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HARVARD UNIVERSITY

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LIBRARY

OF THE

Museum of Comparative Zoology

The Great Basin Naturalist

VOLUME 32, 1972

Editor: Stephen L. Wood

Published at Brigham Young University, by
Brigham Young University

Volume 32 Nos. 1-4

TABLE OF CONTENTS
Volume 32

Number 1 — March 31, 1972

Fossil palm materials from the Tertiary Dipping Vat For-
mation of central Utah. William D. Tidwell, David A.
Medlyn, and Gregory F. Thayn 1

A new subspecies of Crotalus lepidus from western Mexico.
Wilmer W. Tanner, James R. Dixon, and Herbert S.
Harris, Jr .- 16

Two new subspecies of Crotaphytus (Sauria: Iguanidae).

Nathan M. Smith and Wilmer W. Tanner 25

A new species of Stenamma (Hymenoptera: Formicidae )

from Utah. Robert E. Gregg 35

New synonymy in the bark beetle tribe Cryphalini (Coleop-

tera: Scolytidae). Stephen L. Wood 40

The contribution of the 1822 works of Jarocki and Fleming
to herpetological nomenclature. Robert C. Feuer and
Hobart M. Smith .- 55

Number 2 — June 30, 1972

Prey stage distribution, a factor affecting the numerical
response of Typhlodromus occidentalis to Tetranychus
mcdanieli and I'etranychus pacificus. B. A. Croft 61

A revision of the Psoralca lanceolata complex. C. A. Toft

and S. L. Welsh 7C^

Observations on the nesting biology of the long-billed

curlew. Dennis M. Forsythe 88

Vegetation zones around a small ])ond in the White Moun-
tains of Arizona. B. Ira Judd 91

A new species of Gymnodomaeus from Colorado (Acarina:

Cryptostigmata, (jymnodomaeidae). Tyler A. Woolley ... 97

The Sonoran subspecies of the lizard Ctenosaura hcmi-

lopha. HobarJ M. Smith 104

Dead great blue heron found at 11,000 foot Utah elevation.

Gary L. Plickman 112

Five new varieties of Erigonurn (Polygonaceae). James L.

Reveal 113

Plants new to the Utah flora. Larry C. Higgins 117

Notes on Nevada .solpugids. Dorald M. Allred 120

Number 3 — September 30, 1972

vi;

On the rodent-infesting Anoplura of Panama. Phyllis T.

.lohnson 121

Notes concerning Mexican Saldidae, including the descrip-
tion of two species (Hemiptera). John T. Polhemus 137

Scorpions of Utah. John D. Johnson and Dorald M. Allred 154

Relict stands of pinyon hybrids in northern Utah. Ronald

M. Lanner and Earl R. Hutchinson 171

Studies in nearctic desert Decticidae (Orthoptera: Tetti-

gonioidea). Part I. Ernest R. Tinkham 176

Number 4 — December 31, 1972

Aquatic Phycomycetes of Lily Lake, Utah. Hugh M.

Rooney and Kent H. McKnight 181

New synonymy in American bark beetles (Scolytidae:

Coleoptera), Part IL Stephen L. Wood 190

Flights of the western thatching ant, Formica obscuripes
Forel, in Nevada (Hymenoptera: Formicidae). Wil-
liam H. Clark and Peter L. Comanor 202

A new species of soil mite from western Colorado (Acari:
Cryptostigmata, Pelopidae). Harold G. Higgins and
Tyler A. Woolley 208

A checklist of the macroinvertebrates of the Provo River,
Utah. Parley V. Winger, Edward J. Peters, Michael
J. Donahoo, James R. Barnes, and David A. White 211

A record of an indigo bunting and a wintering Say's

phoebe for northern Utah. Gary L. Worthen 220

Comments on two names in an early Utah flora. James A.

Reveal 221

Nearctic desert sand dune Orthoptera. Part XIV: A new

Eremopedes (Decticidae). Ernest R. Tinkham 223

Seasonal food habits of barn owls in LTtah. Dwight G.

Smith, Charles R. Wilson, and Herbert H. Frost 229

Germination of Pinus aristata Engelm. William H, Reid 235

Notice of a new report entitled "Systematics collections: A

national plan." J. C. Dickinson 238

Yifb.K

The

MUS. COMP. ZOO?

LIBRARY Volume 32, No. 1

March 31, 1972

MAY I 9 1972

liARVARD

' '•\U\/>--r-,c;:-rV

Great Basin

nmmim

pubushed by
Brigham Young University

GREAT BASIN NATURALIST

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

Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W.
Tanner, Zoology; Joseph R. Murdock, Botany; Vernon J.
Tipton, Zoology; Ferron L. Andersen, Zoology

Ex officio Editorial Board Members:

A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences

Ernest L. Olson, Director, University Press, University
Editor

The Great Basin Naturalist was founded in 1939 by Vasco M.
Tanner. It has been continuously published from one to four times
a 3^ear since then by Brigham Young University, Provo, Utah. In
general, only original, previously unpublished manuscripts pertain-
ing to tiie biological natural history of the Great Basin and western
North America will be accepted. Manuscripts are subject to the ap-
proval of the editor.

Subscription: The annual subscription is $5.00 (outside the
United States $5.50). The price for smgle numbers is $2.00 each.
All matters pertaining to the purchase of subscriptions and back
numbers should be directed to Brigham Young University Press,
Publication Sales, 205 UPB, Provo, Utah 84601. Libraries or other
organizations interested in obtainiag this journal through a contin-
uing exchange of scholarly publications should contact the Brigham
Young University Exchange Librarian, The Library, Provo, Utah
84601.

Manuscripts: All manuscripts and other copy for the Great
Basin Naturalist should be addressed to the editor. Contributors
should consult the instructions printed on the back cover of a recent
number.

The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

Volume 32 March 31, 1972

No. 1

FOSSIL PALM MATERIALS FROM THE TERTIARY
DIPPING VAT FORMATION OF CENTRAL UTAH

William D. Tidwell,! David A. Medlyn.i and Gregory F. Thayni

hUU^^'J^"''!'^ 7^^ ^hi-opalmoxylon (petrified palm roots) species from the
aI' k 1 ^^i"^°"^' Utah were originally reported as from the Jurassic

Arap en Shale. However, the Utah State Geologic Map upon which this was
largely based is erroneous, and the strata contaming the palm roots are in the
lertiary Dipping Vat Formation, rather than in the Jurassic Arapien Shale
^rom the palm roots previously reported, three new species have been identified:
Khizopalmoxylon behummi, R. blackii, and R. scottii.

Specimens of the new species, Palmoiylon gustavesonii, were collected from
the Dipping Vat Formation east and south of Redmond. This formation uncon-
formably overlies the Arapien Shale in the vicinity of the previously reported

ShalfV'.nn ?K ' "^^^ P'T^"] '''"^- '^^ P^^"^ ^^^^ '•^P^'-'^d from the Arapien
^hale cannot be conclusively demonstrated to have come from the Dipping Vat

Srinn^t F°'^''''r' '^^ P/r'""'^ °^ ^"°!^"'' 'P^^i^^ °f Petrified palm in the
Shafe l^ormation is sufficient to cast doubt on their existence in the Arapien

The species of fossil palm material {Rhizopalmoxylon and
Haimoxyion) discussed in this paper are from the Dipping Vat For-
mation of Tertiary age near Redmond, Utah.

D 7^^ Rhizopalmoxylon species from the Redmond Hills west of
Redmond (Figs. 1, 2b) were originally reported as from the Jurassic
Arapien Shale (Tidwell et al, 1970b, 1971a). l^his was based large-
ly upon these strata being mapped as Arapien Shale (Utah State
(jeologic Map). This mapping has since been demonstrated to be
erroneous (Scott et al., in press), and the strata containing the palm
roots are, without question, in the Dipping Vat Formation.

Specimens of the new Palmoxylon species (Palmoxylon gustave-
sonii) were collected from the Dipping Vat Formation approximately
^ur miles east and south of Redmond in central Utah (Fig 1)
Ihese specimens consist of axes which were embedded within a
resistant tuffaceous sandstone, as well as eroded, fragmental surface
samples.

The specimens in the Dipping Vat Formation, which uncon-
tormably overlies the Arapien Shale Formation in the vicinity of

^Q«Q''^o7^n ?nlnu?''^'"' '^'^'''^ previously reported (Tidwell et al.,
I^t)9^j970a^7()b), were shown to us by Mr. Jay Gustaveson of

•Department of Botany and Range Science. Brigham Young University, Prove, Utah 84601.

\SIN NATURALIST

Vol. 32, No. 1

V

t-'W

Un^'

^■~-\

''

(\ r-^

VSalt Lore City

— 1

UTAH

Redmond Area

Palmoxylon

Rhizopalmoxylon

Fig. 1. Index map of sites of palm material in tlie Tertiary Dipping Vat
Formation.

Salt Lake City some time after articles on the Jurassic palms were
published. These papers would have been seriotisly moditied had
the presence of petrified palms in the Tertiary beds been known.
The surrounding hills had been seriously searched previously and no
fossil palm materials were encountered at that tmie, geologically or
topographically above the Arapien Shale. The reason for not pre-
viously encountering fossil palm materials appears to be the result
of (1 ) not searching far enough into the valley formed in the Dipping
Vat Formation to encounter the palm localities from which the
amateur collectors had previously collected palm specimens and
(2) the collectors removal of float in this area, resulting in a paucity
of specimens.

Although Palmoxylon simpcri and P. pristina, described Jroni the
Arapien Shale, are distinct species, there is a possibility that they

March 1972 tidwell, et al: fossil palms 3

may also have come from the younger Dipping Vat Formation. It is
not certain as to how they arrived in the Arapien Shale. But, to
have them eroded, roll approximately a mile over a near vertical
cliff of about 100 feet, and remain intact as axes three to four feet
in length does not seem possible.

.^^^'^^.^'^^^^^P^^— The Dipping Vat Formation (McGookey,
1960) IS one of a series of formations deposited in association with a
large Late Eocene Lake in central Utah. Clastic fragments of the
formation were derived largely from volcanic activity to the south
or southwest of the central Utah locality. The formation here con-
sists of coarse-grained, gray to white, sometimes gray-blue, locally
hydrothermally altered to red, evenly bedded tuffaceous sandstones
which contain varying amounts of pyroclastic fragments. Minor
white clay and silty limestone are also interbedded.

The Dipping Vat Formation is overlain by the Bullion Canyon
Volcamcs at the type section and is underlain by the Bald Knoll
Formation. In the Redmond Hills, the Dipping Vat Formation is
hydrothermally altered to reddish gray. These sediments are strik-
ingly similar in appearance to some horizons in the Arapien Shale.

Rhizopalmoxylon Felix, 1883

Rhizopalmoxylon is the form genus for petrified roots referable to
the family Palmae. Diagnostic criteria for determination of species
and genera have been variously treated. Tomlinson (1961), Drabble
(1904), and Cormack (1896) considered them not to be definite
enough for differentiation. Mahabale and Udwadia (1959) ". . . sug-
gested that since many of these characteristics [concerning palm
roots] are of a stable nature, they can be used for the analysis of
the artificial genus of palm roots, Rhizopalmoxylon:' Palm roots'
possess some peculiar features such as a so-called polystele due to
dissolution of the stelar system into strands of various shapes, changes
in their anatomy due to varied habitats, etc. Some of these features
are distinctive for certain species and genera which will help in
identifying them on the basis of their anatomy (Mahabale and
Udwadia, 1959).

Roots of living palms arise generally from the basal portion of a
tree trunk where they form a dense, surrounding network. Position
on the plant and function of roots of palms can be distinguished
Ihere are large-sized, aerial roots which function as prop or stilt
roots. Others are small and grow downward, serving as absorbina
roots. Cluster-forming aerial roots arise a little above ground, grow
downwards at various angles, and form a thick mat around the tree
trunk. In some palms the secondary roots grow vertically upward
and form pneumatophores. Each of these vary as to their anatomy
/i^i have distinctive criteria useful in recognizing species of palms
(Mahabale and Udwadia, 1959).

The anatomy of Rhizopalmoxylon has been discussed in conjunc-
V^^.^o , P^^'^oxylon species to which they are attached (Stenzel,
1904; Sahni, 1938; Gothan, 1942; Stockmans and WiUiere 1943-

4 GREAT BASIN NATURALIST Vol. 32. No. 1

Shukla, 1946; Ogura, 1952; Lakhananpal, 1955; Rao and Menon,
1965; Menon, 1968; Tidwell et al., 1971b).

The well-preserved palm roots constituting this report are im-
bedded in growth position at several horizons within sandstone strata
of the Dipping Vat Formation. In some areas, they form an intricate
pattern of large and small roots similar to the network developed in
living arborescent monocots.

Rhizopalmoxylon bchuninii. n. sp.
Figfures 2a. 3

Description. — These consist of silicified roots, 5-10 mm in
diameter, interwoven among the sediments. The epidermal layer,
epiblema of Mahabale and Udwadia (1959), or limiting layer of
Drabble (1904) and the hypodermal layers are generally not
preserved.

The three parts of the cortex lie beneath the slight remains of
the hypodermis. The outer cortex is about 0.05 mm in thickness
and consists of one to four layers of thin-walled, blocky, parenchy-
matous cells.

The lacunar, middle cortex contains slender, radiating air spaces
which are separated by irregular, thin radially extended dia-
phragms. The short, thick-walled parenchyma cells forming the
diaphragms are radially aligned, usually one to three cells in thick-
ness. This zone is approximately 2 mm wide.

Fourteen layers of blocky or oblong-shaped, compact, thin-walled
parenchyma cells compose the inner cortex, which has a total thick-
ness of about 0.05 mm. Beautifully preserved brachysclereids are
sometimes present in this tissue.

The single cell layer of the endodermis is poorly preserved and
resembles the single-layered pericycle. Cas])arian strips occur, where
visible, only on radial walls. Some endodermal cells contain a dark
substance.

There are upward of 26 exarch xylem strands present in the
stele. Each strand contains two or three metaxylem vessels, wall to
wall, or alternating in two to three concentric rings. Areas of phloem
occur between the protoxylem elements near the pericycle. The
metaxylem elements are surrounded by compact, thin-walled paren-
chyma cells, in which an occasional fiber can be observed.

The pith is formed of compact, thin- walled parenchyma cells
and rarely contains ceils with secondary walls. The center portion
of the pith breaks down, leaving an irregularly sha{)ed hollow center
surrounded by parenchyma.

This species is named for the late Mr. Homer Behunin of Red-
mond, Utah, who was the original discoverer of these fossil palm
localities.

Repository. — Brigham Young University, 920 (Holotype)

Locality.— Redmond Hills, Utah

March 1972 tidwell, et al: fossil palms

Fig. 2. A. Cross section of Rhizopalmoxylon behuninii. B. A Rhizopal-
moxylon specimen in place.

GREAT BASIN NATURALIST

Vol. 32, No. 1

Fig. 3. Cross section of Rhizopalmaxylon behuninii The blackened portions
indicate areas of poor preservation. (Cells are slightly enlarged.)

March 1972 tidwell, et al: fossil palms 7

Horizon.— Dipping Vat Formation
Age. — Late Eocene

Rhizopalmoxylon blackii, n. sp.
Figure 4

Description. — Silicified roots are approximately 10 mm in
diameter.

„ \t- ./. n-F l^^ -L

Fig. 4. Cross section of Rhizopalmoxylon blackii. Note the thick-walled
conjunctive tissue. (Cells are slightly enlarged.)

8 GREAT BASIN NATURALIST Vol. 32, No. 1

Epidermis and hypodermis are not preserved and the outer cortex
is nearly obliterated. The outer and middle cortex is 2.5 mm wide
with the composition of the outer being compact, thin-walled paren-
chyma. The middle cortex is not well preserved. Where present it
consists of radiating diaphragms separating air cavities. The dia-
phragms are made of thin-walled, isodiametric parenchjnia cells.

The inner cortex is 0.05 mm in width, and, although poorly pre-
served, relatively thick-walled, isodiametric cells are observable.
This relatively uniform cortical tissue is approximately eleven cells
in thickness.

The endodermis is one cell thick with some darkened areas on its
radial cell walls which may be the remains of casparian strips. The
pericycle is not preserved but appears to have been about one cell
in thickness-

The stele is well preserved. Patches of phloem with approxi-
mately eight sieve tubes per area occur between strands of pro-
toxylem. Thirty protoxylem points are observable with twenty
metaxylem elements in each of two concentric rings. The pro-
toxylem and first formed metaxylem elements are wall to wall with
later developing metaxylem elements alternating.

Compact, small, thick-walled cells make up the conjunctive tissue
with some thin-walled parenchyma ensheathing some of the vessel
elements. Pith contains large, thin-walled cells. A lacuna occurs in
the pith, but this appears to be due to preservation rather than a
breakdown of parenchyma cells.

Rhyzopalmoxylon blackii was named for Mr. Robert Black of
Redmond, Utah, for his contribution to this study.

Repository. — Brigham Young University, 921 (Holotype)

Locality. — Redmond Hills, Utah

Horizon. — Dipping Vat Formation

Age. — Late Eocene

Rhizopalmoxylon scottii. n. sp.
Figure 5

Description. — Petrified roots, 7-9 mm in diameter, embedded
within sandstone layers.

The epidermis and hyj)odermis are not preserved. The outer
cortex, comprised of layers of compactly arranged j)arenchyma cells,
is 0.05 mm wide. The 2 nmi middle cortex in some sjiecimens is
composed of compact, thick-walled, round to hexagonal, isodiametric
parenchyma cells. In other specimens, the middle cortex consists
of diaphragms of these isodiametric cells separated by air cavities.
Some isolated fibers are present in the middle cortex.

The thin-walled, isodiametric-celled, inner cortex is four cells and
0.05 mm in thickness. The cells become smaller toward the endo-
dermis. The endodermis and peri( ycle are uniseriate, and the peri-
cycle is poorly preserved. Casj)arian strips may be observed on the
lateral and inner walls of the endodermal cells.

March 1972

TIDWELL, ET AL: FOSSIL PALMS

iP^^X-'^

'^^

Wr

^J^^-^^

^§

fifi

'I'i^'"

^'""

''^^^L

^V- ■

-im

^^^^mi

¥

Fig. 5. Cross section of Rhizopalmoxylon scottii. Note the compact middle
cortex, the smaller stele size, and the C-shaped casparian strips on the endodermis.
(Cells slightly enlarged.)

In the stele, which is 2 mm in diameter, patches of phloem occur
between the protoxylem strands. There are fourteen exarch xylem
strands arranged in two concentric rings surrounded by thin-walled
conjunctive tissue. Xylem elements are wall to wall or alternating.
Parenchyma, ensheathing the vessel elements, contains some type of
dark substance.

The thin-walled parenchyma cells of the pith are larger than the
cells composing the conjunctive tissue. A lacuna occurs but appears
to be due to preservation.

Rhizopalmoxylon scottii was named for Dr. Richard Scott of the
U.S. Geological Survey for his interest and contribution to this study.

10 GREAT BASIN NATURALIST Vol. 32, No. 1

Repository — Brigham Young University, 922 (Holotype)

Locality. — Redmond Hills, Utah

Horizon. — Dipping Vat Formation

Age. — Late Eocene

Discussion of the Rhizopalmoxylon species. — The palm
roots of this study are found in growth position within the sediments
of the Dipping Vat Formation. The roots appear to be subterranean
and probably do not represent aerial roots or pneumataphores, which
would be more inclined to be destroyed because of exposure. They
appear to be absorbing roots, all serving the same function, and
possessing a different anatomical structure than the other root types.

In comparing the species discussed here with other described
forms, they were found to be dissimilar in their stelar configuration.
Rhizopalmoxylon hehuninii, R. hlackii, and R. scottii have two or
three concentric rings of metaxylem elements, whereas those to
which they were compared had only one.

The stelar configuration and conjunctive tissue of Rhizopalmoxy-
lon behuninii and R. scottii are similar, with the exception of the
much smaller stele size of the latter. The overall root size is the
same in both species, and therefore R. scottii does not appear to rep-
resent a small secondary root of R. behuninii- Smaller secondary
roots possess a compact middle cortex as in R. scottii; however, the
middle cortex of some specimens of R. scottii also contain air cavities,
indicating that they are distinctly different from R. behuninii.

These sjiecies differ from Rhizopalmoxylon blackii in their stelar
pattern and conjunctive tissue. Mahabale and Udwadia (1959, p.
93) states, "The different patterns of the stele in palm roots with its
conjunctive parenchyma form a very characteristic feature of each
species. . . ."

The endodermal cells of R. scottii have the typical C-shaped
casparian strips of Russow, whereas those on R. blackii and R.
behuninii appear to be only on the radial walls.

The conjunctive tissue of Rhizopalmoxylon behuninii and R.
scottii is composed of thin-walled parenchyma, contrasting with the
thick-walled conjunctive tissue of R. blackii. The conjunctive tissue
of R. behuninii varies little from its pith, while the pith of R. scottii
is composed of thin-walled cells which are larger than the cells of its
conjunctive tissue. The j)ith of R. blackii is composed of thin- walled
cells which (hffer from its thick-walled, smaller-celled conjunctive
tissue. R. hlackii also has more metaxylem elements, and the inner
cortex consists of isodiametric cells, rather than the elongated hexa-
gonal cells of R. behuninii.

The Palmoxylon si)ecies to which these roots may be related is
uncertain at the present time. Although all Palmoxylon specimens
collected by the authors from the Dipping Vat Formation are assign-
able to P. gustavesonii, other species of this genus may have also
been present. Eventually, the basal portion of P. gustavesonii and
other possible species may be uncovered and collected, and the af-
finities of these Rhizopalmoxylon species may then become known.

March 1972 tidwell, et al: fossil palms 11

Roots of the living palm genus Latania are similar to these
Rhizopalmoxylon species in stelar configuration. They vary, how-
ever, by the roots of Latania having lysigenous air cavities in the
middle cortex, whereas the middle cortex of the Rhizopalmoxylon
species described in this paper appears to be formed schizogenously.
This is similar to those found in some palms growing in swampy
situations, such as Raphia and Nypa (Tomlinson, 1961).

Palmoxylon Schenk, 1882

Palmoxylon gustavesonii, n. sp.

Figures 6a, 6c

Description. — All that is anatomically preserved of these
specimens are the central and subdermal zones of the petrified axes
which vary from 25 to 30 cm in diameter. The central zone is char-
acterized by irregularly oriented vascular bundles. These bundles
are 750/i at their widest, and vary from 800 to 950/t in height. There
are approximately 67 bundles per cm-. The f/v ratio is commonly
1:1 but can differ slightly (1.5:1, 1:1.2). The bundle shape is
blocky, oblong to infrequently ovate. The bundle cap is reniform
in shape with a shallow median sinus. Auricular sinuses are marked-
ly absent, with auricular lobes being rounded.

Vascularization consists of two metaxylem elements per bundle,
occasionally three. The elements vary from 155 ta 200/1 in diameter.
In longitudinal section, the vessels are 500 to 600/t in length with
simple perforation plates and scalariform thickenings on the wall.
Protoxylem elements were not observed in the central region of our
specimens. The phloem is usually not preserved, but in some bundles
the outline of elements can be determined, and they range from 40
to 65/1 in diameter. Both the vascular portion and the fibrous bun-
dle cap are encased by tabular parenchyma. The ground tissue con-
sists of thin-walled, compact, isodiametric cells, lacking lacunae.
Numerous fibrous bundles 150 to 259;u. in diameter are present, but
lack stegmata.

The bundles of the subdermal zone, although poorly preserved,
tend to be more or less regularly aligned near the dermal zone, and
irregularly oriented near the central zone. The bundles tend to be
larger in the subdermal zone (1 mm x 1.25 mm). There are ap-
proximately 84 bundles per cm- with a f/v ratio of 1:2, rarely 1:1.

The vascularization and size of the metaxylem elements are simi-
lar to those of the central zone.

This species was named in honor of Mr. Jay Gustaveson of Salt
Lake City, Utah, for his aid in this study.

Repository. — Brigham Young University, 923 (Holotype)

Locality. — Four miles east and south of Redmond, Utah

Horizon. — Dipping Vat Formation

Age. — Late Eocene

Discussion and conclusions. — Some collected or observed
axes of Palmoxylon gustavesonii were eight to nine feet long and

12

GREAT BASIN NATURALIST

Vol. 32, No. 1

B

Pig. 6. A. Longitudinal view of simple perforation plate on vessel of Pal-
moxylon gustavesoni. B. Longitudinal section of scalarifonn perforation plate on
vessel of Palmoxylon simperi. C. Cross section of bundles of Palmoxylon gus-
tavesom (enlarged) illustrating typical oblong bundle shape.

March 1972 tidwell, et al: fossil palms 13

were embedded within a resistant tuffaceous sandstone. Samples
were studied from various points along these axes. There does not
appear to be any appreciable anatomical differences between them.

Palmoxylon gustavesonii and P. simper i Tidwell are not similar,
but, because their collection sites are within the same general area,
a detailed comparison of these two species is necessary.

Palmoxylon gustavesonii is distinct from P. simperi in that the
vascular bundles of P. simperi tend to be larger (1.25 mm to 2 mm),
less numerous, and more widely spaced with 53 bundles per cm-.
The bundle caps of both species are reniform, although P. simperi
has prominent auricular sinuses and a rounded to somewhat angular
median sinus. The f/v ratio in the central zone of P. simperi is
2.5:1 compared to 1:1 in P. gustavesonii. Vessels of P. simperi have
both simple and scalariform perforation plates (Fig. 6b) and its
vessels tend to be longer (about 1 mm in length) in longitudinal
section.

Palmoxylon pristina Tidwell is also found in the same general
area as both P. simperi and P. gustavesonii^ but, due to the poorly
preserved specimens of P. pristina., a more detailed comparison is not
possible. The bundles of P. pristina are smaller and more rounded
with the bundle height being the same as the width. The bundles of
P. gustavesonii are mostly oblong, whereas those of P. pristina are
pear-shaped to triangular. The f/v ratio of P. pristina is 1.7:1 to
2.7: 1 as compared to 1 : 1 to 1.5: 1 for P. gustavesonii.

Based on the comparison of the central zones, Palmoxylon gus-
tavesonii, P. pristina, and P. simperi are distinct species. Although
the statement was originally made that P. pristina and P. simperi
may have been from different parts of the same plant and may rep-
resent the same biological species (Tidwell et al., 1970a), they are
distinct and appear to represent separate biological as well as form
species.

The bundle shape in the central zone of Palmoxylon pondicher-
riense Sahni is similar to P. gustavesonii, but it differs mainly in the
occurrence of stegmata on the fibrous bundles and the presence of
large lacunae in the ground tissue of the central zone. The vascular
bundles of P. pondicherriense decrease in size from the dermal to
the central zone, a character which is the reverse of most Palmoxy-
lons.

Palmoxylon edwardsi Sahni looks very similar, but only the
dermal and subdermal zones were present and described. This makes
a comparison with P. gustavesonii difficult because of the poorness of
preservation in the subdermal zone and the lack of the dermal zone
in the latter. Some of the observable differences are the lack of tabu-
lar parenchyma, the occurrence of small air spaces in the ground
tissue, and the tendency for some of the bundles in the subdermal
zone of P. edwardsi to show a radiating pattern.

Because of the interest shown for the possibility or impossibility
of palms occurring in the Jurassic, this paper on petrified palm axes
and roots from the Tertiary Dipping Vat Formation, which in some
areas overlies the Jurassic Arapien Shale, has an added significance.

14 GREAT BASIN NATURALIST Vol. 32, No. 1

The responsibility of any errors in the prior pubhcations relating to
Jurassic palms (Tidwell et al, 1969, 1970a, 1970b, 1971a) remains
with the senior author. Although these publications appear to be due
to hasty conclusions, this was not the case. Much time and effort
was spent in trying to arrive at the truth. Several expeditions with
as many as twenty students were made into the hills surrounding
the collecting localities of Palmoxylon simperi and P. pristina look-
ing for any petrified material above the Arapien Shale and, as pre-
viously stated, none was found. Many persons were questioned
concerning the possibility of fossil palm materials above the Arapien
Shale, and in most cases their answers were later proven to be either
false or evasive for whatever reason each one had. However, no one
admitted to the petrified materials being present in the younger beds
until Mr. Gustaveson came forward and conducted us to their loca-
tion after our earlier papers were published.

The palm roots are definitely from the Dipping Vat Formation,
whereas the palm axes previously reported from the Arapien Shale
cannot be conclusively demonstrated, at the present time, to have
come from this younger formation. However, the possibility of their
derivation from the younger strata, as suggested by petrified palm
axes from the Dipping Vat, is sufficient to cast doubt on their exis-
tence in the Arapien Shale.

Acknowledgments

The authors wish to thank the following individuals for their
time and efforts during the study. Naomi Hebbert for aid in the il-
lustrations, A. Daniel Simper of the University of California at
Davis, Dr. J. Keith Rigby and Dr. James L. Baer of the Department
of Geology at Brigham Young University, and Dr. S. R. Rushforth of
the Botany Department, Brigham Young University, as well as those
previously cited.

LiTER.\TURE Cited

CoRMACK, B. G. 1896. On polystelic roots of certain palms. Trans. Linn. Soc.

Lond. Bot. ser 2. 5:275-286.
Drabble, E. 1904. On the anatomj' of the roots of palms. Trans. Linn. Soc.

Lond. Bot., ser 2. 6:427-490.
GoTHAN, E. 1942. Uber palmenwurzelholzer aus der Braunkolile von Bohlen

(Sachsen). Zeit. F. Glach.
Lakhananpai., R. 1955. Palmoxylon surangei, a new species of petrified palms

from the Deccan Intertrappean series. Palaeobot. 4:15-21.
Mahabale, T. S., and N. N. Udwadia. 1959. Studies on palms: Part IV —

Anatomy of palm roots. Proc. Natn. Inst. Sci. India. 20B: 73-104.
McGooKEY. D. P. 1960. Early Tertiary stratigraphy of part of Central Utah.

Bull. American Assoc. Petroleum Geol. 44:589-615.
Menon, V. 1968. On a new petrified palm wood from Mohgaon Kalan area.

Palaeobot. 16:197-205.
Ogura, Y. 1952. A fossil palm in Kenroku Park in Kanazawa. Trans. Proc.

Palaeont. Soc. Japan, n. s.. 8:225-230.
Rao, a. R., and V. K. Menon. 1965. A new species of petrified palm stem

from the Deccan Intertrappen series. Palaeobot. 14:256-263.
Sahni, B. 1938. Recent advances in Indian Palaeobotany. Proc. 25tli Indian

Sci. Congress, Calcutta. 11:133-176.

March 1972 tidwell. et al: fossil palms 15

Scott, R., P. Williams, E. Barghoorn, L. Craig, L. Hickey, and H. Mac-

GiNiTiE. Pre-Cretaceous angiosperms from Utah: evidence for Tertiary age

of the Pahn woods and roots, (in press)
Shukla, V. B. 1946. Palmoxylon sclerodermum Sahni from the Eocene Beds

of Nawargaon, Wardha District, C. P., J. Indian Bot. Soc. 15:105-116.
Stenzel, K. G. 1904. Fossile Palnienholzer. Beitr. Palaeont. Geol. Ost. — Ung.

16:107-287.
Stockmans, F., and Y. Williere. 1943. Palmoxylon paniselianse de la Bel-

gique. Mem. Mus. Hist. Nat. Belg. 31:1-35.
TiDWELL, W. D.. S. R. Rushforth, and J. L. Reveal. 1969. Petrified palmwood

from the Arapien Shale. Utah. (abst. )Geol. Soc. Am. Special paper.
TiDWELL, W. D., S. R. Rushforth, and A. D. Simper. 1970b. Pre-Cretaceous

flowering plants: further evidence from Utah. Science 170:547-548.
. 1971a. Rhizopalmoxylon from the Arapien Shale (Jurassic) of Utah.

(abst.) Am. J. Bot. 58:473.
TiDWELL, W. D., A. D. Simper, and D. A. Medlyn. 1971b. A Palmoxylon

from the Green River Formation (Eocene) of Eden Valley, Wyoming.

Botanique 2:93-102.
Tomlinson, P. B. 1961. Anatomy of Monocotyledons. II — Palmae. Oxford.

A NEW SUBSPECIES OF CROTALUS LEPIDUS
FROM WESTERN MEXICO

Wilmer W. Tanner,i James R. Dixon^ and Herbert S. Harris, Jr.^

Abstr.\ct.— An examination of additional specimens of Crotalus lepidus
from western Durango and the adjacent parts of Smaloa and Nayarit have
demonstrated the validity of Klauber's (1956) suggestion that a new subspecies
may occur. The new subspecies (C. /. maculosus) is described and compared to
other subsf>ecies.

Klauber (1956:37) recognized the uniqueness of three specimens
of Crotalus lepidus from areas near the junction of the Mexican
States of Durango, Sinaloa, and Nayarit. Although lepidus from
that area were considered to belong to the subspecies klauberi, or
were at least in the designated range for that subspecies, Klauber
keyed such specimens to the subspecies lepidus or morulus and sug-
gested that an adequate series from this rough and inaccessible area
might justify the recognition of a new subspecies.

The area northwest, west, and southwest of El Salto, Durango, is
some of the most rugged terrain in Mexico. In this area with its
deep canyons, high ranges, plateaus, and its diverse climate ranging
from wet to dry, there has been produced a variable and challenging
environment which has seemingly been responsible for the develop-
ment through adaptation of a number of subspecies. Perhaps of equal
importance in this study is the fact that within this general area at
the higher elevations are found some of the more primitive members
of the genus Crotalus. It is not, therefore, surprising to find a popu-
lation of C. lepidus which has, in its adaptation to this area, devel-
oped a series of unique characteristics.

We now have available a series of 19 specimens from these
states and consider them to be sufficiently distinct to warrant a
description of a new subspecies. Because of the distinctive char-
acter of the series of small dorsal spots we choose to name it

Crotalus lepidus r?iaculosus, subsp. nov.

Typk.— BYII No. 33328. 15 miles (24 km) west of La Ciudad,
near Highway 40, Durango, Mexico. An adult male collected on
31 July 1970 by Richard T. Basey.

Pak.mypes.— BYU 40188, 1 km W Los Bancos, HSH 71-24 and
HSH 71-133, near La Ciudad. MVZ 59310, 16 miles (22 km) SW
La Ciudad ajid UCM 46011, 11 miles W La Ciudad, Durango;
TCWC 13 358, 2 (lavs bv nude E from 1 luajicori, Navarit; BYU
40189, Puerto El Ala/aiu BYU 40190-91, 3 miles E Puerto El Ala-
zaii, HSH 71-23, 10 miles W Durango-Sinaloa line (Hwy 40), KU
78973, 19.2 km NE Santa Lucia. KU 79232, 5 km SW Palmito,

'Department of ZooloRy, RriRham YoiuiK University, Provo, Utah 84601.

'Department of Wildlife Science, Texas A & M University, College Station. Texas 77843.

TTie Natural History Society of Maryland, Baltimore, Md. 21218.

16

March 1972 tanner, et al: rattlesnake 17

LACM 50960, 4.8 miles E Santa Rita (Hwy 40), TCWC 33354-6
Rancho Carrizo 12.5 miles (20 km) W Palmito, TCWC 33357, 2
miles (3 km) W Palmito and TCWC 33359, near El Maguey (East-
ward from Huajicori), Sinaloa, Mexico.

Diagnosis. — A medium to small subspecies of lepidus differing
from other subspecies in having a dorsal series of small spots often
longer than wide and in a primary series averaging 29-31 spots.
It also differs from other subspecies in that the dorsal spots do not
extend laterally on each side of the body to form cross bars, except
near the tail where the spots may form moderate cross bars (Fig. 1 ).
C. I. maculosus differs from klauheri and lepidus in having an in-
crease in the average number of ventrals and caudals, and from
klauheri in that the ground color is heavily pigmented between the
spots, a character shared with lepidus and morulus. It is further
distinguished from klauheri by the heavy flecking on the ventrals,
reduced number of scales between the supraoccipitals, presence of a
postocular stripe, reduction in number of scales in internasal-
prefrontal area, and by its smaller size.

Fig. 1. Lateral view of the head of the type (BYU 33328) showing loreals,
anterior subocular and postocular stripe.

Crotalus I. maculosus shares with lepidus and morulus the dark
ground color, postocular stripe and with morulus an increase in body
spots, ventrals and caudals and paired occipital blotches. It is most
distinct from morulus in that the upper preocular is divided and the
anterior subocular is usually in contact with the upper labials.

Description of Type. — An adult male, total length (snout to
first rattle) 591 mm, tail (vent to first rattle) 56 mm, head length
26.2, head width 15.5, fang length 4.1, rattle small, lateral width
of proximal rattle 2.5, dorsoventral depth 5.4; ratio of tail into total
length 10.55 percent, head into body length 20.42, head into total
length 22.56, fang into head length 6.39, and proximal rattle into
head length, 4.76.

Scale rows on body 23-23-17, on tail at tenth caudal 12 rows,
fringe scales around first rattle 10; ventrals 169, caudals 30, anal

18 GREAT BASIN NATURALIST Vol. 32, No. 1

complete; supralabials 12-13, infralabials 10-11; preoculars 2-2
elongate, upper one divided horizontally, postoculars 3-3, suboculars
2-2, anterior subocular round, enlarged and in contact with the orbit
and the fourth and fifth supralabials; one row of interoculabials
between postsubocular and supralabials; anterior nasal largest and
in broad contact with first supralabial and intemasal; loreal angular;
three small prefo veals between loreal and first two supralabials; three
small postfoveals along lower margin of pit and above third and
fourth supralabials.

Rostral wider (3.8) than high (3.0), clearly visible from dorsal
view, not deeply wedging between intemasals; two internasals, not
in contact with loreals and nearly twice as wide (2.7) as long (1.7),
no canthal ridge on edge of intemasals from snout to canthal; can-
thai large, approximately round and in contact with internasal,
nasal, loreal, upper preocular, supraocular, and three intercanthals;
four small scales (two rows) between canthals and posterior to in-
temasals; supraoculars large and oblong, three rows of small scales
between supraoculars near their middle, head scales posterior to
interocular area smaller than those on anterior dorsal surface.

First pair of infralabials in broad contact on the midventral line,
completely surrounding triangular mental and wedging posteriorly
between genials; genials large, elongate, and in broad contact pos-
teriorly, in contact with first three infralabials anteriorly, and wedg-
ing between gulars posteriorly; five gulars between genials and
first ventral.

Dorsal pattern a series of 31 dark brown spots extending from
head to tail; first spot (post temporal) divided and last four extended
laterally to form broken or continuous transverse bars; spots often
longer than wide and margined with a fine edging of light yellowish
or golden brown; spots 2-3 scales long and 4-5 scale rows wide, be-
coming smaller posteriorly, until fusion with lateral spots forms
bars. One row of lateral spots which becomes more obvious pos-
teriorly and fuses with dorsal spots; no intermediate spots between
primaries.

Ground color a dark brown, nearly uniform, but with some dark
flecking particularly between the dorsal and lateral spots; venter
heavily marbled with dark markings and flecks; head without marks
on dorsum; a dark stripe from eye to temporal area; labials marbled,
with less light markings on supralabials; snout a uniform dark
brown.

Tail with one complete bar and four lateral spots on each side,
dorsal area posterior to first bar light brown, noticeably lighter than
dorsum of body; proximal rattle rust red.

Variations. — Wo have used freelv the scale counts given by
Gloyd (1940) and Klauber (1952 and 1956) in our analysis of the
characters used in this study. Obviously, we have not deemed it
necessary to recount specimens previously studied. We have, there-
fore, examined in detail only those specimens not previously ex-
amined with the exception of a few specimens from critical areas.

March 1972

TANNER, ET AL: RATTLESNAKE

19

Twenty additional specimens of klauberi have been examined
from Mexico — six from Chihuahua, seven from Durango, one from
Jalisco, one from Nayarit, and five from Zacatecas. Variations
from these fit well into the range of variations as established
by Klauber (1956). In a few specimens there are variations worthy
of note, such as the light color and color pattern of two specimens
from 4 miles (6 km) N of La Campana (MSU 2831) and 3 miles
(5 km) E of Conjos (MSU 3613), Durango. In both, the dorsal bars
are faint or obsolete anteriorly and the ground color is a light cream
or grey. We conclude that these specimens are western representa-
tives of the palmcri population, which seemingly occurs in the desert
basin between eastern Durango and east central Coahuila near the
type locality of Monclova as established by Garman (1887). Speci-
mens from central Chihuahua do not show this faded color indicat-
ing that the palmeri phase occurs only in the central desert region

Fig. 2. Dorsal view of type specimen showing the dorsal spots
(left) and posterior of body and tail (right).

mid-body

20 GREAT BASIN NATURALIST Vol. 32, No. 1

of these Mexican states. There are no unusual variations in the
scale patterns of these specimens.

Taylor (1944) in an attempt to understand variations in this
species applied the name semicornutus to a population of lepidus
occuring in southwestern Chihuahua. We have seen additional
specimens of this population and recognize it as a variant of klauberi
but with the distinct possibility of having retained some of the
lepidus characteristics such as a greater diffusion of spotting between
the bars and with the accompanying reduction of the greenish ground
color so evident in most klauberi.

A specimen (BYU 14244) taken 1.5 km W of Cerocouhui, Chi-
huahua, has the greenish color above and a shade of pink laterally
which becomes more intense on the first two rows of dorsals and
the tips of the ventrals. This pinkish color also occurred on the
supraoculars and across the canthals to the rostral. The supraoculars
are raised and extended giving them a very noticeable hornlike ap-
pearance and producing a concavity to the dorsum of the head.
Both the color pattern and shape of the head plates were described
by Klauber (1956) for populations of klauberi in southern Arizona.
The influence of the Arizona i)opulations apparently extends south-
ward along the western slope of the Sierra Madre Occidental, but we
have found no evidence of this influence occurring in klauberi on
the eastern slope in Chihuahua and northern Durango.

We have seen no specimens between west central Chihuahua
and west central Durango, a distance of approximately 350 km. In
this area of the western slope, the ruggedness of the area does not
change; however, there is a warming and an increase in precipita-
tion from north to south. It is not surprising, therefore, for popula-
tions on the Pacific slopes of these mountains in western Durango,
eastern Sinaloa, and northern Nayarit to show noticeable variations
when comj)ared to lepidus found on the drier eastern slope.

A summary of the comparative variations is listed in Tables 1
and 2.

Klauber (1956) recognized the variations in three specimens of
C. lepidus taken nr^ar the Sinaloa-Durango-Nayarit border but was
not able to differentiate them from other subspecies. We have seen
one of these three specimens (SDSNII 45145) taken ap|)roximately
70 miles (112 km) W of El Salto by Darling. Although this speci-
men was collected in the area of rriaculosus, it is one of a few we
have seen in which the anterior subocular does not contact a sufira-
labial and has laterally extending dorsal spots. In other characters,
it shows an increase in dorsal sj)ots — 28, the dark color on the venter
and between the dorsal spots, and an increase in caudals.

A specimen from Santa Teresa, Nayarit (USNM 46333) also
has many intergrading characteristics. There are only 21 body
spots which form short bars and the ventrals (159) and caudals (21)
are low. The color is dark, quite in contrast to klauberi and the
fringe scales are reduced to 10 around a small rattle (5.5x2.5).
Two s[)ecimens from near Coyotes. Durango. show indications of
intergradation with maculosus. Their location near the continental

March 1972

TANNER, ET AL: RATTLESNAKE

21

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22 GREAT BASIN NATURALIST

Vol. 32, No. 1

divide east of El Salto is in the proximity of a probable area for
intergradation. In these specimens, as in USNM 46333, the spots are
large and fewer as in klauberi, although they do not involve as
many lateral rows as in most klauberi. A suffusion of pigment
between the dorsal spots and on the ventrals also reflects the ap-
proach to maculosus.

We concur with Klauber (1956) that this population appears
to be more closely related to the more eastern subspecies than to the
western subspecies klauberi. We recognize the subspecies lepidus
as the most primitive group in the species and suspect that its earlier
distribution extended south to include not only areas now occupied
by lepidus but also areas in Zacatecas and adjacent Durango, Sina-
loa, and Nayarit, which are now occupied by klauberi.

Present distributions indicate that klauberi has recently expanded
its range by moving southward along the eastern slopes of the Sierra
Madre Occidental. In so doing, it is possible that the range of lepidus
was not only invaded but actually divided, leaving a small popula-
tion on the western flank of the mountains. From this isolated popu-
lation the new subspecies maculosus has been derived, perhaps in
much the same way as morulus on the eastern edge of lepidus.

We consider klauberi to be of a more recent origin than lepidus.
If this is not the case, then we must assume that lepidus moved south
and then west at a much earlier date than klauberi. At any rate,
we believe that it would be very difficult to understand the relation-
ships of maculosus in its present distribution if lepidus did not pre-
cede klauberi in its distribution to the south and then have the more
vigorous klauberi proceed to replace it, at the same time dividing its
range, leaving a small pocket isolated on the Pacific slopes. Speci-
mens seen from southern Chihuahua, central Durango, and Zaca-
tecas show no indication that they have been strongly influenced by
a lepidus gene flow and suggest that the southward thrust of klauberi

Fig. 3. A dorsolateral view of BYU 40188 (paratype) taken 1 km W Los
Bancos, Durango. Mexico, by Mr. and Mrs. Thomas Walker. Sr.. 28 Aug. 1968.

March 1972 tanner, et al: rattlesnake 23

quite thoroughly replaced any previous lepidus stock. Only in the
edges of the distribution do we find noticeable variations suggesting
hybridization with other populations. It should also be noted that
any intergradation occurring is toward the southern limits of distri-
bution and therefore fits well our suggestion that the distribution of
lepidus has in reality been divided by klauberi, thus isolating the
population on the Pacific slopes and producing a biotic situation
which permitted the adaptive development of maculosus.

At present the subspecies maculosus is confined to a rather nar-
row area extending along the high mountain slopes west of the con-
tinental divide in Durango-Sinaloa-Nayarit and perhaps as far south
as adjacent northern Jalisco. Its habitat is one of rocky slopes high
in the mountains and perhaps descending to the moderately low foot-
hills of the Pacific slopes. Such a habitat is located in an area of
greater rainfall than generally occurs in the areas occupied by the
subspecies lepidus and klauheri. We suggest that such climatic con-
ditions have contributed to the darker ground color in maculosus
and perhaps also in morulus.

Materials

Crotalus I. klauheri. BYU 13416, 2 miles W Colonia Juarez, BYU
14247, 15281-2, 5.5 miles NE Colonia Juarez, BYU 14244, 2 miles E
Cerocouhui, BYU 16866, 25 miles SE Creel, Chihuahua; FMNH
1494 and HSH 71-261, Coyotes, USNM 46349, Guanacevi, HSH 71-
231, 70-90 miles SW Torreon (Hwy 40), Durango; USNM 46472,
Bolanos, Jalisco; USNM 46333, Santa Teresa, Nayarit; USNM
46354, Berriozabal, USNM 46470-1, Plateado, HSH '71-25 (alive)
and HSH R-784 Southern Zacatecas. Zacatecas, Mexico.

Acknowledgments

We are most grateful to the many persons and institutions who
have furnished us data and specimens. The senior author is par-
ticularly indebted to Mr. Richard T. Basey of Arcadia, California,
for providing the type specimen and for information concerning
other available specimens of maculosus. We are also indebted to
Mr. Louis Pistoia, Curator of Reptiles, Columbus Zoological Gardens,
Ohio, for the data provided on four live specimens and the loan of
one other. Mr. Allan J. Sloan, San Diego Society of Natural History
(SDSNH), was helpful in providing one of the three specimens
seen by Klauber (1956) and in furnishing other pertinent informa-
tion. Mr. Henry A. Molt also provided us data. The following have
aided us by loaning specimens: Hymen Marx, Field Museum of
Natural History (FMNH), William E. Duellman, Kansas Univer-
sity Museum of Natural History (KU), John Wright, Los Angeles
County Museum (LACM), Rollin Baker, Michigan State University
(MSU), Robert C. Stebbins, Museum of Vertebrate Zoology (MVZ),
T. Paul Maslin, University of Colorado Museum (UCM), James A.
Peters, United States National Museum (Smithsonian) (USNM),
and our own institutions Brigham Young University ( BYU), Natural

24 GREAT BASIN NATURALIST Vol. 32, No. 1

History Society of Maryland (HSH), and Texas A and M Univer-
sity (TCWC) for their continued courtesies. We also appreciate
the data provided by Dr. Robert S. Simmons.

Literature Cited

Garman, S. W. 1887. Reptiles and Batrachians from Texas and Mexico. Bull.

Essex Inst. 19:119-138.
Gloyd, H. K. 1940. The Rattlesnakes, Genera Sistrurus and Crotalus. Chi.

Acad. Sci., Spec. Pub. 4:280.
Klauber, L. M. 1952. Taxonomic studies of the rattlesnakes of mainland

Mexico. Bull. Zool. Soc. San Diego 26:1-143.

. 1956. Rattlesnakes. University California Press. Vol. 1:1-708.

Taylor, E. H. 1944. Two New Species of Crotalid Snakes from Mexico. Univ.

of Kansas Sci. Bull. 30:47-56.

TWO NEW SUBSPECIES OF CROTAPHYTUS
(SAURIA: IGUANIDAE)

Nathan M. Smiths and Wilmer W. Tanner^

Abstract. — Analysis of the Crotaphytus collaris complex reveals at least
eight subspecies in two distinct groupings. The co//arzi-complex containing C. c.
auriceps, C. c. baileyi, C. c. fuscus, and C. c. collaris lies essentially to the east of
the Colorado River. The greatest portion of the western-complex containing C. c.
bicinctores, C. i. vestigium, C. c. dickersonae and C. i. insularis lies to the west of
the Colorado River except through western Arizona and northern Mexico. Two
previously undescribed subspecies, C. c. bicinctores and C. i. vestigium, are de-
scribed and named.

This paper presents a brief portion of an extensive study of the
western collared lizards from the Great Basin and the Baja California
Peninsula. The complete report will soon appear in the Biological
Series of the Brigham Young University Science Bulletin.

The range of the western collared lizard, Crotaphytus collaris
baileyi, includes a vast area of the western United States and Mexico.
One of us (Tanner, in Fitch and Tanner, 1951; Ingram and Tanner,
1971) has suspected for a number of years that the lizards occurring
in this vast area represent a heterogenous population. Stejneger
(1890) described Crotaphytus baileyi as a species; however, as data
became available it was recognized to be a western subspecies of
Crotaphytus collaris Say (1823), the form found east of the Con-
tinental Divide. Two insular forms, Crotaphytus dickersonae Schmidt
(1922) from Tiburon Island, and Crotaphytus insularis Van Den-
burgh and Slevin (1921) from Angel de la Guardia Island, Mexico,
although given species rank, have long been recognized as closely
related to C. c. baileyi. Allen (1933) referred to the collared lizards
from Tiburon Island as C. c. dickersonae. Burt (1928) considered
all collared lizards to be closely related.

Fitch and Tanner (1951) established C. c. auriceps from the
upper Colorado River Basin. Ingram and Tanner (1971) reaffirmed
C. c. auriceps as a subspecies and, upon redefinition of the char-
acteristics of C. c. baileyi, established C. c. fuscus as a subspecies
found in the Chihuahuan Desert south of the range of baileyi. The
type locality for that subspecies being 6.5 miles N and 1.5 miles W
of Chihuahua City, Chihuahua, Mexico. The techniques described
by Ingram and Tanner (1971) were used to compare four subspecies
of the western-complex with four subspecies of the collaris-complex.
This technique permitted both an intercomplex and an intracomplex
comparison.

Relationships

To determine relationships, Ward's Minimum Variance Cluster
Analysis (Wishart, 1968) was used to group lizards in clusters of
highest morphological similarity. Lizards were chosen to represent

'Department of Zoology, Brigham Young University, Provo, Utah 84601 .

25

26 GREAT BASIN NATURALIST Vol. 32, No. 1

the range of the geographic locations available, and 150 lizards were
used as input. A random sample was taken from each area and in-
cluded- Great Basin (N=20); Baja California Peninsula (N = 20)
Angel de la Guardia Island (N = 17); Tiburon Island (N = 13)
Upper Colorado River Basin (N = 20); Central Arizona (N = 20)
Chihuahuan Desert (N=20); and the Great Plains (N = 20). A
contingency table was prepared comparing these groupings with the
eight groupings formed by the cluster analysis (Table 1). The null
hypothesis (the two classifications, one by closest morphological re-
semblance and the other by geographical location, are completely in-
dependent of each other) was tested by a chi-square of 49 degrees of
freedom. The test is significant at the 0.001 level.

HI n

A'^(l-oc,49) = 2 2(0ij-Eii)yEu

i=i ;'=!

A:'(1-oc,49) =485.7
a:- (0.999,49) =85.4

Therefore AT- (1- ex ,49) >X'( 0.999,49) and the null hypothesis is re-
jected, and the morphological relationships of the lizards examined
form essentially the same groups as those proposed in Table 1.-

Table 1. A contingency table testing the independence of Ward's clustering
method and the propos^ groups as follows: A = Great Basin, B = Baja Calif.,
C = Angel Is., D = Tiburon Is., E = Upper Colo., F = Central Ariz., G = Chih.
Desert, H = Great Plains.

PROPOSED GROUPS

Western-Complex Co//a/-j5-Complex

Clusters ABCD E'FGH

1

4

6

0

6

2

0

12

0

7

3

16

0

0

0

4

0

2

17

0

5

0

0

0

0

6

0

0

0

0

7

0

0

0

0

8

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

9

9

2

4

9

5

4

4

2

-)

12

0

1

3

3

11

'The data for the Upper Colorado Rive rBasin and central Arizona does not contain the key
characters which distinguish C. c. auriceps and C. c. Baileyi.

It is noted that none of the western-complex clustered with those
of the collar is-complex; however C. dickersonae which had been
given species rank was not as distinctly clustered. This lends addi-
tional support to the suggestion (Burt, 1928; Allen, 1933) for re-
vision of this group. We propose that it is a subspecies of C. collaris
and should be called Crotaphytus collaris dickersonae. We have col-
lected on Tiburon Island and also on the Sonoran Coast opposite the
island and have found specimens from these two areas to be indis-

'We have done little with the populations occurring on the Great Plains. Our data does, however,
suggest that a careful study may reveal additional collared lizard populations worthy of subspecific
recognition. We have deferred the study of these populations to R, Montanucci.

March 1972 smith, tanner: crotaphytus 27

tinguishable. Schmidt (1922:639) stated this possibility when he
said of C. dickersonae "it is possible that this species will be found to
extend on the Mexican mainland and that it will ultimately be re-
ferred to a subspecific rank under C. collaris. . . ."

The other insular form, C. insularis, remains a very distinct
population closely related to the collared lizards occurring on the
Baja California Peninsula. These two populations are very similar
in pattern and external morphology. The Angel Island form has
the second collar much reduced or absent, and with 39-45 subdigital
lamellae on the fourth toe. Individuals on the Baja California
Peninsula extend their range north to Palm Springs, California, and
form a narrow zone of sympatry with the Great Basin subspecies at
the most northern part of their range. Because of these morphologi-
cal and biological relationships, the population from Angel Island is
designated Crotaphytus insularis insularis and those from Baja Cali-
fornia as Crotaphytus insularis vestigium.

There are several characters which distinguish the western-
complex from the co//«m-complex. In the western-complex, the
second collar does not extend onto the arm, there is a prominent
black groin patch in the males, spots are absent in the dorsum of the
first collar, and there are more scales between the interparietal and
the anterior edge of the first collar. Diagnostic characters within the
western-complex are discussed in the diagnoses of the new subspecies.

Crotaphytus collaris bicinctores, subsp. nov.

Holotype.^ Adult male Brigham Young University no. 23883,
collected at Mercury Pass, Nevada Test Site, Nye Co., Nevada, 14
June 1966, by W. W. Tanner.

Paratypes.— BYU nos. 22191, 23629-30, 30587 (all topotypes)
from the Nevada Test Site, Nye Co., Nevada; BYU 12715, Western
Beaver Co., Utah; BYU 12696, 12701, N Wash, W side Colorado
River, Garfield Co., Utah; BYU 18921, Crossing of the Fathers, Kane
Co., Utah; BYU 21000-02, Black Rock Mountain, Millard Co., Utah;
BYU 32097, 52 miles N Yuma, route 95, Yuma Co., Arizona; CAS
37055, Caliente, Lincoln Co., Nevada; CAS 22705, Dayton, Lyon
Co., Nevada; LACM 63187, Big Rock Creek at Dorr Creek, Los An-
geles Co., Calif.; LACM 63182, Sweetwater Spring, Ord Mts., San
Bernardino Co., Calif.; SDSNH, 1444, Hot Springs, Owyhee Co.,
Idaho; CSCLB 2736, 4 miles N Yermo, San Bernardino Co., Calif.;
CSCLB 2735, Mule Canyon, Calico Mts., San Bernardino Co., Calif.

Description of Type. — Snout-vent length 95 mm, tail length
177 mm, width of head at angle of jaw 20.5 mm, hindleg length
from midline to tip of fourth toe 88 mm, femoral pores 17-19, supra-
labials 16-15, infralabials 14-15, fused interorbitals 0, frontoparietals
3, loreal-lorilabial series 9, postmentals not in contact with infra-
labials, gular scale rows from angle to angle of jaw 61, scales from
rostral to interparietal 16, scales from interparietal to anterior edge
of first collar 32, scales from anterior edge of first collar to posterior
edge of second collar 32, total dorsals 161, total ventrals 195, scales

28

GREAT BASIN NATURALIST

Vol. 32, No. 1

within dorsal separation of first collar 1, scales within dorsal separa-
tion of second collar 0, number of spots within dorsal separation of
first collar 0, subdigital lamellae of second toe of right hind foot 19,
subdigital lamellae of fourth toe 39. subdigital lamellae of fifth toe
18 (Fig. 1 and 2).

Type a preserved specimen in which coloration closely resembles
living organisms. Reticulation of the chin and parietal area re-
sembles freckles with this pattern extending to posterior dorsum of

/,/ .</ -% 'U^

Fig. 1. A dorsal view from left to right of Crotaphytus insularis vestigium,
male BYU 23338, female BYU 23337 and Crotaphytus collaris bicinctores, male
BYU 23883, female BYU 23629.

f^^ f'^nvir ;^^c''rfn""■^^'''??^'./r'"''"' '^^^ ^° ""'^^^ of Crotaphytus collaris bicinctores.
BYU 23337^ male BYU ^2f3 38^ """^ Crotaphytus insularis vestigium, female

March 1972 smith, tanner: crotaphytus 29

head. Body dorsum light brown, never green, venter cream with
black patches in groin and axillary regions. First collar nearly com-
plete dorsally and merging ventrally with black of gular area. Second
collar complete dorsally, not extending onto upper arm. Dorsal body
pattern rather uniformly dotted with light spots and with two diffuse
cross bands. Forelegs, hindlegs, and tail brown, reticulated with
white. Dorsum of tail ridged and tail compressed laterally.

Diagnosis. — The Great Basin population differs from C. c.
baileyi of the collaris-cova\)\ex in many of the 32 characters mea-
sured and particularly in the following which are listed with their
coefficient of difference (Mayr, 1969): dorsal separation of the first
collar (1.63); number of spots in dorsum of first collar (1.45); num-
ber of enlarged median internasals (1.39); and brown dorsal ground
color (1.33). Other characters include second collar not extending
onto the arm, greater number of scales from interparietal eye to
anterior edge of first collar, and more extensive black coloration of
the ventral groin.

The Great Basin form can be distinguished from the Baja Cali-
fornia and Angel de la Guardia Island populations by a more com-
plete first (1.4) and second (1.84) collar dorsally. It is distinguished
from C. c. dickcrsonae by its dorsal brown coloration, back pattern,
and several other characters which considered alone are not signifi-
cant but that give good population separation when computed to-
gether in the clustering program (Table 1).

Range. — Extends throughout the Great Basin region. Apparent-
ly does not occur east of the Colorado River in Utah, but does cross
the Green River and may intergrade with C. c. auriceps in this
region. Yhe Colorado River continues as a barrier south to Oatman,
Mohave Co., Arizona, where we have captured the Great Basin sub-
species. Twenty miles to the east in Kingman, Arizona, C. c. baileyi
is found. This is an area of possible intergradation that needs further
investigation. The Great Basin form continues on the east side of
the Colorado River down to Yuma Co., Arizona. The area of inter-
gradation with C. c. fuscus and C. c. dickersonac is incompletely
known. In southern California, it extends to the San Gorgonio Wild
Area and the Whitewater River Canyon northwest of Palm Springs,
Riverside Co., and apparently crosses the the narrow sandy valley
of the San Gorgonio Pass to reach the eastern foothills of the San
Jacinto Mountains where C. i. vestigium is found (Fig. 3).

Crotaphytus insularis vestigium, subsp. nov.

HoLOTYPE. — Adult male, Brigham Young University no. 23338,
collected in Guadelupe Canyon, Juarez Mountains, Baja California,
15 July 1965, by James W. Heinrichs.

Paratypes.— BYU nos. 23337, 23339, 22298-99 (all topotypes)
Guadalupe Canvon, Juarez Mts., Baja Calif.; BYU 33321, Mt.
Springs, Imperial Co., Calif.; BYU 2422, 2432, 2435, Chino Canyon,
Palm Springs, Riverside Co., Calif.; BYU 2429-30, Snow Canyon,
Palm Springs, Riverside Co., Calif.; SDSNH 1687, 11951, Palm Can-

30

GREAT BASIN NATURALIST Vol. 32, No.

Fig. i. Geographical distribution of the western subspecies of Crotaphytus
collaris: 1, C. c. bicinctores; 2. C. i. vestigium; 3, C. i. insularis; 4, C. c. dicker-
sonae; 5, C. c. auriceps; 6, C. c. bailryi; and 7. C. c. fuscus.

March 1972 smith, tanner: crotaphytus 31

yon, San Diego Co., Calif.; SDSNH 11088, Pinon Mt., San Diego Co.,
Calif.; SDSNH 22327, Sentenac Canyon, San Diego Co., Calif.;
LACM 16997-98, Los Angeles Bay, Baja Calif.; CAS-SU 17048,
Sierra San Pedro Martir, Baja Calif.; CAS-SU 18822, 9 miles W San
Ignacio, Baja Calif.; CSCLB LWR670619-1 and 670619-2, 11 miles
W Santa Rosalia; CUM 45855-6 and 45858, 14.8 miles S Puerto
Citos; CUM 45857, Okies Landing; and CUM 45859-61, S end San
Luis Gonzalga, N Puerta Final, Baja California.

Description of Type.— Snout- vent length 111 mm, tail length
252 mm, width of head at angle of jaw 23.8 mm, hindleg length from
midline to tip of fourth toe 102 mm, femoral pores 18-21, supra-
labials 15-16, infralabials 16-15, fused interorbitals 0, frontoparietals
3, loreal-lorilabial series 10, postmentals in contact with infralabials,
gular scale rows from angle to angle of jaw 67, scales from rostral to
interparietal 16, scales from interparietal to anterior edge of first
collar 40, second collar absent, total dorsals 180, total ventrals 205,
scales within dorsal separation of first collar 1 7, number of spots
within dorsal separation of first collar 0, subdigital lamellae of second
toe of right hind foot 20, subdigital lamellae of fourth toe 40, sub-
digital lamellae of fifth toe 23.

The type is described from a museum specimen in which the
coloration compares favorably with live specimens seen from the
foothills NW of Palm Springs. Reticulation of head present as in
C. c. bicinctorcs but with larger freckles; body dorsum brown with
large oblong spots and three nearly complete light colored cross bars.
First collar widely separated dorsally and merging ventrally with
black of gular area. Second collar absent. Forelegs light, spotted
with brown hindlegs and tail brown, spotted with white. Dorsum of
tail ridged and compressed laterally.

Diagnosis. — Similar to C. c. bicinctores in many respects but
with greater dorsal separation of the first and second collars, with
distinct dorsal light cross bars, and a smaller second collar length/
snout-vent length index. The Baja subspecies can be distinguished
from C. i. insularis Van Denburgh and Slevin (1922) to which it is
most similar, by fewer toe lamellae on the second and fourth toe,
larger second collar/s-v index, and a smaller dorsal separation of
both the first and second collars.

Range. — East side of the mountains from central Baja Cali-
fornia, north to Palm Springs, Riverside Co., California. It does not
occur on the western Pacific slopes in California, and apparently
does not intergrade with C. c. bicinctores, although these papulations
are in close proximity in the Palm Springs area.

Specimens Examined

A total of 150 specimens from eight populations were used to
make comparisons. Twenty from each population except C. i. insu-
laris and C. c. dicker sonae, where only 17 and 13 were available
respectively for the computer analysis.

32 GREAT BASIN NATURALIST Vol. 32, No. 1

C. c. hicinc tores

California: Kern Co., BYU 31948; Riverside Co., BYU 32099.

Idaho: Butte Co., BYU 30772; Owyhee Co., BYU 2834.

Nevada: Clark Co., BYU 461; Nye Co., BYU 18815. 22194,
23882, 30088.

Utah- Emery Co., BYU 20090; Millard Co., BYU 8755, 8883,
21000, 21703; Tooele Co., BYU 14818, 14820-21; Utah Co., BYU
1455, 13041; Washington Co., BYU 12875.

C. c. dickersonae

Tiburon Isl-and: CAS 14005-06, 14008-12, 53263-64; SDSNII
46003-06.

C. i. insularis

Angel de la Gu.ardia Island: CAS 21948, 50874-76, 50878-79,
86754-55, 86783-84; CAS-SU 22712; LACM 4001-02, 9854; SDSNII
46001.

C. i. vestigium

Baja California: Canyon Guadalupe, Juarez Mtns., BYU
23337-39, and BYU 22298-99; North of Canipole. SDSNH 30109-
111; Coyete, SDSNII 52999; Baja Sur, SDSNH 30107; Los Angeles
Bay, SDSNH 19789-90, 41612, 52950; Las Palmitos, SDSNH 17052,
North Gonzaga Bay, SDSNH 19791-92; Sierra San Pedro, CAS-SU
17048; Santa Rosalia, CSCLB LWR670619-1 and 670619-2.

C. c. baileyi

Arizona: Apache Co., BYU 497; LACM 16895; UIMNH 7524;
USNM 29184. Coconino Co., BYU 506, 11388; UIMNH 6543, 35945;
USNM 15821. Graham Co., UIMNH 24507; USNM 5153, 51737.
Mohave Co., BYU 32116; UIMNH 74778, 74781. Navajo Co., BYU
13574. Pima Co., LACM 3983; SDSNH 15214. Yavapai Co., BYU
33322; UIMNH 5900.

C. c. auriccps

Colorado: Mesa Co., BYU 11342. San Miguel Co., CUM 1333,
4448, 4450, 4451, 4453.

Utah: Grand Co., BYU 1625, 10338, 12854. San Juan Co., BYU
1461, 1464. 12619. H()()6-()8, 16484, 21706, 32088; UU 1461, 2427.

r. r. fuse us

Mexico: Chihuahua, BYIT 13383-86, 13410, 14211; KU 3378,
33789; UC 70704; USNM 14242.

