

than half those of striae. Vestiture confined to declivity, stout, almost
scalelike except for median rows of equally short bristles on each
interspace.

Male. — Similar to female except frons narrower and fifth visible
sternum longitudinally impressed and pubescent.

Type Locality. — Forty miles west of Toluca, Mexico. Mexico.

HosT.^ — Pinus sp.

Type Material. — The female holotype. male allotype and three
para types were taken at the type locality on July 15, 1953, at an
elevation of 7.900 feet, from the base of a pine about 14 inches in
diameter, by S. L. Wood. In addition. 64 para types were taken 10
miles west of El Salto. Durango. Mexico, during July 1964. in flight,
by J. B. Thomas.

The holotype, allotype and some of the paratypes are in my col-
lection, other paratypes are in the Canadian National Collection.

Cactopinus mexicanus, n. sp.

Although not very closely related, this species is more nearly like
huhhardi Schwarz than other species. It may be readily separated
by the stouter body, by the tuberculate elytral interspaces, by the
much steeper, more deeply sulcate elytral declivity, and. in the
male, by the shorter, widely separated frontal horns.

Male. — Length 1.6 mm. (paratypes 1.3-1.6 mm.). 2.1 times as
long as wide; color black.

Frons deeply excavated from eye to eye and from epistoma to
vertex, the margin at vertex rather sharp; epistoma armed by two
long, tapered spines, each about equal in width at base to one-fourth
width of epistoma and in length to distance between epistoma and
vertex, the spines separated at bases by a distance equal to basal
width of one spine; spines and margin of excavation bearing a few
rather coarse, hairlike setae. Eye oval, entire. Antennal club sub-
circular, the sutures bisinuate. marked by rows of setae.

Pronotum wider than long; widest at base, sides rather strongly
arcuate on basal half, strongly constricted behind the rather narrow-
ly rounded anterior margin; summit well behind middle, its eleva-
tion continuing behind basal margin, ending posteriorly in a single
projecting cusp; surface rather coarsely punctured, each puncture
with a large granule or small asperity on its anterior or lateral mar-
gin, those toward summit slightly larger; vestiture scanty, hairlike.

Elytra 1.2 times as long as wide; sides weakly arcuate to base
of declivity, then more strongly rounded to lateral elevations of de-
clivity, deeply, rather broadly emarginate behind; striae not im-
pressed, coarsely, very deeply punctured; interstriae about as wide
as striae, the punctures in uniseriate rows, about one-third as large
as those of striae, deep, each armed on its posterior margin by a high,
pointed or rounded tubercle from base to apex. Declivity rounded,
vertical, deeply sulcate; interspace 1 moderately raised, striae 1 im-

The Great Basin Naturalist

38 s. L. WOOD Vol. XXVII, No. 1

pressed; lateral areas elevated, the summit reached near middle of
declivity on interspaces 3 and 4.

Female. — Similar to male except frons shallowly impressed,
epistoma unarmed, and general sculpture finer.

Type Locality. — Thirteen miles north of Juchitlan, Jalisco,
Mexico.

Host. — Giant cactus.

Type Material. — The male holotype, female allotype and 112
paratypes were taken at the type locality on July 2. 1965, at an
elevation of 3,000 feet, by S. L. Wood, from just under the epidermis
of a dying giant cactus.

The holotype, allotype and paratypes are in my collection.

Pseudothysanoes thomasi, n. sp.

This species is unique among described species in the genus. The
male has a prominent declivital elevation on the third interspaces
that ends abruptly iust below the middle of the declivity. The female
declivity is simple as in other species of the genus.

Male. — Length 1.1 mm., 2.5 times as long as wide; body color
light brov^Ti.

Frons broad, weakly convex below upper level of eyes, with
epistomal area distinctly elevated, more strongly convex above eyes;
surface rugulose and wdth a small, rounded granule at base of each
seta; vestiture consisting of moderately long and abundant fimbriate
setae, those above directed dorsad, those on epistoma directed orad.
Antennal scape short, subcircular. scarcely longer than pedicle, club
1.4 times as long as wide; pubescent, wdth sutures indistinctly in-
dicated.

Pronotum 0.94 times as long as wide; sides strongly arcuate,
widest just behind middle; summit distinct, the asperites moderately
large; anterior margin armed by eight coarse, basally separate teeth;
surface behind summit minutely reticulate, with a few small,
rounded, squamiferous granules; vestiture consisting of semierect
scales and a few inconspicuous bristles.

Elytra about 1.5 times as long as wdde; sides straight and sub-
parallel on basal two-thirds, rather broadly rounded behind; striae
feebly impressed, the punctures minute and distinguished with dif-
ficulty from those of interstriae; interstriae subrugose, the punctures
indistinct. Declivity convex, moderately steep; interspace 2 strongly
narrowed below, interspace 3 with a large unarmed elevation ex-
tending about two-thirds the length of declivity from base, ending
abruptly below, its base as wide as interspaces 1 and 2 combined,
its height slightly less than its basal width. Vestiture consisting of
rows of short, recumbent, strial, hairlike setae, and slightly longer
scalelike setae, each scale on posterior half at least as wide as long.

Female. — Similar to male except frons more strongly impressed,
strial punctures more strongly impressed, elytral declivity devoid of

April 22. 1967 new species of bark beetles 39

elevation, interspace 2 almost normal, and elytral scales slightly
more slender.

Type Locality. — Mazatlan. vSinaloa, Mexico.

Type M.\terial. — The male holotype, female allotype and seven
paratypes were taken at the type locality, evidently in August 1964,
from an unidentified tree by J. B. Thomas.

The holotype. allotype and part of the paratypes are in the
Canadian National Collection, other paratypes are in my collection.

Pityophthorus festus, n. sp.

This species is closely related to ponderosae Blackman, but is
easily distinguished by the regular striae on the basal half of the
elytra, by the more narrowly rounded elytral declivity, and by the
smaller average size.

Male. — Length 1.4 mm. (paratypes 1.3-1.5 mm.), 2.6 times as
long as wide; color dark brown.

Frons convex; epistoma broadly emarginate; surface coarsely,
deeply punctured, a sharply elevated, rather prominent median
carina of almost uniform height extending from just above upper
level of eyes to espistomal margin; vestiture sparse, inconspicuous
except on epistomal margin. Antenna as in ponedrosae.

Pronotum equal in length and width; widest on basal third, the
sides weakly arcuate and converging to weak constriction just be-
hind narrowly rounded anterior margin; anterior margin armed by
eight small teeth; summit at middle, moderately impressed behind;
posterior area rather coarsely, deeply, closely punctured; vestiture
short, sparse, inconspicuous.

Elytra 1.7 times as long as wide; sides straight and subparallel
on more than basal two-thirds, rather narrowly rounded behind;
striae in definite rows to base, the punctures small, rather deep;
interspaces flat, as wide as striae, marked by some surface lines,
impunctate. Declivity narrowly bisulcate, moderately steep; declivi-
tal punctures largely obsolete; interspace 1 moderately elevated, un-
armed, striae 1 sharply impressed, lateral area gradually raised and
rounded with the summit at striae 3; interspace 3 with a few very
minute granules; general surface almost smooth, shining. Vestiture
hairlike, fine, short, sparse.

Female. — Evidently indistinguishable from male except by seg-
mentation of abdominal terga.

Type Locality. — Eighteen miles west of El Salto, Durango,
Mexico.

Host. — Pinus sp. (type). P. ayacahuite and leiophylla (para-
types).

Type Material. — The male holotype, female allotype and two
paratypes were collected at the type locality on June 3, 1965. from
pine branches, at an elevation of 7.500 feet, by S. L. Wood; two
other paratypes bear the same data but were taken 3 miles west of
EU Salto from P. ayacahuite. Four paratypes were taken 23 miles

The Great Basin Naturalist

40 s. L. WOOD Vol. XXVn. No. 1

west of Durango. Durango, on June 4. 1965. from P. leiophylla;
five paratypes were taken 14 miles northwest of Guadalajara, Jalisco,
on July 19, 1953, from pine branches, all by S. L. Wood. Four addi-
tional paratypes were collected 10 miles west of El Salto, Durango.
during July 1964, by J. B. Thomas.

The holotype, allotype and some of the paratypes are in my col-
lection, other paratypes are in the Canadian National Collection.

Pityophthorus foratus, n. sp.

Among described species this species is more closely related to
schwerdtfergeri (Schedl) than to any other, but it is easily dis-
tinguished by the second declivital interspace which is almost twice
as wide as in Schedl's species.

Female. — Length 2.5 mm. (paratypes 2.1-2.5 mm.). 2.5 times
as long as wide; body color dark brown, the elytra a lighter brown.

Frons plano-convex from upper level of eyes to epistoma, rather
coarsely, closely punctured; vestiture very fine, hairlike, moderate
in length and abundance, as in schwerdtfergeri.

Pronotum very slightly wider than long; widest at base, sides
weakly arcuate and converging toward the narrowly rounded an-
terior margin; anterior margin armed by about eight low teeth;
summit at middle, definite; posterior area with rather coarse, close,
deep punctures, surface not entirely smooth, shining; vestiture fine,
inconspicuous.

Elytra 1.5 times as long as wide, 1.8 times as long as pronotum;
sides straight and subparallel on basal three-fourths, abruptly, very
broadly rounded behind; strial and interstrial punctures confused on
basal half of disc, striae 1 impressed near declivity and punctures of
1 and 2 recognizable, rather small, deep; punctures of interspace 1
recognizable at base of declivity, slightly smaller than those of striae.
Declivity flattened, steep; strial punctures reduced but distinct on
all striae; interspace 1 narrow, moderately elevated, unarmed. 2
impressed, very broad, three times as wide as 1, subreticulate, im-
punctate. 3 very narrow, narrower than 1, summit of lateral eleva-
tion reached on 4, unarmed. Vestiture limited to sides, fine, hairlike,
rather long.

Male. — Not represented in series at hand.

Type Locality. — Ten miles west of El Salto, Durango, Mexico.

Type Material. — The female holotype and six female paratypes
were collected at the type locality during July, 1964, in flight in a
pink-oak forest, at about 9.000 feet in elevation, by J. B. Thomas.

The holotype and some paratypes are in the Canadian National
Collection, other paratypes are in my collection.

Pityophthorus quercinus, n. sp.

Among described species this species should be placed near rhois
Swaine in Blackman's group IL although it is not closely related.

April 22. 1967 new species of bark beetles 41

The much larger size and entirely different sculpturing of pronotum
and elytral declivity distinguish it.

Female. — Length 2.1 mm. (paratypes 1.8-2.2 mm.). 3.0 times
as long as wide; color dark brown.

Frons broadly flattened from vertex to epistoma, with epistomal
area slightly elevated, the lateral margins not sharply defined; sur-
face closely, rather coarsely granulate-punctate and uniformly orna-
mented by moderately abundant, long, yellow hair of uniform
length. Antennal club rather small. 1.2 times as long as wide;
sutures 1 and 2 straight and septate, widest through segment 2, 1
and 3 about equal in vsddth.

Pronotum 1.2 times as long as wide; sides on basal half straight
and subparallel; anterior margin broadly rounded and armed by an
indefinite row of basally fused teeth; summit in front of middle, in-
definite; asperities on anterior area rather small, confused, not ex-
tending to base laterally; posterior area subshiiiing. closely, deeply,
rather finely punctured, and with numerous impressed points; disc
glabrous.

Elytra 1.8 times as long as wide; sides straight and subparallel
on more than basal two-thirds, rather narrowly rounded behind;
striae 1 impressed to base, others feebly if at all impressed, the
punctures rather small, deep, close; interspaces shining, weakly con-
vex, impunctate but with rather abundant impressed points.

Declivity bisulcate. moderately steep; all striae with punctures
distinctly impressed but slightly smaller than on disc; interspace 1
rather strongly elevated and armed by three wddely spaced, moder-
ately large, pointed granules. 2 impressed, flat, impunctate, 3 mod-
erately elevated but not equal to 1. armed by a row of about seven
rather large granules, lateral areas with punctures confused. Vesti-
ture hairlike, confined to sides and to setiferous granules on declivity.

Male. — Similar to female except frons convex, somewhat pro-
tuberant on upper half, more deeply, closely punctured, glabrous,
and elytral declivity conspicuously deeper and wider, with striae
obscure, the granules larger.

Type Locality. — ^Three miles west of El Salto, Durango, Mexico.

Host. — Quercus sp.

Type Material. — The female holotype, male allotype and 41
paratypes were taken at the type locality from the phloem tissues of
the bole of a very large oak^ on June 7. 1965. at an elevation of
7.500 feet by S. L. Wood. Twelve additional paratypes were col-
lected at 33 miles east of Moralia. Michoacan. Mexico, on June 14,
1965, at an elevation of 9,000 feet, from the same oak. by S. L. Wood.
Ten paratypes were also taken 10 miles west of El Salto. Durango.
Mexico, during July 1964. from oak. by J. B. Thomas.

The holotype. allotype and some of the paratypes are in my col-
lection; other paratypes are in the Canadian National Collection.

The Great Basin Naturalist

42 s. L. wcx)D Vol. XXVII. No. 1

Pseudopityophthorus hondurensis, n. sp.

Using Blackman's (1931, J. Ent. Soc. Washington 21:225) key
this species is placed near agrifoliae Blackman. However, it differs
in frontal vestiture in both sexes and in lacking the short setae of
striae and interstriae on the elytra.

Male. — Length 1.3 mm. (paratypes 1.2-1.6 mm.), about 2.8
times as long as wide; body color very dark brown, summit of pro-
notum somewhat lighter.

Frons plano-convex over a broad area, with a marginal fringe of
long, yellow hair, the median area punctured. Antennal club widest
through segment 3.

Pronotum 1.05 times as long as wide; sides straight and subpar-
allel on basal half, then moderately constricted before the broadly
rounded anterior margin; anterior margin armed by about 14 low
teeth; summit poorly developed, at middle; posterior area smooth
and shining, with rather sparse, coarse, deep punctures and more
numerous minute points; almost glabrous.

Elytra 1.5 times as long as wide; sides approximately straight
and parallel on basal two-thirds, almost semicircularly rounded be-
hind; striae not impressed, the punctures, fine, distinct but not deep,
in definite rows; interstriae almost flat, smooth and shining, with a
few fine lines and impressed points. Declivity convex, moderately
steep; strial punctures obsolete; interspaces except 2 with a few
minute setiferous grarmles. Vestiture mostly confined to declivity,
consisting of slender, erect, hairlike, interstrial setae.

Female. — Almost indistinguishable from male, but evidently
with frontal vestiture less abundant and shorter.

Type Locality. — Buenos Aires. Cortes, Honduras.

Host. — Quercus sp.

Type Material. — The male holotype. female allotype and 28
paratypes were taken from a dying oak branch at the type locality
on May 7, 1964. at an elevation of 7.000 feet by S. L. Wood.

The holotype, allotype and paratypes are in my collection.

Pseudopityophthorus granuUfer. n. sp.

This species is closely related to granulatus Blackman, but it is
much larger and has longer and more abundant frontal vestiture in
both sexes.

Female. — Length 1.8 mm. (paratypes 1.6-2.0 mm.). 2.8 times
as long as wide; color almost black.

Frons plano-convex over a broad area, with a dense marginal
fringe of long, yellow hair as in the male of most species. Second and
third segments of antennal club equal in width.

Pronotum 1.2 times as long as wide; sides gradually, arcuately
narrowed toward the rather broadly rounded anterior margin; an-
terior margin armed by about 14 teeth; summit indefinite, at middle;
posterior area smooth and shining, with fine, rather sparse, deep

April 22, 1967 new species of bark beetles 43

punctures and more abundant minute points; vestiture limited to
sides and asperate area.

Elytra 1.5 times as long as wide; sides almost straight and sub-
parallel on basal two-thirds, rather narrowly rounded behind; striae 1
weakly impressed, the others not at all impressed, the })unctures very
small, shallow, in rather indefinite rows; interstriae obscure, with a
few punctures, points and surface lines. Declivity convex, rather
steep, with interspace 2 impressed; strial punctures obsolete; inter-
spaces 1. 2 and 3 each armed by a row of rounded granules that de-
crease in size and number toward apex; declivital surface smooth,
dull. Vestiture consisting of rather sparse, erect bristles arranged in
fine strial and coarse interstrial rows; some interstrial bristles on
declivity strongly flattened.

Male. — Evidently exactly like the female, distinguishable only
by segmentation of abdomen.

Type Locality. — Zamorano, Morazan, Honduras.

Hosts. — Quercus sapotaefolia (type), and hondurensis (para-
tvpe) .

Type Material. — The female holotype, male allotype and 10
paratypes were taken from Quercus sapotaefolia and five paratypes
from Q. hondurensis at the type locality on April 18, 1964, at an ele-
vation of 2.200 feet, by S. L. Wood. One other paratype was taken in
flight at Guatemala City. Guatemala, May 30, 1964. A specimen
probably belonging to this species, but not designated as a paratype.
was collected at San Cristobal de las Casas. Chiapas, Mexico, on July
5. 1956, by D. D. Linsdale.

The holotype. allotype and paratypes are in my collection.

Pseudopityophthorus tropicalis, n. sp.

This species evidently is allied to pulvereus Blackman, but is
readily distinguished by the rows of erect declivital scales.

Male. — ^Length 1.7 mm. (paratypes 1.5-1.7 mm.), 2.8 times as
long as wide; color rather dark brown with white vestiture.

Frons plano-convex from eye to eye from vertex to epistoma. and
ornamented at margins by a fringe of abundant, long, yellow hair,
central area concealed in all males at hand. Antennal club elongate,
about 1.5 times as long as wide, with segments 2 and 3 equal in
width.

Pronotum 1.1 times as long as wide; widest at base, sides weakly
arcuate and gradually narrowed anteriorly, weakly constricted just
before anterior margin; anterior margin rather broadly rounded and
armed by about 14 low teeth; summit indefinite; posterior area fine-
ly, closely, deeply punctured, and with abundant impressed points;
vestiture largely confined to sides and as})erate area.

Elytra 1.5 times as long as wide; sides straight and parallel on
basal two-thirds, semicircularly rounded behind; striae and inter-
striae indistinguishable, the punctures fine, shallow, confused. De-

The Great Basin Naturalist

44 s. L. WOOD Vol. XXVII. No. 1

clivity convex, rather steep, interspace 2 rather strongly impressed.
Vestiture abundant, consisting of short semierect bristles, narrow an-
teriorly, almost scalelike posteriorly, and rows of longer erect scales
on declivital interspaces 1, 3 and 4, each scale shorter than twice
the length of the ground vestiture and only slightly wider.

Female. — ^Similar to male except frons more coarsely punctured,
with the vestiture shorter, less abundant and more uniformly distri-
buted.

Type Locality. — Zamorano, Morazan, Honduras.

Hosts. — Quercus sapotaefolia (type), and hondurensis (para-
type) .

Type Material. — The male holotype. female allotype and 39
paratypes from branches of Quercus sapotaefolia and 9 paratypes
from Q. hondurensis were taken at the type locality on April 19,
1964, at an elevation of 2,200 feet, by S. L. Wood. Five other para-
types were collected at Volcan de Agua. Guatemala, on May 19,
1964, from Quercus branches, at an elevation of 3.000 feet by S. L.
Wood. Six additional paratypes were taken 7 miles northeast of
Copala, Sinaloa, Mexico, on .luly 22, 1953. at an elevation of 3,000
feet, by S. L. Wood.

The holotype, allotype and paratypes are in my collection.

Pseudopityophthorus micans, n. s]).

This species evidently is allied to opacicollis Blackman, but it is
unique in having the pronotum and elytra glabrous, smooth and
shining.

Male. — Length 2.2 mm. (paratypes 2.0-2.5 mm.). 2.7 times as
long as wide; color dark brown.

Frons plano-convex on three-fourths of a circle, smooth and shin-
ing at center with a marginal fringe of closely placed, long, erect,
hairlike setae, marginal pubescent area closely, finely granulate-
punctate. Eye deeply emarginate; finely granulate. Antennal club
widest through segment 3.

Pronotum 1.1 times as long as wide; sides weakly arcuate, almost
parallel on basal half, then moderately constricted before the rather
narrowly rounded anterior margin; anterior margin armed by about
14 small teeth; summit indefinite; posterior area behind summit
smooth and shining with rather abundant fine punctures and minute
points; glabrous except at margins.

Elytra 1.6 times as long as wide; sides straight and subparallel
on basal two-thirds, abruptly narrowed at posterolateral angles,
rather narrowly rounded behind; elytral punctures largely limited to
anterior two-thirds of disc. fine, confused except for striae 1 ; surface
smooth and shining but marked with a few fine lines. Declivity
abruy^t with apical and sutural area somewhat produced posteriorly;
surface smooth and shining. Glabrous except for a few setae at an-
terolateral angles.

April 22. 1967 new species of bark beetles 45

Female. — Similar to male except frons finely. deej)ly punctured
except for a narrow median line; frontal vestiture sparse, scattered;
and declivital punctures minute but evident.

Several paratypes are more distinctly punctured on the elytral
declivity than the type. Some also appear dull, rather than shining;
the shine evidently disappears with advancing age.

Type Locality. — Sixty miles west of Durango, Durango, Mexi-
co.

Host. — Quercus spp.

Type M.\terial. — The male holotype, female allotype and 23
paratypes were taken at the type locality on June 5, 1965, at an
elevation of 7,000 feet, from a dying oak tree about 10 inches in
diameter, by S. L. Wood. Other paratypes were collected in Mexico
as follows: seven from 20 miles west of Durango, Durango; four
from 18 miles west and one from three miles west of EJ Salto. Du-
rango; four from 33 miles east of Moralia, Michoacan; and three
from four miles south of Carapan. Michoacan; all were taken from
oak trees, between 7.000 and 9,000 feet in elevation during June
1965. by S. L. Wood. Thirteen additional paratypes were taken 10
miles west of El Salto. Durango, in July 1964. by J. B. Thomas.

The holotype. allotype and most of the paratypes are in my col-
lection; other paratypes are in the Canadian National Collection.

Gnathotrichus dentatus, n. sp.
Fig. 1

This unique species differs from all previously described species
in having the elytral declivity of the male rather deeply sulcate and
armed by a pair of large hi- or tridentate processes. The pubescence
of pronotum and declivity is also much more conspicuous than in
previously known species.

M\le. — Length 3.4 mm. (paratypes 3.1-3.6 mm.), 3.1 times as
long as wide; color dark brown.

Frons very broad, convex; surface rather coarsely, shallowly
punctured becoming increasingly aciculate toward median portion of
epistoma; a short, transverse, elevated carina in median area at
upper level of eyes and from it an impunctate line extending dorsad;
vestiture short, inconspicuous.

Pronotum 1.2 times as long as wide; sides straight and parallel
on basal half, rather broadly rounded in front; anterior margin
armed by 10 low serrations; summit indefinite, in front of middle;
posterior area smooth, subshining, finely, shallowly. rather sparsely
punctured; vestiture fine, short, rather abundant on sides.

Elytra 2.0 times as long as wide; sides almost straight and paral-
lel on basal three-fourths, narrowly rounded behind and narrowly
notched at suture; punctures obsolete, striae and interstriae not in-
dicated; surface very finely marked by surface lines and indefinite,
shallow punctures, dull. Declivity with a broad, moderately deep

46

S. L. WOOD

The Great Basin Naturalist

Vol. XXVIl, No. 1

I. G. dentatus

2. M. quercicolens

5. Micr. desus

4. M, bicavum 3, M, querneus

Figs. 1-5. Posterolateral aspect of male elytral declivity; 1, Gnathotrichus
dentatus; 2, Monarthrum quercicolens; 3, M. querneus; 4, M. bicavum; and 5,
Microcorthylus desus.

sulcus between the pair of prominent processes borne on interspace
3; lateral processes supported on prominent lateral elevations, the
processes arising from median edge of elevation and bearing on dor-
sal margin a series of (two or) three rather large teeth; the teeth
directed caudad and somewhat mesad. increasing slightly in size
posteriorly; about three small dentitions on third interspace anterior
to process, and with three similar small tubercles on interspace 1 at
base of declivity; elytral apex explanate and narrowl}^ notched. Ves-
titure consisting of rather abundant minute hairs and some longer,
erect, coarse, hairlike setae.

Fem.\le. — Similar to male except frons less strongly convex and
devoid of carina; anterolateral angles of pronotum wdth a small tuft
of hair; and elytral declivity with sulcus shallow, lateral processes
not strongly developed, and dentitions absent except for one or two
small, rounded granules.

Type Locality. — Eighteen miles west of El Salto, Durango,
Mexico.

Host. — Quercus sp.

Type Material. — The male holotype, female allotype and 20
paratypes were taken at the type locality on June 7, 1965, from the
bole of a very large oak. at an elevation of 7,500 feet, by S. L. Wood.
Other paratypes include: 11 same data as holotype but taken three
miles west of El Salto; four taken 10 miles west of El Salto during
July 1964, by J. B. Thomas; one taken 16 miles east and 21 taken
33 miles east of Moralia. Michoacan, Mexico, on June 14. 1965,
from a Quercus bole, at an elevation of 9,000 feet by S. L. Wood.

The holotype. allotype and some paratypes are in my collection,
other paratypes are in the Canadian National Collection.

April 22, 1967 new species of bark beetles 47

Gnathotrichus nimifrons, n. sp.

Tliis species is allied to dentatus Wood, described above, but is
readily distinguished by its smaller size, by the narrower and
longer declivital sulcus with the lateral processes undeveloped.

Male. — Length 3.0 mm. (allotype 2.7 mm.), 3.3 times as long
as wide; color brown.

Frons very broad, strongly convex; surface smooth and shining
in central area, shallowly punctured laterally, weakly aciculate to-
ward epistoma; vestiture sparse, inconspicuous.

Pronotum 1.3 times as long as wide; as dentatus except posterior
areas more finely, shallowly and closely punctured.

Elytra 1.9 times as long as wide; sides straight and subparallel
on basal three-fourths, rather narrowly rounded behind, shallowly
notched at suture; striae and interspaces not indicated; surface
minutely irregular, with rather sparse, shallow, fine punctures. De-
clivity rather narrowly, deeply sulcate, moderately steep; suture
not raised, lateral area including interspace 3 strongly elevated from
declivital base about two-thirds the distance to apex, the crest armed
by about seven small, pointed granules; elytral apex explanate, nar-
rowly notched. Vestiture consisting of fine, short hairlike setae on
disc and sides, becoming much longer and abundant on declivity.

Female. — Similar to male except frons less strongly convex and
with fine aciculate lines extending at least to upper level of eyes; and
declivity shallowly sulcate, the lateral elevations poorly developed,
the granules fewer in number and smaller.

Type Locality. — Eighteen miles west of El Salto, Durango,
Mexico.

Host. — Quercus sp.

Type Material. — The male holotype, female allotype and one
male paratype were collected at the type locality on June 7, 1965,
at an elevation of 7.500 feet, by S. L. Wood, from the bole of the
same large oak tree that contained the holotype of dentatus. One
paratvpe is from Carr Canyon, Arizona, taken at blacklight, on
August 8, 1962, by S. L. Wood.

The holotype, allotype, and paratypes are in my collection.

Gnathotrichus perniciosus, n. sp.

This species is allied to, but not closely related to deleoni Black-
man. It is distinguished by the smaller size, by the more slender
form, by the very different frons. by the striate elytral disc, by the
sculpture of the elytral declivity, and by other characters.

Male. — Length 2.4 mm. (paratypes 2.2-2.6 mm.), 3.2 times
as long as wide; color dark brown, the elytral bases somewhat light-
er.

Frons moderately convex; surface convergently aciculate from
upper level of eyes to median point on epistoma, two of the ridges
on median line (one in some paratypes) more strongly raised, par-

The Great Basin Naturalist

48 s. L. WOOD Vol. XXVII, No. 1

ticularly above, the grooves between ridges and above aciculate area
rather sparsely, deeply, coarsely punctured.

Pronotum 1.2 times as long as wide; widest at base, the sides
straight, converging anteriorly very slightly on basal three- fourths,
then broadly rounded in front; anterior margin armed by about 12
low, blunt teeth; summit well in front of middle, marked by a raised
transverse line; posterior area finely reticulate, dull, very finely,
rather sparsely punctured; vestitute inconspicuous, confined to sides.

Elytra 1.8 times as long as wide; sides almost straight and paral-
lel on basal three-fourths, narrowly rounded behind, with a very
small notch at suture; striae not impressed, the punctures in semi-
definite rows, small, shallow; interspaces with surface lines, points,
and an occasional puncture, subshining. Declivity steep, narrowly
and shallowly sulcate; striae 1 and 2 indicated by fine, shallow
punctures; interspaces 1 and 2 very narrow, impressed, 3 moderate-
ly, rather abruptly elevated and armed by about four rounded
granules. Vestiture limited to declivity, fine, hairlike, short and a
few longer setae intermixed.

Female. — Similar to male except outer margin of antennal
funicle and club bearing a few long setae.

Type Locality. — Six miles south of Carapan, Michoacan, Mexi-
co.

Hosts. — Pinus spp.

Type Material. — The male holotype, female allotype and 69
paratypes were collected at the type locality on June 18, 1965. at an
elevation of 7,000 feet, from the base of a pine 12 inches in diameter,
by S. L. Wood. Other paratypes. all taken in Mexico, include: two
from Tres Rios, Chihuahua, July 13, 1959, by W. W. Tanner and G.
W. Robison; one from 20 miles northeast of Copala, Sinaloa, July 22,
1953, from Pinus, by S. L. Wood; three from the southeast slopes
of Mt. Colima, Colima, Dec. 2, 1948, by E. S. Ross; two from near
Cuernavaca, Morelos, August 5, 1949. by J. P. Perry; one from
seven miles south of Mazamitla, Jalisco, December 1, 1948, by H. B.
Leech; and one from 15 miles south of El Guarda, Distrito Federal,
November 14, 1946, by E. S. Ross. This species was also taken at
Cerro Pena Blanca. San Lucas, Yuscaran. and Zamorano, Honduras,
from Pinus oocarpa and pseudostrobus, but the specimens from these
localities are not designated paratypes.

The holotype, allotype and some of the paratj^pes are in my col-
lection, other paratypes are in the California Academy of Sciences.

Gnathotrichus imitans, n. sp.

This species evidently is more closely allied to perniciosus
Wood, described above, than to other known forms, but it is easily
distinguished by the black or dark brown color, by the impressed
lower half of the frons, and by the more strongly striate elytra.

Adult. — Length 3.0 mm. (paratypes 2.9-3.1 mm.), 3.3 times as
long as wdde; color dark brown.

April 22. 1967 new species of bark beetles 49

Frons impressed on more than median half from upper level of
eyes to epistoma, convergently aciculate toward median point on
epistomal margin on most of impressed area; surface coarsely, rather
sparsely, deeply punctured over most of front of head to vertex; a
median carina extending dorsad from uj)per level of eyes; vestiture
sparse, inconspicuous.

Pronotum 1.4 times as long as wide; widest at base, sides almost
straight on basal three-fourths and slightly converging anteriorly,
rather broadly rounded in front; anterior margin with an irregular,
indistinctly serrate, raised rim; summit well in front of middle,
clearly marked by a transverse elevation; posterior area almost
smooth and shining, rather closely, deeply, finel)^ punctured; vesti-
ture restricted to sides.

Elytra 1.9 times as long as wide; sides straight and subparallel
on basal three-fourths, rather narrowly rounded behind; striae not
impressed, the punctures small, deep; interstriae little wider than
striae, marked by a few surface lines and points, subshining. De-
clivity moderately steep, convex; all punctures minute, not clearly
impressed; interspace 1 weakly elevated, 2 impressed, 3 slightly
elevated and armed by a series of about five very small granules;
apical margin not extended as in most representatives of the genus.
Vestiture confined to sides and declivity; longer and in distinct rows
on declivital interspaces 3 and 4.

Sexual differences are not apparent in the material at hand, al-
though both sexes probably are represented.

Type Locality. — Three miles west of El Salto, Durango, Mexico.

Type Material. — The holotype (a female?) and six paratypes
were collected at the type locality on June 7, 1965, at an elevation
of about 7,500 feet, in flight, by S. L. Wood. Other paratypes were
collected in flight near the type locality as follows: 4 from El Salto,
June 3, 1937, by Juan Manual, and 69 from 10 miles west of Ell
Salto. during July 1964. by J. B. Thomas.

The type and some of the paratypes are in my collection, other
paratypes are in the California Academy of Sciences and the Can-
adian National Collection.

Monarthrum quercicolens, n. sp.
Fig. 2

Of the described species known to me this species evidently is
more closely alhed to scutellare (Leconte), but it is not closely
related. The third pair of elevated processes on the declivity and the
pubescent female frons serve to distinguish it from other species.

Male. — Length 4.0 mm. (paratypes 3.7-4.3 mm.), 3.1 times as
long as wdde; pronotum brown, elytral base light brown, posterior
half of elytra dark brown.

Frons convex, with a feeble median elevation below; surface
reticulate below, subshining above, finely, deeply, rather closely

The Great Basin Naturalist

50 s. L. WOOD Vol. XXVII No. 1

punctured; glabrous. Antennal club 1.8 times as long as wide, widest
one-third from apex.

Pronotum 1.4 times as long as wide; sides almost straight and
parallel on basal two-thirds, broadly rounded in front; anterior mar-
gin not clearly armed although several irregular, submarginal asper-
ities present; asperities restricted to slightly more than anterior third;
posterior area subreticulate, dull, with sparse, minute, shallow punc-
tures; glabrous.

Elytra 1.9 times as long as wide; sides straight and subparallel
on basal three-fourths, abruptly rounded behind, shallowly emar-
ginate toward suture; elytral punctures very fine, more or less in
rows, those of striae and interstriae similar in size and depth; surface
marked by irregular surface lines and points, subshining. Declivity
broadly, concavely excavated, abrupt, moderately steep; interspaces
2 and 3 each armed at base by a rather large, sharply pointed tooth;
lateral margin raised, abruptly rounded from second tooth to apex
with its middle third abruptly produced forming a long, flattened
process on lateral margin about equal in height to teeth on upper
margin; sutural margin feebly elevated, the concavity broadly con-
cave, its surface rather coarsely, deeply punctured. Elytra glabrous,
except in declivital excavation rather densely pilose, the setae fine,
rather long, with marginal row of setae distinctly longer.

Female. — Similar to male except frons less strongly convex, wdth
a small, shallow, median impression and subtriangular patch of
rather dense, fine, erect hair on median third between upper level
of eyes and epistoma; declivity not excavated, with two pair of small
tubercles on interspace 2; and antenna with tuft of hair characteristic
of genus.

Type Locality. — Thirty-three miles east of Moralia, Michoacan,
Mexico.

Host. — Quercus sp.

Type Material. — The male holotype, female allotype and three
para types were taken at the type locality on June 14. 1965, from a
large oak log, at an elevation of 9,000 feet, by S. L. Wood. Other para-
types were taken in Mexico from oak as follows: five from Carapan,
Michoacan. June 18, 1965, 7,000 feet elevation; one from three miles
west of El Salto, Durango, June 7, 1965, 7,500 feet elevation, all by
S. L. Wood; six from 10 miles west of El Salto. during July 1964,
by J. B. Thomas.

The holotype, allotype and some of the paratypes are in my col-
lection, other paratypes are in the Canadian National Collection.

Monarthrum querneus, n. sp.

Fig. 3

This species is related to quercicolens Wood, described above, but
is readily distinguished in the male by the smaller size, by the shal-
lower, more widely flattened elytral declivity, and by the slightly

April 22, 1967 new species of bark beetles 51

longer third pair of declivital processes; the female frons is glabrous
and has a moderately large cavity on the vertex.

Male. — Length 3.1 mm. (para types 2.8-3.4 mm.), 3.3 times as
long as wide; color dark brown with sides and base of pronotum
and basal two-thirds of elytra lighter.

Frons convex, with a low median elevation below; surface coarse-
ly reticulate below, becoming smooth above with rather close, shal-
low punctures and numerous fine points; glabrous.

Pronotum and elytral disc as quericicolens. Declivity about as
in quercicolens but concavity more broadly flattened, the apical
margin less strongly produced, and the flattened lateral processes
a little shorter and higher with the declivital vestiture less well de-
veloped.

Female. — Similar to female of quercicolens except frons convex
and glabrous with moderately large impression on vertex as wide
as antennal club and shorter than wide, the impression abrupt above,
gradual below, its depth equal to less than one-fourth its width.

Type Locality. — Thirty-three miles east of Moralia, Michoacan,
Mexico.

Host. — Quercus sp.

Type Material. — The male holotype, female allotype and 42
para types were taken at the type locality on June 14, 1965, at an
elevation of 9,000 feet, by S. L. Wood, from the same large oak log
that contained the holotype of quercicolens.

The holotype, allotype and paratypes are in my collection.

Monarthrum bicavum, n. sp.
Fig. 4

While the declivital structure of this species has a superficial re-
semblance to that of vittatum Blandford, the two are not closely re-
lated. In this species the apical margin of the declivity forms a finely
raised, continuous line across the entire posterior margin of the
declivity without a sutural emargination. The female frons is also
unique in being shallowly concave from eye to eye and from vertex
to epistoma with a low median elevation dividing it.

Female. — Length 2.9 mm. (paratypes 2.9-3.3 mm.), 3.0 times
as long as wdde; color dark brown.

Frons shallowly concave from eye to eye and from vertex to
epistoma, its margin abruptly rounded; concavity divided by a
median carina, subacute above, broad below; each half of concavity
with an almost hemispherical impression at upper level of eyes about
equal in diameter to greatest width of eye; entire surface of concav-
ity and carina densely covered by very short pilose hair. Antennal
club 1.4 times as long as wide, widest one-third from distal end,
marked by two strongly arcuate sutures; posterior face bearing tuft
of hairlike setae as in other species of the genus.

The Great Basin Naturalist

52 s. L. WOOD Vol. XXVII. No. 1

Pronotum 1.2 times as long as wide, subquadrate, the sides
weakly arcuate, anterior margin only slightly more strongly arcuate,
the anterior angles only slightly more broadly rounded than those of
base; siimmit two-fifths of pronotum length from anterior margin,
poorly developed; posterior area minutely reticulose-granulose. im-
punctate; glabrous.

Elytra 1.5 times as long as wide; sides almost straight and paral-
lel on basal two-thirds, then slightly convergent posteriorly to
declivital base, then rather abruptly angled and broadly rounded be-
hind; strial and interstrial punctures not in definite rows, but not
entirely confused, the punctures small, not deep; surface shallowly
reticulate and with surface lines, subshining. Declivity rather abrupt,
very steep, sulcate on upper half; punctures not clearly evident but
sutural interspace feebly raised, 2 narrow, impressed about to middle
of declivity, 3 rather strongly raised above but disappearing by
middle of declivity and armed by a small tubercle near upper limit
and another at middle of declivity, the lower tubercles more widely
separated from one another than the upper; the lower third of lateral
margin marked by an acute, low. elevated line continuing without
interruption to apex, not emarginate at suture; surface minutely
reticulose-granulose. Vestiture scanty, restricted to sides and decliv-
ity.

Male. — Similar to female except frons weakly convex, with a
broad irregular median carina, the surface granulose with sparse,
shallow punctures; posterior area of pronotum with shallow punc-
tures indicated; and declivity more broadly flattened above, the ele-
vation of interspace 3 forming an indefinite crescent on middle third
of declivity, and three or four fine tubercles may occur.

Type Locality. — Laguna Santa Maria, Nayarit, Mexico.

Host. — Quercus sp.

Type Material. — The female holotype, male allotype and 19
paratypes were collected at the type locality on July 16, 1965, from
the bole of a fallen oak, at an elevation of 3,000 feet, by S. L. Wood.

The holotype, allotype and paratypes are in my collection.

Microcorthylus desus, n. sp.
Fig. 5

Of the described species known to me none are closely related
and, at best, only superficially resemble this species. Superficially
it might be placed intermediate between Monarthrum egenum
Blandford and M. edentatum Eggers. The subvertical declivity and
nonemarginate elevated apex of the elytra distinguish this species.
In some respects it might resemble M. bicavum Wood, described
above.

Male. — Length 1.9 mm. (paratypes 1-8-2.1 mm.). 2.5 times as
long as wide; color brown.

April 22. 1967 new species of bark beetles 53

Frons convex, coarsely, not closely or deeply punctured, with a
small, elongate median tubercle at upper level of eyes; vestiture in-
conspicuous. Antennal club oval, 1.5 times as long as wide, widest
at middle, two sutures very weakly angulate; funicle one-segmented.

Pronotum 1.3 times as long as wide; sides straight and subparal-
lel on basal half, rather broadly rounded in front; anterior margin
unarmed; summit at middle; posterior area reticulate, dull, very
finely, sparsely punctured; disc glabrous.

Elytra about 1.5 times as long as wide; sides straight and parallel
to declivital base, posterior margin straight, the posterior angles
curving caudomesad to form lower declivital spines with their apices
directed mesad; elytral punctures small, indistinct, confused, the
surface reticulate, dull. Declivity abrupt, subvertical. somewhat more
gradually rounded at base near suture; face broadly, rather shallow-
ly concave, the lateral margin rather strongly, broadly elevated and
armed by a rather large, pointed tubercle on upper third about in
line with normal position of striae 3 and a larger incurved, hooked
spine two-thirds of declivital length from top; height of lower spine
about equal to its basal width, its apex directed mesad; apical mar-
gin acutely, not strongly elevated on median half of elytral width,
not at all emarginate at suture. Vestiture not conspicuous, limited to
sides and declivity.

Female. — Similar to male except elytral declivity less strongly
sculptured, only moderately impressed, the lateral areas not high, the
tubercles small; apical margin of elytra, frons and antenna as in
male.

Type Locality. — Sixteen miles east of Moralia. Michoacan,
Mexico.

Host. — Quercus sp.

Type Material. — The male holotype. female allotype, and 18
paratypes were collected at the type locality on June 14, 1965, at
an elevation of 7.600 feet, by S. L. Wood, from the dying limb of
an oak.

The holotype, allotype and paratypes are in my collection.

Amphicranus quercus, n. sp.
Figs. 6-7

This species is allied to torneutes Blandford. but it is much small-
er, it lacks a frontal callosity, and it has only one rounded and one
pointed tooth near suture at base of male declivity.

Male. — Length 4.5 mm. (para type males 4.4-4.6, females 3.4-
3.8 mm.). 3.4 times as long as wide; color dark brown.

Frons convex, the surface rather coarsely, shallowly punctured
except small area at center; epistoma bearing a sparse, poorly de-
veloped brush of hair directed orad. Antennal funicle two-seg-
mented; club with sutures 1 and 2 evenly, rather weakly arcuate.

54

S. L. WOOD

The Great Basin Naturalist

Vol. XXVIL No. 1

Pronotum 1.8 times as long as wide; sides straight and parallel
on basal two-thirds, then anteriorly narrowed slightly to an abrupt
subangulate point and broadly rounded in front; anterior area rather
abruptly declivous, the indefinite sununit one-third from anterior
margin; asperities fine, confused except a small area equal in size
to antennal club medially from summit forward, slightly impressed
and devoid of asperities, its surface reticulate; posterior area indis-
tinctly reticulate, dull with fine. s|)arse punctures; glabrous.

Elytra 2.1 times as long as wide; sides straight and almost paral-
lel from base to level even wdth apex of third declivital teeth, very
slightly wider just before apex of elytra, broadly rounded behind
and very deeply, broadly emarginate; depth of cleft equal to one-
fifth elytral length, its width equal to one-half its depth; striae very
indistinctly indicated by minute, shallow punctures, interstriae with
a few confused similar punctures, surface essentially smooth, dull.
Declivity abrupt, sharply margined above, excavated, with lateral
areas very strongly produced, floor of cavity broadly flattened,
rather steep; upper third sharply margined by a low elevation
about as high as thick, its sutural half bearing two almost equal
dentitions, one at interspace 2, pointed, the other bluntly rounded at
sutural interspace; lower two-thirds of lateral margin abruptly, very
strongly produced posteriorly, upper margin of this process horizon-
tal to level equal with sutural apex, then armed by a large, short,
flat, blunt posterodorsally projecting process, and continuing postero-
ventrally to acute, very deep sutural emargination. Vestiture con-
fined to declivity, consisting of rather abundant short, fine, erect
hair in the excavated area, with much longer setae on margins of
expanded lateral areas.

7A. quercuE

9. C. p e t i I u s

8, A. rameus

Figs. 6-9. Male elytral declivity of: 6-7, Amphicranus quercus, posterolateral
and dorsal aspects; 8, A. rameus, dorsal aspect; and 9, Corthylus petilus, postero-
lateral aspect.

April 22. 1967 new species of bark beetles 55

Female. — Similar to male except inner face of antennal club
bearing a tuft of hair, impressed reticulate area in front of pronotal
summit absent, and declivity simple, flattened, with a low, acute
margin on lower half only and armed by two pair of moderately
small, pointed tubercles on interspace 3, one near upper margin, the
other near middle of declivity; declivity subglabrous.

Type Locality. — Three miles west of EJ Salto, Durango, Mexi-
co.

Host. — Quercus sp.

Type Material. — The male holotype. female allotype and 44
para types were collected at the type locality on June 27, 1965, at
an elevation of 7,500 feet, by S. L. Wood, from the boles of several
large oak trees.

The holotype, allotype and paratypes are in my collection.

Amphicranus rameus, n, sp.

Fig. 8

This species is similar to propugnatus Blandford, but it may be
distinguished by the presence of a specialized, raised, coarsely reti-
culate, frontal area, and by very different arrangement of the elytral
declivity.

Female. — Length 3.7 mm. (paratypes 3.5-3.7 mm.), 3.7 times
as long as wide; color of pronotum in front of summit, elytral de-
clivity and abdomen almost black, remainder of pronotum reddish
brown, remainder of elytra yellowish brown.

Frons convex, its surface reticulate and rather coarsely, indis-
tinctly punctured, median third with a low, sharply raised, flat,
coarsely reticulate area having outline of a cup (more nearly heart-
shaped in paratypes); vestiture inconspicuous. Antennal funicle two-
segmented; club with two non-septate weakly arcuate sutures.

Pronotum 1.5 times as long as wide; sides straight and parallel
on basal three- fifths, rather narrowly rounded in front; anterior
margin projecting downward very slightly at center, armed by
about 10 low teeth, the median pair more prominently extended;
summit three- fifths of pronotum length from base, indefinite; rather
strongly declivous and coarsely asperate in front of summit, posterior
area finely reticulate vsdth small, shallow, rather sparse punctures;
glabrous.

Elytra 2.2 times as long as wide; sides straight and parallel to
a point even with sutural apejf. posterolateral angles convergently
projecting behind beyond the broadly emarginate elytral apex; disc
irregularly smooth, shining, subsurface punctures suggested but not
attaining surface. Declivity abrupt, moderately steep, broadly ex-
cavated; margin subacutely elevated from interspace 1 about two-
thirds of distance from top of declivity, the elevation not as high
as wide, then continued on lower third to projecting subquadrate.
converging lateral processes, the processes about one and one-half

The Great Basin Naturalist

56 s. L. WOOD Vol. XXVII. No. 1

times as long as their width in lateral profile; lateral margin armed
by small, equal, pointed spines on interspaces 1 and 3 at base of
declivity, and at interspace 4 by a slender blunt tooth about twice
as long as upper ones; lateral processes without special armature;
median two-thirds of posterior margin emarginate, the emargination
about half as deep as wide. Vestiture fine, scanty, confined to de-
clivity.

Male. — Similar to female except frontal raised callus absent; a
smaller tuft of hair present on antennal club.

Type Locality. — Three miles west of El Salto, Durango. Mexico.

Host. — Quercus sp.

Type Material. — TTie female holotype. male allotype and two
female paratypes were collected at the type locality on June 7, 1965.
at an elevation of 7.500 feet, by S. L. Wood, from a limb four
inches in diameter of the same large oak species that produced
quercus Wood (above).

The holotype. allotype and paratypes are in my collection.

Corthylus petiliis, n. sp.
Fig. 9

Although this species is truncate behind as in compressicornis
(Fabricius), it is not at all closely related. It differs from com-
pressicornis by the smaller, more slender body, and in the female,
by the absence of a tuft of hair on the antennal club and the presence
of a pale, spongy area on the epistomal area.

Female. — Length 2.0 mm. (paratypes 1.9-2.1 mm.). 2.5 times
as long as wide; color dark brown.

Frons rather deeply concave from eye to eye and from epistoma
to vertex; upper three-fourths rather coarsely reticulate, finely,
shallowly punctured, shining; lower fourth spongy, dull, pale in
color; margin above eyes ornamented by a fringe of rather long,
plumose hair. Antennal club much as in compressicornis, without a
tuft of hair.

Pronotum about equal in length and wddth; widest on basal
third, sides weakly arcuate, very boardly rounded in front; summit
well in front of middle, rather steeply declivous in front; posterior
area reticulate, becoming almost smooth and shining at base, finely,
sparsely punctured; glabrous.

Elytra 1.7 times as long as wide; sides almost straight and
parallel to truncate declivity, very weakly rounded behind; strial
and interstrial punctures small, distinctly impressed, confused ex-
cept striae 1 and 3 almost discernible toward declivity; surface with
a few obscure lines and points, shining. Declivity subvertical. trun-
cate, the margin abruptly rounded on upper half, acute below; rather
strongly impressed on a complete circle, particularly below; sutural
interspace strongly elevated on middle two-thirds, gradually reduced

April 22. 19(i7 new species of bark beetles 57

above, more abruj)tly reduced below and obsolete before apex; sur-
face smooth and shining except coarsely reticulate on sutural inter-
space, the punctures coarse, deep; sutural interspace armed by a row
of about nine rounded granules. Almost glabrous.

M.\LE. — Similar to female except frons convex, reticulate, with
a few deep, coarse punctures; anterior margin of pronotum more
strongly rounded and armed by two closely set, moderately large
teeth.

Type Locality. — Three miles west of El Salto, Durango, Mexi-
co.

Host. — Quercus sp.

Type Material. — The female holotype, male allotype and three
para types were collected at the type locality on June 7. 1965, at an
elevation of 7,500 feet, by S. L. Wood, from the same oak branch
that contained the type series of Amphicranus rameus Wood
(above).

The holotype, allotype and paratypes are in my collection.

The Great Basin Naturalist

Founded in 1939 by Vasco M. Tanner

A journal published from one to four times a year by Brigham
Young University, Provo, Utah.

Manuscripts: Only original, unpublished manuscripts, pertain-
ing to the Great Basin and the v/estem United States in the main,
will be accepted. Manuscripts are subject to the approval of the
editor.

Illustrations: All illustrations should be made with a view to
having them appear within tlie limits of the printed page. The illus-
trations that form a part of an article should accompany the manu-
script. All half-tones or zinc etchings to appear in this journal are
to be made under the supervision of the editor, and the cost of the
cuts is to be borne by the contributor.

Reprints: No reprints are furnished free of charge. A price list
for reprints and an order form is sent with the proof.

Subscription: The annual subscription is $2.50 (outside the
United States $3.25). Single number, 80 cents.

All coirespondence dealing witli manuscripts should be addressed
to the Editor, Vasco M. Tanner, Great Basin NaturaHst, Brigham
Young University, Provo, Utah. Other matters such as subscriptions,
reprints, exchanges and other business should be addressed to Ernest
L. Olson, Chairman of University Publications.

Reprints Schedule of The Great Basin Naturalist

Each
Additional

2 pp.

4 pp.

6 pp.

8 pp. 10 pp. 12 pp.

2 pp.

50

copies

$6.00

$7.00

$8.00

$9.00 $10.00 $11.00

$2.00

100

copies

7.00

8.00

9.00

10.00 11.00 12.00

200

copies

8.00

9.00

10.00

11.00 12.00 13.00

300

copies

9.00

10.00

11.00

12.00 13.00 14.00

Covers: $10.00 for first 100 copies, $4.00 for additional 100 copies.

TABLE OF CONTEI^TS

On Two Little-Known Species of the Earthworm Genus

Diplocardia. G. E. Gates 1

Spiders of the Nevada Test Site. Illustrated. Dorald M.

Allred and D Elden Beck _ 11

Problems of Sampling Desert Arthropods Before and After
a Thermonuclear Cratering Test. Illustrated. Clayton S.
Gist 26

New Species of Bark Beetles (Coleoptera: Scolytidae), Mostly

Mexican. Part VII. Illustrated. Stephen L. Wood 36

7

The

MUG. COMP. 20C:..
LIBRARY

MAR I 2 1Q69

HARVARD
UNIVERSITY

Volume XXVn, No. 2
September 5, 1967

Great Basin

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: "Vasco M. Tanner, Department of Zoology and Entomology
Brigham Young University, Provo, Utah

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:
J. V. Beck, Bacteriology

WiLMER W. Tanner, Zoology, Chairman of the Board
W. Derby Laws, Agronomy
Stanley Welsh, Botany
C. Lynn Haywakd, Zoology
Howard C. Stutz, Botany

Ex officio Members:
Rudger H.

CultUTf

Ernest L. Olson, Chairman, University Pubhcations

Rudger H. Walker, Dean, College of Biological and Agri-
cultural Sciences

^ \j- ^ K^ V LIBRARY

MAR 1 2 "089

The Great Basin NaturaIisI?Y

Published at Provo, Utah by
Brigham Young University

Volume XXVII September 5, 1967 No. 2

A COMPARATIVE STUDY OF THE MOUNTAIN BRUSH
VEGETATION IN UTAfP

Elray S. Nixon-

Abstract: a comparative study was made of an exclosure in
the mountain brush vegetation of the Wasatch Mountains, Utah.
The exclosure is located in Pole Canyon, near Provo, Utah, and was
initially analyzed in 1948 and 1949 by Allman (1952, 1953). The
dominant woody species of the vegetation in the exclosure were big-
toothed maple {Acer grandidentatum) and Gambel oak {Quercus
gambelii) . Results indicated that the maple had increased in relative
cover whereas the oak decreased Serviceberry {Amelanchier alni-
folia) had increased in both relative cover and frequency. Frequen-
cy data of forbs and grasses showed a general increase in early
flowering annuals and perennials. Starflower {Tellima parviflora)
and wyethia {Wyethia amplexicaulis) were two of the more promi-
nent species which were new in the exclosure. The complete disap-
pearance of western wheat grass (Agropyron smithii) from 67
percent frequency and the increase of Kentucky bluegrass {Poa
pratensis) represented the greatest changes of grasses in the exclo
sure. Precipitation was recorded from May 1. 1958, to April 30,
1959. Soil data obtained included pH, soluble salt content, cation
exchange capacities, mechanical analyses, and content of available
calcium, magnesium, and potassium. Soils were analyzed at six-
inch intervals to a depth of 24 inches.

Introduction

Allman (1952, 1953) presented a history of the study area and
indicated that during the years that followed the first settlements
in and around Provo, Utah, the area became heavily overgrazed by
livestock. It was not until the late 1930's that the study site was
included within the boundaries and management of the Uinta Na-
tional Forest. As a result, grazing by livestock was restricted until
1949, when the study area was fenced and analyzed by Allman

1. Appreciation is extended to Dr. Earl M. Chrislensen (Brigham Young University, Provo, Uuh)
for sugge<ition<i and guidance, the late Dr. Merrill J. Ilallaiu for aid in •soil analyses, and to Mark
GaiTelt for assistance with the field work.

1. Stophrn !•'. Austin Stair ('ollogr. Naropdorhos. 'I'evas.

59

The Great Basin Naturalist
60 ELRAY S. NIXON Vol. XXVII, No. 2

1952, 1953). The present study was proposed to determine any
changes occurring within the exclosure since the initial study and to
obtain certain environmental data which might supplement these
results.

The exclosure is located in Pole Canyon, which is situated in the
mountain brush vegetation to the northeast of Provo, Utah. It is
specifically located in Section 9 of Township 6 South, Range 3 East,
and is at an elevation of approximately 6,500 feet. The dominant
woody species of the area are big- toothed maple (yAcer grandidenta-
tum) and Gambel oak {Quercus gambelii). Of less prominence are
such species as serviceberry {Amelanchier alnifolia), ninebark
{Physocarpus malvaceus), and snowberry (Symphoricarpos vacci-
nioides). The study was begun during the spring of 1958, and it
continued through the summer of 1959.

Methods and Procedures

The exclosure consisted of 132 five-meter-square contiguous
quadrats arranged on a grid. These quadrats were permanently
marked and used to acquire frequency data for all species. Cover
and relative cover data of the woody vegetation were obtained by the
line-intercept method using a steel tape stretched along the grid lines
separating the quadrats.

Three sites were chosen just outside the exclosure for collecting
soil samples. At each site there were three collection points, one be-
neath an oak canopy, one beneath a maple canopy, and one beneath
a herbaceous cover. Samples were taken at depths of 6, 12, 18, and 24
inches. Analyses of duplicate samples included pH, a mechanical
analysis by the Bouyoucos hydrometer method, soluble salt content
with the use of a salt bridge, cation exchange capacity by the am-
monium acetate extraction method, and the amount of available
calcium, magnesium, and potassium present by use of a Beckman
flame-spectrophotometer. The moisture equivalents were determined
by the centrifuge method. The wilting point of the soil was found
by growing sunflower and wheat plants in glass tumblers in a
greenhouse.

A rain gauge was placed inside the exclosure and precipitation
measurements were taken from May 1, 1958, until April 39, 1959.
These measurements were made the day after the storm unless the
study area was inaccessible due to muddy roads. During the winter
months of January, February, and March the area was entered on
snowshoes; consequently, measurements were taken at the end of
each month.

Results
Vegetation

Results concerning the woody vegetational cover of the exclosure
indicated that the area could be divided into two vegetational types.
The west part was located on a ridge and contained high proportions
of both oak and maple (Table 1). The east part was located on a

Sept. 5. 1967 mountain brush vegetation 61

Table 1 . A comparison of the canopy cover of the east and west parts of the
exclosure in 1949 and 1958

West part (ridge) East part (slope)

% Relative % Cover % Relative % Cover
Species cover cover

1949 1958 1949 1958 1949 1958 1949 1958

Maple

40.7

50.9

33.8

44.6

74.6

71.6

60.6

58.9

Oak

55.9

43.8

46.4

38.4

22.1

21.8

17.9

17.9

Serviceberry

1.0

2.9

0.8

2.5

2.0

3.4

1.6

2.8

Ninebark

0.0

0.4

0.0

0.4

0.0

0.9

0.0

0.7

Snowberry

0.0

0.5

0.0

0.4

0.0

0.4

0.0

0.3

Rose

0.0

0.2

0.0

0.2

0.0

Trace

0.0

Trace

Douglas fir

2.4

1.3

2.0

1.1

0.0

0.0

0.0

0.0

WTiite fir

0.0

0.0

0.0

0.0

1.2

1.8

1.0

1.5

Juniper

0.0

0.0

0.0

0.0

0.1

0.0

0.1

0.0

Total

100.0

100.0

83.0

87.6

100.0

99.9

81.2

82.1

east-facing slope and the canopy was predominantly maple. An
analysis of the east part showed very little change during the nine-
3'ear period in regard to oak and maple cover (Table 1). The area
upon the rise, however, revealed some significant changes. There
was a definite increase of maple in the canopy and a decrease of oak.
Along with this, the total vegetative cover increased by 4.6 percent.
Young maple trees invaded and spread in the areas which were in-
habited mainly by oak in 1949. The oak. on the other hand, did
not invade the areas of older maple trees. There were few young
oak plants under the older maple in the exclosure.

Although maple, oak. rose {Rosa woodsii), and snowberry in-
creased somewhat in frequency, the only shrub showing any marked
change was serviceberry which had increased 14 percent (Table 2).
In addition, 50 seedlings of serviceberry were counted in 132 one-
meter-square quadrats located at random in the five-meter- square
quadrats. Although white fir (Abies concolor) showed no change in
frequency, there were six seedlings noted within the exclosure.

In general, an increase in frequency occurred in the majority of
forbs (Table 2). Spring beauty {Claytonia lanceolata), dog-tooth
violet {Erythronium grandiflorum) , blue-eyed Mary {Collinsia
parviflora). waterleaf {Hydrophyllum capitatum), nemophila (Ne-
mophila breviflora) , and starflower (Tellima parviflora) increased
throughout the exclosure. Plants which increased under the tall
maples in the east part of the exclosure were Engelmann aster (Aster
engelmanni). larkspur (Delphinium nelsonii). peavine (Lathyrus
spp.), and violet (Viola praemorsa) . Dandelion (Taraxacum offici-
nale) and mullein (Verbascum thapsus) decreased in the east part.
In the west part of the exclosure, which is more open, larkspur and
wyethia (Wyethia amplexicaulis) increased, while arrowleaf bal-
samroot (Balsamorhiza sagittata) geranium (Geranium fremontii).
goldenrod (Solidago altissima), and mullein decreased.

Species found in the exclosure in 1958 which were not recorded
in the preliminary study were starflower. western valerian (Valeri-
ana occidentalis) , wyethia, western wallflower (Erysimum asper-

62

ELRAY S. NIXON

The Great Basin Naturalist

Vol. XXVII, No. 2

Table 2. Comparative frequency data of species

% Frequency

% Frequency

Species

1949

1958

1949

1958

Abies concolor

11.4

11.4

Nemophila breviflora

62.1

84.8

Acer grandidentatum

98.5

100.0

Osmorhiza divaricata

84.8

93.2

Achillea lanulosa

34.8

31.1

Osmorhiza occidentalis

1.5

1.5

Agastache urticifolia

18.9

14.4

Physocarpus malvaceus

16.7

18.9

Agropyron smithii

67.4

0.0

Poa curia

15.2

12.1

Agropyron sub secundum 69.7

59.1

Poa pratensis

15.2

32.6

Amelanchier alnifoUa

46.2

60.6

Poa secunda

1.5

9.1

Aster engelmanni

32.6

37.1

Polemonium albiflorum

1.5

0.0

Balsamorhiza sagittata

5.3

0.0

Pseudotsuga taxifolia

3.0

3.0

Bromus carinatus

59.1

45.5

Quercus gambelii

89.4

95.5

Calochortus nuttallii

0.8

0.0

Rosa woodsii

2.3

5.3

Car ex spp.

90.9

91.7

Rudbeckia occidentalis

4.5

2.3

Cirsium undulatutn

0.8

0.8

Sambucus coerulea

0.8

0.8

Claytonia lanceolata

0.8

80.3

Scrophularia occidentalis 6.1

8.3

Collinsia parviflora

32.6

71.9

Smilacina spp.

93.2

86.4

Delphinium nelsonii

3.0

14.4

Solidago altissima

6.8

0.0

Elymus glaucus

32.6

50.8

Stellaria jamesiana

89.4

95.5

Erigeron speciosus

25.8

18.2

Stipa Columbiana

18.2

9.8

Erysimum asperum

0.0

0.8

Stipa lettermani

1.5

9.8

Erythronium

Symphoricarpos

grandiflorum

14.4

88.6

vaccinioides

6.1

9.9

Fragaria bracteata

11.4

14.4

Taraxacum officinale

34.9

25.8

Fritillaria atropurpurea

0.8

0.0

Tellima parviflora

0.0

39.4

Galium aparine

83.3

94.7

Thalictrum fendleri

0.8

1.5

Galium boreale

81.1

82.6

Valeriana occidentalis

0.0

2.3

Geranium fremontii

18.2

8.3

Verbascum thapsus

50.0

18.2

Helianthella uniflora

1.5

4.5

Vicia trifida

87.1

93.9

Hydrophyllum capitatum 22.7

90.2

Viguiera multiflora

31.1

28.8

Lathy r us spp.

78.8

99.3

Viola adunca

37.1

32.6

Mahonia repens

47.7

45.5

Viola praemorsa

14.4

23.5

Melica bulbosa

7.6

2.3

Woodsia sp.

0.8

0.8

Mertensia brevistyla

0.0

0.8

Wyethia amplexicaulis

0.0

5.3

Montia perfoliate

0.8

0.0

um), and shortstyle bluebell (Mertensia brevistyla) . Arrowleaf bal-
samroot, goldenrod, and white polemonium (Polemonium albi-
florum) were forbs recorded in the initial study which were not
found in the exclosure during 1958.

