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HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

The Great Basin Naturalist

VOLUME 33, 1973

Editor: Stephen L. Wood

Published at Brigham Young University, by
Brigham Young University

TABLE OF CONTENTS
Volume 33

Number 1 — March 31, 1973

Helminths of Sceloporus lizards in the Great Basin and
upper Colorado plateau of Utah. Richard C. Pearce
and Wilmer W. Tanner 1

Miscellaneous chromosome counts of western American

Plants — II. James L. Reveal and Eloise L. Styer 19

Incidence of spotted fever in wood ticks of Utah recreational

sites. C. Selby Herrin 26

Notes on the nesting behavior of Steniolia elegans (Hy-

menoptera: Sphecidae). Howard E. Evans 29

A Taxonomic revision of Physaria (Cruciferae) in Utah.

Sheldon B. Waite 31

Two new species of Gymnodamaeus from Colorado (Aca-
rina: Cryptostigmata, Gymnodamaeidae). Tyler A.
Woolley and Harold G. Higgins 37

Neartic desert Decticidae (Orthoptera). Part II. A new

genus and species from Arizona. Ernest R. Tinkham 43

An unusual population of spiders in Utah. Dorald M. Allred .... 51

Some helminths from mink in southwestern Montana, with
a checklist of their internal parasites. Delbert L. Bar-
ber and Lawrence L. Lockard 53

Number 2 — June 30, 1973

Three new species of Palmoxylon from the Eocene Green
River formation, Wyoming. William D. Tidwell,
David A. Medlyn, and Gregory F. Thayn 61

On the taxonomic status of Platypodidae and Scolytidae

(Coleoptera). Stephen L. Wood 77

Studies on Utah stoneflies (Plecoptera). Richard W. Bau-

mann 91

The male brachycistidine wasps of the Nevada Test Site

(Hymenoptera: Tiphiidae). Marius S. Wasbauer 109

Notes on aquatic and semiaquatic Hemiptera from the
southwestern United States (Insecta: Hemiptera).
John T. Polhemus 113

A new subfruticose Eriogonuni (Polygonaceae) from west-
ern Colorado. James L. Reveal 120

Records of Coreidae (Hemiptera) from the Nevada test

site. Dorald M. Allred 123

Density, growth, and home range of the lizard Uta Stans-
buriana stejnegeri in southern Dona Ana County, New

Mexico. Richard D. Worthington and Edward R.

Arviso 124

Small bones of the hypsilophodontid dinosaur Dryosaurus
alius from the upper Jurassic of Colorado. Peter M.
Galton and James A. Jensen 129

Number 3 — September 30, 1973

Ecology of Sceloporus magister at the Nevada Test Site,

Nye County, Nevada. Wilmer W. Tanner and John

E. Krogh 133

Further studies on the wasps of Jackson Hole, Wyoming

(Hymenoptera, Aculeata). Howard E. Evans 147

Additional records of mutillid wasps from the Nevada Test

Site. Dorald M. Allred 156

The effects of soil texture on species diversity in an arid

grassland of the eastern Great Basin. John W.

Wyckoff 163

New synonymy in American bark beetles (Scolytidae:

Coleoptera). Part III. Stephen L. Wood 169

Undescribed species of Neartic Tipulidae (Diptera). XI.

Charles P. Alexander 189

Neartic desert Decticidae (Orthoptera). Part III: The

true tjrmpanum in certain genera, with key. Ernest

R. Tinkham 197

Notes on reproduction in Lampropeltis triangulum and

Coluber constrictor in Utah. William L. Grogan and n

Lloyd C. Pack, Jr 202

Courtship behavior among white-tailed and black-tailed

jackrabbits. Del F. Blackburn 203

Number 4 — December 31, 1973

Local distribution and interspecies interactions in micro-
tines. Grand Teton National Park, Wyoming. Tim

W. Clark 205

Notes on the occurrence and distribution of Pteronarcys
calif ornica Newport (Plecoptera) within streams. John

A. Elder and Arden R. Gaufin 218

Prehistoric bighorn sheep in the northern Sierra Nevada,

California. Peter D. Schulz and Dwight D. Simons 221

Artemesia arbuscula, A. longiloba, and A. nova habitat

types in northern Nevada. B. Zamora and P. T. Tueller .... 225

Significant bird records from Utah. William H. Behle 243

Small mammals of the National Reactor Testing Station,

Idaho. Dorald M. Allred 246

Scorpions of the National Reactor Testing Station, Idaho.

Dorald M. Allred 251

Density changes and habitat affinities of rodents of shad-

scale and sagebrush associations. Earl J. Larrison and

Donald R. Johnson 255

New species of American Microcorthylus (Coleoptera:

Scolytidae). Stephen L. Wood 265

The nominal snake genera Mastigodryas Amoral, 1834, and
Dryadophis Stuart, 1839. Hobart M. Smith and Ken-
neth R. Larsen 276

Vokune 33, No. 1
March 31, 1973

The

iARVARD

Great Basin

Published by
Brigham Young University

GREAT BASIN NATURAUST

Editor-. Stephen L. Wood, Department of Zk)ology, Brigham Young

UniversitT, Provo, Utah

>. I ixuey L. Welsh, Botany, Chairman; Wilmer W.
r, Zoology; Joseph R. Murdock, Botany; Vernon J.
1 ipton, Zoology; Ferron L. Andersen, Zoology

Ex officio Editorial Board Members:

A. Lester Allen, Dean, College of Biological and AgricuJ-

tural Sciences

Ernest L. Olson, Director, University Press, University

Editor

Tlic Great Basin Naturalist was founded in 1939 by Vasco M.
Tanner. It has been continuously published from one to four times
a yeor since then by Brigham Young University, Provo, Utah. In
!j:{\uora], only original, previously impublished manuscripts pertain-
ing to Xho biological, natural history of the Great Basin and western
North Ariierica will be accepted. Manuscripts are subiect to the ap-
! roval of the editor.

ipTiON: The annual subscription is $5.00 (outside the

lies $5.50). The piice for single numbers is $2.00 each.

Iters pertaining to the purchase of subscriptions and back

Jinnl,^ be directed to Brigham Young University Press,

205 UPB, Provo, Utah 84601. Libraries or other

' d in obtaining this journal through a contin-

;irly publications should contact the Brigham

rian. The Library, Provo, Utah

y for the Greet

to ttie editor. Contributors

i>n tlie back cover of a recent

The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

Volume 33 March 31, 1973 No. 1

HELMINTHS OF SCELOPORUS LIZARDS IN THE
GREAT BASIN AND UPPER COLORADO PLATEAU OF UTAH

Richard C. Pearce^ and Wilmer W. Tanner^

Abstract. — Fifty-five Sceloporus lizards representing the four species oc-
curring in Utah were dissected. For each lizard specimen and species, helminth
parasites were tabulated and classified. New host and locality records were
recorded. An appendix containing literature citations of reptilian parasitology for
the American Southwest is provided.

The literature dealing with the parasitic fauna of the south-
western United States contains little information on helminths of
Sceloporus lizards. Previous studies are concerned more with a
single genus or species of parasite than with the total parasitic fauna
of a particular host. Even more scarce are studies of parasitic fauna
from hosts in defined geographical areas.

Literature on reptilian parasites from Sceloporus lizards of the
Great Basin and Upper Colorado Plateau of Utah is sparse and scat-
tered. Grimdmann (1957) examined 47 lizards belonging to six
Utah species, one of which was S. occidentalis; he reported the para-
sites to be host specific. Grundmann (1959) later reported specific
names of hosts and parasites, and locations of parasites within the
host. Woodbury (1934) studied three species of Utah reptiles, in-
cluding S. graciosus. He recovered only one species of nematode,
probably because he studied only the stomach. Burkholder (personal
communication) examined over 300 stomachs of S. graciosus and
found no nematodes. He did, however, recover a cestode from the
intestines of each of two lizards.

Parasites from Sceloporus of neighboring areas have been ex-
amined and provide useful comparisons with Great Basin and Upper
Colorado Plateau populations. Waitz (1961) studied the parasites
of Idaho reptiles, including S. graciosus and S. occidentalis. He re-
covered Oochoristica scelopori from the former and Skrjabinoptera
phrynosoma from the latter. Harwood (1932) studied the helminth
parasites of reptiles neai? Houston, Texas, and found in S. undulatus
the nematode Oswaldocruzia pipiens. His paper was the first major
study on reptilian parasites from a specific geographical area in the
United States. Tanner and Hopkin (1972) studied the ecology of

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

2 GREAT BASIN NATURALIST Vol. 33, No. 1

S. occidentalis at the Nevada Test Site and reported 41 of 145 lizards
with unclassified round worms in the stomach.

Telford (1970) reported on 1485 lizards of southern California,
the coastal islands, and Mexico, including S. magister, S. graciosus,
and S. occidentalis. He lists many helminths, but he generally did
not report the degree of infestation or the location of the parasite in
the host. Stebbins and Robinson (1946) analyzed lizard populations
in parts of California which included S. graciosus. They found three
species of helminths in nine of the 15 animals examined. The para-
sites were classified but were described as being only from the
digestive tract.

The present project was designed as a survey of helminth infesta-
tions in Sceloporus lizards of the Great Basin and Upper Colorado
Plateau of Utah. It also contains a bibliography of literature on
reptile parasitology in the southwestern United States (Appendix).

We gratefully acknowledge the help and advice of Drs. Ferron L.
Andersen and Glen Moore, Departments of Zoology and Botany,
respectively, Brigham Young University. Mr. Gary L. Burkholder
supplied data from his study on 5. graciosus from central Utah.

Methods and Procedures

Host lizards were noosed, removed to the laboratory, and either
anesthetized with ethyl ether or pithed. They were opened from the
throat to the vent and microscopically examined for parasites in the
coelom, mesenteries, and intact organs. The liver, lungs, esophagus,
stomach, and small and large intestines were then removed and
placed in a 75 percent physiological saline solution in preparation
for dissection. Approximately 25 minutes after dissection, the organs
were reexamined and placed in numbered vials. The dissected
lizards were hardened in 1 0 percent formalin and added to the herpe-
tology museum. The method outlined by Meyer and Penner (1962)
for processing helminths was followed. Worms from preserved
lizards were simply removed and placed in numbered vials. Para-
sites were cleared in a solution of lactophenol for identification.

Literature used in the classification of helminths include llyman
(1951) and Yorke and Maplestone (1926) for the higher taxonomic
categories. For the specific identification of helminths of the order
Oxyuroidea, Gambino (1957a, 1957b), Lucker (1951), Read and
Amrein (1953), Telford (1965), and Thapar (1925) were used.
Spiruroid worms were identified by means of Babero and Kav
(1967), Morgan (1942), Ortlepp (1922), and Telford (1965).

Results

Results of this study are presented in four sections, one for each
species of Sceloporus studied. They are based on the dissection of 55
Sceloporus, 49 of which were anesthetized; the remaining six had
previously been preserved.

Sceloporus graciosus Baird and Girard. — Literature review re-
veals S. graciosus to be parasitized by four species of helminths —

March 1973 pearce, tanner: lizard parasites 3

one cestode and three nematodes (Table 4). The three nematodes
represent two orders: Spiruroidea {Skrjabinoptera phrynosoma) and
Oxyuroidea {Pharyngodon giganticus and Cyrtosomum penneri).

Of 22 dissections, 11 were of males and 11 females. The only
helminth recovered from the small intestine of S. graciosus was the
tapeworm Oochoristica scelopori. Although this worm was as long
as the entire small intestine (about 40 mm), it was contained within
the duodenum of its host (about 10 mm) .

All worms in the cecum were of Cyrtosomum and are considered
pin worms of reptiles (Gambino, 1957b). Sceloporus graciosus is a
new host for C. readi and C. heynemani, and, in addition, C. penneri
is reported for the first time from Utah lizards. The worms were
living free in the cecum of the large intestine and were in greatest
numbers around and inside of fecal material on which they ap-
peared to feed. Cyrtosomum species were not recovered from other
regions of the intestine.

Only 60.6 percent of the total helminth population in cecums
were adults. The remaining 39.4 percent were larval Cyrtosomum
not identifiable as to species.

The S. graciosus specimens used in this study were from Provo
Canyon and from the Little Sahara sand dunes southwest of Jericho,
Utah. Of nine specimens from the Provo area, seven harbored
helminths. It was in this group that O. scelopori was found. Six
lizards harbored only Cyrtosomum. Four of the 13 lizards from the
Jericho area were infested by Cyrtosomum. With the exception of
O. scelopori, helminths of all species were recovered from lizards of
these two areas. There were some differences in population composi-
tion in the worms recovered from the two areas (Table 3) .

Sceloporus undulatus elongatus Stejneger. — Three species of
nematodes, one from the stomach and two from the large intestine,
have been reported from S. undulatus as summarized in Table 1.
Oswaldocruzia pipiens belongs to the order Spiruroidea, and the two
species of Cyrtosomum are Oxyuroidea.

The results for this species are based on the dissection of 11
adults, 8 males and 3 females. Nine of these were noosed; the other
two had been preserved. Data for individual species of nematodes,
with the host being considered as one unit, is recorded in Table 4;
total helminth infestation per lizard is considered in Table 2.

The stomach worms, Skrjabinoptera phrynosoma, caused deep
welts in the stomach lining by imbedding their heads, which contain
a pair of intemolateral teeth. This mode of attachment was so strong
that the entire weight of the gastrointestinal tract could be supported
by a single worm when held with forceps.

Cecal worms varied in their life styles. Physaloptera retusa had
a similar mode of attachment to that of Sk. phrynosoma. Pharyngo-
don giganticus were free in the cecum and appeared to feed on fecal
material. Although P. retusa and P. giganticus both occur in the
cecum, the two did not occupy the same cecum. Skrjabinoptera
phrynosoma occurred in one individual w^ith P. retusa and in another

GREAT BASIN NATURALIST

Vol. 33, No. 1

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March 1973 pearce, tanner: lizard parasites 5

host with P. giganticus. These were the only cases of multispecies
infestations in S. undulatus.

All three lizards from the Zion National Park area harbored
helminths. One of these contained both the stomach worm Sk.
phrynosoma and the cecal worm P. giganticus. All lizards from
Arches National Park carried Sk. phrynosoma, and one a female
P. retusa in the cecum. None of the hzards from the St. George-
Santa Clara area had helminths. The one from Price, Utah, har-
bored S. phrynosoma (Table 3).

Sceloporus occidentalis biseriatus Hallowell. — Literature reveals
S. occidentalis to harbor six species of helminths (Table 4). Oocho-
ristica scelopori was the only cestode recovered. The nematodes
were Skrjabinoptera phrynosoma and Physaloptera retusa, order
Spiruroidea; and Cyrtosomum penneri, Pharyngodon giganticus,
and Thelandros iguanae, order Oxyuroidea.

Seven anesthetized and four preserved specimens were dissected.
Three species of helminths were recovered: O. scelopori, P. retusa,
and P. giganticus. Nine larval nematodes {Skrjabinoptera or Physa-
loptera) were recovered from a single cecum. Table 1 provides data
for the individual species of nematodes, whereas the total helminths
per infested host is given in Table 2.

The stomach worm Physaloptera retusa was always found in the
anterior third of the stomach. The mode of attachment was similar
to that previously noted for Sk. phrynosoma in Sceloporus undulatus.
There was, however, a major structural difference in that P. retusa
had the cuticle reflected over the lips, and the teeth were dissimilar

Table 2. Total helminths per lizard and infestations of host organs ex-
pressed in maximum, minimum, and average for the four species of Sceloporus
occurring in Utah.

Maximum

Minimum

Average Per
Infested Host

Helminths Per Lizard

S. graciosus
S. magister
S. occidentalis
S. undulatus

250

.240

1
2
1

140

102

5.7

Helminths Per Organ
Cecum

S. graciosus
S. occidentalis
S. undulatus
Duodenum

250

10
1
1

154
4
5

S. graciosus
S. occidentalis
S. undulatus

Large Intestine
S. magister

Small Intestine

1
1

200

1
1
1

150

4.4

150

S. magister
Stomach

100

S. magister
S. occidentalis

2
1

10.4
5

GREAT BASIN NATURALIST

Vol. 33, No. 1

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PEARCE, TANNER: LIZARD PARASITES

in number and structure (Ortlepp, 1922). As described for Sk.
phrynosoma, attachment was secure and caused welts in the stomach
Uning. When the stomachs were empty, the bodies of the worms
laid straight and usually passed anteriorly into the esophagus. When
food was present, their bodies were coiled in the anterior of the
stomach.

The tapeworm Oochoristica scelopori inhabited the duodenum of
the small intestine. This is the first report of O. scelopori in S. occi-
dentalis of Utah; it was previously reported from California (Tel-
ford, 1970).

Pharyngodon giganticus were only found to inhabit the cecal por-
tion of the large intestine and to move freely in the fecal material.
This is the first time P. giganticus has been reported in 5. occidenta-
lis of Utah. It was previously found in S. occidentalis of California
(Telford, 1970).

The cecum of one preserved lizard contained the eight larval
nematodes previously mentioned. They were not attached to the
intestinal wall when recovered, and no welts were seen on the cecal
lining. Anatomical characters are not sufficiently known to identify
them.

Host lizards were captured at four localities. Four came from
an area northeast of Dividend, Utah. Two of these were parasitized
by P. retusa, and one by P. giganticus. Two came from the Dugway
Mountains; one of these harbored P. retusa in the stomach and P.
giganticus in the cecum. Northwest of Elberta, Utah, four lizards
were collected, two of which harbored worms. One had the tape-
worm O. scelopori in the duodenum, and the other contained P.
retusa in the stomach and P. giganticus in the cecum. One lizard

Table 4. Record of helminths which have infested Sceloporus hosts.

Host Lizard

Reported in this Study

Reported from other Studies

S. graciosus

S. undulatus

S. occidentalis

S. magister

Oochoristica scelopori
Cyrtosomum penneri
Cyrtosomum readi*
Cyrtosomum heynemani*

Skrjabinoptera phrynosoma*
Physaloptera retusa*
Pharyngodon giganticus*

Oochoristica scelopori
Physaloptera retusa
Pharyngodon giganticus
Spiruroidea larva

Thubunaea iguanae
Cyrtosomum penneri
Cyrtosomum readi*
Physaloptera retusa*
Skrjabinoptera phrynosoma*

Oochoristica scelopori
Cyrtosomum penneri
Skrjabinoptera phrynosoma
Pharyngodon giganticus

Oswaldocruzia pipiens
Crytosomum heynemani
Cyrtosomum readi

Oochoristica scelopori
Physaloptera retusa
Pharyngodon giganticus
Skrjabinoptera phrynosoma
Cyrtosomum penneri
Thelandros iguanae

T hubunaea iguanae
Cyrtosomum penneri

* Indicates new host record.

8 GREAT BASIN NATURALIST Vol. 33, No. 1

was from the St. George-Santa Clara area; it had 13 P. retusa in the
stomach and one P. giganticus in the cecum.

Sceloporus magister Hallowell. — Literature on S. magister
(Table 4) reports two species of nematodes: Thubunaea iguanae, a
spiruroid, and Cyrtosomum penneri, an oxyuroid.

The nematodes S. phrynosoma, P. retusa, and C. readi are added
to the host lists given by Gambino (1957b) and Telford (1965,
1970). Thubunaea iguanae and C. penneri are also reported for the
first time to infest 5". magister from Utah (Table 4). Both nema-
todes were reported in S. magister from California (Telford, 1965).
Our data were obtained from 11 specimens (three male and eight
female) of S. magister.

Forty-five percent of the hosts were parasitized by Sk. phryno-
soma and T. iguanae; however, they were not found together in the
same host. Both were similarly attached to the stomach lining but
differed in their cutical structures, oral papillae, teeth, and repro-
ductive organs (Babero and Kay, 1967; Telford, 1970). Thubunaea
iguanae was not as deeply imbedded in the stomach lining as was
Sk. phrynosoma, although both worms were attached in the anterior
half of the stomach in all but one of the lizards. In the one excep-
tion, a preserved specimen, Sk. phrynosoma was free in the stomach.

Two hosts were infested by Cyrtosomum (Table 3). Two pre-
served lizards had these worms in the small intestine and cecum.
We have observed these parisites migrating when hosts were being
preserved; this may account for their presence in the small intestine.
An anesthetized specimen had approximately 150 Cyrtosomum free
in the cecum and one female P. retusa attached to the lining.

We examined 11 lizards; of the five from the St. George-Santa
Clara area, one contained S. phrynosoma. Of the three from Arches
National Park, all contained helminths in the stomach, two had
worms in the cecum, and one had worms in both the small intestine
and the cecum. The three remaining lizards came from eastern
Utah near Price. Two of these harbored helminths, one containing
six stomach worms, T. iguanae, and the other containing approxi-
mately 100 Cyrtosomum in the small intestine and 100 in the cecum.

Helminths recovered from their respective hosts as reported in
this study are compared to those reported in the literature in Table 4.
A summary of host relationships for the helminths recovered by us,
with results of past studies, are recorded in Table 1 .

Discussion

The prime objective of this study was the identification of
helminths in Sceloporus lizards of the Great Basin and Upper Colo-
rado Plateau of Utah. Fifty-five lizards representing four species of
Sceloporus were dissected and eight h(>lniinth species recovered.
The following is a discussion of each helminth species.

Oochoristica scelopori. — This is a tapeworm of reptiles. Two
specimens were recovered, one from the duodenum of S. graciosus

March 1973 pearce, tanner: lizard parasites 9

and one from the duodenum of S. occidentalis. Both hosts were
adult females.

This cestode has been reported from 5. graciosus and S. occi-
dentalis in southern California (Telford, 1970) and S. graciosus in
Idaho (Waitz, 1961). A single cestode was found in one of the nine
S. graciosus from Provo Canyon. Burkholder (personal communica-
tion) autopsied more than 300 Utah specimens of S. graciosus and
found one cestode each in two lizards also from Provo Canyon. The
probable explanation of this apparent nidus, or nested distribution,
lies with the unknown intermediate host. The intermediate host
presumably is an arthropod having a narrower environmental tol-
erance than S. graciosus. This possibility is strengthened by the
absence of this parasite in 13 lizards from the Little Sahara sand
dunes southwest of Jericho, Utah.

Telford (1970) attempted to measure the effect of altitude on
frequencies of parasitic infestations. He reported that the occurrence
of cestodes in lizards was due to the sympatric occurrence of inter-
mediate and definitive hosts and that the presence of intermediate
hosts was dependent on the altitude. Telford's measurement of alti-
tude was actually the sum total of climatic factors. The infested
specimens of S. graciosus from Provo Canyon are from an area
where water and vegetation are relatively abundant compared to
dryer areas west of the Wasatch Front such as the Little Sahara
sand dunes. Therefore, the unknown, intermediate host is probably
an arthropod inhabiting cooler, moister habitats.

The habitat of O. scelopori is seemingly the duodenum of the
small intestine. Since this is the main organ of digestion (Villee and
Dether, 1971), this parasite has an advantage in securing nutrients
formed by the digestive processes of the host.

The effects of O. scelopori on the hosts were apparently negli-
gible. Neither S. graciosus nor S. occidentalis appeared to have
structural damage. Although there may have been some competi-
tion between hosts and parasites for the nutrients, the hosts were
adults with fat bodies in the coelum, indicating an excess of nutri-
ents. It is, of course, possible that multiple infestations of this worm
would damage the host.

In both host lizards there were no other helminths harbored. The
possibility exists that O. scelopori has some mechanism which limits
or interferes with other helminths. The literature is inconclusive on
this point, and laboratory experimentation would be necessary to
establish this hypothesis.

A wide distribution is shown by O. scelopori. Telford (1970)
reports this cestode in lizards of Iguanidae and Xantusiidae, for
which families it is seemingly host specific.

Skrjabinoptera phrynosoma. — This species is the best known
stomach worm of reptiles in the southwestern United States. We
have recovered 47 specimens from two species of Sceloporus. Seven
of 11 specimens of S. undulatus and three of 11 specimens of S.
magister harbored this worm. This appears to be the first report of
this parasite in these species.

10 GREAT BASIN NATURALIST Vol. 33, No. 1

A possible nidus distribution exists for Sk. phrynosoma within
the Great Basin and the Upper Colorado Plateau of Utah. Six of
seven lizards, representing both host species, from Arches National
Park were parasitized. In the St. George-Santa Clara area, however,
only one of eight hosts harbored the worm.

In infested stomachs of anesthetized lizards, Sk. phrynosoma
was always located in the first third of the stomach with the head
deeply buried in the stomach lining. The reason for this anterior
attachment is not known. There may be less movement of the
stomach anteriorly, or it may allow the parasite to extend without
moving into the environment of the small intestine where the pH
is quite different.

Externally, lizards with Sk. phrynosoma could not be distin-
guished from those without. Internally, however, the worms caused
great welts in the stomach lining. There were no signs of tissue in-
flammation or disintegration, but infestation was light. In some
horned lizards, reported infestations are as high as 500 worms, with
an average of 50 worms per lizard, and tissue damage is severe
(Babero and Kay, 1967). However, the maximum number of
worms in S. magister was 10, and in S. undulatus eight; the average
for both species was about five worms per host. It seems that Sk.
phrynosoma is not as abundant in Sceloporus as it is in Phrynosoma
platyrhinos. The latter is a notorious ant eater, and since ants are
intermediate hosts (Lee, 1957), we suspect that one or more species
of ants may be involved in the parasitic cycle.

Skrjabinoptera phrynosoma is limited to the family Iguanidae
but is widely scattered through many of its genera (Telford, 1970).
Physiological host specificity exists at the intermediate host level
(Lee, 1957), but apparent preference for certain definitive hosts is
ecologically determined by the dietary specializations of the host
(Telford, 1970).

Physaloptera retusa. — This species was recovered from S. occi-
dentalism S. undulatus, and S. magister, the latter two constituting
new host records. Physaloptera retusa is typically a stomach worm
of S. occidentalis (Telford, 1970), but in both of the new hosts P.
retusa was recovered from the cecum.

In this study, specimens of 5. occidentalis were captured from the
foothills west of Utah Lake and S. undulatus and S. magister were
taken from Arches National Park.

Sceloporus occidentalis was first reported to harbor P. retusa by
Grundmann (1959), who examined seven specimens from Utah.
Since then, Telford (1970) has reported this nematode to be the
most common stomach worm i)arasitizing S. occidentalis in southern
California. Perha[)s it is now justifiable to say that P. retusa is the
most abundant stomach worm of 5". occidentalis in the southwestern
states. Sceloporus magister and 5. undulatus harbored five and ten
specimens respectively of the stomach worm Skrjabinoptera phryno-
soma. It is interesting to note that P. retusa occurs in the cecum of
these species and in the stomach of iS. occidentalis.

March 1973 pearce, tanner: lizard parasites 11

The effects of P. retusa on its Sceloporus hosts appeared negli-
gible. Attachment of worms to host stomachs was similar to that
described for Sk. phrynosoma. Grundmann (1959) thought P. retusa
was host specific for S. occidentalis, but Telford (1970) has shown
it to be in genera as diverse as Crotaphytus, Uta, Callisaurus, Eu-
meces, and Gerrhonotus. He suggested that host specificity of P.
retusa is dependent on ecological factors similar to those in Sk.
phrynosoma, the main factor being the diet of host lizards, which
explains the distribution of this parasitic worm. Telford attributed
the lack of P. retusa in S. magister from southern California to the
absence in their diet of an unknown intermediate host. The presence
of P. retusa in both 5. magister and S. undulatus from Arches Na-
tional Park indicates that the intermediate host is a common item
in these diets.

Thubunaea iguanae. — This spiruroid nematode was found in the
stomachs of two specimens of S. magister. One, from Arches National
Park, had 30 worms; the other, from the Price area, had six. This
is a sizable range extension for this nematode. Telford (1970)
originally reported its occurrence in S. magister.

The effects of T. iguanae on its host appeared negligible. The
head of the worm was imbedded, but the attachment did not appear
as strong as that reported above for Sk. phrynosoma and P. retusa.
Both hosts were adults, apparently in good health. The effects of
T. iguanae on other species of helminths could not be determined;
however, there is a possibility that Sk. phrynosoma may have af-
fected the distribution of T. iguanae. These helminths were not
recovered from the same host. As noted above, P. retusa was not
recovered from stomachs containing Sk. phrynosoma. These two
facts support a theory that Sk. phrynosoma affects the presence
and/or distribution of other nematodes within the host.

This nematode is harbored by several species from each of the
following families of reptiles: Iguanidae, Xantusiidae, Teiidae, and
Gekkonidae (Telford, 1970).

Pharyngodon giganticus. — This nematode belongs to the order
Oxyuroidea, and was found in the cecum. It was recovered from
two specimens of S. undulatus from Zion National Park, as well as
from four specimens of S. occidentalis- — three from west of Utah
Lake and one from St. George, Utah. Sceloporus undulatus repre-
sents a new host record for P. giganticus.

The effects of P. giganticus upon hosts were apparently negli-
gible. This helminth lives in the fecal material of the host's cecum.
The host-worm relationship appears to be commensal. There is the
possibility that under heavy infestations this worm may be injurious
to the host; however, such was not noted by us.

The effect of P. giganticus on other oxyuroid helminths is a
possibility, since there were no other oxyuroid worms sharing the
cecum with it; however, there were no other oxyuroid species found
in specimens of S. undulatus or S. occidentalis.

12 GREAT BASIN NATURALIST Vol. 33, No. 1

Oxyuroid nematodes show a greater degree of host specificity
than do the spiruroids previously mentioned. Telford (1970) reports
P. giganticus to be restricted to Sceloporus, with the exception of two
widely separated island populations of Uta, which he suggests may
be a relict distribution.

Cyrtosomum penneri, C. heynemani, and C. readi. — These three
oxyuroids are considered together because of the many similarities
in host, habitat, and life style. The Cyrtosomum species were usually
found in association with either one or both of the other Cyrtosomum
species. All three species were recovered from specimens of Scelopo-
rus graciosus taken from Provo Canyon and the Little Sahara sand
dunes. Cyrtosomum penneri and C. readi were recovered from one
specimen of S. magister taken from Arches National Park and also
from one specimen from the Price area. This study reports 5.
graciosus as a new host for C. readi. Gambino (1957a, 1957b)
describes and discusses the morphology of these nematodes.

All three Crytosomum nematodes infested the ceca of their hosts.
In only two specimens of S. magister did these worms occur outside
of the cecum; since both lizards were preserved specimens, this
occurrence could be explained by a migration at the time of preser-
vation. Dissections and observations of infested hosts revealed that
Cyrtosomum worms probably live exclusively in the cecum. Ecologi-
cal barriers such as peristaltic movement, scarcity of food, and re-
striction in space may be limiting factors. Physiological barriers in
the large intestine may limit worm distribution past the cecum,
where the colon removes most of the water and concentrates meta-
bolic wastes. These would cause osmotic and pH gradients that
might impair the survival of parasites.

The effects on the host by Cyrtosomum nematodes appeared
negligible. At times, 300 worms occurred in a single host. Such
heavy infestations of Cyrtosomum worms often caused the walls of
the cecum to move as the worms pushed against it. This may have
caused irritation; however, there was no tissue damage noted.
Worms were seemingly feeding on the fecal matter, and thus were
not competing with the host for food but living in a commensal
relationship.

The presence or absence of other nematodes showed no effect
on the presence or abundance of Cyrtosomum worms. Cyrtosomum
penneri was the most commonly occurring Cyrtosomum. It shared
all the infested lizards harboring either C. readi or C. heynemani
and was found in two 5. magister lizards by itself. Cyrtosomum
heynemani was the only species which did not occur in S. magister.
This may not be significant, because some experts are not certain
that C. readi is a distinct species from C. heynemani (Babero and
Kay, 1967).

Cyrtosomum are host specific to the family Iguanidae, infesting
many genera including Dipsosaurus, Crotaphytus, Sauromalus,
Phrynosoma, Uta, Callisaurus, and Sceloporus (Telford, 1970).

March 1973 pearce, tanner: lizard parasites 13

Conclusions and Summary

Dissections of 55 lizards representing four species of Sceloporus
resulted in the recovery of eight species of helminths. These in-
clude: a cestode, Oochoristica scelopori; three species of spiruroid
nematodes, Skrjabinoptera phrynosoma, Physaloptera retiisa, and
Thubunaea iguanae; and four species of oxyuroid nematodes,
Pharyngodon giganticus, Cyrtosomum penneri, C. heynemani, and
C. readi. Eight new host records were reported, and the geographic
distribution was extended for most of the parasitic species.

Host specificity was not restrictive enough to indicate phylo-
genetic relations between host species. The occurrence of helminths
in Sceloporus lizards apparently was correlated with the distribution
of intermediate hosts and the diets of definitive hosts. The effects
of helminths on their Sceloporus hosts were apparently slight; how-
ever, infestations of cestodes or spiruroid nematodes was light. The
possibility exists that lizards with high infestations have a lower
survival rate. Oxyuroid nematodes did occur in great numbers, but
in all cases the hosts appeared healthy.

The effects of helminths on other helminths in the same host
were indicated. Stomachs were parasitized by a single species of
nematode even when other helminths, usually inhabitants of the
stomach, were recovered from the same lizard. Skrjabinoptera
phrynosoma was apparently dominant to both Physaloptera retusa
and Thubunaea iguanae.

There were some differences in number and types of parasites
found in the same host from different collecting sites. Specimens
from the St. George-Santa Clara area were almost free of helminths,
whereas those from Arches National Park were all infested. This
nested distribution evidently resulted from different diets and the
presence or absence of intermediate hosts in the locality.

Appendix

The literature available on reptilian parasitology from the
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Inf. Dis. 74:48-67.
. 1944b. Saurian malarial parasites of United States and Mexico. J.

Inf. Dis. 74:68-79.
Vincent, I. 1948. Studies on the endoparasites of the Texas horned lizards

Phrynosoma cornutum. (Harlan). Proc. Texas Acad. Sci. 30:250-252.
Walton, A. 1927. A revision of the nematodes of the Leidy collections. Proc.

Nat. Acad. Sci. Philadelphia 79:49-163.
Walton, A. C. 1941. Distribution of the genus Thelandros (Nematoda:

Oxyuroidea). Proc. Helminth. Soc. Washington 8:15-18.
WiECzoROWSKi, Elsie. 1939. Parasite lesions in turtles. J. Parasitol. 25:395-399.
WiTENBERG, G. 1934. Studies on the cestode genus Mesocestoides. Arch. Zool.

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Wood, W. F. 1935. Some observations on the intestinal protozoa of California

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Literature Cited

1. Babero, B. B., and F. R. Kay. 1967. Parasites of horned toads {Phryno-

soma spp.), with records from Nevada. J. Parasitol. 53:168-175.

2. Gambino, J. J. 1957a. Cyrtosomum readi n. sp. and Cyrtosomum heyne-

mani n. sp. (Oxyuroidea: Atractidae) two new pinworms of iguanids.
J. Parasitol. 44:439-445.

3. . 1957b. Cyrtosomum penneri n. sp. (Oxyuroidea: Atractidae). J.

Parasitol. 43:76-80.

4. Grundmann, a. W. 1957. A preliminary study of the intestinal parasites

of some Utah lizards. Proc. Utah Acad. Sci. 34:147-148.

5. . 1959. Parasites recovered from six species of Utah lizards. J.

Parasitol. 45:394.

6. Harwood, p. D. 1932. The helminths parasitic in the amphibia and

reptilia of Houston, Texas, and vicinity. Proc. U.S. Nat. Mus. 81:1-67.

7. Hyman, L. H. 1951. The invertebrates: Acanthocephala, Aschelminthes,

and Entoprocta. Vol. Ill, 1st ed. McGraw-Hill Book Co., New York.

8. Lee, S. H. 1957. The life cycle of Skrjabinoptera phrynosoma (Ortlepp)

Schulz, 1927 (Nematoda: Spiruroidea), a gastric nematode of Texas
horned toads, Phrynosoma cornutum. J. Parasitol. 43:66-75.

9. Lucker, J. T. 1951. Some new Thelandros (Nematoda: Oxyuridae) from

the island night lizard, Xantusia riversiana reticulata Smith, from San
Clemente Island, California. J. Parasitol. 37(suppl.): 14-15.

10. Meyer, M. C, and L. R. Penner. 1962. Laboratory essentials of parasi-

tology. Wm. C. Brown Co., Dubuque, Iowa.

11. Morgan, B. B. 1942. The nematode genus Skrjabinoptera Schulz, 1927.

Lloydia 5:314-319.

18 GREAT BASIN NATURALIST Vol. 33, No. 1

12. Ortlepp, R. 1922. The nematode genus Physaloptera Rud. Proc. Zool.

Soc. London 72:999-1107.

13. Read, C. P., and Y. U. Amrein. 1953. North American nematodes of the

genus Pharyngodon Diesing (Oxyuridae). J. Parasitol. 39:365-370.

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16. Telford, S. R., Jr. 1965. A new species of Thubunaea (Nematode:

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MISCELLANEOUS CHROMOSOME COUNTS
OF WESTERN AMERICAN PLANTS— IP

James L. Reveal- and Eloise L. Styer*

Abstract. — Chromosome numbers are reported for 19 species of western
American plants. Reports for previously uncounted species include Nitrophila
mohavensis Munz and Roos, n=18; Eriogonum contiguum (Reveal) Reveal, n =
16; Mentzelia leucophylla Brandeg., n=18, and M. torreyi A. Gray, n = 12;
Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg., n = 8; Astragalus
beatleyae Barneby, n = ll; Angelica kingii (S. Wats.) Coult. and Rose, n = 22;
Gilia nyensis Reveal, n==9; and Arnica parryi A. Gray var. sonnei (Greene)
Cronq. in Ferris n==19. New ploidy levels are reported in Oxystylis lutea Torr.
and Frem., n=10; Lupinus aridus Dougl. ex Lindl., n = 12; and Castilleja mar-
tinii Abrams var. clokeyi (Pennell) N. H. Holmgren, n = 12. Seven additional
counts are given which confirm those published by others. These are Opuntia
chlorotica Engelm. and Bigel., n = ll; Eriogonum inf latum Torr. and Frem. var.
inf latum, n = 16, E. trichopes Torr., n = 16, and E. bifurcatum Reveal, n = 20;
Stanleya pinnata (Pursh) Britt. var. inyoensis (Munz and Roos) Reveal, n = 28;
Lupinus argenteus Pursh, « = 24; and Castilleja viscidula A. Gray, n = 12.

In this series of papers, of which this is the second, chromosome
counts of randomly gathered western American plants will be re-
ported as part of the Intermountain Flora Project (headed by Arthur
Cronquist of the New York Botanical Garden and aided by Arthur
H. Holmgren, Noel H. Holmgren, and James L. Reveal) and the
proposed Southwest Flora Project (headed by Noel H. Holmgren and
James L. Reveal). These counts will be presented from time to
time by Reveal and various others of his colleagues or students and
will be based on materials gathered by him or others. The present
paper reports on some of the miscellaneous plants gathered in south-
ern Nevada in 1970 and 1971.

Flower buds were collected in developmental series from plants
growing in their native habitats. The buds were fixed in ethanol
and glacial acetic acid (3:1) and refrigerated; the buds were not
transferred to 70 percent ethanol after 24 hours, as is the usual tech-
nique. Anthers were squashed in acetocarmine and illustrations
drawn using a camera lucida mounted on a Wild M20 research
microscope. Most of the counts reported here were made by Reveal.
Voucher specimens are deposited in the herbarium at the Nevada
Test Site, with duplicates deposited in several major herbaria.

Chromosome numbers of 19 species of western American plants
are reported here, nine for the first time, with three counts of new
ploidy levels. One count is given for a species already reported in
the literature but not figured, and six counts confirm those made
previously by others.

Opuntia chlorotica Engelm. and Bigel. 72=11. Figs. 1, 2. This
count confirms those made by others (Stockwell, 1935; Pinkava and

'The first paper in this series was published in the Southwestern Naturalist.

^Department of Botany, University of Maryland, College Park 20742; National Museum of
Natural History, Smithsonian Institution, Washington, D. C. 20560.

^Department of Botany, University of Maryland, College Park 20742.

20 GREAT BASIN NATURALIST Vol. 33, No. 1

McLeod, 1971). The voucher is Beatley and Reveal 11231, west side
of Spring Mountains [Charleston Mountains], Clark Co., Nevada,
17 June 1970.

Nitrophila mohavensis Munz and Roos. « = 18. Figs. 3, 4. This
rare species is known only from a small site in extreme eastern Inyo
Co., California, where it is a locally dominant species. It would
appear that A^. mohavensis is a tetraploid, since Covas and Schnack
(1947) have reported a count of 72 = 9 for the South American species
A^. australis Chod. and Wile, although a second North American
species, A^. occidentalis (Nutt.) Moq., should be counted to confirm
this. Nevertheless, it seems possible to state that the base number
for Nitrophila is t = 9. The voucher collection is Reveal 2291, Amar-
gosa Desert, southern Ash Meadows, along the road between Ash
Meadows and Death Valley Junction on the saline flats of Carson
Slough, Inyo Co., California, 5 May 1971.

Eriogonum inf latum Torr. and Frem. var. inf latum. «=16. Fig.
5. This count confirms those made by Stone and Raven (1958) and
Reveal (1965). The voucher is Reveal 2290, Stewart Valley, 2 miles
north of Nevada Highway 52 on road to Ash Meadows, Nye Co.,
Nevada, 5 May 1971.

Eriogonum contiguum (Reveal) Reveal. n=\6. Figs. 6, 7.
When this plant was counted and found to equal that of E. inflatum
and E. trichopes, and when fieldwork showed no hybridization be-
tween the three, it was decided to elevate E. inflatum var. contiguum
to the species rank (Reveal, 1972). The voucher of this new count
is Reveal 2288, Stewart Valley, 2 miles north of Nevada Highway
52 on road to Ash Meadows, Nye Co., Nevada, 5 May 1971.

Eriogonum trichopes Torr. n=16. Figs. 8, 9. This count con-
firms that made previously (Reveal, 1965). The voucher is Reveal
2289, Stewart Valley, 2 miles north of Nevada Highway 52 on road
to Ash Meadows, Nye Co., Nevada, 5 May 1971.

Figs. 1, 2: Opuntia chlorotica, ti=\\ — metaphase I. Figs. 3, 4: Nitrophilia
mohavensis, n=\S — diakinesis. Fig. 5: Eriogonum inflatum var. inflatum. n = 16
— telophase II. Figs. 6, 7: Eriogonum contiguum. n = \6 — anaphase I, Fig. 6;
metaphase I, Fig. 7. Figs. 8, 9: Eriogonum trichopes, « = 16 — diakinesis. Fig. 8;
anaphase I, Fig. 9. Fig. 10: Eriogonum bifurcatum. n^20 — diakinesis. Figs. 11,
12: Mentzelia leucophylla, n=\S — diakinesis. Figs. 13, 14: Mentzelia torreyi,
„ = 12— telophase II, Fig. 13; anaphase II, Fig. 14. Figs. 15, 16, 17, 18: Oxy-
stylis lutea, n=10 — metaphase I. Figs. 15, 17, 18; diakinesis, Fig. 16. Figs. 19,
20: Stanleya pinnata var. inyoensis, n^28 — diakinesis. Fig. 19; anaphase I, Fig.
20. Figs. 21, 22, 23: Prunus virginiana var. melanocarpa, n = 8 — metaphase I,
Figs. 21, 22; anaphase I, Fig. 23. Figs. 24, 25: Astragalus beatleyae. n = ll —
anaphase I. Figs. 26, 27: Lupinus argenteus, n = 24 — metaphase I, Fig. 26;
anaphase I, Fig. 27. Figs. 28, 29: Lupinus aridus. n=\2 — metaphase I, Fig.
28; metaphase II. Fig. 29. Figs. 30, 31: Angelica kingii. n = 22 — anaphase
I, Fig. 30; metaphase I, Fig. 31. Fig. 32: Gilia nyensis, n = 9 — anaphase I.
Figs. 33, 34: Casiilleja viscidula, n=\2 — anaphase I, Fig. 33; telophase II,
Fig. 34. Figs. 35, 36: Castilleja martinii var. clokeyi, n = l2 — anaphase I. Figs.
37, 38: Arnica parryi var. sonnei, n=19 — anaphase I.

March 1973 reveal, styer: plant chromosomes

22 GREAT BASIN NATURALIST Vol. 33, No. 1

Eriogonum bifurcatum Reveal. « = 20. Fig. 10. When this
species was proposed (Reveal, 1971), the chromosome number was
noted but a figure not provided. The voucher of this count is Reveal
2283, Pahrump Valley, 1 mile east of the California-Nevada state
line just south of Nevada Highway 52, Nye Co., Nevada, 1 3 June
1970.

M entzelia leucophylla Brandeg. n=18. Figs. 11, 12. As current-
ly understood, this is a rather rare species of eastern California and
adjacent southern Nevada. The count indicates the species to be
a polyploid (most likely a tetraploid), but one would like more
counts of the species from throughout its range to confirm this point.
The voucher is from the type location in Ash Meadows, Nye Co.,
Nevada, based on Reveal 2247, 11 June 1970.

Mentzelia torreyi A. Gray. n—\2. Figs. 13, 14. This count is
most unusual, since this number is unknown (at least to us) for the
genus. A more reasonable number would be n=ll, but our report
was confirmed by a number of smears. Bud material was obtained
by Reveal and Beatley on 14 June 1970 from the same place Reveal
made a collection earlier. The voucher is Reveal 1980, Kawich Val-
ley, Nye Co., Nevada, 31 August 1968.

Oxystylis lutea Torr. and Frem. n=10. Figs. 15, 16, 17, 18.
Raven, Kyhos, and Hill (1965) recently reported the number of
2a2 = 20 pairs for this species, but our counts would indicate their
plants to have been tetraploids, since our count is just half theirs.
Based on these data, the base number for the genus would be x=10.
Our voucher is Reveal 2248, west of Ash Meadows, along a dirt road
southwest of the California-Nevada state line from California High-
way 127 toward the abandoned Tonopah and Tidewater Railroad
tracks, Inyo Co., California, 11 June 1970.

Stanley a pinnata (Pursh) Britt. var. inyoensis (Munz and Roos)
Reveal, stat. nov., based on S. pinnata ssp. inyoensis Munz and Roos,
Aliso 3:115. 1965 « = 28. Figs. 19, 20. Our counts are in agree-
ment with those recently obtained from root tips by Rollins and
Riidenburg (1971). The voucher is Reveal 2277, Stewart Valley,
Inyo Co., California, 12 June 1970.

Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg. « = 8.
Figs. 21, 22, 23. This represents a new report, since no number for
var. melanocarpa has been previously published. It is also a new
count for the species, since the only report of var. virginiana we
know of is n = 16 (Sax, 1931), and such a count may raise some
questions as to the relationship between the western American en-
tities, var. melanocarpa and var. demissa (Nutt.) Sarg., and the
eastern chokecherry, var. virginiana. More data are needed and
hopefully can be obtained during the next few years. Our voucher is
Beatley and Reveal 11208, Upper Eden Creek Canyon, east slope of
the Kawich Range, Nye Co., Nevada, 15 June 1970.

Astragalus beatleyae Bameby. n=\\. Figs. 24, 25. This new
count joins those of two other species in subsection Aridi of the

March 1973 reveal, styer: plant chromosomes 23

section Inflati; all are n—\l (Barneby, 1964), which is the most
common number in the North American species of the genus (see
bolkhovskikh et al., 1969). Our voucher of this recently proposed
species (Barneby, 1970) is Beatley and Reveal 10907, flatrock area
-Z.d, miles north of Pahute Mesa Road on the Plateau Road, Pahute
Mesa, Nye Co., Nevada, 4 June 1970.

Lupinus argenteus Pursh. n = 24. Figs. 26, 27. This complex
species is found throughout much of the western United States and
is variously divided into species, subspecies, or varieties by several
authors. C. P. Smith (1944) most likely called this plant L. corym-
bosus Heller. Later, these plants were called L. alpestris A. Nels.
(Clokey, 1951) or L. argenteus var. tenellus (Dougl. ex G. Don)
D. Dunn (Dunn, 1956). More recently, these plants have been re-
ferred to L. argenteus var. stenophyllus (Nutt. ex Rydb.) R. J.
Davis (Beatley, 1969). We are still not clear what name should be
applied to the collections from southern Nevada; L. argenteus is
certainly the oldest name, although L. alpestris would have priority
in our vicinity. The voucher is Beatley and Reveal 10909, south rim
of Pahute Mesa, Nye Co., Nevada, 4 June 1970.

Lupinus aridus Dougl. ex Lindl. /2=12. Figs. 28, 29. This
species has been variously defined in recent years. C. P. Smith
(1944) and Dunn (1956) have considered this form a distinct spe-
cies, while Detling (1951), who monographed the group, referred
L. aridus to a subspecific rank under L. lepidus Dougl. ex Lindl.;
this latter treatment was followed by that of Hitchcock (1961), who
used the varietal rank instead. To date, all entities referred to this
species complex have had a single chromosome number, « = 24.
Among the related species are L. lyallii A. Gray [or L. lepidus var.
lobbii (A. Gray) C. L. Hitchc], « = 24 (Phillips, 1957); L. caespito-
sus Nutt. ex Torr. and Gray [or L. lepidus var. utahensis (S. Wats.)
C. L. Hitchc], ^2 = 24 (Heiser, 1963); and L. lepidus itself, with
72 = 24 (Phillips, 1957). Dunn has seen our voucher and other
specimens from the same site and has identified our material as L.
aridus. However, according to Detling's monograph, our material
would be far out of the known range of the L. lepidus complex, the
nearest confirmed site being in the high Sierra Nevada of California
(L. lobbii A. Gray and L. lyallii) or in southeastern Oregon (L.
aridus). One collection from northern Nye Co., Nevada, was tenta-
tively assigned to L. aridus by Detling, but he was uncertain as to
the exact identity of the collection. Based on the unusual chromo-
some number, geographical disjunction, and the observable morpho-
logical differences seen by both Beatley and Reveal, it would seem
the plants from southern Nye Co., Nevada, may represent a new
taxon. Our voucher is Beatley and Reveal 11138, southern Kawich
Valley just below Gold Flat, Nye Co., Nevada, 14 June 1970.

Angelica kingii (S. Wats.) Coult. and Rose. n = 22. Figs. 30, 31.
This represents a new count for the species. Our voucher is Beatley
and Reveal 11284, Cold Creek, eastern slope of Spring Mountains
[Charleston Mountains], Clark Co., Nevada, 17 June 1970.

24 GREAT BASIN NATURALIST Vol. 33, No. 1

Gilia nyensis Keweal. n = 9. Fig. 32. A new count. This recently
proposed species (Reveal, 1969), found in sandy places in southern
Nye Co., Nevada, is related to G. hutchinsifolia Rydb., which also is
reported to be /2 = 9 (Munz and Keck, 1959). The voucher is Beatley
and Reveal 11147, Kawich Valley, Nye Co., Nevada, 14 June 1970.

Castilleja viscidula A. Gray. ^2=12. Figs. 33, 34. Our count
agrees with that recently published by Holmgren (1971). Our
voucher is the southernmost population of the species: Beatley and
Reveal 11068, Kawich Range, northwest of Kawich Peak, Nye Co.,
Nevada, 6 June 1970.

Castilleja martinii Abrams var. clokeyi (Pennell) N. H. Holm-
gren. n = 12. Figs. 35, 36. Our count represents a new ploidy level
for var. clokeyi. Holmgren (1971) reported n = 24 for plants
gathered on the east slope of the Spring Mountains (Holmgren and
Reveal 2988), while our count, w^hich was obtained from specimens
gathered on the west side of the same range, indicates a diploid situ-
ation. This polyploid condition is to be expected, since Holmgren
reports /2=12, 24, and 36 for var. martinii. The voucher is Beatley
and Reveal 11282, Trough Springs, north end of Spring Mountains,
Clark Co., Nevada, 17 June 1970.

Arnica parryi A. Gray var. sonnei (Greene) Cronq. in Ferris.
72=19. Figs. 37, 38. Our count is the first reported for this taxon
and represents a new ploidy level in the species. Ornduff et al.
(1963) have reported var. parryi to be n = csi. 36, so it is possible
that the typical variant might be n = 38, or twice the value found
for var. sonnei. Our voucher is Beatley and Reveal 10975, near the
stream at Longstreets Ranch, west of Kawich Peak, Kawich Range,
Nye Co., Nevada, 6 June 1970.

Laboratory work on this project was supported by NSF Grant
GR-22645 to the senior author. Fieldwork in Nevada was conducted
under Contract No. AT (04-1) Gen- 12 between the University of
California and the Division of Biology and Medicine, United States
Atomic Energy Commission, under the direction of Janice C. Beatley,
research ecologist. We are grateful to her for her aid with this work.

Literature Cited

Barneby, R. C. 1964. Atlas of North American Astragalus. Mem. New York
Bot. Gard. 13:1-1188.

. 1970. A new Astragalus from Nye County, Nevada. Aliso 7:161-163.

Beatley, J. C. 1969. Vascular plants of the Nevada Test Site, Nellis Air Force
Range, and Ash Meadows. UCLA 12-705. Laboratory of Nuclear Medi-
cine and Radiation Biology, University of California, Los Angeles.

BoLKHovsKiKH, Z., V. Grif, T. Matvejeva. AND O. Zakharyeva. 1969. Chromo-
some numbers of flowering plants. Academy of Sciences of the USSR, Lenin-
grad.

Clokey, L W. 1951. Flora of the Charleston Mountains, Clark County, Nevada.
Univ. Calif. Publ. Bot. 24:1-274.

Covas, G., and B. Schnack. 1947. Estudios cariologicos en Antofitas. Revista
Argent. Agron. 14:224-231.

DetlinGj L. E. 1951. The caespitose lupines of western North America.
Amer. Midi. Nat. 45:474-499.

March 1973 reveal, styer: plant chromosomes 25

Dunn, D. B. 1956. Leguminosae of Nevada, II — Lupinus. Contr. toward Flora
Nevada 39:1-64.

Heiser, C. B. 1963. Numeracion cromosomica de plantas ecuatorianas. Cienc.
y Naturaleza 6:2-6.

Hitchcock, C. L. 1961. Leguminosae. In Hitchcock, C. L., et al., Vascular
plants of the Pacific Northwest. Univ. Wash. Publ. Biol. 17(3): 194-377.

Holmgren, N. H. 1971. A taxonomic revision of the Castilleja viscidula group.
Mem. New York Bot. Gard. 21 (4) : 1-63.

MuNz, P. A., AND D. D. Keck. 1959. A California flora. University of Cali-
fornia Press, Berkeley.

Ornduff, R., p. H. Raven, D. W. Kyhos, and A. R. Kruckeberg. 1963.
Chromosome numbers in Compositae. III. Senecioneae. Amer. J. Bot. 50:
131-139.

Phillips, L. L. 1957. Chromosome numbers in Lupinus. Madrono 14:30-36.

PiNKAVA, D. J., AND M. G. McLeod. 1971. Chromosome numbers in some cacti
of western North America. Brittonia 23:171-176.

Raven, P. H., D. W. Kyhos, and A. J. Hill. 1965. Chromosome numbers of
Spermatophytes, mostly Califomian. Aliso 6:105-113.

Reveal, J. L. 1965. Eriogonum. In Documented chromosome numbers of
plants. Madrono 18:124.

. 1969. New species in Eriogonum and Gilia from southern Nevada.

Bull. Torrey Bot. Club 96:476-484.

. 1971. A new annual Eriogonum (Polygonaceae) from southern Ne-
vada and adjacent California. Aliso 7:357-360.

1972. Descriptions of new species and combinations in Eriogonum

(Polygonaceae). Phytologia 23:163-178.

Rollins, R. C, and L. Rudenberg. 1971. Chromosome numbers of Cruciferae.
II. Contr. Gray Herb. 201:117-133.

Sax, K. 1931. The origin and relationships of the Pomoideae. J. Arnold Arbort.
12:3-22.

Smith, C. P. 1944. Lupinus. In Abrams, L., Illustrated flora of the Pacific
States. Vol. II. Stanford University Press, Stanford. California.

Stockwell, p. 1935. Chromosome numbers of some of the Cactaceae. Bot.
Gaz. 96:565-570.

Stone, D. E., and P. H. Raven. 1958. Eriogonum. In Documented chromo-
some numbers of plants. Madroiio 14:237.

INCIDENCE OF SPOTTED FEVER IN WOOD 1 ICKS
OF UTAH RECREATIONAL SITES'

C. Selby Herrin^

Abstract. — In 1964, 135 male and 223 female Dermaccntor andersoni Stiles
were collected at 48 national forest recreation areas in Utah. Using guinea pigs
immunological tests were performed on each of 62 samples to determine the
presence of the Rocky Mountain spotted fever rickettsia. Thirteen of the samples
(21 percent) were positive. These 13 samples contained 35 males and 34 females
yielding an infection rate of at least 3.6 percent, assuming one infected tick was in
each sample.

The Rocky Mountain wood tick, Dermacentor andersoni Stiles,
has been recognized as the principal vector of Rocky Mountain spot-
ted fever (RMSF) in the Rocky Mountain states since shortly after
the turn of the century. Much human suffering and many deaths
have resulted from this disease in Utah as well as throughout the
western United States. The presence of D. andersoni in recreational
sites of the foothills, canyons, and mountains of the West offers a
potential threat to the health of man. Expanding human population
and increasing use of recreational facilities enhances this potential.

A previous paper (Herrin, 1968) dealt with the prevalence of
D. andersoni adults in national forest recreational sites of Utah.
This research note reports the incidence of Rickettsia rickettsi (Wol-
bach) in adult ticks collected from these recreational areas.

Using a white flannel drag, 358 adult D. andersoni (135 males
and 223 females) were collected from 48 recreational sites during
the spring and summer of 1964. Ticks from 54 collections were
divided into 62 samples, rinsed several times in sterile physiological
saline, and preserved in vials of sterile, nonfat skim milk at below
-30 C. Subsequently, the ticks were thawed, removed from the milk,
triturated, and diluted with 10 ml of sterile, nonfat skim milk. One
guinea pig per sami)le was inoculated with 1 ml of supernatant.
Guinea pigs were observed for scrotal reactions, and rectal tempera-
tures were taken every other day for 28 days. If the guinea pig
died, a second was subsequently inoculated with the original super-
natant. After 28 days, blood was taken and serum was extracted
from the guinea pigs. Four complement fixation (CF) screen tests
were performed on each sample, followed by an additional CF test to
determine the antibody titer of positive samples. As a final test, all
guinea pigs were inoculated with a lethal dose of virulent R. rickettsi
to determine if they were immune to RMSF. In this test, one guinea
pig per sample was challenged, and immunity was judged solely on
survival.

Abnormal temperatures in guinea pigs resulted after injections of
triturated tick material from 18 samples. Seven showed abnormally

>This paper is taken from a master's thesis submitted to the Department of Zoology and the
Graduate School of Brigham Young University (1966).

^Center for Health and Environmental Studies, Brigham Young University, Provo, Utah 84001.

March 1973

HERRIN: SPOTTED FEVER

Table 1. Collections of D. andersoni adults determined to be positive for
R. rickettsi.

Number

Complement

National forest

Collection

and sex

fixation

Immunity

Recreational site

date

in sample

titer

test*

Cache

Friendship and

Spring

July

2M, 2F

Fish Lake

Bowery

June

2M, 2F

City Creek

2 June

128

—

Monrovian Park

June

128

12F

Ponderosa

June

—

lOF

128

Manti-Lasal

Manti Community

July

128

lOF

—

lOF

—

Uinta

Altamont

June

2M, 4F

Aspen Grove

June

2M, 5F

Mutual Dell

June

2M, 3F

128

Silver Lake Flat

June

IM, 4F

Timpooneke

June

128

IIF

Whiskey Springs

June

2M, 3F

128

Wasatch

Sunset

July

128

* Legend: + = immune; — = susceptible.

high temperatures immediately and throughout the observation
period, whereas 1 1 had an increase in temperature after several
days. None demonstrated scrotal reactions or necrosis. Eight guinea
pigs died within 3 to 14 days after inoculation. Thirteen of 62
serum samples (21 percent) yielded positive CF tests (see Table 1).
The titers of these samples ranged from 1:32 to 1:128, whereas con-
trols gave titers of 1:512. Eleven of 62 guinea pigs challenged with
virulent spotted fever organisms survived, indicating immunity to
RMSF. These 11 correspond with 11 of the 13 serum samples yield-
ing positive CF results. In two cases of positive CF tests, guinea
pigs were susceptible to the challenge. This close correlation be-
tween results of the CF tests and the immunity tests suggests that
11, and probably 13, tick samples contained R. rickettsi. Three of
the CF positive guinea pigs showed slightly abnormal temperatures.
The remaining 15, and possibly 18, instances of abnormal tempera-
tures are not attributable to spotted fever, since there was no correla-
tion between samples that showed positive CF and immunity tests.
Of the eight guinea pigs that died, two were from samples deemed
positive by CF and immunity tests. Observations made at necropsy
indicated that death was probably caused by bacterial infection.

28 GREAT BASIN NATURALIST Vol. 33, No. 1

The 13 samples of ticks positive for spotted fever represented 69
ticks (35 males and 34 females). If each positive sample yielded
one tick infected with spotted fever, then 3.6 percent of all ticks col-
lected were infected. Infected ticks were from sites in the northern
half of Utah near human population centers (Provo, Salt Lake City,
Ogden, and Logan). Infected ticks were collected from early June
to late July and from every elevation range at which collections
were made (6000 to 8800 ft). Further studies relative to incidence
and virulence of R. rickettsi in wood ticks would help to determine
the enzootic status of RMSF in Utah. Such studies should include
identification of R. rickettsi by fluorescent antibody staining and
recovery of rickettsiae in egg cultures prior to immunological tests
with guinea pigs.

Literature Cited

Herrin, C. S. 1968. Dermacentor Andersoni in national forest recreation sites
of Utah. Great Basin Nat. 28(1): 30-43.

NOTES ON THE NESTING BEHAVIOR OF
STENIOLIA ELEGANS (HYMENOPTERA: SPHECIDAE)

Howard E. Evans ^

Abstract. — The nesting behavior of a small aggregation of the digger
wasp Steniolia elegans near Fort Collins, Colorado, is described. The prey was
found to consist primarily of bee flies (Bombyliidae), the egg being laid erect on
the first fly placed in the cell. Nests were shallow, unicellular, and were closed
at all times when the female was away.

Several years ago Evans and Gillaspy (1964) reviewed what is
known of the nesting behavior of digger wasps of the genus Steniolia
(tribe Bembicini). One of the more common and widely distributed
species, elegans Parker, was largely omitted from this review, since
little was known concerning it. The present report is based on a
two-day study of a nesting aggregation of this species just west of
the city of Fort Collins, Colorado.

This aggregation was discovered on 4 July 1971 on the slag pile
of an abandoned mine on a south-facing slope. There were an esti-
mated 50 individuals of each sex. Males were most active during
the morning (8:30 a.m. -12: 30 p.m.), flying close to the ground in
irregular patterns and landing here and there with their legs out-
stretched and their antennae extended rigidly forward. Now and
then they hovered, with their middle legs extended, over females
occupied at their nests, and on several occasions attempted copula-
tions were observed.

The nests of the females were located in sloping, moderately fri-
able but very stony soil. During the morning (9:30-11:30 a.m.)
each female reopened her nest, first hovering over the entrance
briefly and then digging through the closure, remaining inside for
20-60 seconds, then emerging and making a fresh closure. Then
she hovered over the nest, flew off, and returned one or more times
to hover again over the nest. These were evidently "inspection trips"
serving to inform the wasp of the needs of the larva and perhaps to
reinforce her memory of the nest site.

Within a few minutes to an hour or two following the inspection,
each female returned with her first prey, carried with the middle
legs in the usual manner of Bembicini. Prey-laden females pro-
duced a loud whine and often hovered briefly over the nest before
digging through the closure. They remained in the nest very brief-
ly (usually less than a minute) before coming out and restoring the
closure. The prey consisted mostly of bee flies, as is common in the
genus. The following species, in the numbers indicated, were taken
from nests or from provisioning females (determinations by L. V.
Knutson) :

Bombyliidae

Anthrax irroratus Say 1

Poecilanthrax signatipennis (Cole) 5

'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138.

30 GREAT BASIN NATURALIST Vol. 33, No. 1

Systoechus vulgaris Loew 4

Villa sinuosa Wiedemann 2

Villa spp. 8
Asilidae

Holopogon atripennis Back 1
Syrphidae

Eupeodes volucris Osten Sacken 1

Microdon coarctatus Loew 1

Volucella sp. 1

Provisioning was found to be fully progressive, the number of
flies brought in per day being determined by the size of the larva.
Nests dug out in the morning, before provisioning began, contained
only the remains of flies.

Several females were seen starting new nests in the afternoon,
after having made the final closure of their previous nests. Much
use was made of the mandibles in breaking through the soil and in
dragging pebbles from the burrow and depositing them on the
mound. From time to time, the female backed out of the entrance
scraping soil, then remained on the mound for a few seconds, turn-
ing to one side or the other while still scraping soil. As the burrow
deepened, the wasp appeared less frequently (at 5- to 10-minute in-
tervals), each time she appeared taking either a brief flight in a
small loop or a much larger loop with a diameter of 3-6 meters. I
observed no leveling movements following completion of the nest,
and the mound of earth at the entrance was left intact, measuring
about 3 cm wide by 4 cm long and 0.5-1.0 cm deep. Several hours
were required for completion of a nest.

When the nest was finished, each female made a thick closure
from the outside by digging into that portion of the mound close to
the entrance. Thus, newly completed nests could be recognized by
the fresh mound which had been dug away on one side, facing the
covered entrance. The egg was found to be laid erect on the side
of the initial fly placed in the cell, in the common manner of
Bembicini.

Six nests were excavated, and all were found to be unicellular —
a simple, oblique burrow leading to a terminal cell 4.5-9 cm deep
(mean 6.6 cm). Some burrows were straight, while others were
curved in various ways to avoid stones. Burrow length varied from
7 to 17 cm (mean 11 cm). None of the nests excavated aj)peared to
be parasitized.

It was evident that wasps did not spend nights or rainy periods
inside their nests, and it is known that this species forms clusters
on vegetation in the manner of other Stcniolia species (Evans and
Gillaspy, 1964). However, an extensive search of surrounding ter-
rain during the evening failed to reveal any such clusters.

Literature Cited

Evans, H. E., and J. E. Gillaspy. 1964. Observations on the ethology of digger
wasps of the genus Steniolia (Hymenoptera: Sphecidae: Bembicini). Amer.
Midi. Nat. 72: 257-280.

A TAXONOMIC REVISION OF PHYSARIA
(CRUCIFERAE) IN UTAH

Sheldon B. Waite^

Abstract. — The Physaria species which occur in Utah are revised. This
study is based on the treatment of tliis genus by Rollins (1939) and Mulligen
(1967). Keys, descriptions, distribution maps, herbarium specimen citations, and
discussions of the taxonomy of the genus are presented.

The genus Physaria is confined to western North America. It
occurs in the upper sonoran, transition, montane, and lower Canadi-
an life zones, chiefly on high plateaus and lower mountain elevations
(Rollins, 1939). This distribution extends from the southwestern
United States to the Canadian border and from the Great Plains to
the Sierra Nevada and Cascade mountain ranges. The genus, accord-
ing to Rollins (1939), is of very recent origin. Mulligen (1967)
verifies this view with studies he conducted using chromosome
counts. He infers from his chromosome studies that several species
are so closely related that a single species could include them all.
One such cluster includes three of the four species which occur in
Utah: P. acutifolia, P. chambersii, and P. newberryi. The other
closely related species which belong to the group are P. didymocarpa,
P. bellii, P. rollinsii, P. brassicoides, P. condensata, and P. floribunda.
Since morphological characteristics distinguish these groups and all
species within them are virtually isolated geographically, Mulligen
(1967) believes they should be retained as individual species.

Some integration can be observed morphologically in southern
Utah between P. chambersii and P. acutifolia. Integration is also
evident between P. chambersii and P. newberryi in the extreme
southern counties of Utah.

Mulligen (1967) corrected the name applied by Rollins (1939)
to the Physaria of eastern Utah from P. australis to P. acutifolia.
This was due to specimens studied from Colorado which have since
been named P. rollinsii (Mulligen, 1966).

Rollins described P. chambersii var. membranacea from speci-
mens taken from Red Canyon, 16 miles west of Rryce Canyon Na-
tional Park, Garfield County, Utah. The classification was based on
the membranaceous nature of the siliques. Plants taken from this
region and cultivated in different soils and environmental conditions
showed a wide variation in the texture of the valves. Mulligen
(1967) concluded that the specimens treated as var. membranacea
did not, therefore, warrant varietal status.

Specimens examined in this study are in the herbaria of Brigham
Young University (bry), Utah State University (utc), and the
University of Utah (ut). Those labeled only as Physaria sp. were
lacking fruits and were not included in citations.

^Department of Botany, Brigham Young University, Provo, Utah 84601 .

32 GREAT BASIN NATURALIST Vol. 33, No. 1

Physaria inhabits dry barren regions where sunlight is intense
and competition is minimal. The adaptation of this plant to survive
xeric conditions is reflected in the heavy pubescence on the gray
green leaves.

Physaria (Nutt.) Gray

Perennial, caespitose, silvery stellate; stems simple, arising later-
ally on a somewhat elongated caudex; basal leaves usually numerous,
often terminating the caudex or its branches in rosette form, petiolate
oblanceolate to obovate or the blade rotund, entire dentate, or divided
into segments; cauline leaves present, usually few, entire or dentate;
inflorescence congested to somewhat elongated, usually elongating
in fruit; pedicels rigid; sepals linear-oblong, pubescent, often cucul-
late at apex; petals yellow or rarely purplish, usually spatulate, glab-
rous; siliques didymous, pubescent, often highly inflated, apical sinus
present; ovules 2-6 in each loculus; style persistent; seeds brown
wingless.

Key to the Species Known in Utah

1. Style less than 3.5 mm long P. newberryi

Style more than 4 mm long 2

2(1 ) . Sinuses of silique equal above and below, valves nearly

orbicular P. acutifolia

Sinuses of silique equal above and below, valves not as
above - 3

3(2). Silique highly inflated, 1.5-3 cm wide, valves mem-
branaceous P. chambersii

Silique moderately inflated less than 1.5 cm, plants
loosely pubescent P. grahamii

Physaria acutifolia Rydb.

Physaria acutifolia Rydb., 1901, Bull. Torr. Bot. Club 28:279.

Physaria didymocarpa (Hook.) Gray var. australis Payson, 1918. Ann. Gard.

5, 144.
Physaria australis (Payson) Rollins, 1939, Rhodora 11, 408.

Description. — Small, tufted })eronnial with a deep, rather
slender taproot; basal leaves numerous 1.5-2 cm long, oblanceolate
or obovate, acute entire or slightly wavy and very finely stellate
stem; leaves rather few, oblanceolate; flowering stems 4-6 cm long,
ascending or depressed; fruit small, obtuse or slightly cordate at the
base, deeply divided above; cells inflated almost spherical, 4-6 mm
in diameter; style about 5 mm long. This species differs from P.
didymocarpa in its smaller acute leaves and in its smaller fruit,
which is more deeply divided above as well as below.

The siliques are characteristic features of this species, being
highly inflated and strongly contracted toward the replum. The

March 1973

WAITE: PHYSARIA

1. Physaria acutif olia Rydb.

2 , Physaria newberryi Gray

3. Physaria chambersii Rollins 4. Physaria srahamii Morton

Figs. 1-4. Physaria distribution in Utah: 1, P. acutif olia Rydb.; 2, P.
newberryi Gray; 3, P. chambersii Rollins; 4, P. grahamii Morton.

34 GREAT BASIN NATURALIST Vol. 33, No. 1

apical and basal sinuses are well developed and almost equal. The
valves are rounded.

Distribution. — This species grows in eastern Utah in dry,
sparsely populated plant communities at ranges from 2,000 feet up
to 9,000 feet in elevation. Members of the genus are found outside
of Utah in northwestern New Mexico, western Colorado, southeast-
ern Idaho, and most of Wyoming.

Specimen citations include the following counties: Grand, B. F.
Harrison 5968, 9-V-1933 (bry); San Juan, S. L. Welsh 2898, 31-V-
1964 (bry); Uintah, S. L. Welsh 49, 29-IV-1955 (bry); Emery,
B. F. Harrison 8076, 8-V-1936 (bry); Kane, J. R. Murdock 403,
3-V-1962 (bry); Carbon, N. D. Atwood 1300, 29-IV-1968 (bry);
Garfield, B. Maguire 19,098, 25-VI-1940 (utc); Duchesne, A. O.
Garrett 7784, 19-VII-1938 (ut); Salt Lake, A. O. Garrett 6355a, 28-
VI-1933 (ut); Tooele, J. Reveal 191 A, 15-VI-1961 (utc); Rich,
J. W. Harrison no number, 24-VI-1927 (ut) (labeled as Cache
County); and Cache, S. Flowers 223, VI-1924 (ut). Note: The
University of Utah specimens used here were labeled by the pre-
viously used name of P. australis or, in some instances, by the name
of P. didymocarpa, which is not known to grow in Utah.

Physaria newberryi Gray

Physaria newberryi Gray, 1860, in Ives' Report Colo. River, pt. 4, 6.
Coulterina newberryi (Gray) O. Kuntze, 1891, Revis. Gen. 2, 931.
Physaria didymocarpa (Hook.) Gray var. newberryi (Gray) Jones, 1895, Proc.
Calif. Acad. Sci. 2, 5.

Description. — Perennial, caespitose, silvery stellate throughout;
caudex simple branched; stems several to numerous, erect, simple,
arising laterally, 0.5-1 dm long including the fruiting raceme; basal
leaves obovate, incised, or merely dentate with broad teeth, slender
petioled, 4-8 cm long, 3-4 mm wide; petals yellow ligulate, often
truncate at apex, 10-13 mm long, 2-3 mm wide; fruiting raceme
dense, 3-5 cm long; pedicels rigid, straight, divaricate, 5-10 mm long;
siliques didymous, highly inflated, apical sinus broad, the shoulders
angular and evenly compressed with appressed stellae; valve keeled
on both outer margins, each valve 8-12 mm wide, 12-16 mm long;
replum linear, acute at apex, 8-10 mm long, 1-1.5 mm wide; style
2-3 mm long; ovules 2-4 in each locule; seeds obovate, light brown
marginless, 2-3 mm wide, 3-4 mm long.

Distribution." — This species is found only in the extreme south-
ern part of Utah growing in dry, open areas. It is found in north
central and northern Arizona and in the extreme northwestern part
of New Mexico. The distinctive characteristics of this species are
its V-shaped apical sinus, short style, and straight-sided silique.

Specimen citations include the following counties: Washington,
S. L. Welsh and G. Moore 6865, 6-IV-1968 (bry); Iron, W. P. Cot-
tarn 4724, 22-IV-1930 (ut); San Juan, B. F. Harrison 11600, 19-V-

March 1973 waite: physaria 35

1950 (bry); Kane, Cottam 4289, 12-VI-1929 (bry); and Beaver,
Cottam 8032, 8-VI-1940 (ut) (labeled as P. didymocarpa) .

