HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

The Great Basin Naturalist

VOLUME XXV, 1965

Editor: Vasco M. Tanner

Assistant Editor: Stephen L. Wood

Assistant Editor: Wilmer W. Tanner

Published at Provo, Utah by
Brigham Young University

VOLUME XXV NOS. 1-2

TABLE OF CONIENIS

A New Net- winged Midge from Idaho (Blepharoceridae.

Diptera). Charles P. Alexander 1

A List of Some Beeflies of the Nevada Test Site. Dorald M.
Allred, D. Elmer Johnson, and D Elden Beck.
Illustrated 5

Tularemia and Deer Flies in the Environs of Utah Lake.

Utah. Kaye B. Cox. Illustrated 13

The Genus Eupagiocerus Blandford (Scolytidae, Cole-

optera). Stephen L. Wood 31

Note of Phalangids at the Nevada Test Site 37

A New Genus and Special in the Chilopod Family Tam-

piyidae. Ralph V. Chamberlin. Illustrated 39

Two Isopods of the Nevada Test Site. Dorald M. Allred

and Stanley Mulaik 43

Note: Prairie Falcon Imitates Flight Pattern of the Logger-
head Shrike. Gerald L. Richards 48

Numbers 3 4 — December 31, 1965

Undescribed Species of Nearctic Tipulidae (Diptera) VI.

Charles P. Alexander 49

Two New Mites from the United States (Acari: Oribatei,
Microzetidae, and Oribatellidae). Harold G. Higgins.
Illustrated 55

A New Genus and Species of Oribatid Mite from Colorado
(Acari: Orbatei, Ceratoppiidae). Tyler A. Woolley
and Harold G. Higgins. Illustrated 59

Studies in Nearctic Desert Sand Dune Orthoptera, A New
Genus and Species of Stenopelmatince Crickets from
the Kelso Dunes with Notes on its multi-annual life
history and key. Part X. Ernest R. Tinkham. Illus-
trated 63

Records of Atherinid Fishes at Inland Localities in Texas

and Northern Mexico. W. L. Minckley. Illustrated 73

A List of Scarabaeidae Beetles of the Nevada Test Site.

Dorald M. Allred and D Elden Beck 77

Angus Munn Woodbury, 1886-1964. Vasco M. Tanner.

Illustrated 81

Index 89

II

INDEX TO VOLUME XXV

The new genera and species described in this volume appear in bold
face type in this index.

A List of Scarabaeidae Beetles of the

Nevada Test Site, 77.
A List of Some Beeflies of the Ne-
vada Test Site, 5.
A New Genus and Species of Oribatid

Mite from Colorado (Acari: Ori-

batei, Ceratoppiidae), 59.
A New Genus and Species in the

Chilopod Family Tampiyidae, 39.
A New Net-winged Midge from Idaho

(Blepharoceridae, Diptera), L
Abatorus, New genus, 40.
Abatorus allredi. New species, 40.
Alexander, Charles P., Article by,

1, 49.
Allozetes, 55.

Allozetes harpezus n. gen., n. sp., 55.
Allred, Dorald M., Articles by, 5, 37,

43, 77.
Ammopelmatus kelsoensis, n. sp., 67.
Angus Munn Woodbury, 1886-1964, 81.
Beck, D Elden, see Allred, 5, 77.
Bibiocephala nigripes sp. n., 2.
Chamberlin, Ralph V., Article by, 39.
Cox, Kaye B., Article by, 13.
Eupagiocerus clarus, n. sp., 33.
Eupagioceurs vastus, n. sp., 34.
Higgins, Harold G., Article by, 55.
Higgins, Harold G., see Woolley, 59.
Index, 89.

Johnson, D. Elmer, see Allred, 5.
Limonia (Dicranomyia) dreisbachi,

53.
Minckley, W. L., Article by, 73.
Mulaik, Stanley, see Allred, 43.
Note of Phalangids at the Nevada

Test Site, 37.

Ophidiotrichus exastus, 57.

Paenoppia n. gen., 59.

Paenoppia forficuia, n. sp., 59.

Prairie Falcon Imitates Flight Pat-
tern of the Loggerhead Shrike, 48.

Records of Atherinid Fishes at In-
land Localities in Texas and North-
ern Mexico, 73.

Richards, Gerald L., Note by, 48.

Studies in Nearctic Desert Sand Dune
Orthoptera, A new Genus and Spe-
cies of Stenopelmatine Crickets
from the Kelso Dunes with notes
on its multi-annual life history and
key. Part X., 63.

Tanner, Vasco M., Article by, 81.

Tipula Mercedensis, n. sp.. 50.

The Genus Eupagiocerus Blandford
(Scolytidae: Coleoptera), 31.

Tinkham, Ernest R., Article by, 63.

Tipula (Nippotipula) metacomet n.
sp., 49.

Tipula (?Pterelachisus) simondsi, n.
sp., 51.

Tularemia and Deer Flies in the En-
virons of Utah Lake, Utah, 13.

Two New Mites from the United
States (Acari: Oribatei, Microzeti-
dae and Oribatellidae), 55.

Two Isopods of the Nevada Test Site,
43.

Undescribed Species of Nearctic Tip-
ulidae (Diptera) VI., 49.

Wood. Stephen L., Article by, 31.

Woolley, Tyler A., Article by, 59.

The

f V iU

Great Basin

Volume XXV July 8, 1965 Nos. 1-2

TABLE OF CONTENTS

A New Net-winged Midge from Idaho (Blepharoceridae,

Diptera). Charles P. Alexander 1

A List of Some Beeflies of the Nevada Test Site. Dorald M.
Allred, D. Elmer Johnson, and D Elden Beck. Illus-
erated 5

Tularemia and Deer Flies in the Environs of Utah Lake,

Utah. Kaye B. Cox. Illustrated 13

The Genus Eupagiocerus Blandford (Scolytidae, Coleoptera).

Stephen L. Wood 31

Note of Phalangids at the Nevada Test Site. Dorald M.

Allred 37

A New Genus and Species in the Chilopod Family Tampiyidae.

Ralph V. Chamberlin. Illustrated 39

Two Isopods of the Nevada Test Site. Dorald M. Allred and

Stanley Mulaik 43

Note: Prairie Falcon Imitates Flight Pattern of the Logger-
head Shrike. Gerald L. Richards 48

4

X939

The Great Basin Naturalist

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

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

Illustrations: All illustrations should be made vsdth a view to
having then appear v/ithin the limits of the printed page. The ill-
ustrations that form a part of an article should accompany the
manuscript. All half-tones or zinc etchings to appear in this jour-
nal are to be made under the supervision of the editor, and the cost
of the cuts is to be borne by the contributor.

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

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

All con-espondence dealing with manuscripts, subscriptions,
reprints and other business matters should be addressed to the Editor,
Vasco M. Tanner, Great Basin Naturalist, Brigham Young Univer-
sity, Provo, Utah.

Reprints Schedule of

' The Great Basin Naturalist

50

too

copies
copies

2 pp. 4 pp.

$6.00 $7.00

7.00 8.00

Each
Additional

6 pp. 8 pp. 10 pp. 12 pp. 2 pp.
$8.00 $9.00 $10.00 $11.00 $2.00
9.00 10.00 11.00 12.00

200

copies

8.00 9.00

10.00 11.00 12.00 13.00

300

copies

9.00 10.00

11.00 12.00 13.00 14.00

Covi'Rs: $10.00 for first 100 copies, $4.00 for additional 100 copies.

The Great Basin Naturalist ^""^^^

UNIVERSITY.

Published at Provo, Utah by

Brigham Young University
Volume XXV July 8, 1965 Nos. 1-2

A NEW NET-WINGED MIDGE FROM IDAHO
(BLEPHAROCERIDAE, DIPTERA)

Charles P. Alexander*

An interesting undescribed species of net-winged midge belonging
to the genus Bibiocephala Osten Sacken was taken in Idaho by my
friend Mr. James Baker, of Baker, Oregon. I am very deeply indebted
to the collector for many new and rare crane-flies taken during the
past twenty-five years in several of the western states and provinces.
The unique type is preserved in my personal collection.

General Account of Bibiocephala

The genus Bibiocephala Osten Sacken (see Bibliography) now
includes five nominal species, one occurring in Japan, the remaining
four names pertaining to western Nearctic species. In attempting
to clear the identities and synonymy of the Ajnerican species I am
reviewing briefly the series of circumstances under which the vari-
ous names were proposed, the papers cited being included in the
appended bibliography. Attention may be directed to two recent
papers by the writer (1958, 1963) which include more detailed
references to the family.

Osten Sacken (1874:564-566, figure, venation) proposed the
genus Bibiocephala, based on the new species, grandis, taken at high
altitudes in the mountains of Colorado, 8,000 to 10,000 feet, August
1873, represented by a single male specimen taken by Lieutenant
W. L. Carpenter. Garrett (1922:91) described Bibiocephala kelloggi
from a single female specimen taken in the city of Cranbrook, British
Columbia, July 13, 1921, b}^ Cecil B. D. Garrett. In defining the
species Garrett recorded it as being a male but actually this type is
a female. The specimen is in my personal collection having been
acquired by purchase from Garrett. It still is uncertain whether this
fly is distinct from grandis. Curran (1923) described Bibiocephala
grisea, as type of a supposed new genus Bibionus, based on a single
male taken at Nordegg, Alberta, June 26, 1921, by James Mc Dun-
nough. This evidently is identical with kelloggi which is the prior
name.

1. Amherst, Massachusetts.

The Great Basin Naturalist
2 CHARLES P. ALEXANDER Vol. XXV, Nos. 1-2

In 1890 von Roder proposed the genus Agathon, for A. elegantula
von Roder, of Nevada, based on the short vein /?., of the wings and
the glabrous thoracic pleura. Kellogg (1903:192-195), in describing
two new species from California with shortened vein /?3, referred
these species to Bibiocephala as B. comstocki and B. doanei, not rec-
ognizing the distinctness of the two genera Agathon and Bibiocephala.
This misinterpretation of the generic limits of Bibiocephala led to the
further errors by Garrett and Curran, as mentioned. Walley (1927)
recognized both genera and separated the various species correctly
except for referring Agathon comstocki (Kellogg) to Bibiocephala.

I'he Japanese species was originally described by Matsumura
(1916) as a species of Liponeura, under the name Liponeura injus-
cata. Kitakami (1950) again followed Kellogg's interpretation and
considered that the Japanese species required a new generic name
Amika (the Japanese name for these insects), thus creating a second
synonym in Bibiocephala.

The presently known members of the genus are as follows:

Bibiocephala grandis Osten Sacken (1874); cf-

Bibiocephala kelloggi Garrett (1922); $ (status in question).

synonym: Bibiocephala grisea (Curran, 1923); cT.
Bibiocephala nigripes sp.n.; d" •
Bibiocephala injuscata (Matsumura, 1916); d"-

Generic synonymy:
Bibiocephala Osten Sacken (1874).
Bibionus Curran (1923).
Amika Kitakami (1950).

Bibiocephala nigripes sp.n.

Male. — Length, about 11 mm.; wing 9x4 mm.; antenna, about
1.2 mm.

Head very large, especially the eyes which are broadly contigu-
ous above; eyes with reduced lower section only about one-third to
one-fourth the upper division. Antennae 14-segmented, short, black
throughout; scape short, pruinose, pedicel much longer, dilated at
apex; proximal two flagellar segments united, nearly equal in length
to the succeeding three combined; eighth and succeeding segments
transverse, broader than long, the penultimate about one-half broad-
er than its length, terminal segment short-oval. Head gray.

Mesonotal praescutum light gray with two brown stripes, their
anterior ends much widened; posterior sclerites and pleura generally
gray. Halteres with stem light yellow, knob dark brown. Legs witn
coxae infuscated; trochanters yellowed, darkened apically beneath;
femora yellowed on about the basal two-thirds, the tips and remain-
ing segments intensely black; tibial spur formula 1 - 2 - 2. Wings
very broad, especially on proximal portion, the anal region pro-
duced backward; wings rather strongly infuscated, the whitened
secondary wing folding more conspicuous than in grandis but nar-
rower and less evident than in infuscaia.

i> r^i - vy ii-<nji:-u iviLuvjii, r nijivi lUAttKJ

Basal abdominal tergites broadly light gray on sides, narrower on
posterior borders, the midregion more blackened, the subterminal
segments more uniformly darkened; hypopygium yellowish brown.

HoLOTYPE, cT, near Featherville, Elmore County, Idaho, in Saw-
tooth State Forest, 4,900 feet, July 13, 1964 (James Baker).

Bibiocephala nigripes is most readily distinguished from the pre-
viously described American species by the intensely blackened tibiae
and tarsi which have suggested the specific name.

Bibliography

Alexander, C. P. 1958. Geographical distribution of the net-winged midges

(Blepharoceridae, Diptera). Proc. Tenth Internat. Congr. Entom., 1 (1956)

813-828, 3 maps, 23 figs.
. 1963. Blepharoceridae and Deuterophlebiidae, in Guide to the insects

of Connecticut, Part VI, fasc. 8, 39-83, pis. 8-15, with 36 figs. (Bull. 93,

State Geol. & Nat. Hist. Sui-vey of Connecticut).
CuRRAN, C. H. 1923. Studies in Canadian Diptera II. The genera of the family

Blepharoceridae. Canad. Ent., 55:266-269 {Bibionus; Bibionus griseus) .
Garrett, C. B. D. 1922. Two new Blepharoceridae (Diptera). Insec. Inscit.

Menst., 10:89-91 {Bibiocephala kelloggi).
Kellogg, V. L. 1903. The net-winged midges (Blepharoceridae) of North

America. Proc. California Acad. Sci., (3) Zool. 3; 187-221, 5 pis., 1 fig.
KiTAKAMi, SiRO. 1950. The revision of the Blepharoceridae of Japan and ad-
jacent territories. Jour. Kumamoto Women's Univ., 2:15-80, 5 pis., 3 figs.

with subfigs.) {Amika).
Matsumura, S. 1916. Thousand Insects of Japan. Additamenta 2:185-474, pis.

16-25 (reference pp. 443-444, pi. 24, fig. 7 {Bibiocephala infuscata, as

Liponeura) .
OsTEN Sacken, C. R. 1874. Report on the Diptera collected by Lieut. W. L.

Carpenter in Colorado during the summer of 1873. In Hayden's U. S. Geol.

& Geogr. survey of Colorado for 1873, pp. 561-566, 1 fig. (venation). {Bibio-
cephala; Bibiocephala grandis).
Roder, V. von. 1890. Zwei neue nordamerikanische Dipteren. Wien. Entomol.

Zeitg, 9:230-232 {Agathon).
Walley, G. S. 1927. Review of the Canadian species of the dipterous family

Blephariceridae. Canad. Ent., 59:112-116, 8 figs. (Key to genera and species).

A LIST OF SOME BEEFLIES OF THE
NEVADA TEST SITE'

Dorald M. Allred', D. Elmer Johnson', and D Elden Beck"

During ecological investigations at the Nevada Test Site (refer to
Brigham Young University Sci. Bull., II (2): 1-52, 1963), several
thousand beef lies were collected between March, 1961, and August,
1962. Specimens were taken by members of our field staff at the
test site, and to a large extent by D. Elmer Johnson, who also identi-
fied the flies.

This reports 2,573 identifications representing 111 species of 24
genera. In addition several undescribed species were taken but are
not listed here.

The species, numbers of individuals collected, months of occur-
rence, and ecological distribution are shown in Table I. The validity
of some identifications made on the basis of descriptions and keys in
the literature is open to question, and these names are followed by
a question mark in the table.

Species which were taken in the most abundant numbers at the
test site are Lordotus albidus, L. nigriventris, and Poecilauthrax
apache. Those most widely distributed ecologically are Paracosmus
morrisoni, Poecitanthrax apache, and Villa aenea. The greatest
numbers of species and individuals were found in the Mixed and
Larrea-Franseria communities (Fig. 1). Seasonally, the greatest

I00_,

LA-FR GR-LY CO AT -KO P I - J U CS MIXED

Fig. 1. Relative numbers of species of beeflies found in each of seven plant
communities.

1. AECBYU Report No. COO-1 355-7. Field work cdinpleted under Atomic Energy Ccminnssion
Contract No. AT(11-1 )78().

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

3. Ecology and Epizoology Research, University of Utah, Dugway, Utah.

ALLRED, JOHNSON, BECK

The Great Basin Naturalist

Vol. XXV, Nos. 1-2

numbers of species and individuals occurred in May, June, April,
and September, respectively (Fig. 2).

Taxonomic Note

Material in the genus Lordotus collected for this study, plus that
collected in other strategic localities, clarifies the relationships be-
tween some of the taxa in the genus and necessitates changes in the
nomenclature of several of them.

Lordotus luteolus Hall, new combination
L. pulchrissimus luteolus Hall

Collection of a copulating pair of this species at Walker Pass,
Kern County, California, on September 12, 1961, by D. E. Johnson
60_

5 0-

4 0_

3 0_

2 0.

I 0_

MAR

APR

MAY

JUN

JUL

AUG

SEP

OCT

Fig. 2. Relative numbers of species of beeflies found during each of eight
months of the year.

linked the two sexes of this species and made possible the certain
separation of this species and Lordotus pulchrissimus Williston. In
L. pulchrissimus the hypopleura are hairy while in L. luteolus they
are bare. This holds true for both sexes.

Lordotus rnelanosus Johnson and Johnson, new combination
L. miscellus rnelanosus Johnson and Johnson

Examination of much material in this species and L. miscellus
Coquillett has failed to reveal any intergrades between the two.

Lordotus nigriventris Johnson and Johnson, new combination
L. sororculus nigriventris Johnson and Johnson

As in the species above, examination of many specimens of this
species and L. sororculus Williston and finding the two occurring at
the same place, without intergrades being evident, are indicative
that the two are best considered as distinct species. Lordotus sororculus
is the more southerly and westerly in distribution, the two coming
together in southern Nevada.

Lordotus striatus Painter, new combination
L. gibbus striatus Painter

This species and L. gibbus Loew have been found associated to-
gether in a number of places. They differ from each other as much
as any of the related species in the group. Therefore, these are con-
sidered as distinct species.

Table 1. Occurrence of beeflies in plant communities
at the Nevada Test Site

(* = Occurrence; P = Predominance; Month in boldface = Period of
greatest abundance.)

Plant Community or Locality'

Species

u

>,

o

x)

h

J

W

3

•-5

X

ti

u

o

■*->

W

hJ

O

U

<

Oh

u

^

Arvastoechus hessei Hall

41-Sept., Oct.
A. melanohalteralis Tucker

2-Sept.
Anthrax albofasciatus Macquart

11-April, May
A. limatulus Say

3-June, Sept.
A. nidicola Cole ?

18-April, May
A. oedipus Fabricius

23-April through October
A. seriepunctatus (Osten Sacken)

9-May, June, July

The Great Basin Naturalist
8 ALLRED, JOHNSON, BECK Vol. XXV, NoS. 1-2

Aphoebantus ubnormis Coquillett * * *

11-Aug., Sept., Oct.
A. altercinctus Melander ? *

37-June
A. arenicola Melander ♦ *

3-May
A. argentifrons Cole *

6-Aug.
A. borealis Cole ? *

2-May
A. brevistylus Coquillett ? ♦

1-July
A. desertus Coquillett ♦ * * P

28-April, May
A. eremicola Melander P *

17-April, May
A. fmnosus Coquillett P * * *

26-April
A. interruptus Coquillett P * * *

15-April, May, June
A. marcidus Coquillett * * p

15-March, May, June
A. marginatus Cole ? *

5- July
A. marmon Melander * P * P

41-March, Aug., Sept., Oct.
A. mus Osten Sacken * P

7-April, May, July
A. pavidus Coquillett *

8-Aug.
A. parkeri Melander P *

8-July, Aug., Oct.
A. peodes Osten Sacken * * p

51-Mar., April, May
A. scalaris Melander P P *

36-May, June, July
A. scriptus Coquillett *

1-May
A. tardus Coquillett * P

14-May, June
A. Hmberlakei Melander *

1-July
A. transitus Coquillett P * * *

20-April, May
A. Ursula Melander p * ♦

15-April, May
A. varius Coquillett ? * P

9-June
A. vasatus Melander ? *

1-May
A. vittatus Coquillett * P

18-May, June, Aug.
A. vulpecula Coquillett P *

25-May, June
Astrophanes adonis Osten Sacken *

1-May
Bombylius lancifer Osten Sacken *

47-May, June
Conophorus fenestratus

(Osten Sacken) * P *

38-AprU, May, July

July 8, 1965 beeflies of Nevada

Desmatoneura argentifrons
Williston *

8-Aug.
Dipalta serpentina Osten Sacken

7-Sept.
Empiidideicus humeralis Melander * *

45-March, May
Epacmus connectens Melander *

1-May
E. labiosus Melander *

11-July, Aug., Sept.
E. litus Coquillett ?

34-Sept.
E. pulvereus Melander * P

11-March, April, May
Eucessia reubens Coquillett

1-July
Exepac^nus johnsoni Coquillett * *

20-March, April, May
Exoprosopa arenicola Johnson
and Johnson

8-Aug.
E. caliptera Say *

8-April, May, Aug., Sept.
E. divisu Coquillett P *

25-June, July, Aug.
E. dorcadion Osten Sacken *

11-April, June, Aug., Sept.
E. doris Osten Sacken P *

68-July, Aug., Sept.
E. sharonae Johnson and Johnson

20-Aug., Sept.
E. utahensis Johnson and Johnson

16-July, Aug., Sept.
Geminaria canalis (Coquillett) * *

32-March, May, June
G. pellucida Coquillett

2-June, July
Geron argutus Painter *

12-May, July, Aug.
Heterostylwrn robustum

(Osten Sacken) * *

12-April, May, June
H. sackeni (Williston) P *

41-April, May
H. vierecki Cresson *

8-April, May, June, Oct.
Lepidanthrax agrestis (Coquillett) P

51-May, June, July, Aug.
L. angnlus Osten Sacken *

1-May
L. hyalinipennis Cole P

60-May, June, July
Lordotus abdominalis Johnson P

and Johnson

28-April, May, June, Sept.
L. albidus Hall P *

208-April, May, June
L. apicula Coquillett

63-May, June
L. singulatus Johnson and Johnson P

85-Sept., Oct.

The Great Basin Naturalist
10 ALLRED, JOHNSON, BECK Vol. XXV, Nos. 1-2

L. gibbus Loew ♦

10-Sept.
L. junceus Coquillett *

7-May, June
L. luteolus Hall P *

39-April, May, Sept., Oct.
L. melanosus Johnson and Johnson *

42-Sept.
L. nigriventris Johnson and

Johnson P * * *

116-March, April, May
L. perplexus Johnson and Johnson P *

22-April, May, Sept., Oct.
L. pulchrissimiis Williston *

9-Sept.
L. sororculus WilUston *

2-M:ay
L. striatus Painter * P

16-Sept., Oct.
Oligodranes ater (Cresson) * *

5-April, May
O. cinctura (Coquillett) *

1-April
O. distinctiis Melander * *

4-May
O. dolorosus Melander ? * * ♦

4-May, June
O. fasciola (Coquillett) * * p

14-June, Sept., Oct.
O. ryius (Bigot) *

4-Sept.
O. pulcher Melander ? *

4-June
O. pullatus Melander ? * P ,.

15-May, June
Pantarbes capita Osten Sacken * * *

6-April, May
P. pusio Osten Sacken *

7-April, May, June
P. willistoni Osten Sacken * *

5-April, May
Paracosnxus insolens Coquillett *

3-May
P. -inorrisoni Osten Sacken p * * * p

61-March, April, May, June,
July, Aug.
Poecilanthrax alpha

(Osten Sacken) *

11-Aug., Sept, Oct.
P. apache Painter and Hall * * * * P

127-Sept., Oct.
P. calif orniciis (Cole) ♦ * * p

77-Sept., Oct.
P. moffitti Painter and Hall *

29-Aug., Sept.
P. poecilagaster (Osten Sacken) *

1-Aug.

