Vol. 61

January 1985

THE

No. 1

Pan-Pacific Entomologist

ARNOLD, R. A.—Geographic variation in natural populations of Speyeria callippe (Boisduval)

(Lepidoptera: Nymphalidae). 1

JOHNSON, J. B .—Aleiodes sp. (Hymenoptera: Braconidae) reared from an anomalous host,

an adult of the Eremochrysa punctinervis species group (Neuroptera: Chrysopidae). 24

POLHEMUS, D. A. and J. T. POLHEMUS—Myrmecomorphic Miridae (Hemiptera) on mistle¬

toe: Phoradendrepulus myrmecomorphus, n. gen., n. sp., and a redescription of Pilopho-

ropsis brachypterus Poppius. 26

GOEDEN, R. D. and D. W. RICKER—The life history of Ophraella notulata (F.) on western

ragweed, Ambrosia psilostachya De Candolle, in southern California (Coleoptera: Chrys-

omelidae). 32

CAMPBELL, C. L. and K. S. PIKE—Life history and biology of Pyrausta orphisalis Walker

(Lepidoptera: Pyralidae) on mint in Washington. 42

WILSON, S. W. and R. J. GILL—The first record of the delphacid Liburniella ornata in

California (Homoptera: Fulgoroidea). 48

MARI MUTT, J. A.—A new species of Orchesella from Manitoba, Canada (Collembola:

Entomobryidae). 50

KRAUS, B. —Oviposition on the backs of female giant water bugs, Abedus indentatus: the

consequence of a shortage in male back space? (Hemiptera: Belostomatidae). 54

MASON, P. G.—The larva of Cyphomella gibber a Saether (Diptera: Chironomidae). 58

WHITFIELD, J. B.—The Nearctic species of Deuterixys Mason (Hymenoptera: Braconidae). 60

KAMM, J. A. —Cutworm defoliators of Ryegrass. 68

WILSON, S. W.—Descriptions of the immature stages of Delphacodes bellicosa (Homoptera:

Fulgoroidea: Delphacidae). 72

TURNER, W. J.—Checklist of Pacific Northwest Tabanidae with new state records and a

pictorial key to common species (Diptera: Tabanidae). 79

WILKERSON, R. C.—A new genus and species of horse fly (Diptera: Tabanidae) from Bolivia 91

SCIENTIFIC NOTES .38, 40

SAN FRANCISCO, CALIFORNIA • 1985

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

EDITORIAL BOARD

J. A. Chemsak, Editor

R. S. Lane, Associate Editor

W. J. Pulawski, Treasurer J. T. Doyen

R. M. Bohart J. A. Powell J. E. Hafernik, Jr.

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addressed to the Treasurer, Dr. Wojciech J. Pulawski, California Academy of

Sciences, Golden Gate Park, San Francisco, CA 94118-9961.

Application for membership in the Society and changes of address should be

addressed to the Secretary, Vincent F. Lee, California Academy of Sciences, Gold¬

en Gate Park, San Francisco, CA 94118-9961.

Manuscripts, proofs, and all correspondence concerning editorial matters should

be addressed to Editor, Pacific Coast Entomological Society, 201 Wellman Hall,

University of California, Berkeley, CA 94720. See back cover for instructions.

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Pacific Coast Entomological Society

OFFICERS FOR 1984

H. I. Scudder, President W. J. Pulawski, Treasurer

J. Gordon Edwards, President-Elect V. F. Lee, Secretary

Statement of Ownership

Title of Publication: The Pan-Pacific Entomologist.

Location of Office of Publication, Business Office of Publisher and Owner: Pacific Coast Entomological

Society, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-9961.

Editor: J. A. Chemsak, 201 Wellman Hall, University of California, Berkeley, California 94720.

Managing Editor and Known Bondholders or other Security Holders: None.

This issue mailed February 20, 1985

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 1-23

Geographic Variation in Natural Populations of Speyeria callippe

(Boisduval) (Lepidoptera: Nymphalidae)

Richard A. Arnold

Department of Entomology, University of California, Berkeley, California 94720.

Abstract.— The geographic variation in 8 wing characters in 35 populations of

Speyeria callippe (Boisduval) is described and the covariation of the characters

summarized. Univariate (SS-STP), discriminant function, and principal compo¬

nents analyses were performed to describe the variation, interpret its origins, and

group together geographically adjacent localities that are similar in their charac¬

teristics. These techniques are used to test the ability of characters to differentiate

subspecies. All three techniques corroborate that these characters, which have

previously been used to differentiate 16 subspecies, cannot do so with any statis¬

tical significance. Thus, the subspecific taxonomy of S. callippe is revised. The

presently recognized 16 subspecies are reduced to 3, whose morphological features

correlate with their geographic distributions.

The subspecies concept and its utility have been critically evaluted by numerous

systematists (Bohme, 1978; Mayr, 1969; Wilson and Brown, 1953), yet many

problems remain in defining and naming subspecies. At issue is a growing dis¬

content with an arbitrary taxonomic category, the subspecies, which often fails

to accurately describe infraspecific variation. In certain animal groups, such as

butterflies, where the ratio of systematists to species is high, minor differences in

morphology are accorded relatively high value in determination of taxonomic

rank. Mayr (1969) defined the subspecies as “an aggregate of local populations of

a species, inhabiting a geographic subdivision of the range of the species, and

differing taxonomically from other populations of the species.” Clearly the de¬

termination of how distinctive geographical segregates of a species must be to

warrant formal taxonomic recognition is an arduous task. Most butterfly subspe¬

cies are named on the basis of one or a few wing characters, principally slight

differences in color, maculation pattern, or size. Generally these differences are

intuitively perceived by a worker, rather than supported by statistical analyses of

character variation throughout the geographic range of the species.

This study describes the geographic variation in 8 wing characters of S. callippe.

Characters that have previously been used to differentiate subspecies are analyzed

as well as others to discover which are most useful for defining patterns of variation

that relate to geographic distribution of the butterflies. Hovanitz (1941, 1943),

Howe (1975), Moeck (1957), and Sette (1962) speculated that the variation in

several characters, including size, wing color, and silver-spot patterns, of the

numerous subspecies of Speyeria callippe was clinal. Also they proposed that the

pattern of variation in several characters partitioned populations into several

geographical segregates or subspecies. Despite the large degree of overlap in the

PAN-PACIFIC ENTOMOLOGIST

Distribution of Subspecific Taxa

Figure 1. Geographic distribution of the formerly recognized subspecific taxa of Speyeria callippe.

Taxa in large capital letters, with a Great Basin distribution, are green-disced populations, whereas

those represented by the smaller lower-case letters, with a Califomia-Oregon distribution, are brown-

disced populations.

range of variation of numerous wing characters, 16 subspecies are currently rec¬

ognized (dos Passos, 1964; Miller and Brown, 1981).

Several univariate and multivariate numerical taxonomic techniques were used

to analyze the geographic variation in 35 populations scattered throughout the

range of S. callippe. Based on these analyses and other numerical taxonomic

studies (Arnold, 1983, 1985) new conclusions about phenotypic character vari-

VOLUME 61, NUMBER 1

ation and the infraspecific nomenclature of Speyeria callippe are presented. The

presently recognized 16 subspecies (Fig. 1) are reduced to three whose morpho¬

logical features correlate with geography.

Data Collection and Analysis

Materials.— Initially, 35 populations or operational taxonomic units (OTU’s),

were selected for this study (Table 1). Samples of 10 males from each population

were chosen by use of a random numbers table. Each sample was composed of

individuals collected on the same date at the same locality. The subspecific identity

(sensu dos Passos, 1964) of each sample was determined. Nineteen additional

populations were later used as unknowns to test the effectiveness of the characters

under analysis in distinguishing subspecific taxa.

Table 1. List of 35 Speyeria callippe populations (OTU’s) analyzed using univariate and multi¬

variate statistical techniques. OTU’s associated with two subspecific names (e.g., OTU number 2

comstocki: callippe), indicate that these are intermediate populations.

Former

OTU subspecific

OTU no. identity Locality data

CALC

comstocki

CHEW

comstocki: callippe

MOCH

callippe : comstocki

MILL

near callippe

LIVE

callippe : comstocki

NCAL

callippe : comstocki

CALL

callippe

NAPA

Uliana

LCAL

Uliana

LRCA

Uliana: rupestris

GLEN

Uliana

MEND

Uliana

RCAL

rupestris

TRIN

elaine : rupestris

ECAL

elaine

CALS

sierra

PLUM

sierra: juba

BUTT

juba

FIDD

juba

ICAL

juba

PION

laura

ALPI

nevadensis

MLCA

laurina

LAUR

laurina

MCAL

macaria

GREE

macaria : laurina

TEHA

macaria

SCAL

semivirida

NEVA

nevadensis

HARM

harmonia

CALG

calgariana

MEAD

meadii

UTAH

harmonia

GALL

gallatini

GUNN

meadii

San Diego Co., CA

Chew’s Ridge, Monterey Co., CA

Arroyo Mocho, Alameda Co., CA

Oakland, Alameda Co., CA

Los Mochos Cyn., Alameda Co., CA

Del Puerto Cyn., Stanislaus Co., CA

San Bruno Mtn., San Mateo Co., CA

Mt. Veeder, Napa Co., CA

Boggs Mtn., Lake Co., CA

Eel River, Mendocino Co., CA

Mendocino Pass, Glenn Co., CA

Shasta-Trinity Co. line, CA

Mt. Shasta, Siskiyou Co., CA

Siskiyou Summit, Jackson Co., OR

Red Clover Valley, Plumas Co., CA

Feather River, Plumas Co., CA

Big Battle Creek, Butte Co., CA

10 mi. E. Fiddleton, Amador Co., CA

Washoe Co., NV

Monitor Pass, Alpine Co., CA

Tulare Co., CA

Glenville, Kern Co., CA

Tehachapi Mtns., Kern Co., CA

Harney Co., OR

several counties, NV

several counties, UT

Calgary, Alberta, Canada

several counties, CO

several counties, UT

several counties, WY

Boulder Co., CO

PAN-PACIFIC ENTOMOLOGIST

Table 2. Characters used in univariate, discriminant function and principal component analyses.

Character types are continuous (C) and nonordered (NO) multi-state. Non-correlated characters that

were used in the numerical taxonomic analyses are denoted by a *.

Character

Name

Type

Veins measured or states

*1) FWLE

forewing length

base of R to distal end of Ml

2) FWWD

forewing width

R4 to 2A at distal margin

*3) FWLW

ratio of forewing

length/width

ratio of FWLE/FWWD

4) FDCL

forewing discal cell

length

base of M to its junction

with Ml

5) FDCW

forewing discal cell

width

Ml to M3 at distal end of cell

*6) FDLW

ratio of forewing

discal cell

length/width

ratio of FDCL/FDCW

7) HWLE

hindwing length

base of M to distal end of M1

8) HWWD

hindwing width

Rs to 2A at distal margin

*9) HWLW

ratio of hindwing

length/width

ratio of HWLE/HWWD

*10) HDCL

hindwing discal cell

length

base of M to its junction

with Ml

11) HDCW

hindwing discal cell

width

Ml to M3 at distal margin

12) HDLW

ratio of hindwing

discal cell

length/width

ratio of HDCL/HCDW

*13) DISC

disc color of

hindwing venter

green, brown, or green and brown

*14) SUHW

silver spot pattern

on hindwing

none, only near body, only

distal, or entire wing

*15) USGC

ground color of

dorsal facies

pale yellow, yellow, orange, or

reddish-orange

Characters .—Twelve quantitative (continuous) and 3 qualitative (non-ordered)

wing characters (Table 2) were measured for each of the 350 individuals. Scores

for the 10 males in each population were averaged to derive a population score

for each character. Continuous characters were measured along specified veins

with dial calipers to the nearest 0.1 mm. These characters were selected based on

their ease of measurement. Overall size has been previously used to delimit some

of the subspecies (Howe, 1975; dos Passos and Grey, 1947), thus the continuous

characters were analyzed in an attempt to define the range of size variation in

each subspecies. Length and width raw data were transformed logarithmically to

standardize the data prior to analysis. Thus each character contributes toward the

overall resemblance inversely in proportion to its variability among the entire set

of OTU’s. Characters with a small range of variation contribute equally as char¬

acters with a large range of variation. The characters described by ratios were, by

definition (Clifford and Stephenson, 1975), transformed. A histogram for each

continuous character was plotted to verify that its distribution was normal. For

the three non-ordered characters, each specimen was compared to a series of

standards representing each character state. Pearson product-moment correlation

coefficients were calculated for each pair of characters to determine the extent

VOLUME 61, NUMBER 1

each character imparts unique information as opposed to information in common

with the other characters being analyzed.

Data analysis .—The approach used to analyze geographic variation in the mor¬

phological characters was as follows:

1) partitioning of the variation of each character by analysis of variance (uni¬

variate or SS-STP analysis);

2) correlation of characters in order to represent the variation in them by a

smaller number of factors (principal components analysis);

3) simultaneous comparison of the variation in all characters to portray the

similarities between populations and assess the degree of geographic segregation

of similar populations (discriminant function analysis);

4) test the ability of the characters used to differentiate the 16 subspecies and

determine the best subspecific classification.

Results

Character correlations .—Mean character states for each population were used

to compute the correlation coefficient between every pair of the 15 characters

originally measured. The correlation between characters measures the similarity

(concordance) of their patterns of geographic variation. Seven of the characters

were highly correlated with one or more other characters (>0.70). These were

eliminated from further numerical taxonomic analyses. Thus the geographic vari¬

ation in 8 uncorrelated or weakly correlated characters, FWLE, FWLW, FDLW,

HWLW, HDCL, DISC, USGC, and SUHW, was analyzed (Table 2).

Univariate analysis .—The recognition of 16 subspecies (dos Passos, 1964) and

previous analyses of infraspecific variation (Hovanitz, 1941, 1943) suggest that

the character variation is discontinuous and that the range of callippe can be

divided into different homogeneous regions. This categorization can be tested by

using multiple comparison techniques (Sokal and Rohlf, 1969).

In order to ascertain which characters could partition the 35 populations into

statistically significant subsets, analysis of variance over all localities was per¬

formed for each of the 8 morphometric characters using Power’s (1970) version

of Gabriel’s (1964) simultaneous test procedure (SS-STP), a multiple comparison

test between means (Gabriel and Sokal, 1969). This procedure calculates character

means of populations, ranks the means for each character, and computes ho¬

mogeneous subsets of means based on the variances. Computations were made

using the computer program UNIVAR (Moss, 1969).

Population means were plotted for each character to examine patterns of vari¬

ation. Following the technique of Doyen (1973), shaded circles appear to the left

of the ranked means on the accompanying distribution maps (Figs. 2-9). These

circles do not correlate with the overlapping subsets of the STP technique, but to

a second grouping of the means based on dividing the total range of ranked means

for each character into 5 equal intervals. Largest values are represented by open

circles, and smallest values by closed circles.

Only one character, ventral disc color (DISC) exhibited variation concordant

with geography. All brown OTU’s (depicted by open circles in Fig. 9) occur from

the Sierra Nevada Mountain range to the west coast, while all green populations

(black circles) are located in the Great Basin and Rocky Mtns. (Fig. 9). The green-

brown intermediates (half open-half black circles) are from southern Oregon, an

PAN-PACIFIC ENTOMOLOGIST

L POPULATION

MEAN

GRADIENT

BUTT

21.5

PION

21.1

- ICAL

21.1

HARM

21.0

III

SCAL

20.7

IIII

NEVA

20.6

inn

CALL

20.6

inn

FIDD

20.5

nun

UTAH

20.5

mini

MILL

20.5

nnnn

GUNN

20.2

nnnni

LIVE

20.2

minim

MEAD

20.1

minimi

GALL

19-7

ninninn

CALG

19-6

ninninn

MOCH

19.4

minimi

ALPI

19.4

minim

NAPA

19-3

minim

mlca

19-3

nnnni

CALS

18.9

nnnni

CALC

18.9

nnnn

LAUR

18.9

mini

CHEW

18.8

mini

PLUM

18.8

mini

NCAL

18.7

nnnn

RCAL

18.7

mini

TEHA

18.6

mini

ECAL

18.5

TRIN

18.4

GREE

18.4

MCAL

18.3

inn

LCAL

17.9

LRCA

17-7

MEND

17-3

GLEN

17.2

Figure 2. Results of univariate (SS-STP) analysis for character FWLE. The circles are used to check

for clinal variation in FWLE, after the technique of Doyen (1973).

VOLUME 61, NUMBER 1

S...7 “ - - .

jpj

SYMBOL POPULATION

SCAL

CALC

NEVA

CALL

LIVE

HARM

ALPI

UTAH

PLUM

CHEW

MCAL

MEAD

MULL

GALL

MOCH

■MLCA

BUTT

MEND

NCAL

GLEN

LAUR

ECAL

TEHA

GREE

MEAN

1.27

1.25

1.24

1 .24

1.24

1.23

1.22

1 .21

1.21

1.20

1 .20

1.17

1.16

1.15

GRADIENT

III

IIII

inn

nnn

mini

nnnn

innni

nnnn

ilium

nnnn

mini

inn

Figure 3. Results of univariate (SS-STP) analysis for character FWLW. The circles are used to

check for clinal variation in FWLW, after the technique of Doyen (1973).

PAN-PACIFIC ENTOMOLOGIST

J ^

* +3t++-J. : m

mm..

..7 - - - . .

10 .

IL 4 "

#:;*• ..

»•; ••

L POPULATION

MEANS

GRADIENT

PLUM

3.55

CHEW

3.50

MILL

3.49

III

NAPA

3.47

1III

CALG

3-43

inn

BUTT

3-43

inn

MOCH

3.43

nnn

UTAH

3-42

nnn

TEHA

3-42

mini

GALL

3-39

nnnn

TRIN

3.38

imiiiii

ALPI

3-38

iiiiiiim

GLEN

3.37

iiiiiiiiin

LIVE

3-32

iiiiiiiiiin

HARM

3.30

iiiiiiiiiiin

LCAL

3.28

iiiiiiiiiiin

MEND

3.28

iiiiiiiiiiin

MLCA

3.26

iiimimm

SCAL

3-24

iiiiiiiiiiin

GREE

3.23

imiiiimm

NCAL

3.22

iiiiiiiiiiiin

RCAL

3.22

iimiiiiim

LRCA

3.21

nnnnnin

FIDD

3.15

iiiiiiiiiin

CALC

3.12

minimi

MEAD

3.11

minim

GUNN

3-08

nnnni

PION

3.08

nnnn

LAUR

3.06

mini

MCAL

3.06

nnn

CALS

3.06

inn

ECAL

3-03

ICAL

2.99

NEVA

2.96

CALL

2.78

Figure 4. Results of univariate (SS-STP) analysis for character FDLW. The circles are used to

check for clinal variation in FDLW, after the technique of Doyen (1973).

VOLUME 61, NUMBER 1

* 6 \

.. -

4<'I i

V * 0

- J

V\ /

S'l \

SYMBOL

-&■

POPULTION

MEND

MEAN

1 .25

GRADIENT

ALPI

1.23

PLUM

GALL

UTAH

1.16

1.15

III

IIII

— r

CHEW

1.14

IIII

TEHA

1. 14

IIII

(

GREE

1.13

IIII

CALG

1.13

inn

MOCH

1.13

(

MEAD

1.12

j \

ICAL

1.12

inn

I t

LIVE

1.12

inn

( \

MILL

1.12

inn

V i

BUTT

1.12

linn

NEVA

1. 12

PION

1.11

mini

S <5

HARM

1.10

mini

NCAL

1.10

mini

S |

NAPA

1.09

mini

FIDD

1.09

mini

GUNN

1 .09

mini

\—# \

MLCA

1.09

niinii

'' h

CALS

1.09

nnnni

SCAL

LAUR

1.08

nnnni

GLEN

1.07

nnnni

TRIN

1 .07

nnnn

ECAL

1.03

mini

CALL

1.03

LCAL

1.03

inn

NCAL

LRCA

1.03

1.01

RCAL

0.99

CALC

0.98

Figure 5. Results of the univariate (SS-STP) analysis for character HWLW. The circles are used

to check for clinal variation in HWLW, after the technique of Doyen (1973).

PAN-PACIFIC ENTOMOLOGIST

•V>

MEANS GRADIENT

NEVA

9.96

HARM

9.78 -

CALG

9.77

III .

CALL

9.39

IIII-

SCAL

9.37

iiiii.

I CAL

9.25

min

CALS

9.21

min

MLCA

9-17

LAUR

9.02

UTAH

8.72

mini

MCAL

8.52

mini

RCAL

8.52

mini

GALL

8.51

mini

MEAD

8.49

mini

LCAL

8.48

mini

GUNN

8-37

mini

CALC

8.36

mini

LCAL

8:05

nnnn

NCAL

7-94

ninni

PION

7.93

nnnn

LRCA

7-87

nnnn

MOCH

6.84

nnnn

MILL

6.72

mini

CHEW

6.70

LIVE

6.63

iiiii

ALPI

6.54

BUTT

6.53

FIDD

6.42

PLUM

6.32

NAPA

6.27

GREE

6.21

TEHA

6.15

TRIN

5-93

GLEN

5.60

MEND

5.58

Figure 6. Results of the univariate (SS-STP) analysis for character HDCL. The circles are used to

check for clinal variation in HDCL, after the technique of Doyen (1973).

VOLUME 61, NUMBER 1

** - — 1

.*•

r .

, i ------

MEANS gradient

BUTT

4.00

CALS

4.00

TRIN

4.00

ECAL

3-95

LCAL

3-90

III

RCAL

3-88

III

FIDD

3-80

nil

PION

3-50

inn

MILL

3-50

inn

LRCA

3.50

inn

CALL

3-40

I CAL

3.35

inn

MEND

3-35

min

GLEN

3-05

MOCH

3.00

MLCA

2.95

inn

NAPA

2.95

NCAL

2.90

CHEW

2.70

LAUR

2.60

CALC

2.60

LIVE

2.50

MCAL

2.05

TEHA

1.80

ALPI

1.75

GREE

1.65

MEAD

1.40

GUNN

1.10

GALL

1.06

CALG

1.06

HARM

1.05

SCAL

1.00

PLUM

1.00

UTAH

1 .00

NEVA

1.00

Figure 7. Results of the univariate (SS-STP) analysis for character USGC. The circles are used to

check for clinal variation in USGC, after the technique of Doyen (1973).

PAN-PACIFIC ENTOMOLOGIST

• l -&

m mmim K

Ss@

f^-w

■#

’ 7 “ - -

4 -,,

SYMBOL

MEAN GRADIENT

O-

CALC

CHEW

MOCH

MILL

LIVE

NCAL

CALL

NAPA

LCAL

PLUM

ALPI

SCAL

NEVA

HARM

CALG

MEAD

UTAH

GALL

GUNN

CALS

TEHA

ECAL

MEND

LRCA

GLEN

MCAL

GREE

TRIN

RCAL

MLCA

LAUR

PI ON

BUTT

FIDD

ICAL

4.0

3.8

3.6

3.5

3.4

3.3

3.2

3.0

2.2

2.0

1.0

1.6

1.3

1.0

III

ini

iiiii

inn

iiiiii

iiiii

Figure 8. Results of the univariate (SS-STP) analysis for character SUHW. The circles are used to

check for clinal variation in SUHW, after the technique of Doyen (1973).

VOLUME 61, NUMBER 1

-vffriU

Rft

•Vl'v-:: : .V;:::

;V:^;:;-;7. ; .

• - ^ *

>:&} : .‘ : . : ."-s;;.

W‘

,7 - - -

- a._

SYMBOL POPULATION

CALC

CHEW

MOCH

MILL

LIVE

HCAL

CALL

NAPA

LCAL

LRCA

GLEN

MEND

PCAL

TRIM

ECAL

CALS

PLUM

BUTT

FIDD

ICAL

PI ON

ALPI

MLCA

LAUR

MCAL

GREE

TEHA

-e-

SCAL

NEVA

HARM

CALG

MEAD

UTAH

GALL

GUNN

MEAN

2.C

3.0

2.0

1.0

GRADIENT

Figure 9. Results of the univariate (SS-STP) analysis for character DISC. The circles are used to

check for clinal variation in DISC, after the technique of Doyen (1973).

PAN-PACIFIC ENTOMOLOGIST

area where Great Basin and Cascade populations may naturally come into contact.

Discordant variation was noted for the remaining 7 characters (Figs. 2-8), each

of which was characterized by abrupt discontinuities. Only one character, FWLE

(Fig. 2), offered a hint of localized clinal variation. The SS-STP subsets of the

other 7 characters included considerable overlap of green and brown populations.

Discriminant function analyses .—With this technique, measurements of two or

more characters are weighed and combined linearly to provide maximal separation

for two or more groups. Each analysis begins with groups of known identity, in

this case the 16 subspecies sensu dos Passos (1964) and Miller and Brown (1981).