Physaria chambersii Rollins

Physaria chambersii Rollins, 1939, Rhodora 41, 403.

Physaria chambersii var. membranacea Rollins, 1939, Rhodora 41, 405.

Description. — Perennial, caespitose, silvery stellate throughout;
stems numerous from a simple caudex, arising laterally, erect or
very often decumbent, simple, 5-15 cm long including the fruiting
raceme; radial leaves entire or dentate, obovate to orbicular, slender
petioled, 3-6 cm long, 1-2 cm broad; cauline few, entire, spatulate,
often acute, 1-2 cm long, 3-6 cm wide; inflorescence rather lax;
sepals linear-oblong, pubescent 6-8 mm long, 1 mm wide; petals
yellow, spatulate, 10-12 mm long, 3-4 mm wide; fruiting raceme
congested, 2-10 cm long; pedicels divaricate, slightly sigmoid, 8-15
mm long; siliques didymous, greatly inflated, evenly and often
densely pubescent, often purplish at maturity, obtuse to slightly
cordate at base; apical sinus deep and open, crests rounded; valves
subreniform, each valve 1-1.5 cm long, 1 cm wide; style 6-8 mm
long; ovules 2-6 (mostly 4) on each side of the replum; seeds orbicu-
lar, flattened brown, 2-3 mm broad, 2-4 in each loculus margin.

P. chambersii is related to P. newberryi but differs in having
round-sided siliques and apical margins, a cordate or nearly truncate
base, sinus crests rounded, style 8-13 mm long and replum 3-6 mm
long compared to keeled apical margin siliques, truncate base, sinus
crests decidedly angular, style 2-3 mm long, replum 8-10 mm long
with acute apex and straight-sided valves for P. newberryi.

Distribution.- — This species is found in the western half of
Utah, most of Nevada, northwestern Arizona, extreme southeastern
Oregon, and southwestern California.

Specimen citations include the following counties: Utah, B. F.
Harrison 8326, 18-V-1938 (bry); Juab, W. P. Cottam 7187, 28-V-
1937 (ut);* Tooele, C. Edwards 123, 15-V-1968 (bry); Garfield,
W. O. Stanton 153, 28-VI-1930 (bry); Sanpete, A. O. Garrett no
number, 29-IV-1911 (bry); Kane, Cottam 4289, 12-VI-1929 (ut);
Summit, S. L. Welsh 6263, 20-VI-1967 (bry); Millard, W. P. Cot-
tam 7347, 14-V-1939 (ut);* Washington, P. Plummer 5429, 9-V-
1939 (ut);* Sevier, R. Stevens 134, lO-IV-1966 (bry); Beaver, M.
Milner 9212, 30-IV-1946 (ut); Iron, Cottam 3922, 31-V-1926 (bry);
Piute, W. P. Cottam 7005, 15-V-1936 (ut);* and Salt Lake, S.
Flowers 1390, 1927 (ut). Plants marked with an asterisk(*) were
mislabeled as P. didymocarpa.

A specimen labeled A. O. Garrett 8333a, 9-VI-1940 (ut), from
Duchesne County, was sent to Reed Rollins, who stated in a letter
now in the herbarium that this is a new species closely related to P.
oregoni or a variety of the same. However, P. oregoni is found no
farther south than central Idaho. No further information was avail-
able concerning this specimen.

36 GREAT BASIN NATURALIST Vol. 33, No. 1

Physaria grahamii Morton
Physaria grahamii Morton, 1937, in Ann. Carneg. Mus. 26:220.

Description. — Perennial, caespitose, densely pubescent through-
out with spreading stelae; stems simple, somewhat decumbent, about
1.5 dm long; basal leaves numerous, broadly oblanceolate to broadly
spatulate, obtuse irregularly pinnatifid, 10-15 cm long, about 3 cm
broad, distal lobes large and variable, cauline, few, dentate or rarely
entire; pedicels divergent, 5-15 mm long; sepals linear-oblong, pubes-
cent, about 5 mm long; petals yellow, spatulate, 6-8 mm long;
siliques erect, didymous, inflated but not highly so, shallow sinus
below, deep sinus above; replum linear-oblong, somewhat constrict-
ed, ovules 2 on each side; style 6-8 mm long; mature seeds unknown.

The type of this species is not altogether satisfactory because the
fruits are immature. Its distinctiveness rests upon the fact that the
entire plant is covered with loose, spreading stelae and large basal
leaves deeply lobed along the margins. P. grahamii is at present
known only from the type collection.

Distribution.— This species is known from specimens collected
at Chandler Canyon, Uinta Basin, Uintah County, Utah, 3-VIII-
1935. Graham 9976 (us type).

References

Index Kewensis. 1895 to present. Clarendon Press, Oxford, England.

Kearney, T. H., and R. H. Peebles. 1951. Arizona Flora. University of
California Press, Berkeley and Los Angeles.

MuLLiGEN, G. A. 1966. Two new species of Physaria (Cruciferae) in Colo-
rado. Can. J. Bot. 44:1661-1665.

. 1967. Cytotaxonomy of Physaria acutifolia, P. chamber sii, and P.

newbe-ryi (Cruciferae). Can. J. Bot. 45:1887-1898.

Rollins, R. C. 1939. The cruciferous genus Physaria. Rhodora 41:392-415.

TiDESTROM, I. 1925. Flora of Utah and Nevada. Government Printing Office,
Washington, D.C.

Welsh, S. L., and M. Treshaw and G. Moore. 1964. Guide to common Utah
plants. Brigham Young University Press, Provo. Utah.

TWO NEW SPECIES OF GYMNODAMAEUS

FROM COLORADO

(ACARINA: CRYPTOSTIGMATA, GYMNODAMAEIDAE)^

Tyler A. Woolley- and Harold G. Higgins'^

Abstract. — Gymnodamaeus plokosus and G. leurolomasus, n. spp., are de-
scribed from soil at an ecological study site near Hayden, Colorado. Their rela-
tionship to allied species is discussed.

A review of the literature indicates that the first species of
Gymnodamaeus were described by Berlese (1910, 1916). Woolley
(1972) reviews the literature of known soil mites in the genus
Gymnodamaeus and describes a new species, Gymnodamaeus chala-
zionus, from the motmtainous areas of northern Colorado. The two
new species described below are additions to this genus.

The specimens of the first species described below were taken in
soil beneath sage and grass along the undisturbed margin of a spoil
bank near one of the coal strip-mining sites of the Hayden, Colorado,
power plant. Comparisons of these specimens with others in the
collections of the writers and with descriptions in the literature indi-
cate that the species is new. It is described below and compared
with other known species.

Those species of Gymnodamaeus from North America with which
this new species were compared are G. veriornatus Higgins, 1961,
and G. chalazionus Woolley, 1972. The new species is smaller in
size than the above species but averages very close to the size of G.
gildersleeveae Hammer, 1952, and G. elegantulus Hammer, 1958;
it is smaller than G. minor Banks, 1947, slightly larger than G.
pearsei Banks, 1947, and much larger than G. quadricaudiculus
Jacot, 1937. The details of this size comparison are given below with
collection data and measurements of the new species.

Gymnodamaeus plokosus, n. sp.

(Figs. 1, 2)

Diagnosis." — Most easily recognized by the crenulated posterior
margin of notogaster with four nubbins, each bearing an ornate,
curved hair (Figure 1); differs from G. gildersleeveae Hammer,
1952, in this feature as well as in the marginal placement of the pos-
terior notogastral hairs; without reticulate pattern on dorsum of
notogaster as in gildersleeveae, but with an arched, transverse bar on
prodorsum just anterior to interlamellar hairs which is not present
in gildersleeveae. Differs from G. quadricaudiculus Jacot, 1937, in
the absence of the dorsal oval sculpturing with rays. The inter-
lamellar hairs of the new species are short and most nearly resemble

^Research supported by the Yampa Project; Ecology Consultants, Inc.

-Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado.

^Biology Department, Granger High School, Granger, Utah.

GREAT BASIN NATURALIST

Vol. 33, No. 1

those of G. chalazionus Woolley, 1972. The sensillum of the new
species is most similar to G. gilder sleeveae, broadly flabelliform and
spined for its entire length.

The trivial name plokosus is from the Greek, meaning "a lock of
hair," or "curled," and refers to the four ornate, posterior notogas-
tral hairs that arise from the nubbins and are so characteristic of the
species.

Description. — Color golden brown with a tuberculated, yellow-
ish cerotegument giving a halo effect around the margins of body
and legs; this cerotegument on prodorsum, notogaster, venter, legs,
prodorsal hairs (except for interlamellar hairs), and notogastral
hairs; prodorsum less than half as long as notogaster, broadly trian-
gular in shape, with blunt, squarish rostrum; rostral and lamellar
hairs of about equal length, decurved, covered with tuberculated
integument; rostral hairs inserted in slight notches in anterolateral
margin of prodorsum; lamellar hairs inserted in short apophyses at
lateral margins, a transverse, sclerotized bar extending between in-
sertions, hairs also covered with tuberculated integument, but
tubercles of minute size compared to those of prodorsum; an arched,
slightly U-shaped bar in posterior third of prodorsum immediately
anterior to interlamellar hairs, extending in width about distance
between interlamellar hairs; interlamellar hairs short, simple, in-

Figs. 1-2. Gyninodaniaeus plokosus: 1, dorsiil aspect, legs partiallj emitted;
2, ventral aspect, legs omitted.

March 1973 woolley, higgins: new mites 39

serted in inverted V-shaped apophyses posterolaterad of arched bar;
pedotecta I and II as in Figure 1, with tuberculated surface due to
cerotegument; pseudostigmata rounded, cup-shaped, with lip raised
above surface of prodorsum, covered externally and internally with
small tubercles; sensillum flabelliform, flattened in one plane, head
broad, pedicel narrow, spined throughout length.

Notogaster oval in shape, with nearly straight anterior margin;
surface covered with tuberculated cerotegument; 8 to 10 large, more
flattened tubercles near median area of anterior margin, other tu-
bercles of surface smaller, more rounded; four short, sclerotized nub-
bins posteriorly, forming a crenulated posterior margin; each nubbin
with an inserted curved, ornate hair covered with small tubercles,
a similar, less curved hair anterolaterad of nubbins on each side
(placement of these six hairs apparently a specific characteristic in
species of Gymnodamaeus) ; other hairs and surface features as in
Figure 1.

Camerostome somewhat triangular, with heavily sclerotized mar-
gins; ventral setae, apodemata as seen in Figure 2; ventral surface
with tubercles, but ventral tubercles only a third as large as dorsal
tubercles; ventral setae simple, short; apodemata IV arched anterior-
ly over genital opening as in other species of Gymnodamaeus; geni-
tal and anal openings contiguous throughout widths; genital covers
with cerotegument of tubercles, each cover with six genital setae;
aggenital setae short, simple, laterad of genital opening; anal covers
with tubercles, each cover bearing two simple anal setae; adanal
setae ornate, covered with cerotegument of smaller tubercles
(Figure 2).

Legs heterotridactylous; surface of legs covered with cerotegument
similar to dorsum; hairs of legs also covered with tuberculated cero-
tegument, but tubercles of proportionately smaller size.

Collections and measurements. — Ten males and two females
of G. plokosus were collected by the writers at the Seneca site num-
ber 2, Hayden, Colorado, lO-IV-1971. The males ranged in size
from 432-372 i.l x 216-186 /t, with an average of 399 /i x 207 ^u. Four
females and three males were collected from the same site on 8-VI-
1971 by the writers. These males ranged in size from 420-408 jx x
210-204 fi, averaging 410 /x x 208 /*,. The females ranged in size from
444-420 jx X 238-210 jx, with an average size of 432 /x x 228 fx.

Additional specimens were taken in the project areas as follows:
3 males (425-415 /x x 212-205 /x) from under scrub oaks, 4 miles N
Seneca Road, l-VIII-1971, by H. G. Higgins; 2 males (425-430 /t x
210-205 jx) from under chokecherries, one-fourth mile N Hayden
Power Plant, l-VIII-1971, by H. G. Higgins; 1 male specimen (395
IX X 205 ix) under serviceberry on spoil bank, lO-IV-1971, by H. G.
Higgins and T. A. Woolley; 1 male (380 /x x 210 /x) from beneath
sagebrush, two miles S Yampa Valley Airport, 9-VI-1971, by H. G.
Higgins and T. A. Woolley. Three specimens (2 females and 1
male) of this species were collected at Terry Falls, Trail Lake Ranch,
Dubois, Wyoming, 20-Vn-1964, by H. and M. Higgins. They
ranged in size from 432-420 /x x 210-204 /x. A single specimen was

40 GREAT BASIN NATURALIST Vol. 33, No. 1

also collected by H. and M. Higgins at Soapstone, Wasatch County,
Utah, on 4-IX-1955. The specimen is so broken, however, that sex
and size are not distinguishable. Two females were collected by
T. A. Woolley in South Dakota, one (426 /<. x 204 ix) at Black Hills,
28-VIII-1968, and one (426 f/. x 316 /i) in the Nordbeck Game Pre-
serve, 28-VIII-1968.

The measurements of the width of all measurable specimens
were taken at the level of legs IV. Generally speaking, females ap-
pear to be larger and more elongated in shape than males.

The drawings were made of a male specimen collected 10-IV-
1971. The type and a para type specimen will be deposited in the
U.S. National Museum.

The second new species was collected in soil beneath aspens and
scrub oak in an undisturbed area about four miles south of the spoil
banks mentioned above. Compared to G. plokosus, it is much larger.

Gymnodamaeus leurolomasus, n. sp.

(Figs. 3, 4)

Diagnosis. — Larger size (.70 mm) median edge of genital plate
smooth, lacking the interlocking dentes found in both G. chalazionus
and G. veriornatus; each genital cover with seven setae; entire body
and setae covered with a fine, granular cerotegument. The words
leuros 'smooth' and loma 'border,' from the Greek, refer to the smooth
median edge of the genital covers. This feature, along with the seven
pairs of genital setae, distinguishes the new species from the other
larger representatives of Gymnodamaeus from North America.

Description.— Color dark reddish brown, entire dorsal and
ventral surfaces covered with a granular cerotegument; prodorsum
about half as long as notogaster, broadly triangular in shape; rostral
and lamellar setae about equal in length, rostral setae inserted dorsal-
ly, posterior to rostral margin but anterior to transverse sclerotized
line between lamellar hairs; interlammellar hairs short, simple, coni-
cal, each inserted near tip of curved, sclerotized, median ridge an-
terior to pseudostigmata; in some specimens, a faint, transverse,
curved sclerotized bar can be seen antoriomediad of interlamellar
hairs (Figure 3); costulae absent; pseudostigmata rounded, with
edge erected above surface of prodorsum; sensillum clavate, surface
of club spined; entire organ slightly longer than prodorsum.

Hysterosoma longer than wide, with a wrinkled surface; posterior
margin with six setae and two pairs of dorsal setae of different sizes,
with muscle scars and j)ores as shown in Figure 3; no distinct dorsal
ornamentation as seen in G. veriornatus Higgins or G. ornatus
Hammer.

Camerostome oval in outline, montum broad, rutella narrowed
anteriorly; ventral surface with apodemata and setae as shown in
Figure 4; genital and anal apertures contiguous; genital covers
smooth on medial edge, each cover with seven simple setae; anal
covers much longer than wide, each cover with two simple setae;

March 1973

WOOLLEY, HIGGINS: NEW MITES

Figs. 3-4. Gymodamaeus leurlamasus: 3, dorsal aspect, legs omitted; 4 ven-
tral aspect, legs omitted.

entire ventral surface covered with a granular cerotegument that
obscures the location of coxistemal setae.

Legs long, surface and hairs covered with cerotegument; all legs
heterotridactylous, median claw larger than laterals.

Collections and measurements.- — Length, .65-. 70 mm; width,
.36-.41 mm. The type, a male, and eight other paratype specimens
(six males, two females) were taken from under quaking aspens,
4 miles S Seneca Road, Hayden, Colorado, 8-X-1971, by H. G. Hig-
gins. Five specimens were taken from under scrub oaks associated
with aspens, 4 miles S Seneca Road, Hayden, Colorado, 1 -VIII- 1971,
by H .G. Higgins. The type and one paratype will be deposited in
the U.S. National Museum.

Discussion and Ecological Notes

It is interesting that two species of smaller Gymnodamaeus, G.
gildersleeveae and G. plokosus, should be found in the project area.
The latter species was first found under sagebrush along the edge
of a spoil bank in rather dry soil. Subsequent collections have shown
that this species is fairly well distributed throughout the western
United States (Colorado, North Dakota, Utah, and Wyoming) as
well as being found in the litter of several plant communities. For
example, G. plokosus has been taken in rather dry soil and litter

42 GREAT BASIN NATURALIST Vol. 33, No. 1

under scrub oaks, serviceberry, sagebrush, and chokecherry in the
Hayden project area. G. gildersleeveae has been found only under
quaking aspens or in more moist scrub oak litter closely associated
with aspens. This suggests that the populations of G. gildersleeveae
prefer a more humid habitat, while G. plokosus is found mainly in
a drier habitat.

Gymnodamaeus leurolomasus must be listed among the larger
North American members of this genus, but it is smaller than either
G. veriornatus or G. chalazionus. It is nearer the size of G. ornatus
Hammer from Canada but lacks the dorsal ornamentation so well
known for that species. It differs from both G. veriornatus and G.
chalazionus in that it lacks the interlocking dentes on the median
edges of the genital plates and possesses seven pairs rather than six
pairs of genital setae. This disparity in the number of genital hairs
is considered a specific characteristic, inasmuch as the generic fea-
tures match other representatives known from North America.

Literature Cited

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.

Hammer, M. 1952. Investigations on microfauna of Northern Canada. Acta

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

and 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 (1-2) :27-28.
Jacot, a. p. 1937. Journal of North American moss mites. J. New York

Entomol. Soc. 45(3-4) : 353-375.
WooLLEY, T. A. 1972. A new species of Gymnodamaeus from Colorado. Great

Basin Nat. 32(2):97-103.

NEARTIC DESERT DECTICIDAE (ORTHOPTERA). PART II.
A NEW GENUS AND SPECIES FROM ARIZONA

Ernest R. Tinkliami

ABSTRACT. — Plalyoplus, n. gen., is erected for P. gilaensis, n. sp., from
the Gila Mountains, Yuma Co., Arizona. The method of preservation for these
large, soft-bodied insects is described.

The genus and species herein described was discovered in the
Gila Mountains, 20 miles east of Yuma, Arizona, on 26 April 1958.
The first specimen, found in a crevice of a giant boulder on a rocky
ridge, eluded capture; however, other specimens were taken later.

Platyoplus, n. gen.

This new eremophilous, xerophilous, and petrophilous genus
portrays characters that would place it between Ateloplus Scudder
and Inyodectes Rentz and Birchim, with closest relationships indi-
cated to Ateloplus. There appears to be little relationship shown to
Eremopedes Cockerell and Pediodectes Rehn and Hebard. A review
of the cereal features found in Ateloplus shows that four species,
namely, notatus, luteus, hesperus, and splendidus have rather slender
cerci with inner apical uncinate hook, while minor and schwarzi
have shorter, quadrate cerci with inner apical tooth. The cerci of
Platyoplus, with an inner subapical projection with uncinate tooth,
is somewhat like that in Oreopedes Rehn and Hebard but is amply
distinct in many ways. On the other hand, the features of the
pronotum are quite ateloploid in nature; but even here the diagnostic
differences are the shallower lateral lobes, which are quite outwardly
flared and with those portions of the lateral lobes of the metazona
tumidly raised and semitransparent above the greatly enlarged tym-
panum, which in itself is diagnostic for the genus. Such features,
the writer believes, warrant generic recognition. Experience based
on many years of study convinces the author that any member of a
particular genus must have the same characters as the other species
of that genus. Thus the cercus of Ateloplus has an inner apical
tooth, whereas in other genera such as Platyoplus and Eremopedes
the inner cereal tooth is subapical or intermediate in position, al-
though in some Eremopedes the cercus is also undulate and quite
distinctive in character. Likewise, the pronotum in the various
genera such as Platyoplus, Ateloplus, Eremopedes, Pediodectes, In-
yodectes, and Oreopedes are different in each genus. Added to these
and other characters not mentioned is the true tympanum, which is
distinctive in the various genera mentioned above and portrays its
greatest development in Platyoplus.

Description. — Head broad and rather shallow, face with great-
est breadth just below compound eyes, its breadth 1^4 times depth

'81-441 Date Palm Avenue, Indio, California 82201

44 GREAT BASIN NATURALIST Vol. 33, No. 1

from vertex to clypeal suture. Fastigium with frontal costa broad,
flat, abruptly terminated intermediately between the antennal
sockets.

Pronotum diagnostic, flat and broad, its breadth equal to its
length, with shallow, reflexed lateral lobes; metazona shorter than
prozona, its lateral lobes tumidly swollen over upper half of greatly
enlarged true tympanum.

Penultimate abdominal notite, with shallow U-shaped median
groove. Cerci moderately slender, with an inner quadrate subapical
prominence bearing an uncinate hook.

Legs average, strong; caudal femora stout and shorter than in
related genera, with both ventral keels bearing six widely spaced
teeth in the apical half. Caudal tibiae fully spined, with numerous
teeth along the entire ventral keels and with four pairs apically on
the upper carinae of the caudal tibiae.

Ovipositor stout, apical portion very slightly recurved, apex
barely surpassing geniculae of caudal femora. Subgenital plate with
a small, V-shaped median notch situated apically in the male; in
female, deep median emargination, its margins very strongly raised
with keels continuing basadly and convergent with an additional
very high and narrow median keel connecting from base of deep,
U-shaped emargination to base of plate; this character quite diagnos-
tic for genus and not observed in related genera.

Coloration pale brownish with very fine mottling of pale purplish
dots and with genicular areas of caudal femora black.

Type species. — Platyoplus gilaensis Tinkham, by monotypy.
This genus is named after the very broad and flat pronotum which
partially hides the very large true tympanum. The species is named
after the Gila Mountains, that barren range of rocks about 20 miles
east of Yuma, Arizona.

Platyoplus gilaensis, n. sp.

Description. — Male: head concolorous, unmarked; eyes sub-
globose, their depth about equal to their breadth. Antennae fully
twice body length. Head broad and short, facial breadth just below
compound eyes 1 1/4 times depth from vertex to clypeal suture.
Fastium angularly rounding into flat and smooth frontal costa, its
margins convergent forward and roundly terminated medianly be-
tween antennal sockets.

Pronotum diagnostic, quite flat with breadth equal to length,
very shallowly rounded into flaring lateral lobes; metazona nearly
flat due to tumid swelling of metazonal lobe above upper half of
greatly enlarged true tympanum; tympanum broadest in its apical
half under swollen metazonal flange. Metazona shorter than pro-
zona; prozona with a prominent but shallow, very broadly V-shaped
sulcus, about two-fifths of prozonal length posteriad of anterior mar-
gin. Anterior margin of the pronotum very smoothly arcuate for its
entire breadth to the anterior lateral angle of lateral lobes; posterior
margin almost squarely truncate, with at most only a slight sugges-

March 1973

TINKHAM: DESERT ORTHOPTERA

9 p^^^f ,.1-7- ^^«^ropZi/5 gilaensis Tinkham: 1, Male holotype, profile view
oiaSTl^'t. l"""^" ^°^°'^P" showing notite, cerci, titillator? 'arfd subgemt'l
plate d, Lateral view of pronotum of male holotype; 4, Tympanum of male holo-

rtv?eVTatf l'"- '^P'f^ ^' P-t--/ P0^*^°" ^f -bgeSital plat oFfemale

46 GREAT BASIN NATURALIST Vol. 33, No. 1

tion of emargination. Lateral lobes, seen from above, arcuately
rounded, metazonal area outwardly bowed due to convexity of meta-
zonal lateral lobe area above enlarged tympanum — a feature dis-
tinguishing Platyoplus from all other desert decticids. Tympanum
very large.

Entire body surface smooth. Abdomen with ultimate notite
bearing a moderate, median, V-shaped notch. Cercus moderate in
build, broadest at base and very slightly tapered to apical two-thirds,
inner margin of this area with a quadrate projection bearing a small,
inner, slightly uncinate tooth. Subgenital plate with posterior lateral
margins straight and with a median, posterior, V-shaped notch
flanked outwardly with a short stylus. Plate distinguished further
by a prominently rounded ridge extending anterior-laterally from
base of each stylus, the ridges merging with basal portions of plate.

Wings exposed for a length equal to that of the metazona.

Forelegs with prominent procoxal spur; profemora with one to
three very small inferior spines on exterior margin (leg forward)
and four slightly larger spines on inner ventral keel. Protibiae with
six or seven usually paired larger spines on ventral keels and two
outer external spines dorsally, one of which is basal. Mesoleg with-
out mesocoxal spine; mesofemora with five outer and two inner sub-
apical small spines on ventral keels (legs in backward position);
mesotibiae with six pairs of ventral spines and two pairs dorsally, as
in protibiae. Hind legs with caudal femora bearing six spines on
each ventral keel. Caudal tibiae heavily spined; 27-28 external
and 24-26 internal strong spines on ventral keels and eight outer and
inner attingent, widely spaced, slender spines on upper keels (ven-
tral in position on folded legs). All tarsi three-segmented.

Living Coloration (notes made 1965): Head with face faintly
purple gray; eyes with incomplete cross-striations of purplish brown.
Antennae reddish brown, nodes narrowly ringed in darker reddish
brown. Pronotum dull grayish brown, tinged more definitely with
burnt umber on central portions of metazona and more narrowly on
front central edge of prozona. Tegmina pale purplish gray, with
cells in exposed central portions piceous. Abdomen above citrus
yellow, profusely and finely mottled with dark j)urplish brown, pos-
terior marginal areas with about 12 washed or running spots of dark
reddish brown. Sternites of thorax and abdomen citrus yellow.
Fore and middle legs pale reddish brown; caudal femora pale reddish
brown, outer pagina with a single central piceous band heaviest pos-
teriorly; geniculae and pregenicular area dark piceous or almost
black; spines of lower keel dark reddish brown. Caudal tibiae pale
reddish brown, piceous at extreme base with both dorsal spine rows
tipped in apical half with reddish brown; fewer scattered ventral
spines similarly colored.

Female: Slightly larger than male, but otherwise closely similar
in coloration and spination of legs. Ovipositor slightly shorter than
length of abdomen, rather heavy in build and very gently recurved,
apex of ovipositor barely exceeding apices of caudal femora. Sub-
genital plate diagnostic, with a large and deep quadrangular emargi-

March 1973 tinkham: desert orthoptera 47

Fig. 8. Photograph of living Platyoplus gilaensis Tinkham on rock of native
habitat.

nation occupying two-fifths of total length of subgenital plate.
Lateral margins of emargination prominently raised and extending
basadly to terminate in an arcuate fashion about half the distance
to base of plate, plus a prominent median keel running from center
of quadrangular emargination to base of subgenital plate.

HoLOTYPE. — Male, Telegraph Pass, Gila Mountains, Yuma Coun-
ty, Arizona, 20 miles E Yuma, Arizona, 800 ft elevation, 30-V-1965
(night collecting on Creosote Larrea divaricata on steep rocky slopes;
Ernest R. Tinkham). Caliper measurements in mm: body length
22.0; pronotum 7.75 long x 7.2 broad; tegmina exposed 2.0; caudal
femora 24.3; antennae 51.5. Holotype deposited in the Tinkham
Eremological Collection.

Female allotype: Same data as holotype. Caliper measurements
in mm: body length 24.9; body length to apex of ovipositor 39.2;
ovipositor 16.0 x 1.6 in middle; pronotum 7.7 x 7.7:, antennae 51.6;
caudal femora 24.9 x 5.2. Allotype in the Tinkham Eremological
Collection.

Male para types: Twelve males same data as holotype; 2, 25-1 V-
1960. Range in calipered measurements (mm): body length 21.6-
28.2; pronotum 7.2-8.1 (length) x 7.2-7.2 (breadth); caudal femora
23.1-24.0. Paratypes identical to the holotype male in every respect
including coloration.

Female paratypes: Twelve females same data as allotype. Range
in calipered measurements (mm): body length 23.6-27.6; total body
length to apex of ovipositor 38.5-42.0; pronotum 8.2-8.8 x 7.2-7.8
(breadth); caudal femora 24.1-26.0; ovipositor 15.7 x 15.7.

48 GREAT BASIN NATURALIST Vol. 33, No. 1

Paired para types will be presented to such major orthopterologi-
cal museums as the following: Academy of Natural Sciences of
Philadelphia, British Museum, California Academy of Sciences, Uni-
versity of Michigan Museum of Zoology, Smithsonian Institution,
and the Tinkham Eremological Collection.

Biology.- — With the advent of the necessary winter rains in the
Colorado Desert, it is believed that the nymphs hatch out either in
the late fall or early winter. By late May the nymphs are in their
last nymphal stadia and become adults in late June or very early
July. The author has kept them alive in his home until late Novem-
ber, but whether they would survive the torrid heat of their rocky,
inhospital habitat in the Gila Mountains is questionable.

Faunal Designation: Platyoplus gilaensis is a member of the
Colorado Desert fauna, since the Gila Mountains represent the east-
ern periphery of that desert. The area ranging from just east of the
Gila Mountains and extending 100 miles to Gila Bend is Gila Desert.
Both are eremological components of the Great Sonoran Desert,
which is composed of seven different deserts. At or in the region
of Gila Bend, the Gila Desert merges with the Sahuaro Desert.

Floral Characteristics: In addition to Larrea divaricata, and
Franseria dumosa growing sparsely on boulder-strewn slopes, Bursera
microphylla is here more commonly represented than in any other
desert sierra in the United States. Occasional Cercidium microphyl-
lum and Hyptis emoryi, Hoffmanseggia microphylla, Franseria ilici-
folia, Ditaxis lanceolata, Eneclia farinosa, and rarely Carnegica
gigantea are all characteristic of these barren, but very interesting,
mountains.

Orthopteran associates: The orthopteran fauna is naturally
quite impoverished, even in good years when up to three inches of
rain may fall. However, since drought years have prevailed for
the past decade, it is likely that some of the members present in the
early forties are now extinct. Thus, Tanaocerus reported as koebe-
lei in 1947 has not been seen since 1942. Likewise, Oedomerus coral-
lipes, discovered in 1942 and reported for the first time in the
United States in 1947, may also be close to extinction. Other orthop-
terans taken in the forties, including Arphia aberrans, Tytthotyle
maculata the Malpais Lubber, and even Trimerotropis p. pallidi-
pennis, have not been seen since 1949. Two other orthopterans are
known from the Gila Mountains, namely, Schistoccrca vaga and
Capnobotes fuliginosus, both considered very rare.

Host Plants: Platyoplus gilaensis was first discovered hiding in
a crevice of a giant boulder by day, in 1958, and it was not until
1960 that the first specimens were taken by night collecting. The
subadult nymphs were found feeding in creosote bushes, about the
only plant available in that habitat. On another occasion, also on
creosote, a fine colony was found in a small mountain canyon. As
far as is known, creosote is the host plant in nature.

Song: The stridulated note of Platyoplus is a soft "zee-zee-zee,"
long continued but scarcely audible beyond ten feet. The very large

March 1973 tinkham: desert orthopter a 49

tympanum characteristic of Platyoplus may be necessary in order
to hear the feebly stridulated song.

Specimen preservation. Considerable has been written about the
preservation of desert decticids, which are admittedly hard to pre-
serve while maintaining colors that are close to those existing in the
living creature. The usually large bodies filled with body fluids and
the delicate coloration make good preservation difficult. The secret
to fine preservation of body and color is the immediate gutting of
the specimen upon its death. The decticids are collected alive in the
field in specially designed cages, which the author has designed for
the occasion. In this particular case, the large series of subadults was
collected in the mountains in late May and removed to the author's
home for proper study. Many biological notes were taken while the
subadult specimens were developing into adults. In August, when
the adults had been imagos for a month and all tissues were well
hardened, small series at a time were killed with potassium cyanide.
Each specimen killed was then immediately gutted by removing the
alimentary tract and the gonads through a snip centrally or laterally
on the three or four basal abdominal stemites. A little careful swab-
bing should be done to remove excess liquid, but care should be taken
not to damage or disturb the subcutaneous color structure. A small
fluff of cotton can then be inserted through the incision, although
this is not necessary. If cotton is used, care should be taken that it
not be rolled, because if a roll of cotton were pushed into the tho-
racic cavity, insertion of the insect pin could cause the whole to be
pushed through the thoracic sternites, to the ruination of the speci-
men.

The specimens are now ready to be pinned, with legs and an-
tennae arranged with the use of additional pins, on a sheet of balsa
or white plastic cellulose. When the legs have been positioned by the
pins and the antennae pulled back and positioned so that they lie
along the dorsolateral line of the body, the creatures are ready for
drying.

The pinning block is now placed in a gas or electric range. If
gas, the oven should be only at pilot heat; that is, no burners on and
the temperature no more than 125 F. Here they are watched and
removed after several hours when dry. Care must be taken not to
use more than gentle heat, since color injury can result. After
several hours the specimens should be dry and the coloration in life
almost perfectly preserved. Living color notes should be made of
the freshly killed creatures.

One other method, relatively new, is also available. This is the
freeze-dry method, where the creature is frozen immediately upon
killing or death and left frozen for a long period of time, during
which time it is completely desiccated by the cryoprocess. If this
method is used, it may be necessary to relax the specimen or speci-
mens overnight unless the specimens were pinned and legs arranged
for the cryoprocess, which all depends upon the space available in
the freezer.

50 GREAT BASIN NATURALIST Vol. 33, No. 1

References

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the Gulf of California in 1921. Dermaptera and Orthoptera. Proc. California

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the Mexican boundar}\ Part III. Revisionary data and new North American

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Dermaptera and Orthoptera, with new records and corrections of the litera-
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of North American Decticinae (Orthoptera: Tettigoniidae) . Trans. Amer.

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Rentz, D. C, and J. D. BuRCHiM. 1968. Revisionary studies in the Nearctic

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keys and descriptions of seven new Chinese species. Notes d'Entomologie

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pis.
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katydids of the North American deserts. Amer. Midi. Nat. 31 (2):257-328,

figs. 1-28.
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in Arizona. Amer. Midi. Nat. 38(1) : 127-149, pis. 1-4.
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Bend region of Trans-Pecos Texas, with especial reference to the Orthopteran

zones and faunae of midwestern North America. Amer. Midi. Nat. 40(3):

521-663, figs. 1-37.
. 1960. Studies in desert sand dune Orthoptera. Part I. A new species of

Plagiostira from eastern New Mexico, with key and notes. Great Basin Nat.

20(1 & 2): 39-47, 10 figs.
. 1962. Studies in Nearctic desert sand dune Orthoptera. Part VII. A

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Eremopedes (Decticidae). Great Basin Nat. 32(4): 223-228.

AN UNUSUAL POPULATION OF SPIDERS IN UTAH

Dorald M. Allred'

An unusual population of spiders belonging to the species Neo-
scona oaxacensis (Keyserling) was observed on West Mountain in
Utah County, Utah, by Miss Mary Fenley and her mother, Mrs.
Ed. J. Fenley, of Provo, Utah, while on a Labor Day outing in
1971. They kindly directed my attention to the phenomenon.

West Mountain is situated in a north-south axis along the south-
eastern side of Utah Lake. Its three major peaks have elevations of
6083, 6813, and 6904 feet, respectively. The level of the lake is at
approximately 4487 ft. The predominant vegetation consists of
various grasses along the foothill areas, and sagebrush and rabbit
brush merging with a scattering of juniper trees at the higher eleva-
tions.

The spiders were densely distributed over several acres in the
sagebrush and rabbit brush about midway up the mountain on the
northern slopes. Only adults were present, and the females were
predominant in number. The webs were not ornate as is typical of
some of the other species of orb weavers. The radii of the webs were
almost exclusively situated in some part of the individual shrub
rather than in the spaces between shrubs. However, bridge lines
frequently extended between separate plants. The bridge lines were
of unusual strength, causing the limbs of plants to which they were
attached to bend toward one another. As I walked between the
plants and consequently broke the bridge lines, the tension of each
line against my legs was almost like that of a lightweight twine.

Several random counts were made of the unusually high popula-
tion of spiders. These varied from 10 to as many as 60 individuals
per square meter. In more than 20 years of extensive field investi-
gations in temperate desert areas I have never observed such a high,
localized population of spiders of any species. Some of my colleagues
stated that they had noted that populations of spiders of other species
in Utah were much higher this year than they had ever seen pre-
viously; but they had not observed such a density as reported here.
Dr. Willis J. Gertsch, who kindly identified the spiders and pro-
vided some information on their biology, indicated that many years
ago the same species occurred in great numbers on the old Saltair
Pavilion on the shore of Great Salt Lake west of Salt Lake City.
Incidentally, Ralph V. Chamberlin at that time named the species
Neoscona saleria, now a synonym.

According to Dr. Gertsch, Neoscona oaxacensis is a common orb
weaver of the southwestern United States, ranging southward
through Mexico to Panama. The species has had several names,
probably the most familiar of which is A^. vertehrata McCook, based
on specimens from California. Neoscona oaxacensis is probably the

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

52 {}REAT BASIN N.'.TURALIST Vol. 33, No. 1

largest of the species of the "smaller" round-shouldered araneas with
elongate oval abdomens.

It is a species that often lives in clusters, with males, females,
and the young stages living close together in webs. However, they
are not necessarily tied by social habits, and their webs are single
and not communal ones.

Continued annual observations of this species on West Mountain
in order to determine cyclical population fluctuations related to
environmental influences would be desirable.

SOME HELMINTHS FROM MINK IN SOUTHWESTERN

MONTANA, WITH A CHECKLIST OF THEIR

INTERNAL PARASITES

Delbert L. Barber^'- and Lawrence L. Lockard^

Abstract. — Thirty-four percent of 100 mink examined from Gallatin and
Madison counties, Montana, revealed the presence of Perostrongylus pridhami An-
derson, 1962 (Anderson, 1963), in the lungs. This is the first report of P. prid-
hami in the United States. Thirteen percent of tlie mink were infected with
Taenia mustelae Gmelin, 1790. This is the first report of T. mustelae in south-
western Montana. A checklist of internal parasites of Mustela vison is included.

Parasites of mink in North America have been reported by nu-
merous authors. No previous parasite surveys have been conducted
on Mustela vison in southwestern Montana. The mink necropsied
in this study were trapped in the Madison River, upper Gallatin
River, and headwaters region of the Missouri River drainages
during the winters of 1969 and 1970.

A total of 100 mink were examined. These animals were eviscer-
ated and the viscera were placed in plastic bags and frozen. An NaCl
fecal flotation was prepared to determine the presence of cestode
and nematode eggs or larvae prior to examination of the animals.
The gastrointestinal tract was dissected using an enterotome device
(Figure 1). The contents were washed onto a 200-mesh screen.

Fig. 1. Enterotome Device. Seam ripper modified with plastic bead placed
on end of point to facilitate intestinal incisions.

then transferred to an illuminated tray for examination (Figure 2).
Cestodes were fixed in 10 percent formalin, stained in Delafield's
hematoxylin, dehydrated in ethanol, cleared in beechwood creosote,
and mounted in HSR (Hartman-Leddon Co.). Lungs, liver, and kid-
neys were dissected and each was placed in a jar with water and
agitated on a mechanical shaker for 20 minutes. The contents were
poured onto a 200-mesh screen, washed, transferred to an illumi-
nated tray, and examined. Lung tissues that appeared to contain
cysts or capsules were pressed between glass plates and observed
under a dissecting microscope. Adult nematodes removed from
lung parenchyma were fixed in 70 percent alcohol-5 percent glyc-
erol and mounted in glycerol. Skulls, when available, were ex-
amined for nasal nematodes.

^Present address; 3131 E. 12th, Casper, Wyoming 82601.

^Department of Zoology and Entomology, Montana State University, Bozeman, Montana 59715.

GREAT BASIN NATURALIST

Vol. 33, No. 1

IHBJH^^^^HH^H

■ - " f

I'^BhHKI

^^HB'l ^1

iiHmiii

HHR^^^^v

I^H

^ flHJ^^H

^^^1

Fig. 2. Illuminated tray used in postmortem recovery of helminth parasites.

Results and Discussion

Fecal examinations revealed that 34 percent of 100 mink were
passing first-stage nematode larvae. Three male nematodes and
portions of two females were dissected out of the lung tissue; larvae
were removed from the gravid females. These larvae were compared
with those found in the fecal samples and it was determined that
they were of the same type. The adult males were identified as

Table 1. Checklist of parasites reported from inink.
Parasite Bibliography number

Treniatoda (Flukes)

Alaria freundi

A. minuta

A. mustelae

14, 15, 30, 31, 61

Apophallus muehlingi

Baschkirovitrema incrassatum

Cephalophallus obscurus

Cryptocotyle concava

C. lingua

57, 58

Enhydridiplostomum alarioides

Euparyphium beaveri

E. incrme

1, 51, 69, 90

¥.. mclis

11, 30, 31, 38, 39, 46, 51

Euryhelmis monorchis

4, 30, 31, 61

E. pacificus

61, 80

E. pyriformis

E. squnrnula

38, 54, 61

Fasciola hepatica

March 1973

BARBER, LOCKARD: MINK PARASITES

Table 1 (continued)

Fibricola cratera
Metagonimoides oregonensis
Metorchis conjunctus
Nanophyetus salmincola
Neodiplostomum lucidum
Paragonimus kellicotti
P. westermani
Parametorchis canadensis
Procyotrema marsupiformis
Sellacotyle mustelae
S. vitellosa
Tocotrema lingua
Troglotrema acutum

47, 61, 81

30, 31, 35

10, 38, 46, 78

2, 3, 12, 30, 31, 35, 38, 49, 61, 79, 93

30, 31, 61, 94

61, 84

Cestoda (Tapeworms)
Diplogonoporus tetrapterus 71

Mesocestoides litteratus 30, 31

Moniezia sp. 58

Taenia mustelae { = tenuicollis) 5, 29, 30, 31, 32, 33, 44, 45, 52, 59,

61, 66, 70, 72, 77, 83, 92

Centrorhynchus conspectus
Corynosoma hadweni
C. semerme
C. strumosuni
C. sp.
Macracanthorhynchus ingens

Eimeria mustelae
Isospora bigemina

:anthocephala
61
58
28, 67

28, 58,

17, 61

Protozoa

Nematoda (Roundworms)

Aelurostrongylus falciformis
Ascaris sp.

Capillaria mustelorum
Crenosoma hermani
Dictyocaulus filaria
Dioctophyme renale

Dranunculus insignis
D. medinensis
Epomidiostomum sp.
Eustrongylus gigas
Filaroides bronchialis
F. mustelarum ( =martis)
Gnathostoma spinigerum
Heterakis isolonche
Molineus patens
Muslelivingylus skrjabini
Muellerius capillaris
Perostrongylus pridhami

( = Aelurostrongylus)
Physaloptera sp.
Seurocyrnea sp.
Skrjabingylus nasicola
Soboliphyme baturini
Strongyloides sp.
Trichinella spiralis

6, 8

1,35, 51, 61, 90

30, 31, 35, 58, 61, 68, 79

6, 26

1, 12, 21, 24, 30, 31, 35, 36, 40, 50, 55,

56, 60, 61, 75, 76, 79, 95, 96, 97, 98

19, 22, 35, 42

13, 20, 30, 31, 62

34, 79

1, 6, 25, 30, 31, 40, 48, 61, 88, 89

17, 38, 40, 99

30, 31, 46, 61, 64, 82

46, 74

6, 7, 8, 87, 88

30, 31, 63, 65

34, 41, 79, 91

1, 51, 58, 90

16. 23, 27, 35,40, 100

56 GREAT BASIN NATURALIST Vol. 33, No. 1

Perostrongylus pridhami Anderson, 1962 (Anderson, 1963), by
Dr. Roy C. Anderson (personal correspondence). This parasite was
previously reported by Anderson (1962) in Ontario, Canada. To
our knowledge this is the first report of this species in the United
States.

Thirteen percent of the M. vison were infected with the cestode
Taenia mustelae Gmelin, 1790. Identification of these specimens
was confirmed by Dr. Gerald D. Schmidt (personal correspondence).
This is the first report of adult T. mustelae in mink from Gallatin
and Madison counties in southwestern Montana.

Acknowledgments

The authors express their appreciation to Dr. Roy C. Anderson
for identification of the lung nematodes, to Dr. Gerald D. Schmidt
for identifying the cestode, to Marvin Donahue for providing the
mink viscera, to Dr. David E. Worley for assistance in preparing
the manuscript, and to Donald H. Fritts for photographs.

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giant kidney worm occurring in mink, from the southern counties of
Michigan. J. Parasitol. (suppl.) 25:23.

99. YosHiDA, S. 1934. Observation on Gnathostoma spinigerum Owen, 1836,

cause of esophageal tumor in Japanese mink (Lutreola itatsi itatsi
Temmenick, 1844), with especial reference to its life history. Japan.
J. Zool. 6:113-122.

100. ZiMMERMANN, W. J., E. D. HUBBARD, L. H. SCHWARTE, AND H. E. BlESTER.

1962. Trichinella spiralis in Iowa wildlife during the years 1953 to
1961. J. Parasitol. 48:429-432.

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The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

Volume 33 June 30, 1973 No. 2

THREE NEW SPECIES OF PALMOXYLON FROM THE
EOCENE GREEN RIVER FORMATION, WYOMING

William D. Tidwell,^ David A. Medlyn/ and Gregory F. Thayn^

Abstract. — Silicified, upright axes of Palmoxylon are abundant in the
Green River Formation in Eden Valley, Wyoming. Three new species of Pal-
moxylon, as well as the previously described Palmoxylon macginitiei Tidwell
et al., were collected. These new species are P. edenense, P. contortum, and
P. colei. They are compared to P. macginitiei and other anatomically similar
Palmoxylon species. Palmoxylon edenense appears to be the most abundant
species at this locality.

Silicified remains of palm axes are very abundant in the Green
River Formation in Eden Valley, Wyoming, near the collection site
of Palmoxylon macginitiei (Tidwell et al., 1971). For the most part,
these remains consist of axes in growth position surrounded by tough,
silicified Chlorellopsis algae. However, one axis (P. colei) was col-
lected lying down, as though it had fallen prior to fossilization.
Weathered fragments eroded from these axes are found on the
ground surface throughout the area.

The upright specimens occur in groups of threes and are ar-
ranged in either a triangular or linear alignment. Although the in-
vestigated specimens in these groupings were of the same species
(either P. macginitiei or P. edenense), no rhizomous or soboliferous
connections were uncovered between them.

With the exception of Palmoxylon contortum, the specimens
vary from 6 to 12 inches in diameter and are generally 2 to 3 feet
high. The incomplete horizontal stem of P. colei is nearly four feet
in length. The preservation of the specimens is essentially the same
as that discussed for P. macginitiei (Tidwell et al., 1971 ) .

Palmoxylon edenense Tidwell, n. sp.
Figures 1, 2, 5A, 9
Stem

Central Zone: This zone is characterized by the usual ran dor
arrangement of its vascular bundles. The bundle"^ vary in size from
700 to 900 jx high by 500 to 600 /i wide. There are approximately
85 bundles per cm-. The f/v ratio of these bundles is 3:1, although

'Department of Botany and Range Science, Bnglcdm Young University, Provo, Utah 84002

GREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 1. Pnlmoxylon cdencnsc. Cros.s-sortions illustrating buiullos of flio
(A) subdermal and (B) dermal zones (30X each).

June 1973

TIDWELL ET AL: FOSSIL PALMS

\f- <%- ^V^iTi*^^

•/' *»«ii

yfr --^^

Fig. 2. Palmoxylon edenense. A. Cross-section of the central zone (SOX).
B. Enlarged bundles of the central zone (60X).

64 GREAT BASIN NATURALIST Vol. 33, No. 2

this may vary slightly (3.5:1.0). The fibrous bundle cap is oval to
flabellate, having a shallow median sinus with rounded auricular
lobes. Auricular sinuses are shallow to absent (Figure 2B). The
cap is encased by one or two layers of tabular parenchjTna, whereas
radial parenchyma surrounds the vascular tissue. The bundles are
typically bivascular, with metaxylem elements 78 to 91 jx in diameter
and protoxylem elements varying from 25 to 35 /x. The phloem is
not structurally preserved. The ground tissue is tightly compacted,
consisting of elongated parenchyma cells. Numerous fibrous bun-
dles, 350 per cm", varying in diameter from 52 to 104 /j. are present
but lack stegmata.

SuBDERMAL ZoNE: Bundles of this zone tend to be irregularly
oriented near the central zone and more or less regularly aligned
close to the dermal zone. These bundles are similar in overall shape
to those of the central zone, although their bundle caps are larger.
The f/v ratio of the bundles is 5:1. There are approximately 100
bundles per cm^ ranging in size from 600 to 700 /x high to 400 to
500 /x wide. The reniform bundle cap is usually as high as it is
wide. The bundles are bivascular, and the size and shape of their
vessel elements is similar to those of the central zone.

Cortical and Dermal Zone: The bundles are all regularly
oriented with their caps towards the stem periphery. The bundles
are more tightly compacted than are those of the subdermal zone,
although they are not contiguous. There are approximately 205
bundles per cm-. Their fibrous bundle caps tend to be radially
elongated. The bundle f/v ratio of this zone is 7:1. The bundles,
which are 600 to 800 /i high by 280 to 470 /x wide, are smaller than
those in the other zones. As in the other zones, the fibrous cap is
surrounded by tabular parenchyma, and radial parenchyma en-
sheaths the vascular tissue. The xylem contains two metaxylem ele-
ments that average about 50 /x in diameter. The radial and tabular
parenchyma composing the ground tissue is compact. Numerous
fibrous bundles and leaf traces appear throughout his zone.

Repository. — Brigham Young University, 916 (Holotype)
Locality. — Eden Valley, Wyoming
Horizon. — Green River Formation
Age. — Eocene

Palmoxylon colei, Tidwell, n. sp.
Figures 3, 4, 5B
Stem

Central Zone: The vascular bundles of the central zone are
irregularly oriented and loosely compacted. This zone has approxi-
mately 115 bundles per cm-. The bundles range in size from 500 to
600 fjL in both height and width. Their f/v ratios vary from 2.5:1
to 3:1. The bundle cap fits Stenzel's Rcniforrnia group (Stenzel,
1904) by having rounded auricular lobes with shallow auricular
sinuses. The median sinus is only slightly indented, giving the

June 1973

TIDWELL ET AL: FOSSIL PALMS

^ % t

Fig. 3. PalmoxyLon colei. Cross-sections of the (A) subdermal and ;,B)
dermal zones (30X).

GREAT BASIN NATURALIST

Vol. 33, No. 2

* - m'- V

•^'» ". ~".

■ 'i^Ai

•.Mt]

% V

;'^|

Fig. 4. Palmoxylon colei. A. Cross-section of the central zone (30X). B.
Longitudinal section of a fibrous bundle demonstrating tJie stegmata along its
surface (240X).

June 1973 tidwell et al: fossil palms 67

bundle cap its characteristic kidney shape (Figure 4A). The fibrous
caps are encased by one or two layers of tabular parenchyma, but
the vascular portion is encircled by radial parenchyma. The bun-
dles are typically bivascular and the metaxylem elements are 65 to
85 /x at their widest diameters. The protoxylem, when present, varies
from 30 to 40 [x in diameter. The phloem is not structurally
preserved.

The ground tissue consists of thin-walled, tightly compacted,
tabular and radial parenchyma, and has approximately 290 fiber
bundles per cm-. These bundles exhibit characteristic stegmata, and
their diameters vary from 78 to 120 /x (Figure 5B) .

SuBDERMAL ZoNE: The bundles of this zone are irregularly
aligned near the central zone and regularly aligned near the dermal
zone. There are approximately 160 bundles per cm" in this region.
The bundles are slightly larger than those of the central zone, being
approximately 600 to 750 /x high and 500 to 650 /x wide. They have
an f/v ratio of about 6:1, although this may vary slightly from
bundle to bundle. The bundles are basically the same shape as those
of the central zone, although the fibrous cap may approach a sagitate
form in some. The presence of departing leaf traces that are at-
tached to the vascular portion of several of the bundles is also notable.
The bundles are commonly bivascular but may be trivascular. The
metaxylem vessels are generally about 75 fi in diameter, although
they vary from 65 to 85 /i. Fibrous bundles with their characteristic
stegmata are present.

Cortical and Dermal Zone: The bundles of the cortical and
dermal zone are tightly compacted, with approximately 230 bundles
per cm-. They are regularly aligned and have an f/v ratio of about
8:1. The bundle caps are reniform in shape but are elongated
radially. The caps have rounded lobes and the auricular sinuses of
the cap are shallow to nearly absent. The bundles are approximately
650 [X high and 400 /x wide. There are commonly two, occasionally
three, metaxylem elements per bundle. Each element has an average
diameter of 52 /x.

This specimen was named for its collector, Mr. Walter N. Cole,
of Provo, Utah.

Repository. — Brigham Young University, 917 (Holotype)
Locality. — Eden Valley, Wyoming
Horizon. — Green River, Formation
Age. — Eocene

Palmoxylon contortum Tidwell, n. sp.
Figures 6, 7
Stem

The vascular bundles of all the zones are essentially the same
size and shape, with the only observable difference being the degree
of compaction. The number of bundles per cm- varies from 350 in
the central zone to 500 in the dermal zone. They are tightly ap-

GREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 5. A. Cross-section of a fibrous bundle of Palmoxylon edcnense. B.
ibrous bundle of P. colei illustrating the stegmata surrounding the fibers.

A fib

Both are 160X.

June 1973

TIDWELL ET AL: FOSSIL PALMS

Fig. 6. Palmoxylon contortum. Cross-sections of the (A) dermal and (B)
central zones showing the extreme closeness of the bundles causing their distor-
tion (30X).

GREAT BASIN NATURALIST

Vol. 33, No. 2

J^i^y-W ^^^

I b #i'v /I

.vT.l^

^\'

*•-■- -^

^••t*

-- ^V

,-^ ^»

Fig. 7. Palmoxylon contortum. A. Cross-section of the dermal zone (SOX).
B. Longitudinal section of scalariform thickenings on some protoxylem vessels
(240X).

June 1973 tidwell et al: fossil palms 71

pressed in all three zones but are distorted in the subdermal and
dermal zones due to extreme bundle compaction. Where not com-
pressed, the bundles are blocky to ovate in appearance. They typi-
cally lack auricular lobes and sinuses but have definite median
sinuses. These bundles commonly contain more vascular than
fibrous tissue, with an f/v ratio between 1:1 and 1:1.5. Both the
vascular and fibrous cells are surrounded by tabular parenchyma.
Bundle size varies from 320 to 400 jx high and 300 to 500 /x wide.
The bundles are characteristically bivascular, with the large rneta-
xylem elements averaging about 55 /x in diameter. The ground tissue
is tightly compact, consisting mostly of thin-walled columnar paren-
chyma. Numerous sclerotic bundles ranging from 40 to 50 /x in
diameter, without stegmata, are present. Specimens of this species
were donated by Mrs. Marian Whitehead of Salt Lake City, Utah.
They were approximately two inches in diameter.

Repository. — Brigham Young University, 918 (Holotype)
Locality. — Eden Valley, Wyoming
Horizon. — Green River Formation
Age. — Eocene

Comparisons

Palmoxylon macginitiei Tidwell et al. is similar in general ana-
tomical features to both P. edenense and P. colei. P. contortum is
dissimilar due to the disfiguration of the bundles and to the larger
number of the vascular bundles per cm^ and its overall size. All of
these species have scalariform perforation plates on their metaxylem
elements.

In order to delineate the above species, to give additional infor-
matipn, and to correct an error in the previous description of Palmo-
xylon macginitiei, a brief description of that species is included. P.
macginitiei is characterized by the shape and size of its fibrovascular
bundles (Figure 8A). They are oval, rarely rounded, and have a
reniform bundle cap. The vascular portion of the bundle is blocky
in appearance and is typically bivascular with tyloses common in the
metaxylem elements (Figure 8B). The metaxylem vessels have
scalariform perforation plates. The bundle size varies from 600 to
700 /A high by 450 to 500 /x wide in the central zone to approximately
420 /x high by 600 /x wide in the cortical and dermal zone. The
numlDer of bundles per cm^ ranges from 104 in the central portion
to 212 in the outer dermal zone.

Palmoxylon edenense, although similar to P. macginitiei, should
be considered a separate species because of the difference in degree
of bundle compaction and bundle shape. P. edenense has an fV ratio
of 3:1 to 3.5:1 in the central zone, whereas the ratio for P. mac-
ginitiei is 1.5:1 to 2:1. Therefore, the bundle caps of P. edenense
are larger than those for P. macginitiei. Hence, more definite
auricular lobes and sinuses are present than in P. macginitiei, and the
overall outline of the vascular portion is more rounded and not as
blocky in appearance. The fibrovascular bundles of P. edenense are

GREAT BASIN NATURALIST

Vol. 33, No. 2

. ^^''T)^-M

Fig. 8. Palmoxylon macginitiei. A. Cross-section of the central zone (30X).
B. Enlarged cross-section of bundles showing tyloses in the vessels (60X).

June 1973 tidwell et al: fossil palms 73

less compacted, with 85 bundles per cm" in the central zone to 205
bundles per cm- in the cortical and dermal zone.

Palmoxylon colei is unique in that it possesses stegmata, an ana-
tomical structure which sets it apart from P. macginitiei. It also
differs from the latter in bundle shape, with less vascular tissue in
the central zone. Also both P. colei and P. edenense characteristically
lack tyloses in the metaxylem elements.

Palmoxylon edenense is similar to P. macginitiei. Therefore, the
comparisons between P. macginitiei and other previously described
Palmoxylon species (Tidwell et al., 1971) would also be valid for
P. edenense. Although further study has shown that P. crassipes
Unger is similar, it differs in that it has a complanate bundle cap
and less vascular tissue in the central zone. The bundle cap of P.
edenense tends to be less rounded and more flabellate with a deeper
median sinus. P. hungaricum Greguss (Greguss, 1959) shows a
close affinity to P. edenense but has larger and more widely spaced
fibrovascular bundles.

The presence of stegmata (stegmata is an anatomical feature
found only in fossil palms [Stenzel, 1904]) in Palmoxylon cold
restricts the number of similar species for comparison. Stenzel
(1904) described three species exhibiting stegmata: P. densum, P.
confertum, and P. astrocaryoides. These all differ from P. colei in
their characteristic bundle shape, with P. densum also having dis-
tinctly smaller bundles. Sahni (1964) described P. pondicherriense,
a palm with stegmata from India. This species is dissimilar from
P. colei in that the bundles in the latter are less numerous and that
they have a reniform bundle cap instead of the lunarian form of
P. pondicherriense.

Palmoxylon contortum is a unique fine-bundled palm, and only
a few species of Palmoxylon are even similar. P. liebigianum Shenk,
which Sahni regards as synonymous with P. kvishna, also exhibits
compact bundles, 150 per cm- in the central zone to 250 per cm- in
the dermal zone, although the fibrovascular bundles of P. liebigianum
are not appressed and disfigured, and its vascular bundles have
lunarian-shaped bundle caps rather than reniform. P. ceylanicum
Unger was considered by Stenzel (1904) as a variety of P. liebigi-
anum. However, Sahni (1964) concluded that P. ceylanicum should
be maintained as a separate species. He based this on the vascular
bundles of P. ceylanicum being thinner and more crowded than in
P. liebigianum. P. contortum differs from the above by its distinc-
tive bundle shape, its degree of compaction, and its characteristic
bundle distortion.

Palmoxylon colei varies from P. simper i Tidwell, P. pristina
Tidwell, and P. gustavsoni Tidwell et al. by having stegmata which
the latter species lack; and P. contortum varies from them by its
smaller size and by having more compressed, distorted bundles.

Discussion

In comparing Palmoxylon edenense and P. macginitiei with P.
simperi Tidwell and P. gustavsoni Tidwell et al., it is a case of com-

74 GREAT BASIN NATURALIST Vol. 33, No. 2

paring species representing the basal portion of palm stems, or the
so-called "stemless" varieties, from the Green River Formation with
the distal portions of P. simperi and P. gustavsoni. Tomlinson and
Zimmermann (1967) illustrated the different vessel types they had
mascerated from a species of Sabal. The base of the stem had rather
long vessels with scalariform perforation plates similar to the Eden
Valley taxa; whereas those near the middle and top of the same axis
had short vessels with simple pores like P. simperi and P. gustavsoni.
P. simperi and P. gustavsoni, whose stem bases have not been ob-
served, are from the same geographical area in central Utah (Tid-
well et al., 1972). Scott et al. (1972) implied that all specimens
from this area are of the same species; however, they most probably
represent the middle to upper portions of P. simperi and P. gustav-
soni. Thus, in comparing these latter two species to each other at
about the same stem elevation and from the same zone (central),
they are quite distinct and probably portray more than a delineation
between form species.

As further discussed by Tomlinson and Zimmerman (1967), the
apical portion of a palm stem is always softer than the basal part.
Consequently, when dead, they generally rot from the apex to base.
The anatomy of the upper portions of the stem of living palms is
easily sectioned and thus studied, whereas the basal portions are most
often fossilized. Therefore, the matching of the anatomy of fossil
and living forms becomes a dubious procedure, and comparisons be-
tween the different parts of the stem may result in rather inaccurate
conclusions. Therefore, comparisons between species representing
the basal portion of the palm stems with those of the upper part
would not be conclusive.

The specimens of Palmoxylon macginitiei and P. edenense ap-
pear to be complete palm axes. They do not indicate any truncation
of the stems as would be anticipated if the stem had been originally
taller (Figure 9). The bases of these specimens are surrounded with
roots, whereas the middle and upper portions of these same axes have
numerous attached petiole bases. The apices of these specimens
consist of overlapping petiole bases. In progressing up the stem,
each successive base becomes smaller and closer to the center of
the axis.

These stems may represent either very young stems or the so-
called "stemless" palms similar to Nipa, some species of Acantho-
cocos and Serenoa. Several forms of Palmae do not have any trunks
above ground and thus are similar to the specimens from Eden
Valley. In Serenoa repens, leaves appear to come right out of the
ground, whereas Attalea cohune grows for many years before its
trunk shows and has been mistaken as trunkless (McCurrah, 1960).
Living Nipa palms balance enormous stemless rosettes on the treach-
erous semiliquid mud of estuaries by means of a stout horizontal
trunk (Corner, 1966). These spread by means of branching root-
stocks. In the Philippine Islands, a Nipa marsh has been reported
covering approximately 20,000 acres (McCurrah, 1960).

The particularly numerous upright specimens from Eden Valley

June 1973

TIDWELL ET AL: FOSSIL PALMS

Fig. 9. A specimen of Palmoxylon edenense drawn after collecting. Note
the attached petioles. Roots are present in the lower portion of the specimen.

76 GREAT BASIN NATURALIST Vol. 33, No. 2

appear to represent the stemless types rather than the younger forms
of taller trunked species. With the number of specimens at this
locality, it would seem that intermediate stages of the trunked forms
would also be present and these have not been observed. However,
it is anticipated that further collecting will provide additional infor-
mation concerning these forms and their possible placement.

Acknowledgments

The authors wish to acknowledge the assistance of the following:
Naomi Hebbert and Paul Smith for aiding in preparing the illustra-
tions and collecting, Mrs. Marian Whitehead of Salt Lake City and
A. Daniel Simper of the University of California at Davis for their
help in collecting and thin-sectioning of the specimens, Mr. Walter
Cole of Provo for donating a specimen of Palmoxylon colei, and Dr.
J. Keith Rigby of the Department of Geology and Dr. S. R. Rushforth
of the Department of Botany at Brigham Young University for re-
viewing the manuscript.

References

Corner, E. J. H. 1966. The natural historj- of palms. Univer. Calif. Press,
Berkeley and Los Angeles.

Greguss, p. 1959. A palmtrunk from the Miocene Coal Basin of Salgotarjan.
Palaeobotany 8:19-21.

McCuRRAH, J. C. 1960. Palms of the world. Harper and Brothers, New York.

Sahni, B. 1964. Revisions of Indian fossil plants. Part III — Monocotyledons.
Monogr. Birbal. Sahni Inst. Palaeobot. 1:1-89.

Scott, R. A., P. L. Williams, L. C. Cr-mg, E. S. B.arghoorn, L. J. Hickey, and
H. D. MacGinitie. 1972. "Pre-Cretaceous" angiosperms from Utah: Evi-
dence for Tertiary age of the palm woods. Amer. Jour. Bot. 59:886-896.

Stenzel, K. G. 1904. Fossile Palmenholzer. Beitr. Palaeont. Geol. Ost. — Ung.
16:107-287.

Tidwell, W. D., a. D. Simper, and D. A. Medlyn. 1971. A Palmoxylon from
the Green River Formation (Eocene) of Eden Valley, Wyoming. Botanique
2:93-102.

Tidwell, W. D., D. A. Medlyn, and G. F. Thayn. 1972. Fossil palm materials
from the Tertiary Dipping Vat Formation of Central Utah. Great Basin Nat.
32:1-15.

Tomlinson, p. B., and M. H. Zimmermann. 1967. The wood of monocotyle-
dons. Bull. Intemat. Assoc. Wood Anatomists 2:4-24.

ON THE TAXONOMIC STATUS OF PLATYPODIDAE AND
SCOLYTIDAE (COLEOPTERA)

Stephen L. Woodi

Abstract. — The reduction of Plat;yT)odidae and Scolytidae to subfamihes of
Curculionidae by certain recent authors is questioned. Several fundamentally im-
portant anatomical characters that could not possibly have been derived from
Curculionidae are cited. Platypodidae (including Coptonotidae) is recognized
provisionally as a derivative of Scolytidae; Scolytidae, with subfamilies Hylesi-
ninae and Scolytinae (including Ipinae), is also recognized as a family. Proto-
platypus, new genus, for P. vetulus, n. sp., a primitive platypodid from New
Guinea, and Proiohylastes, new genus, for Pr. annosus, n. sp., a primitive scolytid
from Queensland, are described.

Apparently commencing in 1954 with Crowson (1967:155), a
number of coleopterists have reduced the traditionally recognized
families Scolytidae and Platypodidae to the rank of subfamily within
the Curculionidae without establishing a foundation for this action.
The recent generic classification of the Platypodidae (Schedl, 1972)
neglected either to acknowledge or to respond to this change.

In reviewing more than a dozen classifications of higher cate-
gories within the Curculionoidea, the most striking feature noted in
them is the lack of agreement on even the most fundamental divi-
sions of the group, apparently due to a lack of detailed knowledge
on the morphology of this enormous superfamily. It would appear
that only Crowson's (1967) major division, based on separate or
confluent gular sutures, is the only phylogenetically sound division
of the group that has been presented. His classification of families
in the first division (couplets 1 to 5) is reasonable (although the
Oxycorynidae and Proterhinidae are unknown to me). His classifi-
cation of the second division, groups having only one median gular
suture (couplets 6 to 8), is questioned. The basis for this question
is an unpublished and incomplete comparative anatomical study of
the Curculionoidea that was started many years ago but interrupted
due to the lack of specimens for dissection in critical groups. The
following comments relating to the pregula and pregular sutures
were drawn from that study.

In the superfamily Curculionoidea five families (Crowson, 1967)
have at least the posterior indications of widely separated gular
sutures. In the Belidae these sutures continue separately to near the
ventral apex of the rostrum, where they diverge and direct their
course to or at least toward the anterior tentorial pits above the an-
terior articulation of the mandible as in other insects. In the An-
thribidae these sutures are obsolete except for minute irregularities
on the margin of the occipital foramen; in Nemonychidae and ap-
parently in Oxycorynidae (not seen) and Proterhinidae (not seen),
separate sutures (Crowson, 1967: Figures 201-202) extend anteriorly

'Department of Zoology, Brigliam Young University, Provo, Utah 84602. Scolytidae Contribii-
tion No. 49.

78 GREAT BASIN NATURALIST Vol. 33, No. 2

to the posterior tentorial pits. In all other Curculionoidea the gular
sutures are confluent, with only one median suture (usually) visible
from near the occipital foramen to the single, median, posterior
tentorial pit at the ventral base of the rostrum. Apparently all
workers have overlooked the minute postgula on the margin of the
occipital foramen at the base of the gular suture (Figures 1-19).
Some workers, realizing that a pregula should be present (c.f. Hop-
kins, 1909:16, and Stickney, 1923), either created pregular sutures
for their illustrations of curculionids or misinterpreted longitudinal
ridges on the rostrum as being pregular sutures. In a survey of
several hundred genera of Attelabidae, Brenthidae, Apionidae, and
Curculionidae (s. str.) I was unable to find any representative hav-
ing pregular sutures accompanied by internal ridges that resulted
from the inflection of the cuticle to form those sutures. However,
pregular sutures are conspicuously present in all Platypodidae and
in all Scolytidae except a few Micracini, etc., that bore into exceed-
ingly hard wood (in which case the gular suture is also obliterated).

It apparently has been traditional among those coleopterists who
have considered the matter (Crowson, 1967:155; Schedl, 1972) to
presume that the Scolytidae were derived from the tribe Rhyncolini
of the subfamily Cossoninae and that the rostrum of the ancestral
form was lost when scolytid habits were established. Yet in no
truly primitive platypodid or scolytid is there a rostrum resembling
that of Cossoninae. Furthermore, no representative of the Rhynco-
lini or of any other cossonid known to me (including an apparently
unnamed Australian cossonid genus with absolutely no rostrum
whatever and large mandibles) has pregular sutures (Figure 4).
In this latter character I have found no intergradation, either ex-
ternally or in the much more complex internal structure. The recog-
nition of this fact suggests that other indicators of phyletic relation-
ship be examined.

Primitive Platypodidae, primitive representatives of all primitive
tribes of Scolytidae, and many Cossonini have interstriae 10 on the
elytra broad throughout its length to near the elytral apex. In all
Rhyncolini known to me the portion of interstriae 10 posterior to the
level of the hind coxa is strongly constricted or entirely obsolete.
Therefore, based on this character in living material examined,
there is no possibility that the Rhyncolini could have given rise to
either the Platypodidae or Scolytidae.

Crowson (1967:158) indicated that all adult Curculionoidea hav-
ing one gular suture, except Attelabidae, have the adult maxillary
palpus 2- or 3-segmented. Browne (1971:49) reported a 4-segmented
maxillary palpus in Austroplatypus BrowTie, a platypodid. It is
very doubtful that this geiuis was derived from an ancestral form
having a 3-segmented maxillary palpus.

In all Rhyncolini and most (all?) Cossoninae known to me there
is a conspicuous spine on the margin of the oral fossa that arises
between the posterior margin of the mandible and the base of the
maxilla (Figure 4). There is no com])arable structure or irregularity
in this area in Platypodidae or Scolytidae (Figures 12, 14, 19).

June 1973 wood: platypodidae and scolytidae

1. Stenoscelis

5. Hylurgops

2. Stenoscelis

6. Hylurgops

7. Hylurgops

10. Schedlarius

3. Stenoscelis

11. Schedlarius
12. Gnathotrupes

8. Platypus

4. Rhyncolus

Figs. 1-12. Head capsules of beetles with the tentorial appartus indicated
by dotted lines: 1-3, Stenoscelis brevis (Boh.), Cossoninae, 1 lateral, 2 posterior,
and 3 dorsal aspects; 4, Rhyncolus knowltoni (Thatcher), Cossoninae, ventral
aspect; 5-7, Hylurgops rugipennis (Mannerheim), Hylesininae, 5 lateral, 6 pos-
terior, and 7 dorsal aspects; 8, Platypus lucasi Chapuis, Platypodinae, dorsal
aspect; 9-11, Schedlarius mezicanus (Duges), Coptonotinae, 9 lateral, 10 posterior,
and 11 dorsal aspects; 12, Gnathotrupes sp., Scolytinae, ventral aspect.

GREAT BASIN NATURALIST

Vol. 33, No. 2

Figs. 13-19. Head capsules of beetles with the tentorial appartus indicated
by dotted lines: 13-16, /p? mcxicanus (Hopkins). Scolytinao. H lateral. 11 ven-
tral, 15 posterior, and 16 dorsal aspects; 17-19, Platypus lucasi Chapuis. Platy-
podinao. 17 lateral, 18 posterior, and 19 ventral aspects.

Kuschel (1966:6) suggested that a series of sj)iiies on the lateral
margins of the tibiae in Scolytidae and in the Araucariini (Cosso-
ninae) indicate a relationship between these groups. Evidently it
was not recognized that the most primitive genera of Scolytidae lack
these spines and have tibiae more nearly like the (>ossonini or even
like certain Brenthidae. The supernumerary tibial spines in these

June 1973 wood: platypodidae and scolytidae 81

groups apparently were acquired independently as they adapted to
a common niche and, therefore, are not primitive. Kuschel also
called attention to the similarity in habits of the Araucariini and
Scolytidae. In the two species of Araucariini (presumably Coptoco-
cynus spp.) examined in the field, the habits do not resemble in any
way those of more than 2000 species (about 30 percent of the known
fauna) of Scolytidae I have studied in the field. The habits of certain
neotropical Rhyncolini (unidentified) are infinitely more similar to
primitive scolytids.

Larval characters that separate the Scolytidae from Curculionidae
have not been reported, presumably due to inadequate research
rather than to the absence of characters. Most larval Platypodidae
have the clypeus reduced or absent and the labrum somewhat en-
larged. The cercus-like processes in Trachyostus (Browne, 1967:
Figure 20) and the tenth abdominal segment of Dolgopygus (Browne,
1967: Figure 23) require investigation.

It is my contention that the Scolytidae and Platypodidae repre-
sent a phyletic line that diverged from other Curculionoidea before
pregular sutures were lost and before a rostrum developed. The
Cossoninae (Figures 1-4) are true curculionids in the structure of the
gular area, tentorial apparatus, head, legs, and body form, and
probably resemble scolytids largely because they independently oc-
cupied the same or a similar niche. While the scolytids and platypo-
dids clearly fall within the Curculionoidea, it is as logical to recog-
nize them as an independent group as it is any other family within
this superfamily. There is great difficulty, however, in separating
the Scolytidae from the Platypodidae, and I find it difficult to give
independent family status to the latter group.

In order to emphasize the significance of the above items, it is
necessary to describe the most primitive platypodid and the most
primitive hylesinine scolytid known to me.

Protoplatypus, n. gen.

The phloeophagous habit and normal tarsi of this remarkably
primitive genus suggest that it should be placed in the family Scoly-
tidae; however, the head, tibiae, pronotum, and other characters in-
dicate a closer relationship to primitive Platypodidae. Although not
closely related to either genus, its phylogenetic position probably lies
between Mecopelmus Blackman (Platypodidae) and Craniodycticus
Blandford (Scolytidae). For convenience of reference I tentatively
place it in the Mecopelmini (Platypodidae) .

Characters of particular significance in phylogeny found in this
genus include the cylindrical, 3-segmentod maxillary palpus, an
antenna intermediate between that of Platytarsulus (Platypodidae)
and Craniodycticus (Scolytidae), a protibia intermediate between
that of Schedlarius (Platypodidae) and Protohylastes (Scolytidae),
an eye similar to that of platypodids, and tarsi similar to those of
scolytids. The phloeophagous and polygamous habits are, for the
most part, typical of scolytids.

