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A. z < ES Ws = * = x = Zz = Yip, = — AS" 2. = 27 fk ENS? ‘a > = > = ays > 3 2 n z ap) = INOSHLINS S31YVYGI7 LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI_N¥INOS LIBRARIES SMITHSONIAN y Gy o NOILNLILSNI LIBRARIES NOLLMIILSNI LIBRARIES NOILALILSNI f AITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYVYSIT LIBRARIES | SMITHS 4X VOL. 92 JANUARY 1990 NO. 1 4 (ISSN 0013-8797) PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS DAVIS, D. R.—Neotropical Microlepidoptera XXIII. First report of the family Eriocottidae from the New World, with descriptions of new taxa .............2 220 e eee eee eee eee DUFFIELD, R. M. and C. H. NELSON.—Seasonal emergence patterns and diversity of Ple- coptera on Big Hunting Creek, Maryland, with a checklist of the stoneflies of Maryland... 120 EASTOP V. F. and D. J. VOEGTLIN.—Taxonomic notes on some North American aphids 115 GAGNE, R. J. and R. W. DUNCAN.—A new species of Cecidomyiidae (Diptera) damaging shoot tips of yellow cypress, Chamaecyparis nootkatensis, and a new genus for two gall MNGPES ONaGHPESSACEAG, fa. cde MEA er Era eas atime cs Bae Savaciea cin teers eis = 146 HALSTEAD, J. A.—Review of Haltichella Spinola in the Nearctic Region (Hymenoptera: Chal- CICA AG) Pepe tae a tee tats peat de Mie ea eo Mae ote a IA POT dre oak ice srsfetetgs ele Fake Swe 153 HODGES, R. W. and V. O. BECKER—Nomenclature of some neotropical Gelechiidae (Lepi- GOPtera) be ake ote ae teens erate keer ett each de et Peationtphes Sone laithaygete atyausteis ale, ob scayete 3% 76 HUNG, A. C. F. and P. W. SCHAEFFER.—Isozyme analysis in six populations of Pediobius foveolatus (Crawford) (Hymenoptera: Eulophidae) ......................000002200005- 160 JENKINS, J.— Mating behavior of Aciurina mexicana (Aczél) (Diptera: Tephritidae) ........ 66 KROMBEIN, K. V.—Systematic notes on some Bethylidae from Botswana: Epyrinae (Hyme- HOUSE ANAS on ae Mihi ap hPa ona ab cn ae'r ea doe One Se BIT REST: Bee eaers 4 G98 MAIER, C. T.—First distributional records of Tabanidae (Diptera) in Connecticut .......... 139 MASON, W. R. M.—Cubitus Posterior in Hymenoptera .......... Ag get Te Sn Roepe tte 93 MATHIS, M. L. and D. E. BOWLES.—Three new species of microcaddisflies (Trichoptera: iydraptilidae) from the Ozark Mountains W.S-Agy jee ns: 2 isos bak Palle eek oa was eek 86 MORSE, J. C. and C. B. BARR.— Unusual caddisfly (Trichoptera) fauna of Schoolhouse Springs, Louisiana, with description of a new species of Diplectrona (Hydropsychidae) .......... 58 NORRBOM, A. L.—The identity of the Genus Hexaresta Hering (=Hyponeothermara Hardy, HaSyis) Goi pteraepHritiGae) eas crs). csi en coh benedeni 4 op ram eee OE athe vis Ateneo (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1990 JEFFREY R. ALDRICH, President NorMAN E. WooDLEY, 7reasurer Davip R. SmiTH, President-Elect Gary STECK, Program Chairman RICHARD G. Rossins, Recording Secretary GEOFFREY B. WHITE, Membership Chairman HO us B. WILLIAMS, Corresponding Secretary F. CHRISTIAN THOMPSON, Past President JAMES B. STRIBLING, Custodian ROBERT D. Gorpon, Editor THOMAS J. HENRY, Associate Editor Publications Committee DONALD R. DAvis DONALD R. WHITEHEAD GEORGE C. STEYSKAL F. CHRISTIAN THOMPSON Honorary President Curtis W. SABROSKY Honorary Members Louise M. RUSSELL ALAN STONE THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M. 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PLEASE SEE P. 318 OF THE APRIL, 1989 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. Editor: Robert D. Gordon, Systematic Entomology Laboratory, ARS, % Department of Entomology, Smith- sonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: T. Henry, Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 30 January 1990 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 1-35 NEOTROPICAL MICROLEPIDOPTERA XXIII. FIRST REPORT OF THE FAMILY ERIOCOTTIDAE FROM THE NEW WORLD, WITH DESCRIPTIONS OF NEW TAXA DONALD R. DAviIs Department of Entomology, U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Abstract.—Crepidochares subtigrina Meyrick from Brazil is redescribed and transferred to Eriocottidae. Formerly this monotypic genus and species had been included in Tineidae. In addition, four new species of Crepidochares, C. aridula and C. austrina from Chile, C. colombiae from Colombia, and C. neblinae from Venezuela are described for the first time. The discovery of these taxa marks the first record of the primitive tineoid family Eriocottidae for the New World. Supplemented by numerous illustrations, the morphology of these and related Old World taxa are summarized. Key Words: Recent investigations on the tineoid com- plex by the author together with an earlier study by Nielsen (1978) have shown the pre- dominantly Old World family Eriocottidae to be among the most ancient of the ditry- sian Lepidoptera. Because the Eriocottidae are the only ditrysian moths known to pos- sess microtrichia randomly scattered over all wing surfaces, I consider this family to be the most primitive member of the Tine- oidea. More importantly, this implies that among the extant Lepidoptera, they most resemble the stem ancestor of the Ditrysia. Prior to this paper, no Eriocottidae were reported to occur in the New World. An eriocottid, Crepidochares subtigrina, was previously described by Meyrick (1922) from the Amazon, but this species had been regarded as a tineid. Fieldwork in southern Chile by the author, R. E. Brown, O. Kars- holt, and E. S. Nielsen in 1981 resulted in collections of two new species. A recent multidisciplinary biological survey of Cerro de la Neblina in Venezuela produced another undescribed species. The latter appears to Lepidoptera, Eriocottidae, Crepidochares, biogeography be a sister species of yet another new mon- tane species from Sierra del Libano, Colom- bia, thus bringing the total species of Erio- cottidae known for South America to five. To supplement the descriptions of the new American taxa, a brief review of the biology and morphology of the family is provided. Depositions of specimens referred to in this paper are: ANIC for Austalian National Insect Collection, CSIRO, Canberra, Aus- tralia, BMNH, British Museum (Natural History), London, England; NHNS, Museo Nacional de Historia Natural, Santiago, Chile; USNM, National Museum of Natu- ral History (formerly the United States Na- tional Museum), Smithsonian Institution, Washington, D.C., UCVM, Universidad Central de Venezuela, Maracay, Venezuela; and ZMUC, Zoologisk Museum, Univer- sitets Kobenhaven, Copenhagen, Denmark. BIOLOGY Distribution.—Prior to this report, the Eriocottidae as reconstituted by Nielsen (1978), were known to occur only in the Old 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = colombiae @Neblinae * subtigrina Map |. World. The genus Eriocottis ranges from southern Europe (Spain, Italy), northern Af- rica (Algeria), southern USSR, Asia Minor to Taiwan. I have examined the Taiwanese species, Eriocottis flavicephalana Issiki (Figs. 8, 115-119), and found it to agree in all respects to E. fuscanella Zeller, with the no- table exception that the maxillary palpi are five segmented and the ocelli are reduced (thus agreeing with Issiki’s (1930) original description). Another recognized genus of the subfamily Eriocottinae, Deuterotinea, is believed to be less widely distributed and confined largely to southern Europe and Asia Minor, from Spain eastward to Syria, Israel, Iraq and southern USSR. A single genus and species of Eriocottinae, Eucryptogona tri- chobathra Lower, occurs in New South Wales, Australia (Nielsen, in litt.). Distibution of Crepidochares in northern South America. In the New World, Eriocottidae are re- stricted to South America where two genera are recognized, both allied to Eriocottis. Thus far, most of the South American species have been found only in temperate forests or high elevations (Maps 1, 2). Cre- pidochares subtigrina, however, was col- lected at Parintins along the Amazon River, which suggests the existence of more species through the vast neotropical lowlands. In 1981 I collected two, very distinct species of Eriocottinae in two different biotic re- gions of Chile (Map 2). Crepidochares arid- ula, new species, was found only in the drier, more northern, Central Valley and envi- rons, whereas Crepidochares austrina, new species, was collected at several localities in the wetter, Nothofagus dominant, Valdivi- an forests to the south (see Davis, 1986, VOLUME 92, NUMBER 1 Map 2, for limits of biotic regions). At ap- proximately the same time, Nielsen and Karsholt collected C. austrina near Valdivia and Anticura, Chile. Compsoctena, the largest genus and sole representative of the Old World subfamily Compsoctenidae, appears the most diver- sified through the Ethiopian Region (sub- Saharan Africa), where a majority of the known species occur, with numerous species also reported from India to mainland China and Taiwan and through portions of In- donesia. As shown by Dierl (1970), most of the known species of Compsoctena were proposed in other genera, often in Melasina or in genera now synonymized under Compsoctena. Several of these species are yet to be studied and still reside in their original genus. One such species, A/avona thaitesii Walsingham, has been examined in this study and transferred to Compsoctena. Life history.—As is true for most families of moths, one of the most pressing needs for fieldwork among the Eriocottidae is for studies on life history and the immature stages. As pointed out by Nielsen (1978), little is known of their biology. Apparently few members of Eriocottidae have been reared, and their larval habits have largely been speculated (e.g. possibly feeding in de- cayed wood, leaves, or as stemborers). Adults appear to be univoltine and are ac- tive in the spring, both in the northern and southern hemispheres. Adult females of the middle eastern, steppe inhabiting Deutero- tinea are wingless. Their larvae are detri- tophagous and construct silken tunnels often with ventilation tubes amongst grass litter (Zagulayev 1973, 1988). All specimens of Crepidochares collected by me were taken in ultraviolet light traps. The general struc- ture of their highly extensible ovipositors indicates that the eggs are inserted into crev- ices within the host substrate. Only slightly more is known about the habits of Compsocteninae. Dierl (1970) provides some evidence that the larvae live underground in silk and earthern tubes, or * aridula ea: @ austrina ees * ‘ aS | | eo 7 | x — \ Ae 9 \ | batt | \_ 9 | e Ss ——/196) _¢ ie as | | 136 ke Rs ‘a i \ | | pw 5 - i im 7 \ SA c= : SNS Net =) ~ 0 eT, > N\ + 40 | 40 | \ | \ | Lm 44) | | Map 2. Distribution of Crepidochares in Chile. (as may be true for Compsoctena reductella (Walker)) bore into decaying plants such as Artemisia and Rubus. MorRPHOLOGY The major morphological features of Er- iocottidae, as they relate to systematics, have been summarized by Dierl (1970) and Niel- sen (1978). As discussed by these authors, the principal synapomorphy of the family is the presence of a fourth pair of short, anterior apophyses located dorsally within the eighth segment of the female abdomen. In the subfamily Eriocottinae these are mostly fused in an X-shaped configuration (Figs. 121, 127, 130). The dorsal anterior apophyses may be either X-shaped or sep- arate in Compsoctena. The new Chilean species, Crepidochares aridula, is unique in possessing a fifth pair of ventral apophyses (Fig. 130) within the eighth segment in ad- dition to the fourth, X-shaped, dorsal pair. No other species of Lepidoptera is known to have developed this many pairs of ab- dominal apophyses. The Compsocteninae are easily distin- guished from Eriocottinae in being generally larger, without ocelli, with greatly reduced, two segmented maxillary palpi and a mi- nute haustellum (Figs. 20-21). Further- more, the male antenna is shortly bipecti- nate in Compsocteninae, compared to simple in female Compsoctena and in both sexes of Eriocottinae. The antenna of male Crepidochares subtigrina 1s unusual in pos- sessing two ventral pairs of short tubercles bearing elongate sensilla chaetica (Figs. 58— 60). This relatively aberrant species also has lost the ocelli, and possesses reduced, three segmented maxillary palpi and porrect la- bial palpi (Figs. 14, 15), similar to the Aus- tralian Eucryptogona trichobathra. Accord- ing to Nielsen (in litt.), the latter differs from Crepidochares subtigrina in possessing more slender wings, flagellomeres without lobes, and male valvae more similar to Eriocottis. Although Nielsen described the antennal pecten as absent in Friocottis, | have found it present in all genera, although sometimes less distinct in Compsoctena. The presence of an antennal pecten is the plesiomorphic condition in Tineoidea as well as the non- ditrysian moths. The wings of Eriocottidae are unique among the ditrysian moths in retaining mi- crotrichia randomly scattered over all wing surfaces. This resembles the plesiomorphic condition present in nearly all non-ditrysian families. In Eriocottidae the microtrichia are relatively short and sparsely distributed (Figs. 26, 27, 44, 48-50, 64, 70, 72), with some variation to be noted. In at least one species, Eriocottis fuscanella Zeller, micro- trichia are largely absent from the dorsal surface of the hindwing and restricted main- ly to the wing base. Microtrichia have been PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lost over nearly all of the wing surfaces in all other Ditrysia, with the notable excep- tion of the tineid genus Eudarcia, where they are absent only over the dorsal surface of the forewings. The possibility exists for microtrichia to be even more developed in other Tineidae not yet examined. If such a condition were found, then this would ne- cessitate a re-evaluation of the supposed basal position of the Eriocottidae among the Ditrysia. The male retinaculum was similar in all genera examined in consisting of an elon- gate, flaplike fold from the ventral costal margin and extending over the base of Sc (Figs. 24, 46, 47, 66-68). The outer margin of the retinaculum in Crepidochares aridula (Figs. 46, 47) appears more revolute than in Eriocottis or Compsoctena. In addition to sharing a similar, plesiomorphic reten- tion of wing microtrichia, Eudarcia also possesses a male retinaculum similar to that of Eriocottidae. The retinacula of 7 other tineid genera examined, representing major subfamilies, were found to resemble the more typical ditrysian type (Davis, 1n press) consisting of a slender cuticular lobe arising from the underside of the subcostal vein. Although the male retinacular lobe arises slightly anterior to Sc in other tineoids and apparently in most ditrysian families, in some genera (e.g., Afteva) it originates on Sc as in the Tineidae examined. The curled apex of the lobe forms a short tube for firmly clasping the male frenulum. As is true in the case with microtrichia, too few tineid genera have been examined to determine the systematic significance between the sub- costal costal fold (Eudarcia, Eriocottidae) and the curled subcostal lobe types (most Tineoidea and higher Ditrysia). Both types of male retinacula occur within the subdi- vision Monotrysia, as well as within a single family (Palaephatidae, Davis 1986, Psychi- dae). Studies to date indicate the subcostal costal fold type to represent the plesio- morphic state. Although Eudarcia displays VOLUME 92, NUMBER 1 certain plesiomorphic similarities to Erio- cottidae, no female Eudarcia examined to date has been observed to possess a fourth pair of abdominal apophyses. Consequent- ly, this genus has not been included within Eriocottidae. The mesofurcasterna of Eriocottidae are similiar in possessing relatively broad, stout, secondary arms (Figs. 88, 92). The apices of the secondary arms differ in Compsoc- tena in having the lateral branch reduced to a small tubercule with an attached tendon (Fig. 92) and in the greater elongation of the mesal branch. The metafurcasterna of the various genera are also of similar mor- phology, with the furcal apophyses of Compsoctena more attenuated (Figs. 93, 94). The apices of the metafurcal apophyses of Crepidochares aridula and C. austrina are truncate, with those of other Crepidochares and Eriocottis being somewhat intermedi- ate in development (Fig. 91). Ifthe examples studied are typical of their respective subfamilies, then the Eriocottin- ae and Compsocteninae may also differ in leg structure. The pretarsal unguitractor plates of Crepidochares neblinae and C. ar- idula are less developed with only two to three ranks of scutes per transverse row (Figs. 33, 57). As is typically the condition in large moths, the unguitractor plate of Compsoc- tena thwaitesii (Walsingham), new combi- nation, is larger and with a much greater number of scutes (8 to 12 ranks, Fig. 75). Apparently what often occurs in this, as well as in other families I have examined, is that as the plate enlarges or decreases with body size, the relative size of the scutes does not change proportionally but individual scutes are added or lost in number to cover the appropriate area. A more significant differ- ence between the two groups may involve the epiphysis, which is lost in some species (Nielsen 1978). The epiphysis of Crepido- chares (Figs. 29-31, 52-55), and Eriocottis are of the standard form, with a comb (or pecten) of stout spines along the inner, cleaning edge and covered elsewhere with tightly appressed, imbricated, scale-like spines. In Compsoctena thwaitesii (Figs. 78- 81) the epiphysis is more elongate and more specialized in lacking both pecten and im- bricated spines. The stiff, tibial scales nor- mally opposite an epiphysis also appear to be lacking in this species. The apex is slight- ly broadened and, similar to all other sur- faces of the epiphysis, densely covered with deciduous scales. The only spines present are minute in size and scattered along the anterior surface (Fig. 81). The two principal morphological systems probably used most frequently in Lepidop- tera systematics, wing venation and male genitalia, appear relatively conservative among Eriocottidae. Wing venation varies little among all known genera, with no stalked or branched veins present and with R5 terminating at or slightly above the apex of the forewing. The male genitalia appear rather uniform within genera (as is typical for another tineoid family, Psychidae), with a prominent spinose lobe arising from either the ventral margin of the cucullus in the Old World genera (and in many Psychidae), or from the distal margin of the sacculus in the only recognized Neotropical genus, Crepi- dochares. The male genitalia of the latter exhibit greater morphological differences between species than is generally true for other eriocottid genera, particularly with re- gard to the development of the spinose, sac- cular lobe. Crepidochares Meyrick Crepidochares Meyrick, 1922: 601.— Fletcher, 1929: 58.—Clarke, 1970: 36.— Davis, 1984: 4, 21. Type species.—Crepidochares subtigrina Meyrick, 1922; by monotypy. Adult.—Small, pale yellowish to brown moths with forewings variably and often in- distinctly banded with brown to fuscous. Ocellus usually present; maxillary palpus 3- 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5 segmented; labial palpus porrect or up- curved. Male valva with a spinose process from sacculus. Female genitalia with usually four pairs of apophyses or five pairs in C. aridula. Length of forewing 4.6-9.5 mm. Head: Vestiture rough, consisting of long piliform scales with simple, acute apices. Antenna usually filiform, with two pairs of ventral tubercles per segment in C. subti- grina, 0.4-0.7 the length of forewing, 31- 40 segmented; scape with pecten of 6-10 piliform scales; flagellum usually with dor- sal half covered with moderately broad scales, ventral half naked except for dense sensilla, completely encircled by scales in C. neblinae. Eye round, relatively well devel- oped; interocular index ranging from 0.63- 1.1; eye index 0.73-0.92. Ocellus usually present, absent in C. subtigrina. Chaetose- mata absent. Pilifers (Fig. 37) well devel- oped, bearing 7-8 elongate bristles directed mesally and nearly touching at midline. Mandible either vestigial or absent. Max- illary palpus usually 5-segmented and equalling or slightly longer but more slender than labial palpus and about half the length of haustellum; basal three segments the shortest and apical two the longest and ap- proximately equal in length; maxillary pal- pus less than one fifth the length of labial palpus and composed of three short seg- ments in C. subtigrina. Haustellum elon- gate, often twice the length of labial palpus; distal half to two thirds externally covered with short overlapping plates (Fig. 40). La- bial palpus 3-segmented, moderately well developed, usually slightly upcurved (Fig. 13) with subapical sensory pit reduced (Figs. 42, 43); palpus larger, relatively smooth and porrect in C. subtigrina (Figs. 14, 15). Thorax: Forewing moderately slender; length 3—3.6 the width. Radius 5-branched; all veins arising separate; RS usually ter- minating just before apex, or at apex (in C. subtigrina), R1 arising usually from basal third of discal cell, or mesad (in C. austrina). Media 3-branched, all veins separate. Dis- cal cell 0.55-0.63 the length of forewing. Accessory and intercalary cells usually de- veloped. 1A and 2A separate at basal 0.3- 0.4, forming an anal loop. Retinaculum of male composed of a broad fold arising im- mediately under costal margin and partially extending over base of Sc (Figs. 24, 46, 47); retinacular fold absent in female, instead retinaculum consisting of a row of elongate, piliform scales from base of Sc. Microtri- chia generally distributed over all wing sur- faces in C. neblinae, most concentrated over basal half of discal cell or underside of fore- wing in C. aridula, not examined closely in other species. Hindwing nearly as broad as arising separate; base of M usually forked within cell, rarely entire. Frenulum single in male (Fig. 51), 2-4 bristles in female. Foreleg with pectinated epiphysis (Figs. 29- 31, 52-55) approximately 0.4 the length of tibia; outer surface covered with flat, mod- erately broad, imbricate spines (Figs. 31, 55). Midleg with an elongate, apical pair of tibial spurs of unequal length. Hindleg with two pairs of elongate tibial spurs of unequal length, one pair apical and other pair arising from outer 4; basal tarsomere with row of 6-8 small spinose setae and apices of all tarsomeres with 3 small setae. Prothorax (Fig. 86) with sternum moderately devel- oped, lightly sclerotized on either side of basisternum; patagium greatly reduced, nearly touching opposite member at dorsal midline; scutum greatly reduced, triangular, tapering to form slender, poorly differen- tiated scutellum. Mesothorax (Fig. 88) with secondary arms of furcasternum relatively broad and stout, abruptly terminating in a pair of short, acute processes; forked ends of secondary arms widely spaced, a distance about equal to width of mesothoracic phrag- ma. Metafurcasternum (Fig. 90) with a pair of stout, either truncate or attenuate furcal arms, each with a single tendon directed anteriorly from anterior apex. Abdomen: Relatively simple, without specialized process, coremata, or corethro- gyne. Second sternum of tineoid type, with VOLUME 92, NUMBER | a pair of slender apodemes projecting an- teriorly from sternum; a minute tubercule and tubercular plate immediately laterad to base of apodeme. Male genitalia: Uncus divided into two short, acute lobes, otherwise not differen- tiated from relatively broad, hoodlike tegu- men. Vinculum moderately short, either V-shaped or attenuated into a distinct sac- cus. A more or less sclerotized, plate like subscaphium sometimes present which is fused medially to slender U- or V-shaped gnathos. Juxta and soci absent. Valva with basal half (sacculus) moderately broad, either equal to or twice the width of usually more slender distal half; (cucullus); ventral margin of cucullus without prominent spi- nose lobe (pollex) but with a variably de- veloped spinose lobe from distal margin of saccular lobe; lobe largest in C. neblinae, most reduced in C. aridula. Aedoeagus rel- atively slender, short, without cornutt; phal- lobase well developed, nearly twice the length of aedoeagus and enclosing distal part of ejaculatory duct. Female genitalia: Ovipositor greatly elongated, telescoping, with usually four or rarely five pairs of rodlike apophyses: pos- terior pair the longest, extending from A7 to caudal apex of abdomen (A10); a much shorter ventral pair located entirely within A10; anterior pair elongate and extending caudad into A8; a single pair of shorter, often mostly fused, ““X-shaped”’ dorsal apophyses located entirely within A8, and an additional short, separated pair located within A8 of C. aridula; apex of ovipositor soft, trilobed (one lobe minute), and setose. Ductus bursae highly variable, extremely short and broad in C. austrina to long and slender in C. neblinae; ductus seminalis usually joined midway along ductus bursae. Corpus bursae moderately enlarged, usually with a single, variably shaped signum; sig- num absent in C. aridula. Discussion.—A single synapomorphy— the presence of a slender, spinelike process arising from the saccular lobe at the base of the male valva—distinguishes this South American genus from its Old World sister- group, Eriocottis. The type species of Cre- pidochares, however, exhibits several apo- morphies that strongly suggest further division within the New World species. Ma- jor among these features which sets C. sub- tigrina apart are the loss of ocelli, fasciculate and pedicellate antennal sensilla, reduction of the maxillary palpi to three short seg- ments, and the relatively smooth and por- rect labial palpi. Because no synapomorphy is known to link the other four species, all have been retained within Crepidochares. KEY TO THE SPECIES OF CREPIDOCHARES 1. Ocellus absent. Maxillary palpus reduced, three segmented. Labial palpus porrect eee _ C. subtigrina Meyrick ~ Ocellus present Maxillary palpus five seg- mented. Labial palpus disinctly upcurved 2. Male genitalia with saccular process minute, less than one third the width of valva (Figs. 106, 110). Distribution southern Chile .. 3 — Male genitalia with saccular spine elongate, more than half the width of valva (Figs. 97, 103). Distribution northern South America 4 3. Forewing with R1 arising from middle of discal cell. Male genitalia (Fig. 84) with subscaphium relatively broad and elongate, arising from gna- thos near insertion of valva; valva with a small spinose process arising from saccular lobe free of valva. Female genitalia (Figs. 126, 127) with four pairs of apophyses; signum present C. austrina Davis, new species - Forewing with R1 arising from basal third of discal cell. Male genitalia (Fig. 108) with sub- scaphium reduced, arising from gnathos above insertion of valva; valva with minute spinose process not projecting beyond margin of valva. Female genitalia (Fig. 130) with five pairs of apophyses; signum absent aE _C. aridula Davis, new species 4. Male with length of saccular process equalling width of valva and terminating in a broad trun- cate spine (Fig. 97); apical half of valva (cu- cullus) broader than basal half. Female with fourth pair of apophyses within A8 mostly fused and X-shaped (Fig. 121) Nm C. neblinae, new species - Male with length of saccular process less than width of valva and terminating in a slender, minute spine (Fig. 103); apical half of valva more slender than basal half. Female with fourth pair of apophyses convergent but not fused (Fig. 1X) AE ee oe ey Bare ee C. colombiae, new species Crepidochares neblinae Davis, New SPECIES Figs. 1, 16, 17, 22-33, 82, 95-99, 120-122; Map 1 Adult (Fig. 1).—Length of forewing: 4, 8 mm; 2, 8.2-8.6 mm. A small moth with grayish forewings marked by three more or less distinct, fuscous cross bands and scat- tered spots; ocellus present; labial palpus slightly upcurved; male valva with a prom- inent, blunt tipped, spinose lobe arising from apex of sacculus; female with X-shaped apophyses within eighth abdominal seg- ment. Head: Vestiture mixed, mostly fuscous near middle bordered by tufts of cream to buff scales laterally and at lower part of frons. Ocellus well developed. Antenna 0.55-0.6 the length of forewing, 39-40 segmented; scape light to medium brown, with a pecten consisting of 8-10 dark brown piliform scales; scales not forming an eyecap; flagel- lum alternately ringed with dark fuscous and light brown; scales encircling each segment with basal ring fuscous; flagellomeres smooth except for a minute, apical mid- dorsal process (Fig. 23); sensilla relatively short and not fasiculate nor born on tuber- cules (Figs. 22, 23). Maxillary palpus elon- gate, 5-segmented: vestiture variable, light to dark brown. Labial palpus slightly up- curved, mostly dark brown to fuscous lat- erally and pale buff to cream mesally, with apices of second and third segments pale buff; numerous cream to fuscous bristles clustered near apex of second segment. Thorax: Pronotum light gray strongly ir- rorated with fuscous tipped scales; tegula mostly brownish fuscous. Venter grayish white to cream. Forewing light gray heavily irorrated with fuscous, most scales with dark fuscous tips; 3 more or less distinct, irreg- ular bands of dark fuscous traversing outer PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON half of wing; subapical band parallel to ter- men and divergent from medial band; small patches of cream scales scattered mostly along costa and termen and extending out into fringe. Hindwing uniformly gray. Fe- male frenulum consisting of two closely set bristles. Fore- and midlegs gray to dark fus- cous dorsally, buff to cream ventrally, gen- erally darker on tibia and tarsus with con- spicuous buff apices to each segment and a median ring on tibia. Hindleg much paler, generally gray with tarsomeres darker and indistinctly ringed with cream. Abdomen. Dark to light gray dorsally, pal- er ventrally. Male genitalia: As shown in Figs. 95-99. Uncus lobes reduced. Subscaphium poorly sclerotized, indistinct. Gnathos slender, forming a deep U. Vinculum abruptly con- stricted to form a moderately elongate, slen- der saccus. Transtilla relatively broad, light- ly sclerotized and highly arched. Valva moderately broad; cucullus rounded; sac- culus with a prominent, elongate spinose lobe arising distally; a single, broad, short, truncate spine arising from apex of lobe. Aedoeagus moderately slender, nearly as long as valva, with a small bulbous lobe at base. Female genitalia: As shown in Figs. 120- 122. Four pairs of apophyses present, in- cluding moderately long anterior and ex- tremely long posterior pairs, a short ventral pair within A10, and a short dorsal pair within A8 which are fused approximately half their length along middle. Ductus bur- sae elongate (about equal to length of pos- terior apophyses), moderately slender, with ductus seminalis joined slightly anterior to middle. Corpus bursae moderately en- larged, with a single, broad, diamond-shaped signum bearing a pair of short, caudally di- rected spines (Fig. 122). Holotype.—Female. Camp VI, 1850 m, Cerro de la Neblina, Territorio Federal Amazonas, Venezuela; 2-4 Dec 1984, R. L. Brown (USNM). VOLUME 92, NUMBER | Paratypes.— VENEZUELA: Same data as holotype; 2 6, 3 9, slides USNM 23672, 29987, 30347, 30420. Paratypes deposited in UCVM and USNM. Host.— Unknown. Flight period.— December. Distribution (Map 1).—Known only from one collecting site on Cerro de la Neblina, Venezuela, which is situated near the Bra- zilian border at 1850 meters and 0°51'N, 6°58'W. Etymology.—The specific epithet is de- rived from the name of the general type locality, Cerro de la Neblina (Mountain of the Mist). Discussion.— Both male and female gen- ital morphology easily distinguishes this species. The spinose lobe of the male valva is the largest of the five currently recognized species of Crepidochares, with an apical spine which is not only the largest, but also the only one that is truncate. Rather intensive collecting during the Cerro de la Neblina expeditions at the Am- azonian basecamp site (130 m), did not re- veal the presence of this species at lower elevations. Collections on the rather large and topographically diverse massif of Neb- lina itself were relatively sparse and un- doubtedly inadequate for Lepidoptera. Only the earliest (December) collections at camp VII (1850 m) resulted in specimens of C. neblinae. All were attracted to ultraviolet lights. Crepidochares colombiae Davis, NEw SPECIES Figs. 2, 100-103, 123-125; Map 1 Adult (Fig. 2).—Length of forewing: 4, 9- 9.5 mm; 2, 11 mm. A small moth with light brown forewings heavily mottled with dark brown striae and bands; ocellus present; la- bial palpus slightly upcurved; male valva with long slender spinose lobe arising from apex of sacculus; female with fourth pair of dorsal apophyses converging caudally but not fused. Head: Vestiture mostly brown, slightly paler and more buff near occiput. Ocellus well developed. Antenna 0.4—0.5 the length of forewing, 40 segmented: scape with 8-10 dark brown piliform scales forming distinct pecten; scales not forming an eyecap; scape and flagellum with dorsal half uniformly covered with dark brown scales, ventral half naked except for dense pubescence of sen- silla. Maxillary palpus elongate, 5-segment- ed; vestiture light brown dorsally, dull white ventrally. Labial palpus uniformly brown laterally, dull white to pale buff mesally; dorsal apex of second segment with a tuft of brown bristles, a few scattered bristles also along dorsal margin; ventral margin rough, with numerous bristles. Thorax: Pronotum uniformly dark brown. Venter white to pale buff. Forewing pale buff, heavily mottled with dark brown; three dark brown bands usually distinct across distal 7; of wing; basal two bands strongly oblique and parallel; distal band parallel to termen; all 3 bands sometimes coalescing to form a ““W” shaped patten; a fine retic- ulate network of dark brown lines and spots scattered between bands and along costal margin; termen mostly dark brown inter- rupted with 3-4 light brown to buff spots. Hindwing uniformly dark gray. Female frenulum consisting of four bristles. Foreleg dark brown dorsally, light brown ventrally with apices of tarsal and tibial segments and middle of tibia ringed with buff. Midleg sim- ilarly marked but generally paler. Hindleg very pale, uniformly pale buff except for slight brownish banding on tarsomeres. Abdomen: Dark brown dorsally, buff ven- trally. Male genitalia: As shown in Figs. 100- 103. Uncus lobes slender, acute, and widely spaced. Subscaphium poorly sclerotized, in- distinct. Gnathos indistinct, membranous. Vinculum constricted to form a short, slen- der saccus. Transtilla a slender arch between bases of valvae. Valva moderately broad, gradully narrowing to apex; an elongate, spi- 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nose process arising from apex of sacculus; apex of process with a slender spine. Ae- doeagus moderately slender, slightly curved, especially at base, and approximately % the length of valva. Female genitalia: As shown in Figs. 123- 125. Four pairs of apophyses present, in- cluding moderately long anterior and ex- tremely long posterior pairs, a short ventral pair within Al0, and a short dorsal pair within A8 which converge at their caudal ends but do not fuse. Ductus bursae mod- erately long, approximately half the length of posterior appophyses, with ductus sem- inalis joined at middle. Corpus bursae great- ly enlarged, with a single, highly irregular, transverse signum (Fig. 125). Holotype.— Male. Sierra del Libano, 6000 ft. [1829 m], Colombia; May 1899, H. H. Smith, 68622, slide 19243 (BMNH). Paratypes.—COLOMBIA: Same locality as holotype, 10 4, 1 2, nos. 6861 3-21, 68623- 24, 68720-21; slides BMNH 19247, USNM 30423. Paratypes deposited in BMNH and USNM. Host.— Unknown. Flight period.— May. Distribution (Map |).—Known only from the type locality, Sierra del Libano, also known as El Libano, which according to Paynter and Traylor (1981) is a dense sub- tropical forest and a spur of the Cuchilla San Lorenzo on the southwestern Sierra Ne- vada de Santa Marta in Magdalena Prov- ince (ca. 11°10’N, 74°W). Etymology.—The specific name is de- rived from the country of origin, Colombia. Discussion.—This species is the largest and darkest in color within the genus. It is also the only species of Crepidochares (of which females are known) in which the dor- sal apophyses (fourth pair) of the eighth ab- dominal segment do not fuse but remain separate, although strongly convergent. Crepidochares colombiae appears most al- lied to C neblinae on the basis of general morphology, particularly the well devel- oped saccular process of the male. Crepidochares austrina Davis, NEw SPECIES Figs. 3, 84, 104-107, 126-128; Map 2 Adult (Fig. 3).— Length of forewing: 3, 4.6— 6 mm; 2, 5.8-6.1 mm. A small moth with buff to light brown forewings variably marked with reddish brown to dark fuscous spots and costal strigulae; male valva with a small spinose process arising from sac- cular lobe; female ovipositor with four pairs of apophyses. Head: Vestiture pale grayish white to buff with a slight concentration of more brown- ish scales across upper frons between an- tennal bases. Antenna 0.6-0.7 the length of forewing, approximately 38 segmented; scape fuscous at base, pale buff apically; scales not forming an eyecap; pecten with 6-8 long light brown piliform scales; flagel- lum with ventral half naked and densely ciliate, dorsal half covered with alternating bands of pale buff and fuscous scales. Max- illary palpus pale buff. Labial palpus pale buff, lightly irrorated with fuscous on sec- ond segment, more heavily so on third; apex and outer side of second segment with 4-6 long, fuscous, bristlelike scales. Thorax: Pronotum light bronzy brown, irrorated with darker and paler brownish scales. Forewing of similar color with a complex pattern of pale buff to reddish brown and dark fuscous scales; distal half of costa with an alternating pattern of about six buff strigulae interspersed by fuscous; oblique fuscous banding slightly evident but obscure due to rubbed condition of speci- mens. Hindwings uniformly shiny gray with an elongate fringe about *4 the width of wing. Venter of thorax uniformly dull white to pale buff. Legs generally dark grayish fus- cous dosally and whitish to buff ventrally, gradually becoming almost entirely whitish to buff on hindleg; tibia and tarsus of fore and midleg with grayish to fuscous banding. Abdomen: Shiny grayish fuscous dorsally, pale buff ventrally. Male genitalia: As shown in Figs. 104— VOLUME 92, NUMBER 1 107. Uncus lobes moderately long and stout. Subscaphium relatively elongate and broad, approximately 0.5 the width of genital cap- sule, joined to arms of gnathos near inser- tion of valvae. Vinculum tapering gradually anteriorly, V-shaped. Valva with ventral margin deeply emarginate near middle, abruptly marking end of sacculus; a small spinose process arising from inner angle of saccular lobe; ventral margin along more narrow, distal half with 5-6 large spinose setae; basal apophysis of valva broadly tri- angular. Aedoeagus without cornuti, ap- proximately *4 the length of valva. Female genitalia: As shown in Figs. 126- 128. Four pairs of apophyses present with only a single, dorsal pair in A8, approxi- mately 65% fused. Ductus bursae short, broad, barely perceptible from moderately enlarged corpus bursae; signum consisting of a single cluster of minute, stout spines (Fig. 128). Holotype.—Female. Anticura, 350 m, Parque Nacional Puyehue, Osorno Prov., Chile; 18 Nov. 1981, Nielsen and Karsholt (ZMHUV). Paratypes.—CHILE: Maule Prov: Paso Garcia, 300 m, ca. 23 km NW Cauquenes: 1 6, 29-30 Nov. 1981, D. R. Davis, slide USNM 29581. Rio Teno, 800 m, ca. 40 km E.. Curico: 1 ¢, 25-27 Nov. 1981, D. R. Davis, slide USNM 29583. Nuble Prov: Alto Tregualemu, 500 m, ca. 20 km SE Chovel- len: 1 6, 1-3 Dec. 1981, D. R. Davis, slide USNM 29580. Osorno Prov: Same data as holotype, | 4, side DRD 3708; 1 6, 17 Dec. 1981, Nielsen and Karsholt. Valdivia Prov: Rincon de la Piedra, 180 m, 20 km S. Val- divia: 1 6, 5 2, 15 Nov. 1981, Nielsen and Karsholt, slides USNM 23599, 29504. Paratypes deposited in ANIC, BMNH, USNM, and ZMHU. Host.— Unknown. Flight period.—November 15 to Decem- ber 17, univoltine. Distribution.—This species occurs fur- ther south than does aridula, and is found within the principal range of Nothofagus. It 11 has been found as far north as the Rio Teno, which approximately marks the northern limits of the Northern Valdivian Forest zone, and extends as far south as Valdivia and Osorno Provinces, which are located well within the more southern Valdivian Forest zone. Etymology.—The specific name is de- rived from the latin austrinus (southern) as suggested by its more southern distribution when compared to the only other Chilean eriocottid, aridula. Discussion.—Crepidochares austrina 1s easily distinguished from the other mem- bers of Crepidochares by major differences in venation and both male and female gen- italia, as summarized in the key to species. The first radial vein in the forewing is char- acteristic in arising more distad from the middle of the discal cell. In contrast to C. aridula, the ovipositor of C. austrina pos- sesses the normal (for Eriocottidae) four pairs of apophyses, including a short, most- ly fused dorsal pair in A8. Present distri- butional data suggest that it may also be more adapted to the wetter Valdivian for- ests of southern Chile. Crepidochares aridula Davis, NEw SPECIES Figs. 4, 5, 18, 19, 34-57, 85-90, 108-112, 129, 130; Map 2 Adult (Figs. 4, 5).—Length of forewing: 6, 5.2-6.1 mm; 2, 4.5 mm. A small moth with light gray forewings variably marked with reddish brown to fuscous spots and oblique bands; male valva relatively simple, with a minute spinose lobe from distal end of sacculus; female ovipositor with five pairs of apophyses. Head: Vestiture pale grayish white to buff with a small lateral tuft of darker scales aris- ing immediately posterior to ocellus and another near front rim of eye. Antenna 0.65 the length of forewing, approximately 31 segmented; scape pale gray irrorated with brown; scales not forming an eyecap; pecten with 6-8 long brownish piliform scales; fla- 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gellum with ventral half naked and densely ciliate, dorsal half covered with alternating bands of grayish white and fuscous scales. Maxillary palpus grayish white to buff. La- bial palpus light brown over basal segments; apical segment mostly covered with diver- gent, pale whitish buff scales; a few bristle- like brown scales arising from apex of sec- ond segment. Thorax: Pronotum pale buff to grayish white, irrorated with darker reddish brown to fuscous scales. Forewing of similar color with a distinct pattern of oblique bands (Figs. 1-2) evident on fresh specimens. Hindwing uniformly pale gray with elongate fringe nearly equalling width of hindwing. Venter of thorax grayish white. Legs mostly dull white ventrally, brown dorsally with two broad fuscous bands usually evident on tib- ia; base of all tarsal segments dark fuscous. Abdomen: Uniformly shiny pale buff to grayish white. Male genitalia: As shown in Figs. 108- 112. Uncus lobes slender, short. Subsca- phium moderately slender and short, joined to arms of gnathos well above insertion of valvae. Vinculum narrowing abruptly an- teriorly to form a moderately broad saccus. Valva relatively simple, ventral margin with saccular lobe slightly developed and bearing a single minute spinose lobe; basal saccular area approximately twice the width of distal half} basal apophysis of valva elongate and slender. Aedoeagus simple, without cornuti, approximately equalling valva in length. Female genitalia: As shown in Figs. 129, 130. Five pairs of apophyses present, in- cluding elongate anterior and posterior apophyses, a short ventral pair within A10, and two pairs of dorsal apophyses within A8, one pair of which are fused about 70% their length. Ductus bursae extremely slen- der. Corpus bursae moderately enlarged, without signum. Holotype.— Male. Rio Colorado, ca. 40 km SE Santiago, 1100 m, Metropolitan Re- gion, Chile; 29-31 Oct. 1981, Don and Mi- gnon Davis, USNM. Paratypes.— CHILE: Same data as holo- type; 7 6, slides USNM 29503, 29587, 22140. Curico Prov. Potrero Grande, 35 km SE Curico, 35°12.5'S, 71°W: 1 4, 6 Dec. 1982, R. L. Brown, USNM slide 29584. 10 km NW Rauco, 34°52’S, 71°21'W: 1 9. 2 Dec. 1982, USNM slide 29582. Paratypes de- posited in ANIC, MHNS, and USNM. Host.-Unknown. Flight period.—October 29 to December 6; univoltine. Distribution.—Known only from the Central Valley Biotic Province and lower elevations of the adjacent Central Andean Cordillera Province of central Chile. The forests of these areas are relatively dry and are situated just north of the principal northern limits of Nothofagus. Etymology.—The specific name is de- rived from the Latin aridulus (diminutive of dry) in reference to the more xeric habitat of this species. Discussion. —Crepidochares aridula dif- fers considerably from the only other Chil- ean eriocottid, C. austrina, by major differ- ences in venation and the genitalia of both sexes, aS Summarized in the key to species. The forewing venation of C. aridula agrees more with the other species of Crepido- chares and Eriocottis in possessing a more basal origin for R1. Of the five recognized species of New World Eriocottidae, the male valva of C. aridula has the most reduced saccular spine. The female ovipositor is unique among Lepidoptera by the presence of five pairs of apophyses. The extra fifth set is believed to be derived from the caudal ends of the anterior apophyses. In addition, C. aridula exhibits a generally paler color than C. austrina. Its range may also be con- fined to the drier, more northern non-Noth- ofagus forests of central Chile. All speci- mens were collected in UV light traps. Because the sole female specimen of this species has not been collected in close as- sociation with any males and is slightly smaller in size with more distinctly marked forewings, its identity remains somewhat in VOLUME 92, NUMBER | doubt. Association of males and females needs confirmation in particular because the most unusual apomorphy (i.e. the fifth pair of apophyses) of the species occurs only in the female. Crepidochares subtigrina Meyrick Figs. 6, 58-63, 83, 113, 114; Map 1 Crepidochares subtigrina Meyrick, 1922: 601.—Fletcher, 1929: 58.—Clarke, 1955: 298; 1970: 36.—Davis, 1984: 21. Adult (Fig. 6).—Length of forewing: 4, 7.8 mm. A small moth with pale yellowish fore- wings irregularly marked with slightly dark- er, more brownish transverse bands and spots; ocellus absent; labial palpus porrect; male valva with a stout spinose lobe arising from apex of sacculus; female unknown. Head: Vestiture pale cream to white. Ocellus absent. Antenna 0.7-0.8 the length of forewing, approximately 40 segmented; scape pale cream, with approximately 12 piliform cream colored scales forming a pecten; scales dorsad to pecten extending forward to form a broad eyecap; flagellum dorsally covered with pale cream scales; ventrally the flagellum is strongly ciliate with most sensilla clustered on two pairs of short pedicels (Figs. 58-60), with apical pair aris- ing more approximate. Maxillary palpus greatly reduced, 3-segmented with apical segment slightly larger, covered with a few pale cream scales. Labial palpus pale buff to cream, irrorated with slightly darker tipped scales; vestiture broad near base, gradually tapering to apex; strongly porrect, completely smooth and without bristles. Thorax: Pronotum cream, slightly darker and more brown over tegula. Venter cream. Forewing cream to pale yellow crossed by several, slightly darker yellow brown bands or strigulae, basal three slightly darker and 13 most distinct; subapical two bands very in- distinct and interrupted; apical band very pale but entire; fringe cream. Hindwing un1- formly pale cream. Foreleg cream with darker brown suffusion dorsally over tibia and tarsus; hindleg uniformly pale cream. Abdomen: Color not examined (on slide). Male genitalia: As shown in Figs. 113, 114. Uncus lobes moderately long and stout. Subscaphium poorly sclerotized, indistinct. Gnathos slender, forming a deep V. Vin- culum tapering anteriorly, with a slight con- striction near base. Transtilla slender. Valva slender, with 3-4 stout, spinose setae arising near middle immediately distad to spinose lobe from sacculus. Aedoeagus moderately stout, approximately 0.6 the length of gen- ital capsule, without cornuti. Type.— Holotype. ¢; BMNH. Distribution (Map 1).—Known only from the holotype, which was collected in Oc- tober by Parish along the Amazon River at Parintins, Amazonas, Brazil. Discussion.—Since its discovery, Crepi- dochares subtigrina has been consistently regarded as a member of the Tineidae. As stated in the key to species and elsewhere, numerous apomorphies distinguish this species, still represented by only the male holotype. The male genitalia, especially the prominent spinose process arising from the sacculus, clearly associates C. subtigrina with the other members of the genus. Although Meyrick describes the ocelli as “posterior,” no ocelli were observed on the holotype. In addition to its rather aberrant mor- phology, C. subtigrina is of further interest in being the only South American eriocottid discovered thus far from the lowland trop- ics. The other four species described herein are either from montane or southern tem- perate habitats. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ Figs. 1-6. Adult Eriocottidae. 1, Crepidochares neblinae n. sp., female holotype, Venezuela (8.5 mm). 2, Crepidochares colombiae n. sp., male holotype, Colombia (9 mm). 3, Crepidochares austrina n. sp., male holotype, Chile (6 mm). 4, Crepidochares aridula n. sp., male holotype, Chile (5.8 mm). 5, Crepidochares aridula n. sp., female paratype, Chile (4.7 mm). 6, Crepidochares subtigrina Meyrick, male holotype, Brazil (7.8 mm). (Length of forewing in parentheses.) VOLUME 92, NUMBER | 15 I ; 12 Figs. 7-12. Adult Eriocottidae. 7, Eriocottis fuscanella Zeller, male [Europe] (8 mm). 8, Eriocottis flaviceph- alana Issiki, male holotype, Taiwan (8.5 mm). 9, Deuterotinea casanella (Eversmann), male [Europe] (10.2 mm). 10, Compsoctena thwaitesii (Walsingham), male, Sri Lanka (15.5 mm). 11, Compsoctena aethalea (Meyrick), male, India (14.8 mm). 12, Compsoctena aethalea (Meyrick), female, India (18 mm). (Length of forewing in parentheses.) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Figs. 13-15. Lateral view of heads. 13, Crepido- chares colombiae n. sp. 14, Crepidochares subtigrina Meyrick, right side, antenna removed. 15, Left side of Fig. 14, note antennal “‘eyecap.” VOLUME 92, NUMBER 1 Figs. 16-21. Head structure. 16, Crepidochares neblinae n. sp., anterior view (0.5 mm). 17, Maxilla of Fig. 16 (0.2 mm). 18, Crepidochares aridula n. sp., anterior view (0.5 mm). 19, Maxilla of Fig. 18 (0.2 mm). 20, Compsoctena thwaitesii (Walsingham), anterior view (0.5 mm). 21, Maxilla of Fig. 20 (0.2 mm). (Scale lengths in parentheses.) 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON > o aS = > Stiropius Polystenidea Figs. 1-3. 1, Cladogram of Stiropius group of genera; 2, face of C. birostratus, 2, anterior view; 3, fa Polystenidea sp., anterior view. 2r; hind femora and apical tarsomeres en- larged, swollen; second metasomal tergite at least 1.5 as broad posteriorly as anteriorly; hypopygium triangular in profile, protruding P Viridipyge Whitfield 2Rs of fore wing subequal in length with 2r, sometimes longer: hind femora and apical tar- someres not enlarged; second metasomal ter- gite variable but seldom as much as 1.5 as broad posteriorly as anteriorly; hypopygium truncate, short . Stiropius Cameron Malar suture present and distinct between com- pound eye and mandibular base (Fig. 2); malar space relatively short, less than half eye height; pterostigma of fore wing almost always elon- gate distally, so that r arises from stigma well before midlength (Figs. 4, 10); hind femora often face of swollen or otherwise modified (Fig. 5); meta- somal tergite IV without semicircular subbasal groove or depression Choreborogas, new genus Malar suture absent, at least not apparent; ma- lar space long, nearly equal to eye height (Fig. 3); r arising from stigma at about midlength; hind femora not modified or swollen, meta- somal tergite IV with semicircular subbasal groove or depression Polystenidea Viereck Choreborogas, gen. n. Type-species: Choreborogas birostratus, n. sp., described below. Diagnosis.—This genus shares with the other three related genera the following 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON combination of features: antennae 13-14 segmented; reduced notauli, especially pos- teriorly; areolate propodeum; metasomal pseudo-carapace incorporating T1-T3 and, toa lesser extent, T4; ovipositor and sheaths short, partially exserted; gonobase of male subtriangular, nearly as long as broad. There are no really substantiated host records; I expect the hosts, as in the other three genera, are lyonetiid or related leafmining Lepi- doptera, and that the wasps endoparasiti- cally mummify the host larvae or prepupae. Choreborogas differs from Polystenidea in lacking the semicircular, subbasal groove on the fourth metasomal tergite, in having an apically elongate pterostigma (as in Chorebus, hence the generic name), 1n hav- ing distinct malar or subocular sutures and in often having swollen hind femora and apical tarsomeres. It is also distinguished from Stiropius by all of the above differ- ences except the subocular sutures, as well as by lacking vein 2r-m of the fore wing. Finally, it differs from Viridipyge in pos- sessing sublateral, pigmented, more strong- ly sclerotized plates on metasomal sterna 2— 5, in having the apically elongate pterostig- ma, and in lacking the vein 2r-m of the forewing. Stiropius and Viridipyge will both key to Bucculatriplex in Marsh et al. 1987; Polyste- nidea and (sometimes with great difficulty) Choreborogas will key to Polystenidea. Comments.—As far as is known, the species of Choreborogas are all predomi- nantly Neotropical, with several of the species ranging into the southeastern U.S. and probably into parts of Arizona as well. They are quite abundant in some light trap samples from Central America, indicating that they are probably nocturnally active and often numerous. I have seen an esti- mated 20 species in collections. Two new species, the first being the type, are de- scribed below to illustrate the range of fea- tures found in this remarkable, yet essen- tially unknown, group of wasps. Choreborogas birostratus, sp. n. Figs. 2:45-5:56, 7 Female.— Body length: 1.4-2.1 mm, fore wing length 1.6-—2.4 mm. Head: Color orange-brown, often with darker brown regions around ocelli, facial carinae and occipital carina. Supraoral depression enlarged to include lower 0.3— 0.5 of frons; clypeus embedded within this depression and flattened; depression marked dorsally and laterally by carina, which is produced submedially into two flattened pointed noselike projections. Mandibles en- larged and strongly overlapping when closed. Inner margins of eyes parallel to diverging ventrally. Malar space less than 0.3 eye height, with strong malar suture. Antennae 14-segmented, slightly shorter than body, with apical flagellomeres 3-4 x as long as broad. Maxillary and labial palpi pale yel- lowish, slender. Hypostomal and occipital carinae remaining separate to mandibular bases. Ocelli roughly equidistant from each other. Mesosoma: Entirely pale orange-brown (occasionally darker brown in generally darker specimens). Pronotum with shallow, narrow dorsal and ventral grooves, other- wise nearly smooth. Mesoscutum very fine- ly granular, matte, with weak indentations indicating courses of notauli. Scutoscutellar scrobe composed of two transverse exca- vations, narrower medially and traversed medially by a thin ridge. Scutellar disc finely punctate, flat. Mesopleuron weakly granu- lar, with shallow, sinuate logitudinal groove. Propodeum with narrow but distinct meta- postnotal-propodeal groove, medial longi- tudinal carina over anterior 0.4-0.5, and posterior medial gothic-arch-shaped areola; costulae obsolescent or entirely absent. Wings: C+Sc+R and stigma pale yellow- brown; stigma occasionally whitish in some specimens. Remainder of prominent ve- nation evenly brownish. Fore wing (Fig. 4) with r originating in basal 0.3-0.4 of stigma, VOLUME 92, NUMBER 1 2r-m absent. Stigma 4 as long as broad. 3Rs meeting wing edge about 0.6 of distance between distal end of stigma and wing tip, just short of end of R1 (metacarp). Legs: All legs pale yellow-brown with ul- timate tarsomeres dark brown apically. Hind femora (Fig. 5) swollen, each with subbasal ventral toothlike projection. Hind tibiae armed with considerably longer setae than remainder of legs. Front and hind apical tarsomeres enlarged, about 2 = as broad as preceding tarsomeres. Metasoma (Fig. 6): T1, T2, anterior edge of T3 light orange-brown and mostly with granular sculpturing. T3 and T4 darker brown. Tergite 1 weakly broadening pos- teriorly, about 1.3 as long as posteriorly broad, with semicircular basal carina and longitudinal medial carina. T2 1.4 as broad posteriorly as long, 1.6 as broad posteriorly as anteriorly, separated from T3 by crenulate furrow. T2 1.5 as broad as T1 at junction with T1. T3 2 as broad as long, weakly rounded posteriorly and strongly overlapping T4. T4 much narrower than T3, anteriorly transversely striate, pos- teriorly granular. Hypopygium short, trun- cate, not projecting. Ovipositor and sheaths short, subexserted. Males. —Essentially same size range as fe- males but with some pronounced morpho- logical differences: lower portion of frons not incorporated into large supraoral depression (thus facial carinae and projec- tions are absent); mandibles not enlarged apically; stigma broader (3 as long as broad); hind femora less strongly swollen and without subbasal, ventral toothlike pro- jection. Material examined.— Holotype 2: MEX- ICO: Guerrero, 17 mi. E. Tixtla, 1 1.vii.1985 (Woolley, Zolnerowich) (TAMU, deposited in USNM). Paratypes: MEXICO: Colima: 1 4, Parque Nac. de Volcan Colima, 8.2 mi. from Hwy. 54, 12.vii.1984 (Woolley) (TAMU). Guerrero: 3 2, same data as ho- lotype; 1 9, 3 6, 6 mi. E. Xochipala, 41 Figs. 4-10. 4, Fore wing of C. birostratus, sp.n., 2; 5, hind leg of C. birostratus, sp. n., 2; 6, metasomal tergites of C. birostratus, sp. n., 2; 7, propodeum of C. birostratus, sp. n., 2; 8, propodeum of C. andeanus, sp. n., 2; 9, metasomal tergites of C. andeanus, sp. n., 2; 10, fore wing of C. andeanus, sp. n., 2. 13.vii.1985 (Woolley, Zolnerowich) (TAMU); 4 9, 3 6, 6.2 mi. SW Xochipala, 13.vii.1985 (Woolley, Zolnerowich) (TAMU); 1 4, 2 mi. E. Ocotito, 11.vii.1985 (Woolley, Zolnerowich) (TAMU); 1 @, 32 mi. SE Petatlan, 10.vii.1985 (Woolley, Zolnerowich) (TAMU). Michoacan: | 2°, 49 mi. SE Aguila, 13.vii.1984 (Woolley, Zolne- rowich) (TAMU); 1 9, 28.5 mi. S. Nueva Italia, 9.vii.1985 (Woolley, Zolnerowich) (TAMU); 1 2, 10 mi. S. Uruapan, 7.v1i1.1985 (Woolley, Zolnerowich) (TAMU). Oaxaca: 1 9°, 4.4 mi. NE San Pedro Mixtepec, 16.vii.1985 (Woolley, Zolnerowich) (TAMU). Tamaulipas: 2 2, 5 46, 5 mi. W. Gomez Farias, 20.iii1.1986 (Wharton) (RAW, deposited in USNM). Comments.—This species 1s as remark- able for its pronounced sexual dimorphism 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as for its unusual facial features. At least one other undescribed species from Central America shares a similar facial modification (along with even more extreme modifica- tions of the hind legs), but many species are similar to C. birostratus males in both sexes. This species can be easily separated from the following one by the shape of the meta- somal tergites (Fig. 6), the small size (usually less than 2 mm), the distally less elongate fore wings, as well as by the striking mod- ifications of the face and hind femora in the females. Choreborogas andeanus, sp. n. Figs. 8, 9, 10 Females. — Body length: 2.0-2.4 mm; fore wing length 2.7-3.1 mm. Head: Entirely deep brown except light yellow-brown labrum, mandibles and palpi. Supaoral depression round, small; clypeus apically concave. Frons finely granular; in- ner margins of eyes parallel. Antennae 14- segmented, apical 5 flagellomeres more te- rete. Malar spaces about 0.3 eye height, with strong malar suture. Ocelli roughly equidis- tant from each other. Mesosma: Mostly weakly punctate/gran- ular dorsally, shinier laterally, entirely deep brown. Pronotum smooth, polished, with narrow, shallow dorsal and ventral grooves. Mesoscutum with no sign of notauli, evenly and finely punctate. Scutoscutellar scrobe composed of two broad rectangular exca- vations separated by a narrow ridge. Scu- tellar disc sculptured as mesoscutum, weak- ly convex, subtriangular. Mesopleuron highly polished, with very faint to absent longitudinal depression. Propodeum (Fig. 8) with narrow but distinct metapostnotal- propodeal groove, medial longitudinal ca- rina over anterior 0.5—-0.6, and horseshoe- shaped to narrower pointed medial areola posteriorly. Costulae absent. Wings: Wings disproportionately large relative to body size. Venation entirely pale brownish. Fore wing (Fig. 10) with apically extremely elongate stigma (stigma 7x as long as broad), r arising in its proximal 0.2. 3Rs strongly curved, reaching wing margin approximately halfway between distal end of stigma and wing tip. Legs: All legs pale yellow-brown with darker, strongly swollen ultimate tarso- meres. Hind femora swollen but otherwise unmodified, with sparse setae of average length. Metasoma (Fig. 9): Tergite 1 finely punc- tate, 1.3 as long as posteriorly broad, weakly broadening to spiracle then parallel- sided, with semicircular carina anteriorly and obsolescent medial longitudinal carina. Tergite 2 finely granular, 1.5 = as broad at T1/T2 junction as T1, slightly longer than apically broad, separated from T3 by shal- low, crenulate groove. T3 smooth, polished, 1.9 as broad as long, with strongly round- ed posterior margin. T4 and succeeding ter- gites smooth, more flexible, telescoped un- der T3. Hypopygium short, truncate, not projecting ventrally. Ovipositor and sheaths short, subexserted. Males. —Essentially similar to females, but with slightly more parallel-sided T2 and rel- atively less transverse T3, which tends to have more (albeit weak) sculpturing as op- posed to the smoother T3 in the female. Material examined.— Holotype 9°: CO- LOMBIA: Putumayo, 2900 m, 1°10’N, 77°15'W, 2.x1ii.1972 (Helava) (CNC). Para- types: 1 2°, 6 4, same data as holotype; CO- LOMBIA: Antioquia, | 2, 1800 m, 7°5'N, 76°30'W, (no date) (Helava) (CNC); Quin- dio, 1 2, 11 km E. Calarca, 7000’, 5.111.1974 (Peck & Peck) (CNC). PERU: Amazonas, 1 4, 2800 m, 6°48’S, 77°38'W, 13.11.1973 (He- lava) (CNC). Comments. —At first glance, the peculiar long-winged and polished appearance of this species makes it seem entirely unrelated to C. birostratus. In many structural features, however, the two species are quite similar, one or the other being more extreme in some respect. The peculiar appearance of both species is produced by exaggeration of ten- dencies found in most species of the genus VOLUME 92, NUMBER 1 to some degree. It would be quite interesting to know what advantage, if any, the long wings of C. andeanus confer. ACKNOWLEDGMENTS I would especially like to thank Michael J. Sharkey (Ottawa) and Robert A. Wharton (College Station) for repeatedly obtaining and sorting out numerous specimens be- longing to the Stiropius group, especially from unsorted Neotropical collections. In addition, Kees van Achterberg (Leiden), Mark R. Shaw (Edinburgh) and Roy A. Shenefelt provided helpful and encouraging discussions on the systematics and biology of Rogadinae. Robert A. Wharton and Mi1- chael S. Arduser offered useful comments on an earlier draft of this paper. The early period of study of this group was supported by a fellowship awarded in 1985 by the North Atlantic Treaty Organization to the author. LITERATURE CITED van Achterberg, C. 1983. Revisionary notes on the Palaearctic genera and species of the tribe Exo- thecini Foerster (Hymenoptera, Braconidae). Zool. Meded. 57: 339-355. Belokobyl’skii, S. A. 1984. [On the division of the tribe Exothecini s.l. (Hymenoptera: Braconidae) in two with the description of a new genus and subgenus.] Zool. Zhurn. 63: 1019-1026. [In Rus- sian, English summary.] Maddison, W. P. and D. R. Maddison. 1987. MacClade, Version 2.1. A phylogenetics computer program distributed by the authors. 43 Marsh, P.M., S. R. Shaw, and R. A. Wharton. 1987. An identification manual for the North American genera of the family Braconidae (Hymenoptera). Mem. Entomol. Soc. Wash. 13. 98 pp. Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): A phylogeny and reclassification of Microgastrinae. Mem. Entomol. Soc. Canada 115. 147 pp. Papp, J. 1975. Three new European species of Co- lastes Hal. with taxonomic remarks. Acta Zool. Acad. Sci. Hung. 21: 411-423. Shaw, M.R. 1983. One evolution of endoparasitism: The biology of some genera of Rogadinae (Bra- conidae). Contrib. Amer. Entomol. Inst. 20: 307- 328. Swofford, D. L. 1980. PAUP— Phylogenetic Analysis Using Parsimony. Version 2.4.1. Computer pro- gram available from the author, Illinois Natural History Survey. Viereck, H. L. 1912. Descriptions of one new family, eight new genera, and thirty-three new species of ichneumon-files. Proc. U.S. Nat. Mus. 43: 575- 593. Whitfield, J. B. 1988a. Revision of the Nearctic species of the genus Stiropius Cameron (= Bucculatriplex auct.) with the description of a new related genus (Hymenoptera: Braconidae). Syst. Entomol. 13: 373-385. 1988b. Taxonomic notes on Rhyssalini and Rhysipolini (Hymenoptera: Braconidae) with first Nearctic records of three genera. Proc. Entomol. Soc. Wash. 90: 471-473. Whitfield, J. B. and van Achterberg, C. 1987. Clar- ification of the taxonomic status of the genera Can- tharoctonus Viereck, Noserus Foerster and Pseu- davga Tobias (Hymenoptera: Braconidae). Syst. Entomol. 12: 509-518. Whitfield, J. B., Johnson, N. F., and Hamerski, M. R. 1989. Identity and phylogenetic significance of the metapostnotum in the non-aculeate Hyme- noptera. Ann. Entomol. Soc. Amer. 82. (In press.) PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 44-57 DESCRIPTIONS, DISTRIBUTION, AND HOST-PLANT RECORDS OF EIGHT FIRST INSTARS IN THE GENUS TOUMEYELLA (HOMOPTERA: COCCIDAE) B. J. SHEFFER AND M. L. WILLIAMS Department of Entomology, Alabama Agricultural Experiment Station, Auburn Uni- versity, Alabama 36849-5413; (current address, BJS) Department of Pediatrics, SUNY- Health Science Center, 750 Adams Street, Syracuse, New York 13210. Abstract. —Detailed morphological descriptions of first-instar nymphs of 8 species in the genus Joumeyella are presented. Included are illustrations, a key to described species, and discussions of general morphology and species relationships. The first instar of a ninth species 7. sonorensis (Cockerell and Parrott), historically included in the genus, proved not to be congeneric with the type or other species within the genus. Host plants and distributions are also given. Key Words: The soft scale insect genus Toumeyella was first proposed as a subgenus of Lecani- um Burmeister by Cockerell in 1895 with Lecanium mirabile as its type species. Cock- erell (1902) later elevated it to generic rank. As currently recognized, the genus Tou- meyella is composed of 11 described species, 8 of which occur in North America. Identification of species of Toumeyella has been a problem because of a lack of ade- quate descriptions and keys, as well as ap- parent host induced and geographical vari- ation in some species. Williams and Kosztarab (1972) and Hamon and Williams (1984) provided descriptions, illustrations, hosts, distributions and biological notes on 5 species occurring in Virginia and Florida, respectively. Included were Toumeyella cerifera Ferris, T. liriodendri (Gmelin), T. parvicornis (Cockerell), 7. pini (King), and T. virginiana Williams and Kosztarab. Previous taxonomic studies of Toumey- ella species have focused primarily on adult females, with little attention to other de- velopmental stages. Ferris (1919), in a re- Homoptera, Coccoidea, Coccidae, Toumeyella, first instars description of 7. mirabilis (Cockerell), il- lustrated the first instar in some detail and commented on the spiracular setae, mar- ginal setae, and anal plate reticulation. Hei- del and Kohler (1979) presented a brief de- scription and illustration of the first instar of 7. cubensis Heidel and Kohler. These papers represent the only attempts of de- scribing and illustrating immature stages of Toumeyella. Studies of immature stages of Coccidae are needed to develop complete and sound classification, yet relatively few such studies have been conducted (Howell and Tippins 1973). Classification based solely on adult female characters may produce erroneous phylogenies. For example, the genera Chionaspis and Pseudaulacaspis were con- sidered closely related until examination of second instar males revealed characters that indicated the genera arose from separate phylogenetic stocks (Takagi and Kawai 1967). Additionally, extreme host-induced di- morphism can lead to erroneous classifi- VOLUME 92, NUMBER 1 cation. For example, individual Chionaspis nyssae Comstock feeding on two different hosts exhibited such diverse morphological characters that they were placed 1n separate genera (Knipscher et al. 1976). Mobile, non- feeding first instars are not so influenced by their host as are later developmental stages (Howell 1981). Sibling species and species in complexes may so closely resemble each other as adult females that utilization of im- mature stages is the only means of separa- tion (Howell 1981). More accurate phylog- enies may be produced with the aid of first instar nymphs which exhibit characters often lost or reduced in adult females. Presented in this paper are detailed de- scriptions, illustrations and an identifica- tion key to first-instar nymphs of 8 species of Toumeyella. Distribution and host plant information are given. A discussion of first- instar general morphology, as well as species relationships, is included. The species of Toumeyella treated herein occur in North America except 7. nectandrae Hempel, which is found in Brazil. Three species could not be studied for lack of specimens: 7. cu- bensis Heidel and Kohler, a Cuban species; T. paulista Hempel, a Brazilian species; and T. pinicola Ferris, from the Western United States. MATERIALS AND METHODS Specimens (slide mounted and/or dry) were borrowed from the following institu- tions: Auburn University (AUEM), Florida State Collection of Arthropods (FSCA), University of California Davis (UCDC), United States National Museum of Natural History (USNM), and Virginia Polytechnic Institute and State University (VPIC). A minimum of 10 slide-mounted first- instar nymphs were measured for each species description. Measurements were made utilizing a phase-contrast microscope fitted with an ocular micrometer. For each structure measured, the mean and range (parenthetic) are given in microns in each species description. Terminology used in 45 descriptions is from Williams and Koszta- rab (1972). Drawings are not made to the same scale in all species nor are dermal structures and enlargements in direct pro- portion to each other. The scale bar shown in each figure refers only to body size and not to enlargements. In the Specimens Studied section, the first number indicates the number of slides and the second number (parenthetic) the num- ber of specimens, if different. Collection ab- breviations are utilized to indicate speci- men deposition in the Specimens Studied section. GENERAL MORPHOLOGY OF FIRST INSTAR NYMPHS Fig. 1 Body (Fig. 1-A).—Slde-mounted speci- mens generally oval to elongate-oval, 597 (330-965) long and 354 (215-681) wide. Derm membranous throughout. Append- ages, mouthparts, pores and microducts sclerotized. Segmentation.—Head, thorax and abdominal segments closely fused. Seg- mentation not readily apparent. Antennae (Fig. 1-B).—Antennae well developed, 5-segmented, 3rd and 5th segments gener- ally about equal length. Slender hairlike se- tae on all segments, with enlarged “stout” sensory setae (Fig. 1-C) on segments 4 and 5. A simple sensory pore (Fig. 1-D) on seg- ment 2. Eyes (Fig. 1-E).—Located on mar- gin just above antennal scape, reduced to a single facet. Mouthparts (Fig. 1-F).— Mouthparts lie between the procoxae, con- sisting of clypeolabral shield, one-segment- ed labium with 6-8 setae, and stylet loop. Legs (Fig. 1-G).—Well developed, without tibiotarsal sclerotization or free articula- tion. Two sensory pores (Fig. |-H) on each side of trochanter. Various hairlike setae on each segment, and knobbed digitules in pairs on tarsi (Fig. 1-I) and claws (Fig. 1-J) except prothoracic legs with | tarsal digitule seti- form. Microctenidia (Fig. 1-K) at tibial apex present or absent. Tarsal claw (Fig. 1-L) simple or with a denticle. Spiracles (Fig. 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Q 1 \} C | w NK | E >h ~ A \ a = | Avy 1 A a os A =" / \ ~ a ne a nN .\ fox —~4 i YJ? s \ Gn — K | Vest a i |" 4 ieee ] ~P (oy | | ic @.44 . fh (\) lsat oN R . T b A Ss NG ; y | \ ‘ yy res / | ele) ba) ww oa K \ | () ii) \y Vs ey) s wr Paar | 100 Hm Fig. 1. star. General morphology, 7oumeyella first in- 1-M).—Two thoracic pairs, associated with ventral quadralocular pores (Fig. 1-N) in spiracular furrows; 2-3 pores anteriorly, 3— 4 posteriorly. Occasional quinquelocular or 6-locular pore. Spiracular setae (Fig. 1-O).— Three stout spiracular setae in each spirac- ular furrow in all species except T. parvi- cornis (Fig. 6). Spiracular setae are taxo- nomically important in separating species. Anal plates (Fig. 1-Pa—c).— Two anal plates, well developed, triangular with rounded an- gles. Dorsum with surface microspines (Fig. 1-Pa), coarse (Fig. 1-Pb) or dense (Fig. 1-Pc) reticulation. Four dorsal setae per plate; | on mesal margin, 3 apical. Thick median apical seta approximately ' body length. One ventral subapical seta per plate and | pair fringe setae on anal fold. Dorsal surface texture of anal plates important in separat- ing species. Anal ring (Fig. 1-Q).—Subcir- cular to roundly hexagonal with 6 stout anal ring hairs and a row of irregularly shaped pores. Pores.—Small (2 wm dia.), usually simple, dorsal disc pores (Fig. 1-R) in sub- median and submarginal longitudinal rows. Dorsal bilocular pores (Fig. 1-S) in sub- median and often submarginal longitudinal rows. Bilocular pores categorized as small (longest length 2 um) to large (longest length 4-8 um). Dorsal trilocular pore (Fig. 1-T) on derm, anterior to each antennal scape. Ducts.— Ventral microducts (Fig. 1-U) in 2 submarginal longitudinal rows of 5 each on the abdomen, | between anterior and pos- terior spiracular furrows and | posterior to each eye. Body setae.— Marginal setae (Fig. 1-V) slender to stout, distribution: 8 ante- riorly between eyes, 2-3 on each side be- tween eye and anterior spiracular setae, 2-3 on each side between anterior and posterior spiracular setae, 16 posteriorly on abdo- men. Additionally, 7. parvicornis has 2 se- tae on the body margin at the apex of each spiracular furrow, which are undifferentiat- ed from marginal setae. Ventral body setae bristlelike, of 2 lengths: submarginal setae (Fig. 1-W) short, in 2 longitudinal rows of 7 each on abdomen, | seta between anterior and posterior spiracular furrows and | pair at head apex; body setae (Fig. 1-X) long, in 2 submedian rows of 3 each on posterior abdominal segments; with 2 interantennal setae. Other structures.— Ventral micros- pines (Fig. 1-Y) on posterior abdominal seg- ments. Key TO 8 First-INSTAR NYMPHS OF THE GENUS TOUMEYELLA 1 Dorsal bilocular pore clusters present (Fig. 6-C); 44 marginal setae around body; spi- racular setae undifferentiated from margin- al setae (Fig. 6-B) ...... I. parvicornis 1g Dorsal bilocular pore clusters absent; 32-36 marginal setae around body; spiracular setae distinctly different from marginal setae, 3 in each spiracular furrow (Fig. 1-O) .......... 2 2.(1') Body with 36 slender marginal setae (Fig. FE) Bl cet a ae mR RR Se a T. cerifera VOLUME 92, NUMBER 1 2’. Body with 32 slender to stout marginal setae 3.(2') Dorsal bilocular pores (2 um) present in 2 submedian longitudinal rows only (Fig. 4-D); median spiracular setae 56 times longer than anterior set of lateral spiracular setae (Fig. 4-C); marginal setae stout T. mirabilis Bf Dorsal bilocular pores (2-8 um) present in submarginal and submedian longitudinal rows (Fig. 1-S); median spiracular setae ap- proximately 2 times longer than lateral spi- LAGU ATSCLAC ME ere ieee eee te 4 4.(3') Small dorsal bilocular pores Q An) present in submarginal and submedian longitudinal rows; tibial microctenidia present (Fig. 1-K) EL ee Ra rn eer er Fart 5 4. Larger dorsal bilocular pores (>3 um) pres- ent in submarginal and submedian longi- tudinal rows, predominantly bow shaped (Fig. 8-D); tibial microctenidia absent Pee eat ratios T. quadrifasciata Dorsal bilocular pores (2 um) present in submarginal, submedian, and intermediate longitudinal rows (Fig. 9-D) T. virginiana Di Dorsal bilocular pores (2 um) present in submarginal and submedian longitudinal rows only 6.(5') Dorsum of anal plates densely rencolated: (Fig. 1-Pc); marginal setae stout; not on Pi- nus sp. .. 7 (op Dorsum of anal plates not reticulated, but with sparsely distributed microspines (Fig. 7-H); marginal setae slender (Fig. 7-B); on Pinus sp. . ‘ . T. pini Dorsal quinquelocular disc pores (2 um) usually present in submarginal and sub- median longitudinal rows (Fig. 3-E); North American T. liriodendri qe Dorsal Giinguelocaler disc pores absent; dorsal disc pores simple (Fig. 5-E); Brazilian . T. nectandrae 5.(4) 7.(6) Toumeyella cerifera Ferris Fig. 2 Toumeyella cerifera Ferris 1921: 90. Stein- weden 1929: 227, Williams and Koszta- rab 1972: 160, Hamon and Williams 1984: 117. Specimens studied. — Cephalanthus occi- dentalis: 3(13), Sussex Co., Airport Pond, Wakefield, VA, 16 Aug 1969, Michael L. Williams (AUEM); 1, Macon Co., AL, 31 May 1975, MLW (AUEM); 2(8), Macon Co., AL, 21 Jun 1975, MLW (AUEM); 2(21), —<-- kK B q OS) 4 7 Wy 2 es [yy al c al ie Bike ee )\ A ( pe ( fo Ape / i < Wo. 6) ; t saa = (2) WwW (@ ey s @ nn ° RS ry \ \ TCT =e Fig. 2. First instar, Towmeyella cerifera Ferris 1921. Macon Co., AL, 13 Jul 1975, MLW (AUEM); 1(3), Macon Co., AL, 24 Aug 1975, MLW (AUEM); 1(2), Macon Co., AL, 13 Oct 1975, MLW (AUEM). Additional host plants and distribu- tion.—Toumeyella cerifera was first de- scribed from A/bizzia occidentalis collected in Baja California, Mexico. Reported to oc- cur in AR, FL, LA, and NC. General appearance.—Body (Fig. 2-A) oval, 642 (514-939) long, 391 (296-610) wide. Dorsum.— Marginal setae (Fig. 2-B) 14 (10-19) long, slender, curved posteriorly, distribution: 8 anteriorly between eyes, 3 on each side between eyes and anterior spirac- ular setae, 3 on each side between anterior and posterior spiracular setae, 16 on pos- terior of body. Three spiracular setae (Fig. 2-C) in each spiracular furrow; median setae 24 (19-28) long, lateral setae 14 (1 1-18) long. Small (2 um) bilocular (Fig. 2-D) and simple disc (Fig. 2-E) pores in submedian and sub- 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON marginal longitudinal rows. Anal plates (Fig. 2-H).—Each plate with dorsum densely re- ticulated, 68 (59-78) long, 34 (28-42) wide: cephalolateral margin 44 (40-48) long, cau- dolateral margin 41 (28-47) long. Venter. — Antennae (Fig. 2-L) 170 (157-179) long. Legs (Fig. 2-M) 250 (234-268) long, mi- croctenidia (Fig. 2-N) at tibial apex; tarsal digitules (Fig. 2-O) 49 (42-56) and 35 (32- 37) long; claw digitules (Fig. 2-P) 23 (17- 26) long; claws (Fig. 2-Q) with denticle. Di- agnosis.— Thirty-six marginal setae occur only in 7. cerifera and serve to separate it from all other Towmeyella. Forty-four mar- ginal setae are found in 7. parvicornis, and all other species possess 32 marginal setae. Toumeyella liriodendri (Gmelin) Fig. 3 Coccus liriodendri Gmelin 1789: 2220. Lecanium tulipiferae Cook 1878: 192. Lecanium liriodendri (Gmelin). Cockerell 1899: 271, Herrick 1911: 12, Carnes 1906: 40. Eulecanium tulipiferae (Cook). King 1902: 59. Eulecanium (?) liriodendri (Gmelin). Fer- nald 1903: 190. Lecanium (Toumeyella) liriodendri (Gme- lin). Pettit and McDaniel 1920: 10. Toumeyella liriodendri (Gmelin). Sanders 1909: 447, Jarvis and Guelph 1911: 70, Dietz and Morrison 1916: 249, Houser 1918: 301, Berger 1922: 68, Harned 1923: 26, Hollinger 1923: 63, Merrill and Chaf- fin 1923: 273, Trimble 1925: 6, Wells 1926: 257, Trimble 1928: 44, Steinweden 1929: 227, Felt and Rankin 1932: 460, Doane et al. 1936: 380, Dodge and Rick- ett 1943: 407, Craighead 1950: 144, Mil- liron 1959: 28, Pirone et al. 1960: 473, Burns 1970: 1, Burns and Donley 1970: 228, 1971: 532, Donley and Burns 1971: 1, Williams and Kosztarab 1972: 164, Kosztarab 1977: 184, Gill 1982: 1, Ha- mon and Williams 1984: 119, Gill 1988: ality Specimens studied.— Liriodendron tulip- ifera: 1(4), Valley Mills, IN, H. Morrison (UCDC); 1(5), Knoxville, TN, let. 24 Sep 1941, G. M. Bentley (USNM); 1(2), Mont- gomery Co., Blacksburg, VA, 16 Sep 1968, MLW (AUEM); 1(10), Simpson, IL, 7 Aug 1969, J. E. Appleby (USNM); 1, Green Co., Mt. Morris, PA, 1 Apr 1970, D. P. Burns (FSCA). Liriodendron: 2(9), College Park, MD, 9 May 1938, H. S. McConnell (USNM); 1(11), San Jose, CA, 16 Nov 1945, let. E. O. Essig, Foster (USNM). Tulip pop- lar: 1(12), Kennett Sq., PA, 21 Aug 1950, C. A. Thomas (USNM). Tulip tree: 1(16), Kingston, NY, 10 Aug 1949, J. A. Naegele (USNM). Additional host plants and distribu- tion.—Toumeyella liriodendri probably is native to the North American yellow-poplar area ranging from New York and Connect- icut to Florida and west through the Mis- sissippi River Valley (Burns and Donley 1970). It also occurs in California on shade and ornamental plantings of yellow-poplar and magnolia (Williams and Kosztarab 1972). Toumeyella liriodendri occurs on numer- ous hosts including: Magnolia acuminata, M. grandiflora, M. nigra, M. obovata, M. sinensis, M. soulangiana, M. soulangiana var. alexandrina, M. stellata, M. virginiana, Michellia fuscata, and cape jessamine (Gar- denia jasminoides?). Numerous authors re- port varied hosts such as Cephalanthus spp.., Gardenia jasminoides, Gordonia lasianthus, Juglans spp., Tilia spp. (Donley and Burns 1965), Magnolia lennei (Merrill 1953), Magnolia kobus (Sleesman 1945), Ascyrum edinianum, A. hypericoides, A. tetra- petalum, Carya cordiformis, Cassia fasci- culata, and Hypericum cistifolium (Hamon and Williams 1984). It is doubtful that 7. liriodendri occurs on Cephalanthus spp., but rather is a misidentification of 7. cerifera (Williams and Kosztarab 1972). Records on Ascyrum spp. and Hypericum spp. are most likely misidentifications of an undescribed Toumeyella species. General appearance.—Body (Fig. 3-A) VOLUME 92, NUMBER 1 oval, 559 (506-724) long, 333 (293-521) wide. Dorsum.— Marginal setae (Fig. 3-B) 24 (19-31) long on head tapering to 10 (7- 12) long near anal cleft, stout, tapering to a point, often curved posteriorly, distribu- tion: 8 anteriorly between eyes, 2 on each side between eyes and anterior spiracular setae, 2 on each side between anterior and posterior spiracular setae, 16 on posterior of body. Three spiracular setae (Fig. 3-C) in each spiracular furrow; median seta 29 (25- 33) long, lateral setae 7 (6-8) long. Small (2 um) bilocular (Fig. 3-D) and small (2 um) quinquelocular (Fig. 3-E), occasionally 4- and 6-locular, pores in submedian and submarginal longitudinal rows. Anal plates (Fig. 3-H).—Each plate with dorsum dense- ly reticulated, 66 (58-69) long, 32 (28-37) wide; cephalolateral margin 43 (37-47) long, caudolateral margin 36 (30-41) long. Ven- ter.—Antennae (Fig. 3-L) 161 (150-171) long. Legs (Fig. 3-M) 238 (229-253) long, microctenidia (Fig. 3-N) at tibial apex; tar- sal digitules (Fig. 3-O) 48 (42-52) and 34 (27-38) long; claw digitules (Fig. 3-P) 24 (22-25) long; claws (Fig. 3-Q) with denticle. Diagnosis.— Toumeyella liriodendri is most similar to 7. nectandrae, though dorsal mul- tilocular disc pores have only been observed in T. liriodendri. All other species have sim- ple dorsal disc pores. Toumeyella mirabilis (Cockerell) Fig. 4 Lecanium mirabile Cockerell 1895: 3. Toumeyella mirabilis (Cockerell). Cockerell 1902: 452, Fernald 1903: 179, Ferris 1919: 45, Ferris 1921: 91, MacGillivray 1921: 181, Steinweden 1929: 227, Wil- liams and Kosztarab 1972: 158, Taber et al. 1975: 439, Ward et al. 1977: 100. Specimens studied. — Mesquite twigs: 1(7), Nogales, AZ, 21 Apr 1940 (USNM). Pro- sopis juliflora var. velutina: 3(8), Tucson, AZ, 11 May 1950, M.E. Elve (UCDC). Pro- sopis sp.: 3(10), Galiuro Mts., AZ, 26 May 1897, Hubbard (USNM); 3, Cochise Co., AZ, 27 Jul 1969, M. Kosztarab (AUEM). 100 wm A First instar, Toumeyella liriodendri (Gme- Fig. 3. lin) 1789. Additional host plants and distribu- tion.— Toumeyella mirabilis has also been recorded on Prosopis glandulosa and P. ju- liflora var. glandulosa. Toumeyella mira- bilis has been recorded from NM, TX and Mexico. General appearance.—Body (Fig. 4-A) oval, 616 (514-847) long, 370 (319-503) wide. Dorsum.— Marginal setae (Fig. 4-B) 17 (11-27) long, stout, tapering to a point, curved posteriorly, distribution: 8 anterior- ly between eyes, 2 on each side between eyes and anterior spiracular setae, 2 on each side between anterior and posterior spiracular setae, 16 on posterior of body. Three spi- racular setae (Fig. 4-C) in each spiracular furrow; median setae 78 (32-128) long: in anterior set, lateral setae 13 (10-20) long: in posterior set, anterior lateral seta 12 (7- 17) long, and posterior lateral seta 21 (9- 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. erell) 1895. First instar, Toumeyella mirabilis (Cock- 33) long. Small (2 um) bilocular pores (Fig. 4-D) in a submedian longitudinal row. Sim- ple disc pores (Fig. 4-E) in submedian and submarginal longitudinal rows. Anal plates (Fig. 4-H).—Each plate with dorsum dense- ly reticulated, 78 (72-84) long, 33 (25-38) wide; cephalolateral margin 48 (43-52) long, caudolateral margin 43 (35-48) long. Ven- ter.—Antennae (Fig. 4-L) 170 (157-179) long. Legs (Fig. 4-M) 299 (267-323) long, microctenidia at tibial apex absent: tarsal digitules (Fig. 4-N) 58 (54-61) and 43 (38- 46) long; claw digitules (Fig. 4-O) 28 (23- 33) long; claws (Fig. 4-P) with denticle. Di- agnosis.— The spiracular setae will serve to separate 7. mirabilis from all other Tou- meyella. In T. mirabilis the median spirac- ular setae are 5-6 times longer than the an- terior set of laterals, small dorsal bilocular pores occur in submedian rows only, anal plates are densely reticulated, and microcte- nidia are absent. Toumeyella mirabilis has only been collected on Prosopis. Toumeyella nectandrae Hempel Fig. 5 Toumeyella nectandrae Hempel 1929: 64, Lepage 1938: 347. Specimens studied. — Nectandra sp.: 3(24), Sao Roque, S. Paulo, Brazil, 27 Oct 1931, H. S. Lepage (AUEM). General appearance.—Body (Fig. 5-A) oval, 490 (430-546) long, 282 (247-314) wide. Dorsum.— Marginal setae (Fig. 5-B) 30 (25-37) long, stout, slightly curved pos- teriorly, distribution: 8 anteriorly between eyes, 2 on each side between eyes and an- terior spiracular setae, 2 on each side be- tween anterior and posterior spiracular se- tae, 16 on posterior of body. Three spiracular setae (Fig. 5-C) in each spiracular furrow; median setae 34 (31-38) long, lateral setae 8 (6-19) long. Small (2 um) bilocular (Fig. 5-D) and simple disc (Fig. 5-E) pores in sub- median and submarginal longitudinal rows. Anal plates (Fig. 5-H).—Each plate with dorsum densely reticulated, 61 (53-67) long, 30 (27-32) wide; cephalolateral margin 37 (28-41) long, caudolateral margin 38 (32- 42) long. Venter.— Antennae (Fig. 5-K) 159 (152-165) long. Legs (Fig. 5-L) 231 (211- 242) long, microctenidia (Fig. 5-M) at tibial apex; tarsal digitules (Fig. 5-N) 43 (41-46) and 30 (26-32) long; claw digitules (Fig. 5-O) 21 (20-22) long; claws (Fig. 5-P) with den- ticle. Diagnosis. — Toumeyella nectandrae is similar to 7. liriodendri, but Toumeyella nectandrae has simple dorsal disc pores, whereas T. liriodendri has multilocular dor- sal disc pores. Toumeyella parvicornis (Cockerell) Fig. 6 Lecanium parvicorne Cockerell 1897: 90. Toumeyella parvicornis (Cockerell). Cock- erell 1902: 452, Fernald 1903: 179, Wil- VOLUME 92, NUMBER 1 son 1917: 59, Ferris 1920: 42, Mac- Gillivray 1921: 181, Merrill and Chaffin 1923: 274, Williams and Kosztarab 1972: 171, Williams and Cobb 1982: 93, Ha- mon and Williams 1984: 122. Lecanium (Toumeyella) numismaticum Pettit and McDaniel 1920: 8. Toumeyella numismaticum Pettit and McDaniel. Steinweden 1929: 227, Craig- head and Middleton 1930: 17, Orr and Hall 1931: 1087, Felt and Rankin 1932: 390, Doane et al. 1936: 375, Dodge and Rickett 1943: 485, Sleesman 1945: 44, Craighead 1950: 144, Merrill 1953: 110, Rabkin and Lejeune 1954: 570, McIntyre 1960: 325, MacAloney 1961: 1, Smirnoff and Valero 1975: 236. Specimens studied.—Long leaf pine: 1(10), Alachua Co., FL, 15 Mar 1954, G. Merrill (USNM). Pinus sp.: 1(8), Houston Co., AL, 16 Apr 1976, Reafield Vester (AUEM). P. elliottii: 1(6), Baldwin Co., Per- dido, AL, 12 May 1978, Charles H. Ray (AUEM). P. montana: 1(15), Washington, D.C., 17 Jul 1947, Wester (USNM). P. pa- lustris: 1(3), Miami, FL, 6 Sep 1977, CHR (AUEM). P. rigida: TYPE, 1(2), Lake City, FL, 10 Apr 1897, A. L. Quaintance (AUEM). P. taeda: 2(4), Auburn University Insectary, Auburn, AL, 24 Oct 1974, CHR (AUEM). P. virginiana: 1(4), Blount Co., nr. Oneonta, AL, 7 Aug 1981, CHR (AUEM). Spruce: 1(3), Mathews Co., VA, 31 May 1968, T. E. Dinwiddie (AUEM). Additional host plants and distribu- tion.— Toumeyella parvicornis occurs pri- marily on pine species including: Pinus ni- gra, P. strobus (Williams and Kosztarab 1972), P. banksiana, P. caribaea, P. clausa, P. densiflora, P. echinata, P. glabra, P. het- erophylla, P. inops, P. mugo, P. mugo var. pumilio, P. mugo var. rostrata, P. pinea, P. ponderosa, P. radiata, P. resinosa, P. sinen- sis, P. sylvestris, and Zygocactus truncatus. Toumeyella parvicornis has been record- ed in most states east of the Mississippi Riv- er and north to Manitoba, Canada (Wil- | i ee ; 7 sf a a a ae —— MS = \-I ] OS — 4 A Siz Q 4 fe, |\ ms a 27 y KS J Pa aN c A - \ At {IK ‘> Ty v\ Lr. : \ / om oF neuen | SF Ve ease Yy N { - \i4 --N \) DON BS 0) ) Cy J: gin Ww VY ¥ H pia ‘al ay “ / (len / Fig. 5. First instar, Toumeyella nectandrae Hempel 1929. liams and Kosztarab 1972) as well as IA, NB, ND, SD, and WI (MacAloney 1961). General appearance.— Body (Fig 6-A) oval, 617 (487-902) long, 348 (284-561) wide. Dorsum.— Marginal setae (Fig. 6-B) 17 (12-21) long on head tapering to 13 (9- 16) long near anal cleft, slender, curved pos- teriorly, distribution: 8 anteriorly between eyes, 3 on each side between eyes and an- terior spiracular furrow, 3 on each side be- tween anterior and posterior spiracular fur- row, 16 on posterior of body. Two setae at apex of each spiracular furrow undifferen- tiated from marginal setae. Bilocular pore clusters (Fig. 6-C) in a submarginal longi- tudinal row. Simple disc pores (Fig. 6-D) in submedian and submarginal longitudinal rows. Anal plates (Fig. 6-G).—Each plate 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. First instar, Toumeyella parvicornis (Cock- erell) 1897. with sparsely distributed microspines on dorsum, 64 (56-70) long, 27 (25-31) wide; cephalolateral margin 36 (30-43) long, cau- dolateral margin 39 (36-43) long. Venter. — Antennae (Fig. 6-K) 183 (162-207) long. Legs (Fig. 6-L) 249 (231-271) long, mi- croctenidia (Fig. 6-M) at tibial apex; tarsal digitules (Fig. 6-N) 49 (40-57) and 35 (30- 39) long; claw digitules (Fig. 6-O) 24 (21- 30) long; claws (Fig. 6-P) simple, without denticle. Diagnosis.— Dorsal bilocular pore clusters, spiracular setae undifferentiated from marginal setae, and simple tarsal claws are characteristics unique to 7. parvicornis. Toumeyella pini (King) Fig. 7 Lecanium pini King 1901: 334. Toumeyella (?) pini (King). Cockerell 1902: 452, Fernald 1903: 179. Lecanium corrugatum Thro 1903: 216, Fer- nald 1903: 179. Lecanium (Toumeyella) corrugatum (Thro). Pettit and McDaniel 1920: 6. Toumeyella corrugatum (Thro). Harned 1923: 26, Doane et al. 1936: 375. Toumeyella pini (King). Ferris 1920: 42, MacGillivray 1921: 181, Steinweden 1929: 227, Craighead and Middleton 1930: 17, Doane et al. 1936: 375, Dodge and Rickett 1943: 485, Craighead 1950: 145, MacAloney 1961: 3, Williams and Kosztarab 1972: 171, Hamon and Wil- liams 1984: 124. Specimens studied.—Pinus mugo: 1(2), Oxford, CT, let. of 13 Jul 1939, R. C. Brown (USNM); 1(9), Waynesboro, VA, 31 May 1941, F. R. Freund (USNM). P. pungens: 1(2), Blain, PA, 30 July 1959, A. T. Drooz (USNM). P. taeda: 1, Auburn, AL, 25 Jul 1974, CHR (AUEM); 1(4), Airport Marsh, Dauphin Island, Mobile Co., AL, 13 May 1978, CHR (AUEM); 1(2), Lee Co., Au- burn, AL, 21 Jun 1977, CHR (AUEM). P. virginiana: 1(10), V.P.I. Plot 531/d, Blacks- burg, VA, 11 May 1969, MLW (VPIC). Red pine: 1(10), Cole Nursery, Tazewell Co., VA, Jun 1957, F. R. Freund (USNM). Pine: 1(3), McConnelsburg area, PA, 10 June 1948, G. Sleesman (USNM). Additional host plants and distribu- tion.—Toumeyella pini has been reported on numerous pine species including: Pinus contorta, P. resinosa, P. sylvestris (Williams and Kosztarab 1972), P. palustris, P. seroti- na (Hamon and Williams 1984), P. austria- ca (Jarvis and Guelph 1911), P. divaricatus. P. echinata, P. elliottii, P. pinaster, P. rigida, and P. strobus. The type specimen was collected in NY with subsequent collections in TX (Hamon and Williams 1984), DC, FL, GA, MD, MI, MS, NJ, OH, and SC. General appearance.—Body (Fig. 7-A) oval, 594 (441-921) long, 336 (264-494) wide. Dorsum.— Marginal setae (Fig. 7-B) 22 (14-32) long on head tapering to 12 (9- 15) long near anal cleft, slender, curved pos- teriorly, distribution: 8 anteriorly between VOLUME 92, NUMBER | eyes, 2 on each side between eyes and an- terior spiracular setae, 2 on each side be- tween anterior and posterior spiracular se- tae, 16 on posterior of body. Three spiracular setae (Fig. 7-C) in each spiracular furrow; median setae 21 (19-25) long, lateral setae 7 (5-10) long. Small (2 um) bilocular (Fig. 7-D) and simple disc (Fig. 7-E) pores in sub- median and submarginal longitudinal rows. Anal plates (Fig. 7-H).—Each plate with sparsely distributed microspines on dor- sum, 63 (57-73) long, 30 (23-35) wide; cephalolateral margin 38 (31-47) long, cau- dolateral margin 38 (31-47) long. Venter. — Antennae (Fig. 7-L) 166 (143-183) long. Legs (Fig. 7-M) 237 (219-261) long, mi- croctenidia (Fig. 7-N) at tibial apex; tarsal digitules (Fig. 7-O) 49 (33-56) and 33 (27- 36) long; claw digitules (Fig. 7-P) 21 (16- 25) long; claws (Fig. 7-Q) with denticle. Bi- ological notes.—Toumeyella pini is often found in mixed infestations with 7. parvi- cornis (Williams and Kosztarab 1972). Di- agnosis.— Toumeyella pini appears similar to T. virginiana. Toumeyella pini has small dorsal bilocular pores in submarginal and submedian longitudinal rows, whereas 7. virginiana has bilocular pores in submar- ginal, submedian, and intermediate longi- tudinal rows. Toumeyella quadrifasciata (Cockerell) Fig. 8 Lecanium quadrifasciatum Cockerell 1895: 3. Toumeyella quadrifasciata (Cockerell). Cockerell 1902: 452, Fernald 1903: 179, MacGillivray 1921: 181. Specimens studied.— Robinia neomexi- cana: TYPE, 7(37), Organ Mts., NM, Ed Owen (AUEM). General appearance.—Body (Fig. 8-A) oval, 579 (514-622) long, 324 (309-343) wide. Dorsum.— Marginal setae (Fig. 8-B) 22 (19-27) long on head tapering to 15 (12- 18) long near anal cleft, slender, curved pos- teriorly, distribution: 8 anteriorly between eyes, 2 on each side between eyes and an- terior spiracular setae, 2 on each side be- Fig. 7. First instar, Toumeyella pini (King) 1901. tween anterior and posterior spiracular se- tae, 16 on posterior of body. Three spiracular setae (Fig. 8-C) in each spiracular furrow; median setae 27 (25-32) long, lateral setae 8 (6-9) long. Large (4 um) bilocular (Fig. 8-D) and simple disc (Fig. 8-E) pores in sub- median and submarginal rows. Bilocular pores often bent in a bow-shaped configu- ration. Anal plates (Fig. 8-H).—Each plate with dorsum densely reticulated, 67 (63-70) long, 35 (31-37) wide; cephalolateral mar- gin 44 (38-47) long, caudolateral margin 41 (40-44) long. Venter.— Antennae (Fig. 8-L) 169 (157-181) long. Legs (Fig. 8-M) 270 (261-279) long, microctenidia at tibial apex absent; tarsal digitules (Fig. 8-N) 47 and 32 long; claw digitules (Fig. 8-O) 24 (23-26) long; claws (Fig. 8-P) with denticle. Diag- nosis.—Dorsal bilocular pores (>3 pm) which are predominantly bow shaped, in submarginal and submedian longitudinal rows occur only in 7. quadrifasciata. 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON : S WY AN) s Nad \ | ce yc s fe a n= 100 ws (g) 4) A iC e, Fig. 8. First instar, Toumeyella quadrifasciata (Cockerell) 1895. Toumeyella virginiana Williams and Kosztarab Fig. 9 Toumeyella virginiana Williams and Kosz- tarab 1972: 182, Hamon and Williams 1984: 126. Specimens studied.— Pinus: 1(2), Au- burn, AL, 26 Jun 1974, L. C. Ray (AUEM); 1(7), Pine Mt. Camp 35A, Harris Co., GA, 5 May 1979, K. Manuel (AUEM). P. echin- ata: 2(4), old water works road, Auburn, AL, 15 Aug 1974, MLW, J. Gilder, CHR (AUEM),; 1(4), Bullock Co., Perote, AL, 26 Apr 1975, MLW (AUEM). P. taeda: 1(2), Dorchester Co., MD, 16 Sep 1971, MLW (AUEM); 2(4), Auburn, AL, 19 Jun 1974, L. C. Ray (AUEM); 1(2), Elmore Co., AL, 10 Aug 1974, CHR (AUEM). Fig. 9. First instar, Toumeyella virginiana Wil- hams & Kosztarab 1972. Additional host plants and distribu- tion.— Toumeyella virginiana has been col- lected on Pinus elliottii, P. palustris, and P. virginiana. Hamon and Williams (1984) re- port occurrence on P. clausa and P. glabra. Toumeyella virginiana has been collected in FL and VA. General appearance.—Body (Fig. 9-A) oval, 618 (330-965) long, 398 (318-681) wide. Dorsum.— Marginal setae (Fig. 9-B) 12 (10-16) long, slender, curved posteriorly, distribution: 8 anteriorly between eyes, 2 on each side between eyes and anterior spirac- ular setae, 2 on each side between anterior and posterior spiracular setae, 16 on pos- terior of body. Three spiracular setae (Fig. 9-C) in each spiracular furrow; median setae 20 (16-25) long, lateral setae 8 (5-12) long. VOLUME 92, NUMBER 1 Small (2 um) bilocular pores (Fig. 9-D) in submedian, submarginal and intermediate longitudinal rows. Simple disc pores (Fig. 9-E) in submedian and submarginal longi- tudinal rows. Anal plates (Fig. 9-H).— Each plate with dorsum coarsely reticulated; 73 (68-77) long, 34 (27-38) wide; cephalolat- eral margin 47 (38-52) long, caudolateral margin 45 (35-52) long. Venter.— Antennae (Fig. 9-L) 178 (152-196) long. Legs (Fig. 9-M) 232 (225-270) long, microctenidia (Fig. 9-N) at tibial apex; tarsal digitules (Fig. 9-O) 45 (36-51) and 29 (25-31) long; claw digitules (Fig. 9-P) 21 (19-25) long; claws (Fig. 9-Q) with denticle. Diagnosis. — Tou- meyella virginiana appears quite similar to T. pini except for the presence of small dor- sal bilocular pores in submedian, submar- ginal, and intermediate longitudinal rows. Toumeyella pini has dorsal bilocular pores in submedian and submarginal longitudinal rows only. DISCUSSION The first-instar nymphs of all species of Toumeyella included in this study can be distinguished by morphological characters, but no means of distinguishing sexual di- morphism in the first instar were found. Also, no group characteristics were ob- served which would differentiate the Pinus feeding species of Toumeyella (T. parvicor- nis, T. pini, and T. virginiana) from those species associated with non-pine hosts (7. cerifera, T. liriodendri, T. mirabilis, T. nec- tandrae, and T. quadrifasciata). The 8 first-instar Toumeyella described in this paper share many features with first instars of Pseudophilippia quaintancii Cockerell and Neolecanium cornuparvum Thro, most significantly 5-segmented an- tennae. Steinweden (1929) considered Tou- meyella, Neolecanium, and Pseudophilippia as constituting one genus, while Williams and Kosztarab (1972) treated all three as valid genera. Comparison of Toumeyella first instars with descriptions of first instars of Pseudophilippia quaintancii and Neole- 55 canium cornuparvum by Ray and Williams (1980, 1983) confirms the close relationship of the Toumeyella to these two genera and further justifies the placement of Touwmey- ella, Neolecanium, and Pseudophilippia in the tribe Toumeyellini. The invaginated bi- locular pores of P. quaintancii are unique to that species and will separate it from all other Coccidae, but N. cornuparvum seems to fit well with Toumeyella based on com- parisons of the first instar nymph and adult male. The dense pattern of bilocular pores of the adult female, however, suggests a closer relationship to other Neolecanium species. Preliminary investigations by the junior author indicate that some of the species currently placed in Neolecanium are actually closer to Toumeyella than other members of Neolecanium. Because of such confusion within the genus we feel it is pre- mature to move N. cornuparvum from its current placement until a review of Neole- canium is conducted. A ninth species, 7. sonorensis (Cockerell and Parrott 1899), included in this study, has 6-segmented antennae and is not con- generic with the genotype, 7. mirabilis or the remaining species studied, which have 5-segmented antennae. Preliminary inves- tigations of the adult female morphology seem to indicate that 7. sonorensis should be in a different genus than Toumeyella, but further revisionary work is needed before determining the placement of this species historically included in the Toumeyella. Additionally, distribution gaps within the United States and among temperate, neo- tropical, and tropical regions suggest further collecting in these areas is needed. ACKNOWLEDGMENTS Thanks are extended to the museum staffs and institutions listed in the Materials and Methods section for the loan of specimens. Also, thanks to W. E. Clark and G. R. Mul- len for manuscript review. This paper is published as Alabama Agricultural Exper- iment Station Journal Series No. 17-881785. 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Berger, E. W. 1922. Another apparently new ento- mogenous fungus from the hammock. Fla. State Horticultural Soc. 35: 68-71. Burns, D. P. 1970. Insect enemies of yellow-poplar. NE. Forest Exp. Sta. USDA Forest Serv. Res. Pa- per NE-159. 15 pp. Burns, D. P. and D. E. Donley. 1970. Biology of the tuliptree scale Toumeyella liriodendri (Homop- tera: Coccidae). Ann. Entomol. Soc. Amer. 63: 228-235. 1971. Quantitative field collection of hon- eydew. Ann Entomol. Soc. Amer. 64: 532-534. Carnes, E.K. 1906. The Coccidae of California. Calif. State Commission of Horticulture. 70 pp. Cockerell, T. D. A. 1895. New North American Coc- cidae. Psyche 7 (1894-1896) (sup.): 1-4. 1897. New and little-known Coccidae from Florida. Psyche 8: 89-91. . 1899. 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Quart. Bul. 2: 2-65. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 58-65 UNUSUAL CADDISFLY (TRICHOPTERA) FAUNA OF SCHOOLHOUSE SPRINGS, LOUISIANA, WITH DESCRIPTION OF A NEW SPECIES OF DIPLECTRONA (HYDROPSYCHIDAE) JOHN C. MORSE AND CHERYL B. BARR (JCM) Department of Entomology, Clemson University, Clemson, South Carolina 29634; (CBB) Department of Entomology, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803. Abstract. —The caddisflies (Trichoptera) of Schoolhouse Springs, Jackson Parish, Lou- isiana, have been studied since 1959, first by H. H. Ross and some of his colleagues and subsequently by several of his students. At least 43 species of caddisflies have been captured in or beside the spring, one of which (belonging to the genus Diplectrona, Hydropsychidae) is described in this paper. The Springs are the type locality for five caddisfly species and one stonefly species. Five of these species of insects are known from nowhere else. Lep- idostoma serratum Flint and Wiggins occurs here apparently as an isolated population, far removed from populations in Connecticut and North Carolina. Key Words: Trichoptera and other aquatic insects have been studied sporadically at Schoolhouse Springs, Jackson Parish, Louisiana (T17N, R1W, about middle of Sec. 12, about 6 miles north of Eros), at least since 1959 by H. H. Ross and several of his colleagues and stu- dents. The fauna and flora of the Springs are sufficiently unusual that the Louisiana Nature Conservancy recently purchased the site to help assure perpetual protection for it and its biota. Schoolhouse Springs are located about a mile west of the community of Indian Vil- lage, named for the band of Choctaws which inhabited the area until the 1820’s. The Springs were named by Ross in 1973 for Springhill Academy which once stood above them on the hill about 100-200 m east (Brown and Asken 1982). For years the site was relatively unknown and undisturbed except by aquatic entomologists. However, in 1987 the land surrounding the Springs Trichoptera, spring faunas, Louisiana, Diplectrona was sold and slated for logging and devel- opment. When the authors urged protection of the area because of its unique aquatic biota, the Louisiana Nature Conservancy immediately launched a thorough investi- gation and named the Springs a high priority site for acquisition. In September 1988, 30 acres, including the Springs and a portion of the bayhead community along the spring- brook, were purchased and designated as a preserve by the Conservancy. At least five distinct, permanent, small springs of cool, clear water bubble from white sands at the base of moderately steep slopes and converge to form Schoolhouse Branch, a shallow stream averaging less than 10 cm depth. Vegetation surrounding the Springs is primarily a mixed hardwood-pine forest; however, a bayhead swamp com- munity is supported in thick accumulations of peaty muck surrounding the springhead and along Schoolhouse Branch. VOLUME 92, NUMBER 1 At least 43 species of Trichoptera have been recognized from Schoolhouse Springs (Table 1), including three endemics: Cheu- matopsyche morsei Gordon, 1974; Hydrop- tila ouachita Holzenthal and Kelley, 1983; and Chimarra holzenthali Lago and Harris, 1987. Herein, JCM adds a fourth endemic caddisfly species, belonging to the genus Di- plectrona. The stonefly Leuctra szczytkoi Stark and Stewart, 1981 (Plecoptera, Leuc- tridae) apparently is endemic to the Springs, also. In addition, Schoolhouse Springs is the type locality for Agarodes libalis Ross and Scott, 1974. The range of the sister species of each of the endemic caddisflies lies to the east or north of Schoolhouse Springs. The sister species of the new Diplectrona species is D. modesta Banks, whose range includes east- ern North America from Florida to New Hampshire and Quebec to Illinois to west- ern Arkansas, northern Louisiana (School- house Springs), and southern Mississippi. The sister species of C. holzenthali is Chi- marra feria Ross (Lago and Harris 1987), a species whose range includes eastern Texas, northwestern Louisiana, the Ouachita and Ozark Mountains, and the Great Lakes re- gion. Speaking of C. holzenthali Ross (1965) noted that “it is possible that [this] local endemic species originated from wind- blown vagrants. It is also possible that during the glacial maxima some of the con- necting streams from the Ouachita Moun- tains were cool enough to afford avenues of dispersal by which caddisflies reached their present spring habitats.”” The sister species of C. morsei is Cheumatopsyche virginica Denning (Gordon 1974), from Coastal Plain localities in New Jersey, Delaware, Virginia, South Carolina, Georgia, Florida, and southern Mississippi (Gordon 1974, Morse et al. 1980, Harris et al. 1982, Lago et al. 1982, Lake 1984). That of H. ouachita is Hydroptila poirrieri Holzenthal and Kelley, from Mississippi and southeastern Louisi- ana (Holzenthal and Kelley 1983). Thus it appears likely that the endemic species of 59 Schoolhouse Springs evolved in isolation here after having arrived either as exten- sions of ranges of their ancestors from the North during Pleistocene glaciation (D. rossi and C. holzenthali) or as relict popu- lations near the edge of the Mississippi Em- bayment and southeastern Coastal Plain (C. morsei and H. ouachita). Showing a pattern similar to the latter two species, although yet without allopatric spe- ciation, Agarodes libalis occurs throughout the southern Coastal Plain from Delaware to Florida to Louisiana; Schoolhouse Springs, its type locality, is at the western edge of its range (Ross and Scott 1974, McEwan 1980, Holzenthal et al. 1982, Lake 1984). Lepidostoma (Mormomyia) serratum Flint and Wiggins apparently is the only species of this large genus occurring in the Springs. According to Weaver (1983, 1988), the only other populations known for the species are found in Connecticut and North Carolina. Collections of adults were made from 30 March through 19 May in the Springs; three prepupae taken on 24 August suggest that a second generation may appear here in the autumn. Diplectrona rossi Morse NEw SPECIES Mature Larva: Length 9.0-15.0 mm; head capsule width 1.08-1.33 mm. Similar to D. modesta in general structure. Head (Fig. 1) generally dark brown to near black with three conspicuous reddish yellow regions on frontoclypeus, including pair of regions in its lateral broadened areas and single region at its dorsal apex, variably light brown re- gions in center and anterior margin of frontoclypeus not as conspicuous. Occipital region and dorsolateral areas light brown. Frontoclypeus evenly convex anteriorly, broadened laterally at mid-length to level of eyes, and with pair of variably obtuse angles near anterior tentorial pits. Mandi- bles resembling those of D. modesta, left mandible without high thumb-like dorso- 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Adult Trichoptera of Schoolhouse Springs, Louisiana. * = Species for which the type locality is Schoolhouse Springs. Hydropsychidae Cheumatopsyche Diplectrona Hydropsyche Potamyia Hydroptilidae Hydroptila Orthotrichia Oxyethira Lepidostomatidae Lepidostoma Leptoceridae Ceraclea Leptocerus Oecetis Triaenodes Limnephilidae Pycnopsyche Molannidae Molanna Philopotamidae Chimarra Polycentropodidae Neureclipsis Nyctiophylax Polycentropus Phylocentropus Psychomyiidae Lype Sericostomatidae Agarodes Ss burksi Ross *morsei Gordon pasella Ross pettiti (Banks) modesta Banks *rossi Morse, n. sp. decalda Ross orris Ross prob. placoda Ross flava (Hagen) novicola Blickle & Morse remita Blickle & Morse *ouachita Holzenthal & Kelley waubesiana Betten spp. nr. consimilis Morton sp. maculata (Banks) Group aegerfasciella (Chambers) novasota Ross serratum Flint & Wiggins cancellata (Betten) protonepha Morse & Ross spongillovorax Resh transversa (Hagen) americanus (Banks) cinerascens (Hagen) ditissa Ross inconspicua (Walker) nocturna Ross ochracea (Curtis) osteni Milne ignitus (Walker) spp. blenda Sibley tryphena Betten aterrima Hagen *holzenthali Lago & Harris p. affinis (Banks) crassicornis Walker lucidus (Hagen) placidus (Banks) diversa (Banks) *libalis Ross & Scott 3 Jun-14 Sep 27 May-3 Jun 3 Jun 30 Mar-24 Aug 14 Apr-24 Aug 14 Apr-7 Jul 14 Apr 8 May-24 Aug 3 Jun 7 Jul 14 Apr 3 Jun-24 Aug 30 Mar-24 Aug 3 Jun—24 Aug 7 Jul 3 Jun 3 Jun-24 Aug 14 Apr-24 Aug 30 Mar-19 May 3 Jun 14 Apr 3 Jun 14 Apr 8 May-7 Jul 14 Apr-24 Aug 14 Apr 14 Apr-24 Aug 8 May-3 Jun 24 Aug 14 Apr-27 May 14 Apr—24 Aug (larvae only) 14 Apr-8 May 23-28 Apr—pupae 8 May-24 Aug 7 Jul-14 Sep 3 Jun 8 May-3 Jun 30 Mar 24 Aug 30 Mar 30 Mar-24 Aug 19 May-7 Jul VOLUME 92, NUMBER 1 lateral projection. [Left mandible of D. mo- desta with dorso-mesal brush of setae pres- ent but not illustrated by Ross 1944, fig. 286.] Pupa: Length 9.0 mm. Similar to D. mo- desta in general structure. Left mandible with five teeth, basal tooth as far from others as next most basal tooth is from apex; right mandible with four teeth. Mesal fork of each apical process 1.5 = as long as lateral fork. Male: Length 8.8-9.5 mm; forewing 7.6- 7.9 mm. Similar to male of D. modesta in general structure. Warts and other struc- tures on dorsum of head (Fig. 3) similar in size and shape to those of male of D. mo- desta (Fig. 4), except with diagonal lines of posterior vertex behind pair of conical pro- tuberances ending on epicranial stem (ep.su.), anteriad of transverse postoccipital sulcus (po.oc.). Antennae each with scape and pedicel usually at least as dark as, or darker than, flagellum. Eyes not unusually large; in dorsal view, ratio of greatest width of one eye to narrowest width of vertex (eye : vertex ratio) about 0.475; malar space (Fig. 5) slightly broader than for male of D. mo- desta (Fig. 6). Extreme lateral corners of pronotum, laterad of lateral warts, each with generally conspicuous pale cream-colored spot (Fig. 5, spt). Male genitalia (Figs. 7-10): Similar to those of D. modesta in general structure. Anterolateral corners of sternum V each with long slender glandular structure. Tergum IX divided anteriorly (Fig. 8, IX), with pair of rugose patches. Superior appendages not distinct, represented by region of setae on pair of dorsolateral lobes (do.lat., = terga IX-X?), each lobe with apex truncate in lat- eral view (Fig. 7), acute dorsally. Tergum X divided longitudinally on meson, each half tapered in dorsal view (Fig. 8, X), rounded in lateral view (Fig. 7, X), lightly sclerotized and setose apically. Inferior appendages each with basal segment bent slightly caudad near apex in lateral view (Fig. 7, inf.app.), curved mesad and clavate in caudal view (Fig. 9), apical segment curved mesad and spatulate 61 in caudal view. Phallus simple (Fig. 10), similar to that of D. modesta. Female: Length 9.4-11.1 mm, forewing 8.0-9.2 mm. Similar to female of D. mo- desta in general structure. Warts and other structures on dorsum of head (Fig. 11) sim- ilar in size and shape to those of female of D. modesta, except with diagonal lines of posterior vertex behind pair of conical pro- tuberances ending on epicranial stem, an- teriad of transverse postoccipital sulcus. Eyes not unusually large. Extreme lateral corners of pronotum, laterad of lateral warts, with- out conspicuous pale cream-colored spots. Female genitalia: Generally resembling those of D. modesta very closely. Median plate (Figs. 12-13) spatulate anteriorly, without longitudinal dorsal carina. Holotype: Male, LOUISIANA: Jackson Parish; Schoolhouse Springs; RI W, T17N, Sec. 12; 6 miles north of Eros; 14 April 1988, mercury vapor and ultraviolet lights, C. B. Barr. Deposited in U.S. National Mu- seum of Natural History (USNM). Paratypes: All same locality: Same data as holotype, | male, 3 females, deposited in Louisiana State University Insect Collec- tion (LSUC); 7 May 1987, C. B. and J. E. Barr, 4 females (USNM and LSUC); 27 May 1972, at light, Ross and Smith, 5 females, deposited in Clemson University Arthro- pod Collection (CUAC); 3 June 1973, J. C. Morse and J. A. Louton, | male, | female (CUAC); 30 March 1973, H. H. Ross et al., 26 larvae, 1 female pupa (CUAC); 14 April 1988, C. B Barr, 12 larvae, 1 prepupa (LSUC); 23 April 1982, R. W Holzenthal and S. W. Hamilton, 4 larvae (USNM); 28 April 1973, J. C. Morse, C. E Dunn, and J. A. Louton, 5 larvae (CUAC); 8 May 1987, C. B. Barr, 2 larvae (LSUC); 7 July 1973, J. C. Morse and J. A. Louton, 4 larvae (USNM). Etymology: named for Dr. Herbert H. Ross, founder of modern trichopterology in North America, discoverer of the unusual aquatic insect fauna of Schoolhouse Springs and author of their name, and naturalist 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 Figs. 7-10 Figs. 1-10. Characters of Diplectrona rossi and D. modesta larvae and males. 1, D. rossi larval head, dorsal view: 2, D. modesta larval head, dorsal view; 3, D. rossi male head, dorsal view; 4, D. modesta male head, dorsal view: 5, D. rossi male head and prothorax, left lateral view; 6, D. modesta male head, left lateral view; 7, D. rossi male genitalia, left lateral view; 8, D. rossi terga IX and X and phallus, dorsal view; 9, D. rossi left inferior appendage, caudal view; 10, D. rossi phallus, left lateral view. do.lat. = dorsolateral lobe of male genitalia (paired), ep.su. = epicranial suture, inf.app. = male inferior appendage (paired), IX = abdominal segment IX, po.oc. = postoccipital sulcus, X = abdominal segment X. Scale lines each 0.5 mm. VOLUME 92, NUMBER 1 63 13 16 15 Fig. 11-16. Characters of Diplectrona rossi and D. modesta females. 11, D. rossi head, dorsal; 12, D. rossi median plate of genitalia, left lateral view; 13, D. rossi median plate of genitalia, dorsal view; 14, D. modesta median plate of genitalia, left lateral view; 15, D. modesta median plate of genitalia, dorsal view; 16, same, variation. Scale lines each 0.5 mm. who first brought the senior author’s atten- tion to this species. Diagnosis: Larvae of this species are readily distinguishable from those of D. me- taqui in that the frons is evenly convex api- cally (versus notched, cf. Ross 1944, fig. 338, “Genus A’) and the left mandible does not have a high thumb-like process (versus with this process, cf. Ross 1944, fig. 282, “Genus A”). The head is relatively broader than that of D. modesta (Fig. 2), but relatively nar- rower than that of D. metaqui (Ross 1944, fig. 338). Unlike both D. metaqui and D. modesta, the frons has a pair of variably obtuse angles near the anterior tentorial pits (Fig. 1; versus evenly curved, Fig. 2), and three conspicuous reddish yellow regions occur laterally and posteriorly on the frontoclypeus, sometimes visible even in the field. The larva of D. californica Banks, 1914, is unknown. The pupa of this species differs from that of D. modesta in that the left mandible has five teeth (four in D. modesta, cf. Ross 1944, fig. 316), with the basal tooth as far from others as the next most basal tooth is from the apex; the right mandible has four teeth (five in D. modesta); and the mesal fork of each apical process is 1.5 = as long as the lateral fork (subequal in D. modesta, cf. Ross 1944, fig. 310A). The pupae of D. californica and metaqui have not been described. Males of this species resemble those of D. modesta in the relative size of warts on the vertex and in the antennae with flagellum lighter than scape and pedicel (middle wart slightly smaller [Ross 1970, fig. 4B] and scape and pedicel lighter than flagellum in D. metaqui). The eyes of D. rossi are slightly smaller than those of D. modesta (eye : ver- tex ratio of D. modesta between 0.55 and 0.65) and clearly larger than those of D. me- taqui (e:v = 0.39-0.40). Diplectrona rossi differs from both D. modesta and D. me- taqui in having the diagonal lines of the pos- terior vertex (behind the pair of conical pro- 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tuberances) ending on the epicranial stem, anteriad of the transverse postoccipital sul- cus (Fig. 3; ending on the postoccipital sul- cus in the other two species, Fig. 4). The genitalia of the male of this species differ from those of D. modesta and D. metaqui in the more nearly truncate apex of each dorsolateral lobe (Fig. 7; cf. Ross 1944, fig. 339). Also, in many specimens of D. mo- desta, the inferior appendage is nearly straight apically in lateral view, but curved posteriad in the three male specimens seen of this species (Fig. 7). The male genitalia of D. metaqui, modesta, and rossi all differ from those of californica in the pair of dor- solateral lobes distinct from the more mesal lobes of tergum X, not clearly separated in californica (Denning, 1965, as D. marga- rita; Flint 1966). Females of this species differ from those of both D. modesta and D. metaqui in the same characters of the head mentioned for the males. Additionally, the pronotum of females of the type series all lack the con- spicuous pale, cream-colored spot on the extreme lateral corners, laterad of the lateral warts, present in males of all three species (Fig. 5) and usually in females of D. mo- desta. The median plate of the genitalia (Figs. 12 and 13) lacks a longitudinal carina pres- ent on the median plate of D. modesta (Figs. 14-16). Characters of the pronotum and median plate of females of D. californica and metaqui are unknown. Further comments: Associations of larvae and pupae with adults of this new species are circumstantial. All these life history stages differ from those of known species and occur uniquely in Schoolhouse Springs. Two larvae of a Diplectrona species have been collected in a spring-fed stream in north-central Tennessee by D. Gillis and T. Kollers, students of Dr. S. W. Hamilton. These larvae resemble those of D. rossi in the presence of conspicuous light-colored spots on the frons and obtuse angles near the anterior tentorial pits. However, the background color of the head of these spec- imens is lighter than for D. rossi, the lateral spots on the frons are much larger, and a median spot is much more conspicuous than the posterior spot. A fifth North American species of this genus is suspected, but should be confirmed with data from other life his- tory stages. ACKNOWLEDGMENTS We are grateful to Drs. H. H. Ross and H. B. Boudreaux for bringing our attention to this fascinating locality and its insect fau- na. We thank J. C. Barr, C. E. Dunn, D. Gillis, S. W. Hamilton, R. W. Holzenthal, T. Kollers, J. A. Louton, and S. E. Morse for their efforts and companionship in the field collections, former owner J. E. Moore for access and historical information, the respective staffs of the Louisiana Nature Conservancy and the Louisiana Depart- ment of Wildlife and Fisheries Natural Her- itage Program for geological and biological information about the area, M. Mathis and A. D. Huryn for the loan of Diplectrona modesta and D. metaqui specimens for comparison with D. rossi, and Mrs. Yang Lian-fang for preparing the illustrations. This is Technical Contribution No. 2909 of the South Carolina Agricultural Experiment Station, Clemson University. LITERATURE CITED Banks, N. 1914. American Trichoptera—notes and descriptions. Can. Entomol. 46: 149-156, 201- 205, 252-258, 261-268, pls. 9, 10, 15, 20. Brown, S. and M. Asken, eds. 1982. Jackson Parish, Louisiana. Jackson Parish Chamber of Com- merce, Jonesboro, Louisiana. 402 pp. Denning, D. G. 1965. New Hydropsychidae (Tri- choptera). J. Kansas Entomol. Soc. 38: 75-84. Flint, O. S., Jr. 1966. Notes on certain Nearctic Tn- choptera in the Museum of Comparative Zoology. Proc. U.S. Natn. Mus. 118(3530): 373-390. Gordon, A E. 1974. A synopsis and phylogenetic outline of the Nearctic members of Cheumato- psyche. Proc. Acad. Natr. Sci. Philadelphia 126: 117-160. Harris, S. C., P. K. Lago, and J. F. Scheiring. 1982. An annotated list of Trichoptera of several streams on Eglin Air Force Base, Florida. Entomol. News 93: 79-84. VOLUME 92, NUMBER 1 Holzenthal, R. W.,S. C. Harris, and P. K. Lago. 1982. An annotated checklist of the caddisflies (Tri- choptera) of Mississippi and southeastern Loui- siana. Part III: Limnephiloidea and conclusions. Proc. Entomol. Soc. Washington 84: 513-520. Holzenthal, R. W. and R. W. Kelley. 1983. New micro-caddisflies from the southeastern United States (Trichoptera: Hydroptilidae). Florida Ento- mol. 66: 464-472. Lago, P. K. and S. C. Harris. 1987. The Chimarra (Trichoptera: Philopotamidae) of eastern North America with descriptions of three new species. J. New York Entomol. Soc. 95: 225-251. Lago, P. K., R. W. Holzenthal, and S. C. Harris. 1982. An annotated checklist of the caddisflies (Tri- choptera) of Mississippi and southeastern Loui- siana. Part I: Introduction and Hydropsychoidea. Proc. Entomol. Soc. Washington 84: 495-508. Lake, R. W. 1984. Distribution of caddisflies (Tri- choptera) in Delaware. Entomol. News 95: 215- 224. McEwan, E. M. 1980. Biology and life history of the genus A garodes (Trichoptera: Sericostomatidae) in the southeastern U.S. M.S. thesis, Clemson Uni- versity. 67 pp. Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey. 1980. Aquatic insects of Upper Three Runs Creek, Savannah River Plant, South Caro- 65 lina. Part I: Orders other than Diptera. J. Georgia Entomol. Soc. 15: 73-101. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Illinois Natr. Hist. Surv. 23: 1- 326. . 1965. Pleistocene events and insects, pp. 583- 596. In Wright, Jr., H. E. and D. G. Frey, eds., The Quaternary of the United States. Princeton University Press, Princeton, New Jersey. 1970. Hydropsychid genus A, Diplectrona (Trichoptera: Hydropsychidae). J. Georgia Ento- mol. Soc. 5: 229-231. Ross, H. H. and D. C. Scott. 1974. A review of the caddisfly genus 4 garodes, with descriptions of new species (Trichoptera: Sericostomatidae). J. Geor- gia Entomol. Soc. 9: 147-155. Stark, B. P. and K. W. Stewart. 1981. Leuctra szczyt- koi, a new stonefly from Louisiana (Plecoptera: Leuctridae). Entomol. News 92: 91-92. Weaver, J. S., HI. 1983. The evolution and classifi- cation of Trichoptera, with a revision of the Lep- idostomatidae and a North American synopsis of this family. Ph.D. dissertation, Clemson Univer- sity. 411 pp. . 1988. Asynopsis of the North American Lep- idostomatidae (Trichoptera). Contrib. Amer. Entomol. Inst. 24(2): 1-141. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 66-75 MATING BEHAVIOR OF ACIURINA MEXICANA (ACZEL) (DIPTERA: TEPHRITIDAE) JOHN JENKINS Department of Entomology, University of Arizona, Tucson, Arizona 85721 (Present address: Department of Zoology, Natural Science Building, Michigan State University, East Lansing, Michigan 48824-1115). Abstract. —Observations on the mating behavior of Aciurina mexicana (Aczél) in the laboratory are presented. Courtship and agonistic displays are named and described in detail. These displays include stereotypic body postures and wing movements, nuptial feeding, and abdominal inflation with odor production by males. Preliminary observations suggest that the male’s nuptial gift is produced from the crop, and the source of the male odor is the abdominal pleura. Mating behavior of A. mexicana is compared with other Aciurina species and tephritid flies generally. Potential use of mating behavior in fruit fly systematics is noted. Key Words: omone, systematics At least 12 species of Aciurina occur in the western United States and northern Mexico (Steyskal 1984, Dodson and George 1986). Larvae of the species for which host plants are known form galls on asteraceous plants, primarily Chrysothamnus species (Steyskal 1984, Dodson 1987b). Mating behavior has been observed for a number of Aciurina species. Tauber and Tauber (1967) reported the reproductive behavior and biology of Aciurina ferruginea (Doane) from California. Wangberg (1981) commented on the mating behavior of 4. ferruginea, A. maculata (Cole), A. semilu- cida (Bates), A. trixa Curran, and an un- described species (probably 4. idahoensis Steyskal—see Steyskal [1984]) from Idaho. Dodson (1987b) described the mating be- havior of A. trixa in New Mexico. Dodson (1987b) predicted that most species of Aciurina would exhibit a similar mating strategy which he termed the “‘male- searching mating system.” In this system, Tephritidae, Aciuwrina, mating behavior, sexual displays, nuptial gift, pher- males move about the host plant scanning for conspecifics and attempt to copulate with any females encountered. Evidently, court- ship is limited to a few brief wing displays and may not precede attempts at copula- tion. Successful mating is dependent more on a male’s ability to maintain a mounted position on a female (Dodson pers. comm.). Aciurina mexicana (Aczél) occurs in southern Arizona, southern California, and northern Mexico where larvae form stem galls on Baccharis sarothroides Gray (Stey- skal 1984). Contrary to what has been re- ported for other Aciurina species, precop- ulatory behavior in A. mexicana is protracted and involves a number of com- plex sexual displays. This paper describes the mating behavior of A. mexicana. MATERIALS AND METHODS Specimens used in this study were swept from B. sarothroides located 14.3 km SE of Continental, Ariz. (Pima Co.) on 16 Feb. VOLUME 92, NUMBER 1 and | Mar. 1986. Flies were separated by sex and caged, one to several, in 0.3-1 clear plastic cups. Cages were fitted with a cotton wick for water and ventilated by a series of small punctures around the top and bottom. Flies were fed a diet of honey containing a small quantity of nutritional yeast. The honey/yeast mixture was provided ad /ib on a paper strip suspended in the cage; water was supplied twice daily by saturating the cotton wick. Flies were kept in the labora- tory at ambient temperature, relative hu- midity, and photoperiod. Observations of mating behavior began by placing a pair of flies ina 100 x 15 mm plastic petri dish. Trial length varied with the flies’ activities. Trials were discontinued when one or both flies became unresponsive or agonistic. When copulation occurred, the pair was observed at least until they uncou- pled. Petri dishes were changed between trials. Observations were made between 1025 and 1627 hours (MST) on nine dates between 25 Feb. and 10 Mar. 1986. Dura- tion of mating activities was recorded to the nearest minute. Video recordings were used to help analyze behaviors. RESULTS Courtship and agonistic displays.—Sev- eral displays were typical of one or both sexes of 4. mexicana during courtship and agonistic interactions. To facilitate discus- sion, these displays are named and de- scribed below. Rotation of the wing refers to twisting the wing so its ventral surface is brought into an anteriorly directed position and the costal margin is pointing upward (Fig. 1). Rotating the wing 90° results in the wing blade being more or less perpendicular to the substrate. Angular measurements of wing movements are visual approxima- tions. Slow signal: One wing is brought slowly forward to an angle of ca. 90° to the long axis of the body (Fig. 2). As the wing is moved forward the wing blade is rotated ca. 90°. The forward movement of the wing may 67 be smooth or by intermittent jerks accom- panied by slight rotational adjustments. The wing is then slowly returned to its original position and the other wing brought forward in a similar manner. Both sexes exhibited this display. Wing fanning: Both wings are held out- stretched at an angle of ca. 45° to the long axis of the body, with the wing blades ro- tated between ca. 45° and 90° (Fig. 3). Both wings are then brought forward and re- turned in short, very rapid, coupled strokes. Bouts of wing fanning were brief (ca. 1 s or less) and sometimes occurred in rapid succession. Only males exhibited this be- havior. Wing flicking: Both wings are held out- stretched at an angle of ca. 45° to the long axis of the body, with the wing blades ro- tated ca. 90° (Fig. 4). Both wings are then brought forward and returned in short, quick strokes, with a brief pause between strokes. Only courting males used this display. Wing thrust: Both wings are simulta- neously and quickly brought forward to an angle of ca. 90° to the long axis of the body; wings are rotated ca. 90° as they are brought forward (Fig. 5). Wings are held in this po- sition as the fly charges toward an intruder. Wing thrusts may be accompanied by wav- ing the forelegs in an aggressive manner (“sparring”), with or without making con- tact. Wing thrusts and sparring were ob- served for both sexes. Wing waving: Wings are alternately brought forward to an angle of ca. 90° to the long axis of the body; the wing is rotated ca. 90° as it is brought forward (Fig. 6). Each wing is returned to its initial position over the back of the fly as the other wing is brought forward. Wing waving is similar to the slow signal, but the wings are brought forward and returned in rapid succession. Wing waving was the basic component of female courtship. Abdominal inflation: The abdominal pleura of males become swollen during courtship and agonistic intrasexual dis- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON b b a ~ a meres nes LAN = See 2 Shh aa \ an \ ys SW TSS \ ; lee wing fanning 3 4a = / SS wing flicking sss R { ) of / = \ geal es \ 2G spe) Tie St ae, \ wing thrust wing waving 5 6 Figs. 1-6. 1. Wing rotation in Aciurina mexicana (left anterolateral view). The wing blade goes from parallel to the substrate to perpendicular to the substrate (rotation = 90°). a, wing apex; b, wing base; c, costal margin. Figs. 2-6. Wing displays in Aciurina mexicana (dorsal view). 2, Slow signal. 3, Wing fanning. 4, Wing flicking. 5, Wing thrust. 6, Wing waving. See text. VOLUME 92, NUMBER | plays. Inflation of the pleura is accompanied by the release of an odor, presumably a pheromone, that is easily detectable by hu- mans. Nuptial gift: During courtship males pro- duce a clear fluid from their mouthparts and dab it onto the substrate with their labellum. Females are attracted to and feed on the fluid while males attempt mounting. The fluid becomes sticky as it dries. Nuptial gifts that were not entirely consumed were readi- ly eaten by both sexes. Mating behavior.—A total of 33 trials were conducted and 20.9 h of observations recorded. Courtship displays were observed for either one or both sexes during 26 trials: only males displayed in 11 trials, only fe- males displayed in two trials, and both sexes displayed during 13 trials. No courtship was observed during seven trials. Copulation (n = 7) occurred only in trials where both sexes exhibited courtship displays. Males initi- ated courtship in 19 trials with two of these resulting in copulation; females initiated in three trials, one resulted in copulation; both flies began courting simultaneously during four trials, all resulted in copulation. Trials in which courtship did not lead to copula- tion (n = 19) were discontinued when one or both flies were unresponsive (n = 5) or agonistic (n = 14). After being placed in a petri dish, flies walked about randomly, often with their wings outstretched, or groomed until one or both became cognizant of the other. Both sexes rhythmically extended and retracted their mouth parts during mating as well as nonmating activities. Courtship and mating were not limited to a particular surface with- in the dish. The most typical and complete sequence of mating behaviors from courtship through copulation is given below. Deviations from this pattern are then discussed. The body of males during noncourtship activities was held close to the substrate. During courtship, however, males extended 69 their legs and held their bodies well above the substrate. In this raised posture males rocked from side to side and displayed wing flicking and abdominal inflation (n = 24). While displaying, males remained station- ary or moved forward by short, uncoupled steps. Males typically approached females from the front. When within a few centi- meters of the female, the male turned, moved a short distance away, turned back to face the female, and reapproached. This sequence was repeated several times prior to producing a nuptial gift. Females responded to courting males by wing waving (n = 13). The intensity of wing waving was affected by the male’s proxim- ity. As the male moved away from the fe- male, wing waving was less vigorous or ceased; as he reapproached, wing waving became more vigorous or was resumed. After a period of reciprocal displays, the male, while continuing to display, became stationary and produced a nuptial gift (n = 7). Although males usually continued to court females that became unresponsive (n = 2) or agonistic (n = 4), nuptial gifts were produced only after vigorous wing waving by females. While continuing her wing wav- ing, the female approached the male and began to feed on the nuptial gift. As the female fed, her wing waving became very rapid. Males displayed for a mean of 8.7 min (n = 7, median = 9 min, range = 2-17 min) before producing a nuptial gift. One male placed his gift on the side of the petri dish, three males placed their gifts on the dish bottom, and three placed a gift on the dish lid. The male ceased displaying soon after the female began feeding on the nuptial gift. He then retreated a short distance and reap- proached the female to attempt mounting. As the male moved away the female ceased to display but continued feeding. As he reapproached the female, his body was close to the substrate and his wings were folded tightly over his abdomen. Males usually 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mounted from the side or rear by simply grabbing the female, or by pushing their head under one of the female’s wings and climb- ing upon her abdomen. Females usually continued feeding and remained stationary while the male attempted to mount. Males probed the tip of the female’s ovi- scape with their surstyli after mounting. Prior to intromission the male grasped the female tightly by hooking his foretarsal claws over the anterolateral margin of her synter- gum 1+2; his meso- and metatarsi re- mained on the substrate. The male main- tained a grip on the female’s syntergum after intromission; his mesotarsi were held loose- ly about her fifth or sixth tergum, and his metatarsi were held loosely about her ovis- cape or trailing behind, touching the sub- strate or not. The male’s abdomen returned to its precourtship size after mounting. After coupling (n = 7), a pair usually moved a short distance before becoming stationary and usually remained in one place if undisturbed. The male’s wings were held over his abdomen during copulation; the female’s wings were spread to accommo- date his body. Copulation lasted a mean of 1.5 h (n = 6, range = 1.1-1.9 h). (Mean duration of copulation does not include one pair that became uncoupled when inadver- tently disturbed.) Males remained motion- less during copulation except for adjusting their posture when disturbed by intermit- tent grooming by the female. Females also were seen to slow signal during copulation. Uncoupling was brief and seemingly without difficulty. Both sexes spent a short period grooming themselves after uncou- pling. Males usually reinitiated courtship, but females either were unresponsive or ac- tively rejected them. No male was seen to remount a female following copulation. Several deviations from the pattern of courtship described above were observed. Males did not produce a nuptial gift in two of the seven trials that resulted in copula- tion. Duration of copulation for these two pairs was 1.4 and 1.5 h. During two trials where gifts were produced and the males were unsuccessful in their first attempts to mount, they reinitiated wing flicking and added more fluid to the gifts. One male added to the gift twice, the other six times; copulation followed in each case. In two other trials the female elicited and fed on a nuptial gift, but then vigorously evaded at- tempts by the male to mount. In four trials, males showing limited or no displays at- tempted to mount females; none of these resulted in copulation. During two trials fe- males exhibited wing waving without pre- vious courting by males. In one of these trials the male was unresponsive; in the oth- er, the male attempted mounting and then exhibited agonistic behavior (wing thrusts) during the remainder of the trial (ca. 6 min). Agonistic behavior.— Wing thrusts were used in an aggressive manner at close dis- tances, inter- or intrasexually. Wing thrusts or sparring, or both, were often used by fe- males to reject courting males (n = 11). Wing fanning accompanied by abdominal infla- tion and odor production was commonly observed between males in the same con- tainer. Wing fanning was observed between a male and female only once. Females evaded a male’s attempt to mount by simply walking or running away. While moving away, females sometimes held or flicked their wings over their ab- domen. Females prevented mounted males from copulating by either kicking with their hind legs or pushing the tip of their oviscape to the substrate, or both. Females did this while stationary or while dragging the male. DISCUSSION Courtship and agonistic displays.—A. mexicana appears to share a number of wing displays with other Aciurina species as well as other fruit flies in general. Tauber and Tauber (1967) described wing movements of A. ferruginea that are similar to the slow signal of A. mexicana. They suggested that the display functioned in intraspecific rec- ognition, courtship, and copulation. This VOLUME 92, NUMBER | display also was reported for 4. ferruginea by Wangberg (1981). Dodson (1987b) de- scribed similar wing movements for 4. frixa. The slow signal of 4. mexicana occurred when flies were together or alone and could not be associated with any specific activity. Slow signals were often seen while flies, re- gardless of sex, faced each other at a short distance. This suggests that slow signals may in part operate in conspecific recognition. A similar display occurs in a number of fruit flies (e.g. Nation 1972, Piper 1976, Cav- ender and Goeden 1982, 1984). Displays like wing fanning and wing flick- ing may also occur in 4. bigeloviae (Cock- erell) and 4. trixa(G. Dodson pers. comm.). “Rowing” of the wings described for A. trixa (Dodson 1987b) is similar to wing fanning and wing flicking described here (G. Dodson pers. comm.). Wing fanning by 4. mexicana may help direct male odor toward intruders. Dis- persal of a sex pheromone by a similar be- havior has been suggested for Anastrepha suspensa (Loew) and Ceratitis capitata (Wiedemann) (Nation 1972, Prokopy and Hendrichs 1979). Alternatively, Sivinski et al. (1984) convincingly demonstrated that wing fanning by males of 4. suspensa pro- duces sounds that are sexually important, intraspecific signals. No sound was noted during wing fanning by Aciurina mexicana. Wing flicking also may help to disperse and direct male odor. Prior to and inter- mittently during courtship, males rubbed their hind legs against their abdominal pleu- ra and in turn rubbed their wings. This may transfer odor from the pleura to the wings where it then could be dispersed by wing flicking. The activity may be coincidental with grooming. Male Anastrepha suspensa exhibit a similar behavior, and Nation (1972) noted that “this cleaning behavior may spread a sex attractant over the body and wings and provide a greater surface from which it can evaporate.” Piper (1976) sug- gested that wing movements in general may help direct pheromones toward either sex. vA Copulating males of Aciurina ferruginea may “‘flick”’ their wings at approaching males (Tauber and Tauber 1967). In the context of agonistic behavior, this display is similar to the wing thrusts of 4. mexicana. Sparring, but not wing thrusts, has been reported for A. trixa (as “grappling,” Dodson [1987b]). A display resembling wing thrusts also has been reported for Trupanea bisetosa (Co- quillett) and Paracantha cultaris (Coquil- lett) (Cavender and Goeden 1982, 1984). In these flies, however, the display functions in courtship. Wing displays comparable to wing wav- ing have been reported for A. ferruginea and A. trixa (Tauber and Tauber 1967, Dodson 1987b) as well as other fruit flies (e.g. Tau- ber and Toschi 1965, Cavender and Goeden 1982, Dodson 1987b). Dodson (1987b) ob- served 11 virgin female 4. frixa to mate after frequently waving their wings (‘“‘ad- vertising behavior’). Conversely, none of nine once-mated females exhibited the dis- play (Dodson 1987b). Tauber and Toschi (1965) reported that during the courtship of Euleia fratria (Loew), the frequency of “wing waving” and displacement of the wings are indicative of the level of sexual excitation of females. As noted above, frequency and duration of wing waving by 4. mexicana females was affected by the proximity of a courting male, and males produced nuptial gifts only in the presence of a displaying female. Although fruit fly wing movements usu- ally are discussed in terms of intraspecific displays, there also is strong evidence for an interspecific role. It was recently shown that wing movement and wing pattern of Rhagoletis zephyria Snow and Zonosemata vittigera (Coquillett) are important in de- terring predation by mimicking the flies’ salticid spider predators (Mather and Roit- berg 1987, Greene et al. 1987). Further, these authors suggested that spider mimicry may be widespread in the Tephritidae. Abdominal inflation and odor produc- tion have not been reported for other Aci- 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON urina species. Tauber and Tauber (1967) refer to ‘“‘pumping of the abdomen” in male homosexual encounters of 4. ferruginea, but it is unclear whether this represents abdom- inal inflation as discussed here. Abdominal inflation accompanied by odor production has been observed in males of the fruit flies Dirioxa (= Rioxa) pornia (Walker), Anas- trepha ludens (Loew), A. suspensa, Trupa- nea bisetosa, and Toxotrypana curvicauda Gerstacker (Pritchard 1967, Nation 1972, Cavender and Goeden 1982, Landolt et al. 1985, Robacker and Hart 1985b). Appear- ance of the inflated pleura of Aciurina mex- icana was quite similar to that illustrated for T. bisetosa(Cavender and Goeden 1982, Fig. 2). Glandular epidermal cells have been identified in the pleura of abdominal seg- ments 3, 4, and 5 of males of D. pornia (Pritchard 1967), seven Anastrepha species, and two Ceratitis species (Nation 1981). The cells are sex-specific, occurring as a thick band in male pleura; female pleural epider- mis is uniformly thin and undifferentiated (Pritchard 1967, Nation 1981). These glan- dular cells are, at least in part, the probable source of male odor in D. pornia, A. sus- pensa, and A. ludens (see Pritchard 1967, Nation 1974, 1981; Robacker and Hart 1985a). Pleura removed from male Aciurina mexicana were visibly thicker than those removed from females when compared un- der a dissecting microscope, regardless of the size of the fly. Based on the above ob- servations, and because odor was detectable only when the pleura were distended, it is likely that the pleura of male 4. mexicana contain glandular cells that are associated with the male odor. Odors produced by male D. pornia, Anas- trepha suspensa, A. ludens, and Toxotry- pana curvicauda are attractive to females (Pritchard 1967, Nation 1983, Landolt et al. 1985, Robacker and Hart 1985a). The odor produced by 7rupanea bisetosa is pre- sumably also a sex pheromone (Cavender and Goeden 1982). Courting males of Aci- urina mexicana always produced odor, but it was not shown that the odor was attractive to females. Because male odor was pro- duced during wing fanning, it may also func- tion intrasexually. The male odor of A. mexicana is distinc- tive, but difficult to describe precisely. The male odor of 7. bisetosa was reported as a “yeasty or musty smell” (Cavender and Goeden 1982). The odor of 4. mexicana is likewise yeasty or musty. Nuptial feeding has not been reported for other Aciurina species. Dodson (1987b) re- ported that adult feeding in general is ‘“‘neg- ligible” for A. trixa. He also noted that no flowers or “obvious exudates” occur on the host plant (Chrysothamnus nauseosus [Pal- las] Britton) when adults are present. In con- trast, flies used in the present study often became so replete with the honey/yeast mixture that ordinary movement, let alone mating, was quite limited. Moreover, stems and leaves of B. sarothroides are coated with a sticky material that is attractive to many adult insects (Meyer et al. 1979) and on which adult 4. mexicana may feed. Adults of A. ferruginea also feed (Tauber and Taub- er 1967). Nuptial feeding has been observed (Freid- berg 1981 and references therein) or sus- pected (Cavender and Goeden 1984) for a number of fruit flies. In five species, nuptial gifts consist of an erect, white, frothy mass produced from the mouthparts of males and deposited onto the host plant (Freidberg 1981 and references therein). In two of these species nuptial gifts originate in large, sex- ually dimorphic salivary glands (Pritchard 1967, Freidberg 1981). In Spathulina sicula Rondani, a species with postcopulatory trophallaxis, the salivary glands of males are much larger than those of females (Freid- berg 1982 [species as tristis (Loew)], Fig. 5). Sexually dimorphic salivary glands also are present in males of Anastrepha and Ceratitis (Nation 1981), but, at least for 4. suspensa, A. ludens, and C. capitata, nuptial feeding has not been reported (Nation 1972, Pro- VOLUME 92, NUMBER 1 kopy and Hendrichs 1979, Robacker and Hart 1985b). Preliminary examination of the alimentary tract of five male and five female A. mexicana showed essentially no intersexual difference in size and shape of the salivary glands. Both sexes possess a rel- atively large crop. Unlike the nuptial gifts discussed above, that of A. mexicana consisted of drops of a clear fluid. When initially produced it was similar to the fluid in the crops of dissected flies. The amount of fluid produced was greater than would be expected from the salivary glands alone. It seems probable then, that contents of the crop contributed to the nuptial gift of 4. mexicana. Mating behavior.— The courtship posture assumed by male 4. mexicana has not been reported for other Aciurina species. The posture is similar to that illustrated for D. pornia (Pritchard 1967, fig. 2). An erect pos- ture also has been observed for Anastrepha ludens (Robacker and Hart 1985b). Mounting and copulatory positions of 4. mexicana were like those described for 4. ferruginea (Tauber and Tauber 1967). How- ever, once mounted, males did not contact the female’s dorsum with their mouth parts as in A. ferruginea. Mounting in A. trixa is facilitated by the male grasping the female’s hindlegs between the tibia and femur of his forelegs (termed “‘leglock,’’ Dodson [1987a, b]). The forefemora of males are enlarged and males with larger forefemora are more successful in securing copulations (Dodson 1987a). This method of mounting was not seen for 4. mexicana and subsequent mea- surements indicated no significant sexual dimorphism in forefemora (Table 1). Dodson (1987b) reported multiple mat- ings by female A. trixa. Females of A. mex- icana may also be polyandrous. On one oc- casion a female mated again after | d, while on another occasion a female mated again after 1 wk. Conversely, in two trials initiated 2 d after copulation, one female was unre- sponsive to and the other actively rejected male courtship. Mean duration of copula- 7S) Table 1. Comparison of mean maximum length and width of forefemora among male and female Aciurina mexicana (two-tailed t-test). Measurements taken with ocular micrometer and dissecting microscope. Means expressed in arbitrary units (35 units = 1 mm). X (n) t df iB Femur length Males 26.9 (11) 0.37 20 >0.25 Females 26.6 (11) Femur width Males 8.4 (11) 0.93 20 >O0115 Females 8.1 (11) tion for 4. trixa (2.2 h [Dodson 1987b]) was longer than that observed for 4. mexicana (1.5 h). Evasive actions taken by female 4. mex- icana to prevent copulation are like those described for A. ferruginea (Tauber and Tauber 1967) and 4. trixa (Dodson 1987b). Females of Anastrepha suspensa similarly press their oviscape to the substrate to pre- vent copulation (Nation 1972). It is unknown to what extent the mating behaviors observed in the laboratory for Aciurina mexicana occur in nature. Obser- vation of behavior in the field was unsuc- cessful because of low fly densities. How- ever, it is very unlikely that the complex behaviors reported here were laboratory ar- tifacts. Mating behavior and systematics. —Aci- urina mexicana, A. aplopappi (Coquillett), and 4. thoracica Curran form Steyskal’s (1984) Aplopappi species group. Unlike oth- er Aciurina species, members of the Aplo- pappi group have host plants other than species of Chrysothamnus, and lack surface specializations on the membranous portion of sternum 8 of the ovipositor (Steyskal 1984). The observations reported herein raise the possibility that members of the Aplopappi group also have distinctive mat- ing behaviors that further distinguish them from the remainder of the genus. Eldredge and Cracraft (1980) noted that 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON “with the exception of the findings of com- parative anatomy, no other kind of simi- larity has been utilized by systematists as much as that of behavior.’”’ Current super- generic classifications of the Tephritidae (e.g. Foote and Steyskal 1987), which are based largely on adult morphology, are problem- atical. It is now well documented that many fruit flies have characteristic mating behav- iors. As shown here, sexual displays are a potentially rich (but unused) source of com- parative data. The interspecific distribution of these displays may help resolve or cor- roborate evolutionary patterns used to con- struct classifications. ACKNOWLEDGMENTS Adult A. mexicana were collected with the help of J. M. Sirota. N. Buck and J. M. Sirota assisted in making video recordings. Earlier drafts of the manuscript were re- viewed by T. Burk, G. Dodson, R. D. Goe- den, G. Henderson, W. L. Nutting, C. A. Olson, J. M. Sirota, and an anonymous re- viewer. Their assistance is gratefully ac- knowledged. Arizona Agricultural Experi- ment Station Manuscript No. 7050. LITERATURE CITED Cavender, G. L.and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Ann. Entomol. Soc. Am. 75: 400-406. 1984. The life history of Paracantha cultaris (Coquillett) on wild sunflower, Helianthus annuus L. ssp. lenticularis (Douglas) Cockerell, in south- ern California (Diptera: Tephritidae). Pan-Pac. Entomol. 60: 213-218. Dodson, G. 1987a. The significance of sexual di- morphism in the mating system of two species of tephritid flies (Aciurina trixa and Valentibulla dodsoni) (Diptera: Tephritidae). Can. J. Zool. 65: 194-198. 1987b. Biological observations on Aciurina trixa and Valentibulla dodsoni (Diptera: Tephrit- idae) in New Mexico. Ann. Entomol. Soc. Am. 80: 494-500. Dodson, G. and S. B. George. 1986. Examination of two morphs of gall-forming Aciurina (Diptera: Te- phritidae): Ecological and genetic evidence for species. Biol. J. Linn. Soc. 29: 63-79. Eldredge, N. and J. Cracraft. 1980. Phylogenetic pat- terns and the evolutionary process. Columbia University Press, New York. Foote, R. H. and G. C. Steyskal. 1987. Tephritidae, pp. 817-831. Jn McAlpine, J. F., ed., Manual of Nearctic Diptera, Vol. 2. Agriculture Canada, Monograph No. 28, Ottawa. Freidberg, A. 1981. Mating behaviour of Schistopte- rum moebiusi Becker (Diptera: Tephritidae). Israel J. Entomol. 