P. willistoni (Coquillett) * * P

62-Aug., Sept., Oct.
Toxophora pellucida Coquillett *

5-April, May, June

July 8, 1965 beeflies of Nevada 11

T. vasta Coquillett * P

12-June
T. virgata Osten Sacken * *

3-June, July, Oct.
Villa aenea Coquillett * * * * p

27-June, July, Aug., Sept.,
Oct.
V. arizonensis (Coquillett) *

4-June
V. atrata (Coquillett) *

1-July
V. cantor (Coquillett) P * P *

26-Sept., Oct.
V. crocina (Coquillett) P * * *

72-June, July, Aug., Sept.
V. cypris (Meigen) * * ♦

5-May, June, July
V. junctura (Coquillett) * *

20-April, May
V. lepidota (Osten Sacken) * P

37-July, Aug., Sept., Oct.
V. viira (Coquillett) *

1-July
V. morio (Linnaeus) p ♦ * *

37-April, May
V. scitula (Coquillett) * P

20-Sept.
V. sinuos^a (Wiedeniann) *

2-July
V. supina (Coquillett) * P

38- June, July
V. utahensis Maughan * * * P

49-April, May

'I.a-Fr = I-airea-Fianseiia; Gr-I.y = Grayia-I.yc iuni; Co = Oileogyne; At-Ko — Atriplex-
Kochia; Pi-Ju = Pinyon-Juniper; CS = Cane Springs; Mixed = areas not applicable to the desig-
nated coniniiuiities.

TULAREMIA AND DEER FLIES IN THE
ENVIRONS OF UTAH LAKE, UTAH'

Kaye B. Cox"

Introduction

Deer flies have been implicated as mechanical vectors of tula-
remia in Utah. Although infected flies heretofore have not been
found in nature, there is little doubt of their importance in trans-
mitting tularemia to man. Two species present in this area, Chrysops
discalis Williston and Chrysops noctifer Osten Sacken, have been
shown experimentally to transmit the disease. The presence of deer
flies in the environs of Utah Lake where tularemia is endemic offers
a potential health threat to man, and the expanding human popula-
tion and development of recreational facilities adjacent to the lake
increase this potential. Despite the fact that deer flies have been
implicated with tularemia in Utah, little is known about their dis-
tribution or seasonal occurrence in the environs of Utah Lake.

The objectives of this study are: (1) to determine the distribu-
tion and seasonal occurrence of deer flies in the environs of Utah
Lake; and (2) to determine the incidence of tularemia pathogens in
the deer flies.

Grateful acknowledgment is made to Dr. Dorald M. Allred for
the valuable suggestions and help given during this investigation, and
to Dr. Don H. Larsen for his assistance and criticism of the manu-
script. Special acknowledgment is made to Dr. Bert D. Thorpe,
director of the Epizoology Research Laboratory, Institute of Environ-
mental Biological Research, University of Utah, for the use of labora-
tory facilities in the bacteriological investigation, and for identifica-
tion of the pathogen. Verification of deer fly identifications was
made by Dr. Cornelius B. Philip, Principal Medical Entomologist,
Rocky Mountain Laboratory, United States Public Health Service,
Hamilton, Montana.

Review of Literature

Tularemia was first described by McCoy (1911) as a "plague-
like disease" of rodents in Tulare County. California. McCoy and
Chapin (1912) isolated the causative agent of the disease and named
it Bacterium tularense. The taxonomic position of the pathogen is
questionable. It is regarded by some as closely related to the pleuro-
pneumonia group of organisms, and by others as more closely re-
lated to the genus Brucella than to its present position in the genus
Pasteur ella (Burrows, Porter, and Moulder, 1959). The name Pas-
teurella tularensis (McCoy and Chapin), as listed in Bergey's Manu-

1. This investigation was supported in part 1)V a Predodoral (Intermediate) Fellowship, 1-Fl-
i'iM-11. <(18-01, from the National Institute of (kmeral Mcdiral Sciences, National Institutes of
Health. Rcthesda, Maryland.

1. Department of '/oology and Entomology, Brighani Young University, Piovo, Utah.

13

The Great Basin Naturalist

14 KAYE B. COX Vol. XXV, Nos. 1-2

al of Determinative Bacteriology, 7th ed. (Breed. Murray, and
Smith, 1957), will be used in this paper.

The taxonomy, description, and characteristics of Pasteurella
tularensis are found in publications of Breed, et^l. (1957), and Bur-
rows, et al. (1959). Hesselbrock and Foshay (1945) described the
morphology of the organism. Diagnostic procedures were described by
Burrows, et al. (1959), Davidsohn and Wells (1963), and Frankel,
Reitman, and Sonnenwirth (1963). Lillie and Francis (1936) de-
scribed the gross pathology of tularemia in laboratory animals. The
virulence of tularemia isolates from nature has been measured in
laboratory animals by Green (1931), Davis, PhiHp, and Parker
(1934), and Philip and Davis (1935). Definitive virulence tests have
been developed by Francis and Feltan (1942), Bell, Owen, and Lar-
son (1955), and Owen, et al. (1955). Jellison, et al. (1961) separated
North American isolates of P. tularensis from nature into two major
groups on the basis of virulence, reservoir host, seasonal distribution,
and geographical occurrence: ( 1 ) tick-borne tularemia of rabbits, and
(2) water-borne tularemia of rodents. Both groups are known to
occur in Utah (Woodbury, 1955; Jellison, Kohls, and Philip, 1951).
Tularemia in Utah was first recognized by Pearse (1911).
Francis (1919, 1921, 1929, 1937) listed numerous cases of human
infection, but it was not until 1937 that the disease was sufficiently
recognized by Utah physicians to accurately report its occurrence to
state health authorities (Woodbury, 1955). The Utah State Depart-
ment of Health reported 839 human cases of tularemia from 1937
to 1964. of which 26 were fatal (Thompson and Wright. 1964;
Wright, 1965). During this period. 136 cases were attributed directly
or indirectly to deer flies (Jenkins, 1965); 14 occurred in Utah
County. The seasonal incidence of tularemia in Utah for a 17-year
period coincides with the seasonal distribution of deer flies (Wood-
bury, 1955).

Tularemia in Utah was initially referred to as "deer-fly fever"
when Pearse (1911) associated the deer fly Chrysops discalis with
its transmission. Francis (1919, 1921), while investigating a disease
of unknown etiology in Utah, recognized "deer-fly fever," isolated
the pathogen, and named the disease tularemia. Although deer flies
have not heretofore been found infected with tularemia in nature
(Jellison, 1950). Francis and Mayne (1921) demonstrated that
female C. discalis could be experimentally infected and were capable
of transmitting the disease to laboratory animals. Chrysops noctifer
has also been implicated experimentally as a mechanical vector of
tularemia (Parker. 1933). Although only two species in the United
States have been shown to be experimental vectors of the disease.
Philip (1931) stated that other deer flies probably were capable of
mechanical transfer of tularemia.

Jellison and Parker (1944) concluded that the primary reservoir
of tularemia was rabbits, and Burroughs, et al. (1945) listed ani-
mals of 28 species as natural hosts of tularemia in the United States.
Tularemia has been found in 33 mammal and 34 bird species in the
Great Salt Lake Desert region of Utah (Bode, 1963), but Marchette,

July 8, 1965 tularemia and deer flies 15

et al. (1961) concluded that tularemia in this region is primarily a
disease of jack rabbits. Thirty-two of the mammal and 33 of the
bird species listed by Bode (1963) are known to occur in the Utah
Lake area (Bee, 1947; Woodbury, Cottam, and Sugden, 1949; Dur-
rant, 1962; Berrett, 1958; Hay ward, 1965).

Correlation of the geographical distribution of human tularemia
infections with the distribution of Chrysops discalis was shown by
Jellison (1950). He concluded that although no infected deer flies
had been found in nature and none had been found feeding on rab-
bits, circumstantial evidence indicated that C. discalis must be ac-
cepted as a vector of tularemia. Roth, Lindquist, and Mote (1952)
subsequently observed deer flies biting the ears of wild rabbits.

Deer flies are placed taxonomically in the subfamily Pangoniinae
of the family Tabanidae. The monograph of Brennan (1935) and
catalog of Philip (1947) with its supplement (1950) summarize the
present taxonomy of the Nearctic Pangoniinae and Tabanidae. The
Pangoniinae of Utah were described by Howe and Knowlton (1936).
Lewis (1949) listed the taxonomy of the Tabanidae of Salt Lake
County, Utah.

Distributional records of deer flies in Utah have been published
by Knowlton and Thatcher (1934), Philip (1947, 1950), and Mid-
dlekauff (1950). Chrysops fulvaster, C. aestuans, and C. discalis
were listed by Rowe and Knowlton (1936) as the common species
of deer flies in salt-marsh areas of Utah.

The biology and ecology of deer flies in the western United
States and Canada have been studied by Cameron (1926), Gjullin
and Mote (1945), Roth and Lindquist (1948), Lewis (1949), and
Roth, Lindquist, and Mote (1952).

Methods and Procedures

Eleven collecting sites in the environs of Utah Lake were chosen
following an initial survey in April, 1964. Selection was made on
the basis of plant association, larval habitat, and geographical loca-
tion. Locations of the sites are shown in Figure 1, and a brief de-
scription of each follows:''

1. Orem Quadrangle, R. 2 E. x T. 6 S., Section 28, west center of sec-
tion, elevation 4,500 feet. Station located 300 yards south-southwest
of Orem City Sewage Disposal Plant and end of Powell Slough ac-
cess road of the Utah Department of Fish and Game. Scirpus spp.
and Distichlis sp. present. Fresh-water springs and ponds.

2. Saratoga Springs Quadrangle, R. 1 W. x T. 5 S., Section 25. south-
east corner of section, elevation 4,500 feet. Station located 50 yards
south of Saratoga Springs resort swimming pools. Scirpus spp. and
Distichlis sp. present. Fresh-water springs and ponds.

3. Pelican Point Quadrangle, R. 1 E. x T .6 S., Section 32, west center
of section, elevation 4,500 feet. Station located 0.9 mile south-
southwest of Pelican Point pumping station. Scirpus spp. and
Distichlis sp. present. Fresh-water springs and drainage.

3. Topographical description of these sites was taken from United States (leolofiiial Survey
Topographical Maps, 7.5 minute series, scale 1:24,000.

The Great Basin Naturalist

16

KAYE B. COX Vol. XXV, Nos. 1-2

®

WH I TE

LAKE

- GOSHEN
(V} WARM

SPRINGS

Fig. 1. Deer fly collection sites in the environs of Utah Lake, Utah.

July 8, 1965 tularemia and deer flies 17

4. Soldiers Pass Quadrangle, R. 1 W. x T. 8 S., Section 9, southeast
corner of section, elevation 4,500 feet. Station located 1.1 miles south-
east of Clyde Knoll. Scirpus spp. and Distichlis sp. present. Inter-
mittant backwater of lake, no fresh-water springs.

5. Goshen Valley North Quadrangle, R. 1 W. x T. 9 S., Section 26,
center of section, elevation 4,490 feet. Station located 3.5 miles north-
northwest of Goshen school. Scirpus spp. and Distichlis sp. present.
Intermittant fresh-water drainage stream.

6. Santaquin Quadrangle, R. 1 E. x T. 10 S., Section 8, west center of
section, elevation 4,520 feet. Station located at Goshen Warm Springs.
Scirpus spp. and Distichlis sp. present. Fresh-water springs and
ponds.

7. West Mountain Quadrangle, R. 1 E. x T. 9 S., Section 17, north
center of section, elevation 4,500 feet. Station located 2.2 miles west
of West Mountain peak, VABM 6904, and end of Lebaron Site ac-
cess road of Utah Department of Fish and Game. Scirpus spp. and
Distichlis sp. present. Fresh-water springs and ponds.

8. West Mountain Quadrangle, R. 1 E. x T. 9 S., Section 20, east center
of section, elevation 4,500 feet. Station located 2.1 miles west-north-
west of Kiegley on east border of White Lake. Scirpus spp. and
Distichlis sp. present. Fresh-water springs, small ponds and drainage.

9. Lincoln Point Quadrangle, R. 1 E. x T. 8 S., Section 11, center of
section, elevation 4,490 feet. Station located 0.7 mile south-southeast
of Lincoln Point benchmark BM 4526, near mouth of Benjamin
Slough. Scirpus spp. and Distichlis sp. present. Fresh-water slough,
springs, small ponds and drainage.

10. Provo Quadrangle, R. 2 E. x T. 8 S., Section 4, north center of
section, elevation 4,500 feet. Station located 0.3 mile north of inter-
section of highway U 228 and Springville road. Scirpus spp. and
Distichlis sp. present. Fresh-water springs, drainage and standing
roadside water.

11. Provo Quadrangle, R. 3 E. x T. 7 S., Section 17, west center of sec-
tion, elevation 4,500 feet. Station located 100 yards west of General
Offices, Pacific States Cast Iron Pipe Company. Scirpus spp. and
Distichlis sp. present. Fresh-water springs, ponds, drainage and small
stream.

Each site was visited for a one-hour period between 9 a.m. and
2 p.m. every two weeks from May through October. Collections
were made on three consecutive days: Sites 1-4 were visited one
day, Sites 5-8 the following day, and Sites 9-11 the succeeding day,
alternating the hours of visitation of the areas collected on any given
day. Adult female deer flies were attracted to me as I walked briskly
through each area in a systematic pattern, stopping for several min-
utes every ten paces. As the attracted flies rose to attack or alighted
on me, they were captured with an aerial insect net. Flies were
frequently attracted to my automobile and were taken from its
metal surfaces. Attempts at aerial capture of hovering flies or sweep-
ing of vegetation proved futile. Captured flies were placed in an
ethyl acetate killing jar.

After collections were completed at each site, the deer flies were
removed from the killing jar and identified. The specimens were sep-
arated to species, thoroughly washed in sterile physiological saline
solution, and put into four-dram screw-cap vials containing sterile,
non-fat skim milk solution. Each vial was labelled externally with
collection data, and frozen and stored at -30 C for subsequent bac-

The Great Basin Naturalist

18 KAYE B. COX Vol. XXV. Nos. 1-2

teriological testing for the tularemia pathogen. Collection and other
appropriate data were recorded for each site and collection. Tem-
perature data were obtained from United States Department of Com-
merce Weather Bureau Climatological Data reports for Utah Lake,
taken at Lehi, Utah.

Bacteriological investigations were conducted at the Epizoology
Research Laboratory, Institute of Biological Environmental Research,
University of Utah, Salt Lake City. Frozen deer fhes were thawed
and triturated in a mortar and pestal. Sterile, non-fat skim milk
was added when required to insure sufficient innocula. Seventy-three
pools were prepared, each containing not more than 15 flies of the
same species, separated by area and date of collection. The super-
natant from each pool was innoculated into one adult I lartley strain
guinea pig and four adult Swiss-Webster strain white mice. Intra-
peritoneal injections of 1.0 cc per guinea pig and 0.5 cc per white
mouse were given. Injected animals were caged and observed twice
daily for a period of 28 days. Dead animals were removed on dis-
covery, placed into separate containers, and stored at -30 C to await
subsequent processing. Animals found dead within 24 hours after
innoculation were discarded.

Aseptic necropsy procedures were used in the examination and
processing of the dead animals. Each carcass was swabbed with
70 percent ethyl alcohol, the outer skin peeled back, and the thoracic
and peritoneal cavities exposed. The gross pathology of the spleen
and liver was noted, particularly for the swelling and mottling
characteristic of tularemia infections. Sections of the spleen and
liver were removed and plated on culture medium. After plating,
the tissues were put into sterile vials and frozen for subsequent use.
The culture medium used in testing for the tularemia pathogen was
blood cystine heart agar (BCHA). It was prepared by dissolving 51
g of Difco Cystine Heart Agar (B47) in 950 ml of distilled water by
heating in an 80 C water bath, adjusting to pi I 7. autoclaving, and
adding 50 cc of outdated, citrated human blood after cooling to 50 C
in a water bath.

The plated BCIIA media were incubated for 48 hours at 37 C
and examined for characteristic colony growth. Typical colonies
were subjected to Gram stain for determination of morphology and
stain reaction. Slide agglutination tests were made on cultures dem-
onstrating characteristic colony growth, morphology, and stain re-
action. A slide agglutination titer of 1:80 or higher with complete
agglutination was considered positive. Plates with positive slide ag-
glutination test results, typical growth colonies, characteristic mor-
phology, and Gram negative reaction were considered positive for
PasteurelJa tularensis.

Immediately after identification of the pathogen, the original
tissue samples composing the positive isolation were examined and
the pathogen reisolated from the infected tissues, confirming the
initial isolation. Determination of the LD-„, was made by testing
various dilutions of the positive isolates in mice, rats, guinea pigs,
and rabbits. Maximum virulence was established as an LDr,,, of nine

July 8, 1965

TULAREMIA AND DEER FLIES

19

organisms or less for the test animals used. Test animals surviving
the 28-day observation period were bled and the sera collected. The
sera were subjected to a standard tube agglutination test for tula-
remia, using a known positive of 1:640 titer as a control. Final
identification of Pasteurella tularensis was made on the basis of typi-
cal colony characteristics, morphology, animal pathogenicity, pathol-
ogy, and slide agglutination test reactions.

Results

A total of 823 deer flies representing three spices was collected
during this study TTable 1;. Of the total number taken. 97.8 percent
were Chrysops fulvaster van der Wulp. 1.92 percent were C. aestuans
Osten Sacken. and 0.24 percent were C. discalis Williston. Chrysops
fulvaster was collected from ten, C. aestuans from seven, and C.
discalis from two sites. The greatest population occurred during the

Table 1. Distribution of deer flies by species, site, and collection
period.

Collection Period

c.gi

Site

S i

Si ">-

>■._>^

_>^ tf

ti b£

ti't

•^ t;

■s ^•

Site

Species'

--

■=-.-.

^.-.

-.<

<<

<y.

X Ji

Ji C

Total

1

Cf

1

8

60

2

71

2

Cf
Ca

2

6

28
1

56
3

2

94
4

3

Cf
Ca

1

6

30
1

1

38
1

4

None observed

5

Cf

4

4

6

Cf
Ca

1

4

27
1

21

1

1

3

57
2

7

Cf
Cd

2

4

43

114
1

102

17

11

1

294
1

8

Cf

3

29

15

35

15

1

1

99

9

Cf
Ca

2

5

11

1
1

19
1

10

Cf
Ca
Cd

3

9

14
5

5

1

1

31
6

1

11

Cf
Ca

1

7
2

68

22

98
2

Totals

Cf
Ca
Cd

15

73
2

272
7

290
7
2

121

21

12

1

805

16

2

^Cf = Chrysops fulvaster, Ca = Chrysops aestuans, Cd = Chrysops discalis.

20

KAYE B. COX

The Great Basin Naturalist

Vol. XXV, Nos. 1-2

July 26-August 8 period. Female deer flies first appeared during
the June 14- June 27 period, approximately one month after the last
freezing temperature in May. Greatest populations coincided with
weekly average maximum temperatures of 75 F or higher (Fig. 2).
Deer flies were not observed after the September 20-October 3 period
when temperatures of 32 F or below occurred. Positive thermotaxic
response was noted for C. fulvaster.

i_ ^i_ ^w >i_ >> i/i> t/>co 0</> CO oo

c cc cc cc cc (PC ci) n rv(B on rtrt
3 —3 (D — <Q<0 10<0 TJ'O -•"D -^ " -• -»

2> 22 22 '-^

Q-O Q Q Q S C 0

*<"* "^^ ^"^ 3^ -* ~J — :j ^ «j — lii'ij y\ij u>J -*u ~^ ~^ -*■-*•

— <— • CO— •— '(JIO— — •Kj'*^— ' K3IO »o— K)-*..CJ— •

KJ>0 OOJ O^ to— SI *>• —00 l-nw 00O> lO-O OlO) <3 O oJO vl *^ ^00

I II 1^1 II I I I I 1 I I

■ — . = Ave maximum temperature ^_^__ = Total deer flies O- 33-34 F during week

= Ave minimum temperature ■ = 32 F or below during

week

Fig. 2. Total numbers of deer flies captured each two-week period and
weekly average maximum and minimum temperatures.

July 8, 1965 tularemia and deer flies 21

Sera tube agglutination tests of laboratory animals surviving the
28-day observation period were negative. Gross pathology was in-
consistent. Pasteurella tularensis of maximum virulence was isolated
from three of 73 pools tested. Isolations were made from one pool of
five Chrysops fulvaster collected at Site 9, Lincoln Beach, on July
3, one pool of three C. aestuans collected at Site 2, Saratoga Springs,
on July 27, and one pool of four C. fulvaster collected at Site 5,
Goshen, on July 29 (Table 2). Slide agglutination tests of isolated
and reisolated pathogen organisms demonstrated a titer of 1 : 80 or
higher. Virulence tests of the isolates resulted in an LD^u of 10"
(1 to 9 organisms).

Table 2. Isolation of Pasteurella tularensis from deer flies.

V,

§■

1

0)

1

4)

"2^

o

^1

a1

CO

z.s

4)

Q

jiy

2D2

Chrysops aestuans

July 27

2

3

6

10"^

5D

Chrysops fulvaster

July 29

5

4

5

10°

9B

Chrysops fulvaster

July 3

9

5

4

10°

'Day of death of laboratory animal following injection for initial isolation.
'Nine or less organisms causing death.

Discussion
Occurrence and Distribution of Deer Flies

The three species of deer flies collected during this study, Chry-
sops aestuans, C. discalis, and C. fulvaster, were previously noted in
the Utah Lake area by Knowlton and Thatcher (1934), and were
listed by Rowe and Knowlton (1936) as those most common to salt-
marsh areas in Utah. Other species have been listed for Utah from
widely separated geographical areas and varied habitats, but the ex-
tent of their distribution in the state is largely unknown (Knowlton
and Thatcher, 1934; Rowe and Knowlton, 1936; Philip, 1947, 1950;
Middlekauff, 1950). One record of Chrysops lupus Whitney was
listed for Provo by Knowlton and Thatcher (1934), but no subse-
quent collections of this species have been reported in this area.

The period of activity of the deer flies in this study corresponds
with the observations of Gjullin and Mote (1945) at Summer Lake,
Oregon, who noted that deer flies were active from the middle of
June to the first part of September. Roth and Lindquist (1948) noted
that the peak abundance of flies for a two-year period occurred during
the last part of July and the first part of August. Cameron (1926)
noted activity of C. aestuans from the first part of June into July.

The seasonal occurrence of the flies collected during this study
showed variation with different species. The flies first observed were

Die (iieat Rnsin Niiluivilist

22 KAYE B. COX Vol. XXV. Nos. 1-2

C. fulvaster on June 24. These were subsequently taken at various
sites until September 26, the peak abundance occurring during the
July 26-August 8 period. The limited number of C. aestuans collected
precludes conclusions as to their seasonal occurrence and distribution,
but their activity during June and July corresponds with the obser-
vations of Cameron (1926). Comparative numbers of C. aestuans
and C. fulvaster collected in this study support the contention of
Rowe and Knowlton (1936) that C. aestuans is often taken with C.
fulvaster but is less abundant. Insufficient collections of C. discalis
preclude any conclusion on its activity other than that it was present
in the study area. Its absence in habitats with which it is usually
associated suggests that some factor or combination of factors resulted
in the minimal number noted in this study.

The most abundant species, C. fulvaster^ was collected at all sites
at which activity was noted (Table 1), which suggests that the con-
ditions at these sites were favorable for this species. Chrysops aestu-
ans was taken at six sites. Its presence at some sites and absence
from others of almost identical habitat is unexplained other than
that the species is not abundant in the study area, as noted by Rowe
and Knowlton (1936). Chrysops discalis was coptured at two sites.
No explanation is offered as to the scarcity of this species in typical
habitat sites in the study area other than that extrinsic factors may
have reduced its numbers.

The daily activity of deer flies between 9 a.m. and 2 p.m. cor-
responds with that noted by Roth and Linquist (1948) and Roth
et al. (1952). The period of activity may be attributed to warming
of cool morning air to an optimum temperature. Few flies were ac-
tive between 2 p.m. and 5 p.m., but a slight increase was noted at
dusk. Little activity was observed during periods of strong wind, re-
gardless of time of day.

The positive thermotaxic response to the metal surfaces of my
automobile when stationary in open sunlight, particularly during
early morning hours, as noted for Chrysops fulvaster, is similar to
that observed for C. aestuans by Cameron (1926) and for C. discalis
by Lewis (1949).