New Mexico: Dona Ana Co., LACM 3971; USNM 22268. Luna
Co., BYU 31940; USNM 44955, 80072. Sierra Co., LACM 3981.

Texas: EI Paso Co., USNM 59351, 59352; UTEP 52; UU 493.

March 1972 smith, tanner: crotaphytus 33

C. c. collaris

Colorado: Los Animas Co., CUM 1292, 2939.

Kansas: Montgomery Co., BYU 22167. Wilson Co., KU 41, 45,
46, 48, 54.

New Mexico: Chaves Co., LACM 3974, 3975. Lincoln Co.,
LACM 16990. Quay Co., USNM 44940.

Oklahoma: Carter Co., BYU 500, 1574.

Texas: Garza Co., CUM 32277. Randall Co., CUM 13554-56.
Roberts Co., USNM 32866. Stephens Co., BYU 13177.

Acknowledgment

We wish to thank Dr. W. E. Duellman, University of Kansas
(KU); Dr. J. M. Legler, University of Utah (UU); Dr. A. E. Levi-
ton and Dr. S. C. Anderson, California Academy of Sciences (CAS)
Dr. R. B. Loomis, California State College Long Beach (CSCLB)
Dr. T. P. Maslin, University of Colorado Museum (CUM); Dr. J. A
Peters and Dr. G. R. Zug, United States National Museum (USNM)
Mr. J. A. Sloan, San Diego Society of Natural History (SDSNH)
Mrs. Dorothy Smith, University of Illinois Natural History Museum
(UIMNH); Dr. R. G. Webb, University of Texas at El Paso
(UTEP); and Dr. J. W. Wright, Los Angeles County Museum
(LACM) for the generous loan of specimens. Thanks also to Mr.
William Ingram for making his coUans-cova\Ae-x. data and statistical
computer programs available, to James Davis, Palm Springs, Cali-
fornia, for specimens from that area, to J. W. Heinrichs for material
collected on the Baja California Peninsula, and to Brigham Young
University for use of specimens and facilities.

References

Allen, M. J. 1933. Report on a collection of amphibians and reptiles from

Sonora, Mexico, with description of a new lizard. Occ. Pap. Mas. Zool. Univ.

Mich. 259:1-15.
Burt, C. E. 1928. The synonymy, vanation and distribution of the collared

lizard, Crotaphytus collaris (Say). Occ. Pap. Mus. Zool. Univ. Mich. 196:

1-19.
Fitch, H. S., and W. W. Tanner. 1951. Remarks concerning the systematics

of the collared lizard {Crotaphytus collaris), with a description of a new sub-
species. Trans. Kans. Acad. Sci. 54:584-599.
Ingram. W., and W. W. Tanner. 1971. A taxonomic study of Crotaphytus

collaris between the Rio Grande and Colorado Rivers. Brigham Young Univ.

Sci. Bull. Biol. Ser. 13 (2): 1-29.
Mayr, E. 1969. Principles of systematic zoology. McGraw-Hill Book Co.. New

York. pp. xi, 428.
Say, T. 1823. In Long's account of an expedition from Pittsburgh to the Rocky

Mountains, performed in the years 1819 and '20. H. C. Carey and I. Lea.

Philadelphia, 1823, Vol. 2, pp. i-xcviii, 6-442.
Schmidt, K. P. 1922. The amphibians and reptiles of Lower California and

the neighboring islands. Bull. Amer. Mus. Nat. Hist. 46:607-707.

34 GREAT BASIN NATURALIST Vol. 32, No. 1

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

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

WisHART, D. 1968. A fortran II programme (CLUSTAN) for numerical classi-
fication. Computing Laboratory, St. Andrews, Fife, Scotland, pp. 1-50.

A NEW SPECIES OF STENAMMA (HYMENOPTERA:
FORMICIDAE) FROM UTAH

Robert E. Gregg^

Abstract. — Stenamma knowltoni is described as a new species from various
localities in northern Utah and southern Idaho. The new species falls clearly into
the large-eyed group of M. R. Smith's monograph of Stenamma.

Recently a number of excellent collections of ants from Utah were
received, and I am indebted to Dr. George F. Knowlton for these
specimens. Among the numerous vials are examples of a new species
of Stenamma which is herewith described.

Stenamma knowltoni, sp. nov.

Worker. — Length, 3.36 mm; head length (excluding mandi-
bles), 0.86 mm; head width, 0.76 mm; head index, 0.884; thorax
length, 1.056 mm.

Head distinctly longer than broad, subrectangular in shape; oc-
cipital border flat, occipital corners definite but rounded, sides of
head straight and parallel; median lobe of clypeus elevated and pro-
duced forward, anterior border denticulate, and lateral lobes receding
diagonally to mandibular insertions; frontal area de})ressed, triangu-
lar; frontal carinae parallel in front, slightly divergent behind, and
lobes partially covering antennal insertions. Antennae 12-segmented,
funicular segments 2-6 very slightly broader than long, last four seg-
ments forming a club whose terminal segment almost equals in
length two preceding it; scapes fail to reach occipital corners by an
amount less than length of funicular segment 1. Eyes very large for
a Stenamma, with about 12 facets across greatest diameter, approxi-
mately 70 facets in entire eye; eyes placed a little anterior to middle
of head from occiput to mandibles, and measuring 0.165 mm in
diameter. Mandibles bear seven teeth gradually reduced in size from
apex to basal border.

Thorax moderate in width, humeral angles rounded, dorsal sur-
face in profile gently convex, and meso-epinotal impression distinct
but shallow. E])inotal base coordinate with i)lane of promesonotum,
but sloping downward and backward to spines. Epinotal spines short
(0.033 mm and half as long as their interbasal distance), triangular
sharp, and pointing abruptly upward. Epinotal declivity vertical.
Petiolar node rounded, subconical, wider than long from above,
peduncle long, slender and constricted; in profile anterior face of
node sloping but distinct from peduncle; ventral surface of petiole
straight and furnished with a minute spine at anterior end. Post-
petiole globular from above, as wide as long; in profile convex above,
flat below. Gaster of usual myrmicine shape, first segment occupy-
ing most of this tagma.

'Department of Biology, University of Colorado, Boulder, Colorado 80302.

35

36

GREAT BASIN NATURALIST

Vol. 32, No. 1

Fig. 1. Head of Stenarnma knowltoni Gregg.

Sculpture: Head covered with longitudinal rugae, becoming
vaguely reticulate on posterior portion; interspaces rather coarsely
punctate, general appearance opaque to subopaque, or at most only
slightly shining. Clypeus with multiple carinulae. Mandibles finely
rugulate or striate, with widely spaced punctures, and shining. Dor-
sum of thorax longilu(hnally rugose, rugae widely spaced, but with
a few transverse rugae at anterior end. Pronotal collar jiunctate.
Pronotum lightly punctate, mesonotum heavily jiunctate between
rugae. Epinotal base punctate, declivity smooth and shining. Pro-
pleurae almost devoid of rugae, smooth and shining; mesopleurae
coarsely punctate especially in lower half, opaque; epinotal pleurae
rugose and coarsely punctate. Petiole and ])ostpetiole more finely
punctured dorsally and laterally, postpetiole having in addition
coarse longitudinal rugae on its dorsolateral as[)ect. Gaster smooth
and shining (faint shagreening), and with extremely short but
distinct basal striae. Whole insect oj)a(|ue to subojiaque, except
gaster, epinotal declivity smooth and shining, and pronotal dorsum
and extreme occipital border quite shining despite their sculpturing.

Pilosity: Delicate, rather long, pointed yellowish hairs on all
parts of body including appendages. Hairs uneven in length, shorter
and numerous on head, longer and sparser on thorax, pedicel, and
gaster.

Color: Head and gaster dark brown, thorax and pedicel slightly
lighter and with reddish tinge. Legs, antennae, mandibles, clypeus,
and pronotal collar light reddish brown.

P'l-.MALr. (deiilate). — Length, 4.4 mm; head length (excluding
mandibles), 0.92 mm; head width, 0.79 mm; head index. 0.857;
thorax length, 1.32 mm.

March 1972 gregg: a new ant 37

Similar to the worker in most respects except size and propor-
tions of thorax. Eye very large, 16 to 17 facets in greatest diameter,
and with a total of 135 to 140 facets. Ocelli distinct, not protuberant.
Scape fails to reach occipital border by an amount less than length
of funicular joint 1, and is similar in other respects to that of
worker.

Sculpture like that of worker in most details, but differing in that
promesonotum has more widely spaced rugae with fewer punctures
and therefore a more shining surface; propleurae rugose but inter-
spaces shining and without punctures; mesopleurae rugose dorsally,
almost sculptureless and shining ventrally; epinotal pleurae with
strong wavy rugae, few punctures, and shining. Base and declivity of
epinotum without sculpture and strongly shining. Petiolar and post-
petiolar nodes rugose. Epinotal spines longer, proportionately more
slender than in worker, and blunt at tips. Fore coxae rugose. Color
and pilosity very similar to that of worker.

Discussion. — A single alate female among the material sent to
me was compared to figures of the wing venation given by Smith
(1957) on page 1 37. The venation of the anterior wing of this speci-
men fits precisely that shown for Figure 2, a female of brevicorne,
and thus belongs to the ''brevicorne type," as designated by Smith,
for it possesses one submarginal cell only in addition to the discoidal,
and this condition is produced by the disappearance of vein r-m.
Mf3 and Mf4 are retained. The pattern of venation is thus consis-
tent with other characters which cause the new species (knowltoni)
to key out to the couplet separating brevicorne and meridionale.

When Dr. Smith monographed the genus Stenamnia he distin-
guished and described eleven species, and among these described
three as new. These ants are small, slow moving, soil and litter spe-
cies, secretive in habits, and generally not easily noticed in their
normal habitats. The total number of forms known is small (less
than 30 for the world according to Smith) and mostly Holarctic in
their general distribution. No new species in the United States
fauna have been discovered since Smith wrote, but the recent speci-
mens from Utah prove to be a heretofore unknown form and quite
distinct from any of those recognized previously. Smith went to
some pains to divide the group into species with small eyes and those
with large eyes, but this is only partially satisfactory; for, while it
is convenient, there is enough gradation in eye-size from the smallest
to the largest that no sharp break occurs among the forms on this
basis alone. Nevertheless, it is a useful procedure, and anyone wish-
ing to identify these ants must consider this character. In Smith's
key the new species comes out on the "large eye" couplet (5-12 om-
matidia), and then runs to the alternate separating brevicorne from
meridionale. All of the very small-eyed species are therefore easily
eliminated, but of the remainder several require that careful distinc-
tion be made. To judge from the characters given, knowltoni might
be confused with huachucanum because of a somewhat bicolored
appearance, but can be separated on the basis of its sculpture which
is longitudinally rugose rather than punctulate, the epinotal spines

38 GREAT BASIN NATURALIST Vol. 32, No. 1

which are not tuberculate or vestigial as in huachucanum, the petio-
lar node which is not unusually high, and the presence of basal gas-
tric rugulae or striae. S- carolinense has one of the largest eyes of
any of the species (10- 12 facets in greatest diameter), but it also
has transverse pronotal and mesonotal rugae, a more pronounced
meso-epinotal impression, and extremely blunt tuberculate spines,
all of which traits differ from knowltoni. S. brevicorne has a large
eye but only 8 to 10 facets across its greatest diameter. Further, it
has a strong meso-epinotal impression, and the epinotum is furnished
with a distinct transverse welt immediately behind the impression.
The epinotal spines are relatively long and sharp. The whole aspect
of the insect is subopaque with coarse longitudinal rugulae on the
dorsum of the promesonotum, and the color of the body is reddish
brown with the gaster light at the base. S. mendionaU\ also among
the larger-eyed species, has, again, only 8 to 10 ommatidia in the
greatest diameter. The meso-epinotal impression is pronounced, but
an epinotal welt is lacking. Epinotal spines are large, digitiform
(Smith), and sharp. The petiolar node is antero-posteriorly com-
pressed and rather quadrate from above (not conical), and the post-
petiole is subcampanulate (not oval).

HoLOTYPE. — Worker, in the author's collection. Paratypes: four
workers and one deiilate female deposited in the author's collection.
These four workers and one queen were collected in mossy sagebrush
duff at Cedar Creek Junction, near Kelton Pass, Box Elder Co.,
Utah, on 16 April 1969, by George F. Knowlton. The species is dedi-
cated to Dr Knowlton who has generously given me many fine
series of ants from various localities in Utah. Additional paratype
material also collected by Knowlton is available as follows: one
worker, sagebrush duff, Kelton Pass, Box Elder Co., Utah, 1 May
1969; six workers, rangeland. Cedar Creek, Box Elder Co., Utah, 1
April 1969; seven workers, dead grass, Kelton Pass, 5000 feet. Box
Elder Co., Utah, 16 Aj)ril 1969.

Specimens belonging to the new species but not part of the type
material were found by Knowlton at the following stations: three
workers, sagebrush duff in canyon, 6 miles east of Holbrook, Oneida
Co., Idaho, 16 April 1969; four workers, juniper duff, desert, 10
miles southwest of Twin Springs, Elmore Co., Idaho, 1 November
1969 (^389); two workers, greasewood duff, southeast of Black Pine,
Oneida (]o., Idaho, 18 October 1969 (-354); two workers, sagebrush
duff, Holbrook Sunmiit, 6115 feet, Oneida Co., Idaho, 22 October
1969 (*359); one alate female, desert biome. Curlew Reservoir, Cur-
lew National Cirassland, Oneida Co., Idaho, 22 October 1969 (-367).

S[)ecimens of Stcnamma knowltoni have been deposited in the
collections of W. S. Creighton and the United States National
Museum.

Before undertaking the description of this new species, I sub-
mitted specimens to Dr. (Creighton who was able to compare them
vvith exam})les of all known North American species of Stcnamma
(including the little known joveolocephalum), except carolinense.
I am indebted to him for confirming my supposition that knowltoni

March 1972 gregg: a new ant 39

represents a new species. He also drew my attention to a curious
situation in Smith's paper concerning eye-size in Stenamma, which
could be a potential hazard to anyone not aware of it. I take the
liberty to quote from his letter: "Cushman is a fine illustrator and
his figures can be trusted implicitly, but this is not the case in the
four outline drawings (5,6,7,8) on page 143 of Smith's monograph.
I am ready to agree that these figures are not drawn to the same
scale, S- huachucanum and impar are much smaller ants than occi-
dentale and schmitti. But in Smith's descriptions you will find that
in all four the maximum diameter of the eye is given as 0.10 mm.
Furthermore, the number of ommatidia across the greatest diameter
of the eye is closely similar for all four species; huachucanum 5-7,
impar 5-6, occidentale 4-6, schmitti 4-6. In short, they are all small-
eyed species. In my opinion the eyes of schmitti are shown as too
small and those of impar and huachucanum as too large. On the
basis of the figures, however, one would wonder why impar and
huachucanum were not brought out on the key split with the large-
eyed species. The only one of the four figures in which the eye-size
is correctly shown is occidentale."'

Despite the use of eye-size by Smith as a convenient method for
grouping the species of Stenamma, it appears that he relied heavily
upon thoracic sculpture in his revision. The extremely detailed and
accurate rendering of this sculpture by Cushman greatly facilitates
the separation of these species. I have indicated above how knowl-
toni differs from its most closely allied species, but it is well to em-
phasize here that its thoracic sculpture differs markedly from almost
all the other forms of Stenamma. The dorsum of the thorax is longi-
tudinally rugose for the most part, with a few transverse rugae an-
teriorly, and the only species with which it might be confused are
impar and brevicorne. Of these two ants, however, the former has
very small eyes and the latter has intermediate sized eyes as well as
other differences.

Finally, although eye-size will continue to be a useful character
in the taxonomy of Stenamma. it should be stressed that no clear-
cut distinction is possible between a small-eyed and a large-eyed
group of species. This can be demonstrated by expressing the eye-
size for all species, as Smith has done, in terms of the number of
facets in the greatest diameter, as follows: heathi. sequoiarum. impar.
diecki. schmitti, and occidentale 4-6; huachucanum 5-7; foveolocepha-
lum 7-8; brevicorne and meridionale 8-10; carolinense and knowl-
toni 10-12.

R

EFERENCES

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool.

104:1-585 (pp. 133-138; PL 17. fig. 1-4).
Smith, M. R. 1957. Revision of the Genus Stenamma Westwood in America

north of Mexico. Amer. Midi. Nat.. 57:133-174.

NEW SYNONYMY IN THE BARK BEETLE TRIBE
CRYPHALINI (COLEOPTERA: SCOLYTIDAE)^

Stephen L. Wood-

Abstract. — A lectotype designated for Bostrichus asperatus Gyllenhal result-
ed in the clarification of the status of Cryphalus Erichson and the removal of
Trypophoeus Fainnaire from synonymy. AH species described by A. D. Hopkins
in Hypothcnemus Westwood and Stephanoderes Eichhoff were reviewed. New
synonymj' included: Bostrichus asperatus Gyllenhal (= B. abietis Ratzeburg);
H. birmanus (Eichhoff) (= H. maculicollis Sharp. S. perkinsi Hopkins. S. ster-
culiae Hopkins. 5. psidii Hopkins); B. crudiae (Panzer) ( = B. plumeriae Nord-
linger, Cryphalus hispidulus LeConte, 5. differens Hopkins, S. paraguayensis
Hopkins. S. polyphagus Ck)sta Lima. 5. uniseriatus Eggers, S. hiyaoea Beeson, >*>.
lebronneci Beeson); H. calif amicus Hopkins (= H. tritici Hopkins); H. diptero-
carpi Hopkins (^ H. mangarivanus Beeson); H. erectus LeConte (= H. validus
Blandford, S. puncticollis Hopkms, S. cubensis Hopkins); H. eruditus Westwood
( = Cryphalus basjoo Niisima, Cosmoderes schwarzi Hopkins, H. bradfordi Hop-
kins. H. ferrugineus Hopkins, H. flavipes Hopkins, H. flavosquamosus Hopkins.
H. heathi Hopkins. H. koebelei Hopkins. H. lineatifrons Hopkins. H. mali Hop-
kins. H . myristicae Hopkins, H. nigricollis Hopkins, H. parvus Hopkins, H. puncti-
pennis Hopkins. H. sacchari Hopkins. H. tenuis Hopkins. H. webbi Hopkins,
S. elongatus Hopkins, S. flavicollis Hopkins, S. pygmaeus Hopkins. S. subconcen-
tralis Hopkins. S. unicolor Hopkins, H. intersetosus Eggers, S. erythrinae Eggers.
H. dubiosus Schedl).

On the following pages are presented (a) a lectotype designation
for Bostrichus asperatus Gyllenhal, 1813, type-species of the genus
Cryphalus Erichson, 1836, and the consequent effect of this act on
the genera Cryphalus and Trypophoeus Fairmaire, 1868, and (b) a
review of all species of bark beetles described by A. D. Hopkins in
the genera Hypothenernus Westwood and Stephanoderes Eichhoff.
Comments on the synonymy of Ernoporus Thomson are also in-
cluded.

Cryphalus Erichson

Cryphalus E.v'xchson (1836:61). Three species.

As originally proposed by Erichson (1836) the genus Cryphalus
contained three species, Apate tiliae Panzer, A. fagi Fabricius, and
Bostrichus asperatus Gyllenhal, as defined by references to Fabricius
(1801:383) for the first two species and to Gyllenhal (1813:368)
for the third species. Thomson (1859:46) designated B. asperatus
as the type-species of the genus Cryphalus and transferred A. tiliae
to his new genus Ernoporus. Thomson (1865: S6()) later transferred
A. fagi to Ernoporus, thus leaving B. asperatus as the oidy original
species remaining in Cryplialus. By definition the antennal funicle
of Cryphalus contained four segments.

Ratzeburg (1839:199) named Bostrichus (Cryphalus) ahietis and
commented that until then this species had been known by his co-
workers as B. asperatus Ciylleidial, presumably includijig Erichson.

'Tho research on which this paper was based was supported by the National Science Foundation.
^DeiKirtnient of Zoology, Brigham Young University, Provo, Utah 84601. Scolytoidea contribu-
tion no. 34.

40

March 1972 wood: bark beetle synonymy 41

Based on syntypes in the Germar collection, now in the Zoologisches
Museum, at Berlin, Ratzeburg (1839:198-199) redefined the true
B. asperatus (iyllenhal and described several new species, one of
which was B. {Cryphalus) binodulus.

On the basis of the 5-segmented antennal funicle, Faimiaire
(1868:105) transferred B. binodulus Ratzeburg, 1839, to his new
genus Trypophoeus. Later, Eichhoff (1878:139) placed B. binodulus
as a junior subjective synonym of B. asperatus in the genus Glyp-
toderus Eichhoff, 1878, = Trypophoeus Fairmaire, 1868, even though
B. asperatus was the type-species of Cryphalus Erichson, 1836. Eich-
hoff's usage continued until the error was pointed out by Wood
(1954:988).

In an effort to clarify the nomenclatural confusion and to con-
serve the traditional usage of the name Cryphalus, an appeal to the
International Commission on Zoological Nomenclature was prepared
(Wood, 1967) requesting that B. abietis Ratzeburg, 1839, be desig-
nated the type-species of Cryphalus since Erichson's concept of B.
asperatus apparently was based on erroneously identified specimens
of B. abietis. When the appeal was considered by the Commission, a
request was made that all type material be examined by me before
any action be taken on this appeal. The following is a report on my
examination of that material.

The Fabricius collection at the Universitetets Zoologiske Museum,
at Copenhagen, contains several s})ecimens of A. tiliae Panzer, all of
which are of the species currently known as Ernoporus tiliae (Pan-
zer); also present there is one damaged specimen labeled A. fagi
Fabricius, although it actually is of /I. tiliae, and almost certainly is
not the type of the Fabricius species. The Germar collection at the
Berlin Museum did not contain original specimens of A. fagi either.

Six syntypes of B. asperatus Gyllenhal and two of B. asperatus
var. B of Gyllenhal were located; three syntypes and the two vari-
ants are at the University of Uppsala, in Gyllenhal's Insecta Suecia
collection, and three syntypes are at the Berlin Museum in the Ger-
mar material. The latter three syntypes apparently are those used
by Ratzeburg (1839:198-199) for his redescription of the species.
All six syntypes and the first of the two variants are identical
and represent the same species described as B. (Cryphalus) abietis
Ratzeburg, 1839. The other variant of B. asperatus Gyllenhal is of
Trypophoeus spiculatus Eggers, 1927. Therefore, B. binodulus Ratze-
burg, 1839, and Trypophoeus Fairmaire, 1868, the genus for which
it is the type-species, have no bearing whatever on the synonymy of
Cryphalus Erichson, 1836, or on its type-species B. asperatus Gyllen-
hal, 1813. I here designate as the lectotype of B. asperatus Gyllenhal
the first syntype in the above-mentioned Gyllenhal Insecta Suecia
series at the University of Uppsala Museum.

Cryphalus asperatus (Gyllenhal)

Bostrichus asperatus Gyllenhal (1813:368). Lectotype, male; presumably from
Sweden; Univ. Uppsala Mus., present designation, above.

42 GREAT BASIN NATURALIST Vol. 32, No. 1

Bostrichus (Cryphalus) abietis Ratzeburg (1839:198). Syntypes; presumably
destroyed with the Hamburg Museum. I^ew synonymy.

As indicated in the above discussion, the type series of Bostrichus
asperatus Gyllenhal was incorrectly identified by Eichhoff (1878)
and subsequent workers. These specimens are of the same species
that has been known since 1839 as abietis Ratzeburg. Specimens of
abietis at the U.S. National Museum and the British Museum (Nat-
ural History) compared to the types by Eggers, Eichhoff, and, pre-
sumably, Blandford were used as a basis for this species.

Ernoporus Thomson

Ernoporus Thomson (1859:147). Type-species: Apate tiliae Panzer, monobasic.

Schedl (1962:92-94), apparently using antennal characters ex-
clusively, treated Ptilopodius Hopkins, 1915, Stephanorhopalus Hop
kins, 1915, Ernoporicus Berger, 1917, AUernoporus Kurenzov, 1941,
and Eocryphalus Kurenzov, 1941, as synonyms of Ernoporus. Au-
thenticated specimens of the type-species of each of these genera are
at hand or were recently studied by me, as well as all known species
of Ernoporus^ four additional species of Ptilopodius (sensu Hopkins),
and two species erroneously placed in Ptilopodius by Schedl, except
for Eocryphalus which is known to me only from the description.
From this material it is apparent that Ptilopodius and Stephanor-
hopalus are completely unrelated to the other genera; the similarity
of the antennae to other species mentioned here is superficial at best.
(These genera will be treated in greater detail at a later date.)
Ernoporicus spessivtzevi Berger, type-species of the monotypic genus
Ernoporicus, is a typical Ernoporus except for the smaller antennal
club and very obscure sutures on the club; I agree with Schedl in
transferring it to Ernoporus. My specimen of AUernoporus evonymi
Kurenzov is very closely allied to Ernoporus, but the antennal funi-
cle is 3-segrnented and the club is totally devoid of sutures; until
more material is available for study I prefer to retain this species in
AUernoporus.

Hypothenemus Westwood

Hypothenemus We&twood (1836:34). Type-species: Hypothenemus eruditus West-
wood.

Hopkins (1915) described 106 species in the genera Hypothene-
mus Westwood, 1836, and Stephanoderes Eichhoff, 1871. Since then
Stephanoderes has been placed in synonymy (Browne, 1963:53)
under Hypothenemus. The species named by Hopkins in this taxon
were based on unique females, and most of them have not been ex-
amined by specialists of the group since then. Recently it was my
j)rivilege to study holotypes of all of the Hopkins species named in
these genera as well as to study the tyi)es of a few other species of
special interest in a review of the Hopkins material. Several of the
American species were previously placed in synonymy (Wood,

March 1972 wood: bark beetle synonymy 43

A review of all species named by Hopkins follows. Valid names
are presented in alphabetical order with synonyms and my com-
ments listed beneath them. Of the 106 species named by Hopkins 21
are considered valid and 85 are treated as synonyms.

Hypothenemus africanus (Hopkins)

Stephanoderes africanus Hopkins (1915:30). Holotype, female; Capetown, South
Africa; USNM, 7542.

This species is allied to setosus (Eichhoff). Specimens have been
examined from the following new localities. Jamaica, 13-IX-35, in
Poinciana pods; Lagunillas, Merida, Venezuela, 12-1-70, 1000 m ele-
vation. Mimosa twig, S. L. Wood; Buitenzorg, Java, 7-VIII-35 (host
not legible), L. G. E. Kalshoven; Singapore, Malaya, IX-64, Man-
gifera indica, N. L. H. Krauss.

The above Java specimen was received from Kalshoven and
bears an unsigned label in his handwriting ''Stephanoderes multi-
punctatus Schedl." This specimen bears the same data as Schedl's
type of multipuncatus. Although Schedl's type was not available for
study, it is highly probable that it is a junior synonym of africanus.

Hypothenemus brunneus (Hopkins)

Stephanoderes brunneus Hopkins (1915:31). Holotype, female; Texas; USNM.
7545.

The name Stephanoderes frontalis Hopkins is a synonym of
brunneus (Wood, 1954:1031). This species almost certainly was
introduced to America from Africa where all near relatives appear
to have originated, although no African specimens have been ex-
amined. It is closely allied to 56^/051/5 (Eichhoff).

Hypothenemus birmanus (Eichhoff)

Triarmocerus birmanus Eichhoff (1878:42, 486). Holotype, female; Burma; pre-
sumably lost in Hamburg Mus.

Hypothenemus maculicollis Sharp (1879:101). Syntypes; Oahu, Hawaiian Islands;
British Mus. Nat. Hist. New synonymy.

Stephanoderes perkinsi Hopkins (1915:31). Holotype, female; Honolulu, Hawaii;
USNM, 7594. New synonymy.

Stephanoderes sterculiae Hopkins (1915:32). Holotype, female; Calapan, Philip-
pine Islands; USNM, 7551. New synonymy.

Stephanoderes psidii Hopkins (1915:32). Holotype, female; Calapan, Philippine
Islands; USNM, 7552. New synonymy.

Previously designated synonyms include Stephanoderes alter
Eggers, S. pacificus Beeson, and S. castaneus Wood (Wood, 1960:35).
The syntypes of maculicollis Sharp, and the holotypes of S. perkinsi
Hopkins, S. sterculiae Hopkins, and S. psidii Hopkins were studied
and all agree with my specimens that were compared to material
Schedl compared to the type of birmanus. It is a common species in
Indonesia, southern Asia, the Pacific Islands, Central America, and
southern Florida.

44 GREAT BASIN NATURALIST Vol. 32, No. 1

Hypothenemus calijornicus Hopkins

Hypothenemus calijornicus Hopkins (1915:19). Holotype, female; Pomona, Cali-
fornia; USNM, 7364.

Hypothenemus tritici Hopkins (1915:19). Holotype, female; Dallas. Texas;
USNM, 7526. New synonymy.

This species occurs in the southern United States, California,
Mexico, and one series was seen from Liberia in western Africa. It
is very closely allied to other African species, including albipilus
Reitter, and probably was introduced to America from Africa. Wood
(1954:1055) treated H. tritici Hopkins as a subspecies and H. thora-
cicus Hopkins as a synonym of tritici. In view of the extended dis-
tribution into Mexico, with intergradation, and its discovery in
Liberia, the status of tritici should be reduced to that of a junior
subjective synonym.

Hypothenemus ceihae Hopkins

Hypothenemus ceibae Hopkins (1915:20). Holotype, female; Cayamas, Cuba;
USNM. 7583.

Apparently this is a distinct species similar to but larger than
H. eruditus Westwood, with very slender interstrial scales.

Hypothenemus columhi Hopkins

Hypothenemus columbi Hopkins (1915:18). Holotype, female; Columbus. Texas;
USNM, 7361.

This common distinctive species occurs from the southern United
States to Colombia and Venezuela. Previously published synonyms
of Hopkins's species include H. ahdominales, H. rufopalliatus, H.
brunneipennis, and //. amplipennis (Wood, 1954:162).

Hypothenemus crudiae (Panzer)

Bostrichus crudiae Panzer (1791:35-38). Syntypes.

Stephanodercs ohscurus: Eggers (1929:50, nee Fabririus. 1801).

Bostrichus plurnrriae Nordlinger (1856:74). Syntypes: Venezuela.

Cryphalus hispidulus LeConte (1868:156). Syntypes. New synonymy.

Stephanoderes differens Hopkins (1915:25). Holotype, female; San Bernardino,
Paraguay; USNM, 7541. New synonymy.

Stephanoderes paraguayensis Hopkins (1915:26). Holotype, female; San Ber-
nardino, Paraguay; USNM, 7^77. New synonymy.

Stephanoderes polyphagus Costa Lima (1924, nee Eggers, 1924). Syntypes? New
synonymy.

Stephanoderes uniseriatus Eggers (1924:103). Lectotype, female; Luebo, Congo;
USNM, 60169. New synonymy.

Stephanoderes hivaoea Beeson (1935:105). Holotype, female; Tahauku Hivaoa,
Marquesas Islands; Bishop Mus. New synonymy.

Stephanoderes lebronneci Beeson (1935:104). Syntypes. New synonymy.

The types of crudiae (Panzer) and plumeriae Nordlinger have
not been examined by me. The usage of these names is based on

March 1972 wood: bark beetle synonymy 45

series in the Eggers collection, at the U.S. National Museum, that
apparently were based on authentic specimens. These specimens
agree with syntypes of hispidulus LeConte, polyphagus Costa Lima,
lehronncci Beeson, the lectotype of uniseriatus Eggers, and the holo-
types of diffcrcns Hopkins, paraguayensis Hopkins, and hivaoea
Beeson. The series of three specimens of Hylesinus obscurus Fa-
bricius in the Copenhagen Museum does not include a specimen of
this species (see H. obscurus, below). Hopkins's names previously
placed in synonymy under this species include brasiliensis, guata-
malensis, and lecontei (Wood, 1954:1041). The origin of this pan-
tropical species is uncertain, but it probably is American.

Hypothenemus cylindricus (Hopkins)

Stephanoderes cylindricus Hopkins (1915:25). Holotype, female; Trece Aguas,

Alta Verapaz, Guatemala; USNM, 7564.
Hypothenemus pallidus Hopkins (1915:18). Holotype, female; Mount Coffee,

Liberia; USNM, 7590. New synonymy.
Stephanoderes transatlanticus Eggers (1941:99). Holotype, female; Trois Rivieres,

Guadeloupe; Paris Mus. New synonymy.

Hopkins's holotypes of cylindricus and pallidus and two cotypes of
transatlanticus Eggers were compared directly and apparently all
represent the same species. The option available to me to ignore
page priority is exercised and I select cylindricus as the name for
this species, because large series of American specimens are available
for study.

Hypothenemus dipterocarpi Hopkins

Hypothenemus dipterocarpi Hopkins (1915:17). Holotype, female; Calapan, Min-

doro, Philippme Islands; USNM, 7588.
Hypothenemus mangarevanus Beeson (1940:196). Holotype, female; Aukena,

Margareva Islands; Bishop Mus. New synonymy .

The holotypes of both dipterocarpi Hopkins and mangarevanus
Beeson were compared to my Micronesia specimens to establish the
above synonymy. The interstrial bristles are scalelike on the disc
and hairlike on the declivity of this distinctive species.

Hypothenemus dolichocola Hopkins

Hypothenemus dolichocola Hopkins (1915:19). Holotype, female; Canton, China;
USNM, 7580.

This species resembles eruditus Westwood in all respects, except
the frons. The frons is shallowly, transversely impressed; it is about
intermediate between vafer Blandford and eruditus on the lower half
of the frons. Apparently it is a valid species, but additional material
should be examined.

Hypothenemus erectus LeConte

Hypothenemus erectus LeConte (1876:356). Lectotype, female; Texas; Mus.
Comp. Zool., present designation.

46 GREAT BASIN NATURALIST Vol. 32, No. 1

Hypothenemus validus Blandford (1904:228). Holotype, female; Motzorongo,
Veracruz. Mexico; British Mus. Nat. Hist. New synonymy.

Stephanoderes puncticollis Hopkins (1915:32). Holotype, female; Tampico, Ta-
maulipas. Me.xico; IJSNM. 7547. New synonymy.

Stephanoderes cubensis Hopkins (1915:32). Holotype, female; Cayamas, Cuba;
USNM, 7553. New synonymy.

The holotypes of H. validus Blandford, S. puncticollis Hopkins,
and S. cubensis Hopkins and a syntype of erectus LeConte were all
examined and were found to represent the same species. The only
syntype of erectus LeConte remaining in the LeConte collection and
labeled as the type, is here designated as lectotype of this species. A
cotype of Stephanoderes samhesianus Eggers, in the Eggers collection
at the U.S. National Museum, apparently is conspecific with erectus.
It is also noted that three cotypes of iS. mozamhiquensis Eggers and
one cotype of S. dispar Eggers are doubtfully distinct from this spe-
cies. More material from additional African localities should be
studied before this synonymy is established.

Since this species has no close relative in America that was not
introduced through commerce, and since all closely allied species are
from Africa, I susj>ect this species is of African origin. Its introduc-
tion into America evidently occurred long before it was described.
It is common from southern Texas to Venezuela.

Hypothenemus eruditus Westwood

Hypothenemus eruditus Westwood (1834:36). Syntypes, England?

Cryphalus hasjoo Niisima (1910:9). Syntypes; Tokyo, Japan. New synonymy.

Cosmoderes schwarzi Hopkins (1915:11). Holotype. female; Haw Creek, Florida;

lost except slide mount of antenna in USNM. New synonymy.
Hypothenemus bradfordi Hopkins (1915:15). Holotype, female; Honolulu, Ha-
waii; USNM, 7567. New synonymy.
Hypothenemus ferrugineus Hopkins (1915:20). Holotype, female; Trece Aguas.

Aha Verapaz, Guatemala; USNM, 7584. New synonymy.
Hypothenemus flauipes Hopkins (1915:18). Holotype, female; Cayamas. Cuba;

USNM, 7575. New synonymy.
Hypothenemus fhwosquamo.sus Hopkins (1910:15). Holotype, female; Mount

Coffee. Liberia; USNM, 7591. New synonymy.
Hypothenemus heathi Hopkins (1915:20). Holotype. female; Independencia.

Parahyba. Brazil; USNM. 7521. New synonymy.
Hypothenemus koebelei Hopkins (1915:17). Holotype, female; Brazil; USNM,

7572. New synonymy.
Hypothenemus lineatifrons Hopkins (1915:17). Holotype. female; Cayamas.

Cuba; USNM. 7570. New synonymy.
Hypothenemus mali Hopkins (1915:17). Holotype, female; Capetown, South

Africa; USNM, 7573. New synonymy.
Hypothenemus myristicoe Hopkins (1915:16). Holotype, female; Buitenzorg,

.Fava; USNM. 7589. New synonymy.
Hypothenemus nigricoUis Hopkins (1915:16). Holotype. female; Capetown. South

Africa; USNM. 7568. New synonymy.
Hypothenemus parvus Hopkins (1915:17). Holotype. female; Cayamas, Cuba;

USNM, 7574. New synonymy.
Hypothenemus punctipennis Hopkins (1915:20). Holotype. female; Capetown,

"West" Africa; USNM, 7585. New synonymy.

March 1972 wood: bark beetle synonymy 47

Hypothenemus sacchari Hopkins (1915:17). Holotype, female; Nevis, West

Indies; USNM, 7379. New synonymy.
Hypothenemus tenuis Hopkins (1915:16). Holotype, female; Trece Aguas, Alta

Verapaz, Guatemala; USNM, 7569. New synonymy.
Hypothenemus webbi Hopkins (1915:17). Holotype, female; Calapan. Mindoro,

Philippine Islands; USNM, 7587. New synonymy.
Hypothenemus inter setosus Eggers (1928:85). Lectotype, female; Sao Paulo,

Brazil; USNM, 60153. New synonymy.
Stephanoderes elongatus Hopkins (1915:25). Holotype, female; Cayamas, Cuba;

USNM, 7561. New synonymy.
Stephanoderes flavicollis Hopkins (1915:24). Holotype, female; Cayamas, Cuba;

USNM, 7559. New synonymy.
Stephanoderes pygmaeus Hopkins (1915:24). Holotype, female; Pagbilao, Philip-
pine Islands; USNM, 7560. New synonymy.
Stephanoderes subconcentralis Hopkins (1915:25). Holotype, female; Cayamas,

Cuba; USNM, 7563. New synonymy.
Stephanoderes unicolor Hopkins (1915:25). Holotype, female; Cayamas, Cuba;

USNM, 7562. New synonymy.
Stephanoderes erythrinae Eggers (1936:628). Holotype, female; Sakalaspur,

India; British Mus. Nat. Hist. New synonymy.
Hypothenemus dubiosus Schedl (1940:207). Syntypes; Hamburgfami, Ebene

Limon, Costa Rica. New synonymy.

In establishing the above synonymy, the following specimens
were examined and compared to my material and to one another:
Cryphalus basjoo Niisima, 4 cotypes; Hopkins's holotypes of brad-
fordi, jerrugineus, flavipes, flavosquamosus, heathi, koebelei, lineati-
frons, mail, myristicae, nigricollis, parvus, punctipennis, sacchari,
tenuis, webbi elongatus, flavicollis, pygmaeus, subconcentralis, and
unicolor; the lectotype of intersetosus Eggers ; a cotype of erythrinae
Eggers; and a syntype labeled "type" of dubiosus Schedl. In addi-
tion, the balsam mount of the antenna of Cosmoderes schwarzi Hop-
kins was examined (the type was lost). Since this antenna is entirely
typical of cruditus, because Hopkins did not have a taxonomic knowl-
edge of the group with which he was working, and in view of the
fact that the description of the type fits eruditus (except for the er-
roneously described antenna), schwarzi is placed in synonymy under
eruditus.

Into this species I have grou{)ed similar forms in which the hair-
like setae of the elytra vary from uniseriate and sparse strial rows
to confused and moderately abundant. There appears to be a com-
plete transition from one extreme to the other. With that exception,
interstrial scales, features of the head, pronotum and elytra are
rather uniform.

Hypothenemus flavus Hopkins

Hypothenemus flavus Hopkins (1915:17). Holotype, female; Java; USNM, 7571.

This species evidently is distinct. It is allied to eruditus West-
wood.

Hypothenemus glabripennis (Hopkins)

Stephanoderes glabripennis Hopkins (1915:32). Holotype, female; Angat, Philip
pine Islands; USNM. 7548.

48 GREAT BASIN NATURALIST Vol. 32, No. 1

This distinctive species is well known to specialists.

Hypothcncmus gossypii (Hopkins)

Stephanoderes gossypii Hopkins (1915:25). Holotvpe, female; Cayamas, Cuba;
USNM, 7557.

The synonymy between gossypii Hopkins and H. beameri Wood
has been established (Wood, 1962, Gt. Basin Nat. 22:78). Since H.
gossypii Sampson evidently is a nomen nudum Hopkins's name is
used here.

Hypothenemus hampei (Ferrari)

Cryphalus hampei Ferrari (1868:11. 12). Syntypes.

Stephanoderes punctatus Eggers (1924:101). Lertotype. female; Eala, Congo;
USNM, 60160. New synonymy.

The lectotype of punctatus Eggers and the holotype of cooki
Hopkins were compared to my series of this well-known species and
were found to be identical. It is an important pest of coffee.

Hypothenemus interstitialis (Hopkins)

Hypothenemus interstitialis Hopkins (1915:28). Holotype, female; Victoria.

Texas; USNM, 7555.
Stephanoderes obliquus Hopkins (1915:30). Holotype, female; Cayamas, Cuba;

USNM, 7538. New synonymy.

The holotypes of interstitialis Hopkins and obliquus Hopkins
were compared directly to establish the above synonymy. Wood
(1954: 1033) also placed Hopkins's interpunctus. approximatus, flave-
scens, opacipennis, and quadridentatus in synonymy under inter-
stitialis.

Hypothenemus liberiensis (Hopkins)

Stephanoderes liberiensis Hopkins (1915:31). Holotype, female; Mount Coffee,
Liberia; USNM, 7593.

This species is very closely allied to erectus (LeConte), but the
pronotal asperities are smaller, and the lateral areas of the pronotal
disc are rugulose. The holotype is 1.8 mm in length.

Hypothenemus mallyi (Hopkins)

Stephanoderes mallyi Hopkins (1915:32). Holotype, female; Capetown, South

Africa; USNM. 7549.
Stephanoderes soussouensis Eggers (1943:74). Holotype. female; Sone, Zambeze;

Paris Mus. New synonymy .

The holotype of mallyi Hopkins and the cotype of soussouensis
Eggers in the Eggers collection at the U.S. National Museum, were
compared and found to represent the same species. It is allied to
rotundicollis (Eichhoff ) .

March 1972 wood: bark beetle synonymy 49

Hypothenemus rnultidcntatus (Hopkins)

Stephanoderes multidentatus Hopkins (1915:28). Holotype, female; Tampico.
Tamaulipas. Mexico; USNM, 7532.

Stephanoderes ferrugineus Hopkins (1915:29, nee Hopkins, 1915:20). Holotype,
female; Livingston, Guatemala; USNM, 7535. New synonymy.

Stephanoderes nitidifrons Hopkins (1915:31). Holotype, female; Tampico. Ta-
maulipas, Me.xico; USNM, 7546. New synonyrrjy.

The holotypes of Hopkins's multidentatus, ferrugineus, and nitidi-
frons were compared directly to one another. They all represent a
species that is very close to inter stitialis Hopkins.

Hypothenemus ohscurus (Fabricius)

Hylesinus obscurus Fabricius (1801:395). Lectotype, female; Essequibo. British

Guiana; Copenhagen Mus., present designation.
Hypothenemus kiinnemanni Reitter (1902:140). Lectotype, female; Breman,

Germany, in Brazil nuts; Budapest, Mus., present designation. New

synonymy.
Stephanoderes moschatae Schaufuss (1905:8, reprint p. 2). Holotype, female;

Guadeloupe; presumably lost with Hamburg Mus. New synonymy.
Stephanoderes rufescens Hopkins (1915:29). Holotype, female; Allegheny, Penn-
sylvania; USNM, 7527. New synonymy.
Stephanoderes buscki Hopkins (1915:30). Holotype, female; Trinidad, West

Indies; USNM. 7537. New synonymy.
Stephanoderes amazonicus Eggers (1934:78). Lectotype, female; Manaos, Brazil;

USNM, 60142. New synonymy.

The entire type series of obscurus Fabricius, kiinnemanni Reitter,
rufescens Hopkins, and buscki Hopkins were examined, as well as
the lectotype of amazonicus Eggers. Several specimens of moschatae
Schaufuss compared by Eggers to the type were also examined. All
of these were compared to my homotypes.

This very common species occurs from Costa Rica and Puerto
Rico to Brazil where it breeds in twigs, nuts, and fruits of a wide
variety of hosts. It is best known from infested Brazil nuts that are
transported through commerce to virtually all parts of the world.

The type series of obscurus Fabricius consists of three female
specimens in the Copenhagen Museum. The first two are of this
species, the third (Kiel specimen) is in poor condition and probabh^
is of pulverulentus Eichhoff, but might possibly be of crudiae Pan-
zer. Since the second specimen is in better condition than the first,
I designate it as lectotype of Hylesinus obscurus Fabricius; a red,
printed lectotype label was placed on the pin bearing this specimen.

The Reitter syntypes of kiinnemanni consisted of five identical
females labeled "Bremen, XII - 1900, Paraniise." The second speci-
men was in the best condition and was labeled and is here designated
the lectotype of Hypothenemus kiinnemanni .

Hypothenemus parallelus Hopkins

Hypothenemus parallelus Hopkins (1910:25). Holotype, female; Tampico, Mexi-
co; USNM, 7556.

50 GREAT BASIN NATURALIST Vol. 32, No. 1

1 his form is essentially identical to eruditus Westwood except
for the frons which bears a small, low nodule at the center. It prob-
ably represents a genetic variation within a normal population, but
due to the breeding habits and (suspected) partial parthenogenesis
the frontal character appears to have greater importance than is
warranted. Series in my collection are from Mexico and the Ha-
waiian Islands.

Hypothenemus pilosus Hopkins

Hypothenemus pilosus Hopkins (1915:20). Holotype. female; Cayamas, Cuba;

USNM. 7586.

In this distinctive species the rows of interstrial setae are hairlike
from the elytral base to the apex. Apparently it is very rare.

Hypothenemus pubescens Hopkins

Hypothenemus pubescens Hopkins (1915:19). Holotype. female; Key West,

Florida; USNM. 7524.
Hypothenemus subelongatus Hopkins (1915:19). Holotype, female; Victoria,

Texas; USNM. 7581. New synonymy.
Stephanoderes opacifrons Hopkins (1915:25). Holotype. female; Aguadilla. Puerto

Rico; USNM, 7565. New synonymy.

This species is almost identical with sparsus Hopkins, but it pos-
sesses rows of strial hair and lacks interstrial granules. In addition
to the localities cited above it occurs in Mexico and Hawaii. It
breeds in the axis of fruiting stems of various grasses, including the
genera Andropogon, Cynodon. and Paspalum.

Hypothenemus pulverulentus (Eichhoff)

Stephanoderes pulverulentus Eichhoff (1871:33). Syntypes(?), female; Mexico;

presumably lost with Hamburg Mus.
Stephanoderes vulgaris Schaufuss (1897:209). Syntypes. female; La Digue.

SechcUe Islands; presumably lost witli Hamburg Mus.
Stephanoderes georgiae Hopkins (1915:27). Holotype, female; Georgia; USNM,

7385. New synonymy.
Stephanoderes tamarindi Hopkins (1915:27). Holotype, female; Manila. Philip-
pine Islands; USNM, 7530. New synonymy.
Stephanoderes niger Hopkins (1915:31). Holotype. female; Brownsville. Texas;

USNM, 7382. New synonymy.
Stephanoderes nitidipennis Hopkins (1915:29). Holotypye. female; Cayamas.

Cuba; USNM. 7533. New synonymy.
Stephanoderes fiehrigi Hopkins (1915:27). Holotype, female; San Bernardino,

Paraguay; USNM. 7387. New synonymy.
Stephanoderes minutus Hopkins (1915:26). Holotype, female, evidently defective;

Cayamas, Cuba; USNM, 7366. New synonymy.
Hypothenemus emarginatus Schedl (1942b: 11). Syntypes, female; Buitenzorg,

.lava. New synonymy.
Stephanoderes darwinensis Schedl (1942a: 178). Syntypes; Australia. New

synonymy.
Stephanoderes andersoni Wood (1954:1045). Holotype, female; Coconut Grove,

Florida; USNM. New synonymy.

March 1972 wood: bark beetle synonymy 51

Stephanoderes liquidambarae Wood (1954:1046). Holotype, female; Jacksonboro,
South Carolina; Snow Ent. Mus., Univ. Kansas. New synonymy.

This abundant, widely distributed species is distinguished with
difficulty from ohscurus (Fabricius) by characters of the frons and
elytral surface. It is possible that multidcntatus Hopkins is a popu-
lation variant at the upper limits of size. The above synonymy was
based on a study of the holotypes of Hopkins's georgiae, tamarindi,
niger. nitidipennis, fichcrgi, and rninutus; on syntypes of emargina-
tus Schedl and darwincnsis Schedl; and on holotypes of andersoni
Wood and liquidomhari Wood. The holotype of rninutus evidently
is defective. There is a certain amount of variation in the minute
details of sculpturing of the frons and in the shape of the interstrial
scales. After examining many hundreds of specimens from America,
the Pacific area, and the Indo-Australian region, I see no alternative
to grouping all of the above under one name. The difficulty is com-
plicated by the intensive inbreeding coupled with (suspected) partial
parthenogenesis which may produce morphologically uniform local
populations. When all of these local po])ulations are studied, how-
ever, they intergrade completely. Previously published synonymy of
Hopkins's species was established for texanus, pini, salicis, floridensis,
ficus, soltaui lucosi, vircntis, pecanus, and nigcr (Wood 1954:1035,
1048). The basis for the names pulverulentus and vulgaris was
specimens identified by and presumably compared to the types by
Eggers.

Hypothencmus rotundicollis (Eichhoff)

Stephanoderes rotundicollis Eichhoff (1878:45, 145). Syntypes (?), female; Ameri-
ca septentrionalis; presumably lost with Hamburg Mus.

The synonymy of this species with one Hopkins's species, quer-
cus, has been established (Wood, 1054:1024). In addition to its dis-
tribution in the eastern and southern United States, it occurs in the
states of Tamaulipas and Nayarit in Mexico. Allied species occur
in Central America.

Hypothcnemus setosus (Eichhoff)

Hypoborus (?) setosus Eichhoff (1867:391). Syntypes; Guadeloupe.
Stephanoderes bananensis Eggers (1922:167). Two syntypes; Banana, Congo; one

in Eggers collection. New synonymy.
Stephanoderes kalshoveni Schedl (1939:35). Syntypes; Pasoeroean, Java; Buiten-

zorg Mus. New synonymy.
Stephanoderes subagnatus Eggers (1940:101). Holotype, female; Eala. Congo;

Tervuren Mus. New synonymy.

Hopkins's species for which synonymy previously was established
include ohesus and philippincnsis (Wood, 1957:402). The above
new synonymy was based on a syntype of bananensis and two syn-
types of subagnatus. Several specimens of kalshoveni received from
Kalshoven and taken by him from the same branch as the syntypes,
were used to establish the synonymy of Schedl's species.

It is difficult to establish the origin of this species at the present

52 GREAT BASIN NATURALIST Vol. 32, No. 1

time, but it evidently reached America from Africa where several
allied species occur, or less probably from the Indo-Malayan area.

Hypothencmus squamosus (Hopkins)

Strphanoderes squamosus Hopkins (1915:26). Holotype, female; Cayamas. Cuba;

IJSNM, 7566.

This distinctive species occurs from southern Florida and Cuba
to Mexico. Its nearest relative occurs in Mexico.

Hypothenemus sparsus Hopkins

Hypothenemus sparsus Hopkins (1915:20). Holotype. female; Columbus, Te.xas;
USNM. 7368.

Two of Hopkins's species, similis and tridentatus, have been
placed in synonymy under this name (Wood, 1954:1040). It is rare
and distinguished with difficulty from pulverulentus (Eichhoff) and
pubescens Hopkins.

Hypothenemus vafer Bland ford

Hypothenemus vafer Blandford (1896:241). Syntypes; Noumea, New Caledonia;

British Mus. Nat. Hist.
Stephanoderes polyphagus Eggers (1924:104). Syntypes. New synonymy.
Hypothenemus heterolepsis Costa Lima (1928:117). Syntypes. New synonymy.
Stephanoderes subvestitus Eggers (1940:232). Holotype, female; Mosolo Kwenge,

Kwongo, Congo; Tervuren Mus. New synonymy.
Stephanoderes martiniquensis Eggers (1941:99). Holotype. female; St. Pierre,

Martinique; USNM. 60156. New synonymy.

In view of its present distribution and abundance, it is most re-
markable that Hopkins did not encounter this species in his study.
No examples of it were included in the collection he studied. This
is the species to which I previously have referred as areccae 1 lornung
{= fungicola Eggers, hispid us Eggers, etc.). Since a question has
been raised as to the true identity of areccae, the next oldest name
known to me for this species, vafer, will be used until the types can
be examined.

The above synonymy was based on the type series of vafer
Blandford, on two syntyj)es of polyphagus Eggers, two syntypes of
heterolepsis Costa Lima, one cotyjie of suhvestitus Eggers, and the
holotyf)e of martiniquensis Eggers.

It apparently is now established in southern Florida. It previous-
ly has been reported from Brazil, Martinique, Plawaiian Islands,
Micronesia, Philippine Islands, the Indo-Malayan region, Chana,
and the Congo. It is polyj)hagous and, presumably, it can breed in
nuts, twigs, or bark.

I'risrhidids atoma (Hopkins)

Hypothenemus atomus Hoi)kins (1910:15). Holotype, female; Morgantown, West
Virginia; USNM, 7565.

March 1972 wood: bark beetle synonymy 53

Hopkins's species Hypothenemus impressifrons, marylandicae,
rohiniae, and toxicodendri were placed in synonymy under atomus
by Wood (1954:1068) and transferred to the genus Trischidias
Hopkins.

References Cited

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Bull. B. P. Bishop Mus. 142:115-121.
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Occ. Pap. B. P. Bishop Mus. 15:191-203.
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Ent. Beligique 40:241-245.
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Berichten 23:53-59.
Eggers, H. 1922. Neue Borkenkafer (Ipidae) aus Afrika. Ent. Blatt. 18:163-174.

. 1924. Neue Borkenkafer (Ipidae) aus Afrika. Ent. Blatt. 20:99-111.

. 1928. Ipidae (Coleoptera) da America do Sul. Archiv. Inst. Biol. Def.

Agric. Anim. 1:83-99.
. 1929. Zur synonymie der Borkenkafer (Ipidae, Col.). Wiener Ent.

Zeit. 46:41-55.

. 1934. Borkenkafer (Ipidae, Col.) aus Siidamerika. Ent. Blatt. 30:78-84.

. 1936. Neue Borkenkafer (Scolytidae, Col.) aus Indien. Ann. Mag. Nat.

Hist. (10)17:626-636.
. 1940a. Neue Borkenkafer (Col., Scolytidae) aus Afrika. Nachtrag IX.

Rev. Zool. Bot. Afr. 33:99-108.
. 1940b. Neue Borkenkafer (Col., Scolytidae) aus Afrika. Nachtrag X.

Rev. Zool. Bot. Afr. 33:227-239.
. 1941. Borkenkafer aus Siidamerika (Coleoptera: Ipidae). IX. Insel

Guadeloupe. Arb. morph. taxon. Ent. Berlin-Dahlem 8:99-109.
. 1943. Neue Borkenkafer (Ipidae) aus Afrika. Nachtrag VIII. Ent.

Blatt. 39:70-76.
Costa Lim.\, A. M. da. 1924. J. do Commercio, Oct. 19 (Not seen).
EiCHHOFF, W. J. 1867. In: Kraatz, G., Beitrage zur Kenntniss der deutschen

Kaferfauna. 4. Stuck. Berliner Ent. Zeitschr. 11:391.
. 1871. Neue exotische Tomiciden-Arten. Berliner Ent. Zeitschr. 15:

131-136.
. 1878. Ratio, descriptio, emendatio eorum tomicinorum qui sunt in Dr.

medic. Chapuisii et autoris ipsius collectionibus et quos praeterea recognovit.

Mem. Soc. Sci. Liege (2)8:1-531 (1879, preprint 1878).
Erichson, W. F. 1836. Systematische Auseinandersetzung der Familie der

Borkenkafer (Bostrichidae). Archiv. Naturg. 2(l):45-65.
Fabricius, J. C. 1801. Systema eleutheratorum. Kilia: Bibliopolii Academici.

Vol. 2, 687 p.
Fairmaire, L. 1868. In: Jacquelin du Val and Fairmaire. Genera des coleop-

teres d'Europe. Vol. 4, 292 p.
Ferrari, J. A. 1867. Die Forst- und Baumzuchtschadlichen Borkenkafer (Tomi-

cides Lac.) aus der Familie der Holtzverderber (Scolytides Lac), mit beson-

derer Beriicksichtigung vorziiglich der europaischen Formen, und der Sam-

mulung des k. k. zoologischen Kabinetes in Wien. Ceroid: Wien. 96 p.
Gyllenhal, L. 1813. Insecta Suecica descripta. Classis I. Coleoptera Siue

Eleuterata. Vol. 1, pt. 3, 730 p.
Hopkins, A. D. 1915. Classification of the Cryphalinae, with descriptions of

new genera and species. U.S. Dept. Agric, Sec Rept. 99:1-75.
LeConte, J. L. 1868. Notes and appendix. In: Zimmermann, Synopsis of the

Scolytidae of America north of Mexico. Trans. American Ent. Soc 2:141-178.
NoRDLiNGER, H. 1856. Nachtrage zu Ratzeburgs Forstinsekten. Stuttgart. 83 p.

(Not seen).
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Nat. Hist. Soc. 3:1-18.

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Panzer, G. W. F. 1791. Beschreibung eines sehr kleinen Kapuskafers. Natur-

forscher 25:35-38. (Not seen).
Ratzeburg, J. T. C. 1839. Die Forstinsekten der Abbildung und Beschreibung

der in den Nachbarstaaten als schadlich oder niitzlich bekannt gewordenen

Insekten. Vol. 1, Die Kafer (Borkenkafer, p. 168-232).
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Zeit. 21:137-141.
ScHAUFUss, C. 1897. Beitrag zur Kaferfauna Madagascars. III. Tijdschr. Ent.

40:209-225.

. Borkenkaferstudien. Insekt. Borse 22: (Reprint, p. 1-12).

ScHEDL, K. E. 1939. Scolytidae und Platypodidae. 47 Beitrag. Tijdschr. Ent.

82:30-53.
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taxon. Ent. Berlin-Dahlem 7:203-208.
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australischen region. Mitt. Miinchner Ent. Ges. 32:162-201.

. 1942b. Neue Scolytidae aus Java. Tijdschr. Ent. 85:1-49.

1962. Zur synonymie der Borkenkafer X. Mitt. Miinchner Ent. Ges.

52:85-107.
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Westwood, J. O. 1836. Description of a minute Coleopterous insect, forming

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. 1957. Distributional notes on and synonvmies of some North American

Scolytidae (Coleoptera). Canadian Ent. 89:396-403.

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24:121-122.

THE CONTRIBUTIONS OF THE 1822 WORKS OF JAROCKI
AND FLEMING TO HERPETOLOGICAL NOMENCLATURE

Robert C. Feuer" and Hobart M. Smith'

Two great synoptic works on amphibians and reptiles, of con-
siderable unfamiliarity to herpetologists yet of marked historical
importance, appeared in 1822: one in English, by Flemmg, the
other in Polish, by Jarocki. Both works are rare and seldom exam-
ined, but Jarocki's has the additional handicap of being written
in a language very little known to systematists in general. An
analysis of the nomenclatural contributions of these two works,
and of their mutual priority, is long overdue.

The question of priority is important because in both works
some new names were proposed for the same taxa. Neave credited
Fleming with priority, without documentation, but our evidence
conclusively gives Jarocki priority. According to the Director of
the Biblioteka Narodowa in Warsaw, Zbigniew Daszkowski, a
short review of Jarocki's "Zoologia" with a statement that "three
volimies have been published so far" appeared in the number (3)
of Gazeta Liter acka [the Literary Gazette] dated 19 Feb. 1822.
Number 5 of the same gazette, dated 5 Mar., gave a bibliographical
description of volume 3 of the "Zoologia" in a column entitled
"Chronicle of National Literature— New Works Which Have Ap-
peared in Print." The evidence justifies the conclusion that volume
3 of Jarocki's work, in which amphibians and reptiles are treated,
appeared no later than February 1822.

The work by Fleming, on the other hand, was indicated by Miss
Ann Lucas, British Museum (Natural History) Library, as having
appeared no earlier than May and no later than June 1822. A
letter received from the National Library of Scotland states that
Fleming's work was entered "at Stationer's Hall" on 28 June 1822
and that an undated advertisement received during the period May-
July 1822 advised that "this day is published, in two handsome
volumes octavo, with engravings, price LI. 10s in boards. The Phi-
losophy of Zoology ..."

Several inquiries elsewhere have yielded no further information
pertinent to date of publication. Since the available data clearly
indicate publication of Jarocki's work at least by 19 Feb. 1822,
and of Fleming's work sometime during May or June but no later
than 28 June, acceptance of nomenclatural priority of Jarocki over
Fleming is inescapable, except by exercise of the plenary powers
of the International Commission on Zoological Nomenclature.

In conjunction with a review of the names in which priority
is involved, the entire nomenclatural contribution of both works,
at least at the generic level, should be reviewed. We here deal with

^Philadelphia College of Pharmacy and Science, 43rd St., KingsessLng and Woodland avenues,
Philadelphia, Penn. 19104; and Department of Biology, University of Colorado, Boulder 80.302.

55

56 GREAT BASIN NATURALIST Vol. 32, No. 1

the herpetological names only, leaving those applied to birds, mam-
mals, and other groups for investigation by others.

Jarocki

Sixty-five generic names were used by Jarocki, including 4
for anurans (Bufo, Hyla, Pipa, Rana), 5 for salamanders (Axo-
lotus, Proteus, Salamandra, Triton, Siren), 1 for caecilians {Coe-
cilia), 6 for turtles (Chelonia, Chelys, Emys, Ophichelone, Testudo,
Trionyx), 3 for crocodilians {Alligator. Crocodilus. Gavial), 3
for amphisbaenians {Amphisbaena, Bipes, Chirotes), 22 for lizards
(Acontias, Agama, Ameiva, Angins, Anolius. Basiliscus, Chalcides,
Chamaeleo, Cordylus, Draco, Dracoena, Gecko, Jguana, Lacerta,
Lacertus. Lophyrus, Monitor, Ophisaurus, Polychrus, Scincus, Stel-
lio, Uroplatus), and 21 for snakes (Acanthophis, Acrochordus, Boa,
Bungarus, Coluber, Crotalus, Elaps, Erix, Erpeton, Hurria, Hydrus,
Langaia, Naja, Platurus. Pseudoboa. Python, Scytale, Tortrix, Tri-
mesurus, Typhlops, Vipera).

Of these 65 names, 3 were not listed by Neave: Angvis, Dra-
coena and Jguana. Neave explicitly equated Angvis and Anguis
and implicitly equated Jguana and Iguana; both of these alternate
names have priority over Jarocki's usages, the different characters
of which are character-equivalents with early classical Latin (u was
written as v, j as /). Dracoena, not listed by Neave or Romer, has
apparently been regarded as an erroneous subsequent spelling,
judging from the fact that it was explicitly based upon the tautonym
Lacerta dracoena Linnaeus, which was originally spelled dracaena.
The least disturbing and therefore most acceptable conclusion at
this point is to assume that both spelling variants {Dracoena, dra-
coena) in Jarocki were inadvertent errors which have no nomencla-
tural status. Even if the generic spelling were regarded as an oc-
cupied name, it would remain a junior synonym of Dracaena Dau-
din, 1802.

The only acceptably new names, at least not previously used,
are Axolotus, Langaia, Ophichelone, and Trimesurus. Axolotus was
suppressed by the International Commission (1963:102) in order
to conserve Amby stoma Tschudi, 1838. Ophichelone, based upon
Testudo serpentina Linnaeus, is antedated by Chelydra Schweigger,
1814. Langaia is a junior synonym and an invalid emendation of
Langaha Brugniere, 1784, but has previously been attributed, er-
roneously, to Fleming (1822). Iriniesurus, erroneously attributed
first to Gray, 1840, is a junior synonym and an invalid emendation
of Trimeresurus Lacopede, 1804. Had Jarocki's I^acertus, based on
Lacerta orbicularis Linnaeus, not been antedated by Lacertus Lace-
pede, 1788, an invalid emendation of Lacerta Linnaeus, 1758, it
would be available for Phrynosoma Wiegmann, 1828.

Fleming
Seventy-two generic names were used by Fleming, including
4 for salamanders {Apneumona, Salamandra, Sirena, Triton), 4

March 1972 feuer, smith: nomenclature 57

for anurans (Bufo, Hyla, Pipa, Rana), 1 for caecilians (Cecilia), 8
for turtles {Chelonia, Chelonura. Chelys, Cistuda, Coriudo, Emys,
Testudo, Trionix), 3 for crocodilians (Alligator, Crocodilus, Gaviala),
3 for amphisbaenians (Amphisbaena, Bipes, Chirotes), 24 for lizards
(Acontias, Agama, Ameiva, Anguis, Anolius, Ascalabotes, Basiliscus,
Calotes, Chalcides, Chameleon, Cordylus, Casta, Dracaena, Draco,
Iguana, Lacerta, Lophurus, Mastigura. Monitor, Ophisaurus, Poly-
chrus. Scincus, Stellio, Trapelus), and 25 for snakes (Acanthophis,
Acrochordus, Boa. Caudisona, Cerastes, Chersea, Cobra, Coluber,
Crotalus, Elaps, Erpeton, Hemachatus, Hydrophus, Langaia, Natrix,
Naja, Pelamis, Platurus, Pseudoboa, Scytalus, Tortrix, Trigonoce-
phalus, Trimeresura, Typhlops, Vipera).

Neave regarded 14 of these generic names as new and not previ-
ously used: Apneumona, Chameleon, Chelonura, Chersea, Cistuda,
Coriudo, Custa, Gaviala, Hemachatus, Hydrophus, Langaia, Lophur-
us, Mastigura, Trimeresura. As indicated in the preceding discus-
sion, Jarocki's Langaia antedates and therefore replaces Fleming's
Langaia, leaving but 13 names not used before Fleming. Romer
rejected Ascalabotes of Cuvier, 1817, originating the name with
Lichtenstein, 1823. Were the rejection of Cuvier, 1817, as a source
for this name upheld (we are aware of no reason, however), Flem-
ing's usage would antedate Lichtenstein's, restoring the total of 14
names originating with Fleming.