This multivariate method uses the non-overlapping information contributed by

each character to produce a linear function that will classify the known specimens

with a minimum probability of misidentification. Different combinations of char¬

acters may be used to achieve statistical distinction between groups. These pro¬

cedures assist in determining which characters most effectively partition the OTU’s

into groups.

Direct and step-wise discriminant function analyses were performed on the

same data matrix using the Mahalanobis generalized distance, D 2 (Klecka, 1975),

to measure phenetic similarity of groups. The 8 characters were first analyzed

using the direct method, where all variables are simultaneously entered into the

analysis. Discriminant analysis defines axes through the multi-dimensional clus¬

ters of individuals resulting in the maximum separation of the clusters.

Later, a step-wise analysis was performed, in which characters were added or

removed from the discriminant function analysis based on their contribution to

the total distance among multivariate means. This criterion is measured by their

F-values, a statistical indication of the relative importance of each character.

Characters having the largest F-values are repeatedly selected for partitioning

groups of individuals (i.e., clusters) as they contribute more to the classification

process. Each step adds the character with the next highest F-value to yield the

greatest separation of clusters. Thus the step-wise procedure indicates a character’s

importance in separating groups, and the least number of characters necessary to

separate the clusters.

Both analyses revealed that ventral disc color (DISC), a non-parametric char¬

acter, had the greatest power of separation based on the F-values for each of the

8 characters (Table 3). DISC could separate all green populations from all brown

populations, but could not significantly separate any subsets within the green or

brown populations. According to the F-values (Table 3), the next most powerful

separation was based upon a parametric character, HDCL, but it could not separate

the populations into meaningful clusters. DISC was the only character which

could significantly separate local populations.

Linear discriminant function analysis also provides an objective means by which

unknown populations can be classified based on the analyses of the reference

groups. The scores for individuals from each group form distributions, each with

a mean score. The midpoint between mean scores is used to determine group

membership, because this point gives the minimum misclassification. Unless the

separation of known groups is complete, scores for some specimens will fall within

the range of scores delimiting another population, and these individuals would

be misclassified. The rate of misclassification of known specimens provides a

measure to the effectiveness of a set of characters in identifying new specimens.

VOLUME 61, NUMBER 1

- 8.00 - 4.25 - 0,50 - 3,25 - 7.00

7.000

- 5.125

■ 3.250

- 1,375

- 0.500 £

- 2.375

- 4.250

■- 6,125

- 8.000

Figure 10. Territorial map of individuals in the 35 Speyeria callippe populations (represented by

letters A-Z and numbers 0-9) based upon discriminant function scores 1 and 2 (axes are labelled DF

1 and DF 2). Centroids of the 35 populations are indicated by stars. Note the extensive amount of

overlap by individuals of different populations.

- 6.125

- 1.875

1,875

5,125

DF 1

The classification routine correctly identified only 82 of 190 unknowns (43.2%)

as members of the local populations to which they actually belong. As a double

check, the 350 individuals which composed the reference populations were then

run as unknowns. Only 161/350 (46.0%) of the individuals were correctly clas¬

sified. Thus most individuals were classified as having their highest group prob¬

ability as members of populations other than that to which they actually belong.

The territorial map of discriminant score 1 vs. score 2 illustrates this degree of

overlap among populations (Fig. 10).

In order to remove any bias introduced into the discriminant functions due to

the three non-parametric (qualitative) characters, a third analysis using only the

5 continuous characters to partition OTU’s was performed. During the classifi¬

cation routine only 60/190 unknowns (31.6%) and 119/350 (34.0%) reference

individuals were correctly classified. This strongly suggests that overall size is a

poor character for distinguishing subspecies of S. callippe.

Principal components analysis (PCA ).—The prior assignment of populations

into arbitrary reference groups is not necessary with this technique. Combinations

of character axes (eigenvectors) representing OTU’s are transformed to uncorre-

PAN-PACIFIC ENTOMOLOGIST

Table 3. Ranking of characters based on their ability to partition populations (OTU’s) as determined

by step-wise discriminant function analysis. Higher F-values indicate characters which best discrim¬

inate between groups.

Character

F to enter or remove

13) DISC

129.9

10) HDCL

35.9

15) USGC

24.6

1) FWLE

7.9

9) HWLW

1.2

3) FWLW

1.1

14) SUHW

1.0

6) FDLW

0.9

lated axes represented in an n-dimensional character hyperspace. The first com¬

ponent accounts for the greatest percentage of variation, the second accounts for

the next greatest percentage of vaiiation, etc., until all variation among all OTU’s

is expressed. Thus, the spatial relationships between OTU’s in the n-dimensional

character space are preserved while the axes are rotated. If a major proportion of

the variability between characters is explained by the first 3 eigenvectors, the

OTU’s can be represented in 3-dimensions without much distortion to their real

relationships. Factor loadings for all characters on each component provide a

basis for inferring which characters are discriminatory and which are uncorrelated.

The first three principal components accounted for 83.2% of the variance ob¬

served (Table 4). Factor loadings for the three non-ordered state characters are

all near zero along the first and second principal components. This indicates that

the qualitative characters contribute little to the character variation along these

components. In contrast, the five continuous characters exhibit higher loadings

along the first and second components. The relative homogeneity of eigenvalues

suggest that the continuous characters along these principal components reflect

general size factors for the OTU’s.

Continuous characters along the third component have low loadings, while the

non-ordered state characters exhibit higher loadings. This indicates that the non-

ordered characters are instrumental in explaining the variation along this com¬

ponent. Continuous characters express differences due to size, while the non-

Table 4. Factor loadings (i.e., eigenvalues) as determined by the principal components analysis.

Factors with the highest eigenvalues explain most of the observed variance.

Factor

Eigenvalue

Percent of

variance

Cumulative

percentage

6.05

43.2

3.09

23.0

66.2

2.38

17.0

83.2

0.80

5.7

88.9

0.73

5.2

94.2

0.39

2.8

95.9

0.25

1.8

98.7

0.18

1.3

100.0

VOLUME 61, NUMBER 1

Table 5. Communalities (range 0.0-1.0) of 7 wing characters as determined by principal compo¬

nents analysis. Characters with lower communalities should partition the OTU’s (populations) into

more meaningful groups, i.e., they are most diagnostic.

Character

Communality

1) FWLE

9) HWLW

3) FWLW

6) FDLW

14) SUHW

10) HDCL

15) USGC

13) DISC

0.94540

0.91284

0.75851

0.73140

0.68258

0.61904

0.54793

0.21685

ordered color (DISC and USGC) and silver-spotting (SUHW) characters provide

a more reliable means to discern groupings of OTU’s. Figure 11 graphically

represents the characters in relation to the second and third principal components.

The communalities (Table 5) indicate that most of the size characters are highly

intercorrelated, i.e., in total they explain most of the observed variability. This

is not surprising as 5 of the characters were continuous and displayed a wide range

of variation. In contrast, characters DISC, USGC and SUHW have relatively low

communalities, i.e., when combined with the other 5 characters, they explain a

lesser proportion of the observed variance in the wing characters. As these only

poorly correlate with any size characters, the PCA demonstrates that the three

non-parametric characters, DISC, USGC and SUHW, are best at distinguishing

the groups of OTU’s. These results may be somewhat biased due to the non-

parametric nature of qualitative characters. Nonetheless, other numerical taxo¬

nomic techniques confirm that characters DISC, USGC and SUHW can best

partition the OTU’s (Arnold, 1983, 1985).

Factor loadings of the first three principal components of each OTU were used

to compute factor score for the 35 OTU’s. Figure 12 depicts interrelationships of

OTU’s in the first three principal components and represents 83.2% of the variance

in the original data matrix. The remaining 17% of the variance might modify

positions of the OTU’s as they are depicted in Figure 12. Based primarily on size

characteristics, all green morphs (OTU’s #22, 29-35) cluster together near the

center of the plot. However, several morphs from the North Coast Range (OTU’s

#12, 14), South Coast Range (#2), Sierra Nevada (#16, 17) and Tehachapi Moun¬

tains (#23, 24, 26, 27) also lie in the same general area. Only the 3 S. callippe

juba OTU’s and the S. callippe laura OTU’s (#18, 19, 20, 21) form a discrete

cluster. S. callippe callippe, S. c. comstocki, and S. c. Uliana OTU’s (#1-12) are

bisected by the cluster of green-morph OTU’s. Thus the principal components

analysis corroborates the finding of the discriminant function analysis that overall

size is a useless character for distinguishing subspecies of Speyeria callippe. Qual¬

itative characters such as DISC, USGC, and SUHW can better partition individ¬

uals into populations than size characteristics.

Discussion

The description and explanation of variational patterns may be examined one

character at a time or with many characters simultaneously. The trend in geo-

PAN-PACIFIC ENTOMOLOGIST

graphic variation studies has been from univariate and bivariate analyses to multi¬

variate analyses (Gould and Johnston, 1972). Multivariate studies reveal not just

correlations between characters, but correlations among suites of characters within

the sampled populations.

My univariate and multivariate analyses of geographic variation, indicate that

many of the characters previously used to distinguish subspecies of S. callippe

are useless as taxonomic characters. This study, one on phenetic variation in non-

ordered state characters (Arnold, 1983), and another on the relationship of geo¬

graphic variation in morphological and climatic variables (Arnold, 1985), dem¬

onstrate that most of the previously recognized subspecies of callippe are invalid

based on the currently used characters. If subsets of OTU’s derived via the uni¬

variate analysis are homogeneous for the characters in question, and there is a

pattern to the variation, it should belong to geographically contiguous regions

(Sneath and Sokal, 1973). Those sets of localities that are statistically homogeneous

and geographically contiguous can be categorized as being biologically homoge¬

neous and recognized taxonomically as subspecies. The univariate analysis dem¬

onstrates that the variation in 7 of the 8 characters is discordant. The pattern of

variation in only one character, ventral disc color (DISC), correlates with geog¬

raphy and can be used as a basis for delimiting subspecies. Similarly, the multi¬

variate techniques corroborate that DISC is the best character for partitioning

populations, while the 5 continuous characters vary discordantly and the OTU’s

cannot be effectively grouped. DISC, USGC, and SUHW exhibit less variability,

and thus can more effectively partition the populations. One advantage of PCA

is that the relationships between the OTU’s can be viewed without prior clustering,

in this study, the grouping of populations into subspecies. Another important

feature is that the trends in variability can be related to the actual characters

which cause them (Moulton, 1973).

PCA has been used successfully in examining populations for phenetic inter¬

mediacy (Rising, 1968, 1970); however, since the technique does not maximize

differences between biologically meaningful reference samples, it is not as powerful

a tool for studies of hybrid or blend zones, or zones of abrupt phenetic transition

as is discriminant analysis (Rohwer, 1972). Indeed, the discriminant analysis

indicated that the previously recognized 16 subspecies were so poorly defined that

only 43.2% of the individuals of unknown subspecific identity could be correctly

identified based upon the same characters other lepidopterists have used to dis¬

tinguish these “subspecies.”

One of the major objections expressed by the opponents of the subspecies is

that the trinominal system forces biologists to make arbitrary decisions that fre¬

quently distort the real nature of character variation and bias subsequent analysis

(Hagmeier, 1958; Wilson and Brown, 1953). Also, most of the thorough studies

of geographic variation show that independent characters vary discordantly in

space (Gillham, 1956), in time (Doutt, 1955; Packard, 1967; Mason, 1964), and

that single-character variation tends to be clinal (Sibley, 1954). As the environ¬

ment exerts differing selective forces on differing characters one should expect

that discordant variation will occur more frequently than concordant variation.

The phenotypes thus assembled under the name of a subspecies might be so

heterogeneous that the subspecies concept loses nearly all meaning. Often the only

practical way to identify subspecies, such as those recognized by many butterfly

VOLUME 61, NUMBER 1

taxonomists, is by geographic location. Yet this may obscure the true patterns of

geographic variation in a species.

Statistical analyses of geographic variation have advantages over more subjec¬

tive methods. They are repeatable, knowledge obtained subsequently can be in¬

corporated, and are objective to the extent that they attempt to describe variation

before partitioning into subspecies. A thorough character analysis can demonstrate

that discontinuities are correlated with disjunct distributions or with abrupt changes

in the environment. These qualities are all too often absent in systematic procedure

at the infraspecific level.

Many subspecies have been based on too few characters, poorly chosen char¬

acters, or inadequate analysis of geographic variation of characters (Wilson and

Brown, 1953). Diagnostic markings and other slight external characters of neces¬

sity serve as prime criteria for distinguishing museum specimens. Nevertheless,

the objective of many describers of subspecies is to discern minute typological

differences in samples of populations from different localities rather than to study

the similarity of samples or variability within a population (Burt, 1954; Hubbell,

1954). The mere pigeonholing of populations at infraspecific levels, especially

when the range of variation has not been ascertained for several characters, con¬

veys a false impression of uniformity (Bogert, 1954). Subspecific names should

be employed to designate recognizably different assemblages of populations within

a species (Mayr, 1969).

Revision of the Subspecific Tax a of Speyeria callippe

Results of the several numerical taxonomic techniques employed in this study

and others (Arnold, 1983, 1985), provide a basis for making taxonomic deci¬

sions at the infraspecific level. The 16 currently recognized subspecies of Speyeria

callippe are reduced to 3 taxa: callippe callippe (Bdv.), callippe nevadensis (Ed¬

wards), and callippe semivirida (McDunnough).

Ventral disc color (DISC) is the best character for distinguishing the 3 subspecies.

All populations with brown ventral discs, no matter what shade of brown, should

be referred to as S. callippe callippe. These include silvered and unsilvered pop¬

ulations and those with several shades of brown scaling varying from reddish-

brown to chocolate brown in color, and ranging from the Mt. Shasta-Trinity Alps

region to San Diego, California.

Populations from southeastern Oregon, with greenish-brown or brownish-green

ventral discs are grouped as S. callippe semivirida. All green disced populations

from the east slope of the Sierra Nevada Mtns. and Great Basin are grouped as

S. callippe nevadensis.

Most characters previously used are too variable to diagnose subspecies. These

include general size, silver-spot pattern, spot color, dorsal melanization, dorsal

ground color, and ventral margin banding characters (Arnold 1983, 1985). The

revised taxonomy is as follows:

callippe callippe { Bdv.), 1852

c. comstocki (Gunder), 1925 New Synonym.

c. Uliana (H. Edwards), “1876” (1877) New Synonym.

c. elaine dos Passos and Grey, 1945 New Synonym.

c. rupestris (Behr), 1863 New Synonym.

PAN-PACIFIC ENTOMOLOGIST

HORIZONTAL FACTOR 2 vs. VERTICAL FACTOR 3

15 = d isc

Figure 11. Graphic representation of principal components numbers 2 and 3 based upon characters

FWLE, FWLW, FDLW, HWLW, HDCL, USGC, SUHW, and DISC. The underlying cause of hori¬

zontal factor number 2 is attributed to overall size, whereas vertical factor number 3 is largely due to

wing coloration characters.

c. juba (Bdv.), 1869 New Synonym.

c. laura (Edwards), 1879 New Synonym.

c. sierra dos Passos and Grey, 1945 New Synonym.

c. macaria (Edwards), 1877 New Synonym.

c. laurina (Wright), 1905 New Synonym.

callippe nevadensis (Edwards), “1870-71” (1870)

c. harmonia dos Passos and Grey, 1945 New Synonym.

c. meadii (Edwards), “1872-73” (1872) New Synonym.

c. gallatini (McDunnough), 1929 New Synonym.

c. calgariana (McDunnough), 1924 New Synonym.

callippe semivirida (McDunnough), 1924.

Implications for Conservation

The San Bruno Mountain (San Mateo County, California) population of Spey-

eria callippe callippe was proposed for recognition as an Endangered Species by

the U.S. Fish and Wildlife Service in 1978. Although the butterfly is no longer

known from its type locality, San Francisco, Howe (1975) believed that the name

Speyeria callippe callippe applied to the population at nearby San Bruno Moun¬

tain. Housing developments threaten to destroy a significant portion of the but¬

terfly’s habitat at this site. In 1978, the Endangered Species Act of 1973 could

legally recognize invertebrates as Endangered Species at the population, subspe¬

cies, or species level. However, the Tellico Dam-Snail Darter controversy resulted

VOLUME 61, NUMBER 1

Figure 12. Three-dimensional plot of 35 Speyeria callippe populations based on the first three

principal components. Labels PC 1, PC 2, and PC 3 on the axes refer to the first three principal

components. Populations are coded as in Table 1.

in several amendments to the Endangered Species Act, including nomination of

only formally named taxa, i.e., unnamed invertebrate populations could no longer

be proposed for protection. Considerable ecological study of San Bruno Mountain

S. callippe has been conducted since 1976 (Arnold, 1981; Reid and Arnold,

unpubl. data). Although this population of S. callippe is no longer considered a

distinct subspecies whose range is restricted to only San Bruno Mountain, it may

represent a unique ecotype, similar to the situation recently elucidated by Murphy

and Ehrlich (1980) for Euphydryas editha (Bdv.) in the San Francisco Bay Area.

However, comparable ecological studies of other Bay Area S. callippe populations

have not been completed.

Summary

Univariate and multivariate statistical analyses of geographic variation in 8

wing characters of Speyeria callippe revealed that only one character, ventral disc

PAN-PACIFIC ENTOMOLOGIST

color (DISC), can partition populations into meaningful groups. Other size and

color characters used to differentiate previously recognized subspecies do not

correlate with geography. These analyses provide a basis for revision of the sub¬

specific taxonomy of S. callippe. The 16 previously recognized subspecies are

reduced to three, whose morphology correlates with geography.

Acknowledgments

Several people lent me specimens used in this study: Dr. P. H. Amaud, Jr.

(California Academy of Sciences), J. DeBenedictis, Dr. R. W. Garrison, Dr. J. E.

Hafemik (San Francisco State University), S. O. Mattoon, and Dr. S. R. Sims

(University of California, Davis). My wife, Debbie, accompanied me on several

of the field trips to procure specimens and was an invaluable assistant. Drs. H.

Y. Daly, J. T. Doyen, T. Duncan, P. R. Ehrlich, and J. A. Powell reviewed earlier

drafts of this manuscript and agree with my conclusions. An anonymous reviewer

provided insightful comments that improved this paper. The Computer Center

and College of Natural Resources at U.C. Berkeley provided several computer

research grants. Dr. Duncan also provided a considerable amount of computer

time through his user’s group at U.C. Berkeley.

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PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 24-25

Aleiodes sp. (Hymenoptera: Braconidae) Reared from an

Anomalous Host, an Adult of the Eremochrysa punctinervis

Species Group (Neuroptera: Chrysopidae)

James B. Johnson

Division of Biological Control, Department of Entomological Sciences, Uni¬

versity of California, Berkeley, California 94720. Present address: Department of

Plant, Soil and Entomological Sciences, University of Idaho, Moscow, Idaho

83843.

During the course of a study concerning the bionomics of some symbiote-using

Chrysopidae (Insecta: Neuroptera) approximately 200 adults of a population be¬

longing to the Eremochrysa punctinervis species group were collected in Gates

Canyon, Solano Co., California. The adult chrysopids were attracted to mixed

ultra violet/visible lights between 23 April and 25 September during the summers

of 1978-1980.

The newly collected adults were housed in Dixie® unwaxed 8 oz squat containers

with clear plastic lids and were supplied with artificial honeydew (Johnson and

Hagen, 1981, Nature, 289:405-407) and water. Although the containers were free

of other insects and plant material, on 2 occasions solitary hymenopterous larvae

were observed in the cartons within 2 days of the collection date. Unfortunately,

only one was noticed before the dead adult chrysopids were removed from the

carton. In this case, a female was found with a collapsed abdomen and an emer¬

gence hole behind tergite VIII. This larva died, but the other larva matured. The

adult was identified as Aleiodes sp. (Hymenoptera: Braconidae). However, after

the subfamily Rogadinae is revised, this species will not remain in Aleiodes

(Shenefelt, pers. comm.).

The Rogadinae are known to parasitize only lepidopterous larvae (Krombein et

al., 1979, Cat. Hymenop. Amer. N. of Mex., Vol. 1). Therefore, this would be a

radical departure from the known host range of the subfamily. Still, it seems

certain that the adult chrysopids were the hosts since there were no other insects

in the cartons and the larvae were green, as would be expected if they had fed on

the chrysopids. In addition, other Braconidae are known to parasitize a wide

variety of insects and life stages, including adult Chrysopidae (Clancy, 1946, Univ.

Calif. Pubs. Ent., 7(13):403-496; Krombein et al., 1979, Cat. Hymenop. Amer.

N. of Mex., Vol. 1). So an adult chrysopid host would not be a major shift in the

context of the family as a whole. However, it seems probable that this was merely

a case of incidental parasitism of an atypical host. While the observed rate of

parasitism was approximately 1% for E. punctinervis adults, many, apparently

conspecific, adult wasps came to the mixed ultraviolet/visible lights. Therefore,

it may be that the wasp normally parasitizes some lepidopterous larvae, but

occasionally oviposits in chrysopid larvae. This could explain the apparent dis¬

crepancy between the low rate of parasitism observed and the abundance of adult

VOLUME 61, NUMBER 1

parasites, and the emergence from an adult host, which is unusual for the Ro-

gadinae.

I wish to thank Dr. P. A. Adams, Department of Biological Sciences, California

State University, Fullerton, California 92634, for identifying the Eremochrysa

and Dr. R. D. Shenefelt, 630 Oak St., Oregon, Wisconsin 53575, for identifying

the Aleiodes, plus his comments on the classification of the Rogadinae.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 26-31

Myrmecomorphic Miridae (Hemiptera) on Mistletoe:

Phoradendrepulus myrmecomorphus, n. gen., n. sp., and a

Redescription of Pilophoropsis brachypterus Poppius

Dan A. Polhemus and John T. Polhemus

(DAP) Department of Biology, University of Utah, Salt Lake City, Utah 84112;

(JTP) 3115 S. York Street, Englewood, Colorado 80110.

Recent collections in Arizona have revealed a diverse and bizarre array of

Miridae inhabiting desert mistletoe ( Phoradendron californicum). Several myr¬

mecomorphic forms are present, including a new genus and a species that has

previously been described under two different names for the different sexes. In

this paper, Phoradendrepulus myrmecomorphus, n. gen., n. sp. is described, Pil¬

ophoropsis balli Knight is placed as a junior synonym of Pilophoropsis brachyp¬

terus Poppius, and the female of Pilophoropsis brachypterus is redescribed.

All specimens treated herein are held in the Polhemus collection (JTP) unless

otherwise noted.

Phoradendrepulus , New Genus

Description.— Head broad, short, triangular as viewed from above; antennae

long, slender, apical segments slightly thickened. Pronotum large, raised, quadrate;

collar well defined; calli indistinct. Mesoscutum well exposed; scutellum sharply

elevated, conical. Hemelytra short, brachypterous, coriaceous, barely attaining

base of abdomen, posterior margins raised, rounded, clavus and cuneus indistin¬

guishable. Abdomen constricted basally, globose posteriorly with conspicuous

pleural fold. Legs long, slender; coxae and femora stout; length of tarsal segment

3 subequal to lengths of basal two combined; claws small, parempodia hair-like,

pul villi minute. Rostrum long, extending past hind coxae. Male genitalia of phy-

line-type; right clasper small, leaf-shaped (Fig. lb); left clasper larger, cup-shaped

(Fig. la). Overall body ant-like in form; both sexes brachypterous and very similar

in general facies.

Discussion.— On the basis of pretarsal structure and form of the genitalia, Phora¬

dendrepulus, n. gen. belongs in the subfamily Phylinae. It appears most closely

allied to Cyrtopeltocoris, in particular the head shape, conical scutellum, and ant¬

like habitus are extremely reminiscent of Cyrtopeltocoris females. Phoradendre¬

pulus, n. gen. may be separated immediately from other North American phylines

by its unique ant-like form and brachypterous condition in both sexes, and by its

large, quadrate pronotum.

Etymology .—The name Phoradendrepulus (m.) is derived from Phoradendron,

the host plant genus, and epulo, Latin for feaster, alluding to the association of

these bugs with mistletoe.

Phoradendrepulus myrmecomorphus , New Species

Description.—Brachypterous male: Small, ant-like; length 2.48 mm; width across

pronotum 0.76 mm; width across abdomen 1.12 mm.

VOLUME 61, NUMBER 1

Figure 1. Phoradendrepulus myrmecomorphus, n. gen., n. sp., male genitalia, a. Left paramere. b.

Right paramere.