15: 89-95. 1982. Courtship and post-mating behaviour of the fleabane gall fly, Spathulina tristis (Diptera: Tephritidae). Entomologia Generalis 7: 273-285. Greene, E., L. J. Orsak, and D. W. Whitman. 1987. A tephritid fly mimics the territorial displays of its jumping spider predators. Science 236: 310- 312. Landolt, P. J., R. R. Heath, and J. R. King. 1985. Behavioral responses of female papaya fruit flies, Toxotrypana curvicauda (Diptera: Tephritidae), to male-produced sex pheromone. Ann. Entomol. Soc. Am. 78: 751-755. Mather, M. H. and B. D. Roitberg. 1987. A sheep in wolf's clothing: Tephritid flies mimic spider pred- ators. Science 236: 308-310. Meyer, R. P., F. G. Zalom, T. L. McKenzie, and P. H. Mason. 1979. Notes on insects associated with desert broom (Baccharis sarothroides Gray) (Com- positae) in southeastern Arizona. Southwestern Naturalist 24: 603-612. Nation, J. L. 1972. Courtship behavior and evidence for a sex attractant in the male Caribbean fruit fly, Anastrepha suspensa. Ann. Entomol. Soc. Am. 65: 1364-1367. . 1974. The structure and development of two sex specific glands in male Caribbean fruit flies. Ann. Entomol. Soc. Am. 67: 731-734. 1981. Sex-specific glands in tephritid fruit flies of the genera Anastrepha, Ceratitis, Dacus and Rhagoletis (Diptera: Tephritidae). Int. J. Insect Morphol. and Embryol. 10: 121-129. . 1983. Sex pheromone of the Caribbean fruit fly: Chemistry and field ecology. Proc. Sth Int. Conf. Pest. Chem. 2: 109-110. Piper, G. L. 1976. Bionomics of Euvarestoides acu- tangulus (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 69: 381-386. Pritchard, G. 1967. Laboratory observations on the mating behaviour of the island fruit fly Rioxa por- nia (Diptera: Tephritidae). J. Aust. Entomol. Soc. 6: 127-132. Prokopy, R. J. and J. Hendrichs. 1979. Mating be- havior of Ceratitis capitata on a field-caged host tree. Ann. Entomol. Soc. Am. 72: 642-648. Robacker, D. C. and W. G. Hart. 1985a. (Z)-3-no- nenol, (Z,Z)-3,6-nonadienol and (S,S)-(-)-epi- anastrephin: Male produced pheromones of the VOLUME 92, NUMBER 1 Mexican fruit fly. Entomol. Exp. Appl. 39: 103- 108. 1985b. Courtship and territoriality of labo- ratory-reared Mexican fruit flies, dnastrepha lu- dens (Diptera: Tephritidae), in cages containing host and nonhost trees. Ann. Entomol. Soc. Am. 78: 488-494. Sivinski, J., T. Burk, and J.C. Webb. 1984. Acoustic courtship signals in the Caribbean fruit fly, Anas- trepha suspensa (Loew). Anim. Behav. 32: 1011- 1016. Steyskal, G.C. 1984. A synoptic revision of the genus Aciurina Curran, 1932 (Diptera, Tephritidae). Proc. Entomol. Soc. Wash. 86: 582-598. 75 Tauber, M. J.and C. A. Tauber. 1967. Reproductive behavior and biology of the gall-former Aciurina ferruginea (Doane) (Diptera: Tephritidae). Can. J. Zool. 45: 907-913. Tauber, M. J. and C. A. Toschi. 1965. Bionomics of Euleia fratria (Loew) (Diptera: Tephritidae). I. Life history and mating behavior. Can. J. Zool. 43: 369-379. Wangberg, J. K. 1981. Gall-forming habits of Aci- urina species (Diptera: Tephritidae) on rabbit- brush (Compositae: Chrysothamnus spp.) in Ida- ho. J. Kans. Entomol. Soc. 54: 711-732. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 76-85 NOMENCLATURE OF SOME NEOTROPICAL GELECHIIDAE (LEPIDOPTERA) RONALD W. HODGES AND VITOR OSMAR BECKER (RWH) Research Entomologist, Systematic Entomology Laboratory, Agriculture Re- search Service, USDA; % National Museum of Natural History, MRC-168, Washington, D.C. 20560; (VOB) Ing, EMBRAPA-CPAC, P.O. Box 70-0023, 73300—Planaltina, D.F., Brazil. Abstract. —Five generic and six specific synonymies are established, and 12 new com- binations are made. Neotypes are designated for four names and a lectotype for one name. Symmetrischema tangolias (Gyen) is established as the valid name for S. plaesiosema (Turner), a pest of potato tubers. Key Words: Many species and genera of neotropical Gelechiidae have remained unknown since their publication. Meyrick (1925) treated all taxa proposed to that date and made many nomenclatural decisions. In many instances he did so on the basis of written descrip- tions; he did not see or make an effort to obtain type specimens beyond those in his collection. Clarke (1969a, b) illustrated the adults and genitalia of type specimens of species of Gelechiidae described by Meyrick and held by the British Museum (Natural History). His work is immensely helpful to gain preliminary, and sometimes final, un- derstanding of a large number of species. Becker (1984) relied mainly on literature, original descriptions and revisions, to as- sociate species with genera and to place gen- era in higher taxa. Because characters nec- essary to define species and genera in the Gelechioidea often are in the male and/or female genitalia and the number of taxa in the Neotropical Region is very large, many described taxa are unrecognized, sometimes at the family level. This paper, based on study of several type specimens, clarifies nomenclature, synonymy, gelechiinae, gelechiid moths, Neotropical Region knowledge of nine generic names and 18 specific names. Kieffer and Jorgensen (1910) published on plant galls, the primary gall makers, and parasites reared from the gall makers that had been observed and collected in Argen- tina (primarily in the province of Mendoza). Nearly all the insects were described as “new species” or as “new genus, new species” combinations. Eight of the insects were Lep- idoptera, and four were Gelechiidae. The gelechiid adults were sent to Embrik Strand for description (Strand 1911). He provided manuscript names for the moths to Kieffer and J6rgensen, and they attributed the names to Strand in the text of their paper (Kieffer and Jorgensen 1910). However, they presented adequate information about each species to validate the names and thus be- came the authors of the Strand names. Be- cause they were not intentionally describing new species and/or genera, they may not have had adults in their possession nor la- belled specimens as types. Their descrip- tions are limited to galls and to larvae or pupae when present; no mention is made of VOLUME 92, NUMBER 1 adult characters. It appears that no type ma- terial of these four species is extant in Ar- gentina or elsewhere. The homonymous names published by Strand are supported by specimens provided by Kieffer and are highly suitable to serve as material from which neotypes can be designated. Type material of the Strand names is well pre- served in the Museum fiir Naturkunde der Humboldt-Universitat, Berlin, East Ger- many. We borrowed the types of the Strand names, dissected the abdomens, and endea- vored to relate their identities to other taxa. The results follow. — . Gnorimoschema (Tuta) atriplicella Kief- fer & J6rgensen 1910: 363. (Figs. 1, 6). The neotype male, present designation, bears the following labels: 1) Argentina/ Mendoza/Kieffer G. 2) male genitalia/ slide 5109/R W Hodges. 3) Neotype by Hodges & Becker 1989. Gnorimoschema (Tuta) atriplicella isa valid species of Phthorimaea Meyrick, NEW COMBINATION; and it is the type species of Tuta Kieffer & Jorgensen. Thus, 7uta is a junior synonym of Phthorimaea, NEW SYNONYMY; and it is removed from the synonymy of Gnorimoschema Busck where Meyrick (1925: 89) had placed it. The moth (Fig. 1) is indistinct, as are most Phthori- maea, and has pale yellow-brown forewings and pale straw-yellow hindwings. The male genitalia are as illustrated (Fig. 6). 2. Tecia mendozella Kieffer & Jorgensen 1910: 375. (Figs. 2, 7). The neotype male, present designation, bears the following labels: 1) Argentina/ Mendoza/Kieffer G. 2) male genitalia/ slide 5107/R W Hodges. 3) Neotype by Hodges & Becker 1989. Tecia mendozella is a junior synonym of Topeutis venosa Butler, NEW SYNONY- MY; and it is the type species of Tecia Kief- fer & Jérgensen. Topeutis venosa Butler is the type species of Orsotricha Meyrick, 77 which thus becomes a junior synonym of Tecia, NEW SYNONYMY; and Tecia ve- nosa (Butler) is a NEW COMBINATION. 3. Fapua albinervella Kieffer & Jorgensen 1910: 378. (Figs. 3, 8). The neotype male, present designation, bears the following labels: 1) Argentinien/ Prov. Mendoza/Kieffer G. 2) male geni- talia/slide 5108/R W Hodges. 3) Neotype by Hodges & Becker 1989. Fapua albinervella is a valid species of Te- cia, and it is the type species of Fapua Kief- fer & JOrgensen. Thus, we confirm Mey- rick’s (1925: 89) placement of the species and treatment of Fapua as a junior synonym of Tecia. The wing pattern of the moth (Fig. 3) superficially resembles that of a species of Coleophora Hiibner (Coleophoridae). The forewings are pale yellow orange with white on the veins; the hindwings are pale yellow with a pale-orange fringe. 4. Tecia (Lata) kiefferi Kieffer & Jorgensen 1910: 398. (Figs. 4, 9). The neotype male, present designation, bears the following labels: 1) Argentinien/ Prov. Mendoza/Kieffer G. 2) male geni- talia/slide 5110/R W Hodges. 3) Neotype by Hodges & Becker 1989. Tecia (Lata) kiefferi is a valid species of Tecia, and it is the type species of Lata Kief- fer & Jérgensen. Thus, we confirm Mey- rick’s (1925: 89) placement of the species and treatment of Lata as a junior synonym of Tecia. The moth (Fig. 4) is similar to Tecia venosa (Butler); however, in the male genitalia (Fig. 9) the anteromesial margin of the tegumen is straight in kiefferi; it is rounded in venosa (Fig. 2). Another hitherto unrecognized, mono- typic genus is Brachypsaltis Meyrick with type species subalbata Meyrick. The holo- type male of subalbata (Fig. 5) was bor- rowed from the Naturhistorisches Museum, Vienna, and the genitalia were dissected. B. subalbata proves to be a valid species of 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S) Figs. 1-5. Wings of Gelechiidae species. 1, Phthorimaea atriplicella (Kieffer & Jorgensen), neotype, m, Argentina. 2, Tecia venosa (Butler), [neotype of Tecia mendozella (Kieffer & Jorgensen)], m, Argentina. 3, Tecia albinervella (Kieffer & Jorgensen), neotype, m, Argentina. 4, Tecia kiefferi (Kieffer & Jorgensen), neotype, m, Argentina. 5, Tecia subalbata (Meyrick), holotype, m, Argentina. VOLUME 92, NUMBER | 19 i 1 { j ‘ : Fig. 6. Phthorimaea atriplicella (Kieffer & Jérgensen), neotype, m, Argentina. a, genitalia with aedeagus removed (ventral aspect). b, aedeagus. Tecia and is transferred to that genus as Tecia subalbata (Meyrick), NEW COMBI- NATION. Thus, Brachypsaltis becomes a junior synonym of Tecia, NEW SYNON- YMY. The male genitalia are as illustrated (Fig. 10). Becker borrowed eight syntypes of Hol- cocera baccharisella Bréthes from the Mu- seo Argentino de Ciencias Naturalis “Ber- nardino Rivadavia,” Buenos Aires. Bréthes did not indicate the provenance nor the number of specimens he had when he de- scribed baccharisella. The syntypes bear the same labels: 1) Bs Aires/iv.1916/J.B. 2) Holcocera baccharisella Br. We designate a male as lectotype and have added a third 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Tecia venosa (Butler), [neotype of Tecia mendozella (Kieffer & Jorgensen)], m, Argentina. a, genitalia with aedeagus removed (ventral aspect). b, aedeagus. label, Lectotype by Hodges & Becker 1989, ler, NEW SYNONYM Y. It is thus trans- to it. Paralectotype labels were added to the ferred to Tecia as Tecia baccharisella remaining seven specimens. Study of the (Bréthes), NEW COMBINATION. male genitalia of baccharisella shows it to Povolny (1980) described Scrobipalpop- be ajunior synonym of Topeutis venosa But- _ sis (Scrobischema) vergarai on the basis of VOLUME 92, NUMBER | 81 Fig. 8. (ventral aspect). b, aedeagus. adults reared from larvae that caused “... hyperplastic deformation of the terminal shoots of Baccharis macrantha HBK.” Te- cia venosa is a gall maker on Baccharis ser- rulata Pers. (Kieffer & Jorgensen 1910: 375). Study of Povolny’s illustrations and discus- sion convince us that S. vergarai is a junior Tecia albinervella (Kieffer & J6rgensen), neotype, m, Argentina. a, genitalia with aedeagus removed synonym of 7. venosa, NEW SYNONY- MY. Further, because vergarai is type species of Scrobischema Povolny, the latter is a ju- nior synonym of Tecia, NEW SYNONY- MY; and Tecia vergarai Povolny is a NEW COMBINATION. Study of the male and female genitalia of 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 9. aspect). b, aedeagus. Gelechia petasitis Pfaffenzeller, the type species of Scrobipalpopsis Povolny, lead us to the conclusion that Scrobipalpopsis is a junior synonym of Tecia, NEW SYNON- YMY, and that Gnorimoschema tetradymi- ella Busck, Gnorimoschema petrella Busck, Gnorimoschema arnicella Clarke, ‘‘Scrobi- palpopsis” chili Povolny, and Scrobipalpop- sis solanivora Povolny are NEW COMBI- NATIONS in Tecia. Hodges (1983: 22) erred in making Scro- Tecia kiefferi (Kieffer & Jorgensen), neotype, m, Argentina. a, genitalia with aedeagus removed (ventral bipalpopsis a junior synonym of Ptycerata Ely. He used a very similar, but incorrectly identified, specimen for his concept of Pty- cerata busckella Ely, the type species of Pty- cerata. The genitalia of the holotype of Pty- cerata busckella Ely show that it is related to Monochroa Heinemann and Isophrictis Meyrick, not to Gnorimoschema Busck and allies. Scrobipalpa Janse, with type species Ge- lechia heliopa Lower, is also very similar to VOLUME 92, NUMBER 1 83 Fig. 10. Tecia subalbata (Meyrick), holotype, m, Argentina. a, genitalia with aedeagus removed (ventral aspect). b, aedeagus. Tecia. We do not propose to synonymize Scrobipalpa with Tecia at this time; how- ever, we draw attention to the similarity. Trichotaphe tangolias Gyen has remained in Trichotaphe Clemens (Becker 1984: 51), which is a junior synonym of Dichomeris Hiibner (Hodges 1986: 10), because no type material has been recognized subsequent to the original description. In the U.S. Na- tional Museum of Natural History we found a microscope slide with the left wings of Gelechia (Trichotaphe) tangolias Gyen made by Busck in 1916 from a specimen received from Prof. Silva Figueroa of Chile. The fore- wing definitely is that of Symmetrischema plaesiosema (Turner). Gyen’s paper (1913) indicates that he received the specimens on which the description was based from C. Silva Figueroa. A few pages further on, in the same publication, Silva (1913) pub- lished on the life history of 7. tangolias, illustrating the immature stages on potato. Silva (1915) published a short note and il- lustrated the adult, pupa, and larva of 77i- chotaphe tangolias. The illustration of the adult is a crude representation of S. plae- siosema. On the basis of the original de- scription, the pair of wings received from Silva, the host, and the illustration by Silva, we conclude that Trichotaphe tangolias 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Gyen is a senior synonym of Symmetris- chema plaesiosema (Turner), NEW SYN- ONYMY, and is a species of Symmetris- chema Povolny [Symmetrischema tangolias (Gyen), NEW COMBINATION]. NOMENCLATURAL SUMMARY GELECHIIDAE Gelechiunae Phthorimaea Meyrick 1902 Tuta Kieffer & Jorgensen 1910, n. syn. [from Gnorimoschema] atriplicella (Kieffer & J6rgensen 1910), n. comb. [from Gnorimoschema] Symmetrischema Povolny 1967 tangolias (Gyen 1913), n. comb. [from Trichotaphe] plaesiosema (Turner 1919), n. syn. melanoplintha (Meyrick 1926), n. syn. tuberosella (Busck 1931), n. syn. Tecia Kieffer & Jorgensen 1910 Fapua Kieffer & Jorgensen 1910 Lata Kieffer & J6rgensen 1910 Orsotricha Meyrick 1914, n. syn. Brachypsaltis Meyrick 1931, n. syn. Scrobipalpopsis Povolny 1967, n. syn. [from Ptycerata] Scrobischema Povolny 1980, n. syn. albinervella (Kieffer & Jorgensen 1910) arnicella (Clarke 1942), n. comb. [from Ptycerata] chili (Povolny 1967), n. comb. [from Ptycerata] kiefferi (Kieffer & Jorgensen 1910) petasitis (Pfaffenzeller 1867), n. comb. {from Scrobipalpopsis] petrella (Busck 1915), n. comb. [from Ptycerata] solanivora (Povolny 1973), n. comb. [from Ptycerata] subalbata (Meyrick 1931), n. comb. [from Brachypsaltis] tetradymiella (Busck 1903), n. comb. [from Ptycerata] venosa (Butler 1883), n. comb. [from Orsotricha] mendozella (Kieffer & J6rgensen 1910), n. syn. baccharisella (Bréthes 1917), n. syn., n. comb. {from Holcocera, Blastobasidae] vergaral (Povolny 1980), n. syn., n. comb. [from Scrobischema] ACKNOWLEDGMENTS We thank Dr. Axel Bachmann (Museo Argentino de Ciencias Naturalis ‘‘Bernar- dino Rivadavia,” Buenos Aires), Dr. H.-J. Hannemann (Museum fiir Naturkunde der Humboldt-Universitat, Berlin), and Dr. Fritz Kasy (Naturhistorisches Museum, Vi- enna) for allowing us to study type speci- mens in their care; Dr. J.-F. Landry (Bio- systematics Research Centre, Ottawa) and Dr. K. Sattler [British Museum (N.H.), Lon- don] for review of the manuscript; and Mr. Victor Krantz (National Museum of Nat- ural History, Washington) for the photo- graphs of adult moths. The line drawings were done by the junior author. LITERATURE CITED Becker, V. O. 1984. Gelechiidae. /n Heppner, J. B., ed., Atlas of Neotropical Lepidoptera, Checklist: Part 1: 44-53. Bréthes, J. 1917. Description d’une galle et du pa- pillon qui la produit. Physis 3: 449-451, 3 figs. {unnumbered]. Busck, A. 1931. Two new Peruvian microlepidoptera of economic importance (Gelechiidae and Oeco- phoridae). Proc. Ent. Soc. Wash. 33: 59-63, figs. 1-6. Clarke, J. F.G. 1969a. Catalogue of the type speci- mens of microlepidoptera in the British Museum (Natural History) described by Edward Meyrick, 6: 219-537, pl. 109-267. Trustees of the British Museum (Natural History), London. . 1969b. Op. cit. 7: 1-531, pl. 1-265. Gyen, W. J. A. K. van 1913. Descriptions of Chili microlepidoptera. Bol. Mus. Nac. [Chile] 5: 338- 340. Hodges, R. W. 1983. Gelechiidae, pp. 19-25. Jn Hodges, R. W. et al., eds., Check List of the Lep- VOLUME 92, NUMBER | idoptera of America North of Mexico. E. W. Clas- sey Ltd. and Wedge Entomological Research Foundation, London. . 1986. Gelechioidea, Gelechiidae (in part). /n Dominick, R. B. et al., The Moths of America North of Mexico 7.1: i-xii + 1-[196], text fig. 1- 31, mon. pl. A-HH, col. pl. 1-4. Kieffer, J. J. and Jérgensen, P. 1910. Gallen und Gal- lentiere aus Argentinien. Centralblatt fiir Bakteri- ologie, Parasitenkunde u. Infektionskrankheiten 27: 362-444, figs. 1-61. Meyrick, E. 1914. Exotic Microlepidoptera 1(9): 257- 288. 1925. Lepidoptera Heterocera fam. Gele- chiadae. Jn Wytsman, P., ed., Genera Insectorum 184: 1-290, pl. 1-5. . 1926. Exotic Microlepidoptera 3(9): 257-288. 1931. Exotic Microlepidoptera 4(2): 33-64. 85 Povolny, D. 1967. Genitalia of some nearctic and neotropic members of the tribe Gnorimoschemini (Lepidoptera, Gelechiidae). Acta Ent. Mus. Pragae 37: 51-127, figs. 1-131. . 1980. Scrobipalpopsis (Scrobischema subgen. n.) vergarai sp. n., a potential pest species in Co- lombia. Acta Ent. Bohemoslovaca 77: 55-63, figs. 1-10. Silva, F. C. 1913. Un nuevo microlepidoptero chi- leno. Bol. Mus. Nac. [Chile] 5: 349-353, figs. 1M. 1915. Un nuevo microlepidoptero chileno. Rey. Chilena His. Nat. 19: 41-42, fig. 7a-c. Strand, E. 1911. Sechs neue Gelechiidae aus Argen- tinien. Berl. Ent. Zeits. 55: 165-173, figs. 1-11. Turner, A.J. 1919. The Australian Gelechianae (Lep- idoptera). Proc. Roy. Soc. Queensland 31: 108- 172. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 86-92 THREE NEW SPECIES OF MICROCADDISFLIES (TRICHOPTERA: HYDROPTILIDAE) FROM THE OZARK MOUNTAINS, U.S.A. MICHAEL L. MATHIS AND DAviD E. BOWLES (MLM) Department of Zoology, University of Arkansas, Fayetteville, Arkansas 72701; (DEB) Department of Entomology, University of Arkansas, Fayetteville, Arkansas 72701. Abstract.—Three new species of hydroptilid caddisflies are described from the Ozark Mountains of Arkansas and Missouri. Hydroptila artesa n. sp. and H. sandersoni n. sp. are members of the H. tineoides species group and are closely related to H. amoena, H. paramoena, and H. oneili. A key separating these five species is presented. Neotrichia arkansasensis n. sp. is most closely related to N. sonora from the desert Southwest, but is easily distinguished by the shape of the inferior appendages. Key Words: tains, new species The Interior Highlands encompasses mountainous areas of southern Missouri, northern Arkansas, eastern Oklahoma, eastern Kansas, and southwestern Illinois. The region includes the Ozark and Ouachita Mountains and a number of smaller ranges in Oklahoma and Kansas. Much of the area in Arkansas, Oklahoma, and Missouri is un- developed or used for light agricultural pur- poses such as livestock ranching. Streams and springs are abundant and many are un- polluted. The area includes five streams classified as National Scenic Riverways and some of the largest volume springs in North America. Surprisingly, the region is one of the more-poorly studied areas in the United States with regard to its trichopteran fauna. Only Kansas and northwest Arkansas have been sampled intensively (Unzicker et al. 1970, Schuster and Hamilton 1978, Ham- ilton and Schuster 1978, 1979, 1980, Ham- ilton et al. 1983). In order to increase our knowledge of the trichopteran fauna of this microcaddisfly, Hydroptila tineoides group, taxonomic key, Ozark Moun- region, we have initiated surveys in the mountainous areas of Arkansas, Missouri, and Oklahoma. We describe herein three new microcaddisflies collected from the Ozark Mountains. Specimens were collected using a UV-light trap with the exception of a single sample from Mammoth Springs, Arkansas, that we obtained from the Illinois Natural History Survey. Terminology and higher taxonomy follow Marshall (1979). In characterizing some species, we use the ratio of the length of the aedeagus to that of the abdomen. The length of the abdomen was measured dor- sally from the posterior margin of the meta- scutellum to the apex of tergite X. Types and voucher specimens are deposited at the American Museum of Natural History (AMNH), Illinois Natural History Survey (INHS), National Museum of Natural His- tory, Smithsonian Institution (NMNH), and University of Arkansas Insect Collection (UAIC). VOLUME 92, NUMBER 1 87 Figs. 1-7. Hydroptila artesa. Figs. 1-4. Male genitalia. 1, Ventral. 2, Dorsal. 3, Lateral. 4, Aedeagus (lateral). Figs. 5-7. Female genitalia. 5, Ventral. 6, Internal apparatus. 7, Eighth sternite (ventral). IA, Inferior Appendage; BpP, Bilobed Process. Hydroptila artesa Mathis and Bowles NEw SPECIES Figs. 1-7 This species belongs to the H. tineoides group and is most similar to H. paramoena Harris. The new species is distinguished from the latter species by the shape of tergite X, inferior appendages, and aedeagus. Male.—Length 1.9-2.8 mm. Antennae 30-34-segmented. Color yellowish brown in alcohol. Abdominal segment VII with long ventral process extending to middle of segment VIII. Segment IX with dorsum broadly rounded, lacking setae; with narrow lateral lobe bearing apical setae; excised deeply anterodorsally and gently postero- and anteroventrally; apodeme long and narrow, arising from dorsal one-half of seg- ment, extending into, but never past seg- ment VII. Segment X fused dorsally with segment IX; lateral margins lightly sclero- tized; hood-shaped in lateral view, but not sharply upturned, widening posteriorly in dorsal view, apex emarginate. Inferior ap- 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pendages short, beak-like, relatively wide in lateral view; distal portion separated and basal portion fused along meson in ventral view; with broad, thumb-like dorsal pro- jection bearing long setae. Bilobed process present. Aedeagus short, less than one-third length of abdomen; widest at base, with two long, apical processes; process bearing eja- culatory duct long and slender, duct pro- truding at apex; other process flattened and pointed apically; titillator spirally one-third turn anteriorly. Female.— Length 2.4—3.4 mm. Antennae 22-26-segmented. Similar to male in gen- eral appearance. Abdominal segment VI with broad, spur-like ventral process. Ab- domen with three pairs of apodemes; pos- terior-most pair extending length of seg- ment IX; mesial pair with anterior end expanded and curved toward midline, ex- tending from posterior margin of segment VIII into anterior one-half of segment VII; lateral pair straight, extending from poste- rior margin of segment IX to posterior mar- gin of segment VI. Segment VIII tapering; sternite with pair of sclerotized lobes pos- teriorly, each bearing 3 (rarely 4) stout setae, membranous anterior to lobes. Segment IX tapering slightly. Segment X short, ovoid; bearing pair of short cirri subterminally and numerous small setae. Internal apparatus lyre-shaped, with a star-like mesial config- uration. Immatures. — Unknown. Etymology.—French, referring to the typ- ical spring habitat of the species. Holotype, male and allotype.— Missouri, Shannon County, Alley Spring, Ozark Na- tional Scenic Riverways (O.N.S.R.), 5 mi W Eminence, Hwy 106, 16 August 1987, M. Mathis, S. Tedder (NMNH). Paratypes.—Same data as holotype, 18 34, 9 92 (NMNH, INHS, UAIC); Carter Coun- ty, Big Spring, O.N.S.R., 7 mi S Van Buren, Hwy 103, 18 August 1987, 15 43, M. Math- is, S. Tedder (INHS, NMNH); Dent Coun- ty, Current River, Montauk State Park, 23 mi SE Salem, Hwy 119, 15 October 1988, 7 48, M. Mathis, D. Bowles (INHS); Oregon County, Eleven Point River, Hwy 19 bridge, 1.5 mi NE Greer, 6 July 1988, 8 48, M. L. Mathis, D. E. Bowles (NMNH); Ozark County, Althea Spring, 8 mi NW Caulfield, H Hwy, 8 August 1988, 5 43, M. Mathis, S. Tedder, L. Tedder (AMNH); Ozark County, North Fork White River, H Hwy bridge, 8 mi NW Caulfield, 8 August 1988, 10 99, 8 August 1988, M. Mathis, S. Tedder, L. Ted- der (AMNH, INHS); Arkansas, Fulton County, Mammoth Spring, 19 July 1969, 4 46 (INHS). Discussion. — Males of Hydroptila artesa, as well as those of H. amoena Ross, H. par- amoena, and H. oneili Harris, have a long ventral process on abdominal segment VII, beak-like inferior appendages, and the phal- lus divided into two apical processes. Hy- droptila hamata Morton also has been re- ported to share these characters, but close examination will reveal a third, spur-like apical phallic process (see Ross 1944; fig. 512D). Hydroptila artesa differs from all three species in that the phallus is less than one-third the length of the abdomen and the apical phallic process that lacks the ejac- ulatory duct is flattened into a broad point- ed apex. It is distinguished easily from H. paramoena, the most closely related species, in that tergite X is not upturned strongly in lateral view, becomes wider posteriorly and is broadly excised in dorsal view, and the inferior appendages are separated along the meson. Feniales of H. artesa may be dis- tinguished from those of H. amoena by the presence of a pair of subterminal lobes on sternite VIII rather than well-developed ovate plates. They differ from those of H. hamata in lacking a pair of transverse scler- otized bars anterior to the apex of sternite VIII. Hydroptila sandersoni Mathis and Bowles NEw SPECIES Figs. 8-11 This species, like the preceding one, is a member of the H. tineoides group and is most closely related to H. oneili and H. amoena. It is distinguished by the shape of VOLUME 92, NUMBER 1 Figs. 8-11. IA, Inferior Appendage; BpP, Bilobed Process. tergite X, inferior appendages, and the length of the apodemes. Male.—Length 2.1-2.3 mm. Antennae 29-30-segmented. Color brown in alcohol. Abdominal segment VII with long, medial, ventral process extending to posterior mar- gin of segment VIII. Segment VIII short, flexed ventrally. Segment IX excised deeply anterodorsally and ventrally and shallowly posteroventrally; dorsum broadly rounded, setation absent; with long narrow lateral lobe bearing many setae along posterior and ven- tral margins; apodeme long, narrow, ex- tending into segment VI. Tergite X long and slender in dorsal view, tapering posteriorly, apex complete; cap-shaped in lateral view, apex straight or slightly up-turned; fused dorsomedially to posterior of tergite IX, Hydroptila sandersoni, male genitalia. 8, 89 Ventral. 9, Dorsal. 10, Lateral. 11, Aedeagus (lateral). ventrally to lateral lobes of segment IX. In- ferior appendages ‘‘beak-like,” with dor- solateral thumb-shaped projection bearing several setae; main axis slender in lateral view, fused basally and contiguous along meson in ventral view. Bilobed process short. Aedeagus widest basally, produced into two slender apical processes; process bearing ejaculatory duct thicker of two, gently curv- ing dorsally; ejaculatory duct not protruding or bent; apex of other process narrow, curv- ing sharply dorsally; titillator turning three- quarter revolution anteriorly. Female. — Unknown. Immatures. — Unknown. Etymology. — Named in honor of Dr. Mil- ton W. Sanderson for his many contribu- tions to the study of Trichoptera. 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Holotype. — Arkansas, Stone County, Syl- amore Creek, Gunner Pool Recreation Area, 20 July 1988, C. Carlton, R. Leschen (NMNB). Paratypes.—Same as above, | ¢(NMNH); same, but 22 July 1987, 2 6 (UAIC); John- son County, Mulberry River, 5 mi W Oark, Hwy 215, 23 July 1986, 1 6, D. Bowles (INHS); Carrol County, Osage Creek, Hwy 68 bridge, 4 August 1985, | ¢, D. Bowles, M. Mathis (INHS). Discussion. — Hydroptila sandersoni bears a close resemblance to both H. amoena and H. oneili, but differs in a number of features. Unlike these species, tergite X of H. san- dersoni is slender and complete apically, the ventral axis of the inferior appendages is narrow in lateral view, and the apodemes of segment IX are relatively longer and al- ways extend into segment VI. The apex of the ejaculatory duct of H. sandersoni is straight and does not protrude, but in H. oneili, it is protruding and bent. Hydroptila sandersoni was collected from the head- water reaches of warm-water streams in northern Arkansas. The H. tineoides species group encom- passes 20 species, all of which are Nearctic except H. tineoides Dalman (Palaearctic) and H. moselyi Ulmer (Oriental). In this group, the inferior appendages are somewhat C-shaped, typically consisting of a short, beak-like ventral axis and a broad dorso- lateral projection bearing one or more long setae. The distal portion of the aedeagus is divided into two or three processes that may be variously modified. Tergite X is well- developed and usually hood-shaped in lat- eral view. The medial process of sternite VII may be short and spur-like or elongated. A bilobed process is present in at least some species. Within the tineoides group there are five species that closely resemble H. amoena. In these species, the medial process of ster- nite VII is elongate, a bilobed process is present, and the aedeagus is divided into two long, slender apical processes. Follow- ing is a key to the Nearctic species of the H. tineoides group that are identified as H. amoena using either Ross (1944) or Blickle (1979). 1. Aedeagus short, less than one-third as long as abdomen; phallic process lacking ejaculatory duct with apex flattened and pointed .. H. artesa — Aedeagus longer, greater than one-third as long as abdomen; apical process not as above .... 2 2. Apex of tergite X complete; ventral axis of in- ferior appendage slender . H. sandersoni — Apex of tergite X excised; ventral axis of in- ferior appendage stout... cece ao. caer oreo 3 3. Apex of tergite X with deep, wide excision (see Harris 1985; fig. 8E); tips of phallic processes simple H. amoena — Apex of tergite X ‘only slightly emarginate; phallic process bearing ejaculatory duct with tip modified or bent ........ 4 4. Tergite X strongly upturned in lateral \ view (See. Harris 1985; fig. 8A); phallic process with an apical, rattle-like structure (see Harris 1985; fig. 8D) .... H. paramoena — Tergite X only slightly upturned i in lateral view (see Harris 1985; fig. 9A); phallic process with ejaculatory duct protruding and noticeably bent (see Harris 1985; fig. 9D) H. oneili Neotrichia arkansasensis Mathis and Bowles New SPECIES Figs. 12-15 Neotrichia arkansasensis closely resem- bles N. okopa Ross, N. sonora Ross, and N. osmena Ross, but it is distinguishable from these species by the shape of tergite X and the inferior appendages. Male.—Length 2.0-2.4 mm. Antenna 18- segmented. Color brown in alcohol. Ab- dominal segment VIII small, subquadrate in dorsal view. Segment IX annular, apo- deme short; with complex network of in- ternal sclerotization; dorsum membranous, with many small setae; extending poste- riorly and covering tergite X. Tergite X con- sisting of a pair of sclerotized, pointed pro- cesses curving posteroventrally and a pair of heavily-sclerotized basal pieces extend- ing ventrally. In dorsal view, basal piece produced into a short, mesial process joined to a concave lateral shoulder; piece sub- VOLUME 92, NUMBER | 91 Figs. 12-15. Neotrichia arkansasensis, male genitalia. 12, Dorsal. 13, Lateral. 14, Ventral. 15, Aedeagus (dorsal). IA, Inferior Appendage; SgP, Subgenital Plate; Br, Bracteole. triangular in lateral view. Inferior append- ages dark brown to black; tapering through- out length in ventral view, curving slightly dorsad in lateral view; basally with a down- curving, finger-like projection on mesial face anda tooth-like process just posterior to this projection. Bracteoles spatulate, with con- cave mesial face, bearing many setae. Subgenital process well developed, with rounded mesial lobe flanked on both sides by long setae; bearing two pairs of short setae on midline of ventral surface. Aedea- gus with distinct proximal and distal re- gions; proximal portion long, wide, and cy- lindrical basally, with a short, tapering neck; distal portion tapering slightly, with a dis- tinct ejaculatory duct and subterminal gen- ital pore; spiral process large, making one revolution anteriorly before extending pos- teriorly one-half revolution. Female.— Unknown. Immatures. — Unknown. Etymology.— Latin: of Arkansas. Holotype.—Arkansas, Madison County, Kings River, 5 mi S Kingston, NW 4, SW Vy Sect. 4, T 15 N, R 24 W, 2 June 1985, D. Bowles (NMNH). Paratypes.—Same as above, | 4 (INHS); Johnson County, confluence of Little Piney and Sulfur Creeks, 12 mi N Hagarville, Hwy 123, 7 June 1986, 3 4, C. Robotham (NMNH, UAIC); Johnson County, un- named spring, 5 mi W Oark, Hwy 215, 12 June 1986 1 4, D. Bowles (INHS); Sharp 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON County, Spring Creek, Spring Creek Wild- life Management Area, 8 June 1988, 1 4, R. Leschen (UAIC). Discussion. — Neotrichia arkansasensis is most closely related to N. sonora, but also shares a number of characters with N. okopa and N. osmena. In N. arkansasensis and N. sonora, tergite X is produced into two long, downturning, sclerotized points that are poorly developed in both N. okopa and N. osmena. The darkly sclerotized base of ter- gite X viewed laterally forms a large trian- gular-shaped plate in N. arkansasensis and N. sonora, but it is reduced in N. osmena and forms a narrow point in N. okopa. The most outstanding difference between N. ar- kansasensis and N. sonora is the shape of the inferior appendages. In N. arkansasen- sis, they are straight or slightly upturned in lateral view, taper throughout their length and are straight or converging in ventral view, and have a finger-like process arising basally from the mesial face. The inferior appendages of N. sonora are sharply up- turned in lateral view, taper abruptly near the apex and are diverging in ventral view, and have the finger-like process arising near the midlength of the mesial face. The two species also differ in the shape of the scler- otized basal portion of tergite X; in ventral view, it forms a sharply angular triangle in N. sonora, but in N. arkansasensis it has a concave lateral face and forms a blunt point distally. ACKNOWLEDGMENTS We thank S. C. Harris for his advice and R. T. Allen, C. E. Carlton, W. C. Yearian, and E. H. Schmitz for reviewing the manu- script. We are further indebted to R. A. B. Leschen, C. E. Carlton, C. D. Rowbotham, S. K. Tedder, and L. K. Tedder for con- tributing specimens to this study. LITERATURE CITED Blickle, R. L. 1979. Hydroptilidae (Trichoptera) of America north of Mexico. Bull. N.H. Agric. Exp. Stn. 509: 1-97. Hamilton, S. W. and G. A. Schuster. 1978. Hydrop- tilidae from Kansas (Trichoptera). Entomol. News 89: 201-205. 1979. Records of Trichoptera from Kansas, II: The families Glossosomatidae, Helicopsychi- dae, Hydropsychidae and Rhyacophilidae. Tech. Publ. State Biol. Surv. Kansas 8: 15-22. 1980. Records of Trichoptera from Kansas, Ill: The families Limnephilidae, Phryganeidae, Polycentropodidae and Sericostomatidae. Tech. Publ. State Biol. Surv. Kansas 9: 20-29. Hamilton, S. W., G. A. Schuster, and M. B. DuBois. 1983. Checklist of the Trichoptera of Kansas. Trans. Kansas Acad. Sci. 86: 10-23. Harris, S. C. 1985. New microcaddisflies (Trichop- tera: Hydroptilidae) from Alabama. Proc. Ento- mol. Soc. Wash. 87: 606-621. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. British Mus. (Nat. Hist.), Entomol. 39: 135-239. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Ill. Nat. Hist. Surv. 23: 1-326. Schuster, G. A. and S. W. Hamilton. 1978. Records of the trichopteran families Hydroptilidae, Phil- opotamidae and Psychomyiidae from Kansas. Tech. Publ. State Biol. Surv. Kansas 7: 36-47. Unzicker, J. D., L. Aggus, and L. O. Warren. 1970. A preliminary list of the Arkansas Trichoptera. J. Ga. Ent. Soc. 5: 167-174. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 93-97 CUBITUS POSTERIOR IN HYMENOPTERA W.R. M. Mason Biosystematics Research Centre, Agriculture Canada, Research Branch, Central Ex- perimental Farm, Ottawa, Ontario K1A 0C6, Canada. Abstract. — Forewing veins of Hymenoptera named by Ross (1936) Cu, Cu 1, Cu la are really Cu A. Ross’s vein Cu Ib is a crossvein, 2cu-a, necessitating that Ross’s cu-a be called Icu-a. The rarely seen vein Cu 2 (Ross 1936) in Hymenoptera should be called Cu P. An apparent distal section of Cu P is readily seen in Rhopalosomatidae, and can be seen in spectral form in many other Apocrita. Key Words: When Comstock (1895) published his system for naming the veins of Hymenop- tera his nomenclature rested on the phylo- genetically unsound practice of interpreting the venation of Hymenoptera by compari- son with that of higher Diptera (Comstock 1918, p. 383 ff). The serious flaws in the resulting scheme may well have accounted for the general reluctance of hymenopterists of the early 20th century to use Comstock’s system (Rohwer and Gahan 1916). These flaws were not corrected until 40 years later when H. H. Ross (Ross 1936) reinterpreted hymenopterous venation by comparing ve- nation of primitive Hymenoptera (Sym- phyta) with what he then believed to be the most closely related extant orders, Mega- loptera, Trichoptera, and Mecoptera. The soundness and brilliance of his interpreta- tion can be seen by the widespread accep- tance, with no essential modification, of the Ross system today, over 50 years later. There were some points about which Ross expressed doubt. One was the identity of the branches of Media and Cubitus. He was familiar with Lameere’s and Martynov’s system of naming convex veins “anterior” and concave ones “posterior” because he Hymenoptera, venation, wing, Cubitus Posterior, Comstock-Ross system cited Martynov’s work and labels veins of many of the non-hymenopterous wings as MA and MP. He decided to call the single Media vein of Hymenoptera ‘“M”’ mainly for lack of evidence and for convenience. For branches of Cubitus he considered the evidence equivocal, apparently because the posterior branch of the Cubital vein of Salis is neutral in profile, even though falling near the claval furrow in a concave part of the wing. As a compromise he retained the Comstock names for the branches of Cubi- tus, Cu 1 and Cu 2, the former subdivided into Cu la and Cu 1b. Subsequent research has clarified the doubts felt by Ross so that one can no longer justify using Comstock- Ross nomenclature for the branches of Cu- bitus in Hymenoptera (Carpenter 1966, Wootton 1979, Rasnitsyn 1980). The La- meere hypothesis and its background is best summarized by Carpenter (1966). Com- stock’s hypothesis is criticized by Lameere and by Martynov (op. cit.). Veins Cu 1 and Cu 2 of the Ross system should be called Cu A and Cu P, respectively, to align hy- menopterous vein nomenclature with mod- er usage and opinion among students of other orders, and especially with usage in 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Icu-a 2cu-a Figs. 1, 2. Symphyta forewings. 1, Orussus occi- dentalis Cr. (Orussidae) to show basal, concave nebular section of Cu P and possible apical, spectral section of Cu P (weakly defined and often absent). 2, Yvela bakeri Konow (Xyelidae) to show nebular basal section of Cu P, claval furrow and cu-a crossvein. (Conventions of delineation follow Mason (1986). Scale lines = | mm). the study of those fossil groups most prob- ably including the sister group of Hyme- noptera (Rasnitsyn 1980). Ross, following Comstock’s system, named the two branches of his first cubital vein Cu laand Cu Ib. In his figures of non- hymenopterous wings (Ross 1936, figs. 2, 6, 8, 20-22, 24) he shows both Cu la and Cu 1b extending to the wing margin far dis- tad of the claval notch (which is found on the margin between the apices of 1A and Cu 2). In his figures of Hymenoptera, how- ever (Ross 1936, figs. 3-5, 23), he shows Cu 1b turning abruptly caudad, crossing the claval furrow, where a bulla is formed, and meeting 1A proximad of the claval notch (Fig. 2). Because the vein called Cu 1b by Ross follows such a radically different course in Hymenoptera compared to that in the older orders, Megaloptera, etc., Ross’s in- terpretation is questionable. Furthermore, if my interpretation of the distal section of Cu P in Hymenoptera is correct (see below), Ross’s hypothesis calls for his Cu 1b to cross Cu P and meet 1A, a highly improbable course. Another interpretation of “Cu 1b” in Hy- menoptera is that it is a second cu-a cross- vein. The Megaloptera (Ross 1936) and the extinct Miomoptera (Rasnitsyn 1980), groups postulated as possibly ancestral to Hymenoptera, are copiously supplied with crossveins. The second cu-a crossvein in Hymenoptera could well have a compound origin similar to that suggested for the first cu-a by Ross (1936, p. 106), i.e. crossveins extending from Cu A to Cu P and from Cu P to 1A lined up with one another during the reduction and loss of Cu P. Modern thought (summarized in Woot- ton 1979) is that Cu P is closely associated with the claval furrow. In light of this it seems to me unreasonable to postulate that Cu A should have a branch crossing the site of Cu P (and the extant claval furrow) to join 1A. I think the existence of a second cu-a crossvein is a more tenable hypothesis for Hymenoptera. A truly branched Cubitus (Cu Al, Cu A2) can be seen in Stephanidae (Fig. 3). Certainly Cu P existed in the forewing of many extinct Neopterous insects and 1s eas- ily seen in extant forms, where it closely parallels the claval furrow. Ross (1936) drew attention to the trace (nebulous, Mason 1986) of a concave vein along the basal part of the claval furrow in forewings of Xyelidae and called it Cu 2 (Fig. 2), his equivalent of what recent authorities call Cu P. Signifi- cantly, there is a similar nebulous vein in Orussidae (Fig. 1). Recently (Mason 1986), I noticed a usually spectral concave vein in forewings of several groups of Apocrita, running distally from the junction of the claval furrow and 2nd cu-a (= Cu 1b, Ross). Further searching has revealed a concave spectral vein in phylogenetically old mem- bers of all apocritous major groups that have most of the venation preserved. At least some species of the following families have the vein present: Stephanidae, Megalyridae, Trigonalidae, Aulacidae, Monomachidae, Roproniidae, Ibaliidae, Cynipidae, Bethyli- dae, Scolebythidae, Tiphiidae, Sapygidae, VOLUME 92, NUMBER | imm 95 Figs. 3-10. Forewing of diverse Apocrita showing apical trace of Cu P branching from claval furrow. 3, Schlettererius cinctipes Cr. (Stephanidae); note 2 branches of Cu A. 4, Monomachus sp. (Monomachidae). 5, Ropronia garmani Ashm. (Roproniidae). 6, Orthogonalys pulchella Cr. (Trigonalidae), note minute jugum. 7, Liosphex varius Tow. (Rhopalosomatidae), note nebular Cu P, fusion of C and R, retention of Ir and loss of stigma. 8, Pristaulacus sp. (Aulacidae); note jugum defined by convex wing fold. 9, Pristocera atra Klug (Be- thylidae); note nebular adventitious vein (Ad.) between | cu-a and 2 cu-a, spectral concave combination of vein 2-M and medial furrow, concave spectral 2m-cu, c.v.-concave vein. 10, Exeristes roborator Grav. (Ichneumon- idae); note nebular | — Rs + M, a unique feature for this family. (Conventions of delineation follow Mason (1986). Scale lines = | mm). Anthoboscidae, Vespidae, Bradynobaeni- dae, Rhopalosomatidae, Astatidae, Hylaei- dae, Ichneumonidae (Figs. 3-10). The Cu P vein is so widespread among primitive Apocrita that I suggest it to be a basic char- acter of the Apocrita. The absence of the “vein” in Symphyta is puzzling. Either the distal part of Cu P disappeared completely in Symphyta and the vein in Apocrita is a newly evolved structure or Cu P is sup- pressed by some genetic mechanism in modern Symphyta. A poorly defined impression in some specimens of Orussus (Fig. 1) may be interpreted as Cu P and may hint that the suppressing mechanism was lost early in the evolution of Apocrita (per- haps among ancient Orussoidea?), thus al- lowing the vein to reappear (i.e. a reversal). Which choice one postulates is of little phy- logenetic consequence; both mechanisms result in an apomorphy for Apocrita and the name to be used for the vein might as 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON well be Cu P in either case because its po- sition and profile fit perfectly into a normal full venation. DISCUSSION Ross (1936) uses elements of the two in- compatible schemes; the Lameere nomen- clature for branches of Media and the dis- credited Comstock scheme for branches of Cubitus. Therefore Ross’s Cu 1 and Cu 2 should be now called Cu A and Cu P. The vein Cu P is general in putative sister- groups of Hymenoptera and a basal relict is visible in some Symphyta (Xyelidae, Orus- sidae). An apical part of Cu P may be pres- ent as a trace vein in many generalized Apocrita but 1s absent in Symphyta with the possible exception of Orussidae. Its pres- ence may be due to reversal of a character suppressed in Symphyta and is probably a synapomorphy for Apocrita. The vein Ross called Cu 1b is probably not a branch of Cu A but most likely is a compound crossvein like the more proxi- mal cu-a, composed of cua-cup and cup-a crossveins inherited from Permian or Trias- sic ancestors and aligned during the dete- rioration of Cu P. My hypothesis is that some Hymenoptera have relicts of Cu P and that there are two cu-a crossveins. I recommend the following modifications to the Ross system. Ross 1936 Amendments Cu Ist Cu A (1Cu) Cull 2nd Cu A (2Cu) Culla 3rd Cu A (3Cu) Cu |b 2 cu-a cu-a 1 cu-a Cu 2 in Neoptera Cure For sake of brevity vein Cu A in almost all Hymenoptera might as well be called Cu, just as the putative MA in Hymenoptera universally receives the appellation M. I suggest that the terms Cu A and Cu P could be retained only for forms that have both. It is unfortunate that the final version of the logical and orderly system of Lameere (1922) (designating all concave veins with the prefix sub-) was never followed, but usage has now firmly fixed remnants of 3 different systems for naming 4 main concave veins: ““Subcosta” (Redtenbacher 1886), the only concave vein that he named; “Radial Sec- tor’ (Comstock 1895), merely a conve- nience term to. substitute for “R 2+3+4+5”; “Media Posterior” and ‘“‘Cub- itus Posterior” (Lameere 1922, and many earlier papers on the Commentry fossils, and Martynov 1924), 2 concave veins not rec- ognized by Comstock. Other recently used names (empusal, plical) seem to be unnec- essary innovations that probably add noth- ing to an understanding of phylogeny and needlessly complicate nomenclature (Woot- ton 1979). I agree with Wootton that con- servation is a more sensible policy than the coining of new names. Specialists in Hy- menoptera using other systems should not lose sight of the strength of Ross’s system: the names of veins designate structures be- lieved to be homologous throughout Insec- ta. Phylogenetic comparisons, even within Hymenoptera, are extremely difficult with- out such a universal system. It is not surprising that the spectral distal part of Cu P should escape the notice of researchers dealing with Aculeata s. |. or Ichneumonidae for they rarely need to deal with spectral venation, but the vein is nebu- lous and plain to see in Rhopalosomatidae. Systems of nomenclature for venation used by most aculeate workers have presumably allowed the phylogenetic significance of a Cu P vein in Rhopalosomatidae to escape attention. Using a traditional naming sys- tem, Cu P would be called Brachius or some other term, and homology would be masked by the inadequacies of the traditional no- menclature system. LITERATURE CITED Carpenter, F. M. 1966. The lower Permian insects of Kansas II. Psyche 73: 43-88. VOLUME 92, NUMBER 1 Comstock, J. H. 1918. The Wings of Insects. Com- stock, Ithaca, N.Y. 430 pp. Comstock, J. H. in Comstock, J. H. and A. B. Com- stock. 1895. A Manual for the Study of Insects. Comstock, Ithaca, N.Y. 701 pp. Lameere, A. 1922. Sur la nervation alaire des in- sectes. Bul. Acad. Roy. Belg. (Cl. Sci.) 1922: 38- 149. Martynov, A. V. 1924. Sur l’interpretation de la ner- vuration et de la trachéation des ailes des Odo- nates et des Agnathes. Rev. Russe d’Ent. 18: 145- 174. Mason, W. R. M. 1986. Standard drawing conven- tions and definitions for venational and other fea- tures of wings of Hymenoptera. Proc. Ent. Soc. Wash. 88: 1-7. OF Rasnitsyn, A. P. 1980. Proikhozhdenie 1 evolyutsiya pereponchatokrylykh nasekomykh. Trudy paleon- tologicheskovo instituta vol. 174, pp. 189 + 2. Redtenbacher, J. 1886. Vergleichende Studien uber das Flugelgeader der Insecten. Ann. K.K. Hofmus. 1: 153-232. Rohwer, S. A. and A. B. Gahan. 1916. Horismology of the hymenopterous wing. Proc. Ent. Soc. Wash. 18: 20-76. Ross, H. H. 1936. The ancestry and wing venation of the Hymenoptera. Ann. Ent. Soc. Am. 29: 99- ae Wootton, R. J. 1979. Function, homology and ter- minology in insect wings. Syst. Entomol. 4: 81- 93. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 98-105 SYSTEMATIC NOTES ON SOME BETHYLIDAE FROM BOTSWANA: EPYRINAE (HYMENOPTERA: ACULEATA) KARL V. KROMBEIN Department of Entomology, NHB Stop 105, Smithsonian Institution, Washington, D.C. 20560. Abstract.—Calyozina caperata, new species, is described from a unique male from Botswana; this is the first record of the genus in Africa. Epyris breviscapus Kieffer is transferred to Trachepyris, and redescribed from both sexes. Key Words: The present contribution describes two of the more interesting species of Epyrinae col- lected for the Smithsonian in Malaise traps by Per Forchhammer, Serowe, Botswana. In an earlier paper on several species of Pris- tocerinae (Krombein 1989) I gave some notes on the ecology of the area where the collections were made. The abbreviations used in the descrip- tions are as follows: LH—length of head from middle of clyp- eal margin to midpoint of vertex; WH-—width of head including eyes: WF—width of front (i.e. least interocular distance); HE—height of eye measured in lateral view; EV —distance from top of eye to crest of vertex in lateral view; WOT—width of ocellar triangle including posterior ocelli; OOL-— ocello-ocular line, least distance be- tween posterior ocellus and inner eye margin; LT—length of thorax, collar excluded, from anterior margin of pronotal disk to posterior end of propodeum. Hymenoptera, Bethylidae, Calyozina, Trachepyris Calyozina caperata Krombein New SPECIES Figs. 2, 3, 5-8 Male.—Length ca 2.5 mm (terminal ab- dominal segments estimated, removed be- fore measurement), forewing 1.7 mm. Black, moderately shining, head and thorax finely alutaceous; mandible except teeth, scape, pedicel and first four flagellar segments be- neath light red; basal antennal segments above, all of terminal segments, tegula and apical tarsal segments light brown; legs ex- cept tarsi darker brown; first four tarsal seg- ments white. Vestiture of head and thorax sparse, short and suberect; flagellar seg- ments above with short, suberect, moder- ately dense setae, apices of processes on first five flagellar segments with somewhat long- er setae. Wings clear, stigma light brown, veins almost colorless. Head: WH 1.09 x LH; WF 1.22 x HE and 0.61 x WH; EV 0.41 x HE; mandible (Fig. 5) stout, not so robust as figured be- cause of foreshortening on micrograph, quinquedentate, apical tooth much longer than inner four; eyes not protuberant, inner orbits diverging slightly above, ocular setae short and quite sparse; front with tiny scat- VOLUME 92, NUMBER 1 Figs. 1-5. holotype from Botswana. |, Antenna, C. staphylinoides (?), 33x; 2, antenna, C. caperata, 115; 3, dorsum of thorax, C. caperata, 75 x; 4, mandible and base of antenna, C. staphylinoides (?), 90 x ; 5, mandible, C. caperata, 455%. tered punctures and a slight protuberance above each antennal insertion, with a weak carina on each side extending obliquely from protuberance to lower inner eye margin; ocelli in a low triangle, frontal angle about 120°, OOL 0.79 x WOT; vertex broadly rounded; occipital carina complete; antenna 13-segmented (Fig. 2), dorsal length of scape, pedicel and first two flagellar segments in a ratio about 25:12:11:13, first flagellar seg- 99 Males of Calyoza staphylinoides Westwood (?) from Kenya, and Calyozina caperata Krombein, ment dorsally as long as wide; basal seg- ments modified beneath, in profile pedicel is roundly protuberant beneath, flagellar segments 1-7 have an elongate projection at apex about three-fourths as long as seg- ment on 2—4, somewhat shorter on | and 5, very short on 6 and 7, segments 8-11 not modified, subcylindrical, 11 about 1.4 as long as 10. Dorsum of thorax (Fig. 3), LT 1.8 = great- 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-8. lateral aspect, apices of paramere and volsella; 7, subgenital plate; 8, forewing. est width (at pronotal lobes); pronotal disk with median length a fourth as long as great- est width, lateral fourth strongly carinate anteriorly, middle section a subtriangular raised area, surface adjacent to anterior ca- rina and median raised area with short ra- diating rugae presenting a wrinkled appear- ance, disk posteriorly with sparse, scattered, tiny punctures, margin without transverse groove, sides not carinate, rounded to lat- eral declivous surface; side of pronotum carinate anteriorly on upper half; scutum with scattered tiny punctures, notauli well Calyozina caperata Krombein, holotype. 6a, Genitalia, ventral aspect at left, dorsal at right; 6b, developed on posterior two-thirds, con- verging slightly toward apex, parapsidal lines weak, present only on posterior half; scu- tellum anteriorly with a pair of pits con- nected by a deep narrow groove; propodeal disk 1.5 as wide as median length, mar- gined laterally by a weakly crenulate groove and carina, posterior margin strongly cari- nate, posterolateral corner foveolate, me- dian discal carina complete, more strongly sculptured basal area about as long as wide, limiting carinae weak, rounded toward apex, surface with a few weak longitudinal carinae VOLUME 92, NUMBER | at base, irregularly and mostly transversely rugulose posteriorly, areas adjacent to basal sculptured section closely, obliquely carin- ulate, a narrow area anterior to posterior carina with weak, short, longitudinal cari- nae; posterior propodeal surface lacking a median carina; forewing (Fig. 8). Abdomen not petiolate; subgenital plate (Fig. 7) short, apical margin weakly emar- ginate; genitalia (Fig. 6). Female. — Unknown. Discussion.—Calyozina Enderlein 1s presently known only from males of six species, the type-species, ramicornis Ender- lein, from Taiwan, four Neotropical species described by Evans, amazonica, azurea, mexicana, neotropica, and caperata from Botswana. The included species are similar to males of Calyoza Westwood in having pectinate antennae with processes beneath at the apices of many of the flagellar seg- ments (cf Figs. 1, 2). They differ at once from Calyoza in having a well-developed first flagellar segment distinctly separated from the second, whereas the first flagellar segment in Ca/yoza 1s short, broadly joined to the second, forming a ring joint (cf Figs. 2, 4), so that the antennae appear superfi- cially to be only 12-segmented. The following combination of characters distinguishes caperata from its congeners: small size, 2.5 mm long as compared to 5.0- 7.0 mm; mandible quinquedentate; pedicel roundly protuberant beneath; pectinations present on flagellar segments 1-7, each shorter than the length of the segment; pro- notal disk strongly carinate anterolaterally and with short radiating rugae on median raised area; scutellum anteriorly with lateral pits connected by a deep, narrow groove; and posterior surface of propodeum without a median carina. Holotype.—é; Botswana, Serowe, Farm- er’s Brigade, July 1987, malaise trap, Per Forchhammer (USNM). Etymology.—From the Latin caperatus, wrinkled, in allusion to the distinctive 101 sculpture of the median part of the pronotal disk (Fig. 3). Trachepyris breviscapus (Kieffer) NEw COMBINATION Figs. 9-19 Epyris breviscapus Kieffer, 1904: 402-403; (6; Cape Verde Islands: holotype in Gen- oa).— Kieffer, 1908: 27 (listed). — Kieffer, 1914: 333 (redescribed in German). Acanthepyris spinitarsis Kieffer, 1904: 402 (2; Portuguese Guinea, now Guinea-Bis- sau; holotype in Genoa).— Kieffer, 1914: 404 (redescribed in German). Epyris spinitarsis (Kieffer) Kieffer, 1908: 28 (transferred to Epyris). Acanthepyris propinquus Turner, 1928: 134— 135 (2; Mossel Bay and Queenstown, South Africa; syntypes in London). Acanthepyris breviscapus (Kieffer) Benoit, 1957: 11 (6; Zaire; synonymized 2 spini- tarsis and 2 propinquus). This relatively common species of Tra- chepyris occurs from the Cape Verde Islands eastward to eastern Zaire and southward to Botswana and South Africa. Specimens were not available when I prepared a paper on the Ceylonese Trachepyris (Krombein 1987), so I take this opportunity to make the ge- neric transfer and a description supple- mented by scanning electron micrographs. References in the description, e.g. (Krom- bein 1987, Fig. 5), contrast the condition in haemorrhoidalis (Kieffer) with that in brevi- SCapUS. Female.—Length 6.1-8.1 mm, forewing 3.7-4.3 mm. Black, the following light red: mandible except base and inner and outer margins, antenna except apical segments in- fuscated above, legs variable, mid and hind coxae and rest of all legs light red, or only the tarsi light red, rest of legs light to dark brown, first four abdominal segments black except occasionally apical two-thirds or half of fourth, and all of fifth and sixth segments 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-13. mandible, apical half, outer surface, 215 surface, 115 x. light red. Wings clear to slightly infumated, stigma medium brown, veins lighter brown. Head shining above, delicately aluta- ceous, WH 1.20-1.23 x LH, not carinate posterolaterally, posterior margin broadly Trachepyris breviscapus (Kieffer), female. 9, Foretarsus, 85 x; 10, foretarsal claws, 215; 11, x: 12, mandible, apical half, inner surface, 215 =; 13, scape, upper and slightly incurved; apical half of man- dible (dorsal, Fig. 11, ventral, Fig. 12) rounded at apex, inner margin with a long, relatively slender subapical tooth (T1) and a shorter, blunt tooth (T2) somewhat basad, VOLUME 92, NUMBER 1 Figs. 14-17. Trachepyris breviscapus (Kieffer), male. 30x; 16, mandible, 150 x; 17, antenna, basal segments, four modified, flattened sensilla chaetica (S) on ventral surface below subapical tooth; clypeus short, raised along midline but not carinate, apical margin of lobe rounded but with narrow emargination in middle; WF 1.52-1.61 x HEand 0.69-0.71 x WH; front with or without a short, weak median groove anteriorly, usually with small, scattered punctures mostly separated from each other by twice or more a puncture diameter, the punctures rarely somewhat deeper and less separated; scape (Fig. 13) 2.75 = as long as wide, longer than in haemorrhoidalis (Krombein 1987, Fig. 5), upper surface 14, Dorsum of thorax, 30x; 15, head and pronotum, 75x. rather flattened and smooth, margined an- teriorly by a row of stout, short bristles that are denser than in haemorrhoidalis and pos- teriorly by sparser, longer bristles; ocelli small, posterior pair almost at hind margin of head, OOL 1.31-1.46 x WOT, front an- gle of ocellar triangle about 90°. Thoracic dorsum delicately alutaceous except propodeal disk glossy; pronotum not carinate anteriorly or laterally, impunctate in middle, elsewhere with punctures of moderate size separated by once or twice the diameter of a puncture; scutum and scu- tellum practically impunctate; dorsal pro- 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18 | Figs. 18, 19. subgenital plate. podeal surface 0.57 x as long as wide, cen- tral area with five longitudinal carinae, the two lateral pairs converging gradually pos- teriorly, median and lateral carinae reaching margin, intervening pair almost reaching margin, areas between carinae with trans- verse carinules; posterior surface with com- plete median carina; forefemur |.96-2.00 x as long as wide, stouter than in haemor- rhoidalis (2.2 x as long as wide); foretarsus (Fig. 9) with pecten about as in haemor- rhoidalis; tarsal claw cleft (Fig. 10), inner ray shorter than in haemorrhoidalis (Krom- bein 1987, Fig. 8); costa with short setae only, transverse median with short stub. Male. —Length 4.8-6.1 mm, forewing 3.1- 3.6 mm. Coloration almost like that of fe- male except mandibular teeth also dark, legs except coxae usually light red, mid and hind femora infrequently medium brown, and seventh abdominal segment also light red. Head glossy, in frontal view (Fig. 15), WH 1.08-1.21 =x LH, not carinate posterolater- ally, posterior margin slightly rounded out; mandible (Fig. 16) quinquedentate, upper Trachepyris breviscapus (Kieffer), male. 0.5mm 18, Genitalia, ventral aspect at left, dorsal at mght; 19, and lower teeth longer than three rounded intermediate teeth; clypeus raised along midline but not carinate, apex of median lobe rounded; front with a short, weak, me- dian groove from antennal insertions, punc- tures small, closer anteriorly and separated by about twice a puncture diameter over most of surface; WF 1.04-1.10 x HE and 0.55 x WH; ocelli in a low triangle, OOL 1.03-1.17 x WOT, front angle of ocellar triangle about 115°; antenna with third seg- ment 1.52-1.55x= as long as wide, a bit shorter than fourth (Fig. 17), ratio of first four segments ranging from 20:5:17:20 to 25:5:20:26. Thoracic dorsum (Fig. 14) glossy; pro- notal disk without anterior or lateral cari- nae, impunctate along a narrow median area, laterally with small punctures separated from each other by one to two puncture diameters; median length of dorsal propo- deal surface about half its greatest width, laterally and posteriorly with a crenulate groove adjacent to marginal carina, central area with three to five longitudinal carinae, VOLUME 92, NUMBER | the three inner carinae rather close and usu- ally reaching posterior crenulation, lateral pair more separated, often present only an- teriorly, curving toward each other when longer but not reaching apex, surface be- tween carinae smooth or with radiating car- inules; posterior propodeal surface with complete median carina. Subgenital plate (Fig. 19); genitalia (Fig. 18). Remarks.— 7. breviscapus is rather sim- ilar to haemorrhoidalis in coloration but is somewhat larger, 2 6.1-8.1 mm long com- pared to 4.7-5.8, ¢ 4.8-6.1 compared to 4.4— 4.8, and differs in other details. The female has a relatively longer scape with a row of stout, denser setae anteriorly on upper sur- face, a stouter forefemur and a shorter inner ray of the tarsal claw. The male genitalia of the two species are quite different, the par- amere of breviscapus being considerably broader and thinner, and clothed on outer, upper margin with close setae, which are lacking in haemorrhoidalis (Krombein 1987, Fig. 19) except for a pair at apex. Specimens examined (all USNM).—6 8, 28 4, Botswana, Serowe, Farmer’s Brigade, malaise trap, Per Forchhammer, dated as follows: Feb (2), 18-30 June (é) and Sep (4), 1986; Jan (2), June (2), Jul (3 4), Aug (2, 2 6), Sep (2, 4), Oct (4), Nov (2 4) and Dec (9, 17 4), 1987. Dates of collection suggest that 105 seasonal activity begins in June and extends through February. ACKNOWLEDGMENTS I am grateful to Beth Norden, Depart- ment of Entomology, for skillful prepara- tion of uncoated specimens for SEM study, and to Susann Braden, Scanning Electron Microscope Laboratory, for making the mi- crographs. I thank George Venable, De- partment of Entomology, for making the line drawings, and for mounting the plates. LITERATURE CITED Benoit, P. L. G. 1957. Hymenoptera—Bethylidae. Exploration du Pare National Albert, Mission G. F. de Witte 1933-1935. 88: 1-57. Kieffer, J.-J. 1904. Description de nouveaux Dryin- inae et Bethylinae du Musee de Génes. Ann. Mus. Civ. Stor. Nat. Genova 41: 351-412. . 1908. Hymenoptera, family Bethylidae. Gen- era Insectorum 76: 1-50. 1914. Bethylidae. Das Tierreich. 41: 1-595. Krombein, K. V. 1987. Biosystematic studies of Cey- lonese wasps, XVIII: The species of Trachepyris Kieffer (Hymenoptera: Bethylidae: Epyrinae). Pan- Pac. Entomol. 63: 135-144. 1989. Systematic notes on some Bethylidae from Botswana: Pristocerinae (Hymenoptera: Aculeata). Proc. Entomol. Soc. Wash. 91: 620- 631. Turner, R. E. 1928. New Hymenoptera of the family Bethylidae. Ann. & Mag. Nat. Hist. (10)1: 129- 152: PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 106-108 NINTH REPORT ON APHID-HOST RELATIONSHIPS AT THE LOS ANGELES STATE AND COUNTY ARBORETUM (HOMOPTERA: APHIDIDAE) MANYA B. STOETZEL AND LouIseE M. RUSSELL Systematic Entomology Laboratory, PSI, Agriculture Research Service, U.S. Depart- ment of Agriculture, Beltsville, Maryland 20705. Abstract.—This is the ninth report on collections of aphids from plants in the Los Angeles State and County Arboretum at Arcadia, California. Sixteen species are listed with information on their hosts, date of collection, morphs collected, and abundance. This report completes the identification of all material collected by the late Dr. Harry G. Walker, formerly with the Department of Arboreta and Botanic Gardens, Los Angeles State and County Arboretum, Arcadia, California, and submitted to us for identification. Key Words: Because aphids are important as plant feeders and vectors of plant viruses, infor- mation on their distribution and host plant relationships is of great importance. This is the ninth in a series of papers on the aphids and their host plants in the Los Angeles State and County Arboretum at Arcadia, California. This and four previous papers (Leonard et al. 1972, Leonard and Walker 1973, 1974, Walker et al. 1978) deal with a variety of aphids and their host plants. Four previous papers (Leonard et al. 1970, 1971a, b, c) dealt with six specific aphids and their hosts. The late Dr. Harry G. Walker, formerly with the Department of Arboreta and Bo- tanic Gardens, Los Angeles State and Coun- ty Arboretum, Arcadia, California, was re- sponsible for collecting the aphids identified in this paper. The following list includes under each aphid name the host plant, the date of collection, morphs collected, and the relative abundance. This report contains collection information not previously re- ported and completes the identification of aphids, host plants, California all material collected by Dr. Walker and submitted to us for identification. The following abbreviations are used for morphs: al = alata, ap = aptera, and ny = nymph. Aphis craccivora Koch Amorpha fruticosa L. 28/1V/67 ap, ny Abundant Artemisia maritima L. 23/V/66 ap, al, ny Abundant Aphis fabae Scopoli Nyssa sinensis D. Oliver 31/V/67 ap, ny Moderate Aphis gossypii Glover Carya ovata (Mill.) C. Koch 4/V/67 ap Scarce Gossypium thurberi Tod. 7/VIII/66 al Scarce Salix babylonica L. 1/V/67 ap, ny Scarce Aphis spiraecola Patch Acanthopanax trifoliatus (L.) Voss 20/1V/67 ap, al, ny Moderate Aloe sp. VOLUME 92, NUMBER 1 28/III/66 al Scarce Amorpha fruticosa L. 28/IV/67 ap, ny Moderate Anacampseros telephiastrum DC. 29/11/66 ap, al, ny Abundant Bauhinia variegata L. 28/1V/67 ap, al, ny Scarce Buddleia davidii Franch. 25/III/66 al Scarce Bursaria spinosa Cav. 27/XII/66 ap, al, ny Abundant Hypericum hookeranum Wight & Arn. 20/IV/67 al Scarce Lonicera fragrantissima Lindl. & Paxt. 1/11/67 ap, al, ny Scarce Pittosporum daphniphylloides Hayata 20/X11/66 ap, al, ny Abundant Prunus lyoni (Eastw.) Sarg. 22/V/66 al, ny Moderate Rubus palmatus Thunb. 1/VII/66 ap, al, ny Scarce Aphis sp. Wisteria floribunda (Willd.) DC. 6/V/67 al Scarce Aulacorthum solani (Kaltenbach) Aralia cordata Thunb. 4/V/67 al, ny Scarce Carya ovata (Mill.) C. Koch 4/V/67 al Scarce Chrysanthemum ‘Rambler’ 15/V/67 ap, al, ny Scarce Liquidambar formosana Hance 6/V/67 al, ny Scarce Brachycaudus helichrysi (Kaltenbach) Chrysanthemum ‘Rambler’ 15/V/67 ap, al Scarce Gazania linearis (Thunb.) Druce 3/1/67 al Scarce Cavariella pustula Essig Salix sp. 1/V/67 ap, ny Scarce Chaitophorus sp. Salix babylonica L. 1/V/67 al Scarce Eulachnus rileyi (Williams) Hibiscus arnottianus A. Gray kauaiensis 29/X/66 ap, al Scarce Hysteroneura setariae (Thomas) 107 Cynodon dactylon (L.) Pers. 27/XII/66 ap, al, ny Abundant Tllinoia sp. Spiraea longigemmis Maxim. 4/V/67 ap, ny Scarce Macrosiphum euphorbiae (Thomas) Anacampseros telephiastrum DC. 29/III/66 al Scarce Macrosiphum rosae (L.) Hex cornuta Lindl. & Paxt. 12/1V/66 ap, al, ny Moderate Macrosiphum sp. Gazania linearis (Thunb.) Druce 3/1/67 ap, al, ny Abundant Myzus persicae (Sulzer) Anacampseros telephiastrum DC. 29/11/66 ap, al, ny Scarce Carya ovata (Mill.) C. Koch 4/V/67 al Scarce Ilex cornuta Lindl. & Paxt. 12/IV/66 al Scarce Lonicera fragrantissima Lindl. & Paxt. 1/11/67 ap, al, ny Moderate Lonicera maackii (Rupr.) Maxim. 2/V/67 ny Scarce Salix babylonica L. 1/V/67 ap, ny Scarce LITERATURE CITED Leonard, M. D. and H. G. Walker. 1973. Aphids collected in the Los Angeles State and County Ar- boretum (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 75(2): 209-212. . 1974. Additional aphids collected in the Los Angeles State and County Arboretum (Homop- tera: Aphididae). Coop. Econ. Ins. Rpt. 24(39): 778-779. Leonard, M. D., H. G. Walker, and L. Enan. 1970. Host plants of Myzus persicae at the Los Angeles State and County Arboretum, Arcadia, California (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 72(3): 294-312. . 1971a. Host plants of Aphis gossypii at the Los Angeles State and County Arboretum, Arca- dia, California (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 73(1): 9-16. . 1971b. Host plants of three polyphagous and widely distributed aphids in the Los Angeles State and County Arboretum, Arcadia, California (Ho- moptera: Aphididae). Proc. Entomol. Soc. Wash. 73(2): 120-131. 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1971c. Host plants of Toxoptera aurantii at Walker, H. G., M. B. Stoetzel, and L. Enari. 1978. the Los Angeles State and County Arboretum, Ar- Additional aphid-host relationships at the Los An- cadia, California (Homoptera: Aphididae). Proc. geles State and County Arboretum (Homoptera: Entomol. Soc. Wash. 73(3): 324-326. Aphididae). Proc. Entomol. Soc. Wash. 80(4): 575- 1972. Host plants of aphids collected at the 605. Los Angeles State and County Arboretum during 1966 and 1967 (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 74(1): 95-120. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 109-110 NOTE The Identity of the Genus Hexaresta Hering (= Hyponeothermara Hardy, n. syn.) (Diptera: Tephritidae) Hering (1941. Siruna Seva 3: 18) de- scribed the monotypic genus Hexaresta and its type species H. juanita from a single specimen of unknown sex, supposedly from Paramaribo, Surinam. I have examined this specimen, deposited in the collection of the Museum fiir Naturkunde der Humboldt- Universitat zu Berlin, through the kindness of Dr. H. Schumann. It is actually the pa- laeotropical species described by Walker (1859. J. Proc. Linn. Soc. London, Zool. 3: 119) as Trypeta multistriga, which is the type species of Hyponeothemara Hardy (1986. Pacif. Insects Monog. 42: 71). Hex- aresta is thus a senior synonym of Hypo- neothermara, and juanita Hering is a junior synonym of Hexaresta multistriga (Walk- er), n. comb. The holotype of juanita 1s in poor con- dition, with its thorax broken and its ab- domen, most thoracic setae, left foreleg and right hindleg missing. Despite this, it can be clearly recognized as a specimen of multi- striga because of Hardy’s (1986) thorough redescription of the latter. I have also com- pared the type with a female from New Guinea in the National Museum of Natural History collection which Hardy determined as multistriga. The juanita type easily runs to Hyponeothermara in the key to the gen- era of Acanthonevrina in Hardy (1986): most of the diagnostic characters in the key, including the shape of the facial carina and the nonsetulose scutellum, can be observed on the juanita holotype. Although the tho- rax 1s broken, the mesonotal color pattern, which is distinctive of multistriga, is also evident in the juanita holotype. It differs only slightly from the pattern in Hardy’s fig. 44c and that of the female examined; the presutural dark brown spots are slightly larger and the postsutural spots are fused at the dorsocentral setae. The color of the head agrees closely with Hardy’s description; there is no frontal vitta as in Hexaresta formosa (Malloch), n. comb., the only other species that Hardy (1986) placed in Hyponeothe- mara. The wings are in good condition and their pattern 1s almost identical to that of the specimen examined, differing only by the lack of the small marginal hyaline spot between the two large hyaline spots in cell r, (compare also fig. 15 of Hering (1941) with fig. 44d of Hardy (1986)). Hardy (1986) states that this spot is variable in multistri- ga. Hering’s figure 1s erroneous in showing the base of cell c dark; it is subhyaline in the juanita holotype like most of the rest of the cell. Hexaresta multistriga almost certainly is not native to the Neotropical Region. Hardy (1986) reported its distribution to be Sula- wesi, the Moluccas, and New Guinea, and species that he considered closely related, such as Hexaresta formosa and the species of Neothemara Malloch and Pseudoneothe- mara Hardy, also occur in the Oriental and Australasian Regions. No closely related species are known from the New World. Hering (1941) accurately recorded the data on the label of the holotype of jwanita, which reads “S. Amerika, Surinam, 5.08, Bezirk [district of] Paramaribo, C. Heller S. V.” These data are doubtful, however, unless this species has been introduced into Suri- nam. More likely, considering the poor con- dition of the juanita holotype, is that it was placed on its present pin after falling off a different one. I am grateful to A. Freidberg (Tel Aviv Univ.), D. E. Hardy (Univ. of Hawai1), and D. A. Nickle and N. E. Woodley (Systematic 110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Entomology Laboratory) for reviewing this Allen L. Norrbom, Systematic Entomol- paper, and to H. Schumann (Humboldt- ogy Laboratory, USDA, ARS, PSI, % U.S. Universitat) for the loan of the holotype of | National Museum, NHB 168, Washington, H. juanita. D.C. 20560. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 111-114 MORPHOLOGICAL VARIATIONS IN THE HEMELYTRA OF CRYPHOCRICOS HUNGERFORDI USINGER (HETEROPTERA: NAUCORIDAE) ROBERT W. SITES Department of Agronomy, Horticulture, and Entomology, Texas Tech University, Lub- bock, Texas 79409-2134. Abstract.— Morphological variations in the hemelytra of Cryphocricos hungerfordi Usin- ger are described and illustrated. The distal margins of hemelytra of brachypterous forms are highly variable and may be straight, concave, convex, and asymmetrical. Additionally, a submacropterous form, intermediate between the brachypterous and macropterous forms, is described and illustrated. The northeasternmost record of the distribution of C. hun- gerfordi is now the South Llano River in central Texas. Key Words: The Naucoridae, or creeping water bugs, consists of predacious, aquatic bugs that pri- marily are pantropical in distribution. These bugs are common components of both lotic and lentic faunas. The genus Cryphocricos is restricted to the New World and La Rivers (1971, 1974, 1976) listed 14 species. Members of the genus are strongly dimorphic in thoracic development (Parsons 1974) which is associated with the brachypterous and ma- cropterous conditions (Usinger 1941, 1947). The brachypterous form is more common than the macropterous form (Usinger 1941), and Parsons & Hewson (1974) considered macropters very rare. The only member of this genus known to occur in the United States 1s Cryphocricos hungerfordi Usinger. The range of this species 1s from Mexico north to central Tex- as. Thus far, in the U.S. it has been recorded from only the Frio, Nueces, and Pecos Rivers in Texas (Polhemus & Polhemus 1988). Usinger (1947) described this species from both brachypterous and macropterous forms, and the type specimen is brachyp- Insecta, creeping water bug, polymorphism, wing terous. Usinger (1947) used the morpho- logical condition of hemelytra in three of eight couplets in his key to the species of brachypterous forms of Cryphocricos. Variations in hemleytral morphology of brachypterous C. hungerfordi are presented herein, and a third morphotype is described and illustrated. Additionally, the north- easternmost known limit of the range of C. hungerfordi is extended to the South Llano River in central Texas. Voucher specimens of each morphotype and the extremes of variation in the brachypterous condition are deposited in the Texas Tech University En- tomological Collection. Stupby SITE A total of 790 adults of C. hungerfordi was collected from a single population in the South Llano River on the Texas Tech University Center campus in Junction, Kimble Co., Texas, from April 1988 through January 1989. The South Llano River is on the Edwards Plateau and is north of the Bal- cones Fault Zone. This locality is ca. 160- 250 km north and northeast of the previous 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14. Morphological variations in the hemel- ytra of brachypterous Cryphocricos hungerfordi. records of C. hungerfordi, which were in the vicinity of the Balcones Fault Zone (Pol- hemus & Polhemus 1988) and represents the northeasternmost known limit of the range of this species. MORPHOLOGICAL VARIATIONS IN HEMELYTRA Asymmetry.— Asymmetry in the hemely- tra is common in the brachypterous forms. Specimens preserved in alcohol were ex- amined and iliustrated prior to pinning; thus, the asymmetry is not an artifact of pinning. Generally, asymmetry is evidenced as one wing shorter than the other (Fig. 1). This was apparent in many specimens with straight distal margins. Additionally, asym- metry was noted as hemelytra of different shapes (e.g. one straight and one sinuate dis- tal margin [Fig. 2]). The illustration of the brachypterous male in the original descrip- tion (Usinger 1947) appears to approach this condition. Whether this was intentional or an imperfection in drawing is not known, as there is no mention of asymmetry in the text. Brachyptery.—Symmetrical morpholog- ical variations in the hemelytra of brachyp- terous specimens are continuous, rather than falling into discrete categories of variation described here. The distal margins may be concave with the posteromedial corners produced (Fig. 3). In addition to the con- cavity, the angle of the distal margins may slope caudad toward the midline. A second variation is that of highly rounded distal margins (Fig. 4). Curvature is continuous from the posteromedial corners and the dis- tal margins merge with the costal margins. The posterolateral corners generally are poorly defined. The form that typically is described in keys (e.g. Polhemus 1984) has truncate, squared-off hemelytra (Fig. 5). Il- lustrations in the original description (Usin- ger 1947) showed the female to have straight distal margins of the hemelytra. Submacroptery.—In addition to the vari- able brachypterous forms and macropter- ous form (Fig. 7), a submacropterous form exists (Fig. 6). Usage of the term submac- ropterous is consistent with terms proposed by Slater (1975) to classify the major types of hemelytral structure. In this condition the wings extend to the 5th abdominal tergum and are represented by both corium and membrane, whereas membrane is absent in the brachypterous form. The hemelytral apices are produced and overlap. The per- centage of each hemelytron represented by the embolar area (14.0), measured at the level of the apex of the scutellum, is inter- mediate between that of the wider brachyp- terous (14.9 + 0.2 [y + SE]) and narrower macropterous (11.5 + 0.9) forms. The pos- terior margin of the pronotum and the hu- meral region of each hemelytron, including the embolar suture, are intermediate in de- gree of development as compared with bra- VOLUME 92, NUMBER 1 Us} Figs. 5-8. 5-7, Brachypterous, submacropterous, and macropterous morphotypes, respectively, of Crypho- cricos hungerfordi. 8, Dorsolateral view of humeral angle of left hemelytron in brachypterous (a), submacropterous (b), and macropterous (c) morphotypes of Cryphocricos hungerfordi. Note intermediate level of development of sutures and pronotum of submacropterous form. chypterous and macropterous forms (Fig. 8). Because only one male submacropterous specimen was collected, it was not dissected to examine the extent of intermediate con- ditions of the thorax and for presence of hindwings. DISCUSSION A total of 790 adults was collected in the South Llano River; six were macropterous (0.8%) and one was submacropterous (0.1%). The remainder (783 specimens) were bra- chypterous. With the discovery of the sub- macropterous form, this species should be referred to as polymorphic rather than di- morphic. Lindroth (1949) concluded that for cer- tain carabids, environmental uncertainty favors alary dimorphism, but in a stable environment brachyptery predominates. 114 Additionally, Slater (1972) suggested that the proportion of species with alary poly- morphism may not only indicate ecological stability of that area, but also stability in terms of evolutionary time. The South Llano River is a stable environment for naucorids (water temperature no colder than 11.5°C during the winter and a constant abundance of prey). Therefore, because the brachyp- terous form is predominant and the mac- ropterous form is present at such a low fre- quency, it 1s unknown whether the submacropterous form is aberrant or a con- sistent morphotype at a very low frequency of occurrence. Despite the diverse variations in the bra- chypterous condition, C. hungerfordi still may be identified with the key prepared by Usinger (1947). However, if this kind of variation is present in brachypterous forms of C. barozzii Signoret, C. breddini Mon- tandon, C. peruvianus De Carlo, or C. rufus De Carlo, many specimens cannot be iden- tified properly with the key because couplet six uses the shape of the “apical margins of hemelytra.” Studies on populations of those species are needed to determine the extent of this morphological variation. ACKNOWLEDGMENTS I am grateful to Ms. Becky Nichols for assistance with field collection of speci- mens. I thank Thomas J. Henry (Systematic Entomology Laboratory, USDA-ARS, % United States National Museum) for allow- ing me to examine USNM naucorid speci- mens and literature. I also thank L. Chan- dler, H. G. Thorvilson (Texas Tech Uni- versity), and J. T. Polhemus (3115 S. York, Englewood, Colorado 80110) for critical re- views of this manuscript. I would also like to thank Robert C. Albin for providing sup- port through Texas State Organized Re- search funds and C. Len Ainsworth for sup- port from the State of Texas Special Line Item: Texas Tech University Center at PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Junction. This 1s Contribution No. T-4-263, College of Agricultural Sciences, Texas Tech University, Lubbock, Texas. LITERATURE CITED La Rivers, I. 1971. Studies of Naucoridae (Hemip- tera). Biol. Soc. Nevada Mem. 2. it1 + 120 pp. . 1974. Catalogue of taxa described in the fam- ily Naucoridae (Hemiptera) supplement no. 1: Corrections, emendations and additions, with de- scriptions of new species. Biol. Soc. Nevada Occas. Pap. 38: 1-17. 1976. Supplement no. 2 to the catalogue of taxa described in the family Naucoridae (Hemip- tera), with descriptions of new species. Biol. Soc. Nevada Occas. Pap. 41: 1-17. Lindroth, C. H. 1949. Die Fennoscandischen Carab- idae. Eine tiergeographische Studie. III. Allge- meiner Teil. Meddel. Goteborgs Mus. Zool. Avd. 122: 1-911. Cited in Vepsalainen, K. 1974. The life cycles and wing lengths of Finnish Gerris Fabr. species (Heteroptera, Gerridae). Acta Zool. Fenn. 141: 1-73. Parsons, M. C. 1974. Modification of the interseg- mental region in the pterothorax of Cryphocricos (Heteroptera: Naucoridae). Psyche 81: 42-50. Parsons, M. C. and R. J. Hewson. 1974. Plastral re- spiratory devices in adult Cryphocricos (Naucor- idae: Heteroptera). Psyche 81: 510-527. Polhemus, D. A. and J. T. Polhemus. 1988. Family Naucoridae Leach, 1815. The creeping water bugs, pp. 521-527. In Henry, T. J. and R. C. Froeschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Bnill, Leiden and New York. Polhemus, J. T. 1984. Aquatic and semiaquatic He- miptera, pp. 231-260. /n Merritt, R. W. and K. W. Cummins, eds., An Introduction to the Aquat- ic Insects, 2nd ed. Kendall/Hunt Pub. Co., Du- buque. Slater, J. A. 1972. Lygaeid bugs (Hemiptera: Ly- gaeidae) as seed predators of figs. Biotropica 4: 145-151. 1975. On the biology and zoogeography of Australian Lygaeidae (Hemiptera: Heteroptera) with special reference to the southwest fauna. J. Aust. Entomol. Soc. 14: 47-64. Usinger, R. L. 1941. Key to the subfamilies of Nau- coridae with a generic synopsis of the new subfam- ily Ambrysinae (Hemiptera). Ann. Entomol. Soc. Am. 34: 5-16. 1947. Classification of the Cryphocricinae (Hemiptera: Naucoridae). Ann. Entomol. Soc. Am. 40: 329-343. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 115-119 TAXONOMIC NOTES ON SOME NORTH AMERICAN APHIDS V. F. EASTOP AND D. J. VOEGTLIN (VFE) British Museum (Natural History), Cromwell Road, London SW7 5BD, United Kingdom; (DJV) Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody, Champaign, Illinois 61820. Abstract.—Typic material of four poorly known aphid species from North America, Amphorophora singularis Hottes & Frison, Capitophorus corambus Hottes & Frison, Pho- rodon scrophulariae Thomas and Kakimia mimulicola Drews & Sampson, has been ex- amined. Appropriate synonymy or generic placement is suggested for each species. Key Words: There are nominal aphid species in the fauna of every region that have been de- scribed from one or a few specimens and have rarely, if ever, been identified after the initial collection. The reasons for this vary, but possibly the most common reason is that the original generic placement, based on earlier generic concepts and often erro- neous host records, is wrong. The four species discussed below fall into this cate- gory, and the synonymy or appropriate ge- neric placement for each is given. For each of the cases below, careful comparisons were made between the typic material and spec- imens of the species to which we believe they belong. Measurements and photo- graphs taken from the typic material are provided to support the placement. The quality of these photographs varies with the condition of the specimen on the slide. No attempt was made to remount the type spec- imens. Amphorophora singularis Hottes and Fri- son (1931) was described from a single ap- tera, taken on an unknown species of grass in Golconda, Illinois. We cannot distin- guish it from Microparsus (Megouroparsus) kislankoi Smith & Heie 1963, for which we believe it is an earlier name. Characters of Homoptera, Aphididae, taxonomy, North America A. singularis fit those of the subgenus Me- gouroparsus and match those in the original description of Aislankoi. Most obvious are the distinct sclerotization pattern, shape of siphunculi, diverging antennal tubercles, short setae on body, head, antennae and legs, and spinules on frontal tubercles and femora. Measurements taken from the type specimen (Table 1) closely match those from the type series of VW. kislankoi as does the photograph of the holotype of A. singularis (Fig. 1) when compared to the photograph of M. kislankoi in the original description (Smith & Heie 1963). We therefore consider Microparsus (Megouroparsus) kislankoi Smith & Heie a synonym of Microparsus (Megouroparsus) singularis (Hottes & Fri- son). M. kislankoi is reported to have as its hosts several species of Lespedeza (Legu- minosae). It is possible that the single aptera of singularis was only accidentally on the grass. Capitophorus corambus Hottes and Fri- son, 1931 was described from an alata (the holotype), one aptera (labelled morphotype) and one alata (paratype) collected from Rosa sp. in Galena, Illinois. The very slightly swollen siphunculi, sensoria only on anten- nal segment III in both aptera and alatae, 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. slightly capitate setae on head and body and shape of head and cauda indicate that they are the Ribes feeding species Hyperomyzus (Neonasonovia) ribiella (Davis 1919). Other specimens on the slides are one aptera with- out antennae which is Rhodobium porosum 4 Holotype of Amphorophora singularis on slide #10381, INHS (7.8 x). (Sanderson) and one nymph of Chaetosi- phon (Pentatrichopus) sp.? supporting the recorded host as rose. Photographs of the holotype and morphotype of C. corambus are shown in Figs. 2 and 3 and measure- ments are given in Table 1. We therefore Fig. 2. Holotype of Capitophorus corambus on slide #10657, INHS (7.8 x). VOLUME 92, NUMBER 1 Fig. 3. consider Capitophorus corambus Hottes & Frison a synonym of Hyperomyzus (Neon- asonovia) ribiella (Davis). A slide labelled Myzus scrophulariae bears the single specimen taken on Scrophularia nodosa at Carbondale, Illinois, and de- scribed as Phorodon scrophulariae Thomas Fig. 4. 117 Morphotype of Capitophorus corambus on slide #10658, INHS (6.25 x). 1879. Characters such as converging, rugose frontal tubercles, swollen and imbricated si- phunculi and strongly wrinkled abdominal dorsum place this specimen in the genus Hyalomyzus. We believe it is the species described as Rhopalosiphum monardae Da- vis 1911, now known a Hyalomyzus monar- Lectotype of Phorodon scrophulariae on slide #2798, INHS (6.25 x). 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. dae (Davis) and for which it would be an older name. We think it is undesirable to replace a well known and appropriate name with a previously unrecognised and inap- propriate name and will request the Inter- national Commision on Zoological Nomen- clature to supress the name scrophulariae. Fig. 6. Cotype of Kakimia mimulicola, slide from Essig Collection at U.C. Berkeley (6.25 x). A photograph of the lectotype of Phorodon scrophulariae is shown in Fig. 4, measure- ments are given in Table 1. Kakimia mimulicola Drews and Samp- son 1937 was described from Mimulus sp. (Scrophulariaceae). The short, distinctly shaped cauda, W-shaped front and rhinarial Cotype of Kakimia mimulicola, slide from Essig Collection at U.C. Berkeley (6.25 x). VOLUME 92, NUMBER 1 Measurements taken from holotype, lectotype or cotype specimens of species mentioned above, all measurements are in millimeters. Abbreviations are as follows: Ant. = antennal segment, Ant. VIb = base of sixth antennal segment, Ant. VIpt = process terminalis of sixth antennal segment, URS = ultimate rostral segment, Ht II = second segment of hind tarsus, Set. URS = accessory setae on ultimate rostral segment, Set. Cauda = setae on cauda, n.m. = not measurable. Table 1. Ultumate Rostral Segment Second Antennal Segment Caudal Hind Tarsus Length # of Setae Setae VIpt VIb Cauda Il IV Siphunculi Body Species Name Amphorophora singularis Hottes & Frison 0.1 n.m. 0.54 0.45 0.92 0.43 7 Holotype, aptera Capitophorus corambus Hottes & Frison 0.14 0.15 0.77 0.1 n.m 0.095 0.33 0.3 0.47 0.57 1.26 1.47 Holotype, alata 0.31 0.097 0.1 0.57 0.27 Morphotype, aptera Phorodon scrophulariae Thomas 0.1 0.12 0.41 0.12 0.26 0.3 0.42 1.56 0.35 Kakimia mimulicola Drews & Sampson Lectotype, aptera 0.13 0.14 0.09 > 0.3 5 0.1 0.20 0.14 0.39 0.29 0.1 1.6 Cotype, alata 0.08 0.28 0.12 0.19 1.98 Cotype, aptera 119 distribution in alatae suggest it belongs in the genus Myzodium. No clear differences can be found between it and either Myzo- dium modestum (Hottes) 1926 or Myzo- dium knowltoni Smith and Robinson 1975 both of which have as their hosts a moss (Bryophyta). It is regarded here as Myzo- dium mimulicola (Drews & Sampson), but its separate identity requires experimental confirmation. Photographs of an alata and aptera taken from cotype slides are shown in Figs. 5 and 6 and measurements are given in Table 1. ACKNOWLEDGMENTS This paper was supported in part by NSF Grant #BSR84-11418 which funded a month-long visit by the senior author to the Illinois Natural History Survey. We would also like to thank George Godfrey and John Bouseman of the Illinois Natural History Survey for reviewing this manuscript and Jerry Powell, University of California, Berkeley for the loan of the cotypes of Kaki- mia mimulicola. LITERATURE CITED Davis, J. J. 1911. Williams’ “The Aphididae of Ne- braska”; a critical review. University Studies of the University of Nebraska 1 1: 253-291; also paged as 1-39. 1919. Miscellaneous aphid notes 1. Can. Entomol. 51: 228-234. Drews, E. A. and W. W. Sampson. 1937. A new species of aphid from California. Pomona Coll. J. Entomol. Zool. 29: 29-30. Hottes, F.C. 1926. Two new genera and a new species of Aphididae. Proc. Biol. Soc. Wash. 39: 115-119. Hottes, F.C. and T. H. Frison. 1931. The plant lice, or Aphiidae, of Illinois. Bulletin of the Illinois State Natural History Survey 19: 121-447. Smith, C. F. and O. E. Heie. 1963. Megouroparsus, new genus, related to Microparsus Patch and Me- goura Buckton (Homoptera: Aphididae). Ann. Entomol. Soc. Amer. 56: 401-406. Smith, C. F. and A. G. Robinson. 1975. The genus Myzodium with the description of 7. knowltoni, new species (Homoptera: Aphididae). Proc. Ento- mol. Soc. Wash. 77: 481-486. Thomas, C. 1879. Noxious and beneficial insects of the state of Illinois. Rep. State Entomol. (Illinois) 8: 1-212. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 120-126 SEASONAL EMERGENCE PATTERNS AND DIVERSITY OF PLECOPTERA ON BIG HUNTING CREEK, MARYLAND, WITH A CHECKLIST OF THE STONEFLIES OF MARYLAND RICHARD M. DUFFIELD AND CHARLES H. NELSON (RMD) Department of Zoology, Howard University, Washington, D.C. 20059; (CHN) Department of Biology, University of Tennessee at Chattanooga, Chattanooga, Tennessee 37403. Abstract. —Species diversity and prevalence patterns of adult stoneflies were studied on Big Hunting Creek, Maryland, primarily by adult sampling. Twenty-two genera repre- sented by 39 species emerged throughout the year. The greatest number of species occurred from April through June. Seventeen of the species were recovered from stomach pump samples of trout from Big Hunting Creek. Twenty-five of the species are new records for the state of Maryland. An updated list of the stoneflies of Maryland is provided. Key Words: This study summarizes four years of adult stonefly collections on Big Hunting Creek, Maryland, situated within the boundaries of Catoctin Mountain Park and Cunning- ham Falls State Park. Included in our data are specimens obtained from stomach pump samples from trout caught in Big Hunting Creek and donated by local fishermen. Since a number of the species of stoneflies are new records for the state, an updated list of the Plecoptera of Maryland is given. Although a number of recent stonefly sur- veys have been conducted on streams in North America, none has been reported for Maryland. Investigations on eastern streams include those of Harper and Magnin (1969) [Quebec], Harper and Pilon (1970) [Que- bec], Woodall and Wallace (1972) [North Carolina], White (1974) [Kentucky], Neves (1978) [Massachusetts], Tkac and Foote (1978) [Ohio], Stark (1980) [South Caroli- na], Masteller (1983) [Pennsylvania], and Singh et al. (1984) [Ontario]. Plecoptera di- versity studies on mid-western and western streams include those of Sheldon and Jewett Plecoptera, stonefly, species list (1967) [California], Radford and Hartland- Rowe (1971) [Manitoba], Narf and Hilsen- hoff (1974) [Wisconsin], Kerst and Ander- son (1974) [Oregon], Ellis (1975) [Alaska], and Friesen et al. (1984) [Manitoba]. This investigation is part of a larger long- term documentation of the insect fauna of relatively unpolluted Big Hunting Creek. Specific aquatic species can be used as in- dicators of water quality (Hynes 1970). Hence, it is important to know not only which species are present in the stream, but also to understand the sequence in which they emerge, their life histories, and their ecological requirements. As stream char- acteristics change due to the encroachment of man, so do the components of benthic communities. Species data bases thus be- come important to document stream changes. Stupy AREA Big Hunting Creek (39°37'N, 77°27'W) originates in the Catoctin Mountains in VOLUME 92, NUMBER 1 western Frederick County, Maryland, and runs into the Monocacy River. This general area of Frederick County receives 76-102 cm of rain per year with approximately 102 cm of snow in the winter. The stream is bordered by Catoctin Mountain Park to the north and Cunningham Falls State Park to the south. In Cunningham Falls State Park the stream flows into and out of the 17.4 hectare Cunningham Lake. Minimum flow from the lake in the late summer and fall is reported to be 0.045 m?*. During the winter months the stream occasionally freezes over (Frederick County Planning and Zoning Commission 1969). The stream bed is composed of a range of particle sizes from small rocks and gravel to medium size boulders. Several sections of the stream bed exhibit bedrock outcrops. There are approximately 22 different genera of trees represented in both Catoctin Moun- tain Park and Cunningham Falls State Park (Buchart and Horn Consulting Engineers and Planners 1964). Dominant species of the drainage basin include those in Quercus (oak), Carya (hickory), Acer (maple), and Liriodendron (poplar). The majority of the trees are second and third growth hard- woods averaging 20-30 cm in diameter. MATERIALS AND METHODS The field work was carried out on Big Hunting Creek over a four year period from February, 1984, through September, 1988. The data are based primarily on adult spec- imens. Most of the collecting was in the areas adjoining the stream between the east- ern entrance of Catoctin Mountain Park on Maryland Route 77 and Hunting Creek Lake and above Cunningham Falls to the west entrance of Catoctin Mountain Park. With- in the study area, several intermittent streams run into Big Hunting Creek. No effort was made to collect along these feeder streams. Many of the specimens were hand collected at the Camp Pineil Bridge on Maryland Route 77. Additional material was collected along the edges of the stream on 121 rocks as well as on vegetation. Several of the smaller species were collected using a mouth-operated aspirator. Specimens were also obtained from stom- ach samples of either rainbow or brown trout caught in the study area of Big Hunting Creek and provided by local fishermen. The stom- ach pump consists of a tube approximately 0.8 cm (O.D.) by 20 cm with a rubber squeeze bulb attached to the end. To obtain a sample of the stomach contents, the tube was placed into the mouth of the trout and gently forced down the esophagus. After in- sertion, a small volume of water (1 ml de- pending upon the size of the fish) was slowly released into the trout’s stomach and drawn back up into the stomach pump. The stom- ach pump was then withdrawn and the fish released unharmed. As a general rule fish under 25-28 cm were not pumped. The sample was then placed in a vial, the water poured off and 70% ethanol added. All collected material was preserved in 70% ethanol. Each sample was labelled with collection data and an accession number. The material studied was stored in the au- thors’ collections, either in the Department of Zoology at Howard University or in the Department of Biology at the University of Tennessee at Chattanooga. Representative material has been deposited at Catoctin Mountain Park. RESULTS AND DISCUSSION Figure | lists the species of stoneflies and the time periods during which adults were collected. A total of 4286 specimens were collected during the course of the study. The number of specimens identified as well as the number of different collections for each species are also listed. The latter usually rep- resented different collecting dates but oc- casionally, if separate collections were made in different sections of the park on the same date, they were labelled as separate collec- tions. Altogether thirty-nine species were col- lected, representing 22 genera. Adults oc- J F M A M J 123412341234123412341234123412341234123412341234 Allocapnia recta Allocapnia nivicola Taeniopteryx maura Paracapnia angulata Allocapnia ricken Prostoia similis Oemopteryx contorta Paraleuctra sara Taenionema atlanicum Allocapnia granulata Pteronarcys biloba Paranemoura perfecta Soyedina carolinensis Leuctra ferruginea Leuctra sibleyi Amphinemura nigritta Sweltsa onkos Acroneuria carolinensis Pteronarcys proteus Ostrocerca truncata Remenus bilobatus Tallaperla elisa J PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A s 10) N D NO. OF NO. OF SPECIMENS COLLECTIONS pais 575 44 —— 1291 58 246 26 16 7 62 rt 67 12 3 3 104 17 67 22 1 1 2 2 3 1 4 3 = 146 28 ————— 46 21 eS 765 32 ———. 13 7 = 3 3 = 11 7 — 8 2 as 9 7 ——. 60 19 123412341234123412341234123412341234123412341234 J F M A M J Fig. 1. 1988. curred during every month of the year, but the majority of species were collected from April through June. Several genera were represented by more than one species, including Leuctra (six), Allocapnia and Isoperla (five), Acroneuria, Amphinemura, Ostrocerca, and Pteronarcys (2). Fifteen genera were each represented by one species, eight of those being represented by a single specimen. The five most abundant species in de- scending order were A/llocapnia_nivicola (Fitch) [1291], Amphinemura nigritta (Pro- vancher) [765], Allocapnia recta (Claassen) [575], Leuctra tenuis Pictet [550], and Tae- niopteryx maura (Pictet) [246]. These five species represented 80% of all the material J Inclusive collection dates for adult stoneflies based on collections from February 1984 to September A Ss oO N D collected. The most abundant species col- lected, A. nivicola, represented approxi- mately one-third of all the material collect- ed. Three of the five most abundant species emerged in the winter or early spring and were present for long periods. This may ac- count for the larger number of specimens collected. Several species represented by multiple collections over several months and by large numbers of individuals indicated that males were present in greater numbers than fe- males at the beginning of the emergence pe- riod. Species exhibiting this pattern include Allocapnia nivicola, A. recta and A. rickeri. Table | lists the ten earliest collections for A. nivicola. Chi-square analysis using the VOLUME 92, NUMBER 1 Jy oF eM SA PUM 4 123412341234123412341234123412341234123412341234 Ostrocerca albidipennis Leuctra tenella - Isoperla burksi Leuctra duplicata Eccoptura xanthenes Isoperla similis Acroneuria abnormis Isoperla sp. Isoperla holochlora Paragnetina media Diploperla duplicata Leuctra carolinensis Isoperla gibbsae Amphinemura wut Leuctra tenuis Perlesta placida Allocapnia aurora 123 J A Ss Oo N OD NO. OF NO. OF SPECIMENS COLLECTIONS 7 5 1 1 = 1 1 5 1 1 = 1 1 = 7 4 — 63 20 — 10 5 oe eens 88 17 5 1 1 = 1 1 = 4 1 = 9 4 a 9 3 — 550 26 = 1 1 = 31 1 TOTAL 4286 123412341234123412341234123412341234123412341234 J F M A M J Species Beginning Emergence 0 2 3) 2) 17 5 Total Species 2 4 9 14 22 17 Total New Emergences 0 2 i 12 29 34 total number from these collections indi- catesa significant deviation from the expected 1:1 ratio of males to females (P = 0.0001). Slightly earlier male emergence patterns for adult stoneflies have been reported by a number of workers including Brink (1949), Sheldon and Jewett (1967), Harper and Pi- lon (1970), Nebeker (1971), Neves (1978) and Masteller (1983). In contrast, other species in our study also represented by multiple collections and large individual numbers did not show a dominance of males at the beginning of the emergence period. These included Leuctra ferruginea, L. ten- uis, Amphinemura nigritta and Taeniopter- yx maura. Seventeen species were obtained from ap- proximately forty separate stomach pump samples. Two species, A//ocapnia granulata and Paranemoura perfecta, were added to J A Ss oO N D 0 2 0 0 2 1 the species list based on specimens obtained solely from this sampling technique. 4//o- capnia granulata was represented by a sin- gle specimen and P. perfecta was represent- ed by four specimens (2 males, | female, and | nymph). Stomach pump samples also provided a major portion of the specimens for two additional species, Soyvedina caro- linensis and Oemopteryx contorta. Of the four adult specimens recorded for S. caro- linensis, two stomach pump samples each provided a single adult specimen. Of the twelve specimens recorded for O. contorta, three adults were collected by hand and a total of nine nymphs were recovered from three separate stomach pump samples. An in depth presentation of the stomach pump data is being published separately. There are a number of variables which must be considered when making conclu- 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Partial collection data for Allocapnia nivi- cola: The ten earliest collections. Date Males Females Nov. 22, 1984 Nov. 24, 1984 Nov. 30, 1984 Dec. 9, 1984 Dec. 26, 1986 Dec. 8, 1987 Dec. 13, 1987 Dec. 24, 1987 Dec. 24, 1987 Dec. 24, 1987 Totals aS Ae DAWN Wwe nb ~ lon in nn sions about emergence based on adult col- lections. The total number of specimens col- lected is not necessarily a correct indication of the abundance of a particular species in the stream. Some species emerge by crawl- ing out of the water on rocks, others fly directly off the surface of the water to resting spots in the tops of trees. Secondly, the adults of some species are available for several days to a week while others are available for sev- eral months. Harper and Pilon (1970) report two types of emergence patterns: a short synchronous period where 90% of the population emerges within several days, and an extended emer- gence where 50% of the population emerges half way through the emergence period. A/- locapnia nivicola and Leuctra ferruginea are examples of species with extended emer- gences. Due to this long period of adult availability, specimens are collected more frequently when making random collec- tions. The periods that adults are present for those species with extended emergence pe- riods should not be misinterpreted. The greater longevity of females over males has been reported by Nebeker (1971), Finni (1975) and Lillehammer (1975). Although adult specimens were being collected, the actual emergence of adults may well have been completed. Thus, adult emergence pe- riod should not be confused with adult life span or adult duration. The data reported in Fig. 1 represents initial emergence dates and reports the duration that adults were detected in the vicinity of the stream. Stark et al. (1986) reported 33 species of Plecoptera from Maryland. During this study we collected an additional 25 new state records which are indicated by an * in the updated list of 58 species that follows. Order Plecoptera Suborder Arctoperlaria Group Euholognatha Superfamily Nemouroidea Family Taeniopterygidae Subfamily Taeniopteryginae Taeniopteryx burksi Ricker and Ross T. lonicera Ricker and Ross T. maura (Pictet) Subfamily Brachypterinae *Oemopteryx contorta (Needham and Claas- sen) Taenionema atlanticum Ricker and Ross Family Nemouridae Subfamily Amphinemurinae *4Amphinemura nigritta (Provancher) *4. wui (Claassen) Subfamily Nemourinae *Ostrocerca albidipennis (Walker) O. truncata (Claassen) *Paranemoura perfecta (Walker) *Prostoia similis (Hayen) Shipsa rotunda (Claassen) * Sovedina carolinensis (Claassen) Family Leuctridae Subfamily Leuctrinae *Leuctra carolinensis Claassen *L. duplicata Claassen *L. ferruginea Walker L. sibleyii Claassen *L. tenella Provancher *L. tenuis Pictet *Paraleuctra sara (Claassen) VOLUME 92, NUMBER 1 Family Capniidae Allocapnia aurora Ricker A. curiosa Frison . granulata (Claassen) . maria Hanson . nivicola (Fitch) . pygmaea (Burmeister) . recta (Claassen) . rickeri Frison . vivipara (Claassen) . wrayl Ross Paracapnia angulata Hanson mime ww ww Pw Group Systellognatha Superfamily Pteronarcyoidea Family Pteronarcyidae *Pteronarcys biloba Newman *P. proteus Newman Superfamily Peltoperloidea Family Peltoperlidae Subfamily Peltoperlinae *Tallaperla elisa Stark T. maria (Needham and Smith) Superfamily Perloidea Family Perlodidae Subfamily Isoperlinae Clioperla clio (Newman) *Tsoperla burksi Frison *7. gibbsae Harper I. holochlora (Klapalek) I. similis (Hagen) Subfamily Perlodinae *Diploperla duplicata (Banks) Isogenoides hansoni (Ricker) *Remenus bilobatus (Needham and Claas- sen) Family Perlidae Subfamily Acroneurinae Acroneuria abnormis (Newman) A. arenosa (Pictet) * 4. carolinensis (Banks) A, filicis Frison Attaneuria ruralis (Hagen) Eccoptura xanthenes (Newman) *Perlesta placida (Hagen) ‘complex’ Perlinella drymo (Newman) P. ephyre (Newman) Subfamily Perlinae Agnetina annulipes Stark A. flavescens (Walsh) *Paragnetina media (Walker) Family Chloroperlidae Subfamily Chloroperlinae Alloperla atlantica Baumann Haploperla brevis (Banks) *Sweltsa onkos (Ricker) ACKNOWLEDGMENTS We thank Mr. Tom McFadden (super- intendent) and Jim Voigt of Catoctin Moun- tain Park (National Park Service) for per- mission and help in carrying out these studies. We would like to thank Jim Gilford for loaning background literature on the Big Hunting Creek drainage basin. We would also like to thank a number of individuals who over the course of the study occasion- ally helped collect specimens. They include Shawn Bowen, Eric Riddick, Don Dunbar, Andrew Maglott, Mark Stolt, and George Middendorf. Lastly, we thank a number of local fishermen for generously providing stomach pump samples from trout caught from Big Hunting Creek and released un- harmed. Manuscript preparation was com- pleted while the senior author was a visiting professor in the Department of Entomolo- gy, Cornell University, Ithaca, New York. LITERATURE CITED Brink, P. 1949. Studies on Swedish stoneflies (Ple- coptera). Opusc. Ent. Suppl. 11: 1-250. Buchart and Horn Consulting Engineers and Planners. 1964. Master recreational development plan for Cunningham Falls State Park, Frederick Co., MD. 19 pp. Ellis, J. R. 1975. Seasonal abundance and distribu- tion of adult stoneflies of Sashin Creek, Baranof Island, southeastern Alaska. Pan-Pac. Ent. 51: 23- 30. Finni, G. R. 1975. Feeding and longevity of the win- ter stonefly A//ocapnia granulata (Claassen) (Ple- 126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON coptera: Capniidae). Ann. Entomol. Soc. Am. 68: 207-208. Frederick County Planning and Zoning Commission. 1969. Natural physical characteristics, a back- ground study. 140 pp. Freisen, M. K., J. F. Flannagan, and D.G. Cobb. 1984. Emergence of stoneflies (Plecoptera) from South Duck River and Cowan Creek in the Duck Moun- tain region, Manitoba. Amer. Midl. Nat. 111: 69- 80. Harper, P. P. and E. Magnin. 1969. Cycles vitaux de quelques Plécopteres des Laurentides (insectes). Can. J. Zool. 47: 483-494. Harper P. P. and J. G. Pilon. 1970. Annual patterns of emergence of some Quebec stoneflies (Insecta: Plecoptera). Can. J. Zool. 48: 681-694. Hynes, H. B. N. 1970. The Ecology of Running Waters. Univ. of Toronto Press, Toronto. 555 pp. Kerst, C. D. and N. H. Anderson. 1974. Emergence patterns of Plecoptera in a stream in Oregon, USA. Freshw. Biol. 4: 205-212. Lillehammer, A. 1975. Norwegian stoneflies. III. Field studies on ecological factors influencing distribu- tion. Nor. Entomol. Tidskr. 22: 71-80. Masteller, E.C. 1983. Emergence phenology of Ple- coptera from Sixmile Creek, Erie County, Penn- sylvania, USA. Aquatic Insects 5: 1-8. Narf, R. P. and W. L. Hilsenhoff. 1974. Emergence pattern of stoneflies (Plecoptera) in Otter Creek, Wisconsin. Great Lakes Entomol. 7: 117-125. Nebeker, A. V. 1971. Effect of high winter water tem- peratures on adult emergence of aquatic insects. Water Res. 5: 777-783. Neves, R. J. 1978. Seasonal succession and diversity of stoneflies (Plecoptera) in Factory Brook, Mas- sachusetts. J. N. Y. Ent. So. 86: 144-152. Radford, D. S. and R. Hartland-Rowe. 1971. Emer- gence patterns of some Plecoptera in two moun- tain streams in Alberta. Can. J. Zool. 49: 657-662. Sheldon, A. L. and S. G. Jewett, Jr. 1967. Stonefly emergence in a Sierra Nevada stream. Pan-Pac. Ent. 43: 1-8. Singh, M. P.,S. M. Smith, and A. D. Harrison. 1984. Emergence patterns of the stoneflies (Plecoptera) from a wooded stream in southern Ontario. Aquatic Insects 6: 233-243. Stark, B. P. 1980. Plecoptera, pp. 85-86. Jn Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey, eds., Aquatic Insects of the Upper Three Rivers Creek, Savannah River Project, South Car- olina. Part I. Orders other than Diptera. J. Ga. Ent. Soc. 15: 73-101. Stark, B. P., S. W. Szcezytko, and R. W. Baumann. 1986. North American stoneflies (Plecoptera): Systematics, distribution, and taxonomic refer- ences. Great Basin Nat. 46: 383-397. Tkac, M. A. and B. A. Foote. 1978. Annotated list of stoneflies (Plecoptera) from Stebbins Gulch in northeastern Ohio. Great Lakes Entomol. | 1: 139- 142. White, D. 1974. The distribution of stoneflies (In- secta: Plecoptera) of the Salt River, Kentucky. Trans. K. Acad. Sci. 35: 17-23. Woodall, W. R., Jr. and J. B. Wallace. 1972. The benthic fauna in four small southern Appalachian streams. Amer. Midl. Nat. 88: 393-407. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 127-134 NEW SPECIES OF PREDACEOUS MIDGES OF THE TRIBE CERATOPOGONINI FROM SUBANTARCTIC ARGENTINA (DIPTERA: CERATOPOGONIDAE) GuSTAVO R. SPINELLI AND WILLIAM L. GROGAN, JR. (GRS) Instituto de Limnologia “Dr. Raul A. Ringuelet,” Casilla de Correo 712, 1900 La Plata, Argentina; (WLG) Department of Biological Sciences, Salisbury State University, Salisbury, Maryland 21801. Abstract. —Four new species of predaceous midges of the tribe Ceratopogonini from subantarctic Argentina are described and illustrated: Diaphanobezzia patagonica, Macru- rohelea fuscipennis, M. similis, and Notiohelea pilosa. A key to the Neotropical species of Macrurohelea is presented. Key Words: gentina, predaceous midges Wirth and Grogan (1988) listed five gen- era of predaceous midges of the tribe Cer- atopogonini whose distributions are exclu- sively subantarctic or nearly so: Austrohelea Wirth and Grogan, with 7 species, 6 are from Australia, Campbell Island, New Zea- land and Tasmania, and one species from southern Argentina; Diaphanobezzia In- gram and Macfie, with 2 species from Ar- gentina; Isthmohelea Ingram and Macfie, with one species from Chile; Macrurohelea Ingram and Macfie, with 12 species, 9 from Argentina and Chile and 3 from Australia; and Notiohelea Grogan and Wirth, with one species from Chile. The purpose of this paper is to describe 4 new species in the tribe Ceratopogonini belonging to 3 rarely collected genera, that were recently taken by GRS in the temper- ate subantarctic region of southwestern Ar- gentina. The types of these new species are deposited in the collection of the Museo de La Plata, La Plata, Argentina (MLP); para- types of Macrurohelea fuscipennis and No- tiohelea pilosa will be deposited in the Na- Diptera, Ceratopogonidae, Ceratopogonini, Neotropical, subantarctic, Ar- tional Museum of Natural History, Washington, D.C. (USNM). In addition, a key to the Neotropical species of Macru- rohelea is provided which includes the fe- male of M. paracaudata Grogan and Wirth (1980) that was recently described by Spi- nelli (1987). For an explanation of general ceratopo- gonid terminology, see Downes and Wirth (1981); for special terms dealing with genera in the tribe Ceratopogonini, see Wirth and Grogan (1988), Grogan and Wirth (1979), and Spinelli and Grogan (1984). Diaphanobezzia patagonica Spinelli and Grogan, NEw SPECIES Fig. 1 Diagnosis. — Distinguished from females of Diaphanobezzia spinellii Wirth and Gro- gan (1988) by its sparsely pubescent eyes, longer 4th palpal segment, and wing with longer costa and R4+5. Holotype female.—Head: Dark brown. Eyes broadly separated above, sparsely pu- 128 bescent between the lower ommatidia. An- tennal scape pale brown, bearing 3 pairs of setae; pedicel dark brown; flagellum (Fig. la) dark brown, lengths of flagellomeres in proportion of 11-7-7-7-7-6-6-7-8-8-9-10- 14; antennal ratio 0.84; flagellomeres 1-8 bearing a pair of trichoid sensilla. Palpus (Fig. 1b) uniformly dark brown; segment lengths in proportion of 3-10-14-15; palpal ratio 3.5. Mandible with 8 coarse teeth. Thorax: Dark brown. Scutum covered with strong setae, anterior spine absent, 8 prealar setae and | postalar seta; scutellum apparently with 6 similar sized bristles. Legs dark brown; tarsomeres 1+ of mid and hind legs slightly paler brown: femora and mid and hind tibiae armed apically with 1-2 long spines, hind tibia with 7-9 spines on exten- sor side; hind tibial comb with 6 spines; apices of tarsomeres I—3 and base of hind tarsomere | with a pair of ventral spines, ventral surface of mid tarsomere | with 3 pairs of widely spaced spines; hind tarsal ratio 2.3; claws small, evenly curved, equal sized with basal inner teeth. Wing (Fig. Ic) whitish hyaline; anterior veins brownish, distal '2 of costa and R4+5 dark brown, all other veins barely perceptible; r-m cross- vein interrupted at middle; cell RS without intercalary veins; costal finge dense: costal sections I-II-III in proportion of 28-25-9, R4+5 40; costal ratio 0.92; wing length 1.06 mm, breadth 0.40 mm. Halter (Fig. 1d) pale in color, sac-shaped, lacking a distinct con- striction below knob, as is typical of the genus. Abdomen: Dark brown. Sternites 3-5 en- tire, with a broad rounded caudal notch; sternite 8 (Fig. le) deeply notched caudally with broadly rounded posterior lobes; each arm of sternite 9 (Fig. le) slightly hooked; sternite 10 (Fig. le) with 2 pairs of setae, the posterior pair longer, cerci short. Sper- mathecae (Fig. le) subequal, slightly ovoid with short necks, each about 0.053 mm in diameter; a small vestigial 3rd spermatheca present. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Type material. — Holotype female labeled “Argentina, Santa Cruz, Rio Pinturas, 15- I-1988, G. R. Spinelli, entomological net,” (MLP). Distribution. — Argentina. Etymology.—The specific epithet, pata- gonica, refers to the southernmost region of Argentina where the type was collected. Remarks.—Females of Diaphanobezzia spinellii Wirth and Grogan (1988) differ from those of D. patagonica n. sp. by their densely pubescent eyes, shorter 4th palpal segment that is less than '2 the length of the 3rd palpal segment, its wing with shorter costa (costal ratio 0.82) and R4+5, complete r-m crossvein, and different costal proportions. Diaphanobezzia pellucida Ingram and Macfie (1931) is known only from a single male but it differs from D. patagonica n. sp. by its larger size (wing length 1.7 mm), hav- ing intercalary veins in cell R5, and its r-m crossvein is complete. This new species, known only from the single female holotype, was collected in the Valley of the Rio Pinturas located in the Patagonian steppe (47°S), approximately 150 km E of the subantarctic forest. Macrurohelea fuscipennis Spinelli and Grogan, NEw SPECIES Fig. 2 Diagnosis. — Distinguished from females of other species in the genus by its bare eyes, wing darkly infuscated on distal *4, proxi- mal '3 hyaline, and spermathecae with long slender necks. Female.— Head: Yellowish brown. Eyes bare, nearly contiguous. Antennal scape yel- low, bearing 2 pairs of setae; pedicel dark brown; flagellum (Fig. 2a) brownish; lengths of flagellomeres in proportion of 30-18-17- 17-18-18-19-20-20-20-22-25-32; antennal ratio 0.78 (0.75-0.81, n = 3); flagellomere 1 with 4 sensilla coeloconica, flagellomeres 1-8 with a pair of trichoid sensilla. Palpus (Fig. 2b) yellowish; lengths of segments in VOLUME 92, NUMBER 1 CIOBEESESSO SOE’ SD Fig. 1. Scale bars = 0.1 mm. proportion of 10-15-16-10-18; segment 3 with small sensory pit; palpal ratio 1.70 (1.40-2.00, n = 4). Mandible (Fig. 2c) in- fuscated at apex, with 8 teeth. Thorax: Yellowish, humeral pits infus- cated, prealar areas dark brown. Scutum (Fig. 2d) covered with fine pubescence, 4 prealar setae, | postalar seta; scutellum (Fig. 2d) with 3-5 bristles; postscutellum dark brown. Legs yellowish, tarsomeres 4—5 a Diaphanobezzia patagonica. a, flagellum; b, palpus; c, wing; d, halter; e, distal portion of abodomen. slightly infuscated; hind tibial comb with 6 spines; hind tarsal ratio 2.1 (n = 4); pali- sade setae on tarsomere | of fore and hind legs; tarsomere 4 cordiform; tarsomere 5 of fore leg 4x longer than broad, 3 longer than broad on mid and hind legs; claws small, equal sized without basal inner teeth. Wing (Fig. 2e) with proximal '3 whitish hya- line, veins yellowish, distal 7; with mem- brane and veins darkly infuscated; 2 radial 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. cells, 2nd 1.5 x longer than Ist; cell RS with- out intercalary veins; costal ratio 0.69 (0.68- 0.70, n = 4); wing length 1.18 (1.10-1.26, n = 4) mn, breadth 0.47 (0.45-0.54, n = 4) mm. Halter whitish. Abdomen: Brown. Sternites 2-6 (Fig. 2f) divided by broad unsclerotized area; ster- nites 7-8 (Fig. 2f) entire; each arm of ster- nite 9 (Fig. 2f) slender with truncate tip; Macrurohelea fuscipennis. a, flagellum; b, palpus; c, mandible; d, scutum and scutellum; e, wing; f, distal portion of abodomen. Scale bars = 0.1 mm (a, d-f), 0.75 mm (b), 0.02 mm (c). sternite 10 (Fig. 2f) with a pair of apical setae, cerci short; segments 9 and 10 elon- gated and bent forward ventrally as 1s typ- ical for the genus. Spermathecae (Fig. 2f) subequal, spheroid, with long slender necks, each 0.034 mm in diameter and the neck 0.019 mm long; a small 3rd vestigial sper- matheca present. Type material.—Holotype female, 3 fe- VOLUME 92, NUMBER 1 male paratypes labeled “Argentina, Chubut, Parque Nacional Los Alerces, ‘El Alerzal,’ 22-I-1988, G. R. Spinelli, entomological net,” (MLP). Distribution. — Argentina. Etymology.—The specific epithet, fusci- pennis, refers to the distinctive wing of this species that is infuscated on its distal por- tion. Remarks.—The wing that is darkly in- fuscated on its distal 73 and hyaline on its proximal ' readily distinguishes this species from all other species of Macrurohelea. In addition, all other species of Macrurohelea (13 species presently known) have pubes- cent eyes and further differ from M. fusci- pennis n. sp. in having spermathecae with shorter necks. Macrurohelea dycei Grogan and Wirth (1985) from Australia is the only other species that has spermathecal necks nearly as long, but its spermathecae are ovoid with moderately tapering necks. See the key to the Neotropical species of Ma- crurohelea below for further ways that this new species differs from others in the genus. This new species 1s presently known from only 4 females which were collected in the temperate subantarctic forest of Argentina. Macrurohelea similis Spinelli and Grogan, NEw SPECIES Fig. 3a, b Diagnosis. — Distinguished from males of other species of Macrurohelea by the fol- lowing combination of characters: small size (wing length 0.95 mm), wing without inter- calary veins in cell R5, gonostylus extending Ys of its length beyond tergite 9. Holotype male. — Head: Dark brown. Eyes pubescent, nearly contiguous. Antenna with pale scape; pedicel dark brown; flagellum and plume brown, flagellomeres distinctly separated with lengths in proportion of 20- 8-7-7-7-7-6-6-6-7-12-15-9; flagellomere 1 with 2 sensilla coeloconica; antennal ratio 0.44. Palpus brown; segment lengths in pro- 131 portion of 5-8-9-6-11; segment 3 with well defined sensory pit. Thorax: Dark brown. Scutum with 4 prealar setae and | postalar seta; scutellum with 3 large and 2 small bristles. Legs uni- formly brown; hind tibial comb with 6 spines; hind tarsal ratio 2.2: palisade setae on fore and mid tarsomere |: claws small, equal sized with bifid tips. Wing with mem- brane slightly infuscated, veins brown; Ist radial cell subequal to 2nd; cell R5 without intercalary veins; costal ratio 0.66; wing length 0.95 mm, breadth 0.36 mm. Halter stem brown; knob white. Abdomen: Brown. Genitalia as in Fig. 3a, b. Sternite 9 moderately long, twice as broad as long, tapered basally, posterior margin straight; tergite 9 moderately short, trian- gular, gradually tapering distally with a quadrate posterolateral extension and slen- der pubescent cerci which are divergent and extend beyond tip of quadrate extensions. Gonocoxite slightly curved, 2.5 = longer than broad with a small mesobasal lobe; gonostylus shorter than gonocoxite, greatly curved, gradually tapering to a slender pointed tip. Aedeagus short, triangular, slightly broader than long, basal arch about + of total length; basal arm nearly straight, heavily sclerotized; distal portion tapering toa moderately pointed tip. Parameres (Fig. 3b) heavily sclerotized, separate; basal arm short, broad basally, pointed apically; distal portions slender, closely approximated proximally, divergent distally, tapering near apex to a slightly bent pointed tip. Type material.—Holotype male labeled “Argentina, Rio Negro, Parque Nacional Nahuel Huapi, camino a cascada de los Al- erces, 24-I-1988, G. R. Spinelli, entomo- logical net,” (MLP). Distribution. — Argentina. Etymology.—The specific epithet, similis, is a reference to the similarity of the geni- talia of this new species to those of Macru- rohelea caudata Ingram and Macfie (1931). Remarks.—Macrurohelea similis n. sp. most closely resembles M/. caudata Ingram PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Macrurohelea similis, a,b, and Notiohelea pilosa, c-f. a, genitalia (parameres removed); b, parameres; c, flagellum; d, palpus; e, wing; f, spermathecae. Scale bars = 0.1 mm (a-*, e), 0.05 mm (d), 0.025 mm (f). and Macfie (1931) by virtue of their similar male genitalia. However, M. caudata differs from this new species by several features of its genitalia such as, aedeagus broader dis- tally with a square tip, cerci that arise from ventral surface of tergite 9 but do not extend beyond tergite 9, gonocoxite that is straight, and parameres with different shaped basal arm and distal portions parallel. In addi- tion, M. caudata has eyes that are widely separated and bare except on extreme inner margins, and the 3rd palpal segment is lon- ger than Sth (Sth segment longer than 3rd in M. similis n. sp.). See the key to the Neo- tropical species of Macrurohelea below for further ways in which this new species dif- fers from other species in the genus. This new species, which is presently known only from the male holotype, was collected in the temperate subantartic forest of Argentina. KEY TO THE NEOTROPICAL SPECIES OF M4CRUROHELEA Nod REE Goosonadsonbeodosanie bf Sorc ante ae ee PD =JaMalest. £0 Sect cman So ee eer 11 25 Onexspermatheca" sae eee ae eee ee 3 —¥ diworspermathecacartense mer eet ieee 4 VOLUME 92, NUMBER | . Wing with intercalary veins in cell R5; wing membrane infuscated, veins dark brown Bia Sct 4 monotheca Spinelli and Grogan Wing without intercalary veins in cell R5; wing membrane whitish hyaline, veins pale ... See cmos gentilii Spinelli and Grogan : Wing with intercalary veins in cell RS : 5 Wing without intercalary veins incell RS .. 6 Second radial cell of wing 2.5 = longer than Ist, veins brown; antennal ratio 1.61 . ee wirthi Spinelli and Grogan Second radial cell of wing 3 x longer than Ist, veins pale; antennal ratio 1.00 ; Nyon ee ee caudata Ingram and Macfie Second radial cell of wing twice as longas Ist 7 Second radial cell of wing at least 3 = longer thanwlisteancwiasec sec ees mies.e eee 210 . Flagellum very short, flagellomeres 9- 12 broader than long, antennal ratio 0.59; very small species, wing length 0.94 mm ... mh Sas kuscheli Wirth Fiagelln longer, flagellomeres 9-12 twice as long as broad, antennal ratio 0.75—1.16; small species, wing length 1.02-1.42 mm . 8 Wing membrane hyaline, veins gray ... Been etna ecaereredsicts irwint Grogan and Wirth Wing membrane infuscated on at least distal 7s, veins brown Wing membrane infuscated on 1 distal 7/3, PrOxX- imal 3 hyaline; tips of claws pointed; sper- mathecae with very long necks fuscipennis, new species Wing membrane entirely infuscated; tips of claws bifid; spermathecae with shorter necks tk: . paracaudata Grogan and Wirth Flagellomeres 5-8 with apical sensilla coe- loconica; legs with inconspicuous setae; wing membrane and veins pale . 2 ee ars oak ee thoracica Ingram and Macfie Flagellomeres 5-8 without apical sensilla coeloconica; legs with numerous long bristly setae; wing membrane and veins infuscated dark brown setosa Wirth . Large species, wing length 2 2.1mmorgreater 12 Smaller species, wing length 1.5 mm or less 13 Legs with long bristly setae; wing membrane and veins infuscated dark brown setosa Wirth Legs with inconspicous setae; wing mem- brane and veins pale .... Esa thoracica Ingram and Macfie . Very small species, wing length 0.90-0.95 mm . BA nena ys eee een nies Saati 14 Small species, wing length 1.3 mm orgreater 15 . Sternite 9 with caudomedial notch; gono- stylus extends to apex of tergite 9 SEMAN erates ter ces paracaudata Grogan and Wirth Sternite 9 without caudomedial notch; gono- stylus extends ' of its total length beyond apex of tergite 9 _ similis, new species 133 15. Wing with intercalary veins in cell RS, 2nd radial cell subequal to 1st; aedeagus more or less crescent shaped : caudata Ingram and Macfie - Wing without intercalary veins in cell RS, 2nd radial cell 1.7—2.0 x longer than Ist; ae- deagus triangular ............. 16 16. Gonostylus bent abruptly subapically at 90°: sternite 9 with deep caudomedial excavation : gentilii Spinelli and Grogan ~ Gonostylus curved subapically, not bent at 90°; sternite 9 with shallow caudomedial ex- cavation irwint Grogan and Wirth Notiohelea pilosa Spinelli and Grogan, New SPECIES Fig. 3c-f Diagnosis. — Distinguished from its only known congener, N. chilensis Grogan and Wirth (1979), by its pubescent eyes, more darkly infuscated wing with the 2nd radial cell over 3 longer than Ist, shorter pro- boscis, more slender 3rd palpal segment, and ovoid spermathecae. Female. — Head: Brown. Eyes pubescent, narrowly separated. Antennal scape pale, bearing 3 pairs of setae; pedicel dark brown; flagellum (Fig. 3c) brown, lengths of flagel- lomeres in proportion of 22-13-14-14-14- 13-13-13-13-14-15-16-20; antennal ratio 0.71 (0.67—-0.74, n = 3); flagellomere | with 5 sensilla coeloconica, flagellomeres 1-8 bearing a pair of trichoid sensilla. Palpus (Fig. 3d) brown; segment lengths in pro- portion of 9-16-20-8-14; palpal ratio 2.10 (2.05-2.25, n = 3); segment 3 with a mod- erately broad apical sensory pit bearing 4— 5 capitate sensilla. Proboscis short, probos- cis/head ratio 0.42 (0.40-0.44, n = 2). Man- dible reduced, vestigial, without teeth. Thorax: Brown. Scutum densely covered with coarse setae, humeral pits present, no anterior spine, 3 prealar setae and | postalar seta; scutellum with 4 similar sized bristles and 6 smaller setae. Legs brown, nearly identical with those of N. chilensis in form and features; hind tarsal ratio 1.75 (n = 3). Wing (Fig. 3e) membrane darkly infuscated (infuscation not depicted in Fig. 3e), cov- ered with coarse microtrichia; veins brown, 134 veins RI and R4+5 with macrotrichia; 2 radial cells present, the 2nd 3.2 x longer than Ist. Halter brown. Abdomen: Brown. Moderately broad proximally, tapering distally at segment 5; sternite 10 with one pair of large setae; 2 ovoid, subequal spermathecae (Fig. 3f), each 0.045 mm long with neck and 0.03 mm wide. Type material. — Holotype female labeled “Argentina, Chubut, Parque Nacional Los Alerces, ‘El Alerzal,’ 22-I-1988, G. R. Spi- nelli, entomological net,” (MLP); 2 female paratypes, | with same data as holotype, | with same data as holotype except taken 23- I-1988 by CDC light trap. Distribution. — Argentina. Etymology.—The specific epithet, pilosa, is a reference to the coarse setae that cover the scutum of this species. Remarks.— The only other known species in the genus, Notiohelea chilensis Grogan and Wirth (1979), differs from N. pilosa n. sp. by its bare eyes, longer proboscis (pro- boscis/head ratio 0.56), broader 3rd palpal segment (palpal ratio 1.82), spherical sper- mathecae, and more lightly infuscated wing with the 2nd radial cell only twice as long asithe: Ist... This species is currently known only from female specimens that were collected in the temperate subantarctic forest of Argentina. ACKNOWLEDGMENTS This paper is Scientific Contribution no. 406 of the Instituto de Limnologia “Dr. Raul PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A. Ringuelet,”’ La Plata, Argentina. This re- search was supported by CONICET grant PID 3-074200/85 awarded to G. R. Spi- nelli. LITERATURE CITED Downes, J. A. and W. W. Wirth. 1981. Chapter 28. Ceratopogonidae, pp. 393-421. Jn McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 1, 674 pp. Agric. Canada Monogr. 27, Ottawa. Grogan, W. L., Jr. and W. W. Wirth. 1979. Notiohe- lea, a new genus of biting midges of the tribe Cer- atopogonini from Chile (Diptera: Ceratopogoni- dae). Pan-Pacific Entomol. 52(1978): 283-286. . 1980. Two new species of Macrurohelea from Chile with a key to the Neotropical species (Dip- tera: Ceratopogonidae). Pan-Pacific Entomol. 56: 137-143. 1985. Two new Australian species of Ma- crurohelea, with a description of the male of M. commoni (Diptera: Ceratopogonidae). Int. J. Entomol. 27: 93-100. Ingram, A. and J. W. S. Macfie. 1931. Ceratopogon- idae. Diptera of Patagonia and South Chile. Part II. Fasc. 4, pp. 155-232. British Museum (Nat. Hist.), London. Spinelli, G. R. 1987. Notas sobre Ceratopogonidae (Diptera, Nematocera) de la Republica Argentina. VI. Las hembras de Paradasyhelea brevipalpis y de Macrurohelea paracaudata. Limnobios 2: 667— 670. Spinelli, G. R. and W. L. Grogan, Jr. 1984. Three new species of Macrurohelea from Argentina with a key to the Neotropical species (Diptera: Cera- topogonidae). Proc. Entomol. Soc. Wash. 86: 961- 967. Wirth, W. W. and W. L. Grogan, Jr. 1988. The Pre- daceous Midges of the World (Diptera: Cerato- pogonidae; Tribe Ceratopogonin1). Flora and Fau- na Handbook no. 4. E. J. Brill, New York, xv + 160 pp. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 135-138 IDENTITY OF ENCARSIA SPP. (HYMENOPTERA: APHELINIDAE) INTRODUCED INTO WESTERN SAMOA FOR BIOLOGICAL CONTROL OF PSEUDAULACASPIS PENTAGONA (TARGIONI-TOZZETTI) (HEMIPTERA: DIASPIDIDAE) D. P. A. SANDs, R. BROE AND W. J. M. M. LIEBREGTS CSIRO, Division of Entomology, Long Pocket Laboratories, Private Bag No. 3, P.O. Indooroopilly, Queensland 4068, Australia. Abstract. —Cultures of Encarsia spp. thought to be E. berlesei (Howard) from the USA, France and Tonga, were imported into Western Samoa for biological control of Pseu- daulacaspis pentagona (Targioni-Tozzetti), a pest on passion fruit vines (Passiflora edulis var. flavicarpa Degener). Comparison of specimens from the cultures with the syntypes of E. berlesei and E. diaspidicola (Silvestri) revealed both species were present. Both species were released but only EF. diaspidicola proved to be an effective biological control agent. Key Words: trol, Western Samoa White peach scale, Pseudaulacaspis pen- tagona (Targioni-Tozzetti) 1s a serious pest on several unrelated plants in many coun- tries (Clausen et al. 1978). In Western Sa- moa P. pentagona is a recently-introduced pest of passionfruit (Passiflora edulis var. flavicarpa (Degener), where it has contrib- uted to a reduction of 43% in the value of pulp produced between 1984 and 1986 (Pe- ters et al. 1985, Rasch 1986). The aphelinid parasitoid Encarsia berle- sei (Howard) has been recorded as an effec- tive biological control agent for P. penta- gona in several countries (Clausen et al. 1978). Unlike E. berlesei, the closely-related E. diaspidicola (Silvestri) has not been rec- ognized as an effective biological control agent (Greathead 1971, Waterhouse and Norris 1987). Moreover, E. diaspidicola has been considered a synonym of E. berlesei by Flanders (1960) and Greathead (1976) despite characteristics noted by Silvestri Encarsia, diaspidicola, berlesei, Pseudaulacaspis, Passiflora, biological con- (1930) that enabled its separation from LE. berlesei. In 1986, three cultures of Encarsia spp., all presumed to be E. berlesei, were im- ported into Western Samoa from the USA, France and Tonga for biological control of P. pentagona. One species, subsequently identified as E. diaspidicola, became an ef- fective biological control agent for this scale insect (Liebregts et al. 1989). Observed dif- ferences between specimens from the USA and those from the other two localities, and difficulties with their identification, prompted a comparison of specimens from all cultures imported in Western Samoa with type specimens of E. berlesei and E. dias- pidicola. MATERIALS AND METHODS The following were examined: E. berlesei: Six syntypes on slide bearing two labels (white)—‘*9942, Prospaltella ber- 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Comparative forewing measurements of Encarsia spp. No. of Measurement* Measurement? Specimens Examined x Range x Range E. berlesei Washington, D.C., USA (syntypes) 5 0.12 0.11-0.12 0.37 0.35-0.37 Gainsville, Florida, USA 6 0.10 0.09-0.12 0.37 0.36-0.37 Upolu, W. Samoa (field recoveries) 3 0.12 0.11-0.12 0.36 0.36-0.37 E. diaspidicola Capetown, South Africa (syntypes) 4 0.21 0.20-0.22 0.31 0.30-0.31 France (CIBC culture) 7 0.21 0.20-0.22 0.32 0.30-0.32 Tongatapu, Tonga 5 0.21 0.20-0.22 0.32 0.31-0.32 Upolu, W. Samoa (field recoveries) 1/ 0.22 0.21-0.23 0.32 0.31-0.32 * Maximum fringe length/wing length. > Maximum wing width/wing length. lesei How. on lilac, Washington, D.C., E. R. Sassocei, 2 June 1906”; (red) “Type No. 9942 U.S.N.M.” in U.S. National Museum, Washington, D.C., USA. E. diaspidicola: Nine syntypes on slide bearing two labels (white)—‘*Prospaltella diaspidicola Silv. (Cotypes) Encarsia dias- pidicola (Silv.) Capetown, S. Afr.”; (red) “Cotype No. 41387 U.S.N.M” in U.S. Na- tional Museum, Washington, D.C., USA and four specimens on slide bearing one label ‘‘Prospaltella diaspidicola 10, 1% ge- neraz. italia Portici—vil. 1909” in the Uni- versity of Naples, Italy. Specimens of Encarsia spp., cultures from Gainsville, Florida, USA (R. I. Sailer), France (CIBC), Tonga (field collected) and Western Samoa (field collected after estab- lishment) were slide mounted for compar- ison with the syntypes of FE. berlesei and E. diaspidicola. All cultures were uniparental. RESULTS AND DISCUSSION We have compared the syntypes of E. ber- lesei and E. diaspidicola with specimens from the three cultures introduced into Western Samoa (Table 1). Despite slight dif- ferences observed (in forewing fringe length) between material from Gainsville, Florida, USA and the syntypes of E. berlesei from Washington, D.C., USA, we conclude that the two are conspecific. We also conclude that materials obtained from France and from Tonga are conspecific with the syn- types of EF. diaspidicola from South Africa. We have had no difficulty in distinguish- ing the two Encarsia spp., based on Silves- tri’s (1909) original description of E. dias- pidicola, his (1930) subsequent comparison of the two species and the differences noted by Gahan (1925). In addition to forewing measurements (Table 1), we have found the following characteristics most useful for dis- tinguishing the two species: the outer fore- wing margin of EF. diaspidicola (Fig. 1) is more narrowly rounded than E. berlesei (Fig. 2) and there is an area almost free of setae close to the outer margin in E. diaspidicola, whereas on the forewing of FE. berlesei the discal setae extend uninterrupted to the marginal fringe. Funicle segment 2 of the antenna of EF. diaspidicola (Fig. 3) is clearly longer than segment | whereas in E. berlesei (Fig. 4) segments | and 2 are subequal. We consider that the identification of these two parasitoids of P. pentagona may have been confused in biological control pro- grams. Flanders (1960) considered EF. dias- pidicola to be asynonym of E. berlesei with- out explanation and Greathead (1971) stated that the former was unable to control P. pentagona in Europe. In view of the success of Tongan and French E. diaspidicola in controlling P. pentagona in Western Samoa, VOLUME 92, NUMBER 1 137 0.2mm Figs. 1-4. Encarsia spp. 1, 2, Forewings of syntypes: E. diaspidicola (Silvestri) and E. berlesei (Howard), respectively. 3, Antenna of EF. diaspidicola (Tonga). 4, Antenna of E. berlesei (Florida, USA). Arrows indicate areas clear of setae (1, 2) and second funicle segment (3, 4). some examples attributed to E. berlesei may actually refer to E. diaspidicola. A taxonom- ic re-assessment of the Encarsia spp. from biological control programs is clearly war- ranted. In Western Samoa, FE. diaspidicola has maintained biological control of P. penta- gona since its establishment in 1986 (Lie- bregts et al. 1989). Apart from early field recoveries over a five month period follow- ing its release, we have no further evidence for establishment of E. berlesei. ACKNOWLEDGMENTS We are grateful to the late Professor R. I. Sailer, the CAB International Institute of Biological Control and Dr. C. Benassy, In- stitut National de la Récherche Agrono- mique, France for the cultures and speci- mens of Encarsia spp. used in this study and to Professor G. Viggiani, Institute of Agricultural Entomology, University of Na- ples, Italy and Dr. E. Grissell, Systematic Entomology Laboratory, U.S. National Museum, Washington, D.C. for loans of type material. This study was supported by the Australian Centre for International Agri- cultural Research. LITERATURE CITED Clausen, C. P. (ed.), B. R. Bartlett, P. DeBach, R. G. Goeden, E. F. Leger, J. A. McMurtry, E. R. Oat- man, E. C. Bay, and D. Rosen. 1978. Introduced parasites and predators of arthropod pests and weeds: A world review. Agric. Handbook U.S. Dept. Agric. 480: 545 pp. Flanders, S. E. 1960. The status of San Jose scale parasitization (including biological notes). J. Econ. Ent. 53: 757-759. Gahan, A.B. 1925. Some new parasitic Hymenoptera with notes on several described forms. Proc. U.S. Natl. Mus. 65: 1-23. 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Greathead, D. J. (ed.). 1971. A review of biological control in the Ethiopian region. Tech. Comm. 5. Commonwealth Agricultural Bureaux, London, 162 pp. . 1976. A review of biological control in west- ern and southern Europe. Tech. Comm. 7. Com- monwealth Agricultural Bureaux, London, 182 pp. Liebregts, W. J. M. M., D. P. A. Sands, and A. S. Bourne. 1989. Population studies and biological control of Pseudaulacaspis pentagona (Targioni- Tozzetti) (Hemiptera: Diaspididae) on passion fruit in Western Samoa. Bull. Ent. Res. 79:163-171. Peters, A., I. Aloalil, R. G. Hollingsworth, and H. Hammans. 1985. Insecticidal control of white peach scale (Pseudaulacaspis pentagona) on pas- sion fruit (Passiflora edulis var. flavocarpa) in Western Samoa. Alafua Agric. Bull. 10: 13-17. Rasch, A. 1986. Review of the current situation of the passion fruit industry in Western Samoa. Working Paper Dev. Bank West. Samoa, Apia, Western Samoa, 24 pp. Silvestri, F. 1909. Notizie e descrizioni preliminari di insetti parasiti della Diaspis pentagona. Ren- diconti Reale Acad. dei Lincei 18: 563-565. . 1930. Contributo alla conscenza della specie orientali del genere Prospaltella (Hym.: Chalcidi- dae). Boll. Lab. Zool. Portici 25: 49-68. Waterhouse, D. F.and K. R. Norris. 1987. Biological control: Pacific prospects. Inkata Press, Mel- bourne, 454 pp. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, p. 138 NOTE Baetis jesmondensis McDunnough, a New Junior Synonym of Baetis tricaudatus Dodds (Ephemeroptera: Baetidae) In their revision of the Nearctic Baetis species, Morihara and McCafferty (1979. Trans. Am. Entomol. Soc. 105: 139-221) inadvertently omitted Baetis jesmondensis McDunnough from their account of species synonymies. They had found the species equivalent to Baetis tricaudatus based on an examination of the type material and larvae tentatively assigned to B. jesmon- densis. Our recent rediscovery and review of this material leads to the same conclu- sion, fully substantiating the previous con- clusion of Morihara and McCafferty. Members of the Holarctic Baetis rhodani species group, to which B. tricaudatus and B. jesmondensis belong, demonstrate con- siderable morphological variation with re- spect to developmental temperature gradi- ents and geographic distribution. The adult types of B. jesmondensis are clearly within the known range of variation of B. tricau- datus. Furthermore, our examination of nu- merous larval series from the northwestern United States and southwestern Canada, in- cluding those assumed to be B. jesmonden- sis and taken from its type locality, showed that all larvae fell within the concept of B. tricaudatus. All of these data lead us to re- affirm that the two names are synonymous. We therefore designate B. jesmondensis McDunnough as ajunior synonym of Baetis tricaudatus Dodds, New Synonym. This paper is Number 11856 of the Pur- due Agriculture Experiment Station. Robert D. Waltz, Division of Entomology and Plant Pathology-IDNR, 613 State Of- fice Building, Indianapolis, Indiana 46204; W. P. McCafferty, Department of Ento- mology, Purdue University, West Lafayette, Indiana 47907. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 139-145 FIRST DISTRIBUTIONAL RECORDS OF TABANIDAE (DIPTERA) IN CONNECTICUT Curis T. MAIER Department of Entomology, The Connecticut Agricultural Experiment Station, P.O. Box 1106, New Haven, Connecticut 06504. Abstract. —First distributional records from Connecticut are given for Aty/otus sphag- nicola Teskey, Chrysops celatus Pechuman, C. hinei Daecke, C. nigribimbo Whitney, Goniops chrysocoma (Osten Sacken), Hybomitra daeckei (Hine), H. frosti Pechuman, H. longiglossa (Philip), H. lurida (Fallen), H. minuscula (Hine), H. nitidifrons nuda (Mc- Dunnough), H. trepida McDunnough, H. zonalis (Kirby), Stonemyia isabellina (Wiede- mann), Zabanus fulvicallus Philip, and 7. vivax Osten Sacken. Habitats and flight periods of most species are discussed. Key Words: Fairchild (1950) summarized distribu- tional records of deer flies and horse flies in Connecticut. His synopsis, however, is out- dated as a result of nomenclatural changes, descriptions of new species, and new state collection records. A current assessment of tabanids in Connecticut would be valuable in light of their considerable importance as nuisance pests and carriers of infectious agents (Krinsky 1976). During the last de- cade, I have collected tabanids throughout Connecticut to provide new data for my planned revision of Fairchild’s (1950) monograph. My extensive sampling of bogs and fens and examination of museum spec- imens have resulted in many new tabanid records. Here I report the first distributional data for 16 tabanid species in Connecticut to make the information available for the database of North American Diptera which is now in preparation. MATERIALS AND METHODS Sampling sites in Connecticut are iden- tified by county, town (geographical sub- division of a county), and additional infor- mation if available. The number and sex of bog, fen, Atvlotus, Chrysops, Goniops, Hybomitra, Stonemyia, Tabanus tabanids collected on a given date or by a certain method are given in parentheses. Adults were captured with a hand-held in- sect net or with a Malaise or canopy trap. The use of the horizontal Malaise trap (D. Focks and Co., P.O. Box 12852, Gaines- ville, Florida 32608) is discussed by Maier (1984). The canopy trap, which 1s pictured by Pechuman (1981, Fig. 9), was baited with ca. 4 kg of dry ice to increase the number of specimens captured. Voucher specimens of tabanids except Stonemyia tsabellina (Wiedemann) are de- posited in the insect collection at The Con- necticut Agricultural Experiment Station, New Haven. The adult specimen of S. is- abellina is retained in the author’s collec- tion. COLLECTION RECORDS AND COMMENTS Atylotus sphagnicola Teskey Collection records.— Hartford Co., Bur- lington, bog near Covey Road, 27 May 1986 (1 4). Litchfield Co., Salisbury, shrubby bog by Bingham Pond, 14-16 June 1983, hor- izontal Malaise trap (1 4), 21 June 1985, 140 horizontal Malaise trap (1 2), 12 July 1984, horizontal Malaise trap (1 2). Based on collection records given by Tes- key (1983) and those presented here, this species apparently is restricted to sphagnum bogs. In Connecticut, adults fly in parts of bogs dominated by ericaceous shrubs. Aty- lotus sphagnicola was recorded previously from northwestern Connecticut (Teskey 1983), but the record is erroneous (Teskey, personal communication 1985). My collec- tion localities are the two southernmost to date. In Connecticut, adults have been col- lected between 27 May and 12 July. Chrysops celatus Pechuman Collection records.—Litchfield Co., Cornwall, Mohawk Pond, 19 July 1979 (1 2), near Hollenbeck River at junct. Conn. Highways 43 and 63, 4 August 1987 (1 2); Kent, Leonard Pond Swamp, 9 July 1979 (1 2), Mud Pond, 20 July 1979 (4 2); Nor- folk, marsh at edge of Beckley Pond, 7 July 1986, horizontal Malaise trap (2 2). Mid- dlesex Co., East Haddam, bog at north end of Lake Hayward, 3 July 1984, canopy trap (8 2), 17 July 1977 (1 2), 1-2 August 1984, canopy trap (2 2). New Haven Co., Bran- ford, 30 July 1985 (2 ), near Stony Creek, 28 July 1948 (1 2); Guilford, 4.5 km NNW junct. Conn. Highways 77 and 80, 27 July 1985 (1 2); Hamden, 12 July 1928 (1 2); New Haven, 9 June 1911 (1 2), 28 June 1911 (1 2); Madison, 14-15 June 1979, Malaise trap baited with dry ice (1 9); Meriden, South Meriden, 7 July 1935 (1 2). New London Co., Voluntown, fen in Atlantic white cedar swamp by Beachdale Pond, 19 June 1985, horizontal Malaise trap baited with dry ice (2 2), 10 July 1984, horizontal Malaise trap (5 2), 16 July 1986, canopy trap (2 2), 16- 17 July 1986, horizontal Malaise trap baited with dry ice (6 2), 7 August 1984, hovering over human (5 2), horizontal Malaise trap baited with dry ice (2 2), 17 August 1984, hovering over human (1 2), Pachaug St. For- est, near Hodge Pond, 21 June 1986 (4 °). Tolland Co., Hebron, Tom 2-Pony, Kinney Road, 7 July 1979 (2 2); Mansfield, Mt. Hope PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON River, Laurel Lane, 24 June 1979 (1 9); So- mers, 12 July 1979 (1 2); Tolland, near Tol- land Marsh Pond, 28 June 1988, canopy trap (5 2); Union, Bigelow Hollow St. Park, 16 July 1979 (2 2); Willington, 1.5 km NNE West Willington, black spruce bog, 30 June 1982 (1 9). Windham Co., Ashford, Elliot Road, 24 June 1979 (1 2); 28 June 1979 (5 2), Great Oak Road, 6 July 1979 (1 2); Plain- field, cedar swamp, 13 July 1979 (1 @); Put- nam, 8 July 1953 (4 2); Woodstock, Old Turnpike Road, 16 July 1979 (1 9). Pechuman (1949) originally described C. celatus as a subspecies of C. flavidus Wiede- mann. Fairchild (1950) recorded C. flavidus from Connecticut, but specimens examined by him are a mixture of C. celatus and C. flavidus. Chrysops celatus is a common species that occurs in a variety of habitats throughout Connecticut. Adults have been captured between 9 June and 17 August. Chrysops hinei Daecke Collection records. — Middlesex Co., Deep River, Cockaponset St. Forest, 12-13 Au- gust 1977, horizontal Malaise trap (3 2); East Haddam, bog at north end of Lake Hay- ward, 1-2 August 1984, canopy trap (1 9); Killingworth, 3 km NW junct. Conn. High- ways 81 and 148, 13 August 1985 (3 2). New Haven Co., North Haven, 27 August 1983 (1 2). New London Co., North Stonington, 3 km WNW Clarks Falls, 14 August 1985 (2 9). This late season deer fly is found mainly in boggy or swampy areas of Connecticut. Jones and Anthony (1964) also reported that it was restricted to swamps. My records in- dicate adults fly from 1 to 27 August in Connecticut. Lawrence et al. (1976) cap- tured adults in Maryland from | August to 12 September. Chrysops nigribimbo Whitney Collection records.—Litchfield Co., Cornwall, Mohawk Pond, 19 July 1979 (2 2). Middlesex Co., East Haddam, bog at north end of Lake Hayward, 17 July 1985 (2 2), 7 August 1986 (1 2). New London Co., VOLUME 92, NUMBER | Voluntown, 2 km NE town center, 7 August 1984 (1 2). Windham Co., Ashford, Elliot Road, 24 June 1979 (3 2), 28 June 1979 (3 9). This species has been captured between 24 June and 7 August in Connecticut, where it is distributed widely. Goniops chrysocoma (Osten Sacken) Collection records.—New Haven Co., Guilford, 4.5 km NNW junct. Conn. High- ways 77 and 80, 21 July 1983 (1 2), 4.0 km NW junct. Conn. Highways 77 and 80, 21 July 1987 (1 9). Both females were resting on vegetation at the edge of a large coastal red maple swamp. My distribution records are the first from New England. Hybomitra daeckei (Hine) Collection records.—New Haven Co., Guilford, Chaffinch Island, 14 June 1986 (3 2), 19 June 1985 (12 2), 19 June 1986 (12 2), 26 June 1986 (2 2), 3 July 1985 (9 2), 15 July 1989 (7 2), all from canopy traps. New London Co., Old Lyme, 1 km S junct. In- terstate Highway 95 and Conn. Highway 156, 3 July 1986, canopy trap (2 9). All Connecticut localities with H. daeckei are salt-marshes where larvae apparently develop. In Connecticut, adults have been trapped between 14 June and 15 July. This flight period is similar in length to the one of 30 May to 26 June recorded in Maryland (Lawrence et al. 1976). Hybomitra frosti Pechuman Collection records.