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4 | VOL. 104 JULY 2002 NO. 3 =f, 9 kK (ISSN 0013-8797)

; PROCEEDINGS

of the

ENTOMOLOGICAL SOCIETY of WASHINGTON

> PUBLISHED 1S ay, \ QUARTERLY

CONTENTS

CONTENTS BLANK, STEPHAN M.—Taxonomic notes on Strongylogasterini (Hymenoptera: Tenthredinidae) 692 BUFFINGTON, MATTHEW L.—Description of Aegeseucoela Buffington, new name, with notes

on the status of Gronotoma Forster (Hymenoptera: Figitidae: Eucoilinae) .................... 589 CHABOO, CAROLINE S.—Range extensions of New World tortoise beetles (Coleoptera:

Ghrysomelidac2CassiGinaey ies cre acre le fete Serelels Hie zeus one SIS eine een st ec eres 716 FLORES, GUSTAVO E. and CHARLES A. TRIPLEHORN—Entomobalia, new genus, the first

member of Nycteliini (Coleoptera: Tenebrionidae) from Brazil .....................-....00 ees 602

GOEDEN, RICHARD D.—Life history and description of immature stages of Oxyna palpalis (Coquillett) (Diptera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern GalitOrnii al rests ce ee erecta tis acs Annee mttete ms ROP arated ci vce ha stl ig een ne aM Cte nae eR eee ae 537

GOEDEN, RICHARD D.—Life history and description of immature stages of Goedenia rufipes (Curran) (Diptera: Tephritidae) on Jsocoma acradenia (E. Greene) E. Greene in southern (CPAIIFOVTINE es ee ad bes Sania bed te anaes ti A Ga SO ee Att Rae ROPE AES SRA ino RSE OuE Roos atone Broa. 576

GOEDEN, RICHARD D.—Life history and description of immature stages of Goedenia setosa (Foote) (Diptera: Tephritidae) on Ericameria brachylepis (A. Gray) H. M. Hall in southern @alifommi as pie acenet: eh oak ears sehes Gh ete ele aaa oe ete sates, be zieiae aye aS REE BS Ae Ce eats aR SERe hala eae 629

GOEDEN, RICHARD D.—Life history and description of adults and immature stages of Goedenia stenoparia (Steyskal) (Diptera: Tephritidae) on Gutierrezia californica (de Candolle) Torrey and A. Gray and Solidago californica Nuttall (Asteraceae) in southern (Bealeton ope hee a oor aaa olotesrak ig Gas tans biadielait Sidhe. Lalagate Beanie as. RETA EMM eRe aayorere eo a Rena Ee 702

GOEDEN, RICHARD D.—Life history and description of adults and immature stages of Goedenia steyskali, n. sp. (Diptera: Tephritidae) on Grindelia hirsutula Hooker and Arnott var. halli (Steyermark) M. A. Lane (Asteraceae) in southern California ...................... 785

HARRISON, B. A., P. B. WHITT, S. E. COPE, G. R. PAYNE, S. E. RANKIN, L. J. BOHN, F. M. STELL, and C. J. NEELY—Mosquitoes (Diptera: Culicidae) collected near the Great Dismal Swamp: New state records, notes on certain species, and a revised checklist for Virginia .... 655

(Continued on back cover)

THE

ENTOMOLOGICAL SOCIETY OF WASHINGTON

)

OFFICERS FOR 2002

GABRIELA CHAVARRIA, President MICHAEL G. PoGuE, Treasurer JONATHAN R. Mawps.Ley, President-Elect RONALD A. OcHoa, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho us B. WILLIAMS, Corresponding Secretary JOHN W. Brown, Past President

Jon A. Lewis, Custodian Davip R. Smirn, Editor

Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MaruHis

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The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 247-248 OF THE JANUARY 2002 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October).

Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168.

Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168.

Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168.

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PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 537-553

LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF OXYNA PALPALIS (COQUILLETT) (DIPTERA: TEPHRITIDAE) ON ARTEMISIA TRIDENTATA NUTTALL (ASTERACEAE)

IN SOUTHERN CALIFORNIA

RICHARD D. GOEDEN

Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu)

Abstract.—Oxyna palpalis (Coquillett) is a univoltine, circumnatal tephritid uniquely reproducing as an inquiline in rosette galls of Rhopalomyia florella Gagné (Diptera: Ce- cidomyiidae) of terminal buds on branches of Artemisia tridentata Nuttall. Its larvae also routinely function as facultative predators of R. florella larvae, novel behavior for Te- phritidae. The egg, first-, second-, and third-instar larvae, and puparia are described and figured for the first time. The egg is distinguished by a pedicel circumscribed by one complete and a partial second ring of irregularly shaped micropyles. All three larval instars of O. palpalis are compared to and distinguished from those of O. aterrima (Doane), its only other known congener in North America. Oviposition occurs in spring (June) in southern California in nearly fully formed galls of R. florella containing young larvae of this cecidomyiid. The young larvae of O. palpalis pass the summer (June—September) as first instars singly in small, central, ovoidal cells basad of the cecidomyiid larvae. Second instars occupy their still-small, separate chambers until late fall/early winter (September— October), when some begin to molt to third instars. By mid-winter (February) all larvae are third instars, which continue to overwinter and grow slowly until the resumption of the spring flush of new plant growth (March). At this time, one to six or more third instars enlarge the central gall chamber to accommodate their faster growth and feed gregariously. Cecidomyiid larvae encountered during this third stadium are killed and devoured; sur- viving immature gall midges usually occupy the periphery of the galls. Pupariation follows in early April, and adults emerge by mid-April. Eurytoma sp. (Hymenoptera: Eurytomi- dae) and Eupelmus sp. (Hymenoptera: Eupelmidae) were individually reared from puparia of O. palpalis as primary, solitary, probably larval-pupal endoparasitoids. Lyrcus sp. (Hy- menoptera: Pteromalidae) was reared from individual puparia as a gregarious, primary endoparasitoid.

Key Words: Insecta, Oxyna, Asteraceae, Artemisia, nonfrugivorous Tephritidae, Rho- palomyia, Cecidomyiidae, biology, taxonomy of immature stages, galls, inquiline, circumnatal life cycle, parasitoids, insect predation

Oxyna palpalis (Coquillett) (Diptera: Te- O. aterrima (Doane) was reviewed by phritidae) is one of two species of Oxyna Foote et al. (1993) and studied by Goeden now known from North America (Foote et (2002b), who synonymized it with O. utah- al. 1993, Goeden 2002b). The other species, ensis Quisenberry.

MATERIALS AND METHODS

The present study was based in large part on dissections of samples of galls of the gall midge, Rhopalomyia florella Gagné (Diptera: Cecidomyiidae), on Artemisia tri- dentata Nuttall prob. ssp. parishii (A. Gray) H. M. Hall and Clements (Asteraceae) col- lected mainly 0.2 km north of the hamlet of Mile High and just south of the hamlet of Largo Vista; 1580-m elevation; Township 4N, Range 9W, Section 4; Angeles National Forest, Los Angeles Co., during 1996 and 1997. Excised R. florella galls, containing eggs and early-instar larvae of O. palpalis, and later, overwintered galls containing third instars and puparia of O. palpalis were sampled mid-monthly from gall-bearing plants during 1996 and 1997. Samples were transported in cold-chests in an air-condi- tioned vehicle to the laboratory and stored under refrigeration for subsequent dissec- tion, photography, description, and mea- surement. Eight eggs, 18 first-, seven sec- ond-, and 19 third-instar larvae and nine puparia dissected from galls were preserved in 70% EtOH for scanning electron micros- copy (SEM). Additional prepuparia and pu- paria in excised, opened galls were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hydrated to distilled water in a de- creasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, |-h immersions in hexa- methyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied and photographed with a Philips XL-30 scanning electron micro- scope in the Central Facility for Advanced Microscopy and Microanalysis, University of California, Riverside.

Most adults reared from isolated prepu- paria and puparia were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reser-

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voir and provisioned with a strip of paper toweling impregnated with yeast hydroly- zate and sucrose. These cages mainly were used for studies of longevity in the insec- tary of the Department of Entomology, University of California, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Two pairs of virgin flies, each consisting of a male and a female obtained from emer- gence cages also were held in a clear-plas- tic, petri dish provisioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observations of their courtship and copulation behavior.

Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993); cecidomyiid names and gall terminology follow Gagné (1989). Ter- minology and telegraphic format used to describe the immature stages follow Goe- den (2001, 2002a, b), Goeden and Norrbom (2001), Goeden and Teerink (1999), and earlier works cited therein. Means + SE are used throughout this paper. All remaining voucher specimens and reared parasitoids of this tephritid reside in my research col- lections. Digital photographs used to con- struct text figures were processed with Ado- be Photoshop® Version 6.

RESULTS AND DISCUSSION

Adult.—Oxyna_ palpalis originally was described as a Tephritis by Coquillett in Baker (1904) from a single male specimen from Ormsby County, Nevada. Quisenberry (1949) in his revision of Oxyna redescribed this species from three females and two males variously from California, Idaho, and Nevada. Foote and Blanc (1963) and Foote et al. (1993) pictured the right wing.

Immature stages.—The egg, first-, sec- ond-, and third-instar larvae, and puparium of O. palpalis are described below.

Egg: Thirteen eggs of O. palpalis dis- sected from field-collected galls of R. flo- rella galls also bearing cecidomyliid larvae were white, opaque, smooth, ellipsoidal,

VOLUME 104, NUMBER 3

Fig. 1.

Egg of Oxyna palpalis: (A) habitus, anterior to left; (B) pedicel 1, (C) pedicel 2, (D) pedicel 3. (The

three pedicels show variation in size, shape, and placement of micropyles.)

0.65 + 0.02 (range, 0.62—0.68) mm long, 0.20 + 0.00 (range, 0.20—0.20) mm wide, smoothly rounded at basal end, with a 0.2- mm, buttonlike pedicel at anterior end (Fig. 1A). The pedicel was circumscribed suba- pically by one complete and a partial sec- ond ring of irregularly shaped micropyles (Bigs IBC LD)

This is the first Oxyna egg pictured at high magnification (Goeden 2002b). On av- erage, the egg of O. palpalis is slightly lon- ger and slightly narrower than that of O. aterrima (Goeden 2002b).

First instar larva: White, ellipsoidal, flat- tened anteriorly and posteriorly (Fig. 2A); body segments with hemispherical or pos- teriorly-directed, short-spinose, minute acanthae on intersegmental areas of meta- thorax and abdominal segments Al through A6 as well as pleura and lateroventrum of

Al through A6; prothorax and gnathoce- phalon smooth, the latter conical (Fig. 2B), both circumscribed by verruciform sensilla (Figs. 2B-1, -2); dorsal sensory organ, well- defined, flat pad (Fig. 2C-1); anterior sen- sory lobe (Fig. 2C-2) bears terminal sen- sory organ (Fig. 2C-3), lateral sensory or- gan (Fig. 2C-4), supralateral sensory organ (Fig. 2C-5), and pit sensory organ (Fig. 2C- 6); stomal sense organ reduced to two ver- ruciform sensilla ventrolaterad of terminal sensory organ (Figs. 2C-7, D-1), not fused with flattened, protrudent, lateral integu- mental petal (Figs. 2C-8, D-2) above each mouthhook, one medial, papillate, integu- mental petal (Figs. 2C-9, D-3) between an- terior sensory lobes and lateral integumen- tal petals; mouthhook bidentate (Figs. 2B- 3, C-10, D-4); median oral lobe laterally compressed, apically rounded (Figs. 2B-4,

540

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

trolateral view, 1—verruciform sensilla on gnathocephalon, 2—verruciform sensilla on prothorax, 3—mouth-

First instar of Oxyna palpalis: (A) habitus, anterior to left, (B) gnathocephalon and prothorax, ven-

hook, 4—median oral lobe; (C) gnathocephalon, frontal, close-up view, 1—dorsal sensory organ, 2—anterior

sensory lobe, 3—terminal sensory organ, 4

lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory

organ, 7—stomal sense organ, 8—lateral integumental petal, 9—medial integumental petal, 10—mouthhook;

(D) oral cavity, ventral view, 1—stomal sense organ,

—lateral integumental petal, 3—medial integumental

petal, 4—mouthhook, 5—median oral lobe, 6—labial lobe, 7—pores.

D-5); labial lobe (Fig. 2D-6) broad, sepa- rated from median oral lobe, and with two pores ventrally (Fig. 2D-7); anterior spira- cle absent; lateral spiracular complexes not seen; caudal segment (Fig. 3A) with a ste- lex sensillum dorsolaterad (Fig. 3A-1), lat- (Figs 3A-2, B), ventrolaterad

erad and

(Figs. 3A-3, C-1) of posterior spiracular plate (Figs. 3A-4, D); posterior spiracular

plate bears two rimae (Fig. 3D-1), ca. 0.005 mm long, and four, unbranched or bifurcate, spinose or apically toothed (some bifurcate) interspiracular processes (Fig. 3D-2), the longest process measuring 0.007 mm; in- termediate sensory complex (Figs. 3A-5, C-

2, D-3) consists of stelex sensillum (Fig. 3C-3) and medusoid sensillum (Fig. 3C-4).

The habitus of the first instar of O. pal- palis is similar to the first instar of O. ater- however, as with Trupanea_ spp. (Goeden and Teerink 1999), Neaspilota spp. (Goeden 2001), and Tephritis spp. (Goeden 2002a), the incidence and patterns of minute acanthae on the thorax and ab- domen show interspecific differences. Ox- yna palpalis has fewer minute acanthae in the intersegmental areas of the thorax and

rima;

abdominal segments Al and A2, and unlike O. aterrima, none on or between abdominal segments A3 through A6 (Goeden 2002b).

VOLUME 104, NUMBER 3

Figs 3: lateral verruciform sensillum, 3—ventrolateral stelex sensillum, 4—posterior spiracular plate, 5—intermediate

First instar of Oxyna palpalis, continued: (A) caudal segment, 1—dorsolateral stelex sensilla, 2—

sensory complexes; (B) lateral stelex sensillum (arrow); (C) 1—ventrolateral verruciform sensillum, 2—inter-

mediate sensory complex (composed of), 3—stelex sensillum, 4—medusoid sensillum; (D) posterior spiracular

plate with 1—two rimae, 2—four interspiracular processes, and 3—two intermediate sensory complexes.

The first instar of O. aterrima has two, medial, integumental petals (Goeden 2002b); whereas, O. palpalis has only one (Figs. 2C-9, D-3). The single, lateral inte- gumental petal above each mouthhook is fused with the stomal sense organ in O. aterrima (Goeden 2002b), but they are not fused in O. palpalis (Figs. 2C-8, D-2). The

mouthhooks are tridentate in the first instar

of O. aterrima (Goeden 2002b), but are bi- dentate in O. palpalis (Figs. 2B-3, C-10, D- 4). The lateral sensilla surrounding the pos- terior spiracular plate of O. aterrima are verruciform (Goeden 2002b; whereas, these are stelex sensilla in O. palpalis (Figs 3A- 2, B). The interspiracular processes on the caudal segment of O. aterrima are un- branched (Goeden 2002b); whereas, some

processes of O. palpalis are two-branched (Fig. 3D-2).

Second instar larva: Ovoidal, rounded anteriorly, truncated posteriorly (Fig 4A), body segments with short-spinose, posteri- orly-directed, minute acanthae (Fig. 4B-1) circumscribing intersegmental areas of tho- rax and abdomen; thorax and gnathoce- phalon smooth (Fig. 4B), the latter conical, the former circumscribed around middle with verruciform sensilla (Fig. 4B-1); dor- sal sensory organ, a well-defined, flat pad (Figs. 4C-1, D-1); anterior sensory lobe (Fig. 4C-2) with terminal sensory organ (Fig. 4C-3), lateral sensory organ (Fig. 4C- 4), supralateral sensory organ (Fig. 4C-5), and pit sensory organ (Fig. 4C-6); two pairs of foliose, protrudent, lateral integumental

542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Second instar of Oxyna palpalis: (A) habitus, anterior to left; (B) prothorax and gnathocephalon, frontolateral view, 1—minute acanthae, 2—verruciform sensillum; (C) gnathocephalon, frontolateral view, 1—

dorsal sensory organ, 2

anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supra-

lateral sensory organ, 6—pit sensory organ, 7—lateral integumental petals, 8—medial integumental petals; (D)

gnathocephalon, ventrolateral view, 1—dorsal sensory organ, 2 integumental petals, 4—oral ridges, 5—-stomal sense organ, 6—mouthhook, 7—median oral lobe, 8

9—pores.

petals (Figs 4C-7, D-2) above each mouth- hook, two pairs of papillate, medial inte- gumental petals (Fig. 4C-8) between each anterior sensory organ, the ventral pair elongate (Fig. 4D-3); at least four oral ridg- es (Fig. 4D-4) ventrolaterad of each anterior sensory lobe and stomal sense organ (Fig. 4D-5); mouthhook (Fig. 4D-6) with two teeth; median oral lobe laterally flattened (Fig. 4D-7); labial lobe (Fig. 4D-8) broad, separated from median oral lobe, with two pores ventrally (Fig. 4D-9); anterior thorac- ic spiracle (Fig. 5A) with five, rounded, wedge-shaped papillae; lateral spiracular complexes not seen; caudal segment (Fig. 5B) with stelex sensillum dorsolaterad

lateral integumental petals, 3—ventral, medial labial lobe,

(Figs. 5B-1), laterad (Fig 5B-2), and ven- trolaterad (Figs. 5B-3, C-1) of posterior spi- racular plate (Figs. 5B-4, D); posterior spi- racular plate with four, elongate, upright, foliose, interspiracular processes (Fig. 5D- 1), the longest process measuring 0.011 mm; intermediate sensory complex (Figs. 5B-5, C-2) consists of stelex sensillum (Fig. 5C-3) and medusoid sensillum (Fig. 5C-4).

One major difference between the second instars of O. palpalis and O. aterrima is the absence of a black marking on the abdom- inal ventrum of the former species, where the ventrum and pleura of the latter species are densely covered with knoblike minute acanthae (Goeden 2002b). In O. palpalis

VOLUME 104, NUMBER 3

Fig. 5.

543

i Wh u

(UGE Se Ne aS 4) ae ‘i A aj \\ ie, ¥ f az ‘“

ES a iff aS fh) - ‘ i a \

Second instar of Oxyna palpalis, continued: (A) anterior spiracle; (B) caudal segment; caudal seg-

ment, 1—dorsolateral stelex sensilla, 2—lateral stelex sensilla, 3—ventrolateral stelex sensilla, 4—posterior spiracular plate, 5—intermediate sensory complexes; (C) 1—ventrolateral stelex sensillum, 2—intermediate sen- sory complex (composed of), 3—stelex sensillum, 4—medusoid sensillum; (D) posterior spiracular plate, 1—

four interspiracular processes.

the minute acanthae are short-spinose and confined mainly to the intersegmental areas of the thorax and abdomen. Oxyna palpalis has two pairs of medial integumental petals between each anterior sensory lobe (Fig. 4C-8); whereas, O. aterrima has only one such pair (Goeden 2002b). The inner, ven- tral oral ridge among the four such ridges of O. aterrima is ventrally toothed and fused with the stomal sense organ (Goeden 2002b); whereas, none of the four oral ridg- es of O. palpalis is toothed nor fused with the stomal sense organ (Figs. 4D-3, 4). Third instar larva: Pale yellow or white, ovoidal, tapering anteriorly, truncated pos- teriorly, distinctly segmented (Fig. 6A), short-spinose, posteriorly-directed, minute acanthae in transverse bands on dorsopos-

terior fifth of gnathocephalon (Figs. 6A, B- 1), minute acanthae also circumscribe an- terior fourth of thoracic segments and an- terior fourth of abdominal segment A1, dor- sum, ventrum and anterior third of pleura of A2-A7, and all but posterior of spiracular plate A-8 (Fig. 7D-1); gnathocephalon smooth, conical (Fig. 6-B); dorsal sensory organ well-defined, hemispherical (Figs. 6B-2, C-1); anterior sensory lobe bears ter- minal sensory organ (Figs. 6B-3, C-2), lat- eral sensory organ (Fig. 6C-3), supralateral sensory organ (Fig. 6C-4), and pit sensory organ (Fig. 6C-5); nine or 10 oral ridges (Fig. 6D-1) laterad and ventrolaterad of an- terior sensory lobe and stomal sense organ, at least six lobes ventrally toothed and sep- arate from prominent stomal sense organ

544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Third instar of Oxyna palpalis: (A) habitus, anterior to left; (B) gnathocephalon, ventrolateral view, I—minute acanthae, 2—dorsal sensory organ, 3—terminal sensory organ, 4—stomal sense organ, 5—mouth- hook, 6—median oral lobe, 7—lateral integumental petal, 8—inner, lateral integumental petals; (C) gnatho-

cephalon, dorsolateral view, 1—dorsal sensory organ, 2—terminal sensory organ, 3—lateral sensory organ, 4— supralateral sensory organ, 5—pit sensory organ, 6—stomal sense organ, 7—medial integumental petals, 8— lateral integumental petals, 9—inner, lateral integumental petals; (D) gnathocephalon and oral cavity, ventrolat- eral view, 1—oral ridges, 2—three-toothed mouthhook, 3—median oral lobe, 4 5—inner, lateral integumental petal, 6

lateral integumental petals,

labial lobe, 7—pores; (E) anterior spiracle with three papillae; (F) anterior spiracle with four papillae.

VOLUME 104, NUMBER 3

(Fig. 6B-4, C-6); mouthhook (Fig. 6B-5, D- 2) tridentate (Fig. 5D-2); median oral lobe laterally flattened, apically rounded (Figs. 6B-6, D-3); three pairs of medial integu- mental petals in vertical row between an- terior sensory lobes (Fig. 6C-7); five, lateral integumental petals between each mouth- hook and anterior sensory lobe, including four foliose, lateral petals (Fig. 6B-7, C-8, D-4) and single, inner, elongate, papillate, lateral petal (Figs. 6B-8, C-9, D-5); labial lobe (Fig. 6D-6) broad, separated from me- dian oral lobe, and with two pores ventrally (Fig. 6D-8); anterior thoracic spiracle with three (Fig. 6E, 7A-1) or four, rounded, wedge-shaped papillae (Fig. 6F); lateral spiracular complex of mesothorax with closed, relict spiracle (Fig. 7A-2) and six verruciform sensilla (Figs. 7A-3) in vertical row posteriorad of spiracle, additional ver- ruciform sensillum posteriorad of fourth-

most-vertical sensillum; lateral spiracular

complex of metathorax similarly composed of closed, relict spiracle (Figs. 7A-4, B-1) and four verruciform sensilla (Figs. 7A-5,

B-2) similarly positioned; lateral spiracular

complex of first abdominal segment com- posed of closed, presumably relict spiracle (Figs. 7A-6, C-1) and three verruciform sensilla (Figs. 7A-7, C-2) similarly posi- tioned; posterior spiracular plate (Figs. 7D- 1, 8A) bears three, broadly elliptical rimae (Fig. 8A-1), ca. 0.03 mm long, and four, unbranched, spiniform, interspiracular pro- cesses (Fig. 8A-2), each ca. 0.007 mm long; stelex sensilla dorsolaterad (Figs. 7D-2, 8B), verruciform sensilla laterad (Figs. 7D- 3, 8C), and stelex sensilla ventrolaterad (Fig. 7D-4) of posterior spiracular plate; in- termediate sensory complexes (Figs. 7D-5, 8D) consist of stelex sensillum (Fig. 8D-1) and medusoid sensillum (Fig. 8D-2). Fortunately, the third instars of both spe- cies of Oxyna known from North America have now been described in considerable detail, facilitating comparison between them. For example, the minute acanthae on third instars of O. aterrima are fewer in number, occupy fewer body segments, and

545

different (Goeden 2002b) than the minute acanthae on O. palpalis. Oxyna palpalis has at least nine or 10 oral

form patterns

ridges (Fig. 6D-1), six of which have ven- trally toothed margins; whereas, O. aterri- ma has only two oral ridges (Goeden 2002b). All five lateral integumental petals of third instar O. aterrima are papillate (Goeden 2002b); whereas, only one of the five lateral integumental petals of the third instar of O. palpalis is papillate, the rest are foliose (Figs. 6B-7, C-8, D-4). The lateral spiracular complex of the mesothorax of O. has four verruciform sensilla (Goeden 2002b); whereas, this same com- plex in O. palpalis has seven verruciform sensilla (Figs. 7A-3). Similarly, the lateral spiracular complex of the first abdominal segment of O. aterrima has four verruci- form sensilla, two pairs in separate vertical rows, (Goeden 2002b); whereas, this same complex in O. palpalis has three, vertical, verruciform sensilla (Figs. 7A-7, C-2). The caudal segments of third instars of these congeners also differ considerably. The dorsolateral, lateral, and ventrolateral sensilla surrounding the posterior spiracular plate of O. aterrima are all verruciform. The intermediate sensory complex of this species also uniquely consists of two ver- ruciform sensilla, and this composition dis- tinguishes O. aterrima from third instars of all other nonfrugivorous tephritids exam- ined by my coworkers and me to date (Goe- den 2002b). In O. palpalis, the dorsolateral and ventral lateral sensilla are stelex in form (Figs. 7D-2, -4E, 8B), and the inter- mediate sensory complex is comprised of a stelex sensillum (Fig. 8D-1) and a medu- soid sensillum (Fig. 8D-2), like all other third instars described by us to date. Differences noted between the first and second instars of O. palpalis include the usual acquisition of an anterior spiracle in the second instar, the increase in the number of lateral integumental petals from one to three, and the drastic change in shape of the interspiracular processes (Figs. 3D-2, 5D- 1). Differences noted between the second

aterrima

546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Bice a metathoracic spiracle, 3—verruciform sensillum, 4—mesothoracic spiracle, 5—verruciform sensillum, 6—first abdominal segment spiracle, 7—verruciform sensillum; (B) part of lateral spiracular complex of metathorax, 1— spiracle, 2—verruciform sensillum, (C) part of lateral spiracular complex of metathorax, 1—spiracle, 2—ver- ruciform sensillum; (D) anal segment, 1—posterior spiracular plate, 2—dorsolateral stelex sensillum, 3—lateral verruciform sensillum, 4—ventrolateral stelex sensillum, 5—intermediate sensory complexes.

and third instars include a change in the in- cidence of the minute acanthae as described above. Other changes include increases from three (Fig. 4C-7) to five (Figs. 6B-7; B-8; C-9; D-4, -5) in the number of lateral integumental petals, which all are foliose in the second instar, but add a central, papillate petal in the third instar. Two pairs of medial integumental petals are present in the sec- ond instar (Fig. 4C-8); whereas, three pairs occur in the third instar (Fig. 6C-7). The anterior spiracles with five papillae in the second instar (Fig. 5A) compare with three (Figs. 6E, 7A-1) or four papillae (Fig. 6F) in the third instar. The sensilla surrounding the posterior spiracular plate are the same in number as in the second instar; however,

Third instar of Oxyna palpalis, continued: (A) lateral spiracular complexes, 1—anterior spiracle, 2—

in the second instar the lateral sensilla are stelex (Fig. 5B-2), not verruciform, as in the third instar (Figs. 7D-3, 8C). The inter- spiracular processes on the posterior spirac- ular plate of the second instar are larger, foliose, and upright (Fig. 5D-1) compared to the small, resupinate, spiniform process- es of the third instar (Fig. 8D-2). Puparia: Reniform-ellipsoidal, light-, yellow-, or reddish-brown, rarely white (Fig. 9B), anterior end bears the invagina- tion scar and anterior thoracic spiracles; caudal segment bears posterior spiracular plates (Fig. 9C), each with three broadly el- liptical, raised rimae (Fig. 9C-1) and four, interspiracular processes (Fig. 9C-2). One hundred and five puparia averaged 3.04 +

VOLUME 104, NUMBER 3

Fig. 8. interspiracular processes, (B) dorsolateral stelex sensillum, (C) lateral verruciform sensillum, (D) intermediate sensory complex, 1—stelex sensillum, 2—medusoid sensillum, 3—minute acanthae.