GREAT BASIN NATURALIST

Vol. 33, No. 2

Description. — Head about as in Schedlarius Wood; eyes sub-
circular, moderately convex; pregula rather large, as in most Platy-
podidae; antennal scape long, slender, funicle 5-segmented, club
rather small, somewhat flattened, entirely devoid of sutures, gla-
brous except for a marginal fringe of abundant, short hair (much
as in Platytarsulus Schedl but much more restricted to actual mar-
gin). Pronotum elongate, sides constricted as in most Platypodidae;
precoxal area of prosternum elongate, almost half as long as entire
prosternum, precoxae small, very widely separated. Scutellum sub-
acutely pointed behind. Elytral bases rounded, not precipitous,
striate, interstriae 10 attaining declivital area; posterior area decli-
vous, sculpture simple. Protibiae of platypodid type; tarsi with seg-
ment 1 only slightly longer than 2 or 3, very slightly shorter than 5,
entire tarsus slightly longer than tibia, all segments cylindrical.

Type species. — Protoplatypus vetulus Wood, described below.

Protoplatypus vetulus. n. sp.
Figures 20, 21, 26

This species is distinguished from other known forms by char-
acters summarized in the above description of the genus. It could
be placed in either the Platypodidae or Scolytidae, depending upon
which characters are emphasized.

Male. — Length 1.2 mm (paratypes 1.2-1.5 mm), 4.0 times as
long as wide; color light brown.

Frons strongly, rather evenly arched from epistoma to vertex,

Figs. 20-21. Protoplatypus vetulus Wood: 20, dorsal aspect; 21, lateral aspect
of head and prothorax. leg omitted. Figs. 22-24, Protohylastes annosus Wood:
22, lateral aspect; 23, dorsal aspect; 24. anterior aspect of left metathoracic tibia.

June 1973 wood: platypodidae and scolytidae 83

rather broadly convex, somewhat intermediate between Schedlarius
and Coptonotus Chapuis; surface reticulate, with sparse, fine, shal-
low punctures; vestiture very sparse, hairlike; eye broadly oval to
subcircular, mioderately convex; pregula rather large, as in Platy-
podidae; maxillary palpi cylindrical, 3-segmented.

Pronotum 1.5 times as long as wide; widest on basal third, sides
conspicuously constricted just in front of middle, anterior margin
only slightly narrower than base; surface shining, reticulate at base
and sides, longitudinally, subreticulately strigose on discal area,
punctures minute, shallow, moderately close. Glabrous.

Elytra 2.3 times as long as wide, 1.6 times as long as pronotum;
sides straight and parallel on basal three-fourths, broadly rounded
behind; scutellum acute; elytral bases rather weakly, not precipi-
tously, elevated; entire surface strongly reticulate; striae not im-
pressed, punctures small, moderately deep, spaced within row by
about two diameters of a puncture; interstriae twice as wide as striae,
punctures very small, widely, regularly spaced. Declivity confined
to posterior fourth, broadly convex; striae about as on disc; interstriae
1 and posterior half of 9 moderately, continuously elevated and meet
at sutural apex, 3 less strongly elevated and ending before attaining
9. Vestiture of sparse, very minute, almost scalelike setae.

Female. — Similar to male except frons with a triangular area
on lower two-thirds smooth, shining, its upper margins marked by
an irregular row of small punctures; scape with setae near apex
distinctly longer.

Type Locality. — Five miles or 8 km NW Bulolo, Morobe, New
Guinea.

Type Material. — The male holotype, female allotype, and 96
paratypes were collected at the type locality in the LATEP logging
area, on 9-VIII-72 from the bole of a recently cut Har pallia pedi-
cellaris, by S. L. Wood. The parental galleries were in the cambium
region but did not engrave the wood; they were of the radiate type,
and the beetles appeared to be polygamous. Larval mines were
parallel to the grain of wood and were visible on the inner surface
of peeled bark.

The holotype, allotype, and several paratypes are in the Aus-
tralian National Collection at Canberra; the other paratypes are in
the British Museum (Natural History) and in my collection.

Protohylastes, n. gen.

This genus superficially resembles Hylurgops LeConte or Pseudo-
hylesinus Swaine, except that the tibiae are completely different
from any other genus of Hylesininae and the protibiae are more
nearly like a curculionid (Curculionoinae) than a scolytid. The
bases of the elytra are similar to those of Hylurgops, without a defi-
nite marginal row of crenulations, except that the submarginal crenu-
lations are even more poorly developed.

84 GREAT BASIN NATURALIST Vol. 33, No. 2

Diagnosis and Description. — Head much as in Pseudohyle-
sinus; eye oval, short; antennal scape short, funicle 7-segniented,
shorter than scape, club as long as scape, small, subconical but dis-
tinctly flattened, entirely devoid of sutures. Prothorax about as in
Pseudohylesinus except coxae rather widely separated, an acutely
elevated ridge extending from anterolateral margin of coxa to antero-
lateral margin of prothorax (as in Hylurgops). Elytra much as in
Hylurgops except basal margins not armed and interstriae 10 ex-
tends to declivity. Anterior tibia slender, entirely unarmed on
margins, a terminal spine next to tarsal insertion at center of apex,
a very small spine on lateral and median apical angles; meso- and
metathoracic tibiae similar except obliquely truncate at apex, with a
very short, blunt spine on both inner and outer angles (Figure 24).
Tarsal segments 1 and 5 each about as long as 2 and 3 combined,
3 broadly bilobed.

rvPE-SPECiEs. — Protohylastes annosus Wood, described below.

Protohylastes annosus, n. sp.
Figures 22-25

This species resembles a very large Pseudohylesinus, but it is
distinguished from all previously known Scolytidae by the very
different tibiae.

Adult. — Sex not determined. Length 9.8 mm, 2.6 times as long
as wide, color very dark brown, vestiture pale.

Frons convex, epistomal area transversely impressed, a small,
median impression at upper level of eyes; surface smooth, shining,
punctures moderately coarse, close, their interiors apparently reticu-
late; subglabrous, a few scales on lateral margins, a few hairlike
setae in epistomal area. Eye oval, about 1.5 times as long as wide.
Antenna as described above.

Pronotum 0.80 times as long as wide; widest at base, sides arcu-
ately converging to a rather strong constriction just behind very
broadly rounded anterior margin; basal margin bisinuate; surface
smooth and shining on most of discal area, becoming reticulate in all
marginal areas and on sides; punctures moderately coarse, close,
deep, weakly subcrenulate at base and in lateral areas. Vestiture
of rather sparse scales, each scale about six times as long as wide.

Elytra 1.7 times as long as wide, 2.3 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather narrow-
ly rounded behind; striae weakly impressed, punctures small, close,
deep; interstriae twice as wide as striae, surface finely rugose-
subreticulate, with numerous small, confused, transverse crenula-
tions, each up to one-third width of an interstriae; crenulations near
basal margins much as in Hylurgops, but margin without a definite
row of crenulations. Declivity moderately steep, occupying posterior
third of elytra, broadly convex, with areas at apices of interstriae
5-7 slightly elevated; striae more deeply impressed; interstriae
10 reaching declivity. Vestiture of rather abundant, small scales,

June 1973 wood: platypodidae and scolytidae 85

each scale very slightly longer than wide, their color pale except
small patches of dark brown on declivity.

Type Locality. — Eungella National Park, Queensland, Aus-
tralia.

Type Material. — The unique holotype was taken at the type
locality on lO-XII-65, by G. Monteith. It is in the Queensland
Museum.

Platypodidae

Most platypodids belong to a compact subfamily of highly modi-
fied ambrosia beetles. In fundamental structure they differ from
scoiytids only in degree, not in the kind of characters represented.
They share with primitive scoiytids the same basic head structure,
including a large, well-defined pregula (Figures 8, 17-19). The
mouthparts in most of them are highly modified to the ambrosial
habit, but primitive forms (Coptonotinae) are comparable to those of
scoiytids. Virtually identical antennae and pronota are found
among primitive forms in the two groups. The protibiae at first
appear unique until it is noted that tubercles and ridges appear on
the posterior face of the protibiae of some representatives of almost
all groups of scolytid woodboring ambrosia beetles. If the minute
ridges and tubercles on the posterior face are ignored and the spines
and denticles on the apical and lateral margins are emphasized, or if
only bark-infesting genera are examined, a graded series of steps
apparently bridging the gap between primitive scoiytids and primi-
tive platypodids can be demonstrated (Figures 25-33). Since socketed
tibial spines characteristic of most scoiytids occur only in that group
and not in platypodids, only unsocketed spines are considered primi-
tive here.

When the posterior face of the prothoracic tibiae are viewed
from exactly the same aspect it is noted that the tarsal insertion is
visible near the apex on all Platypodidae, including Coptonotinae, on
Protohylastes, and on all Scolytini. In Protoplatypus, Mecopelmus,
and Protohylastes the tarsus evidently is capable of movement
through an arc of almost 180 degrees from the lateral to posterior to
mesal positions. In Coptonotus and most Scolytini the action is simi-
lar but the arc is somewhat smaller. In some Scolytini the tarsal
insertion is closer to or even on the apical margin, and greater free-
dom of movement may occur. In Schedlarius the lateral margin of
the tarsal insertion is slightly elevated, thus restricting tarsal action
more nearly to the posterior to mesal arc; in Platypodinae this arc
evidently is even more strongly restricted. In Aricerus, Scolytopla-
typus, and many of the higher Scolytinae, the tarsal insertion is on
the apical or lateral margin, and tarsal action apparently is through
an arc from the lateral to anterior and possibly posterior positions.
In most Hylesininae the tarsal insertion is clearly on the anterior
face, and tarsal action is restricted to an arc from the lateral to
anterior position or less. A submarginal tubercle on the posterior
face near the tarsal insertion in primitive forms appears to have
special significance; it is absent in Scolytini and Cossoninae.

GREAT BASIN NATURALIST

Vol. 33, No. 2

Coptonotus

31.
Scolytodes

32. ^ 33.

Diamerus Camptocerus

Figs. 25-33. Posterior face of prothoracic tibiae: 25, Protohylastes annosus
Wood, with tarsus. Hylastini; 26, Protoplalypus veiulus Wood, with tarsus,
Coptonotinae; 27, Mecopelmus zeteki Blackman, with first segment of tarsus,
Coptonotinae; 28. Schedlarius mexicanus Duges, wdth first segment of tarsus,
Coptonotinae; 29. Coptonotus cyclops Chapuis, witli first and second tarsal seg-
ments, Coptonotinae; 30, Tricolus peltatus Wood, Scolytinae; 31, Scolytodes sp.,
Scolytinae; 32, Diamerus impar Chapuis. Hylesininae; 33, Camptocerus auri-
cnrnus Blandford, Scolytinae. All drawings were made at different scales so as to
be reproduced at a uniform size.

In platypodids the first tarsal segment is supposed to be as long as
the remaining segments combined; in some it is actually less than
half as long (Schedl, 1939). Representatives of the Coptonotinae
(Coptonotidae of Schedl) have the tibiae and tarsi as in some Platy-
podinae (Platypodidae of Schedl) except that there is greater vari-
ability, particularly in the length of the first tarsal segment. One
of these, Schedlarius^ has wood-boring habits, but the larvae, unlike
Platypodinae, form independent mines that wander through deep
xylem tissues; fungal activity definitely associated with the beetles
alters the character of the wood, although there is no mycelial growth
in the tunnels that could be used for food as occurs with ambrosia
beetles. Another coptonotid, Mecopelmus Blackman, is phloeopha-
gous; a large nuptial chamber is formed by the ]iarcnts in which
clusters of eggs are deposited, and larvae then form individual mines
that wander in a radiating pattern from this chamber. The genus
Protoplalypus described above is anatomically rather closely allied

June 1973 wood: platypodidae and scolytidae 87

to Mecopelmus except that the tarsi are typical of scolytids; as noted
above, it is phloeophagous and polygamous.

In most male platypodids, abdominal tergum 8 is of reduced size
and is largely or entirely covered by tergum 7 as in the female. In
Schedlarius, male tergum 8 is more nearly intermediate between
the reduced state seen in other platypodids and the normal condi-
tion of most scolytids and many curculionids. A reduced male ter-
gum 8 also occurs in the scolytid tribes Ipini (Ips, etc.) and Carpho-
dycticini {Craniodycticus, etc.). It is doubtful that this reduction
in these two tribes indicates a close relationship to platypodids.

The larvae of platypodids and scolytids are inadequately known,
but most of them can be segregated using features of the labrum and
clypeus. In platypodids the clypeus is broad and strikingly reduced
in length or entirely absent and the labrum is proportionately
lengthened. In Schedlarius and Protoplatypus of the Platypodidae
and Carphodycticus Wood in Scolytidae, the labrum and clypeus are
intermediate in size and shape.

In my opinion the platypodids are an aberrant group of am-
brosia beetles, as are most other groups of ambrosia beetles within the
family Scolytidae, that arose long after scolytid-platypodid char-
acters had been well established. The ambrosial habit has arisen
repeatedly within the Scolytidae; for example, Camptocerus Erich-
son (Scolytini), Scolytoplatypus Schaufuss (Scolytoplatypini), Both-
rosternus Eichhoff (Bothrosternini), Hyleops Schedl (Hylesinini),
and numerous genera in the more highly evolved tribes Xyleborini,
Xyloterini, and Corthylini. For this reason, the appearance of the
ambrosial habit very early in scolytid phylogeny is not unusual.

Every character on which the family Platypodidae is based,
whether anatomical, behavioral, or ecological, intergrades with the
Scolytidae. The only justification I see, at present, for retaining it as
a family separate from Scolytidae is tradition and the fact that most
forms encountered in the field are easily recognized.

Scolytidae

As conceived here, the family Scolytidae consists of the two sub-
families Hylesininae and Scolytinae (including Ipinae). The adult
forms of Scolytidae ( 1 ) have a definite pregula and pregular sutures,
(2) lack a spherical head and rostrum, (3) have a distinctive man-
dibular articulation (quite different from curculionids?), (4) never
have a petiolate point of labial articulation, and (5) have tibiae that
are adapted to a bark- or wood-boring habit and, in primitive forms,
are no more similar to Cossoninae than they are to those of certain
Brenthidae. In part, the similarity between Scolytidae and Rhyn-
colini is due to convergence or more probably to parallel evolution
that commenced when both groups entered part of the same broad
ecological niche. The short pseudorostrum of the Hylastini and allied
forms of Hylesininae appears to have been acquired independently
and differs in fundamental structural detail from that seen in
Curculionidae (Figures 1-7).

88 GREAT BASIN NATURALIST Vol. 33, No. 2

The division of the Scoiytidae (including Platypodidae) into
three major lines of development occurred very early m its phyletic
history but well after the basic characteristics of the group were
established. The earliest living genera representing those lines, Pro-
toplatypus and possibly Mecopelmus for Platypodidae, Protohylastes,
and, doubtfully, certain Cnemonyx (galeritus Eichhoff) for Hyle-
sininae, could appropriately be placed as primitive Scolytinae since
they are intermediate and lack some of the most diagnostic characters
of the groups I presume they represent.

The argument for the above classification, and also for the re-
tention of Platypodidae and Scoiytidae as families, must be con-
sidered tentative and biased. It is based largely on a limited number
of external adult characters. The larvae, habits, and internal char-
acters of most primitive genera of concern are totally unknown.
Collecting experience in tropical areas has demonstrated that speci-
mens of these primitive genera are exceedingly rare and suggests
that several more equally rare genera await discovery. It should
also be mentioned that in a review of this problem careful attention
should be given to the sporadic occurrence of a very short antennal
scape, broadly bilobed tarsi, scalelike vestiture, gnathal, and other
characters commonly found in primitive genera of these groups.
The significance of a complete gula, with two totally separate gular
sutures found in several species of the neotropical genus Gnatho-
trupes (Scoiytidae) (Figure 12), and the possible cercus-like struc-
tures and the tenth abdominal segment of larval platypodids, cited
above, must also be investigated. Convergent and parallel evolution
among wood-boring Curculionoidea must be examined very carefully
in order to sift the meaningful from the misleading superfluous char-
acters found in these families.

Among primitive Coleoptera the tentorial structure arises from
two separate tentorial pits on the gular sutures about midway be-
tween the foramen magnimi and the oral fossa (Stickney, 1923).
From these pits the posterior tentorial bridge arises, forming a clear-
ly identifiable internal landmark. The anterior tentorium extends
forward from this point for some distance, then branches to form (1)
the dorsal arms that extend to but do not fuse with the wall of the
head near the anteromesal margin of the eye (connected by liga-
ments only) and (2) the anterior arms that extend to the anterior
tentorial pits near the anterior articulation of the mandibles. In all
Curculionoidea having an anterior tentorium the anterior arms ex-
tending to the anterior tentorial pits are entirely absent. The dorsal
arms vary from well-develo[)ed to obsolete. The posterior tentorium
in Curculionoidea is carried inward on a Y-shaped apodeme having
a median basal portion arising from the median gular suture from
the posterior tentorial bridge to the postgula; the two arms of this
basal piece branch from its inner margin. The anterior extremity
of this internal gular structure forms a single median ])illar in all
Curculionoidea having a single gular suture and lacking pregidar
sutures (Figures 1-4). In all Platypodidae and Scoiytidae this an-
terior tentorial structure forms two pillars that continue antero-

June 1973 wood: platypodidae and scolytidae 89

laterally along the strongly inflected apodemal ridges formed by the
pregular sutures (Figures 5-19). In my opinion this character alone
is sufficiently unique and striking, in combination with the external
pregular sutures, to warrant family recognition of the group repre-
sented by the Platypodidae and Scolytidae.

Summary and Conclusions

The tibial and other characters used as a basis for reducing the
Platypodidae and Scolytidae to subfamilies of Curculionidae (Crow-
son, 1967; Kuschel, 1966) were adaptive characters not found in
primitive representatives of the group.

A definite pregula clearly defined by sutures associated with an
internal inflection of the cuticle occurs in Platypodidae and Scoly-
tidae but not in any other Curculionoidea having only one gular
suture. It is postulated that pregular sutures occur only in those
groups in which a rostrum is lacking and has never been developed;
therefore, since pregular sutures are absent in all groups with a
rostrum, or in which it was secondarily lost, their absence indicates
a specialization.

One genus of Platypodidae has the adult maxillary palpus 4-
segmented. Since no known representative of the Curculionidae (s.
str.) has more than a 3-segmented palpus, the ancestral stock from
which the Platypodidae were derived must predate that of the
Curculionidae.

Three of the most primitive genera of Platypodidae for which
habits are known are either phloeophagous or xylophagous and are
not associated with an ambrosial fungus. Since the ambrosial habit
has arisen independently many times in the Scolytidae on almost
every major phyletic line, it is postulated that the platypodids arose
very early from the Scolytidae and now constitute an aberrant group
within or very near that family. Tibial, tarsal, and gnathal char-
acters in primitive genera also tend to intergrade in the two groups.
Therefore, three major, equally distinctive phyletic lines of develop-
ment are recognized in Platypodidae (including Coptonotidae),
Hylesininae, and Scolytinae (including Ipinae). Platypodidae is
tentatively retained as a family separate from Scolytidae for reasons
of tradition until exhaustive studies clearly indicate the need for
a change.

References

Browne, F. G. 1972. Larvae of the principal old world genera of the Platy-
podinae (Coleoptera: Platypodidae). Trans. Roy. Ent. Soc. London. 124:167-
190.

Crowson, R. a. 1967. The natural classification of the families of Coleoptera.
Classey: Hampton, England. Reprinted from Entomologists Monthly Maga-
zine, 1950-1954, and from Lloyd, London, 1955.

Hopkins, A. D. 1909. The genus Dendroctonus. U.S. Dept. Agric, Bur. Ent.
Tech. Bull. 17(1):1-164, 8 pi.

Kuschel, G. 1966. A cossonine genus with bark-beetle habits, with remarks on
relationships and biogeography (Coleoptera Curculionidae). New Zealand
.1. Sci. 9(l):3-29.

90 GREAT BASIN NATURALIST Vol. 33, No. 2

ScHEDL, K. E. 1939. Die Einteilung und geo^aphische Verbreitung der Platy-
podidae. Proc. VII Internat. Cong. Ent., Berlin, p. 377-410.

. 1972. Monographie der Faniilie Platvpodidae Coleoptera. Junk, Den

Haag. 322 p.

Stickney, F. S. 1923. The head-capsule of Coleoptera. Illinois Biol. Monogr.
8(1):1-105.

STUDIES ON UTAH STONEFLIES (PLECOPTERA)

Richard W. Baumann^

Abstract. — Questionable Utah distribution records are reviewed and new
state records are given. The female of Capnia cygna Jewett is described and
figured and an allotype 9 designated. Detailed synonomies and nomenclatural
notes are provided for Capnia vernalis (Newport), Utacapnia poda (Nebeker and
Gaufin) and Oemopteryx fosketti (Ricker). Descriptions are given of the female,
nymph, and egg of Isogenoides zionensis Hanson and an allotype 5 designated.
"Hie male, nymph, and egg of Pictetiella expansa (Banks) are described and an
allotype 5 designated. Sweltsa gaufini, sp. nov., is described for the male, female,
and egg stages and a holotype $ designated. Descriptions are either supported
with original figures or reference is made to acceptable figures in the literature.
Besides line drawings and halftones, useful photographic figures are provided
which were prepared by using a scanning electron microscope.

An annotated list of Utah species is given following the revised nomenclature
of lilies (1966) and Zwick (1973).

The publication of a monograph on the stoneflies of Utah
(Gaufin, Nebeker, and Sessions, 1966) contributed greatly to the
knowledge of intermountain Plecoptera. Since this time, however,
additions and corrections have been discovered. This study contains
this information and includes an annotated list of Utah species.

Detailed synonymies are given for recent nomenclatural changes
and where special clarification is necessary. The listing of type
specimens and their depositories indicates that they were studied.
For further information and complete taxonomic treatment, see lilies
(1966) and Zwick (1973).

Malenka flexura (Claassen)

NemouTa flexura Claassen (1923:284). Boulder, Colorado.

This species was recorded by Gaufin et al. (1966) as occurring
in Utah. The record was based on one female in the collection of
Dr. William E. Ricker, which was collected at Huntsville, Weber Co.
The specimen was examined as part of this study and found to
belong to Malenka californica (Claassen). This does not preclude
the possibility that M. flexura might be found in the future, but no
confirmed records are presently available.

Podmosta decepta (Prison)

Nemoura decepta Prison (1942:13). Estes Park, Colorado.

This species is found in most of the Intermountain states. It was
first collected in Utah at the following locality in the Uinta Moun-
tains: small creek 7 miles N Mirror Lake, Hwv. 150, Simimit Co.,
21-VII-1967, R. W. Baumann, 1 d (NMNH).

'Department of Entomology, Smitlisonian Inslltiilion. Wnsliington. D.C. 20560.

92 GREAT BASIN NATURALIST Vol. 33, No. 2

Podmosta delicatula (Claassen)

Nemoura delicatula Claassen (1923:285). Boulder, Colorado.

Specimens of this species were collected by the author during a
survey of the stoneflies of the Wasatch Mountains. Since the species
was fairly common in small creeks at high altitudes, a closer study
was made of the collection at the University of Utah. This resulted
in the discovery of other Utah specimens previously included with
Prostoia besametsa (Ricker). The confirmed records are as follows:
Davis Co., Farmington Canyon Creek, 23-VIII-1965, R. W. Bau-
mann, 8 9? (UU). Duchesne Co., Mirror Lake, 12-VII-1947, L.
T. Nielsen, 5 5 9 (UU). Rich Co., Allen Canyon, 19-VI-1972,
G. F. Knowlton, 1 9 (NMNH). Salt Lake Co., Big Cottonwood
Creek, Brighton, 16-Vn-1952, A. R. Gaufin, 6 d^ c?, 3 9 9 ; 16-VIII-
1962, 1 d'; 6-Vm-1965, R. W. Baumann, 1 9 ; 14-VII-1966, 1 9 ;
29-VII-1967, 1 d (UU) (NMNH). Summit Co., Beaver Creek,
Beaver Creek Campground, 1 -VIM 959, A. R. Gaufin, 11 cTc?, 29

9 9 (NMNH); Hoop Lake, 30-VM958, D. B. Cahill, 1 9 (USU);
Provo River, below Trial Lake, 5-Vin-1947, A. R. Gaufin, 1 9 ; 6-
VIII-1962, 1 cT (UU); Smith Morehouse Creek, South Fork Guard
Station, 19-Vn-1961, A. R. Gaufin, 4 cf cf, 3 9 9 (UU); Weber
River, junction Smith-Morehouse Creek, 30-VI-1959, A. R. Gaufin,
12 cT cf, 29 9 9 (NMNH). Wasatch Co., Provo River, Soapstone,
21-Vn-1947, A. R. Gaufin, 2 9 9; 6-Vn-1959, 9 cfc^, 15 9 9

(UU) (NMNH).

Zapada frigida (Claassen)

Nemoura frigida Claassen (1923:285). Sitka, Alaska.

Zapada frigida is never abundant but has a scattered distribution
throughout most of the western states. A single male from the Manti-
La Sal National Forest is the first Utah record: Johnson Creek, 19
miles N Blanding, Abajo Mountains, San Juan Co., 18-VI-1946, S. B.
Muliak (NMNH).

Zapada oregoncnsis (Claassen)

Nemoura oregonensis Claassen (1923:288). Blitzen Valley, Harney Co., Oregon.

This species was recorded from Utah by Ricker (1952), Gaufin
(1955, 1964), Gaufin et al. (1966), and Baumann and Gaufin
(1971). The author examined all available specimens of Z. ore-
gonensis while studying the Rocky Mountain Nemouridae and found
them all to bo ZMpada haysi (Ricker). The species has been coiifirmed
from Idaho, Wyoming, and Colorado, so it is possibly also present
in Utah.

Capnia cygna Jewett

Capnia cygna Jewett (1954:546). Washington (?).

The type locality of C. cygna is believed to be Washington. Nebe-
ker and Gaufin (1966a) recorded it from Idaho and gave a brief

June 1973 baumann: utah plecoptera 93

description and drawing of a female dissected from a mature nymph.
Mature females have since been collected, including a single speci-
men from Mueller Park, Davis Co., Utah, 25-11-1949, R. B. Selander,
which is here designated as the allotype $ (NMNH). A detailed
description and new drawing are included because of the teneral
condition of the specimen studied earlier.

Female. — Macropterous. Length of forewings 8-10 mm; length
of body 8-10 mm. Body and appendages dark brown almost black,
broad membranous dorsal stripe on abdominal tergites 1-8. Wings
hyaline; venation typical of genus, with 1-3 crossveins between Ri
and R2 beyond cord. Eighth stemite with subgenital plate quite
simple, heavier sclerotization on both median margins, posterior
median margin produced but not extending beyond distal margin of
segment, produced portion broadly rounded or slightly angular
(Figure 2).

Capnia elongata Claassen

Capnia elongata Claassen (1924:56). Caribou, Plumas Co., California.

The confirmed distribution of this species is along the Pacific
Coast, so it probably does not occur in Utah. The specimens that
contributed to the Utah records by Knowlton and Harmston (1938),
Gaufin (1955), and Gaufin et al. (1966) were found to be Capnin
gracilaria Claassen.

Capnia vernalis (Newport) ^

Capnia vernalis Newport (1851:451). Lectotype, male; Albany River, Ontario.

Canada (BMNH).
Capnia limata Prison (1944:155). Holotype, male; South Platte River, Littleton,

Colorado (INHS).
Capnia vernalis: Zwick (1973: ?).

The name Capnia limata has been given to specimens from the
western United States, while the name Capnia vernalis is used in
Canada and the northeastern United States. After I examined both
type specimens, it was apparent that a single, widely distributed
species was involved. This synonymy is recorded in Zwick (1973).

Isocapnia hyalita Ricker

Isocapnia hyalita Ricker (1959:648). Hyalite Creek, Gallatin Co., Montana.

The discovery of this species in Utah is a large range extension,
since all previous records are from Montana. It appears to be quite
common if sought in the correct habitat at the right time. All records
to date are from mountain streams during the months of April and
May. The new Utah records are: Utah Co., American Fork Creek,
near Timpanogos Cave National Monument, 8-IV-1967, C. D. Bjork
and B. R. Oblad, 2 long- winged d"c?, 1 9; lO-IV-1967, 2 long-
winged d'cf, 2 short-winged d'cf, 2 9 9; 12-IV-1967, 1 short-
winged d", 1 9 (NMNH). Tooele Co., South Willow Creek, Upper

GREAT BASIN NATURALIST

Vol. 33, No. 2

Figs. 1, 3-5, 8. Sweltsa gaufini, n. sp.: 1, male abdomen, dorsal view; 3,
female terminalia, ventral view; 4, epiproct, a. lateral, b. dorsal, c. ventral; 5,
leaflike structure on aedeagus; 8, head and pronotum. Fig. 2. Capnia cygna
Jewett: Female tentninalia, ventral view. Fig. 6. Sweltsa albertensis (Needham
and Claassen): Leaflike structure on aedeagus. Fig. 7. Sweltsa lamba (Needham
and Claassen): Leaflike structure on aedeagus. Fig. 9. Pictetiella expansa (Banks):
Right wings of male.

June 1973 baumann: utah plecoptera 95

Narrows, 15-V-1967, R. W. Baumann and B. R. Oblad, 5 long-
winged cf cT , 6 short- winged dd,^ ? 9 (NMNH).

Utacapnia poda (Nebeker and Gaufin)

Capnia poda Nebeker and Gaufin (1965:475). Holotype, male; Gunnison River,

Colorado (UU).
Capnia (Utacapnia) poda Nebeker and Gaufin (1967:226).
Utacapnia poda: Zwick (1973:?).

This species is widely distributed in Colorado, and the collection
of a specimen in Utah near the border is not too surprising: Green
River, Jensen, Uintah Co., 24-11-1968, R. W. Baumann, 1 9

(NMNH).

Oemopteryx fosketti (Ricker)

Brachyptera fosketti Ricker (1965:475). Holotype, male; South Saskatchewan

River, Clarksboro, Saskatchewan, Canada (CNC).
Brachyptera zelona Ricker (1965:477). Holotype, male; My ton, Utah (INHS).
Oemopteryx fosketti: Zwick (1973:?).

Oemopteryx zelona was described from one male collected at the
Duchesne River in Myton, Utah, which Ricker (1965) erroneously
listed as the Green River. Since the holotype was a single male, a
special effort was made to collect additional specimens. The exami-
nation of several males from Utah led to the synonymy of O. zelona
under O. fosketti included in Zwick (1973). The following records
have become available since the original description: Duchesne Co.,
Duchesne River, Myton, 24-11-1968, R. W. Baumann, 1 ? (NMNH) ;
Duchesne River, near Randlett, 24-11-1968, R. W. Baumann, 1 cf
(NMNH). Uintah Co., Uinta River, Hwy. 40, near Roosevelt,
5-III-1963, A. R. Gaufin, 1 $ (UU); Green River, Dinosaur Na-
tional Monument, 25-III-1967, S. L. Jensen, 3 $ 2 (NMNH);
Green River, Jensen, 24-11-1968, R. W. Baumann, 9 d'cf, 48 9 9
(NMNH); Green River, Ouray, 24-11-1968, R. W. Baumann and
G. Z. Jacobi, 9 d^c^,8 9 9 (NMNH).

Isogenoides zionensis Hanson

Isogenoides zionensis Hanson (1949:109). Holotype, male; Zion National Park,
Utah (UMA).

Female. — Macropterous. Length of forewings 21-23 mm;
length of body 20-22 mm. Body dark brown dorsally, yellow ven-
trally; legs brown. Pronotum brown, with broad yellow median
stripe. Subgenital plate large, extending 1/3 length of eighth sternite,
posterior margin mostly straight, sometimes with small, rounded
median projection, lateral corners slightly rounded, forming nearly
right angles.

Nymph. — Length of mature d" 22-25 mm; mature 9 26-29
mm; antennae 7-9 mm; cerci 10-12 mm; with single submental gills
6-7 times as long as wide. Body and legs light brown; occipital ridge

GREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 10. Isogenoides zionensis (Hanson): Mature nymph.

June 1973 baumann: utah plecoptera 97

heavy on lateral margins, becoming somewhat thinner towards
median line; thin rows of long, light hairs along middorsal line of
thorax and abdomen; faint color pattern with small setae on dorsal
surface of head and thorax (Figure 10). Mouthparts similar to
/. elongatus (Hagen). Labium typical in shape for genus, para-
glossae covered with long, dense hairs. Maxillae elongate; lacinia
with two short teeth, one apical and one subapical, inner margin
with sparse row of thin spinules; galea long and narrow; tip rounded
and bearing tuft of short spines on ventral half (Figure 130), ex-
tending to base of subapical tooth; palpus five-segmented (Figure
13P). Mandibles rather similar, with two apical and one subapical
tooth on each cusp, ventral basal margin of outer cusp with row of
stout denticles, narrow patch of fine hairs extending from denticles
to base of mandible (Figure 14Q,R), basal margin of inner cusp
with dense tuft of long, thin hairs, dorsal surface with rectangular
patch of hairs at base of teeth (Figure 14S,T) .

Egg. — Average width 350 /(,, average length 550 /x. Oval and
somewhat elongate, triangular in cross-section, sides equilateral.
Collar located at one end, round in shape, with three stabilizing ribs,
one at each angle, with broadly rounded, membranous anchor plate
when fresh. Microstructure of surface composed of numerous round-
ed knobs of almost equal size. Micropiles situated at midline in rows
of 4-6 running perpendicular to long axis, present on all three sides
(Figure 12A,B,C,D).

Isogenoides zionensis was described from five males collected at
the Virgin River in Zion National Park. Gaufin et al. (1966) gave
figures of a cT abdominal tip, 5 subgenital plate, and nymphal
mouth parts but did not include descriptions. They also gave excel-
lent figures of the male genitalia, which were mislabeled as being
those of Isogenus elongatus. Their figure of the mandible fails to
show the denticles on the basal margin of the outer cusp. The labium
is shown as bare without the conspicuous covering of dense hairs.
The egg is figured in Knight et al. (1965) .

The nymph is almost indistinguishable from /. elongatus (Hagen)
and keys out as such in Ricker (1952), the only known difference
being the very light brown almost yellow color of /. zionensis as
compared to the darker brown of /. elongatus. Nymphs are very
common throughout the Virgin River drainage of southwestern
Utah, but few adults have been collected. The author, after many
attempts, was finally able to obtain 2 cf c^ and 4 ? 9 from a culvert
at the following locality: East Fork Virgin River, Hwy. 89, near
Glendale, Kane Co., 23-V-1970, R. W. Baumann (UU) (NMNH).
One of the females is here designated as the allotype 9 (NMNH) .

Pictetiella expansa (Banks)

Perla expansa Banks (1920:317). Holotype, female; Grant, Colorado (MCZ),

10,816.
Perla expansa: Needham and Claassen (1925:81, 313, 325) description of $ and

9; figures of wings. $ and 9 genitalia {$ incorrectly associated).

GREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 11. Pictetiella expansa (Banks): Mature nymph.

June 1973 baumann: utah plecoptera 99

Perla expansa: Claassen (1931:55) description of nymph (incorrect nymphal

association) .
Isogenus (Pictetia) expansus: Ricker (1952:120-122) description of $ and nymph;

figures of nymphal maxilla and $ genitalia.
Pictetiella expansella: lilies (1966:375) (incorrect specific name).
Isogenus {Pictetia) expansus: Gaufin, Nebeker, and Sessions (1966:62, 64) figures

of $ and 9 genitalia.

Male. — Slightly brachypterous. Length of forewings 9-12 mm;
length of body 13-16 mm. Body dark brown, last two abdominal
segments yellow dorsally; legs yellowish brown; antennae brown;
cerci yellow. Head as broad as prothorax, ocellar triangle equilateral,
posterior ocelli slightly closer to eyes than to each other, anterior
area light yellow, dark brown U-shaped bands connecting lateral
ocelli to anterior ocellus, bands extending forward beyond anterior
ocellus and laterally from lateral ocelli to form rocking "H," pos-
terior median area yellow, lateral posterior corners brown, palpi
brown. Pronotum slightly wider than long, brown with broad yel-
low median stripe, rugosities rather coarse and restricted to inner %,
marginal groove present only at anterior and posterior margins,
forming broad anterior and posterior bands. Wings hyaline, with
small infuscated area near cord, veins brown, venation slightly aber-
rant (Figure 9). Abdominal segments normal to ninth, which is
narrowed dorsally and greatly extended ventrally at apical margin,
ventral extended margin broadly rounded and clothed with fine
brown hairs; tenth tergite nearly bisected by deep, sclerotized groove
which shields epiproct when not extended. Hemitergites set off from
remainder of tergite by angular membranous areas, anterior half of
hemitergites covered with long, fine hairs. Epiproct long and narrow
with pointed apex, mostly membranous with thin, sclerotized areas
visible on dorsal and ventral median areas, patches of small setae on
dorsolateral margins.

Nymph. — Length of mature cT 15-16 mm; mature 9 17-21
mm; antennae 8-9 mm; cerci 7-8 mm; with single submental gills
3-4 times as long as wide. General color brown, but quite strikingly
patterned; legs mostly yellow; antennae slightly darker at apex;
apical half of cerci very dark brown almost black. Head as wide as
pronotum; maxillae barely visible from above; hind ocelli closer
to eyes than to each other; occipital ridge as few sparse hairs near
lateral margins. Thin rows of hairs running along middorsal line of
thorax and abdomen. Head pattern light on dark background: bulb-
shaped structure in ocellar triangle, M-line, front of head, lateral
tubercles and elongate spot near each eye. Pronotum transversely
oval, almost twice as wide as long, outer borders with narrow, dark
band fringed with long setae, marginal groove complete but dark
along anterior and posterior margins, with distinct light reticulations
on dark background. Wing pads on mesonotum and metanotum well
developed, outer margins running slightly convergent to body axis,
with distinct patches of dark spines along lateral anterior borders,
light reticulate markings medially. Abdominal segments fringed
with row of dark hairs on posterior margins; faint, narrow, dark

100

GREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 12. Eggs. AD, Isogenoides zionensis Hanson: A, collar, end view
(240X); B, lateral view (200X); C, micropiles (620X); D, micropile, closeup
(6,000X). E-H, Picteticlla expansa (Banks): E. collar, end view (270X); F.
dorsal view (195X); G, ventral view (180X); H, collar opening, closeup (385X).

June 1973 baumann: utah plecoptera 101

bands on anterior margins which are interrupted medially and al-
most fade out on terminal segments. Cerci with fringes of short
hairs at apex of each segment, fringe of long hairs running dorsally
throughout entire length (Figure 11). Mouthparts quite distinctive.
Labium typical of family Perlodidae, paraglossae covered with long
dense hairs. Maxillae elongate; lacinia with two long, narrow teeth,
one apical and one subapical, inner margin with few long thin
spinules; galea short but elongate, tip pointed and bearing tuft of
short spines on ventral half (Figure 13L), extending to base of sub-
apical tooth, palpus five-segmented (Figure 13M,N). Mandibles
rather similar, with one subapical and two apical teeth on each cusp,
middle teeth with serrated margins, ventral basal margins of outer
cusp smooth and bare except for large patch of fine hairs at base of
cusp (Figure 14U,V), basal margin of inner cusp with dense tuft of
long, thin hairs, dorsal surface with rounded patch of hairs at base of
teeth (Figure 14W,X).

Egg. — Average width 300 /x, average length 550 /i. Dorsal aspect
nearly oval, axillary lateral ridges extending from median ridge at
broadly rounded end to form skewed, cross-like structure, collar end
enlarged and rounded. Triangular in cross-section, almost equi-
lateral, but ventral side longest. Collar modified as flat, hood-like
structure, convex dorsally, nearly flat ventrally with large opening,
anchor plate large, flat, and covering entire ventral surface. Micro-
structure of surface composed of numerous rounded knobs of almost
equal size, knobs arranged in loose, almost circular, designs. Micro-
piles situated near midline in rows of 5-7 running perpendicular to
long axis, those on ventral surface not as obvious. Surface of un-
cleaned egg covered with small puffball-like structures which, ac-
cording to Brinck (1949), are special adhesive bodies (Figure 12E,F,
G,H).

Pictetiella expansa was named from Colorado from a single
female. Since the time of the original description, much confusion
has existed as to the identity of this species in both the adult and
nymphal stages. Most specimens previously assigned to this species
were examined as part of this study, and four genera were repre-
sented. This indicated a need for the detailed descriptions of the
male, nymph, and egg which are included here. Kicker's (1952)
description of a nymph and of teneral male genitalia seem to be
correctly associated but are rather general. The figures of the male
and female genitalia by Nebeker (Gaufin et al., 1966) were done
from specimens used in this study and are excellent. Saether (1970)
figures a nymphal maxilla labeled Isogenus (Pictetia) expansiis
which does not appear to belong in the genus Pictetiella.

The genus Pictetiella was monotypic and recorded as nearctic
until the recent description of Pictetiella asiatica from Siberia by
Zwick et al. (1972). Records of P. expansa have been confirmed
from Colorado, Montana, Utah, and Wyoming. The Utah localities
are: Salt Lake Co., Parley's Creek above Mountain Dell Reservoir,
5~IV-1963, A. V. Nebeker, nymph (UU); Lambs Canyon Creek,

102 GREAT BASIN NATURALIST Vol. 33, No. 2

19-VI-1966, R. W. Baumann, nymphs; 18-VIII-1966, exuviae (UU);
Mill Creek, Aug. and Sept. 1965, R. W. Baumann, 12 d' cT, 15 9 $ ,
nymphs; July and Aug. 1966, 7 <S <S , 8 ? 9 nymphs (UU)
(NMNH); Big Cottonwood Creek, Cardiff Fork, 12-VIII-1966, R. W.
Baumann, 2 cfcf, nymphs (NMNH). Utah Co., Deer Creek,
American Fork Canyon, ll-Vni-1966, R. W. Baumann, 1 d", 1 9 •
One of the males from Mill Creek Canyon is designated as the allo-
type cf (NMNH).

Alloperla delicata Frison

Alloperla delicata Frison (1935:334). Oak Creek, Corvallis, Oregon.

Alloperla delicata is restricted to the Pacific Northwest in the
Coast, Cascade, and Sierra Nevada mountains. The record in Gaufin
et al. (1966) from Box Elder Co., Utah, proves to be the common
Alloperla sever a.

Sweltsa gaufini, sp. nov.

Male. — Macropterous. Length of fore wings 8-9 mm; length of
body 8-10 mm. General color yellow with brown markings; legs
yellow; antennae yellow at base and light brown towards apex; cerci
yellow. Head slightly narrower than prothorax, ocellar triangle equi-
lateral, posterior ocelli the same distance from each other as from
eyes, dark reticulated markings around ocelli which continue to an-
terior margin and also appear at lateral basal margins (Figure 8).
Pronotum wider than long, completely encircled by brown border
which is fainter along anterior and posterior margins, median stripe
faint, lateral rugosities quite distinctive, posterior border very broad
(Figure 8). Wings hyaline, venation typical for genus. Abdomen
with broad dark dorsal stripe; ninth tergite with sclerotized, V-shaped
process at anterior margin, deeply excavated at posterior margin,
ninth stemite greatly expanded and forming ventral plate; tenth
tergite completely bisected, bearing lateral sclerotized bars; epiproct
mostly membranous: dorsal aspect long and narrow, carina
narrower than body with constriction near apex which grades down-
ward to pointed ventral tip; lateral aspect with narrow constriction
near broad base, which has lateral discoid appendages, anterior por-
tion becoming very broad just behind apex which bears the
pointed tip, basal area with narrow sclerotized ventral projection
which is quite broad in lateral view (Figure 4A,B,C). Aedeagus with
sclerotized, leaf-like appendage, rectangular in shape, with deep
median incision, lateral basal extensions with pointed processes
(Figure 5).

Female. — Macropterous. Length of forewings 9-10 mm; length
of body 9-11 mm. Color and general morphology similar to male.
Subgenital plate formed from posterior median portion of seventh
sternite, broad at base, tapering to broadly rounded tip, usually with
definite lateral undulations, extending over complete width of eighth
stemite, covered with numerous fine hairs, especially apparent at
apex (Figure 3).

June 1973

BAUMANN: UTAH PLECOPTERA

103

Fig. 13. IK. Sweltsa gaufini n. sp.: I, egg collar, end view (330X); J, egg,
lateral view (310X); K, collar, closeup (1,170X). L-N. Pictetiella eipansa
(Banks): L, galea, tip (1,100X); M, left maxilla, ventral (llOX); N, left maxilla,
dorsal (70X). O and P, Isogenoides zionensis Hanson: O, galea, tip (1,100X);
P, left maxilla, dorsal (60X).

104 GREAT BASIN NATURALIST Vol. 33, No. 2

Egg. — Average width 250 /x, average length 400 /x. Oval and
slightly elongate, round in cross-section. Collar terminal, highly
modified into many ridges and projections (Figure 13K), covered
with round, membranous anchor plate when fresh. Microstructure
of surface generally smooth with many small rounded pits, shallow
near midline and deeper near poles. Micropiles located below mid-
line on end opposite the collar, occurring as complete, uneven ring
(Figure 13I,J).

Sweltsa gaufini is most similar to S. albertensis and S. lamba.
Males can be most easily separated by comparing the leaflike ap-
pendage on the aedeagus. The appendage is large, rectangular, and
lightly sclerotized in S. gaufini (Figure 5); smaller, elongate, and
darkly sclerotized in 5. albertensis (Figure 6); short, broad, and
darkly sclerotized in S. lamba (Figure 7). Females are presently
difficult to separate.

This species is known only from the type locality: holotype <S ,
allotype ? , and 19 c? d" and 20 $ ? paratypes. Ricks Spring, Logan
Canyon, Cache Co., Utah, 29-Vin-1964, A. V. Nebeker (UU)
(NMNH). Holotype cT No. 72484 and allotype ? deposited at the
United States National Museum of Natural Histor}^, Washington,
D.C.

Sweltsa gaufini is named for Dr. Arden R. Gaufin of the Univer-
sity of Utah. Dr. Gaufin introduced me to the Plecoptera and pro-
vided direction in my beginning years. It is a pleasure to name a
species in the family Chloroperlidae in his honor.

Utah List

The following is a checklist of the 75 Plecoptera species known to
occur in Utah using the taxonomic arrangement of lilies (1966)
and Zwick (1973).

Nemouridae

Amphinemura banksi Baumann and Podmosta decepta ( Prison )

Gaufin P. delicatula (Claassen)

A. mogollonica Baumann and Gaufin Zapada cinctipes (Banks)

Malenka californica (Claassen) Z. columbiana (Claassen)

M. coloradensis (Banks) Z. frigida (Claassen)

Prostoia besametsa (Ricker) Z. haysi (Ricker)

Capniidae

Capnia confusa Claassen Claassen)

C. cygna Jewett /. grandis (Banks)

C. gracilaria Claassen /. hyalita Ricker

C. nana wasatchae Nebeker and Gaufin /. missourii Ricker

C. uintahi Gaufin /. veddcrensis (Ricker)

C. utahensis Gaufin and Jewett Mesocapnia frisoni (Baumann and

C. vernalis Newport Gaufin)

C. wanica Prison IJtacapnia logana (Nebeker and

Gaufin)

Eucapnopsis brevicauda (Claassen) U. lemoniana (Nebeker and Gaufin)

hnrapnia crinila (Needham and U . poda (Nebeker and Gaufin)

June 1973

BAUMANN: UTAH PLECOPTERA

105

Fig. 14. Q-T, I sogenoides zionensis Hanson: Q, right mandible, ventral
(200X); R, left mandible, ventral (200X); S, left mandible, dorsal (95X) ; T,
right mandible, dorsal (lOOX). U-X, Pictetiella expansa (Banks): U, right man-
dible, ventral (260X); V, left mandible, ventral (260X) ; W, left mandible, dorsal
(BOX); X, right mandible, dorsal (130X).

106

GREAT BASIN NATURALIST

Vol. 33, No. 2

Taeniopterygidae

Doddsia occidentalis (Banks) T. pacifica (Banks)

Oemopteryx fosketd (Ricker) T. pallida (Banks)

Taenionema nigripennis (Banks)

Paraleuctra jewetti Nebeker and

Gaufin
P. occidentalis (Banks)
P. rickeri Nebeker and Gaufin

Leuctridae

P. sara (Claassen)
Perlomyia utahensis Needham and
Claassen

Pteronarcella badia (Hagen)
Pteronarcys californica Newport

Pteronarcidae

P. princeps Banks

Perlodidae

Megarcys signata (Hagen)
Skwala parallela (Frison)
Cultus aestivalis (Needham and

Claassen)
Isogenoides colubrinus (Hagen)
/. elongatus (Hagen)
/. zionensis Hanson
Kogotus modestus (Banks)
Pictetiella expansa (Banks)

Diura knowltoni (Frison)

Isoperla ehria (Hagen)

/. fulva Claassen

/. longiseta Banks

/. mormona Banks

/. patricia Frison

/. petersoni Needham and Christensen

/. pinta Frison

/. quinquepunctata (Banks)

Paraperla frontalis Banks
Utaperla sopladora Ricker
Alloperla severa Hagen
Suwallia pallidula (Banks)
Sweltsa borealis (Banks)
S. coloradensis (Banks)

Chloroperlidae

S. fidelis (Banks)

S. gaufini Baumann

S. lamba (Needham and Claassen)

Triznaka di versa (Frison)

T. pintada (Ricker)

T. signata (Banks)

Perlidae

Doronuria californica (Banks)

D. t head or a (Needham and Claassen)

Hesperoperla pacifica (Banks)
Claassenia sabulosa (Banks)

Acknowledgments

Thanks are given to the following institutions and curators for
providing specimens for study: Dr. Arden R. Gaufin, University of
Utah (UU); Dr. W. J. Hanson and Dr. George F. Knowlton, Utah
State University (USU); Mr. D. E. Kimmins and Mr. Peter Ward,
British Museum (Natural History) (BMNH); Dr. John F. Law-
rence, Museum of Comparative Zoologv, Harvard University
(MCZ); Mr. J. E. H. Martin, Canadian National Collection of In-
sects (CNC); Dr. L. L. Pechuman, Cornell University; Dr. William
E. Ricker, Fisheries Research Board of Canada, Nanaimo, B.C.; Dr.
Donald W. Webb, Illinois Natural History Survey (INHS). Speci-
mens at the United States National Museum of Natural History are
abbreviated (NMNH) and those at the University of Massachusetts,
Amherst, Massachusetts, as (UMA).

June 1973 baumann: utah plecoptera 107

The photographic figures were prepared with the aid of Mr.
Walter R. Brown and Mary-Jacque Mann of the Smithsonian Insti-
tution, Scanning Electron Microscope Laboratory, and Mr. Jarrett L.
Cross. The excellent drawings were made by Mr. George L. Ven-
able, staff artist. Department of Entomology, Smithsonian Institution.

References

Banks, N. 1920. New Neuropteroid insects. Bull. Mus. Comp. Zool., Harvard
Coll. 64:299-362.

Baumann, R. W., and A. R. Gaufin. 1971. The stoneflies (Plecoptera) of the
Wasatch Mountains, Utah. Proc. Utah Acad. Sci., Arts, and Lett. 46(1969):
106-113.

— . 1970. The Capnia Projecta Complex of western North America (Ple-
coptera: Capniidae). Trans. Amer. Entomol. Soc. 96:435-468.

1972. The Amphinemura Venusta Complex of western North America

(Plecoptera: Nemouridae). Natur. Hist. Mus. Los Angeles Co., Contri. Sci.

266:1-16.
Brinck, p. 1949. Studies on Swedish Stoneflies (Plecoptera). Opuscula En-
tomol. Supp. 11. 250 p.
Claassen, p. W. 1923. New species of North American Plecoptera. Can.

Entomol. 55:281-292.
. 1931. Plecoptera Nymphs of America (North of Mexico). Thomas

Say Found. Entomol. Soc. Amer. 3. 199 p.
Prison, T. H. 1935. New North American species of the genus Alloperla

(Plecoptera: Chloroperlidae) . Trans. Amer. Entomol. Soc. 61:331-344.
. 1942. Descriptions, records, and systematic notes concerning western

North American stoneflies (Plecoptera). Pan-Pac. Entomol. 18:9-16.

1944. Three new species of Capnia from Colorado (Plecoptera: Cap-

niidae). Trans. Amer. Entomol. Soc. 69:151-157.
Gaufin, A. R. 1955. The stoneflies of Utah. Proc. Utah Acad. Sci., Arts, and

Lett. 32:117-120.
. 1964. Systematic list of Plecoptera of Intermountain region. Proc.

Utah Acad. Sci., Arts, and Lett. 41:221-227.

1970. Type-species designation for the subgenus Utacapnia (Plecoptera:

Capniidae). Entomol. News, Entomol. Rec. 81:197.
Gaufin, A. R., A. V. Nebeker, and J. Sessions. 1966. The Stoneflies (Ple-
coptera) of Utah. Univ. Utah Biol. Ser. 14. 93 p.
Hanson, J. F. 1949. Studies on the Plecoptera of North America: V. Notes on

Isogenoides. Bull. Brooklyn Entomol. Soc. 44:109-116.
Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82. Walter de

Gruyter and Co., Berlin. 632 p.
.Iewett, S. G., Jr. 1954. New stoneflies (Plecoptera) from western North

America. J. Fish. Res. Bd. Can. 11:543-549.
Knight, A. W., A. V. Nebeker, and A. R. Gaufin. 1965. Further descriptions

of the eggs of Plecoptera of western United States. Entomol. News 76:

233-239.
Knowlton, G. F., and F. C. Harmston. 1938. Notes on Utah Plecoptera and

Trichoptera. Entomol. News 49:284-286.
Nebeker, A. V., and A. R. Gaufin. 1965. The Capnia Columbiana Complex

of North America (Capniidae: Plecoptera). Trans. Amer. Entomol. Sor.

91:467-487.

. 1966a. New Stoneflies from Idaho (Plecoptera). Entomol. News 77:36-43.

. 1966b. New Paraleuctra from the Rocky Mountains (Plecoptera: Leuc-

tridae). Entomol. News 77:255-259.

1967. New Capnia from the Rocky Mountains (Plecoptera, Capniidae).

Trans. Amer. Entomol. Soc. 93:235-247.
Needham, J. G., AND P. W. Claassen. 1925. A Monograph of the Plecoptera

or Stoneflies of America North of Mexico. Thomas Say Found. Entomol.
Soc. Amer. 2. 397 p.

108 GREAT BASIN NATURALIST Vol. 33, No. 2

Nelson, C. H., and J. F. Hanson. 1971. Contribution to the anatomy and

phylogeny of the family Pteronarcidae (Plecoptera). Trans. Amer. Entomol.

Soc. 97:123-200.
Newport, G. 1851. On the anatomy and affinities of Pteronarcys regalis,

Newm.: with a postscript, containing descriptions of some American Perlidae.

together with notes on their habits. Trans. Linn. Soc. London 20:425-452.
Richer, W. E. 1952. Systematic Studies in Plecoptera. Indiana Univ. Publ.

Sci. Ser. 18. 200 p.
. 1959. The species of Isocapnia Banks (Insecta, Plecoptera, Nemouri-

dae). Can. J. Zool. 37:639-653.

1965. New records and descriptions of Plecoptera (Class Insecta).

J. Fish. Res. Bd. Can. 22:475-501.
Saether, 0. A. 1970. Chironomids and other invertebrates from North Boulder

Creek, Colorado. Univ. Colorado Stud., Ser. Biol. 31:57-114.
ZwiCK, P. 1973. Ordo Plecoptera: Phylogenetisches System und Katalog.

Walter de Gruyter and Co., Berlin (In Press).
ZwiCK, P., I. M. Levanidova, and L. a. Zhiltzova. 1971. On the fauna of

Plecoptera from the Soviet Far East. Rev. Entomol. USSR. 50:849-869.

THE MALE BRACHYCISTIDINE WASPS OF

THE NEVADA TEST SITE

( HYMENOPTER A : TIPHIIDAE) '

Marius S. Wasbauer-

Brachycistidine wasps are relatively common inhabitants of arid
regions in the western United States and may be very abundant
locally. There is a strong sexual dimorphism throughout the sub-
family. The males are fairly strong fliers and are attracted to var-
ious light sources. Because of this, they are well represented in col-
lections of western material. The females, on the other hand, are
wingless, strongly fossorial, and probably spend a large proportion
of their life cycle underground. They are poorly known. As might
be expected, the classification is based principally on the males
(Wasbauer, 1966, 1968).

The present study is based on 111 specimens of males from the
Nevada Test Site which were made available to me through the
kindness of Dr. Dorald Allred, Brigham Young University, Provo,
Utah.

The collections made thus far have not been sufficiently inten-
sive to provide information on ecological relationships or seasonal
occurrence. The data given here are thus in the form of a list of the
brachycistidine taxa found within the boundaries of the Test Site
study areas. A statement of general distribution and specific locality
records are given for each species. A code designation provided for
each specimen collected during the course of the sampling at the
Test Site is given in parentheses as part of the specific locality rec-
ord, and where applicable, a plant association is indicated. Addition-
al information on code designations and biotic communities of the
Nevada Test Site may be found in Allred, Beck, and Jorgensen
(1963, 1963a).

Genus Acanthetropis Wasbauer

Acanthetropis aequalis (Fox). — This species is widespread in the
Great Basin. It occurs at least as far north as Idaho and extends
eastward into Kansas and Nebraska. Five specimens were seen
from the Nevada Test Site. They were all taken from a Grayia-
Lycium community approx. 35 miles NW Mercury, 22 July 1965,
incandescent light and black light [18M(T) and 18M(TB)].

Acanthetropis noctivaga (Bradley). — This species is common in
the Colorado and Mojave deserts of California, extending through
southern Nevada to Arizona. It occurs also in the Sonoran Desert of
mainland Mexico and the Vizcaino Desert of northern Baja Cali-
fornia. It has been taken at four areas in the Nevada Test Site: one

'Brigham Young University — Atomic Energy Commission Report COO-786-61.

^Laboratory Services, California Department of Agriculture, 1220 N Street, Room 340, Sacra-
mento, California 95814.

109

110 GREAT BASIN NATURALIST Vol. 33, No. 2

specimen, 13 miles NW Mercury, 21 July 1965, Larrea-Franseria
community, black light [401 (M)TB]; two specimens, approx. 8
miles N Mercury, 19 July 1965, Larrea-Franseria community,
black light [5M(TB)]; one specimen, vicinity Mid Valley, approx.
20 miles NW Mercury, 8 August 1965, Coleogyne community,
black light [17M(TB)]; four specimens, approx. 26 miles NNW
Mercury, 27 July 1965, Coleogyne community, incandescent light
[1M(T)].

Genus Brachycistis Fox

Brachycistis glabrella (Cresson). — A common, widespread, and
quite variable species, B. glabrella has been taken in every state
west of the lOOth meridian and south of the 45th parallel. In Mexi-
co, it occurs in the Chihuahuan Desert and extends south along the
great Central Plateau. It has been taken at two areas in the Nevada
Test Site: one specimen, vicinity Rainier Mesa, approx. 36 miles
NNW Mercury, 9 August 1965, Pinon- Juniper association, black
light [12M(TB)]; two specimens, approx. 20 miles NW Mercury,
8 August 1965, Coleogyne community, black light [17M(TB)].

Brachycistis inaequalis Fox. — The distribution of this species is
limited to southern and western Arizona, southern Nevada, and the
San Joaquin Valley and Colorado and Mojave deserts of California.
It has been taken at three areas in the Nevada Test Site: one speci-
men, vicinity north portion Yucca Flat, approx. 34 miles N Mer-
cury, 28 June 1963, Allrcd live-catch rodent trap [lOSHll(L)];
four specimens, approx. 8 miles N Mercury, 19 July 1965, Larrea-
Franseria community, black light [5M(TB)]; eight specimens,
approx. 26 miles NW Mercury, 16 July 1965, Coleogyne com-
munity, incandescent light [1M(T)].

Brachycistis ioachinensis Bradley. — This is a somewhat variable
species, widespread in the western United States. It has been taken
from the Cascades and northern Rocky Mountains, through the
Great Basin, the southern deserts as far east as the panhandle of
Texas, and well into the Sonoran and Chihuahuan deserts of Mexi-
co. It has been taken at two areas in the Nevada Test Site: one
specimen, approx. 26 miles NNW Mercury, 27 July 1965, Coleo-
gyne community, incandescent light [1M(T)]; four specimens,
approx. 8 miles N Mercury, 19 July 1965, Larrea-Franseria com-
munity, black light [5M(TB)].

Brachycistis linsleyi Wasbauer. — This is a species of the Sono-
ran and Chihuahuan deserts of Mexico. It occurs in the Mojave and
Colorado deserts of California, through the southern portions of Ne-
vada, Arizona, and New Mexico, and in western Texas. It has been
taken at four areas in the Nevada Test Site: two specimens, vicinity
Cane Springs, 18 miles NW Mercury, 21 July 1965, Larrea-Fran-
seria community, black light [401 M (TB)]; one specimen, Mercury,
29 August 1965, Larrea-Franseria community, incandescent light
[M(T)]; one specimen, approx. 8 miles N Mercury, 19 July 1965,
Larrea-Franseria community, black light [5M (TB)]; one specimen.

June 1973 wasbauer: tiphiidae 111

approx. 26 miles NNW Mercury, 16 July 1965, Coleogyne com-
munity, incandescent light [1M(T)].

Brachycistis triangularis Fox. — This species is strictly a desert
inhabitant, and its distributional pattern coincides almost exactly
with that of B. linsleyi given above. It has been taken at four areas
in the Nevada Test Site: four specimens, approx. 8 miles N Mer-
cury, 19 July 1965, Larrea-Franseria community, black light
[5M (TB)]; three specimens, vicinity Cane Springs, 18 miles NW
Mercury, 12 June 1965, Larrea-Franseria community, black light
(CTB) ; one specimen, same locality, 21 July 1965, incandescent
light [401M(T)]; one specimen, approx. 35 miles NW Mercury,
8 August 1965, black light [19M(TB)]; four specimens, approx. 20
miles NW Mercury, 8 August 1965, Coleogyne community, black
light [17M(TB)].

Genus Brachycistina Malloch

Brachycistina acuta Malloch. — This species occurs in the deserts
of southern California, southern Nevada, and western Arizona. It
has been taken at four areas in the Nevada Test Site: one specimen,
approx. 8 miles N Mercury, 19 July 1965, Larrea-Franseria com-
munity, black light [5M(TB)]; one specimen, approx. 26 miles
NNW Mercury 16 July 1965, Coleogyne community, incandescent
light [1M(T)]; one specimen, vicinity Cane Springs, 18 miles NW
Mercury, 21 July 1965, Larrea-Franseria community, black light
[401M(TB)].

Genus Colocistis Krombein

Colocistis brevis (Fox). — This species is a Sonoran Desert in-
habitant and occurs in southern California, southern Nevada, and
western Arizona. It has been taken at one area in the Nevada Test
Site: two specimens. Mercury, 15 August 1965, Larrea-Franseria
community, incandescent light [M(T)].

Colocistis castanea (Cresson). — This is a large, conspicuous
wasp, very abundant in the deserts of southern California, southern
Nevada, Arizona, New Mexico, and Texas. This was the most fre-
quently collected brachycistidine wasp at the Nevada Test Site in
terms of numbers, but it was taken only in three areas: four speci-
mens, vicinity Cane Springs, 18 miles NW Mercury, 21 July and 9
August 1965, Larrea-Franseria community, black light and incan-
descent light [401M(TB) and 401 M(T)]; three specimens, approx.
8 miles N Mercury, 19 July 1965, Larrea-Franseria community,
black light [5M(TB)]; one specimen, same locality, 20 July 1965,
incandescent light [5M(T)]; eight specimens. Mercury, 4-5 August
1965, Larrea-Franseria community, black light [M(TB)]; 19 speci-
mens, same locality, 4-25 August 1965, incandescent light [M(T)].

Colocistis crassa (Bradley). — This is another large species which
occurs in the Great Basin and throughout the southern deserts of
California and Nevada, east to Texas. It has been taken at four areas

112 GREAT BASIN NATURALIST Vol. 33, No. 2

in the Nevada Test Site: one specimen, Mercury, 4 August 1965, Lar-
rea-Franseria community, incandescent light [5M(T)]; one speci-
men, vicinity Cane Springs, 12 June 1965, Larrea-Franseria com-
munity, black light (CTB) ; one specimen, vicinity Mid Valley,
approx. 20 miles NW Mercury, 8 August 1965, Coleogyne com-
munity, black light [5M(TB)]; seven specimens, approx. 8 miles N
Mercury, 19 July 1965, Larrea-Franseria community, black light
[5M(TB)].

Colocistis eremi (Bradley). — As far as is known, this species
occurs only in the southern deserts of California and Nevada. There
is one record for the Nevada Test Site: one specimen, 13 miles NW
Mercury, 21 July 1965, Larrea-Franseria community, black Hght

[401M(TB)].

Genus Quemaya Pate

Quemaya paupercula (Bradley). — This is a small, relatively in-
conspicuous wasp which is restricted, as far as is known, to the
southern deserts of California and Nevada. It has been taken at
one area in the Nevada Test Site: four specimens, approx. 8 miles
N Mercury, 19 July 1965, Larrea-Franseria community, black light
[5M(TB)].

Literature Cited

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. 2(2): 1-52.

. 1963a. Nevada Test Site study areas and specimen depositories. Brig-
ham Young Univ. Sci. Bull., Biol. Ser. 2(4): 1-15.

Wasbauer, M. S. 1966. Revision of the male wasps of the genus Brachycistis
in America north of Mexico (Hymenoptera: Tiphiidae). Univ. Calif. Publ.
in Entomol. 43:1-96.

. 1968. New genera of male BracJiycistidinae with a redescription of

Brachycistellus Baker and a key to North American genera (Hymenoptera:
Tiphiidae). Pan-Pac. Entomol. 44(3) : 184-197.

NOTES ON AQUATIC AND SEMIAQUATIC HEMIPTERA

FROM THE SOUTHWESTERN UNITED STATES

(INSECTA: HEMIPTERA)

John T. Polhemus^

Abstract. — Notes are given concerning distribution and habitat for Och-
terus barberi Schell, Hydrometra aemula Drake, Metrobates artus Anderson,
Metrobates denticornis Champion, Rhagovelia becki Drake and Harris, Velia
beameri Hungerford, Velia summersi Drake, Abedus (Abedus) breviceps Stal,
Abedus {Deinostoma) herberti Hidalgo, Ambrysus buenoi Usinger, and Ambrysus
thermarum La Rivers. The following synonymies are noted: Hydrometra aemula
Drake 1956 = Hydrometra ciliata Mychajliw 1961, new synonymy; Gerris
paludum Fabricius 1794 = Gerris uhleri Drake and Hottes 1925, new synonymy.
The macropterous form of Velia beameri is described.

These notes concern the synonymy and distribution of aquatic
and semiaquatic Hemiptera collected mostly by myself during
numerous trips to the southwestern states, including southwestern
Colorado and parts of Texas. Additional notes are given concerning
habitat preferences, and the description of the winged form of one
of our Velias is included. The material covered here did not seem
appropriate for papers already published or in preparation but is
limited to those items that seem to offer a significant contribution.

Unless otherwise noted, all specimens are in the Polhemus col-
lection; specimens in other collections will be indicated by the insti-
tutional abbreviations given in the acknowledgments.

I am indebted to the following people for help in various ways
during the work that led to the preparation of this paper: H. C.
Chapman, Lake Charles, La.; the late C. J. Drake; R. C. Froeschner,
Smithsonian Institution; J. Herring, USDA, United States National
Museum (USNM); H. R. Leech, California Academy of Sciences
(CAS) ; A. S. Menke, USDA, United States National Museum; J. C.
Schaffner, Texas A & M University (TAM); J. Schuh, Klamath
Falls, Oreg.; P. Wygodzinsky, American Museum of National His-
tory (AMNH).

Ochterus barberi Schell

Ochterus barberi Schell, 1943, J. Kansas Ent. Soc. 16:41 (Type 5, Colorado
Canyon, Arizona; USNM).

Heretofore known from Arizona, California, and Mexico, this
species was taken in New Mexico. This ochterid is common in the
desert areas of Arizona.

Material examined.— Arizona: 2 d'cT, 1 9,1 nymph, Ara-
vaipa, 8-X-1964, CL315, J. T. Polhemus; 2 c^ cf , 5 miles NE Castle
Hot Springs, 7-X-1964, J. T. Polhemus; 1 9, Indian Hot Springs,
9-X-1964, CL314, J. T. Polhemus; 1 cf, Whiteriver, 15-IV-1965,

'Associate in entnmnlogy, University of Colorado Museum, 3115 S. York, Englewood, Colorado
80110.

113

114 GREAT BASIN NATURALIST Vol. 33, No. 2

CL317, J. T. Polhemus. New Mexico: 2 cT cT, Gila River, near Gila
National Monument, 17-IV-1965, J. T. Polhemus; 1 ?, Sherman,
17-IV-1965, CL322, J. T. Polhemus.

Hydrometra aemula Drake

Hydrometra aemula Drake, 1956, Proc. Biol. Soc. Wash. 69:153 (Type 9, Her-
mosillo, Mexico; Drake Collection, Smithsonian Institution).

Hydrometra ciliata Mychajliw. 1961, J. Kansas Ent. Soc. 34:29. (Type 5,
Guerrero, Mexico; F. H. Snow Entomological Museum, Lawrence, Kansas).
New synonymy.

The examination of numerous specimens that I had identified as
H. ciliata has convinced me that ciliata is a synonym of aemula. ~
Drake described the latter from a single female from Hermosillo,
Sonora, Mexico, and Mychajliw described ciliata from six specimens
from Mexico. The turned-up abdomen of the female and hairy ven-
ter of the male separate it immediately from any other species found
in the United States; it is new to our fauna.

Material examined. — Arizona: 13cfcr,ll $9,5 miles NE
Castle Hot Springs, CL312, 7-X-1964, J. T. Polhemus; 1 d', 1 ?,
E of Castle Hot Springs, CL541, 16T-1972, J. T. Polhemus.

Gerris paludum Fabricius

Gerris paludum Fabricius, 1794. Ent. Syst. 4:188 (Type sex. locality, and dispo-
sition unknown to me).

Gerris uhleri Drake and Hottes, 1925, Proc. Biol. Soc. Wash. 38:69 (Type 5,
"Ariz"; USNM). New synonymy.

For a decade I have been disturbed by the inability of myself and
others to secure additional specimens of Gerris uhleri, which was
described, presumably, from "Arizona." Also disturbing was the
great similarity between this species and Gerris paludum Fabricius
from Europe. I examined the type of uhleri and confirmed that it is
indeed synonymous with paludum., but the script label "Ariz" was
still puzzling.

Fortunately I found specimens of Old World Velia in the collec-
tions of the United States National Museum bearing script labels
"Ariz" in the same handwriting, reportedly Uhler's, as the label on
the type of uhleri. Unquestionably the Velia originated in the Old
World, since the females possess an anal plate which is lacking in
the New World species of this genus. Three of the Velia were sent
to Dr. Livio Tamanini, Rovereto, Italy, who very kindly determined
them to be sauli Tamanini and caprai Tamanini. These specimens
correspond with material examined from southern France and west-
em Spain, he said, and there is a river Arize in the Toulouse region
in France which would be pronounced "Ariz." It seems most prob-
able that this is the originating locality for the type of uhleri. As it

-I li.ive now studied tlic types of aemula and rilinla and (onfmiied llioir synonymy.

June 1973 polhemus: aquatic hemiptera 115

now seems certain that uhleri originated in the Old World, its
synonymy with paludum is defensible.

The events described above underscore the importance of main-
taining historical collections (all specimens described above bear
"PR Uhler Collection" labels) and forewarn of difficulties that may
arise if and when the information content of biological collections
is computerized and then utilized without proper regard to its origin
or to the possibility of error.

Material examined. — Denmark: 2 cT cf , 2 9 ? , Logso, Rude
Skov, 28-VII-1941, K. O. Leth and E. W. Kaiser. England: 1 cf,
Surrey, 6-VII-1941; 1 cT, Rickmansworth, HT., Batchworth Lake,
26-VII-1953, PNL. France: 1 o% (Holotype, Gerris uhleri Drake
and Hottes), Ariz, P. R. Uhler Collection (USNM); 1 9, Umg.
Arcachon, Etang de Cazaux, ll-VI-1954, Weber; 1 cT, Pyrenees or.,
Umg, Banyula, 30-V to lO-VI-1953, Gallia mer., H. H. Weber leg.

Metrobates artus Anderson

Metrobates artus Anderson, 1932, J. Kansas Ent. Soc. 5:56 (Type 5, Cameron
County, Texas; F. H. Snow Entomological Museum, Lawrence, Kansas).

This little gerrid is apparently restricted to central and southern
Texas, being replaced by Metrobates trux Bueno to the west and
north and overlapping in range with Metrobates denticornis Cham-
pion in southern Texas.

Material examined.- — Texas: 2 cTcT, 3 ? 5, Altair, Colorado
River, CL390, 8-VIII-1967, J. T. Polhemus; 32 cf d", 23 5 9 , Ottine,
San Marcos River, CL396, 8-VIII-1967, J. T. Polhemus; 1 cf , 3 9 9 ,
Llano, Llano River, 7-IX-1963, H. R. Burke (TAM, JTP); 1 d ■,
1 9 , Brownsville, 22-IV-1945.

Metrobates denticornis (Champion)

Trepobatopsis denticornis Champion, 1898, Biol. Cent. Amer. 2:158 (Type $,
Mexico; Stockholm Museum).

Metrobates denticornis has been previously reported from the
United States (New Mexico, Texas) but is apparently rare here.
It is a distinctive species with a ventral tubercle on the anterior
femur.

Material examined. — Texas: 5 cTd", 4 9 9, Big Bend Na-
tional Park, Reed Camp, Rio Grande River, 25-XL1971, CL539,
J. T. Polhemus.

Rhagovelia becki Drake and Harris

Rhagovelia becki Drake and Harris, 1936, Proc. Biol. Soc. Wash. 49:106 (Type
5 , Sabinas Hidalgo, Nuevo Leon, Mexico; Drake Collection, Smithsonian
Institution) .

Described from Mexico, this species has been reported previously
from Texas. The record from Nevada represents a significant west-

116 GREAT BASIN NATURALIST Vol. 33, No. 2

ward extension of its range; the specimens were taken from a run-
off stream from hot springs.

Material examined. — Nevada: 12 cf cT, 20 9 9 , all apterous,
Moapa, Clark Co., CL281, 22-11-1964, J. T. Polhemus.

Velia beameri Hungerford

Velia beameri Hungerford, 1929, Ann. Ent. Soc. Amer. 22:759 (Type 5, Santa
Rita Mtns., Arizona; F. H. Snow Entomological Museum, Lawrence,
Kansas).

Velia beameri was originally described from one apterous male
and one apterous female from the Santa Rita Mountains, Arizona,
and has heretofore been known only from the type specimens.