Head dark brown, rugose; frons with transverse striae; a pair of stout setae

present anteriorly between eyes; clypeus thickly set with fine gold hairs; vertex

two times dorsal width of an eye, set with four stout dark setae; eyes reddish

brown, anterior margins well separated from antennal sockets. Antennae brown;

segment I short, stout, bearing three stout setae; segments II and III longer, slender;

all segments clothed with short recumbent golden pubescence; lengths of segments

I-IV (in mm): 0.24; 0.80; 0.64; missing.

Pronotum orange brown, raised, quadrate, posterior angles infuscated in some

specimens; surface rugose, covered with very fine, short, upright, golden pubes¬

cence; calli indistinct. Mesoscutum broadly exposed, sloping downward poste¬

riorly to scutellum; scutellum raised, conical, covered with fine gold hairs; a long

seta present on each side of conical apex. Hemelytra orange brown, brachypterous,

a small pallid area present centrally in some specimens; posterior margins up¬

turned, smoothly rounded, barely attaining base of abdomen; surface with fine,

recumbent gold hairs; several long setae present distad along hemelytral com¬

missure.

Abdomen black, polished, shining, globose, basal segments constricted; dorsal

surface with fine, recumbent gold hairs, intermixed with longer upright gold setae;

pleural fold prominent basally.

Ventral surface orange brown; rostrum light brown, set with very fine, erect,

short hairs; length 1.52 mm, reaching beyond hind coxae. Legs orange brown;

fore coxae red, pallid basally, middle and hind coxae pallid; femora and tibiae

clothed with fine recumbent gold hairs; tibiae set with stout spines; middle tibiae

with a row of evenly spaced, fine, erect hairs on inner face; tarsal segment 3 as

long as basal two segments combined. Venter of abdomen black, polished, set

with numerous short, recumbent gold hairs and scattered long, upright hairs.

Male genitalia of phyline-type (see Fig. la, b); right clasper small, trough-shaped;

left clasper larger, cup-shaped, with two spinose projections.

Brachypterous female: Generally similar to male in color and structure (see Fig.

2); stridulatory apparatus present. Length 2.88 mm; width across pronotum 0.80

mm; width across abdomen 1.28 mm. Lengths of antennal segments I-IV (in

mm): 0.28; 0.84; 0.56; 0.48. Hind femora with sclerotized ridge (plectrum) on

inside face of basal half, positioned to rub against microserrate costal margin of

PAN-PACIFIC ENTOMOLOGIST

hemelytra (strigil). Ovipositor sheath long, extending caudad two-thirds length of

abdomen to base of stemite 8.

Discussion.—Phoradendrepulus myrmecomorphus, n. sp. inhabits mistletoes

(Phoradendron californicum ) parasitizing mesquite ( Prosopis juliflora). The in¬

sects are invariably taken in the company of myrmicine ants ( Crematogaster sp.),

and this, along with an exceptional ant-like habitus that makes the insects quite

difficult to distinguish from ants in the field, suggests that the species is myrme-

VOLUME 61, NUMBER 1

cophilic. Another notable feature of this unusual mirid is a stridulatory mechanism

consisting of a raised ridge located basally on the hind femur (plectrum) positioned

to contact a striate costal margin on the hemelytra (strigil). The striations of the

costal margin are very fine and difficult to see, and appear to be present only on

the distal portion in the male.

The antennal proportions given in the description were taken from specimens

having four segments on each side, however an unusually high percentage (25%)

of the specimens in our small sample exhibit antennal oligomery. The antennal

proportions of the specimens exhibiting oligomery are as follows:

Antennal segment

Sex

Date collected

III

X-13-82

L&R

1.2

3.8

2.9

VIII-10-82

1.2

5.8

—

1.1

4.0

missing

In the associated series of Pilophoropsis brachypterus collected from the same

trees, only one specimen of 17 exhibited unilateral oligomery. Antennal oligomery

is not an unusual phenomenon in Heteroptera (Leston, 1952), however the un¬

usually high occurrence in Phoradendrepulus myrmecomorphus leads us to spec¬

ulate that the extremely restricted habitat and mobility of these flightless insects

has led to extensive inbreeding and further that this extensive inbreeding is the

causal factor in the oligomery. The less frequent occurrence in Pilophoropsis

brachypterus would consequently be explained by the greater interdeme gene flow

permitted by the mobility of the macropterous males of the latter species. Of the

dozens of mesquite trees sampled, only a few large trees had populations of these

two species of myrmecomorphic bugs, and these were in general separated by a

mile or more of harsh desert. Given the apparently restricted vagility of Phora¬

dendrepulus myrmecomorphus individuals, gene exchange between local popu¬

lations of this species is undoubtedly very infrequent.

Etymology.— Derived from the greek myrmex, ant, alluding to the ant-like

habitus.

Material examined.— Holotype, 6, and allotype, 2: ARIZONA, Maricopa Co.:

Ariz. Hwy. 87 at Verde River, Ft. McDowell Indian Res., VIII-10-82, J. T.

Polhemus (JTP). Paratypes: 2 2, same data as type; 2 2, same locality as types,

V-24-82; 1 <3, 1 2, same locality as types, X-13-82 (all JTP).

Pilophoropsis brachypterus Poppius

Pilophoropsis brachypterus Poppius, 1914, Ann. Soc. Entomol. Belg., 58:249 (n.

gen., p. 249; n. sp., p. 250).

Pilophoropsis balli Knight, 1968, Brig. Young Univ. Sci. Bull., 9:176. New syn¬

onymy.

Pilophoropsis brachypterus was originally described by Poppius from a bra-

chypterous female specimen taken at “Hot Springs,” Arizona. Knight, over 50

years later, examined several macropterous males from various localities in Ar¬

izona and described them under the name Pilophoropsis balli. One of us (JTP)

has studied the types of both species at the USNM. A long series taken from

PAN-PACIFIC ENTOMOLOGIST

i mm

Figure 3. Pilophoropsis brachypterus Poppius, adult female, dorsal view.

mistletoe, Phoradendron californicum, at Ft. McDowell, Arizona, near Phoenix,

now reveals that the two species are synonymous, with P. balli the junior synonym

of P. brachypterus. Knight’s 1968 description of the male, along with a figure of

its bizarre genitalia, need not be repeated here, but a redescription of the female,

in English, is provided to facilitate future identification of this unusual species.

The insect is apparently confined to the Sonoran desert; to date no macropterous

females have been found.

Brachypterous female: Small, ant-like (see Fig. 3); length 2.96 mm; width across

pronotum 0.56 mm; width across abdomen 1.00 mm.

Head brown, shining, concave along posterior margin; width 0.76 mm; eyes

small, not protrusive; vertex wide, equal to two times the dorsal width of an eye;

frons smooth, set with fine gold setae; lora prominent, expanded. Antennae brown,

segment I paler; segment I with a pair of stout setae; segments II-IV thickly set

VOLUME 61, NUMBER 1

with short, stiff setae; segments III and IV thicker than segment II, subequal in

thickness to segment I; lengths of segments I-IV (in mm): 0.20; 0.56; 0.36; 0.36.

Pronotum dark brown, shining, weakly campanulate, convex in side view; calli

large, tumid, glabrous, with broad longitudinal sulcus between; surface covered

with fine, short golden hairs. Scutellum dark brown, shining; anterior lobe up¬

turned, almost vertical; posterior lobe flat, with a band of silvery, scale-like hairs

running transversely across middle.

Hemelytra short, brachypterous, reaching only to posterior margin of tergite 4;

dorsal surface brown, mostly dull, finely rugose, sparsely set with long, erect,

bristle-like black setae; a transverse band of silvery, scale-like hairs across base

of clavus joins a similar band on scutellum; two small patches of silvery hairs

present anteriorly between costal and radial veins; another transverse band of

silvery hairs present distad on corium near apex of clavus, broadly interrupted

across clavus; clavus with three transverse bands of silvery hairs on apical half;

cuneus dark brown, polished, with a patch of fine gold hairs present on posterior

margin; membrane greatly reduced, dark brown, polished, inner margin rugulose.

Abdomen black, shining, rounded, sparsely set with fine, erect gold hairs; pleural

fold present basally.

Ventral surface dark brown; rostrum length 1.12 mm, reaching between middle

coxae; abdomen with fine recumbent gold hairs intermixed with longer, upright

gold hairs; ovipositor sheath extending posteriorly three-quarters the length of

segment 7. Legs brown; coxae, trochanters, and bases of femora pallid; set with

short golden hairs, femora with a few longer hairs on posterior faces; tibiae with

stout dark spines; terminal tarsal segment longer than preceding two taken to¬

gether.

Records.— ARIZONA, Pima Co.: Tucson, IX-22-20, 1 6 , E. D. Ball (USNM);

Rincon Mountains, 1056 m (3300'), IX-2-38, 1 <3, A. A. Nichol (USNM); Catalina

Springs, IV-15, 1 <3, E. A. Schwarz (USNM). Maricopa Co.: Ft. McDowell Ind.

Res., Ariz. Hwy. 87 at Verde River, CL 1632, VI-2-81, 2 <3, 1 nymph; VIII-10-

82, 2 <3, 4 2, 1 nymph; X-13-82, 1 <3, 8 2; all J. T. Polhemus (JTP).

Acknowledgments

We wish to thank Thomas J. Henry, of the U.S. National Museum, Washington,

D.C., and Randall T. Schuh, of the American Museum of Natural History, New

York, for their help and advice in the preparation of this manuscript.

Literature Cited

Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the western United

States. Brigham Young Univ. Sci. Bull., 9(3): 1-282.

Leston, D. 1952. Antennal oligomery in Heteroptera. Nature, 169:890.

Poppius, B. 1914. Ubersicht der Pilophorus -Arten nebst beschreibung verwandter Gattungen (Hem.

Het.). Ann. Soc. Entomol. Belg., 58:237-254.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 32-37

The Life History of Ophraella notulata (F.) on Western Ragweed,

Ambrosia psilostachya De Candolle, in Southern California

(Coleoptera: Chrysomelidae)

R. D. Goeden and D. W. Ricker

Department of Entomology, University of California, Riverside, California

92521.

Despite an extensive and intensive survey of the phytophagous insect fauna of

western ragweed, Ambrosia psilostachya, in southern California during 1968—

1970, we failed to detect and hence did not include Ophraella notulata among

the 130+ insect species reported from this widespread native plant (Goeden and

Ricker, 1976c). However, while sweeping plants along Kitchen Creek at the south¬

ern end of the Laguna Mountains in San Diego Co. on October 17, 1979, we

encountered this beetle on western ragweed from which it is now reported, pre¬

sumably for the first time from California (Wilcox, 1965; Welch, 1978).

Distribution and host plants. — Wilcox (1965) listed O. notulata in North Amer¬

ica from as far west as Arizona. Welch (1978) described its distribution as “. . .

throughout most of the continental United States, parts of Canada . . . , and as

far south as Guatemala ...” exclusively on so-called “common ragweed,” A.

artemisiifolia (F.). He noted the apparent absence of common ragweed and O.

notulata from southern California, the latter supposition based on our still valid,

negative collection records from several other native ragweeds (Goeden and Rick¬

er, 1974, 1976b). We also have collected eggs, larvae, and adults of O. notulata

at Kitchen Creek from cocklebur, Xanthium strumarium L., a close ragweed

relative, from which this leaf beetle is newly reported (Hilgendorf and Goeden,

1982, 1983). In the insectary, O. notulata additionally has been reared from egg

to adult on potted plants of A. chenopodiifolia (Bentham) Payne, A. confertiflora

De Candolle, A. dumosa (Gray) Payne, A. eriocentra (Gray) Payne, and A. ilicifolia

(Gray) Payne. Apparently, none of these ragweeds is attacked by O. notulata in

nature (Goeden and Ricker, 1975, 1976a, 1976b).

Biology. — The biology of O. notulata was studied in the field on western ragweed

at Kitchen Creek during 1979-1982, and in the insectary of the Division of

Biological Control, University of California, Riverside, at 27 ± 1°C, 40-70%

relative humidity, and a 16/8-hr (light/dark) photoperiod. This supplements the

study by Welch (1978) of O. notulata on common ragweed in Connecticut.

Egg.— Welch (1978) described the pyriform eggs (Fig. la) and illustrated the

microscopic, hexagonal sculpturing of the chorion. Fifty eggs from our insectary

culture measured 0.05 mm smaller in mean greatest width, but otherwise fit his

description.

In insectary cagings, most fertile eggs were lightly, but firmly glued at their

larger ends in clusters to the leaves of potted plants or bouquets of freshly excised

branches. A few (<1%) infertile eggs also were found within these egg masses.

Infertile eggs produced by unmated females and by older females towards the end

of their oviposition periods were loosely scattered individually or in small, irreg-

VOLUME 61, NUMBER 1

ular clusters on the foliage and cage surfaces. Of 456 masses of fertile eggs ex¬

amined, 296 (64.9%) were compact, elongate clusters (Fig. la); 91 (20.0%) were

linear, single or double-ranked series; and 69 (15.1%) were loose, irregular clusters.

Of 654 egg masses examined, 590 (90.2%) were laid on the undersides of leaves;

the remainder, on the adaxial leaf surfaces. Many of the latter clusters were laid

on leaves bent sharply upward or downward from their usual, nearly horizontal

positions. Of 679 egg clusters examined, 212 (32.3%) were attached to a lateral

lobe of a leaf alongside or straddled a primary vein, 143 (21.1%) were deposited

on the terminal leaf lobe and commonly straddled the midrib, 57 (8.4%) were

located in the angle of the midrib and a primary lateral vein, 44 (6.5%) were laid

on a basal leaf lobe, and 4 (0.6%) were attached to a petiole wing. A simple

experiment indicated that this egg placement partly was a response to gravity, as

all 22 egg clusters laid on the leaves of 5 straight branches held vertically, but

reversed 180°, were attached to the upside-down, adaxial leaf surfaces.

An average of 13.3 ± 0.2 (±SE) (range: 2-41) eggs were counted in 833 clusters

recovered from insectary cagings. The distribution of eggs per mass was skewed

such that only 41 (ca. 5%) of these clusters contained 23 or more eggs. Few fertile

eggs were laid singly. Welch (1978) reported much larger masses averaging 36

eggs in his cagings.

Field observations on egg mass placement and size supported our insectary

findings. At Kitchen Creek, 21 egg masses were found attached to the underside

of cauline leaves located an average of 18 ± 1.7 (range: 5-35) cm below the apical,

staminate inflorescences and 13 ± 1.6 (range: 0-30) cm above the lowest, living

leaves on mature plants.

Larva .—Eggs hatched in 5 or 6 days in the insectary. The first instars emerged

headfirst through irregular holes chewed in the chorions just below and lateral of

the apical papillae (Fig. lb). Larvae from the same egg mass usually hatched

within a few hours of each other. Some masses apparently consisted of eggs laid

by the same or different females on successive days. Embryonic development and

hatching were delayed for a distinct segment of the eggs in such masses, which

probably were artifacts of insectary culture. Eclosion took as little as 10 min. The

empty chorion was abandoned and leaf feeding began as soon as 1 min after

eclosion.

Welch (1978) reported head capsule measurements for each of the 3 instars. In

insectary rearings of 21 isolated larvae on bouquets of fresh branch terminals,

larval development lasted 12 ± 0.6 (range: 12-14) days. An average of 3.5 ± 0.1

(range: 3-4) days (ca. 30%) at the end of this period was spent as a nonfeeding

prepupa in a coarse silk cocoon. This prepupal period averaged a day longer than

Welch (1978) reported. On western ragweed, the first larval stadium lasted 4 days;

the second stadium, 2-3 days; and the third stadium, 5-7 days. The larvae grew

in length from 1.0-2.4 mm as first instars to 3.5-7.0 mm as third instars.

Feeding symptoms of the first instars were small “shot-hole” lesions in the

nearer epidermis and mesophyll to, but not through, the opposite epidermis. This

instar fed somewhat gregariously for a day or so, then began to disperse both

upward and downward on the stems to feed as scattered individuals or in small

groups. Molting, like feeding, usually occurred on the underside of a leaf, although

some first instars fed and molted atop leaves. Second instars fed as scattered

individuals and usually molted on the underside of leaves. Third instars skele-

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Life stages of Ophraella notulata. (a) Egg mass, 13 x. (b) Newly hatched larvae, 17.3 x.

6.7 x. (e) Gravid female, 8 x. (f) Adults in copula, 6.8 x.

tonized the leaves from the upper and lower surfaces and left the midribs, primary

lateral veins, and opposite epidermis largely intact (Fig. lc). Dark brown-black,

subcylindrical, fecal pellets littered the foliage upon which the larvae fed, the

foliage beneath, as well as the cage floors. Unlike Welch (1978), we observed no

tendency for the larvae to migrate at dusk toward the terminal growth of potted

plants caged either in the insectary or in a sunlit greenhouse.

The fully grown third instar usually began construction of its cocoon away from

feeding areas on the upper surface of a leaf along and over a midrib or a lateral

vein. Cocoon formation was initiated at various times of the day and night and

continued to completion during both the photophase and scotophase. The first

step in cocoon formation was the stringing of several, transverse, silk strands

VOLUME 61, NUMBER 1

between the leaf or leaf lobe margins across the larval dorsum. Contrary to Welch

(1978), who reported the silk to be a “maxillary secretion,” we found it to issue

as a clear, viscous liquid from an opening on the prostemum between the fore

coxae. The gland orifice is a narrow, circular, sclerotized ring that appears two¬

lipped when closed. The head of the larva periodically dipped postero-ventrally

as it secured additional mouthfuls of this secretion. The thoracic sterna and coxae

shone from a coating of the secretion. The freshly formed, clear, silk strands

darkened to golden yellow, then to reddish brown. After several transverse strands

were laid, the process began of joining them with other strands and connecting

these to the leaf surface and margins. The strands from which the cocoon was

fashioned were irregular in length and diameter and formed an open meshwork

through which the contents could be seen (Fig. Id). Strands were thickened (1)

by running a single strand between the mouthparts while depositing a coating of

the secretion, (2) by coating in a similar manner 2 or more strands held together

with the mouthparts, and (3) by incorporating plant hairs broken or chewed from

the leaf surface into the strand coating. The fore legs and mouthparts were used

to manipulate the strands while spinning. The surface of the leaf upon which the

cocoon was formed was kept relatively free of silk and rubbed smooth by the

body movements of the larva. The larva moved actively about within the cocoon

under construction by adhering to the leaf surface with its posterior, ventral

“proleg” or by grasping the webbing from inside with its middle and hind legs.

Turning itself end-over-end, the larva arched its dorsum upward against the

forming cocoon and thus used its body as a template to shape this structure. The

larva rested periodically during cocoon formation and these resting periods be¬

came longer as the cocoon neared completion. The finished cocoon (Fig. 1 d) had

a flattened teardrop shape inside that was roughly twice the width and one and

a half the height of the larva. Externally, the cocoon lay appressed to the leaf

surface and tapered at one end to a semicircular opening; the other end was

rounded and closed inside, but flared outward externally. One larva was observed

to take 7Vi hr to complete its cocoon.

Pupa. — Prior to pupation the larva partially backed out through the open mouth

of its cocoon and voided a pile of dark, liquid feces on the leaf surface. Molting

subsequently occurred inside the cocoon, where the cast exuviae usually was

deposited behind the exarate pupa at the inner mouth of the cocoon. The pupal

stage of 19 individuals averaged 4 ± 0.1 (range: 3-4) days in the insectary.

Adult .—The adult (Fig. le, f) was described in detail by Horn (1983). Welch

(1978) illustrated an adult in dorsal view and the terminal, abdominal, sternal

characters useful in separating the sexes.

Beetles emerged from cocoons through holes usually chewed in the end opposite

the cocoon opening, but occasionally exited through the top or at the same end

as the opening. Like Welch (1978), we noted that adults emerged only during the

photophase. Depending on the generation involved, the emerged adults feed on

leaves, mate, and either reproduce or overwinter as discussed below. Also like

Welch (1978), we noted that mating could occur within 1 or 2 days after beetle

emergence in the insectary.

All of 6 females individually reared from eggs in the insectary initially oviposited

5 days after they had molted to adults within their cocoons. Each of 21 mating

pairs collected at Kitchen Creek on October 17, 1979, were caged separately with

PAN-PACIFIC ENTOMOLOGIST

an excised branch of western ragweed which was checked for eggs and replaced

daily during the oviposition period of each female. These beetles normally would

have overwintered and not have reproduced until the following summer; however,

after feeding under insectary conditions, the females began to oviposit in an

average of 9 ± 1 (range: 3-21) days. Fourteen of the 21 females that subsequently

were not accidentally killed or escaped laid an average of 667 ± 60 (range: 385—

1206) eggs during a 66 ± 6 (range: 32-106) day oviposition period. Based only

on the days when they oviposited, an average of 45 ± 4 (range: 25-67) days each,

these 14 beetles laid 15 ± 0.4 (range: 1-57) eggs daily. These females were observed

in copula (Fig. If) an average of 5 (range: 3-11) times and lived an average of 9

(range: 1-14) days after they ceased oviposition. The 16 males that did not die

prematurely or escape lived 151 ± 14 (range: 83-260) days after they were col¬

lected; the 14 females that oviposited lived 83 ± 7 (range: 42-123) days after

they were collected.

Seasonal history. —Ophraella notulata is multivoltine in southern California.

The beetles overwinter as mated adults, which Welch (1978) also reported from

Connecticut. The earliest date that we swept beetles from vegetative shoots of

western ragweed at Kitchen Creek was June 17. The leaf beetle was readily cultured

in glass-topped sleeve cages as described by Gilstrap and Goeden (1974) on potted

western ragweed, which itself is readily propagated vegetatively. Under insectary

conditions, O. notulata completed a generation each month under continual cul¬

ture for 27 generations. Ophraella notulata probably produces at least 3 genera¬

tions annually at Kitchen Creek. The last generation, developing on mature plants

in October and early November, was most numerous. Distribution of immatures

and adults was patchy in 1981, but quite uniform in 1982, judging from sweep

net samples along the margins of Kitchen Creek.

Mortality factors.— No parasites were reared from egg masses collected in the

field. Egg predation by a large, red mite (Acarina: Anystidae) was observed. Partly

and wholly collapsed eggs were commonly collected at Kitchen Creek. Larvae

were fed upon by nymphs and adults of Perillus splendidus (Uhler) (Hemiptera-

Heteroptera: Pentatomidae) in the field. Chaetonodexodes vanderwulpi (Town¬

send) (Diptera: Tachinidae) was reared as a solitary, endoparasite from prepupae

collected in cocoons.

Acknowledgments

Ophraella notulata was identified by Dr. T. N. Seeno, Insect Taxonomy Lab¬

oratory, California Department of Food and Agriculture, Scaramento. Dr. J. A.

McMurtry, Department of Entomology, University of California, Riverside, iden¬

tified the anystid mite. Dr. D. Wilder, Systematic Entomology Laboratory, IIBIII,

% U.S. National Museum of Natural History, Washington, D.C., identified the

“very interesting and uncommon tachinid.”

Literature Cited

Gilstrap, F. E., and R. D. Goeden. 1974. Biology of Tarachidia candefacta, a Nearctic noctuid

introduced into the U.S.S.R. for ragweed control. Ann. Entomol. Soc. Am., 67:265-270.

Goeden, R. D., and D. W. Ricker. 1974. The phytophagous insect fauna of the ragweed, Ambrosia

acanthicarpa, in southern California. Environ. Entomol., 3:827-834.

-, and-. 1975. The phytophagous insect fauna of the ragweed, Ambrosia confertiflora,

in southern California. Environ Entomol., 4:301-306.

VOLUME 61, NUMBER 1

-, and-. 1976a. The phytophagous insect fauna of the ragweed, Ambrosia dumosa, in

southern California. Environ. Entomol., 5:45-50.

-, and-. 1976b. The phytophagous insect faunas of the ragweeds, Ambrosia chenopo-

diifolia, A. eriocentra, and A. ilicifolia, in southern California. Environ. Entomol., 5:923-930.

-, and-. 1976c. The phytophagous insect fauna of the ragweed, Ambrosia psilostachya,

in southern California. Environ. Entomol., 5:1169-1177.

Hilgendorf, J. A., and R. D. Goeden. 1982. Phytophagous insects reported worldwide from the

noxious weeds, spiny clotbur, Xanthium spinosum, and cocklebur, X. strumarium. Bull. Ento¬

mol. Soc. Am., 28:147-152.

-, and-. 1983. Phytophagous insect faunas of spiny clotbur, Xanthium spinosum, and

cocklebur, Xanthium strumarium, in southern California. Environ. Entomol., 12:In press.

Horn, G. H. 1893. The Galerucini of boreal America. Trans. Am. Entomol. Soc., 20:57-144.

Welch, K. A. 1978. Biology of Ophraella notulata (Coleoptera: Chrysomelidae). Ann. Entomol. Soc.

Am., 71:134-136.

