Vol. 52 JULY 1976 No. 3

THE

PAN-PACIFIC ENTOMOLOGIST

GARRISON—Multivariate analysis of geographic variation in Libellula luc-

tuosa Burmeister (Odonata: Libellulidae) — 181

POLHEMUS—Notes on North American Nepidae (Hemiptera: Heteroptera) 204

CHENG—A new species of Hermatobates (Hemiptera: Heteroptera) 209

YOUNG—A new species of Ischalia from southeastern China (Coleoptera:

ey MOGIMOLG AC): (aie anon OIE eS eT hie ee te tt a 213

CHEMSAK and LINSLEY—The genus Triacetelus Bates (Coleoptera:

Geranthycidae), seo wer aie Se pan | hl ee 216

POTTS—New species of North American Anomala (Scarabaeidae: Anoma-

LLTTAC)) een ct et AE Mache oR. ae inbleh tn eee ee Soe oe, SI SNe 220

LEWIS—A new species of Evondinus from Wyoming (Coleoptera: Cerambyc-

TG EEVE:)) | yee SR cen ok IT, A Pibal EP GO Rae LORNA SLT sien OO, ck 227

COVILLE—Predatory behavior of the spider wasp, Chalybion californicum

CElymenoptera st ophecidae) mae tens EB A 229

TURNER—Fannia thelaziae, a new species of eye-frequenting fly of the ben-

Jamini group from California and description of F. conspicua female

GWintera-ye Viuscidac ea" 48-20 ot NS ERT Ras i oe ae 234

WIRTH—A new species of brine fly from California rice fields (Diptera:

Folin nrclae) a mee wales. OSC ale Ohad) WP lated Wis Sn 242

ALEXANDER—Additions to the crane flies of California (Diptera: Tipulli-

CLA) Veet eee oe IE a ee oe PR wide AN Oey bated Te ke ate eo oh 244,

GOEDEN AND RICKER—Life history of the ragweed plume moth, Adaina

ambrosiae (Murtfeldt), in southern California (Lepidoptera: Ptero-

jo) TCD Ve ESS ek ee Ao ee. OS ae ee ee a. ads ae 251

HEPPNER—A new Choreutis from the San Francisco Bay Area, California

(Lepidoptera: Glyphipterygidae) — 256

SS CoRR aIPTEC IG AN DIGS i aes Dae edie Ie Me ie Lp BT ne 203, 212, 262, 264

Oe ele els eACIU His nob. 2 ce eer eNO eee oe 228, 243, 250

TOU CIS ARNE Je, 2c ea ge eR ls 1D bc, 233

ZOOTOCIGATANG® VEBIN GIMAUR WR Eien ee de 241

TAs OEP NOME CLs eee Loe een 2 Ue ey ae OI Ae ee 264

SAN FRANCISCO, CALIFORNIA ¢ 1976

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

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The Pan-Pacific Entomologist

VoL. 52 JuLty 1976 No. 3

Multivariate Analysis of Geographic Variation

in Libellula luctuosa Burmeister

(Odonata: Libellulidae)

Rosser W. GARRISON

University of California, Berkeley

The study of intraspecific variation in dragonflies has rarely pro-

ceeded beyond the naming and describing of new taxa. This is partly

due to the paucity of material at hand: dragonflies are seldom collected

and the ranges even of many common species are poorly known. Most

of the papers analyzing geographic variation in odonates (Bennefield,

1965; Huggins, 1927; Johnson, 1972) have involved tabulation of

characters, with taxonomic decisions usually based on observable dif-

ferences in features such as color, size, shape, or wing maculation.

Multivariate methods of classification may be extremely helpful to the

odonate systematist, especially when populations intergrade and varia-

tion among different characters is discordant. This paper analyzes

geographic variation in a common and widespread species and compares

the results of four multivariate methods of analysis for consistency.

Libellula luctuosa Burmeister is a large, dark dragonfly with black

basal wing bands which occurs over most of the United States except

the Great Basin and Florida. It occurs in British Columbia, Quebec,

Manitoba, and Nova Scotia and is known in Mexico from Chihuahua

and Durango. Variation in coloration, maculation, or size might be

expected, since it ranges from mesic northeastern deciduous forests to

cattle ponds of the hot, arid Southwest.

Three different phenotypes of L. luctuwosa were subjectively recognized

by the author; their distribution is shown in Figure 1. From southern

Canada and the Atlantic coast to the eastern edge of the Great Plains,

L. luctuosa is characterized by uniformly dark hind wing bands (Fig.

4a). From the edge of the Great Plains south through Oklahoma, east-

ern Colorado, Texas, New Mexico, southeastern Arizona, and Mexico,

populations possess various degrees of hind wing clearing (Fig. 4a—d).

Hagen (1861) described these paler individuals as a new species,

Libellula odiosa, but subsequently (1875) reduced L. odiosa to a race

Tue Pan-Paciric ENTOMOLOGIST 52: 181-203. JuLy 1976

182 THE PAN-PAcIFIC ENTOMOLOGIST

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Fic. 1. Map of specimen localities, Libellula luctuosa. Solid squares = eastern

luctuosa morphs, solid triangles = odiosa morphs, open circles = Colorado River

phenotypes. More than one specimen may have been taken at some localities.

of L. luctwosa. Most authors since have considered L. odiosa a variety

or synonym of L. luctuosa (Calvert, 1906; Needham and Westfall, 1955;

Ris, 1910). These individuals are hereafter called odiosa, though the

taxon is believed to be synonymous with eastern L. luctuwosa. The odiosa

Fic. 2. Coded examples for wingtip coloration width and length. a = present,

b = 1 (absent), c = 3,d = 5, e = 8, f = 9 (brown to midpoint of pterostigma).

VoL. 52, No. 3, Juty 1976 183

Fic. 3. Thoracic patterns of Libellula luctuosa. a. male Colorado River morph

(Blythe, Riverside Co., Calif.), b. female Colorado River morph (Blythe, Riverside

Co., Calif.), c. male (Pefa Blanca Lake, Santa Cruz Co., Ariz.), d. female (Pena

Blanca Lake, Santa Cruz Co., Ariz.). Both sexes of the odiosa and luctuosa morphs

have thoracic patterns as in c. and d.

phenotype intergrades with nominate L. luctuosa west of the Mississippi

River and in the Edwards Plateau region of central Texas. Allopatric

populations of the odiosa phenotype also occur within the Central

Valley and foothills of the Sierra Nevada of California north of the

Tehachapi Mountains to southern Oregon. A single male from Robson,

British Columbia, has no clearing in the hind wing band and is classified

in this study as nominate L. luctuosa.*

A third group of populations inhabits the Sonoran, Colorado, and

Mojave deserts and is separated from the Central Valley populations by

the Tehachapi Mountains, and from the southeastern Arizona popula-

tions by the eastern edge of the Sonoran Desert. This phenotype, known

1 Both the British Columbia (American Museum of Natural History) and Oregon males (H. G. Dyar

collection at the U.S. National Museum) are probably mislabeled. The Oregon locale is Crater Lake,

29 July 1920. Libellula luctuosa has been taken at the Klamath River immediately south of the

Oregon border.

184 THE Pan-PaciFic ENTOMOLOGIST

Fic. 4. Coded examples for hindwing fenestration. a = 1 (no clearing), b = 3,

c = 8, d = 12 (maximum extent of hindwing clearing).

hereafter as the Colorado River (or desert) morph, has the thorax com-

pletely pruinose blue in males (Fig. 3a), while pruinosity is localized on

the mesepisternum in luctuosa and odiosa phenotypes (Fig. 3c). Mature

females possess a violaceous pruinosity over the typical brown areas

found in odiosa and luctuosa (Fig. 3b and 3d). The hind wing bands

are more fenestrated than those of most odiosa, and the females always

possess dark brown wing tips (Fig. 2e and 2f), sometimes present in

odiosa and luctuosa females. The pale color of individuals from desert

areas agrees with Gloger’s rule (Mayr, 1963).

The author subjected the data from nearly 200 Libellula luctuosa,

intuitively classified by the characters discussed above and enumerated

in Table 1, to the following multivariate means of classification: 1)

principal component analysis, 2) step-wise discriminant analysis, 3)

linear discriminant analysis, and 4) hierarchical numerical taxonomic

methods. Libellula luctuosa, with its broad spectrum of geographic

variation and widespread distribution (Fig. 1), is a suitable species for

this study. The purpose of the study was to test compatibility of results

of various methods and to compare those results with the author’s

intuitive concept of recognizing the species as three geographical enti-

ties, as well as to describe and explain patterns of morphological varia-

tion.

METHODS AND MATERIALS

Variation in L. luctwosa was analyzed using 116 males and 72 females

from 26 states and provinces shown in Figure 1. A total of 10 specimens

(five males and five females), if available, was chosen from each state.

More individuals were chosen from Texas due to its size, and from

California and Arizona because the Colorado River morph is apparently

restricted to those two states.

VoL. 52, No. 3, Juty 1976 185

TABLE 1. Characters.

A. Measured Characters

B.

Ld

. HINDWING: Linear distance from humeral plate to tip of wing.

PTEROSTIGMA: Linear distance along costal margin of hindwing.

INTERPOLATED ANKLE CELLS IN ANAL LOOP: Number of en-

closed cells between veins Ai and Az but not adjacent to those veins.

Ankle cells lie on either side of the midrib.

GAFF: Linear distance between hind angle of hindwing triangle and

heel of anal loop.

SOLE: Linear distance from heel to toe of anal loop.

MIDDORSAL THORACIC STRIPE WIDTH: Width measured near

the antealar sinuses. Absence of stripe, as in Colorado River phenotype

males, = 1; for other individuals, a value of 1 was added to the width

measurement.

METATHORACIC FEMUR: Length along ventral side.

METATHORACIC TIBIA: Length along ventral side from distal end

to concavity before articulation point.

ABDOMINAL SEGMENT 5: Length along lateral carina.

SUPERIOR CAUDAL APPENDAGES: Length along dorsal surface of

superior caudal appendages of males or cerci of females.

Coded Characters

11.

12.

13.

14.

15.

16.

Ld,

WINGTIP COLORATION WIDTH: Brown in anterior one-fourth of

wingtip. Coded 1 (absent) (Fic. 2b) or 2 (present) (Fic. 2a).

Present only on some male Colorado River morphs.

WINGTIP COLORATION LENGTH: Length of brown on wingtip.

Character states ranged from 1 (no brown) to 9 (brown to midpoint

of pterostigma) (Fic. 2b-f).

THORACIC COLORATION: Absence (1) or presence (2) of pale

mazarine (pruinose) blue on sides of synthorax (mesepimeron, mete-

pisternum, metepimeron). Present only on male Colorado River pheno-

type (Fic. 3a).

FOREWING FENESTRATION: Absence (1) or presence (2) of

clearing within brown forewing band of males. As almost all individuals

possessed some degree of clearing, this character was included to test

its importance relative to other characters. The light area in the mid-

basal space in fore- and hindwings is typical of all populations and was

not included.

HIND WING FENESTRATIONS: Degree of fenestration in the hind-

wing band, coded 1 (no fenestration posterior to mid-basal space) to

12 (hindwing clear from midbasal space to two or three cells anterior

to toe of anal loop).

FRONS: Color coded: 1) maize yellow, 2) aniline yellow, 3) Isabella

color, dark olive buff, or deep chrome, 4) olive brown or dark olive,

and 5) metallic black. Frons coded by the color covering 50% or more

of its surface.

ANTECLYPEUS: Color coded as above.

186 THE PAn-Paciric ENTOMOLOGIST

(TaBLE 1. Cont.)

18. LABIUM: Color coded as above.

19. MIDDORSAL |THORACIC STRIPE COLOR: Coded as for frons with

one additional color state: 6) pale mazarine (pruinose) blue.

20. PROTHORACIC FEMUR: Color of ventral sides coded as for mid-

dorsal thoracic stripe color.

Appropriate portions of the U.C.L.A. BIOMED program series as

modified for the U.C.B. computer center were used in this study. Pat-

terns of variation were also analyzed by numerical taxonomic methods

(Sneath and Sokal, 1973), using the NT-PAK (Numerical Taxonomy

Package by W. W. Moss and L. N. Bell). The data were run on the

CDC-6400 computer |at the University of California Computer Center at

Berkeley. Because fully mature LZ. luctuosa are sexually dimorphic,

males and females were run separately.

CHARACTERS

Only mature or juvenile natural adult specimens were used in this

study. Presence of pruinosity, a characteristic whitish bloom present on

many male Libellulidae, was the basis for selecting mature males.

Pruinosity is often destroyed on preserved specimens by heat (Pinhey,

1951), chemical solvents, or leakage of body oils. Excessive heat can

also cause the wings |and body to glisten like tenerals immediately after

eclosion. The types of L. odiosa are in this condition and were so de-

scribed by Hagen (1861): “Entirely brassy-black ...,”? and Muttkowski

(1908) separated L. odiosa from L. luctuosa by this means. No females

with glistening wing membranes nor teneral specimens were used be-

cause their coloration and maculation differ from mature adults and

because many tenerals were in poor morphological condition, preventing

accurate measurements. Juvenile specimens possess a hard cuticle but

lack fully developed pruinosity patterns. The accessory genitalia of

males, usually of great taxonomic importance in dragonflies, showed no

consistent differences, and were not used. Post-mortem color changes

in a few poorly preserved specimens, especially females, were recorded,

even though these changes were probably unnatural. Missing characters

for about 6% of the specimens were substituted by values from similar

specimens of the same size. These conditions constitute some of the

inherent errors common to any phenetic study of organisms.

All continuous characters except hind wing length were measured

to the nearest 0.01 mm using an ocular micrometer. Repeated measure-

VoL. 52, No. 3, Jury 1976 187

TABLE 2. Factor Loadings for Characters of L. luctuosa. Important

loadings (—.200 > « > .200) are underlined.

Males Females

Principal Component Principal Component

Character I I TI I Il

Hindwing —.239 —.385 .099 —.376 180

Pterostigma 325, Ler —.004 376° 136

Ankle Cells ~ 047 -.317 27 —.158 150

Gaff —.134 2638 025 ~.269 101

Sole 136 431 089 —1Br .349

Middorsal 332 005. -.033 -.188 =.123

Stripe Width i, rs

Femur —.258 —.275 .093 —.360 129

Tibia =302 181. 031 318 097

Abdominal Segment —- oO

5 Length —.226 —.264 033 —.312 .203

Caudal Appendages e182 267° —.086 113 ~ (145

Wingtip Width ~.190 ~ (185° .001 — —

Wingtip Length .098 —.063 .013 —.156 —.358

Thoracic Color ~.362 234 -.103 — —

FW Fenestration 041 ~ 020 -.021 ~.034 —.056

HW Fenestration —.335 117 —.030 —.279 -.172

Frons 076 052 620 ~ 188 379

Clypeus 171 -.119 ~ 296 215 346,

Labium 221 -.181 ~ 290 017 229

Middorsal Stripe 263 101 ~ (297 141 346

Color a ar =,

Prothoracic Leg 048 123 478 .149 306

Color —- a

Percentage of

Total Variation 28% 14% 8% 27% 18%

ments revealed an error of about 5%. A standard millimeter rule was

used for the hind wing measurement. Of the 20 characters listed in

Table 2, only 18 were recorded for females. The blue thoracic coloration

and wingtip width characters were not present in females. Color ter-

minology is after Ridgway (1912) and morphological terminology is

after Needham and Westfall (1955).

188 THE Pan-PAciFIC ENTOMOLOGIST

ANALYTICAL TECHNIQUES

Multivariate techniques simultaneously compare all character values

over each OTU (Operational Taxonomic Unit). Many authors have

utilized these techniques for biological material, including Moss (1967),

Moulton (1973) and Rohlf (1968) for principal components, Barlow,

Graham and Adisoemarto (1969) and Bigelow and Reimer (1954) for

linear’ discriminant analysis, and Rohwer and Kilgore (1973) and

Rohwer (1972) using both methods. Numerical taxonomic methods are

elaborated in Sneath and Sokal (1973).

Principal component analysis was first used because a prior assign-

ment of specimens into arbitrary reference groups is not necessary.

Combinations of characters representing OTU’s are transformed to un-

correlated axes represented in an n-dimensional hyperspace. Principal

axes are then established within this hyperspace so that each OTU can

be expressed by its spatial relationship to n-number of components. The

first component accounts for the greatest percentage of variation, the

second accounts for the next greatest percentage of variation, and so on

until all variation over all OTU’s is expressed. Factor loadings for all

characters provide a basis for determining which characters are dis-

criminatory and which are uncorrelated. One hundred sixteen males

(20 characters) and 72 females (18 characters) were analyzed.

Step-wise discriminant and linear discriminant analysis require an

assignment of OTU’s into arbitrary groups of known entities. The first

involves a multiple discriminant analysis in a step-wise manner, whereby

variables having the highest F-values (discriminatory ratio value of

each value compared to the sum values of previously entered variables)

are repeatedly selected for discrimination between groups. Each suc-

ceeding step utilizes the preceding combination of variables, adding

another variable either with the next higher F-value, or with an F-value

which, when combined with the previous set of variables, yields the

greatest separation of groups. With each iteration, variables yielding

the highest F-values are determined and the overlap and misclassifica-

tion of OTU’s between groups are recorded. Since many museum

specimens are affected by post-mortem color and pruinosity pattern

changes, only continuous characters were utilized to determine their

value in classifying OTU’s. The data were run in two groups for each

sex, one with 10 characters (hindwing, pterostigma, ankle cells, gaff,

sole, middorsal thoracic stripe width, femur, tibia, abdominal segment 5,

and superior caudal appendages), the other including an eleventh char-

acter, hindwing fenestration, to determine its importance in further

separating the groups. Twenty-five specimens from each phenotypic

VoL. 52, No. 3, Juty 1976 189

group, if available, were compared. Since only 13 female Colorado

River females were available, 32 odiosa females were included so that

the total number of OTU’s would approximate that for the males.

Linear discriminant analysis for two groups defines an axis through

multidimensional clusters of OTU’s resulting in a maximum separation

of the clusters. Coefficients for each character are provided and these

multiplied by their respective character value and summed, resulting in

a single score or z-value for each OTU which can then be plotted on a

histogram. From the histogram, unassigned specimens can be plotted in

the same manner prescribed for reference specimens, and their position

relative to the two end groups determined. Linear discriminant analysis

provides an objective means by which unknown OTU’s can be classified ;

only specimens with z-values intermediate between the two reference

samples cannot with assurance be assigned to either group.

Calculation of z-values for each taxon is most advantageous when only

a few characters are used. The use of many characters complicates the

computational procedure and defeats the simplicity of the method.

Bicoded qualitative characters which are easily discernible need not be

used, since the observer can easily segregate his samples into like phena.

Therefore, only four continuous variables were used: sole length, hind-

wing fenestration, middorsal thoracic stripe width, and femur length.

The four characters repeatedly ranked with high F-values above the

other seven variables in the step-wise discriminant program.

The NT 11 package used in this study performs character standardiza-

tion by variance, ranking of similarity coefficients, cluster analysis

using unweighted pair group averages (UPGA), histogram, and mini-

mally connected network (Primnet: Prim, 1957). The phenograms

were based on average linkage and represent a one-dimensional cluster-

ing of OTU’s by similarity coefficients. The Primnets represent a one-

dimensional linkage of OTU’s by lines of similarity based on taxonomic

distance. The longer the line between OTU’s, the less similar the OTU’s.

The relative position of non-linked OTU’s to each other is arbitrary and

does not denote similarity.

RESULTS

Principal Component Analysis.—The spatial relationships of the three

phenotypes are depicted in Figures 5 and 6 for males and Figure 7 for

females. The squares, triangles, and circles throughout this study repre-

sent luctuosa, odiosa, and Colorado River morphs, respectively. The

first scattergram for each sex shows the OTU’s in relation to the first

and second components; the second set for males shows the OTU’s in

190 THE PAN-PaciFIC ENTOMOLOGIST

COMPONENT 2

COMPONENT £1

exe,

Fic. 5 (above) and Fic. 6 (right). Scattergram of principal component scores

for males. Open circles = Colorado River morphs, solid triangles = odiosa, and

solid squares = eastern luctuosa. Distribution of symbols indicates overall phenetic

similarity to other morphs. Continuous measured characters (Table 2) are pri-

marily responsible for separation of Colorado River and odiosa-luctuosa morphs

in Fig. 5. Color-coded characters (‘Table 2) are primarily responsible for separation

of morphs in Fig. 6.

relation to the first and third components. Clusters of OTU’s for males

are defined by combining both graphs.

Factor loadings for males (20 characters) are listed in Table 2. Only

wingtip color length and forewing fenestration resulted in uniformly low

values (—.100 < « <.100) for the first three components, indicating

that these characters were not important in segregating the groups. A

combination of continuous and coded characters was important in sepa-

rating the various groups. Measured characters primarily showed the

highest factor loadings (—.300 > « > .300) on the first two compo-

nents. Highest coefficients were for hindwing (component 2), ptero-

VoL. 52, No. 3, JuLy 1976 191

co ee

COMPONENT 3

a

COMPONENT 1

oo

Fic. 6. See Fig. 5 for explanation.

stigma (component 1), sole (component 2), middorsal stripe width

(component 1), tibia (component 1), thoracic color (component 1),

and hindwing fenestration (component 1). Character coefficients on the

third component revealed that color variation (frons, clypeus, labium,

middorsal stripe, and prothoracic leg color) was mostly responsible for

separating the groups; the ankle cell number was the only measured

character with high value. The first three components accounted for

28%, 14%, and 8%, respectively, or a total of 50% of the variation.

Components 4 through 18 accounted for the remaining 50%, but each

component averaged 3.3% (range = 6% to 1%).

192 THE Pan-PaciFic ENTOMOLOGIST

The scattergrams for males show luctuosa and odiosa to form a single

cluster, although individuals of odiosa approach Colorado River indi-

viduals in appearance. The third component (Fig. 6) was minimally

useful in further delineating these groups. Three males were misclassi-

fied: one odiosa clustered with the Colorado River forms (components |

and 2); another odiosa clustered correctly in relation to components

1 and 2, but was greatly separated by component 3; a single Colorado

River individual clustered with the latter odiosa specimen, forming a

separate subgroup. The last two males were juvenile, which probably

accounts for their aberrant location in the scattergram. Since the third

component represents primarily color variation, color and maculation of

these juveniles are probably responsible for their misclassification. Blue

pruinosity on the juvenile Colorado River morph had not obscured the

yellow middorsal stripe and brown and yellow sides of the synthorax. In

addition, both juveniles had an olive-brown frons and lighter facial

colors, instead of a black frons and darker facial colors common to all

other individuals. The important factor loadings (Table 2) for these

characters on component 3 probably contributed to their misclassifica-

tion. The thoracic color coefficient was also relatively high on com-

ponents | and 2, and it probably contributed to the odd placement of the

juvenile Colorado River male, since it was the only Colorado River

morph lacking the blue pruinosity on the sides of the pterothorax. Of

the 116 males, 2% were misclassified according to the author’s sub-

jective treatment of the groups. The odiosa and luctuosa phenotypes

are not phenotypically distinct enough to warrant their status as separate

taxa.

Results for the females were similar to those for the males, except that

the 13 Colorado River females did not separate as clearly. Factor load-

ings for the first two components (Table 2) show forewing fenestration.

as in the males, to be least important in separating the phena. All other

characters were important in explaining the variation; however, higher

coefficients (—.300 > « >.300) prevailed for continuous characters

(hindwing, pterostigma, femur, tibia, and abdomen 5) on component 1,

while the same range of values predominated for coded character coeffi-

cients (wingtip length, frons, clypeus, middorsal stripe color, and pro-

thoracic leg color) for component 2. Only one measured character

(sole) resulted in a high value on component 2. Because body color

patterns in female L. luctuosa vary more than in mature males, it is

logical to expect greater importance for coded color characters. The

southwestern desert forms are lighter than their eastern counterparts.

The other components were not useful in further delineating between

VoL. 52, No. 3, Juty 1976 193

COMPONENT 2

COMPONENT 1

ee

Fic. 7. Scattergram of principal component scores for females. See Fic. 5 and

6 for explanation of symbols.