—Litchfield Co., Cornwall, Mohawk St. Forest, black spruce bog, 12 August 1986, canopy trap (2 2); Nor- folk, bog by Beckley Pond, 22 July 1987 (1 9). This species is restricted to sphagnum bogs and possibly fens (Pechuman 1960, 1981, Baribeau and Maire 1983a, b). I collected my three specimens in bogs where the sphagnum mat was shaded partially by black spruce, Picea mariana (Miller) Britton, Sterns, and Poggenburg, and by ericaceous 141 shrubs. Shade may be necessary for larval survival because Baribeau and Maire (1983b) found most of their larvae in a for- ested portion of an ombrotrophic bog. Based on Connecticut records, adult activity lasts from 21 July to 11 August. This flight period is similar in time and length to that ob- served in Quebec by Baribeau and Maire (1983c). Hybomitra longiglossa (Philip) Collection records. —Litchfield Co., Nor- folk, bog by Beckley Pond, 23 May 1988 (1 6), 27 May 1988 (1 4, 1 2), 29 May 1987, canopy trap (13 @). Adults are active at peat pools in the bog mat where ericaceous shrubs are sparse and rarely exceed 0.5 m in height. Males ap- parently take stations at pool margins be- cause they chase flying insects that pass by them and they mate by pools. This species probably is confined to sphagnum bogs where larvae develop (Teskey and Burger 1976). The specimens captured in Con- necticut extend the known range of H. /on- giglossa southward by one state. In Ontario, adults are active from 3 to 29 June (Pechu- man et al. 1961). Hybomitra lurida (Fallen) Collection records.—Litchfield Co., Ca- naan, junct. Conn. Highway 126 and Page Road, 23 May 1988 (1 2); Norfolk, bog by Beckley Pond, 29 May 1987 (1 2); Salisbury, near Bingham Pond, 23 May 1987 (1 9). Adults of this uncommon species are known only from the three localities in northwestern Connecticut listed above. These collection sites are the southernmost ones recorded for this species. Its 2-week flight period in May is about one-half the length of that observed elsewhere in north- eastern North America (Lewis and Bennett 1977, Maire 1984a, White et al. 1985). Hybomitra minuscula (Hine) Collection records.— Hartford Co., Bur- lington, 15 July 1976 (5 2), 19 July 1976 (3 2), bog by Lamson Corner, 25 June 1986, 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON horizontal Malaise trap baited with dry ice (2 2), 9 August 1985 (11 2); East Windsor, bog at junct. Morris and Wapping Roads, 22 August 1985 (3 4, 3 2). Litchfield Co., Norfolk, bog by Beckley Pond, 30 June 1958 (1 4), 7 July 1986, canopy trap (3 2), hori- zontal Malaise trap baited with dry ice (2 2), 7-8 July 1986, horizontal Malaise trap (2 2), 17 July 1958 (2 2), 17 July 1970 (1 4, 3 2), 2 August 1960 (1 2), 12 August 1986 (2 2), bog by Pond Hill Pond, 12 August 1986 (1 2), bog by Tobey Pond, 12 August 1986 (2 2), 5 km S town center, 3 August 1984 (1 2): Salisbury, bog by Bingham Pond, 12 July 1984, canopy trap (2 2), 21-23 July 1983, horizontal Malaise trap (3 ¢, 2 2), 3 August 1984, canopy trap (1 2). Middlesex Co., East Haddam, bog at north end of Lake Hayward, 3 July 1984 (2 9), 24 July 1985 (2 4, 1 2), 1-2 August 1984, canopy trap (1 2), 13 August 1985 (1 4, 2 9), 21 August 1984, canopy trap (1 2); Killingworth, 1.3 km SE Kroopa Pond, 19 July 1973 (1 2). New Haven Co., Bethany, 4 August 1951 (2 2). New London Co., North Stonington, 3 km WNW Clarks Falls, 14 August 1985 (1 4); Voluntown, fen in Atlantic white cedar swamp by Beachdale Pond, 16-17 July 1986, horizontal Malaise trap (1 2), 17 August 1984 (1 2). Tolland Co., Willington, | km SSW junct. Interstate Highway 84 and Conn. Highway 32, black spruce bog, 30 June 1982 (5 2), 12 August 1982 (2 4, 2 2), 0.6 km N junct. Conn. Highways 74 and 320, 13 Au- gust 1985 (3 6, 3 2). Windham Co., Plain- field, 4 km E town center, 10 July 1984 (1 2), 20 July 1983 (1 4, 2 2), 4 August 1982 (1 2), 19 August 1983 (1 2); Windham, 1.8 km W town center, bog by Plains Road, 13 August 1985 (3 2). In Connecticut, this species occurs exclu- sively in sphagnum bogs which are the only known larval habitats (Teskey 1969). Bar- ibeau and Maire (1983b) found H. minus- cula to be the most common tabanid in open areas of bogs. My observations agree fully with their assessment. In open areas of Con- necticut bogs, males commonly hover about 1 m above the bog mat while they seek fe- males. My observations indicate adults are active between 25 June and 22 August in Connecticut. The flight period in south- western Ontario is nearly identical in time and duration (Judd 1958). Hybomitra nitidifrons nuda (McDunnough) Collection records.—Hartford Co., Bur- lington, bog by Lamson Corner, 27 May 1986 (3 2). Litchfield Co., Canaan, Robbins Swamp, 23 May 1985 (1 2); Cornwall, Mo- hawk St. Forest, black spruce bog, 11 June 1987 (1 2); Norfolk, bog by Beckley Pond, 23-27 May 1988, horizontal Malaise trap (5 2), 29 May 1987, canopy trap (4 2), 4 km WSW town center, 7 June 1984 (1 2), 5 km S town center, 7 June 1984, canopy trap (3 9); Salisbury, Benton Hill Fen, 11 June 1987, canopy trap (2 2), bog by Bingham Pond, 23 May 1987 (2 2), 4 June 1986 (1 9), 16 June 1983 (2 2), 20 June 1984 (1 2). New London Co., Voluntown, | km N junct. Conn. Highways 49 and 165, 15 May 1985 (1 2), 29 May 1986 (1 2). Tolland Co., Wil- lington, | km SSW junct. Interstate High- way 84 and Conn. Highway 32, black spruce bog, 20 May 1986 (1 2), 3 June 1983 (1 2). In Connecticut, adults inhabit a variety of bogs and fens bordered to some extent by woodland swamps which are the typical larval habitats (Teskey 1969). Adults have been captured between 15 May and 20 June in Connecticut. Smith et al. (1970), Golini and Wright (1978), and Leprince et al. (1983) collected adults over a 4- to 6-week period in Ontario or Quebec. Hybomitra trepida McDunnough Collection records. —New London Co., Colchester, 1.5 km N Conn. Highway 16 by Salmon River, 3 July 1985, canopy trap (1 2); Voluntown, | km N junct. Conn. High- ways 49 and 165, 19 June 1985 (1 2), 16 July 1986 (1 2), 16-17 July 1986, horizontal Malaise trap (2 2). Most specimens are from a fen surround- ed by aswamp dominated by Atlantic white VOLUME 92, NUMBER 1 cedar, Chamaecyparis thyoides (Linnaeus) Britton, Sterns, and Poggenburg. Like the larval habitats described by Teskey (1969) and Baribeau and Maire (1983b), the fen has abundant sphagnum. In Connecticut, adults have been taken between 19 June and 17 July. Flight periods recorded elsewhere in northeastern North America vary from 2 weeks to 2 months (Pechuman and Burton 1969, Smith et al. 1970, Lewis and Bennett 1977). Hybomitra zonalis (Kirby) Collection records. — Fairfield Co., New- town, Hopewell Road by Aspetuck River, 19 June 1988, canopy trap (1 2). Litchfield Co., Salisbury, bog by Bingham Pond, 20 June 1984, canopy trap (2 2), fen by Beeslick Pond, | 1-23 June 1987, horizontal Malaise trap (1 2), | km SSE Bald Peak at Wacho- castinook River, 4 June 1986, canopy trap (1 2). New Haven Co., Guilford, Beaver Head Swamp, 7 June 1984 (2 2), 9 June 1984(1 2), 1O-11 June 1984, horizontal Ma- laise trap (1 2), 14 June 1987, on dog (1 9). New London Co., Voluntown, fen in At- lantic white cedar swamp by Beachdale Pond, 29 May 1985 (1 2), 29 May 1986, horizontal Malaise trap baited with dry ice (5 2), 10 June 1984 (1 2), 19 June 1985, horizontal Malaise trap baited with dry ice (24 2), 10 July 1984 (1 9). Adults frequent bogs, fens, and coastal red maple swamps. Teskey (1969) and Bar- ibeau and Maire (1983b) found larvae in Canadian bogs, fens, or both. Larvae prob- ably develop in a wider range of habitats because collection localities in red maple swamps in Connecticut were > 10 km from the nearest bog or fen. Nonetheless, based on trapping records, the largest populations of H. zonalis apparently occur in fens as- sociated with coastal Atlantic white cedar swamps. Adults are on the wing at least from 29 May to 10 July, which is a comparable period in length to the one noted in Quebec by Maire (1984b). Smith et al. (1970), Lewis and Bennett (1977), and Baribeau and Maire 143 (1983c) captured adults over a 3- to 4-week period in other areas in northeastern North America. Stonemyia isabellina (Wiedemann) Collection records.—New Haven Co., Hamden, Lockwood Farm, 17-18 July 1982, horizontal Malaise trap (1 9°). This lone female was captured in a re- growth forest that is described by Maier (1984). My specimen is the first recorded from New England. Pechuman (1981) has suggested that this species may be uncom- mon throughout its range. The collection date of my specimen falls within the flight period of 24 June to 31 July reported for six Pennsylvania specimens (Frost and Pe- chuman 1958). Tabanus fulvicallus Philip Collection records. — Litchfield Co., Salis- bury, fen by Beeslick Pond, 18-22 July 1987, horizontal Malaise trap (1 2). Middlesex Co., East Haddam, bog at north end of Lake Hayward, 24 July 1985 (1 9). Both collection sites have a limited num- ber of small pools on the sphagnum mat, which resemble the areas where Teskey (1969) found larvae. The locality in Salis- bury is a rich fen, and the one in East Had- dam a minerotrophic bog (Maier 1988) or medium fen. Based on adult captures be- tween 29 June and 20 July in Ontario (Pe- chuman et al. 1961), the flight period in Connecticut is probably brief, not exceeding 3 weeks in July. Tabanus vivax Osten Sacken Collection records.—Litchfield Co., Ca- naan, 1.2 km SSW South Canaan, 18 July 1987 (1 2); Cornwall, near Kellog Corners, 22 June 1988 (1 2); Goshen, Allyn Road, 19 July 1979 (1 2); Harwinton, Campville, Naugatuck River, 11 July 1985, canopy trap (1 2), 18 July 1987, canopy trap (2 2); Nor- folk, 5 km S town center, 3 August 1985, canopy trap (2 2). Middlesex Co., East Had- 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dam, bog at north end of Lake Hayward, 3 July 1984, canopy trap (1 2°). All localities with 7. vivax are near streams or rivers. Teskey (1969) also emphasized the presence of streams near his larval col- lection sites. In Connecticut, adults fly at least from 22 June to 3 August. Pechuman et al. (1961) observed a shorter flight period of 12 June to 17 July in Ontario. Although Pechuman (1981) did not give exact dates of capture in New York, he noted that adults had been captured during June, July, and August. ACKNOWLEDGMENTS I thank the South Central Connecticut Regional Water Authority and The Nature Conservancy (Connecticut Chapter) for al- lowing me to collect tabanids on their prop- erty. The following people provided valu- able assistance when I examined tabanids in collections under their care: David G. Furth, Peabody Museum, Yale University, New Haven, CT; Jane E. O’Donnell, Uni- versity of Connecticut, Storrs, CT; Scott R. Shaw, Museum of Comparative Zoology, Harvard University, Cambridge, MA; Louis N. Sorkin, American Museum of Natural History, New York, NY; Kenneth A. Welch, The Connecticut Agricultural Experiment Station, New Haven, CT. Also, I appreciate the help of John F. Burger, University of New Hampshire, Durham, NH, L. L. Pe- chuman (retired), Cornell University, Ith- aca, NY, and H. J. Teskey (retired), Bio- systematics Research Institute, Ottawa, ON, who aided me in identifying tabanids. Louis Magnarelli reviewed an earlier draft of the manuscript. LITERATURE CITED Baribeau, L. and A. Maire. 1983a. Latitudinal dis- tribution of Quebec Tabanidae. Mosq. News 43: 7-13. 1983b. Spatial distribution of Tabanidae (adults and larvae) in two bogs of southern Que- bec. Mosq. News 43: 24-29. . 1983c. Abundance and seasonal distribution of Tabanidae in a temperate and in a subarctic locality in Quebec. Mosq. News 43: 135-143. Fairchild, G. B. 1950. Family Tabanidae. Guide to the insects of Connecticut. Part VI. The Diptera or true flies of Connecticut. St. Geol. Nat. Hist. Surv. Bull. 75: 3-31. Frost, S. W. and L. L. Pechuman. 1958. The Taban- idae of Pennsylvania. Trans. Amer. Entomol. Soc. 84: 169-215. Golini, V. I. and R. E. Wright. 1978. Relative abun- dance and seasonal distribution of Tabanidae (Diptera) near Guelph, Ontario. Can. Entomol. 110: 385-398. Jones, C. M. and D. W. Anthony. 1964. The Taban- idae (Diptera) of Florida. USDA Tech. Bull. 1295: 1-85. Judd, W.W. 1958. Studies of the Byron Bog in south- western Ontario. V. Seasonal distribution of horseflies and deerflies (Tabanidae). Can. Ento- mol. 90: 255-256. Krinsky, W. L. 1976. Animal disease agents trans- mitted by horse flies and deer flies (Diptera: Ta- banidae). J. Med. Entomol. 13: 225-275. Lawrence, R. S., W. E. Bickley, and J. Mallack. 1976. The seasonal distribution of biting flies in St. Mary’s County, Maryland in 1975 (Diptera: Tabanidae, Muscidae). Maryland Agric. Exp. Stn. Misc. Publ. 904: 1-18. Leprince, D. J., D. J. Lewis, and J. Parent. 1983. Biology of male tabanids (Diptera) aggregated on a mountain summit in southwestern Quebec. J. Med. Entomol. 20: 608-613. Lewis, D. J. and G. F. Bennett. 1977. Biting flies of the eastern Maritime Provinces of Canada. I. Ta- banidae. Can. J. Zool. 55: 1493-1503. Maier, C. T. 1984. Habitats, distributional records, seasonal activity, abundance, and sex ratios of Boreidae and Meropeidae (Mecoptera) collected in New England. Proc. Entomol. Soc. Wash. 86: 608-6 13. . 1988. Life cycle of Coptodisca negligens (Lep- idoptera: Heliozelidae) on cranberry. J. Econ. Entomol. 81: 497-500. Maire, A. 1984a. Les Tabanidae (Diptera) a limite septentrionale des foréts (domaine maritime du Nouveau-Québec). Can. Entomol. 116: 227-233. . 1984b. Les taons (Diptera: Tabanidae) d’une toubiére a palses située 4 la limite des foréts (Nou- veau-Québec). Can. Entomol. 116: 1679-1682. Pechuman, L. L. 1949. Some notes on Tabanidae (Diptera) and the description of two new Chrysops. Can. Entomol. 81: 77-84. 1960. Some new and little-known North American Tabanidae (Diptera). Can. Entomol. 92: 793-799. 1981. The horse flies and deer flies of New VOLUME 92, NUMBER 1 York (Diptera, Tabanidae), 2nd edition. Search: Agriculture (Cornell Univ. Agric. Exp. Stn.) 18: 1-68. Pechuman, L. L. and J. J. S. Burton. 1969. Seasonal distribution of Tabanidae (Diptera) at Texas Hol- low, New York in 1968. Mosq. News 29: 216- 220. Pechuman, L. L., H. J. Teskey, and D. M. Davies. 1961. The Tabanidae (Diptera) of Ontario. Proc. Entomol. Soc. Ontario 91: 77-121. Smith, S. M., D. M. Davies, and V. I. Golini. 1970. A contribution to the bionomics of the Tabanidae (Diptera) of Algonquin Park, Ontario: Seasonal distribution, habitat preferences, and biting rec- ords. Can. Entomol. 102: 1461-1473. 145 Teskey, H.J. 1969. Larvae and pupae of some eastern North American Tabanidae (Diptera). Mem. Entomol. Soc. Can. 63: 1-147. 1983. A revision of eastern North American species of Aty/otus (Diptera: Tabanidae) with keys to adult and immature stages. Proc. Entomol. Soc. Ontario 114: 21-43. Teskey, H. J. and J. F. Burger. 1976. Further larvae and pupae of eastern North American Tabanidae (Diptera). Can. Entomol. 108: 1085-1096. White, D. J.,C. D. Morris, and K. Green. 1985. Sea- sonal distribution of northern New York State an- thropophilic Tabanidae (Diptera) and observa- tions on the dispersal of several species. Environ. Entomol. 14: 187-192. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 146-152 A NEW SPECIES OF CECIDOMYIIDAE (DIPTERA) DAMAGING SHOOT TIPS OF YELLOW CYPRESS, CHAMAECYPARIS NOOTKATENSIS, AND A NEW GENUS FOR TWO GALL MIDGES ON CUPRESSACEAE RAYMOND J. GAGNE AND R. W. DUNCAN (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560; (RWD) Pacific Forestry Centre, Forestry Canada, 506 West Burnside Road, Victoria, B.C. V8Z 1M5, Canada. Abstract.—A new species of gall midge (Diptera: Cecidomyiidae) infesting the shoot tips of yellow cypress, Chamaecyparis nootkatensis, in British Columbia is described and illustrated. A new genus, Chamaediplosis, is erected for the new species and one other, previously described species, Contarinia rugosa, known from Cupressus arizonica. The new genus belongs to the tribe Cecidomyiini. Key Words: A new species is described here that was first discovered in 1987 damaging the shoot tips of yellow cypress, Chamaecyparis noot- katensis (D. Don) Spach (Cupressaceae) in British Columbia. The large numbers of in- fested and ultimately killed shoot tips at one site indicate that this species could become a serious pest (Fig. 1). The new gall midge has one generation per year at Saanichton, British Columbia. Most overwintering lar- vae (Fig. 2) pupate within the galls in late winter. Adults emerge soon after (Fig. 3); others pupate in the spring or early summer. Each gall usually contains one larva, but occasionally two, rarely three, are found. A new genus is erected for the new species and Contarinia rugosa Gagné, another species that lives in shoot tips of Cupres- saceae (Gagné 1986b). A separate paper by R. W. Duncan on the biology of the new species in British Columbia is in prepara- tion. gall midges, western North America, taxonomy METHODS Branches with infested shoot tips were collected in the field in February, 1987 near Saanichton, B.C. To maintain the galls in fresh condition during rearing they were placed in sealed polyethylene bags and kept at a constant 20°C until adults emerged. Adults began emerging from the galls after five days and continued for several days afterwards. Immature and adult specimens were preserved in 70% ethanol and mount- ed for microscopic study in Canada balsam using the method outlined in Gagné (1989). Adult terminology follows usage in Mc- Alpine (1981) and larval terminology that in Gagné (1989). The new genus is to be attributed to Gagné, the new species to Gagné and Duncan. Chamaediplosis Gagne, NEw GENUS Adult.— Head: Eyes 5-7 facets long at vertex, separated by '2 to 1 facet diameter; VOLUME 92, NUMBER 1 facets circular, closely adjacent except near midheight of eye where they may be as far as ' facet diameter apart. Vertex of occiput rounded, without dorsal protuberence, with 2-3 rows of setae parallel to the periphery. Frons with several setae. Labella hemi- spherical, with scattered setae. Palpus 4-segmented. Male antennal flagellomeres (Gagné 1986a: Fig. 1) binodal, bicircumfi- lar, the circumfilar loops regular. Female flagellomeres (Gagné 1986a: Figs. 2-3) pro- gressively shorter towards antennal apex, the circumfila appressed. Thorax: Scutum with 2 lateral and 2 dor- socentral rows of setae and setiform scales. Scutellum with a group of setae on each side. Mesanepisternum with 0-3 scales. Mesep- imeron with 6-10 setae. Claws. slightly shorter than empodia, the empodia broad, about as wide as Sth tarsomere. Wing with RS curved apically to join C posterior to wing apex, C broken at juncture with RS. Male abdomen (Figs. 4-6): Tergites 1 to 6 entire, rectangular, with mostly single, un- interrupted, posterior row of setae, 4-10 lat- eral setae anterior to posterior row, a few scattered scales, and pair of trichoid sensilla on anterior margin: tergite 7 as for preceding except weakly sclerotized posteromedially, posterior setae usually present only later- ally; tergite 8 sclerotized only anteriorly to anterolaterally, usually bare except for an- terior pair of trichoid sensilla; pleura with sparse scales; sternites 2—6 rectangular, with mostly single, caudal row of setae, with mixed setae and scales grouped near mid- length, and anterior pair of trichoid sensilla; cerci broadly rounded posteriorly, with ven- trolateral setae; hypoproct deeply divided, its lobes broad, rounded apically, with api- cal and ventral setae; aedeagus attenuate, narrowly rounded apically, with lateral sen- silla; gonocoxites stout, apodeme variously shaped; gonostylus long, narrowing slightly from base to apex, mostly striate, setulae present only near base, chiefly on venter with scattered setae and completely setu- lose. 147 Female abdomen (Figs. 7-9): Tergites 1 to 7 and sternites 2—7 generally as for male, but tergal setae and scales more numerous. Tergite 7 with mostly double row of pos- terior setae, about 7 length of distal half of ovipositor; tergite 8 approximately as long as 7, with anterior pair of trichoid sensilla and 0-10 short posterior setae. Ovipositor short, protrusible, proximal half anteriorly with scattered lateral and ventral setae, dis- tal half posteriorly with scattered short se- tae, completely setulose, unstriated: cerci broad at base, tapering gradually to rounded apex, completely setulose, setae concen- trated at base and apex; hypoproct divided into 2 lobes. Third instar (Figs. 10-12).—Integument rugose. Spatula present, variously shaped, with 2 anterior lobes. Papillae with basic complement of papillae for supertribe (Gagné 1989), but with very short setae; ter- minal papillae with short setae, the usually large, corniform pair found in Cecidomyiini reduced in size, barely larger than remaining three pairs. Type species.—Chamaediplosis nootka- tensis Gagné and Duncan. Etymology.—The name Chamaediplosis combines ““Chamae” (dwarf, creeping) from Chamaecyparis with “‘diplosis” (double, a doubling), a commonly used sufhx for gall midge genera of the supertribe Cecidomyi- idi. Remarks. — Chamaediplosis contains C. nootkatensis, C. rugosa, and a third, un- described species, known only from a series of specimens in the National Museum of Natural History in Washington, D.C. That series, from Cupressus macrocarpa Hartw. in California, is in poor condition and un- suitable for description. These species all infest shoot tips of Cupressaceae. Erecting Chamaediplosis is a step in re- solving the problem of polyphyly in Con- tarinia by dividing that genus into smaller units whose species occur on related plants and share what one can convincingly argue are shared, derived characters. Chamaedi- 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 1 149 5 Figs. 4-6. Genitalia (left side, dorsal). plosis belongs to the tribe Cecidomyiini and differs from other genera in that tribe by its short ovipositor with relatively large and completely setulose cerci and the nearly uni- form terminal papillae of the larva. That the ovipositor is short is plesiotypic, but its — Figs. 1-3. 6 Male, C. nootkatensis. 4, Abdominal segments 6 to end (lateral view). 5, Genitalia (mesal). 6, distinctive shape and setation (Figs. 7-9), presumably well-adapted to its use, can be regarded as apotypic. One pair of larval ter- minal papillae that in Cecidomyiini are much larger than the three remaining pairs and has recurved corniform setae, are in 1, Normal and infested shoot tips of yellow cypress. 2, Infested shoot tip cut open to reveal larva of C. nootkatensis. 3, Newly emerged female of C. nootkatensis and its pupal exuviae protruding from infested shoot tip. 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10 11 1B Figs. 7-13. C. nootkatensis. 7-9, Female: 7, Abdominal segments 7 to end; 8, cerci and hypoproct (ventral view); 9, same (lateral). 10-12, Larva: 10, spatula with associated papillae; 11, spatula; 12, segments 8-9 (dorsal). 13, Spatula of C. rugosa. VOLUME 92, NUMBER 1 Chamaediplosis only slightly larger than that of the remaining papillae. Its small size may be a reduction or, alternatively, the primi- tive condition for the tribe. Except for the shape of the ovipositor and the larval terminal papillae, Chamaediplo- sis could fit into Contarinia, a genus that is used as a catch-all category and has grown to include most species of Cecidomyiuini with elongate, strongly tapered ovipositors (Gagné 1973). It now appears that long ovi- positors suitable for laying eggs in narrow crevices of buds and flowers evolved sepa- rately many times (Gagné 1989). One con- spicuous similarity between the new genus and Contarinia is the loss of one of the three circumfila on the male antennal flagello- meres. That is a character state that appears many times within the Cecidomyiini and is not necessarily evidence for particularly close kinship. The number of circumfila has become reduced separately elsewhere 1n the Cecidomyiini (Taxodiomyia), as well as in the Clinodiplosini (4 metrodiplosis), Lesto- diplosini (Endaphis, Dentifibula), and My- codiplosini (Mycodiplosis: Gagné and Rios de Saluso 1987). Other apotypic character states besides the bicircumfilar flagello- meres that Chamaediplosis and some other Cecidomyiini share but which may be the result of convergence are 1) the loss of the dorsal occipital projection of the head, 2) a certain amount of reduced setation and sclerotization in male abdominal tergites 7 and 8, 3) the deeply divided hypoproct with 2 short, cylindrical lobes, 4) the short ae- deagus, 5) the presence of setulae only at the base of the gonostylus, and 6) the large em- podia. The last is a character that this genus shares with most other conifer gall midges, regardless of their affinities. KEY TO SPECIES OF CHAMAEDIPLOSIS Larvae and pupae in shoot tips of Chamaecyparis nootkatensis; larval spatula with triangular an- terior lobes, occasionally the lobes secondarily toothed, the shaft slightly narrowed near mid- length (Figs. 10-11); female tergite 8 with 0-5 setae along posterior margin (Fig. 7); aedeagus acute at apex (Fig. 6) ...C. nootkatensis Gagné and Duncan Larvae and pupae in shoot tips of Cupressus ari- zonica; larval spatula with rounded anterior lobes, the shaft much narrowed posteriorly (Fig. 13); female tergite 8 with 10-12 setae along pos- terior margin; aedeagus broadly rounded at apex _.C. rugosa Gagné Chamaediplosis nootkatensis Gagne and Duncan, New SPECIES Adult.— Wing length, 2.0-2.5 mm. Tho- rax: anepisternum usually with 0, occasion- ally 1 scale; anepimeron with 6-9 setae. Male postabdomen as in Fig. 4, genitalia as in Figs. 5-6, the apodeme variable, bifurcate (as shown, Fig. 7) or entire. Female postab- domen as in Fig. 7, cerci and hypoproct as in Figs. 8-9. Third instar.— Orange. Spatula (Figs. 10- 12) short and broad, anteriorly with two triangular lobes, these sometimes second- arily divided. Papillae all on mamelons. Posterior segments as in Fig. 12. Holotype.— Male, ex Chamaecyparis nootkatensis, Saanichton, British Colum- bia, III-3-1987, R. Duncan, deposited in the Canadian National Collection in Ottawa. Paratypes, all ex Chamaecyparis nootkaten- sis from Saanichton, B. C.: 2 males, 2 fe- males, II-23-1987; 2 males, | female, III- 3-1987:; 4 males, 3 females, III-10-1987; 5 males, 2 females, [V-22-1988; and 5 larvae, IV-11-1987. The paratypes are divided among the U.S. National Museum of Nat- ural History, Washington, D.C., the Pacific Forestry Centre, and the Canadian National Collection. Remarks. — Except in the shape of the lar- val spatula, this species 1s very similar to C. rugosa. Adults of the two species can be separated with the help of the minor differ- ences outlined in the key given earlier. ACKNOWLEDGMENTS We are grateful to P. Malikul for making the slide preparations, L. Manning for tech- 152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nical help, D. L. Roney for inking Figs. 4— 13, and K. M. Harris, N. E. Woodley, and J. Yukawa for their review of the manu- script. LITERATURE CITED Gagné, R. J. 1973. A generic synopsis of the Nearctic Cecidomyiidi (Diptera: Cecidomyiidae). Ann. Entomol. Soc. Am. 66: 857-889. 1986. A new Nearctic species of Contarinia (Diptera: Cecidomyiidae) recently introduced into Hawaii on Cupressus (Cupressaceae). Proc. Ento- mol. Soc. Wash. 88: 127-130. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ith- aca, New York. xii of 355 pp. and 4 pls. and M. L. A. Rios de Saluso. 1988. A new rust-feeding Mycodiplosis (Diptera: Cecidomyi- idae) associated with rust on Avena sativa (Po- aceae) in Argentina. Ser. Tecn. Inst. Nac. Tecnol. Agropecu. Parana Entre Rios Argent. 54(1987): 1- 112% McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds. 1981. Manual of Nearctic Diptera. Vol. 1. Re- search Branch, Agriculture Canada. Monograph No. 27. vi + 674 pp. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 153-159 REVIEW OF HALTICHELLA SPINOLA IN THE NEARCTIC REGION (HYMENOPTERA: CHALCIDIDAE) JEFFREY A. HALSTEAD California State University, Fresno. Present address: 110 W. Barstow #112, Fresno, California 93704. Abstract. —Five species of Haltichella Spinola are recognized in the Nearctic region: H. onatas (Walker), H. ornaticornis Cameron, H. perpulcra (Walsh), H. rhyacionia Gahan, and H. xanticles (Walker). The females and males of each species are diagnosed. A neotype is designated for H. perpulcra. The males of H. onatas, H. ornaticornis, H. perpulcra, and H. xanticles were previously undescribed; voucher specimens of each are designated. Biological and distributional information is summarized for each species. A key to the Nearctic species and characters to distinguish Haltichella from other Nearctic Chalcididae are presented. Key Words: Haltichella Spinola is one of fifteen genera of Chalcididae in America north of Mexico and is known from all zoogeographical re- gions. This genus was described by Spinola (1811) with Chalcis pusilla Fabricius as the type-species. Taxonomic treatments of Hal- tichella include the European fauna (Boucek 1951), the Russian fauna (Nikolskaya 1952, 1960), the African fauna (Schmitz 1946), and the Japanese fauna (Habu 1960, 1962). As no comprehensive taxonomic work for the Nearctic species exists, I present a re- view of Nearctic Haltichella. Past literature on Haltichella is cataloged in Peck (1963), Burks (1979), and DeSantis (1979). Haltichella are among the smallest Chal- cididae, length rarely exceeding 4 mm. They are black and commonly with orange or or- ange-brown on the legs, antennae, and rare- ly, the tegulae. The wings vary, being hya- line, smokey, or clouded in a specific pattern. Males differ from females by having a short- er abdomen with a blunt apex (Fig. 2), ro- bustly filliform antennae (Fig. 3), and in Haltichella, review, Chalcididae, Nearctic some species, in color and forewing cloud- ing. A generic revision of the American Chal- cididae is in progress (Boucek pers. comm.); therefore, a generic description of Haltich- ella is omitted. However, to facilitate iden- tification of this genus for the Nearctic re- gion, the following characters are diagnostic (Fig. 1): hindtibia truncate distally, two spurs present (Haltichellinae); marginal vein reaching anterior margin of wing, post- marginal and stigmal veins present (Hal- tichellini); tergite 1 dorsally with longi- tudinal carinae at base, originating from a transverse carina; scutellum rounded, slightly bilobed, or rarely, strongly bi- lobed posteriorly but never with large, long projecting processes or a median tooth; vertex of head not produced into horns; frontal carina weak, not forming an arch above anterior ocellus; color predomi- nantly black. 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Useful species characters include the length of tergite 2 (medially); sculpture of tergite 1 (dorsally); color of the antennae, tegulae, and legs (especially the hindfemur); and presence or absence of clouding in the forewing. Besides the above characters, few diagnostic and constant characters exist to distinguish the Nearctic species. Rarely, in- dividual specimens are difficult to identify because of variation in the sculpture of ter- gite 1 and coloration. If possible, a series of specimens from a locale should be exam- ined. However, more than one species can be taken together such as Haltichella rhy- acionia Gahan and Haltichella xanticles (Walker). Haltichella are primary parasitoids of Lepidoptera, secondary parasitoids of brac- onid wasps, and once recorded as a second- ary parasitoid ofa tachinid fly (Freeman and Berisford 1979). Four of the five Nearctic species have been reared with three species being both primary and secondary parasit- oids and the fourth species as only a sec- ondary parasitoid. Primary hosts include several economically important lepidopter- ous pests (see host sections). During this revision, many new host records were ob- tained from label data. In the examination of several thousand specimens, the distributional information on Haltichella was greatly improved and shows the species to be widely distributed in the Nearctic region (Fig. 5). Haltichella are commonly collected in Malaise-type traps and pan traps or by sweeping vege- tation. Some species (e.g. H. xanticles) oc- cur in a variety of habitats, ranging from deserts to coniferous forests. Haltichella are common in collections, ranking third in number only to the Chalcididae genera Spi- lochalcis and Invreia. Abbreviations include T1 for tergite 1, etc. All measurements were made in the flattest plane possible. Specimens were examined at 30 to 100 x. A mylar, glare reducing screen was used in lighting specimens. Collections examined and museum ac- ronyms are as follows: American Museum of Natural History, New York; Bernice P. Bishop Museum, Hawaii; British Museum of Natural History, London (BMNH); Cal- ifornia Academy of Sciences, San Francisco; California Collection of Arthropods, Cali- fornia Department of Food and Agricul- ture, Sacramento; California State Univer- sity, Fresno; California State University, Sacramento; Canadian National Collection, Ottawa; Florida Collection of Arthropods, Florida Department of Agriculture and Consumer Affairs, Gainesville; Fresno County Department of Agriculture, Fresno, California; Illinois Natural History Survey, Champaign; Los Angeles County Museum of Natural History, California; Natural His- tory Museum of San Diego, California; Or- egon Department of Agriculture, Salem; Royal Ontario Museum, Toronto; Texas A&M University, College Station; Tulare County Agricultural Commissioner’s Of- fice, Visalia, California; United States Na- tional Museum of Natural History, Wash- ington, D.C. (USNM); University of California, Berkeley; University of Califor- nia, Davis; University of California, Riv- erside; J. A. Halstead personal collection; H. A. Hespenheide personal collection; R. D. Haines personal collection. Key TO NEARCTIC SPECIES OF HALTICHELLA SPINOLA Females, ovipositor present: <..,.c..:.ccsecunreeiete 2 Males, ovipositor absent Se) T2 medially less than /% the length ofTl PP er eae, perpulcra (Walsh) - 12 Stal greater than 4 the length of Tl .. 3 3. Tl punctate to coriaceous dorsally ......... ee oer cot rhyacionia Gahan ~ TI polished dorsally, without sculpture ...... 4 4. Forewing with one or two clouded spots, rarely hyaline; tegula orange, hindfemur black or with only apex orange ......... ornaticornis Cameron — Forewing hyaline, tegula black, hindfemur not as above .... 5. Legs and antennae black, forewing hyaline or smokey xanticles (Walker) — Legs (except apical ' of hindfemur occasion- nN l= VOLUME 92, NUMBER | ally) and antennae orange, forewing hyaline . Mae ei eames atcPer atebre casters etcciavaxeetiy ct a onatas (Walker) 6. Tl basally with 2 longitudinal carinae, flagel- lum 2'2x height of head . perpulcra (Walsh) — TI basally with 3 or more longitudinal carinae, flagellum less than 22= height of head |... 7 7. T1 punctate to coriaceous dorsally eC ee rhyacionia Gahan — TI polished dorsally, without sculpture .... 8 8. Tegula orange, rarely black; forewing with one or two clouded spots, rarely hyaline ints Nee eee ornaticornis Cameron — Tegula black, forewing hyaline or smokey but never with a clouded spot 9 9. Hindfemur with basal ' to '2 orange, fore and middle legs and scape orange onatas (Walker) — Hindfemur black, or with orange markings at base and/or apex; fore and middle legs and scape black or brown xanticles (Walker) Haltichella onatas (Walker) Fig. 5 Hockeria onatas Walker, 1843: 146, °. Haltichella onatas (Walker); Walker, 1846: 7. Conura onatas (Walker); Walker, 1871: 41. Haltichella longicornis Ashmead, 1887: 185; Burks, 1975: 164. Haltichella onatas (Walker); Burks, 1975: 164, Lectotype designation. Haltichella onatas (Walker), MALE DI- AGNOSIS. Diagnosis.—H. onatas is the only Nearc- tic species with the antennae (in male scape only) and legs (in male only basal 4 to 2 of hindfemora and apex of hindtibiae) orange. The characters: length of T2 medially great- er than 4 that of T1, T1 dorsally polished, with three or more (usually) longitudinal ca- rinae at base, hyaline forewing, black teg- ulae, and orange color are distinguishing. Female.—Black, with antennae and legs orange to orange-brown. Male.—Like female except antennae ro- bustly filiform, base of hindfemora and apex of hindtibiae orange, and abdomen shorter with apex blunt. Variation.—Hindfemur coloration is somewhat variable as noted in the key. (2) Length 2-4 mm. The hindfemur of one fe- 155 male (Type No. 2627 U.S.N.M., originally included in the syntype series of H. xanti- cles) is black, except basally. (¢) Length 2- 3 mm. Type and voucher specimens.—Lecto- type 2 with data: “B.M. Type Hym. 5. 553, 1477a, St. Jon’s Bluff, Hockeria Onatas Walker.” I designate a male specimen as a voucher specimen—red label marked: “VOUCHER MALE, Haltichella onatas (Walker), 4, det. J. A. Halstead 1987” and with data: “Crescent City, Fla. Apr’ 08, Van Duzee, MCVanDuzee Collector, MCVan- Duzee Collection, Collection of the CALI- FORNIA ACADEMY OF SCIENCES, San Francisco, Calif.” Both specimens in the United States National Museum. Hosts.— LEPIDOPTERA, Cosmopteryg- idae: Pyroderces rileyi (Wals.). Olethreuti- dae: Lasperyresia pomonella (L.) pupa, Gra- Pholitha molesta (Busck). Gelechiidae: Isophrictis similiella (Chamb.). Psychidae: Prochalia pygmaea Barnes & Mc- Dunnough. A specimen (New Orleans, Lou- isiana) was reared from a leaf skeletonizer on Phalaris canariensis (Canary Grass). Ashmead (1887) reared a specimen from the gall of Xanthoteras politum (Bassett) (Hymenoptera: Cynipidae). It is likely that a lepidopteran was inhabiting the gall. A specimen (Monticello, Florida) was reared from a stalked braconid wasp cocoon on Catocala (Lepidoptera: Noctuidae); thus, it is also a secondary parasitoid of Braconidae. Haltichella ornaticornis Cameron Fig. 5 Haltichella ornaticornis Cameron, 100, °. Haltichella ornaticollis Cameron; Howard, 1885: 36. Haltichella ornaticornis (Cameron); De- Santis, 1979: 69. Haltichella ornaticornis Cameron, MALE DIAGNOSIS. Diagnosis.—H. ornaticornis is the only Nearctic species with one or two clouded 1884: 156 spots in the forewing. The characters: length of T2 medially greater than 4 that of T1, T1 dorsally polished, with three or usually more longitudinal carinae at base, orange tegulae, black hindfemora, orange and black antennae (in female), and black antennae (in male) are distinguishing. Female.— Black, with the following areas orange: basal ' of scape, flagella 1-3, tarsi, base and apex of tibiae, trochanters, and tegulae. Forewing with a clouded spot from under marginal vein and stigma to middle of wing. Male.—Like female except antennae black and robustly filiform, and abdomen shorter with apex blunt. Forewing like female. Variation.—(?) Length 2-4 mm. Fore- wing clouding somewhat variable. Most specimens are like the holotype, but some with forewing hyaline or with two clouded areas (one under the marginal vein and another fainter spot between it and apex of wing). (4) Length 2-3 mm. Commonly, the forewing is hyaline. The tegulae are rarely black. Comments. — Females that lack forewing clouding can be distinguished by coloration and T1 and T2 characters. Type and voucher specimens. — Holotype 2 with data: ““‘Bugaba, Panama, Champion. BM Type Hym. 5. 288, Haltichella orna- ticornis Cameron.” I designate a male spec- imen as a voucher specimen—red label marked: “VOUCHER MALE, Haltichella ornaticornis Cameron, 4, det. J. A. Halstead 1987” and with data: “MEXICO, Chiapas, Palenque, 10 Sept 1974, GBohart, WHan- son, UTAH STATE UNIVERSITY.” Both specimens in the British Museum of Natural History. Host.— Unknown. Haltichella perpulcra (Walsh) Figs. 4, 5 Hockeria perpulcra Walsh, 1861: 258, @. (Paper not seen by author.) Hockeria perpulchra Walsh, Cresson, 1862: 228 (erroneous subsequent spelling). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Conura perpulchra (Walsh); Walker, 1871: 41. Hockeria perpulcra Walsh; Thomas, 1881: 39, Original description repeated. Haltichella perpulchra (Walsh), Howard, 1885: 37. Haltichella perpulcra (Walsh), NEOTYPE DESIGNATION 2°, MALE DIAGNO- SIS: Diagnosis.—H. perpulcra is the only Nearctic species with the length of T2 me- dially less than 4 that of Tl (Fig. 4), and (in male) flagellum 2! the height of head. The characters: hyaline forewing, T1 dor- sally polished, with two longitudinal carinae at base (in both sexes) and between these carinae 0-5 minute carinae (in female), and black color are distinguishing. Female.—Black, with base and apex of fore and middle tibiae, apex of hindtibiae, and tarsi orange. Male.—Like female except antennae longer and robustly filiform, and abdomen shorter with apex blunt. Variation. —(@ and 4) Length 2-4 mm. Type and voucher specimens. —The type specimen(s) could not be found and are be- lieved to have been destroyed in the Chi- cago fire of 1873. I was unable to obtain Walsh’s (1861) paper of the original de- scription. Thomas (1881), republishing Walsh’s (1861) description, presented key characters: ‘‘abdomen ovate, glabrous, first joint equal to three-fifths of its entire length, and highly polished, intermediate joints very narrow,” “wings hyaline,”’ and “general col- or black” (in combination with other char- acters in the description), which are ade- quate to distinguish this species. Walsh’s description appears to have been based on a female. I designate a female specimen as NEOTYPE-— yellow label marked: ““NEO- TYPE, Haltichella perpulcra (Walsh), ¢, det. J. A. Halstead 1987” and with the data: “MEXICO, Nyarit, San Blas, II-14-1974, G. E. Bohart, UTAH STATE UNIVERSI- TY.” I designate a male specimen as a voucher specimen—red label marked: VOLUME 92, NUMBER | “VOUCHER MALE, Haltichella perpulcra (Walsh), 4, det. J. A. Halstead 1987” and with data: “MEXICO, Chiapas, Palenque, 10 Sept 1974, GBohart, WHanson, UTAH STATE UNIVERSITY.” Neotype and voucher specimen of male deposited in the USNM. Host.—Cocoons of Apanteles militaris (Walsh) (Hymenoptera: Braconidae) on Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae). Haltichella rhyacionia Gahan Figs. 1-3, 5 Haltichella rhyacioniae Gahan, 1927: 545, 2 and 6. Haltichella rhyacionia Gahan; Burks, 1979: 861. Haltichella rhyacionia Gahan, MALE DI- AGNOSIS. Diagnosis.—H. rhyacionia is the only Nearctic species with T1 dorsally coria- ceous to punctate. The characters: length of T2 medially greater than '4 that of T1 (Figs. 1-2), Tl with three or usually more longi- tudinal carinae at base, hyaline forewing, and black legs and tegulae are distinguish- ing. Female.—Black, with apices of tibiae or- ange. Male.—Like female except antennae ro- bustly filiform and abdomen shorter with apex blunt. Variation. —(2 and 6) Length 2-4 mm. T1 coriaceous to punctate. The sculpture on T1 dorsally is rarely faint and confined to the basomedial area. The flagellum is rarely or- ange. Types.—Holotype ° and allotype ¢ in USNM with data: “ex Rhyacionia frustana Comst, Falls Ch Va, 7/16/24, R. A. Cush- man coll., Type No. 40178 U.S.N.M., Hal- tichella rhyacioniae Gahan Type.” Hosts.—LEPIDOPTERA, Olethreuti- dae: Rhyacionia bushnelli (Busck), R. frus- trana, R. rigidana (Fern). Lyonetiidae: Buc- culatrix thurberiella (Busck). DIPTERA, Tachinidae: Lixophaga mediocris Towns. LSi7, ly: Figs. 1-4. Haltichella rhyacionia Gahan. 1, Habi- tus of female. 2, Abdomen of male (lateral view). 3, Antenna of male (lateral view). 4, Abdomen of Hal- tichella perpulcra female (lateral view). Scale line 1.5 mm. Haltichella xanticles (Walker) Fig. 5 Hockeria xanticles Walker, 1843: 147, @. Haltichella xanticles (Walker); Walker, 1846: 7. Conura Xanticles (Walker); Walker, 1871: 41. Haltichella americana Howard, 1885: 36: Burks, 1975: 165. Haltichella xanticles (Walker); Burks, 1975: 165, Lectotype designation. Haltichella xanticles (Walker), MALE DI- AGNOSIS. Diagnosis.—H. xanticles is similar to H. rhyacionia, but differs in coloration and the 158 o—ONATAS »—ORNATICORNIS *—PERPULCRA a—RHYACIONIA e—XANTICLES Fig. 5. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution of Nearctic species of Haltichella. One species’ range includes the Neotropical region. A symbol in a state, province, or country indicates the species is widely distributed in that region. sculpture of Tl dorsally. The characters: length of T2 medially greater than '4 that of T1, Tl dorsally polished, with three or usually more longitudinal carinae at base, hyaline forewing, and black tegulae and legs are distinguishing. Female.— Black, with base and apex of fore and middle tibiae, apex of hindtibiae, and tarsi orange. Male.—Like female except antennae ro- bustly filiform and abdomen shorter, with apex blunt. Variation. —(2 and 4) Length 24 mm. The hindfemur is rarely dark red-brown. Type and voucher specimens.—Lecto- type 2 with data: “B.M. Type Hym. 5.554, 1478a, St. Jon’s Bluff, Hockeria Xanticles Walker.” I designate a male specimen as a voucher specimen—red label marked: “VOUCHER MALE, Haltichella xanticles (Walker), 4, det. J. A. Halstead 1987” and with data: “Mill Valley, Marin Co., Cal., 2. VII. 50, H. B. Leech Collector, Collection of the CALIFORNIA ACADEMY OF SCI- ENCES, San Francisco, Calif.’ Both spec- imens in the USNM. Hosts.—LEPIDOPTERA, Lymantriidae: Lymantria dispar (L.). Olethreutidae: Rhy- acionia buoliana (Schiff.), Rhyacionia sp., Grapholitha molesta. Gelechiidae: Exote- leia pinifoliella (Chamb.). Coleophoridae: Coleophora laricella (Hbn.). Lyonetiidae: Bucculatrix canadensisella (Chamb.). Psy- chidae: Solenobia walshella Clem. Pyrali- dae: Diorctria disclusa Heinr. HY MENOP- TERA, Braconidae: Cotesia melanoscelus VOLUME 92, NUMBER 1 (Ratzeburg). Label data on one specimen (Texas, Westaco) indicates emergence with Antonina graminis (Mask.) (Homoptera: Pseudococcidae) from a breeding cage. Another specimen (Maine, Mtn. Desert Is.) was bred from a parasite of Coleophora sal- mani Heinr. (Coleophoridae). ACKNOWLEDGMENTS I thank the museums, institutions, and individuals who graciously loaned speci- mens. I specially thank R. D. Haines, Tulare County Agricultural Commissioner’s/Seal- er’s Office, Visalia, California and D. J. Bur- dick, California State University, Fresno, for collecting specimens for this revision and for reviewing this manuscript. I thank also W. J. Pulawski, California Academy of Sci- ences, San Francisco, and W. J. Hanson, Utah State University, Logan, for permis- sion to deposit voucher males in other mu- seums. I thank E. E. Grissell, Systematic Entomology Laboratory, ARS-USDA, % USNM for working facilities during my USNM visit, and J. S. Noyes, British Mu- seum of Natural History for loaning the ho- lotype of H. ornaticornis. I thank also an anonymous reviewer for editorial com- ments. LITERATURE CITED Ashmead, W. H. 1887. Studies on the North Amer- ican Chalcididae, with descriptions of new species, chiefly from Florida. Trans. Amer. Entomol. Soc. 14: 183-203. Boucek, Z. 1951. The first revision of the European species of the family Chalcididae (Hymenoptera). Acta Entomol. Mus. Natl. Pragae 27(Suppl. 1): 108 pp. Burks, B. D. 1975. The species of Chalcidoidea de- scribed from North America north of Mexico by Francis Walker (Hymenoptera). Bull. Brit. Mus. Nat. Hist. (Ent.) 32(4): 137-170. 1979. Chalcididae, pp. 860-874. In Krom- bein, K. V. et al., eds., Catalog of Hymenoptera in America North of Mexico. Vol. I. Smith. Instit. Press. Wash., D.C. 1198 pp. Cameron, P. 1884. Biologia Centrali-Americana. In- secta. Hymenoptera. (Families Tenthredinidae— Chrysididae). Vol. I. 487 pp. 159 Cresson, E. T. 1862. Catalog of the described species of North American Hymenoptera. Proc. Entomol. Soc. Phil. 1: 227-266. DeSantis, L. 1979. Catalago de los himenopteros cal- cidoideos de America al sur de los Estados Unidos. Comision de Investigaciones Cientificas de la Pro- vincia de Buenos Aires, La Plata, Argentina. 1- 488. Freeman, B. L. and C. W. Berisford. 1979. Abun- dance and parasitic habits of some parasitoids of the Nantucket pine tip moth (Lepidoptera: Tor- tricidae). Canad. Entomol. 111: 509-514. Gahan, A. B. 1927. Four new Chalcidoid parasites of the pine tip moth, Rhyacionia frustrana (Com- stock). J. Agric. Res. 34(6): 545-548. Habu, A. 1960. A revision of the Chalcididae (Hy- menoptera) of Japan, with descriptions of sixteen new species. Bull. Nat. Inst. Agric. Sci., ser. C, No. 11: 131-363. 1962. Fauna Japonica: Chalcididae, Leuco- spididae and Podagrionidae (Insecta: Hymenop- tera). Biogeogr. Soc. Japan, Nat. Sci. Mus., Tokyo, Japan. 232 pp. Howard, L. O. 1885. Descriptions of North Ameri- can Chalcididae from the collection of the U.S. Department of Agriculture and of Dr. C. V. Riley, with biological notes. Bull. Bur. Entomol., U.S. Dept. Agric. 5: 5-47. Nikolskaya, M. N. 1952. Chalcid fauna of the U.S.S.R. (Chalcidoidea). Zool. Inst. Akad. Nauk SSR. Mos- cow No. 44, 240 pp. 1960. Fauna USSR: Hymenoptera, Vol. VII (No. 5). Chalcidoids, families Chalcididae and Leucospidae. Zool. Inst. Akad. Nauk SSR, Mos- cow (n.s.) No. 76, 221 pp. Peck, O. C. 1963. A catalog of the Nearctic Chalci- doidea (Insecta: Hymenoptera). Canad. Entomol. Suppl. 30: 1092 pp. Schmitz, G. 1946. Exploration du Parc National Al- bert, Mission G. F. De Witte (1933-1935), Chal- cididae (Hymenoptera: Chalcididae). Inst. Des Parcs Nat. Du Congo Belge, Bruxelles, 48: 191 pp. Spinola, M. M. 1811. Essal d’une nouvelle classifi- cation generale des Diplolepaires. Ann. Mus. Hist. Nat. Paris, 17: 147-148. Thomas, C. 1881. Noxious and beneficial insects of the state of Illinois. State Entomol. Rpt. 10: 5-43. Walker, F. 1843. Description des Chalcidites trou- vees au Bluff de Saint-Jean, dans la Floride ori- entale, par MM. E. Doubleday et R. Forester. Soc. Entomol. France, Annals (2)1: 145-162. 1846. List of the specimens of Hymenopter- ous insects in the collection of the British Museum. Part I—Chalcidites. vi, 100 pp. London. 1871. Notes on Chalcidiae. 129 pp., 62 figs. London. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 160-165 ISOZYME ANALYSIS IN SIX POPULATIONS OF PEDIOBIUS FOVEOLATUS (CRAWFORD) (HYMENOPTERA: EULOPHIDAE) AKEyY C. F. HUNG AND PAUL W. SCHAEFER (ACFH) Beneficial Insects Laboratory, Agricultural Research Service, USDA, Beltsville, Maryland 20705; (PWS) Beneficial Insects Research Laboratory, Agricultural Research Service, USDA, Newark, Delaware 19713. Abstract. —Sixteen enzyme loci were used to determine the degree of genetic variability and taxonomic status of Pediobius foveolatus (Crawford) (a hymenopterous parasite of Epilachna spp.) from Japan, Korea, China, Hong Kong, Guam, and India. Two loci, ACON-1! and MDH-1 were found to be polymorphic. However, no intra-population variation was found for any of the 16 loci studied. Based on these isozyme analyses, three biotypes were recognized in P. foveolatus, namely, the China biotype with its unique ACON-1 genotype, the Northern biotype with MDH-1° allele, and the Southern biotype with MDH-I' allele. Key Words: Pediobius foveolatus (Crawford) is a gre- garious parasite of larvae of Epilachna spp. (Coleoptera: Coccinellidae). It is found nat- urally in widespread areas of the Asiatic, Australasian, and African regions (Kerrich 1973). In recent years it has been reported in Japan (Tachikawa 1976), Sumatra (Ab- bas and Nakamura 1985), and China (Schaefer et al. 1986). Recent collections have recovered it in South Korea and in Hong Kong (Schaefer, unpubl. data). In the early 1950s, under the name Pleu- rotropis epilachnae Rohwer, this parasite was intentionally introduced into Guam from the Philippines to combat the introduced Epilachna philippinensis Dieke (Peterson 1955). It was introduced into North Amer- ica from Bangalore, India, for the control of Mexican bean beetle (MBB), Epilachna varivestis Mulsant, in 1966 (Angalet et al. 1968). It failed to overwinter in the U.S. but has repeatedly shown promise as it readily attacks MBB larvae during the season of its allelic repression, biotype, electrophoresis, Epilachna release. In the belief that a race from tem- perate areas might be capable of surviving winters in North America, one of us (P.W.S.) obtained P. foveolatus (hereafter referred to as Pediobius) from Honshu, Japan, and first released it in North America in 1980 but this race also did not permanently establish (Schaefer et al. 1983). We have concluded that Pediobius probably requires an alter- nate host Epilachna sp. which overwinters as a larva or pupa. In parts of Asia there are some species with this trait. The only three species in the Epilachninae in the eastern U.S. all overwinter as adults. At the time of the release of the Japanese Pediobius, we attempted to distinguish this race from the Indian one which, at the time, was being released in widespread projects in several mid-Atlantic states (thus neces- sitating our releases in the Mississippi River valley). Being unable to distinguish these two races morphologically, we turned to electrophoretic analysis to provide a marker VOLUME 92, NUMBER | Table |. 161 History of cultures of Pediobius foveolatus at BIRL. _—_ eee eee ee Designation Locality No. Generation Date of Receipt Starter Sample Size in Laboratory —_—_—_—_—_——_——_.:. nn Japan (A) Kurashiki, Honshu 8/15/79 2878 M&F 131 Japan (B) Yashiro, Honshu 8/27/80 1497 M&F 108 Korea Seoul 10/04/82 259 M&F 63 China Bering & Taiyuen 11/07/84 3M, 79F 23 Hong Kong Fanling, New Territory 8/05/82 2271 M&F | 44 Fanling, New Territory 8/06/83 352 M&F | Guam Mangilao /24/85 18M, 86F 11 - Devanahalli, Bangalore 5/24/72 398 M&F a) Avali, Bangalore 7/5-8/21/73¢ 2814 M&F | 23 India (B) Re-colonized subculture from (A) (Same as above) 286° @M = males, F = females. * Only one of four 1972 shipments (with about 30 females) contributed to the laboratory culture. © Four shipments including 1596 females. ¢ Approximate generation number based on calculations. which might identify the Japanese popula- tion. As more races became available, we extended this study into an assessment of the genetic heterozygosity of Pediobius from widely scattered geographical locations (Ja- pan, Korea, China, Hong Kong, India, and Guam) as well as some of the inter-cross races maintained in culture at Beneficial In- sects Research Laboratory (BIRL), Newark, Delaware. We report on the electrophoretic means of distinguishing some populations from others and on the overall genetic het- erozygosity of this species based on reared material which originated from the loca- tions given. MATERIALS AND METHODS Living material was obtained from cul- tures being reared at BIRL. The origin and brief history of each culture, date(s) of im- portation, starter sample size, and the gen- eration number (or approximation) of each culture are presented in Table |. The ulti- mate rearing procedure was to maintain each culture in unwaxed paper cups (112 mm ID at top, 55 mm deep, capacity 470 ml) with clear plastic lids. Sting units were set up approximately every two to three weeks by aspirating ca. 20-30 adults (ca. equal sex ratio) from old cups and expelling them into new cups which contained 25 fourth instar MBB larvae. These sting units sat for ca. 6 h, after which all Pediobius were collected by aspiration and destroyed. Care was taken never to return specimens into cups once they were removed or escaped. This was done as a precaution to prevent any genetic mixing of indistinguishable wasp strains. The host larvae were then transferred to reusable rigid polyethylene cylindrical cages (158 mm ID, 120 mm deep) with three fine mesh screen portals (5 cm dia.) on the sides for air circulation and inverted glass pie plates for covers. Food was provided as a bouquet of Tendergreen snapbean plants in a glass vial of water with cotton plugging firmly holding the plant stems. Food was replaced during the first week if needed. During the second week host larvae were transferred into new paper cups to await adult emergence. Just prior to expected emergence (two to three weeks depending on the seasons since ambient indoor tem- perature and humidity was not regulated), the inner surface of the lid was smeared with honey as the only food source. Water was not usually provided but on occasion if drying was evident, water was injected through the cup wall using a hypodermic needle. Material used in the electrophoretic anal- ysis was taken from the cup after a subsam- 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Gel and electrode buffer systems. Designation Description Source MC N-(3-Aminopropyl)-morpholine-citrate (pH 6.0) Clayton and Tretiak (1972) PK Discontinuous Tris-citrate (Electrode: pH 8.2; Gel: pH 8.7) Poulik (1957) LG Tris-citrate (pH 7.0) Siciliano and Shaw (1976) TVB Tris-versene-borate (pH 8.0) Siciliano and Shaw (1976) ple was removed to begin the next genera- tion. Specimens in their intact paper cups were shipped or hand-carried to Beneficial Insects Laboratory (BIL), Beltsville, Mary- land, and samples were then frozen and stored at —65°C until electrophoresis. Electrophoretic Methods Electrophoresis was performed on hori- zontal starch gels for 4 h using 6% Elec- trostarch, 6% Sigma starch, 5% sucrose, 10 mg NADP, and 400 ml gel buffer (see Table 2). The 12 enzyme systems examined are listed in Table 3, each followed by the buffer systems used. At least six female and six male wasps per enzyme per culture were electrophoresed. Sample sizes were in- creased to over 20 for loci that exhibited polymorphisms. At least 18 female progeny were used in the controlled progeny analyses of genotypes at ACON-1 and MDH-1 loci. We assumed that discrete zones of en- zyme activity were controlled by single loci coding for specific products. The genetic ba- sis of observed isozyme variations (see ACON and MDH below) were confirmed by progeny analyses. Loci coding for the same enzyme are numbered sequentially from the most anodal to the most cathodal regions of activity. The allelic proteins are designated alphabetically, with ‘‘A” the fastest running allele. The allelic designa- tions in ACON-1 and MDH-1 were inferred from progeny analyses and were labelled as SES? OLS RESULTS AND DISCUSSION In addition to the enzymes shown in Ta- ble 4, we also examined acid phosphatase, aldehyde oxidase, esterase, galactose-6- phosphate dehydrogenase, glutamate de- hydrogenase, leucine aminopeptidase, and superoxide dismutase. However, these en- zymes had either very weak or streaky bands Table 3. Enzymes analyzed in P. foveolatus, with buffer conditions employed. EC* no. Enzyme Buffer System 4.2.1.3 aconitase (ACON) MC 1.1.1.49 glucose-6-phosphate dehydrogenase (G6PD) MC 2.6.1.1 glutamate-oxaloacetate transaminase (GOT) MC a) 3 alpha-glycerophosphate dehydrogenase (GPDH) TC, DVB Qe hexokinase (HK) PK, TVB LET 42 isocitrate dehydrogenase (IDH) AKG Ie) an 777 lactate dehydrogenase (LDH) ING: eS) malate dehydrogenase (MDH) MC, TC 1.1.1.40 malic enzyme (ME) ANG; 1.1.1.44 6-phosphogluconate dehydrogenase (6PGD) MC 5.3.3.9 phosphoglucose isomerase (PGI) MC, TC D2) phosphoglucomutase (PGM) MC, PK * Enzyme Commission. VOLUME 92, NUMBER 1 Table 4. Isozyme phenotypes of six Pediobius fove- olatus populations. Population** Enzyme Hong locus Japan Korea China Kong Guam India ACON-1 lala EE SS FE FE FE ACON-2 AA AA AA AA AA_ AA HK-2 AA AA AA AA AA_ AA IDH AA AA AA AA AA _ AA LDH AA AA AA AA AA_ AA ME AA AA AA AA AA_ AA PGM AA AA AA AA AA_ AA PGI AA AA AA AA AA_ AA 6PGD AA AA AA AA AA _ AA ** Japan (A) and Japan (B) were combined as there were no differences between them. India (A) and India (B) were also combined for the same reason. that could not be clearly scored. Therefore, only data on 12 enzyme systems were used in this analysis (Table 4). All the enzymes in this study migrated anodally with the buffer conditions em- ployed except ACON-2, MDH-2 and 6-PGD which migrated cathodally (MDH-1 and MDH-2 also migrated cathodally in the MC buffer). Only four enzyme systems had more than two loci and with the excep- tion of ACON-1 and MDH-1, all other loci were monomorphic (Table 4). Progeny analyses also revealed that ACON is mo- nomeric and MDH is dimeric in Pediobius. Although the five populations we studied were identical at 14 loci, there are some differences at the other two loci. As shown in Table 4, two alleles were found at both ACON-1 and MDH-1 with each population fixed for one of the two alleles at each locus. The China population was unique in being fixed for the slow allele at ACON-1 while the others were fixed for the fast allele at this locus. Guam, Japan, and Korea popu- 163 lations were fixed for the MDH-1> allele while samples from China, Hong Kong, and India were fixed for the MDH-1' allele. Although Hung et al. (1986) reported high levels of genetic heterozygosity in the hy- perparasitic wasp, Mesochorus nigripes, very low levels of electrophoretic variation have been found in most hymenopteran species. Some species even lack variation altogether (Wagner and Briscoe 1983). The genetic ho- mogeneity found within each population at these 16 loci possible is not the result of founders effects, because even the smallest sample (China culture) originated from 79 female wasps collected from two widely sep- arated localities. This lack of enzyme vari- ation might be due to inadvertent selection during long periods of laboratory rearing as in the case of the screwworm fly (Bush et al. 1976). Since the starter materials were not analyzed electrophoretically, the valid- ity of this assumption cannot be ascer- tained. Pleurotropis epilachnae, described as a separate species from India by Rohwer, was synonymized under Pediobius foveolatus (Crawford) by Kerrich (1973), because the differences in size and color did not hold up in specimens other than the type series. Therefore, our samples were all identified as P. foveolatus and no morphological dif- ferences were found among them (M. E. Schauff, pers. comm.). According to Peterson (1955), Pleurotro- pis epilachnae Rohwer (= P. foveolatus) was successfully introduced into Guam from the Philippines during 1954 to control the phy- tophagous ladybeetle, Epilachna philippin- ensis Dieke. The BIRL quarantine records also show that the only recorded shipments to Guam occurred in 1974 (three ship- ments) and 1975 (three shipments) and are recorded as origin “India.” The Japan cul- ture was first obtained in August 1979, and has never knowingly been imported into Guam. Therefore, it is rather puzzling that the Guam population was the same as the Japan and Korea populations in having only 164 the slow allele at MDH-1. Over 30 female wasps from the Guam culture analyzed all had only the slow allele and it is not likely that this culture was contaminated. We can- not explain the apparent contradiction be- tween our findings and the implications based on historical records. Our cultures of these six populations crossed successfully in both directions un- der laboratory conditions. Peng (1988) also reported that there was no reproductive 1so- lation between his Beijing Pediobius and those from Hong Kong, India, Japan, and Korea that he received from PWS. How- ever, he reported that ““malic dehydroge- nase (sic)’’ was one of three enzymes that were monomorphic in the five cultures he studied. This is different from our results as shown in Table 4. It is possible that the lack of variability in his malate dehydrogenase study is due to the poor resolution of the buffer system and supporting medium he used (see Hung and Vinson 1977). Four patterns of allelic repression have been reported in interspecific as well as in- tertribal hybrids (Avise and Duvall 1977, Hung and Vinson 1977): (a) repression of paternal protein synthesis; (b) repression of maternal protein synthesis; (c) repression of both maternal and paternal protein synthe- sis; and (d) caste specificity of differential parental protein synthesis in social insects. Although the slow allele at the ACON-1 lo- cus was found only in the China population, our controlled progeny analyses of geno- types at ACON-1 locus did not reveal any allelic repression as is frequently observed in interspecific hybridization (e.g. Hung and Vinson 1977, Hung 1985, Hung and Nor- den 1987). Both maternal and paternal genes were fully expressed in F, and F, progeny in our two-way crosses. Therefore, we do not recognize the China population as a sep- arate species. Differences in the percentage of ovipositing females, average fecundity and longevity of females, and number of hosts parasitized were found between the Indian and the Japan “races” of P. foveo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON latus (L. Nong, pers. comm.). However, we believe these biological differences only sig- nify that they are two biotypes. Based on these isozyme analyses, we concluded that there are three biotypes in P. foveolatus: I. China biotype with its unique ACON-1 ge- notype; II. Northern biotype (Japan and Korea) with MDH-1° allele; and III. South- ern biotype (India and Hong Kong) with MDH.-I' allele. The introduced population from Guam apparently belongs to the northern biotype contrary to the expecta- tions based on the historical shipment rec- ords. ACKNOWLEDGMENTS We thank Joseph M. Tropp and David L. Vincent for technical assistance. We also thank Alan C. Bartlett, Michael E. Schauff, and Gary J. Steck for reviewing the manu- script. LITERATURE CITED Abbas, I. and K. Nakamura. 1985. Adult population parameters and life tables of an epilachnine beetle (Coleoptera: Coccinellidae) feeding on bitter cu- cumber in Sumatra. Res. Pop. Ecol. 27(2): 313- 324. Angalet, G. W., L. W. Coles, and J. A. Stewart. 1968. Two potential parasites of the Mexican bean beetle from India. Jour. Econ. Entomol. 61: 1973-1975. Avise, J. C. and S. W. Duvall. 1977. Allelic expres- sion and genetic distance in hybrid macaque mon- keys. J. Hered. 68: 23-30. Bush, G. L., R. W. Neck, and G. B. Kitto. 1976. Screwworm eradication: Inadvertent selection for non-competitive ecotypes during mass rearing. Science 193: 491-493. Clayton, J. W. and D. N. Tretiak. 1972. Amine-ci- trate buffers for pH control in starch gel electro- phoresis. J. Fish. Res. Board Can. 29: 1169-1172. Hung, A.C. F. 1985. Isozymes of two fire ant species and their hybrid. Biochem. Syst. & Ecol. 13: 337- 339. Hung, A. C. F., R. C. Hedlund, and W. H. Day. 1986. High level of genetic heterozygosity in the hyper- parasitic wasp, Mesochorus nigripes. Experientia 42: 1050-1051. Hung, A. C. F. and B. B. Norden. 1987. Biochemical systematics of bees in the Ceratina calcarata—du- pla complex. Biochem. Syst. & Ecol. 15: 691-693. Hung, A. C. F. and S. B. Vinson. 1977. Interspecific VOLUME 92, NUMBER 1 hybridization and caste specificity of protein in fire ants. Science 196: 1458-1460. Kerrich, G. J. 1973. A revision of the tropical and subtropical species of the Eulophid genus Pedio- bius Walker (Hymenoptera: Chalcidoidea). Bull. British Mus. (Nat. Hist.) Entomology 29(3): 133- 199. Peng, H. 1988. Preliminary studies on the taxonomic status of Pediobius spp. emerging from larvae and pupae of the potato 28-spotted beetle, Henose- pilachna viginiioctomaculata (Motschulsky) (Ab- stract). Proc. XVIII Intern. Congr. Entomol. p. 52. Peterson,G.D. 1955. Biological control of Epilachna philippinensis Dieke in Guam. J. Econ. Entomol. 48: 758-759. Poulik, M. D. 1957. Starch gel electrophoresis in a discontinuous system of buffers. Nature 180: 1477. Schaefer, P. W., R. J. Dysart, R. B. Flanders, T. L. Burger, and K. Ikebe. 1983. Mexican bean beetle (Coleoptera: Coccinellidae) larval parasite Pedio- 165 bius foveolatus (Hymenoptera: Eulophidae) from Japan: Field release in the United States. Environ. Entomol. 12: 852-854. Schaefer, P. W., H. Peng, and X. Gou. 1986. Prelim- inary survey of parasites of Epilachna sp. in China. (in Chinese, Engl. summ.) Chinese J. Biol. Control 2(8): 112-115. Siciliano, M. J. and C. R. Shaw. 1976. Separation and visualization of enzymes on gels. Jn I. Smith, ed., Chromatographic and electrophoretic tech- niques. Vol. 2, William Heinemann Med. Books Ltd., New York, pp. 185-209. Tachikawa, T. 1976. Occurrence of Pediobius foveo- /atus (Crawford) (Hymenoptera: Eulophidae) parasitic on Henosepilachna (Coleoptera: Cocci- nellidae) in Japan. Trans. Shikoku Entomol. Soc. 13(1/2): 61-63. Wagner, A. E. and D. A. Briscoe. 1983. An absence of enzyme variability within two species of T7i- gona (Hymenoptera). Heredity 50: 97-103. INSTRUCTION TO AUTHORS FOR PREPARATION OF MANUSCRIPTS GENERAL POLICY Publication in the Proceedings is gener- ally reserved for members. Manuscripts should be in English and not be so lengthy that they would exceed 15 printed pages in- cluding illustrations (two typewritten pages are approximately equivalent to a printed page.) Manuscripts are peer-reviewed be- fore they are accepted. Acceptance of manu- scripts is the responsibility of the Editor. Papers are published in the order they are received rather than in order of date of ac- ceptance. This eliminates possible bias due to the varying length of time taken to review a paper. Notes and book reviews are pub- lished as space is available, usually in the next issue prepared. Immediate publication can be had for payment of full page charges, but this provision should be reserved for papers with some justification for expedited handling. These papers do not lengthen the waiting period of regular manuscripts be- cause they are published in addition to the regularly budgeted number of pages. TYPING THE MANUSCRIPT Type manuscripts on bond paper with double-spacing and ample margins. Num- ber pages consecutively beginning with the title page. Do not use all capitals for any purpose. Underscore only where italic type is intended in the body of the text. Type on separate pages the title page, abstract and key words, literature cited, tables, and figure legends. See Figs. 1-4 for proper format for most of those pages. SUBMITTAL OF THE MANUSCRIPT Submit the original manuscript and two copies to the Editor. State membership sta- tus in a cover letter. Onginal drawings should be retained until the manuscript is accepted. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 166-168 NAMES OF ORGANISMS The first mention of a plant or animal should include the full scientific name with the author of a zoological name unabbrevi- ated. Use only common names approved in Common Names of Insects and Related Or- ganisms (1983 Revision) published by the Entomological Society of America. Do not abbreviate a generic name when it begins a sentence: e.g. use “Xus albus is . . .” rather thanvsXeralDusiSesrcrs CITATIONS When citations are made in the text, a space separates author from date. A comma separates citations. Examples: (Smith 1976), (Smith and Jones 1972), (Smith et al. 1980), (Smith 1970, Roberts 1971, Jones 1985), (Smith 1971, 1972). ILLUSTRATIONS No extra charge is made for line drawings or half tones. Authors should plan illustra- tions for reduction to the dimensions of the printed page and allow room for legends at the top and bottom. Do not make plates larger than 14 x 18”. 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Doe A new species of Family) Xus (Order injurious to hollies, Ilex spp. (Aquifoliaceae) (JRD) Resident Biologist, 315 State St., Meriden, Connecticut 06420; City Parks, (JS) Entomologist, Hartford, Connecticut 06540 Xus albus, a new species of is described, illustrated, Abstract.- ompared with and Key Words. Distribution, ornamental shrub, damage, leaf roller enitalia (lateral view) Fig. 5. Damage to holly leaves Literature Cited Holly Insects Entomol (The above citations are fictitious Figs. 1-4. Sample pages for submitted manuscripts. 1, Title page. The mailing address and phone number of the person(s) to whom the page proofs will be sent should be typed in the upper left-hand corner. A proof can be sent to each author of a two- or multi-authored paper if they desire. Affiliation and full address are typed as a paragraph below the author name(s). Spell out the state name. 2, Abstract page. Note that “Abstract” begins the paragraph. On same page, follow abstract with 3-5 key words. 3, Figure legend page. Note punctuation. Start a new paragraph for legends describing each plate. 4, Literature Cited page. Note punctuation and spacing. Authors with access to Serial Sources for the Biosis Data Base that is provided with Biological Abstracts are expected to use its journal title abbreviations. 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON except for printer’s and editorial errors, will be charged to the author. CHARGES Authors will be assessed a page charge to cover editing and publication expenses. These charges are in addition to those for reprints and author’s correction in proof. Charges are at a higher rate for an excess of printed pages over 15, papers by non-mem- bers, and papers for immediate publication. Member authors who are retired or not af- filiated with an institution may request to have charges waived and should do so when the manuscript is submitted. A waiver may delay publication. Charges for author errors and/or changes in proof, for reprints if or- dered, for immediate publication, and for non-members are not waived. Acceptance of papers is based only on their scientific merit without regard to the author’s finan- cial support. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 169-170 Book REVIEW Combating Resistance to Xenobiotics: Bi- ological and Chemical Approaches. M. G. Ford, D. W. Holloman, B. P.S. Khambay and R. M. Sawicki. Ellis Horwood Series in Biomedicine, Ellis Horwood Publish- ers, 320 pp. The book is essentially the published out- come of a conference held by the Society of Chemical Industry at Southhampton Uni- versity in 1986. The purpose of the confer- ence was to discuss the contribution fun- damental studies can make to the understanding of pesticide resistance and review how knowledge was being utilized to overcome the problem. The editors have brought together a broad selection of topics addressing these goals. In addition to en- tomological applications, the book encom- passes resistance to antibiotics and a variety of pesticides including fungicides, and her- bicides in addition to insecticides. In the forward, D. W. Holloman states that ex- perts from several disciplines sought to link themes of their work on resistance to var- ious antibiotics and pesticides. Accordingly the book is intended to take an interdisci- plinary approach. The introductory chapter “Resistance to pesticides and antibiotics: how far is it com- prehensible and manageable?” by I. J. Gra- ham-Bryce does indeed take an interdisci- plinary tact, but thereafter the subject matter is presented in a more compartmentalized manner. I will focus my comments on the utility of this opus to entomologists. Of 25 chapters, 10 exclusively address insecticides and two cover insecticides within the broad- er pesticide context. The parts of the book on resistance to insecticides, mechanisms of resistance, and structure-activity relationships provide some good general background in these sub- ject areas and some thorough specific in- formation on management of selected pests (e.g. Heliothis armigera, Lucilia cuprina, and Haematobia irritans). For the beginning student the chapter, by R. Sawicki on def- inition, detection and documentation pro- vides a concise introduction to terms, pa- rameters, and measurement of resistance. An introduction to the decision making pro- cesses for management of resistance in in- sect pests, by T. J. Dennehy, complements the chapter by Sawicki. The author encour- ages the reader to know thy: system (is the bioassay appropriate to the agricultural sys- tem?); chemical (in addition to mortality, does it have sublethal effects? effects on the rest of the ecosystem? etc.); and target or- ganism (distribution, biology, behavior, stress factors). A very brief portion of the chapter is devoted to the decision making process per se. It should be construed more as an outline than a protocol in this regard. Using Musca domestica as their model, I. Denholm et al. present an excellent chap- ter on laboratory simulation of selection for resistance including a short section on com- puter modeling of selection rates. They em- phasize the importance of understanding the various biological and operational param- eters acting at the insect-insecticide inter- face before selection rates can be accurately modeled. A short introduction on the genetic as- pects of selection for resistance is given by C. F. Curtis. Although the chapter provides a good introduction to the use of numerical and computer models in the genetics con- text, it does not provide the breadth one is led to expect from its title. The value of pesticide rotation for management of resis- tance 1s a useful addition to this chapter and broadly covers the genetic implications of this strategy. The three chapters on specific resistance management strategies for Heliothis armi- 170 gera, Lucilia cuprina, and Haematobia ir- ritans are among the best in the book. Al- though they each address only one species they give the reader an appreciation for the complexity of the problems and the atten- tion to detail that must be exercised in pro- ducing a sustainable solution. In part 5, Mechanisms of Resistance, only two chapters cover resistance in insects. The chapter by A. C. Baillie is a good introduc- tion to how insecticides work, but falls a bit short on why they fail. Five short para- graphs are devoted to resistance and one to overcoming it. Conversely, the chapter by A. L. Devonshire on biochemical studies of organophosphorus and carbamate resis- tance in houseflies and aphids provides a detailed account of research conducted on this subject at the Rothamsted Experimen- tal Station. Although not a review of all the mechanisms responsible for resistance in the two test insects to these two classes of in- secticides, it does acquaint the reader with an in depth appreciation for two of the mechanisms (e.g. insensitive acetylcholin- esterase and insecticide detoxifying ester- ase) and their measurement. In the remaining part of the book, struc- ture-activity relationships, only one of the three chapters covers insecticides. In a con- cise chapter on selectivity and resistance to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON non-ester pyrethroids and N-alkylamides in houseflies, A. W. Farnham et al. give the reader an understanding of structural effects on insecticidal activity in this increasingly important group of insecticides. One can truly say this book has something for everyone that may be concerned with resistance to one substance or another. It may have, however, provided excessive material for the IPM specialist concerned only with combating resistance in insects. On the other hand the beginning toxicology student would benefit from the information provided in the non-insecticide portions of the book as well as those dealing with in- secticides. However, the book is fairly lim- ited in the depth required for an introduc- tory toxicology course. It should be supplemented with greater detail in the area of structure and activity, the genetics of re- sistance, and the mechanisms responsible for it. Examples are for the most part drawn from studies of medically important insects. A more detailed sequel should include more examples of resistance in agricultural insect pests. Lawrence A. Lacey, Vector Biology and Control Project (USAID/MSCI), 1611 N. Kent St., Suite 503, Arlington, Virginia 22209. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 171-172 Book REVIEW Novel Aspects of Insect-Plant Interac- tions. Edited by Pedro Barbosa and Deb- orah K. Letourneau. Wiley-Interscience Publications, John Wiley & Sons; New York, Chichester, Brisbane, Toronto, Singapore. 1988. Price: $59.95. Sixteen scientists have written nine arti- cles on the subject of allelochemical inter- actions with nearly all of them from the insect ecology point-of-view. The book is divided into four parts each with an intro- duction by Letourneau. Letourneau says that “... this volume represents the forefront of two rapidly advancing areas of ecology: three-trophic level interactions and chemical ecology” (p. 6). Most of the chap- ters deal with predation and herbivory. Part I, Conceptual Framework of Three- Trophic-Level Interactions, contains two chapters which discuss plants, insects and allelochemicals. Chapter one is by D. Whit- man and is on allelochemical interactions among the three-trophic levels. This chap- ter categorizes and gives examples of dif- ferent types of interactions. The second chapter, by D. Nordlund et al., is on alle- lochemicals and selection behavior of en- tomophagous insects. Part II, Microorganisms as Mediators of Intertrophic and Intratrophic Interactions, contains two chapters on the complex in- volvement of microbes in the interactions between macroscopic organisms. Coevolu- tion of microbes and the insects and mi- crobes and the plants is discussed by M. Berenbaum. M. Dicke tackles the problem of microbial allelochemicals that affect predator behavior. Part II, Theory and Mechanisms: Plant Effects via Allelochemicals on the Third Trophic Level, discusses the effects of al- lelochemicals on the food chain. H. Wil- liams et al. cover the topic of parasitoids in the three-level system, emphasizing cotton. Barbosa, in the second paper, proposes the hypothesis for parasitoids, that generalists will suffer greater losses than will specialists from exposure to plant-derived toxins in their hosts. Part IV, Key Roles of Plant Allelochem- icals in Survival Strategies of Herbivores, covers two topics. The first two chapters discuss the ecology of unpalatable prey. J. Pasteels et al. discuss plant-derived defenses in chrysomelid beetles and M. Bowers dis- cusses plant allelochemistry from the stand- point of mimicry. Finally, the last chapter, which was written by L. Brattsten, brings into the discussion the somewhat unrelated topic of man’s interference, insecticide re- sistance. Most of the articles are up-to-date re- views and they all appear to have excellent reference sections that will be of use to any- one trying to gain some insight into the lit- erature. I think all but the very well in- formed allelochemical investigator could learn quite a lot reading the volume. Most of the chapters have an introduction that defines terms and they all end with a short discussion on the future. However, I had two problems with this volume, first as a botanist and a systematist I had difficulty with the terminology, much of which was new to me. I found that different authors meant different things although they used the same terms. This is to be expected but because there is no glossary and only a very short subject index I had to search back and forth through the text for definitions. Also, there is no citation index so if one 1s trying to find if or where a certain reference is cited then one has to look in the reference section of every chapter and then search through that chapter until the location(s) is found. For instance, I wanted to know if Q. Wheel- er’s fungi-beetle interaction research had 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON been cited in the text and if so in what con- text. The second, and more serious, problem I encountered is that as a systematist I kept asking questions about whether or not these various chemicals were historical, inherited from an ancestral taxon and thereby plesio- morphic, or autapomorphic and therefore candidates for consideration as possible adaptive scenerios. This information was not available and I was left feeling disap- PROC. ENTOMOL. SOC. WASH. 92(1), 1990, p. 172 Book Cyanide Compounds in Biology. Edited by David Evered and Sara Harnett. John Wiley and Sons Ltd., Chichester, U.K. 1988, IX + 261 pp., $54.95. This volume covers all of the 15 papers presented at Ciba Foundation Symposium 140 held in Canterbury, England in 1988. Each of the presentations is given in toto together with the verbatim discussions fol- lowing the presentations. All presentations and discussions are in English. Included in the volume are an introduction, three pre- sentations covering the microbial metabo- lism of HCN and organic cyanide com- pounds, three presentations on cyanogenesis in higher plants, one presentation on cyan- ogenesis in insects, one on cyanogenesis in plant-animal interactions, one on methods for determining cyanide and cyanogenic compounds in biological systems, one on the influence of nutritional and biochemical factors on the biological effects of cyanide, one on the mammalian detoxification of cy- anide, one on the mechanism of cyanide intoxication and its antagonism, and, last but not least, two general discussion ses- sions and the chairman’s discussion of all pointed because it seemed that although much work had been done, basic phyloge- netics had been ignored. Ultimately the ones who suffer are those of us who wish to know, not just the facts about the current ecolog- ical relationships but also something about the evolution of the organisms involved. Vicki A. Funk, Department of Botany, Smithsonian Institution, Washington, D.C. 20560. REVIEW presentations. Each of the chapters includes a list of references given alphabetically by author. This volume should be of considerable interest to biochemists and botanists. Un- fortunately, the only presentation of direct (and considerable) interest to entomologists and insect physiologists is that covering cyanogenesis in insects, given by the re- nowned German insect biochemist Adolf Nahrstedt. However, this presentation is excellent in that it covers very well the in- formation discovered mainly since 1978 concerning the cyanogenic defensive com- pounds utilized by various beetles (Lepto- coris isolata) and several species of Lepi- doptera (Zygaena spp., Agraulis vanillae, two species of Heliconius, and Dryas julia). All graphs, line drawings, and other il- lustrations are of high quality, and I could find no grammatical or factual errors in this comprehensive book. Martin Jacobson, Insect Chemical Ecol- ogy Laboratory, USDA (Retired), 1131 Uni- versity Blvd. W., Silver Spring, Maryland 20902. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, pp. 173-175 Book REVIEW The Torre-Bueno Glossary of Entomol- ogy, compiled by Stephen W. Nichols. New York Entomological Society, % De- partment of Entomology, American Mu- seum of Natural History, Central Park West at 79th Street, New York, New York 10024. Library order, $45.00; members of the New York Entomol. Soc., $35.00; non-members, $40.00. J. B. Smith started it with his 154 page “Explanation of Terms Used in Entomol- ogy” in 1906. Then in 1937, J. R. de la Torre-Bueno produced his ‘*A Glossary of Entomology, Smith’s ‘An Explanation of Terms Used in Entomology,’ completely re- vised and rewritten.’ This was reprinted in 1950. Now we have what purports to be an amplified version of the Torre-Bueno work. By the title it was given, we may assume that its predecessor was uncritically added to, which does indeed seem to be the case. There is a list of 50 “Editorial Contribu- tors.” The original Torre-Bueno work had 323 pages of glossary, 2.1 times as much as the Smith work. The present work has 823 pages of glossary on pages of the same size and with the same size of type as in the earlier work, meaning that it is slightly more than 2.5 times as large. The old work had ap- pendices of abbreviations, symbols, and 9 plates of figures. These are lacking in the new work and no abbreviations are included in the text. This material will be greatly missed. Bigger is not necessarily better. Has the subject actually grown to the extent that the Glossary has increased? Let us make a few comparisons. The old work devoted 5 lines to ‘aedea- gus,’ and one phrase ‘aedeagal apodeme,’ based on that term. The new work has 43 lines, nearly an entire page, on ‘aedeagus’ and 24 phrases with ‘aedeagus’ and ‘aedea- gal.’ In the old work, ‘aedoeagus’ was cited with ‘v. aedeagus.’ This is as it should be because ‘aedoeagus’ is no more than a vari- ant of ‘aedeagus,’ but in the new work ‘ae- doeagus’ is cited and defined as a separate word with no cross-reference to ‘aedeagus.’ And, the entries for ‘penis’ and phrases with ‘penis’ and ‘penial’ are even more expan- sive. These and many other cases indicate to what extent the new work is uncritical. The multiplicity of cited phrases is quite evident throughout the new work, although the meanings of many of them are quite obvious from the meanings of the compo- nent terms. The many phrases with ‘lobe(s)’ on p. 407 for the same reason have little claim to inclusion, and Latin ‘lobus (pl. lobi)’ should appear merely as an equivalent of ‘lobe(s).” Much of such expansion of single term definitions results in little more than confusion. Although many of the main entries in the new work have been expanded, ‘venation’ has the same inadequate treatment it had in the old work, viz., “the complete system of veins of a wing.” It would have been helpful at this point to mention that there have been many systems of terminology for the veins of the wings of insects of various orders, all of which have been virtually superseded by the Comstock-Needham sys- tem, one that is convenient to use and ap- plied to all insect wings. Comstock-Need- ham is entered in the new work, but many workers, especially in orders where the ve- nation is highly specialized, still cling to more or less older systems, particularly in Hymenoptera. It is here that a few figures would have been especially useful. It would at least have obviated the need for making entries for all 6 of the ‘longitudinal veins’ of the old Schinerian terminology used in Diptera and show at the same time that there 174 were that many longitudinal veins. A figure would also do much to explain what are ‘recurrent veins’ (Hymenoptera, etc.). Words are included that are not or are only marginally entomological, such as ‘epi- phyte’ (botany), ‘acid’ (chemistry), and ‘mineral’ (general), as well as the many names cited together with the etiology of diseases of plants (plant pathology), and an- imals other than insects (medicine). A few truly entomological subjects have received rather short shrift. One of them is the technique of collection, examination, and preservation of insects. The following are absent: balsam, Canada balsam, Eupa- ral, Cellosolve, medium (mounting, pl. me- dia), macerate, polyporus, silicone, minuten nadel, polyethylene, McPhail trap, Steiner trap, interception trap, even the lepidopter- ists’ old standbys, light trap and light sheet. Some terminology, such as that for mus- cles, is still strictly in Latin and should be printed in italics. Latin for descriptions and other uses in entomology has, however, for at least a century been virtually restricted to nomenclature, the scientific names of or- ganisms. Therefore, citations of adjectival terms such as ‘nigrescent, nigrescens, nigri- cante’ and ‘ferrugineous, ferruginosus, fer- ruginous’ in both the old and the new Glos- saries would better have appeared respectively as ‘nigrescent (L. nigrescens, ni- gricante), and ‘ferrugineous, ferruginous (L. ferrugineus, ferruginosus).’ In other words, distinction should have been made between English words and the Latin words from which they have been derived. The defini- tions for ‘efferent’ and ‘afferent’ in both the old and the new Glossaries are poor; i1n- cluding with their definition something about their derivation in such a manner as ‘afferent (L. ad + ferens ‘toward-carrying,’ and ‘efferens (L. ex. + ferens ‘away-carry- ing’)’ in itself furnishes a good definition; it is then easy to understand that the duct from the testes to the sperm pump in Diptera is an afferent duct of the sperm pump, while the duct from the sperm pump to the ae- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON deagus is efferent, as well as that the duct from the testes is an efferent duct of the testes. The entry ‘ater, atrous, atrus’ would better have been omitted entirely; ‘ater’ is strictly Latin, ‘atrous’ is long-dead English (not in Webster III, and in the Oxford dic- tionary as rare (Nat. Hist.), and ‘atrus’ is only an erroneous form of ‘ater.’ Such words as ‘phallosome’ and ‘phallo- soma,’ instead of being entered separately as if they were distinct words, would better be treated merely as English (with -e) and Latin (from Greek) (with -a) variants of the same word and therefore having different plural forms. The following important works do not seem to have been used in the compilation of the new Glossary: Crampton, G. C. 1947. The external mor- phology of the Diptera. Guide to the Insects of Connecticut, Pt. VI, Diptera; fasc. 1: 1 10- 165 (incl. 14 Pls. of figs. and 13 pp. of bib- liography). Fernald, H. T. 1939. On type nomencla- ture. Ann. Entomol. Soc. Amer. 32: 689- 702. Hammen, L. von der. 1980. Glossary of Acarological Terminology. Vol. 1, 179 pp. Dr. W. Junk B.V., The Hague. Lindroth, C. H. 1957. The principle terms used for male and female genitalia in Co- leoptera. Opusc. Entomol. 22: 241-256. And, finally, a few items of miscellaneous comment: Although the definitions of ‘pollinose’ and ‘pruinose’ are correct, mention should have been made that they are often erroneously used for ‘microtrichose’ and ‘microtomen- tose.” ‘Meskatespisternum’ (with extra s) 1s used instead of mesokatepisternum, although McAlpine used merely katepisternum and anepisternum in the Diptera because the corresponding parts of the prothorax and metathorax are not present in that order. VOLUME 92, NUMBER 1 ‘Aculeus’ as used in the female Tephrit- idae (Diptera) is not mentioned. ‘Vertexal’ is cited correctly, but no indi- cation is made that it is sometimes used instead of ‘vertical’ because bristles, etc., so- called are on the vertex and usually are not vertical in the ordinary sense of the word. ‘Tragplatte’ is a term used in Hennig’s earlier work, but supplanted later by ‘hy- pandrium’; it is not the aedeagal apodeme. ‘Calcipala’ is an organ found in both sexes of Simuliidae. Special uses of ‘cornu’ and ‘ramus’ in the Coleoptera are not cited, nor are the simple meanings of these Latin words, viz., horn and branch, respectively. The entry ‘cornua, pl. cornuae’ should have appeared under ‘cornu’ and the fact that ‘cornua’ is already plural and that ‘cornuae’ is an incorrect dou- ble plural made evident. 175 ‘Gymnopedia’ should appear with refer- ence to ‘pedium.’ “Mesophoba’ should appear with refer- ence to ‘phoba.’ The new Glossary will do much to fill the gap left by the seriously out-of-date old work, but it 1s to be fervently hoped that the com- puter base on which the new work is stored will in not-too-many years from now afford the possibility of producing a much more critical and accurate new edition of it. George C. Steyskal, Cooperating Scien- tist, Systematic Entomology Laboratory, Agricultural Research Service, U.S.D.A., % National Museum of Natural History, NHB- 168, Washington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 92(1), 1990, p. 176 Book REVIEW Cotton Insect Pests and Their Manage- ment. By G. A. Matthews, Longman Sci- entific and Technical, Essex, England, Copublished with John Wiley and Sons, New York. 1989. x + 199 pp. $49.95. Cloth. Improvement of production agriculture in the developing nations of the world is the major emphasis of many international re- search organizations as well as Third World governments. The development and imple- mentation of integrated, rather than solely chemical pest management is of major im- portance in many of those countries. Al- though research on integrated control is far from complete, many advances in the com- patible use of insecticides with other control mechanisms are being made. This book attempts to provide a concise, up-to-date discussion of tropical cotton in- sect management. It gives the reader the best information available on chemical con- trol methods and shows how these methods affect other control strategies and crop pro- duction practices. Research from Africa is emphasized, but management strategies from all cotton production areas are men- tioned where appropriate. The author pro- vides a needed, straightforward view of cot- ton pest management in areas of subsistence agricultural production. The text is in English and is well written, with the exception of a few minor gram- matical/typographic errors. The contents are presented in a logical sequence using nine chapters, with descriptions of the crop and insect pests given first, followed by control tactics and pest management discussions. Figures and tables are of good quality and provide visual illustrations in needed places. Color plates of insects and damage symp- toms are exceptionally clear but unfortu- nately are not available for all of the insects discussed. Inclusion of the additional plates would have been helpful for diagnosing in- sect pest problems. The number and types of appendices is especially pleasing. These appendices can be used to determine ap- propriate insect sampling methods, to con- vert metric units to English units for cali- brating sprayers and mixing chemicals, and to determine insect distributions and eco- nomic impacts. The Reference section is comprehensive and contains most of the more important citations relating to cotton insect management. In general I find this book to be a worthy contribution to the science of insect man- agement in cotton. It is an excellent book particularly for people with little or no for- mal training in insect pest management. It will become a valuable addition to libraries, especially in developing countries, where integrated pest management is in its infan- cy. Pest management specialists will rapidly adopt this book and it will provide the basis for development of pest control programs on cotton. Laurence D. Chandler, /nsect Biology and Population Management Research Labo- ratory, Agricultural Research Service, USDA, P.O. Box 748, Tifton, Georgia 31793. PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Cynipid Galls of the Eastern United States, by Lewis H. Weld Cynipid Galls of the Southwest, by Lewis H. Weld. RG EIBADCESON GY pic pallse sens eee wee ee RS Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman........ Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. A Short History of the Entomological Society of Washington, by Ashley B. Gurney Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. SSS ee ee eee Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. Le ySK alee st Se MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939. No.2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. ht - 10. pals Pl. ilies IS) fey NE ie a ee eh ee Be ee eee ee ee The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. 253 pp. 1949 A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952.00 A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Takahasi. 230 pp. 1969 Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. The North American Predaceous Midges of the Genus Pal/pomyia Meigen (Diptera: Cera- topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 fam), IMEI see SE ee a tO ge Sk Se SE ee ee ee ee Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982... A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. (OU. yeh LORE cat ea od OE SRE) SO RE AE Se Lee, ee An Identification Manual for the North American Genera of the Family Braconidae (Hy- menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 2.00 $15.00 (out of print) 15.00 15.00 15.00 15.00 11.00 12.00 10.00 11.00 18.00 5.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) SANDS, D. P. A., R. BROE, and W. J. M. M. LIEBREGTS.—Identity of Encarsia spp. (Hy- menoptera: Aphelinidae) introduced into Western Samoa for biological control of Pseu- daulacaspis pentagona (Targioni-Tozzetti) (Hemiptera: Diaspididade) SHEFFER, B. J. and M. L. WILLIAMS.—Descriptions, distribution, and host-plant records of eight first instars in the genus Toumeyella (Homoptera: Coccidae) SITES, R. W.—Morphological variations in the hemelytra of Cryphocricos hungerfordi Usinger (Heteroptera: Naucoridae) SPINELLI, G. R. and W. L. GROGAN, JR.—New species of predaceous midges of the tribe Ceratopogonini from Subantartic Argentina (Diptera: Ceratopogonidae) STOETZEL, M. B. and L. M. RUSSELL.—Ninth report on aphid-host relationships at the Los Angeles State and County Arboretum (Homoptera: Aphididae) WALTZ. R. D.—Baetis jesmondensis McDunnough, a New Junior Synonym of Baetis tricaudatus Dodds (Ephemeroptera: Baetidae) WHITFIELD, J. B.— Phylogenetic review of the Stiropius group of genera (Hymenoptera: Bra- conidae, Rogadinae) with description of a new neotropical genus BOOK REVIEWS CHANDLER, L. D.—Cotton Insect Pests and Their Management FUNK, V. A.—Novel Aspects of Insect-Plant Interactions JACOBSON, M.—Cyanide Compounds in Biology LACEY, L. A.—Combating Resistance Xenobiotics: Biological and Chemical Approaches .... STEYSKAL, G. C.—The Torre-Bueno Glossary of Entomology MISCELLANEOUS INSTRUCTIONS TO AUTHORS 135 44 111 VOLE. 92 APRIL 1990 NO. 2 (ISSN 0013-8797) PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS BAKER, G. T.—Morphology of the puparium of Lipoptena mazamae (Diptera: Hippoboscidae) 224 BOTTORFF, R. L., S. W. SZCZYTKO and A. W. KNIGHT.—Descriptions of a new spe- cies and three incompletely known species of western nearctic Jsoperla (Plecoptera: Per- ACEI AG) ener MT ec, Sy ee II cs ore often ticle recs ciols Fis sosuy aiecehe Wale @ alee cele Aves 286 BROWN, B. V.—Triphleba vitrinervis (Malloch), an unrecognized species of Crinophleba Borg- RTETE NP LELAsOMIGAG) ims. 5 yore pee ets Mca ore his see crap Pare oabe ele oiels Sle ey ale Bee ann de 271 DAVIDSON, R. L. and H. J. LEE, JR.—Distribution and habitat of Dyschirius campicola Lindroth (Coleoptera: Carabidae) with new state records for Ohio and Illinois, first records CASO MES NAISSISSIPDIN UV.CTA oe. era creer rome toke ai ue a: te eon miele ayers) tiie mio wie Sy Staneichal 205 DAVIS, D. R.—First record of a bagworm moth from Hawaii: description and introduction of Brachycyttarus griseus De Joannis (Lepidoptera: Psychidae) .....................---45- 259 DIETRICH, C. H. and S. H. MCKAMEY.—Three new idiocerine leafhoppers (Homoptera: Cicadellidae) from Guyana with notes on ant-mutualism and subsociality .............. 214 FREIDBERG, A. and W. N. MATHIS.—A new species of Craspedoxantha and a revised phy- IGPeuva Ome, Penusi (Diptera: DephnritiGae) meer & etary pete. le: as eke are vnikhans ws cota Siete es 325 GRISSELL, E. E. and M. E. SCHAUFF.—A synopsis of the seed-feeding genus Bephratelloides Penal eidotdedsBUnyTOMIGAE)) |p eel uence tort, o chegs mis Be Sachets steed ehiiets a aE aoe es 177 GROGAN, W. L., JR. and W. W. WIRTH.—A new species of the minute predaceous midge genus Nannohelea from Sri Lanka (Diptera: Ceratopogonidae) ...................2.... 347 HERATY, J. M. and K. N. BARBER.—Biology of Obeza floridana (Ashmead) and Pseudo- chalcura gibbosa (Provancher) (Hymenoptera: Eucharitidae) .......................4.. 248 HOFFMAN, K. N. and J. C. MORSE.—Descriptions of the females of three Polycentropus Species (buichopteras LolycemtropOgidae)) v2.22). ). sk scle sidayeles eed doin os wcities Ome eh el etee ers 274 JOHNSON, N. F.—Telenomus (Hymenoptera: Scelionidae) egg parasites of Erinnyis ello (Lep- He GYR Spal SAIN) ig bao eee aaa a eet on GO RR E AN OOO: aU BCEN Seen a CC ne oe hone Soe 306 LASALLE, J. and M. E. SCHAUFF.—New subfamily placement for some North American Bopiidae (Hymenoptera) Chaleidoidea)).. AED meal> l= <> Sr w= 5116S) > Os 30 Ss Figs. 30-35. Dasyhelea hutsoni. 30-33, female; 34-35, male: 30, antenna; 31, palpus; 32, spermatheca; 33, genital sclerotization; 34, parameres; 35, genitalia, parameres omitted. a spoonlike rather than bifurcate tip, and aedeagus which has the lateral sclerites much more slender and lacks the dorsal arcuate sclerite. Dasyhelea hutsoni Wirth, New SPECIES Female holotype. — Wing length 0.66 mm; breadth 0.31 mm; costal ratio 0.52. Head pale brown, antennae darker brown. Anten- na (Fig. 30) with lengths of flagellar seg- ments in proportion of 27-22-23-25-25-25- 26-27-36-34-32-34-56, antennal ratio 0.96; segments sculptured proximally, last seg- ment elongate with tapering tip. Palpus (Fig. 31) short and relatively stout, lengths of seg- ments in proportion of 19-30-15-12. Tho- rax pale brown, scutellum and sides of mesonotum yellowish; legs uniformly stra- mineous. Hind tarsal ratio 3.0. Wing smoky grayish, veins slightly darker; macrotrichia moderately numerous over entire wing, long, coarse and dark brown; first radial cell ves- tigial, second well formed, square-ended. Halter brownish. Abdomen brownish; gen- ital sclerotization as in Fig. 33; spermatheca (Fig. 32) one, well pigmented; oval with short oblique neck; measuring 0.054 by 0.026 mm and neck 0.009 mm long. Male allotype.—Wing length 0.78 mm; costal ratio 0.48. Similar to the female, with the usual sexual differences. Antenna with sparse plume of long brown verticils; seg- ments 7-12 fused; flagellar segments with lengths in proportion of 30-25-23-23-23- 23-23-23-23-50-50-50-62, antennal ratio (12-15/3-11) 0.98. Palpus with lengths of segments in proportion of 15-30-16-17. Hind tarsal ratio 2.4. Genitalia (Fig. 35): Ninth sternum with evenly rounded pos- terior convexity abutting base of aedeagus; ninth tergum short and tapering, about as long as basal breadth, apicolateral processes a pair of angular lobes ending 1n a somewhat beadlike setigerous process, the caudal mar- gin between the lobes deeply concaved. Ba- sistyle somewhat swollen proximally, with- VOLUME 92, NUMBER 2 241 Gt ee eseeeeee 38 40 Figs. 36-41. Dasyhelea tamsi female: 36, antenna; 37, palpus; 38, wing; 39, genital sclerotization; 40, sper- matheca; 41, mesonotal pattern. out special lobe or armature; dististyle about as long as basistyle, base only moderately enlarged, tapering and curved distally to moderately stout, pointed tip. Aedeagus of diagnostic shape; deeply pigmented; main body slightly broader than long, anterior margin slightly concave, with short, slender anterolateral arms; posteriorly a pair of short, stout processes tapering to blunt tips slightly turned ventrolaterad. Parameres (Fig. 34) with stout, slightly asymmetrical basal apodemes; posterior median portion swollen on proximal half with a short, strongly pigmented, slender, pointed pro- cess abruptly bent back ventrolaterad from near midlength; distal half strongly nar- rowed to a pointed tip directed ventrola- terad to side opposite the proximal process. Types.—ALDABRA ATOLL: holotype 2, allotype 6, 4 6 and 1 2 paratypes; South Island, Takamaka Pool, 1—17.11.1968, (Co- gan & Hutson), at light; 1 2 paratype, Taka- maka, in mangroves, otherwise same data. Ile Michel, 16.11.1968, (Cogan & Hutson), 1 2 paratype. (Holotype and allotype in BMNH: paratypes in BMNH, PARIS, USNM.) Distribution. —Aldabra. This species is dedicated to A. M. Hutson of the Department of Entomology, British Museum (Nat. Hist.), London, in appreci- ation of his interest and assistance in my study of the Aldabra Ceratopogonidae. Dasyhelea hutsoni closely resembles D. labourdonnaisi Clastrier (1959) from Ile Ré- union in general appearance and in the structure of the male aedeagus, but the re- lated species differs in its larger size (male wing length about 1.2 mm), straight caudal margins of the ninth sternum and tergum, the apicolateral processes of the tergum long and slender; the aedeagus with longer an- terolateral arms; the dististyle tapering to more slender tip; the posterior portion of the paramere shorter and evenly tapered to the tip and lacking the retrorse proximal process, and the basal apodemes very slen- der and asymmetrical. Dasyhelea tamsi Wirth & Messersmith, 1977. Dasyhelea tamsi Wirth & Messersmith, 1977: 305 (2; Seychelles; figs.); Clastrier, 1983: 36 (2 redescribed from type series). 242 Diagnostic characters.—A small _polli- nose pale gray species; wing whitish with slightly darkened stigma. Mesonotum (Fig. 41) pale brown with three broad dark brownish gray vittae, humeri extensively pale yellowish gray; on slide-mounted spec- imens five narrow opaque lines of internal pigmentation form borders to the vittae (as noted by Clastrier 1983). Scutellum yellow- ish. Legs whitish, knee spots blackish. An- tenna (Fig. 36) short, with segments in a continuous series, moniliform proximally to slightly elongate on distal segments, last segment without terminal stylet; surface of segments conspicuously reticulated. Palpus (Fig. 37) short and stout. Wing (Fig. 38) milky whitish, radial cells forming a dark stigma; costa ratio 0.49; veins forming ra- dial cells greatly strengthened, the first ra- dial cell obsolete, second short with small lumen; macrotrichia long and stout, very sparse, forming lines along veins. Halter grayish infuscated. Abdomen grayish brown, terga with pairs of small round hyaline non- pigmented spots. Genital sclerotization (Fig. 39) with slightly infuscated, quadrate, me- dian lobe and a slender pair of oblique lat- eral arms. Spermatheca (Fig. 40) one, ovoid, tapering to short stout neck; heavily scler- otized; small, measuring 0.060 by 0.045 mm. Male unknown. Distribution. — Aldabra, Seychelles. Material examined.—ALDABRA ATOLL: South Island, Dune Jean-Louis, 13- 20.11.1968, at light, 2 2; Anse Cedre, 17- 19.11.1968, 1 @; Takamaka Grove, 1- 17.11.1968, 1 @ (all Cogan & Hutson) (BMNH, PARIS, USNM). West Island (Ile Picard), Settlement, 12—22.111.1986 (D. Adamski), 23 2 (USNM). SUBFAMILY CERATOPOGONINAE Tribe Culicoidini Culicoides adamskii Wirth, NEw SPECIES Female holotype.— Wing length 1.16 mm; breadth 0.55 mm; costal ratio 0.58. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Head: Brown, antennal flagellum yellow. Eyes (Fig. 45) bare, narrowly separated, by width of 2 facet. Antenna (Fig. 42) with lengths of flagellar segments in proportion of 35-24-26-26-27-28-27-30-50-50-50-57- 75, antennal ratio 1.25; sensilla coeloconica present on segments 3, 5, 7,9, 11-15. Palpus (Fig. 43) dark brown; lengths of segments ment moderately swollen distally, with a moderately large, round, moderately deep sensory pit; palpal ratio 2.02. Proboscis moderately long; P/H Ratio 0.79. Mandible with 16 teeth. Thorax: Brown; mesonotum subshining dark brown with grayish pollinosity; three dark brown vittae, a narrow median vitta on anterior half, a pair of broad sublateral bands from humeral pits to above wing bas- es; a pair of small dark brown spots 1n pre- scutellar depression. Legs (Fig. 47) dark brown, knee spots blackish; fore and mid femora with subapical, and all tibiae with sub-basal, narrow pale rings, tarsi pale; hind tibial comb with five spines, second from spur longest. Wing (Fig. 14, 44): Strongly infuscated, veins dark brown. Pattern as figured; two prominent but small pale spots on anterior margin, first over r-m crossvein continued cephalad to costal margin, second just past tip of costa and extending caudad half the width of cell R5; faint rounded pale spots at wing margin in each of cells R5, M1, M2, and M3+4, the latter filling distal half of cell; small pale spot at wing base just distad of basal arculus; a faint pale spot in cell M2 just behind medial fork and another just in front of mediocubital fork; anal cell with a double pale spot distally extending broadly to posterior wing margin. Macrotrichia long and coarse, abundant, covering entire wing; radial cells well-formed, with distinct lu- mens. Halter dark brown. Abdomen: Brown. Spermathecae (Fig. 49) two plus sclerotized ring and vestigial third; subequal, spherical with long slender necks; each measuring 0.087 by 0.087 mm plus neck 0.032 mm long. VOLUME 92, NUMBER 2 243 EE EE AS CE EES DAD GED DD EE FD) | (- 44 I Figs. 42-49. CD l 42 43 Sa oe) 46 Culicoides adamskii, 42-45, 47, 49, female; 46, 48, male: 42, antenna; 43, palpus; 44, wing; 45, eye separation; 46, genitalia, parameres omitted; 47, femora and tibiae of (left to mght) fore, mid and hind legs; 48, parameres; 49, spermathecae. Male allotype.—Similar to female with usual sexual differences. Genitalia (Fig. 46): Ninth sternum with broad, shallow, cau- domedian excavation, ventral membrane not spiculate; ninth tergum about as long as basal breadth, tapering to small, pointed, moderately separate, apicolateral processes, the caudal margin between them straight, with only a hint ofa median notch. Basistyle about twice as long as broad, with unusual modification of mesal margin consisting of a broad, distally pointed, platelike basal lobe extending from ventral root to half the length of basistyle; ventral root long and slender, slightly curved; dorsal root curved, half as long, with blunt tip; dististyle somewhat swollen proximally, gradually tapering dis- tally to slender, incurved tip. Aedeagus with basal arms slender and evenly curved, form- ing a high slender arch to % of total length; distal process small and slender, with sim- ple tip. Parameres (Fig. 48) each with strong basal knob bearing a short anterior process, constricted a short distance just past knob; mid portion nearly straight, slightly bowed outwardly, gradually tapering distad, the short distal portion abruptly bent ventro- laterad and slightly expanded in a rounded, flattened, spoonlike tip. Distribution. — Aldabra. Types.—ALDABRA ATOLL: Holotype 2, West Island (Ile Picard), Settlement, 1 2- 22.111.1986, (D. Adamski), in UV light trap (USNM). Allotype ¢, South Island, Dune Jean-Louis, 13-20.11.1968 (Cogan & Hut- son) (BMNH). Paratypes, 7 4, 36 2, as fol- 244 lows: Same data as holotype, | 2; same data as allotype, 4 4, 4 2 (BMNH, PARIS, USNM). South Island, Dune D’Messe, 21.11.1968, (Cogan & Hutson), 21 2; Taka- maka, 1—17.11.1968 (Cogan & Hutson), | 4, 7 2. Middle Island, near East Channel, 6- 7.11.1968, (Cogan & Hutson), | 2. West Is- land, near Settlement, 21—31.111.1968, (Co- gan & Hutson), 1 é, 1 2 (BMNH, PARIS, USNM). This species 1s dedicated to David Adam- ski of Mississippi State University, who sent me a small but important collection of Al- dabra Ceratopogonidae for study. Culicoides adamskii is closely related to C. eriodendroni Carter, Ingram, & Macfie (1921) and C. nigripennis Carter, Ingram & Macfie (1920), the taxonomy of which is still in some confusion, with a number of closely related mainland Subsaharan species in the complex remaining to be described (Cornet, Glick, Meiswinkel, Phelps, in litt.). C. adamskii resembles C. nigripennis in an- tennal sensillar pattern 3,5,7,9,1 1-15, pal- pal proportions and shape of the sensory pit, shape and size of the spermathecae, leg color pattern and dark halter, but differs markedly in wing pattern which in C. ni- gripennis 1s restricted to two small anterior pale spots, and the longer costa (costal ratio 0.64) and presence of only four tibial spines in C. nigripennis. Culicoides eriodendroni resembles C. adamskii in wing pattern, leg pattern, and dark halter, but has antennal sensillar pattern 3,11—15, four tibial spines, and unequal ovoid spermathecae without necks. The male genitalia of C. nigripennis (as figured by Boorman & Dipeolu 1979) re- semble those of C. adamskii but have the dististyle like that of C. eriodendroni, ba- sistyle like that of C. /amborni Ingram & Macfie (1925), and the parameres are more slender than in either species and lack the spoonlike swelling at the tip. The male gen- italia of C. eriodendroni (as figured by In- gram & Macfie 1921) differ in the large basal swelling on the dististyle, lack of the basal PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lobe on the mesal margin of the basistyle, and shorter, stouter parameres. Culicoides lamborni, known from the male only, has a wing pattern like that of C. nigripennis, and the genitalia have the dististyle and aede- agus like those of C. adamskii, but the par- ameres have a slender apex and the basistyle lacks the basal lobe. The described species of the C. nigripen- nis Group have been reared from rot holes in various trees, and it is reasonable to ex- pect to find the immature stages of C. adam- skii in similar habitats. Tribe Ceratopogonini Metacanthohelea cogani Wirth & Grogan, 1988. Metacanthohelea cogani Wirth & Grogan, 1988: 66 (6, 2; Aldabra; figs.). Diagnostic characters. — A small, dull dark brown midge; wing length 0.90 mm. Eyes broadly separated, bare. Female antenna with distal five segments elongated: distal three segments of male antenna elongated; male antennal segments 3-10 fused, with sparse plume. Palpus 5-segmented: third segment short, swollen, with well-defined sensory pit. Legs moderately stout; hind fe- mur swollen, bearing 14 large spines on dis- tal half; fourth tarsomeres short but sub- cylindrical, bearing single apical sinuate hyaline sensillum; female claws small and equal with basal inner teeth. Wing milky, with two narrow radial cells, second 1.5 times length of first; costal ratio 0.67; vein M2 narrowly interrupted at base. Two ovoid spermathecae with oblique long slender necks. Aedeagus triangular, broad and short; parameres fused basally with long slender distal portions recurved at their tips. Distribution. —Aldabra, Kenya. Types.—ALDABRA ATOLL: Holotype 2, allotype 4, South Island, Takamaka, |- 17.11.1968, (Cogan & Hutson) (BMNH). Paratypes, 2 6, 2 9, same data; 1 6, South Island, Dune Jean-Louis, at light, 13- 20.11.1968 (Cogan & Hutson); | 4, West VOLUME 92, NUMBER 2 Island, near settlement, at light, 21- 31.11.1968 (Cogan & Hutson). One @ para- type, KENYA, Marsabit Nature Reserve, 4200 ft, 8.xii.1969 (Irwin & Ross) (Califor- nia Acad. Sci.). Material examined.—ALDABRA AT- OLL: West Island (Ile Picard); Settlement, 12-22.111.1986, (D. Adamski), 1 ¢(USNM). The occurrence of this highly modified genus and species in only two widely dis- junct localities, —The Indian Ocean atoll of Aldabra, and at 1250 m elevation at the Marsabit Nature Reserve in Kenya, has in- teresting biogeographical implications which at present are not readily explained, except to point out the close relationship of the Aldabra fauna to that of the African main- land. Stilobezzia spirogyrae Carter, Ingram & Macfie, 1921. Stilobezzia spirogyrae Carter, Ingram & Macfie, 1921: 325 (all stages; Ghana; figs.). Distribution.— Aldabra, Gambia, Ghana, Sao Tome, South Africa. Material examined.—ALDABRA ATOLL: Ile Michel, 3.11.1968, 1 6. South Island, Cing Cases, 3-16,23-29.1.1968, 4 9, Flamingo Pool, 21-22.1.1968, 10 92; Frigate Pool, 20,1.1968, 1 6; Takamaka, Takamaka Grove, Takamaka Pool, 1-17.11.1968, 76 4, 23 2 (all Cogan & Hutson) (BMNH, PARIS, USNM). Tribe Sphaeromiini Homohelea stuckenbergi (de Meillon), 1961. Sphaeromias stuckenbergi de Meillon, 1961: 51 (9; Madagascar; figs.). Homohelea stuckenbergi (de Meillon); Wirth et al. 1980: 170 (combination); de Meil- lon & Wirth, 1981: 543 (in key). Distribution. — Aldabra, Madagascar. Material examined.—ALDABRA ATOLL: South Island, Cing Cases, 23-29.1.1968, 2 2; Frigate Pool, 20.1.1968, 1 2; Takamaka, 1-17.11.1968, 2 2 (all Cogan & Hutson) (BMNH, PARIS, USNM). This species is distinguished from its Sub- saharan congeners by its large size (wing 245 length more than 3.0 mm), ornamented mesonotum and abdomen, fore femur with 9 spines, mid and hind femora with 4-6 spines, and only one talon of the fore claws barbed. Tribe Palpomyiini Bezzia africana Ingram & Macfie, 1923 Bezzia africana Ingram & Macfie, 1923: 71 (2; South Africa; figs.); de Meillon, 1943: 107 (4; Transvaal; figs.); Haeselbarth, 1975: 357 (redescribed; figs.; distribution; synonymy) Distribution.—Aldabra, Cameroun, Madagascar, South Africa, Zimbabwe. Material examined.—ALDABRA ATOLL: South Island, Takamaka Pool, 1-17.ii.1968, (Cogan & Hutson), | 2. ACKNOWLEDGMENTS Iam especially grateful to Brian H. Cogan and A. M. Hutson of the British Museum (Natural History) in London for their kind- ness in making their collections available for study, and to Richard Lane for the loan of types and for information concerning types in that museum. I wish to thank Dr. John Boorman for his kindness in exam- ining syntypes of Thysanognathus mono- stictus Ingram & Macfie in that museum and sending photographs. I also wish to thank David Adamski of Mississippi State Uni- versity, University, Mississippi, for his kindness in furnishing his ceratopogonid collections from Aldabra for study. The as- sistance of Molly A. Griffin in making the illustrations is gratefully acknowledged. LITERATURE CITED Abbott, W. L. 1893. Notes on the natural history of Aldabra, Assumption and Glorioso Islands, In- dian Ocean. Proc. U.S. Nat. Mus. 16: 759-764. Boorman, J. and O. O. Dipeolu. 1979. A taxonomic study of adult Nigerian Culicoides Latreille (Dip- tera: Ceratopogonidae) species. Occas. Publ. Ento- mol. Soc. Nigeria 22: 1-121. Carter, H. F., A. Ingram, and J. W. S. Macfie. 1920. Observations on the ceratopogonine midges of the 246 Gold Coast with descriptions of new species. Ann. Trop. Med. Parasitol. 14: 211-274, 2 plates (Part II); 14: 309-331 (Part III). Carter, H. F., A. Ingram, and J. W. S. Macfie. 1921. Observations on the ceratopogonine midges of the Gold Coast with descriptions of new species. Part IV. Ann. Trop. Med. Parasitol. 15: 177-212. Clastrier, J. 1956. Notes sur les Cératopogonides. I— Quatre Forcipomyia d Algerie et de Tunisie. Arch. Inst. Pasteur Algérie 34: 496-512. 1959. Notes sur les Cératopogonides VIII.— Cératopogonidés de IIe de la Reunion. Arch. Inst. Pasteur Algérie 37: 412-446. 1983. Cératopogonidae des Iles Seychelles (Diptera, Nematocera). Mém. Mus. Natl. Hist. Nat. N.S., Sér. A, Zool. 126: 1-83. Clastrier, J. and W. W. Wirth. 1961. Notes sur les Cératopogonidés XIV. Cératopogonides de la Re- gion éthiopienne (2). Arch. Inst. Pasteur Algérie 39: 302-337. Cogan, B. H., A. M. Hutson, and J. C. Shaffer. 1971. Preliminary observations on the affinities and composition of the insect fauna of Aldabra. Phil. Trans. R. Soc. London B 60: 315-325. Dessart, P. 1961. Contribution a etude des Cera- topogonidae (Diptera) (II) Revision des Forcipo- myia congolaise decrits par le Dr Goetghebuer. Bull. Ann. Soc. R. Entomol. Belgique 97: 315- 376. 1963. Contribution a l’étude des Ceratopo- gonidae (Diptera) (VII) Tableaux dichotomiques illustrés pour la détermination des Forcipomyia Africains. Mem. Inst. R. Sci. Nat. Belgique Deux. Sér. 72: 1-151, 16 plates. Downes, J. A. and W. W. Wirth. 1981. Ceratopo- gonidae, pp. 393-421. Jn Manual of Nearctic Diptera. Vol. 1. McAlpine, J. F., et al., eds. Res. Br. Agric. Canada Monogr. 27: 1-674. Frith, D. W. 1979. A twelve month study of insect abundance and composition at various localities on Aldabra Atoll. Phil. Trans. R. Soc. London B 286: 119-126. Goetghebuer, M. 1933. Ceratopogonidae et Chiro- nomidae du Congo Belge. Revue Zool. Bot. Af- ricaines 24: 129-151. Haeselbarth, E. 1975. Zur Kenntnis von Bezzia af- ricana und verwandten Gnitzen-Arten aus der Ae- thiopischen Region (Diptera: Ceratopogonidae). Entomol. Germanica |: 352-370. Ingram, A. and J. W. S. Macfie. 1921. West African Ceratopogoninae. Ann. Trop. Med. Parasitol. 15: 313-374, | plate. . 1923. Notes on some African Ceratopogon- inae. Bull. Entomol. Res. 14: 41-74. 1924. Notes on some African Ceratopogon- inae—Species of the genus Forcipomyia. Ann. Trop. Med. Parasitol. 18: 533-594. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1925. New Ceratopogoninae from Nyasaland (Dipt.). Bull. Entomol. Res. 15: 283-288. Kieffer, J. J. 1911. No. XV.—Diptera, Chironomidae der Seychellen-Inseln, aus der Sammlung von Mr H. Scott. Trans. Linn. Soc. London 2nd Ser. Zool. 14: 331-366, | plate. Legrand, H. 1965. Lépidoptéres des Iles Seychelles de d’Aldabra. Mem. Mus. Natl. Hist. Nat. Nouv. Ser. Ser. A, Zool. 37: 1-210. Macfie, J. W. S. 1938. Notes on Ceratopogonidae (Diptera). Proc. R. Entomol. Soc. London Ser. B Tax. 7: 157-166. Meigen, J. W. 1818. Systematische Beschreibung der bekannten europaischen zweiflugeligen Insekten. Vol. 1, 333 pp., 11 plates. Aachen. Meillon, B. de. 1939. Notes on Ceratopogonidae from Southern Africa II. J. Entomol. Soc. S. Africa 2: 7-17. . 1943. Simulidae and Ceratopogonidae (Dipt. Nematocera) from the Colony of Mocambique. Estacao Anti-Malarica de Lourenco Marques, 31 pp. 1961. The Madagascan Ceratopogonidae. Revta. Entomol. Mocgambique 4: 37-64. Meillon, B. de and W. W. Wirth. 1981. Subsaharan Ceratopogonidae (Diptera) VI. New species and records of South African biting midges collected by A. L. Dyce. Ann. Natal Mus. 24: 525-561. 1983. Subsaharan Ceratopogonidae (Dip- tera) IX. New species and records from southern Africa. Ann. Natal Mus. 25: 347-381. Peake, J. F. 1971. The evolution of terrestrial faunas in the western Indian Ocean. Phil. Trans. R. Soc. London B 260: 581-610. Scott, H. 1933. General conclusions regarding the insect fauna of the Seychelles and adjacent islands. Trans. Linn. Soc. London 2 Ser. Zool. 19: 307- 391. Stoddart, D. R. and T. S. Westoll, Eds. 1979. The terrestrial ecology of Aldabra. Phil. Trans. R. Soc. London B. 286-1-263. Westoll, T. S. and D. R. Stoddart, Eds. 1971. A dis- cussion of the results of the Royal Society Expe- dition to Aldabra 1967-68. Phil. Trans. R. Soc. London B 260: 1-654. Wirth, W. W. 1952. The Heleidae of California. Univ. California Pubs. Entomol. 9: 95-266. 1956. New species and records of biting midges ectoparasitic on insects (Diptera, Hele- idae). Ann. Entomol. Soc. America 49: 356-364. . 1972. The Neotropical Forcipomyia (Micro- helea) species related to the caterpillar parasite F. fuliginosa (Diptera: Ceratopogonidae). Ann. Ento- mol. Soc. America 65: 564-577. Wirth, W. W. and W. L. Grogan, Jr. 1988. The pre- daceous midges of the world (Diptera: Ceratopo- gonidae, Ceratopogonini). Flora and Fauna Hand- book 4: 1-160. E. J. Brill, Leiden. VOLUME 92, NUMBER 2 Wirth, W. W., B. de Meillon, and E. Haeselbarth. 1980. 10. Family Ceratopogonidae, pp. 150-174. In Crosskey, R. W., ed. Catalogue of the Diptera of the Afrotropical Region, 1437 pp. Brit. Mus. (Nat. Hist.) London. Wirth, W. W. and D. H. Messersmith. 1977. Notes on the biting midges of the Seychelles (Diptera: Ceratopogonidae). Proc. Entomol. Soc. Washing- ton 79: 293-309. Wirth, W. W. and N. C. Ratanaworabhan. 1976. A 247 new species of parasitic midge (Forcipomyia (Pte- robosca)) from Aldabra, with description ofits pre- sumed larva and pupa and systematic notes on the subgenera of Forcipomyia (Ceratopogonidae). Syst. Entomol. |: 341-345. Wirth, W. W., N. C. Ratanaworabhan, and F. S. Blan- ton. 1974. Synopsis of the genera of Ceratopo- gonidae (Diptera). Ann. Parasitol. (Paris) 49: 595—- 613. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 248-258 BIOLOGY OF OBEZA FLORIDANA (ASHMEAD) AND PSEUDOCHALCURA GIBBOSA (PROVANCHER) (HYMENOPTERA: EUCHARITIDAE) JOHN M. HERATY AND KEVIN N. BARBER (JMH) Department of Entomology, Texas A&M University, College Station, Texas, 77843; (KNB) Forestry Canada, Forest Pest Management Institute, P.O. Box 490, Sault Ste. Marie, Ontario P6A 5M7, Canada. Abstract. —Brief descriptions of life history are given for Obeza floridana (Ashmead) and Pseudochalcura gibbosa (Provancher) from Florida and northern Ontario, respec- tively. Obeza floridana oviposits into fruits of Cyanococcus simulatus Small and P. gibbosa into overwintering floral buds of Ledum groenlandicum Oeder. and leaf buds of Arbutus menziesi Pursh. Descriptions of eggs and first-instar larvae of each species are provided and compared to other Eucharitidae. Pseudochalcura gibbosa was reared from Camponotus herculeanus (L.) and details of complete life history and descriptions of all larval stages are supplied. Comparisons are made with the Old World genus Sti/bula and their taxo- nomic interrelationships discussed. Key Words: life history, larval stages Obeza floridana (Ashmead) and Pseudo- chalcura gibbosa (Provancher) belong to the family Eucharitidae (Hymenoptera: Chal- cidoidea). The genus Obeza Heraty was re- cently erected to hold the New World species that were previously considered to be mem- bers of the widespread Old World genus Stilbula Spinola (Heraty 1985). The genus Stilbula, and the genera Obeza, Lophyro- cera Cameron (considered here to include Tetramelia Kirby) and Pseudochalcura Ashmead comprise a monophyletic group (Heraty 1985, 1986). Systematic relationships within this clade have already been discussed (Heraty 1985, 1986) and are summarized as follows. Obeza has been regarded as the sister group to Lo- phyrocera + Pseudochalcura while Stilbula was considered the sister taxon to these three New World genera (Heraty 1985). Lophyro- cera was again recognized as a sister group to Pseudochalcura by Heraty (1986) al- though this was questioned at the time. Ex- act placement of the genus Pseudochalcura is problematic since it is the only member lacking lateral propodeal processes and short, bifurcating spines issuing from the frenum. Close taxonomic relationship be- tween Pseudochalcura and Obeza or Stil- bula is supported by synapomorphies which include similar structure of the pronotal- prepectal area and musculature of the meso- soma (Heraty 1989). A recent discovery of two undescribed species of Pseudochalcura from the Orient suggest a closer relationship to the Old World genus Sti/bula and to the recently described genus Sti/buloidea Bou- éek. Exact systematic relationships of these taxa have yet to be assessed but the mon- ophyly of the group is certain (Heraty 1986). As with other members of the Euchariti- dae, the above-mentioned genera are spe- cialized parasites of mature larvae and pu- pae of ants. Adult females deposit their eggs VOLUME 92, NUMBER 2 away from the host into plant tissue. The active first-instar larva, termed a planidi- um, must make its way back to the ant nest, usually phoretically on an adult ant, where it can attack the brood (Clausen 1940a, b, 1941). Morphology of the immature stages, and in particular, the well-sclerotized pla- nidium, is highly conservative and can be useful in positing relationships at the higher taxonomic levels (Heraty and Darling 1984). Such an approach can serve to test, supple- ment, and refine hypotheses derived from studies of adult morphology. Lophyrocera (including Tetramelia) in- cludes six described Neotropical species and L. apicalis Ashmead in the western United States (Heraty 1985). Nothing is known of the biology or immature stages of any of these seven species. The genus Obeza is widespread in South America with two species represented in North America: Obeza_ septentrionalis (Brues) in Arizona and New Mexico and O. floridana in Florida and Georgia (Heraty 1985). The biology and immature stages of Obeza have remained completely unknown until this time. Pseudochalcura is also widely distributed in the Neotropics (ten species) with three other species occurring in the Nearctic (Her- aty 1986). Pseudochalcura gibbosa is wide- spread in North America and has a typical Boreal distribution to the north and occurs throughout the Rocky Mountains in the west (Heraty 1986). Adults of P. gibbosa have been recorded as ovipositing into flower buds of Gossypium (Malvaceae) in Arizona (Pierce and Morrill 1914) and Arbutus (Er- icaceae) in California, along with collecting associations made with various other plants (Heraty 1986). Pseudochalcura gibbosa has been reared from Camponotus novaebora- censis (Fitch) in Michigan (Wheeler 1907), Camponotus laevigatus (F. Smith) and Camponotus sp. possibly vicinus Mayr in California (Heraty 1986). The only descrip- tion of immature stages of P. gibbosa was by Wheeler (1907) for the female pupa and 249 position of two pupae in a cocoon of C. novaeboracensis. In contrast to the paucity of information on the biology of the New World genera discussed here, there are detailed life history documentations and descriptions of im- mature stages for Stilbula cyniformis Ros- si (Parker 1932, 1937, Parker and Thomp- son 1925), Stilbula manipurensis (Clausen) (Clausen 1928, 1940a, b), and Stilbula ten- uicornis (Ashmead) (Clausen 1923, 1940b, 1941). Some peculiar features in the life his- tories of Sti/bula species are useful in dis- tinguishing this group from other Euchari- tidae and for comparison to the two species in this paper. Females deposit their large egg masses of more than one thousand eggs in a single oviposition, and in S. fenuicornis and S. cyniformis, the eggs have the ability to overwinter (Clausen 1928, 1940b, Parker 1937). As in other eucharitids, planidia are mobile and attach themselves externally to mature ant larvae. The first-instar larvae are unique in their ability to complete devel- opment on the mature larva within the co- coon, and in S. fenuicornis, are cast off with the host’s larval exuvium. The second-in- star larvae then relocates to the mid-section of the pupa to resume feeding (Clausen 1923, Parker 1928). Finally, the third-instar larva has a distinctive morphology with respect to other known genera of Eucharitidae, and more than one eucharitid can develop on a single host pupa (Clausen 1923, 1940a, Par- ker 1932). The recorded ant hosts include Camponotus herculeanus japonicus Mayr and Camponotus herculeanus obscuripes Mayr for S. tenuicornis (Clausen 1923, 1941), Camponotus sp. for S. manipurensis (Clausen 1928), and Camponotus aethiops (Latr.) for S. cyniformis (Parker 1932, 1937). The discovery of populations of O. flor- idana and P. gibbosa allowed us to gather information on the oviposition habits and life histories of these species. It also pro- vides an opportunity to make comparisons between the New World genera and the pro- 250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON posed outgroup genus, Szi/bu/a, and to re- flect upon the phylogenetic relationships of these taxa. Obeza floridana (Ashmead) (Figs. 2, 4, 8) Location and habitat.— The collection site was located just outside of the Apalachicola National Forest, 3.0 kilometres south of Sopchoppy, Wakulla Co., Florida. The host plant was located within a narrow windrow of Pinus sp. that paralleled the highway and bordered a large tree-less bog. Within the windrow, the sandy soil was covered by a thick mat of pine litter and scrub vegetation that consisted of sporadic clumps of Cyano- coccus simulatus Small. (Ericaceae), Dia- spyros sp. (Sapotaceae), Cyrilla racemiflora L. (Anacardiaceae) and Sabal glabra (Mill.) Sarg. (Palmae). General observations and collections. — Collections of adult O. floridana, Cyano- coccus fruits, and ants were made on two separate occasions in 1987—25-26 May and 4-5 June. Adult O. floridana were localized in their distribution and although some males were collected in a broader area, most males and all of the females except one were collected from one isolated C. simulatus bush. All but one of the females were devoid of eggs. Most adults were collected 4-5 June while no females and only two males were collected 25-26 May after a similar collect- ing effort. No copulating pairs were ob- served. Cyanococcus simulatus fruits were col- lected and returned to the laboratory. Egg masses were maintained at 75% RH over a saturated salt solution, either within intact berries or isolated in cotton stoppered glass vials, fora one month period. Although they remained viable over this period, no hatch of mature planidia occurred. Eggs retained for a longer period succumbed to desicca- tion and/or fungus. Strays of Camponotus abdominalis flor- idanus (Buckley) were found but no asso- ciation could be made. Therefore, no ad- ditional immature stages of O. floridana became available for study. Obeza floridana has been reared from the pupae of C. ab- dominalis floridanus near Gainesville, Flor- ida (Lloyd Davis, Insects Affecting Man and Animals Research Laboratory, Gainesville, FL, personal communication). Life history.— Adult females oviposit into developing fruits of C. simulatus in May and June. Adults were observed hovering around the edges of the Cyanococcus or set- tled on the fruits themselves. The single gravid female readily oviposited into green berries and later dissection showed the ova- ries to contain about 3500 fully developed eggs. Egg masses in the fruits consist of small clusters of 25-180 eggs (mean = 100.9, SD = 47.3,n = 15) that are deposited in pockets within the epidermal layer of the fruit (Fig. 2). They can be seen through the skin of the fruit and are associated with a minute ovi- position scar. A sample of 20 berries yielded 15 egg masses with 1-5 egg masses per fruit. The eggs were found in varying states of development within a single berry from pure white (recently deposited) to a darkened col- oration owing to the mature planidium in- side. These observations indicate multiple Ovipositions in individual fruits by different females. No information is available on how pla- nidia escape the fruit or enter the host ant nest. Adults of Camponotus femoratus Fabr. are known to collect independently the pulp and seeds of berries from a number of plants (Davidson 1988). The berries may be direct attractants to the host with planidia col- lected along with the fruit pulp. Deposition of eggs into a perishable fruit, as early as June, indicate that planidia complete de- velopment within the season, even though egg hatch could not be stimulated in the laboratory. Description of the immatures. — Egg (Fig. 4): Undeveloped eggs white with a smooth chorion. Length of egg body about 0.13 mm (SD = 0.01, n = 10); of caudal stalk 0.15 mm (SD = 0.02). Mature eggs with well- VOLUME 92, NUMBER 2 sclerotized first-instar larva occupying al- most entire egg body; head oriented toward stalk and small yellowish yolk mass at- tached to abdominal apex (Fig. 4). Except for the more rounded appearance of the egg body, the egg is similar to those of other Eucharitidae as described in Heraty and Darling (1984). First instar (as dissected from egg; Figs. 8, 10, 12): As described for other Eucharit- inae (Heraty and Darling 1984) but distin- guished as follows. Length 0.14 mm (SD = 0.01, n = 10); maximum width 0.06 mm, widest medially and circular in cross-sec- tion. Cranium with labial plates present, in- cluding hatchet-shaped posterior labial plate; two pairs of dorsal sensilla; large, paired, weakly sclerotized cranial processes arising dorsally from an unsclerotized region. Ter- gite I + II separated ventrally, fused dor- sally; tergopleural line absent. Tergite I + II with two dorsal and one ventral pair of setae; lacking one pair of ventral setae along ventral margin of tergite III; ventral mar- gins of tergites V-VI extended posteriorly as long narrow processes; posterior-ventral margins of tergites V—XI strongly scalloped. Additional information.—A second, un- described species of Obeza was collected on 21 July 1987, in a dry cactus/scrub wood- land habitat, 39.1 kilometres southeast of Teotitlan del Camino, Oaxaca, Mexico. Adults were observed ovipositing into the green berries of Tragia volubilis L. (Eu- phorbiaceae) which are similar in size and texture to the fruits of C. simulatus. The manner of egg deposition, relative size of the egg mass, and morphology of the first- instar larva were identical to those of O. floridana. Pseudochalcura gibbosa (Provancher) (Figs. 1, 3, 5-7, 9-15) Location and habitat.—The collection site was located northeast of Sault Ste. Marie, Ontario approximately 18 kilometres northeast of Searchmont, at about mile 15 on Whitman Dam Road. The habitat is a 251 disturbed, cutover area strewn with consid- erable fallen dead wood, slash, and stumps. The ground cover is predominated by Vac- cinium angustifolium Ait., V. myrtilloides Michx., and Ledum groenlandicum Oeder. Other plants include Polytrichum commune Hedw. (Polytrichaceae), Sphagnum angus- tifolilum (Russ.) C. Jens. (Sphagnaceae), Gaultheria hispidula (L.) Muhl., Epigaea re- pens L. (Ericaceae), Cornus canadensis L. (Cornaceae), Clintonia borealis (Ait.) Ras. (Convallariaceae), Rubus sp. (Rosaceae), and Ribes glandulosum Grauer. (Grossulari- aceae) with scattered individuals of Pinus strobus L., Abies balsamea (L.) Mill., Picea mariana (Mill.) B.S.P., and Larix laricina (DuRoi) K. Koch (Pinaceae) and Prunus pensylvanica L. f. (Rosaceae). General observations, collections, and phenology. — Adults of P. gibbosa were first observed | August 1986 as occasional males in sweep nets, in flight, or on branch tips of young spruce. No copulating pairs were ob- served. Female oviposition activity was lo- calized on the floral buds of Ledum groen- landicum in an area of roughly 10 metres square and directed a concentration of fu- ture observations and collections here. In addition, occasional observations were made of females ovipositing in a wider area. In 1987, a Malaise trap was installed approx- imately 150 metres from this focus on another aspect of the edge of the cutover forest edge. Trap catches were accumulated over 7-14 day periods and those with P. gibbosa are summarized in Table I. From these collections a minimum flight period of adults, particularly gravid fe- males, occurred from 15 July to 12 August. Oviposition presumably occurs throughout this period and was supplemented by direct observation on 22 July and 11 August. The female-biased sex ratio (50:4) probably re- flects differential behaviour of the sexes with concentration of activity and abundance of males in the vicinity of parasitized ant col- onies where they might wait for emerging females. The lower number of spent females 252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. 1, Sub-sagittal section of floral bud of Ledum groenlandicum with single egg mass of Pseudochalcura gibbosa. 2, Sub-saggital section of fruit of Cyanococcus simulatus with three egg masses of Obeza floridana in different stages of development. 3, Egg of P. gibbosa. 4, Egg of O. floridana. 5-7, Planidium of P. gibbosa: 5, Ventral view of head; 6, Ventral habitus; 7, Lateral habitus. 8, Planidium of O. floridana (dor = dorsal, ven = ventral). All scale bars 0.02 mm. VOLUME 92, NUMBER 2 Table |. trap, 1987. Occurrence of adult P. gibbosa ina Malaise Females Trapping Pernod Days Gravid Spent Males 7-15.vii 9 1 0 0 16-22.vu1 7 5 y) 0 23-30.vil 8 15 1 2 31.vii-l levi 12 4 2 2 12-25.vil 14 8 6 0 Totals 50 33 17 4 relative to gravid females (17:33) is likely due to reduced vigour and probability of flight interception after oviposition. Several Ledum buds were observed to have a dead spent female with its ovipositor still embed- ded. Ledum buds were collected in August of 1986 and 1987 in order to obtain eggs and to attempt rearings of planidia. Eggs were held under a variety of conditions including cold storage at 2°C for four months, but even though the masses darkened relatively quickly after initial deposition (larvae ma- tured internally), the eggs never hatched. A small sample of egg masses adhering to the dehiscing bracts of blooming Ledum on 29 May 1987 also failed to hatch. Eight separate sample collections were made of Camponotus colonies (larvae, pu- pae, and adults) from 24 June to 22 July 1987. Two colonies of C. novaeboracensis about 20 metres from the P. gibbosa ovi- position focus were found on 24 June not to be parasitized by P. gibbosa. The six col- lections of C. herculeanus (L.), on the other hand, were made within 3—5 metres of the focus. In total, four of these were eventually found to be parasitized. Ant larvae with planidia were evident in collections made on 24 June and 15 July while late-pupal or preadult P. gibbosa were found on 22 July. Life history.— Adult females oviposit into the floral buds of Ledum groenlandicum (Ericaceae) in July and August. The ovi- positor is used to penetrate directly through 253 the bracts on the apical half of the buds. Some females are known to maintain this position on the bud for over one hour while others are found dead and devoid of eggs. These buds enclose an undeveloped inflo- rescence which overwinters and blooms in May-June the following season. Small, darkened, rose-brown spots visible on the external surface of the bracts could repre- sent Ovipositional scars but this was not evaluated. Eggs are laid in the extrafloral cavities of the bud and not within plant tissue per se. One to five masses were found near the centre of the bud (not necessarily from the same female as evidenced by differential de- velopment) under several layers of bracts. The caudal stalks of the eggs were all joined at the centre of the egg mass by the apical swelling. The number of eggs in a single mass ranged from 400-1800 eggs with a mean of 943 (SD = 442, n = 10). Dissection of ovaries in unemerged females of P. gib- bosa yielded 2000-2500 fully developed eggs. These relatively large egg masses sug- gest that no more than two or three ovi- positions are made per female on average. It is possible that only a single oviposition is modal and some of the smaller egg masses may have been the result of interrupted ovi- position during collection. Eggs overwinter within the overwintering floral buds of Ledum and become exposed the following spring as the inflorescences expand. The failure of spring-collected eggs to hatch may have been because they were already dead, representing those egg masses left after dispersal of the successful planidia. The overwintering of eggs is well docu- mented for S. tenuicornis. Clausen (1923) presented several ideas, which can be equal- ly applied to P. gibbosa, on how the planidia eventually gain access to the Camponotus nest. Larvae dissected from the eggs of P. gibbosa after four months cold storage were viable and capable of limited (voluntary) movement of the head region. If represen- tative of the condition ofa normally eclosed 254 planidium, then this sharply contrasts with other members of the Eucharitidae whose planidia must actively search for their re- spective hosts. The means by which P. gib- bosa enters the nest of C. herculeanus is unknown. In June 1988, buds of Ledum groenlan- dicum were collected that were swollen to the point that egg masses could be observed protruding between the bracts. Egg hatch was observed shortly thereafter and the rup- tured eggs produced copious amounts of liq- uid. The planidia were observed to undulate through the liquid and were able to occa- sionally lift the anterior region of the body. Camponotus adults were confined with the hatched egg masses and were observed to palpate the liquid mass resulting in the transfer of several larvae to the mouthparts of the ant. In one case, dead mosquitoes were also provided and the following day, macerated tissue was found with several ac- tive planidia attached. It is postulated that the egg fluid liberated by the hatch of the first instar serves to attract foraging ants which results in the transfer of several lar- vae to the ant. Further foraging for food or eventual tending of ant brood likely results in the transfer of several planidia to the host larva, possibly along with a food bolus. This method appears more likely than the trans- fer of an entire egg mass as suggested for S. manipurensis which should result in a tre- mendous number of planidia within a single colony (Clausen 1928). In colonies of ants that were parasitized, mature larvae had an average of 3.35 pla- nidia (SD = 1.56, n = 20). This sample represented a total of 67 planidia primarily concentrated along the creases between seg- ments II-IV (Fig. 9). Planidia remain at- tached externally to the ant larva and do not burrow into the cuticle as observed for some eucharitids. They did not show any signs of feeding (as expansion of body segments) while on the ant larva until the cocoon was formed and pupation of the host initiated (Fig. 10). Clausen (1923) believed that it PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON may be the early stages of hystolysis that trigger development of the eucharitid. En- gorged first-instar larvae (Fig. 12) were found attached to the exuvium of an ant pupa within the cocoon and are presumed to be initially “cast off’ and then either the first- or second-instar larvae migrate to the ven- tral midsection of the ant pupa (Fig. 11) where they resume feeding. No first-instar larvae were found feeding on the ant pupa. In S. tenuicornis, the engorged first-instar larvae were shed in a similar manner and Clausen (1923) reported that larvae moult- ed to a second instar before assuming the new position. The second-instar larva was found at- tached to the posterior ventral portion of the thoracic region on the host pupae (Fig. 11) while the feeding position of the third instar was usually shifted dorsolaterally to the pleural region of the host. A maximum of four second-instar larvae were observed feeding on one host pupa, and as many as four pupae were found in a single cocoon although one or two were more common. Wheeler (1907) also found one cocoon with four pupae. One engorged first instar was seen feeding upon a second-instar of P. gibbosa. Remains of second- or third-instar eucharitid larvae were found in some cocoons along with ma- ture pupae of P. gibbosa. This suggests a cannibalistic mechanism of limiting super- parasitism although the mean number of planidia per host (3.35) would generally not suggest this. The psithergate pupae of the ants were not as strongly deformed as found in some ants attacked by eucharitids (Wheeler 1907), but did show some fusion and enlargement of the thoracic segments and poor definition of the leg segments (Fig. 11). Especially where only one P. gibbosa was found in a cocoon, remains of the ant host were not always devoured and one callow adult ex- hibited a strong resemblance to a deformed psithergate. Description of the immatures. — Egg (Fig. VOLUME 92, NUMBER 2 3): Undeveloped eggs white with smooth chorion. Mean length of egg body 0.17 mm (SD = 0.01, n = 10); of caudal stalk 0.24 mm (SD = 0.02). Mature eggs with well- sclerotized (black) planidial larva occupying less than half of overall volume with head oriented toward stalk; yellowish yolk mass attached to abdominal apex, filling other half of egg body. The small size of the planidium with re- gard to the size of the egg body and large yolk mass may be adaptations to maintain- ing energy stores for an overwintering egg or act as an attractant to the ant host with the simultaneous emergence of an egg mass in the spring. A similar difference in size was not found in either S. fenuicornis or S. cyniformis which were both suspected to overwinter in the egg stage (Clausen 1923, Parker 1937). First instar (Figs. 5-7, 12): Mean length of planidial stage, 0.08 mm (SD = 0.06, n = 10), maximum width 0.05 mm; laterally compressed, oval in cross section. Cranial sensilla absent; hatchet-shaped posterior la- bial plate (found in all other Eucharitinae) either absent or reduced to a barely dis- cernible sclerite below anterior labial scler- ite; pleurostomal spines and circular labial sclerite present. Reduction in number of ter- gites to five (Fig. 7) (12 in other Euchariti- dae); tergopleural line absent. Tergites II-V (numbering not indicative of homology with same tergites in other Eucharitidae) scal- loped along posterior ventral margin; strong ventral extensions of terga absent. Tergite I with three pairs of setae; all other tergal se- tae and enlarged caudal setae absent. The morphology of the first-instar larva of P. gibbosais markedly different from any other described eucharitid. Second instar (Figs. 11, 13): More typi- cally hymenopteriform, white, very weakly sclerotized, with a single pair of mesotho- racic spiracles (propneustic). First-instar exuvium remaining attached to ventral sur- face (Fig. 13). Head region weakly delin- eated; containing a pair of small, pincer-like 255) mandibles. This stage is virtually identical to that of S. cynipiformis Rossi (Parker 1932) and Kapala terminalis Ashmead (as figured in Clausen 1940a) which is the only other described second instar within the Eucha- ritinae. Kapala is distantly related indicat- ing a general conservatism of morphology in the second instar. Third instar (Fig. 14): White and poorly sclerotized. Two thoracic and eight abdom- inal sclerites present; entire dorsal surface minutely tuberculate. Head region defined but well-developed mandibles not ob- served. Exuvium of first instar remaining attached to ventral thoracic region. The morphology of the third instar is identical with S. cyniformis (Parker 1932) and S. tenuicornis (Clausen 1923, 1940a). Pupa (Fig. 15): Typically chalcid-like with exception of series of raised ridges along metasomal tergites in common with other Eucharitidae. Additional information.