0.03 (range, 2.13—3.70) mm in length; 1.60 + 0.016 (range, 1.14—1.99) mm in width.

DISTRIBUTION AND HOosTs

Oxyna_ palpalis uniquely reproduces as an inquiline in the rosette galls of Rhopa- lomyia florella of terminal buds on branch- es of Artemisia tridentata in southern Cal- ifornia. In this capacity it also functions as a facultative predator on the larvae and pu- pae of R. florella (see below), which also renders this tephritid rare and fascinating in the annals of tephritidology. Novak et al. (1967) reported that the larvae form small, succulent, polythalamous galls on small branches of A. tridentata in Idaho, but as documented in the next section of this pa- per, this interpretation was incorrect, as O. palpalis is not a gall-former. Foote et al. (1993) mapped the distribution of O. pal-

Third instar of Oxyna palpalis, continued:

(A) posterior spiracular plate, 1—three rimae, 2—four

palis to include California, Idaho, Nevada, Oregon, Utah, Washington, and Wyoming. Thus, like that of O. aterrima (Goeden 2001b), the distribution of O. palpalis may coincide wholly with A. tridentata sensu latu, or in part with one or more of its sub- species (Hickman 1993). This is a shrub that inhabits dry soils, valleys, slopes from 300\to: 3000-- mi im the western U:S.)1-e:, north to Washington, the North Central States and south to New Mexico (Hickman 1993). The distribution of A. tridentata also extends into southwestern Canada (Barkley 1986).

BIOLOGY

Egg.—In each of 31, nearly fully formed rosette galls of Rhopalomyia florella (Gag- ne 1989) already containing larvae of this cecidomyiid (Fig. 1OA), most of 51 eggs of

548

Fig. 9. anterior to left; (B) puparium, habitus, anterior to left; (C) posterior spiracular plate, 1—three rimae, 2—four interspiracular processes.

Oxyna palpalis, (A) prepuparium, habitus,

O. palpalis were found inserted separately, a few in small clusters, pedicel-last, in leaf bases to depths of half to all of the egg lengths (Fig. 10B). Leaves at the margins or in the centers of the galls received the

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

most oviposition, but a few eggs were layed upon, but not penetrating the tissues be- tween adjacent bases of inner or outer leaves. About half of the eggs lay with their long axes parallel to the long axes of the galls (Fig. 10B); the remainder lay at angles of 5° to 45° to the long axes. Oviposition in galls was scattered throughout a stand of A. tridentata, like and sympatric with the galls of R. florella, and not confined to certain individual host-plants galled repeatedly Over successive years (Headrick and Goe- den 1998). This is similar to the pattern of the incidence of galls of O. aterrima (Goe- den 2002b). Some galls bore tephritid eggs on opposite sides that showed different de- grees of embryony, presumably indicative of oviposition by different females at dif- ferent times. An average of 3.4 = 04 (range, 1-10) eggs was found in these 31 galls. Galls containing eggs of O. palpalis (Fig. 1OA) averaged 6.3 + 0.4 (range, 4.3— 11.5) mm in length, and 4.0 + 0.2 (range, 0.6—6.8) mm in width. The linear leaves in- vesting these galls averaged 3.5 + 0.3 (range, 2.0—8.6) mm in length and 0.9 + 0.04 (range, 0.6—1.3) mm in width. Larva.—Upon eclosion, most first instars of O. palpalis tunneled into the center of the gall basad of the young, cecidomyiid larvae and leaf bases (Fig. 10C), where each first instar remained for up to 3 months within an individual, ovoidal, open, smooth-walled cell. An average of 2.3 + 0.2 (range, 1—5) first instars of O. palpalis were found in a total of 53 infested galls sampled monthly during the first 3 months following oviposition, during which time the subspheroidal (smaller) (Fig. 1OA) to hemispheroidal (larger) galls averaged 6.9 + 0.3 (range, 3.1—11.5) mm in length and 4.8 + 0.2 (range, 2.9—7.4) mm in width. The linear leaves laterally surrounding the galls averaged 4.4 + 0.2 (range, 1.4—8.6) mm in length and 0.9 + 0.03 (range, 0.6— 1.4) mm in width. Seventy-three ovoidal to spheroidal cells containing the first instars of O. palpalis averaged 0.78 + 0.09 (range, 0.28—4.6) mm in length and 0.48 + 0.18

VOLUME 104, NUMBER 3

(range, 0.21—0.99) mm in width. The size of most of these cells containing first instars slowly increased during the first stadium. Unfortunately, the number of cecidomyiid larvae was not recorded in most galls; how- ever, 14 galls containing first instars of O. palpalis also were noted to contain an av- erage of 2.2 + 0.5 (range, 1-8) larvae of R. florella. Also, it was subsequently noted that most galls of R. florella without O. pal- palis contained only a single, centrally lo- cated larva (Fig. 11D). Thus, some galls containing first instars of O. palpalis also contained dead, centrally located, cecido- mylid larva(e) presumably killed after ac- cidental contact with a tephritid larva. In- deed, at least for later instars of O. palpalis in those many galls found to be lacking R. florella (see below), this relationship may represent nutritionally advantageous, facul- tative predation.

The larva next molted to the second in- star (Fig. 10D), evidenced as the cast ce- phalopharyngeal skeleton remaining in the cell. This instar may also last about 3 months and is one stage found in overwin- tering galls in southern California (see be- low). The external dimensions of 28 galls found to contain second instars measured 6.8 + 0.3 (range, 3.7—11.4) mm in length by 5.7 + 0.3 (range, 2.9-8.6) mm in width. The linear leaves investing these galls av- eraged 4.6 += 0.3 (range, 1.7—9.1) mm in length and 1.1 + 0.04 (range. 0.7—1.7) mm in width. These 28 galls each contained an average of 1.8 + 0.2 (range, 1—6) second instars of O. palpalis (Fig. 10D). Two of these galls (7%) contained two and three larvae clustered together in a central, com- mon, open cavity. The remaining 26 galls contained one or more second instars in separate, open, ovoidal cells (Fig. 10D) that averaged 1.3 + 0.06 (range, 0.6—2.0) mm in length and 0.8 + 0.04 (range, 0.4—1.1) mm in width. Thus, the galls grew little on average during the second stadium, as did the cells containing individual second in- stars of O. palpalis (Fig. 10D). Ten galls were recorded to contain dead larvae of R.

549

florella or none, the latter of which was pre-

sumed to reflect complete consumption by O. palpalis, but again, cecidomyid inci- dence was not recorded in another 10 of these galls, so the incidence of cecidomyiid mortality may actually have been higher. Oxyna palpalis also overwinters as third instars in southern California (Figs. 1OE, F). This is the stage during which the greatest amount of larval growth and attendant in- crease in gall cavity size takes place (Figs. 1OE, F). Seventy-eight galls containing third instars of O. palpalis averaged 8.9 + 0.2 (range, 5.1—-13) mm in length by 7.4 + 0.2 (range, 2.1—4.5) mm. The linear leaves laterally surrounding the galls averaged 5.3 + 0.2 (range, 2.2—10.8) mm in length and 1.2 + 0.03 (range, 0.9-1.7) mm in width. Thus, on average, galls increased little, if any, in size during the equally slow growth of the third instars during the winter through early spring, when in response to renewed plant growth following winter rainfall, the cavities that contained the third instars expanded to accomodate the fast- growing, O. palpalis larvae. The 65, frass- lined, ovoidal or irregularly shaped, cen- trally located, open cells that contained sin- gle instars of O. palpalis within these 78 galls averaged, 2:6 = O31 @ange; 1yl=5-1) mm in longest measurement by 1.7 + 0.1 (range, 0.57—2.9) mm in shortest measure- ment, or about twice as large as cells con- taining single second instars. Moreover, in galls containing two or more third instars (Figs. 10E, F) these central cavities aver- aged 3.3 + 0.2 (range, 1.7—5.1) mm in lon- gest length by 2.6 + 0.2 (range, 0.85—4.6) mm in shortest width, or, again, more than twice as large on average than cavities con- taining single second instars, or those with single third instars, sometimes occupying much of the interior of smaller galls. All told, the 78 galls contained an average of 1.7 + 0.1 (range, 1—5) third instars (Figs. 1OE, F). The lateral thickness of the walls of these galls averaged 1.8 + 0.6 (range, 1.1—2.9) mm, which afforded insufficient spatial protection from parasitoids (see be-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Life stages of Oxyna palpalis in galls of Rhopalomyia florella: (A) partly grown gall of R. florella that contained egg of O. palpalis, (B) egg of O. palpalis (arrow) in gall of R. florella, (C) newly eclosed, first instar of O. palpalis (arrow) tunneling into center of gall, (D) two, overwintering second instars of O. palpalis in separate chambers (arrows), (E) four third instars of O. palpalis in common, central cell in gall, (F) 1—newly

formed puparium and 2—full-sized larva of O. palpalis, and 3—pupa of R. florella in same gall. Lines = 1 mm.

low) that attacked the third instars and pu- paria of O. palpalis. All told, third instars were found in samples over an 8-month pe- riod, 6 months into which, puparia first ap- peared in monthly gall samples. Pupa.—Near the end of the third larval stadium, the third instar transformed into a preparium (Fig. 9A), which in O. palpalis was of relatively short duration. No window to facilitate future adult egress from the

galls was formed in the gall wall, unlike the windows made by O. aterrima third instars prior to prepupal formation (Goeden 2002b). The prepuparia of O. palpalis (Figs. 9A, 10F-2) transformed into puparia within the larval cells or among the cottony tomentum (Fig. 1OF-1), which by then also contained pupae (Fig. 1OF-3) or empty pu- pal exuviae of emerged, surviving R. flo- rella, usually found on the periphery of a

VOLUME 104, NUMBER 3

51

Fig. 11.

gall (Fig. 1OF-3). The anterior end of the puparium of O. palpalis usually faced dis- tad, away from the base of the gall (Figs. 10F-1, 11). Eighty-four mature galls (Figs. 11B, C) sampled over a 2-mo period con- tained an average of 1.5 + 0.1 (range, 1—5) puparia of O. palpalis. These galls exter- nally averaged 9.1 + 0.2 (range, 5.7—15.0) mm in length by 7.9 + 0.2 (range, 5.0- 11.4) mm in width. The linear leaves lat- erally surrounding the galls averaged 7.1 + 0.3 (range, 2.6—-16) mm in length and 1.4

Life stages of Oxyna palpalis in galls of Rhopalomyia florella, continued, (A) two puparia of O. palpalis in gall, (B) full-size gall that contained puparia of O. palpalis, (C) single central puparium of O. palpalis in gall, (D) single, central pupa of R. florella in gall, (E) male adult of O. palpalis, (F) female adult of O. palpalis. Lines = 1 mm.

+ 0.03 (range, 0.8—2.5) mm in width. Sixty (71%) of the 84 galls each contained a frass-lined, ovoidal or irregularly shaped, central cavity that held a single puparium of O. palpalis (Fig. 11C), which averaged 3.4 + 0.1 (range, 2.3—6.3) mm in longest measurement by 2.0 + 0.1 (range, 1.1—3.4) mm in shortest measurement. Whereas, in galls containing two or more puparia (Fig. 11A) these central cavities averaged 4.1 + 0.2 (range, 2.9—-5.7) mm in longest length by 3.2 + 0.1 (range, 2.3—4.6) mm in short-

S52

Table 1. Incidence of Oxyna palpalis in galls of Rhopalomyia florella on Artemisia tridentata on sam- ple dates indicated.

R. florella Galls Sampled

No. Without No. With Sample Date O. palpalis (%) O. palpalis (%)

10.iv.1996 49 (84) 9 (16) l.v. 1996 51 (84) 10 (16) 7.v. 1996 55 (79) LES (2119) 5.vi. 1996 u ils) (2) 18.vi. 1996 12 (38) 20 (62) 16.vi1. 1996 15 (56) 12 (44) 13.vili. 1996 p25) (7/\\)) 10 (29) 16.1x. 1996 76 (87) MGS) 16.x. 1996 35 (81) 8 (19) 13.x1.1996 60 (79) 16 (21) 18.x11. 1996 40 (70) 17 (30) Lone 997 20 (71) 8 (29) 12997 34 (63) 20 (37) 1S 997 65 (81) Pas) (UL) Sits IL 76 (76) 24 (24) Guan 997 69 (78) 19 (22) 30.iv. 1997 45 (80) 11 (20)

est width, again, sometimes occupying much of the interior of smaller galls. The central location of a single pupa of R. flo- rella in a gall without O. palpalis (Fig. 11D), when compared to a similarly located puparium of O. palpalis typical of the 60 galls containing single individuals noted above (Fig. 11C), most of which lacked any sign of R. florella, strongly suggests a high incidence of facultative, or at least acciden- tal predation by the tephritid on the ceci- domyiid. Table | records the incidence of O. palpalis in galls of R. florella on differ- ent sample dates.

Adult.—Adults exited galls presumably by pushing aside, between, and outward through the enfolding, apical, immature leaves and tomentum (Figs. 11A, C). Under insectary conditions, 17 males (Fig. 11E) lived an average of 48 + 5 (range, 21-84) days, and 12 females (Fig. 11F) an average of 41 + 6 (range, 12-75) days. These lon- gevities are relatively long in duration for a circumnatal tephritid (Headrick and Goeden 1998). For example, males and females of

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

O. aterrima on average respectively lived half as long (Goeden 2002b).

Mating behavior.—The premating, mat- ing, and postmating behaviors of O. pal- palis were not studied in the field, nor were these behaviors observed in petri dish are- nas of the type found to be so useful with many other nonfrugivorous, tephritid spe- cies (Headrick and Goeden 1994, Goeden 2002b). In two such arenas, adults exhibited a few behaviors similar to O. aterrima (Goeden 2002b) and typical of other cir- cumnatal, gallicolous species, cf., Proceci- dochares, previously studied in southern California, i.e., a lack of courtship behavior, the exhibition of enantion type wing move- ments by both sexes, and male stalking of females prior to mating (Goeden and Norr- bom 2001; Headrick and Goeden 1994, 2000).

Seasonal history.—Oxyna palpalis is a univoltine, circumnatal species (Headrick and Goeden 1994, 1998) reproducing in ro- sette galls of Rhopalomyia florella of ter- minal buds on branches of A. tridentata in southern California (Gagné 1989). Eggs are laid in nearly fully formed galls containing young larvae of R. florella in early summer (June) and O. palpalis passes the summer (June-September) as first instars. These lar- vae molt to the second instar beginning in mid-September in early fall, and to the third instar beginning in mid-October, initially overwintering as both second and third in- stars, but solely as third instars by mid-win- ter (February) of the following year. The third instars complete this instar beginning in early spring (March) at the time of the renewed flush of host-plant growth. Pupar- iation and adult emergence follow in April and the adults mate and probably oviposit in newly formed galls of R. florella on or near the same plants from which both spe- cies of flies emerged.

Natural enemies.—Many individual Eur- ytoma sp. (Hymenoptera: Eurytomidae) and six individuals of Eupelmus sp. (Hymenop- tera: Eupelmidae) were reared separately from individual puparia as primary, soli-

VOLUME 104, NUMBER 3

tary, probably larval-pupal endoparasitoids. One, three, three, and seven individuals of Lyrcus sp. (Hymenoptera: Pteromalidae) were reared from individual puparia of O. palpalis as gregarious, primary endoparas- itoids.

ACKNOWLEDGMENTS

My thanks to Andrew C. Sanders, Cu- rator of the Herbarium, Department of Bot- any and Plant Sciences, University of Cal- ifornia, Riverside, for identifications of plants mentioned in this paper. Krassimer Bozhilov in the Central Facility for Ad- vanced Microscopy and Microanalysis, University of California, Riverside, greatly facilitated my scanning electron microsco- py and digital imaging. The parasitoids ini- tially were identified by Harry E. Andersen, Huntington Beach, California, now de- ceased, and later were confirmed or further identified by John Heraty and Jung-Wook Kim, Department of Entomology, Univer- sity of California, Riverside. I also am grateful to Jeff Teerink for technical assis- tance and to David Headrick for his helpful comments on earlier drafts of this paper.

LITERATURE CITED

Baker, C. EF 1904. Reports on California and Nevadan Diptera, |. Invertebrata Pacifica 1: 17—39.

Barkley, T. M., ed. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence.

Blanc, F L. and R. H. Foote. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7: 1-117.

Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon.

Foote, R. H. and F L. Blanc. 1963. The fruit flies or Tephritidae of California. Bulletin of the Califor- nia Insect Survey 7: 1-117.

Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York.

Gagné, R. J. 1989. The Plant-feeding Gall Midges of North America. Comstock Publishing Associates, Cornell University Press, Ithaca and London.

Si nn ee)

Goeden, R. D. 2001. Life history and description of immature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occi- dentalis (Nuttall) (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 191—206.

. 2002a. Description of the immature stages of Tephritis stigmatica (Coquillett) (Diptera: Tephri- tidae). Proceedings of the Entomological Society of Washington 104: 335-347.

. 2002b. Life history and description of im- mature stages of Oxyna aterrima (Doane) (Dip- tera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 104: 510-526.

Goeden, R. D. and A. L. Norrbom. 2001. Life history and description of adults and immature stages of Procecidochares blanci, n. sp. (Diptera: Tephriti- dae) on /socoma acradenia (E. Greene) E. Greene (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 517-540.

Goeden, R. D. and J. A. Teerink 1999. Life history and description of immature stages of Trupanea vicina (Wulp) (Diptera: Tephritidae) on wild and cultivated Asteraceae in southern California. Pro- ceedings of the Entomological Society of Wash- ington 101: 742-755.

Headrick, D. H. and R. D. Goeden. 1994. Reproduc- tive behavior of California fruit flies and the clas- sification and evolution of Tephritidae (Diptera) mating systems. Studia Dipterologica 1(2): 194— 22,

——.. 1998. The biology of nonfrugivous tephritid fruit flies. Annual Review of Entomology 43: 217-241.

. 2000. Behavior of flies in the subfamily Te- phritinae. Chapter 2. Jn Aluja, M. and A. L. Norr- bom, eds. Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, London, New York, and Washington, D.C.

Hickman, J. C., ed. 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles.

Novak, J. A., W. B. Stolzfus, E. J. Allen, and B. A. Foote. 1967. New host records for North Ameri- can fruit flies (Diptera: Tephritidae). Proceedings of Entomological Society of Washington 69: 146— 148.

Quisenberry, B. F 1949. The genus Oxyna in the Ne- arctic Region north of Mexico. Pan-Pacific Ento- mologist 25: 71-76.

PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 554-562

BIOLOGICAL OBSERVATIONS OF CENTISTES GASSENI SHAW (HYMENOPTERA: BRACONIDAE), A PARASITOID OF DIABROTICA SPP. (COLEOPTERA: CHRYSOMELIDAE)!

R. EF W. SCHRODER AND M. M. ATHANAS

(RFWS) Research Entomologist, retired, (MMA) Entomologist, Insect Biocontrol Lab- oratory, U.S. Department of Agriculture, Agricultural Research Service, Bldg. 011A, Rm. 107, BARC-West, Beltsville, MD 20705, U.S.A. (e-mail: athanasm @ba.ars.usda.gov)

Abstract.—In 1992-1993, Centistes gasseni Shaw was imported into the United States, and in the laboratory successfully parasitized: southern corn rootworm, Diabrotica un- decimpunctata howardi Barber; banded cucumber beetle, D. balteata LeConte; western corn rootworm, D. virgifera virgifera LeConte; and striped cucumber beetle, Acalymma vittatum (FE). Males and females of C. gasseni lived an average of 15.4 and 12.9 days (with a maximum of 30 and 29 days), respectively. A single female oviposited in 383 host Diabrotica over her lifetime, from which 158 cocoons were recovered. Additional

observations on the biology and rearing of the parasitoid are presented.

Key Words:

Acalymma vittatum, biological control, Centistes gasseni, Diabrotica bal-

teata, Diabrotica barberi, Diabrotica speciosa, Diabrotica undecimpunc- tata howardi, Diabrotica virgifera virgifera, host specificity

The northern corn rootworm (NCR), Diabrotica barberi Smith and Lawrence, and the western corn rootworm (WCR), Diabrotica virgifera virgifera LeConte are the most damaging and costly pests of corn in North America (Sutter and Lance 1991). To achieve effective control of these chrys- omelids, soil insecticides routinely have been used on 50—60% of the corn acreage, or 12-16 million ha annually (Metcalf 1986). These insecticides are generally ap- plied prophylactically and are frequently unnecessary (Lance and Sutter 1992). Ac- cording to Metcalf (1986), the cost from crop loss and treatment due to the corn rootworm (CRW) complex is approximate- ly one billion dollars per year.

'This article reports the results of research only. Mention of a proprietary product does not constitute an endorsement or a recommendation by the USDA for its use.

In addition, southern corn rootworm (SCR), Diabrotica undecimpunctata ho- wardi Barber; striped cucumber beetle, Acalymma vittatum (FE); western striped cu- cumber beetle, Acalymma trivittatum (Man- nerheim); and banded cucumber beetle, Diabrotica balteata LeConte cause 50—100 million dollars in damage to other crops (Cucurbitaceae, Fabaceae) (Metcalf et al. 1962).

Lance and Sutter (1992) cite health risks to growers, livestock and wildlife due to soil insecticides used for CRW _ control. They also refer to reports of these insecti- cides being detected in ground and surface water. Therefore, it is evident that there is a need for alternative control measures that are environmentally safe and cost effective.

In 1990, while searching for biocontrol agents of Colorado potato beetle (Coleop- tera: Chrysomelidae) in Rio Grande do Sul,

VOLUME 104, NUMBER 3

Brazil, we learned of the existence of an undescribed braconid parasitoid of the adult Neotropical leaf beetle, Diabrotica speciosa Germar (Gassen 1986). It was subsequently described by Shaw (1995) as Centistes gas- seni Shaw, a solitary endoparasitoid of the adult. As there are no known effective par- asitoids of the CRW complex in the United States, we were interested in acquiring and studying C. gasseni as a potential new bi- ological control agent. We traveled to Bra- zil in 1992—93, and, in cooperation with Dirceu Gassen, EMBRAPA-CNPT, collect- ed and imported adults of D. speciosa par- asitized by the braconid. In this article, we present observations on the biology, rear- ing, mating behavior and host specificity of C. gasseni, obtained during initial attempts to colonize the parasitoid in quarantine.

MATERIALS AND METHODS

Approximately 6,000 D. speciosa adults were collected from April 24—26, 1993, at the University of Passo Fundo, Passo Fun- do, Rio Grande do Sul, Brazil. Collections were made primarily from potato plants, but some adults were also collected from near- by plots of pepper and sweet potato plants. Parasitoids were also recovered from bee- tles collected from flowering weeds and wild cucurbit plants, 98 km west of Passo Fundo. Sub-samples of field collected bee- tles were dissected to determine percent parasitism and the number of parasitoids per host. In subsequent collections, Hei- neck-Leonel and Salles (1997) recovered C. gasseni from D. speciosa collected in corn, beans, melon, cucumber, cabbage, broccoli, spinach and lettuce.