Factors Affecting Deer Fly Populations

Deer flies are associated by oviposition with particular plants.
In studies at Summer Lake, Oregon, oviposition was noted on the
sedges Scirpus americanus and S. paludosus (Gjullin and Mote.
1945; Roth and Lindquist, 1948). Lewis (1949) described oviposi-
tion by C. discalis on the salt-marsh grass Distichlis stricta in Salt
Lake County. Cameron (1926) observed oviposition by C. aestuans
on Scirpus sp. and other emergent aquatic plants. The major cri-
terion for the selection of collection sites in this study was the pres-
ence of Scirpus sp. and Distichlis sp., although such factors as water
source and area, larval habitat, and geographical location also were
considered. No flies were observed at Site 4 even though it possessed
the plant species usually associated with deer flies. Although Roth

July 8, 1965 tularemia and deer flies 23

and Lindquist (1948) encountered flies up to two miles from breed-
ing areas, the absence of flies from an area with typical associated
plant species suggests that other factors may be essential for deer
fly populations. All collecting sites except Site 4 were characterized
by fresh-water springs, ponds, or drainage. The water at Site 4 was
brackish, intermittent backwater of Utah Lake, with a sand and
gravel bottom. The w^ater sources at all other sites varied in size,
but all drained alkaline soil, and the bottom of the water area was
covered with thick, black muck, rich in organic debris typical of
lentic habitats. The abundance of deer flies in these areas substanti-
ates findings in their life history by Philip (1931) and Roth and
Lindquist (1948) who discovered larvae and pupae in similar habi-
tats. Philip (1931) described larval habitats of deer flies and associ-
ated C. aestuans with lentic conditions, and C. fuivaster with both
lotic and lentic conditions. Lewis (1949) found C. discalis in brack-
ish, alkaline ponds in lentic associations. The life cycle and abun-
dance of deer flies appears to be dependent upon proper larval and
pupal habitats as well as particular plant species for oviposition.
The absence of flies at one site where appropriate plant species were
present but where proper water habitat and larval and pupal en-
vironments were missing, supports this allegation.

The size of the breeding area may influence deer fly populations.
Except for Site 4, all sites had alkaline drainage and bodies of water
of various sizes. Sites 1.6, 7, and 8 had very large water areas, and
a comparison of the number and extent of deer fly collections at
these sites (Table 1 ) reveals that all except Site 1 were character-
ized by large numbers of flies, an activity period of long duration,
and a near normal curve of fly activity during the study period.
Sites 3, 9, 10, and 11 had relatively small water areas, and were
characterized by increasing numbers of flies until the July 26-August
8 period when the activity decreased abruptly, and then ceased en-
tirely by the August 9-August 22 period. At Site 5, where a small
intermittent stream drained an alkaline meadow, there was no ac-
tivity during the study except for the July 26-August 8 period when
large amounts of water were present and overflowed the stream into
the adjacent meadow. The flies captured at this site may have been
transients from nearby areas. Comparison of the number and activi-
ty of deer flies with the size of the water area and type of drainage
indicates a correlation between the two factors.

Comparison of the number of deer flies collected each two-week
period and the average weekly maximum and minimum tempera-
tures (Fig. 2) suggests a relationship between temperature and deer
fly activity. The weekly average maximum temperature rose to ap-
proximately 70° F before initial deer fly activity was observed, and
maximum activity was noted when the weekly average maximum
temperatures were 75° F or higher. This agrees with the findings of
Roth et al. (1952) who observed greater activity of C. discalis when
the temperature was above 75° F, and Davies (1959) who noted
that peak abundance of Chrysops spp. occurred when the maximum
temperature rose to 80° F, especially when this temperature was

The Great Basin Naturalist

24 KAYE B. COX Vol. XXV, Nos. 1-2

several degrees higher than the maximum of the preceding few days.
The activity of the flies dropped sharply when the first near-freez-
ing temperatures occurred during the week of August 23-29, and
activity ceased with freezing temperatures during the last weeks of
September. Freezing temperatures in the weeks of April 19 to May
9 may account for the minimal collections of C. discalis in this study.
According to the observations of Lewis (1949), this species is nor-
mally active from April to September. If adult flies had emerged
and were active as described, the freezing temperatures of April and
May would have killed most of the emergent flies, and only those
surviving or emerging after the freezing temperatures would have
been present during this study. Chrysops fulvaster and C. aestuans
apparently were not affected by the freezing spring temperatures,
suggesting that these species emerge later than C. discalis. The sharp
decline in activity at some sites when temperatures near 100° F
occurred during the last three days of July agrees with the findings
of Jamnback and Wall (1959) who observed that extended periods
of high temperatures resulted in a sharp decrease in deer fly activity.
The gradual decrease in activity at Sites 6, 7, and 8 may have re-
sulted from an insulating effect of the large water areas at these
sites and offered some protection from the effects of high tempera-
ture.

Deer fly activity at Site 1. where extensive water area, alkaline
drainage, and associated plant species were present, declined abrupt-
ly after aerial spraying for mosquito control on July 27. A total of
68 Chrysops fulvaster had been collected at Powell Slough during
the previous collecting period, but on the day of spraying only two
specimens were taken, and none thereafter. The Utah County Health
Department entomologist (Davis, 1965) revealed that the spray used
was Parathion, applied at one pound per acre in an oil emulsion, and
that spraying also had been done in the areas of Sites 1, 2, 9, and 11
during May and the first week in June. The effect of the May and
June insecticide applications on C. fulvaster and C. aestuans is un-
known, since no activity was noted for these species until later in
June and July, several weeks after spraying. If the seasonal dynam-
ics of C. discalis in the study area were similar to the activity noted
by Lewis (1949), the May and June applications of insecticide may
well account for the small numbers of this species collected during
this study. The apparent complete absence of deer flies following the
July 27 application of insecticide suggests that aerial spraying may
effectively control adult female deer fly populations.

The species occurrence and geographical distribution of deer
flies observed in this study may be related to several factors. Even
though the 1 1 collecting sites in the environs of LTtah Lake were se-
lected on the basis of specific plant associations, larval habitat, and
general environment similar to those noted by previous workers
(Cameron, 1926; Crjullin and Mote, 1945; Roth and Lindquist, 1948;
Lewis, 1949; Roth et al.. 1952), the sites may not have possessed
habitats favorable to all species of deer flies. In order to insure as
large a number of deer flies as possible for subsequent bacteriological

July 8, 1965 tularemia and deer flies 25

examination, sites with apparent atypical habitats were not selected.
An attempt was made to collect from a variety of habitats and to
encompass the periphery of the lake and its environs. When possible,
sites were selected on the basis of their proximity to areas of human
habitation, agriculture and recreation. Collections from additional
sites were limited by time, methods, and transportation requirements.

Isolation of Tularemia Pathogens

The isolation of Pasteurella tularensis from three pools of flies
containing Chrysops fulvaster and C. aestuans substantiates the con-
clusion of Philip (1931) that deer flies other than C. discalis and
C. noctifer, which had been implicated as mechanical vectors in
laboratory experiments (Francis and Mayne, 1921; Parker, 1933),
may be involved as vectors of tularemia. Pathogens were found in
flies during the time of greatest fly populations and seasonal period
when the greatest numbers of human tularemia cases were recorded
over a 17-year period in Utah (Woodbury, 1955). Pathogens were
also found in deer flies taken during an earlier period when fly
populations were small, but well within the seasonal period of high-
est incidence in humans as noted by Woodbury (1955). The corre-
lation of deer fly populations observed during this study with the
seasonal incidence of human infections strongly implies that deer
flies are a major source of human infections in Utah. Utah State
Department of Health statistics list 136 of 839 cases of human in-
fections as being directly or indirectly associated with deer flies, but
703 reports did not state the source of infection (Jenkins, 1965). It
is possible that many of the unstated cases could be attributed to
deer flies.

J he geographical distribution of the infected flies does not show
a significant pattern, although the distribution suggests that an en-
demic tularemia focus may be located at the southern end of Utah
Lake.

The presence of a highly virulent strain of the pathogen indi-
cates that a natural reservoir exists among the native fauna. Whether
such reservoir animals are resistant to the strain is not known. No
evidence of an epizootic was observed during the study in any of
the areas, although many of the animal species known to be infected
with tularemia in Utah (Bode, 1963) are known to occur in the
Utah Lake area (Bee, 1947; Woodbury et ai, 1949; Durrant, 1952;
Berrett, 1958; Hay ward, 1965). Rodents and lagomorphs were oc-
casionally observed in collecting areas, particularly near Site 2,
where large numbers of squirrels were present at a garbage dump
adjacent to a resort.

Although it is possible that only three of 823 deer flies were in-
fected, the isolation of the pathogen from deer flies is medically sig-
nificant. Previous implications of deer flies as mechanical vectors
of tularemia have been based on the results of experimental evi-
dence for two species, C. discalis and C. noctifer (Francis and
Mayne, 1921; Parker, 1933), and to this time no infected flies have

The Great Basin Naturalist

26 KAYE B. COX Vol. XXV, Nos. 1-2

been taken in nature (Francis and Mayne, 1921; Jellison, 1950).
It has been assumed on the basis of circumstantial evidence (Jelli-
son, 1950), that deer flies are responsible for the transmission of
tularemia to humans. Tularemia has been found in rodents, lago-
morphs, and other vertebrates (Burroughs et al., 1945; Bode, 1963);
deer flies have been observed biting rabbits (Roth and Lindquist,
1948) and have been associated with human tularemia infections
(Pearse, 1911; Francis, 1919, 1929; Jellison, 1950; Jenkins, 1965);
and now to complete the chain of infection, Pasteurella tularensis
has been found in deer flies in nature.

Recommendations for Further Study

The isolation of tularemia pathogens from deer flies in areas
where human infection is probable is significant, and on this basis
additional studies should be undertaken. Suggestions for further in-
vestigation arising from this study are:

( 1 ) A comprehensive study should be made of a wide selection
of habitats and geographical areas over several years to determine
the occurrence, distribution, and seasonal dynamics of all the species
of biting flies in the environs of Utah Lake.

(2) An extensive investigation of deer flies should be made to
determine the time, place, and duration of infection by Pasteurella
tularensis.

(3) An investigation of the pathogen isolates should be made to
determine the strains present and their virulence.

(4) A comprehensive study of the potential reservoir animals
in the environs of Utah Lake should be made to determine species,
populations, distribution, and seasonal occurrence of the animals,
and the incidence and virulence of tularemia in the native fauna.

(5) An investigation of the human population in the environs
of Utah Lake should be made to determine the extent of clinical
and subclinical infections.

Summary and Conclusions
On the basis of this study the following data are presented:

1. A total of 823 adult female deer flies representing three
species was collected during the spring, summer, and autumn
of 1964 from 11 collecting sites in the environs of Utah Lake,
Utah. Chrysops julvaster was the most abundant, Chrysops
aestuans the next, and Chrysops discalis the least abundant.

2. Deer flies were found only in areas with fresh-water springs,
ponds, or drainage.

3. Deer flies appeared approximately one month after the last
freezing temperatures in May, and disappeared after the first
freezing temperatures in September. Nlaximum activity of

July 8, 1965 tularemia and deer flies 27

the flies occurred between July 26 and August 8 when the
weekly average maximum temperature was 75° F or higher.

4. The abundance of deer flies appeared to be dependent upon
the size of breeding areas, typical larval habitats, and the
presence of particular plant species.

5. Positive thermotaxic response was noted for Chrysops ful-
vaster.

6. The causative agent of tularemia, Pasteurella tularensis, was
isolated from three of 7?) pools of deer flies. Isolation was
made from one pool of five Chrysops ful vaster collected at
Lincoln Beach on July 3. one pool of three C. aestuans col-
lected at Saratoga Springs on July 27, and one pool of

four C. julvaster collected at Goshen on July 29.

7. The presence of tularemia pathogens in deer flies near areas
of human habitation, agriculture, and recreation in the en-
virons of Utah Lake, Utah, represents a public health threat
for which appropriate preventive measures should be taken.

8. Aerial spraying of appropriate insecticides may control adult
female deer fly populations.

LiTER.ATURE Cited

Bee, J. W. 1947. Mammals of Utah County. Unpublished Master's thesis,
Brigham Young Univ., Provo, Utah. 316 pp.

Bell, J. F., C. R. Owen, .-^nd C. L. L.^rson. 1955. Virulence of Bacterium
tularense. I. A study of virulence of Bacterium tularense in mice, guinea
pigs and rabbits. J. Infect. Dis., 97:162-166.

Berrett, D. G. 1958. Bird Populations of the Wasatch Foothills. Unpublished
Master's thesis, Brigham Young Univ.. Provo, Utah. 156 pp.

Bode, D. D. and Staff. 1963. A Study of the Ecology and Epizoology of the
Native Fauna of the Great Salt Lake Desert. Annual report of the director
and staff, Ecol. and Epizool. Ser. No. 100, Ecol. and Epizool. Res., Univ. of
Utah, Salt Lake City, Utah. 156 pp.

Breed, R. S., E. G. D. Murray, and N. R. Smith. 1957. Bergey's Manual of
Determinative Bacteriology. 7th ed. Williams & Wilkins Co.. Baltimore.
1094 pp.

Brennan, J. M. 1935. The Pangoniinae of Nearctic America. Diptera:
Tabanidae. Univ. Kansas Sci. Bull., 22:249-401.

Burroughs, A. L., R. Holdenreid, D. S. Longanecker, and K. F. Meyer. 1945.
A field study of latent tularemia in rodents with a list of all known natural-
ly infected vertebrates. J. Infect. Dis., 76:115-119.

Burrows, W., R. J. Porter, and J. W. Moulder. 1959. Te.xtbook of Micro-
biology. 17th ed. W. B. Saunders Co., Philadelphia. 954 pp.

Cameron, A. E. 1926. Bionomics of the Tabanidae of the Canadian prairie.
Bull. Entomol. Res., 17:1-42.

Davidsohn, I. and B. B. Wells. 1963. Todd-Sanford Clinical Diagnosis by
Laboratory Methods. 13th ed. W. B. Saunders Co., Philadelphia. 1020 pp.

Davies, D. M. 1959. Seasonal variations of Tabanids in Algonquin Park. On
tario. Can. Ent., 91:548-553.

Davis, G, E.. C. B. Philip, and R. R. Parker. 1934. Isolation from the Rocky
Mountain wood tick of strains of Bacterium tularense of low virulence for
guinea pigs and domestic rabbits. Amer. J. Hyg., 19:449-456.

Davis. T. 1965. Personal communication, March 4.

DuRRANT, S. D. 1952. Mammals of Utah. Univ. Kansas Pub., Mus. Natur.
Hist., 6:1-549.

The Great Basin Naturalist

28 KAYE B. COX Vol. XXV, Nos. 1-2

Francis, E. 1919. Deer-fly fever; a disease of man of hitherto unknown eti-
ology. U. S. Public Health Repts., 34:2061-2062.

. 1921. Tularemia Francis 1921. I. The occurrence of tularemia in nature

as a disease of man. U. S. Public Health Repts., 36:1731-1738.

. 1929. Arthropods in the transmission of tularemia. Trans. Fourth Int.

Congr. Entomol., 2:929-944.

. 1937. Sources of infection and seasonal incidence of tularemia in man.

U. S. Public Health Repts., 52:103-113.

. AND L. D. Felton. 1942. Antitularemic serum. U. S. Public Health

Repts., 50:44-55.

AND B. Mayne. 1921. Tularemia Francis 1921. II. Experimental trans-

mission of tularemia by flies of the species Chrysops discalis. U. S. Public

Health Repts., 36:1738-1746.
Frankel, S., S. Reitman, and A. C. Sonnenwirth. 1963. Gradwohl's Clinical

Laboratory Methods and Diagnosis. 6th ed. C. V. Mosby Co., St. Louis.

2093 pp.
Green, R. G. 1931. Occurrence of Bacterium tularense in the eastern wood

tick, Dermacentor variabilis. Amer. J. Hyg., 14:600-613.
Gjullin, C. M. and D. C. Mote. 1945. Notes on the biology and control of

Chrysops discalis Williston. Proc. Entomol. Soc. Wash., 47:236-244.
EUyward, C. L. 1965. Personal communication, March 11.
Hesselbrock, W. and L. Fosh.\y. 1945. Morphology of Bacterium tularense.

J. Bact., 49:209-231.
Jamnback, H. and W. Wall. 1959. The common salt-marsh Tabanidae of

Long Island, New York. New York State Mus. Bull, 375:1-77.
Jellison, W. L. 1950. Tularemia; geographical distribution of "Deer-fly fever"

and the biting fly, Chrysops discalis Williston. U. S. Public Health Repts.,

65:1321-1329.
. AND R. R. Parker. 1945. Rodents, rabbits and tularemia in North

America: Some zoological and epidemiological considerations. Amer. J. Trop.

Med., 25:349-362.
. G. M. Kohls and C. B. Philip. 1951. Tularemia: Muskrats as a

source of human infection in Utah. Rocky Mountain Med. J., 48:594-597.
C. R. Owen, J. F. Bell, and G. M. Kohls. 1961. Tularemia and ani-

mal populations: Ecology and epizootiolog>'. Wildlife Dis., 17:1-19.
Jenkins, A. A. 1965. Personal correspondence, March 19.
Knowlton, G. F. and T. O. Thatcher. 1934. Utah horseflies. Proc. Utah

Acad. Sci., Arts, and Letters, 11:291-294.
Lewis, W. 1949. The Taxonomy and Notes on Bionomics and Control of the

Tabanidae of Salt Lake County, Utah. Unpublished Master's thesis, Univ. of

Utah, Salt Lake City, Utah. 86 pp.
LiLLiE, R. D. AND E. Francis. 1936. The pathology of tularemia in mammals.

Nat. Inst. Health Bull., 167:83-202.
Marchette, N. J., D. L. Lundgren, P. S. Nicholes. and E. D. Vest. 1961.

Studies on infectious diseases in wild animals in Utah. I. Susceptibility of

wild mammals to experimental tularemia. Zoonoses Res., 1:49-73.
McCoy, G. W. 1911. A plague-like disease of rodents. U. S. Public Health Bull..

43:53-71.
. AND C. W. Chapin. 1912. Studies of plague, a plague-like disease, and

tuberculosis in California. V. Bacterium tularense the cause of a plague-like

disease of rodents. U. S. Public Health Bull., 53:17-23.
Middlekauff, W. W. 1950. The horseflies and deerflies of California. Bull.

Calif. Insect Surv., 1:1-26.
Owen, C. R., J. F. Bell, C. L. Larson, and R. A. Ormsbee. 1955. Virulence

of Bacterium tularense. II. Evaluation of criteria of virulence of Bacterium

tularense. J. Infect. Dis., 97:167-176.
Parker, R. R. 1933. Recent studies of tick-borne diseases made at the United

States Public Health Service Laboratory at Hamilton, Montana. Proc. Fifth

Pacific Sci. Congr., 6:3367-3374.
Pearse. R. a. 1911. Insect-bites. Northwest Med., 3:81.
Philip. C. B. 1931. The Tabanidae (horse-flies) of Minnesota with reference

to their biologies and taxonomy. Minn. Agr. Exp. Sta. TecJi. Bull., 8:1-32.

July 8, 1965 tularemia and deer flies 29

. 1947. A catalog of the blood-sucking fly family Tabanidae (Horse-flies

and deer-flies) of the Nearctic region north of Mexico. Amer. Midland Natur,.
37:257-324.

. 1950. Corrections and addenda to a catalog of Nearctic Tabanidae.

Amer. Midland Natur., 43:430-437.

AND G. E. Davis. 1935. Tularemia: Observations on a strain of low

virulence from rabbit ticks. U. S. Public Health Repts., 50:909-911.
Roth, A. R. and A. W. Lindquist. 1948. Ecological notes on the deer-fly at

Summer Lake, Oregon. J. Econ. Entomol., 41:473-476.
, , AND D. C. Mote. 1952. Incidence and habits of Tabanidae at

Summer Lake, Oregon. J. Econ. Entomol., 45:527-531.
RowE, J. A. AND G. F. Knowlton. 1936. Pangoniinae of Utah (Tabanidae:

Diptera). Ohio J. Sci., 36:253-259.
Thompson, G. D. C. and J. W. Wright. 1964. Annual Report, 1963, Division

of Vital Statistics. Utah State Department of Health, Salt Lake City, Utah.

136 pp.
Woodbury, A. M. 1955. Ecological studies of tularemia in Utah. In Symposium

on Ecology and Disease Transmission in Native Animals. U. S. Army

Chemical Corps, Dugway, Utah. 112 pp.
, C. CoTTAM, and J. W. SuGDEN. 1949. Annotated checklist of the birds

of Utah. Bull. Univ. Utah, 39(16): 1-40.
Wright, J. W. 1965. Personal communication, March 4.

THE GENUS EUPAGIOCERUS BLANDFORD'
(SCOLYTIDAE: COLEOPTERA)

Stephen L. Wood"

A recent opportunity to study the habits of three species of
Eupagiocerus Blandford in Central America makes possible the illu-
cidation of the biology of this genus for the first time. Two pre-
viously undescribed species are also added to the two previously
known in the genus. New synonymy for one species is included.

Eupagiocerus Blandford

Eupagiocerus Blandford, 1896, Biol. Centr.-Amer. 4(6): 133.
Nemopagiocerus Schedl, 1962, Mitt. Miinchner Ent. Gesellsch. 52:85
(new synonymy).

On the basis of the antennal characters, the entire eye, and the
tuft of pleural hair on the prothorax, Schedl (1962:85) erected the
genus Nemopagiocerus for his Eupagiocerus nevermanni. This
species and the two described below as new were studied in the field,
and all three species were compared to the type of dentipes Blandford
All three species breed in the pith of woody vines where they have
habits that fall well within the limits of the closely allied genus
Cnesinus. Since those species of Cnesinus associated with ambrosial
fungi may also have a pleural tuft of setae on the prothorax similar
to that of Schedl's nevermanni, while those not associated with
fungi lack this character, its presence or absence should have no
generic significance. The strongly procurved sutures of the antennal
club in clarus, while more nearly like Schedl's species, are inter-
mediate between nevermanni and dentipes Blandford. The inner
margin of the eye in Schedl's species varies from entire to sinuate,
and in clarus, dentipes and vastus from rather strongly sinuate to
shallowly emarginate. In view of the small number of species in-
volved and the variability of characters it appears that Nemopagio-
cerus Schedl represents only an unusual form of Eupagiocerus Bland-
ford and should be placed in synonymy under the latter name.

Key to the Species of Eupagiocerus

1. Sutures one and two of antennal club strongly pro-
curved, with two extending at least to middle of
club; second declivital interspace impressed or at
least less strongly elevated than either one or three
and, except at upper margin, devoid of tubercles 2

1 . Field work for this research was made possible by a grant from the National Science
Foundation, number GB-532.

2. Department of Zoology and Entomology, Brigham Young University, Provo, Utah. Scolytoidea
contribution number 27.

31

The Great Basin Naturalist

32 STEPHEN L. WOOD Vol. XXV, Nos. 1-2

Sutures one and two antennal club moderately pro-
curved, with two not extending beyond basal third
of club; declivital interspaces equally convex and
each bearing a single row of rounded tubercles 3

2. Smaller, 2.1-2.6 mm.; discal interspaces strongly convex

posteriorly, almost as high as wide at upper margin
of declivity, each elevation ending posteriorly as a
small sharply pointed spine, a row of similar pointed
tubercles continuing down each declivital inter-
spaces (except two); Costa Rica to Venezuela and
Peru ater

Larger 2.6-3.0 mm.; discal interspaces moderately
convex, not ending abruptly posteriorly; second de-
clivital interspace and perhaps lower part of third,
somewhat impressed or flat and devoid of tubercles;
declivital tubercles small, rounded; Panama clarus

3. Elytral striae narrowly impressed, the punctures ob-

scure; interspaces smooth and shining; declivital
granuals more sparsely placed, the vestiture fine,
hairlike; pronotal punctures smaller and more close-
ly placed; type 2.8 mm.; Guatemala dentipes

Elytral striae feebly impressed on disc, rather strongly
at base of declivity, the punctures almost obliterat-
ed; interspaces finely reticulate, dull; declivital
granules more numerous, the vestiture stout, almost
scalelike; pronotal punctures larger, close, but less
crowded; 3.0-3.8 mm.; Costa Rica vastus

Eupagiocerus ater Eggers

Eupagiocerus ater Eggers, 1931, Ent. Rlatt. 27:14 (Venezuela).
Epagiocerus nevermanni Schedl, 1952, Dusenia 3:350 (Costa Rica)

(new synonymy) .
Eupagiocerus serratus Wood, 1961, Gt. Basin Nat. 21:104 (Panama)

(new synonymy).
Nemopagiocerus nevermanni, Schedl, 1962. Mitt Miinchner Ent.