Of the grasses present, western wheatgrass (Agropyron smithii)
underwent the greatest change. It decreased from 67.4 percent fre-
quency to 0 percent. Other grasses which decreased were bearded
wheatgrass (Agropyron subsecundum) , Columbia needlegrass (Stipa
Columbiana), and mountain brome (Bromus carinatus). Kentucky
bluegrass (Poa pratensis), blue wild rye (Elymus glaucus), letter-
man needlegrass (Stipa lettermani). and Sandberg bluegrass (Poa
secunda) increased.

Precipitation and Soil

Rain and snow measurements recorded within the study area
showed a total of 16.18 inches of precipitation during the year with
about four-fifths of it being recorded from November to May (Fig. 1 ) .
Only 3.09 inches of rain fell from May 14 to November 7 and this

Sept. 5. 1967 mountain brush vegetation

Precipitation

2 -

63

u

^1

R

n.

May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr

Fig. 1 . Precipitation data obtained from within the exclosure.

was a result of 1 1 different storms. The highest amount of precipi-
tation. 2.4 inches, was recorded in February. The snow depth at
that time averaged about 30 inches. The smallest amount of precipi-
tation fell in October when .12 of an inch was recorded. July was
also very dry with only .23 of an inch of rainfall. Only one storm
occurred in each of these months.

The parent material of the soil of the study site was composed of
subordinate limestones and quartzites (Allman 1952). The A hori-
zon was fairly well developed but varied in depth extending in places
to more than 24 inches. On the ridge area in the exclosure it was as
shallow as 6 inches. The B horizon was not as variable and in some
places was poorly developed. The C horizon was as close to the sur-
face as 15 inches but in most cases was at a depth of 24 inches or
more. Rocks occurred throughout the profile.

The texture of the soil was a loam or silt loam and in every case
bordered the dividing line between these two classes (Table 3). Soil

Table 3. Average soil data under oak, maple, and herbaceous vegetative types.

Depth in

inches

0-6

6-12

12-18

18-24

Mechanical analysis

% Sand

34.7

36.5

37.9

38.2

% Silt

53.3

49.2

46.4

44.7

% Clay

11.9

14.3

15.7

17.1

Soil tyj)e

Silt Loam

Loam

Loam

Loam

pH

6.6

6.5

6.5

6.7

Soluble salts (ppm)

150.7

124.8

148.9

171.1

Cation exchange

(me/ too gms)

40.3

31.4

27.4

24.2

Ca (ppm)

768.0

631.0

574.0

547.0

Mg (ppm)

617.0

416.0

380.0

404.0

K (ppm)

379.0

172.0

103.0

71.0

Moisture equiv. (%)

31.0

25.0

24.5

22.4

Wilting point (%)

10.4

9.4

9.2

9.4

The Great Basin Naturalist
64 ELRAY S. NIXON Vol. XXVII, No. 2

pH ranged from 5.9 to 7.4 with an average of 6.6. Generally there
was little variation to a depth of 24 inches, and there was no signifi-
cant difference in pH of soils under maple, oak, or herbaceous cover.
Soluble salt content of the soils was comparatively low since, in most
instances, they contained less than 200 ppm. The concentration was
found to increase with depth with the exception of the first 6 inches
which usually contained higher amounts than the 12- and 18-inch
levels. Cation exchange capacities of the soils ranged from 22.5 to^
42.9 meq. per 100 g of soil. They always decreased with depth being
highest in the top 6 inches. The cation exchange capacity was lower
in the surface soil of the herbaceous sites than it was under oak and
maple where more organic matter was present. An increase in soil
depth was usually accompanied by a decrease in available calcium,
magnesimn, and potassium. Each of these elements appeared to be
relatively abundant with the possible exception of potassium at the
24-inch level (Table 3). The moisture equivalents and wilting points
of the soils showed little variation under the different canopy types.

Discussion

The mountain brush-type vegetation in the canyons and on the
mountain sides in the vicinity of the study area is, in most places,
dominated by Gambel oak {Quercus gambelii). Allman's observa-
tions of the exclosure made in 1948-49 led him to conclude that
maple {Acer grandidentatum) would eventually replace oak and
serviceberry {Amelanchier alni folia) in the lower ravines. Christen-
sen (1958) studied succession in reference to growth rates of maple
and oak in Pole Canyon and also concluded that maple was becoming
more significant in the composition of the vegetation. The results of
this study support these conclusions. On the ridge area where oak
was more prominent, young maples were spreading throughout the
oak. Maple freely invaded the oak brush, but it appeared that oak
had difficulty invading the maple.

The reasons maple increased at the expense of the oak are still
hypothetical. Christensen (1950) and Allman (1952) suggested that
competition with shade-tolerant species could cause the oak to de-
crease. Some support for this suggestion exists since there is a greater
abundance of forbs under the maple canopy than under the oak.
Allman (1953) found as many as 31 different species in one five-
meter-square quadrat under the maple. In this study there was an
average of 19 species per quadrat in the east half which was pre-
dominantly maple and an average of 15 species per quadrat in the
west half where more oaks were present. There was not only a
greater variety of species present under the maple, but also the den-
sity of these species was greater. Since oak brush in the exclosure
and in other areas (Moinat, 1956; Brown, 1958) has fewer herbs
beneath its canopy, maple stolons and seedlings could invade this
area more freely.

Both maple and oak reproduce vegetatively and by seeds. The
maple, however, produces many more seeds than the oak; and ap-

Sept. 5, 1967 mountain brush vegetation 65

parently these seeds are not as subjected to biotic influences as are
the oak acorns (Christensen, 1955). Seedling data has indicated that
maple is quite efficient in reproducing by seed, especially in new
areas, giving it a decided advantage over oak brush in reproductive
potential (Christensen and Nixon, 1964).

Among other woody species in the exclosure, serviceberry was
the only one which changed markedly. The fact that serviceberry
increased 14 per cent in frequency and that there were many seed-
lings and young plants present is evidence that it may continue to
increase in the future. Cottam and Evans (1945) reported a density
of .39 for serviceberry in an ungrazed canyon and a density of .19
in a grazed canyon (the term density, as used by Cottam and Evans,
represents percentage of plant cover obtained by the point-observa-
tion-plot method).

The exclusion of grazing has brought about various changes in
the forbs and grasses of the exclosure. Undesirable species such as
mullein (Verbascum thapsus) and dandelion {Taraxacum officinale)
decreased considerably in frequency. The increase of annuals and
small perennials denoted one of the interesting changes. The an-
nuals, nemophila {Nemophila breviflora). bedstraw {Galium apa-
rine), and the winter annual blue-eyed Mary {Collinsia parviflora) .
increased considerably in frequency. Small perennials such as spring
beauty {Claytonia lanceolata), dog-tooth violet {Erythronium. grandi-
florum.), and waterleaf {Hydrophyllum capitatum) also increased
markedly. McKell (1950) found that annual plants increased
greatly the first season after a fire in the mountain brush vegetation
but that perennials were slower in their recovery. The increase of
larger perennials such as Engelmann aster {Aster Engelmanni),
sweet anise {Osmorhiza divaricata). and figwort {Scrophularia occi-
dentalis) has been slow. The large perennial forbs are generally
found among the more open-type vegetation such as the tall maples.
Therefore, their increase will probably accompany the increase in
maple.

Kentucky bluegrass {Poa pratensis) spread under the tall maple
and into the open areas. In Colorado. Moinat (1956) reported that
Kentucky bluegrass was the dominant plant under the oak. Cottam
and Evans (1945) found that the density of the palatable grasses,
mountain brome {Bromus carinatus) , Kentucky bluegrass. and Sand-
berg bluegrass {Poa secunda), was five times greater in an ungrazed
canyon than in a grazed canyon. Sandberg bluegrass increased in
this study, but mountain brome decreased in frequency. Ellison and
Houston (1958) found that production of Bromus carinatus in aspen
openings was greater than under aspen canopy. Fheir results also
showed an increase of production with increase in altitude. The
altitudinal location of the exclosure is probably near the lower limits
of the distribution of this grass. The decrease of western wheatgrass
{Agropyron smithii) was one of the greatest changes. Its high fre-
quency at the time of the preliminary study is evidence that it can
compete under grazing conditions. Moinat (1956) and Brown
(1958) noted that western wheatgrass was the dominant grass in the

The Great Basin Naturalist
66 ELRAY S. NIXON Vol. XXVII, No. 2

openings between the oak, but it produced very little under the oak.

The exclosure received a total precipitation of 16.18 inches, most
of which arrived during the winter months. The summer storms
were of light intensity and occurred at fairly long intervals. This
resulted in one of the driest summers in Utah history. Only 3.09
inches of rainfall fell from May 14 to November 7. Price and Evans
(1937) recorded 7.01 inches in Ephraim Canyon, and AUman
(1952) registered 7.79 inches of rain from May 14 to August 25.
Allman also found that precipitation measured in the exclosure dur-
ing a ten-month period amounted to 22.57 inches. This was one of
the wettest years in Utah history. Baker and Korstian (1931), Price
and Evans (1937), Lull and Ellison (1950), Moinat (1956), and
Brown (1958) recorder measurements between 15 and 20 inches of
precipitation for oak brush areas.

Soil results indicated that conditions were generally favorable
for plant growth. The findings were usually similar to those of
Allman (1953) and, in some instances, comparable to Baker and
Korstian's (1931) and to Price's (1938) results from mountain brush
areas. Although soil characteristics varied, there were no significant
differences under the various canopy types.

List of References

Allman, V. P. 1952. A preliminary study of the vegetation in an exclosure in
the chaparral of the Wasatch Mountains, Utah. M.S. thesis, Brigham Young
University.

. 1953. A preliminary study of the vegetation in an exclosure in the

chaparral of the Wasatch Mountains, Utah. Proc. Utah Acad. Sci., Arts,
and Letters, 30:63-73.

Baker, F. S. and C. F. Korstian. 1931. Suitability of brushlands in the inter-
mountain region for the growth of natural or planted western yellow pine
forest. U. S. Dept. Agr. Tech. Bull., 256:1-82.

Brown, H. E. 1958. Gamble oak in west central Colorado. Ecology, 39:317-327.

Christensen, E. M. 1950. Distributional observations of oak brush (Quercus
gambelii) in Utah. Proc. Utah Acad. Sci., Arts, and Letters, 27:22-25.

. 1955. Ecological notes on the mountain brush in Utah. Proc. Utah

Acad. Sci., Arts., and Letters, 32:107-111.

1958. Growth rates and vegetation change in the oak-maple brush in

lower Provo Canyon, Utah. Proc. Utah Acad. Sci., Arts, and Letters, 35:
167-168.

Christensen, E. M. and E. S. Nixon. 1964. Observations on reproduction of
bigtooth maple. Leafl. Western Bot., 10:97-99.

CoTTAM, W. P. AND F. R. EvANs. 1945. A comparative study of the vegetation
of grazed and ungrazed canyons of the Wasatch Range, Utah. Ecology,
26:171-181.

Ellison, L. and W. R. Houston. 1958. Production of herbaceous vegetation
in openings and under canopies of western aspen. Ecology, 39:337-345.

Lull, H. W. and L. Ellison. 1950. Precipitation in relation to altitude in
central Utah. Ecology, 31:479-484.

McKell, C. M. 1950. A study of succession in the oak brush {Quercus gam-
belii) zone after fire. M.S. thesis. University of Utah.

MoiNAT, A. D. 1956. Comparative yields of herbage from oak scrub and inter-
spersed grassland in Colorado. Ecology, 37:852-854.

Price, R. 1938. Artificial reseeding on oak brush range in central Utah. U. S.
Dept. Agr. Circ, 458:1-18.

Price, R. and R. B. Evans. 1937. Climate of the west front of the Wasatch
Plateau in central Utah. U. S. Monthly Weather Rev., 65:291-301.

PREDACEOUS-SCAVENGER ANTS IN UTAH

D Elden Beck>, Dorald M. Allred', and William J. Despain-

Introduction

During the last twenty years the Brigham Young University
Department of Zoology and Entomology has sponsored extensive
field surveys throughout Utah to collect parasitic arthropods. In
most instances this involved trapping the host. While in the traps
many small rodents were preyed upon by ants, especially during the
night. This report is a summary of data accumulated over the years
on the predaceous activities of these ants. Those which we have con-
sidered as predaceous-scavengers in the following pages are arranged
first in phylogenetic sequence, then alphabetically with dates of col-
lection, localities, numbers of specimens, and prey associates listed
by specific name (Table 1).

Our use of the term predaceous-scavenger refers to those ants
for which we have actual evidence of their eating on the body of a
live animal or one recently killed. It does not include ants in de-
fensive or protective action.

In this study rodents were most often collected with Museum
Special snap traps. Traps were set out and baited in early evening
and retrieved early the following morning. Occasionally a trapline
was checked during the night. When animals were found with ants
eating them, the ants were placed in a paper bag along with the prey.
Cotton soaked in chloroform was used to kill the ants which were
then placed in vials containing 70 percent ethyl alcohol, and a label
showing field number, locality, prey, date and collector was added.
Further details on all collections were recorded in a field book.

All ants were identified by Dr. A. C. Cole. University of Ten-
nessee, to whom we are grateful for this courtesy. During periods of
the natural history surveys involving parasitic arthropods, some re-
search projects were supported by the National Institutes of Health
(Contracts E-103, E-1273. and AI-01273-8). Gratitude is expressed
for this su{)port. In the main, however, the collections were accumu-
lated by field surveys supported by the Brigham Young University
Department of Zoology and Entomology. Students and colleagues too
numerous to mention have been associated with the field operations.
Their valuable services are greatly appreciated.

Literature Review

The only extensive studies of ants in Utah are by Rees and
Grundmann (1940) of the University of Utah. Cole (1942) of the
University of Tennessee, and Olsen (1934) of Colorado State Univer-

I. Staff iiienil)crs, Deparlnient of /oology and Entoniolf)gy, BiigliHiii ^'oung University.
J. GraHualp stndcnl. Oeparlmcnt of '/.oology and F.nlomology. Rrigliani Young University.

67

68

BECK, ALLRED, AND DESPAIN

The Great Basin Naturalist

Vol. XXVII, No. 2

«3

-Q

3

3

'£ 3

3

35

a

3

3

3 -w

-S

a

■^ "3

-S

T3 a

Q

</3 lO <3

-2

a -<i

a -3

3 5 H ^"3

g

"a

is's

S 3

<o"3

2 3

3

<^

"3

.o ,

3 o

., ., • ~ <J ..

a. <-i.

• ~ <j , ^ .

■-^ tj

3

emic
rridi

emit
arvu
anic

to."-"

'Sc

5'

■ ■ |1'

|"i' ' ^

:3'S'3
■5 3.S

3 C

S

=^S

^E

•gS

1 s

•^ ff

1i:b

feS .^ ^5

5S

c

60 C

mias qu
myscus

si

mias qu
m.yscus

.. ..

^3

ohomys
mias sp
myscus
gnathus
myscus

11

5 °

o

o

O O

a o

o o

a o

S o

C <3 0 0 0
^- •5 k, l> k.

(^.

k.

t. k.

■^^ (■*

a. >^

"^ i»*

a. 1^

a. 1^

a.

c^

a, ft.

3 ^

3 5^

Qc^

C 3 5^ ^ t;)

Ot«ja,a,a,

^<^

O)

O O 00 XO

^mmrovnvnvri'r-

^ <£) ro -^ (T> XT) Oi

CS^-H

a)-Hooiomoo<ocN

--cs

<N

■.-H T-

fO

■^ 0 -^ fT

■^

^ o .

be S

o !=; S

tlH

>, to

►^3

3 s

B

d "5 OJ

. co-o-r;

^ •--.

o

^ tOt^COt-. t^ts,OOoO'^^Ot-t>.»-~QOOOOOOOOOOO

(N CN) -r^ 55 -r^ CN (N -rH ^ -^ 01 (>0 ^^^^

u^ vT) >ri ^ vn
d 00 ■^ s tv. 05 ;
t^ 00 N. t^ to

«

s

<

a

i4-M

<»

0

>- S

5 e

^

.2P t'

0)

a ^

Q<

0 3

-/D

<o

</5

Q. ».

0 <»

0 s>

-s §^

§-^

0 Cl.

^ ^

S<^

c

OS

n.

3

Sept. 5, 1967

PREDACEOUS-SCAVENGER ANTS

69

(^ g

11

S c

o o

3-~

)5 c

^ 3 <.

c a ^

coo
a (a 1^)

3

3

3 3 3

--i

3

*- &\ »j

v! .-

Q

N «

^ ^

^ tJC'

-2

3 -->

3 ^

^

l.

S ^ a

J £

"3

^

JJ

■- ^2 2

85 ^ c:~

S <o

•~. c'

o

■3 fe.ii §

^1

o

^
2

0, 3 <o §

^yis^

3l

!j O

x§

^

O C O i"'

i^iiy^

ll^

iE-'

O o o O

Q. k k p 1^ 6.

11

1i'

1

n. k. o. k.

•^ T*

Q. i^

o

• — ^a ■-• ^

<^ ^

•~ &)

<;

Qa-Qa,

Qa^a.T^a.Q

Q, a;

Qti!

hi

oovnoQOxnvn'N m^o vnvn^rimvnoioo'oooo-^'^ooo-^fovfivn oi
•^ csi csi csi -^ •.-I ro CM xn •*■ <n (N <N <o \n -ri (T) 1-

§1)

Xl

5i> S

"13
2>

— - ,<rz2 s-

^ CO ^ -

,S to

Wig

O
01

(0

©2

« g (J * u5

^ g o"^ ■55.S ?
<! u Qh ffi ffi t=<i O cc

^2 3 >^^^'o t, 3 a, -h;

3 !^ C

00 rO 05 O vn Cs) (

vn in tJ- to vn in <
C> t>. 00 00 XI s t^ 00 ■* (O ^D <r> t^ t^ t^ t^ 00 00 00 00 00 00 00 :

o.cu'^

O) (£) -"J- <0 00 00 CT>

a

c

a

■p' S'

13
5.= .2

o

The Great Basin Naturalist
70 BECK, ALLRED, AND DESPAIN Vol. XXVII, No. 2

acsa a 3sa!2<o'o 3«'«3 gi- a

.o fc-t:_(j <o.<j- , , , . .vj, -^.ij 333 .0, ^^.«j - '^ c .'J

8'3?^38'^5 2,3 ''Sjp^^'S 32,3 5=^.52.3?

os|S5-|SS' ''''-§S' SS^2S5|S' SiS'^SS-iH-l

^

"3 sH »i « i ?o

C/D

TTJ

r<

^

V

S

ij

"ffl

W

(D!/5

JT

C «

0

*j 0)

0) 5 2

C O 3

h-1 2 CC 2 fc.S| « ^CQS P^QhO^I^O^ CL, hJ Oh !^ Oh td tf

fTit^ Oi-^ •rtOO«->O^^OOOOC^^c<^■<J■■^-"^^oo■^■^^^^^v^■^oooOfr)oooo■^C>^>•

a

>>.

a

p

1^ ~

£'

3

3-S

Sept. 5, 1967

PREDACEOUS-SCAVENGER ANTS

71

(0

I

3 ;2

«^

S 3

•2 X,

« o

3fe

3

3 3

<3

a a

3

<o 3 <o 3

3.ii 3.ii
C C £ C

a <3 2 a
c 5 c S

o o

•l'^

S3SS

<^ (J

o , (J

a <^ 2 "^

3 X 3 X

P £ ? 5

O O

h o h o

a. S

• ii "--hi >-

Q^

5^^c^

« a c
3 3 c: 3

II

<o 3

^ bo

fc c

£-

5 5

<-5 ^

s|s«

« £ e

r» k. a

: So ci

S^ «*
T? ^ o

^ • ~ ^^

2:QS

l3-i

3 ^g

? O *

-Si^ £
-« £'

(S «^ vr

1

fth'
C .

' " 5 >

'■ So

o

c

0)
bo rj

bo-g

.5 Q S

c c

-i« u

S so

J> — . O to c
a; is tfi to

-_uco.i:(jto(o3

5^ CO

s- '«:

S =

c
S «

CO 0- ,

to c t_,

L> (0

CO -r lU '

PQ

:=: 3

t- o

0) CO

air E 2

■C-S"!;^ CO o*-

^^^>.<»ex> oo tot^cx) i^t^(3o!Ot^t^t^«^oo ^;o^<0>^^-f^oooooo°o;

f^t^CSlCM ^- a; — f<^ t^t^cNCSl-.— t^CX) — CM t^t^c»OOvDt>-OC<)^00

.5= s o

o o

72

BECK, ALLRED, AND DESPAIN

The Great Basin Naturalist

Vol. XXVII, No. 2

CO

3

1

3|

1

5)

W 'ii

§•3

^3

S c

O bo

p

4

n s^

a o

? "^

~ 1^

^ "il

3 (w

:e:a.

fcqo.

a "fa

(*♦ S fs

•■5 •« e

S 3 K "^ ?3 i^ >«-^

<o-Si '^ 3

s § s ^

a ® D t;

"3

S 3

5 ■

= ^ r1

?-ii^

?.^ 3
O -c

3

^5

3 c
■Si 5

■^

^•^

X

^o >^

fc c

3

5 e
c

^ ^"i: ^

bo S ^ fc 2 O bo^ bo S

3 ^^

5. is §-S

o s o 2 o
S « S ^ S

.6. ■g.O. ^.6.

■ 0>ooo^o■^T^o mdvrQooooif)0<r)vnvn-^0'^ ovo c^vn

<£> 0000 -o(£>t>.t^t^'^xr>^^toio^t^ t^t^t^oor

"-V ^~,\ ^\ ~\~\ -\^^^^-\ ^>^ \^\^.

00 00t£) t^OltNiO-^t^fOt^OOOO^'^CSl ro^OOOf^

Tj- csi 00 in to 00 «^

— CN CM

■^ rrs^D XT) 00 O) C

C «;; ::

s

5

«B X

!>.

T3-C

X

.<o

S'

'i

0
c

0

•"'-'"

s.

4i

0

0

a.

Co

Sept. 5. 1967 predaceous-scavenger ants 73

a 5 5 3

"a ^"3 "3 o, "3 ,, 2

:;s 0 c .fee org o a>s o.c e

•g^l 1^, |e>^, 11 ^ll

S £ o 2 £ o2 £ .b |£2 £

§ ¥

c

bCc E M £

>:ia:^ 5 CO C ^lO.S .„.

0? p_ > CO to c 1- 5P^ — „ >

= J2 .'^'^^ CO & ,^C> t.^ !^

s o^ , CO > ^ o'^c/:'^^ ^
V o .2,'^ i--2 5^ CO c V- .?>■-- ^-

J2

S

O > O-c

Oh Q Z cc cQ « H: cc hj (1, w hJ eu W ^5 h

— <T. (N — — C>4 — ^^^

65.

The Great Basin Naturalist
74 BECK, ALLRED, AND DESPAIN Vol. XXVII. No. 2

sity. Although these studies include a large listing of ant species for
the state, little is mentioned about their feeding habits.

Creighton's work (1950) on the ants of North America makes
general references to food habits for some spedies. and in a few in-
stances gives specific reference to others. Several direct references
involve species that we have observed, whereas others relate to spe-
cies not known from Utah. Some of Creighton's data related to
scavenger-predaceous species are quoted below, followed by our
comments.

"Platythyrea punctata (F. Smith): The workers are active and
forage singly. The colonies are small consisting of from fifty to two hun-
dred individuals. It is both carnivorous and predatory" (p. 34).

This species occurs in the extreme southern part of the United
States.

"Cerapachys augustae Wheeler: It is virtually certain that these ants
are carnivorous, and it is probable that they are predaceous" (p. 58).

The range of this species is from western Texas to southern
Arizona.

"At certain seasons these insects [ants of subfamily Dorylinae] be-
come nomadic, and the entire colony sets out on an expedition which
becomes a series of raids against animals that may happen to be in
the vicinity . . . although there has been much exaggeration of the
capacity of these insects for attacking large vertebrates. Undoubtedly,
they would do so if given the opportunity, but unless the animal was
badly crippled or comatose, it could easily avoid the attack. The main
victims of these raids are other insects which are secured in prodigious
numbers" (p. 60).

"There is a persistent belief that in the days when the West was
wilder than it is now, Indians would sometimes stake out a human victim
across a nest of Pogonomyrmex. If this was actually done it would be
hard to imagine a more excruciating death" (p. 110).

We observed Pogonomyrmex occidentalis demonstrating the scav-
enger-predaceous habit in only two instances, yet it is one of the
most widely distributed ants in Utah. It has a ferocious habit of
attacking and stinging a victim as a protective action. The sting is
painful to humans.

"Despite their preference for a graminivorous diet, many species
of Pheidole will accept other food as well. They seem less attracted to
honey-dew than do many ants but will often feed voraciously on animal
tissue when the opportunity offers" (p. 161).

We have records of four species of Pheidole being scavenger-
predaceous in habit. They are P. ceres, P. bicarinata, P. dentata and
P. hyatti.

"Because of their omnivorous habits, they [Solenopsis geminata and
5. saevissima'] are always turning up in unexpected situations. They
have been known to damage the buds and tender twigs of young fruit
trees and kill quail which are too young to leave the nest" (p. 227).

We observed Solenopsis molesta validiuscula as a scavenger-
predator. These ants are the popularly known Fire Ants, a name

Sept. 5, 1967 predaceous-scavenger ants 75

given to them because of their painful sting. We included this
reference because Creighton mentions the term omnivorous; to kill
does not mean the ant is a predator or scavenger.

"The ants {Dorymyrmex pyramicus and D. bicolor] are very active
and predaceous but will feed on honey-dew when thy can get it. They
have a strong odor of butyric acid which is particularly noticeable when
they are crushed" (p. 348).

Dorymyrmex pyramicus and D. bicolor definitely are predaceous-
scavengers.

"Of Myrmecocystus ... a considerable proportion of the species
. . . api>ear to be c£imivorous" (p. 354).

We found this to be true for M. mexicanus hortideorum, M.
pyramicus and M. mojave.

Results

The taxonomic arrangement of subfamilies and genera follows
that of Creighton (1950). In a few instances in the list below, only
generic determination was possible.

Subfamily Myrmicinae

Myrmica brevinodis discontinua Weber Pheidole hyatti Emery

Myrmica lobicornis fracticornis Emer>- Crematogaster depilis Wheeler

Pogonomyrmex occidentalis (Cresson) Crematogaster lineolata emery ana
Aphenogaster subterranea valida Creighton

Wheeler Crematogaster punctulata Emery

Aphenogaster subterranea occidentalis Crematogaster minutissimai Mayr

(Emery) Crematogaster mormonum Em.ery

Pheidole sp. Monomorium minimum (Buckley)

Pheidole ceres Wheeler Solenopsis molesta validiuscula Emery

Pheidole bicarinata Mayr Leptothora muscorum (Nylander)
Pheidole dentata Mayr

Subfamily-Dolichoderinae

Iridomyrmex pruinosum analis Dorymyrmex pyramicus (Roger)

(E. Andre) Tapinoma sessile (Say)

Dorymyrmex bicolor (Wheeler)

Subfamily Formicinae

Camponotus sp. Myrmecocystus mojave Wheeler

Camponotus herculeanus modoc Myrmecocystus pyramicus Smith

Wheeler Formica sp.

Camponotus sansabeanus torrefactus Formica cinerea lepida Wheeler

Wiieeler Formica criniventris Wheeler

Camponotus vicinus Mayr Formica fusca Linne

Paratrechina sp. Formica integra haemorrhoidalis
Lasius sp. Emery

Lasius alienus Mayr Formica limata Wheeler

Lasius crypticus Wilson Formica neoclara Emery

Lasius niger Mayr Formica neorufibarbis gelida Wheeler

Lasius sitkaensis Pergande Formica perpilosa Wheeler

Acanthomyops claviger (Roger) Formica pruinosa Wheeler
Myrmecocystus mexicanus hortideorum Formica obscuripes Forel

McCook

76

BECK, ALLRED, AND DESPAIN

The Great Basin. Naturalist

Vol. XXVII, No. 2

Table 1 lists the species collected, dates of collection, specific
localities (towns or other geographic locations), counties, numbers of
specimens collected, and animals upon which the ants were feeding.
Dates of collections are arranged by day, month and year. When a
species was collected several times during the year, the dates are
listed in chronological order.

Six species were found only in the Great Basin, fourteen in the
Colorado River Drainage Basin, and twenty-four species were gen-
erally distributed in both basins. See Table 2.

For the most part, small rodents were the animals upon which
the ants were observed feeding. In a few instances rabbits were in-
volved. Occasionally small ground-dwelling birds were caught and
killed in snap traps, and ants preyed upon them. In other cases ants
invaded the nests of rodents and attacked their young.

TABLE 2
Geographic Distribution

Great Basin Only

Colorado River Basin
Only

Both Basins

Acanthomyops claviger
Aphenogaster subterranea

occidentalis
Crematogaster Uneolata

emeryana
Crematogaster mormonum
Formica pruinosa
Pheidole ceres

Camponotus sansabeanus

torrefactus
Crematogaster depilis
Crematogaster punctulaia
Crematogaster

minutissima
Formica criniventris
Formica neoclara
Formica perpilosa
Iridomyrmex pruinosum

analis
Lasius alienus
Myrmecocystus

pyramicus
Paratrechina sp.
Pheidole dentata
Pheidole hyatti
Pogonomyrmex

occidentalis

Aphenogaster subterranea

valida
Camponotus vicinus
Camponotus herculeanus

modoc
Dorymyrmex bicolor
Dorymyrmex pyramicus
Formica cinerea lepida
Formica fusca
Formica Integra

haemorrhoidalis
Formica limata
Formica neorufibarbis

gelida
Formica obscuripes
Lasius crypticus
Lasius niger
Lasius sitkaensis
Leptothorax museorum
Monomorium minimum
Myrmica brevinodus

discontinua
Myrmica lobicornis

fracticornis
Myrmecocystus mexicanus

hortideorum
Myrmecocystus mojave
Pheidole sp.
Pheidole bicarinata
Solenopsis molesta

validiuscula
Tapinoma sessile

Sept. 5. 1967 predaceous-scavenger ants 11

Discussion

In the several studies of ants of Utah the schemes of classification
have varied. In so far as we can determine from the literature, ap-
proximately 126 kinds of ants combined under species, subspecies,
and a variety of other categories are known for Utah. We list 42
kinds representing 41 species in 17 genera. The genus Paratrechina
was the only one for which specific identification could not be made.

The followdng 23 species and subspecies and one genus are herein
reported from Utah for the first time: Acanthomyops claviger,
Aphaenogaster subterranea valida, Camponotus vicinus, Cremata-
gaster depilis, C. lineolata emeryana, C. punctulata, C. minutissima,
Formica cinerea lepida, F. Integra haemorrhoidalis, F. limata, F.
neorufibarbis gelida, F. neoclara, Lasius alienus, L. crypticus, Lepto-
thorax muscorum, Myrmica brevinodis discontinua, Myrmecocystus
pyramicus, M. mojave, Paratrechina sp., Pheidole ceres, P. bicari-
nata, P. dentata, and P. hyatti. It is unusual to have more than half
of our collections represent new distribution records.

In many years of field surveys, and especially those involved with
parasitic arthropod investigations, we have sampled most of the
major types of ecological situations which occur in Utah. This may
account in part for the many new distributional records.

Of the approximate 1 26 kinds of ants previously reported, 1 9 have
been found by us to be predaceous-scavengers. This indicates that
the 107 other kinds do not have this habit, or we have failed to dis-
cover such activities for these species. Although the latter is possible,
it seems unlikely when one considers the number of years involved
in our surveys and the thousands of animals trapped in varying
types of habitats.

One should not classify an ant as a predaceous-scavenger kind if
the ant simply assumes a defensive or protective action. Such a de-
fensive pose is taken when Pogonomyrmex occidentalis is disturbed.
One of the most abundant ants in Utah, this insect is responsible for
mounds scattered throughout the valleys and foothills. Yet, our rec-
ords show only two instances where this species was observed con-
suming animal flesh.

Those ants which we consider as predaceous-scavengers and are
widespread in Utah are Camponotus vicinus, Dorymyrmex pyrami-
cus, Lasius niger, Myrmecocystus mexicanus hortideorum, Pheidole
bicarinata. and Tapinoma sessile. Some forms, such as Iridomyrmex
pruinosum analis which was encountered only in the southeastern
part of Utah in lowland desert situations, could be considered geo-
graphically restricted. Creighton (1950:343) stated that "the north-
ern limit of the range appears to lie in southern Idaho." Although
not restricted to any part of Utah. Lasius sitkaensis occurs at higher
elevations on mountains, in canyons and in valleys.

There is little evidence that any of the ants observed in this study
are prey-specific in their association. We have trapped a number of
species of rodents in high mountain situations many times over the

The Great Basin Naturalist
78 BECK, ALLRED, AND DESPAIN Vol. XXVII, No. 2

years. At these higher elevations the numbers of species of preda-
coous-scavcnger an.s are comparatively fewer than at lower eleva-
tions and in the desert

Geographic distributional records were included only for our col-
lections. Seasonal collecting on a year-round schedule in localities
ecologically similar and at similar altitudes is desirable. Collection
data certainly are not complete, for example, when records for
Pogonomyrmex occidentalis are known only from two localities at
opposite ends of the state. The same is true for other species such as
Formica neorufibarbis gelida which shows only an extreme east and
west distribution.

Bibliography

Cole, A. C., Jr. 1942. The Ants of Utah. Amer. Midland Nat., 28(2): 358-388.
Creighton. W. S. 1950. The Ants of North America. Bull. Mus. Comp. Zool.

Harvard College, v. 104.
Olsen, W. O. 1934. Notes on the North American Harvesting Ants of the

Genus Pogonomyrmex Mayr. Bull. Mus. Comp. Zool., Harvard College, v.

78(8).
Rees, D. M. and A. W. Grundmann. 1940. A Preliminary List of the Ants of

Utah. Bull. Univ. Utah, Biol. Ser., 6(2).

NEW RECORDS AND SPECIES OF NEOTROPICAL
BARK BEETLES (SCOLYTIDAE: COLEOPTERA)'

Stephen L. Wood-
While preparing a taxonomic review of the bark and ambrosia
beetles (Scolytidae) of Costa Rica, it was necessary to examine all
available species of this group from Mexico, all of Central America,
and northern South America. As a result, a large number of species
new to science were recognized from areas outside of Costa Rica. In
order to make names available for these species to facilitate this and
other work. 20 species new to science are described below. The new
species represent the genera Cnesinus (11). Leperisinus ( 1 ) , Chra-
mesus {5) . Phloeotribus (I), and Chaetophloeus (2). The type series
of these species were collected in the following countries: Mexico
(7). Guatemala (4). Honduras (3), Panama (4), and Colombia (2).

Cnesinus elegantis, n. sp.

This species is very closely related to elegans Blandford, but is
easily distinguished by the larger size, by the more strongly strigose
pronotum, by the more coarsely, deeply punctured declivital inter-
spaces, and by the more distinctly striate elytral declivity.

Female. — Length 3.3 mm. (paratypes 3.0-3.4 mm.). 2.7 times as
long as wide; color brown.

Frons strongly, transversely impressed at level of antennal bases,
convex above, impressed area with a pair of sublateral calluses on
either side of deepest point of impression, epistomal callus evident
and armed by a pair of rather widely placed, large, pointed tubercles
larger than those of elegans; surface reticulate and minutely, sparsely
punctured; vestiture restricted to sides below level of antennal bases,
and epistomal area, fine, hairlike; eyes separated by a distance equal
to twice the greatest width of an eye.

Pronotum 1.06 times as long as wide; sides widest in front of
middle; surface with closely set, deep, elongate punctures, more than
half longitudinally confluent; glabrous.

Elytra 1.8 times as long as wide, 1. 9 times as long as pronotum;
sides straight and subparallel to declivital base, rather broadly round-
ed behind; striae moderately impressed, the punctures rather small,
deep, separated by distances equal to their own diameters; interstriae
twice as wide as striae, the punctures variable in size, rather con-
fused. Declivity rather steep, impressed between third interspaces;
striae not impressed, the punctures clearly, rather deeply impressed;
interspace 1 moderately elevated. 2 flat and wider than 1 or 3; all

1. Part of the field work relating to this study was sponsored by n reseanh grant from the
Nationel Science Foundation, No. GB-532.

1. Department of Toologv and Entomology. BriKhaiii Young I'nivprsily. Provo, Utah. Scolytoidea
(Contribution No. iV

79

The Great Basin Naturalist
80 STEPHEN L. WOOD Vol. XXVII, No. 2

interspaces armed by a few fine, rounded, irregularly placed gran-
ules. Vestiture largely confined to declivity, consisting of long,
rather fine interstrial bristles, and shorter strial and interstrial hair;
the longest bristles equal in length to combined widths of interspaces
2 and 3.

Male. — Similar to female except frontal callus slightly more
prominent and unarmed.

IVpe Locality. — Volcan Zunil, Quezaltenango Prov.. Guatemala.

Host. — Quercus sp.

Type Material. — The female holotype, male allotype, and 31
paratypes were collected from oak twigs at the type locality on May
27, 1964, at an elevation of 1,000 m.. by S. L. Wood.

The holotype. allotype, and paratypes are in my collection; one
paratype is in the Schedl collection.

Cnesinus hicornus, n. sp.

This species is closely related to elegans Blandford, but is readily
distinguished by the much larger size, by the stouter body, by the
broadly concave frons in both sexes, by the presence of interstrial
granules on the elytral disc, and, in the female, by the much larger
epistomal tubercles.

Female. — Length 3.5 mm. (paratypes 3.3-3.8 mm.). 2.5 times as
long as wide; color dark reddish -brown.

Frons rather deeply concave from eye to eye from vertex to
epistoma, lateral margins acute below eye; epistoma armed by a
pair of very large, pointed, conical spines, each longer than half of
greatest width of eye; concavity on lower half with a pair of lateral
calluses; surface substrigose-reticulate, with small, rather sparse,
obscurely granulate punctures; vestiture fine, obscure, except a row
of long and erect hairlike bristles on lateral margin; eyes separated
by 2.7 times greatest width of an eye.

Pronotum 1.1 times as long as wide; sides almost straight on
basal tvvo-thirds, very slightly wider in front of middle, broadly
rounded in front; surface longitudinally strigose, the punctures
usually not evident, grooves dull, ridges shining; median line evident
on posterior half; vestiture hairlike, inconspicuous except on all
marginal areas.

Elytra 1.6 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and parallel to declivital base, rather broadly
rounded behind; striae narrowly, not strongly impressed, the punc-
tures small, impressed; interstriae three or more times as wide as
striae, subcrenulate toward base, the remaining surface shining, not
clearly punctured, irregular, with small rounded granules in some-
what indefinite uniseriate rows. Declivity rather steep, convex,
except flattened between third interspaces; striae 1 strongly. 2 mod-
erately, 3 not impressed; interspaces subshining, weakly convex,
about four or five or more times wider than strial punctures, each
interspace with a median row of rather large rounded granules.
Vestiture hairlike, more abundant but not longer on declivity; con-

Sept. 5. 1967 neotropical bark beetles 81

sisting of median rows of fine, erect interstrial hair, and more abun-
dant, almost equally long, semirecumbent hair; the longest setae
about equal to distance between rows.

Male. — Similar to female except frontal excavation slightly
shallower and less extensive; epistomal callus developed but
unarmed.

Type Locality. — Thirteen miles west of Morelia, Michoacan.
Mexico.

Type Material. — The female holotype, male allotype, and 43
para types were collected at the type locality on June 15, 1965, at an
elevation of about 2,300 m., by S. L. Wood, from small stems of a
large herb 2-3 m. tall.

The holotype. allotype, and paratypes are in my collection.

Cnesinus foratus, n. sp.

Closely allied to electus Wood, but distinguished by the much
more closely, coarsely, substrigose punctures of the pronotum, by the
steeper, much less strongly impressed elytral declivity, and by the
narrower male epistomal callus.

Male. — Length 2.0 mm. (paratypes 2.0-2.1 mm,), 2.4 times as
long as wide; color dark brown.

Frons transversely impressed at level of occular emargination.
convex above, flattened below this point; upper surface faintly, trans-
versely strigose. shining, with sides reticulate (entire area reticulate
in one paratype) ; lower area flattened, with a low, broad, triangular
epistomal callus glabrous and shining below, a band of closely set.
erect, reddish hair on upper slope, remaining area reticulate; gla-
brous except for epistomal brush and a single, sparse row of hair
along lateral margin to upper level of eyes; eyes separated by a dis-
tance equal to about twice the greatest width of an eye.

Pronotum 1.04 times as long as wide; sides weakly arcuate on
posterior half, widest at about middle, anterior margin broadly
rounded; surface closely, coarsely, shallowly punctured, the punc-
tures longitudinally confluent, the interspacial ridges occupying less
than a third of surface; vestiture consisting of long, coarse bristles
in marginal and median areas.

Elytra 1.6 times as long as wide. 1.7 times as long as pronotum;
sides almost straight and parallel to declivital base, broadly rounded
behind; striae impressed, the punctures partly obscured by conflu-
ence; interstriae somewhat wider than striae, surface irregular, the
punctures rather small, somewhat uniseriate, subcrenulate toward
base. Declivity steep, convex; striae 1 impressed, others indicated,
the punctures reduced but distinct; interspaces as wide as striae, 1
slightly elevated and subgranulate. 2 and 3 and others with a row of
moderately large, rounded granules. Vestiture consisting of long,
stout (subscalelike). interstrial bristles from base to apex, distance
between bristles and between rows slightly less than length of a
bristle.

The Great Basin Naturalist
82 STEPHEN L. WOOD Vol. XXVII, No. 2

Female. — Not represented in the material at hand.

Type Locality. — El Bosque, Caicedonia, Valles, Colombia.

Type Material. — The male holotype and four male paratypes
were collected at the type locality on June 23, 1959, by J. Restrepo,
from dead branches. Two other male paratypes are from Finca la
Tribuna, at the same locality, taken in May, 1959. by Gregorio
Vargus from coffee berries.

The holotype and paratypes are in my collection.

Cnesinus electinus, n. sp.

Very closely related to electus Wood but eyes much more widely
separated and epistomal callus and brush more elaborately de-
veloped.

Male. — Length 2.4 mm. (paratypes 2.1-2.5 mm.), 2.7 times as
long as wide; color dark brown, the elytra somewhat lighter.

Frons transversely impressed at level of eye emargination, flat-
tened below, and convex above that point; epistomal callus broad,
low, lower margin abrupt, precipitous, small triangular median area
impunctate and glabrous, remaining area densely covered by a plush
stand of erect, reddish bristles; convex area almost smooth, subshin-
ing, impunctate except at sides and above; vestiture scanty, limited
to lateral areas except for epistomal brush; eyes separated by a dis-
tance slightly more than twice the greatest width of an eye.

Pronotum 1.1 times as long as wide; sides almost straight and
parallel on anterior two-thirds, very slightly wider in front of middle,
anterior margin broadly rounded; surface smooth and shining, with
rather numerous, minute points and moderately large, deep, oval
punctures, the punctures separated by about one-half to three times
their own diameters, none confluent; vestiture sparse, inconspicuous,
confined to marginal areas.

Elytra 1.7 times as long as wide, 1.7 times as long as pronotum;
striae impressed, the punctures moderately large, deep; interstriae
about twice as wide as striae, convex, the punctures small and stag-
gered in indefinite, uniseriate rows, the punctures granulate on 2 and
3 near declivity and on 1 behind middle of disc. Declivity steep, im-
pressed between third interspaces; striae 1 strongly impressed, others
not impressed, the punctures moderately large and deep; interspace
1 somewhat elevated, 3 convex, 1 with fine punctures, 2 and 3 with
moderately large, rounded granules. Vestiture almost restricted to
declivity, consisting of uniseriate rows of erect bristles on all inter-
spaces except 1, each bristle one to two times as long as distance be-
tween rows of bristles.

Female. — Similar to male except epistomal callus and brush
more poorly developed.

Type Locality. — Fifteen miles south of Mazamitla. Jalisco,
Mexico.

Type Material. — The male holotype, female allotype, and 28
paratypes were collected at the type locality on June 22. 1965. at an

Sept. 5. 1967 neotropical bark beetles 83

elevation of about 2,400 m., by S. L. Wood, from twigs of a small
tree.

The holotyi)e. allotype, and paratypes are in my collection.

Cnesinus niger, n. sp.

This species is rather closely allied to panamensis Blackman, but
is readily distinguished by the dorsad extension of the frontal im-
pression, by the deeper, nonconfluent strial punctures on the elytral
disc, by the more finely sculptured elytral interspaces, and by the
much smaller punctures of all declivital striae.

Male. — Length 2.0 mm. (paratypes 2.0-2.5 mm.), 2.6 times as
long as wide; color almost black.

Frons convex above, transversely impressed at level of occular
emargination, lower half of area between impression and epistoma
bearing an elevated callus, the callus almost reaching lateral margins,
sharply defined above, densely pubescent on its upper slope, with a
small, median, glabrous, triangular area below; surface subreticulate,
shining, punctures not evident below upper level of eyes; vestiture
limited to callus and a few inconspicuous setae along sides. Eyes
separated above by a distance equal to about two and one-half times
the greatest width of an eye.

Pronotum 1.04 times as long as wdde; widest just behind middle;
surface almost smooth and shining, the punctures small, rather close,
almost round at base, each puncture becoming elongate anteriorly
but not more than twice as long as wide, rarely confluent; glabrous
behind, a few short setae on anterior third.

Elytra 1.7 times as long as wide, twice as long as pronotum; sides
straight and subparallel on more than basal half, rather narrowly
rounded behind; striae impressed, the punctures large, deep, clearly
separated from one another; interstriae as wide as striae, the punc-
tures fine, shallow, not strongly confused, surface subshining. De-
clivity rather steep, convex; interspace 1 distinctly elevated, median
half of 2 flat, impressed (less obvious on left side), lateral half nar-
rowly convex, 3 narrowly convex, 2 and 3 each bearing a row of
fine granules. Vestiture confined to declivity, consisting of inter-
strial rows of moderately long, stout bristles, except missing on lower
two-thirds of 1.

Female. — Similar to male except frontal callus and brush greatly
reduced.

Type Locality. — Cerro Punta near Volcan Baru (or Chiriqui),
Panama.

Type Material. — The male holotype. female allotype, and 41
paratypes were taken at the type locality on January 11, 1964. at an
elevation of 1,800 m.. by S. L. Wood, from the twigs of three dif-
ferent tree species.

The holotype, allotype, and most of the paratypes are in my col-
lection; one para type is in the Schedl collection.

The Great Basin Naturalist
84 STEPHEN L. WOOD Vol. XXVII, No. 2

Cnesinus myelitis, n. sp.

This species is allied to punctntus Blandford, but may be distin-
guished by the smaller size, by the more slender form, by the larger,
more shallow punctures of the pronotum. and by frontal and elytral
characters.

Male. — Length 2.4 mm. (para types 2.0-2.4 mm.). 2.4 times as
long as wide; color dark brown.

Frons transversely impressed at level of occular emargination,
convex above and flattened below this point; upper surface minutely,
transversely aciculate to upper level of eyes, lower area subreticu-
late and with a large, broad epistomal callus covered by a brush of
rather long, erect, reddish bristles, the entire callus setiferous, smaller
and less strongly elevated than in punctatus.

Pronotum 1.05 times as long as wide; sides feebly arcuate on
basal two-thirds, very slightly wider in front of middle, anterior
margin rather broadly rounded; surface subshining, evidently with
very minute points and moderately large, shallow, oval punctures,
the punctures largely confluent on anterior half, rather widely
separated on posterior half; subglabrous.

Elytra 1.5 times as long as wide, 1.6 times as long as pronotum;
sides straight and parallel on anterior two-thirds, rather broadly
rounded behind; striae rather strongly impressed, the punctures
moderately large, deep; interstriae about twice as wade as striae,
convex, the punctures uniseriate, fine in middle area, subcrenulate
toward base, forming rather large rounded granules on posterior half
of 1 and posterior fourth of 2 and 3. Declivity rather steep, convex;
striae 1 strongly impressed, others not impressed; interstriae 1 con-
vex, punctured except at top, 2 higher than 1 (except median half
on right side) and bearing a row of rather large, rounded granules,
3 as high as 2 and similarly armed. Vestiture almost limited to de-
clivity, except on interspace 1, consisting of interstrial rows of erect,
tapered bristles, absent on lower three- fourths of 1 ; each bristle
about one and one-half to two times as long as distance between rows.

Female. — Similar to male except epistomal callus and brush
poorly developed.

Type Locality. — Five miles northeast of Teziutlan, Puebla,
Mexico.

Type Material. — The male holotype, female allotype, and 16
para types were collected at the type locality on June 27, 1953, at an
elevation of 1,500 m., by S. L. Wood, from a woody vine less than
1 cm. in diameter.

Cnesinus colombianus, n. sp.

Although not closely related, this species is more closely allied to
punctatus Blandford than other species known to me. It may be
distinguished from Blandford's species by the larger size, by the dif-
ferent sculpture of the epistomal callus, by the more coarsely sculp-

Sept. 5. 1967 neotropical bark beetles 85

tured pronotum, by the granulate interspace 1 on elytral declivity,
and by other characters.

Male. — Length 2.9 mm. (paratypes 2.8-3.2 mm.), 2.3 times as
long as wide; color dark reddish-brown with elytra somewhat lighter.

Frons transversely impressed at level of occular emargination,
convex above and flattened below this point; surface above complete-
ly smooth and brightly shining, with a few minute punctures
laterally and toward vertex, lower area largely covered by a dense
brush of reddish bristles, reticulate and punctured at sides; pubescent
area with a low. sharply elevated, median carina, this carina divided
on lower third of flattened areas and the two branches continued as
an epistomal carina below brush but distinctly above epistomal mar-
gin, subhorizontal arms of carina smooth, shining, glabrous. Eyes
separated by a distance equal to 2.2 times the greatest width of an
eye.

Pronotum 0.95 times as long as wide; sides feebly arcuate on
basal two-thirds, widest in front of middle, broadly rounded in front;
surface coarsely, deeply punctured, most punctures longitudinally
confluent except at base; vestiture restricted to marginal areas.

Elytra 1.5 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather nar-
rowly rounded behind; striae impressed, the punctures obscured by
confluence; interspaces about twice as wide as striae, surface irregu-
lar, the punctures small, confused, some finely granulate on their
anterior margins. Declivity steep, convex, narrowly flattened on
lower half; striae impressed, the punctures clearly evident; inter-
striae slightly wider than striae, convex, 1 and 3 very slightly higher
than 2, each uniseriately, rather finely granulate with some punc-
tures intermixed. 1 and 2 reaching almost to apex, 3 joining 5 and
ending before 2, 4 ending just below middle of declivity. Vestiture
confined to a few scattered bristles in lateral area of declivity.

Female. — Similar to male except epistomal brush reduced, and
median frontal carina absent.

Type Locality. — El Bosque. Caicedonia, Valles. Colombia.

Type Material. — The male holotype, female allotype, and 24
paratypes were collected at the type locality on June 23. 1959. by
J. Restrepo, from dead branches.

The holotype, allotype, and paratypes are in my collection.

Cnesinus retifer, n. sp.

This species is allied to gracilis Blandford. but is readily distin-
guished by the very strongly convex frons. by the completely reticu-
late pronotum and elytra, and by the confluent punctures of the
declivital striae.

Adult. — Presumably a male, length 2.0 mm. (paratypes 1.9-
2.0 nun.), 2.9 times as long as wide; color very dark brown, almost
black.

The Great Basin Naturalist
86 STEPHEN L. WOOD Vol. XXVII, No. 2

Frons strongly convex almost to epistoma; surface coarsely reticu-
late, with minute, rather sparse granules; vestiture short, sparse,
inconspicuous. Eye very feebly sinuate on anterior margin.

Pronotum 1.1 times as long as wide; widest two-fifths of length
from anterior margin, sides straight and diverging from base, broadly
rounded in front; a conspicuous, smooth, shining callus extending
from anterolateral angles at widest point more than halfway toward
median line just behind anterior margin; remainder of surface
reticulate, dull, with rather large, shallow oval punctures in posterior
area, and small, deep, elongate punctures anteriorly; anterior punc-
tures often confluent, posterior punctures usually separate; glabrous.

Elytra 1.8 times as long as wide. 1.7 times as long as pronotum;
sides straight and subparallel to base of declivity, rather narrowly
rounded behind; striae moderately impressed, the punctures conflu-
ent; interstriae about one and one-half times as wide as striae, except
1 narrower, the punctures very fine, deep, uniseriate except at base
of 3. Declivity convex, rather steep; interstriae narrower than striae,
convex. 1 very slightly more strongly elevated. Vestiture confined
to declivity, consisting of interstrial rows of flattened bristles; each
bristle as long as distance between rows of bristles.

Only one sex represented in the type series.

Type Locality. — Fort Clayton, Canal Zone, Panama.

Type Material. — The holotype and five paratypes were collect-
ed at the type locality on December 22, 1963, at an elevation of about
30 m., by S. L. Wood, from the central axis of an unknown vine
that was about 0.5 cm. in diameter.

The holotype and paratypes are in my collection.

Cnesinus annectens, n. sp.

According to Schedl, this species is closely allied to guadeloupensis
Erggers. The description of Eggers' species, however, indicates that
the declivital interspace 2 is narrowed and eliminated before the
elytral apex. In this species it is wider than either 1 or 3 and extends
to the apex. The lack of anatomical specialization makes characteri-
zation of this species difficult; however, biologically it is unique, as
noted below, in that it constructs egg tunnels in the cambium rather
than in the central axis of the host.

Female. — Length 2.3 mm. (paratypes 1.9-2.5 mm.). 2.2 times
as long as wide; color brown with white vestiture.

Frons broadly impressed, almost concave, from upper level of
eyes to epistoma, an obscure epistomal callus indicated; surface
reticulate, obscurely below, coarsely above, with small indefinite
punctures rather uniformly, not closely distributed; vestiture rather
widely distributed in lateral areas, consisting of fine, long, yellow
hair, much longer above, some setae equal to three-fourths the dis-
tance between eyes; distance between eyes equal to twice greatest
width of an eye.

Pronotum 0.93 times as long as wide; very slightly wider at base,
sides very feebly arcuate on about basal two-thirds, broadly rounded

Sept. 5. 1967 neotropical bark beetles 87

in front; surface coarsely, not deeply punctured, the punctures evi-
dently reticulate, almost all confluent, the very narrow, irregular
interspaces smooth, shining; median line narrow, weakly raised from
anterior fourth to base; vestiture short, abundant, fine except almost
scalelike near all margins.

Elytra 1.4 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather broadly
rounded behind; striae rather shallowly impressed, the punctures
moderately large, deep; interspaces about twice as wide as striae,
convex, the punctures small, numerous, subgranulate, in three ob-
scure ranks. Declivity rather steep, laterally convex, impressed be-
tween third interspaces; striae and strial punctures impressed, not
reduced; interstriae as wide as striae, with granules slightly larger
than on disc, uniseriate. 1 and 2 continuing to apex, 3 obscured on
lower third. Vestiture abundant, erect, consisting of stout interstrial
bristles, median row on each interspace a third longer than ground
cover, but shorter than distance between rows.

Male. — Similar to female except frontal impression shorter, not
as deep, the frontal vestiture much shorter.

Type Locality. — Zamorano, Morazan, Honduras.

Host. — Verbisina agricolorum.

Type Material. — The female holotype, male allotype, and 116
para types were collected at the type locality on April 18, 1964, at an
elevation of 700 m., by S. L. Wood, from the lower parts of green
stems about 1-3 cm. in diameter of the host plant.

The holotype, allotype, and most of the paratypes are in my col-
lection; two paratypes are in the Schedl collection.

Habits. — The egg tunnel was cut in the cambium region about
half in the xylem and half in the phloem tissues. It was small and
rather broad, much as in most Scolytodes species, with about three
to eight large egg niches placed along one or more sides. The eggs
were mixed with frass and packed into these niches. The larvae fed
more or less in congress in the phloem tissues, scarcely engraving
wood, until approximately the second instar was completed. The
female parent in most of the older galleries had cut a tunnel from
the egg chamber to the pith along the central axis of the stem. All
third instar larvae present were in these pith tunnels. Larvae older
than the third instar were not present in the material studied. After
cutting the pith tunnel the parent adults appeared to abandon the
tunnels, evidently to begin a second brood.

Cnesinus adusticus, n. sp.

This species is very similar to adustus Schedl. but is distinguished
by the larger size, by the widely separated eyes, by the slightly deep-
er impression of the elytral declivity, and by the more deeply im-
pressed striae 1 on declivity. Males only compared; an authentic
female of adustus not at hand.

The Great B^lsin Naturalist

88 STEPHEN L. WOOD Vol. XXVII, No. 2

Female.— Length 2.9 mm. (paratypes 2.6-3.3 mm.), 2.4 times
as long as wide; color reddish -brown, with white and cinereous
vestiture.

Frons with an arcuate, sharply elevated, transverse carina at
level of occular emargination occupying median third; weakly con-
vex above, flattened below carina; most of lower area covered by a
brush of erect reddish bristles; upper area minutely, transversely
strigose, punctured and granulose at sides and above; lateral areas
above carina and below upper level of eyes bearing rather abundant,
long, erect, plumose, yellow setae; eyes separated by a distance equal
to 1.6 times the greatest width of eye (1.0 times in male adustus).

Pronotum 1.05 times as long as wide; widest in front of middle,
the sides weakly arcuate, rather broadly rounded in front; surface
longitudinally rather coarsely strigose, most punctures confluent;
vestiture scanty except in marginal areas, partly scalelike at base
and near anterior margin.

Elytra 1.5 times as long as wide, 1.5 times as long as pronotum;
striae impressed, the punctures rather large, very close; interstriae
about one and one-half times as wide as striae, surface irregular,
the punctures confused, some median ones partly granulate. De-
clivity moderately steep, impressed between third interspaces; striae
not impressed, the punctures rather large, deep; intrstriae 1 rather
weakly elevated, 2 impressed, essentially flat, 3 and evidently part
of lateral area forming a rather abrupt, moderately high crest, the
lateral interspaces ending in this rounded summit, declining and
obsolete before apex; each interspace bearing a row of small setiferous
granules. Vestiture fine, almost hairlike on anterior two-thirds of
disc, becoming coarse and scalelike behind; consisting of rather abun-
dant, short interstrial setae and median rows of longer, erect setae,
none as long as distance between rows.

Male. — Similar to female except frontal carina absent, upper
area more strongly convex, reticulate, lower area more broadly flat-
tened; epistomal brush somewhat reduced, yellow.

Type Locality. — Los Amendros, Paraiso (near Zamorano),
Honduras.

Host. — Acacia pennatula.

Type Material. — The female holotype, male allotype, and 37
paratypes were collected at the type locality on April 18, 1964, at an
elevation of about 700 m., by S. L. Wood, from tunnels in the central
axis of branches of the above host 0.5-4.0 cm. in diameter.

The holotype, allotype, and most of the paratypes are in my col-
lection; two paratypes are in the Schedl collection.

Cnesinus carinatus, n. sp.

Very close to flavopilosus Schedl, but distinguished by the more
widely separated eyes, by the more coarsely strigose pronotum to
base, by the much finer, essentially hairlike elytral pubescence, by
the female frons being slightly inflated and pubescent in the median

Sept. 5. 1967 neotropical bark beetles 89

area above the frontal carina, and by the male frons being much
more strongly, subconcavely impressed on a much larger area.

Female. — Length 2.7 mm. (paratypes 2.5-3.1 mm.). 2.7 times as
long as wide; color brown.

Frons with a transverse carina at level normally occupied by
occular emargination in this genus, rather weakly convex above,
flattened below this point; surface above and below carina reticulate,
finely punctured and pubescent; vestiture above yellow, shorter in
median area, below carina reddish, erect, arranged in a transverse
brush as in flavopilosus, not reaching epistomal margin; eyes separat-
ed by a distance equal to 2.3 times greatest width of eye. Eye entire.

Pronotum 1.09 times as long as wide; sides almost straight on
basal two-thirds, very slightly wider in front of middle; surface
rather coarsely, longitudinally strigose to base; vestiture confined to
marginal areas.

Elytra 1.8 times as long as wdde, 2.0 times as long as pronotum;
sides almost straight and parallel to base of declivity, broadly rounded
behind; striae impressed, the punctures small, impressed; interstriae
about twice as wide as striae, convex, surface somewhat irregular,
the punctures fine, confused, median row evidently subgranulate.
Declivity steep, rather strongly impressed between third interspaces
as in flavopilosus; striae 1 moderately impressed, others not impres-
sed, the punctures reduced; interspace 1 convex, moderately elevated,
2 impressed, elevated laterally, the elevation continuing to moderately
high summit on 3. each interspace with a row of fine granules.
Vestiture hairlike, those on median row of each interspace longer,
particularly on declivital interspace 3 and laterally.

Male. — Similar to female except frontal carina absent; frons
concavely impressed on lower two-thirds of area below upper level
of eyes, entire frons pubescent, but setae longer and more abundant
at sides and along epistomal brush.

Type Ixx:ality. — Four miles west of Ciudad Hidalgo, Michoacan,
Mexico.

Type Material. — The female holotype, male allotype, and 14
paratypes were collected at the type locality on July 16. 1953, at an
elevation of 2,000 m., by S. L. Wood, from twigs of a rosaceous tree.

The holotype. allotype, and paratypes are in my collection.

Leperisinus guatemalensis, n. sp.

This species is very closely related to californicus Swaine and
eventually may prove to be a subspecies. It is distinguished by the
presence of an almost complete submarginal row of small asperities
just behind the anterior margin of the pronotum, by the smaller
pronotal asperities, by the smaller crenulations along the elytral
bases, and by the complete absence of a median frontal carina in the
female.

Female. — Length 3.0 mm. (paratypes 2.4-3.1 mm.), 1.8 times
as long as wide; color as in californicus except the light scales usually
luteous.

The Great Basin Naturalist
90 STEPHEN L. WOOD Vol. XXVII, No. 2

Frons more broadly flattened to upper level of eyes, more strong-
ly convex above eyes, and devoid of median carina, otherwise as in
californicus.

Pronotum with an almost continuous row of small submarginal
asperities connecting groups of large anterolateral asperities, and
aspertities smaller, otherwise as in californicus.

Elytra with basal crenulations smaller, and general sculpture
f ner, otherwise as in californicus.

Male. — Similar to female except frons shallowly concave;
frontal sculpture variable, but evidently epistomal emargination
w^der than californicus.

Typf, Locality. — Volcan Pacaya, Esquintla Prov.. Guatemala.

Type Material. — The female holotype, male allotype, and 90
pa^atypes were collected at the type locality on June 1, 1964. at an
elevation of about 1,300 m., by S. L. Wood, from cut branche3 of
three very different host species none of which resembled Fraxinus.
The transverse biramous egg galleries were typical of the genus.

The holotype, allotype, and paratypes are in my collection.

Chramesus incomptus, n. sp.

This species resembles other species of Chramesus only in the
antennal club. It has a slender form, a unique sculpture of the pro-
notum, fine vestiture, and a sculpture of the male frons not found
elsewhere in the genus. Tentatively, it is placed in the subgenus Pro-
chramesus although it exhibits much more primitive characters than
other species placed there.

Male. — Length 2.0 mm. (paratypes 1.9-2.1 mm.), 2.3 times as
long as wide; body color brown.

Frons strongly, transversely impressed just below middle, upper
area flattened to vertex, lower area strongly raised to epistoma,
lateral areas rounded, unarmed; epistomal lobe conspicuous, with a
pair of small tubercles at its base; surface reticulate, with small
deep punctures at sides and above; vestiture fine, short, somewhat
more abundant at sides and above, rather inconspicuous. Antennal
club rather small, apically pointed.

Pronotum 0.9 times as long as wide; widest about one-third of
pronotum length from base, the sides moderately arcuate, distinctly
constricted just behind anterior margin; surface minutely granulate
and with small, moderately close, shining, rounded granules; vesti-
ture consisting of rather long, moderately abundant hair.