Field collected adult beetles were held in 30.5 cm collapsible aluminum screen cages (BioQuip® Products, Gardena, CA). Each cage was inverted and the hinged top served as the bottom. A fitted piece of moistened felt was placed on the bottom as a liner, and covered with a ca 0.5—0.6 cm layer of moistened sand with a 2.5 cm sand- free margin on all sides. The sand and felt served as a substrate in which the parasitoid

Si Nn nN

larvae spun cocoons. Approximately one cm above the bottom of the cage, we fitted a pre-cut piece of number 12 mesh stainless steel screen, that separated the adult Dia- brotica from emerging C. gasseni larvae, which passed through to pupate.

The adult beetles, and 31 parasitoid co- coons recovered prior to departure, were taken to the Maryland Department of Ag- riculture quarantine facility, Annapolis, MD, and held at 21.1°C, 60% RH, and a 16:8 LD photoperiod. The host beetles were kept in the modified cages which were ser- viced three times a week. Parasitoid co- coons were carefully removed from the sand and placed in petri dishes, along with a moistened, 3.8 cm cotton roll to maintain humidity.

The containers used for oviposition, and to house the C. gasseni adults, were plex- iglass tubes (7.6 cm in diameter and 10.2 cm long) modeled after those used by Whistlecraft et al. (1984) in their braconid rearing program. To confirm mating, a sin- gle female was confined in an aspirator tube with multiple males until copulation was observed. With Cotesia melanoscela (Ratz- eburg), Weseloh (1977) found that 20 sec- onds was necessary to successfully transfer sperm to the female. We therefore consid- ered a successful mating session as one in which copulation was observed for 20 sec- onds or more. We used several of the meth- ods discussed by Singh (1982) to induce successful mating, including shaking the in- sects to the bottom of the tube, increasing the number of males per female, and peri- odically chilling the females prior to ex- posure to the males.

The following procedures were used in quarantine to conduct studies on the biolo- gy, mating behavior, and host preferences of the parasitoid.

Host exposure methods.—I/ndividual ex- posure: To expose host Diabrotica adults individually, we placed a parasitoid female into a 35 X 10 mm polystyrene culture dish and introduced a single Diabrotica adult. The host beetle was observed until ovipo-

556

sition occurred, then it was removed. Ten to twenty Diabrotica were exposed to a fe- male per session. Following each session, the female was held for one hour in a tube or petri dish, and given access to water and honey solution. Each female had no more than two exposure sessions per day. A sep- arate cage (#1—3) was set up for the Dia- brotica exposed to each of the three, mated C. gasseni females used.

Group exposure with immediate remov- al: Approximately 35 Diabrotica beetles were placed in an oviposition tube, into which a C. gasseni female was then intro- duced. As soon as a host beetle was para- sitized, it was aspirated out of the tube. As above, following each session females were permitted to rest, and limited to two ses- sions per day. A separate cage (#4—6) was set up for the Diabrotica exposed to each of the three, mated C. gasseni females used.

Group exposure for one hour: Six C. gasseni females were used to sting a total of 394 BCB over a 7-day span. The females were confirmed mated, and all had emerged in association with other males and females. Beetles were exposed in groups of 15—20 to individual females for one hour, then placed into a single cage (#7).

Group exposure for four hours: Six C. gasseni females, selected from a group of females, were used to sting a total of 140 BCB over a 4-day span. All the females had been confined with an equal number of males (17 6 & 17 2) for 72 hours preced- ing exposure to Diabrotica, but mating was not confirmed. As above, all had emerged in association with other parasitoid males and females. BCB were exposed in groups of 20—25 to single C. gasseni females, for four hours duration, then placed into a sin- gle cage (#8).

Host species preference.—Adult SCR and BCB were sexed, and three males and three females of each species were placed into an oviposition tube. A C. gasseni fe- male reared from BCB host was introduced and exposed to the beetles for one hour, af- ter which time she was removed and held

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

as previously described. Using nine C. gas- seni females, a total of 792 Diabrotica were exposed.

Host sex preference.—To determine if C. gasseni exhibited a preference for either male or female hosts, SCR reared parasit- oids (n = 10) were exposed individually to six male and six female SCR for one hour. One group of hosts (204 2 & 6, cages | & 2, respectively) was exposed to C. gas- seni females 1—3 days old, another group of hosts (372 2 & 6, cages 3 & 4, respec- tively) was exposed to C. gasseni females, the majority of which were I—5 days old. Two cages of unexposed male and female Diabrotica were maintained as controls.

Host specificity—Host specificity tests were conducted to determine whether SCR, WCR, BCB, the related SCB and Cerotoma trifurcata (FE) were suitable hosts for C. gasseni. The criteria for these tests were based on observing the parasitoids’ ovipo- sitional behavior for one hour after expo- sure to the hosts, and then holding parasit- oid and hosts together (25—50 per exposure) over night. The female was then removed and the hosts held for parasitoid emergence for 30 days.

Additional non-target insects that may occur in the same habitat, including bene- ficial and phytophagous coccinellids were also exposed to C. gasseni to determine their suitability as hosts. In each test, a sin- gle C. gasseni female, less than seven days old, was placed in an oviposition tube. Five Diabrotica (BCB or SCR), which served as controls, and 25—50 adult host beetles (hosts were obtained from laboratory and field sources) were then introduced. Each tube was observed for one hour to confirm any oviposition that occurred. The Diabro- tica were removed after they were stung and set up in separate parasitoid emergence containers. After one hour, observations were discontinued, and females were kept with the test beetles for 24 hours. Host bee- tles were then removed, and held for para- sitoid emergence. Thirty days after expo-

VOLUME 104, NUMBER 3

Mable: 41.

speciosa.

nN nN ~~

Emergence, sex ratios and hyperparasitism of Centistes gasseni from field collected Diabrotica

Sex Ratio No %o Centistes

Date No. Cocoons No No ; Mesochorous Emergence 4/92 1LO8 1] 19 1:2 12 28 4/93 Bil 5 6 {<1 2 és) 5/93 338 138 64 Zo) 3 60 5/94 LO] 10 13 1h 2) 23 6/944 rele) 8 8 Le] 14 19

“Data from cocoons collected at the ARS, South American Biological Control Laboratory, Buenos Aires,

Argentina.

sure, host beetles were dissected to deter- mine if they had been parasitized.

RESULTS

From the 31 cocoons brought back from Brazil, 11 C. gasseni and two hyperparasi- toids, Mesochorous sp. (Hymenoptera: Ich- neumonidae) emerged. From the ~ 6000 D. speciosa collected in Brazil, we recovered 338 C. gasseni cocoons, 66 C. gasseni lar- vae, and 11 puparia of Celatoria bosqui Blanchard (Diptera: Tachinidae) (Table 1). As the number of host beetles collected was approximated, we could not accurately de- termine the percent parasitism. However, in one sub-sample (n = 111) 10.0% were par- asitized. In Pelotas, RS, Brazil, Heineck- Leonel and Salles (1997) reported that par- asitism, of D. speciosa by C. gasseni, ranged from 0-18.9% (* = 8.7, SD = 7.1) in samples obtained at monthly intervals from May 1994—April 1995.

Mating data were obtained on a sub-sam- ple of P and F generation parasitoid fe- males. Forty-one percent of the P sample (n 17) successfully mated (Table 2). All C.

gasseni pupae obtained from D. speciosa were set up individually, thus all females that emerged were isolated from other males and females. Of those that mated, 29% did so on the first attempt, 43% on the second, and 28% required three or more at- tempts to mate. Unsuccessful mating at- tempts were observed for a minimum of 10 minutes. Seventy-one percent of the F sam- ple (n = 90) successfully mated (Table 2). Of these, 72% occurred on the first attempt, 20% on the second, and 8% required three or more attempts. In the majority of cases, the female would aggressively fight off the advances from any males after she had suc- cessfully mated. Nevertheless, 9% of the fe- males did mate a second time, all within 2.1 minutes of the first. When we exposed pre- viously mated females to new males, 8% mated a second time.

Parasitoid female progeny were obtained only from positively mated parental fe- males. When confined with a suitable host, the female mounted the beetle from the side and thrust her ovipositor into the abdomen. The beetle responded to the attack by trying

Table 2. Reproductive behavior of Centistes gasseni reared from native and alternative host Diabrotica. %1so2 % % & % 2 with Mean % Mean ? with & with d 2 &o Mean Expos. Mean Length Mean No. Mean Age 6 Host 2 Mated Age (Days) Mated Mated Mated Mated (Min) (Sec) 3 Used (Days) Pgen> 41 Dalia lesry 1) N/A N/A N/A 32852) S20) 93102428 eee 4S Fa °4 BESS Al LES == OLO88 87:8 86 68 57 Spe ee EI BY PS) series "ete 22 I 2.4 + 0.4

“Tsolated from other females and males. °C. gasseni reared from D. speciosa.

°C. gasseni reared from North American Diabrotica.

558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Parasitism by Centistes gasseni using various host exposure methods. Days ? ? Emerged Hosts No. %e No. 3 No. @ % Cage in Use with 3, 9 Exposed Cocoons Parasitism Emerged Emerged Emerged

1 1—5 None 103 SCR? 62 60 q 39 74 2, 1-2 296 12 61 SCR 45 80 Si 0) 82 3p 1=3 296 12 47 SCR 28 60 14 1 54 4 2-13 aXe) 3) 232 SCR 154 73} 102 0) 66 5 5-9 36 32 129 BCBs 61 62 D1 0) 93 6° 2-12 le 383 BCB 158 52 93 0) 59 7 1-9 — 394 BCB 163 52 68 14 50 8 4—6 — 140 BCB 67 64 36 5) 61

“Cages 1-3, hosts exposed individually, cages 4—6, hosts exposed in groups with immediate removal, cage 7 hosts exposed for one hour, cage 8 hosts exposed for four hours. > Female did not mate until 37 of the 47 hosts had been exposed.

© Female mated twice.

4 Southern corn rootworm, Diabrotica undecimpunctata howardi Barber.

© Banded cucumber beetle, D. balteata LeConte.

to brush the C. gasseni from its back, be- coming highly active or agitated, or by dropping to the bottom of the container.

Dissections of field collected D. speciosa revealed that parasitized beetles contained a single C. gasseni larva per host. However, laboratory dissections indicated that super- parasitism did occur, with only one larva completing development. At 21.1 + 0.5°C, the parasitoid would complete its life cycle in 33—41 days (x = 34.7 + 1.7 days). At 23.9 + 0.5°C, it took 26-32 days (* = 27.8 10.7, days): At 21.) =O "Ce sasseni females and males lived an average of 12.9 25/0 days and 15/4 =" 2.8 "days; respec- tively.

Host exposure methods.—I/ndividual ex- posure: In the hosts exposed individually (cages 1-3), the mean percent parasitism, based upon the number of hosts exposed and the number of parasitoids (cocoons and/or larvae) recovered, was 66.7% (SD = 11.54), and the mean parasitoid emergence from the resultant cocoons was 70% (SD = 14.4) (Table 3). At 32 + 5 days post ex- posure, 89.3% (SD = 5.03) of the host SCR were dead. The female of cage 1, which emerged alone, produced offspring with a sex ratio of 6 2:1 d, the highest ratio of females to males obtained in this study.

Group exposure with immediate remov- al: The females of cages 4 and 6 oviposited

in 232 SCR and 383 BCB respectively, over a 12 day period (Table 3). These yield- ed 154 and 158 cocoons, respectively, and 102 males and 93 males, respectively. This was the largest number of hosts stung, and number of cocoons and progeny obtained from individual C. gasseni females.

Pre-cocoon mortality was higher in the BCB hosts, averaging 22.0% versus 8.5% in the SCR. Eliminating cage 6 from the computations, since this female was ex- posed to an excessive number of hosts to determine oviposition limits, percent para- sitism using this method of exposure aver- aged 67.5% (SD = 7.78). The mean para- sitoid emergence from the cocoons was 72.6% (SD = 18.0), and at 29 + 4 days post exposure, 91.5% (SD = 3.5) of the ex- posed hosts were dead.

Parasitoid females 10—12 days old, were the oldest from which viable progeny were recovered. The oldest age attained by a C. gasseni male and female, was 30 and 29 days, respectively. Over the course of the female’s life, she was exposed to 114 SCR, both in groups and individually. As she aged, all hosts were exposed individually. On her 24th day, she stung two SCR, this was the oldest age at which we observed attack behavior. A cocoon was recovered from a SCR parasitized by a female 14—24 days old. There was no emergence from the

VOLUME 104, NUMBER 3

Table 4. Host specificity of Centistes gasseni.

Para. Emerge from Host

Host Species Tested

Para. Dissected from Host

Para. Emerge from Control

Coccinellidae Coleomegilla maculata (DeGeer) Hippodamia variegata (Goeze) Propylea quatuordecimpunctata (L.) Epilachna varivestis Mulsant Cycloneda munda (Say) Chilocorus stigma (Say) Hippodamia convergens Guerin Harmonia axyridis (Pallas)

Chrysomelidae Cerotoma trifurcata (F.) Acalymma vittatum (FP.)* Diabrotica balteata LeConte* D. undecimpunctata howardi Barber* D. virgifera virgifera LeConte*

| +++4¢4+4+ 44

++ ++

“The Acalymma and Diabrotica tested were suitable hosts, dissections and controls were therefore not used.

cocoon, and a male was subsequently dis- sected out. This proved to be the oldest C. gasseni from which both cocoon, and fully formed offspring have been recovered.

Group exposure for one and four hours: The percent parasitism obtained by expo- sure for one (cage 7) and four (cage 8) hours, was 52% and 64%, respectively (Ta- ble 3). From cage 7, 82 parasitoids with a sex ratio of 5 d:1 2, were recovered. From cage 8, we recovered 41 parasitoids with a 7 3:1 2 sex ratio. This was the first in- stance in which parental females that were not confirmed mated, produced female off- spring. Pre-cocoon mortality averaged 23% for cages 7 and 8. Parasitoid emergence from the cocoons was 50% and 61% for cages 7 and 8, respectively.

Host species preference.—Although all parental C. gasseni used were BCB reared, they did not exhibit a preference for BCB hosts. The percentage parasitism averaged 32% in both the BCB and SCR. Fifty-seven parasitoids, including seven females, were recovered from SCR hosts, and 55 parasit- oids, including three females, were recov- ered from BCB hosts. All C. gasseni female offspring were recovered from BCB fe- males and SCR males. Pre-cocoon mortal-

ity was 14% for both SCR and BCB. Par- asitoid emergence from cocoons was 51% for BCB males and females, 65% for SCR males and 42% for SCR females.

Host sex preference.—In cages | and 2, the percent parasitism for female and male hosts was 32% and 23%, respectively. In cages 3 and 4, it was 45% and 38% for the female and male hosts, respectively. Emer- gence averaged 70% from the cocoons ob- tained from female hosts, and 78% from co- coons obtained from the males. For both C. gasseni age groups, all female offspring were recovered from female SCR hosts.

Beetle mortality appeared to be excessive with this generation: pre-cocoon mortality averaged 46% for cages | and 2, and 53% and 60% for cages 3 and 4, respectively. Thus it is uncertain whether the recovery of female parasitoids from female hosts was due to parasitoid preference, or selective mortality.

Host specificity—The SCR, WCR, BCB and SCB exposed to C. gasseni were suc- cessfully parasitized, resulting in the recov- ery of parasitoids from all four hosts. Nei- ther Cerotoma trifurcata (FE), nor any of the exposed coccinellids were parasitized (Ta- ble 4). In every test, given the choice be-

560

tween the control Diabrotica and the test insect, C. gasseni always searched for, and attacked the Diabrotica first, indicating that the parasitoid was active. The time it took to parasitize all control Diabrotica varied from 10-30 minutes. C. gasseni females approached the test beetles numerous times during the observation period. Usually, they would rapidly tap their antennae on the bee- tle, then move on to another host. Occa- sionally the female would mount the test beetle and probe with the tip of the abdo- men at the same location where she would normally oviposit.

DISCUSSION

The survival of C. gasseni larvae found during the maintenance of the host cages was extremely low, regardless of whether they were returned to the original cage or set up in petri dishes containing moistened sand. Vance (1932) observed that any Che- lonus annulipes Wesmael larvae that emerge from the host and completes its fi- nal feeding, is unable to construct a cocoon if it is disturbed in any way. Harrison et al. (1993) observed a similar situation with lar- vae of Microplitis croceipes (Cresson). Those larvae died within two hours of ex- iting the host if they had not spun cocoons by then.

When inducing C. gasseni to mate, most of the successful copulations occurred when modified, open-ended aspirator tubes were used, and an air current provided. This seemed to aid the males in orienting to the virgin females (Vinson 1978). This suggests the presence of a sex pheromone in C. gas- sent (Matthews 1974). In several cases in which females did not mate, or mating oc- curred after multiple attempts, the males used were =24 hours old. Laing and Cal- tagirone (1969) observed that females of the braconid Habrobracon lineatellae (Fi- scher) will not mate until they are at least 24 hours old, and that insemination is more likely to occur when older males (5—6 days old) are used. As Schlinger and Hall (1960) reported that the sperm supply drops rap-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

idly with each successful mating, whenever possible, we removed each male after it had mated.

The host beetle apparently dies soon after parasitoid emergence; when harvesting C. gasseni cocoons we would find almost an equal number of dead beetles. Loan and Holdaway (1961) observed that the curcu- lionid, Sitona sp., would stop laying eggs 1—2 days after being stung, and usually die in 3—4 hours following emergence of the endoparasitic braconid larva, Pygostolus

falcatus (Nees). No female parasitoid prog-

eny were obtained using group exposure of hosts. The large host to parasitoid ratio (Grinberg and Wallner 1991), and rapid rate of oviposition (Flanders 1956) may have been factors. When exposed to hosts in groups, the females would oviposit at a rate of one host per 0.92 + 0.38 min. When presented with hosts individually, the rate slowed to one host per 2.2 + 0.78 min.

Centistes gasseni was successfully reared through nine generations. During this peri- od, the sex ratio varied widely, ranging from all males to 6 2:1 3d. An adequate number of females was obtained to conduct experiments and propagate the colony. By the tenth generation, only males were pro- duced. Although our results showed that fe- males contacting each other did not pre- clude the production of female offspring, it may have affected the sex ratio. With the scelionid Trissolcus, contact with other fe- males of the same species (Viktorov 1968), or trace pheromones of other females (Vik- torov and Kochetova 1973, Buleza 1975), resulted in production of significantly more male offspring. In addition, the fluorescent lighting utilized in quarantine may have hindered successful mating. Nealis and Fra- ser (1988), observed that the braconid Apanteles fumiferanae (Viereck), mated more frequently under natural rather than artificial light conditions.

The SCR, WCR, BCB, and SCB were readily attacked and parasitized by C. gas- seni, Whereas other phytophagous and ben- eficial beetles tested proved unsuitable as

VOLUME 104, NUMBER 3

hosts. Due to the small number of C. gas- seni females and availability of host beetles, the specificity tests were limited in scope, but generic specificity was evident.

Centistes gasseni has potential for use as a biocontrol agent of the North American CRW complex, and the striped cucumber beetle, A. vittatum. The biology, population dynamics and behavior of C. gasseni needs to be studied on its native host D. speciosa in Brazil.

ACKNOWLEDGMENTS

We gratefully acknowledge the assistance and cooperation of: Ann Sidor, USDA, In- sect Biocontrol Laboratory, Beltsville, Maryland; Guillermo Cabrera Walsh, USDA, Agricultural Research Service, South American Biological Control Labo- ratory, Buenos Aires, Argentina; Dirceu Gassen, EMBRAPA-CNPT, Passo Fundo, RS, Brazil; the Maryland Department of Agriculture, Annapolis, Maryland; and the USDA Vegetable Research Laboratory, Charleston, South Carolina.

LITERATURE CITED

Buleza, V. V. 1975. Influence of trace pheromones on the sex ratio in some species of Trissolcus, pp. 15—19. In Pristavko, V. P., ed. Insect Behavior as a Basis for Developing Control Measures Against Pests of Field Crops and Forests. Amerind Pub. Co., New Delhi, vii + 238 pp.

Flanders, S. E. 1956. The mechanisms of sex ratio reg- ulation in the parasitic Hymenoptera. Insectes So- ciaux 3: 325-334.

Gassen, D. N. 1986. Parasitos, patogenos e predadores de insetos associados a cultura do trigo. EMBRA- PA-CNPT Circular Tecnica No. 1, 86 pp.

Grinberg, P. S. and W. E. Wallner. 1991. Long term evaluation of Rogas lymantria: A braconid endo- parasite of the gypsy moth, Lymantria dispar. En- tomophaga 36: 205-212.

Harrison, W. W., D. A. Herbert, and D. D. Hardee. 1993. Effect of parasitoid and host age on ovi- position and emergence of Microplitis croceipes (Hymenoptera: Braconidae) an endoparasitoid of Helvicoverpa zea (Lepidoptera: Noctuidae). Jour- nal of Entomological Science 28: 343-349.

Heineck-Leonel, M. A. and L. A. B. Salles. 1997. In- cidencia de parasitoides e patoge em adultos de Diabrotica speciosa (Germar) (Coleoptera: Chry-

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somelidae) na regiao de Pelotas, RS. Anais da So- ciedade Entomologia do Brasil 26: 81-85.

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Metcalf, R. L. 1986. Foreword, pp. 7-15. /n Krysan, J. L. and T. A. Miller, eds. Methods for the Study of Pest Diabrotica. Springer-Verlag, New York, 260 pp.

Metcalf, C. L., W. P: Plint, and Ri. Metcalf. 1962: Destructive and Useful Insects. 4th edition. Mc- Graw-Hill, New York, 1087 pp.

Nealis, V. G. and S. Fraser. 1988. Rate of development, reproduction, and mass-rearing of Apanteles fu- miferanae Vier. (Hymenoptera: Braconidae) under controlled conditions. Canadian Entomologist 120: 197-204.

Schlinger, E. I. and J. C. Hall. 1960. The biology, be- havior, and morphology of Praon palitans Mue- sebeck, an internal parasite of the spotted alfalfa aphid, Therioaphis maculata (Buckton) (Hyme- noptera: Braconidae, Aphidiinae). Annals of the Entomological Society of America 53: 144—160.

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Chrysomelidae) with a key to new world species. Proceedings of the Entomological Washington 97: 153-160.

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the braconid Chelonus annulipes Wesm., a para- site of the European corn borer. United States De- partment of Agriculture, Technical Bulletin No. 294, 48 pp.

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of trace pheromones in the regulation of sex ratio moth parasite, Apanteles melanoscelus. Annals of in Trissolcus grandis (Hymenoptera: Scelionidae). the Entomological Society of America 70: 549— Zhurnal Obshchei Biologii 34: 559-562. 554.

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PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 563-570

A NEW SPECIES OF EUTARSOPOLIPUS BERLESE (ACARI: PODAPOLIPIDAE) FROM THE GALAPAGOS ISLANDS, A PARASITE OF AGONUM CHATHAMI VAN DYKE (COLEOPTERA: CARABIDAE)

ROBERT W. HUSBAND

1035 Scottdale Drive, Adrian, MI 49221, U.S.A. (e-mail: rhusband @ adrian.edu)

Abstract.—Eutarsopolipus brettae, n. sp. (Acari: Podapolipidae), is described from Agonum chathami Van Dyke (Coleoptera: Carabidae) from the Galapagos Islands and compared with related species of Eutarsopolipus. Characters of E. brettae do not fit with

existing subgroups of Eutarsopolipus.

Key Words:

Mites in the family Podapolipidae (Acari: Heterostigmata) are highly specialized ecto- and endoparasites of the insect orders Blat- taria, Orthoptera, Heteroptera, Hymenop- tera, and especially Coleoptera. The genus Eutarsopolipus Berlese is restricted to hosts in Carabidae (Coleoptera) and occurs worldwide. Along with the type species, E. lagenaeformis Berlese 1913, more than 40 others have been discovered, of which 21 were described by Regenfuss (1968, 1974). This is the first record of Eutarsopolipus from the Galapagos Islands.

The purpose of this paper is to describe a new species collected from Agonum chathami Van Dyke and compare it with other species of Eutarsopolipus parasitizing the carabid genus Agonum, with Eutarso- polipus from Central and South America, and with related Eutarsopolipus worldwide.

MATERIALS AND METHODS

Males, larval and adult females, and eggs of Eutarsopolipus brettae were collected from Agonum chathami Van Dyke _ bor- rowed from the California Academy of Sci- ences, San Francisco, California, U.S.A. The technique for removing mites from mu-

beetle, Carabidae, parasitic mites, Acari, Podapolipidae, Galapagos, Ecuador

seum specimens is described in Husband and Dastych (1998).

Measurements, in micrometers (wm), were taken with the aid of a Zeiss micro- scope with a stage micrometer and drawing tube. The terminology used follows Lind- quist (1986).

Eutarsopolipus brettae Husband, new species (Figs. 1—3)

Diagnosis.—Adult female E. brettae lack stigmata, cheliceral stylets are long, 75-85, and ambulacral II, III claws are well devel- oped (10). The genital capsule of male EF. brettae has concave lateral margins. Plates C and D are separate in larval female E. brettae, cheliceral stylets are long (60—64), setae of the idiosoma and legs are long (Ta- ble 1), and the larval female has 3 femur I setae in combination with | genu I seta (Ta- ble:2):

Adult female (Fig. 1).—Gnathosoma: Length 62—69, width 60—67. Palp length 22-25; cheliceral stylet length 75-85 with basal sclerite 12-15, pharynx width 17, walls of pharynx thick, dorsal gnathosomal

564

Fig. 1.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

H 1} ii

@ 4d |p iit fit {

0.1mm

Eutarsopolipus brettae, adult female: left, ventral aspect; right, dorsal aspect.

