Vol. 65

January 1989

THE

No. 1

Pan-Pacific Entomologist

LIEBHERR, J. K.—Review of the Palaearctic genus Paranchodemus Habu (Coleoptera: Ca-

rabidae: Platynini). 1

MANLEY, D. G. and J. L. NEFF — Pseudomethoca ilione (Fox), a new synonym of P. gila

(Blake) (Hymenoptera: Mutillidae). 12

SUGDEN, E. A.—A semi-natural, manipular observation nest for Exoneura spp. and other

allodapine bees (Hymenoptera: Anthophoridae). 17

HESSEIN, N. A. and J. A. McMURTRY—Biological studies of Goetheana parvipennis (Gahan)

(Hymenoptera: Eulophidae), an imported parasitoid, in relation to the host species He-

liothrips haemorrhoidalis (Bouche) (Thysanoptera: Thripidae). 25

SIMONS, L. H.—A second record of Tarantula parasitism by Notocyphus dorsalis arizonicus

Townes (Hymenoptera: Pompilidae). 34

DEYRUP, M.—A new species of Pegomya (Diptera: Anthomyiidae) attacking Boschniakia

(Orobanchaceae). 38

OLSEN, S., M. DEYRUP, and I. DEYRUP-OLSEN-Biology of a Pegomya fly (Diptera:

Anthomyiidae) attacking the parasitic plant Boschniakia (Orobanchaceae). 43

MILSTEAD, J. E.—Observations on the host spectrum of the California oakworm, Phryganidia

californica Packard (Lepidoptera: Dioptidae). 50

MARSH, P. M.—Notes on Braconidae (Hymenoptera) associated with Jojoba ( Simmondsia

chinensis ) and descriptions of new species. 58

VICKERY, V. R. and D. B. WEISSMAN —Neonemobius eurynotus (Rehn and Hebard) (Gryl-

loptera: Trigonidiidae: Nemobiinae), a cricket of the San Francisco Bay area, California

. 68

CARMEAN, D., J. C. MILLER, and B. SCACCIA—Overwintering of Phryganidia californica

in the Oregon Cascades and notes on its parasitoids (Lepidoptera: Dioptidae). 74

AKRE, R. D„ C. RAMSAY, A. GRABLE, C. BAIRD, and A. STANFORD-Additional range

extension by the German yellowjacket, Paravespula germanica (Fabricius), in North

America (Hymenoptera: Vespidae)..... 79

RILEY BORDEN, E. E.—The phoretic behavior and olfactory preference of Macrocheles mus-

caedomesticae (Scopoli) (Acarina: Macrochelidae) in its relationship with Fannia cani-

cularis (L.) (Diptera: Muscidae)___ 89

SCIENTIFIC NOTES. 15, 77

SAN FRANCISCO, CALIFORNIA • 1989

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PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 1-11

Review of the Palaearctic Genus Paranchodemus Habu

(Coleoptera: Carabidae: Platynini)

James K. Liebherr

Department of Entomology, Comstock Hall, Cornell University, Ithaca, New

York 14853-0999.

Abstract.— The genus Paranchodemus Habu, new status, is diagnosed and de¬

scribed. It comprises the type species, Anchomenus calleides Bates of Japan, and

P. davidis, n. sp. from Szechwan Province, China. The two species are diagnosed,

and the new species is described. A lectotype is designated for P. calleides, n.

comb. Based on shared-derived characters of the tarsi and the female reproductive

tract, Paranchodemus is phylogenetically similar to the North American genera

Rhadine LeConte and Tany stoma Motschulsky. The characters forming the basis

for this decision are illustrated, and discussed in relation to character-state dis¬

tributions across the tribe Platynini.

In 1978, Habu monographed the Japanese Platynini, proposing placement of

Anchomenus cyaneus Dejean of Europe and the Japanese Anchomenus calleides

Bates in the genus Anchodemus Motschulsky. Because of differences in various

characters, A. calleides was recognized as the type species of a new subgenus within

Anchodemus—Paranchodemus Habu (1978:7).

In the course of revising the closely related platynine genera Anchomenus Bo-

nelli, Sericoda Kirby, Elliptoleus Bates, and Chlaeniomimus Semenov, I have

discovered synapomorphies of the female reproductive tract that necessitate place¬

ment of A. cyaneus in Anchomenus, which is based on Carabus dorsalis Pontop-

pidan. Thus, Anchodemus should be considered a junior synonym of Anchomenus.

A. calleides is recognized as type species of the distinct genus Paranchodemus,

new status. The mainland Asian Paranchodemus davidis, n. sp. is recognized as

a second species of Paranchodemus. Below, I present a diagnosis and description

of Paranchodemus, and a description of the new species. The generic diagnosis

is based on several characters judged to be synapomorphous for the genus, as well

as synapomorphies of greater generality that allow placement of Paranchodemus

in the Rhadine-Tanystoma lineage (Liebherr, 1986:19) of the carabid tribe Pla¬

tynini, subtribe Platyni. This placement is based largely on characters of the tarsi,

which are illustrated and discussed in the context of platynine carabid character

evolution.

Materials and Methods

Taxonomic material for this study was obtained through the courtesy of Nigel

E. Stork, British Museum (Natural History), London (BMNH) and Helene Perrin,

Museum National d’Histoire Naturelle, Paris (MNHP).

Dissection methodology follows Liebherr (1986, 1987). Scanning electron mi-

PAN-PACIFIC ENTOMOLOGIST

crographs were made on an Amray 1000A scanning electron microscope using

gold-palladium coated specimens. The terminology for female ovipositor setation

is drawn from Ball and Hilchie (1983).

Paranchodemus Habu, New Status

Anchodemus (.Paranchodemus ) Habu, 1978, Fauna Japonica, Carabidae: Platyn-

ini, p. 7.

Type species. —Anchomenus calleides Bates.

Diagnosis.— Pronotum lacking setae at hind angles (Figs. 1, 2); mentum with

bidentate median tooth (Figs. 3,4); fourth metatarsomere without dorsal subapical

setae, only ventral setae and apical setae (Figs. 7, 8, 28); seventh elytral stria with

2-4 setae near apex (Figs. 9, 10); abdominal stemites III-V with 2-4 setae each

side (Figs. 11, 12); female apical gonocoxite with 9 furrow pegs in apical pit-like

depression (Figs. 21, 22, 25, 26) and lacking dorsal ensiform seta (Figs. 21, 25);

apical nematiform setae very short and stout (Figs. 21, 25); body dorsum with

metallic blue reflection.

Description. —Head: Eyes convex, with dorsal surface convex relative to deep

supraorbital groove; 2 supraorbital setae; frontal grooves broad, shallow, not

reaching anterior supraorbital setae; labrum with concave anterior margin and 6

apical setae, the median 4 approximate (Figs. 5,6); mentum with bidentate median

tooth (Figs. 3, 4), with well-developed lateral pit-like depressions; antennal scape

robust, third antennomere elongate, subequal in length to scape plus pedicel;

microsculpture well developed, vertex either with rugose wrinkles ( P . calleides )

or rugose punctulae ( P. davidis).

Prothorax: Pronotum cordate, lateral margins broadly arcuate anteriorly and

sinuate in posterior third, hind angles glabrous (Figs. 1, 2); lateral margin narrow

throughout, lateral depression crenulate to punctate; laterobasal depressions heavily

rugose, the surface both wrinkled and punctate; basal marginal bead absent except

near hind angles; median longitudinal depression deep, joined by transverse wrin¬

kles at least basally; anterior transverse impressions shallow; anterior marginal

bead absent, anterior angles slightly projecting; prostemal projection broadly

rounded to slightly acuminate apically in ventral view, not carinate.

Elytra: Basal groove broadly rounded on humeri, lateral marginal depression

narrow throughout; subapical sinuation well developed; sutural apex evenly

rounded ( P. calleides, Fig. 9) to obtuse-rounded (P. davidis, Fig. 10); elytral in¬

tervals broadly rounded, moderately convex; striae deep, well developed; scutellar

seta present at base of sutural stria; 3 or 4 dorsal setae associated with third

interval, the anterior seta in third stria, the more posterior setae in second stria;

2-4 setae in apical portion of seventh stria (Figs. 9, 10); a single seta near apex

of sutural (or first) interval; 17-27 setae set laterad eighth stria from humerus to

subapical sinuation; internal elytral plica obsolete; elytral intervals with strong

isodiametric microsculpture.

Pterothorax: Metepisternum elongate; flight wings fully developed, venation

complete.

Legs: Femora elongate, slender; mesocoxae with single seta on ventrolateral

ridge, mesofemora with 2 setae on anteroventral margin, occasionally the inner

seta doubled; metacoxae bisetose, one seta near anterior margin at base of lateral

VOLUME 65, NUMBER 1

Figures 1-10. 1, 3, 5, 7, 9. Paranchodemus calleides. 2, 4, 6, 8, 10. Paranchodemus davidis, n. sp.

1, 2. Pronotum, dorsal view. 3, 4. Mentum, ventral view. 5, 6. Labrum, dorsal view. 7, 8. Fourth and

fifth left metatarsomere, outer view. 9, 10. Apex of left elytron.

expansion, the other ventrad coxal-trochanteral articulation; protarsi with fourth

tarsomere emarginate apically with two subequal lobes, lobes about 0.50 x as long

as basal portion in females, 0.70-0.85 x as long as basal portion in males; well-

developed dorsolateral sulci on basal 3 tarsomeres of meso- and metatarsi (Fig.

27) , the inner sulcus somewhat weaker; the fourth meso- and metatarsomere

emarginate apically, bearing apical setae but lacking subapical setae (Figs. 7, 8,

28) ; tarsal claws slender, arcuate; apical tarsomere appearing glabrous ventrally,

the ventral setae extremely short (visible at 125 x).

Male genitalia: Parameres glabrous, right or ventral paramere smaller to much

smaller than left paramere (Figs. 16, 18); parameres basally melanistic; median

shaft of aedeagus melanistic, wrinkled basally (Figs. 15, 17); aedeagal internal sac

with very fine spicules; sac as long as apical straight portion of median lobe.

Female reproductive tract: Spermatheca broadly joined to common oviduct at

PAN-PACIFIC ENTOMOLOGIST

Figures 11-14. 11, 13. Paranchodemus calleides. 12, 14. Paranchodemus davidis, n. sp. 11, 12.

Female abdominal stemites III-VI, ventral view. 13, 14. Female reproductive tract, ventral view, sg

= Spermathecal gland; sp = spermatheca; gc = basal gonocoxite.

bursa copulatrix (Figs. 13, 14); spermathecal gland duct entering directly into

reservoir; basal gonocoxite with apical fringe of 15-23 setae (Figs. 13, 14, 19, 23);

apical gonocoxite bearing 3-7 lateral ensiform setae, lacking a dorsal ensiform

seta; apical pit-like depression of apical gonocoxite containing 2 nematiform setae,

and 9 furrow pegs (Figs. 22, 26).

Color: Dorsum of head, pronotum, and elytra piceous with metallic blue re¬

flection; ventral body surface, antennae, palps, and legs piceous to brunneous.

Length: 10.8-13.3 mm.

Paranchodemus calleides (Bates), New Combination

Anchomenus calleides Bates, 1883, Trans. Entomol. Soc. London, p. 256.

Diagnostic combination. — Dorsal body surface piceous with metallic blue-green

reflection; mentum with shallow uniformly sloped mentum pits (Fig. 3); pronotal

disc with strong transverse wrinkles throughout (Fig. 1); elytral striae on disc with

distinct punctulae; seventh stria with 3-5 (usually 4) setae near apex in addition

to single seta near apex of second stria (Fig. 9); visible abdominal stemites III—

V with 2, rarely 3, closely set setae each side (Fig. 11); abdominal stemite VI with

one seta each side in male, two setae each side in female.

Male genitalia: Parameres basally melanistic, the smaller ventral or right par-

VOLUME 65, NUMBER 1

Figures 15-18. 15, 16. Paranchodemus calleides. 17, 18. Paranchodemus davidis, n. sp. 15, 17.

Median lobe of aedeagus with internal sac everted, dextroventral view. 16, 18. Aedeagal median lobe

with internal sac everted and parameres, anterior view, % scale of 15 and 17. gp = Gonopore.

amere rounded apically (Fig. 16); shaft of aedeagal median lobe evenly curved,

apex rounded-acuminate (Fig. 15); aedeagal internal sac covered with very weakly

developed spicules.

Female reproductive tract: Spermatheca apically expanded, globose (Fig. 13);

basal gonocoxite with apical fringe of 17-23 setae on ventral surface (Figs. 13,

19, 20); apical gonocoxite with 2 long acuminate setae basally on lateral margin

(Figs. 13, 19, 20) and 5-7 shorter blunt setae apically and more dorsally on lateral

margin (Fig. 21).

Length: 10.8-11.7 mm.

Lectotype. —9 (BMNH); [card mounted]; Morioka; Japan, G. Lewis, 1910 - 320;

Syn-type [blue bordered label]; Lectotype, Anchomenus calleides Bates, J. K.

Liebherr, 1988.

Distribution. — Bates (1883) described P. calleides from a type series comprised

of specimens from Morioka and Midzusawa, Japan. Habu (1978) reports it to be

generally distributed on the island of Honshu.

Notes. — Habu (1978:7-9) provides a complete synonymy and detailed descrip¬

tion of this species. He incorrectly describes the apical gonocoxite as laterally

bisetose, apparently missing the more apical lateral ensiform setae found dorsad

the lateral margin (Figs. 20, 21).

Habitat.— Reported from “under stones in the Kitakamigawa (Bates, 1883).”

Habu (1978) states that it lives on river beds.

Paranchodemus davidis f New Species

Diagnostic combination. — Body dorsum brilliant metallic blue to blue-green;

lateral depressions of mentum with deep narrowly rounded pit (Fig. 4); strong

PAN-PACIFIC ENTOMOLOGIST

Figures 19-22. Paranchodemus calleides, 9. 19. Left gonocoxa, ventral view, 239 x. 20. Left gon-

ocoxa, outer lateral view, 272 x. 21. Right gonocoxa, dorsal view, 312 x. 22. Apical pit-like depression

with 2 nematiform setae and 9 furrow pegs, 2047 x.

VOLUME 65, NUMBER 1

transverse wrinkles intersecting median longitudinal depression of pronotum, but

discal areas more laterad with at most weak transverse wrinkles (Fig. 2); elytral

striae slightly wavering, but lacking distinct regular punctulae; seventh stria with

2-3 setae near apex, lacking seta near apex of second stria (Fig. 10); visible

abdominal stemites III-V with 2-4 setae in a row near posterior margin each

side; abdominal stemite VI with 1-2 setae each side in male, 2-6 setae each side

in female (Fig. 12).

Description. —Head: Eyes convex, margined mesad by well-defined supraorbital

depression, supraorbital setae separated from supraorbital depression by 1.5 x the

distance of depression from margin of eye; frontal depressions broad, somewhat

irregular, continuous to setae at anterior angles of clypeus; apical margin of clypeus

strongly concave (Fig. 6); clypeus and mandibles brunneous, maxillae and palps

redder; mentum with deep pit in deepest portion of lateral depression (Fig. 4);

diameter of antennal scape 1.33 x the diameter of pedicel apex; third antennomere

1.30-1.45 x length of scape and l.lOx length fourth antennomere; basal 5 an-

tennomeres piceous, apical 6 segments rufopiceous; neck constricted, depressed

on dorsum, the depression densely punctate; microsculpture on vertex weak, an

isodiametric to slightly stretched isodiametric mesh; vertex of head with green to

blue reflection, punctate area of neck with blue to purple reflection.

Prothorax: Pronotum narrow, greatest width 1.10 x width across eyes; lateral

depression narrow in apical half; single lateral seta situated % length anterad hind

angle; hind angles sharply obtuse, distinctly margined; laterobasal depression with

dense rugose wrinkles; mediobasal elevated region rugosely wrinkled; median

longitudinal depression deep, finely engraved, broader and more irregular basally;

strong transverse wrinkles intersecting base of longitudinal depression, transverse

wrinkles weaker forward near anterior transverse depressions and on elevated

portions of disc; anterior transverse depressions strongly bordered anteriorly,

traversed by longitudinal wrinkles toward apex near front angles; front angles

rounded, slightly projecting; proepistemum punctate ventrally; prostemal projec¬

tion broadly rounded apically; discal pronotal microsculpture a transverse mesh;

disc of pronotum with metallic green to blue reflection, anterior and posterior

margins and sternum piceous to rufopiceous.

Elytra: Humeri broadly rounded; elytra widest just behind middle; basal margin

weakly recurved, slightly irregular near bases of striae; scutellar striole long, about

% length distance of anterior dorsal elytral seta from base of third stria; lateral

margin of elytra narrow, subapical sinuation evident (Fig. 10); elytral striae com¬

plete, well impressed, slightly wavering but smooth, not punctate; elytral intervals

only slightly convex, outer intervals flat medially; 3 (or less commonly 4) dorsal

elytral setae, anterior seta in third stria, posterior setae in second stria; apex of

seventh stria with 2-3 setae, single seta near apex of sutural interval (Fig. 10);

16-21 setae laterad eighth elytral interval before subapical sinuation; elytral mi¬

crosculpture a strong isodiametric mesh; elytral margins and sutural intervals with

strong green to blue metallic reflection, elytral epipleura brunneous.

Legs: Mesofemora with 3 setae along anteroventral margin, 4 if the inner seta

is doubled (3 specimens of 7); metafemora bisetose anteroventrally; protarsi of

female with bisulcate first tarsomere; second tarsomere with weak dorsolateral

sulci, median area longitudinally strigose; third tarsomere triangular, with weak

median carina, slightly emarginate apically; fourth tarsomere emarginate apically,

PAN-PACIFIC ENTOMOLOGIST

lobate; protarsi of male with basal tarsomeres 1.6 x as wide apically as in female,

more apical tarsomeres only slightly wider; basal 3 tarsomeres strigose dorsally,

with weak median carina; fourth tarsomere strongly emarginate apically, lobes

O. 70-0.85 x as long as basal portion of tarsomere; basal 3 tarsomeres of meso-

and metalegs with 2 well-developed dorsolateral sulci (Fig. 27); fourth metatar-

somere with broader and slightly longer outer lobe, and a narrower inner lobe

bearing an apical seta (Fig. 28).

Male genitalia: Parameres melanistic; right or ventral paramere very small

relative to spatulate left paramere (Fig. 18); aedeagal median lobe recurved near

apex, short and upturned (Fig. 17); surface of median lobe inside curve melanistic,

wrinkled; internal sac globose, with stronger spicules near apex, gonopore opening

on left side (Fig. 18).

Female reproductive tract: Spermatheca broad and hook-shaped, with acumi¬

nate apex (Fig. 14); basal gonocoxite with apical fringe of 15-20 setae on ventral

surface (Figs. 14, 23); apical gonocoxite with 3-6 ensiform lateral setae set in

groove along lateral margin (Figs. 14, 24); dorsal ensiform seta lacking (Fig. 25);

apical depression with 9 furrow pegs (Fig. 26); apical nematiform setae short and

stout (Figs. 25, 26).

Length: 11.7-13.3 mm.

Holotype.—S; Mou-pin [Mu-Ping, Szechwan, China, 30°30'N, 102°40'E, 1825

m elev.], A. David 1870, Museum Paris (NMHP).

Allotype.— 2; same locality and deposition.

Paratypes. — Mou-pin, A. David, 1870, Museum Paris (2 6, 1 2, MNHP); Thibet,

Chasseurs de Ta-tsien-lou [Kangting, Szechwan, China, 30°10'N, 102°5'E, 2600

m elev.], 1895 (1 <3, 1 2 , MNHP).

Notes. — The type series was found in the main collection of the Paris museum

and bears a hand-written determination of “Anchomenus davidis .” This combi¬

nation has never been published, and I have exhaustively searched the literature

and have found no description of a Chinese platynine carabid that fits this species.

P. davidis appears superficially similar to the European Anchomenus cyaneus

Dejean, but setational, tarsal, and reproductive tract characters indicate its true

affinities.

Habitat. — The valleys around Mu-Ping and Kangting are extremely deep with

steep walls and high-gradient rivers at the bottoms. Based on the habits of P.

calleides in Japan, P. davidis is likely to be found along rivers on bare sandy or

muddy shoreline habitats, where it would hide under stones by day.

Phylogenetic Affinities of Paranchodemus

Placement of Paranchodemus within the tribe Platynini subtribe Platyni can

be accomplished using synapomorphies of the female reproductive tract and tarsal

configuration. The broad duct of the spermatheca observed in Paranchodemus

(Figs. 13, 14) is one basis for recognition of the Rhadine-Tanystoma lineage,

which includes the North American genera Rhadine LeConte, Tanystoma Mot-

schulsky, Atranus LeConte, and Anchus LeConte (Liebherr, 1986). The entry of

the spermathecal duct into the spermathecal reservoir is shared by Paranchodemus

and Anchus. Based on spermathecal configuration, Paranchodemus is not closely

allied to Anchomenus. Anchomenus, Sericoda, and Elliptoleus possess a sper-

VOLUME 65, NUMBER 1

Figures 23-26. Paranchodemus davidis, n. sp., $. 23. Right gonocoxa, ventral view, 239 x. 24.

Right gonocoxa, outer lateral view, 202 x. 25. Left gonocoxa, dorsal view, 232 x. 26. Apical pit-like

depression with 2 nematiform setae and 9 furrow pegs, 1827 x.

PAN-PACIFIC ENTOMOLOGIST

Figures 27-33. 27, 28. Paranchodemus davidis, n. sp. 27. Left basal metatarsomere, dorsal view,

52 x. 28. Left fourth metatarsomere, dorsal view, 72 x. 29. Tanystoma maculicolle, left fourth metatar¬

somere, dorsal view, 73 x. 30. Rhadine caudata, left fourth metatarsomere, dorsal view, 61 x. 31-33.

Apical pit-like depression of female gonocoxite. 31. Tanystoma striata, 1662 x. 32. Tanystoma mac¬

ulicolle, 2532 x. 33. Rhadine caudata, 1194x.

matheca with a long apical filament that is at least several times longer than the

basal reservoir.

The fourth metatarsomere of Paranchodemus species lacks subapical setae (Fig.

28), a character documented across the Platynini by Habu (1978). Subapical setae

are situated on the dorsoapical margin of the fourth metatarsomere, and are nearly

as long as the apical setae found at the ends of the tarsal lobes. Subapical setae

occur in many other Platynini (e.g., species of Calathus, Olisthopus, Anchomenus,

most Agonum, and some Platynus). Paranchodemus shares the derived absence

of subapical setae with Tanystoma (Fig. 29), which possesses only short irregularly

spaced setae on the dorsum of all tarsomeres, and Rhadine (Fig. 30). Anchus and

Atranus species possess subapical setae. This suggests a closer relationship of

VOLUME 65, NUMBER 1

Paranchodemus to Rhadine and Tanystoma than to the other taxa of the Rhadine-

Tanystoma lineage (Liebherr, 1986).

The subtribe Platyni is characterized, among various characters, by gonocoxae

with an apical pit-like depression, in which two long thin nematiform setae and

a variable number of furrow pegs can be found. The number of furrow pegs has

been surveyed in a variety of taxa of the Platynini, and only in the two Paran¬

chodemus species, and in Tanystoma striata Dejean and T. maculicolle Dejean

have as many as nine furrow pegs been observed (Figs. 22, 26, 31, 32). Rhadine

caudata LeConte possesses only two furrow pegs (Fig. 33), the lowest number

observed. Other Tanystoma species and most other platynine taxa exhibit three

to six furrow pegs. The shared possession of the larger number of furrow pegs by

Paranchodemus and some Tanystoma may indicate phylogenetic affinity. By this

interpretation, furrow peg number conflicts with the sister-group relationship for

Rhadine and Tanystoma proposed by Liebherr (1985). A comprehensive cladistic

analysis including all known taxa sharing the above derived characters will be

necessary in order to determine the relationships among these groups.

Acknowledgments

I thank W. L. Brown, Jr. for information on the localities of P. davidis in China,

V. L. Saunders for typing, S. E. Pohl for proofreading, and E. R. Hoebeke for

critically reading the manuscript. A Dean’s Traveling Fellowship from the College

of Agriculture and Life Sciences supported travel to the Museum National d’His-

toire Naturelle, Paris. This research was supported by N.S.F. grant BSR-8614628

and Hatch project NY(C) 139406.

Literature Cited

Ball, G. E., and G. J. Hilchie. 1983. Cymindine Lebiini of authors: redefinition and reclassification

of genera (Coleoptera: Carabidae). Quaest. Entomol., 19:93-216.

Bates, H. W. 1883. Supplement to the geodephagous Coleoptera of Japan, chiefly from the collection

of Mr. George Lewis, made during his second visit, from February, 1880, to September, 1881.

Trans. Entomol. Soc. Lond., 1883:205-290.

Habu, A. 1978. Fauna Japonica, Carabidae: Platynini (Insecta: Coleoptera). Keigaku Publ. Co.,

Tokyo.

Liebherr, J. K. 1985. Revision of the platynine carabid genus Tanystoma Motschulsky (Coleoptera).

J. New York Entomol. Soc., 93:1182-1211.

-. 1986. Cladistic analysis of North American Platynini and revision of the Agonum extensicolle

species group (Coleoptera: Carabidae). Univ. California Publ. Entomol., 106:198 pp.

-. [1987], A taxonomic revision of the West Indian Platynus beetles (Coleoptera: Carabidae).

Trans. Am. Entomol. Soc., 112(1986):289—368.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 12-14

Pseudomethoca ilione (Fox), A New Synonym of P. gila (Blake)

(Hymenoptera: Mutillidae ) 1

Donald G. Manley and John L. Neff

(DGM) Department of Entomology, Clemson University Pee Dee Res. & Educ.