Elytra 1.35 times as long as wide, 1.7 times as long as pronotum;
sides almost straight on basal two-thirds, very slightly wider pos-
teriorly, broadly rounded behind; striae rather weakly impressed,
the punctures rather large, deep; interstriae slightly wider than
striae, weakly convex, surface irregular, the punctures rather numer-
ous, confused, most granulate or subgranulate on anterior side; inter-
space 2 subcrenulate to well behind base. Declivity rather steep,
broadly convex; only striae 1 weakly impressed, the punctures some-

Sept. 5. 1967 neotropical bark beetles 91

what reduced in size; interstriae almost smooth, 3 and upper third of
1 and 2 with sparse, uniseriate, rounded granules. Vestiture moder-
ately long, largely abraded, consisting of coarse, delicate, hairlike
setae, toward declivity median interstrial rows becoming distinctly
longer than similar, shorter, ground vestiture.

Female. — Similar to male except frons irregularly convex with
an indefinite, subfoveate, median impression, surface reticulate-
granulate; epistomal tubercles present; elytral declivity more nar-
rowly convex, the interstrial tubercles on 2 and especially 1 extend-
ing to middle or below.

Type Locality. — Twenty-one miles west of Morelia. Michoacan.
Mexico.

Host. — Clematis sp.

Type Material. — I'he male holotype, female allotype, and 15
paratypes were collected at the type locality on June 16, 1965, at an
elevation of about 2,200 m., by S. L. Wood, from tunnels in the
cambium region of stems of the above vine larger than 1 cm. in
diameter. The biramous tunnels were diagonal; usually two females
were associated with each male.

The holotype. allotype, and paratypes are in my collection.

Chramesus bicolor, n. sp.

This is the only species in the genus having a sharply defined,
scale, color pattern. It evidently is allied more closely to crenatus
Wood than to other species; however, it differs by the more deeply
punctured pronotum, by the larger scutellum and scutellar notch, by
the deep strial punctures and by other characters.

Male. — ^Length 1.5 mm. (paratypes 1.4-1.7 mm.), 1.6 times as
long as wide; body color very dark brown, with white scales except
dark scales on a subcordate area occupying median third of pro-
notum anterior to basal fourth, and on elytra a pair of dark areas on
posterior half of disc from interspace to 2 to 8 but not extending on
declivity.

Frons broadly excavated from vertex to epistoma, the lateral
margins below eye acute, armed just below level of antennal base by
a pair of small teeth; surface reticulate, punctures very fine; vesti-
ture short, sparse.

Pronotum 0.83 times as long as wide; widest at base, the sides
arcuately converging, then distinctly constricted just behind the
broadly rounded anterior margin; surface rather coarsely and deeply
punctured, dull; surface largely obscured by scalelike vestiture. each
scale at least four times as long as wide, a few slender bristles inter-
mixed.

Elytra 1.0 times as long as wide, 1.3 times as long as pronotum;
sides straight and parallel on basal half, broadly rounded behind;
striae somewhat impressed, the punctures large, rather deep; inter-
striae slightly wider than striae, moderately convex, the punctures
squamiferous. confused, the median row subvulcanate. largely ob-

The Great Basin. Naturalist

92 STEPHEN L. WOOD Vol. XXVII, No. 2

soured by the vestiture. Declivity steep, convex; sculpture as on disc
except striae and interstriae somewhat narrower. Vestiture consist-
ing of short, broad scales, each scale as wide as long, and median
interstrial rows of erect longer scales, each about five times as long
as wide and almost as long as distance between rows of scales.

Female. — Similar to male except frons weakly convex, unarmed;
anterolateral areas of pronotum more coarsely asperate.

Type Locality. — La Lima. Cortez. Honduras.

Host. — Cestrum scandens.

Type Material. — The male holotype, female allotype, and 61
para types were collected at the type locality on May 5, 1964, at an
elevation of about 50 m., by S. L. Wood, from stems of the above
host less than 0.5 cm. in diameter.

The holotype, allotype, and paratypes are in my collection.

Chramesus vastus, n. sp.

This species differs from all others in the genus known to me by
its large size, by the sculpture of the pronotum. and by the scalelike
ground vestiture of the elytra with median interstrial rows of longer,
hairlike setae.

Male. — Length 2.4 mm. (paratypes 2.4-2.7 mm.), 1.7 times as
long as wide; color dark brown.

Frons broadly, shallowly excavated, the lateral margins acutely,
rather strongly elevated and armed just below level of antennal in-
sertion by a pair of tubercles; epistomal margin moderately, gradual-
ly elevated; surface coarsely reticulate, with fine punctures on mar-
ginal areas; vestiture inconspicuous.

Pronotum 0.8 times as long as wide; widest just behind middle,
the sides strongly arcuate, moderately constricted just behind an-
terior margin, broadly rounded in front; surface reticulate, dull, with
numerous small, isolated, shining asperities uniformly distributed
from base to anterior margin; vestiture consisting of short, sparse,
coarse hair.

Elytra 1.1 times as long as wide, 1.6 times as long as pronotum;
sides straight and parallel on basal half, broadly rounded behind;
striae 1 and 2 deeply, others moderately impressed, the punctures
small, subconfluent on 1 and 2; interstriae about twice as wide as
striae, weakly convex, the punctures shallow, minute, and each with
a median row of large tubercles except small on base of 1 and a par-
tial double row at base of 2. Declivity rather steep, convex; striae
and interstriae narrower than on disc; the tubercles narrower and
higher, except absent on lower half of interstriae 1 and 2. Vestiture
consisting of very small, abundant scales, each scale slightly longer
than wide except hairlike along suture, and median rows of erect,
interstrial. hairlike bristles, each bristle slightly shorter than distance
between rows.

Sept. 5. 1967 neotropical bark beetles 93

Female. — Similar to male except frons convex; pronotum with
punctures on disc impressed and feebly or not at all asperate toward
base, lateral areas more strongly asperate than in male; elytral striae
more deeply impressed, interstrial tubercles continuing to apex on
1 and 2.

Type Locality. — Cerro Punta. Chiriqui Prov., Panama.

Host. — Inga sp.

Type Material. — The male holotype. female allotype, and 17
paratypes were collected at the type locality on January 11-19, 1964,
at an elevation of about 2.000 m., from branches of the species of
Inga common on the hills west of Rio Viejo. The egg and larval
mines were in the woody tissues.

The holotype, allotype, and paratypes are in my collection.

Chramesus demissus, n. sp.

Somewhat intermediate between vastus Wood (above) and tumi-
dulus Blandford. It may be distinguished from tumidulus by the
reticulate, sparsely asperate pronotum, by the much larger elytral
tubercles, and by the less abundant elytral ground vestiture. From
vastus it is distinguished by the smaller size, by the less deeply im-
pressed, wider elytral striae, and by the very shallowly impressed
male frons.

Male. — ^Length 2.1 mm. (paratypes 2.0-2.3 mm.), 1.7 times as
long as wide; color dark brown.

Frons irregularly flattened, the lateral margins weakly elevated
and armed by a pair of very small granules below level of antennal
nsertion; surface rather coarsely reticulate, the punctures minute;
vestiture short, inconspicuous.

Pronotum 0.8 times as long as wide; widest behind middle, sides
strongly arcuate, moderately constricted behind the narrowly round-
ed anterior margin; surface reticulate, dull, with rather numerous
small, isolated asperities uniformly distributed, except median basal
area, with fine, shallow punctures; vestiture consisting of sparse,
slender scales.

Elytra 1.1 times as long as wide, 1.6 times as long as pronotum;
sides straight and parallel on slightly more than basal half, broadly
rounded behind; striae moderately impressed, the punctures rather
large, deep; interstriae wider than striae, essentially smooth, the
punctures very fine, each with a uniseriate row of coarse tubercles,
interspace 2 with double row of tubercles at base. Declivity steep,
convex; declivital granules smaller than on disc. Vestiture consisting
of small scales on posterior part of disc and on declivity, each scale
longer than wide; and median rows of erect, interstrial bristles, each
bristle slightly shorter than distance between rows of bristles.

Female. — Similar to male except lateral margins of frons not
elevated.

Type Locality. — Volcan Pacaya, Elsquintla Prov.. Guatemala.

The Great Basin Naturalist
94 STEPHEN L. WOOD Vol. XXVII, No. 2

Type Material. — The male holotype, female allotype, and 11
paratypes were collected at the type locality on June 1, 1964, at an
elevation of 1,300 m., by S. L. Wood, from small twigs of an un-
known tree seedling.

The holotype, allotype, and paratypes are in my collection.

Chramesus punctatus, n. sp.

This species is closely related to demissus Wood, described above,
but is readily distinguished by the very coarse strial punctures and
interstrial granules, by the longer declivital bristles on the elytra
(except usually absent on interspace 2). and by the deeper frontal
excavation of the male

Male. — ^Length 1.9 mm. (paratypes 1.5-2.1 mm.), 1.7 times as
long as wide; color dark brown.

Frons broadly, rather deeply excavated from upper level of eyes
to epistoma. the lateral margins rather strongly, acutely elevated and
armed by a pair of small tubercles just below level of antennal bases;
vestiture inconspicuous. Antennal scape with a small tuft of hair.

Pronotum 0.8 times as long as wide; widest on basal fourth, the
sides moderately arcuate, feebly constricted just behind the narrowly
rounded anterior margin; surface reticulate, dull, with numerous
small, isolated, shining asperities and in posteromedian area with a
few small punctures; vestiture consisting of short, semirecumbent
bristles.

Elytra 1.1 times as long as wide. 1.5 times as long as pronotum;
s'des straight and parallel on more than basal half, broadly rounded
behind; striae moderately impressed, the punctures very large, deep;
interstriae about as wide as striae, convex, irregular, the punctures
minute, each interspace with a row of very coarse, isolated tubercles.
Declivity moderately steep, broadly convex; striae and interspaces
somewhat narrower; interspaces 2 and 4 devoid of tubercles (some
paratypes have one or more on each). Vestiture consisting of numer-
ous small scales on declivity, and median interstrial rows of erect
bristles from base to apex; each bristle as long as distance between
rows of bristles and between adjacent bristles in a row; declivital
interspaces 1 and 4 (usually) devoid of bristles (in a few paratypes
both bear one or more bristles) .

Female. — Similar to male except frons weakly convex.

Type Locality. — Volcan Pacaya, Esquintla, Guatemala.

Host. — A vine, evidently Canavalia sp.

Type Material. — The male holotype, female allotype, and 90
paratypes were collected at the type locality on June 1. 1964. at an
elevation of about 1.300 m., by S. L. Wood, from a vine thought to
be Canavalia sp.; 29 additional paratypes bear similar data but were
taken 14 km. west of Guatemala City, on May 30. 1964. from the
same host.

The holotype. allotype, and paratypes are in my collection.

Sept. 5. 1967 neotropical bark beetles 95

Phloeotribus simplex, n. sp.

This species is more closely related to opimus Wood than to other
representatives of the genus known to me, but may be distinguished
by its much more finely sculptured pronotum and elytra, by the re-
ticulate surface of the pronotum. by the longer vestiture. and by
other characters.

Female. — Length 1.5 mm. (paratypes 1.4-1.7 mm.), 2.1 times as
long as wdde; color brown.

Frons convex, with a transverse callus at level of antennal inser-
tion; surface coarsely reticulate, with rather coarse, deep, moderately
abundant punctures; vestiture inconspicuous. Antennal club with
segment 1 about twice as wide as long.

Pronotum 0.9 times as long as wide; widest one-fourth from base,
sides feebly arcuate on basal half, rather strongly constricted just
behind broadly rounded anterior margin; surface unarmed, reticu-
late, the punctures small, deep, separated by distances of one to three
times the diameter of a puncture; vestiture consisting of short, fine,
and longer, coarse, hairlike setae.

Elytra 1.3 times as long as wide, 1.7 times as long as pronotum;
basal crenulations in a single row. rather high; striae not impressed,
the punctures deep, coarse; interstriae as wide as striae, almost flat
and smooth with median rows of small setiferous granules. Declivity
steep, convex, rather narrow; striae and interstriae narrower than on
disc. Vestiture consisting of rows of fine, recumbent, strial hair, and
longer, erect, uniseriate rows of interstrial bristles, each bristle some-
what flattened at its apex and spaced in the rows and between rows
by distances about equal to length of a bristle.

Male. — Not represented in the series at hand.

Type Locality. — Fort Clayton, Canal Zone. Panama.

Type Material. — ^The female holotype and 6 paratypes were
collected on December 22. 1963. at an elevation of about 30 m.. by
S. L. Wood, from the inner tissues of a large vine about 5 cm. in
diameter. Six other paratypes were taken at Barro Colorado Island,
Panama Canal Zone, on December 27, 1963, at an elevation of about
60 m.. by S. L. Wood, from the same species of vine. Scolytus
nodatus also infested this same species of vine.

Chaetophloeus minimus, n. sp.

The small size and vestiture of this species are more nearly like
Liparthrum than other species in this genus, but the antennal and
other characters clearly place it in Chaetophloeus. It is allied to
meiicanus (Blackman). but may be distinguished by the smaller
size, by the more slender form, by the less strongly impressed striae,
by the vestiture. and by the less strongly impressed male frons.

Male. — Length 1.2 mm. (paratypes 1.1-1.4 mm.). 1.9 times as
long as wide; color almost black, with white vestiture.

Frons shallowly concave from eye to eye and from epistoma to
vertex; surface reticulate-granulate, vestiture consisting of moderate-

The Great Basin Naturalist
96 STEPHEN L. WOOD Vol. XXVII, No. 2

ly long, coarse hair, more abundant and longer at sides and above.
Antennal club large. 1.8 times as long as wide; three sutures indi-
cated.

Pronotum 0.6 times as long as wide; widest one-fourth of prono-
tum length from base, sides moderately arcuate on basal half, rather
weakly constricted just behind anterior margin; surface subreliculate-
granulate. with moderately large, obscure punctures; lateral area
armed by one paired group of asperities; vestiture consisting of short,
stout, hairlike bristles and equally abundant, erect scales, each scale
as wide as long.

Elytra 1.3 times as long as wide, 2.0 times as long as pronotum;
basal crenulations as in mexicanus, with two pair of submarginal
ones on interspaces 1 and 2; striae not impressed, the punctures
moderately large, deep; interstriae feebly convex, as wide as striae,
the punctures fine, confused. Declivity steep, convex; striae and
interstriae somewhat narrower than on disc, the interstrial punctures
evidently subgranulate. Vestiture consisting of shor;;, slender nter-
strial bristles along each interspatial margin, and median, interstrial
rows of erect, round scales; scales separated within a row by dis-
tances equal to length of a scale, and between rows by distances equal
to twice the length of a scale.

Female. — Smilar to male except frons weakly convex, minutely
granulate, the vestiture short, sparse; pronotum armed by two pairs
of groups of asperities.

Type Locality. — Two miles east of Armeria, Colima, Mexico.

Type Material.- — The male holotype, female allotype, and 15
para types were collected at the type locality on June 28, 1965. at an
elevation of about 60 m., by S. L. Wood, from an unidentified, thorn-
less shrub with simple leaves.

The holotype, allotype, and paratypes are in my collection.

Chaetophloeus struthanthi, n. sp.

This species is closely related to phoradendri Wood, but is readily
distinguished by the shorter, erect, elytral scales, by the more
strongly convex, granulate, female frons, by the shorter, yellow
pubescence on upper part of male frons, and by the smaller, mandi-
bular processes in both sexes.

Male. — Length 1.7 mm. (paratypes 1.6-1.9 mm.), 1.6 times as
long as wide; color very dark brown.

Frons shallowly concave from eye to eye and from vertex to the
broadly emarginate epistomal margin; surface with rather close, fine,
subgranulate punctures; vestiture largely confined to marginal areas,
longer on upper margin, but longest hairs not reaching ep'stoma,
yellow. Antennal club 2.1 times as long as wide; sutures very
obscure.

Pronotum 0.6 times as long as wide; widest one- fourth of pro-
notum length from base, sides very strongly arcuate on basal half.

Sept. 5. 1967 neotropical bark beetles 97

moderately constricted just behind the broadly rounded, subemar-
ginate. anterior margin; surface subrugulose, rather finely, closely
punctured, lateral areas armed by three pairs of groups of asperities;
vestiture consisting of short, erect scales, about eight much longer
ones in median basal area.

Elytra 1.1 times as long as wide, 2.1 times as long as pronotum;
outlines and basal asperities as in phoradendri; striae weakly im-
pressed, the punctures rather small, shallowly. distinctly impressed;
interspaces convex, about twice as wide as striae, the setiferous punc-
tures small, confused. Declivity convex, steep; striae and interstriae
narrower and less clearly marked than on disc. Vestiture consisting
of short, semierect. small, slender scales of equal length on disc,
median row on each interspace slightly longer on declivity.

Female. — Similar to male except frons convex above, flattened
below, more nearly granulate; mandibular process reduced; antennal
club 1.7 times as long as wide.

Type Locality. — Volcan de Colima, Jalisco. Mexico.

Host. — Struthanthus, probably venetus.

Type Material. — The male holotype, female allotype, and 86
paratypes were collected at the type locality on June 23, 1965, at an
elevation of about 2,500 m.. by S. L. Wood, from the above mistletoe
that grew in an oak tree.

The holotype, allotype, and paratypes are in my collection.

A NEW SPECIES OF SPINIBDELLA FROM
UTAH (BDELLIDAE: ACARINA)i

Clive D. Jorgensen-

Atyeo (A revision of the mite family Bdellidae in North and
Central America [Acarina, Prostigmata], Bull. Univ. Kansas 40(8):
345-499. 1960 reported Spinibdella corticus (Ewing) and cronini
(Baker and Balock) from Utah in his revision of the family Bdellidae.
Apparently additional records of this genus have not been reported
from the state since. The following description is of a species of
Spinibdella collected from apple tree bark at Spring Lake, Utah Co.,
Utah. I am grateful to Warren T. Atyeo for his examination of this
species.

Spinibdella mali, n. sp.

Diagnosis. — The new species is similar to depressa (Ewing), but
can easily be separated by the presence of two pairs of eyes. It is
separated from bif areata (Atyeo) by the first interspace which is
more than twice as long as the length of the internal dorsals. Repeat-
ed efforts to collect additional specimens have not been successful.

Female. — Color unknown. Body striated throughout; divided by
a suture into propodosoma and hysterosoma; length, including
gnathosoma, 696/a. Gnathosoma length, 165/a; chelicera (Fig. 3)
striated; length, 134ju,; two strong setae; small sharp chela. Palpi
(Fig. 4) short, striated; tibiotarsus extending beyond hypostome;
palpal segment lengths: I, 10/x; II plus III, 111/x; IV, IG/x; V, 41 /x,;
ventral end seta, 116/a; dorsal end seta, 152fx. Hypostome with stria-
tions transverse at proximal end, longitudinal at distal end; two pairs
of strong ventral setae; length, 149ju,. Dorsal propodosoma (Fig. 1)
with infrequently broken striae; two pairs of eyes, the posterior pair
being larger than the anterior. Propodosomals slightly plumose; an-
terior 41 /A long, posterior 31jU, long; distance between anterior pro-
podosomals 75/a; distance between posterior propodosomals 46 /x; dis-
tance between anterior sensillae, 39ju,; distance between posterior
sensillae, 95/a; striae directed forward between posterior propodoso-
mals. Dorsal hysterosoma striations only occasionally broken. Setae
all slightly plumose; length of external humerals, 44/i; internal hu-
merals, 31 /a; internal dorsals, 31/x; internal lumbrals, 34ju,; internal
sacrals, 34/x,; external sacrals, 39/x; internal clunals 36^; post anals,
36;ii. Distance between external and internal humerals, 83fx; between
internal humerals and internal dorsals (first interspace). 77 ix; be-
tween internal dorsals and internal lumbrals. 85/a; between internal
lumbrals and internal sacrals, 106/a. Anal border striae parallel; two

1. This study was supported in part by a grant (12-14-100-8029(33) from the U. S. Department
of Agriculture to Brigham Young University.

2. Department of Zoology and Entomology, Brigham Young University, Provo, Utah.

98

Sept. 5, 1967 new species of spinibdella

99

Figure 1. Dorsal view of Spinibdella mali.

100

CLIVE D. JORGENSEN

The Great Basin Naturalist

Vol. XXVII, No. 2

Figure 2. Genital area of Spinibdella mali.
Figure 3. Chelicera of Spinibdella mali.
Figure 4. Left palpus of Spinibdella mali.

Sept. 5. 1967 new species of spinibdella i 101

pairs of anal setae, one pair of paranals; anal setae undulate but not
plumose; paranals slightly plumose; anterior anals length 23/i,; pos-
terior anals length, 28/a; paranals length, SA-fi. Genital plates (Fig. 2)
with six pairs of genital setae; twelve pairs of paragenitals; setae in
genital area appear brittle, several broken, always blunt and un-
dulate. Legs with small strong unadorned claws, shorter than pre-
tarsus. Measurements: tibia I-IV. 54. 59. 61. 72/i,; tarsus I-IV. 75.
77, 88, 90ju. Chaetotaxy: trochanter I-IV, 1, 1. 2, 1; basifemur 1-IV.
5, 5. 6. 3; telofemur I-IV. 4. 4. 4, 4; genu I-IV. 5, 5, 6, 7; tibia I-IV,
12, 11, 11, 10; tarsus II-IV. 16. 17, 16; one small blunt sensory rod
of tibia II deeply recessed. Condition of specimen prevented more
detailed descriptions of leg chaetotaxy.

Male. — Unknown.

HoLOTYPE. — A single female was collected from the bark of an
apple tree at Spring Lake, Utah Co., Utah, on September 28, 1965.
The type specimen is deposited in the author's collection at Brigham
Young University. Provo. Utah.

NOTES ON ERIOGONUM — III
ON THE STATUS OF ERIOGONUM PAUCIFLORUM PURSH

James L. Reveal '• -

In 1814, Fredrick Pursh published in the appendix to his Flora
Americae Septentrionalis, several interesting and new taxa that had
been collected in North America by John Bradbury. Among the
new species was Eriogonum pauciflorum. In the spring of 1966, for
a history class, I reviewed Susan Delano McKelvey's book. Botanical
Exploration of the Trans-Mississippi West, 1790-1850, and realized
that the range of E. pauciflorum, as understood by the Great Plains
and Rocky Mountain botanists did not get near the Missouri River,
nor into North Dakota where Bradbury (1817) stated that he made
his collection. To these botanists, the epithet E. pauciflorum referred
to a plant of southeastern Wyoming and adjacent northern Colorado.

Through the kindness of Dr. A. E. Schuyler. I was able to study
the holotype of Eriogonum pauciflorum in the Pursh Herbarium
which is now deposited at the Academy of Natural Sciences in Phila-
delphia. The examination confirmed earlier suspicions that the name
E. pauciflorum was misapplied. The plant that Bradbury had collect-
ed has been known for several years as E. multiceps Nees or E.
gnaphalodes Benth. in Hook.

A Resume of the Discoveries

John Bradbury is a relatively unknown early botanist, although
as an early writer on the area of the upper Missouri River, he is
far better known. He was born near Stalybridge in Lancashire, Eng-
land, in 1768. but did not come to America until 1809 when he was
41. Following the suggestions of Thomas Jefferson, Bradbury went
to Saint Louis where he established his headquarters rather than
at New Orleans as he had originally planned. This change was
fortunate as it certainly allowed him to gather a far more interesting
and unknown flora under the auspices of the Liverpool Botanic
Garden (Rickett, 1950).

Unknown to Bradbury, a fellow Englishman. Thomas Nuttall,
was setting out to study the western flora at the same time. Nuttall
was born in 1 786 in the village of Long Preston in Craven, England,
and came to America ". . .in the spring of 1807 or 1808, when he
was twenty-one or twenty-two. . ." (Pennell, 1936). Almost as soon

1. The author wishes to thank the United States National Herbarium and the Smithsonian
Institution which sponsored his Predoctoral Internship program in Washington, D.C.. from^ Septem-
ber I'Kifi to Februarv- ]967 where this paper was basically prepared. He is grateful to C. V. Morton
and S. 11. Shelter tor their many helpful suggestions and assistance in obtainmg rntual herbarium
material. The assistance of Dr. C. L. Porter of the University of Wyoming on various aspects
concerning the Wyoming taxa is gratefully acknowledged. The author also wishes to thank the
(urators whose facilities were visited or who kindly sent loan material. This paper has been sub-
mitted t'l the Department of Botany. Brigham Young University, as partial fulfillmenl of two
credits of Spc<ial Problems given during the Summer Se.s.sion of 1967.

1. Depiirtmrnt of Bnlany. Brigham 'NDung University, Provo. Utah

102

Sept. 5. 1967 notes on eriogonum - in 103

as Nuttall arrived, he became acquainted with Benjamin Smith
Barton of Philadelphia. Nuttall. who had collected every curious
plant that he saw walking into the city, had taken his unknown col-
lection to Barton for determination. Barton's previous assistant,
Fredrick Pursh. had recently left, and Barton must have been im-
pressed by Nuttall for he soon hired the young man to collect plants
for him.

In the year 1810. while Bradbury was busy collecting in the
Saint Louis area, Nuttall was setting out to collect in the old "North-
west" under an exact set of directions given to him by Barton (Grau-
stein. 1950). Nevertheless, Nuttall was soon to forget his directions.

How Nuttall and Bradbury met is not known, nor is it exactly
known how the two men came to arrange to go up the Missouri River
with the Overland Astorians. In a series of letters reprinted by Rickett
(1950), Bradbury reveals that in late January the boat on which he
had sent his 1810 collection to New Orleans had run aground below
Saint Louis (in Bradbury's Travels, he says "autumn") and it was
upon his return to the city following his investigation of the condi-
tion of the boat and his collection that he learned of the Astorians.
Wilson Price Hunt, the leader of the Astorians, had been in the
Saint Louis vicinity since September 1810. but to avoid the expense
of over-wintering in the city, he had established a w'nter camp near
St. Joseph. Missouri (Irving, 1836). Apparently Hunt returned to
Saint Louis around the 20th of January, 1811, and no doubt it was
at this time that Bradbury (no mention of Nuttall) was informed of
Hunt's intention to ascend the Missouri River. On the first of
March, Hunt returned to Saint Louis from his winter camp to make
the necessary final arrangements for his fur trapping expedition
into the Pacific Northwest (Bradbury, 1817).

The competition for the fur trade in the Oregon Country was
intense between all companies engaged. The North West G)mpany
and the Hudson's Bay Company in Canada and in the Oregon
Country were locked in bitter, and occasionally bloody, competition.
Likewise, the Missouri Fur Company under Manuel Lisa and the
American Fur Company under John Jacob Astor were rapidly be-
coming arch rivals. And in the middle of the American rivalry were
two botanists, John Bradbury and Thomas Nuttall.

Hunt, during his trips to Saint Louis, was trying to obtain ex-
perienced men who had worked for the Missouri Fur Company in
the past. At the same time, he was also attempting to purchase equip-
ment for his trip. Manuel Lisa, the leader of the Missouri Fur Com-
pany, quickly realized that Hunt was causing a drain of men and
equipment that could reflect in the fur take of his own company.
One of the men that Hunt hired was Pierre Dorion. a half-breed who
had been Lisa's interpreter. When Lisa found that E>orion was going
to work for the Astorians, he had a warrant issued for Dorion's
arrest as he had an outstanding liquor bill which had been incurred
in the mountains. Bradbury and Nuttall, who had remained behind
in Saint Louis for the last mail after Hunt had left, overheard the

The Great Basin Naturalist
104 JAMES L. REVEAL Vol. XXVII, No. 2

sheriff's plans to apprehend Dorion at St. Charles. By leaving shortly
after midnight on March 13th and traveling rapidly overland the
two botanists, now turned spies, informed Hunt of the pending
arrest, and Dorion with his wife and child fled into the woods and
were not found by the sheriff. The incident involving Dorion has
been interpreted in two ways. First. Irving suggests that Lisa's
actions were to prevent Hunt from using the services of Dorion;
Oglesby ( 1 963 ) , on the other hand, suggests that Lisa simply wanted
to travel with Hunt and his large party in order to ward off Indian
attacks.

The slow trip up the Missouri River has been discussed at some
length by various writers on both expeditions (Brackenridge, 1814.
with Lisa; Bradbury. 1817, with Hunt), and by various authors
afterwards (Irving, 1836; Oglesby, 1963). Parts of the expeditions
have been discussed in the botanical literature by Pennell (1936).
Rickett (1950), McKelvey (1955), and Stevens (1946, 1959).

It is obvious from these various authors that Nuttall and Brad-
bury remained distant. In only four places does Bradbury even
mention his fellow naturalist in his Travels. Although the two men
\yere both good botanists, they had widely different outlooks on
their trip and what was necessary to observe. Irving, who unfortu-
nately has been critized as an historian, has written an excellent
summary on the two men which is rather revealing:

Mr. Nuttall seems to have been exclusively devoted to his scientific
pursuits. He was a zealous botanist, and all his enthusiasm was awak-
ened at beholding a new world, as it were, opening upon him in the
boundless prairies, clad in the vernal and variegated robe of unknown
flowers. Whenever the boats landed at meal times, or for any temporary
purpose, he would spring on shore and set out on a hunt for new speci-
mens. Every plant or flower of a rare or unknown species was eagerlj'
seized as a prize. Delighted with the treasures spreading themselves out
before him, .he went groping and stumbling along among a wilderness
of sweets, forgetful of every thing but his immediate persuit, and had
often to be sought after when the boats were about to resume their
course. At such times he would be found far off in the prairies, or up the
course of some petty stream laden with plants of all kinds.

The CanacUan voyageurs, who are a class of people that know noth-
ing out of their immediate line, and with constitutional levity make a jest
of any thing they caimot understand, were extremely puzzled by this
passion for collecting what they considered mere useless weeds. When
they saw the worthy botanist coming back heavy laden with his speci-
mens, and treasuring them up as carefully as a miser would his hoard,
they used to make merry among themselves at his exp^ense, regarding
him as some whimiscal kind of madman.

Mr. Bradbury was less exclusive in his tastes and habits, and com-
bined the hunter and sportsman with the naturalist. He took his rifle or
his fowling piece with him in his geological researches, conformed to
the hardy and rugged habits of the men around him, and of course
gained favor in their eyes. He had a strong relish for incident and ad-
venture, was curious in observing the savage manners, and savage life,
and ready to join any hunting or other excursion.

Thus it was that much of their collection was duplicated, but it
was also this desire of Bradbury's for adventure that set him out for
Fort Lisa on horseback while Nuttall. certainly much less of an

Sept. 5. 1967 notes on eriogonum - iii 105

adventurer, remained behind to go up the river by boat. It was
this trip that possibly allowed Bradbury to discover Eriogonum
pauciflorum, and for Nuttall to miss it.

On the 3rd of June 1811, Lisa, who had left St. Charles on April
2nd. finally caught up with Hunt just below the Arikara Villages.
From now on the small flotilla of five boats moved upstream, but
with Hunt always in the front so that Lisa could not pass him. The
parties finally arrived at the Villages, which were located near
present-day Wakpala, Corson County, South Dakota, on the west
bank of the Missouri River some six miles above the mouth of the
Grand River.

On the 19th of June, Hunt and Lisa settled some of their dif-
ferences, and a trade of Hunt's boats for some of Lisa's horses had
been arranged. It was decided that a party should be sent overland
for the horses which were at the Missouri Fur Company's fort sev-
eral miles upstream. When Bradbury learned of the pending trip,
he ". . . declared to Mr. Hunt that, unless he absolutely refused
me the privilege, I was determined to accompany them." Hunt gave
his consent, and mounted on a bare wooden saddle on a poor horse,
Bradbury set off. often collecting plants and putting them under
his hat until he had a moment to dismount and press them. On the
evening of the 22nd, the party reached the fort.

There has been considerable confusion in the botanical literature
as to the locations of the forts and Indian villages on the upper
Missouri River, and to some degree, this is due to a similar confusion
in the historical literature on this period. While I certainly do not
wish to proclaim to have the final word on this subject, some new
and hopefully enlightening information can be presented.

In attempting to determine the exact localities mentioned by
Bradbur>\ I wrote the State Historical Society of North Dakota' at
the suggestion of Dr. LeRoy R. Hafen, Professor of Far Western
History. Brigham Young University. The information that was
obtained from Sperry contradicts, to some degree, wdth the traditional
data given as to the location of the forts and Indian villages, and
this is presented here in some detail so that a clearer picture of their
locations might be understood.

In 1804. Lewis & Clark built Fort Mandan where they over-
wintered in 1804-1805. Nevertheless, in 1811 this site was aban-
doned, for Lewis & Clark reported (1814) that when they returned
in August 1806. thev found that a large portion of the fort had
been burned. Sperry believes that the site was probably still obvious
when Bradbury visited the area. When Alexander Philip Maxi-
milian, a Prussian soldier and scholar, later visited the area in the
winter of 1883-1834 at Fort Clark, he mentioned (1841) that the
river had changed its course and that the site of Fort Mandan was
either on the opposite bank (the east bank) or in the middle of

\. In rospon'ie Id niv letlei , Ml. James K. Speny, Reseanh Archeologist, virrole on 1') January
|4()7 hi» reply whirh is the major basis for the disfussion on the locations of the forts and Indian
villages, ilis letter is deposited in my personal rorrcspondenrr and in that of C V. Morton's,
Smithsonian In.stitiilion. Washington. D.C.

The Great Basin Naturalist
106 JAMES L. REVEAL Vol. XXVII, No. 2

the river. Sperry comments that the exact site of Fort Mandan has
not been discovered, but according to the maps of Lewis & Clark
and of Maximilian, he believes that the fort was on the east bank
of the river and about 1 5 miles northeast of the site of Fort Clark in
present-day McLean County, North Dakota.

Sperry says that the site of Fort Lisa, which was later named
Fort Vanderbaugh, also has not been definitely located. Bradbury
(1817) indicated that after crossing the Knife River, they had ". . .
seven miles still to travel in order to reach the Fort. . ." If his dis-
tances are correct, that would put the site of the fort just over a mile
south of Mannhaven on the west bank of the Missouri River in
Mercer County.

In the story of Eriogonum pauciflorum, the locations of the
Indian villages are also important. Sperry writes that in 1811 there
were only two Minetaree Villages, both of which are still quite
evident. These villages were simply called the upper and lower
villages by Bradbury. The lower village was a small site about
one mile north of the town of Stanton on the south side of the Knife
River near the present-day Stanton Cemetery. The upper village,
which was considerably larger, was located about 2.8 miles north
of Stanton, or on the north side of the Knife River near the present
site of the Olds School. Thus, the type area of E. pauciflorum ap-
pears to be north of Stanton, Mercer County, rather than south of
Mannhaven (or the location of Fort Lisa) as was stated by Stevens
(1946).

From Bradbury's journal, it is apparent that he was in the
vicinity of the Minetaree Villages on three different days of June:
the 22nd when he rode through in the late evening; the 24th, at
which time he collected on the bluffs north of the upper village;
and the 28th when he was actually in and around the Minetaree
Villages. While Bradbury might have been able to collect the Erio-
gonum on the 22nd, it seems unlikely as it was late in the evening
and he knew that he was going to be in the area for a few days. The
bluffs north of the upper village which he visited on the 24th are
an excellent place for the Eriogonum to grow. However. Bradbury
specifically stated that the Eriogonum came from near the Minetaree
village, and it is assumed that the buckwheat was not collected on
this date. Therefore, it would seem most likely that Bradbury col-
lected the type of E. pauciflorum on the east facing slopes of the
low hills northwest of the upper village when he visited it on the
28th of June 1811.

The discovery of Eriogonum multiceps by Alexander Philip
Maximilian. Prince of Wied-Neuwied. can be traced with a little
more exactness. The holotype deposited at Jardin Botanique de I'fitat,
Bruxelles, Belgium, consists of two specimens, one simply says "Am
uberen Missouri" while the other gives the same information, a
collection number (#114), and a date, "6 July." On July 6th,
Maximilian was at Fort Union, a post near the North Dakota-Mon-
tana state line, where he had been since the 24th of June 1833.

Sept. 5, 1967 notes on eriogonum - iii 107

However, on the 6th, he reported that he was preparing to leave
for Fort McKenzie further up the river, and probably the collection
was not actually made on this day, but on a day before while still at
Fort Union as he does mention collecting plants there. It seems
likely from comparing recent specimens that have been collected in
this area, that the Maximilian collection could have come from the
Fort Union area, although in his book (1841), no mention of an
Eriogonum is made.

The entire collection of some two hundred plants was turned
over to Nees von E^enbeck of Poland, who prepared the botanical
appendix which appears in Maximilian's book. When Nees described
Eriogonum multiceps, he compared it with E. pauciflorum, and
unlike Pursh. adequately described the species. Unfortunately, the
publication Reise in das inner e Nord- America in den Jahren 1832
bis 1854. was not seen by the American botanists until the middle of
the century, and the name E. multiceps was not noted in print
until Torrey & Gray's 1870 revision of Eriogoneae.

In the meanwhile. Bentham had named Eriogonum gnaphalodes
which had been collected by Carl Andreas Geyer, a German col-
lector, probably in the Guernsey area, Platte Q)unty, Wyoming,
which is near Fort Laramie (Goshen County) which is the loca-
tion given by Geyer (1845-1846) for this plant. When Torrey &
Gray discovered that E. gnaphalodes and E. multiceps were basically
the same kind of plant, they placed the Bentham name into sy-
nonymy, and their concept of E. multiceps for the Great Plains
species has been followed since.

Key to the Species

A. Perianth pubescent externally; leaves grayish or whitish
tomentose on both surfaces, sheathing up the stems; com-
mon in eastern Wyoming and adjacent Nebraska northward
to extreme southern Canada.

1. E. pauciflorum

AA. Perianth glabrous externally; leaves densely tomentose
below, subglabrous or glabrous above; southeastern Wyoming
and Colorado.

B. Leaves linear to linear-oblanceolate, less than 3 mm
wide, tightly revolute; involucres usually glabrous ex-
ternally; gumbo clay to granitic soils, mostly below
8,500 feet; Grand Co., Colorado northward to Albany
and Larimer cos.. Wyoming.

2. E. exilifolium

BB. leaves oblanceolate to lanceolate or narrowly elliptic,
3-6 mm wide, not revolute; involucres usually tomen-
tose externally; granitic talus or gravelly slopes and
flats from 8,500 to 12,500 feet; El Paso Co. westward
to Gunnison Co.. Colorado.

3. E. coloradense

108

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 2

1. Eriogonum pauciflorum Pursh, Fl. Am. Sept. 2: 735. 1814.

Low spreading leafy-stemmed perennial herbs which form loose
mats up to 2 dm across; leaves linear-oblanceolate to oblanceolate. 3-5
(8) cm long, (1) 2-6 mm wide, grayish to whitish tomentose below,
less so to subglabrous and green above, sheathing up the stems 1-5
cm. tapering gradually to petioles (0.5) 1-2.5 cm long, the petiole-
bases appressed and clasping, 1.5-3 mm wide; flowering stems 5-15
(20) cm long, tomentose; inflorescences single capitate cluster of
several involucres or umbellate with rays simple and up to 5 cm
long, the involucres or the first node subtended by a whorl of 2-6
linear to semifoliaceous lanceolate bracts, 1.5-20 mm long, the inflo-
rescence head ca 1 cm broad; involucres narrowly turbinate. (3.5)
4-5 mm long, tomentose, the 5 acute teeth 0.5-0.8 mm long, triangu-
lar; perianth whitish-brown, cream, pink, or rose-tinged, usually
densely pubescent on the lower half or rarely glabrous, 2-2.5 mm
long, the calyx-segments similar, oblong, connate about l^ to 1/3 the
length of the perianth, the bases rounded or slightly acute, the apices
rounded; stamens 2.5-3 mm long, the filaments pilose at the bases,
the anthers 0.3-0.4 mm long, oblong; achenes brownish, 2 mm long,
the globose to subglobose bases tapering to scabrous 3-angled beaks.

• ••

• •

••

E. pauciflorum
var. pauciflorum •
va r. gnaphalodes a
var. nebraskense ▲
var. canum ^

E. ex i I ifol iu m n
E. coloradense o

*• •

\A *

o o O o

Figure 1. Distribution of Eriogonum paciflorum Pursh.

Sept. 5, 1967 notes on eriogonum - iii 109

Key to Varieties

A. Inflorescences capitate or nearly so, rays not compoundly

divided nor subtended by bracts.

B. Leaves tomentose below, somewhat less tomentose
above, linear-oblanceolate to oblanceolate, mostly 3-5
(8) cm long and up to 6 mm wide, not densely pube-
scent with long white, densely tangled and matted
hairs; inflorescences ca 1 cm broad; south-central
Canada south to eastern Wyoming.

la. var. pauciflorum

BB. Leaves densely matted with long white tangled hairs
on both surfaces of the leaves, the leaves narrowly
spathulate, mostly 2-3.5 cm long and up to 10 mm
wide; inflorescences ca 1-1.5 cm broad; eastern Wyom-
ing and adjacent western Nebraska.

lb. var. gnaphalodes

AA. Inflorescences compoundly branched, cymose.

B. Flowers white; extreme western Nebraska and ad-
jacent northeastern Colorado.

Ic. var. nebraskense
BB. Flowers yellow; central Montana.

Id. var. canum
la. Eriogonum pauciflorum var. pauciflorum

E. parviflorum Nutt., Journ. Phil. Acad. 1: 35. 1817, a superflu-
ous substitute for E. pauciflorum Pursh.

E. dioecium Raf., Herb. Raf. 38. 1833, nom. nud.; New Fl. N.
Amer. 4: 53. 1838. Type: On River Platte." Bradbury s.n. See com-
ments below.

E. multiceps Nees in Wied-Neuwied, Reise Nord-Amer. 2: 446.
1841. Type: Fort Union, Williams Co.. North Dakota, 6 Jul 1833,
Prince Alexander Philip Maximilian 114. Holotype: BR!

E. depauperatum Small. Bull. Torrey Bot. Club 25: 40. 1892.
Type: Hermosa, Custer Co., South Dakota, 23 Jun 1892, Rydberg
970. HoLOTYPES NY! Isotypes: NEB! US; A form with glabrous
flowers.

E. multiceps ssp. typicum S. Stokes, Gen. Eriog. 94. 1936.

Low spreading leafy-stemmed, loosely matted, perennial herbs;
leaves linear-oblanceolate to oblanceolate, leaf-blades 1-4 cm long,
( 1 ) 2-6 mm wide, grayish to whitish tomentose below, less so to sub-
glabrous and green above, gradually tapering to petioles 2-5 cm long;
flowering stems 5-20 cm long; inflorescences loosely capitate and ca
1 cm broad or umbellate with simple rays up to 5 cm long.

Type: NORTH DAKOTA: Mercer Co.: About 2.8 mi n of Stan-
ton, possibly on the east facing slopes northwest of the present site of

The Great Basin Naturalist
1 10 JAMES L. REVEAL Vol. XXVII, No. 2

the Olds School, on the north bank of the Knife River, elevation about
1800 feet. 28 Jun 1811. Bradbury s.n. Holotype: PH! Isotype: GH!

Representative Specimens: CANADA: MANITOBA: La Riviere
Dawson s.n. (GH, K, MO, UC, US). SASKATCHEWAN: Bracken,
Campbell 38 (MONTU); Wood Mts., Macoun 12949 (NY). UNITED
STATES: IDAHO; Bonner Co.: Kootenai, 1887, Sandberg et al. 997
(CAS, NY). MONTANA: Big Horn Co.: 23 mi n of Grassland, Ander-
son & Wright s.n. (MONT). Blaine Co.: 9 mi e of Zurich, Booth 57568
(MONT). Carter Co.: 2 mi s of Alzada, Booth 2664 (MONT); Box
Elder Cr.. Rose 395 (MONTU). Custer Co.: 9 mi w of Miles City, Hanna
2559 (MONT). Dawson Co.: Glendive, 1900, Blankenship s.n. (MONT,
MONTU), 1903, Blankinship s.n. (MONTU, RM, UTC); Colgate, 1892,
Sandberg et al. 997 (CAS, DS, GH, MO, NY, RM, UC, US). Fergus
Co.: Big Snowy Mts., Canby s.n. (MO, NY, PH, US); 30 mi ne of Roy
Mackie s.n. (MONT). Garfield Co.: Squaw Cr., Holmgren 2076 (MONT,
NY, UTC); 30 mi e of Jordan, Ripley & Barneby 8243 (NY). Golden
Valley Co.: 2 mi nw of Levina, Booth 55205 (MONT). McCone Co.:
20 mi w of Circle, Booth 57220 (MONT). Meagher Co.: 28 mi w of
Harlowton, Booth s.n. (RM). Musselshell Co.: 10 mi nw of Roundup,
Booth s.n. (MONT). Park Co.: White Beaver Cr., Tweedy s.n. (NY).
Petroleum Co.: Mosby. Booth 57205 (MONT); 10 mi sw of Winnett.
Cole s.n. (MONT); 15 mi ne of Grass Range, Kirsch s.n. (MONT).
Phillips Co.: 14 mi sw of Malta, Atwater s.n. (MONT). Powder River
Co.: 13 mi nw of Broadus, McVaugh 6465 (GH, UC). Roosevelt Co.:
Culbertson, Coey 50 (NEB). Rosebud Co.: Forsyth, DeCock s.n.
(MONT). Sheridan Co.: 1 mi ne of Plenty wood. Booth s.n. (MONT).
Valley Co.: Glasgow, Booth 57622 (MONT). Wheatland Co.: 3 mi nw
of Shawmut, Booth 55305 (MONT). Yellowstone Co.: Billings, 1902,
Blankinship s.n. (MONTU, RM), 1903, Blankinship s.n. (RM, UTC):
Pompey's Pillar Butte, Wright 80 (MONT). NEBRASKA: Dawes Co.:
10 mi n of Crawford, Tolstead 952 (GH, NEB). Sioux Co.: Harrison,
Bates s.n. (GH); Orella, Pool & Folsom s.n. (NEB); Hat Cr. Basin,
Webber s.n. (CAS, NEB, US). NORTH DAKOTA: Billings Co.: Men-
dora, Larsen 184 (GH, MO). Bowman Co.: Rhame, Stevens s.n. (MONT,
NDA, RM, US). Dunn Co.: Killdeer Mts., Stevens & Moir s.n. (NDA).
Golden Valley Co.: Sentinel Butte, Stevens 2583, 2584 (NDA, UC).
Grant Co.: sw of Shields, Bell 203, 1259 (NDA). McLean Co.: Ellbo-
woods, Heidenreich s.n. (NDA, UC). Mercer Co.: near Mannhaven,
Stevens 908 (NDA, UC). McKenzie Co.: North Roosevelt Park, Stevens
s.n. (NDA); s of Williston, Waldron 2360 (NDA). Morton Co.: 6 mi e
of Glen Ullin, Rollins & Munoz 2813 (GH, US, UTC). Slope Co.:
Marmarth, Brenckle & Stevens 39-377 (NDA). Stark Co.: Belfield,
Bergman s.n. (NDA); Dickinson, Stevens 1503 (NDA, UC, US). Wil-
liams Co.: Fort Union, Hayden s.n. (GH, MO, NY). SOUTH DAKOTA:
Fall River Co.: 18 mi w of Edgemont, Porter 6706 (RM); Hot Springs,
Rydberg 971 (NEB, NY, US). Harding Co.: 4 mi s of Redig, Jones
36607 (COLO); Moreau, Visher 130 (RM). Hyde Co.: Peno Hills,
Williams s.n. (US). Jackson Co.: Interior, Brenckle 40-43 (UTC);
Cedar Pass, Palmer 37653 (A, MO, US). Lawrence Co.: Lawrence
County, Over 13783 (US). Meade Co.: Fort Meade, For wood 322 (US).
Mellette Co.: Mellette County, Over 15920 (RM, US). Pennington Co.:
1 mi e of Wall, Mclntoch 1389 (NY); se of Rapid City, Over 15921 (RM,
US). Perkins Co.: Badlands, Goodman 3291 (GH, NY, UC), Hapeman
s.n. (NY, UC, UTC). Shannon Co.: White River Valley, Visher 2183
(NY). Washabaugh Co.: Bear Cr., Over 2326 (COLO). WYOMING:
Campbell Co.: 10 mi nw of Gillette, Pennell 21376 (PH, RM). Converse
Co.: near Bill, Ownbey & Lang 1055 (RM). Crook Co.: Rockyford,
A. Nelson 2215 (GH, MO, NY, RM, US); Sundance, A. Nelson 9600
(DS, GH, RM, US); 10 mi n of Devil's Tower, Seig 19 (RM). Johnson
Co.: 10-12 mi n of Buffalo, Pennell & Schaeffer 24416 (PH). Niobrara

Sept. 5. 1967 notes on eriogonum - hi 111

Co.: Lusk, Osterhout 7862 (RM); Cheyenne River, 7. A. Williams s.n.
(RM). Park Co.: Sage Cr., T. A. Williams s.n. (NY, RM). Sheridan Co.:
Ucross, A. Nelson 9740 (MO, RM, UC) ; e of Sheridan, Rollins 559 (GH,
NY). Weston Co.: Newcastle, A. Nelson 8429 (DS, GH, MO, NEB).
NY, RM. US); 10 mi s of Newcastle, Porter 3401 (GH, RM. UC, US).

Eriogonum pauciflorum is a member of the section Capitata
Torr. & Gray of the subgenus Eucycla (Nutt.) Kuntze in Post &
Kuntze, and will be selected as the type of the section in a forth-
coming paper on the subgenera of Eriogonum (Reveal, in press).
The variation in E. pauciflorum is excessive, and the exact nature
of the overall variation is still not entirely surveyed. Within var.
pauciflorum, the degree of tomentum on the leaves is variable,
but never is it as dense as in var. gnaphalodes. Although in the
vicinity of Newcastle, Weston Co., Wyoming, the upper leaf sur-
faces are often totally glabrous, throughout most of the range of
this taxon, the leaves are subglabrous to sparsely tomentose above.
The inflorescence of var. pauciflorum varies from a fairly tight
capitate cluster of involucres to one in which the peduncles have
elongated and form umbellate heads. This latter condition is scattered
throughout the range of the variety, and does not seem to be of any
taxonomic significance. The species, E. depauperatum Small, was
based on a form of var. pauciflorum with glabrous flowers. How-
ever, as nearby populations have pubescent flowers, and as this is
the only differential character, it seems best not to recognize it as
a distinct taxon.

In 1838. Rafinesque published Eriogonum dioecium. The only
Eriogonum specimen of the Bradbury collection, which was cited
as the type, that could fit the species description is the type collection
of E. pauciflorum. H. W. Rickett of the New York Botanical Garden
(1950; per. comm.) has indicated that possibly some of the Brad-
bury specimens were distributed by William Roscoe of Liverpool
who Bradbury had originally sent his collection, and that some dupli-
cates may have come into the possession of Rafinesque. Rafinesque
states that as early as 1833 he had obtained some Bradbury speci-
mens, although he made no reference as from whom he had received
them Rickett indicated that the Pursh collection which was originally
part of the Lambert Herbarium in England, was not sold until 1842,
and as this collection did not come to Philadelphia until some years
after that, it seems impossible that Rafinesque could have seen the
holotype of E. pauciflorum. He might have seen another duplicate,
but this I have been unable to locate. The Rafinesque collection was
sold after his death to Elias M. Durand, then the botanist of the
Academy of Natural Sciences in Philadelphia, but whether or not
Durand discarded the collection as he did with so much of the
Rafinesque collection, or kept it and is now deposited in Durand's
personal collection which he gave to the Museum National d'His-
torie Naturelle in Paris, France, has not been determined. The
name, E. dioecium, is therefore questionably referred to E. pauci-
florum until the type carl be seen.

The Great Basin Naturalist
112 JAMES L. REVEAL Vol. XXVII, No. 2

The one problem in referring Eriogonum dioecium to E. pauci-
florum is the supposed location of the collection site given by Ra-
finesque. Bradbury collected only two Eriogonum species, and the
only two known to grow in the area, E. flavum Nutt. in Fras. and
E. pauciflorum. Both came from near the Minetaree Villages. Ha-
finesque gave the location of his specimen as coming from the Platte
River, but as Bradbury usually only gave the location as the "Up-
per Louisiana" on his specimens, it may have been that Rafinesque
was simply guessing a more exact locality in order to be more spe-
cific.

The transfer of the concept of the name Eriogonum pauciflorum
away from the type to that of a plant in Wyoming and Ck)lorado
may have been due to John Torrey's misidentification of an Engel-
mann collection of Albany Co., Wyoming, that was made in 1856,
and a Parry collection from Middle Park, Q)lorado made a few
years later. When Pursh described the Bradbury collection, he did
not indicate that the flowers were pubescent, and as the collection
was of a series of immature specimens, it is possible that Pursh never
knew of this condition. Pursh must have hoarded nearly the entire
collection as according to Rickett (1950), there are no specimens in
European herbaria, and the only collection in the United States was
deposited in the Pursh Herbarium in Phila<lelphia. Later, a single
plant specimen was cut from the Philadelphia sheet and sent to
Asa Gray at Harvard University by Durand in 1866. It was on this
specimen in the Gray Herbarium that Torrey wrote that the Brad-
bury, Engelmann, and the Parry collections were all the same, and
thus the concept of E. pauciflorum was transferred. This interpre-
tation which appeared in the lorrey & Gray 1870 revision has been
followed since.

lb. Eriogonum pauciflorum var. gnaphalodes (Benth. in Hook.)
Reveal, stat. & comb. nov.

E. gnaphalodes Benth. in Hook., Journ. Bot. & Kew Misc. 5:
263. 1853.

Low-spreading leafy- stemmed, densely matted, perennial herbs;
leaves narrowly spathulate to elliptic, the leaf-blades 1-2.5 cm long,
4-10 mm wide, densely white- tomentose on both surfaces with
long tangled and matted hairs, the leaf-blades usually abruptly
tapering to petioles 1-3 cm long; flowering stems 3-10 (15) cm
long; inforescences densely capitate, mostly 1-1.5 cm broad.

Type: WYOMING: Platte Co.: near Guernsey on cliffs which
are west of the type locality of Fort Laramie, Goshen Co.. which was
cited by the collector, Jul 1843, Geyer 150. Holotype: K! Isotype: K!

Representative Specimens: NEBRASKA: Without definite locality:
Between Pole Cr. and Cedar Bluffs, Engelmann s.n. (GH, MO, NY).
Cheyenne Co.: Bayardito, Baker s.n. (MO); Courthouse Rock, Rydberg
334 (NY, US). Morrill Co.: Chimney Rock, Hapeman s.n. (CAS, MO.
NY, RM, UC). Scotts Bluff Co.: Scotts Bluff, Rydberg 334 (NY);
WYOMING: Converse Co.: Bed Tick Cr., E. Nelson 504O (MONT, RM).

Sept. 5, 1967 notes on eriogonum - in 113

Big Horn Co.: Trapper Basin, Finley 31 (RM); Clark, Pearson 224
(RM). Johnson Co.: Buffalo, Tweedy 3265 (NY, RM). Platte Co.: Lake
Guernsey State Park, Porter 3989 (DS, RM), Porter 4317 (GH, RM,
UC, US). Washakie Co.: Spring Cr.. s of Ten Sleep, Gooding 354 (DS,
GH, NEB, NY, RM, UC, US).

The var. gnaphalodes may be distinguished by its greater reduc-
tion and compaction of the mat, and by its leaves which are wider
and more densely tomentose than in var. pauciflorum. The inflore-
scence is correspondingly more compacted and the involucres are
always tightly held together and not at all umbellate as in var.
pauciflorum. As the var. gnaphalodes tends to occur on the western
and southern edge of the distribution of the species, and is rather
distinctive in the field it is now recognized as a variety.

The placement of Eriogonum gnaphalodes under E. multiceps
by Torrey & Gray (1870) must have come without a chance to see
the type of E. multiceps, or additional material from the upper
Missouri River basin where the type of E. multiceps was collected.
As the Geyer collection and the Maximilian collection are similar in
several general features, Torrey & Gray must have been working
only with the description of E. multiceps which compares somewhat
favorably with the type of E. gnaphalodes which Gray had seen in
the Bentham collection in England.

Ic. Eriogonum pauciflorum var. nebraskense (Rydb.) Reveal, stat.
& comb. nov.

E. nebraskense Rydb., Fl. Rocky Mts. 224, 1061. 1917.

Low spreading, leafy-stemmed, perennial herbs, 2-3 dm high;
leaves oblanceolate, the leaf-blades 1.5-3.5 cm long, 3-5 mm wide,
densely white-tomentose below, grayish-tomentose above, gradually
tapering to short petioles 0.5-1.5 (2) cm long; flowering stems 1-2 dm
long; inflorescences compoundly branched, cymose; perianth rose to
dark red or brown.

Type: NEBRASKA: Kimball Co.: prairies in Kimball County,
12 Aug 1891, Rydberg 337. Holotype: NY! Isotype: US!

Representative Specimens: COLORADO: Hugo Co.: Pawnee Buttes,
Johnston 111 (RM). Weld Co.: Keota, Osterhout 5928 (RM, UC), Oster-
hout 6037 (RM). NEBRASKA: Scotts Bluff Co.: Scotts Bluff County,
Winter 93 (NEB).

The var. nebraskense is a narrowly restricted population that
is readily distinguished from the rest of the section Capitata by its
compound cymose inflorescence.

The reduction of Eriogonum nebraskense to a variety under E.
pauciflorum comes somewhat as a reluctant move. The only dis-
tinguishing characteristic that separates the umbellate forms of var.
pauciflorum from var. nebrasken'ie is the compound cymose inflor-
escence. However, in reviewing the associated specices in the area
where var. nebraskense occurs in part, it is possible that var. nebras-

The Great Basin Naturalist
114 JAMES L. REVEAL Vol. XXVII, No. 2

kense may represent a population that has been genetically influ-
enced by E. effusum Nutt. so that the distinct inflorescence was
introduced. Until detailed field work can be carried out, I am follow-
ing what the gross morphology dictates, and that is to treat E. nebras-
kense as a variety of E. pauciflorum.

Id. Eriogonum pauciflorum var. canum (S. Stokes) Reveal, stat.
& comb. nov.

E. multiceps Nees ssp. canum S. Stokes, Gen. Ej-iog. 94. 1936.

I_iOw spreading, leafy-stemmed, perennial herbs, 1.5-2.5 dm high;
leaves narrowly oblanceolate to lanceolate, the leaf-blades 1.5-3 cm
long, 2-4 mm wide, densely white-tomentose on both surfaces, grad-
ually tapering to short petioles 0.5-1.5 cm long; flowering stems up
to 1 dm long; inflorescences compoundly branched, cymose; perianth
yellow.

Type: MONTANA: Treasure Co.: Custer, 6 Jul 1890, Blank-
inship 113. Holotype: UC! Isotype: US! Stokes cites the date as 30
Jun 1890, but as no sheet at UC with this date has been found, it is
assumed that she made an error in citing the date of the collection.

Representative Specimens: MONTANA: Without definite location:
Montana, 1873, Allen s.n. (US). Treasure Co.: Custer, Brenckle &
Stevens 39-371 (A, GH, NDA).

The var. canum differs from var. nebraskense in its yellow flow-
ers and distinct distribution.

Like var. nebraskense, the var. canum tends to suggest the effects
of possible hybridization, but this time with Eriogonum brevicaule
Nutt. However, as E. brevicaule is not known to be from this area,
considerable field work is necessary on this taxon, and the present
taxonomic arrangement merely reflects the morphological relation-
ships.

2. Eriogonum exilifolium Reveal, spec. nov.

E. pauciflorum of authors, not Pursh.

A Eriogonum coloradense Small foliis linearibus vel lineari-
oblanceolatis, revolutis, inflorescentiis pro parte maxinis compaotis
cymosis, bracteis membranaceis in scapis 3-10 cm longis, involucris
siaepe glabris, perianthiis 2-3.5 mm longis differt.

Planta pulvinata perennis; caules basi per 1.5 cm foliosi; laminae
foliorum lineares vel lineari-oblanceolatae. revolutae, (2) 3-5 (6)
cm longae, 1-2 (3) mm latae, subtus dense albo-tomentosae, supra
subglabrae vel glabrae, petiolis brevibus, 5-15 mm longis, basi mem-
brancea, 5-10 mm longa et 3-5 mm lata, glabris vel pubescentibus;
scapi 3-10 cm longi, glabri vel tomentosi; inflorescentiae cymosae
subcapitatae, brevibus radiis 1-4 mm longis, vel capitatae involucris
3-6 dense compactis, bracteis ternatis, semifoliaceis, membranaceis,
glabris vel parce tomentosis, 3-5 mm longis, 2-3 latis; involucra

Sept. 5. 1967 notes on eriogonum 115

campanulata, 2.5-3.5 (4.5) mm longa, 2-3 mm lata, 5-lobata; per-
ianthia alba vel rosea, 2-3.5 mm longa, segmentis similibus, ellipti-
cis; stamina 2-2.5 mm longa, filamentis basi pilosis, antheris 0.3
mm longis, oblongis; achaenia brunnea, 2-3.5 mm longa, ovata.*

Perennial herbs with strong taproots and branched woody under-
ground caudices up to 8 cm long, forming densely pulvinate mats
up to 2 dm across, sparsely tomentose to glabrous througout; leaves
numerous, linear to linear-oblanceolate, tightly revolute, (2) 3-5
(6) cm long, 1-2 (3) mm wide, densely white-tomentose below,
less so to glabrous and green above, basal or sheathing up the stems
1.5 cm. the apices acute, the bases long-cuneate, gradually tapering
to short petioles 5-10 mm long, the petiole-bases membranaceous,
light brown to tan, expanded and elongated, 5-10 mm long, 3-5 mm
wide, glabrous without, pubescent with long dense white hairs within
and along the margins; scapes 3-10 cm long, glabrous or sparsely
tomentose. ± wirey; inflorescences compact cymose-umbels of 3-7
subcapitate involucres with the short stout rays 1-4 nmi long, the
inflorescences subtended by 3 semifoliaceous membranaceous,
glabrous or sparsely pubescent at the bases and within, triangular
bracts, 3-5 mm long, 2-3 mm wide at the bases, connate; involucres
campanulate, 2.5-3.5 (4.5) mm long, 2-3 mm wide, glabrous vsdth-
out except for small patches of cottony tomentum at the base of
the lobes in some. 5-lobed, the lobes 1-1.5 mm long, rounded, trun-
cate or triangular, distinctly margined with thin membranaceous
margins, erect or more often spreading and reflexed; perianth white
to rose, 2-3.5 mm long, glabrous within and without except for a
few scattered hairs on the lower half within on some, the segments
nearly equal, oblanceolate to elliptic, connate 14 to 1/3 the length
of the perianth, the perianth tubes dark red to brown, occasionally
angled; stamens exserted. 3-4 mm long, the filaments pilose at the
bases, anthers 0.3 mm long, oblong; achenes brown, 2-3.5 mm long,
ovate, the globose bases tapering to 3-angled beaks.

Type: WYOMING: Albany Co.: 2 mi n of Laramie, 21 Jul
1945, C. L. Porter 3706. Holotype: RM! Isotypes: DS, GH, MO, NY,
UC, US.

Representative Specimens: COLORADO: Grant Co.: Middle Park,
Parry s.n. (GH, NEB, NY, US); nw of Tabernash, Ripley & Barneby
10489a (CAS, NY). Jackson Co.: North Park, Crandall 133 (GH, MO,
NY); Lake John, Ramaley & Johnson 890 (CAS, COLO, MO). WYOM-
ING: Albany Co.: From Laramie River along the Medicine Bow Mts.,
Engelmann s.n. (GH, MO, NY); Red Buttes, Greene s.n. (NY); Laramie
Plains, A. Nelson 2794 (GH. MONT, NY, RM, US); Laramie, A. Nelson
7637 (COLO, GH, NEB, NDA. NY); Laramie River, E. Nelson
3385 (BM, MO, NY. RM, US). Larimer Co.: North Fork, Goodding
1924 (COLO, NY. UC, US).

This taxon has long been recognized as an excellent species,
and the naming of it as such after so many years is simply due to the
misapphcation of the name Eriogonum pauciflorum for it. The new
species is closely related to E. coloradense from which it differs

■•The author is grateful to C. V. Morton for his assistance with the I.atiii description.

The Great Basin Naturalist
116 JAMES L. REVEAL Vol. XXVII, No. 2

mainly in its leaf-shape and general size of its parts. The distinct
ecological requirements also differ. The plants in Wyoming are
found mainly on hard gumbo clay hills, but in Q)lorado, they are
often found on rolling granitic sand hills below 8,500 feet eleva-
tion.