<roups. The first and second components accounted for a total of 45%

of the variation. Components 3 through 17 accounted for the remaining

55%, with each component averaging 3.4% (range = 10% to 1%) of

the variation.

As in males, there is no clear separation between odiosa and luctuosa

morphs. Of the 13 desert OTU’s, one clustered with odiosa (Fig. 7).

This specimen was the only juvenile female of the group (lacking most

of the violaceous pruinosity on thorax and abdomen), but otherwise its

color and maculation did not differ appreciably from the other 12.

However, it and one other female were the smallest specimens of the

croup. The other female was mature, but its location in Figure 7

(PC, = —.367, PC. = —.288) places it near the juvenile. Phenetic gaps

between the desert and odiosa-luctuosa phenotypes are not as great as

for the males.

Step-wise Discriminant Analysis—Of the 10 or 11 measured charac-

ters used, only four, with two exceptions, repeatedly emerged with high

194, THE PAn-PAcIFIC ENTOMOLOGIST

TABLE 3. Classification of Assigned OTU’s into Respective Groups

with 10 and 11 Characters by Step-wise Discriminant Analysis—Males

(M) and Females (F).

Number Percent

Number of Colo. Misclas- Misclas-

Characters Morphs luctuosa odiosa Riy. Total sified sified

M 10 luctuosa 20 5 0 25 5 20

odiosa 7 16 2 25 9 36

Colo. Riv. a 0 24 25 1

M il luctuosa 24 1 0 25 1 4

odiosa 5 19 1 25 6 24

Colo. Riv. 0 1 24. 25 1 4,

F 10 luctuosa 20 4 1 25 5 20

odiosa 8 22 2 32 10 31

Colo. Riv. 1 1 11 13 2 15

F 11 luctuosa 23 2 0 25 2 8

odiosa 6 24 2 32 8 25

Colo. Riv. 0 1 12 13 1 7

F-values. Highest of the F-values was for hindwing fenestration (14.36

< F < 706.18), followed by middorsal stripe width (9.16 < F < 58.82),

then tibia (8.33 < F < 37.46), and finally sole length (0.90<F<

28.17). Abdominal segment length (F = 4.02) and femur length (F =

7.21) were the only other characters selected in the first three steps.

Classification of assigned specimens into their respective groups

showed odiosa morphs to be more frequently misclassified (up to 36%)

than the other two groups. This shows the odiosa sample to be a more

heterogeneous group, a conclusion supported by the high variability of

hindwing fenestration. Table 3 shows that most of the misclassified

odiosa were placed with luctuosa.

Linear Discriminant Analysis.—Principal component analysis re-

vealed insignificant differences between odiosa and luctuosa, so these

forms were treated as one group in this analysis. The four characters

selected by step-wise discriminant analysis were used to classify un-

assigned L. luctuosa. A histogram of z-values for males, using the fol-

lowing four discriminant function coefficients, is shown in Fig. 8:

sole = .036, hindwing fenestration = —.006, middorsal thoracic stripe

width = .116, and tibia length = —.025. The mean z-value for the odiosa-

luctuosa group was .208, and for the Colorado River males, .105. While

the mean of the two was .156, it could not be used to separate the groups,

because four odiosa had values less than .156; but a critical z-value of

VoL. 52, No. 3, JuLy 1976 195

> 10

re)

Zz n- 53

Lu

a)

‘) 5

uJ

o

Le

e)

+

15

nz98

~-

o)

z 10

int

a

i

= 5

Te

0 4

28 24 20 16

DISCRIMINANT SCORE

Fic. 8. Histograms of discriminant scores for reference males (below) and

unassigned males (above). White = luctuosa (reference) and odiosa (unas-

signed), black = Colorado River morph. Means for all groups are marked by

small arrows; larger medial arrows indicate critical z-value.

.130 segregated the two reference samples. Therefore, OTU’s with a

z > .130 are classified as odiosa-luctuosa phenotypes, while OTU’s with

z<.130 are classified as Colorado River morphs. Two misclassified

Colorado River morphs had scores of .255 and .164 (Fig. 8). The

former specimen was the juvenile previously misclassified by principal

component analysis. The second male was similar to the others: no

reason for its misclassification is apparent. As with principal com-

ponent analysis, 2% of the reference males were misclassified.

Fifty-three unassigned male L. luctuosa comprising mostly of odiosa

phenotypes from Texas, New Mexico, Arizona, California (Central

Valley), as well as nine Colorado River phenotypes from Yuma and

Phoenix, were plotted using discriminant constants for reference males

(Fig. 8). All but one of the specimens were classified correctly, though

the odiosa phenotypes are heavily skewed to the right of the reference

males. The odiosa sample lacked any eastern luctuosa phenotypes and

indicates that the odiosa morphs are intermediate between the end

groups but are more similar to nominate L. luctuosa than to the Colo-

rado River phenotype. Only one odiosa morph had a z-value falling

within the critical range and could not be classified with certainty.

196 THE PaAn-PactFIC ENTOMOLOGIST

Fic. 9. Minimally connected network (Primnet) for males (n = 116) based

on taxonomic distance. Open circles = Colorado River morphs, triangles = odiosa,

squares = luctuosa. Connecting lines between OTU’s indicate phenetic similarity,

placement of non-connected OTU’s is arbitrary and indicates no phenetic relation-

ship. Superimposed solid curvilinear lines indicate a priori clusterings of Colorado

River morphs, dotted lines indicate those misclassified. See text for further

explanation.

Linear discriminant values for females were not useful in separating

Colorado River forms from odiosa-luctuosa. Only 13 Colorado River

females were available for the study, and four of them were classified as

intermediate between odiosa-luctuosa phenotypes. Two others fall on

the critical z-value (between —.39 and —.41), and the remaining seven

form one end group. Of the 70 females, 8% were misclassified, but 46%

of the Colorado River females were misclassified, compared to 5% for

males.

Numerical Taxonomic Results—Primnets for males and females are

shown in Figures 9 and 10. The curvilinear lines superimposed over

the Colorado River morphs (open circles) indicate the author’s subjec-

tive classification of this group; dotted lines indicate misclassifications

in the Primnet. The results are largely concordant with principal com-

ponent and linear discriminant analysis. Little distinction was made

between odiosa and luctuosa males (Fig. 9) ; there were several connec-

tions between the two. The Colorado River morphs are on a separate

sidebranch, with two odiosa OTU’s connected to it. On the other hand,

two Colorado River morphs are connected to odiosa OTU’s in remote

VoL. 52, No. 3, Juty 1976 197

Fie. 10. Minimally connected network (Primnet) for females (n = 70) based

on taxonomic distance. Symbols and designations as for Figure 9. See text for

further explanation.

parts of the network. One of these is the same juvenile specimen mis-

classified by principal component and linear discriminant analysis. The

other specimen was not misclassified by any of the previous methods,

and the reason for its misclassification is unclear. The taxonomic dis-

tance between OTU’s is relatively uniform, with a range of 0.38 to

1.96, with 104 OTU’s having values less than 1.00. If odiosa and luctu-

osa morphs are considered the same, only two Colorado River morphs

out of 116 OTU’s (2%) were misclassified according to a priori treat-

ment by the author.

Classification of the females (Fig. 10) was similar to preceding re-

sults but was much more confused. The Colorado River phenotypes do

not form a separate sidebranch, but are linked to odiosa OTU’s in

different parts of the network. Two females of the Colorado River

morph are connected only to odiosa phenotypes, the same two specimens

misclassified by principal component and linear discriminant analysis.

Taxonomic distances range from 0.33 to 2.10, but 65 of the OTU’s had

distances less than 1.00. In both Primnets, no Colorado River pheno-

types are connected to any eastern luctuosa morphs.

198 THE PaAn-PAcIFIC ENTOMOLOGIST

As with the previous programs, phenetic gaps between Colorado River

morphs are obvious in males, but not in females. Most misclassified

OTU’s were identical in all methods, which indicates discrepancies in

size and coloration due to juvenile condition to be the most likely causes

of misclassification.

DISCUSSION

The author subjectively recognized three distinct morphs of Libellula

luctuosa, with phenotypic intermediates occurring in the central United

States. A definite phenetic gap was observed only between the highly

pruinose desert populations and the less pruinose luctuosa-odiosa pheno-

types. The lack of intermediate forms may be due in part to inadequate

collecting, but is mostly a result of the absence of specimens from the

Tehachapi Mountains of California and from the eastern edge of the

Sonoran Desert in Arizona.

Results of the various analyses show a high degree of concordance in

the classification of intraspecific forms. The a priori distinctions be-

tween odiosa and luctuosa are not salient and the names do not warrant

separate taxonomic rank. The only recognizable difference between the

two is the degree of clearing in the hindwing bands, but an adequate

sampling of populations of the odiosa morph shows this condition to be

highly variable. Within the same deme are found individuals with litile

or no clearing (Fig. 4a) and with hindwing bands approaching dark

rings (Fig. 4d). The pruinosity patterns of male odiosa-luctuosa morphs

are identical, confined largely to the mesepisternum and abdomen (Fig.

3c). The middorsal thoracic carina and mesopleural regions are

‘always black, and the rest of the synthorax is dark brown.

The California populations of odiosa, although geographically isolated

from their midwestern counterparts by the Great Basin, are not pheno-

typically distinct. It seems likely that the allopatric populations of

odiosa at one time shared a common gene pool instead of the two forms

having arisen independently of one another. Specimens from the

Pacific Northwest are rare, and the two individuals from British Colum-

bia are typical of eastern luctuosa in appearance. While this suggests

that L. luctuosa may range across southern Canada, no specimens are

known from Alberta or Saskatchewan.

No captures of L. luctwosa are known from Nevada (La Rivers, 1940,

1941), Utah (Brown, 1934; Larsen, 1952; Musser, 1961), Wyoming,

South Dakota, or Montana (Bick and Hornuff, 1972, 1974), where its

absence may be due to adverse environmental effects of temperature,

precipitation, or altitude. The Colorado River forms are allopatric with

VoL. 52, No. 3, Juty 1976 199

the odiosa morph, and no intermediate forms are known; but this may

be due to scanty collecting in the Tehachapi Mountains and the eastern

Sonoran Desert. Before the drying of the Southwest during the later

Tertiary (Axelrod, 1967; Antevs, 1955), forms similar to odiosa may

have inhabited the area now occupied by the desert forms. Ancestral

desert forms, then, may have at one time provided a continuous distri-

bution of odiosa morphs from California to the central United States.

Libellula luctuosa has a Nearctic distribution and does not penetrate

Neotropical regions of Mexico (Calvert, 1906) or peninsular Florida

(Byers, 1930). Its absence from the Neotropics and the Great Basin is

probably not due to physical barriers, since L. luctuosa is a vagile species

which regularly frequents temporary bodies of water in the Southwest.

The frequency for extensive brown wingtips in females increases from

east to west, so that all Colorado River females possess the condition

shown in Figures 3e and 3f. In contrast to these, 29 of 110 (26%)

odiosa females west of the Mississippi River and two of 30 (6.6%) L.

luctuosa east of the Mississippi showed wingtip coloration comparable

to the desert females. The sole and metathoracic tibia length of the

Colorado River morphs and eastern L. luctuosa show an inverse rela-

tionship in size. The hindwing width of the Colorado River forms is less

than that of the eastern forms, but the metathoracic tibiae are longer in

the desert forms. A t-test performed on the means of both characters for

both sexes was significant at the 0.05 level. The reason for this variation

is unknown, but similar patterns of geographic variation have been re-

ported by Alpatov (1929) for honeybees and by Rensch (1943) for

carabid beetles.

Thirteen of 188 OTU’s (7%) were misclassified by one or more

methods of analysis. Eight of those were females. Of the 13, only three

were misclassified by all methods—a juvenile male Colorado River

morph and two female Colorado River morphs. The male differed

significantly from the others in its group because pruinosity had not yet

obscured its thoracic pattern. The females were probably misclassified

due to age and size differences. One was juvenile, the only one of its

sroup in this condition, and both were relatively small—their hindwing

lengths were 37 mm, compared to a mean of 39.3 mm for the entire

group. One of the other males misclassified by principal component

analysis was also juvenile, but no other anomalies were observed for the

remaining nine specimens, and the reasons for their misclassification

remain obscure.

Pruinosity patterns are responsible for the greater distinctness of

male than female desert forms. The presence of pruinosity is reflected

200 THE Pan-PAcIFIC ENTOMOLOGIST

in three male characters: thoracic coloration, width of middorsal stripe,

and prothoracic leg color. The thoracic coloration in females is brown

and yellow with only a violaceous pruinose tint (not present in odiosa-

luctuosa phenotypes) covering the usually dark brown patterns of the

thorax and abdomen (Fig. 3b). The yellow middorsal stripe is always

present in all females, and the prothoracic leg color is yellow. The ex-

tensive pruinose condition of the males and, to a lesser extent, females, is

probably of some adaptive significance. Unfortunately, the chemical

nature and biological significance of pruinosity are unknown. Johnson

(1973) speculates that “it is a process of nitrogen elimination acted on

by sexual selection producing mate recognition clues.” There is strong

evidence that pruinosity does provide mate recognition cues (Jacobs,

1955; Johnson, 1962a, 1962b), but it is also present on old females of

some species (Longfield, 1960; personal observations), so its pro-

nounced condition on the southwestern desert forms may also indicate a

physiological function. The southwestern morphs conform, as do many

other invertebrates, to Gloger’s rule, which states that races from cool

areas are more heavily pigmented than those from warm areas. There

is no undisputed explanation for this phenomenon, but solar reflectance

by pale coloration may be one possibility (Bodenheimer, 1954).

ACKNOWLEDGMENTS

I wish to express my sincere thanks to John T. Doyen for his patience

and guidance in answering numerous questions and for his help in

completing this work. His aid with the various computer techniques,

and his reading and constructive criticism of the entire manuscript are

most gratefully acknowledged. Special thanks are also due Woodrow

Middlekauff, Jerry A. Powell, and Dennis R. Paulson for reading the

manuscript and offering constructive criticisms.

The following persons and institutions loaned specimens of Libellula

luctuosa so that this study could be more complete: E. Aitcheson,

Museum of Northern Arizona, Flagstaff; F. Byers, University of Kansas,

Lawrence; O. S. Flint, United States National Museum, Washington,

D.C.; L. Floge, American Museum of Natural History, New York;

S. Frommer, University of California, Riverside; L. K. Gloyd, Univer-

sity of Michigan, Ann Arbor; F. Hasbrouck, Arizona State University,

Tempe; C. L. Hogue and F. S. Truxal, Los Angeles County Museum;

H. B. Leech and P. H. Arnaud, California Academy of Sciences, San

Francisco; J. A. Powell and J. Chemsak, University of California,

Berkeley; S. S. Roback and W. W. Moss, Academy of Natural Sciences

of Philadelphia; F. G. Werner, University of Arizona, Tucson; M. J.

VoL. 52, No. 3, JuLy 1976 201

Westfall, Jr., University of Florida, Gainesville; and H. E. Evans of the

Museum of Comparative Zoology, Cambridge, for the loan of the types

of L. odiosa. I wish to express my sincere thanks and appreciation for

their cooperation. D. R. Paulson of the University of Washington and K.

Tennessen of the University of Florida kindly made available material

from their private collections.

The photographs were executed by Herman G. Real; and I especially

wish to thank my wife, Jo, who typed the manuscript and aided me with

its completion.

Computer time was provided by a grant from the University of Cali-

fornia Computer Center. This study was funded in part by a traineeship

from the National Institutes of Health.

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317-335.

Axetrop, D. I. 1967. Geologic history of the California insular flora, in Pro-

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Bartow, C. A., Granam, J. E. Anp ApisoemaARTO, S. 1969. A numerical factor

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BENNEFIELD, B. L. 1965. A taxonomic study of the subgenus Ladona (Odonata:

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Bick, G. H. anp Hornurr, L. E. 1972. Odonata collected in Wyoming, South

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Bick, G. H. anp Hornurr, L. E. 1974. New records of Odonata from Montana

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BiceLtow, R. S. anp Rermer, C. 1954. An application of the linear discriminant

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BopENHEIMER, F. S. 1954. Problems of physiology and ecology of desert animals,

in Biology of deserts. J. L. Cloudsley-Thompson, Institute of Biology,

London, pp. 162-167.

Brown, C. J. D. 1934. A preliminary list of Utah Odonata. Occas. Pap. Mus.

Zool. Univ. Mich., 291: 1-17.

Byers, F. C. 1930. A contribution to the knowledge of Florida Odonata. Univ.

Fla. Pubs. 1G) 1-327.

CALveRT, P. C. 1906. Biologia Centrali Americana: Neuroptera, R. H. Porter

and Dulau Co., London, pp. 145-212.

Hacen, H. A. 1861. Synopsis of the Neuroptera of North America. Smithsonian

Miscellaneous Collections, 4(1): 1-344.

Hacen, H. A. 1875. Synopsis of the Odonata of America. Proc. Boston Soc.

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202 THE PAn-PAciFIc ENTOMOLOGIST

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Jacosps, M. E. 1955. Studies on territorialism and sexual selection in dragonflies.

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Pachydiplax longipennis Burmeister (Odonata: Libellulidae), South-

western Nat., 7(3-4): 191-197.

Jounson, C. F. 1962b. Reproductive isolation in damselflies and dragonflies

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Argia apicalis (Zygoptera: Coenagrionidae). Can. Entomol., 104(10) :

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SCIENTIFIC NOTE

A Female Specimen of Acanthocinus (Canonura) leechi.—In 1956 (Ann.

Entomol. Soc. Amer., 49:228), Lawrence S. Dillon described a new species of

Canonura, C. leechi, from a single male specimen from Jerome, Yavapai County,

Arizona. The author recently collected a female in the Hualapai Mountains,

Mojave County, Arizona, at black light, in pinyon pine juniper forest, at approxi-

mately 5,300 feet. Since the female has been unknown until this time, a brief

description follows.

Form moderately robust, similar to the male. Pronotum at sides beneath

lateral tubercles deeply punctate, lacking fuscous maculae. Antennae twice as

long as body, segments with the following ratio: 1.0; .2; 1.4; 1.3; 1.2; 1.2;

1.1; 1.1; 1; 1; 1. Abdomen with fifth sternite prolonged, slightly shorter than

sternites 2 through 4 combined, ovipositor strongly produced, extending 5 mm

beyond tips of elytra. Length: 16 mm, not including ovipositor.

This insect is uniformly speckled with numerous small dark spots. These

spots are much more discernable due to the absence of well defined hoary

pubescence which is so pronounced in A. spectabilis (LeConte) and A. princeps

(Walker). A. leechi also differs from A. princeps by the absence of fulvous

markings on the elytra and by the color of the pubescent maculae of the pronotum.

—A. E. Lewis, 1360 Paseo Redondo, Burbank, CA 91501.

Notes on North American Nepidae

(Hemiptera: Heteroptera)

JoHn T. PoLHEMUS

3115 S. York, Englewood, Colorado 80110 *

- In connection with continuing studies aimed at an eventual compre-

hensive treatment of the zoogeography and systematics of North and

Middle American Aquatic Hemiptera, numerous collections and investi-

gations have been made resulting in data that are needed by other

workers. This paper is one of a series of short notes presenting selected

data on systematics and distributions.

In 1930, Hungerford considered it remarkable that a new Ranatra

would be found in the United States. His R. texana Hungerford is now

known to occur also in Middle America (see below) with the Texas

records being the northernmost extension of its range. With much

more comprehensive data available to us now, it is perhaps more

remarkable that there is an apparent endemic relict new species of

Ranatra in Arizona, which I describe below.

The description follows roughly the format of Lansbury (1972)

rather than Hungerford (1922), in order to preserve a continuity of

style with the latest major reviser in the group. Lansbury’s lora of the

head is equivalent to Hungerford’s jugum. All measurements given in

units have 60 units = 1 mm. Types held in the Polhemus Collection

are irrevocably committed to later placement in a designated type reposi-

tory. All material upon which this paper is based is in the Polhemus

Collection unless otherwise noted. I am indebted to A. S. Menke, U. S.

National Museum (USNM) and J. Donahue, Los Angeles County

Museum of Natural History (LACM) for their assistance.

Ranatra montezuma, n. sp.

Male.—body length 21-24 mm, respiratory siphon 13-16 mm; Female: body

length 25-28 mm, respiratory siphon 19-21 mm.

Vertex clearly raised above eyes, smoothly rounded. Eye width less than inter-

ocular space (4/4.5). Clypeus and lora about equally high. Elongations of second

and third antennal segments equal.

Anterior lobe and posterior lobe unicolorous. Anterior lobe almost twice as

long as posterior lobe (7/4), humeral width about one-fourth greater than anterior

width (13/10.5). Prothorax without strong median keel or sulcus. Scutellum

about twice as long as broad (8.3/4.5). The median finger-like posterior projection

* Contribution from the Department of Environmental, Population and Organismic Biology, Uni-

versity of Colorado, Boulder, 80302 and Martin Marietta Corp. Denver 80201.

Tue Pan-Pactric ENTOMOLOGIST 52: 204-208. JuLy 1976

VoL. 52, No. 3, JuLty 1976 205

1/2 ym ”

V4 mm

fmm

d

Fic. 1. Ranaira montezuma n. sp., male. a. metasternum. b. left paramere. c.

antenna. d. foreleg.

of the metasternum not elongated, reaching only to middle of hind coxae, slightly

raised medially (Fig. 1).

Fore femur with stout median tooth and small apical tooth. Posterior femur

slightly longer than intermediate femur, reaching only to basal fifth of abdominal

sternite 5. Tips of posterior tarsi not exceeding middle of siphon in male, basal

third of siphon in female.

Female operculum barely exceeding connexivum. Male paramere with a dis- .

tinct tooth on inner margin.

Material examined: Holotype (¢), Allotype (@), and paratypes 214 6,

11@ 9, Artzona, YAvApar Co., Nr. Rimrock, MONTEZUMA WELL, CL695, III-27-’75,

J. T. Polhemus (JTP Coll.). Paratypes, 42¢ ¢, 339 9, same location, June 19,

1968, A. S. Menke and O. Flint (USNM). Other specimens, not paratypes, 8¢ ¢,

42 9, same location, A. S. Menke and L. Stange, April 1, 1958 (LACM).

Discussion: Ranatra montezuma belongs to the same group of species

as brevicollis Montandon, fusca Palisot Beauvoir and quadridentata

Stal, which occur in the same region, but is smaller than any of them.

In this group, the last abdominal segment in the male does not extend

ventrally to embrace the distal part of the operculum, and the respira-

tory siphon of the females is not as long as the body. Of these species,

montezuma most closely resembles brevicollis in the short appendages

and general facies, but differs from the latter in having a relatively

206 THE PAn-PAcIFIcC ENTOMOLOGIST

longer anterior lobe, second antennal segment with digitate process

subequal to third (as opposed to half as long in brevicollis), and hav-

ing a much stouter male paramere of different shape. R. fusca has a

longer siphon, the lora are less elevated, and the eyes are larger and

more transverse than montezuma. R. quadridentata has a differently

shaped paramere (see Hungerford 1922) and the tylus is not as promi-

nent, appearing much shorter than in montezuma.

RANATRA TEXANA HUNGERFORD

Ranatra texana Hungerford, 1930. Can. Entomol. 62:217 (type, ¢, Bee Co, Texas,

in Snow Entomological Museum, Lawrence, Kansas)

Material examined: UNITED STATES. Texas: 364, 49-9, Christoval,

Conchos River, CL1113, V-9-64, J. T. Polhemus; 3¢ 6, 19, Ottine, CL396,

VIII-6-’67, J. T. Polhemus. MEXICO. Chiapas: 19, Puente La Flor, CL1247,

XIT-19-’69, J. T. Polhemus. Oaxaca: 18, Juchitan, CL1244, XII-18-’69, J. T.

Polhemus. San Luis Potosi: 1¢, 35 km. N. Tamazunchale, CL1241, XII-16-’69,

J. T. Polhemus. Veracruz: 16, W. of Veracruz, CL1335, I-16-’70, J. T. Polhemus.

GUATEMALA. 32 4, 49 2, 10 mi. N. Ascuncion Mita, CL1313, I-10-’70, J. T.

Polhemus; 2¢ 4, 192, Puerto San Jose, CL1250, XII-20-’69, J. T. Polhemus.