The 13 names we believe properly credited as originating with
Fleming are allocated as follows: Apneumona — \\xn\OT synonym of
Proteus Laurenti, 1768; Chameleon = ]\xrnoY synonym and invalid
emendation of Chamaeleo Laurenti, 1768; Chelonura = junior syno-
nym of Chelydra Schweigger, 1812; Chersea = ]unior synonym of
Vipera Laurenti, 1768; Cistuda = junior synonym of Terrapene
Merrem, 1820; CorzWo=: junior synonym of Dermochelys Blain-
ville, 1816; Custa = junior synonym of Tupinambis Daudin, 1803
(Peters et al., 1970:271); Gaviala = junior synonym and invalid
emendation of Gavialis Oppel, 1811; Hemachatus = \a\id, recognized
name; Hydrophus=^ junior synonym of Aipysurus Lacepede, 1804;
Lophur us — junior synonym of Lyriocephalus Merrem, 1820; Mas-
tigura = junior synonym of Uromastix Merrem, 1820; and Trimere-
sura = junior synonym and invaHd emendation of Trimeresurus La-
cepede, 1804.

Type-Species

Neither Jarocki nor Fleming indicated author for their generic
names, and likewise neither usually indicated whether any given
name was proposed as new or not. Accordingly, we regard as "new"
only those names not previously used in that exact spelling. For
such names it is important that type-species be known in order to
permit an incontrovertible nomenclatural allocation.

A total of 17 occupied (i.e., acceptable for nomenclatural con-
sideration) generic names originated with either Jarocki or Flem-
ing. Three other names used by Jarocki are not considered as occu-
pied and therefore do not exist nomenclaturally: Angvis, Dracoena,

58 GREAT BASIN NATURALIST Vol. 32. No. 1

J guana (see preceding account for explanation). The 17 occupied
names have the following type-species:

1. Apneumona Fleming (1822:303), type-species by monotypy
"/I. anguina' = Proteus anguinus Laurenti.

2. Axolotus Jarocki (1822:179), type-species by subsequent
designation (Smith and Tihen, 1961:216) '"Axolotus pisciformis,"
ex Siren pisciformis Shaw = Amhystoma mexicanum Shaw. For an
English translation of the section of Jarocki on Axolotus, see Szarski,
Smith, and Smith (1970:6-7).

3. Chamelon Fleming (1822:272), no species listed; as an
apparent unjustified emendation of Chamaeleo Laurenti, 1768, it
thereby takes the type-species of the latter name, to wit, Chamaeleo
parisiensium haurenti ^Chartiaeleo chamaeleon (Linnaeus, 1758).
The earlier generic name Chamaeleon Gronovius, 1 763, was elimi-
nated by the International Commission on Zoological Nomenclature
(Opinion 89, 1925) through rejection of Grovonius's work of 1763
for nomenclatural purposes.

4. Chelonura Fleming (1822:270), type-species by monotypy
"Testudo serpentina'' [Linnaeus, 17 58} = C he lydr a serpentina (Lin-
naeus) .

5. Chersea Fleming (1822:295), type-species by monotypy
"C. vulgaris'' ex Vipera vulgaris Sonnini and Latreille, 1802 =
Vipera aspis (Linnaeus, 1758).

6. Cistuda Fleming (1822:270), no species mentioned; type-
species by present designation Testudo Carolina Linnaeus, 1758 =
Terrapene Carolina (Linnaeus). The name Cistudo Say, 1825, long
used for this genus in the 19th century literature, was an emenda-
tion of Fleming's Cistuda and therefore invalid at the outset, since
Article 69 of the International Code states that generic names pub-
lished without species names, jirior to 1931, are not thereby un-
available but take as tyj)e-s{)ecies the first one subsequently desig-
nated for it.

7. Coriudo Fleming (1822:271), type-species by monotypy
''Testudo coriacea" [Linnaeus, l766]=Dermochelys coriacea (Lin-
naeus).

8. Casta Fleming (1822:274), type-species "L. teguexin" ex
Lacerta teguixin Linnaeus, \758 = 7upinamhis teguixin (Linnaeus),
by subsequent designation (Peters and Donoso-Barros, 1970:271).

9. Gaviala Fleming (1822:276), type-species by monotypy
Lacerta gangrtica [Gmolin, \789']=Gavialis gangeticus (Gmelin).

10. Hemacfiatus Fleming (1822:295), type-species by mono-
tyj)y '7/. vulgaris,"' a new sj)ecies-group name (although not so
designated by Fleming), a junior synonym, almost universally over-
looked, of Coluber haemachata Lacepede, \7S9 = Hemachatus hae-
machatus ( I .acepede ) .

11. Hydrophus Fleming (1822:292), type-species by mono-
type '7/. ayspisurus:" Neave regarded this name as an emendation

March 1972 feuer, smith: nomenclature 59

of Hydrophis Latreille, 1801; since Fleming did not so state, and
did cite a species name, the latter must be regarded as the type-
species, and Hydrophus an independently new name, not a sub-
stitute new name. The species name is also new, although not so
designated, and has almost universally been overlooked. It appears
to be derived, with a misspelling, from Lacepede's Aipysurus laevis
of 1804, of which it is here designated a synonym. Therefore Hy-
drophus ayspisurus Fleming, \^22^=^ Aipysurus laevis Lacepede,
1804, and the genus Hydrophus Fleming, 1822, becomes a junior
synonym of Aipysurus Lacepede, 1804, instead of Hydrophis La-
treille, 1801.

12. Langaia Jarocki (1822:102), type-species Langaia nasuta
ex Langaha nasuta Brugniere, 1 784, by monotypy. It is by no means
certain whether the Jarocki spelling was a deliberate emendation
(and therefore nomenclaturally occupied) or an inadvertent mis-
spelling (and therefore unoccupied); Neave and most others con-
sidered it occupied, and we accept that decision.

13. Lophurus Fleming (1822:278) was treated without any
species-group names. It might be regarded as an emendation of
Lophyrus Dumeril, 1806, itself unavailable as a junior homonym
of Lophyrus Poli, 1791, a molluscan; to so conclude, however, would
effect replacement of Gonocephalus Kaup, 1825, a long-established
genus. Precisely what species Fleming had in mind is not evident,
but it is illuminating to observe that the two species Jarocki placed
in his probably equivalent genus Lophyrus were Lacerta scutata
and Lacerta superciliosa, both Linnaean names. The latter is now
placed in the monotypic genus Uranoscodon Kaup, 1825; the for-
mer is placed in the monotypic genus Lyriocephalus Merrem, 1820.
The least disruptive allocation of Lophurus Fleming is to the genus
Lyriocephalus, of which it would be a junior synonym; accordingly,
we here designate Lacerta scutata Linnaeus the type-species of Lo-
phurus Fleming, 1822, a junior synonym of Lyriocephalus Merrem,
1820.

14. Mastigura Fleming (1822:277), type-species by monotypy
"M. spinipes"" = Stellio spinipes Daudin, 1802 = Uromastix spinipes
(Daudin), type-species also of Uromastix Merrem, 1820 (the name
was also spelled Uromastyx by Merrem in the same work, but that
spelling has not been adopted and is thus invalid) .

15. Ophichelone Jarocki (1822:21), type-species by monotypy
Testudo serpentina Gmelin ex Testudo serpentina Linnaeus, 1758 =
Chelydra serpentina (Linnaeus).

16. Trimeresura Fleming (1822:291) is presumably an emen-
dation of Trimeresurus Lacepede, 1804, the type-species of which
(Vipera viridis Daudin, 1803) automatically becomes that of Tri-
meresurus also. The result would be the same if Fleming's name
were regarded as independently proposed, for the only species
placed in it is 'T. viridis'^ = Trimeresurus gramineus (Shaw, 1802).

17. Trimesurus Jarocki (1822:103) is also clearly an emenda-
tion or lapsus for Trimeresurus Lacepede, 1804. Neave stated that

60 GREAT BASIN NATURALIST Vol. 32, No. 1

the orthography Trimesurus originated with Gray, 1840, perhaps
regarding one usage dehberate, the other inadvertent. We see no
reason for discrimination in this way and regard the spelling as
deliberate with Jarocki and therefore originating in 1822. The
type-species is therefore the same as for Trimeresurus Lacepede,
namely Vipera viridis Daudin, \^{)i = Coluber gramineus Shaw,
18Q2 = Tnmeresurus gramineus (Shaw).

Summary

Jarocki (1822) has priority over Fleming (1822), the former
appearing at least by February, the latter in May or June. The only
new generic names originating from Jarocki are Axolotus. Langaia,
Ophichelont\ and Trimesurus. none of which is valid today: all are
junior synonyms except Axolotus. which has been suppressed in
order to conserve Amhystoma Tschudi, 1838. The only new generic
names originating from Fleming are Apneumona. Chameleon, Chel-
onura, Chersea, Cistuda, Coriudo. Custa, Gaviala, Hemachatus, Hy-
drophus, Lophurus, Mastigura, and Trimeresura, only one of which
{Hemachatus) is vaHd today. The type-species and present alloca-
tion of all genera are summarized. Lacerta scutata Linnaeus, 1758,
is here designated the type-species of Lophurus Fleming in order to
prevent replacement of C^onocephalus Kaup, 1825, or Uranoscodon
Kaup, 1825. Two generally overlooked species-group names were
created by Fleming: Hemacfiatus vulgaris, a junior synonym of
Hemachatus haemachatus (Lacepede), and Hydrophus ayspisurus,
a junior synonym of Aipysurus laevis Lacepede. The spelling vari-
ants Angvis. Iguana, and Dracaena occurring in Jarocki are re-
garded as having no nomenclatural status: they are "unoccupied."

Literature Cited

Fleming, John. 1822. The philosophy of zoology. London, Constable. 2 vols.
May-June.

International Commission on Zoological Nomenclature. 1963. Opinion
649. Ambystoma Tschudi, 1838 (Amphibia); validation under the plenary
powers. Bull. Zool. Nomencl.. 20(2): 102-104.

Jarocki. Feli.x Pawel. 1922 Zoologia czyli Zwierzetopismo ogolne podlug
Najnowszego Systematu ulozone przez Feliza Pawla Jarockiego, . . . Tom
trzeci. Gady i Plazy. Warsaw, Latkiewicza. [6], 184, [11] pp., 3 pis. Febru-
ary.

Neave, Sheffield Airey. 1939-1950. Nomenclator zoologicus. London, Zool.
Soc. London. 5 vols. (A 6th vol. was prepared by different authors.)

Peters, James A., and Roberix) Donoso-Barros. 1970. Catalogue of the Neo-
tropical Squamata. Part II. Lizards and amphisbaenians. Bull. U.S Nat.
Mus. 297:i-viii, 1-293, ill.

Romer, Alfred Sherwood. 1956. Osteologj^ of the reptiles. Chicago. Univ.
Chicago Press, xxiv, 772 pp., 248 figs.

Smith, Hohart M., and Joseph A. Tihen. 1961. Tigrina (Salamandra)
Green, 1825: prof>osed validation under the plenary powers (Amphibia,
Caudata). Bull. Zool. Nomencl. 18(3):214-216.

SzARSKi, Henryk. Hobart M. Smith and Rozella B. Smith. 1970. Polish
(ontributions to the study of the axolotl, Ambystoma mezicanum. Bull.
Philadelphia Herp. Soc. 17: 4-16, fig. 1 (1969).

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TABLE OF CONTENTS

Fossil palm materials from the Tertiary Dipping Vat Forma-
tion of central Utah. William D. Tidwell, David A.
Medlyn, and Gregory F. Thayn 1

A new subspecies of Crotalus lepidus from western Mexico.
Wilmer W. Tanner, James R. Dixon, and Herbert S.
Harris, Jr 16

Two new subspecies of Crotaphytus (Sauria; Iguanidae).

Nathan M. Smith and WiLmer W. Taimer 25

A new species of Stenamma (Hymenoptera: Formicidae)

from Utah. Robert E. Gregg 35

New synonjony in the bark beetle tribe Cryphalini (Coleop-

tera: Scol3rtidae). Stephen L. Wood 40

The contributions of the 1822 works of Jarocki and Fleming
to herpetological nomenclature. Robert C. Feuer and
Hobart M. Smith 55

The

MUS. CO MP. ZOOL
LiB.RARY

A^R 1 Wo

UNiV_;.RSll-Y

Volume 32, No. 2
June 30, 1972

Great Basin

mmmm

Published by
Bhigham Young University

GREAT BASIN NATURALIST

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

Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W.
Tanner, Zoology; Joseph R. Murdock, Botany; Vernon J.
Tipton, Zoology; Ferron L. Andersen, Zoology

Ex officio Editorial Board Members:

A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences

Ernest L. Olson, Director, University Press, University
Editor

The Great Basin Naturalist was founded in 1939 by Vasco M.
Tanner. It has been continuously published from one to four times
a year since then by Brigham Young University, Provo, Utah. In
general, only original, previously unpublished manuscripts pertain-
ing to the biological natural history of the Great Basin and western
North America will be accepted. Manuscripts are subject to the ap-
proval of the editor.

Subscription: The annual subscription is $5.00 (outside the
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should consult the instructions printed on the back cover of a recent
number.

The Great Basin Naturalist

Published at Provo, Utah, by

Brigham Young University

Volume 32

June 30, 1972 No. 2

PRFY STAGE DISTRIBUTION, A FACTOR AFFECTING THE
^^^^ ^SuMERICAL RESPONSE

OCCIDENT ALIS TO TETRANYCHUS MCDANIELI
AND T£;T/?A/VyC//Z75 PACIFICUS^

B. A. Croft2

Abstract — The ability of Typhlodromus occidentalis to oviposit and nu-

most ux^avoraiSeTt high densities or late season after spider mxtes overexploited
their host plant.

A predator's reproductive rate is greatly infl^^"?^^,^/^ /^.^^
density, nutritional suitability, and behavioral acceptability of its
p?ey For example. Chant (1961) reported that the rate of
^position of the phytoseiid mite Typhlodromus occidenfis^^
bitt, was closely correlated with the number of Pacific spider nutes
{Tetranychus pacificus McGregor) it consumed. Burrell and Mc-
Cormick (1964) reported differential oviposition rates for 1 occi-
dZTaHs when feeding on various tetrany chid prey; ^^d f mWr^^^^
potentillae (Garman) consumes fewer prey/day P^^.7™\^f^j;*^
of egg production when feeding on Panonychus ulmi (K^^^h) reared
on leaves having a high nitrogen level, as compared to P-^/m.
reared on leaves with low nitrogen content (van de Vrie and Boers-

"^^Many^ phytoseiid mites prefer feeding on a If T^jf "1?^ /tage
or staged of certain tetranychid mites (Chant, 1959; Putnam,

"^^^^^oxlV^Ai^l^Su.e University, East I^nsin.. Michigan 48823.

61

62 GREAT BASIN NATURALIST Vol. 32, No. 2

1962; Croft and Jorgensen, 1969; van de Vrie and Boersma, 1971).
Croft and McMurtry (1972a) reported that T. occidentalis preferred
feeding on the egg, larval, and protonymphal stages of Tet. pacificus,
rather than on late nymphal-male adult or female adult mites.
When provided with different stage groups of Tet. pacificus [(a)
eggs, (b) imfed larvae, (c) deutonymphs-male adults, (d) female
adults], predators exhibited marked differences in oviposition rates.
Estimates were twofold greater when T. occidentalis females were
fed prey eggs or larvae as compared to deutonymphs — male adults
and female adults. The relationship between the prey stage con-
sumed and the oviposition rate of predators suggested that the
stage distribution of the prey mites might affect the ability of T. occi-
dentalis to numerically increase and control Tet. pacificus and
Tetranychus mcdanieli McGregor in the field. The investigations
reported herein were conducted to test this hypothesis. This rela-
tionship was studied at several interaction levels; in the laboratory
on confined 6x6 cm paper substrates, apple leaves, and 2.5 cm di-
ameter apple-leaf disks and in the field on apple leaves, in apple
trees, and in an apple orchard throughout a growing season.

General Methods

Rearing methods for the predator and prey populations have
been described elsewhere (Croft, 1970). Stock colonies of T. occi-
dentalis originally were collected from an apple orchard at Wenat-
chee, Washington. Tetranychus pacificus was started from a colony
maintained by L. R. Jeppson at the University of California, River-
side. All laboratory experiments were conducted, unless specified
otherwise, at 75 ± S^F and 50± 10% RH in the insectary at the
University of California, Riverside. Field experiments, samples and
populations of Tet. mcdanieli were all taken in apple orchards at
Oak Glen, near Yucaipa, California. With respect to taxonomy,
morphology, and behavior, Tet. pacificus and Tet. mcdanieli are
closely related species, and T. occidentalis is well adapted to either
prey (Flaherty, 1967;^ Hoyt 1969).

Predators Reared on Paper Substrates

Methods. — The potential of T. occidentalis to numerically in-
crease when fed four life stage groups of Tet. pacificus was tested
on waterproofed construction paper units (6x6 cm). These arenas
were placed on a water- saturated, plastic foam base and were con-
tained in a 15x15x3 cm stainless steel pan. Cellucotton strips bor-
dered each unit and were saturated with water to insure the iso-
lation of each predator-prey population. Life stage groups of Tet.
pacificus: (a) eggs-larvae, (b) larvae-protonymphs, (c) deuto-
nymphs-male adiJts, and (d) female adults were collected by the
method described by Scriven and McMurtry (1971). An abimdance

'Flaherty, D. 1967. The ecology and importance of spider mites on grape vines in the
southern San Joaquin Valley with emphasis on the role of Metaseiulus occidentalis (Nesbitt).
Doctoral Dissertation, Univ. of Calif., Berkeley.

June 1972 croft: mite predation 63

of each prey group was added three times weekly to units where
30 newly oviposited eggs of the predator previously had been
placed. A single pan contained four replicates of each treatment.
After female predators had developed to maturity, morality (U)
and oviposition (mx) measurements were collected daily durmg the
entire oviposition period (16-29 days). Six parameters, includmg
(1) mean eggs produced/female, (2) mean egg production per fe-
male/day, (3) mean length of the oviposition period, (4) mean
generation time (T), (5) net reproductive rate per female/gener-
ation (/?o), and (6) instrinsic rate of increase (r), were measured.
A previously reported sex ratio (Croft, 1970) of 1: 1.7 ( cT : ? ) for T.
occidentalis was used in all calculations.

Results. — Table 1 contains the six respective parameters for
adult females of T. occidentalis when fed on each stage group of
Tet. pacificus. The mean total egg production for females fed on
each prey group were similar. Mean egg production per female/
day and the mean length of the oviposition period were inversely
correlated; high oviposition rates were associated with short ovi-
position periods and vice versa. If similar mortality (Ix) values
had been obtained for each stage type, mean generation time (7)
would have positively correlated with the length of the oviposition
period. However, a greater mortality occurred among predators
when feeding on deutonymphs-male adults and female adults as
compared with those predators feedings on eggs-larvae or larvae-
protonymphs. It is not known if these mortality differences are also
associated with the consumption of the larger prey stages as oc-
curs in the field. The lower mortality (Ix) values also caused the
mean generation time (7) for predators fed on deutonymphs-male
adults and female adults to be similar to those values obtained for
mites fed on eggs-larvae and larvae-protonymphs. The number
of progeny produced per female (/?o) in each generation (T) was
greater among mites feeding on eggs-larvae and larvae-protonymphs
as compared to those feeding on the larger prey stages. Intrinsic
rates of increase {r) were positively correlated with the mean esti-
mates for egg production per female/day.

Table 1. Oviposition and rate of increase parameters for Typhlodromus
occidentalis when fed each of four life stage groups of Tetranychus pacificus.

Life Stage
type

Mean Total
Eggs/Female

Mean Eggs/ Mean Length of
Day/Female Oviposition (Days)

T

Ro

r

Egg-larvae

36.8

2.3

16.0

14.4

21.6

0.213

Larvae-proto-
nymph

35.7

1.9

19.0

16.4

21.3

0.186

Deutonymphs-
male adult

38.0

1.7

23.0

15.4

15.0

0.176

Female adult

37.7

1.3

29.0

16.7

16.8

0.169

64 GREAT BASIN NATURALIST Vol. 32, No. 2

Predator Oviposition Responses on Apple Leaves
Methods. — The oviposition response of T. occidentalis when fed
Tet. mcdanieli was tested on field-collected apple leaves. Leaves with
densities of ca. 60 and 200 prey mites/leaf were selected to insure
that searching for prey was not a factor affecting predator egg pro-
duction. These prey also provided sufficient food to allow the preda-
tors to oviposit for four to six days. Leaves were placed on foam plas-
tic pads in pans of water. Cellucotton strips bordered individual
leaves to discourage predator or prey migrations from the unit. The
initial stage distribution of the prey population was noted for each
leaf and then recounted on successive days during the experiments.
A mean stage distribution value, as the percentage of all preferred
stages (i.e., eggs, larvae, and protonymphs) present during the inter-
action period was estimated from the running percentage means
of the preferred stages present each day. Sufficient female preda-
tors were added to each leaf to insure the destruction of the prey
populations before six days had elapsed. Leaves with lower prey
densities (60 mites/leaf) received 1-3 female predators; those with
higher levels (200 mites/leaf) received 6-14. Oviposition rates per
female/day were calculated, and mean oviposition rate for the en-
tire period was estimated from the daily means, excluding data
from the first day of oviposition and all days after prey density
was lower than predator density. Prior to test termination, preda-
tor populations were largely composed of eggs, larvae, and nymphs
which had a low prey-consumption rate. Searching for prey was
not a significant factor affecting oviposition during the testing
period. Although these experiments were helpful in relating prey
stage distribution to the numerical increase of the predators, the
limitations of a detached leaf and the methods of estimating the
stage distribution values are emphasized.

Results. — When first introduced to a leaf, predators initially
fed on the smaller, more preferred prey stages of Tet. mcdanieli.
Predators consumed over 90 percent of the adult female prey and
oviposition was 44 percent less during the latter half of the test
periods as the more preferred stages were not available. Figure 1
presents a linear regression fit for the relationship between prey
stage distribution and predator oviposition rates for 20 leaf inter-
actions at each density level. A positive regression slope was found
at both density levels, and the sample regression coefficients {b)
for both density treatments were not significantly different. Preda-
tor oviposition rates were high when the mean percent of preferred
prey was high and, conversely, were low when prey populations
had a low proportion of the preferred stages. Estimated oviposition
rates {Y) also were not significantly different between the two prey
density levels.

Feeding Responses of Predators After Periods of Starvation

Several factors may contribute to the differential oviposition
responses of T. occidentalis females to differing stage distributions of

June 1972

CROFT: MITE PREDATION

65

100

80

H 20

2.5 3.0 10 15

EGGS/ DAY/ FEMALE

30

Fig. 1. Regression relationships between the combined percentage of eggs,
larvae, and protonymphs (preferred prey stages) available for food and the ovi-
position rates of Typhlodromus occidentalis for 20 predator-prey interactions on
apple leaves at two prey density levels.

Tet. pacificus and Tet. mcdanieli: Croft and McMurtry (1972a) esti-
mated that T. occidentalis females after feeding on Tet. pacificus eggs
produced five times more eggs than did predators fed on an equiva-
lent weight of adult female prey. On the basis of equivalent number
of prey, predators oviposited only 41 percent as many eggs when fed
eggs as compared to feeding on adult female prey. Unfed larvae were
1.9 times and 16 percent for the same respective parameters. Also,
comparisons of oviposition and prey consumption rates for T. occi-
dentalis fed on prey eggs v. larvae (Croft and McMurtry, 1972a) in-
dicated that an increased feeding response occurred as predators con-
simied about 2.5 times as many larvae as eggs, yet maintained a
similar level of egg production (2.3 eggs per female/day). When
preying on adult female prey, predator oviposition rates were
lower (1.3 eggs per female/day) even though an abundance of food
was provided. Data suggested that eggs and larvae were more ac-
ceptable stages to the predator females than were the larger prey
stages.

Methods. — To test the above hypothesis, standard-sized (2.5
cm in diameter) leaf disks without prey were placed on water-
saturated, foam plastic pads. A uniform starvation schedule prior
to the initiation of the experiments provided female adult preda-
tors in a similar hunger state. These mites were placed singly on
each disk, and at logarithmically-spaced time intervals they were
offered a particular prey stage (eggs, larvae, or female adults).
All tests were made concurrently; 20 predators per replicate and
three replicates (60 total mites) were tested for each stage group at
each time interval. Predators which had been offered a particular

66

GREAT BASIN NATURALIST

Vol. 32, No. 2

prey stage (e.g., eggs only) in previous tests were given a different
type (e.g., larvae or female adult mites) at the next time period to
insure that treatment conditioning did not occur. The percentage of
predators effecting a successful capture after two contacts with a
particular prey stage was recorded at each test period. Direct frontal
approaches to adult female prey nearly always resulted in an
avoidance response by the predators at all starvation times. These
contacts were not counted. After a predator had captured a particu-
lar prey stage and had assumed the feeding position, the prey was
withdrawn. Predators obtained little, if any, food and were further
starved until the next test period.

Results and Discussion. — Figure 2 shows the feeding re-
sponse curves of T. occidentalis females to each prey stage after
varying periods of starvation. Successful capture and acceptance of
prey larvae were extremely high at all starvation periods, reaching
98 percent at 48 hours after the initiation of the tests. Eggs were
slightly less acceptable than larvae at most test periods. Adult female
prey were almost unacceptable during the early periods of the tests
(3-6 hr), increasing to 65 percent successful captures at the most
responsive period (48 hr). In the initial tests (3-6 hr), predators
often made normal feeding approaches to adult female prey, holding

100

80

(/)
m
en

z

2

C/)
111

oc
CD

z

p 60

w 40

C/)

20

T^-

^,1^

^j-^tA

I^

J/

/ \

T /

\

T

Jy' eggs

-'' larvae

I--

female

, adults ,

6 12 24 48

STARVATION TIME (hrs- log scale )

96

Fig. 2. Feeding response curves for Typhlodromus occidentalis to 3 prey
stages of Tetranychus pacificus after varying periods of starvation.

June 1972 croft: mite predation 67

them with their front pair of legs; however, they usually released
the larger mites and did not feed.

During the late stages of starvation (48-96 hr), many predators
ceased searching and remained still until a prey encountered them
or until they died. If a prey larva contacted a starving predator, the
predator would immediately respond and a high percent of successful
captures (95 percent at 96 hr) was effected. Few eggs were en-
countered by predators in this condition. The decline in adult female
consumption at 96 hr resulted from the inability of the starved
predators to overcome and feed on the larger prey stages.

Although these tests were helpful in comparing the hunger
thresholds of T. occidentalis to each prey stage, the interpretation
of these data in terms of field populations should be qualified. Preda-
tors were never 100 percent successful in obtaining prey at any test
period. However, under field conditions, these mites may contact the
prey mites more than twice during each time interval or more than
12 times in 96 hours of searching. We may interpret these data in
terms of field populations by suggesting that when prey densities
are sufficiently high, and a high proportion of the preferred prey
stages are present, predators feed mainly on eggs and larvae. Adult
prey consumption mainly occurs when the density of the preferred
stages is reduced and the hunger level of the predators becomes
high enough to elicit a feeding response on the larger prey stages.

Colonization Pattern and Changes in Prey Distribution

The data discussed previously in this paper have dealt with the
numerical response of T. occidentalis to different stage distribution
of Tet. pacificus and Tet. mcdanieli on a paper substrate and apple
leaf surfaces. Predator-prey interactions at tree and orchard levels
are of greater interest if spider mite control is to be attained. Subse-
quent portions of this paper deal with the effect the previously de-
scribed response on individual leaves may have, in toto, on the larger
sample units in the field.

In southern California, adult females of Tet. mcdanieli overwin-
ter at the base of apple trees and under bark scales in the trunk
region. During April and May, females move up the trunk onto
leaves on water-sprout growth and to leaves on the inner branches
of the scaffold limbs. The initial distribution of prey mites during
this period is typically confined to a low percentage of leaves (1-5
percent) in the lower central portion of the tree. Progeny of the
first generation complete their development on this foliage. At ma-
turity, many of the newly molted females migrate to noninfested
leaves in the upper central portions of the tree and a lateral spread
occurs in subsequent generations. By 1 July, from 20-60 percent of
the leaves in a tree are infested by prey mites (Croft and Barnes,
1971; Croft and McMurtry, 1972b").

The colonization habits of the predator, T. occidentalis are
adapted to those of Tet. mcdanieli. Adult females overwinter in the
debris at the base of trees and under bark scales on the trunk and

68 GREAT BASIN NATURALIST Vol. 32, No. 2

scaffold branches (Hoyt, 1969; Leetham and Jorgensen, 1969). Fe-
male predators move from these hibemacula during the same period
as do the prey mites and initially disperse to the lower, central
region of the tree where the prey population also occurs.

The initial density and distributional relationships between prey
and predator populations vary. On occasion, these relationships
favor the predators, which reduce the initial prey populations to
a low density level (Croft and Barnes, 1972). When no alternate
prey species is present, the early prey reduction results in a starva-
tion decline in predator populations, a later increase in prey levels,
and a second numerical response by the predators.

Occasionally an equilibrium ratio of prey: predators is present,
and a less fluctuating interaction develops with the predator popula-
tion maintaining a reduced density just sufficient to maintain the
prey at a low equilibrium level. The author has observed this con-
dition in naturally developing populations (Croft and Barnes, 1972)
and during minimum release tests (Croft and McMurtry, 1972b).

Most often, the initial numerical and distributional relationships
of predators and prey favor the increase of prey populations. Preda-
tors contact the prey, increase on a limited number of leaves, and
initially have a much lower density and poorer distribution than do
the prey mites. However, some time after prey density has surpassed
a certain minimal level {ca. 2-5 mites/leaf. Croft and McMurtry,
1972b), predator populations numerically respond to the extent of
overcoming and reducing the prey to a low level.

In the latter case, the interaction sequence is generally similar
on individual leaves and in the entire tree. The overwintering fe-
male adult predators disperse to individual leaves in the lower cen-
tral region of the tree and encounter a prey population of a certain
density and stage distribution. Either one or both of these two fac-
tors affect the predator's rate of numerical increase during the
period of interaction on the leaf. The mean rate of predator increase
or decrease on all leaves determines the prey density attained and
the length of time a particular prey level persists in the tree.

If prey density on the leaf is sufficiently high and predator
searching is not a limiting factor, the stage distribution and the ac-
ceptability of these prey to the predators influence the rate of in-
crease on that leaf. The r values in Table 1 give the possible range
of effects this factor might have. One might conclude from mean
tree estimates that the above conditions occur only at high prey
densities; but even at low mean levels (1-5 mites/leaf) the few
leaves with prey have many mites present and the majority are
without prey. If prey density on a leaf is low or zero, searching for
prey is the major factor limiting the rate of numerical increase or
decrease among predators. During this period, the effect of a differ-
ential stage distribution is not greatly expressed. This also applies to
those periods of low prey density on leaves after the predators have
numerically increased and reduced the prey to a low level.

The following interaction sequence in early season was observed
in repeated studies of the numerical response of T. occidentalis to

June 1972 croft: mite predation 69

populations of Tet. mcdanieli on leaves where prey initially were
abundant (Croft and Barnes, 1971, 1972; Croft and McMurtry,
1972b): the immature progeny from eggs oviposited by the over-
winter female predators completed development on the originally
colonized leaf. If prey density was still sufficiently high, a second
generation developed on the leaf. Prior to the complete extincting
of all prey, the original female predators and/or gravid females of
subsequent generations migrated to other infested leaves. Further
prey consumption by developing immature predators most often re-
sulted in the complete extinction of prey from the leaves until later
migrations occurred. At maturity the newly molted female predators
were mated and shortly thereafter dispersed to leaves with prey and
the interaction sequence was repeated. As the predator density in-
creased in the tree, population distribution (percentage of leaves
infested) also increased at a proportional rate (Croft and Barnes,
1971; Croft and McMurtry, 1972b) . Eventually a predator population
numerically increased and was distributed throughout the tree to the
extent of overcoming further prey increases, and control was at-
tained.

Assuming that the above sequence describes a typical numerical
response of T. occidentalis to populations of Tet. mcdanieli in an
individual apple tree, tests were conducted to measure changes in the
stage distribution of the prey mites as related to density and time
upon leaves, in trees, and in an apple orchard. These measurements
were taken to suggest the stage distributions that dispersing predator
populations would encounter in enacting control of the pest.

Prey Infestation on Individual Apple Leaves

As the density and distribution of Tet. mcdanieli initially in-
creased in the tree, a common development was for a single gravid
female prey to disperse to an uninfested leaf. Later, several females
would disperse to uninfested leaves as density further increased
and intraspecific competition intensified. At high density levels,
large numbers of mites would occupy almost all available leaves,
and the remaining uninfested leaves would soon be colonized by
high densities of migrant female mites. These infestations assume
single migrations by various densities of prey mites, but also an in-
finite number of continuous migrations may occur at any time (i.e.,
a single female colonizes a leaf but is joined by one or more addi-
tional females at day 2, day 3, etc., or any other random time).

Methods. — To simulate these infestations, leaf spurs without
mites were collected from a single branch of a "Standard Delicious"
apple tree. All leaves except one of uniform size were removed from
each spur. The spurs were placed in leaf cages and maintained in a
greenhouse under long day photoperiods at 70-90 F, and 50-70 per-
cent R.H. At the initiation of the test, a leaf was infested with either
1, 2, 4, or 16 field-collected prey mites {Tet. mcdanieli, female
adults) of an unknown age. One treatment contained a leaf on
which 1 adult female was added the first and each successive day.

70

GREAT BASIN NATURALIST

Vol. 32, No. 2

Prey density and stage distribution were counted every three days
until test termination. Each of the five treatments was replicated 10
times and the reported data are mean values for all replicates in
each treatment.

Results. — Figure 3 A-F presents the results from the infestation
experiments. The following generalizations were made from these
data: (1) At the lower density levels after colonization had begun,
the percentage of prey in the egg, larval, and protonymphal stages
was extremely high and would be optimally favorable for the
numerical increase of predator populations (Fig. 3 A-E). (2) As
density further increased, the prey population contained a lower
percent of preferred stages (Fig. 3 A-E). (3) If 1, 2, 4, or 16 female
prey mites dispersed to an uninfested leaf and no other mites moved
to that leaf before the first generation was completed, there was a
short period of time (3-6 days in the laboratory) at the end of the
first generation when the stage distribution of the prey was mostly
composed of the large prey stages and would be less favorable for the
numerical increase of the predator (Fig. 3 A-D). (4) The periodic
introduction of additional females to a previously infested leaf tended
to dampen fluctuation (extreme variations in successive sample dates)
in stage distribution on the leaf (Fig. 3E). However, the slope of de-
creasing stage favorability (Fig. 3E) was not markedly different from
the other treatments which supported similar densities, but were

tt 100

00
75

[i\r-,r----

;::„

IV.

^■' .

50

JM^\

D

/ \ ■

F

n

1/ ,v^

[J \

0
10OO

10 20 30

20 30 40 10 20 30

DAYS

Fig. 3 A-F. Prey stage distribution and density changes for individual leaf
infestations of Tetranychus mcdanieli originating fro msingle introductions of 1
(3A), 2 (3B), 4 (3C), 16 (3D female mites, one periodic release treatment of 1
adult femede mite/day (3E) and a combined slope comparison of all treatments
(3F).

June 1972 croft: mite predation 71

started by a single infestation rather than a periodic introduction of
prey mites (Fig. 3 A-D). (5) Depending on the number of prey in-
troduced initially, the period of time that preferred stages would be
available to predators was lessened and the slope of decreasing stage
favorability was steeper as the introduction density of mites was in-
creased (Fig. 3F).

Discussion. — These experiments had implications to the colo-
nization of apple leaves by prey populations under field conditions.
Previous population studies in a T. occidentalis-Tet. mcdanieli sys-
tem (Croft, 1970;* Croft and Barnes, 1972) indicated that most
often the initial contact and subsequent numerical increases of
predator populations were begun in early spring as prey popula-
tions were beginning to increase and total tree densities were low
(<5 prey mites/leaf, but individual leaf densities may be high).
During this period, the stage distribution of the prey would be
optimally favorable for the numerical response of the predator. This
factor undoubtedly contributes to a rapid numerical increase of
predators and the remarkable ability of T. occidentalis to control
Tet. mcdanieli populations during early season (Croft and McMur-
try, 1972b).

Also, during early season, populations of Tet. mcdanieli show
considerable developmental synchronization as generations are some-
what discrete. Samples (total tree estimates) taken prior to the
maturity of the first generation of prey mites will often include
leaves where the proportion of late nymphal newly and molted adult
mites is high (Croft, unpublished data). These conditions of less
favorable stage distribution only persist for short periods in early
spring when temperatures are low.

A comparison of slope differences at each introduction density
(Fig. 3 E) indicated that the period of time in which a highly
favorable stage distribution was present on a leaf, lessened as the
introduction density was increased. Data suggest that as prey in-
creased, the prey stage distribution at the leaf, tree, and orchard
levels would also become less favorable to predator increases.

Seasonal Prey Distribution on Apple Leaves

Methods. — Field samples collected throughout the season were
sampled to test the above hypothesis. Leaves were randomly col-
lected from 5 trees at one-month intervals throughout the grow-
ing season. A sufficient number of leaves were collected at each sam-
ple period to insure that 50 were selected with prey mites present.
Mean total tree densities during the summer increased from 0.88-
162.3 prey /leaf, but individual leaf densities ranged between 1-1000
prey of all stages.

Results. — Figure 4 represents a plot of the combined percent-
age of prey in the egg, larval, and protonymphal stages (preferred

*Croft, B. a. 1970fr. Comparative studies on foiir strains of Typhlodromus occidentalis
Nesbitt (Acarina:Phytoseiidae). Doctoral Dissertation, Univ. Calif., Riverside.

72 GREAT BASIN NATURALIST Vol. 32, No. 2

prey stages) as related to the density of the total prey population on
individual leaves. The broken line in Figure 4 represents the mean
percent of preferred prey stages at increasing density levels. A large
variation in stage distribution was present on individual leaves at
all leaf density levels (Fig. 4). The mean curve declined rapidly as
prey density increased from 0-200 mites/leaf, flattened to an almost
horizontal line from 200-700 prey/leaf, and upturned slightly be-
tween 700-1000 prey/leaf. Although a curve decline occurred at in-
creasing prey densities, the mean percentage of preferred prey re-

100

>

%:.

<

• \

_J

.*• •

^

75

J •;

Q

. •■

>

*. .

n

-. •

z

7

O

w

t?

^'

<

SO

12 3 4 5 6 7 8 9

PREY MITES (hundreds of mites/ leaf )

Fg. 4. A scatter plot of the combined percentage of eggs, larval and proto-
nymphal stages (preferred stages) of Tetranychus mcdanieli as related to the
density of spider mite f>opulations on individual apple leaves at Oak Glen, Cali-
fornia (1970).

mained above 25 percent at all levels and did not attain <10 percent
which would result in the magnitude of r differences as were reported
for eggs and female adult prey in Table 1 . The slight upturn at high
prey densities (Fig. 4, 700-1000 host/leaf) occurred as the highest
prey densities and the more uniform stage distributions developed
just before the plant was overexploited by T. mcdanieli. As leaves
became damaged and were unsuitable food for the spider mites,
their density declined and only the larger mature stages were pres-
ent on the leaves.

June 1972

CROFT: MITE PREDATION

73

Seasonal Prey Distributions in Trees and an Orchard

Methods. — Changes in prey distribution also were measured
throughout the summer of 1970 in the 5 trees selected for the pre-
vious test. Twenty leaves were collected as described by Croft and
Barnes (1971) from each tree at two-week intervals throughout the
season. The number of each stage type on all leaves were counted
at each interval. Density relationships at tree and orchard levels
were also estimated.

Results and Discussion. — Figure 5 presents the combined
percentage of eggs, larvae, and protonymphs (preferred prey stages)
present in two of the trees where fluctuations in stage distribution
were the greatest and a mean measurement for the entire orchard
(5 trees). As expected, data for the individual trees were more vari-
able than was the orchard measurement. Individual tree and orchard
data did not reflect the extreme stage distribution differences that
were observed in the individual leaf samples. This probably was due
to the temporal heterogeneity between individual leaf interaction
within any given tree and the masking effects of lumping individual
leaf counts together. However, individual tree and orchard curves
did show a rapid rise to a peak of host favorability during the initial
phases of colonization, a slow decline during the midseason, and a
rapid decline at the end of the growdng season.

From a biological control standpoint, the most favorable time for
predators to numerically respond and enact control of the prey
would be when prey density had just surpassed the minimum den-
sity required to maintain an increasing predator population (Fig. 5).
Subsequent increasing predator population would encounter less
favorable prey stages and denser populations. Also, Tet. mcdanieli
populations at Oak Glen in 1970 (Fig. 5) attained a lower density
and developed later in the season when compared to the dynamics
of this species in 1968 or 1969. In seasons that prey populations at-
tain high densities earlier in the season, less favorable stage distri-

2 Individual

Trees

5 Trees (

Combined )

\

-stage distribution
.prev density

/

/\""""

,.

/

"\.

J/

1

V

'\

: /

'\

7^,

i

..-'

..--'/

■\-

— ;- - -r

\i

724 821 917 10 1

Fig. 5. The combined percentage of eggs, larvae, and protonymphs (pre-
ferred prey stages) and total population density of Tetranychus mcdanieli in 2
individual apple trees and an apple orchard (mean of 5 trees) at Oak Glen, Cali-
fornia (1970).

74 GREAT BASIN NATURALIST Vol. 32, No. 2

bution probably would be present earlier and over a greater period
of time.

Conclusions

The studies discussed in this paper demonstrate that prey stage
distribution of Tet. mcdanieli and Tet. pacificus is an important
factor influencing the numerical increase of T. occidentalis in small
interaction arenas (apple leaves, 6x6 cm construction paper sub-
strates). The direct effect of this factor at larger interaction levels
(apple trees and orchards) was not obtained. This effect is likely a
complex function of stage distributions, densities, dispersal patterns,
and spatial distributions for both predators and prey and other inter-
acting features of the entire predator-prey-host plant system. How-
ever, measures of prey stage distribution, in the absence of predators,
suggested possible types that T. occidentalis would encounter in en-
acting control of Tet. mcdanieli.

At low and intermediate densities, prey stage distributions were
optimally and generally favorable for the numerical increase of pred-
ators. During short periods early in the season as prey generations
were synchronized and as prey densities became high or overwinter-
ing conditions approached, the stage distribution became less favor-
able for the numerical increase of predators.

It is likely that the prolonged periods of prey stage favorability
contributes to the ability of T. occidentalis to numerically respond
at a rapid rate in early or midseason. However, it is unclear from
these studies if the period of prey stage unfavorability is ever of a
duration or magnitude to have a significant effect on the rate of
numerical response in predator populations. Also, the temporal
asynchrony between leaf interactions may be sufficient to dampen
the effect of these differences on the mean rate of predator increase
at the tree and orchard levels. From an applied biological control
consideration, the effects of unfavorable stage distributions would
most often occur at high densities. Effective control interactions
should have occurred long before these destructive prey levels were
attained. For mass release programs, the most favorable period for
predator release would be shortly after a minimum prey density to
sustain a numerical increase of predators was attained. Not only
would pest density be reduced, but a highly favorable prey stage
distribution would be available to the released predators.

Literature Cited

BuRRELL, R. W.. AND W. J. McCoRMicK, 1958. TyphlodroTTius and Amblyseius
as predators on orchard mites. Ann. Ent. Soc. Amer. 57:483-87.

Chant, D. A. 1959. Phytoseiid mites (Acarina:Phytoseiidae) Part I. Bionomics
of seven species in southwestern England. Part II. A taxonomic review of
the family Phytoseiidae, with descriptions of 38 new species. Canadian Ent.
91:Suppl. 12, 164 p.

— . 1961. The effect of prey density on prey consumption and oviposition

in adults of Typhlodromus (T.) occidentalis in the laborato^>^ Canadian J.
Zool. 39:311-5.

June 1972 croft: mite predation 75

Croft, B. A. 1970. 0>mparative studies on four strains of Typhlodromus
occidentalis Nesbit (AcarinarPhytoseiidae). I. Hybridization and reproduc-
tive isolation studies. Ann. Ent. Soc. Amer. 65:1558-1563.

Croft, B. A., and M. M. Barnes. 1971. Comparative studies of four strains of
Typhlodromus occidentalis. III. Evaluations of releases of insecticide resis-
tant strains into an apple orchard ecosystrm. J. Econ. Ent. 64:845-850.

. 1972. Comparative studies on four strains of Typhlodromus occidentalis.

VI. Persistence of insecticide: resistant strains in an apple orchard eco-
system. Ibid. 65:211-216.

Croft, B. A., and C. D. Jorgensen. 1969. Life history of Typhlodromus mc-
gregori (Acarina:Phytoseiidae) . Ann. Ent. Soc. Amer. 62(6) : 1261 -1267.

Croft, B. A., and J. A. McMurtry. 1972a. Comparative studies on four strains
of Typhlodromus occidentalis Nesbitt (AcarinarPhytoseiidae). IV. Life his-
tory studies. Acarologia (In press).

. 1972b. Minimum releases of Typhlodromus occidentalis to control Te-

tranychus mcdanieli on apple. J. Econ. Entomol. 65:188-191.

HoYT, S. C. 1969. Population studies of five mite species on apple in Washing-
ton. Proc. 2nd Intern. Cong. Acri: 1 1 7-1 33.

Leetham, J. W., AND C. D. Jorgensen. 1969. Overwintering phytoseiid mites
in central Utah apple orchards. Great Basin Nat. 29:96-104.

iPuTMAN, W. L. 1962. Life history and behavior of the predaceous mite
Typhlodromus (T.) caudiglans Schuster (AcarinarPhytoseiidae) in Ontario
with notes on the prey of related species. Canadian Ent. 94:163-177.

Scriven, G. T., and J. A. McMurtry. 1971. Quantitative production and pro-
cessing of tetranychid mites for large scale testing or predator production.
Jour. Econ. Ent. 64:1254-1258.

VAN de Vrie, M., and a. Boersma. 1970. The influence of the predaceous mite
Typhlodromus {A.) potentillae (Garman) on the development of Panony-
chus ulmi (Koch) on apple grown under various nitrogen conditions. En-
tomophaga 15:291-304.

A REVISION OF THE
PSORALEA LANCEOLATA COMPLEX (LEGUMINOSAE)^

C. A. Toft^ and S. L. Welsh^

Abstract. — The Psoralea lanceolata complex is widely distributed in western
North America from Saskatchewan to Texas and westward to Washington, Oregon,
Nevada, and Arizona. The complex has received many interpretations, mostly on
the basis of provincial floras and manuals. The present paper attempts to sum-
marize the treatments of previous workers and to present an overview of the
complex. The Psoralea lanceolata var. stenophylla is proposed. Distribution maps
and illustrations of all entities are included.

The lemon scurf-pea or lemon-weed, Psoralea lanceolata, is a
plant of arid regions and of sandy soils. It often invades disturbed
sites and is an important pioneer on sand dunes, making it a signifi-
cant plant in erosion control (Hopkins, 1951; Huelett and Coupland,
1966). The P. lanceolata complex is notable for its intraspecific
variability (Isely, 1962; Matthews, 1969); indeed the variation with-
in the complex has never been well understood or well documented.
The treatment by Rydberg (1919) is the most recent work on
Psoralea of North America, and his proliferation of species within
this complex attests to its wide variation and to the inadequate under-
standing of that variation. More recent treatments of P. lanceolata
have been in the form of regional floras (Boivin, 1967; Harrington,
1954; Hitchcock et al., 1966; Kearney and Peebles, 1960; Munz and
Keck, 1959), which in the large part do not have to deal with the
overall variation in the lanceolata complex. In any case, they are
mostly only reiterations of Rydberg's original key, modified very
little if at all. The most comprehensive and most recent treatment
of P. lanceolata is that by Isely (1962). His treatment of P. lanceo-
lata is not complete because of its regional nature, and the population
of the north central states varies far less than the more western con-
geners of P. lanceolata.

Indications that the knowledge of variation within the lanceolata
complex is inadequate are provided by specimens which do not fit
well into the established concept of P. lanceolata. These specimens
have been (ollected by a number of people at unrelated times and
places and were consequently housed at various herbaria. It is the
purpose of this study to examine in detail the variation within the
P. lanceolata complex and to elucidate a current concept of this
grouj). From the data gained in this study, it has been necessary to
modify the key to embrace the actual rather than the purported vari-
ation within the group and to propose a new combination in recogni-
tion of a distinct infraspecific population of P. lanceolata.

'Study supported by NSF and undergraduate research funds. Grant No. GY9052.
'Department of ZooloRy and F.ntomolopy, Iowa State University, Ames, Iowa 50010.
"Department of Botany and Range Science, Brigham Young University, Provo, Utah 84601.

76

June 1972 toft, welsh : psoralea 11

Methods

Herbarium specimens of P. lanceolata from the herbaria of
Brigham Young University, Utah State University, University of
Utah, and Iowa State University were examined Acknowledgments
are hereby made to the curators of the herbaria of these institutions.
Various measurements and data were recorded from the specimens
in an attempt to enumerate evident variation within the complex.
The following data were recorded: (1) measurements of the length
and width of the leaflets and calculation from the measurements of
the ratios and the means of the ratios for various populations; (2)
measurements of raceme and peduncle length and number of flower
nodes per raceme and calculation of the ratios of the measurements
to each other; (3) flower color; (4) the condition of pubescence of
the pod; (5) time of flowering; (6) collection locality of each speci-
men; (7) chemotaxonomy of the alkaloids, flavanoids, and phenols
of selected populations.

In addition, opportunity was taken to observe populations in the
field, because the authors acknowledge the limitations of herbarium
specimens in this type study. Collections for herbarium specimens
are usually of only one plant or occasionally a small number of
plants, and often these are not of the average type but rather of the
most robust or most unusual specimen. Nearly the opposite condi-
tions are required for an objective study of variation, specifically:
large, random, and unbiased samples. However, the herbarium speci-
mens are still the most important tool, given that their limitations
are taken into account and if accurate field observations are used to
supplement them.

The composite measurements were evaluated and attempts to
correlate the above characters were made using several techniques
including polygonal graphs, histograms, scatter diagrams, and range
maps. In addition to the evaluation of the statistics of each plant,
the authors beforehand separated the herbarium sheets into various
piles on a purely visual basis, using "intuition" and accumulated
experience with the group. Later, these segregates could be compared
with the evaluated data, and in this way not only provide another
means of correlation but also help to reduce any bias which might
occur.

Taxonomy

Psoralea lanceolata Pursh Fl. 1814. Amer. Sept. 475.

Erect perennial from creeping branched rootstocks; stems 1-6
dm high (usually 2-3 dm), single or clustered, diffusely branched
and often forming bushy clumps, inconspicuously strigose, glandular-
punctate, aromatic; leaves mostly palmately 3-foliate; leaflets 1.5-4
cm long, the basal ones obovate to oblanceolate, narrower upward,
the uppermost linear or remaining oblanceolate, acute to rounded or
mucronate at apex, rounded to cuneate at base, strongly glandular-
punctate, petioles 1-2.5 cm long; stipules linear-lanceolate to subu-
late, 3-10 mm long; peduncles 2-24 cm long, from barely exceeding

78

GREAT BASIN NATURALIST

Vol. 32, No. 2

40

30

Per cent

of

sample

20

10

-n — ' — I — ' — I

1-14 15-28 1-14 15-28 1-14 1528 1-14 15-28 1-14 15-28
My Jn Jl Ag Sp

Date of flowering

Hg. 1. Distribution of flowering dates of Psoralea lanceolata var. steno-
phylla.

the leaves to nearly as tall as the plant; racemes 0 5-17 cm long,
from compact and short to lax and elongate; bracts minute, early
deciduous; calyx campanulate, inconspicuously strigose, 2 mm long,
not enlarging in fruit, teeth nearly equal, shorter than the tube,
obtuse, glandular-punctate; corolla 3-7 mm long, whitish with
purple-tipped keel or entire corolla dark blue to violet, banner almost
orbicular, but with distinct auricles, blades of the wings obliquely
oblong-oblanceolate, keel-})etals scarcely lobed at base; stigma capi-
tate, fruit exposed, subglobose when mature, with short beak, 4-6

June 1972

TOFT, WELSH: PSORALEA

79

40

30

Per cent

of
sample

20

10

1-14 15-28 1-14 15-28 1-14 15-28 1-14 15-28 1-14 15-28
My Jn Jl Ag Sp

Date of flowering

Fig. 2. Distribution of flowering dates of Psoralea lanceolata var. lanceo-
lata within same geographical area as var. stenophylla.

mm in diameter, conspicuously glandular-warty, sparingly strigose
to densely villous, pericarp rigid, not breaking at maturity.

Distribution and Habitat. — Saskatchewan, North Dakota,
South Dakota, south to Texas, west to Washington, Oregon, Cali-
fornia, Arizona. This species occupies a variety of habitats, almost
all of which show signs of disturbance. In the eastern portion of its
range, it may occupy native prairie (Isely, 1962). In arid regions,
it is often found on sand dunes and in other sandy areas. However,
where moisture is readily available, it may become weedy and ag-

80 GREAT BASIN NATURALIST Vol. 32, No. 2

gressive. It is often found on roadsides, bottomlands, and other dis-
turbed areas (Isely, 1962).

Discussion. — In addition to the well-documented variation in
leaflet width (Isely, 1962), the present study has indicated two sig-
nificant types of phenotypic variation in the complex that were
previously overlooked. They are flower color and raceme length.
No flora or treatise which the authors have studied has acknowledged
that P. lanceolata may have dark blue or violet flowers. Indeed, in
regions where P. lanceolata and P. tenuiflora occur together, the
regional keys are written in such a way that flower color separates
the two species (Isely, 1962; Rydberg, 1919; Tidestrom and Kittell,
1941), with P. lanceolata having white flowers with purple-tipped
keels and P. tenuiflora having dark blue or violet flowers However,
in the populations west of the Rocky Mountains, the blue flowered
form may be far more prevalent than the white. In P. lanceolata
var. purshii. the blue flowered individuals seem to be more common
over most of its range. Also, in some small pockets of the Great
Basin P. lanceolata, the two color phases grow side by side. This
condition occurs in the P. lanceolata population of the white sand
dunes of Juab County, Utah. In this isolated population, the speci-
mens of P. lanceolata, which grow densely in the interdune areas,
are mostly blue flowered; however, a white flowered form occurs on
the edge of a blue flowered population. A similar situation occurs
in Garfield County, Utah, and probably in other isolated pockets
as well.

The variable raceme was recognized by Rydberg who named
two species from the lanceolata complex on the basis of the lax and
elongate raceme, P. stenostachys and P. stenophylla, respectively.
However, subsequent authors have ignored raceme variability. In
the present study raceme variation was investigated and found to be
significant.

For the most part the total inflorescence length is quite variable;
however, the flowers are almost always densely clustered on the
raceme (Fig. 3). An exception to this was found in the population
in southern and eastern Utah. In Utah there are two distinct forms
of lanceolata, one which is typical with a dense raceme and one
which has a very lax raceme (Fig. 3). The authors believe that this
latter form is distinct and a separate entity from typical lanceolata
for the following reasons: (1) It occurs only in a restricted geo-
graphic location, whereas typical lanceolata occurs uniformly over
a much wider area (Figs. 3, 4). (2) The authors have never ob-
served the phase with the lax raceme growing in the same habitat
with typical P. lanceolata, c\e\\ though the two types are apparently
sympatric when plotted on distribution maps. It appears, therefore,
that there is segregation of the phases on an ecological basis. (3) The
phase with the lax raceme appears by all evidence to bloom at an
earlier date than the tvpical P. lanceolata in that area (Figs. 1 and 2
summarize collection dates taken from specimens in flower on her-
barium sheets for all plants collected in the area where the two forms
are sympatric, i.e., Kane. Garfield, San Juan, Wayne, and Grand

June 1972

TOFT, WELSH: PSORALEA

81

Fig. 3. Summary of variation within the Psoralen lanceolata complex. A.
var. purshii. Flowers (predominantly blue), racemes compact, pods densely vil-
lous, leaflets broad, uniform. B. var. purshii - var. lanceolata intermediates. Pods
strigose, leaflets intermediate. C. var. lanceolata. Flowers predominantly white,
racemes compact, pods sparingly strigose, leaflets narrow, variable. D. var. steno-
phylla. Flowers predominantly blue, racemes elongate, pods sparingly strigose,
leaflets narrow, variable. E. var. lanceolata, western phase. Flowers commonly
blue, racemes compact, pods sparingly strigose, leaflets narrow, variable.

counties of Utah). (4) The entity with lax racemes is morpho-
logically well separated from typical P. lanceolata, and intermediate
forms have not been observed.

The phase with the elongate raceme is the same entity which
was described by Rydberg as Psoralea stenophylla. Thus, the correct
name for this entity is Psoralea lanceolata var. stenophylla (Rydb.)
Toft and Welsh comb, nov., based on Psoralea stenophylla Bull.
Torr. Club 40:46. 1913.

A brief investigation of the alkaloid, flavinoid, and phenol
chemotaxonomy was made. A solvent system of butanol-acetic acid-
water (4:1:5) was used, and spots were detected by Dragendorf's
reagent for the alkaloids and by ultraviolet light for the flavinoids
and phenols. It was found that the infraspecific populations of
P. lanceolata did not differ significantly in any of these. However,
a spot detected by Dragendorf's reagent, presumably an alkaloid.

82

GREAT BASIN NATURALIST Vol. 32, No. 2

FIG. 4 ^r^-^

/ * • /
/ * /

M /

•
•

~1 I"

•

— 'L

<!

• •

V

V 1 •• • ^

r

•

Fig. 4. Range of Psoralea lanceolata Pursh and P. juncea Eastw.
P. lanceolata var. lanceolata and var. purshii (solid dot).
P. lanceolata var. stenophylla (star).
P. juncea (asterisk).

was present in /^. lanceolata samples at Rf = 0.4 and was missing
for samples of P. juncea. Unfortunately, not all varieties of P.
lanceolata could be investigated to determine if this spot had any
taxonomic significance.

A key which embraces the variation in this study is proposed in
order to distinguish Psoralea tenuiflora and P. juncea from P. lanceo-
lata which may be readily confused with the former two in taxo-
nomic treatments.

Plants dropping all but a few basal leaves by flowering
time; raceme extremely elongate, almost as tall as
plant; flowers usually dark blue; southeastern Utah
and northern Arizona juncea Eastw.

Plants retaining leaves at flowering time; racemes
variable, not usually as tall as plant; flower color vari-
able, white or dark blue; distribution various 2

2.(1 ) Fruits ovoid-elliptical; racemes lax, usually three flow-
ers to a node; corolla dark blue; leaflets nearly uni-
form in size from base to top of plant, narrowly to
broadly obovate to oblong tenuiflora Pursh

June 1972 toft, welsh : psoralea 83

2/ Fruits subglobose; racemes variable, from densely clus-
tered to lax, usually two flowers to a node; corolla vari-
able, either white with a purple keel or entire corolla
blue to violet (latter, common west of Rockies) ; leaflets
variable from top to bottom on same plant, basal ones
obovate to oblanceolate, those near the top of the plant
narrowly linear lanceolata Pursh

Key to the varieties of Psoralea lanceolata Pursh

1. Leaflets obovate at base to oblanceolate at top of plant;
leaflet variation from base to top of plant only slight;
fruits densely villose or, less commonly, sparingly stri-
gose; plants of Idaho, Nevada, Washington, Oregon
var. purshii

1/ Leaflets oblanceolate to linear at base, linear to nar-
rowly linear at top; leaflet variation from base to top
may be pronounced; fruits slightly strigose 2

2. ( 1 ) Racemes dense, flowers clustered closely; flowers white

with purple keel (common in eastern portion of range)
or entire flower dark blue to violet; typical lanceolata
with wide range from Saskatchewan, North Dakota,
South Dakota, south to Texas, and west to Utah and
Arizona var. lanceolata

2/ Racemes lax, the flower nodes widely separated, the
inflorescence extremely elongate; blue flowers pre-
dominating; known only from arid regions of southern
and eastern Utah var. stenophylla

Psoralea lanceolata var. purshii (Vail) Piper, Piper & Beattie. 1901.
Fl. Palouse Reg. 106.

Lotodes ellipticum var. latifolium Kuntze. 1891. Rev. Gen. 1:193.

Psoralea purshii Vail. 1894. Bull, Torr. Club 21:94.

Psoralea scabra Nutt. 1838. In T. and G. Fl. N. Amer. 1:300.

Psoralea lanceolata Pursh ssp. scabra (Nutt. T. & G.) Piper. 1906. Contr.

U.S. Nat. Herb. 11:364.
Psoralidium purshii (Vail) Rydb. 1919. N. Amer. Fl. 24:14.

Description. — Differing from typical P lanceolata in having
broader leaflets, basal leaflets obovate, narrower upward, the upper-
most oblanceolate, leaflet variation from top to base of plant only
slight; racemes dense, flowers predominantly dark blue to violet,
less commonly white with purple keel; fruits densely villose, less
commonly, sparingly strigose.

Distribution and Habitat. — Idaho, Nevada, Washington,
Oregon. Sandy areas often associated with Artemisia (Munz and
Keck, 1959; Tidestrom, 1925).

Flowering Time. — May to July.

Discussion. — Data gained from leaflet measurement in this
study reconfirm the previously recognized var. purshii (Isely, 1962;

84 GREAT BASIN NATURALIST Vol. 32, No. 2

Munz and Keck, 1959; Tidestrom, 1925) in the northwestern part
of the range of the complex, a variety which has broader leaflets
and more uniform leaflets from the top to the base of the plant than
in var. lanceolata. In a restricted area in northern Utah and south-
eastern Idaho var. purshii and var. lanceolata appear to be inter-
grading (Fig. 3).

Representative Specimens. — California. Mono Co.: Reveal
526 (uTc); Idaho. Bingham Co.: Cronquist 2312 (utc); Davis 1508
(uTc); Holmgren s.n. (utc). Butte Co.; Atwood 913 (bry); Atwood
1167 (bry). Elmore Co.: Maguire and Holmgren s.n. (utc). Fre-
mont Co.: Maguire 17178 (utc); Raven and Gregory 19562 (isc).
Jefferson Co.: Anderson 302 (utc); Atwood 996 (bry); Bench s.n.
(bry). Owyhee Co.: Davis 854 (utc); Maguire and Holmgren
26264 (utc). Nevada. Elko Co.: Holmgren 517 (bry, utc). Hum-
bolt Co.: Constance, Molseed, and Ornduff 3695 (bry, utc); Gentry
and Davidse 1539 (bry, utc, isc); Holmgren and Maguire 10605
(utc); Maguire and Holmgren 22481 (utc); Moran s.n. (bry).
Oregon. Morrow Co.: Cronquist s.n. (utc) ; Sherman Co.: Constance
and Beetle 2687 (utc). Washington. Grant Co.: Moran s.n. (bry).
Kittitas Co.: Maguire 17239 (utc); Klickitat Co.: Suksdorf s.n.
(isc).

Psoralea lanceolata var. lanceolata

P. lanceolata Pursh. 1814. Fl. Amer. Sept. 475.

P. elliptica Pursh. 1814. Fl. Amer. Sept. 741.

P. arenaria Nutt. 1818. Gen. 2:103.

P. laxiflora Nutt. 1838. In T. and G. Fl. N. Amer. 1:299.

P. micrantha Gray. 1857. In Torr. Pacif. R.R. Rep. 4:77.

Lotodes elUpticum (Pursh) Kuntze var. angustissiurn Kuntze. 1891. Rev.

Gen. 1:193.
Lotodes micrantha (Gray) Kuntze. 1891. Rev. G«n. 194.
P. stenostachys Rydb. 1913. Bull. Torr. Club 40:46.
Psoralidium lanceolatum (Pursh) Rydb. 1919. N. Amer. Fl. 24:13.
Psoralidium rnicranthum (Gray) Rydb. 1919. N. Amer. Fl. 24:14.
Psoralidium stenostachys Rydb. 1919. N. Amer. Fl. 24:14.

Description. — This is the typical and most widespread form of
P. lanceolata. Leaflets oblanceolate to linear at base, narrower up-
ward; leaflet variation from base to to[) may be pronoimced; racemes
dense, flowers closely clustered or less commonly lax; corolla white
with purple-tipped keel (common in eastern portion of range) or
entire corolla dark blue to violet (common in western portion of
range); fruit sparingly strigose.

Distribution. — Saskatchewan, North Dakota, South Dakota,
south to Texas, and west to Utah and Arizona.