I collected this veliid near Castle Hot Springs, Arizona, in thick
emergent vegetation at the edge of a small pool at the base of a low
dam whose impoundment is completely filled with sand. The in-
sects remained hidden in and under the vegetation until it was vio-
lently disturbed, whereupon they ran out onto the surface of the
water a short distance and then quickly returned to their hiding
place. Included in this series is one macropterous female which is
described below.

One additional specimen and a nymph were taken by violently
sweeping in the dark hollow of a partially submerged tree stump
along a tiny, spring-fed streamlet in Aravaipa Canyon, Arizona.

Description: Macropterous female. Body shape, coloration, and
hairiness as in apterous forms. Humeri more pronounced but not
produced (width, apterous 9 /alate 9; 40/53). Hemelytra brown,
veins prominent; basal white spots of similar length (0.75 mm) to
the wing pads in micropterous specimens; V-shaped white mark on
apical fourth opening caudad.

Material examined. — Arizona: 18 apterous cfcf, 10 apterous
9 9,1 9 macropterous (winged Plesiotype), 4 nymphs, 5 miles
NE Castle Hot Springs, CL312, 7-X-1964, J. T. Polhemus; 1 apterous
9 , 1 nymph, Aravaipa Canyon, CL315, 8-X-1964, J. T. Polhemus.

Velia summersi Drake

Velia summersi Drake, 1951, Rev. Ent. (Brazil) 22:371 (Type 9. Oak Creek
Canyon, Arizona; Drake Collection, Smithsonian Institution).

The holotype, a winged female, was described from Oak Creek
Canyon, Arizona, and the allotype later described from a small series
of specimens taken near Whiteriver, Arizona (Drake, 1957). I took
a long series of specimens along the San Francisco River in New
Mexico from the pebbles of gravel bars. By splashing water on the
shore, the Velia were flushed from their hiding places and easily
collected as they clambered over the stones or ran rapidly over the
water. A diligent search of haunts normally inhabited by Velia,
such as steeply sloping banks, emergent vegetation, and crevices in

June 1973 polhemus: aquatic hemiptera 117

logs failed to yield a single specimen. Several gravel bars along the
river yielded specimens, and a few were taken by splashing water
into pockets of a sheer rock cliff rising out of the river. Additional
specimens were taken from gravel bars in Oak Creek Canyon.

At Seven Springs Canyon north of Phoenix, I searched the gravel
bars for this veliid without success in January of 1972 but found
many specimiens in emergent vegetation in crevices of a sheer rock
wall rising from a deep pool. Violent splashing was necessary to
dislodge the specimens, and they would make every effort to return
to the vegetation. The afternoon temperature was around 50 F in
the canyon, with frosty mornings and ice noted in the road ditches;
so it is possible that these veliids retreat to this habitat and become
more or less dormant in cold periods.

It is worthy of note that all records for this species are from the
Salt River- Verde River system. Several closely related species occur
in Mexico, with Velia alvaradana Drake and Hottes occurring as far
north as Sonora, but I have not seen any material collected between
the Arizona-Mexico border and Alamos, Sonora.

The left paramere of a male is shown in Figure 1.

Figure 1 . Velia sumniersi Drake, left paramere of male from Whiteriver,
Arizona.

Material examined. — Arizona: About 100 specimens, ap-
terous, N Carefree, 18-1-1972, CL542, J. T. Polhemus; 1 d', 1 9
apterous, 1 9 macropterous, Sedona, Oak Creek Canyon, 9-VI-1969,
CL1211, J. T. Polhemus; 1 9 apterous, 8-15 miles NE Whiteriver,
8-11 -VII- 1940, Gertsch and Hood (Part of allotype series, exchange
from AMNH). New Mexico: Several hundred apterous specimens,
Glenwood, 16-IV-1965, CL319, J. T. Polhemus.

Abedus {Abedus) breviceps Stal

Abedus breviceps Stal, 1862, Ent. Zeitung 23:462 (Lectotype 9, Mexico; Riks-
museum, Stockholm).

This species is primarily found in Mexico but has been recorded
from Arizona, New Mexico, and Texas.

Material examined. — Arizona: 3 cfd', 3 9 9,1 nymph.
Camp Verde, 7-X-1964, CL310, J. T. Polhemus. Texas: 1 9, 1

118 GREAT BASIN NATURALIST Vol. 33, No. 2

nymph, Big Bend National Park, Reed Camp, Rio Grande River,
1-25-X-1971, CL539, J. T. Polhemus.

Abedus {Deinostoma) herberti Hidalgo

Ahedus herberti Hidalgo, 1935, Univ. Kansas Sci. Bull. 22:507 (Type 5, Tucson,
Arizona; F. H. Snow Entomological Museum, Lawrence, Kansas).

Abedus herberti is fairly widespread in Arizona but not com-
monly fomid elsewhere. The specimens I have collected vary sub-
stantially in size; the smallest male, from Patagonia, measured 28.5
mm in length, while the largest male, from Sedona, measured
38 mm.

Material examined.- — Arizona: 1 2,2 nymphs, Patagonia,
25-III-1967, CL1217, J. T. Polhemus; 1 d', 4 ? ? , 2 nymphs, Pata-
gonia, 17-III-1967, CL1218, J. T. Polhemus; 1 cf , 2 9 5,2 nvmphs.
Seven Springs, N Carefree, 18-1-1972, CL542, J. T. Polhemus; 1 d",
1 9, Aravaipa, 8-X-1964, CL315, P. T. Polhemus; 1 c^, 4 9 9,2
nymphs, Whiteriver, 15-IV-1965, CL317, J. T. Polhemus; 1 d",
Sedona, 9-VI-1966, CL1211, J. T. Polhemus; 1 9, East Fork Verde
River, Payson to Pine, Gila Co., 6-IV-1966, J. Schuh; 1 cT, 4 miles
S Patagonia, Santa Cruz Co., Santa Cruz River, 4-IV-1966, J. Schuh;
1 9, Santa Cruz Co., 3 miles W Fort Huachuca, 29-V-1969, D.
Schuh. New Mexico: 3 cfd', 3 9 9, Gila River, Gila National
Monument, 17-IV-1965, CL323, J. T. Polhemus.

Ambrysus buenoi Usinger

Ambrysus buenoi Usinger, 1946, Univ. Kansas Sci. Bull. 31:199 (Type 5, Bio
Grande, Brewster Co., Texas; USNM).

This species is one of the "free swimming" Ambrysus that occur
in the slow or still portions of streams. When disturbed, it swims
rapidly away in a manner much like most corixids.

Material examined. — Texas: 31 cfcf,39 9 9,3 nymphs. Big
Bend National Park, Fresno Creek, CL538, 25-XI1971, J. T. Pol-
hemus; 3 cf d', 3 9 9, Alamito Creek, 5 miles SE Presidio, 13-VI
to ll-VII-1968, J. E. Hafernik (JTP, TAM); 1 d', 1 9, Brewster
Co., 21 miles S Alpine, Calamity Creek, 19-VII-1964, D. R. Smith
and C. W. Baker.

Ambrysus thermarum La Rivers

Ambrysus thermarum La Rivers, 1953, Proc. U.S. National Museum 103:1
(Type 5, Hot Springs, Sierra Co., New Mexico; USNM 60987).

The name thermarum for this insect, alluding to its occurrence
in a hot spring area, is indeed unfortunate, for it occurs in colder
water than any other Ambrysus known to me. I have collected it
only in cold mountain streams in the region of the great caldera
northwest of Santa Fe, New Mexico. Presumably, similar streams
exist in the mountains west of Hot Springs (now Truth or Conse-

June 1973 polhemus: aquatic hemiptera 119

quences), the type locality, and the type series was almost certainly
collected from such a habitat. I have not found it in the warmer
or slower reaches of any of the streams. A. thcrmarum inhabits the
stony, swift-flowing portions of streams, where it may be taken by
turning over the rocks and holding the net downstream to catch the
dislodged bugs. At Vallecito the water was so cold that my hands
ached after a few minutes of collecting in this way.

Material examined.^ — New Mexico: 2 cf cT, 3 9 ?, Sandoval
Co., Jemez Cr., La Cueva, alt. 7634 feet, ll-VIII-1965, H. B. Leech
(JTP, CAS); 3 cf cf, 3 $9,3 nymphs, 12 miles W Espanola, 6-VL
1970, CL483, J. T. Polhemus; 3 cT d", 2 9 9 , Canones Creek, 6 miles
N Canones, 31-V-1971, CL530, J. T. Polhemus; 9 dd,^ 9 9,5
nymphs, Rio Tusas, Tusas, 9-X-1971, CL535, J. T. Polhemus; 139
specimens, Vallecito, lO-X-1971, CL536, J. T. Polhemus.

Literature Cited

Drake, C. J. 1957. Notes on the genus Velia (Hemiptera: Veliidae). J. Kansas
Ent. Soc. 30 (4): 138-140.

A NEW SUBFRUTICOSE ERIOGONUM (POLYGONACEAE)
FROM WESTERN COLORADO

James L. ReveaU

Abstract. — A new subfruticose Eriogonum (Polygonaceae) is described as E.
pelinophilum, from specimens obtained near Hotchkiss, Delta Co., Colorado. A
member of the subgenus Eucycla, the new species is most closely related to
E. contortum Small ex Rydb., but differs in having cream-colored flowers instead
of bright 3'ellow flowers. The two species also differ in that E. pelinophilum has
narrowly turbinate involucres which are longer than those of E. contortum and
larger flowers and achenes. The new species is apparently restricted to gumbo
clay hills west of Hotchkiss.

In my series of Notes on Eriogonum^ part VI dealt with the
Eriogonum microthecum Nutt. portion of the Section Corymbosa
Benth. in DC. (Reveal, 1971). In that treatment, I discussed an
undescribed species known only from a single collection which oc-
curred on clay hills near Hotchkiss, Delta Co., Colorado. In spite
of two attempts to find additional material, the plant could not be
located in the field, and the decision was made not to provide formal
recognition of this entity until it could be rediscovered and an exact
location established.

In July 1972, fieldwork in the Hotchkiss area led to the relo-
cation of this long-sought buckwheat which may now be called:

Eriogonum pelinophilum Reveal, spec. nov. A Eriogono contorta
Small ex Rydb. floribus albidis et longioribus (2.5) 3-3.5 mm longis,
involucris anguste turbinatis (2.5) 3-3.5 mm longis differt. Plantae
subfruticosae 5-10 cm altae et 8-20 (30) cm latae; folia oblanceolata,
laminis 5-12 (15) mm longis et 1-2 (3) mm latis, subtus albo-
tomentosis, supra subglabris vel glabris raro floccosis, marginibus
revolutis, petiolis brevissimis, 1 mm longis, floccosis vel glabris;
caules graciles, (3) 5-10 mm longi, floccosi vel glabri; inflorcscentiac
cymosae, 1-2 cm longae et 1-3 cm latae, ramis floccosis vel glabris,
radiis 2-7 (10) mm longis; bracteae ternatae, 0.5-1 mm longae et
latae; pedunculi erecti, 1-5 mm longi, floccosi vel glabri; involucra
solitaria, anguste turbinata, (2.5) 3-3.5 mm longa et 1-1.5 mm lata,
floccosa vel glabra, 5-lobatis 0.3-0.4 mm longis, bracteolis oblan-
ccolatis, 1.8-2.5 mm longis, pedicellis 2.5-4.5 mm longis et glabris;
f lores albidi cum costa et funda rufi, (2.5) 3-3.5 mm longi, glabri,
tepalis similaribus, oblongis, 0.8-1.2 mm latis; stamina exserta, 2.5-4
mm longa, filamentis sparse pilosis basi, antheris albidis, 0.6-0.7 mm
longis, oblongis; achaenia infuscata, 3-3.5 mm longa.

Low-rounded, heavily branched, pulvinate subshrub 5-10 cm
high, 8-20 (30) cm across, the lower woody stems light brown, the
bark exfoliating in long loose strips or in wide plates, leafless, arising
from a stout, woody taproot, the upper branches herbaceous, slender,
floccose to glabrous; leaves solitary, scattered along the entire

'Dopartniont of Rot.uiv, rriivrrsily of Mnrvlniul. CollcKO Pnik JOT-t-J. .iiiil Nntinnnl Museum of
Natural History, Smilhsonian Institution, WashinRton, D.C. 20560.

120

June 1973 reveal: a new eiuogonum 121

length of the herbaceous stems except for the last (3) 5-10 mm,
somewhat closely arranged and congested to widely spaced, the
leaf-blade oblanceolate, 5-12 (15) mm long, 1-2 (3) mm wide,
densely white-tomentose below, the midvein totally obscured by
the tomentum, subglabrous to glabrous and green above, the mar-
gin entire, revolute and complete enclosing the lower surface,
the apex and base acute, the blades persistent throughout the
growing season, the petiole short, 1 mm long, light yellowish-brown
to tan and thinly pubescent when young, becoming glabrous and
brownish at maturity, the petiole-base elongate-triangular, 1-2 mm
long, 0.7-1.2 mm wide, slightly pubescent to glabrous and light
brown without, densely tomentose within; flowering stems erect,
slender, 5-10 mm long, floccose to glabrous; inflorescences cymose,
± compact and congested, 1-2 cm long and 1-3 cm wide, trichot-
omous, the rays 2-7 (10) mm long, once or twice divided, floccose
to glabrous; bracts scalelike, ternate, 0.5-1 mm long and wide tri-
angular, floccose to glabrous without, thinly tomentose within, con-
nate at the base; peduncles, when present, 1-5 mm long, erect,
floccose to glabrous, those of the central involucre longest, 3-5 mm
long, those of the lateral involucres shorter, up to 2 mm long;
involucres solitary, narrowly turbinate, (2.5) 3-3.5 mm long, 1-1.5
mm wide, floccose to glabrous without, glabrous within, the 5 acute
teeth 0.3-0.4 mm long, the bractlet oblanceolate, 1.8-2.5 mm long,
minutely fringed with capitate, gland-tipped cells, the pedicel 2.5-
4.5 mm long, glabrous; flowers white to cream with reddish-brown
midribs and large. cons])icuous brownish-red to greenish-red bases,
(2.5) 3-3.5 mm long, glabrous within and without except for micro-
scopic glands along the midribs within, the tepals essentially similar,
oblong, the outer whorl of tepals 1-1.2 mm wide, the inner whorl
0.8-1 mm wide, the apex of both whorls rounded, united at least
V2 the length of the flower; stamens slightly exserted, 2.5-4 mm long,
the filament sparsely pubescent basally, the anther white to cream,
0.6-0.7 mm long, oblong; achenes light brown, 3-3.5 mm long, the
globose base tapering abruptly to a long, three-angled beak.

Type. — Colorado: Delta Co.: Along Colorado Highway 92, 8.6
miles west of the western edge of Hotchkiss, 0.9 miles east of the
Red Mesa turnoff and 3.1 miles west of Lazear, 11.5 miles east of
U.S. Highway 50 at Delta, on rolling clay hills associated with scat-
tered species of Atriplex and Eriogonum lonchophyllum, at 6400
feet elevation, 16 July 1972, Reveal and Reveal 2780. Holotype,
US! Isotypes, 30 duplicates to be distributed from US.

Additional specimens examined: Colorado: Delta Co.: In desert
just out of Hotchkiss, 23 July 1958, Gentry 2285 (COLO).

The Clay-loving Buckwheat, Eriogonum pelinophilum, was first
thought to be a member of the Section Corymbosa and to be related
to E. bicolor M. E. Jones (Reveal, 1971). Since seeing the plant in
the field, it appears to be more closely related to E. contortum Srnall
ex Rydb., a species which is currently placed in an unnamed section
typified by E. brevicaule Nutt. (Reveal, 1969). The discovery of

122 GREAT BASIN NATURALIST Vol. 33. No. 2

this new species now raises serious questions as to the placement of
these two species within the genus. Stokes (1936) associated E.
contortum with those elements around E. corymbosum, but the re-
duced nature of E. contortum. its yellow flowers, and smooth green-
ish stems seem to exclude it from this group of species. However,
the flower color of E. pelinophilum removes the significance of the
flower color, and one is faced with the problem of whether or not
one should place both E. contortum and E. pelinophilum with E.
brevicaule or with E. clavellatum Small and E. bicolor. Compound-
ing the problem is the highly reduced, diminutive E. acaulc Nutt. of
Wyoming. Further work is needed before the exact placement of
these species can be made, but it is possible that E. pelinophilum,
E. contortum, and E. acaule should be placed in a distinct section
occupying an intermediate position between those species of the
genus typified by E. microthecum-E. corymbosum and those re-
lated to E. brevicaule.

The new species may be distinguished from Eriogonum contor-
tum by its white or cream-colored flowers and its longer involucres,
flowers, and achenes. Eriogonum pelinophilum occurs to the south
of all known locations of E. contortum, which occupies the Grand
Valley of Utah and Colorado. So far as known, E. pelinophilum is
relatively rare in the field, being restricted to gumbo clay hills and
slopes in the desert west of Hotchkiss; no specimens were found to
the east and south of this townsite.

Acknowledgment

Field work on the genus Eriogonum is supported by National
Science Foundation grant GB-22645.

Literature Cited

Reveal, J. L. 1969. A revision of the genus Eriogonum (Polygonareae).
Unpublished doctoral dissertation. Provo.

. 1971. Notes on Eriogonum — VI. A revision of the Eriogonum micro-

thecum comple.x (Polygonaceae). Brigham Young Univ. Sci. Bull., Biol.
Ser. 13(l):l-45.

Stokes, S. G. 1936. The genus Eriogonum, a inolimiuary study based on geo-
graphic distribution. San Franrisro.

RECORDS OF COREIDAE (HEMIPTERA) FROM THE
NEVADA TEST SITE

Dorald M. AUredi

During ecological studies at the Nevada Test Site (Allred, Beck,
and Jorgensen, 1963; Beck and Allred, 1968; Knight, 1968), ap-
proximately 240 coreids were collected. Although most of these
await identification in the U.S. National Museum, Dr. R. C. Froesch-
ner kindly identified some original submittals which are reported
herein.

Arhyssus lateralis (Say). — Two specimens, 31-Vni-1959,
Frenchman Flat, plant host unknown.

Arhyssus sp. — Twelve specimens: Area 12 — 26-Vni-1965 ex
Eriogonum sp.; Area 16 — 5-20-VIII-1965, ex E. deflexum; Area 401
(now Area 26) — 16-VIII-1965, ex E. deflexum; Cane Springs — 25-
VI-1959, ex Asclepias erosa.

Harmostes angustatus Van Duzee. — Four specimens: Mercury
environs — 15-VIII-1965, host unknown; Area 2 — 25-VIII-1964, ex
Pedis papposa.

Harmostes reflexulus (Say). — Five specimens: Frenchman Flat
— 31-VIII-1959, host unknown; Area 12—7, 12-VIII-1965, ex
Chrysothamnus viscidiflorus; Area 16 — ll-VI-1965, ex Eurotia la-
nata; Area 401 (now Area 26) — 18-VI-1965 ex Sphaeralcea sp.

Liorhyssus hyalinus {¥.). — Twenty specimens: Cane Springs —
12-VII-1960, host unknown; Area 5— ll-VI-1964. 1964 ex Sphaeral-
cea ambigua; 22-VI-1964, ex Bailey a multiradiata; lO-VI-1965, ex
Larrea divaricata; Area 6 — 19-VII-1965, ex Eriogonum inf latum;
Area 16— 13-VI-1965, ex Malacothrix glabrata; 5-VIII-1965, ex
Eriogonum deflexum; Area 17 — ll-VI-1965, ex Astragalus lentigi-
nosus; 12-VI-1965, ex Malacothrix glabrata; 17-VI-1965, ex Sphaer-
alcea sp.; Area 18— 23-VIII-1965, ex Sphaeralcea sp.; Mercury en-
virons— 6-VIII-1965, ex Eriogonum deflexum; 15-VII-1965, host
unknown.

Literature Cited

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. 2(2): 1-52.
Beck, D E., and D. M. Allred. 1968. Faunistic inventory — BYU ecological

studies at the Nevada Test Site. Great Basin Nat. 28(3) : 132-141.
Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and

the western United States. Brigham Young Univ. Sci. Bull., Biol. Ser.

9(3): 1-282.

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

123

DENSITY, GROWTH, AND HOME RANGE OF THE LIZARD

UTA STANSBURIANA STEJNEGERI IN SOUTHERN

DONA ANA COUNTY, NEW MEXICO'

Richard D. Worthington- and Edward R. Arvizo'^

Abstract. — Side-blotched lizards. Uta stansburiana, were marked and re-
captured on a study plot near Anapra. Dona Ana County, New Mexico, during
the suiTuner of 1971. The density in July was estimated to be 20.3 individuals
per acre (50.2 per hectare). The home range size for males averaged 0.254
acres and for females, 0.097 acres. Males 48-53 mm s-v increased 0.034 mm/day
in s-v length and 0.067 mm/day in tail length from middle .July to early Septem-
ber. Females 41 .5-46 mm s-v increased 0.028 mm/day in s-v length and 0.058
mm/day in tail length during the same period.

During the past hundred years the vegetation of southern New
Mexico has changed from a desert grassland climax to a desert climax
(York and Dick-Peddie, 1969). Presumably, there have been major
changes in the composition of the vertebrate fauna and the relative
abundance of different species. Preliminary investigations of the
lizard fauna on a study plot located near Anapra, Dona Ana County,
New Mexico, were begun during the summer of 1970 in order to
determine species composition, relative abundance, and other eco-
logical parameters of the different populations. Data gathered on
Uta stansburiana stejnegeri during the summer of 1971 have been
analyzed and are presented and compared with the results of other
investigators.

Methods

A study plot measuring 100 yards to a side (10,000 sq yd; 2.07
acres) was constructed on the morning of 18 July 1971 and marked
with numbered and lettered stakes at 50 ft intervals forming 36
quadrats. Observations were begun on the same afternoon. The
study plot was worked by two observers on 14 visits and by one
observer on two visits through 29 July. Lizards were collected by
noosing and marked by toe clipping. Numbers were painted on the
backs of lizards with a water-base paint to facilitate recognition with
a minimum of recaptures. The point at which the lizard was first
seen was noted, and the number of feet to the nearest stake recorded
in order that the home range could be reconstructed. The sex, snout-
vent length, tail length, and length of any regenerated portion of the
tail was determined for each lizard at the time of first capture.

We attempted to estimate density by marking all individuals on
the study plot and have followed Tinkle and Woodard (1967) in
considering individuals as residents if most of the capture points were
within the study plot. The study plot and its immediate periphery
were patrolled in different patterns to avoid biasing the observations.

'Contribution No 4 from the Museum of Ariel Land Biology, University of Texas at El I'a9f>.
^Department of BioloRiral Sciences, University of Texas at El Pasn. El Paso. Texas 7^008.
'Depaitmrnt of /oolnny, Utali Slate University. Logan. ITtali R\\l\ .

124

June 1973 worthington, arvizo: lizards 125

Only individuals with five or more recapture or sighting points (up
to a maximum of 10) during the July period were used for deter-
mination of home range. The study area was worked again on three
occasions (4-6 Sept.) in order to recapture individuals marked in
July for the determination of growth rates.

Two methods were used to describe the vegetation of the study
plot. A direct count of the larger plants which could supply cover
for the lizards was made. In addition, ten groups of four one-meter-
square plots were selected randomly utilizing a random numbers
table to determine the quadrat, coin tosses to determine the quarter
of the quadrat, and stick toss to determine the one-meter plots. All
of the plants were low; therefore, the basal area (shade area) was
determined for each individual plant present. Frequency and
coverage estimates were calculated. The vegetation analysis was
conducted in September, and the area occupied by annuals and
other plants that sprouted during the summer rainy season was
subtracted in order to approximate plant coverage for July.

Study Area

The study area is in the southern part of Dona Ana County,
New Mexico, along the United States-Mexico boundary (lat. 31°47'
10"N, long. 106°35'05"W). It is situated at the edge of the La Mesa
Surface near the Rio Grande and the town of Anapra at an elevation
of 1250 m. The area consists of low dunes of wdndblown sand which
attain a maximum height of 1.3 m and which are covered sparsely
by vegetation. Larger plants on the study plot providing cover for
lizards are 68 Larrea tridcntata, 52 Yucca elata, 27 Dalea scoparia,
4 Prosopis glandulosa, and 2 Ephedra sp. Plant coverage increased
from about 19 percent at the time the study was initiated to 31
percent in September. Frequency and percentage of total coverage
for plants encountered in 40 one-meter-square plots selected at ran-
dom are presented in Table 1. Botanical preparations of a majority
of the species encountered in the study area have been deposited in
the Museum of Arid Land Biology. University of Texas at El Paso.

Table 1. Relative coverage and frequency of plant species determined from
one-square-meter plots fN^40) selected at random in the studj' area during
September 1971.

Name

Relative

Frequency

No. Plants

Coverage

(%)

Larrea tridentata

46.9

Salsola kali

10.1

Amaranthus sp.

16.7

Sporobolus giganteus

3.1

Sporobolus flexuosus

2.9

Erioneuron pulchellum

1.1

Tidestromia sp.

4.3

Dalea scoparia

9.8

Yucca elala

4.0

Others

1.2

126 GREAT BASIN NATURALIST Vol. 33. No. 2

Weather data for the El Paso International Airport, located about
21 km east of the study area, are available (Orton, 1969). The
average annual precipitation is 20.04 cm, about half of which falls
between June and September. El Paso averages 101 days a year in
which the maximum temperature is 32.2 C or above and only one
day a year in which the maximum temperature is 0 C or below.
The percentage of possible sunshine is 83. Extremes of temperature
range from -22.2 to 41.1 C.

Uta stansburiana is the most abundant lizard on the study plot.
Other reptiles collected on the study plot are Sceloporus undulatus,
Phrynosoma cornutum. Cnemidophorus tigris. Crotaphytus wisUzeni.
Masticophis flagellum, and Crotalus viridis.

Population Density

During July a few hatchlings were beginning to appear in the
population. One hatchling was marked and two others were seen;
however, they were not included in the population estimates. Esti-
mates pertain only to individuals that have overwintered at least
once. The smallest of these individuals was a 39 mm s-v immature
male; however, most of the individuals were sexually mature.

A total of 45 lizards were marked. Population estimates based on
42 individuals having all or a majority of capture points within the
study plot are 20.3 individuals per acre or 50.2 per hectare. The
sex of one individual was not determined, but of the remaining 44
individuals marked, the sex ratio was equal. Turner et al. (1970)
found density estimates for four Nevada study plots over a three-
year period to vary from 25.7 to 113.1 individuals per hectare. The
mean estimates of the four plots for the years 1966 and 1968 show
close agreement to the results reported here with estimates of 56.8
and 42.5 individuals per hectare respectively. The 1967 estimate
of 80.3 is considerably higher than the Anapra estimate and indicates
that wide fluctuations may occur at a given site.

Tinkle and Woodard (1967) compared densities of three popula-
tions of Uta stansburiana located in Texas, Colorado, and Nevada.
Estimates ranged from 12.6 individuals per acre (Nevada, June-July
1964) to 17.5 per acre (Colorado, June- August 1965), which are
lower than the Anapra estimate. Other estimates of the size of Uta
stansburiana populations in Nevada during June (Tanner and Jor-
gonsen, 1963; Tanner and Hopkin, 1972) range from 4.5 to 10.4
individuals per acre and are also lower than reported for Anapra.
The El Paso area was experiencing a severe drought just prior to the
study, with an official rainfall of less than 2 cm for the first six
months of 1971. This suggests the possibility that the estimate of
20.3 individuals per acre may be lower than normal.

Home Range

Home range size estimates were determined from recapture
radii (mean radius plus 2s) for individuals with five or more points
of capture or sighting. This method is considered the most reliable

I luie

1973 WORTHINGTON. ARVIZO: LIZARDS 127

for estimating the home range size of Uta stansburiana, although
there is some disagreement relating to the minimum number of
points to be used (Jorgensen and Tanner, 1963; Tinkle, 1967; Tin-
kle and Woodard, 1967).

The average home range area for 9 males was 0.254 acres (0.046-
0.712 acres). The average home range area of 11 females was 0.097
acres (0.016-0.164 acres). These estimates are consistently lower
than those reported by Jorgensen and Tanner (1963) and Tinkle
and Woodard (1967). Jorgensen and Tanner (1963) report home
range sizes of 0.98 acres (adult males), 0.68 acres (adult females),
and 0.42 acres (juveniles) for Uta stansburiana on the Nevada Test
Site. Tinkle and Woodard (1967) report home range sizes for Uta
in Texas, Colorado, and Nevada ranging from 0.27 acres (Colorado)
to 0.50 acres (Texas) for males and 0.12 (Nevada) to 0.23 (Colo-
rado) acres for females. Density figures for the populations studied
by Jorgensen and Tanner (1963) and Tinkle and Woodard (1967)
are lower than those reported for Anapra. Tinkle (1967) has de-
termined that the juveniles establish home ranges by penetrating and
continually expanding their initial home range until they can no
longer do so. This suggests an inverse relationship between density
and home range size and indicates that the Anapra estimates are
comparable to those obtained by other investigators.

Growth

The study area was worked between 4 September and 6 Sep-
tember 1971 to recapture lizards marked during July in order to
determine summer growth rates. During the period of July through
September, rainfall, plant growth, and presumably insect abundance
are maximal. Most, if not all. of the period is within the breeding
season of Uta. and many of the females collected in July were gravid.
Growth data for males and females are summarized in Table 2.

Tanner and Jorgensen (1963), Tanner (1965), Tinkle (1967),
and Tanner and Hopkin (1972) present growth data for Uta stans-
buriana. Tinkle (1967) presents daily growth rates of adult Uta
from near Kermit, Texas. Daily growth rates of 0.17 mm (males)
and 0.24 mm (females) for the August-September period and 0.10
nun (males) and 0.32 mm (females) during the June-July period are
considerably higher than reported for Anapra lizards (Tinkle, 1967).
It is possible that the discrepancy may represent the inclusion of
many immature individuals in the size classes Tinkle (1967) con-
structed, whereas all of the Anapra lizards are reproductive. Growth
data for Nevada and Utah Uta (Tanner and Jorgensen, 1963; Tan-
ner, 1965; Tanner and Hopkin, 1972) indicate that adults increase
in size at rates more comparable to those reported for the Anapra
population.

Tail-growth rates as high as 0.13 mm/day for 9 males (46-50
mm. s-v) and 0.05 mm/day for 3 females (42-46 mm s-v) are reported
by Tinkle (1967) for Uta collected during September through No-
vember near Kermit. Texas. These rates are comparable to those

128 CREAT BASIN NATURALIST Vol. 33, No. 2

determined for Anapra lizards; however, linkle's June through
August sample rates of 0.01-0.02 mm/day are lower (Table 2).
Rates of increase for total length are included for future comparisons
(Table 2).

Table 2. Daily growth rates of Uta stansburiana stejnegeri in southern
Dona Ana Co., New Mexico, marked between 18 July and 26 July and recap-
tured during 4-6 September 1971.

Sex

S-V Size Range
July

Increase S-V
(mm/day)

Increase Tail
(mm/day)

Total Increase
(mm/day)

Males
Females

48-53
41.5-46

10
10

0.034
0.028

0.067(N = 6)
0.058(N = 5)

0.105(N = 6)
0.099(N = 5)

Literature Cited

JoRGENSEN, C. D.. AND W. W. Tanner. 1963. The application of the density
probability function to determine the home ranges of Uta stansburiana stans-
buriana and Cnemidophorus tigris tigris. Herpetologica 19(2) : 105-1 15.

Orton, R. R. 1969. Climates of the states (Texas). U.S. Dept. Commerce. Env.
Sci. Sciv. Admin.. Env. Data Service, pp. 1-46.

T\NNER, W. W. 1965. A comparative population study of small vertebrates
in the uranium areas of the upper Colorado River Rasin of Utah. Rrigham
Young Univ. Sci. Rull., Riol. Ser. 7(1): 1-31.

Tanner, W. W., and J. M. Hopkin. 1972. Ecology of Sceloporus occidentalis
longipes Raird and Uta stansburiana stansburiana Raird and Girard on
Rainier Mesa. Nevada Test Site. Nye County, Nevada. Rrigham Young Univ.
Sci. Rull., Riol. Ser. 15 (4): 1-39.

Tanner, W. W., and C. D. Jorgensen. 1963. Reptiles of the Nevada Test
Site. Rrigham Young Univ. Sci. Rull., Riol. Ser. 3(3): 1-31.

Tinkle, D. W. 1967. The life and demography of the side-blotched lizard.
Uta stansburiana. Misc. Publ. Mus. Zool. Univ. Mich., No. 132, p. 1-182.

Tinkle, D. W.. and D. W. Woodard. 1967. Relative movements of lizards in
natural populations as determined from recapture radii. Ecology 48:166-168.

Turner, F. R., G. A. Hoddenbach, P. A. Medica, and J. R. Lannom. 1970.
The demography of the lizard, Uta stansburiana Raird and Girard. in south-
em Nevada. J. Anim. Ecol. 39:505-519.

York, J. C, and W. A. Dick-Peddie. 1969. Vegetation changes in southern
New Mexico during tlic past lunidrcd >oat's. pjj. 1 35- 166. In W. G. MrGiiuiies
anfl B..I. Goldman fed.] .'\rid hinds iti perspective. Am. Assoc. Aihaiico.
Sci.. Washington. D.C.

SMALL BONES OF THE HYPSILOPHODONTID DINOSAUR

DRYOSAURUS ALT US FROM THE UPPER

JURASSIC OF COLORADO

Peter M. Gallon' and James A. Jensen^

This note reports the discovery of numerous fragments of a
small ornithopod dinosaur from the Morrison Formation (Upper
Jurassic) of Colorado. This material is referred to the hypsilopho-
dontid Dryosaurus alius (Marsh) and represents juvenile individ-
uals of this species.

Very few specimens of hypsilophodontid dinosaurs have been re-
ported from Colorado, and those were found almost a hundred years
ago. Marsh (1887a) erected Nanosaurus agilis for a specimen (Yale
Peabody Museum No. 1913) from Garden City, a few miles north of
Canon City, with material from the Hallopus Beds of probable Up-
per Jurassic age (Schuchert, 1939). However, the specimen is very
fragmentary, with many of the bones represented by natural molds
(Huene and Lull, 1908), and it is not demonstrably hypsilophodontid.
Marsh (1877b) also described a well-preserved femur (YPM 1915)
from the Morrison FoiTnation of Garden City as Nanosaurus rex,
but Huene and Lull (1908) correctly referred it to the genus
Laosaurus Marsh.

The new material from Colorado consists of many vertebral
fragments and the ends of long bones and metatarsals, the frag-
mentary condition of which is probably due to the fact that it was
collected from the surface rather than in situ.

Occurrence. — Near the base of the Upper or Brushy Basin
Member of the Morrison Formation (Upper Jurassic), approxi-
mately 5.5 miles southwest of Uravan, Montrose County, Colorado.

Collector. — Rodney Scheetz and family.

Repository. — Brigham Yomig University, Earth Sciences Mu-
seimi collections (No. ESM-171R).

A selection of the more diagnostic bones is illustrated in Figure 1 .
All the bones of ESM-171R are very similar to the corresponding
elements of YPM 1876, the holotype of the hypsilophodontid dino-
saur Laosaurus altus Marsh (1878), which is the type species of
the genus Dryosaurus Marsh (1894). This specimen of Dryosaurus
altus is from the Morrison Formation at Como Bluff, Wyoming,
and it has never been adequately figiu-ed. The hypsilophodonts of
the Morrison Formation are currently being studied by one of us
(Galton), who considers Dryosaurus a valid genus. Apart from the
form of the teeth and of the femora, the bones of ESM-171R are
also very similar to the corresponding elements of Hypsilophodon

^Department ol Biology, University ol Bridgeport. Bridgeport, Connecticut 06602.
-Eartli Sciences Musoiini. Brigliam Young University.

129

(IREAT BASIN NATURALIST

Vol. 33, No. 2

Fig. 1. Dryosaurus alius (Marsh), referred specimen BYU ESM-171R,
Morrison Formation, Colorado. A, fragment of left dentary in medial view;
B, distal end of left humerus in posterior view; C, proximal end of left radius
in medial view; D, distal end of left rathus in medial view; E, proximal end
of left femur in posterolateral view; F. distal end of right fenun- in posterior
view; G, proximal end of left fenuu' in anterior view; H, (Hstal end of left
fenun- in anterior view; I, proximal end of right tibia in lateral view; J, distal
end of left tibia in anterior view; K, distal end of right tibia in posterior view;
L, distal end of left fibula in anterior view; M, proximal surface of left astraga-
lus. A X 2 natural size; B-M natural size; horizontal lines represent 10 mm.

June 1973 galton, jensen: fossil dinosaur 131

foxii Huxley (Lower Cretaceous, England), the anatomy of which
will soon be described in detail (Galton, in press) .

There are several jaw fragments containing teeth that are very
similar to those of Dryosaurus. In the largest fragment (Figure lA),
replacement teeth are visible, and, as shown for most other reptiles
by Edmund (1969), the teeth in each alternately numbered series
were progressively replaced in sequence from back to front along
the jaw, i.e., left to right. There are many isolated centra of dorsal
vertebrae, typically ornithopod in form, and the lengths range
from 11 mm to 31 mm. The femora (Figures lE-H) are very sim-
ilar to those of Dryosaurus. Proximally, the lesser trochanter is tri-
angular in cross-section; it is separated from the greater trochanter
by a deep cleft (Figure IG); and distally the anterior intercondylar
groove is quite deep (Figure IH). A rather eroded proximal end is
almost twice the size of those illustrated (Figures IE, IG).

Assuming similar body proportions to those of the much more
complete material of Hypsilophodon (see Galton, in press), it is
possible to give some idea of the sizes of the individuals represented
by the bones of ESM-171R. The smaller of the dorsal centra and
the humerus (Figure IB), radius (Figures IC, ID), tibia (Figure
IJ), fibula (Figure IK), and astragalus (Figure IM) are probably
from animals with a body length of about 3 feet (0.9 m). The femora
(Figures lE-H) and tibia (Figure II) are probably from individuals
that were slightly smaller, and the small tibia (Figure IK) may be
from an animal only 2 feet (0.6 m) long. The large dorsal centra
and the eroded femur are probably from an animal about 5.0 feet
(1.7 m) long. In contrast, the holotype of Dryosaurus altus repre-
sents an animal of about 10.5 feet (3.2 m) in length. Specimens
representing a series of juvenile individuals are also known for the
hypsilophodonts Dysalotosaurus lettowvorbecki Pompeckj (Upper
Jurassic, Tanzania; see Janensch, 1955) and Hypsilophodon foxii
(see Galton, in press).

Grateful acknowledgement is made to Rodney Scheetz and his
family for discovery of the locality and for generously donating
study materials to the Earth Sciences Museum collections. We thank
Dr. John H. Ostrom of the Peabody Museum, Yale University
(YPM), for permission to study material in the collection under his
care. This paper was written while one of us (Galton) was in re-
ceipt of a Faculty Research Grant from the University of Bridge-
port, and was typed by Tempe Reith (University of Bridgeport).

Literature Cited

Edmund, A. G. 1969. Dentition, pp. 117-200. In C. Cans, A. d'A. Bellairs,

and T. S. Parsons (ed.). Biology of the Reptilia, Morphology A, Volume I.

Academic Press, New York.
Gai^ton, p. M. In press. The ornithischian dinosaur Hypsilophodon from the

Wealden of the Isle of Wight. Brit. Mus. (Nat. Hist.) Bull., Geol.
HuENE, F. VON, AND R. S. Lui.i,. 1908. Neubesclireibung des Originals von

Nanosaurus agilis Marsh. N. Jahrb, Mineral, Geol. u Palaont. 1908: 134-144.
Janensch, W. 1955. Der Ornithopode Dysalotosaurus der Tendaguruschichten.

Paleontographica Suppl. 7(3) : 105-176.

132 GREAT BASIN NATURALIST Vol. 33, No. 2

Marsh, O. C. 1877a. Notice of some new vertebrate fossils. Am. J. Sci. 14(3):
244-256.

. 1877b. Notice of new dinosaurian reptiles from the Jurassic Forma-
tion. Am. J. Sci. 14(3):514-516.

. 1878. Principal characters of American Jurassic dinosaurs. Am. J. Sci.

16(3):411-416.

1894. The typical Ornithopoda of the American Jurassic. Am. J. Sci.

48 (3): 86-90.

ScHUCHERT, C. 1939. The geological horizon of the dinosaurs Hallopus and
Nanosaurus agilis. Am. J. Sri. 237:19-26.

GREAT BASIN NATURALIST

Notice to Contributors

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TABLE OF COINPTENTS

Three' 'few f.pecies of Pahnoxylon from tlie Eocene Green

IMvor formation, Wyoming. William D. Tidwell, David

A. Medlyn, and Gregory F. Thayn

On the laxonomic status of Platypodidae and Scolytid.

(Coleoptera), Stephen L. Wood 77

Studies on Utah stoneflies (Plecoptera), Richard W. Ban-

marm

The male brachycistidine wasps of the Nevada Test Si

(Hyraenoptera: Tiphiidae). Marius S. Wasbauer

Notes on aquatic and seraiaquatic Hemiptera from the souii

western United States (Insecta: Hemiptera). John 1",

Polhemus 1 1 3

A new subfi-uticose Eriogonum (Polygonaceae) from wester

Colorado. James L. Reveal

Records of Coreidae (Hemiptera) from the Nevada Test Si

Dorald M, Allred

Density, growth, and home range of the lizai-d Uta stam-

huriana sleinegeri in soutJiern Dona Ana County, New

vTfixico, Richard D. Worthington and Edv\.

rvizo

Smal] bones of the hypsilophodontid dinosaur Dryosaurus

alius fi-om the upper Jurassic of Colorado, Peter M.

Galton aud James A. Jensen ..

Volume 33, No. 3
September 30, 1973

The

Great Ba

Published by
Brigham Young University

GREAT BASIN NATURAUST

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

Editorial Board: StanJ.ey L. Welsh, Botany, Chairman; Wilmer W.
Tanner, Zoology; Joseph R. Murdock, Botany; Vernon J.
Tipton, Zoologj"^; Ferron L. Andersen, Zoology

Ex officio Editorial Board Members:

A. Lester Allen, Dean, College of Biologicfd 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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All matters pertaining to the purchase of subscriptions and back
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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

riU]Db<^r.

The Great Basin Naturalist

Published at Provo, Utah, by

Brigham Young University

Volume 33

September 30, 1973

No.

ECOLOGY OF SCELOPORUS MAGISTER
AT THE NEVADA TEST SITE, NYE COUNTY, NEVADA

Wilmer W. Tanner^ and John E. Krogh-

Abstract. — The natural history of Sceloporus magister was studied in popu
lations at tlie Nevada Test Site, Mercury. Individuals were marked and kep
under surveillance from 1965 through 1970. Reproductive cycles were examine
by field observation and by autopsy. Hatchlings appear in the population in Jul;
and August. Growth is rapid for the first year. Sexual maturity is reached befor
the second hibernation or soon after emergence from it. Territoriality is a pre
nounced behaviorism; individuals have been observed to remain in a small are
for several years. Food consists of arthropods, with ants the predominant iterr
Reproduction occurs during May and/or June with only one clutch per yeai
Density is variable and depends on the appropriateness of the habitat.

Sceloporus magister are abundant on the rocky foothills, particu
larly where large rocks, wood poles or posts, and tree yuccas ^r
available for basking and hiding. Several other species of the spino
sus group have been studied. Blair (1960) studied the rusty lizar(
{S. olivaceus) in Texas for five years; Mayhew (1963) reported oi
the granite spiny lizard {S. orcutti) from southern California afte
a four-year study. Other reports involving species of the spinosu
group have been limited both as to time of observation and materia
available. Kauffeld (1943) reported a female S. clarki from southen
Arizona which contained 24 embryonate eggs. Davis and Smitl
(1953) stated that S. horridus in Morelos seemingly laid one clutcl
of eggs in late July and the first part of August. Davis and Dixo]
(1961) indicate a longer reproductive period for this species extend
ing from May to September.

Sceloporus magister has not been studied intensively. Severa
studies include references to egg laying, large eggs contained in .
female, and age groups. The studies of Axtell (1959), Mintoi
(1959), Smith, Williams, and Moll (1963), Taylor (1936), Steb
bins (1954), and Johnson, Bryant, and Miller (1948) refer to aspect
of its life history. A recent and more complete study is by Parke
and Pianka (1973).

Fitch (1970) adds information to several species and summarize
available reproductive data for the genus Sceloporus. Stejnege

'Department of Zoology, Brigham Young University, Provo, Utah 84602.
'College of Health, University of Utah, Salt Lake City, Utah.

133

134 GREAT BASIN NATURALIST Vol. 33, No. 3

(1893) provides no reproductive data, but does discuss habitat, be-
havior, and feeding habits. His lengthy comments pertain primarily
to the systematics of the larger spiny lizards of the Southwest.
Sceloporus orcutti and S. boulengeri are described as new and other
related species {clarkii, spinosus, magister) are discussed. Richards(m
(1915) referred only to the habitat and a few behaviorisms.

In 1939 Smith reviewed the genus Sceloporus, described several
new subspecies and set forth for the first time a phylogeny of the
genus. Phelan and Brattstrom (1955) discussed the geographic
variations in S. magister and described three new subspecies. Tanner
(1955) described as new the Upper Colorado Basin population. Lar-
sen and Tanner (1973) are preparing a phylogenetic study of the
entire genus using basically external measurements, scale patterns,
and skull characters in a computerized analysis. The populations of
this species occur in limited pockets, which may account for their
not having been studied as intensively as many other desert species
have been.

The present study was conducted at the Nevada Test Site under
the contract At( 11-1) 1496 between the U.S. Atomic Energy Com-
mission and Brigham Young University, Provo, Utah.

Study Plots

Four study areas have been used in securing the data. In each
area habitat conditions such as wood poles or posts, large rocks, or
trees (tree yuccas or cottonwoods) were present. In two areas the
populations were marked and studied from 1965 through 1969.

The Knoll study plot was established in 1965 and the first lizards
were marked on 3 July. All captures were made by noosing. The
plot consisted of a small rocky ridge rising from the bajada, isolated
from the other ranges by a small valley on the west and surrounded
otherwise by a desert alluvial plain. The lizards inhabit the ledges
and rock slides. Although we succeeded in marking 31 lizards and
had 26 recaptures, only one hatchling was seen in four years. The
area included in this study plot was approximately 2 hectares.

A study plot at Cane Springs was set up in 1966 and the first
lizard marked on 28 May. This plot consisted of two areas. One
contained the walls of an old rock house and frame room adjacent to
it. These are shaded by a large cottonwood tree. Down the slope
to the north are poles and posts and the remains of a corral; nearby
to the west are rocks and ledges. In this area captures were made
by noosing. The other area was south and west of the house. Near
the house is a patch of tall grass and brush, which undoubtedly bene-
fits from the shallow water table near the spring. To the west are
sparse desert shrubs on a rocky hillside. Three rows of cans were
placed through the grass and onto the hill, totaling 39 traps. Each
was placed 12.2 m apart, which provided a plat 147 x 37 m. By
including a narrow outer margin to the plot and the area of the
house, which is adjacent to the plot, there is approximately .5 hec-
tare. The corral area is about half this size. We marked 67 S.

Sept. 1973 TANNER, KROGH: LIZARD ECOLOGY 135

rnagister at Cane Springs and of these 15 were hatchlings. There
were 57 recaptures.

Two areas were used to secure lizards for autopsies. One area
consisted of large rocks along the road between Cane Springs and the
Knoll. The second was along the abandoned road from Mercury
Pass into Frenchman Flat. Along this road are power poles, some
ledges, and large boulders. All areas range in elevation between
1075 and 1230 m and are in approximately the same type of habitat
except as man has modified it.

Home Range and Density

Data for 15 individuals captured and recaptured for 3-10 times
show a definite territoriality. In most instances individuals remained
within a rather restricted area, except females who seemingly leave
their home range to lay eggs. Both sexes may leave if a more favor-
able area for hibernation is near the area used for summer activity.
The general size and shape of some home ranges observed for peri-
ods of 2-4 years were similar to those observed for S. occidentalis .
The areas occupied were surprisingly small for such a large lizard
(Fig. 1). Once a territory was established we observed little shifting
or enlarging. An adult female #1K was marked in 1965 and caught
once in 1966 and 1967. During these three years she was observed
within 4 m of each capture. A juvenile #5K captured 6 times in
1966 and 1967 was within an area having a diameter of not more
than 15 m. Two juveniles (#3K and #8K), marked on 29 May
and 8 June 1966, moved north along the ridge for approximately
310 and 620 m. Both were recaptured in subsequent years and both
remained in a small territory (Fig. 1).

At Cane Springs most of the lizards for which we have adequate
data were those living along the rock walls or the rows of posts.
Thus their territory consisted of a narrow strip which could not be
adequately represented by a polygon. An example is an adult male
(#18) marked on 9 July 1966 and last seen on 7 May 1970. During
five years he was recaptured nine times and observed at a basking
site on numerous occasions. Although our visits to this plot were at
intervals of several days and not for long periods of time, only on
two occasions did we observe him away from the line of posts, once
on a rock pile 20 m west and the last time about 200 m west on a
ledge.

Each of these territories possessed at least one area for basking or
observing, and, in most, several objects were used. Basking sites
were usually near a hiding place. For example, #18 used three
posts for basking and observation. He was usually high on the post
if not on its top. Our approach would send him scurrying from the
post and into one of several large bushes at or near the base of the
post. Posts, poles, trees, or rocks were not regularly used if some
protection (bushes, rock piles, etc.) were not nearby.

Density is difficult to determine because of the variations in
habitat. Although the total area at Cane Springs is smaller than at

136

GREAT BASIN NATURALIST

Vol. 33, No. 3

N 6.

*\\#9C

11.

. S

TO 18

Fig. 1. Home ranges for Sceloporus magister. The prefix K = Knoll study-
plot and C = Cane Springs study plot.

Sept. 1973 TANNER, KROGH: LIZARD ECOLOGY 137

the Knoll, we marked and recaptured 36 more lizards. In 1966
eight lizards were marked at the corral (about 32 per hectare) and
25 at the house and from the can traps (about 50 per hectare). In
1968 seven were at the corral and only 18 at the house and traps.
In 1969 twenty were seen at the house and on the plot. During 1966
and 1967 only 12 and 13 individuals were known to be at the Knoll
plot (about 6 or 7 per hectare).

Of the three areas for which density data are available, the house
and plot area supports the greater density. We recognize in this area
more shelter, adequate areas to climb and hide, and because of the
abundant vegetation more food. In this species the quality of the
habitat is seemingly very important in determining density. Because
habitat plays such an important part in the establishment of terri-
tories, population densities will vary from a few lizards to as many
as 50 per hectare.

Behavior

Desert scaly lizards are active from April to October. Both
adults and juveniles were seen 11 April 1970 and adults and hatch-
lings in mid-September. This species is a persistent basker and may
be seen perched on a rock, pole, or some elevated object soon after
sunup and until late in the afternoon. During the heat of the day
shade is sought. The basking habits are similar to those observed for
S. occidentalis longipes (Tanner and Hopkin, 1972). Their climbing
abilities are remarkable, particularly in trees, where they are very
adept at keeping the tree between you and them. In southeastern
Utan, this species is so proficient in climbing that collecting is diffi-
cult without a gun. In this regard we have found magister to be simi-
lar to clarki and presumably olivaceus (Blair, 1960).

Sceloporus magister is a large, heavy-set lizard which usually
produces a rustling sound as it scurries for cover. Because of its
brisk scampering, ranchers in southern Utah call it the bull lizard.
Although it is not as speedy as some lizards, in its habitat of trees,
boulders, and poles it is well equipped to escape a pursuer.

Individuals have been recorded to remain on an object for several
hours, basking at times and/or loafing in the shade. Often they
cling head down as if watching for prey or an approaching enemy.

For active individuals we have cloacal temperatures ranging from
29 to 35 C and averaging 32.8 C. This is within one-tenth of a degree
from the average determined for S. occidentalis (Tanner and Hopkin,
1972) and only .2 C (32.6) from the mean temperature for a large
series of S. graciosus from central Utah (Burkholder and Tanner,
1973). We suspect that higher maximum temperatures are reached
but question that this species is very tolerant of the high tempera-
tures of other species inhabiting the more open desert area. Scelopo-
rus magister seems to confine its activities to areas where shelter and
shade are available. Their tree climbing may also contribute to some
temperature control by permitting individuals to get well above the
intense heat of the desert floor where breezes provide for air circula-
tion.

138 GREAT BASIN NATURALIST Vol. 33, No. 3

A male and female, particularly in the spring, often occupy the
same basking site. As the season progresses there are fewer pairs
seen. Usually by June the courtship is over and females occur singly.
Copulation was not observed.

Reproduction

The reproductive cycles in S. magister are surprisingly similar
to those recorded for S. occidentalis (Tanner and Hopkin, 1972) and
for S. graciosus (Burkholder and Tanner, 1973). Although the gen-
eral reproductive patterns are similar, there are some variations
which will serve to distinguish magister from other Sceloporus seen
by us.

Male Cycle: Although our gonadal series for some months is not
large, a plotting of the data does provide a curve which seemingly
depicts the cycle (Fig. 2). As in other species in the genus, the cycle
moves slowly except for a brief period in June. At this time there is
a sudden change resulting in a considerable decrease in gonadal
weight and a rapid increase in size and weight of the fat bodies
(Table 1). A fat body of five grams was recovered and is unusually
large; however, we have other records for weights between two and
three grams. In magister these large fat bodies are lobate as the
liver but are readily distinguished by the grey greenish color.

Courting was observed in May, and pairs were regularly seen
occupying the same basking site. The gonadal cycle suggests that
mating occurs during May or early June. By mid-June the semi-
niferous tubes are undergoing a rapid regression, reaching a low
point in late July or August (stage 8, after Mayhew, 1971), after
which lizards are seen singly and courtship has ceased. The cycle is
renewed (stages 1 and 2) in late August and September. We have
not seen gonadal examples for late September or October. Examples
seen in early April are in stages 4 and 5 suggesting that some de-
velopment does occur between mid-September and the time of emer-
gence in March. Stages 6 and 7 occur in May and June.

Males are not sexually mature until after the second hibernation.
Although growth is rapid and continuous during that first full season,
the gonadal cycle does not begin until the late summer and fall
before the second hibernation.

Female cycle: Our earliest gonadal samples are for 8 April 1972.
On this date the size of testes was much larger than ovaries in com-
parably sized lizards. The males were more obvious in the habitat
and usually were in a dominant position at the basking sites. Indi-

Table 1 . Gonadal and fat body cycles in male S. magister from the Nevada
Test Site.

June
Cycle April May early late July August Sept.

Gonadal 0.45(4) 0.52 0.51(3) 0.20(4) 0.08(4) 0.035(8) 0.12(3)
Fat Body 0.71 0.12 1.20 1.62 1.36 0.91

Sept. 1973

TANNER, KROGH: LIZARD ECOLOGY

139

September

Fig. 2. The reproductive cycle as determined by the weight of testes and
fat bodies, plotted in milligrams.

cations are that magister female behavior is similar to other northern
Sceloporus species {occidentalis and graciosus) in that they emerge
after the males have established their territories and are less obvious
in the habitat until after mating.

Gravid females were observed in the field during late May and
June. In 1966 and 1967 they were seen from 28 May until 16 June.
In 1969, they were observed from 10 June to 28 June. Autopsied
lizards that year showed oviducal eggs in mid-June. We note one
exception in which there were 6 yolking ova, 7 mm in diameter,
large fat bodies (1.06), and no corpora lutea for 28 June 1969. She
was 97 mm SVL indicating adult size. We have noted an occasional
female individual in other species to be slower than most others in a
population.

140 GREAT BASIN NATURALIST Vol. 33, No. 3

Records for the Mercury Valley plot indicate that adult S. magi-
ster are in the plot during late May and June; we have only one
record for July. Adults entering the plot were two females to each
male and with few exceptions they were gravid. We saw one
juvenile and five hatchlings during five years. About 300 m west
(up slope) from the plot are some ledges and rocky habitat. We
believe that the plot and adjacent areas were used as an area for egg
laying by migrating females. We have one record in which No. 14
was marked on 11 June 1968, noted to be gravid, and weighed 34.28
g. On 28 June she was not gravid and weighed only 28.54 g. Most
individuals were seen only once. Five were seen twice and with one
exception (above) were caught within one week.

The study plot data do not satisfy completely our belief that the
females of magister leave their home territory for depositing eggs as
was reported for S. olivaceus by Blair (1960). While night driving,
we saw a few female magister along roads on the test site. Invari-
ably these were gravid females, perhaps attracted to the warm roads
as they moved to an area for egg laying.

Four clutches of 4, 7, 7, and 10 eggs were laid in the laboratory
on 25 June, 1972 and 20 and 26 June and 20 July, 1973. The clutch
of 4 eggs averaged 0.86 (.75-.97)g and were 16.5 x 9 to 17.0 x 10
mm in size. Some captive females are reluctant to lay in confined
areas. We suspect that the clutches listed above were held in the
oviducts longer than is normal and thus the later dates. Two clutches
of oviducal eggs were taken from autopsied lizards on 18 June, 1969.
There were five and seven eggs per clutch, averaging 21 x 11 and
18x10 mm in size. On the basis of the size and weight of the one
clutch, we estimate that the female in Mercury Valley (No. 14) laid
six eggs. Available data suggest that the clutch size for magister
in southern Nevada is in the range of 4-10 (avg. 7) eggs per clutch.
This is noticeably less than in such species as occidentalis (Tanner
and Hopkin, 1972) with 12, olivaceus (Blair, 1960) with 10, clarki
(Kauffeld, 1943) with a record of 24, and orcutti (Mayhew, 1963)
with 10-12.

Taylor (1936) reports a female from Sonera containing 18 well-
developed eggs and Stebbins (1954) lists four clutches from lizards
taken in southern California and central Baja California. These
range from 7-18 eggs. The size of the females was not recorded and
we cannot correlate numbers of eggs to the size (and perhaps the
age) of the females. We do note in the size of eggs a relationship to
numbers. Data on egg size are available from seven clutches (Steb-
bins, 1954, 4; our data, 3). The larger eggs were measured in the
smaller and medium sized clutches and the smallest in the clutch of
18 eggs.

Apparently only one clutch is laid per year. Yolking and ovi-
ducal eggs are found in autopsied lizards from late May through
June. None have been found in July. Field data also indicate that
gravid females occur in the })opulation from late May to late June
but with none in July. In all years studied, by far the greater

Sept. 1973 TANNER, KROGH: LIZARD ECOLOGY 141

number of gravid females was seen during the first two weeks of
June.

Records from the literature also place egg laying during late May
and June. Apparently this is the case for those populations occurring
much farther south than southern Nevada, where the seasons are
longer. Although our data and that from the literature are limited,
we can see no indications that S. magister lays more than one clutch
per year.

Soon after ovulation the fat bodies begin to increase in size.
They are smallest in late May and until the eggs are laid in June
(usually by 20 June). There is not always a complete loss of the
fat body. The least we have recorded is 0.06 g on 18 June 1969.
By July six females averaged 0.74 g and in August 1.06 g. In none
of the females is the fat body as large as those recorded for males
during July and August. The fat body cycle is similar to that ob-
served in males (Fig. 2) but with an apparent lag of approximately
two weeks after the eggs are ovulated.

Growth and Longevity
Growth

Hatchlings: This age group has been observed from 1966
through 1969. The earliest record for a hatchling is 27 July
1966. In 1969 the first was seen on 28 July. The smallest SVL
recorded is 32 mm for one measured at the Mercury Valley plot on
22 August 1966. Two measured 34 mm at Cane Springs on 27 July
1966. The largest hatchling measured 41 mm on 7 August 1966
(Table 2). Although our data are not extensive and are incomplete
for the fall months, they do indicate an extended hatching period
of approximately one month from late July to late August.

Size of recent hatchlings seems to range in SVL from about
30-35 mm. From this size they grow rapidly, and some approach
50 mm by the time of hibernation. Growth for individuals was not
determined; however, if we assume that those measuring 32-34 mm
SVL are recent hatchlings (with weights of 1.00-1.50 g) then most
hatchlings at least double their weight by the time of hibernation.
Lizards having an SVL of 38-41 mm in August weighed 1.75-2.12 g.

Juveniles: More juveniles were captured and recaptured than
hatchlings. This was particularly the case for the Knoll, where few
hatchlings were seen. Hatchlings were flighty and readily sought
cover when approached. Juveniles were bold and seemed to reappear
soon after their initial hiding. Growth is rapid after emergence from
the first hibernation. In April and May for most years measure-

Table 2. Range and average SVL for twenty
Springs and Mercury Valley study plots.

hatchlings

from the Cane

Year No.

Males

No.

Females

1966C 5
1968C 2
1966M 3

37.2(34-41)
36.5(36-37)
37.2(35-40)

3
5

36.3(34-37)
36.2(34-38)
34.0(32-36)

142

GREAT BASIN NATURALIST

Vol. 33, No. 3

Table 3. Range and average SVL for thirty-seven juveniles.
C = Cane Springs, F = Frenchman Flat)

Year

No.

Males

No.

(K = Knoll,

Females

1966K
1967K
1968K
1969K
1966C
1967C
1970F
Total /Avg.

5
3

2
2

4
16

65.4(57-70)
63.7(60-66)

73.0(71-78)
67.5(65-70)

58.2(52-66)
64.5(52-78)

2
3
3
3
5
2
3
21

62.5(62-63)
53.7(50-57)
62.3(59-68)
75.0(71-77)
57.6(53-60)
61.0(59-63)
60.0(55-65)
61.2(53-77)

merits for SVL range between 50 and 60 mm, with a few in the low
sixties. In Table 3 juvenile SVL measurements include those for
April through July. August measurements are usually in the seven-
ties and some individuals are approaching the size of adults and
exhibit an adult color pattern.

Table 4 provides growth data for ten individuals. In most cases
growth approximated 0.3 mm per day or a growth during May,
June, and July of 25-30 mm. Such a growth rate is not attained by
all individuals. We also note a slowing of the rate in July, perhaps
because of a reduced food supply resulting from the hot and dry con-
ditions which usually prevail. Late springs may retard the rate
early in the season whereas an early spring provides for a rapid as
well as a longer growth period.

Table 4. Measurements and growth rates for selected juveniles after the
first hibernation. (C = Cane Springs, K = Knoll)

Toe
Number

Date

Sex

SVL

Days
Growth

Total
Growth

Growth
per day

12
17

June 66
July 66

53
65

12mm

0.343

16
25

June 66
July 66

60
75

15mm

0.384

June 66
July 66

12mm

0.480

1-6C

16
7

June 67
Aug. 67

19mm

0.365

2-8C

17
9

June 68
July 68

59
67

8mm

0.364

2-9C

17
28

June 68
June 68

68
70

2mm

0.182

2-13C

July 68
Aug. 68

69
74

5mm

0.208

31
15

May 66
July 66

57
72

15mm

0.333

May 66
July 66

68
83

15mm

0.333

8
29

June 66
June 66

63
65

2nmi

0.095

Total /Avg.

319

lOSmmi

0.333

Sept. 1973 TANNER, KROGH: LIZARD ECOLOGY 143

Table 5. Measurements for adults from four study plots at Nevada Test
Site for the years 1965 through 1971.

Plot

No.

Males

No.

Females

Cane Springs
Frenchman Flat
The Knoll
Mercury Valley

98.6(84-114)

101.6(87-112)

97.4(83-115)

96.0(90-100)

93.2(88-102)
97.6(91-107)
92.0(81-102)
95.8(88-103)

Total/Avg.

99.40

95.04

Juveniles reach adult size by the time they hibernate in October.
In September the gonadal cycle is paralleling that of adults and,
except for a smaller size, these juveniles are functionally adults.

Adults: There is a slight sexual dimorphism in size with males
averaging approximately 5 mm larger in SV lengths (Table 5). The
smallest sexually mature female was 81 mm and the smallest adult
male recorded was 83 mm SVL. The largest individuals in the popu-
lation are males (Table 5). Data indicate that few females yolk eggs
before they are approximately 90 mm SVL.

Adults continue to grow for at least three years. Young adults
emerging from the second hibernation usually have a sustained
growth for the first three months (April through June). This is not
as great a growth in length as in hatchlings and juveniles; however,
it does result in a noticeable increase in weight. By this time they
have reached full adult size (90-100 mm) and have completed the
first reproductive cycle. Examples of growth after the second hiber-
nation are listed in Table 6. Records for individuals more than 30
months old show them to be 90 mm or more in SVL. A male indi-
vidual from Cane Springs (18) marked on 9 July 1966 was 87 mm
SVL, and estimated to be about 23 months old since no other ju-
venile seen by us reached such a size before late August. On 4 June
1968 he was 110 mm and on 7 May 1970, 114 mm. Although some
individuals grow more rapidly than others, our data indicate that
individuals measuring 100 mm or more in SVL are the older mem-
bers of the population perhaps 4 or more years of age.

Table 6. Measurements and growth rates for selected individuals as juve-
niles and as adults after the second hibernation. (K = Knoll)

Toe
Number

Date

Sex

SVL

Total
Growth

Age
(approximation)

29 May 66
1 June 67

62
81

19mm

10 mo
22 mo

31 May 66
2 June 67

57
82

25mm

10 mo
22 mo

29 June 66
10 July 67

22mm

11 mo
23 mo

19K

13 July 67
4 June 68

22mm

11 mo

22 mo

1-5K

19 June 68
28 June 69

60
86

26mm

11 mo
23 mo

144

GREAT BASIN NATURALIST

Vol. 33, No. 3

Longevity

We have one record for a male known to be in his sixth year.
Two males were marked at Cane Springs on 9 July 1966. Number
17 was a large adult, 110 mm SVL weighing 54 g. We estimate
him to be at least 4 years old when marked. He was last seen on
15 June 1968, the same length but only 48 g. Number 18 was
smaller, measuring 87 mm SVL and weighing 24 g. He was in his
first year as an adult. He was last seen on 7 May 1970, appeared
to be healthy, and weighed 59.13 g. Several records are for four
years and two for at least five years.

Records for marked individuals and for those compared when
autopsied indicate that the adult population consists mostly of lizards
3-5 years old but with a few older ones. The two oldest are both
males; however, we do not consider this to imply that males live
longer than females. The oldest females were 4 years. Our data are
too limited to establish longevity differences between the sexes.

Food and Feeding Habits

This species is primarily an insectivorous feeder. Fig. 3 lists
the types of food eaten as a percentage of individual items consumed.
Only stomach contents were analyzed. Knowlton and Thomas
(1934) and Knowlton and Nye (1946) reported on the contents of
49 and 12 stomachs from southern Utah. In both studies, ants were
dominant food items. The desert scaly lizard, as with other scelo-
porines we have examined, is an opportunistic feeder. One adult
male contained 1129 small ants. Four adults had eaten Isoptera and
in each case large numbers (78-330) were consumed. The usual
variety of arthropods are included in the diet throughout the year
with an apparent increase in ants during July. This we do not
understand unless it is attributable to the decline of vegetation
(annuals and spring perennials) and the decline of insects as a result
of the heat and drought; these conditions have apparently little effect

1965

1968

1969

Fig. 3. Precipitation for an eight-year period 1965 to 1972. Dark columns
for Cane Springs, dots for Well 5B (weatherstations in Frenchman Flat).

Sept. 1973

TANNER, KROGH: LIZARD ECOLOGY

145

Li zard ( 1 )

Other Arthropods

Round worm s

Fig. 4. Stomach contents plotted as a percentage of the items found in
twenty-one stomachs in 1969.

in reducing the numbers of ants. Eighteen of 22 stomachs contained
ants, which is higher than the ratio (29 of 49) found by Knowlton
and Nye (1946). Knowlton and Thomas (1934) found one small
Cnemidophorus t. tigris and some seeds and berries in stomach con-
tents. We found one hatchling S. magister; we also found that sev-
eral had consumed a number of plant parts, including 21 Lycium
berries, a number of small rocks, and 18 round worms. A general
list of the items recovered is in Fig. 4.

Literature Cited

AxTELL, R. W. 1959. Amphibians and reptiles of the Black Gap Wildlife
Management area, Brewster County, Texas. Southwestn. Nat. 4:88-109.

146 GREAT BASIN NATURALIST Vol. 33, No. 3

Blair, F. W. 1960. The rusty lizard: a population study. Austin: Univ. Texas

Press.
BuRKHOLDER, G. L., AND W. W. Tanner. 1973. Life history and ecology of

the Great Basin sagebrush swift, Sceloporus graciosus graciosus Baird and

Girard, 1852, in press.
Davis, W. B., and J. R. Dixon. 1961. Reptiles (exclusive of snakes) of the

Chilpanciugo region, Mexico. Proc. Biol. Soc. Washington, 74:37-56.
, AND H. M. Smith. 1953. Lizards and turtles of the Mexican State of

Morelos. Herpetologica 9:100-108.
Fitch, H. S. 1970. Reproductive cycles of lizards and snakes. Mis. Publ. Mus.

Nat. Hist. Univ. Kansas, No. 52, pp. 1-247.
Johnson, D. H., M. D. Bryant, and A. H. Miller. 1948. Vertebrate animals

of the Providence Mountains area of California. Univ. Calif. Publ. Zool.,

48:221-376.
Kauffeld, C. F. 1943. Field notes on some Arizona reptiles and amphibians.

Am. Mid. Nat. 29:342-59.
Knowlton, G. F., and W. L. Thomas. 1934. Notes on some insectivorous

Utah lizards. Copeia 1934 (1):10-14.
, AND W. P. Nye. 1946. Lizards feeding on Ants in Utah. Jour. Econ.

Ento. 39(4) :546.
Larsen, K. R., and W. W. Tanner. 1974. The phylogeny of the genus

Sceloporus. Herpetologica, in press.
Mayhew, W. W. 1963. Reproduction in the granite spiny lizard, Sceloporus

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Minton, S. a., Jr. 1959. Observations on amphibians and reptiles of the Big

Ben region of Texas. Southwestn. Nat. 3:28-54.
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Richardson, C. H. 1915. Reptiles of Northwestern Nevada and adjacent terri-
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Smith, H. M. 1939. The Mexican and Central American lizards of the genus

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FURTHER STUDIES ON THE WASPS OF JACKSON HOLE,
WYOMING (HYMENOPTERA, ACULEATA)

Howard E. Evans^

Abstract. — In a report published in 1970, 190 species of wasps were reported
from Jackson Hole, Wyoming, and notes were presented on the biology of several
ground-nesting species. In this paper, records of 18 additional species are pre-
sented. A short account of the biology of a ground-nesting species, Tachysphex
aethiops Cresson, is included. Also included are accounts of four species that
accepted wooden trap nests during the summer of 1971: Symmorphus cristatus
(Saussure), Ancistrocerus adiabatus (Saussure), both Eumenidae; Trypoiylon
aldrichi Sandhouse, Passaloecus cuspidatus Smith, both Sphecidae. Notes are pre-
sented on nest structure, prey, and parasites in each instance.

This is a brief supplement to my study of the ecology and be-
havior of the wasps of Jackson Hole, Wyoming (Evans, 1970).
That paper summarized three summers of study over a six-year
period (1961-1967). I returned to Jackson Hole in the summer of
1971, primarily to round out my work on the digger wasp genus
Philanthus. My data on Philanthus will be saved for a proposed
survey of the comparative behavior of members of that genus. The
present paper includes additions to the faunal list and notes on the
nesting behavior of five species not or only briefly studied earlier.

In my 1970 paper I reported 190 species of wasps from Jackson
Hole. Eighteen species are here added to the list, bringing the total
to 208. Since my earlier studies were concerned with ground-nesting
wasps, I put out 300 wooden trap nests in 1971, hoping to learn
something of the distribution and behavior of twig-nesting species.
Although the percentage of acceptance was high, only four species
of wasps occupied these nests. The results of the trap-nesting work
are summarized in the last section of this paper.

A general description of the study areas will be found in my
1970 paper. Study area no. 3, listed as 4 miles SW of the Elk post
office and indicated by the word Elk in the text of that paper, should
perhaps be qualified, since there is evidently no longer a post office
at Elk. This area is on the Snake River some 9.5 km southwest of
Moran Junction, about 1 1 km south of the research station. For the
sake of consistency, I have continued to call it Elk in the present
paper.

I am, as usual, much indebted to the authorities of Grand Teton
National Park for permission to collect specimens for identification,
and particularly to the staff of the Jackson Hole Biological Research
Station for use of their facilities. I am indebted to several specialists
for identifying certain Sphecidae: Frank Parker (Astatinae), David
Vincent (Passaloecus), Richard Bohart (Gorytes), and Joanne E.
Slanky {Nitelopterus) . Insect prey were identified by specialists
of the Entomology Research Laboratory, U.S. Department of Agri-

*Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138.

147

148 GREAT BASIN NATURALIST Vol. 33, No. 3

culture, as indicated in the text. Spider prey were identified by
H. W. Levi of the Museum of Comparative Zoology.

I. Additions to the Faunae List

Family Chrysididae

Omalus cressoni (Aaron). Pilgrim Creek, 1 9, note no. 2299B: trap-nest collected
22 Aug. 1971, 1 9 emerged 7 May 1972 (nest was that of Passaloecus cuspi
datus; female of host had emerged 24 April 1972).

Family Tiphiidae

Publication of Allen's (1971) study of the genus Tiphia in western North
America permits me to clarify the species listed as "near essigi" and to add two
additional species to the list.

Tiphia unguis Allen. This species was described by Allen (1971), the $ holotype
and several paratypes (2 9 9,3 $ $) being from Elk, 4-10 Julj' 1964, with
additional paratypes from Moran (1 9,1 5,11 Aug. 1964) and from Jermy
Lake (1 9 , 20 July 1936).

Tiphia barberi Allen. Allen reports 3 9 9 and 1 5 from the Grand Tetons,
collected by E. C. VanDyke in June 1938.

Tiphia nevadana Cameron. This is a senior synonym of T. essigi Allen, and the
records for "sp. near essigi" from Moran and Elk should be transferred to
this species; they are so recorded by Allen (1971).

Family Masaridae

Pseudomasaris marginalis (Cresson). Death Canyon, 17 July 1971, 3 9 9,2 $ $,

9 9 on Phacelia sericea.
Pseudomasaris zonalis (Cresson). Pilgrim Creek, 2 Aug. 1971, 2 9 9.2 5 5;

Death Canyon, 17 July 1971, 2 9 9,1 $.

Family Pompilidae

Anoplius (Pompilinus) insolens (Banks). Signal Mt., 7300-7700 feet, 28 July
1971, 1 9.

Family Sphecidae
Subfamily Pemphredoninae

Passaloecus relativus Fox. This species should be deleted from my original list.
David Vincent, of Utah State University, is currently studying this genus
and has examined the specimens reported as P. relativus and found them to
consist of the following three species.

Passaloecus cuspidatus Smith. Moran, 6 July - 23 Aug. 1964, 1967, 4 9 9,2 5 5,
1 9, note no. 1992: taken as prey of Philanthus pulcher; Huckleberrj' Hot
Springs, 11-31 Aug. 1967, 299; Snake River, Elk, 9 Aug. 1967, 1 9. note
no. 2144: taken as prey of Philanthus pacificus. This species accepted trap
nests in considerable numbers during 1971; the results of trap nesting are
discussed on a later page.