RANATRA SPATULATA KUITERT

Ranatra spatulata Kuitert, 1949. J. Kansas Entomol. Soc. 22:32 (Type, 4, listed

as being from “Key West, Florida, May 8, 1919,” in U. S. National Museum.)

Kuitert (1949) gives the type locality as Key West as noted above,

and further notes that the specimen is from the Frank Lutz Collection

and is in the American Museum of Natural History. In fact, the speci-

men is from the J. C. Lutz Collection of Philadelphia, now in the

USNM, where I studied the type.

Over the years I have become increasingly suspicious of the validity of

the locality label on this type, the label being photographically repro-

duced from a typed original, probably by Lutz. In 1947 the assiduous

collector Raymond Beamer made a trip to the Keys to try for Ranatra

(Hungerford 1958), but was unsuccessful. Later I went to the Keys

for the same purpose, and determined that the only permanent fresh-

water habitat there is a pond on Big Pine Key which I worked without

finding any Ranatra. Finally, I studied the type in the USNM, and

found that it apparently is not allied to the American fauna, having the

metasternum and male parameres of a form typical of Old World species,

but not seen in any New World species.

Having just collaborated with Dr. Ivor Lansbury of Oxford in work-

ing out the differences in the Old and New World Ranatra, I asked

him to borrow the type and give me an opinion on this species. (See

VoL. 52, No. 3, Juty 1976 207

d

Fic. 2. Ranatra spatulata Kuitert. a. genital capsule. b. paramere, slightly

enlarged. c. paramere, distal portion. d. foreleg.

Lansbury 1974 for a discussion of Old and New World Ranatra.) Dr.

Lansbury very kindly obliged, and graciously offered the use of his

sketch figures (Fig. 2) and comments, which are given here in part:

“T have now examined the type of Ranatra spatulata Kuitert and find that it is

an African species which has been erroneously labelled, possibly by Frank Lutz.

As far as I can discover . . . spatulata is a species of the grandocula Bergroth group.

Comparison of the parameres shows a remarkable similarity with R. capensis con-

goensis Poisson, but the fore leg of the latter is quite different from that of spatulata

and is quite similar to grandocula.s.1. The metasternal plaque is very similar to

congoensis. As a matter of interest, the type of spatulata does not have its front

legs attached to the body, they are stuck on a piece of card (Hungerford det label)

so the possibility exists that they are not the right legs at all!! When Kuitert

described the species he partially removed the male genital capsule to see the

parameres, in doing so, he somehow damaged the internal genitalia, all that is

left is the bridge and lateral arms, the lamina ventralis and associated structures

have all been removed, consequently I was not able to make as full a study as I

would have normally done in these circumstances.

I am sorry not to be more specific about the real identity of spatwlata. I can how-

ever assure you that it is not a North American species or an Oriental one.”

As it seems certain that Ranatra spatulata is not of American origin,

it should be stricken from the U. S. list and carried instead as an Afri-

can species, with its exact origin unfortunately not known.

CURICTA PRONOTATA KUITERT

Curicta pronotata Kuitert, 1949. J. Kansas Entomol. Soc. 22:66 (type, ¢, Hua-

chuca Mts., Ariz., U. S. National Museum)

Vincent Roth rediscovered this species in northwestern Sonora and

collected it in numbers in Canyon de Evans. Searches in other locali-

208 THE PAn-PAcIFIC ENTOMOLOGIST

ties in upper Sonora and Arizona have not revealed other colonies.

Previously known only from the type series of three males, it is new

to Mexico. A good series reportedly collected in Canyon de Evans

(Roth, in litt.) has not been seen by me, but I have some that he and

I collected.

The female is of the same general facies as the male, but slightly

larger; body length 25 mm, respiratory siphon 10 mm.

Material examined: MEXICO. Sonora: 2¢ ¢, 19,6 nymphs, Canyon de Evans,

Sierra de los Ajos, Pine-oak, N. end, 31°N-110°W, VI-1-’71, V. Roth; 14,

Canyon de Evans, 17 mi. S. E. Cananea, CL570, III-30-’73, J. T. Polhemus.

CURICTA HOWARDI MONTANDON

Curicta howardi Montandon, 1910. Bull. Soc. Sci. Bucharest 18:181 (type, Vic-

toria, Texas in U. S. National Museum).

Curicta drakei Hungerford, 1922. Kansas Univ. Sci. Bull., 14 (18) :432.

This little species is widespread in southeastern Texas, and has been

reported from Louisiana by Hungerford (1922) and Ellis (1952). I

have found it in abundance only in Skull Creek (see below), but dili-

gent collecting resulted in specimens from several other localities.

Material examined: Texas: 55 adults and nymphs, Rock Island Co., Nr. Altair,

Skull Creek, CL392, VIII-6-'67, J. T. Polhemus; 1¢, 2 nymphs, Sheridan, CL394,

VIII-6-’67, J. T. Polhemus; 1¢, 19, Ganado, CL451, VI-5-’69, J. T. Polhemus;

19, College Station, VI-1-’31, H. Mills; 1¢, Brazos Co., College Station, X-10-196?,

J. Sweet.

LITERATURE CITED

Exuis, L. L., 1952. The aquatic Hemiptera of Southeastern Louisiana (exclu-

sive of Corixidae) , Amer. Midl. Nat., 48:302-329.

Huncerrorp, H. B., 1922. The Nepidae of North America, Kansas Univ. Sci.

Bull., 14(18) :425-469.

Huncerrorp, H. B., 1930. Two new water bugs from the Western U.S.A. (Nepi-

dae and Notonectidae) , Can. Entomol., 62: 216-218.

Huncerrorp, H. B., 1958. Raymond Hill Beamer, J. Kans. Entomol. Soc., 31(2):

57-66.

Kurrert, L. C., 1949. Some new Ranatra from the Americas, J. Kans. Entomol.

Soc., 22(1) :24-34.

Lanspury, I., 1972. A review of the Oriental species of Ranatra Fabricius (Hem-

iptera-Heteroptera: Nepidae), Trans. R. Entomol. Soc. Lond., 124(3) :

287-341, 262 figs.

Lansgury, I., 1974. Notes on Ranatra (Amphischizops) compressicollis Montan-

don with a review of its systematic position within the American Ranatra

(Hemiptera-Heteroptera, Nepidae), Zoologica Scripta, 3:23-30.

A new species of Hermatobates

(Hemiptera: Heteroptera)

LANNA CHENG

Scripps Institution of Oceanography

University of California, La Jolla, California 92093

_Hermatobates is a genus of rarely collected and little known marine

insects. The first species, H. haddoni Carpenter was described in 1892.

Several additional species were described subsequently (Coutiére and

Martin, 190la, 1901b; China, 1956, 1957; Herring, 1965) and they

were all assigned to the Gerridae although Matsuda (1960), in his

review of the World Gerridae, excluded Hermatobates from that family.

The status of this genus is thus still debatable; it will be discussed else-

where (Andersen, Cheng & Polhemus, in preparation). A review of the

relevant literature, with a list of the known species, their distributions

and their biology, was published by Cheng (1976).

The specimens to be described in the present paper were collected off

the west coast of Singapore; they were referred to briefly, without spe-

cific epithet, by Cheng (1966, 1976).

Hermatobates singaporensis new species

Holotype male— (Figs. 2, 5 and 6). Apterous, dark brown dorsally, paler brown

ventrally, without distinct markings. Body covered with short, velvety hairs. Legs

with front femora greatly swollen, bearing thirteen small teeth between basal and

distal tubercles; distal tubercle interlocks with tubercles borne at proximal end

of front tibia which bears two oblique hair combs at distal end, with a broad

groove between them (Fig. 3). Length, 3.5 mm; width, 2.0 mm; ratios of various

antennal and leg lengths as follows (10 units = 1 mm): Antennae, 9: 9.5: 6: 6;

front leg, 14: 14: 1: 1.5: 3; middle leg, 23: 12: 1: 6.5: 5; hind leg, 22.5:

13: 1: 6: 5.5 (there are 3 tarsal segments on each leg).

Allotype and paratype female—Color pattern as in male; meso- and meta-nota

differ from male in structure, with lobe-like extensions almost reaching apex of

abdomen (Fig. 1). Front femora not swollen, bearing a row of small teeth on ven-

tral edge but without pronounced tubercles (Fig. 4). Length, 3.6 mm; width

1.8-1.9 mm; ratios of various antennal and leg length as follows: Antenna, 7: 7: 5:

6; front leg, 11: 11: 1: 2: 3; middle leg, 18: 10: 1: 6: 5; hind leg, 18:

10: 1: 55+ 6.5.

Type locality—Holotype male, allotype and one female paratype from approxi-

mately 01° 17’ N and 103° 52’ E, about 5 km off the Siglap coast of Singapore

near an offshore “Kelong” (fishing platform). The type specimens, one male and

two females, known only from the type locality, were collected by dip netting from

a small boat on 3 October, 1963. All three type specimens deposited at the U.S.

National Museum, Washington, D.C.

THe PAn-PactFic ENTOMOLOGIST 52: 209-212. Juty 1976

210 THE PAN-PaciFIC ENTOMOLOGIST

Fics. 1-2. Hermatobates singaporensis. Fig. 1. Female. Fig. 2. Male.

Diagnosis.—This species can be distinguished from the 8 known species of Herm-

atobates by the arrangements of the teeth and tubercles on the male front femur

and tibia. In this respect it appears to be closer to H. weddi China (1957),

described from Monte Bello Island, and H. marchei (Coutiére et Martin) (1901b) .

from the Philippines. However, the front femur of the former species has 14 small

teeth between the basal and apical spurs and the latter has 12. There are 13 in

H. singaporensis. The number and arrangement of tubercles on the front tibia are

quite different in the three spcies.

I thank Dr. Jon L. Herring, U.S. Department of Agriculture, U.S. National

Museum, Washington, D.C. for confirming this is a new species.

1mm 4.

SS

Fies. 3-4. Hermatobates singaporensis. Fig. 3. Foreleg, male. Fig. 4. Foreleg,

female.

VoL. 52, No. 3, JuLty 1976 211

sw

veel ie

psig ets

eee

egies

oes

seid PTS

ei

Fics. 5-6. Hermatobates singaporensis, holotype male (scale in mm). Fig. 5. Dor-

sal aspect. Fig. 6. Ventral aspect.

LITERATURE CITED

Carpenter, G. H. 1892. Reports on the zoological collection made in the Tor-

res Straits by Professor A. C. Haddon, 1888-1889: Rhynchota from

Murray Island and Mabuiag. Proc. Roy. Dublin Soc. (new ser.),

7: 137-146.

Cuenec, L. 1966. A curious marine insect, Hermatobates. Malay. Nat. J., 19(5):

283-285.

Cuenc, L. 1976. The elusive sea-bug Hermatobates (Heteroptera). Pacific

Insects, 17: in press.

Cuina, W. E. 1956. A new species of the genus Hermatobates from the Hawaiian

Islands (Hemiptera-Heteroptera, Gerridae, Halobatinae). Ann. Mag.

Nat. Hist. ser. 12, 9(101) : 353-357.

Cuina, W. E. 1957. The marine Hemiptera of the Monte Bello Islands, with

descriptions of some allied species. J. Linn. Soc. Lond., Zool., 40(291) :

342-357.

CoutigrReE, H. and Martin, J. 190la. Sur un nouvel Hémiptére Halophile. Bull.

Mus. Nat. Hist. Paris, 4(ser. 1) : 172-177.

Coutirre, H. and Martin, J. 1901b. Sur un nouvel Hémiptére Halophile, Herm-

atobatodes marchei, n. gen., n. sp. Bull. Mus. Nat. Hist. Paris, 5:

214-226.

212 Tue PAn-PaAciFiIc ENTOMOLOGIST

Herrinc, J. L. 1965. Hermatobates, a new generic record for the Atlantic

Ocean, with descriptions of new species (Hemiptera: Gerridae). Proc.

U.S. Nat. Mus., 117:123-130.

Matsupa, R. 1960. Morphology, evolution and a classification of the Gerridae

(Hemiptera-Heteroptera). Univ. Kansas Sci. Bull., 41(2): 25-632.

SCIENTIFIC NOTE

Confirmation of the beach habitation of Apatolestes actites Philip and

Steffan (Diptera: Tabanidae) on the California coast.—The original descrip-

tion of Apatolestes actites Philip and Steffan (tribe Pangoniini) was based on

unique specimens of each sex that apparently were taken on or near two Cali-

fornia beaches (1962, Pan-Pac. Entomol., 38: 41-43). The type locality of A.

actites is Goleta Beach, near Santa Barbara, Santa Barbara Co., Calif. A third

specimen that is present in the insect collection of the California Department of

Health, Vector Control Section, Berkeley, was collected along the California

coast at Montana de Oro State Park, San Luis Obispo Co. Its somewhat teneral

condition suggests that it had emerged near the collection site.

One of us (VFL) collected a female A. actites on a sandy beach near Bolinas

Point, Marin Co., on 16 June 1974. Further collections made there in 1974 yielded

one male, one female, and about seven tabanid pupal skins; the latter were found

among driftwood in the supralittoral zone. On 4 July 1975, a cool, overcast day,

30 flies (8¢, 2292) were collected on this beach by five persons. Flies were preva-

lent below the high tide mark where clumps of seaweed were present. Numerous

tabanid pupal skins and desiccated pupae (tribe Pangoniini) were found in, on,

or protruding from sand above the high tide mark. Characteristics of the imago

of A. actites (e.g., pilose body, bare eyes) were visible through the cuticle of the

dead pupae. In the same area larvae of a species of Pangoniini were collected from

sand approximately 8 to 10 cm below the surface. Dr. W. W. Middlekauff, Mr.

T. Sukekane, and Mrs. S. Lane assisted in the July 4th collections.

Immatures also believed to be those of A. actites were collected at Manchester

State Beach, Mendocino Co., California, on 14 and 27 July 1975. This record, if

confirmed, would extend the northernmost distribution of A. actites approximately

140 km beyond Bolinas Point.

The foregoing observations support our belief that A. actites breeds on beaches

along the California coast. We are attempting to rear larvae obtained from both

the Marin and Mendocino Co. sites to conclusively associate them with pupae

and adults——V. F. Ler, California Academy of Sciences, Golden Gate Park, San

Francisco, Calif. 94118; R. S. Lang, California Department of Health, Vector Con-

trol Section, Berkeley, Calif. 94704; and C. B. Putiip, California Academy of

Sciences, San Francisco, Calif. 94118.

A New Species of Ischalia from Southeastern China

(Coleoptera: Pyrochroidae)

DANIEL K. YouNG

Department of Entomology, Michigan State University

East Lansing, Michigan 48824

Ischalia (Ischalia) chinensis new species

(Figures 1, 3, 4)

Description.—Length 4.9-5.8 mm. Body, exclusive of abdominal tergites, mod-

erately covered with yellowish setae, those of elytra more erect than those associ-

ated with remainder of body.

Male—Head sooty yellowish-brown unevenly suffused with dark brown to pice-

ous markings; frontal aspect with shallow longitudinal groove mesally, between

eyes. Antennae dark brownish to piceous basally, gradually becoming lighter

toward apices. Terminal segment of maxillary palpus cup- or scoop-like, with

outer surface convexly rounded and inner aspect concavely depressed.

Pronotum, scutellum, and ventral thorax sooty yellowish-brown unevenly suf-

fused with brownish to piceous coloration. Pronotum irregularly concave in lat-

eral view; gibbose anteriorly, sloping downward posteriorly for two-thirds of its

length, then gradually upward toward hind margin. Anterior margin of pronotum

(Fig. 1) weakly sinuate mesally, lateral margins evenly rounded from front poste-

riorly to the acute, weakly produced hind angles. Pronotal disk with prominent

median carina which is produced posterad of hind margin, and 2 circular im-

pressions or pits on either side of the carina: one near the carina and another

near the postero-lateral margin. Mesepisterna widely separated. Legs sooty yel-

lowish-brown to nearly piceous. Elytra elongate and covering abdomen, somewhat

broader posteriorly with apices slightly separated along suture, surface densely

and coarsely punctate to nearly reticulate; entirely pale yellowish-brown to yel-

lowish-brown with pale brownish sutural margins. Sutural, humeral, lateral discal,

and lateral carinae present with humeral carinae about one-third the length of

lateral discal carinae, the later gradually curving inwardly toward the sutural

carinae but terminating distant from them. Hind wing (Fig. 3) fully developed,

pale yellowish in color.

Abdomen sooty yellowish-brown ventrally, unevenly suffused with brownish to

piceous patches. Parameres ventrad of aedeagus, fused along entire length and

broadly emarginate apically with 6 setae near apex, 3 on either side (Fig. 4).

Female——Differs from the male by having the terminal antennal segment exca-

vated on outer surface, with the excavation densely setose.

TYPE INFORMATION: HOLOTYPE: (¢), Hone San, SE; Kiancsi, CHINA;

15 July 1936; L. Gressitt Collector; L. Gressitt Collection (CASC).

ALLOTYPE: (@), same data as holotype except date: 16 July 1936 (CASC).

PARATYPE: (1 9), Moxansan, CutnA; CheKiang Pr.; 19 September 1927;

Mrs. Dora E. Wright Collector (CASC).

DISTRIBUTION: As detailed above, southeastern China: KiangSi and Che-

Kiang provinces.

Tue Pan-Paciric ENTOMOLOGIST 52: 213-215. JuLy 1976

214, THe PAn-PAcIFIC ENTOMOLOGIST

1.0 mm

0.05 mm

Fics. 1-4. Structures of Ischalia. Fig. 1. I. chinensis Young, pronotal outline,

dorsal aspect. Fig. 2. I. suturalis Blair, pronotal outline, dorsal aspect. Figs. 3-4. I.

chinensis. Fig. 3. Right hind wing of male. Fig. 4. Distal portion of fused para-

meres, dorsal aspect.

REMARKS: The presence of widely separated mesepisterna places J/.

chinensis in the subgenus /schalia as redefined by Paulus (1971).

This species comes closest to the description provided by Blair (1912)

for his suturalis of Assam, India. Through the kindness of C. M. F.

VoL. 52, No. 3, JuLy 1976 215

von Hayek and the British Museum of Natural History, two syntypes

of suturalis were made available to the author for comparison.

The most striking difference between the species is the shape of the

pronotum, with that of chinensis (Fig. 1) slightly more transverse than

that of suturalis (Fig. 2), and the hind pronotal angles of the former

but weakly produced (strongly so in sutwralis). The lateral pronotal

margins of chinensis are evenly rounded from front to back and the

anterior margin gradually and weakly impressed mesally, while in

suturalis the lateral margins are slightly produced midway along their

length and the anterior margin is more abruptly and deeply sinuate

mesally.

Another major difference is the shape of the terminal segment of the

maxillary palpus, with that of chinensis convexly rounded on its outer

aspect and the inner surface deeply excavated to form a cup- or scoop-

like structure (in suturalis both surfaces are convexly rounded, thus

forming a bulbous structure).

The presence of an excavation in the terminal antennal segment of

the female makes chinensis unique among the Ischaliinae known to the

author, since secondary sexual dimorphism has not previously been

recorded for the group. Young’s (1975) description of the genus must

be expanded to include this feature.

I should like to acknowledge D. H. Kavanaugh of the California

Academy of Sciences (CASC) and H. B. Leech, formerly of the Cali-

fornia Academy of Sciences, for the loan of undetermined Asian Pyro-

chroidae which produced the specimens of chinensis. Also, my thanks

are extended to C. M. F. von Hayek of the British Museum of Natural

History for the loan of two syntypes of /schalia suturalis Blair, and to

S. G. Wellso of Michigan State University for reviewing the manuscript.

LITERATURE CITED

Biair, K. G. 1912. Descriptions of new species of Pyrochroidae. Ann. & Mag.

Nat. Hist. (8) 9: 527-533.

Pautus, H. F. 1971. Neue Pyrochroidae aus Nepal (Coleoptera, Heteromera) ,

mit einer Diskussion der verwandtschaftlichen Verhaltnisse der Familie.

Ztschr. Arbeits. Osterr. Entomol. 23: 75-85.

Younc, D. K. 1975. A revision of the family Pyrochroidae (Coleoptera: Hetero-

mera) for North America based on the larvae, pupae, and adults.

Contrib. Amer. Entomol. Instit. 11(3): 1-39.

The Genus Triacetelus Bates

(Coleoptera: Cerambycidae)

Joun A. CHEmsAK and E. G. LINSLEY

University of California, Berkeley

The purpuricenine genus 7J'riacetelus was proposed by Bates (1892)

to accommodate a single new species, sericatus, from Iguala, Guerrero,

Mexico. Zajciw in 1964 described a new genus, Paramphionthe, for a

species from El Salvador. This species was considered a synonym of 7’.

sericatus by Monné and Martins in 1972. In addition to the type species,

sericatus, we now consider the genus Triacetelus to consist of one new

species and another previously described one, both from Mexico.

This study was supported by the National Science Foundation

(GB-BM574.) for a monograph of North American Cerambycidae. We

gratefully acknowledge the following institutions and individuals for

the loan of specimens from collections in their care: California Acad-

emy of Sciences, San Francisco; Canadian National Collection, Ottawa;

Cornell University, Ithaca; Essig Museum of Entomology, University of

California, Berkeley; University of Kansas, Lawrence; Los Angeles

Museum of Natural History; Ohio State University, Columbus; and

G. H. Nelson.

GENUS TRIACETELUS BATES

Triacetelus Bates, 1892, Trans. Entomol. Soc. London, 1892:176; Monné and

Martins, 1972, Rev. Brasileria Entomol., 16:75.

Paramphionthe Zajciw, 1964, Ann. Acad. Brasileria Cienc., 36:437; Monné and

Martins, 1972, Rev. Brasileria Entomol., 16:75 (synonymy). Type: P. moestula

Zajciw, monobasic).

Form slender, sides subparallel. Head with front subvertical, short, deeply

impressed transversely; palpi subequal, short, apical segments cylindrical; mandi-

bles short, barely arcuate, broadly, obliquely truncate at apices; antennal tuber-

cles slightly elevated; antennae slender, about twice length of body in males

usually a little longer than body in females, segments slightly expanded at apices,

third much longer than first, fourth shorter than or subequal to third, fifth longer

than or equal to fourth, eleventh of males very long. Pronotum with acute lateral

tubercles at middle; disk convex, middle with a vague linear callus, base with a

shallow swelling each side of middle; prosternum not impressed, intercoxal

process abruptly declivous, apex narrow, coxal cavities wide open behind; meso-

sternal process slightly tuberculate at middle, gradually declivous in front;

episternum of metathorax broad, slightly tapering posteriorly. Elytra strongly to

vaguely bicostate on each side, surface sericeous to metallic; apices strongly to

shallowly emarginate to bi-emarginate. Legs slender; hind femora extending

Tue Pan-Paciric ENTOMOLOGIsT 52: 216-219. JuLy 1976

VoL. 52, No. 3, Juty 1976 217

beyond apices of elytra, dentate at apices; hind tarsi slender, first segment longer

than two following together, third segment cleft to about middle.

Type species: Triacetelus sericatus Bates (monobasic).

This genus is closely related to Psewdodeltaspis Linsley but the two

may be separated by the strongly five-callused pronotal disk, lack of

dense pubescence on the elytra, and by the strongly metallic color of

the elytra of Pseudodeltaspis.

Three species of Triacetelus are presently known, all from Mexico

with one species extending to El Salvador.