Representative Specimens. — Arizona. Coconino Co.: Carter
1599 (utc); MacDougal 217 (isc); Toft 144 (bry). Mohave Co.:
Deaver 6251 (bry). Colorado. Brown 120 (bry); Alamosa Co.:
Isely 8184 (isc); Ramaley and Johnson 14994 (bry). Denver Co.:
Eetterman s.n. (isc). Douglas Co.: Osterhout and Clokey 3803 (utc,
isc). Elbert Co.: Owneby 1369 (utc). EI Paso Co.: Jones 142 (utc).
Morgan Co.: Isely 6488 (isc). Weld Co.: Welsh, Moore, and Mat-

June 1972 toft, welsh: psoralea 85

thews 9320 (bry). Kansas. Fairchild s.n. 1888 (isc); Hurr E287
(isc). Edwards Co.: Welsh 665 (bry, isc). Osborne Co.: Shear 114
(uTc). Montana. Dawson Co.: Aeton s.n. (isc). Nebraska. Banner
Co.: Welsh 1067 (isc). Box Butte Co.: Churchill s.n. (isc); Welsh
1077 (bry, isc). Cherry Co.: Tolstead 4-470 (isc); Tolstead 380
(isc). Dawes Co.: Tolstead s.n. (isc). Holt Co.: Burzlaff s.n. (utc).
Kearney Co.: Hapeman s.n. (utc). Lincoln Co.: Brown 53-6 (isc).
Morril Co.: Maguire s.n. (bry, utc). Rock Co.: Clements 2874
(isc). Sioux Co.: Rittenhouse 206 (utc). Sheridan Co.: Buchanan
s.n. (isc). Thomas Co.: Isely 6070 (isc). New Mexico. Colfax Co.:
Foster s.n. (isc). Rio Arriba Co.: Flowers and Hall 130 (bry). Okla-
homa. Cimarron Co.: Jespersen 2715 (utc). Logan Co.: Carleton
151 (utc). Texas Co.: Isely 6394 (bry, isc). Woods Co.: Stratton
1326 (isc), 6383 (isc). Saskatchewan. Ledingham and Yip 2225
(isc). South Dakota. Badlands Natl. Mon.: Lindstrom 329 (bry).
Butte Co.: Brown 52-22 (isc). Clay Co.: Van Bruggen 5604 (bry).
Todd Co.: Isely 6058 (isc); Tolstead 4-30 (isc); Welsh 1095 (isc).
Utah. Daggett Co.: Smith s.n. (utc); Smith s.n. (utc). Emery Co.:
Welsh and Atwood 9841 (bry, isc). Garfield Co.: Atwood 459 (bry);
Coles 123 (bry); Collotzi, John, and Atwood 512 (utc); Holmgren,
Reveal, and La France 2044 (bry, utc); Stanton 335 (bry, ut);
Woodruff 1179 (bry). Grand Co.: Jotter 2098 (ut); Rydberg and
Garrett s.n. (ut); Welsh and Atwood 9949 (bry, isc); Welsh and
Moore 2027 (bry). Juab Co.: Harrison 11350 (bry); Harrison 6550
(bry, isc); Harrison 354H (bry, isc); Harrison 11777 (bry, isc,
utc); Smigelski 63 (bry); Smigelski 67 (bry); Toft 17 (bry); Toft
16 (bry); Toft 148 (bry); Toft 149 (bry); Welsh and Moore 5128
(bry). Kane Co.: Barnum 1324 (bry); Castle 170C (bry); Cottam
2712 (bry); Harrison 11077 (bry); Holmgren and Nelson 7191
(utc); Maguire 12292 (bry, utc); Maguire 18893 (utc); Toft 135
(bry); Toft 136 (bry); Toft 137 (bry); Toft 139 (bry); Toft 142
(bry); Toft 143 (bry); Welsh 1696 (bry, isc); Welsh and Welsh
9431 (bry). Millard Co.: Cottam 1006 (bry); Cottam 3782 (bry,
ut); Garrett 3937 (bry, ut). Salt Lake Co.: Dunn 1940 (bry);
Harrison 10586 (bry, ut, utc); Howard s.n. (ut); McKnight 10586
(isc). San Juan Co.: Cottam 5962 (ut); Welsh, Moore, and Canter
2927 (bry); Welsh, Moore, and Canter 3010 (bry, isc). Tooele Co.:
Flowers 898 (ut); Flowers 8 (bry, ut). Uintah Co.: Jones s.n.
(utc); Welsh 446 (bry). Washington Co.: Pendleton s.n. (ut);
Weight 9192 (ut); Weight 10903 (ut). Wayne Co.: Beck and
McArthur 150 (bry); Cottam 4476 (bry, ut); Holmgren, Boyle,
and Will 7792 (utc); Maguire 19251 (utc); Markham s.n. (bry);
Welsh and Atwood 9850 (bry, isc); Welsh and Moore 3595 (bry).
Weber Co.: Brizzee 7814 (ut); Flowers 1277 (ut); Garrett 6289
(bry, ut). Wyoming. Crook Co.: Porter and Porter 7574 (utc).
Carbon Co.: Porter 3725 (utc). Natrona Co.: Gooding 177 (utc,
isc); Isely 6535 (isc); Welsh, Moore, and Matthews 9217 (bry).
Niobrara Co.: Isely 3948 (bry, isc). Park Co.: Greever 32 (bry).
Psoralea lanceolata var. stenophylla (Rydb.) Toft and Walsh Comb,
nov.

86 GREAT BASIN NATURALIST Vol. 32, No. 2

Psoralea stenophylla Rvdb. 1913. Bull. Torr. Club 40:46.
Psoralidium stenophyllum Rydb. 1919. N. Amer. Fl. 24:14.

Description. — Differing from typical lanceolata in having elon-
gate and lax racemes, flower nodes widely separated; flowers most
commonly dark blue to violet; leaflets oblanceolate to linear at base,
narrower upward, leaflet variation from base to top may be pro-
nounced; fruits sparingly strigose.

Distribution and Habitat. — Known only from arid regions of
southern and eastern Utah, i.e., Emery, Wayne, Garfield, Kane,
Grand, and San Juan counties. This form is almost always found
growing in sandy soils: on roadsides, in canyon and wash bottoms,
talus slopes, and sandstone cliffs. It has been reported to be associ-
ated with Amelanchier, Chrysothamnus, Coleogyne, Artemisia,
Pinus, Juniperus, Quercus, Phragmites, Rhamnus. and Vanclevia.

Flowering Time. — This form appears to bloom at an earlier
date than typical lanceolata (Figs. 1,2), beginning in early May and
tapering off by mid-June.

Discussion. — Records for P. lanceloata var- stenophylla and
those for P. juncea, an endemic plant of southeastern Utah and
northern Arizona are largely sympatric (Fig. 4). This range corre-
lation and a similarity in morphology, where var. stenophylla ap-
pears to be intermediate in raceme length between P. juncea and
P. lanceolata var. lanceolata, suggest the possibility that var. steno-
phylla may be the product of hybridization between these two. How-
ever, this assertion cannot safely be made until more detailed study
is made and actual hybridization attempted; thus, the authors are
merely posing the question here as an impetus to further study.

Representative Specimens. — Utah. Garfield Co.: Holmgren
and Goddard 9955 (ut); Kaneko 28 (bry); Welsh and Moore 7110
(bry, isc). Grand Co.: Harrison 11367 (bry). Kane Co.: Beck and
Tanner s.n. (bry); Harrison 12114 (bry); Murdock 371 (bry);
Welsh 1689 (bry). San Juan Co.: Welsh 5387 (bry, isc); Wilson
120 (uTc). Wayne Co.: Beck s.n. (bry, isc); Cottam 9275 (ut);
Welsh and Atwood 9879 (bry, isc).

References

BoiviN, B. 1967. Flora of the prairie provinces: In Provancheria. II. Memoires
de I'Herbier Louis-Marie Faculte d'Agriculture, part I, Universite Laval.

CoRRELL, D. S., AND M. C. JoHNSON. 1970. Manual of the vascular plants of
Texas. Texas Research Foundation, Renner, Texas.

Hitchcock, C. L., A. Cronquist, M. Ownbey, J. W. Thompson. 1961. Vascu-
lar plants of the Pacific Northwest 3:349. Seattle, University of Washing-
ton Press.

Hopkins, H. H. 1951. Ecology of the native vegetation of the loess hills in
central Nebraska. Ecol. Monogr. 21 (2): 125-147.

Huelett, G. K., and R. T. Coupland. 1966. The vegetation of dune sand
areas within the grassland region of Saskatchewan. Can. Jour. Bot. 44(10):
1307-1331.

Isely, D. 1962. The Leguminosae of the north-central United States. IV.
Psoraleae. Iowa State Jour. Sci. 37:103-162.

June 1972 toft, welsh : psoralea 87

Ke.\rney. T. H.. and R. H. Peebles. 1951. Leguminosae. In: Arizona flora.

University of California Press, Berkeley.
Matthews, E. 1969. A preliminary revision of the genus Psoralea in Utah.

Brigham Young University, unpublished manuscript. 47 p.
MuNz, P. A., AND D. D. Keck. 1959. Leguminosae. In: A California flora.

University of California Press, Berkeley.
Rydberg, p. a. 1919. Fabaceae. In: North American Flora. 24(1): 12-17.
Rydberg, p. a. 1922. Fabaceae. In: Flora of the Rocky Mountains and ad-
jacent plains. New York.
Tidestrom, I. 1925. Flora of Utah and Nevada. Contr. U.S. Nat. Herb. 25:

304-305.
Tidestrom, I., and T. Kittell. 1941. Flora of Arizona and New Mexico.

Edwards Bros. Inc., Ann Arbor.
ToRREY, J., AND A. Gray. 1838. Flora of North America. Vol. 1. Wiley and

Putnam, New York.

OBSERVATIONS ON THE NESTING BIOLOGY
OF THE LONG-BILLED CURLEW

Dennis M. Forsythei

Abstract. — Observations were made on four nesting attempts by Long-billed
Curlews {Numenius americanus) in 1965 and 1966 in Box Elder and Cache
Counties, Utah. The location, nest composition, and number of eggs of each nest
are presented. In one instance eggs were laid at intervals of forty-eight hours or
less. Data on weight and measurement and color of eggs are given. Data for the
only two completed nests indicates incubation periods of twenty-seven and twenty-
eight days respectively. The distraction display of the incubatng family is de-
scrbed for the first time.

Both Palmer (1967) and Graul (1971) emphasized the lack of
infoimation on the breeding biology of the long-billed curlew {Nu-
menius americanus). For this reason the following nesting data
are presented on four pairs of curlews gathered during the 1965 and
1966 breeding seasons in Box Elder County and Cache County, Utah.

On 15 May 1965, Jack Andersen discovered a curlew nest in a
grass pasture, elevation 4500 ft, situated on a bench representing the
shoreline of old Lake Bonneville, two miles N of Mendon, Cache
County. The nest consisted of a grass-lined depression with an inside
diameter of 190 mm, which was located in a clump of grass (Gra-
mineae) and sedge (Cyperaceae). When found, it contained two eggs
whose color and pattern were as described by Reed (1965:125) and
Bent (1929:101). At 17:45 on 17 May, I visited the nest and found
three eggs which I color-marked. Between 17 May and 23 May,
when the nest was destroyed by a farm tractor, no additional eggs
were laid. Palmer (1967: 183-184) gives a clutch-size of four for this
species. The above data indicate the greatest interval possible be-
tween the addition of the second and third egg was 48 hours which
is somewhat greater than Graul's (1971:193) observation. Also the
eggs had been rotated in the nest at least once between 17 May and
22 May. On three daytime visits out of five I found a bird of unde-
termined sex incubating. On 17 May when I flushed a bird from
the nest, it silently flew low over the ground with its tail spread,
both wing tips touching the ground and neck perpendicular to the
ground. No other curlews were seen in the vicinity of this nest dur-
ing any visit.

K. L. Shirly found a second nest in the salt flats, elevation 4000
ft, adjacent to the Bear River, eight miles W of Brigham City, Box
Elder County, at 0600 on 1 May 1966. The nest cup, constructed of
desert saltgrass (Distichlis stricta), was located in a clump of dead
saltgrass and samphire {Salicornia rubra). The nest contained three
eggs of normal color and pattern. On 7 May at 0600, K. L. Shirly
and I visited the nest and found the female incubating. This species
is noticeably sexually dimorphic in size, especially the length of the

'Department of Biology, The Citadel, Charleston, South Carolina 29409.

88

June 1972 forsythe: long-billed curlew 89

exposed culmen (unpublished data). When we approached the cur-
lew, it flushed from the nest in a manner similar to that described
above. When 50 meters from the nest, the curlew stopped, then
dragged one wing on the ground, then the other, then both. On
occasion it would jump off the ground, then run along the ground
with neck outstretched and bill touching the ground. This behavior
lasted about one minute until the male appeared and circled over
our heads giving the Ki-keck call (Forsythe 1970:215). The female
answered with the Curluoo call (Forsythe 1970:215). They were
joined by two males giving the Ki-kcck call. The four birds pro-
ceeded in mobbing behavior as described by LaFave (1954:48). This
lasted until we left the area 15 minutes later. When I visited the nest
on 24 May, it had been destroyed by unknown causes leaving no
trace of nest or eggs. No adult birds were observed in the vicinity.

I found a third nest of this species at 1600 on 3 May 1966 in an
irrigated grass pasture, 4000 ft elevation, 3.3 miles W of Logan Post
Office, Cache County. The nest, containing four eggs, was composed
of various grasses and situated on a slight rise in the field. Both
sexes incubated but only the female gave diversionary displays when
flushed from the nest. These displays were similar to those already
described except the bird looked back at the observer with its neck
perpendicular to the ground. The eggs hatched between 0700 on 21
May and 1600 on 23 May. If we assume that they hatched at the
earlier time and that eggs were laid at two-day intervals, the incuba-
tion period would be around 28 days.

K. L. Shirly and I found the last nest at 0700 on 7 May 1966
in the salt flats next to the Bear River, 10 miles W of Brigham City,
Box Elder County. The location and placement of this nest were
similar to those described for the 1 May nest. The nest contained four
eggs. On 24 May at 0600 as I got out of the car about 150 meters
south of the nest, the male was sitting on a fence post 30 meters from
me. As I approached the nest, the male gave several Curuloo calls.
The female, neck outstretched with body low and bill parallel to the
ground, walked slowly off the nest toward the north. When I ap-
proached, the female flushed and gave the diversionary display with
both wings. Then the male along with several other curlews mobbed
me giving the Ki-keck and Arc Display calls (Forsythe 1970:214).
I returned to the nest and found one egg pipped. I marked the eggs
and removed them to the laboratory where they were weighed and
measured (Table 1). The eggs were a darker green than described
by Reed (1965:125). The first egg hatched at 1300, the second at
2025 on 25 May, and the third at 0625 on 26 May. The fourth egg
was preserved before hatching and deposited in the vertebrate col-
lection of Utah State University. The egg measurements were simi-
lar to those given by Bent (1929:101). All eggs decreased in weight
during hatching (Table 1). Other aspects of the hatching process
have been described by Forsythe (1967:340). Using a two-day
interval between the laying of each egg, and assuming that the
fourth egg would have hatched on 26 May, gives an incubation
period for this nest of 27 days.

90 GREAT BASIN NATURALIST Vol. 32, No. 2

Table 1. Weights (g) and measurements (mm) of long-billed curlew eggs.

36

Hours prior to hatching

24 12 0

Egg 1

Weight
Length
Width

Egg 2

Weight
Length

Width

Egg 3

Weight
Length
Width

Egg 4

Weight
Length
Width

74
51

71.1

71

59

X-'
78
48

69.0

68.9

68.6

67.1

67.1

56.6

55.6

57.6

-Egg 4 was preserved before it could hatch.

Acknowledgments

I wish to thank J. Andersen, J. Forsythe, K. L. Shirly, J. Wood-
son, and especially Dr. K. L. Dixon for assistance in various aspects
of this study. G. McNerney typed the manuscript. This study was
supported by a National Defense Education Act Title IV Predoctoral
Fellowship at Utah State University.

Literature Cited

Bent, A. C. 1929. Life histories of North American shore birds. US. Natl.

Mus. Bull. 146.
KoRSYTHE, D. M. 1967. Egg teeth and hatching methods of the Long-billed

Curlew. Wilson Bull. 79:3+0-341.
Forsythe, D. M. 1970. Vocalizations of the Long-billed Curlew. Condor 72:

213-224.
Graul, W. D. 1971. Observations at a Long-billed Curlew nest. Auk 88:182-

184.
LaFave, L. D. 1954. Breeding of the northern Long-billed Curlew at Sprage

Lake. Murrelet 35:48.
Palmer, R. S. 1967. Pp. 183-184 In Stout. C. D. (Ed.). The shorebirds of

North America. Viking Press, New York.
Reed, C. A. 1965. North American bird eggs. Dover Publications. Inc. New

York.

VEGETATION ZONES AROUND A SMALL POND IN THE
WHITE MOUNTAINS OF ARIZONA

B. Ira Juddi

Abstract. — Because of a previous study of wet meadows, an investigation of
Carnero Lake, White Mountains of Arizona, which is ecologically advancing
toward a wet meadow, was undertaken. Information on eutrophication and
successional patterns in ponds of the ponderosa pine is needed for better under-
standing of that ecosystem. Aerial infrared photographs were taken to delineate
areas containing aquatic vegetation and as an aid in identifying zones around the
pond. Five zones from open water to ponderosa pine were found. Abundance of
vegetation by species was estimated in each zone. Protein analyses are given for
29 of the 32 species listed.

The White Mountains of Arizona encompass many small ponds
which are used by migrating waterfowl for resting, feeding, and
sometimes as nesting sites. Arizona is located in the Pacific Water-
fowl Flyway, with most of the birds in the fall migration ar-
riving in middle to late October. Although the state is not known
for its abundance of waterfowl, small mountain ponds provide a
quality type of waterfowl himting for local people. Pond is defined
here as being an area where wave action is feeble, thereby al-
lowing a continuous vegetation cover to develop around the shore-
line.

Deer, elk, and turkey can be seen around the fringes of these
ponds, obtaining forage and water. Intergradations of plant com-
munities from the emergent aquatics in the littoral zone to the
ponderosa pine on dry land also provide many niches for small
birds and mammals. A previous study- prompted the investigation
of a pond that was geologically advancing toward a wet meadow.
Information on eutrophication and successional patterns in ponds in
the ponderosa pine is needed for a better understanding of that
ecosystem.

The reduction of the Apache menace in the 1870s opened the
territory of Arizona to farming and ranching. Concurrently, with
the increased use of range lands, rivers and streams were dammed
and water diverted for irrigation and mining purposes; one such
structure in the White Mountains of Arizona was Tyler Reservoir,
created by building a dam on Carnero Creek in 1878. Somewhere
along the way, the name was changed to Carnero Lake, possibly
by the oldtime Basque sheepherders.

Carnero Springs which empties into the lake is an old Indian
prayer spring. In earlier times many prayer beads and arrowheads
could be found there. The name Tyler was for Jack (Poker) Tyler,
who was an eccentric character of the local neighborhood. The

^Professor of Agronomy, Division of Agriculture with Arizona State University, Tempe,
Arizona.

^Patton, D R . ANn B. I. Judd. 1970. The role of wet meadows as wildlife habitat in
the Southwest. J. Range Mgmt. 23(4) :272-275.

91

92

GREAT BASIN NATURALIST

Vol. 32, No. 2

Fig. 1 . CaiTiero Lake, Apache National Forest. Arizona.

reservoir did not contain much water until a filling ditch was con-
structed in 1913.

Camero Lake is in the upper edge of the ponderosa pine forest
type at an elevation of 8500 feet. It is located approximately three
miles east of Green's Peak (Fig. 1). The lake occupies a portion of
the eastern third of the SW ^4 of section 6, Township 8 north. Range
27 east of the Gila and Salt River Meridian.

Techniques

It was evident from reconnaissance visits that there were rather
distinct vegetation zones from the pond into the ponderosa pine.

June 1972

JUDD: POND SUCCESSION

93

E3 SUBMERGED PLANTS
P EMERGENTS
a OPEN WATER

r=160 feet

Fig. 2. Location of aquatic vegetation in Camero Lake.

Abundance of vegetation by species was estimated in each zone as
very rare, rare, infrequent, abundant, and very abundant.^ Each
species was collected for reference and a sample taken for protein
analysis. Protein samples were composite samples collected in July
and August. Aerial infrared photographs were taken to delineate
areas containing aquatic vegetation and as an aid in identifying
zones around the pond.

Results and Discussion

Total surface area of Carnero Lake is 7.83 acres. Infrared photo-
graphs showed two areas containing aquatic plants. Submerged
aquatics occupied 1.76 acres, emergents 0.31 acres, and open water
5.96 acres (Fig. 2).

Five vegetation zones were identified (Fig. 3):

1. Water — containing aquatic plants.

2. Wet — shoreward zone, frequently inundated.

3. Mesic — occasionally inundated, moist part of the time.

4. Intermediate — slightly moist.

5. Forest edge — drier than intermediate.

^OOSTIIS'G, H.

San Francisco.

of plant communi

(2nd Ed). W. H. Freeman,

94

GREAT BASIN NATURALIST

Vol. 32, No. 2

Fig. 3. Vegetation zones from open water to ponderosa pine.

Table 1 lists the species found in each zone with their abun-
dance and protein content. The five zones contained 32 different
plant species. Vegetation was well defined on the north side of
the pond but the south side had a steeper bank and the zones were
not well defined. One of the first species which gains a foothold
on sandbars or sloping beach is Needle spikesedge, a small perennial.
It is found most abundantly in the Mesic Zone.

It is interesting that Arizona fescue, normally one of the prin-
cipal grasses under ponderosa pine is also found in a thin belt along
the waterline. When the area was visited in early August the
fescue next to the water had mature seed heads while plants of
the drier site were still green.

Protein is considered the most important animal nutrient and
crude protein content of plants can give an indication of forage
value for wildlife. From Table 1 it is obvious that protein is variable
but in general all zones leading to the forest edge contain some
plants high in protein. Two species, watercress and curlydock, had
exceptionally high protein content. Little information is available
on watercress as a wildlife food but it is not uncommon to see deer
foraging on curlydock in meadows.

The zones shown in Fig. 3 represent successful development from
open water to ponderosa pine. As the pond fills with sediment
and debris, mats form at the edge. Conditions are made unsuitable
for the submerged aquatics and sedges and rushes advance centripe-
tally with each zone being repeated from wet to dry conditions.

June 1972

JUDD: POND SUCCESSION

95

iC^c<ics)c>^(NCvic^oorooorO'^-<J--^'^vnvrirovr)vr>vnvn>r)vr!invnxri

xnvnoovn(N'>*'C^ooin-*'<J-triT}-c<i-<j-ro'*vn-<J--^vn-^fr>"^rO'^-*-^POrr)ej'<--i

§- is

a &

-13 2

a* ., c^ -c

ater-milfoil
ater butterc
aked sedge
ater ladysth
ilifornia bul

dge

ghead spikes
-izona fescu*
jrthem mar
ire rush
icky Mt. rus
ood-betony
ort foxtail

^^cS^Uc^«<Z^rt^c^

l2

« c-S g ?

p h i^J

cut.

i^B^S

— -t:-^ £ t.p< "5 u o

3 O i; C3

3 ,n

o i

•2 ■"
a) ^

96 GREAT BASIN NATURALIST Vol. 32, No. 2

With some modification of abundance of individual species suc-
cession should follow the pattern below.

Pond

Water-milfoil
Zone 1 Water buttercup

Water ladysthumb

Northern marmagrass
Wire rush
ry f, Rocky Mt. rush

Short foxtail
Field mint

Zone 3

Redtop bentgrass
Western yarrow
Inland loish
Piney clover

Winter bentgrass
Fleabane
Zone 4 Rocky Mt. iris

Aster
Northern meadow barley

Zone 5 Forest

Acknowledgment

The author is indebted to David R. Patton, research wildlife bi-
ologist, Rocky Mountain Forest and Range Experimental Station,
in cooperation with ASU, for his helpful suggestions during the
course of the study and wishes to express his sincere appreciation
for this assistance.

A NEW SPECIES OF GYMNODAMAEUS FROM COLORADO
(Acarina: Cryptostigmata, Gymnodamaeidae) ^

Tyler A. Woolley-

Abstract. — The taxonomic history of the genus Gymnodamaeus is reviewed
for North American species. A new species, Gymnodamaeus chalazionus, is
described from Colorado and compared with species from both the western and
eastern U.S. Among other features, the new species differs from G. veriornatus
Higgins, 1961, in the shorter interlamellar hairs and the surface structure of the
notogaster and genital plates. The new species is also compared with G. quadri-
caudatus Jacot, 1937, G. pearsei and minor Banks, 1947; with G. gildersleeveae
and ornatus Hammer 1952, from Canada, and with G. elegantulus Hammer,
1958, from South America. Twelve scanning electron micrographs are used to
delineate some of the details of integument and other features.

Relatively few species of the genus Gymnodamaeus have been
described in the years subsequent to Berlese's earlier descriptions in
1910, 1916. Grandjean (1928) described Gymnodamaeus hispanicus
from Spain and differentiated it from all other species described
prior to his article on the basis of the ornamentation of the notogas-
ter. He assumed that the surface structure was the result of secreted
materials. He also indicated that no descriptions of larvae or nymphs
had been made prior to his article, in which he included some de-
scriptions of these stages.

Jacot (1937) described, but did not figure, the species G. quadri-
caudatus from Bent Creek Experimental Forest, N. C, a small form
(0.40mm by 0.23mm) with four posterior notogastral nubbins.
Banks (1947) described and figured G. pearsei and G. minor from
Duke Forest and Durham, N. C. Hammer (1952) found G. ornatus
and G. gildersleeveae in Canada. Later (1958), she described G.
elegantulus from the Andes mountains. In the same year, Balogh
(1958) described, but did not figure, G. tunicatus from tropical Afri-
ca.

Higgins (1961) described G. veriornatus as a new species from
Utah and compared it with Hammer's species from the McKenzie
Delta of northern Canada.

Grandjean (1965) discussed the Gymnodamaeoidea as a new
superfamily in the context of a subnormal tracheal system, but did
not delineate any new species within the family or superfamily.

A collection of mites from Mt. Meeker Camp Ground, Colorado,
in 1952 had a series of nine specimens of the genus Gymnodamaeus,
which at the time were not assignable to a species category. These
specimens were later observed and determined to constitute a new
species but have remained undescribed until now. The new species
has been compared with specimens of G. veriornatus and checked
against other described species in the literature. With the discovery

'Research supported in part by NIH TG AI00094-70; also in part by a research grant from
Great Western Sugar Company, 31-1880-6257.

^Department of Zoology, Colorado State University, Fort Collins, Colorado 80521.

97

98 GREAT BASIN NATURALIST Vol. 32, No. 2

of additional specimens from other locations, and the use of repre-
sentatives of this species in SEM research, it is expedient that the
description be published.

Gymnodamaeus chalazionus, n. sp.

(Figs. l-5;6-17)

Diagnosis. — With ornate tubercles on both dorsal and ventral
surfaces and legs, hence the name chalazionus, from the Greek: hail-
stones, tubercles or knobs (all of which are applicable to the appear-
ance of the tubercles in this instance); interlamellar hairs short,
conelike, smooth compared to short, fine, filiform interlamellar hairs
in G. veriornatus; sensillum clavate, head about twice as broad as in
veriornatus; medial margins of genital plates toothed, as in veriorna-
tus, but plate shorter and wider (see Figs. 4 and 5); without ar-
rangements of six posterior notogastral hairs and reticulate dorsal
sculpturing as in G. veriornatus, slightly smaller in dimensions than
the latter, but both of these species are larger than all other North
American species described so far.

Description. — Color dark reddish brown, surface of prodorsum,
dorsum of notogaster, venter covered with cerotegument consisting
of tubercles; tubercles domelike on dorsum, bothridium and legs
(Fig. 6-10), rounded, reticulate, mammiform; tubercles of ventral
surface thimble shaped (Figs. 12-15); prodorsum about half as long
as notogaster, broadly triangular in shape; rostral and lamellar hairs
of about equal length, decurved; lamellae or costulae absent, but a
sclerotized line extending medially between lamellar hairs (Fig. 1);
interlamellar hairs simple, short, conical, inserted in medial apex
of sclerotized, arched, curved bars anteromediad of pseudostigmata;
pseudostigmata rounded, with lip erected above surface of prodor-
sum, covered internally and externally with mammiform tubercles,
sensillum clavate, spined, length nearly twice as long as distance
between pseudostigmata; tectopedia I and II covered with tubercles.

Hysterosoma nearly rounded in outline, with crenulated margin
and wrinkled surface surrounding slightly raised central dome, tu-
bercles arranged in somewhat racHating lines as in drawing (Fig. 1)
and SEM micrograph (Fig. 8); posterior margin of notogaster with
six hairs and two lyriform jiores as indicated in Figure 1.

Camerostome about half the length of propodosoma, mentum
broad, rutella narrowed anteriorly; apodemata as shown in Figure 2,
ventral setae obscured by integumental covering; apodermata IV
arched anteriorly over genital aperture, consisting of a double-
ridged arrangement; genital and anal ai)ertures contiguous along en-
tire width; genital covers serrated on medial margin, each cover
with 6 genital setae and with nine short medial dentes interlocking
covers, surface of covers with tuberculated cerotegument; a sclero-
tized ring around genital aperture and continuous with similar ring
around anal aperture; anal covers longer than wide, with tubercu-
lated cerotegument.

June 1972

WOOLLEY: A NEW MITE

99

Figs. 1-4. Gymnodamaeus chalazionus: 1, dorsal aspect, legs partially omit-
ted; 2, ventral aspect, legs omitted; 3, tibia and tarsus of leg I of female specimen;
4, genital plates.

Fig. 5. Gymnodamaeus veriornatus genital plates, drawn to same scale as
Figure 4.

Legs with elongated articles, surface covered with mammiform
tuberculated integument, similar tubercles on hairs; tibia of leg I
with elongated apophysis from which solenidion emerges (Fig. 3);
all legs heterotridactylous, median claw larger than laterals.

Measurements. — Of the measurable specimens, the size

•Magnifications of all micrographs are pla
may alter this dimension slightly.

size. Reductions of the micrographs for printing

100

GREAT BASIN NATURALIST

Vol. 32, No. 2

Figs. 6-9. Gymnodamaeus chalazionus scanning electron micrographs: 6,
prodorsum, 200X'; 7, pseudostigmata and sensillum. anterior margin of noto-
gaster, 500X; 8. notogaster and leg III, 300X; 9, rim of pseudostigmata and
enlarged view of integumeiual tuhorcles inside pseudostigmata! cup. lO.OOOX.

ranges are for length: 804-768 /.i; for width: 510-450 ^li; average
length 791 /i; average width 553 /t.

Nine specimens of the species (6 males, 2 females, 1 undeter-
mined sex) taken from Mt. Meeker Camp Ground. Boulder Co.,
(Colorado, 17 July 1952, by T. A. Woolley; nimierous additional spec-
imens taken near Fort Collins on different dates, most recently
(1971) taken near Hayden. (>olorado. The male holotype and one
paratypc are in the U.S. National Museum, the remaining paratypes
are in n\\ collfHtion.

1972

WOOLLEY: A NEW MITE

[01

Figs. 10-13. Gymnodamaeus chalazionus scanning electron micrographs: 10,
mammiform tubercles of notogastrial surface, 10,000X; 11, ventral aspect, 150X;
12, en face view of infracapitulum, showing dorsal chelicerae, veneral rutella
and lateral palps, 700X; 13, tubercles of ventral integument, 10,000X.

Discussion. — Seven species of Gymnodamaeus were reviewed
for this paper, among which is a species of Koch, illustrated and
described by Schweizer (1922) under the name Damaeus femoratus.
The illustrations show that this should be changed to the genus
Gymnodamaeus as it does not belong in the genus Damaeus. Schwei-
zer indicated that the species was found in Algeria, Italy, and Ger-
many.

Of the North American species Gymnodamaeus veriornatus Hig-
gins is the largest in size (0.93mm x 0.54mm), with G. chalazionus.

102

GREAT BASIN NATURALIST

Vol. 32, No. 2

Figs. 14-17. Gymnodarnaeus chalazionus scanning electron micrographs: 14,
genital and anal plates, 300X; 15, serrate margin of genital plates ;i,OOOX;
16, leg joint, 300X; 17, leg and leg hair showing integumental tubercles,
10,000X.

next in size (804 /i x 510 /i. Hammer's species G. ornatus is in-
termediate in dimensions (0.68mm x 0.37mm) ; G. gildersleeveae and
G. elegantulus have identical measurements (0.41mm x ().23mm).
Banks's species are in between others in size, with G. minor at 0.35
mm long and G. pearsci measuring 0.50mm in length.

Specimens of G. chalazionus were used in scanning electron mi-
croscopic research and the accompanying micrographs of this species
show a three-dimensional detail that is not possible with the dia-
grams. Although the micrographs are self-explanatory in many re-
spects, it shoukl be noted that the tubercles of the dorsum, the interi-

June 1972 woolley: a new mite 103

or of the bothridium, and the surface of the legs and leg hairs are
similar — appearing as perforated mammifomi structures (Figs. 7-
10). The tubercles of the venter are more elongated than those of
the dorsum or the other locations (Figs. 11-15).

As stated earlier, Grandjean (1928) assumed that the surface
integument of Gymnodamaeus was a secreted material. This is borne
out by the SEM micrographs (Figs. 6, 8, 11-15) of G. chalazionus.
Specimens of this species were treated with xylene to determine if
the tubercles could be separated or removed from the integument.
They remained in place and appear to be wax blooms of different
configurations on the dorsum and the venter. Further investigations
with the SEM should disclose more details about the structure and
chemical composition of these integumental structures in this new
species of oribatid.

Literature Cited

Balogh, J. 1958. Oribates Nouvelles de L'Afrique tropicale. Revue de Zool.

et de Bot. Africaines 58(i/2) : 1-34.
Banks, N. 1947. On Some Acarina from North Carolina. Psyche 54(2) : 110-

141.
Berlese, a. 1910. Brevi diagnosi di generi e specie nuovi di Acari. Redia

6:346-388.

. 1916. Centuria prima di Acari nuovi. Redia 12:19-67.

Grandjean, F. 1928. Deux nouvelle Oribatei d'Espagne. Bull. Soc. Zool. 53

(6): 424-441.
. 1965. Nouvelles observations sur les Oribates (4e Serie). Acarologia

7(1):91-112.
Hammer, M. 1952. Investigations on Microfauna of Northern Canada. Acta

Arctica 4:108p.
— — — . 1958. Investigations of Oribatid Fauna of Andes Mts.-I The Argentine

& Bolivia. Biol. Skr. Dan. Vid. Selsk. 10(1): 1-129.
HiGGiNs, H. G. 1961. A new beetle mite from Utah (Oribatei-Gymnodama-

eidae). Great Basin Nat. 21 ()^): 27-28.
Jacot, a. p. 1937. Journal of North American Moss Mites. Journal N. Y.

Entomol. Soc. 45(3/4): 353-3 75.
ScHWEizER, J. 1922. Beitrag zur Kenntnis der Terrestrischen Milbenfauna der

Schweiz. Verhandl. Naturforsch. Gesell. in Basel. 33:23-112.

THE SONORAN SUBSPECIES OF THE LIZARD
CTENOSAURA HEMILOPHA

Hobart M. Smith'

Abstract. — Five subspecies of Ctenosaura hemilopha are recognized, in-
cluding C. h. hemilopa from Baja California; C. h. insulana from Isla Cerralvo,
Gulf of California; C. h. conspicuosa from Isla San Esteban and (?) Isla Lobos.
Gulf of California; and two new subspecies: C. h. macrolopha from Sonora and
northern Sinaloa, and C. h. nolascensis from Isla San Pedro Nolasco, Gulf of
California. The species is thought to have evolved from C. pectinata, through
the somewhat similar C. h. macrolopha or its forerunner. Its range probably ex-
tended at an earlier time northward around the Gulf of California, thence south-
ward through Baja California, with populations reaching certain Gulf islands from
the peninsula, not from mainland Mexico, despite greater proximity of some of
the islands inhabited to Sonora than to the peninsula.

It has long been known that populations of Ctenosaura hemilopha
(Cope) of Sonora and adjacent areas differ from those of Baja Cali-
fornia (Smith, 1935: 140-141; Lowe and Norris, 1955: 90). The
peninsular subspecies was identified as C. h. interrupta Bocourt,
1882, by Lowe and Norris {loc. cit.), the mainland subspecies as
C. h. hemilopha (Cope), 1863. The type-locality of Cyclura {Cteno-
saura) hemilopha Cope is, however, "Cape St. Lucas," Baja Califor-
nia, and therefore Bocourt's Ctenosaura interrupta (type-locality
Baja California) is a junior synonym of it, as noted by Hardy and
McDiarmid (1969: 122). Accordingly, the mainland subspecies is
presently nameless. It is here named

Ctenosaura hemilopha macrolopha, subsp. nov.

HoLOTYPE. — Chicago Natural History Museum 108705 (former-
ly E. H. Taylor 235), La Posa, San Carlos Bay, 10 mi NW Guay-
mas, Sonora, 30 June 1934, E. H. Taylor coll. Paratypes: Univ. Illi-
nois Mus. Nat. Hist. 20232 (formerly E. H. Taylor 121), 5 mi SW
Hermosillo, Sonora, 26 June 1934, E. H. Taylor; Univ. Colorado
Mus. Nat. Hist. (UCMNH) 42536, Yepachic, Chihuahua, 5800 ft,
summer 1970, Campbell Pennington; Brigham Young Univ. 14616-
24, Urique, Chihuahua.

Diagnosis. — A mainland Mexican geographic segment of C.
hemilopha with a long middorsal crest reaching to groin and usually
with 8 or more whorls of enlarged caudal scales separated from
adjacent such whorls by no more than a single intercalary row of
scales at any point.

Description of holotype. — A mature male, 194 mm s-v; pat-
tern as depicted in Smith {op. cit.: pi. 23) and measurements as
given in the same work {op. cit.: 141).=

'Department of Biology, University of Colorado, Boulder, Colorado 80302.

-'Art. li i>f the Iiiloni.ilionnl Cooo (if Zoological Nomenclature permits "a definite bibliographic
rrfrreiKc " even to llie required ".statement that purports to give characters differentiating
the taxon" — i.e., the "diagnosis" of a new name Since thai statement is given here, a biblio-
graphic reference to the nomenclaturally less essential description is presumably acceptable.

104

June 1972

SMITH: SONORAN LIZARDS

105

Comparisons. — The species C. hemilopha differs trenchantly
from C. pectinata, its nearest neighbor, by having the whorls of
small scales between the whorls of enlarged scales reduced to less
than two complete whorls (except on the middorsal line, where
there is often one spine) over a large part of the tail, whereas on no
part of the tail in pectinata are there fewer than two complete
whorls of small scales (except at the middorsal line). In addition,

)

J

, 1 -

■\

\

^

-4

I 1 \(

11]

&

'^

I&

Xv

^

.%

r-

^ ^

%.

Fig. 1. Ranges of the subspecies of Ctenosaura hemilopha and of nearby
populations of C. pectinata. The dots indicate localities of record. 1, C. h. hemi-
lopha; 2. C. h. conspicuosa on Isla San Esteban; 2a, C. h. conspicuosa (?) on Isla
Lobos; 3, C. h. nolascensis on Isla San Pedro Nolasco; 4. C. h. insulana on Isla
Cerralvo; 5, C. h. macrolopha; P, C. pectinata. The projected range of C. h.
hemilopha corresponds with the limits of the San Lucan herpetofaunal area as
mapped by Savage (1960:194). Localities and ranges in Sinaloa were drawn
largely from Hardy and McDiannid (1969:231). The projected range in Sonora
is based largely upon Nelson and Goldman (1926: fig. 13). Base map courtesy
Roger Conant.

106 GREAT BASIN NATURALIST Vol. 32, No. 2

the middorsal crest in all forms of hemilopha extends no farther
posteriorly than a point somewhat anterior to the level of the groin;
whereas in pectinata the crest extends to the level of the sacrum,
terminating anywhere from the level of its anterior border to the
base of the tail. C. hemilopha is also brownish in adults and sub-
adults; whereas pectinata exhibits grayish or greenish tones through-
out life, developing black markings on a gray background in adults.
These distinctions hold throughout the 20 C. pectinata examined
from the states of Sinaloa, Nayarit, Michoacan, Morelos, and Oaxaca,
and 28 C. hemilopha representing all subspecies.

The mainland examples of hemilopha differ from all other pop-
ulations of that species in possessing the longest dorsal crest occur-
ring in the species, extending posteriorly nearly to the level of the
groin, not quite reaching the level of the sacrum (as in pectinata).,
in all other populations of hemilopha^ the dorsal crest ends at a level
no more than 4/5 the distance from axilla to groin. In addition,
the mainland population differs from all other subspecies of hemi-
lopha except insulana and hemilopha in usually having no evidence
whatever of a second whorl of small scales between at least 8 ad-
jacent whorls of enlarged scales, in the central region of the tail.
Two of 10 macrolopha examined for this character failed to meet
this criterion, having one or no whorl-set at all completely reduced
to one intercalary row. In the other eight, the incomplete second row
begins at points varying from the 16th whorl-set to the 24th, and
in three the second row is not evident in any distal whorl posterior
to the basal 4 or 5 whorl-sets. Adults retain narrow light bars on the
forelegs.

Remarks.— The subspecific name macrolopha (Gr., long crest)
refers to the exceptionally long crest, reaching to level of groin, in
the mainland population, as contrasted with the short crest (pre-
sumably the implication of hemilopha, Gr., half crest), reaching
only a little beyond midtrunk in other populations of the species. In
this respect macrolopha is intermediate between pectinata, wdth a
crest uninterrupted or narrowly interrupted at sacrum, and insular-
peninsular C. hemilopha. C. pectinata extends northward as far as
about parallel 25 °N, north of Culiacan about 20 miles, where its
range meets, blends with or overlaps that of C. hemilopha (Hardy
and McDiarmid, 1969:119-124, 231, fig. 44). Possibly, although the
trenchant differences do not suggest it, the two populations hitherto
designated as the species C. pectinata and C. hemilopha intergrade
in the Culiacan area; if not, a certain amount of interspecific intro-
gression seemingly occurred prior to attainment of reproductive iso-
lation, accounting for the pectinata-\\ke long crest in the mainland
populations of hemilopha. Further field work will be required for a
definitive conclusion.

Range. — The subspecies C. fi. macrolopha ranges from the vicin-
ity of Hermosillo, Sonora, through the northern third of Sinaloa,
and inland as far as extreme western Chihuahua (Batopilas, Urique,
Yepachic). Bogert and Oliver (1945:334) conjecture that C. hemi-

June 1972 smith: sonoran lizards 107

lopha may once have occurred well north of its present range limits,
the peninsular and mainland ranges being continuous north of the
Gulf of California. The hypothesis is perhaps strengthened by the
long-unconfirmed record of the species from Nogales (Bailey,
1928:21, et. al.). If valid in the context at least of former distribu-
tion, the record is no doubt referable to macrolopha. It occurs on no
island, so far as now known.

Contrary to comments by Smith {op. cit.) and Lowe and Nor-
ris {op. cit.), the population on Isla San Pedro Nolasco, Sonora
(about 13 mi W of the coast near Guaymas), differs sharply from
C. h. macrolopha in having a short middorsal crest and at least par-
tial second rows of small scales between most whorls on the tail. The
specimen figured by Bailey (1928: pi. 5) from that island has a
short crest, contrary to Smith's {op. cit.) interpretation. The great-
est similarity in scutellation is to conspicuosa, but the pattern of the
San Pedro Nolasco population is distinctive. That population is here
named

Ctenosaura hemilopha nolascensis, subsp. nov.

HoLOTYPE.— Univ. Colo. Mus. Nat. Hist. No. 26391, a subaduh
male, taken on Isla San Pedro Nolasco, Sonora, 16 August 1964, by
T. Paul Maslin et. al. Paratypes: Thirteen, all topotypes: UCMNH
Nos. 26388-90, 26392-4, 35174-7, and one unnumbered; Mus. Comp.
Zool. 13178-9.

Diagnosis. — A geographic segment of C. hemilopha restricted to
Isla San Pedro Nolasco, with enlarged scales of middorsal crest ex-
tending posteriorly no farther than a level about two-thirds the dis-
tance from axilla to groin; with rare exception, all whorls of en-
larged scales separated from adjacent whorls by at least a partial
second row of small scales on ventral surface of tail; a second row
at least partially represented on dorsal surface through anterior 3-12
whorls (seldom more than 6); a third row at least partially repre-
sented in 5-7, usually 6 whorls; ventral surfaces of subadults with
scattered rounded dark spots about ^3 diameter of tympanum; arm
all black, lacking light bars, in adults; hind leg reticulated with
black above, bars scarcely evident, in adults.

Description of holotype. — Enlarged scales of dorsal crest ex-
tending posteriorly about 4/7 distance from axilla to groin; at
least one complete intercalary row of scales between each pair of
adjacent whorls of enlarged scales; a second intercalary row ex-
tending across ventral surface between all whorls, and represented
dorsally also in the basal 6 sets of whorls, and in the 18th and fol-
lowing sets; at least part of a 3rd intercalary row present in the
basal 7 sets of whorls, 4 in the basal 5, 5 in the basal 3, 6 in the basal
2 (the first set of whorls is arbitrarily counted as the one following
the anteriormost whorl containing scales at least twice as large as
those immediately preceding it); femoral pores 6-7. Snout-vent 167
mm; tail complete except for the terminal regenerated 50 mm, un-
regenerated part 248 mm.

108 GREAT BASIN NATURALIST Vol. 32, No. 2

Color in preservative a brownish slate; body unicolor above ex-
cept for numerous small rounded dark flecks (each covering 1-9
scales, on sides of body, dim or absent near median line), two
broad brownish black crossbands (one on shoulder, one just posterior
to axilla, joined laterally), and a small round blackish brown spot
on middorsal line at rear of neck; a few faint light spots on a dorso-
lateral line on neck; forelegs black above, with poorly defined, nar-
row, light crossbars; hind leg essentially dark reticulated on a
lighter background, the reticulum evidencing the broad dark bars
and intervening light bands (about 1/3 as wide as dark bars) of ju-
veniles; tail with broad dark rings, 4 to 5 times as wide as interven-
ing light rings, beginning at about 15th set of whorls. Throat and
chest to near level of axilla dusky, with scattered lighter brown dots;
rest of chest progressively less dusky posteriorly, lighter midventral-
ly; ground color still lighter on ventral surfaces of legs and base of
tail; numerous, randomly scattered, rounded, dark brown spots on
ventral surfaces of abdomen, hind legs and base of tail, especially
prominent on legs, somewhat larger than those on sides of abdomen,
in general about 1/5 the lesser diameter of tympanum.

Variation. — Smaller individuals have hght crossbars on the
back, 7 between axilla and groin; the legs are more prominently
barred, the dark spotting on the ventral surfaces more extensive
(the spots often arranged in transverse rows on abdomen) and the
chin and lips prominently dark-and-light barred. A large adult
male (estimated 400 mm s-v) is solid black over the throat, chest,
forelegs, axilla, and shoulders; chin, lips, sides, and dorsal surface
of neck gray brown; black crossing dorsum in two bands on shoulder;
rounded median black spot on nape prominent; abdomen, both
dorsally and ventrally, a light gray brown, with numerous very
small black flecks laterally; groin rather heavil}^ pigmented, but ven-
tral surface of thigh with extensive light areas; dorsal surface of
thigh densely but not coarsely reticulated, very little evidence of bar-
ring; feet and shank reticulate but barred.

Ten of the 11 paratypes examined for characteristics of the
caudal whorls and crest have at least part of a second intercalary
row (ventrally) between all adjacent pairs of whorl-sets ; the one
exception, No. 35176, has one intercalary row only in the 13th
whorl-set. Five to 7 (usually 6) basal whorl-sets have at least parts
of at least 3 intercalary rows, 3-5 (usually 4) have at least 4 or
parts thereof. Three to 12 basal whorl-sets have at least 2 rows of
intercalary scales represented dorsally. The enlarged scales of the
dorsal crest extend posteriorly from 4/7 to 4/5 the distance from
axilla to groin, usually about 4/7.

Comparisons. — C. h. nolasccnsis differs more conspicuously
from its geographically nearest population, in Sonora {C. h. macro-
lopha), than from any other subspecies. In rnacrolopha the dorsal
crest extends nearly to groin, exceeding the limit of 4/5 the distance
from axilla to groin that marks the greatest extent in nolascensis
and other subspecies. In addition, at least 8 whorl-sets in rnacrolopha

June 1972 smith: sonoran lizards 109

possess no more than one intercalary row, whereas that figure in
nolascensis exceeds no more than one.

C. h. nolascensis is less markedly distinct from the other geo-
graphically more remote subspecies. C. h. insulana is similar to
macrolopha (but differs from nolascensis) in having a large number
of whorl-sets with no more than one row of intercalary scales (14
in the single specimen examined) ; its limbs are more prominently
banded, and light crossbars persist on the trunk throughout life. C. h.
conspicuosa loses the light crossbars in the postthoracic region and
on limbs in adults, but is similar to nolascensis in caudal scutella-
tion; the two specimens examined have 4-5 basal whorl-sets with
more than two at least partial intercalary rows, 2-3 basal whorl-sets
with more than three, and 2-4 basal whorl-sets with more than two
intercalary rows dorsally; but it is likely that no real differences in
these characters exist in comparison with other subspecies. C. h.
heniilopha (4 specimens examined) has much the same scutellation
as insulana and retains the light crossbars on the postthoracic re-
gion and hind limbs; in both these characters, it differs from nolas-
censis.

Unfortunately, it has not been possible to examine the caudal
scutellation of an adequate series of all taxa. We have examined
sufficient material of pectinata^ h. macrolopha, and h. nolascensis to
be confident of reliability of the characters observed, but the other
subspecies of hemilopha {insulana., conspicuosa, heniilopha) need
further observation not practical at the present time. Additionally,
fieldwork in the area of contact of pectinata and h. macrolopha will
be required to establish conclusively the interrelationships of these
taxa, although we remain confident that no genetic interchange now
takes place there and that the taxa are of specific rank relative to
each other.

Phylogeny

The evolution of hemilopha and its subspecies is of major inter-
est. Because macrolopha is more like pectinata than is any other race
of hemilopha., and is geographically adjacent, it seems reasonable to
hypothesize that hemilopha is a derivative of pectinata or a pectin-
a/fl-like ancestor and that macrolopha more closely resembles the
ancestral form of hemilopha than does any other race of the species.
Because of the trenchant difference between insular-peninsular races
on the one hand and the mainland race on the other, the close
geographic proximity of the former group to the mainland, and the
relative uniformity of characteristics of the members of the insular-
peninsular group, it seems unlikely that the species radiated across
the Gulf of California to the Peninsula. A more likely probability is
that the species extended northward at one time, and thence into
the Peninsula, and subsequently became more restricted in range,
separating two components, one on the Peninsula and one on the
mainland. C. h. macrolopha, remaining with or reestablishing con-
tact with pectinata or its ancestor, maintained or developed a char-
acter displacement that accentuated the basic difference in caudal

110 GREAT BASIN NATURALIST Vol. 32, No. 2

scutellation between the two species, although macrolopha retained
an intermediate character-state in extent of the crest, or perhaps in-
trogression was responsible for the intermediacy.

The peninsular population diverged in character while in isola-
tion, accentuating the brevity of the dorsal crest. Its range became
essentially relic in nature, restricted to the southern third of the
peninsula, except as a few waif populations in a sweepstake pattern
reached a number of the Gulf islands. In their isolation from contact
with other forms of the genus, these residual populations lost to
varying degrees part of the unique caudal scale pattern, approach-
ing but not achieving the presumably ancestral condition of pectin-
ata.

With taxonomic recognition given to five geographically isolated
populations of hemilopha, virtually all that exist (Soule and Sloan,
1966:140), it is unlikely that any other extant subspecies wdll re-
quire recognition in the future, except perhaps for the population
on Isla Lobos recorded by Lowe and Norris (1955:95). That popula-
tion, so close to Tiburon island (not the Isla Lobos S of Guaymas),
may perpetuate the group that no doubt formerly occurred on the
larger island but is now extinct, probably due to human predation.
Whether the Tiburon-Lobos populations have diverged sufficiently
from others to merit recognition remains to be seen (Soule and
Sloan, 1966:140, refer Lobos material to h. conspicuosa) . In any
event, their relation to the peninsular-insular group as opposed to
the mainland group is of major interest.

Acknowledgments

I am much indebted to Drs. T. Paul Maslin and Wilmer W.
Tanner for the privilege of studying material in the museums of
the University of Colorado and Brigham Young University, respec-
tively; to Richard Holland, Tom Bauer, and Dr. Ronald A. Brandon
for specimens they provided for the museum; and to Alan Savitzky
for aid in studying the material. This work was supported in part by
publication grant LM 00925 from the National Library of Medi-
cine, National Institutes of Health, U.S. DHEW; and in part by
the Biology Department of the University of Colorado.

Key to the Ctenosaura of Northwestern Mexico

1 . All whorl-sets of tail containing at least two complete

intercalary rows of scales (except sometimes on mid-
dorsal line) ; dorsal crest extending to, onto or through
sacral region; general color tone green (in juvenile) to

gray (adults) pectinata

No more than 6 basal whorl-sets of tail containing two
complete intercalary rows of scales (excluding mid-
dorsal line); dorsal crest not reaching sacral region;
general color tone green (very young) to brown
(subadults, adults) — hemilopha 2

June 1972 smith: sonoran lizards HI

2(1). Dorsal crest extending posteriorly nearly to groin, pos-
terior to a point about 4/5 distance from axilla to
groin; usually at least 8 central caudal whorl-sets
with no evidence whatever of a second intercalary

row of scales h. macrolopha

Dorsal crest terminating posteriorly near a point usually
about % distance from axilla to groin, or anterior
thereto; central caudal whorl-sets with a partial sec-
ond intercalary row or not 3

3(2). At least 8 central caudal whorl-sets with no evidence
whatever of a second intercalary row of scales; light
bars on limbs and postthoracic trunk retained through-
out life 4

No more than 1 or 2 caudal whorl-sets with no evidence
whatever of a second intercalary row of scales; pattern
diverse 5

4(3) . Dorsal crest terminating posteriorly near a point about %

distance from axilla to groin h. hemilopha

Dorsal crest terminating about at midabdomen .... h. insulana

5(3). Juvenile and young adults only with round dark spots
(about 1/3 lesser diameter of tympanum) scattered over
ventral surfaces of trunk, hind legs and base of tail,
becoming small dark flecks in adults; dark and light
markings on limbs not sharply contrasted .... Ji. nolascensis
Rounded dark marks on ventral surfaces of juvenile re-
tained at least on hind limbs throughout life; light
and dark markings of hind limbs sharply contrasted
both above and below h. conspicuosa

Literature Cited

Bailey, J. W. 1928. A revision of the lizards of the genus Ctenosaura. Proc.

U. S. Nat. Mus., 73(12): 1-55, pis. 1-30.
BoGERT, C. M., AND J. A. Oliver. 1945. A preliminary analysis of the herpeto-

fauna of Sonora. Bull. Amer. Mus. Nat. Hist., 83:297-426, figs. 1-13, pis.

30-37, maps 1-2.
Hardy, L. M., and R. W. McDiarmid. 1969. The amphibians and reptiles of

Sinaloa, Mexico. Univ. Kansas Publ. Mus. Nat. Hist., 18(3) :39-252, figs.

1-91, pis. 1-8.
Lowe, C. H., Jr., and K. S. Norris. 1955. Analysis of the herpetofauna of

Baja California, Mexico. III. New and revived reptilian subspecies of Isla

de San Esteban, Gulf of California, Sonora, Mexico, with notes on other

satellite islands of Isla Tiburon. Herpetologica, 11:89-96.
Nelson, E. W., and E. A. Goldman. 1926. Mexico. In. Victor E. Shelford,

Naturalist's guide to the Americas. Baltimore, Williams and Wilkins. xv,

761 pp., 16 figs. (Nelson and Goldman: 574-596, fig. 13.)
Savage, J. M. 1960. Evolution of a peninsular herpetofauna. Syst. Zool.,

9:184-212, figs. 1-15.
Smith, H. M. 1935. Miscellaneous notes on Mexican lizards. Univ. Kans.

Science Bull., 22(6) : 1 19-155, pis. 23-25.
SouLE, M., AND A. J. Sloan. 1966. Biogeography and distribution of reptiles and

amphibians on islands in the Gulf of California, Mexico. Trans. San Diego

Soc. Nat. Hist., 14(11): 137-156.

DEAD GREAT BLUE HERON FOUND AT 11,000 FOOT
UTAH ELEVATION

Gary L. Hickman'

Abstract. — A Great Blue Heron (Ardea herodias) was found dead at the
west end of the Utah Uinta Mountains at approximately 11,000 feet. Rain, fog,
and very low cloud ceilings reducing visibility to almost zero causing directly or
indirectly the heron's death was postulated.

On 5 September 1965 I found a dead Great Blue Heron {Ardea
herodias) at the west end of the Utah Uinta Mountains on Utah
Highway 150 just under the north side of the 11,000 foot Hayden
Pass. The heron was found just after sunrise when visibility at
ground level to an undetermined altitude was almost zero because of
fog Rains had fallen periodically throughout the night. The heron
was stiff from rigor mortis and the exact cause of death was not de-
termined, but it appeared the bird had either crashed into the road
or landed on the road and was hit by a vehicle. The heron was not
on the highway at 2200 hours the previous night when I traveled
the road.

The Uinta Mountains form the only major east-west mountain
range in the United States, extending approximately 131 miles. No
Great Blue Herons were observed above 7000 feet in the Uinta
Mountains during my two-year mule deer study or during a two-
year study of waterfowl use of high mountain lakes conducted by
Steven Peterson.

Numerous accounts of migrating birds colliding with man-made
structures, including tall lighted buildings and radio and television
towers, have been reported. Forty-six species of birds were picked
up on the night of 19-20 September 1963 at the television towers at
Eau Claire, Wisconsin. David W. Johnston, at the University of
Florida. Gainesville, estimated that 50,000 birds died on the nights of
7 and 8 October 1954 at the ceilometer of the Warner Robins Air
Force Base near Macon, Georgia. In all instances these mass col-
lisions were associated with rain, fog, and very low cloud ceilings
(John Vosburgh, 1966. Birds in Our Lives. Washington, D.C.,
United States Government Printing Office, p. 361-364).

This observation raises the questions: Do Great Blue Herons
migrate over the Uinta Mountain ranged During migrations over
mountain ranges, under certain climatic conditions is there a mor-
tality resulting from collisions with natural objects and of what
magnitude?

'Utah Coopemtivo Wildlife Research Unit. Utah State University, Logan, Utah. Now at Bureau
of Sport Fisheries and Wildlife, Fish and Wildlife Service, 1031 Miracle Mile, Vero Beach, Florida
^2960.

12

FIVE NEW VARIETIES OF ERIOGONUM (POLYGONACEAE)

James L. Reveal^

As the first volume of Intermountain Flora: The Vascular Plants
of the Intermountain West, by A. Cronquist, A.H. Holmgren, N.H.
Holmgren, and J.L. Reveal, nears completion, a number of minor
nomenclatural changes are needed for the introductory material in
the first volume. Here, five new varieties of Eriogonum (Polygona-
ceae) are proposed, described, and discussed.

Eriogonum brevicaule Nutt. var. cottamii (S. Stokes) Reveal,
COMB. NOV., based on E. tenellum Torr. ssp. cottamii S. Stokes, Gen.
Eriog. 70. 1936.

Plants 1-3.5 dm tall; leaves narrowly oblanceolate to narrowly
elliptic, the leaf-blade 3-7 cm long, 3-5 mm wide, densely tomen-
tose on both surfaces, the margin plane or with thickened edges, not
revolute; flowering stems erect, 0.5-2 dm long, tomentose, the tomen-
tum grayish when young, becoming tannish- to reddish-brown at
maturity; inflorescences cymose, open, 3-7 cm long, tomentose;
peduncles lacking; involucres turbinate, 2-3 mm long, 1.5-2 (2.5)
mm wide, tomentose; flowers yellow, 3-4 mm long.

Type. — Canyons in bottoms of the slopes on West Mountain,
Utah Co., Utah, at 4500 feet elevation, 20 August 1925 (misquoted
as 1924 in the original description), Cottam 411. Holotype, BRY!
Isotype, UT!

Distribution. — Infrequent on dry gravelly limestone slopes
and clay hills in central Utah from eastern Tooele and Juab cos. to
western Utah Co. Flowering from June to September.

The var. cottamii is an easily distinguished entity in the Utah
Buckwheat flora. It differs from var. brevicaule in having densely
tomentose stems, and from var. laxifolium (Torr. & Gray) Reveal
in having a cymose inflorescence instead of a capitate or subcapitate
one. The southwestern Wyoming phase of the species, var. micran-
thum (Nutt.) Reveal perhaps is most closely related to var. cottamii,
the two differing in the color of their tomentum, flower size, and
distribution.

Eriogonum brevicaule Nutt. var. wasatchense (M. E. Jones) Re-
veal, COMB. NOV., based on E. wasatchense M. E. Jones, Contr. W.
Bot. 11:11. 1903.

Plants 3-5 dm tall; leaves narrowly elliptic, 1.5-4 cm long, (3)
4-7 mm wide, densely white-tomentose below, floccose above, the
margin crenulate in most; flowering stems erect, 1-3 dm long, gla-
brous; inflorescences cymose, (8) 10-15 cm long, glabrous; peduncles
rarely present; involucres turbinate, 3-4 mm long, 2-2.5 mm wide,
glabrous; flowers white, 2-2.5 mm long.

^Departments of Botany, University of Maryland, College Park 20742 and National Museum
of Natural History, Smithsonian Institution, Washington, D. C. 20560.

113

114 GREAT BASIN NATURALIST Vol. 32, No. 2

Type.— American Fork Canyon, Utah Co., 27 July 1880, M.E.
Jones 1877. Lectotype, POM! Duplicates of the lectotype, BM, CAS,
GH, MICH, POM, US, UTC!

Distribution. — Infrequent on dry talus slopes and limestone
outcrops along the western flank of the Wasatch Mountains from
Davis Co. south to Utah Co., Utah. Flowering from June to Septem-
ber.

The var. wasatchense is perhaps the most distinct of the several
variants within Eriogonum brevicaule. It is quickly recognized by
its cymose, glabrous inflorescence, glabrous stems and crenulate
leaves sheathing up the lower portions of the stems, and the white
flowers.

Eriogonum shockleyi S. Wats. var. longilohum (M.E. Jones)
Reveal, comb, nov., based on E. longilobum M.E. Jones, Proc. Calif.
Acad. Sci. II, 5:720. 1895.

Low, pulvinate, matted herbaceous perenials forming a dense
mat (1) 2-4 dm across; leaves in rather loose rosettes, the leaf-blade
oblanceolate to spatulate, (3) 5-8 (12) mm long, (2) 3-6 mm wide;
flowering stems up to 3 cm long, tomentose; involucres (3) 4-6 (7)
mm long, the lobes (1) 2-3 (3.5) mm long; flowers white, 3-4 mm
long; achenes mostly densely pubescent.

Type.— Near Price, Carbon Co., Utah, 4 July 1894, M.E. Jones
5590/. Holotype, POM! Isotype, US!

Distribution. — Dry clay flats and lower slopes in eastern Utah
from Duchesne and Uintah cos. southward to northern Arizona in
Coconino, Navajo and Apache cos., and in western Colorado from
Moffatt and Mesa cos. south to San Juan Co., New Mexico. Flower-
ing from May to July.

In the past, three forms within Eriogonum shockleyi have been
maintained. However, as additional material has been collected and
field studies made on the species since 1965, the distinction between
var. shockleyi and the form named candidum by Stokes (1936) be-
came less clear. It now appears impossible to maintain the yellow-
flowered phase {candidum) as a separate entity from the white-
flowered phase {shockleyi) due to a series of pale-yellow popula-
tions across central Nevada. The third element, var. longilobum, can
still be recognized, based on the characteristics outlined below.

A. Leaves 2-5 (6) mm long, 2-4 mm wide; involucres 2-3.5
mm long, the lobes 0.5-1.8 (2) mm long; flowers white or
yellow, 2.5-3.5 (4) mm long; Inyo Co., California, eastward
across central Nevada to western Utah and southern
Idaho var. shockleyi

AA. Leaves (4) 5-12 mm long, 2-6 mm wide; involures (3.5)
4-6 mm long, the lobes (1.5) 2-3 mm long; flowers white,
3-4 mm long; eastern Utah and western Colorado south to
northeastern Arizona and extreme northwestern New
Mexico var. longilobum

June 1972 reveal: erogonum 115

Eriogonum ovalifolium Nutt. var. caelestinum Reveal, var. nov.
A var. nivali (Canby in Gov.) M.E. Jones et var. depressa Blank,
floribus flavis demum rubro-tinctis, involucris solitariis differt.

Low densely caespitose herbaceous perennials forming a mat
0.5-1 dm across; leaves in densely congested basal rosettes, the leaf-
blade mostly elliptic, 2-5 mm long, 1.5-3 mm wide, thinly greenish-
tomentose on both surfaces with a tannish margin, the petiole 1-3
mm long, thinly pubescent; scapes 1-6 cm long, thinly floccose;
involucres solitary or very rarely in pairs, turbinate-campanulate,
2-2.5 mm long; flowers yellow, maturing red or reddish-tinged in
most, 2.5-3 mm long.

Type. — South fork of Pine Creek on the upper ridges of the
Toquima Range, Toiyabe National Forest, Nye Co., Nevada, from
10,900 to 11,800 feet elevation, 23 July 1964, Reveal 629. Holotype,
US! Isotypes, ARIZ, BRY, CAS, DS, GH, MO, NY, OKL, RM, RSA,
UG, UT, UTG, WTU! Distributed as E. ovalifolium Nutt. var.
nivale (Canby in Gov.) M. E. Jones.

Additional Specimens Examined. — Head of Pine Greek Can-
yon, Toquima Range, 11,000 feet elevation, 16 July 1945, Maguire
& Holmgren 25805 (NY, US, UTG).

This lovely addition to Eriogonum ovalifolium has been known
to me for several years, but its distinctiveness has not become obvious
until the late 1960s. At first, the yellow flowers were considered to
be a common feature of the Nevada populations of E. ovalifolium
var. nivale, but the fact was that no other mountain ranges harbored
the yellow-flowered phase. This realization came only after most
of the high mountain ranges of central Nevada were carefully sur-
veyed as part of the Intermountain Flora Project. Thus, the To-
quima Range has another endemic species and can join Astragalus
toquimanus Bameby, Draba arida G. L. Hitchc, Smelowskia holm-
grenii Roll., and an as yet undescribed Geranium.

Eriogonum umbellatum Torr. var. devestivum Reveal, var. nov.
A var. stellata (Benth.) M. E. Jones foliis late ellipticus et omnibus
glabris vel marginibus non nisi pubescentis differt.

Plants forming a large open mat up to 6 dm across; leaves in
loose, ± basal rosettes, the leaf-blade broadly elliptic, 1.5-2 cm
long, glabrous on both surfaces or with the margin slightly pubes-
cent in some, on a long slender petiole; flowering stems erect, slen-
der, 1.5-2 dm long, thinly floccose to glabrous; inflorescences com-
poundly umbellate, 5-15 cm long with foliaceous bracts at the base
of each division, the branches thinly floccose or more commonly
glabrous; peduncles slender, 0.5-2 cm long, essentially glabrous, in-
volucres with tubes 2-3.5 mm long, the reflexed lobes 1-2.5 mm long,
glabrous; flowers bright yellow, 4-7 mm long including the stipe.

Type. — On sagebrush covered hills 10 miles south of Council
along U.S. Highway 95, Adams Co., Idaho, 21 June 1966, Holm-
gren & Reveal 2746. Holotype, US! Isotypes, BRY, CAS, NY, RSA,
UTG, WTU!

116 GREAT BASIN NATURALIST Vol. 32, No. 2

Distribution. — Locally common and sporadic from Adams and
Washington cos. eastward to Blaine Co., Idaho. Flowering from
June to August.

The var. devestivum forms a similar aspect of the species expres-
sion as seen between var. umbellatum and var. aureum (Gand.) Re-
veal. The leaves are totally glabrous (the margins excepted in some
cases). However, unlike the relationship between var. umbellatum
and var. aureum. in which the latter simply replaces the former at
higher elevations throughout much of the range of var. umbellatum,
there are a number of minor morphological differences between var.
stellatum. (Benth.) M. E. Jones and var. devestivum. The most ob-
vious differences are the highly branched inflorescences of the new
variety, the wider leaves, and the longer petioles. While the entire
range of the new entity remains to be fully ascertained, it appears
that var. devestivum occurs a bit to the east of the majority of the
populations of var. stellatum.^ although its stills forms a pocket with-
in the overall distribution of var. stellatum.

Acknowledgment

Field work was conducted under the sponsorship of a National
Science Foundation grant to Arthur Cronquist for the Intermountain
Flora Project. Herbarium studies were conducted, in part, under the
auspices of National Science Foundation grant GB-22645 for studies
of the genus Eriogonum. and of the Intermountain Flora.

Literature Cited

Stokes, S. 1936. The genus Eriogonum. San Francisco.