Passaloecus armeniacae Cockerell and Fox. Moran, Aug. 1967, 2 9 9.

Passaloecus melanocr us Rohwer. Moran, July-Aug. 1967, 1 9, 1 5-

Subfamily Astatinae

Astata mexicana Cresson. Signal Mt, 7300-7700 feet, 28 July 1971, 5 5 5-
Aslata nevadica Cresson. Signal Mt., 7300-7700 feet, 28 July 1971, 1 9.

Sept. 1973 EVANS: Wyoming wasps 149

Diploplectron brunneipes (Cresson). Pilgrim Creek, 2 Aug. 1964, 1 9; String
Lake, 3 Aug. 1961, 1 ?; Snake River, Elk, 14-26 July 1971, 1 $; Moran,
19 July - 17 Aug. 1967, 1 9, 2 5 5, 1 ?, note no. 2163: taken from nest
of Philanthus pacificus, as prey.

Diploplectron fossor Rohwer. Moran, 4-30 July 1961, 2 9 9,2 $ $ ; Snake River,
Elk, 9 Aug. 1967, 1 ?, note no. 2144: taken from nest of Philanthus pacificus,
as prey.

Diploplectron peglowi Krombein. Moran, 18 July 1967, 1 $ .

Subfamily Larrinae

Nitelopterus laticeps Ashmead. Pacific Creek, 12 Aug. 1971, 1 9. This species
has been studied by Powell (1967) under the name A^. calif ornicus
(Ashmead).

Nitelopterus maurus Rohwer. Moran, July 16, 1961, 2 $ $. This record was
originally entered under the name A^. cyanurus (Rohwer), but I am in-
formed by Joanne E. Slansky that they should be called A', maurus.

Subfamily Nyssoninae

Gorytes flagellatus Bohart. Snake River, Elk, 4-10 Aug. 1964, 1 $ ; Moran, 6

Aug. 1964, 1 9 on Perideridia gairdneri.
Gorytes provancheri Handlirsch. Signal Mt., 7300-7700 feet, 28 July 1971, 2 9 9.

Subfamily Philanthinae

Cerceris calcohorti Rohwer. Snake River, Elk, 29 July 1971, 2 $ $, note no.
2260: taken from nest of Philanthus zebratus nitens Banks, as prey.

II. Notes on a Ground-nesting Sphecid

Tachysphex aethiops Cresson

This is a common wasp in sandy soil along the Snake River,
but I was able to report only one incomplete, parasitized nest in 1970.
At 4:00 p.m. on 16 August 1971, I observed a female at a nest in
firm, flat sand about 5 meters from the river (area M0R-A2 of the
1970 paper). The hole was open and there was a low, spreading
mound of sand, 8 cm wide by 5 cm long, in front of the entrance.
The female emerged from the burrow periodically and rose in the
air to a height of 0.8 m, with her feet dangling, facing the burrow.
The significance of this behavior was not apparent, as she soon began
filling the burrow persistently, emerging again and again to scrape
in sand from the entrance. I captured the wasp when the burrow
was nearly filled and excavated the nest. It proved to be surprisingly
shallow, an oblique burrow only 10 cm long terminating in a cell
only 4 cm deep. This cell contained a paralyzed immature acridid
grasshopper, Trimerotropis sp., probably T. suffusa Scudder [det. A.
B. Gumey], bearing the egg of the wasp. Immediately beyond it,
separated by only a thin barrier of soil, there was a second cell, con-
taining an immature acridid of the same species, also bearing an egg.
In both cases the egg was laid on the "throat" of the prey, with one
end glued to the left front coxal membrane, the egg extending trans-
versely over the right coxa, its posterior end free. Both cells ob-
viously contained the full complement of prey, as the female was

150 GREAT BASIN NATURALIST Vol. 33, No. 3

making a final closure, and both grasshoppers exceeded the wasp in
size (12 and 17 mm in length, as compared to 11 mm for the wasp).
This compares favorably with data reported in 1970, for that nest
also contained a single immature grasshopper somewhat larger than
the wasp.

III. Results of Trap Nesting

During the first week in July 1971, I put out 300 trap nests,
about half near the research station at Moran and half along Pilgrim
Creek in Teton National Forest. Techniques were those described
by Krombein (1967) in his well-known book on this subject; for
terminology see his book, p. 18. Of the 300 nests, 200 were pieces of
pine of the type used by Krombein, while 100 were sections of
Sambucus stems cut to about the same length (15 cm), part of them
bored and part with the pith intact. The Sambucus stems proved
unsuccessful, only one being accepted by a wasp and 5 by bees. Of
the 200 standard type, 65 were accepted by wasps in the course of
the summer, 57 by bees. Those accepted by bees were turned over
for study to Stephen L. Clement of the University of California at
Davis; they are not considered here except in cases of supercedure
involving a bee and a wasp. Trap nests wore overwintered in Massa-
chusetts, and all emergence occurred in April and May, 1972, there
being but one generation per year of these species in Jackson Hole.

Although the percentage of acceptance was reasonably high, I
was surprised to find that only four species of wasps (2 Eumenidae,
2 Sphecidae) were involved. Many other twig-nesting wasps are
recorded from Jackson Hole, but evidently they did not lend them-
selves to these techniques. Although these four species occurred in
much the same habitats, the kinds and numbers of parasites emerg-
ing from the nests were very different. Symmorphus cristatus, for
example, was very heavily parasitized by the miltogrammine fly
Amobia distorta Allen, and the other two mud-users had a low inci-
dence of parasitism by Amobia. On the other hand, the resin-user
Passaloecus cuspidatus appeared to be immune from attacks by
Amobia, although it was the only one of the four to be parasitized
by Omalus cuckoo wasps. Table I summarizes the results of the
rearing of parasites.

Table 1. Parasites reared from nests of trap-nesting wasps.

No. Nests Approx. No. Parasites Reared

Species of Wasp No. Cells Amobia Anthrax Chrysididae

Symmorphus cristatus
Ancistrocerus adiabatus
Trypoxylon aldrichi
Passaloecus cuspidatus

^Amobia distorta (Allen)
^A. floridensis (Townsend)
'Chrysis caerulans Fabr.
*Omalus, 3 species

36'

120

Sept. 1973 EVANS: Wyoming wasps 151

Symmorphus cristatus (Saussure)

This was the most abundant wasp in trap nests and also the
most heavily parasitized. Twenty-five trap nests were accepted, all
at Moran. Most were associated with dead wood, either on the walls
of log cabins or on stumps or standing or fallen dead pines and
aspens. Only four of the nests were on live pines, one on a live
willow. Traps accepted were from 0.3 to 2.5 meters high. Of the
25, 18 were in 4 mm borings, 7 in 6 mm borings. The number of
cells per 13 cm boring was small compared to Ancistrocerus adiaba-
tus: from 2 to 6, the majority having 3 or 4. Cell length varied from
8 to 24 mm (mean 15.5). Nest architecture was remarkably varied
and is difficult to summarize. Closing plugs varied in thickness from
2 to 15 mm, most being 3 to 5 mm. As in Krombein's (1967) nests
from New York and Virginia, all had an empty vestibular cell just
inside the closing plug, and several of these cells were subdivided by
partitions. Altogether the vestibular cells varied in length from 1 7 to
55 mm in length, with one exceptional one measuring only 5 mm.
In addition to several having one or two partitions within the vestib-
ular cell, two had barriers 3-4 mm thick (as compared to 1-2 mm
thick partitions) essentially dividing the vestibular cell into two.
For example, one nest had two cells at the end of the boring followed
by two long, empty cells, 35 and 45 mm long, separated by a barrier
4 mm thick and closed on the outside by a plug 5 mm thick.

Intercalary cells, between filled cells, were also variable in oc-
currence and in length. In many cases they consisted of no more
than two partitions separated by a space of only 1-3 mm; two nests
had all or most of the cells separated by such double partitions, and
one of these had one triple partition. One other nest had two distinct
intercalary cells measuring 4 and 9 mm. The first cell was normally
placed at the terminus of the boring, with no preliminary plug, but
several had empty spaces preceding the first partition and cell, these
spaces varying in length from 12 to 73 mm. A number of the more
heavily parasitized nests had partitions that were partly destroyed
by maggots w^hen they were harvested; hence it is not possible to
present accurate quantitative data on nest architecture.

Eggs were found in several cells, without exception suspended
from the roof of the cell by a short filament, near the partition at
the deep end of the cell. The prey are brought in after the egg is laid,
from 3 to 10 (usually 4 to 6) being supplied per cell. Without ex-
ception the prey consisted of a single species of external, leaf-feeding
beetle larvae (Chrysomelidae,Chrysomelinae [det. R. E. White]).

There were four instances of supercedure. In two cases Symmor-
phus cristatus had built 3-5 cells following 3-4 cells of the bee
HopUtis (Formicapis) robusta (Nylander).^ In one case the bee
superceded was an unidentified species of Hylaeus. In still another
case a Symmorphus had filled a single cell deep in a boring and was

^Determination by Dr. G. C. Eickwort of Cornell University. Dr. Eickwort informs me that
this species has usually been called H. dypeata (Sladen) and that H. robusta represents a new
combination. Nothing has been published regarding the biology of this bee, but Stephen Clement
and Richard Rust, of the University of California at Davis, are preparing a paper on this and
several other species of HopUtis.

152 GREAT BASIN NATURALIST Vol. 33, No. 3

superceded by the wasp Passaloecus cuspidatus, which prepared
seven cells. In this instance the Symmorphus male that developed
was eventually found dead in the cell, apparently unable to pene-
trate the resin partitions made by the Passaloecus.

No less than 14 of the 25 nests proved to have been parasitized
by the miltogrammine fly Amobia distorta Allen [det. R. J. Gagne].
In all, 36 flies were reared from these 14 nests, a maximum of 6
flies emerging from each of 2 nests. Within the nests, the maggots
moved freely from cell to cell, in some cases reducing the entire
interior to loose bits of mud, pieces of prey, and maggots or puparia.
A single bombyliid fly. Anthrax irroratus Say [det. L. V. Knutson],
was reared from one nest.

In all, only 12 adult Symmorphus cristatus were reared from
these nests, partly a result of the high incidence of parasitism. The
remaining nests were presumed to belong to this species on the
basis of the prey and, in two cases, on the basis of larvae preserved
when the nest was opened.

Krombein (1967) has studied this species in some detail and has
reviewed previous observations by Fye (1965). My own observa-
tions, so far as they go, are reasonably consistent with the results of
those authors.

Ancistrocerus adiabatus (Saussure)

I obtained six nests of this species, five at Moran and one at Pil-
grim Creek. These were in diverse situations: one on a tree root only
0.3 m high, two on dead trees about 1 m high, one on a cabin 1.3 m
high, one on live willow, and one on live sagebrush 0.3-0.5 m high.
Four of the nests were in 6 mm borings and had from 9 to 11 cells
measuring from 8 to 22 mm in length; each had one or two empty
vestibular cells measuring 10-15 mm in length. A fifth nest in a 6
mm boring had a single cell 15 mm long following several cells of a
bee and followed by a long vestibular cell, 52 mm long. A single
nest in a 4 mm boring had several cells (not counted) measuring
11-14 mm in length and followed by a vestibular cell 14 mm long.
Closing plugs were of mud and measured 2-3 mm in thickness;
partitions between cells were also of mud, but no more than 1 mm
thick; none of the nests had preliminary plugs at the extremity of
the boring. There were no intercalary cells or double partitions in
these nests (in contrast to those of Symmorphus) .

Provisioned cells were packed tightly with small caterpillars,
15-20 per cell. A sampling of these, 56 specimens from two separate
nests, all proved to be Coleotechniies sp. (Gelechiidae) [det. D. M.
Weisman]. Cocoons were whitish, very thin and delicate. The
only case of supercedure was the one mentioned above, in which a
single cell of Ancistrocerus followed several cells of a leafcutter bee
(not reared, but probably Hoplitis rohusta) .

One nest was found to contain maggots in two cells; this nest
later yielded one adult fly, Amobia floridensis (Townsend) [det.
R. J. Gagne]. Another nest contained two brown, parchment-like
cocoons; these later yielded cuckoo wasps, Chrysis caerulans

Sept. 1973 EVANS: Wyoming wasps 153

Fabricius. A third nest yielded a bombyliid fly, Anthrax irroratus
Say [det. L. V. Knutson ]. This fly emerged from a pupa that had
worked its way out of the nest and into the rearing container; a
second, dead pupa was inside the nest.

There are several published accounts of the nesting behavior of
this widely distributed species, mostly under the name A. tigris
(Saussure). My observations are consistent with Krombein's (1967)
much more detailed studies. Chrysis caerulans has been reared from
nests of this species on several occasions, and flies of the genera
Amobia and Anthrax have often been reared from this and other
species of the genus Ancistrocerus. Krombein found the prey to
consist of microlepidopterous larvae of five families.

Trypoxylon aldrichi Sandhouse

Fourteen trap nests were accepted by wasps of this species, 10 at
Pilgrim Creek and 4 at Moran. Five accepted 4 mm borings, 8 ac-
cepted 6 mm borings, and one nested in a 5 mm boring in a Sam-
hucus stem. Traps accepted were from 0.3 to 1.2 meters high and
were in a variety of situations: live alders, standing dead aspens,
standing live or dead pines, and prostrate logs. The number of cells
per 13 cm boring varied from 4 to 7, with two exceptions: one with

1 and one with 2 cells. The nest in the Sambucus stem had two
cells in a boring only 4 cm long; both cells were 8 mm long. In 4
mm borings, cells varied in length from 8 to 20 mm (mean 13.5),
while in 6 mm borings, cells varied in length from 7 to 20 mm
(mean 12.0). An empty vestibular cell, 8-13 mm in length, was
present in four nests. Empty intercalary cells, 2-5 mm in length,
were found in two nests. Closing plugs varied in thickness from

2 to 7 mm, and in several cases were recessed slightly from the
opening of the boring. Partitions between the cells were very thin;
all partitions and plugs were of mud.

From 5 to 16 spiders were provided per cell, the larger number
occurring in larger cells or when the spiders themselves were small
in size. Spiders of four families were utilized, all of them web
spinners, either orb webs, sheet webs, or the irregular webs of
theridiids. The species are chiefly those occurring on tree trunks,
in bushes, close to the ground, or under bark or stones. The follow-
ing is a list of those spiders saved for identification and the numbers
of each [determinations by H. W. Levi].

Theridiidae

Achaearanea sp. 2 9 9

Chrysso nordica (C. and I.) 1 9

Steatoda sp. 2 juveniles

Theridion aurantium Emerton 24 9 9

T. differens Emerton 1 $

LiNYPHIIDAE

Lepthyphantes spp. 7 9 9
Meioneta sp. 3 9 9
Pityohyphantes sp. 12 juveniles

154 GREAT BASIN NATURALIST Vol. 33, No. 3

Araneidae

Genera and spp. ? 24 juveniles

Tetragnathidae

Genus and sp. ? 1 juvenile

The egg of the wasp was found to be laid obliquely on the dor-
sum of the abdomen of one of the spiders deep in the cell. Twenty-
nine T. aldrichi were eventually reared from eight of these nests,
and although none were reared from the remaining six, I feel confi-
dent that all were nests of this species.

Only one instance of supercedurc was found. In this case a bee
{Hoplitis robusta Nylander) [det. G. C. Eickwort] had constructed
four cells at the extremity of the boring, and the Trypoxylon had
then constructed four more, a vestibular cell, and a closing plug.

I noted no maggots or puparia of parasitic flies when these nests
were first examined, but one nest yielded two flies, Amobia distorta
(Allen) [det. R. J. Gagne]. Two nests were, however, found to
contain a small larva of a clerid beetle. In both nests there was evi-
dence that the clerid had destroyed the egg or small larvae of the
wasp; probably some feeding on the spiders also occurred. The
clerids were not successfully reared to the adult stage.

Passaloecus cuspidatus Smith

Twenty trap nests were accepted by wasps of this species, 7 at
Moran and 13 at Pilgrim Creek. These trap nests were from 0.3
to 1.5 m high and were in diverse situations: on our cabin at Moran,
on pine and aspen stumps and logs, and attached to low branches of
live pines. Twelve were in 4 mm borings, 8 in 6 mm borings. Brood
cells in 4 mm borings averaged considerably longer than in 6 mm
borings (mean 11 mm, range 7-17 mm, as compared to mean 8.5
mm, range 5-12 mm in 6 mm borings). The number of cells per
13 cm boring ranged from 2 to 16, all of those having more than 10
cells being in 6 mm borings. Partitions were of resin and were very
thin, no more than 0.3 mm; in no case was there a partition at the
extreme end of the boring. Closing plugs were from 2 to 5 mm thick
and were also of clear resin except that several had dirt particles or
wood chips mixed with the resin. As reported by Krombein (1967)
from New York and Virginia, none of these nests had an empty
intercalary cell, but all had empty vestibular cells; the latter varied
in length from 12 to 58 mm, and one of them was divided in half
by a thin partition.

Provisioned cells were packed tightly with aphids, usually wing-
less forms and often immatures. I counted the aphids in 14 cells and
found the number to vary from 10 to 40 (mean 21), much of this
variation the result of differences in the size of the aphids. The egg
was found to be 1.3 mm long and to be laid on the venter of an aphid,
one end attached just behind the hind coxae and the other end ex-
tending free, obliquely forward and upward.

In most of the nests, all the aphids were dark in color; I preserved
80 aphids from several cells of two nests, and all proved to be Ptero-

Sept. 1973 EVANS: Wyoming wasps 155

comma bicolor (Oestlund) [det. L. M. Russell]. However, two nests
from Moran, collected on 9 and 13 August, were provisioned with
aphids of paler coloration; I preserved 9 of these and all proved to be
Macrosiphum euphorhiae (Thos.) [det. L. M. Russell]. Since there
were over 2000 aphids in the 120 cells of this wasp and I preserved
fewer than 100, I cannot be sure that other species of aphids were
not involved. However, I noted none that were superficially dif-
ferent from these two species.

There were three instances of supercedure, in each case the
Passaloecus usurping the nest of another wasp or bee. In one case
P. cuspidatus built 6 cells following one cell of Symmorphus crista-
tus; in the other two cases P. cuspidatus built 4-10 cells following 3
cells of a bee. The bees were not reared successfully but were prob-
ably Hoplitis robusta.

Passaloecus apj^eared to be free from attacks of miltogrammine
flies, possibly because of the nature of the partitions. One bombyliid
fly. Anthrax irroratus Say [det. L. V. Knutson], was reared from a
nest, but this was one of the nests containing three cells of a bee,
and it was not determined in what cell that parasite had developed.
An eight-celled nest yielded an ichneumon wasp, Poemenia ameri-
cana nebulosa Habeck and Townes [det. H. K. Townes], as well as
a chrysidid, Omalus aeneus Fabr. Both O. aeneus and P. a. americann
(Cresson) have been reared from this host by Krombein (1967).
I reared Omalus aeneus from one additional nest and also reared
O. cressoni (Aaron) from one nest and O. purpuratus (Provancher)
from two.

References

Allen, H. W. 1971. A monographic study of the genus Tiphia (Hymenop-

tera: Tiphiidae) of western North America. Trans. Amer. Ent. Soc. 97:

201-359.
Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole,

Wyoming. Bull. Mus. Comp. Zool., Harvard 140:451-511.
Fye, R. E. 1965. The biology of the Vespidae, Pompilidae, and Sphecidae

from trap nests in northwestern Ontario. Canad. Ent. 97:716-744.
Krombein, K. V. 1967. Trap-nesting Wasps and Bees: Life Histories, Nests,

and Associates. Smithsonian Press, Washington, D.C. 570 p.
Peckham, G. W., and E. G. Peckham. 1905. Wasps Social and Solitary.

Houghton Mifflin, Boston. 306 p.
Powell, J. A. 1967. Behavior of ground nesting wasps of the genus Nitelop-

terus, particularly A', californicus (Hymenoptera: Sphecidae). J. Kansas

Ent. Soc. 40:331-346.

ADDITIONAL RECORDS OF MUTILLID WASPS
FROM THE NEVADA TEST SITE'

Dorald M. Allred-

From 1960 to 1964, more than 800 mutillids, mostly females,
were collected in can pit traps during ecological studies by Brigham
Young University at the Nevada Test Site (Allred, Beck, and Jor-
gensen, 1963). These were submitted to Dr. William E. Ferguson,
San Jose State College, for study. I am indebted to him for making
the identification of those specimens whose records are contained
herein.

Inasmuch as the females of these insects have largely been un-
studied and most species of mutillids have been described from the
males, Ferguson spent part of the summer of 1964 at the test site
in order to correlate the sexes of some species through s])3cialized
collecting. Specimens taken by him were principally males collected
with light traps during August. In 1967 he published records of 31
species of mutillids of the test site, which included descriptions of
4 new to science, 2 new combinations, and 25 synonymies. Most of
his published records were of males.

The records included here deal with females as well as some
males which supplement those listed by Ferguson (1967). Inasmuch
as the females are wingless and likely do not move for great dis-
tances, their occurrence in pit traps may be closely correlated with
habitat and vegetation, whereas the males may fly for some distance
before capture in light traps, and such correlation for them may be
questionable. Notations of such habitat occurrence of these insects
are included under the species listings.

The Nevada Test Site is situated approximately 70 miles north-
west of Las Vegas in the southeastern part of Nye County, Nevada.
It is typified by the Upper and Lower Sonoran life zones, repre-
sentative of the cool and hot deserts of North America. Th-^ northern
part of the test site occurs in the Great Bns'n, whereas the southern
part is in the Mojave Desert.

In their descri{)tion of the biotic communities of the Nevada Test
Site, Allred, Beck, and .lorgensen (1963) delineated six major plant
communities and assigned other less distinct areas to a "mixed"
designation. The Larrca divnricata-Franscrin dumosa community
was designated as belonging to the Mojave Desert. Besides the pre-
dominant species for which the community was named, plants of 37
other species are common therein. The Gray la spinosa-Lycium an-
dersonii and Coleogyne ramosissima plant communities, also of the
Mojave Desert, each have 26 known species of common plants in
their composition. Atriplex confcrtijolia and Kochia americana are
the predominant plants in a conmiunity wherein only three other

iBYU-AEC Publication No. COO-1,355-21 .
*Brigham Young University. Prove, Utah 84602.

156

Sept. 1973 ALLRED: MUTILLID WASPS 157

species are considered as common associates. The Salsola kali com-
munity is typical of disturbed areas and has as many as nine other
species of common plant associates. The Pinyon- Juniper {Pinus
monophylla and Juniperus osteosperma) community is character-
istic of the mesas and higher foothill areas, particularly of the Great
Basin region. Plants of fifteen other species are commonly associated
with these Desert Woodland trees.

Additional Records

Acanthophotopsis falciformis falciformis Schuster. Ferguson
(1967) listed records for six males collected in a light trap in August,
and one in a pit trap in June. Additional record: Area 5 — Larrea
divaricata-Franseria dumosa, July, pit trap. Comment: Ferguson's
(1967) records of this species were taken in the Mixed and Larrea-
Franseria communities.

Acrophotopsis eurygnathus Schuster. Ferguson (1967) reported
19 males from the test site. Additional records: Area 5 — Lycium
pallidum, 4cf July; 3cf 2? June; Larrea divaricata-Franseria du-
most, 1 d" 2 ? July, pit trap. Comments: Ferguson (1967) indicat-
ed the Larrea-Franseria community as the source of his specimens.
His records also include the Mixed vegetative type. The Lycium pal-
lidum is an additional habitat type for this species.

Chyphotes melaniceps Blake. This species has not boen previous-
ly reported from the Nevada Test Site. Records (all males) : Area
1 — Salsola kali, 1 in June, pit trap; Grayia spinosa-Lycium ander-
sonii, 1 in May, 1 in June, 2 in Aug., 2 in Sep., 2 in Oct., pit trap.
Area 4 — Grayia spinosa-Lycium andersonii, 1 in Aug., aerial net.
Area 5- — Larrea divaricata-Franseria dumosa, 6 in July, 1 in Sep.,
1 in Oct., pit trap; Lycium pallidum, 2 in July, 7 in Aug., pit trap;
12 in July, incandescent-light trap. Area 6 — Atriplex confertifolia-
Kochia americana, 1 in Aug., pit trap; Yucca brevifolia-Coleogyne
ramosissima, 9 in Aug., aerial net. Area C — Mixed vegetation, 1 in
Aug., 2 in Sep., pit trap and aerial net, respectively. Area J — Mixed
vegetation, 1 in Sep., pit trap. Comments: A variety of ecological
types are occupied by this species. On the basis of seasonal occur-
rence, the Grayia-Lycium is the most commonly represented. How-
ever, greatest numbers were found in the Lycium pallidum areas.

Chyphotes petiolatus Fox. This species was not reported from the
test site by Ferguson (1967). Records: (all males): Area 1 —
Grayia spinosa-Lycium andersonii, 5 in May, 6 in June, 3 in July,
1 in Aug., 2 in Sep., pit trap; 1 in Mar., aerial net; Salsola kali, 1 in
May, pit trap. Area 5 — Larrea divaricata-Franseria dumosa, 1 in
Apr., 5 in June, 2 in July, 2 in Aug., pit trap; Lycium pallidum, 1 in
Apr., 5 in May, 4 in June, 4 in July, 4 in Aug., 2 in Sep., pit trap.
Area 10 — Coleogyne ramosissima, 2 in June, pit trap. Area CB —
Mixed vegetation, 1 in Sep., pit trap. Comments: This species oc-
cupies a variety of habitats. Seasonally and in abundance it is most
common in the Lycium pallidum and Grayia-Lycium communities.

158 GREAT BASIN NATURALIST Vol. 33, No. 3

Dasymutilla gloriosa (Saussure). Ferguson (1967) reported
three females taken from the test site. Additional records: Area 1 —
Salsola kali, 9 , Jmie, pit trap. Area 4 — Grayia spinosa-Lycium an-
dersonii, 9 , Sep., aerial net. Area 400 — Mixed vegetation, 9 , Tuly,
by hand. Area C — Mixed vegetation, 9 , July, pit trap.

Dasymutilla klugii (Gray). Ferguson (1967) did not report this
species from the test site. Record: Area 5 — Larrea divaricata-Fran-
seria dumosa, <S , July, aerial net.

Dasymutilla satanas Mickel. Ferguson (1967) reported 24 fe-
males from the test site. Additional records: Area 4 — Grayia
spinosa-Lycium andersonii, 2 9 July, 9 Aug., 2 9 Sep., pit trap;
9 Aug., aerial net. Area 5 — Larrea divaricata-Franseria dumosa, cf
3 9 July; Lycium pallidum, 4 9 July, 3 9 Aug., 9 Sep., pit trap.
Area 10 — -Coleogyne ramosissima, 9 Aug.; Salsola kali, 9 Aug., pit
trap. Area C — Mixed vegetation, 9 June, pit trap. Comments:
Ferguson's (1967) records were related to Mixed and Larrea-
Franseria communities. Additional records show a variety of habi-
tats for this species. Seasonally and in numbers it is apparently most
common in the Lycium pallidum and Grayia-Lycium communities.

Odontophotopsis mamata Schuster. Ferguson (1967) reported
100 specimens from the test site. Additional records (all males):
Area 1 — Grayia spinosa-Lycium andersonii, 1 in July, 2 in Aug.,
pit trap. Area 5 — Lycium pallidum, 1 in June, 1 in July, 21 in Aug.,
1 in Sep., pit trap; 4 in July, incandescent-light trap; Larrea divari-
cata-Franseria dumosa, 3 in July, 3 in Aug., pit trap. Area 6 —
Yucca brevifolia-Coleogyne ramosissima, 12 in Aug., aerial net.
Area C — Mixed vegetation, 1 in Aug., aerial net. Area J — Mixed
vegetation, 1 in Aug., pit trap. Comments: This species occurs in a
variety of habitats but is probably most common in the Lycium
pallidum community. Ferguson's (1967) records were from only
the Mixed community type.

Odontophotopsis setifera Schuster. Ferguson (1967) indicated
one specimen from the test site. Additional record: Area 5 —
Lycium pallidum, cf July, incandescent-light trap.

Sphaeropthalma angulifera Schuster. Ferguson (1967) reported
seven specimens from the test site. Additional records: Area 1- —
Grayia spinosa-Lycium andersonii, 2 cT June, cT July, <S Sept.;
Salsola kali, <S June, pit trap. Area 5 — Larrea divaricata-Franseria
dumosa, 2 9 May, 3 9 June, 3 9 July, pit trap. Area J — Mixed
vegetation, 9 June, pit trap. Comments: Ferguson's (1967) records
were taken in Mixed vegetation and Coleogyhe communities. He
indicated that this species does not occur at lower elevations. How-
ever, additional records show it to occur in several habitats, |)rimarily
in the Larrea-Franseria and Grayia-Lycium comnuinities.

Sphaeropthalma blakeii (Fox). Ferguson (1967) listed three
specimens from the test site. Additional records (both males): 1
in June, 1 in Oct., pit trap.

Sphaeropthalma sonora Schuster. Ferguson (1967) recorded 90
specimens from the test site. Additional records (all males): Area

Sept. 1973 ALLRED: MUTILLID WASPS 159

1 — Grayia spinosa-Lycium andersonii, 1 in June, 1 in July, 1 in
Aug., pit trap. Area 5 — Larrea divaricata-Franseria dumosa, 12 in
July, 4 in Aug., pit trap; 6 in Aug., aerial net; Lycium pallidum,
2 in July, 6 in Aug., pit trap; 15 in July, incandescent-light trap;
18 in June, 2 in July, black-light trap. Area 6 — Yucca brevifolia-
Coleogyne ramosissima, 1 in Aug., aerial net. Area 10 — Grayia
spinosa-Lycium andersonii, 3 in July, pit trap. Area CB — Mixed
vegetation, 1 in Sep., aerial net. Comments: Ferguson (1967) indi-
cated that this species is common at lower elevations below the
Pinyon-Juniper and Coleogyne communities. Additional records
show it to be most common seasonally and in numbers in the Lycium
pallidum and Larrea-Franseria communities.

Sphaeropthalma unicolor (Cresson). Ferguson (1967) listed 76
females and 53 males from the test site. Additional records: Area
1 — Grayia spinosa-Lycium andersonii, 10 5 Apr., 23 ? May, 12 9
June, 11 9 July, cf 5 9 Aug., cT 4 9 Sep., 3 9 Oct., 9 Nov., pit
trap; 9 May, aerial net; Salsola kali, 2 9 June, pit trap. Area 4 —
Grayia spinosa-Lycium andersonii, cf Aug., aerial net. Area 5 —
Larrea divaricata-Franseria dumosa, 9 June, 9 Aug., pit trap. Area
10 — Coleogyne ramosissima, <S July, pit trap. Area J — Mixed vege-
tation, 2 9 Apr., 2 9 May, cf 3 9 Aug., pit trap. Comments: This
species was taken by Ferguson (1967) in essentially all communities
from the valley floors to the tops of the mesas. Additional records
show it to be most common seasonally and in numbers in the Grayia-
Lycium community.

Summary ^

Ferguson (1967) listed 31 species of mutillids from the Nevada
Test Site. Three additional records are herein recorded — Chyphotes
melaniceps, C. petiolatus, and Dasymutilla klugii.

On the basis of numbers collected, the following species are con-
sidered as rare at the test site (fewer than 10 specimens collected,
as indicated by the numbers in parentheses) :

Dasymutilla klugii (1) Dasymutilla gloriosa (4)

Dasymutilla paenulata (1) Dilophotopsis concolor (4)

Sphaeropthalma helicaon (1) Sphaeropthalma blakeii (5)

Odontophotopsis setifera (2) Sphaeropthalma ferruginea (6)

Sphaeropthalma macswaini (3) Sphaeropthalma pallida (7)

Sphaeropthalma parapenalis (3) Acanthophotopsis falciformis (8)

Considered as relatively abundant at the test site (over 100 speci-
mens collected, as indicated by the numbers in parentheses) are the
following species:

Sphaeropthalma unicolor (216) Sphaeropthalma sonora (164)

Mutillids of the species Chyphotes melaniceps were the most
widespread ecologically, whereas those belonging to Sphaeropthalma
unicolor were the most widespread seasonally (Table 1). The great-
est varieties of species were found in the Mixed, Larrea-Franseria ,
and Grayia-Lycium communities, in that order, and the fewest in

160

GREAT BASIN NATURALIST

Vol. 33, No. 3

the Atriplex-Kochia (Table 2). The greatest number of species was
taken during July, and the fewest in March and November; none
were found from December through February (Table 3). Greatest

Table 1. Summary of ecological and seasonal distribution of some mutillids
at the Nevada Test Site.

Species

Chyphotes melaniceps
Chyphotes petiolatus
Dasymutilla satanas
Odontophotopsis mamata
Sphaeropthalma sonora
Sphaeropthalma unicolor
Sphaeropthalma angulifera
Dasymutilla gloriosa
Acrophotopsis eurygnathus
Acanthophotopsis falciformis
Dasymutilla klugii
Odontophotopsis setifera
Sphaeropthalma blakeii

No. communities

No.

months

in which

found

collected

Table 2. Number of species and relative abundance of individual mutillids
in seven vegetative types at the Nevada Test Site.

Vegetative type

Mixed

Larrea divaricala-Franseria dumosa

Grayia spinosa-Lycium andersonii

Lycium pallidum

Coleogyne ramosissima

Salsola kali

A triplex confer tifolia-Kochia americana

Individual

Number of

abundance

species

factor*

1.6

11.8

6.5

19.3

2.4

1.0

1.4

•Compared to 1 as representative of the fewest tollected; adjusted to luuiiber of trap nights and
collection attempts.

Table 3. Number of species and abundance of individuals collected during
months of mutillid activity at the Nevada Test Site.

Month

Number
species

Abundance'

Actual

Ratio

numbers

factor^

1.0

'17

5.7

7.7

6.9

196

10.9

145

10.4

4.4

3.5

1.0

March

April

May

June

July

August

September

October

November

2
3
6

14
18
14
12
6
2

'Based on equal collecting efforts for each month over a three-year period, 1959-1962.
The number of individuals divided by the number of species. The higher the factor, the
greater the relative abundance.

Sept. 1973

ALLRED: MUTILLID WASPS

161

numbers of individuals were taken in July, although in considera-
tion of the number of species represented, populations were not sig-
nificantly different between July and August (Table 3). On this
latter basis, a decline in the expected increase in relative populations
in relationship to numbers of species was noted for June (Figure 1).

210

180

150

120

T 1 1 r

Ap My Jn Jy Ap S ON

Fig. 1. Seasonal trend of numbers of individual mutillids, numbers of
species represented, and the ratio factor (number of individuals divided by num-
ber of species represented). The number of species and ratio factor were multi-
plied by 10 to correlate more closely in degree for comparison with population
number trends.

152 GREAT BASIN NATURALIST Vol. 33, No. 3

References

Ferguson W E 1967. Male Sphaeropthalmine wasps of the Nevada Test
Site Brigham Young Univ. Sci. Bull., Biol. Ser. 8 (4): 1-26.

AI.LRED, D. M., D E. Beck, and C. D. Jorgensen. 1963^ Biotic communities
of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser. 2(2):
1-52.

THE EFFECTS OF SOIL TEXTURE ON SPECIES

DIVERSITY IN AN ARID GRASSLAND OF THE

EASTERN GREAT BASIN

John W. Wyckoffi

Abstract. — A primary factor limiting plant species diversity in desert
grasslands seems to be soil texture. Loamy soils consistently support more species
than do adjacent sands. This is probably related to the increased micro-environ-
mental differences associated with heavy textured soils. Such soils would be ex-
pected to have more micro-topographic and soil moisture variations than sandy
soils.

The grassy communities of the Great Salt Lake State Park, situ-
ated on the northern end of Antelope Island of the Great Salt Lake,
Utah, were intensively studied during the period 14 June to 3 Sep-
tember 1971. This study was part of a larger project funded by the
Student Originated Studies Program of the National Science Founda-
tion (Carter et al., 1971). The major aim of my study was to evalu-
ate the relationships of vascular plant diversity to soil texture and
micro-environmental variation within the arid grasslands of the park.

The fact that soil texture is an important ecological factor control-
ling plant populations is well known (Black, 1968). Soil particles
regulate the flow of moisture, retention of nutrients, and movement
of air within a given soil (Russell, 1957). Locally, extensive work
has been done to document characteristics of soils of the Great Salt
Lake- Wasatch Front region (Erickson, 1968; Jennings, 1946; Thome,
1968). Other research in this general geographic area has shown
definite correlations between composition of grassland communities
and edaphic conditions (Kleiner, 1966 and 1968; Ludwig, 1965 and
1969).

The area under consideration is in the northeastern part of the
Great Basin, approximately 30 mi. northwest of Salt Lake City,
Utah. The vegetation of the park is dominated by grassland species
typical of this region. The communities studied have been grazed
by livestock until the recent past and support both native and in-
troduced plant species (Table 1). Soils in the park ranged from sand,
at locations near the lake, to sandy loam in areas farther from the
shoreline. Since most of the precipitation in this part of the Great
Basin is restricted to the winter months, most water available to
vegetation is present for only a short time in the spring. The only
other sources of water during the year are small, infrequent storms
which occur during the drier summer months. Total precipitation
annually averages approximately 10 in. Therefore water is one of
the primary limiting factors for plants in this area, and competition
for water may have important effects on the abundance and distribu-
tion of most plant species.

'Department of Biology, University of Utah, Salt Lake City, Utah.

163

164

GREAT BASIN NATURALIST

Vol. 33, No. 3

Table 1. Species encountered in the grasslands of Great Salt Lake State
Park showing average quadrat frequency expressed as a percentage of the total
within the 15 sections of each 16.2 hectare plot.

Species

Plot 1

Plot 2

Plot 3

Agropyron spicatum

0.3

1.0

41.0

Arisdda longiseta

13.8

50.5

29.3

Artemisia tridentata

5.0

1.3

0.3

Astragalus utahensis

0.3

Bromus tectorum

85.0

99.3

100.0

Castilleja angustifolia

0.3

Chrysothamnus nauseosus

1.5

0.3

Chrysothamnus viscidiflorus

7.3

0.3

Cleome lutea

0.5

1.0

0.3

Cryptantha parva

0.3

Cymopteris longipes

0.3

10.5

Distichlis striata

14.0

0.3

Erigeron englemani

0.3

3.8

Erodium cicutarium

38.5

91.5

20.8

Festuca myoros

0.3

2.5

0.3

Guterrezia sarothrae

2.8

0.3

Hordeum jubatum

2.8

0.3

Lactuca scariola

0.3

Lomatium grayii

0.3

5.5

Oryzopsis hymenoides

3.0

1.0

0.3

Phlox longifolia

8.0

3.0

6.8

Poa sandbergii

15.3

30.3

54.3

Salsola kali

0.5

0.3

Sisymbrium altissimum

0.3

Sitanion jubatum

0.5

0.3

Sphaeralcea coccinea

21.3

2.0

Sporobolus airoides

6.8

0.3

Sporobolus cryptandrus

8.8

18.3

4.8

Stipa comata

3.5

5.5

0.3

Zigadenus paniculata

0.8

0.3

Percent exotics

51.3

61.4

42.7

Percent native

48.7

38.6

57.3

Average Plant Diversity/.007 h

3.5

4.2

3.9

Methods

Three areas in the park were selected for vegetational studies.
The first was comprised primarily of introduced species, the second
was intermediate with approximately equal numbers of introduced
and native species, and the third was dominated by native species.
Each area encompassed 16.2 hectares (40 acres) and was subsampled
by fifteen 0.007 hectare (1/60 acre) circular sections. Within each
of these sections, the vegetation was sampled with 25 quadrats.
Each quadrat had an area of 0.25 m- and was subdivided into 4
units (quartiles) of equal size. At each sampling point the presence
or absence of individual plant species was noted for each quartile.
From these quadrat analyses a frequency of occurrence was estab-
lished for each plant species present.

Soil and surface environmental analyses were made within each
of the 0.007 hectare sections. Soil data taken included particle size,
pH, free carbonates, and salinity. Soil particle size was determined
using Bouyoucos's (1962) method, pH was obtained with a dual

Sept. 1973 WYCKOFF: GRASSLAND PLANTS 165

glass electrode meter (Leeds and Northrup) on a soil paste, and
presence of free carbonates was determined by an effervescence test
using a 10 percent HCL solution. Salinity was evaluated by con-
ductivity of soil paste using a Wheatstone Bridge and soil cup elec-
trode. Surface environmental variables such as surface rock, living
cover, and bare soil were determined quantitatively by an ocular
estimation technique (Warner and Harper, 1972). Average values
for the several variables are reported for the three major study areas
in Table 2.

Using MacArthur and Wilson's (1967) diversity index

Diversity index = 1

1 Pa = frequency of species a

2 frequency of species a-z

it was possible to calculate a workable diversity index for each of the
study sections. The minimum diversity for this index is 1.0 and the
magnitude of the index increases as the number of species and the
uniformity of their frequency values increase.

Results and Discussion

The influence of various environmental parameters on plant
species diversity was evaluated by stepwise multiple regression
(Dixon, 1969). Results of the computer analysis give an indication
of the relative influence of each environmental variable on vegeta-
tional diversity (Table 3) . It is apparent that of those environmental
variables studied, soil texture has the most influence upon plant
diversity. The relationship between soil texture and plant diversity
is graphically shown in Fig. 1. This is probably due to a greater
degree of microenvironmental diversity on more consolidated soils.
Increased diversity might be related, for example, to more small
scale runoff and accumulation of water on the heavier textured soils.
Additionally, in an arid environment one might expect "effective"

Table 2. Avei-age values for environmental variables/16.2 hectare plot.

Plot 1 Plot 2 Plot 3

Percent sand
Percent fines

Conductivity*
Percent surface rock
Percent living cover
Percent surface litter
Percent bare soil
Carbonates
(No. of 0.007 hectare
sections found in)

'Conductivity in ohms is measurement of salinity
fdna = data not available

83.4

62.4

63.4

16.6

37.6

36.6

8.34

7.53

7.34

2235

1391

dna+

1.1

4.9

31.3

73.8

64.8

52.4

15.1

16.8

9.0

15.6

17.0

14.4

166

GREAT BASIN NATURALIST

Vol. 33, No. 3

Erecipitation (i.e., precipitation which can be utihzed by plants) to
e lower on heavier textured soils due to less deep infiltration and
correspondingly larger losses via evaporation as opposed to trans-

r^: .23

100

40
% FINES

Fig. 1. Graphical portrayal of the relationship between plant diversity and
percentage of soil particles classified as fines (percent silt + percent clay).
Diversity indices are calculated using MacArthur and Wilson's (1967) index as
described in text.

Table 3. Results of stepwise multiple regression analysis of environmental
variables affecting plant species diversity. Variables are shown in the order in
which they entered the analysis.

Influence

Increase

R-square

Variable

Diversity

R-value

value

R-square

Soil fines

.4796

.2301

Surface litter

.5383

.2898

.0597

Living cover

.5743

.3298

.0400

Free carbonates

—

.6231

.3882

.0584

Soil sand

.6440

.4148

.0266

Bare soil

.6532

.4267

.0119

.6552

.4292

.0025

Salinity

—

.6580

.4330

.0037

Sept. 1973 WYCKOFF: GRASSLAND PLANTS 167

piration (Black, 1968). The combined effects of heterogeneous
distribution of water and greater aridity of heavy textured soils per
unit precipitation would result in more habitat diversity (niches)
and a more sparse vegetation with the likelihood of reduced compe-
tition between individuals. Both conditions could lead to enhanced
diversity. Data supportive of these hypotheses can also be extracted
from Ludwig's (1969) study of several grassland communities in
northern Utah (Table 4).

Table 4. Community data for grasslands in northern Utah (Ludwig, 1969).

Community type

Average
no. spp./.025 acre

Average
no. spp./quad

Average
percent fines

Agropyron spicatum
Stipa comata
Sporobolus cryptandrus
Oryzopsis hymenoides

26.28

13.74

10.29

9.67

5.80
4.44
4.61
3.12

69
36
30

Conclusions

In Great Salt Lake State Park, sandy sites consistently have lower
diversity than loamy sites. These data suggest that much of the vege-
tational diversity within the community is dependent upon edaphic
factors; this finding is probably related to the uniform environment
of sandy areas. On such sites, the best adapted species could invade
and completely dominate, whereas in areas which foster micro-
environmental diversity (i.e., heavier textured soils, which have
more small scale runoff and therefore small adjacent areas with
very different amounts of "effective" precipitation) any one species
would find it difficult to dominate the entire area.

Literature Cited

Black, C. A. 1968. Soil plant relationships (2nd ed.). New York: John Wiley
& Sons.

BouYOUCOs, C. K. 1962. Hydrometer method improved for making particle size
analyses of soils. J. Agron. 54:464-65.

Carter, C. K., J. C. Bott, J. W. Carter, A. A. Coburn, K. K. Greene, C. R.
Hayes, T. D. Hoagland, J. B. Olsen, R. H. Schulze, S. T. Shipman, E. R.
O'SuLLivAN, J. Vander Meide, AND J. W. Wyckoff. 1971. Some ecologi-
cal considerations of the Farmington Bay Estuary and adjacent Great Salt
Lake State Park. Salt Lake City: University of Utah Press.

Dixon, W. J., ed. 1969. Biomedical Computer Programs. University of Calif.
Publications in Automatic Computation, No. 2. Berkeley: University of
California Press.

Erickson, a. J., AND L. Wilson. 1968. Soil Survey of Davis-Weber Area,
Utah, U.S. Dept. of Agriculture, Soil Conservation Service. Washington, D.C.:
U.S. Government Printing Office.

Jennings, D. S., et al. 1946. Soil survey of the Salt Lake area, Utah. Wash-
ington, D.C.: U.S. Government Printing Office.

Kleiner, E. F. 1966. A study of the vegetational communities of Red Butte
Canyon, Salt Lake County, Utah. M.S. thesis. University of Utah.

. 1968. Comparative study of the grasslands of Canyonlands National

Park. Ph.D. dissertation. University of Utah.

168 GREAT BASIN NATURALIST Vol. 33, No. 3

LuDWiG, J. 1965. Relationships of grasslands to edaphic gradients of the Foot-
hill Zone, Salt Lake County, Utah. M.S. thesis. University of Utah.

. 1969. Environmental interpretation of foothill grassland communities

of northern Utah. Ph.D. dissertation. University of Utah.

MacArthur, R. H., and O. E. Wilson. 1967. The theory of island biogeogra-
phy. Princeton: Princeton University Press.

Thorne, J. P. 1968. Chemical and physical properties of the soils of the
Wasatch Front counties — Weber, Davis, Salt Lake, and Utah. Utah Agric.
Exp. Sta. Logan, Utah Res. Ser. 44.

Warner, J. H., and K. T. Harper. 1972. Understory characteristics related to
site quality for aspen in Utah. Brigham Young Univ. Sci. Bull. Biol. Ser.
16(2):l-20.

NEW SYNONYMY IN AMERICAN BARK BEETLES
(SCOLYTIDAE: COLEOPTERA). PART IIP

Stephen L. Wood-

Abstract. — New synonymy affecting North and Central American Scoly-
tidae is proposed as follows: Araptus Eichhoff {=Neodryocoetes Eggers, Tham-
nophthorus Schedl, Sphenoceros Schedl), Coccotrypes Eichhoff { = Poecilips Schau-
fuss), Gnathotrichus Eichhoff (=Ancyloderes Blackman, Prognathotrichus
Bright), Gnathotrupes Schedl { = Gnathotry partus Wood), Araptus hymenaeae
(Eggers), n. comb., (—Neodryocoetes insularis Eggers, N eodryocoetes caribaeus
Blackman, Neodryocoetes guianae Blackman, Neodryocoetes hoodi Blackman),
Araptus laevigatus (Eggers), n. comb. {=Neopityophthorus insularis Eggers, Neo-
dryocoetes guadeloupensis Schedl), Araptus macer Bright, n. comb. {=N eodryo-
coetes tuberculatus Bright), Araptus politus (Blandford), n. comb. {=N eodryo-
coetes hubbardi Blackman), Araptus tabogae (Blackman), n. comb. {=Neodryo-
coetes vinealis Bright), Cnesinus elegantis Wood (^Cnesinus zapotecus Bright),
Coccotrypes advena Blandford { = Thamnurgides persicae Hopkins), Corthylo-
curus barbatus (Blandford) {=Corthylocurus cincinnatus Bright), Corthylocurus
mexicanus (Schedl) (=Corthylus anomalus Bright), Corthylus mexicanus Schedl
(=Corthylus glabinus Bright), Crypturgus pusillus (Gyllenhal) ( = Crypturgus
atomus LeConte), Dendrocranulus carbonarius (Ferrari) {=Xylocleptes flori-
densis Hopkins, Xylocleptes anonae Hopkins), Dendrocranulus cucurbitae (Le-
Conte) (=Xylocleptes californicus Hopkins, Xylocleptes venturina Hopkins,
Xylocleptes punctatus Hopkins), Dendrocranulus macilentus (Blandford) ( = Den-
drocranulus grossopunctatus Schedl), Hylastes salebrosus Eichhoff {=Hylastes
scobinosus Eichhoff), Hypothenemus seriatus (Eichhoff) {=Stephanoderes pul-
verulentus Eichhoff), Ips spinifer (Eichhoff) {=Orthotomicus sabinianae Hop-
ping), Leperisinus aculeatus (Say) {=Hylesinus imperialis Eichhoff), Micracis
lignator Blackman {^Micracis truncatus Wood), Micracis swainei Blackman
{= Micracis photophilus Wood), Micracis suturalis LeConte {=^Micracis meri-
dianus Blackman), Monarthrum querneus Wood (=Monarthrum bifidus Bright),
Orthotomicus caelatus Eichhoff ( = Xyleborus vicinus LeConte, Xyleborus puncti-
pennis LeConte), Coccotrypes indicus (Eggers) { = Xyleborus conspeciens Schedl,
Coccotrypes insularis Eggers, Poecilips eggersi Schedl), Phloeoborus asper Erich-
son { = Phloeoborus ovatus Chapuis, Phloeoborus rugatus Blandford), Phloeoborus
rudis Erichson ( = Phloeoborus elongatus Chapuis, Phloeoborus rugipennis Eg-
gers), Phloeotribus armatus Blandford {=Phloeotribus mixtecus Bright), Phloeo-
tribus demessus Blandford {= Phloeotribus tuberculatus Eggers), Phloeotribus
pilula (Erichson) {= Phloeotribus obliquus Chapuis, Phloeotribus manni Black-
man), Phloeotribus setulosus Eichhoff {= Phloeotribus asperatus Blandford),
Pityoborus frontalis Wood { = Pityoborus severus Bright), Pityoborus secundus
Blackman (= Pityoborus tertius Blackman, Pityoborus immitus Bright, Pityoborus
intonsus Wood, Pityoborus ramosus Bright), Pycnarthrum hispidum (Ferrari)
{=Nemobius larnbottei Chapuis, Pycnarthrum gracile Eichhoff, Pycnarthrum
quadraticolle Eichhoff, Pycnarthrum transversum Blandford, Pycnarthrum reimo-
seri Schedl, Pycnarthrum reticulatus Schedl), Pseudopityophthorus declivis Wood
{ = Pseudopityophthorus truncatus Bright, Pseudopityophthorus curtus Bright),
Pseudopityophthorus hondurensis Wood {= Pseudopityophthorus montanus Bright),
Pseudopityophthorus micans Wood {^Pseudopityophthorus squamosus Bright),
Pseudopityophthorus opacicollis Blackman {= Pseudopityophthorus aesculinus
Bright), Pseudopityophthorus pruinosus (Eichhoff) {= Pseudopityophthorus pul-
vereus Blackman, Pseudopityophthorus tropicalis Wood, Pseudopityophthorus con-
vexus Bright), Pseudopityophthorus singularis Wood { = Pseudopityophthorus acu-
minatus Bright), Pseudopityophthorus tennis Wood {= Pseudopityophthorus hir-
sutus Bright), Stenocleptus sulcatus (Bruck) (—Stenocleptus ceanothi Blackman,

'The researcli on which this paper was based was sponsored by the National Science Foundation.
^Department of Zoology, Brighani Young University, Provo, Utah 84602. Scolytidae Contribution
No. 50.

169

1 70 GREAT BASIN NATURALIST Vol. 33, No. 3

Stenocleptus rhois Blackman), Thysanoes texanus Blackman { = Thysanoes vachel-
liae Blackman, Thysanoes ratamae Blackman), Trypophloeus striatulus (Manner-
heim) {=^Trypophloeus nitidus Swaine), Xyleborus capucinus Eichhoff { = Xyle-
borus rufithorax Eichhoff), Xyleborus pubescens Zimmermann {= Xyleborus
pini Say of Eichhoff), Xylosandrus curtulus Eichhoff {=^ Xyleborus biseriatus
Schedl). Carphotoreus, n. gen., is described for Chaetophloeus alni Bright. Cocco-
trypes sannio (Schaufuss) is treated as a taxon distinct from Coccotrypes advena
Blandford; and Gnathotrichus consentaneus Blandford is a valid species distinct
from Gnathotrichus sulcatus LeConte. Gnathotrichus bituberculatus Blandford
and all described South American species of Gnathotrichus are transferred to
Gnathotrupes.

In the process of writing a taxonomic monograph of the Scoly-
tidae of North and Central America, the unpubhshed synonymy
summarized in the above abstract and treated on the following pages
was encountered. In order to stabilize nomenclature and fix many
established names, several lectotypes are designated and the resulting
new synonymies are reported. Wherever possible the traditional
usage has been preserved through lectotype designation. In several
instances, however, established names are junior synonyms and
cannot be conserved. In my opinion none of the junior synonyms
affected by this action are sufficiently important to warrant special
action to conserve them. The basis on which the synonymy is
established is cited for each name treated.

Araptus Eichhoff

Araptus Eichhoff, 1878, Mem. Soc. Roy. Sci. Eiege (2)8:305 (Type-species:

Araptus rufopalliatus Eichhoff, monobasic)
Neodryocoetes Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat.,

Paris 1(1):9 (Type-species: Neodryocoetes hymenaeae Eggers. monobasic).

New synonymy
Thamnophthorus Schedl, 1938, Archiv Naturgesch. 7:174 (Type-species: Tham-

nophthorus volastos Schedl, subsequent designation by Blackman, 1942, Proc.

U.S. Nat. Mus. 92:178). New synonymy
Sphenoceros Schedl, 1939, Mitt. Miinchner Ent. Ges. 29:565 (Type- species: Spheno-

ceros Umax Schedl, monobasic). New synonymy

The unique female holotype of Araptus rufopalliatus Eichhoff
in the Institut Royal des Sciences Naturelles de Belgique, Brussels,
lacks antennae and legs. The frons of this specimen has unusual
sculpture and the epistoma is deeply emarginate; consequently, the
phylogenetic position of this genus remained a mystery until a series
of specimens identical to the type was collected at La Carbonera
experimental forest, 50 km NW Merida, Merida, Venezuela, 16-IX-
1969, at 2500 m. No. 21b, from Ncctandra sp., by myself. The
antennae, legs, other anatomical features, and the habits clearly
indicate that this species is congeneric with Neodryocoetes hyme-
naeae Eggers. Neodryocoetes Eggers, therefore, must be placed in
synonymy as indicated above.

In a review of the Central American species of Araptus it was
noted that the antennal and other features used to characterize
Thamnophtliorus Schedl and Sphenoceros Schedl are not consistent
and intergrade completely with those of various other species groups
of Araptus. The species formerly assigned to Sphenoceros are phloe-

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 171

ophagous, not xylomycetophagous as assumed by Schedl. The names
Thamnophthorus Schedl and Sphenoceros Schedl, consequently, are
placed in synonymy under Araptus.

Carphotoreus, n. gen.

This genus superficially resembles certain Chaetophloeus species,
but it is distinguished by the 6-segmented antennal funicle, by the
more widely distributed crenulations on the basal margins of the
elytra, by the absence of submarginal crenulations on the elytral
bases, by the visible scutellum, by the different arrangement and
type of pronotal asperities, and by other characters.

Description. — Length 1.5-1.6 mm, 2.14 times as long as wide.

Frons sexually dimorphic, moderately concave in male, flattened
or convex in female; eye elongate-oval, entire; scape moderately
long, funicle 6-segmented, club flattened, almost symmetrical, three
nonseptate sutures indicated. Pronotum wider than long, antero-
lateral areas with a few asperities. Scutellum small, visible. Elytra
suturally emarginate at base, a row of marginal crenulations at base,
submarginal crenulations entirely absent; striate; declivity simple.
Anterior coxae moderately separated. Third tarsal segments slender.

Type-species. — Chaetophloeus alni Bright.

This genus belongs to the Hylesinini and is tentatively placed
near Alniphagus Swaine, although the relationship is not close.

Coccotrypes Eichhoff ^

Coccotrypes Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:308 (Type species:
Bostrichus dactyliperda Fabricius, subsequent designation by Hopkins, 1914,
Proc. U.S. Nat. Mus. 48:118)

Poecilips Schaufuss, 1897, Berliner Ent. Zeitschr. 42:110 (Type-species: Poecilips
sannio Schaufuss, monobasic). New synonymy

This large diverse genus has been divided into Coccotrypes Eich-
hoff and Poecilips Schaufuss on the basis of pronotal structure and
armature, and on habits. In a review of species assigned to these
generic names, however, there seems to be considerable confusion.
In Coccotrypes the pronotum should be strongly convex, with the
anterior slope asperate, and they are spermophagous. In Poecilips
the pronotum should be more weakly convex, with the anterior slope
unarmed, and most species are supposed to be phloeophagous. Ac-
tually, there is a complete transition in characters within the group
from a strongly to a weakly convex pronotum, and from a coarsely
asperate to smooth anterior flope of the pronotum; in addition, many
species may breed successfully in either fruit (seeds) or phloem.
Coccotrypes indicus (Eggers) is an example where each of at least
two different authors have simultaneously recognized one synonym
as a valid species of Coccotrypes and another synonym as a valid
species of Poecilips, depending on the geographic area of origin of
the specimens examined. Under the circumstances it appears im-

172 GREAT BASIN NATURALIST Vol. 33, No. 3

possible to recognize more than one genus. A majority of species
included in the group are intermediate in characters expressed, al-
though the more familiar, economically important species represent
the extremes.

Gnathotrichus Eichhoff

Gnathotrichus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:75 (Type-species:
Gnathotrichus corthyloides Eichhoff = Tomicus materiarius Fitch, mono-
basic)

Ancyloderes Blackman, 1938, Proc. Ent. Soc. Washington 40:20 (Type-species:
Cryphalus pilosus LeConte, original designation) . New synonymy

Prognathotrichus Bright, 1972, Canadian Ent. 104:1678 (Type-species: Progna-
thotrichus primus Bright, original designation) . New synonymy

The type-species of Ancyloderes Blackman, pilosus (LeConte),
is known only from collections at light. Its relationship to other
species, for this reason, has been doubtful. Recently, several allied
species have been described from Quercus in Mexico, indicating a
clear relationship to Gnathotrichus. These species include obscurus
Wood, dentatus Wood, nimifrons Wood, and primus (Bright). These
species completely bridge the gap between Ancyloderes and Gnat ho
trichus. Since primus falls into the same species group with pilosus,
Paragnathotrichus Bright and Ancyloderes must be placed in
synonymy under Gnathotrichus.

Gnathotrupes Schedl

Gnathotrupes Schedl, 1951, Dusenia 2:125 (Type-species: Gnathotrupes bolivianus

Schedl, monobasic)
Gnathotrypanus Wood, 1968, Great Basin Nat. 28:9 (Type-species: Gnathotry-

panus terebratus Wood, original designation). New synonymy

In a review of this and related genera in North and Central
America, including most of the species described from South Ameri-
ca in Gnathotrichus, it was apparent that Gnathotrichus formed a
compact group quite distinct from another somewhat diverse but
definable genus from Central and South America. The name
Gnathotrupes Schedl apparently is the oldest available name for this
genus. To Gnathotrupes I assign all South American species de-
scribed in Gnathotrichus.. Gnathotrichus hituberculatus Blandford,
from Guatemala, and Gnathotrypanus terebratus Wood and electus
Wood, from Costa Rica. Through this action the genus Gnathotry-
panus becomes a synonym of Gnathotrupes. The characters on
which this division was based are discussed elsewhere.

Araptus hymenaeae (Eggers), n. comb.

Neodryocoetes hymenaeae Eggers, 1933, Mem. Trnv. Lab. d'Ent. Mus. Nat. d'Hist.
Nat., Paris 1(1):9 (Holotype, male; Gourdonville, Guayana Francaise; Paris
Museum)

Neodryocoetes insularis Eggers, 1940, Arb. Morph. Tax. Ent. Berlin-Dahlem
7:128 (Holotype, female; Guadeloupe; Fleuteaux Coll.). New synonymy

Neodryocoetes caribaeus Blackman, 1942, Proc. U. S. Nat. Mus. 92:185 (Holo-
type, female; Trinidad; U.S. Nat. Mus.). New synonymy

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 173

Neodryocoetes guianae Blackman, 1942, Proc. U.S. Nat. Mus. 92:186 (Holotype,
female; British Guiana; U.S. Nat. Mus.). New synonymy

Neodryocoetes hoodi Blackman, 1942, Proc. U.S. Nat. Mus. 92:187 (Holotype,
female; Taboga Island, Panama; U.S. Nat. Mus.). New synonymy

Male cotypes of hymenaeae (Eggers) and insularis (Eggers) and
the female holotypes of caribaeus (Blackman), guianae (Blackman),
and hoodi (Blackman) were examined and compared directly to one
another and to my material. Because of the limited material avail-
able to previous authors the extreme variability of the male frons
of this species was not detected and several synonyms resulted. Each
of several long series from Venezuela, Colombia, and Jamaica con-
tain examples of the described variations clearly indicating that they
all belong to one species.

Araptus laevigatas (Eggers), n. comb.

Pityophthorus laevigatus Eggers, 1933, Mem. Trav. Lab. d'Ent. Mus. Nat.

d'Hist. Nat., Paris 1(1) :6 (Holotype, female; St. Laurent du Maroni, Gua-

yana Francaise; Paris Mus.).
Neopityophthorus insularis Eggers, 1940, Arb. Morph. Tax. Ent. Berlin-Dahlem

7:i30 (Holotype, female; Trois Rivieres, Guadeloupe; deposited in but not

now in Eggers Coll.). New synonymy
Neodryocoetes guadeloupensis Schedl, 1951, Dusenia 2:73 (Replacement nam^e).

New synonymy

Cotypes of laevigatus Eggers and insularis Eggers were examined
and compared to my material from Central and northern South
America. All represent one species. It is common in fallen fruit on
the forest floor. This species is very similar to but distinct from
costaricensis (Schedl, 1938).

Araptus macer (Bright), n. comb.

Neodryocoetes macer Bright, 1972, Canadian Ent. 104:1666 (Holotype, female;

8 km or 5 miles S San Andres Tuxtla, Veracruz, Mexico; Canadian Nat. Coll.)
Neodryocoetes tuberculatus Bright, 1972, Canadian Ent. 104:1665 (Holotype,

female. Lake Catemaco, Veracioiz, Mexico; Canadian Nat. Coll.). New

synonymy

Topotypic para types of macer (Bright) and of tuberculatus
(Bright) were compared to one another and to my material from
Central America. The limited material before Bright evidently did
not include the variability found throughout the range of this species.
His material fits very well within those limits. The name macer
is selected for the valid name of this species under the first revisor
principle.

Araptus politus (Blandford), n. comb.

Pityophthorus politus Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):244

(Syntypes, male; Mexico; British Mus. Nat. Hist.)
Neodryocoetes hubbardi Blackman, 1942, Proc. U.S. Nat. Mus. 92:182 (Holotype,

female; Kingston, Jamaica; U.S. Nat. Mus.). New synonymy

The two syntypes of politus (Blandford) and the female holotype
of hubbardi (Blackman) were compared to my material. All repre-

174 GREAT BASIN NATURALIST Vol. 33, No. 3

sent the same common, rather widely distributed species. It occurs
from Veracruz to Costa Rica, in Haiti, and in Jamaica.

Araptus tabogae (Blackman)

N eodryocoetes tabogae Blackman, 1942, Proc. U.S. Nat. Mus. 92:184 (Holotype,

female; Taboga Island, Panama; U.S. Nat. Mus.).
Neodryocoetes vinealis Bright, 1972, Canadian Ent. 104:1667 (Holotype, female;

Ejipantla, 8 km or 5 miles S San Andres Tuxtla, Veracruz, Mexico; Canadian

Nat. Coll.). New synonymy

The holotypes of tabogae Blackman and vinealis Bright and 10
other specimens from Costa Rica were examined. While the Vera-
cruz record is a significant extension of the range of this species,
the specimen is entirely typical of the species. For this reason
vinealis is placed in synonymy as indicated above.

Cnesinus elegantis Wood

Cnesinus elegantis Wood, 1967, Great Basin Nat. 27:79 (Holotype, female; Vol-

can Zunil, Quezaltenango, Guatemala; Wood Coll.)
Cnesinus zapotecus Bright, 1972, Canadian Ent. 104:1493 (Holotype, female;

5 km or 3 miles N Suchixtepec, Oaxaca, Mexico; Canadian Nat. Coll.). New

synonymy

A topotypic female paratype of zapotecus Bright has the epi-
stomal tubercles slightly smaller and more closely spaced than is
seen in the type series of elegantis Wood, but except for this feature
it and a male paratype are identical to elegantis. However, Bright's
Figure 2 of this species is entirely normal for this species and re-
moves all doubt concerning the synonymy.

Coccotrypes advena Blandford

Coccotrypes advena Blandford, 1894, Trans. Ent. Soc. London 1894:100 (Holo-
type, female; Nagasaki, Japan; British Mus. Nat. Hist.)

Tharnnurgides persicae Hopkins, 1915, U.S. Dept. Agric. Kept. 99:45 (Holotype,
female; Honolulu, Hawaii; U.S. Nat. Mus.). New synonymy

This species has been introduced from the Indo-Malayan area
into .Japan, most Pacific Islands, northern South America, and
several of the Antilles islands, including Cuba. Although numerous
synonyms have been detected, the synonymy of persicae (Hopkins)
with advena Blandford, the oldest available name for the species,
has not been published. The holotypes of both species were ex-
amined and compared directly to my material. Schedl (1961, Rev.
Ent. Mocambique 4[2] : 728) treated persicae as a synonym of sannio
(Schaufuss, 1897), however, the interstrial bristles of a syntype and
of many other African specimens are shorter and more strongly
flattened. In all probability part or all of the African population
constitutes at least a different geographical race. Additional material
must be studied to resolve this problem. Regardless of the outcome,
the name advena has priority and will stand as the valid name of
the species.

Sept. 1973 WOOD: bark beetle synonymy 175

Corthylocurus barbatus (Blandford)

Brachyspartus barbatus Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):265

(Holotype, female; Volcan de Ghiriqui, Chiriqui, Panama; British Mus. Nat.

Hist).
Corthylocurus cincipnatus Bright, 1972, Canadian Ent. 104:1379 (Holotype,

female; 21 km or 13 miles N Ocozocoautla, Chiapas, Mexico; Canadian Nat.

Coll.). New synonymy

The female holotypes of barbatus (Blandford) and cincinnatus
Bright, and 86 other specimens from Puebla to Costa Rica, were
examined. The holotype of cincinnatus has the median frontal ele-
vation slightly higher and the epistomal tufts of hair slightly larger
than is seen in the average specimen, but all characters fall well
within the range of variation of this species. For this reason cincin-
natus is placed in synonymy as indicated above.

Corthylocurus mexicanus (Schedl)

Brachyspartus mexicanus Schedl, 1950, Dusenia 1:163 (Holotype, female; Comi-

tan, Chiapas, Mexico; Schedl Coll.).
Corthylus anomalus Bright, 1972, Canadian Ent. 104:1378 (Holotype, female;

5.6 km or 3.5 miles S Suchixtepec, Oaxaca, Mexico; Canadian Nat. Coll.).

New synonymy

The holotype of mexicanus (Schedl) is a female, not a male as
stated in the original description. This holotype and two topotypic
para types of anomalus were compared directly to my material; all
represent the same species.

Corthylus mexicanus Schedl

Corthylus mexicanus Schedl, 1950, Dusenia 1:159 (Holotype, male; Comitan,

Chiapas, Mexico; Schedl Coll.)
Corthylus glabinus Bright, 1972, Canadian Ent. 104:1372 (Holotype, female;

14 km or 9 miles SW Teopisca, Chiapas, Mexico; Canadian Nat. Coll.). New

synonymy

Two paratypes from 3.5 miles S Suchixtepec, Oaxaca, Mexico,
and the holotype of mexicanus Schedl were compared directly to my
material. One species is represented by these specimens. It is a
rather common species from Puebla and Veracruz to Chiapas.

Crypturgus pusillus (Gyllenhal)

Bostrichus pusillus Gyllenhal, 1813, Insecta Suecica descripta, Coleopt. 1(3):371

(Syntypes?; Sweden; Univ. Uppsala?).
Crypturgus atomus LeConte, 1868, Trans. Amer. Ent. Soc. 2:151 (Holotype, sex?;

New York; Mus. Comp. Zool.). New synonymy

In a review of the genus Crypturgus for North America most of
the Palaearctic species were examined in order to determine phyloge-
netic relationships. In doing this it was noted that atomus LeConte
could not be distinguished from the Eurasian pusillus. The odd pat-
tern of distribution of this species also suggests that it was introduced
into North America at an early date. This synonymy was indepen-
dently noted by Bright and reported to me through correspondence at

1 76 GREAT BASIN NATURALIST Vol. 33, No. 3

about the same time I detected it. The holotype of atomus was com-
pared directly to my North American specimens and these were com-
pared to 26 European specimens of pusillus.

Dendrocranulus carbonarius (Ferrari)

Xylocleptes carbonarius Ferrari, 1867, Die Forst- und Baumsuchtschadichen

Borkenkafer, p. 41 (Syntypes; Cuba; not located)
Xylocleptes floridensis Hopkins, 1915, U.S. Dept. Agric. Kept. 99:43 (Holotype,

female; Biscaj^ne Bay, Florida; U.S. Nat. Mus.). New synonymy
Xylocleptes anonae Hopkins, 1915, U.S. Dept. Agric. Kept. 99:43 (Holotype,

female; Florida; U.S. Nat. Mus.). New synonj-my

The holotypes of floridensis (Hopkins) and anonae (Hopkins)
were compared directly to one another and to my specimens from
Cuba. All represent the same species. These very old Cuban speci-
mens were labeled carbonarius by an unknown authority. They fit
Ferrari's description, but confirmation of their identity is needed.

Dendrocranulus cucurbitae (LeConte)

Xylocleptes cucurbitae LeConte, 1879, Bull. U.S. Geol. Survey 5:519 (Holotype,

female; Utah; Mus. Comp. Zool.).
Xylocleptes californicus Hopkins, 1915, U.S. Dept. Agric. Kept. 99:44 (Holotype,

female; Pomona, California; U.S. Nat. Mus.). New synonymy
Xylocleptes uenturina Hopkins, 1915, U.S. Dept. Agric. Kept. 99:44 (Holotype,

female; Ventura Co., California; U.S. Nat. Mus.). New synonymy
Xylocleptes punctatus Hopkins, 1915, U.S. Dept. Agric. Kept. 99:44 (Holotype,

female; Mesilla, New Mexico; U.S. Nat. Mus.). New synonymy

The holotypes of cucurbitae (LeConte), californicus (Hopkins),
uenturina (Hopkins), and punctatus (Hopkins) were examined and
compared to my material from Utah, California, New Mexico, and
Chihuahua. All clearly represent the same species. It is common in
dying stems of Cucurbita foetidissima and probably will be found
throughout the distribution of that plant.

Dendrocranulus macilentus (Blandford)

Dryocoetes macilentus Blandford, 1898, Biol. Centr. Anier., Coleopt. 4(6):190

(Lectotype, female; Jalapa, Veracruz, Mexico; British Mus. Nat. Hist., present

designation)
Dendrocranulus grossopunclatus Schedl, 1937, Arch. Instit. Biol. Veg. Rio de

Janeiro 3:155 (Holotype, female; San Isidro de Coronado, San Jose, Costa

Rica; Schedl Coll.). New synonymy

Of the two syntypes in Blandford's series the first, a female, is
here designated as the lectotype of macilentus (Blandford). This
lectotype and the female holotype of grossopunctatus Schedl were
compared directly to my female from Tapanti, Cartago, Costa Rica.
All three represent the same unusually slender species.

GnathotricJnis consentaneus (Blandford)

Gnathotrichus consentaneus Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):
247 (Lectotype, male; Totonicapan, Guatemala; British Mus. Nat. Hist.,
present designation).

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 177

Blandford included at least 18 specimens from several localities
under the name consentaneus Blandford when it was described but
stated that it was named from the Totonicapan specimens. In his
series pins 1, 7, and 8 bear Totonicapan specimens of this species;
the male on the first pin has been regarded as the type for many
years and is here designated as the lectotype of consentaneus. Pins
3 and 5 and apparently another specimen of this series now in the
Schedl collection bear specimens from Totonicapan of sulcatus
(LeConte). The remainder of the series will be discussed elsewhere.
The above action reestablishes the name consentaneus in the litera-
ture as a valid species.

Hylastes salebrosus (Eichhoff)

Hylastes salebrosus Eichhoff, 1868, Berliner Ent. Zeitschr. 12:146 (Lectotype,

female; Carolina; U.S. Nat. Mus., present designation).
Hylastes scobinosus Eichhoff, 1868, Berliner Ent. Zeitschr. 12:146 (Lectotype,

male?; Carolina; lost with Hamburg Mus.). New synonymy

Hylastes salebrosus Eichhoff and scobinosus Eichhoff were named
from syntypic series which were destroyed with the Hamburg Mu-
seum except for two cotypes of salebrosus now in the U.S. National
Museum of Natural History. The first of the two cotypes of salebro-
sus is a female of this species, the second is of porculus Erichson. I
here designate the first cotype as the lectotype of salebrosus Eichhoff.
A "cotype" of scobinosus in the U.S. National Museum was sent by
Eichhoff to Schwarz; however, that specimen is from "Tennese" and
cannot be a cotype. Because it is the only known specimen examined
by Eichhoff it is used as the basis of this species. It is a specimen
of salebrosus.

Hypothenemus seriatus (Eichhoff)

Stephanoderes seriatus Eichhoff, 1871, Berliner Ent. Zeitschr. 15:133 (Lectotype,
female; New Orleans, Louisiana; Brussels Mus., present designation).

Stephanoderes pulverulentus Eichhoff, 1871, Berliner Ent. Zeitschr. 15:133
(Syntypes?; Mexico; presumably lost with Hamburg Mus.). New synonymy

The names pulverulentus (Eichhoff) and seriatus (Eichhoff)
were validated on the same page. Since publishing synonymy in-
volving pulverulentus (Wood, 1972, Great Basin Nat. 32:50) I have
examined three female syntypes of seriatus. These syntypes are
identical to the material reported in that paper as pulverulentus.
Because the identity of pulverulentus was based on Eggers's homo-
types, the original series having been destroyed, I select seriatus
Eichhoff as the valid name for this species and designate the first
syntype in the Chapius series as its lectotype.