KEY TO THE SPECIES OF TRIACETELUS

i, Elytra densely clothed with long appressed pubescence which lies trans-

versely on disk, apices bi-emarginate

Elytra densely clothed with very short, appressed ai Aan lying longi-

tudinally, apices deeply emarginate, sutures strongly produced, color metal-

lic greenish to violaceous. Length, 19-21 mm. Michoacan ___. viridipennis

2(1). Pronotum reddish, irregularly punctate, basal lateral callosities not

punctate, disk with a prominent, glabrous, median callus. Length, 12 mm.

a CNP cet: > eee we eee ene. WEA WO ls ee eee Oe gee el oe emarginata

Pronotum black, very densely, confluently punctate throughout, disk with

a vague linear callus extending over most of length. Length, 11-18 mm.

pinaloa, Mexico £6. el Salvador Se reteset needa sericatus

Triacetelus viridipennis, new species

Male.—Form moderate sized, moderately tapering posteriorly; integument black,

elytra dull metallic greenish to violaceous; pubescence short, whitish. Head small,

front deeply impressed transversely, shallowly, irregularly punctate, densely clothed

with pale erect pubescence; median line deep, V-shaped; antennal tubercles

shallow, apices obtusely angulate; antennae slender, extending about 4 seg-

ments beyond elytra, segments three through ten slightly enlarged at apices, seg-

ments finely, densely clothed with very short, reddish-purple, recumbent pu-

bescence, segment three equal to four, fifth slightly longer than fourth, remaining

segments very gradually increasing in length to tenth, eleventh twice as long as

tenth. Pronotum broader than long, sides rounded with a small acute tubercle on

each side; disk convex with a vague, linear, median callus and shallow calluses on

each side near base; base broadly impressed; punctures fine, shallow, very dense,

contiguous, median callus sparsely punctate; pubescence fine, pale, depressed,

dense; prosternum densely pubescent, intercoxal process shallowly carinate at

middle, abruptly declivous, narrow behind, coxal cavities wide open behind;

meso- and metasternum very finely, densely punctate, densely clothed with long,

whitish, appressed pubescence, mesosternal process lying above level of coxae,

shallowly tuberculate at middle, gradually declivous in front. Elytra less than 2%

times as long as broad; each elytron vaguely bicostate; punctures very fine, con-

fluent; pubescence dense, very short, depressed; apices strongly obliquely emar-

ginate, lateral angles dentate, sutural angles elongate, obtuse. Scutellum densely

white pubescent. Legs with front and middle femora moderately clavate, hind

218 THE Pan-PAciFic ENTOMOLOGIST

femora linear, extending to apices of elytra; hind tibiae with a row of dense sub-

erect hairs on inner margin. Abdomen minutely, densely punctate and pubescent

except for glabrous spot at margin of first segment and glabrous margins of seg-

ments 2 to 4; last segment rounded at apex, emarginate at middle. Length, 21 mm.

Female.—Form similar, more parallel-sided. Antennae slightly longer than

body, outer segments moderately serrate, third segment longer than fourth, pu-

bescence dark. Abdomen with sternites narrowly glabrous at apices; last sternite

broadly truncate at apex. Length, 19 mm.

Holotype male, allotype (California Academy of Sciences) from APATZINGAN,

Micuoacan, Mexico, 5 August 1940 (Hoogstraal Expd. 40).

The short pubescence and dull metallic greenish to violaceous overall color of

the elytra will separate this species from other Triacetelus.

TRIACETELUS EMARGINATA (CHEVROLAT), NEW COMBINATION

Cyphosterna emarginata Chevrolat, 1862, Ann. Soc. Entomol. France, (4) 2:758.

Gambria emarginata, Lacordaire, 1869, Genera des coléoptéres, 9:189; Bates, 1880,

Biologia Centrali-Americana, Coleoptera, 5:86.

This species is known only from the unique female type at the British

Museum (Natural History). It greatly resembles sericatus by the

transversely lying pubescence and bi-emarginate apices of the elytra.

The pronotum and appendages are reddish with the elytra dark and

silvery pubescent. Additionally the disk of the pronotum is irregularly

punctate with the basal callosities glabrous. Additional material may

indicate that emarginata represents a population of sericatus. Length,

12 mm.

Type locality.— Yucatan.

TRIACETELUS SERICATUS BATES

Triacetelus sericatus Bates, 1892, Trans. Entomol. Soc. London, 1892:176, pl. 7,

fig. 12; Linsley, 1935, Trans. American Entomol. Soc., 61:100; Chemsak, 1967,

Jour. Kansas Entomol. Soc., 40:80 (Lectotype designation) ; Monné and Mar-

tins, 1972, Rev. Brasileira Entomol., 16:75.

Paramphionthe moestula Zajciw, 1964, Ann. Acad. Brasileria Cienc., 36:438, fig. 4.

Male.—Integument black, elytra and legs frequently pale brownish. Head with

front clothed with fine erect pubescence; antennae about twice as long as body,

third segment subequal to fourth, fifth longer than fourth, segments finely pu-

bescent, more densely at apical expansions. Pronotum finely, densely punctate,

densely clothed with transverse, appressed pubescence; pro-, meso- and meta-

sternum densely silver-pubescent. Elytra distinctly bicostate on each side, sub-

sutural costae not extending to apex; transverse pubescence lying within limits

of costae, longitudinal bands extending from humeri to apex subopaque, finely

pubescent, often submetallic, lateral margins narrowly brownish; apices bi-emar-

ginate, not produced. Legs with hind tibiae possessing a row of erect bristles

along inside margins. Abdomen densely pubescent except for narrow margins of

segments 2 to 4; last sternite emarginate at apex. Length, 11-18 mm.

Female—Form similar. Antennae slightly longer than body, third segment

VoL. 52, No. 3, JuLty 1976 219

longer than fourth, fifth equal to fourth. Pronotum finely pubescent, pubescence

short, not obscuring surface. Abdomen with last sternite broadly truncate, notched

at middle. Length, 12-18 mm.

Type locality—of sericatus, Iguala, Guerrero, Mexico; moestula, Cutuco da

Union, El] Salvador.

Range.—Sinaloa, Mexico to El Salvador.

This species is easily recognizable by the transverse pubescence of the

elytra and the finely, densely punctate pronotal disk.

Geographic variation in color appears to be clinal. The northern-

most specimens are dark with mostly reddish legs. The elytra, particu-

larly along the outside opaque band are submetallic with greenish to

violaceous overtones. In the vicinity of Mazatlan, Sinaloa, the tendency

is toward all black coloration including the legs. Further south, in Na-

yarit, the elytra become brownish and the legs again reddish. This ten-

dency continues southward with the elytra becoming paler brown, and

only faintly metallic.

Long series of Mexican specimens have been examined from the following

localities: 18 miles S Guamuchil, Sinaloa, 7 August 1964, on Croton culicanensis

(J. A. Chemsak, J. A. Powell); 29 miles S Culiacan, Sinaloa, 23 June 1963 (J.

Doyen) ; 18 miles N Mazatlan, Sinaloa, 28 July 1972 (J. A. and M. A. Chemsak,

A. and M. Michelbacher) ; 9 miles N Mazatlan, 25-28 July 1972 (Chemsaks and

Michelbachers) ; 5 miles N Mazatlan, 27 July and 1 August 1972, on Buddleia

wrightii (Chemsaks and Michelbachers), 5-7 July 1964 (J. E. Martin) 14 km S

Mazatlan, 18 July 1965 (R. Snelling); 4 miles S Villa Union, Sinaloa, 23 June

1963 (J. Doyen) ; Arroyo Santiago, nr. Jesus Maria, Nayarit, 5 July 1955 (B. Mal-

kin) ; 35 miles W Tequila, Jalisco, 22 July 1965 (P. H. Freytag, L. P. Gibson) ;

6 miles S Rio Mexcala, Guerrero, 6 August 1965 (G. H. Nelson) ; Mexcala, Guer-

rero, 29 June 1951 (P. D. Hurd, H. E. Evans) ; Bejucos, Temescaltepec, Mexico, 3

July 1933 (Hinton and Usinger) ; 20 miles S Matias Romero, Oaxaca, 2 July 1964

(A. Raske); 21 miles S Matias Romero, 25 June 1961, on Acacia flowers; 14

miles NW Tehuantepec, Oaxaca, 26 June 1961; 3 miles W Tehuantepec, 9 July

1965 (G. H. Nelson).

LITERATURE CITED

Bates, H. W. 1892. Additions to the Longicornia of Mexico and Central Amer-

ica, with remarks on some of the previously recorded species. Trans.

Entomol. Soc. London, 1892: 143-183, pls. 5-7.

Monne, M. A. and Martins, U. R. 1972. Sobre Triacetelus Bates e Beaveria

Lane. Rev. Brasileira Entomol., 16: 75-78, 1 fig.

Zasciw, D. 1964. Novos Longicorneas Neotropicos. X. Acad. Brasileira Cienc.,

36: 433-440, 5 figs.

New Species of North American Anomala

(Scarabaeidae: Anomalinae)

Rosert W. L. Potts

California Academy of Sciences, San Francisco, California

During revisionary study of the species of Anomala of America north

of Mexico the following seven species were discovered to be new. All of

the new species are apparently from relatively small geographic areas,

and at least two or three are from very limited relictual habitats.

Anomala diabla, new species

Holotype male—Medium brown, head and clypeus red-brown, pronotum with

large central red-brown macula extending to base, enclosing paler area medio-

basally, elytra with narrow complete outer margin and sutural interval red-brown,

this extending forward around scutellum and along base; scutellum almost median

in hue between ground color and maculation. Thoracic sternites finely, not densely

pilose. Protibiae tridentate; larger protarsal claw simple; larger mesotarsal claw

finely but rather deeply cleft well back from apex. Antennal club slightly longer

than stem. Clypeus trapezoidal, more than twice as wide as long; sides moderately

divergent, broadly and strongly reflexed marginally; suture slightly depressed;

disc of clypeus together with triangular area of front notably flattened, coarsely,

irregularly punctate (more reticulate under higher magnification). Vertex with

dense, fine, laterally elongate punctures. Pronotum evenly arcuate laterally,

broadest near middle; anterior angles distinct, almost square; posterior angles

broadly rounded; punctures moderately fine, not dense, but becoming more so

laterally; basal bead fine, complete. Elytra slightly broader behind middle; striae

slightly grooved, regularly, finely punctate, the punctures darker, particularly

toward suture; subsutural interval markedly widest, multipunctate basally; other

intervals subequal, 4th and 6th with scattered punctures; striae becoming irregu-

lar laterally. Pygidium punctate, punctures similar to those of pronotum but more

irregular. Length, 9 mm.

Holotype male, Texas, Devit’s River, near Det Rio, 25 May 1932 (CAS); one

male paratype, Texas, Eagle Pass, 2 June 1923 (Howden).

Holotype placed in the collection of the California Academy of Sci-

ences and the paratype in the collection of Henry Howden.

The paratype is remarkably similar to the holotype, except that the

larger mesotarsal claw is simple, and the elytral striae are more roughly

punctate.

Named for the river where the type specimen was collected. The spe-

cies superficially resembles cavifrons, but is without the elongate clypeus

and carinate clypeal suture. It also has distinctly tridentate protibiae.

These characters, with the simple protarsal claw, distinguish it from the

Tue Pan-PaActFic ENTOMOLOGIST 52: 220-226. JuLy 1976

VoL. 52, No. 3, JuLy 1976 221

somewhat similar delicata, and some specimens of undulata, with which

it might be confused.

Anomala eximia, new species

Holotype male.—Head, pronotum and scutellum dark red-brown-black, elytra

straw-colored; margins, punctures, and two vague irregular bands at middle be-

fore apex brown-black. Venter with sparse, moderately long pale pilosity. Meso-

coxal separation moderate, about width of tarsal segment. Protibia without external

tooth, terminal tooth moderately long, terminal articulated spine small, inconspicu-

ous. Larger protarsal claw finely cleft near middle, upper ramus small, spine-like,

closely appressed; larger mesotarsal claw finely cleft near apex. Metatibia as long

as femora, moderately surate. Clypeus subrectangular, about twice as wide as long;

sides slightly divergent, broadly, strongly reflexed; anterior face thick, vertical,

rounded under to small exposed labrum; suture a little depressed; disc of clypeus

together with triangular flattened area of head shallowly punctorugulose. Anten-

nal club somewhat longer than stem. Pronotum slightly arcuate behind, about same

width at base as at middle, then more strongly arcuate to square anterior angles;

posterior angles broadly rounded; discal punctures rather coarse, irregular, con-

tiguous to separated by 1-2 diameters, not becoming larger nor more dense at sides;

basal bead very fine, entire. Elytra widest about middle; strial series very irregu-

lar, formed by individual punctures intermixed with short and long foveate groups

of punctures; intervals scarcely distinct, but 2nd and 4th broader, 2nd very irregu-

larly multipunctate, 4th with uniseriate row of scattered punctures. Pygidium

finely rugose with sparse, moderately long pilosity. Length, 7 mm.

Holotype male, 8 male paratypes, FLoripa, HicHLANps Co., ARCHBOLD BIOLOGI-

CAL STATION, at Malaise Trap, 12-19 June 1969, R and J Matthews, Collrs. (UGA).

The Holotype is deposited in the collection of the U. S. Museum of

Natural History, with paratypes in the collections of the University of

Georgia and the author.

The species varies in size from 6.5 to 7.5 mm. The clypeus commonly

has the sides parallel or even slightly convergent basally. The elytral

sculpture and coloration is quite variable and the two irregular dark

bands are sometimes lacking.

This is the second small species from Florida that has the protibiae

lacking an external tooth, the other being exigua (Schwarz), however,

while the similarity is interesting this species is not close to exigua, but

belongs in the binotata series.

Anomala hardyorum, new species

Holotype male.—Pale straw-colored; clypeal margin, anterior and posterior pro-

notal margins, sides of scutellum and elytral suture very narrowly reddish-brown;

pronotum and elytra immaculate but variably pale due to irregular sclerotization.

Thoracic sterna moderately pilose. Midcoxae narrowly separated. Anterior tibiae

bidentate, lateral tooth not prominent, formed by excavation to terminal tooth;

terminal articulated spine large and prominent; larger protarsal claw narrow,

222 THE PAn-PactFIc ENTOMOLOGIST

barely perceptibly thickened basally, cleft deep, at about middle, with upper ramus

fine, spine-like, much shorter than lower ramus. Metatibia much shorter than femur,

barely longer than first 3 tarsal segments. Antennal club thin, slightly asymmetri-

cal, notably longer than stem. Clypeus trapezoidal, more than twice as wide as

long; strongly reflexed at margins; anterior face steeply oblique, then rounded

beneath to narrow edge of exposed labrum; clypeofrontal suture flat at sides,

depressed medially; front of head broad, subrectangular; both clypeus and front

finely rugosely punctate. Pronotum widest at base, very broadly subangulate before

middle, sides straight posteriorly, barely arcuate anteriorly to nearly square ante-

rior angles; posterior angles obtuse, narrowly rounded; discal puncturation very

fine with punctures separated by up to 3-4 diameters; basal bead fine, entire,

depressed below discal plane. Elytra broadest near middle; 10 primary striae very

weakly or not at all grooved, indicated by small punctures, each in a larger depres-

sion, these contiguous in a row or separated and irregular; subsutural interval

slightly broader, multipunctate anteriorly, thinning posteriorly and punctures

scarcely reaching apex; 4th interval substriately punctate much of length; 3rd,

oth and 7th intervals narrower but irregular, very slightly costate; micropunctures

generally distributed over elytra. Pygidium with large, shallow, irregular puncti-

form depressions becoming finely rugose in part. Length, 9 mm.

Female——Similar to holotype, but with pygidium much more finely and sparsely

punctate; protibiae strongly bidentate with lateral tooth prominent; antennal club

heavy, subequal to stem; larger protarsal claw evenly terminally cleft, upper ramus

as large as and longer than lower ramus.

Holotype male, Catirornia, ImpertAL Co., 3 mi. NW Gtamis, 12 April 1969,

A. Hardy, (Hardy Collection) ; Allotype, same data, 23 April 1970, Hardy &

Pritchard, (Hardy Collection) ; 93 males, 8 females, same data, 3 April to 5 May,

91 males, 10 females, same except various stations nearer Glamis, various dates

between 26 March and 4 May.

Holotype, allotype and paratypes placed at California Academy of

Sciences; paratypes in the collections of California Department of Agri-

culture, University of Arizona, Dave Carlson, J. M. Cicero, Alan Hardy,

H. F. Howden, G. F. Nelson, Ron Mc Peak, and the author.

Male paratypes varied from 7 to 10 mm, the females from 7 to 9 mm;

both were sometimes a little darker, sometimes with faint pronotal cloud-

ing of a darker hue. The size and length of the upper ramus of the larger

protarsal claw was quite variable in both sexes.

Named in honor of Alan and Wanda Hardy who have made many

trips to the Glamis dunes, studying the remarkable relict fauna and flora

of the area.

Anomala imperialae, new species

Holotype male—10 mm. Light brown; with head, clypeus, anterior and posterior

margins of pronotum, broad sutural and lateral margins of elytra and some elytral

punctures dark brown. Beneath strongly pilose with mixture of short and moder-

ately long hairs. Mesocoxae narrowly separated. Protibiae tridentate; larger pro-

and mesotarsal claws simple. Antennal club distinctly longer than stem. Ligula

Voi, 52, No. 3, Jury 1976 223

with more than normal number of erect setae but not enough to be confused with

penicillate ligula of much smaller Leptohoplia. Clypeus trapezoidal, more than

twice as wide as long; sides moderately divergent; margins narrowly, abruptly re-

flexed; discally with fine transverse rugae and small scattered punctures; front

more strongly, irregularly rugosely punctate. Pronotum almost evenly arcuate at

sides; anterior angles almost square, a little rounded; posterior angles broadly

rounded; discal punctures very fine, shallow, becoming much larger and very

irregular laterally; basal bead complete, very fine laterally, heavier medially.

Elytra broadest about middle; striae not at all or only slightly grooved, formed by

rows of pale to dark irregularly spaced fine punctures; subsutural interval broad,

multipunctate anteriorly; other intervals of variable widths but all relatively nar-

row. Pygidium strongly, irregularly punctate.

Female.—11.5 mm, similar except antennal club heavier, about as long as stem;

larger protarsal claws mixed, the left simple, the right broadly cleft; clypeus and

front more roughly, irregularly punctate; pronotum with two small close-set antero-

medial dark brown maculae.

Holotype and 4 male paratypes, CALIFORNIA, [IMPERIAL Co., WINTERHAVEN, 3-4

April 1961, Maxwell, Collr. (CDA); Allotype and 2 male paratypes, Arizona,

Yuma, 27 April 1962, 3 April 1961, 3 May 1962, Yanez, Mellor and Stedman,

Collrs. (UAZ) ; 8 male paratypes, California, Imperial Co., Bard and Laguna Dam,

5 April to 3 May, Harrison, Akins, Miller and Cicero, Collrs. (CDA, Nelson and

Cicero Colls.) ; 9 male paratypes, Arizona, Yuma Co., Laguna Dam and 5 mi. W

Wellton, 10 Aug and 8 Oct, Nutting & Werner and Fisher, Collrs. (UAZ and

Hardy Colls.); 1 male paratype, Arizona, Huachuca Mtns (USMNH); 1 female

paratype, Arizona, Santa Catalina Mtns, Sabino Canyon, 17 May, Woods,

Collr. (UAZ).

Holotype deposited at California Academy of Sciences, and allotype

in University of Arizona Collection, on indefinite loan to California

Academy of Sciences; paratypes in the collections of the above and Cali-

fornia Department of Agriculture, U. S. Museum of Natural History,

J. M. Cicero, Alan Hardy, H. F. Howden, G. H. Nelson and the author.

Paratypes vary from 8 to 11 mm. About half the males have transverse

wrinkles on the clypeus, the others are strongly, irregularly punctate.

The amount of dark maculation is variable, from almost none to very

broadly margined and with the pair of small pronotal maculae, while

the female has the pronotum immaculate. The female also has the larger

claw of the protarsus broadly cleft, but all males have the claw simple.

Anomala kanei, new species

Holotype male—9 mm. Pale straw-colored, head and clypeus brown-black, pro-

notum with pair of small triangular anteromedial spots dark-brown, scutellum

and elytra with extreme marginal edges red-brown and some clouding of brown

inward from lateral margins. Beneath with short, not dense pilosity. Mesocoxal

separation narrow. Protibiae strongly bidentate, terminal tooth long, slender, exter-

nal tooth long, acute; larger protarsal claw strongly but finely cleft about middle;

larger mesotarsal claw finely, deeply cleft well back from apex. Metatibia shorter

224, THE PAn-PAcIFIC ENTOMOLOGIST

than femora, about as long as first 4 segments of tarsi but distinctly more than

twice as long as width at apex. Antennal club much longer than stem, lamellae

thin, somewhat twisted. Clypeus somewhat elongate, about twice as wide as long,

sides subparallel toward base, very strongly and broadly reflexed to margins;

anterior face strongly rounded under to porrect labrum; clypeofrontal suture deeply

impressed in part; clypeus and front very irregularly rugulose and somewhat

punctate, varying from large punctiform craters to small distinct punctures. Pro-

notum evenly arcuately rounded laterally, widest about middle, both anterior and

posterior angles broadly rounded; moderately densely punctate, punctures shal-

low, fairly large; with faint indication of median line; basal bead distinct, entire.

Elytra a little wider behind middle; striae broadly but not deeply impressed, with

spaced, poorly defined strial punctures; entire discal surface with numerous micro-

punctures; subsutural interval distinctly widest but may be apparent only basally

and apically as single row of large punctures forms substriate line at middle.

Scutellum with slight posteromedian depression. Pygidium densely punctate, but

punctures shallow, moderately large, irregular in shape and definition.

Female——9 mm; very similar, differing primarily in sexual characters: antennal

club about length of stem; protibial dentition still more acute and longer; larger

protarsal claw almost evenly cleft with upper ramus a little smaller and shorter;

larger mesotarsal claw with upper ramus a little larger and longer.

Holotype male, allotype, and 94 male paratypes, ARIZONA, Coconino Co., 2 mi.

S. Moenxopt, 3 July 1972, Andrews and Kane, Collrs. (CDA); other paratypes

from Arizona, Coconino Co.: 3 males, 2 mi. S. Moenkopi, 15 July, J. & L. Shep-

pard, Collrs. (Hardy) ; 1 male, Kayenta, 23 July, Maehler, Collr. (Potts) ; 3 males,

Tuba City, 1 July, 1 Aug, Allen, Cazier, Collrs. (CAS, CNC, and USMNH Colls.) ;

and from Arizona, Navajo Co.; 7 males, 1 female, Aug, Duncan, Collr. (UAZ) ;

1 male, Winslow, 5 July, Johnston, Collr. (CNC); and 2 males, Utah, Kanab, 20

July, Knowlton and Harmston, Collrs. (USMNH).

Holotype and allotype, with paratypes in the collection of the Califor-

nia Academy of Sciences; paratypes also in the collections of the Cali-

fornia Department of Agriculture, Canadian National Collection, Uni-

versity of Arizona, U. S. Museum of Natural History, Alan Hardy, and

the author.

The male paratypes vary from 7 to 9 mm, with head and clypeal color

varying from only a little darker than pale straw ground color through

red-brown to brown-black, the pronotum sometimes becoming immac-

ulate and elytral sculpture sometimes very shallow and faint. The female

paratype is heavier, with darker ground color, but dark maculations

paler.

This attractive small species, appearing closest to antennata, is named

in memory of Fred Kane.

Anomala sabinae, new species

Holotype male—15 mm. Light straw-colored, with clypeus, head, and antero-

medial pronotal macula red-brown, posterior margin of pronotum, lateral margins

of scutellum and narrow sutural stripe of elytra brown-black. Beneath with moder-

VoL. 52, No. 3, Juty 1976 220

ately short pilosity on thorax, abdomen almost glabrous. Mesocoxal separation nar-

row, but almost as wide as a tarsal segment. Protibiae strongly bidentate; larger

protarsal claw strongly cleft with smaller ramus fine, slightly longer than larger

ramus; larger mesotarsal claw with cleft nearly similar; metatibiae strongly surate

and metatarsi exceptionally heavy as in delicata, about twice as wide as other tarsi.

Antennal club slender, about as long as stem. Clypeus trapezoidal, more than twice

as wide as long; sides strongly divergent; very narrowly reflexed at margins; ante-

rior face distinctly oblique to exposed narrow portion of labrum; suture flat; front

of head broad, rectangular; clypeus and front finely rugosely punctate. Pronotum

widest at base with sides evenly arcuately narrowing to square anterior angles; pos-

terior angles a little obtuse, slightly rounded, disc very finely, not densely punctured,

punctures separated by 2-4 diameters, not becoming more coarsely nor more

densely punctate at sides; basal bead obsolescent medially. Elytra broadest about

posterior third; strial series with distinct geminate pairs, the 3rd, 5th and 7th inter-

vals very narrow, a little costate, and a similar 9th interval barely discernible;

subsutural and other alternate intervals broad, multipunctate; strial punctures

somewhat smaller than those of intervals, but all formed by a micropuncture in a

pit. Pygidium very finely and densely, but shallowly strigulose.