Wilcox, J. A. 1965. A synopsis of the North American Galerucinae (Coleoptera: Chrysomelidae).

N.Y. State Museum Sci. Ser. Bull., 400:1-226.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 38-39

Scientific Note

Description of the Previously Unknown Male of Ophrynon levigatus

Middlekauff (Hymenoptera: Symphyta, Orussidae)

Recently Dr. Henry A. Hespenheide 1 sent me two Orussidae for identification.

These had been collected at different times and localities in connection with his

studies of the ecology and taxonomy of Buprestidae. One submitted specimen, a

male, is the previously unknown opposite sex to the recently described unique

female, the holotype of Ophrynon levigatus Middlekauff. The second orussid

specimen was an unrelated new genus and species from Panama.

The male specimen emerged from wood of Quercus dumosa Nutt, collected by

D. S. Verity, April 2, 1978 at 3500' elevation in the Santa Rosa Mountains,

Riverside County, California. The scrub oak wood was being held in a cage for

insect emergence, and the following Buprestidae came from the same collection

which produced the male levigatus : Acmaeodera knullorum Barr; A. linsleyi Ca-

zier; A. vulturi Knull; Acmaeoderopsis guttifera (Le Conte); and Hesperorhipis

jacumbae Knull. Verity thought that due to their larger size, one of the Acmaeodera

would most likely be the host of the orussid.

The holotype female was described (Middlekauff, 1984) from a specimen col¬

lected April 3, 1956 at Brawley, Imperial Co., California.

Ophrynon levigatus Middlekauff

Plesioallotype male.— Length 5 mm. Head, antennae, and abdomen black. Knees

(apices of femora), outer faces of tibiae (Fig. 5), and a spot on lower apical portion

of stigma whitish (Fig. 1). Basal two-thirds of femora, and undersides of tibiae

dark amber. Tarsi somewhat lighter amber. Fore wing clear, lacking a pale band

below the stigma. Venation as in Figure 1. Apical veins of the wing beyond M

(forming base of discoidal cell) and a cu-a clear, hardly discernible. Cross vein 2

r-m appears to be absent, but in good light is seen to extend but half way across

the base of cell 2R S . Veins in the hind wing, Figure 2, except for the base of R,

are, if present, so faint as to be indiscernible.

Facial carinae not as pronounced as in the female, but present and enclosing

the frons (Fig. 4). A low, faint carina bisects the frons as shown in the female.

Each shallow pit on the face bears a single, small seta and the area between the

facial pits is smooth and shiny. Postocular carina and surrounding area similar

to female. Legs similar. Scutellum with similar pits, but lacks the anterior notch.

Hind femur swollen (Fig. 5) slightly more than 2.5 times longer than broad.

Hypopygium as shown in Figure 3.

The specimen is in the collection of the California Academy of Sciences, San

Francisco, California.

1 Department of Biology, University of California, Los Angeles, California.

VOLUME 61, NUMBER 1

Figures 1-5. Ophrynon levigatus Middlekauff, plesioallotype, <3. Fig. 1. Forewing. Fig. 2. Hindwing.

Fig. 3. Hypopygium, ventral view. Fig. 4. Head, frontal view. Fig. 5. Hind femur and tibia, lateral

view.

Literature Cited

Middlekauff, Woodrow W. 1983. A revision of the sawfly family Orussidae for North and Central

America (Hymenoptera: Symphyta, Orussidae). Univ. Calif. Publ. Ent., 101:1-46.

Woodrow W. Middlekauff, Department of Entomology, University of Califor¬

nia, Berkeley, California 94720.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 40-41

Scientific Note

Occurrence of the Cotoneaster Webworm, Athrips rancidella, in

California (Lepidoptera: Gelechiidae)

The Palearctic moth, Athrips rancidella (Herrich-SchaefFer), is widespread in

Central and South Europe and occurs in Turkmenia, S.S.R., presuming the syn¬

onymy to be correct (Sattler, 1968, Dt. Ent. Zeit., N.F., 15:115; 1978, ibid., 25:

59). Its larvae feed on Rosaceae: Cerasus, Cotoneaster, and Prunus, according to

records for its synonyms (Spuler, 1913, Kleinschmett. Europas, 361; Schutze,

1931, Biol. Kleinschmett., Verlag Int. Ent. Yerein, 116; Busck, 1934, Proc.

Ent. Soc. Wash., 36:82; Kuznetsov, 1960, Akad. Nauk SSR, Trudy, 27:34). This

species was discovered at Portland and Eugene, Oregon, in 1929, and had been

recorded at several sites in northern Oregon and at Vancouver, Washington, by

1934 (Roaf et al., 1937, J. Econ. Ent., 30:134). In the Pacific Northwest, larvae

were found on Cotoneaster horizontalis, a deciduous, ornamental shrub that orig¬

inated from China; and the adventive population was described as Cremona

cotoneastri by Busck (1934, loc. cit.), in the belief that it was an undescribed

species introduced from the Orient. Subsequently this moth has been collected

in coastal British Columbia, Washington, and Oregon (USNM records; Hodges,

in litt.).

Larvae of A. rancidella were found in Berkeley, California, on Cotoneaster

congesta, by L. E. Casher and R. F. Gill, students in our Immature Insects course,

in early May 1983. C. congesta, which is sometimes known as C. microphylla

var. glacialis, is a low-growing Himalayan shrub with dense foliage, superficially

resembling C. microphylla, and both are popular evergreen ornamentals. Another

collection of larvae and pupae, made May 13, produced adults from May 17 to

June 23 (JAP 83E50). Subsequent surveys of several other species of Cotoneaster

in the vicinity did not reveal presence of the conspicuous larval damage. C.

congesta is not a commonly used ornamental in Berkeley; C. lactea {=Parneyi)

and other slender, arching species with sparse, tomentose leaves are much more

abundant, at times weedy, but evidently are not selected by A. rancidella.

According to Roaf et al. (1937, loc. cit.) the insect is univoltine, and young

larvae spend the winter in silken hibemacula on the branches. In Berkeley, most

of the feeding occurred in spring, when the black larvae covered the branches

with dense silken webs, from which they foraged, encompassing the entire foliage.

Feeding primarily was skeletonizing of the leaf surfaces, so that the whole shrub

became a caked mass of silk, frass, and brown leaves. By late July the plant showed

evidence of recovery, with green foliage distally, and no larvae were discovered

feeding on the new growth.

In 1984, another colony was discovered in Berkeley, by Ward Russell and W.

W. MiddlekaufF. Larvae caused similarly conspicuous damage on the same or¬

namental shrub, and moths emerged in late May and the first week of June (JAP

84E126). The second site is about 1.0 airline km east of the first, at a higher

elevation.

I thank R. A. Beatty, Dept. Landscape Architecture, U.C. Berkeley, for infor¬

mation on the cotoneasters, and R. W. Hodges, Systematic Entomology Labo-

VOLUME 61, NUMBER 1

ratory, U.S. Department of Agriculture, Washington, D.C., for identification of

the moth and literature references.

J. A. Powell, Department of Entomological Sciences, University of California,

Berkeley, California 94720.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 42—47

Life History and Biology of Pyrausta orphisalis Walker

(Lepidoptera: Pyralidae) on Mint in Washington 1

C. L. Campbell and K. S. Pike

Department of Entomology, Washington State University, Irrigated Agriculture

Research and Extension Center, Prosser, Washington 99350.

Abstract.—Pyrausta orphisalis Walker is a foliage feeder of spearmint and pep¬

permint in Washington during its larval stages. Three generations occur per year,

with typically five larval instars per generation. The life history of each generation

is illustrated and various biological data are provided, viz., information on flight

behavior, mating, adult longevity, ovipositional behavior, fecundity, immature

development, larval habitat, feeding damage, overwintering, and natural enemies.

Pyrausta orphisalis Walker is a small (wing span, 14-16 mm), distinctive, orange

and brown day-flying moth, which according to Munroe (1976) occurs along

Canada’s southern tier from Newfoundland to the Fraser Valley of British Co¬

lumbia, south to northern Florida, the Sierra Blanca of New Mexico, Sonoma

and Modoc counties, California. The moth frequents commercial mints [Scotch

spearmint {Mentha spicata ), native spearmint (M. cardiaca), and peppermint {M.

piperita )] for nectar and oviposition, and at times, may be observed by the hundreds

in flight in the field. Population levels will vary from year to year and between

areas in Washington. The larvae feed on mint foliage and have been considered

economically important on occasion by growers.

Descriptions of the life stages of P. orphisalis have been reported previously

(Walker, 1859; Munroe, 1976; Campbell and Pike, 1984), but little information

has been published on the life history or biology of the insect. Frick (1961)

mentioned that there appear to be three generations a year, that the larvae feed

on the terminal growth of mint, and that adults are sometimes extremely numerous

in August. The purpose of our study was to document the life history and basic

biology of P. orphisalis. Our principal objectives were to determine 1) the number

of generations per year and seasonal occurrence of each, 2) mating behavior,

oviposition, and fecundity, 3) larval behavior and feeding damage to mint, 4) the

developmental times for immatures, 5) the overwintering stage, 6) adult flight

behavior and longevity, and 7) the natural enemies of P. orphisalis. Voucher

specimens are on deposit at the Washington State University, Irrigated Agriculture

Research and Extension Center, Prosser, Washington.

Materials and Methods

Field and laboratory studies were conducted over a 2-year period (1981-1982)

at the Washington State University, Irrigated Agriculture Research and Extension

1 Scientific Paper No. 6491, Washington State University, College of Agriculture Research Center,

Pullman. Project 4412. This research was made possible in part by funds provided by the Washington

Mint Commission and the Mint Industry Research Council.

VOLUME 61, NUMBER 1

Center, Prosser. Where practical, observations and studies of P. orphisalis were

carried out on mint under cultivated field conditions. It was often necessary,

however, to observe the same individuals over time, which required the use of

cages or containers to confine the insect. For the field, cylinder cages (15-cm

diameter, 30-cm height), constructed of cellulose-nitrate plastic and vented at the

top with saran screening (400-jum mesh), were employed unless stated otherwise;

for the laboratory, clear plastic (5-cm diameter, 2-cm height) or glass (5-cm di¬

ameter, 2-cm height) petri dishes or cylinder cages (15-cm diameter, 30-cm height,

2.5 x 2.5-cm wire mesh covered with plastic wrap) were used. All adults and

immatures observed in the laboratory were held or cultured under a light and

temperature regime of 16/8 hr (light/dark) and 23-28°C/20.5°C, respectively.

The general life history and habits of P. orphisalis, including the occurrence in

time of its different life stages and number of generations per year, were determined

on the basis of near-daily (May to September) or weekly (April, September to

mid-November) field observations and sampling of spearmint and peppermint.

Mint cuttings, sampled randomly from the field, were usually viewed with a hand

lens or binocular microscope to detect eggs and distinguish first and second larval

instars; third, fourth, and fifth instars, prepupae, pupae, and adults were discern¬

ible without magnification. The overwintering stage of P. orphisalis was deter¬

mined by periodic inspection (October through February) of caged mint, stocked

heavily in late August with second generation adults (100+/cage, two cages used,

each 1.0 x 0.5 x 0.5 m).

Mating, ovipositional period, and fecundity were studied using virgin females

paired with males held on mint under laboratory cage, or on mint leaves in petri

dishes. Fecundity was also assessed on the basis of eggs dissected from virgin

females, reared from collected ultimate instars previously held for ca. 1 week on

caged mint in the field. Egg placement on the plant was determined by field

observation, and systematic sampling and evaluation of mint cuttings. Egg eclo-

sion and mortality were evaluated from eggs on mint in the field and the laboratory.

The developmental times for immatures were determined mainly from labo¬

ratory studies. The feeding sites and habits of each instar were determined through

field observation, as were data on pupation.

Parasitoids were reared from host larvae in the laboratory and subsequently

identified by specialists (Table 1). Predator information was derived from field

observations.

Results and Discussion

There were three generations of Pyrausta orphisalis that developed annually in

Washington. The life history of each is illustrated in Figure 1. The overall gen¬

eration times (in terms of length) were quite similar for the first two generations,

but substantially longer for the third. Similarly, field developmental times, in

terms of equivalent immature life stages for the first and second generations, as

well as the egg, first, second, and third larval instars of the third generation were

quite comparable; the fourth and fifth larval instars and prepupa of the third

generation, however, required more time to develop than their earlier equivalents,

perhaps due to cooler seasonal temperatures. There was overlap in life stages

within and between generations, particularly during the summer (Fig. 1). None¬

theless, there were peak periods of abundance for each life stage in each generation,

PAN-PACIFIC ENTOMOLOGIST

Table 1. Parasitoids of Pyrausta orphisalis Walker in Washington reared from field-collected host

larvae, Prosser, 1981-1982. 1

P. orphisalis

instar

No.

reared

No.

parasitized

% para¬

sitized

spinator 1

% of total parasitization

Cotesia M. cam- S.

sp . 3 pestris 3 bilineatus 2

Unknown

tachinid

sp . 4

100

—

100

—

III

191

—

218

—

384

Total or x

924

115

1 Identification of Ichneumonidae by Drs. J. R. Barron and J. E. H. Martin, Biosystematic Res.

Instit., Agric. Canada, Ottawa, Ont., Canada; Braconidae by Dr. P. M. Marsh, Systematic Entomology

Lab., USDA, Beltsville, Md.

2 Hymenoptera: Ichneumonidae.

3 Hymenoptera: Braconidae.

4 Diptera: Tachinidae.

e.g., first generation, first instar larvae in 1982 were most prevalent during the

first week in June, though some were present before and after that date. Over¬

wintering by the third generation prepupa began in October or November, fol¬

lowed by pupation in April or May, and finally adult emergence in May.

The moth is diurnal; white/blacklight traps operated nightly (concurrent with

peak daytime flight periods) yielded negative results. Striking orange and brown

coloration made the moth distinctive in the field coupled with its characteristic

short, often jerky, rapid flight. The moth seldom ventured far from mint fields,

usually staying just above or within the mint’s upper canopy. Between flights,

short resting periods by the moth commonly occurred on the undersides of mint

leaves. Sound perception in adults was acute; a camera’s shutter click was sufficient

to cause resting moths to take flight. Both sexes of the first two generations took

nectar from mint flowers during daylight hours.

Mating was observed only in the afternoon and at dusk in the field. Periodically,

large field aggregations of males (up to 100 estimated) were observed in flight

around a female, or in flight near a mating pair usually hidden in mint foliage,

suggesting the female released or used a sex pheromone to lure males. Males in

aggregations, were observed with external genitalia extended. In the laboratory,

once a male and female were paired, mating occurred within a period of a few

hours to about 3 days, preceded by presumed calling of the female resting upside

down from a plant or container and with its antovipositor extended. Copulation,

which can last for nearly an hour, was initiated immediately upon approach by

a male to a stationary female. During union, moths faced in opposite directions

and generally moved about short distances. Mating occurred during the same day

as emergence or anytime within the first 6 days following emergence.

Initial oviposition commenced generally two days after mating and lasted from

1 to 6 days, x = 3 (n = 8). Fecundity, based on laboratory studies, was extremely

variable ranging from 1 to 196 eggs, x = 115 (n = 9). We also observed that

unmated females deposited a few eggs. Dissected virgin females contained 17 to

75 mature eggs, x = 35.4 (n = 30).

VOLUME 61, NUMBER 1

20 30 10 20 30 10 20 30 10 20 30 10 20 30

AUG SEPT OCT NOV MAY

Figure 1. Life history and generations of Pyrausta orphisalis Walker in central Washington, 1981-

1982 [n (2-year, 3-generation totals) = eggs, 719; larvae, 1717; adults, 3063],

Females lit frequently to oviposit in the field, usually depositing eggs singly,

but occasionally in clusters of two to three. If four or more eggs occurred at one

site on the plant, it was due to separate visits by the same or different moths.

Females typically probed prospective sites with their antovipositors prior to egg

deposition. Eggs (n = 719) were usually laid on leaf buds, small leaves subtending

the buds, among flowers, and less frequently, on larger terminal leaves or stems.

Interestingly, egg mortality appeared to be related to ovipositional site; egg mor¬

tality on larger leaves and stems reached ca. 60% (n = 128), while mortality for

those among small terminal leaves and flowers was only 5% (n = 509). The ap¬

pearance of unhatched eggs included partial clearing, collapse, color change to

yellow or orange, no development, or partial development but without eclosion.

In the laboratory, eggs hatched in ca. 4 days. The stemmata of the pharate larva

became visible after 3 days and the head capsule darkened on the fourth.

Developmental time from egg to adult was temperature dependent; for larvae

(n = 70) reared in cohorts at 28°C/20.5°C (16/8 hr, light/dark) in the laboratory,

the mean was 32 days. For individually cultured larvae (n= 17) at 23°C/20.5°C

(16/8 hr, light/dark), the mean was 38.5 days. Under July-August field conditions,

the average total time was 34 days (n = 70). There were five larval instars in each

generation. Based on laboratory studies, the stadia of the first four instars were

ca. 4 (n = 50), 3.5 ± 0.7 {n = 29), 3.5 ± 1.1 (n = 27), and 4.0 ± 1.2 (n = 24)

days, respectively. The fifth instar had two distinct phases, an actively feeding

green-phase which lasted 5.3±1.3(« = 24) days, followed by a nonfeeding brown-

PAN-PACIFIC ENTOMOLOGIST

phase which lasted 1.8 ± 0.6 (n = 24) days. The prepupa and pupa lasted 1.7 ±

0.6 (n = 19) and 10.7 ± 0.6 (n= 17) days, respectively. Although five instars

were typical, a sixth (male or female) occurred occasionally (Campbell and Pike,

1984). In such cases, the third and fourth instars lasted about 1 extra day, a green-

phase fifth about a day shorter, followed by a green-phase sixth lasting about 5

days; the remaining stages were comparable to the “typical” individual. Since the

sixth instar was uncommon it was not included in Figure 1. Adults used for

studying fecundity lived 6 to 13 days, x = 9 (n = 13). Virgin females, however,

lived as long as 21 days.

Since eggs were usually laid singly, larvae were usually found one per branch

terminal, but two to three were fairly common. Following eclosion, first instars

bored into leaf buds or fed among the small subtending leaves. Feeding damage

by the first instars was noticeable only with close inspection. Some peppermint

plants showed a slight chlorosis at or near the point of feeding. First instars often

inhabited and fed individually within single flowers making them difficult to

detect. Prior to moulting, first instars constructed single loose-weave silk shelters

in the bud or in the leaves directly below the bud. Ecdysis took place in the shelters

followed by about one day of inactivity by the new larvae. The exuviae were

usually consumed by the larvae.

The second instar was often found in the same locations on the plant as the

first, except not usually inside the flowers. It often hid among the flowers and fed

on the calyxes, or the basal aspects of the leaves. Third instars typically occurred

among small leaves subtending the bud or among larger leaves. These larger leaves

were often silked together by the larvae. Damage by this instar consisted of chewing

holes or removing upper leaf tissue in transverse bands at mid-leaf, causing leaf

dieback distal to the feeding. Leaves tied together by silk occurred not only with

the first three instars, but also the fourth and fifth. These silked together leaves

served not only as ecdysial shelters, but also retreats on and in which larvae fed.

It was not uncommon for a single individual during feeding and development to

construct several shelters. Fourth and fifth instars clearly caused the most plant

damage. These latter instars were highly active and moved freely through the crop

canopy, often feeding on several plants. Feeding ceased with the brown-phase

fifth instar, which would leave the plant to construct pupal shelters on the soil.

The shelters consisted of fallen mint leaves or soil particles silked together. Adults

emerged through the proximal end of the pupal shelters.

A number of natural enemies were found associated with P. orphisalis. The

reared parasitoids included ichneumonids, braconids, and a tachinid (Table 1).

Pristomerus spinator Fabricius (Hym., Ichneumonidae) was the most predomi¬

nant in the rearings. It parasitized commonly third through fifth instars, rarely

first or second. Initial parasitization probably began with thirds, since they were

the first to venture into the more open, vulnerable microhabitats. The host gen¬

erally reached the pupal stage before death and eventual consumption by the

parasitoid. Cotesia sp. (Hym., Braconidae) was about one-fifth as prevalent as P.

spinator. It also attacked third through fifth instar larvae, but differed from P.

spinator in that it caused death in the host earlier, generally by the fourth or fifth

instar. Cotesia sp. pupated in a self-spun cocoon within a feeding shelter, but

outside its host. Meteorus campestris Yier (Hym., Braconidae), a less abundant

parasitoid than Cotesia sp., attacked third through fifth instar larvae and caused

VOLUME 61, NUMBER 1

death to the brown-phase fifth. Generally, the fifth completed its pupal shelter

before death. Only single specimens of Stictopisthus bilineatus Thomson (Hym.,

Ichneumonidae) and an unknown tachinid fly were obtained in the rearings.

Collectively, the five species encountered caused 12% mortality to P. orphisalis

based on the findings averaged over the 2-year study.

Predators of P. orphisalis included minute pirate bugs, Orius sp. (Hemiptera:

Anthocoridae), feeding on the eggs, and big-eyed bugs, Geocoris sp. (Hemiptera:

Lygaeidae), reduviid hemipterans and thomisid spiders feeding on the larvae.

Literature Cited

Campbell, C. L., and K. S. Pike. 1984. Descriptions of the life stages of Pyrausta orphisalis (Lepi-

doptera: Pyralidae). Pan-Pacific Entomol., 60:332-336.

Frick, K. E. 1961. Control of insects and mites attacking mint in central Washington. J. Econ.

Entomol., 54(4):644-649.

Munroe, E. 1976. Pyraloidea (in part). Pp. 119-120, Fasc. 13-28 in R. B. Dominick et al., The

moths of America north of Mexico. The Curwen Press, London.

Walker, F. 1859. List of specimens of lepidopterous insects in the collection of the British Museum.

London, Pt. 17, p. 310.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 48-49

The First Record of the Delphacid Liburniella ornata in

California (Homoptera: Fulgoroidea)

Stephen W. Wilson and Raymond J. Gill

(SWW) Department of Biology, Central Missouri State University, Warrens-

burg, Missouri 64093; (RJG) California Department of Food and Agriculture,

Sacramento, California 95814.

Liburniella ornata (Stal) has been reported from Maine south to Florida and

west to Texas, Nebraska, and Minnesota (Metcalf, 1943) as well as Bermuda (Van

Duzee, 1909) and possibly Ecuador (see Muir, 1926). The only record of its

occurrence west of the Rocky Mountains is that of Downes (1927) for British

Columbia.

L. ornata is common and widely distributed in the eastern United States (Os¬

born, 1938) and is often collected in grassy meadows (Wilson, pers. obs.). It has

been recorded feeding on black walnut (Juglans nigra L.) (Nixon and McPherson,

1977), and has been collected in June and from September to mid-October in

Illinois (Weber and Wilson, 1981). Otherwise, no information is available on its

biology. An adult was illustrated by Metcalf (1923) and the male genitalia were

figured by Muir (1926).

We recently found three specimens of L. ornata from California among un¬

identified delphacids in the B. P. Bliven collection which has been acquired by

the California Academy of Sciences and the California Department of Food and

Agriculture.

The label information for the specimens is as follows:

CALIFORNIA: Humboldt Co., Fieldbrook, 23 July 1939, coll. B. P. Bliven, 1

male, 1 female; Pepperwood, Greenlaw Creek, 9 September 1973, coll. B. P.

Bliven, 1 female [brachypter].

Although L. ornata is of widespread occurrence in the eastern U.S., it has only

been collected in two localities in northern California. These limited collection

records and the substantial geographic and ecological barriers suggest that this

delphacid is an introduced species. Since females have a saw-like ovipositor and

probably insert their eggs in plant tissues as do other delphacids, imported plant

materials containing eggs, such as nursery or house plants or packing straw, may

be a source of introduction. Its spread may be limited by the distribution of

particular introduced plants or by a restricted set of ecological conditions present

in the area where the specimens were collected.

Acknowledgment

We wish to thank Dr. P. H. Amaud, Jr., Department of Entomology, California

Academy of Sciences, San Francisco, for the loan of specimens.

VOLUME 61, NUMBER 1

Literature Cited

Downes, W. 1927. A preliminary list of the Heteroptera and Homoptera of British Columbia. Proc.

Entomol. Soc. British Columbia, 23:1-22.

Metcalf, Z. P. 1923. A key to the Fulgoridae of eastern North America with descriptions of new

species. J. Elisha Mitchell Sci. Soc., 38:139-230.

-. 1943. General catalogue of the Hemiptera. Fasc. IV, Fulgoroidea. Part 3, Araeopidae (Del-

phacidae). 552 pp.