VOLUME 104, NUMBER 3

setae 32-35, ventral setae 20—25, distance between ventral setae 18—20. No stigmata.

Idiosoma: Length 266—450, width 209— 318. Prodorsal plate setae v, 12—15 v, 7-8, sc, 25-30, setae about 10 from posterior margin of prodorsal plate. Distance be- tween setae v, 55—56; v, lateral to line con- necting v, and sc,. Plate C midlength 60— 70, width 237-265, setae c, 7-8, c, 9-10. Plate D length 50—75, width 175-197, setae

0.1mm

d 7-8. Plate EF length 40-53, width 130— 145, setae e 7-8, no setae h,, h,.

Venter: Apodemes | and 2 weakly de- veloped, meeting sternal apodeme medially; sternal apodeme not extending beyond junction with apodemes 2. Coxal setae la 3—4, 2a 5—6; setae la situated equidistant to apodemes 1, 2. Distance between setae la 37-52. Coxal setae 3a not evident, 3b 6-7.

566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ran

BS

oot

we ee eK He ‘ >

Ae, \ \ { \ \ ‘ ‘

1B, BE

Legs: Leg setation as in Table 2. Am- bulacrum I with a terminal stout claw, am- bulacra IH, II with 2 strong claws. Tarsus I subunguinal seta spinelike, 2 terminal spines on each of tarsi II, III. Tarsus I so- lenidion w 5—6; tarsus II solenidion w 5.

= (o

aha)

eer | Gann

== ae

Eutarsopolipus brettae, larval female: left, ventral aspect; right, dorsal aspect.

Tibia I solenidion 8—9, adjacent seta k 4— 6. Tibial I, II, II setae d 45-50, 30—33 and 20—22 respectively.

Male (Fig. 2).—Gnathosoma: length 37—40, width 36-37. Palp length 13-17; cheliceral stylet length 25-31, no_ basal

VOLUME 104, NUMBER 3

sclerite; pharynx width 10—11, dorsal gna- thosomal setae thick 2—4, ventral setae 9— 1S:

Idiosoma: Length 170-188, width 120— 147. Prodorsal setae v, m, v, m, setae sc, 53-65. Distance between setae v, 20-21, distance between setae v, 52; setae v, lateral to line connecting setae v, and sc,. Plates C and D fused, setae c,, v,, d, and e vestigial or not present. Genital capsule length 28— 30, width 30-35 at base, concave lateral margins.

Venter: Apodemes 1, 2 and sternal apo- deme evident; coxae III separated medially. Setae la, 2a, 3b microsetae, 3a not evident.

Legs: Leg setation as in Table 2. Am- bulacrum I with | thick claw, length 4, am- bulacra H, If] with small claws 3. Tarsus I spinelike seta s 5—6, tarsi H, III spinelike setae tc’ and u’ 5—7. Tarsus I solenidion w 5—6, tarsus II solendion w 4—5; tibia I so- lendion & 8, adjacent seta k 2—3. Tibiae I, II, IfI setae d 30-33, 15-22, 19 respective- ly. Genu III seta v” evident in only | male.

Larval female (Fig. 3).—Gnathosoma: length 44-53, width 42—45. Palp length 15—20, cheliceral stylet length 60—64, stylet basal sclerite 7—8, pharynx width 12-13, dorsal gnathosomal setae 31—36, ventral se- tae 13—14, distance between ventral setae 14-20.

Idiosoma: Length 165—192, width 125— 132. Prodorsal plate setae v, 17—20, v, 9- 16, sc, 84-85. Distance between setae v, 34—45; setae v, lateral to line connecting setae v, and setae sc,. Plates C and D sep- arate, setae c, 7-8, c, 8-10, setae d 7-8: distance between setae d 30—33. Setae c, in line with setae c,. Plate EF oval, setae e 7. Plate H broader at base, setae h, 80—90, h, 5—7, distance between setae h, 2.

Venter: Apodemes 1, 2 and sternal apo- deme conspicuous but weakly scerotized. Setae la 3—4, 2a 3-5, 3a m and 3b 5. Dis- tance between setae 3a and 3b 18—20, setae 3a visible only in larval exoskeleton asso- ciated with adult female.

Legs: Leg setation as in Table 1. Am- bulacrum I with 2 small, parallel claws.

567

Ambulacra II, HI with small, diverging claws. Tarsus I spinelike seta s 5, tarsi I, III spinelike setae tc’ and wu’ 5—7. Tarsus | solenidia o 3—5, tarsus II solenidion w 5. Tibia I solenidion @ 9-10, seta k 3. Setae tc’ 7-9, tc” 9-10. Tibiae I, II, III setae d 35-40, 18-23 and 17-25 respectively.

Egg.—Length 210-240, width 120-135.

Type, host, and locality data.—Holotype 2? (RWHO10501-1), allotype ¢ and 24 par- atypes: from Chatham Island, Galapagos Is- lands, Ecuador, from under eleytra of Agon- um chathami Van Dyke (Carabidae) col- lected by F S. Williams, 3 October 1905.

Type deposition.—Holotype, allotype, 9 adult 2 paratypes, 6 ¢d paratypes, 3 larval 2 paratypes and 2 egg paratypes of Eutar- sopolipus brettae deposited in the Depart- ment of Entomology, California Academy of Sciences, San Francisco, California, U.S.A. One 6, 2 adult 2, 1 larval 2 and 1 vial of paratypes in the collection of the au- thor.

Etymology.—The species is named for Roberta Brett, California Academy of Sci- ences, San Francisco, California, in recog- nition of her support of studies of parasitic mites of insects.

DISCUSSION

Before Regenfuss (1968) described 16 new species of Eutarsopolipus trom Ger- many, only E. lagenaeformis from Italy and E. desani Cooreman 1952 from Central Af- rica were known. Regenfuss (1968) defined 7 subgroups of Eutarsopolipus. As new species were described from Pacific Islands, Australia, India, S. Africa, and the Western Hemisphere, characters of existing sub- groups of Eutarsopolipus no longer worked in placing new species. Husband (1995) created the ochoai subgroup and Husband and Macfarlane (1999) created the catad- romi and secundus subgroups. There are now more than 40 species of Eutarsopoli- pus and many more expected from the more than 25,000 known species of Carabidae.

Major characters used to distinguish sub- groups of Eutarsopolipus include: presence

568

Table 1.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Comparisons of characters and maximum measurements of Eutarsopolipus brettae with Eutarso-

polipus species in the pterostichi and agonum groups, E. trichognathi from South America and E. ochoai from

Central America.

Character E. brettae pterostichi agonum E. trichognathi E. ochoai Adult females Stigmata 0) 0) “PF at + Idiosoma 450 510 498 550 580 Amb. II, III claws 10 7 0) O 3, thin Cheliceral stylets 85 48 40 54 67 Dors. gnath. setae 35 19 20 18 29 Vent. gnath. setae 25 6 5 5 25 Setae v, 15 6 9 5 23 Setae v, 8 9 20 5) 18 Femur I v” 19 m 0) O 20 Tibia I sol. 9 10 4 7) 9 Tibia II d 33 10 15 8 31 Genu I v’ 5 2 O O 4 Adult males Idiosoma 188 220) 196 173 222 Amb. HI, HI claws 5) minute 0) minute 3, thin Cheliceral stylets 31 28 22 D3} 31 Dors. gnath. setae 4, thick 13 i 8 13 Vent. gnath. setae 13 4 m m 9 Setae sc, 65 48 S)i/ 32 62 Femur I vy” 0-12 2 0) 0 0 Tibia I d 33 19 13 aT 31 Larval females

Idiosoma 192 270 230 332 AT) Amb. I, II claws 4 minute 0 minute 3, thin Cheliceral stylets 64 49 29 32 49 Dors. gnath. setae 36 30 16 16 26 Vent. gnath. setae 14 7 5 m 15 Femur I v” 14 m 0) 0) 10 Tibia I d 40 8 18 10 26 Tarsus IT sol. w 5S) 2 O 5 5 h,-h, distance 2 d 12 4 O, adj. h, 7 22. 30 25 m Genu I v, 0) 0) 0) 0) 5

or absence of stigmata, shape of the male genital capsule, proportions of cheliceral stylets in relation to gnathosoma, presence or absence of idiosomal setae or setae on legs and presence or absence of idiosomal plates. Major characters used to distinguish species include: relative lengths of chelic- eral stylets, lengths and widths of other structures, idiosomal plates of adult females divided or not, plates of larval females fused or not, idiosomal shape, structure of setae (thick, thin, with or without micro-

spines), setal position, and relatives lengths and position of setae of the legs and idio- soma.

All Eutarsopolipus that are parasites of the carabid genus Agonum fit patterns de- scribed for the biunguis subgroup proposd by Regenfuss (1968). Comparisons of Eu- tarsopolipus brettae with species in the biunguis group yielded more than 40 dif- ferences. All biunguis species have distinct stigmata, no ambulacral I claw, no or mi- nute ambulacra II, HI claws, no femur I se-

VOLUME 104, NUMBER 3

Table 2.

569

Leg setation of femur, genu, tibia and tarsus for larval female E. brettae and related species.

Solenidia are included. Two numbers in a column indicate variation in setal numbers.

Leg I

Leg I

I G Ni la EF G ri Ta F G ri Ta E. brettae | a 7 O | 4 6 0 | 4 5 E. ochoai 3 2 if 9 0 | 4 7 0 | 4 5 E. stammeri O 2 6 9 O | 4 6 0) | 4 5 E. trichognathi 2 0 7 8/9 0 0) 4 6 0) 0) 4 5 E. biunguis subgroup from Agonum 1/2 0) 7 8 0) O 4 5 0 O 4 5 E. lukoschusi 3 0) 6 9 0) 0 4 6 0) 0 4 5) E. pterostichi subgroup 2/3 0/2 5/7 8/9 0) 0) 4 6/7 0) 0) 4 5/6

tae v" and no genu I v” setae. Eutarsopolipis brettae has no stigmata and has the struc- tures which are missing in biunguis species. Patterns of characters do not fit patterns for the biunguis subgroup, any of the other 6 subgroups proposed by Regentuss or any of the 3 subgroups added since 1968.

An attempt was made to place E. brettae with species which have no stigmata: E. stammeri Regenfuss of the stammeri sub- group, E. lukoschusi Husband 1986 and species in the prerostichi subgroup. In con- trast to large claws of female EF. brettae, no claws are present in E. stammeri. In E. lu- koschusi ambulacral claws II, I are minute and ambulacrum I claw is small. Regenfuss (1974) placed EF. inermis Regenfuss from Georgia, U.S.A., within the pterostichi sub- group because it lacked stigmata, the geni- tal capsule of the male was similar to other mites in this group, solenidia were short as was femur I seta d and larval females had slightly separated setae /,. In contrast to other mites in the pterostichi subgroup, adult females of E. inermis have no genu I setae, all ambulacral claws are reduced and coxal seta 3a setae are not present. The ho- lotype of E. pterostichi Regentuss has 3 fe- mur I setae (as does E. brettae), 2 genu I setae in contrast to | genu I setae in E. bret- tae, and short solenidia w in contrast to lon- ger solenida w on tarsi I, Hl in E. brettae. Setae /i, are nearly adjacent in larval female E. brettae and genital capsules of males have concave rather than straight lateral

margins. Although adult female E. brettae do not appear to have setae 3a, ambulacral claws are not reduced as in E. inermis. With the exception of EF. inermis in the pterosti- chi subgroup, cheliceral stylets of all instars of E. brettae are distinctly longer (Table 1).

Geographically, the nearest Eutarsopoli- pus species to E. brettae are E. ochoai Hus- band 1995 parasite of Pasimachus spp. bee- tles from Costa Rica and E. trichognathi Husband and Eidelberg 1996 from Trichog- nathus marginipennis (Latreille) from Bra- zil. The species FE. trichognathi occurs in Ecuador, Columbia, Brazil, Paraguay and Bolivia (Husband 1999). In contrast to no stigmata in E. brettae, both E. ochoai and E. trichognathi have stigmata. Eutarsopo- lipus trichognathi shares characters with the biunguis subgroup of Eutarsopolipus. All instars of E. ochoai have small ambulacra II, II] claws in contrast to larger claws in E. brettae. Genu I of E. ochoai has 2 setae in contrast to | seta of E. brettae. Males of E. brettae have short, thick dorsal gnatho- somal setae (4) in contrast to longer gna- thosomal setae (13) in males of FE. ochoai. I consider, on the basis of these characters, that E. brettae is not closely related to the 2 species of Eutarsopolipus from Central and South America.

Until a reassessment of subgroups of Eu- tarsopolipus is completed, E. brettae cannot be placed in a subgroup. It has combina- tions of unique characters of no other Eu- tarsopolipus.

570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ACKNOWLEDGMENTS

The assistance of Roberta Brett and Da- vid Kavanaugh, Department of Entomolo- gy, California Academy of Sciences, Gold- en Gate Park, San Francisco, California, U.S.A. in acquiring specimens and advice on host beetles is appreciated.

LITERATURE CITED

Berlese, A. 1913. Acari Nuovi. Redia 9: 27-87.

Cooreman, J. 1952. Acariens Podapolipodidae du Con- go Belge. Bulletin Institut de Institute Royal des Sciences Naturelles de Belgique 28(35): 1—10.

Husband, R. W. 1986. New taxa of Podapolipidae (Ac- arina) from S. African Coleoptera: Result of the Namaqualand-Namibia expedition of the King Le- opold Foundation for exploration and protection of nature. Bulletin de I’ Institut Royal des Sciences Naturelles de Belgique: Entomologie 56: 5—14.

. 1995. A new species of Eutarsopolipus (Ac- ari: Podapolipidae) from Costa Rican Pasimachus spp. (Coleoptera: Carabidae). Entomologischen Mitteilungen aus dem Zoologische Museum Ham- burg 11(151): 157-165.

. 1999. American Eutarsopolipus of the biun-

guis group and descriptions of previously un- known male and larval female of Eutarsopolipus trichognathi (Acari: Podapolipidae) from South

America. International Journal of Acarology 25(1): 13-17.

Husband, R. W. and H. Dastych. 1998. A new species of Eutarsopolipus (Acari: Podapolipidae) from Chlaenius sericeus Frost (Coleoptera: Carabidae) from Athens, Georgia, U.S.A. Entomologischen Mitteilungen aus dem Zoologische Museum Ham- burg 12(158): 317-326.

Husband, R. W. and M. Eidelberg. 1996. A new spe- cies of Eutarsopolipus (Acari: Podapolipidae) from Trichognathus marginipennis (Coleoptera: Carabidae) from Brazil. International Journal of Acarology 22(3): 193-197.

Husband, R. W. and D. Macfarlane. 1999. Two new species of Eutarsopolipus (Acari: Podapolipidae) from Catadromus lacordairei (Coleoptera: Cara- bidae) from Australia. International Journal of Ac- arology 25(4): 297-308.

Lindquist, E. E. 1986. The world genera Tarsonemidae (Acari: Heterostigmata): A morphological, phy- logenetic, and systematic revision with reclassifi- cation of family group taxa in Heterostigmata. Memoirs of the Entomological Society of Canada 136: 1-517.

Regenfuss, H. 1968. Untersuchungen zur Morpholo- gie, Systematik und Okolgie der Podapolipidae (Acarina, Tarsonemini). Zeitschrift fiir Wissen- schaftliche Zoologie, Leipzig 177(3/4): 183-282.

. 1974. Neue ektoparasitische Arten der familie

Podapolipidae (Acari: Tarsonemini) von Carabi-

den. Mitteilungen aus dem Hamburg Zoolische

Museum und Institut 71: 147-163.

PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 571-575

NEW HOST RECORD, NEW RANGE INFORMATION, AND A NEW PATTERN OF VOLTINISM: POSSIBLE HOST RACES WITHIN THE HOLLY LEAFMINER PHYTOMYZA GLABRICOLA KULP (DIPTERA: AGROMYZIDAE)

SONJA J. SCHEFFER

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Rm. 137, BARC-West, 10300 Baltimore Avenue, Belts- ville, MD 20705, U.S.A. (e-mail: sscheffe @sel.barc.usda.gov)

Abstract.—The agromyzid leafminer Phytomyza glabricola is reported from the holly Ilex coriacea, a plant not previously reported to host leafminers. The known geographic range of P. glabricola is extended to include Alabama, Florida, Mississippi, North Car- olina, and South Carolina. In these areas, it can be found feeding on sympatric populations of its hosts /. coriacea and I. glabra. Phytomyza glabricola reared from I. coriacea are univoltine, having only a single generation a year, while on /. glabra this species is multivoltine with at least two generations a year. This suggests that either this leafminer exhibits a high degree of host-associated phenotypic plasticity in life history or that host

races are present.

Key Words:

The agromyzid fly Phytomyza glabricola Kulp belongs to a complex of closely re- lated leafminers, all of which feed on var- ious species of holly in the genus //ex (Aquifoliaceae) (Kulp 1968, Scheffer and Wiegmann 2000). Phytomyza_ glabricola was first described from the plant species Ilex glabra (Linnaeus) Gray by Kulp (1968) in his revision of holly-feeding Phytomyza species. On this host, P. glabricola makes a linear-blotch mine and is multivoltine (Kulp 1968, Al-Siyabi and Shetlar 1998). In 1996, I reared P. glabricola from 26 leafmines found on two bushes of /. cori- acea (Pursh) Chapman growing in North Carolina. This plant species has not been previously reported to host agromyzid leaf- miners (Spencer and Steyskal 1986, Spen- cer 1990).

Ilex glabra and I. coriacea are shrubby evergreen hollies that grow sympatrically in

host race formation, sympatry, Aquifoliaceae, gallberry, inkberry

the coastal plain of the southeastern U.S. The range of /. glabra extends from coastal Nova Scotia to mid Florida to eastern Texas and Missouri and is slightly broader than that of /. coriacea which extends only from coastal Virginia to northern Florida to Tex- as [Galle (1997) includes Mexico in the range, but most other sources do not (Stan- dley 1923, Lundell 1961, Gleason and Cronquist 1963, Radford et al. 1968, Cor- rell and Johnston 1970)]. These two plant species are believed to be very closely re- lated and go by the similar common names gallberry (. glabra) and large gallberry (J. coriacea) (Galle 1997).

Phytomyza glabricola has been reported from Connecticut, Massachusetts, New Jer- sey, New York, Ohio, and Washington, D.C. (Kulp 1968, Spencer and Steyskal 1986), but not from the southeastern U.S. where both J. glabra and I. coriacea are

S72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Collection locations, hosts, and dates for P. glabricola. en eee ee ee eee eee Location Host Date Alabama Covington Co. I. glabra Jan. 1998 Covington Co. I. coriacea Jan. 1998 Florida Calhoun Co. I. glabra Oct. 1999 Highlands Co. I. glabra Jan. 1998, Oct. 1999 Marion Co. I. glabra Jan. 1999 Wakulla Co. I. glabra Jan. 1998, Oct. 1999 Wakulla Co. I. coriacea Jan. 1998 Mississippi Forrest Co. I. glabra Jan. 1998 Forrest Co. I. coriacea Jan. 1998 North Carolina Brunswick Co. I. glabra Feb. 1997 Brunswick Co. I. coriacea Feb. 1997 Craven Co. I. glabra Feb. 2000 Craven Co. I. coriacea Feb. 2000 Moore Co. I. glabra Feb. 1996 Moore Co. I. coriacea Feb. 1996 New Hanover Co. I. glabra Feb. 1997, Jan. 1998 New Hanover Co. I. coriacea Feb. 1996, Feb. 1997, Jan. 1998 New York Suffolk Co. I. glabra Aug. 1996

South Carolina Berkeley Co. I. glabra Berkeley Co.

I. coriacea

Feb. 1997, Jan. 1998 Feb. 1997, Jan. 1998

common in the pine woods and bays of the coastal plain. The purpose of this study was to investigate the geographic range of P. glabricola and to determine whether this species commonly feeds on /. coriacea.

MATERIALS AND METHODS

From 1996 to 2000, I periodically trav- eled within the southeastern U.S. looking for populations of /. glabra and I. coriacea. Leaves containing well-developed mines were removed from plants, and adult flies were reared from the pupae found within the mines. Species identity of the leafmi- ners reared from /. glabra and I. coriacea was determined using Kulp (1968) and Spencer and Steyskal (1986).

RESULTS

Adult flies reared from /. glabra and I. coriacea were readily identified as P. gla- bricola. Flies reared from the two hosts did not noticeably differ from each other in morphological characters. Locations where P. glabricola was reared from leafmines on I. glabra and/or I. coriacea are listed in Ta- ble 1. Collection dates for samples are listed in Table 1; eclosion of adults occurred with- in approximately one month of this date.

Leafmines formed by P. glabricola on both I. glabra and I. coriacea are linear- blotch shaped, although in most cases the extensive feeding during the later instars re- moves tissue surrounding the initial linear mine so that only a blotch is apparent. Gen- erally, a leaf will contain only one mine

VOLUME 104, NUMBER 3

with a single larva or pupa present. Occa- sionally, a leaf will have more than one mine, indicating the presence of several lar- vae or pupae. This appeared to be more common on the larger-leafed /. coriacea than on /. glabra. Mines on /. glabra have a tendency to be brown with a blistered ap- pearance, while those on /. more often dark green or reddish-green and do not appear blistered. On both hosts, fly pupation occurs in the mine with the ante- rior pupal spiracles emerging through an “epidermal window,” where just prior to pupation, the larva removes mesophyll ad- jacent to the lower epidermis of the leaf.

During the course of this study, it became apparent that populations of P. glabricola on /. coriacea exhibit a univoltine life cy- cle. Well-developed mines are only found in January and February, and the adults eclose from late January to early March. Observations made at other times indicate that first-instar leafmines are found in the summer, second-instar mines in late fall, and third-instar mines only in the winter prior to adult eclosion. This univoltine life cycle is very similar to that of the native holly leafminer P. ilicicola on its host /. opaca (Kulp 1968, Potter and Kimmerer 1986), but it is in sharp contrast to the mul- tivoltine life cycle of P. glabricola on I. glabra (Kulp 1968, Al-Siyabi and Shetlar 1998), where the generation time can be as short as one month (Scheffer, personal ob- servation).

corlacea are

DISCUSSION

These findings confirm that /. coriacea is a widely used host for P. glabricola in nat- ural settings and extends the known geo- graphic distribution of P. glabricola to Al- abama, Florida, Mississippi, North Caroli- na, and South Carolina.

It is unlikely that further study will un- cover any additional hosts for P. glabricola. Most holly leafminers are strictly monoph- agous (Kulp 1968, Scheffer and Wiegmann 2000), although in artificial situations (e.g., botanical gardens), additional feeding on

573

non-native //ex species or hybrids may be observed (Scheffer, personal observation; host records listed in Spencer and Steyskal 1986, Griffiths and Piercey-Normore 1995). During the course of this study, P. glabri- cola was reared from I. glabra and I. cor- iacea in both natural and ornamental set- tings, but not from other hollies growing in the same locations, including the evergreen species /. cassine Linnaeus, [. myrtifolia Walter, /. opaca Aiton, and I. vomitoria Ai- ton (Scheffer, unpublished data).

Although holly leafmining species are known to exhibit different patterns of vol- tinism, no species previously has been re- ported to exhibit more than one pattern. Many holly leafminers have a multivoltine life cycle with two or more generations a year. In most of these species, the larval feeding period may be as short as a week or two, with a total generation time that of slightly over a month (Scheffer, personal observation; the life cycle of P. opacae Kulp is reported to be even shorter (Kulp 1968)). Multivoltine holly leafminers can be found on either evergreen or deciduous hollies and include the species P. ditmani Kulp, P. opacae Kulp, P. verticillatae Kulp, P. vomitoriae Kulp, and several undescri- bed species (Kulp 1968, Spencer and Steys- kal 1986, Scheffer, unpublished data). In contrast, univoltine holly leafminers have only one generation a year and a highly ex- tended larval developmental period lasting up to ten months that appears to be unique among the leafmining Agromyzidae (Cam- eron 1939, Hering 1951, Potter and Kim- merer 1986). Univoltine holly leafminers have been found only on evergreen hollies and include the species P. ilicicola Loew, P. ilicis Curtis, and an undescribed species on /. myrtifolia (Kulp 1968, Spencer and Steyskal 1986, Scheffer, unpublished data).

There are several possible explanations for the observation that P. glabricola has a different life history pattern depending on host plant identity. At the two extremes, these hypotheses can be classified as those involving phenotypic plasticity and those

574

involving genetic differences between flies on the two hosts. First, the rate of larval feeding could be a direct physiological re- sult of some unknown quality of host leaf tissue, such that P. glabricola larvae in I. glabra always develop quickly, while lar- vae in J. coriacea develop very slowly. This situation would be a form of host-mediated phenotypic plasticity and would not require genetic differences in the flies. Consider- ation of the life history patterns of related holly leafminers suggests that an hypothesis of host-mediated phenotypic plasticity, even if found to be valid in the case of P. glabricola, cannot generally explain voltin- ism differences in the holly leafmining group; P. ilicicola and P. opacae both ovi- posit into the soft new leaves of /. opaca, but P. ilicicola exhibits a univoltine life cy- cle on this host while P. opacae is multi- voltine (Kulp 1968, Scheffer and Wieg- mann 2000).