Female—14.5 mm; similar, however central pronotal macula complete to base,

enclosing pale mediobasal area; pygidium not strigulose, but with very fine, shal-

low, laterally much elongated punctures, about 2-4 diameters apart; elytra with

small dark spot over humeral umbone; other variation as normal for sex: tarsal

claws more evenly cleft, protibiae more strongly dentate, antennal club heavier,

but about as long as in male.

Holotype and allotype, Arizona, SANTA CATALINA Mtwns., SABINO CANYON, in

oak-mesquite-sycamore association along permanent stream, 2500’, 26 July 1948,

Werner & Nutting, Collrs. (UAZ). All paratypes, 28 males, 18 females, also from

Sabino Canyon, various dates between 21 June and 4 September, Werner & Nut-

ting, Butler, Johnson, Potts and Potts, Cicero, O’Brien, Hovore, Collrs. (UAZ,

Potts, Cicero, Woodruff, Howden, and Carlson Colls.)

Holotype and allotype in the University of Arizona collection, on

indefinite loan to the California Academy of Sciences, and paratypes in

the collections of the Academy, as well as in the U. S. Museum of Natu-

ral History, University of Arizona, Carlson, Cicero, Hardy, Howden,

Woodruff, and the author.

The paratypes vary between 13 and 17 mm. and sometimes become

medium brown, the males usually with the pronotal macula antero-

medial, rarely reaching to the base, while in the females the macula is

almost always complete to base. Also, the males rarely show the dark

macula over the humeral umbone, while it is usual in the female.

The species is very similar to, but consistently larger than delicata,

and it has relatives of equal size in southern Mexico.

Anomala suavis, new species

Holotype male——8 mm. Pale straw-colored, with dark brown-black head and clyp-

eus, pronotum wth large central dark brown macula complete to basal margin,

226 THe Pan-PaciFic ENTOMOLOGIST

scutellum and narrow entire margin of elytra dark brown. Moderately short pilose

beneath. Mesocoxal separation narrow. Protibiae strongly bidentate, terminal tooth

slenderly elongate, external tooth acute, longer than normal in genus; larger pro-

tarsal claws distinctly but finely cleft about middle; larger mesotarsal claws finely,

deeply cleft well back from apex; metatibia notably short, shorter than femur,

about length of first 4 segments of tarsus, only barely surate, but definitely more

than twice as long as width at apex. Antennal club much longer than stem. Clyp-

eus elongate, less than twice as wide as long, sides strongly convergent basally,

strongly and broadly reflexed to margins; anterior face thick, strongly rounded

under to small, slightly projecting anterior aspect of labrum; clypeofrontal suture

slightly depressed; irregularly, finely rugosely punctate over clypeus and a small

triangular area of front of head; head very finely punctate over vertex. Pronotum

very evenly arcuately rounded laterally, widest behind middle; anterior angles

obtuse but distinct; posterior angles very broadly rounded; with fine, shallow

median line; discal punctures fine, not dense centrally but becoming more so later-

ally; basal bead not well-defined medially. Scutellum with distinct posteromedial

depression. Elytra widest a little behind middle; striae irregularly, somewhat

broadly impressed, with punctures often indistinct, fine to large; subsutural inter-

val broadest, with scattered very large rough punctures becoming almost striate

series at middle but not continuing to apex. Pygidium with scattered, shallow,

irregular punctures.

Holotype and one male paratype, Texas, WINKLER Co., 4 mi. NE Kermit, 5 June

1971, O’Brien and Marshall. Holotype on deposit at the California Academy of

Sciences and paratype in the collection of the author.

Another of the species that is relatively close to cavifrons, but also

another with well cleft protarsal claws, and the two known specimens are

extremely clearly and neatly marked. In this respect they are quite unlike

the usual cavifrons specimens that show some melanistic maculation.

A New Species of Evodinus From Wyoming

(Coleoptera: Cerambycidae)

ArtHuR E. LEwIs

1360 Paseo Redondo, Burbank, California 91501

Three specimens of a distinct Lepturine Ceramybicid from Beartooth

Plateau, Park County, Wyoming, were received for identification from

the University of Colorado Museum by Dr. U. N. Lanham. A fourth

specimen, included in the following description, was made available

by Dr. John A. Chemsak, University of California, Berkeley. It was

collected in Wyoming in 1897, is faded, and in poor condition, but is

unmistakably conspecific with the original three.

Evodinus lanhami, new species

Female—Form moderate sized, robust; color piceous to black; antennae except

scape, tibiae and tarsi reddish brown, elytra yellow with black markings. Head

trapezoidal, widest across eyes, only slightly narrower than widest portion of

pronotum; frons and tempora shining; pubescence moderately depressed, not

obscuring surface; punctation densely, coarsely, contiguously to rugosely punctate.

Antennae 11 segmented, robust, extending to basal third of elytra; scape black,

remainder of segments reddish brown with apex of fourth through eleventh sub-

opaque, distinctly minutely pubescent without apical cilia; third segment sub-

equal to scape, longer than fourth. Remaining segments subequal.

Pronotum about as wide as long, robust, shining, strongly obtusely tuberculate

laterally; coarsely, densely, contiguously punctate; apex narrower than base; a

poorly defined narrow median line extends from apical sulcus to basal margin;

pubescence moderate, suberect to erect, not obscuring surface. Prosternum moder-

ately densely pubescent and rugulose; meso- and metasternum moderately pubes-

cent, surface not obscured. Elytra less than twice as long as broad, sides to apical

third slightly tapering to subparallel, then gradually rounded to suture; apices

slightly dehiscent; densely, coarsely, confluently puntate with punctation becom-

ing finer apically; pubescence golden, short, uniform, and semirecumbent; color

yellow with black markings as follows: Basal margins triangularly expanded

posteriorly to envelope scutellum, then extending to apex as a sutural vitta, and

continuing over humeri obliquely posteriad as a maculation connecting or not

with the suture; a small medial lateral spot; a larger post median black macula

which may be expanded to reach suture and apices. Legs slender, femora black;

tibiae and tarsi lighter; pubescence fine, uniform, not obscuring surface. Abdo-

men moderately pubescent, shallowly densely punctate; 5th sternite broadly

rounded; 5th tergite subtruncate, narrowly notched. Length 13-14 mm; width

5.0 mm.

A Key to tHe NortH AMERICAN SPECIES OF EVODINUS

1. Pronotum dull, distinctly longer than wide; punctation fine; pubescence mod-

erately dense. Elytra with humeri yellow, without a distinct black sutural

PEER a. ee re eg ik On ieee en Cpe ieiere ls 9 e ron Se monticola

Tue Pan-PActFic ENTOMOLOGIST 52: 227-228. JuLy 1976

228 THE PAn-PactFic ENTOMOLOGIST

Pronotum shining, distinctly coarsely punctate, about as wide as long; pubes-

cence moderate. Elytra with basal margin black extending over humeri. —_.

lanhami

E. lanhami can be easily recognized from its other U. S. counterpart

E. monticola by its more robust form, subparallel sided elytra, trans-

verse pronotum, and elytral maculation. It is apparently taken at high

altitudes.

Holotype female, BEARTOOTH PLATEAU, PARK County, Wyomine, 15-17 June,

1961.

Paratypes, three females—two with above data; one, Pammel, Rapid Creek,

Wyoming, 7-97, R. Hopping Coll. The type is deposited in the Colorado Univer-

sity Museum. Paratypes are deposited in the California Insect Survey, Berkeley;

the California Academy of Sciences, San Francisco; and the collection of the

author.

ACKNOWLEDGMENTS

The author wishes to thank Dr. John A. Chemsak, University of

California, Berkeley, for examining the material and comparing it

with old world members of the genus, and Dr. U. N. Lanham, for

whom the species is named, for making the original specimens available

for study.

RECENT LITERATURE

A BIBLIOGRAPHY AND A KeryYWorD-IN-CONTEXT INDEX OF THE CERATOPOGONIDAE

(DrpTteRA) FROM 1758 To 1973. W. R. Atchley, W. W. Wirth and C. T. Gaskins.

Texas Tech University Press, Lubbock, 1975. 300 pp. $4.00 (paper).

The title accurately describes the contents of this computer produced literature

guide, which will unquestionably be useful to researchers interested in this family

of flies. The bibliography (number of entries not mentioned) occupies 161 pages.

Single line keyword indexing comprises the remaining 130 pages. A short intro-

duction provides instructions for use. An exact description of the data base used

to construct this bibliographic aid is not provided.—£ditor.

Predatory Behavior of the Spider Wasp,

Chalybion californicum

(Hymenoptera: Sphecidae)

Ro.uin E. CoviLLe

Division of Entomology and Parasitology

University of California

Berkeley, Cal.

94720

Chalybion californicum (Saussure) (= caeruleum of authors) is a

common steel-blue mud-daubing spider wasp found throughout most of

North America from southern Canada into Mexico (Bohart and Menke,

1963). Its nesting biology has been studied by Rau (1928a, 1928b),

but only a few isolated observations of its predatory behavior have been

reported. The Peckhams (1898) observed a species of Chalybion prey-

ing on Araneus cornutus Clerck (= Epiera strix). The wasps crawled

into cracks and corners of cottage walls and dislodged the spiders. They

captured spiders with their mandibles and forelegs and stung them at

that time or after taking flight. Rau (1928a) observed a C. californi-

cum attempting to capture a Lycosa spider by landing on its back. How-

ard (1922) reported that Schwarz (presumably Herbert F. Schwarz)

observed C. californicum entangling itself in webs and causing spiders

to run out of their retreats. The wasps would then free themselves and

pursue the spiders. Frost (1944) reported this species engaged in the

same behavior.

Observations reported here were made on eight days from June 27

to July 17, 1974, on the campus of the University of California, Berke-

ley, and confirm the pattern in the accounts of Howard (1922) and

Frost (1944). C. californicum females employ an efficient form of

aggressive mimicry in capturing their prey. This activity took place on

one Garrya elliptica Doug. bush harboring a large population of imma-

ture Zygiella x-notata (Clerck), an araneid spider. Z. x-notata con-

structed orb webs that usually had one of the dorsal pie-slice shaped

sectors missing. From the center of each orb web a strong signal line

ran to a tubular retreat, usually above the orb. The leaves of G. ellip-

tica are oblong with the lateral edges curled under. The sheltered under-

sides of the leaves provided suitable retreats for Z. x-notata, and open-

ings in the foliage provided many locations for their webs. During the

day, the spiders remained in their retreats and used their front legs to

monitor the signal line for vibrations transmitted from the orb. Chaly-

Tue Pan-Paciric ENTOMOLOGIST 52: 229-233. JuLy 1976

230 THE Pan-PActFic ENTOMOLOGIST

Fic. 1. Chalybion californicum female pulling and vibrating the web of an

immature Zygiella x-notata.

bion females were about four or five times as large as Zygiella. In

approximately eleven hours of observations at least three individual

wasps vibrated webs more than a hundred times and captured more

than twenty spiders.

In a typical sequence a Chalybion female flew into openings in the

VoL. 52, No. 3, Juty 1976 231

foliage where she then slowly flew into the exposed face of an orb web

and alighted. After remaining motionless for a few seconds, she began

pulling on the web with her legs (Fig. 1), mimicking an entangled

insect. After several seconds of pulling and vibrating a web, she took

flight and repeated the performance on the next orb she encountered.

This continued until a spider, responding to the vibrations of its web,

ran down the opposite side of the orb from the wasp. Just as a spider

was about to reach her, the wasp lunged forward, seized the spider

with her mandibles and forelegs, curled her abdomen up underneath

her body and stung the spider for several seconds. The exact placement

of the sting could not be observed. One spider avoided capture by

dropping off the web just as a wasp lunged for it, and the wasp did not

pursue it. None of the wasps became entangled in a web.

After a wasp stung a spider, she presumably transferred it to the

other side of the web, but this could not be observed. She then carried

it across the web to a leaf or stem and then manipulated and rolled the

spider with her forelegs and mandibles, in the manner described by the

Peckhams (1898). Occasionally a spider was stung again after manipu-

lation. One wasp was disturbed while manipulating a spider. She flew

to another leaf and started manipulating the spider again but then

dropped it, apparently by accident. Rather than trying to find the spider,

she groomed herself for a few seconds and then flew to two more orbs

before flying away.

Wasps frequently interrupted their hunting and landed on leaves to

groom themselves, and sometimes they walked over the leaves and

stems. Twice wasps walked to the lower edge of an orb and used their

forelegs to vibrate the web. One spider responded to these vibrations

and was captured by the wasp. Several times wasps walked on leaves

that had spider retreats underneath. They often crawled around the

side of such a leaf, as if examining the retreat, but none attempted cap-

ture of spiders in their retreats.

About half the spiders did not respond to the vibrations of their

webs. A smaller proportion left their retreats but then returned before

approaching the wasp. One spider with a relatively exposed web near

the top of the bush repeated this sequence three times in response to

one wasp. Spiders were never pursued. They were captured only if they

ran to the wasp. The wasps were unable to perceive whether a web

contained a spider because they often visited vacant webs.

When attempting to alight on the webs the wasps flew slowly with

their legs outstretched and hanging below their bodies. In this position

the tarsi were oriented approximately perpendicular to the direction of

232 THE PAn-PAcIFIC ENTOMOLOGIST

flight and anterior to the body and wings, enabling contact of the web

by the tarsi, rather than the body or wings. After alighting, wasps used

their tarsal claws to hang onto and pull the webs (Fig. 1). Other con-

tact points may be the arolia and the ventral surface of the tarsal seg-

ments.

Muma and Jeffers (1945) found that Theridiidae and not Araneidae

were the most common prey of C. californicum. They concluded that

the species of spider preyed upon depends on where the wasp hunts.

This is usually on or near the ground and around human habitation,

the haunts of most theridiids, but in several cases they found that the

wasps only collected foliage inhibiting spiders. This suggested that

once a wasp captured a spider, it returned to the same area on sub-

sequent trips. In the observations reported here two female wasps were

marked with a nontoxic paint at the Garrya bush, but only one of them

was seen again, when it returned the following day and capiured a

spider.

This bimodal distribution of prey type suggests flexibility in preda-

tory behavior of C. californicum because there are important biological

differences between many theridiids and araneids (Bristowe, 1971).

Most theridiid webs are 3-dimensional mazes; most araneid webs are

2-dimensional orbs. When araneids subdue their prey, they usually

run to the entangled insect and immediately bite it or wrap it first in

silk. Many theridiids often keep their distance and cast silk threads

over their prey. Only after the insect is covered with silk and immobi-

lized, do they approach to bite it. Recalling the observations of the

Peckhams (1898) and Rau (1928a), it may be that C. californicum

becomes conditioned to certain prey species and this is reflected in vari-

ations in its predatory behavior.

ACKNOWLEDGMENTS

I wish to thank Mr. Charles Griswold for identifying the spiders and

Dr. John L. Strother of the Botany Herbarium, University of Califor-

nia, Berkeley, for identifying the Garrya bush involved in the study. I

also wish to thank Dr. Howell V. Daly and Dr. Jerry A. Powell of the

Department of Entomological Sciences, University of California, Berke-

ley, for their helpful comments and criticisms.

LITERATURE CITED

Bowart, R. M., anp Menke, A. S. 1963. A reclassification of the Sphecinae.

Univ. Calif. Publ. Entomol., 30:91—-182.

BristowE, W. S. 1971. The World of Spiders. Rev. ed. Collins, London. 304 p.

VoL. 52, No. 3, JuLy 1976 230

Frost, S. W. 1944. Notes on the habits of Monobia quadridens (Linn.).

Entomol. News, 55:10-4.

Howarp, L. O. 1922. The Insect Book. Doubleday, Page and Co., Garden City,

429 p.

Muma, M. H., anp Jerrers, W. F. 1945. Studies of the spider prey of several

mud-dauber wasps. Ann. Entomol. Soc. Amer., 38:245-55.

PeckHAM, G. W., AND PeckKHAM, E. G. 1898. On the instincts and habits of the

solitary wasps. Wisconsin Geol. Nat. Hist. Surv., Sci. Ser. 1, Bull. 2.

245 p.

Rau, P. 1928a. The nesting habits of the wasp, Chalybion caeruleum. Ann.

Entomol. Soc. Amer., 21:25-35.

Rau, P. 1928b. Field studies on the behavior of the non-social wasps. Trans.

Acad. Sci. St. Louis, 25:325—489,

BOOK NOTICE

RHOPALOCERA Directory. John R. Beattie. 370 p. (approx.) including introduc-

tion and detailed instructions for use. 128,800 entries plus two supplements of

5,000 entries and 36,700 entries. Offset reproduction, soft cover. $40.00 to libraries,

$30.00 to individuals on 10-day free trial examination. Available June, 1976, from

JB Indexes, 2377 Virginia Street, Berkeley, California 94709.

This first volume in an ambitious undertaking to be called the Insecta Directory

is a computerized index to all generic and specific butterfly names which have

appeared in the Zoological Record Systematic Indices between 1864 and 1971.

It also includes names from the analogous German publication, Berichte tiber die

wissenschaftlichen Leistungen im Gebiete der Entomologie for the period 1834—

1863. The index supplies the year and page number for each reference to each

name in a convenient, easily scanned format, enabling the user to quickly locate

the pertinent references in the Record or in Berichte, without searching each

annual volume.

This should be an extremely valuable literature aid for taxonomic work. Since

the Zoological Record and Berichte concentrate on new-name and status-change

references, with only selective inclusion of other references, it will not be possible

to construct complete synonymies using the directory. Nevertheless the Rhopaloc-

era Directory will provide an advanced point of departure for many projects, par-

ticularly those oriented towards the world literature, and will also supplement the

Nomenclator Zoologicus in placing generic names to family. Additional volumes

are planned to eventually include all orders of insects.—EpITor.

Fannia thelaziae, A New Species of Eye-Frequenting Fly

of the benjamini group from California and

Description of F. conspicua Female?

(Diptera: Muscidae)

Wituiam J. TURNER

Washington State University

Fannia benjamini Malloch is a familiar eye-frequenting fly common to

dry chaparral and oak woodland areas throughout California. Its annoy-

ing habit of frequenting the face and especially the eyes of animals in

search of moisture and other substances has made F. benjamini the

prime suspect as the intermediate host of the mammalian eyeworm,

Thelazia californiensis Price (Winkler and Wagner, 1961). The eyeworm

infests a variety of mammal hosts including deer, rabbits, dogs and

man. Although developing stages of the worm were reported in F.

benjamini (Burnett, et al., 1957), recent transmission experiments pro-

vided no evidence of its association with this fly as an intermediate host

(Weinmann, ef al., 1974).

In recent studies of the biology and oviposition behavior of Fannia

species laboratory colonies were established at the University of Calli-

fornia (Berkeley) (Anderson, J. R., C. J. Weinmann and P. Rubtzoff,

unpubl. data). The original colony stock consisted of wild-caught flies

captured at three sites in central and northern California during 1968.

These laboratory colonies of flies were initially thought to consist only

of F. benjamini, but minor differences in the eggs eventually produced

suggested that a mixed colony with as many as three species was in-

volved (op. cit.). Subsequent isolation and rearing of each type of egg

yielded pure colonies of each species, Ff. benjamini, F. conspicua Mal-

loch, and the new species described here. Only the latter proved to be

suitable for the development of eyeworms to the infective stage (Wein-

mann, et al., 1974).

Chillcott (1960) in his monographic revision of the Fanniinae recog-

nized nine Nearctic species belonging to the Fannia benjamini group

which could be divided further into three subgroups. Fannia clavata

Chillcott and micheneri Chillcott were placed in the vittata subgroup,

setifera Chillcott constituted the setifera subgroup, and six species were

included in the benjamini subgroup: namely F. arizonensis Chillcott,

F. benjamini, F. conspicua, F. neotomaria Chillcott, F. operta Chillcott,

and F. tescorum Chillcott. Species in the latter subgroup are distin-

1 Scientific paper 4561, College of Agriculture Research Center. Research was conducted under

Project 9043.

Tue Pan-Pactric ENTOMOLOGIST 52: 234-241. Juty 1976

VoL. 52, No. 3, Juty 1976 235

guished from other Fannia by their yellow basal antennal segments

and palpi, hind tibiae bearing one anterodorsal and two anteroventral

bristles, and trimaculate abdomen. Additional characters uniting this

subgroup include similarities in male genitalia and female spermathecae.

To facilitate comparisons between species in the benjamini subgroup

treated here, the format for descriptions and illustrations follows that of

Chillcott (1960) and the key to species of the subgroup is based pri-

marily on characters used by him. The key to males is modified to in-

clude F’. tescorum which was inadvertently omitted from the original key

(see Chillcott, p. 43, couplet 15). Standard abbreviations were used for

bristles and leg segments and include: anterodorsal (ad); anteroventral

(av) ; posterodorsal (pd) ; posteroventral (pv); dorsal (d) ; acrosticals

(acr) ; dorsocentrals (dc) ; coxa (C); femur (IF); and tibia (T).

Fannia thelaziae new species

(Figs. 4-8)

Mate: Head.—Front with nine, strong bristles. Parafrontals narrow (0.04 increas-

ing to 0.80 mm), gray pollinose. Parafacials narrow, shining gray, without setulae.

Occipital bristles short, regular; postoccipital setulae on lower third, sparse.

Second antennal segment yellow; third segment brownish black or sometimes very

narrowly yellowish at base, arista brown and weakly pubescent. Palpus yellow,

subspatulate. Theca normal.

Thorax.—Evenly brown pollinose, vittae only faintly indicated behind when

viewed from rear. Scutellum concolorous with scutum, laterobasal spots distinct.

Acrosticals (Acr) evenly triserial. Prealars one or (usually) two strong, short setu-

lae; stigmatal setulae absent. Mesopleural bristling distinctly stronger anterodor-

sally. Squamae pale, lower with margin brownish and slightly larger and projecting

more than upper; halter yellow.

Abdomen.—1.5 by 1.8 mm. Basal three segments distinctly translucent yellow

laterally, grayish white pollen of darker portions reduced in these areas. Median

vitta linear, dark brown, confluent with broad concolorous lateral brown marks

on posterior area of basal segments, dark color encompassing entire fifth and

following segments, color often obscured by grayish pollen.

Legs—Legs brown, femora darkly infuscated basally; tibiae and tarsi black.

F; pvs weak on basal half. Ti ads weak, median bristle absent. Fs avs on basal

half 6 to 8, equally spaced, strong, none reduced to setulae, 3 median bristles

stronger, followed by shorter and close-set comb of 14 to 16 bristles to apex;

pvs with single strong bristle basally followed by shorter, partially double row

of mixed bristles on basal half, then 6 to 8 stronger ones to three quarters, then

decreasing in size to a close-set preapical comb of 6-10 short bristles, the comb

tending slightly postero-dorsally. Ts with biserial ventral mat, half tibial diameter

on apical two-thirds, slightly stronger and longer on apical half: C; lacking pv

setulae. F; (Fig. 4) normal in shape, avs prostrate becoming setulose on basal half

and two (rarely three) strong, separated preapical bristles; pvs basally fine then

with 8-10 long, slender bristles loosely clustered at two-thirds extending basad along

the posterior surface as slender erect setulae. T; with two avs, one ad and no pv

bristles.

236 THE PaAn-PaciFIc ENTOMOLOGIST

0.5mm

ees SERENE a Sag

~ Be Ue ee Se aS

MO SE oo ee

Fries. 1-5. Fic. 1. Hind coxa (posterolateral view) of F. conspicua showing

position of pv bristles. Fics. 2 and 3. Hind femora (anterior view) of F. conspicua

male and female respectively. Fics. 4 and 5. Same for F. thelaziae male and female

respectively.

Hypopygium.— (Figs. 6, 7). Cercal plate simple, divided basally, but fused

apically, slightly elongate, bristling weak. Surstylus broad and simple, elongate

and parallel-sided, without swelling at base in side view. Baciliform process

strongly spiraled, heavy.

FEMALE: Head.—Parafrontals brownish gray pollinose, wider than frontal vitta

and convex along inner margin. Setulae uniserial, extending down onto parafacials

well below aristal base. Third antennal segment entirely brown or very narrowly

yellow at base. Palpus bright yellow, distinctly spatulate. Otherwise as in the male.