Center, Rt. 1, Box 531, Florence, South Carolina 29501; (JLN) Central Texas

Melittological Institute, 7307 Running Rope, Austin, Texas 78731.

Abstract.—Pseudomethoca gila (Blake) (Hymenoptera: Mutillidae) has been

known only from the male sex. Pseudomethoca ilione (Fox) has been known only

from the female sex. Observations over a 6-yr period have provided sufficient

evidence (including mating pair) that the two are the same species. The name P.

gila has precedence over P. ilione. A complete synonymy is included.

Pseudomethoca gila was first described as Mutilla ( Sphaeropthalma ) gila by

Blake (1871). It has been known only from the male sex. Pseudomethoca ilione

was first described as Mutilla ilione by Fox (1899). It has been known only from

the female sex. Mickel (1924) suggested that P. nephele (Fox) might be the female

of P. gila. He offered no evidence for his suggestion.

Observations

Field data on the association of P. ilione and P. gila were obtained over a 6-yr

period (1982-1987) during a study of the biology of the small panurgine bee,

Pseudopanurgus rugosus (Robertson). A full report of the biology of P. rugosus

will be published elsewhere. P. rugosus is a solitary species in the sense that there

is only one female per nest, although the bees commonly form dense, persistent

nest aggregations. The nesting aggregation of interest for this report is located in

a 2 by 3-m plot on the edge of a path at the Brackenridge Field Laboratory (BFL)

of the University of Texas at Austin, Texas. This aggregation has been active for

at least 9 yr although intensive observations did not begin until 1982. P. rugosus

is the only bee nesting in this plot but nests of Halictus ligatus Say, several species

of Melissodes and several sphecids have been observed nearby. Maximum density

of active bee nests in the plot has varied from fewer than 30 to more than 200

nests over the 6-yr period. P. rugosus is univoltine with nesting activity at BFL

normally occurring from the first or second week of May to the third or fourth

week of June. During the 6 yr of observations, 3-4 males of P. gila and 2-3 females

of P. ilione were regularly present at the nest aggregation during the first two-

thirds of the bee nesting season. A maximum of 14 male mutillids (but no females)

was observed in a 1.5-m area on 3 June 1986. While numerous species of mutillids

1 Technical Contribution No. 2810 of the South Carolina Agricultural Experiment Station, Clemson

University.

VOLUME 65, NUMBER 1

have been collected at BFL, P. gila and P. ilione were the only Pseudomethoca

observed at the nest aggregation.

Female wasps were noted at the P. rugosus nest site only during the period of

bee foraging (normally 0900 to 1430 hr CDT) but males were present up to an

hour before bee activity. Female wasps wandered through the nest area in a

seemingly random manner, occasionally probing nest entrances with their anten¬

nae. Actual entrance of open nests by female wasps was rarely observed. Nests

of P. rugosus are normally closed with a loose soil plug except during foraging

periods. Female mutillids were not observed attempting to enter plugged nests

but this could have easily been overlooked given the density of nests, low numbers

of female mutillids and the fact that most observations concentrated on the pro¬

visioning schedules of the bees. Male wasps also walked over the nest area or

perched on grass stems but paid no attention to bee nests. No aggressive inter¬

actions of wasps and bees were observed. Matings, or mating attempts, of P. gila

and P. ilione were occasionally observed but most females were ignored by males.

One pair taken in copula involving a female which had just exited a P. rugosus

nest proved to be a male of P. gila and a female P. ilione. Brothers (1972) has

noted that previously mated females of Pseudomethoca f frigida (Smith) are not

attractive to males of that species. Although the long term association of the bees

and wasps strongly suggests that P. rugosus is a host of P. gila, no mutillids have

yet been recovered from 100+ excavated nest cells ofP. rugosus.

Based on these observations, it is our contention that P. gila (Blake) and P.

ilione (Fox) are, in fact, male and female, respectively, of the same species. Since

the name P. gila has precedence over P. ilione, that name should stand. A complete,

updated synonymy for the species follows.

Pseudomethoca gila (Blake)

Mutilla ( Sphaeropthalma ) gila Blake, 1871:250. 8

Sphaerophthalma (sic) gila Blake, 1886:245. 8

Sphaerophthalma (sic) gila: Cresson, 1887:265. 6

Mutilla gila : Dalla Torre, 1897:43. 8

Mutilla gila: Fox, 1899:225. 8

Pseudomethoca' ? (sic) gila: Andre, 1903:28. 8

Pseudomethoca gila: Mickel, 1924:13. 8

Mutilla ilione Fox, 1899:268. NEW SYNONYM. 2

Ephuta ( Ephuta ) ilione: Andre, 1903:60.

Photopsis ilione: Krombein, 1951:754.

Pseudomethoca ilione: Mickel, 1965:2.

Mutilla aprica Melander, 1903:322. NEW SYNONYM. $

Pseudomethoca aprica: Mickel, 1924:16.

Literature Cited

Andre, E. 1903. Mutillidae. Genera Insectorum, Bruxelles, 1:1-77.

Blake, C. A. 1871. Synopsis of the Mutillidae of North America. Trans. Am. Entomol. Soc., 3:217-

265.

-. 1886. Monograph of the Mutillidae of North America. Trans. Am. Entomol. Soc., 13:179-

286.

Brothers, D. J. 1972. Biology and immature stages of Pseudomethoca f frigida, with notes on other

species (Hymenoptera: Mutillidae). Univ. Kansas Sci. Bull., 50:1-25.

PAN-PACIFIC ENTOMOLOGIST

Cresson, E. T. 1887. Synopsis of the families and genera of the Hymenoptera of America, north of

Mexico. Trans. Am. Entomol. Soc., Suppl. Vol., pp. 106-107, 263-267.

Dalla Torre, K. W. von. 1897. Catalogus hymenopterorum hucusque descriptorum systematicus et

synonymicus. 8(Fossores):l-99.

Fox, W. J. 1899. The North American Mutillidae. Trans. Am. Entomol. Soc., 25:219-292.

Melander, A. L. 1903. Notes on North American Mutillidae, with descriptions of new species. Trans.

Am. Entomol. Soc., 29:291-330.

Mickel, C. E. 1924. A revision of the mutillid wasps of the genera Myrmilloides and Pseudomethoca

occurring in America north of Mexico. Proc. U.S. Nat. Mus., 64:1-51.

-. 1965. New synonymy and records of Mutillidae for America north of Mexico. Proc. Entomol.

Soc. Wash., 67:1-4.

Muesebeck, C. F. W., K. V. Krombein, and H. T. Townes. 1951. Hymenoptera of America north

of Mexico, synoptic catalog. U.S.D.A. Agric. Monograph 2, 1420 pp.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 15-16

Scientific Note

Host Records for Tetratoma concolor LeConte

and Hallomenus scapularis Melsheimer

(Coleoptera: Tetratomidae and Melandryidae)

Fungus feeding by Tetratomidae and Melandryidae has largely been reported

only in general terms by reviewing authors. Specific hosts seem to have been

published only by de Yiedma (1965, Eos, 41:483), Crowson (1963, Entomol.

Mon. Mag., 99:82), Leschen(1988, Coleopts. Bull., 42:338), Paviour-Smith(1964,

Entomol. Mon. Mag., 100:71, 118), and Weiss (1919, Psyche, 26:132). Here, I

report Lentinus lepideus Fr. (Basidiomycetina: Tricholomataceae) as a host fungus

for Tetratomidae and Melandryidae, and confirm fungal relationships for Tetra¬

toma concolor LeConte and Hallomenus scapularis Melsheimer.

Lentinus lepideus is a relatively abundant lignicolous fungus on barkless, weath¬

ered conifer logs, especially Douglas fir ( Pseudotsuga menziesii (Mirb.) Franco)

and grand fir ( Abies grandis (Dougl.) Forbes). In eastern Oregon, sporocarps have

been seen as early as mid-June, but are in their greatest numbers by mid-July.

Sporulation seems to occur shortly after opening of the cap and continues for

several days to almost 2 weeks. Fresh sporocarps have a rubbery texture with a

dry surface and are persistent through summer, becoming hard and desiccated by

late August. Mycetophagous beetles do not arrive in numbers on the fungus until

cap opening.

Tetratoma concolor has been collected in abundance during July throughout

eastern Oregon forests (Grant, Union, and Wallowa counties) on sporocarps of

L. lepideus Fr. Large series of this beetle, up to 95 specimens per sporocarp, have

been collected at: OREGON, Grant County, Beech Creek campground, elev. 1418

m, 24 km NNE Mt. Vernon, 19.VII.1985; Wallowa County, elev. 1200 m, 17

km W Troy, 22.VII.1985; and Union County, elev. 1500 m, 32 km NE Tollgate

(Umatilla County), 22.VII.1985. Single, or few specimens have been found at a

number of other sites. To date, larvae have not been found on the fungus, or

otherwise discovered.

This beetle is typically found between the lamellae or at the stipe base. Grazing

on basidia was observed several times, with a single observation of notching on

a lamella. The beetle is most abundant during the early growth and sporulation

stages of the host, and gradually becomes scarce through summer as the sporocarps

senesce and desiccate.

Additionally, 3-6 specimens per sporocarp have been found on the mycorrhizal

Boletus edulis (Bull. ex Fr.) Steinpilz (Basidiomycetina: Boletaceae) and on a

Boletus sp. On these latter fungi the beetles were active on the hymenium surface

and within the pores.

Hallomenus scapularis has been found in July, August and early September on

L. lepideus, usually in association with T. concolor (see above collection records).

However, this species is much less abundant than the latter with only 1-7 spec¬

imens per sporocarp. One observation suggests that this species may have been

feeding on basidia, but confirmation of this has not been possible. Most specimens

have been found between lamellae, with relatively few at the stipe base. Relative

PAN-PACIFIC ENTOMOLOGIST

abundance seems to peak in late July and early August, only slightly later than

that of T. concolor, but H. scapularis continues through the senescence stages of

the fungus.

Acknowledgments. — My thanks are extended to J. R. LaBonte, Portland, Maine,

and C. C. Lorain, Moscow, Idaho, for collecting assistance and specimens; and

to J. B. Johnson and J. P. McCaffrey, University of Idaho, for their comments

on an early draft.

Paul J. Johnson, Department of Entomology, University of Wisconsin, Madison,

Wisconsin 53706.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 17-24

A Semi-natural, Manipular Observation Nest for

Exoneura spp. and Other Allodapine Bees

(Hymenoptera: Anthophoridae)

Evan A. Sugden

Exotic Pest Analysis Staff, Analysis and Identification Unit, Division of Plant

Industry, California Department of Food and Agriculture, Sacramento, California

94271-0001. 1

Abstract. — Details are given for construction and use of an observation nest for

allodapine bees. The nest can be made for under U.S. $5.00 per unit using readily

available materials and standard tools. Twenty-six colonies of Exoneura asimil-

lima Rayment were introduced into such nests in Australia and survived for

varying periods. The results of several months of experimentation demonstrate

the utility of the design. All major types of behavior were observed through the

transparent walls. Vigorous colonies progressed in development. Colony survival

is positively correlated with the number of eggs inserted in the starter colony and

positively, although insignificantly, correlated with the number of adult females

and larvae inserted. Such colonies of E. asimillima should be initiated with preex¬

isting nests of at least three females and some brood, including eggs.

Bees of the genus Exoneura are the dominant members of the tribe Ceratinini

throughout the southern half of Australia and in Tasmania. Together with their

allodapine allies (chiefly Allodape), they form a highly interesting group of species

which display various levels of social organization in the nest. Reviews of allo¬

dapine and Exoneura biology appear elsewhere (allodapines: Michener, 1971,

1974; Sakagami, 1960; Wilson, 1971; Exoneura: Michener, 1965,1971; Rayment,

1951). Nonparasitic allodapine species excavate tubular nests in pithy twigs, stems,

plant galls, or flower stalks. Such nests are relatively easy to locate, collect, and

dissect. Females will accept substitute nests and/or nest materials and colonies

can be kept and propagated under artificial conditions.

Studies of the nesting biology of these bees are of great value in determining

the origin and function of sociality in such species. Various types of nests have

been used experimentally. Natural nests in native materials have been collected

and displaced without disturbing the integrity of the nest itself (Sugden and Pyke,

1988; Sugden, unpubl.). Nests have also been dissected, modified, and replaced

in the field (Rayment, 1951; Sugden, unpubl.) and natural nesting materials have

been successfully set out to trap Exoneura spp. (Schwarz, 1986; Sugden, 1988).

A variety of artificial nest materials have been used in experiments involving

colony monitoring. Glass or plastic tubes are sufficient for short term laboratory

1 Present address: USDA-ARS, Honey Bee Research Laboratory, 509 W. 4th Street, Weslaco, Texas

78596.

PAN-PACIFIC ENTOMOLOGIST

work (Maeta et al., 1985; Michener, 1972; Schwarz et al., 1987; Skaife, 1953a,

1953b). Such artificial tubes allow complete visibility of behavior in the nest, but

condensation may become a problem since there is no natural vapor absorbency.

[Absorbent paper strips have been inserted in impermeable tubes (Skaife, 1953a).]

Also, natural nest excavation is precluded. Preference for certain types of materials

is exhibited by some species. A recent study compared the success of several types

of nest materials, including natural stems and glass tube variations (Maeta et al.,

1985). There remains a need for a nest design which is artificial yet acceptable to

allodapine bees for nest building, allows continuous observation of all nest activ¬

ities, permits vapor flow through the lumen walls, allows natural excavation, and

is economical and practical to produce. I present here a design for such a nest

and some preliminary results in its use with Exoneura asimillima Rayment.

Materials and Methods

Construction

Materials. — Table 1 lists parts required to build one nest unit. 1) The foundation

of the nest is provided by a balsa block; its dimensions are variable but the

thickness must be enough to prevent warping if the ends are exposed to weather

(i.e., in a window) and long enough to allow reasonable extension of the nest

lumen through pith excavation. Available at model shops. 2) Hardwood strips,

used in wooden models, provide retaining rails for the acrylic tubing pieces.

Available at model shops. 3) Small diameter acrylic tubing is supplied in 2-m or

6-fit lengths. In the U.S., inside diameters (I.D.) are available in Vi 6 -in. graduations

down to Vs in. Available at plastics supply stores. In Australia, the smallest avail¬

able I.D. at this writing is 4 mm (Cut-To-Size Plastics, Sydney). This was the

diameter used in the study discussed here. Milling the half-tube pieces requires

loss of one longitudinal half of each piece (see below). 4) The pith cylinder can

be obtained from small segments of natural nesting material or cut out of relatively

soft balsa. Ideally, it should be slightly softer than the balsa stock. 5) Optional

light-shielding covers can be made from cardboard and aluminum foil. A sanding

rod is required to form the lumen groove (see below). A Vs-in. brass rod is ideal.

Available at model shops for U.S. $0.40 (not included in Table 1). All materials

are readily available in the U.S. and in Australia at this writing.

Milling. — Cut the balsa stock to desired dimensions. Exoneura asimillima fe¬

males may excavate several millimeters per day and natural nests may reach 0.7

m (Sugden, in prep.), so length of the balsa block must reflect possible final length

of the nest lumen. A 1.0-mm-deep “V”-shaped groove is cut down the length of

the block using a straight edge and a sharp knife. This acts as a guide for sanding

the rounded channel. A layer of fine grit sandpaper is wrapped around a Vs-in.

sanding rod and held tight while rubbing the assembly up and down the groove.

A semi-circular cross section will result as the flat edges of the groove are rounded.

The thickness of the sandpaper must be accounted for in arriving at the final

lumen diameter. Attempts to cut a groove with power tools have met with little

success because proper bits are not available. An improvisation made from the

circular-ground end of a conventional router bit was tried, but tended to shred

the balsa, leaving ragged lumen walls. A smooth-walled lumen is requisite, which

also approximates the diameter of the naturally-excavated space of the bee species

to be introduced to the unit.

VOLUME 65, NUMBER 1

Table 1. Parts required for one observation nest unit, dimensions, and costs in U.S. dollars. Price

for metric diameter acrylic tubing converted from Australian to U.S. dollars, November 1987. (U.S.

cost approx. $0.14/ft.) Notes addressed in text under Materials (M) or Assembly (A).

Cost per Source,

Part Dimensions (mm) Number/amount nest unit notes

Balsa block

Wooden strips

Acrylic tubing

120 grit sandpaper

Pith cylinder plug

Aluminum foil

Cardboard

Wood glue

No. 4 brass wood screws

Small rubber bands

Total

2.5 x 7.6 x 305

1.6 x 1.6 x 305

6 (O.D.) x 4 (I.D.) x 305

4 (O.D.) x 100

305 x 500

850 x 300

$1.90

0.32

1.14

1 sheet

0.37

—

0.05

—

1 ml

0.09

0.73

0.03

$4.63

Acrylic tubing should match the rounded channel in the balsa wood in its inside

diameter. Commercial longitudinal cutting of the tubing by the distributor was

not available at the time of this writing due to the small diameter. However, with

care, the necessary work can be done in a wood shop with standard tools. A jig

is necessary to hold the pieces while being ripped and sanded. This is made by

cutting a groove in a piece of hardwood about 10 x 3 x 30 cm. The groove

should be one full outside diameter (O.D.) wide and Vi O.D. deep, a snug fit for

the acrylic tubing. One end of the groove is blocked off by nailing an end piece

to the wood. Cut the acrylic tubing into 15-20-cm lengths. Place one into the jig

groove. The fine-toothed blade of a band saw can then rip the length of the tubing,

using the face of the jig as a guide. The resulting jig-held piece should be slightly

more than a half-diameter. The assembly is then inverted over a fixed belt sander

to smooth the edges of the band saw cut. The acrylic tubing must fit tightly enough

in the groove of the jig to remain in place as it is lowered onto the sander. The

belt should only graze the face of the jig in smoothing the edges of the tubing.

Removed from the jig, the half-diameter length of tubing can be cut sagittally

into segments on the band saw. The ends are squared and smoothed on a disk

sander, with burrs removed by hand with fine grit sandpaper. Optionally, small

sleeves can be attached to the interior of the tubing ends to integrate the assembled

tubing pieces and baffle the joints (Fig. If). They are made from small rectangles

of thin acetate or acrylic film and set with a quick-dry adhesive such as Super

Glue.

Basal pith plugs can be cut as square-end blanks from natural nesting materials

and rounded into cylinders with sandpaper. Also, soft pith can be pushed or bored

out of stems (Maeta et al., 1985). Especially soft pieces of balsa may substitute.

Assembly. — A complete nest unit is illustrated in Figure 1. Glue the wood strips

with sparing amounts of white glue along either side of the half-lumen groove,

leaving a 1-mm or Vi 6 -in. “shoulder” on which the acrylic half-diameter pieces

will rest. Use a piece of cut tubing as a guide. The tubing should fit snugly between

the glued strips but loosely enough to allow easy removal. Determine the lengths

of the tubing pieces to be used and make leader holes along the side of the wood

base for screws corresponding to the center of each piece of tubing. Place a droplet

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Assembled nest unit, to scale (see Table 1). a. Orientation-facilitating pattern, b. Wooden

strip retainer, c. Pre-lumen space or “runway.” d. Rubber band as tubing brace, e. Brass screw as

rubber band cleat, f. Tubing section sleeve or baffle, g. Acrylic half-tube lumen piece, h. Pith plug.

of glue in each leader hole before inserting screw. The glue will prevent the screws

from working loose in the soft balsa while under tension. Leave 1-2 mm of screw

shank above the surface level of the block as rubber band anchor points.

Set the tubing lengths in place. Plan to have the entrance piece stationary, as

the bees will make a permanent adhering collar of wood fragments a few milli¬

meters inside the lumen or flush with the entrance. It is useful, although not

necessary, to have the end of this first piece set back from the face of the balsa

block by a few millimeters. This allows the bees a “runway” and often it catches

interesting debris ejected from the lumen by the bees. Insert the pith cylinder

several centimeters into the basal piece of tubing. Allow enough covered length

for rapid excavation by vigorous colonies. (See Results for excavation rates.) Extra

plug length should be left to either push into the existing lumen or to extend the

lumen over as the pith is chewed up. The tubing pieces should fit snugly together,

leaving no gaps between them. Small cracks will be plugged by the bees. Stretch

rubber bands between the screws to apply downward tension on the tubing. The

entrance end of the wooden base can be patterned to assist free-flying bees in

orienting to their nest.

Introduction of Bees

Adult bees can be inserted individually through the nest entrance or into the

lumen by removing a segment of tubing. Use cotton wool to plug the lumen from

escaping bees during and after introduction. Cotton swabs cut in half and fluffed

to fit make convenient stoppers. Allow one or more days for the bees to adapt

before removing the cotton wool. They may tear at it and pass little balls of fiber

down the lumen in attempts to dig out. Position nest (if free flight desired). Remove

plug.

VOLUME 65, NUMBER 1

Bees can be kept enclosed in the nest for weeks or months or they may be

allowed to fly freely from a nest position at a window in the lab. Observe under

low light conditions; keep the nest covered when not under observation and do

not expose to sunlight.

Enclosed bees can be fed honey or sugar solution from a pipette (Schwarz et

al., 1987; Sugden, unpubl.) and larvae will consume honey bee-collected pollen

moistened with dilute honey (Rayment, 1951; Sugden, unpubl.).

Results

The tests described here were carried out at Nadgee Reserve, southeastern

coastal New South Wales. Natural nests of Exoneura asimillima used to stock

the artificial nests were collected from sites 10 and 17 km distant from the Lab¬

oratory [described in Sugden and Pyke (1988)].

Four nest units were constructed and bees introduced from natural nests on 17

October 1986. Between mid-October and 14 January 1987, 22 more colonies were

initiated. New colonies were continuously selected to replace those which died

before the end of observations. A total of 26 colonies of Exoneura asimillima

were introduced into 12 nest units. Nests were kept in an unheated room and

allowed free flight into the open environment or a flight cage (8-19 January 1987)

through louvered windows. Populations were censused and colony attrition re¬

corded. Observations extended through 13 April 1987.

Population and survival data are displayed in Table 2. Survival of a colony is

defined as the number of days the nest is continuously occupied by at least one

adult female. Among the 26 colonies used in the study, four received supplemental

additions of immature bees and eight (including one of the latter) survived for

less than 2 days for various reasons. These 11 colonies are not considered in the

analysis below. Of the remaining, survival was positively correlated with number

of eggs initially inserted into the nest (r = 0.5898; 0.05 > a < 0.01; n — 15),

although the data are minimal. The number of adult females and larvae inserted

were positively, but not significantly correlated with survival. Square root trans¬

formation of the data and re-analysis produced equivalent results. It is interesting

to note that, from the entire sample, nests which were started with only one or

two adult females did not survive beyond 17 days and 50% of these (n = 12) died

out after the first day. Two of three colonies in which eggs were produced contained

three adult females at nest initiation and survived beyond the mean number of

days for nests lasting longer than 1 day. The colony producing three eggs had the

second longest survival of any colony, however survival was undoubtedly influ¬

enced by the large number of adults which eclosed from inserted pupae.

A number of photographs of bee activity were taken through the acrylic tubing.

Detailed photographic monitoring was possible through the transparent wall.

Image resolution is only slightly blurred; the tubing joints and rubber bands

obscure some of the view of the nest.

The full range of Exoneura nest behavior was observed in the nests, including

exit and return of foragers, trophalaxis between adults, provisioning of brood,

entrance guarding, excavation and pith collar construction, individual develop¬

ment from egg to pupal eclosion, and male transfer between nests. Some of these

observations will be described in more detail elsewhere (Sugden, in prep.)

Mean pith plug excavation rates ranged from 0.0 to 2.4 mm per day. In five

cases, the basal pith plugs were of a harder consistency than the balsa and the

PAN-PACIFIC ENTOMOLOGIST

Table 2. Data from Exoneura asimillima colonies. Bee data represent numbers of individuals

originally inserted into nests. Numbers in parentheses represent eggs produced by colonies while in

confinement. Subscript 1: all data except nests supplemented with individual bees after initial estab¬

lishment; subscript 2: same data except cases where Survival = 1.

Colony

number

Survival (days)

Eggs

Larvae

Pupae

Adult females

Adult males

163

113

(3)2

(1)0

(1)2

n x = 22

Mearij = 33.45

1.86

3.14

2.59

2.68

0.59

n 2 = 15

Mean 2 = 48.60

1.27

3.40

3.60

3.27

0.47

bees excavated downward into the balsa base. This made manipular access and

observation impossible. Cracks in tubing joints were filled with pith particles by

bees from vigorous colonies but not always by small colonies. The rubber bands

deteriorated rapidly, requiring replacement after 1 or 2 mo. A custom made gable

protruding out of the window protected the nest entrances from rain. The card¬

board and aluminum foil covers worked well in protecting the nests from light.

Ants were not implicated in the demise of any sound nests, although the remains

of bees in one dead colony were carried off by workers of Iridomyrmex sp. Iri-

domyrmex glaber and other species of this genus are apparently major predators

of Exoneura under natural conditions (Rayment, 1951; Sugden, in prep.). Skaife

(1953a) mentions I. humilis as a nuisance with regard to captive Allodape colonies

in Southern Africa.

Discussion

The design presented here successfully simulates natural nest structure. It allows

vapor flow through lumen walls, free excavation of natural pith material in lumen

extension by the bees, and continuous unobstructed observation and photography.

The bee colony can be freely manipulated, construction is simple with standard

tools, materials are readily available, and the cost per unit is small.

VOLUME 65, NUMBER 1

The tests discussed above were done during the middle and late seasons of

activity for Exoneura asimillima, when the age distribution of natural colonies

is shifting toward adults and when less vigorous colonies are dying. Greater colony

growth and perhaps lower colony mortality would have occurred if the artificial

colonies had been started earlier in the season. Despite this, it is obvious that

colony survival following insertion into the artificial nest is dependent on the size

of the population of the initiating colony. Some eggs and a minimum of 3 adult

females from preexisting nests give the best assurance of prolonged colony sur¬

vival.