Ips spinifer (Eichhoff)

Tomicus spinifer Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:499 (Holotype,

sex?; California; lost with Hamburg Mus.).
Orthotomicus sabinianae Hopping, 1963, Canadian Ent. 95:64 (Holotype, male;

Middletown, Lake Co., California; California Acad. Sci.). New synonymy

1 78 GREAT BASIN NATURALIST Vol. 33, No. 3

The name spinifer (Eichhoff was placed in synonymy by Swaine
(1918, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(2): 114)
under the name latidens (LeConte). As a result. Hopping presumed
Swaine's action was correct, overlooked the description, and, conse-
quently, named sabinianae in his review of the genus. Eichhoff-
described the pair of large frontal tubercles and the large, cylindri-
cal, declivital spine 3 with sufficient clarity that there is no question
whatever as to the identity of his species even though the type is lost.

Leperisinus aculeatus (Say)

Hylesinus aculeatus Say, 1824, Jour. Acad. Nat. Sci. Philadelphia 3:322 (Syn-

types?; Missouri; evidently lost)
Hylesinus imperialis Eichhoff, 1868, Berliner Ent. Zeitschr. 12:149 (Syntypes;

Wisconsin and Georgia; lost with Hamburg Mus.). New synonymy

A review of all North American Leperisinus indicates that the
only species of this genus that could possibly occur in Wisconsin or
Georgia, and fall within the size limitations of 1^4"!/^ li^i- (2.65-
3.18 mm) set by Eichhoff for his Hylesinus imperialis, is aculeatus
(Say). Since aculeatus is a common and variable species, with re-
spect to the color pattern formed by the abundant scales, because all
syntypes of imperialis evidently were lost with the Hamburg Mu-
seum, and in the absence of all information that might suggest any
other alternative, I place imperialis in synonymy under aculeatus.

Micracis lignator Blackman

Micracis lignator Blackman, 1928, New York St. Coll. For., Syracuse, Tech.

Pub. 25:195 (Holotype, female; Tucson, Arizona; U.S. Nat. Mus.)
Micracis truncatus Wood, 1956, Canadian Ent. 88:152 (Holotype, female; 17 km

NE Jacala, Hidalgo, Mexico; Snow Ent. Mus., Univ. Kansas). New

synonymy

Since 1956, several series have been collected in Arizona and
Mexico that largely fill the gap in characters between lignator Black-
man and truncatus Wood. Although minute differences are still
apparent that might justify the recognition of subspecies, I here
place truncatus in synonymy under lignator.

Micracis swainei Blackman

Micracis swainei Blackman, 1920, Mississippi Agric. Expt. Sta. Tech. Bull. 9:32

(Lectotype, female; luka, Mississippi; U.S. Nat. Mus.)
Micracis photophilus Wood, 1956, Canadian Ent. 88:149 (Holotype, female; El

Salto, San Luis Potosi, Mexico; Snow Ent. Mus., tiniv. Kansas). New

synonymy

After examining more than 463 specimens taken from New York
to Honduras, including the type series of swainei Blackman and
photophilus Wood, I must recognize only one species among this
material. Specimens from the eastern United States are distinguish-
able from those from most of Mexico; however, the characters change
over a gradual, clinal gracHent making the recognition of geographi-
cal races difficult or impossible. It occurs most commonly in Salix
spp., although other hosts are acceptable.

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 179

Micracis suturalis LeConte

Micracis suturalis LeConte, 1868, Trans. Amer. Ent. Soc. 2:165 (Lectotype,

female; Illinois; Mus. Comp. Zool., present designation)
Micracis meridianus Blackman, 1920, Mississippi Agric. Expt. Sta. Tech. Bull.

9:29 (Lectotype, female; Agricultural College, Mississippi; U.S. Nat. Mus.,
present designation). New synon3Tny

The female syntype from Illinois, presently bearing type label
No. 1014 in the LeConte series, is here designated as the lectotype of
suturalis LeConte. The female in the syntypic series of meridianus
Blackman that was labeled "Type" by Blackman, but never so
designated, is here designated as the lectotype of meridianus Black-
man. These lecto types and all other syntypic specimens in the
Musetmi of Comparative Zoology and in the U.S. National Museum,
were examined and compared to my material. I am unable to see
differences in this material and place meridianus in synonymy under
LeConte's name.

Monarthrum querneus Wood

Monarthrum querneus Wood, 1967, Great Basin Nat. 27:50 (Holotype, male;

5 km or 3 miles E Morelia, Michoacan, Mexico; Wood Coll.).
Monarthrum bifidus Bright, 1972, Canadian Ent. 104:1382 (Holotype, male;

13 km or 8 miles E San Cristobal de las Casas, Chiapas, Mexico; Canadian

Nat. Coll.). New synonymy

Two topotypic paratypes of bifidus Bright were compared direct-
ly to the type series of querneus Wood. Only one species is repre-
sented by these specimens. This species occurs in Quercus logs from
Michoacan to Honduras.

Orthotomicus caelatus (Eichhoff)

Tomicus caelatus Eichhoff, 1868, Berliner Ent. Zeitschr. 11:402 (Syntypes;
Pennsylvania, Carolina; lost with Hamburg Mus. except two supposed syn-
types in the U.S. Nat. Mus.).

Xyleborus vicinus LeConte, 1874, Trans. Amer. Ent. Soc. 5:72 (Holotyp>e, female;
British Columbia; Mus. Comp. Zool.). New synonymy

Xyleborus punctipennis LeConte, 1878, Proc. Amer. Philos. Soc. \7:624' (Holo-
type, female; Marquette, Michigan; Mus. Comp. Zool.). New synonymy

The names vicinus (LeConte) and punctipennis (LeConte) have
been retained in the literature as distinct species largely because
their types were not seen by specialists. The holotypes of both
vicinus and punctipennis were examined and were found to repre-
sent normal specimens of caelatus (LeConte). Although the type
series of caelatus was mostly destroyed with the Hamburg Museum,
Eichhoff sent two presumed syntypes to the U.S. National Museum
that serve to fix the identity of his species.

Coccotrypes indicus (Eggers), n. comb.

Thamnurgides indicus Eggers, 1936, Ann. Mag. Nat. Hist. (10) 17:631 (Holo-
type, female; Sakalaspur, Mysore, India; British Mus. Nat. Hist.)
Xyleborus conspeciens Schedl, 1936, Archiv. Inst. Biol. Veg. Rio de Janeiro 3:110

180 GREAT BASIN NATURALIST Vol. 33, No. 3

(Holotype, female; locality not given, presumably Brazil; Schedl Coll.)-

New synonymy
Coccolrypes insularis Eggers, 1940 (nee Eggers 1940:127), Arb. Morph. Tax.

Ent. Berlin-Dahlem 7:129 (Lectotype, female; Trois-Rivieres, Guadeloupe;

U.S. Nat. Mus., designated by Anderson and Anderson, 1971, Smithsonian

Contrib. Zool. 94:15). New synonymy
Poecilips eggersi Schedl, 1952, Dusenia 3:347 (Replacement name). New

synonymy

The holotypes of indicus (Eggers) and conspeciens (Schedl), and
the lectotype of insularis Eggers were all examined and compared to
my material. Only one species is represented. This species has been
introduced from the Indo-Malayan area into northern South Ameri-
ca, Central America, the Antilles islands, and one series was inter-
cepted in Florida, although it probably is not yet established there.

Phloeoborus asper Erichson

Phloeoborus asper Erichson, 1836, Archiv Naturgesch. 2(1):55 (Holotype, male;
Brazil; Zool. Mus. Berlin)

Phloeoborus ovatus Chapuis, 1869, Synopsis des Scoly tides, p. 15 (Lectotype, male;
Cayenne; Brussels Mus., present designation). New synonymy

Phloeoborus rugatus Blandford, 1897, Biol. Centr. Amer. Coleopt. 4(6):153 (Lec-
totype, female; Chontales, Nicaragua; British Mus. Nat. Hist., present
designation). New synonymy

The male holotype of asper Erichson, the three male syntypes of
ovatus Chapuis, and the two female syntypes of rugatus Blandford
were all compared to my long series of this species from Venezuela.
It is obvious that all three names apply to the same, easily recognized
species. The first syntype of ovatus and the first syntype of rugatus
are here designated as lectotype of their respective species as indi-
cated above.

Phloeoborus rudis Erichson

Phloeoborus rudis Erichson, 1836, Archiv Naturgesch. 2(1) :55 (Lectotype, female;

Brazil; Zool. Mus. Berlin, present designation)
Phloeoborus elongatus Chapuis, 1869, Synopsis des Scolytides, p. 13 (Holotype,

male; Brazil; Brussels Mus.). New synonymy
Phloeoborus rugipennis Eggers, 1942, Arb. Morph. Taxon. Ent. Berlin-Dahlem

9:271 (Holotype, female; San Salvador; U.S. Nat. Mus.). New synonymy

Of the specimens of rudis Erichson now in the Zoologische Mu-
seum in Berlin, it appears that only the first two, both females,
were in the original series. Since the first of these two syntypes has
been regarded as the type, it is here designated as the lectotype of
rudis as indicated above. This lectotype was compared directly to
the male holotype of elongatus and to my specimens from Panama
and Peru; my specimens were later compared to the female holotype
of rugipennis. All apparently represent the same slightly variable
species.

Phloeotribus armatus Blandford

Phloeotribus armatus Blandford, 1897, Biol. Centr. Amer.. Coleopt. 4(6): 166
(Holotype, male; Volcan de Chiriqui, Panama; British Mus. Nat. Hist.)

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 181

Phloeotribus mixtecus Bright, 1972, Canadian Ent. 104:1494 (Holotype, female;
26 miles S Juchatengo, Oaxaca, Mexico; Canadian Nat. Coll.). New synonymy

Two topotypic paratypes of mixtecus Bright were compared to
my series that previously had been compared to the male holotype
of armatus Blandford. This species occurs from southern Mexico
to Panama.

Phloeotribus demessus Blandford

Phloeotribus demessus Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6): 165
(Lectotype, female; Toxpam, a probable misspelling of Tuxpan, Veracruz,
Mexico; British Mus. Nat. Hist.).

Phloeotribus tuberculatus Eggers, 1951, Ent. Blatt. 46:147 (Holotype, female;
Turrialba, Cartago, Costa Rica; U.S. Nat. Mus.). New synonymy

The second specimen, a female, in the Blandford series of demes-
sus has been labeled type and has been regarded as the type of this
species, although it has never been so designated. I here designate
that female S3mtype as the lectotype of demessus. This specimen
was compared to my series and some of these were later compared
to the female holotype of tuberculatus Eggers; all clearly represent
the same species.

Phloeotribus pilula Erichson

Hylesinus pilula Erichson, 1847, Archiv Naturgesch. 13(1): 138 (Lectotype, male;

Peru; Zool. Mus. Berlin, present designation)
Phloeotribus obliquus Chapuis, 1869, Synopsis des Scolytides, p. 45 (Syntypes;

Mexique, Nouvelle-Grenada; Brussels Mus.). New synonymy
Phloeotribus manni Blackman, 1943, Proc. U.S. Nat. Mus. 94:385 (Holotype,

female; Rio Madeira, Brazil; U.S. Nat. Mus.). New synonymy

Of three specimens presumed to be syntypes of pilula (Erichson)
now in the Zoological Museum in Berlin, only the first specimen,
a male, is of the species usually designated by this name. The second
and third specimens are females of a different species not presently
known to me and probably were not part of the original series. In
order to avoid possible confusion at a later date, I here designate the
first specimen in the series as the lectotype of pilula. This same
species was named obliquus by Chapuis. Under the name obliquus
in the Brussels Museum were four specimens in 1969; the first syn-
type is of transversus Chapuis (apparently remounted on the wrong
pin when the glue deteriorated) and did not fit the description; the
second syntype did fit the description and it was identical with the
third and fourth specimens labeled "ex-typis." In 1971, when I
reexamined these specimens, the second syntype was missing from
the pin. That specimen previously had been compared to my series
from Peru and to the two Chapuis ex-typis specimens and are of the
same species as the lectotype of pilula. The holotype of manni Black-
man was also compared directly to my specimens from Peru and is
also this species.

This species is abundant in Brosmium sp. from Chiapas, Mexico,
to Peru and Brazil.

182 GREAT BASIN NATURALIST Vol. 33, No. 3

Phloeotribus setulosus Eichhoff

Phloeotribus setulosus Eichhoff, 1868, Berliner Ent. Zeitschr. 12:149 (Lectotype,

male; Colombia; Brussels Mus., present designation).
Phloeotribus asperatus Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6):166

(Holotype, male; Panachel, Guatemala; British Mus. Nat. Hist.). New

synonymy

Two syntypes, a male and a female, of setulosus Eichhoff are in
the Brussels Museum. The male, labeled "Colombie," not Carolina,
as reported by Eichhoff, is here designated as the lectotype of this
species. It is identical to the male holotype of asperatus Blandford.
Both types were compared directly to my material from Central
and South America and they obviously represent the same species.

This is the most abundant and widely distributed Central and
South American species of Phloeotribus; it occurs in a wide variety
of host trees. It is easily misidentified, and numerous specimens in
museums under this name are not of this species.

Pityoborus frontalis Wood

Pityoborus frontalis Wood, 1971, Brigham Young Univ. Sci. Bull. Biol. Ser.

15 (3): 49 (Holotype, female; 13 km S El Cameron, Oaxaca, Mexico; Wood

Coll.).
Pityoborus severus Bright, 1972, Canadian Ent. 104:1676 (Holotype, female;

5 km or 3 miles N Suchixtepec, Oaxaca, Mexico; Canadian Nat. Coll.). New

synonymy

Two paratypes of severus Bright from 8 miles S Miahuatlan,
Oaxaca, Mexico, were compared to the holotype and allotype of
frontalis Wood. They are identical in every respect.

Pityoborus secundus Blackman

Pityoborus secundus Blackman, 1928, Bull. New York St. Coll. For., Syracuse,

Tech. Pub. 25:146 (Holotype, female; La Sal Mts., Utah; U.S. Nat. Mus.)
Pityoborus tertius Blackman, 1942, Proc. U.S. Nat. Mus. 92:202 (Holotype,

female; Chalco, Districto Federal, Mexico; U.S. Nat. Mus.). New synonymy
Pityoborus intonsus Wood, 1958, Great Basin Nat. 28:54 (Holotype, female; 23

km or 14 miles W Texmelucan, Puebla, Mexico; Snow Ent. Mus., Univ.

Kansas) . New synonymy
Pityoborus immitus Bright, 1972, Canadian Ent. 104:1674 (Holotype, female;

68 km W Durango, Durango, Mexico; Canadian Nat. Coll.). New synonymy
Pityoborus ramosus Bright, 1972, Canadian Ent. 104:1677 (Holotype, female;

5.6 km or 3.5 miles S Suchixtepec, Oaxaca, Mexico; Canadian Nat. Coll.).

New synonymy

This species has been collected at 14 rather widely separated
localities from Utah to Oaxaca. The average specimen in each series
is slightly different from those in almost every other series. Several
independent clinal variations between series are also apparent.
However, the differences are so minute and inconsistent within most
series that the only logical solution appears to be the grouping of all
of this material into one species at least until considerably more
material is available for study from many more localities. This

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 183

decision was based on 169 specimens including the holotypes and
type series of secondus Blackman, tertius Blackman and intonsus
Wood, and on female paratypes of immitus Bright and ramosus
Bright. The last four names listed, therefore, are placed in synonymy
under secundus.

Pycnarthrum hispidum (Ferrari)

Hypoborus (?) hispidus Ferrari, 1867, Die Forst.- und Baunzuchtschadlichen
Borkenkafer, p. 19 (Syntypes?, Cuba; presumably in Vienna Mus.)

Nemobius lambottei Chapuis, 1869, Synopsis des Scolytides, p. 42 (Lectotype,
female; Teapa, presumably Tabasco, Mexico; Brussels Mus., present desig-
nation). New synonymy

Pycnarthrum gracile Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:104 (Syn-
types?; Cuba; some syntypes lost with Hamburg Mus., one apparent syn-
type in U.S. Nat. Mus.). New synonymy

Pycnarthrum quadraticolle Eichhoff, 1878,, Mem. Soc. Roy. Sci. Liege (2)8:106
(Lectotype, male; Mexico; U.S. Nat. Mus., present designation). New
synonymy

Pycnarthrum transversum Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6):
177 Lectotype, female; Mirandilla, Guatemala; British Mus. Nat. Hist.,
present designation). New synonymy

Pycnarthrum reimoseri Schedl, 1934, Ent. Blatt. 30:208 (Syntypes, females;
Jimenez on Osa Penninsula, and Volcan Irazu, Costa Rica; Vienna Mus.
and Schedl. Coll.). New synonymy

Pycnarthrum reticulatus Schedl, 1940, An. Esc. Nac. Cienc. Biol., Mexico, 1:335
(Syntypes; Tonala and Mapastec in Chiapas, Tuxtepec in Oaxaca, and
Cardel in Veracruz, Mexico; Schedl and Dampf Collections). New synonymy

This is the only widely distributed, common Pycnarthrum species
in Mexico, Central America, and southern Florida to Guadeloupe.
It breeds in Ficus spp. and it is readily attracted to light. It is the
only species in this genus known to occur north of Honduras and
Guadeloupe Island.

Authentic specimens of hispidus (Ferrari) have not been ex-
amined although several specimens supposedly compared to the
types by Eggers were examined; since no other species of this genus
occurs in Cuba, the association appears certain. The three syntypes
of lambottei (Chapuis) were examined and the first of these, a female
from Teapa, is here designated as the lectotype of Nemobius lambot-
tei Chapuis. A specimen in the U.S. National Museum, sent by
Eichhoff, bears the labels '^'Pycnarthrum gracile Eich., Type, Cuba,
50," and is presumed to be a syntype. That specimen is here desig-
nated as the lectotype of Pycnarthrum gracile Eichhoff, since the
remainder of the Eichhoff series was destroyed with the Hamburg
Museum. From syntypes of quadraticolle Eichhoff in the U.S.
National Museum a male was selected, labeled, and is here desig-
nated as the lectotype of this species. Blandford based transversum
on four female syntypes; the left specimen (with the elytra com-
pletely closed) on the first pin in this series is here designated as the
lectotype of transversum Blandford. One female cotype at the U.S.
National Museum and several other specimens identified by Schedl
as reimoseri Schedl were examined; several specimens identified by
Schedl as reticulatus Schedl were also studied. All of the above

184 GREAT BASIN NATURALIST Vol. 33, No. 3

material and several hundred other specimens from throughout its
distribution were examined and found to represent one species.

Pseudopityophthorus declivis Wood

Pseudopityophthorus declivis Wood, 1971, Brigham Young Univ., Sci. Bull. Biol.

Ser. 15(3):50 (Holotype, female; Laguna Santa Maria, Nayarit, Mexico;

Wood Coll.)
Pseudopityophthorus truncatus Bright, 1972, Canadian Ent. 104:1673 (Holotype,

male; 184 km S Oaxaca, Oaxaca, Mexico; Canadian Nat. Coll.). New

synonymy
Pseudopityophthorus curtus Bright, 1972, Canadian Ent. 104:1674 (Holotype,

female?; km or 8 miles N Ocosingo, Chiapas, Mexico; Canadian Nat. Coll.).

New synonymy

The holotypes of declivis Wood and curtus Bright and two topo-
typic paratypes of truncatus Bright were compared directly to one
another. The specimen of declivis is intermediate in size between
the slightly smaller curtus and the slightly larger truncatus. The
declivital punctures in truncatus are distinctly smaller, particularly
toward the apex and the interstrial bristles are very slightly longer
and finer. The posterior areas of the pronotum of curtus are more
brightly shining, but the prothoraxic joint has been broken and the
anterior unit remounted with the posterior areas; the possibility
exists that the anterior unit came from another slightly larger speci-
men. In view of the paucity of material from widely separated
areas I presently consider truncatus and curtus to be synonyms of
declivis. It is possible that curtus might eventually be restored to
subspecific rank when sufficient material is available from more
localities.

Pseudopityophthorus hondurensis Wood

Pseudopityophthorus hondurensis Wood, 1967, Great Basin Nat. 27:42 (Holotype,

male; Buenos Aires, Cortes, Honduras; Wood Coll.)
Pseudopityophthorus montanus Bright, 1972, Canadian Ent. 104:1667 (Holotype,

male; Mt. Tzontehuitz, Chiapas, Mexico; Canadian Nat. Coll.). New

synonymy

Two topotypic paratypes of montanus Bright were compared to
the type series of hondurensis Wood. Only one species is represented
by this material.

Pseudopityophthorus micans Wood

Pseudopityophthorus micans Wood, 1967. Great Basin Nat. 27:44 (Holotype,
male, 96 km or 60 miles W Durango, Durango, Mexico; Wood Coll.)

Pseudopityophthorus squamosus Bright, 1972, Canadian Ent. 104:1670 (Holotype,
female; 14 km or 9 miles W La Ciudad, Durango, Mexico; Canadian Nat.
Coll.). New synonymy

A topotypic female paratype of squamosus Bright was compared
to the type series of micans Wood. Although most paratypes of
micans have the elytra entirely glabrous, some have declivital scales
exactly as seen in squamosus. Bright's name squamosus must be
placed in synonymy under micans.

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 185

Pseudopityophthorus opacicollis Blackman

Pseudopityophthorus opaciocollis Blackman, 1931, J. Washington Acad. Sci. 21:
235 (Holotype, male; Santa Catalina Mts., Arizona; U.S. Nat. Mus.).

Pseudopityophthorus aesculinus Bright, 1972, Canadian Ent. 104:1672 (Holotype,
female; 112 km or 70 miles N Oaxaca, Oaxaca, Mexico; Canadian Nat. Coll.).
New synonymy

Two topotypic paratypes of aesculinus Bright were compared to
two paratypes and 38 other specimens of opacicollis Blackman taken
from Arizona to Tlaxcala, Mexico. This species is somewhat more
variable than was indicated in Blackman's description, but the para-
types of aesculinus are average specimens and Bright's species must
be placed in synonymy under opacicollis.

Pseudopityophthorus pruinosus (Eichhoff)

Pityophthorus pruinosus Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:187 (Syn-

types; Carolina; most were lost with the Hamburg Mus., one is in the U.S.

Nat. Mus.)
Pseudopityophthorus pulvereus Blackman, 1931, J. Washington Acad. Sci. 21:232

(Holotype, male; Chiricahua Reserve, Arizona; U.S. Nat. Mus.). New

synonymy
Pseudopityophthorus tropicalis Wood, 1967, Great Basin Nat. 27:43 (Holotype,

male; Zamorano, Morazan, Honduras; Wood Coll.). New synonymy
Pseudopityophthorus convexus Bright, 1972, Canadian Ent. 104:1672 (Holotype,

male; 184 km or 115 miles S Oaxaca, Oaxaca, Mexico; Canadian Nat. Coll.).

New synonymy

The only known syntype of pruinosus (Eichhoff), the type series
of pulvereus Blackman and of tropicalis Wood, one topotypic para-
type of convexus Bright, and 83 other specimens of this species from
Arizona and Mexico were examined in a review of all species of
this genus. In attempting to make a key to identify the described
species it was noted that some of the minute characters on which
these species were based were not consistent within or between
series. It became necessary to reevaluate my concept of this species
and broaden it to include pulvereus, tropicalis, and convexus as
synonyms of pruinosus.

Pseudopityophthorus singularis Wood

Pseudopityophthorus singularis Wood, 1971, Brigham Young Univ. Sci. Bull.

Biol. Ser. 15(3) :50 (Holotype, male; 1 km W Las Vigas, Veracruz, Mexico;

Wood Coll.)
Pseudopityophthorus acuminatus Bright, 1972, Canadian Ent. 104:1671 (Holotype,

male; 13 km or 8 miles NE San Cristobal de las Casas, Chiapas, Mexico;

Canadian Nat. Coll.). New synonymy

Two topotypic paratypes of acuminatus Bright were compared
to the type series of singularis. Only one species is represented by
this material.

Pseudopityophthorus tenuis Wood

Pseudopityophthorus tenuis Wood, 1959, Great Basin Nat. 19:1 (Holotype, male;
18 km or 11 miles NE Jacala, Hidalgo, Mexico; Snow Ent. Mus., Umv.
Kansas)

186 GREAT BASIN NATURALIST Vol. 33, No. 3

Pseudopityophthorus hirsutus Bright, 1972, Canadian Ent. 104:1668 (Holotype,
male; 8 km or 5 miles SE Teopisca, Chiapas, Mexico; Canadian Nat. Coll.).
New synonymy

A topotypic paratype of hirsutus Bright was compared to the
type series and to 22 other representatives of this species from Vera-
cruz. The sculpture of the frons is slightly variable throughout the
range of the species, but the specimen of hirsutus fits well within
the limits of that variation.

Stenocleptus sulcatus (Bruck)

Pseudothysanoes sulcatus Bruck, 1936. Bull. S. California Acad. Sci. 35:33 (Holo-
type, male; Mt. Wilson, Los Angeles Co., California; Ohio State Univ.
Coll.).

Stenocleptus ceanothi Blackman, 1943, Proc. U.S. Nat. Mus. 93:358 (Holotype,
female; Yosemite Nat. Pk., California; U.S. Nat. Mus.). New synonymy

Stenocleptus rhois Blackman, 1943, Proc. U.S. Nat. Mus. 93:357 (Holotype,
female; Orange Co., California; U.S. Nat. Mus.). New synonymy

This rare species is known from five short series in which most
of the specimens are in poor condition caused by the adherence of
plant resins and frass to the beetles. Minute differences may be
seen between every series, although their basic features are identical.
Until considerably more material is available I place ceanothi Black-
man and rhois Blackman in synonymy under sulcatus (Bruck). The
entire type series of both of Blackman's species, two paratypes of
sulcatus, and four other specimens were used as the basis for this
synonymy.

Thysanoes texanus Blackman

Thysanoes texanus Blackman, 1943, Proc. U.S. Nat. Mus. 93:353 (Holotype,

female; Brownsville, Texas; U.S. Nat. Mus.).
Thysanoes vachelliae Blackman, 1943, Proc. U.S. Nat. Mus. 93:353 (Holotype,

female; Brownsville, Texas; U.S. Nat. Mus.). New synonymy
Thysanoes ratamae Blackman, 1943, Proc. U.S. Nat. Mus. 93:354 (Holotype,

female; Mexico; U.S. Nat. Mus.). New synonymy

Following a review of all described species of Thysanoes, includ-
ing 160 specimens of this species and a direct comparison of their
holotypes, it was concluded that texanus Blackman, vachelliae Black-
man, and ratamae Blackman all represent one common species that
occurs from southern Texas to Veracruz and Colima. The unique
type of ratamae is in very poor condition, but frontal characters in-
sure the correctness of my identification.

Irypopholeus striatulus (Mannerheim)

Cryphalus striatulus Mannerheim, 1853, Bull. Soc. Imp. Nat. Moscou 26:235
(Holotype, sex?: Lake Skeljanima, Kenai Penninsula, Alaska; presumably

lost)

Trypophloeus nitidus Swaine, 1912, Canadian Ent. 44:349 (Lectotype, sex?;
Weymouth, Nova Scotia; Canadian Nat. Coll., designated by Bright, 1967,
Canadian Ent. 99:679). New synonymy

Sept. 1973 WOOD: BARK BEETLE SYNONYMY 187

The Mannerheim collection contains no specimens under the
name striatulus (Mannerheim) nor are there American specimens
of this or a related genus among his undetermined American ma-
terial. Mannerheim compared this species to the familiar European
Trypophoeus granulatus (Ratzeburg). Considering Mannerheim's
knowledge of the European fauna and the fact that the only species
of American Cryphalini occurring north of British Columbia is
nitidus Swaine, a species very similar to granulatus, I place nitidus
in synonymy under striatulus. My specimen from Cantwell, Alaska,
about 200 miles north of the type locality, is entirely typical of the
species and confirms the existence of this species that far north.

Xyleborus capucinus Eichhoff

Xyleborus capucinus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:281 (Holotype,

female; Guadeloupe Island; Brussels Mus.)
Xyleborus rufithorax Eichhoff, 1869, Berliner Ent. Zeitschr. 12:281 (Holotype

female; Brazil; Brussels Mus.). New synonymy

The female holotypes of capucinus Eichhoff and ruftithorax
Eichhoff were examined and compared directly to my specimens.
The holotype of capucinus is callow, partly crushed, and in poor
condition, and the prothorax of rufithorax is not fully colored, but
there is no question whatever as to their synonymy. The species is
now known from Nayarit and Veracruz, Mexico, to Colombia and
Brazil.

Xyleborus pubescens Zimmermann

Xyleborus pini Say (??): Eichhoff, 1868, Berliner Ent. Zeitschr. 11:401 (Er-

roneus identification)
Xyleborus pubescens Zimmermann, 1868, Trans. Amer. Ent. Soc. 2:145 (Lecto-

type, female; Southern States; Mus. Comp. Zool., present designation).
Xyleborus propinguus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:281 (Lectotype,

female; Amerique boreali; Brussels Mus., present designation). New

synonymy

Various authors have treated as valid a species designated as
Xyleborus pini Eichhoff. However, when the literature is searched
the only Eichhoff reference to such a name is the erroneous identifi-
cation and description of Say's species and the citation of the same
species in synonymy. Neither reference validated this name; con-
sequently, the oldest available name for this taxon is pubescens
Zimmermann. The first of two female syntypes now in the LeConte
collection is here designated as the lectotype of pubescens Zimmer-
mann. The female syntype in the Brussels Museum is here desig-
nated as the lectotype of propinguus Eichhoff; it was compared di-
rectly to my homotypes of pubescens and is identical in all respects.

Xylosandrus curtulus Eichhoff

Xyleborus curtulus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:281 (Holotype,
female; Brazil; Institut Royal des Sciences Naturelles de Belgique, Brussels)

Xyleborus biseriatus Schedl, 1963, Reichenbachia 1:226 (Holotype, female; Nova
Teutonia, Santa Catarina, Brazil; Schedl Coll.). New synonymy

188 GREAT BASIN NATURALIST Vol. 33, No. 3

The holotype of Xyleborus curtulus Eichhoff was compared di-
rectly to specimens which previously had been compared directly to
the holotype of biseriatus Schedl. Only one species is represented.

Because a series from Caicedonia, Valle de Cauca, Colombia,
indicated intergradation between this species and zimmermanni
(Hopkins), they were treated as synonyms (Wood, 1966, Great
Basin Nat. 26: 33) . Since then, many additional series of both species
have been taken in a broad zone of sympatry without additional
indications of intergradation. Both species are now considered valid,
with curtulus occurring from Nayarit and Veracruz, Mexico, to
Brazil, and zimmermanni from Florida and Guatemala to Vene-
zuela.

UNDESCRIBED SPECIES OF NEARCTIC TIPULIDAE
(DIPTERA), XI

Charles P. Alexander^

Abstract. — New species and subspecies of Tipulidae from California and
Alaska are described, including Tipula (Trichotipula) frommeri, Tipula (Tri-
chotipula) sanctaecruzae, and Tipula (Lunatipula) hastingsae diperona, from
California, and Tipula (Pterelachisus) macleani and Limnophila byersi from
Alaska.

The preceding paper in this series was pubhshed in the Great
Basin Naturalist 29:1-10.

The species discussed at this time are derived from three sources:
two from California, the third from Alaska. The first Calif omian
series is from the California Channel Islands in the Channel Islands
National Monument west of Los Angeles including Santa Cruz, San
Miguel, and San Nicolas islands, the materials having been collected
chiefly by Dr. Charles L. Remington, his son, Eric E. Remington,
and L. Serge Matlovsky. The named materials are preserved in the
Peabody Museum, Yale University, New Haven, Connecticut. The
second Californian collection is from various canyons in and near
Death Valley, Panamint Mountains, in Riverside and San Bernardino
counties, chiefly in conjunction with the P. L. Boyd Deep Canyon
Desert Research Center. The survey is under the direction of Dr.
Saul I. Frommer, presently assisted by Messrs. Irwin and LePre.
The materials are to be preserved in the University of California,
Riverside, Research Collection. The Alaskan materials are from Dr.
George W. Byers of the University of Kansas, Lawrence; the speci-
mens having been taken by Stephen M. MacLean, made in conjunc-
tion with the United States Biome Tundra Program at Prudhoe Bay,
Alaska. I express my sincere thanks to all of the above for the
privilege of examining these particularly interesting species.

Tipula (Trichotipula) frommeri, n. sp.

General coloration of thorax yellow, praescutal stripes bordered
by pale brown, in cases the median area paler; legs yellow, tips of
femora narrowly brown; wings weakly darkened, cell Sc and the
stigma light brown, no trichia in outer wing cells; abdominal tergites
conspicuously bicolored, bases broadly yellow, posterior borders
brown, sternites and hypopygium more uniformly yellowed; male
hypopygium with the beak of the inner dististyle long, with con-
spicuous setae.

Male.— Length about 12 mm; wing 10 mm; antenna about 5.5
mm.

Female. — Length about 13 mm; wing 11 mm; antenna about

2.5 mm.

'Amherst, Massachusetts.

189

190 GREAT BASIN NATURALIST Vol. 33, No. 3

Described from alcoholic materials. Frontal prolongation of head
slightly shorter than the remainder, yellow; nasus long and slender;
palpi with proximal two segments brown, third slightly paler, ter-
minal segment elongate, light yellow. Antennae of male elongate;
scape and pedicel yellow, flagellum dark brown, flagellar segments
with conspicuous basal enlargements, verticils shorter than the seg-
ments. Head brownish yellow.

Pronotum yellow, center of scutum restrictedly darker. Meso-
notal praescutum with four yellow stripes bordered by pale brown,
in cases median area paler to produce a vague central stripe, as in
dorsolineata and some others; scutal lobes yellow, ringed with pale
brown; scutellum and postnotum brownish yellow, with erect setae.
Pleura and pleurotergite yellowed, restrictedly darkened on cervical
region, propleura, and pleurotergite. Halteres with stem yellow, knob
infuscated. Legs with coxae and trochanters light yellow; femora
yellow, appearing darker from abundant short darkened vestiture,
tips narrowly brown; tibiae obscure yellow, tarsi passing into dark
brown; claws small, simple. Wings (Fig. 1) weakly darkened, cell
Sc and the stigma light brown; veins darker brown. Stigma with
two or three microscopic trichia, outer cells without trichia. Vena-
tion: Petiole of cell Mi from about one-half to two-thirds m; m-cu
on M4.

Abdominal tergites conspicuously bicolored, bases broadly yel-
low, posterior borders brown, on the proximal segments including
also the narrow bases of the segments; sternites and hypopygium
more uniformly yellow. Abdomen with conspicuous black setae.
Male hypopygium (Fig. 4) with vestiture of tergal lobes, t, restricted
in numbers, setae of emargination longer than a group of about 15
small blackened points near apex of each lobe. Basistyle, b, with a
group of strong setae on outer margin of apex with more sparse
longer bristles on inner side. Dististyles, d, as shown; outer style
with very long black setae, longest exceeding transverse diameter of
style; inner style with beak long, setae of base long and conspicuous,
outer ones progressively shorter, apex of beak with numerous, very
long, more delicate, yellow setae.

Habitat.— California.

HoLOTYPE, cf, alcoholic, P. L. Boyd Desert Research Center,
Riverside County, Thunderbolt, Horsethief Creek, gaging station in
first permanent pool. Malaise, 15 May 1970 (Saul I. Frommer).
Allotopotype, 9 , Deep Canyon, 4 mi. south of Palm Desert, 5680
ft., 24 May 1969 (M. E. Irwin and Saul I.' Frommer) ; 24-471.
Paratopotype, cT, in Alexander Collection, 3.5 mi. south of Palm
Desert, at Marker 58, 15 May 1969 (Frommer and LaPre); 24-472.

This interesting fly is named for Dr. Saul I. Frommer, capable
student of the Diptera, in charge of the Deep Canyon Research
Project. By means of the author's key to the Californian species of
the genus Tipula (Crane flies of California, pp. 56-57; 1967), the
present fly runs to couplets 11 and 12, including three species in the
subgenus Trichotipula that lack trichia in the outer wing cells, these

Sept. 1973

ALEXANDER: NEW CRANE FLIES

191

Figs. 1-3. Wing venation: 1, Tipula (Trichotipula) frommeri, n. sp.; 2,
Tipula {Pterelachisus) macleani, n. sp.; 3, Limnophila byersi, n. sp. Figs. 4-10.
Male hypopygia: 4, Tipula {Trichotipula) frommeri, n. sp.; 5a, Tipula {Trichoti-
pula) sanctaecruzae, n. sp.; 5b, Tipula {Trichotipula) capistrano Alexander; 5c,
Tipula {Trichotipula) beatula Osten Sacken; 6, above Tipula {Lunatipula) hast-
ingsae hastingsae Alexander; 6, below, Tipula {Lunatipula) haslingsae diperona,
n. subsp.; 7, Tipula {Pterelachisus) macleani, n. sp.; 8, Tipula {Vestiplex) ber-
grothiana Alexander; 9, Tipula {Vestiplex) aldrichiana; 10, Limnophila byersi,
n. sp. Fig. 11. Male antenna of Limnophila byersi, n. sp. (Symbols: a, aedeagus;
b, basistyle; d, dististyles; g, gonapophysis; s, stemite; t, tergite).

192 GREAT BASIN NATURALIST Vol. 33, No. 3

being Tipula {Trichotipula) desertorum Alexander, T. (T.) dorso-
lineata Doane, and T. (T.) kennedyana Alexander, all differing evi-
dently in hypopygial structure, especially the dististyles.

Tipula {Trichotipula) sanctaecruzae, n. sp.

Allied to beatula and capistrano; head and thorax patterned with
dark brown and yellow, mesonotal mediotergite with a narrow yel-
low central line; male hypopygium with tergal lobes truncate; appen-
dage of ninth sternite including a long outer lobe, the lateral lobe
low.

Male.— Length about 13 mm; wing 13.5 mm; antenna about
3 mm.

Female. — Length about 14-15 mm; wing 13 mm.

Frontal prolongation of head blackened, narrowly yellow on
sides, ventrally more reddish yellow; palpi black. Antennae with
scape light brown, tip paler, remainder of organ black; flegellar
segments elongate. Head orange on sides, central part of vertex and
occiput brownish black.

Pronotum brownish black. Mesonotal praescutum on disk with
four brownish gray stripes broadly bordered by black, lateral margins
of praescutum light orange; scutal lobes dark brownish gray, with a
narrow central polished black line, median region light yellow;
scutellum light yellow with a narrow central black line, parascutella
black; postnotal mediotergite brownish black with a narrow median
yellow vitta on anterior half, in cases this less distinct, anapleuroter-
gite yellowed, katapleurotergite with central part restrictedly yel-
lowed. Pleura brownish gray, patterned with yellow, more evident
in certain individuals, including the dorsal sternopleurite, ptero-
pleurite, and dorsal meral region. Halteres with stem yellow, base
narrowly orange, knob brownish black, apex restrictedly yellowed.
Legs with coxae brownish gray, trochanters yellow; femora brownish
yellow, apices dark brown; tibiae brownish yellow, tarsi passing
into black. Wings generally as in beatula and capistrano^ differing
only in minor details.

Abdomen with segments conspicuously patterned with brownish
black and yellow, on tergites the dark color including posterior
border and a broad central stripe. Male hypopygium (Fig. 5a) with
ninth tergite, t, having lobes truncate, spinoid setae small and sparse,
as shown; median region bordering emargination with abundant
microscopic setulae. Appendage of ninth sternite, .?, shaped as
shown, including a major outer lobe provided with numerous chiefly
marginal long twisted yellow setae, lateral lobe low. In beatula
(Fig. 5c) tergite generally similar, appendage of ninth sternite, s,
distinctive, with a slender lateral lobe, as shown. In capistrano
(Fig. 5b) tergal lobes, /, more rounded; appendage of ninth sternite,
s, distinctive, obtuse, outer lobe with abundant, very conspicuous,
long, yellow setae.

Habitat. — California.

Sept. 1973 ALEXANDER: NEW CRANE FLIES 193

HoLOTYPE, cT, Santa Cruz Island, Channel Islands, 3% mi. to
Devil's peak, % mi. to U.S.G.S. 1941, woodland, 4 April 1970 (Eric
E. Remington). Allotopotype, ?, 25 March 1970. Paratype, 9,
Santa Cruz Island, Canada del Medio, at University of California
Research Station, 18 April 1970 (Eric E. Remington).

The most similar species are Tipula {Trichotipula) beatula
Osten Sacken and T. (T.) capistrano Alexander, all differing among
themselves in hypopygial structure, especially the tergite and ap-
pendage of the ninth sternite, as discussed and figured.

In the California Bulletin concerning the crane flies, cited under
the preceding species, there are only a few species recorded from the
Channel Islands and several others were added in the collections
made by the Remingtons, now preserved in the Peabody Museum.
The determined species are here listed: Nephrotoma wulpiana (Berg-
roth), Tipula {Bellardina) schizomera Alexander, Tipula (Tripli-
citipula) occidentalis Doane, Tipula (Triplicitipula) planicornis
Doane, Tipula {Triplicitipula) pubera hoew, Tipula {Y amatotipula)
fulvilineata Doane, Limonia (Dicranomyia) defuncta concinna (Wil-
Uston), Limonia {Idioglochina) marmorata (Osten Sacken), and
Erioptera {Trimicra) pilipes (Fabricius).

Tipula (Lunatipula) hastingsae diperona, n. subsp.

Male. — Length about 23 mm; wing 25 mm; antenna about
5 mm.

Female. — Length about 23 mm; wing 20 mm. x

General coloration much as in typical hastingsae Alexander
(Bulletin Brooklyn Ent. Soc, 46:87-89; 1951), described from sev-
eral stations in California, the detailed distribution being given in
the Crane flies of California, as cited earlier, pages 43-44.

Size larger (male 25 mm; in hastingsae, 16-19 mm). Body and
wing coloration generally similar in both subspecies. Chief differ-
ences found in the structure of the male hypopygium, especially the
phallosome, as shown in the figures {hastingsae, Fig. 6 above;
diperona, Figure 6 below) . Gonapophyses, g, in the present fly with
a powerful lateral spine at near midlength, lacking in hastingsae;
aedeagus, a, with posterior spine at apex of organ very long and
slender.

Habitat. — California.

Holotype, cf , Santa Cruz Island, Channel Islands, at University
of California Research Station, Canada del Medio, 18 April 1970
(Eric E. Remington). Allotopotype, 9 , 15 April 1970; Alexander
Collection.

Tipula {Pterelachisus) macleani, n. sp.

General coloration of head and thorax gray, praescutum me-
dially patterned with darker; antennae uniformly black; legs with
femora brownish yellow, tips narrowly black, claws toothed; wings
subhyaline, restrictedly patterned with very pale brown; abdomen

194 GREAT BASIN NATURALIST Vol. 33, No. 3

dark brown, in cases tergites 2-4 reddened; male hypopygium with
central outer portion of tergite depressed to form a shallow yellow
saucer, posterior margin with two approximated triangular points;
eighth stemite with posterior border truncate, with inconspicuous
pale setae.

Male. — Length about 12.5-13 mm; wing 12.5-13 mm; antenna
about 4 mm.

Female. — Length about 17 mm; wing 12.5 mm; antenna about
3.5 mm.

Frontal prolongation of head black, gray pruinose; nasus con-
spicuous, with a terminal tuft of light yellow setae; palpi black.
Antennae black throughout, scape sparsely pruinose; flagellar seg-
ments with small basal enlargements, exceeding verticils in length.
Head light gray; anterior vertex more elevated, low, slightly in-
fuscated; dorsal setae short, black, genae with long light yellow setae.

Pronotal scutum gray, sides of scutellum light yellow. Mesonotal
praescutum light gray, with two intermediate brown stripes that
narrow strongly behind, separated by a vaguely differentiated
ground vitta, lateral stripes very pale to scarcely evident; median
part of transverse suture dark brown; posterior sclerites of notum
clear light gray; scutum with a narrow impressed median line, lobes
slightly variegated with very pale brown; scutellum light gray with
a vague central darkening, parascutella darkened anteriorly, behind
narrowly light yellow; postnotum uniformly clear light gray. Pleura
light gray, dorsopleural membrane restrictedly obscure yellow. Hal-
teres with stem obscure yellow, brighter basally, knob small, brown,
apex obscure yellow. Legs with coxae and trochanters light gray;
femora obscure brownish yellow, tips narrowly black; tibiae light
brown, tarsi darker brown; claw with a small acute point at near
midlength. Wings (Fig. 2) subhyaline, very restrictedly patterned
with pale brown, oval stigma darker; outer two-thirds of cell M
chiefly very pale brown, this with a whitened area at near mid-
length; veins brown. Trichia on certain veins beyond cord, lacking
on most of /?i+2, Rs, M^ and 1st A except at base, vein 2nd A with
extensive trichia. In paratype specimen, veins that comprise long
cell ist M2, with the exception of M3+4, very pale to subobsolete, in
the holotype these elements normally darkened. Venation: Petiole of
cell Ml about one-half m; distal section of Cui strongly recurved at
outer end narrowing the cell.

Abdomen of holotype with basal segment and outer five tergites
dark brown, their posterior borders very narrowly yellowed, tergites
2-4 reddened, grayish sublaterally; all sternites dark brownish gray,
posterior borders very narrowly yellow. Paratype male with ab-
domen uniformly dark brown. Female with tergites 2-7 chiefly
reddened, outer segments darkened, sternites as in the male, the
extreme bases of intermediate segments yellowed. Ovipositor with
cerci long, straight, very slender; hypovalvae stouter, proximal half
blackened, apices rounded. Male hypopygium (Fig. 7) with ninth
tergite, t, dark brown, posterior median portion slightly depressed to

Sept. 1973 ALEXANDER: NEW CRANE FLIES 195

form a flattened yellow saucer, margin produced into two low sub-
median triangular points; setae of tergite pale, on darkened parts
from tiny yellow punctures, vestiture of saucer small and pale, very
inconspicuous. Dististyles, d, as shown; beak of inner style long and
slender, blackened. Eighth sternite, 8s, with posterior border trun-
cate, vestiture pale, discal setae small, sparse marginal bristles only
slightly longer.

Habitat. — Alaska.

HoLOTYPE, cT, Prudhoe Bay, 70° 18' N. Lat., 148° 21' W. Long.,
altitude 15 m, 24 June 1971 (Stephen F. MacLean); Field Catalogue
No. 162; University of Kansas Collection. Allotopotype, $ , with
the type, 13 July 1971; No. 338. Paratopotype, cT, July 1971; No.
228; Alexander Collection.

The species is named for the collector, Stephen F. MacLean, of
the United States Biome Tundra Program. In its general appearance,
including the hypopygium, the fly suggests various regional species
of the subgenus Vestiplex Bezzi, especially Tipula {Vestiplex) ber-
grothiana Alexander (Fig. 8) and T. {V.) aldrichiana Alexander
(Fig. 9), both from Alaska. The drawings of the male hypopygia of
these two flies are based on the type specimens preserved in the
United States National Museum. Other less similar Arctic American
species include Tipula hewitti Alexander, T. johanseni Alexander,
and T. subpolaris Alexander, of the Canadian Northwest Territory,
and T. katmaiensis Alexander. From the structure of the ovipositor
it appears that the present fly is correctly placed in the subgenus
Pterelachisus Rondani.

Limnophila byersi, n. sp.

General coloration of body brownish gray, head light gray, an-
terior vertex broad; antennae of male approximately one-third
length of body; legs with femora yellow, tips brownish black, tarsi
black; wings brownish yellow, stigma oval, dark brown; cell R2
at margin slightly less than one-half cell R^; cell M^ very small, from
about one-fifth to one-sixth its petiole; male hypopygium with outer
margin of basistyle with numerous very long setae, lobe of mesal
face oval; outer dististyle narrowed on outer fourth, terminating in
two small approximated points; phallosome with gonapophyses ap-
pearing as simple slender sickles, outwardly narrowed to an acute
spine; aedeagus simple, long and slender, straight, nearly three times
the apophyses.

Male. — Length about 9 mm; wing 7.8 mm; antenna about 3.2
mm.

Female. — - Length about 12 mm; wing 9 mm.

Rostrum short and broad, transversely oval, dark brown, sparsely
pruinose, with long whitish setae; palpi dark brown. Antennae of
male (Fig. 11) long, nearly one-third the body; scape and pedicel
dark brown, flagellum brownish black; flagellar segments with
short dense pale setae and sparse longer bristles; intermediate fla-

196 GREAT BASIN NATURALIST Vol. 33, No. 3

gellar segments about 4-5 times as long as broad, verticils shorter
than the segments; outer segments about three times as long as
broad, subequal to their longest verticils; outer segment oval. Head
light gray; anterior vertex very broad, about three times exposed
diameter of eyes.

Pronotum brown, sparsely pruinose. Mesonotum almost uni-
formly brownish black, pruinose, praescutal stripes not differen-
tiated; a single median black pit on praescutum, pseudosutural
foveae larger, black; parascutella obscure yellow. Pleura gray;
dorsopleural membrane obscure yellow. Halteres with stem yellow,
knob small, weakly darkened. Legs with coxae and trochanters yel-
low; femora yellow, tips brownish black, on foreleg including about
one-third the length, on posterior pair about one-fifth; tibiae brown-
ish yellow, tips very narrowly darkened; tarsi black. Wings (Fig. 3)
brownish yellow, base and costal region clearer yellow; stigma oval,
dark brown; veins brown. Longitudinal veins beyond cord with
black trichia, lacking on veins comprising cell 1st Mo; further
trichia on Rs, outer third of basal section of Cui, and on most of
2nd A. Venation: cell R, at margin slightly less than one-half cell
Rz; cell Ml very small, about one-fifth to one-sixth its petiole; m-cu
shortly beyond one-third M3+4.

Abdomen brownish black, sparsely pruinose, including the hy-
popygium. Ovipositor with valves stout, cerci obscure yellow, hypo-
valvae black. Male hypopygium (Fig. 10) with basistyle, b, stout,
outer margin thickened, with numerous very long setae, longest
about one-half outer dististyle; cephalic end of mesal face with an
oval lobe provided with long pale setae. Outer dististyle, d, glabrous,
gently curved, outer fourth narrowed, terminating in two small ap-
proximated points, axial one stouter; inner style with base stout,
with conspicuous setae, outer half narrowed, setae microscopic.
Phallosome with gonapophyses, g, appearing as slender simple curved
sickles, narrowed gradually into an acute spine; aedeagus simple,
very long and slender.

Habitat. — Alaska.

HoLOTYPE, cT, Prudhoe Bay, 70° 18' N. Lat., 148° 21' W. Long.,
altitude 15 m, June 1971 (Stephen F. MacLean); No. 345. Alloto-
POTYPE, 9 , with type; No. 298. Paratopotype, cT, with types.

The species is named for my long time friend and fellow student
of the Tipulidae, Dr. George W. Byers, of the University of Kansas.
The closest regional ally is Limnophilo poctica Osten Sacken, widely
distributed in northern North America, from Massachusetts to British
Columbia, thence north to Alaska. This is readily told from the
present fly by the yellow body coloration, longer antennae, vena-
tion, especially the normal cell M,, and in details of the male
hypopygium, including the basistyle, and the gonapophyses and
aedeagus of the phallosome. The microscopically bidentate apex of
the outer dististyle is found in both species.

NE ARCTIC DESERT DECTICIDAE (ORTHOPTERA)

PART III THE TRUE TYMPANUM IN CERTAIN

GENERA WI FH KEY

Ernest R. Tinkhami

This is a report on the nature of the true tympanum in Decticidae,
its location and taxonomic value. The decticids are a most difficult
group to evaluate generically, and any new character that will aid
in defining and keying out genera should be of definite taxonomic
value in addition to adding to our knowledge on sound reception in
these most interesting creatures.

Apparently no recognition has been given in the literature to the
location and character of the true tympanum in the Decticidae or
related Tettigonoidea or in the Oecanthidae or other orthopteroid
families. Textbooks have for many decades illustrated and de-
scribed the tympanum as located in the basal portion of the pro-
tibia where there is a small, hollow, swollen area with internal and
external longitudinal slots that are supposed to permit the entrance
of sound vibrations. All members of the Tettigonoidea that stridu-
late are supposed to possess this protibial organ.

Fulton (1928) conducted experiments on certain tettigoniids, in-
cluding Neoconocephalus nebrascensis, Amblycorypha rotundifolia
brachyptera, and Oecanthus niveus, in which half of his caged speci-
mens had their fore tibiae amputated at their bases; the checks were
normal. His experiments proved that those with amputated protibiae
could still hear but that an asynchronization had developed, while
in the control or untreated cages synchronization of stridulation to
distant stridulating males proceeded normally. These experiments
indicated that tettigoniids and oecanthids with snipped-off fore
tibiae could still hear, but the obvious meaning was obscured. Fulton
summarized his conclusions thus: "The tympanal organs of the front
tibiae are auditory organs, an assumption based on circumstantial
evidence of their structure and the fact that they are present in all
stridulating species."

It is obvious that such an assumption can be erroneous. It is like
amputating the arms but not observing the ears which hear. Cer-
tainly, decticids, tettigoniids, oecanthids, and others can still hear
sound, even with their front legs cut off, because the tympana are
present on the thoraces of these creatures.

Snodgrass (1925) states that "if they [protibial organs] are not
ears what are they?"

Such assumptions and conclusions can explain the oversight in
recognizing the true tympanum. Obviously, the protibial organ in
the decticids, tettigoniids, and oecanthids, as shown by Dr. Fulton's
experiments, must be a substation or synchronometer that interprets
and passes on the sound received by the true tympanum, herein de-

'81-441 Date Palm Avenue, Indio, California 92201,

197

198 GREAT BASIN NATURALIST Vol. 33, No. 3

scribed as located on the thorax, and flashes on a response to the teg-
mina, which stridulate their challenge to the producer of the re-
ceived sound or song.

That the organ which lies just caudad of the prothoracic spiracle
is the tympanum is further corroborated by the position of the tym-
panum in all those sound-producing acridids in many subfamilies of
grasshoppers, where its location is just caudad of the metathoracic
spiracle and only two segments removed from that in the Decticidae
and just above the metacoxal joint of the hind leg.

It is interesting to note that the two new genera recently described
by the author, namely, Petropedes and Platyoplus, portray the great-
est development of the tympanum, but nature exhibits two different
ways to increase the receptivity of sound. In Platyoplus (Figure 3)
though the pronotum is very broad and very shallow, it still hides
half of the very large, auricular-shaped tympanum. To overcome
this effect, the pronotum just above the covered upper portion of the
tympanum at the posteroventral emargination of the lateral lobes is
convexly swollen to facilitate the penetration or reception of sound.
On the other hand, in Petropedes (Figure 4) although the lateral
lobes are very deep, the posteroventral margin is more strongly
emarginate or excised than in any other eremicolous decticid genus,
so that the huge tympanum is almost completely exposed for the
reception of sound.

Although both genera possess a very large tympanum, relation-
ship stops at this point; for the two genera represent two different
sections of the Decticidae. Platyoplus apparently has its nearest
relationships with Ateloplus, especially the largest species A. splen-
didus, whereas Petropedes seems allied to Inyodectes and perhaps
Eremopedes and Pediodectes as well.

Likewise, it is interesting to note that both recently described
genera inhabit barren, inhospitable rocky mountain slopes under a
blazing sun, and the enormous tympana would appear to be adap-
tions to such an environment. Inyodectes (Figure 7) seems to bear
out this observation, for it also has a relatively large tympanum in
relationship to its medium size. Although it does not dwell in such
a hot desert as Petropedes and Platyoplus, which are both members
of the fauna of the hottest desert, the Colorado, it does dwell in piles
of rocks at the base of talus slopes in the western portions of West-
guard Pass at about the lower limits of the pines in the Inyo Moun-
tains, which likewise, because they lie in the rainshadow of the
vSierra Nevadas, are very barren and xeric.

In Ateloplus (Figure 1), a review of the known species shows
that the tympanum is oval in shaj)e and not as large or expanded in
its upper half as in Platyoplus. In Ateloplus, the species A. schwarzi
and A. notatus show tympana that are roundly oval in shape;
whereas in A. hesperus, A. luteus, A. minor, and A. splendidus, the
tympana are more narrowly elliptical in relationship to their size.

In Eremopedes (Figures 2, 10), the tympana range from a nar-
row elliptical slit in E. shrevei, E. balli, E. covilleae, E. ephippiata.

Sept. 1973

TINKHAM: DESERT DECTIDIDAE

199

eye /. zr/P ~r/>^ i- ^

Figs. 1-10. Decticid pronota as seen in lateral aspect and cross-sections
from cephalic aspect: 1, Ateloplus splendidus, male topotype; 2, Eremopedes
ephippiatus sonorensis, male holotype; 3, Platyoplus gilaensis, holotype; 4, Petro-
pedes santarosa, holotype; 5, Capnobotes fuliginosus, male, Mulligan Canyon,
Franklin Mts., Texas; 6, Oreopedes cryptoptera, female, Westguard Pass; 7, Inyo-
dectes pallidas, female topotype, Westguard Pass; 8, Zacycloptera atripennis,
topotype, Walker Lake, Nevada; 9, Plagiostira utahensis, female, western Utah;
10, Eremopedes bilineatus, male, Santa Ana, Sonora, Mexico.

200 GREAT BASIN NATURALIST Vol. 33, No. 3

and E. pallidas to a narrowly oval form in E. bilineatus, E. scudderi,
and E. ephippiata sonorensis.

In most of the other genera, such as Oreopedes (Figure 6), Cap-
nobotes (Figure 5), Zacycloptera (Figure 8), Plagiostira (Figure 9),
Neduba, and Aglaothorax, the tympana are rather oval elliptical,
and some of these, especially Zacycloptera, Plagiostira, and Cap-
nobotes, have the fore margin of the tympanum quite irregular due
to the presence of the prothoracic spiracle.

Another feature of the tympanum that should be noted here is
whether its periphery is bare or lined with an even row of very fine,
short-tapered hairs. In Platyoplus, Petropedes, Inyodectes, Capno-
botes, and Anoplodusa, the peripheral margin is barren of any mi-
nute hairs; in Ateloplus, the margin is very finely hirsute in the
smaller species but bare in A. splendidus, the largest species; in
Oreopedes, the margin is very finely hirsute; in Eremopedes, it is
barren in some species and hirsute in others. The function of these
hairs is not known or surmised at this time; a study of their function
and structure would be most interesting and perhaps revealing.

Key to Certain Eremicolous Genera Based on
Tympana and Pronota

1. Tympanum extremely large, located at or near sinuation

of posteroventral margin of lateral lobes of pronotum 2

Tympana mostly medium to small in size 3

2. Tympanum enormous, broadly oval, and largely exposed

by strongly excavate posteroventral emargination of

pronotum, although upper quarter still concealed

Petropedes

Tympanum very large, auricular in outline, largely con-
cealed by very broad, very shallow lateral lobes of pro-
notum; pronotum convexly swollen above concealed
portion of tympanum Platyoplus

3. Pronotum barrel-shaped, without lateral carina 7

Pronotum with partial or complete rounded lateral carinae .... 4

4. Pronotum extremely large, ovally rounded in dorsal out-

line, its dorsal surface gently convex 5

Pronotum not as above, more narrowly linear, sometimes

quadrate in dorsal outline; dorsal surface irregular 6

5. Pronotum extremely large, oval in dorsal outline, highly

colored Aglaothorax

Pronotum moderately large, colored more drably Neduba

6. Size very large; tegmina and wings far surpassing apex

of abdomen; metazona with strongly arched lateral

ridge or keel Capnobotes

Sept. 1973 TINKHAM: DESERT DECTIDIDAE 201

Size large, tegmina and wings slightly longer than pro-
notum; lateral carina of pronotum crenulate-arcuate;
disc of pronotum irregular Plagiostira

7. Size very small Oreopedes

Size medium to larger 8

8. Body large and heavy; wings jet black; tegmina and

wings slightly longer than pronotum Zacycloptera

Body much more slender, wings not black; tegmina and

wings shorter than pronotum 9

9. Tympanum relatively large for medium size and half

exposed Inyodectes

Tympanum small, usually oval elliptical; size ranging

from small to large - 10

10. Size mostly medium to large, mostly heavy bodied; pro-
notum with deep lateral lobes Eremopedes

Size medium large to small; pronotum broad, with disc

convex and with rather shallow lateral lobes Ateloplus

References

Fulton, B. B. 1928. A demonstration of the location of auditory organs in

certain Orthoptera. Ann. Ent. Soc. Amer. 31:445-448.
Snodgrass, R. E. 1925. Insect musicians, their music and their instruments.

Ann. Rpt. Smithsonian Inst. 1923:405-452.
TiNKHAM, E. R. 1972. Nearctic desert Decticidae (Orthoptera). Part I. A new

genus from California. Great Basin Nat. 32:176-179.
. 1973. Nearctic desert Decticidae (Orthoptera). Part II. A new genus

and species from Aiizona. Great Basin Nat. 33:43-50.

NOTES ON REPRODUCTION IN LAMPROPELTIS
TRIANGULUM AND COLUBER CONSTRICTOR IN UTAH

William L. Grogan^ and Lloyd C. Pack, Jr.^

A search of the hterature has revealed no published records of
egg laying in Lampropeltis triangulum taylori Tanner and Loomis.
On 8 July 1967 a female with a snout- vent length of 494 mm was
collected in Hobble Creek Canyon, 7 mi. east of Springville, Utah.
On 20 July four eggs were laid. After four weeks of incubation the
eggs were attacked by fungi and were discarded.

Van de Velde, Martan, and Risley (1962) first described the
hatching of eggs of Coluber constrictor mormon Baird and Girard.
Since then no other hatchings have been recorded. On 12 July 1967
one male and three gravid females were collected at the same locality
as above. Clutches of seven eggs were laid on 21 July, six on 24
July, and six on 28 July. The eggs were placed on moist sand in a
jar for incubation. The last laid clutch was discarded because of
fungi. Two eggs of that clutch were dissected and found to contain
embryos. Two eggs of the first clutch hatched on 26 September and
five on 27 September. Three eggs of the second clutch hatched on
1 October and one on 3 October. The remaining two eggs in this
clutch did not hatch. The time of hatching varied from 67 to 70
days, with eight of the eleven hatching in 68 days.

The color pattern of our specimens was similar to that described
by Van de Velde, Martan, and Risley (1962); however, there were
a few important differences. The dorsal saddles ranged from 46 to
51 as compared with their 69. The chin, throat, infralabials, and
supralabials are a vivid white with dark reddish brown spots on the
posterior margins of the supralabials and the last infralabial. The
color of the head also differs in that the ground color is light gray
brown with darker brown spots on the prefrontals, supraoculars,
frontal and parietals.

Four of the hatchlings are preserved in collections of Brigham
Young University (BYU 30808-30811). Their respective snout- vent
and total lengths in mm are: 192-263, 222-284, 202-280, 201-274.

We are indebted to Wilmer W. Tanner and Dorald M. AUred
for criticism of the manuscript.

Literature Cited

Van de Velde, R. L., J. Mart.'VN, and P. L. Risley. 1962. Eggs and hatchlings
of the snake Coluber constrictor mormon from Oregon. Copeia 1962(1),
212-13.

'Department of Zoology, Brigham Yoimg University, now at Departniciil of Entomology, Univer-
sity of Maryland, College Park, Maryland 20742.

^Department of Zoology, Brigham Young University, Provo, Utah 84602.

202

COURTSHIP BEHAVIOR AMONG WHITE-TAILED
AND BLACK-TAILED JACKRABBITS

Del F. Blackburni

This paper represents a portion of a research project on behavior
of black-tailed and white-tailed jackrabbits, Lepus calif ornicus and
L. townsendii, of the Castle Rock area of Eastern Oregon. The full
study is described elsewhere (Blackburn, 1968).

The observations made of jackrabbit courtship behavior from
April through July, 1967-68 were quite similar to those described by
Severaid (1941), Janson (1946), Lechleitner (1959), Haskell and
Reynolds ( 1 947 ) , and Pentrelli ( 1 968 ) .

I observed both white-tailed and black-tailed jackrabbits in vary-
ing habitat and behavioral situations. This provided the opportunity
to extend many of the reports just mentioned.

Both Pentrelli and Lechleitner reported black-tailed jackrabbits
as having very intense courtship behavior involving circling, male
and female approaches, and long chases. In this study urine emission
was observed during the jumping and chasing activity. This complex
behavior involving combinations of all the above listed components
was observed to last from 5 to 20 minutes. Copulation usually fol-
lowed. After copulation the male jackrabbit was observed on several
occasions to jump slightly backwards, fall to the ground, emit a
hissing squeal and leap up again renewing the chase with subsequent
copulations of up to four times.

Although only a few observations of courtship among white-
tailed jackrabbits were possible in this study, their courtship behavior
was observed to be basically the same as that of black-tailed jack-
rabbits, with the exception that jumping behavior was more pro-
nounced.

Tinbergen (1952) states that displacement behavior is caused or
motivated by a surplus of stimulus or drive. A jackrabbit, highly
excited during escape or reproductive behavior, may exhibit this type
of activity. I observed five male black-tailed jackrabbits in pursuit
of one female. After sighting me, two of the males left the pack and
began feeding. They fed vigorously and did not attempt to leave the
area upon my approach. Shortly afterwards, two other males aban-
doned the female and began attempting to copulate with each other.
Upon collection, both had their penises fully extended. Both of these
behavior types suggest a displacement type of behavior. A single
remaining male was observed to copulate twice with the female. The
entire area surrounding this field of activity was covered with hair,
the presence of which suggested fighting.

I thank the Bureau of Land Management, Vale, Oregon, for their
support of this study. It was conducted under the guidance of the
Department of Zoology, University of Idaho, Moscow, Idaho.

^Biology Department, Clark College, Vancouver, Washington.

203

204 GREAT BASIN NATURALIST Vol. 33, No. 3

Literature Cited

Blackburn, D. F. 1968. Unpublished M.S. Thesis, University of Idaho, Mos-
cow, Idaho.

Haskell, H. S., and Reynolds, H. G. 1947. Growth, developmental food re-
quirements, and breeding activity of California jackrabbits. Journ. Mammal.
28:129-136.

Janson, R. G. 1946. Survey of the native rabbits of Utah, with reference to
their classification, distribution, their history, and ecology. M.S. Thesis,
Utah State University.

Lechleitner, R. R. 1959. Sex ratio, age classes and reproduction of the black-
tailed jackrabbit. Journ. Mammal. 40:63-81.

Pentrelli, M. J. 1968. Mating behavior of the black-tailed jackrabbits. Journ.
Mammal. 49:785-786.

Severaid, J. H. 1941. The life history and artificial propagation of the snow-
shoe hare, Lepus arnericanus strutherus bangs. M.S. Thesis, University of
Maine.

Tinbergen, N. 1952. Derived activities; their causation, biological significance,
origin and emancipation during evolution. Quart. Rev. Biol. 27:1-32.

GREAT BASIN NATURALIST

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Volume 33 December 31, 1973 No. 4

LOCAL DISTRIBUTION AND

INTERSPECIES INTERACTIONS IN MICROTINES,

GRAND TETON NATIONAL PARK, WYOMING

Tim W. Clark'

Abstract. — Some ecological relationships and interspecies interactions (i.e.,
habitat, foods, and reproduction) between Microtus pennsylvanicus pullatus, M.
montanus nanus, M. longicaudus mordex, and Clethrionomys gapperi galei were
investigated in Grand Teton National Park, Wyoming. Trapping was conducted
from June-July, 1968 and May-July, 1969 yielding 110 M.p.p., 171 M.m.n.,
17 M.l.n. and 41 C.g.g. Six plant communities were defined structurally; all
communities contained voles but varied considerably in numbers of species and
individuals. Analyses of stomach contents showed similar diets. Embryo counts
indicated that there was no significant interspecies differences in litter sizes.
Females of all species were pregnant in about equal proportions. Almost all
adult males showed descended testes and were reproductively active. Testicular
and seminal vesicle weights and lengths fluctuated. M.p.p. mean body weights
were largest: M44.7g (N = 31). F38.1g (N = 29); this was nearly twice the
mean weights of C.g.g.: M20.4g (N = 7), F23.3g (N = 3). Weights for M.m.n.
and M.l.m. fell between these extremes and were similar to each other. Trends
in differential habitat use were clearly demonstrated and evidence suggests the
four microtines are at least partially incompatible.

Four common rodents in Grand Teton National Park are:
Microtus pennsylvanicus pullatus (meadow vole), M. montanus
nanus (mountain vole), M. longicaudus mordex (long-tailed vole),
and Clethrionomys gapperi galei (red-backed vole). Except for
Findley (1951 and 1954), Negus and Findley (1959), and Stoecker
(1970), little is known on distributions and ecology of these four
species in the park. It is well documented that population levels
are directly linked with relative numbers of other species in the
community (Wirtz and Pearson, 1960; Curry-Lindahl, 1959;
DeLong, 1966; Lidicker, 1966; Whitaker, 1967; Batzli, 1968; Shure,
1970). This paper further elucidates some ecological relationships
and interspecies interactions (i.e., habitat, food habits, and repro-
ductive characteristics) between these microtines in Grand Teton
National Park, Wyoming.

Methods

The study was conducted in June and July, 1968, and May
through July, 1969. A total of 339 specimens were collected: 171

^Department of Zoology, University of Wisconsin, Madison 5370fi.

205

206

GREAT BASIN NATURALIST

Vol. 33, No. 4

Microtus montanus, 110 M. pennsylvanicus, 17 M. longicaudus,
and 41 Clethrionomys gapperi. Sherman live traps and snap traps
were used, and each trapline (1 trap every 10 meters) was left
for 3 to 6 days, then moved to a new site. Traplines ranged from
25 to 50 traps each. For each Sherman trapline set, two snap trap-
lines were set in each community. Thus the relative proportion of
live traps to snap traps was uniform between communities.

Table 1 summarizes the six lowland plant communities trapped
to determine habitat affinities. More detailed descriptions are given
by Clark (1971) and Reed (1952).

Food uses were ascertained by stomach content examination.
Contents were identified and a volumetric estimate made to the
nearest 5 percent for each item. The mean volumetric estimates
and frequencies of occurrence of each item were calculated (Clark,
1968).

In the field, all males were checked for position of testes; fe-
males were examined for condition of mammae (relative prom-
inence and possible lactation). In the laboratory, male reproductive
systems were removed; testes and seminal vesicles were examined,
weighed, and measured. In the females, the number of embryos
was counted.

Table 1. Comparison of plant communities sampled for microtines in Grand
Teton National Park, Wyoming.

Prominent

Plant Community

Species

Characteristics

Sedge-Meadow

Car ex spp.

Underlain by firm sod; often cov-

Aleopecurus

ered with several cm of water; no

aequalis

woody plants, only sedges, grasses,

Hordeum

and forbs

bachy anther um

Sedge-Grass

Carex spp.

Slightly elevated ground; very pro-

Meadow

Agropyron

ductive, 30 spp. of forbs; no woody

trachycaulum

plants, only sedges, grasses, and

Bromus, Poa,

forbs; rarely covered by water

Agrostis

Shrub-Swamp

Salix sp.

Water table near surface; stable;

Bromus, Poa,

reduced light intensity under

Calamagrostis,

shrubs along with increased mois-

Aster, Aconitum,

ture; woody plants-shrubs, sedges.

Castelleja.

grasses, and forbs; shrubs over 50

Ranunculus

percent ground cover

Shrub-Sedge-

Same as (3) above

Shrubs less than 50 percent ground

Grass Savanna

cover

Aspen

Populus, Aster,

More level sites adjacent to

Helianthella,

swamps and meadows; closed tree

Lupinus

canopy, dense shade; soil moist in
spring but diminishes in summer

Big Sagebrush

Artemisia, spp..

Most widespread and conspicuous

Purshia

community; shrubs, forbs, and

Stipa, Poa,

grasses; very dry, little to no wa-

Bromus, Aster,

ter

Eriogonum,

Potentilla

Dec. 1973 clark: Wyoming rodents 207

Measurements of tail, hind foot, and total lengths and body
weights were taken on all specimens. All M. pennsylvanicus and
M. montanus were identified on the basis of molar structure. Rep-
resentative specimens were prepared as museum skins and deposited
in the Jackson Hole Biological Research Station collection and in
the Museum of Natural History, University of Wisconsin-Stevens
Point.

Results

Habitat affinities. — Habitat affinities were based on the plant
communities in which capture per species was greatest (Maxell and
Brown, 1968). All communities contained voles but varied con-
siderably in numbers of species and individuals (Figure 1). Two
communities (Sedge-Meadow and Sedge-Grass Meadow) were ex-
clusively occupied by M. montanus, while the other four com-
munities showed multispecies occupancy. The Big Sagebrush Com-
munity yielded only two species (M. montanus and M. pennsyl-
vanicus), the Shrub-Swamp Community three species (all but M.
longicaudus) , and the Shrub-Sedge-Grass Savanna and Aspen Com-
munities all four microtine species.

M. montanus was found in all six plant communities, its popu-
lation numbers exceeding M. pennsylvanicus (the second most
abundant species) in all communities except the Shrub-Swamp,
where M. pennsylvanicus greatly outnumbered other voles. M.
pennsylvanicus showed the second widest distribution in four com-
munities. Clethrionomys gapperi occupied three plant communities
and M. longicaudus only two (Figure 1 ) .

Five of the six plant communities (excluding Big Sagebrush)
are in successional sequence on the wetter sites in Jackson Hole
(Reed, 1952). Some general characteristics of these communities
in relation to vole distribution are given in Figure 1. The closer
the plant community to climax, the greater the vole species diver-
sity [species diversity refers to the number of species in each com-
munity (M'Closkey, 1972)}; correlated to this is a trend toward
mesic soil-moisture conditions and more complex plant community
structure (i.e., potentially more niches). The Aspen Community,
the highest serai community of the sequence, possessing mesic soil-
moisture conditions and three vegetative strata (herbs, shrubs, and
trees), contained all four microtines. The simpler the community
structurally — by virtue of its early successional position (Sedge-
Meadow or Sedge-Grass Meadow) or limited moisture (Big Sage-
brush)— the smaller the species diversity, one and two species,
respectively.

However, the greatest species diversity was not correlated with
greatest total populations numbers. The Shrub-Swamp Community
is characterized by only two vegetational strata (herbs and shrubs),
a 50-100 percent ground cover of shrubs, and some areas inundated
by several cm of standing water; it is located near the middle of
the successional sequence. It was this community that showed the
greatest absolute numbers of voles (N=117) and the greatest CI,

208

GREAT BASIN NATURALIST

Vol. 33, No. 4

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Dec. 1973

CLARK: WYOMING RODENTS

209

with almost twice the total catch as the second highest community
(Nrr60). Nearly all voles in this community were M. pennsylvan-
icus. Population numbers were more or less comparable among the
Sedge-Meadow, Sedge-Grass Meadow, Shrub-Sedge-Grass Savanna,
and Aspen Communities, with a CI (captures per 100 trap days)
of .031, .037, .027, and .025, respectively. The Big Sagebrush Com-
munity, a dry community, showed the lowest total population
number (N=13) and a CI of only .008, over 90 percent lower than
the most populous community (Shrub- Swamp, with Nr=117).