—oviposition habits of P. gibbosa have been observed by Dr. K. Hagen (University of California, Berkeley, CA) at Amador Pines and Pi- oneer, Amador Co., California. Females were observed oviposited into overwinter- ing leaf buds of Arbutus menziesi Pursh. in a manner which is identical to that de- scribed above. The morphology of the first- instar larva was identical with the larvae taken from Ontario. DISCUSSION Obeza is the first member of the subfam- ily Eucharitinae which has been recorded to Oviposit into fruit and have the egg chamber formed completely within the epidermal layer of the fruit as the egg mass expands during oviposition. In most other Eucha- ritinae, females deposit their eggs into ex- isting cavities in plant tissue such as be- tween the scales of an overwintering flower bud by S. manipurensis and S. tenuicornis (Clausen 1923, 1928) or into flower heads of Picris among the bracts or achenes as in S. cyniformis (Parker 1937). In contrast 256 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ? > sy >» BOC. . Figs. 9-15. Pseudochalcura gibbosa: 9, Cumulative distribution of planidia on 20 mature ant larvae; 10, Early feeding stages of planidia on ant host in preparation for pupation (cocoon removed); 11, Second instars feeding on ant pupa; 12, Engorged first instar; 13, Second instar; 14, Third instar; 15, Pupa of male. Scale bars in mm. VOLUME 92, NUMBER 2 to both Stilbula and Pseudochalcura, O. floridana and the undescribed species from Mexico deposit relatively few eggs in each egg mass with each female undertaking many separate ovipositions. Morphologically, adults of Obeza are very similar to Stilbula and generic differentiation is based on the possession of lateral propodeal processes and posterior extension of the genae in Obeza (Heraty 1985). If other species of Obeza share similar oviposition habits, this bio- logical distinction serves to support recog- nition of the two genera. The planidium of Obeza shows a number of distinctive features that provide recog- nition at the generic level but otherwise it is fairly typical of most Eucharitinae (see Heraty and Darling 1984). The pair of large hook-like cranial processes is similar to the anterior cranial spines found in Peri/ampus of the Perilampidae (Heraty and Darling 1984). However, the processes are unscler- otized and therefore regarded as of different derivation (not homologous) from those of Perilampus. The planidium of P. gibbosa is highly de- rived within the Eucharitidae. In a family where conservatism of first-instar larvae 1s extreme, the reduced number of terga and lateral compression of body segments are anomalous. The reduction of segmentation of this planidium would be expected to al- low very limited mobility in comparison with other eucharitids including Sti/bu/a and Obeza. In contrast to the derived condition of the planidium, life history information and morphology of the other larval stages is virtually identical to S. tenuicornis (the reader is referred to Clausen (1923) for a full description) and again suggests a close relationship within the lineage regardless of their very different adult morphology. Poor descriptions for planidia of Sti/bula render them as generally uninformative for comparison with Obeza or Pseudochalcura. However, it is notable that the tergopleural line (a desclerotized line that runs laterally across the tergites and found in some form in all other Eucharitidae) was absent in DiI Obeza and Pseudochalcura, and not figured in any drawings of Stilbula planidia. Clau- sen (1940a) was the first author to illustrate the tergopleural line in three other genera of Eucharitinae. In that paper, he referred to the absence of a line in S. cyniformis as probably due to an oversight by Parker (1932) but neglected to mention the absence in his own earlier illustrations of S. mani- purensis and S. tenuicornis. Heraty and Darling (1984) similarly considered the ter- gopleural line as a synapomorphy of the Eu- charitidae. It is now apparent that the ab- sence of a tergopleural line may be a common feature of a clade including Sti/- bula, Obeza, and Pseudochalcura. The po- sition of this group with respect to other Eucharitidae will need to be determined to assign correct polarity to this state. With few exceptions, the immature stages of the Eucharitidae have proven to be very conservative in the amount of morpholog- ical change and behavioural adaptation. This allows the immature stages to provide use- ful evidence of higher level relationships. Stilbula and Obeza share similar adult mor- phology but dissimilar oviposition habits; Stilbula and Pseudochalcura share similar life histories and morphology of immatures but dissimilar adult morphology. Some of the larval and ecological characters de- scribed and compared here could represent synapomorphies or ground-plan characters of the clade discussed (e.g. general similarity of morphology of larval stages including ab- sence of tergopleural line in planidium), thus supporting the monophyly of the clade but not providing any resolution of the rela- tionships among the four genera. In the ab- sence of data for other taxa, particularly Lophyrocera, other characters appear to be autapomorphies (e.g. cranial processes, Ovi- position into plant tissue in Obeza; reduc- tion of segmentation in planidia in Pseu- dochalcura) that furnish no insight into inter-generic relationships. Information on the within-nest biology of Obeza, Lophy- rocera, and other species of Pseudochalcura as well as oviposition habits and ant host 258 selection of these taxa throughout their geo- graphical ranges will provide invaluable in further resolving the relationships of these taxa. The description of the life history of P. gibbosa, though more extensive than that of O. floridana, is not exhaustive. A number of key questions still remain unanswered and certain suggestions or speculations re- quire verification. How does the planidium travel from the Ledum buds to the ant nest and when is this accomplished? Reader (1977) listed only flies and bees visiting Le- dum flowers and made no mention of ants. Do the planidia discriminate between young ant larvae (maturing in the following sea- son) and mature larvae (pupating in same year) (Sanders 1964, 1972)? If not, how do the non-feeding planidia survive if they must remain on the host for at least a full year? These are likely to be answered only with additional observation and possibly exper- imentation. ACKNOWLEDGMENTS We thank J. B. Woolley (Dept. of Ento- mology, Texas A&M University), C. J. Sanders (Great Lakes Forestry Centre, Sault Ste. Marie), and G. J. Umphrey (Dept. of Biology, Carleton University, Ottawa) for their comments on this manuscript. S. Tay- lor (Great Lakes Forestry Centre) kindly provided identifications of plant specimens collected in Ontario. P. Fryxell (U.S.D.A. Cotton Laboratory, College Station, TX) provided identifications of plant specimens collected in Florida and Mexico. Dr. K. Ha- gen (Division of Biological Control, Uni- versity of California, Berkeley) provided specimens of first instar P. gibbosa for examination. This paper was approved as article number TA-24649 of the Texas Agricultural Experimental Station. Repre- sentatives of adults and slide mounts of first- instar larvae are deposited as voucher num- ber 469 in the Department of Entomology Insect Collection, Texas A&M University, College Station, TX. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Clausen, C. P. 1923. The biology of Schizaspidia ten- uicornis Ashm., a eucharid parasite of Campo- notus. Ann. Entomol. Soc. Am. 16: 195-217. 1928. The manner of oviposition and the planidium of Schizaspidia manipurensis n. sp. (Hymen., Eucharidae). Proc. Entomol. Soc. Wash. 30: 80-86. 1940a. The immature stages of the Euchar- idae. Proc. Entomol. Soc. Wash. 42: 161-170. 1940b. The oviposition habits of the Eu- charidae (Hymenoptera). Wash. Acad. Sci. 30: 504— 516. 1941. The habits of the Eucharidae. Psyche 48: 57-69. Davidson, D. W. 1988. Ecological studies of neo- tropical ant gardens. Ecology 69: 1138-1152. Heraty, J. M. 1985. A revision of the Nearctic Eu- charitinae (Hymenoptera: Chalcidoidea: Eucha- ritidae). Proc. Entomol. Soc. Ont. 116: 61-103. Heraty, J. M. 1986. Pseudochalcura (Hymenoptera: Eucharitidae), a New World genus of chalcidoids parasitic on ants. Syst. Entomol. 11: 183-212. Heraty, J. M. 1989. Morphology of the mesosoma of Kapala (Hymenoptera: Eucharitidae) with em- phasis on its phylogenetic implications. Can. J. Zool. 67: 115-125. Heraty, J. M. and D.C. Darling. 1984. Comparative morphology of the planidial larvae of Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea). Syst. Entomol. 9: 309-328. Parker, H. L. 1932. Notes on a collecting spot in France and a chalcid larva (Stilbula cynipiformis Rossi) (Hymenoptera: Eucharidae). Entomol. News 43: 1-6. Parker, H. L. 1937. The Oviposition habits of Sti/- bula cynipiformis Rossi (Hymen., Eucharidae). Proc. Entomol. Soc. Wash. 39: 1-3. Parker, H. L. and W. R. Thompson. 1925. Notes on the larvae of the Chalcidoidea. Ann. Entomol. Soc. Am. 43: 384-395. Pierce, W. D. and A. W. Morrill. 1914. Notes on the entomology of the Arizona wild cotton. Proc. Entomol. Soc. Wash. 16: 22. Reader, R. J. 1977. Bog ericad flowers: self-incom- patibility and relative attractivness to bees. Can. J. Bot. 55: 2279-2287. Sanders, C. J. 1964. The biology of carpenter ants in New Brunswick. Can. Ent. 96: 894-909. Sanders, C. J. 1972. Seasonal and daily activity pat- terns of carpenter ants (Camponotus spp.) in northwestern Ontario (Hymenoptera: Formici- dae). Can Ent. 104: 1681-1687. Wheeler, W. M. 1907. The polymorphism of ants, with an account of some singular abnormalities due to parasitism. Bull. Am. Mus. Nat. Hist. 23: 1-93. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 259-270 FIRST RECORD OF A BAGWORM MOTH FROM HAWAII: DESCRIPTION AND INTRODUCTION OF BRACHYCYTTARUS GRISEUS DE JOANNIS (LEPIDOPTERA: PSYCHIDAE) DONALD R. DAvis Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. Abstract. — Until recently no representative of the bagworm family Psychidae was known to occur in the Hawaiian Islands. The first infestation of Brachycyttarus griseus De Joannis was discovered during 1984 in a residential area of Haiku on Oahu. The species has since spread to several areas on Oahu and Kauai, where the larvae feed on the introduced grass, Paspalum conjugatum Berg. Originally described from Vietnam, B. griseus has also be- come established in Guam where it is parasitized by a tachinid fly, Stomatomyia species. All stages of B. griseus are described and illustrated and a map showing its present Hawaiian distribution is included. Key Words: Until 1984 no species of the bagworm family Psychidae was known to occur in the Hawaiian Islands. In 1984 I received spec- imens, which I later determined as Brachy- cyttarus griseus De Joannis, from Po-Yung Lai of the Hawaii Department of Agricul- ture. The infestation was discovered in a residential area of Haiku (Kaneohe, see Map 1), Oahu. This site is about 15 mi [24 km] from Honolulu Harbor, the largest shipping dock in Hawaii, and about 17 mi [27 km] from Honolulu International Airport. The bagworm is now established on Oahu and has been encountered at nine different sites in or near Honolulu and Kaneohe (Map 1). The larva is known to feed on Hilo grass, Paspalum conjugatum Berg, another intro- duced species that has already interfered with the propagation of several rare endem- ic plants (Vitousek et al. 1987). On Guam the larva is reported to feed on Zoysia pun- gens Willd. (= japonica Steud.) and ““mixed native grasses” (Muniappan, in litt.). Lepidoptera, Psychidae, bagworm, introduced species, Hawaii Brachycyttarus griseus was first reported from Hanoi, Vietnam (De Joannis 1929). Considering the magnitude of trafhe and shipping from Vietnam, particularly within the previous two decades, it is likely that fertile females or eggs were introduced into Honolulu during that period. However, lit- tle is known about the present species dis- tribution. From adults examined in the Smithsonian Institution (USNM), it is ap- parent that B. griseus has existed in the Phil- ippines for some time. Several specimens were collected at Los Banos, Luzon, some as early as 1918. Labels indicate that a few specimens were reared from grass, thus agreeing with all other host records of B. griseus. Also present in the same collection is a series of grass-covered cases from Ser- dany, Malaysia that is identical to the Hawaiian material. No adults were reared from the Malaysian cases. On the basis of specimens and reports received from R. Muniappan, L. Stevens, and R. Shook of 260 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON O‘ahu b———J 5km Map. |. Distribution of Brachycyttarus griseus in Oahu. Localities represented are: 1, Ewa; 2, Waipahu; 3, Pearl City; 4, Waimalu; 5, Ajea/Halawa; 6, Aliamanu/Moanalua; 7, Salt Lake; 8, Kalihi; 9, Kaneohe. the University of Guam, this grass-feeding bagworm was well established on Guam by 1977. Thus, the Hawaiian population could have originated from several sources, with either Vietnam or Guam perhaps the most likely. No parasites have been reared from the Hawaiian bagworm population. Dipterous parasites reared from B. griseus in Guam and submitted by R. Shook have been iden- tified by C. W. Sabrosky as Tachinidae: Sto- matomyia species. To facilitate the recognition of B. griseus, which has become more widely distributed and, thus, of greater public concern, a de- scription of all developmental stages is pro- vided. Brachycyttarus griseus De Joannis Figs. 1-48; Map 1 Brachycyttarus griseus De Joannis, 1929: 543.—Dierl, 1971: 61. Acanthopsyche (Brachycyttarus) griseus (De Joannis).— Gaede, 1933: 735. Adult (Figs. 1-2).— Length of forewing: 4, 6.5-7.4 mm. A moderately small, slender, broad wing moth with uniformly dark gray wings except for short, grayish white fringe on outer margin of both wings. Apex of cu- cullus on male genitalia with 3 small spines. Female vermiform, naked, and without wings or segmented appendages. Larva with 6 stemmata, the most caudal 3 pairs with vestigial, flattened lenses. VOLUME 92, NUMBER 2 Head: Vestiture sparse, gray. Mouthparts absent. Antenna 18-20 segmented, bipec- tinate with long slender branches; antennal sensilla long, length about 6-7 the di- ameter of supporting branch. Thorax: Sparsely covered with brownish gray to gray piliform scales dorsally over dark cuticle, scales light gray to white ven- trally. Forewing uniformly dark gray dor- sally and ventrally with grayish white, outer fringe scales. Hindwing slightly lighter gray dorsally, mostly white ventrally; fringe grayish white. Legs with femora dark gray, tibiae and tarsi light brown to stramineous. Foreleg the longest; epiphysis long and slen- der, about 0.8 the length of tibia. Midleg and hindleg without tibial spurs. Abdomen: Vestiture dark gray dorsally, grayish white ventrally; cuticle dark reddish brown to black. Eighth tergite (Fig. 41) and sternite (Fig. 42) as illustrated. Male genitalia (Figs. 39-40): Tegumen and vinculum relatively elongate, broadly rounded. Valva approximately half the length of genitalia, divided apically into a smoothly rounded costal lobe and a smaller cucullar lobe bearing 3 short spines. Ae- doeagus approximately same length as gen- italia, straight, relatively stout, with caudal end slightly enlarged. Female genitalia (Fig. 44): Anal papillae reduced to a pair of faintly setose lobes. Apophyses absent. Corpus bursae reduced to a small, digitate lobe anterior to short, broad ductus seminalis. Ductus spermathe- cae elongate, slender, not coiled. Egg (Figs. 5-6).—Length approximately 0.44-0.56 mm; width 0.35-0.4 mm. Cho- rion smooth with stellate micropyle at one end; micropyle with 22-25 slender low ndges radiating out from central disk; arms oc- casionally anastomosing, forming closed cells. Larva (Figs. 7-22, 31-38).—Length of largest larva 15 mm; maximum diameter 2.6 mm. Body color mostly white with scat- tered dark pigmentation over head and tho- racic plates. 261 Head: Maximum width 1.5 mm. Pig- mentation as in Fig. 32. AF2 elongate, ex- tending to labrum. Six pairs of stemmata present; anterior three pairs (3-5) normal, posterior three (1—2, 6) slightly reduced with flattened corneas. S2 arising equidistant be- tween stemmata | and 6 or closer to 1. La- brum with four pairs of almost equal size epipharyngeal setae. Mandible irregularly truncate; distinct cusps barely discernible. Sensilla of antenna as in Figs. 15-17; max- illary palpus as in Figs. 11-12. Thorax: Lightly pigmented as shown in Fig. 31. Pronotum with D1 approximate to XD1. Coxal plates fused medially, with a pair of prominent lobes projecting ante- riorly between coxal setae; Cl the longest. Meso- and metanotum with L2 separate from pinacula bearing L1 and 3. Tarsal claw relatively long and straight, with a small axial spine (Figs. 19-20). Abdomen: Pinacula usually poorly de- fined. D1 and 2 on separate pinacula except on A8 where they arise together on a rela- tively large pinaculum. SD2 minute, sepa- rate from pinaculum bearing SD1 and usu- ally slightly above and anterior to spiracle. L1 and 2 together on same pinacula on Al- 2, separate on A3-9. A9 with D2 and SD1 on same pinacula. Prolegs A3-6 with 20- 22 crochets in lateral penellipse; anal proleg with 15-17 crochets. Larval case (Fig. 3).—Length 10-15 mm; diameter approximately 5-6 mm. Exterior of case densely covered with brownish grass fragments, roughly spirally arranged, thus imparting a very shaggy appearance. For pupation, case is suspended by a slender, silken strand about 0.5-1.0 the length of case (Fig. 4). Male pupa (Figs. 23-26, 47-48).— Max- imum length 6 mm; width 1.6 mm. Young pupa light brown, darkening with age until head and thorax and all appendages dark fuscous to black as well as narrow dorsal and ventral interrupted band composed of terga and sterna of Al-8. Vertex smooth, subtruncate; frontal ridge absent. Antennal 262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Brachycyttarus griseus. 1, Male, Kaneohe, Oahu, length of forewing 6.9 mm. 2, Male holotype, Hanoi, Vietnam, length of forewing 6.6 mm. 3, Larval case, length 13 mm. 4, Larval cases, attachment for pupation. 5, Egg (176 um). 6, Micropyle of egg (43 um). (Scale lengths in parentheses; bar scale for Fig. 6 = Fig. 5.) VOLUME 92, NUMBER 2 sheath extending to A3. Wing sheaths ap- proximately same length, to middle of A3. A6-8 with a relatively short anterior row of dorsal spines as follow: A6 = 7-9 spines; A7 = 10-13; A8 = 7-10. Posterior row of dorsal spines absent. Cremaster composed of a stout pair of acute, anteriorly directed spines from venter of AlO (Figs. 25-26). Female pupa (Figs. 27-30, 45-46).— Maximum length 7 mm; width 2.8 mm. Light brown to stramineus in color. All body appendages either absent or reduced to mi- nute tubercules. Abdomen with both ante- rior and posterior dorsal spine rows reduced and present but never together on same seg- ment; anterior row present only on A6 (= 11-14 spines) and A7 (= 10-13); posterior rows present on Al (= 35-40), A2 (= 17- 21), A3 (= 17-20), A4 (= 10-12); AS with- out either anterior on posterior spine rows. Cremaster composed ofa relatively slender, more reduced pair of anteriorly directed spines from venter of A10 (Figs. 29-30). Type.— Holotype, 4; in the Muséum Na- tional d’Histoire Naturelle, Paris. Type locality.— Vietnam: Hanoi. Host.—Poaceae: Paspalum conjugatum Berg, Zoysia pungeus Willd. (= japonica Steud.) and probably other grasses. Parasite.—Tachinidae: Stomatomyia species. Distribution (Map 1|).— Definitely known to occur in Vietnam, Philippines, Guam, and Hawaii, but probably widely distrib- uted in southeast Asia. In Hawaii, reported only from Oahu and Kauai below 150 me- ters, but in time will undoubtedly spread to other major islands. Discussion. —In his review ofa few Asiat- ic Psychidae, Dierl (1971) treats five species of Brachycyttarus. Most of the species are similar in general appearance, with similar male genitalia. Consequently, species iden- tification within this complex is difficult and still hindered by some uncertainties. Only one species, B. fuscus Dierl, lacks the white scales on the underside of the hindwing. Dierl illustrates the wing venation of four 263 of the five species and indicates that both B. griseus and B. fasciatus Dierl are similar in having M2 and 3 converge at the outer margin of the hindwing (see Fig. 43). Al- though I have noted some venational vari- ation in the specimens examined, the wing structure of the Hawaiian species appears closest to that illustrated for B. fasciatus. This is particularly evident in the stalking of R2+3 with R4+5 in the forewings and the abbreviated length of the basal radial cell in the hindwing. Lacking in Dierl’s drawing of fasciatus is the presence of a bas- al stem of R that is separate from Sc as shown in Fig. 43. Partly because the male genitalia of the Hawaiian species most re- semble that of B. griseus, I have considered them conspecific. Although the hosts of B. fasciatus are not stated, Dierl’s comments that the larva is polyphyagous and lives only in forests under large trees and small shrubs do not suggest the species attacking grasses in Guam and Hawaii. De Joannis (1929) likewise did not state the host of B. griseus, but he did describe the larval case as being covered with grass fragments, which sug- gests grass as the foodplant. The type lo- cality of B. fasciatus (Jhawani, Nepal, 200 m) also suggests a more northern species than B. griseus. The wing fringe of fasciatus was described as brown by Dierl, compared to grayish white in the Hawaiian specimens. This character has to be used with discre- tion, however, as only the outer fringe scales are pale colored, and these are frequently lost in rubbed specimens, as is true for the type of B. griseus (Fig. 2) and in most of the specimens I have examined from Guam and Hawaii. Dierl has introduced another problem in this small but troublesome genus with re- gard to the relatively unknown Javan species, “Pteroma” reijnvaanii Van Leeu- wen. Because the original illustrations of the adult and larval case of this species most resemble B. griseus, reijnvaanii could be the senior synonym of the former. Presently this is impossible to resolve because of the in- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON is griseus, larva. 7, Anterior view of head (0.43 mm). 8, Labial palpi and spinneret Maxilla and labium, ventral view (200 um). 11, Maxillary llary palpus (5 um). (Scale lengths in parentheses; bar scale Figs. 7-12. Brachycyttaru (60 um). 9, Ventral view of head (0.46 mm). 10, palpus (50 wm). 12, Apical sensilla basiconica of maxi for all photographs = Fig. 7.) VOLUME 92, NUMBER 2 Figs. 13-18. Brachycyttarus griseus, larva. 13, Lateral view of head (0.3 mm). 14. Detail of stemmatal area (100 nm). 15, Antenna (60 um). 16, Detail of Fig. 15 (30 um). 17, Apical view of antenna (27 um). 18, Mesothorax, ventral view (0.43 mm), (Scale lengths in parentheses; bar scale for all photographs = Fig. 13.) 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON upa. 19, Prothoracic tarsal claw (50 wm). 20, Detail of axial 22. Anal crochets, A10 (60 um). 23, Male pupa, ventral f dorsal spines A8 (86 um). (Scale lengths in parentheses; Figs. 19-24. Brachycyttarus griseus, larva and p seta in Fig. 19 (5 um). 21, Crochets of A6 (60 um). view of head (0.6 mm). 24, Male pupa, anterior row 0 bar scale for all photographs = Fig. 19.) VOLUME 92, NUMBER 2 267 Figs. 25-30. Brachycyttarus griseus, pupae. 25, Male cremaster, A10, caudal view (136 um). 26, Lateral view of Fig. 25 (150 um). 27, Female, dorsum of A6 and 7, with anterior spines on A7 (231 um). 28, Detail of A7 spines in Fig. 27 (75 wm). 29, Female cremaster, A10, anterior view (100 um). 30, Lateral view of Fig. 29 (60 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 25.) 268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ( | s 83) } S$S34SS¥ rin >“ $2 36 Figs. 31-38. Brachycyttarus griseus, larval chaetotaxy. 31, Lateral view of prothorax, mesothorax, and abdominal segments 1, 6, 8, and 9. 32, Dorsal view of head (0.5 mm). 33, Ventral view. 34, Lateral view. 35, Dorsal view of A8-10. 36, Mandible (0.2 mm). 37, Labrum, dorsal view (0.2 mm). 38, Ventral view showing epipharyngeal setae. (Scale lengths in parentheses.) VOLUME 92, NUMBER 2 Figs. 39-48. Brachycyttarus griseus. 39, Male genitalia, ventral view (0.25 mm). 40, Aedoeagus. 41, Tergite A8 (0.25 mm). 42, Sternite, A8. 43, Wing venation. 44, Female genitalia (0.25 mm). 45, Female pupa, dorsal view (1.0 mm). 46, Ventral view. 47, Male pupa (1.0 mm). 48, Ventral view. (Scale lengths in parentheses.) > 270 adequacy of the original description and the subsequent disappearance of all type ma- terial. All eggs examined in this study, including Figs. 5 and 6, were removed from the bodies of preserved females. Consequently, their relative dimensions or even surface texture may not be typical of deposited eggs. These data eventually need to be compared with externally collected eggs for possible dis- crepancies. Because this report includes the first de- scription of the egg, larva, and pupa for any member of Brachycyttarus, it is not possible to compare these stages between related species. Possible diagnostic larval charac- ters for B. griseus involve the elongate AF2 seta, the relative reduction of the posterior three stemmata (Fig. 14), and the reduction of the SD1 pinaculum on A9 with the con- sequent separation of SD2 and L1 from SD 1. Comparisons involving the sensilla of the larval maxilla and antenna must await con- siderably more SEM work with psychid lar- vae. The relative amount or distribution of larval head and body pigmentation could be of some significance, although this can vary both within and between instars. The male pupa of B. griseus is unusual in completely lacking the posterior row of an- teriorly oriented, dorsal abdominal spines typical of most Psychidae studied (Davis 1975). The anterior row is present but only on segments A6-8. As is typical for ver- miform females, the dorsal spines are great- ly reduced in size in the female pupa, but their distribution is unusual. No abdominal segment possesses a full complement of both anterior and posterior rows and AS is totally lacking in spines. Again, whether these characters are significant at the specific or generic level must await discovery of other Brachycyttarus pupae. ACKNOWLEDGMENTS I wish to thank Po-Yung Lai of the Pest Control Branch, Hawaii Department of Ag- riculture and several of his staff for speci- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mens and data of Brachycyttarus griseus from Oahu. In particular, I am grateful for the efforts of George Funasaki, Ron Hue, and Bernarr Kumashiro. Similarly, I also wish to acknowledge Scott Miller of the Bishop Museum and Hidiyuki Chiba of the University of Hawaii for helpful assistance. I am grateful to Dan Otte of the Philadel- phia Academy of Natural Sciences for the outline map of Oahu and to George Funa- saki for the latest distributional records of B. griseus on Oahu. Thanks also go to R. Muniappan, L. Stevens, and Rosalie Shook of the University of Guam for study ma- terial from Guam. I am indebted to Vichai Malikul and Young Sohn of the Department of Entomology, Smithsonian Institution, for the line drawings and to Susann Braden and Brian Kahn of the Smithsonian SEM Lab and Victor Krantz of the Smithsonian Pho- tographic Laboratory for photographic as- sistance. Iam thankful to Allan Watson and the British Museum (Natural History) and Joel Minet of Muséum National d’Histoire Naturelle for allowing me to study type ma- terial in their collections. I am grateful to Silver West of our Department for prepar- ing the final draft of the manuscript. LITERATURE CITED Davis, D. R. 1975. A review of the West Indian moths of the family Psychidae with descriptions of new taxa and immature stages. Smithsonian Contributions to Zoology 188: 1-66, figs. 1-206, tables 1-5. Dierl, W. 1971. Biologie und Systematik einiger Asia- tuscher Psychidae-Arten (Lepidoptera). Entomolo- gischen Abteilung der Zoologischen Staatssamm- lung Miinchen, Khumbu Himal 4(1): 58-79, figs. 1-21. Gaede, M. 1933. Psychidae. Jn A. Seitz, The Macro- lepidoptera of the World, 10 (Fauna Indoaustral- ica): 729-742. Joannis, J. De. 1929. Lépidoptéres Héterocéres du Tonkin, pt. 2. Annales de la Société Entomolo- gique de France 98(4): 361-557, pls. 3-6. Vitousek, P. M., L. L. Loope, and C. P. Stone. 1987. Introduced species in Hawaii: Ecological effects and opportunities for research. Trends in Ecology and Evolution 2(7): 224-227. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 271-273 TRIPHLEBA VITRINERVIS (MALLOCH), AN UNRECOGNIZED SPECIES OF CRINOPHLEBA BORGMEIER (DIPTERA: PHORIDAE) BRIAN V. BROWN Department of Entomology, University of Alberta, Edmonton, Alberta T6G 2E3, Can- ada. Abstract. — Triphleba vitrinervis (Malloch) belongs to the genus Crinophleba Borgmeier (new combination), based on a comparison with the male of C. angustifrons Goté. The male terminalia are illustrated and characters are given to allow the recognition of male Crinophleba in the Manual of Nearctic Diptera key to Phoridae. C. vitrinervis may represent the undescribed male of C. rostrata Borgmeier, known only from female specimens. Key Words: The taxonomy of phorid flies is plagued by genera described from one sex, leaving open the possibility that the male sex may be described in one genus and the female in another (e.g. Brown 1986). An example is the genus Crinophleba Borgmeier, de- scribed only from female specimens (Borg- meier 1967). It was compared to the genus Anevrina, based on the setulose Rs vein of the wing, but the female of Crinophleba had a much larger proboscis, shorter tergite 6, different terminalia and weaker tibial setae than Anevrina. No described males of North American phorids could be linked with Crinophleba, and it was not until a second species from Japan, C. angustifrons Goto, was described from both sexes (Gotd 1983) that a male was known. The newly de- scribed male had several distinctive char- acters, including a narrowed frons and a lack of cerci, that were not known in any North American species of phorid. Examination of the terminalia of 7. vitrinervis, however, showed that it was a male Crinophleba (NEW COMBINATION). That this was not recog- nized earlier is not surprising, since C. vi- Crinophleba, Triphleba, Phoridae, Diptera, taxonomy trinervis lacks the narrowed frons of C. an- gustifrons and since in the past characters of the male terminalia were not used exten- sively. Although they have not been definitely associated, C. rostrata may be conspecific with C. vitrinervis. Definitive evidence of this relationship would be furnished by col- lecting the adults im copula. If they are con- specific, the valid name would be C. vitri- nervis. Below, the genus Crinophleba is diag- nosed; the male terminalia of C. vitrinervis are briefly described and illustrated. Genus Crinophleba Borgmeier Type species C. rostrata Borgmeier (by original designation). Diagnosis.—Frons without median fur- row. Female with proboscis elongate. An- episternum bare. Fore tibia with short, api- cal setae. Mid and hind tibiae with 2 anterior and | dorsal setae. Wing vein Rs setulose. Male terminalia lack cerci. Basiphallus ex- panded. 272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 2 Crinophleba vitrinervis (Malloch) Trupheoneura vitrinervis Malloch 1912: 419. Triphleba vitrinervis, Brues 1950: 41; Borg- meier 1963: 32. Description. — Male terminalia: Epan- drium continuous anteriorly, ring-shaped (Fig. 1). Left side of epandrium setose (Fig. 2); surstylus separate, with small ventral projection. Right side of epandrium (Fig. 3) setose, with posteroventral process. Hypan- drium deeply cleft (Fig. 1), right lobe broad- er, each lobe with small lateral tooth. Basi- phallus with expanded ventral portion (Fig. 5); posteriorly extended (Fig. 4). Sclerite be- hind basiphallus recurved above basiphal- lus, expanded into large lateral plate with two dorsal projections on left side and nar- row lateral projection on right side. Poste- rior, transverse sclerite present. Cercus ab- sent. Material examined: CANADA. Alberta: Opal, 53°59'N, 113°13'W, 1 4, 20-22.vii. 1989, B. V. Brown, Malaise trap, sand, jack pine. Ontario: Guelph, South Arboretum, | 4, 11-16.v.1985, 1 6, 24-30.v.1985, 34, 19- 24.vi.1985, 1 4, 6-10.viii. 1985, Malaise trap, forest edge, 1 4, 7-11.vi.1985, Malaise trap, wet shrubby meadow, B. V. Brown; Stouff- ville, 2 6, 26.v.-2.v1.1985, B. V. Brown, Ma- laise trap (all specimens deposited in col- lection of the author). This species has also been collected in the states of Maryland, Michigan and New Hampshire in the U.S.A. (Borgmeier 1963). Remarks: The wider frons of C. vitrinervis easily separate it from the male of C. an- gustifrons. Characters for separating female C. rostrata from C. angustifrons are given by Goté (1983). In order to facilitate recognition of the genus Crinophleba in the Manual of Nearc- 273 tic Diptera, Volume 2 key to Phoridae, cou- plet 8 should be changed to the following (figure numbers refer to figures in the Man- ual of Nearctic Diptera, Phoridae chapter): 8. Male lacking cerci. Female with proboscis greatly elongated, rigid (Fig. 4). Arista subapi- cal. Scutellum with two strong posterior bristles and two much shorter coarse setae anteriorly. Tergite 6 of female short. Tibial bristles weak ; Crinophleba Borgmeier 2 spp.; widespread in Canada and northern U.S.A. Male with cerci present. Proboscis short, broad. Arista clearly dorsal (Fig. 35). Scutellum with four subequal bristles. Tergite 6 of female elon- gate. Tibial bristles strong (Fig. 65) ........ Anevrina Lioy 9 spp.; widespread in Canada and U.S.A. ACKNOWLEDGMENTS The author would like to thank Dr. G. E. Ball for reviewing this manuscript and Dr. T. Got6 for originally suggesting that 7. vi- trinervis may have been a Crinophleba. LITERATURE CITED Borgmeier, T. 1963. Revision of the North American phorid flies. Part I. The Phorinae, Aenigmatiinae and Metopininae, except Megaselia. Studia Ent. 6: 1-256. 1967. A new genus and species of phorid fly from Canada (Diptera: Phoridae). Can. Ent. 99: 221-223. Brown, B. V. 1986. Striking sexual dimorphism and its taxonomic consequences among New World phorid flies (Diptera: Phoridae). Proc. Entomol. Soc. Wash. 88: 787-788. Brues, C. T. 1950. Family Phoridae. /n Guide to the Insects of Connecticut. Part IV. Diptera, 4th fas- cicle. Bull. Conn. Geol. Nat. Hist. Survey 75: 33- 85. Goto, T. 1983. A new phorid fly of the genus Crino- phleba from Japan (Diptera: Phoridae). Kontyt, Tokyo 51: 376-383. Malloch, J. R. 1912. The insects of the dipterous family Phoridae in the United States National Mu- seum. Proc. U.S. Nat. Mus. 43: 411-529. — Figs. 1-5. Crinophleba vitrinervis. Figs. 1-3, male terminalia (bar = 0.1 mm, all figs. to same scale). 1, ventral view. 2, left lateral view. 3, right lateral view. Figs. 4-5, aedeagus (bar = 0.1 mm, both figs. to same scale). 4, left lateral. 5, frontal. ABBREVIATIONS: e—epandrium, h—hypandrium, p—posterior extension of basiphallus, t—hypandrial tooth, ts—transverse sclerite, y—ventral expansion of basiphallus. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 274-281 DESCRIPTIONS OF THE FEMALES OF THREE POL YCENTROPUS SPECIES (TRICHOPTERA: POLYCENTROPODIDAE) KEVIN M. HOFFMAN AND JOHN C. MORSE Department of Entomology, Clemson University, Clemson, South Carolina 29634- 0365. Abstract. —The females of Polycentropus blicklei Ross and Yamamoto and Polycentropus carlsoni Morse are described and illustrated for the first time, and the female of Polycen- tropus maculatus Banks is redescribed and illustrated. Diagnoses are provided to distin- guish these females both from each other and from those of other Polycentropus species in the confusus group. Some additional life-history information for P. carlsoni males is provided. Key Words: The confusus group is one of five New World species groups within the genus Poly- centropus, and contains fourteen named and one unnamed species (Hamilton 1986). The need for more associations of both females and larvae with the described males of the confusus group was stressed by Hamilton (1986), who reviewed but did not revise the group. To date, the females of eight species have been described, and larvae have been associated with but one species. The only key to females of the confusus group was provided by Ross (1944, pp. 63-64, cou- plets 21-26) for the females of seven species, one of which was not associated with males and was referred to as Polycentropus species a. The female of one other species, Poly- centropus neiswanderi Ross, has been de- scribed in the intervening years (Ross 1947). Polycentropus blicklei Ross and Yama- moto, Polycentropus carlsoni Morse, and Polycentropus maculatus Banks are all members of the confusus group. Both P. blicklei and P. maculatus have been re- ported from several states in the eastern United States as well as from provinces in eastern Canada (Hamilton 1986), while P. Trichoptera, Polycentropodidae, Polycentropus, females, descriptions carlsoni has been reported from only three first to second order streams in the upper Piedmont of South Carolina and lower Val- ley and Ridge of Alabama (Morse 1971, Lago and Harris 1987). Of these three species, only the female of P. maculatus has been described (Ross 1944). The efforts to discover females of P. blicklei and P. carl- soni were in response to a preliminary re- view considering P. carlsoni for possible protection under the United States Endan- gered Species Act of 1973, as amended. The present paper describes the females of both P. blicklei and P. carlsoni for the first time, redescribes the female of P. maculatus, and provides additional life-history information on males of P. carlsoni. MATERIALS AND METHODS The associations of females with males of P. blicklei and P. carlsoni were accom- plished in two steps. First, intensive collec- tion efforts for adults were made during 1988 and early 1989 using Malaise traps at and near the type locality of P. carlsoni. Malaise traps similar to those described by Townes (1972) were deployed from April 1988 to VOLUME 92, NUMBER 2 February 1989 at the type locality on Wild- cat Creek (site 2, see below) and on four other nearby streams within the Clemson University Experimental Forest surround- ing Lake Issaqueena, Pickens County, South Carolina. Trap collections were examined weekly until the end of August, biweekly from September to the end of November, and monthly from December 1988 to the end of February 1989. Locality data of the five sites and dates of trap operations are as follows: unnamed tributary of Indian Creek (site 1), elevation 230 m, 7 April to 27 June 1988: Wildcat Creek (site 2), elevation 235 m, 4 April 1988 to 28 February 1989; un- named tributary #1 of Sixmile Creek (site 3), elevation 225 m, 5 April to 20 June 1988; unnamed tributary #2 of Sixmile Creek (site 4), elevation 220 m, 24 April 1988 to 28 February 1989; unnamed creek at Holly Springs Picnic Area (site 5), elevation 220 m, 5 April 1988 to 28 February 1989. In addition, five localities on streams sur- rounding Lake Issaqueena were sampled during May 1988 using an Ellisco® light trap with a 15-watt ultraviolet bulb. The trap was operated for a two hour period following sunset. The five localities and dates of sampling are as follows: site 5 on 8 May, Sixmile Creek on 11 May, site 4 on 20 May, site 2 on 21 May, and an unnamed first order stream on 22 May. The second step consisted of identifying Polycentropus females by using the key pro- vided by Ross (1944) and by comparing un- determined females with those associated with males during three previous surveys in South Carolina, North Carolina, and Geor- gia. One of these surveys was conducted on Upper Three Runs Creek and its tributaries on the Savannah River Site, Aiken County, South Carolina; another was conducted on the Lake Jocassee catchment in Oconee County, South Carolina, and Transylvania County, North Carolina; and the third sur- vey was conducted at Spring Creek in Craw- ford County, Georgia. The methodology and results of the studies at Upper Three Runs 275 Creek and Lake Jocassee catchment have been published elsewhere (Morse et al. 1980, 1989), while only partial results of the Spring Creek study have been published (Roths- child et al. 1986, Hamilton and Holzenthal 1986). The survey at Spring Creek was con- ducted from April to October 1983 and con- sisted of a Malaise trap operated continu- ously and supplemented by ultraviolet light trap collections two to three times each month. A number of additional specimens housed in the Clemson University Arthro- pod Collection (CUAC), Department of Entomology, were also examined, most no- tably those collected during a previous study at the type locality of P. car/soni (Carlson 1971). Specimens were examined under a Wild® M8 stereomicroscope. Measurements were taken using an ocular micrometer calibrated at 20 and are presented as a range fol- lowed by the number of specimens mea- sured. External structures were described from uncleared specimens examined in 80% ethanol, while internal structures were de- scribed from abdomens cleared in hot 10% potassium hydroxide and examined in glyc- erin. Terminology of the wings follows that presented by Hamilton (1972) and Ross et al. (1982), while that of the female genitalia follows Nielsen (1980). Because so few data have been published on P. carlsoni, fore- wing lengths and flight periods of both gen- ders are included below. All specimens ex- amined were preserved in 80% ethanol and deposited in the CUAC. RESULTS AND DISCUSSION The males of three species in the confusus group were collected during the survey around Lake Issaqueena. These males were P. blicklei, P. carlsoni, and Polycentropus confusus Hagen. Collected females belong- ing to the confusus group keyed to either P. confusus or P. maculatus when using the key provided by Ross (1944). Females of P. con- usus were easily distinguished using the fig- ure in Ross (1944, fig. 257) and are not con- 276 sidered further. Examination of the genitalia of those females which keyed to P. macu- latus revealed two distinct forms, both of which were different from P. maculatus fe- males collected during the Lake Jocassee catchment survey. The assignment of one of these two forms to P. blicklei was accom- plished by comparison of the genitalia with those of females previously collected with P. blicklei males at localities where no males of either P. carlsoni or P. maculatus were collected. The studies at Upper Three Runs Creek and Spring Creek provided such sites, and females from those localities had been previously identified tentatively as P. blick- lei by S. W. Hamilton and one of us (JCM). Females in the remaining group of macu- latus-like individuals were therefore con- cluded to be females of P. carlsoni. Females of P. blicklei, P. carlsoni, and P. maculatus can be distinguished from those of other species in the confusus group by the elon- gate, parallel to subparallel, internal parts of gonopods VIII (Figs. 1, 5, 9), which are visible in uncleared specimens through ven- ter VIII. Polycentropus blicklei Ross and Yamamoto, 1965 Figs. 1-4 Coloration (in alcohol).—Eyes purple, glazed; dorsum of head, prothorax, meso- thorax, and tegulae brown with erect brown to pale yellow setae; antennae, mouthparts, remainder of thorax, and legs brown to pale yellow, femora, tibiae, and tarsi with brown setae; abdominal sclerites light brown, membrane dull white. Wing membranes and veins light brown, with scattered brown se- tae; forewing with s-m, m, basal fork of M, m-cu, and terminal end of P clear-white, forewing margin lacking setae in spots, ap- pearing mottled; hind wing with s-m, basal fork of M, and m-cu clear-white. Forewing length.—Females 5.1-8.6 mm (n = 70). Female genitalia (Figs. |4).— Venter VIII with ventral plates (v.pl.) narrow, blade-like, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tapering apically (Fig. 1); external parts of gonopods VIII darkly sclerotized in narrow crescent anteriorly (e.gon.VIIIb), transpar- ent posteriorly (e.gon. VIIa) with posterior margin triangular and rounded mesally; lat- eral margins comprised of portions of seg- ment IX (IXb + IXc), darkly sclerotized to ¥4 length. Internal vaginal apparatus mostly comprised of “cushion” (Fig. 3) involving internal parts of gonopods VIII (i.gon. VIII), external parts of gonopods IX (e.gon.IX), and internal portion of segment X (Xd) (Nielsen 1980). Internal parts of gonopods VIII (i.gon. VIII) dark, nearly parallel, ellip- tical, visible through venter VIII (Fig. 1); anterior ends oblique, extending to darkly sclerotized external part of gonopods VIII (e.gon. VIIIb) at basal third of ventral plates (Fig. 1) and to base of internal portion of segment X (Fig. 3); posterior ends acute, falling distinctly short of apices of ventral plates (Fig. 1); ventral surfaces longitudi- nally wrinkled. External parts of gonopods IX (e.gon.IX) weakly sclerotized, not fused with internal portion of segment X (Fig. 3). Internal portion of segment X (Xd) extend- ing anteriad only to anterior margin of ter- gum IX (Nielsen 1980, fig. 22, d.[X), with sclerotized lateral plates widely separated, each with rounded transverse medial pro- jection extending nearly to midline (Fig. 2). Anterior part of genital chamber (g.ch.a.) sclerotized, semicircular, attached by pos- tero-dorsal membranes to antero-ventral edge of internal portion of segment X (Fig. 3, Xd); trough-like with concavity extend- ing full length of postero-ventral surface. Processus spermathecae (p.sp.) ovoid with clear central elevation bearing opening of ductus spermathecae (op.dt.sp.) (Fig. 4), ventral and lateral margins enclosed by an- terior part of genital chamber. Diagnosis.—Females of P. blicklei are distinguishable externally from those of both P. carlsoni and P. maculatus by the exten- sion of the internal parts of gonopods VIII (visible through venter VIII) only to the bas- al third of the ventral plates of venter VIII VOLUME 92, NUMBER 2 and from those of P. car/soni by the darkly sclerotized lateral edges of venter VIII. In- ternally, the short, widely separated lateral plates of the internal portion of segment X and the internal parts of gonopods VIII being as long as these plates are both diagnostic. Notes. — Females were collected from 12 April to 8 November both at ultraviolet lights and in Malaise traps. Material examined.—GEORGIA: Craw- ford County: Spring Creek, approx. 5 mi SSE of Roberta, 7-12.v.1983, 1 2; same data, 11.v.1983, 1 2; same data, 21-28.v.1983, 2 2; same data, 10.vi.1983, 1 2; same data, 11-30.vi.1983, 1 9; same data, 8.1x.1983, 1 9; same data, 20.x.1983, 3 2; RHODE IS- LAND: Richmond, 20.vi.1971, 2 2; same data, 30.vi.1971, 1 ¢; SOUTH CAROLI- NA: Aiken County: Savannah River Plant, Upper Three Runs Creek, 12.iv.1977, 1 9; same data, 8.vul.1977, 1 2; Anderson Coun- ty: Pendleton, Aldwood, |.v.1976, 1 2; Pen- dleton, Tanglewood Spring, springbrook and disturbed site, 225 m el., 30.iv.1988; 6 2; same data, 9.v.1988, 5 2; same data, 17.vi.1987, 1 2; same data, 30.viii.1987, 1 2; same data, 3.xi1.1987, 1 2: Oconee County: Salem, Burgess Creek, 28.1x.1969, 1 9; Pick- ens County: Clemson University Experi- mental Forest surrounding Lake Issa- queena, sites 1-5 and an unnamed creek, 13.1v.1968, 1 2; same data, 14.iv.1968, 1 2; same data, 17.1v.1968, 1 9°; same data, 20.iv.1968, 3 2; same data, 23.iv.1968, 1 @; same data, 24.iv.1968, 2 9; same data, 24.iv.-1.v.1988, 1 2; same data, 26.1v.1968, 3 2; same data, 27.1v.1968, | 2; same data, 6.v.1968, 1 9; same data, 8-15.v.1988, 2 2; same data, 9.v.1968, 1 2; same data, 15- 22.v.1988, 1 9; same data, 18.v.1968, 1 2; same data, 21.v.1988, 2 9; same data, 22.v.1968, 1 9; same data, 22.v.1988; 2 2; same data, 23.v.1968, 1 9; same data, 27.v.1968, 1 9; same data, 28.v.1968, 1 2; same data, 30.v.1968, 1 2; same data, 30.v.- 5.vi.1988, 1 9; same data, 5—12.vi.1988, 3 2; same data, 20-27.v1.1988, 1 2; same data, 2.1x.1968, 1 9; same data, 19.1x.1968, 1 9; 277 same data, 21.1x.1968, 1 2; same data, 26.x.— 8.x1.1988, 1 2; Keowee-Toxaway State Park, unnamed creek, 255 m el., 19.vi.1988, 2 2. Polycentropus carlsoni Morse, 1971 Figs. 5-8 Coloration (in alcohol).—Eyes purple, glazed; dorsum of head, prothorax, meso- thorax, and tegulae brown with erect brown to pale yellow setae; antennae, mouthparts, remainder of thorax, and legs brown to pale yellow, femora, tibiae, and tarsi with brown setae; abdominal sclerites light brown, membrane dull white. Wing membranes and veins light brown, with scattered brown se- tae; forewing with s-m, m, basal fork of M, m-cu, and terminal end of P clear-white, forewing margin lacking setae in spots, ap- pearing mottled; hind wing with s-m, basal fork of M, and m-cu clear-white. Forewing length.— Males 4.7-6.0 mm (n = 20), females 4.5-7.3 mm (n = 35). Female genitalia (Figs. 5-8).— Venter VIII with ventral plates narrow, blade-like, ta- pering apically (Fig. 5); external parts of gonopods VIII darkly sclerotized in narrow crescent anteriorly, transparent posteriorly with posterior margin triangular and round- ed mesally; lateral margins not conspicu- ously darkened. Internal parts of gonopods VIII dark, parallel, rectangular; anterior ends oblique, extending to darkly sclerotized part of external gonopods VIII at basal fourth of ventral plates (Fig. 5) and to basal third of internal portion of segment X (Fig. 7); pos- terior ends oblique, falling distinctly short of apices of ventral plates (Fig. 5); ventral surfaces longitudinally wrinkled, some- times with groove extending entire length of surface. External parts of gonopods IX fused with internal portion of segment X (Fig. 7). Internal portion of segment X ex- tending anteriad only slightly into segment VII, nearly solid sclerite dorsally except for inconspicuous triangular opening medially (Fig. 6), anterior margin darkly rebordered and narrowly cleft medially, posterior mar- gin inconspicuous and notched medially. 278 Sclerotized anterior part of genital chamber attached by postero-dorsal membranes to antero-ventral edge of internal portion of segment X (Fig. 7). Processus spermathecae ovoid with clear central elevation bearing opening of ductus spermathecae (Fig. 8), ventral and lateral margins enclosed by an- terior part of genital chamber. Diagnosis.—Females of P. carlsoni are distinguished externally from those of P. blicklei by the extension of the internal parts of gonopods VIII to the basal fourth of the ventral plates of venter VIII and from those of P. maculatus by the unpigmented lateral margins of venter VIII. Internally, the fol- lowing characters are diagnostic: the darkly rebordered anterior margin and narrow me- dial cleft of the internal portion of segment X, the extension of the internal portion of segment X to the anterior margin of tergum VIII, and (like P. maculatus) the extension of the internal parts of gonopods VIII only to the anterior third of the internal portion of segment X. Notes.—Males were captured from 20 April to 26 October, while females were captured from 15 April to 8 November. All specimens examined were captured in Mal- aise or modified emergence traps on first to second order streams in mixed hardwood- pine forests at elevations between 215 and 245 meters. The only collections of this species at ultraviolet lights were of two males in Alabama (Lago and Harris 1987). Material examined.—SOUTH CARO- LINA: Anderson County: Pendleton, Tan- glewood Spring, springbrook, 17- 24.v1.1987, 1 9; same data, 15—22.vii.1987, 1 2; Pickens County: Clemson University Experimental Forest surrounding Lake Is- saqueena, sites 1-5, 15.iv.1968, 1 9; same data, 20.iv.1968, 2 4; same data, 24.iv.1968, 1 9; same data, 24.iv.-1.v.1988, 1 6; same data, 12.v.1968, 1 9; same data, 15- 22.v.1988, 1 2; same data, 22-30.v.1988, 1 6; same data, 30.v.—5.vi.1988, 4 2; same data, 5-12.vi.1988, 3 6 and 4 2; same data, 12- 20.v1.1988, 6 6 and 2 2; same data, 20- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 27.v1.1988, 1 6 and 1 9; same data, 27.vi-— 4.vii.1988, 2 6 and 3 2; same data, 4— 11.vii. 1988, 3 2; same data, 11-18.vii.1988, 1 ¢éand 5 2; same data, 18-25.vi1.1988, 1 9; same data, 27.vii.—l.vili.1988, 1 92; same data, 8-15.vii.1988, 1 9; same data, 15- 22.viii.1988, 1 6; same data, 31.viii.— 18.ix.1988, 1 4, same data, 18—26.1x.1988, 1 2; same data, 6-13.x.1988, 1 2; same data, 13-26.x.1988, 1 éand 1 2; same data, 26.x.— 8.xi.1988, 1 9. Polycentropus maculatus Banks, 1908 Figs. 9-12 Coloration (in alcohol).—Eyes purple, glazed; dorsum of head, prothorax, meso- thorax, and tegulae brown with erect brown to pale yellow setae; antennae, mouthparts, remainder of thorax, and legs brown to pale yellow, femora, tibiae, and tarsi with brown setae; abdominal sclerites light brown, membrane dull white. Wing membranes and veins light brown, with scattered brown se- tae; forewing with s-m, m, basal fork of M, m-cu, and terminal end of P clear-white, forewing margin lacking setae in spots, ap- pearing mottled; hind wing with s-m, basal fork of M, and m-cu clear-white. Forewing length.—Females 5.3-8.5 mm (n = 15). Female genitalia (Figs. 9-12).— Venter VIII with ventral plates narrow, blade-like, tapering apically (Fig. 9); external parts of gonopods VIII darkly sclerotized in narrow crescent anteriorly, transparent posteriorly with posterior margin triangular and round- ed mesally; lateral margins darkly sclero- tized to ¥4 length. Internal parts of gonopods VIII dark, subparallel, rectangular; anterior ends oblique, extending to darkly sclero- tized part of external gonopods VIII at basal fourth of ventral plates (Fig. 9) and to basal third of internal portion of segment X (Fig. 11); posterior ends oblique, extending near- ly to apices of ventral plates (Fig. 9); ventral surfaces longitudinally wrinkled, with groove extending entire length of surface. External parts of gonopods IX fused with VOLUME 92, NUMBER 2 279 e.gon.Villa é 3 IXb+IXc Xd cS e.gon.IX #4 i.gon.VIll i.gon.VIll pr.sp. g.ch.a. op.dt.sp. r———— - I 1 0.20 mm 4 : 6 0.25 mm Figs. 1-12. Female genitalia of Polycentropus spp. P. blicklei: 1, venter VIII with right ventral plate removed, ventral view. 2, internal portion of segment X, dorsal view. 3, internal genitalia, left lateral view. 4, processus spermathecae and anterior part of genital chamber, caudo-ventral view. P. carlsoni: 5, venter VIII with nght ventral plate removed, ventral view. 6, internal portion of segment X, dorsal view. 7, internal genitalia, left lateral view. 8, processus spermathecae and anterior part of genital chamber, caudo-ventral view. P. maculatus: 9, venter VIII with right ventral plate removed, ventral view. 10, internal portion of segment X, dorsal view. 11, internal genitalia, left lateral view. 12, processus spermathecae and anterior part of genital chamber, caudo- ventral view. v.pl. = ventral plates, e.gon. VIIa and e.gon.VIIIb = external parts of gonopods VIII, i.gon. VII = internal parts of gonopods VIII, IXb + IXc = portions of segment IX, e.gon.IX = external parts of gonopods IX, Xd = internal portion of segment X, g.ch.a. = anterior part of genital chamber, op.dt.sp. = opening of ductus spermathecae, pr.sp. = processus spermathecae. 280 internal portion of segment X and forming conspicuous twisted flange anteriorly (Figs. 10, 11). Internal portion of segment X ex- tending anteriad halfway through segment VII and fused medially (Fig. 10), anterior margin inconspicuous and narrowly cleft medially, postero-medial margin recurved anteriorly and forming pocket. Sclerotized anterior part of genital chamber attached by postero-dorsal membranes to antero-ven- tral edge of internal portion of segment X (Fig. 1 1). Processus spermathecae ovoid with clear central elevation bearing opening of ductus spermathecae (Fig. 12), ventral and lateral margins enclosed by anterior part of genital chamber. Diagnosis.— Females of P. maculatus are distinguished externally from those of P. blicklei by the extension of the internal parts of gonopods VIII to the basal fourth of the ventral plates of venter VIII and from those of P. carlsoni by the darkly pigmented lat- eral margins of venter VIII. Internally, the following characters are diagnostic: the deep, narrow, nonrebordered antero-medial cleft of the internal portion of segment X; the extension of the internal portion of segment X anteriorly to the middle of abdominal segment VII; the conspicuous, twisted an- tero-lateral flanges of the internal portion of segment X; and (like P. carlsoni) the exten- sion of the internal parts of gonopods VIII only to the anterior third of the internal portion of segment X. Notes.— Females were collected from 18 May to 15 September at ultraviolet lights and in Malaise traps. Material examined.—GEORGIA: Union County: Vogel State Park, Wolf Creek, 22.vii.1972, 1 9; NORTH CAROLINA: Transylvania County: Bearcamp Creek at 420 m el., 20-21.vii.1987, 2 2; SOUTH CAROLINA: Oconee County: Coley Creek at 420 mel., 20-21.vii.1987, 2 2; same data, 14-15.1x.1987, 3 2; E. Fork Chattooga Riv- er, U.S. Fish Hatchery, 13.vii.1969, 2 2; Thompson River at NC border, ca. 420 m el., 18-19.v.1987, 1 9; same data, 15- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 16.v1.1987, 1 2; Pickens County: Rocky Bottom, Eastatoe Creek, 5.vii.1969, 3 9; Ta- ble Rock, Carrick Creek, 10.vi1.1969, 1 °. The similarity of the females of P. blicklei, P. carlsoni, and P. maculatus reinforces the close relationships between these species al- luded to in the original descriptions of the males of both P. blicklei and P. carlsoni. Ross and Yamamoto (1965) compared and contrasted the male genitalia of P. blicklei with those of P. maculatus, and Morse (1971) stated that males of P. car/soni most closely resembled those of P. blicklei. How- ever, characters of the female genitalia point to a closer relationship between P. car/soni and P. maculatus than between either of these and P. blicklei. The long, medially- fused internal portion of segment X; the deep, narrow antero-medial cleft of this structure; and the long rectangular internal parts of gonopods VIII appear to be homo- logues unique to the former two species. Thus, characters of the female genitalia may well prove useful in constructing a phylog- eny for the confusus group and may add significantly to phylogenetic analyses of oth- er New World species groups of Polycen- tropus as well. ACKNOWLEDGMENTS We thank Steven W. Hamilton (Austin Peay State University, Tennessee) for both information on the Spring Creek study and advice on morphological terminology, and Steve C. Harris (University of Alabama, Tuscaloosa) for information regarding the Alabama collection localities of P. carlsoni. Gratitude is extended to Joseph D. Culin, John A. DuRant (both of Clemson Univer- sity), and one anonymous reviewer for their helpful comments on this manuscript. This study was funded by a grant from the U.S. Fish and Wildlife Service and administered by the South Carolina Wildlife and Marine Resources Department, and this support is gratefully acknowledged. This is Technical Contribution No. 2981 of the South Caro- VOLUME 92, NUMBER 2 lina Agricultural Experiment Station, Clem- son University. LITERATURE CITED Banks, N. 1908. Some Trichoptera, and allied insects, from Newfoundland. Psyche 15: 61-67. Carlson, P. H. 1971. Emergence and seasonal distri- bution of Ephemeroptera from Wildcat Creek, Pickens County, South Carolina. M.S. thesis, Clemson Univ., Clemson, South Carolina. Hamilton, K. G. A. 1972. The insect wing, part III. Venation of the orders. J. Kans. Entomol. Soc. 45: 145-162. Hamilton, S. W. 1986. Systematics and biogeography of the New World Polycentropus sensu stricto (Tn- choptera: Polycentropodidae). Ph.D. dissertation, Clemson Univ., Clemson, South Carolina. Hamilton, S. W. and R. W. Holzenthal. 1986. Two new species of caddisflies from Georgia (Trichop- tera: Polycentropodidae, Hydroptilidae). Proc. Entomol. Soc. Wash. 88: 163-166. Lago, P. K. and S. C. Harris. 1987. An annotated list of the Curvipalpia (Trichoptera) of Alabama. Entomol. News 98: 255-262. Morse, J.C. 1971. New caddisflies (Trichoptera) from the southeastern United States. J. Ga. Entomol. Soc. 6: 77-84. Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey. 1980. Aquatic insects of Upper Three 281 Runs Creek, Savannah River Plant, South Caro- lina. Part I: Orders other than Diptera. J. Ga. Ento- mol. Soc. 15: 73-101. Morse, J. C., S. W. Hamilton, and K. M. Hoffman. 1989. Aquatic insects of Lake Jocassee catchment in North and South Carolina, with descriptions of four new species of caddisflies (Trichoptera). J. Elisha Mitchell Sci. Soc. 105: 14-33. Nielsen, A. 1980. A comparative study of the genital segments and the genital chamber in female Tn- choptera. Biol. Skr. Udgivet K. Dan. Vidensk. Selsk. 23: 1-200. Ross, H. H. 1944. The caddis flies or Trichoptera of Illinois. Ill. Nat. Hist. Surv. Bull. 23: 1-326. 1947. Descriptions and records of North American Trichoptera, with synoptic notes. Trans. Am. Entomol. Soc. 73: 125-168. Ross, H. H., C. A. Ross, and J. R. P. Ross. 1982. A textbook of entomology, 4th ed. Wiley, New York. Ross, H. H. and T. Yamamoto. 1965. New species of caddisfly genus Polycentropus from eastern North America (Trichoptera, Psychomyiidae). Proc. Biol. Sci. Wash. 78: 241-246. Rothschild, M., R. W. Holzenthal, and S. W. Hamil- ton. 1986. The diversity and phenology of Tn- choptera from Spring Creek, Georgia. Bull. North Am. Benthol. Soc. 3(2): 68 (abstr.). Townes, H. 1972. A light-weight Malaise trap. Ento- mol. News 83: 239-247. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 282-285 NEW SUBFAMILY PLACEMENT FOR SOME NORTH AMERICAN EULOPHIDAE (HYMENOPTERA, CHALCIDOIDEA) JOHN LASALLE AND MICHAEL E. SCHAUFF (JL) CAB International Institute of Entomology, 56 Queen’s Gate, London SW7 SJR, UK; (MES) Systematic Entomology Laboratory, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560. Abstract.—The tetrastichine genus Parachrysocharis Girault (Eulophidae), previously treated in North American literature as a member of the subfamily Entedoninae, does not occur in North America. The one North American species that was included in this genus, P. semiflava Girault, is transferred to the tetrastichine genus Chaenotetrastichus Graham. This is the first record of this genus from North America. The genus Apterolophus Gahan, previously placed in the Eulophinae (Elachertini), is transferred to the Tetrastichi- nae and synonymized with Tetrastichomyia Girault. Tetrastichomyia orygiae Girault is a junior synonym of 7. clisiocampae (Ashmead) and 7. si/vensis is reinstated into Te- trastichomyia from Tetrastichus. Key Words: rastichus In the course of independent studies in the family Eulophidae, we realized that cer- tain North American genera have been tra- ditionally misunderstood, either as to their proper identity, their subfamily placement, or both. This paper arose from the need to correct two such mistakes in order to facil- itate the delimitation of eulophid subfam- ilies necessary for future work. Genus Chaenotetrastichus Graham Chaenotetrastichus Graham, 1987: 25. Type species: Tetrastichus grangeri Erdés (original designation). The genus Parachrysocharis, described in the Entedoninae by Girault (1913), recently was characterized and discussed by Boucek (1988), who transferred it to the Tetrastichi- nae. It presently contains only the type species, P. javensis Girault (1913), previ- ously treated under the name Tetrastichus first record, Parachrysocharis, Apterolophus, Tetrastichomyia, Chaenotet- pyrillae Crawford, which is a parasite of Pyrilla eggs in southern Asia. Girault (1917) described a North American species in the genus Parachrysocharis, P. semiflava. This species clearly does not belong in the genus Parachrysocharis, which is very distinct due to the unique longitudinal striations on the mid lobe of the mesoscutum (see Boucek 1988). Girault’s North American species has been treated in the Entedoninae in all cat- alogues since its description (Peck 1951, 1963, Burks 1979). Examination of the types of P. semiflava showed us that it actually belongs in the recently described tetrastich- ine genus Chaenotetrastichus Graham (1987). This genus was known previously only from Europe. The genus Chaenotetrastichus was de- scribed based on a single European species, Tetrastichus grangeri Erdés. Diagnostic characters for the genus, given in the text and in the generic key presented in the same work (Graham 1987), are as follows: sub- VOLUME 92, NUMBER 2 marginal vein with | dorsal seta; scutellum with 5-6 pairs of setae; mandible with a long, falcate outer tooth and two very small, closely approximated inner teeth; dorsal surface of thorax dull, with raised reticu- lation; mid lobe of mesoscutum without a median line, with 2—3 irregular rows of long, erect setae on each side; dorsal surface of gaster wholly, finely reticulate; body bright- ly metallic green to blue-green. Parachrysocharis semiflava Girault be- longs in the genus Chaenotetrastichus, al- though it differs in several key characters (based on grangeri). The scutellum only possesses 3-4 pairs of setae; the mesoscu- tum only has | row of 3-4 setae along each lateral margin (in both species the setae on the mesoscutum and scutellum are long, whitish, and semi-erect); the gaster is dis- tinctly reticulate on the first tergum, but only faintly reticulate after that; the body is me- tallic green dorsally, yellow ventrally. Oth- erwise, it matches the diagnosis of Chaeno- tetrastichus well, particularly the peculiar mandible shape, which is unique to this ge- nus, the single seta on the submarginal vein, and the distinct reticulation of the thorax. Chaenotetrastichus presently contains two species: C. grangeri (Erdés). Tetrastichus grangeri Erdés, 1958 (1957): 286-287. Holotype 2, FRANCE, Chartrettes, 11.vi.1950, C. Granger. C. semiflavus (Girault), New ComBINa- TION. Parachrysocharis semiflava Girault, 1917: 129. Lectotype ° (present designa- tion), TEXAS, Austin, 16.vi1.1909, C. Hartmann [USNM type no. 20803, ex- amined]. The lectotype 2 of C. semiflavus is mount- ed on a point and has been labelled as lec- totype. The USNM collection also contains a female paralectotype with the same data as the lectotype and a slide on which Girault mounted a head and several pieces of a body from a third specimen. The rest of this spec- imen is missing. Genus Tetrastichomyia Girault Tetrastichomyia Girault, 1916a: 48. Type species: Miotropis clisiocampae Ashmead, 1894 (original designation). Apterolophus Gahan, 1919: 3-4. NEw SYNONYM. Type species: Apterolophus pulchricornis Gahan (original designation). The type species of Apterolophus was set through original designation, not monotypy as stated in North American catalogues (Peck 1951, 1963, Burks 1979). Girault (1916a) described the genus Ter- rastichomyia based on the single species Miotropis clisiocampae Ashmead. He later described two more species in this genus, 7. orgyiae and T. silvensis (Girault 1916b). These three species were subsequently as- signed to Miotropis Thomson by Peck (1951) when he synonymized Tetrastichomyia un- der Miotropis and later were transferred to Syntomosphyrum (Burks 1967, 1979). Gra- ham (1987) provided a key to European genera of Tetrastichinae, in which he res- urrected the genus Tetrastichomyia, and presented the following diagnostic charac- ters: dorsellum divided medially by a groove or ridge; propodeum with a sharp carina on the callus, raised reticulation with rugosity or wrinkles, and a small spiracle; third anel- lus larger than the preceding two and setose (at least in European and North American species); scutellum without submedian lines, sublateral lines deep with lateral edge car- inate; vertex with transverse ridge posterior to ocellar triangle; lower edge of antennal toruli level with ventral edge of eyes; mid lobe of mesoscutum without medial line; frons without transverse suture. Gahan (1919) described the genus Apter- olophus in the subfamily Elachertinae (now considered the tribe Elachertini of the Eu- lophinae). It has since been maintained in Eulophinae, even though at the time of its description Gahan mentioned its close sim- 284 ilarity to Miotropis clisiocampae Ashmead, which has since been transferred to the Tet- rastichinae. Our studies indicate that Ap- terolophus is indeed a tetrastichine, and we are assigning it to this subfamily, where we are synonymizing it with Tetrastichomyia, a genus based on the species M. clisiocam- pae. Examination of Apterolophus pulchricor- nis Gahan, the type species and only in- cluded species in the genus Apterolophus Gahan, shows that it agrees with all the key characters given by Graham. It differs from other species in Tetrastichomyia in that the female is brachypterous. The male is un- known. At present there are three North Ameri- can species placed in the genus Tetrasticho- myia: T. clisiocampae (Ashmead). Miotropis cli- siocampae Ashmead, 1894: 341. Lecto- type 2 (present designation), WEST VIR- GINIA, Morgantown, 28.v1.1891, A. D. Hopkins, ex. Clisiocampe americana on apple [USNM type no. 2183, examined]. As the type of Tetrastichomyia, this com- bination was revived through implication by Graham (1987: 28) when he resur- rected the genus Jetrastichomyia. T. orgyiae Girault, 1916b: 112. New SyNonyM. Tetrastichomyia orgyiae Girault. Holotype °, WASHING- TON, D.C., x1.1915, R. M. Fouts, ex. Orgyia leucostigma [USNM type no. 20399, examined]. T. orgyiazele Burks, 1979: 1005. NEw SynonyM. Unnecessary replacement name. Burks (1979) assigned Synto- mosphyrum orgyiazele as a replace- ment name for Jetrastichomyia or- gylae Girault (nec Syntomosphyrum orgyiae Ashmead) when he trans- ferred 7. orgyiae Girault to Synto- mosphyrum. T. pulchricornis (Gahan), NEw COoMBINA- TION. Apterolophus pulchricornis Gahan, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1919: 3-4. Holotype 2, NEW YORK, Leeds, vi.1918, W. M. Mann [USNM type no. 21910, examined]. T. silvensis Girault, REVIVED COMBINA- TION. Tetrastichomyia silvensis Girault, 1916: 111. Holotype 2, MARYLAND, Glenndale, 16.vui.1915 [USNM type no. 20398, examined]. This species was de- scribed in 1916, in the same paper as or- gylae; however, the date of publication has been incorrectly given as 1919 in North American catalogues (Peck 1951, 1963, Burks 1979). The lectotype of clisiocampae Ashmead is point mounted on a pin with three other specimens (paralectotypes) of the same species. The lectotype point has been marked with a black dot to indicate the proper spec- imen and a lectotype label has been added to the pin. ACKNOWLEDGMENTS We thank E. E. Grissell, J. Heraty, J. Huber, G. A. P. Gibson, and T. Henry for reviewing the manuscript and for their com- ments. LITERATURE CITED Ashmead, W. H. 1894. Description of new parasitic Hymenoptera. Trans. Am. Entomol. Soc. 21: 318- 344. Boucek, Z. 1988. Australasian Chalcidoidea (Hy- menoptera). CAB International, Wallingford. 832 pp. Burks, B. D. 1967. Superfamily Chalcidoidea, pp. 213-282. In Krombein, D. V. and B. D. Burks, eds., Hymenoptera of America North of Mexico. Synoptic Catalog. Second Supplement. USDA, Agriculture Monographs No. 2. 584 pp. . 1979. Family Eulophidae, pp. 967-1022. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds., Catalog of Hymenoptera in America North of Mexico. Vol. I. Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, D.C. 1198 pp. Erdos, J. 1958. Eulophidae novae gallicae (Eulophi- dae nouveaux de France) [Hym.]. Bull Soc. Ento- mol. France 62(1957): 279-287. Gahan, A. B. 1919. A new genus of chalcid-wasp VOLUME 92, NUMBER 2 belonging to the family Eulophidae. Proc. Ento- mol. Soc. Wash. 21: 24. Girault, A. A. 1913. Australian Hymenoptera Chal- cidoidea—IV. The family Eulophidae with de- scriptions of new genera and species. Mem. QsInd. Mus. 2: 140-296. —. 19l6a. Descriptions of miscellaneous North American chalcidoid Hymenoptera of the family Eulophidae. Proc. U.S. Natl. Mus. 51: 39-52. 1916b. New species of parasitic Hymenop- tera. Bull. Brooklyn Entomol. Soc. 11: 111-113. 1917. The occurrence of the genus Parach- rysocharis Girault in the United States. Can. Ento- mol. 49: 129. 285 Graham, M. W.R. de V. 1987. A reclassification of the European Tetrastichinae (Hymenoptera, Eu- lophidae), with a revision of certain genera. Bull. Brit. Mus. (Nat. Hist.), Entomol. Ser. 55(1): 1- 392. Peck, O. 1951. Superfamily Chalcidoidea, pp. 410- 594. In Muesebeck, C. F. W., K. V. Krombein, and H. K. Townes, eds., Hymenoptera of America North of Mexico. Synoptic Catalog. U.S.D.A., Agric. Monograph 2: 1-1420. 1963. A catalogue of the Nearctic Chalci- doidea (Insecta: Hymenoptera). Can. Entomol., Suppl. 30: 1-1092. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 286-303 DESCRIPTIONS OF A NEW SPECIES AND THREE INCOMPLETELY KNOWN SPECIES OF WESTERN NEARCTIC [SOPERLA (PLECOPTERA: PERLODIDAE) RICHARD L. BoTToRFF, STANLEY W. SzCZYTKO, AND ALLEN W. KNIGHT (RLB, AWK) Department of Land, Air, and Water Resources, University of California, Davis, California 95616; (SWS) College of Natural Resources, University of Wisconsin, Stevens Point, Wisconsin 54481. Abstract.—The male, female, nymph, and egg of Jsoperla miwok, new species, are described from a Sierra Nevada intermittent stream, California. Distinguishing features include male aedeagal shape and spination, female subgenital plate shape, adult and nymphal pigment patterns, and microstructure of the collarless egg. This new species is placed in the Jsoperla sobria (Hagen) complex. The male aedeagus, female, nymph, and egg of Isoperla acula Jewett are described, and this species is moved to the /soperla quinquepunctata (Banks) complex. Males have a unique patch of sclerotized scales encir- cling the aedeagus near the apex. Other features include the triangular subgenital plate of the female, the pigment pattern of the nymph, and the collarless egg. Jsoperla acula is closely related to Jsoperla mormona Banks, but differs by inhabiting small intermittent streams. Nymphs of Jsoperla adunca Jewett and Isoperla bifurcata Szczytko and Stewart are described, and both species are retained in the /soperla sordida Banks complex. Isoperla adunca nymphs are unique among western Nearctic Jsoperla species by lacking longitu- dinal abdominal stripes. Key Words: plexes Twenty-two species of /soperla are cur- rently known from the western Nearctic re- gion (Stark et al. 1986), and relationships within this group have been elucidated in recent revisions by Szczytko and Stewart (1979, 1984). Using morphological char- acters of adults, nymphs, and eggs, they erected five species complexes for 20 Jso- perla species, and two species remained un- assigned. Placement of some species within these complexes was tentative because cer- tain life stages were unknown. Presently, one male, one female, ten nymphs, and two eggs of the 22 Jsoperla species are unknown or incompletely described. Szczytko and Stew- Plecoptera, Perlodidae, /soperla, new species, western Nearctic species com- art (1979) have stressed the desirability of completing these life stage descriptions for a better understanding of phylogenetic re- lationships in this group. Further descrip- tions of several Jsoperla species endemic to California are especially needed because this region 1s important in the group’s evolution, past dispersal, and zoogeography. During a study of stoneflies in the central Sierra Nevada of California, we collected and associated many adults and nymphs of Isoperla. In the process, a new species was discovered, plus the undescribed life stages of three rarely collected species were col- lected, Jsoperla acula Jewett (male aedea- VOLUME 92, NUMBER 2 gus, female adult, nymph, and egg), [soperla adunca Jewett (nymph), and Jsoperla bifur- cata Szczytko and Stewart (nymph). The objectives of this paper were to (1) describe all life stages of a new Jsoperla species from California, (2) complete the descriptions of all life stages of J. acula, I. adunca, and IJ. bifurcata, and (3) relate these new data to the five /soperla species com- plexes. MATERIALS AND METHODS This study was based on collections of Isoperla nymphs and adults from many habitats in the Cosumnes River basin (sea level to 2249 m elevation) on the western slope of the central Sierra Nevada, Califor- nia. Positive nymph/adult associations of all four species were made by rearing mature nymphs to emergence in the field and/or laboratory using small growth containers. Cool water temperatures (4—1 2°C) were nec- essary to maintain J. bifurcata nymphs until they emerged, but the other three species collected from low elevation habitats emerged successfully at much warmer water temperatures (15-29°C). Teneral adults were held in the laboratory to obtain eggs. Adult and nymphal drawings were made with a Wild MS-A stereo dissecting micro- scope and camera lucida. The microstruc- tures of nymphal mouthparts, terga, and eggs were examined using compound and scan- ning electron microscopes (SEM). Eggs ovi- posited into holding containers or dissected from preserved gravid females were pre- pared for SEM study as described by Szczyt- ko and Stewart (1979), and micrographs were made with an ISI Super III SEM. Male terminalia were treated for study according to the methods of Szczytko and Stewart (1979). Aedeagal armatures were examined from temporarily mounted sections in glyc- erol, and were studied with a Zeiss Standard RA, Routine and Research compound mi- croscope. To facilitate study of the males, the aedeagus of living specimens was everted just prior to preservation. 287 Voucher specimens of all four /soperla have been deposited in the National Mu- seum of Natural History (USNM), Wash- ington, D.C.; California Academy of Sci- ences (CAS), San Francisco, Calif.; and Brigham Young University, Provo, Utah. Additional specimens are in the collections of S. W. Szczytko and R. L. Bottorff. RESULTS AND DISCUSSION Isoperla miwok Bottorff and Szczytko, NEw SPECIES Male.—Macropterous. Body length 8-10 mm, forewing length 7.5-9.5 mm, slightly exceeding abdomen. General body color light to medium brown. Dorsum of head creamy yellow, dark band connecting me- dian and lateral ocelli; interocellar area light; light spot anterior to median ocellus: occi- put light brown behind lateral ocelli, with reticulations (Fig. 1). Antennae brown, ped- icel and scape margins dark. Pronotum with light median stripe, disks medium brown, rugosities dark, anterior and posterior mar- gins dark, anterolateral corners light (Fig. 1). Meso-metanota with light median stripe or spot anteriorly. Wings light brown. Fem- ora with light-dark distal bands. Abdominal terga with two mesal longitudinal rows of dots. Vesicle absent. Paraprocts pointed, deflected outward at tips, recurved over posterior margin of tenth tergum, slightly crenulated and bearing short setae (Figs. 2, 3). Aedeagus membranous with expanded balloon-like apical section bearing 2 small rounded lobes and a long, narrow, tail-like, anteromedian tube (difficult to see and evert in preserved specimens) covered with fine rounded spinulae (Fig. 4B) and a postero- median truncated lobe void of spinulae (Fig. 4); posteroapical patch of large, heavy, red- dish brown spines (Fig. 4C); mesal area cov- ered with stout, evenly spaced, proximally projecting, golden brown spinulae (Fig. 4A) and posterior band of large, heavy, proxi- mally projecting, reddish brown spines which grade into smaller, lighter spines 288 proximally (Fig. 4D); proximal area with fine shallow scales bearing microtrichia and small fine spinulae (Fig. 4E). Cercal seg- ments with a long posteroventral seta. Female.— Macropterous. Body length 10- 12 mm; forewing length 9-11 mm, slightly exceeding abdomen. Body color and exter- nal morphology similar to male. Subgenital plate truncate, wide at base, produced at least 2 length over 9th sternum; posterior ’; dark brown, posterior margin evenly rounded; scattered long fine hairs mesally (Fig. 5). Nymph.—Body length of mature nymph 10-14 mm. General body color medium brown. Dorsum of head with a dark wide lateral band between ocelli and antennal bases, which extends anterolaterally and en- closes a small light spot; interocellar area light; triangular light area anterior to me- dian ocellus connecting to transverse light band across frontoclypeus, posterior cor- ners of triangle extending as thin light lines to antennal bases; epicranial suture light; occiput bearing irregular row of short spi- nulae (Fig. 6). Lacinia triangular, bidentate: 1 axillary seta; 8-9 long marginal setae be- low subapical tooth (1 thin seta at tooth base, then 7-8 equally-spaced stout setae); 2 long stout submarginal setae below base of main tooth; sparse marginal and sub- marginal fine setae extending to lacinia base (Figs. 7, 8). Mandibles with 6 teeth, outer 3 teeth serrated; wide ventral patch of long setae extending between base of outer tooth and mandibular base, inner mandibular surface with row of long stout marginal setae (Figs. 9, 10); mandibles with brush of stout setae from base of inner teeth to marginal setal row (left mandible brush dense and medium length, right mandible brush sparse and short) (Figs. 11, 12). Antennae 80-100% of body length, 60-69 segments. Pronotal median stripe, lateral margins, and rugosi- ties light; disks brown; margins fringed with short to long setae; angles rounded (Fig. 6). Legs with a dorsal fringe of long fine hairs; dark band distally on femora, proximally PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON on tibia. Abdominal terga with 3 longitu- dinal brown stripes, | mesal and 2 lateral; anterior and posterior margins dark; 8 lon- gitudinal rows of dots, 2 mesal and 3 each laterally (Fig. 6). Cerci 70-80% of body length; 28-31 segments, each with posterior whorl of short setae, and one long dorsal and ventral seta; complete dorsal fringe of long hairs after 17th segment. Egg.—Length 400-450 um; width 300- 350 um. General shape a prolate spheroid, cross section circular (Fig. 13). Color white. Collar and eclosion line absent (Fig. 13). Chorion covered with irregularly rounded to hexagonal follicle cell impressions (FCI’s); FCI walls thick, raised; FCI floors flat with 3-5 medium-sized aeropyles (Fig. 14). Mi- cropylar row subequatorial; orifices with small lips, positioned on FCI floors and walls, some associated with rosettes of 4—5 FCI’s (Fig. 14). Distribution. — This species 1s known only from the Sierra Nevada foothills, Califor- nia. Types. — Holotype male, allotype female, and three paratype nymphs collected from California, El Dorado Co., Indian Creek, 3.3 km NE of Michigan Bar bridge, 13-IV- 1987, R. L. Bottorff, deposited in the National Museum of Natural History, Washington, D.C. Paratypes (R. L. Bottorff, collector).— California: Amador Co.: Little Indian Creek, 3 km W of Plymouth, | fe- male 18-IV-1986, | female 9-V-1986. El Dorado Co.: Indian Creek, 3.3 km NE of Michigan Bar bridge, 97 nymphs 25-III- 1987, 21 males, 4 females, 79 nymphs, 6 males & | female lab-reared 8-IV-1987, 2 males, 3 females lab-reared 9-IV-1987, 2 males, 4 females lab-reared 1 1-IV-1987, 17 males, 8 females, 49 nymphs 13-IV-1987, 2 males, 4 females lab-reared 20-IV-1987, 1 male, 2 females lab-reared 23-IV-1987, 1 nymph 10-III-1988; unnamed creek tribu- tary to N bank of Cosumnes River, 2.9 km upstream of Michigan Bar bridge, 4 males, 5 females, 2 nymphs 9-IV-1986, 1 female lab-reared, | nymph 10-IV-1986, 2 males VOLUME 92, NUMBER 2 lab-reared 1 1-IV-1986, 2 females lab-reared 15-IV-1986; unnamed creek tributary to N bank of N Cosumnes River, 6 km N Nash- ville, 1 exuvium 1-V-1987. Sacramento Co.: Burgoyne Creek, 1.3 km NE of Michigan Bar bridge, 1 male, 1 female 9-IV-1986, 1 female lab-reared 10-IV-1986, 1 female lab- reared 22-IV-1986, 1 nymph 25-III-1987; Cosumnes River at Michigan Bar, | nymph 27-I-1982, 2 nymphs 5-III-1982, | nymph 23-IV-1982, 1 male 22-III-1984, 1 male 14- IV-1984, 1 female 22-IV-1984, 1 male, 1 female 11-IV-1986, 1 male 18-IV-1986, 1 male, | female 29-IV-1986, 1 male lab- reared 31-III-1986, 1 male lab-reared 5-IV- 1986, 1 male & 1 female lab-reared, | nymph 11-IV-1986; Cosumnes River at Slough- house, | nymph 5-III-1982; unnamed creek tributary to S bank of Cosumnes River, 0.3 km upstream of Michigan Bar bridge, 2 males 29-III-1986, | male lab-reared, | nymph 30-III-1986, 1 female lab-reared 3 1- III-1986, 1 nymph 25-III-1987. Paratypes are in the collections of the California Acad- emy of Sciences, Brigham Young Univer- sity, S. W. Szcezytko and R. L. Bottorff. Etymology.—This species is named in honor of the Miwok tribe of California In- dians, whose tribal area includes the type locality. Biological notes.—/soperla miwok pri- marily occurred in small intermittent streams at lower elevations (20-350 m) of the Sierra Nevada foothills, California. Only a few nymphs were found in nearby larger perennial streams despite extensive search- ing in these habitats. When this species emerged in March and April, the intermit- tent streams had low flow and warm water (23-29°C), causing nymphs to exhibit a “push-up” behavior to aid oxygen uptake. Although microhabitats of coarse substrate and fast current were present, mature nymphs usually were collected from pools which had aquatic macrophytes and a silty substrate. Nymphs held in the laboratory several days or weeks at room temperatures (20-25°C) and without water current nor- 289 mally emerged successfully. Emergence started in late March, reached a peak in mid April, and finished by early May, soon after which the stream habitat completely dried. Other Plecoptera found in these intermit- tent streams included, Jsoperla acula, I. adunca, Oemopteryx vanduzeea (Claassen), Suwaillia (new species), Sweltsa californica (Jewett), and more rarely, Cosuwmnoperla hypocrena Szezytko and Bottorff. [soperla miwok emerged several weeks before J. acu- la and I. adunca. Diagnosis. —/soperla miwok is placed in the /soperla sobria (Hagen) complex, which has three other western Nearctic species (Szezytko and Stewart 1979): I. gravitans (Needham and Claassen), J. sobria, and I. tilasqua Szczytko and Stewart. It shares the following characteristics with these species: (1) a large body size, (2) male aedeagus membraneous, tubular, and bearing patches of small stout spinulae and longer hair-like spinulae, (3) male vesicle reduced or absent, (4) female subgenital plate truncate or broadly rounded, wide at base, and (5) egg chorion with evenly spaced aeropyles. The male aedeagus of J. miwok is most similar to I. tilasqua because both have a long apical tube(s). However, J. miwok can be separated from all members of this species complex by the pigment patterns of adults and nymphs, the shape and spinule pattern of the male aedeagus, the single long tube on the aedeagus, the shape and pigment of the female subgenital plate, and the collarless egg. The egg of J. miwok is most similar to J. acula because both lack a collar and have raised FCI walls, but can be distinguished by its larger overall size, fewer aeropyles in the FCI floors, micropyles with small lips, and some micropyles positioned in FCI floors. Within this species complex, only /. mi- wok and I. sobria occur in California. Both species were found in the same major drain- age basin, but were separated by stream type and elevation. Jsoperla miwok occurred in 4 5 Figs. 1-5. Jsoperla miwok adults. 1, Head and pronotum. 2, Male terminalia, dorsal. 3, Male paraproct, lateral. 4, Aedeagus, lateral; A, mesal band of short, stout golden brown spinulae; B, fine rounded spinulae on anteromedian tail-like tube; C, posteroapical patch of large, heavy reddish brown spines; D, posteromesal patch of large, heavy, proximally projecting reddish brown spines; E, proximal patch of fine, shallow scales with microtrichia and fine spinulae. 5, Female subgenital plate, ventral. Scale lines: 1 = 1 mm; 2, 4, and 5 = 0.5 mm; 3 = 0.2 mm; 4A, B, C, D, and E = 25 um. 290 VOLUME 92, NUMBER 2 291 roy iF 7 Py bi, a “2 Fig. 6. Jsoperla miwok, mature nymph, habitus; scale line = 1 mm. low elevation, intermittent, small streams, while J. sobria occurred in medium or high elevation, perennial, small streams. Jsoper- Isoperla acula Jewett 1962: 18. Holotype la miwok coexisted with J. acula and I. male; Fresno Co., California (CAS). adunca, which are in different species com- /soperla acula Mlies 1966: 393. plexes, emerged later, and have different J/soperla acula Szezytko and Stewart 1979: drumming calls (Bottorff et al. in press). Ti. Isoperla acula Jewett 292 Male.—Jewett (1962) and Szczytko and Stewart (1979) have described the male ex- ternal morphology of J. acula. Aedeagus with narrow apical tubular section with sclero- tized patch of flat scales which are finely divided anteriorly (Fig. 15B), apex with a small nipple, spinulae absent (Fig. 15); large anteromedian lobe with narrow anterior band of small, fine spinulae (Fig. 15A); mesal section void of spinulae; area below mesal section covered with stout, evenly spaced, proximally projecting, golden brown spi- nulae (Fig. 15C); proximal area with fine, shallow scales bearing microtrichia (Fig. 15D). Female.—Macropterous. Body length 10- 13 mm; forewing length | 1-12.5 mm. Body color and external morphology similar to male. Subgenital plate triangular, produced posteriorly 2 length over 9th sternum; apex variable (normally narrowly rounded, but some broadly rounded, pointed, or rarely notched); mesal patch of long fine setae (Fig. 16). Nymph.—Body length of mature nymph 10-14 mm. General body color medium brown, covered with dark clothing hairs. Dorsum of head with strongly contrasting pigment pattern (Fig. 17); dark lateral bands extend from ocelli to antennal bases, then anterolaterally; triangular light spot anterior to median ocellus connecting to thin trans- verse light band across frontoclypeus; in- terocellar area light; posterior margin of head dark brown, with dark curved bands ex- tending anteriorly to lateral ocelli; stem of epicranial suture light; large irregular light area between each compound eye and lat- eral ocellus; occiput bearing sinuous row of short spinulae (Fig. 17). Lacinia triangular, bidentate; 1 axillary seta; 20-25 marginal setae below subapical tooth (1 thin seta at tooth base, then 6-8 long equally-spaced stout setae, then | 2-16 smaller setae); 3 long stout submarginal setae below base of main tooth, then a narrow continuous band of fine submarginal setae to lacinia base (Figs. 18, 19). Mandibles with 6 teeth, most ser- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rated; wide ventral patch of long setae ex- tending between base of outer tooth and mandibular base, inner mandibular surface with row of long stout marginal setae (Figs. 20, 21); mandibles with brush of stout setae from base of inner teeth to marginal setal row (left mandible brush dense and medium length, right mandible brush sparse and short) (Figs. 22, 23). Pronotum with median stripe, disk stripes, and lateral margins light; anterior and posterior margins dark; mar- gins fringed with short and occasional long setae; angles rounded (Fig. 17). Meso-meta- nota each with 4 dark pointed bars extend- ing posteriorly from anterior margin toward 2 isolated dark bars (Fig. 17). Thoracic ster- na with numerous chloride cells (Fig. 26). Legs with a dorsal fringe of long fine hairs; tibia with proximal dark spot. Abdominal terga with 3 longitudinal dark brown bands, median band narrow, lateral bands wide and flared anteriorly and posteriorly (Fig. 17); anterior margin dark; posterior fringe of medium-long setae and scattered interca- lary spinulae (Figs. 17, 27, 28). Cercal seg- ments with posterior whorl of short setae; complete dorsal fringe of long hairs after 17th segment. Egg.—Length 350-370 um; width 230- 260 um. General shape a prolate spheroid, cross section circular (Fig. 24). Color cream. Collar and eclosion line absent (Fig. 24). Chorion covered with irregularly rounded to pear-shaped FCI’s; FCI walls thick, raised; FCI floors flat and finely punctate with 18- 28 small aeropyles (Fig. 25). Micropyles subequatorial, associated with rosettes of 4— 5 EFGEs(Figs 25): Distribution. —This species is known only from the Sierra Nevada foothills, Califor- nia. Material examined.—California (R. L. Bottorff, collector, except where noted): Amador Co.: Big Indian Creek, 6 km N of Plymouth, | male 13-VI to 9-VIII-1982, R. Fouch, 7 nymphs 11-IV-1986, | female lab- reared 22-IV-1986, 1 female lab-reared 24- IV-1986, 2 females, 2 nymphs 25-IV-1986, VOLUME 92, NUMBER 2 2 males, 2 females lab-reared 28-IV-1986, 1 male, 3 females 9-V-1986, | male lab- reared 12-V-1986, 2 males lab-reared 14- V-1986, 3 females lab-reared 16-V-1986, 1 female lab-reared 17-V-1986, 1 female lab- reared 26-V-1986, 1 female 30-V-1986, 3 nymphs 25-III-1987, 1 female lab-reared, 7 nymphs 1-V-1987; Little Indian Creek, 3 km W of Plymouth, 3 nymphs 18-IV-1986, 11 nymphs 25-IV-1986, | male, 2 females 29-IV-1986, 1 female 9-V-1986, 2 males, | female 30-V-1986, 24 nymphs 25-III-1987, 18 nymphs 1-V-1987, 6 males, 12 females lab-reared 4-V-1987, 8 males, 9 females, | nymph 12-V-1987. Butte Co.: 9 mi. N Oro- ville, 1 male, 5 nymphs 24-IV-1955, S. W. Hitchcock (USNM). El Dorado Co.: Acorn Creek, 6 km S Pilot Hill, 8 nymphs 3-IV- 1988: Cooper Canyon, 3 km W Pilot Hill, 26 nymphs 2-IV-1988; Deadman Creek, 3.8 km SE of El Dorado, 6 nymphs 1-V-1987; Knickerbocker Creek, 5 km NW Pilot Hill, 35 nymphs 6-II-1988; Skunk Canyon, 6 km S Pilot Hill, 1 nymph 3-IV-1988; Sweet- water Creek, 10 km S Pilot Hill, 20 nymphs 3-IV-1988: unnamed creek tributary to N bank of N Cosumnes River, 6 km N Nash- ville, 5 nymphs 1-V-1987, 90 nymphs 12- V-1987; unnamed creek tributary to Fol- som Lake at Rattlesnake Bar, 6 km SW Pilot Hill, 2 nymphs 2-IV-1988. Fresno Co.: Dry Creek, 7 mi. NE of Academy, | male (Ho- lotype, CAS) 19-IV-1955, D. L. Abell. Sac- ramento Co.: Cosumnes River at Michigan Bar, 1 nymph 5-III-1982, 1 nymph 23-IV- 1982, 1 male 31-V-1982. Biological notes. —/soperla acula primar- ily occurred in small intermittent streams at lower elevations of the Sierra Nevada foothills, California (see Biological Notes for I. miwok). Only a few nymphs were found in nearby larger perennial streams. It co- existed with J. adunca and I. miwok, both members of other Jsoperla species com- plexes. Emergence started in mid April, reached a peak in early May, and finished by late May. Diagnosis.—The phylogenetic relation- 293 ships of J. acu/a have remained unclear since the original description of the male because of the incomplete aedeagal description, the scarcity of specimens, and the undescribed female, nymph, and egg. Szczytko and Stew- art (1979) tentatively placed /. acula in the I. sordida complex based on the male pig- ment pattern and Jewett’s (1962) partial de- scription of an aedeagal sclerotized struc- ture. It is now clear that J. acu/a males have a band of sclerotized scales which encircle the aedeagus apex, but lack the distinctive sclerotized process which projects from the membraneous aedeagus of all species in the I. sordida complex. Isoperla acula should be included in the I. quinquepunctata (Banks) complex, which includes four other western Nearctic species: I. jewetti Szczytko and Stewart, J. /ongiseta Banks, /. mormona Banks, and J. quinque- punctata (Szezytko and Stewart 1979). It shares the following characteristics with these species: (1) 9th and/or 10th male ab- dominal terga with patches of stout hairs or spinulae, (2) a row of occipital spinulae on nymphal head, (3) a dorsal fringe of long hairs on nymphal legs, and (4) three longi- tudinal stripes on abdominal terga of nymphs. Tsoperla acula is closely related to J. mor- mona based on similarities in the shape and spination of the male aedeagus, in the bi- partite patch of spinulae on the male 9th tergum, and in the pigment patterns of the adult and nymphal head-pronotum. Nymphs will key to /. mormona in Szczytko and Stewart (1979). However, J. acu/a can be distinguished from this species by (1) a band of sclerotized scales on the aedeagus, (2) a male vesicle wider than long, (3) male paraprocts long and thin, (4) a triangular female subgenital plate which is produced over sternum 9, (5) larger sized nymphs and adults, and (6) eggs lacking a collar, but with distinct FCI walls. Isoperla mormona occurs throughout the western Nearctic region, while /. acula is restricted to California (Szczytko and Stew- 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-14. mandible, ventral. 10, Left mandible, ventral. 11, Detail of mght mandible, ventral. 12, Detail of left mandible, ventral. 13, Egg. 14, Detail of egg chorion and micropyles. Scale lines: 7, 8, 9, and 10 = 0.2 mm; 11, 12, and 13 = 0.1 mm; 14 = 20 um. art 1979). Both species occur at low eleva- tions in the Sierra Nevada, but emerge at different times and inhabit distinctly differ- ent stream types. Jsoperla acula inhabited small intermittent streams, while J. mor- mona inhabited large perennial rivers. The morphological similarity of the two species suggests a recent divergence, possibly as- sociated with the drier climates and increas- ingly intermittent flow conditions in small streams of the Sierra Nevada foothills fol- lowing the Pleistocene epoch. Most stone- flies in this region emerge prior to the sum- mer warming of streams; however, /. mormona 1s one of the last to emerge, often when water temperatures exceed 20°C in June-July. The ability of J. mormona nymphs to cope with warm water temper- atures in perennial streams may have pre- adapted variants for life in nearby inter- ew PR, wae! Isoperla miwok nymph and egg. 7, Right maxilla, ventral. 8, Left maxilla, ventral. 9, Right mittent streams and led to the recent evolution of 7. acula. The male drumming calls of these two morphologically-similar species are distinctly different in beat num- ber and interval, suggesting that drumming behavior has diverged faster than morpho- logical traits and has been an important iso- lating factor (Bottorff et al. in press). Isoperla adunca Jewett Isoperla adunca Jewett 1962: 19. Holotype male, allotype female; Santa Clara Co., California (CAS). Isoperla adunca Mies 1966: 393. Tsoperla adunca Szczytko and Stewart 1979: 80. Nymph.—Body length of mature nymph 9-12 mm. General body color uniform me- dium brown. Dorsum of head with a square VOLUME 92, NUMBER 2 15 Figs. 15-16. 16 Isoperla acula adults. 15, Aedeagus, lateral; A, anteromedian band of small fine spinulae; B, sclerotized patch of flat scales; C, patch of stout, evenly spaced, proximally projecting golden brown spinulae; D, proximal patch of fine, shallow scales with microtrichia. 16, Female subgenital plate, ventral. Scale lines: 15 and 16 = 0.5 mm; 15A, B, C, and D = 25 um. light spot anterior to median ocellus, spot margined laterally by medium brown an- teriorly-pointed lobes; light lines extend be- tween lobes and antennal bases; small light spot anterior to lateral ocelli; frontoclypeus with a light transverse band; center of in- terocellar area light; occiput with reticula- tions and an irregular row of short spinulae which is interrupted medially (Fig. 29). La- cinia quadrate, bidentate, small gap be- tween bases of main and subapical teeth; | axillary seta; definite marginal shelf below subapical tooth with 10-12 stout setae (6- 7 long, 4-5 shorter); 8-9 long stout sub- marginal setae below main tooth, first 3 in gap; scattered marginal and submarginal fine setae extending to lacinia base (Figs. 30, 31). Mandibles with 6 teeth, most serrated; nar- row ventral row of long setae extending be- tween base of outer tooth and mandibular base, inner mandibular surface with row of long stout marginal setae (Figs. 32, 33); mandibles with brush of stout setae from base of inner teeth to marginal setal row (left mandible brush dense and medium length, right mandible brush sparse and short) (Figs. 34, 35). Antennae light; margin of scape dark. Pronotum uniform brown, lateral margins light; rugosities darker than disks; margins fringed with short and occasional long setae; angles rounded (Fig. 29). Meso- metanota brown, with a few light reticula- tions. Thoracic sterna with mesal sclera lacking dense hairs; membranes with chlo- ride cells (Fig. 37). Legs with a dorsal fringe of long fine hairs. Abdominal terga uniform brown, becoming lighter near pleura, each segment with scattered intercalary spinulae and a posterior fringe of medium length se- tae (Fig. 36). Cercal segments with a pos- 296 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = a di Figs 17. terior whorl of short setae and | long dorsal and ventral seta; sparse intrasegmental se- tae, continuous dorsal fringe absent. Distribution. — This species is known only from the Sierra Nevada foothills and the Coast Range, California. Material examined.—California (R. L. Bottorff, collector): Amador Co.: Big Indian Creek, 6 km N of Plymouth, 3 nymphs 25- IV-1986, 1 female, 3 nymphs 9-V-1986, 1 Tsoperla acula, mature nymph, habitus; scale line = 1 mm. female lab-raised 10-V-1986, 1 male, | fe- male lab-raised 12-V-1986, 1 female lab- raised 13-V-1986, 4 males, 6 females, 1 nymph 30-V-1986, 2 nymphs 1-V-1987; Little Indian Creek, 3 km W of Plymouth, 10 nymphs 18-IV-1986, 1 male, 5 nymphs 25-IV-1986, 1 male lab-reared, 2 nymphs 27-IV-1986, 1 male, 2 females 9-V-1986, 2 males, 6 females, 5 nymphs 30-V-1986, 2 males, | female lab-reared 31-V-1986, 5 VOLUME 92, NUMBER 2 Figs. 18-28. nymphs 25-III-1987, 3 males, 4 females 1-V-1987, 13 males, 8 females, 62 nymphs 12-V-1987. El Dorado Co.: Acorn Creek, 6 km §S Pilot Hill, 1 nymph 3-IV-1988; Coo- per Canyon, 3 km W Pilot Hill, 7 nymphs 2-IV-1988; Indian Creek, 3.3 km NE of Michigan Bar bridge, 34 nymphs 25-III- 1987, 50 nymphs 8-IV-1987, | male, 48 nymphs 13-IV-1987, 1 male, 4 nymphs 21- 297 Isoperla acula nymph and egg. 18, Right lacinia, ventral. 19, Left lacinia, ventral. 20, Right mandible, ventral. 21, Left mandible, ventral. 22, Detail of ght mandible, ventral. 23, Detail of left mandible, ventral. 24, Egg. 25, Detail of egg chorion and micropyles. 26, Chloride cells on thoracic sterna. 27, Abdominal terga. 28, Detail of abdominal tergum. Scale lines: 18, 19, 20, 21, and 27 = 0.2 mm; 22, 23, and 24 = 0.1 mm; 25, 26, and 28 = 20 um. IV-1987, 6 nymphs 10-III-1988; unnamed N. bank tributary to Cosumnes River, 2.9 km upstream of Michigan Bar bridge, 16 nymphs 9-IV-1986; unnamed creek tribu- tary to N bank of N Cosumnes River, 6 km N Nashville, 6 nymphs 1-V-1987, 53 nymphs 12-V-1987; unnamed creek tribu- tary to Folsom Lake at Rattlesnake Bar, 6 km SW Pilot Hill, 13 nymphs 2-IV-1988. 298 Sacramento Co.: Burgoyne Creek, N bank tributary to Cosumnes River, 1.3 km up- stream of Michigan Bar bridge, 25 males, 25 females, 27 nymphs 9-IV-1986, | male, 1 female lab-raised 10-IV-1986, 1 female 1-V-1986, 6 nymphs 25-III-1987; Cos- umnes River at Michigan Bar, 7 nymphs 5-ITI-1982, 30 nymphs 23-IV-1982, 3 nymphs 21-VI-1982; Cosumnes River at Sloughhouse, | nymph 24-IV-1982; un- named S bank tributary to Cosumnes River 0.3 km upstream of the Michigan Bar bridge, 13 nymphs 21-III-1986, 18 nymphs 29-III- 1986, 1 female lab-reared 2-IV-1986, 4 males, 4 females, 10 nymphs, | male lab- reared 11-IV-1986, 17 males, 13 females 29-IV-1986, 15 nymphs 25-III-1987; un- named S bank tributary Cosumnes River 0.7 km downstream of the Michigan Bar bridge, 2 nymphs 14-III-1986, 1 female lab- reared 26-III-1986, 1 male lab-reared 28- III-1986. Biological notes.—/soperla adunca pri- marily occurred in small intermittent streams at lower elevations of the Sierra Ne- vada foothills, California (see Biological Notes for J. miwok). Only a few nymphs were found in nearby larger perennial streams. Emergence usually started in late March and extended through April-May, but varied somewhat in different intermit- tent streams and years as influenced by water temperature and flow duration. Diagnosis. — Based on the nymphal mor- phology described in this study, J. adunca is retained in the J. sordida complex. All nymphs in this group have a pronotal fringe of short stout hairs and occasional longer hairs (Szczytko and Stewart 1979). Addi- tionally, most nymphs in this complex lack a continuous dorsal fringe of long cercal hairs. Based on adult characters, Szczytko and Stewart (1979) found J. adunca most closely related to J. denningi Jewett; how- ever, the nymph of J. denningi is unknown. Szczytko and Stewart (1979) stated that most Isoperlinae nymphs can be separated from the Perlodinae by the presence of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 29. Isoperla adunca, mature nymph, head and pronotum; scale line = | mm. nymphal abdominal stripes. In contrast to all other western Nearctic /soperla, I. adunca nymphs are unique in lacking longitudinal abdominal stripes (some eastern Nearctic species also lack abdominal stripes, includ- ing J. burksi Frison, I. marlynia (Needham and Claassen), and J. signata (Banks)). J[s- operla adunca nymphs also differed from the other /soperla in this study in their mouthpart setation and shape. The lacinia of I. adunca has a broad marginal shelf and numerous stout submarginal setae, while their mandibles have a narrow row of ven- tral setae. These differences were unexpect- ed because Szczytko and Stewart (1979) re- ported little interspecific variation in the nymphal mouthparts of western Nearctic Tsoperla. In the central Sierra Nevada of California, I. adunca and I. bifurcata, members of the same species complex, occurred in the same major drainage basin, but they emerged at different times and inhabited different stream types. Jsoperla adunca occurred in VOLUME 92, NUMBER 2 Figs. 30-37. ventral. 33, Left mandible, ventral. 34, Detail of right mandible, ventral. 35, Detail of left mandible, ventral. 36, Abdominal terga. 37, Chloride cells on thoracic sterna. Scale lines: 30, 31, 32, 33, and 36 = 0.2 mm; 34 and 35 = 0.1 mm; 37 = 20 um. low elevation, intermittent, small streams, while J. bifurcata occurred in medium or high elevation, perennial, headwater springs. Tsoperla adunca coexisted with I. acula and I. miwok, which are in different species complexes and have different drumming calls (Bottorff et al. in press). Tsoperla bifurcata Szczytko and Stewart Isoperla sordida Gaufin et al. 1966: 71. Tsoperla sordida Gaufin et al. 1972: 119. Tsoperla sordida Baumann et al. 1977: 152. Tsoperla_ bifurcata Szczytko and Stewart 1979: 80. Holotype male, allotype female; Union Co., Oregon (USNM). Nymph.—Body length of mature nymph 10-13 mm. General body color medium 299 Tsoperla adunca nymph. 30, Right lacinia, ventral. 31, Left lacinia, ventral. 32, Right mandible, brown, with a median light stripe extending longitudinally along entire dorsum, mem- branes of fresh specimens reddish and densely covered with chloride cells. Dorsum of head with a dark brown square-shaped band connecting ocelli, dark triangular patches extend posteriorly over occiput; center of interocellar area light; frontocly- peus with a light transverse band; light M- shaped line anterior to median ocellus; large oval light spot posterior to and small light spot anterior to lateral ocelli; occiput with an irregular row of short spinulae (Fig. 38). Lacinia triangular, bidentate; | axillary seta; 12-16 marginal setae below subapical tooth (1 fine seta at subapical tooth base, then 5- 6 long equally-spaced stout setae, then 6-9 smaller setae); 3 long stout submarginal se- tae below base of apical tooth; sparse row of fine marginal and submarginal setae ex- tending to lacinia base (Figs. 40, 41). Man- dibles with 6 teeth, most serrated; wide ven- tral patch of long setae extending between base of outer tooth and mandibular base, inner mandibular surface with row of long stout marginal setae (Figs. 42, 43); mandi- 300 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ iv \ ‘ N \ \\ \ N a cuiituantnariaay Seat iy hide eS eS ; 1a 5 i/ J mm e S a e a — Fig. 38. Jsoperla bifurcata, mature nymph, habitus; scale line = | mm. bles with brush of stout setae from base of inner teeth to marginal setal row (left man- dible brush dense and medium length, right mandible brush sparse and short) (Figs. 44, 45). Pronotum with light median stripe bor- dered by dark brown stripes, lateral margins and rugosities light, disks light brown, an- terior and posterior margins dark; margins VOLUME 92, NUMBER 2 Fig. 39. scale line = | mm. Isoperla bifurcata nymph, thoracic sterna; fringed with short stout setae and occasional long setae; angles rounded (Fig. 38). Meso- metanota with light median stripe and re- ticulations. Meso-metasterna with distinct mesal sclera with dark clothing hairs and patches of stout spinulae (Fig. 39). Legs with sparse dorsal fringe of fine hairs. Abdominal terga with longitudinal median light band and two lateral dark brown bands (Fig. 38); numerous long and short intercalary spi- nulae (Figs. 46, 47); posterior fringe of me- dium length setae. Cercal segments with a posterior whorl of short setae and longer ventral seta; sparse intrasegmental setae, continuous dorsal fringe absent. 301 Distribution. — This species is known from California, Idaho, Oregon, and Washing- ton. Material examined.—California (R. L. Bottorff, collector): El Dorado Co.: Single- ton Springs at headwaters of N Cosumnes River, 25 km E of Grizzly Flat, 3 nymphs 6-VIII-1980, 2 nymphs 15-II-1981, 5 nymphs 27-IIJ-1981, 21 nymphs 24-IV- 1981, 10 nymphs 31-V-1981, 25 nymphs 2-X-1981, 15 nymphs 9-III-1982, 37 nymphs 20-IV-1982, 18 nymphs 17-V- 1982, 15 nymphs 2-VII-1982; N Cosumnes River, 30-100 m downstream of Singleton Springs, 39 nymphs 6-VIII-1980, 1 female, 15 nymphs 5-IX-1980, 18 nymphs 24-IX- 1980, 25 nymphs 31-X-1980, 5 nymphs 26- XII-1980, 6 nymphs 15-H-1981, 20 nymphs 27-III-1981, 6 nymphs 24-IV-1981, 1 nymph 31-V-1981, 109 nymphs 10-VII- 1981, 1 male, 1 female, 7 nymphs 16-VIII- 1981, 1 female in trap 16-VIII to 2-IX-1981, 74 nymphs 2-X-1981, 9 nymphs 15-XII- 1981, 40 nymphs 16-I-1982, 42 nymphs 9-III-1982, 66 nymphs 20-IV-1982, 33 nymphs 17-V-1982, 1 male, | female, 59 nymphs 2-VII-1982, 5 males, 4 females 7-VII-1982, 1 male, 3 females 16-VIH-1982, 1 male, 1 female 2-VIII-1982, 27 nymphs 10-VIII-1982, 1 female 23-VIII-1982, 12 nymphs 22-VI-1987, 1 male field-reared 9- 16-VII-1987, 1 male, 2 females field-reared 16-26-VII-1987, 2 females field-reared 26- VII to 8-VIII-1987; unnamed spring stream tributary to N bank of N Cosumnes River, 2.4 km upstream of Meiss Ranch and 18 km E of Grizzly Flat, 5 nymphs 19-VI-1986, 9 nymphs 30-VI-1986, 15 nymphs 8-VI- 1987, 1 male field-reared 8—22-VI-1987, 2 males, 2 females field-reared 22-VI to 9-VII- 1987; unnamed N bank tributary to N Cos- umnes River, 2.3 km upstream of Meiss Ranch and 18 km E of Grizzly Flat, 1 nymph 14-VI-1986, 1 nymph 18-VI-1986. Biological notes.—/soperla bifurcata oc- curred in small spring-fed streams (order |- 2) at medium or high elevations (> 1500 m) in the Sierra Nevada, California, and nor- Figs. 40-47. mally did not coexist with other /soperla species. Emergence started in mid June, reached a peak in July, and continued into August. A few adults were collected in early September, primarily ovipositing females. Other common stoneflies in these habitats included Soyedina nevadensis (Claassen) and Sweltsa borealis (Banks). Diagnosis.— Based on the nymphal mor- phology described in this study, J. bifurcata is retained in the J. sordida complex because all members have similar pronotal and cer- cal setation. Szczytko and Stewart (1979) found a close relationship between adults of I. bifurcata and I. fusca Needham and Claassen. The nymphs of these two species share a similar pattern of longitudinal ab- dominal stripes and a reduced or absent dorsal fringe of leg hairs. [soperla bifurcata PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tsoperla bifurcata nymph. 40, Right lacinia, ventral. 41, Left lacinia, ventral. 42, Right mandible, ventral. 43, Left mandible, ventral. 44, Detail of mght mandible, ventral. 45, Detail of left mandible, ventral. 46, Abdominal terga. 47, Detail of abdominal tergum. Scale lines: 40, 41, 42, 43, and 46 = 0.2 mm; 44 and 45 = 0.1 mm; 47 = 20 um. nymphs can be distinguished from all other members in this species complex by their pigment pattern, reduced dorsal fringe of leg hairs, and sclerotized thoracic sterna. ACKNOWLEDGMENTS We thank J. A. Stanger for nymph and adult drawings, T. Remnsen and the Great Lakes Research Facility for the use of their SEM and lab, the E. Ruman family of Mich- igan Bar, California, for access to streams on their property, and L. D. Bottorff for assistance with field collections. This study was supported in part by Jastro-Shields Re- search Grants from the University of Cal- ifornia, Davis, and the University of Wis- consin/Stevens Point Faculty Research Fund #5760. VOLUME 92, NUMBER 2 LITERATURE CITED Baumann, R. W., A. R. Gaufin, and R. F. Surdick. 1977. The stoneflies (Plecoptera) of the Rocky Mountains. Mem. Am. Entomol. Soc. 31: 1-208. Bottorff, R. L., S. W. Szczytko, A. W. Knight and J. J. Dimick. Submitted. Drumming behavior of four species of western Nearctic /soperla (Plecoptera: Perlodidae). Ann. Entomol. Soc. Am. Gaufin, A. R., A. V. Nebeker, and J. Sessions. 1966. The stoneflies (Plecoptera) of Utah. Univ. Utah Biol. Ser. 14: 1-89. Gaufin, A. R., W. E. Ricker, M. Miner, P. Milam, and R. A. Hays. 1972. The stoneflies (Plecoptera) of Montana. Trans. Am. Entomol. Soc. 98: 1-161. Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich 82: 1-632. 303 Jewett, S. G., Jr. 1962. New stoneflies and records from the Pacific Coast of the United States. Pan- Pacific Entomol. 38: 15-20. Stark, B. P., S. W. Szczytko, and R. W. Baumann. 1986. North American stoneflies (Plecoptera): Systematics, distribution, and taxonomic refer- ences. Great Basin Nat. 46: 383-397. Szcezytko, S. W. and K. W. Stewart. 1979. The genus Tsoperla (Plecoptera) of western North America; holomorphology and systematics, and a new stonefly genus Cascadoperla. Mem. Am. Entomol. Soc. 32: 1-120. . 1984. Descriptions of Ca/liperla Banks, Rick- era Jewett, and two new western Nearctic /soperla species (Plecoptera: Perlodidae). Ann. Entomol. Soc. Am. 77: 251-263. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 304-305 THE STATUS OF THE GENUS MINEUS STAL, 1862 (HETEROPTERA: PENTATOMIDAE: ASOPINAE) DONALD B. THOMAS, JR. USDA-ARS Screwworm Research, Apartado Postal 544, Tuxtla Gutierrez, Chiapas, Mexico. Abstract.—The synonymy of the genus Mineus Stal is confirmed. Mineus strigipes is placed in the asopine genus Perillus Stal. Mineus triangularis (Walker) is transferred to the pentatomine genus Mormidea Amyot & Serville, where it is the senior synonym of Mormidea kirkaldyi Rolston. Key Words: The genus Mineus was erected by Stal (1867) to hold a single species, Podisus strig- ipes Herrich-Schaefer, 1851. The species oc- curs throughout the eastern United States with records for New Mexico (Ruckes 1937) and Arizona (Froeschner 1988) probable misidentifications of Perillus exaptus (Say). A second species, Strachia triangularis Walker, was transferred to Mineus by Dis- tant (1900). The locality of origin given by Walker (1867) was Eucador. In his revision of the Asopinae, Schouteden (1907) provid- ed a key to the genera which included Mi- neus and an excellent figure of Mineus stri- gipes. Schouteden listed Mineus triangularis with a question mark, and in a footnote he stated that based on Walker’s description the species did not appear to belong in Mi- neus. Nothing further has been published on Mineus triangularis, and in spite of the excellent drawing, Mineus strigipes is often misidentified in collections, being confused with Perillus exaptus. The source of the con- fusion is that Schouteden (1907) and later Torre-Bueno (1938) separate Peri/lus and Mineus by the presence of a subapical spine on the profemur. In Perillus exaptus, how- ever, the spine is reduced to a small tubercle Mineus, Pentatomidae, Asopinae, taxonomy which in some cases is absent altogether. The dorsal markings are similar enough so that someone using Knight’s (1952) revi- sion of Perillus would easily confound the two species. It is generally overlooked (e.g. McPherson 1982, Froeschner 1988) that Hoffman (1971) synonymized Mineus un- der Perillus, an arrangement with which I am in complete accord. The genitalic struc- ture of strigipes is identical to that of the other species of Perillus, there being little interspecific variation in either males or fe- males of the genus. McDonald (1966), who studied the genitalia, also concluded that strigipes belonged in Perillus, but it was not within the scope of his study to make formal nomenclatural changes. The species of Per- illus can be distinguished by the dorsal markings as described in Knight’s (1952) revision and key. Perillus strigipes and Per- illus exaptus can be distinguished because strigipes always has a median, longitudinal stripe of yellow to red color on the midline of the pronotum, which is lacking in ex- aptus. The synonymy of Mineus under Perillus leaves Mineus triangularis without a genus. I recently examined the holotype of Strach- VOLUME 92, NUMBER 2 ia triangularis Walker, which is located in the British Museum of Natural History. The species belongs in the genus Mormidea Amyot & Serville, subgenus Melanochila Stal. The specimen is a female and has an ivory callous traversing the pronotum along the posterior margin of the cicatrices. Also, the specimen has a complete ivory callous along the margins of the scutellum, and a prominent linear ivory callous on the disc of the corium following the embolar suture. The specimen is missing all legs except one which is mounted on a card below it. The tibia is black with a median yellow band. Based on these characters and using the re- vision of Mormidea by Rolston (1978), the species can be placed as Mormidea mon- tandoni Kirkaldy, 1902. Subsequently, however, Rolston (1984) reported that 7. montandoni was misidentified by him and was a senior synonym of M. bridarolli Piran, which has the tibia yellow with black spots. The species identified in his revision by having the black tibia with the median yel- low band was therefore unnamed and he proposed the name Mormidea kirkaldyi. Strachia triangularis, with its new combi- nation Mormidea triangularis (Walker 1867), is therefore a senior synonym of Mormidea kirkaldyi Rolston (1984). The placement of Mineus as a junior syn- onym of Peril/us is thus confirmed and the following synonymy is proposed: 1. Perillus strigipes (Herrich-Schaefer). Podisus strigipes Herrich-Schaefer 1851: 338. Mineus strigipes: Stal 1867: 48. Perillus strigipes: Hoffman 1971: 55. 2. Mormidea triangularis (Walker). Strachia triangularis Walker 1867: 323. Mineus triangularis: Distant 1900: 55. Mormidea kirkaldyi Rolston 1984: 342. New SyYNONYMY. Mormidea triangularis: NEw COMBIN- ATION. 305 ACKNOWLEDGMENTS I am grateful to William R. Dolling, Brit- ish Museum Natural History, for lending the type of Strachia triangularis Walker, and David E. Rider for help locating Distant’s reference to Mineus triangularis. LITERATURE CITED Distant, W. L. 1900. Rhynchotal notes. V. Heter- optera: Asopinae-Tessaratominae. Ann. Mag. Nat. Hist. 7: 55-65. Froeschner, R. C. 1982. Jn Henry, T. H. and R. C. Froeschner, eds., Catalog of the Heteroptera or true bugs of Canada and the continental United States. E. J. Brill Publ., Leiden. Herrich-Schaefer, G. A. 1851. Die Wanzenartigen Insecten, Vol. IX. Nurnberg. Hoffman, R. L. 1971. The Insects of Virginia No. 4. Shield Bugs (Hemiptera; Scutelleroidea: Scuteller- idae, Corimelaenidae, Cydnidae, Pentatomidae). Virginia Polytechnic Inst. Res. Div. Bull. 67. Knight, H. H. 1952. Review of the genus Perillus with description of a new species (Hemiptera, Pen- tatomidae). Ann. Entomol. Soc. Am. 45; 229-232. McDonald, F. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera: Heterop- tera). Quaest. Entomol. 2: 7-150. McPherson, J. E. 1982. The Pentatomoidea (Hemip- tera) of northeastern North America with empha- sis on the fauna of Illinois. Southern Illinois Univ. Press, Carbondale. Rolston, L. H. 1978. A revision of the genus Mor- midea (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 84: 161-219. 1984. New synonymy and a new species in the genus Mormidea (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 92: 342-343. Ruckes, H. 1937. An annotated list of some penta- tomids (Heteroptera) from New Mexico. Bull. Brooklyn Entomol. Soc. 32: 32-36. Schouteden, H. 1907. Heteroptera: Fam. Pentatom- idae: Subfam. Asopinae (Amyoteinae). Jn Wyts- man, P., ed., Genera Insectorum Fasc. 52. Bru- xelles. Stal, C. 1867. Bidrag till Hemipternas Systematik. Ofv. Svenska Veten.-Akad. Forhandl. 24: 491- 560. Torre-Bueno, J. R. de la. 1938. A synopsis of the Hemiptera-Heteroptera of America north of Mex- ico. Part I. Families Scutelleridae, Cydnidae, Pen- tatomidae, Aradidae, Dysodiidae and Temitaphi- didae. Entomol. Amer. 19: 141-304. Walker, F. 1867. Catalogue of the specimens of Het- eropterus-Hemiptera in the collection of the Brit- ish Museum, Vol. 2. London. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 306-309 TELENOMUS (HYMENOPTERA: SCELIONIDAE) EGG PARASITES OF ERINNYIS ELLO (LEPIDOPTERA: SPHINGIDAE) NORMAN F. JOHNSON Department of Entomology, 1735 Neil Avenue, The Ohio State University, Columbus, Ohio 43210. Abstract.— Telenomus dilophonotae Cameron, T. connectans Ashmead, and T. monili- cornis Ashmead (= sphingis auct.) are diagnosed. Telenomus dilophonotae (from Costa Rica, Guyana, and Brazil) has been reared from the eggs of Erinnyis ello (L.) and Perigonia stulta Herrich-Schaffer (Lepidoptera: Sphingidae); 7. connectans (Florida, Dominican Republic, Grenada, Costa Rica, and Brazil) from E. ello, Sphinx merops Boisduval, Xylophanes tersa (L.), and X. neoptolemus (Stoll) (Sphingidae); 7. monilicornis has been reared from E. e//o in Costa Rica and the Dominican Republic. The species Telenomus puticulus Johnson is considered a junior synonym of connectans. Key Words: The moth Erinnyis ello (L.) (Lepidoptera: Sphingidae) is a common Neotropical and southern Nearctic species that often reaches pest status on a number of plants, in par- ticular, manioc (Manihot esculenta) and other Euphorbiaceae. The reported egg parasitoids of EF. ello comprise three species: Telenomus connectans Ashmead, Teleno- mus dilophonotae Cameron, and Teleno- mus monilicornis Ashmead. All have pre- viously been recorded as parasites of various sphingids, including E. e//o, but identifica- tion on the basis of the literature has been impossible. They were described around the turn of the century, and no revision or key to the Te/enomus species of the Neotropical region is available. My objectives are to pro- vide diagnoses of these species to assist in their identification and to present taxonom- ic notes on the species. The morphological terminology follows that outlined in John- son (1984) and Bin and Dessart (1983). The biology of connectans and dilophonotae and the impact upon their host will be discussed parasitic wasps, Telenomus, Erinnyis in a separate paper by Dr. Joao Manuel de Abreu. Telenomus connectans Ashmead Telenomus connectans Ashmead, 1895: 792, 4, 2. Lectotype (examined) in British Mu- seum (Natural History). Liophanurus connectans: Kieffer, 1926: 79. Telenomus connectans: Masner, 1965: 111. Telenomus puticulus Johnson, 1984: 54, 8. Holotype (examined) in Canadian Na- tional Collection of Insects, Arachnids and Nematodes. New SYNONYMY. Diagnosis: Legs, including coxae, and basal antennal segments yellow; occipital carina complete medially; 2 clava 5-merous, claval formula Al 1-A7/1,2,2,2,2; 6 antenna 12-merous; preocellar pit present; T1 with 2-3 pairs of sublateral setae; 4 genitalia with 3 large teeth per digitus, penis valves and laminae volsellares strongly melanized (Fig. 2; see Johnson 1984 for illustrations of habi- tus and female antenna). VOLUME 92, NUMBER 2 Telenomus connectans was earlier re- ported by Gahan (1930) to parasitize the eggs of E. e//lo in the Dominican Republic. Dr. William Haber has also reared this species from the eggs of Sphinx merops Boisduval, Yylophanes tersa (L.), X. neop- tolemus (Stoll) and Perigonia stulta Herrich- Schaffer in Monteverde, Costa Rica. The species Telenomus puticulus, known only from Florida, was described because of the unusual presence of the preocellar pit and multiple pairs of sublateral setae on T1. The first of these structures commonly occurs in the genus Trissolcus Ashmead (all of which are parasites of the eggs of Pentatomomor- pha), but is extremely rare in Te/enomus (see Bin and Dessart 1983). In all respects puticulus seems to correspond with the abundant Neotropical material now avail- able to me. Thus the distribution of 7. con- nectans extends from Brazil through both the Lesser Antilles and Central America to Florida. The structure of the male genitalia indi- cates that connectans belongs to the califor- nicus group of species (large ditigal teeth, laminae volsellares approximated medially, aedeagal lobe short). As I earlier pointed out (Johnson 1984), it is distinguished from practically all of those species by the pre- ocellar pit and sublateral setae. In may re- spects it is quite similar to species of Te/en- omus that parasitize the eggs of Pentatomidae (Hemiptera), not, however, with the New World species of the podisi group, but rather with those of the Old World (e.g. T. chloropus Thomson, T. seychellensis Kieffer, 7. triptus Nixon and 7. cyrus Nix- on). Telenomus dilophonotae Cameron Telenomus dilophonotae Cameron, 1913: 133, 2. Holotype (examined) in British Museum (Natural History). Telenomus dilophonotae: Masner, 113: Diagnosis.—Legs and antennae dark brown, occipital carina complete medially; 1965: 307 2 clava 6-merous (Fig. 3), claval formula (A11-A7/1,2,2,2,2; 6 antenna 10-merous (Fig. 4); preocellar pit absent; T1 with | pair of sublateral setae; 6 genitalia with 4 small teeth per digitus, generally weakly mela- nized throughout (Fig. 1). This species is noteworthy among Te/en- omus in that the female antennal clava is composed of 6 antennomeres (see below), and the male antenna possesses but 10 an- tennomeres. The genus Pseudotelenomus (now considered a synonym of Telenomus) was described by Costa Lima (1928) to con- tain another Brazilian species, P. pachycoris (a reduviid egg parasite) in which the male antenna lacked a segment; in this case the fusion of antennomeres 1s still clearly visi- ble. I can find no indication of fusion in dilophonotae. A5 is the sex segment in te- lenomines generally (see Bin and Vinson 1986) and in dilophonotae this segment is unusually enlarged. Thus reduction of an- tennomeres has occurred distal to this seg- ment. There is some disagreement among workers as to which segments are taken to comprise the clava or club of the female antenna. I am using here the definition out- lined in Johnson (1984): I consider A6 to be a clavomere, not because it is greatly broader than the preceding antennomere, but because its apical surface is excavated and parallels the basal surface of A7. The antennae of this species, particularly those of the male, seem to be rather weakly scler- otized and often collapse when air-dried. Cameron’s original description of diloph- onotae (females only) was based on material reared from Erinnyis (then known as Dil- ophonota) in Guyana; Haber has also reared this species from the eggs of Perigonia stulta and an unidentified sphingid in Costa Rica (Monteverde, and Santa Rosa National Park in Guanacaste Province). Telenomus monilicornis Ashmead Telenomus monilicornis Ashmead, 1894: 203, 6. Holotype (examined) in British Museum (Natural History). 308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1, 2. Male genitalia, ventral view. 1. Telenomus dilophonotae. 2. Telenomus connectans. Scale line = 0.10 mm. Telenomus monilicornis: Kieffer, 1912: 22. of sublateral setae; é genitalia with four small Phanurus monilicornis: Kieffer, 1926: 63. teeth per digitus, weakly melanized, similar Telenomus monilicornis: Johnson, 1983: to T. dilophonotae (for illustration see John- 446. son 1984). Telenomus monilicornis: Johnson, 1984: 11. Telenomus monilicornis (= sphingis auct.) is a widely distributed species, most com- Diagnosis.—Legs and antennae dark monly encountered because it parasitizes the brown; occipital carina broadly interrupted eggs of Manduca spp. in the southeastern medially; ° clava 5-merous; 6 antenna 12- United States. Ashmead (1887) originally merous; preocellar pit absent; T1 with 1 pair described the species Teleas sphingis as a VOLUME 92, NUMBER 2 Figs. 3, 4. Telenomus dilophonotae. 3. Female an- tenna. 4. Male antenna. Scale line = 0.25 mm. parasite of Manduca; however, examina- tion of the holotype proved that this name actually and unfortunately belongs to a species of the crassiclava species group, all parasites of the eggs of Homoptera (Johnson 1984). Thus the correct name for this com- mon species is monilicornis. I have speci- mens reared from the eggs of FE. e//lo from Costa Rica (Puntarenas, 2 km N Cuatro Cruces) and the Dominican Republic (San Cristobal). ACKNOWLEDGMENTS I thank J. M. de Abreu (CEPLAC, Ita- buna, Bahia, Brazil), W. Haber (Monte- verde, Costa Rica), and L. Masner (Ottawa, Ontario) for the opportunity to study their material; N. D. M. Fergusson (London) and P. M. Marsh (Washington) for assistance with the study of the type material of Ash- mead and Cameron; and D. C. Darling (To- ronto) for comments on the manuscript. This material is based upon work supported by the National Science Foundation under Grant No. BSR-8516579. 309 LITERATURE CITED Ashmead, W. H. 1887. Report on insects injurious to garden crops in Florida. Bull. Div. Entomol. U.S. Dept. Agric. 14: 9-29. . 1894. Report on the parasitic Cynipidae, part of the Braconidae, the Ichneumonidae, the Proc- totrypidae, and part of the Chalcidinae. Part III. Zool. J. Linn. Soc. Lond. 25: 188-254. 1895. Report on the parasitic Hymenoptera of the island of Grenada, comprising the families Cynipidae, Ichneumonidae, Braconidae, and Proctotrypidae. Proc. Zool. Soc. London 1895: 742-812. Bin, F. and P. Dessart. 1983. Cephalic pits in Proc- totrupoidea Scelionidae and Ceraphronoidea (Hy- menoptera). Redia 66: 563-575. Bin, F. and S. B. Vinson. 1986. Morphology of the antennal sex-gland in male Trissolcus basalis (Woll.) (Hymenoptera: Scelionidae), and egg para- sitoid of the green stink bug, Nezara viridula (He- miptera: Pentatomidae). Int. J. Ins. Morphol. & Embryol. 15: 129-138. Cameron, P. 1913. The Hymenoptera of the George- town Museum, part v. Timehnri, J. Roy. Agr. Comm. Soc. British Guiana 3: 105-137. Costa Lima, A. da. 1928. Nota sobre o Pseudotelen- omus pachycoris (n. g., N. sp.) parasito dos ovos de Pachycoris torridus (Scop.). Bol. Mus. Nacional 4: 51-53. Gahan, A. B. 1930. Synonymical and descriptive notes on parasitic Hymenoptera. Proc. U.S. Natl. Mus. 77(2831): 1-12. Kieffer, J. J. 1912. Proctotrypidae (3e partie). Jn Andre, E., Species des Hyménoptéres d'Europe et d’Algérie. Vol. 11. Pages 1-160. . 1926. Scelionidae. Das Tierreich. Vol. 48. Walter de Gruyter & Co., Berlin. 885 pp. Johnson, N. F. 1983. Types of Neotropical Telenom- inae described by W. H. Ashmead and P. Cameron (Hymenoptera: Scelionidae). Proc. Entomol. Soc. Wash. 85: 439-449. 1984. Systematics of Nearctic Telenomus: Classification and revisions of the podisi and phy- matae species groups (Hymenoptera: Scelionidae). Bull. Ohio Biol. Surv. 6(3). 113 pp. Masner, L. 1965. The types of Proctotrupoidea (Hy- menoptera) in the British Museum (Natural His- tory) and in the Hope Department of Entomology, Oxford. Bull. Brit. Mus. (Nat. Hist.), Entomol. Suppl. 1. 154 pp. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 310-313 A SURVEY OF THE COCCINELLIDAE (COLEOPTERA) ASSOCIATED WITH NURSERY STOCK IN MARYLAND C. L. STAINES, JR., M. J. ROTHSCHILD, AND R. B. TRUMBULE Maryland Department of Agriculture, Office of Plant Industries and Resource Conser- vation, Plant Protection Section, 50 Harry S Truman Parkway, Annapolis, Maryland 21401. Abstract. — Twenty-eight species of adult Coccinellidae were collected in Maryland nurs- eries from 1986 to 1988. Host plants, presence of prey, and active predation (if any) were recorded for each species. The most commonly detected species were Coccinella septem- punctata L., Coleomegilla maculata lengi Timberlake, and Hippodamia convergens Guér- in. Coccinellidae populations were seldom high and the number of times beetles were collected was low compared to the number of nurseries visited. Key Words: host plants, prey Recently there has been a shift in the pest control practices of commercial nurseries away from cover sprays to spot spraying. Under this new regime, pest control by nat- ural enemies is both desirable and feasible. The Coccinellidae have long been known as major predators of various Homoptera and Acari. Gordon’s (1985) treatise on North American Coccinellidae has facilitated adult identification to species. A survey of the Coccinellidae found in Maryland nurseries was undertaken to determine the variety of species present, their frequency of occur- rence, and abundances. MATERIALS AND METHODS Adult Coccinellidae were collected during routine nursery inspections. In addition to the typical collection data, host plants, pres- ence and type of prey, and feeding activity by the coccinellids were also noted. Speci- mens were placed in 70% ethyl alcohol and taken to the laboratory for mounting. Adults were identified by the senior author using Gordon (1985). No efforts were made to identify larvae. Voucher specimens are de- posited in the Maryland Department of Ag- riculture collection. RESULTS AND DISCUSSION Twenty-eight species representing eigh- teen genera were collected. Coccinellidae were found in 67 locations in 1986, 82 lo- cations in 1987, and 37 locations in 1988. One reason for the low detection rate in 1988 is that several field personnel only re- ported unusual coccinellids. Possibly another reason was that the drought con- ditions and high temperatures during the summer made most adult coccinellids seek protected locations. Prey numbers were also low, possibly due to the same conditions. Four species were found in high numbers, but most were observed as occasional adults scattered over large areas. The species found are listed below with the counties, host plants (names as listed in Hortus III, 1976), prey associations or feeding activities, and the months in which specimens were collected. The relative frequency of each species is in- dicated. VOLUME 92, NUMBER 2 SCYMNINAE Scymnini Scymnus cervicalis Mulsant: Anne Arun- del, Prince George’s. Catalpa sp., Quercus sp. Associated with aphids (Homoptera: Aphididae). May. Two locations. S. tenebrosus Mulsant: Anne Arundel, Montgomery. Catalpa sp., Malus sp. As- sociated with aphids. May. Two locations. Stethorus punctum punctum (LeConte): Prince George’s. Gleditsia tricanthos L., Platanus sp., Quercus sp. April, July. Two locations. Hyperaspini Brachicantha felina (Fab.): Montgomery, Prince George’s. Betula sp., Malus sp. As- sociated with aphids. May, June, August. Four locations. B. rotunda Gordon: Harford, Montgom- ery. Betula sp., Malus sp., Robinia pseudo- acacia L. Associated with aphids. Four lo- cations. B. quadripunctata quadripunctata (Melsheimer): Prince George’s. May. One specimen. B. ursina (Fab.): Baltimore. June. One specimen. Hyperaspis binotata (Say): Baltimore, Prince George’s. Prunus sp., Quercus sp. April, May. Associated with Melanaspis ob- scura (Comstock) (Homoptera: Diaspidi- dae). Two locations. H. signata signata (Olivier): Anne Arun- del, Carroll. May, June. Juniperus scorulo- rum Sarg. Feeding on Carulaspis sp. (Ho- moptera: Diaspididae). Three specimens. H. proba (Say): Baltimore, Kent, Prince George’s. Quercus sp. July, November. As- sociated with WZ. obscura. Three locations. CHILOCORINAE Chilocorini Chilocorus kuwanae Silvestri: Specimens of this species were released against various Diaspididae. Recoveries were made in Prince George’s and Worcester. //ex cor- 311 nuta Lindl. & Paxt., I. crenata Thunb., Quercus spp. Feeding on Hemiberlesia la- taniae (Signoret), Lopholeucaspis japonica (Cockerell), 14. obscura. Two locations. C. stigma (Say): Baltimore, Carroll, How- ard, Kent, Montgomery, Prince George’s, St. Mary’s, Somerset. Acer sp., 4. palmatum Thunb., Euonymus alata (Thunb.) Sieb., Fraxinus sp., Prunus sp., Quercus sp., Q. palustris Muenchh., Q. robur L. Associated with Lepidosaphes yangagicola Kuwana, M. obscura, Pseudaulacaspis sp. (Diasipidi- dae), Eulecanium cerasorum (Cockerell) (Coccidae). March, April, May, June, July, September, November. Nineteen locations. This species was common in only one lo- cation. All other collections represent only a few individuals. Exochomus marginipennis (LeConte): Kent, St. Mary’s. Robinia pseudoacacia. May, November. Two specimens. COCCIDULINAE Coccidulini Rhyzobius lophanthae (Blasidell): Mont- gomery. Pinus sp. October. One specimen. COCCINELLINAE Coccinellini Adalia bipunctata (L.): Anne Arundel, Baltimore, Caroline, Carroll, Harford, Kent, Montgomery, Prince George’s, St. Mary’s. Acer rubrum L., Betula sp., Carpinus betulus L., Elageanus umbellata Thunb., Euony- mus alata, Forsythia intermedia Zab., G. triacanthos, Lonicera sempervirens L., Ma- lus sp., Picea abies (L.) Karst., Pinus mugo Turra, P. strobus L., Prunus sp., Pyracantha sp., Quercus bicolor Willd., Rosa rugosa Thunb., Salix sp., Sorbus acuparia L., Spi- rea nipponica Maxim., Viburnum sp., V. rhytidophyllum Hemsl. Feeding on aphids. April, May, June. Adalia bipunctata was the fourth most common species found in the survey (47 locations, five commonly). Anatis labiculata (Say): Anne Arundel, Baltimore, Carroll, Howard, Montgomery, 312 Prince George’s. Acer rubrum, A. saccha- rum Marsh., Betula sp., Cornus florida L., Ligustrum sp., Pyracantha sp., Quercus sp.., Q. acutissima Carruth., Tsuga canadensis (L.) Carriere, U/mus sp., Viburnum sp. April, May, June. Thirteen locations. A. mali (Say): Harford, Prince George’s. Pinus sylvestris L. May. Two specimens. Axion tripustulatum (DeGeer): Prince George’s. Quercus sp. Associated with M. obscura. April. One specimen. Coccinella novemnotata Herbst: Alle- gany, Carroll. June, July. Two locations. This species used to be very common in Maryland, judging by the number of spec- imens in student collections at the Univer- sity of Maryland and observations by the senior author. Since the introduction of C. septempunctata L., the species has only been collected twice in 1986. Whether this is due to a natural decline or competitive displace- ment is not known at present. C. septempunctata L.: Allegany, Anne Arundel, Baltimore, Carroll, Charles, Fred- erick, Harford, Howard, Kent, Montgom- ery, Prince George’s, Queen Anne’s, Som- erset, St. Mary’s, Talbot, Wicomico, Worcester. Acer saccharum, Betula sp., Crategus sp., Elageanus umbellata, Euony- mus sp., Fraxinus sp., Hemerocallis sp., Ilex cornuta Lindl. & Paxt., Juniperus sp., Malus sp., Pieris japonica (Thunb.) D. Don, Pinus sp., P. sylvestris L., P. thunbergiana Franco, Prunus sp., Pyracantha sp., Pyrus callery- ana Decne., Rhododendron sp., Rosa sp., Quercus sp., Q. robur, Spirea sp., Taxus sp., Tilia sp., Tsuga canadensis, Viburnum sp., Wisteria sp., Zelkova serrata (Thunb.) Ma- kino. March, April, May, June, July, Au- gust, September, October, November. This was the most commonly collected species in the survey (87 locations). Feeding on aphids. Coccinella septempunctata was tak- en in large numbers on several occasions. On three separate occasions adult C. sep- tempunctata were collected on the bark of cut Christmas trees of Picea abies, P. pun- gens Engelm., and Pinus sylvestris. The or- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON igin of these trees was eastern Pennsylvania. The movement of cut Christmas trees pro- vides an interesting method of distribution for this species. This species has also been collected in a greenhouse complex. Coleomegilla maculata lengi Timberlake: Anne Arundel, Baltimore, Caroline, Car- roll, Kent, Prince George’s, Queen Anne’s, St. Mary’s, Somerset, Talbot, Wicomico, Worcester. Acer rubrum, Coreopsis sp., Eu- onymus alata, E. japonica Thunb., Forsyth- ia intermedia, Gleditsia tricanthos, Hedera helix L., Hibiscus sp., Ilex crenata, I. opaca Ait., Juglanus sp., Magnolia sp., Phlox sp.., Pinus sp., P. sylvestris, Prunus sp., Pyran- cantha sp., Rosa sp., R. rugosa, Quercus sp., Salix sp., Spirea sp., Taxus sp., Tsuga can- adensis, Vitis sp., Z. serrata. Feeding on aphids; associated with Pineus strobi (Har- tig) (Homoptera: Phylloxeridae) and Dy- micoccus wisteriae (Green) (Pseudococci- dae). April, May, June, July, August. This was the second most common species in this survey (45 locations). Cycloneda munda (Say): Anne Arundel, Baltimore, Charles, Harford, Montgomery, Prince George’s, Queen Anne’s, Wicomico. Acer sp., Betula pendula Roth, Malus sp., Pyracantha sp., Quercus palustris Muenchh., Taxus sp. Feeding on aphids. May, June, July, August. Fifteen locations. Olla y-nigrium (Mulsant): Anne Arundel, Prince George’s, Talbot, Worcester. Mag- nolia sp., Ulmus sp. Associated with Tou- meyella liriodendri (Gmelin) (Homoptera: Coccidae). May, June, September. Five lo- cations. Hippodamia convergens Guérin: Anne Arundel, Baltimore, Carroll, Frederick, Harford, Kent, Montgomery, Prince George’s, St. Mary’s, Washington, Wicomi- co, Worcester. Acer rubrum, Cornus florida, Elageanus umbellata, Euonymus alata, Fraxinus sp., Hedera helix, Hibiscus syria- cus L., Hydrangea sp., Juglanus sp., Malus sp., Picea abies, Prunus sp., Pyracantha sp., Quercus sp., Rosa sp., Viburnum sp. Feed- ing on aphids. April, May, June, August. VOLUME 92, NUMBER 2 This was the third most common species collected (39 locations). H. glacialis (Fab.): Kent, Somerset. July. Two locations. Mulsantia picta (Randall): Caroline, Howard, Kent. May, June. Three speci- mens. Neoharmonia venusta venusta (Melshei- mer): Anne Arundel. Salix sp. May. One specimen. Psylloborini Psyllobora vigintimaculata (Say): Charles, Kent. May, September. Three locations. ACKNOWLEDGMENTS We would like to thank K. M. Berry, S. M. Bohlen, T. S. Creel, J. Fishback, K. Hog- sten, D. C. Laughlin, L. C. Miles, S. C. Par- ry, J. P. Smokonich, S. J. Stevenson, J. A. 313 Wolinski, and M. E. Zastrow for their as- sistance in collecting specimens in the field. We thank J. J. Drea, R. M. Hendrickson, and the rest of the staff of the Beneficial Insects Lab, USDA for supplying the Chilo- corus kuwanae released in Maryland nurs- eries. W. F. Gimpel, Jr., Maryland Department of Agriculture, and J. A. Davidson, Uni- versity of Maryland, commented on earlier drafts of this manuscript. Maryland Department of Agriculture Contribution Number 60-89. LITERATURE CITED Bailey, L. H. and E. Z. Bailey. 1976. Hortus Third. Macmillian Publ. Co., New York. 1290 pp. Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. New York Ento- mol. Soc. 93: 1-912. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 314-324 ANOMALIES IN CORNICLES OF APHIDS (HOMOPTERA: APHIDIDAE) LoulsE M. RUSSELL AND MANYA B. STOETZEL Systematic Entomology Laboratory, Plant Sciences Institute, USDA, ARS, Beltsville, Maryland 20705. Abstract.—Anomalies in number, size, shape, and other characteristics of cornicles (siphuncul1) of aphids are described and illustrated. Variations range from complete ab- sence of cornicles to a total of four in species that normally have two cornicles. Key Words: This article portrays assorted anomalies in cornicles (siphunculi) of aphids. The re- corded aberrations occur in 14 species that typically have two similar, tubelike corni- cles and in one species that has truncate cornicles. Abnormalities in cornicles occur infrequently but they are not rare, and they are seldom mentioned in literature, a con- dition that appears to justify a short treat- ment of the subject. Modifications in cornicles may be in numbers, size, shape, imbrications, reticu- lations, apical openings, and flanges. Vari- ations in number are less frequent than oth- er abnormalities. Flanges are absent when apices of cornicles are almost or completely closed. The closing of cornicle apices doubt- less is detrimental to survival of the insects, as was indicated by Dixon and Stewart (1975) who stated that exudates from the cornicles drove away predators. Earlier, Nault, Edwards and Styer (1973) indicated that cornicles of certain species emit pher- omones that repel other insects from feed- ing sites. Abnormalities are sometimes present in more than one individual ofa collection and other abnormalities may occur in speci- mens with modified cornicles. We noted aborted antennae in two species with ab- normal cornicles. Aphididae, cornicles, anomalies, aphids SPECIES WITHOUT CORNICLES One intermediate vivipara of Dysaphis foeniculus (Theobald) (Fig. 1) (on Daucus carota L., Heyden Trust, Sandy’s Parish, Bermuda, 24-III-1988, D. J. Hillburn, M. J. Mello, M. B. Stoetzel) is devoid of cor- nicles, while other specimens (Fig. 2) of the lot have two normal ones. SPECIES WITH ONE CORNICLE An alate vivipara of Hyalopterus pruni (Geoffroy) (Fig. 3) (on an unidentified plant, Bombay Hook, Delaware, 25-VI-1975, G. Angelet) has one typical cornicle. An apter- ous vivipara of Aphis masoni Richards (Fig. 4) (sweeping mixed meadow, Churchill, Manitoba, Canada, 24-VII-1975, A. G. Robinson) also has one normal cornicle. In an alate vivipara of Myzus ascalonicus Don- caster (Fig. 5) (on Sedum sp., England, in- tercepted at Seattle, Washington, 8-V-1972, J. D. Kail), the one cornicle 1s 4 shorter and more slender than the cornicles in another alata from the same collection; it is slender throughout, irregular in shape, and rounded apically with a minute opening. Stoetzel (1986) apparently was the first to record the presence of only one cornicle in an aphid, Cinara terminalis (Gillette and Palmer). VOLUME 92, NUMBER 2 315 e é Figs. 1-2. Dysaphis foeniculus. 1, Intermediate vivipara without cornicles. 2, Apterous vivipara from same collections with normal cornicles. Fig. 3. Hyalopterus pruni. Alate vivipara with only one cornicle. Fig. 4. Aphis masoni. Apterous vivipara with only one cornicle. Fig. 5. Myzus ascalonicus. Alate vivipara with only one cornicle. ’ 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Aphis craccivora. Apterous vivipara with dwarfed, nipple-shaped cornicles. Fig. 7. Aphis spiraecola. Alate vivipara with one cornicle shortened and rounded apically. Verbascum thapsus L., Athens, Georgia, 4-V-1973, O. R. Pagus), both cornicles are most unusual in that they are no longer than their basal diameters and are nipple-shaped, with apices closed. In an alate vivipara of SPECIES WITH TWO CORNICLES When two cornicles are present, one or both may be modified. In an apterous vi- vipara of Aphis craccivora Koch (Fig. 6) (on VOLUME 92, NUMBER 2 Figs. 8-9. 317 yo Y eee Aphis spiraecola. 8, Alate vivipara with one cornicle narrowed at midlength. 9, Alate vivipara with one cornicle strongly constricted just proximad of midlength. Figs. 10-11. Aphis fabae. 10, Apterous vivipara with one cornicle curved and unusually stout. 11, Apterous vivipara with one cornicle abnormally wide and with a conspicuous hole in its lower surface. Aphis spiraecola Patch (Fig. 7) (on Prunus lyonii (Eastw.) Sarg., Arcadia, California, 22-III-1966, H. G. Walker), one cornicle is about '2 the length of the normal one, is rounded, and apparently is closed apically. An alate vivipara of Aphis spiraecola Patch (Fig. 8) (on Taxodium distichum (L.) L. Rich., Chevy Chase, Maryland, 1-VI-1976, E. J. Hambleton) has one cornicle that ta- pers to its midpoint and then enlarges to its end; it is slightly shorter than the normal one, has few imbrications, and its apex has an opening but lacks a flange. Another alate vivipara of A. spiraecola (Fig. 9) (on Malus sp., Princess Anne, Maryland, 20-VI-1968, C. W. McComb) has one cornicle of nearly normal length that is strongly constricted just proximad ofits midlength, has very few imbrications, and has a typical apical open- ing but lacks a flange. A second alate vivip- ara (not shown) of this collection has one cornicle that is 2 the length of the normal one, has few imbrications, and 1s rounded and closed apically. An apterous vivipara of Aphis fabae Scopoli (Fig. 10) (on Papaver orientale L., Polur, Iran, 16-VII-1974, G. Buckingham) has a curved cornicle that is somewhat shorter and stouter than the nor- mal one and apparently is closed apically. In another apterous vivipara of 4. fabae (Fig. 11) (on Papaver sp., Greeley, Colorado, IX- 1932, W. J. Zaumeyer), one cornicle is slightly shorter than the other and has a subcircular hole in its lower surface; it has a slight indication of a flange. An apterous vivipara of Myzus persicae (Sulzer) (Fig. 12) (on Blechnum occidentale L., Arcadia, California, 26-III-1969, H. G. 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-13. Myzus persicae. 12, Apterous vivipara with one cornicle shortened and club-shaped. 13, Alate vivipara with one cornicle finger-shaped and one somewhat thimble-shaped. Walker) has a shortened, club-shaped cor- nicle that is closed apically. In an alate vi- vipara of M. persicae (Fig. 13) (from yellow pan trap, Fort Kent, Maine, 25-VII-1974, F. R. Holbrook), both cornicles are much less than normal length: one is somewhat thimble-shaped and the other finger-shaped; they have few imbrications and their apices are closed. Antennae of this insect terminate abruptly at the end of the fifth segments; the primary sensorium is present in one anten- na and absent from the other. In an alate vivipara of Nasonovia ribisnigri (Mosley) (Fig. 14) (on Lactuca sativa L., Spain, in- tercepted at New York, New York, 1-V- 1967, F. Planer), one cornicle is 2 the length of the normal one and tapers gradually to an acute, closed tip; its distal portion is de- void of imbrications. In an apterous vivip- ara of Dactynotus sonchi (Geoffroy) (Fig. 15) VOLUME 92, NUMBER 2 319 Fig. 14. Nasonovia ribisnigri. Alate vivipara with one cornicle shortened and tapered. Fig. 15. (on Sonchus oleraceus L., Honolulu, Ha- wall, 20-VI-1975, R. Mau), one cornicle is Ys the length of the normal one, tapers, then enlarges and terminates in a knob; the pres- ence or absence of a terminal opening can- not be determined because of debris. The cornicle has imbrications but lacks reticu- lations. Dactynotus sonchi. Apterous vivipara with one cornicle shortened and knobbed apically. In two specimens of Acyrthosiphon kon- doi Shinji (on Medicago sativa L., Quetta Valley, Pakistan, 19-VI-1978, S. Hamid), one cornicle in an alate vivipara (Fig. 16) is about '2 the length of the normal one and is constricted near its fingerlike end which has a small opening but no flange. In an alatoid nymph (Fig. 17), one greatly abbre- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 16-18. Acyrthosiphon kondoi. 16, Alate vivipara with one cornicle shortened and tapered. 17, Alatoid nymph from same collection with one cornicle dwarfed and bearing a stublike protrusion on its distal margin. 18, Alate vivipara with one cornicle greatly dwarfed. viated cornicle lacks imbrications and a flange; it has a stublike extrusion on its dis- tal margin. And in another alate vivipara of A. kondoi (Fig. 18) (on Medicago sativa L., St. George, Utah, 11-V-1975, D. Hu- ber), one cornicle is a stub no longer than the basal diameter of the normal cornicle; it has few imbrications and an apical open- ing but it lacks a flange. In an apterous intermediate of Macrosi- phum euphorbiae (Thomas) (Fig. 20) (on Rosa sp., England, intercepted at Philadel- phia, Pennsylvania, 10-VI-1964, F. Harvey and N. Arehart), one cornicle 1s about 2 the length of the normal one, tapers slightly to a rounded, closed apex, and has few imbri- cations but no reticulations. There is a transverse, oblong opening on its dorsal sur- face (Fig. 19). This specimen has an atypical cauda, the apex of the abdomen resembling that of a nymph instead of an adult. In an alate vivipara of M. euphorbiae (Fig. 21) VOLUME 92, NUMBER 2 Figs. 19-21. w in Macrosiphum euphorbiae. 19, Abnormal cornicle of apterous intermediate enlarged to show hole in its upper surface. 20, Apterous intermediate with one cornicle modified. 21, Alate vivipara with one cornicle dwarfed. (from yellow pan trap, Beltsville, Maryland, 10-X-1975, W. W. Cantelo), a cornicle dwarfed to about 4 normal length has few imbrications, no reticulations and a small apical opening without a flange. Two alate viviparae of Macrosiphum ro- sae (L.) (on greenhouse roses, New Bruns- wick, New Jersey, 28-X-1975, H. T. Streu) have modified cornicles and antennae. In one specimen (Fig. 22), the cornicles are greatly shortened and their ends are closed; one tapers gradually and 1s pointed apically while the other is of nearly uniform diam- eter and is rounded apically; both have few w NR i) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON * a »- \ a b | - Figs. 22-23. Macrosiphum rosae. 22, Alate vivipara with both cornicles dwarfed and mis-shaped. 23, Alate vivipara from same collection with both cornicles atypical in shape. VOLUME 92, NUMBER 2 323 Fig. 24. Aphis sambucifoliae. Alate vivipara with three cornicles, one greatly reduced in size proximad of a normal one. Fig. 25. Euceraphis punctipennis. Alate vivipara with four cornicles, the anterior pair normal, the posterior pair supplementary. imbrications and no reticulations. In this insect, one antenna terminates and is weak- ly nipplelike at the end of the fourth seg- ment; the other antenna is slightly shorter and is curved with an invagination at its end. In the other specimen (Fig. 23), one of the unusually slender cornicles 1s narrowed abruptly, has four indistinct rows of reticu- lations, an abnormal apical opening, and no flange. The other cornicle, narrowed nearer 324 the distal end, has 10 distinct rows of reti- culations, a normal opening, and a flange. In the latter aphid, one antenna 1s aborted at the distal end of the third segment, and the other antenna has only the proximal portion of the fourth segment; apices of both antennae are rounded. Malformations similar to, or varying slightly from, those discussed are present in other specimens. SPECIES WITH THREE CORNICLES Remaudiére (1964) noted the presence of three cornicles on nymphs and adults of Aphis sp. near esulae Boerner. Of 19 spec- imens studied, two adults and six nymphs had three cornicles. Medler and Ghosh (1967) noted three cornicles on an alate vi- vipara of Macrosiphum sp. Russell (1975) observed three cornicles on an alate vivip- ara of Aphis sambucifoliae Fitch (Fig. 24) (prey of Asilidae, Baltimore County, Mary- land, VIII-1973, A. G. Scarbrough). The ad- ditional cornicle is shorter and narrower than the other two and 1s nearly devoid of im- brications; the apical opening 1s replaced by a conical invagination and there is no flange. SPECIES WITH FOUR CORNICLES Remaudiére (1964) found four cornicles in three nymphs of Aphis sp. near esulae Boerner, while two adults and six nymphs also in his collection had only the normal pair of cornicles. Leonard (1967) observed a pair of adventitious cornicles on an ap- terous vivipara of Aphis sambucifoliae Fitch. Russell (1975), who recorded four cornicles in an alate vivipara of Euceraphis puncti- pennis (Zetterstedt) (Fig. 25) (on Betula sp., Mesa, Colorado, 8-VI-1967, F. C. Hottes), stated **.. . it is the only one of 80 individ- uals of the lot that exhibits a duplication of cornicles.” The adventitious cornicles ap- pear to differ from the typical pair only in their smaller size. In all specimens, the ad- ditional cornicles are located on the sixth abdominal segment, posterior to, and usu- ally shghtly proximad of, the normal pair. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BRANCHED CORNICLES Zirnits (1930) recorded conspicuous branching of cornicles in apterous viviparae of Megoura viciae Buckton. Perhaps the stublike extrusion we observed on the aborted cornicle of a nymph of Acyrthosi- phon kondoi (Fig. 17) exhibits this anomaly. Branched and supernumerary cornicles are seen less frequently than cornicles mod- ified in size, shape and other characteristics. ACKNOWLEDGMENTS For their review of this paper, we thank the following individuals: James B. Kring, University of Florida, Bradenton, FL 34216; Clyde F. Smith, Department of Entomol- ogy, Box 5215, North Carolina State Uni- versity, Raleigh, NC 27650; and Richard E. White, Systematic Entomology Laboratory, Agricultural Research Service, U.S. De- partment of Agriculture, % U.S. National Museum of Natural History, Washington, DC 20560. LITERATURE CITED Dixon, A. F. G. and W. A. Stewart. 1975. Function of the siphunculi in aphids with particular refer- ence to the sycamore aphid, Drepanosiphum platanoides. J. Zool. Soc. London 175: 279-289. Leonard, M. D. 1967. An aphid with four cornicles (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. (1966) 68: 320. Medler, J. T. and A. K. Ghosh. 1967. An alate aphid with an extra cornicle (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 69: 366. Nault, L. R., L. J. Edwards, and W. E. Styer. 1973. Aphid alarm pheromones: Secretion and recep- tion. Environ. Entomol. 