One possible shortcoming of this design is that it requires the insertion of pre-

established colonies. The artificial nests were not tested to see whether they would

be selected by free-flying foundress females, but given their apparent preference

for natural materials (Sakagami et al., 1985; Sugden, in prep.), this seems unlikely.

[However, Skaife (1953a) coaxed foundresses of African Allodape spp. to select

artificial nests.] Certain reversible modifications of the observation nest entrance

such as the attachment of pieces of predrilled natural nest material might be more

conducive to nest establishment by foundresses. This was not tried.

It is not known whether other species of Exoneura or Allodape would accept

the observation nest described here, although it seems highly probable, based on

similarities in nest biologies of most allodapines.

Acknowledgments

I thank the staffs of the Carpentry and Preparations Departments of the Aus¬

tralian Museum for advice and assistance in developing the observation nest

described here. Keith and Claire Aliendi assisted in data collection and nest

maintenance. Use of the field site and laboratory space was provided courtesy of

the New South Wales National Parks and Wildlife Service. This work was funded

in part by a grant from the Ian Potter Foundation (Australia) and another anon¬

ymous source.

Literature Cited

Erickson, R., and T. Rayment. 1951. Simple social bees of Western Australia. W. Australian Nat.,

3(3):45—59.

Maeta, Y., S. F. Sakagami, and C. D. Michener. 1985. Laboratory studies on the life cycle and

nesting biology of Brauns apis sauteriella, a social xylocopine bee (Hymenoptera: Apidae).

Sociobiology, 10(1): 17-41.

Michener, C. D. 1965. The life cycle and social organization of bees of the genus Exoneura and

their parasite, Inquilina (Hymenoptera: Xylocopinae). Univ. Kansas Sci. Bull., 46(9):317-358.

-. 1971. Biologies of African allodapine bees (Hymenoptera, Xylocopinae). Bull. Amer. Mus.

Nat. Hist., 145(3):221—301.

-. 1972. Activities within artificial nests of an allodapine bee. J. Kansas Entomol. Soc., 45(2):

263-268.

-. 1974. The social behavior of the bees. Harvard Univ. Press, Cambridge, Massachusetts.

Rayment, T. 1949. New bees and wasps—part VIII. A new species of Exoneura, with notes on other

reed-bees from the Grampians. Vic. Nat., 65(1):208-212.

-. 1951. Biology of the reed bees. With descriptions of three new species and two allotypes of

Exoneura. Australian Zool., 11:285-313.

Sakagami, S. F. 1960. Ethological peculiarities of the primitive social bees, Allodape Lepeltier and

allied genera. Insectes Sociaux, 7(3):231-249.

Schwarz, M. P. 1986. Persistent multi-female nests in an Australian allodapine bee, Exoneura bicolor.

Insectes Sociaux, 33:258-277.

PAN-PACIFIC ENTOMOLOGIST

-, O. Scholz, and G. Jensen. 1987. Ovarian inhibition among nestmates of Exoneura bicolor

Smith (Hymenoptera: Xylocopinae). J. Aust. Entomol. Soc., 26:355-359.

Skaife, S. H. 1953a. Subsocial bees of the genus Allodape Lep. & Serv. J. Entomol. Soc. S. Afr.,

16(1):3—16.

-. 1953b. African insect life. Longmans Green and Co., London.

Sugden, E. A. 1988. Secret societies of native bees. Australian Nat. Hist., 22(8):381—384.

-, and G. H. Pyke 1988. Effects of honey bees on the reproduction and nest biology of Exoneura

asimillima Rayment (Anthophoridae: Ceratinini), an Australian native bee. Australian J. Ecol.,

Submitted.

Wilson, E. O. 1971. The insect societies. Harvard Univ. Press, Cambridge, Massachusetts.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 25-33

Biological Studies of Goetheana parvipennis (Gahan)

(Hymenoptera: Eulophidae), an Imported Parasitoid,

in Relation to the Host Species

Heliothrips haemorrhoidalis (Bouche)

(Thysanoptera: Thripidae)

Nawal A. Hessein and James A. McMurtry

Department of Entomology, University of California, Riverside, California

92521.

Abstract. — Observations were conducted on the biology of the greenhouse thrips,

Heliothrips haemorrhoidalis (Bouche), and the imported parasitoid, Goetheana

parvipennis Gahan. Time required for the development of the unparasitized and

parasitized host larvae was recorded. Parasitism extended the second host larval

instar and the prepupa, and prevented molting of the host to the pupa. Devel¬

opmental time of the host and the parasitoid was similar. Temperatures ranging

from 21 to 24°C appeared close to the optimum for the parasitoid in terms of

development, progeny production, especially on avocado leaves, and percentage

of adult emergence.

The greenhouse thrips, Heliothrips haemorrhoidalis (Bouche), is a serious pest

on avocados in California (Boyce and Mabry, 1937; Ebeling and Pence, 1953;

Ebeling, 1959). Only one indigenous parasite, Megaphragma mymaripenne Tim-

berlake, is known for this thrips in California (Ebeling, 1959; McMurtry, 1961;

McMurtry and Johnson, 1963). Although M. mymaripenne attacks a considerable

percentage of greenhouse thrips eggs, its ability to control thrips populations is

questionable (Ebeling, 1959; Hessein and McMurtry, 1988).

The introduction and establishment of additional parasitoids seemed necessary

for more effective biological control of the greenhouse thrips. The eulophid, Goe¬

theana parvipennis Gahan, parasitizes greenhouse thrips as well as the red-banded

thrips or the cacao thrips, Selenothrips rubrocinctus (Giard). Goetheana parvi¬

pennis was introduced from West Africa to Trinidad in 1935, and became estab¬

lished there as well as in other parts of the Caribbean (Callan, 1943; Bennett and

Baranowski, 1982). In 1962, G. parvipennis was introduced to California from

Trinidad (McMurtry and Johnson, 1963) and again in 1982 from the Bahamas.

Although the parasitoid has been recovered from one release site, permanent

establishment cannot be documented at present.

This paper reports on preliminary studies on the biology of G. parvipennis in

relation to that of the greenhouse thrips.

Materials and Methods

Observations on developmental biology of G. parvipennis.—An avocado leaf

was placed dorsal side up on a foam pad in a stainless steal pan containing distilled

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Rearing unit for the parasitoid, G. parvipennis Gahan, with the cover on.

water (McMurtry and Scriven, 1965). The leaf was bordered by a moistened strip

of Cellucotton®, 1 cm wide. A circle, 5 cm in diameter, was created on the leaf

surface by a similar moistened strip of cellucotton. Two hundred early second-

instar thrips larvae, reared on orange fruits, were placed inside this circle. Six

fertilized adult females and five adult males of the parasitoid were then placed

on the same avocado surface with the host larvae. The surface of the circle was

covered by a clear plastic, 13 5-ml food container. For ventilation, four holes were

made in the container and covered by cheese cloth. Searching, ovipositing time,

and frequency of oviposition of the parasitoids were observed through the plastic.

The parasitoids and host larvae were left for 3 days, after which the parasitoids

were removed. Twenty of the host larvae were then placed singly in 1-cm 2 areas

formed by moistened thin cellucotton strips on two avocado leaves. The remaining

host larvae were placed in groups of three on areas, each 2 cm 2 , created as above,

on other avocado leaves. When leaves showed signs of deterioration, the host

larvae were moved to fresh ones. After the parasitoids formed black pupae, they

were transferred to vials, either singly or in groups of three, as above. Behavior

VOLUME 65, NUMBER 1

Figure 2. An adult parasitoid G. parvipennis ovipositing laterally in the first part of the host

abdomen.

of the parasitized host, developmental time, morphology and emergence of the

pupae of the parasitoid were observed. Studies were conducted in the laboratory,

at 21.1-24.4°C and 20-28% RH.

Effect of temperature on developmental time, progeny production, sex ratio and

longevity of G. parvipennis.—To study the effect of temperature on developmental

time, progeny production and sex ratio, ripe Valencia oranges were placed singly

inside inverted 1-liter plastic food containers (Fig. 1) to delay desiccation of the

orange. For ventilation, two circles 5 cm in diameter were cut in the side of the

container and covered by cheese cloth glued to the container. Each orange was

supplied with 200 host larvae and 10 fertilized females and 10 males of the

parasitoid. Four replicates were used for each of three temperatures (17.8, 22.2

and 29.4°C) maintained in small, controlled-temperature cabinets.

To determine longevity in the absence of the host, adult females or males of

the parasitoid were placed in groups in small plastic vials, where a streak of honey

PAN-PACIFIC ENTOMOLOGIST

Figure 3. Emergence of mature parasitoid larva anteriorly from the host.

was supplied. The vials were covered with cloth-ventilated plastic covers. Water

was supplied daily by placing a small piece of cotton, soaked in distilled water,

on top of the cloth. Besides the three temperatures used before, the effect of 13°C

was studied using a similar temperature cabinet.

Developmental period and adult longevity ofYL . haemorrhoidalis.—Five ripe

Valencia oranges were placed singly under 1-liter plastic food containers (Fig. 1).

On each orange, an 8 x 4-cm area was delineated by Tanglefoot®. This area was

subdivided into eight spaces (2x2 cm) by Tanglefoot®. A single adult thrips

was placed in each space. Forty adults were allowed to oviposit for 48 hr and

then removed. After the eggs hatched, all but two of the resulting larvae were

removed. The length of the incubation period, each instar and the longevity of

the adults were determined by daily observations. During the active stages of the

thrips, oranges were replaced by fresh ones every 10 days. Studies were conducted

in small temperature cabinets (Platner et al., 1973) at 23.3°C, 15-60% RH, and

12:12 L:D photoperiod.

VOLUME 65, NUMBER 1

Figure 4. Swollen host thrips with single G. parvipennis larva (right) and superparasitized thrips

(left) with two larvae of the parasitoid.

Results and Discussion

Observations on developmental biology of G. parvipennis. —General observa¬

tions indicated that the adult females usually preferred the early second-instar

thrips for oviposition, although sometimes first or late second instars also were

stung. The prepupa of the host occasionally was parasitized but the parasitoid

did not complete its development and the host always died before reaching the

pupal stage. Entwistle (1972), in his review, stated that G. parvipennis females

will attack nearly mature host larvae but reproduce best in younger stages. The

Table 1. Time required for development of parasitized H. haemorrhoidalis and its parasite, G.

parvipennis, at 21.1-24.4°C.

No. of

Estimated required time in days

Stage

individuals

Avg.

Min.

Max.

2nd to 3rd instar (prepupa)

Unparasitized host

4.7

Parasitized host

11.0

Host 3rd instar (prepupa) to parasite pupa

4.7

Parasite pupa darkening to adult emergence

12.5

Total time required for parasite development 1

a. Males

27.8

b. Females

28.3

1 Oviposition to adult emergence.

Table 2. Time required for adult emergence, percentage emergence, sex ratio and progeny production rate/female of G. parvipennis at two different temperatures.

Temperature

RH%

Time to adult emergence (days)

% adult emergence

Sex ratio (?:S)

No. of pupae formed

Avg.

Min.

Max.

Avg.

Min.

Max.

Avg.

Min.

Max.

Avg.

Min.

Max.

22.2 °e

10-65%

31.6

31.0

33.0

75.5

50.7

83.1

1.99:1

1.32:1

2.45:1

10.10

7.00

13.00

29.4°C‘

20-40%

20.3

17.0

26.0

59.1

42.9

63.2

4.83:1

4.00:1

6.00:1

3.43

1.40

5.70

1 Four replicates, each with 10 fertilized females and 10 males.

PAN-PACIFIC ENTOMOLOGIST

VOLUME 65, NUMBER 1

Table 3. Longevity of Goetheana parvipennis Gahan at different temperatures.

Temperature (° C )

Number of days

No. of adults used

Avg.

Min.

Max.

12.8

46 2

11.3

27 3

21.7

17.8

61 2

16.6

24 3

20.8

22.2

34 2

9.3

36 3

12.4

29.4

44 2

1.1

31 3

2.8

adult female inserted its ovipositor laterally in the anterior part of the host ab¬

domen (Fig. 2). Oviposition time ranged from 23 to 48 sec. Stung hosts were

motionless for a few seconds. Up to 25 host larvae were stung by a single female

in a period of 10-25 min, alternating oviposition with short intervals of cleaning

the ovipositor. Parasitized larvae molted to the prepupal stage. Dohanian (1937)

stated that the development of the parasitoid larvae is at first very slow, until the

host develops nearly to the prepupal stage. After reaching the prepupal stage, the

host was motionless and slightly swollen. The end of the abdomen darkened 2-

3 days later. When the parasitoid completed its larval development, it emerged

from the host anterior through a slit between the prothorax and the mesothorax

(Fig. 3). The pupae of the parasitoid were first whitish in color, but in a few hours

turned black, with the shrunken host integument beneath. Superparasitism oc¬

curred occasionally, where two larvae developed in one host (Fig. 4). In such

cases, one of the pupae was usually atrophied, while the other completed devel¬

opment.

The parasitized second-instar larvae took an average of 11 days to reach the

prepupa stage, compared to only 4.7 days for unparasitized second instars (Table

1). After the host thrips reached the prepupa stage, the parasitoid took an average

of 4.7 days to emerge from the host and pupate. Its position was either perpen¬

dicular or parallel to the host integument. Adult parasitoids emerged in 12-13

days after the pupae darkened. Total developmental time from oviposition to

adult emergence averaged 27.8 days for males and 28.3 days for females at 21.1-

24.4°C. Symptoms of parasitism appeared about 14 days after stinging. Dohanian

(1937) stated that the pupa of G. ( =Dasyscaphus ) parvipennis is jet black and

shining, and that its duration is 10 or 11 days until the emergence of the adults.

Under laboratory conditions, he found that its life cycle ranges from 17 to 21

days.

Mean progeny production of G. parvipennis, where avocado leaves were used

as the substrate, was 25.3/female (for the six fertilized females used in the begin¬

ning of the experiment). Mean percentage emergence was 64.47% and the sex

ratio was 6.54 5:1 6.

Effect of temperature on development, sex ratio, fecundity and longevity of G.

parvipennis.—At 17.8°C, the parasitoid did not reach the pupal instar. The average

developmental time was 31.6 and 20.3 days at 22.2°C and 29.4°C, respectively.

PAN-PACIFIC ENTOMOLOGIST

Table 4. Length of developmental stages of Heliothrips haemorrhoidalis (Bouche) in days at 23.3°C.

Stage

Number of days

Number of individuals

Avg.

Min.

Max.

Egg

18.69

1 st larva

3.42

2nd larva

4.66

Prepupa

1.60

Pupa

2.60

Total

30.97

Though the developmental time was short at 29.4°C, adult emergence was low,

averaging only 59.1%. At 22.2°C, emergence was 75.5% (Table 2). A higher sex

ratio occurred at 29.4°C (average of 4.8 2:1 6) than at 22.2°C (average of 2.8 2:1

<$) (Table 2).

A higher rate of progeny production occurred at 22.2°C (10.1) than at 29.4°C

(3.4) (Table 2). However, the average of 10.1 progeny/female was much lower

compared to the rate on avocado leaves, where it was 25.3 progeny/female when

six fertilized females were used. This last rate was usually the case in the rearing

units of the parasitoid on avocado leaves, suggesting that the host plant might

have an effect on progeny production rate and sex ratio for G. parvipennis. Do-

hanian (1937) also obtained about 25 progeny/female when culturing G. parvi¬

pennis on cacao thrips. Adamson (1936) stated that up to 70 progeny have been

obtained from a single female G. ( =Dasyscaphus ) parvipennis in the laboratory

in Trinidad.

Longevity of the adult parasite was considerably reduced at higher temperatures

(Table 3). Mean longevity of males exceeded that of females at all 4 experimental

temperatures.

The data in Tables 1-3 suggest that temperatures of 21.1-24.4°C are near the

optimum range for development and reproduction of G. parvipennis. Low tem¬

perature, such as 17.8°C, or high temperature, such as 29.4°C, are not favorable.

From the above studies, it is obvious also that this parasitoid, coming from the

Bahamas, might be more successful in tropical areas, where temperature ranges

are moderate.

Developmental period and adult longevity of H. haemorrhoidalis.—The length

of the developmental stages of H. haemorrhoidalis is summarized in Table 4. The

total developmental time averaged 30.97 days, including an incubation period of

18.69 days at 23.3°C. The longevity of the adults ranged from 20 to 58 days, with

an average of 40.6 days. One adult lived to 89 days. Rivnay (1935) reported that

the total developmental time of the greenhouse thrips was 42 days at 23°C, 36

days at 25°C, and 30.8 days at 28°C. He found that the longevity of individuals

reared in test tubes was 40 days at 29-30°C, 55 days at 25-27°C, and 110 days

at 18-20°C. At similar temperature ranges, he found that the individuals reared

on plants did not live as long as those reared on leaf tissue in test tubes.

Literature Cited

Adamson, A. M. 1936. Progress report on the introduction of a parasite of the cacao thrips from

the Gold Coast to Trinidad, B.W.I. Trop. Agric. Trin., 13:62-63.

VOLUME 65, NUMBER 1

Bennett, F. D., and R. M. Baranowski. 1982. First record of the thrips parasite Goetheana parvipennis

(Gahan) (Eulophidae: Hymenoptera) from the Bahamas. Fla. Entomol., 65(1): 185.

Boyce, A. M., and J. Mabry. 1937. The greenhouse thrips on oranges. Calif. Citrogr., 33(1): 19-20,

28-29.

Callan, E. M. 1943. Natural enemies of the cacao thrips. Bull. Entomol. Res., 34:313-321.

Dohanian, S. M. 1937. Life history of the thrips parasite Dasyscapus parvipennis Gahan and the

technique for breeding it. J. Econ. Entomol., 30:78-80.

Ebeling, W. 1959. Subtropical fruit pests. Univ. Calif. Div. Agr. Sci., 436 pp.

Ebeling, W., and R. J. Pence. 1953. Avocado pests. Calif. Agr. Exp. Stn. Cir., 428:1-35.

Entwistle, P. F. 1972. Thysanoptera. In pests of cocoa. Trop. Sci. Series, Longmans, pp. 333-362.

Hessein, N. A.,andJ. A. McMurtry. 1988. Observations on Megaphragma mymaripenne Timberlake

(Hymenoptera: Trichogrammatidae), an egg parasite of Heliothrips haemorrhoidalis (Bouche)

(Thysanoptera: Thripidae). Pan-Pac. Entomol., 64:250-254.

McMurtry, J. A. 1961. Current research on biological control of avocado insect and mite pests.

Yearbk. Calif. Avocado Soc., 45:104-106.

-, and H. G. Johnson. 1963. Progress report on the introduction of a thrips parasite from the

West Indies. Yearbk. Calif. Avocado Soc., 47:48-51.

-, and G. T. Scriven. 1965. Insectary production of phytoseiid mites. J. Econ. Entomol., 58:

282-286.

Platner, G. R., G. T. Scriven, and C. E. Braniger. 1973. Modification of a compact refrigerator for

bio-ecological studies under controlled physical parameters. Environ. Entomol., 2(6): 111 8—

1120.

Rivnay, E. 1935. Ecological studies of the greenhouse thrips, Heliothrips haemorrhoidalis Bouche

in Palestine. Bull. Entomol. Res., 26:267-278.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 34-37

A Second Record of Tarantula Parasitism by

Notocyphus dorsalis arizonicus Townes

(Hymenoptera: Pompilidae)

Lee H. Simons

Department of Zoology, Arizona State University, Tempe, Arizona 85287.

Present address: Graduate Group in Ecology, University of California, Davis,

California 95616.

Abstract.— A second account of parasitism by Notocyphus dorsalis arizonicus

Townes on Aphonopelma chalcodes Chamberlin is described. This pompilid wasp

parasitizes juvenile hosts which remain active until near their death. Larval growth

is rapid with entire consumption of the host within about 1 mo. Notocyphus

dorsalis arizonicus larvae can live either internally or externally on hosts, and

pupal duration can vary from less than 2 mo to 15 mo. Variable pupal duration

may allow wasps to synchronize emergence with optimal seasonal conditions, but

position of the larvae (internal versus external), or uncontrolled proximate factors,

such as temperature or humidity, may have also caused the divergent reproductive

behavior observed in this subspecies.

Notocyphus dorsalis arizonicus Townes is a small (~ 15 mm total length) pom¬

pilid wasp which parasitizes the tarantula Aphonopelma chalcodes (Minch, 1979a).

Here I provide a second account of this wasp’s parasitic behavior including pho¬

tographs at several stages in larval development. I then compare my observation

with Minch’s (1979a) original account.

Between 20 and 21 June 1986, I captured a juvenile Aphonopelma chalcodes

(greatest length from head to abdomen = 18 mm) in a pitfall trap about 30 km

east of Phoenix, Arizona. The trap was set in Sonoran Desert habitat at an ele¬

vation of 500 m. Although the spider behaved normally, a minute white larva

was attached to the right side of the tarantula’s dorsum at the thoracic-abdominal

junction (Fig. 1A).

The tarantula was housed at Arizona State University in a 10-cm square by

5-cm deep clear plastic box. The box had 1 cm coarse sand on the bottom and

several small holes in the lid for ventilation. The tarantula caught and ate several

crickets while the larva increased dramatically in size (Table 1, Fig. 1). The larva

was not seen to release its hold on the tarantula or reorient its position, nor was

the tarantula seen attempting to remove the parasite. The larva seemed strateg¬

ically placed on the tarantula to avoid being dislodged by the host. The larva

pupated on 19-20 July 1986 and emerged as an adult 45-48 days later—on or a

few days before 6 September. The adult wasp was identified with Krombein (1979)

and by comparison to other pompilids in the Zoology Department’s insect col¬

lection at Arizona State University. Dr. Mont Cazier confirmed my identification.

My observations match those of Minch (1979a) in several ways. Again the host

VOLUME 65, NUMBER 1

Figure 1. Six stages in the development of Notocyphus dorsalis arizonicus Townes on the host

Aphonopelma chalcodes Chamberlin. The tarantula remained active until the period between photos

C and D. Tarantula is alive but flaccid in photo D. The larva continued feeding on amorphous remains

of the tarantula in diagram E. The adult wasp emerged 45-48 days after pupating. See Table 1 for a

detailed chronology.

was an immature tarantula active away from its burrow (as evidenced by capture

in a pit-fall trap). Both hosts remained active and fed on crickets while parasitized.

Both larvae grew quickly and completely consumed their hosts (except for frag¬

ments) within about 1 mo of capture. Both larvae spun small brown cocoons

which they exited through circular openings. Two notable differences were 1)

duration of the pupal period: 15 mo in Minch’s case but only 48 days in this

account, and 2) position of the larva on the host: internal in Minch’s case, but

external in my observation.

Apparently, Notocyphus dorsalis arizonicus larvae can parasitize hosts either

internally or externally. Minch fed and handled his tarantula without sign of a

parasite until shortly before the larva emerged from inside the host, at or very

near the time of the host’s death (Minch, pers. comm.). Yet, the larva I observed

developed externally (Fig. 1). These contrasting modes of development within

the subspecies seem remarkably divergent.

Williams (1928, cited in Minch, 1979a) reported a pupal duration of 52 days

for Notocyphus tyrannicus Smith in Ecuador, another parasite of theraphosid

spiders. Minch (1979a) suggested that the shorter pupal duration of Notocyphus

PAN-PACIFIC ENTOMOLOGIST

Table 1. Chronology of development of Notocyphus dorsalis arizonicus on a juvenile tarantula

Aphonopelma chalcodes. Larval length refers to greatest rectilinear length, and larval diameter is greatest

diameter.

Date

Observation

21 June 1986

27 June 1986

1 July 1986

3-4 July 1986

July

1986

July

1986

July

1986

July

1986

July

1986

July

1986

July

1986

July

1986

July

1986

Sept.

. 1986

Larva is 0.5 mm long.

Larva is 0.7 mm long. Tarantula ate 2 subadult (10-15 mm) crickets.

Photograph taken (Fig. 1A). Tarantula active.

Tarantula ate 2 subadult (10-15 mm) crickets. Abdomen bloated to capacity.

Tarantula active.

Larva is 2.2 mm long and 0.8 mm in diameter.

Photograph taken (Fig. IB). Tarantula still active.

Larva is 4.0 mm long and 1.5 mm in diameter.

Larva is 8.0 mm long and 3.0 mm in diameter.

Photograph taken (Fig. 1C). Slight deflation of abdomen beginning around site

of larval attachment. Larva is 8.5 mm long and 3.2 mm in diameter.

Photograph taken (Fig. ID). Tarantula with only partial movement in legs. Ab¬

domen nearly completely deflated.

Drawing made (Fig. IE). Tarantula entirely consumed except for some amor¬

phous tissue nestled in larva’s ventral posterior half.

Larva building cocoon. At least some amorphous tissue still attached to larva.

Larva no longer visible through cocoon wall.

Adult wasp emerged by cutting a 4.5 mm circular exit hole at one end of the

cocoon. Right front wing damaged. Wasp mounted on pin.

tyrannicus resulted from the nearly aseasonal climate of the tropics, while a long

pupal duration in Notocyphus dorsalis arizonicus was an overwintering stage of

this temperate wasp. My observation indicates that pupal duration in Notocyphus

dorsalis arizonicus can actually be quite short.