PLANTS NEW TO THE UTAH FLORA

Larry C. Higgins^

Abstract. — New additions to the Utah flora include Cryptantha dumetorum
Greene, C. data (Eastwood) Payson, C. longiflora (Nels.) Payson (Boraginaceae) ,
Hulsea heterochroma A. Gray. Parthenium incanum H. B. K. (Compositae),
Arctostaphylos pringlei C. C. Parry (Ericaceae), Eremocarpus setigerus (Hook.)
Benth., Euphorbia micromera Boiss, Tragia ramosa (Euphorbiaceae), Leptochloa
filiformis (Lam.) Beauv., Schismus arabicus Nees. (Gramineae), Epilobium
nevadense Munz (Onagraceae), Lomatium megarrhizum (A. Nels.) Mathias
(Umbelliferae), and Aloysia wrightii (Gray Heller) Verbenaceae.

Collections over the last six years provide the following additions
to the Flora of Utah and Nevada (Ivar Tidestrom, 1925) and Com-
mon Utah Plants (Welsh, Treshow, and Moore, 1965). Voucher
specimens are deposited in the herbaria of Brigham Young Uni-
versity (BRY) and West Texas State University (WTSU).

Boraginaceae

Cryptantha dumetorum Greene, Bush-loving cryptantha. Wash-
ington Co.: black stationary sand dunes just southwest of Ivins Res-
ervoir; associated with Coleogyne and Hymenoclea; 17 May 1971.
L. C. Higgins 4130 (BRY, WTSU).

Cryptantha elata (Eastwood) Payson, Tall cryptantha. Grand
Co.: near milepost 39 along Utah hwy 128; clay soil; Atriplex com-
munity; 3 June 1968. L. C. Higgins 1479 (BRY, WTSU). Atriplex
community; along hwy 124, ca. 3 miles south of U.S. hwy 50-6; on
mancos shale; 2 May 1968. S. L. Welsh 6952 (BRY).

Cryptantha longiflora (Nels.) Payson, Long-flowered cryptan-
tha. Grand Co.: near milepost 32 along Utah hwy 128; sand-loam
soil; associated with Populus and Atriplex; 3 June 1968. L. C. Hig-
gins 1478 (BRY, WTSU). Near Cisco, along U.S. hwy 6-50, clay
soil; Atriplex community; 3 June 1970. L. C. Higgins 3314 (BRY,
WTSU). Morrison formation; along Colorado River, ca. 32.5 miles
from Moab along Utah hwy 128; 3 May 1968. S. L. Welsh 6989
(BRY).

Compositae

Hulsea heterochroma A. Gray, Various-colored hulsea. Wash-
ington Co.: summit of the Beaverdam Mountains, ca. 3 miles south-
east of the television relay tower; limestone soil; pinyon- juniper
community; 28 May 1968. L. C. Higgins and N. D. Atwood 1410
(BRY).

Parthenium incanum H.B.K., Mariola. Washington Co.: Beaver-
dam Mountains, at Castle Cliffs along U.S. hwy 91; rocky, south-

'Department of Biology, West Texas State University, Canyon, Texas 79015.

117

118 GREAT BASIN NATURALIST Vol. 32, No. 2

facing limestone slope; associated with Buddleja-Agave-Larrea-Psilos-
trophe; 17 May 1971. L. C. Higgins 4162 (BRY, WTSU).

Ericaceae

Arctostaphylos pringlei C. C. Parry, Pringles manzanita. Wash-
ington Co.: near the television relay tower at the summit of the
Beaverdam Mountains; associated with Quercus-Pinus-Juniperus;
limestone soil; 27 May 1966. L. C. Higgins 665, 3414 (BRY).

Euphorbiaceae

Eremocarpus setigerus (Hook.) Benth. Turkey mullein. Wash-
ington Co.: Beaverdam Mountains, near the Utah- Arizona border
along U.S. hwy 91; sandy soil; associated with Hymenoclea-Fran-
seria-Larrea; 29 November 1968. L. C. Higgins 1809 (BRY).

Euphorbia micromera Boiss. Washington Co.: Beaverdam Moun-
tains, near the Utah-Arizona border along U.S. hwy 91; sandy
soil; associated with Hymenoclea-Larrea-Franseria; 29 November
1968. L. C. Higgins 1802 (BRY).

Tragia ramosa Muell. Arg. Nosebum. Washington Co.: Beaver-
dam Mountains, near summit along U.S. hwy 91; gravelly wash
bottom; J uniperus-Cercocarpus-Purshia community; 15 May 1966. L.
C. Higgins 602 (BRY).

Gramineae

Leptochloa filiformis (Lam.) Beauv. Sprangle-top. Washington
Co.: Beaverdam Mountains, 1 mile west of U.S. hwy 91 along the
Jackson road; sandy soil; Larrea-Prosopis-Coleogyne community;
19 August 1966. L. C. Higgins 823 (BRY).

Schismus arabicus Nees. Arabian schismus. Washington Co.:
Beaverdam Mountains, near the Utah-Arizona border along U.S.
hwy 91; gravelly soil; Larrca-Franseria-Yucca community; 8 April
1966. L. C. Higgins 331 (BRY).

Onagraceae

Epilobium nevadense Munz, Nevada willow herb. Washington
Co.: summit of the Beaverdam Mountains near the television relay
tower; westfacing, rocky, limestone hillside; pinvon-juniper-oak
association; 9 July 1966. L. C. Higgins 778 (BRY).'

Umbelli ferae

Lomatium megarrhizum (A Nels.) Mathias, Large-rooted lo-
matium. Emery Co.: San Raphael Swell, about 1 mile south of
the San Raphael River Bridge; clay hillside; desert scrub and
Bouteloua gracilis community; 30 April 1968. L. C. Higgins and
J. Reveal 1268 (BRY, WTSU).

June 1972 higgins: new utah plants 119

Verbenaceae

Aloysia wrightii (Gray) Heller, Oreganillo. Washington Co.:
Beaverdam Mountains, about 5 miles west of Castle Cliffs along
the road to Terry's Ranch; rocky limestone cliffs; associated with
Ferocactus-Coleogyne-Larrea; 26 May 1966. L. C. Higgins 615
(BRY). Rare, only 4 to 5 shrubs were seen in the entire area.

Literature Cited

TiDESTROM, I. 1925. Flora of Utah and Nevada. Verlag Von J. Cramer.
Welsh, S. L., M. Treshow, and G. Moore. 1965. Common Utah Plants.
Brigham Young University Press.

NOTES ON NEVADA SOLPUGIDS

Dorald M. Allredi

Muma (1963, Brigham Young Univ. Sci. Bull., Biol. Ser. 3 (2);
1970, Florida Dept. Agr. Contr. No. 154) listed distribution
records of solpugids for the Nevada Test Site and other parts of
North America, Central America, and the West Indies. In a series
of solpugids collected at the Nevada Test Site subsequent to his
1963 report are two specimens which represent records worthy of
mention.

In his 1963 report Muma recorded a single male of Hemero-
trecha bronchi taken at the Nevada Test Site, collected in Area 1
in a Grayia-Lycium plant commimity, 28 June 1960. An additional
specimen was taken at Cane Springs at the test site, 12 June 1965.
According to Muma (1970) this species also occurs in Arizona,
California, and New Mexico.

Muma (1970) listed Hemerotrecha jacintoana as known only
from California. A specimen was taken in a Coleogyne plant com-
munity approximately 2400 m north of the Sedan project crater at
the Nevada Test Site, 22 June 1964; this extends the known range
of the species to Nevada.

^Department of Zoology, Brigham Young University, Prove, Utah 84601.

120

GREAT BASIN NATURALIST
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TABLE OF CONTENTS

Prey stage distribution, a factor affecting the numerical
response of Typhlodromus occidentalis to Tetranychus
mcdanieli and Tetranychus pacificus. B. A. Croft 61

A revision of the Psoralea lanceolata complex. C. A. Toft

and S. L. Welsh 76

Observations on the nesting biology of the long-billed curlew.

Dennis M. Forsythe 88

Vegetation zones around a small pond in the White Moun-
tains of Arizona. B. Ira Judd 91

A new species of Gymnodamaeus from Colorado (Acarina:

Cryptostigmata, Gymnodamaeidae) . Tyler A. Woolley .... 97

The Sonoran subspecies of the lizard Ctenosaura hemilopha.

Hobart M. Smith 104

Dead great blue heron found at 11,000 foot Utah Elevation.

Gary L. Hickman 112

Five new varieties of Eriogonum (Polygonaceae). James L.

Reveal 113

Plants new to the Utah flora. Larry C. Higgins 117

Notes on Nevada solpugids. Dorald M. Allred 120

Volume 32, No. 3
MU'^. CO MP. ZOOL. Sept. 30, 1972

LIBRARY

The

0€C 21972

HARVARD
UNIVERSITY

Great Basin

nmmm

Published by
Brigham Young University

GREAT BASIN NATURALIST

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

Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W.
Tanner, Zoology; Joseph R. Murdock, Botany; Vernon J.
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Ex officio Editorial Board Members:

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The Great Basin Naturalist was founded in 1939 by Vasco M.
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a year since then by Brigham Young University, Provo, Utah. In
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ing to tiie biological natural history of the Great Basin and western
North America will be accepted. Manuscripts are subject to the ap-
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The Great Basin Naturalist

Published at Provo, Utah, by

Brigham Young University

Volume 32 September 30, 1972 No. 3

ON THE RODENT-INFESTINC, ANOPLURA OF PANAMA

Phyllis T. Johnson'

Abstract. — The species of Anoplura found on indigenous Panamanian
rodents belong to the genera Hoplopleura (8 species), Fahrenholzia (3 species),
N eohaematopinus (1 species), and Polyplax (1 species). As is the case with the
Panamanian rodents, the anopluran fauna contains both Nearctic and Neotropical
elements. Two new species of Hoplopleura are described: H. scodnomydis from
Scotinomys xerampelinus, and H. mendezi from Oryzomys sp. (probably O.
alfaroi or O. albigularis) . A host-parasite list is included.

Panama, the land bridge between North and South America,
has an unusually rich fauna whose study is important to an under-
standing of the movements and distribution of the animals of both
continents, as Fairchild (1966) pointed out in his introduction to
Ectoparasites of Panama. As well as for its intrinsic value, a study
of the Panamanian anopluran fauna can be a useful adjunct to the
study of movements and relationships of their mammalian hoSvts.

A preliminary paper on the Anoplura of Panama was published
in Ectoparasites of Panama (Wenzel and Johnson, 1966). Since
1966, I have had the opportunity to study the numerous specimens
collected during the surveys mentioned in the above volume. The
present paper is based on those studies. Species discussed by Wen-
zel and Johnson, for which there is no additional information, are
merely enumerated in the "Host-Parasite List," which is based on
the present survey as well as on the records of Wenzel and John-
son. Place names and the species names of mammals mentioned
herein may be traced in Fairchild and Handley (1966) and Hand-
ley (1966), respectively. The species groups of Hoplopleura re-
ferred to in this paper are discussed by Johnson (in press).

The holotypes and allotypes of new species, and the majority
of other specimens are deposited in the collections of the National
Museum of Natural History, Smithsonian Institution, Washington,
B.C.

Hoplopleura Enderlein

The hesperomydis group.

There are two, perhaps three, Panamanian representatives of
this group. All the known species occur on mice of the New

'National Marine Fisheries Service, Oxford Laboratory, Oxford, Maryland 2l(i54.

121

122 GREAT BASIN NATURALIST Vol. 32, No. 3

World tribe, Peromyscini. Characters held in common by members
of the hesperomydis group are given below:

Adults. — Head lacking accessory dorsal seta (ADHS of Kim,
1965). All abdominal setae inserted on posterior margins of sternal
and tergal plates; para tergal plates III- VI with two truncate apical
lobes; plate III with two large apical setae; plates IV-VI with dorsal
apical seta small to minute and ventral one longer, though seldom
surpassing apices of apical lobes. Females with two apical lobes on
plate VII; aberrant males of described species occasionally with
ventral lobe missing or vestigial; this lobe always missing in males
of a new species described below. Thoracic sternal plate elongate;
posterior apex narrowed, usually acute.

Nymphs. — Lacking abdominal spiracles and paratergal or
other abdominal plates. First instar with one terminal abdominal
seta on each side; second and third instars with one pair plus one
single long terminal seta on each side. All instars with accessory
dorsal head seta absent; one of dorsolateral head setae often en-
larged (outer sutural head setae of Kim, 1965).

Characters that are too variable to assist greatly in species iden-
tification within the hesperomydis group are as follows: shape of
the male genitalia, which depends on position; placement of setae
on the lateral postantennal head margins, which depends to some
extent on the amount of flattening during mounting; and shape of
the apical lobes and length of setae of the paratergal plates. Shape
of the thoracic sternal plate and presence or absence of a definite
mesal keel on its posterior apex may be good characters for
determination of some species. According to the statistical analysis
of Kim et al. (1966), relative length and breadth of the thoracic
sternal plate and total body length of the male are reasonably
good discriminatory characters for males of two species, ferrisi
Cook and Beer, and hesperomydis (Osborn). However, if a male
specimen is not completely extended, measurement of body length
is useless, and the reproductive state of a female may make a con-
siderable (hfference in total length. In most cases, as suggested
by Cook and Beer (1959) and Kim (1965), nymphs will be neces-
sary for a definitive study of the hesperomydis group.

HoplopU'ura emphereia Kim
Figs. 1, 7, 10, 14
Hoplopleura ferrisi emphereia Kim, 1965, J. Parasit. 51:882, figs. 37-42.

Hoplopleura hesperomydis: Wenzel and Johnson, 1966, Ectoparasites of Panama,
p. 274 (err. det., records from Peromyscus nudipes, and probably that from
Reithrodontomys sp.).

The female holotype, male allotype, 6 male and 20 female para-
types were taken from Peromyscus nudipes. El Hato, Province of
Chiriqui, Panama, and 1 male and 2 female paratypes from P.
guatamalensis and P. mexicanus saxatilus, Guatemala.

Sept. 1972 JOHNSON: anoplura of Panama

123

Figs. 1-8. Hoplopleura hesperomydis-group: 1, paratergal plates II-VIII,
H. emphereia Kim, female ex Peromyscus nudipes, no. 10247; 2, same, H. scoti-
nomydis, n. sp., female paratype ex no. 10245. same, male paratype ex no.
10155; 4, same, H. hesperomydis (Osborn), female ex Peromyscus leucopus. New
Hampshire; 5, aedeagus, H. hesperomydis ex Peromyscus maniculatus, Washing-
ton; 6, same, H. reithrodontomydis Ferris (?) ex Reithrodontomys megalotis,
California; 7, same, H. emphereia ex Peromyscus nudipes, Chiriqui, Panama; 8,
same. H. scotinomydis, n. sp., holotype.

124 GREAT BASIN NATURALIST Vol. 32, No. 3

Panamanian specimens. — From P. nudipes, 145 males, 248 fe-
males, and 1 third instar nymph (female), in 52 collections; most
from near or at the type locality. A few specimens were taken in
the Province of Bocas del Toro.

Comparison of emphereia and hesperomydis. — The adult of
emphereia differs as follows: thoracic sternal plate broader in pro-
portion to length, slightly bulbous laterally, almost always lacking
indication of mesal posteroapical keel (Figs. 12, 14). Usually larger
{hesperomydis, 8 males and 16 females measured: male, 0.8 to 0.91
mm; female, 1.17 to 1.20 mm; emphereia, 20 males and 20 females
measured: male, 0.95 to 1.10 mm; female, 1.15 to 1.25 mm). The
dorsal apical lobe of paratergal plate VII is always subtruncate, with
oblique, sometimes notched, apex (hesperomydis with this lobe
usually evenly acute apically) (Figs. 1,4). Aedeagus somewhat larg-
er; often with parameres more convex laterally; pseudopenis more
angulate laterally (Figs. 5, 7). Placement of small laterodorsal
head setae and relative lengths of apical setae of paratergal plates
variable in specimens examined.

Status of the name emphereia Kim. — The third-instar n\TTiph
from P. nudipes is unlike the nymph of ferrisi Cook and Beer, thus
the name emphereia does not pertain to the latter species.

Kim (1965) resurrected the name reithrodontomydis Ferris,
1951, on the basis of a study of 8 nymphs collected from Reithro-
dontomys sp. in Chiriqui Province at El Hato, by Tipton. I have
seen a female from Reithrodontomys sp. with collection data essen-
tially as the nymphs studied by Kim, plus 13 males and 19 females
in 14 collections from Reithrodontomys creper, and 3 males and 7
females from R. mexicanus, Chiriqui, 3 miles (5 km) W Cerro
Punta, March, 1962. I did not find consistent differences between
adults in the population from Reithrodontomys species, and that from
Peromyscus nudipes. Further, my third instar n\Tnph from P.
nudipes (Fig. 10) appeared similar to Kim's (1965) drawings of
a nymph from Reithrodontomys sji. except that the head capsule
had been split and the mechodorsal portion, which would have con-
tained the large dorsomedian seta figured by Kim, was missing. Be-
cause of the fully developed adult abdomen within, small ventral
abdominal setae were not visible. One cannot, without furth(*r speci-
mens, state unequivocally thai emphereia from P. nudipes is con-
specific with the form from Panamanian Reithrodontomys species,
but it is highly probable and I make that assumption here.

As a further complication, I have seen a male and female of
Hoplopleura from Reithrodontomys megalotis, California, that are
smaller than the Panamanian specimens from Reitlirodontomys
species (male, 0.85 mm; female, 1.08 nmi) and the thoracic sternal
plate (Fig. 13) and aedeagus (Fig. 6) are more like those of true
hesperomydis from North American Peromyscus maniculatus and
P. leucopus than those of Panamanian s{)ecies from P. nudipes and
Reithrodontomys species.

Sept. 1972 JOHNSON: ANOPLURA OF PANAMA 125

If all the above specimens are of the same species and prove to
be the same as the population from Reithrodontomys chrysopsis
that was originally given the name reithrodontomydis Ferris, this
name has priority over emphereia Kim.

Hoplopleura scotinomydis, n. sp.

Figs. 2, 3, 8, 9, 11, 15-18

Type data. — Male holotype, female allotype, 1 male and 2 fe-
male para types ex Scotinomys xerampelinus, Panama: Province of
Chiriqui, Boquete Trail, 3 miles (5 km) W Cerro Punta, 7700 ft
(2700 m), 9-III-1962, PMMC 10275. Additional paratypes, all
from Scotinomys xerampelinus. Province of Chiriqui, collected by
the Preventive Medicine and Malaria Control Unit, U.S. Army, as
follows: 1 female, Martinez Dairy, Cerro Punta, 6800 ft (2400 m),
8-1-1966, no. 6862. A series from Boquete Trail, 3 miles (5 km) W
Cerro Punta, 6900 to 7800 ft (2400 to 2700 m), 7-14-III-1962,
as follows: 3 males, 7 females, plus 1 third-stage nymph, no. 10155;

1 female, no. 10172; 1 male, 1 female, no. 10238; 4 females, no.
10245; 2 males, no. 10253; 1 male, plus 1 third-stage nymph, no.
10257; 1 female, plus 1 third-stage nymph, no. 10258; 1 female, no.
10260; 2 males, 1 female, no. 10276; 1 female, no. 10277; 1 male,

2 females, no. 10405; 1 male, 2 females, no. 10406; 1 female, no.
10424; 5 males, 5 females, no. 10454; 1 male, no. 10481; 2 females,
no. 10492.

Diagnosis. — A member of the hesperomydis group. Separable
by having apical lobes of paratergal plate II both short, not with
ventral greatly prolonged (Fig. 2) and in lacking sword-shaped
setae on abdomen. Female further separable by having genital seta
rather short and blade- shaped (Fig. 11). Male further separable
by having only one apical lobe on paratergal plate VII (Fig. 3)
(occasional males of related species also lack the ventroapical lobe
of this plate) .

Lengths. — Male: holotype, 1.0 mm; paratype, 0.95 to 1.10
mm. Female: allotype, 1.45 mm; paratype, 1.30 to 1.45 mm.

Description. — Male (Fig. 16). Head (Fig. 17). First antennal
segment not particularly enlarged, third segment not modified; post-
antennal margins slightly rounded, converging somewhat posteriorly;
accessory dorsal seta missing. Thorax. Sternal plate (Fig. 15) broad,
posterior apex bluntly rounded, short and broad, lacking a mesal keel;
plate less than one and one-half times as long as broad. Dorsal meso-
thoracic seta long. Abdomen. All setae long, flexible, none sword-
shaped; tergal and sternal plates well developed, all setae confined
to their posterior margins; lacking tergal plate and setae on seg-
ment 1. Paratergal plates (Fig. 3) II with short acute apical lobes
of similar length and two stout setae extending beyond apices of
lobes; plates III-VI with equal, scaly, quadrate, apical lobes; plate
VII with short triangular dorsal lobe; plate VIII lacking apical

126

GREAT BASIN NATURALIST

Vol. 32, No. 3

Figs O-l"). Honlopleura hesperomrdis-Krouii: 9, H. scotmomydis. n. sp.,
third-stage nymph ex no. 10155; 10, H. emphereia Kim third-stage "ymph ex
PeromyTcus nudipes, Chiriqui, no. 10254. Note that the dorsomedian part of the
head is missing; 11. //. scotinomydis, n. sp., female genitalia, allotype; 12, thoracic
sternal plate, H. hesperomydis (Osborn) ex Peromyscus mamculatus.a, male,
Washington; b, female, Washington; c, female, Utah; 13, same, A/, reithrodonto-
mydis Ferris (?), California ex Reithrodontomys megalotis: a, female; b, male;
14, same, H. emphereia Kim, Chiriqui ex Peromyscus nudipes no. 6814: a. male;
b, female; 15, same, H. scotinomydis, n. sp.: a. holotype; b, allotype.

Sept. 1972 JOHNSON: anoplura of Panama 127

lobes; plate III with two stout apical setae extending about to apices
of lobes; IV-VI with small to minute ventral apical seta and stout
dorsoapical seta extending almost to apices of lobes. Aedeagus. (Fig.
8) with parameres somewhat broadened basally, lateral outlines
convex; pseudopenis with elongate apex.

Female (Fig. 18). Head and thorax as in male. Abdomen as
in male except for usual sexual dimorphism. Apical setae of para-
tergal plates (Fig. 2) often relatively shorter than in male; plate
VII with two narrow apical lobes, the dorsal one the longer, obliquely
truncate; ventral lobe acute. Genital seta shorter than in related
species, blade-shaped (Fig. 11).

Nymph (Fig. 9). Third instar. Head, scaly dorsally, usual
ventral tubercles; principal dorsal seta short, stout; other head setae
minute. Dorsal mesothoracic seta about same size and shape as princi-
pal dorsal head seta. Four minute setae ventrally on abdomen. Ter-
minal setae of abdomen consisting of a pair and single on each side,
all these short, stout; anal lobe extended, short, bluntly rounded.

The travassosi group.

H. travassosi Werneck and allies are typical parasites of South
American rodents. Species of this group are found throughout that
continent. The northernmost representatives known are H. angulata
Ferris, from Panama, and H. similis Kim, which is found in Panama
and as far north as tropical Mexico.

Hoplopleura angulata Ferris

Hoplopleura angulata Ferris, 1921, Stanford Univ. Publ. Biol. Sci. 2:73, Figs.
40, 41A-C, E; Johnson, in press, Brigham Young Univ.. Sci. Bull., Biol. Ser.
17(3), Figs. 58-69.

The type series was from Rhipidomys venezuelae, Venezuela.
This louse species is common on several species of Rhipidomys in
Venezuela, and also occurs on species of Rhipidomys in Peru, Co-
lombia, and Trinidad.

Panamanian specimens. — Two males from the rare Nyctomys
sumichrasti. Province of Chiriqui, Cerro Punta. Occurrence of angu-
lata on a mouse other than Rhipidomys is a surprising finding con-
sidering the rigid host specificity of this species in Venezuela.

Hoplopleura similis Kim

Figs. 19-21

Hoplopleura similis Kim, 1965, J. Parasit. 51:884, Figs. 47-53.

The female holotype and a series of female paratypes were
from Oryzomys fulvescens, Veracruz, Mexico. The male allotype
was from Bolivian Oryzomys chaparensis.

Panamanian specimens. — From Oryzomys fulvescens: 1 female,
Province of Bocas del Toro, Rancho Mojica, 500 ft (1700 m), 8-IX-

128

GREAT BASIN NATURALIST

Vol. 32, No. 3

1961, no. PMMC 8017; 1 female, Province of Chiriqui, Boquete
Trail, 3 miles (5 km) W Cerro Punta, 7750 ft (2700 m), lO-III-

1962, no. PMMC 10314; 4 nymphs (first and third instar). Province
of Chiriqui, 3 miles (5 km), W Cerro Punta, 7800 ft (2700 m),
7-III-1962, no. PMMC 10177.

Figs. 16-18. Hoplopleura scotinornydis,
type; 18, allotype.

sp.: 16, holotype; 17, head, holo-

Sept. 1972

JOHNSON: ANOPLURA OF PANAMA

129

Figs. 19-23. Hoplopleura species: 19, H. similis Kim third-stage nymph; 20,
same, first-stage nymph; 21, same, paratergal plate VII, f«"?«l«^,«' ^^^^ "°;,^«^^i^d'
b, ex no. 10314; 22, H. mendezi, n. sp., third-stage nymph; 23, same, second-
stage nymph.

The Panamanian females are like the holotype except the
ventral apical lobe of paratergal plate VII is very short (Fig. 21).
The nymph has not been described previously. It is very like that

130 GREAT BASIN NATURALIST Vol. 32, No. 3

of travassosi except the dorsal head setae are less thornlike. The
third instar lacks the dorsal mesothoracic seta. If the seta is missing,
not broken off, this is a major difference between travassosi and
similis.

Description. — Nymph. Abdomen of both first and third instar
faintly reticulate; thoracic cuticle and anterior ventral abdominal cu-
ticle spiculate. First instar (Fig. 20) with one pair of long terminal
abdominal setae on each side; anal lobe somewhat extended; head
with usual tubercles ventrally, principal dorsal head seta well de-
veloped, its accessory seta short, stout. Other dorsal setae of head
small, not thornlike. Dorsal mesothoracic seta very long. Third
instar (Fig. 19). Head much as in first instar but principal dorsal
seta relatively shorter. Dorsal mesothoracic seta aparently not pres-
ent. Two pairs and a single terminal abdominal seta on each
side.

The quadridentata group.

As is true of the travassosi group, species of Hoplopleura re-
lated to quadridentata (Neumann) are typically Neotropical. How-
ever, distribution of known species of the group is generally more
northern. In South America true quadridentata-group species ap-
parently do not occur south of the tropical regions, while to the
north one species, oryzomydis Pratt and Lane, is common on rice
rats {Oryzomys palustris) in the southern United States. In Panama
there are three representatives of the quadridentata group, one
of them new.

Hoplopleura nesoryzomydis Ferris

Hoplopleura nesoryzomydis Ferris, 1921, Stanford Univ. Publ., Biol. Sci. 2:90,
Fig. 53A; Wenzel and Johnson, 1966, Ectoparasites of Panama, p. 275.

Panamanian specimens. — From Zygodontomys niicrotinus, 9
males, 11 females, 2 nymphs, taken in 13 collections. Canal Zone
and Azuero Peninsula. Also present in 2 collections from Oryzomys
capito talamancac. Canal Zone (4 males, 3 females, 15 nymphs)
and Cerro Punta, Chiriqui (1 female, 1 nymph).

Hoplopleura oryzomydis Pratt and Lane

Figs. 27, 31-34

Hoplopleura oryzomydis Pratt and Lane, 1951, J. Parasit. 37:141, Figs. 1-3, 5;
Wenzel and Johnson, 1966, Ectoparasites of Panama, p. 275. Johnson, in
press, Brigham Young Univ. Sci. Bull., Biol. Ser. 17(3), Figs. 129-137.

The type series of oryzomydis was taken from Oryzomys palustris,
Florida and South Carolina. Venezuelan specimens from Nectomys
squamipes are similar to the North American form (Johnson, in
press).

Panamanian specimens. — One pair from Oryzomys caliginosus.
El Valle; 1 pair from Oryzomys capito. Province of Darien, Santa
Fe, no. OICS 2286.

Sept. 1972

JOHNSON: ANOPLURA OF PANAMA

131

Figs 24-28. Hoplopleura quadridentata-grouiK 24 H. ^^"^^f ' ""^ . f g'
capita, Darien, Panama; 28, H. mendezi, aedeagus, holotype.

132 GREAT BASIN NATURALIST Vol. 32, No. 3

Morphological differences among the populations: paratergal
plates of Panamanian specimens scalier than those of types. Thoracic
sternal plate (Fig. 31) somewhat broader than in U.S. specimens;
more like that of Venezuelan series. Aedeagus (Fig. 27) similar to
types. Abdominal tergal and sternal plates narrower than in
Venezuelan and U.S. specimens, approaching obsolescence. Penul-
timate tergal plate of female smaller than in U.S. specimens, but
like them in having two rather than three or four apical setae
(compare Figs. 32-34).

Hoplopleura mendezi, n. sp.
Figs. 22-26, 28-30

Type data. — Male holotype, female allotype, 2 male paratypes
from Oryzomys sp., Province of Chiriqui, Bambito, El Hato, 15-T
1961, Keennn and Wislocki collectors, RML 40451. One female
para type as above but Chiriqui Viejo, Casa Shannon, 17-T1961,
Wislocki collector, RML 40450. Also examined: 2 males, 1 female,
3 nymphs (second and third instar), lacking data but possibly RML
no. 40445.

According to Handley (1966) the type host could be either
Oryzomys alfaroi, which is rare, or Oryzomys albigularis. Both
these species are found at the type locality. A third species, Ory-
zomys fulvescens, occasionally occurs at high elevations like the
type locality, in this area of Panama, but as it carries Hoplopleura
similis Kim, there is less likelihood that it is the type host of
mendezi.

Diagnosis. — A member of the quadridentata group. Adult
separable by lacking long setae on paratergal plate IIL Nymph dif-
fers (from others known) by having antennal segments 3 to 5
incompletely separated; and further from nesoryzomydis by having
the small dorsolateral head setae minute.

Lengths. — Male: holotype, 1.0 mm; paratypes 0.95 to 1.05
mm. Female: 1.3 mm.

Description. — Male (Fig. 24). Head (Fig. 26). Postantennal
angles well developed; antennae unmodified, first segment not par-
ticularly enlarged; accessory dorsal seta present. Thorax. Sternal
plate (Fig. 30) elongate, squared apically. Dorsal mesothoracic seta
long. Abdomen, l^rgal and sternal plates arranged as usual; seg-
rnent 1 with indistinct tergal plate and one minute seta to either
side of the plate. Typical abdominal plates well developed, their
posteroapical setae long, straight, slightly sword-shaj)ed, some setae
off plates dorsally and ventrally. Paratergal plates (Fig. 29, fe-
male) IH-VI each with two deeply subdivided, rounded apical
lobes; VII with truncate, divided dorsal lobe and lacking ventral
lobe; VIII lacking apical lobes. Plate II with two long apical setae;
plates III-VI with apical setae missing, extremely minute, or minute
and removed from plate margin. Aedeagus (Fig. 28) with para-

Sept. 1972

JOHNSON: ANOPLURA OF PANAMA

133

Figs. 29-33. Hoplopleura quadridentata-group: 29, H. menAezi, n. sp., para-
tergal plates, female para type; 30, same, thoracic sternal plate, allotype; 31,
H. oryzomydis Pratt & Lane, thoracic sternal plate, ex Oryzomys capita, Darien,
Panama: a, male; b, female; 32, H. oryzomydis, penultimate abdominal tergal
plate, female, paratype; 33, same, ex O. capita, Darien; 34, same, ex Nectomys
squamipes, Venezuela.

meres not noticeably convex laterally, pseudopenis flared and ser-
rate laterally.

Female (Fig. 25). Head, thorax and abdomen as in male except
for usual sexual dimorphism. Paratergal plate VII with long
rounded-acute ventral apical lobe and truncate, apically divided
dorsal lobe. Genital seta long, not blade-shaped.

Nymph. Second and third instar similar except for size and the
fact that third instar (Fig. 22) has a pair plus a single terminal ab-
dominal seta on each side while the second instar (Fig. 23) has
only one terminal seta on each side. Anal segment slightly pro-
longed. Principal dorsal head seta well developed, its accessory
seta short but not minute; other dorsal setae small to minute.
Thorax with long mesothoracic seta; abdomen spiculate, lacking
plates or spiracles.

This species is named for my friend and colleague Eustorgio
Mendez, Gorgas Memorial Laboratory, Panama, in recognition of

134 GREAT BASIN NATURALIST Vol. 32, No. 3

his contributions to the taxonomy of the Mallophaga and Siphon-
aptera of South and Central America.

Fahrenholzia Kellogg and Ferris

Fahrenholzia ferrisi Werneck

Fahrenholzia ferrisi Werneck, 1952, Rev. Bras. Biol. 12:73, Fig. 7; Johnson, 1962,
Ann. Ent. Soc. Amer. 55:417, Figs. 10-13, 30, 37.

Fahrenholzia fairchildi: Wenzel and Johnson, 1966, Ectoparasites of Panama,
p. 275 (lapsus, specimens from Heteromys desmarestianus. Cerro Punta).

The type series was from Heteromys goldmanL Veracruz, Mex-
ico. This species also occurs on Heteromys sp., Guatemala.

Panamanian specimens. — From Heteromys desmarestianus, 20
collections (43 males, 57 females, 39 nymphs). Seventeen collec-
tions were from Province of Chiriqui, near Cerro Punta; two from
Boquete; one from Cerro Campana. Also examined: one nymph,
probably this species, from H. desmarestianus, Pina, Canal Zone.

It seems evident that F. ferrisi is the normal anopluran parasite
of Heteromys desmarestianus in Panama. See further comment be-
low.

Fahrenholzia fairchildi Johnson

Fahrenholzia fairchildi Johnson, 1962, Ann. Ent. Soc. Amer. 55:419, Figs. 18-21,
32, 39. Wenzel and Johnson, 1966, Ectoparasites of Panama, p. 275.

The type series was supposedly from Heteromys desmarestianus,
Santa Fe, Panama. Para types were from Liomys adspersus. Fort
Kobbe and Summit Road, Canal Zone. These were all collected in
1955-57. In the 1961-63 surveys, this species was taken only from
Liomys adspersus. There were 34 collections, including 29 males,
44 females, and 67 nymphs, Azuero Peninsula, Los Santos Province,
and a collection of 1 male and 2 nymphs from .Juan Mina, Canal
Zone. The collection data of the type collection probably were er-
roneous, since fairchildi was found consistently on Liomys in the
present series, while Heteromys desmarestianus carried only F.
ferrisi. Thus the normal host of fairchildi is Liomys adspersus.
Through a lapsus, specimens of ferrisi from //. desmarestianus,
Cerro Punta, were reported as fairchildi bv Wenzel and Johnson
(1966).

Polyplax Enderlein

Polyplax auricularis Kellogg and Ferris

Polyplax auricularis Kellogg and Ferris, 1915. Stanford Univ. Publ., Univ. Ser.,
p. 13, Fig. 4; pi. 4, Fig. 8. Ferris, 1921, Stanford Univ. Publ., Biol. Sci.
2:206.

The type series was from Peromyscus maniculatus, California.
This species is also found on other species of Peromyscus and from
Onychomys sp. in North America. It has been reported from
Reithrodontomys mexicanus and Neotomodon sp., Mexico.

Sept. 1972

JOHNSON: ANOPLURA OF PANAMA

135

Panamanian specimens. — From Reithrodontomys creper, Pro-
vince of Chiriqui, near Cerro Punta, 15 males, 19 females, 7 nymphs,
in 10 collections. P. auricularis often occurred together with Hoplo-
pleura emphereia Kim.

Panamanian specimens do not differ consistently from North
American individuals taken from species of Peromyscus, Neotoma,
and Onychomys. This aberrant species is the southernmost repre-
sentative of the genus Polyplax in the New World. Although its
northern populations apparently have a broad host range, in Pan-
ama— doubtless at the southernmost point of its range — auricularis is
associated only with Reithrodontomys creper.

Host-Parasite List

Order Rodentia
Family Sciuridae

Sciurus granatensis

Sciurus variegatoides

Family Heteromyidae
Liomys adspersus
Heteromys desmarestianus

Family Cricetidae
Oryzomys sp.
Oryzomys caliginosus
Oryzomys capita

Oryzomys julvescens
Nyctomys sumichrasd
Reithrodontomys creper

Reithrodontomys mexicanus
Peromyscus nudipes
Zygodontomys microtinus
Scotinomys xerampelinus
Sigmodon hispidus

Family Muridae
Rattus rattus

Family Echimyidae

Hoplomys gymnurus

Enderleinellus sp. (longiceps group)
N eohaematopinus semifasciatus F., s.
N eohaematopinus semifasciatus F., s.

Fahrenholzia fairchildi J.
Fahrenholzia ferrisi W.
Fahrenholzia hertigi J.

Hoplopleura mendezi, n. sp.
Hoplopleura oryzomydis P. & L.
Hoplopleura oryzomydis P. & L.
Hoplopleura nesoryzomydis F.
Hoplopleura similis K.
Hoplopleura angulata F.
Hoplopleura emphereia K.
Polyplax auricularis K. & F.
Hoplopleura emphereia K.
Hoplopleura emphereia K.
Hoplopleura nesoryzomydis F.
Hoplopleura scotinomydis, n. sp.
Hoplopleura hirsuta F.

Polyplax spinulosa (B.)
Hoplopleura audax F.

Literature Cited

Cook, E. F., and J. R. Beer. 1959. The immature stages of the genus Hoplo-
pleura (Anoplura: Hoplopleuridae) in North America, with descriptions of
two new species. J. Parasit. 45:405-416.

F.-^iRCHiLD, G. B. 1966. Introduction. In R. L. Wenzel and V. J. Tipton (eds.),
Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago, p. 1-7.

, AND C. O. Handley. 1966. Gazetteer. In R. L. Wenzel and V. J. Tip-
ton (eds.). Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago, p. 9-20.

Ferris, G. F. 1921. Contributions toward a monograph of the sucking lice.
Part II. Stanford Univ. Publ., Univ. Ser., Biol. Sci. 2:55-133.

Handley, C. O. 1966. Checklist of the mammals of Panama. In R. L. Wenzel
and V. J. Tipton (eds.). Ectoparasites of Panama. Field Mus. Nat. Hist.,
Chicago, p. 753-795.

136 GREAT BASIN NATURALIST Vol. 32, No. 3

Johnson, P. T. 1962. The species of Fahrenhohia Kellogg and Ferris from

spiny pocket mice (Anoplura: Hoplopleuridae) . Ann. Ent. Soc. Amer. 55:

415-428.
. In press. The sucking lice of Venezuelan rodents, with remarks on

related species (Anoplura). Brigham Young Univ. Sci. Bull., Biol. Ser. 17

(3).
Kellogg, V. L., .\nd G. F. Ferris. 1915. The Anoplura and Mallophaga of

North American mammals. Stanford Univ. Publ., Univ. Ser., p. 1-74.
Kim. K. C. 1965. A review of the Hoplopleura hesperomydis complex (Ano-
plura, Hoplopleuridae). J. Parasit. 51:871-887.
, B. W. Brown, and E. F. Cook. 1966. A quantitative taxonomic study

of the Hoplopleura hesperomydis complex (Anoplura, Hoplopleuridae), with

notes on a posteriori taxonomic characters. Syst. Zool. 15:24-45.
Pratt, H. D., and J. E. Lane. 1951. Hoplopleura oryzomydis new species, with

notes on other United States species of Hoplopleura (Anoplura: Haema-

topinidae). J. Parasit. 37:141-146.
Wenzel, R. L., and p. T. Johnson. 1966. Checklist of the sucking lice of

Panama. In R. L. Wenzel and V. J. Tipton (eds.). Ectoparasites of Panama.

Field Mus. Nat. Hist., Chicago, p. 273-279.
Werneck. F. L. 1952. Contribui^ao ao conhecrmento dos Anopluros. I. Rev.

Bras. Biol. 12:69-78.

NOTES CONCERNING MEXICAN SALDIDAE, INCLUDING
THE DESCRIPTION OF TWO NEW SPECIES (HEMIPTERA)

John T. Polhemus'

Abstract. — A complete description and discussion of the genus Enalosalda
Polhemus is given, and the males of E. mexicana Van Duzee and Saldula
hispida are described. Saldula saxicola and Saldula durangoana are described
as new. Saldula suttoni Drake and Hussey is transferred to loscytus (n. comb.);
Salda hispida Hodgden is considered a subspecies (n. comb.) of Saldula sulcicollis
Champion.

The new taxa and nomenclatural changes proposed here have
resuhed from a comprehensive study of Mexican Saldidae. As the
larger work may not be pubHshed for some time, it seems advisable
to make this information available to other workers.

The work upon which this paper is based was supported in part
by a grant from the University of Colorado Museum.

T\ie specimens utilized are primarily from the Polhemus collec-
tion (JTP) and the University of Colorado Museum Collections
(CU). A small amount of additional material was borrowed from
Texas A & M University (TAM), the California Academy of Sci-
ences (CAS), and the University of Michigan (UM); and I am
indebted to J. C. Schaffner, P. H. Arnaud, and T. E. Moore re-
spectively for their help in this regard.

The genus Enalosalda Polhemus was recently named in a paper
by Polhemus and Evans (1969); however, a complete description
and discussion were inappropriate in that paper and are given
below.

Enalosalda Polhemus
Enalosalda Polhemus, 1969, in Polhemus and Evans, Pacific Insects 11:575.

Hemelytra with membrane not differentiated from corium ex-
cept at inner margin; cell pattern variable, usually indistinct. Hind
wings reduced to small, membranous strips. Female ovipositor
short, broad, normally with six small teeth. Tube leading from
spermatheca bulb with tapering walls, thinning toward apex, with-
out flange. Larval organ absent. Ocelli, measured from center,
equidistant from each other and inner margins of eyes.

All other generic characters typical of the Chiloxanthinae (Cob-
ben, 1959), i.e., base of filum gonopori forming one closed ring,
median sclerotized structure of aedeagus paired, apicolateral struc-
ture of aedeagus absent, posterior margin of female subgenital plate
truncate with hind margin weakly indented, head without post-
clypeus. (The ductus ejaculatorius has not been studied.) This
genus falls in the subfamily Chiloxanthinae.

Type species: Orthophrys mexicanus Van Duzee, 1923.

'University of Colorado .Museum, Boulder. Colorado «05(L'.

137

138

GREAT BASIN NATURALIST

Vol. 32, No. 3

Discussion. — Van Duzee (1923) described mexicanus from a
single female taken under kelp on Angel de la Guardia Island in
the Gulf of California and placed it in the genus Orthophrys. Its
generic position was considered by Drake and Hoberlandt (1950),
who placed it in Pentacora, and later Drake and Hottes (1954)
figured the type and published a note on it. Recently, Lattin and
Cobben (1969) reexamined the type and tentatively assigned the
species to Pentacora., however, the rediscovery of the species at
Guaymas, Sonora, Mexico, has permitted the examination of a long
series, and I concluded that the taxon represents a distinct genus,
hence the name and a brief diagnosis were recently published
(Polhemus and Evans, 1969). Table 1 gives a comparison between
Enalosalda and the other genera in the Chiloxanthinae of which fe-
males were available. (Only a single male of Pelachoris leucographa
[Rimes] was studied, but it and Drake's description [1962] indi-
cate that Pelachoris is a fairly typical member of the subfamily.)

Enalosalda is divergent from other Chiloxanthinae, as it lacks
some of the specialized structures of Pentacora, Chiloxanthus, and
Paralosalda. The great reduction of flight (metathoracic) wings and
lack of larval organs could be interpreted as an indicator of the
age of split off of Enalosalda from the rest of the subfamily, which
is a fairly homogeneous unit with the exception of the annectant
Paralosalda. Flight wings are of high importance to many saldids,
and only those species occupying stable ecological situations over
a long time period are likely to lose the ability to fly. Even then,
many brachypterous forms retain relatively well-developed posterior

Table 1. Comparison of the genera of Chiloxanthinae.

Enalosalda

Spermatheca tube lead-
ing from bulb having
tapering walls.
Ovipositor short, having
six small teeth and no
well-developed ridges.

Flight (metathoracic)
wings vestigial, being re-
duced to small mem-
branous strips.
Female hemelytra with
only slight modification
to receive male coupling
plate, tlie modification
being a glabrous area
with a few erect hairs on
the underside of emboli-
um.

Larval organ absent.
Intertidal.

Paralosalda

Spermatheca tube lead-
ing from bulb having
tapering walls.
Ovipositor of moderate
length, having 8 well-
deveJoped teeth. Ridges
along ovipositor well de-
veloped.

Flight wings well devel-
oped.

Female hemelv-tra modi-
fied to receive male
coupling plate.

Larval organ present.
Intertidal.

Pentacora, Chiloxanthus

Spermatheca tube wdth
thickened structure
(flange) at terminus.
Ovipositor long, with 13-
14 well-developed teeth.
Ridges along ovipositor
well developed.

Flight wings more or
less well developed, near-
ly attaining apex of
hemelytra in all species.
Female hemelytra al-
ways modified to some
degree to receive male
coupling plate; at least a
thickening of the embo-
lar region.

Larval organ present.
Littoral or inland.

Sept. 1972 POLHEMus: Mexican saldidae 139

wings, although a reduction in hind wing structure, similar to that
in Enalosalda, is seen in some other saldids colonizing the ocean
shore (i.e., Orthophrys and Halosalda), and in the intertidal Omania
they are absent. The parallelism between Enalosalda and Orthophrys
has been discussed at length in another paper (Lattin and Cobben, loc.
cit.). The lack of a flange on the spermatheca is a character of un-
known significance; however, in his studies of the female genitalia of
the Heteroptera, Pendergrast (1957) stated that for the Saldidae a
single deep pump flange exists. The species that he studied were
Saldula saltatoria (L.) and Salda littoralis (L.). Concerning the
Heteroptera as a whole, he states that the spermatheca may possibly
be of use at the generic level of classification. The species studied
by this author in the Chiloxanthinae, other than Enalosalda^ all
possess the flange.

The reduced ovipositor structure, tending toward the mesoveliid
type, may have resulted from laying eggs in rock crevices or soft
material rather than in plant stems. While the eggs have not been
found, the habitat surrounding the intertidal rocks, in which these
bugs were studied, provided no visible terrestrial plant life that
could be used for egg deposition. Also, nymphs of almost all in-
stars were found in the intertidal zone.

I am indebted to G. G. E. Scudder for studying the female geni-
talia in detail. His figures (lA and B) are reproduced here, and
the nomenclature is that of Scudder (1959, 1961). His comments,
in part, are: "It seems to me that this is a fairly typical saldid,
both in external structure and detail. . . . The attachment of
the gonoplacs is a little different to that I have studied in Salda
and Saldula: The 'connecting piece' is quite free in your new genus
and not in Salda and Saldula.''

As the same condition of the "connecting piece" is found in
Pentacora signoreti Guerin, it is possible that this character may be
typical of the subfamily Chiloxanthinae; however, in Paralosalda it
is not free. The figure by Drake and Hottes (1954) does not
properly depict the antenna of Enalosalda mexicana, and as no male
has been available until the present series came to hand, a re-
description of the species follows. The type, a female, has been
studied.

Enalosalda mexicana (Van Duzee)

Orlhophrys mexicanus: Van Duzee, 1923, Proc. Calif. Acad. Sci. 12:165 (Type:

Puerto Refugio, Angel de la Guardia Island, Gulf of California, Mex.; Calif.

Acad. Sci. type no. 1045).
Pentacora mexicana: Drake and Hoberlandt, 1950, Acta Ent. Mus. Nat. Pragae

26:5.
Pentacora mexicana: Drake and Hottes, 1954, Occ. Papers Mus. Zool., Univ.

Mich. 553:5.
Pentacora mexicana: Lattin and Cobben, 1968, Ent. Berichten. 28:130.
Enalosalda mexicana: Polhemus and Evans, 1969, Pacific Insects 11:575.

Coloration. — Head, pronotum, scutellum, sutures of thoracic
venter, hemelytral markings and veins, dorsal surface of antennal

140 GREAT BASIN NATURALIST Vol. 32, No. 3

segment 1 and all of segments 3 and 4 brown to deep brown; small
spot either side of vertex of head, elongate area along inner margin
of each eye, medial spot on frons and apex of frons, rostrum, ante-
clypeus, labrmn, broad areas along lateral pronotal margins, nar-
row area caudad from collar, two triangular pronotal spots ad-
jacent to anterior femora, dorsal surface of middle and hind femora,
apices of tibia, tarsi ochreous; hemelytral ground color flavous to
ochreous; thoracic underparts leucine; legs, coxae, tibia luteous; an-
tennal segment 2 luteous, becoming lighter distally; femora luteous,
knees lighter.

Head. — Shining, frons and vertex rugulose. The elongate
ochreous areas along eyes interrupted by a dark brown sulcus start-
ing between ocelli and eye and extending anterolaterally almost to
eye, deeper ahead of ocelli; rostrum reaching between hind coxae;
ocelli equidistant from each other and eyes; with usual three pairs
of tricobothria.

Thorax. — Pronotum shining, rugulose; anterior lobe scarcely
raised, weakly differentiated from posterior lobe; with broad de-
pression medially and a depression on each side of middle caudad of
medial depression; lateral margins straight, angles rounded, anterior
width slightly less than eyes, posterior width/anterior width, 60/43;
posterior lobe/anterior lobe, 5/16; width/median length, 60/21;
posterior margin slightly indented over scutellum. Scutellum width/
length, 21/26.

Wings. — Hemelytra semi-brachypterous; broad, irregular spots
along margin two-fifths and three-fifths of the distance from base
to apex, a smaller spot four-fifths toward apex, irregular spots on
inner corium one-third and two-thirds toward apex, and at apex,
irregular, elongate brown areas along vestigial claval suture on both
inner corium and clavus and along margin of scutellum on clavus;
clavus fused to corium, vestigial suture ochreous; membrane largely
wanting, existing narrowly from apex of claval commissure to apex
of hemelytra, remainder of hemelytra rather uniformly coriaceous;
cell structure indistinct, apparently with three cells; flight wings
vestigial, consisting of short membranous strips (Fig. 2).

Extremities. — Antenna 1 segments clothed with short, dark hairs
and scattered longer hairs; pr()})()rti()iis (60 units ^ 1 mm) I, 16;
II, 32; III, 26; IV, 29. Tibiae and tarsi with usual scattered dark
spines; all leg structures covered with semi-long light brown hairs.

Genital structures. — Parandria, paramere, sclerotized struc-
tures of aedeagus, filum gonopori. and coupling plate as shown in
Fig. IC, E, F, G, H, and I.

Size. — Length 2.7 mm. width 1.4 mm.

Female. — Similar to male, but slightly larger (see table);
subgenital plate truncate (Fig. ID). Hemelytra with polished area
on underside of embolium about two-thirds toward apex from base,
polished area having several short decumbent hairs directed caudad.

Sept. 1972

POLHEMUS: MEXICAN SALDIDAE

141

Q c<si^^iU!iJji^

Fig. 1. Enalosalda mexicana (Van Duzee) : (A) Female genitalia: Ts,
tergite 8; Tg, tergite 9; PTs, paratergite 8; Ga, gonangulum; 1 Gx, first gonocoxa;
IR, first ramus; 1 Gpo, first gonopophysis. (B) Female genitalia: Gpl, gonoplac;
2 Gx, second gonocoxa; 2R, second ramus; 2 Gpo, second gonopophysis; Cp,
connecting piece. (C) Parandria. (D) Female subgenital plate. (E & F) Left
male paramere, two views. (G) Male coupling plate. (H) Aedeagus. lateral view.
(I.) Aedeagus, dorsal view.

Ovipositor short, broad, with six small teeth (Fig. lA, 1 Gpo). An-
tennal proportions, I, 18; II, 40; III, 30; IV, 30.

Mean length of lOd'd': 2.68mm (max 2.7; min 2.6)
1.36mm (max 1.4; min 1.3)
3.06mm (max 3.1; min 2.9)

Mean width of 10 d" d
Mean length of 10$ 9

142 GREAT BASIN NATURALIST Vol. 32, No. 3

Mean width of 10? 9: 1.59mm (max 1.7; min 1.5)

Material examined. — 104cf cT, 66 $ $ , 15 nymphs, San Carlos
Bay, Guaymas, Sonora, Mexico, CL1202, 28-V-1966 to 5-VI-1966
(JTP); 9crcf, 2$ ?, 3 nymphs, Ensenado Lalo, Sonora, Mexico,
27 56' 15" N X 111 17' 30" W, 29-X-1966, W. G. Evans. On high
mid-tide rocks (JTP); 1 2, Angel de la Guardia Island, Gulf of
Lower California, 29-VI-1921, E. P. Van Duzee (Holotype, CAS);
Id', Pelican Point, Sonora, Mexico, 31 20' N, 113 38' W, on
reef, water's edge, 27-III-1969, V. Roth (JTP).

The Ensenado Lalo and Pelican Point series were kindly fur-
nished by the collectors, and I am indebted for the opportunity to
study this material.

Variation. — The Ensenado Lalo series exhibits a greater vari-
ation in eunomy than the San Carlos series, with the brownish
areas often less extensive but much darker where they occur, al-
most piceous. A very dark male has head markings darker, in the
same pattern as the described male, but has the pronotum only nar-
rowly margined with ochreous, the light area along the collar in-
terrupted medially, and the remainder piceous; the scutellum is con-
colorous; the hemelytra are largely deep brownish black, with
scattered light areas and broad elongate light area on the basal
three-fourths of the outer corium interrupted medially by a weakly
shining deep brown wedge; other markings similar to the described
male, but the brown is darker.

In a female from the same series, the brown markings are deep
as in the male, and the pattern is similar to the described male ex-
cept the hemelytra, which are largely leucine to ochreous, with deep
brown markings as follows: lightly shining spot on embolium and
adjacent irregular contiguous spots on outer and inner corium one-
fifth of the distance from base to apex; slight markings along claval
commissure, and a tiny spot on clavus near apex of scutellum.

The veins delineating the cells of the membrane have a rather
random pattern, creating from three to five irregularly shaped
cells on each hemelytra (Fig. 2). This is true throughout all series.

Fig. 2. Enalosalda mexicana (Van Duzee), left hemelytron.

Sept. 1972 poLHEMus: Mexican saldidae 143

Habitat. — Enalosalda mexicana (Van Duzee) was found at
San Carlos Bay in the intertidal zone on porous volcanic rock. The
rock abounded with tiny pockets and crevices where the bugs hid
when disturbed. None were observed feeding; however, it is pre-
sumed they utilize the abundant sea life clinging to the rocks. In
June of 1966 the saldids made their appearance on the rocks as
soon as the tide receded sufficiently for the surfaces to start drying,
and were most abundant on dry rock. None were found where
substantial spray was evident. The habitat seems restricted to the
mid-tidal zone, where the full tide covers the rocks with several
feet of water and low tide leaves several feet of rock exposed. All
suitable areas along the rocky headland south of the trailer park
near the Yacht Club at San Carlos Bay had abundant populations
of the bugs, but a few miles distant in areas that appeared suitable
none could be found. The rocky headland where the colonies are
found is protected by a nearby island, which could be important
since it protects the area from heavy seas.

When the low tide occurs at night, it will be interesting to de-
termine whether or not the saldids feed, as most saldids are posi-
tively phototropic.

(In March of 1967 the habitat described above and another har-
boring this species on nearby Deer Island were examined. At this
time of year the tide has little variation and is very low, so the
saldids followed a pattern consisting of activity in sunlight and re-
treating to the rock crevices during darkness, the normal pattern
of activity followed by littoral saldids. )

loscytus suttoni (Drake and Hussey), n. comb.

Saldula suttoni: Drake and Hussey, 1951, Occ. Papers Mus. Zool. Univ. Mich.
536:1-3 (Type: Michoacan, Mexico; Museum of Zoology, Univ. Mich.).

I have examined the type Saldula suttoni and find that it must
be transferred to the genus loscytus because of the nature of the an-
tennae, dorsal hairiness, pronotum, and general facies. The left
hemelytron is shown in Fig. 5.

Material.^ — Mexico: \ <S (Holotype), Michoacan, 26 km S
Patzcuaro, under stones, 10,000 ft 2-III-1949. G. M. Sutton (UM).

Saldula sazicola, n. sp.

Of moderate size, quite slender, general color black, macrop-
terous. (For all measurements 60 units =1 mm.)

Coloration. — Hemelytra dull black, barely shining; head,
scutellum, thorax, and venter of abdomen deep brown to piceous,
with golden pubescence on dorsum and fine silver pubescence on
venter; spot on each side of vertex of head extending from hind
margins of ocelli to eye, anteriorly diverging bands on each side
of frons, outer corium anteriorly, spots on clavus and hemelytra
yellowish; lateral pronotal margins whitish yellow, sharply be-

144 GREAT BASIN NATURALIST Vol. 32, No. 3

coming black anteriorly and posteriorly; embolium whitish yellow,
becoming leucine posteriorly, there expanding to a large concolorous
spot adjacent to membrane; clypeus and labrum ochroleucus; an-
terior and middle coxae margined with subhyaline; hind coxae
acetabula, ventral segments of abdomen brown margined; outer
corium yellowish anteriorly; inner corium with yellowish spot
mediolaterally, two yellowish spots between the apex of the clavus
and the membrane along hemelytral commissure. Narrow whitish
yellow band or spot at apex of inner corii m along anterior margin
of membrane. Clavus with basal portion brownish yellow, and a
yellowish spot near the apex. Legs hyaline to leucine, anterior fe-
mora brownish on apical four-fifths, middle and hind femora
brownish on apical one-third, all femora light at apex; knees dark,
middle and hind tibiae slightly embrowned over apical half, be-
coming lighter before apex which is brown.

Head.- — Vertex raised medially in ocellar region; frons with
median sulcus, and a cavity between the posterior portion and
eye; a pair of conspicuous long black hairs on vertex, another pair
on frons; rostrum brownish, extending between hind coxae.

Thorax. — Shining, faintly rugose, clothed with conspicuous
golden pubescence; lateral margins straight, narrowing moderately
anteriorly; callus strongly raised, with circular deep impression on
each side; posterior lobe shorter than anterior lobe, midline ratio
8:13.

Scutellum about as wide as long (W, 51 ; L, 47) .

Wings. — Hemelytra fully developed, covered with scattered
golden pubescence, the longest subequal to width of the hind tibia;
membrane long, with four cells, clouded with ochreous to brown,
veins darker. For the eunomy, see Figs. 3F, G, and H.

Extremities. — Antennae long, slender; segment 1 stoutest, tes-
taceous; segments 2, 3, and 4 deep brown; all segments clothed with
short, inconspicuous hairs; antennal proportions:

cf: I, 19; II, 44; III, 29; IV, 27.
?: I, 20; II, 47; III, 31; IV, 30.

Tibiae with a few scattered dark spines.

Genital structures. — Ovipositor of female with very stout,
well-separated teeth on saw. Mole paramere (Figs. 4C and D)
clothed with fine hairs, filum gonopori coiled one and one-half
times; coupling plate and parandria of male. (Figs. 41 and J.)

Holotype (d)- length 3.5 mm, width 1.4 mm.
Allotype ( $ ) : length 3.7 mm, width 1.6 mm.
Mean length of lOcT cT: 3.28 mm (max 3.5; min 3.1)
Mean width oflOd'cT: 1.31 mm (max 1.4; min 1.3)
Mean length of 10 $ $ : 3.76 mm (max 3.9; min 3.4)
Mean width of 10 9 9: 1.59 mm (max 1.7; min 1.5)

Sept. 1972 poLHEMus: Mexican saldidae

145

Fig. 3. Saldula dewsi group, hemelytra: (A, B. C) Saldula abdominalis
(Champion). (D, E) Saldula ventralis (Stal). (F, G, H) Saldula saxicola n. sp.
(I, J, K, L) Saldula dewsi (Hodgden).

Material examined. — HolotypecT, allotype?, and paratypes
'^d d and 4 ? $ , 5 miles (8 km) NE Castle Hot Springs, Arizona, CL
312, 7-X-1964 (JTP); 12cf d', 16 9 9,2 nymphs, Cuchajaqui Arroyo,

146

GREAT BASIN NATURALIST Vol. 32, No. 3

J

Fig. 4. Saldula dewsi group, genitalia: (A, B) Saldula dewsi (Hodgden),
left m^e paramere, two views. (C, D) Saldula saxicola n. sp., left male paramere,
two views. (E, F) Saldula abdominalis (Champion), left male paramere, two
views. (G, H) Saldula ventralis (Stal), left male paramere. two views. (I, J)
Saldula saxicola n. sp.. male coupling plate (1) and parandria (J).

near Alamos, Sonora, Mexico, CL 1215, 21-III-1967, J. T. Polhemus
(JTP); 6crcf, 3 9 $, Cuchajaqui Arroyo, near Alamos, Sonora,
Mexico, CL 1264, 29-V-1966, .1. T. Polhemus (JTP); 4crcf, 2$ 9,
1 mile (2 km) SW Ixhuatan, Chiapas, Mexico, CL 1098, 5-V-1964,
J. T. and M. S. Polhemus (CU); ScTcT, 2 9 9, 2 miles (3 km)

Sept. 1972 poLHEMus: Mexican saldidae

147

Fig. 5. Hemelytra: loscytus suttoni (Drake and Hussey), type.

NE Ixhuatan, Chiapas, Mexico, CL 1099, 5-V-1964, J. T. and M. S.
Polhemus (CU);6d"cr,2 § $, Santa Fe, Chiapas, Mexico, CL 1101,
5-V-1964, J. T. and M. S. Polhemus (CU); 3cf c^, 3 9 9, 3 miles
(5 km) W El Naranjo, 1200 ft (367 m), San Luis Potosi, Mexico, 5-
VM965, Burke, Meyer, and Schaffner (TAM) ; 1 cT, Rio Quezalapan,
2 miles (3 km), E Lago Catemaco, Veracruz, Mexico, 21 -VI to S-VIL
1964, J. R. Meyer (TAM); IcT, Ciudad Victoria, Mexico, 14-Vn-
1950, Drake and Hottes ( JTP) ; 1 9 , Cola de Caballo, 3000 ft (917
m), Nuevo Leon, Mexico, 6-Vn-1959, R. B. Selander and J. C. Schaff-
ner (JTP); Icf, Real de Arriba, Temescaltepec, Mexico, 4-VM 933;
H. E. Hinton and R. L. Usinger (JTP); 1 d", 22 miles W Ptazcuaro,
Michoacan, Mexico, CL 1033, 23-IV-1964, J. T. and M. S. Polhemus
(CU). Holotype, allotype, and paratypes in the University of
Colorado Museum collection; paratypes will be deposited in the
collections of Texas A & M University, California Academy of
Sciences, and the author.

Comparative notes. — The most obvious external character
separating Saldula saxicola from 5. dewsi (Hodgden) is the long,
hairy dorsal vestiture of dewsi. The male parameres are helpful in
diagnosing the group, those of dewsi usually being larger, having a
tuft of heavier black hairs (Figs. 4A and B), and a shape different
from either saxicola or Saldula abdominalis (Champion).

Saldula saxicola was first recognized in the Arizona material and
later additional specimens were found in the Mexican material
being held under abdominalis. While the second antennal segment
in some of the southern Mexico forms occasionally overlaps the
proportions found in abdominalis, the eunomy will help to separate
the two. The hemelytral markings in abdominalis (Figs. 3A, B,
and C) tend to form an eye-shaped spot arising from the embolium
at the forepart of the wing, whereas the tendency in saxicola is to
form a solid fascia arising from the embolium, and in darker speci-

148 GREAT BASIN NATURALIST Vol. 32, No. 3

mens the light area draws away from the embohum and degenerates
into a line parallel with it. Rarely is saxicola found without a
line or thin spot of white in the inner corium along the anterior
margin of the membrane, whereas this is lacking in abdominalis;
and two white spots are often found close to the hemelytral com-
missure in saxicola^ whereas never more than one is present in ab-
dominalis. Also the fascia at the apex of the corium along the em-
bolium is usually solid white in saxicola, and if darkening occurs it
is in the center. In abdominalis this spot is usually U-shaped, open
at the rear; as the eunomic series progresses, the lighter colored
specimens have the U filled in with white, and the eye-shaped spot
well developed.

In the northern populations (Arizona, Sonora, Tamaulipas) of
S. saxicola, the ratio of the length of the second antennal segment
to the width of the head through the eyes will easily separate them
from abdominalis. as the ratio is .80 or more in saxicola and a
maximum of .60 in abdominalis. In the southern specimens, how-
ever, the ratio varies widely even within a given series, and an oc-
casional overlap is found. Of the 51 specimens of saxicola examined
for this character, only 4 had ratios of .60 or less, the lowest being
.57. Of the 21 specimens of abdominalis studied, only 4 had ratios
of .57 or greater, the highest being .60. Of the 8 specimens in the
overlap zone, none presented a great problem in separation because
of the long series available to work out the eunomy of each species.

Habitat. — In Arizona, S. saxicola was found on exposed sloping
sandstone in a streambed. The water was seeping from the base of
an abandoned dam, wetting the roughly sculptured rock surfaces
and forming pools where Trepobates becki Drake and Harris was
abundant. On the more steeply sloping damp surfaces, S. saxicola
was collected, along with Saldula pexa Drake.

At Cuchajaqui Arroyo, Mexico, the specimens were found in a
similar habitat, on vertical stone surfaces beside a stream flowing
into a large vertical-walled, sandstone basin and on a large log in the
stream. Other locations in Mexico produced saxicola from simi-
lar situations, the specimens almost always being found on vertical
or steeply sloping damp stone surfaces.

Distribution. — In the Saldula dewsi species group, S. ab-
dominalis, S. dewsi, S. saxicola, and Saldula ventralis (Champion)
are found in Mexico. S. ventralis is the southernmost component
of these and is not easily confused with the others. S. dewsi is
found in Arizona, Mexico, and Central and South America, roughly
paralleled in distribution by 5. abdominalis; but the latter has not
been found north of Central Mexico. S. saxicola apparently has its
distributional center in Mexico and ranges into Arizona.

Saldula duran^oana, n. sp.

Moderately large, robust; general color black and light brown,
macropterous. (All measurements: 60 units = 1 mm.)

Sept. 1972 POLHEMUS: MEXICAN SALDIDAE 149

Coloration. — Hemelytra dull black, not shining; head, scutel-
lum, thorax, and venter of abdomen black, covered with fine de-
cumbent golden pubescence on dorsum and thick silver pubescence
on venter; clypeus, labrum, rostrum testaceous to brownish; an-
terior and middle acetabulae, coxae, ventral abdominal segments
posteriorly margined with subhyaline; base of embolium, corium
next to clavus, two medial elongate spots between corium and em-
bolium, and small spot at apex of embolium deep brown to black;
remainder of corium and embolium luteous, inner corium lighter at
median corial suture, outer corium subhyaline. Clavus black, with
fine golden pubescence, apex with faint brownish area. Legs testa-
ceous to ochreous, anterior and middle femora blackish beneath on
apical three-fourths, posterior femora blackish beneath on apical
half, femora ochreous basally, with brownish spots on sides.

Head. — Shining with scattered long hairs on vertex and frons.
Vertex with usual small yellow spot between ocelli and eye on each
side, faintly rugose; ocelli large, seperated by approximately the
width of an ocellus. Rostrum extending between hind coxae.