In the two communities where M. montanus was most abundant,
other species were absent. The community where M. montanus was
almost totally absent was the community showing the largest
numbers of both M. pennsylvanicus and C. gapperi. The C. gapperi
were situated on drier sites in peripheral areas of this community,
and M. pennsylvanicus tended to occupy wetter areas. The driest
community yielded two species, with M. montanus outnumbering
M. pennsylvanicus seven to one. In the two communities containing
all four microtines, the numbers of all species were considerably
less than in communities that were occupied by only one, two, or
three species. In all six communities, for every 10 M. montanus
taken, 7.5 M. pennsylvanicus, 2.5 C. gapperi, and 1 M. longicaudus
were captured.

Stomach contents. — Among the more obvious possibilities for
competition between species is maintenance of adequate levels of
energy and nutrition for meeting metabolic requirements. Analyses
of stomach contents showed a similar diet in the four microtines
(Table 2). Green plant materials comprised the major identifiable
items in all stomachs; seed fragments were the other important
food. The herbivorous habits of all species suggest potential competi-
tion for food, at least during seasons when food may be scarce.
During the months of this study, food was abundant and did not
seem to influence distributions.

Microtus montanus and M. pennsylvanicus, the two species
probably most directly in competition, showed comparable levels of

Table 2. Comparative stomach contents of some microtines in Grand Teton
National Park, Wyoming. Figures are mean volumetric estimates calculated to
the nearest 1 percent; figures in parentheses are percent frequencies of occur-
rence; T indicates trace.

Microtus Microtus Microtus Clethrionomys

Stomach montanus pennsylvanicus longicaudus gapperi

Contents (N = 139) (N=107) (N=14) (N = 25)

Green plant

materials

63(80)

94(80)

38(55)

59(76)

Seed fragments

8(13)

1(9)

1(1)

9(19)

Arthropods

T(l)

Parasitic

roundworms

4(8)

2(6)

Hair

6(18)

Unidentified

25(100)

3(100)

61(100)

26(100)

210 GREAT BASIN NATURALIST Vol. 33, No. 4

parasitic roundworm infestations. Both M. longicaudus and Cleth-
rionomys gapperi lacked worms. The infested voles otherwise
looked healthy; the parasitic load did not seem to be responsible
for any observable mortality.

Reproduction. — Differential reproduction may greatly en-
hance the competitive advantage of one species over another. The
breeding status of females of the four microtine species is given in
Table 3. Embryo counts indicated that there was no significant
interspecies difference in litter sizes. The percentage of pregnant
adult females in all species was similar except for Clethrionomys
gapperi, which showed a pregnancy rate about one-half that of the
other three. Throughout May, June, and July, females of all
species were pregnant in about equal proportions, indicating con-
tinuous breeding activity of the population during this time.

Almost all adult males of the four species showed descended
testes; males wdth scrotal testes are frequently used as an indication
of population breeding (Packard, 1968). The mean testicular lengths
and weights and mean seminal vesicle lengths and weights varied
over the periods of study for all four species (Tables 4 and 5).
Figure 2 shows testis lengths in millimeters and weights in milli-
grams for the four microtines. A sharp increase in lengths and
weights was evident in Microtus montanus from June 16 to 30, cor-
responding to a slight decrease in the other three species. The re-
lationship was not clear. In the other three microtines, testes param-
eters were more or less stable, showing no such sudden increase
or decrease. Seminal vesicles showed a pattern of weights and
lengths corresponding with that of testicular weights and lengths
(Figure 2) . The mean seminal vesicle parameters for M. montanus
again increased sharply in late June, corresponding to the increase
in testes size. The mean seminal vesicle length in M. longicaudus
increased slightly during the same period (June 16-30), but a con-
comitant increase in seminal vesicle weights was absent. Testicular
and seminal vesicle characteristics in C. gapperi showed no peak of
activity, perhaps indicating a constant level of spermatogenic activity.

Morphological characteristics. — Relative body size may be
significant in determining the outcome of agonistic interactions
and, thus, species distributions. The four microtines varied in mor-
phological characteristics (Table 6). Microtus pennsylvanicus mean
weights were largest: males 44.7 g, females 38.1 g; this was nearly

Table 3. Comparative female reproductive statuses of four microtines. Grand
Teton National Park, Wyoming.

Reproductive

Microtus

Clethrionomys

Characteristic

montanus

pennsylvanicus

longicaudus

gapperi

Total number examined

Number pregnant

Mean number of embryos

6.35

6.59

6.00

Range in embryo number

4-9

5-7

4-7

Standard deviation

±1.29

±1.34

±1.16

±1.19

Dec. 1973

CLARK: WYOMING RODENTS

211

Table 4. Testis weights in mg and testis lengths in mm for four microtine
species in Grand Teton National Park, Wyoming.

Periods

Mean

Range

S.D.

Testis

Lengths

(mm)

Testis

Weights (mg;

)

Mean

Range

S.D.

Microtus montanus

May 15-31

680

526-729

±124

13.0

9-15

±2.86

June 1-15

535

261-676

±101

10.8

9-12

±0.80

June 16-30

1449

850-1818

±420

15.7

13-18

±2.06

July 1-15

535

500-570

± 35

10.7

10-12

±0.50

July 16-31

586

497-621

±321

11.8

9-14

±1.26

Total

Microtus pennsylvanicus

May 15-31

1225

731-1185

±159

15.1

13-17

±1.27

June 1-15

1389

568-1849

±322

15.7

11-18

±1.81

June 16-30

6274

967-1501

±173

14.0

13-16

±1.73

July 1-15

1275

1060-1495

±178

15.0

14-17

±1.22

July 16-31

1190

1136-1238

±192

14.8

13-16

±1.64

Total

Microtus longicaudus

May 15-31

541

436-582

± 89

12.0

11-13

±1.0

June 1-15

679

610-694

± 47

12.0

11-13

±1.0

June 16-30

535

489-569

±110

10.0

9-11

±0.8

July 1-15

587

502-621

±126

10.0

9-11

±0.8

July 16-31

610

—

11.0

—

Total

Clethrionomys gapperi

May 15-31

364

?30-380

± 23

9.7

9-10

+ 0.50

June 1-15

366

287-410

±218

10.0

—

June 16-30

409

362-440

±167

9.2

8-11

±1.09

July 1-15

318

289-354

±148

10.6

10-12

±0.89

July 16-31

392

—

9.3

9-10

±0.57

Total

twice the mean weights of Clethrionomys gapperi: males 20.4 g,
females 23.3 g. Weights for M. montanus and M. longicaudus fell
between these extremes and were similar to each other.

The linear measurements of C. gapperi and M. montanus were
similar, both smaller than the other two microtines. Even though
M. pennsylvanicus had the largest body weight, its length was ex-
ceeded by that of M. longicaudus. M. longicaudus total and tail
lengths are greatly expanded by its disproportionately long tail.

Discussion

Animal distributions are influenced by a constellation of factors
ranging from a complex of abiotic environmental gradients to biotic
regulators including diseases, predators, and intra- and interspecific
competition. The importance of these factors varies from area to
area; it is difficult to dissect out each factor and determine its rela-
tive influence in controlling population distributions and numbers.
This study examined some aspects of the environment as well as
the relationship of other species in the community to distributions
of some microtines. Distribution patterns suggest that some mecha-

212

GREAT BASIN NATURALIST

Vol. 33, No. 4

Table 5. Seminal vesicle weights in nig and lengths in mm for four micro-
tine species in Grand Teton National Park, Wyoming..

Seminal

Vesicle

Lengths

jminal Vesicle Weights
Mean Range

(mg)
S.D.

(mm)

Periods

Mean

Range

S.D.

Microtus montanus

May 15-31

379

286-399

± 69

19.0

14-22

±1.97

June 1-15

359

262-420

± 73

16.3

14-19

±1.38

June 16-30

824

270-1153

± 89

23.7

15-29

±6.18

July 1-15

364

287-435

± 61

17.2

14-20

±2.54

July 16-31

342

268-411

± 89

18.2

14-21

±3.16

Total

Microtus pennsrlvanicus

May 15-31

484

325-760

±118

20.1

17-24

±1.89

June 1-15

733

316-1210

±281

22.1

14-26

±3.45

June 16-30

623

410-875

±179

20.5

17-23

±2.51

July 1-15

936

561-1420

±357

20.5

17-24

±2.89

July 16-31

782

528-926

±286

20.7

17-22

±2.44

Total

Microtus longicaudus

May 15-31

340

289-390

± 51

12.3

11-14

±1.53

June 1-15

205

176-256

± 44

14.0

13-15

±1.00

June 16-30

240

210-314

± 94

17.0

14-18

±2.00

July 1-15

320

286-354

± 72

14.0

13-16

±1.41

July 16-31

280

—

14.8

—

Total

Clethrionomys gapperi

May 15-31

230

100-251

± 62

16.0

14-17

±1.50

June 1-15

235

140-252

± 72

15.0

14-16

±1.00

June 16-30

210

126-246

±117

15.4

15-16

±0.55

July 1-15

279

224-343

± 69

14.6

13-17

±1.52

July 16-31

261

242-289

± 24

16.0

15-17

±1.00

Total

nism of active separation is operative, especially between Microtus
montanus and M. pennsylvanicus. Some of the habitats contained
multiple species, while others only one. In the habitats containing
one species, interspecific relations may be a factor keeping other
species out. In situations where two or more species occur, competi-
tion is probably occurring. As discussed by Whitaker (1967) when
competition is great enough one or both species may tend to reduce
the population of the other species. Then we would expect to find
lower population numbers in areas where both species occur than
in situations where only one species is found. Grant (1970a) con-
cluded from a series of laboratory studies supported by fieldwork
(Grant, 1969, 1970b) that when adults of two species that normally
occupy different habitats in mainland regions meet in the wild they
will interact aggressively and tend to disperse. This habitat segrega-
tion is maintained on islands where both genera (Clethrionomys
and Microtus) are represented by at least one species; but on islands
where only a single species of one of the genera occurs, it often
occupies the habitat of the absent genera (Ota and Jameson, 1961;

Dec. 1973

CLARK: WYOMING RODENTS

213

M. montonus
M. pennsylvanicus
M. longicoudus
C. gopperi

m 100

•

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• • • ■■

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1 1 1

MAY

JUNE

JULY

MAY

JUNE

JULY

15-31

1-15

16-30
PERIODS

1-15

16-31

15-31

1-15

16-30
PERIODS

1-15

16-31

Fig. 2. Changes in testis lengths (mm) and weights (mg) and seminal
vesicle lengths (mm) and weights (mg) for four microtines, Grand Teton Na-
tional Park, Wyoming.

Cameron, 1964; Corbet, 1964). This implies some form of popula-
tion interaction in areas of sympatry.

Ecological segregation in my study could be a result of each
species responding to microenvironmental requirements and may
not be related to other species in the community. Whereas several
of the plant communities sampled in this study are very similar
and since the four microtines are closely related taxonomically, it
is likely that they share some similar ecological requirements;
therefore, distributions exhibited are probably the result of competi-
tion for these requirements. When few or no ecological differences
are present between two or more species occurring together, and

214

GREAT BASIN NATURALIST

Vol. 33, No. 4

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Dec. 1973 clark: Wyoming rodents 215

when species numbers are much lower than expected, the species
may be adversely affecting one another (Whitaker, 1967) as is
suggested by this study.

M. montanus occupied a wide range of plant communities and
seems capable of existing in large numbers in all of them (with
one exception) regardless of the other species jiresent. Competitive
interactions seem to be occurring with M. pcniisylvanicus, resulting
in the restriction or total elimination of M. pcnnsylvanicus from
certain situations. Murie (1969) and Hodgson (1972) showed that
M. pennsylvanicus demonstrated a wet substrate preference while
M. montanus did not. In my study, M. pennsylvanicus occurred
in largest numbers in the wettest situations and was present in
limited numbers or totally absent from areas tending to the drier
end of the spectrum. This concurs with Findley (1951, 1954) who
concluded: (1) that where the range of M. pennsylvanicus overlaps
that of M. montanus, a species well adaj)ted to dry mountain grass-
lands, M. pennsylvanicus is forced to retire to its optinmm niche,
the hydrosere community, and (2) that the distribution of M.
pennsylvanicus, in part at least, is due to competition with closely
related microtines.

The relationships of M. longicaudus and C. papperi are less
clear; M. longicaudus is the most restricted of the four microtines.
It is apparently capable of coexisting with the other three species,
being found in the wetter, more structurally complex plant commu-
nities. A similar tendency was exhibited by C. gapperi, but it is ca-
pable of occupying a slightly wider latitude of situations. Voles of the
genera Clethrionomys and Microtus tend to occupy different habi-
tats, the former woodlands and the latter grasslands (Hall and
Kelson, 1959; Southern, 1964). Evidence that M. pennsylvanicus
and C. gapperi tend to inhibit each other from using the opposite
species habitat is abundant (Grant, 1969; Morris, 1969; Iverson
and Turner, 1972).

Results of my stomach analyses suggest a similarity in food
uses, but microtine populations seldom critically deplete their food
supplies (Negus and Pinter, 1966). The quality, rather than the
quantity, of available food was shown to exert considerable influ-
ence on initiation, duration, and success of the breeding season in
natural populations of .1/. montanus (Negus and Pinter, 1966). A
comparative reproductive investigation showed litter sizes arnong
the four microtines to be similar, with no species having a disad-
vantage in this respect, except possibly C. gapperi, which showed
about one-half the pregnancy rate of the other three microtines.
This low numerical productivity may be offset by a greater sur-
vivorship, or the population may suffer suppression by the superior
numbers produced by the other three species. However, the latter
probably was not the case, as suggested by the wider latitude of
occupancy of plant communities than M. longicaudus, which showed
greater productivity.

Even though litter sizes are comparable, the first species coming
into breeding in the spring would have an advantage, especially if

216 GREAT BASIN NATURALIST Vol. 33, No. 4

the females establish a breeding territory excluding all other species
as suggested by Stoecker (1970) . Over the three months of this study,
nearly the entire adult populations of all four microtines were in
breeding condition.

Other factors that may be involved in segregation are behavioral
interactions (Murie, 1963; Stoecker, 1970) and possible physiological
differences in water balance (Getz, 1963). Stoecker (1970), in an
analysis of sympatry in M. pennsylvanicus and M. montanus. found
that because of co-occurrence in the same situations and because of
the similarity in their time of activity, behavioral contacts unques-
tionably occur. He found M. montanus to agonistically dominate M.
pennsylvanicus. Social intolerance between the two species was fur-
ther suggested by the low recruitment of young and by immigration
of one species when the other was removed by trapping. Stoecker's
(1970) results contrast with those of Grant (1970a), who noted that
the outcome of interspecies encounters depends on w'here and when
they occur and upon the relative body sizes of the interactants (lar-
ger species almost always wins) .

The existence of trends in differential habitat use was clearly
demonstrated in this study, and evidence suggests that the four mi-
crotines are at least partially incompatible. However, a multiplicity
of ecological and behavioral factors are probably operating syner-
gistically to produce the observed species distributions.

Acknowledgments

This study was supported by the Theodore Roosevelt Memorial
Fund of the American Museum of Natural History. Field assistance
of D. D. Skryja, F. J. Camenzind, and J. W. Huckabee was invalu-
able. Jackson Hole Biological Research Station facilities were used to
prepare specimens.

D. E. Casey, F. J. Camenzind, and C. G. MacFarland offered
critical advice on the manuscript. Illustrations were done by C.
Hughes. Assistance of National Park Service personnel, especially D.
B. Houston, was appreciated.

Literature Cited

Batzli, G. O. 1968. Dispersion patterns of mice in California annual grass-
lands. J. Mammal 49:239-250.

Cameron, A. W. 1964. Competitive exclusion between the rodent genera.
Microtus and Clethrionomys. Evolution 18:630-634.

Cl.'^rk, T. W. 1968. Food uses of the Richardson ground squirrel (Spermophi-
lus richardsonii elegans) in the Laramie Basin of Wyoming. Southwestern
Nat. 13:248-249.

1971. Ecology' of the western jumping mouse in Grand Teton Nation-
al Park, Wyoming." Northwest Sci. 45:229-238.

Corbet, G. B. 1964. Regional variation in the bank vole, Clethrionomys
glareolus, in the British Isles. Proc. Zool. Soc. Lond. 143:191-219.

CuRRY-LiNDAHL, K. 1959. Notes on the ecology' and periodicity of some ro-
dents and shrews in Sweden. Mammalia 23:389-422.

DeLong, K. T. 1966. Population ecology of feral house mice: Interference by
Microtus. Ecology 47:481-484.

Dec. 1973 clark: wyomin(; uodknts 217

FiNDLEY, J. S. 19'>1. Habitat preforenco of four species of Microtus in Jarkson

Hole, Wyoming. J. Mammal. 32:118-120.
■ . 1954. Competition as a possible limiting factor in the distribution of

Micro/us. Ecology 3'): 41 8-420.
Gp:tz, L. L. 1961 A comparison of the water balance of the prairie and mea-
dow voles. Ecology 44:202-207.
Grant, P. R. 1969. Experimental studies of competitive interactions in a two

species system. I. Microtus and Clethrionomys species in enclosure. Can. J.

Zool. 47:1059-1082.
— . 1970a. r^xperimenfal studies of competitive interactions in a two species

system. W. The behavior of Microtus, Peromyscus and Clethrionomys species.

Anim. Behav. 18:411-426.

1970b. Experim(>ntal studies of competitive interactions in a two spe-

cies system. III. Microtus and Clethrionomys species in enclosure. (In

preparation)
Hall, E. R.. and K. Kelson. 1959. The Mammals of North America. Ronald

Press, New York.
Hodgson, J. R. 1972. Local distribution of Microtus rnontanus and M. pennsyl-

vanicus in southwestern Montana. .1. Mammal. 51487-499.
IvERSON, S. L., AND B. N. TuRNER. 1972. Winter coexistence of Clethrionomys

gapperi and Microtus pennsylvanicus in a grassland habitat. Amer. Midi.

Nat. 88:440-445.
LiDicKRR, W. Z. 1966. Ecological observations on a feral house mouse popula-
tion declining to extinction. Ecol. Monogr. 36:27-50.
Maxri.i., M. H., AND L. N. Brown. 1968. Ecological distributions of rodents

on the high plains of easter-n Wyoming. vSouthwestern Nat. 13:143-158.
M'Ci.oskey, R. T. 1972. Temporal changes in [)opulations and species diversity

in a California rodent community. .1. Mammal. 5 3:657-676.
Morris, R. D. 1969. Competitive exclusion between Microtus and Clethri-
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Montana. Unpublished M.A. Thesis, Univ. Montana, Missoula.
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voles {Microtus). Amer. Midi. Nat. 82:622-625.
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Negus, N. C, and A. J. Pinta. 1966. Reproduction responses of Microtus

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Ota, K., and E. W. .Iameson, .1r. 1961. Ecological relationships and economic

importance of Japanese microtinae. Ecology 42:184-186.
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Stoecker, R. E. 1970. An analysis of sympatry in two species of Microtus.

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NOTES ON THE OCCURRENCE AND DISTRIBUTION OF

PTERONARCYS CALIFORNICA NEWPORT (PLECOPTERA)

WITHIN STREAMS

John A. Elder* and Arden R. Gaufin*

Abstract. — During a physiological study that required the collection of
several thousand stoneflies, Pteronarcys californica, observations were made on
the distribution of these aquatic insects within streams. It was noted that P.
californica occur primarily in large rivers that have loose, unconsolidated sub-
strates and swift currents. They occur from depths of six inches to several feet.
They are exclusively vegetarian, feeding on detritus, algae, and diatoms. P.
californica congregate in dense clumps, but vsdthin the clumps they segregate
by size. The largest specimens are found in the quietest water. There is evidence
of some seasonal variation in their behavior.

Pteronarcys californica is a large stonefly (Order Plecoptera)
of the mountain and Pacific west. Because of its large size (often in
excess of one gram) and its hardiness compared to other stoneflies,
it is an ideal organism to use in physiological studies.

During one such study, observations were made on the occur-
rence and distribution of this species within river systems. The
study required several thousand of these large insects, and collec-
tions were made throughout the year. Most of the observations
were made in Utah in the lower Provo River below Deer Creek
Reservoir. The section containing Pteronarcys californica was en-
tirely within Utah County from the immediate vicinity of Vivian
Park to the mouth of Provo Canyon. Residents in the area reported
that Pteronarcys californica had at one time existed close to Deer
Creek Reservoir but had been replaced by the smaller Pteronarcella
hadia in recent years. Upon investigation of the tailwater at this
point, it was discovered that the fauna and flora had the appear-
ance of a stressed environment. The algae were long, filamentous
greens, and the most common invertebrate was an amphipod (many
times more common than any other organism). This led us to the
conclusion that there was at least one period during the year when
poorly oxygenated water was discharged from Deer Creek Reservoir.
Apparently Pteronarcella can outcompete Pteronarcys in this type
of stress.

Other rivers where we collected Pteronarcys californica, to a
limited extent, were the Bear River upstream from Evanston, Wyo-
ming, Uintah County, to near the Utah border; North Fork, Teton
River, Idaho, at Idaho 32 bridge, Teton County; and Henry's Fork,
Idaho, near the U.S. 20, 191 bridge near Ashton in Fremont County.

Claassen (1931) states that Pteronarcys universally inhabits
"small, cold, upland spring brooks." This is not true of Pteronarcys
californica. It inhabits the larger rivers of the west that have large
volumes of water throughout the year. Man's alteration of the habi-

^Departnient of Biology, University of Utah, Salt Lake City 84112.

218

Dec. 1973 elder, gaufin: plecoptera 219

tat can adversely affect its distribution. Pteronarcys calif ornica
once was numerous on the Gunnison River, Colorado (Knight,
1965), prior to the construction of several dams. In 1972 when we
visited the river, Pteronarcys californica had not only disappeared
below the reservoirs but was very rare above them. Why this has
occurred is not known.

Usually Pteronarcys californica occurs in dense clumps in larger
streams. Occasionally, a few scattered individuals can be found, but
stream drift is probably responsible for this. They do not seem to
exhibit any drifting behavioral patterns, but in our holding tanks
Pteronarcys californica were nocturnally active. Heaton (1966)
found a 22: 1 night-day drift ratio for this species. It is possible that
Pteronarcys californica drift at night. Their typical behavior when
dropped into flowing water or when dislodged from the substrate is
to "ball up" and drop to the bottom.

Large populations of Pteronarcys californica are typically found
in swift water but rarely in "white" water. We have found them
in depths ranging from six inches to three feet, and they probably
occur deeper. The rocks are notably free of large clumps of algae,
and the bottom of rooted aquatics. Loose, unconsolidated bottoms
seem to be preferred, with the minimum size of stones approxi-
mately six inches and maximum size unknown. In Henry's Fork
we collected in excess of 100 individuals from one rock of approxi-
mately three feet in diameter.

Pteronarcys californica seem to prefer loose rock where terres-
trially derived detritus collects, or rocks with a coating of diatoms
and other small green algae. Richardson (1965) reported that
Pteronarcys californica is a detritus feeder. Our analysis of gut con-
tents from more than 1000 individuals has not revealed a single
case of carnivorous feeding.

Pteronarcys californica nymphs tend to segregate by size (size
is determined by age and sex). The aquatic stage of this species
lasts three years. In a single stretch of water on the Provo River at
Vivian Park, third-year females, the largest size, were found
farthest upstream in the calmer water with the largest rubble.
Pteronarcys californica density was the lowest in this type habitat.
Downstream, as the depth decreased, the rubble became smaller
and looser, and the organisms became smaller and more numerous.
Here, while measurements were not made, it appeared that second-
year females were mixed with second- and third-year males. First-
year specimens were found commonly with the second-year females
and second- and third-year males but were most common in areas
lacking the larger classes. In the coldest winter months, when the
rivers were lowest, there was some indication that the size segrega-
tion partially broke down. This was probably due to lack of avail-
able habitat.

During the spring, May- July, when Pteronarcys californica
emerge, the rivers rise to such an extent that sampling is virtually
impossible. However, it has been noted by Gaufin that the emerging
insects migrate to shallow water, where the rocks are partially ex-

220 GREAT BASIN NATURALIST Vol. 33, No. 4

posed. Here they congregate and wait for the cue to emerge. Emer-
gence is accomplished by crawhng out of the water onto a rock or
shore, spHtting the nymphal exoskeleton, and emerging as an aduh.
On most rivers where we collected, Pteronarcys californica.
when found, was extremely abundant, but in very limited areas of
one hundred meters or less. On the Henry's Fork, Pteronarcys cali-
fornica was much more abundant than elsewhere we collected.
Nevertheless, in any given riffle, a sample of approximately one
meter square might yield more than 100 organisms, while only a
few meters away on rubble that was superficially identical an area
of similar size might yield few, if any, individuals.

References Cited

Claassen, p. W. 1931. Plecoptera nymphs of America (north of Mexico).
Thomas Say Foundation, Inc., Springfield, III.

Heaton, J. R. 1966. The benthos and drift fauna of a riffle in the Madison
River, Yellowstone National Park. Ph.D. Dissertation. Montana State
University.

Knight, A. W. 1965. Studies of the stoneflies (Plecoptera) of the Gunnison
River drainage in Colorado. Ph.D. Dissertation. University of Utah.

Richardson, J. W., Jr. 1965. The food habits of plecoptera. M.S. Thesis. Univer-
sity of Utah.

PREHISTORIC BIGHORN SHEEP IN
THE NORTHERN SIERRA NEVADA, CALIFORNIA

Peter D. Schulz' and Dwight D. Simons'

Abstract. — Data from pictograph and archaeological sites in northeastern
California are used to verify the postulated former existence of small resident
populations of bighorn sheep (Ovis canadensis) in this region. The reason for
the disappearance of bighorns froin this area remains unclear, with climatic
change, Euro-American settlement, or hunting pressure existing as possibilities.

The known historic range of bighorn (Ovis canadensis) in Cali-
fornia has been reported by Jones (1950). In the 160 miles between
Observation Peak in east central Lassen Cotinty and the southeastern
portion of Alpine County to the south, there is no record of bighorn
sheep except for an isolated sighting east of Donner Pass, Nevada
County (Wistar, 1914:113). Some authors (Cowan, 1940; Hall and
Kelson, 1959; Buechner, 1960) have posited the entire Sierra Ne-
vada as pristine bighorn range, but no evidence exists in the zoologi-
cal literature to support this suggestion. Archaeological research
in this area during the last decade and a half sheds considerable
light on the matter. This note wall present a brief review of relevant
findings and a description of new material analyzed by the authors.

Three archaeological sites in northeastern California contain
rock carvings of bighorns (Heizer and Baumhoff, 1962; Payen,
1966). These are: site 4-Las-38 in Ball's Canyon, 8 miles north of
Standish, Lassen County; site 4-Sie-l, V4 mile southeast of Hawley
Lake, Sierra County; and site 4-Pla-26, 11 miles south of Old High-
way 40 at Soda Springs, Placer County. Heizer and Baumhoff
(1962) and Grant et al. (1968) discuss the relevance of pictograph
distributions to the prehistoric range of mountain sheep, particularly
in Nevada and southeastern California. Their research suggests
that these pictographs functioned in hunting rituals. The paucity of
such sites in our area of concern, however, lends little aid to range
determination.

Direct evidence of late prehistoric sheep populations exists in
the faunal remains from five archaeological sites (Figure 1). Age
estimations for all the sites are based on artifact typology and are
considered reliable, since these are correlated with radiocarbon-
dated deposits in neighboring areas of California and Nevada.

Bare Cave (4-Las-S228) is located in northeastern Lassen County
at the southernmost end of Surprise Valley, 10.5 miles southeast
of Eagleville. Although the fauna from this site has not yet under-
gone systematic analysis, the authors have identified remains of
several individuals of Ovis canadensis represented by abundant
cranial and postcranial elements. Stratigraphic records on the faunal
remains appear to have been lost, and thus changes in sheep abun-

'Department of Anthropology, University of California, Davis, Calif. 95616.

221

222 GREAT BASIN NATURALIST Vol. 33, No. 4

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Fig. 1. Map of the counties of northeastern California showing archaeological
sites with bighorn sheep remains (circles) and bighorn pictographs (triangles).
Stippling indicates historic range (after Jones, 1950).

dance through time cannot be demonstrated. Brown (1964) dates
the deposit between 2000 b.c. and a.d. 1200.

The Karlo site (4-Las-7) is approximately 20 miles northeast
of Susanville, Lassen County, near the Karlo siding on the Southern
Pacific line between Susanville and Alturas. Riddell (1960) de-
scribes numerous bone tools made from metapodials and scapulae
of Ovis canadensis. Unmodified bones of this species w^ere also pres-
ent in the deposit. Sheep remains are more abundant in the upper
levels of the site, but this correlates with other bone remains and
cannot be used as evidence of faunal change. It is suggested that
the site was occupied seasonally from ca. 2000 B.C. until the nine-
teenth century a.d.

Tommy Tucker Cave (4-Las-l) lies 4 miles southeast of Wen-
del, Lassen County, on the south slope of Hot Springs Peak. Riddell
(1956) reports 11 bones assignable to Ovis canadensis. Remains
from the deposit range in age from ca. a.d. 1000 to the his-
toric period.

Chilcoot Rockshelter (4-Plu-44) is situated 7.5 miles north of
Vinton, Plumas County. Bone preservation was extremely poor,
and Payen and Boloyan (1961) report only a single specimen at-
tributable to bighorn. The deposit is dated from a.d. 1400 to 1850.

The Loyalton Rockshelter (4-Sie-S43) is located on Elephant
Head Peak, 2 miles northeast of Loyalton, Sierra County. Faunal
remains were fairly abundant, although the deposit was less than

Dec. 1973 schulz, simons: bighorn shkep 223

a foot deep. We recorded 7 adult calvaria definitely assignable to
Ovis canadensis and 2 foetal crania assignable to Ovis and almost
certainly associable with the bighorn remains. Postcranial elements
were also abundant in the deposit but were so badly smashed for
marrow extraction and tool use that few were identifiable. The
presence of adults and foetuses indicates occupation of the area by
a resident sheep population rather than by stray individuals. Wilson
(1963) attributes the deposit to the period between ca. a. d. 1000
and the Euro- American immigration.

The presence of bighorn remains at these sites provides signifi-
cant indication that the species was widely present in the northern
Sierra Nevada of California during the late prehistoric period. The
reason for the disappearance of the form over all of this range is
unclear. At present, factors such as late climatic change in the area,
withdrawal of sheep before immigrating Euro-Americans, or ex-
tinction of local populations by hunting are possibilities. Since most
of the sites contain very few individuals and only three sheep picto-
graphs have been recorded for the entire area (as opposed to more
than 7,000 in Inyo County alone), it is fairly certain that bighorns
were never as abundant in this region as they were farther south
and east. Thomas (1972) notes that if local bighorn bands are
hunted out, their area will be slowly or never repopulated. This re-
sults from the maintenance of similar home ranges by bighorn
bands from generation to generation. This trait, coupled with low
population density, would have facilitated extinction through any
of the agencies mentioned above.

We wish to thank D. L. True and R. L. Rudd for criticizing the
manuscript, and M. A. Baumhoff for discussions which stimulated
our work in this area.

Literature Cited

Brown, W. R. 1964. The prehistory of Surprise Valley. Unpublished M. A.
Thesis. Univ. California, Davis.

BuECHNER, H. K. 1960. The bighorn sheep in the United States: Its past, present,
and future. Wildlife Monographs 4:1-174.

Cowan, I. McT. 1940. Distribution and variation in the native sheep of North
America. Amer. Midi. Nat. 24:505-580.

Grant, C, J. W. Baird, and J. K. Pringle. 1968. Rock drawings of the Coso
Range, Inyo County, California. Maturango Mus. Publ. 4:1-145.

Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Ronald
Press New York.

Heizer, R. F., and M. A. Baumhoff. 1962. Prehistoric rock art of Nevada and
eastern California. Univ. California Press, Berkeley. --, . d n

Jones, F. L. 1960. A survey of the Sierra Nevada bighorn. Sierra Club Bull.
35-29-76.

Payen, L. a. 1966. Prehistoric rock art in the northern Sierra Nevada, Califor-
nia. Unpublished M. A. Thesis. Sacramento State College.

Payen L A, and D. S. Boloyan. 1961. Archeological excavations at Chilcoot
Rockshelter, Plumas County, California. California Dept. Parks and Recrea-
tion Archeol. Rep. 4:1-14. ry. i /->

Riddle, F. A. 1956. Final report of the archaeology of Tommy Tucker Lave.
Univ. California Archaeol. Survey Rep. 35:1-25. ^ ^ ,., • tt •

. I960. The archaeology of the Karlo site (Las-7), Califorma. Univ.

California Archaeol. Survey Rep. 53:1-133.

224 GREAT BASIN NATURALIST Vol. 33, No. 4

Thomas, D. H. 1972. Unmodified faunal renrains from NV-Pe-104. In Cowan,
R. A. [Ed.] The Archaeology of Barrel Springs Site (NV-Pe-104), Pershing
County, Nevada. Archaeol. Res. Facil., Dept. Anthropol., University Califor-
nia, Berkeley, pp. 41-48.

Wilson, N. L. 1963. The archaeology of the Loyalton Rock Shelter, Sierra
County, California. Unpublished M. A. Thesis. Sacramento State College.

WisTAR, I. J. 1914. Autobiography of Isaac Jones Wistar. Wistar Inst, of Anat.
and Biol., Philadelphia.

ARTEMISIA ARBUSCULA, A. LONGILOBA, AND A. NOVA
HABITAT TYPES IN NORTHERN NEVADA

B. Zainora^'2 and P. T. Tueller^

Abstract. — Artemisia arbuscula, A. longiloba, and A. nova are dwarf sage-
brush species that occur extensively throughout the shrub steppe of northern
Nevada. These species are similar ecologically in that they occupy habitats
strongly influenced by edaphic factors. Nine major habitat types on which these
shrubs are dominant are recognized in this region. The A. arbuscula habitat
types are most prevalent in extreme northern Nevada. Southward, they generally
become restricted to altitudes above the Pinus-Juniperus woodland zone. A single
A. longiloba habitat type is described, occurring in northeastern Nevada. The A.
nova habitat types are most prevalent in north central and east central Nevada.
Four features appear consistently among soils of these habitat types: (1) shal-
lowness, (2) high profile gravel volume, (3) presence of a clay B horizon close
to the soil surface, and (4) presence of large quantities of mineral carbonates
throughout profiles of most A. nova habitat types.

Artemisia arbuscula, A. longiloba, and A. nova are dwarf sage-
brush species which dominate approximately one-half of the sage-
brush vegetation in Nevada (Beetle, 1960). The most extensive com-
munities of these species in Nevada occur in the northern portions
of the state and form very distinct and important components of the
shrub steppe of this region. Excessive livestock grazing in past years
has greatly altered the herbaceous composition of the vegetation of
many of these communities. Virtually no synecological studies of
undisturbed A. arbuscula, A. longiloba, and A. nova vegetation have
been undertaken in Nevada (Passey and Hugie, 1962). Such infor-
mation would be useful in ascertaining the biotic potential and limi-
tations of the habitats occupied by these shrubs and would greatly
aid in the management and manipulation of these communities for
grazing and watershed purposes. The purpose of this study was to
dehmit the characteristics of the major A. arbuscula, A. longiloba,
and A. nova habitat types in the shrub steppe of northern Nevada.

Study Area

The study encompassed areas throughout northern Nevada where
these species of sagebrush were abundant and where undisturbed
vegetation would likely be found. The region referred to as northern
Nevada is that portion of the state generally north of the 39th paral-
lel. The physiography of much of this region consists of numerous,
short mountain ranges and intermontane basins with internal drain-
age. Extreme northern Nevada consists of an upwarped plateau of
hilly and mountainous terrain with major drainage into the Snake
River to the north in Oregon and Idaho. Soils of this region belong
predominantly to the Aridisol, Entisol, Mollisol, and Vertisol orders.
The climate is arid. The mean annual temperature is 8.9 C, with the
coldest temperatures occurring in December through February

^Renewable Resources Center, University of Nevada, Reno 89507. , . . , ^^

^Present address: Agricultural Research Service, U.S. Department of Agriculture, Department
of Agronomy and Soils, Washington State University, Pullman 99163.

225

226 GREAT BASIN NATURALIST Vol. 33, No. 4

(inean = 2.2 C) and the warmest in July and August (mean=20.8
C). The mean total annual precipitation is 239.2 mm. The long-
range normals (in mm) for total monthly precipitation are as fol-
lows: January 29.2, February 26.4, March 23.9, April 22.9, May
24.6, June 18.0, July 9.9, August 7.6, September 9.9, October 18.8,
November 20.6, December 27.4.

The shrub steppe in extreme northern Nevada appears as a uni-
form, continuous shrub cover occasionally interrupted by Juniperus
or Cercocarpus woodlands. In north central Nevada, shrub steppe
communities occupy basins and lower pediment slopes of the moun-
tain ranges. At higher altitudes, the shrub steppe is interrupted by a
zone of Pinus monophylla and Juniperus osteosperma woodlands
(Billings, 1951). Above this zone, shrub communities again appear,
particularly those dominated by Artemisia.

Methods

A preliminary reconnaissance of the vegetation dominated by
Artemisia arbuscula, A. longiloba, and A. nova gave an indication of
the community types, distribution, and pattern of occurrence. Lists
of the dominant species \'\dthin each stand were taken during the
reconnaissance. These species were then ranked according to their
abundance. Nine community types were subjectively delineated by
examination of these lists in stand tables. Based on this initial clas-
sification, 39 stands representing the various community types were
selected for detailed study on the basis of the following criteria: ab-
sence of disturbance due to grazing, erosion, and wildfire; absence of
a tree overstory; presence of a uniform shrub and grass cover.

Vegetation data were collected from a 15-by-30-m macroplot lo-
cated within each of the 39 stands. Ten 15-m transects were ran-
domly located perpendicular to the long axis of the macroplot. Fre-
quency data for plant species were collected from 10 3-by-6-dm ob-
servation plots placed at 1-m intervals along each transect. Only
species rooted within each observation plot were tallied. The same
observation plots were used to sample basal area of each species along
4 of the 10 transects. Species not recorded in the observation plots
but present within the macroplot w-ere listed. Shrub measurements
were collected from 1-by-l-m plots placed contiguous and parallel to
the same 4 transects from which basal area data were collected.
Crown cover, density, and maximum heights were recorded for each
shrub species within each meter-square plot.

Physiographic features recorded at each location were: percentage
slope, slope aspect, position of the stand on the slope, elevation, land-
form on which the stand occurred, and terrain of the general area in
which the stand occurred. A profile description of the soil underlying
each stand was made and identified to the family level (Soil Survey
Staff, 1967). One soil sample from each horizon of each profile was
collected for laboratory analysis. Soil reaction w^as measured on a
saturated paste with a glass electrode pH meter. Electrical conduc-
tivity of an extract from a saturated paste was determined only on

Dec. 1973 zamora, tueller: artemesia 227

the Al and B2 horizons of each profile. Calcium carbonate equiva-
lents were determined by gravimetric loss of carbon dioxide upon
treatment wdth 3NHC1 for all horizons.

The vegetation data from the 39 stands were summarized in stand
tables and mean values for the various characteristics calculated.
The vegetation and associated environmental characteristics were in-
terpreted according to the association and habitat type synecological
concepts (Daubenmire, 1968). Stands similar in both shrub and herb
dominance were considered representative of an association. A habi-
tat type is a unit of land that now supports or is still capable of sup-
porting one plant association. The name of each habitat type con-
sists of the name of the principal shrub and herb of the association
which occupies it. Species nomenclature follows that of Ward (1953)
and Beetle (1960) for Artemisia and Holmgren and Reveal (1966)
for other vascular plants.

Description of Habitat Types

Artemisia arbuscula is most abundant throughout extreme north-
ern Nevada. Southward into central Nevada. A. arbuscula is found
predominantly at higher altitudes above the Pinus monophyUa-J uni-
perus osteosperma woodland zone. A. longiloba occurs almost en-
tirely within a small area in north central Elko County in north-
eastern Nevada. Occasional small, insular stands may be found in
surrounding counties. A. nova is most abundant in the north central
and eastern portions of the state. Scattered, insular stands may be
found throughout northern and southern Nevada.

The physiognomy of the vegetation is quite similar throughout
the entire range of these species in Nevada. The shrubs appear as a
low, uniformly distributed population interspersed by perennial,
caespitose grasses. Separate shrub and grass layers are not always
distinguishable because of the similarity in height of most shrubs
and grasses. In most areas, individual stands of these dwarf sage-
brush species are contiguous to extensive communities of A. tridenta-
ta, the predominant sagebrush species in Nevada. Ecotones between
contiguous stands often appear abrupt. The low, compact gro\\th
form of A. arbuscula, A. longiloba, and A. nova contrast sharply with
the taller, robust, and diffusely branched growth form of A. tridcnta-
ta.

Nine major habitat types were recognized on the basis of shrub
and grass dominance in this region. The 39 stands studied in detail
represent the vegetation of 8 of the habitat types. An additional habi-
tat type is described on the basis of reconnaissance data only. Con-
stancy, basal area, and frequency of major plant species in each as-
sociation are given in Tables 1 and 3. Shrub characteristics are shown
in Tables 2 and 4.

(1) Artemisia arbuscula-Festuca idahoensis habitat type
This habitat type is found on slopes and ridgetops of hilly and
mountainous terrain throughout much of northern Nevada at alti-

228

GREAT BASIN NATURALIST

Vol. 33, No. 4

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Table 4. Mean values for shrub characteristics" of three Artemisia nova
associations.

A. nova- A. nova-

Agropyron Agropyron A. nova-

spicatum inerme Stipa

'No Ht C^ No Ht Cv li^ Ht Cv

Artemisia nova 181 23 13 133 14 11 216 15 12

Chrysothamnus viscidiflorus 17 20 1 7 15 T 18 13 T

Eurotia lanata 7 6TT 9T 1514T

Ephreda viridis T 30 T T 1 1 T 1 49 T

Eriogonum microthecum 4 12 T — — — — — —

Tetradymia canescens 2 22T — — — — — —

Leptodactylon pungens 2 27 T — — — — — —

Grayia spinosa — — — T19T — — —

Atriplex confertifolia ___ 2 19T 2 16T

tudes ranging from 1900 to 2000 m. Artemisia arbuscula is the domi-
nant shrub of the association. Festuca idahoensis is the dominant
perennial herb, with Poa sandbergii and Agropyron spicatum the
principal subdominants. Low scattered bushes of Chrysothamnus vis-
cidiflorus occur in all stands. Bromus tectorum, the only annual
grass, occurs in very minute amounts but becomes abundant in most
serai stands. Forbs in all Artemisia arbuscula associations are very
low in basal area but often attain relatively high frequencies. Soils
supporting stands studied in detail were divided among the subgroups
Typic Haploxerolls, Typic Argixerolls, Typic Durixorolls, and Xerol-
lic Camborthids. The association is similar to those described by
Eckert (1957), Tueller (1962), Culver (1964), Hall (1967), and
Dealy (1971) in central and eastern Oregon.

(2) Artemisia longiloba-Festuca idahoensis habitat type

In north central Elko County in northeastern Nevada, stands of
Artemisia longiloba may be found on slopes of gently rolling to hilly
terrain at ahitudes ranging from 1900 to 2100 m. Only two stands
were found that were not extensively altered by grazing. Except for
the substitution of Artemisia longiloba for Artemisia arbuscula, the
vegetation closely resembles that of the Artemisia abruscula-F estuca
idahoensis association in both floristics and abundance of herbaceous
species. Soils of the two stands belong to the subgroups Xerollic
Camborthids and Mollic Paleargids. Tisdale, Hironaka, and Fosberg
(1965) described a pristine Artemisia longiloba-Festuca idalioensis-
Stipa thurberiana community in southern Idaho. The abundance of
5. thurberiana was the only major difference between the community
in Idaho and the two stands in Nevada. Hugie, Passey, and Wd-
liams (1964) described an Artemisia longiloba community in the
Snake River Plain, Idaho, in which F. idahoensis, Agropyron spica-
tum, and Poa sandbergii were the principal grasses. Robertson, Niel-
sen, and Bare (1966) report that Agropyron spicatum is the pre-

234

GREAT BASIN NATURALIST

Vol. 33, No. 4

dominant grass associated with Artemisia longiloba on range site in
Colorado.

(3) Artemisia arbuscula-Stipa thurberiana habitat type

The Artemisia arbuscula-Stipa thurberiana habitat type occurs on
gently rolling to hilly terrain primarily in northwestern Nevada at
altitudes varying around 1800 m. Stands are distinguished by the
dominance of the perennial grass Stipa thurberiana. Subdominant
grasses include Poa sandbergii and Sitanion hystrix. Festuca idahoen-
sis is low in constancy. Mat-forming forbs are a conspicuous compo-
nent of the vegetation, the most prevalent species being Eriogonum
ovalifolium. Phlox hoodii. Aster scopuJorum, and Astragalus purshii.
Soils underlying stands studied in detail belong to the subgroups
Mollic Paleic Durargids, MoUic Paleargids, Mollic Haplargids, and
Typic Durixerolls.

(4) Artemisia arbuscula-Agropyron spicatum habitat type

The Artemisia arbuscula-Agropyron spicatum habitat type occurs
on slopes and ridgetops of hilly and mountainous terrain in north-
eastern and east central Nevada (Figure 1). Altitudes of stands
range from 1900 to 2700 m. The association is characterized by the
abundance of Agropyron spicatum. Festuca idahoensis is usually ab-
sent but may occur in minute amounts. In some stands, Poa sand-

Fig. 1. Stand representative of the Artemisia arbuscula-Agropyron spica-
tum association in northern Nevada. The meter stake is marked in decimeters.

Dec. 1973

ZAMORA, TUELLEn: ARTEMESIA

235

bergii may equal or exceed Agropyron spicatum in both basal area
and frequency. Chrysothamnus viscidiflorus is the only other shrub
of high constancy. In east central Nevada, this association was found
only above the Pinus-J uniperus woodland zone. The growth form of
Artemisia arbuscula was greatly altered at these higher altitudes,
appearing as an almost prostrate shrub with dense, flattened cano-
pies. Scattered bushes of Artemisia frigida and Artemisia nova may
be present in these stands. Soils supporting the stands studied in de-
tail are divided among the subgroups Lithic Argixerolls, Typic Hap-
loxerolls, and MoUic Haplargids. Eckert (1957), Culver (1964), and
Hall (1967) described similar associations in central and eastern
Oregon.

(5) Artemisia arbuscula-Purshia tridentata- Agropyron spicatum

habitat type

The Artemisia arbuscula-Purshia-Agropyron habitat type was
found only on the pediment slopes of Duck Creek Basin in the Schell
Creek mountain range northeast of Ely, Nevada (Figure 2). These
slopes are dissected, forming many narrow to broadly rounded ridges
extending from the mountain range into the basin. Stands occur be-
tween 2400 and 2500 m elevation. All soils described belong to the
subgroups Typic Argixerolls. Codominance of the shrubs Artemisia
arbuscula and Purshia tridentata characterizes the association. Pursh-

Fig. 2. Stand representative of the Artemisia arbuscula-Purshii tridentata-
Agropyron spicatum association in the Duck Creek Basin in east central Nevada.

236

GREAT BASIN NATURALIST

Vol. 33, No. 4

ia tridentata has a slightly larger crown cover than Artemisia arbus-
cula but considerably lower density (Table 2). Scattered individuals
of Artemisia tridentata, Symphoricarpos longiflorus, Tetradymia ca-
nescens, and C hrysothamnus viscidiflorus occur throughout the vege-
tation. Agropyron spicatum is the dominant perennial herb. Stands of
this association occur at the lower edges of the Piniis monophylla-
Juniperus osteosperma zone within the basin. Dealy (1971) described
a Purshia tridentata-Artemisia arhuscula-Stipa thurberiana commu-
nity in south central Oregon which occurred near the lower edge of
the Finns ponderosa forests.

(6) Artemisia nova- Agropyron spicatum habitat type

The Artemisia nova-Agropyron spicatum habitat type occurs
throughout north central Nevada, i)rincipally on slopes of genth'
rolling to hilly terrain at altitudes ranging from 2000 to 2300 m
(Figure 3). This habitat type may also occur at higher altitudes on
certain mountain ranges where soils are derived from calcareous
parent materials. Such areas are found in the Snake, Schell Creek,
and Egan ranges of east central Nevada. Artemisia nova is the prin-
cipal shrub and Agropyron spicatum the principal herb. Poa sand-
bergii occurs in small amounts in most stands but increases in abun-
dance at higher elevations. Other grasses such ns Poa nevadensis.

Fig. i. Stand representative of the Artemisia nova— Agropyron spicatum as-
sociation on slopes of the Schell Creek Range in east central Nevada. The meter
stake is marked in decimeters.

Dec. 1973 zamora, tuelleu: artf.mf.sia 237

Bromus tcctorum. Oryzopsis hymenoides, Sitanion hystrix, and Stipo
thurbcriana are present in very limited amounts. Chrysothnmnus
viscidiflorus is the only other shrub of high constancy. Stands of this
association and Artemisia tridcntata communities often constitute
the major vegetation directly below and above the Pinus niono-
phylla-J uniperus osteosperma woodland zone in east central Neva-
da. Artemisia nova may also occupy openings within the woodlands
and form a shrub layer beneath the tree canopies. The soils support-
ing the stands studied in detail were divided among the subgroups
Typic Torriorthents, Pachic ArgixeroUs, Typic Durixerolls, Pachic
Haploxerolls, and Xerollic Paleorthids.

(7) Artemisia nova-Agropyron inerme habitat type

This habitat type occurs on foothill slopes of the Pancake moun-
tain range predominantly in Nye County in central Nevada. It
ranges in altitude from 1800 to 2100 m. Agropyron inerme is the
dominant herb. The tall, robust, caespitosc growth form of this grass
is the most conspicuous feature of the physiognomy. Poa sandbergii
and Sitanion hystrix are the only other grasses with high constancy
among the stands. Stands of the Artemisia nove-Agropyron inerme
association are commonly associated with extensive communities of
Atriplex confertifolia and Grayia spinosa. Influence of the Atriplex
and Grayia vegetation on these stands is evident in the presence of
shrubs such as Atriplex confertifolia, Eurotia lanata. Ephedra viridis.
and Grayia spinosa, which are characteristic of the Atriplex and
Grayia vegetation. The soils described belong to the Aridisol sub-
groups Mollic Paleargids, Mollic Haplargids, and Lithic Mollic Hap-
largids.

(8) Artemisia novaStipa comata habitat type

The Artemisia nova-Stipa habitat type is found in White Pine
County on pediment slopes of hilly terrain. The altitudinal range is
2000 to 2200 m. The association is characterized by the abundance
of Stipa comata. Poa sandbergii and Sitanion hystrix are highly con-
stant grasses but are low in frequency and basal area. Bromus tec-
torum and HiJaria jamesii may be present in very small amounts.
A. nova has a high density in comparison to other A. nova associa-
tions (Table 2). Scattered individuals of Chrysothamnus viscidi-
florus and Eurotia lanata are generally present. Soils underlying
these stands belong to the subgroups Typic Torriorthents and Xerol-
lic Paleorthids.

(9) Artemisia nova-Oryzopsis hymenoides habitat type

The Artemisia nova-Oryzopsis habitat type is described only
from reconnaissance data. The vegetation of this habitat type has
been severely grazed by livestock in the past. Because of the nature
of the terrain, the vegetation is easily accessible and conducive to
fall and winter grazing by livestock. No stands were found that had

238

GREAT BASIN NATURALIST

Vol. 33, No. 4

not been affected by grazing to various degrees. However, the spe-
cies composition of the vegetation is quite constant over large areas
even though the herbage is often reduced to very small quantities
by livestock.

The habitat type occurs extensively on undulating and gently
rolling pediment slopes in the intermontane basins in central and
east central Nevada (Figure 4). These slopes are generally less than
5 percent in steepness. Areas of the Artemisia nova-Oryzopsis hab-
itat type generally occur over an altitudinal range of 1800-2300 m.
Soils are predominantly of the Aridisol and Entisol orders.

Artemisia nova is the dominant species of the vegetation. Chryso-
thamnus viscidiflorus and Eurotia lanata are the only shrubs com-
monly occurring with Artemisia nova. The following additional
shrubs may occur and are listed in order of decreasing presence:
Ephredra viridis, Atriplex confertifolia, Eriogonum microthecum,
Artemisia tridentata, Leptodactylon pungens and Tetradymia canes-
ccns. The herb layer is very sparse, composed principally of the
caespitose grass O. hymenoides. Other grasses that may be occasion-
ally found are Poa sandbergii, Sitanion hystrix^ Stipa comata, and
Hilaria jamesii. Forb species are generally scarce throughout the
vegetation. The most abundant, in order of decreasing presence, are
Phlox hoodii, Eriogonum oval i folium, Haplopappus acaulis, Caulan-
thus crassicaulis, Erigeron pumilis. Phlox longi folia, Sphaeralcea

Fig. 4. Stand representative
association in central Nevada.

of the Artemisia nova-Oryzopsis hymenoides

Dec. 1973 zamora, tuellf.h: ahtf-mhsia 239

grossulariaefolia, and Astragalus calycocus. Stands of the Artemisia
nova-Oryzopsis association are commonly contiguous with stands of
Atriplex, Eurotia lanata, and Grayia spinosa.

Soils

Considerable variation in morphology was encountered among
the soils of the Artemisia arbuscula, Artemisia longiloba and Artemi-
sia nova habitat types. Of the 39 soils described, 25 different families
were identified. This was expected, since many of the soils are dis-
tributed over a wdde geographical area and occur in a variety of
topographic positions. The more prominent features of tho collective
soils are described.

The sola thickness of soils of the Artemisia arbuscula and Arte-
misia longiloba habitat types varied from 23 to 87 cm, with an aver-
age of 55 cm. Sixty-five percent of these soils have dense clay hori-
zons (B2t). The upper boundaries of these horizons varied from 7
to 36 cm below the soil surface, with an average of 19 cm. These
prominent clay horizons did not form a root-restricting layer, since
roots were abundant throughout their depth. Sixty-five percent of the
soils have sola with high gravel volumes ranging from 20 to 62 per-
cent. The majority of these soils belong to the Artemisia arbuscula-
Agropyron and Artemisia arbuscula-Purshia-Agropyron habitat
types. Those soils lacking dense clay horizons have sola with high
gravel volumes. Root-restricting layers (duripans or bedrock) were
found in only 21 percent of the soils, occurring at depths of 23-87
cm. The remaining soils were generally underlain by gravelly loam
strata. The occurrence of these layers was not restricted to soils of
any particular habitat type. Sola of the Artemisia arbuscula and
Artemisia longiloba soils were not calcareous nor alkaline. Soil re-
action of the Al and B2 horizons varied from pH 5.5 to 7.4, with
electrical conductivities ranging from 0.1 to 1.0 mmhos per cm.

The sola thickness of soils of the Artemisia nova habitat types
varied from 7 to 89 cm, with an average of only 26 percent of the
Artemisia nova soils described having dense clay horizons. The up-
per boundaries of these horizons ranged from 7 to 15 cm below the
soil surface. Eighty-one percent of the Artemisia nova soils have
high volumes of gravel in their sola, varying from 26 to 52 percent.
Root-restricting layers (duripans, petrocalcic horizons, or bedrock)
were found in 50 percent of the soils at depths of 29-68 cm and did
not occur predominantly in any one habitat type. The remaining
soils were generally underlain by gravelly and sandy loam strata.
The majority of the soils of the Artemisia nova habitat types are cal-
careous throughout their sola. Percentage CaCO.s equivalents of the
Al and B2 horizons varied from 9 to 36 percent. Soil reaction of
these horizons ranged from pH 6.6 to 8.5, with electrical conductivi-
ties varying from 0.2 to 0.7 mmhos per cm. However, sola of soils
of the Artemisia nova-Agropyron inerme habitat type are generally
free of detectable mineral carbonates. Carbonates are present in C
horizons at depths of 23-41 cm. Soil reaction of the Al and B2 hori-

240 GREAT BASIN NATURALIST Vol. 33, No. 4

zons in these soils ranged from pH 6.3 to 1 .1 , with electrical con-
ductivities varying from 0.2 to 1.1 mmhos per cm.

Discussion

Artemisia arhuscula, A. longiloha, and A. nova vegetation is pres-
ent in numerous states surrounding Nevada (Ward, 1953; Beetle,
1960). Studies in these areas have generally indicated that the dis-
tributions of these species are edaphically determined.

Eckert (1957), Tueller (1962), Culver (1964), Hall (1967), and
Dealy (1971) found A. arbuscula associations in central and south-
eastern Oregon to occur on soils that were shallow, stony, and had
moderately to strongly structured B2t horizons high in clay and very
close to the soil surface. Hugie, Passey, and Williams (1964) and
Tisdale, Hironaka, and Fosberg (1965) in southern Idaho and Rob-
ertson, Nielsen, and Bare (1966) in Colorado report similar mor-
phology for soils underlying A. longiloba communities. Fosberg and
Hironaka (1964) compared soils supporting A. arbuscula and A. tri-
dentata communities in southern Idaho. A. arbuscula was restricted
to soils that had a clay B horizon or bedrock within 33 cm from the
surface or to weakly developed soils that were as deep as 50 cm but
had over 30 percent gravels and cobbles distributed throughout their
profiles. Soils supporting A. tridcntata communities were deeper and
had weaker profile development. They suggested that the properties
of the A. arbuscula soils result in conditions of poor a'^ration in the
rooting zone during the winter and spring months due to the devel-
opment of a perched water table above the dense clay horizon or
bedrock. Summerfield (1969) concluded that the depth of a clayey
horizon of prominent, compound prismatic, and blocky structure be-
low the soil surface was the only consistent morphological difference
between soils supporting A. arbuscula and A. trident ata communities
in northwestern Nevada. These horizons occurred from 7 to 31 cm
under A. arbuscula communities and 31 to 53 cm under A. tridentata
communities.

Shantz (1925), Shreve (1942), Fautin (1946), Thatcher (1959),
and Richards and Beatley (1965) describe A. nova communities oc-
curring on gravelly, shallow soils on ridgetops or slopes. Andre,
Mooney, and Wright (1965) observed that in the White Mountains
of California A. nova was most prevalent on limestone areas. Hirona-
ka (1963) found that A. nova occurred on highly calcareous soils
and shallow gravelly soils developed from limestone parent material.

Although there is considerable variation in the soils of the habitat
types described in this study, four salient features appear consistent-
ly among the soils. These features are: (1) shallow^ness and thin-
ness of the sola, (2) presence of a dense clay B horizon close to the
soil surface, (3) a high volume of gravel throughout the profile,
and (4) presence of mineral carbonates throughout most sola of the
A. nova habitat types. The regularity of occurrence of these features
is confirmed by the studies of Blackburn et al. (1968, 1969) of the
vegetation and associated site characteristics on watersheds through-

Dec. 1973 zamora, tui.i.lkh: artf.ivif.sia 241

out northern Nevada. The presence of a dense clay B horizon was
the most commonly occurring feature among the A. arbuscula and
A. longiloba soils. In soils lacking this prominent horizon, very high
gravel volumes occurred throughout the profile. Not all of the soils
that lacked this dense clay B horizon were shallow, however . Several
of the soils contained no restricting layer within a meter of the soil
surface, indicating that extreme droughtiness may be the dominant
physical factor of these soils. Shallowness, high gravel volumes, and
presence of large quantities of mineral carbonates were the most
commonly occurring features of the A. nova soils.

Acknowledgments

This paper is based on a portion of a thesis prepared for the
Master of Science degree in range science by the senior author. Fi-
nancial support was provided by Regional Project W-89, Characteri-
zation of Habitat Types on Sagebrush Ranges. The authors are
grateful to Dr. G. H. Ward, Knox College, Galesburg, Illinois, and
Dr. A. A. Beetle, University of Wyoming, Laramie, for verifying
identification of Artemisia specimens and to Mr. L. N. Langan, Soil
Conservation Service, Reno, Nevada, for assistance in identification
of the soils.

Literature Cited

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land zone in the White Mountains of California. Amer. Midi. Natur. 73:225-
239.

Beetle, A. A. 1960. A study of sagebrush, the section Tridentata of Artemisia.
Wyoming Agr. Exp. Sta. Bull. 368.

Billings, W. D., 1951. Vegetation zonation in the Great Basin of western
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Blackburn, W. H., P. T. Tueller, and R. E. Eckert, Jr. 1968a. Vegetation
and soils of the Duckwater Watershed. Agr. Exp. Sta., Univ. of Nev., R40.

. 1968b. Vegetation and soils of the Crowley Creek Watershed. Agr. Exp.

Sta., Univ. of Nev., R42. , , .

1968c. Vegetation and soils of the Mill Creek Watershed. Agr. Exp.

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. 1969a. Vegetation and soils of the Churchill Canyon Watershed. Agr.

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1969b. Vegetation and soils of the Pine and Mathews Canyon water-

sheds. Agr. Exp. Sta., Univ. of Nev., R46.
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and soils of the Coils Creek Watershed. Agr. Exp. Sta., Univ. of Nev.. R48.
1969b. Vegetation and soils of the Cow Creek Watershed. Agr. Exp.

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the eastcentral Owyhee Uplands of Oregon. M.S. Thesis. Oregon State Univ.,

Corvallis. , , r i u

Daubenmire, R. 1968. Plant communities: A textbook of synecology. Harper

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Dealy, J. E. 1971. Habitat characteristics of the Silver Lake mule deer range.

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in northern Harney and Lake counties, Oregon. Ph.D. Thesis. Oregon State
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Fautin, R. W. 1946. Biotic communities of the northern desert shrub biome in
western Utah. Ecol. Monogr. 16:251-310.

FosBERG. M. A., AND M. HiRONAKA. 1964. Soil properties affecting the distri-
bution of big and low sagebrush communities in southern Idaho, pp. 230-
236. In Forage plant physiology and soil-range relationships. Amer. Soc.
Agron. Spec. Pub. No. 5.

Hall, F. C. 1967. Vegetation-soil relations as a basis for resource manage-
ment on the Ochoco National Forest of Central Oregon. Ph.D. Thesis. Oregon
State Univ., Corvallis.

HiRONAKA, M. 1963. Plant-environment relations of major species in the sage-
brush-grass vegetation of southern Idaho. Ph.D. Thesis. Univ. of Wisconsin,
Madison.

Holmgren, A. H., and J. L. Reveal. 1966. Checklist of the vascular plants of
the intermontane region. U. S. Forest Service Res. Paper INT-32, Inter-
montane Forest and Range Exp. Sta., Ogden, Utah.

Hugie, V. K., H. B. Passey, and E. W. Williams. 1964. Soil taxonomic units
and potential plant community relationships in a pristine range area of
southern Idaho, pp. 190-205. In Forage plant physiology and soil-range re-
lationships. Amer. Soc. Agron. Spec. Pub. 5.

Passey, H. B., and V. K. Hugie. 1962. Sagebrush on relict ranges in the
Snake River Plains and northern Great Basin. J. Range Mgt. 15:273-278.

Richards, W. H., and J. C. Beatley. 1965. Canopy-coverage of the desert shrub
vegetation mosaic of the Nevada Test Site. Ecology 46:524-529.

Robertson, D. R., J. L. Nielsen, and N. H. Bare. 1966. Vegetation and soils
of alkali sagebrush and adjacent big sagebrush ranges in North Park, Colora-
do. J. Range Mgt. 19:17-20.

Shantz, H. L. 1925. Plant communities in Utah and Nevada, p. 15-23. In
Tidestrom, I., Flora of Utah and Nevada. Contri. U.S. Nat. Herb. 25.

Shreve, F. 1942. The desert vegetation of North America. Bot. Rev. 8:195-246.

Soil Survey Staff. Soil Conservation Service. 1967. Supplement to Soil Clas-
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SuMMERFiELD, H. B., Jr. 1969. Pedological factors related to the occurrence
of big and low sagebrush in northern Washoe County, Nevada. M.S. Thesis.
Univ. of Nevada, Reno.

Thatcher, A. P. 1959. Distribution of sagebrush as related to site differences
in Albany County, Wyoming. J. Range Mgt. 12:55-61.

TiSDALE, E. W., M. HiRONAKA, AND M. A. FosBERG. 1965. An area of pristine
vegetation in Craters of the Moon National Monument, Idaho. Ecology
46:349-352.

TuELLER, P. T. 1962. Plant succession on two Artemisia habitat-types in south-
eastern Oregon. Ph.D. Thesis. Oregon State Univ., Corvallis.

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taxonomic study. Contri. from the Dudley Herb. 4(part 6): 155-205.

SIGNIFICANT BIRD RECORDS FROM UTAH

William H. Behlei

Abstract. — New or additional records of six species of birds are reported for
Utah: Red-throated Loon, Black-legged Kittiwake, Vaux's Swift, Blue Jay, Mag-
nolia Warbler, and Rose-breasted Grosbeak. All are substantiated by museum
specimens. Unless reported for the first time, other records for the state for these
rare birds are summarized.

Three birds either new or rare in Utah were taken in the spring
of 1972 by a field party from the University of Utah to the Beaver
Dam Wash in extreme southwestern Utah. In addition, specimens
of three birds new to the state have been submitted to the museum in
recent years by interested individuals. These six kinds are hereby
placed on record in order to update the checklist of the birds of the
state.

Red-throated Loon

A loon of this species, Gavia stellata^ was first seen at the Bear
River Migratory Bird Refuge by refuge personnel on 28 July 1973.
It remained several days and was observed by many others. Becom-
ing languid, it was eventually captured and taken to headquarters
for observation, but it died on 21 August. An autopsy revealed that
a catfish spine was lodged in the esophagus and had perforated the
wall. By implication, the bird died from starvation. It was emaci-
ated. However, the bird was also host to many endoparasites. It
was a male. The specimen was presented to the University of Utah
by Lloyd Gunther, refuge manager, and is now a study skin in the
collection.

Black-legged Kittiwake

A male of Rissa tridactyla tridactyla was picked up dead, although
in fresh condition, at the Fish Springs National Wildlife Refuge, 40
miles southwest of Dugway, Juab County, Utah, on 12 March 1972,
by a trapper, Jim Harrison, and was relayed to the University by
John Venegoni and Lloyd Gunther, refuge managers. The specimen
weighed 265 g and the testes measured 6x2 mm. This is the only
record of the species for Utah; hence, it is of accidental status.

Vaux's Swift

A female of Chaetura vauxi vauxi was taken at the Terry Ranch
in the Beaver Dam Wash, 2500 ft elevation, five miles north of the
Utah- Arizona border, Washington County, Utah, on 19 May 1972,
by William Buntin, M.D. It was a lone bird. The ova measured 0.5
mm in diameter. This is the first unquestionable specimen of record
for the state. Woodbury et al. (1949:18) note the existence of a

^Department of Biology, University of Utah, Salt Lake City 84112.

243

244 GREAT BASIN NATURALIST Vol. 33, No. 4

miunmified skin found on 28 October 1939 in a stove in a cabin at
the New State Gun Club or Jordan Fur Farm, Davis County, near
the mouth of the Jordan River. The bird had presumably been
trapped there sometime during the late spring or summer of 1939.

In addition to these two specimens there are several sight records
for the species. Two were seen in a flock of other kinds of swifts
and swallows on 10 May 1959 at Provona Beach, Utah Lake, near
the mouth of the Provo River (National Audubon Society, 1959: 391 ) .
One was seen at Utah Lake on 12 May 1963 (National Audubon
Society, 1963b:422). Three were seen at the Gap, near Parowan,
on 31 August 1964 (National Audubon Society, 1965:64), and three
were seen at Springdale near the mouth of Zion Canyon from 11 to
15 September 1965 (National Audubon Society, 1966:77). Thus it
appears that the species is a regular but uncommon transient through
the state in both spring and autumn.

Blue Jay

The first specimen of Cyanocitta cristata cyanotcphra from the
state, a female, was taken by John Bushman at Holladay, Salt Lake
County, Utah, 4500 ft elevation, on 30 April 1970. A jay of this
species, possibly the same individual, had been observed in scrub
oaks in the same area off and on since 5 January 1969. Another
example was observed in the area during the winter of 1970-71.
There are some earlier sight records for Utah, one by Dr. and Mrs.
Carl L. Hubbs {in lift.) on 25 June 1950 on the Markagunt Plateau,
Iron County, beside State Highway 143 near its junction with the
road leading to the summit of an eminence known as Brian Head.
The location was in coniferous forest immediately north of the
northern boundary of Cedar Breaks National Monument at an ele-
vation of about 10,000 ft. Another sighting in the same general vi-
cinity was made by the late Stuart Murie (National Audubon
Society, 1967:63) on 29 October 1966 at a locahty that he simply
designated as Cedar Mountain east of Cedar (^ity.

Magnolia Warbler

A male of Dcndroicn magnolia was taken by the writer in a low
desert shrub at the Terry Ranch, 2500 ft elevation, Beaver Dam
Wash, five miles north of the Utah-Arizona border, Washington
County, Utah, on 19 May 1972. It weighed 6.9 g, the testes mea-
sured 5x4 mm, and the bird was moderately fat. Several other
species of migrating warblers were seen, but this was the only one
of this species. While this is the first specimen from Utah, the spe-
cies has been seen several times before. Two were observed at Salt
Lake City on 14 October 1962 (National Audubon Society,
1963a: 54); one at Green River, Emery County, Utah, on 30 May
1968 (National Audubon Society, 1968:561); and some at the Bear
River Refuge, Box Elder County, on 26 September 1971, (National
Audubon Society, 1972:98). The species therefore appears to be an
uncommon migrant.

Dec. 1973 behle: utah birds 245

Rose-breasted Grosbeak

A fourth record specimen of Pheucticus ludovicicmus for Utah
was obtained by WiUiam Buntin, M.D., at the Terry Ranch, 2500
ft elevation, Beaver Dam Wash, five miles north of the Utah- Arizona
border on 19 May 1972. It was a male with testes that measured
10 X 8 mm. It was the male of a pair. Records of the species in
Utah have been slow in accumulating until recent years, but it now
seems that the species is of casual occurrence in the state and may
breed, at least upon occasion. The first record was an observation
of the species at Kanab on 26 April 1935 (Behle, et al, 1958). A
second record, not heretofore recorded, pertains to a juvenile with
soft mandibles and conspicuous yellow gape trapped by Calvin D.
Wilson of the Tracy Aviary at his home in South Salt Lake on 4
August 1955. It was kept in a cage of native birds at the aviary
until it died on 12 December. It was not saved.

The first specimen from the state was obtained by Roland Wauer
(Wauer and Carter, 1965:78) at the Springdale Ponds, Washington
County, near the entrance to Zion National Park on 3 May 1965. It
is in the collection at the park. Wauer also saw one at Kanab (oral
report) on 7 June 1965. A second speciment from Utah now in the
collection at the University of Utah was killed by flying into a win-
dow at the residence area of Arches National Park near Moab, Grand
County, on 26 May 1965 (Behle, 1966:396). The third specimen
from the state, also in the University of Utah collection, was found
dead at Fish Springs National Wildlife Refuge on 2 June 1965
(ibid.). Two days later another was seen there but not taken.

Literature Cited

Behle, W. H., J. B. Bushman, and C. M. Greenhalgh. 1958. Birds of the
Kanab area and adjacent high plateaus of southern Utah. Univ. Utah Biol.
Ser. 11 (7): 1-92.

Behle, W. H. 1966. Noteworthy records of Utah birds. Condor 68(4) : 396-397.

National Audubon Society. 1959. Great Basin, central Rocky Mountains
regional report. Audubon Field Notes 13(4) : 390-391.

. 1963a. Great Basin, central Rocky Mountains regional i-eport. Audu-
bon Field Notes 17(l):53-54.

. 1963b. Great Basin, central Rocky Mountains regional report. Audu-
bon Field Notes 17(4) : 422-423.

. 1965. Great Basin, central Rocky Mountains regional report. Audu-
bon Field Notes 19(1): 63-64.

. 1966. Great Basin, central Rocky Mountains regional report. Audu-
bon Field Notes 20(1): 76-77.

. 1967. Great Basin, central Rocky Mountains regional report. Audu-
bon Field Notes 21 (l):62-64.

1968. Great Basin, central Rocky Mountains regional report. Audu-

bon Field Notes 22(4) :560-562.
, 1972. Great Basin, central Rocky Mountains regional report. American

Birds. 26(1) -96-98. . . ^ ,

Wauer, R. H., and D. L. Carter. 1965. Birds of Zion National Park and

vicinity. Zion Nat. Hist. Ass'n.:l-92.
Woodbury, A. M., C. Cottam, and J. W. Sugden. 1949. Annotated checklist

of the birds of Utah. Univ. Utah Biol. Ser. 11 (2): 1-40.

SMALL MAMMALS OF THE NATIONAL REACTOR
TESTING STATION, IDAHO^

Dorald M. Allred=

Abstract. — During studies of ectoparasites in 12 plant communities in 1966
and 1967, five types of traps were used to capture 2,478 mammals of the follow-
ing 1 1 species: Dipodomys ordii, Euiamias minimus, Microtus montanus,
Onychomys leucogaster, Perognathus parvus, Peromyscus maniculatus, Reithro-
dontomys megalotis, Sorex merriami, Spermophilus townsendii, Neotoma cinerea,
and Thomomys talpoides. The most abundant species was D. ordii and the least,
M. montanus. Plant communities which contained the greatest number of species
were the Chrysothamnus-Artemisia and Chrysothamnus-grass Tetradymia. Fewest
species were found in the grass and Juniperus communities. Greatest populations
were in the Juniperus and grass communities, and lowest populations in the
Artemisia-Chrysothamnus, Artemisia- Atriplex, and Chrysothamnus-grass-Tetrady-
mia associations.

Between June 1966 and September 1967, ectoparasites were col-
lected from mammals at the National Reactor Testing Station by
personnel of Brigham Young University. The ectoparasites taken in
those studies were reported by Allred (1968a, 1968b, 1970, 1971).
In his 1968 report, AlLred described the physical and ecological
characteristics of the station, specific techniques used, and location
of the study sites.

The station, situated in southeastern Idaho approximately 30
miles west of Idaho Falls, is in the Snake River Section of the Colum-
bia River Basin. The vegetation is typical of the cool, northern
desert shrub biome and is characterized predominantly by sagebrush
(Artemisia), rabbitbrush (Chrysothamnus), and grasses of several
genera.

Five types of traps were utilized to collect the small mammals:
(1) the YAW live-catch trap, 15 inches long by 4^2 inches square
and made of three-mesh galvanized hardware cloth with a galvanized
sheet metal door and reinforcement bands; (2) a modified Hubbard
trap, 8 inches long and 3 inches square and made of galvanized
sheet metal; (3) the can pit trap, which consists of an outer galvan-
ized metal case 7 inches in diameter and 14 inches long, with a
stainless steel, flanged inner can of slightly smaller size; (4) the
Museum Special break-back trap; and (5) the California gopher
trap. YAW, Museum Special, and Hubbard traps were baited with
rolled oats; other traps were not baited. The latter traps were ar-
ranged in a radiating pattern (refer to Allred, 1968), with each
ty])e of trap arranged in two lines, 10 stations in each line, each sta-
tion 10 meters apart with two traps per station (except pit cans,
which consisted of one line with one can per station). All traps
were operated simultaneously for a three-night period once each
month.