If larvae are established early enough in the season, Notocyphus dorsalis ari¬

zonicus can complete development and probably breed in the same year. Tarantu¬

las are most active in the Sonoran Desert in late summer and early fall (Minch,

1979b), so hosts should be abundant for wasps emerging in the fall. Other de¬

velopmental patterns may occur at other times of the year, so that Notocyphus

dorsalis arizonicus may be multivoltine. If development varies solely to synchro¬

nize with optimal seasonal conditions, however, it is unclear why Minch’s larva

would then remain encapsulated over the entire period during which my larva

transformed.

Minch’s wasp was from an elevation about 350 m higher than mine, so ele-

vational differences may have led to the variation reported here. Internal versus

external position of the larvae may also influence development by these wasps.

Finally, proximate factors such as temperature or humidity may have altered

normal development in either of these laboratory observations. Additional study,

ideally with controlled experiments, are needed to assess these possibilities.

Acknowledgments

I thank the Department of Zoology, Arizona State University for support, and

John Alcock, Mont Cazier, Edwin Minch, and Laurie Stuart Simons for helpful

comments.

VOLUME 65, NUMBER 1

Literature Cited

Krombein, C. 1979. Catalog of Hymenoptera in America north of Mexico. Smithsonian Institution

Press, Washington, D.C.

Minch, E. W. 1979a. Notocyphus dorsalis arizonicus Townes (Hymenoptera: Pompilidae), a new

host record of theraphosid spiders. The Wasmann Journal of Biology, 37 (1&2):24 -26.

-. 1979b. Annual activity patterns in the tarantula, Aphonopelma chalcodes Chamberlin. Novi-

tates Arthropodae, 1(1): 1—34.

Williams, F. X. 1928. Studies in tropical wasps—their hosts and associates (with descriptions of

new species). Bulletin of the Experiment Station of the Hawaiian Sugar Planters’ Association,

19:1-179.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 38-42

A New Species of Pegomya (Diptera: Anthomyiidae)

Attacking Boschniakia (Orobanchaceae)

Mark Deyrup

Archbold Biological Station, P.O. Box 2057, Lake Placid, Florida 33852.

Abstract. — The species Pegomya hyperparasitica is described from specimens

reared from flowering stalks and ovaries of the parasitic plant Boschniakia hookeri

Walpers.

Boschniakia hookeri Walpers is a perennial parasitic plant associated with sev¬

eral species of Ericaceae in the West Coast region of North America from North

California to North British Columbia. The subterranean tuber-like perennial por¬

tion (called the soma) of mature parasitic plants is attached to the root of a host,

and sends up one or more fleshy flower stalks between early spring and June (Olsen

and Olsen, 1979); during this time an anthomyiid fly attacks the inflorescence.

The larvae of the fly riddle the flower stalks and the developing seed capsules,

reducing or preventing seed production. Although the fly is possibly the most

important natural enemy of Boschniakia hookeri, the attack is restricted to the

inflorescence, leaving the soma unharmed, hence the fly may be considered a

parasite of the parasite. The fly is described here to make the name available for

use in a publication on the biology of Boschniakia. At the very time that these

biological studies were occurring, Griffiths was preparing his exhaustive mono¬

graph on Nearctic Pegomyia (Griffiths, 1982, 1983, 1984a, 1984b). I regret that

I was unaware of this revision, as it would have been preferable to have the

Boschniakia fly described by Griffiths in his monograph.

Pegomya hyperparasitica, New Species

Male.—Length: From frons to apex of basal stemite of hypopygium 5.0 mm.

Coloration: Entirely black except for reddish brown frons and buccae; head

densely white microtomentose except for ocellar triangle and postocciput; thorax

thinly white microtomentose with three faint dark notal vittae barely visible in

oblique lighting; tergites more densely white pollinose than thorax, pygidium and

wide median stripe on tergites 1 + 2-4 black. Wings tinted blackish.

Head (Fig. 1): Width of frons at narrowest point 0.65 as wide as distance between

outer edges of posterior ocelli; frons in lateral view 0.57 as wide as distance from

lower edge of eye to oral margin; third antennal segment 0.56 as wide as long;

arista with minute slender hairs as short as the wide hairs on the third antennal

segment; 8 convergent parorbital bristles, no cruciate bristles on disk of frons,

anterior ocellars proclinate, postocciput with numerous small bristles below post¬

ocular series, buccae along oral margin with numerous bristles posteriorly, reduced

to a single row of strong bristles in anterior half; proboscis short, excluding la-

bellum about as long as front coxae; palps slender, covered with black setulae.

VOLUME 65, NUMBER 1

Thorax: The strong dorsal bristles on each side are 3 humerals, 4 posthumerals,

2 presuturals, 2 presutural dorsocentrals, second presutural acrostical, 1 prealar,

1 supralar, 2 postalars, 2 interalars, 3 postsutural dorsocentrals, 1 basal submar¬

ginal scutellar, 1 discal scutellar, 1 apical scutellar. Prealar as long as first noto-

pleural. Acrosticals except for second pair weak, similar to marginal scutellar

series. Strong pleural bristles are 2 notopleurals, 1 mesopleurals on anteroventral

comer, 2 smaller mesopleurals below first notopleural, 6 mesopleurals in series

along posterior border, the uppermost bristle conspicuously weaker and shorter

than the others, 5 stemopleurals, the ventral pair about half as long as the upper

3. Pteropleuron completely bare.

Legs: Forefemora with 4 rows of posterior bristles; foretibia with medial pos¬

terior bristle, 1 apical posterior ventral bristle, 1 dorsal preapical bristle; foretarsi

with a short stout basal ventral bristle on first segment, first segment ventrally

concave in lateral view, all tarsal segments with a ventral pad of short setulae;

midfemora with posterior ventral, anterior ventral, and anterior rows of bristles

along length of femur, subapical posterior and subapical posterior dorsal bristles;

midtibiae with 1 posterior bristle at end of first third of tibia, 1 posterior and 1

posterior ventral bristle at second third, 1 anterior dorsal bristle at second third,

this bristle about one-half the length of posterior dorsals, a ring of apical bristles;

midtarsi similar to foretarsi; hindfemora like midfemora; hindtibiae with 1 pos¬

terior dorsal and 1 anterior dorsal bristle at end of first third of tibia, 1 anterior

dorsal and 1 posterior dorsal near middle, 1 posterior dorsal at apical eighth of

tibia, 1 apical anterior dorsal, a series of about 10 shorter anterior bristles along

apical half; hindtarsi similar to foretarsi, but ventral bristle at base of first segment

about 2.5 times longer.

Abdomen: Tergites 1-5 with a subapical row of about 6 strong bristles, basal

sclerite of hypopygium with a subbasal row as well, lateral border of tergite 5 with

a row of 5 strong bristles; processes of stemite 5 (Fig. 3) with elongate apices

recurved dorsally, with a strong series of flattened marginal setae in medial section,

a posteriorly projecting flat projection at basal third of processes; postabdomen

(Figs. 6, 7) with mesolobus divided halfway to base, surstyli in dorsal view with

apices bent down, interior lobes expanded and bent upwards.

Female.—Length: From frons to apex of fifth tergite 5.3 mm.

Coloration: Sides of head reddish brown except for black ring around eyes, ring

broken ventrally, and gray lower posterior comer of buccae; frons reddish brown,

becoming black near ocellar triangle; ocellar, parafrontals, postocciput blackish,

grayish brown microtomentose, remainder of head densely white microtomen-

tose; body completely densely grayish brown microtomentose; trochanters, fem¬

ora, tibiae, light reddish brown; tarsi black; wings tinted brownish.

Head (Fig. 2): Width of frons at narrowest point 3.4 times distance between

outer edges of posterior ocelli; antennae and bristles as in male.

Thorax: Strong dorsal bristles as in male; strong pleural bristles as in male

except only 1 lower weaker stemopleural bristle.

Legs: Strong bristles on front legs as in male except for additional dorsal tibial

bristle at second third; forebasitarsus concave with setulae as in male; midfemora

as in male except anterior row of bristles has a gap of about 3 bristles in apical

third; midtibiae and midtarsi as in male except anterior dorsal bristle at second

third of tibia as long as posterior dorsals; metafemur as in male; hindtibiae with

PAN-PACIFIC ENTOMOLOGIST

Figures 1-6. Pegomya hyperparasitica, n. sp. 1. Frontal view of head, male. 2. Frontal view of

head, female. 3. Fifth stemite of male. 4. Gonostyli and mesolobus in posterior view. 5. Gonostylus

and mesolobus in left lateral view. 6. Postgonite and pregonite in right lateral view.

VOLUME 65, NUMBER 1

Figures 7, 8. Pegomya hyperparasitica, n. sp. 7. Hypoproct and cerci of female. 8. Epiproct and

cerci of female.

1 posterior dorsal, 1 anterior dorsal at end of first third, one anterior dorsal and

1 posterior dorsal bristle near middle, 1 posterior dorsal at beginning apical eighth,

1 anterior dorsal and 1 anterior ventral at beginning of apical fourth, 1 anterior

apical, 1 anterior ventral apical; hindtarsi as in male.

Abdomen: Tergites 1 + 2-5 with subapical row of strong bristles, tergite 1 +

2 with a medial band of strong bristles on each side extending a short distance

onto disk; ovipositor in dorsal and ventral view (Figs. 4, 5) conical with a few

long marginal bristles; cerci long and slender, each with 3-4 long apical bristles.

Type material. — Holotype and 4 paratype males from Bear Lake, Bremerton,

Kitsap Co., Washington, reared from inflorescence of Boschniakia hookeri Wal-

pers 21 April 1979. Allotype female same site, ovipositing on flower of Boschni¬

akia hookeri, 11 May 1979. Seventeen paratype males and 24 paratype females,

same site, same host, emerged 18 April; 1 male paratype same site, same host,

emerged 31 March 1979; 1 male paratype same site, same host, emerged 14 April

1979. Holotype, allotype, 10 paratype males, 10 paratype females deposited in

the U.S. Museum of Natural History, Washington, D.C.; 4 paratype males, 4

paratype females deposited in the collection of the California Academy of Sciences,

San Francisco; 5 paratype males, 5 paratype females deposited in the Canadian

National Collection, Ottawa, Ontario; 4 paratype males, 5 paratype females de¬

posited in the collection of the Archbold Biological Station, Lake Placid, Florida.

Etymology. — The specific epithet “hyperparasitica” refers to the parasitic re¬

lationship of the fly to a plant which is itself parasitic.

Discussion.— There is little chance of confusing the male of this species with

other Pegomya as the unusual configurations of the fifth stemite, easily seen in

normally pinned specimens, are diagnostic. The description of the processes of

P. umbripennis Huckett (Huckett, 1966) might give the impression that they are

similar to those of P. hyperparasitica, but the processes of P. umbripennis are not

PAN-PACIFIC ENTOMOLOGIST

reflexed so that the apices are directed anteriorally, nor do they have the large

strongly flattened setae of P. hyperparasitica, and the base of stemite 5 is not

membranous as P. hyperparasitica. The fifth stemite and hypopygial structures

of P. umbripennis are illustrated by Griffiths, 1983. The female can be keyed to

P. sombrina Huckett in Huckett’s 1971 key, but the presence of 4 stemopleurals

and 4 strong metatibial anterodorsals in P. hyperparasitica separates the two

species. There are no other Anthomyiidae or Muscoidea reported as breeding in

Boschniakia or other Nearctic Orobanchaceae. P. hyperparasitica does not appear

to have close morphological affinities to any of the species or species groups

described by Griffiths (George Steyskal, pers. comm.). The strongly enlarged and

expanded seta of the postgonite suggests a closer relationship with the mushroom-

inhabiting rather than the leaf-mining groups of Pegomya.

Acknowledgments

All specimens were reared or collected from host plants by the late Sigurd Olsen

of Seattle, Washington, and by Dr. Ingrith Olsen, of the University of Washington,

Seattle. Dr. Paul Amaud supplied a specimen of Pegomyia umbripennis from the

collection of the California Academy of Sciences. The author is greatly indebted

to the eminent and versatile dipterist, George Steyskal (Cooperating Scientist,

U.S. National Museum), for his encouragement and assistance in preparing this

description.

Literature Cited

Griffiths, G. C. D. 1982. Anthomyiidae. Pp. 1-160 in G. C. D. Griffiths (ed.), Flies of the Nearctic

region, Part 2, No. 1, Vol. 8. Schweizerbart, Stuttgart.

-. 1983. Anthomyiidae. Pp. 161-288 in G. C. D. Griffiths (ed.), Flies of the Nearctic region,

Part 2, No. 2, Vol. 8. Schweizerbart, Stuttgart.

-. 1984a. Anthomyiidae. Pp. 289^408 in G. C. D. Griffiths (ed.), Flies of the Nearctic region.

Part 2, No. 3, Vol. 8. Schweizerbart, Stuttgart.

-. 1984b. Anthomyiidae. Pp. 409-600 in G. C. D. Griffiths (ed.), Hies of the Nearctic region.

Part 2, No. 4, Vol. 8, Schweizerbart, Stuttgart.

Huckett, H. C. 1966. California Anthomyiidae and Muscidae. Proc. Calif. Acad. Sci. Ser. 4, p. 34.

-. 1971. The Anthomyiidae of California exclusive of the subfamily Scatophaginae (Diptera).

Bull. Calif. Ins. Serv., 12:1-121.

Olsen, S., and I. Olsen. 1979. Growth of host root establishes contact with parasitic angiosperm

Boschniakia hookeri. Nature, 279:635-636.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 43-49

Biology of a Pegomya Fly (Diptera: Anthomyiidae)

Attacking the Parasitic Plant Boschniakia (Orobanchaceae)

Sigurd Olsen, f Mark Deyrup, and Ingrith Deyrup-Olsen

(SO) 6542 55th Avenue N.E., Seattle, Washington 98115; (MD) Archbold Bi¬

ological Station, P.O. Box 2057, Lake Placid, Florida 33852; (IDO) Department

of Zoology, University of Washington, Seattle, Washington 98195.

Abstract.— The anthomyiid fly Pegomya hyper parasitica Deyrup deposits its

eggs in the flowers of Boschniakia hookeri Walpers, a parasitic plant growing on

roots of various Ericaceae in western Washington State. Larvae feed in the ovaries,

then bore into the fleshy stem of the inflorescence. Pupae are in soil near the base

of the plant. Adults emerge from late March through April. The fly greatly de¬

creases seed production, without injuring non-reproductive parts of the plant.

In the preceding paper in this issue the anthomyiid fly Pegomya hyper parasitica

Deyrup was described from specimens reared from the plant Boschniakia hookeri

Walpers. B. hookeri is a perennial parasitic plant that attacks roots of a number

of species of Ericaceae along the coastal regions of Washington, Oregon, and Brit¬

ish Columbia. The plant occurs as a tuber-like growth that sends up one or more

fleshy flowering stalks during spring (Figs. 1, 2).

Methods

P. hyperparasitica was studied from the spring of 1977 through the summer of

1979 at Bear Lake (Section 36 of T122N, R1W) in the Puget Sound Lowlands,

on the Kitsap Peninsula in western Washington State. The topsoil is an easily-

permeated gravelly sandy loam. Observations were concentrated in a property lot

of about 270 m 2 , belonging to S. and I. Olsen. An unusually dense population of

B. hookeri, several hundred plants, occurs on this site. The climate of the site is

characterized by mild winters (avg. temp. 4°C) with heavy precipitation (around

1100 mm), and summers that are cool (avg. temp. 18°C) and dry (around 250

mm precipitation).

Pupae were collected in the field by digging below damaged inflorescences during

June-August. Pupae were placed in plastic petri dishes with a thin overlay of soil

or on moistened filter paper, and kept over the winter in an unheated shed.

Cylindrical enclosures covering an area of ground about 225 cm 2 were made from

wire screening covered with mosquito netting (Fig. 3). These cages were set up in

early spring 1979 over the site of plants that had been damaged in 1978. Since

the flies emerge before the plants, the same enclosures could be used to exclude

flies in 1979. Smaller enclosures, about 80 cm 2 were also used to exclude flies,

t Deceased.

PAN-PACIFIC ENTOMOLOGIST

Figures 1-3. 1. Flowering stalk of B. hookeri. 2. Complete plant of B. hookeri. 3. Cages used to

both trap and exclude flies.

VOLUME 65, NUMBER 1

B. hookeri

Figure 4. Emergence of P. hyperparasitica (n = 132) and blooming period of B. hookeri (n = 585).

but were ineffective in trapping. Six large and 7 small traps were set out on 19

March 1979 and examined for flies weekly through June 1979.

Details of the effects of larvae on the flowers and the stem were photographed

with a Nikon-F camera using Nikon PB-4 bellows and Zeiss Luminar lenses (100,

63, 40 mm). Illumination was provided by a Bowens Multitec Tecturelite.

Life History and Behavior of Fly

In the field adult emergence begins in late March and continues into early June,

based on records of 132 flies captured in 1979. Seventeen flies emerged from

pupae kept in petri dishes over the winter; these began to emerge 4 days earlier

than flies collected in the field, and had completed emergence 9 days earlier. The

first flies emerging in the field appeared 23 days before the first flowers opened,

and emergence was complete long before the blooming season reached its peak

(Fig. 4). This implies that the flies require considerable maturation time, or that

there is a strong advantage associated with ovipositing on the earliest flowers.

Even in the laboratory, where flies were maintained in closed containers and not

fed, the flies lived for several weeks, easily spanning the time between eclosion

and availability of Boschniakia flowers. Except for the flies emerging in enclosures,

no flies were seen in the field until ovipositing females appeared on the flowers

in May.

Oviposition was observed repeatedly and the typical oviposition stance pho¬

tographed (Fig. 5). The female backs into the flower until about half the abdomen

is concealed in the flower. A single egg is laid on an anther (Fig. 6). The ovipositor

is rather simple compared with that of Pegomya species that insert eggs into the

tissues of plants, but is equipped with long hairs that might be used to locate an

PAN-PACIFIC ENTOMOLOGIST

Figure 5. P. hyperparasitica ovipositing in flower of B. hookeri.

anther. Upon hatching, the larva bores into the ovary (Fig. 7), where it feeds on

the developing seeds (Fig. 8). When the young seeds have been eaten, the larva

moves into the soft stem of the inflorescence, working downward while feeding

(Fig. 9). The larvae leave the stem near ground level and burrow down to about

the level of the tuber-like body of the plant. Pupation occurs in this zone.

The feeding habits encompassed in the genus Pegomya are diverse, including

leaf-mining, boring in canes of Rubus, stems of Equisetum, and stems of mush¬

rooms (Griffiths, 1982, 1983, 1984a, 1984b). Of the 94 species listed by Griffiths,

hosts are known for 49; 27 species are leaf-miners, and 18 are mushroom feeders.

The morphological characteristics of most of the species with unknown hosts

strongly suggest that these are also leaf-miners or mushroom feeders. P. hyper¬

parasitica is the only species known to feed on seeds and flower stalks and is thus

ecologically as well as morphologically isolated from other Pegomya. The structure

of the male pregonite and postgonite of P. hyperparasitica, especially the enlarged,

flattened seta of the postgonite (likely to be a derived character state), suggest a

closer relationship with some of the fungus-feeding species groups. Since B. hookeri

is somewhat fungoid in appearance, lacks chlorophyll (possibly of nutritional

significance), and can be attacked without any specialized structures on the ovi¬

positor, there is some ecological justification for considering P. hyperparasitica a

radical offshoot of a mycetophagous lineage.

VOLUME 65, NUMBER 1

Figures 6-8. 6. Egg of P. hyper parasitica on anther of B. hookeri. 7. Entrance hole of fly larva. 8.

Fly larva among seeds.

We strongly suspect that P. hyper parasitica is a monophagous species. The

specific oviposition site and the two sequential larval feeding sites suggest a spe¬

cialized species. We have reared a few of these flies from pupae collected around

B. strobilacea Gray in Oregon, but since this plant is often considered a southern

variant of B. hookeri (Hitchcock et al., 1959), these records do not affect the

monophagous status of the fly. We examined local specimens of two other parasitic

plants, Allotropa virgata Torrey and Gray (Ericaceae) and Hemitomes congestum

Gray (Ericaceae), but found no evidence of attack by flies. These are the only

local parasitic plants with succulent stems like that of Boschniakia, though floral

and stem morphology are quite different from Boschniakia.

Effects of P. hyperparasitica on its Host

Some insects, such as yucca moths and fig wasps, whose larvae feed on deve¬

loping seeds, guarantee seed set by pollinating the host plant. The hairy abdomen

of P. hyperparasitica may transfer pollen, but this transfer is not necessary for

seed set, as the pollen of each flower is shed directly onto the stigma. Seed pro¬

duction and seed viability of flowers on plants in exclusion cages is equal to that

of undamaged flowers on plants left in the open. The principle cross-pollinators

appear to be bumble bees (Bombus spp.). If a substantial number of fly eggs fail

to develop, or if the flies often make abortive oviposition attempts, the flies might

be significant cross-pollinators, but this seems highly unlikely.

The larva has a great effect on seed production. Seed counts from 10 randomly

collected plants taken in 1978 and again in 1979 indicated that about 90% of the

seed crop was destroyed by flies both years. Undamaged capsules remaining on

the plant usually develop normally, in spite of the damage to the stem. B. hookeri

PAN-PACIFIC ENTOMOLOGIST

Figure 9. Larval damage to stem of inflorescence.

is not usually an abundant plant, and the fly may well be partly responsible for

its scarcity, though there is no way to test this idea.

Although the fly has a great effect on the inflorescence, it does not attack the

underground tuber-like body of the plant. The fly does not exhaust the plant’s

resources, as no nutrient-storing portions are attacked, and may actually reduce

VOLUME 65, NUMBER 1

the nutritional drain by the inflorescence if ovaries are consumed before their

allocation of nutrients is complete. In this sense we consider the fly parasitic, as

it never kills its host plant.

Conclusion

P. hyperparasitica is previously unknown species that appears to be morpho¬

logically and ecologically divergent from any known species or species groups of

Pegomya. It attacks the reproductive parts of a species of plant which is itself

morphologically and ecologically isolated. Although these flies can be easily found

once the host is known, both fly and host are relatively uncommon organisms.

This furnishes another example of a species that would not be known to ento¬

mologists were it not for the observations of botanists.

Acknowledgments

We thank George Steyskal (Cooperating Scientist, U.S. National Museum) for

his assistance in preparing the description of P. hyperparasitica, and Sean Morris

(Oxford Scientific Films Ltd.) for providing the photograph of the fly used in Fig¬

ure 5.

Literature Cited

Griffiths, G. C. D. 1982. Anthomyiidae. Pp. 1-160 in G. C. D. Griffiths (ed.), Flies of the Nearctic

region, Part 2, No. 1, Vol. 8. Schweizerbart, Stuttgart.

-. 1983. Anthomyiidae. Pp. 161-288 in G. C. D. Griffiths (ed.), Flies of the Nearctic region,

Part 2, No. 2, Vol. 8. Schweizerbart, Stuttgart.

-. 1984a. Anthomyiidae. Pp. 389-408 in G. C. D. Griffiths (ed.), Flies of the Nearctic region,

Part 2, No. 3, Vol. 8. Schweizerbart, Stuttgart.

-. 1984b. Anthomyiidae. Pp. 409-600 in G. C. D. Griffiths (ed.), Flies of the Nearctic region,

Part 2, No. 4, Vol. 8. Schweizerbart, Stuttgart.

Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1959. Vascular plants of the

Pacific northwest, Vol. 4. Ericaceae through Campanulaceae. Univ. Washington Press, Seattle,

510 pp.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 50-57

Observations on the Host Spectrum of the California Oakworm,

Phryganidia californica Packard (Lepidoptera: Dioptidae)

James E. Milstead

Division of Entomology and Parasitology, University of California, Berkeley,

California 94720.

The California oakworm, Phryganidia californica Packard, is a herbivore com¬

monly found on coast live oak in the San Francisco Bay Area. The larvae are

reported to occasionally overwinter on deciduous oaks (Burke, 1919) but the

species appears to require evergreen foliage in order to maintain spatial and

temporal continuity within the boundary limits of its potential hosts.

The host spectrum of the California oakworm appears to consist primarily of

members of the family Fagaceae. Kellogg and Jack (1895) recognized 5 native

species of oaks: Quercus agrifolia, Q. lohata, Q. kelloggii, Q. dumosa and Q.

douglasii that could serve as hosts. This list was subsequently expanded to include

Q. chrysolepis (Kellogg, 1908), Q. suber (Essig, 1915), Castanea dentata and Eu¬

calyptus globulus (Burke and Herbert, 1920), Q. tomentella, Q. bicolor, Q. rubra,

Q. robur, Lithocarpus densiflora, Carya ovata, C. tomentosa, Castanea sativa,

Betula papyifera (Sibray, 1947), Q. durata (Harville, 1955), Q. velutina, Q. vir-

giniana (Tietz, 1972), Rhododendron occidentale (Fumiss and Carolin, 1977) and

Castenopsis chrysophylla (Wickman and Kline, 1985).

While it is generally held that all native and introduced oaks can support

oakworm development, temporary feeding on unrelated native and exotic species

(here termed incidental hosts) may occur during periods of high population density

when preferred foliage has been depleted (Sibray, 1947) and larval starvation is

imminent. Cursory observation of such behavior may lead to unwarranted as¬

sumptions of nutritional adequacy for these incidental hosts which are never

subsequently tested by controlled rearing studies.

More than 150 species including native flora associated with oaks as well as

fruit trees and ornamentals have been tested on newly hatched and nearly mature

larvae (Sibray, 1947) with generally negative results. Unfortunately this author

failed to provide a complete list of species tested so that it is impossible to verily

many of his findings.

The current study reports on rearing trials conducted between 1983 and 1987

on native and exotic oaks as well as other oak associates located primarily on the

University of California Campus at Berkeley.

Materials and Methods

Two types of rearing trials were conducted: 1) in which a single host species

was utilized throughout the entire larval period and 2) in which outbreak con¬

ditions were simulated and last instar larvae were transferred from Q. agrifolia

leaves to leaves of other species (terminal hosts).