Muir, F. 1926. Contributions to our knowledge of South American Fulgoroidea (Homoptera). Part

I. The family Delphacidae. Bull. Hawaiian Sugar PI. Assoc. Exp. Stn., 18:1-51.

Nixon, P. L., and J. E. McPherson. 1977. An annotated list of phytophagous insects collected on

immature black walnut trees in southern Illinois. Great Lakes Entomol., 10:211-222.

Osborn, H. 1938. The Fulgoridae of Ohio. Bull. Ohio Biol. Surv., 35:283-349.

Van Duzee, E. P. 1909. Notes on some Hemiptera taken in the Bermudas by W. J. Palmer. Can.

Entomol., 41:126-128.

Weber, B. C., and S. W. Wilson. 1981. Seasonal and vertical distributions of planthoppers (Ho-

moptera:Fulgoroidea) within a black walnut plantation. Great Lakes Entomol., 14:71-75.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 50-53

A New Species of Orchesella from Manitoba, Canada

(Collembola: Entomobryidae)

Jose A. Mari Mutt

Department of Biology, University of Puerto Rico, Mayagiiez, Puerto Rico

00708.

Several months ago I received six microscope slides containing seven specimens

of Orchesella collected at Fort Whyte, Manitoba, Canada. The specimens were

escorted by a letter from Dr. Kenneth Christiansen, Department of Biology, Grin-

nell College, Iowa, who stated that they were unlike any member of the genus

from North America. A detailed study indicated that the specimens belong to the

new species described below. The holotype and two paratypes are deposited in

the Museum of Comparative Zoology, Cambridge, Massachusetts; two paratypes

are in Dr. Christiansen’s collection, and the remaining two paratypes are in my

collection.

The system of nomenclature used for head macrochaetotaxy was described by

Mari Mutt (1979). The system for the macrochaetotaxy of the second and third

abdominal segments follows the terminology proposed by Christiansen and Tuck¬

er (1977; see also Christiansen and Bellinger, 1980) for the third segment, and

which I have modified only slightly to use it also for the second abdominal segment

(Mari Mutt, 1984).

Orchesella manitobae Mari Mutt, New Species

(Figs. 1-6)

Length to 1.8 mm (x = 1.56 mm, n = 7). Antennae 3.9-4.4 x length of head

and 0.74-0.82 x length of head plus body. Antennal segments 1 to 4 (Ant. 1-4)

with violet pigment restricted mainly to sides and apices, segments 5 and 6 evenly

pigmented. Head with median V-shaped band and sometimes a rounded patch

anterior to this band. Pigment also laterally behind eye patches, around antennal

bases, and on an intense spot between the bases of the antennae. Body evenly

covered by light violet pigment (Fig. 1) except for a clear median streak on Th.

2 and Th. 3 (not visible in the photograph). Legs lightly but evenly pigmented or

pigment restricted to coxae. Collophore pigmented distally, furcula without pig¬

ment.

Apex of Ant. 6 with a 3-pointed pin seta and without papilla or protruding

structures. Anterior head macrochaetotaxy (Fig. 5) follows formula: An = 4-5,

A = 7, M = 4, S = 10. Four macrochaetae along midline of head. Prelabral setae

rarely bifurcated. Labral papillae (Fig. 6) with pointed tips. Differentiated seta of

outer labial papilla well developed but not reaching apex of its papilla (Figs. 3,

4), latter with 2 anterior setae and a single posterior seta external to the differ¬

entiated seta. Setae of anterior labial row smooth and subequal in length. Posterior

labial row internal to seta E with 4-7 setae per side, all ciliated. Setae E, L, and

L 2 ciliated. Macrochaetal formula for second abdominal segment (Abd. 2): IA =

VOLUME 61, NUMBER 1

Figure 1. O. manitobae. Holotype, the specimen measures 1.8 mm.

4-7, OA = 1-2, MP = 3 (4 on one side of one specimen), L = 2. Formula for

Abd. 3: IA = 3-5, OA = 1, M = 2-4, L = 3. Corpus of tenaculum with 1 smooth

seta. Inner margin of unguis with basal pair of teeth and 2 distal unpaired teeth

(Fig. 2), outer margin with a small median tooth and 2 large lateral teeth. Un-

guiculus with a tooth placed near center of an outer lamella. Male genital plate

circinate, with about 17 smooth setae arranged in 1 row around periphery of plate.

Manubrium and dentes with many ciliated setae and no smooth setae. Mucro

with 2 teeth and basal spine.

Diagnosis.— In coloration the new species comes closest to the North American

O. carneiceps Packard and O. folsomi Maynard. The first is almost black except

for the pale head (Christiansen and Bellinger, 1980:807) and the second is also

dark but both head and mesothorax are yellow (Maynard, 1951:189). In O. man¬

itobae, pigment is almost evenly distributed over head and body and is much

lighter. The unguicular tooth of O. manitobae and O. carneiceps is placed near

the middle of an outer lamella but in O. folsomi the tooth arises distally.

In Stach’s 1960 revision of Orchesella, the new species keys out to O. folsomi.

None of Stach’s 117 figures illustrating the pigmentation of European species

match closely the distribution of pigment of O. manitobae.

Comments.— The color pattern described above is based on the three largest

specimens. Two small specimens possess very little pigment but it is still evenly

distributed.

Head macrochaetotaxy was studied fully in only one specimen, in all others

the head is distorted and it is impossible to work out the complete pattern. The

macrochaetotaxy of Abd. 2 and Abd. 3 was studied in almost all the specimens

52 PAN-PACIFIC ENTOMOLOGIST

Figures 2-6. O. manitobae. 2. Prothoracic claws. 3. Right outer labial papilla with its differentiated

seta and accompanying setae, note presence of one basal seta. 4. As preceding, left outer labial papilla.

5. Anterior head macrochaetotaxy. 6. Labral papillae.

VOLUME 61, NUMBER 1

and it revealed much variation, which limits the use of this character for separating

closely related species. This variation may be due to changes in the number of

setae during growth, inherent variability, and the asymmetric distribution of setae

which is rather frequent in members of this genus.

Material examined .—All the specimens were collected by J. Aitchison on the

surroundings of Fort Whyte, Manitoba, Canada. Below I give the accession num¬

ber provided by Dr. Christiansen and additional data for each specimen.

Number 5507—holotype (2), pitfall trap placed on small marshy meadow,

20.1.1982; 5508A-1 2, 25.VI.1980, 725 m; 5506-1 5, 1 2, 20.V.1981; 5511-1 2,

15.X. 1980, an aspen-bur oak wood litter, 805 m; 5512—1 2, 20.VIII.1980, small

marshy meadow being invaded by trees, 737 m; 5516—1 <3, 20.VII.1979, in a

meadow.

Literature Cited

Christiansen, K., and P. Bellinger. 1980. The Collembola of North America north of the Rio Grande,

vol. 3, family Entomobryidae. Grinnell College, Grinnell, Iowa, pp. 789-1042.

-, and B. E. Tucker. 1977. Five new species of Orchesella (Collembola: Entomobryidae). Proc.

Iowa Acad. Sci., 84:1-13.

Mari Mutt, J. A. 1979. A revision of the genus Dicranocentrus Schott (Insecta: Collembola: Ento¬

mobryidae). Univ. Puerto Rico Agr. Exp. Stn. Bull. 259, 79 pp.

- 1984. Five new species of Orchesellini from Central Mexico (Collembola: Entomobryidae:

Orchesellinae). Proc. Entomol. Soc. Wash., 86:808-820.

Maynard, E. A. 1951. A monograph of the Collembola or springtail insects of New York State.

Comstock Publ. Co., Ithaca, N.Y., 339 pp.

Stach, J. 1960. The apterygotan fauna of Poland in relation to the world-fauna of this group of

insects. Tribe Orchesellini. Panstwowe Wydawnictwo Naukowe, Krakow, 151 pp.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 54-57

Oviposition on the Backs of Female Giant Water Bugs,

Abedus indent at us: The Consequence of a Shortage in

Male Back Space? (Hemiptera: Belostomatidae)

Bill Kraus

Department of Biology, C-016, University of California, San Diego, La Jolla,

California 92093.

Females of the subfamily Belostomatinae (Lauck and Menke, 1961) character¬

istically fasten their eggs onto the dorsum of their mates, the males subsequently

carrying the clutch until hatching. The use of males as oviposition sites led R. L.

Smith (1979a) to suggest that at times a female’s reproductive output may more

likely be limited by the availability of unencumbered males on which to oviposit

than by her own egg production. If this is true, then a gravid female unable to

secure a “free” male may opt instead to lay a small clutch on some alternative

substrate rather than not lay at all. Recently, Kruse and Leffler (1984) have

observed egg-bearing females in a population of Belostoma flumineum. In this

note I report a similar observation for Abedus indentatus.

Of 706 sexually-mature adult females 12 of A. indentatus Haldeman examined

from 3 May to 15 August 1983 at Deep Canyon Creek, Palm Desert, California,

1 observed three females bearing eggs on their dorsum. (Females are identified by

the possession of two tufts of setae located mesally near the apical margin of the

genital plate [Menke, 1960].) The first female was collected on 17 May and carried

13 eggs (2.9 mm each), the second female was collected on 31 May and carried

2 eggs (2.6 mm each), and the third female was collected on 6 June and carried

52 eggs (3.6-4.6 mm each). The third female was recaptured on 7 July carrying

27 eggs, 14 of which had hatched as indicated by the presence of 14 empty chorions

with ruptured cephalic caps (Smith, 1974). Whereas egg deposition on males begins

at the apex of the hemelytra and proceeds forward, uniformly covering the dorsum

of the male (Smith, 1974), the placement of eggs on the three females did not

follow this pattern (see Fig. 1).

The fact that egg-encumbered females represent less than one percent of those

examined in the field suggests that this phenomenon is quite rare. Since anatomical

1 All individuals were marked for future identification, some individually with numbers on their

pronotum, others simply by making a small cut in one hemelytron. All three of the egg-bearing females

collected in the field had been individually marked.

2 The 706 females represent females seen for the first time. None of the 229 recaptured females that

was examined carried eggs. An additional sample of 672 recently-eclosed, sexually-immature adult

females was not included here since it was uncertain as to whether their exoskeletons had hardened

enough to bear eggs. No recently-eclosed male or female was ever seen carrying a clutch.

Figure 1. Photographs of three egg-carrying females (A, B, C) collected at Deep Canyon Creek,

California. Photograph (D) depicts the female shown in (C) 31 days later. Note the presence of empty

chorions (arrow).

VOLUME 61, NUMBER 1

!Ja'> 'J-.i

PAN-PACIFIC ENTOMOLOGIST

constraints preclude a female from ovipositing on her own dorsum, the eggs must

necessarily be the product of a second female. Smith (1979a) has reported that

heavily gravid female A. herberti will lay small clutches of eggs on floating aquatic

vegetation in their containers. He suggested that this apparent relaxation in spec¬

ificity for an oviposition substrate was due to a prolonged absence of the appro¬

priate surface, i.e., an unencumbered male. I have seen similar behavior in A.

indentatus, and Menke (1960) has demonstrated that a gravid A. indentatus female

will oviposit at least a few eggs on the back of a male of a different species if a

conspecific is unavailable. In addition, I have observed two instances of egg

carrying by females in the laboratory (clutch sizes of 9 and 3 eggs, respectively),

both instances occurring in aquaria containing only females. Although neither of

these encumbered females were successful at rearing the clutch (both clutches were

discarded before the eggs could hatch), one of the eggs deposited on the female

carrying three eggs expanded to 5.6 mm before it was lost. A well-developed

embryo was found when the discarded egg was dissected.

Judging by their size, the eggs on the three A. indentatus females collected in

the field were laid in May and June 1983, months in which the majority of sexually -

mature A. indentatus males carried broods (86.44%, n = 59, and 74.55%, n = 55,

respectively). In addition, the adult sex ratio during the same period appeared to

be significantly skewed towards females. 3 (Interestingly, Kruse and Leffler [1984]

indicate that a majority of Belostoma flumineum males were also encumbered

when they made their observations.) I therefore suggest that egg-encumbered

females are the consequence of at least a temporary shortage in available male

back space. It remains unclear as to whether females are simply renewing their

oviducal egg supply until “free” males become available (Smith, 1979a), or wheth¬

er they are attempting to rear a small brood in the absence of such males. In

regards to the latter hypothesis, I agree with Kruse and Leffler (1984) that this

phenomenon is not an alternative to male brooding. However, it may be a viable

option in instances where males are locally unavailable and when the only other

alternative is not to lay at all. Finally, although egg-encumbered females could

conceivably be the result of recognition errors made by ovipositing females, the

complexity of courtship that characterizes Abedus (Smith, 1979a; Kraus, pers.

obs.) makes this hypothesis unlikely.

Smith (1979b) has convincingly argued that the repeated bouts of copulation

observed in Abedus are an adaptation to assure paternity through sperm presi-

dence. The presence of hatched eggs on the back of the female recaptured in July

demonstrates that the potential for cuckoldry does exist in nature, since the fertility

of these eggs was dependent on the ovipositing female having retained at least

some viable sperm from a previous mating. In addition, although male-specific

brooding behaviors such as broodpumping appear to significantly increase the

hatching success of a brood (Smith, 1976a, 1976b; Yenkatesan, 1983), they are

apparently not essential for egg development. In fact, the low survival rate of

female-borne clutches may be as much a result of the irregular placement and

small sizes of these clutches as it is a consequence of the disparity in brooding

3 The sex ratio of previously unmarked, sexually-mature adults closely fit a 2:1 ratio in favor of

females (n = 263, x2 = 0.008, df = 1, P > 0.9). Sampling bias in catchability, analyzed over a 2-week

period subsequent to marking, could not be detected (« = 103 recaptures, x2 = 0.60, df = 1, P > 0.2).

VOLUME 61, NUMBER 1

behaviors between the sexes. Even among males, small clutches appear to be

much more susceptible to losing their adhesion and being discarded than large

ones (especially when expansion of the developing eggs increases drag). In this

regard, it is suggestive that the one field-collected female which was observed to

have successfully reared a clutch also carried the largest number of eggs.

Acknowledgments

I wish to thank Robert Gibson and Bob Smith for their helpful comments and

criticisms. I would also like to thank the staff of the Phillip L. Boyd Deep Canyon

Desert Research Center for the facilities made available to me. This work was

supported in part by a National Science Foundation predoctoral fellowship.

Literature Cited

Kruse, K. C., and T. R. Leffler. 1984. Females of the giant water bug, Belostoma flumineum (He-

miptera: Belostomatidae), captured carrying eggs. Ann. Entomol. Soc. Amer., 77:20.

Lauck, D. R., and A. S. Menke. 1961. The higher classification of the Belostomatidae (Hemiptera).

Ann. Entomol. Soc. Amer., 54:644-657.

Menke, A. S. 1960. A taxonomic study of the genus Abedus Stal (Hemiptera, Belostomatidae). Univ.

of Cal. Publ. Entomol., 16:393-439.

Smith, R. L. 1974. Life history of Abedus herberti in Central Arizona (Hemiptera: Belostomatidae).

Psyche, 81:272-283.

-. 1976a. Brooding behavior of a male water bug Belostoma flumineum (Hemiptera: Belo¬

stomatidae). J. Kansas Entomol. Soc., 49:333-343.

-. 1976b. Male brooding behavior of the water bug Abedus herberti (Hemiptera: Belostomat¬

idae). Ann. Entomol. Soc. Amer., 69:740-747.

-. 1979a. Paternity assurance and altered roles in the mating behavior of a giant water bug,

Abedus herberti (Heteroptera: Belostomatidae). Anim. Behav., 27:716-725.

-. 1979b. Repeated copulation and sperm presidence: paternity assurance for a male brooding

water bug. Science, 205:1029-1031.

Venkatesan, P. 1983. Male brooding behavior of Diplonychus indicus Venk. & Rao (Hemiptera:

Belostomatidae). J. Kansas Entomol. Soc., 56:80-87.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 58-59

The Larva of Cyphomella gibbera Saether (Diptera: Chironomidae)

P. G. Mason

Research Station, Agriculture Canada, 107 Science Crescent, Saskatoon, Sas¬

katchewan S7N 0X2, Canada.

Abstract .—The larva of Cyphomella gibbera Saether is described. It differs from

the other described Cyphomella sp. in having a smaller antennal ratio and in the

colouration of the lateral notches of the median mental tooth.

Saether (1977) first described the male and pupa of Cyphomella gibbera based

on material from South Dakota. In a recent study of Saskatchewan River Chi¬

ronomidae (Mason, 1978) a number of imagines were reared from 4th instar

larvae collected from sand, gravel, pebble and cobble substrates covered with silt

or muck.

The larva of C. gibbera differs from that of Cyphomella sp. (Saether, 1977) in

antennal and mental features. The former species has a smaller antennal ratio

(AR of C. sp. is 1.78 and that of C. gibbera is 1.20-1.55) and the lateral notches

of the median mental tooth are lighter-coloured than the lateral teeth. Cympho-

mella sp. has the lateral notches of the median mental tooth concolourous with

the lateral teeth (Saether, 1977, Fig. 38H).

Cyphomella gibbera Saether, 1977, Bull. Fish. Res. Board Can., 196:103.

Head capsule brown, except tormae, U-shaped sclerite, lobes of premandibles,

base of mandibles, lateral mental teeth and postoccipital margin which are dark

brown. Head width 0.21-0.25, 0.23 mm. Ventral head length 0.09-0.10, 0.10

mm. Labral sensillum (Fig. 1) three-segmented. SI simple; length, 20-28, 23.8

pm. SII 45-60, 49.8 gm long. Pecten epipharyngis a small scale with 2-3 teeth.

Premandibles (Fig. 2) length: 58-70, 63 gm. Antenna (Fig. 3) with apical three

segments minute; total length 66-77, 71.3 gm; lengths of individual segments

(gm) 35.0-45.0, 40.6; 16.0-28.0, 21.5; combined lengths of segments three to

five, 5.0-10.0, 8.2; AR 1.2-1.6, 1.4; ring organ on basal 0.26-0.44, 0.33 of first

segment; antennal blade length 26.0-37.0, 32.5 ^m; antennal style on apex of

segment two. Maxillary palp (Fig. 5) three-segmented, with 7-8 styles at apex of

basal segment; ratio of length to basal width of first segment 1.5-2.0, 1.7. Mandible

(Fig. 4) with three inner and apical tooth light-coloured; total length 90.0-112.5,

103.0 gm; ratio of length to basal width 0.6-1.2, 0.9; seta subdentalis narrowed

apically, length 14.0-22.0, 17.8 gm; inner mandibular margin smooth. Mentum

(Fig. 6) 94.5-130.5, 111.6 gm wide. Ventromental plates with smooth anterior

margin and acute inner angle; striations distinct; ratio of width to length 2.3-2.9,

2.6. Posterior parapods with 15-16 simple hooks. Each procercus short, with eight

anal setae. Anal tubules narrowed and rounded apically, 0.8-1.6 times as long as

basal width.

VOLUME 61, NUMBER 1

Figures 1-6. Cyphomella gibbera Saether. 1. Labral sensillum. 2. Premandible. 3. Antenna. 4.

Mandible. 5. Maxillary palp. 6. Mentum and premental plates (scale, 100 jum).

Acknowledgments

I would like to thank D. M. Lehmkuhl who provided laboratory facilities and

NSERC funds for materials and travel. The Institute for Northern Studies and

the College of Graduate Studies and Research at the University of Saskatchewan

provided scholarship funds for my work on chironomids.

Literature Cited

Mason, P. G. 1978. A biosystematic study of larval and pupal Chironomini (Diptera: Chironomidae)

in the North and South Saskatchewan Rivers. M.Sc. thesis, Univ. of Saskatchewan, Saskatoon,

456 pp.

Saether, O. A. 1977. Taxonomic studies on Chironomidae: Nanocladius, Pseudochironomus and the

Harnischia complex. Bull. Fish. Res. Board Can. 196, 143 pp.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 60-67

The Nearctic Species of Deuterixys Mason

(Hymenoptera: Braconidae)

James B. Whitfield

Department of Entomological Sciences, University of California, Berkeley, Cal¬

ifornia 94720.

Abstract. —The three Nearctic species of the genus Deuterixys are described and

keyed for the first time: D. quercicola Whitfield, D. pacifica Whitfield and D.

bennetti Whitfield. Comparisons are made to the described Palearctic species.

The genus Deuterixys was proposed by Mason (1981) to include the members

of Nixon’s (1965, 1976) carbonarius-group of Apanteles Foerster: A. carbonarius

(Wesm.), A. rimulosus Niez., A. nixoni Papp and A. patro Nixon, all Old World

species. Below I describe and key three new species from North America, all

reared from Bucculatrix spp. (Lepidoptera: Lyonetiidae).

Materials and Methods

The 149 examined specimens came from the collections mentioned in the

acknowledgments and from my own collection. All measurements were made to

the nearest 0.1 mm at 50 x or 100 x using a Leitz dissecting microscope with an

ocular micrometer. The wing figures were prepared from slide-mounted material

using a microprojector. Morphological terminology follows that of Mason (1981).

Deuterixys Mason, 1981

A detailed generic description is provided by Mason (1981). The genus can be

distinguished from other Microgastrinae by the following combination of features:

vein r-m2 of fore wing (2r-m of Mason, 1981) absent; propodeum polished and

bearing a strong longitudinal medial carina; tergite I of metasoma with mediobasal

longitudinal groove; tergites II and III broad, subrectangular and usually densely

sculptured; ovipositor sheaths short, decurved, subexserted. In addition, the species

treated here share the following characteristics which may not be universal within

the genus: metanotum sublaterally withdrawn from scutellum, exposing meso-

thoracic postphragma; sublateral setiferous lobes small but projecting anteriorly

to near scutellum; tibiae of metathoracic legs with 10-12 spines on outer face;

tergite II of metasoma with a more or less distinct raised medial section.

As far as is known, all Deuterixys species parasitize the larvae of Bucculatrix

spp. and are unusual among Microgastrinae in emerging as adults from the host

cocoon. In addition to the described Old World and the new Nearctic species

discussed here, I have seen several undescribed species from Central and South

America.

VOLUME 61, NUMBER 1

Deuterixys quercicola Whitfield, New Species

(Figs. 1, 4)

Holotype female. — Overall length 1.8 mm; fore wing length 1.9 mm.

Head: Antennae slightly longer than body, entirely dark; apical 9 flagellomeres

with only one rank of placodes; 2nd flagellomere 3.0 x longer than broad, 14th

flagellomere 1.4 x longer than broad. Frons 1.4 x broader at midheight than

midlength; inner margins of eyes converging towards clypeus. Punctation of head

indistinct.

Mesosoma: Mesoscutum black, weakly punctate anteriorly, becoming nearly

smooth and polished posteriorly. Scutellar disc mostly nearly impunctate with

weak punctation appearing peripherally. Propodeum highly polished, virtually

unsculptured except for strong medial longitudinal carina and weak transverse

sculpturing near spiracles and along anterior third of medial carina; declivous

over most of length.

Legs: Prothoracic legs fulvous except darker brown coxal bases, femoral bases

and apical tarsomeres. Mesothoracic legs dark brown proximally except yellowish

distal trochanters; femoral apices, tibiae and tarsi fulvous. Metathoracic legs dark

brown except in vicinity of trochanters and bases of hind tibiae (here fulvous).

Hind apical tibial spurs short, whitish, subequal in length.

Wings: Tegulae dark brown, weakly translucent. Metacarp (Rl) of fore wing

virtually same length as stigma; 2r and IRs subequal in length and meeting at

rounded, approximately 135 degree angle. Pigmentation of fore wing veins yellow-

brown proximally, becoming more grey-brown distally.

Metasoma: Tergite I about 1.2 x longer than posterior width, broadening slightly

posteriorly, coarsely aciculorugose, with strong longitudinal groove surrounded

by broad depression over basal 0.3. Tergite II quadrate, 1.6 x broader than long,

aciculorugose with stronger overlay of longitudinal sculpturing than in tergite I;

medial raised portion indistinct. Tergite III separated from II by crenulate groove,

becoming abruptly wider than tergite II but with rounded lateral margins; length

somewhat less than that of tergite II; sculptured as in II. Tergite IV short, apically

rounded, overlapped by III and with fine, weak longitudinal sculpturing over most

of visible surface. Remainder of tergites telescoped under anterior 4 in dried

specimens. Hypopygium short, blunt, with sparse long hairs. Ovipositor sheaths

short, decurved, subexserted, apically hairy.

Male. — Similar to female except antennae longer, apical flagellomeres more

slender (flagellomere 14 2.2 x longer than broad), with only apical 4-5 flagello¬

meres having a single rank of placodes. Venation of fore wing slightly darker.

Variation .—Fore wing length 1.6-2.1 mm, with little or no sexual size dimor¬

phism. Coloration comparatively uniform in California material (see Comments

below for Eastern specimens).