The species name is selected to denote the narrow leaves.

3. Eriogonum coloradense Small, Bull. Torrey Bot. Club 33: 53.
1906.

E. multiceps Nees ssp. coloradense S. Stokes, Gen. Eriog.
94. 1936.

Perennial herbs with strong taproots and branched woody under-
ground caudices up to 4 cm long, forming loose pulvinate mats up
to 1.5 dm across, tomentose or glabrous almost throughout; leaves
few to many, oblanceolate to lanceolate or narrowly spathulate. not
revolute but the margins sometimes thicker than the blades or cris-
pate, 1-4 (5) cm long. 3-6 (8) mm wide, densely white-tomentose
below, lightly floccose to glabrous and green above, basal or sheath-
ing up the stems 1 cm. the apices acute to rounded, the bases long-
cuneate, gradually tapering to short petioles 2-8 mm long, the petiole-
bases membranaceous, light tan, expanded and elongated. 2-4 mm
long, 1-1.5 (2) mm wide, sparsely to densely tomentose without,
densely tomentose within; scapes 3-5 (6) cm long, glabrous or den-
sely matted tomentose; inflorescences capitate or nearly so, 3-4
involucres on short rays up to 1 mm long, subtended by 3 semi-
foliaceous membranaceous, glabrous or tomentose. triangular bracts,
1.5-3 (4) mm long, 1-1.5 mm wide at the bases, connate; involucres
turbinate-campanulate. 2-3 mm long. 1.5-2.5 (3) mm wdde, glabrous
without except for small patches of cottony tomentum at the base
of each lobe, 5-lobed, the lobes 0.3-0.7 (1) mm long, mostly acute
and triangular, margins membranaceous, erect or reflexed; perianth
white to rose, 2.5-3.5 mm long, glabrous within and without, the
segments nearly equal, oblong to ovate, connate about 1/4 of the
length of the perianth, the perianth tube dark brown to red-brown,
occasionally angled; stamens exserted, 3.5-4 mm long, the filaments
pilose at the bases, the anthers 0.3 mm long, oblong; achenes brown,
2.5-3.5 mm long, ovate, the globose bases tapering to 3-angled beaks.

Type: COLORADO: Chaffee Co.: Mt. Harvard, 1896. F. E.
Clements 66. Holotype: NY!

Representative Specimens: COLORADO: El Paso Co.: Pikes Peak,
Snow s.n. (PH). Gunnison Co.: Virginia Basin, Langenheim 285 (UC).
Langenheim 1062 (COLO); nw of Castle Peak, Ewan 11750, 11757
(COLO); North Pole Basin, Weber & Barclay 9184 (COLO); 2 mi ne
of Gothic, Wherry s.n. (GH). Park Co.: South Park, Canby s.n. (GH,
NY, PH); 2 mi sw of Glentivar, litis & litis 18830 (COLO, WIS); 6 mi
n of Fairplay, Penland 1320 (CAS); between Red Hill Pass and Como,
Weber 8758 (COLO); Sulphur Spr., South Park, Wolf & Rothrock 27
(DS, PH, US). Saguache Co.: Cochetopa Pass, Weber 5795 (COLO),
Weber 9419 (DS); 9 mi se of the Flying-M- Ranch, Wherry s.n. (A).
Eriogonum coloradense is a high alpine species that is most

Sept. 5, 1967 notes on eriogonum 117

closely related to E. exilifolium, which is found to the north and at
a much lower elevation.

Bibliography

Brack ENRiDGE, H. M. 1814. Views of Louisiana; together with a journal of

a voyage up the Missouri River in 1811. Pittsburgh.
Bradbury, J. 1817. Travels in the interior of America, in the years 1809,

1810, 1811 ... Liverpool.
Geyer, C. a. 1845-1846. Notes on the vegetation and general character of the

Missouri and Oregon territories . . . London Joum. Bot. 4: 479-492, 653-662.

1845; 5: 22-41, 198-208, 285-310, 509-524. 1846.
Graustein, J. E. 1950. Nuttall's travels in the Old Northwest. An unpublished

1810 diar>'. Chronica Botanica 14: no. V2- 86 pp.
Irving, W. 1836. Astoria; or, anecdotes of an enterprise beyond the Rocky

Mountains. 2 vols. Philadelphia.
Lewis, M. & W. Cl-^rk. 1814. History of the expedition under the command

of Captains Lewis and Clark . . . Prepared for press by Paul Allen, Esquire;

edited by Nicholas Biddle. 2 vcJs. Philadelphia.
Maximiuan, a. p. 1841. Reise in das innere Nord-America in den Jahren

1832 bis 1834. 2 vols. Coblenz.
McKelvey, S. D. 1955. Botanical exploration of the Trans- Mississippi west,

1790-1850. Jamaica Plains, Massachusetts.
OoLESBY, R. E. 1963. Manuel Lisa and the opening of the Missouri fur Irade.

Norman, Oklahoma.
Pennell, F. W. 1936. Travels and scientific collections of Thomas Nuttall.

Bartonia 18: 1-51.
Rafinesque, C. S. 1833. Account of the botanical collections of Professor C.

S. Rafinesque. Atl. Joum. & Friend of Knowl. 167-170.
Reveal, J. L. In press The subgeneric concept in Eriogonum (Polygonaceae).

Bull. Torrey Bot. Club.
RiCKETT, H. W. 1950. John Bradbury's explorations in Missouri Territory.

Proc. Am. Philos. Soc. 94: 59-89.
Stevens, O. A. 1946. Botanical visits to forts Clark, Mandan and Union in

North Dakota. Rhodora 48: 98-103.
. 1959. Bradbury and Nuttall, pioneer Dakota botanists. North Dakota

Hist. Quart. 26:159-169.
Torrey, J. a A. Gray. 1870. A revision of the Eriogoneae. Proc. Amer. Acad.

8:145-200.

The Gireat Basin Naturalist

Founded in 1939 by Vasco M, Tanner

A journal published from one to four times a year by Brigham
Young University, Provo, Utah.

Manuscripts: Only original, unpublished manuscripts, pertain-
ing to the Great Basin and the western United States in the main,
will be accepted. Manuscripts are subject to the approval of the
editor.

Illustrations: All illustrations should be made with a view to
having them appear within the limits of the printed page. The illus-
trations that form a part of an article should accompany the manu-
script. All half-tones or zinc etchings to appear in this journal are
to be made under the supervision of the editor, and the cost of the
cuts is to be borne by the contributor.

Reprints: No reprints are furnished free of charge. A price list
for reprints and an order form is sent with the proof.

Subscription: The annual subscriptian is $2.50 (outside the
United States $3.25). Single number, 80 cents.

All correspondence dealing with manuscripts should be addressed
to the Editor, Vasco M. Tanner, Great Basin Naturalist, Brigham
Young University, Provo, Utah. Other matters such as subscriptions,
reprints, exchanges and other business should be addressed to Ernest
L. Olson, Chairman of University Publications.

Reprints Schedule of The Great Basin Naturalist

Each
Additional

8 pp. 10 pp. 12 pp. 2 pp.

$9.00 $10.00 $11.00 $2.00
10.00 11.00 12.00
11.00 12.00 13.00
300 copies 9.00 10.00 11.00 12.00 13.00 14.00

Covers: $10.00 for first 100 copies, $4.00 for additional 100 copies.

2 pp.

4 pp.

6 pp.

50 copies

$6.00

$7.00

$8.00

100 copies

7.00

8.00

9.00

200 copies

8.00

9.00

10.00

TABLE OF CONTENTS

A Comparative Study of the Mountain Brush Vegetation
in Utah. Illustrated. Elray S. Nixon 59

Predaceous-Scavenger Ants in Utah. D Elden Beck, Dorald
M. Allred, and William J. Despain 67

New Records and Species of Neotropical Bark Beetles (Scoly-
tidae: Coleoptera). Stephen L. Wood — 79

A New Species of Spinihdella from Utah (Bdellidae: Aca-
rina). Illustrated. Clive D. Jorgensen 98

Notes on Eriogonum - III on the Status of Eriogonum pauci-

florum Pursh. Illustrated. James L. Reveal 102

Volume XXVn, No, 3
November 30, 1967

L/3RARY

The

W

Great Basin

PUBUSHED BY

Brigham Young UNivERsmr

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Zoolc»y and Elntomology

^ooic»y
, Utah

Brigham Young University, Prove

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:
J. V. Beck, Bacteriology

WiLMER W. Tanner, Zoology, Chairman of the Board
W. Derby Laws, Agronomy
Stanley Welsh, Botany
C. Lynn Hayward, Zoology
Howard C. Stutz, Botany

Ex officio Members

RUDGER H,

cultur
EIrnest L. Olson, Chairman, University Publications

RuDGER H. Walker, Dean, College of Biological and Agri-
cultural Sciences

^ / (S fc>, 2S LIBRARY

MAR I 2 1963

HARVARD

The Great Basin Naturalisf ^'^^

Published at Provo, Utah by
Brigham Young University

Volume XXVII November 30, 1967 No. 3

NEW RECORDS AND SPECIES OF NEOTROPICAL BARK
BEETLES (SCOLYTIDAE: COLEOPTERA), PART II '

Stephen L. Wood-
While preparing a taxonomic review of the bark and ambrosia
beetles (Scolytidae) of Costa Rica it was necessary to examine all
available species of this group from Mexico, all of Central America,
and northern South America. As a result, a large number of species
new to science were recognized from areas outside of Costa Rica. In
order to make names available for these species to facilitate this and
other work. 23 species and one genus (Stegomerus) new to science
are described below. The new species represent the genera Loganius
(1), Scolytus (1). Scolytodes (10), Stegomerus (5), Xyleborus (3),
Corthylus (1 ). and Tricolus (2). The type series of these species were
collected in the following countries: Mexico (6), Guatemala (6),
Honduras (3). Costa Rica (8). Panama (8), Colombia (2), and
Bolivia ( 1 ) . One species occurred in two countries and five were col-
lected in three countries.

Loganius exilis, n. sp.

This species is rather closely related to liratus Wood, but it is dis-
tinguished by the less strongly serrate costal margins of the elytral
declivity, by the more strongly elevated alternate interstriae on the
elytral declivity, by the poorly developed funicular tuft of hair, and
by the somewhat narrower frons.

Male. — Length 1.4 mm. (paratypes 1.3-1.6 mm.). 2.7 times as
long as wide; color very dark brown.

Frons flattened or very feebly concave on lower half, convex
above; surface rather coarsely, closely, deeply punctured in im-
pressed area, more sparsely punctured above; median third of epis-
tomal margin bearing a shining, elevated, procurved carina as in
liratus; vestiture limited to impressed area, consisting of abundant,
erect, stout, uniformly rather short setae.

1. Part of the field work relating to tins study was sponsored by rcsearcli grants GB-5?2 and
GB-5()78 from the National Science Foundation.

2. Department of Zoology and Entomology, Brigliam Young University, Provo, Utah. Scolytoidea
Contribution No. 35.

119

The Great Basin Naturalist
120 STEPHEN L. WOOD Vol. XXVII, No. 3

Pronolum equal in width and length; widest near base, the sides
weakly arcuate and converging slightly toward the rather narrowly
rounded anterior margin; surface smooth, shining, the punctures
rather large, deep, moderately close, oval; glabrous.

Elytra 1.8 times as long as wide. 1.7 times as long as pronotum;
sides straight and parallel on basal two-thirds, narrowly, subserrately
rounded behind; striae weakly impressed, the punctures moderately
large, deep; interstriae shining, slightly wider than striae, very weak-
ly convex, the punctures uniseriate, small, widely spaced. Declivity
rather steep, convex, the punctures uniseriate, small, widely spaced.
Declivity rather steep, convex; striae narrower than on disc; inter-
striae subshining, 1, 3, 7. and 9 moderately elevated, 7 joins 9 and
continues almost to 3, elevated interspaces and 2 (somewhat less)
uniseriately granulate, the granules moderately large, rounded,
rather widely spaced; costal margin subserrate toward apex. Vesti-
ture restricted to declivity, sparse; consisting of rows of erect, inter-
strial scales, each scale about five times as long as wide.

Female. — Similar to male except frons less strongly impressed
below, more broadly rounded above.

Type Locality. — Volcan de Agua, Guatemala.

Type Material. — The male holotype. female allotype, and 81
paratypes were collected at the type locality on May 19, 1964, at
an elevation of 1,000 m., by S. L. Wood, from the bole and branches
of a common small tree that awaits identification.

The holotype, allotype, and })aratypes are in my collection.

Scolytus aztecus, n. sp.

This species appears to be allied to ventralis Leconte, but it is
easily recognized by the very much larger size, by the two pair of
projecting lobes at elytral apex; by the broad, recurved, epistomal
elevation, and by other characters. This is the largest American
representative of the genus.

Adult. — Presumably a female. length 5.7 mm. (})aratypes 4.7-
6.3 mm.), 2 A times as long as wide; color very dark brown, elytra
reddish brown.

Frons convex, transversely impressed just above ej)istoma, a
slight median impression at vertex; surface shining, the punctures
small, rather close, substrigose above; epistoma with a distinctly ele-
vated, dorsally arched ridge above epistomal brush, its lateral ex-
tremities blending into a marginal callus; vestiture short, sparse, in-
conspicuous.

Pronotum 1.03 times as long as wide; widest just behind middle,
the sides very weakly arcuate, not converging until anterior third,
rather strongly constricted at sides just before anterior margin, an-
terior margin boardly, very feebly emarginate; surface smooth,
shining, the punctures fine, deep on disc, about twice as large in
lateral areas; vestiture confined to lateral areas.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 121

Elytra 1.4 times as long as wide, 1.4 times as long as pronotum;
sides almost straight and parallel on basal two thirds, then converg-
ing very slightly to tlie rather abrupt posterolateral angles; median
half of j)osteri()r outline occupied by two pairs of broad serrations,
each serration half as long as wide; striae not im[)ressed. the punc-
tures small, deep, regular; interstriae about three to four times as
wide as striae, almost flat, shining, the ])unctures moderately large,
deep, uniseriate except confused on 2; declivity descending slightly;
glabrous, except a few hairs on declivity. Venter as in female ven-
tralis. except more coarsely punctured, the feeble, carinate elevation
at posterior margin of sternum 2 present on all four specimens.

Male. — Not represented in the series at hand.

Type Locality. — Twenty-seven miles east of Morelia, Michoa-
can, Mexico.

Host. — Abies religiosa.

Type Material. — The female holotype and three female para-
types were taken at the type locality on June 14. 1965, at an eleva-
tion of about 2.600 m.. by S. L. Wood, from biramous. transverse egg
tunnels in the bole of a standing fir tree about 60 cm. in diameter.

The holotype and paratypes are in my collection.

Scolytodes clusiacolens, n. sp.

This species is more closely allied to clusiae Wood than to other
known species, but it may be distinguished by the larger size, by
the coarsely, deeply punctured pronotum and elytra and. in the
female, by the weakly convex frons.

Female. — Length 2.3 mm. (paratypes 1.8-2.9 mm.), 2.5 times
as long as wide; color black.

Frons plano-convex from a point well below upper level of eyes
to epistoma. convex above that point, the transition rather abrupt;
surface reticidate. finely, deeply, rather closely punctured; vestiture
inconspicuous, consisting of very fine, moderately abundant, uni-
formly distributed hair.

Pronotum 1.1 times as long as wide; sides almost straight or very
feebly constricted on basal two-thirds, broadly rounded in front;
dorsal profile weakly arched from base, more strongly declivous on
anterior fifth; surface reticulate, with rather coarse deep punctures
from base to near anterior margin, gradually replaced by smaller
punctures laterally and minute asperities medially on anterior fifth;
glabrous except near lateral and anterior margins.

Elytra 1.5 times as long as wide. 1.5 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather narrow-
ly rounded behind; striae impressed, the punctures rather large,
deep; interstriae slightly narrower than striae, convex, shining, the
punctures fine, sharply but not deeply impressed, uniseriate. some of
them slightly subgranulate on anterior margins (variable in the
series). Declivity convex, rather steep; striae 1 only distinctly im-
pressed, the punctures distinctly .smaller than on disc; interstrial

The Great Basin Naturalist
122 STEPHEN L. WOOD Vol. XXVII, No. 3

punctures minute, clearly impressed. Vestiture fine, hairlike, abraded
on disc of type, in regular strial and interstrial rows on teneral speci-
mens.

Male. — Similar to female except frons convex to transverse im-
pression just above epistoma, the i)unctures larger, deeper, the vesti-
ture almost confined to epistomal area; more nearly subgranulate
interstrial punctures.

Type Locality. — About 10 km. east of Volcan Paracutin, Mi-
choacan. Mexico.

Host. — CJusia sp.

Type Material. — The female holotype, male allotype and 66
para types were taken at the type locality on June 19, 1965, at an
elevation of about 2,500 m., by S. L. Wood, from Clusia twigs and
branches. Other paratypes include 16 from Volcan Pacaya, Guate-
mala. June 1, 1964, 1,300 m.. from Clusia; 1 from Cerro Peha Blanca,
Honduras, May 23, 1964, 2.000 m., from Clusia.

The holotype, allotype, and paratypes are in my collection.

Scolytodes clusiavorus^ n. sp.

This species is rather closely related to clusiae Wood, but may be
distinguished by the smaller size, by the more sharply, somewhat
more deeply punctured pronotum and elytra, particularly the elytral
declivity and, in the female, by the less deeply, less extensively con-
cave frons.

Female. — Length 1.5 mm. ('j)aratypes 1.3-1.6 mm.), 2.6 times as
long as wide; color black.

Frons shallowly, broadly plano-concave from just below upper
level of eyes to epistoma; surface subshining, closely, finely, deeply,
uniformly punctured from vertex to epistoma; vestiture consisting of
rather coarse, moderately long, uniformly but rather sparsely dis-
tributed hair.

Pronotum 1.1 times as long as wide; sides almost straight or very
feebly constricted on posterior two-thirds, broadly rounded in front;
surface reticulate, the punctures rather fine, deep, moderately close,
the anterior fifth more strongly declivous with a few minute asperi-
ties in median area, the punctures attaining anterior margin only in
lateral areas; vestiture confined to anterior and lateral areas.

Elytra 1.5 times as long as wide. 1.6 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, narrowly
rounded behind; striae not impressed, the junctures moderately
large, dee[); interstriae slightly narrower than striae, almost smooth,
with some irregular lines, the punctures minute, almost uniseriate,
not at all granulate. Declivity convex, rather steep; strial punctures
smaller and shallower than on disc. Vestiture consisting of rows of
minute strial hairs and longer, erect rows of coarser interstrial hair.

Male. — Similar to female exce[)t frons strongly convex, reticu-
late, the punctures rather coarse, s])arse, vestiture restricted to episto-
mal margin.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 123

Type Locality. — Volcan de Agua. Guatemala.

Host. — Clusia sp.

Type Material. — The female holotype. male allotype, and three
paratypes were collected at the type locality on May 19. 1964, at
an elevation of 1.000 he. by S. L. Wood, from the i)hloem of small
branches of a small Clusia tree.

The holotype, allotype, and paratypes are in my collection.

Scolytodes pcrditus, n. sp.

This species is more closely allied to cecropiacolens Wood than to
other known species, but it is easily distinguished by the more slender
body form, by the less deeply impressed, finer punctures on pro-
notum and elytra and, in the female, by the dense, long brush of
frontal hair arising above the eyes.

Female. — Length 2.0 mm. (paratypes 1.6-2.1 mm.), 2.1 times as
long as wide; color yellowish brown, the anterior fourth of prothorax
darker.

Frons flattened from well above upper level of eyes to epistoma,
with low. submarginal, longitudinal carinae extending from epistoma
about half the distance to upper level of eye; central area minutely
granulate, lateral areas with a few punctures; vestiture consisting
of alnmdant. coarse, long, subplumose setae mostly arising above
eyes, none as low as level of antennal bases, some of the setae suf-
ficiently long to reach epistoma.

Pronotum 1.1 times as long as wide; widest at base, sides almost
straight, converging very slightly anteriorly, rather narrowly
rounded in front; slightly less than anterior half very finely, closely
asperate; posterior area reticulate, the punctures rather fine, shallow;
glabrous.

Elytra 1.2 times as long as wdde. 1.2 times as long as pronotum;
sides straight and parallel on slightly more than basal half, rather
narrowly rounded behind; striae not impressed, the punctures small,
sharply but not deeply impressed, the strial punctures almost in-
distinguishable from the similar, abundant, confused, interstrial
punctures; surface almost smooth, shining. Declivity convex, rather
steep; strial punctures deeper than on disc, not reduced; interstrial
punctures distinctly smaller, uniseriate except near upper area.
Vestiture restricted to a few setae on sides.

Male. — Similar to female except frons convex, with a transverse
impression iu«;t above epistoma but interrupted by a small subcari-
nate median elevation; surface smooth, shining, the punctures rather
coarse, deep; strial punctures on disc deeper and somewhat larger
than those of striae.

Type Material. — Fort Clayton, Canal Zone. Panama.

Host. — Cecropia sp.

Type M.^^terial. — The female holotype, male allotype, and 16
paratypes were collected at the type locality on December 22. 1963,

The Great Basin Naturalist
124 STEPHEN L. WOOD Vol. XXVII, No. 3

at an elevation of about 30 m., by S. L. Wood, from the terminals of
Cecropia branches.

The holotype. allotype, and paratypes are in my collection.

Scolytodes nanellus^ n. sp.

This species is somewhat allied to pumilus Wood, but it is much
smaller, much less coarsely punctured, and the female frons is armed
by a pair of submarginal. longitudinal carinae below and by a brush
of long hair above upper level of eyes.

Female. — Length 1.0 mm. (female para type 1.0 mm.; males
0.9 mm.), 2.4 times as long as wide; color medium brown.

Frons flattened from upper level of eyes to epistoma, with sub-
marginal, low carina extending from epistoma almost to upper level
of eyes; surface subshining, very finely and closely punctured; vesti-
ture consisting of rather abundant, coarse, subplumose setae, con-
fined to area near or above upper level of eyes, some of setae almost
reaching epistomal margin.

Pronotum 1.1 times as long as wide; widest just behind middle,
the sides feebly arcuate, rather broadly rounded in front; summit
distinctly in front of middle, very finely asperate in front, posterior
area rather coarsely reticulate, the punctures rather fine, not deep;
glabrous.

Elytra 1.4 times as long as wide, 1.3 times as long as pronotum;
sides almost straight and parallel on slightly more than basal half,
rather narrowly rounded behind; striae 1 weakly, others not im-
pressed, the punctures small, not deep; interstriae slightly wider than
striae, shining, almost smooth, the punctures fine, not always clearly
impressed. Declivity convex, rather steep; all punctures somewhat
reduced. Vestiture consisting of a few scattered bristles on or near
declivity.

Male. — Similar to female except frons convex, with a slight
transverse impression just above epistoma, surface subreticulate, the
minute punctures obscure, vestiture reduced, confined to epistomal
area.

Type Locality. — Barro Colorado Island. Canal Zone. Panama.

Type Material.- — The female holotype, male allotype, and two
paratypes were collected on December 27, 1963, at an elevation of
about 70 m.. by S. L. Wood, from phloem tissues of an unidentified
tree branch.

The holotype, allotype, and paratypes are in my collection.

Scolytodes venustulus, n. sp.

This species apjiears to be closely related to venustus Wood, but
is distinguished by the smaller size, by the reticulate, less closely
punctured posterior area of the pronotum, by the unimpressed elytral
striae and by the almost equal size of strial and interstrial punctures.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 125

Male. — Length 1.5 mm. (male para types 1.3-1.5 mm.), 2.3
times as long as wide; color dark brown, with a slight reddish cast.

Frons convex, with a transverse im[)ression just above epistoma;
surface smooth and shining to a level above eyes, the punctures very
fine; vestiture fine, hairlike, largely confined to epistomal area.

Pronotum 1.0 times as long as wide; widest just behind middle,
the sides rather weakly arcuate on basal two-thirds, rather narrowly
rounded in front; dorsal profile arched from base, a little more
strongly declivous on anterior fourth; surface reticulate, with rather
coarse, deep, moderately close punctures on basal two-thirds, finely,
closely asperate in median area in front, the punctures reaching
anterior margin in lateral areas; punctures on disc separated by
distances about equal to their own diameters; glabrous.

Elytra 1.5 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and y)arallel on slightly more than basal half,
rather narrowly rounded behind; base of suture just behind scutel-
lum acutely but not strongly elevated; striae not impressed, the punc-
tures moderately large, deep; interstriae slightly wider than striae,
smooth and shining, the punctures fine, distinct. Declivity convex,
rather steep; strial punctures reduced; interspace 2 narrow, not wider
than diameter of a strial puncture (noticeably wider on one para-
type j. Vestiture consisting of minute strial and interstrial hairs, and
longer, erect, bristles on declivital interspaces 1, 3, 5, and 7. Inter-
space 10 ending before level of hind coxa.

Female. — Not represented in the series at hand.

Type Locality. — Cerro Punta on Volcan Baru (Chiriqui), Pan-
ama.

Host. — Oreopanax sp.

Type Material. — The male holotype and three male paratypes
were collected at the type locality on January 11. 1964, at an eleva-
tion of about 1,800 m.. by S. L. Wood, from phloem of a young
Oreopanax sapling about 5 cm. in diameter.

The holotype and paratypes are in my collection.

Scolytodes ficivorus, n. sp.

This species is very closely related to schwurzi (Hopkins), but it
may be distinguished by the more coarsely punctured, less shining
])ronolum and elytra, by the reticulate pronotal disc, by the more
finely asperate anterior area of the pronotum. and by the somewhat
more coarsely punctured marginal areas of the female frons. Addi-
tional data eventually may prove this to be only a subspecies of
schwarzi.

Female. — Length 16 mm. (paratypes 1.5-1.7 mm.). 2.3 times
as long as wide; color yellowish brown.

Except for the characters noted in the above diagnosis this species
appears to be identical to schwarzi.

Male. — Similar to female except frons convex, reticulate,
obscurely punctured, with frontal vestiture greatly reduced.

The Great Basin Naturalist
126 STEPHEN L. WOOD Vol. XXVII, No. 3

Type Locality. — Palin, Escuintla, Guatemala.

Host. — Ficus sp.

Type Material. — The female holotype, male allotype, and two
male para types were collected at the type locality on May 19, 1964,
at an elevation of about 300 m., by S. L. Wood, from a small, broken
fig branch.

The holotype, allotype, and paratypes are in my collection.

Scolytodes facetus^ n. sp.

This species is very closely related to ficivorus Wood, described
above, but is distinguished by the very fine, somewhat obscure punc-
tures of pronotum and elytra, by the slightly longer and more abun-
dant vestiture on the female frons, and by the more narrowly
separated eyes. Except for the fact that the two type series of this
species and ficivorus were taken with two other species of Scolytodes
from the same small branch, the differences are of such a nature
that they might possibly be considered geographical replacements
of one another.

Female. — Length 1.6 mm. (male paratypes 1.4-1.5 mm.), 2.3
times as long as wide; color yellowish brown.

In addition to the characters noted in the above diagnosis, this
species differs from ficivorus in having the eyes separated by 1.6
times the width of an eye (2.6 times in ficivorus), and the elytra are
reticulate, with the punctures shallow and with those of the inter-
spaces obscure. The obscure, minute pronotal punctures are separated
by distances at least twice their own diameters (this distance less
than the diameter of a puncture in ficivorus) .

Male. — Similar to female except male frons convex, reticulate,
obscurely punctured; eyes separated by 2.1 times the width of an
eye (3.1 in ficivorus).

Type Locality. — Palin, Escuintla, Guatemala.

Host. — Ficus sp.

Type Material. — The female holotype, male allotype, and two
male paratypes were taken at the type locality on May 19, 1964, at
an elevation of about 300 m., by S. L. Wood, from the same small
branch of a native fig tree that contained the type series of ficivorus.

The holotype. allotype, and paratypes are in my collection.

Scolytodes ingavorus. n. sp.

This species is rather closely related to ficivorus Wood, described
above, but it may be distinguished by the pubescent elytra, by the
deeper punctures on the pronotal disc and. in the female, by the
much more heavily pubescent frons that lacks a shining glabrous
area at the center.

Female. — Length 1.6 nmi. (paratypes 1.3-1.7 mm.). 2.3 times
as long as wide; color yellowish brown, with anterior half of pro-
notum darker.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 127

Frons flattened from upper level of eyes to the somewhat elevated
epistoma, with a pair of submarginal, low, longitudinal carinae (or
elevations; obscured by vestiture) extending dorsad from epistomal
margin, its upper limits obscured by vestiture; surface smooth and
shining with fine punctures below, becoming finely granulate and
obscurely punctured in central area above; vestiture consisting of
abundant, long, coarse, subplumose setae above upper level of eyes,
continuing on lateral areas to epistomal margin as somewhat shorter
setae, central area bearing shorter, fine, moderately abundant, hair-
like setae.

Pronotum 1.1 times as long as wide; sides almost straight and
parallel on slightly less than basal two-thirds, rather broadly rounded
in front; summit just in front of middle, the anterior area rather
coarsely asperate, the interior margin armed by an almost continuous
ridge; posterior areas reticulate, the punctures coarse, deep, close,
separated from one another by less than half the diameter of a punc-
ture; scanty hairlike setae confined to sides and asperate area.

Elytra 1.3 times as long as wide, 1.2 times as long as pronotum;
sides almost straight (feebly arcuate) and parallel on slightly less
than basal two-thirds, broadly rounded behind; striae not impressed,
the punctures coarse, distinctly but shallowly impressed; interstriae
slightly wider than striae, the punctures uniseriate, almost identical
to those of striae. Declivity steep, convex; all punctures reduced, in
rows; surface shining. Vestiture consisting of fine, short, erect strial
and interstrial hair, and longer interstrial hair on declivity.

Male. — Similar to female except frons broadly convex, trans-
versely impressed just above epistoma; short vestiture on declivity
stout, almost scalelike.

Type Locality. — Los Corchos, Nayarit, Mexico.

Host. — Inga sp.

Type Material. — The female holotype, male allotype, and 43
paratypes were collected at the type locality on July 10, 1965. at an
elevation of less than 10 m., by S. L. Wood, from the phloem of
Inga branches.

The holotype, allotype, and paratypes are in my collection.

Scolytodes proximus, n. sp.

This species is rather closely related to tenius Wood, but may be
distinguished by the very minute, obscure punctures on the pro-
notal disc, by the finer, shallower, elytral punctures, by the finer
elytral vestiture, and by the less abundant vestiture on the female
frons.

Female. — Length 1.8 mm. (paratypes 1.7-1.9 mm.), 2.8 times
as long as wide; color dark brown.

Frons narrowly plano-concave from below upper level of eyes to
epistoma; surface reticulate, obscurely punctured; vestiture scanty,
consisting of fine, rather sparse, long hair uniformly distributed from
upper level of eyes to epistoma.

The Great Basin Naturalist

128 STEPHEN L. WOOD Vol. XXVII, No. 3

Pronotum 1.1 times as long as wdde; widest slightly in front of
middle, sides feebly constructed on posterior half, rather broadly
rounded in front; summit indefinite, about one-third from anterior
margin; surface reticulate, minutely, sparsely punctured, anterior
third finely asperate; anterior margin armed by a series of low, in-
definite teeth; glabrous except a few setae in asperate area.

Elytra 1.7 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and parallel on slightly more than basal two-
thirds, rather narrowly rounded behind; striae not impressed, the
punctures rather fine, shallow, obscure; interstriae as wide as striae,
somewhat irregular, subshining. the punctures fine, uniseriate, ob-
scure. Declivity convex, steep; the punctures obscure, about as on
disc. Vestiture consisting of minute, recumbent strial and interstrial
hair, and longer, erect, sparse, interstrial hair.

Male. — Similar to female except frons convex, with a transverse
impression above epistoma, the surface reticulate with fine, obscure
punctures.

Type Locality. — Base of Volcan Baru (Chiriqui) near El Hato
del Volcan, Panama.

Host. — Clusia sp.

Type Material. — The female holotype, male allotype, and 31
paratypes were collected at the type locality on January 11, 1964,
at an elevation of about 1,500 m., by S. L. Wood, from the phloem
of twigs and branches of a Clusia tree.

The holotype, allotype, and paratypes are in my collection.

Scolytodes amoenus, n. sp.

Superficially, this species resembles multistriatus Wood, but it is
brightly shining, has the declivity more gradual, the female frons is
entirely different, and the tenth interspace is acutely elevated to the
declivity. Among species with an asperate anterior slope of the pro-
notum, continued tenth elytral interspace, and a tubercle on the
posterior face of the front tibia, this species is unique. The large s^'ze,
the multistriate elytra and the unique female frons also help to dis-
tinguish this unusual species.

Female. — Length 2.0 mm. (paratypes 1.9-2.3 mm.), 2.2 times
as long as wide; color of basal third of pronotum, most of elytra and
most of body yellowish brown, with head, anterior two-thirds of
pronotum, all margins of each elytron and some parts of meso- and
metasternum dark brown (variable in series).

Frons flattened from just above upper level of eyes to epistoma;
most of area between eyes occupied by a very finely granulate, per-
fect circle extending from upper level of eyes to epistoma; lower half
of circle bearing on its lower half a smooth, shining, transverse, pro-
curved band with small, median, orad extension, this area about
equal to half the width of circle; area outs-de of circle rather coarsely,
very closely punctured; vestiture consisting of equally long, fine.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 129

plumose setae on a continuous band surrounding circular area,
numerous additional setae along epistoma; central area glabrous.

Pronotum 1.1 times as long as wide; widest at base, the sides very
feebly arcuate, almost straight, and converging anteriorly very slight-
ly on basal two-thirds, rather narrowly rounded in front; indefinite
summit near middle, anterior fourth finely asperate; anterior margin
armed by a continuous ridge; posterior areas obscurely subreticulate,
coarsely, deeply, closely punctured, the punctures separated by
slightly less than their own diameters; glabrous except near lateral
and anterior margins.

Elytra 1.3 times as long as wide. 1.2 times as long as pronotum;
sides almost straight and parallel on basal half, rather narrowly
rounded behind; elytra with basal margin marked by a fine, raised
line; striae 1 weakly, others not impressed, the punctures rather
fine, deep; interstriae shining, one and one half times as wide as
striae, the punctures rather fine, deep, rather abundant, confused.
Declivity convex, steep; striae 1 strongly impressed; all punctures
reduced, those of striae little larger than the abundant interstrial
punctures. Glabrous except at sides.

M.ALE. — Similar to female except frons convex, with a transverse
impression iust above epistoma. surface reticulate with coarse deep
punctures above, a few granules below.

Type Locality. — Thirty-five km. north of Juchitlan, Jalisco,
Mexico.

Host. — Ficus sp.

Type Material. — The female holotype, male allotype, and 66
para types were collected at the type locality on July 3, 1965, at an
elevation of about 1.300 m., by S. L. Wood, from fig branches.

The holotype, allotype, and paratypes are in my collection.

Stegomerus, n. g.

This genus superficially resembles several genera in the Crypha-
lini and the Micracini. but it is sufficiently unique that a question
exists concerning the tribal group to which it should be assigned.
Schedl (personal communication) placed it in the specialized Micra-
cini because the pronotum lacks a finely raised, basal and lateral
line; I include it with the primitive Cryphalini, near Cryphalomor-
phus Schaufuss. because the anterior coxae are contiguous, because
the eye is emarginate. because of the Cryphalus-\\\e antennae, and
because fimbriate hairs are absent from the head.

Description. — Head subglabrous; frons convex, slightly im-
pressed in male. Eye very long, broadly emarginate, finely granu-
late. Antennal scape rather short, club-shaped; funicle 5-segmented.
the distal segments not conspicuously wider. Pronotum coarsely
asperate on anterior slope, the anterior margin usually armed; sum-
mit well defined; basal and lateral margins rounded, not marked by
a fine lateral line. Elytra elongate, costal margins descending behind.
Anterior tibiae gradually widened on basal two-thirds, obliquely

The Great B-^sin Naturalist
130 STEPHEN L. WOOD Vol. XXVII, No. 3

narrowed toward apex, the lateral margin armed by several small
teeth on distal two-thirds; third tarsal segments narrow, laterally
compressed. Monogamous; phloeophagous in vines and lianas.
Type-species. — Stegomerus vulgaris Wood, described below.

1. Interstrial scales and hair in uniseriate rows on disc

and declivity; striae clearly evident; small

species, usually less than 1.2 mm PYGMAEUS

Elytral vestiture and punctures abundant, confused

at least on declivity; usually larger than 1.4 mm 2

2(1). Setae on basal half of elytral interspaces 9 and 10
consisting of fine, long hair; suture 2 on anten-
nal club much more strongly procurved than
suture 1 ; scales on elytral disc almost in uni-
seriate rows 3

Setae on basal half of elytral interspaces 9 and 10
primarily scalelike, short; sutures 1 and 2 on
antennal club equally, weakly procurved; scales
on elytral disc abundant, confused 4

3(2). Smaller. 1 3-1.6 mm.; scales on declivital interspaces

2 and 3 mostly uniseriate; punctures of striae
and interstriae somewhat smaller, essentially in

in rows; in Canavalia villosa CHIRIQUENSIS

Larger, 1.8-2.1 mm.; scales on all declivital inter-
spaces strongly confused; punctures of striae
and interstriae coarser, completely confused; in
Muelenbeckia tamnifolia MONTANUS

4(2). Scales on at least some of discal interstriae between

3 and 9 in uniseriate rows; declivital scales
broad, usually less than twice as long as wide.

the apex of each scale truncate MEXICANUS

Scales on all discal interstriae multiple, confused; de-
clivital scales more than twice as long as wide,
the apex of each scale rounded VULGARIS

Stegomerus pygamaeus, n. sp.

As indicated in the key. this species is distinguished from others
in the genus by its small size, and by the uniseriate strial and inter-
strial punctures, with the elytral scales and hair in rows.

Male. — Length 1.1 mm. (paratypes 0.9-1.2 mm.). 2.5 times as
long as wide; color dark brown, the summit of pronotum and the
elytra somewhat lighter.

Frons narrow, convex above, flat below, the epistomal margin and
its obtuse, median lobe slightly elevated; surface reticulate above,
subreticulate below, and finely, sparsely punctured; vestiture on
flattened area moderately abundant, consisting of fine, rather long

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 13l

hair. Eye elongate, about three times as long as wide; shallowly,
broadly emarginate; rather finely faceted. Antennal scape shorter
than the 5-segmented funicle; club marked by three equally, slightly
procurved sutures.

Pronotum 1.0 times as long as wide; widest one-third of length
from base, the sides weakly arcuate, broadly rounded in front;
armature on anterior margin poorly developed; summit at middle,
prominent; anterior area rather coarsely asperate; posterior and
lateral areas almost smooth and shining, with small, rounded, isolated
granules; vestiture consisting of hairlike setae over entire surface
with erect scales intermixed on posterior half.

Elytra 1.4 times as long as wide, 1.5 times as long as pronotum;
sides straight and parallel on more than posterior two-thirds, rather
broadly rounded behind; striae 1 feebly, others not impressed, the
punctures small, shallow; interstriae slightly wider than striae, the
punctures fine, uniseriate. Declivity moderately steep, convex; simi-
lar to disc except strial and interstrial punctures smaller and some-
what obscure. Vestiture consisting of uniseriate rows of short, semi-
recumbent, strial hair and erect interstrial scales, each scale about
three times as long as wide.

Female. — Similar to male except lower half of frons more near-
ly convex.

Type Locality. — Los Corchos, Nayarit, Mexico.

Host. — CanavaUa villosa (type), Cesirum scandans (paratype),
and Dioclea megacarpa (paratype).

Type Material. — The male holotype, female allotype, and 35
paratypes were collected at the type locality from stems of the above
host on July 10, 1965. at an elevation of about 20 m.. by S. L. Wood.
Eight paratypes were collected at Zamorano, Morazan. Honduras,
on April 18, 1964. from Dioclea megacarpa, elevation 700 m.; one
paratype bears the same date but was taken from CanavaUa villosa;
two paratypes were taken at La Lima, Cortez, Honduras, on May 5,
1965, from Cestrum scandens; all were collected by S. L. Wood. One
specimen, not included in the type series, was taken at San Jose,
San Jose Prov.. Costa Rica, on October 22, 1963, at an elevation of
1,600 m., evidently from CanavaUa villosa.

The holotype. allotype, and paratypes are in my collection.

Stegomerus chiriquensis, n. sp.

As indicated in the key this species is allied to montanus,
described below, but it may be distinguished by the smaller size, and
by the different arrangement of elytral punctures and scales.

Male. — length 1.4 mm. (paratypes 1.3-1.6 mm.), 2.6 times as
long as wide; color light brown, a few paratypes much darker.

Frons convex above, rather strongly, transversely impressed near
middle, flattened below; a median epistomal lobe present; surface
reticulate, with moderately abundant, fine, obscure punctures; vesti-

The Great Basin Naturalist
132 STEPHEN L. WOOD Vol. XXVII, No. 3

ture fine, rather sparse. Eye elongate, about three times as long as
wide; broadly, rather deeply emarginate; rather coarsely faceted.
Antennal scape shorter than the 5-segmented funicle; club large,
longer than combined length of scape and funicle, three procurved
sutures subangulate at middle, the first partly septate.

Pronotum as in pygmaeus, except anterior margin armed by a
series of about six small teeth.

Elytra 1.6 times as long as wide, 1.7 times as long as pronotum;
punctures of striae and interstriae very close, moderately large and
deep, in obscure rows. Declivity rather steep, convex; strial punctures
slightly smaller and interstrial punctures much smaller on disc,
shallow, in rows. Vestiture consisting of uniseriate rows of short,
fine, recumbent strial hair and longer, erect interstrial scales, each
scale four or five times as long as wide; sutural margin on declivity
with an additional row of shorter scales; an occasional scale on
declivity not in the rows.

Female. — Similar to male except transverse frontal impression
obscure or absent.

Type Locality. — Cerro Punta, Chiriqui, Panama.

Host. — Canavalia villosa.

Type Material. — The male holotype, female allotype, and 47
paratypes were collected at the type locality along Rio Viejo on
January 11, 1964, at an elevation of 1.800 m., by S. L. Wood from
stems of the above host.

Stegomerus montanus, n. sp.

As indicated in the above key this species is distinguished from
the closely allied chiriquensis Wood by the larger size and by the
different arrangement of elytral setae.

Male. — Length 2.0 mm (paratypes 1.8-2.1 mm.), 2.6 times as
long as wide; color dull black or grayish black, with el3"tra and sum-
mit of pronotum having a reddish cast.

Frons, eye, and antenna as in chiriquensis.

Pronotum 1.0 times as long as wide; widest on basal fourth, the
sides very feebly arcuate and converging slightly toward the rather
narrowly rounded anterior margin; summit at middle, impressed
behind; surface and vestiture about as in chiriquensis.

Elytra 1.5 times as long as wide, 1.7 times as long as pronotum;
disc densely, deeply, rather finely punctured, the punctures separ-
ated by distances less than half the width of a puncture, confused.
Declivity rather steep, convex; striae and interstriae not distin-
guishable, the punctures obscure. Vestiture consisting of rows of
minute, strial hair between confused rows of slender scales; setae
on interspaces 9 and 10 from base to declivity consisting of fine hair
only.

Female. — Similar to male except frons flattened, not trans-
versely impressed.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 133

Type Locality. — Volcan Irazu, Cartago Prov., Costa Plica.

Host. — Muelenheckia tarnnifoUa.

Type Material. — The male holotype, female allotype, and 15
paratypes were collected from the above locality and host on June
28. 19(i3. at an elevation of 2,800 m.. by S. L. Wood. Fourteen addi-
tional paratypes were collected at Villa Mills on Cerro de la Muerte,
CarUigo Prov., Costa Rica, on August 1, 1966, at 3,000 m. elevation
by S. L. Wood.

Habits. — This species inhabits the cambium region or the outer
woody layers of stems larger than about 1 cm. of the host, a woody
vine (or liana). The beetles are monogamous; each pair constructs an
ll-shaped system of egg galleries, with the entrance tunnel enter-
ing on the short transverse bar and the arms extending parallel to
the grain of the wood. The larvae feed in the phloem tissues and
usually construct tunnels parallel to the grain of the wood. Young
adults commonly tunnel into the deeper tissues.

Stegomerus mexicanus, n. sp.

This species is closely allied to vulgaris Wood, described below,
but it is distinguished by the shape and arrangement of the elytral
scales as indicated in the above key.

Male. — Length 1.7 mm. (paratypes 1.4-1.7 mm.), 2 5 times as
long as wide; color rather light brown, some specimens dull black,
with light vestiture.

Frons, eye, antennal scape and funicle as in chiriquensis; anten-
nal club large, 1.3 times as long as wide, with three equally, weakly
procurved sutures indicated.

Pronotum as in chiriquensis, but narrowly rounded in front,
armed by four large teeth.

Elytra 1.5 times as long as wide, 1.7 times as long as pronotum;
sides straight and parallel on basal three-fourths, broadly rounded
behind; disc with shallow, confused, close indefinite punctures. De-
clivity rather steep, convex; surface as on disc except punctures evi-
dently very feebly granulate. Vestiture consisting of uniseriate rows
of fine, strial hair and confused rows of erect interstrial scales, at least
some of the rows between interspaces 4 and 9 uniseriate; each scale
about three to four times as long as wide.

Female. — Similar to male except frons not as strongly flattened.

Type Locality. — Twenty-one km. west of Morelia, Michoacan,
Mexico.

Host. — Serjania sp.

Type Material. — The male holotype, female allotype, and 51
paratypes were taken at the type locality on June 15, 1965, at an
elevation of 2.300 m., by S. L. Wood, from Serjania stems less than
1 cm. in diameter.

The holotype, allotype, and paratypes are in my collection.

The Great Basin Naturalist
134 STEPHEN L. WOOD Vol. XXVII, No. 3

Stegomerus vulgaris, n. sp.

In the above key this species is placed near mexicanus Wood, but
it is distinguished by the shape and arrangement of the elytral scales.

Male. — Length 2.0 mm. (paratypes 1.7-2.0 mm.), 2.6 times as
long as wide; color light brown, some specimens with areas almost
dull black.

Frons weakly convex, slightly impressed just above epistoma;
epistomal process evident, projecting very slightly as a median epis-
tomal lobe; surface coarsely reticulate, with moderately large, in-
definite, isolated granules in central area, reduced in marginal areas.
Eye three times as long as wide; coarsely faceted; about one-fourth
divided by a broad emargination. Antennal scape and 5-segmented
funicle short, subequal in length, their combined lengths distinctly
less than length of club; club subcircular, 1.1 times as long as wide,
divided into four subequal segments by three broadly procurved
sutures, suture 1 partly septate at both ends.

Pronotum 1.0 times as long as wide; widest one-third of length
from base; basal angles rounded, sides rather strongly, arcuately
converging toward the narrowly rounded anterior margin; anterior
margin armed by about six teeth; oval area on anterior slope coarse-
ly asperate; summit elevated, near middle; posterior areas sub-
reticulate-granulate, with moderately large, rather close, rounded
granules; vestiture hairlike, except mixed with scales on basal
fourth.

Elytra 1.7 times as long as wide, 1.8 times as long as pronotum;
scutellum rather large triangular, flat; sides straight and parallel on
basal three-fourths, somewhat narrowly rounded behind; disc closely,
obscurely punctured, the punctures shallow, confused. Declivity
rather steep, convex; striae 1 and 2 obscurely indicated (largely by
color), otherwise similar to disc. Vestiture consisting of small,
obscure, uniseriate rows of strial hair and abundant, confused, erect,
interstrial scales, each scale about four times as long as wide.

Female. — Similar to male except frons very slightly, more
strongly convex.

Type Locality. — Volcan Zunil, Quezaltenango, Guatemala.

Host. — An unidentifiable woody vine or liana (type) ; Serjania
mexicana and S. triquetra (paratypes).

Type Material. — The holotype, allotype, and 66 paratypes
were collected at the type locality on May 27. 1964, at an elevation
of 1,000 m., by S. L. Wood, from an unidentifiable cut. woody vine
about 3-5 cm. in diameter. Other paratypes include 15 from San
Lucas, Paraiso. Honduras. April 22. 1964. 800 m., from Serjania
mexicana; 17 from Zamorano. Morazan. Honduras. April 18, 1964,
700 m., from Serjania triquetra; 1 from Palin. Esquintla. Guatemala,
May 19. 1964, 350 m., from Serjania sp.; 33 from San Jose, San Jose
Prov., Costa Rica, October 22. 1963, 1.300 m. evidently from

Nov. 30, 1967 NEOTROPICAL BARK BEETI ES 135

Serjania sp.; and 14 from Tapanti, Cartago Prov., Costa Rica, August
17, 1963, 1.300 m., evidently from Serjania sp.

The holotype. allotype, and paratypes are in my collection.

Habits. — Essentially as described above for montanus.

Xyleborus horridatus, n. sp.

This species is very closely related to horridus Eichhoff, but it
may be distinguished by the more slender body, by the more finely
sculptured pronotum and elytral disc, and by the more broadly im-
pressed, more coarsely and shallowly punctured elytral declivity
that lacks a submarginal, lateral row of fine tubercles. Specimens
labeled horridus from southern Central America probably are of this
species

Female. — Length 4.2 mm. (female paratypes 3.7-4.3 mm.),
2.3 times as long as wide; color rather dark reddish brown.

Frons rather evenly, broadly convex from epistoma to vertex,
the surface reticulate, rather deeply, somewhat obscurely punctured;
almost glabrous, except a few fine setae near epistoma.

Pronotum 1.0 times as long as wide; widest one-third from base,
the sides moderately arcuate from base to the somewhat narrowly
rounded anterior margin; summit behind middle closely asperate
from anterior margin to summit; posterior areas smooth and shining,
with small, sharply but shallowly impressed punctures; disc glabrous,
a few inconspicuous, hairlike setae in marginal areas.

Elytra 1.6 times as long as wide, 1.6 times as long as pronotum;
sides straight and diverging slightly to middle, then converging
shghtly to a level near apex, then rather abruptly, very broadly
rounded behind; striae 1 slightly, others not at all impressed, the
punctures small, clearly impressed; interstriae about four times as
wide as those of striae, shining, transversely wrinkled (particularly
1 and 2). Declivity beginning slightly in front of middle on inter-
spaces 1 and 2. increasing in width and steepness posteriorly; apical
and lateral margins acute on lower third, rather abrupt on middle
third, broadly rounded above, the margins distinctly, not strongly
elevated; declivital face similar to but broader than in horridus, the
median tubercles (2 pair of larger ones near middle, with 1 very fine
pair below and 3 above these) about as in horridus, lateral tubercles
just within lateral margin almost entirely absent. Vestiture fine;
scanty, particularly on declivity (abundant on declivity in horridus).

Male. — Length 3.3 mm. (male paratype 3.5 mm.); frons
medially impressed on upper half; eye reduced; anterior third of
pronotum concavely excavated, the anterior margin elevated and
armed by one median, subacute process, the asperites reduced within
the concavity; elytra evidently somewhat shorter, otherwise similar
to female.

Type Locality. — San Isidro del General, San Jose Prov.. Costa
Rica.

Host. — Citrus sp.

The Great Basin Naturalist
136 STEPHEN L. WOOD Vol. XXVII, No. 3

Type Material. — The female holotype, male allotype, and 16
paratypes were collected at the type locality on December 5, 1963,
at an elevation of about 1,000 m., by S. L. Wood, from Naranjo
dulce. Additional paratypes were taken as follows: 5 at Dominical,
Puntarenas, December 9, 1963, and 1 Pandora, Limon Prov., Costa
Rica, August 23, 1963, by S. L. Wood, from an unknown woody
vine and a log; 1 at Hamburg Farm, Sta. Clara Pr., Costa Rica,
June 10, 1928, by Nevermann; 9 at Finca El Encanto, La Plata
Huila, Colombia. April 28, 1959, by B. Humides. from Naranjo dulce
{Citrus sp.); 1 from Barro Colorado Island, Canal Zone, Panama,
February 26, 1929, by S. W. Frost.

The holotype. allotype, and paratypes are in my collection.

Xyleborus horridicus, n. sp.

The only species known to me that are even remotely related to
this are horridus Eichhoff and horridatus Wood, described above. It
may be distinguished from those species by the larger size, and by
the very different elytral declivity.

Female. — Length 6.5 mm., 2.5 times as long as wide; color
reddish brown.

Frons rather narrow, moderately convex from epistoma to vertex;
surface subshining, punctate-granulate; vestiture fine, rather sparse,
epistomal brush rather well developed. Eye coarsely faceted; one-
third divided by an emargination.

Pronotum 1.1 times as long as wide; widest about a third from
base, very feebly arcuate on basal third, more strongly arcuate and
converging on middle third, rather narrowly rounded in front; sum-
mit just behind middle, anterior area rather finely asperate; posterior
area shining, finely punctured. Vestiture fine, erect, sparse on disc,
more abundant and longer in lateral areas.

Elytra 1.4 times as long as wide, 1.4 times as long as pronotum;
sides straight on basal half, slightly wider just behind middle, arcuate
and narrowed slightly to level of sutural apex, then abruptly, broad-
ly, subemarginately truncate behind; striae not impressed, the punc-
tures small, moderately deej), somewhat staggered; interstriae shin-
ing, irregular, about three times as wide as striae, the punctures
somewhat staggered, some granulate on 1-3. Declivity beginning
very slightly in front of middle, rather gradual; up|)er half broadly
flattened, the margins rounded, not at all elevated; lower half with
lateral margins very strongly elevated, abruptly above, the summit
dorsally pointed as seen in profile, the elevation decreasing gradually
below to apex of suture; punctures in the broadly excavated area
moderately large, shallow, confused; upper third armed by one pair
of moderately large tubercles in line with striae 2, a few additional
small tubercles near declivital margin on interspaces 1-3. Vestiture
fine, confined to lateral areas except for a few minute, recumbent
hairs in excavated area of declivity.

Type Locality. — - Reyes, Bolivia.

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 137

Type Material. — The female holoty})e was collected at the type
locality in October, 1921. by W. M. Mann.
The holotype is in my collection.

Xyleborus bicornatulus, n. sp.

This species is allied to reconditus Schedl, but it is easily dis-
tinguished by the very different sculpture of the declivity. This
species has the row of tubercles on interspace 3 interru{)ted on the
lower half, with the tubercle near the middle of the declivity much
longer; the tubercles on interspaces 4-6 are also larger than in
reconditus.

Female. — Length 2.2 mm. (female paratypes 2.0-2.8 mm.),
2.8 times as long as wide; color almost black.

Frons broadly, transversely convex, longitudinally straight to
well above eyes, the epistoma slightly, gradually raised; surface re-
ticulate, with rather coarse, deep, sparse punctures; vestiture in-
conspicuous except along epistoma. Eye finely faceted; half divided
by a narrow emargination.

Pronotum 1.2 times as long as wide; widest just behind middle,
sides weakly arcuate, feebly if at all converging before the rather
abrupt anterolateral angles, rather broadly rounded in front; anterior
margin subserrate; summit at middle; posterior area smooth and
shining on disc, reticulate laterally, with sparse, very fine punctures;
vestiture sparse, inconspicuous.

Elytra 1.6 times as long as wide. 1.3 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, then broadly
rounded, almost straight on median third of posterior margin, the
arcuate portion strongly serrate; striae not impressed, the punctures
very small, distinct; interstriae shining, not entirely smooth, about
four or five times as wide as striae, the punctures about as small as
those of striae. Declivity beginning very slightly more than one-third
the elytral length from base, gradual, rather broadly excavated, the
sides moderately elevated, rounded; striae 1 and 2 wider than nor-
mal, distinctly punctured; interspaces 1 and 2 shining and uni-
seriately punctured, each with three or four tubercles at declivital
base, 1 weakly elevated. 2 impressed; interspace 3 somewhat ele-
vated, upper third armed by a row of up to five tubercles, an addi-
tional much longer spine at middle of declivity; apex of interspace
3 near declivital margin with one or two tubercles; interspace 4 with
a row of rather coarse, pointed tubercles on upper third, 5 with a
similar series on middle half, 6 and 7 with similar series on most of
lower half, a few tubercles extending almost to apex. Vestiture con-
sisting of rows of interstrial hair, rather fine on disc and sides, stout
on declivity.

Male. — Not represented in the material at hand.

Type Locality. — Moravia. Cartago Prov., Costa Rica.

Hosts. — Ochroma sp., and Theohroma cacao (paratypes).

The Great Basin Naturalist
138 STEPHEN L. WOOD Vol. XXVII, No. 3

Type Material. — The female holotype and 21 para types were
taken at the type locality on March 11, 1964, at an elevation of about
500 m., by S. L. Wood, from an unknown log. Additional paratypes
were taken in Costa Rica as follows: 15 at Finca La Lola. Limon,
Prov., December 27, 1962, J. L. Saunders; 1 at Rio Damitas in the
Dota Mountains, San Jose Prov., Feb. 18, 1964, 250 m ; 3 Finca
Gromaco on Rio Goto Brus, Puntarenas Prov., July 14, 1963, 500 m.
One para type was taken 12 km. southeast of El Ha to del Volcan,
Chiriqui, Panama, January 7, 1964, 1,000 m.; and 5 paratypes were
taken at Pamplona, San Vicente, Santander, Colombia, on June 26,
1959, by S. Betancourt, from balsa.

Corthylus splendens, n. sp.

This species is very closely related to collaris Blandford and
possibly could be a subspecies. It differs from collaris by the shorter
frontal pubescence on the female, by the much smaller cirrus on the
female antennal club, by the shining, more finely punctured elytra,
and by the more shallowly, more narrowly impressed elytra! de-
clivity.

Female. — Length 2.0 mm. (paratypes 1.8-2.1 mm.), 2.3 times
as long as wide; color yellowish to reddish brown, the elytra slightly
darker (variable in series; elytra black in some paratypes).

Frons rather shallowly concave from eye to eye, from epistoma
to vertex well above upper level of eyes, all margins rounded except
lower lateral margins abrupt; surface of concavity evidently minute-
ly subgranulose, obscured by abundant, erect, uniformly distributed,
short hair of uniform length. Eye large, very coarsely faceted; deep-
ly emarginate. Antennal club large, asymmetrically produced an-
teriorly as in collaris, with sutures 1 and 2 indicated by grooves;
cirrus present along distal margin, scanty, not extending beyond
apex.

Pronotum 1.0 times as long as wide; sides almost straight and
parallel on about basal half, anterolateral angles rather abrupt,
somewhat narrowly rounded in front; anterior margin armed by
about a dozen low, irregular teeth; anterior area asperate to in-
definite summit at middle; posterior area reticulate, subshinmg,
punctures not evident; vestiture inconspicuous, confined to anterior
and lateral marginal areas.

Elytra 1.2 times as long as wide, 1.2 times as long as pronotum;
sides very slightly arcuate, not converging behind before declivital
base, rather broadly rounded behind; elytral disc smooth, shining,
with a few lines, the punctures minute, shallow, distinct, sparse,
those on striae almost in identifiable rows. Declivity convex, rather
steep, very slightly impressed on upper half toward interspace 2;
interspace 1 convex, slightly elevated, striae 1 and interspace 2
(medially) impressed, gradually ascending laterally to interspace 3;
punctures of striae 1 moderately large, in rows, those on 2 small,
ahnost entirely obsolete, others not discernible; some punctures of

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 139

interspaces 3 evident. A few, fine, hairlike setae present on sides and
lower declivity.

Male. — Similar to female except frons convex, with a conspicu-
ous, transverse impression just above epistoma, very coarsely, deep-
ly, sparsely punctured, glabrous; pronotum more narrowly rounded
in front and armed by only two large, pointed teeth.

Type Locality. — Sixteen km. southeast of Cartago on the Pan-
American Highway, Cartago Prov., Costa Rica.

Hosts. — Miconia globuliflora (type), M. dodecandra, Conostegia
oersted iana, Phoebe mexicana. Solarium torvum, Theobroma cacao,
and several other woody plants.

Type Material. — The female holotype, male allotype, and 37
para types were collected at the type locality on September 24, 1964,
at an elevation of 1,800 m., by S. L. Wood, from branches of the
above Miconia. Other para types were taken in Costa Rica, in 1963,
by me. as follows: 6 at the type locality, July 3; 2 at Peralta. March
10, 1964. 500 m.; 11 on July 2 and 13 and on Oct. 24 at Tapanti at
1.300 m.; and 1 at Turrialba on July 5, 700 m., Cartago Prov.; 8 at
San Ignacio de Acosta, San Jose Prov., on July 5, 1,500 m.; 14 at
Pandora, Limon Prov., on July 23, 50 m.; and 1 at Volcan, Punt-
arenas Prov., December 11, 700 m. Ten para types were taken De-
cember 27, 1962, and 2 in July, 1963, at Finca La Lola, Limon Prov.,
Costa Rica, by J. L. Saunders; and 4 at 12 km. southeast of El Ha to
del Volcan, Chiriqui Prov.. Panama, on January 7, 1964, 1,000 m., by
S. L. Wood.

The holotype, allotype, and paratypes are in my collection.

Tricolus saundersi, n. sp.

This species is in the same species group as nodifer Blandford,
but it is smaller, reddish brown in color, and has the lateral profile
of the third declivital tooth more nearly quadrate.

Male. — Length 2.3 mm. (paratypes 2.3-2.6 mm.), 2.8 times as
long as wide; color reddish brown.

Frons broadly convex, the surface rather finely reticulate and
finely punctured, with a large, transversely oval, slightly elevated,
coarsely reticulate area occupying central half, its upper limits
slightly above upper level of eyes; vestiture limited to epistomal mar-
gin. Antennal funicle three-segmented; club broadly obovate, suture
1 weakly, 2 moderately procurved, both sutures partly septate.

Pronotum 1.4 times as long as wide; widest near base, sides al-
most straight, almost imperceptibly converging anteriorly on basal
half, then very narrowly, almost subangulately rounded in front;
anterior margin armed by a series of about a dozen teeth, the median
one larger and projecting more nearly forward; asperities decreasing
in height posteriorly, appearing a short distance behind summit as
shining wrinkles; posterior areas reticulate, very finely, sparsely
punctured; glabrous.

The Great Basin Naturalist
140 STEPHEN L. WOOD Vol. XXVII, No. 3

Elytra 1.5 times as long as wide, 1.6 times as long as pronotum;
sides almost straight and subparallel to base of declivity then ar-
cuately narrowed about a fourth to the projecting third declivital
teeth, very broadly, shallowly emarginate behind; surface almost
smooth, shining, the punctures rather fine, not close, strial and inter-
s trial punctures not close, in obscure rows except confused near
suture. Declivity abrupt, broadly excavated, the lateral and upper
margins abruptly elevated; upper margin bearing a small, acute
tooth as tall as wide, in line wdth striae 1, the second tooth located
on lateral elevation just above middle of declivity, subquadrate,
about one and one-half times as long as basal thickness, projecting
caudad and slightly dorsad; teeth 1 and 2 connected by an acutely
margined ridge thicker than high; tooth 3 just above and basally
separated from the acutely marked posterior declivital margin, sub-
quadrate, with upper basal margin expanded, twice as thick and
almost twice as long as 2, directed primarily caudad but slightly
inward and upward; ridge connecting 2 and 3 thicker than above,
but with the same continuous acute inner margin that ends in 3;
excavation broadly, transversely concave, the suture slightly ele-
vated, the punctures rather fine, shallow, not sharply defined, con-
fused. Glabrous.

Female. — Similar to male except posterior face of antennal
club bearing several scattered, long hairs.

Type Locality. — Dominical, Puntarenas Prov., Costa Rica.

Host. — Theobroma cacao.

Type Material. — The male holotype, female allotype, and 4
para types were collected at the type locality on September 29, 1964,
near sea level, by J. L. Saunders, from a branch of cacao.

The holotype, allotype, and paratypes are in my collection.

Tricolus spectibilis, n. sp.

This species is very closely related to speciosus Schedl from
which it is distinguished by the more strongly projecting second
declivital spine which is horizontal on its upper, distal half.