Thorax.—Distinctly gray to faintly brownish gray pollinose with single in-

distinct narrow brown vitta along acr row, lateral vittae present only as spots about

bases of dc bristles and irregular brown markings around the intraalar row.

Scutellum with two small, indistinct brown spots subbasally. Squamae with con-

colorous margins.

Abdomen.—Integumental color distinctly yellow on basal half of second and

usually on third segment of abdomen, remainder gray pollinose with discontinuous

median vitta and large brown lateral spots. Spermathecae two (Fig. 8).

Legs—yYellowish brown. Fi pvs evenly decreasing to base. Fs avs and pvs

with weak suberect setulae. Ts medially with one each av, ad, pv and p subapical

bristles, all subequal to tibial diameter, and 0-2 weak pvs. F; (Fig. 5) avs and pvs

setulose except for two stronger preapical avs; T; ads and avs 1-2, limited to one

median bristle each, two d bristles longer, one median and one subapical.

Type Matertat.—Holotype: Male, Hastings Reservation, Monterey County, Cali-

fornia; laboratory reared from egg of female captured 18-IX-1969 (C. J. Wein-

mann). Allotype female, same data as holotype.

Paratypes, 368 ¢ 6,212 9 9. Catirornia: Russelmann Park (nr. Mt. Diablo),

Contra Costa County, 2 9 2 10-IX-1969 (P. Rubizoff); 17 ¢ 4,4 2 9, laboratory

reared from eggs of females captured 10-IX-1969. University of California, Hop-

land Field Station (4 mi. east of Hopland), Mendocino County, 5 @ @ (11-18)-

IX-1969 (D. Westrom), 1 @ 2-VI-1970, 1 @ 9-X-1969, 2 99 22-X-1969, 1 @

29-VI-1970, 1 @ 14-VIT-1970 (P. Rubtzoff); 3 ¢¢4, 2 @@ laboratory reared

VoL. 52, No. 3, JuLy 1976 at

Fics. 6-9. Fic. 6. Male hypopygium (posterior view) of F. thelaziae. Fic. 7.

Same in profile. Fic. 8. Female postabdomen (ventral view) and spermathecae of

F. thelaziae. Fic. 9. Same for F. conspicua.

from eggs of females collected (11-18) -[X-1969. Hastings Reservation, Monterey

County, 42 9 9 18-IX-1969, 7 @ 9 13-VIII-1970, 2 @ 9 22-IX-1970 (C. J. Wein-

mann); 347 ¢4, 156 @@ laboratory reared from eggs of females collected

18-IX-1969. Guerneville, 5.5 mi. E., Sonoma County, 1 2 9-[X-1970 (P. Rubtzoff) ;

1 ¢,1 @ laboratory reared from eggs of female captured 9-IX-1970.

Holotype, allotype and 10 paratypes deposited in the California Academy of

Sciences, San Francisco. Additional paratypes have been placed in the American

Museum of Natural History, New York; British Museum (Natural History), Lon-

don; California Department of Public Health, Bureau of Vector Control, Berkeley;

Canadian National Collection, Ottawa; Florida State Collection of Arthropods,

Gainesville; Oregon State University, Corvallis; United States National Museum,

Washington D.C.; University of California, Berkeley; Washington State University,

Pullman; and Zoological Museum, Copenhagen.

Males of this species possess characters intermediate between several

recognized species in the benjamini subgroup. In the male, the dark

brown mesoscutum of F. thelaziae is typical only of F. benjamini while

the yellowed abdomen and leg bristles suggest some similarity to F.

arizonensis. Both F. thelaziae and F. arizonensis have several features

of male genitalia in common, including the basic outline of the dis-

tistylus and the strongly curved bacillus process. In fact, the dististylus

of F. thelaziae differs from that of F. arizonensis only in the absence of

the basal swelling.

All females of the benjamini subgroup appear quite similar to each

other, except perhaps to F. benjamini itself which lacks the typical yellow

lateral markings at the base of the abdomen. In F. conspicua the mark-

238 THE PANn-PAcIFIC ENTOMOLOGIST

ings are present but appear yellowish gray and are sometimes quite

difficult to see. Fannia thelaziae females most closely resemble females

of arizonensis. The spermathecae are similar in shape, size and struc-

ture in both species.

Both sexes of F. thelaziae lack posteroventral bristles on the hind

coxae although they are present in all other species included in the

benjamini subgroup by Chillcott. Even though lacking these bristles,

F. thelaziae is retained in the benjamini group on the basis of its color

pattern in both sexes, and the structure of the male genitalia and female

spermathecae. Chillcott suggested an affinity between the benjamini and

pretiosa groups. This latter group is characterized in part by unbristled

metacoxae. The absence of these bristles in F. thelaziae would seem to

strengthen the association of these groups.

As this species has now been incriminated as the vector of the mam-

malian eyeworm, Thelazia californiensis (Weinmann, et al., 1974), the

name F’. thelaziae seems most appropriate.

ADULT SEASONAL AcTIVITY.—Collecting dates extend from 2 June to

22 October. Most specimens taken in the field were captured in early

to mid September.

DistripuTion.—Although collection records of this species are

limited, the species appears to occur primarily at the margins of oak

woodland areas of the San Francisco Bay Area and in adjacent counties

from Mendocino County in the north to Monterey County in the south.

At each locality sampled, F. thelaziae was sympatric and synchronic

with both F. benjamini and F. conspicua.

FANNIA CONSPICUA Malloch

(Figs. 1-3 and 9)

Fannia conspicua Malloch, 1913: 624.

FemaLe: Head.—Parafrontals brownish gray pollinose, about as wide as frontal

vitta and convex on inner margin. Parafrontals tending to silvery gray; setulae

uniserial, extending down onto parafacials well below aristal base. Oral membrane

yellowish brown often making the oral margin beneath antennae appear yellowish.

Third antennal segment broadly yellow at base, especially along inner ventral

margin; base of arista yellowish. Palpus bright yellow, distinctly spatulate.

Thorax.—Gray pollinose, with faint narrow brown median vitta; lateral vitta

reduced to dark brown spots about the bases of strong dcr bristles and irregular

markings in intraalar rows. Scutellum with two more or less distinct brown spots

subbasally. Squamae and their margins concolorous.

Abdomen.—Basal two or three segments faintly yellowish gray becoming gray

laterally and on distal half or entirely gray with no hint of yellow coloration;

remainder of abdomen gray pollinose with discontinuous brown median and large

brown sublateral spots. Spermathecae two (Fig. 9).

VoL. 52, No. 3, Juty 1976 239

Legs.—Yellowish to brownish yellow; base of Fi brownish, tarsi black, pvs

strong apically, evenly decreasing to base. F: avs and pvs present as regular,

weak and suberect setulae; T2 medially with one each av, ad, pd and subapical d,

all subequal to tibial diameter, and 0-2 pvs appearing shorter and weaker. Fs

(Fig. 3) avs and pvs reduced to setulae except for two or three stronger preapical

avs; T; avs 1 or 2, ads one median subequal to tibial diameter, d two, stronger and

definitely longer than tibial diameter.

Malloch (1913) described F. conspicua from a single male specimen taken at

Williams, Arizona. Females of this species are similar to most other females

included in the benjamini subgroup. They are most easily recognized by their

particular leg bristle patterns cited in the key. The yellow sublateral markings

at the base of the abdomen in this species are quite variable. They are usually

apparent in most specimens but in a significant number the color tends to

yellowish gray and the marks appear restricted to the extreme lateral margins

where they are very difficult to see. In still others the basal segments are

entirely gray. In these latter individuals the yellow on the base of the third

antennal segment, as is characteristic of F. conspicua, appears extensive and

nearly always includes the base of arista. These specimens then, while looking

much like females of F. benjamini, may be recognized by this antennal character.

ADULT SEASONAL AcTIVITY.—California records of F. conspicua range

from 14 July to 29 October. A single Arizona record (no collecting

date for the type is given) cited by Chillcott (1960) is considerably

earlier (12 April).

DIsTRIBUTION.—Chillcott (1960) recorded the species from Arizona

and cited but two records, the type locality and Tempe. Since his study

this species has been collected at a few locations in northern and

central California, in particular Mendocino, Contra Costa and Monterey

Counties. Fannia conspicua will probably be discovered at a number

of localities between the extremes of its present known distribution.

MATERIAL EXAMINED.—CALIFORNIA: Russelmann Park (nr. Mt. Diablo), Contra

Costa County, 2 @ 2 29-VII-1969, 1 @ 10-IX-1969 (P. Rubtzoff); 1 2 laboratory

reared from egg of female captured 10-IX-1969. Hopland University of Calli-

fornia, Hopland Field Station (4 mi. east of Hopland), Mendocino County, 5 2 9

(11-18) -[X-1969, 3 29 2 25-IX-1969, 4 9 2 30-IX-1969 (D. Westrom), 3 Q 29-X-

1969, 1 g, 1 @ 14-VII-1970, 1 @ 21-VII-1970 (P. Rubtzoff); 12 ¢ 4, 56 9

laboratory reared from eggs of females captured (11-18) -[X-1969, Hastings Reser-

vation, Monterey County, 4 2 Q 18-IX-1969 (C. J. Weinmann) ; 124 ¢ 3, 204 9 9

laboratory reared from eggs of females captured 18-IX-1969.

Female specimens of F. conspicua have been placed in the American Museum

of Natural History, New York; California Academy of Sciences, San Francisco;

British Museum (Natural History), London; California Department of Public

Health, Bureau of Vector Control, Berkeley; Canadian National Collection, Ottawa;

Florida State Collection of Arthropods, Gainesville; Oregon State University, Cor-

vallis; United States National Museum, Washington, D.C.; University of California,

Berkeley; Washington State University, Pullman; and Zoological Museum, Copen-

hagen.

240

THE PAN-PAcIFIC ENTOMOLOGIST

Kry To NEARCTIC SPECIES OF THE FANNIA BENJAMINI SUBGROUP

10.

11.

12.

(modified from Chillcott, 1960)

Eyes contiguous or nearly so (males) 2

Eyes distinctly separated (females) 8

Thorax dark brown pollinose, not noticeably vittate _ 3

AEs «OAS UP ER EOS rceetc cee ch Ee lta i tg ge oe 4

Abdomen dark gray with pollen basally; third antennal segment black;

Cs pvs present (California) benjamini Malloch

Abdomen yellow laterally on basal segments; third antennal segment yellow

narrowly at base, rarely all black; Cs pvs absent (California) —..

opi Bake NN elec Sees tee Ty act ede Em SOE OR en eee oon, thelaziae n. sp.

F; avs with a distinct cluster of longer bristles medially, only one strong

Pree eas rsh eee, ede ace ee ek en I et ce ET al ne la 5

F; with two to four strong preapical avs, no strong avs medially _..... 6

F; avs and pvs at least as long as femoral diameter where situated (Ari-

CONE ec Et a 611: eee ee ec ee ee eee eee conspicua Malloch

F; av and pv bristles distinctly weaker, the pvs never as long as femoral

diameter (New Mexico) _ neotomaria Chillcott

F; with four strong preapical avs (California) _. operta Chillcott

F; with two (rarely three) strong preapical avs 7

F; with four to six strong pvs forming a tight cluster which does not grade

into setulae basad (Arizona) __ tescorum Chillcott

F; with four or five slender pv bristles in loose cluster continued basad

as erect setulae (Arizona) — arizonensis Chillcott

Basal abdominal segments dark, overlaid with gray pollinosity 9

Basal abdominal segments yellowish, at least yellowish gray laterally;

if abdomen is entirely gray, then arista yellowish basally 10

Femora largely infuscated; thorax mostly dark brown (California) ___.

oS ge Se Lacan te I ra ENS Sada EAN Be Oh RL operta Chillcott

Femora mostly yellowish; thorax gray pollinose (California) —

re dee A et TLS as re OA ee al Ee benjamini Malloch

Cs without pv bristles; third antennal segment yellow narrowly at base,

arista entirely dark (California) thelaziae n. sp.

Cs; with one or two pvs (Fig. 1); arista and third antennal segment

Pirelli Tia A ce at rama dened eaoncnneensnihgeniataaoiacas Atak mh aa 11

Te median bristles longer than tibial diameter (Arizona) —.. tescorum Chillcott

Tz median bristles equal to or shorter than tibial diameter 12

Basal abdominal segments 2-4 distinctly yellowish (Arizona) —

Pe Ae SLE SEN SCT Le TA eR ENN, oe) ho Fiche PTO, arizonensis Chillcott

Abdominal segment 2 and usually 3 yellowish, color often very indistinct

or tending to yellowish gray and confined to lateral areas; segment 4 gray

(Avizonay taliorntay “Jen. ee ee conspicua Malloch

LITERATURE CITED

Burnett, H. S., Parnepe, W. E., Lez, R. D. anp Wacner, E. D. 1957. Observa-

tions on the life cycle of Thelazia californiensis Price 1930. J. Parasitol.,

43 (4): 433.

VoL. 52, No. 3, JuLy 1976 241

‘CuiticoTt, J. G. 1960. A revision of the Nearctic species of Fanniinae (Diptera:

Muscidae). Canad. Entomol., Suppl. 14. 295 pp.

Mattocu, J. R. 1913. Notes on some American Diptera of the genus Fannia,

with descriptions of new species. Proc. U. S. Nat. Mus., 44: 621-631.

Weinmann, C. J., AnpERson, J. R., Ruprzorr, P., CONNOLLY, G. AND LONGHURST,

W. M. 1974. Eye worms and face flies in California. Calif. Agric.,

28(11): 4-5.

WINKLER, L. R. anp Wacner, E. D. 1961. A cultured life cycle of the canyon fly

Fannia benjamini Malloch with observations on the natural history.

Trans. Amer. Microscop. Soc., 80(2): 179-185.

ZOOLOGICAL NOMENCLATURE

The following Opinions recently published by the Commission may be found by

referring to the indicated volume of the Bulletin of Zoological Nomenclature.

1038 Argiope Audouin, 1826 and lobata, Aranea, Pallas, 1772, placed on Official

Lists (Arachnida, Aranes). Vol. 32(2) :105.

1039 Type-species of Uloma Dejean, 1821 (Tenebrio culinaris Linnaeus, 1758)

and of Phaleria Latreille, 1802 (Tenebrio cadaverinus Fabricius, 1792)

designated under the plenary powers; these names, with Alphitobius

Stephens, 1829 and Gnatocerus Thunberg, 1814, and the names of their

type-species, placed on Official Lists (Coleoptera). Vol. 32(3) :136.

1042 Deuterosminthurus Borner, 1901; Smynthurus bicinctus Koch, in Herrich-

Schaeffer, 1840 designated as type-species under the plenary powers (Col-

lembola). Vol. 32(4) :212.

1043 Eusminthurus Borner, 1900; Podura viridis Linnaeus, 1758 designated as

type-species under the plenary powers (Collembola). Vol. 32 (4) :214.

1049 Macgillivraya Grote, 1894 suppressed under the plenary powers; Friesea von

Dalla Torre, 1895 (type-species Triaena mirabilis Tullberg, 1871) placed

on Official List (Collembola). Vol. 32(4) :235.

1050 Pediculus eurysternus Burmeister, 1838 suppressed under the plenary

powers; Solenopotes Enderlein, 1904 (type-species S. capillatus Enderlein,

1904) and eurysternus, Haematopinus, Denny, 1842, placed on Official

Lists (Anoplura). Vol. 32(4) :238.

1051 Rhopalidia Lepeletier, 1836 suppressed under the plenary powers (Hyme-

noptera). Vol. 32(4) :240.

1053 Formica maxima Moore, 1842, suppressed under the plenary powers (Hyme-

noptera). Vol. 32(4) :244.

1054 Crinocerus Burmeister, 1835; Cimex sanctus Fabricius, 1775, designated

as type-species under the plenary powers (Hemiptera). Vol. 32(4) :246.

The Commission cannot supply separates of Opinions.—R. V. MELVILLE, Sec-

retary, International Commission on Zoological Nomenclature.

A New Species of Brine Fly From California Rice Fields

(Diptera: Ephydridae)

Wiis W. WirtH

Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv., USDA*

In my 1971 revision of the North American brine flies (Genus

Ephydra Fallén) I gave a number of California records of what I had

determined as the Palaearctic species, Ephydra macellaria Egger. More

recently (Wirth, 1975) I have revised the Old World species of Ephydra

and have more closely studied the macellaria complex. Ephydra macel-

laria Egger proves to be a seacoast species widely distributed in northern

and western Europe; whereas the Old World rice field species breeding

in habitats of fresher water as well as salt water is EF. helwanensis

Steyskal, described from Egypt and very common and widespread in

the Mediterranean area.

The California species is closely related and very similar to E.

macellaria and helwanensis, as redescribed by Wirth (1975), but with

distinct genitalic differences. My 1971 description and figures are

adequate to characterize the species, and I need merely at this time to

indicate how it may be distinguished from macellaria and helwanensis.

Ephydra usingeri, new species

Ephydra macellaria Egger; Wirth, 1971: 364 (misident.; description and figures;

California records) .

Resembles Ephydra macellaria Egger and helwanensis Steyskal in its heavily

pollinose body, only slightly shiny, with more of a grayish tinge on the sides;

legs yellowish with femora somewhat olivegreen infuscated; setae of body and

legs rather large; prescutellar area of female with a low hump bearing a group

of longer acrostichal setae. Aedeagus with apex slender and rounded in lateral

profile, the anterior membranous lobe with strong ridges only proximally, the

recurved basal process about half as long as straight portion. Surstyli stout

basally, tapering with rounded tips distally; without distinct sclerotized carina

or concavity, with fairly numerous scattered fine hairs except on dorsal side at

apex. Gonite with apex bearing 3 distinct lobes. Fifth tergum with short, stout,

anteroventral sclerotized process. Sternal plate strongly sclerotized and with

very strong transverse ridgelike folds, the whole angularly bent ventrally in middle,

forming a strongly serrate ventral process in genital pouch.

Differs from the two related species as follows: Male surstyli less triangular

in ventral profile and less pointed distally than in helwanensis, the tips rounded

but not constricted subapically as in macellaria. Gonite nearly as in helwanensis,

1 Mailing Address: c/o U.S. National Museum, Washington, D.C. 20560

Tue Pan-Pactric ENTOMOLOGIST 52: 242-243. JuLy 1976

VoL. 52, No. 3, Juty 1976 243

the rounded distal lobe not as prominent as in macellaria. Sternal plate with

large coarse transverse accordianlike pleats nearly as in macellaria, but usually

deeply folded into genital pouch, not forming as strong a process as in helwanensis.

Length, 4 mm.

Holotype, male, allotype, Tracy, SAN Joaquin Co., CatirorniA, 1 June 1951,

J. W. MacSwain (deposited in California Academy of Sciences, San Francisco) .

Paratypes, 21 males, 30 females, as follows: CALIFORNIA: Kings Co., Tulare

Lake, 17 March 1931, E. P. VanDuzee, 4 males, 3 females. San Joaquin Co.,

Tracy, same data as types, 12 males, 8 females. Stanislaus Co., Westly, 1 June

1949, H. H. Keiffer, collected from rice (California Dept. Agr.), 2 females.

Yolo Co., Davis, 30 April, 20-31 May 1960, F. E. Strong, 5 males, 17 females

(Davis) Paratypes in collections of USNM, California Academy of Sciences,

California Insect Survey in Berkeley, California Department of Agriculture in

Sacramento, and University of California at Davis.

The species is named for my teacher and good friend at the Uni-

versity of California in Berkeley, the late Dr. Robert L. Usinger, in

tribute to his leadership in the study of aquatic insects in North

America, and especially in his home state of California.

From the Stanislaus County collection from rice, and from the

nature of the California distribution records, it appears that E. usingeri

is a distinctly freshwater species that may be associated with rice

culture.

LITERATURE CITED

Wirto, W. W. 1971. The brine flies of the genus Ephydra in North America

(Diptera: Ephydridae). Ann. Entomol. Soc. Amer., 64: 357-377.

Wirt, W. W. 1975. A revision of the brine flies of the genus Ephydra of the

Old World (Diptera: Ephydridae). Entomol. Scand., 6: 11-44.

RECENT LITERATURE

FLOWERING PiLants. Evo.tuTion ABOVE THE Species LeveL. G. L. Stebbins.

Belknap Press, Harvard University, 1975. 399 pp., 13 tables, 59 figs. ($18.50,

hardcover).

THE SoctaAL BEHAVIOR OF THE Bees. C. D. Michener. Belknap Press, Harvard

University, 1974. 464 pp., figs. ($25.00, hardcover).

Additions to the Crane flies of California

(Diptera: Tipulidae)

CHARLES P. ALEXANDER

Amherst, Massachusetts

The volume entitled “The Crane Flies of California” by the present

author, was published in 1967 by the University of California Press.1

Somewhat more than 450 species were included, and during the inter-

vening period, numerous collections from the state have been studied

and additions to the list have been made. At this time I am describing

a few further species that were collected by Donald G. Denning, have

added the additional species that were made to the basic list during

this period, and have indicated two important changes in species

names. I wish to thank Doctor Denning for the many crane flies that

he has sent to me in the past that included several records of special

importance.

Rhabdomastix (Sacandaga) brevicellula, new species

General coloration of head and thorax light yellow, praescutum and scutum

patterned with light brown; wings with abundant trichia on vein R:; veins Ri+e

and Rs approximated at costa, the distance between them about ‘one-third vein

Rs; cell Ist Mz unusually short and broad, the second section of vein Mi,2 about

one-fifth longer than the cell width.

Male.—Length about 5.5 mm.; wing 6 mm. Rostrum and antennal scape clear

light yellow; palpi broken. Antennae with pedicel and flagellum black, proximal

flagellar segments subglobular to short-oval, outer segments progressively more

oval; outer segments with a single elongate verticil that is longer than the segment.

Head clear light yellow.

Pronotum clear light yellow, narrowly pale brown medially. Mesonotal praes-

cutum light yellow, with four light brown stripes, the intermediate pair nearly

confluent, ending some distance before the suture; posterior sclerites yellow, each

scutal lobe with two confluent light brown areas, posterior half of postnotal medio-

tergite similarly light brown. Pleura light yellow, ventral sternopleurite and meron

broadly light brown. Halteres yellow. Legs with coxae yellow, fore pair slightly

darker; trochanters yellow; remainder of legs broken. Wings light yellow, with-

out darkened pattern; veins very pale brown. Macrotrichia on R and outer longi-

tudinal veins, including about 20 over the whole length of R:; veins Rs, 2nd and

outer sections of Miy2, Ms and Mz with complete series of trichia; very sparse

scattered trichia on outer section of Cu: and outer ends of both Anal veins, Ist A

with a single trichia, 2nd A with two or three. Venation (Fig. 1) as shown; Sci

1 Alexander, C. P. The Crane Flies of California. Bulletin of the California Insect Survey. Volume

8, pp. 269; 524 figures, 1 plate, 106 maps. University of California Press, Berkeley; November 27,

1967.

Tue Pan-Paciric ENTOMOLOGIST 52: 244-250. Juny 1976

VoL. 52, No. 3, JuLty 1976 245

ending about opposite two-thirds to three-fourths Rs; distance on costa between

veins Ri,2 and Rs about one-third the latter vein; cell Jst Mz unusually short and

broad, as shown, the second section of M1,2 about one-fifth longer than the breadth

of the cell.

Abdomen, including hypopygium, chiefly yellow, faintly patterned with brown,

more evident on outer segments. Male hypopygium (Fig. 4) as shown.

Holotype male, CaLirorniA, Monterey County, Highway 128 at NortH Fork

of NarrAro River, July 6, 1975 (D. G. Denning).

The most similar regional species is Rhabdomastix (Sacandaga)

trichophora Alexander (California, Oregon, Washington) which simi-

larly has trichia on vein Ry of the wings, differing from the present fly

most evidently in venation, including the wide separation of veins

Ri, and Rs at costa and the longer cell Ist Mo.

Hesperoconopa anthracina, new species

General coloration of entire body black, certain areas more pruinose; halteres

brownish black, base of stem restrictedly yellowed; legs black; wings strongly

darkened throughout, stigma not differentiated; abdomen black, sternites slightly

more pruinose.