Thorax. — Pronotum moderately shining, faintly rugose, clothed
with long, erect, black hairs that are not obvious unless viewed
from the side, except a few along the anterior lateral margins; lateral
margins curved, slightly explanate, narrowing moderately anteriorly.
Callus moderately raised with a deep circular median impression;
posterior lobe shorter than anterior lobe (not including collar), mid-
line ratio 12:14, length to width ratio 30:103. Anterior and
middle acetabula, coxae margined with subhyaline. Scutellum as
wide as long (70:70).

Wings. — Hemelytra fully developed, dull, not shining, with
scattered golden pubescence, a little thicker at the base of the em-
bolium, entire dorsum except membrane clothed with long, erect,
black hairs. Membrane faintly brownish, subhyaline, veins slightly
darker; long, with four cells (Fig. 6).

Extremities. — Antennae moderately long, moderately stout;
segment 1 stoutest, testaceous, dark brown beneath on apical three-
fourths; segment 2 longest, dark brownish yellow, lighter on apical
third, apex dark; segments 3 and 4 black, somewhat thicker than
segment 2; thicker than hind tibia; all segments clothed with semi-
short pubescence, longer on segments 1 and 2, scattered longer stiff
hairs on segments 3 and 4; antennal proportions:

d: I, 18; II, 34; III, 24; IV, 30
9: I, 18; II, 43; III, 26; IV, 30

Tibia covered with short, depressed hairs, none obviously longer
than diameter of tibia, and usual dark spines; femora with longer,
thin hyaline hairs beneath.

Genital structures. — Male paramere (Figs. 7A and B) stout,
clothed with long, fine hairs. Parandria as shown in Fig. 7C.

150

GREAT BASIN NATURALIST

Vol. 32, No. 3

Fig. 6. Saldula durangoana n. sp.: (A, B) hemelytra.

Holotype ( ? ) : length 4.5 mm; width 2.1 mm.
Allotype ( cf ) : length 3.8 mm; width 1.7 mm.
Paratype ( 9 ) : length 4.2 mm; width 2.0 mm.

Allotype. — The allotype (male) darker than holotype; outer
corium mostly brown to dark brown with a white spot medially at
the apical four-fifths.

Material examined. — Holotype ( 9 ) and paratype ( 9 ), 39
miles W Durango, Durango, Mexico, CL 1013, 20-IV-1964, J. T.

Fig. 7. Saldula durangoana n. sp.: (A. B) Left male paraniere, two views.
(C) Parandria.

Sept. 1972 poLHEMus: Mexican saldidae 151

and M. S. Polhemus. Allotype (cT), 61 miles W (99 km) W Du-
rango, Durango, Mexico, CL 1014, 20-IV-1964, J. T. and M. S.
Polhemus. (The holotype and allotype are in the University of
Colorado Museum, and the para type in the Polhemus collection.)

Comparative notes. — The species most closely resembles 5a/-
dula luciuosa (Stal) but luctuosa has a faintly shining to shining
hemelytra, while durangoana is dull. This species bears a slight re-
semblance to Saldula andrei Drake; however, andrei is a narrower
species with polished shining scutellum and pronotum, straight
lateral pronotal margins, and a tendency for the apical spot on the
clavus to form a fascia.

Habitat. — Taken from amongst the grasses at two locations,
one a grazing meadow, the other a margin of a farm pond. At both
locations the ground was very wet. The elevation here is approxi-
mately 2000 m (6500 ft).

Saldula sulcicollis hispida (Hodgden), n. comb.

Salda hispida Hodgden, 1949, J. Kansas Ent. Soc. 22(4): 115-156 (Type: Temescal-
tepec, Mexico; Calif. Acad. Sci.)

The comparison of the type of Saldula hispida (Hodgden) with
a long series of Saldula sulcicollis (Champion) from various loca-
tions ranging from Arizona to Costa Rica has revealed that hispida
cannot be considered more than a subspecies of sulcicollis. Speci-
mens from the high mountains of Northern Chiapas in Mexico agree
well with the type of hispida, and a male has been designated as
typical of the subspecies and is described below.

In the series of sulcicollis from San Cristobal las Casas, Chiapas,
specimens vary from the typical form almost to the hispida form
in antennal dilation and coloration, which are the only reliable
characteristics which will separate the two. In the typical form of
sulcicollis^ the last two antennal segments are slightly swollen and
the apical half of the fourth segment is light colored, whereas in the
hispida form the antennal segments are more slender and the last
segment concolorous. The male genitalia of the two forms agree
perfectly, and the other characters used by Hodgden for separa-
tion are too variable to be reliable.

As the type of hispida is from Real de Arriba (about 7000 ft)
and the Chiapas material of hispida is from about the same altitude,
it seems that this subspecies is restricted to the higher elevations,
while the typical subspecies inhabits the valleys and plains from
about 3000 to 5000 ft (900 to 1500 m) elevation. Very recent col-
lections in Guatemala also support this view.

Description of male. — Of moderate size, general color deep
brown to black, macropterous, entire dorsum (except membrane)
covered with long, erect setae, and semilong sparse golden pubes-
cence. (For all measurements, 60 units = 1 mm.) Length 4.5
mm, width 1.8 mm.

152

GREAT BASIN NATURALIST

Vol. 32, No. 3

Coloration. — Hemelytra black to deep chocolate brown with
pruinose and leucine markings; head, scutellum, thorax, and venter
of abdomen black; small spot between ocelli and each eye, clypeus,
anteclypeus, labrum ochreous; rostum flavous to ochreous; clavus
velvety black, with bluish pruinose areas at base, middle and near
apex, narrow area at suture shining; inner corium pruinose black
basally, pruinose area terminating in whitish grey at basal one-
fourth; small whitish spot medially, whitish pruinose area next to
membrane at middle; remainder shining black; outer corium blackish
brown basally, becoming translucent deep chocolate brown apically,
with leucine spot at apical fourth on embolium; legs ochreous to leu-
cine; femora medially darkened, lighter at base and apex, with usual
brown spots; tibia dark at knees, apex and broad medial area darkest
on anterior legs, leucine elsewhere; tarsi with first segment largely
leucine, second segment darker; coxae leucine, dark basally.

Head. — Shining, frons and vertex rugulose. Rostrum reaching
between hind coxae; frons slightly depressed longitudinally at mid-
dle; ocelli approximate, slightly elevated. Eyes large, extending
laterally half their width beyond anterior pronotal margins, with
scattered setae.

Thorax. — Pronotum rugulose, shining callus strongly raised,
with deep medial depression and a tiny, shallow depression on each
side of the medial pit, marked off from collar and posterior lobe by
deep, pitted sulci; collar wide (5) ; anterior lobe longer than posterior
lobe (17/11); lateral margins slightly concave; humeral angles

A

C^^

c.

Fig. 8. Saldula sulcicollis (Champion), male genitalia: (A) Parandria.
(B, C) Aedeagus, lateral view (B) and dorsal view (C). (D, E) Left male para-
mere, two views.

Sept. 1972 poLHEMus: Mexican saldidae 153

sharply rounded; posterior margin broadly convex; posterior width/
anterior width, 88/45. Scutelluni rugulose, shining; width/length,
58/62.

Wings. — Hemelytra fully developed; membrane fumose, with
whitish mottling, veins dark, outer margin shining brown basally
and apically, leucine medially, with four cells.

Extremities. — Antennal segment 1 ochreous, with small brown-
ish anterior spot; segment 2 ochreous. brown at apex, slender; seg-
ments 3 and 4 dark brown, subcqual in diameter to segment 1; seg-
ments 1 and 2 clothed with semilong hairs and scattered long hairs,
segments 3 and 4 clothed with short pubescence and scattered longer
hairs; proportions, I, 23; II, 54; III, 36; IV, 41. All leg structures
covered with short, fine, light-colored pubescence; femora with long
hairs beneath.

Genital structures. — Parandria, paramere, and median
sclerotized structure of aedeagus as shown in Fig. 8; filum gonopori
coiled two and one-half times.

Material examined. — 1 cS <S ■, 2 9 $, Mexico, Chiapas, 5 miles
(8 km) S Tapiula, CL 1095, 4-V-1964, J. T. and M. S. Polhemus
(CU); Id", 1$, Chiapas, 12 miles (19 km) N Bochil, CL 1091,
J. T. and M. S. Polhemus (CU).

Literature Cited

CoBBEN, R. H. 1959. Notes on the classification of Saldidae with the descrip-
tion of a new species from Spain. Zool. Meded. Leiden 36:303-316.

Drake, C. J. 1962. Sjoionymic data and two new genera of shorebugs
(Hemiptera: Saldidae). Proc. Biol. Soc. Washington 75:115-124.

Drake, C. J., and L. Hoberlandt. 1950. Catalog of genera and species of
Saldidae (Hemiptera). Acta. Ent. Mus. Nat. Pragae 26(376) : 1-12.

Lattin, J. D., AND R. H. CoBBEN. 1968. The systematic position of Orthophrys
meiicanum Van Duzee (Hemiptera Heteroptera: Saldidae). Ent. Berichten
28:130-135.

HoDGDEN, B. B. 1949. New Saldidae from the western hemisphere (Hemip-
tera). J. Kansas Ent. Soc. 22(4) : 149-165.

Pendergrast, J. G. 1957. Studies on the reproductive organs of the Heterop-
tera with a consideration of their bearing on classification. Trans. Roy. Ent.
Soc. London 109:1-63.

Polhemus, J. T., and W. E. Evans. 1969. A new genus of intertidal Saldidae
from the eastern tropical Pacific with notes on its biology (Hemiptera).
Pacific Insects 11 (3-4) :571-578.

Scudder, G. G. E. 1959. The female genitalia of the Heteroptera: Morphology
and bearing on classification. Trans. Roy. Ent. Soc. London 111:405-467.

Van Duzee, E. P. 1923. Expedition of the California Academy of Sciences to
the Gulf of California in 1921. The Hemiptera (true bugs, etc.). Proc.
California Acad. Science, Fourth Series. 12(11): 123-200.

SCORPIONS OF UTAH
John D. Johnson' and Dorald M. Allred-

Abstract. — The 736 scorpions representing nine species collected in Utah,
listed in order of greatest to least abundance, are Vaejovis boreus, V. utahensis,
Anuroctonus phaeodoctylus, V. confusus. Hadrurus spadix, V. becki, V. wupatkien-
sis, H. arizonensis and Centruroides sculpturatus. Centruroides sculpturalus, H.
arizonensis, V. becki, V. confusus and V. wupatkiensis are reported from Utah
for the first time. Vaejovis boreus is the most widely distributed of the Utah
scorpions. Vaejovis boreus and V. confusus occur in both the Great Basin
and the Colorado River Basin. Centruroides sculpturatus, Hadrurus arizonensis.
H. spadix, V. wupatkiensis, and V. utahensis occur only in the Colorado River
Basin, whereas Anuroctonus phaeodactylus and V. becki are confined to the Great
Basin. Anuroctonus phaeodactylus, V. boreus and V. confusus occur from the
southern to the northern border of the state.

Both males and females were taken from May through October, with
greatest numbers in May, June, and July. Females were taken in greater
numbers than males. This may be related to their mating habits during the
collecting periods, and does not necessarily indicate other than a 1:1 ratio in
normal populations.

Morphological variations associated with distribution were distinguishable in
two of the species collected — Anuroctonus phaeodactylus and Vaejovis boreus.
The mean and standard error computed for the carapace, preabdomen. postabdo-
men, and telson on these two species indicate a significant morphological varia-
tion in the size of the preabdomen between the northern and southern
populations.

Scorpions first collected from Utah were taken by Stansbury
during the late 1840's, but none were named until 1854 (Ewing,
1928). Nothing more was published on scorpions of this region
until Banks (1900) listed several species. However, most of Bank's
records were of specimens from the Marx collection (now in the
U.S. National Museum), which were incorrectl}' labelled and mis-
identified (Gertsch and Soleglad, 1966). Although more interest
was shown in scorpions since the turn of the century, little published
information and pertinent collecting data for Utah are existent.
Studies by Stahnke (1950) and Williams (1966) dealt with scor-
pions of the western United States, but only those by Gertsch and
Soleglad (1966) and Gertsch and Allred (1965) refer to Utah taxa.

Although scorpions are occasionally seen in desert areas of the
western United States, few systematic studies have been made. Most
specimens collected heretofore in Utah have been used for exhibit
and teaching in high schools and universities, and their collection
data is not existent or available. Specimens from Brigham Yoiuig
University, supj)lemented by collections made by the senior author,
constitute the basis for this listing of species from Utah, their geo-
graphic (Hstribution, seasonal occurrence, and sex ratio, and notes
on some morphological variations associated with distribution. Speci-
mens from other institutions, if existent, were not obtainable.

M)op<irtniorit of niology. Sail Jose City College, San Jose, (California ',)5I25.
-Dcpiirliiicnt of '/oology, Brigham Young University, Provo, Utah 84(i01.

154

Sept. 1972 JOHNSON, ALLRED: SCORPIONS 155

Dr. Stanley C. Williams, San Francisco State College, verified
and/or identified some specimens. Financial assistance was pro-
vided, in part, by a Karl G. Maeser Research Award given to Drs.
D Elden Beck and Dorald M. Allred by the Alumni Association of
Brigham Young University, shortly before Dr. Beck's untimely
death in August, 1967. The Department of Zoology, Brigham Young
University, suppUed laboratory space, equipment, supplies, and some
transportation.

Procedures

Scoq)ions were collected from the eastern part of the Great Basin
and western area of the Colorado River Basin within the political
boundaries of Utah. The ecology and topography of these basins
were described by Durrant (1952), Hay ward, Beck, and Tanner
(1958), Allred, Beck, and Jorgensen (19'63), and Allred (1968) as
high, inland plateaus separated by mountain ranges. These moun-
tains begin at the northwest corner of Wyoming and extend south-
ward approximately two-thirds the length of Utah, then westward
to the southwest corner of the state. They are among the highest in
Utah with elevations from 9,000 to 12,000 ft. Within each of the two
basins are lower mountain ranges oriented in north-south directions.

Scorpions already existent in collections had been taken by their
collectors principally during daylight hours under rocks and other
ground cover. Most of the ones being reported herein were captured
at night by use of the ultraviolet light technique described by
Honetschlager (1965).

Measurements for comparison and identification were taken in
the following manner. The total length is from the anterior tip of
the carapace to the posterior tip of the cauda, excluding the telson.
The length of the cauda is from the anterior margin of the first
caudal segment to the posterior margin of the fifth segment, ex-
cluding the telson. The length of the pedipalp is from the proximal
end of the femur to the distal end of the fixed finger, excluding the
coxa and trochanter. The length of the chela includes the palm and
fixed finger as a single unit.

Adult scorpions were identified by use of keys by Ewing (1928),
Comstock (1948), Gertsch and Allred (1965), Gertsch and Sole-
glad (1966), and Williams and Hadley (1967). Specific keys are
not available for the identification of immature specimens, but
morphological characteristics of most young are so similar to those
of the adults that they can be assigned to species.

The mean and standard error (x, s) were computed to determine
statistical significance in variation (95 percent level) between dif-
ferent populations.

Collectors of the scorpions reported in this study are indicated
by their initials as follows:

BB

B. Bradley

CC

C. Cottam

BM

B. McMorris

CH

C. L. Hayward, et al.

BL

B. R. Lemora

DA

D. M. Allred

BW

B. Weston

DB

D E. Beck

156

GREAT BASIN NATURALIST Vol. 32, No. 3

DU

D. Usher

LH

L. Hook

EC

E. Cottam

MD

M. Dumas

FC

F. Camenzind

MK

M. Killpack

FH

F. Harmston

PH

P. Hastings

GH

G. M. Harmston

SD

S. C. Daines

GK

G. F. Knowlton

SO

S. Oliver

HV

H. E. Yokes

VT

V. M. Tanner

JF

J. Finlev

WG

W. J. Gertsch

JJ

J. D. Johnson

WR

W. Robertson

JR

J. A. Rowe and C.

F.

Smith

Key to Scorpions of Utah and Southern Nevada

1. Middle lamellae of pectine consists of five or fewer irregu-

ularly-shaped, rectangular segments (Fig. 1) 2

Middle lamellae of pectine consists of six or more regularly-
shaped, oval segments (Fig. 2) 3

2. Ridges on fingers of pedipalps black; telson abruptly con-

stricted near base of aculeus (Fig. 3); medioventral
margin of moveable finger of chelicera with conspicuous
brush of hair on basal part Anuroctonus phaeodactylus

Ridges on fingers of pedipalps same color as rest of finger;
telson not abruptly constricted (Fig. 4); medioventral
margin of moveable finger of chelicera with light fringe
of hair Superstitionin doncnsis

3. Medioventral margin of moveable finger of chelicera with

one or two large, conspicuous, dark teeth (Fig. 5) 4

Medioventral margin of moveable finger of chelicera lacks

large teeth, but often finely serrate or irregular (Fig. 6) .. 6

4. Medioventral margin of moveable finger of chelicera with

two large teeth; telson with subaculear spine (Fig. 7)
Ccntruroidcs sculpturatus

Medioventral margin of moveable finger of chelicera with
one large tooth; telson lacks subaculear spine (Fig.
4) - 5

5. Interocular area of carapace pale, much lighter than dark

color of rest of carapace Iladrurus arizonensis

Interocular area of carapace dark brown or black, same

color as rest of carapace Hadrurus spadix

6. Palm of pedipalp has prominent, granulose keels; inner

margin of fixed finger of pedipalp stronglv crenulate

(Fig. 8) : 7

Palm of pedi|)alp essentially smooth, keels not prominent;
inner margin of fixed finger of pedipalp straight or only
slightly crenulate (Fig. 9) 8

7. Palm of p('(h[)aip approximately twice as long as wide;

Sept. 1972 JOHNSON, ALLRED: SCORPIONS 157

telson smooth; fixed finger of pedipalp about same
length as palm (Fig. 10) Vaejovis becki

Palm of pedipalp less than one and one-half times as long
as wide; telson lightly granulate, especially near base;
fixed finger of pedipalp shorter than length of palm
Vaejovis horeus

8. Venter of telson densely covered with long hairs (Fig. 11)

Vaejovis hirsuticauda

Telson with only a few long hairs (Fig. 4) 9

9. Palm of pedipalp not over one and one-half times as long

as wide Vaejovis utahensis

Palm of pedipalp twice as long as wide 10

10. Venter of telson conspicuously granulate over most of area

(Fig. 12) Vaejovis confusus

Venter of telson granulate essentially only at base, and then

only lightly so (Fig. 13) Vaejovis wupatkiensis

Anuroctonus phaeodactylus (Wood)

Centrums phaeodactylus Wood, 1863, Proc. Acad. Nat. Sci. Philadelphia, p. Ill;
1863, J. Acad. Nat. Sci. Philadelphia (2)5:372.

Anuroctonus phaeodactylus Pocock, 1902, Biol. Centrali-Americana, Arachnida,
Scorpions, Pedipalpi and Solfugae, p. 14.

In Utah, scorpions of this species occur over the entire length,
but not the width, of the state. Forty-seven specimens were collected
from the Great Basin, but none were found in the Colorado River
Basin (Fig. 14, I'able 1). Six times as many males as females were
collected, mainly because females were seen only at the entrance of
their burrows, and their quick movements made capture difficult.
This species is also known from California, Colorado, Nevada,
Virginia, Baja California, and Guatemala (Pocock, 1902; Gertsch
and Allred, 1965).

This species in Utah is surpassed in length only by Hadrurus
arizonensis and H. spadi.r. It may be distinguished from other
species in Utah by its relatively stout appearance, heavy pedipalps
bearing short, black fingers, and by a bulbous swelling posterior
to the vesicle on the telson in males. On some small males the
sting is extended as an evenly-curved spine, and resembles that
of the female.

These scorpions typically are burrowing animals that exhibit
a high degree of aggregation; they are not randomly distributed
as individuals, but occur in scattered colonies in relatively large
numbers (Williams, 1966).

The mean and standard error were computed for morphological
variations of the carapace, preabdomen, postabdomen and telson for
specimens of two groups — those from northern Utah and those from

158

GREAT BASIN NATURALIST

Vol. 32, No. 3

Pigs 1-13 (1) Poctines with rectangular, middle lamellae; (2) pectines with
oval middle lamellae; (3) telson of Anuroctonus phaeodactylus; (4) telson
which lacks aculear constriction; (5) ventral view of chelicera with large tooth
on moveable finger; 6) ventral view of chelicera with serration in place of
large tooth on moveable finger; (7) telson with ventral spine at base of aculeus;
(8) pedipalj, with crenulated fixed finger; (9) pedipalp with non-crenulated,
tixed inger; (10) pedipa p with designated limitations of palm and fixed finger;
11) teson oiVaeiovis hirsuticauda; (12) telson witli conspicuous granulation;
{\5) telson which essentially lacks conspicuous granulation

Sept. 1972

JOHNSON, ALLRED: SCORPIONS

159

0 A. phaeodactylus
J^ C. sculpturatus

Fig. 14. Distribution of Anuroctonus
atus and Hadrurus arizonensis in Utah.

:tylus, Centruroides sculptur-

southern Utah (Figs. 15-18, Table 2). No significant differences
occurred in the lengths of the carapace, postabdomen or telson at
the 95 percent level of confidence, but a significant difference in
the length of the preabdomen (1.24 mm) was evident between the
two populations.

160

GREAT BASIN NATURALIST

Vol. 32, No. 3

Table 1. Collection data for Anuroctonus phaeodactylus in Utah.
County and location Date

No. and sex Collector

Box Elder, Lucin

7-VII-68

16 m, 1 f

Iron, Cedar City

4-VI-68

5 m

Millard, Holden

l-VI-68

1 m

Clear Lake

3-VI-68

14 m, 3f

Tooele, Knolls

5-VII-68

1 m

Utah, Provo

VII-58

1 m

DB

VIII-58

1 m

DB

X-67

1 m

GH

Spanish Fork

V-28

1 f

DB

Springville

unknown

1 m

JF

Washington, 10 mi S.

6-VI-68

1 m

JJ

Shivwits Indian Res.

on U.S. Hwy 91

Table 2. Length of the carapace, preabdomen, postabdomen, and
telson of 20 Anuroctonus phaeodactylus from northern Utah and
20 from southern Utah, with computation of the mean and standard
error.

Northerr

I Utah

Southern Utah

Car.

Preab.

Postab.

Tel.

Car.

Preab.

Postab.

Tel.

8.0

16.7

"193^

7.8

8.2

18.8

20.0

8.0

8.0

15.7

19.9

7.4

8.9

13.7

20.7

7.8

8.2

16.2

20.0

7.5

8.4

19.9

20.6

7.8

7.9

17.7

18.6

7.0

8.1

20.7

20.9

8.0

7.9

13.8

19.5

7.5

7.8

13.9

19.2

7.4

8.1

15.9

18.8

7.0

7.8

21.2

19.7

6.9

7.8

15.4

18.8

7.5

7.8

18.6

18.3

7.0

7.7

15.4

19.1

7.3

8.3

20.9

20.0

7.8

8.2

17.6

19.2

7.6

7.9

19.7

19.0

7.4

7.8

13.9

18.6

7.3

8.0

21.1

19.4

7.2

8.2

14.2

19.2

7.1

7.8

20.8

18.7

7.0

8.1

16.4

19.3

7.5

8.2

20.6

19.2

7.3

7.9

18.0

20.0

7.6

8.3

19.8

19.2

6.8

7.8

15.1

19.0

7.1

8.4

16.0

20.8

7.6

7.5

14.9

18.7

6.8

7.7

18.7

19.7

7.0

8.9

17.0

21.4

8.3

7.8

17.8

18.4

6.9

8.5

13.0

21.7

7.7

8.0

18.5

19.0

7.3

8.8

13.0

21.7

7.8

8.0

17.0

19.1

7.2

8.5

12.9

22.9

8.7

7.9

14.0

19.8

7.6

9.0

14.3

21.0

7.9

Mean

8.14

15.38

19.81

7.52

8.10

18.51

19.53

7.37

SE

±.19

±.72

±.55

±.21

±.13

±1.17

±.68

±.23

Sept. 1972

JOHNSON, ALLRED: SCORPIONS

161

21

20

19

1

18

17

1

16

[

15

f

14

13

S

N

8.0 •
7.5 .
7.0

l!

16

h

15

17

Figs. 15-18. The mean, standard deviation and range of length (in mm)
of the (1) preabdomen; (2) carapace; (3) telson; (4) postabdomen of Anuroc-
tonus phaeodactylus from northern and southern Utah.

Centruroides sculpturatus Ewing
Centruroides sculpturatus Ewing, 1928, Proc. U.S. Nat. Mus. 73(2730) :20-21

Members of the species Centruroides sculpturatus occur abun-
dantly throughout southern Arizona (Ewing, 1928), and recent
records have noted their occurrence in Puerto Penasco, Sonora,
Mexico (Williams and Hadley, 1967). In Utah, two females were
collected by D E. Beck from Hole-in-the-Rock, Kane County, May,
1953 (Fig. 14). This is the only known record of this species
in Utah.

These are medium-sized scorpions, comparable in length to
those of Vaejovis boreus. The base color is yellow to orange-brown,
without markings on the body. In general appearance they resem-
ble V . wupatkiensis, but are much longer and have a wider pre-
abdomen. The fingers of the chelae are long, but not as narrow
and slender as in V. wupatkiensis . Members of this species may be
distinguished from others in Utah by their relatively long post-
abdomen and distinct tooth at the base of the vesicle on the telson.

Hadrurus arizonensis Ewing

Hadrurus hirsutus arizonensis Ewing, 1928, Proc. U.S. Nat. Mus. 73:8.
Hadrurus arizonensis Stahnke, 1945, Amer. Mus. Novitates, no. 1298, p. 6.

Scorpions of this species are known from Arizona, southern
Nevada, southern California, and southward into Sonora, Mexico
(Gertsch and Allred, 1965). This is the first time they have been

162 GRKAT I5ASIN NATURALIST Vol. 32, No. 3

reported from Utah. Three specimens were collected from the
Colorado River Basin area of Utah; none were taken from the
Great Basin (Fig. 14, Table 3).

The genus Hadrurus contains the largest Utah scorpions. The
two species which occur in Utah, H. arizonensis and H. spadix, are
similar in length (up to 100 mm in the adult), and both are covered
with red bristles. Specimens of H. arizonensis may be distinguished
from those of H. spadiz by their overall lighter coloring and the
presence of a yellow interocular space extending from the median
eyes to the anterior edge of the carapace.

Hadrurus spadix Stahnke

Hadrurus spadix Stahnke, 1940, Iowa State College J. Sci. 15:102; 1945. Amer.
Mus. Novitates, no. 1298, p. 4.

These animals are known from Nevada, California, and eastern
Oregon (Gertsch and AUred, 1965). Eighteen adults and immatures
were collected in the Colorado River Basin of Utah. Males and
females were taken in a ratio of 1:1 (Fig. 19, Table 4).

Scorpions of this species are structurally similar to those of H.
arizonensis, and color is the only distinguishing characteristic. Mem-
bers of H. spadix are dark brown, almost black, and lack the yellow
interocular space described for those referrable to H. arizonensis.

Table 3. Collection data for Hadrurus arizonensis in Utah.

County and location

Date

No. and sex

Collector

Garfield, Calf Creek

X-46

1 f

BL

Kane, Kanab

6-V-66

1 m

SD

Washington, St. George

1920

1 m

VT

Table 4. Collection data for Hadrurus

spadix in Utah.

County and location

Date

No. and sex

Collector

Garfield, North Wash

V-55

2 m

MD

Grand, Arches Nat. Mon.

VIII-50

1 f

DA

Moab

20-VI-68

2m

JJ

Kane, Hole-in-the-Rock

16-V-53

1 m

DB

San Juan, Navajo Mt.
Trading Post

2-V-55

1 m, 4 f

DB, CH

Glen Canyon

17-V-68

2 f

BM

Canyonlands Nat. Park
(Needles campground)
Washington, Zion
Nat. Park

21-VI-68
7-VII-28

1 m, 2 f
1 f

JJ
VT

Sept. 1972

JOHNSON. ALLRED: SCORPIONS

163

Fig. 19. Distribution of Hadrurus spadix, Vaejovis becki and V. boreus in
Utah.

Vaejovis becki Gertsch and AUred

Vaejovis Koch, 1836; Williams, 1971, Pan-Pacif. Entomol. 47(1): 78-79.
Vaejovis becki Gertsch and AUred, 1965, Brigham Young Univ. Sci. Bull., Biol.

Ser. 5(4): 9-11.

Members of this species are known from southern Nevada and
adjacent California (Gertsch and Allred, 1965). In Utah, six males

164 GREAT BASIN NATURALIST Vol. 32. No. 3

and seven females were taken at St. George, Washington County,
5-VI-1968 (Fig. 19). This is the first time V. becki has been re-
ported in Utah.

Vaejovis becki closely resembles V. boreus, but lacks the charac-
teristic markings and robust, heavily granulated hands of V. boreus.
The base color is yellow to orange-brown. They may be distin-
guished from other species in Utah by the presence of a dusky
patch enclosing the median eyes, which radiates forward almost to
the anterior edge of the carapace. No other markings are present
on the body.

Vaejovis boreus (Girard)

Scorpio (Telegonus) boreus Girard, 1854, in Marcy, Exploration of the Red

River of Louisiana in the year 1852, p. 257.
Vaejovis boreus Ewing, 1928, Proc. U.S. Nat. Mus. 73(9): 12.

Vaejovis boreus is known from California, Arizona, Colorado,
Wyoming, Idaho, Oregon, Washington, Montana, and North Dakota
in the United States, and British Columbia, and Alberta, in Canada
(Gertsch and Soleglad, 1966). It is the most common and most
widely distributed of all scorpions in Utah. It occurs in both the
Great Basin and the Colorado River Basin. In Utah it occurs from
the southern to the northern border of the state. Four hundred and
one specimens were taken in Utah. Males and females were col-
lected in a ratio of 1:1.9 (Fig. 19, Table 5).

This species is medium-sized, comparable in length to V. confusus.
It is the fourth largest species in Utah, surpassed in length only by
Hadrurus arizonensis, H. spadix, and Anuroctonus phaeodactylus. It
may be distinguished from other species in Utah by the character-
istic markings on the carapace and preabdomen, and the robust.

Table 5. Collection data for Vaejovis boreus in Utah.

County and location

Date

No. and sex

Collector

Beaver, Milford

20-IV-30

1 f

VT

Box Elder, Brigham City
Grouse Creek foothills
Lucin

Locomotive Springs
Utah Hwy 70, on
Grouse Creek Road

VIII-28

VI-28

19-VI-52

19-VL52

7-VI-68

2 f
1 f
2f
1 m

2 m, 4 f

WR
VT
DB
DB

JJ

Cache, Blacksmith
Fork Canyon
Logan

9-Vn-35
26-VIII-36

1 f
1 f

JR, CS
FH

Daggett, Sheep Creek
Flaming Gorge

VI-26

28-VI-26

1 f
1 f

EC
EC

Duchesne, Roosevelt

5-Vn-68

11 m, 12 f

JJ

Emery, Ferron
Buckhorn Canyon

IX-28
6-IX-53

2 f
4 f

WR
VT

Sept. 1972 JOHNSON, ALLRED: SCORPIONS 165

Table 5. (continued)

Green River

7TV-46

2 f

GK

7-V-46

1 f

GK

6IX-46

1 m, 5 f

HV

18-VL68

11 m, 13 f

JJ

Garfield, North Wash

V-55

3 f

MD

Grand, Arches Nat. Mon.

12-V-48

1 f

DB

3-V-68

10 m, 70 f

JJ

20-VI-68

2 m, 20 f

JJ

Moab

io-vn-67

1 f

DU

Cisco

18-VL68

6 m, 7 f

JJ

Castle Valley

19-VL68

11 m, 14 f

JJ

Sego

28TV-68

1 f

BW

Iron, Parowan

31-VIII-29

1 m

VT

Newcastle

5-VL68

4 m, 5 f

JJ

Lund

5-VL68

4 m, 5 f

JJ

Kane, Johnson Canyon

30-Vin-53

4 f

VT

Millard, Delta

2-VL68

2 m, 9 f

JJ

14 mi S Deseret

4-VL68

3 m, 10 f

JJ

Piute, Kingston

25-VL52

1 m, 1 f

DB

Salt Lake, Salt Lake

22-VL30

1 m, 2 f

JR

San Juan, Navajo Mt.

Trading Post

5-V-55

1 m, 1 f

CH

Canyonlands Nat. Park

(Needles campground)

22-VI-68

3 m, 18 f

JJ

Hatch Trading Post

23-VL68

1 f

JJ

Sevier, Richfield

19-VI-47

1 f

DB

15-V-40

1 m, 5 f

WG

Sevier Canyon

19-Vn-40

1 f

WG

2 mi E Glenwood

30-VL40

1 f

WG, LH

Fishlake

22-VL30

1 m

WG

Paradise Valley

lO-Vin-52

1 f

MK

Tooele, Skull Valley

19-IV-39

1 f

GK

Desert Range Exp. Sta.

9-XL50

2 f

DB

Knolls

5-Vn-68

1 m, 9 f

JJ

Delle

5-Vn-68

6 m, 9 f

JJ

Uinta, Dinosaur Nat.

Mon. (Quarry)

21-V-50

2 f

VT

6-Vn-68

19 m, 26 f

JJ

Utah, Provo

XL28

1 m

VT

Washington, St. George

19-IIL31

1 m, 1 f

VT

29-in-31

2 f

VT

Zion Nat. Park

9-VL68

3 f

JJ

Wayne, Fruita

17-Vn-31

2 f

WG

Torrey

17-VI-38

3 f

DB, VT

7-Vin-52

1 f

MK

166

GREAT BASIN NATURALIST

Vol. 32, No. 3

Table 6. Length of the carapace, preabdomen, postabdomen, and
telson of 20 Vaejovis boreus from northern and 20 from southern
Utah, with computation of the mean and standard error.

Northern Utah

Southern Utah

Car.

Preab.

Postab.

Tel.

Car.

Preab.

Postab.

Tel.

4.9

16.4

18.0

6.0

5.0

15.1

19 2

6.0

4.9

14.5

16.9

5.6

5.3

16.2

197

6.0

6.0

17.0

23.3

7.0

5.7

11.3

21.2

6.2

4.6

13.9

14.5

4.7

5.6

12.9

20.0

6.9

5.5

17.7

20.0

6.1

5.6

7.8

19.0

6.1

6.0

19.1

21.9

6.9

6.4

12.7

24.0

7.5

6.4

19.4

21.4

6.8

5.9

10.6

22.5

6.7

5.5

13.8

19.0

6.0

5.3

13.8

21.5

6.3

5.8

15.5

21.3

6.0

5.7

15.3

21.8

6.3

5.8

13.0

19.5

6.1

5.7

13.2

22.8

6.7

5.8

15.2

20.6

6.4

5.2

13.0

19.2

6.0

5.7

19.0

20.5

6.0

5.2

13.0

19.4

5.9

4.9

13.8

18.2

5.7

4.5

11.2

17.8

5.2

5.3

13.5

19.9

6.4

5.3

15.0

21.4

6.3

5.4

15.0

19.0

5.8

4.9

14.6

19.0

5.7

6.0

15.6

21.0

6.9

4.9

12.3

13.5

5.7

5.7

17.9

19.3

6.2

4.8

14.5

16.9

5.2

5.0

12.0

16.9

5.3

6.7

18.9

22.8

6.9

5.6

16.8

19.3

6.2

6.7

12.6

22.6

6.6

6.4

17.7

23.0

7.3

5.8

15.9

20.0

6.1

Mean

5.53

15.84

19.68

6.17

5.51

13.50

20.47

6.22

SE

±.26

±.99

±.97

±.28

±.27

±1.08

±.87

±.26

coarsely granulated pedipalps. The base color is pale yellow. Speci-
mens from northern Utah, when compared with those from southern
Utah, were not significantly different at the 95 percent level (Figs.
20-23, Table 6) in the length of the carapace, postabdomen, and
telson. However, a significant difference of 0.27 mm in the length
of the preabdomen was present between j)opulations.

Vaejovis confusus Stahnke

Vaejovis confusus Stahnke, 1940, Iowa State College J. Sci. 15:101.

This species is known from southern Nevada (Gertsch and
Allred, 1965) and southern California (Williams and Hadley, 1967).
This is the first time V. confusus has been reportetl from Utah.
Twenty-three specimens representing adults and innnatures were
collected from localities in the Great Basin and the ('olorado River
Basin. Males and females were taken in a ratio of 1:3.6. This
species occurs along the entire length of the state (Fig. 24, Table 7).

Sept. 1972

JOHNSON, ALLRED: SCORPIONS

167

24

23

22

21

II i

.

20

\ ^

19

1

18

17

S

16

15

N

19

17

1

16

I 1

15

1

1

14

13

» 1

12

11

10

9

8

S

■^

22

I!

20

21

23

Figs. 20-23. The mean, standard deviation and range of length (in mm)
of the (1) postabdomen; (2) preabdomen; (3) carapace; (4) telson of the Vac
jovis boreus from northern and southern Utah.

Table 7. Collection data for Vaejovis confusus in Utah.

County and location

Date

No. and sex

Collector

Box Elder, Lucin

7-VII-68

2 f

JJ

Grand, Arches Nat. Mon.

20-VI-68

1 m

Unknown

Millard, Holden

l-VI-68

4-f

JJ

Clear Lake

3-VI-68

3 m, 5 f

JJ

San Juan, Canyonlands

Nat. Park (Tower Ruins)

22-VI-68

1 m, 4 f

JJ

Monument Valley

24-VI-68

1-f

JJ

Utah, Orem

20-V-68

1 f

SO

Washington, St. George

5-VI-68

1 f

JJ

Vaejovis confusus closely resembles V. wupatkiensis in general
appearance, but V. confusus is slightly larger and lighter in color.
It may be distinguished from other species in Utah by the lack of
definite markings on the body, a relatively thick, fifth caudal
segment, relatively long pincers on the pedipalps, and a coarsely-
granulated, robust telson bearing a short sting.

168

GREAT BASIN NATURALIST

Vol. 32, No. 3

V. confusus

V. wupatkiensis

V. utahensis

Fig. 24. Distribution of Vaejovis confusus, V. wupatkiensis and V. utahensis
in Utah.

Vaejovis wupatkiensis Stahnke
Vaejovis wupatkiensis Stahnke, 1940, Iowa State College J. Sci. 15:101.

This species is known from northern Arizona, and westward into
Nevada and adjacent Cahfornia (Gertsch and Allred, 1965). This
is the first time it has been reported in Utah. The eleven females

Sept. 1972 JOHNSON, ALLRED: SCORPIONS 169

were taken only from the Colorado River Basin in Utah (Fig. 24,
Table 8).

Vaejovis wupatkiensis is a slender scorpion, similar to but smaller
than V . confusus. It lacks the coarse granulation of the cauda and
telson conm^ion to V . confusus. It may be distinguished from other
species in Utah by its uniform yellow to orange-brown color, absence
of any markings on the body, long fingers on the pedipalps, and a
tubercle or tooth beneath the base of the vesicle on the telson.

Table 8. Collection data for Vaejovis wupatkiensis in Utah.

County and location

Date

No. and sex

Collector

Grand, Moab

VI-23

2-f

cc

V-55

5-f

CH

Thompson

3-V-68

1 f

FC

San Juan, Navajo Mt.
Trading Post

2-V-55

2 f

DB

Canyonlands Nat.
Park (Tower Ruins)

VI-67

1 f

PH

Vaejovis utahensis Williams

Vaejovis utahensis Williams, 1968, Pan-Pacif. Entomol. 44(4):313-321.

This species was heretofore known only from Bluff, San .luan
County, Utah (Williams, 1968). An additional 218 specimens rep-
resenting adults and immatures were taken from the Colorado River
Basin in Utah (Fig. 24, Table 9).

Table 9. Collection data for Vaejovis utahensis in Utah.
County and location Date No. and sex Collector

Garfield, 35 mi W Dirty

Devil River on Utah

Hwy 95

25-VI-68

5 f

Grand, Dead Horse Pt.

Road

19-VI-68

25 f

Emery, 30 mi N Hanks-

ville on Utah Hwy 24

25-VI-68

2 f

San .luan. Monument

Valley

24-VI-68

56 f

Bluff

23-VI-68

26 f

Canyonlands Nat. Park

(Needles campground)

20-VI-68

50 f

JJ

lO-VI-68

54 f

BB

170 GREAT BASIN NATURALIST Vol. 32, No. 3

1 his species closely resembles V. boreus, but may be distinguished
from it by the absence of the dusky, transverse bands on the seg-
ments of the preabdomen, and the longer, less robust hands of the
pedipalps.

LITERATURE CITED

Allred, D. M. 1968. Ticks of the National Reactor Testing Station. Brigham

Young Univ. Sci. Bull., Biol. Ser. 10(1): 1-29
Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. Biotic communities

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

ll(2):l-52.
Banks, N. 1900. Synopsis of North-American invertebrates. The scorpions,

solpugids, and pedipalpi. Amer. Nat. 34(9) :421-427.
CoMSTOCK, J. H. 1948. The Spider Book. Revised and edited by W. J. Gertsch.

Comstock Publishing Co., Inc., New York.
DuRRANT, S. D. 1952. Mammals of Utah, taxonomy and distribution. Univ.

Kansas Pub., Mus. Nat. Hist. 6:1-549.
EwiNG, H. E. 1928. The scorpions of the western part of the United States.

with notes on those occurring in northern Mexico. Proc. U.S. Nat. Mus.

73(9):l-26.
Gertsch, W. J., and D. M. Allred. 1965. Scorpions of the Nevada Test Site.

Brigham Young Univ. Sci. Bull., Biol. Ser. 6(4): 1-15.
Gertsch, W. J., and M. Soleglad. 1966. The scorpions of the Vaejovis boreus

group (subgenus Paruroctonus) in North America (Scorpionida, Vaejovidae).

Amer. Mus. Novitates 2278:1-54.
Hayward, C. L., D E. Beck and W. W. Tanner. 1958. Zoology of the Upper

Colorado River Basin. Brigham Young Univ. Sci. Bull., Biol. Ser. 1(3): 11-74.
HoNETSCHLAGER, L. D. 1965. A uew method for hunting scorpions. Turtox

News 43 (2): 69-70.
PococK, R. L. 1902. Scorpions, Pedipalpi and Solfugae. Biologia Centrali

Americana 71:12.
Stahnke, H. L. 1940. The Scorpions of Arizona. Iowa State College J. Sci.

15:101-103.
Williams, S. C. 1966. Burrowing activities of the scorpion Anuroctonus

phaeodactylus (Wood) (Scorpionida: Vaejovidae). Proc. California Acad.

Sci., Fourth Ser. 34(8) :419-428.
Williams, S. C. 1968. Two new scorpions from western North America. Pan-

Pacif. Entomol. 44(4) : 313-321.
Williams, S. C, and N. F. Hadley. 1967. Scorpions of the Puerto Penasco

area (Cholla Bay), Sonora, Mexico, with description of Vaejovis baergi.

new species. Proc. California Acad. Sci., Fourth Ser. 35(5) : 103-1 16.
Williams, S. C. 1971. Clarifications in the nomenclature of some North American

Scorpionida. Pan-Pacif. Entomol. 47(1): 78-79.

RELICT STANDS OF PINYON HYBRIDS
IN NORTHERN UTAH' -

Ronald M. Lanner ■ and Earl R. Hutchison-'

Abstract. — Relict stands of Pinus edulis and P. monophylla, and of natural
hybrids of these species have been found in Cache and Rich counties. The stands
and their sites are described. The P. edulis stand in Rich County extends the
range of this species northward. Possible means of seed dissemination are dis-
cussed and it is speculated that both species formerly migrated into and out of
this area during periods of differing climates.

Two pinyon pine species are native to Utah: pinyon (Pinus
edulis Engelm.) and singleleaf pinyon. (P. monophylla Torr. &
Frem.). Pinyon is typically a species of the Colorado Plateau,
while singleleaf pinyon is characteristic of the Great Basin. Cole^
postulated that the two species hybridize along an ecotone running
north and south through central Utah. Recent field studies have con-
firmed that these pines hybridize along the eastern edge of the Great
Basin and in other areas where their ranges overlap (Lanner, 1971).

Hybrid segregates have also been discovered in Cache and Rich
counties, in a series of relict stands previously thought to be well
outside the range of P. edulis (Critchfield & Little, 1966). The
evidence for hybridization will be fully covered in a later paper.
Major criteria are number of needles per fascicle (P. edulis ^= 2;
P. monophylla = 1; hybrids have fascicles of both types) and num-
ber of resin canals per needle (P. edulis = 2; P. monophylla = 2 -
10; hybrid means are intermediate). The aims of this paper are to
describe the northern relicts and to explain their supposed origin.

The Populations

Four of the relict stands are in Cache County, and one is in the
Crawford Moimtains of Rich County.

1. Mt. Logan. Eleven trees on a spur between Logan and Dry
canyons, in Sec. 31 and 32, TI2N, R2E, Cache National Forest. Of
10 trees examined, 8 appear to be pure P. monophylla, but the
others have 11-35 percent of their foliage in fascicles of two.

2. Logan Canyon. Two trees located 2 miles NE of the above
population, in Sec. 29, T12N, R2E, Cache National Forest. Both ap-
pear to be pure P. monophylla. Reported presence of additional trees
is unverified.

3. Blacksmith Fork. Many trees scattered in drainage of Mollen's

^This research is supported by funds provided through the Mclntire-Steiuiis Cooperative Forestry
Reseanh Program. It is approved as journal paper no. 1270 of the Utah Agricultural E.xperinient
Station

-An abstract of this paper was presented at the Utah Academy of Sciences, Arts, and Letters
meeting at Logan in September 1971.

^Department of Forest Science, Utah State University, Logan, Utah 84321.

■•Cole, Franklin R. The pharmacognosy of Utah pinyon pines. Doctoral dissertation, University
of Utah, 1965.

171

172 GREAT BASIN NATURALIST Vol. 32, No. 3

Hollow, Sec. 27, 28, 29, 32, 33, 34, TION, R4E, Cache National
Forest. Most are pure P. monophylla, but some have up to 68 per-
cent of their foliage in fascicles of two.

4. Porcupine. At least 29 trees, N of Porcupine Reservoir in
Sec. 10 and 11, T9N, R2E. Trees range from pure P. monophylla
to individuals with 90 percent of their foliage in fascicles of two.

5. Crawford Mountains. Eleven trees were sampled 4 miles E
of Randolph in a box canyon on the western scarp of Rex Peak in
Sec. 30, TUN, R8E. Trees range from pure P. monophylla to pure
P. edulis, with individuals of intermediate character.

Unlike the other populations, this is mainly a P. edulis stand and
represents the northernmost limit of the species.

Locations are given in reference to the Salt Lake Meridian. They
fit within a rectangle 34 miles long (E - W) and 15 miles wide (N -
S), the NW corner of which lies at 41°45'30"N and 111°45'30"W;
and the SE corner at 4r31'30"N and 111°05'30"W.

Growth Habits

The relict stands consist of small groups of relatively slow-
growing trees. They are usually on south-facing slopes within a
narrow elevational range and associated with tree species that
characterize the transition from relatively xeric to subhumid con-
ditions (Table 1). The pinyon pines seem unable to compete with
lush north-slope tree and brush cover and are mainly restricted to
drier aspects where plants are widely spaced between large areas of
bare ground. The limited area of these stands and their age dis-
tribution suggests a precarious balance between survival and ex-
tinction. Though regeneration is sporadic, some trees are long-lived
and have many opportunities to reproduce (Table 1). It is possible
that the P. edulis trees, growing at the northern limit of their range,
may be reproductively impaired by pollen that is rendered inviable
by low temperatures at critical times in its development (Chira.
1967). In very small populations, the high incidence of self-polli-
nation may result in the production of inviable seed or non vigorous
seedlings.

Origins

The nearest station of P. monophylla is in the Bhuk Pine Moun-
tains of southern Idaho, about 70 miles WNW of Logan. ' The near-
est P. edulis stands are about 100 miles S of Randolph, in the Uinta
Basin. The disjunct stands are either old relicts, left behind by wide-
s[)read extinction, or they represent new invasions.

Invasion seems unlikely. Pinyon pine seeds are wingless, heavy,
and ballistically unadapted to wind dissemination. Experiments in
this laboratory by R. Warnick indicate that a wind of 100 mph is
required to carry P. monophylla seeds 30 ft downwind from the

*Critchfield and Little (1966) show P. monophylla extensively distributed in the northern
Wasatch, straddling the Utah-Idaho state line, but this report appears to be in error

Sept. 1972 L.-KNNER. HUTCHINSON: PINYON HYBRIDS

173

• 5 '^S"

CO HH

^5! Sd';;^

c.S v f

CO -P 4 >--

S-S

teosperma
opulorum
lenziesii
■difolius
s flexilis
ilus tremu

d

opulorum
lenziesii
difolius
exilis

1

S ^ S ^ 1 1

i

§ £^ ^

§

-; -; ex: o a, Q-

■^

>-; o; o ai

-^

^

w

K

w"

^-

^.

CO

C/5

z

CL, o

I 2

6 o

6 o

s s

..

i2

Kl

S

4-. t- &

a;

S

-d

■a

g

1

iM

CO

U(

d.

U

174 GREAT BASIN NATURALIST Vol. 32, No. 3

top of a 30-ft tree. Long distance dispersion by birds or mammals,
though a possibility, seems unlikely to account for two tree species
on sites far removed from both present distribution areas, one to the
west, the other to the south.

The movement of seeds by man is a possibility worth consider-
ation. Wright^ concluded that a disjunct P. edulis stand in Owl
Canyon, near Fort Collins, Colorado, resulted from "accidental or
intentional planting that took place approximately 400 years ago"
along an ancient Indian trail. Acceptance of this interpretation has
led Weber (1961, 1965) to explain other trees subsequently
found nearby in the Rockies as having originated from seed
transported there, by birds, from the Owl Canyon grove. It seems
equally logical to consider all the northeast Colorado pinyon lo-
cations as relicts of a once-continuous distribution from the south,
but the evidence on this point is by its very nature inconclusive.

In the present case, man seems an unlikely agent. The Paiute
and Shoshoni of the Great Basin often set up their temporary winter
villages in pinyon pine areas, in order to take advantage of the pine-
nut crop (pers. comm. Prof. Julian H. Steward, Univ. of Illinois).
Thus, the Indians went to the trees, rather than attempting to plant
them. The pinyon stands at Mt. Logan and Logan Canyon are
on steep ridges, unsuitable as travel routes or campsites where caches
might be established. And, again, the coincidence of establishing
two species on the same sites seems farfetched.

We lean toward an explanation based on paleoclimates. There is
a rough parallel here with the situation described by Cottam et al.
(1959). According to their analysis, hybrid clones of Quercus gam-
belii Nutt. and Q. turbinella Greene originated early in the latest
postglacial period, when warmer climates in the north permitted
Q. turbinella to persist well outside its present range. Using a simi-
lar line of reasoning, we propose the following sequence of events
to account for the pinyon pine relicts of northern Utah:

(a) P. monophylla migrated into the northern Wasatch and
Crawford mountains at a time when it was cooler and moister
than now. At this time, P. monophylla may have been continuous
along the north rim of the Great Basin from the Raft River Moun-
tains, across the low hills on the north shore of Lake Bonneville, to
the Wasatch.

(b) P. edulis migrated north on the Wasatch from the Uinta
Basin when it was warmer than now. The species interbred. It is
interesting in this connection that an outpost of Gambel oak has
been found several miles east of the Porcupine location where
there are trees appearing to be nearly pure P. edulis. In the Uinta
Basin these species are sometimes associated with each other.

(c) Later changes caused the following to occur:

i. P. monophylla became extinct at lower elevations, leaving

•Wright, C. W. An ecological description of an isolated pinon pine grove. Master's thesis,
University of Colorado, 1952.

Sept. 1972 LANNER, HUTCHINSON: PINYON HYBRIDS 175

populations that have survived to become its major stands on the
Great Basin ranges, and relicts on the Wasatch.

ii. P. edulis retreated southward, persisting in a few suitable
areas where competition was less severe. Thus, both species are
present only as relicts. The species have an overlapping range of
tolerance to environmental conditions, as attested by their frequent
sympatry, so it is not surprising they should share common refugia.
Admittedly, this interpretation of these relict stands is speculative,
but other interpretations require assumptions that are equally specu-
lative.

Literature Cited

Chira, E. 1967. Pollen grains of Pinus edulis with mor^ than the haploid

number of chromosomes. Silvae Genet. 16(1):14-18.
CoTTAM, W. P., J. M. Tucker, and R. Drobnick. 1959. Some clues to

Great Basin post pluvial climates provided by oak distributions. Ecology

40(3):361-377.
Critchfield, W. B., and E. L. Little, Jr. 1966. Geographic distribution of the

pines of the world. U. S. Dept. Agr. Misc. Publ. 991, 97 pp.
Lanner. R. M. 1971. Conifer flora of the Bear Lake area and mountains south

of the Great Salt Lake. Great Basin Nat. 31 (2):85-89.
Weber, W. A. 1961. Additions to the flora of Colorado. Univ. Colorado Stud.,

Ser. Biol. 7:1-26.
. 1965. Plant geography in the Southern Rockies, p. 463. In Wright,

H. E., Jr., and Frey, D. G., Editors, The Quaternary of the United States:

Princeton Univ. Press, 922 pp.

NEARCTIC DESERT DECTICIDAE (ORTHOPTERA)
PARl^ I. A NEW GENUS FROM CALIFORNIA

Ern-st R. Tinkham'

Pctropedes. new genus

This genus appears to have closest relationships to Inyodectes
Rentz and Burchim from which T differs by much larger size,
strongly excavate inferior margin of the lateral lobes of the prono-
tum, the enormous tympanum, the different cerci, bispinose genicular
lobes of the pro- and mesoTemora, the unispined external genicular
lobe of the caudal femora as well as other features; from Eremopedes
Cockerell by the differently shaped pronotrm and the emargination
of the inferior margin of the lateral lobeii of the pronotum, by the
much larger tympanum, by the bispinose and not unispinose genic-
ular lobes of the pro-femora, by the less prominent emargination
of tho ultimate notite of the abdomen with less forcipate arms, by
the cerci and subgenital plate and by the ovipositor; from Oreopedes
Rehn and Ilebard by size, pronoLum, tympanum, spination of the
genicular lobes of all femora and many other features; and from
other cremicolous genera by even more striking dissimilarities.

Diagnosis: — Size, medium large; antennae, at least three times body
length. Pronotum, about twice as long as broad, rounding into rather deep
lateral lobes having inferior margin strongly excavate to expose enormous
tympanum. Tegmina exposed for a length equal to that of metazona, its base
pale, its apical portion jet black, and diagnostic. Ultimate notite of abdomen
v.'ith median, moderately deep-shaped notch formed by moderate forcipate di-
varicating arms, much shorter than thos? of Eremopedes. Cerci moderately
slender with one internal triangular, almost central, projection bearing a piceous.
thin plate with 6-8 fine serrate teeth directed anteriorad. Subgenital plate
subtriangular in male, with rounded ventro-lateral ridges and a shallow median
posterior notch. Pro- and meso-femora wit.h bispinose genicular lobes. Ovi-
positor, about length of body, strong and slightly recurved in apical half.
Subgenital plate in female with a very shallow median posterior emargination.

Description:- — Head slightly broader than deep (at clypeal
suture); eyes roundly oval and subglobose; antennae at least three
times body length. Pronotum with barrel long, rounding into lateral
lobes, about twice as long as broad; anterior and posterior margins
squarely truncate. Prozonal sulcus a very broad v-shaped shallow
groove running to near fore margin of lateral lobes, thence continuing
as a very broad shallow groove submarginally to almost posterior
margin. Prozonal-metazonal juncture, dorsally, with a slight horse-
shoe-shaped depress'on. True tymi)aimm very large, roundly oval,
fore margin slightly irregular due to |)rothoracic spiracle; its margin
devoid of hairs. Tegmina exposed for a length equal to metazonal
length. Abdomen with ultimate notite having divaricating forcipate
arms forming a moderately deep v-shaped emargination. Cerci, diag-
nostic, relatively slender, with internal, almost median, triangular

'81-441 Date Palm Avenuo. Iiul.... Cnlifoinia.

176

Sept. 1972 TINKHAM: DESERT ORTHOPTERA 177

prominence bearing a dark sclerotized thin plate with 6-8 serrate,
fine, teeth. Titillator arm straight, vent'al keel very slightly curved
apically, dorsal keel with row of rotro.-se teeth. Subgenital plate
narrowly triangular with round ventro-latoral ridges and a shallow
U-shaped emargination posteriorly.

Leg spination: Forelegs with strong j^rocoxal spine; fore femora
with 3 strongly appressed aciculate teeth on central portion of ex-
ternal and internal ventral keels plus bispinose geniculae; fore tibiae
with six pairs of long acuminate spines on the ventral keels and three
dorsal external spines (one on so called tympanum of authors, one
median and one apical). Tarsi 3-segmentcd with ungues segment.
Meso-femora with four pairs of ventral sjjinos in apical two-thirds
plus bispinose geniculae; mesotibiae ventrally as in protibiae, dorsally
with 2 anterior and 4 posterior dorsal spines (leg in back position).
Caudal femora with 6-8 external and 6-7 internal spines on
ventral keels all in apical half. Caudal tibiae with 25-26 external
and 23-25 internal teeth on ventral keels and 7 external and 3-5
internal dorsal apical teeth. Leg spination: forelegs with strong
procoxal s})ine; fore femora with 3 strongly appressed aciculate
spines on central portion of internal inferior keel plus bispinose
geniculae; fore tibiae with six pairs long acuminate spines on ventral
keels and three dorsal external spines, one on so-called tympanum of
authors, one median and one apical. Tarsi 3-segmented plus
ungues segment. Meso-femora with four pairs ventral spines on the
apical two-thirds plus bispinose geniculae; mesotibiae ventrally as
in protibiae; dorsally with two external and four internal spines
(leg in back position). Caudal femora with 6-8 external and 6-7 in-
ternal teeth on ventral keel all in apical half. Caudal tibiae with
25-26 external and 23-24 internal ventral teeth and 7 external and
3-5 internal dorsal teeth.

Type species. — Petropedes santarosae Tinkham.

Petropedes santarosae, n. sp.

Coloration generally tawny or pale reddish brown, profusely
mottled on abdominal notites with dark gray flecking, especially
along posterior marginal area. Internal musculature of legs indi-
cated by fine grayish infuscation. Tegmina pale with black in-
fuscation in apical third, leaving posterior marginal area pale buff.

Male Holotype: — Santa Rosa Mountains, Palms to Pines Hiway, 2000
feet elevation, 9-VIII-70; E. R. Tinkham; crossing pavement at night. Calipered
measurements m mm: body length 24.8; head 5.0 x 6.8 at clypeal suture;
pronotum 8.6 x 6.8; caudal femora 26.6; antennae approx 77.0. Type deposited
in the Tinkham Eremological Collection.

Female description: — Size slightly larger than holotype; ovi-
positor as described. Subgenital plate rounding without ventro-
lateral ridges as in male; posterior margin circularly rounded with
very shallow v-shaped emargination.

Female allotype: — Same data as holotype but collected 5-VII-69. Cali-
pered measurements in mm: body length 29.9, length to ovipositor 46.8;

178

GREAT BASIN NATURALIST Vol. 32, No. 3

Figs. 1-5. Petropedes santarosae Tinkhani: 1. male holotype; 2, genitalia
of male holotype showing ultimate notite. cerci. and subgenital plate; 3, lateral
aspect of pronotum and tympanum of male holotype; 4. lateral aspect of ovipositor
of female allotype; 5. posterior portion of subgenital plate of female allotype.

ovipositor 20.2; pronotum 8.2 x 7.2 in maximum breadth at inferior marginal
flange. Allotype in the Tinkham Eremological Collection.

Male p.aratypes: — Same data as holotype except: 3. 26-VI-66; 4, 26-VII-66;
3, 9-VII-70; 2. lO-VII-70. All male paratypes very closely similar to holotype
in every respect. Calipered range measurements in mm as follows: body
length 123.8-25.0; pronotum 8.1 x 5.9 to 9.2 x 7.0 breadth; caudal femora
25.8-29.9, antennae approx. 86.8. Paratypes to be exchanged with major
Orthopterological museums.

Sept. 1972 TINKHAM: DESERT ORTHOPTERA 179

Female paratypes: — Closely similar to allotype except: 2, 26-VI-66; 1,
3-VII-67; 2, 9-VII-70; 1, 4-VII-70 (after light rain). Calipered range measure-
ments in mm: Body length 25.0-30.0; length to apex of ovipositor 41.0-46.8;
ovipositor 23.0-18.8; pronotum 9.5 x 7.5 to 9.1 x 7.3; caudal femora 31.1-
28.8. Deposition as noted above.

Habitat: — Petropcdes inhabits the steep, rocky, northern slopes
of the Santa Rosa Mountains in a rather narrowly defined alti-
tudinal zone ranging from 1900 to 2200 feet elevation; the optimum
being about 2000 feet. This belt, although within the areal limits
of the Colorado Desert, is, duo to its considerable elevation, more
accurately Gila Desert because of the presence of the indicative
Mohave yucca {Yucca schidigera) . Other dominants and codomi-
nants include: desert agave {Agave desert i) ; cheese bush {Hymen-
oclea salsola) ; desert sweet {Bebbia juncea) ; desert sunflower
{Vigueria dehoides) and other rarer plants that occupy their niches
in and among the boulders and rocks of this habitat. Even more
preferred are the rocky road cuts of this highway for Petropedes
is truly a rock-inhabiting decticid.

Biology: — Little is known about their life history at present,
as the creature was only found crossing the pavement at night.
Its saltatorial powers arc considerable. Its song is very weak which
may explain the enormous tympanum. The eggs probably hatch
during the winter rains. In 1971, a drought year, collecting failed
to reveal any specimens in the optimum zone of habitation. In
a xeric environment the Orthopteran associates included: Ateloplus
notatus^ Capnobotes fuliginosus, Poecilotettix sanguineus, Oedomerus
sp., Melanoplus sp., and Stagmomantis calif ornicus .

Petropedes spinosa (Hebard), n. comb.

Eremopedes spinosa Hebard, 1923, Proc. California Acad. Sci. 12(15) :337 figs.
10-13.

For many yea -s the identity of this Mexican decticid described
from Mejia Island in the Gulf of California has been a mystery.
There is now no question that it is a member of the genus Petropedes
Tinkham but at present nothing is known about its biology and
little about its habitat.

Literature Cited

Hebard. M. 1923. XV. Expedition of the California Academy of Sciences to the
Gulf of California in 1921. Proc. California Acad. Sci., Fourth Series 12(15):
337-339, figs. 10-13.

Rentz, D. and J. D. BuHCHiM. 1968. Revisionary studies in the Nearctic
Decticinae. Memoirs Pac. Coast Ent. Soc. 3:1-173, 37 figs.

Tinkham, E. 1944. Biological, taxonomic and faunistic studies on the shield-
back katydids of the North American deserts. Amer. Midi. Nat. 31(2):
257-328. 27 figs.

■ I960. Studies in Nearctic desert sand dune Orthoptera. Part I. A new

species of Plagiostira from eastern New Mexico with key and notes. Great
Basin Nat., 20:39-48, 10 figs.

GREAT BASIN NATURALIST
Notice to Contributors

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TABLE OF CONTENTS

On the rodent-infesting Anoplura of Panama. Phyllis T.

Johnson 121

Notes concerning Mexican Saldidae, including the descrip-
tion of two species (Hemiptei-a). John T. Polhemus 137

Scorpions of Utah. John D. Johnson and Dorald M. Allred .... 154

Relict stands of pinyon hybrids in northern Utah. Ronald

M. Lanner and Earl R. Hutchinson 171

Studies in neartic desert Decticidae (Orthoptera: Tettigoni-

oidea). Part I. Ernest R. Tinkham 176

Volume 32, No. 4
December 31, 1972

The

MUS. COMP. ZOOLl
LIBRARY

MAR 2 3 1973

HARVARD
UNIVERSITY

Great Basin

Published by
Brigham Young University

GREAT BASIN NATURALIST

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

Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W.
Tanner, Zoology; Joseph R. Murdock, Botany; Vernon J.
Tipton, Zoology; Ferron L. Andersen, Zoology

Ex officio Editorial Board Members:

A, Lester AUen, Dean, Collie of Biological and Agricul-
tural Sciences

Ernest L. Olson, Director, University Press, University
Editor

The Great Basin Naturalist was founded in 1939 by Vasco M.
Tanner. It has been continuously pubKshed from one to four times
a year since then by Brigham Young University, Provo, Utah. In
general, only original, previously unpubhshed manuscripts pertain-
ing to ihe biological natural history of the Great Basin and western
North America will be accepted. Manuscripts are subject to the ap-
proval of the editor.

Subscription: The emnual subscription is $5.00 (outside the
United States $5.50). The price for single numbers is $2.00 each.
All matters pertaining to the purchase of subscriptions and back
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Young University Exchange Librarian, The Library, Provo, Utah
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Manuscripts: All manuscripts and other copy for the Great
Basin Naturalist should be addressed to the editor. Contributors
should consult the instructions printed on the back cover of a recent
number.

The Great Basin Naturalist

Published at Provo, Utah, by

Brigii AM Young University

Volume 32 December 31, 1972 No. 4

AQUATIC PHYCOMYCETES OF LILY LAKE, UTAH^

Hugh M. Rooney- and Kent H. McKnight^

Abstract. — During the ice-free period from May to November on two
successive summers v^ater molds were collected on 14 different substrata placed
in Lily Lake, a subalpine bog lake in the Uinta Mountains of Utah. Twentj'-
five collections yielded thirty-four species in 20 genera, 1 1 families, and 6 orders
of aquatic Phycomycetes. Correlation of frequency presence of the species re-
ported with physical and chemical characteristics of the lake showed that the
number of species collected increased with an increase in water temperature
and a decrease in oxygen content during June to August, reaching a peak in
late July. Rhipidium americanum Thaiter (100 percent frequency) and Sapro-
myces androgynus Thaxter (72 percent frequency) were the most common spe-
cies and the species found on more different substrata than any others. Also
in the "very abundant" group were Saprolegnia ferax (Gruith.) Thurent and
Achlya sp.. both with 64 percent frequency.

Lily Lake, a subalpine lake in the Wasatch National Forest of
the Uinta Mountain region, Utah, is located about one-half mile west
of Trial Lake at an elevation of 10,000 feet (T. 3, R. 9 E., sec. 31).
This lake, which is one of the acidic, cold, lentic bogs characteristic
of the area, is the site of a taxonomic-distribution study of aquatic
Phycomycetes. This study is a beginning to our understanding of the
role of fungi in Lily Lake, and it supplies the first information on
record of the water molds of this area. Fungi found on 14 different
substrata are identified and correlated with pH, dissolved oxygen,
and temperature of the lake.