Female. — Length 2.4 mm. (paratypes 2.3-2 5 mm.), evidently
3.0 times as long as wide (elytra slightly spread); color very dark
brown.

Frons not visible, head withdrawn into prothorax in all speci-
mens except allotype, evidently as in male, described below.

Pronotum 1.3 times as long as wide; sides subparallel but feebly
arcuate on basal half then convergently arcuate to the narrowly
rounded, subserrate. anterior margin; asperities decreasing in size
posteriorly, ending at or before summit; posterior area subreticulate.
finely, deeply, rather sparsely punctured; glabrous.

Elytra 1.7 times as long as wide, 1.3 times as long as pronotum;
sides almost straight and parallel to level of middle of declivity, then
very broadly rounded behind, with a rather broad, deep, sutural

Nov. 30, 1967 NEOTROPICAL BARK BEETLES 141

emargination; strial jmnctures fine, shallow, in rather indefinite
rows; interstriae smooth, w^ith a few irregularly placed punctures
identical to those of striae. Declivity beginning two-thirds elytral
length from base, broadly, deeply excavated, with upper and lateral
margins acutely, rather strongly elevated except rounded on inter-
space 1 ; upper margin armed on interspace 2 by a small, pointed
tooth; a second ])rominence on lateral margin just below middle, its
basal margins in lateral profile form a 90 degree angle (somewhat
blunt) ; the acute margin continuing to apex; profile as in speciosus
except the second prominence higher; excavated area broadly con-
cave, shining, sutural inters{)ace finely, not strongly elevated,
smooth, the remaining area finely, rather closely punctured. Glab-
rous.

Male. — Similar to female except serrations on anterior margin
of pronotum much more prominent. Frons rather strongly, broadly
convex; surface reticulate and rather coarsely, deeply punctured
except on a small, more coarsely reticulate, impunctate area below
upper level of eyes equal in diameter to one-fifth width of frons;
vestiture limited to epistomal area.

Type Locality. — Finca La Lola, Limon Prov., Costa Rica.

Host. — Theobroma cacao.

Type Material. — The female holotype. male allotype, and four
paratypes were collected at the type locality on June 22, 1963, by
.1. L. Saunders, collection number 53.

The holotype. allotype, and paratypes are in my collection.

ON THE EARTHWORM FAUNA OF THE GREAT
AMERICAN DESERT AND ADJACENT AREAS^

G. E. Gates2

The Great American Desert, in long gone days when grade school
geography texts were large enough to conceal sundry surreptitious
activities from Teacher, was a conspicuous feature on American
maps. Little is heard today about that desert and a hasty glance
through the present, smaller-sized texts suggests that the region may
have lost much of its significance even if it has not entirely dis-
appeared. Deserts usually are not associated mentally with soft-
bodied, moisture-loving animals such as earthworms. Nevertheless,
some Asiatic, desert-like regions have large populations. Very little
information has been available hitherto about the earthworms of
the Great Basin and adjacent states. Fortunately, Prof. D E. Beck
was interested in this problem and for a number of years collected
oligochaetes as opportunity offered. Now that he is no longer able
to continue that activity he has interested a former student and col-
league. Prof. T. W. Barrett in carrying on the project.

No attempt is made to specify an exact boundary for the region
under consideration herein. All of it does have one character in
common which will become obvious as this contribution is read. Of
California only the arid southern region is included. A small portion
of the northwest corner of Idaho is excluded for reasons stated below.
Some data about states, for which no specimens were available, are
summarized (cf. appendix) to complete a review of available evi-
dence pertinent to the problems under consideration.

The introduction originally was as above but the manuscript was
not for a time submitted for publication because of anticipated ob-
jections to the title and the frequent use of "desert" in the text. Then
Hollon's book (1966) was encountered. According to that author,
the existence of a great American Desert is a fact that must not
be concealed or deprecated. Rather, it should be fully acknowledged,
with pride for past achievement in difficult circumstances and with
justified hope for future control insofar as available water permits.
The approximate limit of desert influence, as shown in Hollon's
frontispiece map, encloses most of the region herein discussed. Al-
though not because of inadequate rainfall, Minnesota, Iowa and
Missouri are included for reasons explained below.

1 . From research financed bj' the National Science Foundation.

2. Department of Zoology, University of Maine, Orono.

142

Nov. 30, 1967 DESERT EARTHWORM FAUNA 143

ACANTHODRILIDAE

Microscolex Rosa, 1877
Microscolex dubius (Fletcher, 1887)

California. — San Diego Co.: Alpine, 1836 feet, bottom of can-
yon, relatively undisturbed area east of city on Route 80, moist
humus type of soil at stream bank, March 10, 1966, 0-0-5. Beck;
Santa Barbara Co.: Santa Barbara. 2 miles south of city limits, high-
ly granular sand covered with about a quarter inch of humus of
stream bank, under Pacific Coast Highway, March 25, 1966, 0-0-2.
Beck.

Arizona. — Maricopa Co.: Mesa, March 6, 1966, 0-0-5 and
2-0-17. T. Barrett and D E. Beck. Wickenburg, March 8, 1966, along
banks of Hassayampa River, March 8, 1966, 0-0-1. Beck.

Remarks. — A/, dubius had been reported previously from Cali-
fornia but like other earthworm species only from a northern part
of the state.

Reproduction is parthenogenetic.

Microscolex phosphoreus (Duges, 1937)?

Arizona. — Luminescent specimens (alive) secured by Barrett
were considerably damaged during transportation. The spermathecae
were like those of M. phosphoreus. No other species of the United
States is known, at present, to have those two characteristics along
with an appropriate size. There is no previous record of the species
from Arizona. An older record of phosphoreus is for Santa Barbara
County, California.

Remarks.— Reproduction in phosphoreus probabl}'^ is partheno-
genetic. Both species of Microscolex are widely distributed anthro-
pochores, probably originally from southern South America, certain-
ly exotic anywhere in North America.

Lumbricidae

Allolobophora Eisen. 1874

Allolobophora chlorotica (Savigny, 1826)

California. — This species was first recorded from California
nearly ninety years ago as ripnria. Less than a quarter of a century
later it was simultaneously recorded from 29 places in the state. That
presumably was the basis for a subsequent erroneous characteriza-
tion (Michaelsen, 1900) of the distribution as "ganz Nord-Amerika."
Actually to this date, the species has only been reported from about
a score of the 49 mainland states and but five of the Canadian prov-
inces.

The species is not now expected to be common in southern Cali-
fornia.

The Great Basin Naturalist
144 G. E. GATES Vol. XXVII, No. 3

Reproduction is obligatorily biparental.

Idaho. — Idaho Falls, September 2, 1952, 5-16-3. Beck; Canyon
Co.: Middleton, from sod below surface of Boise River one mile south
of the town, June 19, 1964, 0-0-5. Beck; Clark Co.: Birch Creek, at
6500 feet, about 15 miles south of Lemhi County border, from turf
under birch, willow and an occasional quaking aspen, June 3, 1952,
1-2-14. Beck; Lemhi Co.: Salmon, 11 miles to the north, ca. 4100
feet, from fibrous root system of grassy turf of a relatively dry
meadow near Salmon River, June 1, 1952, 2-3-8. Beck, Mary Clark,
Grace Grant.

Remarks. — Megascolides americanus Smith, 1897, of course was
the first species to be recorded from within the present political limits
of Idaho. The worms were found just east of the Washington border
near Pullman. The latter is presumed to be the type locality. The
presence of M. americanus in Latah County, if not a result of trans-
portation, puts the place where it was found into a Pacific zoogeo-
graphic region and thereby removes it, as already mentioned above,
from the area under consideration herein.

The records above are the first of an identified megadrile species
for a very large unexcluded portion of Idaho. That state is one of
those in which Seton (1904 or 1929 summarizing 19th century ob-
servations and records) himself could find no earthworms. However,
long ago Wilcox (1884) reported that "At Boise City, Idaho, some
enthusiastic disciples of Izaak Walton imported and successfully
reared the coveted bait in soil suited to the habitat of the Lumbri-
cidae!" The species is unlikely to have been chlorotica but certainly
does not need to be L. terresiris {q. v. below) as some are likely to
think. The introduction was made, according to Wilcox, because
there were no "angleworms" in prairies of the "Northwest."

European settlement began in 1860. Some years probably passed
before angleworms were first introduced.

Nevada. — Elko Co.: Ferguson Springs, ca. 25 miles south of
Wendover (Utah) on Route 50, under boards and rocks along edge
of drain where seep water from the spring coursed down the hillside.
August 5. 1950, 0-0-1. Beck ("Surrounding country extreme desert.")
Lyon Co.: Wabuska, July 5, 1966. 4-6-28. Carson River, south of
Silver Spring. July 15, 1966, 0-1-1. E. V. Komarek. Sr.; Washoe
Co.: Wadsworth, damp porous soil associated with root system of
rank plant growth some distance from edge of the Trukee River,
August 6, 1950, 0-0-13. Beck.

Remarks. — These are the first records of earthworms for Ne-
vada. For reasons now unknown, neither Seton (1929) nor Wilcox
(1884) mention Nevada although it already had been cut off from
Utah as a separate territory in 1861.

Utah. — Sanpete Co.: Mt. Pleasant. Pleasant Creek Picnic
Grounds, at contact line of conifers and aspens, August 5, 1951.
0-0-8. Beck; Summit Co.: Wanship, at 5900 feet, about two feet
above stream bed, six inches under sod kept moist by ephemeral

Nov. 30, 1967 DESERT EARTHWORM FAUNA 145

seepage, July 13. 1962, 0-1-2. Beck; Uintah Co.: Vernal, soil by a
roadside creek, August 27. 1952, 0-0-1. Beck; Utah Co.: Soldier Sum-
mit, upper three inches of heavy clay under cardboard and wooden
boxes of an abandoned garbage dump. May 22, 1952, 1-2-10. Provo,
April 12, 1951, 0-0-3. Beck; Washington Co.: Pine Valley, July 16,
1953, 0-0-3. Zion National Park, detritus and under stones along
one of the side streams near Weeping Rock. Virgin River, September
3, 1950, 0-1-1. Beck.

Remarks. — The records above are the first for Utah of an identi-
fied species. However, earthworms probably have been there for at
least a century. Provo, Utah, according to Hallock. (1877, p. 353),
"is the only place west of the Rocky Mountains where an abundance
of earthworms may be had, the species having been introduced by
an enterprising physician from the east."

The introduced worms are unlikely to have been of chlorotica,
as anglers, for several centuries, have deemed them useless as fish-
bait. Lurnbricus terrestris probably was believed, by the enterprising
physician, to be his species, but one, or more, of the allolobophoras
is much more likely.

Arizona. — Mohave Co.: Cane beds, under rocks in sandy soil
with little humus by a trough at which cattle drink, from an area
no "more than a vard square," July 1, 1953, 0-1-17. Beck; Yavapai
Co.: Prescott, March 7, 1966, 0-6-14. Beck.

Remarks. — A. chlorotica had not been recorded hitherto from
Arizona, one of the states in which Seton (1929) failed to find
earthworms.

Montana. — Mineral Co.: Saltese, 3 miles east of Lookout Pass
on Route 10, at about 4000 feet, mucky black forest soil near a small
stream and/or a coarse sand and gravel mixture by the stream side,
June 25. 1954. 0-0-4. Beck.

Remarks. — This is the first record of an earthworm species for
the large state of Montana. Seton (1929) found no earthworms there.

Colorado. — Routt Co.: Along the Yanipa River, ca. 2 miles
north of Steamboat Springs, under stones, boards, logs and in loose
soil near willow clumps in a small meadow, August 27. 1952, 0-0-8.
Beck.

Remarks. — This species was recorded from Colorado once before
(Smith. 1917) but without specification as to county or town.

In his Colorado searches, Seton (1929) found no earthworms.

Further evidence as to former absence in the state was provided
by Cockerell (1924):

"In the mountain district of Colorado, and the adjacent upland
plains, it appears that earthworms were formerly absent. Old settlers
assure us that when they first came there were none. This seemed
at first almost incredible, but in recent years Prof. Frank Smith of
the University of Illinois has collected earthworms in Colorado,
and found only the widespread presumably introduced types." (c/.
Smith, 1917)

The Great Basin Naturalist

146 G. E. GATES Vol. XXVII, No. 3

Cockerell is but one of the very many biologists who have thought
the facts to be "almost incredible."

Allolobophora longa Ude, 1885

Colorado. — Chaffee Co.: Buena Vista, Rancho Sa watch, August
14, 1953, 0-1-0. Ottys Sanders.

Remarks. — This is the first record of this species for any place
west of the Mississippi River.

Allolobophora trapezoides (Duges, 1828)

California. — Imperial Co.: Holtville, bank of Alamo River
west of town, March 9, 1966, 0-0-2. Beck; Kern Co.: Wasco, resi-
dence of P. D. Pilsbury, dark wet earth under a hedge around outlet
of an air-conditioning hose, August 21, 1950, 4-2-2. Beck. With
fibrous root system of Bermuda grass at edge of a stagnant pool in
an irrigation ditch, August 21, 1950, 0-1-4-6. P. D. Pilsbury per
Beck; Los Angeles Co.: Torrance, six inches of moist organic soil
above sand in a flower bed, August 29, 1950, 5-1-1. Beck. Foothills
of Angeles Mountains, at ca. 1500 feet, December 31, 0-4-8. D.
McKey-Fender. Orange Co.: Capistrano, one mile north in ditch by
highway, December 29, 0-0-4. D. McKey-Fender. San Bernardino
Co.: Victorville, one half mile west of bridge over Mohave River,
in wet, coarse sand under algal compost, September 1. 1950, 0-0-3.
Beck. Santa Barbara Co.: Santa Barbara, two miles south of city
limits on Pacific Coast Highway, very moist, highly granular sand
covered with about a quarter inch of humus on bank of stream under
highway, March 25, 1966, 0-0-1. Beck. San Diego Co.: Pine Valley,
June 21, 1966, 0-0-10. E. V. Komarek, Sr. Alpine, bottom of a shal-
low canyon, at east entrance to city on Route 80, moist humus soil
on banks of small stream. March 10, 1966, 12-9-19. Beck (nine
cocoons secured at this site may have been deposited by A. trape-
zoides) .

Remarks. — A deep brown allolobophora, so common at moist
or swampy places in California, was mentioned by Eisen in several
of his pre- 1900 contributions. The species must have been lacking
in Scandinavia or he would have had a name for it. Indeed, because
of that unfamiliarity he seems at times to have suspected that the
species might be native. Much more recently and with much less
justification L. rubellus has been believed to be native to Washing-
ton State merely because of its commonness there.

The two anthropochorous allolobophoras, often correctly charac-
terized as deep brown, are A. longa and trapezoides. Neither had
reached Scandinavia by 1900. Since then A. trapezoides has been
secured in a northern part of California and once, rather surprising-
ly, at a height of 5800 feet. No evidence indicative of any Cali-
fornia colonization by longa has been found. A. trapezoides now
seems to be originally from a Mediterranean source. As such, the

Nov. 30, 1967 DESERT EARTHWORM FAUNA 147

species could have been introduced at an early date by Spaniards
who established 21 missions from San Diego to Sonoma during 1 769-
1823. The vehicle of transport could have been earth with wine-
grape })lants which were taken to most if not all of the missions.

Absence of earthworms in middle and southern California also
was mentioned in some of Eisen's publications. Seton (1929) found
none in the mountains and the dry parts.

Reproduction, in A. trapezoides, is parthenogenetic. However in
a sperm-maturing. Italian morph embryonic development does not
begin until it is initiated by penetration of sperm into the ovum
(pseudogamy). Polyploidy (cf. Muldal, Omodeo, 1952) has been
reported.

Idaho. — Bannock Co.: McCammon, 3 inches below surface and
in an area ca. 2 feet square, a few feet from bank of Portneuf River
and about a foot above water level. May 30. 1952, 0-1-8. Beck. Butte
Co.: Arco. to the south about 1 mile, heavy black clay loam at
water's edge and a more loose and only slightly moist soil a few feet
above the water's edge. May 31, 1952, 0-2-19. Beck, Mary Clark,
Grace Grant. Canyon Co.: Middleton, four inches below surface of
an area with sod where logs and other detritus had accumulated near
Boise River, June 19, 1954, 0-3-11. Beck. (Some of the worms se-
cured at that site were said to be "balled" and presumably were in
diapause. Two couples were copulating and they may have been of
the amphimictic species found at the same site, though male sterile
individuals do sometimes still copulate with each other. Cocoons
were deeper down and in sandy soil.) Fremont Co.: "Osborne
Springs." 25 miles north of Ashton on Route 91. in heavy black
loam. 6-8 inches below surface and under a thick grass turf but
ground dry at surface, September 1, 1952. 0-0-0-6. Beck (The col-
lector stated that the worms were in the process of "balling up.")
Kootenai Co.: Worley, upper three inches of soil in grassy woodland
area close by a stream, June 24, 1954, 0-1-18. Beck. Latah Co.: Mos-
cow, eight miles to the south, in area surrounding drain-off from
cesspool. May 20, 1950, 0-0-1. C. W. Lame. Lemhi Co.: Salmon, hill-
side seep spring area, on route 93, 1 mile north of town, May 31,
1952, 0-3-16. Beck. 11 miles to the north, ca. 4100 feet, from fibrous
root system of grassy turf of a relatively dry meadow near Salmon
River, June 1, 1952. 0-4-11. Beck, Mary Clark, Grace Grant. Lincoln
Co.: Shoshone Falls. Porous black soil, under rocks in shaded grassy
areas, near the "Falls" area. June 18, 1954, (10?) -4-12-1. Wet to
mucky sites near a spring that is the water supply for the public,
June 18. 1954. 0-0-4. Sandy area beneath a west facing cliff, June
19, 1954. 0-1-3. Beck.

Remarks. — A. trapezoides is the second oligochaete species to
be recorded herein for the first time from Idaho. In that area Seton
(1929) found no earthworms. His conclusion as to absence was
supported by Wilcox (1884). A. trapezoides is more likely to have
been imported to Idaho in the 1880's than L. tcrrestris which at that

The Great Basin Naturalist

148 G. E. GATES Vol. XXVII, No. 3

time })robably had nowhere near reached its present and still limited
American range.

Nevada.- — Elko Co.: Ferguson Springs, on Highway 50, under
boards and rocks along edge of drain channel where seep water from
the spring coursed down the hillside, surrounding country extreme
desert, August 5, 1950, 1-3-5. Beck. Lyon Co.: Wabuska, July 5,
1966, 1-1-0. E. V. Komarek, Sr. Washoe Co.: Wadsworth, from
damp, porous soil associated wdth root system of rank plant growth
some distances from edge of the Trukee River, August 6, 1950,
0-3-11. Beck.

Remarks. — A trapezoides is the second earthworm species to be
recorded herein for the first time from Nevada. Although this state
was not mentioned by Seton (1929) he obviously believed (c/. p.
144) its earthworm fauna to be the same as that of other desert states.

European settlement began around 1849.

Utah. — Beaver Co.: Beaver, soil of a peat-like consistency near
a small stream, July 11, 1952, 0-1-13. Marvin Coffey per Beck.
Wahwah Springs. Moist soil along stream bank, in a marshy soil
and under rocks near the marshy area, September 12, 1952, 8-2-0.
Highly organic soil and under rocks near edge of desert spring and
its runoff. September 12, 1950, 1 (-f 24?) -4-5-1. Beck. Box Elder Co.:
Lucin, seven miles to the north at a desert spring where water waj
salty and vegetation was of the alkaline desert type, June 19, 1952
0-0-3. Beck. Daggett Co.: Willow Creek, at 6000 feet, under plam
cover of sage and juniper, near Uintah Mountains, June 12. 1953
0-0-11. Beck. Garfield Co.: Bear Valley Junction (probably Angus
1952?), 4-4-0. M. A. Coffey per Beck. Iron Co.: Cedar Breaks Na
tional Monument, June 20, 1953. 3-6-20-2. Beck. Juab Co.: Levan
8 miles to the south on Route 28, 3-6 inches below surface, unde:
stones and boards in an area about two yards square, April 19. 1952
1 ( -f ?) 20-13-11. Beck. (The area is extremely dry in the summer
snow had covered the ground 10 days before the collection.) Pint
Co.: Marysvnle, 2 miles south of Big Rock Candy Mountain on Rout
89. with fibrous root system of grass and sedge quite close to wate
level in Sevier River. June 26, 1952, 1-0-10. Beck. Rich Co.: Sag
Creek Junction, bank of a creek, with wet soil and fibrous roots, a
6500 feet. August 21, 1952, 0-3-14. Woodruff, at the edge of
marshland seven miles west and in the confines of a canyon moutl
at 7000 feet, six inches below surface in very porous and moist soi
June 25, 1953. 0-2-4. Beck. San Juan Co.: Kigalia Ranger Statior
Bears Ears, in black soil near stream at 8500 feet, June 8. 195f
0-5-22-9 and June 9, 1955. 0-0-2. Bluff. May 5, 1951, 0-1-4. Bed
Sanpete Co.: Mt. Pleasant, Pleasant Creek Picnic Grounds, beneat
bark and under trees fallen to the ground, August 5, 1951. 3-3-1:
Beck. Summit Co.: Wanship, at 5900 feet, about two feet abo\
stream bed. six inches under sod kept moist by ephemeral seepagi
July 13, 1962. 0-1-1. Echo. July 11. 1952, 0-0-2. Beck. Utah Co
Soldier Summit, upper three inches of heavy clay under cardboar

Nov. 30, 1967 DESERT EARTHWORM FAUNA 149

and wooden boxes of an abandoned garbage dump. May 22. 1952,
16-6-2. (Late juveniles had rudiments of genital tumescences in
rlitellar region but lume were recognizable in ix-xi. Tubercula puber-
tatis unrecognizable.) Lehi. Lehi Sugar Factory area, in wood of a
rotted board near pond, April 25. 1952. 1-0-0. Beck. (Elsewhere in
the area no worms were found.) Weber Co.: Ogden. depressions in
the flood plain plant growth is rank. September 3, 1952, 5(-^?)-5
( + ?)-0. Beck. Washington Co.: Grafton, upper 8 inches of heavy,
sandy, dark-colored loam, under rocks, boards, and logs in an or-
chard Aj)ril 5. 1952, 3-3-26. Beaver Dam Wash. Ed Terry Ranch,
3-10 inches below ground surface but under gopher mounds, in
alfalfa field, at ca. 2500 feet, in the Joshua tree-creosote belt, Feb-
ruary 23. 1952. 0-17-6. St. George or Springdale. Sej)tember 2, or
September 3, 1950, 0-0-1-2. Pine Valley, July 16. 1963. 0-1-14. Leeds,
Oakgrove Campground, at 700 feet, mucky soil along a seep stream,
July 14, 1953. (?)-2-4. Zion National Park, in detritus and under
stones along one of side streams near Weeping Rock, Virgin River,
September 3. 1950, 0-2-2. Beck. Wayne Co.: Elkhorn Ranger Station,
Thousand Lake Mt., at 9700 feet, damp, loose black soil under rocks
and dead timber, and in a forest of quaking aspen, Populus tremu-
loides, August 9. 1952. 3-2-10. Beck.

Remarks. — A. trapezoides is the second species to be recorded
herein for the first time from the state of Utah.

The original home of the taxon probably is in some part of the
Mediterranean region. Professor Beck was asked to provide informa-
tion as to how a Mediterranean earthworm could have been brought
to Utah. He replied {in lit.) that he believed some stock of fruit trees
came along with the first settlers as an agricultural program had
been worked out even before Mormons left Nauvoo. Arrival in Utah
was between 1847 and 1850. So plants started to arrive and were
well in growth soon after 1847 and began to be spread by coloniza-
tion north, south, east and west. Beck, speaking of the history of the
settlement of St. Cieorge with which he is well acquainted, stated
[in lit., Jan. 5. 1965): "Joseph E. Johnson, an early leader in horti-
culture in southwestern Utah, imported many fruit trees right after
settlement. These came by horseback, wagon, buggy, and carried by
hand; some from California and others from northern Utah. This
was between 1861 and 1870. Within that period the pe(jple had no
fruit, to a condition of abundance: Peaches, plums, pears, apples,
figs, nectarines, almonds, quince, raspberries, strawberries, pome-
granates, and apricots. One time he imported 1.000 maple trees
hoping to develop the maple sugar industry in southern Utah."
Plants presumably were brought across the plains from Mississippi
or from Los Angeles by way of the San Bernardino Pass. If wine-
grape plants were imported from Los Angeles the dark brown allolo-
bophora that was so common in California could have been brought
to LUah directly. Another possibility is that the species also could
have come along with plants from southern states or even a southern
part of Illinois.

The Great Basin Naturalist

150 G. E. GATES Vol. XXVII, No. 3

Arizona. — Cochise Co.: Alfalfa field, 4 miles north of Elfrieda
and ca. 25 miles north of Douglas, upper two inches of sandy loam
with perhaps 2^/2% organic matter, September 3, 1966, 18-1-1-15.
Leslie Canyon, McNeal, upper four inches of sandy to clay loam
with 2-3% organic content, along a small stream (no fishing),
under Arizona willow, September 3, 1966, 1-2-3-1. 16 miles north-
east of Douglas Canyon-Ricker Road, 1/4 mile east of road near cattle
pen at 4600 feet, from sod and upper 4 inches of black, heavy, clay
loam nearly saturated by four days of rain (no farms and permanent
streams in the area), September 16, 1966, 8-1-21. 3 miles south of
Junction of Leslie Canyon and main east-west Rucker Canyon road,
among roots in top four inches of loam by abandoned water storage
tank (for cattle, no fishing streams nearby), September 16, 1966,
5-3-6. Barrett per Beck. Tombstone, Costello Ranch, 8 miles from
town at elevation of ca. 5000 feet, in soil rich with manure (pH
ca. 7.5), July 23, 1933, 0-2-6 also 2 fragments, each containing a
clitellum. A. Petrunkevich per G. E. Pickford.

Gila Co.: Sierra Ancha Exp. Forest Headquarters. Two miles
north of Headquarters, along almost dry Rose Creek, in sandy loam
under walnut and oak, August 29, 1966, 0-0-0-9. Ten miles north
of headquarters, at confluence of Rose and Workman Creeks, top
four inches of sandy-clay loam at edge of meadow, under walnut,
sycamore and Arizona willow, at 6200 feet, August 29, 1966, 11-4-
0-1. Top 2 inches of loam to sandy loam under or near maple (but
not pine and fir), within 10 feet of bank on each side of Workman
Creek (but lacking 10-50 feet from the bank), at 6900 feet, August
30. 1966, 6-2-3. A dell under a maple. 3 miles up Workman Creek
from Young Road, at ca. 6500 feet, sandy loam with maple leaves
and grass, September 1, 1966, 9-3-6-3. Barrett per Beck. Mohave
Co.: Cane beds, under rocks in sandy soil with little humus by a
trough at which cattle drink, July 11, 1958, 0-1-8. Beck.

Remarks. — A. trapezoides probably was the member of the
caliginosa congeries that was recorded from Arizona 67 years ago.
Recent collections from the state have contained only one lot of
turgida and none of tuberculata. A very abnormal Tombstone speci-
men, at first tentatively referred to caliginosa (Gates, 1956), enabled
synonymization of the supposedly Lusitanian and very old A.
relictus Souther. 1909, without examination of the type (the only
known specimen). The homoeosis of both worms was maximal.

Arizona is one of the states in which Seton (1929) found no
earthworms. It is unlikely that he searched in gardens or sites to
which the animals could have been deliberately or accidentally in-
troduced.

Montana. — Ravalli Co.: Hamilton, four miles west and near
Blodgett Creek at 3600 feet, upper 3-4 inches of black soil mixed
with sand under Pinus ponderosa, June 1, 1952, 1 ( -|- l?)-2-18. Black,
heavy loam under coarse sandy soil near bank of Bitterroot River,
June 1, 1952, 0-3-2. Beck, Mary Clark, Grace Grant. Victor, moun-

Nov. 30, 1967 DESERT EARTHWORM FAUNA 151

tain meadow stream rising in the Bitterroot Mountains. June 25,
1954, 0-0-6-1. Beck.

Remarks. — No species of earthworm had been reported from
Montana before. The state is one of those in which Seton (1929)
found no earthworms.

The earliest European settlers, after traders, trappers and ex-
plorers, were miners. Farming is said to have been little till after
1880.

Wyoming. — Hot Springs Co.: Wilderness, 20 miles south of
Dubois, along bank of Deniwoodie River where it crosses Route 287,
under short grass turf at edge of stream, August 30, 1952 (2?) -26-5.
Beck. Teton Co.: Teton National Park, about 5 miles north of the
park boundary where Snake River crosses Route 287, sodded bank
of river (where bait had been dumped at end of day's fishing?),
August 31, 1952 (4?)-9-3-l. Beck.

Remarks. — This is the first record of an identified megadrile
species from the state of Wyoming. The area is mentioned as one
of those in which Seton (1929) could find no earthworms.

Probably few Europeans had settled in Wyoming before 1867
when the Union Pacific began to build its railroad through the state.

Colorado. — Boulder Co.: Longmont, under log in pasture at
Boulder Creek and its crossing by Route 287, August 29, 1952, 1-3-2.
Beck. Grand Co.: Hot Sulphur Springs, moist loose dark soil with
fibrous grass roots along bank of Colorado River about 7 miles south-
west of the town, August 28, 1952, 0-0-0-2. Beck. Larimer Co.: Fort
Collins, eight inches of dark, compact, sandy loam above water level
in seepage area at base of South Reservoir, August 29, 1952. 0-2-2-3.
Stonewall Creek, 30 miles south of Wyoming border on Route 287,
loose sandy soil with much humus in sodded area where willow
growth was rank, August 29, 1952, 0-15-8-2. Beck. Routt Co.: Steam-
boat Springs, two miles north, along edge of Yanipa River, also under
stones, boards, logs, August 27, 1952, 0-1-1. Beck.

Remarks. — These are the first records of A. trapezoides for
Colorado, a state in which Seton (1929) found no earthworms (also
cf p. 145).

Gold was discovered in 1850 but the rush came only in 1858.
Before that there probably had been little white settlement.

Iowa. — Jefferson Co.: Fairfield, Parsons College, May 22, 1951,
0-0-5. Von Ohlen.

Remarks. — A. trapezoides had been recorded for Iowa but as
A. iowana Evans. 1948. A cotype from the U. S. Nat. Mus. has been
examined. The Fairfield specimens mentioned above are from the
type locality and were secured by the same person who collected the
types of Evans' species.

Allolobophora tuberculata Eisen, 1874

California. — Santa Barbara Co.: Santa Barbara, March 25,
1966, 0-2-1. Beck.

The Great Basin Naturalist

152 G. E. GATES Vol. XXVII, No. 3

Remarks. — A. tuberculata had not been recorded from Cali-
fornia previously.

Reproduction, in A. tuberculata, is obligatorily biparental.

Idaho. — Canyon Co.: Middleton, four inches below surface of
sod where logs and other detritus had accumulated on bank of Boise
River, June 19, 1954, 0-0-3. Beck. (Worms were congregated in the
humus but cocoons were in sandy soil beneath. Two pair were in
copula. Although soil still was relatively moist, some worms were
"balled." Clark Co.: Birch Creek, about 15 miles south of Lemhi
County border on Route 28, from turf under birch, willow and an
occasional quaking aspen, at 6500 feet, June 3, 1952, 4-3-8. "Osborne
Springs," ca. 25 miles north of Ashton on Route 191, heavy black
loam, 6-8 inches below surface, in process of "balling," surface quite
dry, September 1, 1952, 1-1-3. Beck. Lemhi Co.: Salmon, 11 miles
to the north, ca. 4100 feet, from fibrous root system of grassy turf
of a relatively dry meadow near Salmon River, June 1. 1952, 0-8-11.
Beck, Mary Clark, Grace Grant. One mile north, seep area of a
hillside spring, 0-4-17. Beck. Morgan Co.: Weber Canyon. 6 miles
east of Coalville, heavy loam with humus in shaded woodland down
to depth of 8 inches, at 5600 feet, July 11, 1952, 0-1-20. Beck.

Remarks. — A. tuberculata is the third species to be recorded
herein for the first time from Idaho.

Nevada. — White Pine Co.: Kennicot Duck Creek Station, one
mile to the south, wet heavy clay soil with organic matter, August
5, 1950, 0-2-2. Beck.

Remarks. — A. tuberculata is the third species to be recorded
herein for the first time from Nevada.

Utah. — Cache Co.: Hyrum, north fork of the Blacksmith Fork
Can3'on, near edge of mountain stream at 7500-8000 feet, under
birch and sod, June 27, 1953, 0-5-6. Beck. (From 8-10 inches below
surface, in dark compact soil. No worms in top six inches of sandy
soil. Some were "balled.") Piute Co.: Kingston, bank of ditch at base
of cliff in canyon desert at 6100 feet, June 26. 1-0-2. Beck. Summit
Co.: Wanship, at 5900 feet, about two feet above stream bed. six
inches under sod kept moist by ephemeral seepage, July 13, 1962,
0-0-1. Beck. Utah Co.: Provo, Brigham Young University campus,
January 31, 1966, 0-1-0. Beck.

Remarks. — A. tuberculata is the third sy)ecies to be recorded
herein for the first time from Utah. The state was not mentioned
by Seton (1929) but he obviously believed its earthworm fauna
to be the same as in other desert states, i.e., none.

MoNTANA.^ — Mineral Co.: Saltese. 3 miles east of Lookout Pass
on Route 10. at about 4000 feet, mucky black forest soil near a small
stream and/or a coarse sand and gravel mixture by streamside.
June 25, 1954, 0-0-2. Beck. Ravalli Co.: Hamilton, nine miles south
on Route 93, bank of Bitterroot River, in black layer under sand and
turf, June 1, 1952, 0-21-7. Beck, Mary Clark and Grace Grant. Vic-

Nov. 30, 1967 DESERT EARTHWORM FAUNA 153

tor, from grassy roots of a mountain meadow stream, June 25, 1954,
0-0-16. Beck.

Remarks. — A tuberculata is the third species to be recorded
herein for the first time from Montana.

Wyoming. — Albany Co.: Laramie, August 29. 1952. 1-2-9. Beck.
Fremont Co.: Lander, soil with humus along bank of Papaogie
Creek, August 30. 1952. 0-7-1. Beck.

Remarks. — A. tuberculata is the second species to be recorded
herein for the first time from Wyoming.

Colorado. — Chaffee Co.: Buena Vista, Rancho Sa watch, August
14, 1953, 0-0-1. Ottys Sanders. Grand Co.: Hot Sulphur Springs,
under stones and in fibrous grass roots of moist, loose, dark soil with
much humus along banks of Colorado River, August 28, 1952, 0-3-25.
Beck. Larimer Co.: Fort Collins, seepage area at base of South
Reservoir, in upper eight inches of a dark, compact, sandy loam high
in humus above a wet level, August 29, 1952, 0-14-7. Beck.

Remarks. — These are the first records of A. tuberculata for
Colorado.

New Mexico. — Grant Co.: Fort Bayard, Central City, ca. 6190
feet, arroyo drainage, 6 and more inches below surface, February
27, 1966. 0-0-2. Beck.

Remarks. — This is the first record of a lumbricid earthworm for
the state of New Mexico.

The first permanent European settlement, by Spaniards, was in
1610.

Iowa. — Jefferson Co.: Fairfield, Parsons College, May 22, 1951,
0-3-4. F. W. Von Ohlen.

Remarks. — This is the first record of the species for Iowa. The
present specimens were secured from the type locality of A. iowana.
Evans also had specimens of tuberculata. He recognized that they
were different from the caliginosa he had studied in England and
that they might be of a new species. The taxon already had been
provided a name by Eisen as well as by Friend and Gates, inde-
pendently.

Allolobophora turgida Eisen, 1874

California. — San Diego Co.: Alpine, bottom of a shallow can-
yon, at east entrance to city on Route 80. moist humus soil on banks
of small stream, March 10, 1966, 2-5-25. Beck. (Nine cocoons found
at the site are less likely to be of A. turgida than A. trapezoides.)
Santa Barbara Co.: Santa Barbara, very moist, highly granular sand
covered with about a quarter inch of humus on banks of a small
stream. March 25, 1966. 0-0-1. Beck. Mendocino. June 30, 1966.
l(+4?)-l-l. E. V. Komarek. Sr. (Mendocino is the only locality at
which A. turgida is known to be present along with A. trapezoides
and tuberculata.)

Remarks. — A. turgida had not previously been recorded from
California.

The Great Basin Naturalist

154 G. E. GATES Vol. XXVII, No. 3

Reproduction, in A. turgida, is obligatorily amphimictic.

Idaho. — Bannock Co.: Pocatello, roadside parking and picnic
area to south on Route 91, under shubbery and boards in a very
moist situation, June 3, 1952, 0-0-7. Beck. Canyon Co.: Middleton,
four inches below surface of sod where logs and other detritus had
accumulated on banks of the Boise River, June 19, 1954, 0-0-8. Beck.
(C/. note on this site, tuberculata.) Latah Co.: Moscow, eight miles
to the south, in drain-off from a cesspool. May 20, 1950, 0-0-1. C. W.
Lame. Lemhi Co.: Salmon, 11 miles to the north, ca. 4100 feet, from
fibrous root system of grassy turf of a relatively dry meadow near
Salmon River, June 1, 1952. 0-2-5. Beck, Mary Clark, Grace Grant.

Remarks. — A turgida is the fourth species to be recorded herein
for the first time from Idaho.

Nevada. — Elko Co.: Ferguson Springs, under boards and rocks
along edge of drain channel where seep water from spring coursed
down the hillside, surrounding country extreme desert, August 5,
1950, 0-1-3. Beck.

Remarks. — A. turgida is the fourth species to be recorded herein
for the first time from Nevada.

Utah. — Cache Co.: Hyrum, north fork of the Blacksmith Fork
Canyon, near edge of mountain stream at 7500-8000 feet. 0-2-9.
Beck. (C/. note re site, p. of tuberculata.) Morgan Co.: Weber Can-
yon, 6 miles east of Coalville, heavy loam with humus in shaded
woodland, down to depth of 8 inches, at 5600 feet, July 11, 1952,
0-0-2. Beck. Summit Co.: Wanship, at 5900 feet, about two feet
above stream bed, six inches under sod kept moist by ephemeral
seepage, July 31, 1952, 0-4-16. Beck. Wayne Co.: Elkhorn Ranger
Station. Thousand Lake Mt., at 9700 feet, damp loose black soil,
under rocks and dead timber, and in a forest of quaking aspen, P.
trer?iuloides, August 9, 1952, 0-0-1. Fruita, wet soil near a rock-edge
seep only, January 24, 1951, 0-0-2. Beck.

Remarks. — A. turgida is the fourth species to be recorded herein
for the first time in Utah.

Arizona. — Gila Co.: Sierra Ancha Exp. Forest Headquarters.
Ten miles north of headquarters, at confluence of Rose and Work-
man Creek, top 4 inches of sandy clay loam (with optimum moisture
and an abundandce of leaf mold) at edge of meadow, under walnut,
sycamore and Arizona willow, at 6200 feet, August 29, 1966, 2-11-0.
Barrett.

Remarks. — A. turgida had not been recorded from Arizona.

Montana. — Mineral Co.: Saltese. 3 miles east of Ix)okout Pass
on Route 10, at about 4000 feet, mucky black forest soil near a small
stream and/or sand and gravel mixture by the stream side, June 25,
1954, 0-0-15. Beck.

Remarks. — A. turgida is the fourth species to be rej)orted here-
in for the first time from Montana.

Colorado.— Routt Co.: Along the Yanipa River, ca. 2 miles north
of Steamboat Springs, under stones, boards, logs, and in loose soil

Nov. 30, 1967 DESERT EARTHWORM FAUNA 155

near willow clum|)s in a small meadow. August 27. 1952, 0-0-1.
Be(k.

Remarks. — This is the first record of A. turgida from Colorado.

Bimastos Moore, 1893

Diniastos parvus (Eisen, 1874)

Nevada. — Lander Co.: Austin. 25 miles west on Route 50,
marshland coursing through desert valley, wet organic material,
August 6, 1950. 1-0-2. Beck.

Remarks. — B. parvus is the fifth species to be recorded herein
for the first time in Nevada.

Reproduction, in B. parvus^ is parthenogenetic and at least in
male sterile morphs obligatorily so.

Wyoming. — Fremont Co.: Lander, 44 miles south. Sweetwater
River crossing of Route 287, under old spruce log at sand bar. August
30, 1952. 0-0-5. Beck. Teton Co.: Moran, Teton National Park, under
stones and logs, August 30, 1952, 0-0-3. Beck.

Remarks. — B. parvus is the third species to be recorded herein
for the first time from Wyoming.

Large populations of parvus have not been found anywhere.
Known series are short. Long thought to be native to North America,
the original home of the species is unknown.

B. parvus, the only known North American anthropochore, has
been carried all around the world. More than 67 years ago (Michael-
sen, 1900) the species already had been intercepted at San Francsico
on plants from China. A later interception (Michaelsen, 1910) was
at Hamburg on plants from Japan. Recent interceptions at American
ports, were from soil with plants originating in Taiwan. Japan,
Australia, Mexico. England, and Italy. The species also had been
found in Tibet, Afghanistan. Kazakstan, and St. Paul's Rock (un-
inhabited by man) in the Indian Ocean. The distribution as now
known would seem to indicate that transportation of the species from
this country to other continents and then back and forth around the
world has been much more frequent than carriage within the United
States and has resulted in many more successful colonizations.

Dendrobaena Eisen, 1874
Dendrobaena octaedra (Savingny, 1826)

Colorado. — Boulder Co.: Boulder. Bluebell Canyon, under moss
near spring, September 1914. 0-0-2. (U. S. Nat. Mus.) E. J. Miller.
Boulder City. June 18. 1966. 1-0-3-2. L. Krummholz & W. Osburn
per Beck. Chaffee Co.: Buena Vista. Rancho Sa watch, August 14,
1953. 0-0-2. 0. Sanders.

Remarks. — D. octaedra was recorded for Colorado once before
(Smith, 1917) and from Boulder. Reproduction, in D. octaedra, is

The Great Basin Naturalist

156 G. E. GATES Vol. XXVII, No. 3

parthenogenetic and at least in male sterile morphs obligatorily so.
Polyploidy has been reported.

Nebraska. — Custer Co.: Custer, black soil near creek. June 29,
1946, 0-0-1. .1. L. Macnab per D. McKey-Fender.

Remarks. — D. octaedra, previously unreported from Nebraska,
is the eighth species to be recorded from that state. Others are: A.
caliginosa, B. parvus, E. foetida, Pheretima diffringens, and three
diplocardias of uncertain status.

Dendrobaena rubida (Savigny, 1826)

Idaho. — Butte Co.: Arco, to the south about 1 mile, heavy black
clay loam at water's edge and a more loose and only slightly moist
soil a few feet above the water's edge, May 31, 1952, 0-0-2. Beck,
Mary Clark, Grace Grant.

Remarks. — D. rubida is the fifth species to be reported herein
for the first time from the state of Idaho.

Reproduction, in D. rubida, at least for male sterile morphs is
amictic. Amphimixis seems to be optional in sperm maturing
morphs. Polyploidy has been reported. Only male sterile morphs
were recognized in desert states.

Nevada. — Eureka Co.: Eureka, 10 miles east, 6500 feet, heavy
clayey soil near a mountain spring, August 6, 1950, 0-1-8. Beck.

Remarks. — D. rubida is the sixth species to be reported herein
for the first time from the state of Nevada.

Utah. — Box Elder Co.: Lucin, at a desert spring, seven miles to
the north, where water was salty and vegetation was of the alkaline
desert type, June 19. 1952, 0-0-1. Beck. Clark Co.: Birch Creek, June
3. 1952, 0-0-1. Beck. Juab Co.: Tom's Creek, Callao, moss at stream
side, August 12, 1953, (18?) -0-4. Beck. San Juan Co.: Kigalia Ranger
Station. Bears Ears, June 9, 1955. 4-9-3. Beck. Sanpete Co.: Mt.
Pleasant, Pleasant Creek Picnic Grounds, beneath bark and under
fallen trees at contact line of as])en and conifers. August 5. 1951,
0-0-9. Beck. Summit Co.: Echo. July 11, 1952, 0-0-3. Beck.

Remarks. — D. rubida is the fifth species to be reported herein
for the first time from Utah.

Wyoming. — Fremont Co.: Wilderness. 20 miles south of Dubois,
in moist soil with much humus, under short grass turf at edge of
Deriwoodie River, August 30. 1952. 0-0-7. Beck. Teton Co.: Teton
National Park, about 5 miles north of the park boundary where
Snake River crosses Route 287, sodded bank of river (where bait had
been dumped at end of day's fishing?). August 31, 1952, 0-0-1.
Moran. river bank and bed about five miles to the east. August 30.
1952. 0-0-5. Beck.

Remarks. — D. rubida is the fourth species to be recorded herein
for the first time in Wyoming.

Elsewhere, in the region now under consideration. D. rubida has
been reported from Colorado (Smith, 1917) but under another name
and without specification as to county or town.

Nov. 30, 1967 DESERT EARTHWORM FAUNA 157

Eisenia Malm, 1877
Eisenia foetida (Savigny. 1826)

California. — Los Angeles Co.: Temple City, March 1957. 0-4-1.
P. W. Oman. San Bernardino Co.: Fontana, March 1953, 7-1-12. G.
E. Temi)leton |)er E. W. Price. March 1957. 3-4-2. P. W. Oman. San
Diego Co.: Jacuml)a. March 9. 1966. 0-8-34. Beck. (Purchased from
a bait dealer who obtained them from West Virginia. In this resort
city no earthworms were found in the vicinity of the hot springs,
near a lake, and other spots in and around the town. Local anglers
informed Beck that there are no earthworms locally.)

Remarks. — Previous Californian records of E. foetida were all
from a northern part of the state.

Reproduction, in E. foetida, is obligatorily amphimictic.

Utah.— Utah Co.: Provo, April 12, 1951, 0-1-0. Beck.

Remarks. — E. foetida is the sixth species to be reported herein
for the first time from the state of Utah.

The single specimen may have been an escapee from an artifical
habitat such as the bed of an earthworm farm, a greenhouse, a
potted plant, a sewage bed.

Arizona. — Maricopa Co.: Tempe. March 4, 1966, 0-0-55. Beck.
(Purchased from an earthworm farm. The species was being sold for
bait to produce mulch for gardens, also for introduction into worm-
less areas to enrich the soil.)

Remarks. — E. foetida had not been recorded previously from
Arizona. If presence in artificial habitats such as greenhouses and
earthworm farms is to be counted the species will have to be re-
corded from every one of the 50 states with the possible exception
of Alaska, as E. foetida, under one or more of its popular names, has
been sold for cultivation in every state and Canadian province, and
was recently introduced to Hawaii. Although sold in large numbers
throughout the country and transported back and forth, records for
natural habitats are lacking for many states.

Eisenia rosea (Savigny, 1826)

California. — Kern Co.: Wasco, with fibrous root system of
Bermuda grass at edge of a stagnant pool in an irrigation ditch,
August 21. 1950. 0-0-1. P. D. Pilsbury per Beck. Los Angeles Co.:
Torrance, slightly damp ground at edge of water of a marsh, August
29. 1950, 2-9-5. and 6 inches of moist organic soil above sand in a
flower bed. August 29, 1950. 10-0-1. Beck. San Diego Co.: Pine
Valley. .June 21. 1966. 0-0-45. E. V. Komarek. Sr.

Remarks. — E.. rosea is now reported for the first time from
southern California. There are older records for a northern part of
the state.

Reproduction, in E. rosea, is parthenogenetic and at least for the
many male sterile mor[)hs obligatorily so.

The Great Basin Naturalist

158 G. E. GATES Vol. XXVII, No. 3

Idaho. — Bannock Co.: McCammon, 3 inches below surface and
in an area ca. 2 feet square, a few feet from bank of Portneuf River
and about a foot above water level. May 30, 1952, 0-0-3. Idaho Falls,
September 2, 1952, 5-2-12. Beck. Butte Co.: Arco, to the south about
1 mile, heavy black clay loam at water's edge and a more loose and
only slightly moist soil a few feet above the water's edge, May 31,
1952. 0-0-3. Mary Clark, Grace Grant and Beck.

Remarks. — E. rosea is the sixth species to be recorded herein
for the first time from Idaho.

Nevada. — White Pine Co.: Kennicot Duck Creek Station, one
mile to the south, wet, heavy, clay soil with organic matter, stream
side, August 5, 1950, 2-1-6. Beck.

Remarks. — E. rosea is the seventh species to be recorded herein
for the first time from Nevada.

Utah.^ — Beaver Co.: Beaver, soil of a peat-like consistency near
a small stream, July 11, 1952, 1-1-0. Beck. Piute Co.: Kingston, bank
of ditch at base of cHff in desert at 6100 feet, June 26, 1952, 0-2-6.
Beck. Rich Co.: Sage Creek Junction, stream bank of a creek, only
with wet soil and fibrous grass roots, at 6500 feet, August 21, 1952,
1-0-0. Beck. Sanpete Co.: Mt. Pleasant, August 5, 1951, 0-0-1. Beck.
Summit Co.: Wanship, at 5900 feet, about two feet above stream
bed, six inches under sod kept moist by ephemeral seepage, July 13,
1952, 0-1-1. Beck. Utah Co.: Soldier Summit, upper three inches of
heavy clay under cardboard and wooden boxes of an abandoned
garbage dump. May 22, 1952, 0-0-1. Beck. Washington Co.: St.
George, almost pure clay at edge of an irrigation ditch from Virgin
River, September 2. 1950, 5-0-0. Zion National Park, detritus and
under stones among one of the side streams, near Weeping Rock,
Virgin River. September 3, 1950. 0-2-3. Beck.

Remarks. — E. rosea is the seventh species to be recorded herein
for the first time from Utah.

Arizona. — Cochise Co.: 16 miles northeast of Douglas on Leslie
Canyon-Rucker Road, y^ mile east of road near cattle pen at 4600
feet, from sod and upper four inches of black, heavy clay loam nearly
saturated by four days of rain (no farms and permanent streams in
the area), September 16, 1966, 7-0-1. Three miles south of junction
of Leslie .Canyon and main, east-west Rucker Canyon Road, along
roots and iii top four inches of sandy to gravelly clay loam by aban-
doned water storage tank (for cattle, no farms and fishing streams
nearby), September 16. 1966. 40-1-3. Barrett per Beck. (Remote and
infrequently visited areas without streams, no fishing streams near
sites.) Tombstone, Costello Ranch, at elevation of ca. 5000 feet, in
soil (pIT 7.5) rich with manure. July 23. 1933. 0-0-15. A. Petrunke-
vitch per G. E. Pickford. Leslie Canyon. McNeal. upper four inches
of sandy to clay loam along a small stream (no fishing), under Ari-
zona Willow. September 3. 1966. 6-0-0-3. Barrett. McNeal. March
19. 1966, 2-0-0. Barrett per Beck. Coconino Co.: Payson. June 29,
1966, 4-1-3. Barrett per Beck. Gila Co.: Sierra Ancha Exp. Forest

Nov. 30, 1967 DESERT EARTHWORM FAUNA 159

Headquarters. Two miles north of the Headquarters along almost
dry creek in sandy loam, under walnut and oak, August 29, 1966,
1-0-0. Ten miles north of Headquarters, at confluence of Rose and
Workman Creeks, at edge of meadow, under walnut, sycamore and
Arizona Willow, at ca. 6200 feet, August 29, 1966, 1-0-0-1. Top 2
inches of loam to sandy loam with 4-5 % organic matter, under maple
(but not pine and fir), within 10 feet of bank of Workman Creek
(none in region 10 to 50 feet from bank), at 6900 feet, August 30,
1966, 2-1-0. Barrett. Maricopa Co.: Gila Bend, March 8, 1966, 8-3-0.
Beck. Wickenburg, March 8, 1966, 6-1-0. Tempe, March 5, 1966,
0-14-19. Barrett and Beck. Mesa, March 6. 1966, 6-6-5-1 + one tail
piece and 0-2-1. X. Frost, Beck. March 4, 1966, 0-2-1. Beck. Mohave
Co.: Littlefield. rich soil with about 50% humus, under bridge on
Route 91 over Beaver Dam Creek, Septemper 2, 1950, 6-5-2. Beck.
(The only species secured at the site where worms "were not too
numerous and required diligent searching" to find these.) Pima Co.:
Tucson, June 18, 1966, -13-0 + 17 tail pieces, Beck. (At least 12
specimens were early adolescents i.e., aclitellate.) June 18, 1966,
7-0-17. E. V. Komarek, Sr. Santa Clara Co.: Sondita Creek, Pata-
gonia, June 19. 1966. 0-1-3. E. V. Komarek. Sr.

Remarks. — E. rosea already had been collected in the state be-
fore 1900. Nevertheless, Arizona is one of the states in which Seton
(1929) said he could find no earthworms.

Spaniards could have brought live plants to their missions from
some time in the early 1600's.

Montana. — Mineral Co.: Saltese. 3 miles east of Lookout Pass
on Route 10, at about 4000 feet, mucky, black forest soil near a small
stream or in a coarse sand and gravel mixture by the stream side,
June 25. 1954. 0-0-1. Beck.

Remarks. — E. rosea is the fifth species to be recorded herein for
the first time from Montana.

Wyoming. — Fremont Co.: Lander, soil with humus along banks
of Papaogie Creek, August 30. 1952, 0-0-1. Beck.

Remarks. — E. rosea is the fifth species to be recorded herein for
the first time from Wyoming.

Colorado. — Larimer Co.: Fort Collins. August 29. 1952, eight
inches of dark, compact, sandy loam above water level in seepage
area at base of South Reservoir, August 29. 1-0-10. Beck. Routt Co.:
Along the Yanipa River, ca. 2 miles north of Steamboat Springs,
under stones, boards, logs, and in loose soil near willow clumps in
a small meadow, August 27. 1952, 0-0-5. Beck.

Remarks. — These are the first records of E. rosea for Colorado.

New Mexico. — Chaves Co.: Rio Felix, south of Dexter. February
25. 1966. 0-40-0. Beck.

Remarks. — This is the second species to be recorded herein for
the first time from New Mexico. This is one of those states in which
Seton (1929) himself could find no earthworms.

The first permanent white settlement was by the Spanish in 1610.

The Great Basin Naturalist

160 G. E. GATES Vol. XXVII, No. 3

Eiseniella Michaelsen, 1900
Eiseniella tetraedra (Savigny, 1826)

California. — Kern Co.: Deep Creek, Mojave desert, April 25,
1943, 0-0-2. per D. McKey-Fender. Wasco, residence of P. D. Pils-
bury, dark wet earth under a hedge and around outlet of an air-
conditioning hose under a hedge, August 21, 1950, 0-0-3. Beck. With
fibrous root system of Bermuda grass at edge of a stagnant pool in
an irrigation ditch, August 21, 1950, 0-0-1. P. D. Pilsbury per Beck.
Santa Barbara Co.: Santa Barbara, March 25, 1966, 0-0-3 and 1 tail
piece. Beck. San Bernardino Co.: Victorville, one-half mile west of
bridge over Mojave River, in wet, coarse sand under algal compost,
September 1, 1950, 0-0-1 (Hercynian morph). Beck.

Remarks. — These are the first records of E. tetraedra for the
southern part of the state.

Reproduction, in E. tetraedra, is parthenogenetic and often is
obligatorily so because of male sterility. Although male sterile
morphs of many species had been studied and named by oligochae-
tologists, the sterility for long was not even suspected. Megadrile
parthenogenesis was first proved experimentally in E. tetraedra by
raising isolated hatchlings to maturity in the laboratory.

Idaho. — Butte Co.: Arco, to the south about 1 mile, heavy black
clay loam at water's edge and a more loose and only slightly moist
soil a few feet above the water's edge. May 31, 1952, 2-1-16. Beck,
Mary Clark, Grace Grant. Canyon Co.: Middleton, four inches be-
low surface of an area with sod where logs and other detritus had
accumulated near Boise River, June 19, 1954, 0-0-1. Beck. Clark
Co.: Birch Creek, ca. 15 miles south of Lemhi (bunty border, wet to
moist sand and under stones at stream's edge, at 6500 feet, June 3,
1952, 0-0-1. Beck. Custer Co.: Challis, several miles to the south, at
ca. 6500 feet, moist to wet soil and under rocks at side of a small
mountain streamlet. May 31, 1952, 20-12-52. Mary Clark, Grace
Grant, Beck. Kootenai Co.: Worley, under logs and other debris at
bank of a stream that ran into a marsh land. June 24, 1954, 1-4-13.
Beck. Lemhi Co.: Salmon, 11 miles to the north, ca. 4100 feet, from
fibrous root system of grassy turf of a relatively dry meadow near
Salmon River, June 1, 1952, 0-0-1. Beck. Mary Clark, Grace Grant.
Hill side seep spring area, on Route 93. 1 mile north of town. May
31, 1952, 0-0-2. Beck. Lincoln Co.: Shoshone Falls, beneath rocks at
edge of pools, streams and springs, June 19. 1954, 0-0-24. Beck.

Remarks. — E. tetraedra is the seventh species to be recorded
herein for the first time from Idaho.

Nevada.— Lyon Co.: Wabuska, July 5. 1966. 0-0-25. E. V. Ko-
marek, Sr. Ormsby Co.: Carson River, south of Silver Spring. Julv 5.
1966. 0-0-20. E. V. Komarek. Sr. White Pine Co.: McGill. drain
north of a swimming pool which is northwest of the town, August
5, 1950, 1-1-4. Beck. Kennicot Duck Creek Station, one mile to the

Nov. 30, 1967 DESERT EARTHWORM FAUNA 161

south, at stream edge, wet heavy clay soil with organic matter,
August 5. 1950. 1-2-23. Beck.

Remarks. — E. tetraedra is the eighth species to be recorded
herein for the first time from Nevada.

Utah. — Beaver Co.: Wahwah Springs, September 12, 1952,
moist soil along stream bank, and under rocks near the marshy area,
5-0-12. under rocks near edge of desert spring and its run-off, 0-0-3.
Beck. Box Elder Co.: Lucin, seven miles to the north, at a desert
spring where water was salty and vegetation was of the alkaline
desert type. June 19, 1-0-0. Beck. Washington Co.: Zion National
Park, at 6000 feet, under plant cover of sage and iuniper, near
Uintah Mountains. June 12, 1955, 0-0-1. Beck. Piute Co.: Marysvale,
2 miles south of Big Rock Candy Mountain on Route 89, with
fibrous root system of grass and sedge quite close to water level in
Sevier River. June 26. 1952. 0-1-15. Beck. Rich Co.: Randolph,
abundant along bank of Otter Creek, at 6600 feet, August 21. 1952,
0-0-9. Beck. Sanpete Co.: Mt. Pleasant, Pleasant Creek Picnic
Grounds, at contact line of conifers and aspens, beneath bark and
under fallen trees. August 5, 1951, 0-0-9. Beck. Utah Co.: Mt. Nebo,
at 8600 feet, under stones, logs, boards near a bog on north side of
mountain on the Loop Road. September 21, 1952, 0-0-15. Lehi Sugar
Factory Area, in wood of a rotted board near pond. April 25, 1952,
0-0-1. Beck. (Elsewhere in the area no worms were found.) Wash-
ington Co.: St. George, almost pure clay at edge of an irrigation ditch
from Virgin River, September 2, 1950, 1-0-6. Beck.

Remarks. — E. tetraedra is the eighth species to be recorded here-
in for the first time from Utah.

Montana. — Mineral Co.: Saltese, 3 miles east of Lookout Pass
on Route 10. at about 4000 feet, mucky black forest soil near a small
stream or in a coarse sand and gravel mixture by the stream side,
June 25. 1954. 0-0-1. Beck. Ravali Co.: Llamilton, four miles west
and near Blodget Creek at 3600 feet, upper 3-4 inches of black soil
mixed with sand under Pinus ponderosa, June 1. 1952, 1-0-0. Beck.

Remarks. — E. tetraedra is the sixth species to be recorded here-
in for the first time from Montana.

Wyoming. — Carbon Co.: Spring Creek. 2 miles south of Saratoga
on Route 130. June 7, 1961. 0-0-2. (Hercynian morph). North Brush
Creek. 1 mile west of Medicine Bow Natural Forest Line on Route
130, June 7. 1961. 0-0-1. G. F. Edmunds & W. L. Peters. Fremont
Co.: Lander, sandy soil near Papaogie Creek at south side of city
park. August 30. 1952, 1-1-11 (of which 1-1-8 are of Hercynian
morphs). Beck. Yellowstone National Park. September 1. 1952,
0-0-5. Beck.

Remarks. — E. tetraedra is the sixth species to be recorded here-
in for the first time from Wyoming.

Colorado. — Chaffee Co.: Buena Vista. Rancho Sawatch. August
14. 1953. 0-0-6. Ottys Sanders. Grand Co.: Hot Sulphur Springs,
under stones along edge of Colorado River south and west of the

The Great Basin Naturalist

162 G. E. GATES Vol. XXVII, No. 3

town about seven miles, August 28, 1952, 0-4-38 (0-4-32 of the
Hercynian morph). Beck. Larimer Co.: Fort Collins, August 29,
1952. eight inches of dark, compact sandy loam above water level in
seepage area at base of South Reservoir, 2-0-0. Beck. Routt Co.: Along
the Yanipa River, ca. 2 miles north of Steamboat Springs, under
stones, boards, logs, and in loose soil near willow clumps in a small
meadow, August 27, 1952, 0-0-5 (1 of Hercynian morph). Beck.
Summit Co.: Frisco, one mile to the northeast, August 13, 1953,
0-0-2. Ottys Sanders.

Remarks. — These are the first records of £. tetraedra for named
Colorado localities. The species was recorded (Smith, 1917) for the
state but without specification as to county or otherwise.

South Dakota. — Lawrence Co.: Spearfish Creek, 0.8 miles
south of hydro plant no. 2, June 9, 1961, 0-0-1. G. F. Edmunds &
W. L. Peters.

Remarks. — Although earthworms are known to have been
present in South Dakota for some years (c/. Taylor, 1924) the record
above is the first in print for an identified species.

Dakota, according to Seton (1929), had no earthworms. The
northern state probably had little European settlement till after
1862. First European settlers, other than traders and trappers, ar-
rived in the southern state in 1850.

Taylor stated that earthworms "do not thrive in acid soils such
as are found in the great plains section of North America .... They
are generally plentiful in silt soils along streams .... The most
productive soil I have ever planted was full of them, always was, and
is today. The only fertilizer it ever gets is silt from over-flow of
the river, and the same land has been farmed for over seventy
years."

The soil of 1.310 square miles in 6 counties of eastern South
Dakota is unusually interesting. The average depth is 36 inches in
an area with an east-west width of 16-32 miles and a north-south
length of 75 miles. Because that soil consists almost entirely of worm
casts and filled worm channels it was named (Buntley & Papen-
dick, 1960) Vermisol. Worm activity had almost completely
destroyed horizontal zonation.

Like Darwin (in the title of his last book. "The formation of
vegetable mould through the action of worms") Buntley & Papen-
dick presumably expected us to deduce that worms = earthworms
as they did not once use that word. A request for information
brought the following reply (Buntley. in lit. Oct. 18, 1960), "I did
collect some worms from each of the two worm-worked soils. These
were identified as Lumhricus terrestris by our entomology depart-
ment."

Euroy)ean settlement in South Dakota began after 1850. Less than
a hundred years (c/. dates. 1966) was probably available for the
destruction, by earthworms, of soil zonation to an average depth of
three feet. At first that does seem like too much to expect even of

Nov. 30, 1967 DESERT EARTHWORM FAUNA 163

the kind of animals so highly praised by organic gardeners. L. ter-
restris, according to Olson (1928, p. 59), "has become widely dis-
tributed over Ohio in the last ten years." Such spreading capacity
presumably could enable the required Dakota distribution through
a much smaller area in less than a hundred years. However, the
identity of the species involved is by no means certain. Far too
often, even today, Lumbricus terrestris is THE earthworm. Con-
versely any earthworm easily becomes L. terrestris. Within the last
twelve years, Eisenia foetida and AUolohophora tuberculata have ap-
peared as Lumbricus terrestris in scientific journals, probably also
A. trapezoides and longa and one or more other megadrile species the
identity of which cannot even be guessed.