Material examined. —Holotype 5: CALIFORNIA: Alameda Co., Berkeley Hills,

7-VIII-1981 (J. B. Whitfield), reared ex Bucculatrix albertiella cocoons, JBW no.

81H4, emgd. 15-18-VIII-1981. Paratypes: CALIFORNIA: Alameda Co., Berke¬

ley Hills, 1 6, 6 9, 2-V-1981 (J. B. Whitfield), same host as holotype, emgd. 9-

18-V-1981, 1 8, 1 2, same data except coll. 25-IV-1982, emgd. 20-26-V-1982, 1

<3, same data except coll. 11-IV-1981, emgd. 8-V-1981. Contra Costa Co., Tilden

Regional Pk., 3 9, 8 <5, 28-VIII-1982, emgd. 1-22-IX-1982 (J. B. Whitfield), reared

PAN-PACIFIC ENTOMOLOGIST

Figures 1-3. Fore wings of 1, Deuterixys quercicola, n. sp.; 2, D. pacifica, n. sp.; 3, D. bennetti, n.

sp. Scale = 0.5 mm.

ex Bucculatrix albertiella on Quercus agrifolia, JBW no. 82H12. Holotype de¬

posited in USNM; paratypes in USNM, CNC, UCB.

Other material: CALIFORNIA: Alameda Co., Strawberry Cyn., 1 8, 1 2, 11-IV-

1968 (P. A. Opler), reared ex Bucculatrix on Quercus agrifolia, JAP no. 68D117-8,

VOLUME 61, NUMBER 1

7 6, 6 9 , 6-IX-31-X-1975 (D. S. Green), same host. Contra Costa Co., 2 mi E

Antioch, 1 < 3 , 28-11-1968 (P. A. Opler), reared ex Bucculatrix on Quercus agrifolia ;

Briones Regional Pk., 1 $, 11-VIII-1981 (J. B. Whitfield). Los Angeles Co., 10 mi

N Castiac, 1 < 3 , 1 9 , 26-IV-1977 (D. S. Green), reared ex Bucculatrix on Quercus

agrifolia. Mendocino Co., Univ. Calif. Hopland Field Sta., 880', 1 < 3 , 5-V-1968,

malaise trap (W. J. Turner). Orange Co., O’Neill Park, 1 9 , 25-11-1977, reared ex

Aeaea dulcedo on Q. agrifolia (D. S. Green). San Diego Co., 4 mi W Ramona, 1

< 3 , 26-11-1977, reared ex B. albertiella on Q. agrifolia (D. S. Green). San Luis

Obispo Co., 9 mi SW Atascadero, 1 6, 4-V-1976, reared ex B. albertiella on Q.

agrifolia (D. S. Green). Santa Barbara Co., Santa Cruz Isl. nr. field sta., 1 9 , 1-V-

1976, reared ex B. albertiella on Q. agrifolia (D. S. Green). Sutter Co., Feather

R. at Nicolaus, 1 < 3 , 2-X-1982, reared ex Bucculatrix sp. on Quercus lobata, JBW

no. 82K19 (J. B. Whitfield), 1 9 , 4-X-1982, reared ex Bucculatrix sp. on Artemisia

douglasiana, JAP no. 82K1 (D. L. Wagner). Tulare Co., 3 mi N Kaweah, 1 9 , 28-

IV-1979, reared ex Bucculatrix sp. (J. A. Powell).

Hosts.—Bucculatrix albertiella Bsk. on Quercus agrifolia Nee, Bucculatrix sp.

on Quercus lobata Nee and possibly other Bucculatrix spp. on oaks. The record

from Aeaea dulcedo is probably in error and could easily have resulted from

unseen Bucculatrix contaminants in rearing material. I suspect the record from

Bucculatrix sp. on Artemisia douglasiana Bess, is also mistaken; I know that the

specimen from Bucculatrix from Quercus lobata was taken very nearby and Buc¬

culatrix larvae could easily have spun down from the oaks onto the Artemisia

foliage to pupate. All other Deuterixys from Bucculatrix on Artemisia have been

D. pacifica, n. sp., described below.

Comments. — Several specimens in the USNM collection from Baltimore, Mary¬

land, Washington, D.C. and Tallulah, Louisiana agree with the above description

in all features except the metathoracic femora are entirely fulvous and veins lCul

and 2Cul of the fore wing are less strongly pigmented than in the California

material. The specimens tend to be slightly smaller as well, but I have seen only

4 eastern U.S. individuals. I provisionally include these as D. quercicola, but more

material, including rearing records for the eastern forms, is necessary for a more

certain determination. It is certainly possible that D. quercicola appears widely

throughout the range of oak-feeding Bucculatrix.

This species can be distinguished from D. carbonaria (Wesm.) and D. nixoni

(Papp), its most similar Palearctic relatives, by its possession of the following

combination of features: 1) posteriorly broadening metasomal tergite I; 2) meta-

somal tergite IV with fine longitudinal sculpturing over most of its surface; 3)

tergite IV usually concealing successive terga in dorsal view; 4) propodeum with

strong medial longitudinal carina and 5) hind coxae dark brown. The resemblance

to D. nixoni is striking; it is possible the two may eventually prove to be syn¬

onymous.

Deuterixys pacifica Whitfield, New Species

(Figs. 2, 5)

Holotype female.—Ove rail length 1.6 mm, fore wing length 1.7 mm.

Head: Antennae approximately same length as body, nearly black throughout,

apical 9 flagellomeres with only one rank of placodes; 2nd flagellomere 3.0 x

longer than wide; flagellomere 14 1.3 x longer than wide. Frons 1.4 x broader at

PAN-PACIFIC ENTOMOLOGIST

midheight than long down middle; inner margins of eyes weakly converging to¬

wards clypeus. Punctation of frons, vertex and postgenae indistinct; microsculp¬

ture producing dull metallic reflections in diffused light.

Mesosoma: Mesoscutum shallowly punctate anteriorly, becoming less distinctly

and more sparsely punctate posteriorly. Scutellar disc distinctly punctate, duller

between punctures than mesoscutum, strongly convex posteriorly. Propodeum

highly polished, virtually unsculptured except for strong medial longitudinal Ca¬

rina; weakly convex in profile.

Legs: Prothoracic legs basally dark brown to black up to distal 0.5 of femur,

lighter fulvous beyond this point. Mesothoracic legs dark brown to black except

distal 0.2 of femur and proximal 0.6 of tibia, which are fulvous. Metathoracic

legs very dark brown to black except yellowish proximal 0.6 of tibia. Apical spurs

of hind tibiae short, whitish, subequal in length.

Wings: Tegulae dark brown, weakly translucent. Metacarp (Rl) of fore wing

slightly shorter than stigma. 2r and IRs virtually straight, subequal in length,

meeting at about 130 degree angle. Venation including stigma mainly deep brown;

M+Cu and 1A+2A weakly pigmented proximally.

Metasoma: Tergite I strongly narrowing posteriorly, approximately 2.5 x broad¬

er anteriorly than posteriorly, with strong medial longitudinal groove over basal

0.4-0.5; longitudinally rugose/carinulate over posterior 0.7. Tergite II 2.3 x broad¬

er anteriorly than long down middle, lateral margins converging noticeably pos¬

teriorly; surface longitudinally aciculorugose. Tergite III separated from II by weak

crenulate furrow, abruptly widening, becoming broader posteriorly than tergite II

at broadest point; surface with fine longitudinal sculpturing mediobasally, oth¬

erwise very weakly if at all sculptured. Laterotergites dark yellow-brown. Suc¬

ceeding terga of usual unsculptured, overlapping type. Hypopygium short, blunt

apically, somewhat truncate at tip, with sparse long hairs. Ovipositor sheaths

short, subexserted, decurved, polished, with hairs concentrated apically.

Male .—Similar to female except antennae clearly longer than body, with all but

apical 4-5 flagellomeres with 2 ranks of placodes. Length/width ratio of flagello-

mere 14 = 2.3 (other apical flagellomeres also more elongate than in female).

Variation.— Fore wing length 1.6-2.0 mm. No apparent sexual size dimorphism.

Coloration comparatively uniform except some darker high elevation individuals,

which also tend to have smoother metasomal sculpturing. A few specimens have

the first metasomal tergite less strongly narrowed than in the type series.

Material examined. — Holotype 2: CALIFORNIA: Contra Costa Co., Tilden

Regional Pk., 15-VIII-1982 (J. B. Whitfield). Paratypes: CALIFORNIA: Contra

Costa Co., Tilden Regional Pk., 6 2, 13 5, 15-VIII-1982 (J. B. Whitfield), 8 2, 6

5, 29-VIII-1982 (JBW), 8 2, 2 <5, 25-IV-1983 (JBW); Chabot Regional Pk., 3 mi

N Lake Chabot, 1 <3, 12-VII-1980 (JBW); nr. Clayton, 1 2, 27-11-1982, reared ex

Bucculatrix, JAP no. 82B29, emgd. 21-III-1982 (D. L. Wagner). Alameda Co.,

Berkeley, 11 2, 6 6, 10-IV-1983, reared ex Bucculatrix on Baccharispilularis, JBW

no. 83D19 (J. B. Whitfield). Holotype deposited in USNM; paratypes in USNM,

CNC, UCB.

Other material: CALIFORNIA: Alameda Co., Berkeley Hills, 2 <3, 6/7-VIII-

1982 (J. B. Whitfield); Patterson Reserve, Del Valle Lake, 1 <3, 29-IV-1974, reared

ex microlepidoptera, JAP no. 74D40-41 (J. A. Powell). Contra Costa Co., Pt.

Molate, Richmond, 1 <3, XI-1969, reared ex Iva axillaris, JAP no. 69L9. El Dorado

Co., Fallen Leaf Lake, 1 2, 9-VII-1983, reared ex Bucculatrix on Artemisia tri-

4 5 6

Figures 4-6. Basal tergites of 4, Deuterixys quercicola, n. sp.; 5, D. pacifica, n. sp.; 6, D. bennetti,

n. sp. Scale = 0.1 mm.

dentata, JBW no. 83G67 (J. B. Whitfield). Riverside Co., Thousand Palms, 1 <3,

27-IV-1955 (W. R. Richards); Palm Springs, 1 9, 3-V-1955 (W. R. M. Mason).

San Luis Obispo Co., Oso Flaco Lake, 2 9, 13-VII-1959 (C. A. Campbell). San

Mateo Co., San Bruno Mts., 1 3, 12-III-1982 (J. B. Whitfield), 2 9, 21-IV-1983,

reared ex Bucculatrix on Baccharis pilularis, JADeB. no. 83111 -E (J. A. De-

Benedictis), 1 9, 15-VI-1982, reared ex Bucculatrix on Baccharis pilularis (D. L.

Wagner). Ventura Co., San Nicolas Isl., 1 mi SSE 1000 Springs, 1 9, 6-8-VI-1980

(S. E. Miller, P. M. Mercer). NEW MEXICO: Springer, 1 3 (no date) (C. N. Ainslee).

UTAH: Guardsman Pass, nr. Brighton, 1 3, 10-VII-1981, 9800' (B. H. Poole).

BRITISH COLUMBIA: Robson, 1 9, 13-V-1947 (H. R. Foxlee).

Hosts.—Bucculatrix variabilis Braun and Bucculatrix separabilis Braun on Bac¬

charis pilularis DC, Bucculatrix sp. on Artemisia douglasiana Bess., Bucculatrix

sp. on Artemisia tridentata Nutt, and Bucculatrix sp. on Iva axillaris Pursh. I

suspect a wide range of Bucculatrix spp. on Compositae serve as hosts.

Comments.— No other known species of Deuterixys has the first metasomal

tergite so strongly narrowed apically. The relative lack of sculpturing on the third

tergite is also distinctive among the Nearctic species, although this feature is shared

with the Palearctic D. rimulosa (Niez.). The species appears to be widespread in

Western North America, probably in part due to the broad range of Artemisia

tridentata.

Deuterixys bennetti Whitfield, New Species

(Figs. 3, 6)

Holotype female.— Overall length 1.6 mm, fore wing length 1.7 mm.

Head: Antennae approximately 1.1-1.2 x longer than body; scapes and pedicels

PAN-PACIFIC ENTOMOLOGIST

light yellow-brown, more distal portions of antennae dark brown; apical 9 fla-

gellomeres with only one row of placodes; 2nd flagellomere 3.8 x longer than

broad; 14th flagellomere 1.2 x longer than broad. Frons 1.3 x broader at midheight

than long down middle; inner margins of eyes weakly converging towards clypeus.

Punctation of frons, vertex and postgenae faint; micro sculpture producing dull

metallic reflections in diffused light.

Mesosoma: Mesoscutum shallowly punctate anteriorly, becoming nearly im-

punctate posteriorly. Scutellar disc sparsely, shallowly punctate throughout, evenly

convex. Propodeum highly polished, virtually without sculpturing except for strong

medial longitudinal carina and weak transverse ridging in immediate vicinity of

longitudinal carina and lateral margins; propodeum strongly convex anteriorly in

profile.

Legs: Prothoracic legs fulvous virtually throughout except slightly darker apical

tarsomeres. Mesothoracic legs fulvous virtually throughout except infuscate distal

half of tibiae and apical tarsomeres. Metathoracic legs with darkened coxal bases,

apical third of femora, apical two-thirds of tibiae and entire tarsi; remainder of

hind legs lighter yellow-brown. Hind tibial spurs whitish, subequal in length, 0.4 x

as long as hind basitarsi.

Wings: Tegulae pale yellowish. Metacarp (Rl) 1.2-1.3X length of stigma; 2r

and IRs subequal, 2r very faintly arched, the two veins meeting at a 145 degree

angle. Venation of fore wing, including stigma, translucent pale yellow-brown.

Metasoma: Tergite I weakly narrowing posteriorly, 2x longer than broad at

midlength, with strong medial longitudinal groove over anterior 0.4; coarsely

aciculorugose over most of surface. Tergite II strongly quadrate, slightly broader

anteriorly than posteriorly, 1.8 x broader anteriorly than long down middle; sur¬

face coarsely aciculorugose throughout. Tergite III separated from II by strong

crenulate furrow, wider than II, abruptly widening over anterior 0.4, then parallel¬

sided; 2.2 x broader posteriorly than long down middle; surface coarsely, longi¬

tudinally carinulate over most of surface. Succeeding terga virtually sculptureless,

normally overlapping. Laterotergites translucent orange-brown. Hypopygium short,

blunt but not truncated apically, sparsely clothed with long hairs. Ovipositor

sheaths short, subexserted, decurved, polished, with hairs concentrated apically.

Male. — Similar to female except antennae longer, more slender (14th flagello¬

mere 2.2 x longer than broad), with single placode bands only on apical 4; antennae

generally somewhat lighter brown. Hind legs mostly evenly darker fulvous than

in female with nearly black coxae and lighter proximal portions of tibiae.

Variation. — Fore wing length 1.7-1.9 mm. No apparent sexual size dimorphism.

Coloration rather uniform in limited series available; some specimens in poor

condition and somewhat bleached. Some variation exists in the number of spines

on the outer faces of the hind tibiae.

Material examined. — Holotype 2: FLORIDA: Nassau Co., O’Neil, V-1960,

reared from Bucculatrix on Baccharis halimifolia (F. D. Bennett). Paratypes:

FLORIDA: Nassau Co., 2 2, 3 6, V-1960, same host (F. D. Bennett); Brevard

Co., Melbourne Beach, 2 2, 1 <3, V-1960, same host (F. D. Bennett); Walkulla Co.,

Medart, 1 2, V-1960, same host (F. D. Bennett); St. Johns Co., St. Augustine, 1

2, V-1960, same host (F. D. Bennett); Pinellas Co., St. Petersburg, 1 2, V-1960,

same host (F. D. Bennett), 3 2, X-1960, same host (F. D. Bennett); Pasco Co.,

Elfers, 1 <3, 16-IV-1960 (O. Peck). Holotype deposited in USNM, paratypes in

USNM, CNC collections.

VOLUME 61, NUMBER 1

Hosts. — The only recorded host is Bucculatrix sp. on Baccharis halimifolia L.

It is likely that other Bucculatrix spp. feeding on shrubby composites in the

Southeast may also serve as hosts.

Comments.— The species is named after F. D. Bennett, the collector of most

of the type series. D. bennetti strongly resembles the Palearctic D. rimulosa (Niez.)

(=D. comes (Wilk.) according to Papp, 1971), but differs in the much longer

metacarp and the more strongly sculptured third metasomal tergite.

Key to Nearctic Species of Deuterixys Mason

1. First tergite of metasoma narrowed posteriorly. 2

First tergite broadened posteriorly . D. quercicola, n. sp.

2. Tegulae pale yellow-brown; third metasomal tergite distinctly sculptured

over most of its surface. D. bennetti, n. sp.

Tegulae dark brown; third tergite at most only weakly sculptured antero-

medially . D. pacifica, n. sp.

Acknowledgments

I would like to thank the following curators for the loan of material in their

care: W. R. M. Mason, Biosystematics Research Institute, Ottawa (CNC); P. M.

Marsh, U.S. National Museum, Washington (USNM); L. E. Caltagirone, J. A.

Powell, Univ. California, Berkeley (UCB).

David L. Wagner was of great assistance in rearing the parasitoids from field-

collected host material, and in determining the hosts. I also appreciate the com¬

ments of H. V. Daly and W. R. M. Mason, who reviewed the manuscript.

Literature Cited

Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae):

a phylogeny and reclassification of Microgastrinae. Mem. Ent. Soc. Can., 115, 147 pp.

Nixon, G. E. J. 1965. A reclassification of the tribe Microgasterini (Hymenoptera: Braconidae). Bull.

Brit. Mus. (Nat. Hist.) Ent. Suppl., 2, 284 pp.

-. 1976. A revision of the north-western European species of the merula, lacteus, vipio, ultor,

ater, butalidis, popularis, carbonarius and validus -groups of Apanteles Forster (Hymenoptera:

Braconidae). Bull. Ent. Res., 65:685-732.

Papp, J. 1971. Ergebnisse der zoologische Forschungen von Z. Kaszab in der Mongolei, 265. Bra¬

conidae (Hymenoptera) III. Ann. Hist.-Nat. Mus. Nat. Hung. (Budapest), 63:307-363.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 68-71

Cutworm Defoliators of Ryegrass 1

J. A. Kamm

Forage Seed and Cereal Research Laboratory, USDA ARS, Department of

Entomology, Oregon State University, Corvallis, Oregon 97331.

Abstract. — Cutworm species that constitute a common complex that defoliate

ryegrasses in Oregon are: Dargida procincta (Grote), Pseudaletia unipuncta (Haw¬

orth), Leucania pallens luteopallens Grote, Diarsia pseudorosaria (Hardwick),

Peridroma saucia (Hubner), and Leucania insueta Guenee. In this study D. pro¬

cincta and P. unipuncta rapidly defoliated ryegrass by severing leaves and wasted

more foliage than they ate. D. pseudorosaria and L. p. luteopallens slowly defo¬

liated ryegrass and seldom wasted foliage by severing leaves. Black-light trap

captures provided reasonably good estimates of adult flight of all species except

D. procincta.

Various foliar feeding cutworms infest ryegrasses in the Pacific Northwest, but

little is known about the importance or type of damage inflicted by many species.

Among the cutworms common in ryegrass in the Willamette Valley of Oregon,

Pseudaletia unipuncta (Haworth) and Peridroma saucia (Hubner) are well known

pests (Bohart, 1948; Rice et al., 1982). Lesser known but common grass feeders

include Diarsia pseudorosaria (Hardwick), Leucania pallens luteopallens Grote,

Leucania insueta Guenee, and Dargida procincta (Grote) (Crumb, 1956). Of these

species, Thompson (1943) considered D. procincta to be the most important pest.

The impact of ryegrass defoliators may become more important in seed pro¬

duction with the recent introduction of plant growth retardants that, in effect,

decrease production of foliage and increase seed yields (Chilcote, 1983). This

study was conducted to determine the dominant species and relative abundance

of foliar feeding cutworms in seed fields of ryegrass. Also, greenhouse feeding tests

were made to determine the type of feeding injury inflicted by individual species.

Materials and Methods

Seasonal flight of adults was monitored with battery-powered black-light traps

placed in seed fields of ryegrass near Corvallis, Oregon. Traps were operated 3-

4 nights weekly during the flight season. Then, mean daily trap captures were

computed and multiplied times 7 so weekly data were comparable. Larval pop¬

ulations were estimated by collection of larvae from harvested seed on a field

basis. When fields were windrowed for harvest, larvae sought shelter in, and often

pupated within, the windrow. A large number of these larvae were carried along

with seed into the combine. As harvested seed was unloaded, larvae curled up

1 Contribution of the Agricultural Research Service, USDA, in cooperation with the Agricultural

Experiment Station, Oregon State University, Technical Paper No. 7073 of the latter.

VOLUME 61, NUMBER 1

and rolled down the pile of seed. These larvae were collected until at least 100

specimens were obtained from each field. The larvae were taken to the laboratory

to determine the percentage of each species in a given field.

Feeding tests were conducted in the greenhouse with densely seeded ryegrass

grown in pots (15 cm wide) to a height of 30-40 cm. Larvae from the previously

described collections were temporarily confined overnight in large freezer con¬

tainers and fed leaves of ryegrass. The next day, 20 active and uninjured larvae

(predominantly 4th-5th instars) were caged on individual pots of ryegrass. This

PAN-PACIFIC ENTOMOLOGIST

Table 1. Larval species composition of foliar feeding cutworms (%) obtained from several types

of ryegrass during harvest, Corvallis, Oregon.

Tetraploid

Annual

Perennial

Cutworm

1“

VII

VIII

Dargida procincta

Leucania pallens luteopallens

Pseudaletia unipuncta

Diarsia pseudorosaria

Periodroma saucia

a Fields I, III, V, VI, VIII sampled in 1980; all others sampled in 1981.

procedure was replicated 4 times for each species of cutworm. Infested plants

were observed daily for the type and degree of feeding damage inflicted by the

larvae until the plants were defoliated.

Results and Discussion

Adult flight.— The species of moths captured in light traps in ryegrass were, in

decreasing order of abundance: L. p. luteopallens, D. pseudorosaria, P. unipuncta,

L. insueta, P. saucia and D. procincta (Fig. 1). The abundance of species was

variable from year to year, especially the second generation of moths. For example,

the August generation of L. p. luteopallens was barely detectable in 1981, but

very strong in 1982. A strong second generation seems to occur only when rainfall

is adequate to stimulate grass regrowth soon after harvest.

Larval populations.— The species found in larval samples in decreasing order

of abundance were: D. procincta, L. p. luteopallens, P. unipuncta, D. pseudorosaria

and P. saucia. D. procinta was by far the most abundant larval species (Table 1)

and the least abundant in light traps. Often both larvae and pupae of this species

were easily found in windrows when relatively few or no adults were captured in

light traps. Apparently, the species is only weakly attracted to light traps. Based

on larval samples, 5 species clearly constitute a common cutworm complex in all

types of ryegrass. Three species were present in all fields, and the remaining two

species were found in 8 of 9 fields (Table 1). The relative abundance of larvae of

a given species varied among fields during the same year. For example, L. p.

luteopallens constituted only 1% of the population in one field and 54% in another.

Both fields were the same variety, and no insecticide was used on either field.

Feeding tests.— Larvae of 4 cutworm species were caged on ryegrass in the

greenhouse to evaluate feeding behavior in relation to plant defoliation. Larvae

of D. procincta and P. unipuncta initiated feeding on the margin of leaves near

the base of the plant. After larvae had eaten halfway through the leaf for several

cm, they often completely severed the leaf and then climbed on to a new leaf and

repeated this behavior. The amount of leaf foliage wasted (not eaten) was sub¬

stantially greater than the amount actually consumed by larvae of both species.

The plants (30-40 cm in height) in all pots were completely defoliated by 20

larvae after 2 days, leaving developing seed culms severed 3-5 cm above the soil.

The feeding behavior of both species was very similar.

In contrast, larvae of L. p. luteopallens fed along the leaf margins and often

consumed the tissue on only one side of the leaf midrib. As feeding progressed

VOLUME 61, NUMBER 1

toward the top of the leaf, the midrib was severed but some tissue remained so

the leaf was partially attached. A few leaves were eventually severed but only

after substantial amounts of foliage were consumed. Complete defoliation by 20

larvae required 8 days or 4 times as long as D. procincta or P. unipun eta.

Larvae of L. pseudorosaria also fed along the leaf margins and consumed nearly

all the tissue on one side of the midrib of the leaf. Larvae rarely penetrated the

midrib or wasted foliage by severing the leaf. These larvae consumed foliage rather

slowly and none of the plants was completely defoliated by 20 larvae after 10

days. In fact, plant growth compensated for some defoliation during the test.