Gesellsch. 52:85.

The type of ater Eggers (in the Berlin Zoological Museum) was
compared by Dr. K. E. Schedl to nevermanni Schedl (type?) and to
one of my specimens collected Dec. 22, 1963 at Fort Clayton, Pan-
ama Canal Zone. According to Schedl (letter dated May 10. 1965)
minor differences are apparent in the three specimens, but only
one species is represented. The Panama specimen also agrees in all
essential characters to the paratype of serratus Wood.

A series of this species was collected with the above mentioned
PcMo^ma specimen from a common vine of the genus Serjania. From

July 8, 1965 genus eupagiocerus blandford 33

the entrance hole in a stem 2 cm. in diameter, the tunnels extended
directly to the central axis of pith, then extended about one to three
centimeters in either direction. The older mines contained larvae in
the parental gallery. On the walls of the tunnel grew a thick mat of
white cottony fungus mycelium upon which the larvae fed. This
fungus could not be seen in galleries that contained only eggs. Addi-
tional larval excavations were not apparent.

Eupagiocerus clarus, n. sp.

This species is allied to ater Eggers, but is readily distinguished
by the larger size, by the more slender body form, by characters of
the elytral declivity indicated in the above key, and by other fea-
tures of the frons and prothorax.

Female. — Length 3.0 mm. (paratypes 2.6-3.0 mm.); 2.3 times
as long as wide; body color black.

Frons broadly, evenly convex above and broadly, shallowly con-
cave below level of eye emarginations, the abrupt line of separation
extended forward slightly in median area; upper area reticulate, dull
below, subshining above; lower areas shallowly, evenly concave be-
tween subacute lateral margins to epistomal margin; epistomal mar-
gin slightly raised, shining, with a small bilobed process; concave
area rather densely, minutely pilose; other vestiture inconspicuous.
Eye 2.6 times as long as wide, shallowly emarginate on upper half,
depth of deepest point of emargination about equal to diameters of
two facets. Antennal scape bearing several coarse setae, evidently
shorter than the 7-segmented funicle; club 1.8 times as long as wide,
the sutures strongly procurved, one extending to a point one-third
length of club from base, two extending three-fifths of club length
from base.

Pronotum 1.1 times as wide as long, sides almost straight and
parallel on basal half then abruptly narrowed to the broadly rounded
anterior margin; surface closely, finely, longitudinally strigose over
entire dorsal surface; subshining; glabrous. Hairlike setae in anterior
pleural area longer and conspicuous, but not forming a definite tuft
as in ater.

Elytra 1.4 times as wide as long, sides feebly curved, somewhat
narrowly rounded behind; basal margins slightly raised, neither
crenulate nor acutely produced; striae narrowly, rather deeply im-
pressed, the punctures very obscurely indicated; interstriae feebly
convex anteriorly, moderately so at base of declivity, about three
times as wide as striae, the surface reticulate, subshining and with
numerous confused, small, shallow punctures. Declivity moderately
steep, somewhat flattened between third interspaces; interspace one
slightly elevated, uniseriately punctured, with a few punctures bear-
ing small rounded granules, two flat, appearing impressed, with
confused fine punctures, three and four convex on upper half and
bearing regular rounded tubercles on the convex portion, becoming
flattened on lower half and merging into impression of two, five to

The Great Basin Naturalist

34 STEPHEN L. WOOD Vol. XXV. Nos. 1-2

nine convex and each bearing a row of rounded tubercles. Vestiture
scanty, restricted to a few hairlike setae in posterolateral area.

Male. — Similar to female except frons with transition from
convex to concave areas gradual, the concaveity slightly deeper, not
pilose, but with a continuous tuftlike row of erect plumose setae on
lateral and lower margins, the concave area with rather numerous
small hairlike setae.

Type Locality. — Rio Viejo, Volcan Chiriqui, Panama.

Type Material. — The female holotype, male allotype and 32
paratypes were collected at the type locality on January 11, 1964, at
an elevation of 5500 ft., by S. L. Wood. They were taken from the
central axis of a rough-barked vine in stems 2 cm. in diameter or
less. The tunnels were similar to those of vastus; there was no evi-
dence of an ambrosial fungus.

The holotype, allotype and some paratypes are in my collection;
other paratypes are in the U. S. National Museum and in the British
Museum (Natural History).

Eupagioceurs vastus, n. sp.

This species is closely allied to dentipes Blandford, but is readily
separated by characters summarized in the above key.

Fem.'vle. — Length 5.7 mm. (paratypes 3.0-3.8); 2.1 times as long
as wide; color rather dark brown.

Frons flatteened on lower half, feebly convex above, with median
third remarkably sculptured; lower half with lateral margins acute
and slightly elevated, a narrow marginal and epistomal area irregu-
larly punctured, this area marked dorsally by a transversely elevated
low median carina on median third; remaining area of lower half
densely pilose; pilose area terminated above by a deep, narrow ven-
trally procurved groove on slightly more than median third, above
the groove a meuian subhexagonal impunctate area densely very
closely reticulate-granulate, a shallow transverse groove near lower
margin of hexagonal area; lateral pubescence and median hexa-
gonal area extended to upper level of eyes; dorsal areas head subre-
ticulate and finely shallowly punctured. Eye elongate, shallowly
sinuate on inner margin. Antennal scape elongate, about as long as 7-
segmented funicle; club narrowly ovate, 1.4 times longer than funicle
with two sutures, the first reaching one-fifth of length from base,
broadly procurved, the second narrowly, subagnulately procurved
reaching two-fifths of length of club from base.

Pronotum very slightly wider than long, base bisinuate, sides on
basal half straight deverging anteriorly then narrowed abruptly to a
constriction just in front of anterior margin, anterior margin very
broadly rounded, submarginate at middle; surface dull, rather closely
marked by moderately large elongate punctures, each about twice as
long as wide, the punctures much smaller anteriorly and laterally;
glabrous. Lateral margins acute on basal two-thirds, pleural areas
rather coarsely punctured.

July 8, 1965 genus eupagiocerus blanuford 35

Elytra 1.3 times as long as wide, about 1.3 times as long as prono-
tum; sides almost straight and parallel on basal half, rather narrowly
rounded behind; basal margins bisinuate, slightly elevated, not at all
crenulate; scutellum small, round; striae very feebly impressed on
disc, somewhat more strongly toward declivity, the punctures, indi-
cated but very feebly impressed; interstriae more than twice as wide
as striae, almost flat anteriorly, weakly convex at base of declivity,
surface dull, densely, microscopically punctured (visible at a magni-
fication of 80 diameters) and also finely closely, confusedly punc-
tured, with about 4 to 6 punctures across the width of an interspace.
Declivity moderately steep, convex, the surface (including strial area)
closely reticulate-granulate, each interspace bearing a median row of
small but high, rounded granules, each granule as high as wide and
spaced by distances about equal to distance between rows of granules.
Vestiture, consisting of rows of erect bristles, each arising from base of
a granule, and forming a median row on each interspace, each bristle
almost as long as distance between rows; and irregular rows of short-
er bristles about half as long as median ones, located on each side of
median row. Ninth interspace subacutely elevated,

Male. — Frons subconcavely impressed on lower half and densely
pilose in impressed area, the special sculpturing absent, otherwise
similar to female.

Type Locality. — Puerto Viejo. Heredia Province, Costa Rica.

Type Material. — The female holotype, male allotype and 51
paratypes were collected at the type locality on March 12, 1964,
from an unknown woody vine 4 to 5 cm. in diameter, by S. L. Wood.
From the entrance tunnel the adult beetles constructed linear gal-
leries along the central axis of pith about 2 to 4 cm. The larvae and
young adults then extended these somewhat. There was no evidence
of fungal growth in the tunnels.

The holotype, allotype and some paratypes are in my collection;
other paratypes are in the U. S. National Museum and in the British
Museum (Natural History).

NOTE OF PHALANGIDS AT THE NEVADA TEST SITE'

Dorald M. Allred=

This is another of a series of faunal reports dealing with the
ecology of the Nevada Test Site. A description of the biotic com-
munities of the test site was given by AUred, Beck, and Jorgensen
(1963), and subsequent publications have dealt with specific animal
groups (refer to Beck, et al., 1964, p. 209). The specimens for which
the data are reported here were identified by Willis J. Gertsch,
American Museum of Natural History, New York. Dr. Gertsch
also supplied information on the general distribution of the species.

The Phalangida include the familiar long-legged, spider-like
"daddy-long-legs" or "harvestmen" that occur in both temperate and
tropical climates. Although a number of species are common in most
parts of the United States, relatively little is known of their biology.

Two species are known from the Nevada Test Site. Eurybunus

1000 _

800 .

6 00 _

400 _

200 _

:!vXv!vX

'////////•

0

vXvX'X*

+ =2 SPECIMENS ONLY

T-KO CO GR-LY LA-FR LY MIXED

Fig. 1. Relative abundance of Eurybunus riversi in nine plant communities
at the Nevada Test Site. (AR = Artemisia tridentdta, AT-KO = Atriplex cori-
fertifolia — Kochia americana, CO = Coleogyne ramosissima, GR-LY = Grayia
spinosa — Lycium andersonii. LA-FR = Larrea divaricata — Franseria dumosa,
LY = Lycium pallidum, MIXED = other areas of a diversity which does not
permit assignment to a specific community, PLJU = Pinus monophylla —
Juniperus osteosperma, SA = Salsola kali).

I. BYU-AEC Report No. COO-1 355-9. Field work completed under AEC Contract No. AT(ll-l)

786.

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

37

38

DORALD M. ALLRED

The Great Basin Naturalist

Vol. XXV, Nos. 1-2

riversi Goodnight and Goodnight was described from specimens tak-
en near Reno, Nevada, and occurs in the Mojave Desert of Cali-
fornia. At the test site, specimens of this species are widely distribut-
ed geographically, found in 10 of the 25 areas studied: 1, 4, 5, 6, 10,
12. C, E, J, and T. 1 hey are present in all of the plant communities,
although they occur only rarely in the Atriplex-Kochia, Lycium.
and Salsola communities (Fig. 1). Their seasonal activity is pre-
dominantly during the winter months, and only a few were taken
during the summer (Fig. 2). At least half of the specimens collected
were immature.

Leiobunum townsendi Weed is a common species in the western
United States, but at the test site only four specimens were found.
These were taken in a Pinyon-Juniper community during July and
August. This species likely is more typical of the higher and more
northern communities than of the Lower Sonoran and Mojave areas.

200 _

150 _

I00_

50 _

0 J

AG

JA

MR

AP MY JN JL

^ Fig. 2. Seasonal activity of Eurybunus riversi based on numbers of individ-
uals taken at the Nevada Test Site.

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., D. M. Allred, J. R. Murdock, C. D. Jorgensen, C. L. Hayward,

AND W. W. Tanner. 1964. Nevada Test Site Desert Ecology. Proc. Utah

Acad. Sci., Arts, and Letters, 41 (2) :202-210.

A NEW GENUS AND SPECIES IN THE
CHILOPOD FAMILY TAMPIYIDAE'

Ralph V. Chamberlin"

The large family Sogonidae and the family Tampiyidae, both of
which were first proposed in 1912, pertain to a larger and apparent-
ly homogeneous group of geophilid chilopods forming a characteristic
element in the fauna of the northern tropical and subtropical areas
of America, but so far unknown from the eastern hemisphere. Repre-
sentatives of both of the two families mentioned occur in the south-
ern and southwestern United States where their progenitors must
have arrived by incursion from the south. As at present known the
most readily recognizable characteristic of these chilopods is the
possession of a labrum composed of a single piece which is always
partially and sometimes completely fused with the clypeus with no
indication of sutures setting off the divisions usual in most geophilids.
In connection with the labrum there is often developed a more or
less pronounced extension of the clypeus caudad in a lobe or process
that involves the median section of the labrum which may show
degeneration or be entirely replaced. Because of the very small size
of most sogonids and the obscure habits of the members of the group
as a whole, these animals have been generally overlooked or neglect-
ed by most collectors. As a result there undoubtedly remain many
new forms to be discovered and new groupings to be defined.

The tampiyids in their superficial appearance contrast with the
sogonids in their much larger size and more numerous segments.
Of the four genera presently referred to the Tampiyidae two were
brought to light in the study of a collection of chilopods from the
Nevada Test Site kindly referred to me for identification by Drs.
Dorald M. Allred and D Elden Beck, project directors of the Brigham
Young University ecology project at the Site. One of these two
genera is here first diagnosed; the other one, Eremorus, typified by
E. becki, was previously described (Proc. Biol. Soc. Wash., 76:33-36).
The followdng key will aid in distinguishing the four known genera.

Key to the Genera of Tampiyidae

1. Prosternum armed anteriorly with two stout teeth; last
sternite narrow; pleurocoxal pores small, independent,
and numerous Tampiya Chamb.

— Prosternum unarmed; last sternite very broad; pleuro-
coxal glands composite, opening through one or two
large pores on each side 2

1. AECBYU Report COO-I 355-10. Field work and research condmted under AEC Contracts
AT( 11-1)780 and AT( 11-1 J 1355 with Brigham Young University, Provo, Utah.

2. Institute of Arachnology, University of Utah, Salt Lake City.

39

The Great Basin Naturalist
40 RALPH V. CHAMBERLIN Vol. XXV, NoS. 1-2

2. Two composite glands and two pores on each side
Abatorus gen. n.

— One composite gland on each side, with a single pore 3

3. Median section of labrum presenting three separate den-
tate blocks Eremorus Chamb.

— Median section of labrum bearing teeth in a single un-
broken series Ketampa Chamb.

Abatorus, new genus

Head small, but little longer than broad. Antennae long and fili-
form, only narrowly separated at base.

Labrum fused at middle with the clypeus, but free laterally;
median section with posterior margin convex and bearing a series
of teeth; the lateral pieces not dentate or fimbriate.

First maxillae with coxae forming a coxosternum; palpus typical-
ly thick, distally rounded and setose. Second maxillae with coxae
united at middle by a membrane; palpus ending in a stout claw and
bearing coarse setae.

Prehensors when closed not surpassing the anterior margin of
head, all joints unarmed. Prosternum with pleurosternal suture long,
extending to antero-ectal corner on each side; the sclerotic line or
raphe strongly developed and complete; anterior margin smooth,
unarmed.

Ventral pores present in a transverse band.

Last sternite very broad, trapeziform. The pleurocoxal glands
composite, two glands and pores on each side, concealed beneath
border of the sternite.

No terminal pores.

In the male the legs are long and crassate excepting the tarsal
articles; tarsus biarticulate, bearing a stout unguiform pretarsus.
Type species: Abatorus allredi, new species.

Abatorus all red i, new species

Head small, slightly longer than broad (cir. 10:9), of form
shown in Fig. 1, with no frontal suture. Antennae long and slender-
1} narrowing distad; the terminal article with a subelliptic planate
or slightly depressed area nearly as long as the article; this area
clothed densely with short sensory hairs; similar but smaller areas on
the two preceding articles which taken together somewhat exceed the
ultimate in length.

Labrum with median section moderately convex and bearing a
comb of narrowly acute teeth as shown in Fig. 3; lateral divisions
with caudal margin a little irregularly roughened or crenulate.

First maxillae with coxae fused into a coxosternum; inner pro-
cesses conical; palpus with second or terminal article rounded, bear-

July 8, 1965 chilopod family tampiyidae

41

ing numerous coarse setae. Second maxillae with coxae united at
nuddle; palpus bearing at end a straight claw and many coarse
setae.

Prehensors short, the claws when closed not surpassing the an-
terior margin of head; all joints unarmed. Prosternum unarmed; the
pleurosternal suture long and extending forward to the antero-ectal
corner; sclerotic line strongly developed and essentially complete,
ending distally a litte ectad of the condyle.

Ventral pores in a transverse band in front of the caudal margin
of the sternite.

Last sternite very broad, trapeziform. Pleurocoxal glands com-

Abatorus allredi, new species. Fig. 1, head in outline. Fig. 2, terminal articles
of antennae. Fig. 3, labrum. Fig. 4, caudal end of body, ventral view.

The Great Basin Naturalist
42 RALPH V. CHAMBERLIN Vol. XXV, NoS. 1-2

posite, covered by border of sternite as shown in Fig. 4. No terminal
pores. Gonopods of male biarticulate. Anal legs of male of form
shown in Fig. 4. Pairs of legs 67 or more.

Length: 40-46 mm.

Localities: Nevada: Nye Co., Nevada Test Site. One male
taken Dec. 24, 1961. California: Riverside, Box Springs. Several
specimens taken by Dr. Joseph C. Chamberlin, Dec. 1, 1925.

TWO ISOPODS OF THE NEVADA TEST SITE'

Dorald M. Allied', and Stanley Mulaik'

During ecological studies over a four-year period at the Nevada
Test Site (Allred, Beck, and Jorgensen, 1963a), 490 isopods of only
two species were collected. These were determined by Dr. Stanley
Mulaik as representing 381 specimens of Armadillo arizonicus
(Mulaik) and 109 specimens of Porcellio laevis Latrielle. They were
collected principally in sunken can traps (described by Allred, Beck
and Jorgensen, 1963a) which were operated continuously for at
least one year in nine areas representing seven major plant com-
munities. In 19 other areas of the site, cans were operated only for
a two-week period during a summer. This resulted in a total of
51,992 can trap-nights over the four-year period.

Armadillo arizonicus

(Tables 1 and 2, Fig. 1 )

Isopods of this species were found in seven vegetation types as
shown in Table 2. They were most commonly associated with
Lycium pallidum and least with Atriplex conjertijolia-Kochia ameri-
cana. Within some plant communities the concentration of animals
was somewhat localized. In the Mixed (Xeric) community the two
collecting transects were only 825 feet apart, yet there were more
than five times as many isopods taken from one transect than from
the other. This same situation also occurred in the Larrea-Franseria,
Lycium, and Grayia-Lycium communities. In the latter study area
four collecting transects, each 1.5 miles long, ran NE, SE. SW and
NW, respectively, from a central point (refer to Allred, et al., 1963a,
Fig. 12). Over 88 percent of the isopods taken were found on the
NE transect, and 70 percent of the animals on this transect were
taken at one collecting station. A similar situation was found in the
Lycium study where more than 70 percent of the isopods from one
transect were found at two adjacent stations. On the other transect
over 40 percent of the animals were taken at one station.

In the total collection of adult isopods the sex ratio was 1:1, but
this varied between different plant communities. In the Mixed
(Xeric) community the ratio of males to females was 2:1, in the
Lycium 3:4. in the Grayia-Lycium 1:2, with slight differences in
the other communities.

Seasonally, A. arizonicus was active only from May through
October (Fig. 1 ). Highest populations occurred during August for the
adults and July through September for immatures. Seasonal appear-
ance of males and females was similar in all communities except in

1. Atomic F.iieiRV Cnniiiiission Report No. COOl 355-4, Field work completed under AEC Con-
tract .Vl'l 1 1 -1 )7H(> wltii Brigham Young University.

2. Deparlint'iU iif Zoology and Entomology, Brigham 'S'oung University.
i. Department of '/oology, University of Utah.

43

The Great Basin Naturalist
44 ALLRED AND MULAIK Vol. XXV, NoS. 1-2

Table 1. Collection localities* of isopods at the Nevada Test
Site, Nye County, Nevada.

Armadillo arizonicus

8 d" 15 9 31 im., Study IB, Ground zero 1 to radius of 1.5 mi.
7 cT 11 9 9 im.. Study 5A, 0.2 mi. E. of Mercury Highway

S of Well 5B road.
5 cT 9 9 1 im.. Study 5CQ, 0.3 mi. E. of Mercury Highway,

N of Well 5B road.
34 cf 43 9 90 im.. Study 5E, 1.1 mi. E. of Mercury Highway,

thence 1 mi. S of Well 5B road.

1 cT, Study 6A, 0.5 mi. S of Well 3B, thence 0.6 mi. E.

2 cT 6 9 1 im., Study lOD, 9.5 mi. N. of Well 3B along

Groom Lake road, thence 0.5 mi. E.
16 cf 14 9 1 im., Study CB, Environs of Cane Springs.
50 d' 25 9 2 im., Study JA, 9.3 mi. W. of Mercury along

Jackass Flats Highway, thence 1000 ft. SW.

Porcellio laevis

1 im., Study lOB, 9 mi. N. of Well 3B along Groom Lake

road, thence 0.4 mi. W.
36 cf 58 9 16 im., Study CB, Environs of Cane Springs.

'I'"or specific location refer to Allred, Be<k, and Jorgensen, l')()5b.

Larrea-Franseria where the males did not appear in the collections
until August.

Porcellio laevis
(Table 1, Fig. 2)

All but one of the 109 specimens were taken at Cane Springs in
a Mixed (Mesic) plant community. The one exception was taken in
a Coleogyne area. Over 90 percent of those at Cane Springs were
found at the three adjacent stations closest to a pond.

The ratio of males to females was 1:1.6. Seasonally, highest
populations of adult males appeared during August through Decem-
ber, and females during August through October (Fig. 2).

Discussion

It is unusual that so few species of isopods are represented at
the test site. It is urdikely that other species, if present, are widely
distributed. However, if the isopods tend towara localized and sea-
sonal populations as demonstrated by the two species collected, then
other species may exist at the test site in specific areas where we
have not done extensive collecting on a year-round basis. Some iso-
pods do not roam about as do P. laevis and A. arizonicus. Others
climb onto plants and may be obtained by sweeping. However, our
extensive sweeping of plants for other arthropods has still not yielded
isopods other tnan the two species taken. Furthermore, we collected
the entire plant and ground debris of eleven species monthly over a

July 8, 1965
50 -.

ISOPODS OF NEVADA TEST SITE

45

40 -

30 -

20 -

0 -

/v

/\

\k'

r I

MALES
FEMALES

IMMATURES

Y „ .—

1 1 1 1 r

APR MAY JUN JUL AUG SEP OCT NOV

Fig. 1. Seasonal abundance of Armadillo arizonicus.

year's period (Allred and Beck, 1964), but not a single isopod was
found. In addition to the buried can and other methods of collecting,
considerable searching was made for isopods under rocks, fallen
trees, and in plant humus. Nevertheless, our relatively limited col-
lecting over the 1,000 square-mile area of the test site and our some-
what selective methods of sampling leave some question as to the
complete isopod fauna of the site. It is evident that more specific
investigations on the ecology of isopods in desert areas need to be
made.

The occurrence of isopods around moist situations such as P.
laevis at Cane Springs is not surprising. However, large populations
of those such as A. arizonicus in such xeric habitats are not usually

46

ALLRED AND MULAIK

The Great Basin Naturalist

Vol. XXV, Nos. 1-2

Table 2. Relative abundance of Armadillo arizonicus in several
plant communities at the Nevada Test Site.

« s

1 Li
(8 <D

S

o

■>.a

ft) m

3

(0 u

U (0

o

o

t >■•

« £

>^

u

OJ

Jta

h-1

l1

Is

-s-

IX

Actual number
collected

1

Relative abundance
adjusted by no. of
collecting attempts 2

20

53

53

42

78

168

381

31

53

77

169

expected. The differences in relative abundance of A. arizonicus in
different plant communities are difficult to explain on the basis of
either plant-food relationships or edaphic factors. Little is known of
the feeaing preferences of this species of isopod relative to organic
debris. Rockiness of the soil and moisture conditions vary considera-
bly between some plant communities at the test site. On the high
mesas there are rocky situations and generally a greater degree of
moisture which are more conducive to optimum conditions than in
the drier valleys. Yet not a single isopod of either species was taken
in our mesa studies. In the Lycium area where A. arizonicus oc-
curred in greatest abundance there are essentially no surface rocks
of a size that would provide suitable habitats for the isopods, and the
soil is loose sandy-clay. This area is adjacent to the Frenchman Flat
playa which is dry for most of the season. Other than this area A.
arizonicus was found most abundantly in a Mixed community where
the condition is also xeric but large rocks are abundant. In both
plant communities the total plant cover is considerably less than in

15 _

10 _

Fig. 2. Seasonal abundance of Porcellio laevis.

July 8, 1965 isopods of Nevada test site 47

the Atriplex-Kochia and Coleogyne areas where populations of A.
arizonicus were lowest. Except for the Grayia-Lycium site each of
our study areas was established in areas where the vegetation was
considered to be rather uniform. This was confirmed by vegetation
analyses made by the line-intercept method.