An alternative hypothesis is that the flies exhibit different patterns of voltinism be- cause they are genetically different and rep- resent host races or incipient or cryptic spe- cies. Under this scenario, the difference in life history on the two hosts could be either a cause of genetic divergence (i.e., the life history differences may be incompatible ad- aptations to each of the two host plants thereby favoring host race formation) or a result of genetic divergence (i.e., host races diverged for reasons unrelated to voltinism patterns and voltinism patterns have sub- sequently diverged). Note that even if ge- netic host races have formed, the difference in voltinism could still be due to phenotypic plasticity, if it exists, such that voltinism would depend on the host, but each host race would exhibit only one pattern because it uses only one host.

Whether P. glabricola on I. glabra and I. coriacea represents a single oligophagous species that feeds on two hosts or a com- plex of closely related but genetically di- vergent host races currently is not known. Although the difference in voltinism on the two hosts is suggestive of host races, the

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

voltinism difference by itself does not pre- clude flies from the two hosts from inter- breeding. Adult flies from both hosts emerge in late winter (see dates in Table 1) and can be expected to be present in the same general areas together for several weeks. During this one time each year, in- terbreeding between flies from the two hosts is at least possible.

Anecdotal evidence from the field indi- cates that host plant preferences suggestive of host races may be present within P. gla- bricola. When the univoltine leafminers on I. coriacea, which are often extremely abundant, emerge en mass in February, adults can be readily observed on /. cori- acea foliage but not on adjacent /. glabra foliage (Scheffer, personal observation), suggesting a possible preference for /. cor- iacea. This is despite the fact that P. gla- bricola trom I. glabra can be taken into the lab and easily reared to the next generation on /. glabra with no apparent hesitancy to oviposit and little or no larval mortality (Scheffer, personal observation).

Results to date of phylogenetic analysis of molecular data are inconclusive with re- gard to the question of host races wihtin P. glabricola. Mitochondrial cytochrome oxi- dase sequences of flies from /. glabra and I. coriacea are nearly identical (< 1% pair- wise divergence across more than 2000 bp; Scheffer and Wiegmann 2000), which is typical of sequence variation found within agromyzid species (Scheffer, unpublished data; see also Scheffer 2000). Using this gene region, there is no evidence of host- associated divergence, but given the esti- mated rate of evolution of insect mitochon- drial genes (approx. 2.3% per million years; Brower 1994), mitochondrial sequences would be unlikely to have sufficient varia- tion to track very recent divergence or spe- ciation events.

In conclusion, P. glabricola is widely distributed throughout the coastal plain of the eastern United States. It feeds on two holly species, /. glabra and I. coriacea, which are commonly sympatric throughout

VOLUME 104, NUMBER 3

much of their ranges. Differences exhibited by P. glabricola in patterns of voltinism on these two hosts suggest either host-associ- ated phenotypic plasticity in life history or the presence of genetically differentiated host races. Additional studies of morpho- logical variation, mating and oviposition

behaviors, and/or highly variable molecular

markers are needed to determine the status of P. glabricola \eafminers feeding on /. glabra and I. coriacea.

ACKNOWLEDGMENTS

I thank Leslie Iskenderian and Kevin Omland for excellent assistance in the field. Douglas Miller, Dave Hawthorne, and two anonymous reviewers provided useful com- ments on the manuscript. For permission to collect leafminers, I gratefully acknowledge authorities at Carolina Beach State Park (North Carolina), Francis Marion National Forest (South Carolina), Conecuh National Forest (Alabama), DeSoto National Forest (Mississippi), Croatan National Forest (North Carolina), Apalachicola National Forest (Florida), Ocala National Forest (Florida), and Archbold Biological Station (Lake Placid, Florida). During part of this work, I was supported by a National Sci- ence Foundation/Sloan Foundation Post- doctoral Research Fellowship in Molecular Evolution (Grant no. BIR-9510795). The Holly Society of America provided funds for an extensive holly leafminer collecting trip in 1998.

LITERATURE CITED

Al-Siyabi, A. A. K. and D. J. Shetlar. 1998. Inkberry leaf miner, Phytomyza glabricola Kulp (Diptera: Agromyzidae): Life cycle in Ohio. Ohio State Ex- tension Research Special Circular 165-99.

Brower, A. V. Z. 1994. Rapid morphological radiation and convergence among races of the butterfly Hel- iconius erato inferred from patterns of mitochon- drial DNA evolution. Proceedings of the National Academy of Sciences USA 91: 6491—6495.

Sys)

Cameron, E. 1939. The holly leafminer (Phytomyza ilicis, Curt.) and its parasites. Bulletin of Ento- mological Research 30: 173-208.

Correll, D. S. and M. C. Johnston. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, TX, 1881 pp.

Galle, E C. 1997. Hollies: The Genus //lex. Timber Press, Portland, Oregon, 573 pp.

Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Willard Grant Press, Boston, MA, 810 pp.

Griffiths, G. C. D. and M. D. Piercey-Normore. 1995. A new agromyzid (Diptera) leaf-miner of moun- tain holly (Nemopanthus, Aquifoliaceae) from the Avalon Peninsula, Newfoundland. Field-Naturalist 109: 23-26.

Hering, E. M. 1951. Biology of the Leaf Miners. Junk s’Gravenhage, The Hague, The Netherlands, 420 pp.

Kulp, L. A. holly leaf miners (Diptera: Agromyzidae). Uni- versity of Maryland Agriculture Experiment Sta- tion Bulletin A-155: 1—42.

Lundell, C. L. 1961. Flora of Texas, Vol. 3. Texas Re- search Foundation, Renner, TX, 433 pp.

Potter, D. A. and T. W. Kimmerer. 1986. Seasonal al- location of defense investment in //ex opaca Aiton

Canadian

1968. The taxonomic status of dipterous

and constraints on a specialist leafminer. Oecolo- gia 69: 217-224.

Radford, A. E.; H: Ey Ables;and CR. Bells 1968: Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill, NC, 1183 pp.

Scheffer, S. J. 2000. Molecular evidence of cryptic species within the Liriomyza huidobrensis (Dip- tera: Agromyzidae). Journal of Economic Ento- mology 93: 1146-1151.

Scheffer, S. J. and B. M. Wiegmann. 2000. Molecular phylogenetics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): Species limits, speci- ation, and dietary specialization. Molecular Phy- logenetics and Evolution 17: 244—255.

Spencer, K. A. 1990. Host specialization in the world Agromyzidae (Diptera). Kluwer Academic Pub- lishers, Dordrecht, The Netherlands, 444 pp.

Spencer, K. A. and G. C. Steyskal. 1986. Manual of the Agromyzidae (Diptera) of the United States. United States Department of Agriculture Hand- book No. 638, Washington, DC, 478 pp.

Standley, P. C. 1923. Trees and Shrubs of Mexico. Contributions from the United States National Herbarium, Vol. 23. Washington, 1721 pp.

PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 576-588

LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF GOEDENIA RUFIPES (CURRAN) (DIPTERA: TEPHRITIDAE) ON ISOCOMA ACRADENIA (E. GREENE) E. GREENE IN SOUTHERN CALIFORNIA

RICHARD D. GOEDEN

Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu)

Abstract.—Goedenia rufipes (Curran) is an oligophagous, nonfrugivorous, fruit fly (Dip- tera: Tephritidae) producing at least two, probably three, annual generations altogether in the flower heads of Chrysothamnus teretifolius (Durand and Hilgard) H. M. Hall and Isocoma acradenia (E. Greene) E. Greene in southern California. Both of these confirmed hosts are Asteraceae belonging to the subtribe Solidagininae of the tribe Astereae. The egg, second- and third-instar larvae, and puparia are described and figured, and selected characteristics of these stages are compared with those of G. timberlakei (Blanc and Foote), the only other well-known species of Goedenia. The egg of G. rufipes is the first pictured for this genus; it bears a prominent pedicel with semicircular to fusiform micro- pyles. The second instar is white, but the third instar has a dark brown to black venter on the meso- and metathorax and a similarly darkened caudal segment. The prothorax and gnathocephalon of the second and third instars are smooth, mostly free of the minute acanthae that circumscribe most other body segments. Minute acanthae cover the posterior end of the truncated caudal segment, which also is perforated by scattered, open pores. The third instar lacks oral ridges. The anterior thoracic spiracle of the second instar bears three papillae, which are reduced to two papillae in the third instar. The life cycle is of the aggregative type and overwintering occurs in dead flower heads as prepuparial third instars, aS puparia in an open, central cells loosely surrounded by floret fragments and intact undamaged achenes, and as unmated, sexually immature adults. Eurytoma sp. (Hy- menoptera: Eurytomidae) and Pteromalus sp. (Hymenoptera: Pteromalidae) are reported as solitary, larval-pupal endoparasitoids, and Eupelmus sp. (Hymenoptera: Eupelmidae) as possible solitary endoparasitoids.

Key Words: Insecta, Goedenia, Chrysothamnus, Isocoma, Asteraceae, nonfrugivorous Tephritidae, biology, taxonomy of immature stages, flower-head feeding, aggregative life cycle, seed predation, parasitoids

Most indigenous, western North Amer- _ stages of only one of the eight known spe- ican Myopitini (Diptera: Tephritidae: Te- cies of Goedenia have been described in phritinae) formerly assigned to the Pale- detail, i.e., G. timberlakei (Blanc and arctic genus Urophora Robineau-Desvo- Foote), by Goeden et al. (1995). This pa- idy were reclassified in the genus Goe- per describes the life history and selected denia by Freidberg and Norrbom (1999). immature stages of a second species, G. To date, the life history and immature rufipes (Curran).

VOLUME 104, NUMBER 3

MATERIALS AND METHODS

The present study utilized specimens of adults reared from 1|-liter samples of mature flower heads of Chrysothamnus teretifolius (Durand and Hilgard) H. M. Hall and /so- coma acradenia (E. Greene) E. Greene (As- teraceae) collected throughout southern California since 1980 (Goeden 1987). The life history study and description of the im- mature stages of G. rufipes were based in large part on dissections of samples of ma- ture and immature flower heads of /. acra- denia collected east of Ocotillo at Coyote Wells and at 42-m elevation, southwestern Imperial Co., during 1990-1999, One-liter samples of excised, immature and mature flower heads containing the scarce larvae and purparia were transported in cold- chests in an air-conditioned vehicle to the laboratory and stored under refrigeration for subsequent dissection, photography, de- scription, and measurement. Ten ova dis- sected from a gravid female as well as two second- and 22 third-instar larvae, and nine puparia dissected from flower heads were preserved in 70% EtOH for scanning elec- tron microscopy (SEM). Prepuparia and pu- paria were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temper- ature for adult and parasitoid emergence. Specimens for SEM were hydrated to dis- tilled water in a decreasing series of acid- ulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, I-h immer- sions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, studied and digitally photographed with a Philips XL-30 scan- ning electron microscope in the Institute of Geophysics and Planetary Physics, Univer- sity of California, Riverside.

Five arenas each consisting of a clear- plastic, petri dish were provisioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994). Each arena contained a vir-

S77.

gin male and female obtained from emer- gence cages that were used for observations of courtship and copulation behavior.

Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden (2001la, b, c), Goeden et al. (1993), Goeden and Headrick (1992), Goeden and Norrbom (2001), Goe- den and Teerink (1997), Teerink and Goe- den (1999), and our earlier works cited therein. Means + SE are used throughout this paper. Digitized photographs used to construct text figures were processed with Adobe Photoshop® Version 6.

RESULTS AND DISCUSSION TAXONOMY

ADULT.—Goedenia rufipes was described from Arizona as Aleomyia rufipes by Cur- ran (1932), who shortly thereafter reclassi- fied it as Euribia rufipes (Curran 1934, as cited by Foote et al. 1993). Foote (1965) assigned it to Urophora, which Steyskal (1979) adopted in his key to Myopitinae genera and species of Urophora. Freidberg and Norrbom (1999) redesignated most of the indigenous, western North American species as Goedenia, with A. rufipes Curran as the type species.

The wing was figured by Curran (1934, as. cited! by Foote etealss 1993) sSteyskal (1979), and Foote et al. (1993). Freidberg and Norrbom (1999) provided line draw- ings of the head in lateral and anterolateral view and the hypandrium and phallopode- me in dorsal view.

Immature stages.—The egg, second- and third-instar larvae, and puparium of G. ru-

fipes are described below.

Egg: Sixteen ova dissected from a 17- day-old female were white, opaque, smooth, elongate-ellipsoidal, 0.55 + 0.008 (range, 0.48—0.60) mm long, 0.18 + 0.005 (range, 0.14—0.20) mm wide, smoothly rounded at tapered basal end (Fig. 1A);

578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. terior to left; (B) pedicel showing pattern and shapes

Egg of Goedenia rufipes: (A) habitus, an-

of aeropyles (probably distorted).

pedicel prominent, 0.03 mm long, circum- scribed apically by different-sized, semicir- cular to fusiform aeropyles arranged singly or in two rows with their long axes parallel to the long axis of the egg (Fig. 1B).

The ova of G. rufipes on average equalled in length the ova of G. timberlakei and were slightly wider, but otherwise agreed with the description provided by Goeden et al. (1995). Unfortunately, the scanning electron micrograph of a possibly deformed pedicel of G. rufipes (Fig. 1B) is the only view of this structure obtained for this genus to date. The eggs and first instars of Goedenia spp. are very small, extremely difficult to find and probably are available for only a short period in nature; moreover, their study is further complicated by the tendency of these tephritids to co-occur

with other tephritid genera, i.e., Neaspilota, Procecidochares, Trupanea, in flower heads of the same host (“‘symphagy’’, Goe- den 1997), usually in subordinate numbers, as reported by Goeden (1987).

Second instar larva: White, cylindrical, tapered anteriorly, bluntly rounded posteri- orly (Fig. 2A); gnathocephalon conical (Fig. 2B), smooth, with few minute acan- thae ventrally (Fig. 2B-1), lacking oral ridg- es (rugose pads); dorsal sensory organ well- defined, round, flattened (Figs. 2B-2, C-1); anterior sensory lobes (Figs. 2B-3, C-2), separated by vertical medial cleft, with ter- minal sensory organ (Figs. 2B-4, C-3), lat- eral sensory organ (Fig. 2C-4), supralateral sensory organ (Fig. 2C-5), and pit sensory organ (Fig. 2C-6); stomal sense organ (Figs. 2B-5, C-7) ventrolaterad of anterior sensory lobe and fused (Fig. D-1) with la- teralmost of five, foliose, protrudent, lateral integumental petals (Figs. 2C-8, D-2) dor- sad of each mouthhook, two vertical pairs of medial integumental petals between an- terior sensory lobes (Fig. 2C-9); mouth- hook (Figs. 2B-6, D-3) bidentate (Fig. 2D- 3); median oral lobe laterally compressed, apically tapered (Fig. 2D-4), separated from labial lobe (Fig. 2D-5); verruciform sensilla circumscribe posterior third of gnathoce- phalon dorsomedially, dorsolaterally, and laterally (Fig. 2B-7); anterior spiracle with three, subglobose papillae (Fig. 2E); minute acanthae (Fig. 2F) posteriorly directed, spatulate, apically rounded on anterior fourth of meso- and metathorax venters and circumscribing all but posterior three- fourths of first abdominal segment (A1), most of A2 to A6, all but posterior three- fourths of A7, and anterior half of A8. Pos- terior surface of caudal segment not viewed.

The habitus of the second instar of G. rufipes (Fig. 2A) approximates that of G. timberlakei (Goeden et al. 1995). Differ- ences noted include five lateral integumen- tal petals in G. rufipes (Figs. 2C-8, D-2), not four, as pictured for G. timberlakei (Goeden et al. 1995). Moreover, the latter

VOLUME 104, NUMBER 3 579

Fig. 2. Second instar of Goedenia rufipes: (A) habitus, anterior to right; (B) gnathocephalon, frontolateral

view, |—minute acanthae, 2—dorsal sensory organ, 3—anterior sensory lobe, 4—terminal sensory organ, 5— stomal sense organ; 6—mouthhook, 7—verruciform sensilla; (C) gnathocephalon, close-up, 1—dorsal sensory

organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supralateral sensory

lateral integumental petals, 9—medial integumental

organ, 6—pit sensory organ, 7—stomal sense organ, 8 petals, mouthhook; (D) oral cavity of gnathocephalon, ventrolateral view, 1—stomal sense organ, 2—lateralmost integumental petal, 3—mouthhook, 4—median oral lobe, 5—labial lobe; (E) anterior spiracle; (F) minute acan-

thae on dorsolateral aspect of abdominal segments | and 2, anterior to left.

580

species appears to have only a single pair of medial integumental petals (Goeden et al. 1995), not two pairs, like G. rufipes (Fig. 2C-9). The numbers of these integumental petals were not quantified by Goeden et al. (1995), but their presence and general po- sitions were noted and pictured. Another difference is that the anterior spiracle of the second instar of G. rufipes bears three pa- pillae (Fig. 2E), not two papillae, like G. timberlakei (Goeden et al. 1995).

Third instar larva: Oblong-ellipsoidal, roundly tapered anteriorly, bluntly truncat- ed posteriorly (Fig. 3A), integument white, but venter of meso- and metathorax with dark brown to black infuscation (Fig. 6B); caudal segment dark brown or black (Figs. 6B, C); outwardly or posteriorly directed, conical, bluntly or sharply pointed (Figs. 3B-1, F-1) or hemispheroidal (Figs. 4C-1, E-2, F-2) minute acanthae circumscribe an- terior fourth of meso- and metathorax and first abdominal segment (A1) and cover all venters thereof, and circumscribe all but an- terior and posterior fifths of A2-A5, all but posterior quarter of A-7, and cover poste- rior plate of caudal segment (Figs. 4D-2, E- 2, F-2); prothorax smooth, lacking minute acanthae (Fig 3B), but circumscribed by verruciform sensilla dorsally, dorsolaterally, laterally, and ventrolaterally (Fig. 3B-2); gnathocephalon conical, anteriorly flat- tened, and medially divided by vertical cleft (Figs. 3B-3, C), pore dorsoposteriorad of each dorsal sensory organ (Fig. 3C-1); dor- sal sensory organ well-defined, hemispher- ical (Figs. 3C-2, D-1); anterior sensory lobe (Figs. 3C-3, D) bears terminal sensory or- gan (Figs. 3C-4, D-2), lateral sensory organ (Fig. 3D-3), supralateral sensory organ (Fig. 3D-4), and pit sensory organ (Fig. 3D- 5); two medial, papilliform integumental petals (Figs. 3C-5, D-6, E-1), four, lateral, spatulate or papilliform, integumental petals (Figs. 3C-6, D-7, E-2) above each mouth- hook (Figs. 3C-7, E-3), lower, lateral petal separate from stomal sense organ (Figs. 3C- 8, D-8, E-4) ventrolaterad of anterior sen- sory lobe; none (Figs. 3C, D), or sometimes

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

one, oral ridge (Fig. 3E-5) laterad of each anterior sensory lobe; mouthhook bidentate, anterior tooth, concave ventrally (Figs. 3E- 6); median oral lobe laterally compressed, apically pointed (Figs. 3C-9, E-7); anterior thoracic spiracle on posterior margin of prothorax bears two subglobose papillae (Figs. 3B-4, F-2); mesothoracic, lateral spi- racular complex with six verruciform sen- silla in vertical series (Fig. 4A-1), mesotho- racic spiracle not seen; metathoracic lateral spiracular complex with nearly closed, lat- eral spiracle (Fig. 4B-1) and four verruci- form sensilla (Figs. 4A-2, B-2) in vertical series posterior to spiracle; lateral spiracular complex of first abdominal segment con- sists of nearly closed spiracle (Figs. 4A-3, C-1) and three verruciform sensilla in ver- tical series posterior to spiracle (Figs. 4A- 4, C-2); caudal segment with pair of pos- terior spiracular plates (Figs. 4D-1, E-1, F) surrounded by hemispherical minute acan- thae (Figs. 4D-2, E-2, F-1) interspersed dor- somedially, medially, and less so, ventro- medially with open pores (Figs. 4D-3, E-3); each posterior spiracular plate bears three, smoothly flattened, ovoid rimae (Fig. 4F-2), ca. 0.01 mm in length, and four spinose in- terspiracular processes, each ca. 0.004 mm long (Fig. 4F-3).

The habitus of the third instar of G. ru-

fipes resembles that of G. timberlakei (Goe-

den et al. 1995). In both species, the venter of the thorax and the caudal segment are darkly pigmented (Figs. 6C, Goeden et al. 1995) and minute acanthae circumscribe the meso- and metathorax and abdomen, and especially noteworthy, the posterior surface of the caudal segment, also 1s dotted prom- inently with scattered pores (Figs. 4D-3, E- 3; Goeden et al. 1995). The prothorax and gnathocephalon of both species are smooth and free of minute acanthae (Fig. 3C, Goe- denver ale 1995):

Two medial and four lateral integumental petals are present in G. rufipes (Figs. 3C-5, =6, D=6., -7,, E-1.-2)2 whereas) G. timber- lakei has two medial and six lateral inte- gumental petals (Goeden et al. 1995, un-

VOLUME 104, NUMBER 3 58]

Fig. 3. Third instar of Goedenia rufipes: (A) habitus, anterior to left; (B) gnathocephalon and prothorax,

frontolateral view, |—minute acanthae, 2—verruciform sensilla, 3—gnathocephalon, 4—anterior spiracle; (C)

anterior sensory lobe, 4—terminal sensory

gnathocephalon, frontal view, 1—pores, 2—dorsal sensory organ, 3

organ, 5—medial integumental petals, 6—lateral integumental petals, 7—mouthhook, 8—stomal sense organ, 9—median oral lobe; (D) gnathocephalon, close-up, 1—dorsal sensory organ, 2—terminal sensory organ, 3— lateral sensory organ, 4—supralateral sensory organ, 5—pit sensory organ, 6—medial integumental petals, 7—

lateral integumental petals, 8—stomal sense organ; (E) oral cavity of gnathocephalon, ventral view 1—medial

integumental petals, 2—lateral integumental petals, 3—mouthhook, 4—stomal sense organ, 5—posterior con-

cavity on anterior tooth of mouthhook, 6—median oral lobe; (F) anterior spiracle, 1—minute acanthae, 2—

papillae.

582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

20 ime,

Fig. 4. Third instar of Goedenia rufipes, continued, (A) lateral spiracular complexes, 1—verruciform sensilla on mesothorax, 2—verruciform sensilla on metathorax, 3—spiracle on first abdominal segment, 4—verruciform sensilla on first abdominal segment, (B) 1—metathoracic lateral spiracle, 2—verruciform sensillum; (C) 1— lateral spiracle of first abdominal segment, 2—minute acanthae; (D) caudal segment, 1—posterior spiracular plates, 2—minute acanthae, 3—pores; (E) caudal segment, close-up, 1—posterior spiracular plates, 2—minute acanthae, 3—pores; (F) posterior spiracular plate, |—minute acanthae, 2—rimae, 3—interspiracular processes, 4—ecdysial scar.

VOLUME 104, NUMBER 3

published data). The lateralmost integumen- tal petals are separated from the stomal sense organs in both species (Figs. 3C-8, D- 8, E-4; Goeden et al. 1995).

Like G. timberlakei (Goeden et al. 1995, unpublished data), the third instar of G. ru- fipes lacks oral ridges on either side of the mouth opening, and ventral or ventrolateral to the stomal sense organ (Fig. 3E).

The mouthhooks of the third instars of G. rufipes (Fig. 3E-6), like those of G. tim- berlakei (Goeden 2001c), are bidentate; however, the teeth of the latter species were described as “‘conical’’, but in the former species the anterior tooth is concave ven- trally (Fig. 3E-6). Unfortunately, a vertical view of the oral cavity such as obtained for G. rufipes (Fig. 3E), was not obtained for G. timberlakei (Goeden et al. 1995, unpub- lished data).

The anterior spiracle of both species bears only two papillae (Figs. 3B-4, F; Goe- denvet aly995).

The lateral spiracular complex of the me- sothorax of G. rufipes includes six verru- ciform sensilla in a vertical series (Fig. 4A- 1); whereas, in G. timberlakei, this same complex includes only two verruciform sensilla (Goeden et al. 1995). Likewise, the metathoracic lateral spiracular complex of G. rufipes includes four verruciform sensil- la (Fig. 4A-2), but again, only two such sensilla in G. timberlakei (Goeden et al. 1995). Finally, three verruciform sensilla in a vertical series comprise the lateral spirac- ular complex of the first abdominal segment of G. rufipes (Figs. 4A-4, C-2), but only one verruciform sensillum is found on this segment in G. timberlakei (Goeden et al. 1995):

Differences noted between the second and third instars of G. rufipes include the reduction in the number of papillae on the anterior thoracic spiracle from three (Fig. 2E) to two (Figs. 3B-4, F-2). The number of lateral integumental petals also is re- duced from five in the second instar (Figs. 2C-8, D-2) to four in the third instar (Figs. 3C-6, D-7, E-2). The stability of these num-

583

bers is questionable, and their separation as medial versus lateral integumental petals is problematic because more precise counts and assessments would require consider- ably more replication of similar views of gnathocephala than were available to the author.