Evidence supporting a rapid rate of zone-destruction by earth-
worms may be unpublished in field notes of various pedologists. Be-
tween the spring of 1958 and 1961, according to K. K. Langmaid
{in lit.). Head of Canadian Soil Survey, soil horizons (at a Carling-
ford. New Brunswick, cut-over forest site) had been obliterated by
earthworm action, to a depth of six inches. The species in the sample
from that site were Allolobophora tuberculata (4 specimens), Lum-
bricus terrestris (1).

The Dakotas unfortunately were not visited by Beck. From the
northern state two species have been recorded, Eisenia foetida from
earthworm farms, Microscolez phosphoreus from a greenhouse.
Nothing at all is known about megadrile populations of natural habi-
tats.

Lumbricus Linnaeus, 1758

Lumbricus castaneus (Savigny, 1826)

Idaho. — Lemhi Co.: Bitterroot River, near continental divide on
Route 93, June 1, 1952, 0-0-1. Beck.

Remarks. — L. castaneus is the eighth species to be recorded here-
in for the first time from Idaho. The species appears to be much
less important in the western faunas of the United States (c/. below).
In central Maine, on the contrary, castaneus is fairly common but
rubellus is rarely found.

Reproduction, in L. castaneus, is obligatory biparentai.

Lumbricus rubellus Hoffmeister, 1843

Idaho. — Fremont Co.: "Osborne Springs," ca. 25 miles north
of Ashton on Route 191, heavy black loam. 6-8 inches below surface,
in process of "balling." surface quite dry. September 1. 1952, 3-3-4.
Beck. Lincoln Co.: Shoshone Falls, damp porous black soil, under
rocks in grassy area and 2-8 inches below surface of soil, June 18.
1954, 0-0-1.

Remarks. — L. rubellus is the ninth species to be recorded herein
for the first time from Idaho.

The Great Basin Naturalist
164 G. E. GATES Vol. XXVII, No. 3

Utah. — Weber Co.: Ogden, depressions in the flood plain where
plant growth is rank, September 3, 1952, 0-0-2. Beck.

Remarks. — L. rubellus is the ninth species to be recorded herein
for the first time from Utah.

Colorado. — Routt Co.: Along the Yanipa River, ca. 2 miles
north of Steamboat Springs, under stones, boards, logs, and in loose
soil near willow clumps in a small meadow, August 27, 1952, 1-1-10.
Beck.

Remarks. — This is the first record of L. rubellus for Colorado.

Reproduction, in L. rubellus, is obligatorily amphimictic.

Lumbricus terrestris L., 1758

Idaho. — Bannock Co.: Pocatello, June 3, 1952, 0-0-1. Beck. Latah
Co.: Moscow, Palouse soil in lawns of University of Idaho, March
15, 1950, 0-0-2. C. W. Lame.

Remarks. — L. terrestris is the tenth species to be recorded here-
in for the first time (by its scientific name after a definite identifi-
cation) from Idaho.

Writing about "night crawlers," Painter (1942) said, "Many
people in Twin Falls, Idaho get night crawlers for bait from lawns
by electricity. The worms supposedly had been imported originally
from the East." Night crawler is a common name for Lumbricus
terrestris. Nevertheless, use of that common term is no guarantee
that an identification as terrestris would be correct. The fact that
electricity was used to secure the worms itself is a contra-indication.
Night crawlers, when conditions are favorable, feed and copulate on
the surface at night where they are easily "jacked," without elec-
tricity, by those familiar with their habits. Millions are easily col-
lected that way every year in this country. If conditions are un-
favorable for surface activity, the crawlers are likely to be too deep
down to be brought out by a superficial electric current. American
allolobophoras, on the contrary, are not known to feed on the sur-
face at night. They are geophagous and more likely to be active at
soil levels where they could be stimulated by electricity.

Utah.- — Morgan Co.: Weber Canyon, at 5600 feet, about 6
miles east of Coalville, July 11, 1952, 0-0-1. Beck. Summit Co.:
Wanship, at 5900 feet, two feet above stream bed. six inches under
sod kept moist by ephemeral seepage, July 13. 1962, 4-0-1. Beck.
Utah Co.: Provo, April 12 or May 13, 1951. 0-0-1. Beck. Washington
Co.: Pine Valley, July 16, 1953. 0-0-2. Beck.

Remarks. — L. terrestris is the tenth species to be recorded here-
in for the first time from Utah. The paucity of records above, sur-
prisingly, may be due to the commonness and usefulness of the
species. Beck remembers watching in the fall of 1925 "jacking" of
night crawlers in the long practiced manner. By that time the
species must have been common around Provo at least. Today, Beck
reports, along the highways of the state one sees numerous signs
signifying night crawlers for sale. lie also mentioned that " jacking"

Nov. 30, 1967 DESERT EARTHWORM FAUNA 165

Still is a nightly summer activity unless a fastidious lawn owner
has treated his grassland with Pax or chlordane.

Colorado. — Grand Co.: Hot Sulphur Springs, moist, loose dark
soil with fibrous grass roots along back of Colorado River about 7
miles southwest of the town, August 28, 1952, 9-0-1. Beck. Routt
Co.: Along the Yanipa River, ca. 2 miles north of Steamboat Springs,
under stones, boards, logs and in loose soil near willow clumps in a
small meadow. August 27. 1952, 1-0-3. Beck.

Remarks. — L. terrestris was recorded from Colorado once before
(Smith. 1917) but without specification as to places or sites from
which the worms were obtained.

Iowa. — Jefferson Co.: Fairfield. Parsons College campus, May
22, 1951: Mud flats of a small stream on north side, 1-0-1. Wooded
slope southwest of flats, 2-0-0. F. W. Von Ohlen.

Remarks. — Reproduction, in L. terrestris, is obligatorily bi-
parental.

Octolasion Oerley, 1885

Octolasion cyaneum (Savigny, 1826)

Colorado. — Larimer Co.: Stonewall Creek, 30 miles south of
Wyoming border on Route 287. loose sandy soil with much humus,
in sodded area where willow growth was rank, August, 1952, 0-0-1.
Beck.

Remarks. — This is the second trans-Mississippi record of the
species which is but rarely found anywhere in the American hemi-
sphere.

Reproduction, in O. cyaneum, is obligatorily parthenogenetic.
Polyploidy has been reported.

Octolasion tyrtaeum (Savigny, 1826)

Idaho. — Butte Co.: Arco. to the south about 1 mile, heavy black
clay loam at water's edge and a more loose and only slightly moist
soil a few feet above the water's edge. May 31, 1952, 0-0-1. Beck,
Mary Clark. Grace Grant.

Remarks. — O. tyrtaeum is the eleventh species to be recorded
herein for the first time from Idaho.

Reproduction, in O. tyrtaeum, is obligatorily amictic. Male steril-
ity is common. Polyploidy has been reported.

Utah. — San Juan Co.: Elkhom Ranger Station. Thousand Lake
Mt., at 9700 feet. damp, loose black soil, under rocks and dead timber
and in a forest of quaking aspen, P. tremuloides, August 9. 1952,
0-0-1. Beck.

Remarks. — O. tyrtaeum is the eleventh species to be recorded
herein for the first time from Utah.

Colorado. — Chaffee Co.: Buena Vista. Rancho Sa watch. Augiist
14. 1953. 0-0-1. Ottys Sanders. Larimer Co.: Stonewall Creek. 3 miles
south of Wyoming border on Route 287, loose sandy soil with much

The Great Basin Naturalist

166 G. E. GATES Vol. XXVII, No. 3

humus, in sodded area where willow growth was rank, August 29,
1952, 0-1-16. Beck

Remarks. — Early settlers, according to Cockerell (1924), main-
tained they found no earthworms in mountainous districts or in
adjacent upland plains of the state.

D. tyrtaeum was reported once from Colorado, as O. lacteum.
The same author (Smith, 1917) recorded for the state the following
valid species: A. chlorotica, D. octaedra and rubida, E. tetraedra, L.
terrestris. Except for D. octaedra which had been taken only at
Boulder, no data as to counties, sites and number of specimens was
provided.

Nebraska. — Laurence Co.: Hickman, June 11, 1960, 0-0-3. W.
R. Murchie.

Remarks. — O. tyrtaeum previously had not been recorded from
Nebraska, even as its synonym, O. lacteum.

Species previously reported from th state are seven: A. caliginosa
(now known to have been a complex of four or more species), B.
parvus, E. foetida, Pheretima diffringens, and three diplocardias of
uncertain status. The single record of P. diffringens, soil near Uni-
versity greenhouses at Lincoln, probably is indicative of the way this
oriental species reached the state.

"Settlement began in a small way in 1854," according to P. W.
Gates (in lit.) but increased considerably after 1862 when a Federal
Homestead Act provided free land for veterans.

Iowa. — Sioux Co.: Ireton. under maple trees in yard near farm-
house, 5 miles from the village, April 21, 23-3-25. Mary M. Jinks.

Remarks. — O. tyrtaeum was recorded in 1948 from Iowa as O.
lacteum. Other megadrile records for the state are: S par ganophilus
eiseni (a North American endemic that is widely distributed), Allo-
lobophora trapezoides, A. caliginosa (probably A. turgida), Eisenia
foetida and rosea. Lumbricus terrestris, and Octolasion cyaneum.

There was, according to historian P. W. Gates {in lit.), "Not
much settlement before 1836-1938."

Megascolecidae

Pheretima Kinberg, 1866

Pheretima hawayana (Rosa, 1891)

California. — Los Angeles Co.: Los Angeles, lawn, June 1955,
a number of clitellate individuals. A. W. Bell.

Remarks. — Reproduction, in P. hawayana. probably is obliga-
torily amphimictic. The original home of the species is in eastern
Asia.

Pheretima hupeiensis (Michaelsen, 1895)

Utah. — Utah County: Provo, Rrigham Young University cam-
pus, January 31. 1966, 0-2-14. Beck.

Nov. 30, 1967 DESERT EARTHWORM FAUNA 167

Remarks. — P. hupeiensis is the twelfth species to be recorded
herein for the first time from Utah. The sj)ecies had been recorded
{)reviously from Louisiana but nowhere else west of the Mississippi
River.

The following information about this species was provided by
Beck {in lit., June 20. 1966): liach spring and fall and at periods of
warmth, which occurred in January of this year, these worms
crawled from the lawns by the thousands to the sidewalks, creating
quite a squishy situation for the students walking back and forth
from the buildings.

Rej)roduction, in P. hupeiensis, is parthenogenetic, probably
often because of male sterility.

Directly or indirectly, the species may have come from China.

Ocnerodilidae

Ocnerodrilus Eisen, 1878

Ocnerodrilus occidentalis Eisen, 1878

This species has been reported occasionally from the desert
region. Types were found in California and of course the species was
thought to be native to that state. Somewhat later, specimens from
China and Hawaii, were intercepted at San Francisco. They were
obtained from soil with potted plants. Presumably the species had
been transported around the world for several centuries. Even today,
the original home of O. occidentalis is unknown. Central Africa and
tropical America have been postulated.

Absence of specimens in recent collections may be of little or no
significance. However, colonization could have been temporary.
Similar failures to achieve permanent domicle are believed to have
occurred in the faunas of oceanic islands such as Hawaii and St.
Helena.

Reproduction is parthenogenetic.

Appendix

No collections were available for several states that need consid-
eration herein. Some of Professor Beck's collections were lost in
transit which is much regretted because of our ignorance of earth-
worm faunas.

Kansas. — Earthworms originally were lacking in Kansas ac-
cording to Wilcox (1884). There was little European settlement till
1851. By 1917. Smith was able to report three species of Bimastos,
parvus, another now known as tumidus. and welchi. known only
from the original description based on sections of the single type.
The latter, according to Smith, was immature. None of the three
are native to Kansas. A specimen of a double-tailed worm from
Kansas (Hamly, 1932) was said to be of Lumhricus terrestris but
that too often, even today, means nothing more than "earthworm."

The Great Basin Naturalist

168 G. E. GATES Vol. XXVII, No. 3

Oklahoma. — Earthworms were lacking in the prairies of Indian
Territory according to Wilcox (1884). Since 1954 the following were
reported from Oklahoma, Allolobophora caliginosa (most probably
in large part if not wholly, A. trapezoides) , Eisenia foetida and rosea,
each of which is present in desert states, Bimastos tumidus (the
original home of which is unknown), and four diplocardias of uncer-
tain status, three of which were not reported from Missouri and
Arkansas.

Minnesota. — Lumbricus terrestris was the first earthworm
recorded (Smith, 1917) for Minnesota but as usual without data to
localities and number of specimens. Mickel (1925), who studied
earthworm parasites, provided a little information. A. caliginosa was
the most abundant earthworm at St. Paul. L. terrestris was the sec-
ond most abundant. E. foetida was found only in compost and then
not commonly. Mickel's caliginosa is less likely to have been trape-
zoides than tuberculata or turgida or both. Except for those parasites,
presence of L. terrestris would be the extent of our knowledge of
the earthworm fauna of Minnesota in 1967. E^ch Minnesota species
is present in the desert states.

Missouri. — The megadrile fauna of this state comprises the
following: Allolobophora caliginosa (perhaps in part not A. trape-
zoides), chlorotica and trapezoides, Bimastos heimburgeri (of uncer-
tain status), tumidus and zeteki. Dendrobaena rubida, Eisenia foetida
and rosea, Eiseniella tetraedra, Lumbricus rubellus and terrestris,
Octolasion tyrtaeum of the Lumbricidae, Pheretime agrestis of the
Megascolecidae, Sparganophilus eiseni and 5 diplocardias of uncer-
tain status four of which also are recorded from Arkansas.

Nine of those species are present in desert states, 10 obviously
are exotic. Original homes of the diplocardias. species of Bimastos
and S. eiseni are unknown but now seem unlikely to have been in
Missouri.

Arkansas. — Megadriles recorded for this state are: A. caliginosa
(probably in large part, if not all, A. trapezoides) , Bimastos parvus
(in which beddardi and longicinctus may have to be included),
tumidus and zeteki, Dendrobaena octaedra and rubida. Eisenia foeti-
da, hortensis (recorded as B. venetus), and rosea, Eiseniella tetraedra,
Lumbricus rubellus and terrestris, Octolasion tytraeum, Pheretima
californica, diffringens and hupeiensis, four diplocardias of uncertain
status.

Eleven of those species are present in desert states. Thirteen ob-
viously are exotic, and three pheretimas being of Asiatic origin, the
others of European origin. Species of Bimastos, as well as one diplo-
cardia, are unlikely to have originated in Arkansas.

Discussion

A sample of Httle more than 2500 worms may seem too small to
warrant discussing the megadrile fauna of an area containing more
than a million square miles. As a result, mainly of the activity of

Nov. 30, 1967 DESERT EARTHWORM FAUNA 169

Prof. Beck, we do know much more about the earthworms of the
desert area than before. Each of the following pairs indicates num-
ber of species previously and now known from the mentioned states.
Arizona 3-8. California (southern part only) 1-7, Colorado 6-12,
Idaho (one small section excluded) 0-11 (with no exclusion, 1-12),
Iowa 8-8, Montana 0-6, Nebraska 7-8, Nevada 0-13. New Mexico 0-2,
North Dakota 2-2, South Dakota 0-1, Utah 0-12, Wyoming 0-6.
Numbers of species known from adjacent states are: Kansas 3, Okla-
homa 8. Minnesota 3, Missouri 19. Arkansas 20.

Number of species now known from desert states is 21, larger
than the number recorded for anyone of the adjacent states with
much more rainfall (cf. Table 1).

Every one of those 21 species obviously is e.xotic in the area,
most of the lumbricids having come from Europe, one probably from
somewhere east of the Mississippi, and megascolecids from Asia.
The original home of the ocnerodrilid is unknown, but is unlikely
to have been in North America. Prof. Beck has found absence of
endemics in such a large area hard to believe, as will many others.

Exotics could have gotten to the desert region in two ways. The
first of course is by migration. This is the view of some Europeans
(cf Omodeo. 1963). European species moved across the north Atlan-
tic to Greenland, Iceland and eventually to the American mainland.
There, they alone survived the great glaciations that pauperized,
perhaps ])ermanently, the megadrile fauna of North America. Geolo-
gists seem to be unaware of proof that an Atlantic bridge existed at
the proj)er time (cf. Wright & Frey, 1965). Survival of soft-bodied,
moisture-loving animals through countless millenia on bare nunataks
seems unlikely.

The other way by which the exotics could have gotten to desert
states is by introduction. That is likely to have resulted only from
human activity. Supporting evidence for introduction is of two sorts.
Data of the first sort got into print as a direct result of the publication
in 1881 of Darwin's last book, "The formation of vegetable mould
through the activity of worms." Parenthetically it is well to note
that the worms are earthworms. The conditions Darwin thought re-
sulted only from megadrile activity existed in the great plains of
Canada where there were then no earthworms. Seton (1929) con-
cluded his study of the subject in the following words:

"Since 1882. I have made personal investigations in parts of
Saskatchewan. Alberta, Southeastern British Columbia, Dakota,
Wyoming, Montana. Idaho. Colorado. New Mexico. Arizona, and
the mountains and dry parts of California: and made numberless
inquiries covering the western part of the Mississippi drainage, as
well as all the adjoining mountains without hearing of any earth-
worms excepting in localities where they were introduced. (They
have recently appeared in many highly cultivated parts of Mani-
toba.)"

"Further. I am satisfied that, excluding the narrow humid belt
along the Pacific Coast, earthworms are not native to any part of

The Great Basin Naturalist

170 G. E. GATES Vol. XXVII, No. 3

America south of the Great Slave Lake, or west of the immediate
Mississippi Valley. Probably the true earthworm is not native to any
part of North America." (Presumably by "true earthworm" was
meant L. terrestris.)

As further support for Seton's conclusions, his "narrow humid
belt along the Pacific coast" is where Eisen found endemics unre-
lated to those of any other section of the United States. A northern
portion of that same belt is where Macnab and McKey-Fender were
finding endemics until their research was abandoned. Also accord-
ing to Repenning (1966), "The Pacific Coast Province has remained
a more or less isolated faunal unit since the early part of the mam-
malian history."

Presence of lumbricid exotics in other parts of the world, where
endemics could be expected but were lacking, usually was blamed
on native inability to survive competition. Those ideas were sum-
marized by Stephenson (1930, p. 905) as follows. "The Lumbricidae
are a recently evolved and dominant group which possesses great
powers of adaptation to new surroundings. Numerous species have
been carried by man and have established themselves all over the
world. Their introduction into a new territory frequently causes the
disappearance of the endemic earthworm fauna." The Lumbricidae
no longer can be regarded as recently evolved, the range from the
Mississippi River across Europe and Asia to Japan is the largest of
any megadrile family and must be of considerable age. Furthermore,
when pending revisions are completed American endemics will not
be in the same genera as now are the European species. Nor is the
family as a whole characterized by great powers of adaptation and
dominance. A few species, perhaps less than a dozen, in the struggle
for existence during Quaternary glaciation. may have acquired some
unusual colonizing ability but at the same time lost any ability they
may have previously had to maintain themselves in tropical climates
except at high elevations. Many years ago G. E. Hutchinson col-
lected 268 earthworms (Gates, in MS) in the vicinity of Naples.
Italy, where considerable endemicity could be expected, but 240
belonged to the very species that colonized the Great American
Desert. Seemingly, European endemics in their own home, lack
dominance and marked ability to survive whatever may have been
involved.

Anthropochorous forms very clearly have replaced endemics in
many niches of various regions in Australia. New Zealand. South
Africa and southern South America. By "replaced" nothing more
need be understood than "taken the place of." In some of such
niches, hemerophobic endemics may have disappeared as a result
of human disturbance of their environment before arrival of the
supposed competitors. Innumerable cases of transport and of first
appearance of worms in a new area around farm houses and fields
prove that exotics are hemerophilic and at least do tolerate consider-
able human interference with their environment. In New Zealand,
Lee (1961) thinks replacement rather than competitive extermina-

Nov. 30, 1967 DESERT EARTHWORM FAUNA 171

tion was involved. Miller et al. (1955) say that exotics usually ap-
pear some years after disappearance of the endemic fauna.

A Washington State endemic, according to Altman (1936), was
common in cultivated fields. As he had only ten specimens of the
species, the cited observation may have been given him by some one
unfamiliar with common similarities of many species. McKey-
Fendor {in lit. March 4, 1948) as a result of her experiences be-
lieved that "The natives quickly disappear under cultivation, being
restricted almost entirely to the edges of fields or to uncultivated
areas .... Old timers remark about their disappearance. In recent
years, Ray Albright of Dayton, Oregon has noted them most often
at the borders of his fields."

In South Africa, according to Pickford (1937), "The apparent
displacement of endemic by peregine species in cultivated grounds
is probably due to the fact that many of the latter can tolerate con-
ditions which are inimical to the former, for example the continued
breaking up and drying out of the soil." Pickford also mentioned
instances of South African endemics coexisting, in uncultivated sites,
with the exotic lumbricids.

In southwestern Australia, Michaelsen (1907) found that A.
caliginosa had acquired a wdde distribution and had become numer-
ous near towns within seventy years of the first settlement by
Europeans. Accordingly, the same lumbricids could have had time
enough (cf. previous pages), since first European settlements in
desert states, to acquire their present distribution. These instances
are cited in part because some have thought (cf. Omodeo 1963)
that far too little time since 1500 A.D. had been available for
European species to acquire their present American distributions.

Earthworms are soft bodied animals without exoskeletal protec-
tion against dehydration. Water constitutes around 80 percent of
body w^eight in P. hupeiensis, A. caliginosa (Grant, 1955), A. chloro-
tica and L. terrestris (Roots. 1956). Mechanisms for resisting dessi-
cation would seem to be of special survival value in the desert. A
weight loss of 54 percent by cocoons of Bimastos zeteki did not pre-
vent normal development (Murchie, 1960) when water was added.
Similar ability often is imputed to other megadriles, especially in
popular writings, but experimental proof is lacking. Earthworms
of various species can survive a loss of 50-70 percent of their body
water. Among those studied was A. caliginosa which Grant (1955)
found to have the most efficient mechanism for resisting dessication.
The American section of the caliginosa complex comprised three dis-
tinct species each of which could have differed from the others in
such a mechanism. Which species was used in the experiments un-
fortunately is unknown. A. trapezoides is the species most frequently
obtained in the desert states but is unlikely to have been consistently
available in New England.

How much moisture is needed for normal earthworm activity
and reproduction? Answers were first sought from precipitation data.

172

G. E. GATES

The Great Basin Naturalist

Vol. XXVII, No. 3

TABLE: Precipitation

State

Rainfall

in inches

Remarks

Arizona

3.1 to 32.4

Arkansas

48.64 (norma

1)

California (sou

ithem)

10-20 or 0-10

Colorado

17

7-60

Idaho

8-25

Iowa

31.44

Kansas

27

Missouri

40.15

Montana

10.15

Or 20 in northwest and
on mountains

Nebraska

12.65-27.58

But dry years in
irregular cycles

Nevada

3-12

New Mexico

14.3

North Dakota

14-22

'-

Oklahoma

16.51-46.53

South Dakota

14-24

Utah

5-40

Latter only in Wasatch Mts.

Wyoming

14.79

6 in parts, to 35 on mountains

Figures are from state articles. Encyclopedia Britannica, (1965). Some authors
gave only a single annual state average. Others gave annual averages for smaller
regions.

"The average annual rainfall is one of the most important factors
delimiting distribution of earthworms," according to Pickford (1937)
whose studies were made in South Africa. There she found that
endemic acanthodrilines were almost entirely restricted, in the
south, by the 20 inch isohyet. In the north, localities of her native
wonns were bounded by the 30 inch isohyet. Pickford was most in-
terested in the endemics and provided little information as to rela-
tionship between rainfall and distribution of the exotic lumbricids.
Mention was made of the fact that exotics had been found in areas
with less than 20 inches of annual rainfall. Seven lumbricid species
(Gates, in MS) were represented in her collections. Each of those
species is present in our desert states.

In Manchuria, Kobayashi (1940) found that an annual rainfall
of less than 400 mm {ca 16 inches) was unfavorable to earthworms.
He likewise was mainly interested in native taxa. "In the region
where the amount of annual rainfall is less than 400 mm. no
endemic species can exist." (p. 308). Some at least, if not all. of the
supposed endemics, when revised, will fall, into synonymies of more
or less widely spread anthropochores. Probably no single megadrile
will prove to be autochthonous (evolved in. and not found elsewhere)
in either Manchuria and Mongolia.

Beck collected earthworms in various areas where average annual
rainfall is much less than 16 inches and indeed, may amount only
to 3-10 inches annually. Surprisingly, the species represented in the
desert region by the third largest number of specimens is E. tetraedra.
This form rarely is found away from water and usually is charac-

Nov. 30, 1967 DESERT EARTHWORM FAUNA 173

terized as ainj)hibious. lininic. hygrophile or as a dweller in purely
aquatic habitats. Obviously something more than annual rainfall
must be involved.

Check of habitat data reveals that more than half of the desert
earthworms (all species) were obtained from banks of creeks, rivers,
springs, from seepage areas, marshes, and similar sites. Many other
specimens for which site data are lacking, possibly, if not also prob-
ably, were from similar habitats. The common factor of each of those
habitats is presence of more moisture than would be provided by the
annual rainfall. Important and suggestive data were provided for one
locality by Barrett. He found the worms in a strip of land ten feet
wide, on each side of the creek. In soil 10-50 feet from the creek
bank worms were absent.

A monthly rainfall of 1 1 -22 inches in certain sections of Burma
completely fails to keep many species from going into a state of
hibernestivation (Gates, 1961). Annual averages of rainfall may
then be of little or no significance in some desert areas. If so. amount
of moisture in soil will determine earthworm survival. Involved will
be maintenance of adequate soil moisture for a period long enough to
permit completion of a life cycle. Little information is available
about life cycles in different climates but in Europe information is
being acquired about moisture factors and earthworm survival.

In Hungary. Zicsi (1958) found that individuals of A. caliginosa
are active only when water content of the soil is at or above 30%
of capillary capacity. Four or more species have been involved in the
European section of the caliginosa complex. Similar figures for
validly defined species present in America were, rosea 35 percent,
chlorotica 40 percent, tyrtaeum (as lacteum) 30 percent.

Parthenogenesis usually is thought to favor colonization of new
areas (c/. Muldal. 1952) as only one specimen is needed. However,
a single specimen of an obligatorily amphimictic individual also can
start a new colony — after its battery of spermathecae has been
charged with sperm (millions?) as a result of copulating with an-
other individual. Of the 21 species living in the desert states. 12 re-
produce parthenogenetically. The other 9 are obligatorily amphimic-
tic. Of the 2500+ individuals, over 1800 were of parthenogenetic
species. Less than a third were of amphimictic species. Partheno-
genesis has not enabled lumbricids to colonize tropical lowlands.
Other factors, such as efficiency of drouth resisting mechanisms, may
be of equal or even greater importance especially in desert regions.

If earthworms were absent in plains and desert states when
settled by Europeans, the species now known to be domiciled there
must have been introduced and. for most of the region, within the
last 100-120 years. But how much reliance is to be placed on Seton's
observations? Considerable, according to Wilcox (1884) who con-
firmed and even added to them. Introduction could have been acci-
dental or deliberate. Very many if not most may have been acci-
dental (c/. p. above), in soil or other materials around roots of

The Great Basin Naturalist
174 G. E. GATES Vol. XXVII, No. 3

imported plants. Some introductions were deliberately made in order
to have a supply of bait for angling. Reports of deliberate introduc-
tion (cf. p. 144) provide additional support for previous absence of
angleworms. Probably only a very few of the deliberate introductions
got reported in print elsewhere.

Some reluctance to accept Seton's conclusions completely might
be in order. Travellers have reported absence of earthworms in cer-
tain parts of the tropics or adjacent areas. Observations were made in
the dry season when the soil was sun baked almost to brick hardness.
Worms were estivating at the proper depths in the soil from which
they return when the surface became satisfactorily moist. In some
Asiatic deserts earthworms remain inactive during drouths of several
years. It is of course possible that Seton and Wilcox did make some
of their observations in wrong seasons or that they did not investi-
gate inaccessible areas where conditions might have been more favor-
able. However, Beck's collection provided confirmation. Seton's word-
ing "west of the immediate Mississippi Valley" might also be accept-
able if we knew how far west is meant by "immediate." If an eastern
part of Texas, some portions of Oklahoma and Arkansas are "im-
bediate" that is where endemics are now expected.

Before the recent ice ages, lumbricid earthworms probably were
widely s])read throughout much of the northern hemisphere. Quater-
nary ice sheets thousands of feet thick exterminated all earthworms
for varying distances south of the present pole, in Europe and Asia
as well as in America. Evidence proving that extant endemics mi-
grated north in America since recession of the ice is lacking, though
such migration frequently must have been assumed. Earthworm
extermination was not confined to glaciated portions of Canada and
United States. For as yet unknown distances below the southern ice
face, the climate was too frigid for earthworms. The very few Ameri-
can lumbricids that did survive, except for B. parvus, appear to have
little colonizing ability. A few European species, including those
now domiciled in the desert states, not only survived, perhaps in
nearest proximity to the ice. but even evolved a physiology better
adapted to colonizing Iceland, (ireenland. Siberia, than tropical low-
lands.

Providing an explanation for absence of earthworms in un-
glaciated portions of the Great Basin states, now seems likely to be
more difficult. Until unquestioned endemics are found there, it is
necessary to suppose that these animals never lived there or were
exterminated, before arrival of Europeans, by unfavorable climatic
factors.

References

BuNTLEY, G. .J.. AND R. I. Papendick. 1960. Worn worked soils of eastern
South Dakota, their morphology and classification. Proc. Soil Sci. Soc.
America, 24:128-132.

CocKERELL, T. D. A. 1924. Earthworms and the cluster fly. Nature, 113:193.

Nov. 30, 1967 DESERT EARTHWORM FAUNA 175

Darwin, C. 1881. The formation of vegetable mould through the action of
worms, with observations on tlieir habits. London, vii -|- 328 pp.

Gates, G. E. 1942. Check list and bibliography of North American earthworms.

American Midland Nat. 27:88-108.
. 1956. On another lumbricid earthworm with "relictus" characteristics

and the disposition of Southern's species. Arm. Mag. Nat. Hist. (12)9:369-373.
. 1961. Ecology of some eartli worms with special reference to seasonal

activity. American Midland Nat. 66:61-86.

1966. Requiem — for megadrile Utopias. A contribution toward the

understanding of the earthworm fauna of North America. Proc. Biol. Soc.

Washington, 79:239-254.
Gr.ant, W. C. 1955. Studies on moisture relationships in earthworms. Ecology,

36:400-407.
H.\LLOCK. 1877. The sportsman's gazeteer and general guide. Forest & Stream

Publishing Co., New York.
Harnly, H. J. 1932. Double-tailed earthworm. Trans. Kansas Acad. Sci. 35:99.
HoLLON, W. E. 1966. The Great American Desert, p. 284. New York City.
KoBAYASHi, S. 1940. Terrestrial Oligochaeta from Manchokuo. Sci. Rept.

Tohoku Univ. (4, Biol.), 15:261-315.
Lee. K. E. 1959. The earthworm fauna of New Zealand. New Zealand Dept.

Sci. Indust. Res. Bull. 130, 486 pp.
MiCHAELsiiN, W. 1900a. Abhandl. Naurwiss. Ver. Hamburg, 16.

. 1990b. Oligochaeta. Das Tierreich, 10:575 pp. Berlin.

. 1907. Oligochaeta. In Die Fauna Siidwest- Australians. Jena.

. 1910. Zur Kenntnis der Lumbriciden und ihrer Verbreitung. Ann.

Mus. Zool. St. Petersbury, 15 pp.
MiCKEL, C. E. 1925. Notes on Zygocystis cometa Stein, a gregarine parasite of

earthworms. Jour. Parasitol. 11:135-139.
Miller, R. B. et al. 1955. The biological and chemical changes following shrub

burning on a New Zealand hill soil. New Zealand Jour. Sci. Tech. (B)37:

290-313.

MuLDAL, S. 1952. The chromosomes of the eartliforms. I. The evolution of

of polj'ploidy. Heredity, 6:55-76.
MuRCHiE, W. R. 1960. Biology of the oligochaete Bimastos zeteki Smith &

Gittins (Lumbricidae) in northern Michigan. American Midland Nat. 64:

194-215.
Olson, H. W. 1928. The earthworms of Ohio. Ohio Biological Survey Bull.

17:47-90.
Omodeo, p. 1952. Cariologia dei Lumbricidae. Caryologia, 4:173-275.
. 1963. Distribution of the terricolous oligochaetes on the two shores of

the Atlantic. In: Love and Love, North Atlantic Biota and their History, pp.

127-151.
Painter, R. 1942. Letters. Nat. Hist. 50:225, 276.
PiCKFORD, G. E. 1937. A monograph of the Acanthodriline earthworms of South

Africa. Cambridge, England, 612 pp.
Repenning, C. a. 1966. Tertiary provincial and immigrant terrestrial verte-
brates. Science, 151:863.
Roots. B. I. 1956. The water relations of earthworms. II. Resistance to dessica-

tion and immersion, and behavior when submerged and when allowed a

choice of environment. Jour. Exp. Biol. 33:29-44.
Seton, E. T. 1904. The master plowman of the west. Century Magazine, 68:

300-307.
. 1929. Lives of Game Animals. Vol. 4, part I. New York, Doubleday

Doran & Co., Inc. (Citations are from pages 412-414.)

The Great Basin Naturalist

176 G. E. GATES Vol. XXVII, No. 3

Smith, F. 1917. North American earthworms of the family Lumbricidae in

the collections of the United States National Museum. Proc. U. S. Nat.

Mus. 52:157-182.
Stephenson, J. 1930. The Oligochaeta. Clarendon Press, Oxford, xvi + 978 pp.
Taylor, S. C. 1924. Earthworms. Flower Grower, 11(10):405.
Wilcox, T. E. 1884. On the absence of earthworms from the prairies of the

Canadian northwest. Nature, 29:406-407.
Wright, H. E. and D. G. Frey. 1965. The Quaternary of the United States.

Princeton Univ. Press, x + 922 pp.
Zicsi, A. 1958. Einfluss der Trockenheit und Bodenbearbeitung auf das Leben

der Regenwiirmer in Ackerboden. Acta Agron. Acad. Sci. Hungaricae, 8:67-

75.

A NEW NAME FOR A UTAH LEPIDIUM

James L. Reveal i- -

For several years, Utah State University and the New York
Botanical Garden have been working on the Illustrated Flora of the
Intermountain Region which will have its first volume published
in a few years. As a first step to preparing for the actual writing
of the Flora, a checklist was written to summarize the known plants
which occur in this area by Arthur H. Holmgren, Curator of the
Intermountain Herbarium, and this author (1966). During the time
of the writing of the checklist, several taxa were noted which seemed
to be worthy of more study in the field and in the herbarium. One
such taxon is Lepidium montanum Nutt. ex Torr. & Gray ssp. demis-
sum C. L. Hitchc.

Questions regarding this subspecies were raised by the brief
comments given concerning the possible relationships of it to L.
montanum (sen. lat.) in general, and to L. davisii Rollins in particu-
lar. From the short description of ssp. demissum, and the more com-
plete description of L. davisii (Rollins, 1948) which Hitchcock
(1950) also reduced to the subspecific rank under L. montanum, it
seemed that these two taxa did not fit properly into the L. montanum
complex as previously defined by Hitchcock (1936). Thus, an iso-
lype and other specimens of L. davisii were studied by the author at
Idaho State University in 1963. The following year, attempts were
made to find both ssp. demissum and ssp. davisii in their type locali-
ties, but no plants were found. In the spring of 1965, after a deter-
mined search for the ssp. demissum in its type locality in the lower
end of Indian Creek Canyon, Duchesne Co., Utah, flowering speci-
mens were finally found and studied.

During the author's Predoctoral Internship at the Smithsonian
Institution, herbarium studies upon the entire Lepidium montanum
complex were conducted at the United States National Herbarium,
the New York Botanical Garden, and the Gray Herbarium of Har-
vard University. Additional studies were carried out on specimens
at the various Utah herbaria at Logan, Salt Lake City, and Provo.

In the spring of 1967, the type locality of Lepidium montanum
ssp. demissum was again visited and fruiting material obtained and
studied. From these combined field and herbarium studies, evidence

1. The nulhor would like to .icknowledgc the support given lo liiiu fur fiolil work from
Utah Stiile University and from National Science Foundiilion grants to IJr. Arlluir (ronquisl of
the New York Boldnit.il Garden for studies on the Intcrmountiiin l'"lora, a coopenilivc program
between Utah State University and the New York Botanical Garden. Hcrharium studies were
mainly supported by tlie Smithsonian Research Foundation and conducted duiing the author's Pre-
doctoral Internship proj^rani at the United Slates National Herbarium in Washington, I). C, from
September I'i(>(i to lebruary \'.)h7 . This pai>cr has been subniitlcil to the DcpartiiieiU of Botany,
Brigham Young University, as partial fulfillment of two credits of Special Problems given during the
Summer Session of 1907.

2. Department of Botany, Brigham Young University, Prove, Utah.

177

178

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 3

seemed to mount which indicated that L. davisii as well as the ssp.
demissum were distinct species in their own right. As the epithet,
demissum has been used at the species level for a South American
Lepidium, it is necessary to provide a new name for the Utah plant
which is now proposed as:

Lepidium barnebyanum Reveal, nom. nov.

L. niontanum Nutt. ex Torr. & Gray ssp. demissum C. L. Hitchc,
Madrono 10:157. 19-50. non L. demissum C. L. Hitchc, Lilloa 11:121.
1945.

Pulvinate perennials, (4) 5-15 cm. high and up to 20 cm.
across, the mats often forming raised humps; taproots deep, woody,
once or twice divided and widely spreading, the caudices dichoto-
mously branched 3-6 times. 2-5 cm. below the surface of the ground,
invested with only a few old leaf-bases, each resulting caudex branch
topped by a single, erect, green, leafy and herbaceous stem, the plants
appearing glabrous and glaucous, yet with minute retrorse simple
trichomes which are usually widely scattered nearly throughout the
plants; leaves basal and cauline, the basal leaves linear to linear-
oblanceolate, entire, ± v-shaped, 2.5-3.5 (4.5) cm. long, (1) 2-3.5
(4) mm. wide, the apices acute, the bases tapering gradually to
slightly expanded clasping petiole-bases, these hyaline and mem-
branaceous, 2.5-3.5 mm. wide; cauline leaves alternate along the
stems and about equally spaced. 5-10 mm. apart, the leaves 8-25

IS^^'Mi

^smm,

Fig. 1. General growth pattern of a fruiting plant of Lepidium barnebyanum
showing the pulvinate habit and tiie raised humps.

Nov. 30, 1967

A UTAH LEPIDIUM

179

Fig. 2. Enlarged detail of the inflorescences of Lepidium barnebyanum showing
the oblong-ovate fruits on the ascending pedicels and a few terminal flowers.

(30) mm. long, similar to the basal leaves but more reduced;
racemes 1-2 (3) cm. in early anthesis, ± clustered, becoming 2.5-6
(8) cm. long in fruit although a few late flov^ers may be present,
leafless, the simple trichomes more numerous than below; pedicels
terete, slender, ascending, 3-5 mm. long; sepals greenish, 2 mm. long
and 1-1.5 mm. wide, concave-convex, deciduous shortly after an-
thesis. flowers 3-4 mm. long and u[) to 7 mm. across, the petals white
with the blades rotund. 2-2.5 mm. long and wide, narrowing to
slender claws 0.8-1 mm. long. 0.3-0.5 mm. wide, the blades curving
backwards from their bases; stamens 6. the singles slightly longer
than the pairs, the filaments 2-2.5 mm. long, glabrous, the anthers
yellowish, oblong. 0.8-1 mm. long; silicles oblong-ovate. (3) 4-5 mm.
long. (2) 3-4 mm. wide, less than 2 mm. across, glabrous, not winged,
the apices tapering to long, briefly truncated tips; styles 0.5-1.2 mm.
long; cotyledons incumbent.

Typp:. — Utah: Duchesne Co.: Indian Creek Canyon on white
shale ridge-tops. ca. 4 mi. sw of Duchesne. 15 .lun 1947. D. D. Ripley
& R. C. Barnehy 8699. I lolotvpe dei)osited at W'lU! Isotypes: NY!
US!

Distribution. — Known only from a long ridge-top on the north
edge of Indian Creek Canyon, ca. 1/2 mi. n of Utah Highway 33. 3.5-4
mi. sw of Duchesne. Duchesne Co.. Utah. sec. 16-17. T. 4 S.. R. 5 W.,
elevation 6400-6500 feet. Flowering in Mav atid .Tune, fruiting into
Julv.

180

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 3

Specimens Examined. — Utah: Duchesne Co.: North ridge of
Indian Creek Canyon, 3.5 mi. sw of Duchesne, Holmgren, Reveal &
LaFrance, 1959 (BRY, NY, UTC), Reveal & Reveal 884 (BRY, CAS,
DS, GH, MO, NY, OKL, RM, RSA, UC, US, UT, UTC, WTU).

Hitchcock (1950) considered Lepidiwn barnebyanurn to be a
subspecies of the highly variable and complex species, L. montanum,
and most closely related to L. montanum var. inter gi folium (M. E.
Jones) C. L. Hitchc. However, I feel that the morphological differ-
ences between these two taxa are so great that the species rank
which is now proposed seems much more logical. From L. mon-
tanum var. intregifolium, L. barnebyanum differs in its sparsely
pubescent stems, leaves, and pedicels, the short linear to linear-
oblanceolate entire leaves, the highly branched and compressed
rooting S3^stem which results in the pulvinate mats, and in the
oblong-ovate silicles. In several respects, and especially in the pul-
vinate habit, L. barnebyanum is similar to L. davisii, a species which

Icm

I'

mm

mm

Fig. 3. Sketches showing the salient features of Lepidium barnebyanum.
A-B — front and side views of a basal leaf, C — a cross section through a leaf show-
ing a v-shape. D-E — front and side views of a mature silicle. F- -an outline of a
mature flo\%er petal showing the rotund blade and the narrow claw-like base.

Nov. 30, 1967 A UTAH lepidium 181

is apparently highly restricted to a dry bed of a small playa on a
sagebrush mesa near the lava outcrops on the north rim of the Snake
River Canyon, about 12-14 miles south of Mountain Home, Elmore
Co., Idaho. However, L. davisii may be distinguished by its shorter
leaves which are only 1-2 cm. long, denser pubescence, shorter sta-
ture (mostly under 8 cm. high), and its shorter silicles which are
3-3.5 mm. long. Unlike L. barnebyanum, the basal leaves of L.
davisii are shorter than the cauline leaves.

In the field, the Utah Lepidium occurs with Eriogonum bate-
rnanii M. E. Jones and Silene acoulis L. var. subacaulescens (F. N.
Will.) Fern. & St. John, on a broken white shale ridge in a Pinyon-
Juniper woodland. It might be suggested that the pulvinate condition
of L. barnebyanum is a result of the environment, but as most of the
other plants on the ridge are found on the sides of the ridge and
elsewhere in the Uinta Basin, the effects of the environment of these
plants in the several areas where they have been observed does not
seem to be an ecological response but rather due to a genetic basis
whicli aids the plants to survive in these ecological niches. Thus, it
is suggested that the pulvinate habit and the other morphological
features of L. barnebyanum are not a result of its ecological niche,
but rather a result of natural selection which has allowed for a
genetically distinct species to evolve, probably from the L. montanum
complex. The var. intregifolium does occur in the general area in
more moist habitats, and L. barnebyanum may have originated
from that entity. In searching for the Lepidium on other ridges
in Indian Creek Canyon, several other similar ecological sites were
found, but no other localities of the L. barnebyanum are presently
known to the writer.

The s{)ecies name is selected to commemorate the discoverer of
the plant. Rupert C. Barneby, the authority on the genus Astragalus,
and one of the finest collectors of the Intermountain flora.

Literature Cited

Hitchcock, C. L. 1936. The genus Lepidium in the United Stales. Madrono

-5:265-320.

. 1950. On the subspecies of Lepidium montanum. Madrono 10:155-158.

Holmgren, .\. H. & J. L. Reveal. 1966. Checklist of the vascular plants of

the Intermountain region. Intermountain Forest and Range Experiment

Station, l.niled States Forest Service. Research Paper INT-32.
Rollins. R. C. 1948. On two perennial caespitose Lepidium of western North

America. Madrono 9: 162-165.

The Great Basin Naturalist

Founded in 1939 by Vasco M. Tanner

A journal published from one to four times a year by Brigham
Young University, Provo, Utah.

Manuscripts: Only original, unpublished manuscripts, pertain-
ing to the Great Basin and the western United States in the main,
will be accepted. Manuscripts are subject to the approval of the
editor.

Illustrations: All illustrations should be made with a view to
having them appear within the limits of the printed page. The illus-
trations that form a part of an article should accompany the manu-
script. All half-tones or zinc etchings to appear in this journal are
to be made under the supervision of the editor, and the cost of the
cuts is to be borne by the contributor.

Reprints: No reprints are furnished free of charge. A price list
for reprints and an order form is sent with the proof.

Subscription: The annual subscription is $2.50 (outside the
United States $3.25). Single number, 80 cents.

All correspondence dealing with manuscripts should be addressed
to the Editor, Vasco M. Tanner, Great Basin NaturaHst, Brigham
Young University, Provo, Utah. Other matters such as subscriptions,
reprints, exchanges and other business should be addressed to Ernest
L. Olson, Chairman of University Publications.

Reprints Schedule of The Great Basin Naturalist

Each
Additional

2 pp.

4 pp.

6 pp.

8 pp. 10 pp. 12 pp.

2 pp.

50 copies

$6.00

$7.00

$8.00

$9.00 $10.00 $11.00

$2.00

100 copies

7.00

8.00

9.00

10.00 11.00 12.00

200 copies

8.00

9.00

10.00

11.00 12.00 13.00

300 copies

9.00

10.00

11.00

12.00 13.00 14.00

Covers: $10.00 for first 100 copies, $4.00 for additional 100 copies.

TABLE OF CONTENTS

New Records and Species of Neotropical Bark Beetles (Scoly-

tidae: Coleoptera), Part II. Stephen L. Wood 119

On the Earthworm Fauna of the Great American Desert

and Adjacent Areas. G. E. Gates . 142

A New Name for a Utah Lepidium. Illustrated. James L.
Reveal _ 1 77

Volume XXVn, No. 4
December 30, 1967

The

GYeat^Bami

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Z,oolocy and Entomolt^y
Brigham Young University, Provo, Utaii

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:
J. V. Beck, Bacteriology

WiLMER W. Tanner, Zoology, Chairman of the Board
W. Derby Laws, Agronomy
Stanley Welsh, Botany
C. Lynn Hayward, Zoology
Howard C. Stutz, Botany

Ex officio Members:

RUDGER H.

culture
Ernest L. Olson, Chainnan, University Publications

Rudger H. Walker, Dean, College of Biological and Agri-
cultural Sciences

S- ^-'76 6,^

1VIU3. COIVIP. ZOO!
LIBRARY

MAR 1 2 1968

L^ A en/ A pQ

The Great Basin Naturalist

Published at Provo, Utah by
Brigham Young University

Volume XXVII December 30, 1967 No. 4

NOTES ON ERIOGONUM - V
A REVISION OF THE ERIOGONUM CORYMBOSUM COMPLEX

James L. Reveal^

The E. corymbosum complex, as here defined, includes those
species of the genus which are woody perennial shrubs or subshrubs
with large lanceolate to elliptic or orbicular leaves which are on the
lower half of the stems. The E. microthecum complex (to be treated
in a later paper) differs mainly in having leaves that are narrower
and shorter on generally smaller plants.

Eriogonum corymbosum and its related species occur mainly in
the Colorado and Green River drainage basins with one outlying spe-
cies. The region is one of scant rainfall, warm temperatures, and high
evaporation. Yet, the scanty vegetation of this region possesses
numerous endemic species, and in the genus Eriogonum, the en-
demics are numerous and frequently encountered. The entities occur
in a variety of habitats, ranging from hard gumbo clay hills to sandy
desert flats and steep mountain slopes. However, rarely are the
species of Eriogonum the dominant plant in any given site.

Historical Review

The first species to be described in the complex was Eriogonum
corymbosum. Bentham (1856) named this species from information
supplied to him from John Torrey of New York, who had examined
the 1845 Fremont collection. In order to show the close relationship
between E. microthecum and E. corymbosum, Bentham proposed a
new section, Corymbosa. At the same time, Bentham described E.
microthecum var. fendlerianum from information sent to him by
Asa Gray of Harvard University. This taxon was later elevated to
the species rank by Small (1906). As Torrey & Gra}^ were aware
that Bentham was planning to publish E. corymbosum in de Can-
dolle's Prodromus. they j)roposed E. corymbosum var. divaricatum.
believing that their variety would appear before Bentham's revision
was published. Fortunately, however, the Beck with Report was de-
layed until 1857 and the new variety was published nearly a year

1. Department of Bot«ny, Brigham Young University, Provo, Utah.

183

The Great Basin Naturalist
184 JAMES L. REVEAL Vol. XXVII, No. 4

after Bentham's paper. If the variety had appeared before 1856, the
name would have been illegitimate.

Following these initial discoveries and papers, the few specimens
that were collected were simply called E. corymbosum. In 1895,
Marcus E. Jones distinguished the yellow-flowered plants as E.
aureum. However, in 1902, Aven Nelson raised a question as to the
use of the name E. aureum, noting that in Bentham's 1856 revision,
this name had been cited in synonymy under E. brevicaule Nutt. In
order to correct this seeming error, Nelson proposed the substitute
name, E. fruticosum, for E. aureum. However, as Bentham was cit-
ing the epithet, E. aureum, in synonymy, the name had never been
published and was still available to Jones. The Nelson substitute
name is superfluous.

In a second article. Nelson (1904) described E. salinum from a
small series of specimens from southern Wyoming. As it will be
noted below, this particular collection {Nelson 3753) is the closest
example found in this study to the Fremont collection which is the
type of E. corymbosum.

When Jones described E. aureum, he also proposed two new
varieties, var. ambiguum and var. glutinosum. The first is a form of
E. microthecum from the east side of the Sierra Nevada mountains of
California. By 1903, Jones recognized that his species, E. aureum.,
was nothing more than a variety of E. corymbosum and made the
combination E. corymbosum var. glutinosum for the yellow- flowered
form of the species from southern Utah and adjacent northern
Arizona.

The elevation of E. corymbosum var. divaricatum to the species
rank by Small (1906) as E. diver gens saw a corresponding shift of
the concept of this taxon away from its type to those specimens which
are here called E. corymbosum var. orbiculatum. At the same time,
Rydberg (1912) was modifying the concept of E. jonesii S. Wats, of
northern Arizona to include specimens of var. orbiculatum. Ryd-
berg's concept was subsequently incorporated in his Flora of the
Rocky Mountains and Great Plains (1917). In Tidestrom's (1925)
treatment of the genus, his key is written so that var. orbiculatum
will key out to E. nummulare M. E. Jones. This misapplication has
caused the herbarium folders of E. nummulare to bulge with a total-
ly unrelated species. It was not until 1936 when Susan G. Stokes, in
her monograph of the genus, noted these errors; yet. for the most
part, they have persisted.

It has been difficult to comprehend the species alignments of the
members of this complex as seen by Stokes. She placed most of the
species related to E. corymbosum under the more or less unrelated
High Plains species, E. effusum Nutt. At the same time she referred
E. aureum to E. microthecum. From herbarium studies, several col-
lections have been noted in which Stokes has given two or three dif-
ferent names to duplicates of the same collection. It seems likely
that she never truly understood the species involved in either the
E. corymbosum or E. microthecum complexes.

Dec. 30. 1967 eriogonum corymbosum complex 185

During the course of field work and herbarium studies conducted
independently and together with Jack D. Brotherson and others, a
number of new taxa related to Eriogonum corymbosum were found.
The history of these discoveries will be discussed under each entity.

Acknowledgments

I would like to express my appreciation to Dr. Stanley L. Welsh
of Brigham Young University who made several suggestions on the
various aspects of this paper and the species treated therein. I am
indebted to Jack D. Brotherson. formerly a fellow graduate student
at Brigham Young University, whose master's thesis on the ecology
of two varieties within Eriogonum corymbosum has been freely
drawn upon, and whose assistance in an early part of this study may
be noted below. Field work on this group has been supported both by
private financing and by National Science Foundation grants to Dr.
Arthur Cronquist of the New York Botanical Garden for the Inter-
mountain Flora Project, a cooperative program between the New
York Botanical Garden and Utah State University, and by the Texas
Research Foundation. Critical herbarium material has been bor-
rowed through the support of Brigham Young University. Her-
barium visits which have been made during this study have been
largely supported by Utah State University, the New York Botanical
Garden, and the Smithsonian Research Foundation. In this latter
case. I would like to thank Conrad V. Morton of the United States
National Herbarium who not only assisted me in the preparation of
the Ijatin descriptions, but also freely gave his advice on this and
other subjects during a predoctoral internship program at the United
States National Herbarium of the Smithsonian Institution from Sep-
tember 1966 to February 1967. This paper has been submitted to
the Brigham Young University Botany Department as partial ful-
fillment of three credits of Doctoral Research given during the Fall
Session of 1967-1968. I wish to thank Mrs. Twila Davis Bird whose
excellent illustrations are seen below.

The following herbarium collections have been consulted, and to
the several curators of herbaria that were visited or who sent loan
material for this study. I am most grateful. The abbreviations follow
Lanjouw & Stafleu (1964).

A Arnold Arboretum, Harvard University, Cambridge, Massachusetts.

ARIZ University of Arizona, Tucson, Arizona.

ASC Northern Arizona University, Flagstaff, Arizona.

BM British Museum of Natural History, London, England.

BR Jardin Botanique de I'fitat, Bruxelles, Belgium.

BRY Brigham Young University, Provo, Utah.

CAS California Academy of Sciences, San Francisco. California.

COLO University of Colorado, Boulder, Colorado.

DS Dudley Herbarium, Stanford University, Stanford, California.

GH Gray Herbarium, Harvard University, Cambridge, Massachusetts.

IDS Idaho State University, Pocatello, Idaho.

ISC Iowa State University, Ames, Iowa.

KSC Kansas State University, Manhattan, Kansas.

LY Laboratoire de Botanique de la Faculte des Sciences, Lyon, France.

186

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

MICH

MO

MONTU

NY

OKL

ORE

OSC

P

PH
POM

RM

RSA
SD
TEX
UC

UNM
US

USPS

UT

UTC

WIS

ws

WTU

University of Michigan, Ann Arbor, Michigan.
Missouri Botanical Garden, St. Louis, Missouri.
Montana State University, Missoula, Montana.
New York Botanical Garden, Bronx Park, New York.
Bebb Herbarium, University of Oklahoma, Norman, Oklahoma.
University of Oregon, Eugene, Oregon.
Oregon State University, Corvallis, Oregon.

Museum National d'Histoire Naturelle, Laboratoire de Phanerogamie,
Paris, France.

Academy of Natural Sciences, Philadelphia, Pennsylvania.
Pomona College Herbarium, Rancho Santa Ana Botanic Garden,
Claremont, California.

Rocky Mountain Herbarium, University of Wyoming, Laramie,
Wyoming.

Rancho Santa Ana Botanic Garden, Claremont, California.
San Diego Society of Natural History, San Diego, California.
University of Texas, Austin, Texas.
University of California, Berkeley, California.
University of New Mexico, Albuquerque, New Mexico.
United States National Herbarium, Smithsonian Institution, Washing-
ton, D. C.

Forest Service Herbarium, United States Department of Agriculture,
Washington, D. C.

University of Utah, Salt Lake City, Utah.

Intermountain Herbarium, Utah State University, Logan, Utah.
University of Wisconsin, Madison, Wisconsin.
Washington State University, Pullman, Washington.
University of Washington, Seattle, Washington.

Taxonomy

A. Leaf-apices sharply acute, the leaves mostly lanceo-
late, usually more than 3 cm long.

B. Branches subglabrous to tomentose; involucres
tomentose externally.

C Involucres 2.5-3 mm long; inflorescences
with several short branches; perianth
white, 3-3.5 mm long, the calyx-segments
± dissimilar; leaves 3-5 cm long, the
petioles 3-6 mm long, deciduous on the
lower parts of the stems; Mancos Shale
east of Wellington, Carbon Co., Utah.

1. E. lancifolium

CC. Involucres 3-4 mm long; inflorescences

open with few long branches; basal stem

leaves usually persistent.

D. Involucres 3.5-4 mm long; perianth
white, 3.5-4.5 mm long, the calyx-
segments ± dissimilar; leaves 3.5-7
cm long, the petioles 5-10 (18) mm
long; Bad Land Cliffs, Duchesne
Co., Utah. 2. E. hylophilum

DD. Involucres 3-3.5 mm long; perianth
yellow, 2.5-3 mm long, the calyx-

Dec. 30. 1967 eriogonum corymbosum complex 187

segments similar or nearly so; leaves
2.5-4 cm long, the petioles 4-7 mm
long; Indian Creek Canyon, Du-
chesne Co.. Utah. 3. E. duchesnense

BB. Branches glabrous or floccose; involucres gla-
brous externally.
C. Leaves tomentose below.

D. Involucres 2-3 mm long; perianth
cream to pale yellowish-white, 2-3
mm long, the calyx-segments ob-
lanceolate; inflorescences 1-2 dm
long; Dinosaur National Monument
area. Uintah Co., Utah. 4. E. saurinum

DD. Involucres 2.5-3.5 (4) mm long;
perianth- white, 2.5-3.5 (4) mm
long, the calyx-segments elliptic to
oblong; inflorescences up to 1 dm
long; southern Colorado and ad-
jacent New Mexico and Texas.

5. E. fendlerianum
CC. Leaves as well as the entire plant glabrous
throughout; perianth yellow, 3-4 mm
long; San Rafael Desert. Emery Co.,
Utah. 6. E. smithii

AA. Leaf-apices mostly rounded, the leaves oblanceolate
to lanceolate or elliptic to nearly orbicular, 1-3 (4.5)
cm long; southwestern Wyoming and western Colo-
rado to Utah, south to northern Arizona and north-
western New Mexico. 7. E. corymbosum

1. Eriogonum. lancifolium Reveal & Brotherson, spec. nov.

Fruticulus 3.5-5 dm altus, erectus; caules basi per 1.5-3 dmi foliosi
vel nudi per 5-10 cm; laminae foliorum lanceolatae. 3-5 cm longae,
0.5-1 cm latae, d= revolutae. subtus albo-tomentosae, supra subglabrae
et virides. petiolis brevibus, 3-6 mm longis, basi expansa petioli 2 mm
lata, tomentosa; caules deorsum tomentosa. sursum subglabri vel
parce tomentosi. bracteis 1.5-3 mm longis, linearibus; inflorescentiae
ex fasciculis confertis ramulorum erectorum compositae. 6-14 cm
longae, subglabrae; involucra turbinata, 2.5-3 mm longa. 1.5-2 mm
lata, sessilia, extra tomentosa. intus glabra, 5-lobata. bracteolis ob-
lanceolatis. 1.5-2 mm longis, pedicellis 3-4 mm longis. glabris; peri-
anthia alba, costa brunnea. (2.5) 3-3.5 mm longa, segmentis subsi-
milibus, exterioribus spathulatis, 3-3.5 mm longis. 1-1.3 mm latis,
apice truncato. interioribus oblanceolatis. 2.5-3 mm longis, 0.3-0.6
mm latis; stamina 1.5-4.5 mm longa. filamentis basi glabris. antheris
0.5-0.4 mm longis, oblongis; achaenia brunnea. 2 mm longa.

Shrubby perennials. 3.5-5 dm high, forming densely branched,
erect crowns from woody caudices; leaves on the lower half of the

The Great Basin Naturalist
188 JAMES L. REVEAL Vol. XXVII, No. 4

plants, the lowest most often deciduous so that the first leaves are
5-10 cm up the stems, leaf-blades lanceolate, 3-5 cm long, 0.5-1 cm
wide, entire or ± crenulate, revolute in some, densely white-tomen-
tose below, less so to subglabrous and green above, the petioles short,
3-6 mm long, floccose-tomentose, the expanding petiole-bases 2 mm
wide, subglabrous to sparsely tomentose externally, tomentose in-
ternally; lower stems densely tomentose. the branches becoming sub-
glabrous or sparsely tomentose above, 5-12 cm long; bracts ternate.
scalelike, 1.5-3 mm long, linear, widening from the acute apices to
connate bases, glabrous or subglabrous externally, tomentose in-
ternally; inflorescences dense clusters of short, erect, trichotomous,
subglabrous branches, 6-14 cm long; involucres sessile, turbinate,
2.5-3 mm long, 1.5-2 mm wide, tomentose externally, glabrous in-
ternally, the 5 acute teeth I/4 the length of the tubes, the bractlets
narrowly oblanceolate, 1.5-2 mm long with minute capitate marginal
cells, the pedicels 3-4 mm long, glabrous; perianth white with brown-
ish (greenish at first) midribs and bases, the outer whorl of segments
spathulate with truncate apices, 3-3.5 mm long, 1-1.3 mm wide, the
inner segments narrowly oblanceolate, 2.5-3 mm long, 0.3-0.6 mm
wide; stamens exserted, 1.5-4.5 mm long, the filaments glabrous at
the bases, the anthers 0.3-0.4 mm long, oblong, yellowish-white to
reddish; achenes brown, 2 mm long, the ovoid bases gradually taper-
ing to 3-angled beaks. Figure 1.

Type, utah: Carbon Co. On low, rolling Mancos Shale hills 5
mi E of Wellington, 9 Sep 1967, James L. Reveal & Gerrit Davidse
957 . Holotype deposited at utc. Isotypes distributed to ariz, bry,
CAS, DS, GH, MO, NY, RM, RSA, uc, US, and Other herbaria.

Distribution. Known only from the Mancos Shale hills 3-6
miles east of Wellington and 5.5 miles south of Wellington, Carbon
Co., Utah. Map 1. Flowering mainly from August to September.

Specimens Examined, utah: Carbon Co., Near Price, Flowers in 1926 (ny,
ut), in 1927 (bry, ut, utc), in 1928 (bry, sd, ut); Price, M. E. Jones 6512
(bm, mo, pom, us); 5 mi E of Wellington, Moore 661 (bry, ny, utc); Price,
5. Stokes s.n. (uc); 5.5 mi S of Wellington, Reveal <& Davidse 955 (bry, ny.
utc); 5 mi E of Wellington, Reveal <£; Reveal 727 (bry, ny, utc); 3 mi E of
Wellington, VanCott & Larsen 87 (utc); 5 mi E of Wellington, Welsh & Moore
1836 (bry, isc).

This population was first called to my attention by Dr. Stanley
L. Welsh of Brigham Young University in 1965. Later, as this study
began, a few specimens were seen from the Price- Wellington area
that agreed with the Welsh & Moore collection that had been seen
before. With the assistance of Brotherson, detailed herbarium studies
were carried out in the spring of 1966 which were then followed by
field work in 1966 and 1967.

From the herbarium studies, we quickly realized that other taxon-
omists before us had considered this population to be of some interest.
Dr. Seville Flowers of the University of Utah had collected a large
series of specimens over a period of three years for study by Stokes.
Although both Jones and Stokes had collected the species before, their
material was rather limited.

Dec. 30, 1967 eriogonum corymbosum complex

189

Fig. 1 . Habit sketch of Eriogonum lancifolium showing the general aspect of
the species and a single enlarged segment of the inflorescence with involucres and
exserted flowers.

The Great Basin Naturalist
190 JAMES L. REVEAL Vol. XXVII, No. 4

One of the more interesting features of this plant and some of
the other members in the complex is the fact that the lower leaves on
the stems are deciduous. This condition of early leaf fall gives the
plants a rather bare look. In the fall of the year, wilted leaves may
be seen as well as the old expanded petiole-bases. However, in the
spring, the branches that had leaves the year before were found to
lack leaf buds. Also, it should be pointed out that the leaves on this
species tend to vary with the conditions of the year. In poor years
the leaves tend to be longer and narrower and decidedly more acute,
and the inflorescences are not as densely branched. Thus the type
collection cited above which was collected in a good year has a large,
dense inflorescence with somewhat shorter leaves than most of the
specimens cited in the list of exsiccatae.