Apparently there is no plant association which influences the
occurrence and abundance of A. arizonicus. As far as is known, the
edaphic factors within each of the study areas are generally uniform
throughout. Further detailed studies are needed, however, to deter-
mine why populations of A. arizonicus are localized and more abun-
dant around one small area than around another which is apparently
similar.

The almost exclusive occurrence of P. laevis near a constant
water source indicates that it requires a more mesic habitat than
does A. arizonicus. It is likely that the Cane Springs habitat supplies
more decaying, moldy food than is to be found in drier areas on the
test site.

The seasonal activity of A. arizonicus occurred between May and
October when temperatures are highest and rainfall lowest at the
test site. On the other hand, P. laevis was active every month of the
year in an area where moisture was available the year round, but
where temperatures varied little from those of the more xeric condi-
tions of the habitats of A. arizonicus.

Literature Cited

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

of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser., 11(2):

1-52.
. 1963b. Nevada Test Site Study Areas and Specimen Depositories.

Brigham Young Univ. Sci. Bull., Biol. Ser., 11(4): 1-15.
Allred, D. M. and D E. Beck. 1964. Arthropod Associates of Plants at the

Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser., V(2):l-16.

PRAIRIE FALCON IMITATES FLIGHT PATTERN
OF THE LOGGERHEAD SHRIKE

White (Condor, 64, 1962:439-440) stated, "The normal mode of hunting for
the prairie falcon is the expected falcon method of a swift direct flight about 30
to 200 feet in the air with a long, low angle stoop at the expected prey." On
December 22, 1962, while attempting to trap prairie falcons at the Nevada Atomic
Test Site (United States Atomic Energy Commission Contract AT(1 1-1 )786), an
unusual hunting method used by two prairie falcons was observed. Instead of
hunting in the usual fashion, they left their perches in an undulating flight pattern
similar to that of the loggerhead shrike. The hoax was executed so skillfully that
the falcons were able to approach within 30 feet of their unsuspecting prey
(White-tailed Antelope Squirrels). When the squirrels became alarmed and ran
for cover, the falcons promptly returned to their usual onrushing attack to close
the remaining distance. No squirrels were captured in this manner, even though
six different flights were observed.

Another instance where a falconiform used the flight pattern of an unrelated
species was reported by Mavrogordata (A Hawk for the Bush, 1960:4). He ob-
served a European sparrow hawk using the characteristic wing beat of a lapwing
to conceal its approach on some feeding starlings. In this instance the behavior was
similar in that when the prey became alai-med the predator assumed its usual
flight.— Gerald L. Richards, 1939 North 450 West, Provo, Utah.

48

Great Basin

Volume XXV December 31, 1965 Nos. 3-4MNlV£R^> ^^

TABLE OF CONTENTS

Undescribed Species of Nearctic Tipulidae (Diptera) VI.

Charles P. Alexander 49

Two New Mites from the United States (Acari: Oribatei,

Microzetidae, and Oribatellidae) . Illustrated. Harold G.

Higgins 55

A New Genus and Species of Oribatid Mite from Colorado

(Acari: Orbatei, Ceratoppiidae) . Illustrated. Tyler A.

Woolley and Harold G. Higgins 59

Studies in Nearctic Desert Sand Dune Orthoptera, A new

Genus and Species of Stenopelmatine Crickets from the

Kelso Dunes with notes on its multi-annual life history

and key. Illustrated. Part X. Ernest R. Tinkham 63

Records of Atherinid Fishes at Inland Localities in Texas

and Northern Mexico. W. L. Minckley. Illustrated 73

A List of Scarabaeidae Beetles of the Nevada Test Site.

Dorald M. Allred and D Elden Beck 77

Angus Munn Woodbury, 1886-1964. Vasco M. Tanner. Illus-
trated 81

Index 89

Published by
Brigham Young University

The Great Basin Naturalist

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

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

Illustrations: All illustrations shoLdd be made with a view to
having then appear within the limits of the printed page. The ill-
ustrations that form a part of an article should accompany the
manuscript. All half-tones or zinc etchings to appear in this jour-
nal are to be made under the supervision of the editor, and the cost
of the cuts is to be borne by the contributor.

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

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

All correspondence dealing with manuscripts, subscriptions,
reprints and other business matters should be addressed to the Editor,
Vasco M. Tanner, Great Basin Naturalist, Brigham Young Univer-
sity, Provo, Utah.

Reprints Schedule of The Great Basin Naturalist

Each
Additional

2 pp.

4 pp.

6 pp.

8 pp. 10 pp. 12 pp.

2 pp.

50

copies

$6.00

$7.00

$8.00

$9.00 $10.00 $11.00

$2.00

100

copies

7.00

8.00

9.00

10.00 11.00 12.00

200

copies

8.00

9.00

10.00

11.00 12.00 13.00

300

copies

9.00

10.00

11.00

12.00 13.00 14.00

C>)vers: $10.00 for first 100 copies, $4.00 for additional 100 copies.

-^ ^ ' ^^'0 AUG 18 1966

The Great Basin NaturW^^'^^

Published at Pkovo, Utah by
Brigham Young University

Volume XXV December 31, 1965 Nos. 3-4

UNDESCRIBED SPECIES OE NEARCTIC TIPIUJOAE
(DIPTERA) VI.

Charles P. Alexaiidcr'

In the present paper I am describing three species of the genus
lipula Linnaeus and a single species of Limonia Meigen. All types
of the novelties are preserved in my personal collection through the
kindness of the individual collectors of the materials. Mr. James H.
Baker of Baker, Oregon, Mr. W. ¥.. Simonds of Sacramento, Cali-
fornia, and the late Robert R. Dreisbach of Midland, Michigan.

Tipula {Nippotipula) metacomet, n.sp.
The present fly has been confused with lipula (Nippotipula)
ahdominalis (Say) which it closely resembles but from hypopygial
structure evidently is distinct. In size somewhat smaller than ahdomi-
nalis (wing 23 mm.). Legs uniformly brownish black with no indi-
cation of pale rings on the femora or bases of tibiae. Male hypopy-
gium with both dististyles black, outer style with the basal part be-
fore the oblique thickened ridge short, the outer half or more slightly
angulated and produced into a rectangular blade, its apex truncate,
with short spinoid setae; inner style much smaller, the base enlarged,
outer end suddenly narrowed into a slender blackened club, the sirn-
ple apex without major setae. In abdomin'alis the outer dististyle is
roughly oval in outline, the oblique ridge beyond midlength, re-
stricting the outer part to a smaller semioval lobe with abundant
spinoid setae; inner style relatively large, the pale apex dilated,
more or less bilobed. with several strong setae, at base of style with
a pale blade or flange.

I1.A.BITAT. — Massachusetts (Hampshire County).

HoLOTYPE, d", Amherst, August 15, 1945 (Charles P. Alex-
ander) ; type mounted on two microscope slides.

Named for Metacomet, more commonly known as King Philip,
second son of Massasoit of the Wamponoag tribe of New England
Amerinds. King Philip's War (1675-1676) with the English colon-
ists was waged in the vicinity of the type locality.

1. Anilicist. Miiss;i(liiisolls.

49

The Great Basin Naturalist
30 CHARLES P. ALEXANDER Vol. XXV, NoS. 3-4

Tipula (Lunatipula) mercedensis, n.sp.
Allied to degeneri; male hypopygium with posterior border of
tergite narrowly emarginate; inner dististyle with the beak bilobed,
dorsal crest narrow, blackened, with long setae; outer basal lobe a
powerful curved arm that narrows into a strong spine, the upper
surface of arm with exceedingly long yellow setae.

Male. — Length about 18.5 mm.; wing 17 mm.; antenna about
6.5 mm.

Frontal prolongation of head long, nearly equal to remainder of
head, yellow, narrowly dusted with light gray above, nasus long;
palpi with proximal three segments yellowed, the fourth brownish
black. Antennae elongate; scape, pedicel and most of first flagellar
segment yellow, the remainder black; flagellar segments strongly
incised, producing a conspicuous outer enlargement, subequal in size
to the basal swelling; segments slightly exceeding their longest ver-
ticils. Head brownish gray, orbits paler gray; occipital region huffy.
Pronotal scutum gray, scutellum yellow. Mesonotal praescutum
gray with four entire light brown stripes, the intermediate pair
slightly more cinnamon brown than the laterals; posterior sclerites
of notum gray, each scutal lobe with two brown areas, medioter-
gite with a capillary brown central vitta. Pleura gray, dorsopleural
membrane yellowed. Halteres with stem whitened, the base more
yellowed, knob brown. Legs with coxae light gray; trochanters yel-
low; remainder of legs yellow, the tarsi passing through brown to
black; claws of male toothed. Wings brownish yellow, prearcular and
costal fields clearer yellow, stigma a little darker, brownish yellow;
obliterative areas poorly differentiated; veins brown. Venation: Tip
of /?,+:; atrophied; petiole of cell M, slightly longer than m.

Abdominal tergites yellow with a broad blackened central stripe,
the posterior borders of the intermediate segments narrowly yel-
lowed, sublateral areas very vaguely darkened; sternites yellow;
hypopygium large, brownish yellow. Male hypopygium with pos-
terior border of tergite narrowly and deeply emarginate, fringed
with long setae, the border adjoining the emargination produced into
a flattened blade. Appendage of ninth sternite dilated and weakly
emarginate, the border with strong yellow setae, those of upper lobe
stouter. Basistyle produced virtually as in degeneri. Inner dististyle
with the beak bilobed. dorsal crest low, blackened, with long yellow
setae; outer basal lobe produced into a powerful curved arm. its tip
narrowed outw^ardly into a strong spine; upper or concave surface of
the arm with exceedingly long yellow setae from base almost to the
terminal spine; no basal dilation as in degeneri. Eighth sternite
sheathing, posterior border broadly concave on either side with an
apical lobe provided with abundant long yellow setae, the tips
curved and twisted, the shorter mesal setae forming a dense tangle.
Habitat. — California (Mariposa County).

HoLOTYPE, cT , Yosemite National Park, along Merced River, trail
to Vernal Falls, altitude about 4,200 feet. June 12, 1963 (James
Baker).

Dec. 31. 1965 new nearctic tipulidae 51

The only known relative of this fly is Tipula (Lunatipu/a)
degeneri Alexander, still known only from Sequoia National Park,
California. The present species differs conspicuously in the hypopy-
gial characters, particularly the ninth tergite and the inner dististyle,
as described. In degeneri the outer basal lobe of the inner style is
expanded at base, the outer arm slender and without setae.

Tipula (? Pterelachisus) simondsi, n.sp.

Wings reduced in size, at least in the female; general coloration
of head and thorax light gray, frontal prolongation of head elongate,
with a nasus; mesonotum restrictedly patterned with brown; halteres
chiefly pale yellow; wings about one-half the length of body, stenop-
terous. pale yellow, conspicuously patterned with dark and paler
brown, veins behind R \%athout macrotrichia; abdomen tricolored,
segments dark brown basally, light gray on more than outer half,
the posterior borders of outer segments narrowly light yellow; ovi-
positor with cerci smooth, hypovalvae elongate, with rounded apices.

Female. — Length about 23 mm.; wing 11x2 mm.; abdomen
about 18 mm.; antanna about 2.5 mm.

Frontal prolongation of head unusually long, only a little less than
remainder of head, light gray, with indications of a narrow darker
median line; nasus distinct, about twice as long as broad, with a few-
stout black setae; palpi black, sparsely pruinose. Antennae black,
pruinose; 12-segmented, the proximal two flagellar segments partially
fused, the combined length only a little less than the scape, terminal
segment reduced to a tiny oval structure; verticils a little shorter
than the segments. Head gray, the vertex with vague indications of
a capillary darkened line, more diffuse behind; vestiture of head
reduced to a few scattered microscopic black bristles.

Cervical region huffy. Pronotum gray, center of scutum elevated,
slightly more infuscated. Mesonotum gray, praescutum with indica-
tions of two darkened lines on anterior half, converging behind and
vaguely reaching the suture, evidently representing the usual inter-
spaces, the four gray stripes including two narrow intermediate
vittae and much broader lateral areas; pseudosutural foveae very
small, blackened; scutum gray, more huffy medially, each lobe with
narrow darkened lines; scutellum and postnotum light gray, vaguely
patterned with darker; mesonotum glabrous. Pleura gray, dorso-
pleural membrane abruptly light yellow. Halteres pale yellow, base
of knob weakly darkened. Legs with coxae light gray, with sparse
long white setae ; trochanters brownish gray; femora and tibiae
light yellowdsh brown, tips narrowly brownish black; basitarsi
brown, outer tarsal segments blackened. Wings slightly reduced in
size, stenopterous, to produce an evidently flightless condition; wmgs
of nearly uniform width for most of the length; ground color pale
yellow, restrictedly patterned with dark and paler brown; prearcular
and costal fields, including both cells C and Sc, clear hght yellow;
the darker brown areas include marks at arculus, origin of Rs and
the anterior cord; paler brown markings include a linear dash in cell

The Great Basin Naturalist
52 CHARLES P. ALEXANDER Vol. XXV. Nos. 3-4

R adjoining the vein, and broad areas in cell M placed at midlength
and near outer end; further linear pale brown lines at midlength and
near outer ends of cells Cu and 1st A, and almost the outer two-thirds
of 2nd A; beyond the cord. M,+.. is narrowly seamed with brown;
veins light yellow in the ground areas, pale brown in the darker
brown markings. Macrotrichia on veins C, Sc and R, lacking on all
other veins; squama naked. Venation: Rs about one-half longer than
m-cu; veins /?,+.. and R. widely divergent, cell R. at margin about
twice /?.i; petiole of cell Af, short, about one-third m; m-cu longer
than distal section of Cux; prearcular field of wing elongate.

Abdomen long, the segments conspicuously patterned, the basal
rings dark brown, the broader apical parts light gray; posterior bor-
ders of tergites five through eight and sternites six and seven narrow-
ly light yellow; ovipositor with dorsal shield and the cerci brownish
black, the ventral parts slightly paler. Ovipositor with cerci elongate,
margins smooth, apices subacute; hypovalvae about four-fifths as
long, compressed-flattened, tips obtuse.

Habitat. — California (Inyo County).

HoLOTYPE, 5 , on snow field along trail to Mount Whitney,
Sierra Nevadas (Lone Pine to Whitney), altitude 12,600 feet. July
18, 1965 (W. E. Simonds).

I am pleased to name this interesting fly for the collector. Mr.
William E. Simonds of Sacramento, California. In the absence of
the male sex I hesitate to assign it definitely to any subgenus but
am placing it with some question in Pterelachisus Rondani {Oreo-
myza Pokorny) where it appears to belong. It may possibly belong
to Serratipula Alexander which includes various subapterous species.
Other regional subgenera are excluded by different characters, as
Lindneria Mannheims by the loss of the nasus. and Lunatipula
Edwards and Triplicitipula Alexander by the presence of squamal
setae. I cannot associate the present female with any of the numer-
ous regional fully winged males. The striking tricolored pattern of
the abdomen is somewhat suggestive of Tipula {Pterelachisus) ter-
naria Loew, which is fully winged in both sexes. This is a high
northern Nearctic fly that ranges from Ontario to Quebec, west-
ward to the Yukon, southwards into the more northern New England
states. It is probable that the male sex of the present species likewise
will be found to be fully winged.

Subapterous species of Tipula are well known in California, in-
cluding several species in Serratipula and Triplicitipula. including
the female sex and in a few cases also the male. As indicated above,
the present fly has the reduced condition of the wings only moderate.
By the scale of wing atrophy given by Bezzi in an important paper
on the subject', the fly would fall between his categories 1 and 2
where the wing shape has been deformed but the venation clearly
indicated. Attention may be called to a very recent paper by the

1. Bezzi, Mario Riduzionp e sroniparse delle all iiegli iiiM'lli Dilloii. Riii^ln di S(irn:t- Siitiinili.
",\'ntura", 7:85-182, 10 figs., with numerous siibfigiiros, 11)11).

Dec. 51. 19fl5 NEW NEARCTIC TIPULIDAE 5 3

writer tliat concerns the Californian species oi Tipula and bears on
the present question; reference- below. ^

Limonia {Dicranornyia) dreisbachi, n.sp.

General coloration of head and thorax gray, the praescutum with
a dark brown central stripe on anterior half; antennae black through-
out; wings whitish hyaline, stignia pale yellow; Sc short, Rs less than
the basal section of /?4+.,. cell 1st M. closed; male hypopygium with
the rostral prolongation of the ventral dististyle long, spines sub-
equal in length, placed close together at near two-thirds the length.

Male. — Length about 6.5 mm.; wing 7 mm.

Rostrum yellow, sparsely pruinose above the base, palpi black.
Antennae black throughout; flagellar segments oval, the outer ones
slightly more elongate; verticels shorter than the segments. Head
gray; anterior vertex broad, about three times the diameter of scape.

Pronotum buffy yellow. Mesonotal praescutum dark gray, clearer
light gray on sides; a conspicuous dark brown central stripe on an-
terior half, lateral stripes not or scarcely indicated; scutum gray,
lobes darker gray; scutellum dull yellow, sparsely pruinose at base;
postnotum gray. Pleura gray, dorsal pteropleurite and meral region
buffy; dorsopleural membrane brownish yellow. Tlalteres with stem
whitened, knob brown. Legs with coxae and trochanters yellow;
remainder of legs broken. Wings whitish hyaline, prearcular field
light yellow; stigma vaguely indicated, pale yellow; veins delicate,
brown, those of proximal third of wing more yellowed. Venation:
Sc short, Sc\ ending some distance before origin of Rs, the distance
exceeding Rs, Sc. retracted, Sci alone more than three-fourths Rs;
Rs shorter than basal section of /?4+r,,- cell 1st M-, closed, nearly as
long as distal section of M,+^.,- m-cu longer than outer section of Cu^,
placed shortly beyond the fork of M.

Abdomen dark brown, hypopygium yellowed. Male hypopygium
with the tergite transverse, posterior border shallowly emarginate to
form low obtuse broadly thickened lobes, setae long, pale yellow,
inconspicuous. Basistyle with ventromesal lobe oval, with long yel-
low setae. Dorsal dististyle a gently curved rod that narrows gradual-
ly to a slender point; ventral style subequal in area to the basistyle,
body oval, with pale setae; rostral prolongation long, the apex nar-
rowed, the two spines placed close together at near two-thirds the
length, the spines subequal to the apex beyond their insertion. Gona-
pophysis with mesal-apical lobe slender, gently curved.

Habitat. — Michigan (Midland County).

HoLOTYPE, cf. Midland County, without more exact data. August
4, 1954 (R. R. Dreisbach).

The species is dedicated to the collector, the late Robert R Dreis-
bach, distinguished student of the Hymenoptera. Superficially it is
much like Limonia (Dicranornyia) hrevivena (Osten Sacken) in the
venation, as the short Sc and Rs but actually is a very different fly.

2. Alexander. Charles P. New subgenera and sperie*; nf rrane-flies from (California (Diptera:
lipulidae ) .
I'nrijir Inserts 7. no. 2:?iV38fS, ^', fig<;.: June !0(,5.

TWO NEW MITES FROM THE UNITED STATES
(ACARI: ORIBATEI, MICROZETIDAE AND ORIBATELLIDAE)'

Haiold G. Higgins-

The family Microzetidae is composed of small, "winged" mites
with large, often bizarre lamellae. Until the present time this family
has been reported only once from the United States, but has been
found in many collections from South America. Europe, Africa and
Madagascar. In a collection of oribatid mites from North Carolina
was an undescribed species of microzetid as well as a new species of
Ophidiotrichus. a genus not previously recorded from North America.
Figures and descriptions of these new species follow.

Allozetes n. gen.

Diagnosis: Lamellae long, extending over tip of rostrum, broad,
incurved lateral margins; lamellar hairs heavy, incurved, strongly
setose and extended beyond tip of lamellae by about one-half their
lengths; sensillus with brush-like tip and long pedicle; fixed digit of
chelicerae with a long, racemose, thorny spine as shown in figures
3 and 4.

Allozetes harpezus n. gen., n. sp.
(Figs. 1-4)

Diagnosis: Lamellar hairs heavy, incurved, strongly setose and
extended beyond tip of lamellae by nearly one-half their lengths;
sensillus brushlike, not proclinate.

Description: Propodosoma longer than wide; rostral hairs small,
not visible from above; lamellae covering propodosoma, with smooth,
incurved, lateral margins, anterior ends with rounded medial edges
and a short, lateral point, medial edges nearly parallel, separated
near base by a rectangular space; lamellar hairs heavy, setose, in-
serted under the lamellae and attached nearer the medial edge of
lamellae, curving laterally and then medially, extending beyond tip
of lamellae by about one-half their lengths; insertions of interlamel-
lar hairs located on each side of the rectangular space between the
lamellae, but interlamellar hairs missing in type specimen; pseudo-
stigmata cup-like, located at junction of lamellae and pteromorphs;
sensillus brush-like with a long pedicle.

Hysterosoma round in outline; sclerotized lines extending pos-
teriorly from the straight dorsosejugal suture; pteromorphs not di-
rected forword, each pteromorph with two setae on upper surface;
dorsal setae as shown in figure 1 ; hysterosoma covered with an ir-
regular, reticulate pattern.

Camerostome egg-shaped; chelicerae heavy, fixed digit with a
long, racemose, thorny spine that is about as long as the movable

I Srtll liike Citv. Utah. , <- i i i^ u

1. I'aitMipant in National Sc lenre Foiindalidn Rt-scan h pai tinpation for High School I eachers

55

56

The Great Basin Naturalist
HAROLD G. HIGGINS Vol. XXV, Nos. 3-4

Fg 3 AUn T f '''''"' " ^'"•' "• ^P- "^"^^-^^ ^^^-- legs omitted.

V 1 ^ °-^^^^ /^arp^^«. n. gen., n. sp., chelicera.

^ig. 4. Allozetes harpezus n gen., n. sp., spine on fixed digit of chelicera.

digit, as shown in figures 3 and 4; genital plate about as long as
wide with parallel sides, each plate with six setae (figure 2). ^3 even
with ^4 but located near lateral edge of cover; anal opening about as
long as wide, larger than genital aperture, each anal plate with two
setae; aggenital setae located posterolaterad of genital plate; three
pairs of adanal setae, ada-,, located at anterior level of anal plate; pore
lad located between ada. and ada,.., ventral apodemata and setae as
shown in figure 2.

Size: Length 276 ft; width 204 ix.

Type Locality: One specimen from four miles north of Chero-
kee, North Carolina, 28 May 1957 by W. Mason.

Discussion: Until now only one Microzetid has been reported
from the United States. Jacot (1938) described Microzetes auxiliaris
appalachicola from grass sod at Bent Creek Exp. Forest, North Caro-
lina. However, because of the short description and lack of a figure,
I must agree with Balogh (1962a) that it is impossible to put this
form in its proper place in the genus Microzetes.

Allozetes harpezus n. gen., n. sp. is different from all known
members of this family in the shape of the lamellae, the size and
shape of the lamellar hairs, the brush-like sensillus, and the long,
racemose, thorny spine on the fixed digit of the chelicerae. The name
harpezus means "thicket" or "thorn hedge" and refers to the un-
usual spine on the fixed digit of the chelicerae.

Doc. 31. 1965

NEW MITES FROM UNITED STATES

57

Ophidiotrichus exastus n. sp.
(Figs. 5-6)

Diagnosis: Anterior end of lamellae serrate, median dens longer
than lateral dens; rostrum pointed, entire; interlamellar hairs more
than two-thirds as long as lamellae; no distinct areae porosae.

Description: Propodosoma longer than wide; rostrum pointed,
entire, not visible from above; lamellae long, broad, finely pitted
with a serrate tip, the median dens longer than lateral dens; lamellar
hairs very heavy, broad, finely setose, located in anterior end of
lamellae; interlamellar hairs located close together, medially on pro-
dorsum near dorsosejugal suture, over two-thirds as long as lamellae,
setose, with small insertions; pseudostigmata cup-like; sensillus
weakly setose with nearly parallel sides and fine tip which extends
nearly to ends of lamellae.

Hysterosoma quite round; dorsosejugal suture straight; entire
dorsum finely pitted; setae simple and located as shown in figure 5;
muscle scars around edge of hysterosoma; pseudoporosae anterior to
setae r^; no distinct areae porosae.