Puparia: Light (Fig. 6D) to dark (Fig. 6E), reddish brown with dark brown to black, anterior stripe on venter of meso- and metathorax and similarly dark caudal segment, elongate-ellipsoidal, with smooth- ly rounded anterior end and truncated pos- terior end (Fig. 5A). Anterior end bears in- vagination scar and raised, bilobed, anterior thoracic spiracles (not shown). Flattened posterior end of caudal segment studded with smoothly rounded, hemispherical, mi- nute acanthae (Fig. 5B-1) interspersed with open pores (Fig. 5B-2). A pair of raised, oval, posterior spiracular plates (Fig. 5B-3) each bear three oval rimae interspersed with four, peg-like interspiracular processes (not shown, see Fig. 4F and above description of third instar for details). Ten puparia dis- sected from flower heads of /socoma acra- denia averaged 2.73 + 0.07 (range, 2.50-— 3.17) mmin dength:. 1.157 0:04. @ange; 1.05—1.31) mm in width.

DISTRIBUTION AND HOostTs

To date, Goedenia rufipes only is known from southern California and southwestern Arizona north of Mexico (Foote et al. 1993); however, it probably ranges well into Mexico attacking flower heads at least of Chrysothamnus teretifolius and Isocoma acradenia. The former host is newly re- ported (Goeden 1987, Foote et al. 1993). Both of these confirmed hosts are Astera- ceae belonging to the subtribe Solidagini- nae of the tribe Astereae (Bremer 1994). Accordingly, G. rufipes probably is a nar- rowly oligophagous tephritid, that to date has not been reared by me from flower heads of several other species of these two, common and widespread, plant genera. Chrysothamnus teretifolius occurs on rocky slopes and flats from 600 to 4,000 m in Cal-

584

Figs 5: anterior to left, (B) caudal segment, 1—minute acan-

Puparium of Goedenia rufipes: (A) habitus,

thae, 2—pores, 3—posterior spiracular plates.

ifornia to southern Nevada and northwest- ern Arizona (Hickman 1993), including the higher parts of the Lower Sonoran Zone and arid, lower margins of the Upper Son- oran Zone as delimited by Shreve and Wig- gens (1964). Isocoma acradenia occurs on sandy or clay soils in alkaline or gypsum flats or slopes below 1,300 m in California, Arizona, Nevada, and Baja California, Mexico (Hickman 1993).

BIOLOGY

Egg.—Egegs of G. rufipes are inserted singly, pedicel-last, parallel to the long axes of and between the outer phyllaries of closed, preblossom flower heads of [. acra- denia.

Larva.—Upon eclosion, the first instar tunneled immediately through the inner

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bracts and into an ovule of a preblossom flower head. It fed with its body perpendic- ular to and its mouthparts toward the recep- tacle within an ovule, which it first exca- vated, then exited and entered an adjacent ovule. The receptacle was neither abraded or pitted by such feeding.

Second instars (Fig. 6A) mainly contin- ued feeding on ovules in closed, preblos- som flower heads, but a few were found feeding on soft achenes in open, blossom or postblossom flower heads (Fig. 6A). They usually fed within the ovules or achenes with their bodies perpendicular to the re- ceptacles, but always above the receptacles. Receptacles of 10 flower heads containing second instars averaged 1.45 + 0.12 (range, 0.85—1.99) mm in diameter. These 10 flow- er heads each contained a single larva that had damaged an average of 3.2 + 0.6 (range, 1—6) ovules/achenes, or about 28% of the average total of 11.6 = 0.8 (range, 7-14) ovules/achenes per flower head counted within the 10 flower heads. How- ever, more than 600 uninfested flower heads were individually dissected in order to lo- cate these 10 (1.7%) flower heads infested with second instars.

Third instars in flower heads fed with their long axes oriented perpendicular to the receptacles, and with their mouthparts di- rected towards the receptacles, that they usually scored or pitted deeply (Fig. 6B, C). Fifty flower heads (three, closed preblos- som; six, open blossom; 41 postblossom) were dissected that contained an average of 1.3 + 0.1 (range, 1—4) third instars. These 55 flower heads averaged 1.8 + 0.05 (range, |.1—2.9) mm in diameter and con- tained an average total of 13.9 + 0.4 (range, 8—20) ovules/achenes, of which on average 7.1 + 0.5 (range, 1-15) ovules/achenes were damaged or ~51%. However, well over 1,000 flower heads were individually dissected to locate these 55 infested flower heads. Receptacles that consistently were pitted suggested that sap constituted at least part of the diet of third instars of G. rufipes. Goeden (1988), Headrick and Goeden

VOLUME 104, NUMBER 3

(1990), Goeden and Headrick (1992), Goe- den’ et al. (1993, 1995), Headrick et. al. (1996), Goeden and Teerink (1997) first noted, described, and discussed sap feeding by florivorous species of Tephritidae in the genera Trupanea, Paracantha, Neaspilota, Tephritis, Goedenia (as Urophora), Dioxy-

na, and Xenochaeta, respectively. Upon completing feeding, the larvae oriented

with their anterior ends away from the re- ceptacles, retracted their mouthparts, and formed puparia (Figs. 6D, E).

Pupa.—The receptacles of nine flower heads that contained an average of 1.3 + 0.2 (range, 1—2) puparia (Figs. 6D, E) av- eraged 1.9 + 0.1 (range, 1.4—2.3) mm in diameter. The receptacles were deeply pit- ted in all nine flower heads, further con- firming that sap constituted part of the diet of third instars. The posterior end of the puparium rested in the smooth cup-like de- pression and the middle and anterior part of the puparium was surrounded by excavated floret fragments that formed a close, central cell not glued to the puparium.

Adult.—The premating and mating be- haviors of G. rufipes were not studied in the field, but were observed in petri dish arenas of the type found to be useful with many other nonfrugivorous, tephritid species (Headrick and Goeden 1994). The wings of both sexes were held away from the body at about 45° without supination when at rest (Fig. 6F). Both sexes exhibited wing ha- mation (Headrick and Goeden 1994) throughout the day concurrent with other behaviors, 1.e., grooming, resting, and feed- ing; this also was the most common wing movement reported for G. timberlakei (Goeden et al. 1995). Premating behaviors observed with G. rufipes included males and females tracking individuals of the op- posite sex, during which males sometimes swayed and usually exhibited abdominal pleural distension (Fig. 6F). The male ag- gressively mounted a female by jumping upon her, usually from the front, then turn- ing and forcefully grasping her wing bases, thoracic pleura, and aculeus for purchase,

585

while the female usually struggled, resisted his attempts to part her wings, and pushed against the male with her hind legs and tar- si. The males countered by grasping and raising the oviscape with the mid- and hind tarsi, while positioning the apex of the ex- serted aculeus of a receptive female against his epandrium. Nonreceptive females did not exsert their aculeus or pressed the ex- serted aculeus against the substrate so as to hinder or prevent proper positioning by males. But if receptive, the female allowed the male to raise her oviscape and extended her aculeus to its full length in reponse to his copulatory induction behavior (““CIB”’; Headrick and Goeden 1994, 1999). The CIB mainly consisted of the male rapidly rubbing his hind tarsi, back and forth, along her oviscape. The final mating position commonly had the wings of the male parted at about 20° (Fig. 6H), the wings of the fe- male parted at 60° (Figs. 6G, H), with both pairs of wings centered over the midlines of the flies (Figs. 6G, H). The foretarsi of the male grasped the dorsum of the abdo- men of the female laterally at the thoracic juncture, the midtarsi grasped the oviscape at its base, and the hindtarsi crossed under the oviscape or occasionally rested on the substrate (Fig. 61). The body of the female paralled the substrate with the oviscape raised about 30°, while the extended acu- leus pushed the male upward and backward (Fig. 61). Five pairs were observed to mate once or twice per day for a total of nine matings that lasted an average of 96 (range, 20—285) min. Females became restless be- fore termination of mating by pushing against the males with their hind tarsi, by lofting their wings so as to push them against the males, and by fully extending their aculeus. The male in turn countered this behavior with CIB, rocked from side to side to regain purchase or to avoid the fe- male’s pummeling, and sometimes rapidly vibrated his wings, all of which appeared to calm the female and allow coitus to contin- ue. Females sometimes walked about the arenas carrying the males while remaining

586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Life stages of Goedenia rufipes (A-E in flower heads of [socoma acradenia): (A) second instar (arrow) feeding on floret; (B) third instar (arrow) feeding deeply in receptacle of flower head; (C) third instar showing dark, ventral infuscation (arrow); (D) newly formed, light brown puparium (arrow); (E) dark overwin- tered puparium (arrow); (F) ventral view of adult male with wings in resting position with inflated abdominal pleura; (G) mating pair, dorsal view; (H) mating pair, ventral view; (1) mating pair, lateral view. Lines = | mm.

VOLUME 104, NUMBER 3

in copula. One male was observed to at- tempt mating with a newly dead female continuously for 5 h. Separations of three other pairs were observed, during which the male initially moved slightly forward while pulling the aculeus vertically upward, then rapidly turned and laterally walked off the female, and quickly posteriorly away from the female, while pulling free his genitalia. The separation of one pair took 15 s.

Seasonal history.—The life cycle of G. rufipes in southern California follows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the third instar, pu- parium, and some adults variously are the overwintering stages. Adults emerge from some of the puparia formed in late-fall, ear- ly winter (October-December) and these unmated, sexually immature adults over- winter. The remaining, nonfeeding third in- Stars, prepuparia, and puparia Overwinter in dead flower heads remaining on dormant Chrysothamnus teretifolius and Isocoma acradenia. These overwintered individuals emerge as adults in late winter (February— March) and either aggregate on one or more, as-yet-unknown, spring blooming, al- ternate hosts to mate and produce an as-yet- undetected spring generation. Or, the over- wintered and newly emerged adults contin- ue to pass the following spring and summer (April—August), possibly still as non-repro- ductive individuals feeding in mountain meadows and riparian habitats bordering the low-elevation, Sonoran Desert. They eventually aggregate on preblossom, fall- blooming, C. teretifolius and I. acradenia, mate, and subsequently oviposit in the small, newly-formed, closed, preblossom flower heads.

Natural enemies.—Fourteen Eurytoma sp. (Hymenoptera: Eurytomidae) and 12 Pteromalus sp. (Hymenoptera: Pteromali- dae) were reared from separate puparia of G. rufipes as solitary, larval-pupal endopar- asitoids. Two Eupelmus sp. (Hymenoptera: Eupelmidae) also were reared from insec- tary cagings of mature flower heads as pos- sible solitary endoparasitoids, as reported

587

from G. Goeden et al.

(1995).

timberlakei by

ACKNOWLEDGMENTS

I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. Krassimer Bozhilov in the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. I also am grateful to Jeff Teerink for his technical assistance and to David Headrick for his helpful comments on an earlier draft of this paper.

LITERATURE CITED

Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon.

Curran, C. H. 1932. New North American Diptera, with notes on others. American Museum Novita- tes 526: 1-13.

—.. 1934. The families and genera of North Amer- ican Diptera. C. H. Curran (privately printed). Foote, R. H. 1965. Family Tephritidae, pp. 658-678. In Stone et al., eds. A catalog of the Diptera of America North of Mexico. U. S. Department of

Agriculture Handbook 276. 1,696 pp.

Foote, R. H., KE L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York.

Freidberg, A. and A. L. Norrbom. 1999. A generic reclassification and phylogeny of the Tribe My- opitini (Tephritinae), pp. 581—627 (Chapter 23). Jn M. Aluja and A. L. Norrbom, eds. Fruit Flies (Te- phritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, Florida. 944 pp.

Goeden, R. D. 1987. Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in southern California. Proceedings of the Entomological So- ciety of Washington 89: 269-274.

1988. Life history of Trupanea imperfecta

(Coquillett) on Bebbia juncea (Bentham) Greene

in the Colorado Desert of southern California

(Diptera: Tephritidae). Pan-Pacific Entomologist

64: 345-351.

. 1997. Symphagy among florivorous fruit flies

(Diptera: Tephritidae) in southern California.

Chapter 3. /n K. Dettner, G. Bauer, and W. V6IkI,

eds. Vertical Food Web Interactions: Evolutionary

Patterns and Driving Forces. Ecological Studies

130: 39-52. Springer-Verlag, Heidelberg, Ger-

many.

. 2001la. Life history and description on im- mature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidental- is (Nuttall) Torrey and A. Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 103: 191—206.

. 2001b. Life history and description on im- mature stages of Tephritis joanae Goeden (Dip- tera: Tephritidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 586—600.

. 2001c. Life history and description on im- mature stages of Tephritis teerinki Goeden (Dip- tera: Tephritidae) on Hulsea vestita A. Gray (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 807-825.

Goeden, R. D. and D. H. Headrick. 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 94: 59-77.

Goeden R= Ds) DH? Headricks and Wh vA] Meernmnk. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210— 222

. 1995. Life history and description of imma- ture stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 97: 779-790.

Goeden, R. D. and A. L. Norrbom. 2001. Life history and description of adults and immature stages of Procecidochares blanci n. sp. (Diptera: Tephriti- dae) on /socoma acradenia (E. Greene) E. Greene (Asteraceae) in southern Calfornia. Proceedings of the Entomological Society of Washington 103: 517-540.

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Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Xenochaeta albiflorum Hooker in central and southern Cali- fornia. Proceedings of the Entomological Society of Washington 99: 597-607.

Headrick, D. H. and R. D. Goeden. 1990. Resource utilization by larvae of Paracantha gentilis (Dip- tera: Tephritidae) in capitula of Cirsium califor- nicum and C. proteanum (Asteraceae) in southern California. Proceedings of the Entomological So- ciety of Washington 92: 512-520.

1994. Reproductive behavior of California fruit flies and the classification and evolution of Tephritidae (Diptera) mating systems. Studia Dip- terologica 1(2): 194—252.

—.. 1998. The biology of nonfrugivous tephritid

fruit flies. Annual Review of Entomology 43: 217-241.

——.. 1999. Behavior of flies in the subfamily Te- phritinae, pp. 671-707. In Aluja, M. and A. L. Norrbom, eds. Fruit Flies (Tephritidae): Phyloge- ny and Evolution of Behavior. CRC Press, Boca Raton, London, New York, Washington, D.C.

Headrick, D. H., R. D. Goeden and J. A. Teerink, 1996. Life history and description of immature stages of Dioxyna picciola (Bigot) (Diptera: Te- phritidae) on Coreopsis spp. (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 98: 332-349.

Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles.

Shreve, E and I. L. Wiggens. 1964. Vegetation and Flora of the Sonoran Desert. Volume 2. Stanford University Press, Stanford, California. 1740 pp.

Steyskal, G. C. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Special Publication, Entomological Society of Washington. 61 pp.

Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85.

PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 589-601

DESCRIPTION OF AEGESEUCOELA BUFFINGTON, NEW NAME, WITH NOTES ON THE STATUS OF GRONOTOMA FORSTER (HYMENOPTERA: FIGITIDAE: EUCOILINAE)

MATTHEW L. BUFFINGTON

Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A.; current address: Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: mbuff@citrus.ucr.edu)

Abstract.—Aegeseucoela Buffington, a replacement name for Moneucoela Dalla Torre and Kieffer 1910, is described. Aegeseucoela contains two previously described species, Aegeseucoela flavotincta (Kieffer), n. comb., and A. grenadensis (Ashmead), n. comb., both of which are redescribed. The status of Gronotoma Forster is discussed, and the synonymy of Eucoilidea Ashmead with Gronotoma is formally documented. Gronotoma nigricornata Buffington, new name, is proposed to replace Eucoilidea nigricornis Kieffer 1908. Thirty-two new combinations in Gronotoma are given, and a checklist of the world species of Gronotoma is provided. Species in both Aegeseucoela and Gronotoma are common parasitoids of Agromyzidae (Diptera). Aegeseucoela is restricted to the New World tropics and subtropics, and Gronotoma is common in the Afrotropics, as well as

Asia, Australia, and the Palearctic and Nearctic regions.

Key Words:

Eucoiline wasps are endoparasitoids of cyclorrhaphous Diptera inhabiting a variety of habitats. These wasps are generally shiny black to dark reddish brown and range in size from 0.5 mm to 5 mm. The Eucoilinae contain 82 genera and nearly 1000 species, and are by far the most diverse of all figitid subfamilies (Ronquist 1999). Only two ma- jor bodies of work have attempted to clas- sify all of the eucoiline genera (Dalla Torre and Kieffer 1910, Weld 1952), and for the most part, eucoiline classification schemes have resulted in a great deal of chaos (dis- cussed in Nordlander 1982b). Presently available identification keys to eucoiline genera (Dalla Torre and Kieffer 1910, Weld 1952) are largely useless due to the reliance on a few key features, none of which are very dependable.

Nordlander (1976, 1978,

LOSO 19S 12

Aegeseucoela, Gronotoma, Eucoilinae, Figitidae, Agromyzidae, Cynipoidea

1982a, 1982b) was the first to treat eucoi- line classification from a phylogenetic point of view. Through these works, clear generic and species level definitions were provided for the first tme for a number of Palearctic and cosmopolitan taxa. Nordlander (1982b) summarized his findings by proposing in- formal genus groups defined by explicit morphological criteria, a first step towards a more logical and natural classification scheme.

An investigation into the phylogenetics and classification of one of these informal genus groups, the Gronotoma group (Buf- fington, unpublished data), resulted in the identification of a clade of eucoiline wasps of questionable taxonomic placement. Two previously described species were found to belong in this clade, described here as Ae-

geseucoela, and both species are rede-

590

scribed. Both species are restricted to the New World tropics and subtropics where they have been reared on numerous occa- sions from agromyzid flies (O. Lewis, un- published data).

The first indication of the need for a new eucoiline genus was uncovered during an examination of Kieffer’s eucoiline types. The type specimen of Rhabdeucoela flavo- tincta Kieffer did not possess any of the diagnostic features of the genus Rhabdeu- coela Kieffer, and the species was most likely placed in Rhabdeucoela based on the relatively well-developed mesoscutal keel and large scutellar plate (neither of which are universally diagnostic features of Rhab- deucoela). Furthermore, Rhabdeucoela fla- votincta was later moved to two different genera simultaneously by Weld (1952), Mo- neucoela Dalla Torre and Kieffer 1907 and Tropideucoila Ashmead 1903. This species is not readily accommodated in any of the three genera in which it was previously placed, and it was therefore coded separate- ly in a phylogenetic analysis (Buffington, unpublished data), the results of which in- dicate none of the three genera (1.e., Rhab- deucoela, Moneucoela, and Tropideucoila) will remain monophyletic if this species is included within them. Therefore, Aegeseu- coela is proposed to accommodate this spe- cies and a second species (discussed be- low).

The second species, Diranchis grenaden- sis Ashmead, also has a confusing taxo- nomic history. It was one of two species originally included in Moneucoela Dalla Torre and Kieffer 1910, which is itself a junior homonym of Moneucoela Dalla Tor- re and Kieffer 1907. Rohwer and Fagan (1917) apparently missed Kieffer’s 1907 publication containing the description of Moneucoela, and determined that the genus was described as new in Dalla Torre and Kieffer (1910). Further, Rohwer and Fagan (1917) designated Diranchis grenadensis Ashmead as the type species of Moneucoe- la Dalla Torre and Kieffer 1910. Weld (1952) considered this a mistake, removed

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

grenadensis (Ashmead) from type status of Moneucoela Dalla Torre and Kieffer, and designated tinctipennis Kieffer (one of two species described in Kieffer 1907) as the type species of Moneucoela Dalla Torre and Kieffer 1907. Weld (1952) did not include grenadensis within the included species list for Moneucoela, resulting in the placement of the species as incertae sedis.

In their revision of Moneucoela Dalla Torre and Kieffer 1907, Diaz and Gallardo (1998) did not mention Moneucoela Dalla Torre and Kieffer 1910, nor the two species that were included in this genus when it was proposed (Dalla Torre and Kieffer 1910). Of the two species originally includ- ed in Moneucoela Dalla Torre and Kieffer 1910, one is congeneric with flavotincta and is redescribed below as Aegeseucoela. The second species belongs in Zaeucoila Ashmead (Buffington, unpublished data) and will be treated in a subsequent paper. Furthermore, since grenadensis Ashmead was designated as the type species for Mo- neucoela Dalla Torre and Kieffer 1910 by Rohwer and Fagan (1917), and Moneucoela Dalla Torre and Kieffer 1910 is preoccupied by Moneucoela Dalla Torre and Kieffer 1907, Aegeseucoela is proposed here as a replacement name for Moneucoela Dalla Torre and Kieffer 1910.

The second part of this paper is dedicated to a discussion on the status of the eucoiline genus Gronotoma Forster. Gronotoma re- sides within a basal portion of the eucoiline clade (Fontal et al., in preparation), as is the case with Aegeseucoela (though these two genera do not form a monophyletic group) (Buffington, unpublished data). Similar to Aegeseucoela, species of Gronotoma have all been reared from agromyzid flies (Scheffer unpublished data, Davidson 1963, Harding 1965, Viraktamath et al. 1993), mostly found within the genus Melanagro- myza Hendel (Abe and Konishi 1995; Greathead 1969, 1971). The host preference for agromyzid flies appears to be a plesio- morphic feature within the Eucoilinae (Fontal et al. in preparation).

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Gronotoma presently contains 47 de- scribed species (including synonymies pro- posed below), making this genus the most diverse of all eucoiline genera that special- ize on agromyzid hosts. Species of Grono- toma have a worldwide distribution and the Afrotropics are particularly speciose (Quin- lan 1986). Species are also very common in the Oriental (Abe and Konishi 1995), the Palearctic, and the Nearctic regions (Dalla Torre and Kieffer 1910, Weld 1952). Mem- bers of this genus are rarely recorded from the Neotropical Region but are frequently collected throughout subtropical Mexico (Buffington, unpublished data). Because of their potential usefulness in the biological control of pest Agromyzidae and the rela- tively high global diversity of species with- in the genus, the status of Gronotoma is reviewed and includes an important syn- onymy. Phylogenetic evidence (Buffington, unpublished data) supports the conclusions of Hedicke (1930) and Beardsley (1988) with respect to the synonymy of Eucoilidea Ashmead with Gronotoma. A world check- list of the species of Gronotoma is present- ed, with nomenclatural notes where appli- cable.

BMNH_ The Natural History Museum, London, UK (S. Lewis). Bernice P. Bishop Museum, Hon- olulu, HI, USA.

California Academy of Sciences, San Francisco, CA, USA (W. Pu- laski, R. Zuparko).

Canadian National Collection of Insects, Ottawa, Canada (J. Hub- ec Je Read “Masner):

Cornell University Insect Collec- tion, Ithaca, NY, USA (E.R. Hoe- beke).

Institute of Biology at the Acad- emy of the People’s Republic of Romania.

International Centre of Insect Physiology and Ecology, Nairobi, Kenya (R. Copeland). Laboratory of Entomology, Kyo-

BPBM

CAS

CNC

CUIC

IBPR

ICIPE

KPU

59]

to Prefectural University, Kyoto, Japan.

Musée Royale de Il’ Afrique Cen- trale, Tervuren, Belgium. National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, USA (D. Smith).

Zoological Institute, Russian Academy of Sciences, St. Peters- burg, Russia.

Zoologisches Museum, Humboldt Universitat, Berlin, Germany (EF Koch).

MRAC

USNM

ZIN

ZMHB

Type specimens (holotypes and_ para- types) were generously loaned to me by the CAS (containing a large portion of Kieffer’s eucoiline types originally housed in Po- mona), the USNM (containing many of Ashmead’s and Weld’s eucoiline types), the BMNH (containing Quinlan’s eucoiline types), and the ZMHB (containing many of Forster’s eucoiline types).

While sorting unidentified eucoilines in the AEIC in the summer of 2000, I found an extensive series of eucoilines conspecific with the type specimen of Aegeseucoela fla- votincta, all collected from Costa Rica. Ad- ditionally, a long series of Aegeseucoela

flavotincta and of A. grenadensis were sent

to me for identification by O. Lewis (Sil- wood Park, UK); these specimens were of particular importance since host data ac- companied each specimen. Unidentified specimens of African Gronotoma were gen- erously loaned to me by Dr. Robert Cope- land (ICIPE).

Institutions and individuals that donated ethanol preserved specimens for this study were: CNC; TAMU; and Dr. Owen Lewis, Imperial College at Silwood Park, UK. All SEM images utilized in this study were pre- pared digitally on a JOEL JSM-5600 SEM (operated by James Ehrman, Digital Micro- scope Facility, Mt. Allison University, Sackville, NB, Canada).

Terminology follows largely that of Weld (1921, 1952), Nordlander (1982b), Ron-

592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. to lateral ocellus. C, Mesosoma, dorsal view; A = parapsidal hair line; B = well developed lateraldorsal pro- jections of scutellum; C = pronotal triangle.

quist and Nordlander (1989), and Ronquist (1995), with the following modifications: parapsidal ridges is preferred over parapsi- dal furrows, as found in Weld (1952); or- bital furrows and pronotal triangle are new terms and are defined as such below.

Orbital furrow (Fig. 1B).—A _ distinct

Aegeseucoela flavotincta. A, Side view, habitus. B, Head, anterior view; A = orbital furrow complete

groove originating at either the lateral ocel- lus or the lateral side of the torulus (de- pending on the taxon) and lining the inner orbit of the eye, terminating at the clypeal margin after fusing with or paralleling the malar sulcus.

Pronotal triangle (Figs. 1C, 2C).—An

VOLUME 104, NUMBER 3

area on the dorsal surface of the pronotum bordered by the lateral pronotal carina, the pronotal plate and the anterior margin of the mesoscutum; found in genera with well de- veloped pronotal ridges.

Aegeseucoela Buffington, new name

Moneucoela Dalla Torre and Kieffer 1910: 103, 888. Type species: Diranchis gren- adensis Ashmead, designated by Rohwer and Fagan 1917. Preoccupied by Moneu- coela Dalla Torre and Kieffer 1907.