All larvae were maintained at 20°C in 13-dram plastic vials and fed mature

VOLUME 65, NUMBER 1

Table 1. Influence of incidental host plants on survival of lirst-instar larvae of the California

oakworm.

Host

No. of

trials

Month foliage sampled

n larvae

tested

Age at 100% mortality 1

(days) (x ± SE/trial)

Alnus rhombifolia

March

144

8.0 ± 1.5

Arbutus menziesii

Aug., Sept.

222

5.2 ± 1.0

Arctostaphylos Columbiana

Apr.

10.0 ± 0

Betula papyifera

March, Apr., May

724

34.4 ± 4.6

Eucalyptus globulus

May

4.6 ± 0.4

Fagus grandiflora

Sept.

14.0 ± 0

Fagus sylvatica

Sept.

34.0 ± 0

Garrya eliptica

Jan., Aug.

143

7.6 ± 1.4

Heteromeles arbutifolia

Mar.

6.0 ± 0

Liquidambar styraciflua

Mar.

317

7.8 ± 0.8

Ulmus americana

Apr., May

162

24.0 ± 14.0

Ulmus procera

Oct.

145

14.8 ± 1.8

Starvation

328

6.0 ± 0.3

1 No larvae survived beyond the second instar.

excised leaves selected from the lower crown area of the same donor trees. Foliage

was replaced 3 times per week. No attempt was made to control or monitor the

relative humidity. The initial number of larvae was determined by the egg mass

size and all larvae were reared individually after the first instar.

Feeding arenas were examined daily and mortalities recorded. Insects were

weighed at the onset of pupation and adult eclosion.

Matings were carried out in plastic vial arenas. Q. agrifolia leaves were replaced

3 times per week to provide a fresh ovipositional substrate.

Leaf moisture was estimated by comparing initial weights of fresh leaves with

weights obtained after a 24 hr ovenization at 105°C (Volney et al., 1983).

Table 2. Influence of native and exotic fagaceous host plants on the survival of California oakworm

larvae.

Host species

No. of insects reared

% survival to adult

Quercus agrifolia

273

Q. chrysolepis

Q. douglasii

Q. englemanii

Q. kelloggii

Q. lobata

Q. alba

Q. cerris

Q. ilex

Q. macrocarpa

Q. mongolica

Q. phellos

Q. robur

Q. rubra

Castenopsis sempervirens

Lithocarpus densiflora

Table 3. Influence of native and exotic fagaceous hosts on developmental rate, size and fecundity of California oakworm.

Treat¬

ment

Host species

Duration of larval period (days) (X ± SE)

Pupal weight (mg) [X ± SE)

n S

n 2

Q. agrifolia

54.1 ±

2.8

53 45.7 ± 2.8

60 110.0 ±4.9

Q. chrysolepis

65.4 ±

7.4

6 52.2 ± 2.1

10 86.8 ± 9.4

Q. douglasii

50.9 ±

1.1

28 52.5 ± 2.4

24 104.3 ± 3.3

Q. kelloggii

44.3 ±

0.5

5 55.8 ± 6.3

4 149.8 ± 23.1

Q. lobata

47.0 ±

1.1

16 38.7 ± 0.7

30 132.3 ± 5.9

Q. macrocarpa

45.0 ±

1.0

21 36.8 ± 0.8

22 163.2 ± 5.0

Q. mongolica

47.6 ±

3.5

22 49.1 ±5.9

18 138.7 ± 9.0

Native species

F = 2.73, P

= 0.05

F = 2.53, P = 0.05

F= 5.82, P = 0.01

All species

F = 2.91, P

= 0.01

F = 2.72, P = 0.05

F = 17.77, P = 0.01

Paired contrasts significant at:

P = 0.05

2 vs. 7

1,7 vs. 6

1 vs. 4; 2 vs. 3

P = 0.01

2 vs. 3

1 vs. 5,7; 2 vs. 4

P= 0.001

2 vs. 5,6; 3 vs. 6

2,3,4 vs. 5,6

2,3 vs. 5; 3 vs. 4; 1,2,3,5 vs. 6;

2,3,6 vs. 7

PAN-PACIFIC ENTOMOLOGIST

Table 3. Continued.

Treat¬

ment

Pupal weight (mg) (.? ± SE)

Adult weight (mg) (Jc

± SE)

Eggs laid (X ± SE)

Total

71.4 ± 4.9

92.4 ± 4.7

34.5 ± 1.6

98.6 ± 7.5

56.0 ± 6.1

67.2 ± 9.5

25.8 ± 2.8

79.1 ± 20.2

70.9 ± 2.1

79.2 ± 2.8

32.5 ± 1.9

90.5 ± 8.6

60.7 ± 7.9

109.6 ± 23.3

44.2 ± 8.5

179.5 ± 17.5

87.4 ± 3.7

104.5 ± 5.3

46.4 ± 5.2

122.9 ± 10.8

99.8 ± 3.3

128.3 ± 4.8

50.4 ± 4.1

156.4 ± 13.5

80.8 ± 3.5

115.0 ± 9.6

44.2 ± 2.9

163.9 ± 27.8

F = 6.33, P = 0.01

F= 13.93, P = 0.01

F= 4.52, P = 0.01

F = 8.22, P = 0.01

T = 4.58, P = 0.01

F = 6.51, P= 0.01

T= 2.71, P = 0.05

F= 4.98, P = 0.01

Paired contrasts significant at:

P= 0.05

1 vs. 2; 1,3,4 vs. 7; 5 vs. 6

1 vs. 2,7

2 vs. 4,5

1,2 vs. 4; 3 vs. 7; 2,3 vs. 5

P= 0.01

1,4 vs. 5; 2 vs. 3,6

2 vs. 5,7; 5 vs. 6

1,3 vs. 5; 2 vs. 6; 1,2,3 vs. 7

1 vs. 7; 2 vs. 6; 3 vs. 4

P= 0.001

2,3 vs. 5; 1,2,3,4,7 vs. 6

1,2 vs. 6; 3 vs. 4,5,6,7

1 vs. 6

1,3 vs. 6

C/i

VOLUME 65, NUMBER 1

LT\

Table 4. Influence of terminal host plants on survival, larval longevity and duration of larval period of last instar larvae of the California oakworm.

Treat¬

ment

Host species

No.

leaves

assayed

% moisture

{X ± SE)

No.

larvae

tested

survival

Larval longevity

(days) (£ ± SE)

Duration of larval period (days) (Jc ± SE)

n 2

Arctostaphylos Columbiana

52.0 ± 0.2

9.5 ± 0.2

Betula papyifera

14.3

18.1 ± 2.2

1 24.0 ± 0

22.4 ± 4.8

Eucalyptus pulverulenta

49.1 ± 0.3

10.8 ± 0.3

Garrya eliptica

48.6 ± 0.6

9.9 ± 0.4

Heteromeles arbutifolia

48.1 ± 0.3

1.8

11.6 ± 0.6

Lithocarpus densiflora

46.5 ± 0.4

88.6

12 18.5 ± 0.8

14.9 ± 0.5

Quercus agrifolia

49.9 ± 0.3

92.0

25 17.2 ± 0.7

13.3 ± 0.3

Umbellularia calif or nica

52.9 ± 0.3

11.1 ± 0.7

Starvation

9.9 ± 0.3

F= 38.06, P = 0.01

F = 67.0, P = 0.01

F — 9.89, P = 0.01

F =

19.9, P = 0.01

Paired contrasts significant at:

P =

0.05

1 vs. 8; 3 vs. 5

1 vs. 8; 4 vs. 5

P =

0.01

4 vs. 6

1 vs. 3,5; 2 vs. 8,9

6 vs.

2,7

P =

0.001

1,8 vs. all; 6 vs. 3,5,7;

2 vs. 1,3,4,5

2 vs. 7

5 vs. 7

PAN-PACIFIC ENTOMOLOGIST

VOLUME 65, NUMBER 1

Data were analyzed using an analysis of variance. Means were compared using

the method of least significant difference.

Results

The results of a series of feeding trials in which attempts were made to rear

first-instar larvae on incidental hosts are presented in Table 1. Only Betula, Fagus

and Ulmus supported survival for as long as 2 wk and no larval development to

the third instar was observed. Negative results were also obtained with the fall

foliage of Corylus cornuta var. californica and Ribes divaricatum. These disparities

in larval performance are not attributable to leaf age.

Similar trials were conducted using foliage taken from native and exotic oaks,

tanbark and chinkapin. The results of these trials (Table 2) contrast sharply with

those of Table 1. Only Q. niger and Q. laurifolia failed to support development

to the adult stage and with both species, feeding and frass production was observed.

During the late summer flight period of 1986, male moths were observed hov¬

ering about the canopies of all of these native and exotic species on the Berkeley

Campus with the single exception of Q. mongolica. Oviposition and larval feeding

was also observed in the Tilden Park Botanical Garden, Berkeley, California on

Q. agrifolia, Q. chrysolepis, Q. dumosa, Q. durata, Q. douglasii, Q. dunni, Q.

garryana, Q. garryana var. semota, Q. kelloggii, Q. lobata, Q. x macdonaldi, Q.

parvula, Q. saddleriana, Q. tomentella, Q. turbinella var. turbinella, Q. turbinella

var. californica and Q. wislizenii.

Extensive defoliation also occurred on Q. agrifolia, Q. chrysolepis and chinkapin

in the Huckleberry preserve, Oakland, California.

In addition oakworm were observed to feed on Q. turbinella foliage collected

from the Mohave Desert near Chloride, Arizona and the insects were reared to

adulthood on foliage of Q. gambelii (collected from the Mingus Mountains, Pres¬

cott, AZ) and Q. vaccinifolia (collected from Echo Lake, El Dorado, CO).

The influence of foliage sampled from Lithocarpus densiflora and Quercus species

on developmental rate, insect mass and fecundity is presented in Table 3. Ratios

of pupal wet weight to the duration of the larval period (pw/t) show that growth

rates for females are greater than that for males on comparable foliage. The highest

values for female larvae occurred on the deciduous species Q. macrocarpa, Q.

kelloggii, Q. lobata and Q. mongolica. Feeding trials with the latter species using

small immature leaves, however, consistently resulted in 100% mortality. Ratios

of realized fecundity to pupal weight paralleled pw/t values. The poorest perfor¬

mance occurred on Q. chrysolepis and Q. douglasii.

Tables 4 and 5 present data derived from experiments in which larvae reared

during the early instars on Q. agrifolia foliage were transferred at the onset of the

last instar to foliage of a terminal host.

Table 4 compares the influence of coast live oak and tan oak with native and

exotic incidental hosts on larval/pupal survival, larval longevity and duration of

the larval period. European birch proved clearly superior to the other incidental

hosts tested in supporting development. However, in contrast with Quercus agri¬

folia and Lithocarpus, larvae reared on Betula suffered a much higher mortality,

required a longer larval period to reach pupation and produced smaller pupae

and adults with greatly reduced fecundity (Table 5). A single male survivor on

Heteromeles weighed only 6 mg and probably would have been incapable of

mating.

PAN-PACIFIC ENTOMOLOGIST

Table 5. Influence of terminal host plant on pupal weight, adult weight and fecundity of California

oakworm.

Treat-

Pupal weight (mg) {X ± SE)

ment

Host species

n 2

Betula papyifera

1 72.6 ± 0

47.5 ± 4.6

Lithocarpus densiflora

12 108.6 ± 9.3

75.2 ± 2.9

Quercus agrifolia

25 137.4 ± 6.4

81.4 ± 13.9

Paired contrasts significant at:

P= 0.05

P = 0.01

F = 4.72, P = 0.05

2 vs. 3

F= 14.63, P = 0.01

1 vs. 3

2 vs. 3

Discussion

Although Phryganidia californica is not strictly monophagous (Puttick, 1986)

it does appear to generally restrict itself to fagaceous hosts.

The inability of most native and exotic incidental hosts to support feeding and

development of first-instar larvae precludes the possibility of such species playing

a direct role in oakworm population dynamics since the insect overwinters at this

stage in its life cycle. These plants, however, may provide allomonal protection

to later larval and pupal stages from natural enemies.

While the population density of Phryganidia is generally sparse, heavy infes¬

tations periodically result in severe defoliation to fagaceous hosts and under such

circumstances oakworm larvae may be observed crawling and feeding on nearby

plant associates.

These species can be termed true hosts if larval feeding results in the production

of robust adults capable of mating and producing viable offspring.

Of the nonfagaceous terminal hosts studied, only Betula papyifera, a species

limited to ornamental plantings, would marginally qualify for inclusion as a host.

It is possible that a related species B. occidentals, which may be sympatric with

California oakworm populations reported from Siskiyou County, California (Mil¬

ler, 1987) could augment larval feeding in streamside habitats.

Scriber and Slansky (1981) have shown that relative larval growth rate is a

function of leaf water content. This relationship does not serve to explain the

disparity between Lithocarpus and Arctostaphylos, Eucalyptus, Heteromeles or

Umbellularia and there may be allomonal factors present in the foliage of these

species that act as feeding deterrents for oakworm larvae.

In a comparison of native oak species, Puttick (1986) has reported that Q. lobata

was a better quality food source for California oakworm larvae than Q. agrifolia.

Larvae reared on Q. lobata had greater efficiencies of conversion of ingested and

digested food which led to more rapid larval development, greater pupal mass

and enhanced fecundity. These findings are supported by the results of this study.

Her generalization that deciduous oaks are a better food source than evergreen

oaks is partially supported by evidence indicating that larval performance and

adult fecundity was highest on Q. kelloggii and Q. lobata and lowest on Q. chry-

solepis. However, current data suggest that the foliage of coast live and blue oak

are essentially equivalent in their effects. Since the mean duration of the larval

period ranged between 2 and 3 wk for both males and females it is possible that

VOLUME 65, NUMBER 1

Table 5. Continued.

Adult weight (mg) (X ± SE)

Eggs (x ± SE)

n 2

n Total

1 48.2 ± 0

12 89.0 ± 8.3

25 115.8 ±5.3

F = 4.65, P = 0.05

F= 1.98

26.5 ± 4.5

42.5 ± 2.5

44.5 ± 1.8

1 38.0 ± 0

9 114.0 ± 18.6

22 160.5 ±11.4

F = 4.31, P = 0.05

1,2 vs. 3

1 vs. 2,3

reports of the occurrence of three oakworm generations per year in the San Fran¬

cisco Bay Area may be partially attributable to host mediated differences in larval

developmental rate.

Because information is not yet available on the spatial and temporal variability

of host metabolites and the possible influence of prior herbivory on the allocation

of these metabolites quantitative differences in host nutritional suitability can at

best be only rough approximations.

Results from the current study suggest that all Quercus species, Lithocarpus

densiflora, Castenopsis chrysophylla and C. sempervirens qualify as true host species

whose foliage can support larval development in areas where temperature and

humidity regimes allow oakworm populations to persist.

Literature Cited

Burke, H. E. 1919. Notes on the California oak worm. Proc. Ent. Soc. Wash., 21:124-125.

-, and F. B. Herbert. 1920. California oak worm. Farmers Bulletin, 1076:3-11.

Essig, E. O. 1915. Injurious and beneficial insects. Supp. Bull. California State Host Comm., Vol.

4, Berkeley, California, 503 pp.

Fumiss, R. L., and V. M. Carolin. 1977. Western forest insects. U.S.D.A. Forest Service Misc. Publ.

1339, Washington, D.C., 654 pp.

Harville, J. P. 1955. Ecology and population dynamics of the California oak moth Phryganidia

californica Packard (Lepidoptera: Dioptidae). Microentomology, 20:83-166.

Kellogg, V. L. 1908. American insects, 2nd ed. Henry Holt, New York, 674 pp.

-, and F. J. Jack. 1895. The California Phryganidian ( Phryganidia californica Pack.). Proc.

Calif. Acad. Sci., Series 2, 5:562-570.

Miller, J. S. 1987. A revision of the genus Phryganidia Packard, with description of a new species

(Lepidoptera: Dioptidae). Proc. Entomol. Soc. Wash., 89:303-321.

Puttick, G. M. 1986. Utilization of evergreen and deciduous oaks by the California oak moth

Phryganidia californica. Oecologia (Berk), 68:589-594.

Scriber, J. M., and F. Slansky, Jr. 1981. The nutritional ecology of immature insects. Ann. Rev.

Entomol., 26:183-211.

Sibray, W. S. 1947. Bionomics of the California oak moth. M.S. thesis, Dept, of Entomology, Univ.

California, Berkeley.

Tietz, H. M. 1972. An index to the described life histories, early stages and hosts of the macrolepidop-

tera of the continental United States and Canada, Vol. 1. Allyn Museum of Entomology,

Sarasota, Florida, 536 pp.

Volney, W. J. A., J. E. Milstead, and V. R. Lewis. 1983. Effects of food quality, larval density, and

photoperiod on the feeding rate of the California oakworm (Lepidoptera: Dioptidae). Environ.

Entomol., 12:792-798.

Wickman, B. E., and L. N. Kline. 1985. New pacific northwest records for the California oakworm.

Pan-Pacific Entomologist, 6:152.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 58-67

Notes on Braconidae (Hymenoptera) Associated with Jojoba

(Simmondsia chinensis) and Descriptions of New Species

Paul M. Marsh

Systematic Entomology Laboratory, U.S. Department of Agriculture, Agricul¬

tural Research Service, % U.S. National Museum of Natural History, NHB 168,

Washington, D.C. 20560.

Abstract. — A brief review is given of 13 species of Braconidae that were asso¬

ciated with insects feeding on jojoba, Simmondsia chinensis, in the southwestern

United States. In addition to host and distributional data, four new species and

one new genus are described.

Jojoba, Simmondsia chinensis (Link) Schneider (Buxaceae), a shrub native to

the southwestern United States, has recently been studied for its economic po¬

tential as an oil producer (Sherbrooke and Haase, 1974; Scarlett, 1978; Yermanos,

1979). In conjunction with this, studies have been undertaken recently to survey

the arthropods associated with jojoba (Pinto and Frommer, 1980, 1984; Pinto et

al., 1987). During these studies I was asked to identify the Braconidae that were

collected, and this paper is a review of those species and is intended to provide

names and taxonomic information important for future studies on the biological

control of jojoba pests.

This review covers only those Braconidae that were actually reared from a host

insect attacking jojoba branches or leaves, or where there was little doubt of host

association with an insect from the plant. Of the 13 species collected, four are

described as new, including one new genus. All of the braconid and host specimens

are deposited in the collection of the University of California, Riverside (UCR)

except the holotypes and some paratypes of the new species which are in the U.S.

National Museum of Natural History, Washington, D.C. (USNM) and the Mu¬

seum of Comparative Zoology, Harvard University, Cambridge, Massachusetts

(MCZ).

Key to Species of Braconidae Associated with Jojoba

Keys to restricted groups of parasites such as this one are limited in their

accuracy and may give misleading information to the user. I suggest that braconid

parasites associated with jojoba should also be run through the key to North

American genera provided by Marsh et al. (1987), which also should be consulted

for explanation and illustration of characters mentioned in the key and descrip¬

tions below.

1. Abdominal terga forming a rigid dorsal carapace that covers most of

remainder of abdomen . 2

- Abdominal terga not carapace like, sutures usually distinct. 3

VOLUME 65, NUMBER 1

2. First cubital and first discoidal cells of fore wing separated, basal segment

of cubitus present (Fig. 2) . Ascogaster shawi, n. sp.

- First cubital and first discoidal cells confluent, basal segment of cubitus

absent (Fig. 4). Chelonus ( Microchelonus) periplocae McComb

3. Space between clypeus and mandibles forming a circular opening when

mandibles closed . 4

- Space between clypeus and mandibles absent . 7

4. Fore wing with 2 cubital cells, wings mottled (Fig. 8) .

. Percnobraconoides jojoba, n. gen., n. sp.

- Fore wing with 3 cubital cells, wings not mottled (Figs. 5, 9, 10). 5

5. First intercubitus of fore wing weak or absent, first and second cubital

cells confluent (Fig. 5). Heterospilus frommeri, n. sp.

- First intercubitus present, first and second cubital calls separated (Figs.

9, 10) . 6

6. Abdominal terga granular; stigma in fore wing uniformly clear yellow,

nervulus vein at most slightly beyond basal vein or intersecting basal

vein (Fig. 10) . Xenosternum ornigis Muesebeck

- Abdominal terga longitudinally puncto-striate; stigma brown with basal

half and spot at apex yellow, nervulus well beyond basal vein by distance

greater than its length (Fig. 9) . Aleiodes buoculus, n. sp.

7. Forewing with 2 cubital cells (Fig. 1). 8

- Forewing with 3 cubital cells, second cell sometimes small and triangular

(Figs. 3, 6, 7). 9

8. Ovipositor longer than abdomen; propodeum with a triangular areola

margined by carinae . Apanteles sp.; A. aristoteliae Viereck

- Ovipositor barely exserted and much shorter than first abdominal seg¬

ment; propodeum without and areola, often with a median longitudinal

carina . Cotesia spp.

9. Second cubital cell of fore wing triangular, first cubital and first discoidal

cells confluent (Fig. 3). Bassus binominata (Muesebeck)

- Second cubital cell rectangular or square, first cubital and first discoidal

cells separated (Figs. 6, 7) . 10

10. First abdominal segment narrow at base, abruptly widening at apex, at

least 3 times wider at apex than at base . Meteorus sp.

- First abdominal segment parallel sided, base and apex nearly equal in

width . Homolobus truncator (Say)

Aleiodes buoculus Marsh, New Species

(Fig. 9)

Female. — Length of body, 4-4.5 mm. Color: Body generally honey yellow, eyes

and ocellar triangle black, mesonotal lobes, mesopleuron, propodeum, and apical

abdominal segments frequently light brown; wings hyaline, veins brown, stigma

yellow on basal Vi and at extreme apex. Head: Eye very large, malar space short,

at most Vs eye height; mouth opening small, width only slightly longer than malar

space, clypeus swollen; temple small, about % eye width; ocelli large, ocellar-

ocular distance about x h diameter of lateral ocellus; face, vertex and temple punc¬

tate, frons granulate; antenna with 39 flagellomeres. Thorax: Propleuron rugose;

mesonotum and scutellum punctate, notauli weakly scrobiculate, meeting in a

PAN-PACIFIC ENTOMOLOGIST

Figures 1-10. Wings of braconid species. 1. Apanteles aristoteliae Vier., fore wing. 2. Ascogaster

sham, n. sp., fore andhind wings. 3 .Bassus binominata (Mues.), fore wing. 4. Chelonus ( Microchelonus )

periplocae McC., fore wing. 5. Heterospilus frommeri, n. sp., fore and hind wings. 6. Homolobus

truncator (Say), fore wing. 7. Meteorus sp., fore wing. 8. Percnobraconoides jojoba, n. gen., n. sp., fore

and hind wings. 9. Aleiodes buoculus, n. sp., fore and hind wings. 10. Xenosternum ornigis Mues.

Scale line = 0.5 mm.

wide rugose area; mesopleuron granular-punctate, rugose near wing base and at

area of sternaulus, sternaulus not distinct; propodeum granular-rugose with a weak

median longitudinal carina. Abdomen: First tergum slightly longer than apical

width, granular-strigate; second tergum granular-strigate; third and following terga

VOLUME 65, NUMBER 1

granular-punctate; first, second and sometimes extreme base of third tergum with

median longitudinal carina; ovipositor about l h length of hind basitarsus. Legs:

Tarsal claw without basal tooth; inner spine at apex of hind tibia Vi length of

basitarsus. Wings (Fig. 9): Fore wing with first segment of radius x h length of second

segment and Vi length of recurrent vein, nervulus postfurcal by distance greater

than its length; hind wing with radiellen sinuate and slightly widening at apex,

basella nearly as long as second segment of mediella, postnervellus present.

Male.— Essentially similar to female.

Holotype female.— ARIZONA, Pinal Co., 7 mi W of Superior, 2500 ft, 4 Oc¬

tober 1980, “parasite pupa inside geometrid hosts seen 13 Oct 1980, adult wasp

emerged 24 Oct 1980.” Deposited in USNM.

Paratypes.— 4 9, 2 $, same data as holotype except dates from 4 May 1979 to

7 June 1980. Deposited in USNM and UCR.

The specimens were reared from an unnamed geometrid larva and listed as

Rogas sp. by Pinto et al. (1987).

This species will key out to the genus Rogas in the generic key to North American

Braconidae (Marsh et al., 1987) but, according to van Achterberg (1982), most

of the species now included in Rogas should be placed in Aleiodes Wesmael. The

genus Aleiodes is presently being revised by me and S. R. Shaw. A. buoculus

belongs to a group distinguished by extremely large eyes and ocelli which con¬

tribute to the short malar space and narrow temples; it is separated by its wing

venation with the nervulus in the fore wing being beyond the basal vein by a

distance greater than its length and by the bi-colored stigma.

The specific name is from the Latin prefix bu- meaning large and the Latin

oculus meaning eye.

Apanteles aristoteliae Viereck

(Fig. 1)

Locality. — 1 5, 5 6, CALIFORNIA, Riverside Co., 5.6 mi S of Sage.

Host. —Epinotia kasloana McDunnough.

This species, listed by Pinto and Frommer (1984) and Pinto et al. (1987), has

been reared from various species of Tortricidae and Gelechiidae but this is the

first record for E. kasloana.

Apanteles Species A

Locality. — 3 $, CALIFORNIA, Riverside Co., 5.6 mi S of Sage.

Host. —Epinotia kasloana McDunnough.

This species is listed in Pinto and Frommer (1984) as Apanteles sp. C and in

Pinto et al. (1987) as Apanteles sp. Because the specimens are all males it is

difficult to accurately identify the species.