On most of the early collections, the location data were given
simply as "Price" or "Near Price." Attempts to find this species
near the Price area have been without success, and it is thus suggest-
ed that all of the collections came from the Wellington area, and the
use of "Price" was simply to indicate the largest city in this general
area.

The relationship of E. lancifolium to the other members of the
complex is not certain. Its most closely related species seems to be
E. saurinum, but the pubescence is not at all similar. At the same
time, the Wellington Buckwheat approaches E. hylophilum in the
branching pattern of the basal portions of the plants, but the inflo-
rescences differ, and E. hylophilum has leaves which are shaped
differently and not deciduous.

2. Eriogonum hylophilum Reveal & Brotherson, spec. nov.

Fruticulus 2.5-4 dm altus; caules basi per 1-1.5 dm foliosi; lami-
nae foliorum lineari-lanceolatae vel lanceolatae, 3.5-7 cm longae, 3-6
(8) mm latae, subtus albo-tomentosae, supra viridi-tomentosae, peti-
olis longis, 5-10 (18) mm longis, basi expansa petioli 2-3 (3.5) mm
lata, tomentosa; caules ubique tomentosi, 1-2.5 dm longi, bracteis
superioribus 2-3 mm longis, inferioribus ± folia ceis, 3-20 mm longis;
inflorescentiae ex fasciculis nonnulis brevibus densis ramulorum com-
positae, tomentosae, 3-8 cm longae; involucra turbinata, 3.5-4 mm
longa, 2.5-3 mm lata, extra tomentosa, intus glabra, 5-6 lobata.
bracteolis oblanceolatis, 3.5-5 mm longis, pedicellis 3.5-5 mm longis,
glabris; perianthia alba, costa viridi, (3) 3.5-4 (4.5) mm longa, extra
glabra, intus minute glandulosa, segmentis subsimilibus. exterioribus
spathulatis, 1.3-1.7 (2) mm latis. interioribus oblanceolatis, 0.6-0.9
(1.2) mm latis; stamina 2-3 (3.5) mm longa, filamentis basi pilosis,
antheris 0.3-0.5 mm longis, oblongis; achaenia brunnea, 2.5-3 mm
longa.

Subshrubby perennials, 2.5-4 dm high, forming open, branched
and erect crowns from woody caudices; leaves on the lower fourth of
the plants, persistent, linear-lanceolate to lanceolate, 3.5-7 cm long,
3-6 (8) mm wide, densely white-tomentose below, less so and green
above, the margins entire and rt revolute or rarely crenulate, the

Dec. 30. 1967 eriogonum corymbosum complex 191

petioles long, 5-10 (18) mm long, tomentose, the expanding petiole-
bases 2-3 (3.5) mm wide, tomentose on both surfaces; stems tomen-
tose, 1-2.5 dm long; bracts ternate, scalelike and triangular above,
2-3 mm long, becoming ±: foliaceous and up to 2 cm long below,
widening from acute apices to connate bases; inflorescences open,
the branches short and stout. 3-8 cm long, tomentose; involucres ses-
sile with 1-3 per node, turbinate, 3.5-4 mm long, 2.5-3 nun wide,
tomentose externally, glabrous internally, the 5-6-lobes less than ^4
the length of the tubes, the bractlets oblanceolate, 3.5-5 mm long,
hirsutulous with long marginal cells, the pedicels 3.5-5 mm long,
glabrous; perianth white with greenish midribs, (3) 3.5-4 (4.5) mm
long, glabrous without, minutely glandular along the midribs within,
the calyx-segments =b dissimilar, the outer whorl of segments spathu-
late with acute apices, 1.3-1.7 (2) mm wide, the inner segments
oblanceolate, 0.6-0.9 (1.2) mm wade; stamens included, 2-3 (3.5) mm
long, the filaments pilose basally, the anthers greenish-white, 0.3-0.5
mm long, oblong; achenes brown, 2.5-3 mm long, the ovoid bases
tapering to 3-angled. retrorsely roughened, beaks. Figure 2.

Type, utah: Duchesne Co. Along Utah highway 53 (the
Wellington-Myton Road) in Gate Canyon, 2.7 mi SW of the summit
of the Bad Land Cliffs, Sec. 20, T. 11 S., R. 15 E., elevation 6500 feet,
15 Aug 1966, Noel H. Holmgren <& James L. Reveal 5017 . Holotype
deposited at utc. Isotypes distributed to ariz, bry, cas, ds, gh, mo,
NY, RM, RSA, uc, US, and other herbaria.

Distribution. Locally common on the Bad Land Cliffs near
Gate Canyon, Duchesne Co., Utah. Map 1 . Flowering from late July
to September.

Specimens Examined, utah: Duchesne Co., Suminit of the Bad Land Cliffs,
along Utah highway 53, Holmgren, Reveal, <& LaFrance 2386 (ariz, bry, cas,

DS, GH, KANS, MO, NY, OKL, CSC, RM, RSA, UC, US, UT, UTC, WIs) .

This distinctive species has apparently gone unnoticed by the
several taxonomists who have passed over the summit of the road
between Wellington and Myton. Actually, as far as Eriogonum is
concerned, several of the late season entities are poorly known, and
suspecting that a large and relatively unknown flora might exist in
the Intermountain Region in the fall, several collecting trips have
been made during this period. As this paper and others (Reveal, in
press a. b) demonstrate, the rewards from this late season collecting
have been rich.

Eriogonum hylophilum was first discovered in 1965, but at that
time it was thought to be E. corymbosum. As the nature of E. corym-
bosum was determined, it was possible to see that this material from
the Bad Land Cliffs was distinct. The population extends nearly five
miles down Ciate Canyon on the Wellington side of the summit, but
less than half of a mile on the Myton side. Toward the west, the
species was found to extend less than a mile, but its eastward limits
have not been determined.

In Ciate Canyon, E. hylophilum occurs as scattered plants on
Artemisia hillsides in the Pinyon-Juniper belt. The species is com-

192

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

Map 1. Distribution in eastern Utah of Eriogonum lancifolium (L); E. hy-
lophilum (H); E. duchesnense (D); E. saurinum (S); and E. smithii (SM).

Dec. 30. 1967 eriogonum corymbosum complex

193

monly found with E. corymbosum var. erectum, and in the field, the
two are easily distinguished. However, the separation of E. hylo-
philum and E. lancifolium is not as readily apparent, especially in
the herbarium. These two differ in several technical characteristics
such as perianth size and shape, involucre characters, and the nature
of the inflorescences and growth patterns, yet they seem to be rather

Fig. 2. Habit sketch of Eriogonum hylophilum showing the generally stout
erect stems and the capitate or short-rayed inflorescences of clustered involucres,
with an enlarged involucre and flowers, and of a single achene.

The Great Basin Naturalist
194 JAMES L. REVEAL Vol. XXVII, No. 4

closely related. Actually however, the species that is probably most
closely related to E. hylophilum is E. duchesnense.

3. Eriogonum duchesnense Reveal, spec. nov.

Fruticulus 2-3.5 dm altus; caules basi per 1-1.5 dm foliosi; lami-
nae foliorum lanceolatae vel spathulatae, 2.5-4 cm longae, 0.5-1 cm
latae, subtus albo-tomemosae, supra subglabrae et virides, petiolis
brevibus, 4-7 mm longis, basi expansa petioli 2 mm lata, extra tomen-
tosa, intus subglabra vel glabra; caules deorsum tomentosi, ramis
1-1.5 dm longis; bracteae foliaceae, 3-10 mm longae, lineares vel
lanceolatae, tomentosae; inflorescentiae ex fasciculis nonnullis brevi-
bus densis ramulorum compositae, tomentosae, (3) 5-10 cm longae;
involucra turbinata, 3-3.5 mm longa, 2-2.5 mm lata, sessilia, extra
tomentosa, intus glabra, 5-lobata, bracteolis oblanceolatis, 2-2.5 mm
longis, pedicellis 3-4.5 mm longis, glabris; perianthia flava, 2.5-3
mm longa, extra glabra, intus minute pilosa vel glandulosa. segmentis
subsimilibus, elongato-obcordatis vel oblanceolatis, exterioribus 1.5
mm latis, interioribus 1-1.3 mm latis; stamina 1.5-2 mm longa, fila-
mentis basi pilosis, antheris 0.3-0.4 mm longis, oblongis; achaenia
brunnea, 2-3 mm longa.

Subshrubby perennials, 2-3.5 dm high, forming open crowns
from woody caudices; leaves on the lower third of the plant, per-
sistent, lanceolate to spathulate, 2.5-4 cm long, 0.5-1 cm wide, densely
white-tomentose below, subglabrous and green above, the petioles
short, 4-7 mm long, subglabrous, the petiole-bases 2 mm wide, tomen-
tose to subglabrous externally, subglabrous to glabrous internally;
branches tomentose, densely so among the leaves, less so above, the
branches 1-1.5 dm long; bracts ternate, ± foliaceous, 3-10 mm long,
linear to lanceolate, widening from the acute apices to connate bases,
tomentose within and without; inflorescences open and compoundly
divided, the short, stout branches rather clustered, tomentose, (3)
5-10 cm long; involucres sessile with 1-3 per node, turbinate, 3-3.5
mm long, 2-2.5 mm wide, tomentose externally, glabrous internally,
the 5 acute teeth nearly ^^ the length of the tubes, the bractlets ob-
lanceolate, 2-2.5 mm long, hirsutulous with numerous long marginal
cells, the pedicels 3-4.5 mm long, glabrous; perianth yellow, 2.5-3
mm long, glabrous without, wdth few scattered hyaline hairs or
minute glands within, the calyx-segments ± similar, the outer whorl
of segments elongate-obcordate, 1.5 mm wide, the inner segments
slightly narrower, 1-1.3 mm wide; stamens included, 1.5-2 mm long,
the filaments pilose basally, the anthers 0.3-0.4 mm long, greenish-
yellow, oblong; achenes brown, 2-3 mm long, the ovoid bases tapering
to short 3-angled beaks, ridged with the distinct margins extending
down the entire length of the fruit, roughened with retrorse hairs
along the margins. Figure 3.

Type, utah: Duchesne Co. Ca. 24 mi SW of Duchesne along
Utah highway 33, in Indian Creek Canyon, 0.7 mi below the Indian
Creek Canyon Guard Station on steep clay banks, sec. 28, T. 10 S., R.
7 W., elevation 7800 feet, 2 Sep 1964, James L. Reveal 678. Holotype

Dec. 30. 1967 eriogonum corymbosum complex

195

1mm

1mm|.^^

Fig. 3. Habit sketch of Eriogonum duchesnense showing the erect stems and
comiK>und inflorescences with a single enlarged involucre with several flowers and
a single involucre.

deposited at utc. Isotypes distributed to ariz, bry, cas, ds, gh, mo,
NY, rm, rsa, uc, us, and other herbaria.

Distribution. Infrequent and rare in Indian Creek Canyon,
Duchesne Co., Utah. Map 1. Flowering mainly in August and
September.

Specimens E.xamined. utah: Duchesne Co., 5 mi N of the summit of Indian
Creek Canyon, Maguire & Richards 13287 (gh, uc, utc).

Eriogonum duchesnense has been a puzzling plant to study.
When this species was first discovered by me a large collection was
made, photographs taken, and its distribution in the canyon was
studied. However, since 1964, the State of Utah has rebuilt the upper
part of the highway in the canyon, and for the past three years not a

The Great Basin Naturalist
196 JAMES L. REVEAL Vol. XXVII, No. 4

single plant has been found. Thus, the description of the entity is
based on the material collected by Maguire and Richards, and on the
type collection.

This species seems to be most closely related to E. hylophilum,
from which it may have been derived. As noted in the key, the two
differ in several characteristics of the involucres, calyx-segments, and
leaves. Yet, the two are rather similar in habit, growth patterns, and
ecology, although when comparing the various collections in the
herbarium, these similarities are lost. The most readily discernible
feature separating the two species is the flower color.

4. Eriogonum saurinum Reveal, spec. nov.

Fruticulus 3-5 dm altus; caules basi per 1-1.5 dm foliosi vel nudi
per 5-10 cm; laminae foliorum lanceolatae, 3-6 cm longae, 4-8 mm
latae, subtus albo-tomentosae, supra subglabrae et virides, petiolis
5-10 mm longis, basi expansa petioli 2-2.5 mm lata, tomentosa; caules
ubique glabri, (1.5) 2-3.5 dm longi, bracteis superioribus 1 mm
longis, linearis, inferioribus foliaceis, 1-3 cm longis; inf foreseen tiae
ex fasciculis apertis ramulorum expansorum compositae, 1-2 dm
longae; involucra turbinata, 2-3 mm longa, 1.5-2 m!m lata, subses-
silia vel sessilia, glabra, 5-lobata, bracteolis linearis, 2-3 mm longis,
pedicellis 2.5-3.5 mm longis, glabris; perianthia eburnea, costa
brunnescena, 2-3 mm longa, segmentis subsimilibus, exterioribus ob-
lanceolatis, 3-3.5 mm longis, 0.7-0.9 mm latis, apice rotundo, interi-
oribus anguste oblanceolatis, 2.5-3 mm longis, 0.5-0.7 mm latis;
stamina 2.5-4 mm longa, filamentis basi pilosis, antheris 0.3-0.4 mm
longis, flavis; achaenia brunnea, 2.5-3 mm longa.

Shrubby perennials, 3-5 dm high with open and spreading crowns
from branching woody caudices; leaves on the lower half of the
plants, these often becoming deciduous on the lower 5-10 cm of the
stems, the leaf-blades 3-6 cm long, 4-8 mm wide, lanceolate, densely
white-tomentose below, subglabrous and green above, the petioles
5-10 mm long, floccose-tomentose, the petioles bases 2-2.5 mm wide,
tomentose on both surfaces; lower stems tomentose only among the
leaves, becoming green and glabrous above, (1.5) 2-3.5 dm long;
bracts ternate, scalelike and 1 mm long above, becoming foliaceous
and up to 3 cm long below, these similar to the leaves and usually
only at the first node, the acute apices widening to connate bases,
glabrous externally, tomentose internally; inflorescences cymose,
spreading, the open branches mostly trichotomous, 1-2 dm long; in-
volucres sessile or subsessile, not clustered, turbinate, 2-3 mm long,
1.5-2 mm wide, glabrous, the 5 acute lobes less than ^4 the length of
the tubes, the bractlets linear, 2-3 mm long, the pedicels 2.5-3.5 mm
long, glabrous; perianth cream- white to pale yellowish- white with
light tan midribs and bases, 2-3 mm long, glabrous except for minute
glandular hairs within along the midribs, the calyx-segments ± simi-
lar, the outer whorl of segments oblanceolate with rounded apices,
3-3.5 mm long, 0.7-0.9 mm wide, the inner segments narrower and
shorter, 2.5-3 mm long, 0.5-0.7 mm wide; stamens 2.6-4 mm long.

Dec. 30, 1967 eriogonum corymbosum complex

197

the filaments pilose basally, the anthers 0.3-0.4 mm long, pale
yellow; achenes light brown, 2.5-3 mm long, the subglobose bases
tapering to 3-angIed beaks. Figure 4.

Type, utah: Uintah Co. Along the Island Park Road, 10 mi E
of Vernal along Brush Creek on steep hillsides on the ridges of
Mowry Shale, sec. 1, T. 1 N., R. 22 E., elevation 5200 feet, 15 Aug
1966, Noel H. Holmgren & James L. Reveal 3019. Holotype de-
posited at UTC. Isotypes distributed to ariz, bry, cas, ds, gh, mo,
NY, RM, RSA, uc. US, and other herbaria.

Fig. 4. Habit sketch of Eriogonum saurinum showing the general aspect of
the species with enlarged drawings of the inflorescence, a single involucre with
two exserted flowers, and the bracts subtending the inflorescence.

The Great Basin Naturalist
198 JAMES L. REVEAL Vol. XXVII, No. 4

Distribution. Locally common in the Dinosaur National Monu-
ment area where it is highly restricted to the narrow band of Mowry
Shale, Uintah Co., Utah. Map 1. Flowering from late July to
September.

Specimens Examined, utah: Uintah Co., Diamond Valley, Andrews <£:
Noble s.n. (utc); 6.5 mi N of Vernal, near Steinaker Res., Reveal 674 (ariz, bry,

CAS, DS, GH, KANS, MO. NY, OKL, OSC, RM, RSA, TEX, UC, US, UT, UTC, WIs) ; near

Dinosaur N. M., Welsh 566 (bry); Island Park, Welsh 517 (bry, utc); between
Island Park and Vernal, Wolf & Dever 5261 (utc).

Eriogonum saurinum is an unusual species from an ecological
standpoint. Throughout the area surrounding the Dinosaur National
Monument as well as in the Monument itself, there is a narrow,
often upright, band of acidic Mowry Shale which has a pH often
less than 5 (Hanson, 1962; Brotherson, 1967). On the soil derived
from the Mowry Shale, as well as the band itself, E. saurinum is
found. When standing on some prominence, it is possible to see this
resistant shale standing out as it often forms the backbones of the low
hills. On these hills, and especially along their ridges, E. saurinum
is abundant. In fact, when one views the ridges from the side, they
appear frilled with the green branches of the Eriogonum. When the
Mowry Shale is not exposed, the buckwheat is missing. The Dino-
saur Buckwheat is rarely more than 10 feet away from the shale.
Those plants which grow along side the layer are not as robust as
those closer or directly upon the shale. Welsh (1957) made several
observations on the occurrence of this plant, as did Brotherson (per.
comm.), and they both confirm the strict adherence of this species
to Mowry Shale.

In the Dinosaur National Monument, endemics are not unusual
(Welsh & Christensen, 1957; Holmgren, 1962). One of the more
interesting of the endemics is Astragalus chlooides Barneby. In many
respects the genus Astragalus is similar to Eriogonum in its features
of endemism. Like E. saurinum, A. chlooides is highly restricted
to a single formation (Entrada Sandstone) and has not been found off
of this specific soil type.

For several years. E. saurinum has been incorrectly called E.
lonchophyllum Torr. & Gray. However, this latter species is known
only from northern New Mexico and southern Colorado, and is
actually a member of the section associated with E. brevicaule Nutt.
The two species differ in several characteristics. The leaves of both
Species are lanceolate, but those of E. saurinum are less than 6 cm
long, while the leaves of E. lonchophyllum often range from 5-10
cm long. The involucres and perianth of E. lonchophyllum are both
3-4 mm long, while those of E. saurinum are only 2-3 mm long.

The distinction between E. saurinum and E. corymbosum is most
striking in the field. As Brotherson (1967) has shown, they occupy
two different soil types, with E. saurinum being on the acid soils and
E. corymbosum normally on the basic soils, with notable exceptions
(Welsh 1957). In the herbarium, the two may be distinguished by
the glabrous stems and open, few branched crowns of E. saurinum.

Dec. 30, 1967 eriogonum corymbosum complex 199

while the other species is tomentose and the crowns are decidedly
more densely branched.

5. Eriogonum fendlerianum (Benth. in DC.) Small, Bull. Torrey
Bot. Club 33: 55. 1906.

E. microthecum Nutt. var. ? fendlerianum Benth. in DC, Prodr.
14: 18. 1856. E. effusum Nutt. ssp. fendlerianum S. Stokes, Gen.
Eriog. 79. 1936. E. corymbosum Benth. in DC. var. fendlerianum
attributed to S. Wats, by S. Stokes, Gen. Eriog. 79. 1936, in sy-
nonymy.

E. aimlei Woot. & Standi., Contr. U. S. Nat. Herb. 16: 117. 1913.
E. effusum Nutt. ssp. ainslei S. Stokes, Gen. Eriog. 79. 1936. (Type:
Cimarron. Colfax Co., New Mexico, 20 Sep 1909, C. A^. Ainslie s n.
Holotype: us! Isotypes: us!)

Subshrubby to shrubby perennials, (1) 1.5-4 (5) dm high, form-
ing open and spreading crowns of few to several branches from
woody caudices; leaves on the lower half to two-thirds of the plants,
the leaf-blades lanceolate to elliptic, 1.5-4 (5) cm long, (0.3) 0.5-2 cm
wide, entire or crenulate along the margins, densely white-tomentose
below, subglabrous to glabrous and green above, the petioles long,
5-20 mm long, subglabrous on the upper surface, tomentose on the
lower surface, revolute, the expanding petiole-bases 2-3 mm wide,
glabrous or tomentose externally, tomentose internally; lower stems
densely tomentose among the leaves, the branches becoming glabrous
or rarely floccose above, 3-15 (20) cm long; bracts ternate, mostly
scalelike. 1-2.5 mm long, linear, widening from acute apices to con-
nate bases, glabrous or subglabrous without, densely tomentose with-
in; inflorescences open, mostly trichotomous, 2-10 cm long; in-
volucres sessile, peduncled in the forks of the branches 1-7 mm long,
involucres turbinate to turbinate-campanulate, 2.5-3.5 (4) mm long,
1.5-3.5 mm wide, glabrous or rarely subglabrous externally, often
pilose at the throat of the tubes internally, becoming glabrous below,
the 5 acute lobes less than 14 the length of the tubes, the bractlets
linear-oblanceolate, 2-3 mm long, hirsutulous with minute marginal
cells, the pedicels 3-4.5 (5) mm long, glabrous; perianth white with
brownish midribs and bases, 2.5-3.5 (4) mm long, the calyx-segments
similar or nearly so. the outer whorl of segments elliptical to oblong,
the inner segments slightly narrower and shorter; stamens 2-4 mm
long, the filaments pilose basally, the anthers 0.4-0.5 mm long, ob-
long; achenes brow^n, 2-2.5 mm long, the ovoid bases gradually taper-
ing to 3-angled beaks. Figure 5.

Type, new mexico: Toas Co. Red River at Rock Creek, 1847,
Fendler 767. Holotype deposited at gh! Isotypes: bm, gh, mo, ny!

Distribution. Southwestern Colorado eastward to eastern Colo-
rado and adjacent northeastern New Mexico, and (apparently) in
western I'exas. Map 2. Flowering from July to early September.

Representative Specimens. Colorado: El Paso Co., Ck>lorado Spr., M. E.
Jones s.n. (us); Fountain, Redfield 527 (ny). Fremont Co.. Canon City. Brande-

200

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

Fig. 5. Habit sketch of Eriogonum fendlerianum showing the stout stems and
inflorescence with enlarged drawings of a single involucre with exserted flowers
and the bracts subtending the inflorescence.

Dec. 30. 1967 eriogonitm corymbosum complex

201

Map 2. Distribution of Eriogonum jendlerianum.

gee 527 (ny, uc); Canon City, M. E. Jones 788 (bm, br, cas, ny, pom, us, utc);
3 mi S of Florence, Penland 4621 (colo) ; 6 mi E of Canon Cit3% Reveal &
Davidse 871 (bry, ny, utc); 12 mi E of Canon City, Waterfall 10868 (ariz,
COLO, OKL, utc). Las Animas Co., 10 mi SE of Trinidad, Rollins 1860 (ds, gh,
MO, MONTU, NY, US, UTC, ws). Montezuma Co., Mancos, Eastwood in 1890
(coLo), in 1892 (mo, uc); North Rim of Mesa Verde, Frauke s.n. (rm); Mesa
Verde, Maguire 12701 (ny, okl, utc). Montrose Co., 10 mi SW of Crawford,
Rollins 1961 (ds, gh); Coventry', Walker 540 (gh, mo, ny, pom, us, ws). Otero
Co., La Junta, Rose <£• Fitch 17165 (ny, us). Pueblo Co., Pueblo, Greene in 1874
(gh, ISC, NY, PH, us), in 1889 (uc); 18 mi W of Pueblo, Waterfall 10856 (ariz,
COLO, OKL, UTc). San Miguel Co., 10 mi W of Telluride, Maguire 12775 (gh,
utc). new MEXICO: Colfa,ic Co., Cimmaron, Ainslie s.n. (us); Raton Mts., Bele
50 (gh); near Raton. Rose & Fitch 17554 (ny, us); 6 mi SW of Raton, Reveal &
Davidse 875 (bry, ny, utc); 12 mi SE of Raton, Turner & Melchert 4827 (te.x);
8 mi SE of Raton, Waterfall 12025 (okl, tex). Quay Co., Logan, Fisher 92
(mo, us). Rio Arriba Co.. near Tierra Amarilla, Wooton s.n. (us), texas:
Without definite location. Ramrnel s.n. (us).

Eriogonum jendlerianum occurs in two forms, both of which have
been named. The low, spreading form was named E. ainslei and is
found mainly in southeastern Colorado and on the plains of north-
eastern New Mexico. To the west is the larger and somewhat more
erect form which is represented by the type of E. jendlerianum. In
southwestern Colorado, this kind of plant is at its extreme in stature.
Nevertheless, in studying the plants in the field, these variants are
easily bridged when intermediate geographical areas are visited.
South of Raton. New Mexico, both forms may be seen growing to-
gether. At least in that area, the small form {E. ainslei), seems to
be only younger plants of the larger form {E. jendlerianum). As I

The Great Basin Naturalist
202 JAMES L. REVEAL Vol. XXVII, No. 4

can see no morphological grounds for maintaining both as species,
E. ainslei is now reduced to synonymy.

In the Stokes treatment (1936), she recognized E. effusum Nutt.
ssp. salicinum (Greene) S. Stokes as a form related to what is here
called E. fendleriarium. However, in visiting the type area of E.
salicinum at the Black Canyon of the Gunnison, it was found to be
the same kind of plant that has been called E. scoparium Small.
Eriogonum salicinum, which includes E. scoparium and possibly E.
tristictum Small, seems to be closely related to E. lonchophyllum
Torr. & Gray. However, this group of species is in need of consider-
able critical study.

Eriogonum fendlerianum is seemingly closely related to the more
northern species described above, yet the exact nature of this sug-
gested relationship, if any, has not been determined. From the aspect
of gross morphology, this species resembles E. saurinum, mainly in
the open crown, various features of the involucres and perianth, and
usually in its ecology. At the present time, however, it seems that
E. fendlerianum is actually closer to E. corymbosum, and the simi-
larities with the Dinosaur Buckwheat are presently thought to be
parallel rather than an indication of close relationship. The plants
from the Mesa Verde area are exceedingly large and robust, and in
the leaf features approaches certain specimens which are now tenu-
ously assigned to E. corymbosum var. orbiculatum from southeastern
Utah. The Reveal 687 from 15 miles south of the La Sal Junction
resembles the Mesa Verde plants closely, but differs in the stem
pubescence, and size of the involucres and perianth. The area from
the La Sal Junction southeastward to the Mesa Verde has had little
botanizing in the fall of the year, and until some detailed field obser-
vations of E. fendlerianum can be made, the several questions of
relationships must remain unanswered.

6. Eriogonum smithii Reveal, spec. nov.

Fruticulus (3) 4-8 dm altus, omnino viridis et glabris; caules basi
per 1.5-4 (5) dm foliosi; laminae foliorum ellipticae, 2.5-4.5 cm
longae, 6-10 mm latae, ± revolutae, petiolis 3-5 mm longis, basi
expansa petioli 2.5-3.5 mm lata; bracteae 1-1.5 mm longae; inflores-
centiae cymosae, 2-2.5 cm longae, ± compactae; involucra turbinata
(2.5) 3-3.5 mm longa, 2-2.5 mm lata, glabra margine ciliata excepta,
5-lobata, bracteolis lineari-oblanceolatis, 1.5-2.5 mm longis, pedicellis
3-3.5 mm longis; perianthia lutea, 3-4 mm longae, glabra, segmentis
subsimilibus, exterioribus obovatis, 1.5-2 mm latis, interioribus ob-
lanceolatis, 1-1.5 mm latis; stamina 2-5 (7) mm longa, filamentis basi
pilosis, antheris flavis, 0.4-0.5 mm longis; achaenia brunnea. 3 mm
longa.

Perennial shrubs, (3) 4-8 dm high and up to 2 m across, with
many branches arising from woody caudices, the stems woody only
at the base, becoming herbaceous above, the plants entirely glabrous
throughout except for the axils of the leaves which bear the leaf-bud
primordia, these sparsely pubescent; leaves decurrent and widely

Dec. 30, 1967 eriogonum corymbosum complex 203

separated along the lower % of the plants, the lower leaves often
deciduous so that the first leaves are 5-15 cm or more up the stems,
the leaf-blades narrowly elliptical, 2.5-4.5 cm long, 6-10 mm wide,
± revolute. the leaf-margins always thicker than the blades, bright
green, the petioles short, 3-5 mm long with the expanding petiole-
bases 2.5-3.5 mm wide, glabrous and brown or tan externally, densely
or sparsely white-tomentose internally; bracts temate, scalelike, 1-1.5
mm long, linear to triangular, the acute apices widening to connate
bases; inflorescences open, cymose, mostly trichotomously branched
throughout. 2-25 cm long, the branches short and somewhat compact,
bright green; involucres sessile, turbinate, (2.5) 3-3.5 mm long, 2-2.5
mm wide, glabrous, finely ciliated in the throat in some, sharply
angled with 5 acute teeth, these less than ^4 the length of the tubes,
the lobes of the teeth often with membranaceous margins, the bract-
lets 2 per pedicel, linear-oblanceolate, 1.5-2.5 mm long, hirsutulous
with short acute marginal cells, the pedicels 3-3.5 mm long, glabrous;
perianth yellow, 3-4 mm long, glabrous without, microscopically
glandular along the midribs within, the calyx-segments subsimilar,
the outer whorl of segments obovate, 1.5-2 mm wide, the inner seg-
ments oblanceolate, 1-1.5 mm wide; stamens 2-5 (7) mm long, the
filaments pilose basally, the anthers yellow, 0.4-0.5 mm long, oblong;
achenes brown, lance-ovoid, 3 mm long, the ovoid bases tapering to
roughened 3-angled beak«. Figure 6.

Type, utah: Emery Co. On the east side of a low summit along
a sandy desert road between Little Flat Top and Big Flat Top, San
Rafael Desert, about 10 mi SE of Utah highway 24 from the turnoff
w^hich is 0.5 mi S of the Goblin Valley turnoff, sec. 22, T. 26 S., R.
13 E., elevation 5500 feet, 14 Aug 1966, Noel H. Holmgren & James
L. Reveal 3012. Holotype deposited at utc. Isotypes distributed to
ARIZ, BRY, CAS, Ds, GH, MO, NY, RM, RSA, uc, US, and Other herbaria.

Distribution. Known only from the north and east side of Big
Flat Top and near Little Flat Top, San Rafael Desert, Emery Co.,
Utah. Map 1 . Flowering from late July to September.

Specimens Examined, utah: Emery Co., E of Big Flat Top, Reveal &
Davidse 950 (bry, ny, utc); N of Big Flat Top, Reveal & Davidse 951 (bry,
NY, utc); between Big Flat Top and Little Flat Top, A. D. Smith s.n. (bry, cas,
NY, utc).

Eriogonum smithii can be instantly recognized by its bright green
color, its bright yellow flowers, and its almost total lack of pube-
scence, the latter feature not found in any other perennial species in
the genus. Its relationship to other species in the genus has been
determined on the basis of its overall morphology. It is believed that
E. smithii is related to E. corymbosum.

The site where the type of E. smithii was collected is in a small
circular basin that is surrounded on two-thirds of its sides by steep
red sandstone cliffs of the Summerville Formation. The remaining
side is the edge of the San Rafael Desert which is composed of red
Entrada "blowsand" in this area. In the basin, the plants occur
mainly on the floor where there seems to be a high selenium content.

204

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

Fig. 6. Habit sketch of Eriogonum smithii showing the erect leaves and
coxnix>und inflorescences with enlarged drawings of the petiole base, a cross sec-
tion of a leaf, and involucres with exserted flowers.

Dec. 30, 1967 eriogonum corymbosum complex 205

However, the largest plants occupy the upper edges of the basin and
especially so on the sandy desert floor east of Little Flat Top and thus
north and east of Big Flat Top. In this area the plants are on the
edges of small outwash areas where the infrequent rain water has
formed small gullies and rivlets. Further to the east, on a small mesa
east of Big Flat Top, the plants of E. smithii are much smaller and
more sparsely branched than elsewhere. I have seen this species up
to 5 miles east of Little Flat Top, but how much further to the east
the species extends has not been determined.

The branching habit of E. smithii reflects most clearly the kind
of branching habit found in this group. In the axis of one of the
leaves is a stem bud. This bud is often one decimeter or more up the
branch. In the s])ring, the bud produces a stem which in turn, of
course, has a stem bud. However, as the stem buds are always below
the inflorescence, the new stem with its greater weight causes the
old stem to become nearly prone and thus widely spreading. The
new stems are erect or only slightly spreading outwardly to conform
with the overall shape of the bush. The old inflorescence, that is, the
one produced the previous year, is now brown and dried and sticks
out beyond the new stem. In E. corybosum var. orbiculatum, the
old inflorescences may be easily seen. Thus, as the plants grow, the
distance between the root and the flowering stems gradually in-
creases. In the sandy areas particularly, the older stems and
branches are buried in rounded mounds of sand.

It is a privilege to follow the suggestion of Arthur H. Holmgren,
who called this plant to my attention, by naming this species in honor
of Dr. Arthur D. Smith, professor in the Range Sciences Department
and a member of the Utah State Fish and Game Department. Long
recognized as an authority on range management practices in the
western United States, he has been a diligent plant collector of Great
Basin and Intermountain plants for over thirty years.

7. Eriogonum corymbosum Benth. in DC. Prodr. 14: 17. 1856.

Subshrubby and shrubby perennials, (2) 3-8 (12) dm high, form-
ing erect to hemispherical crowns of few to many branches from
branching, woody, caudices; leaves mainly on the lower half of the
plants, often extending up to the base of the inflorescences, the leaf-
blades oblanceolate to lanceolate or elliptic to nearly orbicular, 1-3
(4.5) cm long, (0.3) 0.5-3 (3.5) cm wide, entire or crenulate, densely
white-tomentose on both surfaces, or often becoming less pube-
scent to subglabrous or nearly glabrous and green above, the
petioles short to long. 2-15 mm long, subglabrous to tomentose. the
petiole-bases 1.5-3 mm wide, subglabrous to tomentose externally,
densely tomentose internally; stems tomentose or subglabrous to
(rarely) glabrous, (8) 10-20 (25) cm long; bracts ternate. scalelike,
1-3 mm long, triangular, widening from acute apices to connate bases,
tomentose or glabrous externally, tomentose internally, the bracts of
the lower nodes of the inflorescences occasionally foliaceous and up
to 2.5 cm long, similar to the leaves only more reduced; inflo-

The Great Basin Naturalist
206 JAMES L. REVEAL Vol. XXVII, No. 4

rescences cymose with few to many tomentose or glabrous dichoto-
mous or trichotomous branches, (1) 2.5-20 cm long; involucres
sessile, turbinate, 1.5-3.5 mm long, 1-2 mm wide, tomentose to
glabrous externally, glabrous internally, the 5 acute lobes from 1/6
to 1/3 the length of the tubes, the bractlets narrowly oblanceolate,
1-3.5 mm long, hirsutulous with long acute marginal cells, often with
scattered capitate cells among the longer acute cells, the pedicels
glabrous, 1.5-4 mm long; perianth white with greenish or reddish
midribs, whitish-brown, pale yellow, or yellow, 2-3.5 (4) mm long,
the calyx-segments ± similar, the outer whorl of segments oblanceo-
late to spathulate with acute or rounded apices, the inner segments
elliptical, slightly shorter and narrower than the outer segments,
glabrous except for a few scattered hyaline hairs or minute glands on
the midribs within; stamens mostly included, 1-4 (5) mm long, the
filaments glabrous or more commonly pilose basally, the anthers
0.3-0.5 mm long, oblong; achenes brown, 2-2.5 (3) mm long, the
ovoid to subglabrous bases tapering to retrorsely roughened or rough-
ened 3-angled beaks.

Key to the Varieties of Eriogonum corymbosum
A. Perianth white or brownish- white.

B. Leaves oblanceolate to elliptic, 1-3 (4.5) cm
long, 1-2 cm wide, the petioles 2-6 mm long.

C. Involucres 1.5-2.5 mm long, 1-1.5 mm
wide; stems spreading into subglobose
crowns, the branches whitish-tomentose;
perianth white; plants found mostly below
6000 feet, northwestern Colorado and ad-
jacent southern Wyoming southward
through northeastern and central Utah to
north-central Arizona. 7a. var. corymbosum

CC. Involucres 2.5-3.5 mm long, 1.5-2 mm
wide.

D. Stems and crowns open and erect,
the branches brownish- tomentose;
leaves 2-3.5 cm long, 0.5-1.5 cm
wide, brownish-tomentose; plants
found mostly above 6000 feet, north-
eastern Utah from western Wa-
satch Co. to extreme western Uintah
Co. 7b. var. erectum.

DD. Stems and crowns spreading, the
branches silvery-tomentose; leaves
3-4 cm long, (0.5) 1-2 cm wide,
silvery-tomentose; known only from
Wellington, Carbon Co., Utah.

7c. var. davidsei

Dec. 30. 1967 eriogonum corymbosum complex 207

BB. Leaves elliptical-oblong to ovate-orbicular, 1-3
(4) cm long, 1-3 (3.5) cm wide, the petioles
5-10 (15) mm long.

C. Plants greenish; leaves densely tomentose
below, subglabrous to glabrous and green
above; perianth 2.5-3 mm long; plants
found mainly on sandy soil, southeastern
Utah and adjacent Colorado south into
northeastern Arizona and extreme north-
western New Mexico. 7e. var. orbiculatum

CC. Plants brownish-white; leaves densely
white-tomentose below, floccose and
brownish- or yellowish- white above; peri-
anth 2-2.5 mm long; plants found mainly
on clay soils, west-central New Mexico.

7f. var. velutinum
AA. Perianth yellowish.

B. Perianth pale yellow to yellow, 2.5-3 mm long;
involucres 2.5-3 mm long; inflorescences less
than 2 cm long; extreme eastern Utah Co. and
adjacent Duchesne Co., Utah 7d. var. albogilvum

BB. Perianth yellow, 1.5-2.5 mm long; involucres
1-2 mm long; inflorescences 3-10 cm long;
southern Utah and northern Arizona.

7g. var. glutinosum

7 Si. Eriogonum corymbosum var. corymbosum

E. corymbosum var. divaricatum Torr. & Gray, Pacif. Railroad
Report 2: 29. 1857. E. divergens Small, Bull. Torrey Bot. Club 33:
55. 1906, non E. divaricatum Hook. E. effusum Nutt. ssp. divarica-
tum S. Stokes. Gen. Eriog. 80. 1936. (Type: Near Green River,
Emery Co., Utah, 1 Oct 1853, Creuzfeldt on the Gunnison Expedition.
Holotype: ny! Isotype: gh!)

E. salinum A. Nels., Bull. Torrey Bot. Club 31: 240. 1904. E.
effusum Nutt. ssp. salinum S. Stokes, Gen. Eriog. 81. 1936. (Type:
Salt Well, Sweetwater Co., Wyoming, 17 Jul 1897, A. Nelson 3753.
Holotype: rm! Isotypes: gh, us!

E. effusum Nutt. ssp. corymbosum S. Stokes, Gen. Eriog. 79.
1936.

E. effusum Nutt. ssp. durum S. Stokes. Gen. Eriog. 80. 1936.
(Type: Sunnyside, Carbon Co., Utah, 13 Sep 1901, M. E. Jones
ni34. Holotype: pom! Isotype: ny!)

Shrubs or subshrubs, 3-8 dm high, the crowns suberect to sub-
globose, up to 1 m across; leaves lanceolate to oblanceolate or elliptic,
1-3 (4.5) cm long, (0.3) 0.5-1 (1.5) cm wide, the petioles short. 2-6
(8) mm long; bracts scalelike above, often becoming foliaceous be-
low; inflorescences mostly cymose, trichtomous nearly throughout.
3-10 cm long, usually densely tomentose but becoming subglabrous

208

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

Fig. 7. Habit sketch of Eriogonum corymbosum var. corymbosum showing
the general aspect of the species with enlarged drawings of the stem leaves and a
single involucre with several exserted flowers.

in some; involucres 1.5-2.5 mm long, 1-1.5 mm wide; perianth white,
2-3 (3.5) mm long. Figure 7.

Type. Colorado: Eagle Co. "On road — Day we reached Grand
River." From the Fremont Report, the type area would be near
Yarmony. NE of Piney Cr., near the Colorado River, 4 Sep 1845,

Dec. 30. 1967 eriogonum corymbosum complex

209

John C. Fremont 248. Holotype deposited at ny! Drawing, frag-
ments, and photographs of this specimen are deposited at bry. Iso-
type: gh!

Distribution. Northwestern Colorado and adjacent southwest-
ern Wyoming south through northeastern and central Utah to north-
central and northwestern Arizona. Map 3. Flowering mainly from
late July to early October.

Map. 1 Distnhution in Utah. Colorado, Arizona, and New Mexico of
Eriogonum corymbosum, var. corymbosum (C) with short dash lines showing the
form of the variety represented by the type, the long dash lines showing the
desert form, and the dotted line which represents the low mountain and southern
form; var. erectum (E); var. davidsei (D); var. albogilvum (A); var. orbiculatum
(O); var. velutinum (V); and var. glutinosum (G).

The Great Basin Naturalist
210 JAMES L. REVEAL Vol. XXVII, No. 4

Representative Specimens. Arizona: Coconino Co., Navajo Bridge, Darrow
2906 (ariz); Jumpup Spr., Darrow 2972 (ariz, cas); 13 mi SE of Fredonia,
Eastwood & Howell 6385 (cas, us); 14 mi W of Cameron, Kearney & Peebles
12S26 (gh, us), 12827 (ariz); 2 mi SE of Fredonia, Reveal 698 (ariz, bry, cas,
Ds, GH, KANs, MO, NY, OKL, RM, RSA, TEX, uc, US, UT, UTC, wis); Moquitch Canyon,
Swapp 44 (usFs); Wapatki N. M., Whiting 1089/5279 (ariz); Ryan, Thackery
569 (ariz, us). Mohave Co., Between Fredonia and Mt. Trumbull, Mann 2
(usFs); Boysag Point, Michaels 23 (■^riz). Colorado: Garfield Co., 20 mi SW
of Rifle, Waterfall 11040 (okl). Mesa Co., DeBeque, Osterhout 4753 (ny, rm),
5848 (okl, rm); 0.2 mi E of the Utah-Colorado line along U. S. Hwy. 50-6,
Reveal & Davidse 948 (bry, ny, utc). Moffat Co., Lodore Canyon, Baker &
Cutler 3489 (okl); Dinosaur N. M., Barmore s.n. (utc); Blue Mt., McLeod
51 A (colo). Montrose Co., Bedrock, Payson & Payson 3924 (gh, rm). utah:
Without definite location. Southern Utah, Siler 141 (gh, isc, mo, ph). Daggett
Co., Hideout Camp, Flowers et al. 216 (ut). Duchesne Co., 2 mi S of Duchesne,
Reveal 676 (bry, ny, utc). Emery Co., 0.5 mi N of San Rafael Bridge, Harrison
9807 (bry, uc, utc); San Rafael Swell, M. E. Jones s.n. (pom); 6 mi SE of Castle
Dale, McVaugh 14593 (cas, mich, ny) ; Cottonwood Canyon, Ware 179 (usfs).
Garfield Co., Bryce Canyon N. P., Eastwood & Howell 7170 (cas, gh, uc, utc) ;
15 mi N of Widtsoe, Holmgren et al. 2253 (bry, cas, ds, gh, isc, mo, ny, osc,
RSA, TEX, uc, utc). Grand Co., Tumbow Cabin, Arches N. M., Welsh & Moore
2709 (bry). Kane Co., E of Zion N. P., Eastwood & Howell 6374 (cas); Mt. Car-
mel, Eastwood & Howell 7127 (cas); Mt. Carmel Junct., Harrison 11067 (bry, uc,
us, utc); Glendale, M. E. Jones 6047b (mo, ny, uc); 2 mi above the Lake
Canyon, Lindsay 103 (wis); N of Kanab, Milner 8959 (us); V4 rni S of Glendale,
Reveal & Holmgren 316 (utc, wtu). Sevier Co., Tony's Hollow, Martineau 119
(usFs); 1 mi up Red Cr., Salina Canyon, Stevens 90 (bry, utc). Uintah Co.,
Dinosaur N. M. Headquarters, Bradley 5371 (colo); 33 mi S of Ouray, Brother-
son 453 (bry); 12 mi S of Ouray, Brotherson 519, 520 (bry); Split Mt. Gorge,
Brotherson 776 (bry); Quarry, Dinosaur N. M., Brotherson 830 (bry); 23 mi
SE of Ouray, Holmgren et al. 2267 (bry, cas, ds, mo, ny, osc, rm, tex, uc, us,
UT, utc); 32 mi SE of Ouray, Holmgren et al. 2269 (bry, cas, ds, gh, mo, ny,
osc, rm, tex, uc, us, ut, utc); Tabyago Canyon, Holmgren et al. 2350 (bry,
CAS, ds, gh, mo, ny, rm, RSA, TEX, UC, US, UT, UTC, wis) ; Upper Hill Cr., Tavaputs
Plateau, Vickery & Wiens 1662 (ds, rsa, uc, ut, wtu) ; Overlook, Split Mt.
Gorge, Welsh 379 (bry). Wayne Co., Fruita, Beck s.n. (bry); 7 mi E of Teasdale,
Harrison et al. 7536 (bry); Grover, Holmgren, et al. 2547 (bry, ny, utc); 5 mi
E of Teasdale, Holmgren et al. 2550 (bry, ny, utc); Wayne Wonderland, Milner
7242 (ut); Rabbit Valley, Ward 561 (us).

The critical part of the discussion of E. corymbosum var. corym-
bosum is in regards to the type collection (fig. 8). First, the Fremont
collection was made in 1845, and the label data given indicates that
he obtained the specimens from near Yarmony, Eagle Co., Colorado.
No other collection of this species is known at present to come from
this area, and a recent visit to this area failed to reveal the plant.
Thus, if the type came from this area, it is the easternmost collection.
The interested reader of far western history of the United States
will recall that following this collection. Fremont made an extended
exploration trip through Utah, Nevada, and California, became in-
volved in the Bear Flag Revolt and for his part in this affair was or-
dered out of the state into Oregon only to return later and become in-
volved in the fight for California's independence from Mexico in the
Mexican War. Needless to say, the type specimen of E. corymbosum
went with him throughout his adventure, and suffered considerably.
The specimens became partially destroyed, broken in places, and
heavily infected with matted mycelium of a fungus. With such
poorly preserved material, it was necessary for me to carry out de-

Dec. 30. 19(i7 eriogonum corymbosum complex

211

"1

. Pro*, llii 17,

Fig. 8. Photograph of the holotype of Eriogonum corymbosum.

tailed morphological studies in order to correctly associate the sub-
sequent specimens.

The holotype is deposited at the New York Botanical Garden and
has few leaves, although the isotype at the Crray Herbarium has a
few more. Thus, except for the leaf studies, all of the details have
been taken from the holotype. The inflorescence of the holotype was
manufactured when the specimen was mounted. The small pieces
which bear the involucres were glued down to give a corymbose
appearance rather than a cymose inflorescence as seen in the isotype.
and in most of the subsequent collections. The pubescence of the
stems, leaves, and involucres is obscured by the fungal mycelium.
However, in a few small places, it is possible to see the kind of
pubescence, and it too compares favorably with modern sj^ecimens.

The Great Basin Naturalist
212 JAMES L. REVEAL Vol. XXVII, No. 4

When Torrey worked on the collection, he made a series of drawings
on a small sheet of blue paper. Although the pencil drawings have
faded over the years, they are still visible, and compare favorably
with the present observations. Fortunately, the concept of E. corym-
bosum as held by most taxonomists is represented by the type, and
through all of the difficulties experienced with the type collection,
it has been possible to ascertain the identity of the holotype and con-
clude that the type represents that part of the overall species as out-
lined in this paper.

This conclusion was aided by several additional key specimens.
The specimen at hand that comes closest to the holotype is another
type collection. Aven Nelson collected the only specimens of E.
corymbosum known from Wyoming- and described it as E. salinum.
In making detailed comparisons with the Fremont collection, the
Nelson specimens were found to be nearly identical. The several
collections from the Dinosaur National Monument area also re-
semble the type closely, but these tend to have bracts which are
somewhat longer and with the stem leaves closer together on the
stem.

The plants from northwestern Colorado and adjacent north-
eastern Utah are generally of a shorter stature than those plants
found elsewhere. The larger plants to the south which are generally
in the foothills and the mountains were recognized by Stokes under
the name E. effusum Nutt^ ssp. durum, while the plants of inter-
mediate stature which generally occur in the desert were described
earlier as E. corymbosum var. divaricatum by Torrey and Gray.

In E. corymbosum var. corymbosum several different popula-
tions may be seen which are associated with corresponding variation
in ecology and geology. As noted above, those plants which are
similar to the holotype range from northwestern Colorado and ad-
jacent Wyoming, westward into northeastern Utah, then south onto
the Tavaputs Plateau (Map 3, p. 309). This southern line ends
where this high plateau breaks off into the Roan and Book cliffs.

At the base of the Book Cliffs is the larger and more robust plant
which Stokes named ssp. durum. The durum phase of var. corym-
bosum extends westward from the Book Cliffs over to the eastern
front of the Wasatch Plateau, then southward along the mountains
to its southern end where the species crosses the mountains and
continues southward into southwestern Utah and adjacent Arizona
(Map 3).

The desert form of corymbosum extends down the Green River
drainage, and southward from the Book Cliffs onto the San Rafael
Desert and the San Rafael Swells, with outlying populations in
Arches National Monument and adjacent areas, and in extreme
western Colorado. This plant was described as var. divaricatum.
The divaricatum form of the variety is found on the clay soils, and

2. Recently (3 Aug 1967) Dr. C. L. Porter of the University of Wyoming recollected this
variety in Wyoming in the same location as Nelson, 35 miles south of Rock Springs, Porter <£•
Porter 1050S (bry, rm). He found, intermixed with this species and E. brevicaule {10509}, a series
of hybrids that would key out to the var. albogilvum (10510). However, unlike the Utah plants of
var. atbogiloum, the Wyoming plants seem to be only first generation hybrids.

Dec. 30, 1967 eriogonum corymbosum complex 213

extends as far south as this kind of soil is found or to where the
sandy desert and E. corymbosum var. orbiculatum. are encountered
(Map 3).

Even though it is possible to discuss these various populations
which have been given names, their distinction in the herbarium
and in the field is not as great nor as consistent for recognition with
as much certainty as those given formal taxonomic rank in this
paper. In coming to this conclusion, no consistent separation was
found possible when the geographical location was disregarded. Ad-
ditional critical and detailed study may show that some of these
forms can be distinguished, but if they are, other populations which
are equally as distinct, will have to be recognized. At present,
plants from Arches National Monument and the Capital Reef area
are different, as are the plants in the foothills north of Bryce Can-
yon. Thus, as I see it. the var. corymbosum is a large and still vari-
able taxon that is undergoing still more geographical isolation and
speciation.

In southwestern Utah and particularly in northern Arizona, the
var. corymbosum is similar to var. glutinosum in several features,
and at times the two seem to differ only in flower color. Both occur
in similar habitats, and often may be found growing in mixed popu-
lations. Considerable work (and probably genetic studies) is needed
on this part of the species. As far as this paper is concerned, the
two varieties will be basically separated on flower color, but realizing
that in some populations, especially some of those in Arizona, the
distinction may be artificial.

The relationships of E. corymbosum to the other species discussed
in this paper show a reticulated pattern of development, and the
linear arrangement of the species is therefore somewhat misleading.
Probably the entity that comes closest to E. corymbosum is E. hylo-
philum; however, as noted in the discussions of the various species,
this relationship is not clearly understood and highly speculative

During recent field studies in western Colorado (1967), E. saro-
thriforme Gandg. {Bull. Soc. Bot. Belg. 42: 192. 1906) was seen and
studied. Stokes (1936) reported that this species was referrable to
E. corymbosum, but in my opinion, E. sarothriforme is a member of
the E. brevicaule complex. The Colorado species has long, narrow,
green basal leaves at the base of a green and glabrous branching
inflorescence. It occurs mainly in Garfield Co., with a single
floccose-tomentose collection {Reveal & Davidse 860) coming from
Elagle Co.

The following varieties of E. corymbosum are all based on gross
morphology, and the relationships expressed between them are main-
ly speculations.

7b. Eriogonum corymbosum var. erectum Reveal & Brotherson,
var. nov.

A var. corymboso plantis fruticosis ad 6-10 dm altis, ubique
brunneo-tomentosis. foliis lanceolatis vel ellipticis, 2-3.5 cm longis,

214

E. duchenense

E. hylophilum

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

E. saurinum / E. fendlerionum

E. smithii

E. lancifolium

war. olbogilvum

var. davidsei

var. erectum

E. corymbosum
var. corymbosum

var velutinum
7-^

vor. glutinosum

Fig. 9. Diagram showing the possible relationships between the various spe-
cies and varieties in the Eriogonum corymbosum complex.

0.5-1.5 cm latis, inflorescentiis brevibus et compactis, densis, 2.5-7
cm longis, erectis, involucris 2.5-3.5 mm longis, 1.5-2 mm latis,
perianthiis albo-brunneis, costa basi rosea, 2.5-3 mm longis differ.

Erect shrubs (3) 6-10 dm high, brownish-tomentose nearly
throughout; leaves lanceolate to elliptic, 2-3.5 cm long, 0.5-1.5 cm
wide, densely tomentose below, less so to subglabrous and green
above, appressed to the stems and erect, the leaf-blades thick and
stiff; inflorescences mostly short and compact, the branches dense,
2.5-7 cm long; involucres 2.5-3.5 mm long, 1.5-2 mm wide; perianth
brownish-white with reddish midribs and bases, 2.5-3 mm long.
Figure 10.

Type, utah: Wasatch Co. Along U. S. Hwy. 40, 5.5 mi E of
Strawberry Reservoir, 33 mi W of Duchesne, sec. 28, T. 7 S., R. 10
W., elevation 7500 feet, 16 Aug 1966, Noel H. Holmgren & James L.
Reveal 3022. Holotype deposited at utc. Isotypes distributed to
ARIZ, BRY, CAS, Ds, GH, MO, NY, RM, RSA, uc, US, and Other herbaria.

Distribution. Widely scattered in northeastern Utah from
Wasatch Co. to extreme western Uintah Co., Utah, mainly above
6000 feet elevation. Map 3. Flowering mainly from August to
September.

Representative Specimen, utah: Duchesne Co., 10 mi up the Tabiona
Road, Atwood 675 (bry, utc); 10 mi N of Altonah, Brotherson 604 (bry); 5 mi
N of Fruitland, Brotherson 477, 478 (bry); 15 mi SW of Myton, Brotherson 720,
735 (bry); Rock Cr., Brotherson 900 (bry); Hanna, Christensen s.n. (bry); 0.6
mi up Grass Hollow, Indian Creek Canyon, Holmgren & Reveal 3021 (bry, ny,
utc); Pine Hollow, Hatchings s.n. (ariz); 16 mi N of Duchesne, Maguire 12523
(gh); 1 mi W of Mountain Home, Stoddart & Passey s.n. (okl, utc); 7 mi N
of Duchesne, Stoddart & Passey s.n. (okl, utc). Summit Co., Uinta Mts., Milner
6766b (ut). Uintah Co., Dry Fork Mt., Andrews & Noble s.n. (ds, ws); 5 mi

Dec. 30, 1967 eriogonum corymbosum complex

215

2mm

Fig. 10. Habit sketch of Eriogonum corymbosum var. erectum showing the
erect stems and stem leaves with an enlarged drawing of a single involucre with
several exserted flowers.

NW of Whiterock. Brotherson 55/ (bry); near Roosevelt, Thome 18926 (rsa).
Utah Co., 12 mi E of Soldier Summit, Reveal & Reveal 724 (bry, ny, utc);
12 mi E of Soldier Summit, Welsh & Moore 5426 (bry). Wasatch Co., Straw-
berry Res., Brotherson 669 (bry); Wolf Cr., Garrett 7487 (cas, ds); between the
head of Daniel's Canyon and Duchesne. Garrett 8037 (mo, uc, ut); 5 mi E of
Strawberry- Res., Maguire 17652 (gh, ids, ny, oki,, uc, utc), 17653 (ny, okl,
uc, UTC, WTu); 2 mi N of the summit of Indian Creek Canyon, Maguire 21324
(gh, utc).

rhe discovery of this variety came as a result of detailed ecologi-
cal studies by Jack D. Brotherson, presently a graduate student at
Iowa State University. Ames. Iowa. Brotherson's master's thesis
(1967). on the ecology of E. corymbosum in the Uinta Basin, dis-

The Great Basin Naturalist
216 JAMES L. REVEAL Vol. XXVII, No. 4

closed that two distinct forms existed in this area. From herbarium
studies, Brotherson and I could easily note var. corymbosum and
var. erectum on the basis of the latter's more erect stems and de-
cidedly brownish tomentose stems and. branches.

In the field, the two taxa are quite distinctive. The var. erectum
has a distinct brownish cast to its tomentum that is readily seen,
and the crowns possess few and more erect stems. The leaves of var.
erectum are generally not as spreading as in the var. corymbosum.
Lastly, as Brotherson (1967) has shown, var. erectum occurs in
scattered patches where it is often associated with Artemisia in
Pinyon-Juniper Woodlands. This is unlike var. corymbosum of the
lower portion of the Uinta Basin which is normally restricted to the
clay hills and slopes where it is often associated with Atriplex. The
distinct elevational differences between the two varieties are prob-
ably best seen in Indian Creek Canyon, southwest of Duchesne.
The upper elevations have scattered plants of var. erectum; however,
as one proceeds down the canyon, var. erectum is reduced in num-
bers and at middle elevations it is missing altogether. At the lower
end of the canyon, long, rolling hills of a clay formation are found,
and on these hills, var. corymbosum is fairly common. There is a gap
of some five to eight miles between the two varieties, as well as
nearly a thousand feet in elevation.

The relationship of var. erectum to var. corymbosum seems to be
rather clear, with the var. erectum occupying a different ecological
niche and geographical range.

7c. Eriogonum corymbosum var. davidsei Reveal, var. nov.

A var. corymboso et var. erecto plantis fruticosis 8-12 dm altis,
ubique argenteo-tomentosis, foliis lanceolatis vel ellipticis, 3-4 cm
longis, (0.5) 1-2 cm latis, inflorescentiis brevibus et compactis,
densis, 3-6 cm longis, patulis, involucris 2.5-3 mm longis, 1.5-2 mm
latis, perianthiis albo-brunnis, costa basi olivacea, 2-2.5 mm longis
differt.

Large spreading shrubs 8-12 dm high and 5-15 (20) dm across,
silvery-tomentose nearly throughout; leaves lanceolate to elliptic, 3-4
cm long, (0.5) 1-2 cm wide, densely tomentose below, less so but
tomentose and whitish-green above, spreading from the stems, the
leaf-blades thin and lax; inflorescences mostly short and compact, the
branches dense, 3-6 cm long; involucres 2.5-3 mm long, 1.5-2 mm
wide; perianth brownish-white with olive-green bases and thin,
nearly indistinct, midribs, 2-2.5 mm long. Figure 11.

Type, utah: Carbon Co. 0.7 mi S of U. S. Hwy. 50-6 at Well-
ington just south of the Price River Bridge on the dirt road to Mound,
on steep dark Mancos Shale hills, 9 Sep 1967, James L. Reveal &
Gerrit Davidse 956. Holotype deposited at utc. Isotypes distributed
to ARIZ, BRY, CAS, Ds, GH, MO, NY, RM, RSA, uc, US, and Other herbaria.

Distribution. Known only from the low Mancos Shale hills on
the river bank along the Price River south of Wellington, Carbon
Co., Utah. Map 3. Flowering from August to October.

Dec. 30. 1967 eriogonum corymbosum complex

217

1cm

Fig. 1 1 . Habit sketch of Eriogonum corymbosum var. davidsei showing the
general aspect of the species with an enlarged drawing of a single involucre with
several exserted flowers.

The Great Basin Naturalist
218 JAMES L. REVEAL Vol. XXVII, No. 4

Specimens Examined. Known only from the type although additional living
material was obtained on 12 Oct. 1967 for illustrations and further study.

The var. davidsei seems to be closely related to var. erectum
from which it differs only in a few technical characteristics. Never-
theless, var. davidsei is distinct in its ecological and edaphic require-
ments. As noted above, the var. erectum commonly occurs in the
mountains on sandy soils associated with Artemisia in pinyon-
juniper. The var. davidsei, on the other hand, is foimd on low rolling
Mancos Shale hills south of Wellington, Utah, where it is associated
with Atriplex. The type area of the new variety is along the banks
of the Price River and the plants are exceedingly large and robust,
most unlike the narrowly erect, few branched forms of var. erectum.
The leaf width and length of var. davidsei is the greatest in this part
of E. corymbosum, but it does not approach the leaf length-width
ratio of either var. orbiculatum or var. velutinum.

The most striking morphological feature observed in the field is
the silvery tomentum which is exceedingly soft and felt-like. This
gives the plants a distinct cast against the otherwise dark clay slopes
and allows them to be easily seen from a distance. The crowns
which tend to be densely branched with numerous short flowering
inflorescences also give the plants a distinct appearance as the re-
sulting round-shaped crowns are a mass of flowers, although the
crowns are not as dense as in var. orbiculatum. When this plant
was collected in September, it was necessary to scare away numerous
honeybees which were probably from local hives in the Wellington
area. Some small flies were also seen on the flowers.

This variety is named for Mr. Gerrit Davidse, a graduate student
in botany from Utah State University, Logan, who collected with
me on an extended trip into Texas and adjacent states in August
and September, 1967.

7d. Eriogonum corymbosum var. albogilvum Reveal, var. nov.

A var. erecto plantis subfruticosis, (1.5) 2-4 dm altis, foliis elhp-
ticis, 1-2 (2.5) cm longis, 4-7 mm latis, supra virido-tomentosis.
subra subglabris et viridis, inflorescentiis brevibus et compactis, 1-3
cm longis, involucris 2.5-3 mm longis, 1.5-2 mm latis, perianthiis
albo-gilvum vel gilvum differt.

Erect subshrubs, (1.5) 2-4 dm high, whitish to brownish tomen-
tose nearly throughout; leaves elliptical, the leaf-blades 1-2 (2.5) cm
long, 4-7 mm wide, densely greenish- white tomentose below, sub-
glabrous and green above, appressed to the stems and erect, the
petioles 1-3 mm long; inflorescences short and compact, 1-3 cm
long; involucres 2.5-3 mm long. 1.5-2 mm wide; perianth pale yel-
low to yellow with reddish-brown midribs and bases. 2.5-3 mm long.
Figure 12.

Type, utah: Duchesne Co. About 3 mi N of the Indian Creek
Canyon Summit along Utah Hwy. 33, on steep hillside, sec. 1, T. 10
S., R. 8 W., elevation 8500 feet, 26 Aug 1966, James L. Reveal &
Caroline G. Reveal 726. Holotype deposited at utc. Isotypes will

Dec. 30. 1967 eriogonum corymbosum complex

21Q

2mm

Fig. 12. Habit sketch of Eriogonum corymbosum var. albogilvum showing
the erect stems and short compact inflorescences with two enlarged sketches of the
basal node of the inflorescence which shows three involucres and several exserted
flowers subtended by bracts, and a single enlarged involucre with several e.xserted
flowers.

be distributed to ariz. bry, cas, ds, gh, mo, ny, rm. rsa, uc, us, and
other herbaria.

Distribution. Infrequent and widely scattered in the upper
end of Indian Creek Canyon, east of Soldier Summit at the head of
Price Canyon, and north of Duchesne, in Duchesne and Utah coun-
ties, Utah. Map 3. Flowering mainly in August and September.

The Great Basin Naturalist

220 JAMES L. REVEAL Vol. XXVII, No. 4

Specimens Examined, utah: Duchesne Co., 8 mi N of Duchesne along
Utah Hwy. 87, Welsh <& Moore 6775 (bry). Utah Co., 12 mi E of Soldier Sum-
mit, Reveal <£• Reveal 725 (bry, ny, utc); 12 mi E of Soldier Summit, Welsh &
Moore i425 (bry).