Diagnosis.—Orbital furrows originating either at lateral ocellus or lateral side of to- rulus. Genal carina well developed, often flanged posterior to compound eye. Meso- scutal keel present, at least anteriorly. Par- apsidal ridges absent. Parapsidal hair lines present. Laterodorsal projections of scutel- lum present to absent. RI never tubular or pigmented (radial cell open). Most similar to Zaeucoila and Agrostocynips, but differs by the presence of the extended orbital fur- rows (in some species), the presence of par- apsidal hair lines and R1 incomplete.

Description.—Head: Nearly glabrous, with a few scattered setae along lower face, clypeus, inner orbits of compound eyes, malar space and gena; orbital hair patches present. Ventral %4 of lower face with ad- medial clypeal furrows converging towards clypeus. Orbital furrows present, originat- ing at lateral side of torulus or at lateral ocelli (species dependent), terminating at malar sulcus. Malar sulcus simple. Malar space smooth with a single prominent con- ical protuberance. Genal carina present, ex- tending from malar space to lateral ocelli, often undulating posterior to compound eye:

Antenna: Female, 13 segments, monili- form, semi-clavate; segments 3—13 sube- qual in length; rhinaria present on segments 3-13. Male, 15 segments, filiform; rhinaria present on segments 3—15; segments 4—15 subequal in length. Segment 3 slightly lon- ger than 4, curved outwardly, excavated lat- erally.

593

Pronotum: Pronotal plate wide, with se- tae along dorsal margin; slightly crested and bifurcate dorsally; pronotal fovea open. Pronotal triangle present (Figs. 1C, 2C). Pronotal impression absent. Lateral aspect of pronotum (below pronotal triangle) smooth and glabrous. Lateral pronotal ca- rina absent.

Mesoscutum: Smooth with some setae. Mesoscutal keel present, reaching posterior margin of mesoscutum; tapering posterior- ly. Parapsidal ridges absent. Parapsidal hair lines present (Figs. 1C, 2C). Parascutal im- pression incomplete, narrow. Notauli ab- sent.

Mesopectus: Upper part and lower part of mesopleuron glabrous and smooth. Dor- sal margin of mesopleural triangle well de- fined, rounded ventrally. Mesopleural cari- na simple. Lower part of mesopleuron bor- dered by distinct precoxal carina; anterior surcoxal depression present, reticulate.

Scutellum: Scutellar plate ranging from medium to large; mid pit placed centrally on plate; plate truncated posteriorly; nearly always bearing tubercles and setae on dor- sal surface. Dorsal surface of scutellum re- ticulate; margined laterally and posteriorly. Laterodorsal projections present, present to nearly absent (Figs. 1C, 2B, 2C); posterior projections absent.

Metapectal-propodeal complex: Meta- pectus nearly glabrous with a few scattered setae posteriorly. Spiracular groove with a well defined dorsal margin and a well de- fined to rounded ventral margin. Posterior margin of metapectus ridged. Metapleural ridge absent; submetapleural ridge variable from present to absent. Anterior impres- sions of metepimeron and metepisternum present. Anteroventral cavity semi-circular and setose. Propodeum covered in long se- tae. Lateral propodeal carinae semi-parallel, bowed at junction with auxiliary propodeal carinae; auxiliary propodeal carinae indis- tinct. Nucha glabrous, reticulate.

Wings: Hyaline, with base of forewing sometimes darkened; usually setose basally, always setose apically. R1 incomplete, mar-

594

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

lateral projections of scutellum. C, Mesosoma, dorsal view; A

projection of scutellum; C = pronotal triangle.

ginal cell as long as deep. Apical fringe pre- sent, short.

Legs: Fore and mid coxae about the same size, hind coxa about twice size of either fore or mid coxae. Fore coxa variably cov- ered in long setae; mid coxa with anterior and posterior dorsoventral setal bands; hind

Aegeseucoela grenadensis. A, Side view, habitus. B, Mesosoma, posterolateral view; A = reduced

parapsidal hair line; B = reduced lateraldorsal

coxa with a prominent setal band on hind margin. Femora and tibiae sparsely setose; tarsomeres with dense, appressed setae. Length of hind tarsomere | equal to 0.5 combined length of remaining hind tarso- meres.

Metasoma: Female: Sub-equal in size to

VOLUME 104, NUMBER 3

mesosoma. Base of syntergum with a hairy ring present, ranging from complete to dor- sally bare, comprised of short, semi-ap- pressed setae and longer erect setae; re- mainder of metasoma glabrous. Micropunc- tures present on posterior 4 of syntergum, and on remaining terga. Terga posterior to syntergum directed posteroventrally, result- ing in a 70 degree angle between syntergum and remaining terga. Male: As in female but terga posterior to syntergum abruptly angled ventrally, resulting in a 90 degree angle between syntergum and remaining terga.

Biology.—I have examined specimens reared by O. Lewis from species of agro- myzid flies in the genera Haplopeodes Steyskal (on Solanaceae) and Calycomyza Hendel (on Fabaceae).

Distribution.—Neotropical Region: Cos- ta Rica, Belize, Mexico (Veracruz, Tamau- lipas), Panama. Nearctic Region: USA: AZ. Previously only known from Guatemala (flavotincta) and Grenada (grenadensis).

Etymology.—Aeges; from Greek my- thology, the name of Athena’s dreaded shield, used here to reference the broad, shieldlike pronotal plate present in this ge- nus; eucoela, a suffix frequently used by J.J. Kieffer in his treatments of the Neo- tropical Eucoilinae.

Comments.—Though this genus is close- ly related to Zaeucoila and Agrostocynips, phylogenetic evidence (to be found in a forthcoming publication by the author) sug- gests that neither of these genera will re- main monophyletic if the species of Aege- seucoela are placed within them. The par- apsidal hair lines, extended lateral protu- berances on the scutellum and incomplete R1 vein on the forewing are reliable auta- pomorphies for the genus, and reliable syn- apomorphies for the species within the ge- nus.

INCLUDED SPECIES

Aegeseucoela flavotincta (Kieffer), n. comb. Rhabdeucoela flavotincta Kieffer

595

1908:46, holotype in CAS (#10537). Re- described below.

Aegeseucoela grenadensis (Ashmead), n. comb. Diranchis grenadensis Ashmead 1900: 248, holotype in BMNH. Rede-

scribed below.

Aegeseucoela flavotincta (Kieffer), new combination (Fig. 1)

Description.—As in description of genus except as follows: Head: Orbital furrows originating at lateral ocelli (Fig. 1B); genal carina continuing to near lateral ocelli, un- dulating posterior to compound eye (Fig. 1A). Pronotum: Pronotal crest prominent, bifurcate; pronotal ridge well developed. Scutellum: Scutellar plate large, nearly round; laterodorsal projections of scutellum sometimes well developed (Fig. IC). Me- tapectal-propodeal complex: Submeta- pleural ridge usually well developed, con- necting ventral margin of spiracular groove with posterior margin of mesopleuron. Wings: Base of forewing occasionally dark- ened; base of forewing ranging from gla- brous to setose; forewing always setose api- cally. Metasoma: Hairy ring at base of syn- tergum often highly reduced (narrow), but always present.

Material examined.—Holotype 2, Champerico, Guatemala. Coll. Baker. CAS #10537; the type specimen is in good con- dition, with Kieffer’s original determination label (a large red label), the collection data labels, depository label and my determina- tion label (slender white label). Additional material: BELIZE: Las Cuevas, Chiquibul Forest, Cayo District, 550 m, (various dates between Oct. 1997 and Sept. 1998), O.T. Lewis (13°23 5"'6)., BOLIVIA:* Yungas; XII.4.84, 2,400 m, Luis Pefia (1 2); COS- TA RICA: S. Rosa Park, Guan., various dates between 27.V.1976 and 10.VIII.1978, D.H. Janzen, Dry Hill (46 6, 90 2). MEX- ICO: Veracruz, 2 km SW Fortin, 8°54'N, 97°00’ W, 2,700’, 23.V1.1997, J.B. Woolley, screen sweep (1 2); Veracruz, El Crucero nr Puente Nacional, 19°20'’N, 96°26'W,

596

13.V1.1997, L.A. Wilson & J.B. Woolley (1 2); Tamaulipas, 97 km Ciudad Victoria, Hwy 70, 3.VII.1986, G. Zolnerowich & R. Trevino (1 2). PANAMA: Colon Prov., 2 km S Sabanitas, 4—15.VII.1999, 120 m, Gillogly & Woolley, MT 99/033 (1 @). U.S.A: Arizona, Portal, 19—23.VIII.1987, H. & M. Townes (4 ¢).

Distribution.—Neotropical and southern Nearctic regions (see above list of locali- ties).

Biology.—I have examined specimens reared from the agromyzid flies Haplopeo- des sp. on Solanum erianthum D. Don (So- lanaceae) and Calycomyza hyptidis Spencer on Hyptis capitata Jacq. and H. urticoides Kunth (Lamiaceae) (data from O. Lewis).

Aegeseucoela grenadensis (Ashmead), new combination (iss)

Description.—As in description of genus except as follows: Head: Orbital furrows originating at torulus; genal carina reduced, non-undulating, terminating at dorsal mar- gin of compound eye (Fig. 2A). Pronotum: Pronotal crest reduced, sometimes absent; pronotal ridge sometimes absent. Scutel- lum: Scutellar plate ranging from medium to small; laterodorsal projections usually re- duced/absent (Figs. 2B, 2C). Metapectal- propodeal complex: Submetapleural ridge completely reduced. Wings: Base of fore- wings occasionally darkened; usually entire forewing surface is setose. Metasoma: Hairy ring at base of syntergum thick and densely pubescent.

Material examined.—Holotype @; Bal- thazar (windward side), Grenada, West In- dies. H.H. Smith (Coll.), BMNH; the type specimen is in poor condition, with the head plus thorax on one end of a pinned card, and the metasoma on the other end. The diagnostic features discussed below for this genus are all visible. Two ‘type’ labels are present on the specimen, one with a red circle and the second labeled ‘BM Type Hym., y. 50’. Under this lies a label with ‘paratype’ printed on it. Below that is Ash-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mead’s original determination label (in Ashmead’s hand). Below that is the collec- tion data, and finally below that, my des- ignation label. Additional material: BE- LIZE: Las Cuevas, Chiquibul Forest, Cayo District, 550 m, (various dates between Oct. 1997 and Sept. 1998); O:1. Lewist@7 a2; 20 3). MEXICO: Veracruz, 0.7 mi N Jilo-

tepecs 19°36 N, 196756 W, 32680" Bscreen Sweep; 14.V1.1997; L.Aq Wilson 5B: Woolley (1 2). PANAMA, Prov. Colon,

Quebrada Lopez, MT, 2—4.VII.1999, A. Gillogly (1 2). WENEZUELA: Merida, Merida City, 8°35'54"N, 71°08'42”W, 1860 m, sweep veg. along trib. to Chama R., 3.V.1981, L. Masner (1 @).

Distribution.—Neotropical Region (see above list of localities).

Biology.—I have examined specimens reared from the agromyzid flies Calycomy- za verbenivora on Verbena sp. (Verbena- ceae); Calycomyza c.f. cassiae (Frost) on Senna cobanensis (Britton) H.S. Irwin & Barneby (Fabaceae); Haplopeodes sp. on Solanum erianthum (Solanaceae) (data from O. Lewis).

Comments.—The type specimen of Dir- anchis grenadensis Ashmead, as noted above, is labeled ‘paratype.’ In the original description of D. grenadensis, Ashmead stated “‘described from 1 female speci- men.” Since the single specimen, the lo- cality data, and the description all agree, this must be the holotype.

Gronotoma Forster

Gronotoma Forster 1869: 342, 346. Type species: Gronotoma sculpturata Forster 1869: 346, by original designation.

Eucoilidea Ashmead 1887: 150, 154. Type species: Eucoilidea canadensis Ashmead, by subsequent designation (Ashmead 1903); synonymy by Hedicke (1930) and Beardsley (1988).

Eucoelidea Dalla Torre 1893: 1901: 159. Emendation.

Afrostilba Benoit 1956: 544. Type species: Afrostilba nitida Benoit, by monotypy. Synonymy by Quinlan (1986).

15; Kieffer

VOLUME 104, NUMBER 3

Ashmead (1887) proposed the genus Eu- coilidea to accommodate E. longicornis and E. canadensis, but did not specify a type species. Later, Ashmead (1903) designated Eucoilidea canadensis as type species. Dal- la Torre (1893) emended the original Ash- mead spelling of Eucoilidea to Eucoelidea, and cites the former in brackets. Eucoelidea was recognized in Kieffer (1901), and Kief- fer (1907, 1909) described four new species (and two varieties) in ‘“Eucoelidea Ashm.’ None of Kieffer’s treatments of Euwcoelidea mention the name’s emendation from Ew- coilidea. Based on Article 33.2.1 of the ICZN (1999), Eucoelidea is an emendation of Eucoilidea and is not available. Burks (1979) reported two apparent misspellings of Eucoilidea, namely Eucoilidia (Kieffer 1907) and Eucoelidia (Kieffer 1909); after consulting the original references, neither of these misspellings were found to exist.

It is apparent from Ashmead’s (1887) original description of Eucoilidea that he made no comparison with Gronotoma be- fore proposing the new genus, and Hedicke (1930) was the first to propose Eucoilidea as a synonym of Gronotoma. Weld (1952) and Quinlan (1986, 1988), however, sub- sequently maintained Eucoilidea as distinct from Gronotoma. Beardsley (1988) again synonymized these two genera. Though the holotype of E. canadensis is in poor con- dition, with great portions of the body ob- scured by blackish glue, features such as the notauli, the lateral pronotal carinae and the scutellar plate are all clearly visible. Comparing these features with those found in Gronotoma sculpturata clearly indicate that indeed, Eucoilidea is a junior synonym of Gronotoma, supporting the decisions of Beardsley (1988) and Hedicke (1930).

Species of Gronotoma can be recognized by the following diagnostic features: lateral pronotal carina present (a distinct ridge, dis- tal to the lateral margins of the pronotal plate, that give species in the this genus the appearance of having a large pronotal plate); notauli present and well-developed in nearly all species; scutellar plate large;

597

hairy ring at base of syntergum absent. Spe- cies of Gronotoma are most easily confused with Diglyphosema Forster, but easily dis- tinguished from that genus by having the scutellum meeting the scutellar plate at an acute angle (meets at a 90 degree angle in Diglyphosema), and the scutellar plate not as elongate (though some species of Gron- otoma tend to have an oval scutellar plate). Previous works treating the synonymy of Eucoilidea with Gronotoma focused pri- marily on the type species; thus, as indicat- ed in the list below, most of the described species have not been formally transferred Hence, 32 new combina- tions are proposed, mainly reflecting the taxonomic work on this genus by Weld (1952) and Quinlan (1986). Species for which type material (holotypes and/or para- types) were examined are indicated by an * The reader should also note that Kieffer (1901) did indeed treat four species in (and we, Gronotoma carinata (Cresson), G. minor (Provancher), G. nigricornis Kietfer and G. ovalis (Thomson) (see in list below).

to Gronotoma.

Gronotoma not Eucoilidea),

Included Species

adachiae Beardsley 1988: 39, holotype in BPBM.

*advena (Quinlan), n. comb. Eucoilidea advena Quinlan 1986: 262, 263, holotype in MRAC, paratypes in BMNH.

allotriaeformis (Giraud). Eucoila allotriae- formis Giraud 1860: 142. Gronotoma al- lotriaeformis: Forster 1869: 346. Type depository presently unknown.

*arcuata (Kieffer), n. comb. Eucoelidea arcuata Kieffer 1909: 65. Type in CAS.

*bakeri (Kieffer), n. comb. Eucoelidea bakeri Kieffer 1907: 107. Type in CUIC.

*bakeri var. cupularis (Kieffer), n. comb. Eucoelidea bakeri var. cupularis Kieffer 1907: 108. Type in CAS.

*bakeri var. flavipes (Kieffer), n. comb. Eu- coelidea bakeri var. flavipes Kieffer 1907: 108. Type in CAS.

*bucca (Quinlan), n. comb. Eucoilidea

598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bucca Quinlan 1986: 263, holotype in MRAC, paratypes in BMNH.

*canadensis (Ashmead). Eucoilidea cana- densis Ashmead 1887: 154. Gronotoma canadensis: Hedicke 1930; Beardsley 1988. Holotype in USNM.

carinata (Cresson). Eucoila ? carinata Cresson 1865: 6. Gronotoma carinata: Kieffer 1901: 159. Type depository pres- ently unknown.

*compressa (Quinlan), n. comb. Eucoilidea compressa Quinlan 1986: 263, 264, ho- lotype in MRAC, paratypes in BMNH.

*conversa (Quinlan), n. comb. Eucoilidea conversa Quinlan 1986: 264, holotype in MRAC, paratypes in BMNH and MRAC.

*crenulata (Kieffer), n. comb. Eucoilidea crenulata Kieffer 1908: 47. Type in CAS.

*dilitata (Kieffer), n. comb. Eucoelidea dilitata Kieffer 1907:108. Type in CAS.

domestica Girault 1932: 3. Type depository presently unknown.

*dubia (Quinlan), n. comb. Eucoilidea du- bia Quinlan, 1986: 264, 265, holotype in BMNH, paratypes in BMNH and MRAC.

extraria (Quinlan), n. comb. Eucoilidea ex- traria Quinlan 1986: 265, holotype and paratype in MRAC.

*fetura (Quinlan), n. comb. Eucoilidea fe- tura Quinlan 1986: 265, 266, holotype in MRAC, paratypes in BMNH.

fulvicornis (Hedicke). Ganaspis fulvicornis Hedicke 1913: 445. Gronotoma fulvicor- nis: Hedicke 1934: 704. Holotype and four paratypes in ZMHB.

*furcula (Quinlan), n. comb. Eucoilidea furcula Quinlan 1986: 266, holotype in BMNH, paratypes in BMNH and MRAC.

*fuscipes (Kieffer), n. comb. Eucoelidea fuscipes Kieffer 1907: 112. Type in CAS.

gracilicornis Cameron 1889: 15. Type de- pository presently unknown.

guamensis (Yoshimoto), n. comb. Eucoili- dea guamensis Yoshimoto 1962a: 107, holotype and paratypes in BPBM.

hiranoi Abe and Konishi 1995: 309-311, holotype and paratypes in KPU.

insularis Ashmead 1895: 743, holotype in BMNH.

lacerta (Quinlan), n. comb. Eucoilidea lac- erta Quinlan 1986: 266, 267, holotype and paratype in MRAC.

lana (Quinlan), n. comb. Eucoilidea lana Quinlan 1986: 267, holotype and _ para- type in MRAC.

*leptis (Quinlan), n. comb. Eucoilidea lep- tis Quinlan 1986: 267, 268, holotype in BMNH.

longicornis (Ashmead), n. comb. Eucoili- dea longicornis Ashmead 1887: 154. Type depository presently unknown.

maquilingensis (Kieffer), n. comb. Eucoil- idea maquilingensis Kieffer 1914: 184— 185. Type depository presently unknown.

*marcellus (Quinlan), n. comb. Eucoilidea marcellus Quinlan 1986: 268, holotype in BMNH, paratypes in BMNH and MRAC.

*maurt (Quinlan), n. comb. Eucoilidea mauri Quinlan 1986: 268, 269, holotype and paratype in BMNH.

melanagromyzae Beardsley 1988: 40, ho- lotype in BPBM.

micromorpha (Perkins). Eucoilidea micro- morpha Perkins 1910: 676. Gronotoma micromorpha: Beardsley 1988: 38. Eu- coilidea rufula Yoshimoto 1962b: 845, synonymy by Beardsley (1988), holotype and paratypes in BPBM.

minor (Provancher). Eucoila minor Pro- vancher 1888: 398. Gronotoma minor: Kieffer 1901: 159. Type depository pres- ently unknown.

nigra lonescu 1963: 10, holotype in IBPR.

*nigricornata Buffington, new name. Eu- coilidea nigricornis Kieffer 1908: 48. Preoccupied by Gronotoma nigricornis Kieffer 1901: 1595 Paratypes miCUIce.

nigricornis Kieffer 1901: 159. Type depos- itory presently unknown.

*nitida (Benoit), n. comb. Afrostilba nitida Benoit 1956: 544, holotype in MRAC, paratypes in BMNH. Eucoilidea nitida: Quinlan 1986: 269.

ovalis (Thomson). Cothonaspis ovalis Thomson 1877: 817. Gronotoma ovalis: Kieffer 1901: 159. Type depository pres- ently unknown.

*pallida (Quinlan), n. comb. Eucoilidea

VOLUME 104, NUMBER 3

pallida Quinlan 1986: 269—270, holotype in BMNH, paratypes in BMNH MRAC.

*parma (Quinlan), n. comb. Eucoilidea parma Quinlan 1986: 270, holotype in BMNH, paratypes in BMNH and MRAC.

and

presently unknown.

*perangusta (Quinlan), n. comb. Eucoili- dea perangusta Quinlan 1986: 270, 271, holotype and paratypes in BMNH.

quadrisulcata (Hedicke), n. comb. Eucoil- idea quadrisulcata Hedicke 1922: 229. Two syntypes in ZMHB.

rufipes (Gillette), n. comb. Eucoilidea ru- fipes Gillette 1891: 205. Type depository presently unknown.

*sculpturata (Forster). Eucoila sculpturata Forster 1855: 257. Gronotoma sculptur- ata: Forster 1869: 342, 346. Holotype in ZMHB.

*seychellensis Kieffer 1911: 309. Holotype and paratypes in BMNH.

sugonjaevi Belizin 1973: 22, holotype in ZIN; not examined but characters pre- sented in the original description suggest this species probably belongs in Nordlan- deria.

*trulla (Quinlan), n. comb. Eucoilidea trul- la Quinlan 1986: 271, holotype in BMNH, paratype in MRAC.

*tyrus (Quinlan), n. comb. Eucoilidea tyrus Quinlan 1986: 271, holotype in BMNH.

*yurundiensis (Benoit), n. comb. Eucoilidea urundiensis Benoit 1956: 548, holotype in MRAC, paratypes in BMNH and MRAC.

Additional material examined.—Sp. 1: Nearctic Region: Canada (AB, BC, MB, ONVOU; PO): USA dE. MD..NY, PA, TX, VA). Neotropical Region: Mexico (Tamau- lipas, Chiapas).

Sp. 2: Nearctic Region: Canada (AB, MB); USA. (AZ, CA, FL: IN, MO; NGC, NM, PA, TN, TX, WA). Neotropical Re- gion: Mexico (Veracruz); Venezuela (Mer- ida).

599

Sp. 3: Nearctic Region: Canada (ON); USA (NC).

Specimens of other species from: Neo- tropical Region: Mexico (Oaxaca, Guerre- ro); Ecuador; Nicaragua. Afrotropical Re- gion: Uganda; Zaire; Kenya.

ACKNOWLEDGMENTS

I thank Dr. Bob Wharton and Dr. Jim Woolley for guidance in this project at Tex- as A&M University; I also thank Dr. Géran Nordlander and Dr. Fredrik Ronquist for additional guidance in understanding eucoi- line morphology, taxonomy, and evolution. Special thanks to all those individuals listed under “‘Materials and Methods” who as- sisted with the loan of material for my ex- amination, including James Ehrman _ for preparation of SEM’s and Dr. Owen Lewis for the use of host records. Additional thanks to Dr. Kathy Schick, Dr. David Smith, and one anonymous reviewer for critical comments on an earlier version of this manuscript. Finally, thanks to my wife for her unending support of my research. This research was entirely funded by the NSF PEET Project, #DEB9712543, award- ed to Drs. Wharton and Woolley. All re- search was carried out in the Department of Entomology, Texas A&M University.

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. 1907. Beschreibung neuer parasitischer Cy-

nipiden aus Zentral- und Nord-Amerika (En

Aleman). Entomologische Zeitschift 21: 70-162.

. 1908. Nouveau Proctotrypides et Cynipides

d’Amerique, recueillis par N. Baker chef de la

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ciété Scientifique de Bruxelles 32: 7—64.

1909. Description de nouveaux Cynipides

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turalle (du Département de la Moselle) de Metz

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. 1911. Hymenoptera, Cynipidae (of the Seys-

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Nordlander, G. 1976. Studies on Eucoilidae (Hyme- noptera: Cynipoidea) I. A revision of the North- western European species of Cothonaspis Htg. with a description of a new species and notes on some other genera. Tijdschrift Voor Entomologie 97: 65-77.

. 1978. Revision of genus Rhoptromeris Fors- ter, 1869 with reference to north-western Euro- pean species. Studies on Eucoilidae (Hymenop- tera: Cynipoidea) II. Entomologica Scandinavica 9: 47-62.

. 1980. Revision of the genus Leptopilina Fors- ter, 1869 with notes on the status of some other genera (Hymenoptera: Cynipoidea: Eucoilidae). Entomologica Scandinavica 11: 428—453.

. 1981. A review of the genus 7rybliographa Forster, 1869 (Hymenoptera, Cynipoidea: Eucoil- idae). Entomologica Scandinavica 12: 381—402.

. 1982a. Identities and relationships of the pre- viously confused genera Odonteucoila, Coneucoe- la and Trichoplasta (Hymenoptera, Cynipoidea: Eucoilidae). Entomologica Scandinavica 13: 269— 292.

. 1982b. Systematics and phylogeny of an in- terrelated group of genera within the family Eu- coilidae (Insecta: Hymenoptera, Cynipoidea). Doctoral dissertation. University of Stockholm, Sweden. 34 pp.