Ascogaster shawi Marsh, New Species

(Fig. 2)

Female. — Length of body, 3.5-4 mm. Color: Black; mandibles brown on apical

half; anterior surface of scape, pedicle, and flagellomeres 1-8 orange or light brown;

palpi yellow; fore and middle coxae, trochanters, femora, and anterior surface of

tibiae honey yellow; apical fifth of hind coxa honey yellow, hind trochanters honey

yellow, hind femur varying from honey yellow with black apex to black on entire

anterior surface and apical half of posterior surface, apical half of hind tibia honey

PAN-PACIFIC ENTOMOLOGIST

yellow, hind tibial spurs yellow, hind basitarsus honey yellow on basal third; fore

wing slightly dusky with darker dusky patch below stigma, veins brown, hind

wing veins clear. Head: Face, vertex, and temple coarsely punctate; clypeus sparse¬

ly punctate and shining, lower margin notched medially; frons rugulose, carina

between antennae extending onto dorsal third of face; antenna with 30 flagello-

meres (broken in all paratypes), flagellomeres 1-8 longer than wide, remainder

shorter and more compact. Thorax: Pronotum with a dorso-medial depression,

coarsely punctate, somewhat areolate anteriorly; mesonotum coarsely punctate,

areolate-rugose postero-medially, notauli scrobiculate; scutellum sparsely punc¬

tate and shining, pre-scutellar furrow 5-6 foveate; mesopleuron smooth to sparsely

punctate medially, areolate-rugose dorsally and along stemaulus; propodeum

strongly areolate-rugose, lateral caudal tubercles blunt, median caudal tubercles

weak or absent. Abdomen: Carapace bicarinate basally, rounded apically, weakly

areolate-rugulose; ventral cavity not reaching carapace apex; ovipositor short, not

reaching carapace apex when exserted. Wings (Fig. 2): First segment of radius

about 1 Vi times longer than second segment, third segment about 3 Vi times longer

than first segment; stigma about 1 Vi times longer than radial cell along wing margin.

Male. — Essentially as in female.

Holotype female. —CALIFORNIA, Riverside Co., Sec. 32, T.7S, R.1E, Site 2,

116°54'W, 33°31'N, 5.6 mi S Sage onR3, 6 February 1980, S. Frommer. Deposited

in USNM.

Paratypes.— 2 2, 3 6, same data as holotype with dates of 26 February 1976,

25 May 1979, and 27 May 1980, and four of them labelled as reared from Epinotia

kasloana McD. Deposited in USNM, UCR, and MCZ.

Host.—Epinotia kasloana McDonnough. Pinto and Frommer (1984) list this

parasite as Ascogaster sp. and Pinto et al. (1987) list it as Ascogaster provancheri

group.

This species will run to provancheri Dalla Torre in Shaw’s (1983) key to the

North American species of Ascogaster (couplet 6) but sham is distinguished by

the more coarsely punctate face, the weak or absent median caudal tubercles on

the propodeum, the less coarsely sculptured abdominal carapace, and the dark

markings on the antennae and legs. This species is named for my colleague Scott

R. Shaw in recognition of his excellent study of the genus Ascogaster.

Bassus binominata (Muesebeck)

(Fig. 3)

Locality. — 1 2, 1 6, CALIFORNIA, Riverside Co., 5.6 mi S of Sage.

Host. —Epinotia kasloana McDunnough.

This species was previously placed in the genus Agathis and listed in Pinto and

Frommer (1984) and Pinto et al. (1987) as Agathis sp., but Sharkey (1985) has

recently shown the distinctness of Bassus from Agathis. This species was known

only from the eastern U.S. and this is an unusual extension of its range. It has

been reared from various leaf mining Lepidoptera including Epinotia, but this is

a new record for E. kasloana.

Chelonus ( Microchelonus ) periplocae McComb

(Fig. 4)

Locality.— CALIFORNIA, Riverside Co., Palm Desert.

Host. —Periploca sp.

VOLUME 65, NUMBER 1

This species, known only from California, was previously recorded by Pinto

and Frommer (1980) and Pinto et al. (1987) as being reared from Periploca sp.

in jojoba leaves. The type series was reared from P. nigra Hodges (McComb,

1968).

Cotesia Species

Species of Cotesia were included in the genus Apanteles prior to the study by

Mason (1981). Five specimens which apparently represent two species were reared

from jojoba. Because the genus is badly in need of revision and only a few

specimens of these species are available, I have chosen to list the species below

without specific names.

Cotesia Species A

Locality . — 3 9, 1 6, ARIZONA, Pinal Co., 7 mi W of Superior.

Host. —Anacamptodes obliquaria Grote.

This species does not agree with other Cotesia reared from Anacamptodes and

it is possibly undescribed. Pinto et al. (1987) list it as Cotesia sp. 1.

Cotesia Species B

Locality. — 1 9, ARIZONA, Pinal Co., 9 mi W of Superior.

Host. — Glaucina eureka (Grossbeck).

I have been unable to find records of any species of Cotesia, or any braconid

for that matter, reared from this geometrid. This species is also apparently un¬

described; it is listed by Pinto et al. (1987) as Cotesia sp. 2.

Heterospilus frommeri Marsh, New Species

(Fig. 5)

Female.— Length of body, 4 mm; ovipositor, 1.5 mm. Color: Head, thorax and

abdomen dark brown, abdomen with light brown patches on second tergum and

apical terga; antenna brown; legs honey yellow. Head: Face, frons and vertex finely

strigate, temple smooth; malar space x h eye height; ocellar-ocular distance about

2.5 times diameter of lateral ocellus; antenna with 22 flagellomeres. Thorax:

Propleuron rugose; mesonotal lobes finely granulate-strigose; notauli scrobiculate,

meeting in a wide rugose area with two distinct carinae; scutellum smooth; me-

sopleuron smooth with striae around stemaulus, stemaulus impressed but smooth;

propodeum rugose with distinct carinae dorsally outlining a central areola and

two basal areas. Abdomen: First tergum rugose, wider at apex than long; second

tergum strigate-rugose; third tergum strigate on basal Vy remainder of terga smooth;

ovipositor about % length of abdomen. Legs: Fore tibia with irregular row of

about 20 stout spines on inner edge. Wings (Fig. 5): Second segment of radius

slightly longer than first segment.

Male. — Unknown.

Holotype female.— CALIFORNIA, Riverside Co., Riverside, UCR Field 4C,

September 1985, S. Frommer, emerged Oct.-Nov. 1985, associated with tunnels

of Amphicerus cornutus (Pallas). Deposited in USNM.

Paratypes. — 7 9, same data as holotype. Deposited in USNM and UCR.

This species is distinctive by its rugose first abdominal tergum and dark col¬

oration. The type series was reared from jojoba branches which contained only

the bostrichid Amphicerus cornutus. Although not definitely reared from this

PAN-PACIFIC ENTOMOLOGIST

beetle, these were the only insects taken from the plant indicating a likely asso¬

ciation of the two species. It is listed as Heterospilus sp. by Pinto et al. (1987).

The species is named for its collector, Saul Frommer.

Homolobus truncator (Say)

(Fig. 6)

Locality. — 1 $, CALIFORNIA, Riverside Co., 5.6 mi S Sage.

Host.—Periploca sp.

This species was listed in the Hymenoptera catalog (Marsh, 1979) as Zele mellea

(Cresson), which is a junior synonym, and was previously recorded by Pinto and

Frommer (1980) as Zele sp. It occurs over the entire U.S. and has been recorded

from various species of Noctuiidae.

Meteorus Species

(Fig. 7)

Locality. — 1 $, 1 ?, ARIZONA, Pinal Co., 9 mi W of Superior.

Host.— Undetermined geometrid.

The genus Meteorus is also in need of revision and the species are not easily

identified. Because of this and the fact that one specimen is missing its abdomen,

I am not able to give a specific name. Pinto et al. (1987) list it as Meteorus sp.

Percnobraconoides Marsh, New Genus

Type species.— Percnobraconoides jojoba Marsh, n. sp.

Diagnosis. — Head cubical, space between clypeus and mandibles circular, distal

margin of clypeus concave, labrum concave; mesonotum strongly declivous an¬

teriorly; fore tibia (Figs. 12, 13) with two irregular rows of stout spines on anterior

side, spines on outer side normally sharp pointed, those on inner side bluntly

pointed; hind coxa with weak basal tubercle on inside; fore wing (Fig. 8) with two

cubital cells, radial cell short a wide, last segment of cubitus arcuate, intercubitus

about as long as recurrent vein, nervulus present and interstitial with basal vein,

brachius weak, first brachial cell open at apex; hind wing (Fig. 8) with radiella

absent, cubitella unpigmented and visible only in reflected light, nervellus present,

hind wing of male with large stigma at base; abdomen somewhat petiolate, first

tergum narrow at base and with strong basal depression (glyma) on each side;

ovipositor curved upward.

This genus is distinctive for North America and will run to couplet 145 of the

recent key to braconid genera (Marsh et al., 1987) where it can be separated from

Polystenidea by the stout spines on the fore tibia (Fig. 12). These spines are

distinctive for the subfamily Doryctinae to which this genus belongs. In addition

to these stout spines, the inner side of the tibia has several rows of unusual spines

which are somewhat flat and bluntly pointed, as distinguished from the normal

sharply pointed spines on the outer side (Figs. 12,13). I have not seen this character

in any other doryctine genera. Percnobraconoides is also similar to the North

American genus Pambolidea but is distinguished by the femora not being swollen.

It is also very similar to the Central and South American genus Percnobracon

Kieffer, but Percnobraconoides can be distinguished by its wing venation, partic¬

ularly the long first segment of the radius and short second segment of the cubitus

in the fore wing and the stigma in the male hind wing, by the first abdominal

VOLUME 65, NUMBER 1

Figures 11-13. Percnobraconoides jojoba, n. gen., n. sp. 11. Female habitus (scale line = 1 mm).

12. Fore tibia. 13. Enlarged view of spines on fore tibia.

PAN-PACIFIC ENTOMOLOGIST

tergum which is much wider at apex than at the base, and by the large glyma at

the base of the first tergum.

The generic name refers to the similarity of this genus to Percnobracon\ the

gender is masculine.

Percnobraconoides jojoba Marsh, New Species

(Figs. 8, 11-13)

Female (Fig. 11).—Body length, 3-4 mm; ovipositor length, 1.5-2 mm. Color:

Head, basal % of antenna, and legs honey yellow; thorax and first abdominal

segment red-brown; apical x h of antenna and remainder of abdomen black; fore

wing banded. Head: Granulate-rugose, malar space strigate; antenna with 16-21

flagellomeres; eye large, malar space about l h eye height, temple about Vi eye

width; ocelli small, ocellar-ocular distance 2.5 times diameter of lateral ocellus.

Thorax: Mesonotum sharply declivous anteriorly; notauli absent, mesonotum

costate-granulate, scutellum granulate; propleuron costate-granulate; mesopleuron

finely granulate, stemaulus deeply sulcate and smooth, subalar groove rugose;

propodeum rugose dorsally and laterally, finely granulate on sides and on dorsal

basal areas on each side of median rugose area. Abdomen: First tergum slightly

longer than apical width, narrowed sharply at base, longitudinally costate-gran-

ulate with two conspicuous raised basal carinae and deep depression (glyma) at

each side; second tergum costate-granulate, remainder of terga granulate; ovi¬

positor a little longer than abdomen and curved upward. Legs: Fore tibia (Figs.

12, 13) with irregular rows of stout spines on anterior side, spines on outer side

normally sharply pointed, those on inner side bluntly pointed; hind coxa with

weak basal tubercle on inside. Wings (Fig. 8): Fore wing with two cubital cells,

radial cell short and wide, last segment of cubitus arcuate, intercubitus about as

long as recurrent vein, nervulus present and interstitial with basal vein, brachius

weak, first brachial cell open at apex; hind wing with radiella absent, cubitella

unpigmented and visible only in reflected light, nervellus present.

Male.— Essentially as in female; hind wing with large stigma at base.

Holotypefemale. — ARIZONA, Pima Co., Tuscon Mountains, Saguaro National

Monument West, Red Hill, 3 March 1980, J. Cicero collector, reared from dead

jojoba branches. Deposited in U.S. National Museum.

Paratypes. — 1 6, same data as holotype. 1 2, MEXICO, Guaymas, 10 April

1938, R. H. Crandall collector. Deposited in USNM and UCR.

The type specimens were reared from branches of jojoba but not associated

with any specific host insect. Frommer (per. comm.) mentions that a bostrichid,

Amphicerus cornutus (Pallas), was also collected from the same jojoba plants and

it could be the host of the wasp.

The specific name comes from the common name for the plant Simmondsia

chinensis (Link) Schneider from which the specimens were reared.

Xenosternum ornigis Muesebeck

(Fig. 10)

Locality.— CALIFORNIA, Riverside Co., Palm Desert.

Host.—Periploca sp.

This species was reported by Pinto and Frommer (1980) and Pinto et af (1987).

VOLUME 65, NUMBER 1

It was previously recorded from the southern plains states and has been reared

from the gelechiid Fascista cercerisella (Chambers).

Acknowledgments

M. J. Sharkey, Biosystematics Research Centre, Ottawa, Canada, provided the

identification for Bassus binominata. S. R. Shaw, Museum of Comparative Zo¬

ology, Harvard University, confirmed the identity of the new species of Ascogaster

and provided comments on the manuscript. S. Frommer, University of California,

Riverside, and A. L. Norrbom, Systematic Entomology Laboratory, USD A, also

read the manuscript and offered many helpful suggestions.

Literature Cited

Achterberg, K. van. 1982. Notes on some type-species described by Fabricius of the subfamilies

Braconinae, Rogadinae, Microgastrinae and Agathidinae (Hymenoptera: Braconidae). Entomol.

Berich., 42:133-139.

Marsh, P. M. 1979. Family Braconidae. Pp. 144-295 in K. V. Krombein et al. (eds.), Catalog of

Hymenoptera in America north of Mexico. Smithsonian Institution Press, Washington, D.C.

-, S. R. Shaw, and R. A. Wharton. 1987. An identification manual for the North American

genera of the family Braconidae (Hymenoptera). Mem. Entomol. Soc. Wash. 13, 98 pp.

Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae):

a phylogeny and reclassification of Microgastrinae. Mem. Entomol. Soc. Can. 115, 147 pp.

McComb, C. W. 1968. A revision of the Chelonus subgenus Microchelonus in North America north

of Mexico (Hymenoptera: Braconidae). Univ. Md. Agr. Exp. Sta. Bull. A-149, 148 pp.

Pinto, J. D., and S. I. Frommer. 1980. A survey of the arthropods on jojoba ( Simmondsia chinensis).

Environ. Entomol., 9:137-143.

-, and-. 1984. Laboratory and field observations on the life history of Epinotia kasloana

McDunnough (Lepidoptera: Tortricidae: Olethreutinae), a moth feeding on jojoba {Simmondsia

chinensis (Link) Schneider). Proc. Entomol. Soc. Wash., 86:199-209.

-,-, and S. A. Manweiler. 1987. The insects of jojoba, Simmondsia chinensis, in natural

stands and plantations in southwestern North America. Southwestern Entomol., 12:287-298.

Scarlett, P. L. 1978. Jojoba in a nutshell. The natural history, cultivation and market demand of

jojoba {Simmondsia chinensis ). Jojoba Int. Crop., Carpinteria, California, 55 pp.

Sharkey, M. J. 1985. Notes on the genera Bassus Fabricius and Agathis Latreille, with a description

of Bassus arthurellus n. sp. (Hymenoptera: Braconidae). Can. Entomol., 117:1497-1502.

Shaw, S. R. 1983. A taxonomic study of Nearctic Ascogaster and a description of a new genus

Leptodrepana (Hymenoptera: Braconidae). Entomography, 2:1-54.

Sherbrooke, W. C., and E. F. Haase. 1974. Jojoba: a wax-producing shrub of the Sonoran Desert.

Literature review and anotated bibliography. Arid Lands Res. Inf. Pap. No. 5, 141 pp.

Yermanos, D. M. 1979. Jojoba—a crop whose time has come. Calif. Agric. 33(7&8):4-7, 10-11.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 68-73

Neonemobius eurynotus (Rehn and Hebard)

(Grylloptera: Trigonidiidae: Nemobiinae), a Cricket

of the San Francisco Bay Area, California

Vernon R. Vickery and David B. Weissman

(VRV) Emeritus Curator, Lyman Entomological Museum and Research Lab¬

oratory, Macdonald College, McGill University, 21111 Lakeshore Rd., Ste. Anne

de Bellevue, Quebec H9X ICO, Canada; (DBW) Department of Entomology,

California Academy of Sciences, Golden Gate Park, San Francisco, California

94118.

Abstract. — The nemobiine cricket, Neonemobius eurynotus (Rehn and Hebard,

1918) is compared with other species of Neonemobius. This species has been

known only from the female holotype. The male sex is described.

Weissman and Rentz (1978) recorded and discussed the saltatorial orthopteroid

fauna of Jasper Ridge, a Biological Preserve of Stanford University, and neigh¬

boring Palo Alto, California. Included was a nemobiine cricket, Neonemobius

eurynotus (Rehn and Hebard, 1918). The species was described from a single

female from Berkeley and was not seen again until the report of Weissman and

Rentz (1978). The male has not been described.

Superficially the cricket looks like Modicogryllus but comparison of females

with the unique holotype (Academy of Natural Sciences, Philadelphia, type H472)

showed clearly that it is N. eurynotus. The ovipositor of the holotype is badly

worn so that the teeth on the dorsal valve are missing (Rehn and Hebard, 1918:

103, fig. 2). A normal ovipositor is shown in Figure 1. The dorsal aspect of the

holotype (Rehn and Hebard, 1918:103, fig. 1) shows all legs present; when ex¬

amined during the present study all right legs were missing (Fig. 2b). In all other

respects the Palo Alto specimens and the holotype are almost identical.

Rehn and Hebard (1918) considered its closest relative to be Brachynemobius

panteli Hebard, a Mexican species, but that more specimens would be required

to determine whether eurynotus should be assigned to a different genus. It resem-

Figure 1

Neonemobius eurynotus, ovipositor.

VOLUME 65, NUMBER 1

Figure 2. Photographs, Neonemobius eurynotus. a. Male, Palo Alto, California, dorsal, b. Female

holotype, Berkeley, California, dorsal, c. Male, lateral.

bles B. panteli in some respects but differs in others and is here placed in the

genus Neonemobius Hebard, 1913.

The male genitalia (Fig. 3a-c), the tibial spines and the distal spurs of the hind

tibia (Fig. 4a, b) are similar to those of other species of Neonemobius. The chro-

PAN-PACIFIC ENTOMOLOGIST

Figure 3. Male genitalia, a. Dorsal, b. Lateral, c. Ventral.

mosome number (2N 8 = 19) (Fig. 5) is the same as in the other species of

Neonemobius described by Lim (1971). The broad robust body is similar to that

of N. mormonius (Scudder) but is different from some slender species, such as N.

palustris (Blatchley). The ovipositor is short and heavy, similar to, but shorter

than, that of N. mormonius.

Neonemobius eurynotus is more robust than other species in the genus (Fig.

2a-c) with the possible exception of N. toltecus (Saussure). Total body length is

greater, male maximum is 11.9 mm and N. mormonius is next at 7.5 mm. Max¬

imum total length, female is 12.1 mm and N. toltecus is next largest at 10.0 mm

(Hebard, 1913). The pronotal length and length of the hind femur of both sexes

of N. eurynotus are greater than in all other species of the genus, with the exception

of the measurements for N. toltecus given by Hebard (1913). The tegmina and

ovipositors are shorter than in any of the other species.

Rehn and Hebard (1918) gave measurements of the holotype. All but one of

the measurements fall within the range of the sample of 10 females measured

(Table 1). The ovipositor length was given as 2.35 mm, while the present sample

ranges from 1.8 to 2.2 mm. Other than this discrepancy, which may be due to

method of measurement, the original description allows accurate description of

the female sex of the species.

Description. —Relatively large, broad species (Fig. 2a, c), proportion of width

Table 1. Specimen measurements (mm).

Males (n — 10)

Females (n = 10)

Holotype 1

Body length

11.0 (9.9-11.9)

11.1 (10.1-12.1)

10.0

Head length

2.0 (1.7-2.2)

2.0 (1.8-2.1)

—

Head width

3.0 (2.7-3.2)

2.9 (2.6-3.1)

3.1

Pronotum length

1.8 (1.7-1.9)

1.8 (1.7-2.1)

1.95

Pronotum width

2.8 (2.6-3.1)

2.7 (2.5-3.0)

3.0

H. femur length

5.2 (5.0-5.4)

5.1 (4.6-5.6)

5.5

H. tibia length

3.9 (3.6-4.4)

3.8 (3.1-4.2)

4.2

Tegmen length

2.8 (2.5-2.9)

2.1 (1.9-2.4)

2.15

Ovipositor length

2.0 (1.8-2.2)

2.35

1 Measurements of female holotype from Rehn and Hebard (1918).

VOLUME 65, NUMBER 1

Figure 4. Hind tibia, a. Tibial spines, b. Distal spurs.

to length (Table 1) greater than for other North American nemobiine genera. Head

large, rounded, distinctly broader than pronotum. Pronotal width slightly less

than twice the length; pronotum rounded; anterior and posterior widths about

equal, sides subparallel. Tegmina short, less than one and one-half times longer

than pronotum. Wings reduced to very small lateral pads. Eyes small in dorsal

aspect, not protruding; antennae very long, about one and one-half times body

length, mostly pale, darker apically; distoventral spurs of hind tibiae typical of

the genus Neonemobius, unequal in length, the interior spur longer (Fig. 4b).

PAN-PACIFIC ENTOMOLOGIST

Description of male. — Characteristics of male stridulatory file as in Table 2.

Hind tibiae with four internal and three external spines plus the glandular spine,

this conical and nearly black (Fig. 4a).

Male genitalia (Fig. 3a-c) subquadrate, tapering on distal two-thirds; anterior

margin of dorsal epiphallic plate with relatively shallow median U-emargination

(shallower than in most nemobiines); transverse epiphallic sulcus anterior in po¬

sition; distal part of dorsal epiphallic plate long and flat with deep distal median

V-emargination; rami long and slender with terminal processes broad, ventral

processes blunt; mesal lobes curved posteriorly.

Color: Front of face dark brown beneath eyes and extending as a dark triangle

dorsally between eyes to include at the apex the dorsal ocellus; area above antennae

to middle of eyes pale; dorsum of head brown with very small darker spots and

pale blotches, cut by four longitudinal paler lines extending to the occiput, the

outer two lines beginning at the inner margins of the compound eyes; labial palps

pale; maxillary palps generally pale with apical half of apical segment infumate,

brownish.

Pronotum pale brown, with two narrow triangular darker areas, with apices

directed laterally, and two raised, darker, brownish areas posterior to the triangles.

Wings hyaline. Legs pale brown with long, thin black spines, except those of hind

tibiae which are pale, robust and very long in both sexes, not short and heavy as

stated by Rehn and Hebard (1918).

Table 2. Characteristics of male stridulatory file (n = 8).

No. of teeth

Length of file (mm)

No. of teeth/mm

160.25

0.99

163.8

(148-175)

(0.80-1.15)

(143-200)

VOLUME 65, NUMBER 1

Abdomen medium brown above with two longitudinal broad pale stripes lat¬

erally on each segment but these are much less distinct than in females; cerci pale

brown.

Cytology

The cytology of two male crickets from Palo Alto, Santa Clara County, was

examined. Diakinesis-metaphase I preparations show a total of 18 autosomes, all

of which appear to be telocentric, plus an XO/XX sex chromosome system, which

gives 2N <3=19 and 2N $ = 20 elements (Fig. 5) as in other species of Neonemobius

(Lim, 1971). The X is a rather large chromosome. One of the autosomal elements

(No. 9 in Fig. 5) is much smaller than the others and separates earlier. Most

bivalents form only a single chiasma, frequently interstitially or distally located.

Desutter (1987) has clarified the general classification of the Grylloidea. We

agree with her in placing the Nemobiinae as a subfamily of the Trigonidiidae

rather than of the Gryllidae.

Acknowledgments

The specimens will be returned to the California Academy of Sciences, San

Francisco. The first author owes thanks to the late Dr. H. R. Roberts, Academy

of Natural Sciences, Philadelphia for the loan of the holotype of Neonemobius

eurynotus. Thanks also to Dr. P. G. Fontana for the photographs of the chro¬

mosomes and to Mr. Pierre Langlois for photographs of specimens.

Literature Cited

Desutter, L. 1987. Structure et evolution du complexe phallique des Gryllidea ( Orthopteres ) et

classification des genres Neotropicaux de Grylloidea. Premiere Partie. Ann. Soc. Entomol. Fr.

(N.S.), 23(3):213-239.

Hebard, M. 1913. A revision of the genus Nemobius (Orthoptera: Gryllidae) found in North America

north of the Isthmus of Panama. Proc. Acad. Nat. Sci. Philad., 65:394-492.

Lim, H.-C. 1971. Note on the chromosomes of the Nemobiinae (Orthoptera: Gryllidae). Can. J.

Zool., 49:391-395.

Rehn, J. A. G., and M. Hebard. 1918. A new species of the genus Nemobius from California

(Orthoptera: Gryllidae: Gryllinae). Ent. News, 29:102-105.

Weissman, D. B., and D. C. F. Rentz. 1978. The Orthoptera of Stanford University’s Jasper Ridge

and neighboring Palo Alto, California. Wasmann J. Biol., 35(1977):87—114.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 74-76

Overwintering of Phryganidia californica in the

Oregon Cascades and Notes on its Parasitoids

(Lepidoptera: Dioptidae)

David Carmean, Jeffrey C. Miller, and Brian Scaccia

Department of Entomology, Oregon State University, Corvallis, Oregon 97331.