Christensen and Harrison (1961) have described the psysiogra-
phy and possible plant succession around Lily Lake. Stutz (1951)
studied the hydrarch succession as well as physical and chemical
properties of water and soil at Moss Lake, a similar subalpine lake
in the Granddaddy Lake Basin nearby. Tanner (1931) reported on
the algae of the Mirror Lake region in the Uinta Mountains.

Little is known about the fungi in subalpine lakes. However,
Tiesihausen (1912) and Rutter (1937) reported on the alpine water
molds of the Swiss Alps. Koob (1966) reported Rhizophidium plank-

'This paper intludes a portion of a thesis piesented in partial fulfillment of the requirements for
a Master of Science degree, Brighaiii ^■ounK ITmversity.

'RioloRv Oepaitnient. Virginia I'olytechni( Institute and State University, Blarkshurg, Virginia.

^IMant Protection Institute, Agricultural Research Serviie, US. Department of .Agriculture,
Beltsville, Maryland 20705.

181

182 GREAT BASIN NATURALIST Vol. 32, No. 4

tonicum Canter on AsterioneUa formosa Hauss. in subalpine lakes of
Colorado. Studies by Chapman (1965j, Coker (1923), Jewell and
Brown (1929), and Willoughby (1961) report fungi in bogs, and
there are a few papers showing seasonal distribution of aquatic
Phycomycetes: Coker (1923j, Suzuki (1960, 1961), and Willoughby
(1962).

Water molds are given little consideration in general treatments
of aquatic ecology (Welsh, 1945; Weston, 1941). However, repre-
sentatives of these fungi have been found in all types of inland
waters that have been studied for them (Crooks, 1938; Graff, 1928;
Johnson, 1956; Koob, 1966; Sparrow, 1960; Suzuki, 1960a, 1961a;
Suzuki and Nimura, 1960, 1961). As saprophytes and parasites on
aquatic plants and animals, they are important in biological inter-
actions in lakes.

Methods and Materials

At approximately weekly intervals during the study period, water
molds were collected and environmental data of the water were
recorded. Baits were placed in the lake during the ice-free period
from May 1965, to November 1965, and from May 1966, to June
1966. The 14 substrata used for bait may be classified in four major
groups as follows: (i) twigs (river birch, weeping white birch, pop-
lar, hackberry, pine); (ii) fruits (apples, rose hips); (iii) chitinous
and keratinous materials (human hair, snake skin, insect exuviae);
and (iv) miscellaneous materials (cellophane. Ulothrix. pine pollen,
pine needles). Of these, the twigs and fruits were collected in the
autumn preceding their use and stored at 5 C until needed.

A wire basket containing the baits was placed II/2 feet below the
surface of the lake each week and removed four weeks later. When
removed from the lake the basket was transferred to a glass jar and
brought to the laboratory, where the baits were rinsed with tap water
and examined for aquatic Phycomycetes. Both temporary water
mounts and i)ormaneiit mounts were prepared for microscopic ex-
amination of the fungi. After this initial examination the baits were
cultured in large jjlastic dishes, using water collected that week from
the lake. Such additional substrata as human hair, pine pollen, and
insect exuviae were added to the cultures, which were then covered
with a glass plate and cultured at 5 C. These cultures were examined
three to seven days later for chytrids and other microscopic species.

For any one collection, species of Phycomycetes were listed
simply as present or absent, and no attempt was made to estimate
the rumiber of thalli; therefore, a record of presence may refer to a
single thallus or a great number of thalli. The term frequency, as
used in this study, refers to th(> number of times the species occurred
in 25 collections and is express(>d as a percentage. For example,
Rhipidiurti thnxteri Minden occurred seven times in 25 collections
and has a frequency of 28%.

Water temperature, {)H, and dissolved oxvgen were measured
on each visit to the lake. The pH was measured in the field with a
Beckman pH meter. Model G, except on July 29 arid August 8. 1965,

Dec. 1972 rooney. mcknight: aquatic phycomycetes 183

when it was measured with a Beckman Model K pH meter in the
laboratory. Dissolved oxygen was measured with a Yellow Spring
Instrument Model 51 oxygen meter.'

Results

Thirty-four species representing 20 genera, 1 1 families, and 6
orders of aquatic Phycomycetes were obtained from the 25 collec-
tions made from Lily Lake during the course of this study. On the
basis of frequency, each species is placed in one of four groups as
follows (Table 1): (i) very abundant (frequency 64-100%) — 4
species; (ii) moderately abundant (frequency 20-36%) — 10 species;
(iii) occasional (frequency 12-20%) — 5 species; and (iv) scarce (oc-
curred only once or twice) — 15 species.

Table 2 shows the number of species in each order and the sub-
strata on which each species occurred. Identification of the 11
species of Saprolegniales made this order the most widely represented
among the orders studied. However, they were not the most abun-
dantly occurring species on substrata of categories (i) twigs, (ii)
fruits, or (iii) chitin-keratin. Seven species of Leptomitales were
found on twigs, fruits, pine needles, and insect exuviae. The Chy-
tridiales had six species occurring on keratin, chitin, cellophane,
Ulothrii\ and pine pollen. Four species of Blastocladiales, four spe-
cies of Monoblepharidales, and two species of Peronsporales were
found on twigs and fruits.

The pH of the lake remained acidic throughout the study, with
an average pH of 6 and a range of 5.4 to 6.5; dissolved oxygen con-
tent had an average of 7.2 ppm and a range of 5.5 to 9.2; and water
temperature ranged from 0 C to 18 C for the 1965 season (Figure 1).

Discussion

The 34 species found in this study indicate that many fungi can
grow under the environmental conditions of this subalpine lake, and
their abundance indicates that they may be important in the biologi-
cal degradation of substrata in these areas.

Since a record of species presence in this report might refer to a
single thallus or, in other cases, to a great many thalli, erroneous
conclusions could result. However, although no counts were made
showing the number of individuals present, we noticed that species
for which a high frequency was recorded were generally very abun-
dant whenever found. The species for which a low frequency was
recorded were often represented by few individuals at the time of
collection. These species grew very s])arsely on the bait, and there
was a considerable likelihood that some were overlooked. They were
seen on only a few occasions in a large number of collections.

The species in the "very abundant" and "moderately abundant"
groups were saprophytes and parasites on a wide range of substrata.

■•Mpntion iif ii trademark name or a proprietary prodmt does not
warranty of the product by the USDA and does not iniph- its approval
products that may also be suitable.

184

GREAT BASIN NATURALIST

Vol. 32, No. 4

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TABLE 2. Water molds of Lily Lake and the substrata on which they
occurred.

Species Substrata

Chytridiales
Olpidiaceae

Olpidium endogenum (Braun) Schroeter insect exuviae, pine pollen

Olpidium pendulum Zopf algae

Rhizidiaceae

Rhizophlyctis rosea

(de Bary and Woronin) Fischer cellophane

Chytridiaceae

Chytridium acuminatum Braun cellophane, Ulothriz, baby

hair, pine pollen
Megachytriaceae

N owakowskiella ramosa E. J. Butler cellophane

Megachytrium westonii Sparrow Ulothrix

Blastocladiales
Blastocladiaceae

Blastocladia sp apples

Blastocladia ramosa Thaxter weeping white birch twigs,

apples

Blastocladia angusta Lund weeping white birch twigs,

rose hips

Blastocladia pringsheimii Reinisch apples, rose hips, twigs of

river birch and poplar
Monoblepharidales
Gonapodyaceae

Gonapodya polymorpha Thaxter hackberrj' twigs, rose hips

Gonapodya prolifera (Cornu) Fischer apples

Monoblepharidaceae

Monoblepharis insignis var. insignis

Thaxter poplar twigs, apples

Monoblepharis polymorpha Corne poplar twigs

Saprolegniales
Saprolegniaceae

Phythiopsis cymosa apples, pine needles, twigs of

river birch, poplar and
hackberry

Saprolegnia delica Coker apples, rose hips, insect

exuviae, twigs of river birch,
poplar, and hackberry

Saprolegnia ferax (Gruith.) Thurent hackberry twigs, apples

Saprolegnia hypogyna Pringsheim twigs of river birch and

hackberry

Leptolegniella sp river birch twigs

Leptolegniella keratinophilum Hunoycutt river birch twigs

Protoachlya paradoxa Coker poplar twigs

Achlya sp apples, insect exuviae, twigs

of river birch, poplar, and
hackberry

Achlya klebsiana Pieters twigs of river birch, poplar,

and hackberry

Achlya oblongata de Bary twigs of river birch and

poplar

Achlya americana Humphrey poplar twigs

Leptomitales
Li'ptomitaceae

Leptomitus lacteus (Roth) Agardh hackberry twigs, apples.

insect exuviae
Apodachlya brachynema (Hildeb.) Pringsh. .. river birch twigs

Dec. 1972 rooney, mcknight: aquatic phycomycetes 187

Table 2. Continued

Rhipidiaceae

Rhipidium americanum Thaxter apples, rose hips, pine

needles, twigs of river birch,
weeping white birch, poplar,
hackberry. and pine

Rhipidium interruptum Cornu poplar twigs, apples

Rhipidium thaxteri Minden pine twigs, apples, pine

needles

Sapromyces androgynus Thaxter apples, twigs of river birch,

weeping white birch, poplar,
hackberry, and pine

Sapromyces elongatus (Cornu) Coker rose hips, pine needles, twigs

of poplar and pine
Peronosporales
Pythiaceae

Pythium sp apples, twigs of river birch,

poplar, hackberry. and pine

Phythophthora sp apples, rose hips

By comparison, the "occasional" and "scarce" groups were usually
specific saprophytes and parasites on a limited range of substrata.
The greater numbers of species present during the peak periods
might be a function of abundance and quality of substrata as well as
physical factors of the lake. Thus, abundance of such naturally oc-
curring substrata as lily pads, algae, and pine pollen available in
June, July, and August could account for the presence of some species
which were not found in May, September, and October, when the
substrata were limited or absent.

The number of species collected increased with an increase in
water temperature and a decrease in oxygen content during June to
August, as illustrated in Figure 1. The number of different species
collected was highest in the month of July 1965, and lowest in May
of 1965.

Aspects of the physical environment measured may account in
part for seasonal population fluctuations, but other factors may also
influence them. For example, these fungi seem to infect substrata
with rough surfaces or with broken or decomposed protective outer
layers. Thus Rhipidium americanum Thaxter infected pine needles
only after the cuticle of the pine needle was decomposed or cracked,
calling to mind Emerson's report (1951) that in 9-11 days Blasto-
cladia infected apples that were wiped with ether but took several
weeks to infect untreated apples.

The two families of Leptomitales both have representatives oc-
curring in Lily Lake. However, the species of Leptomitaceae found
in this study were in group iv (scarce) as opposed to species of
Rhipidiaceae, which were all in groups i and ii (abundantly to very
abundantly occurring). Five species in the Rhipidiaceae had a high-
er percentage of occurrence than those of any other family reported
in this study (Table 1). Rhipidium americanum was the most com-
mon species, being found in every week of the study period, and
Sapromyces androgynus Thaxter was the second most common.

GREAT BASIN NATURALIST

Vol. 32, No. 4

TEMPERATURE C

•

.0. ^" r

OXYGEN PPM

/\ [] NUMBER OF SPECIES —-..^

' \ / ^v / \ -^ ^^

8-

\/ \ y \ /

\ \ ' ^ '

f

>.. \ . ^^

6-

••-.. K ,

"■

-^ \-l

&■' • • ••.. ..•

I

v^-

r

V

\

\

z

\

\

<9

\

u

\ /

o

\ ^ /

' \ F

1 /

■o

1 1

\

/

i .

r

j

^ [

y

i

1

\\

2-

[

1

/

M

o-'

a^^-' —

'

/

8 0 24 1 8 15 22 29 6 13 20 27 3 10 17 24 1 8 1

32

9 1

2 9 27

M

ftY ""-JUNE-

■^ '^

- JU

LY-

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-A

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T-

-/\-

SEP

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-l\

3CT

DBE

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•J M

AY ^

-JUNE-

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Fig. 1. Weekly comparisons of water temperature, pH. and dissolved
oxygen in relation to numbers of species.

These two species were found on more different substrata than any
of the others. The other two species in the "very abundant" group
are Saprolegnia ferax (Gruith.) Thurent and Achlya sp., both of
the Saprolegniaceae. The same two famiUes dominate the "moderate-
ly abundant" group. There were two species of the Pythiaceae,
namely, Pythium sp. and Phytophthora s{)., while the Blastocladio-
ceae is represented by Blastocladia pringsheimii Reinsch and the
Gonapodyaceae is represented by Gonapodya polymorpha Thaxter.
Six species of Chytridiales were collected from the lake, all in the
"scarce" and "occasional" grou[)s. However, our choice of baits and
sampling techniques very likely selected against the detection of
many chytrid species. Six species were found on substrata not men-
tioned by Sparrow (1960). These aro: Mcgnchytrium wrstonii
Sparrow, parasitic on Ulothrix; Rhizophlyctis rosea (deB. and Wor.)
Fischer, on cellophane and Ordogonium sp.; Chytridium acuminatum
Braun, |)arasitic on pine j)ollen; Monohlrpharis insignis Thaxter,
sapro[)hytic on poplar twigs and apples; and Rhipidium americanum
and Rhipidium thaxteri. saprophytic on pine needles.

Acknowledgments

We gratefidly acknowledge the counsel and guidance given dur-
ing the course of this study by Dr. Earl M. Christensen, Def)artment
of Botany and Range Science, Brigham Young University, and the

Dec. 1972 rooney, mcknight: aquatic phycomycetes 189

critical reading of the manuscript by Dr. Christensen and Dr. R. A.
Paterson, Department of Biology, Virginia Polytechnic Institute.

Bibliography

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Intelligencer Press, North Carolina. 281 p.
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. 1961a. Distribution of aquatic Phycomycetes in some inorganic acido-

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Arch. f. Hydrobiol. & Planktonkunde 7:26; Figs. 1-24.

Welsh, P. S. 1952. Limnology. McGraw-Hill, Inc., New York. 381 p.

Weston, W. H. 1941. The role of aquatic fungi in hydrobiology. Univ. Wis-
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Willoughby, L. G. 1961. The ecology of some lower fungi at Esthwaite Water.
Trans, Brit. Mycol. Soc. 44:305-332.

. 1962. Ecology of some lower fungi in the English Lake District. Trans.

Brit. Mycol. Soc. 45:121-136.

NEW SYNONYMY IN AMERICAN BARK BEETLES
(SCOLYTIDAE: COLEOPTERAj, PART IP

Stephen L. Wood-

Abstract. — New synonymy involving American Scolytidae includes: Acan-
thotomicus Blandford (= Mimips Eggersj. Dendroterus Blandford (= Xylochilus
Schedlj, Chramesus dentatus Sciiaeffer ( = Ch. barbatus Eggers), Cnemonyx atra-
tus (Blandford) (= C. nitens Wood), C. errans (Blandford) (= Ceratolepsis
brasiliensis Schedl), C. exiguus (Blandford) (= Loganius pumilus Eggers), C.
minusculus (Blandford) (= Loganius uismiae Eggers), Cnesinus porcatus Bland-
ford (-= Cn. bicostatus Schedl j, Cryptocarenus serialus Eggers (= Cr. adustus
Eggers), Dendroterus luteolus (Schedl) (= D. mundus Wood), D. mexicanus
Blandford (= D. confinis Wood), D. sallaei Blandford (= Xylochilus insularis
Schedl), D. striatus (LeConte) (= Plesiophthorus californicus Schedl), Hylastes
gracilis LeConte (= H. longus LeConte). Hylocurus elegans Eichhoff (= Hy.
minor Wood), Hy. retusipennis Blandford (= Hy. bidentatus Schedl), Hy. rudis
(LeConte) (= Micracis biorbis Blackman). Xyleborus asper Eggers (= X. arnoe-
nus Schedl). X capucinus Eichhoff (= X. capucinoides Eggers), X. caraibicus
Eggers (= X. trinidadensis Schedl). X. declivis Eichhoff (= X. pseudoprocer
Schedl), X. deplanatus Eggers (= X. longideclivis Wood), X. discretus Eggers
(= X. usticus Wood), X. gilvipes Blandford (= X. mexicanus Eggers), X.
godmani Blandford (= A', caelebs Blandford). X. guatemalensis (Hopkins) f =
X. anisandrus Schedl), X. intrusus Blandford (= X. howardi Hopkins, X. scopu-
lorum Hopkins), X. lecontei (Hopkins) (= X. gundlachi Eggers), X. sparsipi-
losus Eggers (= X. inconveniens Schedl), X. spathipennis Eichhoff (= X. coro-
natus Eichhoff, Boroxylon burgdorfi Hopkins. X. curtus Eggers. X. femoraius
Eggeis), X. tumucensis Hagedorn (= X. guayanensis Eggers). X. vespatorius
Schedl (= X. corniculatus Schedl. X. corniculatulus Schedl), and X. volvulus
(Fabricius) (= X. grenadensis Hopkins, X. vagabundus Schedl). Microborus
bicolor Eggers is removed from synonymy, and the new name Acanthotomicus
bidentis is proposed for the preoccupied name Mimips bidens Wood.

While reviewing the Scolytidae of North and Central America
in the preparation of a taxonomic monograph of the family for this
area, the types of numerous species have been examined to confirm
the identity of the various taxa. This has resulted in the discovery of
several new synonyms. Since the monograph will not be published
for several years, the new synonymy is presented below in order
that names might be used in identifications and other work. The
genera and then the species treated are presented in alphabetical
order.

An item of special significance is noted here for the first time.
Two species described by Blandford (1895-1905. Biol. Centr. Amer.,
Coleopt. 4, part 6) from Mexican tobacco refuse intercepted at Paris,
(^nrmonyx atrntus (Blandford) and Hylocurus retusipennis Bland-
ford, have never been found in North or Central America, but both
are reported here as species native to southern Brazil. It is suggested
that all Blandford species described from Mexican tobacco refuse
actually came from Brazil, since none of them have ever been taken
from North or Central America.

''riiis work was supported l>v n iPsonK li (^riiiii fmni llu- National Siieiue I'luindation.
'DcpnilniCMl of /ooIorv. HriRhaiii Vmiiin rnnnsitv. I'lovo. Utah 84()0I.

190

Dec. 1972 wood, bark beetle synonymy 191

Acanthotomicus Blandford

Acanthotomicus Blandford, 1894, Trans. Ent. Soc. London 1894:89 (Type-species:

Acanthotomicus spinosus Blandford, monobasic).
Mimips Eggers, 1932, Rev. Zool. Bot. Afr. 22:33 (Type-species: Ips pilosus Eggers,

original designation). New synonymy.

In species assigned to the genera Acanthotomicus Blandford,
1894, and Mimips Eggers, 1932, the strongly flattened antennal club
varies from minutely pubescent and entirely devoid of sutures to
clearly marked, strongly procurved sutures in both groups. The
elytral declivity is broadly excavated, with the margins armed by
one to six pairs of denticles in species assigned to both groups. In
view of the diversity of the groups and the intergradation between
species assigned to each name, and in the absence of characters that
might support continued separation, I place Mimips in synonymy
under the much older name Acanthotomicus.

Schedl (1964, Reichenbachia 2:218) placed Acanthotomicus and
several other genera {Orthotomides Wood, Pityokteines Fuchs, and
Orthotomicus Ferrari) in synonymy under Ips. Although there is
some justification for his action, I do not feel it is in the interest of
taxonomy or of forestry to support his action. The characters em-
ployed and the justification for my restoration of these genera will
appear in another work.

Dendroterus Blandford

Dendroterus Blandford, 1904, Biol. Centr. Amer., Coleotp. 4(6):233 (Two species);
Hopkins, 1914, Proc. U.S. Nat. Mus. 48:120 (Type-species: Dendroterus
mexicanus Blandford, subsequent designation).

Xylochilus Schedl, 1956, Pan-Pacif. Ent. 32:31 (Type-species: Xylochilus insularis
Schedl = Dendroterus sallaei Blandford, original designation). New syn-
onymy.

The type-species of Xylochilus Schedl, X. insularis Schedl, is
identical to Dendroterus sallaei Blandford (see below), a species
clearly conspecific with D. mexicanus Blandford, the type-species of
Dendroterus Blandford. For this reason Xylochilus must be placed
in synonymy.

Acanthotomicus bidentis, n. n.

Mimips bidens Wood, 1971 (nee Schedl. 1967). Brigham Young Univ. Sci. Bull.,
Biol. Ser. 15(3) :41. Preoccupied.

When Mimips bidens Wood was published, the use of the same
name for an African species by Schedl (1967, Opusc. Zool. Budapest
7:229) was overlooked. In view of the above synonymy involving
the name Mimips and the homonymy, the new name Acanthotomi-
cus bidentis is proposed as a replacement name for Mimips bidens
Wood.

Chramesus dentatus Schaeffer

Chramesus dentatus Schaeffer, 1908, Jour. New York Ent. Soc. 16:221 (Lectotype,
female; Huachuca Mts., Arizona; U.S. Nat. Mus.).

192 GREAT BASIN NATURALIST Vol. 32, No. 4

Chramesus barbatus Eggers, 1931, Ent. Blatt. 26:169 (Holotype. male; Valle de
Mexico; Berlin Zool. Mus.)- ^^ew synonymy.

A pair of specimens bearing identical data to the lectotype of
dentatus Schaeffer and also compared to it, and the holotype of
barbatus Eggers were compared directly to one another. The males
are identical in every respect. The name barbatus. therefore, is here
placed in synonymy.

Cnemonyx atratus (Blandford)

Loganius atratus Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 129 (Lecto-
type, female; Bugaba, Chiriqui, Panama; British Mus. Nat. Hist., present
designation).

Cnemonyx nitens Wood, 1969, Brigham Young Univ. Sci. Bull., Biol. Ser. 10 (2): 9
(Holotype, male; Puerto Viejo, Heredia, Costa Rica; Wood Coll.). New
synonymy.

Blandford named Loganius atratus from three syntypes taken at
Bugaba, Panama. Of these three, two are females and the third
specimen is missing from its pin. Because the specimens of Cne-
monyx nitens Wood taken to the British Museum in 1964 were all
males and Blandford's atratus was represented only by females, the
synonymy was not detected until later when a lectotype was selected.
I here designate the first syntype from Bugaba, Panama, in Bland-
ford's series as the lectotype of Loganius atratus. This specimen was
labeled "Type" many years ago and since then has been regarded
as the type, although it has never officially been so designated.

Cnemonyx errans (Blandford)

Ceratolepsis errans Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 127 (Lecto-
type, male; intercepted at Paris in "Mexican" tobacco refuse; British Mus.
Nat. Hist., present designation).

Ceratolepsis brasiliensis Schedl. 1936. Archiv. Inst. Biol. Veg. Rio de Janeiro 3:104
(Syntypes; Rio Grande do Sul, Brazil; Schedl Coll. and Vienna Mus.). New
synonymy.

Blandford named Ceratolepsis errans from a syntypic series of
several specimens intercepted at Paris in tobacco refuse that sup-
posedly came from Mexico. Four of those syntypes are in the British
Museum (Natural History) where the first specimen, a male, was
labeled "Type" and has generally been regarded as the type, al-
though it has never officially been so designated. I here designate
that male syntype as the lectotype of Ceratolepsis errans Blandford.
This lectotype and two male paralectotypes were compared to three
males and two females of Ceratolepsis brasiliensis Schedl in my col-
lection identified by Schedl, labeled "Brasilian, Nova Teutonia, XII-
1940, F. Plaumann," and were found to be identical. It is noted
that in the original description of brasiliensis the sexes were reversed.

This sj)0(-ies has been reported from southern Brazil and northern
Argentina. There are no records or other reasons for believing it
occurs in Mexico as was indicated by Blandford.

Dec. 1972 wood: bark beetle synonymy 193

Cnemonyx exiguus (Blandford)

Loganius exiguus Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 130 (Lecto-
type, male; Bugaba, Chiriqui, Panama; British Mus. Nat. Hist., present
designation).

Loganius pumilus Eggers, 1929, Wiener Ent. Zeit. 46:65 (Holotype, male; Tur-
rialba, Costa Rica; U.S. Nat. Mus.). New synonymy.

Loganius exiguus Blandford was named from two male and one
female syntypes from Bugaba, Panama. The first male syntype is
here designated as the lectotype of Loganius exiguus Blandford. This
specimen was labeled "Type" many years ago, but has never of-
ficially been so designated. This lectotype and the male holotype of
Loganius pumilus Eggers were both compared to the same male
homotypes in my collection. Since all belong to the same species,
Eggers's name is here placed in synonymy under the senior name
exiguus Blandford.

Cnemonyx minusculus (Blandford)

Loganius minusculus Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 130

(Holotype, male; Volcan de Chiriqui, Chiriqui, Panama; British Mus. Nat.

Hist.).
Loganius vismiae Eggers. 1929, Wiener Ent. Zeit. 46:63 (Holotype, male; La

Caja, 8 km W San Jose, San Jose, Costa Rica; Deutschen Ent. Mus.). New

synonymy.

The male holotype of minusculus (Blandford) was compared
directly to a male cotype and a male topotype of vismiae (Eggers).
They are identical in all respects. An additional specimen from
San Juan, Alta Verapaz, Guatemala, was also examined.

Cnesinus porcatus Blandford

Cnesinus porcatus Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 137 (Six
syntypes; Cerro Zunil, Guatemala, and Volcan de Chiriqui, Panama; British
Mus. Nat. Hist.).

Cnesinus bicostatus Schedl. 1936, Arch. Inst. Biol. Veget. 3:106 (Holotype. male;
Turrialba, Cartago, Costa Rica; Schedl Coll.). New synonymy.

Following a year of collecting in Costa Rica, including several
days at various seasons at Turrialba, only two Cnesinus species
could be found that even remotely resemble bicostatus Schedl. These
were porcatus Blandford (2.8-3.1 mm) and costulatus Blandford
(2.0-2.3 mm) of which 43 and 32 specimens respectively were ex-
amined. Based entirely upon the original description and field ex-
perience in the area of its type locality, it was concluded that bi-
costatus Schedl (2.7 mm), known only from the unique male holo-
type which is not available for loan, must be a male of porcatus.
This proposed synonymy must be considered tentative until the type
of bicostatus is available for study.

Cryptocarenus seriatus Eggers

Cryptocarenus seriatus Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist.
Nat. 1(1): 10 (Holotype, female; Nouveau Chantier, Guyane Francaise; Paris

Mus.).

194 GREAT BASIN NATURALIST Vol. 32, No. 4

Crrptocarenus adustus Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. 1(1): 11
(Holotype, female; St. Jean du Maroni, Guyane Francaise; Pans Mus.).
New synonymy.

The female holotypes of seriatus Eggers and adustus Eggers were
examined and compared to more than 80 specimens from Florida to
Brazil. This material is easily associated with the type of seriatus,
which is 2.0 mm in length. The abraded condition of the type of
adustus, its shorter declivital setae on the elytra, and the concealed
frons led Eggers to regard it as a different species. The size was
reported as being 2.5 mm; however, if one compensates for the
slightly crushed condition of the type it actually is 2.3 mm in length,
well within the size range of seriatus. The removal of a small piece
of the mounting card exposed the previously concealed frons which
is of the typical .seriatus sculpture. The short declivital setae of the
type of adustus occurs commonly in specimens from Venezuela.
For these reasons the name adustus Eggers must be placed in syn-
onymy under seriatus Eggers because of page priority and the option
available to the first revisor.

Dendroterus luteolus (Schedl)

Plesiophthorus luteolus Schedl. 1951, Dusenia 2:111 (Holotype, male; Mexico;

Schedl Coll.).
Dendroterus mundus Wood, 1959, Great Basin Nat. 19:3 (Holotype. male; Tehuit-

zingo, Puebla, Mexico; Snow Ent. Mus.. Univ. Kansas). New synonymy.

The holotypes of both luteolus Schedl and mundus Wood were
examined. Prior to the description of mundus all identified speci-
mens (by Schedl) of luteolus known to me were from Baja California
and actually were of striatus (LeConte). However, the holotype of
luteolus has the evenly convex frons and coarser elytral vestiture of
mundus. Although the exact type locality of luteolus in Mexico is
unknown, the characters are sufficiently clear that the name mundus
should be placed in synonymy.

Dendroterus mexicanus Blandford

Dendroterus mexicanus Blandford, 1904. Biol. Centr. Amer., Coleopt. 4(6):233
(Holotype, female; Tehuantepec, Oaxaca, Mexico; British Mus. Nat. Hist.).

Dendroterus confinis Wood, 1959, Great Basin Nat. 19:6 (Holotype, male; Mag-
dalena, Jalisco, Mexico; Snow Ent. Mus., Univ. Kansas). New synonymy.

The larger average size of this species and the coarser sculpture
of the elytra and frons found in the northwestern part of its range
suggested the existence of a very different species from that de-
scribed by Blandford. However, the examination of 119 specimens
from six widely se[)arated localities in Mexico indicates that these
characters vary within a series and between series to such an extent
that only one species can be recognized. The holotypes of both mexi-
canus Blandford and confinis Wood were examined.

Dendroterus sallaei Blandford

Dendroterus sallaei Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):233 (Holo-
type. female; Veracruz, Veracruz. Mexico; British Mus. Nat. Hist.).

Dec. 1972 wood: bark beetle synonymy 195

Xylochilus insularis Schedl, 1956, Pan-Pacif. Ent. 31:31 (Holotype, male; Ar-
royo Hondo, Maria Madre. Tres Marias Islands, Gulf of California; Cali-
fornia Acad. Sci.). New synonymy.

The holotype of Dendroterus sallaei Blandford, the holotype and
several paratypes of Xylochilus insularis, and 82 other specimens
were examined. Only one species is represented by this material.
The specimen labeled "female holotype" in the Schedl collection has
status only as a paratype of his species.

Dendroterus striatus (LeConte)

Hypothenemus striatus LeConte, 1868, Trans. Amer. Ent. Soc. 2:156 (Syntypes;

Cape San Lucas, Baja California; Mus. Comp. Zool.).
Plesiophthorus californicus Sciiedl, 1952, Pan-Pacif. Ent. 23:123 (Holotype, female;

Angeles Bay, Gulf of California, Baja California; California Acad. Sci.).

Neiv synonymy.

The syntypic series of striatus LeConte, the holotype and several
paratypes of californicus Schedl, and 13 other specimens of this
species were examined. Only one species is represented. It is very
closely related to luteolus Schedl and eventually may be found to
represent only a geographical race of that species.

Hylastes gracilis LeConte

Hylastes gracilis LeConte, 1868. Trans. Amer. Ent. Soc. 2:174 (Two syntypes;

Tahoe Valley, California; Mus. Comp. Zool.).
Hylastes longus LeConte, 1876, Proc. Amer. Philos. Soc. 15:388 (Holotype,

female; Colorado; Mus. Comp. Zool.). New synonymy.

Both LeConte syntypes of gracilis and the holotype of longus
LeConte, and 126 other specimens of this species were examined.
Except for the brighter luster of the holotype of longus and of other
specimens from the southeastern part of the range, there is little
variation in this material. The name longus is here placed in syn-
onymy under the senior name gracilis LeConte.

Hylocurus elegans (Eichhoff)

Hylocurus elegans Eichhoff, 1871, Berliner Ent. Zeitschr. 15:134 (Holotype,
male; Teapa. Tabasco, Mexico; Inst. Roy. Sci. Nat.. Brussels).

Hylocurus minor Wood, 1961, Great Basin Nat. 21:4 (Holotype, female; Finca
Alto Bonito, Caicedonia. Valle de Cauca, Colombia; Wood Coll.). New
synonymy.

The holotypes of elegans Eichhoff and minor Wood and 230 other
specimens were examined. Because of the large size and the lack of
clarity in the original descriptions of elegans, this name could not
be associated with minor. An examination of the types, however,
leaves no doubt as to the synonymy.

Hylocurus retusipennis Blandford

Hylocurus retusipennis Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6):223
(Holotype, male; "Mexican" tobacco refuse; British Mus. Nat. Hist.).

196 GREAT BASIN NATURALIST Vol. 32, No. 4

Hylocurus bidentatus Schedl, 1950, Dusenia 1:149 (Syntypes; Nova Teutonia,
Santa Catarina. Brazil; Schedl and Plaumann Collections). Probable syn-
onymy.

The holotype of retusipennis Blandford that was presumed to
have come from Mexico, was compared to a series of bidentatus
Schedl, identified by Schedl, received from Plaumann from Santa
Catarina. The males are identical in every respect. Since the types
of bidentatus are not available for loan, the confirmation of the
suspected synonymy must be delayed.

Hylocurus rudis (LeConte)

Micracis rudis LeConte, 1876, Proc. Amer. Philos. Soc. 15:369 (Holotype, female?;

Detroit, Michigan; Mus. Comp. Zool.).
Micracis biorbis Blackman, 1920, Mississippi Agric. Expt. Sta. Tech. Bull. 9:22

(Holotype, male; Syracuse, New York; U.S. Nat. Mus.). New synonymy.

Due to an error in identification, Blackman named biorbis from
specimens that are identical to the holotype of rudis LeConte. Black-
man associated the name rudis with a southern species that is now
known as Hylocurus torosus Wood. The holotypes of both rudis
and biorbis were examined.

Microborus bicolor Eggers

Microborus bicolor Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat.
1(1): 19 Holotype. sex?; Bas Carsevenne, Guyane Francaise; Paris Mus.).

This species was placed in synonymy under aberrans Wichmann
by Schedl (1962, Mitt. Miinchn. Ent. Ges. 52:86). However, the
types of aberrans and setulosus Eggers were examined and were
found to represent different sexes of the same species; the type of
bicolor is larger and should be placed near ambitus Wood in a dif-
ferent species group from aberrans.

Xyleborus asper Eggers

Xyleborus asper Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat.

1(1):30 (Holotype, female; Nouveau Chantier, Guvane Francaise; Paris

Mus..)
Xyleborus amoenus Schedl, 1949. Rev. Brasil. Biol. 9:282 (Holotype, female;

Hamburgfarm on Rio Reventazon, Limon, Costa Rica; Schedl Coll.). New

synonymy.

The holotypes of both asper Eggers and amoenus Schedl were
examined and compared directly to my specimens from Costa Rica,
Panama, Colombia, Venezuela, and French Guiana. Only one easily
recognized species is represented by this material. The junior name,
amoenus, is hero placed in synonymy.

Xyleborus capucinus Eichhoff

Xyleborus capucinus Eichhoff, 1868 (1869), Berliner Ent. Zeitschr. 12:281 (Holo-
type, female; Guadeloupe Island; Inst. Roy. Sci. Nat.. Brussels).

Dec. 1972 wood: bark beetle synonymy 197

Xyleborus capucinoides Eggers, 1941, Arb. Morph. Taxon. Ent. Berlin-Dahlem
8:104 (Holotype, female; Gourbeyre, Guadeloupe Island; U.S. Nat. Mus.).
New synonymy.

The holotypes of both capucinus Eichhoff and capucinoides
Eggers and 102 other specimens were examined. Although the holo-
type of capucinus is callow and slightly crushed, it clearly is of the
same species as capucinoides. Eggers's name must be placed in
synonymy.

Xyleborus caraibicus Eggers

Xyleborus caraibicus Eggers, 1941, Arb. Morph. Taxon. Ent. Berlin-Dahlem

8:103 (Holotype, female; Guadeloupe; U.S. Nat. Mus.).
Xyleborus trinidadensis Schedl, 1961, Ann. Mag. Nat. Hist. (13)3:530 (Holotype,

female; River Estate. Trinidad; British Mus. Nat. Hist.). New synonymy.

The holotypes of caraibicus Eggers and trinidadensis Schedl and
24 other specimens were examined and compared to my material.
Only one species is represented by this material. The junior name
trinidadensis is here placed in synonymy.

Xyleborus declivis Eichhoff

Xyleborus declivis Eichhoff, 1868 (1869), Berliner Ent. Zeitschr. 12:280 (Holo-
type, female; Teapa, Tabasco, Mexico; presumably lost with Hamburg Mus.).

Xyleborus pseudoprocer Schedl. 1949, Rev. Brasil. Biol. 9:279 (Holotype, female;
Guatemala; Schedl Coll.). New synonymy.

My three specimens of this species from Costa Rica and Mexico
were compared to Blandford's series from Ciuatemala and to the
holotype of p.seudoprocer Schedl; all clearly represent the same spe-
cies. This material completely fits the description of declivis Eich-
hoff, the type of which evidently is lost. Since it is the only species
of this size (4.0-4.4 mm) from Mexico and Central America in this
species group, since it fits the original description, and because it
agrees with material identified by specialists who studied the Eich-
hoff collection, it is reasonable to assume that this species is declivis
Eichhoff. If this is correct, the name pseudoprocer Schedl must be
treated as a junior synonym.

Xyleborus deplanatus Eggers

Xyleborus discretus Eggers. 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist.

Nat., Paris 1(1): 32 (Holotype. female; Charvein, Guvane Francaise; Paris

Mus.).
Xyleborus longideclivis Wood, 1968, Great Basin Nat. 28:1 (Holotype, female;

Bartica Triangle, British Guiana; British Mus. Nat. Hist.). New synonymy.

The female holotype of deplanatus Eggers was compared directly
to four para types of longideclivis Wood. This species is 2.0-2.3 mm
in length, not 3.0 mm as stated in Eggers's description. In addition
to British and French Guiana it also occurs in Colombia.

Xyleborus discretus Eggers

Xyleborus discretus Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d' Hist.
Nat.. Paris 1(1):29 (Holotype, female; Marcapata, Peru; U.S. Nat. Mus.).

198 GREAT BASIN NATIIRALIST Vol. 32, No. 4

Xyleborus usticus Wood, 1968, Great Basin Nat. 28:3 (Holotype. female; Bartica
District, British Guiana; British Mus. Nat. Hist.). New synonymy.

When usticus Wood was named, the distribution of discretus
Eggers in French Guiana was overlooked. When the holotype of
discretus was compared to a topotypic paratype of usticus, the syn-
onymy was immediately apparent. This species is now known from
Costa Rica, Venezuela, British and French Guiana, and Peru. As
indicated above, the name discretus has priority.

Xyleborus gilvipes Blandford

Xyleborus gilvipes Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6):205 (Holo-
type, female; Zapote, Guatemala; British Mus. Nat. Hist.).

Xyleborus mexicanus Eggers, 1931, Ent. Blatt. 27:19 (Holotype, female; Mara-
vatio, Michoacan. Mexico; Zool. Mus. Berlin). New synonymy.

The holotypes of gilvipes Blandford and mexicanus Eggers were
both examined and compared directly to my specimens. They clear-
ly represent the same species. The junior name mexicanus is here
placed in synonymy.

Xyleborus godmani Blandford

Xyleborus godmani Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6): 197 (Holo-
type, female; Bugaba, Chiriqui. Panama; British Mus. Nat. Hist.).

Xyleborus caelebs Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6): 198 (Holo-
type, male; Volcan de Chiriqui, Chiriqui, Panama; British Mus. Nat. Hist.).
New synonymy.

The holotypes of both godmani Blandford and caelebs Blandford
were compared to definitely associated males and females from Pana-
ma and Costa Rica. Although the two sexes are very different
anatomically, it is entirely clear that only one species is represented
by the two names. The name caelebs is here placed in synonymy
because of page priority and by choice of the first revisor.

Xyleborus guatemalensis (Hopkins)

Ambrosiodmus guatemalensis Hopkins, 1915. U.S. Dept. Agric. Kept. 99:56
(Holotype, female; Trece Aguas, Alta Verapaz, Guatemala; U.S. Nat. Mus.).

Xyleborus anisandrus Schedl, 1954, Dusenia 5:44 (Syntypes. females; Rio Claro.
Brazil; Schodl Coll.). New synonymy.

The holotype of guatemalensis Hopkins and the syntype of ani-
sandrus Schodl in the Schedl collection were both compared to my
material from Costa Rica, Panama, (>)lombia, Venezuela, and Brazil.
I am unable to see even minor differences in the 32 specimens ex-
amined. The junior name ani.sandrus is here placed in synonymy.

Xyleborus intrusus Blandford

Xyleborus intrusus Blandford, 1898. Biol. Centr. Amer., Coleopt. 4(6):213 (Syn-
types, females; San Geronimo, Guatemala; British Mus. Nat. Hist.).

Xyleborus howardi Hopkins, 1915, U.S. Dept. Agric. Kept. 99:65 (Holotype.
female; Washington, D.C.; U.S. Nat. Mus.). New synonymy.

I

Dec. 1972 wood: bark beetle synonymy 199

Xyleborus scopulorum Hopkins, 1915, U.S. Dept. Agric. Rept. 99:66 (Holotype,
female; Black Hills, South Dakota; U.S. Nat. Mus.). New synonymy.

The three syntypes of intrusus Blandford, the holotypes of how-
ardi Hopkins and scopulorum Hopkins, and 42 other specimens were
compared to my material. The material from the western United
States, Mexico, and Guatemala range from 2.3 to 2.7 mm in length
and clearly represent one species. Specimens from the eastern
United States average slightly smaller in size, ranging from 2.2 to
2.5 mm in length. It is also noted that pronotal and elytral char-
acters mentioned by Bright (1968, Canadian Ent. 100:1320) are
not consistent and occur in both populations. In the absence of dis-
tinguishing characters or other means of separating these popula-
tions, I here place howardi and scopulorum in synonymy as indicated
above.

Xyleborus lecontei (Hopkins)

Ambrosiodmus lecontei Hopkins, 1915, U.S. Dept. Agric. Rept. 99:56 (Holotype.

female; Keene, Florida; U.S. Nat. Mus.).
Xyleborus gundlachi Eggers, 1931, Ent. Blatt. 27:20 (Holotype, female; Cuba;

Zool. Mus. Berlin) . New synonymy.

The holotypes of both lecontei Hopkins and gundlachi Eggers
were examined and compared directly to my specimens. They are
identical in all respects. The junior name gundlachi is here placed
in synonymy.

Xyleborus sparsipilosus Eggers

Xyleborus sparsipilosus Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist.
Nat., Paris 1 ( 1 ) : ^4 (Holotype. female; Nouveau Chantier. Guyane Francgise;
Paris Mus.).

Xyleborus inconveniens Schedl, 1948, Rev. de Ent., Rio de Janeiro 19:577 (Holo-
type, female; Hamburgfarm on Rio Reventazon, Limon, Costa Rica; Schedl
Coll.). New synonymy.

The female holotypes of sparsipilosus Eggers and inconveniens
Schedl were both compared to several of my females from Costa
Rica. All represent the same species in all details. The name incon-
veniens must be placed in synonymy under the senior name spar-
sipilosus.

Xyleborus spathipennis Eichhoff

Xyleborus spathipennis Eichhoff, 1868, Berliner Ent. Zeitschr. 12:145 (Syntypes,
female; Peru; Inst. Roy. Sci. Nat., Brussels).

Xyleborus coronatus Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:348 (Holo-
type, male; Brasilia interior; Inst. Roy. Sci. Nat., Brussels). New synonymy.

Xyleborus burgdorfi Hopkins, 1915, U.S. Dept. Agric. Rept. 99:59 (Holotype,
female; Costa Rica; U.S. Nat. Mus.). New synonymy.

Xyleborus curtus Eggers, 1928, Arch. Inst. Biol. Sao Paulo 1:94 (Lectotype,
female; Cachabe, Equador; U.S. Nat. Mus. J. New synonymy.

Xyleborus femoratus Eggers, 1928, Arch. Inst. Biol. Sao Paulo 1:95 (Syntypes,
females; Bahia, Brazil; Zool. Mus. Berlin).

The holotypes of coronatus Eichhoff and burgdorfi Hopkins, the

200 GREAT BASIN NATURALIST Vol. 32, No. 4

lectotype of curtus Eggers, the Chapuis syntypes of spathipennu
Eichhoff, three syntypes of femoratus Eggers, and 74 other speci-
mens were examined and either compared directly to one another or
to specimens in my collection including several series of definitely
associated males and females. Only one species is represented;
coronatus, burgdorfi, curtus, and femoratus are placed in synonymy
as indicated above.

Xyleborus tumucensis Hagedorn

Xyleborus tumucensis Hagedorn, 1905, Bull. Mus. d'Hist. Nat., Paris 6:414

(Three female syntypes; Riviere Lunier. Tumuc-Humac, Guyane Francaise;

Paris Mus.).
Xyleborus guayanensis Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist.

Nat., Paris 1(1):28 (Male and female syntypes; Nouveau Cliantier, Guyane

Francaise; Paris Mus.). New synonymy.

The female syntype of tumucensis Hagedorn that has been
labeled "type," the male syntype and two female cotypes (the female
syntype is missing from the Paris Museum ) of guayanensis Eggers,
and 43 other specimens of this sj)ecies were examined and compared
directly to one another. Only one species is represented. The error
evidently occurred when Eggers (1933) misidentified specimens of
geayi Hagedorn which he reported as tumucensis. The name guay-
anensis Eggers is placed in synonymy under the older name tu-
mucensis.

Xyleborus respatorius Schedl

Xyleborus vespatorius Schedl. 1931, Ann. Mag. Nat. Hist. (10)8:339 (Holotype.

female; San Ignacio, Argentina; Schedl Coll.).
Xyleborus corniculatus Schedl, 1949, Rev. Brasil Biol. 9:275 (Holotype, female;

Santa Catarina, Brazil; Schedl Coll.). New synonymy.
Xyleborus corniculatulus Schedl, 1949, Rev. Brasil Biol. 9:275 (Holotype, female;

Trinidad; Schedl Coll.). New synonymy.

The holotypes of vespatorius Schedl, corniculatus Schedl, and
corniculatulus Schedl were examined and compared directly to one
another and to my homotyi)es. The three names are based on what
I consider to be minor variations of the same species. The declivital
denticles of the holotype of vespatorius have been damaged, pre-
sumably by the chewing of siblings, thereby making recognition
more difficult. The name vespatorius has priority over both cornicu-
latus and corniculatulus.

Xyleborus volvulus (Fabricius)

Bostrichus volvulus Fabricius, 1775, Systema Entomologiae, p. 454 (Syntypes,

females; America ligno Dom v. Rohr. presumably Cuba; Copenhagen Mus.).
Xyleborus lorquatus Ei( hhoff. 18()8. Bcilinor Ent. Zeitschr. 12:146 (Syntypes.

female; Cuba, Brazil, Puerto Rico; presumably lost with Hamburg Mus.).
Xyleborus grenadensis Hopkins, 1915, U.S. Dept. Agric. Rept. 99:65 (Holotype,

female; Grenada, West Indies; U.S. Nat. Mus.). New synonymy.
Xyleborus vagabundus Schedl, 1949, Rev. Brasil. Biol. 9:277 (Holotype, female;

Mexico; Schedl Coll.). New synonymy.

Dec. 1972 wood: bark beetle synonymy 201

Three syntypes of volvulus Fabricius, the holotypes of grena-
densis Hopkins and vagabundus Schedl, and several hundred speci-
mens from Florida and Baja California to Argentina, Africa, Hawaii,
Micronesia, and Australia were examined. Apparently two distinct
geographical races of this species existed prior to the advent of mod-
ern commerce: a northern one (volvulus), from central Mexico
northward including Cuba and Florida; and a southern race (tor-
quatus), from Central and South America. Over the past century,
one or both races of this species were introduced to other areas where
one race or the other predominated or else hybridized to form local
populations either intermediate in anatomical details or highly vari-
able in structure with both extremes and all degrees of intergrada-
tion represented. The southern race apparently has been introduced
repeatedly into the range of the northern race and has maintained
itself locally to some degree. Because of the extreme hybridization
which occurred in areas outside of the American continents and the
mixing taking place in Mexico, I see no possibility of the continued
recognition of definite geographical races.

The types of grenadensis Hopkins and vagabundus Schedl both
represent normal minor variations of the northern race and must be
placed in synonymy.

FLKrHTS OF THE WESTERN THATCHING ANl,

FORMICA OBSCURIPES FOREL, IN NEVADA

( H YMENOPTER A : FORMICIDAE ) •

William H. Clark- and Peter L. Comanor-'

Abstract.— Obsei-\'ations indicate that Formica obscuripes flies earlier
in Nevada than in other parts of the country where the flights have been re-
corded. The alate males are the first to emerge from the nest. These are eventual-
ly succeeded by alate females, and finally only females are seen. F. obscuripes
flew during periods when air temperatures were 20.5-26.5 C, when relative hu-
midities were about 18%, and when solar input was about 1.35 langleys. Slight
winds approaching 3 mph did not keep the ants from flying but may have in-
fluenced their direction of flight.

The nests of F. obscuripes observed near Reno differ from those reported in
other areas in that they do not have such an extensive thatch covering them.

Flight activities have been reported for Formica obscuripes by
McCook (1884). He states that the marriage flight of this ant in
"Dakota" occurs in "the latter part of July, [when] numerous
swarms of 'flying ants' are seen" and that "the appearance of the
swarms continuing throughout August and into September" has
been observed. Cole (1932) states that "winged males and females
appear in large numbers during June and July." In North Dakota,
flights occur in the month of June (Weber, 1935), and in Michigan,
in June and early July (Talbot, 1959; 1972). It has been suggested
that F. obscuripes does not have true nuptial flights and that winged
forms leave the nest one or a few at a time (Weber, 1935). On the
other hand, more recent observation suggests that F. obscuripes has
conspicuous swarming and mating activities (Talbot, 1959; 1972).
This seems to be substantiated by the polygymous habit of this ant
(King and Sallee, 1956). Wheeler and Wheeler (1963) report
winged females of F. obscuripes in nests from 6 June to 8 August
and winged males from 23 May to 12 July in North Dakota.

Records of flight activities of this ant in the Great Basin have not
been found in the literature. The Nevada flights reported below were
first noted on 15 April and lasted until at least 10 May 1972, thus
occurring almost two months earlier than previously recorded for
this species.

F. obscuripes has been recorded in Nevada, but not previously in
Washoe County (La Rivers, 1968).

Materials and Methods

Three nests of F. obscuripes were periodically observed from 15
April to 10 May 1972 in conjunction with other ant studies in the
general areas of the study sites described below.

'Thi'; work was assisted 1)V a uranl from the Research Advisory Board. University of Nevada.
Ken-

-Dopartnictil of nlol^^;v. I'ruversitv of Nevada. Reno, Nevada 81507.

202

Dec. 1972 clark, comanor: ant flight 203

Numbers of winged males and females on the nests, in nearby
vegetation, and actually flying were noted. Temperature data were
taken in C by two methods. At nest number 1, the temperatures
were taken with a YSI 12 channel telethermometer, and tempera-
tures at nests number 2 and 3 were taken with two standard labor-
atory glass thermometers. Ground surface temperatures and air
temperatures at 10 cm above the ground surface were taken. Rela-
tive humidity was taken with a sling psychrometer operated as near
the ground surface as possible, and wind measurements were made
with a hand-held anemometer at 1 meter above ground surface. On
1 May, solar input, in langleys per minute (cal/cmVmin), was
taken with a weathemieasure recording pyranograph.

The three F. obscuripes nests observed during April and May
1972, all located just north of Reno, Washoe County, Nevada, occur
at elevations of about 1550 m and are separated by distances of up
to 15 miles. The vegetation of these areas is dominated by Artemesia
tridentata, basin big sagebrush. Numbers of subordinate species vary
among the sites. The chief ones are Chrysothamnus viscidiflorus, C.
nauseosus. Ephedra viridis. Ribes velutinum, Tetradymia canescens
and Bromus tectorum. Nest number 1 is 1.1 m in diameter and 30
cm high. It is constructed around a living Chrysothamnus nauseosus
shrub, and its thatch is relatively thin and composed mainly of
Ephedra viridis stems, Artemesia tridentata bark, and grass overly-
ing a base of coarse, decomposed granite sand. Nest number 2 is 79
cm in diameter and 25 cm high. This nest is built around living and
dead Artemesia tridentata and Tetradymia canescens. with its thatch
constructed mainly of sage bark and grass. Nest number 3 is 50 cm
in diameter and 1 5 cm tall and is in living and dead Artemesia tri-
dentata and Chrysothamnus viscidiflorus. Its thatch is composed
mostly of Ephedra viridis stems, sage bark, and grass.

These three nests differ from the nests described by Cole (1932),
Weber (1935), and Wheeler and Wheeler (1963) in the extent of
the thatching. These workers state that the thatch material covers
the mound and extends into the soil to varying depths, to a maxi-
mum of about 30 cm. The three nests observed in this study have a
thin layer of thatch on top, with the lower sides composed mostly of
bare gravel and occasional sticks or grass stems. None of the nests
observed has a thatch that extends to or into the ground.

Results

The first winged individuals of F. obscuripes observed were males
noted on the morning of 15 April. About a dozen winged males were
observed in Chrysothamnus nauseosus near nest number 1. After
these winged ants were noticed, two other F. obscuripes nests were
located and occasionally observed. On 18 April, seven males were col-
lected from the surface of nest number 2 in the time period from
10:00 to 10:30 a.m. None were observed in surrounding plants.
Nest number 3 was located on 19 April, but no winged individuals
were found; however, two winged males were seen on nest number

204 GREAT BASIN NATURALIST Vol. 32, No. 4

2. On 29 April at 2:45 p.m., winged males were again observed on
nest number 2.

On 1 May 1972, nest number 1 was observed for a day (see
Table 1 ) , and a maximum of fourteen females flew per minute at
10:25 a.m., representing the height of the flight, when the air
temperature at 10 cm above ground surface was 25 C (see Figure 1 ).

Discussion

According to Weber (1935), when "the air is calm, the sky quite
clear, the temperature above 60 F (15 C), and the humidity above
50%, the ants take flight." These temperatures were taken at 10
inches (25.4 cm) above ground level (Talbot, 1971). Talbot (1959)
found that F. obscuriprs "began flying at about 70 F (21 C), pro-
vided they were not wet, the grass was not swaying, and the sky was
not darkening." The correspondence of flights with rising tempera-
tures in the morning is suj)ported by work on several species of For-
mica elsewhere (Talbot, 1964). The flights observed in Nevada
all occurred within the temperature range given by Talbot (1959;
1972).

During the flights recorded for Nevada, a breeze was always pres-
ent, the sky was clear, and the temperature was always above

.>^^.,,, ,

i* i

5r

Fig. 1. Alate Formica obscuripes females on Chrysothamnus nauseosus
1 May 1972.

Dec. 1972

CLARK, COMANOR: ANT FLIGHT

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206 GREAT BASIN NATURALIST Vol. 32, No. 4

15 C. The humidity during these flights was always below 50%,
with a relative humidity of about 18% during flights which occurred
on 1 May 1972. The wind was not strong, usually averaging 2-3
mph, and did not seem to inhibit the flight of the ants. As they were
observed to fly from the Chrysothamnus stems which were moving
in the wind, the wind did seem to influence the direction of flight.
On 1 May, the wind was from the east and the ants were flying
westward and northwestward. The alate ants flew upward for one
to about five meters, occasionally straight up, but usually at an
angle to the west. Several times females would begin flight in an
easterly or southerly direction and after attaining an altitude of
several meters would shift to a westerly direction. This change in
flight direction could be a result of the light winds present. The
same type of wind conditions prevailed on 4 May, and again the
ants flew in a westerly direction. Three winged females appeared on
the nest surface for a short time at 5:20 p.m., then returned into the
nest. Winged ants were seen outside the nest when the correspond-
ing solar input ranged from 0.98 to 1.52 langleys and were observed
to fly when the solar input ranged from 1.33 to 1.435 langleys.

On 4 May at 10:00 a.m., nest number 1 had 10 females on the
surface and 45 were located in nearby shrubs; three males were ob-
served in shrubs. Sixteen females and one male flew in a one-minute
time period, when the air temperature was 20.5 C. At 10:30 a.m.
nest number 3 had one winged female on the nest surface. On 10
May at 8:45 a.m., nest number 2 had seven winged females on the
surface and one in a sagebrush. At 9:45 a.m., nest number 3 had
10 winged females on the nest and 20 in sagebrush.

An additional F. ohscuripes nest on Austin Summit, Lander
County, Nevada, elevation 2286 m, was found to have no winged
individuals present outside the nest during the morning of 12 May
1972.

Winged males of F. ohscuripes were much fewer than females.
Males were also the first winged forms observed to leave the nest in
April; gradually more females were present, until finally in May
only females were flying. These observations agree with those of
Talbot (1959) for this species.

Our observations indicate that F. ohscuripes flies earlier in Ne-,
vada than elsewhere where it has been observed. Sudd (1967) states
that "ants of deserts and steppes do not fly in late summer, when
the quec^ns would be exposed to hot, dry weather in the time when
they are founding colonies. The flight is delayed till spring." The
earlier flights in Nevada could be an avoidance of the summertime
hot, dry weather; or, it may be that the ants simply develop sooner
in this climate.

Acknowledgements

The authors thank Drs. George C. and Jeanette Wheeler for the
species determination of F. ohscuripes.

Dec. 1972 clark, comanor: ant flight 207

Literature Cited

Cole, A. C, Jr. 1932. The thatching ant, Formica obscuripes Forel. Psyche

39:30-33.
King, R. L.. and R. M. Sallee. 1956. On the half-life of nests of Formica

obscuripes Forel. Proc. Iowa Acad. Sci. 63:721-723.
La Rivers, I. 1968. A first listing of the ants of Nevada ( Hymenoptera. Formi-

cidae). Occ. Pap. Biol. Soc. Nevada 17:1-12.
McCooK, H. C. 1884. The rufous of thatching ant of Dakota and Colorado.

Proc. Acad. Natur. Sci. Phila. Part 1, pp. 57-65.
Sudd, J. H. 1967. An introduction to the behavior of ants. Arnold, London.

200 p.
Talbot, M. 1959. Flight activities of two species of ants of the genus Formica.

Amer. Midi. Natur. 61:124-132.
. 1964. Nest structure and flights of the ant Formica obscuriuentris

Mayr. Animal Behaviour 12:154-158.

. 1971. Flights of the ant Formica dakotensis Emery. Psyche 78:169-179.

. 1972. Flights and swarms of the ant Formica obscuripes Forel. Jour.

Kansas Entomol. Soc. 45:254-258.
Weber, N. A. 1935. The biology of the thatching ant Formica obscuripes Forel

in North Dakota. Ecol. Monogr. 5:165-206.
Wheeler, G. C, and J. Wheeler. 1963. The ants of North Dakota. Univ.

North Dakota Press. Grand Forks. 326 p.

A NEW SPECIES OF SOIL MITE FROM WESTERN COLORADO
(ACARI: CRYPTOSTIGMATA, PELOPIDAE)i

Harold G. Higgins- and Tyler A. Woolley''

Abstract. — A new species of soil mite from Hayden, Colorado, is described.
This species, Peloptulus tanyirichosus, is the first representative of the genus in
North America. The species is described, figured, and compared with P. foveola-
tus Hammer, 1961. Eight specimens were taken from different habitats, including
aspen litter, scrub oak litter, and beneath greasewood and pepper grass.

During the summer of 1971, a number of soil samples were taken
in the vicinity of a coal-burning power plant near Hayden, Colora-
do. During these investigations several new species of soil mites were
taken from different ecological niches in the project area. Among
others is a new species in the genus Peloptulus. This genus is known
from Europe and Peru, South America, but to our knowledge has
never before been recorded from North America.

Peloptulus tanytrichosus, n. sp.

(Figs. 1, 2, and 5)

Diagnosis. — Sensillum much longer than any known species,
extending well beyond level of translamella; lamellar tip without
cusps.

Description.- — Color deep reddish brown; prodorsum about as
long as broad; rostrum squarish, lateral margins flaring somewhat
anteriorly; lamellae long, reaching nearly to tip of rostrum, covering
much of the prodorsum, without cusps or notched tip; translamella
narrow, longer than broad, closer to rostrum than notogaster; lamel-
lar hairs heavy, setose, attached on inner margin of lamellae near
their tips, and flaring outward; interlamellar hairs weak, smooth,
about as long as distance between lamellae at their insertion, in-
serted below anterior border of hysterosoma, in the arch between the
pseudostigmata, extending forward slightly beyond the sclerotized
line connecting the pteromorphs; sensillum long, in some specimens
more than three-fourths as long as lamellae, with swollen, setose
head and long, narrow pedicel; pseudostigmata oval, with heavy,
sclerotized lips rising above surface of prodorsum; pedotectum I with
a long, slender, pointed ti[) that can be seen from the dorsal side.

Hysterosoma rounded in outline with dorsal setae and markings
as shown in Figure 1 ; the two posterior setae are slightly larger,
with more squarish tips than the rest; specimens show some varia-
tion in size and length of dorsal setae.

Camerostome flask-shaped; genital opening squarish, about as
long as wide and shorter than anal opening, each genital plate with

■Research supported by Western Colorado Hesourre Study.
'Department of Biology, Granger High School, Granger." Utah
'Deparlmeni nf Zoology. Colorado Stale University, Fort Collins. Colorado

208

Dec. 1972

HIGGINS, WOOLLEY: A NEW MITE

209

Figs. 1-3. Peloptulus tanytrichosus,
genital area; 3. anal area.

n. sp.: 1, dorsal aspect, legs omitted; 2,

six setae as shown in Figure 2; anal plates somewhat egg-shaped,
inner edge of anal plates wavy; distance between genital and anal
opening equal to about length of anal opening as shown in Figure 3.

Legs heterotridactylous, the middle claw larger than laterals.

Size. — Length, .45 mm; width, .34 mm.

Collection Data. — The type, a male, and on other male speci-
men were collected from litter under aspens, 4 miles S Seneca Road,
Hayden, Colorado, 1-VIIL1971, by H. G. Higgins; 1 specimen from

210 GREAT BASIN NATURALIST Vol. 32, No. 4

under aspens, 4 miles S Seneca Road, Hayden, 8-VI-1971, by H. G.
Higgins and T. A. Woolley; 3 specimens from imder scrub oaks, 4
miles S Seneca Road, l-Vni-1971, by H. G. Higgins; 1 specimen
from under scrub oaks near spoil bank, Wolf Creek North, Hayden,
9-VI-1971, by H. G. Higgins and T. A. Woolley; 1 specimen from
under greasewoods and pepper grass, 4 miles S Hayden, 9-VI-1971,
by T. A. Woolley and H. G. Higgins.

Discission. — The most striking fhfference between this species
and other known species of the genus Peloptulus is the long sensil-
lum. This species differs from the South American species P. foveo-
latus Hammer by lacking the reticulations on the body, a cusped
lamellae, and having a much longer sensillum. The sensillum of P.
foveolatus reaches nearly to the translamella. The trivial name from
the Greek tanytrichosus means "long hair" and refers to the long
sensillum.

This mite appears to live in a number of habitats, since it has
been found in litter under quaking aspen, in litter under scrub oak,
and in rather dry soil under greasewoods and pepper grass. In the
small collection under study it appears that this species shows indi-
vidual variation in the length of sensillum, but the relative length as
it relates to the translamella is a diagnostic feature of importance. It
appears that the sensillum of males is slightly longer than the sensil-
lum of females. In addition, the species from the dryer scrub oaks
and greasewoods seem to have a narrower translamella than those
from the more moist aspen litter.

LiTERATURF. CiTED

Balogh, J. 1963. Identification keys of holarctic oribatid mites (Acari) fami-
lies and genera. Acta Zool. 9(1-2): 1-60.

H.\MMER, M. 1961. Investigations of the oribatid fauna of the Andes Mts., II
Peru. Biol. Skr. Dan. Vid. Selsk. 13(1): 1-57.

WiLLMAN. C. 1931. Moosmilben oder Oribatiden. Tierwelt Deutschlands 22:79-
200.

A CHECKLIST OF THE MACROINVERTEBRATES
OF THE PROVO RIVER, UTAH

Parley V. Winger,^ Edward J. Peters,' Michael J. Donahoo,'
James R. Barnes, ' and David A. White'

Abstract. — A checklist of the aquatic macroinvertebrate species from
the Provo River, Utah, was compiled from field collections and from literature
sources.

Recent water control activities and proposed developments along
the Provo River in Provo Canyon have served to stimidate research
on aquatic life in this river. Many studies of aquatic life deal either
directly or indirectly with the aquatic insects in the stream. Pre-
vious studies of aquatic insects in the Provo River are either unpub-
lished theses or incidental references in papers of a broader scope
(Edmunds, 1952). The most complete studies were conducted by
Gaufin (1949, 1951, 1959).

Students at the University of Utah studied the Trichoptera
(Merkley, 1948), aquatic Coleoptera (Todd, 1952), Chironomidae
(Brooks, 1955), and Plecoptera (Sessions, 1960) of the Provo River.

Incidental references to the species of aquatic insects were made
by Needham and Christensen (1927), Hazzard (1934), Moffet
(1936), Chandler (1941), Tanner (1941), Anonymous (1948),
Dunstan (1951), Anderson (1960), Musser (1961), and Braithwaite
(1962).

The present study was conducted during the winter of 1969-70.
Collections were taken from 10 stations located along the Provo River
from 51/2 miles above Woodland, Utah, downstream to the U. S. 91
highway bridge in Provo, Utah.

The main purpose of this paper is to make available, in one place,
a complete list of the aquatic invertebrates which have been collected
from the Provo River. Table 1 lists all organisms collected during
the present study and from previous pertinent studies (INIerkley,
1948; Gaufin, 1951; Todd, 1952; Gaumer, 1952; Brooks, 1955; Ses-
sions, 1960; Gaufin, Nebeker, and Sessions, 1966).

The collection of Sialis sp. (Sialidae) has not, as far as we know,
been previously reported in the literature for northern Utah. In this
study, immatures of Sialis were collected on several occasions in the
Provo River downstream from Woodland, Wasatch Co., Utah. One
specimen of Sialis has been collected from Hobble Creek, east of
Springville, Utah Co., Utah. Dr. Arden R. Gaufin, University of
Utah (Pers. Comm., 1972), has collected Sialis from the Weber,
Provo, and Bear rivers.

Acknowledgments. — We would like to thank Brigham Young
University for allowing us the use of equipment and facilities used

^Department nf Zoology. Brigham Young University. Provo. Utali a4()()l.

211

212 GREAT BASIN NATURALIST Vol. 32, No. 4

in this study. Thanks are also due to Dr. Roy T. Sawyer, College of
Charleston, Charleston, South Carolina, who identified the Hiru-
dinea.

Table 1. Checklist of the Macroinvertebrates of the Provo River, Utah.
(Parentheses around number indicates specimens not classified to species. 1 -=
present study; 2 = Gaufin, 1951; 3 = Merkley, 1948; 4 = Sessions, 1960; 5 =
Todd, 1952; 6 = Brooks. 1955; 7 = Gaufin, Nebeker. and Sessions, 1966; 8 =
Gaumer, 1952; 9 = Edmunds, 1952.)