Female——Length about 5.3 mm.; wing 5.8 mm.; antenna about 1.1 mm. Ros-

trum and palpi black. Antennae black; flagellar segments subcylindrical, ends

truncated, subequal in length to the longest verticils. Head brownish gray; eyes

small; anterior vertex broad.

Thorax dull black, praescutum with vague still darker stripes, central area of

scutum and the scutellum slightly more pruinose. Pleura dark gray, ventral

sclerites slightly more darkened. Halteres brownish black, base of stem restrictedly

yellowed. Legs with coxae and trochanters black, slightly pruinose; remainder

of legs black. Wings (Fig. 2) strongly darkened throughout, stigma not dif-

ferentiated; narrow whitened lines over veins M, Mise, Msis and Ist A; veins

light brown, Sc, R, Mz and Cu darker. Outer wing cells from R; to Ms with scat-

tered trichia punctures, with further series in cells 1st A and 2nd A as indicated

in the figure. Venation as shown; Ress about one-half longer than Re; m-cu at

fork of M.

Abdomen blackened, sternites slightly more pruinose. Ovipositor with cerci

slender, outer ends slightly upcurved.

Holotype, female, CALiForNIA, ELpoRADO County, SoutH ForK oF AMERICAN

River, July 17, 1975 (D. G. Denning).

Generally similar to Hesperoconopa melanderi (Alexander), like-

wise from California, differing chiefly in the darker body coloration

and especially the more blackened wings. I believe that the male sex

when discovered will provide further hypopygial differences. The very

distinct aquatic immature stages of Hesperoconopa have been described

and figured by C. Dennis Hynes (Pan-Pacific Entomologist, 44: 324-

327, 4 figs.; 1968).

246 THE PAN-PACIFIC ENTOMOLOGIST

7

Fics. 1-7. Fig. 1. Rhabdomastix (Sacandaga) brevicellula sp.n.; venation. Fig.

2. Hesperoconopa anthracina sp.n.; venation. Fig. 3. Ormosia (Ormosia) den-

ningt sp.n.; venation. Fig. 4. Rhabdomastix (Sacandaga) brevicellula sp.n.;

male hypopygium. Fig. 5. Molophilus (Molophilus) spiniapicalis sp.n.; male

hypopygium. Fig. 6. Ormosia (Ormosia) denningi sp.n.; male hypopygium. Fig.

7. Ormosia (Ormosia) loretta sp.n.; male hypopygium.

Symbols: a, aedeagus; 0b, basistyle; bd, basal dististyle; d, dististyle; g, gona-

pophysis; i, interbase; id, inner dististyle; od, outer dististyle; p, phallosome;

t, tergite.

VoL. 52, No. 3, Juty 1976 247

Molophilus (Molophilus) spiniapicalis, new species

Allied to distilobatus; general coloration of thorax light brown, abdomen dark

brown; legs with femora yellow, tarsi brown; male hypopygium with mesal lobe

of basistyle long-extended, narrowed outwardly, the aggregation of blackened

spinoid setae opposite or beyond the apices of the dististyles, the subtending lobe

broad, with numerous setae.

Male.—Length about 44.3 mm.; wing 4.5-5.2 mm. Rostrum and palpi black.

Antennae (broken at fifth segment) with scape and pedicel obscure yellow, flagel-

lum black; flagellar segments oval, more elongate outwardly, verticils long. Head

blackened, gray pruinose.

Pronotum and pretergites light yellow. Mesonotal praescutum light brown,

humeral region vaguely more yellowed; scutum brown, scutellum obscure yellow;

postnotum dark brown to brownish black. Pleura yellowish brown to dark brown,

dorsopleural region still darker. Halteres with stem yellow, knob light brown,

apex more yellowed. Legs with coxae and trochanters yellow; femora yellow,

tibiae more obscure yellow, tarsi brown. Wings with prearcular and costal fields

light yellow, including the veins, remainder more whitish gray; veins brown, trichia

black.

Abdomen dark brown. Male hypopygium (Fig. 5) generally as in distilobatus,

differing in details, especially of the basistyle. Mesal lobe of basistyle, 6, long-

extended and narrowed, the aggregation of blackened spinoid setae lying opposite

or beyond the apices of the dististyles, the subtending lobe broad, with numerous

setae. Aedegus, a, long and slender, extended into a long spine.

Holotype male, CALIFORNIA, MONTEREY COUNTY, PLASKETT CREEK CAMPGROUND,

July 25, 1975 (D. G. Denning). Paratopotype, ¢. Paratype, ¢, California, with-

out further data, August 22, 1966 (C. Dennis Hynes).

The most similar species is Molophilus (Molophilus) distilobatus

Alexander (Washington to California). The distinctive male hypopy-

sium of this is discussed and figured in the Crane Flies of California,

p. 148, fig. 514.

Ormosia (Ormosia) denningi, new species

General coloration of head and thorax dark gray, mesonotal praescutum with

a narrow dark brown median line; wings light brown, stigma darker brown; abdo-

men brownish black; male hypopygium with tergite large, narrowed outwardly

to the shallowly concave apex; inner dististyle black, outwardly divided into two

strong spines, the outer one slightly broader, the margin microscopically rough-

ened; basistyle with interbase a simple long slender blackened spine.

Male.—Length about 6 mm.; wing 5.5 mm. Rostrum and palpi black. Anten-

nae (male) broken at fourth segment, scape and pedicel light brown, flagellum

black; proximal two flagellar segments elongate, broadest basally, narrowed to

the outer end, the antenna when entire evidently relatively long. Head brown-

ish gray. ‘

Thorax almost uniformly dark gray, mesonotal praescutum with a narrow dark

brown central line, behind almost attaining the suture. Halteres light brown,

outer half of knob dark brown. Legs with coxae obscure yellow, trochanters some-

what clearer yellow; remainder of legs broken. Wings (Fig. 3) light brown,

248 THE PaAn-PAcIFIC ENTOMOLOGIST

stigma conspicuously darker brown, wing cells before the stigma slightly more

yellowed; veins dark brown. Venation almost as in burneyana; left wing of type

mounted on slide, with a long subterminal spur near outer end of vein R., not

reaching the margin, the right wing normal.

Abdomen brownish black, sparsely pruinose. Male hypopygium (Fig. 6) with

the tergite, t, large, broadest across base, gradually narrowed outwardly to the

shallowly concave apex, outer lateral angles produced into small pale oval lobes that

are provided with abundant microscopic setae. Outer dististyle, od, simple, broad-

est at near midlength, narrowed outwardly into an obtuse point; inner style, id,

distinctive, black, widened outwardly, divided into two strong spines, the outer

one broader, microscopically roughened. Basistyle, 6, with interbase, i, a simple

long slender blackened spine, the base dilated.

Holotype male, Catirornia, MonterEY County, PLasKeTt CREEK CAMPGROUND,

July 25, 1975 (D. G. Denning).

I take pleasure in naming this interesting fly for Dr. Donald G. Den-

ning, distinguished student of the Trichoptera, to whom I am indebted

for many Tipulidae from the western United States. The nearest rela-

tive is Ormosia (Ormosia) burneyana Alexander, likewise from Calli-

fornia, which differs chiefly in hypopygial structure, including the

tergite, basistyle and inner dististyle. The male hypopygium of this

species was figured in the California Bulletin, p. 144, fig. 481. The

structure of the inner dististyle also is generally similar to that of the

unrelated Molophilus (Molophilus) fenderi Alexander, also of western

North America.

Ormosia (Ormosia) loretta, new species

Antennae of male elongate; body dark brownish gray, abdomen dark brown;

male hypopygium with the inner dististyle blackened, terminating in a major

spine; basistyle with the interbase distinctive, with two long blackened spines;

phallosome with gonapophysis bispinous.

Male.—Length about 5.5 mm.; wing 6 mm. Rostrum and palpi black. Anten-

nae broken beyond the fourth segment, elongate, if entire apparently extending

to the wing root or slightly beyond, black; basal flagellar segments elongate, with

very long erect setae, the longest nearly equalling the segment. Head dark gray.

Prothorax dark brownish gray. Mesonotal praescutum chiefly covered by four

grayish brown stripes, interspaces scarcely indicated, each with a row of micro-

scopic setigerous punctures; pseudosutural foveae and tuberculate pits black,

conspicuous; posterior sclerites of notum dark brown. Pleura brownish gray, dor-

sopleural membrane light yellow. Halteres with stem obscure yellow, knob brown-

ish black. Legs with coxae obscure yellow, bases of middle pair narrowly darkened;

trochanters light yellow; remainder of legs broken. Wings light brown, cells basad

of the dark brown stigma light yellow, beyond the stigma less evidently brightened;

cells C and Sc slightly darker than the ground; veins dark brown. Venation: Sci

ending shortly beyond level of Re; Rs straight, slightly longer than vein Sci; cell

Mz open by atrophy of Mz; m-cu shortly before fork of M; vein 2nd A with distal

third slightly arcuated.

VoL. 52, No. 3, Juty 1976 249

Abdomen dark brown. Male hypopygium (Fig. 7) with the important structures

as shown. Inner dististyle, id, blackened, expanded outwardly, terminating in a

long triangular point, with two smaller more dorsal spines. Basistyle, b, with a

modified interbasal structure, 7, as shown, terminating in an elongate spine with

two microscopic spinules on outer margin, on inner face with a comparable long

slender spine. Phallosome, p, including the small slender aedeagus and the

gonapophyses, g, that appear as a straight yellow spine that bears a long black

lateral rod at near midlength.

Holotype, ¢, California, Santa Cruz County, Watsonville, on Highway 152,

August 1, 1975 (D. G. Denning).

I take pleasure in naming this species for Mrs. Donald Denning,

Loretta. There are several allied and generally similar species in west-

ern North America that have been discussed and figured in the Califor-

nia Bulletin, such including Ormosia (Ormosia) burneyana Alexander,

O. (O.) burneyensis Alexander, O. (O.) denningi sp.n., O. (O.) legata

Alexander, O. (O.) pleuracantha Alexander, and some others.

SPECIES NEW TO CALIFORNIA LIST

Tipula (Trichotipula) frommeri Alexander. Great Basin Naturalist, 33: 189-

192; 1973. California, Riverside County, Deep Canyon, Desert Research Center,

May 15, 1969, Saul Frommer.

Tipula (Trichotipula) sanctaecruzae Alexander. Great Basin Naturalist, 33:

192-193; 1973. California, Santa Cruz Island, Channel Islands, 1941 (Eric Rem-

ington).

Tipula (Lunatipula) hastingsae diperona Alexander. Great Basin Naturalist, 33:

193; 1973. California, Santa Cruz Island, Channel Islands, University of Califor-

nia Research Station, Canada del Medio, April 1970 (Eric Remington).

Tipula (Lunatipula) mecotrichia Alexander. Great Basin Naturalist, 26: 2-3;

1966. California, Fresno County, Kings Canyon, June 1, 1963 (Alexander).

Dicranota (Dicranota) bernardinensis Alexander. Great Basin Naturalist, 24:

4; 1966, California, San Bernardino County, Thurman Flats, Mill River, May 1,

1963 (Alexander).

Dicranota (Rhaphidolabis) nooksackensis Alexander. American Midland Natu-

ralist, 42: 300-301; 1949. Washington (type); Oregon, Alaska. California, Eldo-

rado County, South Fork of American River, July 17, 1975 (Denning).

Dactylolabis (Dactylolabis) postiana Alexander. Pan-Pacific Entomologist, 20:

94-95; 1944. Oregon (types). California, Shasta County, March 31, 1968

(Denning).

Dactylolabis (Eudactylolabis) vestigipennis Alexander. Bull. Brooklyn Ent. Soc.,

45: 45-46; 1950. Types from the Tucson Mountains, Pima County, Arizona, Feb-

ruary and March 1937 (Owen Bryant). California, Los Angeles Co., February 10,

1967 (David Rentz) ; San Bernardino County, near Cajon Pass, at edge of Mohave

Desert, March 1967 (Rentz). For a more detailed account, see Rentz and Gagné,

Ent. News, 78: 261-262; 1967.

Idiognophomyia enniki Alexander. Pan-Pacific Entomologist, 50: 279-281, 3

figs.; 1974 (adult). George W. Byers, the same, 50: 282-287, figs. 1-15 (larva

and pupa); 1974. Types from Ventucopa, western Ventura County, Los Padres

250 THE PAN-PACIFIC ENTOMOLOGIST

National Forest. Larvae and pupae were found by Dr. Franklin Ennik in decaying

Yucca near U. S. Highway 399 on April 9, 1974.

CHANGES IN NAMES

(Trichoceridae)

Diazosma hirtipennis (Siebke) (1863)

Diazosma subsinuata Alexander; North American Catalogue, p. 15; 1965.

Crane Flies of California, p. 13; 1967.

Diazosma hirtipennis Siebke; Christine Dahl, Opuscula Entomologica, 31:

97; 1966.

(Tipulidae)

Ormosia (Ormosia) affinis (Lundbeck)

Rhypholophus affinis Lundbeck; Diptera Groenlandica. Vidensk. Medd. nat.

hist. Copenhagen, p. 266, plate VI, fig. 17 (wing) ; 1898.

Ormosia manicata (Doane), as Rhypholophus; Jour. N. Y. Ent. Soc., 8: 187;

September 1900. See Bo Tjeder, 1970 Entomologiske Meddelelser, 38: 253-

256, 4 figs. (male hypopygium) for detailed synonymy. Synonymy of manicata

given by Alexander in California Bulletin, p. 145.

Changes in generic and subgeneric names.

Limnophila (Euphylidorea) Alexander, Ent. News, 83: 32; 1972.

Four species of Limnophila (Phylidorea) in Craneflies of California, p. 112,

aequiatra Alexander, brevifilosa Alexander, burdicki Alexander, and flavipila

Doane are transferred to the new subgenus Euphylidorea, cited above.

Erioptera (Hespererioptera) Alexander, Ent. News, 35-36; 1972, new subgenus

for Erioptera oregonensis Alexander, California Bulletin, p. 135.

RECENT LITERATURE

REVISION DE LA TRIBU DES RHODOPININI GRESS. DE LA REGION AsIATO-AUSTRALIENNE

(CoLEOPTERA, CERAMBYCIDAE), PREMIERE PARTIE. S. Breuning. 70 pages plus

7 photographs on 2 plates, 21 cm x 30 cm page, spiral bound. “Imprimé en

Xerocopie” by Sciences Nat, 45 rue des Alouettes, 75019 Paris. “Copyright:

SCIENCES NAT 1975.” Price about US $10.

This publication provides a key and brief descriptions to the 75 genera (4 are

new, with also 3 new subgenera) for the area considered. Keys are provided

to the species if there is more than one included species in a genus. 231 species

are briefly described, of which 12 are new. The 7 photographs mounted on 2

pages include illustrations of 3 Pic types—Paut H. ArNaup, Jr.

Life History of the Ragweed Plume Moth, Adaina

ambrosiae (Murtfeldt), in Southern California

(Lepidoptera: Pterophoridae)

R. D. GorpEN anp D. W. RICKER

Department of Etomology, University of California,

Riverside, California 92502

Faunistic surveys of California ragweeds (Ambrosia spp., Compos-

itae) detected numerous species of phytophagous insects associated

with these native plants (Goeden and Ricker, 1974a, 1974b, 1975, 1976,

and unpublished data). Many of these insects were little studied spe-

cies of no economic importance. We have begun life history studies of

the more stenophagous species, some for their intrinsic interest, others

as candidate agents for the biological control of North American rag-

weeds accidentally introduced into eastern Europe and elsewhere, where

they are becoming important agricultural and hayfever-inducing weeds

(Goeden et al., 1974).

Adaina ambrosiae (Murtfeldt) was among the first species selected

for more intensive study in the field and laboratory; however, it was

found to reproduce readily on sunflower, Helianthus annuus L., and

thus was rejected as a candidate biological control agent. Our observa-

tions on the life history of this heretofore little known plume moth are

reported. Unless otherwise indicated, these data were obtained from

cultures maintained in the insectary of the Division of Biological Con-

trol, Department of Entomology, University of California, Riverside.

Insectary conditions were 27 +1°C, 40-70% relative humidity, and

a 12/12-hr (light/dark) photoperiod.

Taxonomy.—Adaina ambrosiae was described as a species of Oida-

matophorus by Murtfeldt (1880). She included descriptions of the

larva and pupa, which were quoted and amplified by Barnes and Lind-

sey (1921). The adult was illustrated in Barnes and Lindsey (1921)

and Essig (1926).

Distribution and Host Planis.—Essig (1926, p. 714) described its

range as “. . . throughout the United States, and in Arizona and along

the Pacific Coast in the west.” He noted that the larvae feed on “rag-

weed.” Barnes and Lindsey (1921) reported this species from Am-

brosia artemisitfolia L.

We collected Adaina ambrosiae larvae during January—October, 1969-

71, from Ambrosia acanthicarpa Hooker, A. chamissonis (Lessing)

Greene, A. confertiflora Decandolle, A. dumosa (Gray) Payne, A. erio-

Tue PAn-Pactric ENToMoOLOoGIsT 52: 251-255. Juty 1976

252 THE Pan-PaciFiIc ENTOMOLOGIST

mas —

shed

Fic. 1. Life stages of Adaina ambrosiae. (a) egg, 50, (b) fifth instar larva

that has fed on leaf of Ambrosia confertiflora, 2, (c) pupa, 7X, (d) adult, 4x.

centra (Gray) Payne, and A. psilostachya Decandolle at the following

locations in southern California: Los Angeles Co.—Azusa, Llano,

South San Gabriel; Orange Co.—Huntington Beach, El Toro, Yorba

Linda; Riverside Co.—Banning, Desert Center; San Bernardino Co.—

Apple Valley, Bloomington, Cedar Canyon, Hackberry Mountain, Moun-

tain Pass, Twenty-nine Palms; San Diego Co.—Chula Vista, Fallbrook,

La Mesa, Piru, and Ventura. Adaina ambrosiae was most commonly and

widely associated with western ragweed, Ambrosia psilostachya, on

which it was cultured in the insectary. We also have collected the larvae

from cocklebur, Xanthium strumarium L., on several occasions and

reared them to adults on this weed.

Biology.—Egg.—The egg (Fig. la) is ellipsoidal, translucent white

to pale yellow, smooth, and lustrous. The mean (+S.E.) length and

width of 50 eggs were 0.354 + 0.002 mm and 0.237 + 0.001 mm, respec-

tively.

The positions of 220 eggs obtained from 5 moths were recorded. All

were attached singly to leaf blades, one to several per leaf, their long

axes parallel to the upper and lower leaf surfaces (Fig. la). Only 4

(1.8%) of these eggs were laid on the superior epidermis. Fifty-seven

VoL. 52, No. 3, Juty 1976 253

(26%), 121 (56%), and 38 (18%) of the remaining 216 eggs were

attached to the basal, middle, and apical thirds of the undersides of

leaf laminae, respectively. Of the 220 eggs, 108 (49%) were laid along-

side a midrib, 60 (27%) were laid in contact with a primary vein, 25

(11%) were laid next to a secondary vein, and the remaining 27 (12%)

touched no veins. The incubation period was about 4 days.

Larva.—Twelve of 15 larvae individually reared from eggs passed

through 5 instars in an average of 18+0.8 days. Three larvae ap-

parently molted only 3 times and pupated after 14, 15, and 15 days.

The maximum lengths attained by the first to fifth instars were 1.15 +

0.03 mm, 1.90 + 0.03 mm, 3.06 + 0.07 mm, 4.94 + 0.03 mm, and 7.44

+ 0.07 mm, respectively. Larvae with 5 instars grew in length an aver-

age of 60% between each successive molt.

The newly hatched larva is pale yellow. After eclosion, it abandoned

the chorion and immediately proceeded to feed on the leaf lamina,

gradually assuming the pale green coloration characteristic of sub-

sequent instars. All instars were leaf skeletonizers (Fig. lb). They

consumed the adjacent epidermis and mesophyll, forming irregularly

shaped scars, but left intact the opposite epidermis, midrib, primary

veins, and at least the bases of the secondary veins. First to third

instars evidenced a slight preference for feeding upon the lower leaf

surfaces. Fourth and fifth instars tended to feed more upon the upper

leaf surfaces. The latter tendency was most pronounced in the last instar

(Fig. 1b).

The larvae apparently fed most actively at night. In the field, later

instars commonly rested by day in shallow grooves excavated the

lengths of their bodies along the midribs on the upper sides of leaves.

From these resting niches, in which pupation frequently occurred, the

larvae apparently foraged by night to return and rest between feedings

by day. Fine silken threads on the foliage marked the course of their

movements. In the insectary, one or more stadia, sometimes the entire

larval period, was spent on the same leaf. The first to fifth stadia

averaged 3 + 0.2 (range: 2-4.) days, 2.6 + 0.1 (range: 2-3), 2.30.1

(range: 2-3), 3.5 40.2 (range: 3-5), and 5.6 +0.2 (range: 5-7)

days, respectively.

The later instars threw their small, dark, fecal pellets over their heads

for distances of several cm by means of rapid upward flips of their

terminal abdominal segments. This behavior kept their feeding areas

relatively clean (Fig. 1b).

A non-feeding prepupal stage began 2 days prior to pupation, during

which time the mature larva contracted slightly in length, took on a

254, THE PAN-PAcIFIC ENTOMOLOGIST

violet hue, and spun a small silken pad upon the leaf surface. The pre-

pupae turned whitish 1 day prior to pupation.

Pupa.—tThe pupa (Fig. lc) is slightly angulate, hairy, and naked.

It is attached posteriorly by a cremaster to the silken pad spun by the

mature larva. When disturbed, the pupa bent its abdomen backward

and arched upward anteriorly. The mean length of 118 pupae was

5.73 = 0.03 mm.

Most pupation occurred atop leaf blades, although a few pupae were

attached to the undersides of living or dead, lower cauline leaves.

Some mature larvae vacated the plants and pupated on the walls and

floors of cages. The pupal periods for 55 individuals averaged 5.5 = 0.1

(range: 2-8) days.

Adult.—Adults were not commonly encountered in nature. They

mostly remained at rest on plants by day, their wings characteristically

outstretched (Fig. 1d), flying short distances when disturbed.

When caged with bouquets of freshly excised, vegetative shoots and

provided with water, 11 fecund females lived an average of 11.8 + 1.1

(range: 7-17) days; 15 males averaged 10.7 + 1.1 (range: 6-22) days.

After a preovipositional period that averaged 2.8 + 0.3 (range: 2-5)

days, these 11 females oviposited for an average of 8.54 1.1 (range:

4—15) days. During these ovipositional periods, they laid an average

total of 119 + 26 (range: 38-283) eggs. The daily numbers of eggs

produced by these females varied considerably. Based only on days

when females oviposited, the daily totals varied from 1 to 67 eggs and

averaged 14.4, + 1.4 eggs. Oviposition rates sharply declined toward the

ends of the oviposition periods. Oviposition apparently occurred only

at night.

Mating and dispersal normally were nocturnal activities, although

4. instances of mating were observed during daylight hours in the morn-

ing and afternoon for protracted, but untimed durations.

Of 109 adults reared from a random sample of insectary-reared F,

generation pupae derived from field-collected larvae, 52 were females

and 57 males.

Seasonal History.—Adaina ambrosiae is multivoltine in southern Cal-

ifornia; however, the number of annual generations is unknown and

probably varies considerably with the host plant species and climatic

zone involved. In the insectary, the egg to egg cycle was completed in

about 30 days.

Natural Enemies.—Field collected larvae commonly were heavily par-

asitized. Apanteles sp. nr. nemoriae Ashmead (Hymenoptera: Bra-

conidae) was reared from larvae collected from Ambrosia confertiflora

VoL. 52, No. 3, JuLy 1976 250

and A. psilostachya. Hyposoter sp. nr. nigrolineatus (Viereck) (Hymen-

optera: Ichneumonidae) was reared from larvae collected on A. con-

fertiflora, A. dumosa, A. eriocentra, and A. psilostachya. Both were

primary, solitary endoparasitoids. A single specimen of Mesochoris sp.

(Hymenoptera: Ichneumonidae) was reared, probably as a _hyper-

parasitoid, from a larva collected on Ambrosia psilostachya.