Abstract. —A population of California oakworm ( Phryganidia californica Pack¬

ard) was observed for 3 yr in the Oregon Cascades. The site was 150 km inland

from the coast, 44°18'N and at an elevation of 750 m. The only parasitoids reared

were Ceranthia sp., Hyphantrophaga virilis, both tachinids, and an ichneumonid,

Mesochorus sp.

The California oakworm (Phryganidia californica Packard) overwinters as a

larva feeding on the foliage of plants in the oak family (Fagaceae). Thus, its

northern distribution may be strongly influenced by the severity of winter con¬

ditions and the availability of a host with persistent (evergreen) foliage. Popula¬

tions of the California oakworm have been reported to overwinter in California

only as far north as southern Mendocino County (about 39°N) and there only in

the areas with a maritime climate (Miller, 1987). Harville (1955) found small

outbreaks outside the overwintering range of the California oakworm and sug¬

gested that vehicles or wind might carry the caterpillars or moths long distances.

In Oregon, the California oakworm has been reported from golden chinkapin,

Castanopsis chrysophylla (Dougl.) (Fumiss and Carolin, 1977). Wickman and

Kline (1985) reported an outbreak that affected many acres of golden chinkapin

in the southwestern Willamette Valley (elev. 150-230 m, 44°6 , N). This population

was observed in November 1971. No subsequent observations were reported, and

they did not comment on overwintering.

Here we report on a population of the California oakworm found overwintering

at 750 m elevation on the west side of the Cascade Mountains.

On 12 August 1985 we found one California oakworm larva on Oregon white

oak ( Quercus garryana Dougl.) in the H. J. Andrews Experimental Forest, Lane

County, Oregon (44°13'N). On 22 August 1985 we found first and second instars

to be very abundant on several golden chinkapins 1.7 km NE of the Oregon white

oak site. This second site was on a SSE-facing slope, 2.2 km NE of the headquarters

of the H. J. Andrews Experimental Forest. We returned to this area on 30 August

and 27 September 1985, collecting many (n > 200) caterpillars each time. Late

instars, indicating overwintering, were also found at this site on 30 April 1986.

Additional larvae were collected here on 19 June 1986, 18 November 1987, and

5 May 1988.

The cold-hardiness of this population merits further study. Sibray (1947) tested

the affect of temperature on a Berkeley population of California oakworm. At

10°C he found an average larval period of 178 days. At 4.4°C there was limited

VOLUME 65, NUMBER 1

development and death after 3 mo. At 1.7°C there was no feeding and negligible

activity. There was no hatch of eggs at 1.7°C and 4.4°C. In the H. J. Andrews

Experimental Forest, at 640 m elevation and 1.8 km SWW of our sample site,

temperatures below freezing occurred each month from November 1985 to April

1986, and from November 1986 to April 1987. The coldest months were No¬

vember 1985 (mean minimum air temperature — 1.6°C with an absolute minimum

of — 9.2°C) and January 1987 (mean minimum air temperature of — 1.4°C with

a minimum temperature of — 9.2°C).

We searched for the California oakworm in other areas of the H. J. Andrews

Experimental Forest and western Oregon. Our only find was a single first instar

in SW Oregon (elev. 250 m, 42°44'N, Daphne Grove Campground, Coos County)

on tanoak, Lithocarpus densiflorus (Hook and Am.) Rehd., on 27 July 1986.

Field-collected larvae were reared to document the occurrence of parasitoids.

The parasitoid species observed were: Ceranthia sp. (Tachinidae); Hyphan-

trophaga virilis (Aldrich and Webber) (Tachinidae); and Mesochorus sp. (Ich-

neumonidae). One pupa was found with an emergence hole of an unidentified

hymenopteron parasitoid. From the collections in 1985 we reared over 200 field-

collected larvae and only 14 were parasitized, all of these by Ceranthia sp. In

addition, one of the Ceranthia sp. was parasitized by Mesochorus sp. Parasitoids

emerged from 7 of 26 larvae collected on 30 April 1986 and 19 June 1986. Four

of these larvae were parasitized by Ceranthia sp. and three by H. virilis.

Ceranthia sp. is a gregarious, larval, endoparasitoid. Usually one but up to

three parasitoids occurred per host. The parasitoid emerged from the last and

next to last instars. Parasitism was as high as 23.5% (n = 17, 30 April 1986).

In our study H. virilis acted as a larval-pupal parasitoid. They emerged from

the pupae of field collected host larvae. Previously H. virilis has been reported to

emerge from California oakworm larvae (Young, 1977). Also, H. virilis is recorded

as a larval parasitoid of various Lepidoptera (Amaud, 1978).

Young (1977) found only 2 (0.17%) of 1150 larvae parasitized by H. virilis

while Harville (1955) found 6 H. virilis from “several thousands of Phryganidia

larvae.” Horn (1974) studied parasitism from 1969 to 1972 and found 453 of

798 field-collected pupae parasitized by various Hymenoptera but did not rear

any tachinid parasitoids.

Voucher specimens have been placed in the Systematic Entomology Laboratory

at Oregon State University.

Acknowledgments

Our thanks to Paul H. Arnaud, Jr., California Academy of Sciences, for deter¬

mining the Tachinidae, Frederick Bierlmaier, H. J. Andrews Experimental Forest,

for weather data, and John D. Lattin, Oregon State University, for contributing

support to the study. Oregon St. Univ. Agric. Exp. Stn. Tech. Paper No. 8767.

Literature Cited

Amaud, P. H., Jr. 1978. A host-parasite catalog of North American Tachinidae (Diptera). U.S.D.A.

Misc. Publ. No. 1319, 860 pp.

Fumiss, R. L., and V. M. Carolin. 1977. Western Forest Insects. U.S.D.A. Forest Service Misc.

Publ. No. 1339, 654 pp.

Harville, J. P. 1955. Ecology and population dynamics of the California Oak Moth Phryganidia

californica Packard (Lepidoptera: Dioptidae). Microentomology, 20:83-166.

PAN-PACIFIC ENTOMOLOGIST

Horn, D. J. 1974. Observations on primary and secondary parasitoids of California Oakworm,

Phryganidia californica, pupae. Pan-Pac. Entomol., 50:53-59.

Miller, J. S. 1987. A revision of the genus Phryganidia Packard, with description of a new species

(Lepidoptera: Dioptidae). Proc. Entomol. Soc. Wash., 89:303-321.

Sibray, W. S. 1947. Bionomics of the California Oak Moth. M.S. thesis, Univ. California, Berkeley,

80 pp.

Wickman, B. E., and L. N. Kline. 1985. New Pacific Northwest records for the California Oakworm

Phryganidia californica. Pan-Pac. Entomol., 61:152.

Young, L. C. 1977. Pupal parasites of the California Oak Moth, Phryganidia californica Packard: a

biological and ecological study. Ph.D. thesis, Univ. California, Berkeley, 126 pp.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 77-78

Scientific Note

First Record of a Dryopid (Coleoptera: Dryopidae) from

Washington and Notes on the Distribution of the

Family in the Northwestern United States

Beginning in early 1986 we began an extensive survey of certain groups of

aquatic Coleoptera and Hemiptera occurring in the Pacific Northwest. Among

the more interesting records are the first collection of a dryopid beetle, Helichus

striatus LeConte, from Washington and also the finding of it in northern Idaho.

Since recordings of dryopid beetles from the northwestern United States are rare

and scattered, we felt this an opportune time to present those records with which

we are familiar.

Hatch (1965, The beetles of the Pacific Northwest, Part IV, Univ. Wash. Press)

recorded Helichus striatus from southwestern British Columbia, southeastern Ida¬

ho, and Oregon. Brown (1976, USEPA, Water Pollution Control Research Series

18050 ELD04/72) recorded H. striatus striatus as occurring across the northern

United States and southern Canada, and H. striatus foveatus LeConte as occurring

along the west coast of the United States north into British Columbia. Again,

Brown (1983, A catalog of the Coleoptera of America north of Mexico, USDA

Agr. Hdbk. 529-49) records both H. striatus striatus and H. striatus foveatus from

British Columbia, Washington, and Oregon. However, Brown had not seen spec¬

imens from Washington or Idaho when compiling these reports. It appears that

most of the records concerning the distribution of Helichus in the northwestern

United States were based on what might occur in the region as opposed to what

had been collected.

In Washington, specimens of Helichus striatus striatus were collected at Douglas

County, East Foster Creek, ca. 13 km SE of Bridgeport (R26E T28N Sec. 3), at

an elevation of 570 m on 31 May 1988. East Foster Creek is perennial for a length

of ca. 20 km and intermittent for another 20-30 km. The stream is fed by a series

of springs and run-off, arising mainly from the area of Foster Coulee. The creek

empties into the Columbia River near Bridgeport. The steppe region vegetation

which dominates the area was classified as an Artemisia tridentata NutUFestuca

idahoensis Elm. association (Daubenmire, 1970, Steppe vegetation of Washing¬

ton, Wash. Agric. Exp. Stn. Tech. Bull. 62). At the collecting site the stream was

ca. 1-2 m wide with areas of short riffles and numerous, larger pools. Water flow,

for this time of year, was unusually slow due to persistent drought conditions

over the past year. Helichus striatus was collected only from areas along the stream

bank where current had undercut the shoreline leaving an overhang (ca. 20-40

cm deep) of soil and vegetation. Kicking-up into this overhang released the beetles

which were then carried by the current into a net held downstream. No dryopids

were taken from the riffle areas. Thirty-four adult specimens were collected from

an overhang area ca. 15 m long. Areas within 300 m up or down stream were

mostly small pools in which no dryopids were found. In addition to H. striatus,

33 specimens of Agabus seriatus Say (Coleoptera: Dytiscidae) were collected at

the overhang site.

In addition to the Washington record we have taken Helichus striatus from two

PAN-PACIFIC ENTOMOLOGIST

PACIFIC NORTHWEST

Figure 1. Collection sites of Helichus striatus in the northwestern United States. A total of 80

specimens were examined.

locations, 4 mi apart, along Deep Creek in Boundary Co., Idaho. And, although

not a record for the state, this find represents a significant northern extension of

the range of the beetle within Idaho. At the Deep Creek locations, the beetles

were collected from inundated grasses along the shore.

The distribution of Helichus striatus in the northwestern United States is pre¬

sented in Figure 1. Specimens examined are from the following counties: IDAHO:

Bannock, Blaine, Boise, Boundary, Lemhi, Madison, Owyhee, Twin Falls, Valley,

and Washington; OREGON: Baker, Crook, Grant, Harney, Lake, and Linn;

WASHINGTON: Douglas.

Acknowledgments.— We thank William H. Clark, Rock Creek Rural Clean Water

Program and Orma J. Smith Museum of Natural History, The College of Idaho;

David Kavanaugh, California Academy of Sciences; James DiGiulio, Oregon State

University; Frank Merikel, University of Idaho; and Richard Haswell for the loan

of specimens. This work was conducted under project 9043, College of Agriculture

and Home Economics, Washington State University, Pullman, Washington.

Richard S. Zack, James Entomological Collection, Washington State University,

Pullman, Washington 99164-6432 and Harley P. Brown, Department of Zoology,

The University of Oklahoma, Norman, Oklahoma 73019.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 79-88

Additional Range Extension by the German Yellowjacket,

Paravespula germanica (Fabricius), in North America

(Hymenoptera: Vespidae ) 1

Roger D. Akre, Carol Ramsay, Al Grable,

Craig Baird, and Alan Stanford

(RDA, CR) Department of Entomology, Washington State University, Pull¬

man, Washington 99164-6432; (AG) Department of Biology, Walla Walla College,

College Place, Washington 99324; (CB) Cooperative Extension, University of

Idaho, Parma, Idaho 83660; (AS) Idaho State Department of Agriculture, 421 W.

Sherman Ave., Nampa, Idaho 82627.

The German yellowjacket, Paravespula germanica (Fab.), has become estab¬

lished in many temperate areas of the world (Edwards, 1976; Brown, 1979; Smith-

ers and Holloway, 1977,1978; Olafsson, 1979; Chiappaetal., 1986; Magunacelaya

et al., 1986a, 1986b), and it continues to become established in new areas as it

moves west across North America (MacDonald et al., 1980; MacDonald and

McDonald (sic), 1980; Dunn, 1980a, 1980b; Stanford, 1984). These latter reports

showed emigrations of German yellowjacket into the West. In the Pacific North¬

west it has become established in southern Idaho around Nampa, and is present

in the Seattle, Washington area. It was also reported in Minneapolis/St. Paul,

Minnesota and Winnipeg, Canada. In addition, these yellowjackets have invaded

California. In 1985 these wasps were reported as nesting in the San Francisco Bay

area with at least 3 suspected colonies (Gambino, 1987).

The German yellowjacket is noteworthy because it has a propensity to nest in

man-made structures in North America. Also, colonies are frequently large and

usually persist late into the fall. These characteristics bring these yellowjackets

into close contact with humans that often result in stinging episodes. German

yellowjackets also have a tendency to become perennial, or at least to continue

into a second year in warmer climates. Some colonies become huge with over a

million cells in some nests in Tasmania, Australia (Spradbery, 1973; see also

Edwards, 1980). This yellowjacket has apparently become the dominant species

in some localities it has invaded (Stanford, 1984), and workers are very aggressive

scavengers. The wasp is always perceived as a serious public health threat.

The purpose of this paper is to report additional range extensions of the German

yellowjacket in North America, and make comparisons of biology and behavior

with that of Paravespula pensylvanica (Saussure), the western yellowjacket.

Previous Distribution in North America

We last reported P. germanica nests occurring in Nampa, Idaho in 1981 and

workers in Puyallup, Washington in 1982 (MacDonald and Akre, 1984). The

1 This work was conducted under project 0037, Washington State University, College of Agriculture

and Home Economics.

PAN-PACIFIC ENTOMOLOGIST

western-most records of this wasp, into the Great Plains of North America, were

at Minneapolis and Winnipeg. No colonies have been reported from anywhere

in the South.

New Distribution

It was inevitable that the German yellowjacket continue its spread westward.

It continues to be the dominant yellowjacket in southern Idaho, and it is becoming

more abundant in western Washington each year. Significant new distribution

records (Fig. 1) of German yellowjacket presence and/or establishment, previously

unreported, are listed briefly below:

Pacific Northwest

The first German yellowjacket nest found in Washington was located 4 August

1983 in a Puyallup home. The colony was killed but the nest was not accessible

for collection.

In July 1984, the first workers were netted in British Columbia, Canada at

Cloverdale (80 km W of Vancouver).

From August to December 1985, German yellowjacket nests were collected

from houses in Seattle, Tacoma, and Chehalis, Washington.

In fall 1986, a colony with a nest of 91 cm x 46 cm was killed in a home in

Clearbrook, British Columbia (near Vancouver).

In August and September 1986, the first workers were collected in Ontario and

Nyssa, Oregon.

In October 1986, the first nest of the German yellowjacket was collected in

Walla Walla, eastern Washington, located in a rotten stump of a willow tree.

In September 1987, the first nest was discovered in Oregon, in a house in Milton-

Freewater.

* On 2 October 1988 a nest was collected in Prasser, Washington, and on 2

November 1988 a nest was removed from a house in Yakima, Washington.

* As of November 1988 the wasp was established from Vancouver to Rosedale

in the Skagit Valley.

Other Significant Occurrences in North America

German yellowjackets were first collected in St. Louis, Missouri in 1974, and

they are now abundant in the St. Louis area (Hunt, 1988). They have not been

collected from other areas.

In 1985 we received specimens of P. germanica workers from personnel at El

Dorado County Vector Control, Department of Agriculture, that had been col¬

lected in 1983 from South Lake Tahoe, California. Perhaps this was an isolated

incident as no additional wasp problems have occurred in this locality.

We found an active colony 7 December 1987, 14.5 km E of East Quincy,

California (elev. ca. 1100 m), in the base of a tree stump. The temperature was

about 4°C and snow was lying in patches on the ground, but workers were flying.

During the summer of 1987 the Parks and Recreation Department of Saskatoon,

Saskatchewan received numerous complaints about yellowjackets, mainly due to

German yellowjackets. Personnel located 6 nests, but all were in homes and the

colonies were destroyed. No nests were collected for analysis. However, a sample

* Added at proof.

VOLUME 65, NUMBER 1

Figure 1. Known distribution oiParavespula germanica in the Pacific Northwest, with the interstate

corridors noted by a dashed line.

of workers and males was sent to us in March 1987 for species verification. The

presence of males indicated colonies were present in the city and reproducing.

The Parks Department will expand a surveillance and monitoring project for this

wasp in 1988 and 1989.

Yellowjacket Trapping

Trapping in Western Washington

Personnel at the State Department of Agriculture became concerned about the

distribution of German yellowjacket in Washington during 1985, and during the

summer of 1986 a trapping network was established to determine areas in western

Washington where this yellowjacket was present.

Two hundred Seabright Yellowjacket Traps were deployed along Interstate 5

from the Oregon to Canadian borders. The traps were baited with either ham or

pork sausage. All traps were checked and rebaited every 3 working days. Traps

were monitored from 15 August through 21 October. All captured yellowjackets

were placed in alcohol for subsequent determination.

The survey showed that German yellowjackets were present along the entire

corridor (Fig. 1). However, apparently they were present only in low numbers

since a total of only 233 workers was captured. The same traps caught 1582

Paravespula vulgaris (L.) and 1707 P. pensylvanica workers. Of course, it is pos¬

sible that the baits or the traps were less effective at attracting or capturing P.

germanica workers.

Trapping in Southern Idaho

As mentioned, the first German yellowjacket nest was collected in Idaho on 24

October 1981, and collections since this time indicated an ever increasing distri¬

bution (Stanford, 1984). Because of these discoveries, efforts were made in 1985

PAN-PACIFIC ENTOMOLOGIST

Table 1. Sites and nest sizes of analyzed Paravespula germanica nests in Washington, Oregon, and

Idaho.

Location

Site

Number of cells

Bellingham, WA

attic

12,579

Renton, WA

ceiling

5251

Auburn, WA

ground

4186

Tacoma, WA

wall

4370

Pacific, WA

ceiling

1518

Puyallup, WA

eaves

13,748

Longview, WA

wall

3953

Longview, WA

ground

4955

College Place, WA

floor

4744

College Place, WA

wall

3153

College Place, WA

wall

10,461

College Place, WA

ground

6431

College Place, WA

crawl space

3980

Walla Walla, WA

basement

12,948

Milton-Freewater, OR

wall

13,654

Milton-Freewater, OR

attic

6353

Caldwell, ID

out building

7309

Nampa, ID

unknown

2078

Nampa, ID

crawl space

10,916

and 1986 to trap or net yellowjackets around Nampa to determine possible spread.

Trapping was conducted with standard heptyl butyrate traps but with ginger ale

as the attractant as meat baits were apparently not attractive. Netting was done

by hand with an aerial net. Traps were placed at 13 sites in 1985 and 9 sites in

1986 on the perimeter of Nampa and Caldwell, Idaho, and along the Interstate

84 corridor from Boise, Idaho to Ontario, Oregon.

Trapping showed German yellowjackets were extremely common in Nampa,

and are the dominant yellowjacket. However, the survey also showed that these

wasps are present in Caldwell, Greenleaf, Wilder, Fruitland, and Parma, Idaho,

and in both Nyssa and Ontario, Oregon. Essentially they are present along the

entire corridor from Boise to just across the Oregon border. However, in 1985

the numbers trapped and netted were low, with 17 females trapped; 6 females

netted. In 1986 low numbers were found again, with 34 females, 3 males trapped;

17 females netted. The period of collection during 1985 was 29 August to 8

November; during 1986-1987 this period was 22 August to 9 January.

Nests

While many reports of German yellowjacket were submitted by county agents

or provincial entomologists from western Washington and Canada, we have been

able to obtain only a few nests for analysis. Usually colonies were killed and the

nests destroyed. However, we collected or received information on 38 nests; 31

of these were built inside structures (Table 1). The other 7 nests were in the soil

or stumps.

A comparison was made between German yellowjacket and western yellowjack¬

et nests collected on about the same Julian dates but in different years. These data

show German yellowjacket colonies are, in general, larger (Fig. 2). The analyses

also showed that colonies of this species tend to persist later into the year (Fig.

VOLUME 65, NUMBER 1

U-

15000 -

12500 -

10000 -

Paravespula pensylvanica

A - August

S - September

O - October

N - November

DATES

7500 -

J1 7 A7 A1 7 A22 A22 A24 A29 S3 S3 S11 S18 S25 03 09 016 027 N8

DATES

Figure 2. Comparison of nest size measured by cell numbers of Paravespula germanica and P.

pensylvanica. P. pensylvanica data are from MacDonald et al. (1974).

3; Tables 2, 3). For example, presence of numerous empty cells or cells with

multiple eggs (indicative of lack of queen control and the presence of egg laying

workers, respectively) in nests of P. germanica occurred later into the year than

in nests of P. pensylvanica (Tables 2, 3). Also in many cases the P. germanica

foundress queen was still alive and functional later into the year (Table 2).

A serious problem in comparing nests of these two species is that all western

yellowjacket nests were collected in the field under unaltered conditions. This is

reflected in the relatively smooth increase in the size of the colonies (Fig. 2). Many

German yellowjacket nests were collected from situations in which colony history

was unknown. The data on colony size suggested that several colonies were treated

with insecticides early in the development of the colony as the nests are unusually

small for the dates of collection (Fig. 1, Table 2). All data on these nests should

be considered with this potential problem in mind.

Similarity in colony sizes exists between P. germanica in North America and

PAN-PACIFIC ENTOMOLOGIST

J1 7 A7 A17 A22 A22 A24 A29 S3 S3 S11 S18 S25 03 09 01 6 027 N8

DATES

Figure 3. Comparison of number of empty cells in nests of Paravespula germanica and P. pen¬

sylvanica. P. pensylvanica data are from MacDonald et al. (1974).

Europe. Spradbery (1971) found a maximum of 12,000 cells per nest in England

and Vasic (1968) cited a nest analyzed by Grozdanic (1965) with 12,632 cells per

nest in Belgrade. This compares favorably to nests collected in Washington, 13,748

cells, Idaho, 10,916 cells, and Oregon, 13,654 cells. North American and European

colonies are very small in comparison with those in the temperate maritime

climatic regions where some nests contain more than a million cells (Spradbery,

1973). Nesting behavior of North American and European colonies also appears

to be similar with nests occurring in structures and in the soil. However more

colonies nest in structures in North America.

Seasonal Activity

The seasonal activity of P. germanica and P. pensylvanica differs only in du¬

ration. Queens become active in the spring from early April to May, depending

on weather, and forage for food and seek nest sites until late June. The first workers

Table 2. Analyses of 19 nests of Paravespula germanica from Washington, Idaho, and Oregon.

Location

Coll, date

Queen

Combs

Cells

Eggs

Larvae

Pupae

Empty cells (%)

Longview

17 July 1987

3953

303

799

2397

454 (11.5)

Auburn

4 Aug. 1985

4186

268

2057

1851 (44.2)

Tacoma

21 Aug. 1985

4370

522

662

1860

1326 (30.3)

Coll. Place 1

21 Aug. 1987

yes

4744

272

866

2548

1058 (22.3)

Pacific

22 Aug. 1985

1518

233

415

721

149 (9.8)

Coll. Place

23 Aug. 1987

yes

3153

725

558

1701

169 (5.4)

Coll. Place 1

23 Aug. 1987

yes

10,461

1535

3256

4856

814(7.8)

Longview 1

1 Sept. 1987

yes

4955

587

1257

1891

1220 (24.6)

Renton 1

2 Sept. 1985

5251

762

1632

1957

900(17.1)

Puyallup 1

12 Sept. 1985

13,748

752 2

2068

3091

7027 (51.1)

Coll. Place 1

16 Sept. 1987

6431

482

1911

3075

963 (15.0)

Milton-Freewater 1

22 Sept. 1987

13,654

0 3

656

279

12,719 (93.2)

Coll. Place 1

13 Oct. 1987

yes

3980

508

1516

1730

226 (5.7)

Milton-Freewater 1

14 Oct. 1987

yes

6353

544

2740

630

2439 (38.4)

Caldwell 1

15 Oct. 1986

7309

260

2109

1769

3171 (43.4)

Nampa 1

3 Nov. 1983

2078

31 2

208

324

1515 (72.9)

Nampa 1

19 Nov. 1983

10,916

—

— —

Walla Walla 1

19 Nov. 1987

12,948

274

2142

340

10,192 (78.7)

Bellingham 1

7 Dec. 1987

12,579

12,579 (100)

1 Colonies with reproductive cells.

2 Colony with >4% multiple eggs per cell (all others < 1%), suggesting queen had died.

3 Colony collected 1 mo after killed, eggs probably too dry to identify.

— No data taken.

VOLUME 65, NUMBER 1

PAN-PACIFIC ENTOMOLOGIST

Table 3. Analyses of 17 nests of Paravespula pensylvanica from Washington (MacDonald et al.,

1974).