Light traps clearly were inadequate to assess adult populations of all species in

the complex of infested ryegrasses. The relative abundance of species varied among

fields and years, but the order of species dominance in light traps was about the

same as in larval samples except for D. procincta, the most important defoliator

of ryegrasses. Sampling larval populations will require visual inspection of lodged

foliage, windrows, or harvested seed because sweeping with an insect net was not

effective. Two considerations are important in assessing potential damage: 1) the

propensity of the dominant species to sever developing culms and 2) relative

abundance. Tests herein have shown that D. procincta and P. unipuncta were

capable of severing leaves and culms, whereas P. pseudorosaria and L. p. luteo-

paliens may contribute to defoliation but plant damage was less severe.

Literature Cited

Bohart, R. M. 1948. Sod webworms and other lawn pests in California. Hilgardia, 17:267-308.

Chilcote, D. O. 1983. Role of growth regulators in seed production. Oregon Crop Sci. Circ. No. 50,

23 pp.

Crumb, S. E. 1956. The larvae of Phalaenidae. USDA Tech. Bull. No. 1135, 356 pp.

Rice, S. E., A. A. Grigarick, and M. O. Way. 1982. Effect of leaf and panicle feeding by armyworm

(Lepidoptera: Noctuidae) larvae on rice grain yield. J. Econ. Entomol., 75:593.

Thompson, B. G. 1943. Cutworm control in Oregon. Oregon Agric. Exp. Sta. Circ., 147:1-4.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 72-78

Descriptions of the Immature Stages of Delphacodes bellicosa

(Homoptera: Fulgoroidea: Delphacidae)

Stephen W. Wilson

Department of Biology, Central Missouri State University, Warrensburg, Mis¬

souri 64093.

Abstract.— The immature stages of Delphacodes bellicosa Muir and Giffard are

described and illustrated. Features useful in separating nymphal instars include

the number of metatarsal segments, the size, shape, and dentation of the metatibial

spur, the increase in body size, wingpad size, and number of pits. Eggs of D.

bellicosa were found inserted in stems and nymphs and adults were observed

feeding on leaves of knotgrass ( Paspalum distichum L.)

Delphacodes bellicosa Muir and Gilfard (1924) is known only from the male

holotype collected at Three Rivers, Tulare Co., California. On 17 May 1981 I

collected 3 males of this species at a manmade lake in Bidwell Park, Chico, Butte

Co., California. Identity of females collected is questionable because two other

Delphacodes, D. consimilis (Van Duzee) and D. foveata (Van Duzee) were also

collected at the lake.

On 20 May 1981 an adult male D. bellicosa was observed feeding on the leaves

of knotgrass {Paspalum distichum L.). Ten plants were returned to the laboratory

and 52 eggs were found inserted in one stem; 10 of these were preserved in 95%

ethyl alcohol and the remaining eggs removed from the surrounding plant tissue

with a fine needle and placed on a strip of filter paper. The strip was placed in a

petri dish that was covered on the bottom with a disc of moistened filter paper.

The dish was covered with plastic secured with an elastic band and covered with

the lid; the plastic prevented newly hatched nymphs from escaping between the

dish and lid. Upon hatching, the nymphs were provided with a cutting of knotgrass

leaf added to the dish and replaced every three to four days. The dish was kept

in an incubator under a 16L:8D photoperiod at 29 ± 1.4°C. All of the eggs hatched.

Because 12 of the 1 st instars died within a few days of hatching (apparently by

drowning in the condensation on the walls and plastic cover of the dish), the

remaining nymphs were transferred to knotgrass that had been planted in a clay

pot; this knotgrass was collected at the same locality and carefully examined for

eggs before transplanting. The pot was covered with a glass cylinder and covered

with a piece of fine mesh gauze secured with an elastic band. The potted knotgrass

was kept in a greenhouse.

In order to obtain specimens for description several nymphs were collected

from the potted plant each week until the emergence of 3 adult males. Specimens

to be described were preserved in 95% ethyl alcohol. The first instar is described

in detail but only major changes from previous instars are described for subsequent

instars. Comparative statements refer to previous instars (e.g., more elongated).

VOLUME 61, NUMBER 1

Figures 1-6. Delphacodes bellicosa Muir and Giffard. 1. Habitus of male macropter, vertical bar =

0.5 mm. Male genitalia. 2. Complete left lateral view. 3. Styles in caudal view. 4. Aedeagus in left

lateral view. 5. Aedeagus in right lateral view. 6. Aedeagus in ventral view.

Dimensions of eggs and nymphs are expressed in millimeters as mean ± SE. For

nymphs, length was measured from tip of vertex to tip of abdomen; width was

measured across the widest part of the body. Thoracic length was measured along

the midline from the anterior margin of the pronotum to the posterior margin of

the metanotum.

Adult (Figs. 1-6).—Muir and Gilfard (1924) described an adult male brachypter

and provided illustrations of the aedeagus. An adult male macropter is illustrated

in Figure 1 and male genitalia in Figures 2-6. Collecting data for all specimens

are: CALIFORNIA: Butte Co., Chico, 17 May 1981 (3 6), 20 May 1981 (11 3;

plus 3 6 “collected as eggs, greenhouse reared”), 27 May 1981 (4 6), 1 June 1981

(1 *).

Egg (Fig. 7). —Length 0.76 ± 0.007; width 0.19 ± 0.003; 10 specimens ex¬

amined. Eggs laid singly; each elongated, curved, subconical, narrowing at ends,

anterior end blunt, posterior end broadly rounded; white; chorion translucent,

smooth.

First instar (Fig. 8).—Length 0.76 ± 0.015; thoracic length 0.24 ± 0.005; width

0.26 ± 0.003; 9 specimens examined.

Form elongated, subcylindrical, widest across mesothorax. Body pale brownish.

Vertex subquadrate, anterior margin barely discernible, extending to or beyond

level of anterior margin of eye. Frons convex and broadly rounded in lateral view;

in frontal view, ovoid, about as wide as long, widest about midway from vertex

to clypeal border; lateral margins convex, ventral margin concave, each lateral

margin carinate (outer carina) and paralleled by a second carina (inner carina),

which originates near anterior border of vertex and extends ventrally to clypeal

border; 9 pits in 2 irregular rows between each inner and outer carina and 2 pits

PAN-PACIFIC ENTOMOLOGIST

Figures 7, 8. Immature stages of D. bellicosa. 7. Egg. 8. 1st instar. A. Nymph, B. Ventral view of

distal end of metathoracic leg. Vertical bar = 0.5 mm.

between outer carina and eye. Clypeus narrowing distally, consisting of basal

postclypeus and cylindrical distal anteclypeus. Beak 3-segmented, extending to

metacoxae; segment 1 almost obscured by anteclypeus, segments 2 and 3 subequal.

Eyes red. Antennae 3-segmented; scape ring-like, short; pedicel subcylindrical,

VOLUME 61, NUMBER 1

ca. 3 x longer than scape; flagellum bulbous basally, filamentous distally, bulbous

portion ca. 2 / 3 to % length of pedicel.

Thoracic nota divided by a longitudinal middorsal line into 3 pairs of plates.

Pronotum longest laterally, extending anteriorly to or beyond posterior margin

of eye; each plate subrectangular, posterior margin slightly sinuate, with a short,

oblique, sinuate carina extending posterolaterally from anteromedial corner; with

a single row of 6 pits. Mesonotum with median length slightly longer than that

of pronotum; each plate subquadrate, posterolateral margin convex, with 2 pits

in median half and 2 pits near lateral margin. Metanotum with median length

slightly shorter than that of mesonotum; each plate subquadrate, with 1 pit in

posteromedial corner. Pro- and mesocoxae elongated, posteromedially directed;

metacoxae smaller, obscured by cup-like trochanters. Metatibiae with 2 black-

tipped spines apically and a very short, moveable, conical, spikelike spur; spur

subequal in length to longest spine. Tarsi 2-segmented, divisions between tarso-

meres very obscure; pro- and mesotarsomere 1 somewhat wedge-shaped, meta-

tarsomere 1 with a row of 3 black-tipped spines apically; segment 2 of all legs

subconical, slightly curved, with a pair of slender apical claws.

Abdomen 9-segmented, subcylindrical, widest across segments 4 and 5; segment

9 elongated vertically, surrounding anus. Segments 1 and 2 with tergites reduced

to obscure plates; each segment with the following number of pits on either side

of midline (lateralmost and caudal pits often not visible in dorsal view): segment

5 with 1 lateral pit on tergite, segments 6-8 each with 2 lateral pits on tergites,

segment 9 with 3 caudal pits.

Second instar (Fig. 9).—Length 0.87 ± 0.003; thoracic length 0.29 ± 0.007;

width 0.39 ± 0.007; 3 specimens examined.

Body slightly dorsoventrally flattened.

Vertex subpentagonal. Frons with 3 pits between each outer carina and eye.

Antennae with pedicel bearing 2 ring-like sensoria on dorsal aspect near apex;

flagellum with bulbous portion ca. V 2 length of pedicel.

Mesonotum with each plate bearing an oblique carina originating on anterior

margin ca. ‘/ 3 distance from midline to lateral margin and extending posterolater¬

ally to posterior margin of mesonotum; with 2 pits between midline and oblique

carina and ca. 3 pits between oblique carina and lateral margin. Metanotum with

each plate bearing 2 pits. Metatibiae bearing 3 black-tipped spines apically; spur

2 x or more length of longest spine. Tarsi with divisions between tarsomeres

distinct; metatarsomere 1 with a row of 4 black-tipped spines apically.

Third instar (Fig. 10). —Length 1.03 ± 0.027; thoracic length 0.36 ± 0.007;

width 0.46 ± 0.009; 10 specimens examined.

Frons, in lateral view, convex and broadly rounded dorsally but nearly straight

vertically. Antennae with pedicel bearing 4 sensoria; flagellum with bulbous por¬

tion ca. V 3 length of pedicel.

Mesonotum with wingpads slightly lobate. Metanotum with each plate appar¬

ently bearing 1 pit. Metatibiae with 2 small black-tipped spines on lateral margin,

1 near base and 1 in basal V 3 to V 2 and a row of 5 black-tipped spines apically;

spur somewhat flattened, more elongated, with 2-3 very small black-tipped teeth

just before apex and 1 tooth at apex. Metatarsomere 1 with a row of 5 black-

tipped spines apically.

Abdominal tergites with the following number of pits on either side of midline

PAN-PACIFIC ENTOMOLOGIST

Figures 9, 10. Immature stages of D. bellicosa. 2nd instar. 10. 3rd instar. A. Nymph, B. Ventral

view of distal end of metathoracic leg. Vertical bar = 0.5 mm.

(lateralmost and caudal pits often not visible in dorsal view): segment 5 with 1

lateral pit on tergite, segments 6-8 each with 2 lateral pits on tergite, segment 9

with 3 caudal pits.

VOLUME 61, NUMBER 1

■o

0 ^

/ * A Jf

j : •' "A /

l O

O /

oi • >-——

-o ■]

1 \i°

■ • 0

_. ° FT w

jSw O . /

\vN. 1 J

’fX'/ 0

/ n.o

O \

0:o

o o y

/ °

\ 0

° \

V/ \

^ • ' • *

' o I

Tooj^-—

0 O of

AqcT^

0 ~0flT

Figures 11, 12. Immature stages of D. bellicosa. 11. 4th instar. 12. 5th instar. A. Nymph, B. Ven¬

tral view of distal end of metathoracic leg. Vertical bar = 0.5 mm.

Fourth instar (Fig. 11). —Length 1.62 ± 0.063; thoracic length 0.52 ± 0.013;

width 0.75 ± 0.043; 6 specimens examined.

Body pale with irregular light brown markings.

PAN-PACIFIC ENTOMOLOGIST

Frons with 4 pits between each outer carina and eye. Antennae with pedicel

bearing 6 sensoria; bulbous portion of flagellum ca. % length of pedicel.

Mesonotum with wingpads distinctly lobate, covering ca. V 2 of metanotal plate

laterally. Metatibial spur elongate, more flattened, with a row of ca. 8 very small

black-tipped teeth, spur extending almost to apex of tarsomere 1. Metatarsi with

tarsomere 1 bearing a row of 6 black-tipped spines apically; tarsomere 2 with a

row of 3 small black-tipped spines near middle partially subdividing tarsomere.

Fifth instar (Fig. 12). —Length 1.72 ± 0.087; thoracic length 0.57 ± 0.020;

width 0.74 ± 0.025; 8 specimens examined.

Frons with length ca. 1 V 2 x width. Antennae with pedicel bearing 9 sensoria;

bulbous portion of flagellum ca. V 6 length of pedicel.

Pronotum with each plate bearing 7 pits. Mesonotum with wingpads extending

to, or almost to, apex of metanotal wingpads laterally (specimens upon which this

description is based are assumed to be macropters, brachypters would probably

have mesonotal wingpads similar to those of 4th instars as in Megamelus davisi

Van Duzee [see Wilson and McPherson, 1981]). Metatibial spur foliose, more

elongated and flattened, with a row of ca. 15-21 very small black-tipped teeth.

Metatarsi 3-segmented; tarsomere 1 with a row of 7 black-tipped spines apically;

tarsomere 2 with a row of 4 black-tipped spines apically; tarsomere 3 similar to

apical tarsomere of earlier instar.

Acknowledgments

I thank Dr. R. Ediger, Department of Biological Sciences, California State

University, Chico (CSUC), for identifying the host plant, Dr. D. H. Kistner, CSUC,

for his helpful comments on the illustrations, and Dr. J. P. Kramer, Department

of Entomology, Smithsonian Institution, Washington, D.C., for confirming my

identification of the adult males.

Literature Cited

Muir, F. A. G., and W. M. Giffard. 1924. Studies in North American Delphacidae. Hawaiian Sugar

PI. Assoc. Bull., 15:1-53.

Wilson, S. W., and J. E. McPherson. 1981. Life history of Megamelus davisi with descriptions of

immature stages. Ann. Entomol. Soc. Am., 74:345-350.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 79-90

Checklist of Pacific Northwest Tabanidae with

New State Records and a Pictorial Key to

Common Species (Diptera, Tabanidae ) 1

William J. Turner

Department of Entomology, Washington State University, Pullman, Washing¬

ton 99164.

Abstract. — An annotated checklist of the horse flies and deer flies of Washington,

Oregon, Idaho, western Montana and southern British Columbia is presented.

New state records for Washington (10 spp.) and Idaho (2 spp.) are also given.

Pictorial keys are provided to assist in identifying females of 47 species in 7 genera

most commonly found in the Pacific Northwest region.

Philip (1965) catalogued 65 species of horse and deer flies in 9 genera from

Washington, Oregon, Idaho, and portions of Montana and British Columbia. In

the nearly 20 years since Philip’s compilation there have been slight adjustments

in the numbers of species, including additions and deletions, some synonymy and

several important range extensions. There remains a dearth of overall taxonomic

works for tabanids specifically of this region. Adult horse and deer flies occurring

in the Pacific Northwest (PNW) have been treated as portions of more extensive

revisionary studies (Brennan, 1935; Philip, 1954, 1955; Stone, 1938), in treat¬

ments of taxonomically difficult groups (McAlpine, 1961; Teskey, 1982), or simply

in descriptions of new species. Descriptions of immatures and their biologies for

many PNW species can be found in Burger (1977), Lane (1975, 1979), Middlekauff

and Lane (1980), and Teskey (1969). These references serve to identify the more

important and recent publications that include PNW tabanids.

The present PNW checklist, preliminary to a more complete study, is the result

of extensive survey/biological studies by me in Washington and recent surveys

in Idaho (Nowierski and Gittins, 1976) and Oregon (Mahmoud, 1980). Additional

work in Alberta (Thomas, 1973), California (Middlekauff and Lane, 1980) and

Illinois (Pechuman et al., 1983) has helped to complete the picture of distribution

patterns in areas adjacent to the region. As a result, a more accurate idea of the

nature and composition of the regional tabanid fauna and its relationships to

adjacent areas is emerging.

The following individuals allowed access to their collections and specimens in

their care or shared important distribution information: P. H. Amaud, Jr. and C.

B. Philip (Calif. Acad. Sci., San Francisco); J. D. Lattin and A. A. Mahmoud

(Ore. St. Univ., Corvallis); L. L. Pechuman (Cornell Univ., Ithaca); J. A. Powell

and E. I. Schlinger (Univ. Calif., Berkeley); and F. C. Thompson (USDA, ARS,

1 Scientific Paper No. SP6571, Washington State University, College of Agriculture Research Center,

Pullman. Project 0209.

PAN-PACIFIC ENTOMOLOGIST

SEA, Washington, D.C.). Their assistance is appreciated. Dr. Pechuman kindly

provided information on tabanid distribution for each western state and the

Canadian Provinces. He along with Dr. H. J. Teskey (Canad. Nat. Coll., Ottawa)

also made important comments about the particular species discussed here. Ap¬

preciation is extended to E. P. Catts for his review of the manuscript and for

testing the keys.

Checklist of Species

Recent specific name changes and descriptions of new species have produced

several new names for western species recognized as occurring here. Some of the

more important changes include the following (all new names or changes since

Philip, 1965, are denoted on Table 1 with an asterisk): Chrysops ater Macquart

now refers to western forms recognized previouly as C. carbonarius (Walker), a

species evidently restricted to eastern North America (Pechuman and Burton,

1969; Pechuman, 1981a; and Pechuman, in Middlekauff and Lane, 1980); Hy-

bomitra lurida (Fallen) supercedes H. metabola McDunnough (Pechuman and

Stone, 1968); Pechuman (1981b) synonymized Atylotus incisuralis Macquart with

A. insuetus Osten Sacken; Tabanus similis Macquart replaces western forms pre¬

viously called T. lineola Fabricius and T. lineola scutellaris Walker as does T.

tetropsis Bigot for T. productus (Middlekauff and Lane, 1980); Hybomitra philipi

Stone was synonymized as a populational variant of H. sonomensis Osten Sacken

and H. enigmatica Teskey was recognized as new and separate from related H.

sonomensis and H. phaenops Osten Sacken (Teskey, 1982); H. typha (Whitney)

was found to be restricted to eastern North America and the western forms were

described as H. pechumani Teskey and Thompson (1979).

Subspecies and variants for three species have been recognized as valid species

or evidence suggests that they should be treated as such. Hybomitra phaenops

Osten Sacken, long recognized as a subspecies or variant of H. sonomensis, was

elevated to specific rank on differences in immature stages and their habits (Mid¬

dlekauff and Lane, 1980; Teskey, 1982). Similarly, Apatolestes willistoni Brennan,

treated as a variant of A. comastes Williston, is now recognized by Middlekauff

and Lane (1980) as a separate species. The latter workers and Mahmoud (1980)

indicate that A. willistoni var. fulvipes Philip is the form occurring in the region

(see comment below). Based on morphology and distributional data, H. osburni

Hine is likely separate from H. rhombica Osten Sacken, a species which is re¬

stricted to Utah and adjacent states (Pechuman, pers. comm.).

In contrast, a number of subspecies and variants are considered here simply as

populational variants. They occur occasionally as individual morphs, melanic

forms, or clinal variants that apparently integrate broadly in color or pollinosity

characters with typical forms in contiguous populations. They have no status and

I see no need to recognize each variant. They are included here so that proper

associations with their respective species can be made. Variants occurring in the

PNW and references to their current status are as follows: Apatolestes willistoni

var. fulvipes Philip (Middlekauff and Lane, 1980); Pilimas californica var. beameri

Philip (Middlekauff and Lane, 1980); Chrysops aestuans var. abaestuans Philip

(Nowierski and Gittins, 1976); C. aestuans var. confusus Krober (listed in the

Nearctic Diptera catalog as a variant of C. callidus Osten Sacken, but Pechuman

(pers. comm.) suggests that C. callidus probably does not occur in the west; also

VOLUME 61, NUMBER 1

Table 1. List of Tabanidae of the Pacific Northwest.

ORE

WASH

IDA

MONT

Apatolestes

albipilosus Brennan

comastes Williston

*willistoni Brennan 3

comastes var. willistoni

Brennan

var. fulvipes Philip

Pilimas

californica (Bigot)

var. beameri Philip

Stonemyia

tranquilla fera (Williston)

Silvius

gigantulus (Loew)

notatus (Bigot)

philipi Pechuman

quadrivittatus (Say)

Chrysops

*aestuans van der Wulp

var. abaestuans Philip

var. pseudoconfusus Philip

callidus var. confusus Krober

asbestos Philip

*ater Macquart

carbonarius Walker

fugax Osten Sacken

carbonarius var. nubiapex Philip

bishoppi Brennan 6

var. gilvus Philip

coloradensis Bigot

discalis Williston

excitans Walker

frigidus Osten Sacken

var. xanthas Philip

fulvaster Osten Sacken

furcatus Walker

var. chagnoni Philip

mitis Osten Sacken

nigripes Zetterstedt

*noctifer Osten Sacken

noctifer pertinax Williston

proclivis Osten Sacken

var. atricornis Bigot

surdus Osten Sacken

var. piceus Philip

wileyae Philip

Haematopota

americana Osten Sacken

Atylotus

* calcar Teskey

PAN-PACIFIC ENTOMOLOGIST

Table 1. Continued.

ORE WASH BC IDA MONT

*insuetus (Osten Sacken)

incisuralis (Macquart)

tingaureus (Philip)

*utahensis (Rowe and Knowlton)

Hybomitra

aasa Philip

affinis (Kirby) b

astuta (Osten Sacken)

atrobasis (McDunnough)

californica (Marten)

captonis (Marten)

*enigmatica Teskey

epistates (Osten Sacken)

frontalis (Walker)

fulvilateralis (Macquart)

hearlei (Philip)

illota (Osten Sacken)

lanifera (McDunnough)

lasiopthalma (Macquart) b

liorhina (Philip)

*lurida (Fallen)

metabola McDunnough

melanorhina (Bigot)

nuda (McDunnough) b

opaca (Coquillett)

*osburni (Hine)

rhombica var. osburni Hine

*pechumani Teskey and Thompson b

typha Whitney, in part

pediontis (McAlpine)

*phaenops (Osten Sacken)

sonomensis var. phaenops Osten Sacken

procyon (Osten Sacken)

rupestris (McDunnough)

sequax (Williston)

*sonomensis (Osten Sacken)

*philipi Stone

tetrica (Marten)

var. hirtula Bigot

var. rubrilata Philip

trepida (McDunnough) b

zonalis (Kirby) 3

zygota (Philip) b

Tabanus

aegrotus Osten Sacken

fratellus Williston

kesseli Philip

laticeps Hine

marginalis Fabricius b

monoensis Hine b

punctifer Osten Sacken

reinwardtii Wiedemann

VOLUME 61, NUMBER 1

Table 1. Continued.

ORE

WASH

IDA

MONT

*similis Macquart

lineola Fabricius

lineola scutellaris Walker

stonei Philip

var. jellisoni Philip

*tetropsis Bigot b

productus Hine

a New Idaho state record.

b New Washington state record.

see Nowierski and Gittins, 1976); C. bishoppi var. gilvus Philip (Middlekauff and

Lane, 1980); C. frigidus var. xanthus Philip (Nowierski and Gittins, 1976); C.

furcatus var. chagnoni Philip (Thomas, 1973); C. proclivis var. imfurcatus Philip

and var. atricornis Bigot (Middlekauff and Lane, 1980); C. surdus var. piceus

Philip (Middlekauff and Lane, 1980); Hybomitra tetrica var. hirtula (Bigot) (Mid¬

dlekauff and Lane, 1980). The subspecies Chrysops noctifer pertinax Williston is

treated likewise (Middlekauff and Lane, 1980).

Total diversity of genera and species seems to vary little among the PNW states

and western Canadian province of British Columbia. All areas share the same

nine genera (if one assumes that Haematopota occurs in Oregon, although it is

not yet recorded from there). The overall diversity of species is likewise very

similar with the greatest number occurring in southern British Columbia (56) and

the least in western Montana (49). Tallies of species for other PNW states are

Washington (54), Idaho (54) and Oregon (52). British Columbia shares similar

coastal habitats and associated fauna with Washington and Oregon, but it also

receives more boreal and montane faunal influences from the north and northeast.

These northern relationships increase the diversity for British Columbia as several

species do not extend farther south into the states. The low number of species

in Oregon is surprising as one would expect its southern portion to be influenced

by California to the south. I suspect that Oregon, Montana and Idaho may show

greater diversity in the final analysis, but their fauna has been insufficiently col¬

lected.