Camerostome egg-shaped; genital opening wider than long and
separated from the larger anal plate by twice its length, each plate
with six setae as shown in figure 6; anal opening larger than genital,
nearly as wide as long, each plate with two setae, an, near medial
edge of plate and an-, nearer middle of plate; three pairs of adanal
setae, ada,, located at level near middle of anal plate; iad at antero-
lateral margin of anal aperture anterior to ada^., aggenital setae
postero-laterad to genital plate; apodemata and setae as shown in
figure 6.

Size: Length 247 jx; width 180 p..

Type Locality: A single specimen from four miles north of
Cherokee, North Carolina, 29 May 1957 by W. Mason.

Fig. 5. Ophidiotrichus exastus n. sp. dorsal view, legs omitted.
Fig. 6. Ophidiotrichus exastus n. sp. ventral view, dorsal plate removed.

The Great Basin Naturalist
58 HAROLD G. HIGGINS Vol. XXV, Nos. 3-4

Discussion: Although the genus Ophidiotrichus (family Oribatel-
lidae) has been previously reported from Europe, this is the first
record from North America. This species resembles somewhat Ophi-
diotrichus connexus vindobondensis Piffl in having a pitted body and
notched lamellae, but differs from all known forms in having a ser-
rate-tipped lamella with long median dens; longer, heavier, inter-
lamellar hairs more medially located; rostrum entire, pointed; and
in the lack of distinct areae porosae. In addition there is a difference
in the location of the genital setae. The name exastus means "rough
edge" and refers to the anterior edge of the lamellae.

References

Balogh, J. 1962. New Microzetids from Eastern Peru (Acari. Oribatei). Ann.

Nat. Mus. Nat. Hung., 54:405-417.
. 1962a. An Outline of the Family Microzetidae Grandjean. 1936.

(Acari: Oribatei). Opusc. Zool. Budapest. 4(2-4) : 35-58.

1965. A Synopsis of the World Oribatid (Acari) Genera. Acta Zool.

Hung.. ll(l-2):5-99.
Berlese, a. 1904. Acari nuvoi. Manipulus III. Redia, 2:10-32.
Grandjean, F. 1936. Microzetes auxiliaris n. sp. (Oribates). Bull, de Mus.

France, 8(2) : 138-145.
. 1936a. Les Microzetidae n. fam. (Oribates) Bull. Soc. Zool. France.

61 (2): 60-93.

1953. Observations sur les Oribates (27*" serie). Bull. Mus. Paris. 2''

serie, 25(5) :469-476.
Jacot, a. p. 1938. Some new Western North Carolina moss mites. Proc Ent.

Soc. Wash., 40:10-15.
PiFFL, E. 1960. Zur Oribatidenfauna des Leopoldsberges von Wien (1. Beitrag).

Ann. Nat. Hist. Mus. Wien, 64:165-172.
WooLLEY, T. A. 1958. Redescription of Ewing's Oribatid Mites. VH. Trans.

Amer. Micr. Soc. 77(2) : 1 35-146.

A NEW GENUS AND SPECIES OF ORIBA 1 II) MITE

FROM COLORADO

(ACARI: ORIBATEI, CERAIOPPIIDAE)'

Tyler A. Woollev- and Harold G. Higgins'

Observations of collections of some montane oribatid mites dis-
closed what appeared to be a new genus and species related to Cera-
toppia bipilis (Herm.) and Pyroppia lanceolata Hammer, yet differ-
ent from either. Additional study and morphological comparisons of
the new species with descriptions of Pyroppia and available speci-
mens of Ceratoppia disclosed some marked differences in the lamel-
lae, chelicerae and other structures. These differences indicated con-
clusively that the mite was distinct and new, so other diagnostic
features were studied.

Although some similarities exist among these genera, several dif-
ferences were observed, most striking of which was the discovery of
pelopiform chelicerae, with a swollen base, narrow shaft, and small
dentate shears. It was this discovery which prompted further inves-
tigation and comparison of the lamellae, infracapitulum and legs of
the new species and those of established genera.

Because of these comparisons other details were disclosed that
extend beyond the scope of this paper. It was apparent that further
study would be necessary to determine final relationships and de-
limitations of these species. Research on this comparative morphology
has begun to assess the generic and specific features of the ceratop-
piids and liacarids. A few pertinent facts will be delineated below,
but disclosure of many others will await completion of the study
currently being conducted.

Paenoppia n. gen.
With sclerotized, tuberculated prodorsum and lamellae; lamellar
cusps smooth, not tuberculated, finger-like, extended above prodor-
sum; chelicerae pelopiform; sensillus setiform, with small barbules
along nearly parallel sides except for filiform tip, slightly swollen at
beginning of terminal flagellum. Differs from Pyroppia in the wider,
tuberculate lamellae; the setiform sensillus compared to the lanceo-
late sensillus of Pyroppia; the rounded body shape compared to the
pear-shaped body of Pyroppia. The generic name is compounded to
indicate a postulated relationship with other ceratoppiids.

Paenoppia forficula, n. sp.
(Figs. 1-7)
Diagnosis: Pelopiform chelicerae; lamellae broader than lamellar
cusps, and each with a longitudinal dorsal ridge extending from base

1 Research sponsored by National Science Foundation grants G-I4333 and GB 3872.
2. Department of Zoology, Colorado State University, Fort Collins.

i. Participant in NSF Research Participation for High School Teachers program, Colorado State
University, simimer 1965.

59

The Great Basin Naturalist
60 T. A. WOOLLEY & H. G. HIGGINS Vol. XXV, Nos. 3-4

of cusp toward insertions of lamellar hairs, surface of lamellae and
prodorsum tuberculate; lamellar cusps stout, finger-like, narrower
than lamellae, cylindrical and smooth, projected above prodorsum;
translammella incomplete, interrupted medially; sensillus bristle-like,
with small barbules along proximal two-thirds, with a thin tenninal
filament. Differs from other species of Ceratoppiidae in the lamellae
and lamellar cusps, but principally in the pelopifomi chelicerae. The
specific name is descriptive of the small shears of the distinctive
chelicerae.

Description: Color yellowish-brown; prodorsum broadly triangu-
lar, surface finely tuberculate, tubercles more prominent between
lamellae; rostrum rounded; rostral hairs plumose, curved, shorter
than lamellar hairs, inserted in raised tubercles at lateral margins
about half their lengths posterior to rostral tip (Figs. 1, 5); lamellae
about as wide as width of pseudostigmata, with tuberculated surface,
a dorsal longitudinal ridge extending from base of cusp to toward
insertion of interlamellar hairs, lamellar cusps smooth, cylindrical,
projected above surface of prodorsum, tapered anteriorly; lamellar
hairs setose, longer and less plumose than rostral hairs, extending
beyond tip of rostrum, inserted in tips of cusps; translamella incom-
plete medially; interlamellar hairs setose, projected slightly beyond
tips of lamellar cusps, inserted posterior to longitudinal ridge of
lamellar cusps, inserted posterior to longitudinal ridge of lamellae,
insertions slightly raised; pseudostigmata cup-shaped, at posterior end
of lamellae, slightly postero-mediad of pedotecta I; sensillus longer
than length of lamella, proximal two-thirds setiform with fine bar-
bules on shaft, slightly swollen near fine, terminal flagellum (Fig.
3) ; pedotecta I as seen in figures 1, 2.

Hysterosoma glabrous, nearly spherical, much more rounded than
in Pyroppia lanceolata; nine pairs of dorsal setae; muscle scars
and/or pseudoporosae ^ and fissures as seen in figure 1.

Camerostome elongated, with sclerotized rim; mentum, mental
hairs, rutellum as in figures 2, 6; chelicerae pelopiform (Fig. 4),
with swollen base, elongated shaft and tiny, serrate shears; post-
camerostomal apodemata and ventral setae as seen in figure 2; pedo-
tecta II smaller than pedotecta I, extended antero-laterally; apode-
mata II divided, a broad, decurved band between coxae, an incom-
plete incurved apodeme anterior to main band; a slightly sclerotized.
vertical, medial apodematal bar extending from posterior margin of
apodemata II to anterior margin of apodemata IV; apodemata IV an
arched band investing anterior margin of genital aperture and con-
fluent with perigenital ring; genital aperture about its length anterior
to anal opening, trapezoidal; each cover angled antero-laterally. with
a diagonal sclerotized bar near anterior margin and extending width
of cover, each cover with six genital setae, g:l, g:2, g:3, g:4 about
equidistant from each other, g:5 displaced laterally on cover, g:6
near postero-medial comer, but in line with first four setae; aggeni-
tal setae inserted about length of genital cover posterolateral to
genital opening; fissure iad posterc^lateral to anterior margin of

Dec. 31. 1965 oribatid mite prom Colorado

61

Fig. 1:

Fig. 2:

Fig. 3

Fig. 3:

Fig. 4:

Fig. 5:

Fig. 6:

Fig. 7:

P^^poppia forficula, n. gen., n. sp., from the dorsal aspect, legs omitted
P. forficula, n. gen n. sp., from the ventral aspect, legs partially om tS

SeudoslS;:;' '".^ '^^^il'"' H forficula\o show ^barbules and
i-seudostigmata and sensillus of P. forficula, n. gen., n sp to show

barbules and terminal flagellum g - • sp. lo snow

Pelopiform chelicera of P. forficula, n. gen., n. sp.
Enlarged detail of lamellae and rostrum of P. forficula. n gen

n. sp. trcm the dorsal aspect ' "

Infracapitulum of P. forficula, n. gen., n. sp
Iibia and tarsus I of P. forficula, n. gen., n sP

The Great Basin Naturalist
62 T. A. WOOLLEY & H. G. HIGGINS Vol. XXV. Nos. 3-4

elongated anal opening; anal aperture with prominent preanal piece;
each anal cover with two setae; three pairs of adanal setae, ada:l.
ada:2 mainly posterior to anal opening, ada: 3 lateral and farther
from anal opening than iad fissure.

Legs heterotridactylous, median claw only slightly larger than
laterals; tibia and tarsus I as in figure 7; trochanter III with thin,
slightly plumose seta, similar to larger, more plumose seta of tro-
chanter II in Ceratoppia and Pyroppia.

Length: 384 /a, prodorsum 102 /x, hysterosoma 282 ^u,; width 312 /x.

Nine specimens of this species were collected near the summit of
Berthoud Pass, Colorado, 13 September 1958, by T. A. Woolley. The
type and one paratype are deposited in the USNM.

Discussion: Aside from its smaller size. Paenoppia forficula,
n. gen., n. sp., differs from known species of Ceratoppia and Pyroppia
in the tuberculate prodorsum and lamellae. The most distinctive dif-
ference is the pelopiform chelicerae of P. forficula as contrasted to
the more robust chelicerae of Ceratoppia and Pyroppia.

Hammer (1955, p. 15) states for Pyroppia lanceolata that "The
long feathered hair on coxa III, which is characteristic of the genus
Ceratoppia, is very short in Pyroppia.'' This seems to be a perpetua-
tion of an error of Michael (1887, p. 357) concerning Ceratoppia
bipilis that the prominent "spine on the coxa of each leg of the third
pair near the proximal end, at the outer comer: . . . usually stands
more or less parallel to the pseudostigmatic organ, and it is from
these two spines on each side that the name is given." Willmann
(1931) indicates that this bristle is located on trochanter III — not
coxa III — and observations of representatives of Ceratoppia and
Paenoppia confirm this location. We did not have access to speci-
mens of Pyroppia for examination, but it appears conclusively from
the literature and observations that these erect hairs are on trochan-
ter III and are common to this complex of mites. Such hairs are
found in Paenoppia forficula^ n. gen., n. sp.. but are less robust than
those found in species of Ceratoppia and Pyroppia.

Although no specific relationships can be established, it appears
that Paenoppia forficula is intermediate between Ceratoppia and
Pyroppia, but probably related within the complex of Astigestes,
Cultroribula, Ceratoppia andPyroppia. Research currently underway
should disclose information that will elucidate these relationships.

LiTER.'^TURE Cited

Hammer, Marie. 1955. Alaskan Oribatids. Acta Arctica, Fasc. VII: 1-36.
Michael, A. D. 1887. British Oribatidae. Vol. II, Ray Society, London.
Willmann, Karl. 1931. Moosmilben oder Oribatiden (Oribatei) In Die Tier-
welt Deustchlands 22(5) : 79-200.

STUDIES IN NEARCTIC DESERT SAND DUNE OR'IHOPTERA

A new (ienus and Species of Stenopehnatine Crickets fioni the Kelso
Dunes with notes on its multi-annual life history and k(>\-. Part X

Eitu'st R. Tinkliiiin'

During the past decade the author has made a score of trips to
the great Kelso Dunes studying its fauna and flora; the summers of
1957-1960 assisted by National Science Foundation grants. As these
dunes lie 155 miles north of Indio. California, by road, a total of 6200
miles has been travelled in these trips during the period 1954-1964.
On the second trip in mid Jime. 1955. tracking a strange trail by day
led to the discovery of a crippled .Jerusalem sand cricket. Due to its
great rarity, only a few specimens have been taken during the years.
Often years go by without the finding of a specimen, especially true
of the drouth years which most have been since 1960. The second
specimen was a young nymph, captured the night of October 25.
1957. when a light drizzle of rain was falling on the dunes.

The discovery of specimens was somewhat expedited, when, in
the spring of 1959, I started teaching Extension Courses for San
Diego State College. One of the requirements for my course NATURE
STUDY OF THE DESER T was an overnight field trip to the Kelso
Dunes, usually held in late April and late October of every year.
With 25 to 50 teachers, accompanied often by many children and
quite a few Coleman lanterns, such a group of searchers covers an
area far greater than any scientist can possibly do. Even with such
groups, only a few specimens have been found through the years.
The last finding was five young nymphs, the night of April 20,
1963. when the sands were icy cold (temperature of the sand next
morning at 7:00 was 34° Fahrenheit). At the time of writing in
August. 1965, almost two and one half years later, three are still
alive and are about one-half to two-thirds grown, so that we may
expect the multi-annual life cycle to run three or possibly four or
more years in length.

In a paper soon to be published by the California Academy of
Sciences, the author has laid the groundwork for future studies in
the genus Stertopelmatus Burmeister. From this research it is obvi-
ous that the worker must rely on the chaetotaxy of the legs as the
most significant taxonomic feature to distinguish the various species.
In this article, I recognized and redescribed S. intermedius Davis
and Smith along chaetotaxical lines. Such nebulous characters as
punctation. length of antennal segments, nature of frontal sutures
and the sternal plates have been discarded, along with the form of
the ovipositor, which appears identical in all species. I have laid
the stress on the spination of the legs.

63

The Great Basin Naturalist
64 ERNEST R. TINKHAM Vol. XXV, Nos. 3-4

Hence, if a certain species shows a marked departure from the
normal spination characteristic of the genus Stenopelmatus, then
that creature must be accorded new generic rank. This I have done
with the Kelso Jerusalem Sand Cricket and this consideration has
been arrived at after years of thoughtful weighing of the problem.
The Provisional Key presented below will amply illustrate the dif-
ference between the genus Stenopelmatus and the new genus based
on chaetotaxical taxonomy.

Provisional Key to Californian Stenopelmatine Crickets

1. Tibial spines vestigial or missing on the apical dorsal
margins of the caudal tibiae. Ringlet of 6 apical caudal
calcars almost even and broadly spathulate for areni-
colous habitus. Median or presubapical spur on the ven-
tral surface of the foretibiae absent. Pronotum not ex-
panding anteriorly. Size medium, coloration uniformly

orangish Ammopelmatus new genus

Tibial spines prominently developed on the apical dorsal
margins of the caudal tibiae. Ringlet of 6 apical caudal
calcars uneven in length, usually conical or subconical

in form, the innermost calcar the longest. Median or
presubapical spur on the ventral surface of the foretibiae
always present. Pronotum expanding anterioradly to
house the posterior portions of the very large head. Size
medium to very large; coloration generally dark, espe-
cially on abdomen Stenopelmatus (2)

2. Size large; color of head and pronotum orange red 3

Size medium to small; color of head and pronotum not

orange red but piceus to shining black 4

3. Calcars of the caudal tibiae forming a semi-ringlet of
6 long spurs, the two innermost much the longest and

cylindrical in form longispina Rrunner

Calcars of the caudal tibiae forming a semi-ringlet of

6 spurs, these spathulate or trowel-shaped on their inner
faces, the three inner relatively equal and longer than
the 3 outer spurs fuscus Haldeman

4. Entire body uniformly dark brown with black abdomi-
nal tergites. Caudal tibiae with 5 internal and 2 external

apical dorsal teeth intermedins Davis & Smith

Upper half of head shining black with tan sutural
areas. Pronotum with dorsum bearing irregular areas
of shining black. Femora marked with pale brown fasci-
ations. Caudal tibiae with 3 to 4 internal and 2 external
apical dorsal teeth pictus Scudder

Ammopelmatus Tinkham. new genus
Generic Diagnosis: Size medium, uniformly pale yellowish
orange on head and pronotum; abdomen more whitish orange with

Dec. 31. 1965 NEARCTIC DESF.HT ORTHOPTI'MA 65

the infuscated annular rings on the dorsum ol the abdominal seg-
ments less conspicuous than observed in Stenopelmdtus. The general
features are those of a medium-sized Stenope/rnatus, from which it
is amply distinct in many features. Form more slender than in
Stvnopelmatus, the cephalic features generally similar but lacking
the typical megacephalism portrayed by Stenopelmatus, nor does
the anterior portions of the pronotum seem to house the posterior
margin of the enlarged head as in Stenopelmatus. The pronotum is
relatively narrower in Ammopelmatus and the posterior angles of
the dorsum of the pronotum more evenly and broadly rounded than
in Stenopelmatus. where this angle has a slight flattening on this
latero-posterior face. Anterior lateral angle of the lateral lobes of the
pronotum inore evenly rounded in Amniopelruatus than in Stenopel-
matus^ where all species have a pronounced downward angulation.
Sternal plates in the new^ genus are similar to those observed in
Stenopelmatus.

Ammopelmatus shows several distinctive generic categories in
the spination of chaetotaxy of the legs which quickly separates it
from Stenopelmatus. These are chiefly found in the apical and sub-
apical tooth-like spurs found on the ventral side of the tibiae. In
Ammopelmatus. the ventral margin of the fore tibiae have only one
or two very small spurs placed one behind the other (See A of plate),
whereas the various species of Stenopelmatus have always three
larger ones placed in linear alignment, the third or most posterior
of which being almost median in placement. The mesotibiae are
alike in both genera, there being only an evenly placed pair of apical
conical spurs. Likewise, in the caudal tibiae in Stenopelmatus, there
is always a pair of apical conical spurs on the ventral surface (see
D, F, H, I and J of plate), w^hereas in Ammopelmatus there is only
one centrally placed apical ventral spur (see B and K of plate); a
feature of considerable generic worth. Thus the two features: single
apical ventral spur of the caudal tibiae and the two apical small
spurs placed in linear alignment on the ventral side of the fore
tibiae, quickly identify Ammopelmatus. the new genus. In addition
the six short spathulate calcars of the caudal tibiae and the lack of
large teeth on the dorsal margins of the caudal tibiae, as well as the
truncate or spathulate nature of the apical dorsal teeth of the caudal
tibiae, further identifies Ammopelmatus. Other features, such as the
ovipositor and supra-anal and subgenital plate, appear to be similar
to that in Stenopelmatus.

The diagnostic characteristics peculiar, to the new genus Am-
mopelmatus have apparently been developed through the ages by
adaptation to an arenicolous or ammophilous environment. The loss
of all tibial teeth on the dorsal margins of the caudal tibiae, except
for the anal ones which have developed into a pair whose posterior
surface has developed a truncate spathulate face, appears to be a
direct response to a sand habitat. The broadening, shortening and
evening of the calcars of the caudal tibiae is. likewise, a development
for excavation of sand and a sort of pushing forward movement along

66

ERNEST R. TINKHAM

The Great Basin Naturalist

Vol. XXV, Nos. 3-4

Explanation of Plate

All drawings of fore tibiae are of left fore tibiae, external face; drawings of
caudal tibiae are left caudal tibiae external face. All drawings nnich enlarged
and then reduced.

Ammopflmatus krlsornsis from Kelso Dunes. A. fore tibiae showing only 2
ventral apical teeth ( a and b). B. Caudal tibiae showing 1 ventral apical tooth (1).

Stenopelmatus pictus from Antioch, California. C. fore tibiae showing 2
apical (a and b) and 1 subapical ventral teeth (c). D. Caudal tibiae showing
dorsal tibial spines, calcars (1 to 6) and pair of ventral apical teeth (1 and 2).

Dec. 31, 1965 nearctic desert orthoptera 67

Stenopelmatus intermedius from Lagunitas. California. E. fore tibiae showing
calcars and 2 apical (a and b) and 1 subapical ventral tooth (c). F. Caudal
tibiae showing dorsal tibial spines, calcars (1-6) and pair of ventral apical teeth

(1 and 2).

Stenopelmatus longispina from Palo Alto, California. G. fore tibiae showing
calcars, 2 apical and 1 subapical teeth. H. caudal tibiae showing dorsal teeth,
calcars and two apical ventral teeth.

I. Ventral view of caudal tibiae of S. intermedius showing the pair of apical
teeth and the six calcars.

J. Ventral view of 5. pictus showing apical pair of teeth and the six calcars.

K. Ventral view of caudal tibiae of Ammopelmatus kelsoensis showing single
apical tooth and the spathulate calcars.

the loose surface of the sand of the magnificent mountainous ridges
of sand known as the Kelso dunes.

Genotype: Ammopelmatus kelsoensis, new species

Having studied the life of the sand dunes of the three great
North American Deserts since 1952, and four years, 1957-1960.
under grant from the National Science Foundation, it would appear
that Ammopelmatus is restricted to the Kelso Dunes.

Ammopelmatus kelsoensis Tinkham, new species

Allotype Female: Kelso Dunes, San Bernardino County, Cali-
fornia. April 27. 1963, Ernest R. Tinkham. Type deposited in the
Tinkham Eremological Collection.

Description: Head typical of the Stenopelmatine crickets, the
eyes more narrowly pyriform than in Stenopelmatus, otherwise
closely similar in all general features of the head to that genus.
Pronotum more evenly quadrate than in Stenopelmatus, the lateral
margins parallel to the very broadly rounding posterior lateral
angles; posterior margin squarely truncate, anterior margin evenly
and conspicuously emarginate with that margin typically hirsute.
Dorsum of the pronotum crossed by a conspicuous groove or sulcus
about one-sixth the pronotal length caudad of the anterior margin
and parallel to it, with a median sulcus commencing on the shoul-
ders and angling forward and downwards to the emargination on
the anterior third of the lateral lobes of the pronotum; the lobe in
front of this emargination more evenly rounding into the anterior
margin of the pronotum than in Stenopelmatus, where it is more
angularly produced downwards. Sternal plates similar to those ob-
served in Stenopelmatus. Supra-anal plate semicircularly rounded;
subgenital plate very broadly triangular; ovipositor typical of Ste-
nopelmatus.

Leg spination as foUowts: forelegs with enlarged femora
smooth, the dorsal margin quite arcuately rounded; tibiae with 5
typical calcars surrounding the apical margin (see A of plate) and

The Great Basin Naturalist
68 ERNEST R. TINKHAM Vol. XXV, Nos. 3-4

with only two ventral apical conical spurs in linear alignment;
tarsi quadrimerous, the first segment equal to the remaining three in
length. Meso legs with mesofemora more slender than fore femora
and less arcuate along the dorsal margin; meso tibiae with five ter-
minal calcars, each dorsal margin bearing a subapical toothlike spur
proximad of each lateral calcar; tarsi as in the forelegs, the first
segment giving evidence that it is formed of two fused segments.
Caudal legs with caudal femora short and heavier than in Stenopel-
matus, the niferior margin straight, the superior or dorsal margin
strongly arched with both dorsal margins bearing long brown setae;
caudal tibiae more tumid than in Stenopelmatus^ the external and
internal dorsal margins without teeth except for a large spathulate
tooth at the internal apex and a smaller similar tooth on the external
apex, the latter having a minute tooth just proximad of it and both
margins lined with long brown setae. Ventral surface with only one
apical spur of generic import. The caudal tibiae bear apically the 6
spathulate calcars so typical of the new genus (see figs. B and K of
plate). Caudal tarsi similar to those in the fore and middle legs, ex-
cept that the first segment is slightly longer.