'BYU-AEC Report No. COO-1559-11.

-Department of Zoology, Brigham Young University, Provo, Utah 84602.

246

Dec. 1973 allred: idaho mammals 247

This paper describes the ecological distribution and relative
abundance of the 2,478 small mammals of 11 species trapped in 12
different plant communities (Tables 1-3).

Results

Dipodomys ordii was much more abundant in the juniper com-
munity where an understory of Chrysothamnus and Eurotia was
present, and in the grass community than in other communities.
In these two communities almost all other species of small mam-
mals were present only in minimal numbers compared to other
communities of the station where the species were found. Two
species, Eutamias minimus and Peromyscus maniculatus, occurred
in next to their lowest populations in the /i/n//9^rw5-mixed-understory
association. D. ordii was found in all 12 communities, but popula-
tions were lowest in the Artemisia-Chrysothamnus— grass associa-
tion. These kangaroo rats were the next to most abundant mammal
in five communities.

Eutamias minimus was most abundant in the Chrysothamnus-
Artemisia-grass association but was not found where grasses were
predominant. These chipmunks were present in all other communi-
ties, although their populations were lowest in the Chenopodium-
Eurotia, Chrysothamnus-grass-forh, and J uni per us-mixed-under-
story associations. They were the third most abundant species in
five of the communities.

Microtus montanus was found only in the Chrysothamnus-
Artemisia and Elymus-iorh associations, and only in small numbers.

Onychomys leucogaster was found most abundantly in the
Chrysothamnus-Artemisia association; in small numbers in the
grass, Chenopodium-Eurotia, and /wAzzp^rw^-mixed-understory as-
sociations; but was absent in the Elymus-iorh, Juniperus (no under-

Table 1. Vegetative composition and cover in 12 communities at the Na-
tional Reactor Testing Station, Idaho.

Study

site and percentage^

cover

Plant taxon

Artemisia

Atriplex

Chenopodium

Chrysothamnus

Elymus

Eurotia

Juniperus

Opuntia

Salsola

Stipa

Tetradymia

Forbs, misc

Grasses, misc.

Shrubs, misc.

Bare ground

'Nearest whole percent.

248

GREAT BASIN NATURALIST

Vol. 33, No. 4

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Dec. 1973

ALLRED: IDAHO MAMMALS

249

Table 3. Sequence of occurrence based on relative abundance index' of
some small mammals within the same plant community at the National Reactor
Testing Station, Idaho.

Plant cormnunity" Relative

abun-

and animal species dance

index'

Artemisia (site 10)

Peromyscus maniculatus

10.0

Dipodomys ordii

8.7

Perognathus parvus

7.9

Eutamias minimus

4.4

Onychomys leucogaster

1.6

Reithrodontomys megalotis

1.0

Artemisia-Atriplex (site 8)

Peromyscus maniculatus

57.3

Perognathus parvus

14.5

Dipodomys ordii

13.3

Eutamias minimus

2.5

Spermophilus townsendii

1.0

Artemisia-Chrysothamnus-

Grass (site 2)

Peromyscus maniculatus

87.0

Perognathus parvus

67.0

Eutamias minimus

41.0

Onychomys leucogaster

12.0

Spermophilus townsendii

8.0

Reithrodontomys megalotis

2.0

Dipodomys ordii

1.0

Chenopodium-Eurotia (site 9)

Peromyscus maniculatus

85.0

Dipodomys ordii

69.0

Spermophilus townsendii

13.0

Perognathus parvus

12.0

Eutamias minimus

2.0

Onychomys leucogaster

1.0

Chry sot hamnus- Artemisia (site 7)

Peromyscus maniculatus

153.0

Perognathus parvus

66.0

Eutamias minimus

22.0

Dipodomys ordii

10.0

Onychomys leucogaster

9.0

Spermophilus townsendii

6.0

Reithrodontomys megalotis

2.0

Sorex merriami

1.0

Chrysothamnus-Artemisia-

Grass (site 1)

Peromyscus maniculatus

123.0

Perognathus parvus

89.0

Eutamias minimus

77.0

Dipodomys ordii

27.0

Onychomys leucogaster

21.0

Microtus montanus

5.0

Reithrodontomys megalotis

2.0

Sorex merriami

1.0

Spermophilus townsendii

1.0

Chrysothamnus-Grass-

Tetradymia (site 11)

Peromyscus maniculatus

68.0

Dipodomys ordii

51.0

Onychomys leucogaster

9.0

Eutamias minimus

4.0

Perognathus parvus

4.0

Sorex merriami

2.0

Reithrodontomys megalotis

1.0

Spermophilus townsendii

1.0

Chrysothamnus-Tetradymia-

Artemisia (site 6)

Peromyscus maniculatus

147.0

Dipodomys ordii

51.0

Perognathus parvus

9.0

Eutamias minimus

8.0

Sorex merriami

3.0

Spermophilus townsendii

1.0

Elymus-Fovhs (site 3)

Peromyscus maniculatus

117.8

Perognathus parvus

9.8

Eutamias minimus

4.3

Dipodomys ordii

, 3.0

Spermophilus townsendii

2.8

Reithrodontomys megalotis

1.3

Microtus montanus

1.0

Juniper us (site 5)

Peromyscus maniculatus

4.6

Dipodomys ordii

4.0

Eutamias minimus

1.3

Perognathus parvus

1.0

Juniperus-Chrysothamnus-

Eurotia (site 12)

Dipodomys ordii

180.0

Peromyscus maniculatus

32.0

Eutamias minimus

4.0

Onychomys leucogaster

1.0

Sorex merriami

1.0

Oryzopsis-Stipa (site 4)

Dipodomys ordii

143.0

Peromyscus maniculatus

13.0

Perognathus parvus

3.0

Onychomys leucogaster

1.0

^With 1 as the minimum number, the higher the index the greater the population.
^The predominant plant in each commimity is listed first, but others listed are also relatively
abundant.

Story), Chrysothamnus- — Artemisia, and Artemisia — Atriplex com-
munities.

Perognathus parvus was present in all of the commmiities ex-
cept the Juniperus-TmxeA-wcvAeTsX.OTj association. Its populations
were highest in the Chrysothamnus- Artemisia and lowest in the

250 GREAT BASIN NATURALIST Vol. 33, No. 4

grass and Chrysothamnus— grass— Tetradymia associations. It was
the second most abundant species in five communities.

Peromyscus maniculatus was present in all 12 communities. It
was most abundant in the Elymus-iorh association and least abun-
dant in the grass community. It was the most abundant species in
10 of the communities and the second most abundant in two.

Reithrodontomys megalotis was most abundant in the Artemisia
community but was not common in any of the six communities
where it was found. It was not found in the grass, Juniperus (no
understory) , Chrysothamnus— Tetradymia— Artemisia, Artemisia—
Atriplex, Chenopodium-Eurotia, and /w/2//7^rw5-mixed-understory
associations.

Sorex merriami was most abundant in the Chrysothamnus-
Artemisia-Tetradymia association. Although not abundant in any
of the communities, it was also present in the Chrysothamnus-Arte-
misia, Chrysothamnus-gvass— Tetradymia, and Juniperus-mixed-
understory associations.

Spermophilus townsendii was most abundant in the Chenopo-
dium-Eurotia association. It was not found in the grass, Juniperus
(no understory), Artemisia, and /w/2z^^A-w5-mixed-understory asso-
ciations. Its lowest numbers occurred in the Chrysothamnus-Arte-
misia and Chrysothamnus— grass-Tetradymia communities. It was
the least abundant mammal in four of the communities in which it
was found.

Neotoma cinerea, the bushy-tailed wood rat, was collected once
in the Juniperus (no understory) community, but in all other col-
lections it was taken near or in volcanic caves and outcroppings.

Thomomys talpoides, the northern pocket gopher, was found
infrequently in the Chrysothamnus-Artemisia, Artemisia-Chryso-
thamnus, Chenopodium-Eurotia, and 7w«//?erw5-mixed-understory
associations.

The communities that contained the greatest number of species
were the Chrysothamnus-Artemisia and C hrysothamnus-grass-Te-
tradymia. Fewest species were found in the grass and Juniperus
(no understory) communities.

Greatest populations of small mammals were found in the Juni-
/7^rw5-mixed-understory and grass communities, and fewest mam-
mals in the Artemisia-Chrysothamnus, Chrysothamnus-Artemisia,
Artemisia- Atriplex, and C hry sot hamnus-grass— Tetradymia associa-
tions.

References

Allred, D. M. 1968a. Ticks of the National Reactor Testing Station. Brigham
Young Univ. Sci. Bull. 10(1).

. 1968b. Fleas of the National Reactor Testing Station. Great Basin Nat.

28(2):73-87.
. 1970. Mites and lice of the National Reactor Testing Station. Brig-
ham Young Univ. Sci. Bull. 12(1).

1971. Mammalian ectoparasite consortism at the National Reactor

Testing Station. Great Basin Nat. 31 (2): 77-82.

SCORPIONS OF THE NATIONAL
REACTOR TESTING STATION, IDAHO^

Dorald M. Allred'

Abstract. — During ecological studies in 1966 and 1967, 282 scorpions of
Paruroctonus boreus (Girard) were collected in can pit-traps in 12 plant commu-
nities. The ideal habitat was in vegetated areas where the total ground cover
was more than 85 percent, of which at least 60 percent was broad-leaf shrubs
and 5 percent grasses. Populations of ants and spiders were highest where
scorpions were most abundant. Scorpions were most active above ground during
August, with about half as much activit}" during July. None were found prior to
May or after September.

Ecological investigations of ectoparasites and other arthropods of
the National Reactor Testing Station were conducted by personnel
of Brigham Young University from June 1966 to September 1967.
Can pit traps that were used to trap ground-dwelling animals cap-
tured 282 scorpions of the species Paruroctonus boreus (Girard) in
12 plant communities. Some ecological data on these arachnids are
reported herein. I am grateful to John Johnson for the identification
of most of the specimens.

The National Reactor Testing Station, situated in southeastern
Idaho approximately 30 miles west of Idaho Falls, is in the Snake
River section of the Columbia River Basin. The vegetation is typical
of the cool, northern desert shrub biome and is characterized pre-
dominantly by sagebrush {Artemisia tridentata), rabbitbrush
(Chrysothamnus) , and grasses of the genera Agropyron, Elymus,
Oryzopsis, and Stipa. Allred (1968) described the ecological aspects
of the station and defined and pictured the 12 study sites included
here. The vegetative cover of each of these study sites is summarized
below. For further details on the vegetation of the station, consult
Atwood (1970).

Site 1: Chrysothamnus 38%, Artemisia 24%, grasses 15%, forbs 10.5%, Opun-
tia 6%, Tetradymia 1.5%, bare ground 5%.

Site 2: Artemisia 50%, Chrysothamnus 15%, grasses 15%, Eurotia 10%, forbs
7%, bare ground 3%.

Site 3: Elymus 50%, forbs 35%, bare ground 15%.

Site 4: Oryzopsis and Stipa 90%, shrubs 5%, forbs 5%.

Site 5; Juniperus 40%, forbs 1%, bare ground 59%.

Site 6: Chrysothamnus 30%, Tetradymia 30%, Artemisia 20%, grasses
5%, forbs 5%, bare ground 10%.

Site 7: Chrysothamnus 41%, Artemisia 30%, Eurotia 9%, forbs and grasses
4%, bare ground 16%.

Site 8: Artemisia 30%, Atriplex 26%, Chrysothamnus 5%, forbs and grasses
1%, bare ground 38%.

Site 9: Chenopodium 40%, Eurotia 35%, Artemisia and Chrysothamnus 2%,
Salsola 2%, forbs 1%, bare ground 20%.

Site 10: Artemisia 68%, Opuntia 7%, grasses 7%, forbs 5%, bare ground 13%.

iBYU-AEC Report No. COO-1559-10. .

^Department of Zoology, Brigham Young Umversity, Provo, Utah 84602.

251

252 GREAT BASIN NATURALIST Vol. 33, No. 4

Site 11: Chrysothamnus 30%, grasses 26%, Tetradymia 22%, Artemisia
5%, Opuntia 5%, forbs, Atriplex 2%, bare ground 5%.

5//g /2.- Juniperus 30%. Chrysothamnus 15%, Eurolia 15%, Artemisia 13%,
grasses 10%, forbs 6%, Opuntia 1%, bare ground 10%.

Ten can pit traps spaced at 15-nieter intervals were placed in a
row at each site. These were open continuously from April to No-
vember. Cans were visited each day of a three-day period approxi-
mately every two weeks to collect the trapped animals. These visits
were correlated with the times that other traps which were designed
to capture rodents were in operation on each study site.

Discussion

Species distribution. — The occurrence of Paruroctonus boreus
at the station is not unusual. This "northern scorpion," as it was des-
ignated by Ewing (1928), occurs from the Mojave Desert of Califor-
nia and southwestern Nevada northward into the Great Basin of Ne-
vada, Utah, and Idaho; in the Upper Colorado River Basin of Ari-
zona, Colorado, and Utah; in the Snake River and Columbia River
basins of Idaho, Oregon, and Washington; in the Missouri River
Basin of Montana, and in the southern part of Canada (Gertsch and
Soleglad, 1966). Although this was the only species found at this
northern latitude, Gertsch and AUred (1965) indicated that Hodru-
rus spadix occurs in eastern Oregon in the Columbia River Basin.
Williams (1970a) also noted the occurrence of //. spadix in south-
western Idaho. Johnson and AUred (1972) indicated that the ranges
of Anuroctonus phaeodactylus and Vaejovis confusus extend into
northern Utah, and Anderson (personal correspondence) stated that
A. phaeodactylus occurs in Idaho. Apparently these three species do
not occur at the station, although perhaps they occupy a much differ-
ent habitat than typified by the twelve study sites investigated and
consequently were not captured in these studios.

Habitat preference. — Williams (1970b) found that the rela-
tive abundance of scorpions was related to habitat type. He observed
that Vaejovis confusus, for example, was predominant in nonrocky,
fine-textured soils, whereas some other s[)ecics occurred jirimarily in
rocky habitats. Undoubtedly the presence and relative abundance
of scorpions arc determined by edaphic factors as well as the vegeta-
tive components with their associated arthropods which serve as
available food. At the Nevada Test Site, Gertsch and Allred (1965)
found scorpions of Paruroctonus boreus most abundant in the Pin-
yon-Juniper community. They were seldom found in other plant
types tyi)ical of the Mojave Desert but did occur somewhat abun-
dantly in the Artemisia community typical of the northern deserts.

At the National Reactor Testing Station, scorpions of P. boreus
varied significantly in abundance among the different plant com-
munities ('{"able 1). In four of the communities none were found,
and in two other communities only one and four specimens were
taken, respectively. The ideal habitat for this species at the station

Dec. 1973 allred: idaho scorpions 253

is in those vegetated areas where the total ground cover is more than
85 percent, of which about 60 percent is broad-leaf shrubs, and where
grasses constitute at least 5 percent. The presence of forbs apparent-
ly is not influential to a significant degree. In all areas where popu-
lations of scorpions were highest, the total ground cover was above
85 percent. However, one exception where no scorpions were found
was Site 4, where the cover was 100 percent but where 90 percent of
this was grass with no shrubs present. In these same high-popula-
tion areas (except the 90 percent grass area mentioned), the cover-
age by broad-leaf shrubs was above 60 percent, with two exceptions:
Site 8, with a 61 percent shrub coverage where no grass occurred;
and Site 7, with a 71 percent shrub coverage where grass coverage
was less than 2 percent.

With one exception, populations of ants and spiders were also
highest at those sites where scorpions were most abundant (Allred,
1969; Allred and Cole, 1971). On sites 4, 5, 8, and 9, where scor-
pions were not found, spiders were essentially absent from sites 4,
5, and 9, and ants were absent from sites 5 and 9.

Sex ratios. — Gertsch and Allred (1965) used can pit traps al-
most exclusively to capture scorpions in Nevada. The sex ratio of
males to females collected there was 6 to 1. Johnson and Allred
(1972) used an ultraviolet light for most of their collections and
found the male to female ratio to be 1 to 1.9. In this study in Idaho,
which utilized pit traps exclusively, the ratio was 5.5 to 1. From
these data I assume that the males wander much more extensively
than do the females and consequently are more apt to fall into the
pit cans. Johnson and Allred (1972) noted that the females of
Anuroctonus phaeodactylus were seen only at the entrances to their
burrows. This same phenomenon of limited movement by the fe-
males, at least for most periods of the year, may also be applicable to
Paruroctonus horeus in Idaho.

Table 1. Seasonal occurrence and abundance of Paruroctonus boreus at 12
sites at the National Reactor Testing Station during 1966 and 1967.

No.

specimens

Site

May*

Jun.

Jul.

Aug.

Sep.

Total

163

282

•Numbers for May are doubled, inasmuch as traps for that month operated only during 1967.

254 GREAT BASIN NATURALIST Vol. 33, No. 4

Seasonal occurrence. — At the National Reactor Testing Sta-
tion, cold temperatures prevail from October to April, and snow fre-
quently lies on the ground for much of that period. Scorpions were
most active above ground during August, with about half as much ac-
tivity during July (Table 1 ) . None were found prior to May or after
September. Immatures were found all summer except in May, and
females were found from May to September (Table 2).

Table 2. Total numbers of Paruroctonus boreus taken seasonally at the
National Reactor Testing Station during 1966 and 1967.

Stage

No.

specimens

or sex

May*

Jun.

Jul.

Aug.

Sep.

Total

Immature

Male

Female

4
6

9
9

61
11

150

3
4
6

224

•Numbers for May are doubled, inasmuch as traps for that month operated only during 1967.

References

Allred, D. M. 1968. Ticks of the National Reactor Testing Station. Brigham

Young Univ. Sci. Bull., Biol. Ser. 10(1).

1969. Spiders of the National Reactor Testing Station. Great Basin

Nat. 29:105-108.
Allred, D. M., and A. C. Cole, Jr. 1971. Ants of the National Reactor Test-
ing Station. Great Basin Nat. 31:237-242.
Atwood, N. D 1970. Flora of the National Reactor Testing Station. Brigham

Young Univ. Sci. Bull., Biol. Ser. 11(4).
EwiNG, H. E. 1928. The scorpions of the western part of the United States,

with notes on those occuning in northern Mexico. Proc. U.S. Nat. Mus.

73 (9): 1-24.
Gertsch, W. J., AND D. M. Allred. 1965. Scorpions of the Nevada Test Site.

Brigham Young Univ. Sci. Bull., Biol. Ser. 6(4).
Gertsch, W. J., and M. Soleglad. 1966. The scorpions of the Vaejovis boreus

group (subgenus Paruroctonus) in North America. American Mus. Nov.

2278:1-54.
Johnson, J. D., and D. M. Allred. 1972. Scorpions of Utah. Great Basin Nat.

32:154-170.
Williams, S. C. 1970a. A systematic revision of the giant hairy-scorpion genus

Hadrurus. California Acad. Sci., Occasional Paper No. 87.
. 1970b. Coexistence of desert scorpions by differential habitat preference.

Pan-Pacific Ent. 46:254-267.
. 1972. Four new scorpion species belonging to the genus Paruroctonus.

California Acad. Sci., Occasional Paper No. 94.

DENSITY CHANGES AND

HABITAT AFFINITIES OF RODENTS OF

SHADSCALE AND SAGEBRUSH ASSOCIATIONS

Earl J. Larrison and Donald R. Johnson'

Abstract. — Rodent trapping was conducted in representative northern
Great Basin habitat types for six consecutive years to deteiTnine the magnitude
of density change, the specific habitat affinities, and the effects of habitat altera-
tion on rodent density. Although species responded differently, total rodent
density was greater in depleted shadscale and sagebrush communities than in
comparable pristine sites. However, disturbed sites with nearly pure stands of
weedy introduced annuals supported few rodents.

Several species exhibited abrupt and concurrent yearly changes in density.
The factors causing these changes were not identified. Until this information is
available, we can develop only crude models to predict the direction and magni-
tude of population change.

Population Geologists lack information regarding the magnitude
of density changes in small mammal populations. Ideally, popula-
tion studies should extend over a sufficient time span to determine
not only a mean density but also provide some estimate of the ex-
pected deviations from that mean. We are not aware of such a study
in the Intermountain West. ^^

Ecologists have gained some understanding of the habitat affini-
ties of the more common rodents. The deer mouse {Peromyscus ma-
niculatus) is recognized as eurytopic, while other species are known
to be restricted to a few habitat types. The presettlement habitat af-
finities of most western species may never be fully known because
of land clearing, livestock grazing, and the establishment of weedy
annuals which are maintained in nearly pure stands by periodic
fires.

The senior author began studies of small mammal populations
in southern Idaho in 1951 in an effort to (1) determine the species
representation in specific habitat types, (2) measure the magnitude
of rodent density changes, and (3) determine the effects of habitat
alteration on both species composition and density.

The press of other work prevented regular yearly sampling; how-
ever, trapping data are available for six consecutive years ( 1955-60)
for several common habitat types of the northern Great Basin. Most
of the trapping was conducted near Malta, Cassia County, Idaho,
with additional investigations in Owyhee, Elmore, and Weiser coun-
ties. Sites were usually trapped during the summer months, although
seasonal trapping was conducted in the Raft River Valley in 1957.

We wish to thank Iain Baxter and Edward Francq for field as-
sistance. The University of Idaho's Special Research Fund supported
part of this study.

'Department of Biological Sciences, University of Idaho. Moscow 83843.

255

256 GREAT BASIN NATURALIST Vol. 33, No. 4

Methods

Transect lines of 50 Museum Special traps were set out at 10-
foot intervals and baited with rolled oats. The lines were maintained
three days and checked twice daily to rebait and recover the catch.
Effort was made to trap in stands of homogeneous vegetation. The
habitat types were identified by the dominant plant species present,
including big sagebrush {Artemisia tridentata) , shadscale (Atriplex
confertifolia), wheatgrass (Agropyron cristatum and Agropyron
desertorum), kochia (Kochia americana), greasewood {Sarcohatus
vermiculatus) , and halogeton (Halogeton glomeratus) .

The number, sex, and age group of each capture was recorded.
An index of density (N/100 trap days) was calculated. Trapping
at most sites was made in replication and the results reported as
means. We made no effort to convert indices of abundance to densi-
ty estimates. However, density and the catch from index lines are
closely correlated (Hansson, 1967; Petticrew and Sadlier, 1970). We
believe that these data accurately reflect density changos in these
populations.

Results

Deer mice were the most abundant and ubiquitous species trapped
in the Raft River Valley, often comprising 60-80 percent of the
catch (Table 1). Depletion of shadscale and sagebrush-grass com-
munities favors its increase. Deer mice were about twice as numer-
ous in depleted shadscale stands in the Raft River Valley as in those
in better condition (Table 1). No truly pristine shadscale stands re-
main in the valley after a long history of heavy grazing (Tisdale
and Zappetini, 1953) and insect defoliation (Mackie, 1958). As a
consequence, deer mice remained the most numerous rodent trapped
in depleted stands as well as in those in better condition. In the more
pristine shadscale and winterfat (Eurotia lanata) communities near
Mountain Home, Elmore County, the catch of chisel-tcothod kan-
garoo rats (Dipodomys microps) and Great Rasin pocket mice (Pcro-
gnnthus parvus) regularly exceeded that of deer mice.

Deer mice were 8-9 times more numerous outside a 40-acre
exclosure established in big sagebrush near Rurley, Cassia County
(Table 2). The grass understory outside the exclosure consisted al-
most entirely of cheatgrass (Bromus trrtorum), while that inside
included perennial species such as Agropyron trachycaulum. Stipa
comata, and Sitanion hystrix. The vegetation within the exclosure
had been protected from livestock since 1930 (Piemeisel, 1945).

Deer mice were the most abimdant rodent trapped in saltsage
{Atriplex nuttalli), black sage {Artemisia arbuscula)^ and Utah
juniper {Juniperus utahensis) in the Raft River Valley and in moun-
tain mahogany {Cercocarpus ledifolius) and western juniper (/. oc-
cidentaJis) communities in Owyhee County.

Least chipmunks (Eutamias minimus) were most common in de-
pleted shadscale stands, where their numbers averaged about 10 times
greater than in stands in better condition (Table 3). Although none

Dec. 1973

LARRISON, JOHNSON: RODENT EC:OLOGY

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258 GREAT BASIN NATURALIST Vol. 33, No. 4

were caught inside the Piemeisel exclosure, 1 1 were trapped in the
sagebrush-cheatgrass outside it (Table 2), further evidence that
range depletion favors an increase in its density.

The yellow pine chipmunk {Eutamias amoenus), which occupies
the higher parts of several ranges in the northern Great Basin (Raft
River Mountains, Cache Peak-Mount Harrison, Jarbidge Mountains),
is absent from the Silver City Mountains of Owyhee County, where
it is replaced by Eutamias minimus. The occurrence of the least
chipmunk in a subalpine fir (Abies lasiocarpa) community on Boul-
der Summit confirms its ability to successfully colonize a variety of
habitats in the absence of a sciurid competitor.

Ord kangaroo rats (Dipodomys ordi) were most abundant in
stands of kochia in the Raft River Valley (Table 4) and along road-
sides where disturbed earth provided easy tunneling (Johnson 1961).
It is more abundant on sandy than on gravel substrates (Fautin,
1946; Maxell and Brown, 1968). This species successfully colonizes
wheatgrass seedings (Table 4). Road building has permitted its dis-
persal into broad stretches of shadscale formerly occupied exclusive-
ly by Dipodomys microps.

In the Raft River Valley, chisel-toothed kangaroo rats w^ere 2-3
times more abundant in "healthy" shadscale than in depleted stands
(Table 5). In the more pristine shadscale and winterfat communi-
ties near Mountain Home it was the most common, and sometimes
the only, rodent trapped. Shadscale leaves comprise most of its diet
(Johnson, 1961), and its distribution closely coincides with that of
shadscale in the Intermountain West. Kenagy (1972) has found
that the chisel-shaped lower incisors are used to strip away the
hypersaline epidermis of shadscale leaves; hence, only the less saline
mesophyll is ingested. This species was also common (6.7/100 trap
days) in a hop sage (Grayia spinosa) community west of Malta.

Western harvest mice (Reifhrodontomys mcgaJotis) were most
abundant in seeded stands of wheatgrass (Table 6) and in giant
wild rye {Elymus cinereus) communities of the Raft River Valley.
Further evidence of its affinity for denser grass habitats was mani-
fest at the Piemeisel exclosure (Table 2) and in a Wvoming studv
(Maxell and Brow^n, 1968).

Great Basin pocket mice were most common in seeded stands of
wheatgrass and in big sagebrush-Idaho fescue {Festuca idahoensis)-
bluebunch wheatgrass (Agropyron spicatum) communities. Its num-
bers are reduced in sage-grass habitat types sustaining heavy graz-
ing (Table 2). This species was common (to 7.4/100 trap days) in
a black sage community south of Albion, Cassia County. Although
often considered a semi-desert species, it was trapped on Boulder
Summit. Owyhee County, in subalpine fir (2500 m) and in a sub-
alpine meadow (2750 m) near Lake Cleveland, Cassia County.

Other rodents trapped in smaller numbers included the sage-
brush vole (Lagurus curtaius), the little pocket mouse (Perognathus
longimemhris), and the desert woodrat (Neotoma lepida) . The habi-
tat affinities of these species and those previously discussed are
summarized in Table 7. We also caught grasshopper mice (Ony-

Dec. 1973

LARRISON, JOHNSON: T.ODENT KC:OLOGY

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Vol. 33, No. 4

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Table 7. Relative abundance of rodents in southern Idaho habitat types.
t = abundant (often > 10/100 trap days); § = common (often > 5/100 trap
days); * = present, caught in small numbers; t = rare or absent. Scientific
names abbreviated.

Habitat Type

P.m.

E.m.

D.o.

D.m.

R.m.

P.p.

L.c.

P.I.

N.l.

Greasewood

Salt sage

♦

Healthy shadscale

♦

Depleted shadscale

♦

Hopsage

Winterfat

•

Kochia

•

Halogeton

Sagebrush-cheatgrass

•

♦

•

Sagebrush-perennial

J.
1

grass

Crested wheatgrass

♦

Black sage

•

♦

Mountain mahogany

Juniper-big sage

♦

chomys leucogaster) in almost all habitat types sampled, but its
catch is not reported here because of the likelihood that it shuns
rolled oats as bait. Small numbers of montane {Microtus montanus)
and long-tailed voles (M. longicaudus) were also taken in grassy
habitats throughout southern Idaho. Townsend ground squirrels
{Citellus townsendi) were trapped in shadscale and winterfat com-
munities. A few antelope ground squirrels {Citellus leucurus) were
taken near the Bruneau Sand Dunes, Owyhee County.

Halogeton, an introduced annual chenopod, has invaded large
areas of depleted shadscale in southern Idaho (Tisdale and Zappe-
tini, 1953). Dense stands of halogeton sometimes support large
numbers of deer mice (to 26/100 trap days) and a few of the other
common species.

Former sage-grass habitat types which support weedy annuals
contain few rodents. Heavy grazing and repeated fires have fos-
tered the establishment of nearly pure stands of cheatgrass over
vast areas of southern Idaho (Stewart and Hull, 1949). Such sites
support only a few deer mice and Great Basin pocket mice. Medusa-
head (Elymus caput-medusae) , a more recent invader, has replaced
cheatgrass over nearly three-quarter of a million acres in southern
Idaho (Hironaka, 1961). Nearly pure stands of this annual support
even fewer rodents: only 8 deer mice were trapped in 900 trap
days east of Weiser, Washington County. Former sagebrush-grass
sites near Mountain Home supporting Russian thistle (Salsola kali)
contained few rodents, but we caught considerable numbers
(12.7/100 trap days) of western harvest mice in a stand of Russian
thistle near Bridge, Cassia County. Former sagebrush-grass sites

262 GREAT BASIN NATURALIST Vol. 33, No. 4

supporting stands of tansy mustard {Descurainia pinnata) or pepper-
grass (Lepidium perjoliatum) contain few rodents.

Much of this sagebrush range can be rehabilitated through re-
seeding with drought-resistant wheatgrasses. Although certain grass-
adapted species are more numerous in wheatgrass seedings, the
total rodent catch remains about the same as that on depleted sage-
brush sites.

Discussion

The effects of grazing on rodent abundance has received con-
siderable attention over the past half-century (reviews by Bond,
1945; Howard, 1953). Most of these studies have been conducted
in central California, the Southwest, or on the Great Plains. Our
data corroborate earlier findings that range depletion favors an in-
crease in deer mice populations (Phillips, 1936; Quast, 1948) and
tends to diminish the numbers of western harvest mice (Quast,
1948). We found that Great Basin pocket mice, like two other kinds
of pocket mice in Arizona (Reynolds and Haskell, 1949), were
most abundant in vigorous stands of perennial grasses. Like them, its
numbers were reduced on depleted ranges.

There are interspecific differences in the responses of kangaroo
rat populations to range depletion. While several species are more
abundant on grazed sites (McCulloch, 1962; Reynolds, 1958; Quast,
1948), the numbers of chisel-toothed kangaroo rats are reduced in
depleted shadscale stands (Table 5). Thus it is sometimes hazardous
to generalize, predicting the response of a rodent population to habi-
tat alteration, basing the prediction on the response expected from
a related species.

In an earlier investigation, Fautin (1946:279) found that deer
mice occurred in relatively low numbers in six communities within
the shadscale and sagebrush associations of west central Utah. We
found deer mice the most abundant rodent in all communities in
the Raft River Valley, probably a result of its long history of heavy
use by livestock. Although we found least chipmunks in all the
habitat types trapped in the valley, Fautin found them restricted
to sagebrush sites in Utah. Again, we ascribe this difference to the
range depletion at our trapping sites.

The highest rodent numbers encountered in this study (46.7/100
trap days) were those of deer mice in a big sagebrush community
in the Raft River Valley in May 1964 (Table 1). This level is greatly
exceeded by microtine populations during irruptions (Piper, 1909;
Federal Cooperative Extension Service, n.d.). As hunters, insec-
tivorous and granivorous species such as the deer mouse search
greater distances for food (McNab, 1963), and it is unlikely that
they cannot achieve the high densities found in some foliage herbi-
vore populations.

Rodent populations often exhibit abrupt changes in density
(Horn and Fitch, 1942; Reynolds, 1958). We found that deer mice
populations regularly changed by factors of 2 or 3 and sometimes

Dec. 1973 larrison, johnson: rodknt ec:ology 263

by a factor of 10 from one year to the next (Table 1). The catch
of deer mice was 18/100 trap days in big sagebrush near Malta in
1958. It fell to 1.3/100 trap days in 1959 and then increased to
15.3/100 trap days the following year. Similar changes occurred in
populations at other trapping sites during the same time interval
(Table 1). These concurrent changes in populations of different
species are in contrast to the independent changes occurring in
rodent populations on desert grasslands in New Mexico (Wood,
1965, 1969).

In the Raft River Valley where trapping was conducted on a
bimonthly basis during the 1957 field season, peak densities occurred
in June for most species. The proportion of juvenile mice in the
catch declined steadily, indicating that breeding terminated in late
spring. Rodent populations reached low levels at these sites the
following year.

Our understanding of the dynamics of small mammal popula-
tions will reach maturity only after we are able to identify those
variables which most affect density. We can then develop sensitive
models to predict population change, one of the goals of the Inter-
national Biological Program, Biome Studies.

Literature Cited

Bond, R. M. 1945. Range rodents and plant succession. Trans. N. Amer.

Wildl. Conf. 10:229-234.
Fautin. R. W. 1946. Biotic communities of the northern desert shrub biome

in western Utah. Ecol. Monographs 16:251-310.
Federal Cooperative Extension Service, n.d. The Oregon meadow mouse

irruption of 1957-1958. Oregon State College, Corvallis.
H\NssoN. L. 1967. Ind-^y line catcher as a basis of population studies on small

mammals. Oikos 18:261-276.
Hironaka, M. 1961. The relative rate of root development of cheatgrass and

medusahead. J. Range Mgt. 14:264-267.
Horn, E. E., and H. S. Fitch. 1942. Interrelations of rodents and other wild-
life on the range. In The San Joaquin Range. Calif. Agric. Exp. Sta. Bull.

663:96-129.
Howard, W. E. 1953. Rodent control on California ranges. J. Range Mgt.

6:423-434.
Johnson, D. R. 1961. The food habits of rodents on rangelands of southern

Idaho. Ecology 42:407-410.
Kenagy, G. J. 1972. Saltbush leaves: Excision of hypersalme tissue by a kan-
garoo rat. Science 178:1094-1096.
Mackie, R. a. 1958. A new species of Eumrsia from southern Idaho. Entomol.

Soc. Wash. 60:5-8.
Maxell. M. H., and L. N. Brown. 1968. Ecological distribution of rodents on

the high plains of eastern Wyoming. The Southwestern Naturalist 13:143-

158.
McCuLLOCH. C. Y., Jr. 1962. Populations and range effects of rodents on the

sand sagebrush grasslands of western Oklahoma. Okla. State Univ. Publ. No.

11.
McNab, B. K. 1963. Bioenergetics and the detennination of home range size.

Amer. Nat. 97:133-140.
Petticrew. B. G., and R. M. F. S. Sadlier. 1970. The use of index trap lines

to estimate population numbers of deermice (Peromyscus maniculatus) in a

forest envirorunent in British Columbia. Canad. J. Zool. 48:385-389.

264 GREAT BASIN NATURALIST Vol. 33, No. 4

Phillips, P. 1936. The distribution of rodents in overgrazed and normal grass-
lands of central Oklahoma. Ecology 17:673-679.

PiEMEiSEL, R. L. 1945. Natural replacement of weed hosts of the beet leaf-
hopper as affected by rodents. USDA Circ. No. 739.

Piper, S. E. 1909. The Nevada mouse plague of 1907-08. USDA Farmers
Bull. 352.

QuAST, J. C. 1948. Habitat preferences of rodents on grazed and ungrazed
foothill pastures in California. M.S. Thesis. Univ. Calif., Berkeley.

Reynolds, H. G. 1958. The ecology of the Merriam kangaroo rat (Dipodomys
merriami Meams) on the grazing lands of southern Arizona. Ecol. Mono-
graphs 28: 111-127.

Reynolds, H. G., and H. S. Haskell. 1949. Life history notes on Price and
Bailey pocket mice of southern Arizona. J. Mammal. 30:150-156.

Stewart, G., and A. C. Hull. 1949. Cheatgrass (Bromus tectorum L.) — An
ecological intruder in southern Idaho. Ecology 30:58-74.

Tisdale, E. W.. and G. Zappetini. 1953. Halogeton studies on Idaho ranges.
J. Range Mgt. 6:225-236.

Wood, J. E. 1965. Response of rodent populations to controls. J. Wildlife Mgt.
29:425-438.

Wood, J. E. 1969. Rodent populations and their impact on desert rangelands.
New Mexico State Univ. Agric. Exp. Sta. Bull. 555.

NEW SPECIES OF AMERICAN MICROCORTHYLUS
(COLEOPTERA. SCOLYTIDAE)^

Stephen L. Wood-

Abstract. — Species described include: Microcorthylus demissus and M. in-
validus n. spp. (Mexico); M. debilis and M. vescus n. spp. (Guatemala); M.
pusillus n. spp. (Guatemala and Honduras) ; M. concisus and M. ocularis n. spp.
(Costa Rica); M. inermis, M. lassus, and M. pumilus n. spp. (Costa Rica and
Panama); M. dilutus n. spp. (Colombia); M. contractus, M. curtus, M. diversus,
M. hostilis, M. umbratus n. spp. (Venezuela).

While treating the genus Microcorthylus for my monograph of
North and Central American Scolytidae, I encountered several un-
described species. In order to make names available for identification
and other work, I have described 16 of those species below. Keys
and other aids to identification will be included in the monograph.
The species were taken in the following countries: Mexico (2),
Guatemala (2), Costa Rica (2), Colombia ( 1 ), Venezuela (5), Guate-
mala and Honduras (1), Costa Rica and Panama (3).

Microcorthylus debilis, n. spp.

In this species the frontal punctures are rather coarse and elon-
gate and the elytral declivity is only moderately impressed and de-
void of an elevation between the costal margin and the lateral mar-
gin.

Female. — Length 1.8 mm (paratypes 1.6-1.9 mm), 2.8 times as
long as wide; color yellowish brown.

Frons convex, with a distinct transverse impression just above
epistoma, a weak, median epistomal process indicated; surface ob-
scurely reticulate in central area, becoming almost rugose-reticulate
in marginal areas; punctures coarse for this genus, elongate; sub-
glabrous. Antennal club 1.1 times as long as scape, 1.4 times as long
as wide, broadly somewhat obovate; sutures 1 and 2 almost straight;
posterior face with small tuft of hair.

Pronotum 1.1 times as long as wide; widest near base, sides par-
allel on basal half, broadly rounded in front; about a dozen weakly
raised serrations on anterior margin; indefinite summit just an-
terior to middle; anterior slope moderately steep, rather finely asper-
ate; posterior areas strongly reticulate, sparse, very minute, shallow
punctures indicated. Glabrous.

Elytra 1.6 times as long as wide, 1.5 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, then arcuately
converging to lateral margin of declivity, broadly rounded behind
on median half; strial punctures very obscurely indicated, interstrial
punctures obsolete, discal surface subreticulate, subshining. Declivity
steep, broadly sulcate; impressed area extending from suture to

'This research was supported by grants from the National Science Foundation.
^Department of Zoology, Brigham Young University, Provo, Utah 84602. Scolytidae lontri-
bution No. 51.

265

266 GREAT BASIN NATURALIST Vol. 33, No. 4

about striae 1 above and to striae 3 or 4 below; sutural interstriae
feebly, abruptly elevated, forming a distinct, continuous crest on
its lateral margin on right elytron; lateral margins increase in
height gradually on upper two-thirds, obsolete below, crest of upper
half armed by two pairs of rather widely separated, small, pointed
denticles; all punctures obsolete; surface reticulate. Glabrous.

Male.— Similar to female except epistomal process forming a
definite median tubercle; serrations on anterior margin of pronotum
larger.

Type Locality.— Volcan Pacaya, Esquintla, Guatemala.

Type Material.-— The male holotype, female allotype, and 28
para types were taken at the type locality on l-VI-64, 4000 ft (1300
m). No. 670, from a broken branch 3 cm in diameter by S. L. Wood;
other paratypes bear similar data except 2 are No. 669, and 5 are
653 from a woody vine; 23 paratypes are from Volcan Zunil, Que-
zaltenango, Guatemala, 27-V-64, 3000 ft (1000 m). No. 625, shrub,
S. L. Wood.

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

Microcorthylus demissus n. sp.

This species is distinguished from debilis Wood by the smaller
size, by the more shining, less strongly punctured female frons,
male frons finely reticulate, and by the less strongly impressed
elytral declivity.

Female. — Length 1.5 mm (paratypes 1.3-1.5 mm), 2.9 times as
long as wide; color yellowish brown.

Frons as in debilis except central half smooth, shining, punc-
tures not elongate; antennal club 1.3 times as long as scape, 1.5
times as long as wide.

Pronotum as in debilis except anterior margin more narrow-
ly rounded.

Elytra as in debilis except declivity less strongly impressed,
lateral margins much less strongly elevated.

Male. — Similar to female except frons uniformly, finely reticu-
late; anterior margin of pronotum armed by six rather coarse serra-
tions; declivital denticles slightly larger.

Type Locality. — Six miles (9 km) NE Teziutlan, Puebla, Mex-
ico.

Type Material. — The female holotype, male allotype, and 22
paratypes were taken at the type locality on 2-VII-67, 4800 ft (1500
m). No. 137, Miconia, S. L. Wood; other paratypes taken at the
same locality by me include 11, No. 149, tree branch and 19 taken
27-VI-53, tree branch.

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

Microcorthylus ocularis, n. sp.

This species is distinguished from demissus Wood by the ab-
sence of a median epistomal tubercle, by the narrow frons, by the

Dec. 1973 wood: new scolytidae 267

enlarged, coarsely faceted eyes, and by the slender body form. Very
similar to dilutus Wood.

Female. — Length 1.4 mm (paratypes 1.3-1.4 mm), 3.0 times
as long as wide; color yellowish brown.

Frons as in demissus except mostly reticulate and epistomal
tubercle reduced or absent. Eye greatly enlarged, coarsely faceted,
frons correspondingly narrower as in dilitus. Pronotum and elytra
more strongly reticulate.

Type Locality. — Tapanti, Cartago, Costa Rica.

Type Material. — The female holotype and two female para-
types were taken at the type locality in 2-VII-63, 4000 ft (1300 m).
No. 10, in Conostegia oerstediana, by S. L. Wood. Four female para-
types bear identical data except No. 7B, in Boehmeria ulmifolia; one
paratype is from 9 miles (15 km) SE Cartago, Cartago, Costa Rica,
24-XI-63, 5600 ft (1800 m). No. 24B Siparuna nicaraguensis, S. L.
Wood.

The holotype and paratypes are in my collection.

Microcorthylus inermis, n. sp.

This species is distinguished from demissus Wood by the
more strongly impressed elytral declivity, with the lateral margins
abruptly but not strongly elevated, by the uniformly, finely, rugose-
reticulate frons, and by the broader antennal club.

Female. — Length 1.5 mm (paratypes 1.5-1.6 mm), 2.8 times as
long as wide; color dark brown.

Frons broadly convex, a conspicuous median tubercle just above
epistomal process; entire surface from epistoma to vertex rugose-
reticulate, minute punctures almost obsolete. Antennal club 1.3 times
as long as scape, 1.3 times as long as wide; broadly obovate; a small
tuft of setae on posterior face.

Pronotum 1.1 times as long as wide; as in debilis except poster-
ior areas much more strongly reticulate.

Elytra 1.8 times as long as wide; as in debilis except entire
surface uniformly reticulate; declivity more strongly impressed,
lateral margins abruptly elevated.

Male. — Similar to female except frontal punctures more dis-
tinct; antennal club slightly smaller and devoid of tuft of hair on
posterior face; serrations on anterior margin of pronotum rather
coarse.

Type Locality. — Nine miles (15 km) SE Cartago, Cartago, Costa
Rica.

Type Material. — The female holotype and 14 paratypes were
taken at the type locality on 24-IX-63, 1800 m, No. 199, Siparuna
nicaraguensis, by S. L. Wood; the male allotype and eight paratypes
bear identical data except they are No. 248. Thirteen paratypes are
from Escasu, San Jose, Costa Rica, 2-X-63, 1300 m, either No. 215
in Guazuma ulmifolia or No. 218 in a cut tree seedhng; and four

268 GREAT BASIN NATURALIST Vol. 33, No. 4

para types are from Cerro Punta (labeled Volcan Chiriqui), Chiriqui,

Panama, 11-1-64, 1800 m, Nos. 381, 399, in woody vines; all by me.

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

Microcarthylus invalidus, n. sp.

This species is distinguished from demissus Wood by the more
nearly reticulate frons, by the much more deeply impressed elytral
declivity with its margins more abruptly elevated, and by the re-
ticulate female scutellum.

Female. — Length 1.6 mm (paratypes 1.5-1.6 mm), 2.8 times as
long as wide; color light brown.

Frons as in demissus except reticulation more intensive in
lateral areas. Antennal club 1.2 times as long as wide.

Pronotum 1.2 times as long as wide; as in demissus.

Elytra as in demissus except declivity much more strongly
impressed; declivity about as in inermis Wood except lateral mar-
gins of impressed area converge toward base at an angle of about
30 degrees (about 45 degrees in inermis).

Male. — Similar to female except frons rugose-reticulate, punc-
tures minute; antennal club smaller, without tuft of hair on poster-
ior face; anterior margin rather coarsely serrate.

Type Locality. — Four miles (6 km) W Tepic, Nayarit, Mexico.

Type Material. — The female holotype and male allotype were
taken at the type locality on 13-Vin-65, 1000 m, No. 239, from a
cut branch, by S. L. Wood. Two paratypes bear identical data ex-
cept they are No. 240.

Microcorthylus pumilus, n. sp.

This species differs from all of the preceding forms in having a
continuous, submarginal, subacute line extending from the suture
to the costal margin to the lateral margin of the declivity; it is also
unique in having the female frons very broad, with the mandibles
proportionately lengthened.

Female. — Length L8 mm (L7-1.9 mm), 2.9 times as long as
wide; color very dark brown.

Frons very broad, particularly below, broadly convex; surface
very finely punctured, finely rugose-reticulate to upper level of eyes,
smooth and shining above. Antennal club 1.1 times as long as scape,
1.3 times as long as wide; obovate, widest through suture 2; poster-
ior face without a tuft of long hair.

Pronotum 1.2 times as long as wide; sides almost straight and
parallel on basal half, very broadly rounded in front; anterior mar-
gin weakly serrate; summit indefinite, on anterior third; anterior
third finely asperate; posterior areas strongly reticulate, punctures
minute, shallow. Glabrous.

Elytra 1.8 times as long as wide, 1.7 times as long as pronotum;
sides almost straight and parallel on basal three-fourths, then arcu-

Dec. 1973 wood: new scolytidae 269

ately converging, almost straight behind on median two thirds; strial
punctures obscurely indicated, almost obsolete; surface reticulate.
Dechvity very steep, almost vertical on lower half; rather narrowly
sulcate on upper third, broadly impressed below; punctures obso-
lete, surface reticulate; suture narrowly elevated, lateral margin of
this line subacuate, continued without interruption along costal sub-
margin to lateral margin of declivity; impressed area somewhat
heart-shaped; spines 1 and 2 almost obsolete. Glabrous except for
a few hairlike setae near declivity.

Male. — Similar to female except lower frons normal, not as
broad; antennal club more slender; anterior margin or pronotum
more strongly serrate, 10 serrations.

Type Locality. — Nine miles (15 km) SE Cartago, Cartago, Costa
Rica.

Type Material. — The female holotype, male allotype, and three
para types were taken at the type locality in 3-VII-63, 1800 m. No.
IZB, in cut Conostegia oerstediana branches, by S. L. Wood. One
paratype is from Tapanti, Cartago, Costa Rica, 2-VII-63, 1300 m,
No. 10, same host; and six paratypes are from Cerro Punta (labeled
Volcan Chiriqui), Chiriqui, Panama, 11-1-64, 1800 m. No. 376, from
a tree branch; all were taken by me.

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

Microcorthylus pusillus, n. sp.

This species is distinguished from pumilis Wood by the larger
size, by the very different frons, by the tuft of hair on the
posterior face of the female antenna, and by the larger declivital
denticles.

Female. — Length 2.0 mm (paratypes 2.0-2.4 mm), 2.9 times
as long as wide; color light reddish brown.

Frons with lateral thirds impressed from epistoma to slightly
above eyes, median third strongly elevated, attaining a rather broad
summit at a level slightly below upper level of eyes; surface reticu-
late in impressed areas, smooth and shining on median third and
on entire area above eyes. Antennal club 1.2 times as long as wide;
a small tuft of hair on posterior face.

Pronotum 1.02 times as long as wide; about as in pumilus;
14 serrations on an anterior margin. Glabrous.

Elytra about 1.6 times as long as wide, 1.7 times as long as pro-
notum; about as in pumilus except lower declivity narrower, and
spines 1 and 2 small, but distinctly larger.

Male. — Similar to female except frons broadly, rather evenly
convex; with a short median carina on lower fourth, central area
very slightly elevated, surface reticulate and with moderately coarse,
shallow punctures; serrations on anterior margin of pronotum slight-
ly larger.

Type Locality. — Buenos Aires, Cortez, Honduras.

Type Material. — The female holotype, male allotype, and six

270 GREAT BASIN NATURALIST Vol. 33, No. 4

paratypes were taken at the type locality on 7-V-64, 2300 m, No.
576B, from a tree seedling, by S. L. Wood. Two paratypes are from
Volcan Pacaya, Esquintla, Guatemala, l-VI-64, 1300 m, No. 668, tree
branch, S. L. Wood.

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

Microcorthylus lassus, n. sp.

This species is distinguished from all preceding species in this
genus by the complete absence of an elevation or crest on declivital
interstriae 1 and by the complete absence of sutures on the male
antennal club, suture 1 and the ends of 2 occurring in the female.

Female. — Length 1.5 mm (paratypes 1.5-1.6 mm), 2.8 times as
long as wide; color dark brown.

Frons as in pumilus Wood except mandibles normal. Anten-
nal club 1.5 times as long as scape, 1.2 times as long as wide, suture
1 complete, 2 obsolete except at extreme margins.

Pronotum and elytral disc as in pumilus. Elytral declivity
essentially as in debilis Wood except interstriae 1 not elevated,
its lateral crest entirely obsolete; all punctures obsolete; surface re-
ticulate.

Male. — Similar to female except sutures on antennal club ob-
solete (one specimen with suture 1 very feebly indicated) ; anterior
margin of pronotum armed by two basally contiguous, coarse,
pointed serrations.

Type Locality. — Tapanti, Cartago, Costa Rica.

Type Material. — The female holotype, male allotype, and two
paratypes were taken at the type locality on 17-VIII-63, 1300 m,
No. 106, from a woody vine, by S. L. Wood. Three paratypes are
from Cerro Punta (labeled Volcan Chiriqui), Chiriqui, Panama, 11-
1-64, 1800 m. No. 388, tree seedling, S. L. Wood.

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

Microcorthylus concisus, n. sp.

This species is distinguished from grandiclavatus Eggers by
the smaller size, by the uniformly convex female frons, and by the
less strongly impressed elytral declivity.

Female. — Length 1.8 mm (paratypes 1.6-1.8 mm), 2.6 times as
long as wide; color dark brown.

Frons broadly convex, a slight transverse impression just above
epistoma, a short, weak, median carina at epistomal margin; central
area almost smooth, lower and lateral areas rugose-reticulate, punc-
tures fine in smooth area, slightly coarser elsewhere. Antenna about
as in M. grandiclavatus.

Pronotum and elytra as in grandiclavatus except declivity
less strongly impressed; punctures and tubercles on declivital striae
1 smaller, varying in number from zero to four.

Male. — Similar to female except frons uniformly rugose-reticu-
late; anterior margin of pronotum serrate, one or two median pairs

Dec. 1973 wood: new scolytidae 271

larger; elytra obscurely to weakly reticulate; tubercles on declivital
striae 1 larger, strial punctures almost or entirely obsolete.

Type Locality. — Volcan, Puntarenas, Costa Rica.

Type Material. — The female holotype, male allotype, and eight
para types were taken at the type locality on ll-XII-63, 1000 m. No.
304, from a tree branch, by S. L. Wood. Seven paratypes are from
San Ignacio de Acosta, San Jose, Costa Rica, 5-VII-63, 1500 m. No.
34, Croton gossypiifolius, S. L. Wood.

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

Microcorthylus vescus, n. sp.

This species is distinguished from concisus Wood by the larger
size, by the absence of a median epistomal carina, by the more
nearly strigose punctures on the frons, and by the strongly reticu-
late male elytra.

Female. — Length 2.1 mm (paratypes 2.0-2.3 mm), 2.5 times as
long as wide; color brown.

Frons as in concisus except epistomal tubercle not at all cari-
na te; punctures and impressed points slightly larger and more
nearly longitudinally strigose.

Pronotum and elytra as in concisus except punctures on de-
clivital striae 1 obsolete, accompanying tubercles minute.

Male. — Similar to female except frons more uniformly rugose-
reticulate; anterior margin of pronotum serrate; elytral disc strongly
reticulate to declivity (also in female) ; tubercles and punctures on
declivital interstriae 1 larger, but smaller than in grandiclavatus
Eggers.

Type Locality. — Volcan Zunil, Quezaltenango, Guatemala.

Type Material. — The female holotype, male allotype, and 24
paratypes were taken at the type locality on 27-V-64, 1000 m, No.
628, from a tree seedling, by S. L. Wood.

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

Microcorthylus dilutus, n. sp.

This species is distinguished from ocularis Wood by the slight-
ly larger size, by the stronger epistomal impression, with the
lower frons flattened, and by the smoother elytra with the strial
punctures in rows.

Male. — Length 1.6 mm (paralypes 1.5 mm), 2.8 times as long
as wide; color yellowish brown (callow?).

Frons rather narrow, less than 1.2 times greatest width of eye;
convex, somewhat flattened below, transversely impressed immedi-
ately above epistomal margin; surface almost smooth, with a few
small, shallow punctures; vestiture very sparse, inconspicuous. Eye
almost twice as large as in other species, very coarsely faceted. An-
tennal club large, 1.5 times as long as scape, 1.3 times as long as
wide; sutures 1 and 2 almost straight.

272 GREAT BASIN NATURALIST Vol. 33, No. 4

Pronotum 1.1 times as long as wide; essentially as in lassus
Wood except anterior margin very weakly serrate.

Elytra 1.9 times as long as wide, 1.8 times as long as pronotum;
sides almost straight and parallel on basal three-fourths, very broad-
ly rounded behind; disc almost smooth, subshining, strial punctures
rather obscurely indicated, in rows. Declivity steep, shallowly sul-
cate, contours about as in demissus Wood; sutural interstriae
weakly elevated, its lateral margin abrupt; surface reticulate.

Female. — Similar to male in all respects.

Type Locality. — Piedras Blancas, 10 km E Medellin, Antioquia,
Colombia.

Type Material. — The male holotype, female allotype, and one
paratype were taken at the type locality on 15-VII-70, 2500 m, No.
658, from a species of Guttiferae, by S. L. Wood.

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

Microcorthylus hostilis, n. sp.

This species is somewhat similar to rufotestaceus Schedl but
is distinguished by the larger size, by having the anterior margin
of the pronotum armed in both sexes, by the slightly different frons,
and by the complete absence of two large pits in declivital striae
1 in the male.

Female. — Length 2.6 mm (paratypes 2.6-2.7 mm), 2.5 times as
wide; color dark brown.

Frons convex, a weak transverse impression just above epistoma,
a low, blunt, median epistomal tubercle almost attaining epistomal
margin, tubercle larger and nearer margin than in rufotestaceus;
surface uniformly rugose-reticulate on lower two-thirds of area be-
low eyes, more nearly reticulate on sides above, central area almost
smooth and shining to vertex, punctures small, shallow, not close.
Posterior face of antennal club with several long setae not exceeding
tip of club.

Pronotum and elytra as in rufotestaceus except anterior mar-
gin of pronotum armed by at least two rather coarse, median serra-
tions; and declivital spines 1 and 2 almost obsolete, lower declivity
with several very small, irregularly placed granules. Fine vestiture
as in rufotestaceus.

Male. — Similar in all respects to female except antennal club
without long setae on posterior face.

Type Locality. — Thirty km N Merida, Merida, Venezuela.

Type Material. — The female holotype, male allotype, and 10
paratypes were taken at the type locality on 8-1-70, 2200 m, No.
226, from a broken branch, by S. L. Wood.

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

Microcorthylus diver sus, n. sp.

This species is distinguished from hostilis Wood by the slight-
ly smaller size, by the idenfinite epistomal tubercle, by the differ-

Dec. 1973 wood: new scolytidae 273

ent frons, and by the row of punctures on declivital striae 1 of the
male.

Male. — Length 2.3 mm (paratypes 2.2-2.4 mm), 2.5 times as
long as wide; color dark brown.

Frons as in hostilis except epistomal tubercle essentially ab-
sent; rugose-reticulate area extending above eyes.

Pronotum and elytra as in hostilis except striae 1 on upper
half of declivity with a series of seven (8 to 11 in paratypes) rather
small, deep punctures; lower area of declivity more nearly reticu-
late and with fewer granules.

Female. — Similar to male except transverse impression above
epistoma more distinct; frons from just below upper level of eyes
to vertex smooth, brightly shining, with moderately abundant fine
punctures and impressed points, antennal club with a few long hairs
on posterior face; punctures on declivital striae 1 minute to obsolete.

Type Locality. — La Carbonera Experimental Forest, 50 km (air-
line) NW Merida, Merida, Venezuela.

Type Material. — The male holotype, female allotype, and 27
paratypes were taken at the type locality on 10-XL69, 2500 m, Nos.
124, 126, 127 (type), from cut Nectantra seedlings, b}'^ S. L. Wood;
28 paratypes bear similar data except 27-X-69, No. 91 A; 18 para-
types are from Colonia Tovar, Aragua, Venezuela, 4-V-70 1700 m,
No. 485, Nectandra, S. L. Wood.

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

Microcoorthylus umbratus, n. sp.

This species is distinguished from diversus Wood by the smaller
size, by the different frons in both sexes, by the much less strong-
ly impressed elytral declivity, and by the much smaller punc-
tures on the male striae 1 .

Male. — Length 1.9 mm (paratypes 1.9-2.2 mm), 2.5 times as
long as wide; color brown to very dark brown.

Frons as in diversus except strongly rugose-reticulate from
epistoma to vertex; rather dull.

Pronotum and elytral disc as in diversus except anterior mar-
gin of pronotum with up to six serrations; elytral declivity much
less strongly impressed, lateral convexities rather weakly elevated,
dull, finely reticulate, striae 1 with about eight (7-11 in paratypes)
minute punctures on middle half of declivity.

Female. — Similar to male except lower third of frons abruptly,
rather shallowly impressed (stronger than in female diversus),
upper area on median half smooth, brightly shining; posterior face
of antennal club with several long hairs; anterior margin of prono-
tum feebly serrate; punctures on declivital striae 1 obsolete.

Type Locality. — La Carbonera Experimental Forest, 50 km
(airline) NW Merida, Merida, Venezuela.

Type Material. — The male holotype, female allotype, and nine
paratypes were taken at the type locality on 10-XL69, 2500 m, Nos.

274 GREAT BASIN NATURALIST Vol 33, No. 4

125, 126 (type), 127, from cut Nectandra seedlings, by S. L. Wood;
14 paratypes bear similar data except 27-X-69, 91A; 3 paratypes
are from the Merida Teleferico, Merida, Venezuela, 27-11-70, 2500
m. No. 332, Nectandra, S. L. Wood.

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

Microcorthylus curt us, n. sp.

This species differs from all of the preceding species in having
the lateral margin of the declivity continued at the base to the su-
ture, and in having the face of the declivity finely punctured, not
reticulate.

Male. — Length 1.6 mm (paratypes 1.5-1.7 mm), 2.5 times as
long as wide; color dark brown.

Frons evenly convex from epistoma to vertex; surface rugose-
reticulate from epistoma to upper level of eyes, upper areas almost
smooth, with numerous impressed points and a few rather fine, shal-
low punctures; vestiture inconspicuous. Antennal club 1.5 times as
long as scape, 1.15 times as long as wide, subcircular; sutures 1
and 2 clearly indicated.

Pronotum and elytral disc essentially as in diversus Wood,
except elytral disc on posterior third weakly reticulate with some
small, smooth, shining areas. Elytral declivity subvertical, broadly
impressed, not sulcate at base; lateral margins moderately, rather
abruptly elevated on upper two-thirds, continued to suture, armed
in usual position by minute spines 1 and 2; declivital face minutely,
densely punctured. Vestiture confined to declivital face, of very
short, fine, abundant hair.

Female. — Similar to male except anterior margin of pronotum
unarmed.

Type Locality. — Twenty km SW El Vigia, Merida, Venezuela.

Type Material. — The male holotype, female allotype, and six
paratypes were taken at the type locality on lO-XII-69, 50 m, No.
190, from a cut tree seedling, by S. L. Wood.

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

Microcorthylus contractus, n. sp.

This species is distinguished from curtus Wood by the more
extensively reticulate, subaciculate frons, by the more numerous
serrations on the anterior margin of the pronotum, and by the less
extensive, more shallowly impressed elytral declivity.

Male. — Length 1.4 mm (paratypes 1.4-1.6 mm), 2.5 times as
long as wide; color brown.

Frons as in curtus except abundant impressed points longi-
tudinally strigose.

Pronotum and elytra as in curtus except anterior margin of
pronotum armed by about 14 serrations of about equal size; elytral
declivity strongly reticulate to declivity. Elytral declivity trangular-

Dec. 1973 wood: new scolytidae 275

ly impressed on a limited area, lateral margins not strongly elevated,
their upper limits attaining area of striae 1; striae 1 with a row
of very small, obscure punctures on lower two-thirds; surface re-
ticulate. Subglabrous.

Type Locality. — Seven km NW Socopo, Barinas, Venezuela.

Type Material.- — The male holotype and four male paratypes
were taken at the type locality on 13-11-70, 200 m, No. 332, from
a cut Nectandra seedling, by S. L. Wood.

The holotype and paratypes are in my collection.

THE NOMINAL SNAKE GENERA MASTIGODRYAS
AM ARAL, 1934, AND DRYADOPHIS STUART, 1939

Hobart M. Smiths and Kenneth R. Larsen^

Abstr-'\ct. — Some recent workers who have combined the genera Mastigo-
dryas Amaral, 1834, and Dryadophis Stuart, 1939, under the former name, have
done so prematurely in view of characters by Amaral that differentiate
them. Either the genera should be maintained distinct, or the younger, but
widely-known name, Dryadophis, should be retained through exercise of the
plenary powers of the ICZN, already requested.

Prompted by Romer's (1956: 577) indication of the synonyniy of
Mastigodryas Amaral, 1934, and Dryadophis Stuart, 1939, a propo-
sal was made (Smith, 1963) that Mastigodryas be suppressed under
the plenary powers of the International Commission on Zoological
Nomenclature in order to preserve the more familiar Dryadophis.
The ICZN never acted upon the proposal, however, because Amaral
(1964) pointed out that the monotypic Mastigodryas should not be
regarded as a synonym of Dryadophis, since it has no apical scale
pits and 70 subcaudals, whereas in Dryadophis the subcaudals num-
ber 79 or more, and paired apical scale pits are present in all species.

Nevertheless Peters and Orejas-Miranda (1970:190) lumped the
two genera, citing all valid Central and South American species-
group taxa of Dryadophis under the generic name Mastigodryas.
No mention was made of the comments by Smith and Amaral that
appeared in the Bulletin of Zoological Nomenclature; but Romer's
observation was noted and the comment added that Peters had seen
the type of M. danieli, that he concluded it is congeneric with Drya-
dophis, and that priority should be obrerved.

We have not seen any specimens of M. danieli, but on the basis
of Amaral's (1964) comments maintain that to regard it as congener-
ic with Dryadophis is premature. Dryadophis merits peri)etviation
until more conclusive evidence of synonymy with Mastigodryas is
available. If such a conclusion is confirmed, reconsideration should
be given to suppression of the nominal geiuis Mastigodryas in order
to {)reserve the widely known name Dryadophis. resubmitting for
ICZN action the proposal first presented in 1963.

LlTr,R..\TURE CiTF.D

Amaral, A. do. 1964. Comment on the proposal to substitute the generic name
Dryadophis Stuart, 1939, for Mastigodryas Amaral, 1934. Bull. Zoo).
Nomencl. 21(1): 13.

Peters, J. A., and B. Ore.jas-Miranda. 1970. Catalogue of the neotropical
Squamata: Part I. Snakes. Bull. U.S. Nat. Mus. 297:1-346, ill.

RoMER, A. S. 1956. Osteology of the reptiles. University of Chicago Press.

Smith, H. M. 1963. Dryadophis Stuart. 1939 (Reptilia, Serpentes): Pro-
posed validation under the plenary powers. Bull. Zool. Nomencl. 20(3):230.

'Department of Environmental, Population, and Organismic Biology, University of Colorado,
Boulder 80302.

276

INDEX TO VOLUME 33

The genera and species described as new to science in this volume
appear in bold type in this index.

A new subfruticose Eriogonum
(Polygonaceae) from western
Colorado, p. 120.

A taxonomic revision of Physaria
(Cruciferae) in Utah, p. 31.

Additional records of mutillid wasps
from the Nevada Test Site, p. 156.

Alexander, Charles P., article by,
p. 189.

Allred, Dorald M., articles by, p. 51,
123, 156, 246, 251.

An unusual population of spiders in
Utah, p. 51.

Artemesia arbuscula, A. longiloba,
and A. nova habitat types in
northern Nevada, p. 225.

Arvizo, Edward R., and Richard D.
Worthington, article by, p. 124.

Barber, Delbert L., and Lawrence L.
Lockard, article by, p. 53.

Baumann, Richard W., article by,
p. 91.

Behle, William H., article by, p. 243.

Blackburn, Del F., article by, p. 203.

Carphotoreus, p. 171.

Clark, Tim W., article by, p. 205.

Courtship behavior among white-
tailed and black-tailed jackrabbits,
p. 203.

Density changes and habitat affini-
ties of rodents of shadscale and
sagebrush associations, p. 255.

Density, growth, and home range of
the lizard Uta stansburinana stej-
negeri in southern Dona Ana Coun-
ty, New Mexico, p. 124.

Ecology of Sceloporus magister at
the Nevada Test Site, Nye County,
Nevada, p. 133.

Elder, John A., and Arden R. Gau-
fin, article by, p. 218.

Evans, Howard E., articles by p. 29,
147.

Further studies on the wasps of
Jackson Hole, Wyoming (Hymen-
optera, Aculeata), p. 147.

Galton, Peter M., and James A.
Jensen, article by, p. 129.

Gaufin, Arden R., and John A. El-
der, article by, p. 218.
Grogan, William L., and Lloyd C.

Pack, Jr., article by, p. 202.
Gymnodamaeus leurolomasus, p. 40.
Gymnodamaeiis plokosus, p. 37.
Helminths of Sceloporus lizards in
the Great Basin and upper Colo-
rado plateau of Utah, p. 1.
Herrin, C. Selby, article by, p. 26.
Higgins, Harold G., and Tyler A.

Woolley, article by, p. 37.
Incidence of spotted fever in wood
ticks of Utah recreational sites,
p. 26.
Jensen, James A., and Peter M.

Galton, article by. p. 129.
Johnson, Donald R., and Earl J.

Larrison, article by, p. 255.
Krogh, John E., and Wilmer W. Tan-
ner, article by, p. 133.
Larrison, Earl J., and Donald R.

Johnson, article by, p. 255.
Larsen, Kenneth R., and Hobart M.

Smith, article by, p. 276.
Limnophila byersi, p. 195.
Local distribution and interspecies
interactions in microtines, Grand
Teton National Park, Wyoming, p.
205.
Lockard, Lawrence L., and Delbert

L. Barber, article by, p. 53.
Medlyn, David A., Gregory F.
Thayn, and William D. Tidwell,
article by, p. 61.
Microcorthylus concisus, p. 270.
Microcorthylus contractus, p. 274.
Microcorthylus curtus, p. 274.
Microcorthylus debilis, p. 265.
Microcorthylus demissus, p. 266.
Microcorthylus dilutus, p. 271.
Microcorthylus diversus, p. 272.
Microcorthylus hostilis, p. 272.
Microcorthylus inermis, p. 267.
Microcorthylus invalidus, p. 268.
Microcorthylus lassus, p. 270.
Microcorthylus ocularis, p. 266.
Microcorthylus pumilus, p. 268.
Microcorthylus pusillus, p. 269.

277

278

GREAT BASIN NATURALIST

Vol 33, No. 4

Microcorthylus umbratus, p. 273.

Microcorthylus vescus, p. 271.

Miscellaneous chromosome counts of
western American plants— II, p.
19.

Nearctic desert Decticidate (Orthop-
tera). Part II. A new genus and
species from Arizona, p. 43.

Nearctic desert Decticidae (Orthop-
tera). Part III, the true tym-
panum in certain genera with key,
p. 197.

New species of American Microcor-
thylus (Coleoptera: Scolytidae), p.
265.

New synonymy in American bark
beetles (Scolytidae: Coleoptera).
Part III, p. 169.

Notes on aquatic and semiaquatic
Hemiptera from the southwestern
United States (Insecta: Hemip-
tera), p. 113.

Notes on reproduction in Lampropel-
tis triangulum and Coluber con-
strictor in Utah, p. 202.

Notes on the nesting behavior of
Steniolia elegans (Hymenoptera:
Sphecidae), p. 29.

Notes on the occurrence and distri-
bution of Pteronarcys californica
Newport (Plecoptera) within
streams, p. 218.

On the taxonomic status of Platy-
podidae and Scolytidae (Coleop-
tera), p. 77.

Pack, Lloyd C, Jr., and WiUiam L.
Grogan, article by, p. 202.

Palmoxylon colei, p. 64.

Palmoxylon contortum, p. 67.

Palmoxylon edenense, p. 61.

Pearce, Richard C, and Wilmer W.
Tanner, article by, p. 1.

Platyoplus, p. 43.

Platyoplus gilaensis, p. 44.

Polhemus, John T., article by, p.
113.

Prehistoric bighorn sheep in the
northern Sierra Nevada, California,
p. 221.

Protohylastes, p. 83.

Protohylastes annosus, p. 84.
Protoplatypus, p. 81.
Protoplatypus vetulus, p. 82.
Records of Coreidae (Hemiptera)
from the Nevada Test Site, p. 123.

Reveal, James L., article by, p. 120.

Reveal, James L., and Eloise L.
Styer, article by, p. 19.

Schulz, Peter D., and Dwight D.
Simons, article by, p. 221.

Scorpions of the National Reactor
Testing Station, Idaho, p. 251.

Significant bird records from Utah,
p. 243.

Simons, Dwight D., and Peter D.
Schulz, article by, p. 221.

Small bones of the hypsilophodontid
dinosaur Dryosaurus altus from
the upper Jurassic of Colorado, p.
129.

Small mammals of the National Re-
actor Testing Station, Idaho, p.
246.

Smith, Hobart M., and Kenneth R.
Larsen, article by, p. 276.

Some helminths from mink in south-
western Montana, with a checklist
of their internal parasites, p. 53.

Studies on Utah stoneflies (Plecop-
tera), p. 91.

Styer, Eloise L., and James L. Re-
veal, article by, p. 19.

Sweltsa gaufini, p. 102.

Tanner, Wilmer W., and John E.

Krogh, article by, p. 133.

Tanner, Wilmer W., and Richard C.
Pearce, article by, p. 1.

Thayn, Gregory F., William D. Tid-
well, and David A. Medlyn, article
by, p. 61.

The effects of soil texture on species

diversity in an arid grassland of
the eastern Great Basin, p. 163.

The male brachycistidine wasps of
the Nevada Test Site (Hymenop-
tera: Tiphiidae), p. 109.

Three new species of Palmoxylon
from the Eocene Green River for-
mation, Wyoming, p. 61.

Tidwell, William D., David A. Med-
lyn, and Gregory F. Thayn, article
by, p. 61.

Tinkham, Ernest R., articles by, p.
43, 197.

Tipula (Trichotipula) frommeri, p.
p. 189.

Tipula (Lunatipula) hastingsae di-
perona, p. 193.

Tipula (Pterelachisus) macleani, p.
193.

Dec. 1973 index 279

Tipula (Trichotipula) sanctaecruzae, Wasbauer, Marius S., article by, p.

p. 192. 109.

Tueller, P. T., and B. Zamora, arti- Wood, Stephen L., articles by, p. 77,

cle by, p. 225. 169, 265.

Two new species of Gymondamaeus Woolley, Tyler A., and Harold G.

from Colorado (Acarina: Crypto- Higgins, article by, p. 37.

stigmata, Gymnodamaeidae), p. Worthington, Richard D., and Ed-

37. ward R. Arvizo, article by, p. 124.

Undescribed species of nearctic Wyckoff, John W., article by, p. 163.

Tipulidae (Diptera), XI, p. 189. Zamora, B., and P. T. Tueller, arti-

Waite, Sheldon B., article by, p. 31. cle by, p. 225.

M :-j\^-.>H','..:''.vf-i.-t\:.-. ^ '';:'.v' . ■- •;■

GREAT BASIN NATUR
Notice to Contributors

Original manuscripts in English pertaininj^ biolospcal

t latural history of western North America ai : i •

tion in the Great Basin Naturalist should 1' <. "^

Young University, Stephen L. Wood, Editor, Great Basin Naturalist,
Provo, Utah 84602.

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TABLE OF CONTENTS

I^ocal distribution and interspecies interactions in microtines.

Grand Teton National Park, Wyoming. Tim W. Clark...! 205
Notes on the occurrence and distribution of Pteronarcys

californica Newport (Plecoptera) within streams. John

A. Elder and Arden R. Gaufin 218

Prehistoric bighorn sheep in the northern Sierra Nevada,

California. Peter D. Schulz and Dwight D. Simons 221

Artemesia arbuscula, A. longiloba, and A. nova habitat types

in northern Nevada. B. Zamora «ind P. T. Tueller 225

Significant bird records from Utah. William H. Behle. 243

Small mammals of the National Reactor Testing Station,

Idaho. Dorald M. Allred. 246

Scorpions of the National Reactor Testing Staticm, Idaho.

Dorald M. Allred ....^ 261

Density changes and habitat affinities of rodents of shadscale

and sagebrush associations. Elarl J. Larrison and Donald

R. Johnson 255

New species of American Microcorthylus (Coleoptera:

Scolytidae). Stephen L. Wood. 265

The nominal snake genera Mastigodryas Amaral, 1834, and

and Dryadophis Stuart, 1839. Hobart M. '=^""»i- -nd

Kemieth R. Larsen. , .. 276

3 2044 072 231 079

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