2: 101-105. Remaudiére, G. 1964. Sur la présence de cornicules surnumeéraires chez un Aphis (Hom. Aphididae). Rev. Path. Veg. Entomol. Agr. France 43: 31-35. Russell, L. M. 1975. Euceraphis punctipennis (Zet- terstedt), the fourth aphid species with four cor- nicles (Hemiptera: Homoptera: Aphididae). J. Wash. Acad. Sci. 65: 40-41. Stoetzel, M. B. 1986. The first report ofan aphid with only one cornicle (Homoptera: Aphididae). Proc. Entomol. Soc. Wash. 88: 389-390. Zirnits, J. 1930. Duplication of cornicles in Megoura viciae Kalt. Fol. Zool. Hydrobiol. 2: 1-3. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 325-332 A NEW SPECIES OF CRASPEDOXANTHA AND A REVISED PHYLOGENY FOR THE GENUS (DIPTERA: TEPHRITIDAE) AMNON FREIDBERG AND WAYNE N. MATHIS (AF) Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel-Aviv University, Tel-Aviv, 69978, Israel; (WNM) Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560, USA. Abstract. —Craspedoxantha bafut, reared from Vernonia calvoana and V. adoensis, 1s newly described from specimens collected in Cameroon and Nigeria. A cladistic analysis of the 10 species of Craspedoxantha Bezzi with Orellia punctata (Schrank) as an outgroup was performed using Hennig86 (c). Using the “implicit enumeration” option of Hennig86, four trees of equal length (16 steps) were calculated from which a Nelson consensus tree was then generated. Using the successive weighting technique of Hennig86 further reduced the number of trees to two, which differed only in the sequence of Afrotropical species of the manengubae group. A Nelson consensus tree of these two trees is the same as the second tree. All trees that were calculated, including the Nelson trees, confirmed the validity of the two previously established species groups, marginalis and manengubae. Key Words: Craspedoxantha, fruit flies, phylogeny, Vernonia Freidberg (1985) recently revised Cras- pedoxantha Bezzi and included a discussion of its biology and phylogeny. His phylogeny divided the genus into two species groups: the marginalis group, with four Afrotropical species, and the manengubae group, with five species, of which three are Afrotropical and two are Oriental. The character data for the phylogeny were analyzed by hand, and the topology of the published tree was based in part on intuition. During a recent field trip to Cameroon and Nigeria, a new species of Craspedoxan- tha was reared and collected from species of Vernonia (Asteraceae). Its discovery pro- vides a good opportunity to test and refine the phylogeny of the genus with the addition of more characters and by using a computer program (Hennig&86 (copyrighted), see Fitz- hugh 1989 for description) for calculating and analyzing trees from the character data. Craspedoxantha bafut Freidberg and Mathis, NEw SPECIES Figs. 1-4 Diagnosis.—This species is placed un- ambiguously in the manengubae group (Freidberg 1985: 189, 202) because of its similarity to the Afrotropical congeners, manengubae Speiser, yaromi Freidberg and vernoniae Freidberg, of that species group. In the key to species (loc. cit.), this species runs to couplet 8, which contains vernoniae and manengubae. It differs from vernoniae by the yellow marginal wing band, which does not widen opposite crossvein r-m. It differs from manengubae by the apical blackish spot on the wing, which does not broaden in cell r4+5 and is less than “4 as wide as the length of the apical section of vein M. The terminalia, however, are more 326 similar to those of C. yaromi, especially the aculeus (Figs. 1-2), which is only slightly more rounded and wider at the tip, the sper- matheca (Fig. 3), and the epandrium (Fig. 4), which is slightly different in lateral view but indistinguishable in ventral view. C. bafut can otherwise be distinguished from C. yaromi by its larger size (length of wing: 5.5-6.5 mm versus 4.5—5 mm in C. yaromi), by having predominantly dark, not pale, setulae on the scutellum, by the marginal wing band not widened opposite crossvein r-m, and by the lack of yellow pattern in and around the discal cell, although a mi- crotrichial pattern, similar to the yellow pat- tern usually present on the wing of C. ya- romi, is usually present in C. bafut. Description. — Fitting the generic descrip- tion (Freidberg 1985), with the following details. Head: Similar in shape to C. marginalis (Freidberg 1985, fig. 1), but with shorter Ist flagellomere, only 1.5-2 times as long as wide and 1.5 times as long as pedicel (in C. marginalis 2.5 times). Dull yellow, except face whitish and ventral facial margin shiny. Thorax: Normal for genus; scutal length to width ratio = 1.1:1; dorsocentral setae aligned about midway between anterior su- pra-alar setae and suture; | anepisternal seta; dorsocentral and prescutellar black spots large; supra-alar spot and presutural spot slightly smaller than dorsocentral spot and often more elongate; scale-like setulae on scutum very dense, white or yellow, hardly extended beyond black, lyre-like pattern, which is discernable with difficulty. Scutel- lum unspotted, predominantly covered by blackish setulae, with yellowish setulae only near insertion of basal scutellar seta; pleura not striate; subscutellum and mediotergite brownish to blackish, densely covered by grayish-yellow microtomentum, except cor- ners of subscutellum and ventral margin of mediotergite, which in posterior view ap- pear less densely covered by microtomen- tum. Calypteres white to yellow, with or without brownish margins; halter yellow. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Legs: Yellow, elongate; femora without dis- tinctly dense setulae ventrally. Wing: Length 5.5-6.5 mm; marginal band uniformly nar- row, without a bump opposite crossvein r-m; apical blackish spot evenly narrow, as wide as 4-4 of terminal section of vein M; cell cup very lightly yellow; wing with micro- trichial pattern in and around cell dm sim- ilar to the yellow pattern of C. yaromi (Freidberg 1985, fig. 13); microtrichia lack- ing from apical '2 of cell br except near pos- terior part of crossvein r-m, from basal part of cell r4+5 except near posterior part of crossvein r-m, from cell bm and from base of cell dm; vein R4+5 with 2-8 setulae dor- sally and ventrally at node. Abdomen: Normal for genus; with pre- dominantly yellow setulae, tergite 3 through last with | or few rows of brown setulae posteriorly; pattern of black spots reduced in 4 to anterior band on tergite 5, which is microtomentose, matt, and a pair of small spots at posterior margin, and often entirely reduced in 2, although in some ¢@ a pair of small spots present at anterior margin of tergite 6 and at base of syntergosternite 7 (oviscape) and tip of syntergosternite 7 nar- rowly blackish; syntergosternite 7 about as long as combined length of posteriormost 4 tergites; ¢ terminalia as in Fig. 4; 2 termi- nalia as in Figs. 1-3. The distiphallus is practically indistinguishable from that of C. vernoniae (Freidberg 1985, Fig. 29). Type material.— Holotype 2: “CAMER- OON, Rt. N6 Bali-Batibo W. of Bamenda 20.X1.1987 A. FREIDBERG.” The allo- type 4, and four paratypes (2 4, 2 2) have the same label data as the holotype, except the collector of one ¢ is Fini Kaplan, and one 2 is also labeled: ex. flowerhead of Ver- nonia calvoana 23 Nov 1987. Additional paratypes are as follows: CAMEROON. Northwest Province: Rt. P16 Mbengwi 25 Km W Bamenda, 23 Nov 1987, Fini Kap- lan, 1 9; Rt. N11, Bafut 20 Km N. Ba- menda,17-24 Nov 1987, A. Freidberg, | 2. NIGERIA. Plateau State: Kurra Falls, 60 Km SE Jos, 5-7 Dec 1987, A. Freidberg, 2 VOLUME 92, NUMBER 2 Figs. 1-4. Craspedoxantha bafut. 1, aculeus. 2, apex of aculeus. 3, spermatheca. 4, epandrium, lateral view. 6 2 2, of which | 4 2 2 are also labeled: ex. flowerhead of Vernonia adoensis, 10 Dec 1987; Kefh, Rt. 234, 4 Dec 1987, Fini Kap- lan, 1 4. The holotype is in excellent con- dition, is pinned directly, and is deposited together with most paratypes in the Zoo- logical Museum, Tel-Aviv University. Paratypes have also been deposited in the BMNH and USNM. Biology and host plants.— All specimens were collected or reared from Vernonia adoensis Sch. Bip. ex Walp. or V. calvoana (Hook. f.) Hook. f. (Asteraceae). The spec- imens from Rt. N6, between Bali and Ba- tibo, were collected together with numerous specimens of C. manengubae, a species that probably also breeds in V. cal/voana, al- though this latter association has not been confirmed. Etymology.— This species is named after Bafut, a picturesque village in the highlands of Cameroon, where this species was col- lected. The specific epithet is a noun in ap- position. PHYLOGENY OF CRASPEDOXANTHA Freidberg (1985) briefly discussed the phylogeny of Craspedoxantha and noted that the genus clearly formed a monophyletic group within the tribe Terelliini. The syn- apomorphies that establish the monophyly of the genus are those Freidberg used to distinguish it in his key to the genera of Terelliini. With one additional character, these synapomorphies are as follows: 1. Eye is 1.5—2 times higher than long. 2. Scutum with three pairs of black spots that are uniquely arranged as follows: at the base or immediately behind the pre- sutural supra-alar seta and at the base of the dorsocentral and prescutellar acros- tichal setae. 3. Scutellar disc with at least some blackish setulae. 4. Wing with a complete, mostly yellow costal band. The apex of the band, from vein R2+3 to slightly beyond vein M, is mostly blackish, and there are usually three, seldom two, blackish, evenly spaced spots in cell rl. Freidberg further noted that among ter- elliines the closest relative of Craspedoxan- tha was probably Orellia Robineau-Des- voidy, especially species in the falcata group, which includes punctata (Schrank) (the type species of Orellia), falcata (Scopoli), and distans (Loew.) Korneev (1985) removed from Orellia all species except those of Freidberg’s falcata group, thus reducing the genus to a more firmly established mono- phyletic group. In the analysis to follow, Orellia punctata, the exemplar we chose to represent Orellia, was selected as the outgroup of Craspedo- xantha. Although O. punctata appears to be a suitable candidate for this analysis, we are not completely satisfied with this selection. Our reservations derive from a lack of phy- logenetic perspective on the tribe in general. Despite several recent publications on the taxonomy of the tribe (Freidberg 1985, Freidberg and Mathis 1986, Korneev 1982, 1985, 1987), very little has been reported on the phylogeny of the included taxa (e.g. Freidberg 1985). A better understanding of the phylogeny of the tribe and its six or seven currently recognized genera (seven in Freidberg (1985), six in Korneev (1987), who relegated Cerajocera Rondani to subgeneric status within the genus Terellia Robineau- Desvoidy) would have greatly facilitated the selection of an outgroup for Craspedoxan- tha. The main impediment to achieving a phylogeny for the tribe is our lack of knowl- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON edge about the outgroup of the Terelliini. Terelliini is now generally thought to belong in the subfamily Tephritinae, and from oth- er studies, we suggest that Yyphosia Robin- eau-Desvoidy could possibly be its sister group, although this and other possibilities await further study and resolution. Despite not having a well-corroborated phylogeny from which an outgroup for Craspedoxantha could be selected, we feel that Orellia ought to be considered for the following reasons. Orellia, as characterized by Korneev (1985) and accepted by us, shares with Craspedoxantha three charac- ters that are probably synapomorphies: (1) a generally similar arrangement of black mesonotal spots, (2) a relatively short and heavily sclerotized distiphallus that lacks elongate, distal tubes (Korneev’s (1985) “paired sclerites of epiduct” —this character is also shared by Chaetorellia Hendel, Chae- tostomella Hendel and some Terellia), and (3) the host plants of Craspedoxantha and Orellia tend to be in Asteraceae other than the tribe Cardueae, a character also shared by Neaspilota Osten Sacken. Nine characters were used in the phylo- genetic analysis, and most of these were il- lustrated previously (Freidberg 1985). In ac- cordance with standard procedures for cladistic analysis, we ordered and polarized the characters. We then coded the charac- ters, with the most plesiomorphic states, such as those of the outgroup, as 0, and the more apomorphic states as 1 and 2. The coding we assigned to character states 1s giv- en in parentheses. For purposes of clarifi- cation, we have included, as needed, an ex- planation of the characters in the listing as follows: 1. Scutellum with 4 (0), 2 (1), or 0 (2) black spots. 2. Anepisternal setae numbering 2 (0), or 1 (1). 3. Femora slender and lacking dense in- vestment of setulae (0), or swollen and densely setulose ventrally (1). VOLUME 92, NUMBER 2 329 Orellia punctata >. ET milleri 7 ol (i ae polyspila : | 2 ees unimaculata ————EE— marginalis BS | —— octopunctata 1(2), _.- itdica o- ee e yaromi 2?,6,8,9 = vernoniae [= manengubae bafut Tree length 17; Consistency Index 64; Retention Index 82 Character: at 2 S 4 5 6 7 8 9 Steps: 3 1 1 1 2 1 3 3 2 Consistency Index: 66 100 100 100 50 100 33 66 50 Retention Index: 83 100 100 100 75 100 50 75 50 Fig. 5. Nelson consensus tree and its analysis for species of Craspedoxantha with Orellia punctata as the outgroup. A ‘*?” denotes characters that are partially homoplasious. = Dorsocentral setae aligned with anterior supra-alar setae (0), or inserted more an- teriorly (1). Cell cup distinctly (0), or indistinctly (1) yellow. . Presutural black spots about as large as (0), or distinctly smaller than (1) dor- socentral spots. wn lon) Table 1. Character matrix for Orellia punctata and species of Craspedoxantha. Missing or unavailable data are indicated by a ?. 123456789 Taxon (Characters) Orellia punctata 000000000 C. unimaculata 1010001?? C. marginalis 101000101 C. milleri 0010000?? C. polyspila 001010100 C. octopunctata 21000000? C. indica 210100000 C. yaromi 210111011 C. vernoniae 210111021 C. manengubae 210111121 C. bafut 210111111 7. Marginal band on the wing approaching (0) or not approaching (1) crossvein r-m. In some species of Craspedoxantha the marginal wing band is uniformly narrow and does not noticeably approach the junction of crossvein r-m and vein R4+5 (coded 1). In most other species the band has a small posterior bump that reaches or almost reaches this point. In C. milleri Freidberg and species of Orellia this bump is actually part of a transverse band. For all the latter taxa it was coded 0. . Posterior margin of the epandrium, the surface from which the cerci arise, in lat- eral view concave (0), straight (1) or con- vex (2). . Host plants: Host plant associations that are in part or exclusively with Vernonia (tribe Vernonieae) are hypothesized as the derived condition (1). We are unable to differentially treat other associations, whether they include plants of the tribe Cardueae (hosts of most Terelliini), Lac- 330 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Orellia punctata milleri | -——_—_—_— polyspia —_ marginalis octopunctata = reas | -— apie | = vernoniae manengubae Tree length 88; Consistency Index 86; Retention Index 94 Character: il! 2 3 4 5 6 7 8 ) Steps: ae 1 3 oa: S2rleasrsseeeeee 66 100 100 100 50 100 33 100 50 83 100 100 100 75 100 50 100 50 Consistency Index: Retention Index: Orellia punctata milleri polyspila ee 3 Tree length 88; Consistency Index 86; Retention Index 94 Character: 1 2 Sige tA De Owe be) eet, Steps: 3 al a: al! 2 al 3 2 2 Consistency Index: 66 100 100 100 50 100 33 100 50 Retention Index: 83) 100 1007 1007 75-1100" 50 1005750 Figs. 6-7. Two trees calculated with the successive weighting technique and their analysis. The second Nelson consensus tree, which was calculated from these two trees, is identical with Fig. 7. tuceae (hosts of Orellia species) or others and considered them all as primitive (0). Trees were calculated from the character data using the “implicit enumeration” op- tion of Hennig&86. This option generates the most parsimonious tree(s), i.e. the tree(s) of ™ minimal length or with fewest number of steps. Four trees of equal, minimal length (16 steps) were calculated, and a Nelson consensus tree (Fig. 5) was then generated from these four trees to demonstrate where branches and relationships are consistent (most of the lineages) or inconsistent (the VOLUME 92, NUMBER 2 African members of the manengubae group). The overall consistency index for this Nel- son tree is 0.64, with a “retention index” of 0.83 (formula for the “retention index” is in Fitzhugh 1989). An analysis of this tree indicates that about half of the characters (2, 3, 4 and 6) have a perfect consistency index and that the characters that are most homoplasious (as judged by the indices) are characters 5, 7 and 9. Characters 5 and 7 deal with wing pattern, a feature that is often subject to homoplasy in Tephritidae. Char- acter 9 involves host-plant data and in ad- dition to having some apparent homoplasy, also suffers from a lack of information for three of the species. As more than one tree resulted from the “implicit enumeration” option, we then used the successive weighting technique (Farris 1969, Carpenter 1988) to further resolve and assist in the selection of a tree. Two trees resulted from this procedure (Figs. 6-7) and were summarized in the form of a second Nelson consensus tree (Fig. 7). The two trees differ only in the African part of the ma- nengubae group, which is the clade con- taining bafut, manengubae, vernoniae and yaromi. Both of these trees place manen- gubae and vernoniae as a monophyletic group (sister species), and the other two species either form an unresolved trichot- omy with this monophyletic group (Fig. 6), or have bafut as the sister group to manen- gubae + vernoniae, and yaromi as a sister group to the other three species (Fig. 7). The second Nelson tree was identical with the second successive weighting tree (Fig. 7). At the moment we prefer the unresolved pos- sibility of the first Nelson consensus tree (Fig. 5) over the other trees, because we feel that the relationships between these four species are as yet unresolved. Two characters that have not been used in the cladistic analysis, should be men- tioned. The first is the superficial similarity between the wing pattern of yaromi and the microtrichial pattern of bafut, which, to- gether with the great similarity in the ter- 331 minalia of both species, may indicate sister- group relationships between these species. The second is the zoogeographical pattern of the four species, with vernoniae and ya- romi apparently restricted to East Africa, and bafut and manengubae apparently re- stricted to West Africa. This zoogeograph- ical pattern may indicate the actual sister- group relationships between these four species, which differs from that suggested by the previous character. Although these species are very closely related and similar, at least in the adult stage, it is possible that studies of immature stages will resolve this quadrichotomy. It is interesting to compare the Nelson trees (Figs. 5, 7) of this study (which in- cludes bafut n. sp.) with Freidberg’s (1985) intuitive tree. The similarity is rather strik- ing. The two previously established species groups (marginalis and manengubae) are as clear in all trees that were calculated using Hennig86 and the composition of the groups is the same. There are, however, two dis- crepancies. In the marginalis group, milleri is placed by Hennig86 as a sister species to the other species of this group; whereas in the intuitive tree it is the sister species to polyspila Bezzi only. In the manengubae group, octopunctata is placed as a sister species to the other species of this group; whereas in the intuitive tree it is the sister species to indica Zaka-ur-Rab only. These discrepancies are mainly the result of pre- viously underestimating single characters, such as the wing pattern of mii//eri, which differs markedly from other patterns of its species group, and using zoogeographical considerations that were not used in the present analysis. C. indica and octopunctata were considered sister species in the intu- itive tree because, in addition to overall morphological resemblance, they are the only Oriental congeners, a fact that was giv- en more weight than some morphological evidence. In summary, the use of Hennig86 or sim- ilar computerized algorithms 1s strongly rec- 332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ommended mainly because of their objec- tivity and their ability to analyze large numbers of taxa and characters quickly. In addition, a prerequisite to using Hennig&86 is the preparation of a well-documented character matrix, which undoubtedly im- proves the thoroughness of revisionary work. ACKNOWLEDGMENTS We wish to thank Fini Kaplan and Y. Zvik for collecting flies for this study, Allen L. Norrbom and Jon K. Gelhaus for criti- cally reviewing this paper and Elaine R. S. Hodges for inking the illustrations. LITERATURE CITED Carpenter, J. M. 1988. Choosing among multiple equally parsimonious cladograms. Cladistics 4(3): 291-296. Farris, J. S. 1969. A successive approximations ap- proach to character weighting. Syst. Zool. 18: 374— 385. Fitzhugh, K. 1989. Cladistics in the fast lane. J. N.Y. Entomol. Soc. 97(2): 234-241. Freidberg, A. 1985. The genus Craspedoxantha Bezzi (Diptera: Tephritidae: Terelliinae). Ann. Natal Mus. 27(1): 183-206. Freidberg, A. and W. N. Mathis. 1986. Studies of Terelliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithson. Contr. Zool. 439: 1-75. Korneev, V. A. 1982. Knowledge of the tephritid fly fauna (Diptera, Tephritidae) of the European part of the USSR. Vest. Zool. 1982(2): 83-84 [In Rus- sian]. . 1985. Fruit flies of the tribe Terelliini Hendel, 1927 (Diptera, Tephritidae) of the fauna of the USSR. Entomol. Obozr. 64: 626-644 [In Russian, English translation in Entomol. Rev. 65(1): 35- 55]. 1987. A revision of the subgenus Cerajocera stat. n. of the genus Terellia (Diptera, Tephritidae) with description ofa new species of fruit flies. Zool. Zh. 1987: 237-243 [In Russian]. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 333-346 METHODS FOR IDENTIFICATION OF ANASTREPHA LARVAE (DIPTERA: TEPHRITIDAE), AND KEY TO 13 SPECIES G. J. Steck, L. E. CARROLL, H. CELEDONIO-HURTADO AND J. GUILLEN-AGUILAR (GJS) Department of Entomology, Texas A&M University, College Station, Texas 77843, and Systematic Entomology Laboratory, USDA-ARS, BARC-Bldg. 476, Beltsville, Maryland 20708 (Correspondence: SEL.) (LEC) Department of Entomology, Texas A&M University, College Station, Texas 77843; (HC, JG) Programa MoscaMed, DGSV, SARH Apdo. Postal 368, Tapachula, Chiapas, 30700, Mexico. Abstract. — Detailed methods are provided for observing useful characters to distinguish among species of Anastrepha fruit flies in their immature stages. Additionally, a key is provided to third instar larvae of 13 species: 4. bistrigata Bezzi, distincta Greene, frater- culus (Wiedemann), grandis (Macquart), interrupta Stone, leptozona Hendel, limae Stone, ludens (Loew), obliqua (Macquart), pallens Coquillett, serpentina (Wiedemann), striata Schiner and suspensa (Loew). There is considerable overlap in many character states among species. Discriminant analysis is necessary to distinguish among species in some couplets. Key Words: Anastrepha is a New World genus of fruit flies (Diptera: Tephritidae) comprising about 180 valid species (Norrbom and Kim 1988). Several species are major fruit pests in the American tropics and subtropics. Descrip- tions are available for immature stages of only 15 species, and several of these are very incomplete. The paucity of taxonomic i1n- formation makes it extremely difficult to identify the larvae of Anastrepha (as well as those of most other fruit flies). This is es- pecially problematical because fruits infest- ed with larvae usually are encountered in the absence of associated adults, as is the case of most interceptions of Tephritidae at U.S. ports of entry (APHIS 1987). Published keys or descriptive works which specifically attempt to discriminate among larvae of two or more Anastrepha species include Greene (1929), Phillips (1946), Berg (1979), Heppner (1984), Steck and Wharton Anastrepha, fruit flies, larvae, discriminant analysis (1988), Steck and Malavasi (1988) and Car- rolland Wharton (1989). Twelve species are included in these works, but not all of them simultaneously. Berg’s key (1979) is the most inclusive and treats those six species con- sidered to be the most serious pests (frater- culus (Wiedemann), /udens (Loew), obliqua (Macquart), serpentina (Wiedemann), stria- ta Schiner, and suspensa (Loew). Unfortu- nately, some of the characters used in Berg’s key are very difficult to interpret. Also, the natural variability within species is such that the key often leads to an incorrect identi- fication. In this paper, we incorporate additional characters not previously utilized. Since the natural variability of larval characters is so poorly documented, especially among geo- graphical regions, we are careful to note the source and sample sizes of all the material utilized to generate the key. We have ad- 334 dressed the problem of variation by ex- amining specimens from more than one lo- cality to the extent that material was available. The economically most important species are all included, as well as several other available species whose identities were cer- tain. Thus, the usefulness of various larval characters could be assessed across a broad taxonomic range within this large genus. The 13 species treated here represent six differ- ent species groups within the genus (Norr- bom and Kim 1988). Among species clas- sified in the same group, the amount of overlap in character states is expectedly high. For some couplets it is necessary to employ discriminant analysis to determine the species to which a particular specimen be- longs. MATERIALS AND METHODS Specimens from which the key was de- veloped are listed below. Accurate associ- ation of larvae with their identifiable adult forms was a critical objective of this study. Identity of larvae cannot be presumed if they are taken from naturally infested fruits, since even an individual fruit may be mul- tiply infested by more than one species. Most museum specimens are not explicitly as- sociated with reared adults from the same collection; thus, their identity must be con- sidered cautiously. Many larvae used in this study were bred in the laboratory from known adults. Others were taken from nat- urally infested fruits from which numerous adult specimens of exclusively one species were reared. Exceptions are noted below. Specimen collectors’ names are given in pa- rentheses after the collection date. Local names for host fruits are given in parenthe- ses after their scientific names. Asterisks de- note specimens used to generate linear dis- criminant functions. Voucher specimens of all larvae and associated adults are housed in the collections of the U.S. National Mu- seum of Natural History, Smithsonian In- stitution (USNM) and/or the Department PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Entomology, Texas A&M University as TAMU voucher numbers 213, 214, 219, 220; 222,223. 225, 226vand 227//, A, bistrigata Bezzi— BRAZIL: Sao Paulo, Universidade de Sao Paulo, XI-1986 (Steck, Malavasi); 23* + 7 specimens from a lab- oratory culture on Psidium guajava L. (gua- va) initiated with adults reared from guava, Campinas, S.P. (See Steck and Malavasi 1988.) A. distincta Greene—VENEZUELA: Merida, Merida, V-1988 (Steck, Norrbom); 5 specimens from /nga sp. BRAZIL: Bahia, Cruz das Almas, VI-1988 (Steck, Con- ceicao); 5 specimens from Jnga sp. MEX- ICO: Chiapas, Tapachula vicinity, Obre- gon, III-1986 (Carroll); 10 specimens from Inga (larval identity presumed from host relationship). HONDURAS: E.A.P., 30 km s.e. of Tegulcigalpa, V-1985 (Sequiera); 10 specimens from /nga (larval identity pre- sumed from host relationship). A. fraterculus (Wiedemann)— BRAZIL: Sao Paulo, Itaquera, XI-1986 (Steck, Mala- vasi); 25* specimens from Eugenia brasi- liensis Lam. (grumichama); larval identity based on numerous previous rearings from the same trees and from which only frater- culus adults emerged (A. Malavasi, personal communication). BRAZIL: Sao Paulo, Uni- versidade de Sao Paulo, XI-1986 (Steck, Malavasi); 5 specimens from culture on ar- tificial medium initiated with adults from Sao Paulo state. BRAZIL: Bahia, Santo Amaro, VI-1988 (Steck, Conceicao); 5 spec- imens from guava. MEXICO: Chiapas, Ta- pachula, Metapa, III-1986 (Carroll); 15* specimens from culture on guava initiated with adults from Tapachula area. COSTA RICA: Puntarenas, Dominical, IV-1986 (Steck, Valerio); 10* specimens from Ter- minalia catappa L. (almendron). VENE- ZUELA: Merida, Merida area, V—-VI-1988 (Steck, Norrbom); 3 specimens from Rubus glaucus Benth. (mora), 3 specimens from Syzygium jambos L. Alston (pomarrosa) and 3 specimens from Coffea arabica L. (café). VENEZUELA: Dto. Federal, Las Caracas, VOLUME 92, NUMBER 2 V-1988 (Rosales); 3 specimens from 7. ca- tappa. A. grandis (Macquart)—BRAZIL: Sao Paulo, Universidade de Sao Paulo, XI-1986 (Steck, Malavasi); 18 specimens from cul- ture on Cucurbita maxima Duch. and 5 specimens from culture on Cucumis melo L. (melon) initiated with adults reared from C. maxima at Sao Roque, S.P. ARGEN- TINA: 11 specimens from USNM, pre- served in alcohol and bearing the following label: ““Anastrepha grandis (Macq.) Pump- kin Argentina. 1-4-37 Houston Tex.-2003 Lot. 37-521.” (See Steck and Wharton 1988.) A. interrupta Stone—USNM: 18 speci- mens preserved in alcohol bearing the fol- lowing label: U.S.A. ““Anastrepha interrupta Homestead, Fla. 3.1.1951 Schoepfia chryso- phylloides berries; 51-997 SPBFLA 109425.” (See Steck and Wharton 1988.) A. leptozona Hendel—MEXICO: Chia- pas, Tapachula vicinity, Huehuetan. III- 1986 (Carroll); 19 specimens from Microph- olis mexicana (Gilly) (baricoco). (Larvae of A. serpentina occurred in low frequency in same collection, but were readily distin- guishable.) A. limae Stone—USNM: 31 specimens preserved in alcohol bearing the following label: ‘PANAMA: Capira 19-20.x.1935 J. Zetek 3552 reared ex Passiflora quadran- gularis.” (See Steck and Wharton 1988.) A, ludens (Loew)—MEXICO: Chiapas, Tapachula, Metapa, IV-1986 (Carroll); 15 specimens from culture on Mangifera in- dica L. (mango) initiated with adults reared from mango in Tapachula area. U.S.A:: Texas, Texas A&M University. IV-1984 (Carroll); 15 specimens from culture on ar- tificial medium. (See Carroll and Wharton 1989.) A. obliqua (Macquart)— MEXICO: Chia- pas, Tapachula, Metapa, IV-1986 (Carroll); 20* specimens from culture on mango ini- tiated with adults reared from Spondias sp. (obo) in Tapachula area. COSTA RICA: Alajuela, F. Baudrit Expt. Stn., [V-1986 335 (Steck, Valerio); 9* specimens from mango. VENEZUELA: Merida, Hwy 7 x Pueblo Nuevo road, VI-1988 (Steck, Norrbom, Holmquist); 5 specimens from mango. BRAZIL: Bahia, Cruz das Almas area, VI- 1988 (Steck, Conceicao); 2 specimens from Averrhoa carambola L. (carambola), 2 spec- imens from Spondias purpurpea L. (caja) and 2 specimens from mango. A. pallens Coquillet-—USNM: 10 speci- mens preserved in alcohol bearing the fol- lowing labels: ““Pseudodacus pallens (Coq.) / A, pallens / laboratory collection Coma berries Pseudodacus pallens Coq. lot no. 35- 19611 FHB/ GVH #35.” A. serpentina (Wiedemann)— MEXICO: Chiapas, Tapachula, Metapa, III-1986 (Carroll); 20 specimens from culture on Manilkara zapota (L.) P. Royen (chico za- pote) initiated with adults from M. zapota, Pouteria sapota (Jacq.) Moore and Stearn (mamey) and Chrysophyllum cainito L. (caimito) from Tapachula area. MEXICO: Veracruz, Los Tuxtlas Biol. Stn., VII-1984 (Steck); 11 specimens from P. sapota. VEN- EZUELA: Aragua, Maracay, V-1988 (Steck, Norrbom, Rosales); 5 specimens from MM. zapota (nispero). BRAZIL: Sao Paulo, Sao Sebastiao, VI-1988 (Amaral); 5 specimens from M. zapota (abrico). A. striata Schin- er—MEXICO: Chiapas, Tapachula, Meta- pa, IV-1986 (Carroll); 20* specimens from culture on guava initiated with adults reared from guava in Tapachula area. COSTA RICA: Cartago, Tres Equis, Hwy 10 be- tween Turrialba and Siquirres, 3-IV-1986 (Steck, Carlson, Valerio); 10* specimens from guava. VENEZUELA: Merida, Meri- da, V-1988, (Steck, Norrbom, Holmquist); 5 specimens from guava. VENEZUELA: Miranda, Guatopo National Park, VI-1988 (Condon); 5 specimens from guava. A. suspensa (Loew)—U.S.A.: Florida, Homestead, TREC-IFAS, I-1985 (Bara- nowski); 30* specimens from culture on ar- tificial medium initiated with adults from southern Florida. 336 In developing the key, complete mea- surements were taken on all specimens as in Steck and Wharton (1988). Those char- acters newly used in this key mostly concern the presence or absence of dorsal spinules on the various segments, and quantitative counts and measurements on the posterior spiracular processes. For convenience, some of the measurement procedures are repeated here as they relate specifically to the use of the key. Only features visible with a dis- secting or compound microscope were ex- amined. Terminology follows Teskey (1981). Oral ridge (ORL) counts and determi- nation of anal lobe shape are taken from whole specimens. Specimens are removed from alcohol and propped in an appropriate position on an alcohol-dampened wad of cotton in a small watchglass. The alcohol evaporates off the surface after a minute or two, and the oral ridges (Fig. 1) become clearly separable and countable. Use of a strong, fluorescent light is recommended; use of an incandescent light requires careful adjustment of the lighting angle. A mini- mum of 80x magnification is necessary for accurate counts On most specimens. Oral ridges are counted along the inner margin adjacent to the oral opening. The terminal upper and lower oral ridges are reduced in size and often difficult to see. The shape of the anal lobes is also best determined just after the alcohol dries off the surface of the whole specimen. (Their shape usually will be still apparent after slide-mounting.) In some species the lobes are almost always obviously bilobed (e.g. /udens, serpentina; Figs. 3, 4); or obviously entire (e.g. suspensa, obliqua; Fig. 6). In other species, such as striata and distincta, the lobes may be wrin- kled or finely grooved, and thus indeter- minate in this respect (Fig. 5). These latter are keyed both ways at the corresponding couplets. Anterior spiracular tubules (ANS) are also counted on whole specimens. Ifthe spiracles are not well exposed on the whole specimen, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON they become so after the specimen is treated in sodium hydroxide (NaOH). Specimens are not perfectly symmetrical; numbers of oral ridges and anterior spirac- ular tubules frequently are unequal on left and right sides. Count data used in the key are the average of the two sides rounded upwards; e.g. a specimen with 10 oral ridges on one side and 11 on the other would be counted as 11 (Fig. 1). (Measurements en- tered into discriminant analysis, however, were not rounded.) If one anal lobe 1s bi- lobed and the other entire, the specimen 1s considered to be bilobed. Dorsal spinules occur in broken, parallel rows. They usually are apparent at 80x magnification with good fluorescent lighting and best seen from a dorsolateral angle. Rows are counted on the dorsum, defined to be that surface bounded by a pair of imag- inary lines drawn lengthwise between the anterior spiracle and posterior spiracle on each side (Fig. 2). Many specimens have rows of dorsal spinules interrupted by a broad, bare hiatus across the medial third of the dorsum. If rows are not visible on whole specimens, one should re-check slide- mounted specimens at 100 x magnification on a compound microscope. For purposes of orientation, dorsal spinules on the second thoracic segment (T2) are those immedi- ately posterior to the insertion of the ante- rior spiracles. Specimens are slide-mounted for all re- maining observations. The body is slit lengthwise along one side from just below an anterior spiracle to just above the anal lobes. Specimens are then left in 10% NaOH overnight at room temperature (or about | hr at 60°C). Afterwards, internal tissues are easily cleared away. The cephalopharyngeal skeleton (CPS) is gently separated from the cuticle and mounted laterally as normally figured (e.g. Steck and Wharton 1988). The cuticle is mounted flat in glycerol (or per- manently mounted in Hoyer’s medium or Euparal). It helps to cut small notches in the VOLUME 92, NUMBER 2 cuticle around the posterior spiracles and anal lobes, and between the anterior spira- cles so the entire specimen will lie flat (see Fig. 7). Thus mounted, rows of dorsal spi- nules can be counted readily. Measurements on the posterior spiracular openings (PSO) and posterior spiracular processes, SP-I and SP-IV (Fig. 8) are made at 400 magnification using an ocular mi- crometer. Use of Nomarski optics provides a 3-dimensional perspective and facilitates counting of trunks and tips of spiracular processes. Measurements and counts are usually made on only one side, right or left, choosing whichever side is best positioned. PSO are measured to the outside edges of the heavily sclerotized rimae; values used in the key for length (LTH) and width (WTH) are the averages of the dorsal and ventral openings. Likewise, number of tips (TIP) and trunks (TRK) is the average of SP-I and SP-IV. The number of tips usually is readily countable. Determination of the number of trunks as clearly separate insertions into the cuticle is sometimes difficult due to orien- tation or crowding. In practice, when the insertion points are obscured, any branch seen as separate beyond about the basal 10% is counted as a trunk. The basal width (BAS) is the distance between outermost trunks at their insertion points; again, the average of SP-I and SP-IV is used. Throughout development of the key we tried to use ratios of measurements on re- lated structures (e.g. basal width of SP-I and SP-IV to length of PSO) as key characters to avoid biases resulting from unusually large or small specimens (related perhaps to type of host fruit utilized). The user of this key will note, however, that numerous couplets rely on absolute measurements, e.g. length of PSO, distal width of anterior spiracles, etc. For these latter characters, ratios did not prove useful in distinguishing among species, whereas the absolute measurements did. Very little has been published on intra- specific geographical variation in larval 337 characters. Some of the complexity of this key arises from such variation. It is possible that other populations besides those sam- pled here will fall outside the key ranges. In our /udens, for example, the range of lengths of PSO for Weslaco specimens did not over- lap with the range for specimens from Ta- pachula. (The key does not use this partic- ular character in arriving at an identification of /udens.) Larvae of A. fraterculus may present an especially thorny problem, since there are long-standing, unresolved ques- tions about the occurrence of cryptic species in different geographical regions. Larvae of other species also display non-overlapping states for various characters among assorted populations. We foresee more such prob- lems arising as other geographical regions are sampled. The key works strictly on the basis of morphological characters. When informa- tion on host fruit and geographic origin of specimens are available, the task of iden- tification 1s considerably simplified. Table 1 summarizes host and distribution infor- mation for the 13 species included in the key. Within-species variability was extensive, and few, if any, single characters could re- liably be used to diagnose species. An ad- equate number of specimens was examined in most cases to allow us to delimit ranges in which most key character states fell. We aimed for a key which would allow accurate identification for 95% or more of all spec- imens examined. Thus, we did not include numerous additional couplets to accom- modate those specimens which displayed extreme character state values. In view of the difficulties, we would consider any de- termination based on a single specimen to be suspect. When several specimens of a collection are examined, the likelihood of a correct determination is greatly increased. In some cases, there was so much overlap in key character states between species, that a simple bifurcating key became unman- ageable. This was true for striata/bistrigata PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distnbution southern Brazil, northern Peru southern Texas to South America southern Texas to South America South America, Panama southern Florida, Bahamas Central and South America Venezuela, Panama, Texas southern Texas to Costa Rica Mexico to South America, Caribbean Texas to Honduras southern Texas to South America Mexico to South America 338 Table 1. Host plants and geographical distributions of Anastrepha species. Species Host Plants bistrigata Psidium distincta Inga fraterculus numerous grandis cucurbits interrupta Schoepfia leptozona Chrysophyllum, Pouteria, Micropholis limae Passiflora ludens numerous obliqua numerous pallens Bumelia serpentina numerous striata numerous suspensa numerous and fraterculus/obliqua/suspensa. Multi- variate statistics were employed to discrim- inate among species at the corresponding couplets. The data were analyzed using SAS (1988). Observations of potentially useful characters were first subjected to univariate analysis to check for normality; non-nor- mally distributed measurements were trans- formed as needed. Characters were then subjected to stepwise discriminant analysis (Lachenbruch 1975). The significance level specified to enter and to keep a character was 0.05. Those characters retained as use- ful discriminating factors in the stepwise analysis were incorporated into discrimi- nant analysis to develop a model for ac- tually identifying specimens. Classification results were cross-validated by a jackknifing technique. The original models were also re- tested using sets of new observations on ad- ditional specimens. Specimens marked with asterisks in the preceding collections list were used to develop the discriminant models. Results of stepwise discriminant analysis are presented in Table 2, in which relevant statistics are shown for only those charac- ters which contributed significantly to the discriminant function. The complete sets of characters used for each stepwise discrim- inant analysis are as follows (significant variables are indicated by asterisks): Cou- Greater Antilles, Bahamas, southern Florida plet 4’—ANS*, TRK*, log,.(BAS)*, ORL, LTH, TIP, squareroot((WTH), RTOI1 (=LTH/WTH), log,,.(RTO2) (where RTO2 = TIP/TRK), and log,,.(RTO3) (where RTO3 = BAS/LTH); couplet 14’—ORL*, ANS*, LTH*, log,.(BAS)*, log,.(RTO2)*, TIP, TRK, RTO1, log,,(RTO3), square- root(WTH); couplet 14’A and couplet 14'B—same characters as 14’, but with dif- ferent significant variables; couplet 15— log,o(BAS)*, log,)(RTO2)*, LTH, TIP, TRK, RTO1, log,,(RTO3), squareroot(WTH). Thus, for example, at couplet 4’, ten char- acters (some transformed) were analysed for discriminating striata and bistrigata. Only three characters (ANS, TRK and log,,(BAS), indicated with asterisks, contributed signif- icantly to the discriminant function. The discriminant models presented in Table 3 can be used to identify individual specimens which key to the corresponding couplet. The character values for a given specimen are substituted into the equation. Whether the equation yields a positive or negative value (except couplet 14; see be- low) indicates to which species the specimen is most likely to belong. Consider, for ex- ample, a hypothetical specimen which keys to couplet 4’ and has 13.5 anterior spiracles (13 on one side, 14 on the other; note that values used in discriminant functions are VOLUME 92, NUMBER 2 Table 2. Stepwise discriminant analysis results. Variable* Step Enter Remove Partial R? F Prob > F Couplet 4’: striata vs bistrigata 1 log,,BAS - 0.388 28.49 0.0001 2 ANS — 0.175 9.32 0.0038 3. TRK = 0.167 8.64 0.0053 Couplet 14’: suspensa vs fraterculus vs obliqua 1 LTH _ 0.548 49.64 0.0001 2 log,o.BAS — 0.391 25.99 0.0001 3 JORE = 0.294 16.68 0.0001 4 log,.RTO2 — 0.315 18.14 0.0001 5 ANS _ 0.120 5.32 0.0068 Couplet 14’A: fraterculus vs obliqua ibe 19 0) 3 - 0.478 58.62 0.0001 2 log,oBAS — 0.242 20.11 0.0001 3 ANS - 0.110 7.68 0.0074 Couplet 14'B: suspensa vs obliqua 1D 3 i) BI — 0.598 65.57 0.0001 2 DIP = 0.244 13.91 0.0006 3) (ORL - 0.137 6.69 0.0133 4 ANS — 0.141 6.74 0.0130 Couplet 15: fraterculus vs suspensa apie - 0.482 62.35 0.0001 2 log,oBAS = 0.159 12.52 0.0007 3 log, ,.RTO2 _ 0.113 8.28 0.0054 4 — TIP 0.012 0.81 0.3729 * Abbreviations: ANS, number tubules on anterior spiracles; BAS, basal width of posterior spiracular pro- cesses; LTH, length posterior spiracular opening; ORL, number oral ridges; TIP, number tips on posterior spi- racular processes; TRK, number trunks on posterior spiracular processes; WTH, width posterior spiracular opening; RTO1, ratio LTH to WTH; RTO?, ratio TIP to TRK; RTO3, ratio BAS to LTH. All measurements in um. not rounded), 18 trunks, and a PSP basal width of 46.8 um (log;) = 1.67). When these values are substituted into the couplet 4’ equation, the calculation yields a value of +2.79. A positive result indicates that the specimen is striata; and, using the cross- validation error rate from Table 4, one would conclude that the likelihood of error is 0.231. In the case of couplet 14, a simple positive or negative result is not possible since three species are involved. A discrim- inant function is provided for each of the 339 three species. Character values for an un- known specimen are substituted into each of the three equations; whichever yields the highest value (C) indicates the most likely identification. Because the natural distri- butions of fraterculus and suspensa do not overlap, couplets 14’ and 15 might not rep- resent likely sets of alternatives. Therefore, we also provide discriminant analyses for the pairwise comparisons of fraterculus and obliqua (couplet 14'A), which overlap throughout mainland Central and South America, and obliqua and suspensa (couplet 14’B) which overlap in the Caribbean (Ta- ble 3). The performance of the classification rule was examined using three error rates (Table 4): (1) the apparent error rate (errors in clas- sifying the original specimens using the clas- sification rule calculated from measure- ments on those specimens); (2) the error rate from cross-validation using a jackknifing technique; and (3) the error rate in classi- fying a different set of test specimens. The apparent error rate underestimates the true error rate, although this bias is reduced if the sample size 1s large enough. The cross- validation method using the jackknife tech- nique is almost unbiased (Lachenbruch 1975, Panel... 1989). The jackknife meth- od gives an assessment of the true proba- bility of misclassification of additional spec- imens taken from the original populations. The error rate from the test data set indi- cates the robustness of the classification rule when applied to other populations of spec- imens. The classification results indicated by the apparent and cross-validation errors for each couplet of Table 4 are very similar indicating that sample sizes were adequate for developing each of the discriminant models. The model for couplet 4’ performed poorly for the test specimens of bistrigata, probably due to the fact that the few test specimens available were in poor condition. Also, models 14’, 14’B and 15 fared rela- tively poorly for suspensa test specimens. This indicates that the data base for sus- 340 Table 3. Linear discriminant functions. Couplet 4’: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 23.5(log,,BAS) — 0.75(ANS) — 0.63(TRK) — 15.00 > 0 striata Couplet 14’: < 0 bistrigata fraterculus 186.07(log,,BAS) + 13.01(ORL) + 39.58(log;.~RTO2) + 8.63(ANS) + 1.10(LTH) — 294.42 = C, suspensa —151.06(log,,BAS) + 17.23(ORL) — obliqua Couplet 14’A: 5.62(log,,RTO2) + 8.51(ANS) + 1.25(LTH) — 285.19 =C, 161.23(log,,BAS) + 14.17(ORL) + 27.96(log,,RTO2) + 9.77(ANS) + 1.58(LTH) — 326.32 =C, 22.70(log,,BAS) — 0.45(LTH) — 0.99(ANS) + 24.00 > 0 fraterculus Couplet 14’B: < 0 obliqua 1.40(ANS) + 0.60(TIP) + 0.24(LTH) — 2.45(ORL) — 30.93 > 0 obliqua Couplet 15: < 0 suspensa 24.20(log,)BAS) + 26.67(log,;)RTO2) — 40.52 > 0 fraterculus < 0 suspensa pensa should be augmented with specimens from additional populations. As noted previously, it has not been pos- sible to construct a key to accommodate all the variability observed in our samples. However, the accuracy of the key is very high. The percentage of study material which keyed correctly (*before discriminant anal- ysis) was as follows (sample sizes for each species in parentheses): *bistrigata (30)/ striata (40)—100%, distincta (30)—97%, *fraterculus (72)/obliqua (40)/suspensa (40)— 100%, grandis (34)— 100%, interrup- ta (18)—100%, leptozona (19)—95%, limae (31)—100%, /udens (30)—100%, pallens (10)—100% and serpentina (41)—98%. The generic description and diagnosis presented below are based on published de- scriptions by other authors (especially Kan- dybina 1977) and on additional unpub- lished observations of our own. With the exception of holarctic species of Rhagoletis, Table 4. Error rates of classification by discriminant analysis. Species Apparent Error Crossvalidation Error Test Error (sample size) Couplet 4’: striata 0.192 0.231 0.125 (n = 8) bistrigata 0.046 0.091 0.500 (n = 4) Couplet14’: Sraterculus 0.026 0.026 0.222 (n = 9) suspensa 0.158 0.158 0.429 (n = 7) obliqua 0.148 0.148 0.167 (n = 6) Couplet 14’A: Larvae originating in Central or South America Sraterculus 0.025 0.025 0.214 (n = 14) obliqua 0.143 0.143 0.182 (n= 11) Couplet 14'B: Larvae originating in Caribbean obliqua 0.074 0.074 0.000 (n = 7) suspensa 0.000 0.000 0.286 (n = 7) Couplet 15: fraterculus 0.136 0.136 0.111 (n= 9) suspensa 0.172 0.172 0.333 (n = 9) VOLUME 92, NUMBER 2 the larvae of relatively few species of fruit- infesting Tephritidae have been adequately described. Larvae of numerous Diptera families may be found in fruits (Keifer 1930), though only a few of these would be found in ripe, healthy fruits suitable for human consumption. Be- sides Tephritidae, only a few species of Lon- chaeidae are likely to be encountered. These are readily distinguished from tephritids by the appearance of the posterior spiracles. In lonchaeids they comprise a pair of promi- nent stumps, round, black, and heavily sclerotized, projecting from the caudal seg- ment; rimae of spiracular openings are at nearly right angles. In tephritid fruit flies, the spiracular openings comprise a pair of three elongate slits, nearly flush with the body surface; their rimae are sclerotized and golden-brown, and their long axes are roughly parallel. Among Tephritidae, other fruit-infesting genera which may be en- countered in the Neotropics and subtropics include endemic Rhagoletis and Toxotry- pana and introduced species of Ceratitis and Dacus. Rhagoletis generally are distinctive in possessing prominent, chitinized teeth or “stomal guards” adjacent to the oral open- ing and strongly developed intermediate and ventral tubercles on the caudal segment (see Phillips 1946, Kandybina 1977), neither of which are seen in Anastrepha. Toxotrypana is largely restricted to papaya, larvae are very large, and all caudal sensilla are greatly reduced (see Heppner 1986). Ceratitis and Dacus both may be recognized by the pres- ence on the caudal segment of a distinct crescent-shaped ridge connecting, or just dorsad of, sensilla I] and I2 (see Heppner 1985, Elson-Harris 1988) and conspicuous dental sclerites (see Exley 1955). The caudal ridge is lacking and dental sclerites usually are not seen in Anastrepha larvae. The di- agnostic and key characters are for third in- stars only, and cannot be applied to earlier instars. Sections I to III of the key will elim- inate most specimens for which the key is not intended. ANASTREPHA GENERIC DESCRIPTION (THIRD INSTAR) Body elongate, 4~7 times longer than wide, pointed anteriorly. Integument thin, smooth, colorless. Spinules separate, conical, in short, staggered rows (occasionally flat, blunt, ba- sally connected in short rows); occurring in discrete fusiform areas ventrally on all ab- dominal segments; also dorsally in bands on T1, T2, usually T3, present or absent on abdominal segments. Antennal and maxil- lary sensory organs on well-developed ce- phalic lobes above mouthhooks. Antennal sensory organ appearing 2-segmented with basal sclerotized, cylindrical collar and api- cal hemispherical to conical sense organ. Maxillary sensory organ cylindrical, trun- cate, apically bearing peg-shaped sensoria. Oral ridges 7-30 per side, well developed. Stomal organ minute cluster of sensilla borne distally on large, simple, oblong lobe ante- rior to mandible. Sclerotized stomal guards absent. Cephalopharyngeal skeleton with clearly separate sclerites as follows: Man- dible falciform (occasionally uncurved and blunt), single-toothed, length to height ratio (lateral view) about 1.0-1.5, heavily scler- otized; dental sclerite apparently lacking or small and inconspicuous; epipharyngeal and labial sclerites present; hypopharyngeal sclerite in dorsal view H-shaped, width at bridge about equal to length (ratio, 0.75- 1.25), and length in lateral view about twice height, anterior forks heavily sclerotized; parastomal bar long and thin, usually bent medially, 0.75-1.0 times length of hypo- pharyngeal sclerite; anterior sclerite irreg- ularly developed and shaped; dorsal cornua narrowly connected at dorsal bridge: ventral cornu trough-shaped, with 7 pharyngeal ridges. Anterior spiracle with distinct, cy- lindrical trunk; sharply flared and bilobed apically with numerous (9-37) tubules. Caudal segment more or less smooth and rounded; intermediate sensilla I] and I2 on relatively developed tubercles; remaining sensilla (D1, D2, 13, L1, V1-V3) on weak Rigel: Fig. 2. lines drawn between anterior spiracles and posterior spiracles; ds, dorsal spinules in 34 rows on segment T2 (note hiatus in rows of spinules across mid-dorsum of segment T3); Tl and T2, first and second thoracic segments. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Oral ridges (or), A. suspensa. or undeveloped tubercles. Posterior spira- cles located above horizontal midline; with three slits having well-developed rimae and trabeculae. Anal lobes entire or bifid; en- circled by spinules. KEY TO ANASTREPHA LARVAE (THIRD INSTAR) I Posterior spiracles prominently raised from the body surface; or most body segments with conspicuous setae or processes; or posterior spiracular openings sinuous Posterior spiracles nearly flush with body sur- face; tubercles, if present, on caudal segment only; posterior spiracular slits elongate or oval II With prominent chitinized teeth (stomal guards) adjacent to oral opening, and strongly developed tubercles on caudal segment; or, with crescent-shaped ridge between sensilla Il and I2 on caudal segment and conspicuous dental sclerite; or, larva taken from papaya, very large (more than 12 mm long), caudal tubercles lacking and caudal sensilla all very reduced tnt re es Lacking stomal guards; caudal tubercles at most moderately developed; lacking crescent not Tephritidae I See not Anastrepha ridge on caudal segment; dental sclerite lack- ing or inconspicuous; not normally attacking papaya; at least caudal sensilla I] and I2 con- spicuous Il Anterior spiracle absent, and posterior spi- racle with only 2 openings; and/or, mandible with well-developed subapical tooth, and posterior spiracular openings short (less than 55 um) and oval; and body short and thin (less than about 6.0 mm long and 1.0 mm diameter) Anterior spiracle present; posterior spiracle with 3 elongate openings at least 65 um long; mandible without subapical tooth; body length and diameter greater than about 6.0 mm by 1.0 mm IV. Anastrepha, third instar Dorsal spinules present on two or more ab- dominal segments RAE rth orn sah Dorsal spinules present on Al, but not be- yond . 4 5 stegei bt dts asicctayerelel seas ce Reo Dorsal spinules absent on all abdominal seg- ments = BE a IETS oes Dorsal spinules separate, conical; in fewer than 5-6 rows on T2 and T3 (except /imae). Pos- terior spiracular processes SP-I and SP-IV with average of 6 or more trunks and bristle length '4 or more times length of spiracular opening Dorsal spinules connected basally in flat, sawtooth pattern, blunt-tipped; in 8 or more Dorsal surface of A. /imae. Abbreviations: as, anterior spiracle, D, dorsum bounded by imaginary Ill not 3rd instar Ini tN VOLUME 92, NUMBER 2 343 Figs. 3-4. Bifid anal lobes, A. serpentina. Indeterminate, grooved anal lobes, 4. distincta. Entire anal lobes, 4. fraterculus. Fig. 5. Fig. 6. rows on T2 and T3, at least 3 rows on Al to A4, 1-4 rows (often with medial hiatus) on T2 with 2-4 rows; T3 with 1-3 rows, often with medial hiatus 6 A5. SP-Iand SP-IV with average of 5 or fewer 5'. Dorsal spinules well-developed on Al, in 2 trunks, and bristle length about '5 times length or more rows, without medial hiatus; both of posterior spiracular opening . _ pallens T2 and T3 with 5-6 rows . limae (part) 3. Anterior spiracle with 12-23 tubules; distal 6. Anterior spiracle with 14-22 tubules. Pos- Width Ol 9=O0:37 MMi sige sees en ye ee ame 4 terior spiracular opening 94-130 um long. 3’. Anterior spiracle with 28-37 tubules; distal SP-I and SP-IV with average of 7-13 trunks width 0.43-0.61 mm NR aie grandis and 17-28 tips . ludens (part)* 4. SP-I and SP-IV with average of 8 12 trunks 6’. Anterior spiracle with 10-13 tubules. Pos- and 12-21 tips; basal width 12-19 um, 0.1- terior spiracular opening 72-84 um long. SP-I 0.2 times length of spiracular opening. Dorsal and SP-IV with average of 4-7 trunks and 5— spinules absent on A3 ............ limae (part) 11 tips interrupta (part)* 4’. SP-Land SP-IV with average of 13-23 trunks 7. Oral ndges 7— HH Leap and 23-49 tips; basal width 19-67 um, 0.2- 7’. Oral ridges 12 or more ... 16 0.5 times length of spiracular opening. Dorsal 8. Anterior spiracle with 15 or more tubules 9 spinules usually present on A3 ......... striata 8'. Anterior spiracle with 9-14 tubules . 14 Rien ee eae deer er eee ae an bistrigata) 9. Anal lobe bifid Peer 0) Bie Cee ee (See Table 3, couplet 4’) 9’. Anal lobe entire euchaoeen, Ll 5. Dorsal spinules weakly developed on A1, in 10. Posterior spiracular opening 74-96 um long. only | row, usually with broad medial hiatus; SP-I and SP-IV with average basal width of 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Slide-mounted cuticle: AL, anal lobes; PS, posterior spiracles; T2 and T3, second and third thoracic segments; Al, first abdominal segment. Fig. 8. Posterior spiracle (right side): SP-I to SP-IV, dorsal to ventral spiracular processes; BAS, basal width of spiracular process; LTH, spiracular opening length; WTH, spiracular opening width. 12-22 um, 0.1-0.2 times length of spiracular 16. Anterior spiracle with 15 or more tubules .. 17 opening; average of 7-11 trunks .......... 16’. Anterior spiracle with 9-14 tubules ....... 20 PASO AREAS Son Ooo as poate serpentina (part) 17. SP-I and SP-IV with average basal width of 10’. Posterior spiracular opening 103-122 um 10-24 um, 0.1-0.3 times length of spiracular long. SP-I and SP-IV with average basal width OPCDING ) ff yas sacra acecmnsts eis eee eee 18 of 29-58 um, 0.3-0.5 times length of spirac- 17’. SP-I and SP-IV with average basal width of ular opening; average of 10-17 trunks ...... 24-65 um, 0.3-0.6 times length of spiracular : Rees PR rs Aoki SEE See distincta (part) OPENING ey here. careers distincta (part)* 11. Anterior spiracle with 17 or more tubules .. 12 18. Anal lobe bifid ......................... 19 11’. Anterior spiracle with 9-16 tubules : 13° 18’. ~Analilobeventire!: ee eke leptozona (part) 12. SP-I and SP-IV with average basal width of 19. Dorsal spinules present on T3 (rows often 29-58 um, 0.3-0.5 times length of spiracular with medial hiatus). Posterior spiracular opening Sa HOO S Hot distincta (part) opening 40-54 um long ......... ludens (part) 12’. SP-I and SP-IV with average basal width of 19’. Dorsal spinules absent on T3. Posterior spi- 14-20 um, 0.1-0.2 times length of spiracular racular opening 31-40 um long........... OPENINGS emer ence cir jeetn, WepLOzONa\(part) «-_ss(“$ "ee Seer eeeer nce serpentina (part) 13. SP-land SP-IV with average of 10-17 trunks 20. Dorsal spinules present on T3 (rows may have and 24-37 tips. Anterior spiracle distal width medialshiatus) eee. ae ere eer 21 OR — D7 Sh nM pert cee erro .. obliqua (part) 20’. Dorsal spinules absent on T3_.. serpentina (part) 13’. SP-I and SP-IV with average of 6-10 trunks 21. SP-I and SP-IV with average of 15 or more and 13-23 tips. Anterior spiracle distal width tips and 7 or more trunks. Up to 16 oral ridges 260=335 jum Feast. .. leptozona (part) SoC ne Re re CR Aeicls abicox 22 4 oe Analllobei bifid: <.s¢.c0--apre eee ee ee 15 21’. SP-Iand SP-IV with average of 5-11 tips and 14’. ‘Analitobeventire: 22 .2eu. eee suspensa (part) 7 or fewer trunks. Up to 20 oral ridges .... San OOS On Oe Gols aoe fraterculus (part) SEE Cee Eee etn SeRIILernuptal (part) rs SCOT ise COS SISEInIA obliqua (part) 22. Anal lobe bifid. Anterior spiracle distal width SA atta (See Table 3, couplets 14’, 14’A, 14’B) 260-347 um. Up to 16 oral ridges ........ l'53) Anteriorspiraclewith,9—13'tubulestandiSR=I= eee eee eee eee ludens (part)* and SP-IV with average of 11 or more trunks 22'. Anal lobe entire. Anterior spiracle distal width eR AA Ha SOHN fraterculus (part) 161-248 um. Up to 12 oral ridges ........ Bete hs isobar eiee suspensa (part) Be ene oe Pen: Bee SUSDEMSA (Dane) tg Shia o oes A TOS e ee (See Table 3, couplet 15) 15’. Anterior spiracle with 13-14 tubules, and SP-I $$ and SP-IV with average of 11 or fewer trunks * indicates that 10% or fewer of the individuals of a ES re ge RM ee rat er te At serpentina (part) given species key to the corresponding couplet. VOLUME 92, NUMBER 2 ACKNOWLEDGMENTS The help of R. A. Wharton (Texas A&M University) in all aspects of initiating and carrying through this and related studies of tephritid immatures is gratefully acknowl- edged. Valuable assistance in acquiring specimens and providing facilities and lo- gistical support was given by the personnel of Programa MoscaMed, especially P. Liedo-F. and A. Schwarz-G. (Tapachula, Mexico); E. Morales-M. and J. Valerio-S. (Organization Internacional Regional San- idad Agropecuaria, San José, Costa Rica); L. F. Jiron (Universidad de Costa Rica, San José); A. Malavasi and family (Universi- dade de Sao Paulo, Brazil); A. Pinto da Cun- hia and J. L. Conceigao (EMBRAPA, Cruz das Almas, Brazil); C. J. Rosales (Univer- sidad Central, Maracay, Venezuela); O. Holmquist and A. Briceno (Universidad de los Andes, Merida, Venezuela) and M. Con- don (Smithsonian Institution). A. L. Norr- bom (Systematic Entomology Laboratory, USDA) provided collection and identifica- tion assistance. S. Blanchard (Statistical Consulting and Analysis Services, USDA) advised us on statistics and performed the analyses. R. A. Wharton, A. L. Norrbom, I. M. White (CAB International Institute of Entomology) and A. Freidberg (Tel Aviv University) provided many useful sugges- tions for improving the manuscript. J. Plas- kowitz (Systematic Botany, Mycology & Nematology Laboratory, USDA) assisted with SEM photographs. R. Brittingham and W. Denny (Plant Germplasm Quarantine Center, APHIS) graciously provided access to their facilities. This research was funded in part by USDA/OICD project No. 58- 319R-5-016. LITERATURE CITED APHIS. 1987. List of intercepted plant pests. Fiscal year 1986. U.S. Department Agric., Animal Plant Health Inspection Serv., Plant Protection Quar- antine, APHIS 82-13. Hyattsville, Md. Berg, G. H. 1979. Pictorial key to fruit fly larvae of 345 the family Tephritidae. Organismo Internacional Regional de Sanidad Agropecuaria. San Salvador. Carroll, L. E.and R. A. Wharton. 1989. Morphology of the immature stages of Anastrepha ludens (Loew) (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 82: 201-214. Elson-Harris, M. M. 1988. Morphology of the im- mature stages of Dacus tryoni (Froggatt) (Diptera: Tephritidae). J. Aust. Ent. Soc. 27: 91-98. Exley, E.M. 1955. Comparative morphological stud- ies of the larvae of some Queensland Dacinae (Trypetidae, Diptera). Queensland J. Agric. Sci. 12: 119-150. Greene, C. T. 1929. Characters of the larvae and pupae of certain fruit flies. J. Agric. Res. 38: 489- 504. Heppner, J. B. 1984. Larvae of fruit flies. I. Anastre- pha ludens (Mexican Fruit Fly) and Anastrepha suspensa (Caribbean Fruit Fly) (Diptera: Tephriti- dae). Fla. Dept. Agric. Consumer Serv., Div. Plant Industry. Entomol. Circ. 260. 4 pp. Heppner, J. B. 1985. Larvae of fruit flies. II. Ceratitis capitata (Mediterranean fruit fly) (Diptera: Te- phritidae). Fla. Dept. Agric. Consumer Serv., Div. Plant Industry. Entomol. Circ. 273. 2 pp. Heppner, J. B. 1986. Larvae of fruit flies. HI. Toxo- trypana curvicauda (Papaya fruit fly) (Diptera: Te- phritidae). Fla. Dept. Agric. Consumer Serv., Div. Plant Industry. Entomol. Circ. 282. 2 pp. Kandybina, M.N. 1977. Larvae of fruit flies (Diptera, Tephritidae). Opredeliteli Po Faune SSSR. No. 114. Leningrad: Nauka, Leningradskoe otd-nie, 1977. (Translated from Russian by Mrs. Geti Saad for OICD, LGU, USDA, 1987). Keifer, H. H. 1930. Synopsis of the dipterous larvae found in California fruits. Monthly Bull. Dept. Agr. Calif., pp. 574-581. Lachenbruch, P. A. 1975. Discriminant Analysis. New York, Hafner Press. Norrbom, A. L. and K. C. Kim. 1988. A list of the reported host plants of the species of Anastrepha (Diptera: Tephritidae). U.S. Dept. Agric., APHIS- PPQ. Hyattsville, Md. 114 pp. Panel on Discriminant Analysis, Classification, and Clustering. 1989. Discriminant analysis and clustering. Stat. Sci. 4: 34-69. Phillips, V. T. 1946. The biology and identification of trypetid larvae (Diptera: Trypetidae). Amer. Entomol. Soc., Mem. No. 12, 177 pp. SAS Institute. 1988. SAS/Stat User’s Guide, Release 6.03 Edition. SAS Institute Inc., Cary, North Car- olina. Steck, G. J. and A. Malavasi. 1988. Description of immature stages of Anastrepha bistrigata (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 81: 1004—- 1009. 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Steck, G. J. and R. A. Wharton. 1988. Description of immature stages of Anastrepha interrupta, A. limae, and A. grandis (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 81: 994-1003. Teskey, H. J. 1981. Morphology and terminology— larvae, pp. 65-88. Jn J. F. McAlpine, B. V. Pe- terson, G. E. Shewell, H. J. Teskey, J. R. Vocke- roth, and D. M. Wood (coords.), Manual of Nearc- tic Diptera, Vol. 1. Research Branch, Agriculture Canada, Monograph 27. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 347-350 A NEW SPECIES OF THE MINUTE PREDACEOUS MIDGE GENUS NANNOHELEA FROM SRI LANKA (DIPTERA: CERATOPOGONIDAE) WILLIAM L. GROGAN, JR. AND WILLIS W. WIRTH (WLG) Department of Biological Sciences, Salisbury State University, Salisbury, Mary- land 21801; (WWW) Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, Florida 32606. Abstract. — Nannohelea tamil, a new species of minute predaceous midge from Sri Lanka is described and illustrated. A key to the extant species of Nannohelea is provided. Key Words: Sri Lanka, Oriental Grogan and Wirth (1980) proposed the minute predaceous midge genus Nannohe- lea composed of 3 species: bourioni (Clas- trier) (1961), as type-species, from France and Algeria (Palearctic); fuscipennis (To- kunaga) (1964), from New Guinea and Ma- laysia (Oriental and Australasian); and c/as- trieri Grogan and Wirth (1980), from Columbia (Neotropical). Recently, Szad- ziewski (1988) described 2 new species from Eocene Baltic amber (ca. 40 million years old), suggested that the genus is of Laurasian origin, and that it probably migrated to South America during the Tertiary. We recently obtained 4 specimens of Nannohelea collected by Ginter Ekis in Sri Lanka during 1973 that represent a new species apparently most closely related to N. fuscipennis. In addition to describing this new species, we also present a key to the extant species of Nannohelea. For an explanation of general ceratopo- gonid terminology, see Downes and Wirth (1981); for special terms dealing with pre- daceous midges in the tribe Ceratopogonini, see Wirth and Grogan (1988). The types of this new species will be deposited in the National Museum of Natural History, Washington, D.C. (USNM). tN Diptera, Ceratopogonidae, Nannohelea, predaceous midges, new species, Key TO THE EXTANT SPECIES OF NANNOHELEA Females Males ee 4 Anal lobe of wing poorly developed, wai Bane of 6-10 short capitate setae; small species, wing length 0.68-0.74 mm; Palearctic Region bourtoni (Clastrier) Anal lobe of wing well developed, with fringe of normal slender setae; very small species, wing length 0.47-0.49 mm, Oriental and Australa- sian Regions ee 3 . Eyes contiguous; wing with 3 vein R243 much shorter than r-m crossvein, radial cell narrow, cubital fork obsolete on distal 7 fuscipennis (Tokunaga) Eyes narrowly separated; wing with vein R2+3 as long or longer than r-m crossvein, radial cell rounded, cubital fork obsolete only at extreme tip at wing margin tamil, new species Flagellum with 8 flagellomeres; small species, wing length 0.50-0.55 mm bourioni (Clastrier) Flagellum with 7 flagellomeres; very small species, wing length 0.37-0.43 mm 5 Eyes contiguous; flagellum with only distal 2 flagellomeres elongated, antennal ratio 0.82— 0.85 fuscipennis (Tokunaga) Eyes separated; flagellum with distal 3 flagello- meres elongated, antennal ratio 1.38 or greater Aedeagus with long slender basal arms; flagel- lum with last flagellomere bearing a single api- cal trichoid sensilla, antennal ratio 1.38-1.39 . tamil, new species 348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON e Fig. 1. Nannohelea tamil. a, female flagellum; b, male flagellum; c, female wing; d, male palpus; e, aedeagus. Scale bars = 0.03 mm (a-b), 0.1 mm (c), 0.01 mm (d-e). — Aedeagus with short stout basal arms; flagel- lum with last flagellomere bearing 2 trichoid sensilla, antennal ratio 1.93 J (Glcrss clastrieri, Grogan and Wirth Nannohelea tamil Grogan and Wirth, New SPECIES (Fig. 1) Diagnosis. — Differs from its congeners by the following combination of characters: very small size (female wing length 0.47 mm, male wing length 0.41 mm); wing with well developed anal lobe, vein R2+3 in nearly a straight line with and as long or longer than r-m crossvein, radial cell rounded; eyes separated; male aedeagus with long slender basal arms and rounded tip. Holotype female.— Head: Dark brown. Eyes pubescent, narrowly separated. Anten- na with dark brown pedicel; flagellum (Fig. la) light brown with darker brown bands on midportion of distal 5 flagellomeres, proximal 8 flagellomeres bearing a pair of stout subapical trichoid sensilla bent at 90° angle near base, flagellomere | with a pair of apical sensilla basiconica, distal 5 flag- ellomeres with subapical pair of sensilla basiconica that are also bent at base; lengths of flagellomeres in proportion of 21-11-12- 11-11-11-11-12-13-13-18-18-20; antennal ratio 0.82. Clypeus with 2-3 pairs of sub- marginal setae. Palpus similar to that of male (Fig. lc) but somewhat disoriented due to mounting; segment 2 very broad with large deep sensory pit bearing capitate sensilla. Mandible very small with 4 small curved teeth. Thorax: Dark brown. Scutum without an- terior spine or humeral pits, covered with short fine pubescence and a few large setae; scutellum with 4 bristles; postscutellum VOLUME 92, NUMBER 2 highly produced. Legs light brown; femora moderately slender, unarmed, stoutest on fore leg; tibiae slender, unarmed, stoutest on hind leg, hind tibia with comb of 6 setae and bifurcated or trifurcated spur; tarsi with apical spines on tarsomeres 1-3 of mid leg only, tarsomere | of hind leg with well de- veloped palisade setae, tarsomeres 4 cylin- drical, tarsomeres 5 slender and bearing small equal sized simple claws. Wing (Fig. lc) infuscated dark brown, membrane covy- ered with microtrichia, macrotrichia con- fined to anterior margin at tip and a few on radial sector; fringe moderately long, ex- tending from base of costa to halfway up anal lobe; anal lobe well developed; a rounded radial cell present; r-m crossvein rather long and set at an oblique angle; vein R2+3 nearly in line with and as long as r-m crossvein; vein M1 straight, obsolete only near wing margin, M2 absent; cubitus fork- ing at level of r-m crossvein, obsolete only near wing margin; anal veins absent; costal ratio 0.59; wing length 0.47 mm, breadth 0.27 mm. Halter light brown. Abdomen: Brown. Two large spermathe- cae, partially collapsed, apparently ovoid with long slender necks, one of which is 0.06 mm in length including the neck. Allotype male.—Smaller but similar to holotype female with the following notable differences: Antennal flagellum (Fig. 1b) re- duced to only 7 flagellomeres; flagellomere 1 with only a single apical sensilla basicon- ica, flagellomere 2 with only a single straight trichoid sensilla, flagellomere 5 with a pair and flagellomere 6 with a single sensilla bas- iconica, flagellomere 7 with a single long straight sensilla trichodea; lengths of flag- ellomeres in proportion of 31-15-13-13-40- 30-29; antennal ratio 1.38. Palpus (Fig. 1d) 3 segmented: segments in proportion of 12- 33-19; palpal ratio 1.32. Mandible vestigial, without teeth. Wing length 0.41 mm, breadth 0.24 mm; costal ratio 0.59. Gen- italia very small, distorted due to mounting: sternite 9 very short, caudal margin straight; tergite 9 apparently very short, extending 349 only 2 the length of gonocoxite. Gonocoxite straight, very short; gonostylus longer than gonocoxite, tapering slightly distally to a narrow rounded tip. Aedeagus highly dis- torted, a reconstruction of that of paratype shown in Fig. le; basal arm slender; distal portion with rounded tip. Parameres not discernible in holotype, but visible in para- type as a slender bridge that connects the bases of the gonocoxites. Type material.—Holotype female, allo- type male, one female and one male para- type labeled “SRI LANKA: Rat. Dist., Gil- imale Lumber Mill, 7 VIII 1973, 115 feet, Ginter Ekis” (USNM). Distribution. —Sri Lanka. Etymology.—The specific epithet, tamil, is a reference to the small Dravidian people that inhabit the forests of Sri Lanka. Remarks.— Nannohelea fuscipennis (To- kunaga) most closely resembles this new species but differs from it in having contig- uous eyes, a wing with a smaller radial cell and the cubitus vein obsolete basally and on the distal 74, a more slender 2nd palpal segment, male antenna with only distal 2 flagellomeres elongated and the aedeagus is pointed apically. Nannohelea clastrieri Gro- gan and Wirth, known only from a single male from Colombia, differs in having an aedeagus with short basal arms and the last flagellomere has 2 apical sensilla trichodea. Nannohelea bourioni (Clastrier) from France and Algeria differs by being a larger species (female wing length 0.68-0.74 mm; male wing length 0.50-0.55 mm), the wing of the female has a poorly developed anal lobe bearing capitate setae, and the male flagel- lum has 8 flagellomeres. The 2 species of Nannohelea that were recently described by Szadziewski (1988) from Eocene Baltic am- ber differ from all extant species in having a 4 segmented palpus. ACKNOWLEDGMENTS Weare extremely grateful to Ethel L. Gro- gan for preparing the illustrations. 350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Clastrier, J. 1961. Notes sur les Ceratopogonides. XV. Ceratopogon et Alluaudomyia de la Region Pa- learctique. Archs. Inst. Pasteur Alger. 39: 401- 437. Downes, J. A. and W. W. Wirth. 1981. Chapter 28. Ceratopogonidae, pp. 393-421. Jn McAlpine, J. F. et al., eds. Manual of Nearctic Diptera. Vol. 1, 674 pp. Agric. Canada Mongr. 27. Ottawa. Grogan, W. L., Jr. and W. W. Wirth. 1980. Nan- nohelea, a new genus of biting midges of the tribe Ceratopogonini, related to Baeohlelea Wirth and Blanton (Diptera: Ceratopogonidae). J. Kans. Entomol. Soc. 53: 373-385. Szadziewski, R. 1988. Biting midges (Diptera, Cer- atopogonidae) from Baltic amber. Polskie Pismo Entomol. 57: 3--283. Tokunaga, M. 1964. Biting midges of the genus Cer- atopogon from New Guinea (Diptera: Ceratopo- gonidae). Pacific Insects 6: 292-299. Wirth, W. W. and W. L. Grogan, Jr. 1988. The pre- daceous midges of the world (Diptera: Ceratopo- gonidae; Tribe Ceratopogonini). Flora and Fauna Handbook no. 4. E. J. Brill, New York, xv + 160 Pp. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 351-357 HOST SPECIFICITY AND ESTABLISHMENT OF APHTHONA FLAVA GUILL. (CHRYSOMELIDAE), A BIOLOGICAL CONTROL AGENT FOR LEAFY SPURGE (EUPHORBIA ESULA L.) IN THE UNITED STATES ROBERT W. PEMBERTON AND NORMAN E. REES (RWP) Asian Parasite Laboratory, Agricultural Research Service (ARS, USDA) Seoul, Korea, % American Embassy, APO San Francisco 96301; (NER) Rangeland Insect Lab- oratory, ARS-USDA, Montana State University, Bozeman, Montana 59717. Abstract. —The potential host plant range of the European flea beetle Aphthona flava, a candidate biological control agent for leafy spurge, E. esula, was evaluated. Fewer than 19 (none of which are rare or legally protected) of the 113 Euphorbia species native to the U.S. appear to be potential host plants for 4. flava. Releases of A. flava were made in Montana (1985-1987), North Dakota (1985) and Idaho (1986). Establishment occurred in four of the eight Montana sites, and at the North Dakota site. Aphthona flava, which increased to 31 beetles/m* at one Montana site in 1988, is part of a complex of insects being introduced in an attempt to control leafy spurge, a serious weed of Great Plains rangelands. Key Words: Leafy spurge (Euphorbia esula L.) isa deep rooted perennial herb native to Europe and Asia, which has become one of the most serious weeds of rangelands in the Great Plains region of North America (Lacey et al. 1985, Watson 1985). Cattle, one of the primary products of the Great Plains, will not eat the plant nor much of the palatable forage growing near it (Messersmith and Lym 1983). Chemical control is usually not economical on the low value land that leafy spurge infests. For this reason, and since leafy spurge was not known to be a problem in its native Eurasian range, a biological control program was begun in the early 1960s by Agriculture Canada (Harris 1984). The USDA’s Agricultural Research Ser- vice joined the effort to discover, test and introduce insects of spurge in the mid 1970s (Pemberton 1985). These programs have re- sulted in the introduction of the following biological control of weeds, flea beetle, Great Plains, rangeland, weed complex of European Euphorbia feeding in- sects to the United States: a foliage and flow- er feeding moth (Hyles euphorbiae L., Sphingidae), a root feeding moth (Cha- maesphecia tenthrediniformis (Den. and Schif.), Aegeriidae), a stem boring beetle (Oberea erythrocephala (Schrank), Ceram- bycidae), a shoot-tip gall midge Spurgia capitigena Gagne, Cecidomyidae) and four root feeding flea beetles Aphthona flava Guill., A. cyparissiae (Koch), A. czwalinae Weise and 4. nigriscutis Foudras (Chrys- omelidae) (Pemberton 1985, Rees et al. 1986, Pemberton and Rees, unpublished data). Aphthona flava is one of a complex of 40 Aphthona spp. recorded to feed on Euphor- bia spp. in Europe (Harris et al. 1985) and Asia (Pemberton and Wang 1989). Like other Aphthona species that feed on Eu- phorbia species, the adults of A. flava feed 352 on leaves and flower bracts, and the larvae feed upon the root hairs and roots. 4. flava has one generation per year. The adults usu- ally emerge in June, feed and lay eggs for several months before dying. The mature larvae overwinter and pupate in late spring or early summer. This flea beetle is native to Europe, from northern Italy, east and north through Yugoslavia, Hungary, Czechoslovakia, Bulgaria, Rumania and Russia (Sommer and Maw 1982). It has been recorded from Euphorbia cyparissias L., E. esula L., E. seguieriana Necker and E. pan- nonica Host. (Kuntze 1930, Sommer and Maw 1982, Harris et al. 1985). Aphthona flava was evaluated as a can- didate biological control agent for leafy spurge by Sommer and Maw (1982), who found that the beetle’s potential host range would be limited to species of Euphorbia. Based on these data, 4. flava was introduced to Alberta and Saskatchewan, Canada (McClay and Harris 1984). Before A. flava could be introduced into the United States, additional host specificity testing was need- ed to better define the beetle’s potential host range within the genus Euphorbia. Host SpEcIFICITY TESTING The United States has 113 native species of Euphoria (sensu lato), including two rare species (E. garberi Engelm. ex Chapm. and E. deltoidea Engelm. ex. Chapm.) that are legally protected and nine other rare species that are under review for protected status (U.S. Dept. Agric. 1982, U.S. Dept. Inter. 1980 and 1983, Pemberton 1985). It is not possible to predict from the European host plant records which of these North Amer- ican Euphorbia species could become host plants of A. flava, because most American Euphorbia species belong to subgenera (Agaloma and Chamaesyce) that are not represented in the European flora. Aphthona flava, from E. esula near Pisa, Italy, was tested against ten North American Euphor- bia species in the USDA-ARS quarantine in Albany, California, during 1984 and 1985 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Pemberton, unpublished data). The Fu- phorbia species used in the testing included representatives of the subgenera that occur in North America! and species that are rare, weedy, ornamental, or sympatric with leafy spurge. Host plant suitability was studied by plac- ing adult beetles with test plant bouquets or potted plants to measure adult feeding and longevity; and by transferring eggs or first instar larvae to potted test plants to see which plants could support full larval develop- ment (to the third instar). Table 1 summarizes the results of these studies. None of the tested members of the subgenera Chamaesyce (3 of the 57 US. species) or Aga/oma (2 of the 26 U.S. species) were suitable hosts by any of the criteria measured. All six tested species of the sub- genus Esula (21 U.S. species) were accepted as adult food. Three of these (EF. incisa En- gelm., E. robusta (Engelm.) Small and E. palmeri Engelm.) supported adult longevity for more than two months. These three species and E. spatulata Lam. supported full larval development. Significantly, neither E. purpurea (Raf.) Fernald or E. telephiodes Chapm., both rare subgenus Esula species under review for protected status, appeared to be suitable hosts, since no larval devel- opment took place on these plants. From these data, we predict that some portion of the remaining 19 subgenus Esula species could be potential host plants for A. flava, if the beetles were to spread through the United States. If A. flava becomes es- tablished throughout the North American range of leafy spurge, eight subgenus Esula species, which are roughly sympatric with leafy spurge, might become host plants for the beetle. The most sympatric of these is E. robusta, a Rocky Mountain species (U.S. Dept. Agric. 1982), which was an acceptable laboratory host plant in the testing. Eu- ‘No species of the small subgenus Poinsettia (3 U.S. species) were used, since the two species tested by Som- mer and Maw (1982) did not support development. VOLUME 92, NUMBER 2 ee) n vs) Table 1. Summary of Aphthona flava host plant specificity testing on native North American Euphorbia species.* % of Plants Accepted % of Adults Living % of Plants Supporting Test Plant Species Subgenus for Adult Feeding 2 Months or Longer Larval 3rd Instar Euphoria esula Esula 100 (10/10) 86 (19/20) 90 (27/30) Euphorbia incisa Esula 100 (10/10) 63 (10/16) 90 (9/10) Euphorbia palmeri Esula 80 (8/10) 53 (8/15) 88 (7/8) Euphorbia robusta Esula 80 (8/10) 63 (10/16) 80 (8/10) Euphorbia spatulata Esula _ 5 (1/20) 20 (2/10)? Euphorbia purpurea” Esula 30 (3/10) 0 (0/16) 0 (0/10) Euphorbia telephiodes” Esula 60 (6/10) 0 (0/16) 0 (0/10) Euphorbia maculata Chamaesyce 0 (0/10) 0 (0/16) 0 (0/10) Euphorbia supina Chamaesyce 0 (0/10) 0 (0/16) 0 (0/10) Euphorbia serpyllifolia Chamaesyce 0 (0/10) 0 (0/16) 0 (0/10) Euphorbia corollata Agaloma 10 (10/10) 6 (0/16) 0 Euphorbia marginata‘ Agaloma 0 0 0 Euphorbia heterophylla: Poinsettia 0 0 0 * From Pemberton unpublished data 1984-85. > Rare species. © Tested by Sommer and Maw 1982. 4 Many small plants were in each pot, single plants are probably too small to support larval development. phorbia incisa and E. palmeri, which were also acceptable laboratory hosts, are south- western species (U.S. Dept. Agric. 1982) that may have some contact with leafy spurge in Nevada or northern Arizona. The other ac- ceptable laboratory host plant, E. spatulata, is a small annual that ranges throughout much of the United States (U.S. Dept Agric. 1982). Larval development of 4. flava oc- curred in 20% of the pots densely planted with E. spatulata; single plants may be too small to support complete larval develop- ment. The other sympatric species are: E. brachycera Engelm., a southwestern peren- nial, and three annuals: E. commutata En- gelm. from the east and south central U.S., E. lurida Engelm. from the northeast, and E. crenulata Engelm). of the Pacific states. Since relatively few of the 113 Euphorbia species native to the U.S. appeared to be potential hosts of A. flava, a petition (Pem- berton unpubl. report) for its release was made to the Federal Working Group of Bi- ological Control of Weeds. Approval for re- lease was received in 1985 and releases be- gan the same year. RELEASE AND ESTABLISHMENT All A. flava beetles intended for release were collected from leafy spurge popula- tions in the Pisa area of northern Italy by M. Stazi and M. Cristofaro (U.S. Dept. Agric.-ARS Biological Control Laboratory, Rome). Each collection was sent to the U.S. Dept. Agric—ARS Biological Control of Weeds Quarantine in Albany, California. In the quarantine, a small number (usually ca. 5%) of the beetles were killed and sent to Consulting Diagnostic Service (Berkeley. Calif.) to check for internal pathogens. None were found. A small number of specimens were also sent to R. White (U.S. Dept. Agric.-ARS U.S. National Museum, Wash- ington) to confirm their identity. The re- maining beetles were paired and placed on bouquets of leafy spurge to observe feeding and record oviposition. Beetles that fed and laid eggs normally were sent to the field for release. There wasa very high mortality (80-95%) experienced by overwintering larvae in lab- oratory cultures which significantly reduced 354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Releases and Establishment of Aphthona flava. Number Number of Adults Site Released Date Recovered Date Montana? North Bozeman (Gallatin Co.) 59 16 July 1985 no recovery (site sprayed 1986) Reed Point (Stillwater Co.) 50 31 July 1985 2 9 June 1987 2 10 June 1988 Columbus Island (Sweetgrass Co.) 57 31 July 1985 1 9 June 1987 Glacier National Park (Flathead 150 2 Aug 1985 no recovery to date Co.) Gallatin River (Gallatin Co.) 46 4 Aug 1985 no recovery to date Lyman Creek-Shade (Gallatin Co.) — 106 25 June 1986 =x: 31/m? 4 Aug 1988 Clyde Park (Park Co.) 240 10 July 1986 no recovery to date Lyman Creek-Sun (Gallatin Co.) 2077 in 6 re- 9 July-6 Aug 51/278 sweeps 22 July 1988 leases 1987 Idaho? Featherville (Elmore Co.) 200 8 July 1986 no recovery to date Rathdrum (Kootenai Co.) 210 24 July 1986 no recovery to date North Dakota‘ Bald Hill Dam (Barnes Co.) 260 in 2re- 11-23 July x 7.5/m? July—Aug 1987 leases 1986 x 14/m? July-Aug 1988 * Montana releases made primarily be N. Rees and R. Pemberton. > Idaho releases made by J. McCaffrey, University of Idaho at Moscow. © North Dakota release made by R. Carlson, North Dakota State University at Fargo. the number of beetles for field colonization. Consequently foreign field collected beetles, instead of laboratory reared material, were released. Since no parasites or pathogens had been found in Italian 4. flava popula- tions (although many are known from other areas (Sommer and Maw 1982)), release of field collected material from this area ap- peared to have few risks, and seemed jus- tified to try to establish the beetle in the U.S. Direct release of foreign collected ma- terial in the U.S., as a normal mode of op- eration, is unwise, since pathogens and par- asites, which could negate successful biological control programs, could easily be introduced. In 1988, after the releases of A. flava from Italy reported here were com- pleted, a pathogenic microsporidian, No- sema sp., and a lethal parasitic mite, 7rom- bidium susteri Feider, were found to be associated with Aphthona spp. collections, originating from Austria, that were intend- ed for release in the U.S. (G. Johnson, El Cerrito, Calif., pers. comm.). Aphthona flava was released in Montana from 1985 to 1987, and in North Dakota and Idaho in 1986 (Table 2). The seven Montana releases, east of the Continental Divide, were made by the authors (primar- ily NER) assisted by N. Poritz. The Glacier Park release was made by D. Lang (National Park Service, West Glacier). All of the Mon- tana sites had dense infestations of leafy spurge, estimated to constitute more than 50% of the above ground dry weight annual plant production. Brief descriptions of the Montana sites are as follows: The North Bozeman site (al- titude ca. 1600 m) is 3.2 km northeast of Bozeman and south of the Bridger Moun- VOLUME 92, NUMBER 2 tains in Gallatin County. It consists of an open, south facing slope cut by a shallow valley, which in addition to dense leafy spurge, had Rosa sp., and mixed annual grasses. This site had a history of herbicide spraying (Tordon and 2-4 D ester) for leafy spurge control and was, unfortunately, sprayed in 1986, the year following the re- lease. The Reed Point site (altitude ca. 1200 m) is 9.3 km east of Reed Point in Stillwater County. The spurge infestation is on the south bank of the Yellowstone River, be- tween the river and highway I-90. The site is a level terrace that has been used as a cattle pasture. The vegetation, except for Salix along the river, was weedy, including Iva xanthifolia Pursh. and Asclepias spe- ciosa Torr., in addition to the leafy spurge. Part of the site was plowed in 1986, the year after the release. The Columbus Island Site (altitude ca. 1100 m) is a rock and sand island, of ca. 2 hectares, in the Yellowstone River adjacent to the town of Columbus in Sweetgrass County. The dominant plants are Populus deltoides Marsh, Salix and Rosa spp. The leafy spurge density was 48 stems/m? in 1983. Goats have been used extensively at this site in an attempt to control leafy spurge. The Gallatin River site (altitude ca. 1600 m) is 9.6 km east of Bozeman in Gallatin County. The site lies on the east side of the Gallatin River and is dominated by Populus angustifolia James, leafy spurge and the ex- otic tansy (Tanacetum vulgare L.). Leafy spurge had a density of 129 stems/m? in 1987. The site has had periodic grazing. The Glacier Park site (altitude ca. 980 m) is located at Big Prairie on the east bank of the Flathead River in the western sector of the Park. In addition to large growths of spurge, this natural prairie has a mixture of small herbs and grasses with patches of in- troduced Linaria vulgaris L. The site had been plowed historically, but has been free of agriculture for many years. The Park Ser- vice has used mowing, burning, and a lim- 355 ited amount of plowing to try to control the leafy spurge at this site. The Lyman Creek Shade site (altitude ca. 1600 m) is located on the northern bank of Lyman Creek on the southern slopes of the Bridger Mountains ca. 8 km north of Boze- man in Gallatin County. Leafy spurge grows in a 50 m? opening, surrounded by Douglas fir (Pseudotsuga menziesii (Mirbel Franco), Rocky Mountain juniper (Juniperus scopu- lorum Sarg.) and Prunus virginiana L. Both Lyman Creek sites are on the City of Boze- man Water Company land and have had no grazing or chemical use for many years. The Clyde Park (altitude ca. 1600 m) is 5.6 km northeast of Clyde Park and south of the Crazy Mountains in Park County. The site consists of a level pasture, with Artemisia spp., Lupinus sp. and some grass, and a 30 m long 45° slope running from the pasture to a Populus trichocarpa T. & G. dominated riparian community. The bee- tles were released on the slope, which was densely covered by leafy spurge. The site, particularly the level portion, has been grazed and treated with herbicides to try to control leafy spurge. The Lyman Creek Sun site is an open, south facing hillside located ca. 200 m down stream from the Lyman Creek Shade site. The open infested hillside (ca. 1400 m7’) is surrounded by Douglas fir and aspen (Pop- ulus tremuloides Michx. On the hillside are Geranium viscosissimum Fish. & Mey. and species of Equisetum, Balsamorhiza, Arte- misia, Rosa and Symphoricarpos. In 1988, leafy spurge accounted for 77.5% (SD + 21.8) of the above ground dry weight plant biomass on the hillside. The x leafy spurge density was 216.5 g/m? (SD + 99.42). The North Dakota release was made by R. Carlson assisted by D. Mundal, North Dakota State University at Fargo. The re- lease site is at Bald Hill Dam in Barnes County. The site 1s adjacent to the reservoir and has zones of prairie and woodland con- sisting of planted shelter belt. The leafy spurge density was 133.4 stems/m? in 1986. 356 The Idaho releases were made by J. McCaffrey of the University of Idaho at Moscow. The Rathdrum site (altitude ca. 760 m) is located in Kootenai County in northern Idaho. Leafy spurge accounted for more than 50% of the annual production at this site, which in addition to spurge, had mixed annual grasses. The Featherville site (altitude ca. 1500 m) is in Elmore County in southern Idaho. The site is a disturbed sagebrush-grass com- munity with an estimated spurge density of 50 stems/m? in about a one hectare infes- tation. The Montana releases at North Bozeman, Reed Point, Columbus and Lyman Creek Shade were made in 3 m x 3 m plastic screen cages to concentrate the beetles and possibly aid their establishment. The re- maining Montana releases were made in the open. The Idaho releases were made within one m? cages and in the open near the cages at both sites. The North Dakota release was made in four 3.3 m? cages. The number of A. flava released ranged from 50 to 260 bee- tles per site, except at the 1987 Lyman Creek Sun site release, where a major collection effort resulted in 6 releases totaling 2077 beetles. This ““mass” collection and release was done to learn if releasing large numbers could promote better establishment and rapid numerical increase in 4. flava popu- lations after establishment. Aphthona flava established at four of the eight Montana sites and at the single North Dakota site. No recoveries have been made to date at the Idaho sites. The best Montana establishment was at the Lyman Creek Shade site, where a mark and recapture study (following unpublished techniques used by A. McClay, Vegreville, Alberta) estimated an A. flava population of 31/m? adults. A visual search and count of A. flava adults at the North Dakota site yielded 14/m? in 1988 (R. Carlson, pers. comm.) Beetles at the Reed Point and the Columbus, Montana sites were not recovered the year following release (1986), but were found in low numbers two PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON years after release (1987). Establishment of A. flava was obtained with the release of only 50, 57 and 106 beetles per site. In Mon- tana, releases made in cages resulted in es- tablishment at three of four sites. The cage release at the North Bozeman site may have failed because of the spraying of herbicides at the site the year following release. The mass release at the Lyman Creek Sun site was the only one of four open releases in Montana that produced an establishment. Good establishment was obtained in North Dakota where cages were used but no re- covery of A. flava has been made in Idaho where both open and cage releases were made. There is no apparent pattern relating es- tablishment of A. flava to known site char- acteristics or release dates. IDENTIFICATION AND DISTINGUISHING FEATURES Aphthona flava is completely orange, has no elytral sculpturing and is large (3—4 mm) for a flea beetle. The only North America flea beetles recorded from Euphorbia species are Glyptina species. Glyptina spuria Le- Conte has been collected from E. maculata L., and E. blodgettii Engelm. ex A. Hitche. from the east and southeastern U.S. (Wheel- er 1981). Glyptina cyanipennis Crotch is re- corded from E. cyathophora Murray in Florida (Schwarz 1890) and G. atriventris Horn adults have been found on flowers of leafy spurge in North Dakota (Julian 1984). All of these G/yptina species are small (<3 mm), have regular rows of punctures on the elytra and are darkly colored (Arnett 1968). The other Aphthona species that have been introduced against leafy spurge are smaller (2-3 mm) and black (4. czwalinae Weise) or brown (A. cyparissiae (Koch) and A. ni- griscutis Foudras. CONCLUSION The ability of A. flava to control leafy spurge has not yet been established. The beetle has not had time to increase its pop- VOLUME 92, NUMBER 2 ulations to the point where they may begin to stress the plants. We think that the utility of A. flava will be as part of a complex of natural enemies, stressing different parts of the plant and in different areas of the weed’s range, that may eventually control leafy spurge. ACKNOWLEDGMENTS We thank Gerald R. Johnson USDA- APHIS, El Cerrito, California, and Noah Poritz, USDA-APHIS, Bozeman, Montana (both former USDA-ARS employees) for technical assistance; Massimo Stazi and Massimo Cristofaro, Biological Control of Weed Laboratory, USDA-ARS, Rome, It- aly for collecting A. flava; Richard E. White, Systematic Entomology Laboratory, USDA- ARS, Washington, D.C. for determination of A. flava; B. Thomas, Berkeley, California for examining 4. flava for pathogens; Rob- ert Carlson, North Dakota State University at Fargo, David Lange, Glacier National Park, Montana, and Joseph McCaffrey, University of Idaho at Moscow for releasing A. flava in their areas and for generously sharing their data relating to these releases; Alec McClay, Alberta Environmental Cen- ter, Alberta, Canada and Joseph McCaffrey for reviewing the manuscript. LITERATURE CITED Arnett, R. H. 1968. The beetles of the United States. Amer. Entomol. Inst., Ann Arbor. 1112 pp. Harris, P. 1984. Euphorbia esula-virgata complex, leafy spurge and E. cyparissias L., cypress spurge (Euphorbiaceae), pp. 159-69. Jn Kelleher, J. S., and M. A. Hulme, eds., Biological control pro- grams against insects and weeds in Canada 1969- 1980. pp. 159-169. Harris, P., P. H. Dunn, D. Schroeder, and R. Vonmoos. 1985. Biological control of leafy spurge in North America, pp. 79-92. Jn Watson, A. K., ed., Leafy spurge. Weed Sci. Soc. Amer. Mongr. No. 3, pp. 104. Julian, J.J. 1984. A survey of insects associated with leafy spurge in North Dakota. Master’s thesis, 357, Entomol., North Dakota State Univ. Fargo. pp. 114. Kuntze, R. 1930. Drugi przyczynek do znajomosci fauny Halticinow (Coleoptera) Polski. Polsk. Pis- mo. Entomol. 9: 40-64. Lacey, C. A., P. K. Fay, R. G. Lym, C. C. Messersmith, B. Maxwell, and H. P. Alley. 1985. Leafy spurge, biology and control. Montana State Univ. Coop. Exten. Serv. (Bozeman) Circular 309. 15 pp. McClay, A. S. and P. Harris. 1984. Biological control of leafy spurge in Canada. Proc. Leafy Spurge Ann. Meeting, Dickinson, North Dakota. pp. 18-20. Messersmith, C. C. and R. G. Lym. 1983. Distri- bution and economic impacts of leafy spurge in North Dakota. North Dakota Farm Research 40: 8-13. Pemberton, R. W. 1985. Native plant considerations in the biological control of leafy spurge, pp. 365- 390. In Delfosse, E. S., ed., Proc. VI Int. Symp. Biol. Contr. Weeds, Vancouver, Canada. pp. 885. Pemberton, R. W. and R. Wang. 1989. Survey for natural enemies of Euphorbia esula L. in northern China and Inner Mongolia. Chinese J. Biol. Contr. 5: 64-67. Rees, N. E., R. W. Pemberton, A. Rizza, and P. Pecora. 1986. First recovery of Oberea erythrocephala on the leafy spurge complex in the United States. Weed Sci. 34: 395-397. Schwarz, E. A. 1890. Food-plants and food-habits of some North American Coleoptera. Proc. Entomol. Soc. Wash. 1: 231-233. Sommer G. and E. Maw. 1982. Aphthona cyparissiae (Koch) and 4. flava Guill. (Coleoptera: Chryso- melidae): Two candidates for the biological con- trol of cypress and leafy spurge in North America. Unpubl. report Commonwealth Inst. of Biol. Contr., Delemont, Switzerland. 60 pp. US. Dept. Agric. 1982. National list of scientific plant names. Soil Conserv. Serv. Tech. Pub. 159 Vol. I, pp. 416. U.S. Dept. Inter. Fish and Wildlife Service. 1980. En- dangered and threatened wildlife and plants: Re- view of plant taxa for listing as endangered or threatened species. Fed. Regis. 45: 82480-82569. . 1983. Endangered and threatened wildlife and plants. Supplement to review of plant taxa for list- ing as endangered or threatened species. Fed. Reg- is. 48: 53640-53670. Watson, A. K. 1985. Introduction—the leafy spurge problem. pp. 1-6. /n Watson, A. K., ed., Leafy spurge. Weed Sci. Soc. Amer. Mongr. No. 3, pp. 104. Wheeler, A. G. 1981. Insect associates of spurges, mainly Euphorbia maculata L., in eastern United States. Proc. Entomol. Soc. Wash. 83: 631-641. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, p. 358 NOTE Two new synonyms in Rhyacophilidae (Trichoptera) Specific identifications of Trichoptera al- most invariably depend on examination of the genitalia, and principally those of the male which exhibit characteristics that are far more conspicuous than those of the fe- male. Regarding specific characters of cad- disflies McLachlan (1874, A monographic revision and synopsis of the Trichoptera of the European fauna. Pt. I:1-46. London) remarked, “Colour, minor points of neu- ration, &c., furnish these characters in part; but the most important are found in the anal appendages, especially of the male.” Over the years this method has become a well established convention in Trichopterology, and today the description of the male gen- italia is essential in virtually all caddisfly species descriptions. However, in the past this method was not so universally accepted and many species that were described solely on the basis of females can only be regarded presently as nomina dubia. Fortunately, subsequent taxonomic contributions have made it possible to identify the females of most of the eastern North American species of Rhyacophila. Recent examination of the female holotypes of two species has re- vealed that Rhyacophila formosa Banks is conspecific with vuphipes Milne, and main- ensis Banks with melita Ross. Formosa is a member of the fuscu/a group that includes one other species, fuscula (Walker). Main- ensis is a member of the siberica group that includes only four other eastern species, amicis Ross, atrata Banks, manistee Ross, and minor Banks. Female descriptions of all of these species have been provided by Schmid (1981, Mém. Soc. Ent. Canada 116: 1-83), with the exception of amicis. How- ever, I have examined the female of amicis and find that, as in the females of all the aforementioned species, it is quite distinct. I am grateful to Scott R. Shaw, then at the Museum of Comparative Zoology [MCZC], Harvard University for the loan of type material, and to Donald S. Chandler, University of New Hampshire, for review- ing the manuscript. Rhyacophila formosa Banks Rhyacophila formosa Banks 1911, Trans. AmerEnt: Soc: 37: 3533/35); 2: Rhyacophila vuphipes Milne 1936, Studies N. Amer. Trich. Cambridge, Pt. 3, pp. 99, 102, 111, fig. 6. New SyNonyM. Examination of the 2 holotype of formosa [MCZC] has revealed that it matches the description of vuphipes provided by Schmid (1981). Thus, the latter is recognized here as a junior synonym of formosa. This species is widespread along the east coast of North America, but it is not especially common. Sherberger and Wallace (1971, New York Ent. Soc., 69: 43-44) mention that larvae occur in small, rocky rivers. Reliable rec- ords are known from Georgia, Massachu- setts, New York, North Carolina, Ontario, Pennsylvania, Quebec, South Carolina, Tennessee, and West Virginia. Rhyacophila mainensis Banks Rhyacophila mainensis Banks 1911, Trans. Amer. Ent. Soc. 37: 354, 8. Rhyacophila melita Ross 1938, Ill. Nat. Hist. Survey Bull. 21: 104-105, f. 6, 6. NEw SYNONYM. Examination of the 2 holotype of mainen- sis [MCZC] has revealed that it matches the description of melita provided by Schmid (1981). Therefore, the latter is recognized here as a junior synonym of mainensis. Re- liable records are known from Maine, Mas- sachusetts, Michigan, New Hampshire, Newfoundland, New Jersey, New York, Quebec, and West Virginia. John S. Weaver III, Department of Ento- mology, University of New Hampshire, Dur- ham, New Hampshire 03824. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 359-360 Book REVIEW The Plant- Feeding Gall Midges Of North America. Raymond J. Gagné. 1989. Cor- nell University Press, Ithaca. 356 pp. $45.00. Gall midges, or cecidomyiids, are intrigu- ing insects to study because they are so small and spend most of their lives wrapped in a specialized chamber provided for them by plants. Adults, the size of a comma, do not feed. They emerge to mate and lay eggs, often dying the same day. There are about 900 species of gall midges in North Amer- ica. Most of them, or their galls, including the one-third which are known but not yet identified to species, are illustrated and keyed in Gagné’s book. A wide range of topics is covered in the eight chapters of this 356-page book. The chapter on biology reviews present knowl- edge about how gall midges select and cope with different hosts. It discusses their strat- egies for survival and the damage they cause to economic plants. The chapter on anato- my gives an overview of the life stages: egg, larva, pupa, and adult. A detailed discus- sion of third instar larval anatomy is post- poned to the chapter containing the larval key. Technical terminology 1s kept to a min- imum, and specialized words are clearly de- fined in the glossary. The chapter on clas- sification reviews the history of gall midge taxonomy in North America, beginning with the Hessian fly in 1817. Problems with the naming of species are mentioned, and a summary of the gall midge genera is pre- sented. A short chapter defines galls and their growth and diversity. It describes pri- mary characteristics for all galls, including those made by wasps, mites, and other flies. The book’s two keys are an indication of the considerable emphasis placed on iden- tification. Contrary to the general view, Gagné tells us morphology of the larvae and knowledge of the host are important for identification to the species level. Gagné does not give us a key to adults because adults are difficult to prepare for study and their characters are difficult to use. Both male and female are required for identification to the species. Females, by themselves, usually cannot be identified even to genus. There- fore, one chapter is a key based on the ma- ture or third instar larvae. Another chapter, comprising over half of the book, is a key to the galls, based on the host plant and gall types, their location on the plant, and dam- age they cause. Couplets in both keys are well constructed and clearly worded. Knowledge of the host plant group is a nec- essary prerequisite for proper use of both keys but otherwise they are not difficult. The gall key may be easily used in the field for quick identifications. Another chapter dis- cusses the collecting, rearing, and general study of gall midges. All information in the book is abundantly documented in a bib- liography of over 450 references. Many of the articles cited are recent and a few are still in press, indicating thorough research and a concern for accuracy. This hardbound book is printed on acid- free paper in an easy-to-read style. The two column format affords wide margins. There are 434 impressive half-tone drawings, all uncluttered, clear, and well labeled. Many of the drawings help to ease the user through the larval and plant keys while others illus- trate important pupa and adult types. There are also four high quality, full page, colored plates showing life stages of several different gall midges and their hosts. A few flaws in organization weaken the book somewhat. The chapters on distribu- tion of gall midges and the morphology of the galls should have been placed before the keys rather than between them because they serve to introduce us to the subject. The chapter on collecting, rearing, and preparing gall midges for study belongs at the end. The 360 valuable information on evolution of gall midges is scattered throughout the book and could have been brought together to make an interesting chapter. This is the first new book on midges in 50 years. This book, intended for the general reader, will also appeal to the professional entomologist and ecologist because it deals well with this unique group of flies and their hosts. Gagné’s purpose in writing is to list PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON all of the plant gall midges of North America and to review the current literature in order to encourage and assist the further study of these insects. He has succeeded very well in achieving these goals. This is a succinct, well conceived, and well written book. Paul E. Boldt, Grassland, Soil, and Water Research Laboratory, USDA, Agricultural Research Service, Temple, Texas 76502. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, p. 360 Note A New Synonym in Hydroptilidae (Trichoptera) In a paper about Trichoptera from Penn- sylvania (Ann. Carnegie Mus. 47: 1-12, 1978) we described Stactobiella solzhenit- syni, unfortunately unaware of the fact that S. martynovi Blickle & Denning (J. Kansas Ent. Soc. 50: 287-300, 1977) was in press at the same time. We recognize here that S. solzhenitsyni Sykora & Weaver is a junior synonym of S. martynovi Blickle & Den- ning. John S. Weaver III, Department of Ento- mology, University of New Hampshire, Dur- ham, New Hampshire 03824 and Jan L. Sy- kora, Department of Environmental Health—IEHS, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, Pennsylvania 15261. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 361-362 Book REVIEW Plant Stress-Insect Interactions. Edited by E. A. Heinrichs. 1988. John Wiley & Sons, New York, 492 pp. Hardcover $59.95. E. A. Heinrichs, Entomology professor and department head at Louisiana State University, has compiled a book of thirteen chapters on plant stress and insect interac- tions. As the author of chapter 1, Heinrichs first reviews and categorizes plant stress, and then defines it as “‘any abiotic or biotic fac- tor of the environment that affects plant physiology, chemistry, growth, and/or de- velopment in such a way that plants per- form below the average fora region” (p. 10). The remaining twelve chapters review cur- rent knowledge of how plant stress (natural or man-induced) affects insects, especially pests of economically important plants, and to a lesser extent, how insects alter the im- pact of plant stress. Chapters 2 through 5 describe abiotic fac- tors of plant stress, specifically mineral nu- trition, water stress, temperature-induced stress, and electromagnetic radiation. Chap- ters 6 through 9 review the (mostly) man- induced plant stress of insecticides, plant growth regulators, air pollution, and me- chanical damage. The final four chapters ad- dress the role of insects in altering the im- pact of environmental stress, plant stress and natural enemy efficacy, and pathogens and weeds as stress-inducers in plant/insect interactions. As with many edited books, the quality of the thirteen chapters varies considerably. For example, Dale’s chapter on soil mineral stress and Campbell’s chapter on plant growth regulators (PGR) are excellent re- views that not only discuss the complexity of the plant stress/insect interactions, but also provide a list of research needs. In con- trast, some of the chapters, such as Beren- baum’s on electromagnetic radiation, Al- tieri’s on weed-induced stress, and Smith’s on mechanical damage are incomplete. Al- though Berenbaum reviews ultraviolet ra- diation and plant chemistry as it affects host plant suitability, she ignores such topics as shade as a stress factor, nutritional as op- posed to allelopathic effects of radiation on insects, and the direct influence of radiation on insects. Similarly, Smith’s chapter on mechanical damage focuses on allelochem- ics as a factor in host plant suitability, but does not discuss how nutritional chemistry changes with damage. Most of the research presented measured the response of insects to varying plant con- ditions, with little focus on the cause for any change in response. As Campbell states, “most reports of effects of PGRs on insect- plant interactions do not explore the chem- istry of these effects and are only observa- tions of insect performance on the treated plants” (p. 234). Entomologists need to fur- ther develop their knowledge of the mech- anisms underlying these changes. In fact, Hughes’s chapter on air pollution would be a good source for a graduate student re- search problem as he devotes about 40% of his text to methods, approaches, and re- search needs. With the focus on individual topics in this book, discussion of the interrelationship of the mechanisms of plant stress falls between the chapters (although see p. 299). In ad- dition, no general theory is developed to summarize our knowledge of plant stress/ insect interactions or to provide testable predictions for future research. However, the absence of a theory may indicate that the subject is still in a descriptive stage of development. Nevertheless, to my knowl- edge no books preceding this one focus ex- clusively on plant stress/insect interactions. Instead, entomologists have had to rely on reviews found in various scientific journals. Thus, Heinrichs’ book is an excellent at- 362 tempt to fit together some of the puzzle pieces of plant stress/insect interaction re- search. But other pieces are still missing, and though this will be a useful reference for insect ecology and pest management courses, it may be especially useful as a basis Book Insect Spiracular Systems. By T. B. Ni- kam and V. V. Khole. Ellis Horwood Limited, Chichester, England and Hal- stead Press, New York. 1989, 136 pp. The intent of this short book is to provide students and researchers with an introduc- tion into the realm of spiracles and their interaction with other elements of tracheal systems. This objective was achieved, how- ever, in a somewhat abbreviated manner and one can only wish for an expanded ver- sion. The strong points of the text are the phys- iological discussions found in Chapter 3 (Environmental compulsions and relative adaptive features of spiracles), Chapter 4 (Exogenous and endogenous factors affect- ing the functioning of spiracles), and Chap- ter 7 (Insects: an air-cooled engine). Al- though these chapters are only 10, 14, and 11 pages, respectively, they are quite good, draw together many recent publications, and are recommended readings. The weak parts of the text generally are found in the critical first part of the book, the comparative morphology and devel- opmental sections. Chapter | on develop- ment and metamorphosis leaves the most unsaid. Although information available on embryological development of spiracles 1s not extensive, more should have been writ- ten because this chapter serves as the basis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for future research in plant/insect interac- tions. William O. Lamp, Department of Ento- mology, University of Maryland, College Park, Maryland 20742. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 362-363 REVIEW for the entire text. In discussing metamor- phosis, the authors correctly indicate that this phenomenon imparts great alterations on the system, yet they concentrate, as ex- haustively as on any other subject, on changes occurring between cyclorrhaphan larval instars (including four pages of plates where one plate would have been sufficient) and only minimal reference is made to changes appearing 1n the pupa and none to adult metamorphosis. Other major criti- cisms of this chapter include taking a sen- tence from Snodgrass’s classic text (p. 426) that is quoted nearly verbatim on page 14 without citation: reprinting the embryo fig- ure of Dixippus from Snodgrass with an ob- vious tracheal invagination on the first max- illary segment, yet stating that none exist in this region; leading the reader to infer that ectodermal invaginations give rise to the tracheal system at about 78 hours for all insects (page 13); referring to the integu- ment as a skin (page 13); and stating that Holometabola undergo incomplete meta- morphosis (page 17). A short discussion on the spiracular cuticle would have added 1m- portant depth and aided in understanding closing mechanisms. Chapter 2 discusses the morphology of spiracles by order and does a commendable job of presenting the comparative structure and physiology of the thoracic and first ab- dominal spiracles of adults. However, this VOLUME 92, NUMBER 2 would have been an opportune place to also discuss the evolution of spiracles, to spec- ulate on which of the four pairs of spiracles in Diplura are homologous (if any) to other insect orders, and to discuss the number of abdominal spiracles of each order (the num- ber is cited only for adults of some orders). Unfortunately there 1s no separate chap- ter, or major subdivision of a chapter, de- voted to larval spiracles, including structure and ordinal differences and similarities. There is a wealth of data on these larval structures making this omission (except for the previously mentioned cyclorrhaphan Book Crop and Plant Protection: The Practical Foundation. By Rudolph Heitefuss. Ellis Horwood Limited, Chichester, England, Distributed by Halsted Press, New York. 1989, 261 pp. Cloth. Crop protection based on the integration of all known pest management techniques is a goal of many crop production systems. To achieve this goal a thorough understand- ing of the principles of all aspects of crop management, along with knowledge of en- vironmental and economic consequences due to a particular management technique, is needed. The book reviewed here attempts to provide a concise outline of current crop management practices that can be used by students and practitioners alike. The current edition is an English trans- lation of the original German Pflanzen- schutz, Second Edition, published in 1987 by Georg Thieme Verlag. In general, the translation is well done with few grammat- ical errors. Several typographic errors were found. At times page references within the text were not complete. The use of “‘(see p. 363 spiracles and a few scattered references) un- fortunate. The writing style of the authors is direct and prolific in its citations; one is quickly reminded of the Annual Review volumes rather than writings in most texts. Perhaps the brevity of writing is partly the result of page limitations imposed upon the authors by publishers; if so, the work suffered from this constraint. Richard J. Elzinga, Department of Ento- mology, Kansas State University, Manhat- tan, Kansas 66506. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 363-364 REVIEW 000)” was noted several times. Obviously the actual page numbers of the references were not provided which caused some con- fusion. The book was written with 11 chap- ters and was illustrated adequately. The author based much of his informa- tion on crop protection on studies con- ducted in western Europe, particularly the Federal Republic of Germany. Most refer- ences cited are German with some citations provided from the United Kingdom and the United States. The book is arranged in a logical manner and provides an easy to fol- low sequence of how crop protection pro- grams are developed, and the means and tools with which to implement such pro- grams. Discussions of the ecological aspects of pest management are provided as are dis- cussions on most control measures. An in- terdisciplinary approach is used with fun- damental ideas presented from the fields of plant pathology, nematology, entomology, weed science and crop fertility. Though the author could not provide comprehensive discussions of all included subject matter, he has done a commendable job of tying 364 together the needed basic information in or- der to make the book worthwhile to all per- sons interested in crop protection. Several of the chapters included in the text provided much needed insight into the growing environmental and economic con- cerns faced by most agricultural producers. I was particularly intrigued with two chap- ters: Chapter 2, Importance of Farm Eco- nomic Management of Plant Protection; and Chapter 7, Consequences of the Use of Chemical Plant Protection Agents. These chapters provided fresh ideas on improving crop management in order to stabilize prof- its and improve environmental conditions associated with agriculture. The chapters provided thought provoking ideas seldom mentioned in many texts. Other chapters dealing with reduction of pest damage and the description of control tactics were well PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON written. The chapter on direct control mea- sures, particularly the description of chem- ical plant protection agents, was a bit long but does give the reader a basic background in how various pesticide family groups were derived and how they work. In general I find this book to be a useful contribution to the science of crop protec- tion. The integration of management tech- niques from all disciplines into one text 1s unique and provides students with an ex- cellent reference manual. It will become a valuable edition to libraries throughout the world in spite of its emphasis on German production practices. Laurence D. Chandler, /nsect Bioliogy and Population Management Research Labo- ratory, Agricultural Research Service, USDA, P.O. Box 748, Tifton, Georgia 31793. PROC. ENTOMOL. SOC. WASH. 92(2), 1990, pp. 365-372 SociETY MEETINGS 949th Regular Meeting— May 4, 1989 The 949th Regular Meeting of the Ento- mological Society of Washington was called to order by President F. Christian Thomp- son in the Naturalist Center, National Mu- seum of Natural History, at 8:03 p.m. on 4 May 1989. Twenty-three members and four guests were present. Minutes of the April meeting were read and approved with one addition and one correction. Membership Chairman G. B. White read the name of the following applicant for membership: Don Harrington, Richardson, Texas. Gaye L. Williams exhibited a gourd dec- orated with arthropod motifs carved in a Mexican vein. The gourd had been sent by a former ESW member, Elaine Lowry, who now resides in the Southwest. Edd Barrows described the Michener Re- tirement Conference, or “MichFest,” held at the University of Kansas on 14-15 April to honor one of this century’s greatest ento- mologists. The warmth of this occasion was amply conveyed in a series of slides that also provided glimpses of the local flora and at least one prominent Kansan declivity. The speaker for the evening was James B. Stribling, Associate Research Professor, Department of Biology, Georgetown Uni- versity, Washington, D.C. His talk, entitled “Life History of Marsh Beetles in Treeholes and Bromeliads,” focused on the ecology of the Scirtidae, whose larvae are aquatic and occur in phytotelmata: plant structures that are capable of holding water, e.g. leaf bases of bromeliads, coconut shells, and depres- sions or hollows in trees. As a group, scirtid larvae are instantly recognizable by their long, multisegmented antennae. However, specific determinations are often impossible without rearing. Adults are terrestrial and may be collected at light traps or by beating vegetation. Dr. Stribling’s extensive field experience in the United States and Central America has yielded data on larval char- acters and habitat specificity that together will constitute a foundation for phyloge- netic studies of this family. J. H. Fales reported that during the last two years the monarch butterfly (Danaus plexippus (Linnaeus)) has been virtually ab- sent as a Spring migrant in southern Mary- land. Similar observations have been made by lepidopterists in the Northeast. Fall mi- grations also have been poor. However, monarchs were seen moving through south- ern Maryland on 26 and 28 April, dates that are slightly ahead of Fales’ average for first sightings, which is | May. Mignon Davis thanked her two children, Marisa and Steven, for helping to make the date-nut bread and other goodies brought to this evening’s meeting. President Thompson urged the member- ship to turn out in force for what promises to be an especially enjoyable annual ban- quet. Visitors were introduced and the meeting was adjourned at 9:18 p.m. Refreshments followed. Richard G. Robbins, Recording Secretary 951st Regular Meeting— October 5, 1989 The 951st Regular Meeting of the Ento- mological Society of Washington was called to order by President F. Christian Thomp- son in the Naturalist Center, National Mu- seum of Natural History, at 8:00 p.m. on 5 October 1989. Twenty members and five guests were present. Minutes of the May meeting were read and approved. 366 President Thompson praised the accom- plishments of outgoing Editor Hiram G. Larew and introduced his successor, Robert D. Gordon, Systematic Entomology Labo- ratory, U.S. Department of Agriculture. Dr. Gordon thanked the Society for the trust it had placed in him and announced that dur- ing his tenure he would refrain from making major editorial decisions by himself. In- stead, he will refer such matters to the Pub- lications Committee, in which there have been some recent changes: Don Davis and Don Whitehead are slated to replace Bob Peterson and Rebecca Surdick. President Thompson added that there are vacancies in the roster of officers for 1990. Prospective candidates are welcome. Recording Secretary R. G. Robbins read the names of the following applicants for membership: Richard L. Bottorff, Placer- ville, California; R. G. Brown, Common- wealth Agricultural Bureaux, Institute of Biocontrol, Curepe, Trinidad; Zapparoli Marzio, Instituto di Difesa Delle Piante, Vi- terbo, Italy; Vicente Carapia Ruiz, Depar- tamento de Parasitologia, Universidad Autonoma Agraria “Antonio Narro,” Bue- navista Saitillo, Coahuila, Mexico; Chang Eon Lee, Department of Biology, College of Natural Sciences, Kyungpook National University, Taegu, South Korea; John S. Weaver III, Department of Entomology, University of New Hampshire, Durham; Charles E. Williams, Department of Biol- ogy, Washington and Lee University, Lex- ington, Virginia; and Michael J. Sharkey, National Institute of Agro-Environmental Sciences, Division of Entomology, Yatabe, Tsukuba, Ibaraki, Japan. Mignon Davis circulated a sign-up sheet for volunteers to bring refreshments to our meetings. In an evening replete with exhibits, Doug Sutherland led off by distributing copies of U.S. House of Representatives Joint Res- olution 411 designating the monarch but- terfly (Danaus plexippus (Linnaeus)) as our national insect. Introduced by Leon E. Pa- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON netta of California on 27 September of this year, the resolution reads: ‘““Whereas the monarch butterfly, native to North Amer- ica, is found throughout the United States; whereas the monarch is a unique represen- tative of over 600 species of butterflies and nearly 90,000 other insects that are an in- tegral part of the natural heritage of the United States; whereas the great diversity of insects play a vital role in the daily lives and ecology of the environment; whereas the population of monarchs is declining un- der pressure from urbanization and loss of habitat which results in the reduction of the host plant (milkweed) and overwintering groves of trees in California and Mexico; whereas conservation efforts are under way in both California and Mexico to maintain these overwintering sites; whereas Pacific Grove, California, holds an annual festival celebrating the return of the monarchs to overwinter until spring when the monarchs begin a northward flight; and whereas the monarchs enhance the beauty of the envi- ronment and signals [sic] the need for pro- tection and conservation of the natural wonders: Now, therefore, be it resolved by the Senate and House of Representatives of the United States of America in Congress assembled, that the monarch butterfly is designated and adopted as the national in- sect of the United States, and the President is authorized and requested to declare such fact by proclamation.” Dr. Sutherland also exhibited this October’s issue of Ranger Rick (a young people’s nature magazine pub- lished by the National Wildlife Federation), which instructs children on how to lobby for passage of H. J. Res. 411; the summer 1989 issue of Wings (vol. 14, no. 2, a pub- lication of the Xerces Society), which is en- tirely devoted to the monarch; and this Sep- tember’s issue of the Reader’s Digest, which on pages 134-136 contains an article about monarchs entitled ““My Butterfly Mystery,” by Jonathan Weiner. Continuing in a lepidopterological vein, Edd Barrows displayed a larva of the tiger VOLUME 92, NUMBER 2 swallowtail (Papilio glaucus Linnaeus) (“very dark this year’), a tomato hornworm (Manduca quinquemaculata (Haworth)) be- decked with the cocoons of a braconid par- asite, a brachypterous female moth of un- known affinity, and a single live female bag- worm (Thyridopteryx ephemeraeformis (Haworth)) together with a quarter pound of preserved suitors. For lepidobibliophiles, Ted Spilman ex- hibited the new paperback Florida Butter- flies by Eugene J. Gerberg and Ross H. Ar- nett, Jr., Natural Science Publications, Inc., Baltimore, Maryland, $9.95, ISBN 0-89140- 031-1. Each of the true butterflies (Papilio- noidea) of Florida is illustrated with a color photograph, followed by brief descriptions of one or both sexes, adult habitats, flight periods, larval food plants, and distribution in Florida. For the confusingly similar skip- pers (Hesperioidea), only distributions and larval hosts are listed. Ninety-seven true butterflies and 67 skippers are known to breed in Florida, which means that for both superfamilies the Sunshine State can claim over 20% of the North American fauna. President Thompson issued a change in orders by exhibiting volume four of Myia (1989, Insect Associates, South San Fran- cisco, California), a tribute to Edward Lu- ther Kessel on the occasion of his 85th birthday. Kessel is the foremost authority on flat-footed flies (Platypezidae). Between 1947 and 1987, he and his coauthors pub- lished 56 papers in which over one third of the world’s 215 known platypezid species were named and described. Dr. Thompson also displayed a souvenir of his attendance at last year’s Nobel Symposium: The Hi- erarchy of Life. Molecules and Morphology in Phylogenetic Analysis (1989, Excerpta Medica, New York, ISBN 0-444-81073-0) contains the proceedings of Nobel sympo- sium 70 held at Alfred B. Nobel’s Bjérk- born, Karlskoga, Sweden, 29 August—2 Sep- tember 1988. A descriptive brochure and photos of the symposium were circulated among the membership. 367 The speaker for the evening was Donald R. Whitehead, Research Entomologist, Sys- tematic Entomology Laboratory, U.S. De- partment of Agriculture, whose talk was en- titled ‘“‘Mimetic Miullipede Madness: Tropical Biology in Eastern North Ameri- ca.’ Dr. Whitehead’s millipede studies have focused on the several species complexes that occur in West Virginia, where problems in geographic variation are often com- pounded by Miillerian mimicry and by dis- tributions that appear to be unrelated to lo- cal drainage systems, soil types, or other environmental variables. One consequence of this chaos 1s that the millipede fauna of eastern North America is not yet amenable to phylogenetic analysis. However, thanks to Bob Gordon’s camera caddying, we now have a good photographic record of these peripatetic arthropods. Visitors were introduced and the meeting was adjourned at 9:20 p.m. Refreshments followed. Richard G. Robbins, Recording Secretary 952nd Regular Meeting—November 2, 1989 The 952nd Regular Meeting of the En- tomological Society of Washington was called to order by President F. Christian Thompson in the Naturalist Center, Na- tional Museum of Natural History, at 8:05 p.m. on 2 November 1989. Twenty mem- bers and 19 guests were present. Minutes of the October meeting were read and ap- proved — pending determination of the cor- rect generic name for the tomato hornworm (vide supra). Membership Chairman G. B. White read the names of the following applicants for membership: John R. Linley, Medical Ento- mology Laboratory, Institute of Food and Agricultural Sciences, University of Flori- da, Vero Beach; Stuart H. McKamey, De- partment of Ecology and Evolutionary Bi- 368 ology, University of Connecticut, Storrs; and Robert W. Pemberton, Asian Parasite Lab- oratory, APO San Francisco, California. Nominating Committee Chairman Ted Spilman presented the following slate of of- ficers for 1990: President, Jeffrey R. Aldrich (automatic); President-Elect, David R. Smith; Recording Secretary, Richard G. Robbins (incumbent); Corresponding Sec- retary, Hollis B. Williams; Treasurer, Nor- man E. Woodley (incumbent); Program Chairman, Gary Steck; Membership Chair- man, Geoffrey B. White (incumbent); Edi- tor, Robert D. Gordon; Associate Editor, Thomas J. Henry; and Custodian, James B. Stribling. Ted thanked Nominating Com- mittee members Edd Barrows and Doug Sutherland for their assistance in formulat- ing this roster, which will be put to a vote at the December meeting. In anticipation of this evening’s presen- tation, Edd Barrows displayed a jar of fleas (presumably Ctenocephalides felis (Bouché)) from his family dog. The speakers for the evening were Ralph P. Eckerlin, Professor, and Harry F. Painter, Professor Emeritus, Natural Sciences Di- vision, Northern Virginia Community Col- lege, Annandale Campus, whose talk was entitled “Studies on the Fleas of Virginia.” For almost two decades, these indefatigable siphonapterists have braved the elements and some quizzical citizens in an effort to catalog and comprehend the Old Domin- ion’s little-known flea fauna. By trapping small mammals, retrieving road kills, and— perhaps most important—processing host nests in Berlese-Tullgren funnels, they have determined that flea diversity is lowest on the warm, humid coastal plain (11 species), increases in the Piedmont (17 species), and is greatest in the high mountains bordering West Virginia (26 species). Moreover, the number of flea species tends to increase from south to north, while the number of host species increases in the opposite direction. Similar observations from other parts of the world appear to confirm the general rule PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that flea species diversity increases as tem- perature and humidity decrease. Drs. Eck- erlin and Painter speculate that as many as 45 flea species may be present in Virginia because much of the state lies in a transition zone between austral and boreal elements. They have recently lengthened the list of fleas from Fairfax County in northern Vir- ginia, and they have conducted detailed in- vestigations of the seasonal dynamics of fleas associated with the southern flying squirrel, Glaucomys volans (Linnaeus), in the Vir- ginia Piedmont. Collections of fleas from any part of the state will be warmly wel- comed as contributions toward an eventual monograph in the /nsects of Virginia series, Virginia Polytechnic Institute and State University, Blacksburg. In keeping with the evening’s subject matter, R. G. Robbins could not resist the itch to introduce Dr. Robert Traub, the world’s foremost authority on fleas and, by recent vote of the American Committee of Medical Entomologists, only the third re- cipient of the prestigious Harry Hoogstraal Commemorative Medal, which will be con- ferred at this year’s Annual Meeting of the American Society of Tropical Medicine and Hygiene, in Honolulu, Hawau. Though justly revered for his siphonapterological studies, Dr. Traub achieved transcendent fame for his central role in the chemopro- phylaxis of chigger-borne rickettsiosis (Rickettsia tsutsugamushi) during and after World War II, work that earned his team a nomination for the Nobel Prize. Beyond being a renowned scientist, Dr. Traub is one of a fast-fading élite of Renaissance men that included his lifelong friend and col- league Dr. J. Ralph Audy (1914-1974), whose charming book Red Mites and Ty- phus (1968, Athlone Press, London) cap- tured the romance of medical acarology’s golden age. On this occasion, Dr. Traub was accompanied by his sprightly wife Renée; both received the warm applause of the membership. R. G. Robbins also announced the im- VOLUME 92, NUMBER 2 minent departure of Dr. Lance A. Durden, who has accepted a position as a Research Associate in the Department of Entomol- ogy, Auburn University. Dr. Durden’s re- search interests center on the systematics and ecology of the sucking lice (order An- oplura). Though only 34 years old, he has already published 12 papers on Anoplura and 33 papers on other ectoparasite groups, especially fleas, ticks, and chiggers. Indeed, at the last National Conference of the En- tomological Society of America (which Dr. Durden was unable to attend), Dr. K. C. Kim of Pennsylvania State University not- ed during an open seminar that of the world’s five experts on Anoplura, Lance Durden is the only one who is under 50 years of age. In other words—and this point could not have been lost on such a perceptive audi- ence—he has already achieved what has cost all others the better part of their careers! For the last two years, Dr. Durden has worked in Dr. Traub’s laboratory at the Smithson- ian’s Museum Support Center in Suitland, Maryland. With a round of applause, the Recording Secretary wished Lance God- speed and a productive tenure in Alabama. Mignon Davis took photographs of this evening’s large audience and of the five ec- toparasitologists present (the latter, a group photo, did not develop). She also circulated copies of a bill (CB-74-1989) recently in- troduced to the Council of Prince George’s County, Maryland, that, if enacted, would require developers to preserve mature trees, specimen trees, or large clumps of trees ex- isting on subdivisible properties. Though Maryland already has proportionately less forest cover than any other mid-Atlantic state save Delaware (2,653,200 acres or just 42% of the Old Line State’s area), this mod- est proposal is being resisted by a powerful construction lobby that caters only to Wash- ington’s ceaseless craving for Lebensraum. Visitors were introduced and the meeting was adjourned at 9:45 p.m. Refreshments included a rainbow cake, compliments of 369 our youngest guests, Marisa and Steven Da- VIS. Richard G. Robbins, Recording Secretary 953rd Regular Meeting— December 7, 1989 The 953rd Regular Meeting of the En- tomological Society of Washington was called to order by President F. Christian Thompson in the Naturalist Center, Na- tional Museum of Natural History, at 8:00 p.m. on 7 December 1989. Twenty mem- bers and nine guests were present. Minutes of the November meeting were read and approved. Membership Chairman G. B. White read the names of the following applicants for membership: Edward F. Connor, Depart- ment of Environmental Sciences, Univer- sity of Virginia, Charlottesville; Donald F. J. Hilton, Department of Biological Sci- ences, Bishop’s University, Lennoxville, Quebec, Canada; and David M. Pollock, Enid, Oklahoma. Chairman White also re- ported that the Society gained 30 new mem- bers in 1989, bringing our total to 561. Annual reports were given by the Cor- responding Secretary, Treasurer, and out- going Editor (the last-named two read by F. C. Thompson). The President then warmly thanked all officers and volunteers for help- ing to maintain our Society as this nation’s premier regional entomological organiza- tion. President Thompson reviewed the slate of nominees for officers in 1990 and called for additional nominations from the floor, of which there were none. A motion was made and seconded that the slate be ac- cepted as presented. The motion was ap- proved by acclamation. Mignon Davis announced that the tree preservation bill described in the Novem- 370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ber Minutes had been passed by the Council of Prince George’s County, Maryland. Corresponding Secretary J. M. Kingsol- ver exhibited three absorbing entomologi- cal texts: Larvas de Coleoptera do Brasil, by Cleide Costa, Sergio A. Vanin, and Sénia A. Casari-Chen, 1988, Museu de Zoologia, Universidade de Sao Paulo; A Manual of Forensic Entomology, by Kenneth G. V. Smith, 1986, British Museum (Natural His- tory) and Comstock Publishing Associates, a division of Cornell University Press, Ith- aca, New York, ISBN 0-8014-1927-1; and Pictorial Guide to Insect Pests of Stored Food Products, by Toshiharu Yoshida, Naoshi Watanabe, and Mochiyuki Sonda, 1989, Zenkoku Noson Kyoiku Kyokai Publishing Co., Ltd., Tokyo. President-Elect J. R. Aldrich displayed the first offspring of a female southern green stink bug (Nezara viridula (Linnaeus) (He- miptera: Pentatomidae)) collected in a field of soybeans (Glycine max (Linnaeus) Mer- rill) in Beltsville, Prince George’s County, Maryland, on 4 October of this year. Ap- parently a victim of hurricane Hugo, this specimen was blown some 600 km north of its known range. It has subsequently pro- vided Dr. Aldrich with six egg masses. Edd Barrows exhibited cocoons and adults of the sawfly Priophorus morio (Lepeletier) (Hymenoptera: Symphyta: Tenthredini- dae), together with photographs of larval damage to raspberry (Rubus idaeus Lin- naeus). Dr. Barrows’ specimens were col- lected in Montgomery County, Maryland, which is south of this species’ range. How- ever, the rarest of Edd’s exhibits this eve- ning were photographs of our intrepid Pres- ident—afield and stalking the wily syrphid! The speaker for the evening was Steven L. Heydon, Visiting Scientist, National Mu- seum of Natural History, Smithsonian In- stitution, whose talk was entitled “‘Desper- ately Seeking Salix: Interactions Between a Willow Stem-galling Agromyzid and its Two Parasitoids.”’ Galls on willow stems are fre- quently the work of larvae of the agromyzid fly Hexomyza salicis (Malloch). Adult fe- males of this univoltine species oviposit in early May, each egg-laying event leaving a distinctive scar. First-instar larvae molt in late June, and the second instar feeds on plant fluids throughout the summer and .{: . sc.c Bo .ths pote Woape fee ee earls eee LAMP: W..E.—Plant Stress-Insect Interactions .... 5.5 .:aeqsn sone see oa ee eee ee SOGIETY MEETINGS (25.505 ra ssc mbites de!« cA. cipteteitl piece heat ie RMeias fe Me Sette eee aria eer X VOL. 92 JULY 1990 NO. 3 (ISSN 0013-8797) PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS ADLER, PETER H.—Two new species of black flies (Diptera: Simuliidae) from North America BAKER, GERALD T. and SONNY B. RAMASWAMY.—Tarsal and ovipositor sensilla of Heliothis virescens and H. subflexa (Lepidoptera: Noctuidae) .......................-- 521 BAO, NONGGANG and WILLIAM H. ROBINSON.— Morphology and mating configuration of genitalia of the oriental cockroach, B/atta orientalis L. (Blattodea: Blattidae) ......... 416 BORKENT, A.—A revision of the Nearctic species of Dicerura Kieffer (Diptera: Cecidomyiidae) 571 COOPER, KENNETH W.-—Likely causes and explanation of probable atavism in a somatically mosaic fly from a wild population (Diptera, Asilidae, Nannocyrtopogon minutus) ....... 373 FREYTAG, PAUL H.—A new genus Nancyana and nine new species with a review of the related genus Rhogosana (Homoptera: Cicadellidae) ............................2.... $30 HEADRICK, DAVID and RICHARD D. GOEDEN.—Resource utilization by larvae of Par- acantha gentilis (Diptera: Tephritidae) in capitula of Cirsium californicum and C. proteanum MASIELACCAL) MISOUERETIN Ca OTN IA ieee cis ore seven tere eis ioe vei. clei sia, adele lads ve tev sisialiacnt gare 512 HESPENHEIDE, HENRY A.—New species of Buprestidae (Coleoptera) from the Dominican RES PIO LIC ere ee rete ee ree Ree etree eae ter ae ty na ERA ees Son h. gait oea) Shateseca beste sale a ctasebeceooeaa 400 HUNG, AKEY C. F.—Scale-like structures on the tibia of the parasitic wasps, Trichogramma RHP ary menOoptera-wniCnOprammMatidae)) vaca ea ckctetsial said oe oeitie ele cueiers wot e wietesise sie 548 KIRIAC, ION, FRANCIS GRUBER, TAD POPRAWSKI, SUSAN HALBERT, and LESLIE ELBERSON.—Occurrence of sexual morphs of Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae), in several locations in the Soviet Union and the northwestern UIBTUSa! SHAVES on enlad odibio 4 edn op aOl SOA Geneon AS ce Seto e aeeee e KONDRATIEFF, BORIS C. and JUDITH L. WELCH.—The Nemomydas of southwestern United States, Mexico, and Central America (Diptera: Mydidae) ...................... 471 KNUTSON, L., R. E. ORTH, and R. ROZKOSNY.—New North American Colobaea, with a preliminary analysis of related genera (Diptera: Sciomyzidae) ......................... 483 LINLEY, JOHN R. and DAVE D. CHADEE.—Fine structure of the eggs of Psorophora columbiae, ps. Cingulata and ps. Ferox (Diptera: Culicidae) ..........................- 497 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1990 JEFFREY R. ALDRICH, President NORMAN E. WoobDLey, Treasurer Davip R. Situ, President-Elect Gary STECK, Program Chairman RICHARD G. Rossins, Recording Secretary GEOFFREY B. WHITE, Membership Chairman Ho Luis B. WILLIAMS, Corresponding Secretary F. CHRISTIAN THOMPSON, Past President JAMES B. STRIBLING, Custodian Rosert D. Gorpon, Editor THoMaAS J. HENRY, Associate Editor Publications Committee DONALD R. DAVIS WAYNE N. MATHIS GEORGE C. STEYSKAL F. CHRISTIAN THOMPSON Honorary President Curtis W. SABROSKY Honorary Members Louise M. RUSSELL ALAN STONE THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $20.00 (U.S. currency) of which $18.00 is for a subscription to the Proceedings of the Entomological Society of Washington for one year. PROCEEDINGS. — The Proceedings are published quarterly beginning in January by The Entomological Society of Washington, % Department of Entomology, NHB-168, Smithsonian Institution, Washington, D.C. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $50.00 per year, domestic, and $60.00 per year, foreign (U.S. currency), payable in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE P. 318 OF THE APRIL, 1989 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. Editor: Robert D. Gordon, Systematic Entomology Laboratory, ARS, % Department of Entomology, Smith- sonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: T. Henry, Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. i ———— This issue was mailed 9 August 1990 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA weer PLL PRL OC ™ A PERRI MPA AIPe rm FRR CUA KEI. OP Ree RPA DER PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 373-384 LIKELY CAUSES AND EXPLANATION OF PROBABLE ATAVISM IN A SOMATICALLY MOSAIC FLY FROM A WILD POPULATION (DIPTERA, ASILIDAE, NANNOCYRTOPOGON MINUTUS) KENNETH W. COOPER Department of Biology, University of California, Riverside. Abstract. — Description is given of an otherwise normal male asilid (of a genus of 28 species having either hyaline or lightly infuscated wings) with its left wing strikingly color- patterned. This appears to be the first recorded not-gynandromorphic, not-parasitized somatic mosaic in Diptera apart from laboratory cultures and experiments. Possible genetic origins of such mosaicism, of phenotypic expression, and their consequences are outlined. Despite lack of relevant fossils, the more plausible conclusion is that the wing pattern is primarily atavistic and not a neomorphism. Mutants calling forth ancestral attributes do not differ qualitatively from those altering familiar, ‘“‘lesser’”” phenotypes. Ancestral phenotypic attributes probably regularly disappear long before their genetic mechanisms pass beyond the capacity for reexpression, as substantiated by disappearance and reoccurrence of R, in Brachycera. Key Words: What is to be made of a chimaeric male robber fly, Nannocyrtopogon minutus Wil- cox and Martin, otherwise normal, having the blade of one wing palely infuscate and normal for the species, the other wing dis- playing a striking color pattern that is un- usual even among asilid species normally having maculated wings (Figs. 1-3, 5)? This in a genus in which the 28 other species do not have color patterns on their wings, being nearly equally divided between those with hyaline and those with lightly tinted wing membranes. The explanation must largely be genetic. Because genetic systems are subject to mutation, errors of mitosis and fertilization, individuals of a population may be viewed as having their bodies potentially subjected to partition into two or more genetically different sectors during development. In most species, individuals are regarded as “normal” if no disparate sector is detected. Atavism, somatic mosaicism, genetics, wing maculation, wing venation Those having from more than 0 to as much as 50% included in such sectors are termed mosaics. Though ordinarily rare, the commonest mosaic detected in wild populations of in- sects is the gynandromorph, in which the body is partitioned into genetically and phe- notypically sexual sectors. Such sexual mo- saics, not to be confused with intersexes which are of uniform genotype, have been found in many orders of insects and in many families of flies, though not in the Asilidae.! 'No museum dipterist of whom I inquired could recollect having read of or seen any mosaic asilid (either gynandromorphic or not-gynandromorphic). It is un- likely that striking anomalies of asilids go unnoticed (e.g. see Weinberg 1973). Yet no asilid gynandromorph or other mosaic is recorded by either Zoological Record or Entomological Abstracts (to Volume 20(5), June 1989) within the years they cover for the interval 1925— 1989, nor did Collin (1927) mention any earlier records in his brief review. 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 3 Mosaics of a single sex, with an aspect suf- ficiently striking to be noticed, are well- known but not common in Lepidoptera (Cockayne 1924, Robinson 1971) where striking individual variations in color pat- terns of wings rarely escape notice. Else- where among insects, as with flies in wild populations, they appear to be of a second order of rarity. Apart from certain Nema- tocera infested with parasites, those recoid- ed for Diptera, of which I am aware, are derived from laboratory cultures and ex- periments. The earliest general account is that by Morgan and Bridges (1919) for Dro- sophila. It is assumed that the genetics of asilids, like that of most functionally diploid insects which have been studied, does not depart in any general or unique way from that of species of Drosophila. The phenotypic ef- fects of asterisked mutant alleles of D. mel- anogaster Meigen mentioned in discussion (e.g. *Lyra), unless another reference is giv- en, will be found in Lindsley and Grell (1963). There are accordingly two probable an- swers to the large question posed by the mosaic asilid, each with more than one pos- sible explanation. They are: the left wing of the mosaic fly may provide a preview of a remarkable apomorphy potentially realiz- able in the future, or it may display in near entirety a purely atavistic trait. THE CAPTURE The male mosaic was one of a total of 5 males and 2 females of N. minutus collected 375 on July 20 and 26, 1988; the second search was made with the hope that others would be found, perhaps with both wings macu- lated. The site is approximately 5.2 km northwest of Fawnskin, San Bernardino Co., Ca., at an altitude of roughly 1890 m, not far from some of the formerly recorded sites at which N. minutus has been collected. The species is probably generally distributed in the San Gabriel and San Bernardino Moun- tains (Wilcox and Martin 1957). At mid-day N. minutus sallied after smaller flies from perches on boulders, 30- 50 cm in diameter, in the dry bed of Hol- comb Creek. Most flies were old, to judge from the torn hind margins of the wings and broken or missing macrochaetae. Oddly, they were found only along a particular length of the creek bed, some 30-40 m long. The total number of individuals along that stretch was almost certainly fewer than two dozen, but more than twelve. NORMAL WING COLORATION AND STRUCTURE Because the left wing of the mosaic is strikingly unusual by having a color pattern, it was necessary to determine whether it also differs in less obvious features. Though Wil- cox and Martin’s (1936b) description of N. minutus portrays the overall appearance of the fly, the account of the wing is not ade- quate for close comparison. The following condensed description is drawn from wings of the six normal individuals collected at Holcomb Creek, ten from the University of California collection at Riverside, and from /—— Figs. 1-5. Figs. 1-3. Mosaic male of Nannocyrtopogon minutus Wilcox and Martin, ca. 13 x magnification (actual wing length 4.2 mm). 1 —Left side, 2—dorsal aspect, 3—tilted to display right wing. Fig. 4. Diagram of flattened wing base—the proximal portion of radius has folded over the basal half of the basisubcostal cell, 2. Veins: C—costa, Sc—subcosta, R—radius, M—media, CuA—anterior cubitus, CuP—posterior cubitus, A— first anal vein. Lettered cells: c—costal, sc—subcostal, br—basiradial, bm—basimedial, cup—posterior cubital, a—anal. Numbered cells and cell-like enclosures at wing base: 1—basicostal, 2—basisubcostal, 3—first basi- medial, stem or prearculus cell, 4—basianal cell, anterior to distal limb of 3rd axillary sclerite, 5—“‘cell” anterior to proximal limb of 3rd axillary sclerite (the only hyaline cell at the base of the wing). Other: al—alula. Fig. 5. Diagram of patterned areas and venation of left wing of mosaic male (cf. Fig. 1) roughly portraying depths of coloring and extent of pattern—see description for details of pattern. 