Caliper measurements in mms.: body length 31.0; pronotum 5.8
long by 7.5 in width; caudal femora 9.0 x 3.2; caudal tibiae 8.8 x 2.2;
ovipositor 2.0 mms.

Paratypes: 12 females from the same habitat as the Allotype at
the Kelso Dunes. 1 female adult, 1 subadult, Oct. 25, 1957, the latter
living until July 15. 1960. 1 adult June 14, 1957, 1 adult Oct. 21,
1961, 5 small female nymphs April 20, 1963, 3 adults April 27,
1963. Two of April 20. 196^, females still alive and subadult on
Sept. 7, 1965.

Paratypes identical to the Allotype except for variations in the
ventral apical spur of fore tibiae and caudal tibiae. Oct. 25, 1957, 1
adult and 1 subadult, have one extra ventral apical spur on caudal
tibiae. One Oct. 20, 1963 female and one April 27, 1963 paratype
possess only single spurs ventrally on the fore tibiae and caudal
tibiae. One paratype from Oct. 20. 196^, has one spur on fore tibiae
but two spurs on caudal tibiae.

Range in measurements of adult paratypes: body length 29.0 —
31.0; pronotum 5.0 x 7.3 — 5.8 x 7.5; caudal femore 8.8 x 3.1 x 9.0 x
3.2; caudal tibiae 8.7 x 2.2 x 8.8 x 2.2; ovipositor 2.2 — 2.0 mms.

Date of Night Collecting Trips

1954: June 25; 1955: June 15; 1956: July 13; 1957: June 14,
June 30, Oct. 25; 1958: April 19, June 10. July 1. Sept. 22; 1959:
March 27, April 9. May 9, May 22, Oct. 17; I960: April 30, Oct. 19;
1961: April 15, Oct. 21; 1962: April 28. Oct. 20; 196S: April 20, 27.
Oct. 21 ; 1964: May 2. Oct. 24.

From 1959 on, all April and October night collecting was assisted
by 25 to 50 teachers with Coleman lanterns and flashlights.

Dec. 31. 1965 nearctic desert orthopteba (i9

Collecting Notes

1955: June 15. one crippled adult crawling across dunes in early
morning.

1957: June 14: 1 small nymph, 1 adult. Air and sand temp. 20°C.
Oct. 25. light drizzle on the dunes commencing about 5 p.m.
1 small nymph, 1 adult.

1961: Oct. 21 adult, temp. 25 °C.

1963: April 20. 5 young nymphs collected between 8-9 p.m. when
air tem. 13.2''C and sand surface 13.0°C. Rained and sleeted
during night. At 7:00 a.m., air 6.0°C and sand in shade 2.0°C.
April 27. 8-10 p.m. 3 adults. Air and sand surface were

irc.

Habit.^t: Ammopehnatus has been found only in a limited area
at the north base of a sand declivity, some 15 to 25 feet high, that
lies about 200 yards southwest of the main camping area and about
1 V2 miles ENE of the main peak of sand that is about 700 feet high.
In this area there are scattered clumps of Sandpaper weed. Peta-
lonyx Thurberi, Croton, Croton californicum. Sand dune grass. Ory-
zopsis hymenoides, and in wetter years a great variety of other
plants such as Dicoria canescens. Euphorbia spp., Astragalus sp. and
many others.

Orthopteran Associates: Nocturnal associates on the sands
include: Macrobaenetes kelsoensis Tinkham 1962, Amtnobaenetes,
n.sp., the sand roach Arenivaga, sp., Ceuthophilus fossor and in the
creosote growing from the sands in the marginal area close by such
decticids as: Anoplodusa arizonensis, Capnobotes fuliginosus, Ere-
mopedes n.sp., as well as the Creosote leaf katydid Insara covilleae.
Diurnal associates include: Coniana snowi on its host Coldenia ph-
cata, Xeracris minimum on Petalonyx Thurberi, the ever present
Trimerotropis p. pallidipennis, Ligurotettix c. kurtzei on Creosote
stems, Tanaocerus k. koebeli on marginal Franseris dumosa. Para-
bacillus coloradus on the common Galleta grass Hilaria rigida and
the ground mantid Litaneutria minor.

Faunae Designation: As camp on the Kelso Dunes lies about
2550 feet elevation, the Kelso Dunes lies entirely within the confines
of the Gila Desert which is characterized chiefly by vast mesas and
bajadas of Creosote.

Life History Studies

Egg: The egg of Ammopelmatus kelsoensis is roundly oval, pearly
white in coloration and measures 5.2 mms. in length by 3.3 mms. in
maximum diameter. Only one egg has been found in captivity dur-
ing the decade; partly due to the fact that the rearing has been
mainly of nymphs.

The Great Basin Naturalist
70 ERNEST R. TINKHAM Vol. XXV, Nos. 3-4

Nymph AL Stages: The small nymph collected the night of Oct.
25, 1957, lived until mid- July, 1960, when it died during my ab-
sence on a trip back east. No record of moltings was obtained.

The cold night of April 20, 1963, when temperatures dropped to
37°F., 5 small nymphs were found immobile on the sand. These
were given Numbers 1 to 5, and fed lettuce nightly, but it was not
always possible, due to my great preoccupation with other duties
to screen the sand as often as it should have been screened. Conse-
quently some molts can be expressed only as "early" or "late" in a
particular month while others were observed in the act of molting.
As the eggs of Macrobaenetes Tinkham and Ammoboenetes Hubbell
hatch in late September or early October with the advent of the first
rains, as do other forms of desert insect life, and as the nymphs have
been found in the fall, all evidence indicates that these five young
nymphs had hatched in the fall of 1962. As such they were either
first stadium nymphs or possibly some that were a little larger may
have been in the second stadium.

The salient features of each will be given below followed by a
short summary of the life history of the new genus and species.

No. 1:

1963: April 20: small nymph collected

late June; molted, small pieces of exuvium found July 4.

chewing up of cast skin occurs some days after molt
Aug. 5: found pieces of complete molt.
Sept.: molt in late Sept.
Oct.: molted night of Oct. 16-17

1964: Sept. molted in late September.

1965: Aug.: molted in early part of month.

Sept.: still living at time of writing manuscript.

No. 2:

1963: April 20: small nymph collected
July 4: found complete exuvium
Sept.: molted night of Sept. 2-3
Oct.: molted night of Oct. 20-21

1964: summer: molted but time not known

Oct.: molted between Oct. 20 and Oct. 30

1965: Sept.: died on fourth, was molting Sept. 2 but failed to get
out of exuvium; tried to remove creature from old skin but
failed to survive.

No. 3:

1963: April 20: collected small nymph

June: found complete skin in late June

Dec. 31. 1965 nearctic desert orthoptera 71

Sept.: late, found parts of 2 legs

Dec: 13, found parts of body. 1 leg. parts of palpus

1964: Aug.: died on the third

No. 4:

1963: April 20: collected small nvmph
June: molted in late June
Aug.: molted on 3rd
Sept.: 25: found lying on side at 10:30 p.m., having just

molted. Temp. 28 °C (82.4°F). Sept. 28 only part of cast

skin remained after feeding on it.
Oct. 29: found feeding on freshly molted exuvium

1964: July: died

No. 5:

1963: April 20, collected small nymph

June: molted in late June

Aug.: found pieces of abdomen, front and coudal tibiae indi-
cating molt in early August.

Nov.: molted at 10:30 p.m. 19-20th. Exuvium at head end
of freshly molted cricket

1964: Sept. 14. molted, lying on side. Sleeping upside down on 17th
at noon.

1965: Sept.: 18, found lying on back at 10:30 p.m. with cast skin.

Life History Summary: As far as is known the egg hatches in
the early part of the fall; probably having been laid the previous
spring. Whether there is a molt in the fall or early spring is not
known at the present time. The first known molt is in late June or
very early July; Second molt in late July or very early August,
usually; Third molt usually in late September or up until late Oc-
tober. The Fourth molt occurs in the summer of the second year and
the Fifth molt about September of that same year. The Sixth molt
happens about early September of the third year from the evidence
at hand. It is believed that adults maturing in September of the third
year live through to the following spring when the eggs will be laid,
thus giving a life span from egg to egg laying of four years.

Bibliography

Davis, Alonzo C. and Smith, Leslie N. 1926. Notes on the genus Stenopel-

matus with description of a new species (Orthoptera). Pan. Pac. Ent. 2(4):

174-181, 7 figs.
Hebard, Morgan. 1916. A study of the species of the genus Stenopelmatus

found in the United States. Journ. N.Y. ent. Soc., 24(1): 70-84 pi. VII.
. 1932. New species and records of Mexican Orthoptera, Trans. Amer.

ent. Soc, 58:342-244.

The Great Basin Naturalist
72 ERNEST R. TINKHAM Vol. XXV, Nos. 3-4

Rehn. James A. G. 1902. Two new species of North American Stenopelmatine.

Ent. News, 13:240-241.
Saussure and Pictet. 1897. Biol. Cent. Amer., Orth., I. pp. 286-291, 1 pi with

23 figs.
ScuDDER, Samuel H. 1899. The Stenopelmatinae of the Pocific Coast. Can.

Ent. pp. 113-121.
Strohecker. H. F. 1945. Notes on and descriptions of Mexican Orthoptera.

Ann. ent. Soc. Amer., 38(2) :207-215. 12 figs.
TiNKHAM, Ernest R. 1965. Critical notes on the genus Stenopehnatus Bur-

meister in California with Redesscription of S. intermedius Davis and Smith.

Calif. Acad. Sci., (awaiting publication).

RECORDS OF ATHERINID FISHES AT INLAND LOCALITIES
IN TEXAS AND NORIHERN MEXICO

W. L. Minckley'-'

On June 3-4, 1964, seining collections from the Rio Conchos,
Chihuahua, Mexico, 1.1 miles east of Las Varas (near Ciudad Deli-
cias), and at Ciudad Camargo, included specimens of the tidewater
silverside, Menidia beryllina (Cope) (Arizona State University Nos.
881 and 894, respectively). The collection from Camargo also in-
cluded one individual of Chirostoma sphyraena Boulenger (ASU
1533), and one specimen of an unidentified Chirostoma (ASU 1534).
These, records published by Til ton and White (1964), some provided
by L. S. Campbell {pers. comm.)^ and localities given in Fisheries
Reports of the Texas Parks and Wildlife Department, are the basis
for this paper (Fig. 1).

Records for M. beryllina from Texas (Fig. 1) were obtained from
Federal Aid to Fisheries Project Reports' as follows: between El
Paso and the Big Bend National Park, in the Rio Grande (presum-
ably above the mouth of the Rio Conchos); and listed as "an invader
that dominates localities in Dove Creek and the South Concho River
[Colorado River system. Texas]. Especially abundant in the reser-
voir created by the Guinn Dam [on Dove Creek]." Campbell {pers.
comm.) provided two record localities for the Pecos River drainage —
Imperial Reservoir, Pecos County, and Red Bluff Reservoir, Reeves
and Loving counties. The following information was provided for
other Texas areas: "We have also taken these silversides [M. beryl-
lina^ from Lake Brownwood, Colorado River drainage. Brown Coun-
ty [reported on by I'ilton and White, 1964], and from the Concho
River. Colorado River watershed, Tom Green County. I cannot ac-
count positively for the spread of these fish, but I suspect that they
were introduced through the actions of bait dealers. Tidewater and
Mississippi silversides [A/, audens Hay] are also reported from some
areas of the Red River system. We did not find silversides in our
earlier work (1957-1958), but apparently once introduced they ex-
perienced an enormous expansion. Since 1963 the populations have
been drastically reduced through unknown factors." Additional
records in the Colorado River (Tilton. 1961; Tilton and White,

1. Department of Zoology, Arizona vState University. I'enipe.

2. I thank Mr. L. S. Campbell of the Texas Parks and Wildlife Department for providing in-
formation used in this report. Colleitions in Mexico were made while en route to collect elsewhere
for NSF Project GB-24(il, the support of which is gratefully acknowledged. I'ermits to collect fi.shes
in Mexico were cordially granted bv the Direc( ion (Jeneral do I'psca e Industrias ('onexas. Mexico
D. F., Mexico. Able assistance in collecting was provided bv Arthur C. Kchternachl. Richard K.
Koehn. and Alan P. Wick.

1 Menidia beryllina is listed in the following Job Completion Reports from the Texas Parks and
Wildlife Department that are available to me: Job B-32, "Fisheries Reconnais.sance." Fed. Aid Proj.
F-5-R-Q (l'»hii, F-5-R-l() Il')(i5l. and F-5-R-I1 (1964); Job B-i5. "A Preimpoundmenl Survey of
Twin Butts Reservoir and its Im luded Watershed," F-5-R-I0 (19(i3); and Job B-H. ".Xn Investigation
of Waters of the El Paso Area in Order to Evolve Efficient Managonient of the (lame Fish Resource,"
F-'i-R-ll flWH).

73

74

W. L. MINCKLEY

The Great Basin Naturalist

Vol. XXV, Nos. 3-4

Fig, 1 . Distributional records for Menidia beryllina in streams of Texas and
northern Mexico. The numerous records of Robinson (1959), from the lower Rio
Grande, are spanned by the five localities plotted; others are from Tilton (1961),
Tilton and White (1964), and from. data in the present paper. The most southern
record station in Mexico is Ciudad Camargo; Chirostoma spp. also were obtained
at that locality.

1964) are from near La Grange, Texas, and from Lakes Inks, Bu-
chanan, and Marble Falls.

Prior to 1951, M. beryllina had been recorded in the Rio Grande
system from the mouth at Brownsville, to 35 miles upstream from
Laredo, Texas (Robinson, 1959; Tilton and White, 1964). Hubbs
(1957), however, suggested earlier that the fish might inhabit most
of the Tamaulipan Biotic Province of Texas (Blair, 1950). and this
would extend its possible range to near the mouth of the Devil's
River, Val Verde County. It is conceivable that it invaded upstream
in both the Colorado and Rio Grande, perhaps in response to de-
creasing discharges and increasing salinities that result from irriga-
tional use, but the distances involved and inhospitable habitats that
exist make this less probable than is direct introduction by man.

Experimental introductions of marine fishes into Imperial and
Red Bluff reservoirs of the Pecos River system, the same impound-

Dec. 31, 1965 atherinid fishes 75

merits from which M. beryllina has been caught, raises the possibility
of accidental introduction of Menidin in those areas as eggs or larvae
in water used for transport of the larger species.' Because atherinids
are difficult to transport as adults it seems unlikely that Menidia was
transferred a long distance as such. The promiscuous spawning of
species of Menidia (Bumpus, 1898; Bigelow and Schroeder, 1953)
would make eggs available for accidental collection, and M. beryllina
occurs abundantly in Texas coastal waters from which many of the
marine fishes were obtained for stocking (Gunter, 1945). Movement
down the Pecos River, then through the Rio Grande and into the
Rio Conchos. may explain the origin of the Mexican stocks. With
styrofoam coolers and other types of bait containers now available,
short-distance transfers to the Colorado River basin, for example,
might have occured as suggested by Tilton and White (1964) and
by Campbell (see above).

Stream occurrences of M. beryllina are few and it seems worth-
while to give the following habitat notes. At Las Varas the Rio
Conchos was 50 to 75 feet wide and four feet deep at the deepest
point. Pools alternated with riffles and bottoms were gravel except
in quiet backwaters where silt occurred. Aquatic macrophytes were
absent, but the shore was lined with a thick gallery of trees and
shrubs. The water had a milky color and was salty to taste; dis-
charge was estimated at about 125 cubic feet per second. At Ciudad
Camargo there was a large pool upstream from the highway bridge
and small pools and riffles near the bridge. Again the water was
Milky in color and was malodorous, appearing polluted with sewage.
Vegetation on the banks was cattail {Typha), low grasses, and
small shrubs. Various aquatic plants were on riffles {Chara, Najas,
and Potamogeton), but pools were unvegetated. Bottoms were of
deep silt except on the swiftest riffles. Discharge was about 80 cubic
feet per second. At both localities the atherinids were concentrated
below riffles in moderate current, or were in eddies. I hey were
obviously in mid-water and appeared to avoid the banks. Five speci-
mens of M. beryllina were obtained at Camargo and 134 at Las
Varas. In the latter collection, specimens are from near 35 to more
than 60 mm. in standard length.

Of the two additional atherinids obtained at Camargo, Chirostoma
sphyraena is endemic to the Rio Lerma basin of south-central Mexico,
in the Pacific watershed (Meek, 1904). It could occur in the relative-
ly well-collected Rio Conchos only through introduction. The un-
identified specimen of Chirostoma may . also have come from the
Lerma basin, from which a number of species of that genus are
described.

Movement of fishes by man from place to place in Mexico is
undoubtedly increasing with construction of reservoirs and develop-

4. For data on the'ie introductions refer to Job Completion Reports F-1. "Experimental Stocking
of Marine Fish Species in Saline Waters of Westem Texas," Fed. Aid Proj. F-5-R-9 (19(i2), F-7-R-<>
(1<«>2), F-7-R-I0 (l*t(i^). and F-5-R-11 (lf)(i4). and other reports listed therein. Species of marine
fishes that are reported as introduced are as follows: Roccus saxatilis (Walbaunil, Cynnsiion nehulosus
(Cuvier), Leiostomus lanthurus Lacepede, Micropogon undulatus (Linnaeus), Sciacnops ocellata
(Linnaeus), Pogonias rromis (Linnaeus), and Paralichthrs Icthostigma Jordan and Gilbert.

The Great Basin Naturalist

76 w. L. MiNCKLEY Vol. XXV. Nos. 3-4

ment of sport fisheries. Solorzano (1963) discussed the biology of
Chirostoma estor Jordan in reference to its possible artificial propa-
gation and introduction into areas other than its native range as a
food fish; perhaps attempts have already been made to introduce
some species of Chirostoma. Eggs of some species of the genus, at least
of those from lake Patzcuaro, float (Solozano, 1961, 1963), and
easily could be pumped into holding tanks meant for other fishes
and inadvertently transferred outside of their native ranges.

Literature Cited

Bi^MR, W. F. 1950. The biotic provinces of Texas. Texas Jour. Sci., 2:93-117.
BiGELOw, H. B.. AND W. C. ScHROEDER. 1953. Fishes of the Gulf of Maine.

U. S. Fish and Wildl. Ser., Fish. Bull., 74. 53:i-viii, 1-577.
BuMPUS, H. C. 1898. The breeding of animals at Woods Hole during the

months of June. July, and August. Science (n.s.), 8:850-858.
GuNTER, G. 1945. Studies on marine fishes of Texas. Publ. Inst. Mar. Sci..

Univ. Texas, 1:1-190.
HuBBS. C. 1957. Distributional patterns of Texas fresh water fishes. SW Nat..

2:89-104.
Meek, S. E. 1904. The fresh-water fishes of Mexico north of the Isthmus of

Tehuantepec. Publ. Field Columbia Mus., Zool. Ser.. 5:i-lxiii. 1-252. 17 pis.
Robinson, D. T. 1959. The ichthyofauna of the lower Rio Grande. Texas and

Mexico. Copeia, 1959 (3):253-256.
Soi-ORZANO Preciado, A. 1961. Contribucion al conocimiento de la biologia del

charal prieto del Lago de Patzcuaro. Mich. (Chirostoma bartoni Jordan y

Everman. 1896). Sec. Indust. Commerc, Dir. Gen. de Pesca Ind. Con..

Mexico D. F.. Mexico. Pp. 1-70.
. 1963. Algunos aspectos biologicos del pescado bianco del Lago de

Patzcuaro, Mich. {Chirostoma estor Jordan. 1879). Ibid. Pp. 1-15.
TiLTON, J. E. 1961. Ichthyological survey of the Colorado River of Texas.

Unpubl. M. A. Thesis. Univ. Texas. Pp. 1-73 (not seen).
. AND R. L. White. 1964. Records of Menidia beryllina from several

central Texas impoundments. Texas Jour. Sci., 16:120.

A LIST OF SCARABAEIDAE BEETLES OF II IE
NEVADA TEST SITE'

Dorald M. Allre<l and D Elden Beck-
In August, 1959, Brigham Young University initiated an ecologi-
cal study at the Nevada Test Site as a basis for determining the ef-
fects of nuclear testing on native animals (refer to Brigham Young
Univ. Sci. Bull., Biol. Ser., 11(2): 1-52; 1963). During the succeeding
five years, studies dealing with selected groups of arthropods yielded
a number of beetles of the family Scarabaeidae. These were submit-
ted to Dr. Henry F. Howden, Canada Department of Agriculture.
Ottawa, to whom we are grateful for the identification of the speci-
mens. For the most part, specimens were trapped in pit cans as de-
scribed by AUred, et al. {Ibid., pp. 8-9). In some cases, however
beetles were taken by hand directly from the host plant.

In the listing that follows, the plant community and the host
plant are given when known (for a description of the plant communi-
ties refer to Brigham Young Univ. Sci. Bull., Biol. Ser.. 11(4): 1-5;
1963). When no host plant is listed, the specimens were taken from
pit cans in the plant community indicated.

Aphodius fucosus Schm.

Of a total of 222 beetles of this species, 219 were taken from the
Grayia-Lycium community during March, April, and May. Three
specimens were taken from the Coleogyne community in March. The
majority were found during April.

Aphodius militaris Lee.

Five of 13 specimens taken were from the Grayia-Lycium com-
munity in March, May, and June. One specimen was taken from
Lycium andersonii in January, and another from Lycium sp. in
November, both in the Larrea-Franseria community. Six were taken
from a Mixed community in November and December.

Aphodius nevadensis Horn

Two of a total of 127 specimens were taken from a Grayia-
Lycium community in February. Seven were taken from a
Coleogyne community — one in January, three in February, one in
November, and two in December. One was collected from the Pinyon-
Juniper community in November. A total of 1 1 7 was taken from the
Mixed community — one in November, 106 in December, and nine

1. BYUAEC Report COO-1355-11, Field work completpd under Contracts .VFd 1 -I i78(i and
.\T(11-1 ,1.52(i between the Atomic Energy Commission and Brigham > oung University.

2. Department of /.oology and Entomology. Brigham Young University, Provo. Uuh.

77

78 ALLRED AND BECK Vol. XXV, Nos. 3-4

in January. Over the whole test site, the predominant number of
specimens was taken in December.

Chnaunanthus flavipennis Horn

One specimen was taken from the Grayia-Lycium community
in May.

Cyclocephala longula Lee.

Two of a total of seven specimens were taken from a Grayia-
Lycium community in August, three from a Larrea-Franseria com-
munity in August, and two from a Coleogyne community in July
and August.

Diplotaxis deserta Fall

A total of 19 specimens was taken from a Mixed community.
Eighteen were taken in June and one in August.

Diplotaxis haydeni Lee.

Five specimens were taken from a Pinyon-Juniper community
in July.

Diplotaxis incuria Fall

Five specimens were taken from the Larrea-Franseria community
in June.

Diplotaxis insignis Lee.

One specimen was taken from the Pinyon-Juniper community
in July.

Diplotaxis moerens moerens Lee.

A total of 77 specimens was taken. Thirty-five were found in the
Larrea-Franseria community in August, and one was taken from
Lycium pallidum in the same community in July. One specimen was
taken from the Atriplex-Kochia community in September, 13 from
the Coleogyne community in August, and four from the Grayia-
Lycium community in July. Twenty-two were taken from a Mixed
community — -1 1 in July, four in August, and seven in September.
With reference to the whole test site, the greatest number of indi-
viduals was taken in August.

Diplotaxis pacata Lee.

One specimen was found in a Larrea-Franseria community in
June.

Diplotaxis subangulaia Lee.

Thirty of 167 specimens taken were from the Grayia-Lycium
community — one in May, four in July, 24 in August, and one in
September. Forty-two specimens were found in a Larrea-Franseria
community — five in June, two in July, and 35 in August. Four were

Dec. 31. 1965 scarabaeidae beetles 79

taken from a Coleogyne community in July, and 52 from a Mixed
conmiunity — one in June. 43 in July, six in August, and two in Sep-
tember. With reference to the whole test site, the predominant num-
bers were taken in July and August.

Ochodaeus sparsus Lee.

One specimen was taken from a Grayia-Lycium community and
two from a Coleogyne community, all in August.

Paracotalpa granicollis llald.

A total of 226 specimens was taken. From the (irayia-Lycium
community 25 were taken in January and 34 in February. Five were
found in the Salsola community in February. In the Larrea-Franseria
community 19 were taken in January and 10 in February. In the
Atriplex-Kochia community 99 were taken in January, four in
•March, and two in April. In the Mixed communit}^ 26 were taken in
January, one in February, and one in March. Over the whole test
site, the predominant number was taken in January.