It is suggested that E. corymbosum var. albogilvum is of a hybrid
origin between E. corymbosum var. erectum and E. brevicaule Nutt.
var. laxi folium. (Torr. & Gray) ReveaP or var. pumilum S. Stokes ex
M. E. Jones. Evidence for this statement is based on field observa-
tions and on pollen analysis. In the field, var. albogilvum is always
associated with the two suggested parents, and is intermediate in
several respects to them. The var. albogilvum is similar to var.
erectum. in its overall morphology, and is not at all similar to E.
brevicaule var. laxifdlium which is low and nearly caespitose with its
tomentose stems less than 10 cm long and topped by a capitate in-
florescence. Nor is the var. albogilvum similar to var. pumilum
which has an umbellate inflorescence and otherwise similar to var.
laxi folium. The flowers of var. erectum are brownish- white, while
those of var. laxifolium and var. pumilum are yellow. The flowers
of var. albogilvum are mostly pale yellow, although the plants from
north of Duchesne {Welsh & Moore 6775) vary from pale yellow to
bright yellow, and thus of an intermediate condition. The stem and
branch pubescence is similar to var. erectum, but the leaf pubescence
which is greenish and nearly glabrous above, is similar to var. laxi-
folium and var. pumilum. The leaf shape, however, is similar to var.
erectum and not at all similar to the leaves found in the two varieties
of E. brevicaule.

From the pollen analysis of the type collection, it has been found
that the var. albogilvum has from 80 to 90 percent of its pollen
viable, and this seems to show some hybridization has occurred but
that its effect has been lessened, possibly by backcrossing. Both of
the supposed parents were found to have about 100 percent pollen
fertility.

In Indian Creek Canyon and north of Duchesne where the popu-
lations are rather extensive, the var. albogilvum appears to have
become stabilized, and apparently is not backcrossing with var.
erectum. However, at the site east of Soldier Summit, the var.
albogilvum appears to be in danger of becoming swamped by var.
erectum, and this population is not nearly as stable as the other
known populations.

i. The large ami <oniple\ species, Kriogonum brevicaule. is in a current stale of reevaluation,
especially in Utah where several distinct forms are found. Although considerable adjustment must
Ik; made in the rest of the species in Utah, this combination is made so that it can be used in this
paper and in llie Eriogonum treatment for the Pacific Nortliwest I'lora. The var. laxijolium ranges
from 5000 feel to more than 10,500 feet elevation, and may be distinguished from the other varieties
in R. brevicaule by iUs capitate inflorescences. Along tlie middle elevations of the VVasatdi Front,
the var. laxijolium is largely replaced by the var. pumilum S. Slokes ex M. E. Jones. The com-
bination may l)e made as follows:

The following are sjiionyms:

Kriugonum brevicaule Nutt. var. I.ajrijoliuin (Toit. & Gray) Reveal, comb. nov.

E. kingii Torr. & Gray var. laxifolium Torr. & Gray, Proc. Amer. Acad. 8:1()5. 1870.
P.. chrysocephalum A. Gray, Proc. Amer. Acad. 11:101. 1876, a substitute name for var. laxifolium.
E. ochrocephalum S. WaLs. var. angustum M. V.. Jones, Contr. West. Bot. 11:9. lOOS. E. medium
Rydb., Jt'lora Rocky Mis. 220. 1917. E. nudicaulc (Torr.) Small ssp. angustum S. Stokes, Gen.
Eriog. 84. \9i6. E. chrysocephalum A. Gi-ay ssp. bannockcnse S. Stokes, l^afl. West. Bot.
3:200. 1941. E. bannockense attrib. to S. Stokes by R. J. Davis, Flora Idaho 246. 1952.

Dec. 30. 1967 eriogonum corymbosum complex 221

The var. albogilvum presents several difficult problems in regards
to nomenclature. The fact that hybrids exist is no longer questioned,
and even now the International Code (1966) provides for their
nonienclatural recognition. The problem with var. albogilvum is
that it is of a hybrid origin between two different species, and back-
crossed onto one of the parents, and while in this immediate point
of time it seems to be stable, it may not continue to exist as a des-
crete and self-reproducing population.

Several approaches to this problem can be taken. First, the plants
can simply be ignored, or mentioned in the discussion as an occa-
sionally found population. Secondly, the plants can be given a
species hybrid name, but as I understand the Code, this is usually
restricted to sterile F, hybrids, and I think such a concept should
continue. This population may be recognized, taxonomically, as a
mere form, but again, for those who prefer the use of forma, of which
I am not one. these are usually single individuals which differ from
the rest of the population in some (often mutable) single characteris-
tic. Actually, the var. albogilvum. does not really fit into any of
these categories. Variation within species are often introduced from
outside species through hybridization. Yet there is no reason to as-
sume that the same thing cannot happen within a definable part of
the species, or as in our case, within a variety of the species. Nor-
mally such occurrences are unsuccessful, or happened so long ago
that the incoming characteristics have been incorporated within the
population and only by experimental evidence is it possible "to dis-
cover this outside influence.

7e. Eriogonum corymbosum var. orbiculatum (S. Stokes) Reveal &
Brotherson, stat. & comb. nov.

E. effusum Nutt. ssp. orbiculatum S. Stokes, Gen. Eriog. 79.
1936.

E. divergens sensu authors, non E. corymbosum var. divaricatum
Torr. & Gray.

E. jonesii sensu authors, non E. jonesii S. Wats.

E. nummulare sensu authors, non E. nummulare M. E. Jones.

Large subshrubs or shrubs, 3-12 dm high, forming compact,
hemispherical crowns of numerous divaricated branches, the shrubs
often bright green; leaves elliptical-oblong to ovate-orbicular, 1-3
(4) cm long, 1-3 (3.5) cm wide, floccose to tomentose on both sur-
faces, often somewhat less dense above than below, the petioles 5-10
(15) mm long; inflorescences of dense, compact, short branches,
often up to 20 cm long, rigid; perianth white with green or reddish
midribs and bases, 2.5-3 mm long. Figure 13.

Type, utah: Emery Co. Green River, 3 Nov 1915, M. E.
Jones s.n. Holotype deposited at pom! Isotypes: ds, pom!

Distribution. Eastern Utah south of the Book and Roan cliffs
and adjacent western Colorado south along the Green and Colorado

222

JAMES L. REVEAL

The Great Basin Naturalist

Vol. XXVII, No. 4

Fig. 13. Habit sketch of Eriogonum corymbosum var. orbiculatum showing
the narrowly erect stems and compact inflorescences with enlarged drawings of
part of an inflorescence with several involucres and exserted flowers and of the
basal node of the inflorescence with large leaves at this node.

Dec. 30. 1967 eriogonum corymbosum complex 223

river drainages into northeastern Arizona and adjacent northwestern
New Mexico. Map 3. Flowering from late July to early November.

Representative Specimens, arizon.a: Apache Co., Between Many Farms
and Round Rock, Barr et al 1991a (.ariz, ut). Coconino Co., 2 mi below Navajo
Bridge. Cutler ilS9 (gh, ny okl); Bass Canyon, Grand Canyon, Thornber 2S21G
(,.\Riz). Navajo Co., 3 mi N of the Totem Poles. Blass 33 (uc); 8 mi E of Kayen-
ta. Cutler 2876 (ds, ny) ; Kayenta. Eastwood & Howell 6547 (cas, ds, gh, ny.
POM. uc, us); 28 mi SE of Kayenta, Hutchinson 7432 (colo); Monument Valley,
Peebles <£• Fulton 11947 (ariz). Colorado: Montrose Co., 1 mi above IJravan,
Ownbey 1506 (ds, gh, ids, mo, montu, ny, ore, rm, uc, usfs, utc, ws, wtu);
il mi SE of Gateway, Weber 3564 (ariz, bm, cas, colo, ds, montu, okl, ph, rm,
rsa, tex, uc, utc, ws, wtu). San Miguel Co., Basin Cr., Ownbey 1499 (cas, ds,
ids, ny. okl, pom, uc, us, utc, ws, wtu). new MEXICO: Without definite loca-
tion. Mesa la Vaca, Marsh 244 (us); Bad Lands, Wooton 2815 (ariz, us). Mc-
Kinley Co., Chaco Canyon, Clark s.n. (nmu). utah: Garfield Co., 17 mi S of
Escalante, Beck s.n. (bry); 50 mi S of Escalante, Beck s.n. (utc); Tickaboo Can-
yon. Colorado River, Lindsay 21 (uc, wis); Mt. Ellen, Osgood s.n. (us). Grand
Co., Moab, M. E. Jones in 1891 (mo, us), in 1913 (pom); 6 mi N of Moab, Ma-
guire d- Maguire 5852 (utc); 6 mi W of Moab, Maguire & Richards 13301 (gh,
NY, OKL, utc); near Castleton Cr.. Moran 1419 (ds); 8 mi NW of Moab, Water-
fall 11407 (oKL, utc); Courthouse Towers, Arches N. M.. Reveal 686 (ariz, bry,
cas, ds, gh. kans, mo, ny, okl, osc, rm, rsa. tex, uc, us, UT, utc, wis). San
Juan Co., Bridge Canyon, Natural Bridges, N. M., Clover 4039 (ariz); 8 mi NW
of Oljeto Post, Cutler 2251 (cas, gh, mo, us); 1 mi NW of Wide Butte, Cutler
2989 (ds, gh, NY, okl), 2999 (ds, gh, ny, okl, wis); Cooper Canyon, Cutler 3169
(ds, ny, okl); Barton Range, Eastwood 129 (gh, mo, uc, us); Cottonwood Wash,
Ferris 11638 (ds); Rainbow Bridge Trail, Goodman & Payson 3261 (uc. wtu);
3 mi N of Red Mesa, Harrison 12189 (bry); Monument Valley, Holmgren 3850
(gh, ny, uc, utc, wis, ws) ; Monument Pass, Monument Valley, Holmgren &
Reveal 2999 (bry, ny, utc); 8 mi S of Bluff, Maguire et al. 5851 (gh, uc, utc);
15 mi S of LaSal Junct., Reveal 687 (.'\riz, bry, cas, ds, gh, kans, mo, ny, okl,
osc, RM, RSA, tex, UC, US, UT, UTC, wis) ; 1 .2 mi E of Utah highway 47 on Utah
highway 262, Reveal 689 (ariz, bry, cas, ds, gh, kans, mo, ny, okl, osc, rm,
RSA. TEX, UC, us, UT, UTC, wis) ; near The Tanks, Rydberg <& Garrett 9875 (gh,
mo, ny, us. wis); 3 mi S of Kane Spr., Welsh c& Moore 1846 (bry, isc, wis);
Elephant Hill, Welsh et al. 3707 (bry); 1 mi E of Upheaval Dome, Welsh &
Moore 3854 (bry). Waj'ne Co., Marvine Laccolite, M. E. Jones 5663 (mo, ny,
POM, uc, us).

Eriogonum corymbosum var. orbiculatum has been known to
taxonomists under a variety of names for several years. In 1906,
Small proposed a new name for E. corymbosum var. divaricatum,
but when he applied the name, E. divergens, to plant specimens,
they represented the var. orbiculatum instead of the kind of plant
typified by the Gunnison expedition type. Rydberg. in his flora on
the Rocky Mountains (1917). applied the name E. jonesii S. Wats,
to this population, and this error has resulted in the reports of this
species being found in Utah. Actually, E. jonesii is a narrow en-
demic species known only from northwestern Arizona (Reveal, in
press c). While E. jonesii does approach var. orbiculatum in its
wide leaves. E. jonesii has leaves that are cordate and nearly always
basal, and not at all like var. orbiculatum.

In her monograph, Stokes (1936) discusses at some length the
variation in the taxon. and yet. for some reason, she selected as the
type a Jones specimen that is on the very edge of the variation within
what is here defined as var. orbiculatum. Certainly the leaves are
rotund, but they are exceedingly small, and the plants in the type

The Great Basin Naturalist
224 JAMES L. REVEAL Vol. XXVII. No. 4

area tend to be somewhat less hemispheric in shape than those found
further to the south.

The hemispheric crowns in this variety are rather interesting to
observe in the field. The new year's growth begins on the stems of
the previous year and grows slowly throughout the spring and early
summer. As the season moves into July, the branches divide into
the inflorescences which are so profuse as to make the top of the
crown exceedingly firm and rigid. In late August the crowns which
are bright green start to flower. The green branches of the inflo-
rescences are gradually masked by the white flowers so that in Sep-
tember massive rounded humps are seen spotting the red sands in
southeastern Utah.

7f. Eriogonum corymbosum var. velutinum Reveal, var. no v.

A var. orbiculato plantis fruticosis ad 5-10 dm altis, ubique albo-
tomentosis, foliis ellipticis vel oblongis, (1.5) 2-2.5 cm longis, (1)
1.5-2.5 cm latis, inflorescentiis compactis, densis, 4-9 cm longis,
perianthiis brunneo-albis, 2-2.5 mm longis differt.

Large, densely grayish-white tomentose shrubs 5-10 dm high, the
tomentum matted and velvet-like, the plants forming rather compact
crowns of several branches; leaves elliptical to oblong, mostly trun-
cate at the bases, (1.5) 2-2.5 cm long, (1) 1.5-2.5 cm wide, densely
white-tomentose below, less so and brownish-floccose above, the
petioles 0.5-1.5 cm long; inflorescences of dense, compact, short
branches. 4-9 cm long, densely tomentose; perianth brownish- white
with light brown midribs and bases, 2-2.5 mm long. Figure 14.

Type, new mexico: Socorro Co. 7.6 mi E of the junction of
Interstate 25 and U. S. Hwy. 380 near San Antonio, at the base of a
small ridge system in dark, heavy soil, 5 Sep 1967, James L. Reveal

6 Gerrit Davidse 919. Holotype deposited at utc. Isotypes distrib-
uted to ARIZ, BRY, CAS, DS, GH, MO, NY, RM, RSA, UC, US, and Other

herbaria.

Distribution. West-central New Mexico from Sandoval and
Santa Fe cos. south to Socorro Co. on heavy clay, gypsum, or oc-
casionally sandy soils. Map 3. Flowering mainly in August and
September.

Specimens Examined, new mexico: Sandoval Co., San Ysidro, Arsene
19424 (p); San Ysidro, Barneby 12828 (cas, ny); ■>an Ysidro Benedict 225'^
2268, 2302 (us); LaVantaua Coal Mine, Casieller 5756 (nmu); 6 mi NW of
San Ysidro, Reveal & Davidse 925 (bry, ny, utc); 4 mi S of San Luis, Spring-
field 429 (us, usFs); 4 mi N of San Luis, Springfield 733 (nmu). Santa Fe Co.,
3 mi NE of I^s Cerrillos, Bennett 8222 (tex); Los Cerrillos, Herrick 778 (us);
1.8 mi N of Los Cerrillos, Reveal & Davidse 922 (bry, ny, utc). Socorro Co..

7 mi E of San Antonio, Barneby 12895 (cas, ny); 3 mi E of San Antonio, Dunn dc
Lint 4630 (nmu).

The var. velutinum is the large and distinctive form of the spe-
cies found in west-central New Mexico and thus largely isolated
from the rest of the species. Like var. orbiculatutn, the var. velu-
tinum is rather uniform throughout its range even though it occurs

Dec. 30. 1967 eriogonum corymbosum complex

225

Fig. 14. Habit sketch of Eriogonum corymbosum var. velutinum showing the
general aspect of the plant with an enlarged drawing of part of the inflorescence.

on a wide variety of edaphic conditions. For the most part, the var.
velutinum is on clay slopes on the low foothills of the mountains,
althouf^h near Los Cerrillos it is fairly common on sandstone out-
crops and northwest of San Ysidro it is usually on gypsum slopes.
The plants tend to be most robust on the clay soils and the least ro-
bust on the gypsum soils.

In the field, this variety is distinguished by its whitish-brown
or whitish-green color of the densely matted tomentose stems and
branches and the brownish-white color of the flowers. The plants
have rounded crowns, but these are not nearly as dense as the var.
orbiculatum. This is due to the fact that the branches of var. velu-
tinum lend to be more erect and do not normally lay on the ground.
The leaves which are spreading, are not as round as var. orbicula-
tum, but tend to be slightly longer than broad. While the tomentum
of var. velutinum is matted and dense, it is not nearly as soft as var.
davidsei.

The Great Basin Naturalist

226 JAMES L. REVEAL Vol. XXVII, No. 4

7g. Eriogonum corymbosum var. glutinosum (M. E. Jones) M. E.
Jones, Contr. West. Bot. 11: 14. 1903.

E. aureum M. E. Jones, Proc. Calif. Acad. IL 5: 719. 1895. E.
fruticosum A. Nels., Bot. Gaz. 34: 23. 1902, a superfluous substitute.
E. microthecum Nutt. ssp. aureum S. Stokes, Gen. Eriog. 76. 1936.
(Type: St. George, Washington Co., Utah, 28 Sep 1894, M. E. Jones
6091. Holotype: pom! Isotypes: mo, ny, uc, us!)

E. aureum var. glutinosum M. E. Jones. Proc. Calif. Acad. II, 5:
720. 1895. E. fruticosum var. glutinosum A. Nels., Bot. Gaz. 34: 23.
1902.

E. crispum L. O. Will., Bull. Torrey Bot. Club 59: 427. 1932.
E. microthecum Nutt. var. crispum S. Stokes. Gen. ELriog. 76. 1936.
(Type: Cedar Canyon, east of Beaver, Beaver Co., Utah, 2 Sep 1931,
A. O. Garrett 6027 . Holotype: rm! Isotyjae: ut!)

Subshrubs or shrubs 2-8 (10) dm high, forming large open to
compact, hemispheric crowns with few to many divaricated
branches; leaves lanceolate to oblanceolate or elliptic, 1-4 cm long,
0.5-1.5 cm wide, the petioles 5-10 mm long; inflorescences cymose,
the branches short, glabrous to tomentose, 3-10 cm long; involucres
1-2 mm long, 1-1.5 (2) mm wide; perianth yellow, 1.5-2.5 mm
long. Figure 15.

Type. Arizona: Navajo Co. Holbrook, 22 Aug 1883, Rusby
808. Holotype deposited at us! Isotypes: mich, mo, ny, ore, uc, us!

Distribution. Southwestern and south-central Utah southward
to north-central Arizona where it occurs mainly on sandy soils.
Map 3. Flowering mainly from late July to early October.

Representative Specimens. Arizona: Apache Co., Petrified Forest N. M.,
Borell s.n. (uc); Chinle, Demaree 38524 (okl); Crazy Cr., Eastwood & Howell
6898 (gas, us); Adamana, Rusby s.n. (ny). Coconino Co., Lee Canyon, Clover
7241 (mich); Houserock Valley, Eastwood & Howell 6482, 6483 (cas); Metor
Crater Eastwood & Howell 6915 (cas, us) ; Sunset Crater, Gaines 1310,
311, 1312 (ws); Cape Royal, Grand Canyon, Goodman 6184 (okl); 20 mi NW
of Winslow, Hall 11178 (uc); Cameron, Hanson A190 (cold, mo, ny, osc);
Wupatki N. M., D. J. Jones 157 (ariz, ny), 54-1939 (ariz, cas); Billings, M. E.
Jones 4708 {.\v.ix, bm, cas, cold, gh, ny, pom, us, utc); i4 mi W of Cameron,
Kearney <& Peebles 12823 (ariz, ny, us), 12824. 12825 (ariz, us); San Francisco
Mts., Knowlton 217 (gh, us), 249 (us); Sunset Mt., Purpus 32, 33 (mo, uc, us);
Point Sublime, Grand Canyon N. P., Reveal 695 (ariz, bry, cas, ds, gh, kans,
MO, NY, osc, RM, RSA, TEX, UC, US, UT, UTC, wis) ; 4 mi N of Cosnino, Wetherill
s.n. (ariz); near Flagstaff, Whiting 916/3214 (ariz, ny). Mohave Co., W of
Peach Spr., Barneby 5005 (c.vs, ny); Toroweap, Cottam 13950- (ariz, cas, ut) ;
3 mi E of Chloride, Darrow & Gould 3756 (ariz). Navajo Co., Mishongnovi,
Cutler 3101 (ds, ny, okl); Little Colorado River, Hough 90 (us); Hopi Indian
Res., Whiting 854/2831 (ariz); Holbrook, Zuck s.n. (us), utah: Beaver Co., 2
mi N of Beaver, Maguire 17538 (gh, okl, uc, utc, wtu), 17539 (okl, utc).
Garfield Co., 8 mi NE of Henrieville, Cottam 14112 (pom, ut) ; 20 mi SW of
Escalante, Holmgren et al. 2413 (bry, cas, gh, mo, ny, osc, rm, rsa, tex. uc, us,
UT, utc); near Widtsoe, Smith s.n. (utc). Iron Co., 8.7 mi SE of Cedar City.
Dress 4758 (bm); 10 mi E of Cedar City, Hitchcock et al. 4621 (cas, ds, isc,
montu, ny, okl, pom. rsa, uc, us, utc, ws, wtu); 5 mi E of Cedar City, Ma-
guire & Richards 13292 (gh, uc, utc). Washington Co., Zion N. P., Bacigalupi
2799 (cas, ds. rsa); Gunlock, Cottam 5358 (bry, ut) ; Beaver Dam Mts., East-

Dec. 30. 1967 eriogonum corymbosum complex

227

Fig. 15. Habit sketch of Eriogonum corymbosum var. glutinosum showing
the general aspect of the plant with enlarged drawings of the inflorescence and an
involucre with exserted flowers.

wood & Howell 6518 (c.\s, ds, uc); Jackson Road, Higgins 810 (bry); 1 mi N of
Hurricane, Maguire & Blood 4574 (gh, uc, utc); St. George, Palmer 6787 (bm,
MO. ny); 5 mi W of Santa Clara, Reveal <& Reveal 586 (c.\s, mo, utc).

The distinction of E. corymbosum var. corymbosum and var.
glutinosum. has been discussed previously under var. corymbosum.
and the only point that needs to be noted here is that the area of
overlap between the two forms is small, and thus the distinction
presented here is valid. While some mixed populations (such as
that west of Cameron, Coconino Co., Arizona) seem to be separated
only on flower color, the vast majority of specimens present no prob-
lems. As the var. glutinosum occupies a distinct geographical area
and can be separated by morphological characteristics, it is recog-
nized at the varietal level.

When Jones (1895) described E. aureum. he recognized two va-
rieties. Two of the taxa proposed, E. aureum and the var. gluti-
nosum, are the same kind of plant, and when he recognized this in
1903, Jones reduced the species to a variety of E. corymbosum. The
third taxon. var. ambiguum. however, is the yellow-flowered form

The Great Basin Naturalist
228 JAMES L. REVEAL Vol. XXVII. No. 4

of E. microthecum from the southern Sierra Nevada, and this has
only been recently noted by Reveal (In Munz, in press). Stokes
must have based the concept of her "ssp. aureurri" to a large degree
on the Sierra Nevada material as she placed this taxon under E.
microthecum rather than under E. effusum Nutt. where the remain-
der of the E. corymbosum complex had been placed.

Jones distinguished the var. aureum from the var. glutinosum
on the basis that the first had glabrous stems while the second had
tomentose stems. This distinction does not seem to be of much im-
portance here, a conclusion that Jones reached some years ago.

In the field, var. glutinosum occurs in a wide variety of habitats.
In the St. George, Washington Co., Utah area, and to some degree,
to the south, the variety occurs on sandy soils. In the Grand Canyon
it may be found clinging to the canyon walls. In much of its Utah
range, it occurs among broken rocks and ledges in protected areas,
unlike var. corymbosum which is usually found in the more open
sites and on clay hills.

Literature Cited

Brotherson, J. D. 1967. A study of community relationships of Eriogonum
corymbosum Benth. in DC. in Uintah Basin, Utah. Unpublished master's
thesis, Brigham Young University.

Davis, R. J. 1952. Flora of Idaho. Dubuque, Iowa.

Gandoger, M. 1906. Le genere Eriogonum (Polygonaceae). Bull. Soc. Bot.
Belg. 42:183-200.

Gray, A. 1876. Miscellaneous botanical contributions. Proc. Amer. Acad. 11:
71-104.

Hanson, C. A. 1962. Perennial A triplex of Utah and the northern deserts.
Unpublished master's thesis, Brigham Young University.

Holmgren, A. H. 1962. The vascular plants of the Dinosaur National Monu-
ment. Utah State University, Logan, Utah.

Jones, M. E. 1895. Contributions of Western Botanv. No. VII. Proc. Calif.
Acad. Sci. II, 5:611-733.

. 1903. Eriogonum. Contr. West. Bot. 11:4-18.

Lanjouw, J. et al., editors. 1966. International Code of Botanical Nomencla-
ture. Regnum Vegetabile vol. 46. Utrecht. Netherlands.

& F. A. Stafleu. 1964. Index Herbariorum. 5th Edition. Regnum

Vegetabile vol. 31. Utrecht, Netherlands.

Nelson, A. 1902. Contribution from the Rocky Mountain Herbarium. III. Bot.
Gaz. 34:21-35.

. 1904. New plants from Wyoming. XV. Bull. Torrev Bot. Club 31:

239-247.

Reveal, J. L. In press a. New species of Eriogonum from Utah. Madrono.

. In press b. A new annual Eriogonum from Utah. Ann. Mo. Bot. Gard.

. In press c. Eriogonum jonesii S. Wats. Southwest. Nat.

. IN: MUNZ, P. A., in press. Eriogonum. Supplement to A California

Flora. University of California Press, Berkeley.

Rydberg, P. A. 1912. Studies on the Rocky Mountain flora. XVII. Bull. Tor-
rey Bot. Club 39:301-328.

. 1917. Flora of the Rocky Mountains and adjacent Plains. New York,

New York.

Dec. 30. 1967 eriogonum corymbosum complex 229

Small, J. K. 1906. Studies in North American Polygonaceae. II. Bull. Torrey
Bot. Club 33:51-57.

Stokes, S. G. 1936. The genus Eriogonum, a preliniinary study based on geo-
graphic distribution. San Francisco, California.

. 1943. Further studies in Eriogonum. V. Leafl. West. Bot. 3:200-202.

TiDESTROM, I. T. 1925. Flora of Utah and Nevada. Contr. U. S. Nat. Herb.
25:1-665.

Torrey, J. & A. Gray. 1857. Botany in: E. G. BECKWITH. Report of explora-
tions for a route for the Pacific railroad, on the line of the forty-first parallel
on north latitude. 2:115-132.

. 1870. A revision of the Eriogoneae. Proc. Amer. Acad. 8:145-200.

Welsh. S. L. 1957. An ecological survey of the vegetation of the Dinosaur
National Monument, Utah. Unpublished master's thesis, Brigham Young
Universit>'.

& E. M. Christensen. 1957. An ecological study of the vegetation of the

Dinosaur National Monument. Proc. Utah Acad. 34:55-56.

Williams, L. 0. 1932. Field and herbarium studies. I. Bull. Torrey Bot. Club
59:427-429.

WooTON, E. & p. C. Standley. 1913. Descriptions of new plants preliminary
to a report upon the flora of New Mexico. Contr. U. S. Nat. Herb. 16:109-196.

D ELDEN BECK (1906-1967)

Vasco M. Tanner'

D Elden Beck, the son of Mitchel Robertson and Ruth Davis
Beck, was born at Spanish Fork, Utah, April 11, 1906, and died
August 9, 1967 at Provo, Utah. He married May 31, 1933, Florence
Robinson, of Provo, Utah. Four children were born to this couple:
Janet (Mrs. Jon G. Clark); Brent Robinson; Linda (Mrs. R. Brent
Bullough) all of Provo, and Larry Robinson of Magna, Utah. Pro-
fessor Beck's father was a descendant from John Forsyth Beck who
came to Utah in 1850 from Freehold Township, Buck County,
Pennsylvania. His mother was the daughter of Charles Augustas
Davis who emigrated from Boston, Massachusetts, to Utah in 1849.
Elden's parents were married in 1904 and resided in Spanish Fork
and environs until 1914 when for economic reasons, they moved to
McGill, Nevada, where his father was employed at the mines.
Young Beck began his grade school work there. Life in a mining
camp was new and different. In his boy scout activities, he min-
gled with youngsters from families of many nationalities. He with
his pals explored the region round-about engaging in boyish pranks,
camping, target practicing, collecting insects, rocks, plants, pine nuts,
and animals.

As World War I ended, mining operations dropped off and the
Beck family was numbered among the many to leave McGill in
1921. The move took them to Lava Hot Springs, Idaho, where they
engaged in road construction work and farming. Here they were
in a new environment for the fifteen-year-old lad who was begin-
ning to manifest an interest in nature. The meadows, Portneuf River,
and hot mineral sj)rings were new and challenging conditions to him.
He often remarked that Roscoe E. Davis, a teacher in the high
school at Lava Hot Springs, was an inspiration to him. They went
into the fields together collecting plants, animals, and rocks, but
above all, Davis encouraged Elden to continue his education after
graduation from high school.

When Beck graduated from the Lava Hot Springs High School
in 1925, he began preparation to enter the Brigham Young Univer-
sity. In his freshman year he elected to take courses in chemistry
and general zoology which gave him an introduction to the sciences.
Due to lack of funds, he lived in Spanish Fork with his grandmother
Matley (on his mother's side), and commuted each day on the elec-
tric interurban train, a distance of 12 miles. In his sophomore year,
he batched in Provo with Fred Rowland, a classTnate and chum from
Lava Hot Springs, until January when due to the death of his grand-
father Davis in Spanish Fork, he decided to help his grandmother by
staying with her and commuted to Provo. At the University, he

1. Professor of Zoology and Kiitoiiiology. lirifjliani Yiiurin University. Provo. Utali.

230

Dec. 30, 1967

D ELDEN BECK

231

D Elden Beck, entomologist, invertebrate zoologist,
Brigham Young University, 1938-1967.

had become deeply interested in entomology. 1 le arrived early at
the laboratory and was one of the last to leave at night. Laboratory
hfe was still in the tradition of close contact between teacher and
student, and the activities of teaching and research were still so
closely allied that intellectual relations struck root deeply. At any
rate, the teachers and students with whom Elden Beck worked in this
period of his career had a j)owerful influence in bringing out his
latent possibilities and ripening his inclination toward natural his-
tory. Such classmates as Clarence Cottam, C. J. D. Brown, Irvin

The Great Basin Naturalist
232 VASCO M. TANNER Vol. XXVII, No. 4

Rasmussen, C. Lynn I lay ward, W. Thalman Hasler, Allen Rowe,
and Wilford Olsen continued their graduate studies and all have
been awarded Ph.D. degrees in zoology or entomology. During
EJden's first year at the University, his social and athletic activities
were neglected. To compensate for this, he enlisted and served for
three years in the Medical Corps of the National Guard.

In the summer of 1928, Beck was chosen to participate in the
third biology field expedition. A group of five — ^Fred Richan, Elden
Beck, Orlin Biddulph, advanced biology students, under the direction
of Drs. Vasco M. Tanner and Walter P. Cottam, spent six weeks in
field survey and collecting in southern Idaho, and along the Utah-
Nevada state line to the St. George region. Beck enjoyed this very
much as he was an excellent collector and field student. During his
senior and graduate years, he was a laboratory instructor in general
zoology. In 1929 he was awarded the Bachelor of Arts degree and in
1930 a Master of Arts degree in zoology and entomology. His out-
look on life was enlarged when in the spring of 1930 he was granted
a research assistantship in entomology at Iowa State College, Ames,
Iowa. The next three years were spent in graduate study with Drs.
Drake, Wellhouse, Harris, Knight and Richardson. Elden also had
the following Utah student friends with whom he associated at
Ames: Francis Mortenson, Arvil Stark, Ara Call, Quinten Anderson,
Delbert Greenwood, and staff members Drs. C. Y. Cannon and
Rudger Walker.

During most of the summer vacation in 1931, Beck and Ara Call
collected insects in Kansas, Oklahoma, New Mexico, Northern Mexi-
co from El Paso to Casas Grandes, Magdalena, and Nogales. This
collection was shared with the entomologists at Iowa State College
and Brigham Young University.

The year 1933 was important in the life of D E. Beck. In the
spring of the year he was appointed head of the Biology Department
at Dixie College; he and Florence Robinson were married on May
31 ; and in June he was awarded the Ph.D. degree.

Beck finished his class work and thesis — A Morphological Study
of the Male Genitalia of Various Genera of Bees and was graduated
with a Ph.D. degree from the Department of Elntomology at Iowa
State College (now Iowa State University) in June 1933. This same
year he was appointed head of the Department of Biology at Dixie
College, St. George, Utah. This position was made vacant by the
death of Dr. William Harrison, a Brigham Young University alum-
nus, who had recently graduated from Iowa State College, with a
Ph.D. degree in biology; Dr. Harrison was a very good teacher and
community worker. President Joseph K. Nicholes of the college was
anxious to fill this position with an equally promising candidate.

Dr. Beck soon ingratiated himself with the students and the
people of the community. He went to considerable effort to take his
students on field trips in order that he might lead them into a first-
hand acquaintance with the flora and fauna of the St. George area.
His trip into Mexico in 1931 so fascinated him that he ventured
again in the summer of 1934 to spend six weeks collecting in North-

Dec. 30. 1967 d elden beck 233

ern Sonora as well as on the plateau area around Mexico City. This
collecting party consisted of his wife Florence and a student, Floyd
Atkin. Their bounteous collection of insects and reptiles was used
to great advantage in teaching.

Elden's artistic and helpful wife, Florence, while studying art
with Mr. Ralph lluntnian. artist at the college, persuaded him to try
landscaj)e painting which resulted in his interest in painting and the
organization in 1934 of a Fine Arts Festival at the college, which
has continued to the present time.

An opportunity to return to his alma mater, the Brigham Young
University, as assistant professor of zoology and entomology was
embraced in 1938. Dr. Beck's addition to the zoology staff was wel-
comed since he added impetus to field biology study, development
of an enlarged program in invertebrate zoology, and research. In
1942, he wrote a laboratory guide for general zoology now in the
4th edition (Burgess Publishing Company).

World War II beckoned Dr. Beck to volunteer and serve in the
Medical Entomological Service. His first assignment was at Camp
Barkley, Abilene. Texas, (April 1943 - July 4, 1943) for basic train-
ing at the Medical Replacement Training Center. On July 4, 1943,
he was transferred to Fort McPherson and Fort Benning. Georgia,
for mosquito survey and control work. Here he remained until
December 11. 1943. when he was again transferred this time to
Camp Ellis, Illinois. At this camp a malaria survey unit was or-
ganized with Lieutenant Beck in command. On January 14, 1944,
he left for Camp Plauche near New Orleans, Louisiana, to receive
training preparatory to going over seas. He set sail from San Fran-
cisco April 20, 1944. arriving in Guadalcanal. Solomon Islands, on
May 19, 1944. During the year of continental service, Lieutenant
Beck made valuable collections of insects, amphibians, and reptiles
in Texas. Georgia, and Louisiana, which he sent back to the Univer-
sity at Provo.

Collecting had long been a passion with Dr. Beck. During the
twelve months — May 19. 1944 to May 22, 1945 — spent on Guadal-
canal, aside from the duties as director of mosquito control activities
at the 20th Station Hospital on the J'enaru River where he distin-
guished himself in this capacity, he did considerable collecting of
insects and reptiles. He not only collected several hundred frogs,
lizards, and snakes, but he made a large general collection of most
of the insect orders. These specimens were so well prepared for
shipping that very little loss resulted from breakage and damage.
All the collecting was done within a radius of five miles of the hos-
pital. This area is in the low. sloping plains consisting of swamps,
grasslands, and jungles. All of the specimens collected by him were
sent to the Brigham Young University where the writer, with stu-
dent help, spent much time pinning, labeling, and preserving the
thousands of specimens he collected.- Dr. Beck not only worked

2. Vascr) M. Tanner, 1948. Pacific islands llcrpctolngy No. 1. - .Mariana Islands. GrrnI Basin
Xaluralisl. Vol. IX, Nos. 1-2, pp. 1-20, 19t\ . Pacific Islands llcrpctolngy. No. V - Guadalcanal,
Solomon Island*: A Check I.i<t of Specio'c. Vol. XI. Nos. ^-4. pp. 6V8(i.

The Great Basin Naturalist
234 VASCO M. TANNER Vol. XXVII, No. 4

hard as a collector but he directed the efforts of many students in
that war area in their collecting and shipping of specimens to the
Brigham Young University. Some of the University alumni mem-
bers who cooperated with Dr. Beck in this project were Ernest
Reimschiissel, Doyle Taylor, Cluff Hopla, and Herbert Frost. Wee-
vils of the tribe Celeuthetini he collected have served as a nucleus
for a critical study of the species of five genera found in the Solomon
Islands.

On May 22, 1945, Captain Beck (Lieutenant Beck was promoted
to Captain in February 1945) returned to the United States, and
arrived in San Francisco on June 19, 1945. From here he was sent
to Camp Carson General Hospital in Colorado where he received
medical care until October 10, 1945. when he was honorably dis-
charged to return to his wife and family. Florence, his faithful and
efficient wife, and four children had, with the help of members of
her family and friends, carried on in a stoic manner. Her worries,
loneliness, and heartfelt anxiety for Elden's safety and whole return
were gallantly borne.

The experience gained in mosquito control methods during the
war was put to good use by Dr. Beck. He was assigned by the Utah
County Commission and Health Department in 1946-47 to develop
a mosquito control program for Utah County. He was also active in
civic affairs serving as President of the Provo Junior Chamber of
Commerce. For his service he was awarded the distinguished service
award by this organization in 1947. He was also editor of the Provo,
Utah, Centennial Souvenir 1849-1949.

In 1951-52 Elden was granted a sabbatical leave from the Uni-
versity for the purpose of doing research work at the American
Museum of Natural History, New York City. Here his contact with
Libby Hyman, great authority on invertebrate zoology, was invalu-
able. Several months were also spent at Lake Placid, Florida, where
he became acquainted with Dr. James G. Needham. A first edition
of an invertebrate zoology laboratory guide was finished at this time.
A second and third edition of this manual with Dr. Lee Braithwaite
as co-author has been widely adopted by more than 200 colleges and
universities of the United States.

Dr. Beck began making a collection of earthworms about 1948,
since he was aware of the lack of in depth knowledge of the earth-
worms fauna of the western United States. He sent his collection of
these worms to the American Museum of Natural History, New
York City. Dr. G. E. Gates, University of Maine, Orono, Maine,
authority on earthworms of the United States, graciously decided to
study Beck's collection. His recently published paper dealing with
The Earthworm Fauna of the Great American Desert and Adjacent
Areas^ is based largely upon Dr. Beck's collection. In a letter to the
writer (October 16, 1967) Dr. Gates comments as follows on Beck's
earthworm collection and his (Gates) article referred to above.
"Data from study of Beck's earthworms will be appearing from time

3. G. E. Gates, I9f)7. The Earthworm Fauna of the Great American Desert and Adjacent
Areas. Great Basin Naturalist. Vol. 27. No. ^. pp. 142-176.

Dec. 30. 1967 d elden beck 235

to time in various contributions, but such data will largely be inci-
dental and not of the importance that it derives from the Great Basin
collections. In a sense, then, I suppose, that this contribution will be
a sort of memorial — at least to his collecting of megadriles. A second
regret re of Beck's work is that one shipment of his earthworm ma-
terial got lost somewhere (he never knew where or how) and never
reached the American Museum to which he had been sending his
collection. The lot or lots probably would have made the MS.
even more massive."

In 1951 he received a grant from the National Institutes of
Health for a study of the ectoparasites of the mammals of the west-
ern United States. Many small mammals were trapped in Utah,
Arizona, and Idaho from which Beck obtained a large collection of
fleas and ticks.

Dr. Beck's recognized success in mosquito control gained for him
a leave from university duties 1956-1958 to serve as an adviser
under the auspices of the World Health Organization on malaria
mosquito control for the Chinese government in Tiawan. Mrs. Beck
and some of the children accompanied him. While there, he made a
lifelong friend of Dr. Hsieh, parasitologist, who is now engaged as a
parasitologist at llarbel. Liberia. West Africa.

Dr. Beck was advanced to Professor of Zoology and Entomology
and made chairman of the department at Brigham Young Univer-
sity, serving from February 1962 to June 1965. During his chair-
manship new' staff members were added and the department in-
creased in enrollment and facilities.

From 1959-1966 Dr. Beck was associated with Dr. Dorald Allred.
associate professor of zoology and entomology at Brigham Young
University, in an ecological study at the Nevada Test Site as princi-
pal investigator 1964-66 and associate investigator 1959-64. This
study was carried forward under an Atomic Energy Commission
grant. The main object of the research project was to make a faunis-
tic inventory of the test site. Many specialists have been involved in
studying the specimens collected by a staff of collectors who worked
under the supervision of Beck and Allred. More than 70 papers
have been published dealing with the invertebrates, vertebrates, and
the ecology of the test site.

Two of the papers merit comment at this time. The Orthoptera
of the Nevada Test Site,* by Andrew H. Barnum, at Dixie College,
1964, is a noteworthy contribution. In this study approximately
8.000 specimens of Orthoptera were collected and studied. Species
from four of the five recognized suborders. 9 families, 41 genera, and
60 s[)ecies are recognized in the study. There are 157 well-executed
morphological figures, 34 distribution maps, along with keys and
description to the taxa involved. This contribution of 134 pages adds
greatly to the knowledge of the American Desert Orthoptera of the
Great Basin. Two species of camel crickets are described as new to
science.

4. Andrew H. Bamum. \'Mt4, Orthiplcra of the .\c'vada Test Site. Brigham Young University
Science Bulletin; Biological Series - Vol. IV, No. i. pp. 1-1 H.

The Great Basin Naturalist
236 VASCO M. TANNER Vol. XXVII, No. 4

The magnum opus of these pubhcations is one by Dr. Harry H.
Knight, entomologist at Iowa State University, Ames, Iowa. This
study now in press is a Taxonomic Review. Miridae (Hemiptera)
of the Nevada Test Site and Western United States. It is an ex-
haustive treatise of the mirids of the test site and the western United
States. Dr. Knight has put forth great effort in deahng with the 612
species and 122 genera covered in this study. He has recognized
160 species and 50 genera as occurring on the Nevada test site; of
which 3 genera and 96 species are described as new to science.

Dr. Beck is honored by Dr. Knight with the following new genus
Beckocoris laticephalus, and five new species as follows: Chlamy-
datus becki; Nevadocoris becki (this is a new genus); Lopidea becki;
Parthenicus becki; and Phytocoris becki. This volume is dedicated
to the memory of Dr. D Elden Beck.

Great credit is due Drs. Beck and Allred for the success achieved
in accomplishing the main objective of the research project. Much
more is now known about the fauna of the Great Basin than 10 years
ago before the initiation of this project.

Dr. Beck was active in the following learned societies: Society
of Sigma Xi; Utah Academy of Sciences, Arts, and Letters; Ameri-
can Society of Parasitologists; American Wildlife Disease Associa-
tion; American Mosquito Control Association; and Utah Mosquito
Abatement Association.

We should not neglect to report that Professor Beck was an avid
and talented photographer. His photographic files contain more
than a thousand negatives of plants, animals, and geological features,
many of which have been published in national magazines.

D E. Beck was proud of his family. His 11 grandchildren were
anxious to visit their grandfather and were always welcomed into the
well-ordered Beck home. Elden met people easily with a ready and
even voluble speech and he possessed a charm of manner that made
him good company. The instinct to help others seems to have been
one of Dr. Beck's most deep-seated characteristics. He went beyond a
willingness to lend a sympathetic ear, to lend a hand to friends and
students alike. His artistic soul drank in the beauty of the seasonal
changes of mountain, stream, and valley. He was in harmony with
the animate and inanimate make-up of his environment. The color
of man did not wean him from the inner warmth and spirit beneath
the human skin. He had an abiding faith in the justice and power of
his Maker. We will miss, but long remember our colleague and
friend D Elden Beck.

The Following Comprise the Principal Articles (63)
Published by Professor Beck on Scientific Subjects, 1929-1967

1929

Bees of the Sub-family Osminae in the collection of the BYU Zoology Dept.
Bull. Brooklyn Ent. Soc, Vol. 24. pp. 303-306, 1929.
1931

Length of the Developmental Stages of the Horn-fly Haemotobia irritans at
Constant Temperature. Jour. Economic Ent., Vol. 24, 1931.

Dec. 30, 1967 d elden beck 237

1933

A Morphological Study of the Male Genitalia of Various Genera of Bees.
Proc. Utah Acad, of ScL, Vol. 10, pp. 89-137, 1933.
1935

The Place of Physiology and Hygiene in the Secondary Curriculum. Utah
Ed Review., Vol. 28^ pp. 279-280, 1935.
1936

Notes on Utah Water Striders. Proc. Utah Acad, of Sci., Arts, Lett., Vol. 13,
pp. 203-206, 1936.
1937

Capitol Reef. Utah Mag., Vol. 2, No. 19, pp. 6, 15, and 20, 1937.
1939

Social Hygiene. Utah Ed. Rev., Vol. 32, pp. 174-186, 1939.
1942

Life History Notes on the California Gull, No. 1. Great Basin Naturalist,

Vol. 3, Nos. 3 - 4, pp. 91-108, 1942.
Gull Banding Notes at Utah Lake, No. 2. Great Basin Naturalist, Vol. 3,
pp. 55-57, 1942. Junior co-author with Vasco M. Tanner.
1943

Coyote Gulch. Improvement Era, Vol. 4, No. 8, pp. 462-463, 503, 1943.
California Gull - A Comparative Plumage Study. Great Basin Naturalist,
Vol. 4, pp. 57-61, 1943.
1946

Down the Escalante. Utah Mag., Vol. 8, No. 4, pp. 24, 34, 45, 49-50; No. 5,
pp. 32-33, 48; No. 6, pp. 30, 45-47; No. 7, pp. 30-31, 42, 44; No. 8, pp.
30-31, 33, 36-37; No. 9, pp. 30-31, 43-44. (Booklet run as a serial, 1946.)
1947

The Seagull in Utah. Utah Mag., Vol. 9, No. 4, pp. 22-25, 1947.
1949

Provo, Centennial Booklet. (Exlitor, and feature articles author) Published by
Provo City, Lorraine Press Printers, Salt Lake City, Utah, pp. 1-75, 1949.

1951

Some Observations on Biology Instructions in Secondary Schools. Proceedings
Eighth Annual Conference on Higher Education: Utah Conferences on
Higher Education. September 1951.

1953

Mite Fauna of Woodrat Nests in Utah. Proc, Utah Acad. Sci., Arts, Lett.,

Vol. 30, pp. 53-56, 1953. Junior co-author with Dorald M. Allred.
Arthropcxi Consortes Found in the Nests of Neotoma cinera acraia (Ord) and

Neotoma lepida lepida Thomas. Proc, Utah Acad. Sci., Arts, Lett., Vol.

30, pp. 43-52, 1953. Senior co-author with A. H. Barnum and Lenard

Moore.
A Study of Some Consortes Found at a Nesting Site of the Northern Cliff

Swallow Petrochelidon albifrons albifrons (Rafinesque). Proc, Utah

Acad. Sci., Arts. Lett., Vol. 30, pp. 39-42, 1953.
A New Species of Acomatacarus (Acarina, Trombiculidae) from Utah.

Great Basin Naturalist, Vol. 13, Nos. 3-4, 1953. Junior co-author with

Dorald M. Allred.
Plague, Plague Distribution and Plague Vectors in Utah. Seventieth Annual

Sigma Xi Address. BYU Chap. Proc, 1953.
1954

Ecological and Distributional Notes on Some Utali Hirudinea. Proc, Utah

Acad. Sci., Arts, Lett., Vol. 31, pp. 73-78, 1954.
Distributional Records of Some Aquatic Coleoptera in Utah. Proc, Utah

Acad. Sci., Arts, Lett., Vol. 31, pp. 52-56, 1954.
Distributional and Natural History Notes on Polycelis coronata in Utah.

Proc, Utah Acad. Sci., Arts, Lett., Vol. 31, pp. 79-82, 1954.

1955

Potential and Capable Vectors of Rocky Mountain Spotted Fever and Plague
in Utah. BYU Sci. Bull., Biological Series. Vol. 1, No. 1, pp. 1-64,
February, 1955.

The Great Basin Naturalist
238 VASCO M. TANNER Vol. XXVII, No. 4

Some Unusual Distributional Records of Ticks in Utah. Juur. uf Parasitology,

Vol. 4-1, No. 2, pp. 1-4, 1955.
The Chiggers of Utah (Acarina: Troinbiculidae) . Great Basin Naturalist,

Vol. 15, Nos. 1-4, pp. 1-26, 1955. Junior co-author with James M.

Brennan.
Seasonal Study of the Tick, Ornithodoros hermsi found in nests of the Desert

Wood Rat, Neotoma lepida lepida in Utah. Proc, Utah Acad., Vol. 32,

pp. 131-135, 1955. Senior co-author with Dorald M. Allred.
1957

Seasonal Population Fluctuations of Mites in Desert Wood Rat Nests in

Central Utah. Ecology, Vol. 38, No. 1, 1957.
1958

Zoology of the Upper Colorado River Basin. 1 . Biotic Communities. BYU Sci.

Bull., Biological Series, Vol. 1, No. 3, pp. 1-74, 1958. Junior co-author

with C. L. Hayward and W. W. Tanner.
Malaria Control and Eradication in Taiwan (Formosa, China). Progress

Report, 1952-1957. (Co-author). Bull. World Health Org. 19., 1958.
1960

Ticks of the genus Ixodes in Utah. BYU Sci. Bull., Biological Series, Vol. 1,

No. 4, pp. 1-42, 1960. Junior co-author witli Dorald M. Allred and

L. D. White.
Mosquito Survey of Central Utah Valley, Utah. Proc, Thirteenth Annual

Meeting of the Utah Mosquito Abatement Assoc, 1960.
Central Utah County Mosquito Survey Studies. Mosquito News, Vol. 21,

No. 1, pp. 6-11, 1960.
Invertebrate Zoolog)' Laboratoiy Workbook. Burgess Publishing Co., Minne-
apolis, Minnesota. 1960. Second Edition, pp. 1-289, 1962. Senior co-
author with h. F. Braithwaite.
Comparative Ecological Studies of Animals Exposed to Nuclear Detonation.

(abstract) Proc, Utah Acad. Sci., Arts, Lett.. Vol. 37, pp. 152-153, 1960.

Junior co-author with D. M. Allred and J. R. Murdock.
1962

Ecological Distribution of Mites on Lizards at the Nevada Atomic Test Site.

Herpetologica, Vol. 18, pp. 47-51, 1962. Junior co-author with D. M.

Allred.
1963

Ecological Distribution of some Rodents at the Nevada Atomic Test Site.

Ecology, Vol. 44, No. 1, pp. 211-214, 1963. Junior co-author with D. M.

Allred.
Comparative Ecological Studies of Animals at the Nevada Test Site. Proc, 1st

Nat. Symposium on Radioecology. Reinhold Publ. Corp., pp. 327-331,

1963. Junior co-author with D. M. Allred.
Range of Movement and Dispersal of Some Rodents at the Nevada Atomic

Test Site. Jour, of Mammalogy, Vol. 44, No, 2, pp. 190-200, 1963.

Junior co-author with Dorald M. Allred.
Ticks of the Nevada Test Site. BYU Sci. Bull., Biological Series. Vol. 4. No. 1,

pp. 1-11, 1963. Senior co-author with D. M. Allred and Elias P. Brinton.
Some Effects of an Underground Nuclear Detonation on Biotic Communities

at the Nevada Test Site. Proc, Utah Acad. Sci., Arts, Lett., Vol. 40, pp.

49-61. 1963. Co-author with D. M. Allred and C. D. Jorgensen.
Nevada Test Site Study Areas and Specimen Depositories. BYU Sci. Bull.,

Biological Series, Vol. 2, No. 4, pp. 1-15, 1963. Co-author with D. M.

Allred and C. D. Jorgensen.
Biotic Communities of the Nevada Test Site. BYU Sci. Bull., Vol. 2, No. 2,

pp. 1-52. 1963. Co-author witli D. M. Allred and C. D. Jorgensen.
Hard Bodied Ticks of the Western United States. BYU Sci. Bull., Vol. 2,

Part 1, No. 3, pp. 1-28, 1963. Co-author with Elias P. Brinton.
Hard Bodied Ticks of the Western United States. BYU Sci. Bull., Vol 2,

Parts 2 - 3, No. 3. pp. 1-21, 1963. Co-author with Elias P. Brinton.
Comparative Ecological Studies of Animals at the Nevada Test Site with

Specific Reference to Their Reaction to Exposures of Nuclear Effects.

Dec. 30, 1967 d elden beck 239

Junior coautlior witli Dorald M. AUred. Off site Ecological Research of

the Division of Biology and Medicine - Terrestrial and Freshwater, U. S.

Atomic Energy Commission, March 1963, pp. 4-6. TID-13358. Edited by

Vincent Schultz.
Mites from Pocket Mice at the Nevada Test Site. Proc. Ent. Soc. Wash.,

Vol. 65, No. 3, pp. 231-233, 1963. Junior co-author with D. M. Allred.
1964

Mites on Reptiles at the Nevada Atomic Test Site. Trans. Am. Micro-

scropical Soc, Vol. 83, No. 2, pp. 266-268, 1964. Co-author with D. M.

Allred.
Observations on Host-Parasite Relationships and Seasonal History of Ticks in

San Mateo County, California. Great Basin Naturalist, Vol. 24, No. 1,

pp. 1-6, 1964. Junior co-author with Carol O. Mohr and Elias P. Brinton.
Arthropod Associates of Plants at the Nevada Test Site. BYU Sci. Bull., Bio-
logical Series, Vol. 5, No. 2, pp. 1-16, 1964. Junior co-author with

Dorald M. Allred.
Close-in Effects of an Underground Nuclear Detonation on Small Mammals

and Selected Invertebrates. Report PNE-266F; Washington: Office of

Technical Services, U. S. Dept. of Commerce, 22 pp. FINAL REPORT.

1964. Co-author with D. M. Allred and C. D. Jorgensen.
1965

A List of Scarabaeidae Beetles of the Nevada Test Site. Great Basin Naturalist,

Vol. 25, Nos. 3-4, 1965. Junior co-author with D. M. Allred.
A List of Some Beeflies of the Nevada Test Site. Great Basin Naturalist, Vol.

25, Nos. 1-2, pp. 5-11, 1965. Co-author with D. M. Allred and E. Johnson.
Ecological Effects of Nuclear Testing on Native Animals at the Nevada Test

Site. Proc, Utah Acad. Sci., Arts, Lett., Vol. 42, Part 2, 1965. Co-author

with D. M. Allred and C. D. Jorgensen.
Identification of the Adults, Nymphs, and Larvae of Ticks in the Genus

Dermacentor koch (Ixodidae) in the Western United States. BYU Sci.

Bull., Biological Series, Vol. 5, No. 4, pp. 1-44, 1965. Co-author with

E. P. Brinton and D. M. Allred.
Nevada Test Site Desert Ecology. Proc, Utah Acad. Sci., Arts, Lett., Vol. 41,

No. 2, pp. 202-209, 1965. D E. Beck et al.
Some Variables to Consider in Field Collections Which Will Affect the Inter-
pretation of Animal Population Data for Parasitic Arthropods. Proc,

Utah Acad. Sci., Arts, Lett., Vol. 42, Part 1, pp. 56-59, 1965.
1966

Mites of Utah Mammals. BYU Sci. Bull., Vol. 8, No. 1, pp. 1-123, 1966.

Junior co-author with D. M. Allred.
Siphonaptera (Fleas) of the Nevada Test Site. BYU Bull., Vol. 7, No. 2,

pp. 1-27, 1966. Senior co-author with D. M. Allred.
Summary of the Ecological Effects of Nuclear Testing on Native Animals at

the Nevada Test Site. Proc, Utah Acad. Sci., Arts, Lett., Vol. 42, No. 2,

pp. 252-260, 1966. Co-author with D. M. Allred and C. D. Jorgensen.
Tingidae, Neididae (Berytidae and Pentatomidae) of the Nevada Test Site.

Great Basin Naturalist, Vol. 26, No. 1-2, pp. 9-16, 1966. Junior co-author

with D. M. Allred.
1967

Predaceous - Scavenger Ants in Utah. Great Basin Naturalist, Vol. 27, No. 2,

pp. 67-78. 1967. Co-author with D. M. Allred and W. J. Despain.
General Zoology Syllabus. Laboratory Workbook, Burgess Publishing Co.,

pp. 1-165. Minneapolis, Minnesota. 4th Edition, 1967.

INDEX TO VOLUME XXVII

The new genera, species, and varieties in this volume appear in bold
type in this index.

A Comparative Study of the Moun-
tain Brush Vegetation in Utah,
p. 59.

Allred, Dorald M. and D Elden Beck,
Article by, p. 11.

Amphicranus quercus, p. 53.

A. rameus, p. 55.

A New Name for a Utah Lepidium,
p. 177.

A New Species of Spinibdella from
Utah (Bdellidae: Acarina), p. 98.

Beck, D Elden, Dorald M. Allred,
and William J. Despain, Article
by, 67.

Cactopinus mexicanus, p. 37.

Chaetophloeus minimus, p. 95.

C. struthanthi, p. 96.

Chramesus bicolor, p. 91.

C. demissus, p. 93.

C. incomptus, p. 90.

C. punctatus, p. 94.

C. vastus, p. 92.

Cnesinus adusticus, p. 87.

C. annectens, p. 86.

C. bicornus, p. 80.

C. carinatus, p. 88.

C. colombianus, p. 84.

C. electinus, p. 82.

C. elegantis, p. 79.

C. foratus, p. 81.

C. myelitis, p. 84.

C. niger, p. 83.

C. retifer, p. 85.

Corthylus petilus, p. 56.

C. splendens, p. 138.

D Elden Beck (1906-1967), p. 230.

Eriogonus pauciflorum var. gnapha-
lodes (Benth. in Hook.) Reveal,
stat. & comb, nov., p. 112.

E. p. var. canum (S. Stokes) Reveal,
stat. & comb, nov., p. 112.

E. p. var. nebraskense (Rydb. ) Re-
veal, stat. & comb, nov., p. 113.

E. p. var. canum (S. Stokes) Reveal,
stat. & comb, nov., p. 114.

E. exilifolium Reveal, p. 114.

Eriogonum corymbosum, Benth. in

DC, p. 205.
E. c. var. albogilvum. Reveal, p. 218.
E. c. var. corymbosum, p. 207.
E. c. var. davidsei. Reveal, p. 216.
E. c. var. erectum. Reveal & Broth-

erson, p. 213.
E. c. var. glutinosum, M. E. Jones,

p. 226.
E. c. var. orbiculatum (S. Stokes)

Reveal & Brotherson, stat. & comb.

nov., p. 221.
E. c. var. velutinum. Reveal, p. 224.
E. duchesnense. Reveal, p. 194.
E. fendlerianum, Benth, in DC, p.

199.
E. hylophilum. Reveal & Brotherson,

p. 190.
E. lancifolium. Reveal & Brotherson,

p. 187.
E. saurinum. Reveal, p. 196.
E. smithil. Reveal, p. 202.
Gates, G. E., Articles by, p. 1, 142.
Gist, Clayton S., Article by, p. 26.
Gnathotrichus dentatus, p. 45.
G. imitans, p. 48.
G. nimifrons, p. 47.
G. perniciosus, p. 47.
Hylastes mexicanus, p. 36.
Jorgensen, Clive D., Article by, p.

98.
Leperisinus guatemalensis, p. 89.
Lepidium barnebyanum. Reveal,

nom. nov., p. 178.
Loganius exilis, p. 119.
Microcorthylus desus, p. 52.
Monarthrum bicavum, p. 51.
M. quercicolens, p. 49.
M. querneus, p. 50.
New Records and Species of Neo-
tropical Bark Beetles (Scolytidae:
Coleoptera), p. 79.
New Records and Species of Neo-
tropical Bark Beetles (Scolytidae:
Coleoptera), Part II, p. 119.
New Species of Bark Beetles (Cole-

241

242

optera: Scolytidae), Mostly Mexi-
can, p. 36.

Nixon, Elray S., Article by, p. 59.

Notes on Eriogonum III - On the
Status of Eriogonum pauciflorum
Pursh, p. 102.

Notes on Eriogonum V - A Revision
of the Eriogonum corymbosum
Complex, p. 183.

On the Earthworm Fauna of the
Great American Desert and Ad-
jacent Areas, p. 142.

On Two Little-Known Species of the
Earthworm Genus Diplocardia,
p. 1.

Phloeotribus simplex, p. 95.

Pityophthorus festus, p. 39.

P. foratus, p. 40.

P. quercinus, p. 40.

Predaceous-Scavenger Ants in Utah,
p. 67.

Problems of Sampling Desert Arth-
ropods Before and After a Ther-
monuclear Cratering Test, p. 26.

Pseudopityophthorus granullfer, p.
42.

P. hondurensis, p. 42.

P. micans, p. 44.

P. tropicalis, p. 43.

Pseudothysanoes thomasi, p. 38.

Reveal, James L., Articles by, p.

102, 177, 183.
Scolytodes amoenus, p. 128.
S. clusiacolens, p. 121.
S. clusiavorus, p. 122.
S. facetus, p. 126.
S. ficivorus, p. 125.
S. ingavorus, p. 126.
S. nanellus, p. 124.
S. perditus, p. 123.
S. proximus, p. 127.
S. venustulus, p. 124.
Scolytus aztecus, p. 120.
Spiders of the Nevada Test Site, p.

11.
Spinibdella mail, p. 98.
Stegomerus, p. 129.
S. chiriquensis, p. 131.
S. mexicanus, p. 133.
S. montanus, p. 132.
S. pygamaeus, p. 130.
S. vulgaris, p. 134.
Tanner, Vasco M., Article by, p. 230.
Tricolus saundersi, p. 139.
T. spectibilis, p. 140.
Wood, Stephen L., Articles by, p.

36, 79, 119.
Xyleborus bicornatulus, p. 137.
X. horridatus, p. 135.
X. horridicus, p. 136.

The Great Basin Naturalist

Founded in 1939 by Vasco M. Tanner

A journal published from one to four times a year by Brigham
Yoimg University, Provo, Utah.

Manuscripts: Only original, unpublished manuscripts, pertain-
ing to the Great Basin and the western United States in the main,
\vill be accepted. Manuscripts are subject to the approval of the
editor.

Illustrations: All illustrations should be made with a view to
having them appear within the limits of the printed page. The illus-
trations that form a part of an article should accompany the manu-
script. All half-tones or sdnc etchings to appear in this journal are
to be made under the supervision of the editor, and the cos-t of the
cuts is to be borne by the contributor.

Reprints: No reprints are furnished free of charge. A price list
for reprints and an order form is sent with the proof.

Subscription: The annual subscription is $2.50 (outside the
United States $3.25). Single number, 80 cents.

All correspondence dealing with manuscripts should be addressed
to the Editor, Vasco M. Tanner, Great Basin Naturalist, Brigham
Young University, Provo, Utah. Other matters such as subscriptions,
reprints, exchanges and other business should be addressed to Ernest
L. Olson, Chairman of University Publications.

Reprints Schedule of The Great Basin Naturalist

Each
Additional

2 pp.

4 pp.

6 pp.

8 pp. 10 pp. 12 pp.

2 pp.

50 copies

$6.00

$7.00

$8.00

$9.00 $10.00 $11.00

$2.00

100 copies

7.00

8.00

9.00

10.00 11.00 12.00

200 copies

8.00

9.00

10.00

11.00 12.00 13.00

300 copies

9.00

10.00

11.00

12.00 13.00 14.00

Covers: $10.00 for first 100 copies, $4.00 for additional 100 copies.

TABLE OF CONTENTS

Notes on Eriogonum - V. A Revision of the Eriogonum

Corynibosum Complex. Illustrated. James L. Reveal .... 183

D Elden Beck (1906-1967). Illustrated. Vasco M. Tanner .... 230

Index _ 240

3 2044 072 231 103

Date Due

-KB-a^mo