376 the right wing of the chimaera. Vein color was viewed by reflected light, wing mem- brane color by transmitted light. The small- est details mentioned, and common to all, were determined at a magnification of 50x, and checked at 250 x in two wings softened with KOH and mounted in euparal.? Where possible, nomenclature of veins and cells (Fig. 4) follows McAlpine (1981). As usual, fluting of wing along longitu- dinal veins pronounced; veins dark sepia, somewhat lighter as they thin distally; vein MA (arculus), crossvein sc-r, veins CuP and A, weakly developed; membrane hyaline throughout, coloring microtrichial (prop- erly acanthal, type b [Richards 1979]); cell c subhyaline to very pale fuscous; cells be, bsc, stem cell [Shannon’s (1924) “‘prearcu- lus cell’’], extreme base of cell a,, pale to light brown; “‘cell”’ bounded above by prox- imal stem of MP to A,-complex, and an- terior basal portion and proximal lateral apophysis of third axillary sclerite, hyaline (Fig. 4, ‘cell’ 5); cell sc ight brown in apical half; short, pale brown streak proximally between veins CuA and CuP in some; a broad, bare hyaline band along length of posterior margin of CuP; remaining mem- branous areas of wing very light to pale brown, gradually paler posteriorly and ba- sally; alula in part very pale brown, or not (Fig. 4, al). THE MosalIc Apart from the surprising 3-partite col- oration of 3 grades of saturation marking the blade of the left wing (Figs. 1, 2, 5), a slightly more exaggerated fluting along its > At 250~, slide preparations show an approxi- mately | 2-partite internal ‘“‘annulation” of MP, or bul- la, immediately before its bifurcation, without a thyrid- ial clear spot, that is not ordinarily detectable at 50 x in pinned specimens. Campaniform sensilla occur along the basal margin of the tegula (ca. 18), at base of Sc (20+) and adjacent to its junction with crossvein h (8- 9), at base of vein R (ca. 70) and widely scattered along its length (6-8) just before and following separation of Rs. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON longitudinal veins, and minor defects to be discussed, the mosaic specimen is a normal male. In appearance it corresponds well with Wilcox and Martin’s description. External morphology of head, antennae,’ thorax, ab- domen, terminalia, legs, patterns of prui- nosity and setation, right wing coloration, sizes and venational patterns of both wings (cf. Figs. 1, 3), and body coloring—even at the regions of the thorax bearing the wings— are typical of N. minutus. Left wing stalk, venation, color of the five small “cells” at the stalk of the wing, and general light to pale acanthal browning of all unaffected areas of the wing blade normal for the species. Both membrane and acan- thae are colored brown in sharply delimited regions of the blade, giving a much darker, large basal area and a smaller, much less dark one apically, separated by a continuous broad region of a normal light to pale tint (Figs. 1, 2, 5). Thus: cell c pale brown, lighter distally; cell sc light brown from origin to apex, cell br dark brown, somewhat lighter along anterior half; cell bm dark brown, nar- rowly paler in proximal third along vein CuA; bases of cells r, and r,,, dark brown nearly to a line connecting the distal end of vein Sc to crossvein r-m; r, lighter along veins R, and R,,,; nearly basal 0.4 of cell d (to a point below cross vein r-m) dark brown, darkest basally and along veins M, , 5 and M;; basal 0.3+ of cell m; and basal third of CuA successively lighter brown. Ex- cept at apex, remainder of blade very light to pale brown as in the unaffected right wing. Basal half of wing therefore presents a strongly contrasting dark brown macula in the shape of a slightly opened fan, given added emphasis by the fluting of the wing along the longitudinal veins. Wing tip with a sharply bounded apical lunule, extending from near apex of cell r, to near midpoint of outer margin of cell m,; greatest width at cell R, nearly one-eighth length of wing; much paler than most of } Broken off during photography. VOLUME 92, NUMBER 3 B) basal “‘fan,” similar in lightness to color- ation at base of cell cua only; everywhere contrasting strongly with the lightly tinted adjacent membrane (Figs. 1, 2, 5); alula palely infuscate. DEVELOPMENTAL DEFECTS OF THE MACULATED WING The left wing’s length (corrected for cur- vature), veins, venational pattern of cells, and outlines of the wing’s margin are all normal. Apart from a slight positive cur- vature of the blade, physical abnormalities occur in pigmented areas only and appear minor; indeed, detectable only at higher magnifications. At 25 =, seven tiny dorsal blisters are visible: two near the distal end of cell bm (70 and 40 um in greatest di- ameters), and five in the lunule: one in cell r; (20 um); one in r, (30 wm); and three in m, (10-20 um). At 50x, the dorsal mem- branes over the maculae seem very slightly thicker than the surrounding not-maculated membrane. These abnormalities are explicable as re- sults of a slight but consistent incoordina- tion during the terminal stages of dorso- ventral epithelial contraction of the pupal wing. Their minor nature contrasts mark- edly with the often extreme abnormalities to be seen among the phenotypes of mutant genes affecting the wings of Drosophila (see Waddington 1940, 1942). AsILID WING PATTERNS Most asilid wings derive their coloration, when present, from type B acanthae, from membrane pigmentation, or from both; rarely is it structural. The wings of the ma- jority of Nearctic asilids range from hyaline through tinged to full color (usually browns to nearly black), or have a gradually deep- ening color along an axis. A minority have discrete, maculated patterns. The common- est of these is a slight clouding or spotting at crossveins and venational branchpoints. Somewhat less frequent, but widespread, is SH a darkening of the wing adjacent to the apex, often in the form of a lunule. Nearly the full range of wing coloration is shown by Nearctic species of Cyrtopogon, of which there are some seventy, and from which Nannocyrtopogon was split by Wil- cox and Martin (1936a). Some, as C. dasyllis Williston, C. maculipennis (Macquart), and the male of C. bimacula (Walker), have large, striking patterns, but in each the principal macula is not proximal, nor is the apex mac- ulate. As Dr. Eric Fisher pointed out to me, however, at least three Palearctic species of Crytopogon do have a truly apical macula separated by clear membrane from a more basal pattern; e.g. C. centralis Loew, the most similar of these (see Engle 1929, fig. 222, p. 355). Nevertheless, the basal macula of C. centralis (apically very similar in outline and extent to that of the mosaic) does not reach the base of the wing, nor is the apical macula alunule. Though wing patterns of some Cyr- topogon seem not far removed, none is wholly like that of the left wing of N. mi- nutus. Indeed, none of the species of the other forty-six genera of Nearctic Dasypogoninae* have a compound basal and apical pattern closely similar to that of the chimaera, nor do the remaining Nearctic asilids. How then is the occurrence of the two differently colored wings of the aberrant male fly to be explained, and how may the uniqueness of the patterned wing be under- stood? INTERPRETATION Because the patterned wing of the male chimaera is free of striking abnormality in form, basal coloring, veins, and venational pattern, it is unlikely to have been the direct result of an asymmetrically directed envi- + The Dasypogoninae of Martin’s and Wilcox’ (1965) classification have been split into three allied subfam- ilies by both Papavero (1973) and Lehr (1988); in North America we have: Dasypogoninae (11 genera), Steno- pogoninae (31 genera), and Trigonomiminae (= Tri- gomiminae of Lehr; 4 genera). Nannocyrtopogon and its allies are stenopogonines. 378 ronmental influence. Even were it so, to have responded to that external stimulus in the manner required, the fly’s genotype neces- sarily included within its repertoire an oth- erwise unexpressed capacity to provide the biochemical and developmental prerequi- sites for production of a nearly unblemished wing with that particular pattern, as later explained. If an external influence was in- volved, most probably it only indirectly in- stigated the necessary genotypic response (see below). Among many conceivable genetic expla- nations, two well-known sporadic events may equally well account for the patterned wing. Because male asilids of known karyo- type are either XY or XO, and females XX (Makino 1951, Cooper, unpublished), the fact that the chimaera is a male places a different restraint on the nature of the chro- mosome involved in each case. These events are: 1) Somatic mutation of a gene in the dif- ferential segment of a sex chromosome (+s/o — s/o),> the new allele’s recessive phenotype therefore being expressible in its present hemizygous state. The mu- tation may have arisen by action of an external agent (e.g. by environmental ra- diation, mutagens, etc.), or internally (by replicative error, transposon, etc.). 2) Somatic crossing-over (see Stern 1968) in an autosomal heterozygote for an al- lele (a) giving a recessive phenotype, namely (+a/a — a/a and +a/+a equal- ly). Studies of such crossing-over in Dro- sophila led Stern (1936) to conclude that it may in fact prove the most likely cause of somatic mosaicism when suitable het- erozygosity 1s present. If the frequency of the allele (a) were as high (but no higher) than 0.17 in the pop- > The “to” in these formulae indicates that there is no genic portion of the alternative sex chromosome, if any, that possesses the wild-type allele (+s), or a gene that suppresses the phenotypic action of (s). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ulation of N. minutus, more than 160 flies (a number considerably larger than that of the reported and probable specimens now in collections) would be required for a 99% likelihood that at least one (a/a) individual with both wings patterned would be includ- ed within the sample. None has been re- ported, or described as a new species, as would be likely had such a specimen been found. The requisite heterozygotes (+a/a), however, would be relatively common (ca. 28% of both sexes). In both cases the genetic change is as- sumed to involve an allele giving a recessive phenotype because most realized mutants with dominant phenotypes are far less com- mon and more likely to produce malfor- mations (catalog in Lindsley and Grell 1968). The change would necessarily occur in a nucleus at an early cleavage division of a preblastodermic egg. In that way a cell of a new genotype (either s/o or a/a) could have given rise to a sufficiently large clone to have formed the imaginal disc of the left wing, and perhaps other tissues of the chimaera. A mosaic arising from somatic crossing-over after the first ““cleavage” division would be a trisectorial mosaic, in contrast to the bi- sectorial mosaic produced by a single so- matic mutation. DISCUSSION However the mosaic arose, it 1s clear that identical modes of genic action may be ascribed to the mutant allele, whether new (s) or preexistent (a). Choices for the results of such genic action in the case of the mosaic are two: (1) a discontinuous phenotypic change qualitatively different from that of wild type, a complex phenotype without precedent; in effect a preview of a potential apomorphy in the descendants of N. mi- nutus; or (2) a recovery of an ancestral wing pattern, or nearly so; an atavistic expression which, were it found characteristic of a pop- ulation today, would no doubt be viewed as an apomorphy. Whether newly mutated or not, a struc- VOLUME 92, NUMBER 3 tural gene does but one thing: it codes for the production ofa single product. For many mutants, perhaps most, that product ulti- mately may play an active role in more than one biochemical pathway in development, giving rise to one or more seemingly unre- lated phenotypic effects. Such an allele is said to be “pleiotropic” in its action. Thus *Lyra of D. melanogaster affects the eyes, body setae, wings, abdominal tergites and color; when homozygous it is lethal. Of the thousand or more loci for which mutant alleles are now known in D. mela- nogaster (Lindsley and Grell 1968—“... reasonably complete through 1966’’), most of the alleles have adverse pleiotropic ef- fects. Alleles at nearly a third of the loci have an effect upon the wings, and about an eighth affect only the wings in one or more ways (catalog in Braver 1956). The phe- notypic changes in the wings are almost al- ways anomalous, among which are minor to extreme abnormalities of the blade, of its margins, of venation, of acanthae, of color, retention of hemolymph, and of expansion of the pupal wing at eclosion. Though many Drosophila species have maculated wings, including males of some members of the melanogaster subgenus (Sophophora) Bock and Wheeler (1972), none of the known phenotypes of male or female D. melano- gaster take the form of a wing with a color pattern. Those mutant alleles which do produce a new coloration of the blade without accom- panying abnormalities of the wing are but a tiny minority of all; e.g. fuliginosus (Buz- zati-Traverso 1947), *lemon, *pallid and *yellow. All such alleles at the four loci, except one (y’’?>, Gianotti 1951), affect both body and wing color in similar ways. Their primary effect is evidently upon the capacity of epidermal cells to produce particular melanins rather than an exclusive effect upon the epidermal cells of the wing itself. The latter appears to have been the case for y*°’”. Ifthe very extensive observations on Dro- sophila reflect in a general way attributes of 379 mutations of flies, then comparable muta- tional changes affecting only coloration of the blade of the wing are expected to be extremely uncommon. Compared with the mutant alleles that affect the wings of D. melanogaster, that presumed in N. minutus to have brought about maculation of the left wing is aston- ishing in the complexity of its phenotype and freedom from gross malformation. The phenotype leaves coloration of the small ve- national “‘cells” in the stalk of the wing (Fig. 4, “cells” 1-5) and most of the membrane of the blade unaffected (Figs. 1-3). How- ever, it selectively heightens the levels of pigmentation, to very different degrees, in two unequally shaped, large, well-separated groups of contiguous epidermal cells (cf. Figs. 3 and 1, 2, 5). The pigmentation of the newly maculated areas is cell-produced and cell-limited; the boundaries between pig- ment cells and adjoining normally colored membrane are therefore sharply defined. Even venational cells r, and r,,;, the bases and apices of which are of greatly different intensities of brown, show not the slightest signs of a decreasing color gradient from dark to light. If all this resulted from a single product coded by a new allele, that product must have enhanced pigment formation (which awakens no problem) yet have benignly ac- tivated a series of coordinated pathways not ordinarily revealed by a difference in pig- mentation basally and apically, nor by any partitioning of the wing into such special domains other than by veins. No mutations recorded for D. melanogaster produce de novo comparably complex, well-ordered phenotypes in any structure without notable abnormality. The circumstances appear to call for another interpretation of the origin of the pattern. In perhaps most populations there is a phenotypically unexpressed retention of ge- netic bases for one or more ancestral attri- butes. Indeed Garcia-Bellido (1983) com- mented that “... it is not impossible .. . 380 that most new patterns found in evolved groups of Drosophila are ancestral pat- terns.”’° In fact, Richards (1958) has dem- onstrated just such a case in Ephestia. Fur- thermore, Sondhi (1962), by continued selection in a strain of D. melanogaster, was able to produce a wholly new pair of bristles, in a particular location, comparable and presumably homologous with those found in the related family Aulacigastridae, and very probably with those in an ancestor of the two families. Causes and means for con- tinuance of such apparently “‘silent”’ genic presences within the genome are discussed by Regal (1977), Riedl (1977), Hall (1984), and Coyne and Prout (1984) among others, along with examples from a variety of reac- tivated phenotypic expressions of such con- cealed bases of ancestral attributes other- wise known only from fossils. Gauld and Mound (1982) have discussed apparently frequent reversals and the problems they necessarily awaken in phyletic analysis. To most there would seem to be an un- bridgeable gap of complexity between most “ordinary” mutations and one that seems to call forth a probable attribute ofan ances- tor of countless generations removed. Is that so? A mutant allele that restores the expres- sion of an ancestral attribute does not differ from other mutant alleles with less striking phenotpes in kind, in degree, in mode of action in a developmental pathway, or even necessarily in the phyletic age of the path- way affected. It differs solely by its chance triggering and disclosure ofa latent, ancient, yet still potentially expressible system with- in the genome. The difference is therefore not the nature of the mutation, but resides in a special peculiarity of the genome it- self—a retained but suppressed integrated system, a “‘prepattern,” in this case for wing maculation. ° 1 would add “perhaps most often in a somewhat modified form because of their reexpression within a changed genetic milieu.” PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The mutant allele codes for a product just as in other cases, but that product makes biochemically possible release and expres- sion, wholly or in part, of the existant co- ordinated but “silent” ancestral pathways within the present genetic system. No chance pleiotropic concatenation of pathways to produce a coordinated wing pattern need be involved—that Achilles heel of the hypoth- esis of a wholly new phenotype. They al- ready exist in a coordinate relation owing to prior evolution. The minor abnormalities expressed in the patterned wing of the mo- saic may owe either to a pleiotropic effect of the mutant, or they may reflect a loos- ening of developmental timing within the retained ancestral system now being reex- pressed against the milieu of new mutations accumulated since suppression of the an- cestral wing pattern became a lineage at- tribute, or both. In any case, the reappear- ance of an ancestral wing pattern (or close thereto, perfect reversion being unlikely) seems to me the more plausible interpre- tation of the left wing of the mosaic N. mi- nutus. In the absence of evidence from fossils’ of likely ancestral stocks, atavism cannot be disproven or proven for the wing of the mo- saic. Nevertheless it does seem plausible be- cause the complex maculated pattern in a nearly perfect wing of N. minutus appears otherwise as a freak of nature, for all 28 species of Nannocyrtopogon have either hyaline or lightly infuscated wings. How- ever, as earlier mentioned certain species of ’ The relevant Oligocene-Miocene fossils are as- signed by their authors to stenopogonine genera con- temporaneous with Cyrtopogon and Nannocyrtopogon (namely Ceraturgus [as Ceraturgopsis], | sp.; Dioctria, 2 spp.; Holopogon, 2 spp.; and Microstylum, 1 or 2 spp.). See Hull (1962) and Papavero (1973) for refer- ences and comment. So far as can be told all have either hyaline, not-maculated, or infumated wings. However, absence of maculations in a fossilized wing is not of itself reliable evidence for a corresponding absence of color pattern in the wing prior to fossilization (see Car- penter 1971). VOLUME 92, NUMBER 3 the presumed sister group, Cyrtopogon, do have strongly maculated wings of interre- lated patterns, some with both a central and apical macula. Those patterns are some- what less complex, and the maculations dif- ferently shaped, defined, and placed than those of the left wing of the mosaic. Because de novo origin of such a complex pattern by a single mutation is highly improbable, and by simultaneous multiple mutations im- plausible, it is reasonble to assume that Cyr- topogon and Nannocyrtopogon shared an ancestor with a patterned wing, and that the means for wing patterning was retained in both lineages.*® In the line from which Nan- nocyrtopogon species were derived, how- ever, expression of pattern was suppressed. Retention of the suite of ancestral mutants involved presumably owes to their still es- sential contribution to one or more stages of development. Only their inessential ac- tions, as those leading to an expression of a pattern, are genetically suppressed. The new mutant (s) or homozygote (a/a) then codes for a product of which the ultimate effect is reactivation of suppressed pattern pathways. I now turn to another atavism, wide- spread among asilids and other Brachycera, that does not appear to have received the attention merited. Hennig (1954) raised the question as to whether the presence of vein R, in the asilid Promachus and its apocleine relatives represents an atavism. He thought not, although he left the question open for mydids and possibly others in which only the distal stub of R, or its trace remains. R, occurs also in the genus Pseudorus which is only remotely related to Promachus (Pa- pavero 1973). Shannon and Bromley (1924) ® Contrary to Meyere’s (1907) opinion, ancestral flies, especially those of the Nematocera and of early Asi- lomorpha, probably did not have hyaline wings (all had epidermal “melanin”-producing pathways). Prob- ably wing maculations of one sort or another were at least as common among them as they are among mod- ern forms. 381 indicated the presence of R, in Pogonoso- ma, another asilid of rather remote affinity to both Pseudorus and Promachus, and in one or another form in other asilids, many bombyliids, some leptids, mydids, tabanids and occasionally in therevids. Very likely R, in these and perhaps other families is a vein tending to widespread reduction and loss at individually different rates through- out the Brachycera (in many families it has already been lost), a kind of “‘orthogenesis,”’ much as appears to be happening to the basal length of M in flight wings of beetles. The isolated, regular occurrence of a com- plete R, in certain species of Pseudorus (as Papavero 1973 suggests; see figs. 4, 5 in Old- royd 1964), and in the 18 or so species of Pogonosoma, may represent recurrences rather than prolonged retention of R, be- yond that of the numerous other members of their subfamilies. Certainly the well- known “‘anomalous” occurrences, most fre- quently asymmetrically, of a remnant of R, (as a “stump vein,” = Tillyard’s 1919 ‘in- terradial crossvein”) or, more rarely, in complete form in individuals of species nor- mally lacking all traces of R;, are to be re- garded as atavisms. Such individuals dem- onstrate that suppression of the R, phenotype is still not complete in their species, and that frequency and penetrance are low for the gene(s) still capable of re- storing the phenotype. Unlike the maculat- ed wing of the mosaic N. minutus, such cases do not require genetic mosaicism. The general notion that the loss of an at- tribute in evolution, especially loss of a complex one, tends to be unrecoverable in later descendants was first proposed by Meyrick (1884), and apparently indepen- dently by Schlosser (1890), Gadow (1893), Dollo (1893) and, for plants, by Arber (1919). Such “lost” attributes, perhaps most of them, probably become unrecoverable in recognizable form only long after complete disappearance by genetic inactivation of their obvious phenotypic expression from a population. That is, only after potential co- 382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ordination of their genetic bases has finally been lost from the genome are they beyond mutational recall. Loss of a phenotypic attribute and total loss of its recoverability have different im- mediate causes, and probably regularly oc- cur stepwise over long but varying intervals of time. Just as that last known occurrence of an extinct form gives an unreliable date for de facto extinction (witness the coel- acanth Latimeria), so also the time of final loss of recoverability of an apparently van- ished attribute within a lineage must gen- erally remain a matter of guesswork. CONCLUSION Though both somatic crossing-over and somatic mutation (or still less frequent ge- netic events) may formally account for the origin of the mosaic’s patterned wing by reactivation of suppressed genetic pathways of distant ancestors, somatic mutation seems the simpler, more likely hypothesis. Both hypotheses predict certain possible outcomes by which they may be differen- tiated: If the mosaicism was caused by somatic crossing-over in a fly of a population car- rying an autosomal allele established at a moderate frequency, whether or not sper- matogonial cells were included within the new (a/a) section, it is possible that a male or female will be found with both wings displaying the striking new color pattern in one or another population of N. minutus. No significant sexual difference in frequency would be expected were numbers of such flies found. Additionally, similar wing mo- saics may turn up in the future because con- ditions for their formation are present in the population. On the other hand, if a newly mutated sex-linked gene (s) were the cause of the mosaicism, no future finding of individuals with both wings maculated would be ex- pected unless the mutant sector included at least some spermatogonial cells in addition to the left wing’s imaginal disc. Even so there would be but a small likelihood of (s) en- tering and persisting in the local population at Holcomb Valley. It would depend on the male’s success in leaving (s/+) female prog- eny, sampling error, and local population size. If (s) did persist in the population, no flies with maculated wings would be ex- pected in the first filial generation. There- after males with maculated wings, though rare, would be greatly more frequent (about two orders of magnitude) than such females. If (s) did not persist in the population, re- occurrence of a similar wing mosaic, or of flies with patterned wings, would require a new mutation to the same allele or an isoal- lele. DEPOSITION OF SPECIMEN For the present the specimen remains in my possession. ACKNOWLEDGMENTS I am indebted to Dr. Eric M. Fisher (Calif. Dept. Agric.) for identifying the asilid. He and Profs. E. Gorton Linsley (U. Cal., Berkeley), John Pinto (U. Cal., Riverside), and my former colleague in genetics, Dean R. Parker, most helpfully commented on an earlier draft of the article. To my colleagues William L. Belser and Karl Fryxell I am indebted for their discussions of certain ge- netic implications awakened by the mosaic. Saul Frommer, curator of insect collections at U. Cal., Riverside, generously provided facilities for the study of the fine collection of Asilidae in his care. To Profs. Hampton L. Carson (U. Hawaii), Henry A. Hespen- heide (U.C.L.A.), R. Lavigne (U. Wyo- ming), C. Riley Nelson (U. Utah), and Drs. Gregg J. Gunnell (Museum of Paleontology, U. Michigan) and Norman E. Woodley (U.S.N.M.), I am grateful for their cordial replies to my queries. None, of course, are responsible for the analysis presented. LITERATURE CITED Arber, A. 1919. On atavism and the law of irrever- sibility. Amer. J. Sci. 148: 27-32. VOLUME 92, NUMBER 3 Bock, I. R.and M. R. Wheeler. 1972. The Drosophila melanogaster species group. Studies in Genetics, VIII, Univ. Texas Pub. 7213: 1-102. Braver, N. 1956. 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Some special features of the wings of Diptera. Insec. Inscit. Menstr. 12: 34-36. Shannon, R. C. and S. W. Bromley. 1924. Radial venation in the Bachycera. Insec. Inscit. Menstr. 12: 137-140. Sondhi, K.C. 1962. The evolution of a pattern. Evo- lution 16: 186-191. Stern, C. 1936. Somatic crossing-over and segrega- tion in Drosophila melanogaster. Genetics 21: 626— 730. 1968. Genetic Mosaics and Other Essays. Harvard Univ. Press, Cambridge, xi + 185 pp. Tillyard, R. J. 1919. The panorpoid complex. Part 3: The wing venation. Proc. Linn. Soc. New South Wales 44: 533-718. Waddington, C. H. 1940. The genetical control of wing development in Drosophila. Jour. Genet. 41: 75-139. . 1942. The pupal contraction as an epigenetic crisis in Drosophila. Proc. Zool. Soc. London (A) 111: 181-188. 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Weinberg, M. 1973. Uncas d’anomalie génitale chez 1936b. The genus Nannocyrtopogon (Dip- un Diptére. L’Entomologiste, Paris, 29: 164-165. tera-Asilidae). Ann. Amer. Entomol. Soc. 29: 449- Wilcox, J. ana C. H. Martin. 1936a. A review of the 459. genus Cyrtopogon Loew in North america. Ento- 1957. Nannocyrtopogon (Diptera-Asilidae). mol. Amer. 16: 1-85. Ann. Amer. Entomol. Soc. 50: 376-392. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 385-399 A MORPHOLOGICAL, ALLOZYMIC, AND KARYOTYPIC ASSESSMENT OF THE PHYLOGENY OF SOME LOWER TERMITES (ISOPTERA: KALOTERMITIDAE) PETER Luykx, DAvip A. NICKLE, AND BRIAN I. CROTHER (PL, BIC) Department of Biology, University of Miami, Coral Gables, Florida 33124; (DAN) Systematic Entomology Laboratory, Agricultural Research Service, National Mu- seum of Natural History, Washington, D.C. 20560; BIC (address: Texas Memorial Mu- seum, 2400 Trinity Street, Austin, Texas 78705). Abstract.—Ten species of termites in the family Kalotermitidae were examined mor- phologically, electrophoretically, and chromosomally. The assignment of the recently described species Neotermes luykxi Nickle and Collins 1989, to the genus Neotermes, originally made solely on morphological grounds, is supported by the electrophoretic data. Neotermes castaneus, however, appears to be a sister group to other species of Neotermes and Incisitermes. Observations on the chromosomes suggest that centric fusions, trans- locations involving the sex chromosomes, and discrete genome amplification events have all been involved in karyotype evolution in these termites, but a phylogeny based on chromosomal changes alone does not agree with the morphological and electrophoretic data. The observations suggest that chromosomal changes may be too rapid and wide- spread to be of use in constructing phylogenies in these insects. Key Words: This morphological, allozymic, and chro- mosomal survey of several kalotermitid ter- mites from Florida (and a few from else- where) was undertaken for two reasons: first, to clarify the systematic position of the new- ly described species Neotermes luykxi (Nickle and Collins 1989); and second, to try to resolve an apparent conflict between the karyotypic relations of some members of the family Kalotermitidae (Luykx and Syren 1979) and the systematic relations as described by Krishna (1961). Preliminary observations on Neotermes luykxi had at first suggested that it might simply be a morphological variant of Neo- termes jouteli (Banks), but chromosome counts and preliminary electrophoretic data suggested a closer relation to another Flor- ida termite, /ncisitermes snyderi (Light). A termites, Neotermes, enzymes, chromosomes, cladistics comparison of the new species with a wider range of other kalotermitids was therefore required. In this paper we report the results of an allozymic, morphological and karyo- typic comparison of the new species with nine other kalotermitid species. Early chromosome studies on kaloter- mitid termites from south Florida (Syren and Luykx 1977, Luykx and Syren 1979) revealed the presence of sex-linked trans- locations in several species belonging to dif- ferent genera, but other species in the same genera lacked them. Neither in the presence or absence of these translocations, nor in the chromosome numbers, did there seem to be much correlation between karyotype and systematic position. Therefore this study was undertaken to re-evaluate, using mod- ern cladistic methods, the phylogeny of the 386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Species of termites used in the electrophoretic studies. Number of Genus, Species Colony i.d. No. Collecting Sites Genomes Sampled Cryptotermes cavifrons Banks 547 Elliott Key, FL 4 Incisitermes milleri (Emerson) 357 Mona Island, PR 4 minor (Hagen) SC Lafayette, CA 4 schwarzi (Banks) 490, 540 N. Miami, FL 8 snyderi (Light) 541 Hollywood, FL 4 Neotermes castaneus (Burmeister) 551 Miami, FL 4 jouteli (Banks) é Southern Florida 14 luykxi Nickle & Collins 466, 543 Hollywood, FL 7 mona (Banks) 361A Mona Island, PR 4 Pterotermes occidentis (Walker) 91-N2-125 Tucson, AZ 4 * Colonies 432 (Knight Key), 454 (Bokeelia), 463 (Fakahatchee Strand), 465 (Dania), and 527 (Hollywood). termites whose karyotypes had already been described in a preliminary way. The karyo- types are described in more detail, and the results are evaluated with regard to the question of the role of chromosome changes in speciation 1n these insects. MATERIALS AND METHODS Termite colonies were maintained in the laboratory on the wood in which they were collected, for periods of time ranging from one month to two years. For the electro- phoretic studies, individual termites were homogenized in one or two drops of dilute buffer (0.03 M Tris-citrate, pH 8.5), and the homogenates were absorbed onto filter pa- per wicks. Electrophoresis was carried out on 12% horizontal starch gels, using a 1:1 mixture of starch from Electrostarch Co. (Madison, Wisconsin) and Connaught Lab- oratories Ltd. (Willowdale, Ontario, Can- ada), with either the Tris-citrate buffer sys- tem (pH 8.5) described by Ridgway et al. (1970), or 0.01 M Tris-citrate at pH 6.7. Gel slices were stained according to stan- dard procedures described by Yang (1971) and Harris and Hopkinson (1976). In all cases where there was any doubt about the relative migration rates of electromorphs from different species, the samples from the questionable species were re-run side-by- side. An average of 5 worker termites from each colony were used. Since each colony is a single family (the king, the queen, and their offspring; Santos and Luykx 1985), a single worker from a colony represents two paren- tal genomes, a sample of two workers rep- resents (on average) three parental genomes, and in a sample of 5 workers the probability is .93 that all four parental genomes are rep- resented. The number of genomes sampled for each species, along with the locations of collecting sites, is given in Table 1. Chromosome preparations of meiotic and mitotic cells were made from the testes of reproductive males by methods described earlier (Luykx and Syren 1979, Luykx 1983). Meiotic cells provided the best material for determining the presence or absence of translocations involving sex chromosomes, while mitotic cells were best for determining the number of acrocentric and metacentric chromosomes. For a few species, only a small amount of material was available, and occasionally the quality of the chromosome VOLUME 92, NUMBER 3 preparations from these species left some uncertainty as to the exact number of meta- centrics present in the karyotype. The num- bers given represent our best estimates of metacentrics vs. acrocentrics; we consider it unlikely that they are in error by more than +1 (haploid), a margin of error that does not affect our general conclusions. For the purposes of this study, the chro- mosomal data reported here have been combined with previously published obser- vations on the chromosomes of Crypto- termes cavifrons, Incisitermes milleri, I. schwarzi, I. snyderi, Neotermes castaneus, and N. jouteli (Luykx and Syren 1979). The observations on the chromosomes of /ncisi- termes minor, Neotermes luykxi, N. mona, and Pterotermes occidentis are new. The ob- servations on the chromosomes of Masto- termes darwiniensis Froggatt confirm these previously published by Bedo (1987). Morphological characters were deter- mined from preserved specimens in the col- lection of the U.S. National Museum of Natural History, Washington, D.C. The morphological and allozymic data sets were analyzed, both separately and combined, with the PAUP (Phylogenetic Analysis Using Parsimony) package (ver- sions 2.4 and Beta test 3.0) written by David L. Swofford (Illinois Natural History Sur- vey, 607 East Peabody Drive, Champaign, Illinois 61820). The morphological data were coded and analyzed as an ordered data set, with the transformation series based on outgroup comparison. The allozyme data were analyzed as an unordered data set. To ensure equal clustering power among char- acters with disparate numbers of states, the characters were weighted according to the number of states. Because each state has potential clustering power, a multi-state character has inherently more clustering power than a simple two-state character (Cranston and Humphries 1988). Therefore multi-state characters were down-weighted to be equal to two-state characters. Best es- timates of relationship were obtained using 387 both BRANCH SWAPPING (GLOBAL) and BRANCH AND BOUND subroutines for comparison. Mastotermes darwiniensis was employed as the outgroup for the morphological anal- ysis and the combined data analysis. No allozyme data were available for M. dar- winiensis, SO no outgroup was employed in the allozyme analysis. Instead, the allozyme tree was rooted at the midpoint and inter- preted as an unrooted tree or network. Be- cause a midpoint root is simply placed half- way between the two farthest points on a tree it does not affect character-state trans- formation series, and its removal results in a network. RESULTS Morphology.—Thirteen morphological characters were examined, seven from ima- goes and six from soldiers. When these char- acters were used for phylogenetic analysis, using Mastotermes darwiniensis as the out- group, a single most-parsimonious clado- gram was obtained (Fig. 1A). With the ex- ception of Neotermes castaneus, the results are reasonably consistent with the taxono- my as reflected in the generic divisions. Ex- amination of the morphological data matrix (Table 2) reveals that morphological char- acters are useful in indicating phylogenetic relations at the generic level, but (especially in the genera Neotermes and Incisitermes) are not generally useful in establishing phy- logenies at the species level, nor even in distinguishing species within genera. For most of the morphological characters examined, Neotermes species appear to be more similar than Jncisitermes species to Mastotermes darwiniensis (family Masto- termitidae), the species used as the outgroup for the cladistic analysis. /. darwiniensis, because of several cockroach-like charac- teristics, is generally considered to be the species most similar to the original ances- tors of modern termites (McKittrick 1965, Grassé 1986). The morphology of Neotermes castaneus 388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Morphological traits of some kalotermitid termites compared with Mastotermes darwiniensis. Neo- termes species names: cas, castaneus; jou, jouteli; luy, luykxi; mon, mona. Incisitermes species names: mil, milleri, min, minor, sch, schwarzi;, sny, snyderi; Pt. occ, Pterotermes occidentis; C. cav, Cryptotermes cavifrons; M. dar, Mastotermes darwiniensis. Imago characters: 1: Left mandible, anterior margin of second marginal tooth (1) equal to or (2) longer than posterior margin of first marginal tooth; 2: Right mandible, posterior margin of second marginal tooth (1) equal to or (2) longer than molar plate; 3: Wing, median vein (1) weakly or (2) strongly sclerotized; 4: Wing, median vein (1) closer to radial sector than to cubitus, or (2) midway between radial sector and cubitus; 5: Wing, median vein (1) extends to tip unbranched, (2) or branched, or (3) joins radial sector at two-thirds length of wing; 6: Wing, veins (1) latticed apically or (2) not; 7: Foot (1) with or (2) without arolium. Soldier characters: 8: Pronotum, shape of anterior margin, (1) concave or (2) incised; 9: Pronotum anterior margin (1) smooth or (2) serrated; 10: Pronotum, posterior margin (1) concave or (2) truncate; 11: Antennal segment no. 3, (1) similar to fourth, (2) greater than fourth + fifth, or (3) greater than fourth + fifth + sixth; 12: Eye pigment (1) absent, (2) slight, or (3) heavy; 13: Head shape (1) oval or pyriform, (2) elongate or reticulate, or (3) phragmotic. Neotermes Character Number jou luy mon Imagoes Left mand. l Right mand. ] Wing vein sc. l ] 2 Wing vein pos. . Wing m. vein . Wing v. latt. Foot UWNDN HWY Soldiers 8. Pron. s. ant. 9. Pron. ant. m. 10. Pron. post. 11. Ant. segm. 12. Eye pigm. 13. Head shape NN = = Se WNN— — i) NRwWwNnee Ne Incisitermes mil min sch sny Pt.occ C.cav M.dar 2 2 2 2 l 2 1 2 2 2 2 l 2 1 2 2 2 2 2 1 1 2 2 2 2 2 2 1 2 2 2 2 l 3 ] 2 2 2 y) 2 2 1 1 2 l 1 2 1 1 2. 2 2 2 2 2 1 2 2 2 2 l 1 1 2 2 2 2 l 1 1 2 3 3 2 2 1 1 1 2 2 2 1 1 1 2 2 2 2 1 3 1 is generally similar to that of other species in the genus, but it differs from other Neo- termes in three of the thirteen traits sum- marized in Table 2: in imago wing venation, in the relative length of the third antennal segment, and in the shape of the pronotum in soldiers (characters 5, 10, and 11). Ac- cording to the results of the cladistic anal- ysis (Fig. 1A), the last two of these traits are primitive traits retained from an ancestor (i.e. like the traits in the outgroup species), while the wing venation (character 5) is a derived trait similar to that in the genus Incisitermes. The cladogram suggests that the similarity of this trait in Incisitermes species and Neotermes castaneus is a result of convergent evolution. Neotermes luykxi differs morphologically from N. jouteli only slightly. It is slightly smaller in size and has a somewhat narrow- er soldier postmentum (Nickle and Collins 1989). As described below, however, the two species can be reliably distinguished on the basis of allozyme patterns and chromosome number. Allozymes.—The electrophoretic data are summarized in Table 3. Phylogenetic anal- ysis of the allozyme data alone resulted in 7 equally parsimonious trees. These 7 trees could be combined via majority-rule con- VOLUME 92, NUMBER 3 jouteli N. mona N. castaneus M. darwiniensis | minor — | snyder — 1 miller) C. cavilrons P. occidentis le — | schwarzi «= i N. luykxi [= east A. 1. minor G EN y C. cavifrons =] | | ro __N. jouleti N. castaneus | 1 | N. mona fechwaci) | =—— Lp. occidentis B. Figure 1 Fig. 1. Hypotheses of relationships based on in- dependent analyses of the morphological and biochem- ical data. A. Cladogram based on the morphological data. It is the single most parsimonious hypotheses, with a consistency index (C.I.; Kluge and Farris 1969) of .708. B. Network based on allozyme data. The tri- chotomies are the result of seven most parsimonious trees (C.I. = .879) combined by the majority rule con- sensus method. sensus to yield a single general hypothesis (Fig. 1B). It is clear that the morphological and allozyme results are not congruent (compare Fig. 1A with Fig. 1B). The mor- phology supports the monophyly of Jncisi- termes, the paraphyly of Neotermes, and the sister status of N. jouteli and N. luykxi. But the unrooted allozyme tree suggests that neither Jncisitermes nor Neotermes are monophyletic, and does not support a N. Jouteli-luykxi sister group. However, combining data from several studies into a single matrix is often prefer- able to consensus methods for reconstruct- ing phylogenies (Miyamoto 1985, Hillis 1987). Accordingly, the allozymic and mor- 389 phological data were combined in a single matrix, and this resulted in a single most parsimonious cladogram (Fig. 2). The com- bined result is entirely congruent with the morphological result. In the combined cladogram all the nodes are well defined and all the terminal taxa are well delineated. Of the 30 characters, 12 (1, 2, 4, 6, 8, 9, 14, 16, 17, 18, 24, and 25) were perfect fits (C.I. = 1.0; see legend to Fig. 1) to the final hypothesis (Fig. 2). Of these 12, only one character (24) had no synapomorphic con- tent; the other 11 were all informative. The morphological characters were not good de- lineators at the species level: only 3 of the 10 taxa—Cryptotermes cavifrons, Ptero- termes occidentis, and Neotermes casta- neus—were delineated by any morpholog- ical characters, and even for these 3 taxa over half of the autapomorphies were al- lozyme characters. The rest of the taxa were identified as unique only by allozyme char- acters. (Every HTU (hypothetical taxonom- ic unit) node is, however, defined by at least one morphological character.) The combined tree is in general agree- ment with the one presented by Krishna (1961, fig. 81) for the genera of the family Kalotermitidae, except for the position of Pterotermes, which is here joined with /n- cisitermes and Cryptotermes to form a monophyletic group, whereas Krishna put Pterotermes on a branch together with Neo- termes. The new species Neotermes luykxi could be distinguished from the morphologically similar N. jouteli at nine of the seventeen allozymic loci examined. This finding, along with the clear chromosome differences de- scribed below, leaves no doubt that the two are distinct species. Nevertheless, NV. /uykxi did appear most similar to N. joufe/i among the other species examined by electropho- resis, being indistinguishable from it at eight of the seventeen loci studied. Three of these loci (characters 15-17: AC-C, ADH, and ALD) are synapomorphies for N. /uykxi and N. jouteli, an additional two (characters 25 390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Allozyme (enzyme) data obtained by starch gel electrophoresis on ten species of kalotermitid termites. For each locus, lower numbers indicate forms of the enzyme that migrated more slowly on the gels, higher numbers forms that migrated more rapidly. All enzymes were anodal, except for those designated ‘*-C,” which appeared on the cathodal side of the origin. 0 = enzyme not detected; X = not tested. Species abbreviations as in Table 2. AC, aconitase; ADH, alcohol dehydrogenase; ALD, aldolase; GAM, galactosaminidase; GK, glucokinase; GNDH, gluconate dehydrogenase; GPI, glucose phosphate isomerase; GR, glutathione reductase; LAP, leucine aminopeptidase; MDH, malate dehydrogenase; ME, malic enzyme; PEP, glycyl-leucine peptidase; PGM, phos- phoglucomutase; SOD, superoxide dismutase; XDH, xanthine dehydrogenase. = Neotermes Character Number Locus luy S jou 14. AC l 3 5 I'S AC-C 4 2 2 16. ADH 0 2 2 17. ALD 3 5 5 18. GAM l 1 1 19. GK 6 1 3 20. GNDH 2 + 1 21. GPI 6 3 4 22: GPI-C 2 2 1 23% GR 4 3 2 24. LAP 4 1 1 25% MDH ] 6 6 26. ME 2 2 2 Dill: PEP 2 6 5 28. PGM 6 4 5 29! SOD 2 4 4 30. XDH 1 2 3 and 29: MDH and SOD) are synapomor- phic for the group that includes N. mona, and the remaining three (characters 18, 24, and 26: GAM, LAP, and ME) appear to be primitive characters shared by other species as well. Chromosomes.—Neotermes luykxi and N. jouteli can be readily distinguished on the basis of their chromosomes (Figs. 3e, f). Not only are the chromosome numbers dif- ferent (2n = 45 and 56, respectively), but the N.F. (“nombre fondamental,” the num- ber of major chromosome arms) is different also (haploid, 25 and 28, respectively). Fur- thermore, N. /uyAxi populations have a sex- trivalent; no sex-multivalents of any kind have been seen in N. jouteli. The results of the chromosome study for all species are summarized in Table 4; sev- eral examples are illustrated in Fig. 3. The karyotypes of the ten kalotermitid species mon Wwe NU WeK NK he —_ PhD vO Incisitermes mil min sch sny Pt. occ C. cav 2 2 4 6 2 2 » 4 3 3 2 1 3 », 4 0 4 5 3 6 Xx 6 3 3} 4 1 2 3 4 3 0 3 + 5 6 3 4 2 6 1 i 6 3 2 3 6 5 ~ 1 2 1 3 4 1 2 1 2 2 2 0 3 3 »4 l 1 1 0 2 4 7 4 2 5 3 5 6 3 4 5 3 5 i 3} 1 4 4 5 3 5 5 l 2 5 1 D 3 4 3 4 3 2 3 0 3 examined for this study show considerable variation. Diploid chromosome numbers range from 28 (Incisitermes milleri) to 79 (Pterotermes occidentis). The diploid chro- mosome sets of some species are composed entirely of acrocentrics, while others con- tain, in addition to acrocentrics, from 3 to 22 metacentrics. Some species are without morphologically differentiated sex chro- mosomes, while others have multiple sex chromosomes that in male meiosis form chains or rings containing from 3 to 14 chro- mosomes. The total number of major chromosome arms (the “‘nombre fondamentale,” N.F.) in different species shows relatively little vari- ation compared to the chromosome number itself (see Table 4). For example, the haploid chromosome numbers of Jncisitermes schwarzi, N. mona, and N. jouteli are 16, 23 and 28, respectively, while the total num- VOLUME 92, NUMBER 3 391 I. minor I. schwarzi a 14 4 D6 a1r4 20 eo 46-7124 pa Mast a 2 8.520 57. 27 [7 2274 2 3 297-2 : 2972 a+, %&-. snyderi 11 3 19 5 14 6 A 2 2071 15 [a2 I. milleri 21 5 16 5 22 3 23 i) 27 3 rt 2 18 a 29 1 26 4 21 3 2675 . 2775 C. cavifrons 12 2 29 5 19 oigs 304-4 ati 5 3 9 2 1 1 10 2 13 1 20 6 16 6 23 2 17 vi A 5 . . 2 a4 i3t+2 P. occidentis N. jouteli N. luykxi 27 1 21 2 28 5 24 2 14 5 Z 2 19 1 22 1 13 1 20 4 23 2 1 2 16 3 21 3 27 5 2 2 17 4 30 2 28 | 3 2 20 3 573 20-1 N. mona 18 3 24 ie} 22 1 25 5 20 5 12 3 28 2 26 5 23 1 14 3 29 3 30 te) 24 3 15 2 26 1 16 2 3 2 30 1 17 5 4 2 6 2 10 2 8 2 16 5} 14 2 ie 2 15 7 20 1 N. castaneus 13 To 25 6 19 4 27 6 26 3 28 4 5 2 19 6 V1 2 21 6 25 3 23 4 27 4 24 4 29 4 28 6 30 3 Figure 2 Fig. 2. The final cladogram constructed from the combined morphologic and allozymic matrix. Excluding uninformative characters, the C.I. is .780. The steps associated with the branch connecting the outgroup to the most recent common ancestor of the study group are not shown because they are unimportant in defining the relationships of the study group. Numbers to the left of the hash marks represent characters; numbers to the right of the hash marks represent character states. bers of major chromosome arms for these species are 27, 27, and 28. When the total numbers of major chro- mosome arms are compared among the dif- ferent species, the numbers appear to fall into distinct groups. I. milleri has N.F. = 14; most of the other /ncisitermes and Neo- termes species have an N.F. ranging from 25 to 28; and Pterotermes occidentis and Mastotermes darwiniensis have N.F. = 49 and 52, respectively. It is possible to order the chromosome changes in plausible evolutionary se- quences, assuming for example that the N.F. changes from low to high by some sort of amplification process, that centric fusions increase the number of metacentric chro- mosomes at the expense of acrocentric chro- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 392 al | _* VOLUME 92, NUMBER 3 Ww ‘oO ww Table 4. Chromosomes of several species of kalothermitid termites and of Mastotermes darwiniensis. The columns in the table are: a, species name; /, the reference for the karyotype (1, this report. 2, Luykx and Syren 1979; 3, Bedo 1987); c, diploid chromosome number in males; d, approximate number of metacentric (two-armed) chromosomes in the diploid set; e, the N.F. (“nombre fondamental,” or total number of chromosome arms) of the haploid set; f, the number of recognizable sex chromosomes (usually multivalent). a b c d e f Species Ref. 2n Meta N.F. Sex Chrom Neotermes castaneus 2 38 14 26 VI Neotermes jouteli 2 56 0 28 — Neotermes luykxi 1 45 3 25 Hl Neotermes mona 1 46 8 27 — Incisitermes milleri 2) 28 0 14 _ Incisitermes minor 1 ST 11 25 Ill Incisitermes schwarzi 2 32 22 27 XIV Incisitermes snyder 15.2 45 8 Dy, I Pterotermes occidentis 1 79 17 49 Ill Cryptotermes cavifrons 2 40 2 21 _ Mastotermes darwiniensis jt 8) 98 6 sy — DISCUSSION mosomes, and that a simple sex-chromo- some pair is built up to multivalent rings and chains by means of successive centric fusions and translocations between sex chromosomes and autosomes. On such as- sumptions plausible phylogenies based on chromosomal changes alone can be con- structed; an example is illustrated in Fig. 4. It is clear that this phylogeny bears little relation to that based on morphology and allozymes (compare Fig. 4 with Figs. | and 2). As explained in the Discussion, there is reason to think that chromosomal changes may occur more frequently and become es- tablished in populations more rapidly than the genic changes that accompany specia- tion and that are reflected in the morpho- logical and allozymic variation between species. Therefore the chromosomal data were not added to the allozymic and mor- phological data matrix, and no attempt was made to arrive at a consensus phylogeny using the chromosomal data. One of the reasons for undertaking this study was to clarify the systematic relation- ship of a new termite discovered in south Florida in 1984. Initially thought to be sim- ply a size variant of Neotermes jouteli, it became the subject of a careful morpho- metric study (Nickle and Collins 1989) when it was later found to have a chromosome number different from that of N. joutel/i. On the basis of the morphometric study (Nickle and Collins 1989), it was recognized as a distinct species and named Neotermes luykxi. The present study established clearly that, on the basis of both chromosome num- ber and enzyme differences, it is a species distinct from N. jouteli. A difference in chromosome number alone is not sufficient to establish species status. ““Chromosomal races” of the same species may also have different chromo- some numbers as a result of variation in the number of centric fusions (John 1983). But — Fig. 3. Male meiosis in several lower termites. Magnification is approximately the same for all cells, about 1400 =. Bar = 10 um. a, /ncisitermes schwarzi, 9 bivalents and a sex-multiple of 14 chromosomes; b, Neotermes castaneus, 16 bivalents and a sex-multiple of 6 chromosomes; c, /ncisitermes snyderi, 22 bivalents and a sex- univalent, upper left; d, Neotermes mona, 23 bivalents; e, Neotermes luykxi, 21 bivalents and a C-shaped sex- trivalent, at top; f, Neotermes jouteli, 28 bivalents; g, Mastotermes darwiniensis, 49 bivalents; h, Pterotermes occidentis, 38 bivalents and a linear sex-trivalent, just below and to the right of center. 394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LUTTTN Tid Incisitermes schwarzi 1=4=7 | sex chromosome lea | UL MThvudd Neotermes castaneus | KEY acrocentric autosome | metacentric autosome metacentric involved in translocation with sex chromosome 1-4-3 Ss { 2T A livid Incisitermes minor 1=5=1 4A | livia Incisitermes snyderi {III {|| | 1-4-0 TL Pterotermes occidentis 2-8-1 ss. Mastotermes darwiniensis 2-3-0 livucnide [hii Neotermes luykxi Neotermes mona 1=1=1 1-4-0 NO fe Cryptotermes cavifrons 1-1-0 [. Neotermes jouteli 1-0-0 fe HANI Incisitermes milleri 0-0-0 Fig. 4. Schematic representation of ordered karyotype changes in 11 species of lower termites (10 Kaloter- mitidae, | Mastotermitidae), based on the data in Table 4. Arrows are labeled by the following karyotypic changes: P, polyploidization; A, autosomal centric fusion; S, sex-chromosome-autosome centric fusion; T, translocation between autosome and sex-chromosome. The 3-digit formula under each species name indicates for each karyotype (ploidy level)}{haploid number of autosomal metacentricsHhaploid number of sex-chro- mosome metacentrics). This tree was constructed by hand, minimizing genome amplification events and avoiding centric fissions on the assumption that these are much less frequent than centric fusions and translocations in karyotype evolution. This and other parsimonious trees of karyotype evolution (constructed by PAUP) are not congruent with the trees (Figs. 1 and 2) derived from morphological and allozyme characters (see text). in such cases the N.F., the number of major chromosome arms, Is the same for all races. This is not the case here; the haploid N.F. for /uykxi is 25, while that for jouteli is 28. This indicates that the two types are not simply different populations of a single polytypic species related by centric fusions. Moreover, fusions between autosomes and sex chromosomes have occurred in /uykxi but are so far unknown in joutel. These chromosomal differences, along with the enzyme differences described in this VOLUME 92, NUMBER 3 paper (9 out of 17 loci studied) and the slight but definite morphological differences de- scribed by Nickle and Collins (1989), leave little doubt that /uyAxi and jouteli do not share a common gene pool, and are there- fore different species. The phylogenetic re- lationships of the two species, as reflected in the cladogram based on the combined morphological and electrophoretic data (Fig. 2), indicate that the two species are never- theless closely related, and therefore /uyAxi is appropriately included in the genus Neo- termes. The cladogram (Fig. 2) is based on 13 morphological characters and 17 enzyme characters. It might be argued that the rel- atively small number of individuals sam- pled in the electrophoretic studies could bias the cladogram, if numerous enzyme poly- morphisms went undetected, and appar- ently fixed differences between species were in fact simply different allelic forms uncov- ered as a result of chance sampling among a small number of genomes. For several reasons, it is unlikely that many of the species differences represent simply allelic differences in polymorphic populations, and that the phylogenetic tree topology is thereby significantly biased. First, levels of enzyme polymorphism in the lower termites are probably low. In the only kalotermitid species studied extensively to date, Incisitermes schwarzi, Santos and Luykx (1985) found only 4 polymorphic loci out of 23 studied (17%). Limited though the samples of each species in the present study were, there were no other clear cases of in- traspecies polymorphism, indicating that levels of polymorphism in the other species are generally low also. Secondly, when 4 genomes are sampled for each species (the minimum number sampled for each species in this study), the probability that two species will appear to have fixed differences when in fact they both are polymorphic at a given locus, is not very high. It can be calculated, for example, that if two species are both polymorphic at a locus, and alternate alleles each occur at a 395 frequency of .95 in the two different species, then the probability is .66 that, when 4 ge- nomes are sampled from each species, one species will appear to be fixed for one allele and the other species will appear to be fixed for the other allele. This means that under these special circumstances— approaching a state of fixed species differences—about 7 of the truly polymorphic loci will show up as fixed differences in the two species; that is, only about 2 loci out of the 17 loci stud- ied, assuming that /ncisitermes schwarzi 1s fairly representative of the family (see above). If the two alleles are both equally frequent at a given locus in each of two species, the probability is less than 1% that the two species will appear to be fixed for the different alleles. Finally, it has been shown that when a relatively large number of loci are used, and the fraction of loci that are polymorphic is relatively low (as appears to be the case here), samples as small as even single individuals from each species give cladogram topolo- gies that are not any different from those obtained when larger species samples are used (Hillis 1987; see also Gorman and Renzi 1979). Admittedly, larger sample sizes for each of these species would resolve these ques- tions. But the above considerations make it improbable that failure to detect enzyme polymorphisms because of limited sample sizes significantly affected the topology of the cladogram. The cladogram presented in Fig. 2, com- bining both morphological and electropho- retic data, was rooted using Mastotermes darwiniensis as the outgroup. Samples of this species were unfortunately not avail- able at the time of the electrophoretic stud- ies, so the rooting of the tree is based on the morphological data alone. In view of the general agreement (similar consistency in- dices) between the morphological and elec- trophoretic data, however, it seems unlikely that the addition of electrophoretic data from Mastotermes would significantly alter the topology of the tree. This is a point that 396 can be investigated more thoroughly in fu- ture studies. The phylogeny as presented in Fig. 2 is in general agreement with that proposed by Krishna (1961) on the basis of classical mor- phological studies. There are, however, two differences. The first is that Pterotermes oc- cidentis is phylogenetically related to the /n- cisitermes-Cryptotermes branch of the kal- otermitids, not to the Neofermes branch as Krishna supposed. The association of P. oc- cidentis with Incisitermes and Cryptotermes rather than Neotermes is supported by 9 de- rived characters, 7 of which are shared with at least one other species of /ncisitermes, and 3 of which are shared with all the /n- cisitermes species studied. These numbers make it highly improbable that Prerotermes is more closely related to Neotermes than to Incisitermes (see Felsenstein 1985). The second difference with Krishna’s phylogeny is that Neotermes appears to be paraphyletic. Neotermes castaneus (unfor- tunately the type species for the genus) in fact appears in Fig. 2 as a sister group to the other Neotermes species and to /ncisitermes species. The data indicate that some revi- sion of the taxonomy of this branch of the Kalotermitidae is required, but it seems pre- mature to revise it until more extensive studies, including more members of the ge- nus, are carried out. It is interesting that the morphological characters and allozymic characters appear to define phylogenetic groupings at different levels: species are defined more by their unique allozyme characters than by their morphology, while the morphological char- acters tend to define higher taxa (genera). This tendency has also been observed in other groups of animals (e.g. see fig. 4 in Hillis 1987). The observation suggests that allozymic changes frequently accompany speciation, while morphology is more con- served—a tendency that might be expected if single base changes are responsible for allozyme differences, while significant mor- phological differences require more exten- sive genetic repatterning. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Some investigators (e.g. Miyamoto 1983) have treated the karyotype as a single char- acter with multiple states (arising from cen- tric fusions, pericentric inversions, etc.), and have combined karylogical data with the morphological and electrophoretic data to generate phylogenetic trees. Treating chro- mosomal data in this way, however, as- sumes that chromosomal changes occur at approximately the same rates and play ap- proximately the same role in speciation as do the gene mutations that lead to changes in morphology and allozymes. Rates of structural changes in chromosomes, how- ever, are several orders of magnitude higher than rates of gene mutations (Jacobs 1981, Van Dyke et al. 1983, Hook et al. 1984). The role of chromosomal changes in spe- ciation is still a controversial subject (e.g. Sites and Moritz 1987), and it therefore seems better to treat chromosomal changes separately, and not combine them with morphological and allozyme data. It is theoretically possible to construct a separate phylogeny based on the chromo- somes of the ten kalotermitid species stud- ied here and of Mastotermes darwiniensis (Fig. 3). As outlined in the following para- graphs, the changes that have apparently led to the karyotypic differences between these species are (i) an increase in number of ma- jor chromosome arms, possibly by a process akin to polyploidization; (ii) centric fusion between autosomal acrocentrics; (i11) centric fusion between sex chromosomes and auto- somes; and (iv) whole-arm translocation be- tween autosomal and sex-chromosomal metacentrics. Evolutionary polyploidization in animals is very rare (White 1973), and it is uncertain whether this process has really occurred in these termites. But the numbers of major chromosome arms (N.F.) in the species studied here seem to fall into distinct cat- egories: Incisitermes milleri has N.F. = 14, most of the other Jncisitermes and Neo- termes species have N.F. ranging from 25 to 28, and Pterotermes occidentis and Mas- totermes darwiniensis have N.F. = 49 and VOLUME 92, NUMBER 3 52, respectively. This is almost a doubling series and suggests, if not polyploidization, at least distinct evolutionary episodes of amplification. The karyotypes of many of these termite species consist of mixtures of acrocentric and metacentric chromosomes. As would be expected if centric fusions or fissions were important in the karyotypic changes exhib- ited by these termites, metacentric chro- mosomes are in general about twice the size of acrocentric chromosomes. And even if the absolute chromosome numbers differ, species with similar N.F. have similar DNA contents. The haploid DNA content of Neo- termes jouteli is 1.30 pg, almost identical with that of [ncisitermes schwarzi, 1.35 pg (Luykx, unpublished data). While the hap- loid chromosomes numbers of these two species are quite different, N. jouteli with n = 28 (all acrocentrics) and J. schwarzi with n= 16 (5 acrocentrics and 11 metacentrics), the N.F. for these two species is almost the same (28 and 27, respectively). Differences in chromosome number are probably due primarily to centric fusions rather than cen- tric fissions, since fusions are much more common than fissions among orthopteroid insects in general (Hewitt 1979), and be- cause there is little doubt that fusions are responsible for the origin of the sex-triva- lents in Neotermes luykxi, Incisitermes mi- nor, and Pterotermes occidentis, as well as in other kalotermitid species (Luykx and Syren 1979). The formation of the multivalent rings seen in male meiosis in Neotermes cas- taneus and Incisitermes schwarzi can be accounted for by a series of whole-arm translocations between autosomal and sex- chromosomal metacentrics. Starting with a metacentric pair of sex chromosomes (which may themselves have arisen by centric fu- sions), each successive translocation ofa sex chromosome with an autosome would in- crease the size of the sex-multivalent by two chromosomes (Syren and Luykx 1981). Thus, one translocation would give a ring of 4 chromosomes, an additional translo- 397 cation would give a ring of 6, and so on. Evidently, two such translocations have oc- curred in N. castaneus, and a total of 6 such translocations have occurred in the /. schwarzi population used in these studies (see Luykx and Syren 1979 and Luykx 1987 for other translocation variants in this species). It seems likely that a karyotype consisting entirely of acrocentric chromosomes was the ancestral condition. This karyotype is the most common one among the kalotermitids (Luykx and Syren 1979, Luykx 1990). The chromosomal variations seen in these ter- mites, then, in accord with the consider- ations discussed above, can be understood as arising from a limited number of pro- cesses acting on a primitive all-acrocentric karyotype: the amplification of the number of chromosome arms (perhaps by poly- ploidization-like events), the fusion of cen- tromeres (between autosomes and sex chro- mosomes as well as among autosomes), and the translocation of whole arms between au- tosomal and sex-chromosomal metacen- trics. A phylogeny of chromosome changes, based on the data in Table 4, can be con- structed on the above principles. A simple phylogeny is shown in Fig. 3. It is obvious that this and other parsimonious trees of karyotype evolution (constructed by PAUP) bear little relation to the cladogram derived from morphological and allozyme charac- ters (Fig. 2). The most reasonable expla- nation for the discrepancy is that chromo- some arrangements in these insects are too labile to be good indicators of phylogeny. In other words, chromosome changes may arise and be fixed in populations more rap- idly than the speciation events themselves, a view supported by the extensive karyotype variation also observed within these species (Syren and Luykx 1981, Luykx 1983, Luykx 1987). Similar karyotype modifications may therefore occur independently on separate branches of the “true” phylogeny (here as- sumed to be approximated by Fig. 2). Thus, 398 since there is no obvious mechanism for halving genome size in a single step, discrete genome amplification events (polyploidiza- tion?) would have to have occurred sepa- rately on all branches except the one leading to Incisitermes milleri. Autosomal centric fusions appear to have occurred to varying extents on different branches, giving the same number of autosomal metacentrics in Incisitermes snyderi and Neotermes mona, for example, quite independently. Similar- ly, frequent centric fusions and repeated translocations between metacentric sex chromosomes and metacentric autosomes, to give multivalent rings in male meiosis, appear to have occurred independently on separate branches leading to Neotermes cas- taneus and Incisitermes schwarzi. It seems likely that these processes of karyotype modification— approximate dou- bling of chromosomal material, centric fu- sion, and whole-arm translocations—are all widespread and common enough that vir- tually every lineage is subject to them. The different karyotypes that are currently ob- served in these various species are therefore probably simply the outcome of variations in the frequency with which these chro- mosome changes occur within lineages, and in population factors that affect the likeli- hood that the changes will be fixed. ACKNOWLEDGMENTS For sending us the colonies of /ncisi- termes minor and Pterotermes occidentis, we are grateful to Mike Haverty (Pacific South- west Forest and Range Experiment Station, U.S.D.A.) and to Tim Myles (University of Arizona), respectively. We also thank Niilo Virkki (University of Puerto Rico) for his assistance in collecting Neotermes mona, and J. A. L. Watson (C.S.I.R.O., Canberra, Australia) for providing the colonies from which the Mastotermes darwiniensis chro- mosome preparations were made. We thank Drs. T. E. Henry and E. E. Grissell (System- atic Entomology Laboratory, National Mu- seum of Natural History, Washington, D.C.) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for reviewing the manuscript. This research was supported by National Science Foun- dation research grant no. BSR-8119692. LITERATURE CITED Bedo, D. G. 1987. Undifferentiated sex chromo- somes in Mastotermes darwiniensis Froggatt (Isoptera; Mastotermitidae) and the evolution of eusociality in termites. Genome 29: 76-79. Cranston, P.S.and C.J. Humphries. 1988. Cladistics and computers: A chironomid conundrum? Cla- distics 4: 72-92. Felsenstein, J. 1985. Confidence limits on phyloge- nies: an approach using the bootstrap. Evolution 39: 783-791. Gorman, G. C. and J. Renzi, Jr. 1979. Genetic dis- tance and heterozygosity estimates in electropho- retic studies: effects of sample size. Copeia 1979: 242-249, Grassé, P.-P. 1986. Termitologia: Anatomie, Phy- siologie, Systematique des Termites. t. III, pp. 446—- 459. Masson, Paris. Harris, H. and D. A. Hopkinson. 1976. Handbook of Enzyme Electrophoresis in Human Genetics. North Holland Publishing Co., Oxford. Hewitt, G. M. 1979. Orthoptera, pp. 1-170. /n John, B. et al., eds., Animal Cytogenetics, vol. 3, Insecta 1. Gebruder Borntraeger, Berlin, Stuttgart. Hillis, D. M. 1987. Molecular versus morphological approaches to systematics. Ann. Rev. Ecol. Syst. 18: 23-42. Hook, E. B., D. M. Schreinemachers, A. M. Willey, and P. K. Cross. 1984. Inherited structural cy- togenetic abnormalities detected incidentally in fe- tuses diagnosed prenatally: Frequency, parental- age associations, sex-ratio trend, and comparisons with rates of mutants. Am. J. Hum. Genet. 36: 422-443. Jacobs, P. A. 1981. Mutation rates of structural chro- mosome rearrangements in man. Am. J. Hum. Genet. 33: 44-54. John, B. 1983. The role of chromosome change in the evolution of orthopteroid insects, pp. 1-110. In Sharma, A. K. and A. Sharma, eds., Chro- mosomes in Evolution of Eukaryotic Groups. CRC Press, Inc., Boca Raton, Florida. Kluge, A. G. and J. S. Farris. 1969. Quantitative phyletics and the evolution of anurans. Syst. Zool. 18: 1-32. Krishna, K. 1961. A generic revision and phyloge- netic study of the family Kalotermitidae (Isop- tera). Bull. Am. Mus. Nat. Hist. 122: 303-408. Luykx, P. 1983. XO:XX sex chromosomes and Rob- ertsonian variation in the autosomes of the wood- roach Cryptocercus punctulatus (Dictyoptera: VOLUME 92, NUMBER 3 Blattaria: Cryptocercidae). Ann. Entomol. Soc. Amer. 76: 518-522. 1987. Variation in sex-linked interchange heterozygosity in the termite /ncisitermes schwarzi Banks (Insecta: Isoptera) on the island of Jamaica. Genome 29: 319-325. 1990. A cytogenetic survey of twenty-five species of lower termites from Australia. Genome, 33: 80-88. Luykx, P. and R. M. Syren. 1979. The cytogenetics of Incisitermes schwarzi and other Florida ter- mites. Sociobiology 4: 191-209. 1981. Experimental hybridization between chromosomal races in Kalotermes approximatus, a termite with extensive sex-linked translocation heterozygosity. 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Holozygosity for sex- linked genes in males of the termite /ncisitermes schwarzi. Biochem. Genet. 23: 729-740. Sites, J. M., Jr. and C. Moritz. 1987. Chromosomal evolution and speciation revisited. Syst. Zool. 36: 153-174. Syren, R. M. and P. Luykx. 1977. Permanent seg- mental interchange in the termite /ncisitermes schwarzi. Nature 266: 167-168. 1981. Geographic variation of sex-linked translocation heterozygosity in the termite Kalo- termes approximatus Snyder (Insecta: Isoptera). Chromosoma 82: 65-88. Van Dyke, D. L., L. Weiss, J. R. Roberson, and V. R. Babu. 1983. The frequency and mutation rate of balanced autosomal rearrangements in man esti- mated from prenatal genetic studies for advanced maternal age. Am. J. Hum. Genet. 35: 301-308. White, M. J. D. 1973. Animal Cytology and Evolu- tion. Third Edition, pp. 454-463. Cambridge Uni- versity Press, Cambridge. Yang, S. Y. 1971. Appendix, pp. 84-90. Jn Selander, R. K., etal. 1971. Biochemical polymorphism and systematics in the genus Peromyscus. |. Variation in the old-field mouse (Peromyscus polionotus). Studies in Genetics VI. University of Texas Pub- lication No. 7103. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 400-406 NEW SPECIES OF BUPRESTIDAE (COLEOPTERA) FROM THE DOMINICAN REPUBLIC HENRY A. HESPENHEIDE Department of Biology, University of California, Los Angeles, California 90024. Abstract.—Three species of Buprestidae are newly described, illustrated, and distin- guished from other Antillean and closely related species in their respective genera: Sam- bomorpha clarki, Agrilus klapperichi, and Neotrachys bilyi. Additional records are given for Acmaeodera cruenta (Olivier), Chrysobothris haitiensis Fisher, Taphrocerus haitiensis Fisher, Leiopleura darlingtoni Fisher and L. gibbipennis (Fisher). A list of Hispaniolan Buprestidae is given in a table. Key Words: Recent collections, primarily in the Do- minican Republic, show the buprestid fau- na of Hispaniola to be relatively poorly known, especially for smaller species. This paper describes three new species, reports recent collections of previously described species, and gives a checklist of forms cur- rently known to occur on the island. Ad- ditional specimens in the Mastogeninae have been seen but remain undetermined. The following species appear not to be described: Sambomorpha clarki, NEw SPECIES (Figs. 1, 2) Description. — Holotype, probably fe- male: Length 6.4 mm, width 2.1 mm; dor- soventrally somewhat flattened, strongly so above; moderately shining; head, pronotum and scutellum purplish red, darker on pronotum postero-medially, more aeneous on anterior angles, elytra dark reddish pur- ple, more reddish at base and on margins; pronotum ornamented with patches of elongate white setae in anterior and poste- rior angles, along basal '2 of midline and in spots on anterior margin ' the distance to margin, elytra ornamented with white setae Agrilus, biogeography, distribution, Hispaniola, Neotrachys, Sambomorpha in a broken line of spots on either side of suture and along lateral margin on basal *4 and in two irregular transverse fascia at api- cal ¥4 and near apex; otherwise with long, thin, dark setae; beneath purplish with gold- en reflections. Head with front shallowly depressed along midline, with indistinct convexities on either side of midline at middle of eye; surface nearly smooth on dorsal '2, especially me- dially, weakly rugose on ventral 1; sparsely clothed with long white setae dorsally, with shorter, denser setae along midline on ven- tral '2; epistoma narrow between antennae, faintly carinate along midline to base, an- gulately emarginate along base; antennae reaching only anterior '3 of pronotal length, serrate from the fifth segment, outer joints rounded-triangular; eyes small, broadly oval, wider ventrally. Pronotum 1 ¥, wider than long, widest just anterior to base; sides slightly sinuate then narrowed to apical angles, posterior angles narrowly rounded-obtuse; when viewed from side lateral margin sinuate, strongly produced, especially for anterior 2; anterior margin broadly rounded; base with narrow, smooth margin, angulate-emarginate at VOLUME 92, NUMBER 3 5 401 Figs. 1-6. Figs. 1, 2. Sambomorpha clarki, n. sp.; line equals 1 mm. 1, dorsal view. 2, lateral view. Figs. 3, 4. Agrilus klapperichi, n. sp.; line equals 1 mm. 3, dorsal view. 4, lateral view. Figs. 5, 6. Neotrachys bilyi, n. sp. 5, dorsal view, holotype—shading indicates area of dark violet; line equals 1 mm. 6, genitalia, allotype; line equals 0.5 mm. middle of each elytron, median lobe mod- erately produced and truncate in front of scutellum; disk broadly convex anteriorly, transversely depressed on each side at base, strongly so interior to prehumeral carinae to anterior angles; prehumeral carinae very strong, parallel to lateral margin, extending from basal margin nearly to anterior margin of pronotum; narrowly depressed between carina and margin; surface somewhat finely imbricate-punctate, intervals smooth. Scu- tellum nearly flat, smooth. Elytra slightly wider than pronotum at base; sides shallowly constricted then slight- ly expanded to middle, then gradually nar- rowed to broadly, separately rounded and denticulate tips; sides of abdomen slightly exposed above; disk strongly flattened, with rather strong transverse basal depressions; surface finely imbricate-punctate. Prosternum confluent-punctate, uni- formly, sparsely clothed with long, white, recumbent setae, deeply transversely de- pressed at base of prosternal lobe, which 1s declivous and broadly, arcuately emargin- ate; sides of prosternal process converging gradually to behind coxal cavities, then acutely angulate. Abdomen convex ven- 402 trally, marked with transverse crenulate lines; sparsely, uniformly clothed with fine, white, recumbent setae, and small spots of denser setae on anterio-lateral portions of segments 2-4; suture between first and sec- ond segments indicated only at sides; last segment broadly, shallowly emarginate at apex. Tibiae nearly straight, armed with a small, thin spine on inner margin at apices; anterior tibiae denticulate on outer margin at base. Tarsal segments equal; tarsal claws similar on all feet, cleft at base, inner tooth straight, much shorter than outer one. Holotype.—Rep. Dominicana, Bani, 30.1X.1985, W. E. Clark, to be deposited in USNM. The genus Sambomorpha was described by Obenberger (1924) for a single species from southeastern Brasil. All members of the genus most obviously share the dorso- ventrally depressed body form, strong and sinuate prehumeral carinae on the prono- tum, and similar diffuse patterns of pubes- cence on the elytra. I have recently (Hes- penheide 1979) transferred to the genus a second Brasilian species described original- ly as an Agrilus, and have seen undescribed material from Mexico. Sambomorpha clar- ki can be distinguished from both Brasilian species by the absence of a posthumeral ca- rina on the elytra, color, and somewhat larg- er size; and from the Mexican forms by its color and more definite pattern of pubes- cence on the elytra. Sambomorpha shares its rather widely disjunct and unusual dis- tribution in the New World (Hispaniola, Mexico, southeastern Brasil), presumably a relictual one, with the genus 7etragonos- chema. Cyphothorax shows a similar dis- junct range between Mexico and southeast- ern Brasil. The type appears to be a female, although it was not dissected. Agrilus klapperichi, New Species (Figs. 3, 4) Description.—Holotype female: Length 5.9 mm, width 1.4 mm; moderately elon- gate and robust, strongly flattened above; PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON weakly shining, black, head, sides of prono- tum, and ventral surfaces with bronzy re- flections; elytra ornamented with five pairs of small pubescent spots. Head with the front wide, a small trans- verse depression below middle and a nar- row one along midline above and below middle, lateral margins nearly straight; sur- face moderately rugose, clothed with mod- erately long setae which radiate from the central depression, becoming very short on occiput; epistoma moderately wide between the antennae, shallowly emarginate along base; antennae serrate from the fourth seg- ment, outer joints triangular; eyes about equally rounded above and below. Pronotum slightly wider than long, sub- equal at base and apex, widest at apical 4; sides subparallel at base, shallowly convex to apical angles; posterior angles nearly per- pendicular; when viewed from side margin- al and submarginal carinae narrowly sepa- rated for their entire length, more broadly so at apex, weakly sinuate; anterior margin sinuate, median lobe moderately produced and broadly rounded; base angularly emar- ginate at middle of each elytron, median lobe slightly produced and weakly emargin- ate in front of scutellum; disk shallowly con- vex, depressed on each side at base, strongly so at anterior angles, two shallow transverse medial depressions and depressed along midline for basal '2; prehumeral carinae subparallel with and narrowly separated from lateral margin, extending for basal '2 of pronotum; surface rather finely, trans- versely rugose, impunctate, setae short and inconspicuous on disc, longer and more conspicuous just interior to prehumeral ca- rinae and lateral margins. Scutellum strong- ly transversely carinate, surface shagreened. Elytra subequal to pronotum at base, wid- est at humeri; sides very shallowly con- stricted and then slightly widened to apical ¥s, then gradually narrowed to tips, which are separately rounded and finely toothed; sides ofabdomen exposed dorsally; disk with moderate basal depressions and shallowly VOLUME 92, NUMBER 3 depressed along sutural margins, except just behind scutellum; surface rather finely im- bricate-punctate, uniformly clothed with short setae, each elytron ornamented with five small round spots of denser setae: in basal depression, along suture at basal ', 73, and near apices, and near sides just be- yond middle. Posterior coxae nearly straight on poste- rior margin. Abdomen with suture between first and second segments distinctly indi- cated at sides; anterior vertical portions of segments 2, 3, and 4 ornamented with oval spots of longer setae; dorsal portions of seg- ments 1, 3, and 4 densely setose. Tibiae unarmed. Posterior tarsi shorter than tibiae, first joint subequal to following three united. Tarsal claws similar on all feet, cleft with inner tooth subparallel to and shorter than outer one. Holotype.—Rep. Dominicana, Bani, 30.1X.1972, J. & S. Klapperich (Basel). Agrilus klapperichi is quite different from the other, much larger species of Agrilus (A. dominicanus Thomson, related to 4. macer LeConte) known from Hispaniola (Fisher 1925). A. klapperichi appears to be related to the group of species that in the United States includes A. obsoletoguttatus Gory, A. exsapindi Vogt, A. limpiae Knull, A. scitulus Horn and A. faeniatus Chevrolat. Among Central American species, A. klapperichi appears most closely related to 4. simulans Waterhouse, 4. femoralis Waterhouse and A. antennatus Waterhouse. All of these species are characterized by similar patterns of pubescence on the elytra and ventral body surfaces and by sexual dimorphism in the antennae, in which males tend to have an- tennae which are relatively more elongate and occasionally modified otherwise. The characters which are unique to A. klapperi- chi compared to the North and Central American species mentioned above are (a) the prehumeral carina subparallel with the marginal carina, (b) the separation of the marginal and submarginal carinae of the pronotum for their entire length, and (c) the 403 transverse medial depression on the front. Because the specimen is mounted venter down on a card, characters on the ventral surface could not be described. Neotrachys bilyit, NEW SPECIES (Figs. 5, 6) Holotype female.—Length 2.75 mm, width 1.3 mm; oval, moderately convex, rounded-quadrate in front, attenuate pos- teriorly; inconspicuously short pubescent; moderately shining and strongly sha- greened; head, pronotum and humeri green- ish blue, elytra blue, becoming dark violet blue on inner apical '2 along suture and be- yond basal 3; beneath black. Head depressed on front along midline between eyes and with transverse depres- sion between epistomal pores; epistomal pores moderate in size; surface moderately densely ocellate-punctate, punctures less dense and finer between eyes; intervals sha- greened; epistoma with anterior margin very shallowly angulate-emarginate; subocular pores large; antennae with 6th segment slightly produced below, conspicuously ser- rate only from 7th segment. Pronotum moderately convex, about 2', times as wide as long, distinctly narrower in front than behind, widest at base: sides arcuately rounded from base, more strongly so at anterior angles, narrowly margined; anterior angles obtuse; posterior angles per- pendicular; anterior margin nearly trans- verse, with obsolete lobe at middle; poste- rior margin transversely feebly sinuate with strong subtruncate median lobe before scu- tellum; surface shallowly depressed along basal margin and more strongly so at an- terior angles; rather densely ocellate-punc- tate along margins, more densely so in pos- terior angles, more finely, simply and less densely so on disk; intervals strongly sha- greened. Scutellum small. Elytra moderately convex, wider than pronotum at base; humeral angles broadly rounded-quadrate; sides nearly parallel to middle, then shallowly attenuate to con- 404 jointly, rather narrowly rounded tips; each elytron with small, shallow depression at base interior to humerus, a deep, narrow, elongate one behind humerus along lateral margin, and an elongate, shallow one along suture for posterior 74; surface regularly punctate, intervals strongly shagreened. Prosternum sparsely, faintly ocellate- punctate, intervals smooth, greenish-blue;: anterior margin shallowly arcuate; proster- nal process moderately broad, broadened slightly behind coxal cavities, and broadly, rounded-quadrate at apex. Hind coxae de- pressed along their length, broadly, shallow- ly biemarginate with broadly, obtusely rounded projection dorsal to attachment of hind legs. Abdomen beneath with dense, large, shallow, fine ocellate punctures, which are elongate and denser at base and sides of first segment, sparser and more rounded on segments 2-5; intervals densely, finely, ob- soletely reticulate-striolate. Allotype male: Similar to female, except bright golden-green above, with darker golden coppery reflections on top of head, disk of pronotum, and central portions of each elytron. Holotype.—Rep. Dominicana, Cazabita, 1250 m, 15.VIII.1972, J. & S. Klapperich (Basel). Allotype.— Dominican Rep., La Vega, 15 km E of El Rio, 26.V.1978, C. W. & L. B. O’Brien & Marshall (USNM). Neotrachys bilyi differs strikingly in ap- pearance from most other Antillean Neo- trachys treated in the revision of the genus (Hespenheide 1980). Although propor- tioned like N. dominicanus, N. fennahi and N. guadeloupensis, it is conspicuously sha- greened (or granular reticulate) like N. hoff- mani. The color of the holotype, that 1s, greenish-blue anteriorly shading to deep vi- olet-blue on the inner posterior portions of the elytra, is unique among the Antillean species. The male is less strikingly colored and is similar to some individuals of Neo- trachys from Puerto Rico. It is not clear PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON whether these latter specimens represent N. hoffmani, in which case that species is more variable than originally treated, or whether they belong to a second, undescribed Puerto Rican species; additional specimens are needed to determine which is the case. N. bilyi can be separated from these question- able forms as follows: N. bilyi N. (hoffmani?) dorsal setae minute conspicuous subocular pores — large inconspicuous pronotum broader, narrower, more shorter elongate elytral intervals strongly sha- mostly smooth greened posterior margin, biemarginate nearly straight hind coxae distribution Hispaniola Puerto Rico N. bilyiis readily distinguished from typical N. hoffmani, which are strongly depressed along the lateral margins of the elytra near the middle and dark olive green in color. The characteristics in the table separate N. bilyi from all Central American species as well (Hespenheide 1982). The beetle is named in honor of Svato- pluk Bily, who allowed me to see and de- scribe this and the preceding species. The following species are represented by additional distributional data: Acmaeodera cruenta (Olivier) Specimens examined.—Dominican Re- public: 6.5 mi W Azua, 13.VI.1968, H. A. Hespenheide, on Prosopis (RLWC); Pera- via, 6 km W Bani, 4, 15.1X.1983, W. E. Clark (AUBU, CHAH). Chrysobothris haitiensis Fisher, 1930:7 Specimens examined: Dominican Re- public: Boca Chica, 10 m, 4.XI.1972, Klap- perich (Basel); Bani, 31.III.1973, Klappe- rich (Basel); Barahona, 6 km NW Fundacion, 1.1X.1983, W. E. Clark (AUBU); Peravia, 13 km NW Bani, 6.VIII.1979, C. W. O’Brien (CHAH). VOLUME 92, NUMBER 3 Table 1. Checklist of species of Buprestidae re- corded from Hispaniola. Literature citations are given only for those species described or reported since Fish- er’s (1925) monograph of Antillean Buprestidae. Polycesta fisheri Obenberger, 1936:105 insulana Fisher, 1930:125 regularis Waterhouse?—type locality uncertain (Fisher 1925) porcata (Fabricius) Paratynaaris antillarum Fisher, 1939:156 Acmaeodera cruenta (Olivier) Hilarotes nitidicollis (Laporte & Gory) mannerheimi (Mannerheim) Psiloptera aurata (Saunders) aurata var. domingoensis Fisher, 1930:126 aurifer (Olivier) Dicerca divaricata (Say)—introduced? (Fisher 1925) Cinyra albonotata (Laporte & Gory) Buprestis hispaniolae Fisher, 1939:157 striata Fabricius maculativentris Say Mixochlorus elegans Fisher Peronaemis insulicola Fisher, 1939:159 Melanophila acuminata (Deg.)— Blackwelder 1944: 313 Tetragonoschema quadrata (Buquet) Chrysobothris tranquebarica (Gmelin) dentipes (Germar) haitiensis Fisher, 1930:7 megacephala Laporte & Gory chlorosticta Thomson parvofoveata Fisher hispaniolae Fisher Actenodes bellula Mannerheim embrik-strandi Obenberger, 1936:138 nobilis (Linnaeus)— Fisher 1930:127 Sambomorpha clarki, n. sp.—this study Agrilus dominicanus Thomson klapperichi n. sp.—this study Taphrocerus haitiensis Fisher, 1949:348 Leiopleura darlingtoni Fisher, 1939:162 gibbipennis (Fisher), 1939:160 Neotrachys bilyi n. sp.—this study Micrasta hispaniolae Fisher, 1939:166 monticola Fisher, 1939:165 Taphrocerus haitiensis Fisher, 1949:348 Specimens examined: Rep. Dominicana: Boca Chica, 10 m, 5.X.1970, 29.XI.1970, Klapperich (Basel); Bani, 65 m, 30.1V.1974, Klapperich (Basel); La Altag., 31 km N Hi- guey, 1. VIII.1979, G. B. Marshall (CHAH). 405 Leiopleura darlingtoni Fisher, 1939:162 Specimen examined: Dominican Repub- lic: La Culata (La Vega), 1500 m, 18.11.1978, H. and A. Howden (CHAH). Leiopleura gibbipennis (Fisher), 1939:160 Specimens examined: Dominican Re- public: Boca Chica, 10 m, 5.X.1970, 29.X1.1970, Klapperich (Basel). With the addition of the three species de- scribed above, the total fauna of Bupresti- dae reported from Hispaniola stands at 39, of which one is probably introduced and a second is questionable. Table | presents the list of species reported to date. ACKNOWLEDGMENTS The author (CHAH) is indebted to Sva- topluk Bily of the Natural History Museum of the National Museum, Prague, Czecho- slovakia (NMPC): Ronald D. Cave and Wayne E. Clark of Auburn University (AUBU); Henry F. Howden (HAHC) of Carleton University, Ottawa, Canada; John Kingsolver of the U.S. National Museum (USNM); Charles W. and Lois B. O’Brien of Florida A&M University; and Gayle H. Nelson (GHNC) for loaning specimens. Gayle H. Nelson also determined Chryso- bothris haitiensis Fisher and commented on a draft of the manuscript. Financial assis- tance was provided in part by the UCLA Academic Senate. Margaret Kowalczyk pre- pared the final illustrations. LITERATURE CITED Blackwelder, R. E. 1944. Checklist of the coleopter- uous insects of Mexico, Central America, the West Indies, and South America, Part 2. U.S. Natl. Mus. Bull. 185: 187-341. Fisher, W. S. 1925. A revision of the West Indian Coleoptera of the family Buprestidae. Proc. U.S. Natl. Mus. 65(9): 1-207. 1930. New West Indian Buprestidae (Co- leoptera). Proc. Entomol. Soc. Washington. 32: 125-129. . 1930. New West Indian Buprestidae and Cer- ambycidae (Coleoptera). Can. Entomol. 62: 7-11. 406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1939. New West Indian buprestid beetles. . 1982. A revision of Central American species Psyche 46: 156-166. of Neotrachys (Buprestidae). Coleopts. Bull. 36: 1949. New buprestid beetles from Mexico, 328-349. Central and South America, and the West Indies. Obenberger, J. 1924. Deuxieme serie de nouveaux Proc. U.S. Natl. Mus. 99: 327-351. genres de buprestides. Sbornik Entomol. Odd. Nar. Hespenheide, H. A. 1979. Nomenclatural notes on Mus. Praze 2: 7-44. the Agrilinae (Buprestidae): IV. Coleopts. Bull. 33: . 1936. Eine Festarbeit zum sechzigjarigen Ju- 105-120. bilaum meines Freundes Univ.-Prof. Dr. Embrik —. 1980. A revision of Antillean Neotrachys Strand. Fest.-schr. 60. Geburtstag E. Strand. 1: 97- (Coleoptera, Buprestidae). J. Kansas Entomol. Soc. 145. 53: 815-824. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 407-410 FIRST NORTH AMERICAN RECORDS FOR HARMONIA QUADRIPUNCTATA (PONTOPIDDIAN) (COLEOPTERA: COCCINELLIDAE); A LADY BEETLE NATIVE TO THE PALAEARCTIC NATALIA J. VANDENBERG 705 Carmel Ave., Albany, California 94706. Abstract.— Harmonia quadripunctata (Pontopiddian), an Old World member of the tribe Coccinellini, is newly reported from three localities in the north eastern United States: Paterson and Westfield, New Jersey, and Mt. Kisco, New York. The North Amer- ican specimens appear to have been derived from a single founding population which was established as early as 1924 and probably as the result of an adventive introduction. Key characteristics are given which will distinguish this species from the rest of the native and introduced North American Coccinellini. Habitat and prey preferences are briefly dis- cussed. Key Words: Harmonia quadripunctata (Pontopiddi- an) is an Old World lady beetle of the tribe Coccinellini (sensu Sasaji 1971), which has become established in New Jersey and New York without any record of a deliberate in- troduction (Gordon 1985). A total of 8 spec- imens with North American collection lo- calities have been recovered from 3 museums and one private collection. These specimens represent a minimum of 3 sep- arate collection events (see map, Fig. 1) spanning a period of 54 and a half years and covering a linear distance of approximately 50 miles (=90 km). Although this is the first literature report on the occurrence of H. quadripunctata in North America, the un- usually detailed specimen labels indicate that it has drawn the attention of earlier observ- ers, some of whom were even apprised of its alien status. The diagnosis below will serve to distinguish the North American populations of H. quadripunctata from the rest of the introduced and native Coccinel- Coleoptera, Coccinellidae, Harmonia, aphidophagous, forest lini. Iablokoff-Khnzorian (1982) should be consulted for a synonymical bibliography. Harmonia quadripunctata (Pontopiddian) Fig. 1 Diagnosis of North American population: Form ovo-elliptic, weakly convex, 5.0 to 8.0 mm in length. Ground color of dorsal sur- faces pale orangy brown; pronotum of fully maculate individuals with eleven puncti- form black spots, one or two pairs some- times faint or absent; elytron immaculate or with a pair of elongate black marks at lateral margin on each side of mid-line. Tib- ial spurs lacking. Postcoxal line of abdomen curved posterolaterally, closely approach- ing or joining posterior margin of segment; oblique dividing line present. Harmonia quadripunctata will key to the genus Mulsantina Weise in Gordon (1985), however it can be readily distinguished from North American members of that genus by the larger body size and the presence of an 408 PENNSYL VANIA Fig. 1. Recorded North American collecting sites for Harmonia quadripunctata (ocellate dot); collection dates are given in parentheses. Habitus drawing of adult showing typical maculation of the North American population. oblique dividing line in the postcoxal region of the abdomen. Although the genus Har- monia Mulsant is included in Gordon’s key, the characters used are diagnostic only of the introduced species H. dimidiata (F.), which was the sole species known to occur in North America at the time. The distinc- tive dorsal color pattern of the North Amer- ican population of H. quadripunctata is the best way to separate it from other members of its tribe. Members of the genus Neohar- monia Crotch, which also lack tibial spurs, possess fewer and larger pronotal maculae than H. quadripunctata. Within its native range, H. qguadripunc- tata exhibits extreme elytral color pattern polymorphism from nearly solid black, to various combinations and confluences of black spots against a pale background (Ma- der 1926-37, Iablokoff-Khnzorian 1982). Melanic individuals are rare in the southern part of the European continent, but a range of color forms can usually be found together at a given locality. This within population variation makes it impossible to speculate on the source area of the North American founding population even though they con- sistently fall at the extreme pale end of the color form spectrum. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON I first became aware of the presence of H. quadripunctata in the United States when my colleague Stuart McKamey invited me to have a look at some coccinellids he had collected ten years earlier. I was discon- certed to find that the largest coccinellid in the box was completely unfamiliar to me, and could not be identified using any avail- able keys to the North American fauna. The single specimen, which constitutes our most recent collection record of this exotic species, has a type-set label with the following data, “Mt. Kisco, NY Westchester Co. July 1979 Stuart McKamey Coll.” and a neatly penned post script ‘only specimen collected, not very common.” I soon began to discover other North American specimens of H. quadripunctata which had previously escaped my notice. In the American Museum of Natural History, New York, a pair of the beetles were nestled inconspicuously in a unit tray of Olla y-ni- grum Mulsant. The specimen labels contain the following data, ‘““Westfield, N.J. Union Co., July 8, 1955 G.R. Ferguson [type-writ- ten, photographically reproduced?]/11192 C.A.F.°59 [hand-printed].”’ One of the spec- imens has two additional hand-printed la- bels sandwiched between the others. The white upper label reads “A very unusual immac. var | 1192” and the lower blue label adds “I ought to swipe.” These two speci- mens were collected twenty four years prior to the example captured by McKamey. Some of the earliest collected specimens of North American H. quadripunctata were retrieved from the drawer of miscellaneous coccinellids in the Essig Museum, Univer- sity of California, Berkeley. The find con- sists of a pair of card mounted beetles top- ping a stack of over-sized labels badly yellowed with age. The upper three labels bear the following information in a fine gray script ‘Paterson New Jersey Feb 27 1924 FM Schott/Bulaea lichatschovi Hummel/A Newcomer to United States found here by writer Native to Mediterranean region.” A fourth label on a piece of torn and folded VOLUME 92, NUMBER 3 paper appended to the bottom of the stack, offers the following correction in a robust black script ““Harmonia quadripunctata Pontop. (North Europe).”’ The latter species identification has since been confirmed by comparing all of the U.S. specimens with a European series of H. quadripunctata from the U.S. National Museum of Natural His- tory. Type specimens of H. quadripunctata have not been located (Gordon 1987b, Ia- blokoff-Khnzorian 1982), although workers seem to be in agreement on the identity of this common palaearctic species. The ento- mology collection of the California Acade- my of Sciences has three additional speci- mens of H. qguadripunctata possibly derived from the series collected by Schott. These are labeled as follows: one specimen, “‘Pa- terson N.J. 11.27.24 [no collector given]/ Nunenmacher collection” and two speci- mens on a single mount, “‘Bulaea lichchat- chovi Paterson NJ Feb 27 [no year or collector given]/R. HOPPING COLLEC- TION.” Someone had correctly filed these examples among the European H. quadri- punctata but had either not published or simply failed to notice the unusual collec- tion locality. In addition to the eight specimens men- tioned above, a search through the ento- mology collection of the Los Angeles Coun- ty Museum revealed a single example of /7. quadripunctata placed among the undeter- mined North American Coccinellidae. This individual showed the same distinctive col- or pattern as specimens collected in New York and New Jersey, but the glossy spec- imen label had faded to a uniform blue- gray. The source of the specimen must therefore be considered unsubstantiated. Although ten years have passed since the last known collection of H. quadripunctata in North America, I am inclined to believe that it is still present. The three known col- lection dates, from earliest to most recent, were separated by intervals of about 30.5 and 24 years respectively during which no additional specimens were found. The re- 409 cent ten year interval is therefore a com- paratively short one, and since my search was by no means exhaustive, other speci- mens may well come to light. Several factors could have contributed to the low collection rate for this species as compared with oth- ers. In Europe, H. quadripunctata is most common in forested regions where fir, pine, poplar and chestnut grow (lablokoff- Khnzorian 1982, Klausnitzer and Klaus- nitzer 1986). It is therefore less likely that the species would turn up in an agricultural setting, which is one of the places where coccinellid activities are most intensely monitored. In addition, the species 1s some- what cryptically colored and its arboreal habits would further limit the chances of a casual sighting. The very reduced elytral markings of the North American popula- tions of H. quadripunctata would also tend to make it resemble lightly marked individ- uals of other more common species, and it might be ignored by a collector who had already “filled his quota.” It would seem unlikely that the North American specimens of H. quadripunctata represent fortuitously intercepted individ- uals transported from Europe, or specimens collected in Europe which were subsequent- ly mislabeled. The proximity of the three collection sites, and the remarkable simi- larity between the color patterns of the eight specimens, strongly suggests that they have descended from a single long established population. Probably the species was acci- dentally introduced to the east coast of North America on board European ships. This method of transport has been suggested for Propylea quatuordecimpunctata (L.) (Laro- chelle and Lariviere 1980) another Euro- pean coccinellid which became established in the vicinity of Montreal. The habit of 1. quadripunctata of forming large dormant aggregations in crevices of tree bark (Klaus- nitzer and Klausnitzer 1986, Bielawski 1961) would facilitate its successful trans- port in this manner. Harmonia quadripunctata is one of four 410 exotic coccinelline species reported as es- tablished in eastern North America in the last couple of decades. The other three species are Propylea quatuordecimpunctata (Chantal 1972), Coccinella septempunctata L.(Angalet and Jacques 1975) and Hippo- damia variegata (Goeze) (Gordon 1987a). In all four cases the established populations were first recorded near the east coast, from Montreal to New Jersey, and appear to have resulted from either undocumented or un- intentional releases. The similarities which exist in the attributes of these four species are also interesting to note. They are all members of the tribe Coccinellini, are broadly endemic to the palaearctic, pri- marily aphidophagous, and form large ag- gregations during periods of dormancy. At this point the similarity ends. The three pre- viously reported exotic species habituate herbs and grasses. Consequently these coc- cinellines have drawn considerable interest for their potential role as biological control agents of the introduced Russian wheat aphid Diuraphis noxia (Mordvilko). In con- trast, H. quadripunctata is almost exclu- sively an arboreal species. Its habitat and prey preferences in the New World will most likely coincide with those of our native species of Anatis Mulsant and Myzia Mul- sant. Harmonia quadripunctata will prob- ably not play an important role in the suppression of aphids in most agricultural settings, with the possible exception of some orchard crops. European populations have been recorded on aphis pomi (Degeer) in apple orchards (Asgari 1966), but the species is most typically associated with Cinara sp. in forest and woodland habitats (Klausnit- zer and Klausnitzer 1986). ACKNOWLEDGMENTS For the use and/or loan of specimens I thank the following individuals and insti- tutions: R. D. Gordon, U.S. National Mu- seum of Natural History; Stuart McKamey, private collection; L. H. Herman, American PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Museum of Natural History; J. A. Chemsak, Essig Museum of Entomology; D. H. Ka- vanaugh, California Academy of Sciences; and C. L. Hogue and R. R. Snelling, L.A. County Museum. I also thank K. S. Hagen for professional encouragement and the use of his reference library, and R.D. Gordon and M. A. Brown for their comments on earlier drafts of this manuscript. LITERATURE CITED Angelet, G. W. and R. L. Jacques. 1975. The estab- lishment of Coccinella septempunctata L. in the Continental United States. United States Dept. Agr. Coop. Econ. Insect Rep. 25: 883-884. Asgari, A. 1966. Untersuchungen uber die im Raum Stuttgart Hohenheim als wichtigste Pradatoren der grunen Apfelblattlaus (Aphidula pomi Deg.) auf- tretenden Arthropoden. Z. angew. Zool. 53: 35- 93. Bielawski, R. 1961. Die in einem Krautpflanzenve- rein und in einer Kieferschonung in Warszawa/ Bielany auftretenden Coccinellidae (Coleoptera). Fragm. faun. Warszawa 8: 485-525. Chantal, C. 1972. Additions a la faune Coleopterique du Quebec. Nat Can. 99: 243-244. Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Jour. New York Entomol. Soc. 93: 1-912. . 1987a. The first North American Records of Hippodamia variegata (Goeze) (Coleoptera: Coc- cinellidae). J. New York Entomol. Soc. 95(2): 307- 309. 1987b. A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occ. Pap. syst. Ent. 3, 46 pp. lablokoff-Khnzorian, S. M. 1982. Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Paris, 568 pp. Klausnitzer, B. and H. Klausnitzer. 1986. Marien- kafer (Coccinellidae). Die Neue Brehm-Bucherei 451, GDR, 104 pp. Larochelle, A. and M.-C. Lariviere. 1980. Propylea quatuordecimpunctata L. (Coleoptera: Coccinel- lidae) en Amerique du Nord: etablissement, hab- itat et biologie. Bull. Invent. Ins. Quebec 2(1): 1- 9. Mader, L. 1926-37. Evidenz der palaarktischen Coc- cinelliden und ihrer Aberrationen in Wort und Bild, I. Teil. Wien, 412 pp. Sasaji, H. 1971. Fauna Japonica: Coccinellidae (In- secta: Coleoptera). Tokyo, 345 pp. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 411-415 A NEW SPECIES OF WATER SCAVENGER BEETLE, GUYANOBIUS SIMMONSORUM, FROM BRAZIL (COLEOPTERA: HYDROPHILIDAE) PAUL J. SPANGLER Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. Abstract.— A new species of water scavenger beetle, Guyanobius simmonsorum, from a tributary of the Rio Xingu in the state of Para, Brazil is described. The beetle is illustrated by pen and ink drawings and SEM micrographs and is distinguished from the only other species described in the genus, Guyanobius adocetus Spangler. Key Words: beetle, Brazil The genus Guyanobius was described for the single species, G. adocetus Spangler (1986) from Guyana. This second species of Guyanobius was collected in Brazil short- ly after the description of the genus was pub- lished and is described to further define the genus. The specimens of this new species were collected from a small shaded tributary of the Rio Xingt where it flows through a lowland tropical rainforest in the Brazilian state of Para. The specimens were collected during a biological survey of the area in ad- vance of the construction of a hydroelectric dam on the river. If the dam is constructed as planned, the habitat from which this new species was collected will be inundated. Guyanobius simmonsorum, NEw SPECIES Figs. 1-11 Holotype 4.— Form and size: Hemispher- ical, strongly convex dorsally (Figs. 1, 2). Length, 2.98 mm; greatest width, 2.24 mm. Color: Shiny black dorsally except narrow band on anterior margin of head, lateral margins of pronotum, and very narrow lat- eral margins and posterior half of sutural margins of elytra dark reddish brown. Ven- Hydrophilidae, Guyanobius simmonsorum, new species, water scavenger ter light reddish brown except metasternal disc and middle of abdominal segments 1 and 2 slightly darker reddish brown. Head: Very finely, sparsely punctate; dis- cal punctures separated by 4 to 8 times puncture diameter; punctures across base of head between eyes smaller and sparser than discal punctures and separated by 3 to 4 times puncture diameter. Clypeus (Fig. 4) strongly, broadly expanded anteriorly and laterally, concealing labrum (Fig. 4); lateral margin extending deeply into eye; anterior margin feebly arcuate apicomedially. Men- tum shallowly concave, moderately broad and moderately emarginate apicomedially; surface moderately coarsely, densely punc- tate; punctures separated by 3 to 5 times puncture diameter. Submentum shallowly concave and densely, finely punctate; each puncture bearing a seta. Thorax: Pronotum widest at posterior third; strongly rounded laterally; shallowly emarginate apically and feebly arcuate ap- icomedially (Fig. 1); truncate posteriorly; narrowly rimmed laterally and anterolat- erally behind eyes; not rimmed posteriorly except at posterolateral angles; sides nearly 412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-9. Guyanobius simmonsorum, new species, 3: 1, habitus, dorsal view, x 30; 2, habitus, ventral view, x 25; 3, prosternum, mesosternum, metasternum, x60; 4, head, ventral view, x60; 5, metasternum and ab- domen, x60; 6, setation, hind margin of abdominal sternum 2, x 1000; 7, hind femur, x 100; 8, hind tibia, x 120; 9, protarsal claws, x 800. VOLUME 92, NUMBER 3 Figs. 10-13. 10, 11. vertical, extending well below prosternum; discal punctures finer and slightly more widely separated than discal punctures of head; most punctures separated by 6 to 8 times puncture diameter; lateral punctures slightly coarser. Prosternum with distinct, keel-like, medial process on anterior third; keel extending beyond anterior margin of prosternum; a tuberculate process on pos- teromedial margin (Figs. 2-4). Mesoster- num with moderately broad, triangular pro- tuberance between and slightly in front of mesocoxae (Fig. 3). Metasternum shiny, glabrous (except a few setae behind meso- coxae); with raised, broadly triangular area medially; sides shallowly concave; metepi- sterna pubescent. Procoxae sparsely finely setose laterally but with 12 very stout, dark- er setae ventroapically. Profemora densely punctate and pubescent ventrally on basal two-thirds; mesofemora and metafemora, 413 Guyanobius simmonsorum, new species, genitalia, 4: 10, ventral view; 11, lateral view. Figs. 12, 13. Guyanobius adocetus Spangler, genitalia, 6: 12, ventral view; 13, lateral view. except apical fourth, densely punctate and pubescent ventrally (Fig. 7); male protarsal claws with broad, tooth-like base (Fig. 9); metatarsal claws without tooth-like base. Elytra with sides nearly vertical, extending well below mesosternum, metasternum, and abdominal sterna (Fig. 2); without sutural striae; finely, sparsely punctate; punctures larger than those on pronotal disc and disar- rayed except as follows. Each elytron with 1 incomplete, indistinct row of moderately coarse, shallow punctures laterally, punc- tures separated by about 2 times puncture diameter; lateral punctate row starting a short distance behind humeral area and ex- tending a distance about equal to % length of elytron; with 4 additional poorly defined rows of widely separated, seta-bearing punctures. Lateral margin of each elytron narrowly rimmed from base to apex. Scu- tellum flat, triangular; surface finely, sparse- 414 ly punctate; punctures separated by 3 to 9 times puncture diameter. Abdomen: Sterna | and 2 strongly con- cave; with coarse, sparse, seta-bearing punc- tures; punctures separated by 2 to 4 times puncture diameter; posterior margin of ster- num 2 with a dense row of robust setae (Figs. 5, 6). Remaining sterna finely and densely punctate and densely pubescent. Male genitalia: As illustrated (Figs. 10, 11). Female.—Similar to male except average size 1s larger. Variations.— Males (N = 10) varied in length from 2.86 to 3.30 mm (=3.05) and in width from 2.07 to 2.39 mm (=2.23); females (N=10) varied in length from 2.94 to 3.53 mm (=3.27) and in width from 2.30 to 2.60 mm (=2.47). The concavity of sterna 1 and 2 of some specimens bears a lens- shaped hyaline mass as is seen on specimens of Laccobius and Chaetarthria. The hyaline mass Is not present on many specimens be- cause it is easily dislodged when specimens are placed in alcohol. Comparative notes.—Guyanobius sim- monsorum is very similar to G. adocetus Spangler (1986) from Guyana but may be distinguished from it as follows (character state for G. simmonsorum stated first). Ely- tral rim present from base to apex vs rim disappearing at apical fourth; elytron with 1 incomplete, indistinct row of moderately coarse shallow punctures laterally and be- hind humeral area vs elytron with 3 rows of coarse, distinct punctures laterally be- hind humeral area. Mentum with entire sur- face coarsely, densely punctate vs surface of mentum moderately coarsely, sparsely punctate laterally and meson almost gla- brous. Parameres (Figs. 10, 11) with apices subtriangular vs apices of parameres round- ed. (Figs; 12, 13), Type data.—Holotype male: BRAZIL: PARA: Altamira (ca. 60 km S), Rio Xingu Camp, 52°22'W, 3°39’S, 14 Oct 1986, P. J. Spangler & O.S. Flint, colln #23, left branch off 1st jungle stream on trail 1; deposited in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Museu de Zoologia, Universidad de Sao Paulo, Brazil. Allotype: same data as ho- lotype. Paratypes: Same data as holotype, 24 4, 18 2: same data except: 3 Oct 1986, colln #6, 11 6, 11 9; 7 Oct 1986, colln #12, 3 4, 2 2. Paratypes will be deposited in the British Museum (Natural History), London; Cali- fornia Academy of Sciences, San Francisco; Canadian National Collection, Ottawa; In- stitut Royal de Histoire Naturelle de Bel- gique, Bruxelles; Museo Argentina de Cien- cias Naturales, Buenos Aires; Museo de Zoologia, Universidad de Sao Paulo, Sao Paulo; National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C.; Zoologische Staatssammlung Min- chen, Miinchen. Etymology.—The specific epithet sim- monsorum 1s named for the Jerry L. Sim- mons family for their interest in and support of aquatic Coleoptera research at the Smith- sonian Institution. Habitat.— The type material was collect- ed from a small, shaded stream that flowed slowly from pool to pool through the jungle near the base camp. The stream was at an altitude of 90 meters; had a velocity of 2 m/min; was clear, a meter wide, and up to 75 cm deep; had a sand and leafy substra- tum, and colorimetric readings of hardness, 0, and a pH 5. The water temperature was 25.5°C and the air temperature was 27.5°C at the time the specimens were collected. The following aquatic insects were asso- ciated with G. simmonsorum in the same habitat: COLEOPTERA: DRYOPIDAE: Pelonomus, Dryops. DYTISCIDAE: Lacco- philus, Derovatellus, Desmopachria, Bides- sines. ELMIDAE: Tyletelmis, Heterelmis; HYDRAENIDAE: Hydraena; HYDRO- PHILIDAE: Derallus, Paracymus, Helocha- res, Hydrochus, Phaenonotum, Notionotus; LUTROCHIDAE: Lutrochus. GYRINI- DAE: Gyretes. HETEROPTERA: BELOSTOMATI- DAE: Belostoma. GERRIDAE. HELOTRE- VOLUME 92, NUMBER 3 PHIDAE: Helotrephes. HYDROMETRI- DAE: Hydrometra. NAUCORIDAE: Ambrysus. NEPIDAE: Ranatra. VELI- IDAE: Microvelia, Paravelia, Rhagovelia. ACKNOWLEDGMENTS For support of the fieldwork during which the specimens of this species were collected, I thank the Consorcio Nacional de Engen- heiros Consultores, S. A., of Brazil for their patronage; Paulo Vanzolini for the invita- tion to participate in the fieldwork; and Maria Beatriz Ribiero do Valle for man- aging logistical problems so effectively. 415 The following individuals contributed to the preparation of this article and I thank them for their assistance: Young T. Sohn, biological illustrator, for the art work; Su- sann G. Braden and Robin A. Faitoute, for the SEM micrographs; and Phyllis M. Span- gler, for typing the manuscript. LITERATURE CITED Spangler, P. J. 1986. Anew genus and species of water scavenger beetle, Guyanobius adocetus, from Guy- ana and its larva (Coleoptera: Hydrophilidae: Hy- drobiinae). Proceedings of the Entomological So- ciety of Washington 88(3): 585-594. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 416-421 MORPHOLOGY AND MATING CONFIGURATION OF GENITALIA OF THE ORIENTAL COCKROACH, BLATTA ORIENTALIS L. (BLATTODEA: BLATTIDAE) NONGGANG BAO AND WILLIAM H. ROBINSON Urban Pest Control Research Center, Department of Entomology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061. Abstract. — Male and female genitalia of the oriental cockroach, Blatta orientalis L., are redescribed and illustrated. New terms are assigned to lobes of the left and right phallo- meres of the male genitalia. Aspects of the mating behavior are presented, including the function of the titillator in initiating copulation. The configurational arrangement of the genitalia during copulation is described and illustrated, including the functions and po- sitions of the three phallomeres of the male genitalia with the valvulae of the female ovipositor. The five modified lobes of the male left phallomere and the scoop-like ventral phallomere function mainly to stabilize the female valvulae on each side, while the serrata of the right phallomere performs to separate the paired first valvulae from the center. This spreading configuration provides for the successful transfer of spermatophore. Key Words: The oriental cockroach, Blatta orientalis L., 1s native to North Africa and now is distributed throughout the temperate re- gions of the world (Cornwell 1968, Cochran 1982, Woo 1987). It is the dominant cock- roach pest species in Great Britain (Ragge 1965, Cornwell 1968). Mampe (1972) and Piper and Frankie (1978) reported it to be a seasonal household pest in portions of the northwestern, midwestern, and southern United States. The pest status of the oriental cockroach has been documented by Thoms and Robinson (1986, 1987). Snodgrass (1933) provided a comprehen- sive description of the female genitalia and related musculature of B. orientalis. Snod- grass (1937) also described and illustrated the male genitalia of this species and pre- sented a hypothetical plan for the phylo- genetic development of the male genitalia and corresponding phallic musculature. Blatta orientalis, morphology, genitalia, copulation Marks and Lawson (1962) and McKittrick (1964) compared and illustrated ovipositors of several cockroach species. They consid- ered the structure of the ovipositor of the oriental cockroach very similar to that of Periplaneta americana (L.) and other Peri- planeta species. The morphology of the gen- italia of B. orientalis has not been re-ex- amined since the early descriptions and interpretations by Snodgrass (1933, 1937) and McKittrick (1964). This study provides an overall revision of the female and male genitalia and describes the mating behavior of the oriental cockroach. The configura- tional arrangement of the genital structure during copulation are also described and il- lustrated. MATERIALS AND METHODS Adult cockroaches were obtained from field and laboratory colonies. Genitalia of VOLUME 92, NUMBER 3 male and female adults were examined and illustrated using an optical dissecting mi- croscope and ocular grid. Specimens were preserved in 70% ethanol and treated with 10% aqueous potassium hydroxide (KOH) for about 24 hours at room temperature be- fore examination and illustration. The gen- italia were preserved in 70% ethanol with a few drops of glycerine for further exami- nation and photography. The terms and abbreviations used in this study to describe the male and female gen- italia of B. orientalis were adopted from those used by Snodgrass (1933, 1937) and McKittrick (1964). New terms were as- signed to some structures of the male gen- italia. For the study of the configurational ar- rangement of genitalia during copulation, male and female adults were separated soon after the final molt and kept segregated for 7 days. Mating behavior was observed and recorded when the male and female were placed together. After coupled for 20-30 minutes, the male and female were anes- thetized with carbon dioxide, then killed with ethyl acetate. The configuration of the male and female genitalia during copulation was determined by dissecting the genital segments of 23 pairs of freshly killed spec- imens or those preserved in 10% KOH for 24 hours. RESULTS AND DISCUSSION Female genitalia.—The reduced eighth and ninth abdominal segments, or the gen- ital segments, bear appendages that form the ovipositor of the female. The seventh abdominal sternum (SVIJ) is expanded and prolonged posteriorly to form two large, valve-like lobes (SVIIL) that conceal the Ovipositor. Dorsal to the SVIIL, the paired clefts are the tenth tergum (TX). The cerci lie beneath the basal corners of the tenth tergum. Beneath the TX at the posterior end of the abdomen are a pair of sclerotized paraprocts (PAPT). These may be remnants of the eleventh abdominal segment. The 417 SV SVIII SVII Fig. 1. Diagrammatic median view of female Blar- ta orientalis genital segment. ACGL, accessory gland; AN, anus; GC, genital chamber; GPR, gonopore; MOD, median oviduct; PAPT, paraproct (T XI); RECT, rec- tum; SPR, spermathecal pore; SPT, spermatheca; SVI, sixth sternum; SVII, seventh sternum; SVIII, eighth sternum; SVIIL, lobe of seventh sternum; TVI, sixth tergum; TVII, seventh tergum; TVIII, eighth tergum; TIX, ninth tergum; TX, tenth tergum; VAI, first val- vula; VAII, second valvula; VST, vestibulum. (Mod- ified after Snodgrass 1933). anus lies centrally in the membrane between the pair of paraprocts and the tenth tergum (Fig. 1). There are two internal chambers, the ves- tibulum (VST) and the genital chamber (GC), which are formed by the modified SVII, SVIII, SIX, SX, and TIX. The ovi- positor lies in the vestibulum, with the sev- enth sternum as the floor of the chamber. Anterior to vestibulum is the smaller genital chamber, with the invaginated eighth ster- num as its floor. The median oviduct (MOD) empties into the floor of the genital chamber by way of the gonopore (GPR). The sper- matheca (SPT) possesses a pore which emp- ties into the roof of the genital chamber. The accessory glands (ACGL) open on the roof of the vestibulum between the second pair of valvifers (VLFII) near their bases (Figs.1, 2). The roof of the genital chamber possess- es a pair of large, lateral sclerites or basi- valvulae (BSV) and a median sclerite, or spermathecal plate (SPPL) (Fig. 2). These 418 Fig. 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Female genitalia of B. orientalis (ventral view). ACGR, acessory gland pore; BSV, basivalvula (SVIII); GPR, gonopore; LTIX, ninth laterosternite (SIX); LTSH, laterosternal shelf (SVIII); PAPT, paraproct (T XI); PATG, paratergite (TIX); SPR, spermathecal pore; SPPL, spermathecal plate; TX, tenth tergum; VAI, first valvula; VAII, second valvula; VAIII, third valvula; VLFI, first valvifer; VLFII, second valvifer. sclerites may represent the secondary sclerotizations between the eighth and ninth segments. The floor of the genital chamber possesses two pairs of sclerites, or the latero- sternal shelves (LTSH), which are derived from the eighth sternum. The gonopore lies in the membrane between the laterosternal shelves. The roof of the vestibulum, consisting of the fused sterna of the ninth and tenth ab- dominal segments, supports the base of the ovipositor. The slender sclerite, or parater- gite (PATG), on the roof edge of the ves- tibulum is derived from the ninth tergum. The irregular sclerites, the ninth lateroster- nites (LTIX), on the roof of the vestibulum, with the anterior ends fused to the first val- vifers (VLFI), are the sternites of the ninth segment. The ovipositor has two pairs of valvifers and three pairs of valvulae. The sclerotized first pair of valvulae (VAI) of the ovipositor are widely divergent at their bases, fused to the first pair of valvifers which may be de- rived from the eighth segment. The scler- otized second pair of valvulae (VAII) arise laterally from the fused, highly sclerotized second pair of valvifers. The pair of third valvulae (VAIII) are largely membranous and arise from the second valvifers, mesad of the second valvulae and beneath the first valvulae (Fig. 2). Male genitalia.—The ninth abdominal segment, or the genital segment, bears ap- pendages that form the genitalia of the male (Fig. 3). The bilobed tenth tergum (TX) forms the roof of the male genital chamber. The ninth sternum (SIX) bears a pair of small styli (STY), which may represent the remnants of the male gonopods. The paired paraprocts (PAPT) lie beneath the tenth ter- gal plate. The anus is situated in the mem- brane between the tenth tergum and the paraprocts. The external genitalia of the male consist of genital lobes or phallomeres (PHM) as- VOLUME 92, NUMBER 3 Fig. 3. 419 Male genitalia of B. orientalis (dorsal view). ACL, acutolobus; DEJR, ejaculatory duct pore; FLX, falax; INL, inner lobe; LPHM, left phallomere; MDL, middle lobe; PAPT, paraproct; PSP, pseudopenis; RPHM, right phallomere; SIX, ninth sternum; SRT, serrata; STY, stylus; TIX, ninth tergum; TLT, titillator; TX, tenth tergum; VPHM, ventral phallomere. sociated with the genital pore. Phallomeres consist of three major parts: the left, right, and ventral phallomere. The right phallomere (RPHM) has two sclerites, the serrata (SRT) and falax (FLX), which lie on the center of the genital cham- ber towards the right. The serrata of the right phallomere, which lies in the center of the genital chamber, bears a fork-like scler- otized structure with two sharp processes on the left and a highly sclerotized hook- like structure on the posterior right. Snod- grass (1937) considered the serrata to be composed of three sclerites. The “right lob- ule” is membranous in nature. Since there are no sutures and sulci present in the ser- rata, there is no evidence to divide the ser- rata into three sclerites. The only difference is the unequal degrees of sclerotization in different portions of the serrata. The falax of the right phallomere is a simple sclerite situated laterally and joined with the right side of the serrata. The left phallomere (LPHM) is the most complicated phallic organ of the male gen- italia. It bears five elongated structures: the titillator (TLT), the outermost elongated and sclerotized lobe with a hook at the base of its pointed tip; the pseudopenis (PSP), mesad of the titillator with a bulbous tip; the middle lobe (MDL), next to the inner side of pseudopenis and partially sclero- tized; the acutolobus (ACL), beneath the middle lobe, with a sclerotized hook and a partially sclerotized tooth-like process on its inner side; and the inner lobe (INL), the small innermost lobe with a sclerotized pro- cess on its posterior right end. The ventral phallomere (VPHM) is a broad, scoop-like lobe projecting to the right from the posterior surface of the genital membrane. The ejaculatory duct (DEJ) empties into the floor of the genital chamber by way of the male gonopore, which is sit- uated on a small membranous elevation on the base of the ventral phallomere. Mating behavior and configurational ar- rangement of genitalia at copulation.—The mating behavior of B. orientalis observed in this study was very similar to previous of Fig. 4. Configurational arrangement of genitalia during copulation (dorsal view). ACL, acutolobus; FLX, falax; LVAI, first left valvula; MDL, middle lobe; PSP, pseudopenis; RVAI, first right valvula; SRT, serrata; TLT, titillator,; VPHM, ventral phallomere. reports by Roth and Willis (1952) and Barth (1970). The description by Barth (1970) is more similar to the observations in this study. A male is usually stimulated by con- tact with a female body or antennae. The aroused male typically raises his wings, and extends his abdomen, and depresses it against the substratum. Simultaneously, the titillator of the left phallomere is rhythmi- cally protruded out of the genital chamber. The male actively searches for females and frequently attempts to back under a female from in front of her. A receptive female will move forward on the male, with her mouth- parts continuously contacting his abdomi- nal terga at the same time the male moves backwards. When the female moves to “feed” on the first abdominal tergum of the male, she spreads apart her vestibulum. Si- multaneously, the male extends his abdo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON men and rapidly raises and inserts his gen- ital segments to the vestibulum of the female. The hooked titillator of the left phallomere is the first phallic organ which fastens to the Ovipositor to achieve the connection. Once the connection is established, the male swings out from underneath the female and rotates 180° to attain an opposed position. Then, the distal segments of both the sexes are tightly hooked together by way of the male genital organs inserted in the vestibu- lum between the ovipositor and the lobes of SVII. Copulation was observed to last 30 to 45 minutes. During copulation the titillator and acu- tolobus of the left phallomere spread a great distance from the original left-side position towards the night and assist the ventral phal- lomere to clasp the first left-valvula (LVAI) of the ovipositor (Fig. 4). Both structures have strong, pointed tips and hooks, which help to clasp the valvulae tightly. The other lobes i.e. the inner lobe, middle lobe, and pseudopenis of the left phallomere hold the first right-valvula (RVAI) on the left-ven- tral side. The pseudopenis has a groove on its inner side, in which the first right-valvula is placed. The ventral phallomere grips the first left-valvula on its groove, which is formed by means of folding its distal perim- eter upwards. The forks of the serrata grip the first right-valvula from the inner side towards outside, which hold the valvula along with the three lobes of the left phal- lomere. The hook of the serrata grips the first left-valvula from the inner side towards the outside. It is associated with the ventral phallomere, the titillator, and acutolobus. The serrata of the right phallomere keeps the first pair of valvulae open from the cen- ter, so that the other two pairs of valvulae and the genital chamber are exposed. The male gonopore, with associated membrane on the base of the ventral phallomere, pro- jects into the female genital chamber where the spermathecal sac and the female gono- pore are located. This spreading of the val- vulae provides for the successful transfer of VOLUME 92, NUMBER 3 a spermatophore from the male ejaculatory duct onto the ventrally projecting sper- mathecal papilla of the female. Gupta (1947) reported a similar mating behavior and con- figurational arrangement of genitalia at cop- ulation for P. americana. He illustrated the coupled structure of the male and female genitalia and pointed out that the titillator of the male genitalia was important in forc- ing open the female vestibulum (gynatrium) to allow the entry of the male genitalia. He also described the importance of the right phallomere which functioned as the main clasping organ to hold the valvulae of the female genitalia during copulation. The configurational arrangement of the genitalia of B. orientalis is similar to that of P. amer- icana during copulation. LITERATURE CITED Barth, R. H. 1970. The mating behavior of Peripla- neta americana (Linnaeus) and Blatta orientalis Linnaeus (Blattaria, Blattinae), with notes on 3 additional species of Periplaneta and interspecific action of female sex pheromones. Z. Tierpsychol. 27: 722-748. Cochran, D. G. 1982. Cockroaches—Biology and control. World Health Org., Vector Biol. Control Div., Pub. 856. Cornwell, P. B. 1968. The Cockroach. Vol. 1. Hutch- inson and Co., Ltd. London. Gupta, P.D. 1947. On copulation and insemination in the cockroach, Periplaneta americana (Linn.) Proc. Nat. Inst. Science, India. 13: 65-71. 421 Mampe, C. D. 1972. The relative importance of household insects in the continental United States. Pest Control 40(12): 24, 26-27, 38. Marks, E. P. and F. A. Lawson. 1962. A comparative study of the Dictyopteran ovipositor. J. Morph. 111: 139-172. McKittrick, F. A 1964. Evolutionary studies of cock- roaches. Cornell Univ. Agri. Exp. Station. Memoir 389. Ithaca, New York. Piper, G. L. and G. W. Frankie. 1978. Integrated management of urban cockroach populations, pp. 249-266. In Frankie, G. W. and C. S. Koehler, eds., Perspective in Urban Entomology. Academic Press, Inc., New York. Ragge, D.R. 1965. Grasshoppers, Crickets and Cock- roaches of the British Isles. Frederick Warne and Co. Ltd., London. Roth, L. M. and E. R. Willis. 1952. A study of cock- roach behavior. Am. Midl. Nat. 47: 66-129. Snodgrass, R. E. 1933. Morphology of the insect ab- domen. Part II. The genital ducts and the ovipos- itor. Smithsonian Misc. Coll. 89: 1-148. 1937. The male genitalia of Orthopteroid insects. Smithsonian Misc. Coll. 96: 1-107. Thoms, E. M. and W. H. Robinson. 1986. Distri- bution, seasonal abundance, and pest status of the oriental cockroach (Orthoptera: Blattidae) and an evanid wasp (Hymenoptera: Evaniidae) in urban apartments. J. Econ. Entomol. 79: 431-436. . 1987. Distribution and movement of the ori- ental cockroach (Orthoptera: Blattidae) around apartment buildings. Environ. Entomol. 16: 731- 131. Woo, F. 1987. Investigations on domiciliary cock- roaches from China. Acta Entomologica Sinica 30: 430-438. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 422-425 A NEW DARJILINGIA (SYMPHYTA: TENTHREDINIDAE) FROM TAIWAN IcHu1 TOGASHI Ishikawa Agricultural College, 1-308, Suematsu, Nonoichi-machi, Ishikawa Prefecture 921, Japan. Abstract. — Darjilingia varia sp. nov. from Taiwan is described and illustrated. Key Words: Darjilingia, Tenthredinidae The genus Darjilingia was known only by its type species, D. gribodoi (Konow), from India (Himalayas) and Burma. It probably does not occur in Borneo as Konow stated (Konow 1896; Malaise 1934). Malaise (1963) questionably recorded the genus from Formosa. Through the courtesy of Dr. A. Shinohara, Department of Zoology, Na- tional Science Museum (Nat. Hist.), Tokyo, I had an opportunity to examine four spec- imens of Darjilingia that he collected in Tai- wan in 1976 and 1977. These specimens represent a new species, and I describe the species below. They are the first definite re- cord of Darjilingia from Taiwan. Darjilingia varia Togashi, NEw SPECIES (Figs. 1-18) Female.— Length 8.5 mm. Head and tho- rax black, with following parts yellowish white: clypeus, labrum, mandible except for apex, maxillary and labial palpi, posterior margin of pronotum, oval macula on me- soscutellum, cenchri, metascutellum, a small spot on upper portion and an elongate mac- ula on lower portion of mesopleuron (Fig. 15), and posterior portion of meseepime- ron. Apex of mandible red. Parapteron, teg- ula, and a small spot on central portion of mesonotal lateral lobes (Fig. 12) reddish yel- low. Wings hyaline, stigma except for basal portion and veins dark brown. Basal por- tion of stigma of forewing whitish (Fig. 1). Legs reddish brown but all coxae, trochan- ters, and basal 3 of hind femur yellowish white; knee of hind femur, apical portion of hind tibia, and hind basitarsus except for apical 3 black; apical 3 of hind basitarsus and following four segments of hind tarsus yellowish white (Fig. 7). Abdomen reddish brown but Ist tergite and Sth to 8th tergites dark brown or with large dark brown mac- ulae (Fig. 1); 9th tergite yellowish white; 8th sternite, 2nd valvifer, and sawsheath dark brown to black. Head seen from above transverse; post- ocellar area slightly convex; lateral furrows distinct and deep; postocellar and interocel- lar furrows slightly depressed (Fig. 2); OOL: POL:OCL = 1.7:1.0:2.4; lateral and median foveae slightly depressed; ratio between an- tenno-ocular distance and distance between antennal sockets about 0.7:1.0; apical mar- gin of clypeus as in Fig. 3; occipital carina distinct but upper portion nearly obsolete (Figs. 2, 4); post-orbital groove distinct (Fig. 4); malar space narrow, nearly '2 as long as diameter of front ocellus. Antenna (Fig. 5) nearly as long as costa of forewing; relative lengths of segments about 17621f07316:3.5:3'2225:2)021-8-2.2: Thorax: normal; mesoscutellum slightly raised. Wing venation as in Fig. 1; anal cell of hindwing sessile. Legs: front inner tibial VOLUME 92, NUMBER 3 423 mm mm 0.5 mm 5 Figs. 1-9. Darjilingia varia Togashi sp. nov. 1, dorsal view (paratype); 2, head, dorsal view; 3, head, front view; 4, head, profile; 5, antenna, lateral view; 6, front inner tibial spur, lateral view, 7, hind tibia and tarsus, lateral view; 8, tarsal claw; 9, sawsheath, lateral view. spur as in Fig. 6; tarsal claw with a large inner tooth, and with distinct basal lobe (Fig. 8); hind basitarsus slightly longer than fol- lowing four segments combined (ratio be- tween them about 1.00:1.07). Abdomen: sawsheath as in Fig. 9. Punctuation.—Head and thorax nearly impunctate, shining; abdominal tergites nearly impunctate, shining. Male.—Length 7.5 mm. Coloration as in female but 2nd to 6th tergites reddish brown, and last two abdominal segments yellowish white. Structures as in female except for sex- ual segments. Male genitalia and penis valve as in Figs. 10 and 11. Holotype: female, 13. HI. 1977, Wushe, Taiwan. Paratypes: | female, 4. V. 1976, Meifeng- Sungkang, Taiwan; | male, 3. V. 1978, Nan- shanchi, Taiwan; 1 female, 13. III. 1979, Wulai, Taiwan. Holotype and two para- types (female and male) are deposited in the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ \, \ Bo po <5 116 Figs. 10-13. valve. Figs. 12-14. Colour pattern of thorax of D. varia sp. nov. 12, typical form (holotype); 13 and 14, variations (paratypes). Figs. 15-18. Colour pattern of mesopleuron of D. varia sp. nov. 15, typical form (holotype); 16 and 17, variations (female paratypes); 18, typical form (male paratype). Department of Zoology, National Science Museum (Nat. Hist.), Tokyo. One paratype is deposited in the U.S. National Museum, Washington, D.C. Variation.—The coloration of the prae- scutum is reddish brown in the paratypes (Figs. 13, 14). The coloration of the meta- scutellum varies from black (Fig. 13) to yel- lowish white (Fig. 12), and the coloration of the mesopleuron varies from black (Fig. 15) to pale reddish yellow (Fig. 17). Also, the coloration of the abdomen is dark to light reddish yellow. Remarks.—This new species closely re- sembles D. gribodoi (Konow 1896) from Borneo (?), Burma, and India (Darjeeling, Khasia Hills, and Assam), but it is easily distinguished from gribodoi by the color- ation of the basal three segments of the an- oa Figs. 10-11. Male genitalia and penis valve of D. varia sp. nov. 10, male genitalia; 11, penis tenna (in gribodoi, the basal three segments of the antenna are reddish yellow to reddish brown), by the coloration of the apical por- tion of the hind tibia and hind basitarsus (in gribodoi, the legs are reddish yellow ex- cept for the basal portion of the coxae, and the hind basitarsus is black), and by the length of the antenna (in gribodoi, the an- tenna is nearly as long as the body length). ACKNOWLEDGMENTS I wish to express my sincere thanks to Dr. A. Shinohara, Department of Zoology, Na- tional Science Museum (Nat. Hist.), Tokyo, for the loan of the valuable specimens and to Dr. David R. Smith, Systematic Ento- mology Laboratory, USDA, Washington, D. C., for reviewing the manuscript. VOLUME 92, NUMBER 3 LITERATURE CITED Konow, Fr. W. 1896. Neue und einige bisher ver- kannte Arten aus der Familie der Tenthrediniden. Entomol. Nachrichten 20: 308-319. 425 Malaise, R. 1934. On some sawflies from the Indian Museum, Calcutta. Rec. Ind. Mus. 36: 453-474. 1963. Hymenoptera Tenthredinoidea, subfamily Selandriinae, key to the genera of the world. Ent. Tidskr. 84: 159-215. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 426-430 HOST SPECIFICITY OF CHAETORELLIA AUSTRALIS (DIPTERA: TEPHRITIDAE) FOR BIOLOGICAL CONTROL OF YELLOW STARTHISTLE (CENTAUREA SOLSTITIALIS, ASTERACEAE) D. M. Mappox,! A. MAYFIELD,! AND C. E. TURNER? Biological Control of Weeds, U.S. Department of Agriculture, Agricultural Research Service, Western Regional Research Center, 800 Buchanan Street, Albany, California 94710. Abstract.—The flower head tephritid fly Chaetorellia australis Hering was studied to determine its host specificity for biological control of Centaurea solstitialis L. (yellow starthistle) in the United States. Flies from flower heads of C. cyanus L. collected in northern Greece were tested for oviposition and development on nine plant species in no-choice host tests during the summers of 1986-87 in Albany, California. Oviposition and development occurred on only two species: flies damaged 93.7% (1986) and 79.6% (1987) of the heads of C. solstitialis and 85.8% of the heads of C. cyanus. No evidence of oviposition and development occurred on the other test plant species: Centaurea amer- icana Nutt., Centaurea rothrockii Greenm., Carthamus tinctorius L., Cirsium occidentale (Nutt.) Jeps., Helianthus annuus L., Zinnia elegans Jacq., and Lactuca sativa L.. More than 92% of the pupal fly-yielding flower heads produced only one pupal fly, while less than 8% of these flower heads had two pupal flies, indicating that the fly is not particularly gregarious. Key Words: Yellow starthistle (Centaurea solstitialis L., Asteraceae) is a winter annual that is a naturalized weed primarily in the western United States. Surveys indicate that it oc- curs in 208 counties in 23 states within the U.S. (Maddox et al. 1985), and infestations in California alone have reached an esti- mated 3.25 million gross hectares (Maddox and Mayfield 1985). The weed is a pioneer- 'D. M. Maddox is a Research Entomologist (retired) and A. Mayfield is a Biological Technician (retired). Current addresses of D.M.M. and A.M. : 18469 Chap- arral Dr., Penn Valley, CA 95946 and 5481 Crittenden St., Oakland, CA 94601 respectively. 2 Please direct all requests for reprints to C. E. Turn- er. biological control, weed, rangeland, insect ing species that is especially invasive on dis- turbed lands. Its primary economic impact is on rangelands where it reduces livestock productivity because of its unpalatability, competitiveness, and toxicity to horses (Maddox et al. 1985). Yellow starthistle is believed to be native to the eastern Medi- terranean Basin and western Asia (Prodan 1930). Multiple introductions have proba- bly occurred in the U.S. (Maddox and May- field 1985). An analysis of seeds contained in adobe bricks from early buildings in Cal- ifornia indicates that yellow starthistle was introduced in the nineteenth century after 1824 (Hendry 1931, Hendry and Bellue 1936, Maddox and Mayfield 1985). A search for natural enemies of yellow VOLUME 92, NUMBER 3 starthistle and other weedy members of the thistle tribe (Cardueae) was begun in Europe in the late 1950’s (Zwolfer et al. 1971). In 1985 the flower head weevil, Bangasternus orientalis (Capiomont) (Coleoptera: Cur- culionidae), was introduced from northern Greece for biological control of yellow star- thistle and is now successfully established in several western states (Maddox et al. 1986). Sobhian and Zwolfer (1985), in their studies of the yellow starthistle flower head— insect host system, show that the larvae of about 20 phytophagous insect species utilize the flower heads of yellow starthistle in the Mediterranean Basin and regions in the northern half of the Balkan Peninsula. Chaetorellia australis Hering is one of those flower head insects that offers promise as a biological control agent. Tephritid fly species, including C. australis, are consid- ered to be one of the most important ele- ments within the guild of insects utilizing the flower heads (Sobhian and Zwolfer 1985). They are important flower head feed- ers on asteraceous species in general, and are often either strongly monophagous or stenophagous on their host plants. Flower head feeders may be especially important as biological control agents because yellow starthistle is an annual weed that relies sole- ly on seed production to reproduce. This fly has previously been referred to as Chaeto- rellia hexachaeta australis Hering (White and Marquardt 1989). BIOLOGY The biology of C. australis in northern Greece is as follows (Sobhian and Zwolfer 1985, Sobhian and Pittara 1988): Three generations of flies occur per year. Females begin oviposition after adult emergence in the spring. Under uncrowded conditions, a female usually oviposits beneath an invo- lucral bract one egg per host flower head. Oviposition occurs preferentially on ma- ture, closed flower head buds. Each egg characteristically possesses a long filament, which can extend beyond the margins of a 427 bract. Under laboratory conditions, females may Oviposit up to 243 eggs during an ovi- positional period of up to 60 days. Hatched larvae tunnel through the involucre into the interior of the flower head, where they tun- nel through and feed on many ovaries and developing achenes. In one limited test, a single larva destroyed an average of 86.3% of the seeds in a flower head (Sobhian and Pittara 1988). According to Sobhian and Pittara (1988), the overwintering generation passes the winter as mature larvae within cocoons made of pappus hairs inside the flower heads of C. solstitialis. The overwin- tering larvae pupate and emerge as adults the following spring in April and May. The first generation larvae develop primarily on C. cyanus as it typically flowers earlier than C. solstitialis; the second and third (over- wintering) generation larvae develop on C. solstitialis. TAXONOMY, Host RANGE, AND GEOGRAPHIC DISTRIBUTION The following information is from White and Marquardt (1989) unless otherwise specified. Chaetorellia is a Palearctic genus of nine known species in the tribe Terelli- inae. Chaetorellia australis is one of prob- ably five Chaetorellia species in the C. ja- ceae species-group. Chaetorellia australis was originally described by Hering (1940) as a subspecies of Chaetorellia hexachaeta (Loew). The known hosts of all known species of Chaetorellia are species of Cen- taurea, Carthamus and Chartolepis in the Cardueae subtribe Centaureinae. The known hosts of species of Chaetorellia in the C. Jaceae species-group are species of Centau- rea. The host records for C. australis include C. solstitialis from Bulgaria, Greece, Hun- gary, Turkey and Moldavian SSR; C. cy- anus from Greece and Hungary; and C. de- pressa Bieb. from Turkey. Centaurea solstitialis is in the subgenus Solstitiaria, and C. cyanus and C. depressa are in the sub- genus Cyanus (Dostal 1976). In an extensive field sample of natural populations of a di- 428 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Chaetorellia australis no-choice host specificity tests in Albany, California, 1986-87. % Damaged Heads (with % Heads with Frass, Lar- Larvae, No. of Plants Total Heads vae, Pupae or % Heads with Pupae, or Plant Species Year Tested Tested Tested Pupal Cases) Frass Only Pupal Cases Centaurea solstitialis 1986 25 320 93.7 3222 61.5 Centaurea solstitialis 1987 16 1S 79.6 15.0 64.6 Centaurea cyanus 1987 20 268 85.8 1S 84.3 Centaurea americana 1986 25 39 0 0 0 Centaurea rothrockii 1986 14 45 0 0 0 Carthamus tinctorius (“Hartman’’) 1986 25 72 0 0 0 Carthamus tinctorius (“4440”) 1987 20 107 0 0 0 Cirsium occidentale 1986 25 73 0 (0) 0 Helianthus annuus 1986 25 68 0 0 (0) Zinnia elegans 1986 13 145 0 0 0 Lactuca sativa 1986 20 800 0 0 0 verse array of thistles throughout mainland Greece in 1985, C. australis was reared only from the flower heads of C. solstitialis and C. cyanus (Turner et al. in press). Host SpEcIFICITY TESTING Host specificity was measured in no- choice cage tests of C. australis oviposition and development on nine test plant species. The adult flies used in all tests were collected in northeastern Greece (by R. Sobhian) as larvae in heads of C. cyanus and shipped to the USDA-ARS quarantine facility of the Biological Control of Weeds Laboratory at Albany, California. Tests were carried out in this quarantine facility during the sum- mers of 1986 and 1987. Host test plant species were chosen on the basis of taxonomic affinity, economic sig- nificance, and place of origin. The test plant species were C. solstitialis, C. cyanus, C. americana Nutt., C. rothrockii Greenm., two varieties of Carthamus tinctorius L., Cir- sium occidentale (Nutt.) Jeps., Helianthus annuus L., Lactuca sativa L., and Zinnia elegans Jacq.. All test plant species are in the Asteraceae, and all are in the thistle tribe Cardueae except H. annuus, L. sativa and Z. elegans. Carthamus tinctorius (safflower) and H. annuus (sunflower) are oilseed crops, and H. annuus is native to the United States. Carthamus tinctorius var. “Hartman” is grown primarily in the northern plains area, while C. tinctorius var. “4440” was devel- oped primarily for California. Zinnia ele- gans (zinnia) is an ornamental, and L. sativa (lettuce) is a leafy food crop. Centaurea americana, C. rothrockii and C. occidentale are thistles native to the United States. Cen- taurea solstitialis and C. cyanus, the known hosts, served as controls. Test plants were grown in 15 cm pots and the plants of each species were arranged on a wood base platform (ca. 1 m?) according to a random number table. The test plants were enclosed by | m? screen cages that rest- ed on the wooden bases. One plant species was tested per cage. Thirteen to 25 plants were tested per plant species (Table 1). One pair (12 14) of newly emerged files were used per test plant; for example 25 plants and 25 pairs of flies (50 flies total) were enclosed by a cage in the test with C. americana (Ta- ble 1). The flies were released into each cage where they had free access to the test plants. Food for the adult flies was provided by a 30 ml shell vial with a wick containing a honey-water solution. The tests were con- ducted under natural light conditions (14.5- 16h light; average of 24°C daytime and 13°C nightime). Tests were terminated when all adult female flies were dead, the longest last- VOLUME 92, NUMBER 3 429 Table 2. Number of pupae per infested flower head in Chaetorellia australis no-choice host specificity tests in Albany, California, 1986-1987. No. Flower Heads % Flower Heads with 1 Fly Pupa per % Flower Heads with 2 Fly Pupae per Plant Host Species (Year Tested) Infested by Pupae Flower Head! Flower Head! Centaurea solstitialis (1986) 143 92.3 ileal Centaurea solstitialis (1987) 55 94.5 ‘ Centaurea cyanus (1987) 198 98.7 1.3 ' Pupal counts include intact living pupae and pupal cases from emerged flies. ing 36 days. The flower heads were then dissected and microscopically examined for evidence of C. australis feeding and devel- opment (frass, larvae, pupae or pupal cases). Tests were conducted between 18 June to 10 September 1986, and 15 June to 24 July 1987. RESULTS AND DISCUSSION Oviposition and larval development oc- curred only on C. solstitialis and C. cyanus, and these species were heavily attacked as evidenced by the presence of frass, larvae, pupae or pupal cases. Next generation adult flies emerged only in the cages containing C. solstitialis and C. cyanus. There was no evidence of host use of any of the other test plant species. For C. solstitialis, 93.7% of the flower heads in 1986 and 79.6% of the flower heads in 1987 were attacked by the fly, while 85.8% of the C. cyanus flower heads were attacked (Table 1). Our results are congruous with the known host records (White and Marquardt 1989, Turner et al. in press). All available information indi- cates that C. australis has a narrow host range with C. solstitialis, C. cyanus and C. depressa as the only known hosts. The re- stricted host range of this fly provides strong evidence that it is safe for introduction into the United States as a biological control agent for yellow starthistle. In the course of the flower head dissec- tions, the numbers of pupae and pupal cases (from emerged flies) per flower head were noted. Chaetorellia australis does not ap- pear to be gregarious as mostly only one or sometimes two pupae were found in infest- ed flower heads (Table 2). For C. solstitialis, 92.3% (1986) and 94.5% (1987) of the in- fested flower heads had only one pupa, and 98.7% of the infested flower heads of C. cyanus had only one pupa (Table 2). ACKNOWLEDGMENTS R. Sobhian collected the C. australis used in the host specificity testing. The California Department of Food and Agriculture pro- vided funding support for this study. L. A. Andres and S. L. Clement critically re- viewed the manuscript. LITERATURE CITED Dostal, J. 1976. 