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. 1988. A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). Bulletin of the Brit- ish Museum 56: 171-229.

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PROC. ENTOMOL. SOC. WASH. 104(3), 2002, pp. 602-613

ENTOMOBALIA, NEW GENUS, THE FIRST MEMBER OF NYCTELIINI (COLEOPTERA: TENEBRIONIDAE) FROM BRAZIL

GUSTAVO E. FLORES AND CHARLES A. TRIPLEHORN

(GEF) Instituto Argentino de Investigaciones de las Zonas Aridas (ADIZA CRIGYT): Casilla de Correo 507, 5500 Mendoza, Argentina (e-mail: gflores@lab.cricyt.edu.ar); (CAT) Department of Entomology, Museum of Biological Diversity, The Ohio State Uni- versity, 1315 Kinnear Road, Columbus, OH 43212, U.S.A. (e-mail: ctriplhrn@aol.com)

Abstract.—Entomobalia, new genus, (Pimeliinae: Nycteliini) is described from north- eastern Brazil and is the first genus of the tribe Nycteliini recorded for that country. This new genus is created on the basis of two previously described species: Asida platynotos Perty and Asida picta Perty (new combinations in Entomobalia), formerly placed in the genus Scotinus Kirby (Pimeliinae: Asidini). Sixteen different character states among the species of Scotinus and the two species assigned to Entomobalia are discussed, 11 of which are shared by Entomobalia with all or some genera of Nycteliini and not with the remaining species of Scotinus. A description for the genus and redescriptions of the two species are provided. Main diagnostic characters for Entomobalia are in sexual dimor- phism and male and female genitalia. Habitus photographs, illustrations of external mor- phology, internal skeletal anatomy, genitalic features, and a distribution map are included.

Key Words: ica, Brazil

Nycteliini is an endemic Neotropical tribe of Pimeliinae (Doyen 1993), with 285 species distributed in Argentina, Chile, Bo- livia, Peru, Paraguay, and Uruguay (Fig. 1) (Flores 1997). The species of Nycteliini are currently arranged in 11 genera, the nine recognized in previous revisions: Gyrioso- mus Guérin-Méneville, Pilobalia Burmeis- ter, Entomoderes Solier, Nyctelia Latreille, Epipedonota Solier, Psectrascelis Solier, Scelidospecta Kulzer, Auladera Solier, Mi- tragenius Solier (Kulzer 1954, Flores 1997), the recently described genus Pata- gonogenius (Flores 1999), and the restored genus Callyntra Solier (Flores and Vidal 2000).

Our examination of a single specimen collected in Brazil by W. Mann from the Field Museum of Natural History (Chicago,

Tenebrionidae, Pimeliinae, Asidini, Nyctelini, Entomobalia, South Amer-

USA) led us to think we had discovered a new species of Nycteliini. More recently we discovered a long series of 80 specimens of this species and another series of 25 spec- imens of another smaller species, both col- lected in northeastern Brazil by W. Mann, deposited in the National Museum of Nat- ural History, Smithsonian Institution, (Washington, DC). One specimen of the longer series had been determined by Kulz- er as Scotinus platynotos (Perty). After re- questing a loan of all the species of Scotin- us present in The Natural History Museum (London, UK), we confirmed that both spe- cies had already been described by Perty (1830), the larger as Asida platynotos and the smaller as Asida picta. Finally, study of the types of Perty from the Zoologische Staatssammlung, Miinich (Germany) con-

VOLUME 104, NUMBER 3

firmed the identity of both series of these species.

When Perty (1830) described these two species, he placed them in the genus Asida Latreille (Asidini). Laporte (1840) consid- ered that they should be included in the ge- nus Scotinus Kirby (Asidini). Scotinus picta had not been mentioned in the literature for 160 years, but in 1935, Blair expressed the opinion that S. platynotos “‘is a somewhat aberrant Pilobalia’. Gebien (1910) listed this species under Scotinus, but later he list- ed it under the nycteliine genus Pilobalia (Gebien 1937). This means that both au- thors believed this species belongs to the tribe Nycteliini rather than the Asidini. However, Kulzer (1954) still accepted this species as a member of the genus Scotinus. Through a detailed discussion of the char- acters, we demonstrate that Scotinus platyn- otos and S. picta are not congeneric with the remaining species of Scotinus, that they are not Asidini, that they share most of these characters with all or some genera of Nycteliini, and that they deserve recogni- tion as a separate genus, which we have named Entomobalia. This new genus exhib- its a mosaic of characters present in other genera of Nycteliini, especially Pilobalia and Entomoderes. The inclusion of these two species in any other known genus of Nycteliini would imply a completely differ- ent concept and redefinition of that genus. In addition, the male and female genitalia show unique apomorphies.

Species of Nycteliini inhabit arid and semiarid environments, mainly in the bio- geographical provinces Chaco, Monte, Pampa, Central and Arid Puna, Prepuna, Coquimbo, Santiago, Payunia, Western, Central and Fueguinan Patagonia, and San Jorge Gulf (Morrone 2001). The discovery of these two species from northeastern Bra- zil, at least 3,000 km from the Chaco, the closest point of distribution of a genus of Nycteliini (Entomoderes), was unexpected. Thus the range of the tribe is greatly ex- panded to the Caatinga biogeographical province (Morrone 2001). It is possible that

603

other species of Nycteliini may be found in the Cerrado biogeographical province, Which lies between Caatinga and Chaco Gries al):

The objectives of this study are to de- scribe this new genus of Nycteliini, rede- scribe the two species assigned to the new genus, and compare the distinctive charac- ter states of the species of Scotinus (Asi- dini) with the two species assigned to En- tomobalia (Nycteliini), which had previous- ly been placed in Scotinus.

Specimens were obtained from the fol- lowing institutions: The Natural History Museum, London, UK (BMNH), Field Museum of Natural History, Chicago, IL, USA (FMNH), National Museum of Nat- ural History, Smithsonian Institution, Washington, DC, USA (USNM), and Zool- ogische Staatssammlung, Miinich, Germa- ny (ZSMC). Thanks to the kind generosity of the USNM, we distributed some speci- mens of both species in the following in- stitutions: California Academy of Scienc- es, San ‘Francisco, GA,. USA (CASE), Field Museum of Natural History, Chica- go, IL, USA (FMNH), Instituto Argentino de Investigaciones de las Zonas Aridas, Mendoza, Argentina (IADIZA), Museo Argentino de Ciencias Naturales Bernar- dino Rivadavia, Buenos Aires, Argentina (MACN), Museu de Zoologia da Univer- sidade de Sao Paulo, Sao Paulo, Brazil (MZSP), and The Ohio State University, Columbus, OH, USA (OSUC).

Body length was measured dorsally, along the midline, from anterior margin of labrum to elytral apex. For paraproct/coxite length (P/C) we used the ratio proposed by Doyen (1993). For basal lamina of the teg- men/lateral styles length (B/E) and median lobe/tegmen length (L/T) we used the ratios proposed by Flores (1996). Drawings were made with a camera lucida adapted to a ste- reoscopic microscope.

TAXONOMIC PLACEMENT OF ENTOMOBALIA

The two species that we have assigned to the new genus Entomobalia (Nycteliini)

604

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs 1

Known distribution of the genera of Nycteliini (lined area). Black square: distribution records for

Entomobalia platynota; black circle: distribution records for E. picta. | and 2: Caatinga and Cerrado biogeo-

graphical provinces (sensu Morrone 2001).

had previously been placed in the genus Scotinus, which belongs to the Asidini, a large tribe of Pimeliinae with more than 1,000 species distributed in North and South America, Africa south of Sahara, Madagascar, and the Mediterranean area (Koch 1955). The species of Scotinus are endemic to Brazil (Gebien 1937). We have studied the following characters within the species of Scotinus, some of which have been used to study the relationships be-

tween the tribes of Pimeliinae (Doyen 1993) and to define the tribe Nycteliini (Flores 1997), finding that Scotinus platyn- otos and S. picta share all of these charac- ters with all or some genera of Nycteliini and not with the remaining species of Sco- tinus. The species of Scotinus examined from BMNH are the following (T includes type specimens): S. crenicollis Kirby (T) (type species), S. crucifer Eschscholtz, S. propius Wilke, S. quadricollis Eschscholtz,

VOLUME 104, NUMBER 3

S. quadricollis incisus Wilke (T), S. tuber- culatus Eschscholtz, S. tuberculatus dispar Wilke (T), S. chapadensis Wilke (T), and S. bajulus Wilke (T).

fe

i)

Size of antennomere I1: Very small, even rudimentary in Scotinus (Figs. 2— 3); in Entomobalia and Nycteliini well developed, sometimes equal to or longer than antennomere 10 (Figs. 4—7). Subgenal process (defined by Doyen 1993: 454): Subadjacent with mentum in Scotinus (Solier 1836: plate XIII, Figs. 1-2, Doyen 1993: Figs. 22—25); in En- tomobalia and Nycteliini it is remote from mentum (Doyen 1993: Fig. 21). Position of ligula: Nearly concealed be- neath mentum in Scotinus (Solier 1836: plate XIII, Figs. 1-2); in Entomobalia it is entirely exposed anterad of mentum (Fig. 8), with the articulating membrane visible. Within Nycteliini, this character state is shared with Gyriosomus, Pilob- alia and Entomoderes.

Relative size of ligula: Less than half the size of mentum in Scotinus (Solier 1836: plate XII, Figs. 1-2); in Entomobalia it is larger than half of mentum (Fig. 8). Within Nycteliini, this character state is shared with Gyriosomus, Pilobalia and Entomoderes.

Shape of mentum: Cordiform in Scotin- us, With anterior margin twice as long as posterior margin (Solier 1836: plate XIII, Figs. 1-2). Subtrapezoidal in En- tomobalia, with anterior margin 1.5 times the length of posterior margin (Fig. 8). Within Nycteliini, this character state is shared only with Pilobalia (Fig. 10). Mesepisternum: Small, not reaching the mesocoxal cavities in Scotinus (Fig. 11); in Entomobalia and Nycteliini it is large, reaching the mesocoxal cavities and closing laterally (Fig. 12). Mesepimeron: Long, reaching the base of elytral epipleuron in Scotinus (Fig. 11); in Entomobalia and Nycteliini it is short, not reaching the base of elytral epipleuron (Fig. 12).

9.

605

Metepimeron: Transverse in Scotinus, widely separating the metacoxal cavities from the elytral epipleuron (Fig. 11); in Entomobalia and Nycteliini it is not ev- ident, since the metacoxal cavities are very close to the elytral epipleuron (Fig. (2).

Lateral Closed laterally by metepimeron and first visible abdominal sternum in Sco- tinus (Fig. 11); closed laterally by me- tepisternum in Entomobalia (Fig. 12). Within Nycteliini, this character state is

closure of metacoxal cavities:

shared with Gyriosomus, Pilobalia and Entomoderes.

10. Distance between meso- and metacox-

ll.

ae: Does not exceed half the metacoxal length in Scotinus (Fig. 11); in Ento- mobalia and Nycteliini it exceeds half the metacoxal length (Fig. 12). Abdominal margin of elytral epipleu- ron: Curved where it contacts meso- and metathorax in Scotinus (Fig. 11); in En- tomobalia and Nycteliini it is straight where it contacts meso- and metathorax (Rise 12):

Additional character states which differ

in Scotinus and Entomobalia:

ha

i)

oS)

Antennal length: Short in Scotinus, not reaching the middle of the lateral margin of pronotum; longer in Entomobalia, ex- tending beyond posterior margin of pronotum.

Shape and width of antennomere 10: Oval, wider than long in Scotinus, wider than more proximal antennomeres (Figs. 2—3); subrectangular, longer than wide in Entomobalia, no wider than more prox- imal antennomeres (Figs. 4—5).

Ventral femoral surface: Covered by sparse setae in Scotinus; densely setose on proximal %4 in Entomobalia.

Elytral epipleuron: Not evident in Sco- tinus (Fig. 11); evident throughout in Entomobalia (Fig. 12). In addition, in Entomobalia the elytral epipleuron is equal in width throughout (Fig. 12); within Nycteliini this character state is

606 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2-12. External structure. 2-7, Antennae in dorsal view. 2, Scotinus crenicollis. 3, S. crucifer. 4, En- tomobalia platynota. 5, E. picta. 6, Pilobalia decorata (Erichson). 7, P. oblonga (Blanchard). 8—10, Labium and mentum in ventral view. 8, Labium of Entomobalia platynota. 9, Mentum of Entomoderes satanicus Waterhouse. 10, Mentum of Pilobalia decorata. 11-12, Mesothorax, metathorax and abdomen in ventral view. 11, Scotinus quadricollis. 12, Entomobalia picta (female). Abbreviations: 1, ligula, m, mentum, p, labial palpus, mes, mese- pisternum, mep, mesepimeron, met, metepisternum, mtp, metepimeron. Scale bar = | mm.

VOLUME 104, NUMBER 3

shared with Gyriosomus, Pilobalia and Entomoderes.

Nn

Sexual dimorphism: Species of Scotinus do not exhibit sexual dimorphism; spe- cies of Entomobalia exhibit four char- acters different in male and female (see below).

In his cladistic analysis of Pimeliinae, Doyen (1993) found that all genera of As- idini (from South African, Madagascan, Mediterranean, North and South American regions) constitute a monophyletic group defined by six synapomorphies, five of which are not present in bridge of tentorium absent or incomplete,

Entomobalia:

antennae with ten segments plus the re- duced eleventh antennomere, abdominal la- terotergites extremely small, apicodorsal lobe of proctiger ending at coxite base, and baculus of proctiger extending proximad, equal to baculus of paraproct. Furthermore Doyen (1993) pointed out that the first two are synapomorphies unique to Asidini with- in the subfamily Pimeliinae.

Within Pimeliinae, Entomobalia must be placed within the Asidine clade (Doyen 1993) by having multiple, long, slender spermathecal tubes which open as a fascicle into the base of the accesory gland duct or into the vagina near the duct (Figs. 17—18). Within the Asidine clade of Doyen (1993), Entomobalia belongs to the subclade of South American tribes Nycteliini, Physo- gasterini and Praocini by having metendos- ternite arms fused with mesocoxal inflex- ions. Entomobalia is placed in the tribe Nycteliini according to the definition of that tribe by Flores (1996, 1997) and the follow- ing change in that tribal concept should be mentioned: in the female genitalia of En- tomobalia the basal lobe of the coxite is separated vertically from apical lobe (Figs. 17-18), while in the remaining genera of Nycteliini it is separated horizontally from apical lobe (Flores 1996).

The most recent key provided for the genera of Nycteliini is that by Flores

607

(1997), which is modified at couplet 8 to separate out Pilobalia and Entomobalia:

8. Pronotum with a short central-posterior carina and two longer longitudinal carinae, lateral margin with lobe (Flores 1997: Fig. 25); fem- ora with umbilicate setae; mentum transverse (Fig. 9) Pronotum lacking carinae, lateral margin con-

Meg eat ca OO, O NC Entomoderes Solier cave, lacking lobe; femora with simple setae; mentum subtrapezoidal (Figs. 8, 10) ..... 8a

8a. Mentum impunctate or with very small punc- tures; antennomere 10 subspherical or pyri- form (Figs. 6-7); male with protibiae straight and not expanded apically (Flores 1997: Fig.

ZG)! + ia coves Beh ee ae

— Mentum with large punctures; antennomere

Pilobalia Burmeister

10 subrectangular (Figs. 4—5); male with pro-

tibiae curved inward and expanded apically

(Figs: 19=20)) 2%. See Entomobalia, new genus

Entomobalia Flores and Triplehorn, new genus

Type species.—Asida platynotos Perty, present designation.

Diagnosis.—Distinguished from other Nycteliini by the following combination of characters: ligula sclerotized, ventrally ex- posed, equal to half mentum area; labial palps inserted on posterior half of ligula; mentum subtrapezoidal; pronotum smooth, without punctures, striae or carinae, lateral margin concave, posterior angles acute, overlapping elytral humeri, epipleuron equal in width throughout; metacoxal cav- ities closed laterally by metepisternum; ventral surface of femora densely setose; median lobe of aedeagus subapically ex- panded and strongly sclerotized on distal third; sexual dimorphism: male with proti- bia curved inward and expanded apically, inner surfaces of tibiae densely setose on distal half, and central area of metasternum with setae; female with protibia straight and not expanded apically, inner surfaces of tib- iae with normal setae, and central area of metasternum glabrous and shiny.

Description.—Length, 12.1—-19.9 mm; width, 6.4—11.3 mm. Body and legs brown to black, with antenna and maxillary palp brown. Pronotum and elytron with two

608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 13-18.

Internal skeletal anatomy and female genitalia of Entomobalia spp. 13-14, Tentoria. 13, En-

tomobalia platynota. 14, E. picta. 15—16, Metendosternites in posterior view. 15, Entomobalia platynota. 16, E.

picta. 17-18, Ovipositor (ventral view), spiculum and internal female reproductive tract. 17, Entomobalia pla- tynota. 18, E. picta. Abbreviations: c, coxite, 0, oviduct, p, paraproct, s, spermatheca, sag, spermathecal accessory

gland, sp, spiculum, v, vagina. Scale bar = | mm.

kinds of short setae, one stout, dark brown, and other finer, golden or light brown. Head: Epipharynx with anterior margin entirely concealed beneath labrum. Labrum, clypeus and frons with abundant short, golden setae. Clypeus with anterior margin concave. Clypeal suture defined by a deep depression with setae at antennal insertion

level. Frons without longitudinal or lateral grooves. Eye reniform. Antenna long, ex- tending beyond posterior margin of prono- tum; antennomeres subrectangular, with two different kinds of pubescence: one short and abundant on entire surface, and second consisting of long, scattered setae; unique apical semicircular tomentose sen-

VOLUME 104, NUMBER 3

Figs. 19, 20.

sory patch on antennomere 9. Ligula scler- otized, articulated with mentum by a nar- row membrane and exposed ventrally, equal to half mentum area (Fig. 8); labial palpi inserted on posterior half of ligula (Fig. 8). Ligula and mentum with setae arising from large punctures. Mentum subtrapezoidal (Fig. 8). Submentum posteriorly continuous with gula. Tentorium with medial straight bridge (Figs. 13-14).

Thorax: Pronotum short (length: prono- tum/elytron = 0.33), pubescent and smooth, without punctures, striae or carinae; disc convex; anterior margin slender and central area not broadened; anterior angles round- ed; lateral margin simple, slender, concave, widest behind mid point; posterior angles acute, overlapping elytral humeri; posterior margin biconcave, as wide as base of elytra. Proepisternum with or without grooves and with sparse not umbilicate setae. Proster- num arched, not extended over mesoster- num. Mesosternum inclined forward, sepa- rated from prosternum. Scutellum visible. Meso- and metepisternum without grooves.

609

Habitus in dorsal view. 19, Entomobalia platynota, male. 20, E. picta, male.

Elytron: Dorsal surface, pseudopleuron and epipleuron impunctate; with one or two carinae, space between external carinae and lateral margin without grooves; lateral mar- gin straight and sharp, epipleuron equal in width throughout, texture similar to that of elytra.

Legs: Procoxal cavity closed posteriorly. Metacoxae separated by one metacoxal width, enclosed laterally by metepisternum. Ventral surfaces of trochanters pubescent, brushlike. Ventral surfaces of femora dense- ly setose on proximal %4. Ventral surfaces of tarsi bearing abundant decumbent setae.

Sexual dimorphism: Male with protibia curved inward and expanded apically, inner surfaces of tibiae densely setose on distal half, and central area of metasternum with setae. Female with protibia straight and not expanded apically, inner surfaces of tibiae with normal setae, and central area of me- tasternum glabrous and shiny.

Internal skeletal anatomy: Mesendoster- nite with a long and slender dorsal arm, lon- ger than horizontal arm. Metendosternite

610

(Figs. 15—16) with arms long, extending be- yond mesocoxal inflections about half dis- tance to tergum, stem equal to metacoxal width, width of stem exceeding length, and stem narrow at middle. Elytral-abdominal fusion accomplished by a ridge in the ely- tral epipleuron which interlocks in a lon- gitudinal groove of abdominal sterna.

Male genitalia: Rods of sternum IX close at basal third, distance between them not exceeding width of aedeagus. Dorsal membrane of proctiger concave, with two sclerotized areas. Basal lamina of tegmen long (B/E > 1.00). Lateral styles of tegmen distally close, with apex straight and long setae on ventral surface; ventral proximal margin convex in ventral view, projecting dorsally over median lobe. Median lobe long (L/T > 1.00), sheath-shaped, one third the width of lateral styles of tegmen, ex- panded subapically and strongly sclerotized on distal third, with apex acute and straight.

Female genitalia (Figs. 17-18): Spicu- lum with arms “V-shaped. Paraprocts moderate (1,2 = P/C = 2,0), glabrous. Cox- ites with setae, basal lobe of coxite extend- ed over paraproct and separated vertically from apical lobe, baculi of coxite inclined 45°: midventral sclerite distally broadened. Proctigeral baculus equal to length of par- aproct baculus. Vagina saccate; spermathe- cal tubes shorter than vagina length, all similar in width and branching pattern: spermathecal accesory gland longer than vagina, with duct annulate and thick.

Etymology.—The name of the genus re- fers to the similarity to the Nycteliine gen- era Entomoderes and Pilobalia. Gender feminine.

Distribution and habitat.—Brazil: States of Ceara, Rio Grande do Norte, Pernam- buco and Bahia, in the Caatinga biogeo- graphical province (Morrone 2001) (Fig. 1). W. Mann, the collector of the large series of both species in Baixa Verde, Rio Grande do Norte, stated that “the country was arid, with much scrub and cacti, but little life in evidence” (Mann 1948: 91).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

KEY TO THE SPECIES OF ENTOMOBALIA

1. Pronotum with lateral margin reflexed; elytron with lateral margin single and protuberances ir- regularly and sparsely distributed on dorsal surface; length usually greater than 15 mm (Fig. 19) E. platynota (Perty)

— Pronotum with lateral margin not reflexed; el- ytron with lateral margin double and lacking protuberances; length usually less than 13 mm (Fig. 20) E. picta (Perty)

Entomobalia platynota (Perty), new combination (Piessaeaes., 135 ise eZ)

Asida platynotos Perty 1830: 56, plate 12, fig. 2.

Scotinus platynotos: Laporte 1840: 208; La- cordaire 1859: 165.

Scotinus platynotus: Gemminger and Har- old 1870: 1880 (cat.); Gebien 1910: 139 (cat.).

Pilobalia platynota: Blair 1935: 104; Ge- bien 1937: 753 (cat.); Blackwelder 1945: 519 (cat.); Kulzer 1954: 254 (rev.); Flo- res 1997: 16 (ist).

Scotinus platynotos: Kulzer 1954: 265.

Redescription.—Length, 15.0-19.9 mm; width, 8.0—-11.3 mm. Body and femora brown to black, with antenna, tibia and tarsi brown. Stout dark brown setae and golden or light-brown finer setae uniformly and sparsely distributed on pronotum and dorsal surface of elytron, both forming velvet-like patches only on posterior third of elytron. Antenna long, extending three antenno- meres beyond posterior margin of prono- tum. Pronotum with lateral margin reflexed. Male metasternum with tuft of setae on cen- tral area, brush-like. Metepisternum with not umbilicate setae. Elytron with lateral margin single and protuberances irregularly and sparsely distributed on dorsal surface, with one feebly raised complete or incom- plete carina, close to lateral margin (Fig. 19): in some specimens with a secondary carina close to suture, consisting of aligned protuberances. Male with protibia expanded apically, but not excavated. Last abdominal segment truncate in males and females. In addition to three characters of sexual di-

VOLUME 104, NUMBER 3

611

Figs. 21-24.

Male genitalia of Entomobalia spp. in dorsal and ventral views. 21, 22, Entomobalia platynota.

23, 24, E. picta. Abbreviations: bl, basal lamina of tegmen, Is, lateral styles of tegmen, ml, median lobe. Scale

bar = | mm.

morphism at generic level, first abdominal sternum with a tuft of brush-like setae on central area in males and glabrous and shiny in females.

Male genitalia.—Lateral styles of tegmen with apex wide, widest at mid point, with long setae on distal half of ventral surface (Fig. 21). Median lobe with apical aperture large, strongly expanded subapically (Fig. D2).

Specimens examined.—Lectotype: [Type] [Brasilia/Scotinus/platynotos/Perty | (ZSMC). To fix the current interpretation of this name and to ensure stability, we are hereby designating this lectotype: [Lectoty- pus/Asida_platynotos/Perty, 1830/Des. G. Flores-/C. Triplehorn 2001]. Non-type spec- imens. BRAZIL: Rio Grande do Norte. Baixa Verde, W. Mann, 75 [55 (USNM), 3

(CASC), 4 (FMNH), 7 CUADIZA), 2 (MAECN), .2)'\(MZSP):'2 (OSUC)]? Ceara- Mirim, W. Mann, | (USNM). Pernambuco. Ouricuri,' ‘TW=1982,"E.C 9 "Gux (ex Bufo stomach), 3 (OSUC). Bahia. Near Queima- das, 11-VI-1915, PG. Russell, 2 (USNM);

S. Salvador, 1918, 2 (USNM); Barra, 1 (BMNH); without more precise data: 2 (BMNH).

Entomobalia picta (Perty), new combination (Figs. ie, 127 A, MG 182205235 24)

Asida picta Perty 1830: 56, plate 12, fig. 3.

Scotinus picta: Laporte 1840: 208; Lacor- daire 1859: 165.

Scotinus pictus: Gemminger and Harold