ORGANISM COLLECTOR

Collembola 2

Coleoptera

Caraboidea
Amphizoidae

Amphizoa lecontei Matthews 5

Dytiscidae

Bidessus af finis (Say) 5

Hydropnrus planiusculus Fall (2), 5

Hydroporus pervicinus Fall 5

Hydroporus occidentalis Sharp 5
Hydroporus transpunctatus Chandler

Hygrotus masculinus (Crotch) 5

Hygrotus patruelis (LeConte) 5

Hygrotus tumidiventris (Fall) 5

Laccophilus decipiens LeConte 5

Laccophilus astristernalis Ciotch 5

Oreodytes crassulus (Fall) 5

Oreodytes congruus (LeConte) 5

Agabus cordatus (LeConte) 5

Agabus hypomelas Mannerheini 5

Agabus seriatus intcrsectus Leech 5

Agabus oblong us Fall 5

Agabus approximatus Fall 5

Agabus austini Sharp 5

Agabus strigulosis (Crotch) 5

Agabus kcnaiensis Fall 5

Agabus rrichsoni Gemmingor & Harold 5

Agabus tristis Aube 5

Ilybius subaenus Erickson 5

Ilybius fralerculus LeConte 5

Rantus divisus (Aube) 5

Rantus longipes Sharp 5

Rantus hoppingi Wallis 5

Dytiscus marginirollis LeConte 5

Dytiscus dauricus Gehl. (1). (2), 5

Dytiscus circumsinctus Ahrens 5

Acilius semisulcatus Aube 5

Cybister explanatus LeConte 5

Haliplidae

Brychius horni Crotch 5

Haliplus imniaculicollis Harris (1), (2). 5

fialiplus Irrchi Wallis 5

Peltodytes callosus (LeConte) 5

Gyrinoidea
Gyrinidae

Gyrinus bifarius VaU 5

Gyrinus consobrinus LeConte 5

Dec. 1972

WINGER, ET AL: CHECKLIST

213

Table 1. (Continued)

Gyrinus pleuralis Fall
Gyrinus picipes Aube
Gyrinus affinis Aube

Hydrophiloidea
Hydrophilidae

Hflophorus oblongus LeConte
Helophorus obscurus LeConte
Telophorus nitidulus LeConte
Berosus infuscatus LeConte
Berosus stylijerous Horn
Hydrous triangularis (Say)
Tropisternus ellipticus (LeConte)
Tropisternus dorsalis (Brulle)
Tropisternus calif or nicus (LeConte)
Hydrobius fuscipes Linnaeus
Hydrobius scabrosus Horn
Paracyrnus subcuprens (Say)
Crepitis moratus (Horn)
Helochares maculicollis Mulsant
Enochrus hamiltoni (Horn)
Enochrus conjunctus (Fall)

Dryopoidea
Dryopidae

Helichus striatus LeConte
Elmidae

Elmis ornata Schaffer

Helrrlimnius quadrimaculatus (Horn)

Heterlimnius corpulentus (LeConte)

Narpus concolor (LeConte)

Narpus angustus Casey

Simsonia quadrinotata (Say)

Zaitzevia parvulus (Horn)

Microcylloepus pusillus LeConte

Diptera

Chironomidae
Calospectra sp.
Chironomus sp.
Dianipsa sp.
Harnischia sp.
Hydrobaenus sp.
Cricotopus sp.
Corynoneura sp.
Psilodiamesa sp.
Syndiamesa sp.
Pentaneura sp.
Tendipes sp.
Tanytarsus sp.
Procladius sp.
Spaniotoma sp.
Tipulidae
Antocha sp.
Eriocera sp.
Dicranota sp.
Hexatoma sp.
Pedicia sp.
Holorusia sp.
Phalacrocere sp.
Tipula sp.
Rhaphidolabis sp.

(2), 5

5

5

5

5

5

5

5

5
(2), 5

5

5

5

5

5

5

(t). 2. 5

5

(1), 2, 5

2, 5

5

2

5
(1), 2, 5

2

(1)
6
2
6
6
6
6
6
6
6

2, 6
6
2
2
2

1, 2

2

2

1, 2

2

2

2

214

GREAT BASIN NATURALIST

Vol. 32, No. 4

Table 1. (Continued)

Psycodidae

Pericoma sp.
Empididae

Hemerodromia sp.

Roederiodes sp.
Simuliidae

Simulium sp.
Tabanidae

Chrysops sp.

Dixidae

Dixa sp.
Rhagionidae

Atherix variegeta Walker
Muscidae

Lininophora aequifrons Stein
Heleidae

Palpomyia sp.
Stratiomyiidae

Euparyphus sp.

Odontotnyia sp.
Deuterophlebiidae

Deuterophlcbia sp.
Blepharoceridae

Agathon sp.

Blepharicera sp.

E|)hemeroptera
Baetidae

Bactis bicnudalus Dodds

Baetis intermedius Dodds

Baetis tricaudalus Dodds

Callihactis fuscus Dodds
Ephonicifllidae

I-'.fthi-nirrrlla coloradensis Dodds

Ephcnicrclla doddsi Needhani

K[>fu'frirrfll(i grandis grandis Eaton

liplwinerrlla hecuba (Eaton)

Ephrnirrclln inerniis Eaton

Ephcnierrlla infrequens McDuiniougli

Ephrmcrella maragrita Necdham

Ephemerrlla tibialis MrDunnonRh
Heptageniidae

Cinygrnula mimus (Eaton)

Cinygmula par (Eaton)

Eprorus (Iron) albertae (McDunnougli)

Eperous dcccptivus (McDunnough)

Epeorus longimanus (Eaton)

Epcorus sj).

Heptagenia criddlei McDuiuiough

Hrptagcnia snlilaria IV1( Dunnough

Heptagenia elegantula (Eaton)

Rhithrngeria doddsi McDonnoungh

Bhithrogrna rnorrisoni (Banks)

Rhithrogena robusta Dodds
Siphlonuridae

Ameletus oregonensis McDunnough

Ameletus uelox Dodds

Ameletus sp.

1, 2

1, 2
2

2

2

1, 2
9

2

1, 2
1, 2
2

1, 2
2
9
2

9

9

2

2

2

2

2

1, 2

2

9

9

9

Dec. 1972 winger, et al: checklist 215

Table I . ( Continued )

Siphlonurus occidentalis Eaton 2

Parameletus columbine McDunnough 9

Leptophlebiidae

Paraleptophlebia debills (Walker) 2

Paraleptophlebia heteronea (McDonnough) 1, 2

Paraleptophlebia memorialis Eaton 9
Paraleptophlebia packi (Needham)

Tricorythidae

Tricorythodes minutus minutus Traver 1. 2

Plecoptera

Chloroperlidae

Alloperla pallidula (Banks)

Alloperla borealis (Banks)

Alloperla lamba Needham and Cleasen

Alloperla piniada Ricker

Alloperla coloradensis (Banks)

Alloperla severa (Hagen)

Alloperla signata (Hagen)

Paraperla frontalis Banks

Utaperla sopladora Ricker
Nemouridae

Brachyptera nigripennis (Banks)

Brachyptera pacifica (Banks)

Brachyptera pallida (Banks)

Capnia confusa Claassen

Capnia columbiana Claassen

Capnia gracilaria Claassen

Capnia lemoniana Nebeker and Gaufin

Capnia ligulata Hansen

Capnia logana Nebeker and Gaufin

Capnia nana wasatchae Claassen

Capnia uintahi Gaufin

Eucapnopsis brevicauda (Claassen)

Paraleuctra occidentalis (Banks)

Paraleuctra sara ((Claassen)

Isocapania crinata (Needham and Claassen)

Isocapania grandis (Banks)

Nemoura californica Claassen

Nernoura cinctipes Banks

Nemoura columbiana Claassen

Nemoura besametsa Ricker

Nemoura oregonensis Claassen

Nemoura haysi Ricker

Perlomyia utahensis Needham and Claassen
Perlidae

Acroneuria pacifica Banks

Claassenia sabulosa (Banks)
Perlodidae

Arcynopteryx signata (Hagen)

Arcynopteryi parallela (Prison)

Isogenus aestivalis (Needham and Claassen)

Isogenus modestus (Banks)

Isoperla ebria (Hagen)

Isoperla fulva Claassen

Isoperla mormona Banks

Isoperla patricia Prison

Isoperla pinta Prison

Diura knowltoni (Prison)

(1)

, 2

, 4, 7

4,

7

2,

7

4,

7

4,

7

4,

7

4,

7

4,

7

4.

7

1,

2

4, 7

2,
l'

4"
7

7

1,

4,

7

2.

4.

7

4.

7

7

7

7

7

7

4.

7

2

7

(2)

, 4

, 7

7

7

(1)

, 4

, 7

2,

4,

7

4,"

7

1.

2.

4, 7

2.

4^

7

7"

7

1,

2

4. 7

1,

2",

4, 7

(1)

2

, 4. 7

1.

'4,

7

2^

4.

7

T

(1)

, (2), 4.

1,

2.

4. 7

4,

7

2,

4,

7

4,

7

4,

7

4,

7

216

GREAT BASIN NATURALIST

Vol. 32, No. 4

Table 1. (Continued)

Pteronarcidae

Pteronarcys californica Newport
Pteronarcella badia (Hagen)

Trichoptera

Brachycentridae

Brachycenirus aspolus Ross
Brachycentrus americanus (Banks)
Brachycentrus occidentalis (Banks)
Brachycentrus sp.
Micrasema bactro Ross

Helicopsychidae

Helicopsyche borealis (Hagen)

Hydropsychidae

Arctopsyche grandis (Banks)
Cheumatopsyche analis (Banks)
Hydropsyche californica Banks
Hydropsyche cockerelli Banks
Hydropsyche occidentalis Banks
Hydropsyche oslari Banks
Parapsyche elsis Milne
Pycnopsyche sp.

Hydroptilidae

Agraylea multipunctata Curtis
A gray lea saltesea Ross
Hydroptila consimilis Morton
Hydroptila sp.

Neotrichia panneus Denning
Neolrichia sp.
Ochrotrichia atylata Ross
Ochrotrichia logana (Ross)
Tascobia brustia (Ross)

Lepidostomatidae

Lepisostoma pluvialr (Milne)
Lepisostorna unicolor (Banks)
Lepisostoma podager (McLachlan)
Lepisostoma cascadense (Milne)

Leptoceridae
Oecetis sp.

Limnephilidae

Chyranda centralis (Banks)
Dicosmoecus atripes (Hagen)
Dicosmoecus unicolor (Banks)
Hesperophylax consimilis (Banks)
Limnephilus arizona Ross
Limnephilus sp.
Neophylai sp.
Neothrrmnia alicia Banks
Oligophlebodes minutus (Banks)

Philoptomidae

Dolophilus garhriclla (Banks)
Trenlonius arqualis I Ranks)
Chiniarra sp.

Psyrornyiidae

Psychomyea flarida Hagen
Tinodes sp.

Rliyac ofjhilidae

Anagapetus debilis Ross
Glossosoma alascense Banks

1. 2, 4. 7
1, 2, 4, 7

3

1
2. 3

1. 2, 3

2. 3

1. 2, 3

(1), 2, 3
(2), 3
(1). (2)

3

3

3

2, 3
3

3
3
3

(2), 3
3
2
3
2
2. 3

(2). 3
3
3
3

1. 2. 3

3

(2). 3

3
(1), 2, 3

2. 3
(1), 3

2
(1). 3

3

2,
2,
2

3
3

3

(t).

Dec. 1972

WINGER, ET AL: CHECKLIST

217

Table 1. (Continued)

Glossosorna parvulum Banks
Glossosoma verdona Ross
Rhyacophila acropedes Banks
Rhyacophila angelita Banks
Rhyacophila coloradensis Banks
Rhyacophila hannstoni Ross
Rhyacophila hyalinata Banks
Rhyacophila lobifera Betten
Rhyacophila pellisa Ross
Rhyacophila verrula Milne

Hemiptera
Corixidae
Gerridae
Mesoveliidae
Saldidae
Belostomatidae

Odonata

Coenagrionidae
Aeshnidae

Aeshna palmata Hagen

Corduliidae

Somatochlora semicircularis Selys

Megaloptera
Sialidae
Sialis sp.

Crustacea
Cladocera
Daphnia sp.

Amphipoda
Gammarus sp.
Hyallella azteca (Saussure)

Isopoda

Asellus tomalensis Harford

Hydracarina

Atractides sp.
Calonyx sp.
Eylais sp.
Hydryphantes sp.
Hygrobates sp.
Laminipes sp.
Lebertia sp.
Limnesia sp.
Megapus sp.
Panisus sp.
Fiona sp.
Sperchon sp.
Testudacarus sp.

Oligochaeta

Hirudinea

Cystobranchus verrilli Meyer
Dina dubia Moore and Meyer
Dina parva Moore
Eropbdella punctata (Leidy)
Helobdella stagnalis (Linnaeus)
Nephelopsis obscura Verrill

2.

3

3

(1)

, (2), 3

3

3

1,

2

1,

2

1,

2

1

1

1,

2

1

1, 2
2

I, 2

1, 2

2

218

GREAT BASIN NATURALIST

Vol. 32, No. 4

Turbellaria

Poly cells sp.

Nematoda

MoUusca
Physdae

Physa sp.
Planorbidae

Gyraulus sp.
Lymnaeidae

Lymnaea sp.
Bulimidae

Fluminicola sp.
Ancylidae

Ferrissia sp.
Sphaeriidae

Sphaerium sp.

1, 2
1, 2

, 2

•Musser, 1961.
'Braithwaite. 1962.

Literature Cited

Anderson, R. D. 1960. Taxonomy, distribution, and biology of the Dytiscidae

of Utah. Unpublished Ph.D. Thesis. Univ. Utah. Salt Lake City, Utah.
Anonymous. 1948. Reclamation on the Prove River. Utah Fish and Game Bull.

5(9): 2-3.
Braithwaite, L. F. 1962. The taxonomic problem of Polycelis in the United

States. Unpublished M.S. Thesis. Brigham Young Ihiiv. Provo, Utah.
Brooks, G. D. 1955. The Chironomidae of the Provo River. Unpublished M.S.

Thesis. Univ. Utah. Salt Lake City, Utah.
Chandler, H. P. 1941. Study of the aquatic and semiaquatic Coleoptera in

the state of Utah. Unpublished M.S. Thesis. Brigham Young Univ. Provo.

Utah.
DuNSTAN, W. A. 1951. A comparative study of a dredged and undredged por-
tion of the Provo River. Utah. Unpubli.shed M.S. Thesis. Univ. Utah. Salt

Lake City, Utah.
Edmunds, G. F., Jr. 1952. Studies on the Ephemeroplera. Unpublished Ph.D.

Thesis. Univ. Massachusetts.
Gaufin, a. R. 1949. A comparative study of the l)ottom fauna of the north

and south forks of the Provo River at St(nv;ut"s Riuu h. I tali. Utah Acad.

Sci., Arts, Letters Proc. 26:1.
. 1951. Production of bottom fauna in llie Provo River. Utah. Unpub-
lished Ph.D. Thesis. Iowa State Coll. Ames. Iowa.

- . 1959. Production of bottom fauna m the Provo River (tali Iowa

State Coll. J. of Sri. 33(3) : 395-41 9.
Gaufin, A. R., A. V. Nebeker, and J. Sessions. 1966. The stoneflies (Plecop

tera) of Utah. Univ. Utah Biol. Ser. 14(1): 1-93.
Gaumer, R. E. 1952. Taxonomy of some Utah aquatic mites (Hydracarina)

with notes on their biology. Unpublished M.S. Thesis. Univ. Utah. Salt

Lake City, Utah.
Ha7.7,ard, a. S. 1934. Quantitative studies of trout food in some Utah streams.

J^oc. Utah Acad. Sci. 11:271.
Merki.ey, D. R. 1948. The adult caddis flies of the Provo River. Unpublished

M.S. Thesis. Univ. Utah. Salt Lake City, Utah.
Moffett. J. W. 1936. A quantitative studv of the hottoni fauna in some Ihah

streams variously affected by erosion. Hull, Univ. of IMah 26(9): 1-32.
Mu.sser, R. J. (Stout). 1961. Dragonfly nvniphs of Utah. Unpublished M.S.

Thesis. Univ. Utah. Salt Lake City, Utah.

Dec. 1972

WINGER, ET AL: CHECKLIST

219

Needham. J. G., AND R. D. Christenson. 1927. Economic insects

streams of northern Utah. Utah Agri. Exp. Sta. Bull. 201.
Sessions. J. 1960. A study of the stoneflies of the Provo River, Utah.

lished M.S. Thesis. Univ. Utah. Salt Lake City, Utah.
Tanner, M. C. 1941. A study of the stoneflies of the Ogden River.

lished M.S. Thesis. Univ. Utah. Salt Lake City. Utah.
Todd, G. K. 1952. The adult water beetles of the Provo River. Unpublished

M.S. Thesis. Univ. Utah. Salt Lake City, Utah.

Unpub-
Llnpub-

A RECORD OF AN INDIGO BUNTING AND A
WINTERING SAY'S PHOEBE FOR NORTHERN UTAH

Gary L. Worthen'

Abstract. — A male indigo bunting (Passerina cyanea) and a female Say's
pheobe (Sayornis saya) were taken at Salt Lake City. Utah on 20 May 1966 and
2 February 1966, respectively. Both birds are the northernmost records for Utah
in the season in which they were taken.

On 20 May 1966 a freshly killed male Indigo Bunting (Passerina
cyanea) was found dead near the Union Building on the University
of Utah campus (4720 feet elevation), Salt Lake County, Utah. The
bird was found beneath some large windows, into one of which it
had apj)arently flown. The bunting seemed otherwise in good physi-
cal condition, weighed 17.6 grams, had no fat, and had testes that
measured 8x6 mm.

Indigo Buntings are considered to be rare but regular summer
residents in extreme southwestern Utah (Woodbury, et al., 1949;
Wauer, et al., 1965), but this account furnishes the first specimen
record of this species for northern Utah.

On 2 February 1966 a female adult Say's phoebe (Sayornis saya)
was taken from a night roost in a small shack at the abandoned Salt-
air Resort on the shore of the Great Salt Lake, 17 miles west of Salt
Lake City (4211 feet elevation). Salt Lake County, Utah (lat. 40°
47'N, long. 112°10'W). The bird was heavy in fat, weighed 19.55
grams, and was in apparently good condition except for the total
absence of the flight feathers of the tail. The loss of the tail was ap-
parently recent, as there was no indication of new feather growth in
the area. The bird was measured and compared to measurements
given by Bishop (1900), and would on that basis be assignable to
Sayornis saya yukonensis.

The bird's crop was empty, but the gizzard had a small amount
of unidentifiable food in it (.69 grams including the grit). Judging
by the bird's apparent good health, heavy fat layer, and gizzard con-
tents, it might be assumed that the bird was finding adequate food
despite the rigorous Utah winter.

Woodbury, et al. (1949) list the Say's phoebe as a casual winter
resident, with records from only central and southern Utah, and it
is believed that this account constitutes the first wintering record for
this species for northern Utah. Both of the above specimens are lo-
cated in the University of Utah Museum of Vertebrate Zoology.

Referknces

Bishop. 1900. Descriptions of three birds from Alaska. Auk 17:113-120.
Wauer. R. H.. and D. L. Carter. 1965. Birds of Zion National Park and

vicinity. Zion Natur. Hist. Assn., 92 p.
Woodbury. A. M. C. Cottam and .1. W. Sugden. 1949. Annotated check-list

of the birds of Utah. Univ. Utah Biol. Ser. 11 (2): 1-40.

'Museum of Natural H stork-. University nf Kansas. Lawionip. Kansas (iliO+f

220

COMMENTS ON TWO NAMES IN AN EARLY UTAH FLORA

James L. Reveal'

Abstract. — Two new varieties described by Durand in 1859 from the Great
Salt Lake area of northern Utah are discussed. One. Erysimum asperum var.
purshii. is the oldest available name for the western United States variant of
the species, while the second name, Acerates decurnbens var. erecta, is a synonym
of Asclepias asperula. Neither name has been included in botanical reference
works nor in monographic studies. One new combination, Erysimum, asperum var.
amoenum, is proposed for the orange-flowered phase of the species found in
southern Colorado and Utah.

In 1859, Elias Durand of the Academy of Natural Sciences of
Philadelphia published an article entitled "A sketch of the botany of
the basin of the Great Salt Lake of Utah," in which he summarized
the flora based on collections made by John C. Fremont (1843 and
1845), Howard Stansbury (1849 and 1850), and Edwin O. Beck-
with (1854), all associated with various expeditions sponsored by
the United States government, and Mrs. Jane Carrington of Salt
Lake City. Little is known about Carrington. Her collections were
delivered to Durand by Colonel Thomas L. Kane, and Durand al-
ludes to her as a "Mormon lady" in an unpublished catalogue of
his plant collection; I have been unable to add more than the 1857
date when she obtained her plants (Reveal, 1972). Since she was the
first woman botanist (or at least plant collector) in Utah, it is hoped
that historians will discover more about this person.

While in Paris recently, I had an opportunity to review Carring-
ton's collections deposited in the Durand Herbarium at the Museum
National d'Histoire Naturelle, Laboratoire de Phanerogamie. While
time did not permit a full investigation of the 59 species attributed
to her by Durand, two specimens representing new entities were
studied.

Erysimum asperum. (Nutt.) DC. var. purshii Durand, Trans.
Amer. Philos. Soc. II, 11:159. 1859. Durand states: "The very same
form as Pursh's specimen in herbarium of Phila. Acad, of Nat. Sci.
Stems simple, several from the same root, smaller than the preceding
[sic, alluding to var. asperum], scarcely 1 foot high, few-flowered;
radical leaves entire, or nearly so; siliques l'/2-2 inches long."

Lectotype: Utah: Salt Lake Co.: Near the Great Salt Lake,
Salt Lake Valley, June 1857, Carrington s.n. Holotype, P!

The Pursh collection, cited above, is also in the Durand Herbar-
ium, but without data. It seems wise to select the Carrington speci-
men as the lectotype although the description could equally apply to
either collection.

Erysimum asperum is a widespread and highly variable species
that is composed of several weakly defined varieties. The type of the

'Department of Botany, University of Maryland, College Park 20742, and National Museum of
Natural History, Smithsonian Institution, Washington, DC. 20560.

221

222 GREAT BASIN NATURALIST Vol. 32, No. 4

species comes from the Great Plains and has more or less spreading
fruits. The western United States material with yellow flowers and
erect fruits should be called var. purshii. Hitchcock (1964) was
unable to determine which name should be applied to this phase, be-
ing unaware of Durand's publication. In most western floras, this
phase has been called E. capitatum (Dougl. ex Hook.) Greene, al-
though Welsh et al. (1965) call this plant simply E. asperum.

Those specimens with orangish or reddish flowers from the
southern Rocky Mountains and high mountains of Utah should be
called E. asperum var. amoenum (Greene) Reveal, comb. & stat.
nov., based on Cheiranthus nivalis var. amoenus Greene, Pittonia
3:137. 1896. Holmgren (1959) and Welsh et al. (1965) have called
this phase E. wheeleri Rothr. Similar plants occur sporadically in
the Pacific Northwest but appear to represent another kind as yet
undescribed (Hitchcock, 1964), while those of the southern Coast
Range of California are called E. asperum var. stellatum J. T.
Howell.

Acerates decumbens (Nutt.) Dene, in DC. var. erecta Durand,
Trans. Amer. Philos. Soc. II, 11:174. 1859. Durand states: "Stem
erect, 3 feet high. Leaves scattered, sometimes verticilate in threes,
ovate-lanceolate, 4-5 inches long and 1 broad, shortly petiolate.
Umbel terminal solitary, globose, 21/2-3 inches in breadth; pedicels
pubescent; calyx and corolla green, crown deep purple."

Type: Utah: Salt Lake Co.: Salt Lake City, 1857, Carrington
s.n. Holotype, P!

This variety is a synonym of Asclepias asperula (Dene in DC.)
Woodson var. asperula^ a common species which occurs in the cen-
tral United States and northern Mexico. Woodson did not account
for this name in his monograph of Asclepias (Woodson, 1954).

Literature Cited

Durand, E. 1859. A sketch of the botany of the basin of the Great Salt Lake

of Utah. Trans. Amer. Philos. Soc. II. 11:155-180.
Hitchcock, C. L. 1964. "Cruriferae." In Hitchcock. C. L., et al. Vascular

plants of the Pacific Northwest. Univ. Wash. Publ. Biol. 1 7(2) : 430-557.
Holmgren, A. H. 1959. Handbook of the vascular plants of the Northern

Wasatch. Lithotype Process Co., San Francisco.
Reveal, J. L. 1972. Botanical explorations in the Interniountain Region, pp.

40-76. In Cronquist, A., et al. Intermountain Flora: the vascular plants of

the Interniountain West. U.S.A. Hafner Publish Co.. New York.
Welsh. S. L.. M. Treshow, and G. Moore. 1965. Common Utah plants.

Brigham Young Thiiv. Press, Provo, Utah.
Woodson, R. E.. .Jr. 1954. The North American species of Asclepias. Ann.

Missouri Bot. Gard. 41:1-211.

NEARCTIC DESERT SAND DUNE ORTHOPTERA
PART XIV. A NEW EREMOPEDES (DECTICIDAE)

Ernest R. Tinkham'

Abstract. — A new species, Ereniopedes kelsoensis, is described from the
Kelso Sand Dunes. San Bernardino Co., California, with notes on its biology,
song, host plants, desert distribution, orthopteran associates, and a key to
species in the genus.

The genus Eremoped.es is one of the best defined of the various
eremophilous genera of our Southwestern Decticidae. The geno-
type E. Scudderi Cockerell is largely confined to the Coahuila
Desert, one of three eremological components of the Great Chihua-
huan Desert; the other two being the northern Pecos and the ex-
treme southeastern Salado deserts. E. covilleae Hebard is known
only from the type locality about 4-5 miles SE of Persimmon Gap
in the Coahuila Desert, not nine miles S of the park entrance as
assumed by Rentz and Birchim. E. shrevei Tinkham is known only
from the type locality in the Salado Desert. E. bilineatus (Thomas)
is widespread in southeastern Arizona and southern New Mexico;
E. halli inhabits the pine zone in the mountains of Arizona and New
Mexico. E. pallidus Tinkham, originally described as a variety of E.
balli, inhabits sand dune areas of the Painted Desert, one of four
eremological components of the Great Basin Desert. The giant E.
ephippiatus sonorensis Tinkham inhabits the Ilermosillo Desert, the
southernmost of seven eremological components of the Great Sono-
ran Desert.

The new species herein described is largely confined to certain
xeric dune areas and their sparse vegetation in the Lahontan Desert,
the westernmost member of the Great Basin Desert.

Eremopedes is easily recognized by the long barrel of the prono-
tum, with its well-developed lateral lobes, and by the characteristic
cerci of the males which demonstrate two types: in scudderi, bilinea-
tus, shrevei, covilleae, and the new species the cercus is long and
slender, somewhat undulate with median internal tooth; in balli,
pallidus, ephippiatus and e. sonorensis the male cercus is much
shorter and heavier. The status of E. spinosa Hebard remains to be
determined.

Key to the Species of Eremopedes

1. Cerci slender, elongate, somewhat undulate, with internal

median subapical prominence and tooth; metazonal

lobes pale 2

Cerci short, heavy, rather quadrate, with internal, subapi-
cal tooth; metazonal lobes black 6

2. Color green or gray; size large to small; form slender.

+41 Date Palm Avenue, Indio, California 92201

223

224 GREAT BASIN NATURALIST Vol. 32, No. 4

with white dorso-lateral stripe running length of prono-

tum and body 3

Color green or gray; size medium, form heavier; dorso-
lateral stripe, if present, confined to abdomen 4

3. Size large; Sahuaro Desert bilineatus

Size small; Great Basin Desert kelsoensis

4. Dorsolateral lobes of pronotum with marginal areas pale;

metazona concolorous with prozona 5

Dorsolateral lobes of pronotum with pale arcuate area;

metazona broadly buff in color; Coahuila Desert .... covilleae

5. Size medium; penultimate abdominal notite very deeply

notched; Coahuila Desert scudderi

Size small; grayish; posterior penultimate notite shallowly

notched; Salado Desert shrevei

6. Size very large; Hermosillo Desert e. sonorensis

Size medium to small 7

7. Outer pagina of caudal femora unmarked; Painted Desert

pallidus

Outer pagina of caudal femora with two black longitu-
dinal stripes 8

8. Cerci of male as broad as long; Oak Zone e. ephippiatus

Cerci more slender; Pine Zone halli

Figs. 1-3. Eremopedes kelsoensis, n. sp.: 1, dorsal view of terminalia of male
holotype showing penultimate abdominal tergite with U-shaped notch, cerci, left
titillator, and cerci of subgenital plate; 2, ventral view of subgenital plate of male
holotype; 3, ventral view of subgenital plate of female allotype.

Eremopedes kelsoensis. n. sp.

Comparison: E. kelsoensis is most closely related to the much
larger bilineatus and is entirely confined to desert regions west of
the Colorado River, whereas E. bilineatus is restricted to desert re-
gions oast of the Colorado River. There is no area of contiguity or
intergrade. Relationships with tiio other species are clearly expressed
in the Key.

Dec. 1972 tinkham: desert orthoptera 225

Description. — Male holotype: Size small, slender, typical of
more slender element of Eremopedes. Face typical of genus. Prono-
tum almost twice as long as broad; fore margin almost squarely trun-
cate and very slightly emarginate; posterior margin almost squarely
truncate; lateral lobes of pronotum fairly deep, especially at prozonal
lobe; shallower in metazonal lobe area, ventral margin showing a
very slight emargination immediately above oval tympanum which
lacks hairy margin.

Abdominal penultimate notite bearing a moderate U-shaped
emargination, but much shallower than very deep V-shaped
emargination of bilineatus. Cerci slender, quite elongate, outer mar-
gin slightly undulate, inner margin bearing an internal, median,
subapical prominence tipped with a minute tooth. Subgenital plate
narrow, keeled, with a deep U-shaped median notch on posterior
margin. Titillators straight, slightly sj)athulate with a row of retrose
teeth on exterior margin.

Leg spination: Fore legs with long procoxal spine; fore femora
unarmed; fore tibiae with ventral keels bearing six pairs of long
aciculate spines (two pairs in basal half, four pairs in apical half),
and outer dorsal ridge with basal spine on apical portion of syn-
chronometer, the other two, one medianly and one apical. Mesotibiae
with six pairs of ventral spines and four internal and two external
dorsal spines widely and irregularly distributed. Caudal femora with
externoinferior keel unarmed; internoinferior keel with six widely
spaced small teeth in apical two-thirds. Caudal tibiae with 20-22
spines on ventral keels and six pairs of dispersed dorsal spines.

Living coloration: General coloration pale viridian. Head with
entire face, antennal scrobes, and fastigium viridian; cheeks very
pale green, occiput same palely tinged with very pale orange. Prono-
tum with median dorsal area dark green, outwardly edged with
rusty orange; dorsolateral area nacreous white; dorsolateral lobar
area deep green; prozonal margin nacreous, metazonal area rusty
brown; meso- and metathoracic areas green. Abdominal notites deep
green, with nacreous dorso-lateral stripe having both margins edged
irregularly with rusty red. Sternites viridian; ventrolateral areas
nacreous. Fore and middle legs greenish with some pale reddish
tinge dorsally. Caudal legs with femora largely green, sulcus ventral-
ly tinged with very pale red; tibiae pale viridian. Tegmina pale red-
dish brown.

Female allotype: Closely similar to holotype in body size and
coloration; fore and middle legs with spination as in holotype. Cau-
dal femora with only one spine on inner ventral keel. Caudal tibiae
with 24-26 ventral teeth on each keel; 5-6 widely spaced dorsal
teeth. Ovipositor long, exceeding the length of the body and slightly
recurved in the apical three-quarters. Subgenital plate circular in
outline with an open V-shaped median notch on the posterior mar-
gin.

Type material. — Male holotype: Kelso Dunes, 10, ll-VI-1958,
nymph which became adult 1-VIII, Creosote margin of dunes, E. R.

226 GREAT BASIN NATURALIST Vol. 52, No. 4

Tinkham. Calipered measurements in mm: body length 21.1; prono-
tum 5.8 X 4.8 max. breadth; exposed tegmina 3.0; caudal femora
19.8 X 2.5; cerci 2.1. Type in the Tinkham Eremological Collection.

Female holotype: Dunes on north side of Owens dry lake, Owens
Valley, California (several miles west of Keeler), 26-VIII-1957; on
Suaeda torreyana, night collecting, E. R. Tinkham. Calipered mea-
surements in mm: total length to tip of ovipositor 43.2; ovipositor
25.1 X 1.4; body length 19.8; caudal femora 20.8 x 2.5. Allotype in
the Tinkham Eremological Collection.

Male paratypes: 1 cf , Kelso Dunes, San Bernardino County, 17-
X-1970, on Hymenoclea in sand dune arroyo; 1, Kelso Dunes in
Creosote margin, 11 -VI- 1958, 2, N side Owens Lake dunes, 26-VIII-
1957, on Suaeda; 2, 1-2 miles W of Tonopah Dunes in Big Smokey
Valley about 15 miles NW Tonopah, 19-IX-1957. All collected by E.
R. Tinkham. 1, Adelante, 28-VIII-1957, Theodore Cohn.

Paratype males closely similar to holotype; size smallest from
Adelante, Owens Lake, and Tonopah, maximum size from Kelso
Dunes. Coloration closely similar except in Adelante male, which is
dark greenish gray in coloration.

Calipered range measurements in mm: body length 16.3-21.5;
pronotum 5.2-5.8 x 3.9-4.4; caudal femora 18.7-20.8.

Female paratypes: 3 same data as the allotype. Coloration same
as in allotype. 2 subadults with shorter ovipositors. Calipered mea-
surements in mm: adult, total length to apex of ovipositor 45.9; ovi-
positor 25.5, body length 23.0; pronotum 5.8 x 5.1; caudal femora
22.2. Eclosure of one subadult imperfect due to confinement. 1 female
subadult: total length to tip of ovipositor 40.0 mm; ovipositor 21.1
mm and straight; body length 20.2 mm; pronotum 5.6 x 4.8 mm;
caudal femora 22.3 mm.

Habitat. — At the Kelso Dunes E. kelsoensis inhabits the creosote
margin, in good wet years, where also Croton californicum. Rumex
hymenclytra, clumps of Petalonyx Thurhcr'u Ililarin rigida. and a
stiff silvery Gilia sp. are always present. In good years Oenothera
dcltoides, primaverens, Gilia aiirea decora, Baileya pauciradiata.
and many other flowers are present. The arroyo that cuts a deep
valley through the towering dunes is dominated by Larrea divaricata.
Acacia Greggii, Hymenoclea salsola, Opuntia echinocarpa. O. ramos-
simima, Haplopappus acradenius, and other plants.

On the north side of dry Owens Lake, rejiresenting the Lahontan
Desert, the principal dominants on the dunes are Atriplex conferti-
folia, A. canescens. Sarcohatus vermiculatus, Suaeda Torreyanna.
Artemesia tridentata, Franscria duniosa. Dalea polyadenia. and
Distichlis sp. One dune had a si)ring flowing out of the top of it, well
surrounded by tules and other water-loving plants. On the west side
of the towering Tonopah Dunes in Big Smokey Valley, about 15
miles NW of Tonopah, Nevada, the habitat ranged 1-3 miles west
of the dunes which carried a stunted vegetation of Atriplex spinifera,
Coldenia plicata, A. canescens, Dalea polyadenia, and perhaps other
rarer f)lants growing on the sandy soil.

Dec. 1972 tinkham: desert orthoptera 227

/'

Figs. 4-5. Eremopedes kelsoensis. n. sp. Female allotype: 4. dorsolateral as-
pect; 5, dorsal aspect.

Eremography. — The Tonoj^ah and Owens Lake dunes (north
and west sides) He within the periphery of the Lahontan Desert, the
westernmost component of the Great Basin Desert. This desert is
dominated by halophilous species of Atriplex in the sahne valley
bottoms and by Artemesia tridentata on the flanks or bajadas of the
valleys. The Great Sonoras Desert is dominated by Larrea divaricata
with its associated subdominants. The Kelso Dunes lying within the
confines of the Gila Desert show many more northern elements,
probably due to their elevation, which ranges between 2400 and 3100
feet, and their very cold winters, which undoubtedly make it possible
for this northern element to exist at a more southern latitude. Thus
the Tenebrionids, such as Lariverius, Trogloderus, and Eusattus, as
well as Ammcbaenetes and Eremopedes kelsoensis and perhaps
others, are examples of this eremographic picture.

Host plants. — At the Kelso Dunes, Hymenoclea salsola was the
host plant, but it is not known which plant serves as the host plant
in the creosote margin. At the north side of the Owens Lake dunes
Suaeda Torreyana was the host, but elsewhere the host plant is not
known.

228 GREAT BASIN NATURALIST Vol. 32, No. 4

Biology. — It is very evident that due to the severe winters of
its environment E. kelsoensis must hatch from the over-wintered
egg in the spring; its time of hatching varying with the character of
the spring and with the moisture in the soil or sand. The first speci-
men captured at the Kelso Dunes on 10-11 June 1957, was a male
nymph which became adult on 1 August 1957. This species was dis-
covered during the first of four summers of study supported by the
National Science Foundation, to which the author is most indebted.
The time of adulthood would also depend on the time of the ad-
ventitious rains, if they do arrive, the emergence ranging anywhere
from August into early October.

On 26 August 1957, at 10:30 p.m., the writer captured a pair that
had just mated in a Suaeda bush at the north Owens Lake dunes;
the male was still in the bush and the female was feeding on the
spermothecal sac. In 1970 the rains came late, and although they
did not touch the Kelso Dunes, heavy rains in the Granite and
Providence mountains to the east sent a big arroyo down through
the dunes for the first time in 15 years. This event accounted for
the discovery of the two males on 1 7 October on Hymenoclea sahola
after long and diligent searching.

Song. — the song is a soft and continuous "zee" characteristic of
the genus Eremopedes.

Orthopteran .assgci.ates. — At the Tonoj)ah location the only
associate was Coniana snowi, which is very rare in this xeric habitat.
At the Owens Lake dunes Anconia Integra, Eremiacris, Ligurotettix
coquilletti, Derotmema, and Aeoloplus were the acridids and the
tettigoniids were the new species and Anoplodusa arizoncnsis, the
latter being the first of this rare decticid reported for Owens Valley,
which is the Zoogeographical Center of this family. At the Kelso
Dunes the associates were more numerous: Acridids were Ciholacris,
Tanaeocerus, Tytthotyle, Coniana, Eremiacris, Bootettix, Xcracris.
and Irinierotropis p. pallidipcnnis. aUhough some of these have not
been seen since the fifties. The tettigoniids were Capnohotcs fidigino-
sus, Anoplodusa arizonensis (1954), and Insara covilleae; the rhaph-
idophorids were Amniobaenetes sp., Macrohaenetcs kelsoensis, Am-
mopelmatus kelsoensis, and Ceuthophilus fossor.

References

TiNKHAM, E. R. 1944. Biological, tiixononiir and faunistic studies on the shield-
back katydids of the North American deserts. Amer. Midi. Nat. 31(2):257-
328, figs. 1-28.

. 1962. Studies in nearctic desert sand dune Orthoptera. Part V. A new

genus and three new species of large sandtreader camel crickets from the
Colorado Desert, with keys and notes. Bull. So. Calif. Acad. Sci. 61(2):89-
111, 4 figs.

. 1965. Studies in nearctic desert sand dune Orthoptera. Part X. A new

genus and species of Stenopelmatine crickets from the Kelso Dunes, with notes
on its multi annual life history and kev. Great Basin Nat. 25(3-4) :63-72.
11 figs.

SEASONAL FOOD HABITS OF BARN OWLS IN UTAH

Dwight G. Smith,! Charles R. Wilson,- and Herbert H. Frost-

Abstact. — The food habits of a small colony of barn owls in central Utah
were investigated from 30 January 1969 to 30 January 1970 to determine seasonal
trends in prey species composition and abundance. An analysis of 783 pellets
yielded 1845 prey individuals, of which mammals, primarily Microtus. were the
most abundant durmg all seasons of the year. A wide variety of avian prey
species revealed an opportunistic aspect of barn owl prey selection.

A majority of the numerous studies of barn owl {Tyto alba) food
habits were determined from sporadic pellet collections deposited by
an unknown number of owls over an indefinite period of time.
Several exceptions include the investigations of the daily and season-
al food of barn owls in Davis, California (Evans and Emlen, 1947),
the report of seasonal food habits of barn owls from 14 locales in
England and Wales (Glue, 1967), the analysis of annual changes in
the diet of barn owls in France (Saint (jirons, 1968), and a long
term study of the food habits of this owl in Germany (Uttendorfer,
1952). Our objective was to determine the seasonal food habits of a
small colony of barn owls in central Utah. Prior to this study the
barn owl was considered to be an uncommon permanent resident in
Utah, and there was little information on its feeding ecology and
economic status from this part of the Great Basin.

The colony was in the abandoned fronton Steel Mill near Spring-
ville, Utah Co., Utah — a complex of 60 major brick and steel struc-
tures on approximately 500 acres of land. The colony numbered 26
individuals in the winter of 1968 but had increased to 38 individuals
by late June 1969. From July 1969 through January 1970 the colony
declined to 10 individuals. Large numbers of pigeons {Columba
livia), starlings (Sturnus vulgaris), and house sparrows (Passer
domesticus) also used the mill. The barn owls hunted within the steel
mill complex and in adjacent habitats. Although the interior of the
steel mill was largely devoid of vegetation, the adjacent habitats in-
cluded extensive cattail {Typha sp.) marshes, cheat grass (Bromus
tectorum) fields, and several shallow ponds. On the basis of 750
trap nights the most common small mammals within these habitats
included meadow mice {Microtus pennsylvanicus) , deer mice (Pero-
myscus maniculatus) , house mice (Mus musculus), and vagrant
shrews (Sorez va grans) .

Methods

The food habits of the barn owls were determined from pellet
collections — the most practicable method, despite necessary limita-
tions, for accumulating massive data on the food habits of nocturnal
raptors (Errington, 1967; Southern, 1969). Pellets were collected

'Department of Biology, Southern Connecticut State College, New Haven, Conn. 05515.
-Department of Zoology. Brigham Yoiing University, Provo, Utah 84601 .

229

230 GREAT BASIN NATURALIST Vol. 32, No. 4

biweekly from beneath the roosts of adult birds of the colony and
separated into four time groupings, each representative of a seasonal
period of barn owl activity and roughly corresponding with one of
the four seasons. The spring period from 2 March to 1 June corre-
sponded with the reproductive activities of the barn owl population;
the summer period from 2 June to 1 September was characterized
by the attentiveness of the adults to the newly fledged but still de-
pendent young; the autumn period from 2 September to 1 December
included the abandonment and subsequent dispersal of the majority
of the young from the colony; and from 2 December through 1
March the remaining owls moved into well-protected winter resi-
dence structures.

Pellet analysis followed methods described by Marti (1969). In-
dividual prey remains were identified by comparison with mammal
and avian specimens of the Brigham Young University Life Sciences
Museum natural history collections.

Pellet Deposition and Composition

Guerin (1928) reported that in France adult barn owls usually
deposited two pellets per 24 hours. The first pellet is dropped about
dawn on the hunting territory, while the second pellet is deposited
at the roosting site before the owl resumes its hunting the following
evening. We did not attempt to locate pellets dropped by the bam
owls of Iron ton on the hunting territory. At their roosting sites they
deposited an average of one pellet per day during spring, summer,
and early autumn, but this rate frequently declined during late
autumn and winter. This decline in the winter pellet deposition rate
correlated with severe weather conditions, and during several periods
of snow cover and extreme cold no pellets were deposited for one to
two days by one or more of the adult owls.

Spring and summer pellets were strikingly larger and averaged
almost twice as many prey individuals per pellet than pellets found
in autumn and winter (Table 1). In addition, summer pellets con-
tained up to eight prey individuals per pellet compared to a maxi-
mum of five individuals i)er pellet from the autumn and wiiiter pel-
lets. The smallest pellets were found during rigorous winter condi-
tions and usually contained but one prey individual. Errington

Table 1. Seasonal trends in pellet composition.

No.

Total

N

o. Indv

Av.

Drv Wt.

Pellets

No.

per

of

Pellets

Season

CoUecte-l

Indv.

Pellet

Range

in

grams*

Spring

346

889

2.6

1-7

10.6

Summer

198

509

2.6

1-8

9.1

Autumn

117

23t-

1.9

1-5

4.6

Winter

122

213

1.7

1-4

3.1

Totals

783

1845

Av.

2.2

Av.

1-6

•Determined from 100 randomly selected pellets from each season, wtiirli were air dried for
10 days.

Dec. 1972 smith, et al: barn owls 231

(1931) and Stewart (1952) also noted a progressive decline in the
size of the pellets deposited by barn owls during adverse climatic
conditions and reported that several of their owls subsequently died
of starvation. Although none of the barn owls of fronton died during
winter, it is probable that their ])opulations in Utah and other more
northern parts of their range may be limited by inherent food pro-
curement difficulties during severe weather conditions.

Composition of Diet

There were 1845 prey individuals of 21 different species, includ-
ing eight mammalian and 13 avian prey species (Table 2). On an
annual basis, mammals comprised 90% of the total prey and were
the most common prey of every season, although the frequency of
their occurrence declined significantly during autumn (x- =: 35.8,
P > 0.001, df = 3). In contrast, the frequency of avian prey, while
comprising only 10% on an annual basis, doubled during autumn.

Meadow mice were the most abundant mammalian prey species
and appear to represent the single most important food item (81%)
of the barn owls at fronton. Other important mammal prey species
included deer mice and house mice, but neither these nor any of the
remaining mammal species comprised over 3% of the barn owl diet.

Only two avian species were present in pellets from every sea-
son: the starling, which was the second most common prey species,
comprising 6.2% of the total annual prey; and the house sparrow,
which comprised 2.4% of the total annual prey. Both species nested
in fronton and adjacent locales, and large flocks roosted within the
buildings during the autumn and winter months. The frequency of
their occurrence as prey rose signficantly during autumn (x- =
16.9, P > 0.001 and x- = 23.4, P > 0.001 for the starling and house
sparrow respectively), coinciding with the sharp rise in their popula-
tions.

The majority of the other avian prey species were migrants taken
infrequently. Especially large numbers of transient birds fed and
roosted in the marshes and fields bordering fronton, and the spring
occurrence of a lesser yellowlegs {Totanus flavipes) and early
autumn occurrence of bank swallows (Riparia riparia) and red-
winged blackbirds (Agelaius phoeniceus) indicate that these mi-
grants occasionally present suitable prey for barn owls. The occur-
rence of lesser yellowlegs and American coot {Fulica arnericono) re-
veals that barn owls may take larger birds, although these must ap-
proach the upper limits of the prey-size capabilities of the owls. Do-
mestic pigeons were rarely found in the prey items, despite their
abundance within the mill.

The American kestrel (Folco sparverius) prey individual was a
recently fledged juvenile. Kestrels frequently perched in open, ex-
posed locations during the late evening hours at the time when barn
owls were initiating their nocturnal hunts; this individual was prob-
ably taken during this slight overlap in the activity periods of the
two species. In contrast, essentially no overlap occurred during the

232

GREAT BASIN NATURALIST

Vol. 32, No. 4

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Dec. 1972 smith, et al: barn owls 233

early morning hours because barn owls began roosting from one to
three hours before kestrels were active.

In addition to the vertebrate prey species, some vegetable matter
was recovered from the pellets. This material almost certainly repre-
sents the gut contents of the prey, and that which could be identi-
fied included fruits of Russian olive {Elaeagnus angustijolia) and
seeds of various grasses (Graminae).

Discussion

The barn owls at Ironton were supported exclusively by the
mammal and avian communities, despite the seasonal availability of
large invertebrate (primarily Insecta) and amphibian populations.
The year-round predominance of small rodents in their diet is in
agreement with the findings of similar investigations from other
areas of the range. This, coupled with the conspicuous lack of in-
vertebrates in their spring and summer diets, indicates selective pre-
dation.

Within the limitations of their food habits, however, the barn
owls at Ironton exhibited a considerable degree of opportunism. Haw-
becker (1945) and Wallace (1948) noted that the owls of their
respective studies tended to prey heavily on the most available ani-
mals of a community. Predation on the basis of availability is re-
flected in the present study by the high frequency of occurrence of
Microtus pennsylvanicus, which was the most common mammal in
the area.

The opportunism of barn owl predation is also revealed by both
the variation in total number of prey species taken in the different
seasons and the changes in seasonal frequency of several of the
prey species. The largest variety of i)rey species were taken in the
spring and summer months when transients and summer nesting
birds greatly increased the potential j)rey available to the local barn
owl population. In contrast, fewer different prey species were record-
ed from the decreased fauna of the autumn and winter months.

The changes in seasonal frequency of several of the prey species
was also a function of their comparative exposure at different times
of the year. This is shown by the autumn increase in the frequency
of occurrence of starlings and house sparrows, whose increased
autumn poi)ulations and i)reroosting flight behavior during the eve-
ning hours undoubtedly heightened their exposure to barn owl preda-
tion.

We conclude that the ability of the Ironton barn owls to ef-
fectively exploit the local prey populations was revealed by ( 1 ) their
heavy utilization of the most abundant prey species, (2) their pre-
dation on additional species when available at specific times of the
year, and (3) their response to local fluctuations of prey po})ula-
tions.

References

Errington, p. L. 1931. Winter killing of barn owls in Wisconsin. Wilson
Bull. 43:60.

234 GREAT BASIN NATURALIST Vol. 32, No. 4

Errington, p. L. 1967. Of predation and life. Iowa State University Press,

Ames. 277 p.
Evans, F. C, and J. T. Emlen, Jr. 1947. Ecological notes on the prey selected

by a barn owl. Condor 49:3-9.
Glue, D. E. 1967. Prey taken by the barn owl in England and Wales. Bird

Study 14:169-183.
Guerin, G. 1928. La vie des chouettes. Regime et croissance de I'effraye com-
mune Tyto alba alba (L) en Vendee. P. Lechevalier, Paris. 157 p.
Hawbecker, a. C. 1945. Food habits of the bam owl. Condor 47: 161-166.
Marti, C. D. 1969. Some comparisons of the feeding ecology of four owls in

north-central Colorado. Southwestern Natur. 14:163-170.
Saint Girons, M. 1968. Analyse des fluctuations du regime de I'effraye, Tyto

alba dans le department de la somme (Nord de la France) pendant une pul-

lulation de Microtus arvalis.
Stewart, P. A. 1952. Winter mortality of barn owls in central Ohio. Wilson

Bull. 64:164-166.
Southern, H. N. 1969. Prey taken by tawny owls during the breeding season.

Ibis 111:293-299.
Uttendorfer, O. 1952. Neue ergebnisse uber die ernahrung der Greifvogel

und Eulen. Eugen Ulmer. Stuttgart.
Wallace, G. J. 1948. The barn owl in Michigan: its distribution, natural

history, and food habits. Tech. Bull., Agric. Exp. Sta., Michigan 208:1-61.

GERMINATION OF PINUS ARISTATA ENGELM.

William H. Reid'

Abstract. — Seeds of Pinus arislata Engelm., bristlecone pine, from the type
locality at the headwaters of Clear Creek in Colorado have a geiTnination of 80
percent. Germination occurs within 10 days with or without vernalization. Scari-
fied seeds are invaded by fungus and less than 10 percent germinate. Seedlings
10 and 180 days old do not survive exposure to -IOC (14F) even when exposed
to gradually decreasing temperatures to -4C (25F) for five weeks before treat-
ment. Rapid germination, scarification mortality, and lack of cold hardness in
seedlings work against the reproduction of bristlecone pine.

In a recent study of the Pinus subsection Balfourianae (the fox-
tail pines) Bailey (1970) described the eastern California and
Nevada populations of P. aristata Engelm. as a new species, P.
longaeva Bailey. A review of the literature shows that nearly all
botanical work on P. aristata has used material from the far western
populations now defined as a separate species. The Great Basin
and Rocky Mountain populations — retained as P. aristata — re-
main virtually unstudied.

In Colorado, bristlecone pine occurs in rather small, pure, open
stands scattered throughout the south central and southwestern parts
of the state. The stands are found between 2500 meters (Cochetopa
Pass) and 3700 meters (Cumberland Pass), most frequently on the
gravelly soil of south- or west-facing slopes. This paper is part of
an analysis of P. aristata from the type locality — the headwaters
of Clear Creek in Colorado — and examines the germinability of
the seed in an effort to understand the limited occurrence of this
pine.

Seeds used in the study were collected from the north and east
slopes of Mount Evans at an altitude of 3000 meters during October
1970. Germination tests were started during April 1971.

Seed Morphology. — The seed of P. aristata is typical of the
genus Pinus (Martin and Barkley, 1961). The wing shows wavy,
fingerprint-like lines. The testa has a mottled, dark brown surface
and a papery brown nucellus. The length of the seeds with wings
averages 16 mm. Seeds in a sample of 100 weighed an average of
24.5 mg with wings and 22.4 mg without wings.

One hundred seeds were opened; 88 had a firm, white endo-
sperm and a pale yellow, well-defined embryo, with from 7 to 12
cotyledons. The 12 seeds without healthy embryos contained only
a flake of dry tissue.

Imbibition. — Fifty seeds with wings removed were submerged
in distilled water. At intervals they were removed, blotter dried,
and weighed. Initial water uptake was rapid: 18 percent during
the first hour. The weight gain continued at a reduced rate through
germination (Fig. 1).

'Department of Biology, University of Colorado, Boulder, Colorado 80302.

235

236

GREAT BASIN NATURALIST

Vol. 32, No. 4

200

100

50

20

10

0.5

12 10 20

ELAPSED TIME (hours)

100 200

Fig. 1. Imbibition of Pinus aristalo: percentage weight gain in water as a
function of time since immersion.

Germination. — Forty seeds were placed on moist filter paper
in petri dishes. Twenty were scarified by making a shallow 2-4 mm
long cut on the flat side of the seeds. All of these scarified seeds
were quickly invaded by fungus and none germinated. Of the 20
unscarified seeds, 15 (75 percent) germinated.

One hundred seeds were planted 10 mm deep in clay pots with
four parts sand to one part peat moss. Of these, 80 were exposed
to -IOC (14F) for 100 hours prior to planting. Thirty-two of these
were scarified as described above. All of the seeds had been col-
lected in October and had, no doubt, been exposed to subfreezing
temperatures prior to collection.

Germination occurred ra{)idly, with the first plants appearing
during the 4th day, and no additional plants occurring after the
1 0th (lay. Of 20 planted as collected, 16 (80 percent) germinated.
Of those vernalized but not scarified, 37 {77 percent) germinated,
and only 3 (8 percent) vernalized and scarified seeds germinated.
All of the seedlings not used for experiments were alive 60 days
later.

Dec. 1972 reid: bristlixone fink (iKrmination 237

Cold Hardness. — Four 14-day-old seedlings were exposed to
-IOC (14F) for 2 hours. All wilted and died. Six 180-day-old
plants were placed in a growth chamber set for a 12 hour day, a
day temperature of 13C (55F), and a night temperature of 7C
(45F). Over a period of 14 days, the day temperature was reduced
to 2C (36F). The day length was reduced to 9 hours. These
temperatures were maintained for another 21 days.

Three of the pine seedlings were then put into a plastic bag to
reduce dehydration and were placed in a freezer at -IOC for 48
hours. The plants were then returned to the growth chamber and
the temperature increased to 20C (68F). Those which had been in
the freezer browned in 5 days and died.

Discussion. — The tests suggest the following:

1. P. aristata germinates rapidly with a high yield (ca. 80 percent);

2. less than 10 percent of the seeds will germinate if they have
been scarified;

3. the seeds do not require long cold treatment prior to germination;

4. seedling bristlecone pine is sensitive to freezing and is killed by
exposure to -IOC (14F).

The seeds and seedlings of P. aristata do not appear to be well
adapted to the climate found in their habitat in Colorado. At tree
line, frost and snow can occur any month of the year. In the
fall, severe storms followed by warm weather and a thaw are fre-
quent, and the spring is often an irregular series of sunny days
and heavy snows lasting into .June (Marr, 1961). All of these
factors work against the successful reproduction of this pine. The
rapid germination must cause the death of many seedlings after
thaws, and those growing in temporarily moist locations must often
desiccate and die. In spite of these disadvantages, P. aristata, a pine
that lives as much as 800 years in Colorado, has survived without
morphological change in this area since the Miocene (Bailey, 1970).

Acknowledgments. — I would like to thank Dr. J. Bock for
assistance in this study, and K. R. Sullivan for making extensive
seed collections.

Literature Cited

Bailey, D. K. 1970. Phytogeography and taxonomy of the Pinus subsection
Balfourianae. Ann. Missouri Bot. Gard. 57:210-249.

Mark, J. W. 1961. Ecosystems of the east slope of the front range in Colorado,
University of Colorado Press. 1 34 p.

Martin, A. C, and W. D. Barkley. 1961. Seed identification manual. Univer-
sity of California Press. 221 p.

SPECIAL NOTICE

NOTICE OF A NEW REPORT. ENTITLED
"SYSTEMATICS COLLECTIONS: A NATIONAL PLAN . . ."

Draft copies of a Report entitled "Systematics Collections: A National Plan
. . ." will be available at no charge until 3/ January 1975 from Philip S.
Flumphrey. Interim Secretary. Association of Sjstematics Collections. Museum
of Natural History, University of Kansas, Lawrence. Kansas 66044.

The draft Report soon to be available, represents the combined efforts of a
great many systematists over a period of more than a year and takes into ac-
count the many constructive and thoughtful comments stimulated by the Sym-
posium on "Systematic Biology — The Development of a National Program on
Resources and Resource Management" held at the National Academy of
Sciences in July 1972. Following the Symposium in late September, a Writing
Conference was held at the Belmont Estate, chaired by representatives of the
Association of Sj'Stematics Collections, with generous support from the Smith-
sonian Institution. The 24 members of the "Belmont Writing Conference," all of
them systematic biologists, were unanimous in their feeling that the systematic
community and other communities of "users" of systematcs collections and as-
sociated resources should have an opportunity to review the Report and comment
on it before its final publication and distribution, whch is now scheduled for
March 1973.

The Report presents a "Natonal Plan" for making systematics collections
and their associated resources a more effective national resource, better able to
serve the needs of systematic biologists and those agencies of society which
depend on systematic information in seeking solutions to various biomedical,
environmental, agricultural, and other problems. The "Natonal Plan" pre-
sented in the draft Report is an important first step which, hopefully, will lead
soon to solutions to some of the most basic problems which have been plaguing
systematics collections and their associated resources for a great man>- years.

The editors responsible for preparing the Report for publication, will be
doing so during the latter part of February 1975. It is hoped that all those
interested in systematics collections and the services they provide will take
the opportunity to read the Report in its draft form and provide the editors
with thoughtful and constructive comment.s, all of which will be considered
by the editors in their efforts to make the Report a more effective instrument
for the communities it is designed to serve.- J. C. Dickinson, Interim Presi-
dent. Association of Systematic Collections.

238

INDEX TO VOLUME 32

The genera and species described as new to science in this volume
appear in bold type in this index.

A checklist of the macroinverte-
brates of the Provo River, Utah,
p. 211.

A new species of Gymnodamaeus
from Colorado, p. 97.

A new species of soil mite from
western Colorado (Acari: Crypto-
stigmata, Pelopidae), p. 208.

A new species of Stenamma (Hy-
menoptera: Formicidae) from
Utah, p. 35.

A new subspecies of Crotalus lepidus
from western Mexico, p. 16.

A record of an indigo bunting and a
wintering Say's phoebe for north-
ern Utah, p. 220.

A revision of the Psoralen lanceo-
lata complex (Leguminosae), p.
76.

Acanthotomicus bidentis, p. 191.

Allred, Dorald M., article by, p. 120.

Allred, Dorald M., and John D.
Johnson, article by, p. 154.

Aquatic Phycomycetes of Lily Lake,
Utah, p. 181.

Barnes, James R., Michael J. Dona-
hoo, Edward J. Peters, David A.
White, and Parley V. Winger, arti-
cle by, p. 211.

Clark, WiUiam H., and Peter L. Co-
manor, article by, p. 202.

Comanor, Peter L., and William H.

Clark, article by, p. 202.

Comments on two names in an early
Utah flora, p. 221.

Croft, B. A., article by, p. 61.

Dead great blue heron found at
11,000 foot Utah elevation, p. 112.

Dickinson, J. C, special notice from,
p. 238.

Dixon, James R., Wilmer W. Tan-
ner, and Herbert S. Harris, Jr.,
article by, p. 16.

Donahoo, Michael J., James R.
Barnes, Edward J. Peters, David
A. White, and Parley V. Winger,
article by, p. 211.

Eremopedes kelsoensis, p. 224.
Feuer, Robert C, and Hobart M.

Smith, article by, p. 55.
Five new varieties of Eriogonum

(Polygonaceae), p. 113.
Flights of the western thatching ant,

Formica obscuripes Forel, in Ne-
vada (Hymenoptera: Formicidae),

p. 202.
Forsythe, Dennis M., article by, p.

88.
Fossil palm materials from the Ter-
tiary Dipping Vat formation of

central Utah, p. 1.
Frost, Herbert H., Dwight G. Smith,

and Charles R. Wilson, article by

p. 229.
Germination of Pinus aristata

Engelm., p. 235.
Gregg, Robert E., article by, p. 35.
Gymnodamaeus chalazionus, p. 98.
Harris, Herbert S., Jr., James R.

Dixon, and Wilmer W. Tanner,

article by, p. 16.
Hickman, Gary L., article by, p. 112.
Higgins, Harold G., and Tyler A.

Woolley, article by, p. 208.
Higgins, Larry C, article by, p. 117.
Hoplopleura mendezi, p. 132.
Hoplopleura scotinomydis, p. 125
Hutchison, Earl R., and Ronald M.

Lanner, article by, p. 171.
Johnson, John D., and Dorald M.

Allred, article by, p. 154.
Johnson, Phyllis T., article by, p.

121.
Judd, B. Ira, article by, p. 91.
Lanner, Ronald M., and Earl R.

Hutchison, article by, p. 171.
McKnight, Kent H., and Hugh M.

Rooney, article by, p. 181.
Medlyn, David A., William D. Tid-

well, and Gregory F, Thayn, arti-
cle by, p. 1.
Nearctic desert Decticidae (Orthop-

tera). Part I: A new genus from

California, p. 176.

239

240

GREAT BASIN NATURALIST

Vol. 32, No. 4

Nearctic desert sand dune Orthop-
tera. Part XIV: A new Eremope-
des (Decticidae), p. 223.

New synonymy in American bark
beetles (Scolytidae: Coleoptera).
Part II, p. 190.

New synonymy in the bark beetle
tribe Cryphalini (Coleoptera: Sco-
lytidae), p. 40.

Notes concerning Mexican Saldidae,
including the description of two
new species (Hemiptera), p. 137.

Notes on Nevada solpugids, p. 120.

Notice of a new report entitled "Sys-
tematics collections: A national
plan," p. 238.

Observations on the nesting biology
of the long-billed curlew, p. 88.

On the rodent-infesting Anoplura of
Panama, p. 121.

Palmoxylon gustavesonii, p. 11.

Peloptulus tanytrichosus, p. 208.

Peters, Edward J., James R. Barnes,
Michael J. Donahoo, David A.
White, and Parley V. Winger, arti-
cle by, p. 211.

Petropedes, p. 176.

Petropedes santarosa, p. 177.

Plants new to the Utah Flora, p.
117.

Polhemus, John T., article by, p.
137.

Prey stage distribution, a factor af-
fecting the numerical and response
of Typhlodromus occidentalis to
Tetranychus mcdanieli and Tetra-
nychus pacificus, p. 61.

Reid, William H., article by, p. 235.

Relict stands of pinyon hybrids in
northern Utah, p. 171.

Reveal, James L., articles by, p.
113, 221.

Rhizopalmoxylon behuninii, p. 4.

Rhizopalmoxylon blackii, p. 7.

Rhizopalmoxylon scottii, p. 8.

Rooney, Hugh M., and Kent H. Mc-
Knight, article by, p. 181.

Saldula durangoana, p. 148.

Saldula saxicola, p. 143.

Scorpions of Utah, p. 154.

Seasonal food habits of barn owls in
Utah, p. 229.

Smith, Dwight G., Herbert H. Frost,
and Charles R. Wilson, article by,
p. 229.

Smith, Hobart M., article by, p. 104.

Smith, Hobart M., and Robert C.
Feuer, article by, p. 55.

Smith, Nathan M., and Wilmer W.

Tanner, article by, p. 25.

Stenamma knowltoni, p. 35.

Tanner, Wilmer W., James R. Dix-
on, and Herbert S. Harris, Jr.,
article by, p. 16.

Tanner, Wilmer W., and Nathan M.
Smith, article by, p. 25.

Thayn, Gregory F., William D. Tid-
well, and David A. Medlyn, article

by, p. 1.

The contributions of the 1822 works
of Jarocki and Fleming to herpe-
tological nomenclature, p. 55.

The Sonoran subspecies of the lizard
Ctenosaura hemilopha, p. 104.

Tidwell, William D., David A. Med-
lyn, and Gregory F. Thayn, arti-
cle by, p. 1.

Tinkham, Ernest R., articles by, p.
176, 223.

Toft. C. A., and S. L. Welsh, article
by, p. 76.

Two new subspecies of Crotaphytus
(Sauria: Iguanidae), p. 25.

Vegetation zones around a small
pond in the White Mountains of
Arizona, p. 91.

Welsh, S. L., and C. A. Toft, article
by. p. 76.

White. David A., James R. Barnes,
Michael J. Donahoo, Edward J.
Peters, and Parley V. Winger, arti-
cle by, p. 211.

Wilson, Charles W., Herbert H.
Frost, and Dwight G. Smith, arti-
cle by. p. 229.

Winger, Parley V.. Edward J. Pe-
ter. Michael J. Donahoo, James R.
Barnes, and David A. White, arti-
cle by, p. 211.

Wood, Stephen L., articles by, p. 40,
190.

Woolley, Tyler A., article by. p. 97.

Woolley. Tyler A., and Harold G.
Higgins, article by, p. 208.

Worthen, Gary L., article by, p. 220.

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TABLE OF CONTENTS

Aquatic Phycomycetes of Lily Lake, Utah. Hugh M. Rooney

and Kent H. McKnight 181

New synonymy in American bark beetles (Scolytidae:

Coleoptera), Part II. Stephen L. Wood 190

Flights of the western thatching ant, Formica obscuripes

Forel, in Nevada (H5Tnenoptera: Formicidae). William

H. Clark and Peter L. Comanor 202

A new species of soil mite from western Colorado (Acari:

Cryptostigmata, Pelopidae). Harold G. Higgins and

Tyler A. WooUey 208

A checklist of the macroinvertebrates of the Provo River,

Utah. Parley V. Winger, Edward J. Peters, Michael J.

Donahoo, James R. Rames, and David A. White 211

A record of an indigo bunting and a wintering Say's phoebe

for northern Utah. Gary L. Worthen 220

Comments on two names in an early Utah flora. James L.

Reveal 221

Nearctic desert sand dune Orthoptera, Part XIV. A new

Eremopedes (Decticidae) . Ernest R. Tinkham 223

Seasonal food habits of barn owls in Utah. Dwight G. Smith,

Charles R. Wilson, and Herbert H. Frost 229

Germination of Pinus aristata Engelm. William H. Reid 235

Notice of a new report entitled "Systematics collections:

A national plan." J. C. Dickinson 238

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