138. Centaurea L., pp. 254-301. In Tutin, T. G., V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters, and D. A. Webb, eds. Flora Europaea, vol. 4. Cam- bridge, University Press. Hendry, G. W. 1931. The adobe brick as a historical source. Agric. History 5: 110-127. Hendry, G. W. and M. K. Bellue. 1936. An approach to south-western agricultural history through adobe brick analysis. Symp. on Prehist. Agric., Univ. of New Mexico Bull., Univ. Press. 8 pp. Hering, E. M. 1940. Neue Arten und Gattungen. Si- runa Seva 2: 1-16. Maddox, D. M. and A. Mayfield. 1985. Yellow star- thistle infestations are on the increase. Calif. Agric. 39(11-12): 10-12. Maddox, D. M., A. Mayfield, and N. H. Poritz. 1985. Distribution of yellow starthistle (Centaurea sol- stitialis) and Russian knapweed (Centaurea re- pens). Weed Sci. 33: 315-327. Maddox, D. M., R. Sobhian, D. B. Joley, A. Mayfield, and D. Supkoff. 1986. New biological control for yellow starthistle. Calif. Agric. 40(1 1-12): 4-5. Prodan, I. 1930. Centaureele Romaniei, Buletinul 430 Academieide Cluj Institulul de Arte Grafice. “Ar- dealul” Strada, Memorandului No. 22. 256 pp. Sobhian, R. and H. Zwolfer. 1985. Phytophagous insect species associated with flower heads of yel- low starthistle (Centaurea solstitialis L.). Z. ang. Entomol. 99: 301-321. Sobhian, R. and I. S. Pittara. 1988. A contnbution to the biology, phenology and host specificity of Chaetorellia hexachaeta Loew (Dipt., Tephriti- dae), a possible candidate for the biological control of yellow starthistle (Centaurea solstitialis L.). J. Appl. Entomol. 106: 444-450. Turner, C. E., R. Sobhian, and D. M. Maddox. Host specificity studies of Chaetorellia australis (Dip- tera: Tephritidae), a prospective biological control PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON agent for yellow starthistle, Centaurea solstitialis (Asteraceae). Proc. VII Int. Symp. Biol. Contr. Weeds, 6-11 March 1988, Rome, Italy. Delfosse, E. S. (ed.) lst Sper. Patol. Veg. (MAF). (In press.) White, I. M. and K. Marquardt. 1989. A revision of the genus Chaetorellia Hendel (Diptera: Tephriti- dae) including a new species associated with spot- ted knapweed, Centaurea maculosa Lam. (Aster- aceae). Bull. Entomol. Res. 79: 453-487. Zwolfer, H., Frick, K. E., and L. A. Andres. 1971. A study of the host plant relationships of European members of the genus Larinus (Col.: Curculioni- dae). Tech. Bull. Commonw. Inst. Biol. Control 14: 97-143. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 431-443 TWO NEW SPECIES OF BLACK FLIES (DIPTERA: SIMULIIDAE) FROM NORTH AMERICA PETER H. ADLER Department of Entomology, Clemson University, Clemson, South Carolina 29634. Abstract.—The larva, pupa, female, and male of two new species of black flies from North America are described and illustrated. Simulium fionae, new species, a member of the S. vernum group, is known from Pennsylvania and New Hampshire. Simulium clar- icentrum, new species, a member of the S. pictipes group, is known from Arkansas, Missouri, Oklahoma, and Pennsylvania. Characters are provided to separate both species from closely related Nearctic taxa. Key Words: As of 1986, the North American simulid fauna north of Mexico consisted of 162 for- mally recognized species (Crosskey 1987); one additional species has since been de- scribed (Adler 1987). In the present paper, I describe two new Nearctic species, one a member of the Simulium vernum group and the other a member of the S. pictipes group. The taxonomic status of the Nearctic S. vernum group has been summarized by Ad- ler (1987). Thirteen species have been de- scribed formally, and the chromosomes of at least three additional species have been resolved in terms of the vernum standard (Brockhouse 1985, Hunter and Connolly 1986). Here, I describe all life stages of one of these latter species, Simulium sp. of Hunter and Connolly (1986). In the S. pic- tipes group, two species have been described (Shewell 1959) and a third, described herein, has been known cytologically as S. pictipes “A” (Bedo 1973, 1975). Procedure and nomenclature follow those used by Adler (1987), although measure- ments of adults of S. sp. were taken from alcohol-preserved specimens and of S. pic- tipes “A” from freeze-dried specimens. What previously were referred to as mandibular Simuliidae, Simulium, black fly, aquatic insect teeth are differentiated in this paper as ser- rations and sensillum, following the ter- minology of Craig and Craig (1986). All il- lustrations and photographs are based on material collected at the type localities. Ho- lotypes and some paratypes are deposited in the United States National Museum of Natural History, Washington, D.C. Addi- tional paratypes are deposited in the Ca- nadian National Collection, Biosystematics Research Centre, Ottawa (all chromosomal photographs are deposited here); the British Museum (Natural History), London; and the Clemson University Arthropod Collection, South Carolina. Simulium fionae Adler, NEw SPECIES Figs. 1-9 Simulium (Eusimulium) furculatum, Adler, 1983, (not Shewell 1952): 197, pupa. Simulium (Nevermannia) species near fur- culatum/croxtoni Adler & Kim, 1986: 29, larva, pupa. Simulium sp. Hunter & Connolly, 1986: 300, chromosomes. Simulium sp. near croxtoni-furculatum Hunter, 1987: 52, chromosomes. 432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4 0.1 mm Figs. 1-6. Simulium fionae new species. 1, Larval head capsule (dorsal view). 2, Larval hypostoma. 3, Larval head capsule (ventral view). 4, Female genital fork (sternite 9). 5, Male terminalia (ventral view with left gonocoxite, gonostylus, and parameres removed). 6, Male dorsal plate. Larva (final instar).— Length 6.3-7.5 mm __ simple setae (visible with phase contrast); (x = 6.8 mm, n = 47). Head capsule (Fig. headspots brown, distinct, delineating in- 1) pale yellowish brown, palest anterodor- fuscated area; eye spots rather large; line sally, covered with numerous, fine, pale, over eye spots brown, leading into heavy VOLUME 92, NUMBER 3 Figs. 7-9. Simulium fionae new species. 7, Larval habitus (dorsal view). 8, Larval cuticular setae (from dorsum of segment 7), as viewed with bright-field compound microscope. 9, Scanning electron micrograph of granules on portion of pupal thorax (dorsal view); ecdysial line is apparent on the left side. brown area posteriorly. Antenna with distal article faintly brown, median article trans- lucent or very pale yellowish brown dor- sally, proximal article pale yellowish brown; approximately '3—'2 of distal article sur- passing labral-fan stalk; proportions of ar- ticles (distal to proximal, excluding apical sensillum) approximately 1.0:1.3:1.0. La- bral fan with 41-51 (% = 46, n = 45) primary rays in New Hampshire specimens [35-40 (X = 37, n = 2) in Pennsylvania specimens]. Hypostomal teeth (Fig. 2) with median tooth and lateral teeth subequal in length and prominence; sublateral teeth variously smaller; lateral margin of hypostoma with 2 paralateral teeth and 2-5 lateral serrations per side; hypostoma with 2-3 prominent and 2-5 small lateral setae per side. Post- genal cleft (Fig. 3) about 1.3-1.5 times as long as wide, extending about /2-%, distance to hypostomal groove, widest at midpoint, rounded apically; subesophageal ganglion unpigmented. Maxillary palpus 2.8-3.5 times as long as basal width. Inner subapical ridge of mandible with double or triple sen- sillum proximal to | elongate serration. Lat- eral plate of thoracic proleg moderately sclerotized, rather broad, elongate, extend- ing almost entire length of apical article. Body (Fig. 7) reddish brown; intersegmental bands clear, distinct; ventral tubercles rounded, about 3 depth of abdomen at at- tachment points; abdominal segments 4-8 (sometimes 5-8) dorsally and laterally with 434 many short, multiply branched, dark brown setae (Fig. 8), ventrally with similar but much sparser dark brown setae; thoracic and remaining abdominal segments with many shorter, multiply branched, translucent se- tae (visible with phase contrast). Antero- dorsal arms of anal sclerite broadly con- nected to and subequal in length to pos- teroventral arms, associated with elongate, translucent, simple setae (visible with phase contrast). Rectal setulae pale, sparse (visible with phase contrast). Posterior proleg bear- ing 9-12 hooks in 61-64 rows. Anal papillae of 3 compound lobes. Pupa.— Length 3.1—3.9 mm (X = 3.4 mm). Head projecting downward, with numer- ous, minute, rounded granules; antennal sheath of female extending almost to pos- terior margin of head; antennal sheath of male extending about '/ distance to poste- rior margin of head. Gill (Fig. 58 in Adler and Kim 1986) about as long as pupa, con- sisting of 8 rather widely splayed filaments; base short, giving rise to 3 short petioles; dorsalmost petiole giving rise to 2 widely divergent filaments; lateral petiole giving rise to.a single lateral (or ventral) filament plus a dorsal pair on a petiole 1—5S times its basal width (rarely sessile); ventral petiole yield- ing a single lateral filament plus a ventral pair on a petiole 1-6 times its basal width (rarely sessile); filaments grayish, long, thin, tapering, with numerous furrows; surface sculpturing of base weakly differentiated. Thorax (Fig. 9) with numerous, minute, dome-shaped granules; trichomes simple (some occasionally bifid), slender, dark, 5— 6 on each side of thorax. Tergite I with 1 pair of setae; tergite I] with 5-6 anteriorly directed setae on each side of midline, and 1-2 minute setae laterally; tergites III and IV each with 4 anteriorly directed hooks on posterior margin on either side of midline, 1 small seta between and anterior to 2 out- ermost hooks, and 2-3 small setae laterally; tergites V to VIII each with row of fine, posteriorly directed spines along anterior margin, and 2-3 minute setae posteriorly on either side of midline; tergite IX with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pair of short, stout, slightly curving, dorsally directed terminal spines. Pleural membrane of segments II to VII usually with 1-3 mi- nute setae per side. Sternite III with about 3 minute setae per side; sternite IV poste- riorly with pair of closely set, moderately heavy, simple or bifid, anteriorly directed setae, and at least 2 fine, minute setae per side; sternite V posteriorly with 1 pair of closely set, anteriorly directed, multifid, hook-like setae, and at least | pair of fine setae per side; sternites VI and VII poste- riorly with | pair of distantly set, anteriorly directed, simple to trifid, hook-like setae, and at least 1 pair of fine setae per side; sternites VIII and IX with at most | pair of fine setae; sternites [TV to VIII with numer- ous, extremely fine microspines. Cocoon (Fig. 41 in Adler and Kim 1986) well formed, rather coarsely woven, with short, irregu- larly woven anterodorsal projection ac- counting for about 4.6-13.9% (x = 9.3%, n = 9) total cocoon length (in lateral view). Female.— General body color brown, with gray pruinosity, and silvery and pale golden pile. Length: body, 2.9-3.2 mm (X = 3.0 mm, n = 5); wing, 3.1-3.3 mm (xX = 3.2 mm, n = 3). Frons at vertex about 1.5-2.0 times broader than at narrowest point, about % width of head, with decumbent and erect, sparse, mixed silver and brown pile. Clypeus about as long as wide, with sparse silvery pile. Occiput with silvery pile reaching pos- terior margin of eye; postocular setae black. Antenna with fine silver pubescence; first flagellomere longest; pedicel and scape light brown; flagellum brown. Mandible with 41- 45 serrations. Lacinia with 31-32 retrorse teeth. Palpus dark brown, with stout, pale golden setae; palpomere V 1.7—2.0 times as long as III. Sensory vesicle elongate, located posteriorly to subcentrally, occupying about ’» of palpomere III; neck short, arising near anterodorsal margin, opening to exterior through rounded, slightly expanded mouth. Median proximal space of cibarium broadly U-shaped, lacking armature. Postpronotum and proepisternum brown, VOLUME 92, NUMBER 3 with long silvery pile. Scutum dark brown, humeral angles light brown; pile recumbent, golden centrally, silvery peripherally. Scu- tellum dark brown, with long, very pale golden pile mixed with black setae. Post- notum dark brown. Anepisternum and kat- episternum dark brown; katepisternum with a small, ventral patch of silvery pile (often rubbed off); membrane and mesepimeron brown; mesepimeral tuft of long, silvery se- tae. Wing veins pale yellowish brown. Setae on stem vein and costal base dark brown, with bronze reflections; setae on other veins primarily brown; subcosta setose ventrally; fringes of calypter and alar lobe silvery. Hal- ter tan, with line of pale golden pile. Coxae and tarsi dark brown; femora and tibiae brown; pile on legs silvery to pale golden; hind basitarsus 6.5-7.3 times as long as broad; calcipala and pedisulcus well devel- oped; claws each with large, thumb-like lobe. Abdominal sclerites dark brown; pile sparse, silvery; additional sparse, long, black setae on terminal tergites; membranous areas gray to brown, with silvery pile. Basal fringe of long, silvery to very pale golden pile. Anal lobe subquadrate in lateral view, rounded anteriorly, with acute posterodor- sal extension. Cercus a broadly rounded tri- angle, about 1.2—1.8 times as broad as long. Hypogynial lobes subtriangular, with space between lobes forming a narrow rectangle. Genital fork (Fig. 4) with stem moderately long and slender; lateral arms rather broad basally, forming suboval space in region of bifurcation; posteromedial areas of lateral arms well developed, and with bluntly acute angles; anteromedial area produced ante- riorly. Spermatheca over 1.5 times as long as broad, with superficial pattern of sub- equal polygons. Male. — General body color velvety black, with gray pruinosity and golden pile. Length: body 3.0-3.3 mm (x = 3.1 mm); wing, 2.7— 2.9 mm (x = 2.7 mm). Frons and clypeus with erect, brown pile. Occiput with long, erect, brown pile. An- tenna dark brown, with fine, light brown pile. Palpus dark brown, with brown pile; 435 palpomere V about twice as long as pal- pomere III. Sensory vesicle subspherical, about length of its segment, located pos- teriorly; neck rather short, slender, opening to exterior through small, rounded mouth. Postpronotum and proepisternum brown, with silvery to pale golden pile. Scutum vel- vety black, with recumbent, golden pile. Scutellum dark brown, with bronze pile. Postnotum dark brown. Anepisternum, kat- episternum, membrane, and mesepimeron dark brown, latter two sometimes paler: katepisternum bare; mesepimeral tuft of long, brown pile with bronze reflections. Wing veins pale yellowish brown. Setae on stem vein and costal base dark brown; setae on other veins brown; fringes of calypter and alar lobe brown with bronze reflections. Halter dark brown basally, paler distally, with bronze pile. Legs dark brown, with midsections of femora and tibiae paler; pile brown and golden brown, sometimes sil- very on forecoxa. Hind basitarsus 5.3-6.0 times as long as broad. Abdominal tergites velvety black, paler along posterior margins, with bronze pile; membranous areas gray, with bronze pile; sternites dark brown, with bronze pile. Bas- al fringe of very long, bronze pile. Termi- nalia as in Fig. 5. Gonocoxite about as long as broad. Gonostylus about as long as gon- ocoxite, about 2.9 times as long as breadth at midpoint, expanded apically into flat- tened, subtriangular, medially directed flange bearing | apical spine. Ventral plate in ventral view subrectangular, about twice as broad as long, slightly narrowing poste- riorly, with posterolateral corners well rounded, and posterior margin medially produced as a small, hirsute tubercle; an- terior margin with slight, medial concavity; arms directed slightly outward, with apices bowed slightly inward; lip in terminal view pronounced, broadly rounded, serrate lat- erally; median sclerite long, slender, forked for about '4—'5 its length; dorsal plate (Fig. 6) well sclerotized, with broad collar-like base, suborbicular distally; paramere in lat- eral view moderately narrow basally, broad- 436 ening medially, and bearing | long, slender, strongly sclerotized spine-like process. Chromosomes (from larval salivary glands; inversions are relative to the S. ver- num standard of Brockhouse [1985], Hunt- er and Connolly [1986]; 20 preparations ex- amined by Hunter and Connolly [1986] from Pennsylvania, 20 preparations examined from New Hampshire).—n = 3; chromo- center present; B chromosomes lacking; IS with inversions JS-/, [S-4, and /S-5; IL with inversions /L-2, IL-3, IL-4, IL-6, and IL-7, and with secondary nucleolar organiz- er (section 41C-42B) generally expressed; IIS standard for vernum sequence; IIL com- plexly rearranged (Fig. 18 in Hunter and Connolly 1986); IIIS with inversion ///S-2; IIIL with inversions ///L-4, IIIL-5, IITL-6, and //IL-8; sex chromosomes differentiat- ed as Y, = IJIL-1 sp, X, = IIIL standard; floating inversions in all arms except IIIS. Types. — Holotype: ¢ (pinned) with pupal and larval exuviae (in glycerin), outlet, Two- Towns Pond, Dixville Notch (The Bal- sams), Coos County, New Hampshire (44°52'N, 71°18'W), 24 May 1988, collected by P. H. Adler. Paratypes: NEW HAMP- SHIRE: COOS COUNTY: same data as ho- lotype, 77 larvae (including 40 mature), 10 chromosome preparations (8 female larvae, 2 male larvae) with photographic negatives, 15 pupae, | pupal exuviae, 4 4 (pinned) with larval and pupal exuviae (in glycerin), | 4 (cleared, in glycerin), 3 2 (pinned) with lar- val and pupal exuviae (in glycerin); same data as holotype, 19 May 1987, 2 larvae, J. F. Burger; same data as holotype, 26 May 1987, 8 larvae, 4 chromosome preparations (3 female larvae, | male larva) with pho- tographic negatives, J. F. Burger; outlet, beaver pond no. 5, Dixville Notch (The Bal- sams), 20 May 1987, | larva, 1 chromosome preparation (female larva) with photo- graphic negatives, J. F. Burger; PENNSYL- VANIA: MONROE COUNTY: outlet, White Heron Lake, Rt. 402, 1.6 km north PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Marshalls Creek (town), 7 May 1983, 4 larvae, | pupa, P. H. and C. R. L. Adler. Additional specimens examined.— PENNSYLVANIA: MONROE COUNTY: pond outlet, Pocono Highland Camp, Rt. 402, 4.8 km north of Marshalls Creek (town), 7 May 1981, 1 pupa, G. E. Jones; NEW HAMPSHIRE: COOS COUNTY: outlet, Round Pond, Dixville Notch (The Bal- sams), 9 June 1988, 4 larvae, J. F. Burger. An additional 83 males (68 pinned with exuviae in glycerin, 15 in alcohol with exu- viae) and 43 females (34 pinned with exu- viae in glycerin, 9 in alcohol with exuviae) were examined from the type locality (same data). Based on the configuration of the pu- pal gill, these specimens appear to be S. fionae. However, because pupal characters exhibit overlap between S. fionae and S. croxtoni and because the larval cuticle did not remain associated, I labeled the speci- mens as Simulium croxtoni/fionae. Etymology.—This species is named in honor of Fiona F. Hunter who originally resolved the chromosomes of this species, and who has contributed significantly to a cytological understanding of the S. vernum group. Diagnosis.—The larva of S. fionae is eas- ily distinguished by the dark, multiply branched setae covering the posterior 4 or 5 abdominal segments. Larvae of the closely related and morphologically similar S. crox- toni Nicholson and Mickel possess dark, simple abdominal setae and generally have a longitudinal, pale brown stripe on the frontoclypeal apotome. Larvae of S. fur- culatum Shewell have simple setae and some scattered, multiply branched (mainly bifid) setae. The pupal gill of S. fionae is generally darker and more splayed than in S. croxtoni and S. furculatum, and the single filament of the ventral petiole typically arises lateral to the doublet, whereas in SS. croxtoni the doublet often arises lateral to the single fil- ament. The anterodorsal projection of the VOLUME 92, NUMBER 3 cocoon is often shorter than in S. croxtoni. No reliable characters have been found to distinguish females of S. fionae and S. crox- toni. In S. fionae, the ventral plate of the male is slightly broader and less tapered than in S. croxtoni. Chromosomally, S. fionae differs from the S. vernum standard by approximately 23 fixed inversions, and from all other known species by at least one fixed inversion in every arm except IIS (Hunter and Connolly 1986). It is most readily distinguished by a chromocenter in combination with a sub- terminal Z marker (section 32A—32B) and a reoriented blister group (sections 75C- 76A), relative to standard. Populations in New Hampshire and Pennsylvania both carry IL-1 sp, HL-1 sp, IIL-2 sp, and the simple Y-linked inversion IIIL-1 sp. Larvae from New Hampshire sporadically carry a small inversion in the middle of IS, two independent inversions in IIS (approximate limits 52-54 inclusive and 50-52C inclu- sive), and a subterminal inversion in IIL (approximate limits 71B-72C2 inclusive). Additionally, the majority (85%, m = 20) of New Hampshire larvae carry at least one of five inversions in IL. Biology.—Simulium fionae was collected from the outlets of two man-made im- poundments in Pennsylvania and from the outlets of two beaver ponds and one man- made impoundment in New Hampshire. In Pennsylvania, larvae and pupae were col- lected in early May from old cattail (Typha) leaves and stalks trailing in the water. In New Hampshire, immatures were found from mid-May to early June almost exclu- sively on the undersurfaces of stones and sticks. At all sites, immatures were found with Cnephia dacotensis (Dyar and Shan- non), and were restricted to within 20 m of the outlet. Stream temperatures at the time of collection ranged from 16.5 to 17°C, and stream widths ranged from 0.6 to 1.2 m. Nearly 12% of the larvae collected at the Two Towns Pond outlet were infected with 437 mermithid nematodes; one larva was in- fected with the fungus Coelomycidium si- mulii Debaisieux while another was infect- ed with an unidentified microsporidium. Simulium claricentrum Adler, New SPECIES Figs. 10-17 Simulium pictipes “A” Bedo, 1973: 12, chromosomes. Simulium “species A” Reisen, 1974a: 19, ecology. (Hagenomyia) “species A” Reisen, 1974a: 72, larval biology. Simulium “species A” Reisen, 1974b: 275, larval biology. Simulium (Shewellomyia) “species A” Rei- sen, 1975a: 949, larval biology. Simulium “sp. a.” Reisen, 1975b: 27, larval biology. Simulium pictipes “A” Bedo, 1975: 1150, chromosomes. Simulium (Shewellomia) [sic] “species A” Reisen, 1977: 325, larval ecology. Simulium (Shewellomyia) pictipes ““cyto- species A” Adler and Kim, 1986: 36, lar- va. Larva (final instar).— Length 7.2-9.9 mm (x = 8.5 mm, n = 47). Head capsule (Fig. 10) whitish yellow to brown with pale areas centrally and anteriorly on frontoclypeal apotome; pigmentation beneath cuticle often visible as dark reticulate pattern; headspots indistinct, or with anterolateral, postero- lateral, and/or posteromedian headspots brown, fairly distinct; eye spots rather large; line over eye spots thin, brown. Antenna with distal article brown, median article pale brown (hyaline band visible subapically in darker specimens), proximal article pale brown; apex of distal article reaching end of labral-fan stalk; proportions of articles (distal to proximal, excluding apical sensil- lum) approximately 1.0:1.7:1.0. Labral fan with 47-60 (x = 54, n = 45) primary rays. Hypostomal teeth (Fig. 11) with median tooth and lateral teeth relatively large; me- 438 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON JN 13 Figs. 10-14. Simulium claricentrum new species. 10, Larval head capsule (dorsal view). 11, Larval hypos- toma. 12, Larval head capsule (ventral view). 13, Female genital fork (sternite 9). 14, Male terminalia (ventral view with left gonocoxite, gonostylus, and parameres removed). 0.1mm dian tooth longest; outermost sublateral and 3-4 lateral serrations per side; hypos- teeth shorter than lateral teeth, but longer toma with 6-7 prominent setae and 1-2 than innermost sublateral teeth; median small lateral setae per side. Postgenal cleft sublateral teeth shortest; lateral margin of (Fig. 12) about as long as basal width, ex- hypostoma with 0-2 small paralateral teeth tending about '2 distance to hypostomal VOLUME 92, NUMBER 3 Figs. 15-17. groove, widest at base, narrowing anteriorly to rounded point; subesophageal ganglion darkly pigmented. Maxillary palpus 3.2—4.0 times as long as basal width. Inner subapical ridge of mandible with single sensillum proximal to | large, subtriangular serration. Lateral plate of thoracic proleg moderately well sclerotized, about as long as wide, ex- tending almost length of apical article. Body (Fig. 15) gradually expanding posteriorly, dark gray and white, piebald, lacking con- spicuous cuticular setae; pigmentation heaviest on dorsum of segments 5-7 (often forming a triangle with apex pointing an- teriorly), and on thorax; gonadal sheath darkly pigmented in male (often visible through integument), unpigmented or pig- mented posteriorly in female. Anterodorsal arms of anal sclerite broadly connected to, 439 Simulium claricentrum new species. 15, Larval habitus (dorsal view). 16, Scanning electron micrograph of base of pupal gill. 17, Scanning electron micrograph of granules on portion of pupal thorax (dorsal view). broader than, and about 3 shorter than pos- teroventral arms. Rectal setulae short, sparse, visible only under phase contrast. Posterior proleg bearing | 8—22 hooks in 92- 100 rows. Anal papillae of 3 compound lobes. Pupa.— Length 3.3-4.5 mm (X = 3.9 mm, n = 15). Head projecting downward, with many minute granules as on thorax; anten- nal sheath of female extending nearly to posterior margin of head; antennal sheath of male extending about ' distance to pos- terior margin of head. Gill short, about length of pupa, consisting of 9 filaments; base extremely short, immediately giving rise to 4 short-petiolate pairs of filaments plus a single filament that curves ventro- laterally and often wraps around head; fil- aments moderately thin, tapering, weakly 440 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON annulate; surface sculpture of base weakly differentiated (Fig. 16). Thorax with nu- merous, irregularly spaced, minute, round- ed granules (Fig. 17); trichomes simple, moderately long, slender, rather dark, 7 on each side of thorax. Tergite I with | pair of setae; tergite II with 4 stout, anteriorly di- rected setae and 3 smaller setae on each side of midline, plus 3 setae laterally per side; tergites III and IV each with 4 anteriorly directed hooks on posterior margin on either side of midline, 2 small setae between and anterior to 2 outermost hooks, and 3-4 small setae laterally per side; tergites V to VII bare or with | minute seta each; tergite VIII with 3-6 hook-like setae along anterior margin on each side of midline, plus 2 minute setae per side and numerous comb-like micro- spines laterally; tergite [IX with pair of very short, stout, slightly curving, dorsally di- rected terminal spines, and numerous comb- like microspines. Pleural membrane of seg- ments II to VII with 0-1 minute setae per side. Sternite III with pair of fine setae; ster- nite IV with 2-3 heavy, anteriorly directed setae and 1-2 fine setae per side; sternite V with pair of stout, anteriorly directed setae and 2 fine setae per side; sternites VI and VII with pair of long, anteriorly directed, hook-like, simple or bifid setae and 1-2 fine setae per side; sternites VIII and IX gen- erally lacking setae; sternites III to VII with numerous rows of extremely fine, comb-like microspines. Cocoon boot-shaped, cover- ing entire pupa and gill, very coarsely woven (especially anteriorly). Female.—Generally grayish pruinose, with black markings on thorax and abdo- men; pile silver, more golden centrally on thorax. Length: body, 2.5-2.9 mm (x = 2.8 mm, ” = 7); wing, 3.3-3.7 mm (X = 3.4 mm, 7” = 7). Frons gray, at vertex about 1.5 times broader than at narrowest point, about 4 width of head, with decumbent, silver pile. Clypeus gray, slightly wider than long, with silver pile. Occiput with long, silver pile reaching posterior margin of eye; postocular setae black. Antenna with fine silver pu- bescence; first flagellomere longest, slightly longer than pedicel; scape and pedicel yel- lowish brown; flagellum dark brown. Man- dible with 33-35 serrations. Lacinia with 26-27 retrorse teeth. Palpus dark brown, with stout, brown setae; palpomere V 1.9- 2.3 times as long as III. Sensory vesicle lo- cated posteriorly, occupying about |” of pal- pomere III, opening directly to exterior through rounded, expanded mouth (neck extremely short or lacking). Median proxi- mal space of cibarium broadly U-shaped, lacking armature. Postpronotum and proepisternum gray, with long silver pile. Scutum gray to grayish black, with 3 fine, black vittae running lon- gitudinally, and with black patch antero- dorsal to wing base; pile recumbent, silver, becoming golden centrally; humeral angle yellowish gray. Scutellum dark brown to grayish black, with long, mixed silver and brown pile. Postnotum dark brown. Anepi- sternum and katepisternum dark brown, with gray pruinosity; membrane and mes- epimeron slightly paler; mesepimeral tuft of long, silver setae. Wing veins pale yellowish brown. Setae on stem vein and costal base mixed brown and silver; setae on other veins brown; subcosta setose ventrally; fringes of calypter and alar lobe silver. Halter tan to pale yellow, with line of fine, pale golden pile. Legs brown, often with paler patches, especially on hind basitarsus; pile primarily silver, brown on tarsi; hind basitarsus 6.4— 7.7 times as long as greatest width; calcipala small; pedisulcus deep; claws simple, mod- erately curved. Abdomen gray, with segments 3-5 black dorsally and segments 3—7 (sometimes 2) with small, partially shiny, brown patches laterally; sclerites of segments 6-9 dark brown, with gray pruinosity, and sparse, sil- ver pile; sternite 7 with long, brown setae along posterior margin. Basal fringe of very long, silver pile. Anal lobe brown, shiny, subrectangular in lateral view, with ventral margin rounded, and with dorsal finger-like VOLUME 92, NUMBER 3 extension nearly as long as body of lobe. Cercus subrectangular, about twice as broad as long, posterior margin straight, corners well rounded. Hypogynial valve a short truncate lobe, space between lobes rather broad, subrectangular. Genital fork (Fig. 13) with stem moderately long, slender, ex- panded at anterior end; lateral arms narrow, forming suboval to subtriangular space be- tween them, posterior areas expanded into subrectangular plates. Spermatheca rather small, subspherical, with cuticular micro- spines. Male.—Generally velvety black, with golden and brown pile. Length: body, 2.7- 3.5 mm (x = 3.0 mm, n = 5); wing, 2.9- 3.1 mm (X = 3.0 mm, n = 7). Frons and clypeus with golden-brown pile. Occiput with long, erect, brown pile. An- tenna brown, with fine, pale golden pubes- cence; pedicel pale brown. Palpus brown, with brown pile; palpomere V 2.5—2.7 times as long as palpomere III. Sensory vesicle oblong, about '4 length of its segment; neck short, opening to exterior through rounded mouth. Postpronotum and proepisternum brown, with long, silver pile. Scutum velvety black, with pair of silvery patches extending from humeral angle posteromedially, and with silvery patch posteriorly, humeral angles pale brown; pile recumbent, golden. Scu- tellum dark brown, with mixed brown and golden pile. Postnotum dark brown. Anepi- sternum, katepisternum, and membrane dark brown; mesepimeron dark brown, pal- er centrally; mesepimeral tuft of silver or pale golden pile. Wing veins pale gray to yellowish brown. Setae on stem vein and costal base dark brown; setae on other veins brown; fringes of calypter and alar lobe brown. Halter dark brown basally, tan to yellow distally, with fine, brown pile. Legs brown, with paler patches, especially on hind basitarsus; pile brown to golden brown. Hind basitarsus 4.9-5.0 times as long as broad; calcipala small; pedisulcus deep. Abdominal tergites velvety black, with 441 reflective, silver patches laterally on seg- ments 2, 6, and 7, and minimally on 8; pile brown to golden brown; membranous areas grayish, with long, brown and golden brown pile; sternites brown, with brown pile. Basal fringe of long brown to golden brown pile. Terminalia as in Fig. 14. Gonocoxite about 1.5 times longer than broad, with ventro- lateral angle produced posteriorly. Gono- stylus narrow, rounded apically, about as long as gonocoxite, 3.6-3.7 times as long as basal breadth, lacking apical spinule. Ven- tral plate finely setose, in ventral view deep- ly and broadly incised posteromedially, about 1.5 times broader than long, with lat- eral margins somewhat parallel and anterior margin smoothly convex; arms short, with apices curving inward; lip in terminal view with three, subequally prominent lobes; me- dian sclerite elongate, triangular; paramere in lateral view subquadrate basally, narrow- ing distally, and bearing numerous, variably long, spine-like processes. Chromosomes (from larval salivary glands: Bedo 1975).—n = 3; chromocenter absent; pseudochromocenter (ectopic pair- ing) occasional; B chromosomes lacking; centromere regions of all chromosomes ex- panded, each bearing sharp centromere band (that of chromosome III included in heavily staining heterochromatic area of expanded region); all chromosome arms standard for S. pictipes group; IIL as sex chromosome, with females homozygous for heavy band at 63A4 and males heterozygous for heavy and thin band at 63A4; floating inversions lacking. Types. — Holotype: 4 (pinned) with pupal exuviae and larval head capsule (in glyc- erin), Sixteenmile Creek, junction of Wash- ington Street and Shadduck Road, North- east (town), Erie Co., Pennsylvania (42°11'N, 79°50’W), 10 August 1988, col- lected by P. H. Adler and C. R. L. Adler. Paratypes: same data as holotype, 174 lar- vae (including 40 mature), 10 chromosome preparations (6 female larvae, 4 male lar- vae) with photographic negatives, 36 pupae, 442 91 pupal exuviae, 24 4 (pinned) with exu- viae (in glycerin), 2 6 (cleared, in glycerin), 2 6 (in alcohol), 25 2 (pinned) with exuviae (in glycerin); same data as holotype, 5 Au- gust 1986, 36 larvae, E. C. Masteller; AR- KANSAS: MARION COUNTY: same data as holotype, 12 August 1989, 36 larvae (in- cluding 20 mature), 42 pupae. Georges Creek, junction of Rt. 62, 6 km west of Yell- ville, 24 April 1989, 9 larvae (including 1 mature), 5 pupal exuviae, P. H. Adler and @aReL Adler: Additional specimens examined.— PENNSYLVANIA: ERIE COUNTY: Fourmile Creek, Wesleyville, near Behrend Campus, 12 May 1983, 1 chromosome preparation (female larva), E. C. Masteller. Etymology.—The specific name is de- rived from the Latin c/ari meaning clear, and centrum meaning center, in reference to the clearly defined centromere bands that readily distinguish this species from closely related taxa. Diagnosis.—Larvae (later instars) of S. claricentrum can be distinguished from those of other members of the S. pictipes group by the piebald pigmentation of the body and the paler head capsule; earlier instars are more uniformly dark. The nine-filamented gill of the pupa lacks the tuberculate surface sculpture on the basal 4 to '3 of each fila- ment, unlike other members of the S. pic- tipes group. The female cannot be reliably distinguished from that of S. pictipes. The ventral plate of the male is about 1.5 times as broad as long, with somewhat parallel lateral margins; in S. pictipes Hagen the ventral plate has divergent lateral margins, and in S. longistylatum Shewell it is nearly as broad as it is long. Chromosomally, the species is most easily identified by the pres- ence of well-defined centromere bands (Bedo 1975). Biology.—Like other members of the S. pictipes group, the immatures of S. clari- centrum are often found in moss-like clumps in swift watercourses. In the Lake Erie drainage basin of Pennsylvania, S. claricen- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON trum was found where water spilled over small shale-bottomed waterfalls, but in one case was found on a coarse cement sluice- way. These streams ranged in width from 10 to 20 m and in temperature from 16 to 27°C. In Oklahoma, S. claricentrum was collected from limestone streams with smooth, travertine substrates (Reisen 1974a, 1975a); larval abundance was highest dur- ing the spring months and was positively correlated with periphyton abundance (Rei- sen 1977). The Oklahoma streams ranged in temperature from 17 to 29°C, in pH from 7.7 to 8.2, in alkalinity from 235 to 250 ppm, and in dissolved oxygen from 7.5 to 8.5 ppm (Reisen 1974b, 1975a). In Arkan- sas, Iimmatures were taken from a calcar- eous siltstone-bottomed stream approxi- mately 10 m wide, with a temperature of 23°C. Simulium claricentrum is multivol- tine, immatures having been collected from early May through August in Pennsylvania and year round in Missouri and Oklahoma (Bedo 1975, Reisen 1974a, 1975a, b). The species passes through six larval instars (Re- isen 1975a). In 1989, I found a mixed pop- ulation of S. pictipes and S. claricentrum in Sixteenmile Creek, Pennsylvania. About 5% of S. claricentrum larvae in Sixteenmile Creek (1989) carried patent infections of the microsporidium Polydispyrenia simulii (Lutz and Splendore), formerly known as Pleistophora multispora (Strickland) (Can- ning and Hazard 1982). Reisen (1974a) recorded mating swarms at the base of rifles; coupled pairs imme- diately dropped to the ground where cop- ulation lasted only a matter of seconds. Fe- males oviposited from 200 to 400 eggs in dense masses during the early evening hours on rocks, grasses, algae, and mosses splashed with water (Reisen 1974a). ACKNOWLEDGMENTS J. F. Burger provided collections of S. fionae from northern New Hampshire, and facilitated my efforts to collect this species. E. C. Masteller provided the first Pennsyl- VOLUME 92, NUMBER 3 vania collection of S. claricentrum, as well as several subsequent larval collections. F. F. Hunter scored three slides of S. fionae for polymorphisms. A. H. Undeen identi- fied the microsporidia. C. I. Dial and J. P. Norton rendered the illustrations, and J. R. Brushwein and J. D. Culin photographed the larvae. C. R. L. Adler and J. C. Morse reviewed the manuscript. To all of these individuals I extend my thanks. This is Technical Contribution No. 2975 of the South Carolina Agricultural Experiment Station, Clemson University. LITERATURE CITED Adler, P. H. 1983. Ecosystematics of the Simulium vittatum complex (Diptera: Simuliidae) and as- sociated black flies. Ph.D. thesis. The Pennsyl- vania State Univ., University Park, Pennsylvania. 307 pp. 1987. A new North American species in the Simulium vernum group (Diptera: Simuliidae) and analysis of its polytene chromosomes. Proc. Ento- mol. Soc. Wash. 89: 673-681. Adler, P. H. and K. C. Kim. 1986. The black flies of Pennsylvania (Simuliidae, Diptera): Bionomics, taxonomy and distribution. Penn. State Agric. Exp. Stn. Bull. 856. 88 pp. Bedo, D. G. 1973. Sibling species and sex chromo- somes in black flies of the Simulium pictipes group (Diptera: Simulidae). M.S. thesis. Univ. Toronto, Ontario. . 1975. Polytene chromosomes of three species of blackflies in the Simulium pictipes group (Dip- tera: Simuliidae). Can. J. Zool. 53: 1147-1164. Brockhouse, C. L. 1985. Sibling species and sex chro- mosomes in Eusimulium vernum (Diptera: Sim- uliidae). Can. J. Zool. 63: 2145-2161. Canning, E. V. and E. I. Hazard. 1982. Genus Pleis- tophora Gurley, 1893: An assemblage of at least three genera. J. Protozool. 29: 39-49. Craig, D. A. and R. E. G. Craig. 1986. Simuliidae 443 (Diptera: Culicomorpha) of Rarotonga, Cook Is- lands, South Pacific. N. Z. J. Zool. 13: 357-366. Crosskey, R. W. 1987. An annotated checklist of the world black flies (Diptera: Simulidae). 72 Kim, K. C. and R. W. Merritt, eds., Black Flies: Ecol- ogy, Population Management, and Annotated World List. The Pennsylvania State University, University Park. Hunter, F.F. 1987. A cytotaxonomic study of species in the Simulium vernum group (Diptera: Simuli- idae). M.S. thesis. University of Toronto, Ontario. 181 pp. Hunter, F. F. and V. Connolly. 1986. A cytotaxo- nomic investigation of seven species in the Eusi- mulium vernum group (Diptera: Simulidae). Can. J. Zool. 64: 296-311. Reisen, W. K. 1974a. The ecology of larval blackflies (Diptera: Simuliidae) in a south central Oklahoma stream. Ph.D. dissertation. Univ. Oklahoma. Nor- man, Oklahoma. 170 pp. 1974b. The ecology of Honey Creek: A pre- liminary evaluation of the influence of Simulium spp. (Diptera: Simuliidae) larval populations on the concentration of total suspended particles. Entomol. News 85: 275-278. . 1975a. Quantitative aspects of Simulium vir- gatum Coq. and S. species life history 1n a southern Oklahoma stream. Ann. Entomol. Soc. Am. 68: 949-954. 1975b. The ecology of Honey Creek, Okla- homa: Spatial and temporal distributions of the macroinvertebrates. Proc. Okla. Acad. Sci. 55: 25- EVE 1977. The ecology of Honey Creek, Okla- homa: Population dynamics and drifting behavior of three species of Simu/ium (Diptera: Simuliidae). Can. J. Zool. 55: 325-337. Shewell, G. 1952. New Canadian black flies (Diptera: Simuliidae). I. Can. Entomol. 84: 33-42. 1959. New Canadian black flies (Diptera: Simuliidae). I]. Can. Entomol. 91: 83-87. Vavra, J. and A. H. Undeen. 1981. Microsporidia (Microspora: Microsporida) from Newfoundland blackflies (Diptera: Simuliidae). Can. J. Zool. 59: 1431-1446. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 444-460 NEW SPECIES AND RECORDS OF PREDACEOUS MIDGES FROM FIJI (DIPTERA: CERATOPOGONIDAE) WILLIS W. WIRTH AND FRANCIS E. GILES (WWW) Systematic Entomology Laboratory, USDA, and Florida State Collection of Arthropods, 1304 NW 94th St., Gainesville, Florida 32606; (FEG) Professor of Biology, Loyola College in Maryland, 4501 N. Charles St., Baltimore, Maryland 21210. Abstract.—Fourteen species of predaceous midges of the tribes Ceratopogonini, Het- eromyiini, Sphaeromiini, and Palpomyiini are reported from Fiji, including eight species described as new: Downeshelea stenochora, Monohelea beaveri, M. fijiensis, M. coloisuvae, M. leveri, Stilobezzia browni, Nilobezzia fijiensis, and Bezzia vitilevuensis. All are new records from Fiji, from which only four species of Forcipomyiinae and two species of Culicoidini had previously been reported. Key Words: The biting and predaceous midges of the family Ceratopogonidae are a conspicuous element of the Dipterous fauna of the Pa- cific islands. Tokunaga and Murachi (1959) recorded 147 species from Micronesia. For unknown reasons practically no collecting or taxonomic study has been done on the ceratopogonids of Fiji. Debenham (1978) made a careful listing of all the species re- corded from the Australasian Region and recorded only six species from Fiji, as fol- lows: Atrichopogon jacobsoni (de Meijere), Forcipomyia fijiensis (Macfie), F. fuliginosa (Meigen), F. indecora Kieffer, Culicoides belkini Wirth and Arnaud, and C. cancri- socius Macfie. In this paper we report on the predaceous species of the tribes Ceratopogonini, Het- eromylini, Sphaeromiini and Palpomyiini. The species of the subfamilies Forcipomyi- inae and Dasyheleinae, and the bloodsuck- ing species of the tribe Culicoidini of the subfamily Ceratopogoninae will be reported on in a second paper. We here record 14 species, of which eight are new species, and all are new Fijian records. predaceous midges, Ceratopogonidae, Fiji Taxonomic characters employed for identification of adult ceratopogonids were described by Wirth et al. (1977), Downes and Wirth (1981), and particularly for the tribe Ceratopogonini by Wirth and Grogan (1988). Wing length is measured from the basal arculus to the wing tip and costal length from the basal arculus to the costal apex. Costal ratio is the costal length divided by the wing length. Antennal ratio of the fe- male is the sum of the lengths of the elon- gated five distal flagellar segments divided by the sum of the lengths of the preceding eight; in the male the antennal ratio is the sum of the lengths of the elongated three distal segments divided by the sum of the lengths of the preceding 10 short segments. Palpal ratio is the length of the third palpal segment divided by its greatest breadth. Tarsal ratio is the value obtained by divid- ing the length of the basitarsus by the length of the second tarsomere. The holotypes and allotypes of the new species herein described are deposited in the U.S. National Museum of Natural History in Washington, D.C. (abbreviated USNM). VOLUME 92, NUMBER 3 Paratypes as available will be deposited in the British Museum (Nat. Hist.), London; the Museum National d’Histoire Naturelle in Paris; the Bishop Museum in Honolulu (BISH); the Australian National Insect Col- lection in Canberra; and the DSIR National Insect Collection in Auckland, New Zea- land. Acknowledgments.—For the loan of ex- tensive Fijian material we are greatly in- debted to Neal Evenhuis, G. A. Samuelson, and the late J. Linsley Gressitt and the Trustees of the Bishop Museum in Hono- lulu. Richard L. Brown of the Mississippi Entomological Museum, Mississippi State University, in Starkville, furnished us ma- terial that he collected in Fiji in 1986. We are also grateful to Noel L. H. Krauss of Honolulu for his continued interest and co- operation in collecting ceratopogonids from the Pacific islands including Fiji. Masaaki Tokunaga of Kyoto, Japan, very unselfishly turned over to us his unpublished notes and drawings on some Fijian ceratopogonids he received from the Bishop Museum. Karen Toohey and Mark Goettel of the University of Alberta in Edmonton sent us some valu- able reared ceratopogonid material from Fiji while they were stationed with the Vector Research Unit of the Fiji Office of Health. The senior author wishes to acknowledge with deep appreciation the support of the Institute of Natural Resources, R. J. Mor- rison, Director, University of the South Pa- cific, Suva, Fiji, for an appointment as Re- search Associate for a two-week period in November 1985. Roger A. Beaver, head of the Biology Department of the same Uni- versity, was very helpful in arranging ac- commodations and laboratory and field support, as well as specimens from his own collecting in Fiji. M. Kamath of the Fiji Ministry of Forests at Colo-i-Suva kindly assisted in transportation and assistance in the field collections. Howard Moore, Technical Assistant at Loyola College, gave invaluable aid in the photographic work. 445 Subfamily Ceratopogoninae Tribe Ceratopogonini Genus A//luaudomyia Kieffer References: Tokunaga, 1959 (New Guinea species); Tokunaga and Murachi, 1959 (Mi- cronesian species); Wirth and Delfinado, 1964 (revision Oriental species); Deben- ham, 1971 (revision of Australia and New Guinea species). Alluaudomyia bipunctata Tokunaga and Murachi Alluaudomyia bipunctata Tokunaga and Murachi, 1959: 356 (male, female; Car- oline Islands; fig. male wing, palpus, an- tenna, genitalia). Distribution. — Caroline Islands; Fiji. New record.—FIJI: Viti Levu, Savura Creek, v.1983, R. A. Beaver, Malaise trap, 2 females. Alluaudomyia tenuistylata Tokunaga Alluaudomyia tenuistvlata Tokunaga, 1959: 296 (male; West Irian); Tokunaga, 1963: 225 (male, female; West Irian, New Guinea); Debenham, 1971: 171 (male, fe- male redescribed; figs.; Queensland). Distribution.—Fiji, New Guinea, Queensland, West Irian. New record.—FIJI: Viti Levu, Savura Creek, v.1983, R. A. Beaver, Malaise trap, 2 females; 14 km w Lami, 7-10.xi1.1986, R. L. and B. B. Brown, UV light trap, 1 male, 2 females. Genus Downeshelea Wirth and Grogan Downeshelea Wirth and Grogan, 1988: 50. Type-species, Monohelea stonei Wirth, by original designation. References (to Downeshelea and Mono- helea): Tokunaga and Murachi, 1959: 404 (Micronesian species); Tokunaga, 1963: 238 (New Guinea species); Debenham, 1972: | (Australia and New Guinea species); Ra- tanaworabhan and Wirth, 1972: 439 (Ori- ental species). 446 Downeshelea stenochora Wirth and Giles, NEw SPECIES Figs. 1-7, 51 Female allotype.— Wing length 1.02 mm; breadth 0.39 mm. Head: Brownish, narrow bases of anten- nal segments 4-10 pale. Eyes contiguous in a point, bare. Antenna (Fig. 3) with lengths of flagellar segments in proportion of 18- 15-14-14-13-13-13-13-22-23-23-23-30; antennal ratio 1.17. Palpus (Fig. 4) dark brown, stout; lengths of segments in pro- portion of 4-8-14-9-13; palpal ratio 1.7. Mandible with 10 coarse teeth. Thorax: Dark brown, scutellum yellowish on each end. Legs (Fig. 1) brown, paler on fore leg, dark brown on hind leg; knees con- spicuously yellowish; hind femur and tibia slightly stouter. Hind leg with lengths from femur to tarsomere 5 as 88-78-50-20-10-9- 12; hind basitarsus (Fig. 2) with abrupt bend near base, with one dense row of short pal- isade setae and a sparse row of longer setae, one strong black ventral spine at base and two slender spines at apex; tarsomeres 2-3 each with pair of slender apical spines; tar- somere 4 subcylindrical, with pair of long, strong, black spines at apex; hind claw sin- gle, simple, length 0.068 mm. Fore tarsus with slender ventral spines as follows: tar- somere | with one at base and two at apex: tarsomeres 2-4, one apical. Mid basitarsus with two similar spines at base and eight along length of segment; segments 2-4 each with two apical. Fore and mid claws paired, subequal, curved; elongate, length 0.043 mm on fore leg and 0.036 mm on mid leg. Wing (Fig. 51) with conspicuous pattern due to enlarged dark gray microtrichia consisting of a broad fascia across midlength at level of first radial cell, and a small subapical fascia in cell R5 behind tip of costa, the two connected by a darkened area in cell M1. Costa unusually elongate, costal ratio 0.85; two radial cells, the first nearly half the length of second, rather broad, the second unusu- ally narrow, almost slitlike distally. Halter intensely dark brown. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Abdomen: Dark brown; pleural mem- brane with microscopic striations due to rows of dense blackish spicules. Sperma- thecae (Fig. 5) two, subspherical with well- developed stout necks; subequal, each 0.065 by 0.038 mm including the neck. Male holotype.— Wing length 0.98 mm; breadth 0.29 mm. Similar to the female with the usual sex- ual differences. Wing much narrower, costa shorter, CR 0.72. Antenna with well-de- veloped, pale brownish plume; lengths of flagellar segments in proportion of 28-10- 10-10-10-10-10-8-8-12-27-25-26, antennal ratio 0.87. Hind leg with lengths from femur to tarsomere 5 as 77-64-40-20-12-8-10. Claws short, equal and similar on all legs, each bent at base and straight distally. Genitalia (Fig. 7): Dark brown, strongly sclerotized, about as long as broad. Ninth sternum narrow, slightly broadened caudad in midportion, there with four strong setae; ninth tergum convex distally, caudal margin nearly straight, with a pair of small, bead- like, apicolateral processes, each with a mi- nute seta. Basistyle stout, about twice as long as broad, without lobes or armament; dis- tistyle 0.65 as long as basistyle, moderately stout distally, slightly curved. Aedeagus complex, typical of the genus; basally a broad median plate about half again as long as broad, with a pair of short, stout, antero- lateral arms, distolateral corners of the plate strongly sclerotized, flangelike; on the con- cave caudal margin of this plate between the flanges arise a pair of long slender pro- cesses nearly as long as the plate, with tips converging caudally on midline. Parameres (Fig. 6) joined on proximal third in a scler- otized plate with irregular outlines as fig- ured, posteriorly expanded in a pair of broad, crescent-shaped processes, each bearing at broadest point on lateral margin a curved blade directed ventromesally with apices nearly meeting on midline at about half the length of the plate. Distribution. — Fiji. Types. — Holotype male, allotype female, VOLUME 92, NUMBER 3 447 =e = Se, \ \) ; 0.10mm | Vv SS S 0.036mm ——— es As Figs. 1-13. Figs. 1-7, Downeshelea stenochora, 1-5, female; 6-7, male; Figs. 8-13, Monohelea beaveri, 8- 11, female; 12-13, male: 1, 11, femora and tibiae of (left to right) fore, mid and hind legs; 2, hind tarsus; 3, 9, antenna; 4,8, palpus; 5, 10, spermathecae; 6, 12, parameres; 7, 13, genitalia, parameres omitted. the Greek: stenos—narrow, and chora— space, referring to the unusually narrow ra- dial cells of this species. Discussion. —Six species of Downeshelea Viti Levu, Savura Creek, Colo-i-Suva, Fiji, iv.1983, R. A. Beaver, in Malaise trap. Paratype, 1 male, same data except v.1983. Etymology.—The specific name is from 448 are known from the Australasian Region (Wirth and Grogan 1988): /eei (Debenham 1972) from New South Wales, mcdanieli (Tokunaga, in Tokunaga and Murachi 1959) from the Caroline Islands, nigra (Tokunaga 1963) from New Guinea, sepikensis (De- benham 1972) from New Guinea, uwnimacu- lata (Debenham 1972) from New South Wales, and xanthogonua (Tokunaga 1963) from New Guinea. Of these species, D. stenochora most closely resembles xantho- gonua, with the same general structure of the male genitalia, especially the setae on the ninth sternum, shape of the apicolateal processes on the ninth tergum, and the gen- eral plan of the aedeagus and parameres. However, D. xanthogonua is readily distin- guished from stenochora by its more re- stricted wing pattern, the shorter and broad- er radial cells, and the different shapes of the aedeagus and the distal portions of the parameres. Genus Echinohelea Macfie Reference: Debenham, 1970: 145 (revi- sion, species of Australia and New Guinea; key). Echinohelea flava Tokunaga Echinohelea flava Tokunaga, 1963: 235 (fe- male; New Britain; figs.); Debenham, 1970: 151 (descriptive notes; male de- scribed and figured is not flava; New Guinea, Solomon Is.). Recorded Distribution.—New Britain, New Guinea, Solomon Islands. New Records.—FIJI: Viti Levu, Lami, 11.1981 (Krauss), 6 males, 7 females (BISH); Naussori Highlands, 500-700 m, xi.1976 (Krauss), | male, | female; Navai, 10.11.1971 (Krauss), | male; Colo-i-Suva, Savura Creek, v.1983 (Beaver), 1 male. Genus Monohelea Kieffer References: Tokunaga and Murachi, 1959 (Micronesian species); Tokunaga, 1959, 1963 (New Guinea species); Debenham, 1972 (revision Australia and New Guinea PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON species); Ratanaworabhan and Wirth, 1972 (revision Oriental species); Clastrier, 1985a (New Caledonia species); Wirth and Gro- gan, 1988 (diagnosis of genus and check list of species for world). Remarks.— Wirth and Grogan (1988) di- vided the traditionally recognized genus Monohelea into three genera, Monohelea Kieffer, A/lohelea Kieffer, and Downeshelea Wirth and Grogan, for which they gave di- agnoses and world lists of species. They re- stricted the genus Monohelea to two groups, the tigrina group, an Australasian group of clear-winged species, and the hieroglyphica group, a large and widespread group with a hieroglyphic type wing pattern and exten- sively banded legs. They listed 10 species of the hieroglyphica group from the Austral- asian Region; species of this grouyp are dif- ficult to separate except for details of the male genitalia. Adults of both sexes of the species occurring in Fiji, however, can be separated by the following key: KEY TO THE FIJIAN SPECIES OF MONOHELEA Hind femur pale except for subapical dark band 2 Hind femur dark at least on proximal half ... 3 Mid femur and tibia pale except apex of tibia coloisuvae new species — Maud femur and tibia brownish Be ee OL Oe fijiensis new species 3. Second radial cell of wing without infuscated area; costa short, costal ratio 0.79; hind femur brown on proximal 0.75, pale on distal 0.25; fore and mid femora and tibiae pale brown except broadly pale at knees . beaveri new species — Second radial cell with large infuscated area; costa long, costal ratio 0.89; hind femur brown except for two distal oblique narrow pale rings; fore and mid femora pale with subapical dark band, tibiae pale with subbasal dark band and apical|dark band <..-5..).- ... leveri new species Nl Monohelea beaveri Wirth and Giles, NEw SPECIES Figs. 8-13, 52 Female allotype.— Wing length 0.76 mm; breadth 0.30 mm. Head: Yellowish, antennae pale brown, palpi whitish. Eyes broadly separated, bare. VOLUME 92, NUMBER 3 Antenna (Fig. 9) with lengths of flagellar segments in proportion of 11-9-10-10-10- 11-11-12-13-13-14-15-20; antennal ratio 0.90. Palpus (Fig. 8) with lengths of seg- ments in proportion of 3-4-6-5-11; third segment with small round sensory pit. Man- dible with eight small teeth. Thorax: Pale brown; mesonotum with dark brown mottling seen in slide-mounted specimen; scutellum yellow. Legs (Fig. 11) pale yellowish with pale brownish bands as follows: proximal 0.6 of femora, all except bases of fore and mid tibiae, and a mod- erately broad subbasal band and distal fifth of hind tibia; bands on hind legs more in- tense. Hind leg with lengths from femur to tarsomere 5 as 114-103-50-24-16-18-13; hind basitarsus with abrupt bend near base and with row of palisade setae; tarsi without prominent ventral spines. Claws nearly straight, subequal on fore and mid legs, a single long claw on hind leg; lengths 0.043 mm on fore leg, 0.040 mm on mid leg, and 0.058 mm on hind leg. Wing (Fig. 52) with pattern typical of hieroglyphica species group; no dark spot in midportion of second radial cell; costal ratio 0.79; radial cells well formed, first 0.55 as long as second. Halter whitish with end of knob brown. Abdomen: Pale brown. Spermathecae (Fig. 10) two; greatly unequal in size and shape, the larger 0.062 mm by 0.046 mm including neck, ovoid with short thick neck; the small- er subspherical, 0.026 mm by 0.023 mm, without neck but with long, threadlike scler- otization of the duct. Male holotype.— Wing length 0.70 mm; breadth 0.30 mm. Similar to the female with the usual sex- ual differences. Wing slightly narrower; cos- ta shorter, costal ratio 0.70. Antenna with sparse brownish plume; segments 6-12 fused; antennal ratio 0.67. Legs with mark- ings as in female; hind leg with lengths from femur to tarsomere 5 as 95-86-48-34-16- 10-12; claws short, equal and similar on all legs, very slender and pointed and nearly straight. 449 Genitalia (Fig. 13): Dark brown; short, about as broad as long. Basistyle stout and tapering; dististyle nearly as long as basi- style, slender and curving gradually to tip. Aedeagus typical of hieroglyphica group, the well-sclerotized, triangular, lateral sclerites joined basally by a sclerotized loop, the me- dian dorsal membrane with bilobed caudal margin, the lobes sharp-pointed. Parameres (Fig. 12) with the winglike anterior lobes joined mesally by a narrow sclerotized bridge, the caudal portions well separated, each a slender, nearly straight column slightly expanded distally, especially in lat- eral view, and abruptly recurved on distal portion in a slender pointed process bent ventrally. Distribution. — Fiji. Types.— Holotype male, allotype female, Viti Levu, Colo-i-Suva, Savura Creek, v.1983, R. A. Beaver, Malaise trap. Para- types, 9 males, 9 females, as follows: VITI LEVU: Same data as types, 4 males, 6 fe- males; same but iv.1983, 4 males, 3 females. Naraiyama, 178°S’E, 17°56’S, 18- 30.xi.1986, R. L. Brown, UV light trap, 1 male. Discussion.—This species is dedicated with pleasure to R. A. Beaver of the Biology Department, University of the South Pacific in Suva, in appreciation of his kindness and assistance to the senior author during his visit to Fiji in 1985. Monohelea beaveri can be distinguished from the three other Fijian species of Monohelea by the characters presented in the key above. Monohelea coloisuvae Wirth and Giles, New SPECIES Figs. 14-20, 53 Female allotype.— Wing length 0.78 mm; breadth 0.33 mm. Head: Yellowish, antenna pale brown, palpi whitish. Eyes broadly separated, bare. Antenna (Fig. 15) with lengths of flagellar segments in proportion of | 2-10-12-13-14- 14-14-15-16-17-17-17-20; antennal ratio 450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Se Sy Da ae eC gE a a Ae b 18 0.16mm 0.017mm Figs. 14-25. Figs. 14-20, Monohelea coloisuvae, 14-18, female; 19-20, male; Figs. 21-25, M. fiyjiensis, male: 14, 21, femora and tibiae of (left to right) fore, mid and hind legs; 15, antenna; 16, spermathecae; 17, 23, palpus; 18, 22, hind tarsus; 19, 24, parameres; 20, 25, genitalia, parameres omitted. 0.83. Palpus (Fig. 17) with lengths of seg- Thorax: Yellowish with brownish mot- ments in proportion of 4-5-8-6-8; third seg- _ tling; scutellum yellow. Legs (Fig. 14) pale ment with inconspicuous small round sen- yellowish, coxae brownish. Hind leg with sory pit. Mandible with eight small teeth. _ three brown bands; femur with an oblique VOLUME 92, NUMBER 3 band just past midlength continued as a nar- row infuscation ventrally to tip of femur; tibia with narrow brown band just before midlength and a broad brown area covering distal fourth. Hind leg with lengths from femur to tarsomere 5 as 1 1 5-100-50-25-16- 13-13. Claws long and slender, slightly curving, subequal on fore and mid legs, a single long claw on hind leg (Fig. 18); lengths 0.044 mm on fore leg, 0.039 mm on mid leg, and 0.064 mm on hind leg. Very slender ventral spines on tarsi as follows: One sub- basally on mid and hind basitarsi; a pair at apices of fourth tarsomeres, and a pair at apices of tarsomeres 1-3 on mid leg. Wing (Fig. 53) with hieroglyphic pattern as in /. beaveri, CR 0.79. Halter whitish with end of knob brown. Abdomen: Pale brownish. Spermathecae (Fig. 16) two, greatly unequal in size and shape; the larger 0.067 by 0.049 mm in- cluding neck, ovoid, tapering to slender neck, with faint perforations in sclerotization on tapering portion near neck; the smaller 0.029 mm in diameter, spherical, without neck but with long, threadlike sclerotization of the duct. Male holotype.— Wing length 0.72 mm; breadth 0.30 mm. Similar to the female with the usual sex- ual differences; costal ratio 0.74. Antenna with sparse brownish plume; segments 6— 12 fused; distal three segments elongated, antennal ratio 0.90. Legs with markings as in female, femora slightly infuscated on fore and mid legs, ventral infuscation at tip of hind femur fainter; hind leg with lengths from femur to tarsomere 5 as 97-90-47-24- 14-11-12; claws short, equal and similar on all legs, very slender and pointed, and nearly straight. Genitalia (Fig. 20): Brown, about as broad as long. Ninth tergum convex distally with small, moderately separated, papilla-like apicolateral processes. Dististyle short, half as long as basistyle, moderately stout and tapering to blunt-pointed tip. Aedeagus with lateral sclerites rather slender distally, each 451 flanked by a slender hyaline blade subequal in length. Parameres (Fig. 19) joined in a narrow bridge near midportions, bases ex- panded winglike as usual in hieroglyphica group; distal portions each moderately slen- der and gradually tapering to ventrolaterally directed distal process, a small rounded ventromesal lobe at the base of the bent distal process. Distribution. — Fiji. Types.— Holotype male, Viti Levu, Na- raiyama, 28-30.xi.1986, R. L. Brown, UV light trap. Allotype female, Viti Levu, Colo- i-Suva, Savura Creek, v.1983, R. A. Beaver, Malaise trap. Paratypes, 2 females, same data as allotype; 2 males, 3 females, same data but iv.1983. Discussion.—The species takes its name from the locality near Suva on Viti Levu where the allotype and paratypes were col- lected. Monohelea coloisuvae can be distin- guished from the three other Fijian species of Monohelea by the characters presented in the key above. It is the Fijian species with the most restricted brownish leg markings, and the shapes of the distal portions of the parameres are diagnostic. Monohelea fijiensis Wirth and Giles, New SPECIES Figs. 21-25, 54 Male holotype.—Closely resembling the preceding species, Monohelea beaveri, but differing as follows: Wing length 0.69 mm; breadth 0.27. Head: Antenna with lengths of flagellar segments in proportion of 20-7-7-7-7-7-6- 5-5-6-20-20-23, antennal ratio 0.82. Palpus (Fig. 23) with lengths of segments in pro- portion of 6-9-17-10-16, third segment with small round sensory pit. Thorax: More extensively brownish, but scutellum almost entirely yellowish. Legs (Fig. 21) pale brownish, knees narrowly pale; hind femur and tibia pale with three broad dark bands, one subapically on femur, sec- ond sub-basally on tibia, and third at apex of tibia. Hing leg with lengths from femur 452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to tarsomere 5 as 92-80-453-23-15-12-12. Claws small, equal and simple on all legs (Fig. 22). Wing (Fig. 54) with dark markings as in beaveri, but not quite as extensive; costal ratio 0.67. Genitalia (Fig. 25): Shorter and broader than in beaveri; ninth tergum more convex distally, apicolateral processes smaller and rounded and set closer together near mid- line. Dististyle short and tapering, only two- thirds as long as basistyle. Aedeagus with lateral sclerites stouter, the basal loop not well developed. Parameres (Fig. 24) short and stout, joined in midportion at about half their length; each distal portion a broad, distally rounded plate, from the apex of which arises a moderately slender, tapering process of about the same length, curving ventrolaterad. Distribution. — Fiji. Types.— Holotype male, one male para- type, Viti Levu, Colo-i-Suva, Savura Creek, iv.1983, R. A. Beaver, Malaise trap. One male paratype, Naraiyama, Viti Levu, 28- 30.x1.1986, R. L. Brown, UV light trap. Discussion. — Monohelea fijiensis can be separated from its three Fijian congeners by the characters given in the key above. Monohelea leveri Wirth and Giles, NEw SPECIES Figs. 33-39, 55 Female allotype.— Wing length 0.70 mm; breadth 0.36 mm. Head: Pale brown, antennae darker, palpi paler. Eyes broadly separated, bare. Anten- na (Fig. 36) with lengths of flagellar seg- ments in proportion of 10-7-7-8-8-9-10- 10-12-12-13-15-20; antennal ratio 1.04. Palpus (Fig. 35) short and stubby; lengths of segments in proportion of 3-4-6-5-8; sen- sory pit on third segment small and round. Mandible with eight strong teeth. Thorax: Brown including scutellum. Legs (Fig. 33) brown; distal third of fore femur and proximal third of fore tibia pale, also a broad but incomplete subapical pale band on fore tibia; mid femur pale on mid third, mid tibia pale on mid half; hind femur and tibia dark brown except narrow pale bands in midportion. Hind leg with lengths from femur to tarsomere 5 as 96-82-43-20-13- 12-12; hind femur and tibia slightly thick- ened and hind femur slightly bowed. Mid and hind basitarsi with a strong spine near base and a pair of slender spines at apex (Fig. 34); a pair of slender spines at apices of tarsomeres 2—4 on mid leg. Claws nearly straight, subequal on fore and mid legs, a single long claw on hind leg; lengths 0.029 mm on fore leg, 0.026 mm on mid leg, and 0.046 mm on hind leg. Wing (Fig. 55) with dark hieroglyphic pattern much more ex- tensive than in other Fijian species, a prom- inent dark mark included in midportion of second radial cell; costa unusually long, cos- tal ratio 0.89. Halter brown. Abdomen: Pale brown. Spermathecae (Fig. 37) two, greatly unequal in size and shape, the larger 0.069 by 0.046 mm including neck, ovoid with short thick neck; the smaller spherical, 0.029 mm in diameter, without neck but with long, threadlike sclerotization of the duct. Male holotype.— Wing length 0.67 mm; breadth 0.27 mm. Similar to the female with the usual sex- ual differences; wing narrower and costa shorter as normal in the genus; costal ratio 0.80. Antenna with sparse, pale brownish plume, segments 6-12 fused as usual; an- tennal ratio 0.77. Legs with markings as in female; hind leg with lengths from femur to tarsomere 5 as 94-80-42-22-15-12-13; claws short, equal and similar on all legs, very slender and pointed and nearly straight. Genitalia (Fig. 39): Slightly broader than long, brownish. Ninth tergum covex distad, with short, papilla-like apicolateral pro- cesses. Dististyle stout and tapering to blunt point, about two-thirds as long as basistyle. Aedeagus with darkly sclerotized basal loop, the lateral sclerites slender distally, flanked laterally by a slender process similar to that of M. coloisuvae. Parameres (Fig. 38) short and broad, the lateral wings at base well VOLUME 92, NUMBER 3 453 0.090mm DS cas = | 0.031mm 34. OFS 0.036mm es, 0.18mm w Oo 0.15mm 38 Figs. 26-39. Figs. 26-32, Stilobezzia browni, 26-31, female; 32, male; Figs. 33-39, Monohelea leveri, 33- 37, female; 38-39, male: 26, 33, femora and tibiae of (left to right) fore, mid and hind legs; 28, 36, antenna; 29, 35, palpus; 30, 37, spermathecae; 31, abdominal terga; 32, genitalia; 38, parameres; 39, genitalia, parameres omitted. 454 developed; main stem portion of each un- usually short, tapering and abruptly bending ventrolaterally in a hooklike process; the stem portion continuing straight caudally in a prominent, rather slender, unequally forked, distal process twice as long as the hooklike ventrolateral process. Distribution. — Fiji. Types.— Holotype male, allotype female, Viti Levu, Colo-i-Suva, Savura Creek, iv.1983, R. A. Beaver, Malaise trap. Discussion.—The species is dedicated to the memory of R. J. A. W. Lever, for many years the Government Entomologist for the British Colony of Fiji, and the foremost au- thority on the economic insects of the south- western Pacific islands. The insect collec- tion at the Agriculture Station at Koronivia remains a well-curated collection of speci- mens, the great majority of which were col- lected by Lever. Monohelea leveri differs markedly from the three other known species of Fijian Mo- nohelea, distinguished by its stout legs, more brownish color with more extensively dark wing markings, more extensive brownish leg markings, and especially by the unusually long costa and second radial cell. The male parameres, with their short, curved, lateral process and long, straight, unequally-forked, distal process, are diagnostic. Monohelea palauensis Tokunaga (in Tokunaga and Mu- rachi 1959), a widespread Pacific species, 1s closely related, with stout legs, wing pattern with a dark mark in the second radial cell, and male genitalia ofa similar structure, but palauensis differs in its shorter costa (female costal ratio 0.79), paler legs, and different proportions on the distal processes of the male parameres. Genus Stilobezzia Kieffer References: Lee, 1948: 345 (key to Aus- tralia and New Zealand species); Tokunaga and Murachi, 1959: 363 (Micronesian species); Tokunaga, 1963: 249 (key to New Guinea species); Das Gupta and Wirth, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1968: 1 (revision of Oriental species; generic diagnosis). Stilobezzia bifurcata Tokunaga Stilobezzia bifurcata Tokunaga, 1959: 307 (male, female; New Guinea; figs.); To- kunaga, 1963: 271 (notes; New Guinea records); Debenham, 1978: 472 (Austra- lasian literature and distribution). Recorded distribution.—Irian Jaya, New Britain, New Guinea. New records.—FIJI: Viti Levu, Ovalau, Levuka, xi.1975, N. L. H. Krauss, | female (BISH); Korotongo, 11.1981 (Krauss), 1 male (BISH); Lami, 11.1981 (Krauss), 1 male; same, i,111.1971, 111.1976, 11.1977, x11.1978 (Krauss), | male, 7 females; 14 km w Lami, 7-10.x11.1986, R. L. & B. B. Brown, UV light trap, 2 males, 4 females. Stilobezzia (Stilobezzia) browni Wirth and Giles, New SpEcIES Figs. 26-32, 56 Allotype female.— Wing length 1.05 mm; breadth 0.36 mm. Head: Yellowish brown; palpi and anten- nal flagellum, except bases of first eight seg- ments, dusky. Eyes contiguous, bare. An- tenna (Fig. 28) with lengths of flagellar segments in proportion of 16-13-13-13-13- 14-14-14-30-30-30-30-40; antennal ratio 1.46. Palpus (Fig. 29) with lengths of seg- ments in proportion of 5-7-12-10-12. Man- dible with seven coarse teeth. Thorax: Pale brownish. Legs (Fig. 26) yel- lowish, hind leg with lengths from femur to tarsomere 5 as 120-110-60-22-7-6-16; without strong setae except a few at tips of femora. Hind basitarsus with 2'2 rows of palisade setae; one row on first two tarso- meres of fore and mid legs and tarsomere 2 of hind leg; fourth tarsomere of fore and mid legs with pair of strong black batonnets. A single long claw on each leg, claw with slender basal tooth nearly half as long as claw; length of claw 0.058 mm on fore leg, 0.080 mm on mid leg and 0.038 mm on VOLUME 92, NUMBER 3 hind leg (Fig. 27). Wing (Fig. 56) pale gray- ish hyaline, without macrotricnia, veins slightly infuscated; first radial cell small, slightly elongate; second radial cell spa- cious; costal ratio 0.68. Halter brownish. Abdomen: Y ellowish; terga brownish, with setae arranged in pattern as figured (Fig. 31). Spermathecae (Fig. 30) two, greatly unequal in size and shape; the larger brownish, 0.072 by 0.058 mm, oval without sclerotized neck; the smaller hyaline, spherical, 0.017 mm in diameter. Holotype male.— Wing length 1.05 mm; breadth 0.33 mm. Similar to the female with the usual sex- ual differences. Eyes broadly separated, bare. Antenna with well-developed plume which is yellowish proximally, dusky distally; lengths of flagellar segments in proportion of 25-14-14-14-14-14-14-14-14-16-43-50- 72; antennal ratio 1.08. Palpus as in female. Thorax darker brown than in female, scu- tellum paler. Legs yellowish, distal third of mid femur slightly darkened; tarsomeres 1- 3 of mid leg with pair of distal spines ven- trally; fifth tarsomeres without batonnets; claws short, equal, distally cleft on all legs. Wing as in female but narrower; costal ratio 0.66. Halter with dark brown knob. Genitalia (Fig. 32): Pale brownish, broad- er than long; ninth tergum convex caudally with a submedian pair of prominent setose cerci. Basistyle short, stout, tapering distal- ly; dististyle halfas long as basistyle, curved, gradually tapered to slender pointed tip. Ae- deagus reduced to a pair of small linear sclerites extending obliquely caudomesad from base of basistyle. Parameres separate, each with prominent basal knob, main por- tion in form of a very elongate, strongly sclerotized, sickle-like blade slightly swollen on proximal third, straighter on distal half. Distribution. — Fiji. Types. — Holotype male, allotype female, Viti Levu, 14 km w Lami, 7-10.x11.1986, R. L. and B. B. Brown, UV light trap. Para- types, 1 male, 1 female, same data; 1 male, 455 Suva, Koronivia Agr. Sta., 6.xii.1968, S. Singh, light trap (BISH). Discussion.—This species is named for Richard L. Brown in appreciation of his in- terest and cooperation in making available to us his fine collection of ceratopogonids taken during his visit to Fiji in 1986. Stilobezzia browni belongs to the subvir- idis group of the subgenus Stilobezzia as characterized by Das Gupta and Wirth (1968), but is not closely related to any of the Oriental species described in that group. It is most similar, especially in the shape of the male parameres, to S. flavizonata To- kunaga (1963), described from New Guinea, but that species (male only) is paler yellow, the abdomen dark with two basal terga pale yellow, and the aedeagus shaped differently. Tribe Heteromyiini Genus Clinohelea Kieffer Reference: Debenham, 1974: 6 (revision, species of Australia and New Guinea; key). Clinohelea tasmaniensis Lee Clinohelea tasmaniensis Lee, 1948: 65 (male, female; Tasmania; figs.); Deben- ham, 1974: 7 (redescribed; Australia rec- ords). Recorded distribution.— Australia (Tas- mania to southern Queensland). New records.—FIJI: Viti Levu, Lami, iii.1955, iii.1976, N. L. H. Krauss, 2 fe- males; Nagali, xi.1957 (Krauss) (BISH), 1 female; Suva, Koronivia, 6.xi1.1968 (Singh), light trap, | female (BISH). Note.—The two Fii females agree well with an Australian female in the USNM, except that the fore femur is dark brown only at the extreme tip in Fiji specimens. Tribe Sphaeromiini Genus Hebetula Wirth and Debenham Reference: Debenham, 1974: 22 (revision Australia and New Guinea spp.; as M/ixo- helea),; Wirth and Debenham, 1977: 282 (diagnosis; list of included species). 456 Hebetula tonnoiri (Lee) Xenohelea tonnoiri Lee, 1948: 66 (female; Tasmania; figs.); Tokunaga, 1966: 116 (female redescribed; New Guinea). Mixohelea tonnoiri (Lee); Debenham, 1974: 22 (combination; male, pupa described; Australia records; figs.). Hebetula tonnoiri (Lee); Wirth and Deben- ham, 1977: 282 (combination). Recorded distribution.— Australia, New Guinea. New records.—FIJI: Ovalua, Levuka, 111.1969 (Krauss), 1 female (BISH). Viti Levu, Lami, 1,111.1971, ii1.1976, 11.1977, xu1.1978 (Krauss), 12 females (BISH). 2 km N Naraiyawa, 27.x.1986, R. L. Brown, at light, 1 male, 1 female (USNM); Nausori Highlands, 500--700 m, 26.1ii.1970 (Krauss), | female (BISH); Suva, Koronivia Agr. Sta., 6.x11.1968 (Singh), light trap, 1 female (BISH); Suva, xi.1957 (Krauss), 2 females (BISH); Tholo-i-Suva, i.1955 (Krauss), 1 female (USNM). Genus Nilobezzia Kieffer Reference: Debenham, 1974: 62 (revision species Australia and New Guinea); Wirth and Ratanaworabhan, 1981: 408 (key to Oriental species). Nilobezzia fijiensis Wirth and Giles, NEw SPECIES Figs. 40-44, 57 Holotype female.— Wing length 2.48 mm, breadth 0.74 mm. Head: Dark brown including antenna and palpus. Eyes broadly contiguous. Antenna (Fig. 40) with lengths of flagellar segments in proportion of 25-13-13-13-13-13-14-16- 42-42-40-40-50, antennal ratio 1.65. Pal- pus (Fig. 41) with lengths of segments in proportion of 10-15-20-8-12; second seg- ment broadest, succeeding segments pro- gressively more slender. Mandibles with five and seven teeth. Thorax: Dark brown including scutellum (pinned paratype was mounted from fluid PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and thorax is dirty, pollinosity obscured if present). Legs dark brown; tarsi yellowish but narrow apices of tarsomeres 1, 3, most of 4, and all of tarsomere 5 brownish. Fem- ora each with one strong black ventral spine at apex; tibiae with 5-6 enlarged bristles scattered along dorsal side; tarsomere 5 (Fig. 42) with 12 black batonnets on fore leg, 14 on mid leg, and 10 on hind leg; claws 0.9 as long as fifth tarsomeres, external tooth 0.27 as long as claws. Wing (Fig. 57) pale smoky grayish, anterior veins yellowish; costal ratio 0.84. Halter brown. Abdomen: Dark brown. Gonopore flanked by several short bristles and 4—5 long black bristles on each side. Spermathecae ob- scured on slide mount; shapes and mea- surements not discernible. Allotype male.— Taken ata different time and place, the specimen described here is the presumed male of N. fijiensis, in spite of minor differences in setation and the pal- er color presumed to be due to prolonged storage in alcohol. Wing length 1.47 mm; breadth 0.45 mm; costal ratio 0.72. Thorax dark brown, head and legs paler brownish; wing grayish, radial veins slightly infuscated; halter brownish. Eyes contiguous. Antenna with sparse pale brownish plume; lengths of flagellar seg- ments in proportion of 28-16-15-15-15-16- 16-16-17-24-28-28-x; last segment not in position to measure. Palpus stubby; lengths of segments in proportion of 3-7-12-5-8. Legs without strong spines at apices of fem- ora. Genitalia (Fig. 44): Dark brown; shape and structure typical of the genus, with short ninth sternum and elongate ninth tergum with irregularly convex posterior margin. Basistyle and dististyle inperceptibly fused in a long, irregularly tapering lobe three- fourths as long as tergum. Aedeagus a tri- angular sclerite half again as long as basal breadth, with very slight anterior concavity, tapering distally to a slightly expanded cap- like tip. Parameres (Fig. 43) fused in another triangular tapering sclerite, slightly longer VOLUME 92, NUMBER 3 457 Se . Se Se Baer 40 da | 0.050mm 0.040mm Figs. 40-50. Figs. 40-44, Nilobezzia fijiensis, 40-42, female; 43-44, male; Figs. 45-50, Bezzia vitilevuensis, 45-48, female; 49, 50, male: 40, 45, antenna; 41, 46, palpus; 42, fifth tarsomere and claws of (top to bottom) fore, mid and hind legs; 43, 49, parameres; 44, 50, genitalia, parameres omitted; 47, femora and tibiae of (left to right), fore, mid and hind legs; 48, spermathecae. 458 and basally narrower than aedeagus, with slender knoblike tip bent posteroventrally. Distribution. — Fiji. Types.— Holotype female, Viti Levu, Lami, 11.1981, N. L. H. Krauss (USNM). Allotype male, Viti Levu, Suva, Koronivia Agr. Sta. 6.x11.1968, S. Singh, light trap (BISH). Paratype, 1 female, Viti Levu, Suva, 111.1956 (Krauss) (BISH). Discussion.—Nilobezzia fijiensis closely resembles the widespread Oriental species N. raphaelis (Salm) in its uniformly dark brown femora and tibiae, dark brown an- tenna and abdomen, and femora with only 1-2 apical spines, but raphaelis differs in having pale palpi and whitish halteres. Ni- lobezzia whartoni Lee from Australia and New Guinea is similar but possesses pale halteres and a whitish abdomen. Tribe Palpomyiini Genus Bezzia Kieffer References: Tokunaga, 1966: 141 (New Guinea species; key); Debenham, 1978: 557 (catalog, Australasian Region); Wirth and Ratanaworabhan, 1981: 413 (Southeast Asia species; key); Clastrier, 1985b: 45 (new species from New Caledonia). Bezzia vitilevuensis Wirth and Giles, NEw SPECIES Figs. 45-50, 58 Holotype female.— Wing length 1.02 mm; breadth 0.43 mm. Head: Dark brown. Eyes broadly contig- uous, bare. Vertex with strong spines curv- ing over eyes. Antenna (Fig. 45) with lengths of flagellar segments in proportion of 15- 12-12-11-10-10-10-10-20-20-20-20-21; antennal ratio 1.12. Palpus (Fig. 46) short and stubby; lengths of segments in propor- tion of 3-5-7-6-7. Mandible with nine coarse teeth. Thorax: Dark brown: mesonotum with scattered erect spinelike setae; supra-alar se- tae especially long and spinelike. Legs (Fig. 47) dark brown, tarsi whitish; femora un- armed; tarsomeres | and 2 of mid leg with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON one row of palisade setae, 1 and 2 of hind leg with two rows of palisade setae. Hind leg with lengths from femur to tarsomere 5 as 124-106-70-26-13-10-15; claws short, curved, equal, each claw 0.032 mm long. Wing (Fig. 58) slightly brownish infuscated due to coarse dark microtrichia; radial veins brownish; wing especially broad; costal ra- tio 0.85. Halter dark brown, knob intensely brown. Abdomen: Dark brown; one pair of gland rods extending anteriorly the length of two body segments. Spermathecae (Fig. 48) two, slightly ovoid with short narrow necks, slightly unequal, 0.039 by 0.032 mm and 0.036 by 0.029 mm including neck. Allotype male.— Wing length 0.79 mm; breadth 0.32 mm; costal ratio 0.82. Similar to the female with the usual sex- ual differences. Antennal plume very sparse and inconspicuous, pale brownish; flagellar segments with lengths in proportion of 20- 10-10-9-8-8-8-9-10-13-22-22-25; antennal ratio 0.65. Palpus reduced to tiny globular segments; proportions 3-4-6-5-6. Hind leg with lengths from femur to tarsomere 5 as 106-94-52-23-13-7-13; palisade setae pres- ent only on tarsomeres | and 1 of hind leg, in two rows. Genitalia (Fig. 50): Typical of the genus Bezzia; about as broad as long; ninth ster- num short with shallow caudomedian ex- cavation; ninth tergum short and convex, with long setose cerci. Basistyle short and tapering, without lobes; dististyle tapering irregularly to distal point. Aedeagus form- ing a nearly equilateral triangle; basal arms slender with low basal arch; distal process triangular, tapering to slender distal point. Parameres (Fig. 49) fused as usual in the genus; basal processes winglike with sharp anterior and lateral points; distal process long and spatuliform with slightly enlarged, rounded tip. Distribution. — Fiji. Types. — Holotype female, allotype male, Viti Levu, 14 km w Lami, 7-10.xii.1986, R. L. and B. B. Brown, UV light trap. VOLUME 92, NUMBER 3 459 Figs. 51-58. Wings of Ceratopogonidae: 51, Downeshelea stenochora; 52, Monohelea beaveri, 53, M. coloisu- vae; 54, M. fijiensis; 55, M. leveri, 56, Stilobezzia brownt;, 57, Nilobezzia fyiensis, 58, Bezzia vitilevuensis. Discussion.— The specific name is from Cah. O.R.S.T.O.M., ser. Ent. Med. Parasitol. 23: Viti Levu, the island where the species was 45-54. irae as . pee Das Gupta, S. K. and W. W. Wirth. 1968. Revision oii a) ; , taken. Bezzia vitilevuensis is readily distin of the Oriental apecies of Stilohezzia Kieller (Dip- guished from all other Bezzia species of the tera, Ceratopogonidae). Bull. U.S. Nat. Mus. 283: Oriental and Australasian Regions by the 1-149. combination of uniformly dark brown fem- Debenham, M. L. 1970. Australasian Ceratopogon- ora and tibiae and lack of femoral armature. idae (Diptera, Nematocera) Part XIII: Australian and New Guinea species of Echinohelea Macfie. Proc. Linn. Soc. New South Wales 94: 145-159. LITERATURE CITED ; 1971. Australasian Ceratopogonidae (Dip- Clastrier, J. 1985a. Ceratopogonidae de Nouvelle- tera, Nematocera) Part XV: The genus Al/uau- Caledonie II]. Genre Monohelea (Diptera, Nema- domyia Kieffer in Australia and New Guinea. Proc. tocera). Ann. Parasitol. Hum. Comp. 60: 747-759. Linn. Soc. New South Wales 96: 128-174. 1985b. Ceratopogonidae de Nouvelle-Cale- 1972. Australian and New Guinea “‘picture- donie III. Genre Bezzia (Diptera, Nematocera). wing” species of the genus Monohelea Kieffer 460 (Diptera: Ceratopogonidae). Australian J. Zool. Suppl. 12: 1-40. 1974. A revision of the Australian and New Guinea predatory Ceratopogonidae) (Diptera: Nematocera) of the tribes Heteromyiini and Sphaeromiini. Australian J. Zool. Supp. 28: 1-92. 1978. An annotated checklist and bibliog- raphy of Australasian Region Ceratopogonidae (Diptera: Nematocera). Univ. Sydney Commw. Dept. Health Monogr. Ser. Ent. Monogr. 1: 1-671. Downes, J. A. and W. W. Wirth. 1981. Chapter 28. Ceratopogonidae, pp. 393-421. Jn McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 1, 674 pp. Agric. Canada Monogr. 27. Ottawa. Lee, D. J. 1948. Australasian Ceratopogonidae (Dip- tera, Nematocera). Part V. The Pa/pomyia group of genera. Proc. Linn. Soc. New South Wales 73: 57-70, | plate. Ratanaworabhan, N.C. and W. W. Wirth. 1972. The biting midge genus Monohelea Kieffer in the Ori- ental Region. (Diptera: Ceratopogonidae). Pacific Ins. 14: 439-473. Tokunaga, M. 1959. New Guinea biting midges (Dip- tera: Ceratopogonidae). Pacific Ins. 1: 177-313. 1963. New Guinea biting midges (Diptera: Ceratopogonidae) 3. Pacific Ins. 5: 211-279. . 1966. Biting midges of the Palpomyiinae from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON New Guinea (Diptera: Ceratopogonidae). Pacific Ins. 8: 101-152. Tokunaga, M. and E. K. Murachi. 1959. Diptera: Ceratopogonidae. Insects of Micronesia 12: 103- 434. Wirth, W. W. and M. L. Debenham. 1977. Hebetula, a new genus of the predaceous midge tribe Sphae- romiini (Diptera: Ceratopogonidae). Proc. Ento- mol. Soc. Washington 79: 281-283. Wirth, W. W. and M. D. Delfinado. 1964. Revision of the Oriental species of Alluaudomyia Kieffer (Diptera, Ceratopogonidae). Pacific Ins. 6: 599- 648. Wirth, W. W. and W. L. Grogan, Jr. 1988. The pre- daceous midges of the world (Diptera: Ceratopo- gonidae; Tribe Ceratopogonini). Flora and Fauna Handbook no. 4. E. J. Brill, New York, xv + 160 pp. Wirth, W. W.and N.C. Ratanaworabhan. 1981. New species and records of predaceous midges (Dip- tera: Ceratopogonidae) from rice paddies in Thai- land. Pacific Ins. 23: 396-431. Wirth, W. W., N. C. Ratanaworabhan, and D. H. Mes- sersmith. 1977. Natural History of Plummers Island, Maryland XXII. Biting midges (Diptera: Ceratopogonidae). |. Introduction and key to gen- era. Proc. Biol. Soc. Washington 90: 615-647. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 461-464 GONATOPUS BARTLETTI OLMI [HYMENOPTERA: DRYINIDAE] IN MEXICO: A PREVIOUSLY UNREPORTED PARASITOID OF THE CORN LEAFHOPPER DALBULUS MAIDIS (DELONG & WOLCOTT) AND THE MEXICAN CORN LEAFHOPPER DALBULUS ELIMATUS (BALL) [HOMOPTERA: CICADELLIDAE] FERNANDO E. VEGA AND PEDRO BARBOSA! Department of Entomology, University of Maryland, College Park, Maryland 20742. Abstract. —The dryinid Gonatopus bartletti Olmi was reared from parasitized Dalbulus maidis (DeLong & Wolcott) and Dalbulus elimatus (Ball) collected in México. This is the first record of this dryinid in México. Key Words: Although the corn leafhopper, Da/bulus maidis, is mainly found at low to mid ele- vations, and the “Mexican corn leafhop- per,”’ Da/lbulus elimatus, is mainly found at high elevations (Barnes 1954, Nault in press), both species are pests of maize in México. These insects are capable of vec- toring 3 maize stunting pathogens: maize rayado fino virus (MRFYV), corn stunt spi- roplasma (Spiroplasma kunkelii Whitcomb et al.), and maize bushy stunt mycoplasma- like organism (Nault 1985). Yield losses due to MRFV transmission by Dalbulus maidis have been estimated at 40-50% of the weight of a mature ear in maize cultivars adapted to Central America (Gamez and Leon 1985). In newly developed cultivars losses can reach 100% (Gamez and Leon 1985). Little is known about natural enemies (Madden et al. 1986) or biological control of Dalbulus spp. In México, remnants of Dalbulus spp. were found in webs of the spider Tetragnatha sp. (Araneae: Tetrag- nathidae). Feeding by this spider on Dal- bulus was confirmed in the laboratory (F. ' Scientific article No. A-5033, contribution No. 8081 of the Maryland Agricultural Experiment Station. corn leafhopper, Dalbulus, dryinids, Gonatopus, México E. Vega, unpublished data). In the labora- tory, Hippodamia convergens Guérin- Méneville nymphs and adults fed on adult Dalbulus (F. E. Vega, unpublished data). In Nicaragua, Ectatomma ruidum (Formici- dae: Ponerinae) has been observed to prey on Dalbulus maidis (Perfecto 1989) while two fungi, Metarhizium anisopliae and Beauveria bassiana, are known to infect Dalbulus maidis (S. Gladstone, personal communication). So far, there is no infor- mation of Dalbulus egg predators or para- sitoids. Pipunculids, strepsipterans, and dryinids are known to attack leafhopper nymphs and adults (Waloff 1975), but in México, only dryinid parasitism has been observed (by F. E. Vega). In El Salvador, a dryinid identified as Agonatopus sp. was found attacking D. maidis (Quezada 1979), although it is sus- pected to be Gonatopus bartletti Olmi (M. Olmi, personal communication). G. bart- letti was first reported from Puerto Rico (Bartlett 1939, Olmi 1984), but Olmi (per- sonal communication) now has records from Nicaragua, Venezuela, Bahamas, and Be- lize. We present the first record of G. bart- letti (Hymenoptera: Dryinidae) in México. 462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. Male. Gonatopus bartletti Olmi. 1. Female. 2. MATERIALS AND METHODS To identify the parasitoids attacking D. maidis and D. elimatus, leafhoppers were collected in maize fields in 6 Mexican states: Jalisco, Guanajuato, Querétaro, México, Morelos, and Veracruz. Although other plant hosts could have been sampled for Dalbulus spp. (e.g. Tripsacum and the teo- sintes), we presumed it most logical to begin a search in maize agroecosystems within the widest geographical area we could sample. Those insects showing symptoms of para- sitism (e.g. a black spot in the abdomen or a bloated abdomen) were separated from the rest with a manual aspirator and taken to the laboratory where they were placed in plastic cups with plastic lids with a screen- covered rectangular hole. The cup con- tained about 1.5 cm of soil and fresh maize leaves which were replaced daily. RESULTS Five specimens identified as Gonatopus bartletti Olmi (Dryinidae: Gonatopodinae) were reared: two apterous females from 2 parasitized D. maidis and one winged male and 2 apterous females from 3 parasitized D. elimatus (for a complete description of the Dryinidae, see Olmi 1984). Parasitized insects were collected at the Colegio de Post- graduados in Montecillos, state of México, and along Road 43, 19 km west of Celaya in the state of Guanajuato. Parasitism by dryinids was observed in the state of Jalisco in the Pacific coast and eastward to the state of Veracruz on the Gulf coast. Only parasitoids in their late stages of de- velopment emerged, as indicated by the big sac on the host’s abdomen (Waloff 1974). Before the parasitoid larva emerged, the leafhopper exhibited sluggish behavior, and its wings and elytron had been pushed up- wards by the parasitoid sac. A few minutes before larval emergence the leafhopper clung to a leaf blade and died. After the parasitoid larva emerged, it moved around the cup, and spun a cocoon, either in the soil, in the leaf, or on the walls of the cup. VOLUME 92, NUMBER 3 Table 1. Dalbulus species found in México. . quinquenotatus . chiapensis maidis . tripsacoides charlesi gelbus . guzmani . longulus . guevarai . elimatus SSoeee es S's'> DISCUSSION México is the putative center of origin for ten of eleven Dalbulus species (Table 1) (Nault and DeLong 1980, Triplehorn and Nault 1985). Due to the number of Da/bulus species, their host plants, and the habitats in which these can be found, it can be argued that there should be a wider diversity of natural enemies of Dalbulus in México than elsewhere. A search in maize plantations in six Mexican states revealed only Gonatopus bartletti Olmi. Although the importance of this dryinid as a biological control agent is not known, a study by Waloff (1974) sug- gests that dryinids have the potential to con- trol leafhoppers. Using different leafhopper species she determined that a female Gona- topus sepsoides Westwood could potentially parasitize 177 leafhoppers over her lifetime. Further research on rearing methods, and the bionomics of this dryinid will determine the feasibility of the use of Gonatopus bart- letti Olmi as a biological control agent. An extended search for natural enemies of all other Dal/bulus species may uncover other parasitoids and predators which might act against Dalbulus maidis and Dalbulus eli- matus. ACKNOWLEDGMENTS We thank Javier Trujillo Arriaga, Alejan- dro Pérez Panduro, and the staff at the Cen- tro de Entomologia y Acarologia of the Co- legio de Postgraduados in Chapingo, México, for their help in conducting this 463 project. Dryinid identification by Arnold Menke (Systematic Entomology Labora- tory, USDA, U.S. National Museum) and Prof. Massimo Olmi (Istituto di Difesa delle Piante, Italy) is appreciated. We thank Ar- nold Menke, Lowell R. Nault, and Massimo Olmi for their comments, and Aileen Hsu (Carnegie Museum, Pittsburgh, PA) for the drawings. Financial support was provided by an Institute of International Education ITT International Fellowship to FEV. LITERATURE CITED Barnes, D. 1954. Biologia, ecologia, y distribucion de las chicharritas, Da/bulus elimatus (Ball) y Dal- bulus maidis (DeLong & Wolcott). Folleto Técnico ll, Secretaria de Agricultura y Ganaderia de Mex- ico, D. F., Oficina de Estudios Especiales y La Fundacion Rockefeller, 112 pp. Bartlett, K. A. (dated 1938 but issued in February 1939). A dryinid parasite attacking Balbulus mai- dis in Puerto Rico. J. Agric. Univ. Puerto Rico 22: 497-498. Gamez, R.and P. Leon. 1985. Ecology and evolution of a neotropical leafhopper-virus-maize associa- tion, pp. 331-350. Jn Nault, L. R., and J. G. Ro- driguez, eds., The Leafhoppers and Planthoppers. Wiley, New York. Madden, L. V., L. R. Nault, S. E. Heady, and W. E. Styer. 1986. Effect of temperature on the pop- ulation dynamics of three leafhopper species. Ann. Appl. Biol. 108: 475-485. Nault, L. R. (In press). Evolution of an insect pest of maize and the corn leafhopper: A case study. May- dica. Nault, L. R. 1985. Evolutionary relationships be- tween maize leafhoppers and their host plants, pp. 309-330. Jn Nault, L. R. and J. G. Rodriguez, eds., The Leafhoppers and Planthoppers. Wiley, New York. Nault, L. R. and D. M. DeLong. 1980. Evidence for co-evolution of leafhoppers in the genus Dalbulus (Cicadellidae: Homoptera) with maize and its ancestors. Ann. Entomol. Soc. Am. 73: 349-353. Olmi, M. 1984. A revision of the Dryinidae (Hy- menoptera). Mem. Amer. Entomol. Inst. 37: 1- 1913, Perfecto, I. 1989. Ants as biological control agents in the maize agroecosystem in Nicaragua. Ph.D. Dis- sertation. University of Michigan. Ann Arbor, Michigan, USA. Quezada, J. R. 1979. Hallazgo de Agonatopus sp.: (Hymenoptera: Dryinidae), parasito del Da/bulus maidis (Homoptera: Cicadellidae) en el Salvador. Ceiba 23: 1-12. 464 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Triplehorn, B. W. and L. R. Nault. 1985. Phyloge- of Dryinidae (Hymenoptera). J. Entomol. (A) 49: netic classification of the genus Dalbulus (Ho- 97-103. moptera: Cicadellidae), and notes on the phylog- Waloff, N. 1975. The parasitoids of the nymphal and eny of the Macrostelini. Ann. Entomol. Soc. Am. adult stages of leafhoppers (Auchenorrhyncha: 78: 291-315. Homoptera) of acidic grassland. Trans. Royal Waloff, N. 1974. Biology and behavior of some species Entomol. Soc. London 126: 637-686. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 465-470 A NEW HOST RECORD FOR EUR YTOMOCHARIS ERAGROSTIDIS HOWARD (CHALCIDOIDEA:EURYTOMIDAE) INFESTING ERAGROSTIS TEF IN SOUTH DAKOTA B. McDANIEL AND A. BOE Department of Plant Science, South Dakota State University, Brooking, South Dakota 57007. Abstract.—The species Eurytomocharis eragrostidis is for the first time recorded from South Dakota infesting Eragrostis tef grown in the Americas. The female ovipositor, propodeum and male reproductive apparatus are illustrated. Key Words: kota Stem-boring Hymenoptera infest cereal crops and numerous other grasses (Essig 1958). The genus Euryvtomocharis was found to infest up to 30% of stems of several range grasses in New Mexico (Watts and Bellotti 1967). Bugbee (1966) redefined the genus Eurytomocharis and described 4 new species from stems of grasses. Teff [Eragrostis tef (Zucc.) Trotter] is the primary grain crop of Ethiopia. This crop is currently being evaluated as a potential new forage crop for the northern Great Plains (Boe et al. 1986). Stunted growth of the crop in South Dakota in 1988 prompted dissec- tion of stems that revealed larvae of Eury- tomocharis eragrostidis Howard feeding in- side. The observations and data reported here describe a new insect-plant relation- ship and the magnitude of infestation of teff by E. eragrostidis at 2 widely-separated South Dakota locations. Anatomical de- scriptions, illustrations of male and female genitalia and additional morphological structures are presented to assist in the iden- tification of E. eragrostidis. MATERIALS AND METHODS In late July 1988, 7 random teff plants from each of 4 replications in field trials at Hymenoptera, Eurytomidae, Eurytomocharis eragrostidis Teff, South Da- Brookings and Highmore, South Dakota, were examined for infestation by E. era- grostidis. The trials were planted on 17 May and | June 1988 at Highmore and Brook- ings, respectively. Tillers of each plant were slit longitudi- nally with a razor blade and numbers of larvae, pupae, and exit holes were deter- mined for each mature internode. Approx- imately 30 internodes containing larvae or pupae were placed in small, covered glass jars and adults that emerged in the summer and fall of 1988 were collected for identi- fication. RESULTS AND DISCUSSION Plant-insect interaction. —F. eragrostidis infested a high percentage of teff tillers at both South Dakota locations in 1988. In- dividual tillers frequently had more than one infested internode, resulting in 28 and 38% internode infestation at Brookings and Highmore, respectively (Table 1). Watts and Bellotti (1967) reported that the 4 Eurytomocharis species found infest- ing range grasses in the southwestern United States had similar life cycles with only | generation per year on any given grass species. In South Dakota, exit holes were 466 Table 1. Infestation rates of teff by Eurytomocharis eragrostidis at two South Dakota locations in 1988. Numbers examined Percent infested! Inter- Inter- Location Tillers nodes Tillers nodes Highmore 132 449 2a We, 38 +4 Brookings 158 434 SOE 7 28et3 ' Mean + standard deviation of 4 replications. found in approximately 30% of the infested teff internodes in late July (Fig. 1) and adults of E. eragrostidis emerged under laboratory conditions in August and September 1988. More research is needed to determine if a second generation can be produced on teff in South Dakota. Teff had been grown at both locations for more than 5 years with no previous indi- cation of stem-boring insect problems. E. eragrostidis has been previously reported from Agropyron spp., Agrostis alba L., An- dropogon saccharoides Swartz, Eragrostis cilianensis (All.) Lutati., E. erosa Scribn., E. poaeoides Beauv. ex Roem. and Schult., Muhlenbergia porteri Scribn., M. wrightii Vasey, Oryzopsis hymenoides (Roem. and Schult.) Ricker, and Sporobolus airoides (Torr.) Torr. (Burks 1979). Watts and Bellotti (1967) found E. era- grostidis most frequently on side-oats grama [Bouteloua curtipendula (Michx.) Torr.] in New Mexico. E. cilianensis and side-oats grama occur commonly throughout South Dakota, but more study is needed to deter- mine if these species are hosts of E. era- grostidis in the northern Great Plains. Several parasitic chalcids have been fre- quently recorded from stem-boring eury- tomids in range grasses (Watts and Bellotti 1967). No parasitic chalcids emerged from teff internodes stored in the laboratory nor were any observed in dissected stems. How- ever, since dissection frequently destroyed the enclosed larvae, few adults were reared from the samples. Exit holes observed in plants in the field in July may have been PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. teff. Exit hole of Eurytomocharis eragrostidis in made by parasites of E. eragrostidis. A more thorough study would be required to deter- mine if FE. eragrostidis is parasitized in teff. The apparent reduction in growth due to E. eragrostidis may be an important factor influencing the potential of teff for forage production in the northern Great Plains. Forage yields of teffat both locations in 1988 were approximately one-fourth of the forage yields obtained for several years prior to the discovery of infestation by FE. eragrostidis in 1988. Watts and Bellotti (1967) observed similar stunting symptons in range grasses from which they collected FE. eragrostidis. External and reproductive structure mor- phology.—The following description is based on 19 specimens collected from teff, of which 2 females and 2 males were sent to the United States National Museum for identification. Female: Head dark black; mesosoma black to reddish black; pronotum, propo- deum, petiole, gastor bluish black; antenna with scape yellow (two specimens had dark scape); flagellum yellow, bordered with brown; maxillary palp white; mandibles white; legs with coxae, trochanters, femora and tibia yellow; tarsi yellow; gena carinate, VOLUME 92, NUMBER 3 head with fairly deep piliferous punctures dorsally between ocelli extending past pos- terogenal region; plesosoma with same pilif- erous punctures; head with frons concave between eyes; antennal scrobes at base of concave frons at or below eye; scape longer than pedicel and first funicular segment to- gether; funicular segments separated by short stalk; club segments fused, 3 in number, first two similar in shape and size to funicular segments, last segment broad at base, nar- rowing to rounded apex; mesosoma collar broad, edge rounded, without punctures; pronotum with piliferous punctures; scu- tellum convex, scutoscutellar sulcus contin- uous, both covered with piliferous punc- tures; marginal vein broader and longer than postmarginal; parastigma with clear region at attachment to marginal; stigmal subequal to postmarginal, shorter than marginal; sub- marginal vein with single sensillum behind first dorsal submarginal setae; dorsal par- astigma with 2 sensilla, pigmented portion extending anterior of sensilla, the remainder clear with light pigmentation in center; cos- tal cell with single row of dorsal setae, ven- tral surface with scattered setae; speculum closed; cubital hair line and medial hair line distinct; propodeum (Fig. 3) with light yel- low tinge, lighter than scutellum, darker than hind coxae, area near coxae setose, plated, center with shallow furrow, asetose, all plates irregular in shape not punctured as found on scutellum (Fig. 3); Ovipositor (Fig. 2) semicircular sheaths (2nd valvifers) with 2 setae near laminated bridge, rami spines not fixed in number (mean numbers of spines were 36.6 + 2.1 and 36.8 + 1.6 for left and right rami of a sample of 6 females); fulcral plate and inner ovipositor plate attached to semicircular sheets; fulcral plate notched with four monitoring spines; attachment of outer ovipositor plate is in-between fulcral plate notch and attachment of fulcral plate and curved ramus edge; outer ovipositor plate fused with eighth tergite; eighth tergite contains button-like cercus with five setae of different sizes and shapes; near cercus a 467 series of setae are found to be dispersed along the eighth tergite, (these are more numerous near the cercus becoming single toward the fulcral plate); eighth tergite setal region plat- ed, bordered by dark line that divides fused outer ovipositor plate into different pig- mented areas; apex of eighth tergite with series of long setae; inner ovipositor plate separated from semicircular sheath by dark- ened region; below darkened region is a groove in which the fulcral plate fits along with monitoring spines; inner ovipositor plate plated to region of fused ovipositor sheath; ovipositor sheath not articulated; ovipositor sheaths lightly plated with lon- gitudial striae with a series of setae at apex. Male: Color similar to female with head dark black; mesosoma, pronotum, propo- deum black; petiole elongated, black; gastor black; ventor dark due to dark coxae in some specimens, other specimens with coxae yel- low similar to female; antennae with scape darker than in female; flagellum lighter than body color, with five funicular segments, long setae and well-developed petiole seg- ments; maxillary palp and mandibles yel- low; legs with coxae dark, similar in color to body, with some yellow; trochanters, femora and tibia yellow, hind femora with some brown in middle section; tarsi yellow; sculpturing similar to female; wings same as female except medial not distinguished by a single row of setae, rather with 2-3 closely set setae marking region of medial hairline; costal cell same as female; basal setae well-developed, joining cubital hair- line closing speculum; male reproductive apparatus (Figs. 4, 5) with aedeagus bilobed, each lobe with 6 sensory pores on ventor; parameres with 2 setae, one associated with apex which is usually hidden between the digiti and the aedeagus, second setae on the narrow arm of the fused parameres, the lat- ter larger than apical setae; single ventral setae located on expanded portion of par- ameres; digiti with 2 spines, digiti with paired pore-like structures and paired dig- ital apodemes; aedeagal apodemes protrude 468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ————— Ful PLSp. ——-Ful.PLNot SUS Wyss LFS =~ 2574 GATES rts = VOLUME 92, NUMBER 3 from caulis; aedeagus dorsally covers digiti and caulis; a covering attached to the eighth tergite is torn away from aedeagus when re- moving whole male reproductive structure (this structure is covered with small setae and may function in insertion of the aede- agus into the female). At apex of aedeagus on dorsal surface are pigmented raised areas (these raised areas are not the same pig- mented pores that are observed on the ae- deagus of members of the genus Bruchopha- gus). Copland and King (1972) stated that the number of rami spines varies with species but did not indicate that they vary within individual females as well as between val- vifers of a single female. In E. eragrostidis reared from teff the mean number of ramus spines was 36.7. The numbers of left and right ramus spines were equal in only two of the six females examined. The largest dif- ference found between number of left and right ramus spines within an individual fe- male was four (39 on the left ramus and 35 on the right ramus). Rami spines are con- sidered to be sensory in nature, serving to monitor the position of the stylets (Copland and King 1972). They are most widely spaced in the region close to the laminated bridge becoming closer together near at- tachment of fulcral plate. In a current study of the genus Bruchophagus, we have found that the mean number of ramus spines can — 469 be used to statistically separate the closely related species that attack leguminous seeds. Arrangement of eighth tergite setae has been found to be of value in separating closely related species of the genus Brucho- phagus and may well be a diagnostic struc- ture in separating species of Eurytomo- charis. However, we have not examined other species of Eurytomocharis to deter- mine if setal arrangment differences do exist within the genus. In E. eragrostidis the ovipositor sheaths are connected by a ligament. This ligament is easily torn during slide preparation, giv- ing the appearance that the ovipositor sheaths are not connected. On slide-mount- ed genitalia the ovipositor sheaths have a lip-like projection that is the broken ovi- positor sheath ligament. It is from these pro- jections that the thin ligament is attached connecting the ovipositor sheaths. ACKNOWLEDGMENTS The authors extend their gratitude to Dr. E. E. Grissell, Systematic Entomology Lab- oratory, ARS/USDA, United States Na- tional Museum for providing identification of Eurytomocharis eragrostidis and to Ka- thy Robbins for assistance in data acquisi- tion. This research was supported by the South Dakota Agricultural Experiment Sta- tion, SDSU, Brookings, project numbers H-277 and H-388, contribution no. 2446. Fig. 2. Ovipositor of Eurytomocharis eragrostidis collected from teffin South Dakota. Abbreviations: (Lam. Br.) Laminated bridge; (Ra) Ramus; (Ra. Sp.) Ramus Spines; (Vf2) 2nd valvifer (Semicircular sheath); (Ful. Pl.) Fulcral plate (1st valvifer); (Ful. Pl. Not.) Fulcral plate notch; (Ful. Pl. Sp.) Fulcral plate spines; (Ops. In. Pl.) Ovipositor Inner Plate (3rd valvulae); (Ops. Ot. Pl.) Outer ovipositor plate (8th tergite); (Ops.) Ovipositor (1st & 2nd valvulae); (Ops. Sh.) Ovipositor sheath; (Ops. Sh. Lg.) Ovipositor Sheath Ligament; (Ce.) Cercus; (8-Teg. Set.) 8th tergite setae; (Ep.) Epipygium; (8-teg.) 8th tergite. Fig. 3. Female propodeum. Abbreviations: (Mt.) Metanotum; (Sp.) Spiracle; (Pli.) Plica; (N.) Nucha (neck). Figs. 4,5. Male reproductive apparatus, dorsal, and ventral respectively. Abbreviations: (Aeg. S. Por.) Aedeagal sensory pores; (Aeg.) Aedeagus; (Dgi.) Digiti; (Dgi. Por.) Digiti pore; (F. Par. Pl.) Fused Paramere Plate; (Dgi. Apd.) Digiti apodemes; (Vt. Por.) Ventral Pore; (Ca.) Caulis; (Vos. Pl.) Volsellar Plate; (Aeg. Apd.) Aedeagus apodemes; (Aeg. Bilo. Ap.) Aedeagus Bilobed apex. 470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Boe, A., J. Sommerfeldt, R. Wynia, and N. Thiex. 1986. A preliminary evaluation of the forage po- tential of teff. Proc. S. D. Acad. Sci. 65: 75-82. Bugbee, R. E. 1966. A revision of the genus Eury- tomocharis Ashmead in North America (Hyme- noptera:Eurytomidae). Amer. Midl. Natur. 75: 367-382. Burks, B. D. 1979. Family Eurytomidae, pp. 835- 860. In Krombein, K. V., P. D. Hurd, D. R. Smith, B. D. Burks and others, Catalog of Hymenoptera in America North of Mexico. Vol. 1: Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington. xvi + 1-1198. Copland, M. J. W. and P. E. King. 1972. The struc- ture of the female reproductive system in the Eury- tomidae (Chalcidoidea: Hymenoptera) J. Zool., Lond. 166: 185-212. Essig, E.O. 1958. Insectsand Mites of Western North America. The Macmillan Co., New York. Watts, J. G. and A. C. Bellotti. 1967. Some new and little-known insects of economic importance on range grasses. J. Econ. Entomol. 60: 961-963. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 471-482 THE NEMOMYDAS OF SOUTHWESTERN UNITED STATES, MEXICO, AND CENTRAL AMERICA (DIPTERA: MYDIDAE) Boris C. KONDRATIEFF AND JUDITH L. WELCH Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. Abstract.—The species of Nemomydas Curran occurring in the southwestern United States, Mexico, and Central America are reviewed. Thirteen species are recognized, in- cluding two new species, N. fronki n. sp. and N. wendyae n. sp. The males of N. brachy- rhynchus (Osten Sacken) and N. sponsor (Osten Sacken) are described for the first time, N. fumosus Hardy is elevated to a full species, and N. panamensis (Curran) is considered a nomen dubium. A key to the males is provided. Key Words: Nemomydas Curran is a distinctive genus of mydas flies found in North and Central America, and contains a number of geo- graphically restricted species. Hardy (1950) revised the North American species, and Steyskal (1956) reviewed the species in the eastern United States. However, our knowl- edge of the Mexican and Central American species is poor. Recent collections from Mexico, Guatemala, southwestern United States, as well as the examination of type specimens, has made it possible to review the fauna of this area and clarify the taxo- nomic status of several regional species. Papavero and Wilcox (1968) transferred Mydas senilis Westwood to Nemomydas. Our examination of the male holotype of this species indicate it as a true Mydas as defined by Wilcox and Papavero (1971), and a member of the interruptus group (Welch and Kondratieff 1990). We also examined the types of N. desideratus (Johnson) (MCZ #7592) and N. jonesii (Johnson) (MCZ #7593) as well as available specimens of N. lara Steyskal and N. melanopogon Steyskal. These four species are restricted to the southeastern United States, especially Flor- Nemomydas key, new species ida. Nemomydas lara, originally described from females, is probably the female of N. melanopogon, a species known only from males. Specimens for this study were provided by the following institutions: British Mu- seum (Natural History) (BMNH); Califor- nia Academy of Sciences (CAS); Canadian National Collection (CNC): Colorado State University (CSU); Florida State Collection of Arthropods (FSCA); Michigan State Uni- versity (MSU); Museum National D’His- toire Naturelle, Paris (MNHN); Museum of Comparative Zoology, Harvard (MCZ): San Diego Natural History Museum (SDNHM)- University of Arizona (UA); University of California, Berkeley (UCB); University of Colorado (UC); University of Kansas (UK); and the United States National Museum of Natural History (USNM). Methods of preparation, and terminology of the male genitalia, follow Wilcox and Pa- pavero (1971) and Wilcox (1981). This study illustrates, for the first time, the male ter- minalia of all species known from this re- gion. The females of N. bifidus Hardy, N. solitarius (Johnson), and N. tenuipes (Loew) 472 are unknown, therefore no key is presented for the females. We follow Snelling’s (1987) political designations for Lower California. KEY TO THE MALES OF NEMOMYDAS OF WESTERN NORTH AMERICA AND CENTRAL AMERICA 1 peoeeels in lateral view, enlarged or expand- Cdi(Pipss2—5) itive: sera sim ree eeterrre neers 2 — Distal section of aedeagus tube-like and elon- gate in lateral view (Figs. 1, 6-13) 2. Distal section of aedeagus abruptly expanded and recurved medially in lateral view (Figs. ORT) FE er ote ncaa abnoogde Jodene boonenoT 3 — Aedeagus swollen, tapering apically in lateral view (Figs. 1, 3, 5) 3. Abdominal tergites yellowish brown (in some specimens, tergites shaded with brownish black); terminalia as Fig. 2. Distribution: southwestern United States (Arizona) and northern Mexico (Sonora) : _brachyrhynchus (Osten Sacken) — Abdominal tergites black with posterior mar- gins of tergites 2-6 yellow; terminalia as Fig. 4. Distribution: Guatemala and Costa Rica 4 sponsor (Osten Sacken) 4. Abdominal ieee: black, posterior margins of 1-6 or 1-7 yellow 5 — Abdominal tergites yellow oat with ace dorsal blackish brown spots or dashes; ter- minalia as Fig. 1. Distribution: Honduras . bequaerti (Johnson) ay Acdedgus in fateral view tongue-like, con- stricted medially (Fig. 3). Distribution: Costa Rica srrcninvt aan eee aia lamia (Séguy) — Aedeagus in lateral view broad basally, ta- pering apically with small flange anteroapi- cally (Fig. 5). Distribution: Mexico . AEE CO nn at. cer wendyae Nn. sp. 6. Tergites entirely black or reddish black .... 7 — Tergites reddish, yellow, yellowish brown or dark and distinctly marked with white or yel- LOW ee tet een ree tance cloteuae siete 8 7. Dorsal digitate process of gonocoxite in lateral view, small originating from inner surface of ventral digitate process (Fig. 6); body and legs entirely black, covered by long black setae. Distribution? Texas: 1.5. sec. oe cei fronki n. sp. — Dorsal digitate process of gonocoxite in lateral view, large originating at base of ventral dig- itate process (Fig. 7); thorax, tergite 1 or 2 with yellow or white setae. Distribution: Cal- ifornia, Mexico (Baja California) . et Ne, SAR Ie ee nin terre tenuipes (Loew) 8. Tergites brownish black or black marked with yellow or white PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Tergites reddish yellow, yellow or yellowish brown 9. Tergites 1-7 black or brownish black with transverse yellow margins posteriorly; basal portions of hind femur and tibia yellow; ter- minalialasinihigalSin. ase venosus (Loew) — Tergites brownish black, tergites 3-5 with tr- angular, whitish spots laterally; basal portions of hind femur and tibia blackish brown; ter- MinaliavasiFig: lileaee eee ere fumosus Hardy 10. Dorsal digitate process of gonocoxite, in lat- eral view, small, originating from inner sur- face of ventral digitate process (Fig. 12). Dis- tribution: Colorado ........ solitarius (Johnson) — Dorsal digitate process of gonocoxite, in lat- eral view, originating at base of ventral digi- tate process (Figs. 8, 9). Distribution: British Columbia south to Mexico (Baja California) 11. Proboscis short, 0.8—0.9 times as long as sub- cranialicavityss.ere act pantherinus (Gerstaecker) — Proboscis long, 1.3 times as long as subcranial cavity . : 12. Abdominal tergites and legs with dense yellow toswhite;setaeia 7. “.naqesecimaees intonsus Hardy — Abdominal tergites and legs with black setae tia eS ote ee bifidus Hardy Nemomydas bequaerti (Johnson) Figs. 1, 14 Leptomydas bequaerti Johnson, 1926: 144. Type locality: Honduras, Depto. Colon, Puerto Castilla. Holotype male (MCZ #7594), examined. Nemomydas bequaerti, Papavero and Wil- cox, 1968: 34.10. Johnson (1926) provided an adequate de- scription of both sexes of this species. In his key, Johnson stated that N. bequaerti has cell r, open; however, our examination of all material available, including the holo- type male, showed that this cell is closed. The following may be added to the original description of the male: proboscis short, 0.6 times as long as subcranial cavity; tergite | with long erect whitish setae; and tergites 2-7 with recumbent, short, black setae. The extent of the middorsal blackish brown dashes or spots on abdominal ter- gites is variable. The holotype and | addi- tional specimen have these marks on ter- VOLUME 92, NUMBER 3 gites 2-7, whereas another specimen has these marks only on tergites 2-3. Nemomydas bequaerti may be easily sep- arated from all others in this study (Central American Nemomydas) by the short stubby aedeagus (Fig. 1), and tergites 2-7 yellow- brown with middorsal blackish brown dash- es or spots. The female of N. bequaerti is very similar to the female of N. brachy- rhynchus Osten Sacken, but may be dis- tinguished by the more elongate second flagellomere (Fig. 14), and apparent geo- graphical distribution (Honduras). The fe- male of N. brachyrhynchus has an expanded second flagellomere (Fig. 15) and is known from northern Mexico (Sonora) to southern Arizona. Material examined.— HONDURAS: Ho- lotype male as noted above; paratype male, same data except 28 III 1924 (MCZ); Puerto Castilla, 2 TV 1926, R. H. Painter, 2 males, 1 female (CAS); same data 1 male, 1 female (CNC); same data but 26 III 1924, 1 male, 1 female (BMNH). Nemomydas brachyrhynchus (Osten Sacken) Figs. 2, 15 Leptomidas brachyrhynchus Osten Sacken, 1886: 69. Type locality: (Northern) So- nora, Mexico, Holotype female (BMNH), examined. Leptomydas brachyrhynchus, Johnson, 1926: 142. Nemomydas brachyrhynchus, Hardy, 1950: DD: Nemomydas brachyrhynchus, Papavero and Wilcox, 1968: 34.10. Male.—Length 12-19 mm. Head shiny black, setae erect, golden yellow to white; antenna 3.2 mm long, reddish black, tinted with orange, especially apically; proboscis short, 0.8 times long as subcranial cavity, reddish brown. Scutum reddish brown to reddish black, 3 broad brownish black stripes slightly converging posteriorly; scu- tellum shiny brown; posterior portion of 473 postnotum blackish brown; wing hyaline, membrane around longitudinal veins tinted with brown; halter yellow; legs yellowish brown, except coxa and trochanters shiny brown, distal portion of hind femur and tib- ia brownish black, tarsus tinted with brown, setae on distal portion of hind femur and hind tibia black, others yellow. Abdominal tergites usually yellowish brown, in some specimens shaded with brownish black, pleural margins blackish brown; bulla black, setae long, erect and whitish to yellow on tergite 1, short, recumbent and black on ter- gites 2-7; sternites yellowish brown, darker posteriorly. Terminalia.— Yellowish-brown, gono- coxite tinted with blackish-brown; dorsal digitate process of gonocoxite slender, curv- ing inward; aedeagus in lateral view, api- cally expanded (Fig. 2). This is the first description of the male. Hardy (1950) suggested that N. brachyrhyn- chus was a possible synonym of N. panth- erinus. The association of the male with the female clearly indicates that this species is distinct. Males of N. brachyrhynchus can be easily distinguished from all other Nemo- mydas by the combination of aedeagus with an enlarged apical section, (Fig. 2) and usu- ally with yellowish brown abdomen. Several specimens from Sonora, Mexico, and Ari- zona have tergites shaded with brownish black. The aedeagus of N. sponsor (Fig. 4) is similar to that of N. brachyrhynchus but the male of N. sponsor has a black abdomen with yellow posterior margins on tergites 2— 6. The female of N. brachyrhynchus 1s sim- ilar to the females of N. bequaerti, N. pan- therinus (Gerstaecker) and the light phase of N. venosus (Loew). However, N. brachy- rhynchus is readily distinguished from N. pantherinus by the following features: (1) lacking long dense postocular setae (N. pantherinus has long dense postocular se- tae); (2) lacking dense lateral scutal setal fringe (N. pantherinus has a dense fringe of long yellow to white setae); and (3) abdom- inal tergites without brown transverse bands 474 (N. pantherinus females have brown ante- rior transverse bands on tergites 2-5). The light phase of N. venosus also lacks the dense, long postocular setae, and the dense lateral scutal setal fringe, but has abdominal ter- gites 2-4 or 2-7 with brown anterior trans- verse bands. Characters for separation of the female of N. brachyrhynchus from N. bequaerti are given in the discussion of the latter species. Material examined.— MEXICO: Sonora, holotype female (BMNH); 7 mi. S. Alamos, Rio Cuchajachi, 20 March 1985, L. Stange and R. Miller, 2 males (FSCA); ARIZONA: Cochise Co., 2 mi. NE of Portal, 30 V 1962, J. Wilcox, | female (CAS); Pima Co., Mad- rona Ranger. Sta., W. Rincon Mts., 15 V 1964, at mud and water, M.L. Noller, J. C. Bequaert, H. Eltom, M. Nurein, F. G. Wer- ner, | female (UA); Sabino Canyon, 26 V 1962, F. D. Parker and L. A. Stange, 2 males (CAS); same data except 8 V 1961, Sharp, 1 male; Santa Catalina Mtns., 9 V 1950, J. Markley, | female (UA); same data except 30 IV 1955, F. G. Werner, | female (CAS); same data except 30 IV 1955, F. G. Werner, 1 male (UA); same data except | V 1956, G. D. Butler, | male (UA); same data except 13 V 1960, Halberg, | female (UA); same data except 5 V 1961, B. Bryce, 1 male (UA); same data except 8 V 1961, R. Band, | male (UA); same data except Wargo, | male (UA); same data except 8 V 1961, C. Jackson, | female (UA); same data except 11 V 1962, Huiseir, | female (UA); same data except 13-14 V 1962, E. Stout, 1 male, | female (UA); same data except 6 V 1966, B. L. O., 1 female (UA); same data except 6 V 1966, G. Roux, | male (UA); same data except 9 V 1966, Donald, 1 male (UA); same data except Molino Basin, 12 V 1980, C. Olson, MacLachlan, 1 male (UA); Tucson, 21 IV 1957, Witman, 1 female (UA); same data except 6 V 1962, D. Parks, 1 male (UA); Santa Cruz Co., Madera Canyon, 17-21 V 1971, J. Wilcox, 2 males, 2 females (CAS); Bear Canyon, 12 V 1961, E. M. Painter, 1 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON male (CAS); Santa Rita Mtns., VIII 1977, C. A. Olson, | female (UA). Nemomydas lamia (Seguy) Fig. 3 Nomoneura lamia Séguy, 1928: 146. Type locality: La Caja, Costa Rica. Lectotype male (here designated and so labelled, MNHN), examined. Nemomydas lamia, Papavero and Wilcox, 1968: 34.11. Male.— Length 13 mm. Head shiny black, setae erect, whitish; orbital margin of com- pound eye whitish; antenna 3.2 mm long, brown, tinted reddish brown and gray pol- linose apically; proboscis long, 2.3 times as long as subcranial cavity, black. Scutum black, with pair of submedian yellowish pollinose stripes converging posteriorly, a pair of lateral yellowish pollinose stripes, setae whitish, sparse, erect; postnotum with lateral yellowish white areas; wings hyaline, longitudinal veins brown; halter brown; foreleg and midleg brown, setae whitish, hindleg brown with basal portion of femur yellowish, setae whitish dorsally, black ven- trally on femur, spines reddish brown; tibia brown. Abdominal tergites shiny black, posterior margins on tergites 1-7 yellow, bulla black, setae long and erect, whitish on tergites 1-2, short and whitish on 3-7; ster- nites shiny black, setae long, whitish and erect. Terminalia.— Reddish-brown; ventral digitate process of gonocoxite thickened, thumb-like; dorsal digitate process slender, curved inward; aedeagus in lateral view tongue-shaped, constricted medially; in ventral view tapering apically (Fig. 3). Female.—Length 15-18 mm. General coloration and structure similar to males except posterior margins on tergites 1-4 or 5 yellow. Material examined.—COSTA RICA: La Caja, Paul Serre, 1920, Lectotype male des- ignated (marked with blue margined lecto- VOLUME 92, NUMBER 3 type label); 6 female paralectotypes, same label data as lectotype (MNHN). Remarks. —Séguy (1928) did not desig- nate a holotype. The syntypes of N. /amia sent to us by D. Baylac (Museum National D’Histoire Naturelle) consisted of a male marked with a red type label and a label ““Nemoneura lamia Seguy, type,”’ (consid- ered here as the lectotype), a female marked also as “type,” another male and 6 females. These 8 specimens have similar Costa Rica locality labels and determination labels of N. Papavero (in 1970). The second male syntype 1s identical to the male of NV. sponsor and included under that species. The 6 fe- males are all considered to be N. /amia. The male of N. /amia can be distinguished from the closely related N. wendyae and N. spon- sor by the distinctive aedeagus (Fig. 3) and the whitish setae of the head. The females of N. /amia, N. sponsor, and N. wendyae are very similar. Nemomydas lamia has blackish brown abdominal ter- gites, with tergites 1-4 or 5 with yellow pos- terior transverse margins. Both N. sponsor and N. wendyae have tergites 5-7 reddish brown. Nemomydas sponsor (Osten Sacken) Fig. 4 Leptomydas sponsor Osten Sacken, 1886: 68. Type locality: San Geronimo, Gua- temala. Holotype female (BMNH), ex- amined. Male.— Length 14 mm. Head shiny black, setae erect, black to reddish brown; orbital margin of compound eye whitish; antenna 3.1 mm long, black, tinted with brown; pro- boscis long, 1.8 times as long as subcranial cavity, black. Scutum black, with pair of submedian gray white pollinose stripes con- verging posteriorly, a pair of lateral gray white pollinose stripes; anterior portion of scutellum gray white pollinose, posteriorly shiny black, setae whitish, sparse, erect; 475 a 3 b Figs. 1-3. 1. Nemomydas bequaerti. Male termi- nalia, a. lateral, b. ventral. Abbrev.: ae, aedeagus; al, anal lamellae, ce, cercus; ddp, dorsal digitate process; ep, epandrium; goncx, gonocoxite; vdp, ventral digitate process. 2. Nemomydas brachyrhynchus. Male termi- nalia, a. lateral, b. ventral. 3. Nemomydas lamia. Male terminalia, a. lateral, b. ventral. postnotum with lateral gray white areas; wings hyaline except membrane around longitudinal veins tinted with brown; halter black; foreleg and midleg black, setae yel- lowish, hindleg black with basal portion of femur yellowish, setae black ventrally on femur, spines reddish brown; tibia reddish black. Abdominal tergites shiny black, pos- terior margins of tergites 1-6 yellow, brown on tergite 8, bulla black, setae long and erect, whitish on tergites 1-2, short and black on 3-7; sternites shiny black, setae long, whit- ish and erect. 476 Terminalia.— Reddish brown, dorsal and ventral digitate processes of gonocoxite elongate; aedeagus constricted medially, ex- panded and recurved apically (Fig. 4). Material examined.—COSTA RICA: 1 male (MNHN); GUATEMALA: Depto. Guatemala, Tacaton, Lago Amatitlan, near Villa Canales, 10 I 1989, B. C. Kondratieff, 1 male, 2 females (CSU); Holotype female, S. Geronimo, Champion (BMNH). Remarks.—The male is described here for the first time. Osten Sacken (1886) pre- sented an excellent descriptions of the fe- male of this species. The male of N. sponsor resembles N. /amia, N. wendyae and N. ve- nosus but may be immediately distin- guished by the form of the aedeagus (Fig. 4). The female is very similar to N. wendyae but can be separated by the distal flagello- meres being reddish brown and brown setae of face, whereas the female of N. wendyae has the antennae distally blackish-brown and white facial setae. Nemomydas wendyae, NEw SPECIES Figa5 Male.—Length 13.5 mm. Head shiny black, setae erect, black; orbital margin of compound eye whitish; antennae 3.9 mm long, scape and pedicel black, flagellomere 1 and 2 brownish red, 3 and 4 black, tip of 4th silvery pollinose; proboscis long, 2.2 times as long as subcranial cavity, black. Scutum black, with pair of submedian gray white pollinose stripes converging poste- riorly, a pair of lateral gray white pollinose stripes; anterior portion of scutellum gray white pollinose, posteriorly shiny black, se- tae whitish, sparse, erect; postnotum with lateral gray white areas; wings hyaline, lon- gitudinal veins brown; halter black; foreleg and midleg blackish brown, setae mainly black, hindleg black with basal half of femur yellowish with whitish setae, distally setae black, spines reddish-brown; tibia basally yellowish, distally blackish-brown, setae black, tarsi black. Abdominal tergites shiny PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON black, posterior margins on tergites 1-6 yel- low, brown on tergite 8, bulla black, setae long and erect, whitish on tergites 1-2, short and black on 3-7; sternites shiny black, se- tae long, whitish and erect. Terminalia.—Reddish-brown, dorsal digitate of gonocoxite tapered, apically acute, directed inward; aedeagus broad basally, wedge shaped, with small flange apically (Fig. 5): Female.—Length 15.0-15.5 mm. Color- ation and structure similar to male except abdominal tergites 4-7 and terminalia or- ange-brown, tergites 1-4 with yellow pos- terior transverse margins. Material examined.— Holotype male, MEXICO: Acapulco, Guerrero, 17 IX 1941, Joseph D. Reed; paratypes, 2 females, same data as holotype. The holotype and para- types will be returned to the University of Colorado Museum, Boulder. Etyomology.— We take great pleasure in naming this species for Wendy Meyer, Col- orado State University, whose knowledge of entomology is inspiring. Remarks.—The male of N. wendyae can be distinguished from the similar appearing N. lamia and N. sponsor by the form of the aedeagus (Fig. 5). The female is also similar to N. sponsor and can be separated by the blackish brown distal flagellomeres and white setae of the face. Nemomydas fronki NEw SPECIES Fig. 6 Male.—Length 12.0-13.5 mm. Head shiny black, setae erect, black; antennae 3.3- 3.5 mm long, black; proboscis short, 0.9 times as long as subcranial cavity, black. Thorax shiny black, setae black, long and erect; wing light brown, veins blackish brown; halter black; legs black, setae long, black. Abdominal tergites black, bulla black, setae black, long, erect on tergites 1-3, long, black, recumbent on tergites 4-7; sternites black, setae black. Terminalia.— Black, dorsal digitate pro- cess of gonocoxite small, originating on in- VOLUME 92, NUMBER 3 ner surface of ventral digitate process; ae- deagus slender in lateral view (Fig. 6). Female.— Length 13-14 mm. Head shiny black, setae short, erect, black; antennae 3.5- 3.8 mm, black tinted with brown; proboscis short, 0.8 times as long as subcranial cavity, black. Scutum reddish brown, 3 faint black dorsal stripes, setae black, recumbent; wing light brown, darker brown tinting around blackish brown longitudinal veins; halter black; leg dark brownish black, setae black. Abdominal tergites 1-5 or 6 brownish or- ange, tergite 6 or 7 black, posterior margins of tergites 4-5 or 6 black, lateral margins of tergites 1-5 black, bulla black, setae very sparse, black, erect; sternites 1-7 blackish brown; terminalia black. Material examined.—Holotype male, TEXAS: Kenedy Co., 5 mi. S. 10-15 mi. E of Sarita, 25 V 1979, H. E. Evans, A. Hook, W. Rubink. Paratypes: 2 males, 2 females, same data as holotype. The holotype and one paratype female will be deposited in the CAS and the re- maining specimens in the Colorado State University Insect Collection. Eytomology.—We take great pleasure in naming this species for Dr. W. Don Fronk, Emeritus Professor of Entomology, Colo- rado State University. He has nurtured nu- merous students of entomology throughout his distinguished career. Remarks.—The male of N. fronkiis easily distinguished from all other species by its totally black coloration (including all setae) and distinctive small dorsal digitate process of the gonocoxite (Fig. 6). The brownish orange abdominal tergites with black pleu- ral margins easily separates the female from all other described females. Nemomydas tenuipes (Loew) Fig. 7 Midas tenuipes Loew, 1872: 61. Type lo- cality: California. Holotype male (MCZ #10654), examined. Leptomydas tenuipes, Johnson, 1926: 142. Nemomydas tenuipes, Hardy, 1950: 33. 477 Figs. 4-6. 4. Nemomydas sponsor. Male termi- nalia, a. lateral, b. ventral. 5. Nemomydas wendyae, n. sp. Male terminalia, a. lateral, b. ventral. 6. Nemo- mydas fronki, n. sp. Male terminalia, a. lateral, b. ven- tral. Remarks.—Only the male is known and Hardy (1950) presented an excellent de- scription. The abdomen ranges from black to reddish-black. Nemomydas tenuipes 1s readily distinguished from the only other black species, N. fronki, by dorsal digitate process of the gonocoxite originating at the base of the ventral digitate process of the gonocoxite (Fig. 7), yellow or white setae on the thorax and tergite 1, and its limited dis- tribution (California and Mexico (Baja Cal- ifornia)). Material examined.— Holotype male, CALIFORNIA: Kern Co., Edwards (MCZ #10654); San Francisco, Presidio Park, 13 V1 1981, W. J. Pulawski, 1 male (CAS); San 478 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-9. nalia, a. lateral, b. ventral. 8. Nemomydas bifidus. Male terminalia, a. lateral, b. ventral. 9. Nemomydas inton- sus. Male terminalia, a. lateral, b. ventral. 7. Nemomydas tenuipes. Male termi- Diego Co., Hot Springs Mountain Peak, 30 VI 1979, JWB, 1 male (SDNHM). MEXI- CO: Baja California (Norte), Vic. Fausino, San Juarez, 7 VI 1981, D. K. Faulkner and Brown, | male (SDNHM). Nemomydas bifidus Hardy Fig. 8 Nemomydas bifidus Hardy, 1950: 22. Type locality: California. Holotype male (CAS), examined. This species is only known from the ho- lotype male. It closely resembles N. intonsus and immaculate male variants (yellow brown tergites lacking the posterior trans- verse blackish brown bands) of N. pantheri- nus. It may be distinguished from N. inton- sus by the black setae of the legs and abdomen, and from N. pantherinus by the longer proboscis (at least 1.3 times the length of the subcranial cavity) and the stouter ae- deagus (Fig. 8). The lateral view of the ter- minalia illustrated by Hardy (1950, Fig. Se) is not accurate. Material examined.—Holotype male: CALIFORNIA: Riverside Co., Idyllwild, VI 1936, E. S. Ross (CAS). Nemomydas intonsus Hardy Fig. 9 Nemomydas intonsus Hardy, 1950: 27. Type locality: Pine Valley, California, Holo- type male (UK), examined. Remarks. — Hardy (1950) provides an ex- cellent description of this rare species, known only from the types, and need not be re- peated here. The proboscis is moderately developed, 1.3 times length of subcranial cavity. Material examined.— Holotype male: CALIFORNIA: San Diego Co., Pine Valley, 27 VI 1938, L. W. Hepner (UK); allotype female, same data as holotype (UK). Nemomydas fumosus Hardy, New STATUS Fig. 10 Nemomydas intonsus fumosus Hardy, 1950: 29. Type locality: San Diego Co., Cali- fornia, Holotype male (UK), examined. Hardy (1950) considered this species to be a variety of N. intonsus. It is considered here as a valid species, and can be separated from similar appearing relatives, N. bifidus and N. intonsus by the blackish brown fem- ora, tergites brownish black with lateral tri- angular whitish spots on tergites 3-5. The proboscis is 1.6 times length of the sub- cranial cavity. The terminalia of N. fumosus and N. intonsus are very similar. Material examined.— Holotype male, CALIFORNIA: San Diego Co., 7 VI 1929, VOLUME 92, NUMBER 3 P. W. Oman; allotype female, same data as holotype. Nemomydas pantherinus (Gerstaecker) Fig. 11 Leptomydas pantherinus Gerstaecker 1868: 85. Type locality: California. Holotype fe- male (Humboldt-Universitat), not ex- amined. Leptomydas pantherinus, Johnson, 1926: 142. Nemomydas pantherinus, Hardy, 1950: 30. Remarks.—This relatively common and quite variable species can be distinguished from other far western, primarily yellow or yellow brown species (N. intonsus and N. bifidus) by the short proboscis, 0.8-0.9 times as long as subcranial cavity. Material examined.—CANADA: British Columbia, Oliver, 22 July 1923, P. N. Vroon, 2 males, 4 females (CNC). MEXI- CO: Baja California Sur, 3 mi NE San Isidro (La Purisima), 2 IV 1985, Bloomfield and D. K. Faulkner, 1 female (SDNHM); Santo Domingo River, 3-4 VII 1979, Brown and D. K. Faulkner, | male (SDNHM); same except Santo Domingo (ruins), 1 male (SDNHM). UNITED STATES, CALI- FORNIA: Humboldt Co., Strong Std., 14 VIII 1938, B. P. Bliven, | female (CAS); Van Duzen River, 19 VII 1936, B. P. Bli- ven, 5 males, 5 females (CAS); Inyo Co., Hoton Creek Campground, Hwy 395, 4 VII 1981, R. M. Brown, | female (CAS); Los Angeles Co., N. Long Beach, 5 VIII 1938, A. Mallis, 1 female (CAS); Orange Co., Ir- vine, 23 VIII 1960, D. Magoi, 1 male (CAS); Santa Ana, 10 VIII 1964, J. Wilcox, | fe- male (CAS); Riverside Co., Herkey Creek, San Jacinto Mtns., 20 VI 1940, 1 male (CAS); Temecula, 30 VI 1956, J. Wilcox, 1 male, | female (CAS); Temecula, 4 VII 1950, J. W. MacSwain, 2 males (CNC); San Ber- nardino Co., Barstow, 24 VI 1914, J. R. Haskin, 1 female (CAS); El Cajon, 2 VII 1974, 1 female, 1 male (CAS); San Bernar- dino, 2 IX 1895, W. G. Wright, 1 female (CAS); Victorville, 2.5 mi. NW at Mojave, 479 Figs. 10-12. minalia, a. lateral, b. ventral. 11. Nemomydas panther- inus. Male terminalia, a. lateral, b. ventral. 12. Ne- momydas solitarius. Male terminalia, a. lateral, b. ven- tral. 10. Nemomydas fumosus. Male ter- 9 VIII 1983, D. Williams, 1 female (CAS); nr. Wrightwood, 21 VII 1954, F. M. Hull, 1 female (CNC); San Diego Co., Borden- fieldis Bay, 6 VIII 1982, B. Parks, 1 female (SDNHM); Lakeside, 20 VII 1965, J. Heppner, | male (FSCA); Mission Gorge (dam), 15 VII 1978, L. Guidry, 4 males (SDNHM); San Diego, 16 IX 1890, F. E. Blaisdell, 2 males, 1 female (CAS); San Die- go, 2-5 VIII 1954, H. E. and M. A. Evans, 1 male, 1 female (CAS); San Diego, 12-13 VII, W. S. Wright, 2 males, 1 female (CAS); Tulare Co., Porterville, 6 VIII 1959, E. Ball, 1 male (FSCA); same but 25 VII 1957, 1 female (FSCA); Springville, 25 VII 1957, E. 480 Ball 1 female (FSCA); Ventura Co., Foster Park, 1 VII 1959, J. L. Bath, 2 males, 1 female (CAS). Nemomydas solitarius (Johnson) Fig. 12 Leptomydas solitarius Johnson, 1926: 142. Type locality, Colorado, Holotype male (MCZ #7391), examined. Nemomydas solitarius, Hardy, 1950: 32. Hardy (1950) suggested that this species . may possibly be the same...” as N. pantherinus. Nemomydas solitarius is known only from the holotype male, apparently collected in “Col” with no further data. De- spite extensive collecting in Colorado, no additional specimens have been collected for study. It is more similar to N. bifidus, N. intonsus, and N. fumosus than N. pan- therinus, and can be distinguished from these species by the dorsal digitate process of the gonocoxite originating on the inner face of the ventral process (Fig. 12). The proboscis is missing from the holo- type. Johnson’s (1926) description omitted any reference to the relative length of this structure. Hardy (1950) evidently consid- ered the ‘“... mouthparts conspicuously short, scarcely, if at all, extended beyond the oral margin...” Material examined.—COLORADO: Ho- lotype male (MCZ #7391). Nemomydas yvenosus (Loew) Fig. 13 Midas venosa Loew, 1866: 15. Type local- ity: Texas. Holotype male (MCZ #10653), examined. Leptomydas venosus, Johnson, 1926: 142. Nemomydas venosus, Hardy, 1950: 34. Remarks.— The large thumb-like ventral process of the gonocoxite (Fig. 13) easily distinguishes the male of this widespread species. There are both light and dark phas- es of both sexes, especially with females. Pairs taken in copula have been mixed. Many of the past misidentifications, espe- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a 13 Fig. 13. Nemomydas venosus. Male terminalia, a. lateral, b. ventral. cially for N. brachyrhynchus and N. pan- therinus (see Curran (1965)), have been due to this color variability. The light form of the female may be confused for these two species and characters of separation are giv- en under N. brachyrhynchus. The dark form cannot be confused with any other species. Material examined. — MEXICO: Chihua- hua, 6 mi. S. Villa Matamoros, 21 VII 1967, R. C. Gardner, C. R. Kovacic, K. Lorenzen, 2 males, 2 females (CAS); Sinaloa, Baviri (playa) W Los Mochis, 9 IX 1986, D. K. Faulkner, Bloomfield, 1 male (SDNHM); Sonora, La Aduana, W. of Alamosa, 18 VIII 1964, T. E. Irwin, 1 male (CAS). UNITED STATES, ARIZONA: Cochise Co., Chiri- cahua Natl. Mon., 11 VIII 1962, J. Wilcox, 1 male (CAS); 3 mi. NE Coronado Natl. Mon., 17 VIII 1966, R. L. Westcott, 1 fe- male, | male (CAS); 8 mi. E Douglas, 4 VIII 1958, R. M. Bohart, 1 female (CAS); same except 8 VIII 1964, P. M. Marsh, | male; Montezuma Canyon, Huachuca Mtns., 19 VIII 1968, G. R. Ballmer, 1 female (CAS); Wilcox Dry Lake, 25 VIII 1967, F. G. An- drews, | male 3 females (CAS); same except E. I. Schlinger, 1 female 1 male (CAS); same except D. J. Culver, 1 male (CAS); Coconi- no Co., Oak Creek Canyon, 15 VI 1936, G. P. Engelhardt, 1 male (CAS); Gila Co., Globe, D. K. Duncan, | female (CSU); Pima Co., Florida Wash, 21 VIII 1979, D. K. Faulkner, 3 males (SDNHM). COLO- RADO: Phillips Co., Holyoke, 26 VII 1946, M. T. James, | male (CSU); Weld Co., Rog- gen, 21 VIII 1976, H. E. Evans, 1 male VOLUME 92, NUMBER 3 I ame 5 C0 Figs. 14, 15. Nemomydas bequaerti. Antenna. 15. Nemomydas brachyrhynchus. Antenna. (CSU); same except 24 VIII 1976, 2 males; same except 4 VIII. 1977, 1 male; same except 17 VIII 1982, 5 males, 4 females; Roggen, 8 IX 1933, M. T. James, | male (CSU); same except 31 VIII 1938, 9 males, 5 females; same except 15-18 VIII 1941, 1 male, | female; KANSAS, Kearny Co., Lak- in, 28 VIII 1951, R. R. Dreishbach, 1 male (UCB); NEW MEXICO, Chaves Co., 14 VIII 1955, R. R. Dreisbach, ! male (MSU); Grant Co., 29 VIII 1935, R. T. Kellogg, 3 males (CAS); Silver City, 14 IX 1935, B. T. Kellogg, | male, 1 female (CAS). TEXAS: Holotype male, Texas (MCZ #10653, ter- minalia missing); Jeff Davis Co., 24 mi. NW Ft. Davis, 24 XI 1965, R. W. Thorp, | fe- male (CAS); Kleberg Co., 20 mi. SE Kings- ville, 1 V 1985, W. J. Pulawski, 1 male (CAS); Riviera Beach, 28 V 1979, H. Evans, A. Hook, W. Rubink, | M (CSU). Nemomydas panamensis (Curran), nomen dubium Nomoneura panamensis Curran, 1934: 165. Type locality: Panama, Canal Zone, Bruja Point. Nemomydas panamensis, Papavero and Wilcox, 1968: 34.11. Curran (1934) never published a formal description for this name, and illustrated only the head. Papavero and Wilcox (1968) considered the name available (authors cit- ed Article 16 (vii) (1964 Code). However, this name was published in 1934, therefore does not satisfy Article 13 (1985 Code). An attempt was made to locate the two males 481 referred to by Papavero and Wilcox (1968). Curran was at the CNC from 1923 to 1928 and from 1928, to 1960 at the AMNH. Cu- rators of these museums (D. Grimaldi, AMNH) and B. E. Cooper, CNC) could not locate these two specimens in their respec- tive collections. We therefore consider this name as a nomen dubium. ACKNOWLEDGMENTS We would like to thank the following per- sons who made valuable material available for study: Paul H. Arnaud, Jr., California Academy of Sciences; M. Baylac, Museum National d’ Histoire Naturelle, Paris; Robert W. Brooks, University of Kansas; J. E. Chainey, British Museum (Natural Histo- ry); B. E. Cooper, Canadian National Col- lection; John T. Doyen, University of Cal- ifornia, Berkeley; David K. Faulkner, San Diego Natural History Museum; David A. Grimaldi, American Museum of Natural History; L. Matile, Museum National d’Histoire Naturelle, Paris; C. Riley Nelson, California Academy of Sciences; Carl Ol- son, University of Arizona; Christopher O’Toole, Hope Entomological Collections, University Museum, Oxford; R. V. Peter- son, National Museum of Natural History, Smithsonian Institution; Scott R. Shaw and C. Vogt, Museum of Comparative Zoology; Howard V. Weems, Florida State Collection of Arthropods; Michael Weissmann, Uni- versity of Colorado; Floyd G. Werner, Uni- versity of Arizona; and Ilan Yarom, Uni- versity of Kansas. David Carlson, Colorado State University prepared the illustrations. Special gratitude is expressed to Dr. and Mrs. Robert MacVean and Dr. Charles MacVean, Guatemala City for making the collecting in Guatemala possible. This manuscript was reviewed by Howard E. Ev- ans, Colorado State University, C. Riley Nelson and R. VY. Peterson. LITERATURE CITED Curran, C.H. 1934. The families and genera of North American Diptera. Ballou Press, New York, 512 pp. 1965. Family Mydidae (Mydaidae, Mydas- idae), pp. 357-360. Jn Stone, A., et al., eds., Cat- alog of the Diptera of America north of Mexico. U.S. Dept. Agric. Handbook 276. 1969 pp. Gerstaecker, A. 1868. Systematische Uebersicht der bis jetzt bekannt gewordenen Mydaiden (Mydasii Latr.). Settin Entomol. Ztg. 29: 65-103. Hardy, D. E. 1950. The Nearctic Nomoneura and Nemomydas (Diptera: Mydaidae). Wasmann J. Biol. 8: 9-37. Johnson, C.W. 1926. A revision of some of the North American species of Mydaidae. Proc. Boston Soc. Nat. Hist. 38: 131-145. Loew, H. 1866. Diptera Americae septentrionalis in- degena. Centuria septima. Berlin. Entomol. Ztschr. 10: 1-54. 1872. Diptera Americae septentrionalis in- digena. Centuria decima. Berlin. Entomol. Ztschr. 16: 49-115. 1886. Diptera, Vol. 1 (part), pp. 1-128. Jn Goodman, F. D., and O. Salvin, eds., Biologia Centrali-Americana. Zoologia-Insecta-Diptera 1. London. Papavero, N. and J. Wilcox. 1968. 34. Family My- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON didae (Mydaidae, Mydasidae), pp. 1-20. A cata- logue of the Diptera of the Americas south of the United States. Dept. Zool., Secr. Agric. Sao Paulo, Brazil. Séguy, E. 1928. Etude ser quelques Mydaidae nou- veaux Ou peu connus. Encycl. Entomol. Ser. II. Diptera 4: 129-156. Snelling, R.R. 1987. Geographical inexactitude. Pan- Pacific Entomol. 63: 339-340. Steyskal, G. C. 1956. The eastern species of Nemo- mydas Curran (Diptera: Mydaidae). Occ. Papers Mus. Zool. Univ. Michigan 573: 1-5. Welch, J. L. and B. C. Kondratieff. 1990. Review of the genus Mydas (Diptera: Mydidae). The xan- thopterus group of southwestern United States and Mexico. Ann. Entomol. Soc. Amer. 83: 142-148. Wilcox, J. 1981. Mydidae, pp. 533-540. In Mc- Alpine, J. F., et al., eds., Manual of Nearctic Dip- tera. Vol. 1. Res. Branch, Agric. Canada. Mongr. 27, Ottawa. Wilcox, J. and N. Papavero. 1971. The American genera of Mydidae (Diptera), with the description of three new genera and two new species. Archos Zool. Est. S. Paulo 21: 41-119. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 483-492 NEW NORTH AMERICAN COLOBAEA, WITH A PRELIMINARY ANALYSIS OF RELATED GENERA (DIPTERA: SCIOMYZIDAE) L. Knutson,! R. E. ORTH, AND R. ROZKOSNY (LK) Systematic Entomology Laboratory, Beltsville Agricultural Research Center, Ag- ricultural Research Service, USDA, Beltsville, Maryland 20705; (REO) Division of Bi- ological Control, Department of Entomology, University of California, Riverside, Cali- fornia 92521; (RR) Faculty of Science, J. E. Purkyné University, Brno, Czechoslovakia. Abstract. —Two new species of Colobaea (Diptera: Sciomyzidae) are described: C. can- adensis from Manitoba, Canada, and C. montana from Montana, United States. The male genitalia, wings, and antennae are illustrated; the distributions, along with that of C. americana Steyskal, are shown in a map; and a key to the world species is included. The major characters of the genera Colobaea, Pherbellia, Pteromicra, and Ditaeniella are compared and characterized as plesiomorphic or apomorphic, as an aid in determining the generic placement of the new species. Pherbellia trivittata (Cresson), P. parallela (Walker), and P. patagonensis (Macquart) are reassigned to Ditaeniella. Key Words: cation Although the classification of the Scio- myzidae has become well established over the past three decades, new species are con- tinually being discovered, particularly the smaller, rarer, and less conspicuous species. The genus Colobaea Zetterstedt consists primarily of small, black and yellow flies and includes the smallest (less than 2 mm) species of Sciomyzidae, Colobaea ameri- cana Steyskal (1954). Prior to this study one species was described from the Nearctic Re- gion and seven were described from the Pa- laearctic Region. There is one undescribed species from Pakistan, Afghanistan, and Iran and one from Nigeria. Species of Colobaea were among the first for which there was at least circumstantial ' Current address: USDA, ARS, Biological Control of Weeds Laboratory-Europe, American Embassy AGRIC, APO NY 09794. Flies, Ditaeniella, Pteromicra, Pherbellia, taxonomy, systematics, identifi- evidence (provided over 60 years ago) that the larvae of Sciomyzidae feed on mollusks. Lundbeck (1923) reared adults of C. pec- toralis (Zetterstedt) and C. punctata (Lund- beck) from puparia found in floating shells of small aquatic snails in Denmark. Roz- kosny (1967) provided brief descriptions of the puparia of Colobaea distincta (Meigen) and C. pectoralis, and Knutson et al. (1973) reared an undescribed species of Colobaea in Iran and presented the essential aspects of the life cycle (as C. iranica Knutson, no- men nudum). Knutson and Bratt (in prep- aration) have reared the Palaearctic species C. bifasciella (Fallén), C. pectoralis, and C. punctata, the Nearctic species C. ameri- cana, and the undescribed species from Iran through their complete life cycles and will describe all of their immature stages. Despite their apparent restriction to freshwater habitats where there are various small species of non-operculate snails, the 484 Table 1. Character states of major generic features of Colobaea and related genera. Colobaea Pteromicra Ditaeniella Pherbellia 1 +.(=) = = = 2 ree) = = = 3 + — (4+) — — eee eee a : aC) 5 =r) = = = 6 = + (-) - - 7 = (=) = = 8 = + = = 9 = ue); a = 10 = = TS) = 11 = = +f — (+) 12 7 = + = 13 = — + = List of characters (+ = apomorphic state, — = ple- siomorphic state; exceptions noted in parentheses): 1. Arista with some dorsobasal hairs slightly to much longer than ventrobasal hairs (same length in C. canadensis). 2. Vein Sc ending close to anterior branch of R, (except in C. canadensis). 3. Vein A, + CuA, evanescent apically (variable in Pteromicra leucothrix Melander). 4. Anterior surstylus with characteristic peglike processes [absent in C. canadensis, present in Pher- bellia mikiana (Hendel)). 5. Hypandrium with narrow, long posterior pro- cess (except in C. bifasciella). 6. Frons entirely shining (tomentose in an un- described species of Pteromicra from Japan and India). 7. Fore femur with pecten (absent in Preromicra anopla Steyskal). 8. Hypandrium with pair of pubescent lobes in posteroventral position. 9. Only one fronto-orbital seta [two pairs in Pre- romicra angustipennis (Staeger) and P. leucopeza (Meigen)]. 10. Prosternum haired (hairs absent in D. trivit- tata). 11. Inner posterior margin of hind coxa with sev- eral hairs (present in Pherbellia seticoxa Steyskal). 12. Anterior surstylus considerably reduced. 13. Paramere with conspicuous apical spines. species of Colobaea are broadly distributed. Colobaea is most abundant and species rich in Europe and species are also known from Central Asia, Pakistan, Iran, and northern Nigeria. In North America, Colobaea ranges from Montana to Manitoba, Quebec, and New York. In his recent review, Rozkosny (1984a) characterized the genus, described a new species from Sweden and Finland, and presented a key to the species. Roz- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON kosny (1984b) illustrated the terminalia of all the species known from Fennoscandia and Denmark. The male genitalia of C. punctata were illustrated by Rivosecchi and Prigioni (1980). To date, the only described Nearctic rep- resentative of Colobaea is C. americana Steyskal. The two North American species described below also apparently belong to Colobaea;, C. montana Knutson and Orth has all of the recognized apomorphic char- acteristics of the genus, but the placement of C. canadensis Knutson and Orth is less certain. Table 1 shows the character states of the major generic features (primarily as recognized by Rozkosny 1984b) of Colo- baeaand the three related genera, Pherbellia Robineau-Desvoidy, Pteromicra Lioy, and the recently resurrected Ditaeniella Sack (Rozkosny 1987); however, we have not been able to identify any synapomorphies for these four genera. Key TO SPECIES OF COLOBAEA ZETTERSTEDT (MODIFIED FROM ROZKOSNY 1984a) 1. Ground color of mesonotum brownish yellow; wing with two transverse dark bands (Palaearc- L1G) eek crashing ake See eee C. bifasciella (Fallén) Mesonotum black; wing without bands ...... 