ACKNOWLEDGMENTS

Adaina ambrosiae was identified by Dr. D. C. Ferguson, Systematic

Entomology Laboratory, U. S. Dept. Agric., Agric. Res. Serv., c/o

U. S. National Museum, Washington, D. C. The parasitoids were

identified by Dr. P. M. Marsh, same address.

LITERATURE CITED

Barnes, W., AnD Linpsey, A. W. 1921. The Pterophoridae of America, north

of Mexico. p. 281-477. In Contributions to the Natural History of the

Lepidoptera of North America. The Review Press, Decatur, Illinois.

Essic, E. O. 1926. Insects of Western North America. The Macmillan Co., New

York. 1035 p.

GoebEN, R. D., anp Ricker, D. W. 1974a. The phytophagous insect fauna of the

ragweed, Ambrosia acanthicarpa, in southern California. Environ.

Entomol. 3: 827-834.

GorpEN, R. D., and Ricker, D. W. 1974b. The phytophagous insect fauna of

the ragweed, Ambrosia chamissonis, in southern California. Environ.

Entomol. 3: 835-839.

GorDEN, R. D., anp Ricker, D. W. 1975. The phytophagous insect fauna of the

ragweed, Ambrosia confertiflora, in southern California. Environ.

Entomol. 4: 301-306.

GorpEN, R. D., anp Ricker, D. W. 1976. The phytophagous insect fauna of the

ragweed, Ambrosia dumosa, in southern California. Environ. Entomol.

5: 45-50.

GorEpEN, R. D., Kovatev, O. V., AnD Ricker, D. W. 1974. Arthropods exported

from California to the U.S.S.R. for ragweed control. Weed Sc. 22:

156-158.

MourtreLtpt, M. E. 1880. New plume moths (Pterophoridae). Amer. Entomol.

3: 235-236.

A New Choreutis from the San Francisco Bay Area,

California

(Lepidoptera: Glyphipterygidae) *

Joun B. HEPPNER?

Department of Entomology and Nematology

University of Florida, Gainesville, Florida 32611

Current revisionary work in the Glyphipterygidae*®, of the Palearctic

fauna by A. Diakonoff (Rijksmuseum van Natuurlijke Historie, Leiden,

Netherlands), of the Japanese fauna by Y. Arita (Meijo University,

Nagoya, Japan), and of the Nearctic fauna by myself, has uncovered

numerous taxonomic and nomenclatural problems at the species and

generic level. Brock (1967) has indicated that the family may be poly-

phyletic, which may require the elevation of the two included sub-

families (Choreutinae and Glyphipteryginae) to family rank, but this

requires further investigation in a world-wide generic revision.

The Holarctic genus Choreutis Hubner is heterogeneous as presently

constituted in North America, necessitating some transfers to the long

unused but distinct Tebenna Billberg. Various characters common to

several western North American species currently in Choreutis, includ-

ing the new species described below, also indicate that this species

sroup may require a new genus. Species of Choreutis and Tebenna have

been almost exclusively placed in the former genus or in Simaethis

Leach. Simaethis is a junior synonym of Anthophila Haworth.

The new species described below, tentatively placed in Choreutis, was

among material sent on loan from the California Insect Survey of the

University of California, Berkeley. Biological studies by J. A. Powell

(University of California, Berkeley) warrant a description of the spe-

cies at this time. The specific name of the new species is derived from

Greek translated as “from hound’s tongue.”

Choreutis apocynoglossa, new species

Male (Fig. 1).—Forewing length 4.8 to 6.0 mm. Head: vertex with dense white-

tipped, fuscous scales directed antero-mesad; frons similar in color. Labial palpus

somewhat upturned, extended anteriorly nearly twice eye diameter; basal segment

1 Florida Agricultural Experiment Station Journal Series No. 7013.

2 Research Associate, Florida State Collection of Arthropods.

3 The International Commission on Zoological Nomenclature is considering a proposal by A. Diako-

noff and myself for plenary power retention of the name Glyphipterygidae, rather than the spell-

ing correction to Glyphipterigidae, and curtailment of the further use of Glyphipteryx Curtis for

Chrysoclista in Blastodacnidae (case Z. N. (S.) 2115).

Tue Pan-Paciric ENTOMOLOGIST 52: 256-262. JuLy 1976

VoL. 52, No. 3, Juty 1976 a7

Fics. 1-2. adults of Choreutis apocynoglossa Heppner: Fig. 1. holotype male;

Fig. 2. allotype female.

white, tufted ventrally; 2nd segment subequal to Ist, fuscous with apical two

scale rows white-tipped, tufted ventrally with fuscous and white scales as long as

2nd segment; apical segment half length of 2nd, white and fuscous. Antenna

somewhat more than half forewing length; scape fuscous, remaining segments

with alternating bands of dark fuscous and white; ventrally with setae (male sex

character) twice length of antennal width. Thorax: dorsum white and fuscous

intermixed. Patagia white ringed, fuscous centrally. Venter white. Legs white

with some fuscous on fore- and midtibiae; tarsal segments basally fuscous, apically

white. Spurs white, inner spurs near twice outer spur lengths. Forewing: length

2.5 greatest width; apex and tornus rounded; termen only slightly convex, oblique

to tornus; costal margin and posterior margin convex; chorda weakly separated

from cell; pterostigma well developed. Ground color dark fuscous, scales white-

tipped. Basal fourth fuscous, some white-tipped scales, with a small patch of

ochreous-tipped scales near base of radius. Antemedial fascia convex, distinctly

dull white, scales basally dark fuscous. Wide median fuscous band; white spot on

costal margin, followed diagonally toward tornus by silvered line ending at cubital

border of cell, then interrupted before continuing to posterior margin, angled

toward anal angle; upper silvered line surrounded by ochreous-tipped scales; two

diffuse patches of black-fuscous scales near median on radius and anal vein. Post-

medially of ground color; distal third with four diffuse, white longitudinal striae

above black tornal mark (striae sometimes more or less merged) ;- costal margin

with white spot just beyond median and another at apical fourth of wing; metallic-

silver scales on radius at median and near apical corner of cell; apical angle with

fuscous curved line followed by metallic-silver curved line; distal linear yellow

spot at apex. Tornal quarter with large black trapezoidal spot, interrupted dis-

tally beyond midpoint by vertical light yellow line, with metallic-silver spot in

anterior half; a distal border of metallic-silver. Tornal angle fuscous, scales white-

tipped, some yellow-tipped scales adjacent to metallic-silver border of large black

spot. Fringe with basal row of fuscous, white-tipped scales, then fuscous to diffuse

white outer margin of termen; longer and oblique at tornus. All metallic-silver

spots elevated above wing surface due to convex scales. Ventrum with costal white

258 THE Pan-PActFIc ENTOMOLOGIST

xc We } WAVY

Ab WS cA

TZ v

Fic. 3. male genitalia of Choreutis apocynoglossa Heppner holotype, ventral

aspect; a, ventro-lateral aspect of aedeagus.

spots repeated; costal half of wing evenly fuscous; posterior half lustrous dull

white to irregular border area with costal half; apex fuscous, extended as a fus-

cous terminal border to tornus. Hindwing: triangular, tornal margin broadly

rounded; costal margin convex; ground color uniformly lustrous white except for

some scattered fuscous scales at base (rarely in central white area) ; fuscous mar-

gin at termen; margin broader at apical angle, becoming narrow toward tornus.

Fringe white, slightly brown and basally fuscous only at apical margin. Ventrum

white; some fuscous at apical angle. Abdomen: fuscous and white ringed dorsally

and ventrally, more white laterally, each segment anteriorly fuscous and posteriorly

white. Genitalia: as in Fig. 3. Tegumen convex posteriorly, becoming a narrow

band laterally. Socii large, well developed. Gnathos absent except for apparently

vestigial and plate between socii. Juxta basally elongated to juncture with

valval sacculus; lateral edges becoming thin. Valva dorso-laterally convex. Vin-

culum broadly convex. Aedeagus with 34 strongly sclerotized, flat, sharply

pointed cornuti almost as long as phallobase length. Genitalia drawn from holotype

(JBH prep. No. 255) ; 9 preparations examined.

Female (Fig. 2).—Forewing length 5.8 to 6.5 mm. As described for male, but

with antenna lacking ventral setae; hindwing terminal fuscous border sometimes

wider than in male; ventrum of forewing with less fuscous on termen than in male.

Genitalia (Fig. 4): ovipositor lobes small, sharply edged, with many setae on

sclerite anterior to lobes. Posterior apophyses longer than anterior apophyses;

basally broad and flattened. Anterior apophyses with a broad flattened central

area with a slight hook-projection ventro-laterally. Ostium round with broadened

antero-ventral margin, bordered anteriorly by extensive setal field. Ductus bursae

sclerotized, nearly as wide as ostium, becoming two times wider at enlarged, dis-

VoL. 52, No. 3, JuLy 1976 259

tinct offset juncture with bursal ventrum. Junction of ductus bursae and ductus

seminalis at one-third ductus bursae length from ostium; ductus seminalis with

oval bulla seminalis. Bursa copulatrix large, heart-shaped, with somewhat elongate-

pointed anterior end. Signum a linear band of spicules merging to smaller spicules

laterally, as a line between heart-shaped lobes, from ductus bursae junction to two-

thirds distance to anterior end dorsally. Bursa with minute spicules over most of

surface. Dorsal margin of ductus bursae-bursa juncture with patch of large

spicules. Genitalia drawn from allotype (JBH pre. No. 256); 7 preparations

examined.

Types.—Holotype male and allotype female: PAtrrerson Reserve, DEL VALLE

Lake [Rocky Ruipceg, 8 air mi. SSE. Livermore], ALAMEDA County, CALIFORNIA,

3 Feb 1974, rearing 74B1, emerged ex Cynoglossum grande 23 Feb 1974 (J. A.

Powell). Holotype and allotype with the University of California, Berkeley,

deposited on indefinite loan at the California Academy of Sciences.

Specimens examined.—45 paratypes from California: Marin County: Alpine

Lake, 25 Apr 1958 (1 9); Lagunitas, 17 Mar 1970 (1 @) (P. A. Opler) .—San

Mateo County: San Bruno Mts., 20 Mar 1970 (2 4) (Opler).—Contra Costa

County: Tilden Park, Berkeley Hills, 16 Feb 1970, rearing 70B6, emerged ex

Cynoglossum grande 28 Feb 1970 to 7 Mar 1970 (4 6, 492) Cynoglossum associ-

ation, 16 Apr to 20 Apr 1969 (6 ¢,13 2, Opler) Merry Go Round Hill, Berkeley

Hills, 20 Feb 1973, rearing 73B2, emerged ex Cynoglossum grande 10-15 Mar 1973

(3 6,29); MGR Hill-Wildcat Creek, Berkeley Hills, 7 Mar 1972, rearing 72C3,

emerged ex Cynoglossum occidentale 22-29 Mar 1972 (5 92). Alameda County:

Patterson Reserve, Del Valle Lake, 3 Feb 1974, rearing 74B1, emerged ex Cyno-

glossum grande 23 Feb 1974 (1 6,19), 24 Feb 1974 (1 2); 17 Feb 1973, rear-

ing 73B1, emerged ex Cynoglossum grande 6 Mar 1973 (1 @).

Additional specimens from California not designated as paratypes: “Placer

Co.”, 20 Apr (4 6,3 9—USNM) (A. H. Vachell); 1 May (1 6—MCZ; 1 9—

USNM) (Vachell) ; Cisco, Placer Co., 1 Jun 1905 (1 9—USNM) (Vachell).

All specimens were collected or reared by J. A. Powell unless indicated other-

wise. Paratypes are deposited in the following collections: University of California,

Berkeley, Canadian National Collection, Florida State Collection of Arthropods,

U.S. National Museum of Natural History, and my own collection.

Specimens from the San Francisco Bay area show little variation,

except that freshly emerged specimens have darker fuscous markings

than flown specimens. There is one anomalous female from the type

locality with the 2nd anal vein fuscous and with a spot at the discal end

of the cell. This female approaches the more extensive fuscous hind-

wing scaling of some of the specimens (¢ and 2) of the Sierra Nevada

foothill population (Placer County). These latter specimens include

some that have white hindwings as in the coastal population, others that

have most of the hindwings fuscous, while some are intermediate. The

dark-hindwing members of this Placer County population greatly resem-

ble Choreutis occidentella Dyar, which is most closely related to C. apoc-

ynoglossa, but they may be distinguished by always having at least the

anal margin to the 3rd anal vein of the hindwings white. C. occidentella

260 THE PAn-PaciFic ENTOMOLOGIST

Fic. 4. female genitalia of Choreutis apocynoglossa Heppner allotype, ventral

aspect; a, dorsal schematic of bursa copulatrix inflated (reduced scale).

VoL. 52, No. 3, Juty 1976 261

has the hindwings uniformly gray-brown with no white scales. C. occi-

dentella also has a consistantly longer and thinner aedeagus, and a val-

val sacculus that is more produced at its distal end than in C. apocyno-

glossa.

Choreutis occideniella occurs throughout montane areas of the Rocky

Mountains, from northern New Mexico to northern British Columbia,

west into northern California and the Sierra Nevada. A series of melanic

Colorado specimens, defined as Choreutis coloradella Kearfott, probably

represents a high altitude form of C. occidenitella, although a final clari-

fication of the status of C. coloradella requires further investigation.

The only known zone of possible contact between C. apocynoglossa and

C. occidentella is in the Placer County area of the Sierra Nevada, al-

though another likely contact zone would be in northern Lake County,

which is adjacent to montane areas of Glenn County, the southern known

limits of C. occidentella in northern coastal California. The darker

hindwings of some members of the Placer County population indicates

possible hybridization with C. occidentella. Since the exact status of

the Placer County population of C. apocynoglossa is uncertain, the

available specimens are not added to the paratype series.

Choreutis caliginosa Braun from Montana is superficially similar to

C. apocynoglossa but may be easily distinguished by the very different

genitalia. The Mexican Choreutis schausiella Busck, from Las Vigas,

Mexico, but erroneously stated to be from New Mexico (Busck, 1906),

is likewise similar to C. apocynoglossa, but the genitalia are very dif-

ferent and the adults are about half the size of C. apocynoglossa.

Flight period.—Mid-March to late April (San Francisco Bay area) ;

late April to early June (Sierra Nevada foothills).

Biology.—Hosts: Cynoglossum grande Douglas and Cynoglossum

occidentale A. Gray (Boraginaceae). The only other Choreutis known

to have a host plant in the Boraginaceae is Choreutis augustella Clarke

of the Pacific Northwest (Braun, 1940). Most other Chorewtis species,

where known, utilize Labiatae and Urticaceae, but the hosts remain un-

known for the majority of the species. Most known Tebenna hosts are

in Compositae. The hosts of species closely related to C. apocyno-

glossa are not known. Characteristic of Choreutis and related genera,

C. apocynoglossa larvae pupate in an elliptical, flattened cocoon of silk

formed in two layers. Pupal shells are extruded with adult eclosion.

Immature stages will be described in my revision of the family. Col-

lection sites in the San Francisco Bay area are typical Coast Range oak

woodland habitats where Cynoglossum often is a common understory

plant. While Cynoglossum occurs in open surroundings as well as

262 THE PAn-PAcIFIC ENTOMOLOGIST

among dense oak stands, larvae of Choreutis apocynoglossa have thus

far been found only on plants growing in deep shade (Powell, personal

communication).

ACKNOWLEDGMENTS

For their helpful comments I wish to thank D. H. Habeck and J. Reis-

kind, University of Florida, and H. V. Weems, Jr., Florida State Collec-

tion of Arthropods, Florida Department of Agriculture and Consumer

Services, Gainesville. In comprising part of my revisionary studies of

the Nearctic Glyphipterygidae, this paper owes its completion in large

part to the generous support provided by the Department of Entomology

and Nematology, Institute of Food and Agricultural Sciences, Univer-

sity of Florida. My thanks also to curators of the Museum of Compar-

ative Zoology (MCZ), Harvard University, and the National Museum of

Natural History (USNM), Washington, D.C., for making material

available for study. Finally I wish to thank J. A. Powell for providing

most of the specimens for study from his collecting and rearing efforts,

and for kindly making available biological information and larvae of

the new species.

LITERATURE CITED

Braun, A. F. 1940. Notes and new species in the yponomeutoid group (Micro-

lepidoptera). Trans. Amer. Ent. Soc. 66:273-282.

Brock, J. P. 1967 [1968]. The systematic position of the Choreutinae (Lep.,

Glyphipterygidae). Ent. Mon. Mag. 103:245-2406.

Buscx, A. 1906 [1907]. New American Tineina. Proc. Ent. Soc. Wash. 8:86—99.

SCIENTIFIC NOTES

Galapagos Moths.—The appearance of Hayes’ well-illustrated publication on

the larger moths of the Galapagos Islands (Proc. Calif. Acad. Sciences, series 4,

vol. 40, pp. 145-208, 1975) represents a landmark in the study of Galapagos

insects. Including the colored frontispiece which illustrates three adult female

and two larval color phases of the common Manduca rustica calapagensis Holland

(long known under the familiar generic name “Protoparce’’), there are 157

photographs of adult moths representing all but two of the species treated, usually

including both sexes and frequently melanic phases and other variants, and the

male genitalia (in one case the female genitalia, also) of a few of the old species

and all but one of the new species described. Thus, the opportunity is provided

for the non-specialist to identify most of the species by habitus, at least tentatively,

while actually engaged in field studies in the archipelago. Hopefully, it will

stimulate use of these moths in ecological studies, including their role in the

pollination of Galapagos plants (see Linsley, 1966, in Bowman, Proc. Symposia

Galapagos Intern. Sci. Project, Univ. Calif. Press, pp. 225-232). Since many

of the species visit flowers in the daytime or at dusk, their activities lend themselves

VoL. 52, No. 3, JuLy 1976 263

ta observation with less difficulty than during the night. With the publication of

Rindge’s treatment of the Geometridae of the Galapagos (American Museum

Novitates, no. 2501, pp. 1-31, 1973), only the Pyralidae and Microlepidoptera

remain to be studied.

An interesting by-product of the Hayes’ paper is an opportunity to judge, at

least roughly, the impact of man in accelerating immigration and establishment of

moth species from the mainland, as well as results from collecting by entomologists

using both “black” and “white” lights and the rearing efforts of those resident

for various periods at the Darwin Research Station on Santa Cruz (notably R. Perry,

formerly Director of the Station, and Tj. deVries of the Zodlogisch Museum,

Amsterdam). In these respects it may be instructive to make comparisons of the

state of knowledge of the larger moths reflected in the list provided by Linsley

and Usinger (Proc. Calif. Acad. Sci., ser. 4, 33: 113-196, 1966), which with

the exception of the Sphingidae' which had been revised in 1962 and 1964 by

Kernbach, essentially reflected the situation prior to the establishment of the

Darwin Station.

If we confine our comparison to noctuids, Linsley and Usinger recorded 42

identified forms from the Archipelago, based primarily on the reports of Schaus

and Richards published respectively in 1923 and 1941. Of these, nearly half (20)

were regarded as cosmopolitan, pantropical or neotropical in distribution, the

remainder as endemic species or subspecies. By contrast, Hayes records 72 species

of which 33 are regarded as endemic forms (26 are treated as distinct species

and seven as subspecies of mainland taxa).

Of the 20 widespread species listed by Linsley and Usinger, only four were

then known to occur on four or more islands. Hayes’ list, which was based to

a large extent on material collected in the 1960’s after the World War II airbase

on Baltra had been established and abandoned in the 1940’s and post-war commerce

between the islands and the mainland had been expanded during the interim,

included 39 species of widespread mainland occurrence,’ 17 of which are reported

from four or more islands. In spite of the fact that Hayes had available large

amounts of material resulting from intensive collecting efforts by many individuals

with more effective techniques and greater access to the individual islands than

their predecessors, this fact alone could hardly explain the large number of

previously unknown immigrant species represented in the recent collections. Much

of it must be the result of the increased transport and commerce with the mainland.

This is further suggested by the fact that almost all of the continental species

recorded by Hayes from three or fewer islands do not appear in the Schaus or

Richards lists. Many of the recent additions to the Galapagos noctuid fauna are

economic pests elsewhere and: some may well prove destructive in the islands. The

well-known nature of the immigrant species is attested by the fact that of the

39 recorded by Hayes, 10 were named before 1800 and 18 before 1860 (of these

8 each were proposed by Guenée, 1852, and Walker, 1856, although three of the

latter were revived or newly interpreted names).—E. G. Linstey, Division of

Entomology and Parasitology, University of California, Berkeley 94720.

1 Hayes doubtfully included an additional sphingid, a specimen of which was labeled ‘‘Galapagos

and Cocos Islands’? which almost surely does not occur in the Galapagos. Now it can only be

removed from the list by cumulative negative evidence, like the cerambycid Taeniotes hayi Mutchler

(see Linsley and Chemsak, Proc. Calif. Acad. Sci., ser. 4, 33: 233, 1966), a somewhat similar case.

2 Four species listed in an addendum are not included in this tabulation since no data are

available. concerning their occurrence on specific Galapagos islands.

264, THE Pan-PAaciFIC ENTOMOLOGIST

Suspected Cannibalism in Survival of Nymphs of Triatoma protracta

navajoensis.—Ryckman (1962, Univ. Calif. Publ. Entomol., 27:168-69) reported

a maximum survival period without a bloodmeal for 4th instar Triatoma protracia

protracta (Uhler) of 309 days (N=4), and 193 days for 3rd instar nymphs

(N= 4). He postulated a 1,069 day maximum developmental period for nymphs

if fed only at the end of the maximum survival period for each instar. Of 15 bugs

recorded for instars 1, 2 and 3, he noticed that the well-fed nymphs which did

not molt appeared to have a survival advantage of 1 to 10 days over those that did.

In moving stocks of Triatoma protracta navajoensis Ryckman in June of 1973

from Los Angeles City College, Los Angeles County, to Thousand Oaks, Ventura

County, one cardboard box remained buried among boxes of laboratory equipment

along the west wall of the garage. Of 658 nymphs sealed in the box, 5 survived

to feed again on 17.1X.1975 at Thousand Oaks. One individual 4th instar survived

894 days, two 3rd instars 895 days, and two 4th instars 903 days after the last

feeding in Los Angeles. These bugs had access to no food except other bugs. In

order to remain alive, the surviving 3rd and 4th instar nymphs must have sucked

the body fluids of their own or larger instars in these cultures. No record was

made at the time of isolation of the stages in each culture. The dead bugs were

recorded as to instar except for 189 which appeared to be mostly 3rd instars in a

culture where there were no survivors.

One surviving 4th instar nymph was associated with six 5th and twenty-six 4th

instar dead bugs. Two 4th were with fifteen 5th, thirteen 4th and five 3rd instars.

Two 3rd were with eight 4th, forty-one 3rd and twelve 2nd instar dead bugs.

Unfortunately, since all surviving nymphs were very flat, no evidence of color

differentiation (body fluids or blood) could be discerned through the tan exo-

skeleton. Since the writer has shown that 5th and 4th instars of T. p. navajoensis

can take meals of 248 and 119 mgm, respectively, their expected survival may be

much longer than all other stages (Wood, 1976, Ann. Entomol. Soc. Amer., In

Press). The maximum blood meal record for 3rd instars was 56 mgm.

Schenone eé al. (1974, Bol. Chil. Parasit., 72:331-32), summarizing experiences

with Triatoma infestans in xenodiagnosis in Chile, reported that the 4th intsar

nymph survives better than other stages. For T. p. navajoensis, two 4th or four

3rd instar nymphs would be as valuable as one 5th for xenodiagnosis because of

their relatively large blood meals (Wood, 1976, loc. cit.).

The cannibalistic tendency of JT. p. navajoensis may have increased survival

value in relation to the canyon wall habitat of such localities as Chaco Canyon

National Monument (Wood & Wood, 1961, Am. J. Trop. Med. & Hyg., 10:155-65).

The numerous cracks and crevices of these massive sandstone walls offer excep-

tional protection to conenose bugs as well as native rodents. In such a location,

much effort or undue exposure——SHERWIN F. Woon, 614 W. Shenandoah Si.,

Triatoma can easily move in a secluded niche from one host to another without

Thousand Oaks, CA 91360.

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