Location

Coll, date 1

Combs

Cells

Eggs

Larvae

Pupae

Empty cells (%)

Pullman

17 July 1973

563

188

179

17 (30.0)

Pullman

7 Aug. 1973

3215

694

1373

1135

13(0.4)

Pullman

17 Aug. 1973

2230

439

952

827

12 (0.5)

Pullman

22 Aug. 1973

3801

725

1509

1537

30 (0.8)

Pullman

22 Aug. 1973

1147

250

499

369

29 (2.5)

Pullman 2

24 Aug. 1973

4935

661

1842

1790

642(13.0)

Pullman 2

29 Aug. 1973

4866

747

2014

1674

431 (8.9)

Pullman 2

3 Sept. 1973

3002

625

1324

1006

47 (1.6)

Pullman 2

3 Sept. 1973

4275

712

1846

1406

311 (7.3)

Pullman 2

11 Sept. 1973

4291

649

1688

976

978 (22.8)

Pullman 2

18 Sept. 1973

6489

741

2741

1394

1613 (24.9)

Pullman 2

25 Sept. 1973

4447

460

2004

1097

886 (19.9)

Pullman 2

3 Oct. 1973

5159

5095 (98.8)

Pullman 2

9 Oct. 1973

3869

3829 (99.0)

Pullman 2

16 Oct. 1973

4499

4489 (99.8)

Pullman 2

27 Oct. 1972

5891

5844 (99.2)

Pullman 2

8 Nov. 1972 1

6128

6128 (100)

1 Only one colony available in November and no colonies were available in December.

2 Colonies with reproductive cells.

of the western yellowjacket appear about 10 June in Pullman, Washington, while

the earliest records of German yellowjacket workers indicate this species may be

a bit later, in mid-late July. However, it is more probable that this late appearance

of workers is simply due to a lack of collection records early in the season when

workers are few in number. Colony longevity, however, seems to differ markedly

as we received a few reports of German yellowjacket workers still very active and

flying late in the fall and winter, with one case in Marysville, Washington with

workers still active into February. This colony was in the wall of an unheated

garage. In addition, two colonies were still active in Eagle, Idaho, 22 km NE of

Nampa, in January 1987. One colony was in the attic of a house and the other

in the wall of a bam. The maximum life span of western yellowjacket colonies is

7 mo, compared to the German yellowjacket which survives up to 11 mo.

Males and new queens of P. pensylvanica are active late September-October,

infrequently into early November. Queens of P. germanica are active from Oc¬

tober into early January.

Summary and Discussion

Analyses of nests of the German yellowjacket show colonies are similar to those

found in Europe, but are considerably larger than typical western yellowjacket

colonies, and they persist later into the year. Since food resources become sparse

later in the year, there is a tendency for these wasps to be in contact with humans

for a longer period of time. Their propensity for nesting in structures also adds

to greater human contact.

We anticipate the German yellowjacket will continue to expand its range into

nearly all of western North America, and will become the dominant, scavenging

yellowjacket in many of these areas. No information was available about German

VOLUME 65, NUMBER 1

yellowjacket in eastern or northern Oregon. However, it is assumed that they are

present. Problems arise in determining range extensions when similar species are

present, because many people do not realize that a new species has become

established. Although interactions between P. germanica and P. pensylvanica are

essentially unknown, within 2-5 yr of becoming established in an area, P. ger¬

manica apparently out-competes western yellowjackets to become dominant in

numbers of workers and colonies. Also, the mild climate in parts of the Pacific

Northwest might be amenable to colonies surviving more than one season. How¬

ever, while we have received reports of at least 3 large nests that reportedly lasted

more than a year, we have been unable to verify these accounts.

The sympatric existence of these two species in many localities of North Amer¬

ica presents a unique opportunity to study the behavioral interactions of two very

closely related species. It would not be at all surprising if they would eventually

mate to produce a hybrid. However, in most areas mating is most likely improb¬

able since male and queen production by the German yellowjacket is later in the

year. Conversely, there is some overlap in reproductive production of colonies,

and eventually hybridization is a possibility.

Acknowledgments

We are indebted to a number of people who collected information, specimens,

and nests of German yellowjackets for the project. We sincerely thank Art An-

tonelli, Sharon Collman, Henry Gerber, Lyle Klostermeyer, Eric LaGasa, Carl

Roush, and Robert Stidham for their efforts. Richard Zack and Dan Suomi are

thanked for critically reading the manuscript and for making suggestions for

improvement.

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-, and-. 1978. Establishment of Vespula germanica (Fabricius) (Hymenoptera: Vespidae)

in New South Wales. Aust. Entomol. Mag., 5(3):5 5-60.

Spradbery, J. P. 1971. Seasonal changes in the population structure of wasp colonies (Hymenoptera:

Vespidae). J. Anim. Ecol., 40:501-523.

-. 1973. The European social wasp Paravespula germanica (F.) (Hymenoptera: Vespidae) in

Tasmania, Australia. Int. Union Study Soc. Insects VII Int. Cong. Proc., 7:375-380.

Stanford, A. E. 1984. Dispersal and behavior of the introduced yellowjacket, Paravespula germanica

(Fab.) (Hymenoptera: Vespidae) in and around Nampa, Idaho. Proc. Wash. St. Ent. Soc., 45:

659-664.

Vasic, Z. 1968. Biological investigations on the German wasp (Vespa germanica F.). Bull. Nat. Hist.

Mus., Belgrade Ser. B., 23:211-224.

PAN-PACIFIC ENTOMOLOGIST

65(1), 1989, pp. 89-96

The Phoretic Behavior and Olfactory Preference of

Macrocheles muscaedomesticae (Scopoli)

(Acarina: Macrochelidae) in its Relationship

with Fannia canicularis (L.)

(Diptera: Muscidae ) 1

Erin E. Riley Borden

Department of Entomology, Washington State University, Pullman, Washing¬

ton 99164-6432.

Abstract. — The phoretic behavior and olfactory preference of Macrocheles mus¬

caedomesticae (Scopoli) (Acarina: Macrochelidae) were studied in regard to their

relationship with Fannia canicularis (L.) (Diptera: Muscidae). The mites and fly

larvae were located in the top 5 cm of poultry manure cones. An average of one

female mite was found on each female F. canicularis collected in the field.

The olfactory preference of the mite was determined to be for the adult, followed

by the egg, larva, and pupa of F. canicularis, respectively. Studies indicated that

the mites have a negative effect on the oviposition and longevity of the flies.

Fannia canicularis (L.) (Diptera: Muscidae), the little house fly, is an important

pest of humans and livestock. The fly is capable of carrying many viral and

bacterial pathogens and therefore is also a public health concern (Axtell, 1985).

It is a prime pest in poultry houses where the rapid accumulation of manure

serves as an excellent breeding medium for the fly.

In San Diego County, California, F. canicularis is the major pest associated

with poultry ranches. Large populations of flies cause severe annoyance to workers

and fecal material produces unacceptable spots on eggs and equipment. However,

the flies are most troublesome when they disperse to surrounding areas where

they constitute a serious nuisance to the people in neighboring homes and busi¬

nesses. This strains community relationships and sometimes leads to lawsuits.

Fannia canicularis also spreads pathogens that cause poultry diseases; of particular

importance is the virus that causes velogenic viscerotropic Newcastle disease

(VVND), a serious problem in poultry (Axtell, 1985).

Macrocheles muscaedomesticae (Scopoli) (Acarina: Macrochelidae), a mite

commonly found in manure, is an effective predator on the eggs and first instars

of F. canicularis (Axtell, 1961; Pereira and de Castro, 1945, 1947; Steve, 1959;

Wicht and Rodriguez, 1970; Willis and Axtell, 1968). In recent years studies have

concentrated on using this mite, selected pesticides, and microbial agents in in¬

tegrated pest management programs for muscid flies (Wicht and Rodriguez, 1970;

Anderson, 1982; Krantz, 1982).

1 Work done at San Diego State University, continued under project 0807 at the College of Agri¬

culture and Home Economics, Washington State University, Pullman, Washington 99164-6432.

PAN-PACIFIC ENTOMOLOGIST

Behavioral studies (Axtell, 1964; Farish and Axtell, 1966, 1971; Kinn, 1966;

Jail and Rodriguez, 1970) have detailed some of the predatory behavior of the

mite on the house fly, and Jail and Rodriguez (1970) discovered that olfactory

cues directed the mite to its host. It preferred adult house flies to eggs. The adult

female of M. muscaedomesticae, in addition to being a predator, is also phoretic

on adult F. canicularis (Steve, 1959; Singh et al., 1966; O’Donnell and Nelson,

1967; Wicht and Rodriguez, 1970; Axtell, 1981). It is unknown if this relationship

is simply phoresy or if parasitism (feeding by the mites) is also involved.

The purposes of this study were to ascertain the location of M. muscaedomes¬

ticae and the immature stages of F. canicularis in poultry manure cones, to de¬

termine if M. muscaedomesticae is found on adult F. canicularis in the field, and

to determine if there is a preference by M. muscaedomesticae for certain stages

of F. canicularis. In addition, the relationship between adult female M. muscae¬

domesticae and adult F. canicularis was examined by studying oviposition and

longevity of the flies.

Materials and Methods

Fly Population

Rearing. —Fannia canicularis (Laboratory stock, University of California, Riv¬

erside) were reared on spent (previously composted non-heat generating which is

suitable for the low temperature tolerance of the little house fly) Musca domestica

(L.) CMSA (Chemical Specialties Manufacturers Association) medium that was

frozen to kill any remaining M. domestica. Four sleeve cages each containing ca.

100 adult flies were maintained throughout the study, and were provided with

water and a powdered sugar-milk mixture (1:1). Fresh water and fresh protein

mixture were provided every third day. Eggs were collected by placing a tablespoon

of spent CSMA medium (which had been sewn pillow-like into a piece of muslin

cloth) into the sleeve cage for 24 hr. Eggs could be removed from the pillows by

a simple rinsing with distilled water.

Mite Population

Rearing. — The mites were collected from the surface layer of chicken manure

from the Hiliker Poultry Ranch in Lakeside, California (San Diego County). The

top 3-5 cm of a manure cone was collected and stored in waxed containers until

the sample could be placed into a modified Berlese funnel (MacFadyen, 1953;

Averbach and Crossley, 1960) to separate the mites.

Stock cultures were initiated by taking samples from jars below the Berlese

funnels, chilling them (2-3 min), and sorting the M. muscaedomesticae from other

arthropods. Female mites, easily recognized by their oval body and ventral plates

(Wade and Rodriguez, 1961), were placed via a camel’s hair brush into plastic

containers (9 x 9 cm with a 4 x 4-cm screened hole in the lid) that contained

moist spent CSMA medium. The mites were maintained in an incubator at 27

± 1°C. Although relative humidity was not controlled, water was added every

third day so that the medium remained slightly damp. Frozen fly eggs were added

to the cultures every other day. Cultures remained in good condition for about 5

wk, until fungal growth developed in the medium and it had to be replaced.

VOLUME 65, NUMBER 1

Collection of Manure Samples

Collections of manure were made to determine where in the manure cone the

M. muscaedomesticae and larvae of F. canicularis were located. Pie-tin (12.5 x

3.5-cm) sized samples were taken from the cone with a wooden spatula (10x6

cm). The samples were stored in wax containers with plastic wrap cover to prevent

desiccation until analyzed. Each sample was divided into 5 parts of ca. 10 g each

for analysis under a dissecting microscope. The samples were picked apart with

a metal probe and visually scored for M. muscaedomesticae and F. canicularis

larvae. Manure samples were taken at 5-cm, 10-cm, 15-cm, and 30-cm levels

from the top of the manure cone. Samples were also taken from directly around

the bottom of the cone and 5 cm out from the bottom of the cone in the dry, flat,

dust area.

Olfactory Orientation Tests

Olfactory response tests were conducted to determine if M. muscaedomesticae

preferred any one stage of the fly. These tests used F. canicularis and M. mus¬

caedomesticae from the laboratory stock.

Apparatus. — The olfactory response of the mite was tested in a glass olfactometer

(Fig. 1) consisting of four horizontal tubes each 1 cm diam and 6 cm long. A fifth

vertical tube, which entered from above and was placed at the center of the

preference tubes, was used to place the mites into the apparatus. Previous re¬

searchers employed a simple choice tube (Jail and Rodriguez, 1970) or an area

type apparatus (Farish and Axtell, 1966). Neither of these designs was suitable

for my investigations.

Preliminary bias tests. -Fifty-two mites were used for this experiment. The

apparatus was rotated a quarter of a turn after each mite was tested to test for

possible responses due to differences in lighting or for other directional responses

due to unknown factors. The mites were introduced into the apparatus individ¬

ually, and each mite was removed before the next was introduced. The procedure

was modified from previous experiments where mites were tested in large groups

(Farish and Axtell, 1966; Jail and Rodriguez, 1970). In these tests, mites were

run separately to eliminate any possible following behavior. Each mite was taken

from its culture with a fine camel’s hair brush, placed into the center of the

apparatus, and given 30 sec to make a choice. If no choice was made, the mites

were removed and discarded. A choice was recorded when a mite crawled into

one of the four arms a distance of at least 2 cm. The mite was then transferred

to a new stock container. None were used repeatedly. Of the 50 mites tested, 12

went down arm 1, 12 down arm 2, 11 down arm 3, and 12 down arm 4. This

indicated the apparatus was suitable for olfactory tests.

Olfactory tests. — This experiment was conducted to determine if any stage of

the fly was preferred by the mite. Viable eggs, larvae, pupae, and adults of F.

canicularis were used. One developmental stage of the fly was live mounted at

the end of one of the tubes with wax. An egg was mounted in arm 1, a larva in

arm 2, a pupa in arm 3, and an adult in arm 4. The mites were introduced one

at a time and removed after making a choice. Again, the mites were given 30 sec

to make a choice after which time they were removed. A choice was defined as

the mite walking 2 cm into one of the arms. The apparatus was again rotated

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Apparatus used to test the olfactory preference of the mite. A = plan of the apparatus;

B = elevation of apparatus.

after each trial to control for differences in room lighting or possible directional

preference by the mites. A total of 200 mites were tested in four replicates con¬

sisting of 48, 52, 48, and 52 mites, respectively. After a replicate was completed,

the fly stages were removed, and the apparatus was washed with 70% ethanol and

rinsed with water. Fresh fly stages were then mounted in the apparatus for the

next test.

Field Collection of Adult Fannia canicularis

Fannia canicularis were held collected to determine the average number of M.

muscaedomesticae found on adult flies. The collection trap consisted of a card¬

board, triangular trap with sides of 9.5 cm with a rectangular sticky board base

VOLUME 65, NUMBER 1

9.8 x 17 cm. Inside the trap a cotton wick hung above the sticky board. The wick

was saturated with “Grandma’s molasses”® obtained locally.

A fresh cotton wick with molasses was furnished every second week. The sticky

boards were removed from the traps when they became covered with insects, or

every 2 wk and were replaced with fresh boards. Boards were placed into plastic

bags for transport to the laboratory for examination under a dissecting microscope.

The number and sex of F. canicularis on the board were noted, along with the

number of attached mites. The collection and examination of flies continued from

17 February through 27 April 1986.

Studies on the Phoretic Behavior of

Macrocheles muscaedomesticae

Since previous experiments had suggested that the mite had a deleterious effect

on the fly, I decided to investigate oviposition and longevity.

The normal, average number of eggs laid by F. canicularis was determined by

placing 32, 7-8-day-old female flies (which had previously not oviposited) into

individual vials. The flies were chilled for 5 min to facilitate handling. Each vial

contained one muslin cloth egging pillow (2x2 cm) filled with previously thawed,

spent CSMA fly medium. The flies were then held at 25°C for 24 hr. After 24 hr,

the number of eggs present was determined.

A second test was performed in which one female mite per fly was added. Since

only female mites are phoretic (Axtell,1964) they were the only sex used. Each

mite was picked at random from a stock container with a fine brush and placed

on an immobilized fly. After 24 hr, a count of fly eggs present was again made.

Longevity experiment.— These tests were conducted to determine if M. mus¬

caedomesticae affected the life span of the fly. Fifty 3-day-old flies were placed

into individual vials. Each fly had one female mite placed on it. A control group

of 50 miteless flies was run simultaneously under the same conditions. The vials

were held at 25°C, and no food was supplied during the experiment (Jail and

Rodriguez, 1970). Longevity was recorded in days.

Results

Collections from Chicken Manure

Macrocheles muscaedomesticae was found together with F. canicularis larvae

only in the top 5 cm of the manure cone (Fig. 2), although fly larvae continued

to be present to at least a depth of 10 cm. None of the other sample sites yielded

either flies or mites.

Olfactory test. — Two hundred mites were tested in replicates of 48, 52, 48, and

52 mites, respectively. Sixty chose the egg, 28 the larvae, 36 the pupa, and 70 the

adult. The mite clearly showed a preference for the adult and egg (chi-square,

25.52 significant to 0.001).

Of the 109 adult F. canicularis collected, only 11% had mites and a maximum

of one female mite per fly was found. In addition all mites were found on female

flies, although both male and female flies were found in the traps.

Nine of the 32 flies tested without mites did not lay eggs, 22 of the flies with

mites did not lay eggs (z-test, z = 2.97, significant at 0.05, two-tailed test) (Won-

nacott and Wonnacott, 1985). Dissection revealed that all the flies that failed to

oviposit had mature eggs in the ovaries. A comparison of the flies alone and the

PAN-PACIFIC ENTOMOLOGIST

flies with mites using a completely randomized analysis of variance test resulted

in an F -value of 7.71 which is significant at 0.01 (Wonnacott and Wonnacott,

1985). A comparison of the two groups of flies using the same statistical test, but

excluding the flies that did not oviposit, results in an F -value of 4.12 which is

significant at 0.10 level.

The average life span of flies without mites was 1.5 days while that of flies with

mites was 1.3 days. This was not a significant difference.

Discussion

The mite showed a definite olfactory preference for the adult and egg stage of

the fly, and the phoretic studies indicated that the mite has a negative effect on

oviposition and perhaps to some extent longevity of the fly.

The locations of the mites and flies in the top 5 cm of the manure cones suggests

the mite may be an important controlling agent of F. canicularis on poultry ranches

in San Diego County. On poultry ranches where the flies are in high population

a staggered removal of manure cones may facilitate maintaining a high predatory

mite population. Also chemical spot treating the manure cones and avoiding the

high mite areas may help decrease fly populations (Axtell, 1985).

The preference of the mite for the adult fly agrees with the results of a similar

study on M. domestica (Jail and Rodriguez, 1970). Perhaps the phoretic attach¬

ment of the mite to the fly takes place when the manure is less attractive (e.g.,

dry) to the fly and losing its ability to support a high mite population. The mites

would benefit by choosing an adult fly for transport to a new manure source, and

hence, a food supply in the form of fly eggs.

A single mite on F. canicularis reduced longevity and oviposition, but there

still is speculation whether there is actual feeding by the mite on the fly. Jail and

Rodriguez (1970) attempted to show that the mite feeds on the fly by weighing

the fly after infestation with mites. They found that flies lost weight compared to

a control, and attributed this weight loss to the mites sucking haemolymph from

the fly. Further studies investigating actual feeding on the flies should test the

VOLUME 65, NUMBER 1

longevity of the mites on flies compared to mites alone, and changes in the water

content of the mites. The relatively high number of mite infested flies that did

not lay eggs suggests that mites may reduce little house fly populations in this

manner. However, since my tests lasted only 24 hr, the oviposition experiment

must be carried out over the total life cycle of the fly to determine if the mite can

reduce the fly population by completely inhibiting oviposition. However, since

ammonia vapor from manure causes the mites to drop from the flies (Pereira and

de Castro, 1947), this may not work.

These studies confirm the potential for the use of M. muscaedomesticae as a

biological control agent of flies. They also show that M. muscaedomesticae has

the potential to be an important part of integrated fly control of F. canicularis in

San Diego County.

Acknowledgments

I would like to thank Dr. Ronald Monroe, Dr. Michael Atkins, Dr. Robert

Yaremko, and Dr. Brad Mullens for their advice. I would especially like to thank

Mr. Harold Hiliker for the use of his poultry ranch in Lakeside, California. I

would also like to thank Dr. Roger Akre and Richard Zack for reviewing this

paper and Mark Borden, Terry Miller, and Elizabeth Myhre for editorial assis¬

tance.

Literature Cited

Anderson, J. R. 1982. Mites as biological control agents of dung-breeding pests: practical consid¬

erations and selection for pesticide resistance. Pp. 99-102 in M. Hoy, G. Cunningham, and L.

Knutson (eds.), Proceedings of a conference on the biological control of pests by mites. Univ.

California, Berkeley, 185 pp.

Averbach, S. I., and D. A. Crossley. 1960. A sampling device for soil microarthropods. Acarologia,

2:281-290.

Axtell, R. C. 1961. New records of North American Macrochelidae (Acarina: Mesostigmata) and

their predation rates on the house fly. Ann. Entomol. Soc. Am., 56:628-633.

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Mesostigmata) to the house fly. Ann. Entomol. Soc. Am., 57:584-587.

-. 1981. Use of predators and parasites on poultry housing. Pp. 26-43 in R. S. Patterson (ed.),

Proceedings of a workshop on the status of biological control of filth flies. Univ. Florida,

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-. 1985. Arthropod pests of poultry. Pp. 269-295 in R. E. Williams, R. D. Hall, A. B. Bruce,

and P. J. Scholl (eds.), Livestock entomology. New York. 335 pp.

Farish, D. J., and R. C. Axtell. 1966. Sensory function of palps and first tarsi of Macrocheles

muscaedomesticae (Acarina: Macrochelidae), a predator of the house fly. Ann. Entomol. Soc.

Am., 59:165-170.

-, and-. 1971. Phoresy redefined and examined in Macrocheles muscaedomesticae (Ac¬

arina: Macrochelidae). Acarologia, 13:16-29.

Jail, M., and J. G. Rodriguez. 1970. Studies of behavior of Macrocheles muscaedomesticae (Acarina:

Macrochelidae) with emphasis on its attraction to the house fly. Ann. Entomol. Soc. Am., 63:

738-744.

Kinn, D. N. 1966. Predation by the mite Macrocheles muscaedomesticae (Acarina: Macrochelidae)

on three species of flies. J. Med. Entomol., 3:151-158.

Krantz, G. W. 1982. Mites as biological control agents of dung breeding flies with special reference

to the Macrochelidae. Pp. 91-98 in M. Hoy, G. Cunningham, andL. Knutson (eds.), Proceedings

of a conference on the biological control of pests by mites. Univ. California, Berkeley, 185 pp.

MacFadyen, A. 1953. Notes on methods for the extraction of small soil arthropods. J. Anim. Ecol.,

22:65-77.

O’Donnell, A. E., and E. L. Nelson. 1967. Predation by Fuscuropoda vegetans (Acarina: Uropodidae)

PAN-PACIFIC ENTOMOLOGIST

and Macrocheles muscaedomesticae (Acarina: Macrochelidae) on the eggs of the little house fly

Fannia canicularis. J. Kans. Entomol. Soc., 40:441-443.

Pereira, C., and M. P. de Castro. 1945. Contribuicao para o conhecimento da especie tipo de

“Macrocheles Latr.” “Acarina”: “M. muscaedomesticae (Scopoli, 1772).” Arch. Inst. Biol., 16:

153-186.

-, and-. 1947. Forese e parthenogenese arrentoca em “M. muscaedomesticae ” (Scopoli)

“Acarina: Macrochelidae” e sua significacao ecologica. Arch. Inst. Biol., 18:71-89.

Singh, P., W. E. King, and J. G. Rodriguez. 1966. Biological control of muscids as influenced by

host preference of Macrocheles muscaedomesticae (Acarina: Macrochelidae). J. Med. Entomol.,

3:78-81.

Steve, P. C. 1959. Parasites and predators of Fannia canicularis (L.) and Fannia scalaris (F.). J.

Econ. Entomol., 52:530-531.

Wade, C. F., and J. G. Rodriguez. 1961. Life history of Macrocheles muscaedomesticae (Acarina:

Macrochelidae), a predator of the house fly. Ann. Entomol. Soc. Am., 54:776-781.

Wicht, M. C., and J. G. Rodriguez. 1970. Integrated control of muscid flies in poultry houses using

predator mites, selected pesticides and microbial agents. J. Med. Entomol., 7:687-692.

Willis, R. R., and R. C. Axtell. 1968. Mite predators of the house fly: a comparison of Fuscuropoda

vegetans and Macrocheles muscaedomesticae. J. Econ. Entomol., 61:1669-1674.

Wonnacott, R. J., and T. H. Wonnacott. 1985. Introductory statistics. J. Wiley, New York, 649 pp.

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Pan-Pacific Entomologist

RESH, V. H.-Donald G. Denning (1909-1988).

FLINT, O. S., JR. and D. G. DENNING—Studies of Neotropical caddisflies, XLI: New species

and records of Austrotinodes (Trichoptera: Psychomyiidae)___

DENNING, D. G. —Eight new species of Trichoptera....

DOWELL, R. V. and R. GILL—Exotic invertebrates and their effects on California.

KNIGHT, A. L. and B. A. CROFT—Host discrimination by the gregarious parasitoid Onco-

phanes americanus (Hymenoptera: Braconidae).

EHLER, L. E.—Observations on Scutellista cyanea Motsch. (Hymenoptera: Pteromalidae).

BOHART, R. M.—North American Pterocheilus. I. Subgenus Onchopterocheilus (Hymenop¬

tera: Eumenidae) ___.___

DOWELL, R. V.—Toxicity of water extracts of Murraya paniculata Jack leaves to immature

citrus blackfly, Aleurocanthus woglumi Ashby (Homoptera: Aleyrodidae)_

YIN, X. and R. L. SMITH—Three new grasshoppers from the western United States (Orthop-

tera: Acrididae)___________

DODSON, G. N. and D. K. YEATES—Male Bembix furcata Erichson (Hymenoptera: Spheci-

dae) behaviour on a hilltop in Queensland__

GARDNER, M. R. and R. M. SHELLEY—New records, species, and genera of Caseyid mil-

lipeds from the Pacific Coast of North America (Diplopoda: Chordeumatida: Caseyidae)

SCIENTIFIC NOTES. 16

SAN FRANCISCO, CALIFORNIA • 1989

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

EDITORIAL BOARD

J. A. Chemsak, Editor

R. S. Lane, Associate Editor

W. J. Pulawski, Treasurer

R. M. Bohart J. A. Powell