The limited distribution of some species may reflect a lack of collecting or

actually represent species that are geographically limited. Four Chrysops species

(C. ater, C. frigidus Osten Sacken, C. fuly aster Osten Sacken, and C. nigripes

(Zetterstedt)) are known from the eastern portions of Montana and Idaho, but

they have not been collected or have questionable records in Washington and

Oregon. Tabanus reinwartii Wiedemann likewise is a midwestern species that has

invaded the eastern foothills of the Rocky Mountains in Montana but no farther

west. While many species have wide distributions throughout the PNW region,

others with similar patterns have not been recorded in certain areas. For example,

it is difficult to explain the following absences: Atylotus tingaureus (Philip) from

Idaho; Tabanus marginalis Fabricius, T. punctifer Osten Sacken, Hybomitra la-

nifera (McDunnough) and H. lurida from Montana; H. pechumani, H. lurida, H.

opaca (Coquillett), and H. zonalis (Kirby) from Washington; Tabanus tetropsis

PAN-PACIFIC ENTOMOLOGIST

TABANIDAE: PICTORIAL KEY TO COMMON GENERA OCCURRING IN

THE PACIFIC NORTHWEST (BASED ON ADULT FEMALES)

WILLIAM J. TURNER

1983

Hind leg with 2

tibial spurs

(Subfamilies Pangoniinae

and Chrysopsinae)

Hind leg without

tibial spurs

(Subfamily Tabaninae)

Frontal callus touching

eye margin

Frontal callus not

touching eye margin

hpatoleatea (3 PNW spp.,

(Subfamily Chrysopsinae)

Chryeopa (16 PNW Spp.,

15 common; see key to

Chryaopa species)

Abdomen orange

Silvius gigantulua

ms giai.

(Loew)

Silvius (4 PNW spp.,

3 common)

Abdomen gray

First antennal segment

swollen and longer

than high

Haematopota (1 PNW spp.

H. americana Os ten

Sacken)

Vertex with distinctly

raised and shining

tubercle

Hybomitra (31 PNW spp.,

17 common; see key to

Hybomitra species)

Basal callus broad and

touching eyes or close

to eye margins

Tabanua (11 PNW spp.,

8 common; see key to

Tabanu8 species)

Wing veins with spots

Wing veins without

(Say)

Silvius notatus

(Bigot)

First antennal segment

short and barely

longer than high

Vertex flat and without

raised tubercle (may

be bare and shining)

Basal callus small and

not touching eye margins

Atylotue (4 PNW spp.,

1 common: A. insuetus

Osten Sacken)

Figure 1. Pictorial key to common genera occurring in the Pacific Northwest (based on adult

females).

from British Columbia; and Haematopota americana Osten Sacken from Oregon.

Finally, it is equally difficult to explain the very restricted and perhaps relict

population of H. astuta (Osten Sacken). Reported from one Oregon locality (Mah¬

moud, 1981), it has not been collected in any other PNW area.

Several primarily northern and eastern species have been recorded from north¬

ern Idaho, northern Washington and British Columbia. They are usually en-

VOLUME 61, NUMBER 1

TABANIDAE: PICTORIAL KEY TO COMMON PACIFIC NORTHWESTERN

SPECIES OF CHRYSOPS (BASED ON ADULT FEMALES)

WILLIAM J. TURNER

1983

Wing clouded beyond crossband

-1

Wing clear beyond crossband

Basal callus entirely black Basal callus mostly brownish

orange

2nd basal cell dark over 1/2

its length

Chrysopa nootifer

Osten Sacken

Chrysops excitana

Walker)

-1

Cu^ cell entirely black basally

Chrysops mitis

Osten Sacken

—1

Basal antennal segment not

swollen

Apical spot ending in R. cell; triangular

hyaline area beyond dark crossband

complete to hind margin of wing

Apical spot continuing into R^ cell and

beyond, usually enclosing hyaline

area beyond dark crossband

(Continued on next page)

Chrysopa wileyae

Philip

Chrysopa fulvaster

Osten Sacken

Figure 2. Pictorial key to common Pacific Northwest species of Chrysops (based on adult females).

countered at higher latitudes in the west. One-half (5 of 10) of the species reported

here as new state records for Washington show this northeastern influence in the

region. These include: Hybomitra affinis (Kirby), H. lasiopthalma (Macquart), H.

nuda (McDunnough), H. trepida (McDunnough) and Tabanus marginalis. Other

new Washington records for Chrysops bishoppi and Tabanus monoensis Hine

represent species that are widely distributed but only recently collected in the

state. The remaining two species new to Washington {Hybomitra zygota (Philip)

and Tabanus tetropsis) have been collected in adjacent areas and represent antic-

PAN-PACIFIC ENTOMOLOGIST

TABANIDAE: KEY TO PACIFIC NORTHWESTERN

CHRYSOPS SPECIES CONTINUED

WILLIAM J. TURNER

1983

(Continued from previous page)

2nd basal cell clear or darkened

only at apex and base

Chrysops disaalis

Williston

Face black with

pollinose stripe

along midiine

Face yellow without

pollinose stripe

along midline

Chrysops aestuans

van der Wulp

Front coxae yellow Front ccxae black

Chrysops bishoppi

Brennan

I—

Abdominal tergites 3-6

with black markings

entire behind

Abdominal tergites

with black marki

scalloped behind

3-6

ngs

Chrysops furcatus

Walker

Chrysops asbestos

Philip

Chrysops frigidus

Osten Sacken

Face broadly yellow

between lateral

lobes

Face mostly black with

narrow median yellow

stripe

Chrysops proclivis

Osten Sacken

Chrysops surdus

Osten Sacken

Figure 3. Key to Pacific Northwest Chrysops species continued.

ipated range extensions into Washington. The two new Idaho records are for

Apatolestes willistoni Brennan and Hybomitra zonalis. The last is another northern

species that eventually should be collected in northeastern Washington.

Pictorial Keys

Among important blood-sucking Diptera, the horse and deer flies represent one

group that is easily recognized by the non-specialist. Unfortunately, the body of

literature available for identifying tabanids is written primarily for the taxonomic

VOLUME 61, NUMBER 1

TABANIDAE: PICTORIAL KEY TO COMMON PACIFIC NORTHWESTERN

SPECIES OF HYBDMITRA (BASED ON ADULT FEMALES)

WILLIAM J. TURNER

Abdomen black, 2nd segment lacking pale dorsal

sublateral spots and faint reddish mark at

side (may be gray pollinose at side and behind)

Subcallus pale pollinose

and dul1

Subcallus bare, black

and shining

Abdomen broadly orange brown or dark and 2nd

segment with pale dorsal sublateral spots

(triangles), faint reddish mark at side or both

Hybomitra sequax

(Williston)

Hybomitra procyon

(Osten S'acken)

Abdomen mostly black at sides, pale

s-pots margined in black or reddish

Abdomen broadly orange brown at

sides, without black margins

R. stump vein

4 absent

stump vein

present

Hind tibia with long

golden fringe

Hind tibia with black

fringe or none

Subcallus pollinose

and dull

Hybomitra tetrica

(Marten)

Subcallus bare and

shining

Notopleural lobe

yellowish brown

Hybomitra melanorhina

(Bigot)

Notopleural lobe

black

(Continued on next page)

Hybomitra califomica

(Marten)

Antenna mostly block

and 3rd segment narrow

Antenna mostly orange

and 3rd segment thick

Eye bare or with few

Hybomitra sonomensis

(Osten Sacken)

Hybomitra phaenops

" (Osten Sacken)

and

Hybomitra enigmatica

Teskey

(Continued on next page)

Figure 4. Pictorial key to common Pacific Northwest species of Hybomitra (based on adult females).

specialist. The keys provided here represent one attempt to develop taxonomic

works for more general use on this important group of biting flies. In format the

keys are modeled after the CDC series of keys to arthropods and other animals

of medical importance (U.S. Dept. Health, Educ., Welfare, 1967). These keys are

easier to use than more traditional ones which are often limited to word descrip¬

tions and a plethora of technical terms. In all cases where precise identifications

are necessary, determinations should be considered tentative until checked by a

specialist.

The included keys have been limited to include the more common species and

PAN-PACIFIC ENTOMOLOGIST

TABANIDAE; KEY TO PACIFIC NORTHWESTERN

HYBOPIITRA SPECIES CONTINUED

WILLIAM J, TURNER

1983

(Continued from previous page)

Subcallus pollinose and dull

(Continued from previous page)

Notopleural lobe black

Antenna mostly orange and

3rd segment thick

Abdominal spots yellow

and nearly round

Hybonritra frontalis

(Walker)

Abdominal spots yellowish

brown and oblique

2nd abdominal tergite

mostly black at sides

2nd abdominal tergite

orange or pink at sides

Hybonritra opaca

(Coquillett)

Hybonritra osbumi

(Hine)

Hybonritra rupestrie

(McDunnough)

Subcallus pollinose

and dull

Subcallus bare and

shining

Hybonritra fulvilateralis

(Macquart)

Hybonritra epistates

(Osten Sacken)

Hybomitra aasa

(Philip)

Hybonritra captonis

(Marten)

Figure 5. Pictorial key to common Pacific Northwest Hybomitra species continued.

genera of tabanids. Whole groups were excluded because they are considered

uncommon (i.e., those that are rarely encountered or poorly collected; for example,

species of Stonemyia and Pilimas). Species were judged to be “common” after

reviewing published and unpublished distribution records, past requests for de¬

terminations or biological information, and lists of data collected from museum

specimens or associated with field survey samples and studies from the region.

In some keys, however, several less common biting species have been included

because either the groups were small (e.g., Silvius ) or the similar-appearing rarer

species might be confused with strictly common ones (e.g., some species of Chry-

VOLUME 61, NUMBER 1

TABANIDAE: PICTORIAL KEY TO COMMON PACIFIC NORTHWESTERN

SPECIES OF TABANUS (BASED ON ADULT FEMALES)

WILLIAM J. TURNER

1983

Abdomen not completely black but with

pale dorsal sublateral spots, median

line (or triangles), or both

R 4 stump vein present

fL stump vein absent

Abdomen black and without pale dorsal

sublateral spots or median line (may

have tufts of white hair at midline)

Scutum with dark hair

Scutum with pale hair

Abdominal tergites (at least

1st) each with small tuft

of white hair along midline

Tabanua kesseli

Philip

Eye bare and without

minute hairs (at 30x)

Eye minutely haired

Abdominal tergites without

any tufts of white hair

along midline

Tabanus aegrotus

Osten Sacken

Hind tibia with dark

outer fringe

Hind tibia with pale

outer fringe

Tabanus latioeps

Hine

Tabanu8 atonei

Philip

Figure 6. Pictorial key to common Pacific Northwest species of Tabanus (based on adult females).

sops; also Hybomitra). The keys also are restricted to females as most records and

identification requests involve females that are regularly attracted to humans,

animals or baited traps. Males of all species, in contrast, do not seek animal hosts

or suck blood and are therefore rarely encountered.

In one couplet of the key to the species of Hybomitra, I have not separated two

species, H. phaenops (Osten Sacken) and H. enigmatica Teskey. Both are com¬

mon, very similar and not readily identified without rehydrating the heads in

order to restore the prominent eye bands visible in living material (see Price and

Goodwin, 1979 for technical details of the method). In H. phaenops the dark

PAN-PACIFIC ENTOMOLOGIST

horizontal bands are wider and extend completely to the lateral eye margin while

the upper and lower margins are darkened. In contrast, the bands in H. enigmatica

are thinner and do not reach the eye margin while the darker margins above and

below are absent. Specimens keyed to H. phaenops/enigmatica should be referred

to a specialist for positive identification.

Literature Cited

Brennan, J. M. 1935. The Pangoniinae of Nearctic America, Diptera: Tabanidae. Univ. Kans. Sci.

Bull., No. 22, pp. 249-401.

Burger, J. F. 1977. The biosystematics of immature Arizona Tabanidae (Diptera). Trans. Am.

Entomol. Soc., 103:145-258.

Lane, R. S. 1975. Immatures of some Tabanidae (Diptera) from Mendocino County, Calif. Ann.

Entomol. Soc. Am., 68:803-819.

-. 1979. Larvae and pupae of two Hybomitra species (Diptera: Tabanidae) from northern

California. J. Med. Entomol., 16:142-149.

Mahmoud, A. A. 1980. The Tabanidae (Diptera) of Oregon. Ph.D. dissertation, Ore. St. Univ.,

Corvallis, 269 pp.

McAlpine, J. F. 1961. Variation, distribution and evolution of the Tabanus ( Hybomitra ) frontalis

complex of horseflies (Diptera: Tabanidae). Can. Entomol., 93:894-924.

Middlekauff, W. W., and R. S. Lane. 1980. Adult and immature Tabanidae (Diptera) of California.

Bull. Calif. Insect Surv., 22:1-99.

Nowierski, R. M., and A. R. Gittins. 1976. The horse flies and deer flies of Idaho. Univ. Ida. Exp.

Sta., Res. Bull. 96, 48 pp.

Pechuman, L. L. 1981a. The horse flies and deer flies of New York (Diptera, Tabanidae). 2nd ed.

Search: Agric., 18:1-66.

-. 1981b. Notes on Atylotus and description of a new species from eastern North America

(Diptera: Tabanidae). Entomol. News, 92:1-6.

-, and J. J. S. Burton. 1969. Seasonal distribution of Tabanidae (Diptera) at Texas Hollow,

New York in 1968. Mosq. News, 29:216-220.

-, and A. Stone. 1968. A new synonymy in Hybomitra. Proc. Entomol. Soc. Wash., 70:302.

-, D. W. Webb, and H. J. Teskey. 1983. The Diptera or true flies of Illinois. I. Tabanidae.

Bull. Illin. Nat. Hist. Surv., 33:1-122.

Philip, C. B. 1954. New North American Tabanidae. VIII. Notes on and keys to genera and species

of Pangoniinae exclusive of Chrysops. Rev. Brasil. Entomol., 2:13-60.

-. 1955. New North American Tabanidae. IX. Notes on and keys to the genus Chrysops Meigen.

Rev. Brasil. Entomol., 3:47-128.

-. 1965. Family Tabanidae. Pp. 319-342 in A. Stone, C. W. Sabrosky, W. W. Wirth, R. H.

Foote, and J. R. Coulson (eds.), A catalog of the Diptera of America north of Mexico. A.R.S.,

U.S. Dept. Agric., Handbk. 276, 1696 pp.

Price, J. O., and J. T. Goodwin. 1979. The taxonomic significance of eye pattern in female Tabanidae.

Ann. Entomol. Soc. Am., 72:725-734.

Stone, A. 1938. The horseflies of the subfamily Tabaninae of the Nearctic Region. Misc. Publ. U.S.

Dept. Agric., No. 305, 171 pp.

Teskey, H. J. 1969. Larvae and pupae of some eastern North American Tabanidae (Diptera). Mem.

Entomol. Soc. Can., No. 63, 147 pp.

-. 1982. A new species of the Hybomitra sonomensis group (Diptera: Tabanidae) with descrip¬

tion of its immature stages. Can. Entomol., 114:1077-1082.

-. 1983. A review of the Atylotus insuetus group from western North America including

description of a new species and immature stages (Diptera: Tabanidae). Can. Entomol., 115:

693-702.

-, and A. W. Thomas. 1979. Identity of Hybomitra typhus and description of a new species

previously confused with it (Diptera: Tabanidae). Can. Entomol., 111:343-350.

Thomas, A. W. 1973. The deer flies (Diptera: Tabanidae: Chrysops ) of Alberta. Quaes. Entomol.,

9:161-171.

U.S. Dept. Health, Education and Welfare. 1967. Pictorial keys to arthropods, reptiles, birds and

mammals of public health significance. Pub. Health Serv., Comm. Disease Center, 192 pp.

PAN-PACIFIC ENTOMOLOGIST

61(1), 1985, pp. 91-94

A New Genus and Species of Horse Fly

(Diptera: Tabanidae) from Bolivia 1

Richard C. Wilkerson

Research Associate, Florida State Collection of Arthropods, Division of Plant

Industry, Florida State Department of Agriculture and Consumer Services, P.O.

Box 1269, Gainesville, Florida 32602.

The following description of a new genus and species presents another example

of the continuing discovery of neotropical horse fly diversity. It indicates to me

the high potential of finding many more unstudied higher taxa in this part of the

world. It is with reluctance that I would describe a new species from a single

specimen, let alone a new genus, but in this case it is quite clear that the species

is distinct and requires generic status.

Roquezia Wilkerson, New Genus

(Tabanidae: Tabaninae: Diachlorini)

Diagnosis. — Frons relatively wide, convergent below. Basal callus square and

protuberant, not touching sides of frons. Eyes bare. A slightly raised tubercle at

vertex bears 3 small ocelli. Antennal basal plate slender, dorsally bluntly rounded,

style 4 annulate. Palpus short, greatly inflated but sharply pointed. Scutum pale

gray and dark brown striped. Wings glass-clear, venation normal, vein R 4+5 with

a short appendix. Basicosta without macrotricheae (bare). Abdomen above black¬

ish brown with a series of large sublateral pale gray spots on tergites 3-6. Tergites

1 and 2 largely gray pollinose.

The overall impression is of a muscoid fly.

Type of genus. —Roquezia signifera Wilkerson, new species.

Etymology .—Named for Roberto Vasquez C., Bolivian botanist and generous

host during my visits to Bolivia.

Roquezia signifera Wilkerson, New Species

(Figs. 1-4)

Diagnosis.— A small muscoid appearing dark brown and pale gray species with

shiny square basal callus, small but distinct ocellar tubercle and short, swollen,

pointed palpi. Scutum pale gray striped, wing glass-clear and legs black. Abdomen

above with large pale gray sublateral spots except on tergite 2 which, except for

a median dark brown triangle is entirely pale gray.

Female.— Length: body 8 mm; of wing 9 mm. Head structures in Figures 1-3.

Frontal index 5.0. Frons narrowed below, width at vertex/width at base = 1.7.

Frons, subcallus, gena, frontoclypeus and postocciput pale gray pollinose with a

faint yellowish tint. Vertex darkened, area around the tubercle shiny black; dorsal

to this shiny area integument dark and sparsely dark brown pollinose. Frontal

1 Florida Agricultural Experiment Station Journal Series No. 5805.

PAN-PACIFIC ENTOMOLOGIST

Figures 1-3. Head structures of female Roquezia signifera Wilkerson, new species. 1, Antenna. 2,

Palp. 3, Frons.

hairs mostly black, short and sparse; those at vertex longer and more numerous;

those on lower Vi of frons quite sparse and pale. Ocellar tubercle small and slightly

raised bearing 3 small yellowish brown ocelli. Ocelli seem to be covered by

pollinosity and are not protuberant. Eyes bare, color and pattern not recorded.

Basal callus a protuberant shiny black square with a slender upward extension.

Subcallus with small sublateral bare spots, apparently the result of rubbing. Nu¬

merous short pale yellow hairs present on gena, less dense laterally on fronto-

clypeus. Beard of numerous short pale yellow hairs. Antenna pale reddish brown,

the annuli darker brown; antennal hairs black. Second palpal segment short, much

swollen basally but apically acutely pointed; integument pale yellowish brown,

sparsely pale gray pollinose and black haired except for a few pale hairs beneath.

Proboscis short, wholly fleshy and black. In this specimen it is retracted but stylets

are short and reach only to tip of palpus. Labella about as long as palpus.

Scutum and scutellum as in Figure 4. Pale areas of scutum pale gray pollinose,

the dark areas with a dark brown integument covered with sparse dark brown

pollinosity. Scutellum blackish brown. Scutal hairs short, sparse and black; hairs

of scutellum also sparse and black but longer. Pleura, and mid and hind coxae,

blackish brown overlain with sparse pale gray pollinosity. Pleural hairs short,

sparse and black; those below wing base longer. A small clump of yellow hairs

VOLUME 61, NUMBER 1

Figure 4. Roquezia signifera Wilkerson, new species. Dorsal view of thorax and abdomen.

present in front of wing base. Fore coxae mostly pale yellowish gray pollinose and

pale yellow haired. Legs black and black haired. Wing including costal cell glass-

clear, venation normal; R 4+5 (fork of 3rd vein) with a short appendix. Basicosta

bare. Squamae pale, with a fringe of white hairs. Halteres brown but tip of knob

yellow.

PAN-PACIFIC ENTOMOLOGIST

Abdominal dorsum as in Figure 4, dark areas blackish brown, pale areas pale

silvery gray pollinose. Dorsum wholly sparsely short black haired. Venter dark

blackish brown in ground color, pale bluish gray pollinose and wholly short black

haired.

Male. — Not known.

Type. — Holotype female, Santa Cruz Department, Bolivia, 10 km E Warnes,

22-VIII-1983, R. Wilkerson coll. To be deposited in the Florida State Collection

of Arthropods, Gainesville, Florida.

Etymology. — Signifera is from the Latin meaning bearing marks, figures or signs.

Discussion. — Tribe Diachlorini (Tabaninae, Tabanidae) remains a confusing

group of genera and subgenera whose relationships are not yet understood (Fair-

child, 1969). The tribe comprises more than half of the neotropical species of

Tabanidae and is united, perhaps artificially, by the following: “Basicosta smooth¬

ly pollinose, without setae. If setae are present, sparse or numerous, and there are

vestiges of ocelli, a strong tubercle at vertex, labella partly sclerotized, a long tooth

on third antennal segment, wings strongly patterned or other striking specializa¬

tions” (Fairchild, 1969).

Roquezia belongs in this assemblage by virtue of its bare basicosta. Several

species or species groups in Diachlorini superficially resemble Roquezia because

of their muscoid fly appearance (mimicry?). Stypommisa marucii (Fairchild) and

most Philipotabanus (Mimotabanus ) Fairchild differ by having a clavate callus

and elongated palpi. In addition they have bluish pruinosity on the first 2 ab¬

dominal segments and P. Mimotabanus has a patterned wing. Myiotabanus Lutz

differs by having a longer proboscis, partly sclerotized labella, partially bare sub¬

callus, a large ovoid basal callus, no vestiges of ocelli and short stubby antennae.

Eutabanus pictus Krober differs by having protuberant and conical notopleural

lobes, inflated fore tibiae, flattened hind tibiae, broad divergent frons, large drop¬

shaped callus and a small discal wing band.

Roquezia seems to be most closely related structurally to Stenotabanus Lutz,

many species of which have glass-clear wings, short fleshy labella, short, basally

swollen palpi, similar antennae, vestiges of ocelli, frons convergent below, and

protuberant square basal callus, sometimes with a thin dorsal extension.

Roquezia, however, differs in 2 significant ways from any Stenotabanus known

to me. First, the frons differs by having a basal callus that does not touch the

sides of the frons and there is no median darkened area on the frons evident in

most St. (Stenotabanus ). Secondly, no Stenotabanus seen has a thoracic and/or

abdominal pattern similar to Roquezia.

Acknowledgments

I wish to thank G. B. Fairchild for initial review of this manuscript and Dorie

Peyton for the fine drawings of the thorax and abdomen. This publication based

on work supported by the National Science Foundation under Grant BSR 8211808.

Literature Cited

Fairchild, G. B. 1969. Notes on neotropical Tabanidae. XII. Classification and distribution, with

keys to genera and subgenera. Arq. Zool., Sao Paulo, 17(4): 199-255.

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Vol. 61

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THE

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WEEKS, L. and H. A. HESPENHEIDE—Predatory and mating behavior of Stichopogon (Dip-

tera: Asilidae) in Arizona. 95

RUST, R. W.—Notes on the biology of Pseudocotalpa giulianii Hardy (Coleoptera: Scara-

baeidae).... 105

HEPPNER, J. B.—Review of the North American genus Araeolepia (Lepidoptera: Plutellidae)

.._. 110

POINAR, G. O., JR .—Silpha aenescens Casey (Silphidae: Coleoptera) as a pest of home grown

strawberries in California. 118

CHANDLER, D, S.—New synonymy in Notoxus (Coleoptera: Anthicidae). 121

EICKWORT, G. C.—The nesting biology of the sweat bee Halictus farinosus in California,

with notes on H. ligatus (Hymenoptera: Halicitidae). 122

ANNOUNCEMENT. 13 8

GUNTHER, K. K.—A new pygmy mole grasshopper from California and Baja California,

Mexico (Orthoptera: Tridactylidae). 139

POLHEMUS, D. A.—A review of Dichaetocoris Knight (Heteroptera: Miridae): New species,

new combinations, and additional distribution records. 146

PARKER, F. D.—Nesting habits of Osmia grinnelli Cockerell (Hymenoptera: Megachilidae). 155

WHARTON, R. A.—A new species of Mallochia (Hymenoptera: Ichneumonidae) introduced

to Texas to control Eoreuma loftini (Dyar) (Lepidoptera: Pyralidae) in sugar cane. 160

POLHEMUS, J. T. and D. A. POLHEMUS—Studies on Neotropical Yeliidae (Hemiptera)

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introduction, and spread. 170

HAAS, G. E., N. WILSON, and T. RUMFELT—Bird fleas, genus Ceratophyllus, from Alaska

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MIDDLEKAUFF, W. W. — Ophrella, a new genus of Orussidae from Panama (Hymenoptera:

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