Jorgensen (Pan-Pacific Ento., 39(3): 154-6; 1963) noted some
aspects of the biology and distribution of this species at the Nevada
Test Site. He mentioned specifically the flight and mating activities
of the adult males and females.

S erica alternata Lee.

Thirty-four specimens were taken from the Pinyon-Juniper com-
munity in July and 15 from a Mixed community in August.

Serica perigonia Dawson

Five specimens were found in the Pinyon-Juniper community
in July.

Summary

In addition to those listed above, others collected for which data
are not complete are Aphodius (near) talpoidesi Brown, Phyllophaga
{Listrochelus) sp., Serica curvata Lee, and Serica jalli Dawson.

In sequence of greatest abundance, the most common species
known to occur at the test site are Paracotalpa granicollis, Aphodius
fucosus, Diplotaxis suhangulata, and Aphodius nevadensis^ respec-
tively. Seasonally, the greatest numbers of species in the adult stage
were most active in July, June and August, and May, respectively,
although greatest numbers of individuals were found in January,
July. April, August, and December.

It is expected that further collecting specifically oriented toward
the scarabeids will reveal additional species.

ANGUS MUNN WOODBURY
1886-1964

Vasco M. Tanner'

Angus Munn Woodbury, Professor of Biology, Emeritus, Uni-
versity of Utah, Salt Lake City, and his wife, (irace Atkin Woodbury,
were killed in a two-car head-on collision Saturday, August 1, 1964,
a mile north of Loveland. Colorado, on U.S. Highway 287. Dr. Wood-
bury was employed, at the time of his death, as a lecturer at the
BSCS Institute at the State University of Colorado, Fort Collins.

Angus M. Woodbury was born in Saint George, Utah on July 11,
1886. His parents. John Taylor Woodbury and Mary Evans Wood-
bury, were early settlers in St. George. They were highly respected
members of this isolated pioneer settlement. Dr. Woodbury early took
advantage of educational opportunities. He attended the Brigham
Young University in 1906, after which he became assistant Forest
Ranger in 1908. He retired from the forest service in 1920, having
advanced to that of deputy supervisor. From 1920 to 1926 he was
engaged in agricultural work and was teaching assistant at Dixie
Junior College. In 1926 he again attended Brigham Young Univer-
sity, graduating in 1927 with a Bachelor of Science degree, with a
major in zoology.

In the fall of 1927 Dr. Woodbury began his graduate work in
zoology at the University of Utah. He was granted a Master of
Science Degree in the spring of 1928. After serving as an instructor
in 1929 he made preparations to continue his training in zoology at
the University of California, Berkeley, from which institution he
graduated with a Doctor of Philosophy Degree in 1931. In 1925 he
was appointed Park Naturalist at Zion National Park which position
he held during the summers until 1933. It was while serving as a
pioneer naturalist that he became interested in ecology which re-
sulted in his two major contributions in this field: Biotic Relation-
ships of Zion Canyon^ Utah with Special Reference to Succession,
1933, and Principles of General Ecology, 1953.

In 1931 Woodbury attained the rank of Assistant Professor at the
University of Utah, and advanced to Associate Professor in 1939, and
Professor in 1942. He was made head of the Department of Vertebrate
Zoology in 1948 and retired as Emeritus professor in 1952.

Retirement had no deterring effects on Woodbury's activities
since he served as director of Ecological Research at Dugway Proving
Grounds from 1952 to 1956. In 1957 he was made director of
Ecological Research on the Colorado River Project which position he
held until 1964. His ability to organize and direct research work on
the Ecology of Disease Transmission in native animals at Dugway
met with great favor by the Army Chemical Corps. The monographic
reports on the flora and fauna of the Upper Colorado River Basin
carried out in connection with several collaborators, laid the founda-

1. I'nif.'ss,,r ,,f •/,,.., lofjv and Knl(>ni<ili>nv. Bn^;lumi ^uiiiit,' rniv.MMlv. I'r.n.i. I'lali.

82

The Great Basin Naturalist
VASCO M. TANNER Vol. XXV, NoS. 3-4

Angus M. Woodbury, general ecologist, biologist;

University of Utah, 1928-1964.

Dec. 31. 1965 angus m. woodbury 83

tion for future work on the ecology of this new recreational water-
way, Lake Powell. One of the convincing arguments made in support
of the protection of Rainbow Bridge without building a dam on Aztec
Creek, was made by Dr. Woodbury — Science, 1961, 133(3464):
1572-1583.

His accomplishments were many. One that stood out was his
teaching ability. He was a devoted leader of his students. His quiet,
kindly manners endeared him to his students and friends. His
lectures were well oragnized and frequently interspaced with clever,
meaningful jokes followed by a "whisper-soft laugh". He drew from
his scholarly background and wide field experience many clinching
examples to buttress his reasoning.

Professor Woodbury's publications, which include several books,
number over 100 and cover a wide range of subject matter. He was
greatly interested in the history of Utah, as evidenced by his several
papers including: The Route of Jedediah S. Smith in 1826 from the
Great Salt Lake; and A History of Southern Utah and its National
Parks. He was well informed on Utah birds. As a senior co-author
with Clarence Cottam and John Sugden, a sizeable manuscript on the
birds of Utah awaits publication. These three authors did, however,
publish in 1949: An Annotated Checklist of the Birds of Utah. In
1945 Woodbury and Henry N. Russell pubHshed: Birds of the Navajo
Country, 158 pages. Woodbury's interest in the reptiles of Utah
resulted in two published reports: Catalogue of the Reptiles of Utah
and Studies of the Desert Tortoise, Gopherus agassizii (with Ross
Hardy), which will long be useful references when dealing with this
class of Utah animals. A complete list of all of Dr. Woodbury's
published papers is included as a part of this paper.

Professor Woodbury was an active and enthusiastic supporter of
a number of professional organizations. He was punctilious as to
his attendance and participation in society meetings, even though it
necessitated his traveling long distances outside of Utah. Some of
the scientific organizations of which he was a member and his rank
follow: Fellow of the American Association for the Advancement of
Science; Fellow of the American Ornithological Union and a member
of the council of this organization; Fellow of the Herpetological
League; Fellow, Utah Academy, Science, Arts and Letters; Member,
Population Reference Bureau; Member, American Society Ichthyolo-
gists and Herpetologists; Member, Ecological Society of America;
Member, Cooper Ornithological Society; Member, Biological Society
of Washington; Member, Nature Conservancy; and Member, Ameri-
can Museum Natural History. He was active in the Audubon Society
Program in Utah.

Angus M. Woodbury and Grace Atkin were married on January
15. 1909. They are survived by their four sons and two daughters:
Lowell Angus; Marian (Mrs. A. Herbert Gold); Max Atkin; Edith
Rae (Mrs. Robert C. Pendelton); Dixon Miles; and John Walter.
The sons all have Ph.D. degrees in the biological sciences, having
followed in their father's footsteps. The daughters did not depart

The Great Basin Naturalist

84 VAsco M. TANNER Vol. XXV, Nos. 3-4

from the field of biology since they are married to Ph.D. biologists.

I have never known a more devoted couple than Angus and
Grace Woodbury. After marriage their ontogony was such that their
lives, home life and research activities resulted in an integrated
family life. They traveled together in field study activities and made
their home a haven for biology students. Mrs. Woodbury once said,
"I early learned to live in peaceful coexistence with a scientist."

Few native sons of Utah have risen to such prominence in the
field of biology as did Angus M. Woodbury. His untimely death
came as a great shock to his family, colleagues, and host of friends.
At the time of his death he was actively engaged in several research
projects. His accomplishments and influence as a teacher will live
on in the enrichment of the lives of others.

List of Angus M. Woodbury Published Papers 1927-1964

1927

Studies in the Sphecoidea of Utah. Utah Acad. Sci., Proc. 4:27-28, 1927.
1928

The reptiles of Zion National Park, [Utah]. Copeia No. 166:14-21, March

23, 1928.
The reptiles of Zion National Park. Utah Acad. Sci., Proc. 5:11-12, 1928.
1929

Notes on the spiders of Washington County, Utah [with *R. V. Chamberlin].

Biol. Soc. Wash., Proc. 42:131-141, Pis. I and II. March 30, 1929.
The snails of Zion National Park. Nautilus 43:54-61, October, 1929.
A new rattlestake from Utah. Univ. Utah Bull. 20(6), Biol. Ser. 1(2), 3 pp.,
2 figs., December. 1929.
1930

Insect studies of Zion National Park. Utah Ed. Rev. 23:382-383, 403, 404,
April, 1930.
Crotalus confluentus concolor (Woodbury). Antivenin Inst. Amer., Bull. 4:23,
May, 1930.
1931

A descriptive catalog of the reptiles of Utah. Univ. Utah Bull. 21(5). Biol.

Ser. 1(4), X -h 129 pp., 58 figs., February, 1931.
The route of Jedediah S. Smith in 1826 from the Great Salt Lake. Utah

Hist. Quart., 4:35-46, April, 1931.
A list of the common birds of Zion National Park. [List XI, p. 169, Bird
watching in the west, R. S. Twining]. Metropolitan Press, Portland,
Oregon, October, 1931.
1933

The scratching of the spurred towhee. Condor 35:70, March, 1933.
Biotic relationships of Zion Canyon, Utah, with special reference to suc-
cession. Ecol. Monogr. 3:147-246, 29 figs., April, 1933.
The bird rookeries of the islands of Great Salt Lake [with W. H. Behle].

Utah .Acad. Sci.. Proc. 10:165, 1933.
Animal life. Conservation and management of wild life. [Editor and contri-
butor Parts IX and X in] Utah resources and activities, Utah State
Board of Education, 1933.
1936

Animal relationships of Great Salt Lake. Ecology 17:1-8. 1 fig., January,
1936.
1937

Planning for the conservation of Utah's wildlife. Utah Ed. Rev. 30:173-174,

February 8, 1937.
An evolutionary time scale. Evolution 4:7-8, illus., June, 1937.

'Senior ,\uthur

Dec. 31. 1965 angus m. woodbury 85

A brown pelican record from Utah. Condor 39:225. September. 1937.

A record of Tadarida macrotis from Utah. Jour. Mamm. 18: 51 '5. November

14. 1937.
1938

The Lower Sonoran in southwestern Utah. Science 87:484-485, May 27, 1938.
Red-naped sapsucker and rufous hummingbird. Condor 40:125, May, 1938.
An hour in the life of a broad-tailed hununingbird [with John W. Sugden].

lbid\ 40:160-162. 1 fig.. July, 1938.
1939

Bird records from Utah and Arizona. Ibid. 41:157-163, July, 1939.
Obsei-vations on the breeding of the western chipping sparrow. Oologist 56:

114-116 (No. 629), October, 1939.
[Abstract] Management of aquatic wildlife in the Great Basin. Utah Acad.

Sci., Arts Lett., Proc. 16:41, 1939.
[Abstract] Reproductive cvcles in the sagebrush lizard Sceloporus g. graciosus

[with Marian Woodbury]. Ibid. 16:65, 1939.
1940

Management of aquatic wildlife in the Great Basin. Sci Monthly 50:307-322.

illus.. April, 1940.
Antler-eating by rodents. Science. 92:127-128, August 9, 1940.
Northern crested lizard collected in Utah [with Ross Hardy]. Copeia 1940

(3):205, November 14. 1940.
The dens and behavior of the desert tortoise [with Ross Hardy]. Science 92:

529. December 6. 1940.
Pigmy conifer limits in Utah. Utah Acad. Sci., Arts Lett., Proc. 17:13. 1940.
1941

Nesting of the gray flycatcher [with *Henrv N. Russell. Jr.]. Auk 58:28-

37, January, 1941. "
Copulation in gopher snakes. Copeia 1941(1):54, March 25, 1941.
Bird habitats of the Salt Lake Region. Audubon Mag. 43:253-264, Illus., May,

1941.
Animal migration — periodic-response theory. Auk 58:463-505, October, 1941.
Changing the "hook-order" in cows. Ecology 22:410-411. October, 1941.
1942

Man's biological future. Science 95:121-122, January 30, 1942.

Studies of the rat snake. ElapHe laeta, with description of a new subspecies

[with Dixon M. Woodbury]. Biol. Soc. Wash., Proc. 55:133-142. 2 figs.,

October 17, 1942.
Notes on migrations of the painted lady butterfly in 1941 [with John W.
Sugden and Clvde Gillette]. Pan.-Pac. Ent. 18:165-176. 1 fig.. October,

1942.
Status of the name Crotalus concolor. Copeia 1942(4): 258, December 28,

1942.
My rattlesnake bite. Utah Acad. Sci. Art. Lett., Proc. 19:179-184. 1942

[printed 1944].
1943

The Joshua tree flower as a source of food [with Maurine Noall, Corliss R.

Kinney, and James Sugihara]. Ibid. 20:49-50, 1943 [printed 1944].
Report of rattlesnake bite of J. Dwain Anderson [with Mrs. J. Dwain An-
derson]. Ibid. 20:185-188, 1943 [printed 1944].
I 944

Type locality of Perisoreus canadensis capitalis Ridgwav (with Clarence

Co\.\.am\ Auk 61:131-132. January, 1944.
Research on the high school level. Nation's Schools 33:47. February, 1944.
Research is needed in high schools and colleges. Utah Eld. Rev. 37:271-273,

306. May 8. 1944.
A history of southern Utah and its national parks. Utah Hist. Quart. 12: 1 1 1-22,

illus., July-October, 1944 [printed in 1945: revised and reprinted in

1950.
Notes on Mexican snakes from Oaxaca [with Dixon M. Woodbury]. Wash.

'Senior .\uthor

The Great Basin Naturalist

86 vAsco M. TANNER Vol. XXV, Nos. 3-4

Acad. Sci., Jour. 34:360-375. November 15, 1944.
Further light on the "fossil" bird nests of the Great Salt Lake Desert [with

'William H. Behle and Walter P. Cottam]. Utah Acad. Sci. Art. Lett.,

Proc, 21:4, 1944.
1 945

Birds of the Navajo country [with Henry Norris Russell, Jr.]. Univ. Utah

Bull. 35(14), Biol. Ser.'9{l), 158 pp., 7 figs., 1 fold, map, March 1, 1945.
A new Gerrhonotus lizard from Utah. Biol. Soc. Wash.. Proc. 58:5-10. 2

figs.. March 21. 1945.
[Abstract] Results of color-banding California Gulls at Egg Island, Great

Salt Lake, Utah [with William H. Behle]. Utah Acad. Sci. Art. Lett.,

Proc. 22:8. 1945.
Life-history studies of the sagebrush lizard Sceloporus g. graciosus with special

reference to cycles in reproduction [with *Marian Woodbury]. Herpe-

tologica 2:175-196, 3 figs., 1945.
1946

Huge migratory flock of purple martins in LTtah. Condor 48:42, January,

1946.
Color-banding California Gulls at Great Salt Lake, Utah [with William H.

Behle and John W. Sugden]. Univ. Utah Bull. 37(3), Biol. Ser. 10(2),

14 pp., 2 figs., June 30, 1946.
1947

Notes on migrations of the Painted Lady butterfly in 1945 [with *John W.

Sugden and Clyde Gillette [. Pan.-Pac. Ent. 23:79-83. April, 1947.
The Mohave rattlesnake in Utah [with Ross Hardy]. Copeia 1947(1 ):66.

April 20. 1947.
Distribution of pigmy conifers in Utah and northeastern Arizona. Ecology

28:113-126, 6 figs., April, 1947.
[Review of] The puma, mysterious American cat. Ibid. 28:328-329, July,

1947.
The speckled rattlesnake in NW Arizona [with Ross Hardy]. Herpetologica

3:169, 1947.
1948

Studies of the desert tortoise, Gopherus agassizii [with Ross Hardy]. Ecol.

Monogr. 18:145-200, 25 figs., April, 1948.
Animals and salinity in the Great Basin. Amcr. Nat. 82:171-187. 3 figs..

May. 1948.
Marking reptiles with an electric tattooing outfit. Copeia 1948(2) : 127-128.

June 30, 1948.
Resource problems of Lhah. [In symposium on conservation of natural re-
sources in Utah.] Utah Acad. Sci. Art. Lett., Proc. 25:55-57. 1948

[printed 1950].
1949

Annotated check-list of the birds of LTtah [with Clarence Cottam and John W.

Sugden]. Univ. Utah. Bull. 39(16). Biol. Ser. \\{2), 39 pp.. March, 1949.
Ecology' and national welfare. Arner. Soc. Prof. Biol., News 3:3, 8. November,

1 949.
1950

[Revision of] A history of southern Utah and its national parks. Utah Hist.

Quart. 12:111-223, illus., July-October 1944 [rinted in 1945; revised and

reprinted in 1950].
Unusual snake records from Utah and Nevada [with Earl W. Smart]. Her-
petologica 6:45-47, 1950.
A snake den in Tintic Mountains. Utah [with Richard M. Hansen]. Ibid.

6:66-70, 2 figs.. 1950.

1951

snake den in Tooele County, Utah. [Introduction — a ten year study].
[Introduction to symposium with Basil Vetas, Gordon Julian. Henry R.
Glissmeyer, F. LaMarr Heyrend and Anson Call. Earl W. Smart, and
Raymond T. Sanders.] Ibid.' 7:\4; 7 figs., March 27, 1951.

"Senior .\uthor

Dec. 31. 1965 angus m. woodbury 87

Results of the Pacific gull color-banding project [with Howard Knight]. Con-
dor 53: 57-77. 11 figs., March 27, 1951.
Meteorological data in ecology. Science 114:216-217, August 24, 1951.
1952

a. Ecological Taxonomy. Science, 115 (2992). May 2.

b. Hybrids of Gopherus berlandi and G. ugassizii. Herpetologica, 8:33-36.

July 15.

c. Amphibians and reptiles of the Great Salt Lake Valley. Herpetologica,

8:42-50. July 15.
1953

Methods of field study in reptiles. Herpetologica, 9:87-92. July 22.
1954

a. Principles of general ecology. McGraw-Hill Book Co., New York. 503 pp.

b. Ecolog>'^ of Disease transmission in native animals. Semi-annual report of

the Director. Ecological Research. University of Utah.

c. Study of reptile dens. Herpetologica, 10:49-35.

1955

1956

1957

1958

An annual cycle of the desert jackrabbit. Ecology^ 36:353-356.

b. Science, population and arid lands. Science, 122:200.

c. Ecology and th population problem. Science, 122:831-834.

d. Ecology of tularemia transmission in native animals. The 1954?! 955

annual report of the Director, Ecological Research, Univ. Utah. 57 pp.

e. [Editor and contributor] Symposium on ecology of disease transmission

in native animals. Army Chemical Corps, Dugway. Utah, 1-112.

f. Ecological studies of tularemia in Utah. In symposium on ecology of

disease transmission in native animals. Army Chemical Corps. Dugwaj',
Utah, 81-87.

[Editor] Ecological check lists (from Dugway, Utah): plants, inverte-
brates, vertebrates. The Great Salt Lake Desert Series, Ecol. Research,
Univeisity of Utah. 125 pp.

The Colorado dam controversy. Sci. Monthly, 82:304-313.

Symposium on uses of marking animals in ecological studies: Introduction,
Ecology 37:665; Marking amphibians and reptiles. Ecology 37:670-674.

[With *Billy N. Day and Harold Egoscue] The Ord Kangaroo rat in
captivity. Science 124 (3220) :485-486.

Comfort for survival. Vantage Press, N.Y., 104 pp., March 25.
b. The Colorado dam controversy. Sci. Monthly, 8:199-200.

[and staff members] Preliminary report of biological resources of the
Glen Canyon Reservoir, Univ. Utah, Anthropological papers, 31:1-219,
(Glen Canyon Series. No. 2).

The name [Crotalus viridis concolor Woodbury.] Copeia, 1958:151.

[and Grace A. Woodbury] Ralph Vary Chamberlin, educational and
cultural contributions. The Biologist, 40:21-26.

1959

[with Stephen D. Durrant and Seville Flowers] Survey of vegetation in
the Glen Canyon Reservoir Basin. Univ. Utah, Anthropological Papers
36:1-53 (Glen Canyon Series No. 5).

Clasping behavior of the leopard lizard. Herpetologica, 15:118.

[Biological editor] Ecological studies of the flora and fauna in Glen Can-
yon. Univ. Utah Anthro. Papers, No. 40:1-226.

Working plan for ecological studies. Glen Canyon Reservoir. Upper Colo-
rado River Basin. Univ. Utah Anthro. Papers, 40:1-20. [Original Report
to National Park Service, 1957].
Amphibians and reptiles of Glen Canyon. Univ. Utah Anthro. Papers,

40:155-148.
Ecological Study of Colorado River in Glen Canyon. Univ. Utah Anthro.
Papers, 40:149-176.

The Great Basin Naturalist

88 vAsco M. TANNER Vol. XXV, Nos. 3-4

1960

a. [with S. D. Durrant and Seville Flowers] A survey of vegetation in the

Flaming Gorge Resei-voir Basin. Univ. Utah Anthro. Papers, 45:1-121.

b. The Colorado River — the physical and biological setting. In the Colorado

. . . River of the West. Utah Hist. Quarterly, 28:199-208.

c. Protecting Rainbow Bridge. Study of dangers to Rainbow Bridge from

waters of Glen Canyon Reservoir brings surprising answers, Sci.. 132:
519-528. Aug. 26.

d. Animal Community. Published in Encyclopedia of Technology and Sci-

ence. McGraw-Hiih N.Y., pp. 409-411.

e. Rainbow Bridge, a letter to the Editor in reply to F. R. Fosberg. Science

132 (3435): 1262, October 28.

f. The desert tortoise needs protection. El Paisano, Banning. Calif. No. 19:

6-7.

1961

1962

1963

[with 'Monroe E. Wall] Steroidal sapogenins from the Joshua tree. /.

Economic Botany 15:79.
Protection of Rainbow Bridge National Monument. Sci. 133(3464) : 1572-

1583.
[with S. D. Durrant and Seville Flowers] Survey of vegetation in the

Navajo Reservoir Basin. Univ. Utah Anthro. Papers, 51:99.
Editor, Ecological studies of the flora and fauna of Navajo Reservoir

basin, Colorado and New Mexico. Univ. Utah Anthro. Papers, 55. Biota,

an introduction and summary, 55:1-14.

[with S. D. Durrant and Seville Flowers] A survey of vegetation in the
Curecanti Reservoir basins. Univ. Utah Anthro. Papers, 56:1-98.

b. [with Clarence Cottam] Ecological studies of birds in Utah. Univ. Utah

Biol. Series, 12(7). 1-28.

c. Editor. Ecological studies of the flora and fauna of the Curecanti Reser-

voir basins, western Colorado. Univ. Utah Anthro. Papers 59:1-285.
Foreword, introduction and summary, p. 1-11.

[with Jean Musser] A limnological study of the Fremont River, Capitol
Reef National Monument. Special Report to Regional Office, U. S. Nat-
ional Park Service. Sante Fe, N. M.. 52 pages.

Editor. Studies of biota in Dinosaur National Monument, Utah and Colo-
rado. Univ. Utah Div. Biol. Sci. Misc. Papers, 1:1-77. Introduction, 2

PP-

[with Delbert W. Argyle] Ecology and limnology of Green River. In
Univ. Utah Div. Biol. Sci. Misc. Papers 1 : 7-48.

Biological-ecological aspects of Betatakin Canyon, Navajo National Monu-
ment. Arizona. Univ. Utah Div. Biol. Sci., Misc. Papers 2:1-56.

Cartoon highlights of Orin Nelson Woodbury, by Grace A. Woodbury.
12 p. Edited and published by Angus M. Woodbury, 248 University St.,
Salt Lake City.

Reminiscences of Ann Cannon Woodbury. Arranged, edited and privately
published by Angus M. Woodbury. 248 University St., Salt Lake City,
Utah.

1964

A review of the ecology of Eniwetok Atoll. Pacific Ocean. Univ. Utah.

Inst, of Env. Biol. Research, 1-123.
Disease dissemination among biotic communities of the Great Salt Lake

Desert. Bull. Univ. Utah No. ; Inst, of Env. Biol. Research, Misc.

Papers, No. 4; Bull, of Utah Engineering Exp. Station. No. . 282 pp.

'Senior Aiithor

3 2044 072 231 111

Date Due