2 2. Proepisternum and anepisternum both, at least In part; VELOW, « ssscc: caretyce or eer Pe OO 3 — Proepisternum and anepisternum usually en- tirely black like rest of thorax .............. 7 3. Thorax mainly black, but with some yellow on proepisternum and anepisternum ........... 4 — Thorax more extensively yellow ............ 5 4. Dorsobasal hairs of arista distinctly longer than ventrobasal hairs (Nearctic) .. trobasal hairs (Nearctic) Sorat C. canadensis Knutson and Orth, n. sp. 5. Third antennal segment 3 times as long as width at base; both crossveins of wing conspicuously infumated (Palaearctic) ..... C. limbata (Hendel) — Third antennal segment 2 times as long as width at base; crossveins not infumated 6. Upper margin of anepisternum with complete dark stripe (Palaearctic) ...... cre SeN a ae AP ce C. pectoralis (Zetterstedt) — Upper margin of anepisternum only with rounded black spot below anterior notopleural seta (Palaearctic) C. punctata (Lundbeck) VOLUME 92, NUMBER 3 7. Third antennal segment 3 times as long as wide basally; both crossveins of wing distinctly in- fumated (Palaearctic) ....... C. beckeri (Hendel) — Third antennal segment 2 times as long as wide basally; crossveins not infumated .. . 8 8. Third antennal segment and arista black; an- terior margin of frons extensively yellow ... 9 — Third antennal segment white at base, arista white; frons entirely black (Palaearctic) C. distincta (Meigen) 9. Propleuron always black; only last tarsal seg- ment of foreleg white (Palaearctic) ....... BAe sctsbS.A evsinve Siu C. nigroaristata Rozkosny — Propleuron usually black, occasionally yellow- ish; last two tarsal segments of foreleg white (Nearctic) C. montana Knutson and Orth n. sp. Colobaea canadensis Knutson and Orth, NEw SPECIES Figs. 1-4, 13 Male.—Body length, 3.0 mm. Head slightly wider than high, width of gena about Y height of eye. Face and gena with short, whitish tomentum. Frons narrowed ante- riorly, yellowish tomentose, with many short, black bristles evenly distributed over anterior 4. Ocellar triangle shiny black, not extending as far as anterior end of shiny, brownish fronto-orbital plates. No orbito- antennal spot, very narrow stripe of fine, whitish tomentum extending along upper orbital margin and fading out near posterior fronto-orbital bristle. Pair of shallow depressions, with dense whitish tomentum, on either side of midline near top of occiput. Occiput blackish brown around foramen. Postgena yellowish to tan. Two pairs of rather strongly recurved fronto-orbital bris- tles, anterior pair slightly shorter than pos- terior pair; ocellar, postocellar, and inner and outer vertical bristles well developed and subequal in length. Many short, black setae on lower '2 of gena, finer anteriorly and extending onto lower parafacial. Short black setae also on ocellar triangle, along outer margin of fronto-orbital plates, and above occipital foramen. Lateral margins of occiput and postgena with several rows of irregularly dispersed, short, stout setae, no such setae mid-dorsally on occiput. Anten- nae light yellow to yellowish brown; seg- 485 ment 3 oval, darkened dorso-apically. Aris- ta brown, lighter basally, plumosity moderately dense, semi-recumbent, about as long as width of arista at base, dorsobasal hairs not setose or longer than ventrobasal hairs. Palpus yellowish white with many strong bristles, especially apically; labella yellowish. Thorax black with faint gray tomentum, postpronotal lobe slightly brownish. Pleura mostly brownish black, lower parts with strong whitish tomentum; ventral part of propleuron and anteroventral part of anepi- sternum yellowish. Thoracic chaetotaxy: | pro-episternal, | postpronotal, 1 presutural intra-alar bristle, 2 notopleural, | supra-alar, 2 subequal postalar, 2 dorsocentral (anterior pair slightly shorter, somewhat mesad of posterior pair and caudad of supra-alar), | weak prescutellar acrostichal behind dor- socentral, | basal scutellar, and | subequal apical scutellar. A few fine hairs posterior to pro-episternal seta. Anepisternum bare. Anepimeron with two fine bristles antero- ventrally, no vallar (subalar) bristles. Pos- terior half of katepisternum with dense coat of long, fine hairs over most of surface, sev- eral bristles along upper margin, two excep- tionally long hairs posterodorsally, well de- veloped bristles ventrally. Pro-episternum bare. Front coxa slightly less than *4 length of fore femur, yellow with whitish tomentum, several strong bristles below middle on ex- ternal margin and one or two near apex; midcoxa brownish dorsolaterally, yellowish ventrally; hind coxa yellowish, bare on pos- terodorsal surface. Fore femur robust, shiny black, with two irregular rows of 6-10 bris- tles on dorsal surface, bristles of anterior row larger than those of posterior row; mid- femur with anterior bristle beyond middle and three short hairlike bristles in row at posteroventral apex. Hind femur with two strong bristles posterodorsally, ventrally with dense coat of short bristles; fore tibia black except extreme base yellowish, with dense coat of greyish tomentum; mid and hind tibiae yellowish. Tarsi entirely yellow 486 except basal segment of fore tarsus darker on sides. Wing length, 2.8 mm. Membrane yellow- ish, hyaline; longitudinal veins yellow, crossveins darker and slightly infuscated. Pterostigma large, R, terminating beyond r-Mm crossvein, r-m crossvein at midlength of discal medial (dm) cell, vein A, + CuA, not reaching margin of wing. A few short setae on R,,; beyond r—m crossvein and on CuA, before dm-cu crossvein. Halter, ca- lypter, and calyptral ciliae yellowish white. Abdominal terga shiny dark brown with faint silvery tomentum, posterior margins yellowish. Bristles longest on posterolateral margins of fifth tergum. Syntergosternite 6- 8 well developed, anterior margin of sixth sternum darkly pigmented, extending across venter to dextral side of abdomen. Termi- nalia shiny blackish brown. Andrium as in Figs. | and 2. Ventral margin of epandrium not oblique posteriorly. Anterior surstylus flat, with one rounded, posterolaterally di- rected lobe and one rounded, ventrally di- rected lobe; without darkly pigmented, me- sal peglike processes. Posterior surstylus broad, scooplike, apex curved at almost right angle to meson; apices of posterior surstyli overlapping. Anterior end of hypandrium straight, broad. Ejaculatory apodeme equal in length to aedeagal apodeme, sinuate, plate rather small. Posterior arms of aedeagal apodeme broad, moderately elongate. Female.— Body length, 3.0-3.2 mm. Frons only slightly narrowed anteriorly. Subshiny orbital plates light brown, only slightly darker than yellowish frons. Occi- put brownish, cervical area yellowish. No strong hairs at base of arista. Postpronotal lobe yellow. Pro-epister- num extensively yellow below spiracle, an- episternum faintly yellow along upper part of posterior margin; anepimeron mottled yellow and brown. Prescutellar acrostichal bristles not larger than other acrostichals. Outer postalar a little larger than inner post- alar. Anepimeron with three or four fine bristles. Katepisternal hairs and _ bristles shorter, sparser than in male. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Abdominal terga lighter than in male, posterior margins narrowly to broadly yel- low. Cercus yellow. Midcoxa brownish to yellowish laterally. One specimen with three posterodorsal and two anterodorsal bristles apically on hind femur. Fore tarsus black to dark brown, api- cal segment lighter; mid and hind tarsi yel- low. Wing length, 2.9-3.1 mm. Diagnosis: Colobaea canadensis differs from all other species of Colobaea by the dorsobasal hairs on the arista being no long- er than the ventrobasal hairs, and the lack of peglike processes on the anterior sursty- lus. It most resembles C. bifasciella in that both species have a dull, tomentose, bris- tled, yellow frons; a similarly shaped head; two pairs of postalar bristles; and R, ex- tending to the level of the rm crossvein. These two species can be readily distin- guished, however; C. canadensis has a mainly shiny black body and unpatterned wings, whereas C. bifasciella has a yellow body with black markings and patterned wings. Type specimens.— Holotype male: Can- ada, Manitoba, Mile 505, Hudson Bay Ry.; 13 June 1952; J. G. Chillcott; ecological data F-D; in Canadian National Collection, Ot- tawa. Allotype: female, Canada, Manitoba, Mile 504, Hudson Bay Ry.; 21 June 1952; J. G. Chillcott; ecological data F-H, along RR; in Canadian National Collection. Para- type: female, Canada, Manitoba, Warkwork Cr. nr. Churchill; 7 July 1952; J. G. Chill- cott, ecological data F-D; LVK Slide Nos. 7044 and 7055, in United States National Museum of Natural History. Distribution map, Fig. 13. Colobaea montana Knutson and Orth, NEw SPECIES Figs. 5-13 Male (female unknown): Body length, 3.0 mm. Head 4 narrower than height; height of gena about 4 height of eye. Face shiny yellow, narrow, with strong median carina; clypeus rounded and somewhat produced 4 ae 0.4 mm Figs. 1, 2. Colobaea canadensis, holotype male. 1, Postabdomen, sinistral view, inverted. 2, Postabdomen, ventral view. Figs. 3, 4. Colobaea canadensis, paratype female. 3, Left antenna. 4, Right wing. 488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 3 anteriorly. Gena yellowish white. Frons strongly narrowed just above antenna, densely matt-tomentose, mostly black, yel- lowish anteriorly, yellow area extended tr- angularly toward anterior ocellus. Ocellar triangle subshiny black, not prolonged an- teriorly; fronto-orbital plates shiny black; tomentose whitish patch between base of antenna and orbital plate. Occiput shiny black over most of surface; postgena whitish yellow. Two subequal, erect fronto-orbital bristles; ocellars and inner and outer ver- ticals well developed, longer than fronto- orbitals; postocellars smaller than fronto- orbitals. Short, black setae in two irregular rows along lower margin of gena extending along groove between gena and face to an- teroventral angle of eye; on anterolateral edge of yellowish part of frons; sparsely be- tween ocellar and postocellar bristles; and in patch above foramen. Upper occipital margin with weak setae only. First and sec- ond antennal segments yellow; second seg- ment with weak bristles on dorsal edge, around apical margin, and several strong bristles ventro-apically; third segment ovoid, black, slightly brownish at base of arista on external surface, more extensively yellowish brown basally on inner surface. Arista brown with a few short, rather thick hairs; some dorsobasal hairs slightly spinose and longer than ventrobasal hairs; all hairs shorter than width of arista at base. Palpus yellowish white, with a few weak bristles; labella yel- lowish white. Thorax including scutellum entirely black except in the Swan Lake specimen, pro- pleuron yellowish; no silvery tomentum. Prosternum yellow. Thoracic chaetotaxy: 1 short pro-episternal, | postpronotal, | post- = 489 humeral, 2 notopleural, | supra-alar, | post- alar, 2 dorsocentral (anterior pair slightly shorter), | weak prescutellar acrostichal, | basal scutellar, 1 apical scutellar. Anepister- num bare, anepimeron with three or four fine bristles anteriorly, no vallar bristles. Katepisternum with one strong bristle mid- dorsally and a few hairs posteroventrally, well developed bristles midventrally. Pro- sternum bare. Fore coxa 7 length of fore femur, sub- shiny yellowish white, one or two strong bristles below middle on external margin and several apically; mid and hind coxae yellowish, darker dorsally; hind coxa bare above. Fore femur robust, basal '2 to 74 yel- lowish, remainder black, more extensively darkened along dorsal surface, two irregular rows of 5-8 bristles on dorsal surface; mid- femur yellow, one anterior bristle beyond middle, three hairlike bristles in row at pos- teroventral apex; hind femur mostly yellow- ish, brownish apically, one strong bristle an- terodorsally; ventrally with double row of widely spaced bristles. Fore tibia black, mid and hind tibiae yellow. Fore tarsus with bas- al segment black, segments 2 and 3 brown- ish, segments 4 and 5 yellowish; mid and hind tarsi yellowish, apical segment darker. Wing length, 1.9-2.1 mm. Membrane grayish hyaline, longitudinal veins yellow- ish to brownish, crossveins not infuscated. Pterostigma small, Sc ending close to R,; R, terminating basad of crossvein r-m; r-m slightly beyond midlength of cell dm; A, + CuA, extending almost to margin of wing. Halter, calypter, and calyptral ciliae yellow- ish white. Abdominal terga and sterna shiny black, posterior margins of posterior sterna nar- Figs. 5, 6. Colobaea montana, holotype male. 5, Postabdomen, sinistral view, inverted. 6, Postabdomen, ventral view. Figs. 7-12. Colobaea montana, paratype male, 4 miles east of Bigfork, Montana. 7, 8, Posterior surstylus, sinistral view, inverted. 7, Outer surface. 8, Inner surface. 9, Left antenna. 10, 11, Anterior surstylus, sinistral view, inverted. 10, Outer surface. 11, Inner surface. 12, Right wing. 490 . CANADENSIS - MONTANA 1000 MILES KILOMETERS LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION Fig. 13. rowly yellow; only posterior terga with a few strong, black bristles. Syntergosternite 6-8 well developed; anterior margin of sixth sternum darkly pigmented, extending across venter to dextral side ofabdomen. Andrium as in Figs. 5 and 6. Epandrium with ventral margin oblique posteriorly, with lobelike process midlaterally on posterior margin. Anterior and posterior surstyli widely sep- arated. Anterior surstylus clublike, curved mesally and approximate, posteromesal margin and posterodorsal surface with dense coat of short, thick, peglike processes. Pos- terior surstylus as in Figs. 7 and 8. Anterior end of hypandrium broad, rounded. Ae- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON CHICAGO HENRY M LEPPARD, EDITOR Collection sites for Colobaea americana, C. canadensis, and C. montana. deagal apodeme lightly pigmented, broad- ened posteriorly, twice as long as ejaculatory apodeme. Diagnosis: Witha mostly matt black frons and R, terminating considerably basad of rm, C. montana is more similar to C. americana, C. nigroaristata Rozkosny, C. punctata, and related species than it is to C. bifasciella and C. canadensis. Although in the key to species, which is based on exter- nal characters, C. montana and C. nigro- aristata run out in the same couplet, males can be readily separated by the internal structures of their terminalia—most nota- bly the shape of the posterior surstylus (see VOLUME 92, NUMBER 3 Rozkosny 1984a). The shape of the poste- rior surstylus, short dorsobasal hairs on the arista, and lack of silvery tomentum along the margin of the eye distinguish C. mon- tana from C. americana, the only other species of Colobaea likely to occur with C. montana. Type specimens.— Holotype male and 2 male paratypes: United States, Montana, 4 miles east of Bigfork; 29 July 1965; B. A. Foote; one male paratype: Montana, 20 mi. south of Swan Lake, 20 August 1968; B. A. Foote. All specimens in the United States National Museum of Natural History. Dis- tribution map, Fig. 13. COMPARISON OF COLOBAEA WITH OTHER GENERA A comparison of the character states of major generic features of Colobaea and the related genera Pteromicra, Ditaeniella, and Pherbellia is shown in Table 1. Pherbellia is probably polyphyletic. In the process of making these comparisons, it became clear that one group of “Pherbellia” species should be transferred to the genus Ditaeniella. Di- taeniella was proposed by Sack (1939) for the Palaearctic species Sciomyza grisescens Meigen. Most subsequent authors placed S. grisescens in Sciomyza or Pherbellia. Stey- skal (1963) noted the similarities (especially in the male genitalia) of S. grisescens, S. humilis Loew (Nearctic) (= S. parallela Walker), and S. patagonensis Macquart (Neotropical) and placed these species in the Pherbellia grisescens group. He noted, “The andrium is of a rather special type in the Sciomyzinae and may be the basis for even- tually segregating the group nomenclatur- ally from Pherbellia.” Rozkosny (1987) res- urrected Ditaeniella as a valid genus, including S. grisescens. We propose the fol- lowing new combinations: Ditaeniella parallela (Walker) 1853 (Nearc- tic and Neotropical). Ditaeniella patagonensis (Macquart) 1851 (Neotropical). 491 Ditaeniella trivittata (Cresson) 1920 (Nearc- tic). ACKNOWLEDGMENTS The contributions of the following per- sons are gratefully acknowledged. Speci- mens were provided by B. A. Foote (Kent State University, Kent, Ohio), and H. J. Teskey (Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario). The manuscript was reviewed by B. A. Foote; S. E. Neff, (Temple University, Philadelphia, Pennsylvania); and W. L. Murphy, A. L. Norrbom, and R. V. Peterson (Systematic Entomology Laboratory, USDA, Beltsville, Maryland). Geographic information was provided by R. V. Peterson. LITERATURE CITED Cresson, E. T., Jr. 1920. A revision of the nearctic Sciomyzidae (Diptera, Acalyptratae). Trans. Am. Entomol. Soc. 46: 27-89, 3 pls. Knutson, L. and A. D. Bratt. In preparation. Biology and immature stages of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae). Knutson, L. V., E. R. Shahgudian, and G. H. Sahba. 1973. Notes on the biology of certain snail-killing flies (Sciomyzidae) from Khuzestan (Iran). Iran. Jour. Publ. Hlth. 2(3): 145-155. Lundbeck, W. 1923. Some remarks on the biology of the Sciomyzidae, together with the description ofa new species of Crenu/us from Denmark (Dipt.) Vidensk. Meddr. Dansk Naturh. Foren. 76: 101- 109. Macquart, J. 1851 [1850]. Diptéres exotiques nou- veaux ou peu connus. Suite de 4° supplement. Mem. Soc. R. Sci., Agr. Arts, Lille 1850. 134-294, pls. 15-28. Rivosecchi, L. and C. Prigioni. 1980. Sciomyzidae (Diptera, Acalyptera) della collezione “Corti” ed ambienti umidi naturali della Provincia di Pavia. Boll. Mus. Civ. St. Nat. Verona 7: 675-689. Rozkosny, R. 1967. Zur Morphologie und Biologie der Metamorphose-stadien mitteleuropaischer Sciomyziden (Diptera). Acta Sci. Nat. Brno. 1: 117- 160. . 1984a. Review of Colobaea Zetterstedt (Dip- tera: Sciomyzidae), with a new species from north- ern Fennoscandia. Entomol. Scand. 15: 85-88. 1984b. The Sciomyzidae (Diptera) of Fen- noscandia and Denmark. Fauna Entomol. Scand. 14: 1-224. 492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1987. A review of the Palaearctic Sciomy- _———. 1963. Taxonomic notes on Sciomyzidae zidae (Diptera). Folia Fac. Sci. Nat. Univ. Purk. (Diptera, Acalyptratae). Pap. Mich. Acad. Sci., Arts Brun. 86: 1-100 + 58 pls. Lett. 48: 113-125. Sack, P. 1939. Sciomyzidae, 87 pp. Jn Lindner, E., Walker, F. 1853. Diptera, Part IV, pp. (253)}-414, ed., Die Fliegen der palaearktischen Region, Lief. pls. 7-8, cont. 7n Saunders, W. W., ed., Insecta 125, Parts 1, 2, 3 (Fam. 37). E. Schweitzerbart. Saundersiana: Or characters of undescribed in- Stuttgart. sects in the collection of W. W. Saunders, Esq. Steyskal, G.C. 1954. Colobaea and Hedria, two gen- Van Voorst. London. era of Sciomyzidae new to America (Diptera: Acalyptratae). Can. Entomol. 86: 60-65. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 493-496 A NEW SPECIES OF DIORYCTRIA (PYRALIDAE: PHYCITINAE) FROM MEXICO H. H. NEUNZIG Department of Entomology, North Carolina State University, Raleigh, North Carolina 27695-7613. Abstract. —Dioryctria cuitecensis n. sp. from the state of Chihuahua is described. A habitus photograph of the male holotype, and drawings of the male and female genitalia of D. cuitecensis, and a key to Mexican species of Dioryctria are included. Key Words: Larvae of all species of Dioryctria Zeller (Pyralidae) are associated with conifers. Be- cause pines and their relatives are a com- mon group of trees in many parts of Mexico, numerous species of Dioryctria may be ex- pected in this part of North America. As recently as the middle of this century, how- ever, only one species of these phycitines was recorded from the Republic (Heinrich 1956), probably because of limited collect- ing. Within the last several decades ento- mologists from the United States and Can- ada have more extensively light trapped in Mexico, and thereby greatly increased the number of Mexican Lepidoptera in collec- tions. In addition, the Mexican Division of Forest Sciences, in cooperation with the Forest Service of the United States Depart- ment of Agriculture, has made a concerted effort to collect and rear cone insects in Mexico (Hedlin et al. 1981, Cibrian-Tovar etal. 1986). Asa result, the number of known species of Dioryctria in Mexico increased to ten. Recently, the following additional Mex- ican species in the genus has come to my attention. Dioryctria cuitecensis, NEW SPECIES (Figs. 1, 2-4) Diagnosis. — D. cuitecensis 1s a large, dark, rather uniformly marked species (Fig. 1). taxonomy, Dioryctria cuitecensis, Dioryctria key The transverse lines are very obscure, con- sisting of only a very few white-tipped scales, the discal spot is only slightly paler than the surrounding scales, and the black scales that characteristically accent the transverse lines are for the most part diffuse. The male and female genitalia of D. cuitecensis are similar to those of D. cambiicola Dyar, a species occurring in the United States, however, the forewing of the latter species has distinct silvery white transverse lines accented with adjacent patches of black, and distinct patches of silvery white scales inside the subbasal scale ridge, on the discal spot, be- tween the discal spot and the postmedial line, and just before the terminal line. Description.—Head: Frons and vertex brown, fuscous or reddish brown. Labial palpus reaching above vertex in both sexes, mostly brown to fuscous with very few white-tipped scales. Maxillary palpus squa- mous, mostly brown and fuscous. Antenna of male filiform with abundant, short sen- silla trichodea. Collar: Brown or reddish brown. Thorax: Dorsum brown to reddish brown, in part suffused with fuscous or black. Forewing: Length 15.5-17.0 mm; above with distinct, strongly raised scales forming subbasal and antemedial patches; postme- dial patch of raised scales also present but scales noticeably less elevated than those of 494 Fig. 1. basal patches; additional smaller patches of raised scales at base and at discal spot; ground color brown; antemedial line very obscure, usually consisting of a few pale brown or white-tipped pale brown scales in inner half; postmedial line also very weak; diffuse, indistinct patches of fuscous and black scales in median area and along costa; numerous red, orangish red or reddish brown-tipped scales in basal, subbasal, in- ner median and terminal area (usually a dis- tinct rust color in basal and subbasal area including the raised subbasal patch of scales); discal spot obscure, only slightly more pale than surrounding scales; terminal line fus- cous to black; undersurface of male with short, basal, pale gray subcostal streak. Hindwing: Above, dark brownish gray. Male genitalia: (Figs. 2, 3) similar to male geni- talia of Dioryctria cambiicola except juxta of D. cuitecensis noticeably smaller than that PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Dioryctria cuitecensis, 6, Holotype. Scale line = 1.0 mm. of D. cambiicola. Female genitalia: (Fig. 4) similar to female genitalia of D. cambiicola. Type material. — Holotype male, Cuiteco, Chih, Mex, VIII 27 1969, T. A. Sears, R. C. Gardner, C. S. Glaser, genitalia slide 2090 HHN, deposited in the Bohart Museum of Entomology, University of California, Da- vis. Paratypes: three females, same collec- tion data as for holotype except IX 3 1969, IX 8 69, genitalia slide 2091 HHN. Para- types deposited in collections of University of California, Davis, National Museum of Natural History, Washington, D.C. and North Carolina State University, Raleigh. Distribution and life history.—Known only from southwestern Chihuahua, Mexi- co. Host plant(s) and behavior of larvae, unknown. Remarks.—D. cuitecensis belongs to the zimmermani group of Mutuura and Mun- roe (1972). VOLUME 92, NUMBER 3 495 Figs. 2-4. Dioryctria cuitecensis. 2, Male genitalia (most of left valva and aedeagus omitted). 3, Aedeagus. 4, Ductus bursae and corpus bursae of female genitalia. Scale lines = 1.0 mm. Key TO MEXICAN SPECIES OF DIORYCTRIA 1. Forewing without raised scales — Forewing with raised scales 2. Forewing with many red or orange scales; val- va with distal part blunt and short, at most slightly falcate; ductus bursae with sclerotized part usually shorter than length of corpus bur- SAC a iterate Tete ere enter ee eee rce — Forewing with only few red or orange scales; valva with distal part distinctly produced api- cally; ductus bursae with sclerotized part about as long as, or longer than, length of corpus Forewing without antemedial line . . ross! Munroe . Forewing heavily dusted with white (moth ap- pearing mostly gray); male antenna weakly SCITALG tar ss he na ae ate pinicolella Amsel Forewing lightly dusted with white (moth ap- pearing mostly brown or fuscous); male an- tenna strongly serrate (appearing almost uni- pectinate) .s<2 ce aeeeee eee mayorella Dyar 496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Uncus constricted at base with distinct lateral protuberances; ductus bursae with proximal part distinctly narrower than distal part ... 6 Uncus not constricted at base and without lateral protuberances; ductus bursae with proximal part about as wide as, or wider than, distal part ..... . Valva with dorsal part broad distally (about 4x as wide as more setiferous ventral part); ductus bursae with distal end simple, well sep- arated from proximal spines of corpus bursae Valva with dorsal part more narrow distally (about 2—2.5 x as wide as more setiferous ven- tral part); ductus bursae with distal end with attached sclerite supporting proximal spines of corpus bursae Forewing mostly reddish Browne ealva with distal part wide and blunt; ductus bursae with sclerotized part about as long as corpus bursae eh Ose ee erythropasa (Dyar) Forewing with few reddish brown scales; val- va with distal part produced into a slender curved hook; ductus bursae with sclerotized part longer than corpus bursae ; martini Mutuura and Neunzig : Forewing gray, densely dusted with white and with distinct transverse lines Be ee eR Aa: ip a a gt albovittella (Hulst) Forewing brown, with only very few scales dusted with white and without distinct trans- verse lines (Fig. 1) .. cuitecensis Neunzig Forewing with narrow, interrupted, black lon- gitudinal line extending most of length of wing (starts near base at CuA, and extends through discocellular area); valva with dorsal projec- tion extending well beyond apex of ventral part of valva cibriani Mutuura and Neunzig Forewing without black longitudinal line ex- tending most of length of wing; valva with dorsal projection barely exceeding apex of ventralipartiofevalVa\ semaine c eee cence 10 Forewing ground color pale gray rand with strongly contrasting black subbasal scale ridge, black outer border of antemedial line, black inner border of postmedial line, and small black scale ridge following antemedial line; ductus bursae with proximal part bulbous . . 5 Ie as DOOR Semen bE subtracta Heinrich — Forewing ground color brown, without black strongly contrasting scales associated with scale ridges and transverse lines; ductus bursae with proximal part not strongly expanded durangoensis Mutuura and Neunzig ACKNOWLEDGMENTS Specimens of D. cuitecensis were sent to me, as part of a loan of phycitines, by A. Porter and R. Schuster of the University of California, Davis. Research funds for this study were provided in part by the USDA, Forest Service. This is paper No. 12186 of the Journal Series of the North Carolina Ag- ricultural Research Service, Raleigh, North Carolina 27695-7643. LITERATURE CITED Cibrian-Tovar, D., B. H. Ebel, H. O. Yates, and J. T. Méndez-Montiel. 1986. Insectos de conos y sem- illas de las coniferas de Meéxico/Cone and seed insects of the Mexican conifers. U.S. Dept. Agr. Gen. Tech. Rep. SE-40. 110 pp. Hedlin, A. F., H. O. Yates, D. Cibrian-Tovar, B. H. Ebel, T. W. Koerber, and E. P. Merkel. 1981. Cone and seed insects of North American conifers. Joint Publication: Environ. Can., Can. For. Ser- vice; U.S. Dept. Agr., For. Service; Sec. Agr. Re- cursos Hidraulicos, Mexico. 122 pp. Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus. Bull. 207. 581 pp. Mutuura, A. and E. Munroe. 1972. American species of Dioryctria (Lepidoptera: Pyralidae) HI. Group- ing of species: species of the auranticella group, including the Asian species, with the description of a new species. Can. Entomol. 104: 609-625. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 497-511 FINE STRUCTURE OF THE EGGS OF PSOROPHORA COLUMBIAE, PS. CINGULATA AND PS. FEROX (DIPTERA: CULICIDAE) JOHN R. LINLEY' AND DAVE D. CHADEE? 'Florida Medical Entomology Laboratory, Institute of Food and Agricultural Sciences, University of Florida, 200 9th Street S.E., Vero Beach, Florida 32962; ?Insect Vector Control Division, Ministry of Health, 3 Queen St., St. Joseph, Trinidad, West Indies. Abstract. —The eggs of Psorophora columbiae, Ps. cingulata and Ps. ferox are described with reference to scanning electron micrographs. In contrast to earlier descriptions, com- plete details of the Ps. columbiae and Ps. ferox eggs are given, including the anterior end and micropyle, posterior end, and intact outer chorion. This is the first account of the egg of Ps. cingulata. Eggs of Ps. ferox from Florida are discernibly different from a Trinidad population in terms of the number and form of the small outer chorionic tubercles in each chorionic cell. Key Words: The known distribution of Psorophora (Grabhamia) cingulata (Fab.) extends through Trinidad, possibly Central America and much of South America (Knight and Stone 1977). Itis a species about which little is known, except for a number of records of the larval habitat (Heinemann et al. 1980). Its egg is described here for the first time. Psorophora (Janthinosoma) ferox (von Humboldt) is a very widespread species, ranging from South through Central Amer- ica, the Greater and Lesser Antilles, the eastern United States and south-eastern Canada. The egg of this species was first described by Horsfall et al. (1952), from eggs stripped of the outer chorion. Subsequently, using eggs prepared in the same way, Hors- fall et al. (1970) published scanning electron micrographs showing the inner chorionic sculpturing. Although eggs lacking the outer chorion are adequate to demonstrate the ba- sic Outline pattern of the outer chorionic cells, the entire structural detail of the outer chorion is lost. In this paper we provide the first description of the intact egg, including Insecta, mosquito, egg, fine structure, chorionic structure details of the posterior end and anterior end and micropyle. The description is based on eggs from Ps. ferox collected in Florida, but we also describe and illustrate differences in eggs from a Trinidad population. In their papers cited above, Horsfall and associates described (as Ps. confinnis) the egg of Ps. (Grabhamia) columbiae (Dyar and Knab). Their material was collected from the eastern United States, which would im- ply that it is Ps. columbiae as currently rec- ognized (Belkin et al. 1970, Darsie and Ward 1981). Bosworth et al. (1983) examined Ps. columbiae eggs in more detail as part of their study of eggs of the Ps. confinnis complex from 7 areas of the country. However, as in the work by Horsfall and co-workers, the outer chorion was first removed, then the sculpturing of the inner chorion in the an- terolateral part of the egg was compared by scanning electron microscopy (SEM). None of these studies, therefore, has presented a description of the egg in its natural form (outer chorion intact), or of all its parts. Fol- lowing the more complete account in this 498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ps. columbiae. Entire egg, anterior end up- permost. Scale = 100 um. paper, we point out some characters of the intact egg that may be of value in evaluating a species complex. MATERIALS AND METHODS Eggs of Ps. cingulata and of Ps. ferox were obtained from females collected in the vi- cinity of Arena, Trinidad, blood-fed and then induced to oviposit in the laboratory. Eggs were allowed to embryonate and were then preserved in 30% ethanol, pre-filtered through a 0.45 um polycarbonate mem- brane, and sent to Vero Beach. On arrival, the eggs were pipetted onto small circles of filter paper and air-dried. Psorophora ferox eggs from Vero Beach were obtained from females collected on the grounds of the Florida Medical Entomology Laboratory. Eggs of Ps. columbiae were from a small laboratory colony established from material collected within a few miles of Vero Beach. When fully embryonated, eggs of both species were suspended in cold, filtered water long enough to pipette them onto cir- cles of filter paper, where they were air-dried. For all three species the paper circles were then attached to stubs with silver paint, completely desiccated over calcium chlo- ride, then coated with gold. Specimens were examined in a Hitachi S-510 SEM. Measurements of living eggs were done with a stereomicroscope and ocular mi- crometer. Dimensions and ranges of struc- tures on the egg surfaces were determined from inspection of 5 eggs in each case. The terminology follows Harbach and Knight (1980), except for the terms “anterior ring” and “outer chorionic cell field,’ defined by Linley (1989). RESULTS Psorophora (Grabhamia) columbiae (Figs. 1-3) Size: dimensions as in Table 1. Color: dull black. Overall appearance: broadly banana- shaped, dorsal curvature greater than ven- tral, widest at about anterior ', posterior portion gradually tapered, anterior taper VOLUME 92, NUMBER 3 499 Table 1. Dimensions of eggs of three species of Psorophora (n = 10). Length um Width pm L/W ratio Species Mean + SE Range Mean + SE Range Mean + SE Range Ps. columbiae 860.9 + 17.8 736.4-936.4 235.5 + 3.4 218.2-254.5 3.66 + 0.05 3.24-3.81 Ps. cingulata 76231, = 1:8 749.9-766.6 240.1 + 1.8 233.3-249.9 3.18 + 0.03 2.99-3.29 Ps. ferox (Fl) 904.5 + 13.7 845.5-963.6 250.2 + 3.6 227.9-265.5 3.62 + 0.03 3.44-3.75 Ps. ferox (Tr.) 918.3 + 1.9 911.0-927.7 S160 1.7 3111-32222 2.90 + 0.02 2.83-3.04 more abrupt and pointed, overall surface appearing spiny, micropylar collar distinct (Fig. 1). Outer chorionic cells longitudinally elongated, each with many small peripheral tubercles and a single large, elongate, an- teriorly inclined tubercle (Fig. 1). Chorion, dorsal, lateral and ventral sur- faces: all surfaces very similar (Fig. 1). Cho- rionic cell structure typified by mid-lateral cells, middle of egg, as follows. Shape most- ly hexagonal, occasionally pentagonal, out- lines quite variable (Fig. 2e), length 17-26 um, width 11-18 wm. Cell fields about 2 um smaller in each dimension. Each cell with a single, large, elongate, anteriorly inclined tubercle, positioned towards posterior part of cell, and a variable number of small tu- bercles around cell periphery (Figs. 2e, 3e; Table 2). Size distribution (largest diameter) of small tubercles bimodal (Fig. 4), size in- crease highly significant from posterior to anterior part of cell (Fig. 5). Each large tu- bercle consisting of a distinct base, upper part of tubercle smooth (but see variations, below), often inclined upward only in an- terior 12, length 7-12 wm, width 3-5 um. Cell floor covered with tiny, more or less round tubercles (Fig. 3e), diameter 0.1—-0.2 um, fewer posterior to large tubercle. Small tubercles tending to be oval, with flared base and distinct upper portion, entire tubercle slightly elongated in same axis as adjacent outer chorionic reticulum (Fig. 3e, f). Cho- rionic reticulum a fine meshwork, width 1.9- 2.7 um, with central line of closely spaced or fused bead-like protuberances, diameter 0.3-0.6 um (Fig. 3f). Outer chorionic cell structure differs from anterior to posterior part of egg. Outer cho- rionic cells close to anterior end smaller, length 16-21 wm, width 9-13 um, large tu- bercles flat or not as erect, often pointed, with rough upper surfaces (Fig. 3a), small tubercles more irregular. At middle of egg large tubercles more tongue-shaped, almost always erect, smooth, small tubercles more regular (Fig. 3b). Approaching posterior end chorionic cells smaller, length 15-19 um, width 13-19 um, large tubercles smaller (Fig. 3c). Cells very close to posterior end even smaller, large tubercles much shorter, cap- like, small tubercles fewer (Fig. 3d). Anterior end, micropyle: outer chorionic cells as described (Fig. 3a), anterior ring present, diameter 41-44 um, width 3-9 um, outer margin with anteriorly curved points (Fig. 2a). Micropylar collar very distinct, almost always complete (no gaps), flared an- teriorly (Fig. 2a), height 11-12 um, outer diameter about 26 um, outer margin irreg- ular (Fig. 2b, c), wall width variable, 1.5-8 um, inner diameter about 14 um, inner edge slightly irregular (Fig. 2c). No micropylar disc visible, micropyle diameter about 2.5 um. Posterior end: outer chorionic cells close to end as described (Fig. 3d), end rounded, large tubercles in cells button-like (Fig. 2d). Psorophora (Grabhamia) cingulata (Figs. 6-8) Size: dimensions as in Table 1. Color: dull black. Overall appearance: broadly cigar-shaped, dorsal curvature greater than ventral, an- terior end somewhat conical, posterior more rounded, surface somewhat spiny but less so than Ps. columbiae, micropylar collar sometimes fairly distinct (Fig. 6), but often not so (Fig. 7a). Outer chorionic cells as in 500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Ps. columbiae. (a) anterior end, dorsal surface; (b) anterior end and micropylar apparatus; (c) detail of micropylar apparatus; (d) posterior end, lateral view; (e) typical outer chorionic cells, lateral surface, middle of egg. Scale = 20 um. VOLUME 92, NUMBER 3 501 Fig. 3. Ps. columbiae, details of lateral outer chorionic cells on different parts of egg (figure letters correspond to positions labelled in Fig. 1). (a) close to anterior end; (b) at about anterior 1/3; (c) at about posterior 1; (d) very close to posterior end; (e) detail of whole cell, middle of egg; (f) detail of outer chorionic reticulum, middle of egg. Scale = 10 um (a, b, c, d, e), = 5 um (f). 14 Ps. columbiae as oN Ps. cingulata 8 6 4 2 0 8 it, 6 5 4 3 2 1 0 Number 14 Ps. ferox Trinidad oN F&F DD @ Tubercle diameter (pm) Fig. 4. Size distributions of small outer chorionic tubercles. Ps. columbiae except that small, peripheral tubercles more numerous (Table 2) and large tubercle sometimes anteriorly inclined, sometimes decumbent (Fig. 6). Chorion, dorsal, lateral and ventral sur- faces: all surfaces very similar (Fig. 6). Outer chorionic cells usually hexagonal, occasion- ally pentagonal, outlines variable (Fig. 7e), length 25—43 um, width 21-25 wm. Cell fields about 2 um smaller in each dimension. Sin- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4.5 Ps. columbiae : b = +0.281 (P<0.001) Length (pm) ne nT enh Se ak? Tubercle number —— posterior anterlor—s Fig. 5. Regressions of small tubercle length (largest dimension) on position in outer chorionic cell (number of tubercle indicates its numerical position relative to posterior cell boundary). gle large tubercle positioned well towards posterior part of cell, often inclined ante- riorly, but often flat, owing to attachment to cell floor at anterior end (Fig. 7e). Small tubercles variable in number (Table 2), size distribution (largest diameter) skewed (Fig. 4), size increasing significantly from poste- VOLUME 92, NUMBER 3 Table 2. Numbers of small outer chorionic tuber- cles in three species of Psorophora. Number of small tubercles* Species n Mean + SE Range Ps. columbiae 49 19 E1053 7-16 Ps. cingulata 31 25.0 + 0.4 20-29 Ps. ferox (FL) 36 32:9 + 127 17-51 Ps. ferox (Tr.) 28 27.5 +016 23-35 * In outer chorionic cells, mid-lateral surface, middle of egg. rior to anterior part of cell, but this pro- gression not as great as in Ps. columbiae (Fig. 5) and not easily discerned in individ- ual cells (Figs. 7e, 8b). Base of attachment of large tubercle fairly small, upper surface (Fig. 8e) fissured at base, at anterior end, and around edge, smoother centrally (but see variations, below), length 8-18 um, width 5-7 um. Cell floor with many tiny, round tubercles (Fig. 8e), diameter 0.1—-0.5 um. Small peripheral tubercles irregular in shape, longest dimension tending to parallel adjacent chorionic reticulum (Fig. 8b), each tubercle having a flared base and distinct upper portion, surfaces smooth (Fig. 8e, f). Outer chorionic reticulum a fine meshwork, only slightly raised, width 0.7-1.5 wm, with central line of closely spaced or intermit- tently fused bead-like protuberances, di- ameter 0.3-0.4 um (Fig. 8f). Variations in outer chorionic cell struc- ture as follows. Cells close to anterior end smaller, length 22—28 um, width 12-16 um, large tubercles usually flat or very little erect (Figs. 7a, 8a), more pointed, length 8-17 um, width 4.5-6.5 wm, upper surfaces en- tirely rough (Fig. 8a). Small tubercles more irregular. Approaching posterior end outer chorionic cells become somewhat smaller, length 21-35 um, width 17-20 um. Large tubercles longer (9-21 um), almost always decumbent (Fig. 7d), attached anteriorly to cell floor, often by ragged, finger-like exten- sions (Figs. 7d, 8c), surfaces smooth except for anterior edges (Fig. 8c). Immediately at 503 Fig. 6. Ps. cingulata. Entire egg, anterior end up- permost. Scale = 100 um. 504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Ps. cingulata. (a) anterior end, dorsal surface; (b) anterior end and micropylar apparatus; (c) detail of micropylar apparatus; (d) posterior end, lateral view; (e) typical outer chorionic cells, lateral surface, middle of egg. Scale = 20 um (a, b, d, e), = 10 um (c). VOLUME 92, NUMBER 3 Fig. 8. Ps. cingulata, details of lateral outer chorionic cells on different parts of egg (figure letters correspond to positions labelled in Fig. 6). (a) very close to anterior end; (b) at about anterior '/; (c) at about posterior '; (d) very close to posterior end; (e) detail of whole cell, middle of egg: (f) detail of outer chorionic reticulum, middle of egg. Scale = 10 um (a, b, c, d), = 5 um (e, f). 506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ; 3 % Fig. 9. Ps. ferox. Entire egg, anterior end upper- most. Scale = 100 um. posterior end large tubercles become scale- like, appressed to cell floors, anterior mar- gins ragged, surfaces smooth (Fig. 8d). Small tubercles few in number. Anterior end, micropyle: outer chorionic cells as described (Figs. 7a, 8a). Anterior ring absent. Micropylar collar sometimes fairly distinct (Fig. 6), but usually not so (Fig. 7a), complete, height 8-10 um, usually tapered, outer diameter about 27 um, outer margin quite regular, but surface rough (Fig. 7b, c), wall width fairly uniform, 5.2-—7 um, inner collar diameter about 13 wm, inner edge formed of a ring of very shallow ex- cavations (Fig. 7b, c). Micropylar disc not clearly visible, micropyle diameter about 2.2 um. Posterior end: outer chorionic cells as de- scribed, pole rounded, large tubercles in cells capping end button-like (Fig. 7d). Psorophora (Janthinosoma) ferox (Figs. 9-11) Size: dimensions as in Table 1. Color: black. Overall appearance: shape somewhat to quite fusiform, dorsal curvature greater than ventral, width greatest just anterior to mid- dle, anterior end especially conical, surface spiny, micropylar collar rather inconspic- uous (Fig. 9). Outer chorionic cells distinct, longitudinally elongated, with many pe- ripheral small tubercles and one long, an- teriorly inclined one (Fig. 9). Chorion, dorsal, lateral and ventral sur- faces: all surfaces very similar (Fig. 9). Cho- rionic cells almost always hexagonal, oc- casionally pentagonal (Fig. 10e), quite uniform in shape (Fig. 8), length 27-43 um, width 17-24 um. Cell fields about 2 um less in each dimension. Each cell with single very long, tongue-like tubercle originating very close to posterior margin of cell field and extending to about anterior 3 (Fig. 10e), tubercle much more sharply inclined from about anterior 3 (Fig. 10e). Small periph- eral tubercles numerous (Figs. 10e, 11b, e) VOLUME 92, NUMBER 3 507 Fig. 10. Ps. ferox. (a) anterior end, lateral surface; (b) anterior end and micropylar apparatus; (c) detail of micropylar apparatus; (d) posterior end, lateral view; (e) typical outer chorionic cells (Florida), lateral surface, middle of egg; (f) typical outer chorionic cells (Trinidad), lateral surface, middle of egg. Scale = 20 um. 508 and numbers variable (Table 2), longest di- mension usually perpendicular to adjacent chorionic reticulum, size distribution skewed (Fig. 4), and size increasing highly significantly from posterior to anterior part of cell (Fig. 5), as easily visible (Fig. 10e). In detail, large tubercles made up of distinct, flared base, with rough sides (Fig. 1 1e), sup- porting tongue-shaped upper portion. Length of tubercle 13-25 wm, width 5.5-7 um, surface covered with irregular bumps, slightly more defined anteriorly, less so pos- teriorly (Fig. 1 le). Cell floor covered with many tiny tubercles (Fig. | le, f), diameter 0.1-0.4 um, tending to be more numerous anteriorly. Small tubercles irregular, but usually tending to be rectangular (Fig. 1 1b, f), with widely flared base and small, cap- like upper portion, surfaces smooth (Fig. 11f). Chorionic reticulum an intricate, fine meshwork, slightly raised, width 1.1—2.2 um, with a double, centrally positioned row of small protuberances, diameter 0.1-0.4 um (Fig. lle, f). Meshwork of reticulum often extending up outer sides of small tubercles (Fig. 11f). Structural differences in chorionic cells as follows. Close to anterior end cells consid- erably smaller, length 16-24 um, width 9- 14 wm. Cell fields become progressively smaller (Fig. 10a, b) until only partially open, usually along one side of large tubercle (Fig. 11b). Inner margins of small tubercles even- tually fuse with large tubercle, forming con- tinuous layer (Figs. 10a, b, 11a). Large tu- bercles not as steeply inclined, becoming more decumbent (Fig. 10a), small tubercles becoming lower anteriorly, shape more ir- regular, bases flatter, caps often with bumpy surfaces (Fig. 11a). Immediately behind an- terior end small tubercles almost disappear (Figs. 10b, 11a), large tubercles appearing as low humps on cell surface (Fig. 10b). Meshwork of chorionic reticulum not dis- tinct, the two central rows of small protu- berances closer, forming prominent ridge (Fig. 11a). Somewhat further from anterior end cells much more as in middle of egg, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON but large tubercles not as steeply inclined (Fig. 11b), meshwork of reticulum visible but not as distinct, and its two central rows not as widely separated. Towards posterior end, progressive changes are similar to those at anterior end (Figs. 9, 10d). Cells become smaller, small tubercles flatter, large tuber- cles completely decumbent (Figs. 10d, 1 1c). Cells at posterior end have partially or al- most completely occluded fields, very in- distinct or almost indistinguishable small tubercles, and reticulum with meshwork scarcely detectable and pronounced central ridge (Fig. 11d). Large tubercles button-like with surface sculpturing less distinct (Figs. 10d, 11d) and almost invisible on most pos- terior ones (Fig. 11d). Anterior end, micropyle: chorionic cells as described. Anterior ring absent, micro- pylar collar visible but not prominent (Fig. 10a, b), not complete (gaps present), height 7 um, outer diameter about 40 um, outer margin bumpy (Fig. 10b, c), wall width vari- able, 0-6 um, inner collar diameter about 27 wm, inner margin quite smooth, with very shallow excavations (Fig. 10b, c). Mi- cropylar disc very prominently domed (Fig. 10b, c), diameter about 14 um, micropyle diameter about 2 um. Posterior end: outer chorionic cells as de- scribed, rather conical, tapering, and end smoothly rounded (Fig. 10d). Examination of eggs of Ps. ferox from Trinidad revealed a number of distinct dif- ferences from the Florida population. Al- though Trinidad eggs were longer (Table 1), they were not significantly so (t-test). They were, however, significantly greater in width (P < 0.001), and the length/width ratio con- sequently was smaller (P < 0.001). The out- er chorionic cells were similar in outline appearance, but differences in the small pe- ripheral tubercles rendered the Trinidad eggs visually distinct (Fig. 10f cf. 10e). The mean number of tubercles per cell in Trinidad eggs was significantly (P < 0.01) smaller (Table 2), and their size, on average, clearly greater (Fig. 4). Normalization by logarithmic VOLUME 92, NUMBER 3 509 Fig. 11. Ps. ferox, details of lateral outer chorionic cells on different parts of egg (figure letters correspond to positions labelled in Fig. 9). (a) very close to anterior end; (b) at about interior '4; (c) at about posterior '; (d) very close to posterior end; (e) detail of whole cell, middle of egg; (f) detail of outer chorionic reticulum, middle of egg. Scale = 10 um (a, b, c, d), = 5 um (e, f). transformation of the size distributions in- dition to this difference in terms of their deed proved the tubercles of Trinidad eggs largest dimension, the shape of the small to be larger (P < 0.001), as easily discernible tubercles also was different. Thirty tubercles in the micrographs (Fig. 10f cf. 10e). In ad- (10 randomly picked from each of 3 eggs 510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for each population) were measured to re- cord their radial (relative to the cell) and circumferential diameters and the ratios (r/ c) calculated. The mean (+SE) Trinidad ra- tio (1.02 + 0.03) was significantly (P < 0.001) smaller than the Florida sample (1.53 + 0.06), implying a much closer approxi- mation to square in the Trinidad eggs (Fig. 10f cf. 10e). DISCUSSION In their survey of the morphology of the inner chorion of eggs of the Ps. confinnis complex, Bosworth et al. (1983) examined 21 eggs of Ps. columbiae from the same lo- cation (Indian River County) as the mate- rial studied here. Their inner chorionic pat- tern type B was the most frequent (67%), with types A (5%), D (14%) and G (14%) also present. Type B is generally consistent with the intact eggs we examined. Notches in the cell outline (corresponding to the midline of the outer chorionic reticulum), some with short ridges extending into the cell field, are visible in Fig. 2e. Some cells seen at higher magnification also accord with type B (Fig. 3a, c, f) in that they appear to have short ridges in the cell fields, not con- nected with the reticulum. Type D as de- scribed by Bosworth et al. (1983) is similar to Type B, except that the cells contain a “cellular disc,” which obviously represents an impression of the base of the cell’s large tubercle on the inner chorion. Why only some cells stripped of the outer chorion show this impression is not clear. It has occurred to us, however, that the presence or absence of an impression could be determined by inconsistencies in preparative technique. These eggs were rolled on sticky tape to re- move the outer chorion (Bosworth et al. 1983) and presumably were subjected to at least some pressure. Differences in this pres- sure could have caused a gradation in the amount of detail impressed onto the inner chorion. Outer chorionic structures are eas- ily removed from these eggs by abrasion. Thus, it is also possible that some or many of the cells lacked the outer chorionic tu- bercles when the egg was applied to the sticky tape and could not have left an impression. Admittedly, cell outlines are not likely to be affected by removal of the outer chorion, but the possibility of preparative artifacts suggests the need for caution when inter- preting inner chorionic patterns. Given that egg surfaces are to be exam- ined by SEM, there is nothing to be gained by removing the outer chorion. Aside from possible artifacts, much structure, along with its potential variation, is lost. This is illus- trated by Ps. ferox eggs from Trinidad and Florida, where the number, greatest diam- eter and form of the small tubercles were all demonstrably different. The small tu- bercles were, of course, removed from eggs studied by Bosworth et al. (1983), but there is suggestive evidence that they may differ in number between populations of the Ps. confinnis complex. A few of the eggs from Arkansas and Texas possessed an unusual inner chorionic pattern (type J), in which the periphery of each cell was lined with small ovoid pits, almost certainly corre- sponding to the small tubercles. Bosworth et al. (1983) state that 15-20 pits were pres- ent in each cell, apparently more than in Ps. columbiae eggs from Florida, where the numbers ranged from 7-16 (mean 11.9) in 49 cells counted. Tubercle numbers are re- lated to cell size, but the anterolateral cells studied by Bosworth and co-workers would tend to be smaller (and therefore have fewer tubercles) than those selected here, which were laterally positioned in the middle of the egg. A re-examination of eggs of the Ps. confinnis complex, making use of the intact structural detail of the outer chorion, should be undertaken. ACKNOWLEDGMENTS We thank Dr. D. Sauerman for providing eggs of Ps. columbiae and Mr. L. C. Persad for laboratory and field assistance in Trin- idad. D. Duzak assisted with the electron microscopy and Bonnie Pattok printed the VOLUME 92, NUMBER 3 electron micrographs. This paper is Uni- versity of Florida, Institute of Food and Ag- ricultural Sciences Experiment Stations Journal Series No. R-00173. LITERATURE CITED Belkin, J. N., S. J. Heinemann, and W. A. Page. 1970. Mosquito studies (Diptera: Culicidae). X XI. The Culicidae of Jamaica. Contrib. Am. Entomol. Inst. (Ann Arbor) 6: 1-458. Bosworth, A. B., S. M. Meola, and J. K. Olsen. 1983. The chorionic morphology of eggs of the Psoroph- ora confinnis complex in the United States. I. Taxonomic considerations. Mosq. Syst. 15: 285- 309. Darsie, R. F., Jr.and R.A. Ward. 1981. Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosq. Syst. Suppl. 1: 1-313. Harbach, R. E.andK.L. Knight. 1980. Taxonomists’ 511 glossary of mosquito anatomy. Plexus Publishing Inc., Marlton, New Jersey. 415 pp. Horsfall, W. R., R. C. Miles, and J. T. Sokatch. 1952. Eggs of floodwater mosquitoes. I. Species of Pso- rophora (Diptera: Culicidae). Ann. Entomol. Soc. Am. 45: 618-624. Horsfall, W. R., F. R. Voorhees, and E. W. Cupp. 1970. Eggs of floodwater mosquitoes. XIII. Cho- rionic sculpturing. Ann. Entomol. Soc. Am. 63: 1709-1716. Heinemann, S. J., T. H. G. Aitken, and J. N. Belkin. 1980. Collection records of the project ‘*Mos- quitoes of Middle America.” 14. Trinidad and To- bago (TR, TRM, TOB). Mosq. Syst. 12: 179-284. Knight, K. L. and A. Stone. 1977. A catalog of the mosquitoes of the world (Diptera: Culicidae). En- tomological Society of America, The Thomas Say Foundation, Vol. VI, 611 pp. Linley, J. R. 1989. Comparative fine structure of the eggs of Aedes albopictus, Ae. aegypti and Ae. ba- hamensis (Diptera: Culicidae). J. Med. Entomol. 26: 510-521. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 512-520 RESOURCE UTILIZATION BY LARVAE OF PARACANTHA GENTILIS (DIPTERA: TEPHRITIDAE) IN CAPITULA OF CIRSIUM CALIFORNICUM AND C. PROTEANUM (ASTERACEAE) IN SOUTHERN CALIFORNIA DAvID HEADRICK AND RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, California 92521. Abstract.—Resource utilization and resource sharing by larvae of Paracantha gentilis Hering were analyzed in capitula of two native thistles (Tribe Cynareae): Cirsium cali- fornicum Gray and Cirsium proteanum J. T. Howell. Guilds described and analyzed for thistle-insect systems in Europe, though more complex than in southern California, ap- parently lack phytophages with the trophic strategy of P. gentilis. This tephritid displays rarely solitary, mainly aggregated attack on immature, closed capitula, but unlike European Tephritidae, does not form galls or otherwise cause host-tissue proliferation. Instead, feeding behavior of third instars of P. gentilis showed a density-dependent change in feeding niche from ovule-feeding alone to ovule, upper receptacle and plant-sap feeding which extended the range of available resources and minimized intraspecific competition. The feeding-niche of P. gentilis in thistle capitula is novel by European criteria, and exemplifies the failure of phytophage communities to converge in structure despite similar resources on different continents. Key Words: sium, evolution, thistles Resource utilization by thistle-head in- sects was described by Zwolfer (1985) as “the percent of flower heads in a sample containing phytophagous insects or show- ing signs of insect damage.” This parameter is more precisely measured as the number of damaged achenes per capitulum (= head). ZwoOlfer (1985) used resource utilization and guild structure to study the feeding ecology of several thistle-insect systems in Europe. Ten species of Cirsium (Asteraceae, Tribe Cynareae) are native to southern California (Munz 1974). Few native stenophagous in- sect species are associated with native Cir- sium spp. in North America (Goeden and Ricker 1986a, b, 1987a, b). In contrast, many stenophagous species are associated with European Cirsium spp. (Zw6lfer 1965). Insecta, resource utilization, guild structure, Tephritidae, gall-formers, Cir- This disparity offers opportunity for com- parative studies of resource allocation and resource utilization by thistle-head insects. Paracantha gentilis feeds in capitula of eight species of native Cirsium thistles and the introduced C. vulgare (Savi) Tenore in southern California (Goeden and Ricker 1986 a, b, 1987a, b) as well as several dif- ferent species of native Cirsium in northern California (Pemberton et al. 1985) and else- where in North America (Steck 1984). In C. californicum Gray, Goeden and Ricker (1986b) recorded P. gentilis as the dominant phytophage reared from 19 (73%) of 26 samples (average 50 capitula/sample). Three other insect species found in less than 50% of the capitula from the same samples were: Rotruda mucidella (Ragonot) (Lepidoptera: VOLUME 92, NUMBER 3 Pyralidae), Platyptilia carduidactyla (Riley) (Lepidoptera: Pterophoridae), and Orellia occidentalis (Snow) (Tephritidae). Paracan- tha gentilis was reported from 62% of sam- ples of C. proteanum capitula by Goeden and Ricker (1986b). Among the three other insect associates mentioned above, R. mu- cidella was the dominant phytophage in C. proteanum capitula, occurring in 92% of the samples (Goeden and Ricker 1986b). Zwolfer (1988) outlined three trophic strategies for thistle capitula-infesting in- sects in Europe: (1) An early-aggregated at- tack in closed young capitula, usually com- bined with gall formation, that lead to gregarious feeding behavior and protection from parasitoids and predators. In Europe, these insect associates are highly host-spe- cific Tephritidae, Cynipidae, and Curcu- lionidae. (2) Feeding on the maturing achenes and receptacle without induction of galls. Oviposition takes place in older ca- pitula, only single eggs are deposited, and capitula already occupied usually are avoid- ed, e.g. Tephritidae and Curculionidae in Europe. (3) Polyphagous species that usu- ally occur singly and oviposit after a capit- ulum has opened. The larvae are highly mo- bile, aggressive, often cause accidental mortality of other individuals in a capitu- lum, and are typically the dominant phy- tophage, e.g. Anobiidae, Pyralidae, and Tortricidae in Europe. Harris (1989) distin- guished between ovule (unfertilized) and soft or hard achene (fertilized ovule)-feeding by insects in knapweed and thistle capitula, a modification of Zwo6lfer’s (1988) scheme which we have used in our discussion. Preliminary field studies by Goeden and Ricker (1986a, b, 1987a, b) suggested dif- ferences from European Tephritidae in the way P. gentilis feeds and interacts with other insects infesting capitula of native Cirsium thistles in southern California. Compari- sons were based on two thistle-insect sys- tems involving (1) a C. californicum Gray population in which P. gentilis was the only capitulum infesting species and (2) a C. pro- 513 teanum J. T. Howell population in which P. gentilis was part of a capitulum-infesting guild composed of as many as four species. MATERIALS AND METHODS Cirsium californicum capitula were sam- pled at Mill Creek, San Bernardino National Forest, San Bernardino Co., CA, and C. pro- teanum capitula were sampled on Sawmill Mt., Angeles National Forest, Los Angeles Co., CA. Current season’s capitula were col- lected during the Spring and Summer of 1987, 1988 and 1989. Overwintered heads from 1986 were collected in early Spring, 1987, and stored at 5°C for later dissection. Serial dissections of capitula at different developmental stages were carried out to determine larval feeding behavior. Dissect- ed eggs and larvae were placed in covered glass petri dishes lined with filter paper and soaked with physiological saline and held in darkened growth chambers at 27°C until identified and then were held for puparia- tion and adult emergence, or were preserved or discarded. Field sites consisted of scattered individ- uals and patchy aggregations of 10-50 this- tles growing along roads and on south-fac- ing hillsides. At the Mill Creek site, 25 plants were inspected on each sampling date, and a total of 10-25 capitula were collected on each date for later dissection. The number of capitula on all plants occurring in a 3 x 3 m-area at the Mill Creek site was counted on four occasions during April-May, 1988, to determine phenology and capitulum pro- duction. Oviposition by overwintered fe- males in the small immature capitula (com- monly and erroneously called “‘buds’’) also was monitored on a weekly basis for the entire season during 1987 and 1988. Individual larvae and puparia dissected from capitula were placed in 60-mI* plastic vials fitted at one end with 100-mesh brass screening for ventilation and held in the in- sectary of the Department of Entomology, University of California, Riverside. Insec- tary conditions were 26° + 1°C, 30 to 60% 514 PED | | CAPITULUM Aen WEEK:11/ 2/3 1415! 6) 7/819 1/10/11 (12113114115 1161 P_GENTILIS STAGES EGG 1ST INSTAR }———-4 2ND INSTAR ———— 3RD INSTAR.) = }-}————4 PUPARIUM ADULT = }—e Fig. 1. Phenology of Paracantha gentilis in capitula of C. californicum. 1. Immature capitulum. II. Blos- som. III. Post-blossom. ACH, achene; FT, floral tube; INV PHY, involucral phyllanes; PAP, pappus; PED, peduncle; REC, receptacle. RH, anda 12-12 (1/d) photoperiod. Voucher specimens of all insect species involved are stored in the research collection of RDG, and eventually will be offered to the collec- tion of the Department of Entomology, University of California, Riverside. RESULTS AND DISCUSSION Resource utilization—The phenology of Paracantha gentilis in capitula of C. cali- fornicum in southern California is shown in Fig. 1. Paracantha gentilis follows the first trophic strategy in Zwé6lfer’s scheme de- scribed above, but with a major difference — although it oviposits in closed young capit- ula and usually feeds gregariously (up to 12 larvae per capitulum), it induces no galls. In the present study, resource utilization was calculated as the percentage of attacked capitula in a sample (Table 1). The accuracy of this statistic will be discussed below with regard to feeding strategies of capitulum- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Resource utilization or percent of C. cal- ifornicum and C. proteanum capitula infested by three phytophagous species.* No. (%) = Capitula Attacked Capitula Sample Dissected P.G. R.M. 0.0. C. californicum? 1986 15 12 (80%) 0 0 1987 110 86 (78) 0 0 1988 30 24 (80) 0 0 1989 40 31 (77) 0 0 Totals 195 153 (78) 0 0) C. proteanums 19824 Di; 18 (66) 20 (74) 21 (78) 1983¢ 27 8 (30) 17 (62) 11 (41) 1986 30 16 (53) 12 (40) 6 (20) 1987 50 37 (74) 43 (86) 26 (52) Totals 134 79 (58) 92 (68) 64 (47) @P.B., P. gentilis, R.M., R. mucidella; O.0., O. oc- cidentalis. > Mill Creek. © Sawmill Mt. * Unpublished data Goeden and Ricker. infesting Tephritidae. Thus, resource utili- zation by P. gentilis in C. californicum was 78%, as 153 of 195 capitula examined were attacked. In C. proteanum with its complex insect guild, resource utilization by P. gen- tilis was 58% (79 of 134 capitula), by R. mucidella was 68% (92 of 134 capitula), and by O. occidentalis was 47% (64 of 134 ca- pitula). In southern California, P. gentilis is an early-aggregated attacker; whereas O. occi- dentalis is a solitary phytophage that ovi- posits after the capitula begin to open and whose larvae feed on the floral tubes and maturing achenes. Thus O. occidentalis em- ploys Zwolfer’s second trophic strategy. Ro- truda mucidella and P. carduidactyla follow Zwolfer’s third trophic strategy as highly mobile, indiscriminate feeders (Goeden and Ricker 1986b, 1987a, b). Resource sharing.— Resource sharing by P. gentilis larvae in capitula of C. califor- nicum involved avoidance of intraspecific competition for food early in their life cycle. Females oviposit eggs singly or in clusters VOLUME 92, NUMBER 3 Fig. 2. Temporal and spatial partitioning (diagram- matic) ofa thistle capitulum by three of the most com- mon insect species comprising a complete guild in southern California. P.G., P. gentilis; O.0., O. occi- dentalis, R.M., R. mucidella. of up to 13 centrally in the capitula (Head- rick and Goeden 1990 and unpublished data). After eclosion, first instars tunnel into separate, nearby floral tubes where they feed for the entire stadium. Second instars leave these initially attacked floral tubes and tun- nel towards the outer margin of the capitula through a series of floral tubes well above the level of the achenes. Thus, central place- ment of the eggs and differences in feeding modes between the first two instars mini- mized competition between these instars 1n capitula of C. californicum. Third instar feeding was confined to the central ovules, and as noted below, scored the upper re- ceptacle at higher larval densities despite the presence of an outer ring of uneaten ovules remaining in the capitulum. Interspecific competition in C. protea- num was avoided among guild members by temporal and spatial division of the capitula resources (Fig. 2). Paracantha gentilis avoided substantial interspecific competi- tion because it attacked early and fed and pupariated centrally at anthesis surrounded by a mixture of dried feces and fragments of floral tubes and pappus hairs. This type 5 imm Fig. 3. Face view of dissected C. proteanum capit- ulum showing peripheral feeding path of R. mucidella (dotted-line), arrows show tops of centrally located P. gentilis puparia. Fig. 4. Face view of receptacle of C. californicum pitted with P. gentilis cups formed by third instars. Fig. 5. Cross-section of receptacle of C. californi- cum with P. gentilis puparium cupped in feeding cav- ity, dotted line denotes receptacle surface. 516 Table 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cirsium californicum capitula dissected (n = 107) containing late 3rd instar larvae or puparia, and the percentage of infested capitula with receptacle scoring by larvae. nN 1 No. of capitula per class 32 27 No. (%) of capitula per class with receptacle scored 2 (6) 4(15) of spatial evasion was first recognized by Zwolfer (1979) for thistle-feeding tephritids in Europe, where he suggested it mitigated the effects of interspecific competition. In contrast to P. gentilis, O. occidentalis and R. mucidella typically feed on the pe- ripheral achenes (Figs. 2, 3). The value of this avoidance behavior was illustrated by dissection of a capitulum in which a P. gen- tilis larva had pupariated angled ca. 45° from a normal position perpendicular to the re- ceptacle. The adult emerged abnormally to- wards the side of the capitulum, crossing the path of a R. mucidella larva which ate through it, leaving the head and abdomen behind! Third instar feeding.— Dissection of in- fested C. californicum capitula also led to the discovery that if three or more P. gentilis third instar larvae were present in a capit- ulum, at least one had scored the upper re- ceptacle centrally. In all capitula dissected, this receptacle feeding formed uniform cup- like depressions, 1.6 + 0.09 (x + S.E.) mm wide and 1-2 mm deep (n = 35) (Figs. 4, 5). The uniform depth and width of these depressions suggested a specialized function as sources of sap upon which the older lar- vae fed at higher larval densities. The mouthparts of the third instar, including the newly discovered “median oral lobe,’ show modification for such liquid food uptake (Headrick and Goeden 1990). Fluid feeding by larvae of Tephritidae in galls, capitula, and other plant parts may be underappre- ciated in its importance and occurrence. Romstéck (1987) reported that larvae of Tephritis conura Loew induced a callus while Infestation Class: No. 3rd Instar Larvae or Pupana per Capitulum 3 4 5 6 7 23 8 4 3 2 15 (65) 5 (62) 3 (75) 3 (100) 2 (100) feeding in the receptacle of Cirsium heter- ophyllum (L.) Hill. This structure then acted as a “‘sink”’ to maintain the nutrient flow to the head, much like a gall (Zw6lfer 1985). That tephritid galls act as metabolic sinks in capitula of knapweeds (Subtribe Cen- taurinae), close relatives of Cirsium thistles (Subtribe Carduinae) was well demonstrat- ed experimentally by Harris (1980). Thus, T. conura larvae similarly were found to induce and use callus formation to extend their resource limits, because the immature capitulum per se was a finite resource that did not contain enough nutrients to support completion of larval development. With P. gentilis, there was no induction of callus tis- sue or a gall, probably because receptacle scoring occurs after the meristematic stage and tissue differentiation (Harris 1980, 1989). Larval feeding apparently involves the slow, continual erosion and re-wound- ing of the upper receptacle during feeding- cavity formation to insure continued sap flow. In this manner, immature closed ca- pitula of limited biomass can sustain several P. gentilis larvae. This feeding mode is distinct from the receptacle feeding exhibited by species of Cirsium-head-infesting tephritids in Eu- rope (Zw6lfer 1988, Harris 1989) and else- where in North America (Steck 1984). As described by Steck (1984), Chaetostomella undosa (Coquillett) larvae bored into and fed on the receptacle tissues during all three instars. When larval densities were high, and after the receptacle was consumed, they mined downwards into the peduncle and stem. The scheme of Harris (1989) limits VOLUME 92, NUMBER 3 Table 2. Extended. Infestation Class: No. 3rd Instar Larvae or Pupana per Capitulum 8 9 10 11 12 1 3 1 0) yp) 1 (100) 3 (100) 1 (100) = 2 (100) receptacle feeding by non-gall formers to immature “buds,” but C. wndosa continues to mine the receptacle through floret and achene growth stages (Steck 1984). Par- acantha gentilis co-existed with C. undosa and was the dominant phytophage in Cir- sium capitula sampled by Steck (1984). No other species of non-gall-forming, ovule or soft achene-feeding Tephritidae has yet been reported to feed on the ovules and then score the receptacle in the manner of P. gentilis, cf. Tauber and Toschi (1965), Stegmaier (1967), Cavender and Goeden (1982, 1984), Lamp and McCarty (1982), Goeden (1987), and Goeden et al. (1987). These data also clearly demonstrate that tephritids associ- ated with Nearctic Cirsium spp. show novel feeding habits distinct from Palearctic te- phritids (Zw6lfer 1988, Harris 1989). This change in third instar feeding pat- terns was analyzed in capitula containing only P. gentilis larvae. The average number of third instars found in 107 infested capit- ula was 3.3 + 0.3 (range, 1-12). The recep- tacle diameter (used as an index of available resources) was highly correlated with the number of achenes per capitulum (corr. coeff. = 0.851, P=0.001, n = 25). However, there was no significant correlation between the diameter of the receptacle and the num- ber of larvae in a given capitulum, so the size of a capitulum did not limit infestation density. Dissections showed, instead, that the age of the third instar as well as larval density determined the feeding mode. When capit- ula contained fewer than three third instar larvae, they usually fed centrally on the 238 8 x 3 § 8 8 TOTAL WO OVULES ATTACKED e 6 i) 1 2 304 5 6 7 8 9 10 " 12 NO. 3RD INSTAR LARVAE PER CAPITULUM Fig. 6. Relationship of total ovules attacked per capitulum and P. gentilis third instar density. ovules and pupariated without scoring the receptacle (Table 2). A change in third instar feeding behavior took place in capitula with three or more individuals; instead of con- tinuing to feed on the ovules, at least one, possibly the smallest and/or the youngest, third instar(s) scored the receptacle. This occurred in 65% of all the capitula dissected containing three third instars. All capitula containing six or more (up to | 2) third instar larvae had scored receptacles (Table 2). Zwolfer (1985) noted that the number of achenes (ovules) attacked was a more pre- cise measurement of resource utilization than the percentage of capitula attacked. However, in capitula of C. californicum containing from three to 12 puparia, com- plete consumption of the ovules never was detected. The number of ovules attacked per larva decreased as densities rose (Fig. 6). This decrease was not due to a depletion of resources (ovules), intraspecific compe- tition and larval starvation, as suggested for P. culta (Wiedemann) by Lamp and Mc- Carty (1982). Instead, this reflected a new resource, i.e. the receptacle, being used along with the central ovules by the younger third instars. Therefore, the precision of the sta- tistic for resource utilization is dependent on the feeding strategy of the species of Te- phritidae involved. Ecological significance.— By not consum- ing all the achenes in a capitulum, P. gentilis followed the strategy of “‘evasion,” as de- 518 X PUPARIAL WIDTH(mm) NO. PUPARIA PER CAPITULUM Fig. 7. Relationship of mean pupal width of P. gen- tilis per capitulum of C. californicum to pupal density (corr. coeff. = 0.059, n.s.). fined by Zw6lfer (1979) for thistle-head-in- festing tephritids. However, the strategy of a “non-interactive grazing system” in which the phytophages merely consume a surplus of seeds produced (Zwo6lfer 1979) does not apply to P. gentilis in C. proteanum capit- ula. At the Sawmill Mt. location, where the guild was complete, i.e. at least three insect species infested the capitula, total con- sumption of achenes occurred in 60% of 50 infested capitula dissected. Paracantha gen- tilis had the advantage in this system by its optional use of an evasion strategy, 1.e. being able to augment ovule-feeding with another, replenishable resource—plant sap from the receptacle. In this manner, P. gentilis avoid- ed most competition from other guild mem- bers as well as among its siblings at high densities. Several parameters were measured on adults reared from puparia dissected from capitula containing one to 12 individuals to determine if there were any effects of upper receptacle-moderated sap-feeding on pu- parial and adult sizes. No significant differ- ence in puparial widths was found among individuals in capitula containing low and high fly densities (Fig. 7). Percentage adult emergence showed no differences among ca- pitula containing different numbers of pu- paria, i.e. 54 of 58 (91.8%) adults emerged from unparasitized puparia. Head widths, hind tibial lengths and oviscape lengths of adult females also showed no significant correlation among flies that emerged from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2s oO " 2 E E : = a w 5 * a = 5 > 2 * 1 *Oe+ OVISCAPE e@— HEAD CAPSULE — HIND TIBIA Oo ° 1 2 3 4 5 6 7 8 9 7o oN” 12 NO. PUPARIA PER CAPITULUM Fig. 8. Relationship of three P. gentilis adult female measurements (max. head width, hind tibia length, oviscape length) to pupal P. gentilis density per capit- ulum (corr. coeff. = 0.181, 0.154, 0.033, respectively, n.s., n = 27). capitula containing low or high numbers of puparia (Fig. 8). CONCLUSION The trophic strategy involving early-ag- gregated attack in capitula by Tephritidae, as now studied with both Palearctic and Nearctic species, shows three types of ad- aptations to extend and partition a finite resource, the capitulum. These adaptations involve tephritids that induce galls in the immature capitula (Zw6lfer 1985); callus- forming tephritids (Romstéck 1987); and as reported in the present study, a receptacle- scoring tephritid that induces no plant tissue growth, and thus directly feeds on assimi- lates channelled to the immature capitulum. Is Paracantha gentilis evolving towards gall formation? According to Zwélfer (1983), the most evolutionarily advanced insect- thistle capitulum relationship is gall for- mation. This implies an intimate, long evolved relationship between insect and host plant. However, Zwé6lfer (1988) noted that tephritid species maintained their trophic preadaptations during host transfer, i.e. gall- formers induced galls on any new host. By this interpretation, P. gentilis was preadapt- ed to feed in Cirsium capitula without in- ducing tissue proliferation. But, is the change VOLUME 92, NUMBER 3 to receptacle feeding by third instars also a pre-adapted feeding response to infestation densities, or an evolved adaptation to re- duce inter- and intraspecific competition within thistle capitula? If it is the latter, P. gentilis may further evolve into a gall for- mer. North American Cirsium thistles lack gall-forming insect associations (Goeden and Ricker 1987b). Supplementing findings by ZwoOlfer (1985, 1988) and Romstoéck (1987), we suggest that receptacle scoring by P. gen- tilis represents just one end of a spectrum of feeding strategies evolved by thistle-head infesting tephritids, i.e. a new category of an early aggregated attacker that makes use ofan immature capitulum in a unique man- ner to support its larval development with- out recourse to gall or callus tissue forma- tion. In this manner the infested, closed, immature capitulum itself acts like a gall, offering the larvae a food source and con- stant micro-environment protected from desiccation, predation, and most parasit- oids (Headrick and Goeden 1989b). The “feeding niche” of P. gentilis in this- tle capitula is thus novel by European cri- teria (Zw6lfer 1988, Harris 1989). There- fore, it and associated herbivores exemplify the failure of phytophage communities to converge in structure despite similar re- sources on different continents, as demon- strated with bracken (Pteridium aquilinium (L.) Khn.) and its herbivores in England, New Mexico, and South Africa by Lawton (1976, 1982) and Compton et al. (1989). ACKNOWLEDGMENTS We thank D. W. Ricker for technical as- sistance with this research, T. S. Bellows for his helpful discussions and manuscript re- view, and G. Gordh, J. D. Hare, P. Harris, J. Lawton, A. L. Norrbom, J. Pinto, and H. Zwolfer for their advice and comments on earlier drafts of our manuscript. LITERATURE CITED Cavender, G. L.and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild 519 sunflower in southern California. Ann. Entomol. Soc. Am. 75: 400-406. 1984. The life history of Paracantha cultaris (Coquillett) on wild sunflower, Helianthus annuus L. ssp. /enticularis (Douglas) Cockerell, in south- ern California (Diptera: Tephritidae). Pan-Pac. Entomol. 60: 213-218. Compton, S. G., J. H. Lawton, and V. K. Rashbrook. 1989. Regional diversity, local community struc- ture and vacant niches: The herbivorous arthro- pods of bracken in South Africa. Ecol. Entomol. 14: 365-373. Goeden, R. D. 1987. Life history of Trupanea con- juncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). Pan- Pac. Entomol. 63: 284-291. Goeden, R. D. and D. W. Ricker. 1986a. Phytoph- agous insect faunas of two introduced Cirsium thistles, C. ochrocentrum and C. vulgare, in south- ern California. Ann. Entomol. Soc. Am, 79: 945- 952. 1986b. Phytophagous insect faunas of the two most common native Cirsium thistles, C. cal- ifornicum and C. proteanum, in southern Califor- nia. Ann. Entomol. Soc. Am. 79: 953-962. 1987a. Phytophagous insect faunas of the native thistles Cirsium brevistylum, C. congdonii, C. occidentale, and C. tioganum, in southern Cal- ifornia. Ann. Entomol. Soc. Am. 80: 152-160. . 1987b. Phytophagous insect faunas of native Cirsium thistles, C. mohavense, C. neomexica- num, and C. nidulum, in the Mohave Desert of southern California. Ann. Entomol. Soc. Am. 80: 161-175. Goeden, R. D., T. D. Cadatal, and G. L. Cavender. 1987. Life history of Neotephritis finalis (Loew) on native Asteraceae in southern California (Dip- tera: Tephritidae). Proc. Entomol. Soc. Wash. 89: 552-558. Harris, P. 1980. Effects of Urophora affinis Frfld. and U. quadrifasciata (Meig.) (Diptera: Tephritidae) on Centaurea diffusa Lam. and C. maculosa Lam. (Compositae). Z. Angew. Entomol. 90: 190-201. Feeding strategy, coexistence and impact of insects in spotted knapweed capitula. In press. Jn Delfosse, E. S., ed., Proceedings of the VII Inter- national Symposium on Biological Control of Weeds, Rome, Italy, 1988. Headrick, D. and R. D. Goeden. 1989. Life history of Pteromalus coloradensis (Ashmead) (Hyme- noptera: Pteromalidae), a parasite of Paracantha gentilis Hering (Diptera: Tephritidae) in Cirsium thistle capitula. Proc. Entomol. Soc. Wash. 91: 594-603. 1990. Description of the immature stages of Paracantha gentilis. Ann. Entomol. Soc. Am. 83: 220-229. Lamp, W. D. and M. K. McCarty. 1982. Biology of 520 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON predispersal seed predators of the Platte Thistle, Cirsium canescens. J. Kans. Entomol. Soc. 55: 305- 316. Lawton, J. H. 1976. The structure of the arthropod community on bracken. Bot. J. Linn. Soc. 73: 187- 216. 1982. Vacant niches and unsaturated com- munities: A comparison of bracken herbivores at sites on two continents. J. Anim. Ecol. 51: 573- 575% Munz, P. A. 1974. A Flora of Southern California. University of California, Berkeley. Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 1985. New host records for tephritid flies (Dip- tera) from Cirsium and Saussurea thistles (Aster- aceae) in California. Proc. Entomol. Soc. Wash. 87: 790-794. Romstéck, M. 1987. Tephritis conura Loew (Diptera: Tephritidae) und Cirsium heterophyllum (L.) Hill (Cardueae). Struktur- und Funktionsanalyse eines dkologischen Kliensystems. Dissertation Report in Natural Sciences, Bayreuth 1987. 147 pp. Steck, G. J. 1984. Chaetostomella undosa (Diptera: Tephritidae): Biology, ecology, and larval descrip- tion. Ann. Entomol. Soc. Am. 77: 660-678. Stegmaier, C. E., Jr. 1967. Notes on a seed-feeding Tephritidae, Paracantha forficula, (Diptera) in Florida. Fla. Entomol. 50: 157-160. Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior of Tephritis stigmatica (Co- quillett) (Diptera: Tephritidae). Pan-Pac. Ento- mol. 41: 73-79. Zwolfer, H. 1965. Preliminary list of phytophagous insects attacking wild Cynareae (Compositae) in Europe. Commonw. Inst. Bio. Cont. Tech. Bull. 6: 81-154. 1979. Strategies and counterstrategies in in- sect population systems competing for space and food in flower heads and gall. Fortschr. Zool. 25: 331-353. . 1983. Life systems and strategies of resource exploitation in tephritids, pp. 16-30. Jn Cavalloro, R., ed., Fruit Flies of Economic Importance. Proc. CEC/IOBC Int. Sym., Athens, Greece. Nov. 1982, A. A. Balkema, Rotterdam. . 1985. Insects in Thistle Heads: Resource Uti- lization and Guild Structure, pp. 407-416. Jn De- fosse, E. S., ed., Proc. 6th Int. Sym. Bio. Cont. Weeds, Vancouver, Canada. 1988. Evolutionary and ecological relation- ships of the insect fauna of thistles. Ann. Rev. Entomol. 33: 103-122. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 521-529 TARSAL AND OVIPOSITOR SENSILLA OF HELIOTHIS VIRESCENS AND H. SUBFLEXA (LEPIDOPTERA: NOCTUIDAE) GERALD T. BAKER AND SONNY B. RAMASWAMY Department of Entomology, Mississippi State University, P.O. Drawer EM, Mississippi State, Mississippi 39762. Abstract. —Three types of sensilla are present on the tarsi of Heliothis virescens and H. subflexa; a long, fluted sensillum chaeticum and short and long sensilla trichodea. The number of these sensilla varies from tarsus to tarsus within a species and between species. Females of H. virescens have significantly more of each type than females of H. subflexa and the males of H. virescens. Tarsus II bears the most of each type of sensillum; tarsus III has the fewest. The ovipositor of each species does not differ in the types and number of sensilla. Long and short sensilla chaetica can be found on most areas of the ovipositor whereas the 5 or 6 sensilla trichodea are situated on the apex of each valve. The surface of each valve is covered by short, pointed microtrichia. Key Words: Olfactory receptors, contact chemorecep- tors, mechanoreceptors and visual receptors that are situated on various insect body re- gions such as the antennae, legs and ovi- positor are involved in host finding (Dethier 1982, Miller and Strickler 1984, Ramaswa- my 1988). Little is known about the role of tarsi and ovipositor in selection of an ovi- position site by adult Lepidoptera. Most of the early morphological and behavioral re- search on adult Lepidoptera dealing with host plant selection is on butterflies (Min- nich 1921, 1922a, b, Fox 1966, Ma and Schoonhoven 1973, Calvert 1974, Calvert and Hanson 1983, Renou 1983). Many species in the family Noctuidae are serious pests on a wide variety of important agricultural crops throughout the world but our knowledge 1s scarce about what role sen- sory receptors on the tarsi and ovipositor play in host-plant finding. Tarsal and ovi- positor sensory receptors are known to have an important role in host plant selection in moths such as Chilo partellus (Swinhoe) and Noctuidae, Heliothis, tarsus, ovipositor, sensilla Spodoptera littoralis (Boisd.), by responding to various chemical and mechanical stimuli (Chadha and Roome 1980, Waladde 1983, Salama et al. 1984, Waladde et al. 1985). There are several important pest species in the genus Heliothis, at present only one pa- per (Ramaswamy et al. 1987) deals with the possible role of sensory receptors on the tar- si and ovipositor in host plant selection for oviposition of H. virescens (F.). The purpose of the present study is to provide infor- mation on the morphology, number and distribution of sensory receptors on the tarsi and ovipositor of H. virescens, which has a wide host plant range, and to compare this species to H. subflexa, which has a very re- stricted host plant range. MATERIALS AND METHODS Specimens for scanning electron micros- copy were fixed in 4% glutaraldehyde in Na- cacodylate buffer (pH 7.1) for 8 h at 4°C. They were washed in the same buffer and post-fixed in 2% osmium tetroxide for 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tarsi of H. virescens female. 1, Tarsus I. 2, Tarsus II. 3, Tarsus III. 3. Figs. l- VOLUME 92, NUMBER 3 Table 1. Comparison of the number of sensilla on the tarsi of H. virescens females. Tarsus A B (e I 21.88 + 0.83b 42.75 + 1.04b 11.46 + 0.52b Il 23.63 + 0.74a 45.25 + 0.89a 12.81 +0.7la Ill 14.88 + 0.64¢ 32°50 25:0:93'¢ 9.43 + 0.67 ¢ A = sensillum chaeticum; B = short sensillum trichodeum; C = long sensillum trichodeum. Means within a column not followed by the same letter are significantly different (P < 0.05) as determined by ANOVA followed by Student-Newman-Keuls Test (n = 8). h. After dehydration in a graded series of ethanol, the specimens were placed in pen- tane overnight and then air dried. The tarsi and ovipositors were sputter-coated with gold-palladium and examined with a JEOL JSM-35 CF scanning electron microscope at 20 kV. The legs of eight female H. virescens and H. subflexa and eight male H. virescens were cleared in 7% KOH and mounted in eu- paral. The data on the number of sensory receptors on tarsi I-III are given as a mean plus the standard deviation and were sub- jected to ANOVA followed by Student- Newman-Keuls Test (P < 0.05). A ¢-test (P < 0.05) was used for the comparison of the number of sensory receptors on each tarsus of H. virescens and H. subflexa females and H. virescens female and male. RESULTS AND DISCUSSION Tarsi.—The tarsi consist of 5 tarsomeres and a pretarsus. Scales cover the dorsal sur- face of each tarsus whereas on the ventral surface of each tarsus there is an area that is devoid of scales (Figs. 1-3). This area 1s the contact region between the tarsus and substrate. The total area that comes in con- tact with the substrate differs for tarsi I-III (Figs. 1-3). It is from this region that the number of various types of sensory recep- tors was counted. Tarsus IT has the largest contact area and the greatest number of sen- sory receptors, followed by tarsi I and III (Figs. 1-3; Tables 1, 2). The long trichoid sensilla are also located on the periphery of the contact region and at the apex of tarsomere V (Fig. 1). Two long trichoid sensilla are apically located on tarsomere V on either side of the tarsomere midline (Fig. 1). This sensory receptor is 65— 85 wm long and curves back in the direction of the body (Figs. 1, 8). A distinct cuticular pattern covers the surface of the sensillum from the base to the apex (Fig. 9). On each tarsus there are 3 rows of sensilla chaetica, 1 row on each side of the contact region and the third row set off-centre on tarsus I and on the midline of tarsi II and III (Figs. 1-3). This pattern is the same for the female and male of H. virescens and H. subflexa female (Figs. 1-7). The sensillum Table 2. Comparison of the number of sensilla on the tarsi of H. subflexa females. Tarsus A B Cc I 18.13 + 0.99 a 37.88 + 0.83 a 9.31 + 0.64 a Il 12.13 + 0.98 b 28.00 + 0.76 b 6.78 + 0.81 b Ill E752 O77 10b 27.88 + 0.64b 6.12 + 0.77b A = sensillum chaeticum; B = short sensillum trichodeum; C = long sensillum trichodeum. Means within a column not followed by the same letter are significantly different (P < 0.05) as determined by ANOVA followed by Student-Newman-Keuls Test (n = 8). 524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-7. male. chaeticum has a socket at the base, tapers and slants downward in the direction of the pre-tarsus (Fig. 1). The surface of this sen- sory receptor has large, longitudinal ridges possessing secondary ridges on its surface and pits are situated at the base of the large ridges (Figs. 10, 11). This type of sensillum ranges from 80-165 um long. Two rows of short trichoid sensilla that are situated on either side of the central row of sensilla chaetica are on each tarsus in the contact region (Fig. 1). There are 2 distinct 4-6, Tarsi of H. subflexa female. 4, Tarsus I. 5, Tarsus II. 6, Tarsus III. 7, Tarsus I of H. virescens types of short trichoid sensilla. The first type if 42-46 um long and 7-9 wm wide at the base and is slightly curved near the apex (Fig. 12). The receptor surface is covered with irregular longitudinal striations (Fig. 13). The second type of short trichoid sen- sillum is 39-43 um long and 5—6.5 wm wide at the base, with the apical third distinctly hooked (Fig. 12). The receptor surface has irregular, lateral striations (Fig. 14). The distribution pattern and morpholog- ical types of sensilla that are found on the VOLUME 92, NUMBER 3 Figs. 8-11. tarsi of female H. virescens and H. subflexa, and male H. virescens are similar in that each tarsus has 2 outer rows of sensilla chae- tica and long trichoid sensilla, a central row of sensilla chaetica and a row of short trich- oid sensilla on either side of the central row of sensilla chaetica (Figs. 1-7). The number of sensory receptors differs on each tarsus. In females of H. virescens and H. subflexa the number of sensilla chaetica and long and short trichoid sensilla show the same pat- tern in that tarsus II > tarsus I > tarsus III (Tables 1, 2). There is a significant difference between H. virescens and H. subflexa fe- males in the number of sensory receptors on each tarsus in that H. virescens has the greater number of each sensillar type (Table 3). The pattern in the number of sensory receptors on the male tarsi of H. virescens differs from the female in that tarsus I has 8, Long sensilla trichodea. 9, Surface pattern on the long sensillum trichodeum. 10, Sensillum chaeticum. 11, Surface pattern on the sensillum chaeticum. significantly more long and short trichoid sensilla and sensilla chaetica than tarsi II and III which have similar numbers of sen- sillar types (Table 4). Females of H. vires- cens have significantly more of all sensillar types than the males (Table 5). Similar sensilla chaetica and trichodea on the tarsi of H. virescens and H. subflexa are also located on the tarsi of other moth species such as Chilo partellus and Eldana sac- charina (Waladde 1983), and Helicoverpa zea (Callahan 1969), but there are no data on the distribution and number of each sen- sillar type on the tarsus of each leg. These types of sensilla are present on the tarsi of females and males of Pieris brassicae and the distribution pattern is similar to what is found on H. virescens and H. subflexa. The number of sensilla chaetica on the tarsi of Chlosyne lacina Geyer and Heliconius 526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | Figs. 12-14. Short sensilla trichodea. 12, Straight and hooked forms. 13, Surface of the straight form. 14, Surface of the hooked form. charitonius L. is greatly reduced, and this type of sensillum is often associated with a cluster of trichoid sensilla (Calvert 1974, Renou 1983). Electrophysiological experiments on sev- eral species of Lepidoptera showed that the trichoid sensilla respond to salt, sugar and plant substances (Morita etal. 1957, Takeda 1961, Ma and Schoonhoven 1973, Renou 1983, Waladde 1983, Waladde et al. 1985). Behavioral tests involving the tarsi indicate the importance of tarsal sensory receptors for host plant acceptance and oviposition (Ma and Schoonhoven 1973, Calvert and Hanson 1983, Salama etal. 1984, Faucheux 1985). The trichoid sensilla on the tarsi of H. virescens respond to salts, sugars and plant extracts (Ramaswamy and Hanson unpub- lished data), and behavioral experiments Table 3. Comparison of the number of sensilla on the tarsi of H. virescens males. Tarsus A I 18.38 + 0.92 b II 22.00 + 0.76a Ill 112163) £20:911''¢ B ¢c 38.50 + 0.93 b 971 10:59'b 42.88 + 0.99 a 11.34 + 0.68 a 26.75 + 1.49 ¢ 8.18 + 0.52¢ A = sensillum chaeticum; B = short sensillum trichodeum; C = long sensillum trichodeum. Means within a column not followed by the same letter are significantly different (P < 0.05) as determined by ANOVA followed by Student-Newman-Keuls Test (n = 8). VOLUME 92, NUMBER 3 52i1 Table 4. Comparison of the number of sensilla on the tarsi of H. virescens and H. subflexa females. Tarsus A Hv. 1 21.88 + 0.83a Ess 18.38 + 0.92 b H. v. I 23.63 + 0.74a Je FoKyel Ul 22.00 + 0.76 b H. y. Ul 14.88 + 0.64 a Je Eaxyen UE 12.63 + 0.92 b B Cc 42.75 + 1.04a 11.46: '0:52/a 38.50 + 0.93 b 9.71 + 0.59 b 45.25 + 0.89 a 12.81 + 0.71 a 42.88 + 0.99 b 11.34 + 0.58 b 32.50 + 0.93 a 9.43 + 0.67a 26.75 + 0.49 b 8.18 + 0.52 b A = sensillum chaeticum; B = short sensillum trichodeum; C = long sensillum trichodeum. Means not followed by the same letter are significantly different (P < 0.05) as determined by Student’s-t-test (n = 8). showed that the sensory receptors on the tarsi are involved in host plant acceptance and oviposition (Ramaswamy et al. 1987). Heliothis virescens oviposits on a wide va- riety of host plants, such as tobacco, cotton, peanut and tomato, whereas H. subflexa uses ground cherry. This difference is the num- ber of host plants for oviposition may be due in part to the difference in the number of sensory receptors found on the tarsi of both species. Ovipositor.—The ovipositor consists of two papillae anales that are on either side of the oviduct and anal openings (Fig. 15). The oviduct opening is surrounded by short and long sensilla chaetica which have ta- pered tips and smooth walls (Fig. 16). These sensilla are also situated on the rest of the Ovipositor surface (Fig. 17). The apex of the ovipositor bears blunt-tipped trichoid sen- silla which have shallow longitudinal grooves on the surface that-fade near the apex of the sensillum (Fig. 18). Each pa- pillae anales has five to six of these trichoid sensilla. The remainder of the ovipositor surface is covered with microtrichia. There are no differences in the morphol- ogy, number and distribution of the sensilla between H. virescens and H. subflexa. The trichoid sensilla on the ovipositor of Phthorimaea operculella (Zell.) (Gelechi- idae) are contact chemoreceptors that are involved in oviposition on a suitable sub- strate (Fenmore 1978, Valencia and Rice 1982). Additionally these sensilla are pres- ent on the ovipositor of noctuid species such as Chilo partellus and Spodoptera littoralis, and morphological and electrophysiological data indicate that they are contact chemo- receptors (Chadha and Roome 1980, Wa- ladde 1983, Waladde et al. 1985). But the exact role(s) these contact chemoreceptors Table 5. Comparison of the number of sensilla on the tarsi of H. virescens females and males. > Tarsus H. v. f1 21.88 + 0.83 a H.v.mI 18.13 + 0.99 b H. v. f I 23.63 + 0.74a H. v.m I 12.13 + 0.98 b H. y. f Ill 14.88 + 0.64a H. vy. m Ill eT Sic Ob B € 42.75 + 1.04a 11.46 + 0.52a 37.88 + 0.83 b 9.31 + 0.64 b 45.25 + 0.89 a 12.81 +0.7la 28.00 + 0.64 b 6.78 + 0.81 b 32.50 + 0.95 a 9.43 + 0.67a 27.88 + 0.64 b 6:12. 20.77 D A = sensillum chaeticum; B = short sensillum trichodeum; C = long sensillum trichodeum. Means not followed by the same letter are significantly different (P < 0.05) as determined by Student-t-test (n = 8). 528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A! Figs. 15-18. Ovipositor of H. virescens. 15, Two papillae anales surrounding the anal and oviduct openings. 16, Short and long sensilla chaetica around the oviduct opening. 17, Sensillum trichodeum amongst the long and short sensilla chaetica. 18, Sensillum trichodeum with a slightly grooved surface. VOLUME 92, NUMBER 3 play in oviposition behavior of the above mentioned species and H. virescens and H. subflexa needs to be determined. ACKNOWLEDGMENTS We would like to thank D. Harvey and B. Perrigin for typing the manuscript. The manuscript 1s assigned no. J-7278. LITERATURE CITED Callahan, P. S. 1969. The exoskeleton of the corn earworm moth, Heliothis zea, Lepidoptera: Noc- tuidae, with special reference to the sensilla as polytubular dielectric arrays. Univ. Georgia, Coll. Agric. Exp. Stn. Res. Bull. No. 54, 105 pp. Calvert, W. H. 1974. The external morphology of foretarsal receptors involved with host discrimi- nation by the nymphalid butterfly, Chlosyne la- cinia. Ann. Entomol. Soc. Am. 67: 853-856. Calvert, W. H. and F. E. Hanson. 1983. The role of sensory structures and preoviposition behavior in oviposition by the patch butterfly, Chlosyne la- cinia. Entomol. Exp. Appl. 33: 179-187. Chadha, G. and R. Roome. 1980. Oviposition and behavior and the sensilla of the ovipositor of Chilo partellus and Spodoptera littoralis (Lepidoptera: Noctuidae). J. Zool., Lond. 192: 169-178. Dethier, V. G. 1982. Mechanism of host plant rec- ognition. Entomol. Exp. Appl. 31: 49-56. Faucheux, M. 1985. Role des tarsomeres et des ap- pendices cephaliques dans le comportement de ponte Tineola bisselliella Humm. (Lepidoptera: Tineidae). C. R. Acad. Sc. Paris 300: 493-498. Fenmore, P. G. 1978. Oviposition of potato tuber moth Phthorimaea operculella Zell. (Lepidoptera: Gelechiidae); the physical nature of the oviposi- tion substrate. N.Z. J. Zool. 5: 591-599. Fox, R. M. 1966. Forelegs of butterflies. Introduc- tion: Chemoreception. J. Res. Lepid. 5: 1-12. Ma, W. C. and L. M. Schoonhoven. 1973. Tarsal contact chemosensory hairs of the large white but- terfly Pieris brassicae and their possible role in oviposition behavior. Entomol. Exp. Appl. 16: 343-357. Miller, J. R. and K. L. Strickler. 1984. Finding and accepting host plants, pp. 127-157. Jn Bell, W. J. and R. T. Carde, eds., Chemical Ecology of Insects. 529 Sinauer Associates, Sunderland, Massachusetts. 524 pp. Minnich, D. E. 1921. An experimental study of the tarsal chemoreceptors of two nymphalid butter- flies. J. Exp. Zool. 33: 173-203. 1922a. The chemical sensitivity of the tarsi of the red admiral butterfly, Pyrameis atalanta L. J. Exp. Zool. 35: 57-81. 1922b. A quantitative study of tarsal sensi- tivity to solutions of saccharose, in the red admiral butterfly, Pyrameis atalanta L. J. Exp. Zool. 36: 445-457. Morita, H., S. Doira, K. Takeda, and M. Kuwabara. 1957. Electrical responses of contact chemore- ceptors on the tarsus of the butterfly Vanessa in- dica. Mem. Fac. Kyushu Univ., Series E (Biol.) 2: 119-139. Ramaswamy, S. B. 1988. Host finding by moths: Sensory modalities and behaviors. J. Insect Phys- iol. 34: 235-249. Ramaswamy, S. B., W. K. Ma, and G. T. Baker. 1987. Sensory cues and receptors for oviposition by He- liothis virescens. Entomol. Exp. Appl. 43: 159- 168. Renou, M. 1983. Les recepteurs gustatifs du tarse anterieur de la femelle d’Heliconius charitonius (Lep.: Heliconiidae). Ann. Soc. Entomol. Fr. 19: 101-106. Salama, H., A. Rizk, and A. Sharaby. 1984. Chemical stimuli in flowers and leaves of cotton that affect behavior in the cotton moth, Spodoptera littoralis (Lepidoptera: Noctuidae). Entomol. Gen. 10: 27- 34. Takeda, K. 1961. The nature of impulses of single tarsal chemoreceptors in the butterfly Vanessa in- dica. J. Cell. Comp. Physiol. 58: 233-245. Valencia, L. and M. Rice. 1982. Contact chemore- ceptors on the ovipositor of the potato moth, Phthorimaea operculella (Zell.) (Lepidoptera: Ge- lechiidae). Int. J. Insect Morphol. Embryol. 11: 121-128. Waladde,S.M. 1983. Chemoreception of adult stem- borers: Tarsal and ovipositor sensilla on Chilo par- tellus and Eldana saccharina. Insect Sci. Appl. 4: 159-165. Waladde, S. M., H. Kahoro, E. Kokwaro, and M. Chimtawi. 1985. Responses of Chilo partellus to material obtained from susceptible and resistant maize cultivars. Insect Sci. Appl. 6: 341-347. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 530-537 A NEW GENUS NANCYANA AND NINE NEW SPECIES WITH A REVIEW OF THE RELATED GENUS RHOGOSANA (HOMOPTERA: CICADELLIDAE) PAUL H. FREYTAG Department of Entomology, University of Kentucky, Lexington, Kentucky 40546-0091. Abstract. —The new genus Nancyana and nine new species, N. agitata, N. gadouae, N. lubrica, N. fernandezi, N. fasciata from Venezuela; N. bordoni from Argentina; N. curva, N. isoclada, N. abluta from the Guyanas are described. Two species transferred from Rhogosana are also included in this genus, N. aldeia (DeLong) (type species) new com- bination, and N. duida (DeLong) new combination. The other species added to Rhogosana by DeLong are transferred as follows: Folicana amazona (DeLong) new combination (= marra Freytag) new synonym, and F. fosteri (DeLong) new combination (= robusta Osborn) new synonym, and Gypona (Marganalana) brazilia (DeLong) new combination. Key Words: pona On reviewing the species of Rhogosana Osborn it was found that all of the species added to this genus (DeLong 1975, 1981) belong to different genera. In this paper a new genus is set up for two of the species and nine new species. This genus 1s closely related to Rhogosana but can be separated from it on external color pattern and male genitalic characters. The remaining three species are trans- ferred to other genera as follows: brazilia DeLong 1975 (transferred to Gy- pona Germar in the subgenus Marganalana Metcalf, New Combination. amazona DeLong 1981 transferred to Fo- licana DeLong and Freytag (= marra Frey- tag 1979), New Combination and New Syn- onym. fosteri DeLong 1981 transferred to Foli- cana (= robusta Osborn 1938), New Com- bination and New Synonym. This leaves Rhogosana monotypic with just the type species rugulosa (Osborn). This species appears quite primitive being very Cicadellidae, Gyponinae, leafhoppers, Nancyana, Rhogosana, Folicana, Gy- large and unicolorous, and unique in that the male genitalia are quite different from other genera in this subfamily. Genus Nancyana, New GENUS Crown short, broadly rounded, three times as wide between eyes at base as median length, with a definite foliaceous margin, disc smooth usually without striae. Head narrower than pronotum. Ocelli prominent, slightly closer to median line than to eyes, slightly nearer posterior than anterior mar- gin of head. Aedeagus with basal processes (paraphyses) and two pair of subapical pro- cesses. Pygofer bifurcate at apex. Usually unicolorous, greenish brown or brown on head and pronotum; forewings darker brown with many spots of brown or black, some- times spots on forewings forming a pattern. Type species: Rhogosana aldeia DeLong. This genus is closely related to Rhogosana and Folicana, but the color pattern and the male genitalia of the species included in this genus are quite different and warrant the VOLUME 92, NUMBER 3 separation of this genus. A comparison of some of the morphological characters of these three genera are given in Figs. 1-6. Most species of this genus can easily be rec- ognized by being over 12 mm in length, with the head foliaceous, with the head, prono- tum and scutellum smooth and evenly col- ored; and with the forewings darker colored and usually speckled or spotted. This genus is named for my first wife who died in 1984. Key TO THE GENUS NANCYANA (Males of gadouae, lubrica, fernandezi, fasciata, bordoni and curva; and females of abluta, agitata, isoclada and duida are not known.) Pe Malesinm women cries cielnainiar waimeities ofreraye 2 1’. Females 2. Aedeagus with both | pairs sot Sutapical pro- cesses sharply pointed (Fig. 34) PE Ssh aten Rroatiencs duida (DeLong) 2'. Aedeagus with one pair of subapical pro- cesses sharply pointed, other pair truncate at apex (Fig. 24) 3. Aedeagus with both pairs of subapical pro- cesses equal in length and lying in same plane in lateral view (Fig. 19) isoclada sp. n. 3’. Aedeagus with both pairs of subapical pro- cesses not equal in length or not lying in same plane in lateral view (Figs. 14, 29) ......... 4 4. Aedeagus with outer pointed subapical pro- cesses wider at base and extending ventrally (ES GSe SecA) soerrscs vor eras sinyeeele. cece agitata sp. 0. 4’. Aedeagus with outer pointed subapical pro- cesses not greatly wider at base and extending dorsally (Figs. 28, 29) ....-... 5 5. Aedeagus with outer pointed subapical pro- cesses shorter than inner truncate subapical processes (Fig. 29) ........... aldeia (DeLong) 5’. Aedeagus with outer pointed subapical pro- cesses nearly same length as inner truncate subapical processes (Fig. 24) .. abluta sp. n. 6. Forewing with a distinct pattern as in Fig. 43 Mee eas eee reese s Bice ROSE E rs curva sp. n. 6h Eon winewaiha an indistinct pattern (Figs. 38- AD) hee apres ote 7. Forewing shiny, polished seventh sternum as in Fig. 12 Ni chs raaets bordoni sp. n. 7’. Forewing not shiny or polished; seventh ster- num with posterior margin usually more deeply emarginate (Figs. 10, 11) .......... 8 8. Seventh sternum with median triangular (Fig. OQ) Peat nastite oe Recerca ce lubrica sp. n. 5311 = ~) é : = ; [mm 3 5 RHOGOSANA NANCYANA FOLICANA 10 i 12 FERNANDEZI FASCIATA BORDONI Figs. 1, Rhogosana rugulosa (Osborn). 1. Dorsal view Se pronotum and scutellum. 2. Lateral view of head, pronotum and scutellum. Figs. 1-6 drawn to same scale. Figs. 3, 4. Nancyana aldeia (DeLong). 3. Dorsal view of head, pronotum and scutellum. 4. Lateral view of head, pronotum and scutellum. Figs. 5, 6. Folicana zella Freytag. 5. Dorsal view of head, pronotum and scutellum. 6. Lateral view of head, pronotum and scutellum. Figs. 7-12. Female seventh sternum, ventral view. 7. Nancyana aldeia (DeLong). 8. N. gadouae sp. n. 9. N. lubrica sp. n. 10. N. fernandezi sp. n. 11. N. fasciata sp. n. 12. N. bordoni sp. n. All drawn to the same scale. 8’. Seventh sternum with median rounded and emarginate (Fig. 11) ae 9 9. Seventh sternum with median obviously emarginate (Fig. 10) _fernandezi sp. n. 9’. Seventh sternum only lightly emarginate (Fig. 118 D rahe ee ee ne Re ae 10 10. Seventh sternum with median same length as lateral margins (Fig. 7) . aldeia (DeLong) 10’. Seventh sternum with median shorter than lateral margins (Fig. 8) .......... 11 11. Seventh sternum with lateral margins round- CU et, a ncusancine cere tecatie severe gadouae sp. n. 11’. Seventh sternum with lateral margins point- CO cir ores ele wicpepecneria es wttoee fasciata sp. n. Nancyana duida (DeLong) New CoMBINATION (Figs. 33-37) Rhogosana duida DeLong, 1975 (type lo- cality— Mt. Duida, Venezuela). Length of male 15 mm, female unknown. Crown broadly rounded, three times as wide at base between eyes as median length. Color: Head, pronotum and scutellum uniformly dull yellow brown. Forewings brown mottled with creamy brown spots. Male genitalia: Pygofer rounded, with tri- angular lobe at apical margin. Genital plate long, nearly four times as long as wide, trun- cate at apex. Style with apex blunt, nearly truncate but slightly protruding on dorsal and ventral margins. Aedeagus with broad shaft, two pairs of subapical processes, half length of shaft, sharply pointed at apex; base robust, with a pair of basal processes from base to ventral side of shaft extending to pointed apices two thirds length of shaft. Type: Holotype male, in the American Museum of Natural History. Note: The holotype was examined and used for the illustrations. No other speci- mens of this species have been seen. The paratype male from Brazil was not exam- ined and can not be verified as belonging to this species. This species is similar to a/deia, except slightly larger and with distinct male genitalia. Nancyana aldeia (DeLong) New COMBINATION (Figs. 7, 28-32, 38) Rhogosana aldeia DeLong, 1975 (type lo- cality—Shudihar R., British Guiana). Length of male 14 mm, female 17 mm. Crown broadly rounded, three times as wide at base between eyes as median length. Color: Head, pronotum and scutellum uniformly dull yellow (appears faded from a yellowish green). Forewings dark brown with many speckles of lighter brown and yellow. Male genitalia: Pygofer rounded with two PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON truncate apical lobes. Genital plate long, four times as long as wide, with rounded apex. Style with apical arm more than five times as long as wide, apex slightly expanded into a foot-like apex. Aedeagus with robust base, a pair of basal processes from base to ventral side of shaft, extending two thirds length of shaft; shaft tubular, ventral margin con- cavely depressed, six times as long as wide, expanded slightly at apex with two pair of subapical processes, outer pointed pair curved dorsad and laterad, inner bluntly rounded pair nearly paralleling shaft, gono- pore apical. Female genitalia: Seventh sternum (Fig. 7) more than twice as wide as median length; posterior margin excavated between round- ed lateral lobes and median rounded pro- jection, latter with slight median emargi- nation. Type: Holotype male, in the American Museum of Natural History Collection. Notes: Other specimens seen are one male, GUYANA, Esseq., 6 mi. S. Wineperu, Pi- erewana Is., March 8-16, 1969, Duckworth & Dietz, and one female, GUYANA, Es- seq., Plantain Is., March 25-26, 1969, Duckworth & Dietz, in the U.S. National Museum Collection. The holotype was used for the illustrations. The paratype male from Brazil was not seen, and can not be verified at this time as being this species. Nancyana agitata sp. n. (Figs. 3, 4, 13-17) Length of male 15-15.5 mm, female un- known. Similar to a/deia in general size and color pattern, but with distinct male geni- talia. Male genitalia: Pygofer and genital plate (Figs. 16, 17) similar to aldeia. Style (Fig. 15) with apical arm more than 5 times as long as wide; apex expanded, foot-shaped, pointed at toe and heal. Aedeagus (Figs. 13, 14) with robust base; a pair of basal pro- cesses from base extending to ventral side of shaft, half way to apex of shaft; shaft tubular, ventral margin concavely de- VOLUME 92, NUMBER 3 pressed, six times as long as wide, expanded slightly at apex with two pairs of subapical processes, outer pointed pair thicker at base, curving ventrad and laterad, inner bluntly rounded pair close to shaft, nearly same width to apex; gonopore apical. Holotype male: VENEZUELA, Sn. Pedro de Cataniapo, T. F. Amazonas, 100 m, Sep- tember 23-27, 1981, En la Luz, G. L. Garcia Coll., in the Universidad Central de Ven- ezuela Collection. Paratypes: Two males, VENEZUELA, Bolivar, carret, Caicara, San Juan de Manapiare, KM 150, 300 m, March 21, 1978, Gadou Coll., one in the Univer- sidad Central de Venezuela Collection and one in the University of Kentucky Collec- tion. Note: This species differs from the other known species of the genus by having the first pair of subapical processes of the ae- deagus thicker at the base giving the outer margin a wavy appearance, and the second pair of subapical processes uniformly nar- row and slightly longer than the first pair. Nancyana isoclada sp. n. (Figs. 18-22, 39) Length of male 13.5 mm, female un- known. Similar to a/deia, but slightly small- er and with distinct male genitalia. Color: Head, pronotum and scutellum brown. Forewings brown, spotted with creamy brown overall and some darker brown spots in apical cells. Male genitalia: Pygofer robust with apical margin with two truncate lobes, dorsal lobe longer. Genital plate three times longer than broad, apex broadest and truncate. Style long, with apex slightly expanded. Aedeagus with stout shaft with two pair of equally long subapical processes, outer pair pointed at apex, inner pair truncate at apex; base ex- panded with basal processes extending to two thirds length of shaft, bending to ventral side of shaft. Holotype male: GUYANES, Ile de Touenké, 19-21-XI-1975, Itani (Guyanes) 533 AGITATA Rov ; DF Ha y) Yi 16 coat \7 ISOCLADA ee A oe Je ‘ | } | / ee ite} 19 L 20 21 Ress 22 ABLUTA — N \ : 8 YY LY a (—*s Y 27 23 «24 2 26 = ALDEIA “a \ es \\ we ue YA io Y/ 28 «429°~=—(30 3i Se Figs. 13-17. Nancyana agitata sp. n. 13. Ventral view of aedeagus. 14. Lateral view of aedeagus. 15. Lateroventral view of style, with lateral view of apex. 16. Ventral view of genital plate. 17. Lateral view of genital capsule. Figs. 13-32 all drawn to the same scale. Figs. 18-22. Nancyana isoclada sp. n. 18. Ventral view of aedeagus. 19. Lateral view of aedeagus. 20. Lateroventral view of style, with lateral view of apex. 21. Ventral view of genital plate. 22. Lateral view of genital capsule. Figs. 23-27. Nancyana abluta sp. n. 23. Ventral view of aedeagus. 24. Lateral view of aedeagus. 25. Lateroventral view of style, with lateral view of apex. 26. Ventral view of genital plate. 27. Lateral view of genital capsule. Figs. 28-32. Nancyana aldeia (DeLong). 28. Ven- tral view of aedeagus. 29. Lateral view of aedeagus. 30. Lateroventral view of style, with lateral view of apex. 31. Ventral view of genital plate. 32. Lateral view of genital capsule. Mission, M. Boulard, P. Jauffret et P. Pom- panon Coll., in the Muséum Paris. Note: This species can be separated from the other known species of the genus by the subapical processes of the aedeagus being equal in length and both curving dorsad. 534 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON DUIDA fe rae duida Pan 33. Ventral view of Pas 34. Lateral view of aedeagus. 35. Figs. 33-37. = I 5 Be | Lateroventral view of style, with lateral view of apex. 36. Ventral view of genital plate. 37. Lateral view of genital capsule. All drawn to the same scale. Nancyana abluta sp. n. (Figs. 23-27) Length of male 13.2 mm, female un- known. Similar to isoclada, but with dis- tinct male genitalia. Color: Head, pronotum and scutellum brown. Forewings brown spotted with creamy brown. Male genitalia: Pygofer robust with apical margin with two truncate lobes, dorsal lobe slightly longer. Genital plate three times longer than broad, apex broadly rounded. Style long, with apex slightly expanded. Ae- deagus with stout shaft with two pair of sub- apical processes, outer pair pointed, nearly same length as inner pair, apex curving dor- sad, inner pair truncate at apex, paralleling shaft. Holotype male: GUYANES, Antécume- pata (Saut Kialo), 22-XI-1975, Itani (Guy- anes) Mission, M. Boulard, P. Jauffret et P. Pompanon, in the Muséum Paris. Note: This species can be separated from isoclada by the rounded apex of the genital plate and the subapical processes of the ae- deagus not paralleling each other. Nancyana gadouae sp. n. (Figs. 8, 40) Length of female 17.5 mm, male un- known. Similar to a/deia, except larger and with different shaped female seventh ster- num. Color: Head, pronotum and scutellum uniformly brown. Forewing dark brown speckled with lighter spots and apically with darker spots. Female genitalia: Seventh sternum nar- row, median lobe shorter than lateral lobes, with a slight median emargination. Holotype female: VENEZUELA, S. [er- rania] de Lema, 1200 m, V-1983, Gadou Leg., in the Universidad Central de Vene- zuela Collection. Notes: This species is larger and darker in color than a/deia and the female seventh sternum is much narrower. This species is named after Mrs. Marilou Gadou, an ex- cellent collector of Auchenorrhyncha. Nancyana lubrica sp. n. (Fig. 9) Length of female 19 mm, male unknown. Similar to a/deia, but much larger and with a distinct seventh sternum. Color: Head, pronotum and scutellum greenish brown, spotted with brown. Spots on pronotum appear as punctures. Fore- wings dark brown, heavily spotted overall with creamy brown, some blackish brown spots at apex. Female genitalia: Seventh sternum with VOLUME 92, NUMBER 3 median triangular lobe extending length of lateral lobes. Holotype female: VENEZUELA, T. F. Amazonas, San Carlos de Rio Negro, 10- XII-1984, R. Brown Coll., in the Univer- sidad Central de Venezuela Collection. Note: This very large species is darker than other known species and the female seventh sternum is triangular instead of rounded. Nancyana fernandezi sp. n. (Fig. 10) Length of female 17.5 mm, male un- known. Similar to a/deia, except larger and with a different shaped female seventh ster- num. Color: Head, pronotum and scutellum uniformly light brown. Forewings dark brown, heavily spotted with clear or creamy yellow spots. Female genitalia: Seventh sternum with median lobe extending equal to lateral lobes, emarginate medially. Holotype female: VENEZUELA, Bolivar, Km 107, El Dorado, Santa Elena, 520 m, 19-VIII-1957, F. Fernandez Y. & C. J. Ro- sales Colls., in the Universidad Central de Venezuela Collection. Note: This species is named after the late Dr. F. Fernandez-Yepes an excellent ento- mologist and friend. Nancyana fasciata sp. n. (Figs. 11, 41) Length of female 15 mm, male unknown. Similar to a/deia in size and color, but with a different shaped female seventh sternum. Color: Head, pronotum and scutellum uniformly light brown. Forewings dark brown spotted and speckled with lighter and darker brown. Female genitalia: Seventh sternum with a small rounded median lobe, shorter than lateral lobes, slightly emarginate medially. Holotype female: VENEZUELA, Bolivar, El Bochinche, 200 m, 6—13-XII-1974, J. Salcedo and R. E. Dietz Colls., in the Uni- versidad Central de Venezuela Collection. Note: This species is very similar to al- 535 deia, but the seventh sternum of the female is overall smaller with smaller lateral lobes. Nancyana bordoni sp. n. (Figs. 12, 42) Length of female 17 mm, male unknown. Similar to aldeia, except larger, with darker markings on wings and a different female seventh sternum. Color: Head, pronotum and scutellum uniformly light brown. Forewings dark brown, spotted with light brown basally, blackish brown apically, mostly shiny, ap- pearing polished. Female genitalia: Seventh sternum with a rounded median lobe as long as lateral lobes, slightly emarginate medially. Holotype female: ARGENTINA, Pto. Iquazu, Misiones, 100 m, 25-XI-8-XII- 1983, C. Bordon Coll., in the Universidad Central de Venezuela Collection. Note: This species is named for Dr. Carlos Bordon, a well-known entomologist and the collector of this species. Nancyana curva sp. n. (Fig. 43) Length of female 16.5 mm, male un- known. Similar to a/deia, except larger, with a distinct color pattern and different female seventh sternum. Color: Head, pronotum and scutellum uniform light brown. Forewings brown spotted with creamy yellow or white, with a large solid brown half circle area near mid- dle along costal margin, and darker blackish brown spots in apical cells. Female genitalia: Seventh sternum sim- ilar to a/deia in shape, except median lobe extending beyond length of lateral lobes. Holotype female: GUY ANE, Riviere-Ca- mopi, Mont Alikene, 11-XI-1969, Piége Lumineux, Guyane Mission, Balachowsky- Gruner, Oct.-Nov. 1969, in the Muséum Paris. Note: This species is easily separated from the presently known species of this genus by the very distinct brown half-circle on the forewing, whereas in most species the fore- 536 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 38-43. Nancyana spp., lateral view. 38. N. aldeia (DeLong), male from Guyana. 39. N. isoclada sp. n., holotype. 40. N. gadouae sp. n., holotype. 41. N. fasciata sp. n., holotype. 42. N. bordoni sp. n., holotype. 43. N. curva sp. n., holotype. Length of each given in species description, approximately 2x. wings are just generally spotted and speck- led with only a slight pattern. ACKNOWLEDGMENTS I thank the following for the loan of the material used in this study: James P. Kra- mer, U.S. National Museum of Natural His- tory; M. Boulard, Muséum Paris; F. Fer- nandez-Yepes, Museo Instituto de Zoologia Agricola, Universidad Central de Venezue- la; Carlos Bordon, El Lemon, Venezuela and Marilo Gadou, El Lemon, Venezuela. I also VOLUME 92, NUMBER 3 thank R. T. Schuh, American Museum of Natural History for the loan of the type ma- terial. This paper is published with the ap- proval of the Director of the Kentucky Ag- ricultural Experiment Station as journal article no. 89-7-202. LITERATURE CITED DeLong, D. M. 1975. The genus Rhogosana (Ho- moptera: Cicadellidae) with descriptions of three new species. Ohio J. Sci. 75(3): 126-129. 1981. New species of Rhogosana and Pona- 537 na, (Homoptera: Cicadellidae) from Central and South America. Entomol. News 92(1): 17-22. DeLong, D. M. and P. H. Freytag. 1971. Studies of the Gyponinae: Rhogosana and four new genera, Clinonella, Tuberana, Flexana and Declivara. J. Kansas Entomol. Soc. 44: 313-324. 1972. Studies of the Gyponinae: The genus Folicana and nine new species. J. Kansas Ento- mol. Soc. 45: 282-295. Freytag, P.H. 1979. Additions to the genus Folicana (Homoptera—Cicadellidae—Gyponinae). J. Kansas Entomol. Soc. 52: 810-819. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 538-543 THE GENUS XIPHOGRAMMA, ITS OCCURRENCE IN NORTH AMERICA, AND REMARKS ON CLOSELY RELATED GENERA (HYMENOPTERA: TRICHOGRAMMATIDAE) JOHN D. PINTO Department of Entomology, University of California, Riverside, California 92521. Abstract.—The genus Yiphogramma is briefly reviewed and compared to related genera, Chaetogramma and Brachygrammatella. Xiphogramma fuscum n. sp., the first species from the New World, is described from southwestern North America. A key to the species of Xiphogramma and a description of the male genitalia are included. Key Words: The genus Xiphogramma Nowicki, with three species included, has been known only from the Old World (Doutt 1974, Hayat 1980). This paper describes a fourth species, from North America, and includes a key to the known fauna. The characters of the new species indicate the artificiality of Chaeto- gramma Doutt, as defined by Hayat (1981). Arguments for and against synonymy of these genera are presented. Hosts of Xiphogramma are unknown. The species described here emerged from grape leaves containing eggs of both Cicadellidae and Miridae. Of its related genera, Chae- togramma and Brachygrammatella Gi- rault, hosts of only the latter are known. Species of Brachygrammatella have been associated with eggs of Cicadellidae, Mem- bracidae and Miridae (Doutt 1968, Viggiani 1968, Yousuf & Shafee 1987). Xiphogramma Xiphogramma was described by Nowicki (1940) for the unique European trichogram- matid, ¥. holorhoptra Nowicki. He afforded the species generic status on the basis of oOvipositor structure—‘“‘abdomen with a powerful ovipositor occupying its entire Hymenoptera, Trichogrammatidae, Yiphogramma taxonomy length and protruded for more than a half of abdomen’s length: the valvae are much broadened before the tip, curved upwards and sabre-like.”’ All Xiphogramma species have an exserted, curved ovipositor, but it is not necessarily as long as in X¥. holorhop- tra. Other features of the genus include an- tenna with two anelli, two subequal, closely- appressed funicle segments, one-segmented club, club not widest at base; maxillary palp one-segmented; vertex wrinkled, vaulted; wing disk densely setate, vein tracks, if pres- ent, usually becoming obsolescent apically; stigmal vein broad, subsessile, marginal vein not densely setate; a dark macula of varying size and intensity beneath venation; tarsal segment I of middle leg elongate, distinctly longer than that of hind leg. An African species, X. anneckei, was added by Doutt (1974), and X. indicum, from India, was described by Hayat (1980). The new species described below occurs in arid and semiarid regions of southwestern North America. Males and females have been collected. Heretofore, the only male of Xiphogramma known was the allotype of X. anneckei. VOLUME 92, NUMBER 3 539 Figs. 1, 2. 0.1 mm. Xiphogramma fuscum, NEW SPECIES The description is based on critical point dried (for color and body length measure- ments) and slide mounted specimens. Quantitative data represent means taken from three specimens from the type locality; the mean is followed by a range, in paren- theses, if variation is considerable. Signifi- cant intraspecific variation among locales was not detected. Female (Fig. 1).—Body length 1.08 mm; 0.95 mm excluding exserted ovipositor. Color: Primarily dark brown except as Xiphogramma fuscum, female. 1, Lateral view (appendages removed). 2, Forewing. Scale bar = follows: head with linear yellow area along medial rim of eye; frons yellow brown; face yellow brown to dark brown. Antenna with scape primarily pale yellow, margined with brown; pedicel light brown; funicle yellow brown; club pale brown or yellow brown. Thorax with narrow linear yellow marking at midline of pronotum and immediately lateral to mid-lobe of mesoscutum; mes- epimeron, mesepisternum with at least some yellow; metanotum, propodeum, segment I of gaster yellow except laterally. Legs brown except apex of coxae, trochanters, base and 540 apex of femora and tibiae, and tarsi whitish (apical tarsal segment may be pale brown). Venation of fore wing bicolored; stigmal vein, apical half of marginal vein pale brown; remainder of venation pale yellow. Wings hyaline except a small fumate area beneath stigmal vein. Head. Length and width subequal; vertex vaulted, arched above eyes, wrinkled; scrobes relatively deep; lower margin of tor- ulus coincident with ventral margin of eye; malar space ca. 0.6 eye length. Mandible with four teeth. Antenna (Fig. 3) with second anellus very short, inconspicuous, closely appressed to funicle; Fl, F2 subequal in length; length/ width of segments as follows: scape—3.42, pedicel—2.07, Fl —0.62 (0.58-0.67), F2— tenna moderately setate, setae longer, stout- er on pedicel and funicle; Fl with 1 trans- verse placoid sensillum, distal portion of sensillum curved toward apex of segment; F2 with 3 oblique placoids; F1, F2 each with several basiconic peg sensilla on apical mar- gin; club with 12 linear placoids, club also with many thin-walled setiform sensilla at apical halfand several basiconic peg sensilla near middle. Thorax. Forewing (Fig. 2) broad, subob- late apically, 0.55 as broad as long, with a very short fringe; venation attaining 0.43 length of wing; setation on disc apical to venation dense; vein tracks becoming ob- solescent apically; RS, represented by 2-3 setae, 2 setae in line with RS, on apex of stigmal vein; basal vein track with 2 setae; costal cell, narrow, with setae along apical half of anterior margin; relative length of veins as follows: subcostal—29, premargin- al—14, marginal—15, stigmal—6; marginal vein stout, slightly, gradually widened api- cally with about 8 setae; stigmal vein poorly defined, stout, subsessile, ca. as long as broad; premarginal vein with 2 setae; sub- costa with | seta at middle. Hindwing mod- erately broad, maximum width of disk 1.4 x length of longest posterior fringe setae; with 2 distinct setal tracks just behind anterior PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON margin, remainder of disk with many scat- tered setae. Thoracic setae elongate, stout; mid-lobe of mesoscutum with 4 setae of subequal length; scutellum with posterior pair of setae ca. 1.5 x as long as anterior pair; lateral lobe of mesoscutum, axilla each with | elongate seta. Foretibia with 5 “teeth” on anterior sur- face, basal 2 weaker than apical ones. Fore and middle femora with a relatively elon- gate seta ventroapically. Hind femur dis- tinctly broader than others; hind trochanter swollen dorsally. Tibial spurs not plumose. Relative length of coxae, trochanters, fem- ora, tibiae, and (tarsi) as follows: fore leg— 26:11:41:32:(9:11:11); middle leg—21:14: 40:49:(19:14:13); hind leg—35:17:40:48:(1 3: 13:13); length of apical tibial spines 4, 7, 8, resp.; length of apical setae on fore and mid- dle femora 8, 11, resp. Gaster (Fig. 1): Gaster (excluding ovi- positor) elongate, apically acuminate, ca. 1.5 as long as thorax; hypogynium attain- ing 0.8 length of gaster. Ovipositor elongate, running along entire length of gaster and beyond, broadly curved dorsally, its length 2.6-3.1 length of mid and hind tibiae; gon- oplac elongate, densely setate, comprising 0.36 length of entire ovipositor, almost its entire length extending beyond apex of gas- ter; gonangulum small, subtriangular, basal 0.10 of ovipositor extending anterior of gonangulae. Male.—As in female except as follows: Head (in 3 of 6 dried specimens) paler with vertex yellow lateral and dorsal to scrobes. Antenna (Fig. 4) with pedicel, Fl more elongate; with only 1 placoid sensil- lum on F2, 5 placoids on club; club seg- ments incompletely fused, an obsolescent U-shaped suture on anterior surface be- tween basal and apical placoids; length/ width of segments as follows: scape—3.50, pedicel— 2.41, Fl —1.07, F2—0.75, club— Dai9y Genitalia (Fig. 5) very similar to that de- scribed for Chaetogramma maculata (Ha- yat 1981, Fig. 5); very elongate, narrow, VOLUME 92, NUMBER 3 541 Figs. 3-5. 6.7 as long as wide, length subequal to that of hind tibia; aedeagus fused to genital capsule, apodemes absent; base of genital capsule attenuate, its ventral region extend- ing anteriorly; posterior border of antero- dorsal aperature not sclerotized, poorly in- dicated; genital capsule with 2 short, stout spines apicoventrally; volsellae broad, un- armed, apically truncate; gonostyli not dif- ferentiated. Types.— Holotype female; from Mexico, Sonora, Caborca; emerging in laboratory from grape leaves collected 2-VI-1989; L. Drake, collr.; deposited in the United States National Museum. Allotype male; same data as holotype; deposited in the United States National Museum. Additional specimens from Caborca (4 22, 7 44) are designated paratypes and deposited as follows: 1 9, 1 6, British Museum (Natural History); 1 2, 1 6, Canadian National Collection (Ottawa); 2 22, 5 46, University of California (River- side). The holotype, allotype and 6 of the Xiphogramma fuscum. 3, Right female antenna (anterior surface). 4, Right male antenna (same). 5, Male genitalia (ventral). Scale bar = 0.1 mm. paratypes (3 2°, 3 44) are mounted on glass slides in Canada balsam. One ¢ and 4 é4 paratypes are card mounted. Diagnosis. — XY. fuscum is similar to_X. in- dicum. Characters separating them are pre- sented in the key below. The most similar species in North America is Chaetogramma occidentalis Doutt. Ovipositor length (short, not exserted in C. occidentalis) separates fe- males. Genital structure will distinguish males. In X. fuscum, the aedeagal apodemes are not expressed and the base of the genital capsule is attenuate anteriorly (Fig. 5); in C. occidentalis the apodemes are well devel- oped, and the genital capsule is truncate ba- sally (Pinto, unpubl.). Also in X. fuscum the two funicle segments are distinct and not partially fused as in C. occidentalis. Etymology.—The specific name is Latin and refers to the dark brown body color. Host.—The host of X. fuscum is un- known. Specimens from Caborca, Sonora, and Tonopah, Arizona, emerged from grape 542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON leaves harboring eggs of leafhoppers and mirid bugs (Parthenicus). Records.—19 99, 12 64. MEXICO. Baja California Sur: Ciudad Constitucion, 11 km N.; 1 2; 27-X-1983; screen sweeping desert vegetation; J. D. Pinto. Sinaloa: Mazatlan, 12 mi. N.; 1 9; 25-X-1982. Sonora: Cabor- ca; 5 92, 8 446; emerging from grape leaves coll. 2-VI-1989, 27-VII-1989 & 2-VIII- 1989; L. Drake. Hermosillo; | 2, 1 4; emerg- ing from grape leaves coll. 2-VI-1989, 9-VII- 1989; L. Drake; & 4 99; 6-X-1985: D. Gon- zalez. UNITED STATES. Arizona: Stan- field, 1 2, emerging from grape leaves coll. 11-VII-1989; L. Drake. Sycamore Canyon, 9 mi. W. Pena Blanca Lk. (Santa Cruz Co.), 4100 ft. elev.; 1 9; 12-VII-1983; R. Ander- son. Tonopah; | 2, | 4; emerging from grape leaves coll. 21-VI-1989; L. Drake. Califor- nia: Baker, 11 km N.; 1 2; 30-HI-1989; screen sweeping desert vegetation; J. D. Pinto. He- met, E of (4000 ft. elev.); 1 2; screen sweep- ing Adenostoma sparsifolium Torr.; 30-VI- 1983; R. Velten. Texas: Ben Bolt, 8 mi. W. (La Copita Res. Sta.); 3 29, 2 64; 20-V-1987; screen sweeping; J. B. Woolley. KEY TO THE SPECIES OF XTIPHOGRAMMA (FEMALES) 1. Gaster elongate, its length at least 2 that of thorax; exserted portion of ovipositor greater than half gaster length oy : ae — Gaster shorter, its length about 1.5 that of thorax, subequal to length of head and thorax combined; exserted portion of ovipositor not greater than half gaster length Dorsum of gaster primarily dark brown, weak- ly marked with yellow at tergal margins only. Marginal vein widened apically. Length of fu- nicle equal to or slightly shorter than pedicel. Europe (Poland) .... X. holorhoptra — Dorsum of gaster primarily yellow, marked with brown; marginal vein not widened apically. Length of funicle distinctly greater than that of pedicel. Africa (South Africa, Tanzania, Ivory COaSt) ee aceeeet ane SHE ee ee eee CIINECKEL 3. Hypogynium elongate, attaining apex of gaster. Antenna with F1 as long as or longer than wide, longerithanP2! India scans coke X. indicum — Hypogynium shorter, attaining 0.8 gaster length. Antenna with FI distinctly wider than long, subequal in length to F2. Southwestern North AMeniGal serena Bare clerk? X. fuscum, N. sp. > te DISCUSSION Xiphogramma is closely related to Brach- ygrammatella and Chaetogramma. Wing and antennal structure is similar in all three genera. Brachygrammatella is distinguished by the densely setate marginal vein, and minor antennal differences (club widest at base, funicles much broader than long); also, the Ovipositor in this genus does not project beyond the gaster (see Doutt and Viggiani 1968). Doutt (1974) described Chaetogramma for an African and a North American species, which he separated from Xipho- gramma primarily by the short ovipositor (not extending beyond gaster), and the fused or partially fused funicle segments. His statement that the number of anelli also sep- arate the two (1 in Chaetogramma, 2 in Xiphogramma) is incorrect. There are 2 anelli in both, as well as in Brachygram- matella. Hayat (1981) divided Chaetogramma into two subgenera, the nominate, which in- cludes both of Doutt’s species, and Chae- togrammina, erected for C. maculata Hayat from India. Chaetogrammina was distin- guished primarily by its completely divided funicle segments, its more distinct vein tracks and better developed costal cell. Male genitalia, not compared by Hayat, provide another difference. In the nominate subge- nus distinct aedeagal apodemes are present (Pinto, unpubl.) as they are in Brachygram- matella (Viggiani 1971, Pinto, unpubl.). In Chaetogrammina, based on descriptions and figures in Hayat (1981) and Viggiani (1984), they are absent or poorly developed. Viggiani (1984) pointed out the similarity of male genital structure in Brachygram- matella and C. (Chaetogrammina) and, on this basis, questioned the validity of Chae- togramma. The male genitalia in Xipho- gramma cast further doubt on the validity of Chaetogramma. They are virtually iden- tical to that in C. (Chaetogrammina) mac- ulata. The only character now separating Xiph- VOLUME 92, NUMBER 3 ogramma and Chaetogramma 1s ovipositor length. The difference in length between _X. anneckei and X. holorhoptra on the one hand, and species of Chaetogramma on the other, although considerable, is bridged substantially by Y. indicum and X. fuscum. For example, in ¥. anneckei the exserted portion of the ovipositor is 0.8-0.9 the length of the gaster, and in C. maculata the ovi- positor does not extend beyond the gaster. In_X. fuscum, however, the exserted portion of the ovipositor is never more than 0.45 gaster length. Although synonymy is sug- gested, I hesitate at present for the following reason. Chaetogramma, as currently de- fined, is paraphyletic. It is distinguished from Brachygrammatella and Xiphogramma only by primitive traits (e.g. absence of a densely setate marginal vein, and a short oviposi- tor). Synonymizing it with Yiphogramma simply results in a larger paraphyletic unit, more difficult to characterize than either is at present. Structure of the forewing, antenna and genitalia suggest that C. (Chaetogrammina) is closer to Yiphogramma than to its nom- inate subgenus. Moving this subgenus to Xiphogramma probably 1s appropriate. The only clearly derived traits currently justi- fying this are associated with the male gen- italia, however. Because the male genitalia are known in only one species of Xipho- gramma, I consider it premature to transfer Chaetogrammina and then define Xipho- gramma solely on male features. ACKNOWLEDGMENTS I thank Dan Gonzalez and John Luhman for providing several collections of the new 543 species, and Rob Velten for preparing spec- imens for study. LITERATURE CITED Doutt, R. L. 1968. The genus Brachygrammatella Girault (Hymenoptera: Tnchogrammatidae). Pan- Pac. Entomol. 44: 289-294. . 1974. Chaetogramma, a new genus of Trich- ogrammatidae (Hymenoptera: Chalcidoidea). Pan- Pac. Entomol. 50: 238-242. Doutt, R. L.andG. Viggiani. 1968. The classification of the Trichogrammatidae (Hymenoptera: Chal- cidoidea). Proc. Calif. Acad. Sci. (4th ser.) 35: 477- 586. Hayat, M. 1980. The genera Neocentrobiella and Xiphogramma from India, with descriptions of two new species (Hymenoptera: Trichogrammati- dae). Boll. Lab. Ent. Agr. Portici 37: 203-207. . 1981. The genera Chaetogramma and Lath- romeromyia from India, with descriptions of two new species (Hymenoptera: Trichogrammatidae). Boll. Lab. Ent. Agr. Portici 38: 73-79. Nowicki, S. 1940. Descriptions of new genera and species of the family Trichogrammidae (Hym. Chalcidoidea) from the Palearctic Region, with notes—Supplement. Zeit. Angew. Ent. 26: 624— 663. Viggiani,G. 1968. Ricerche sugli Hymenoptera Chal- cidoidea XVII. Nuove specie di Trichogrammati- dae. Boll. Lab. Ent. Agr. Portici 26: 251-262. 1971. Ricerche sugli Hymenoptera Chalci- doidea XXVIII. Studio morfologico comparativo dell’armatura genitale esterna maschile dei Trich- ogrammatidae. Boll. Lab. Ent. Agr. Portici 29: 181- DD . 1984. Further contribution to the knowledge of the male genitalia in the Trichogrammatidae (Hym. Chalcidoidea). Boll. Lab. Ent. Agr. Portici 41: 173-182. . 1984. Further contribution to the knowledge of the male genitalia in the Trichogrammatidae (Hym. Chalcidoidea). Boll. Lab. Ent. Agr. Portici 41: 173-182. Yousuf, M. and S. A. Shafee. 1987. Taxonomy of Indian Trichogrammatidae (Hymenoptera: Chal- cidoidea). Indian J. Syst. Entomol. 4(2): 55-200. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 544-547 OCCURRENCE OF SEXUAL MORPHS OF RUSSIAN WHEAT APHID, DIURAPHIS NOXTA (HOMOPTERA: APHIDIDAE), IN SEVERAL LOCATIONS IN THE SOVIET UNION AND THE NORTHWESTERN UNITED STATES Ion KirIACc, FRANCIS GRUBER, TAD POPRAWSKI, SUSAN HALBERT,! AND LESLIE ELBERSON (IK) All Union Research Institute for Biological Methods in Agriculture, Kishinev, Moldavia, USSR. (FG, TP) USDA European Parasite Laboratory, Behoust, France. (SH) University of Idaho Southwest Idaho R/E Center, 29603 U of I Lane, Parma, Idaho 83660, USA. (LE) University of Idaho, Department of Plant, Soils and Entomological Sciences, Moscow, Idaho 83843, USA. Abstract.—Diuraphis noxia were collected in North America and the Soviet Union during autumn 1989. Sexual forms constituted more than half of Moldavian and Crimean collections and about nine percent of collections near and between Odessa and Kherson, Ukraine. Six oviparae were found in Idaho and Oregon, but they represented less than | percent of total collections. No sexuales were found in the Soviet Republic of Kirghizia. Moldavian D. noxia colonies readily produced sexual forms under natural autumn con- ditions, whereas an Idaho isolate of D. noxia produced no males or oviparae after 10 weeks under a 6:18 (L:D) photoperiod at 10°C. Under a photoperiod of 8:16 (L:D) at 20°C, Moldavian D. noxia produced sexual forms, but Syrian, French, Turkish, Jordanian and Kirghizian populations did not. A Kirghizian population did produce sexuales and eggs at 16°C and a photoperiod of 14:10 (L:D). Key Words: Diuraphis noxia, Aphididae, sexuals There are two overwintering strategies among Aphididae. Some populations (termed holocyclic) produce males and Oviparae which must mate to produce vi- able overwintering eggs. Other populations (termed anholocyclic) overwinter in pro- tected locations as viviparous females, and no sexual morphs are produced. Some pop- ulations can utilize either strategy depend- ing upon climatic conditions. The Russian wheat aphid, Diuraphis nox- ia (Mordvilko) (Homoptera: Aphididae), a serious pest of small grains, is indigenous ' Order reprints from Susan Halbert. to the Middle East, the Soviet Union, Af- ghanistan and probably western China (Hewitt et al. 1984). Holocyclic populations of D. noxia are known from the Soviet Union (Grossheim 1914). Sexual forms have been described briefly by Grossheim (1914), but no formal descriptions exist. On 7 IX 1989, Dr. Manya B. Stoetzel found an ap- terous male in a laboratory colony main- tained at the USDA-ARS European Para- site Laboratory, Behoust, France, and originally collected in Kishinev and vicin- ity, Moldavia, USSR, 28 V—2 VI 1989 on wheat and barley by Dr. Tad Proprawski and Francis Gruber (M. B. Stoetzel, per- sonal communication). Additional males VOLUME 92, NUMBER 3 and oviparae have been obtained from this colony. Diuraphis noxia was recently intro- duced into South Africa and North America (Stoetzel 1987, Walters 1984). Prior to this article, no sexual forms of D. noxia have been reported from North America, al- though D. noxia has been reported as far north as 50° latitude in Canada (Jones et al. 1989). No sexual forms have been reported from South Africa. The purpose of the surveys was to com- pare occurrence of the various morphs of D. noxia in the northwestern United States and the Soviet Union where D. noxia 1s native. Our preliminary attempts to force various populations of D. noxia to produce sexuales under laboratory conditions are presented here to support field observations and are not intended to be definitive ex- periments on the nature of triggering mech- anisms for development of sexuales. METHODS Live D. noxia were collected on wheat and barley from two fields in Moldavia (Oc- tober 25-27, 1989), four fields in the Cri- mean Peninsula (October 31—November 2, 1989), five fields in the southern Ukraine (near and between Odessa and Kherson) (November 2-4, 1989), seven fields in the Kirghiz Inner Tian Shan Range of south central USSR (September, 1989), six fields in the Treasure Valley of Canyon County, Idaho and Malheur County, Oregon (No- vember 14-16, 1989) and 2 fields in the Palouse area of Latah and Nez Perce Coun- ties, Idaho (November 25-27, 1989). In the Ukraine and Moldavia D. noxia is quite rare, so every colony found was collected. In the Treasure Valley where D. noxia is more abundant, infested plants were select- ed along a 100 m transect within each field, and additional plants were collected in heavily infested areas of several fields. In the Palouse area, wild oat plants with ob- vious damage symptoms along the perim- eter of a field previously in barley (Latah County) and within a winter wheat field (Nez 545 Perce County) were collected. In Kirghizia, heavily infested wheat and barley plants were selected. In the Soviet Union and the Treasure Valley, immature aphids were kept on fresh wheat plants in plastic containers until they became adults. Nymphs with wing pads which died before reaching maturity were recorded as alatae. Other nymphs which died prior to reaching maturity were recorded as undifferentiated nymphs. Adult aphids were examined and preserved in 70% ethanol. In the Palouse, only adults were examined. Voucher specimens are on de- posit at the All Union Research Institute for Biological Methods in Agriculture, Kishi- nev, Moldavia, U.S.S.R.; the USDA Eu- ropean Parasite Laboratory, Behoust, France, the University of Idaho Southwest Idaho Research and Extension Center, Par- ma, Idaho, U.S.A.; the Pasteur Institute, Paris, France; and the USDA-ARS System- atic Entomology Laboratory, Beltsville, Maryland, U.S.A. In preliminary experiments, we have made attempts to force various populations of D. noxia to produce sexual morphs by subjecting them to short days and cool tem- peratures (Blackman and Eastop 1984). In Idaho, one of us (S.H.) maintained North American D. noxia individually in petri dishes supplied regularly with fresh leaves on moist cotton subjected to a photoperiod of 6:18 (L:D) at 10°C from February—April, 1988. A similar experiment was done at the USDA-ARS European Parasite Laboratory, Behoust, France using Moldavian, Syrian, Jordanian, French, Turkish and Kirghizian D. noxia kept at a photoperiod of 8:16 (L: D) at 20°C from October-December, 1989 (F.G. and T-.P:). RESULTS AND DISCUSSION Collections in the USSR.—No alatae or alatoid nymphs were found in the Ukraine or Moldavia (Table 1). In Moldavia all 26 of the adult D. noxia found or reared were oviparae. In the Crimean Peninsula, the only location where males were found, four fifths 546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Morphs of Diuraphis noxia (Mordvilko) found in 6 locations in the southern Soviet Union and northwestern United States. 1989. Morphs Found Number Alate Apterous Undifferenti- Location Month of Collection of Sites Viviparae! Viviparae Oviparae Males ated Nymphs? Moldavia October 2 0 0 26 0 1 Crimea November 4 0 6 21 3 6 Ukraine? November 5 0 89 9 0 0 Kirghizia September 7 76 843 0 0 43 Treasure Valley* November 6 107 401 5 0 70 Palouse November 2 8 193 1 0 - ‘Includes alatoid nymphs in Treasure Valley collections. 2 Aphids collected were maintained in cages to allow them to become adults before they were scored. Nymphs without wing pads which died before becoming adults are scored as undifferentiated nymphs. 3 Fields near and between Odessa and Kherson. 4 Fields in Canyon Co., Idaho and Malheur Co., Oregon. ‘ Fields in Latah and Nez Perce Counties, Idaho. Only adults were counted. of the 30 adult aphids recovered were sexual forms (21 oviparae, 3 males). In the south- ern part of the Ukraine 9 of the 98 adult D. noxia recovered were oviparae. No sexuales were found in Kirghizian collections. Diuraphis noxia is not common in Mol- davia or the Ukraine in the autumn and is not considered an important pest. The in- festations we observed affected isolated plants or patches up to 2 m in diameter. Diuraphis noxia was most common in vol- unteer grain. In winter wheat, infestations were found most often near field borders or in places where plants were relatively sparse. Plants showing characteristic D. noxia damage typically had one or two D. noxia. Other cereal aphids, particularly Rhopalo- siphum padi (L.), Rhopalosiphum maidis (Fitch) and Sitobion avenae (Fabricius), were common. In North America and Kirghizia, D. noxia is much easier to find, and infested plants often have large colonies of aphids. Collections in North America.—It was surprising to find oviparae in the Treasure Valley and the Palouse in Idaho and Ore- gon. In all, 6 oviparae were found from two fields in the Treasure Valley and one field in Latah Co., Idaho among 785 D. noxia examined from a total of 8 fields. No males have been found to date, but the presence of oviparae opens the possibility that a small percentage of the North American D. noxia population is now holocyclic. Males, which were much less common than oviparae in the Soviet Union, may have been over- looked in our limited North American col- lections. The fact that oviparae were found in several fields in Idaho and Oregon in- creases the possibility that an occasional male could find a mate. The unusually harsh winter of 1988/9 in the Pacific Northwest USA could have provided a heavy selective advantage to holocyclic populations which resulted in their reaching detectable levels this year. Another possible explanation for the presence of oviparae follows from Black- man (1974). He has reported clones of My- zus persicae (Sulzer) which he terms andro- cyclic because they produce occasional males but no oviparae. Similarly, it is possible that North American D. noxia are gynocyclic, occasionally producing oviparae but not males. Preliminary laboratory experiments. — North American D. noxia subjected to a photoperiod of 6:18 (L:D) at 10°C for 10 weeks produced no sexual morphs. Syrian, Jordanian, French, Turkish and Kirghizian D. noxia produced no sexual morphs after three months at a photoperiod of 8:16 (L: D) at 20°C, but a Moldavian population kept VOLUME 92, NUMBER 3 under the same conditions produced ovipa- rae, males and eggs. A Moldavian popula- tion collected in August, 1989, and propa- gated under natural autumn conditions produced abundant sexuales and eggs by 23 October. Surprisingly, a Kirghizian popu- lation maintained for two months at 16°C and a photoperiod of 14:10 (L:D) at the European Parasite Laboratory produced sexuales and eggs. More research is needed on mechanisms for triggering production of sexuales in D. noxia. We thank Joyce Sorrell (University of Idaho) and Eva Rey (European Parasite Laboratory) for technical assistance and Thomas Mowry and James B. Johnson for reviewing the manuscript. We thank USDA- APHIS, USDA-ARS, the Idaho Wheat Commission and the University of Idaho for funding. This is University of Idaho Ag- ricultural Experiment Station Scientific Pa- per Number 9073. LITERATURE CITED Blackman, R. L. 1974. Live cycle variation of Myzus persicae (Sulz.) (Hom., Aphididae) in different parts of the world, in relation to genotype and environ- ment. Bulletin of Entomological Research 63: 595— 607. 547 Blackman, R. L. and V. F. Eastop. 1984. Aphids on the World’s Crops. Chichester, New York, Bris- bane, London, Toronto and Singapore: John Wi- ley & Sons. 466 pp. Grossheim, N. A. 1914. The barley aphid Brachy- colus noxius Mordvilko. Memoirs of the Natural History Museum of the Zemstvo of the Govern- ment of Taurida, Simferopol II, pp. 35-78. Hewitt, P. H., G. J. J. van Niekerk, M. C. Walters, C. F. Kriel, and A. Fouché. 1984. Aspects of the ecology of the Russian wheat aphid, Diuraphis noxia, in the Bloemfontein district. I. The colo- nization and infestation of sown wheat, identifi- cation of summer hosts and cause of infestation symptoms, pp. 3-13. Jn Walters, M. C., ed., Pro- gress in Russian wheat aphid (Diuraphis noxia Mordw.) research in the Republic of South Africa, Republic of South Africa Department of Agricul- ture Technical Communication No. 191. Jones, J. W., J. R. Byers, R. A. Butts, and J. L. Harris. 1989. Anew pest in Canada: Russian wheat aphid Diuraphis noxia (Mordvilko) (Homoptera: Aphid- idae). Canadian Entomologist 21: 623-624. Stoetzel, M. B. 1987. Information on and identifi- cation of Diuraphis noxia (Homoptera: Aphididae) and other aphid species colonizing leaves of wheat and barley in the United States. Journal of Eco- nomic Entomology 80: 696-704. Walters, M. C. 1984. Introduction, p. 2. /n Walters, M. C., ed., Progress in Russian wheat aphid (Di- uraphis noxia Mordw.) research in the Republic of South Africa, Republic of South Africa De- partment of Agriculture Technical Communica- tion No. 191. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 548-551 SCALE-LIKE STRUCTURES ON THE TIBIA OF THE PARASITIC WASPS, TRICHOGRAMMA SPP. (HYMENOPTERA: TRICHOGRAMMATIDAE) Akey C. F. HUNG Beneficial Insects Laboratory, USDA-ARS, Beltsville, Maryland 20705. Abstract. —Nine scale-like structures were found at the distal end of the hind tibia in both sexes of Trichogramma. No pore can be detected in these structures and they are so thin that they appear transparent. Behavioral observations indicate that they are probably used as brush to transfer a secretion from the abdomen to the wings to keep them from drying and thus increases the aerodynamic function of the wings, or it might be that some unknown behavioral semiochemical is transferred to the wing. Key Words: A scale-like structure in the parasitic wasps, 7richogramma spp., was first re- ported by Hung at the First International Symposium on 7Jrichogramma and Other Egg Parasites in 1981 (Hung 1982). This unique structure was subsequently found in other Trichogramma by Cals and Cals-Us- ciati (1987) and Schmidt and Smith (1987). However, none of these papers gave a de- tailed description. Since this structure has never been observed in any other group of insects, further description and discussion are given here. MATERIALS AND METHODS Seven species of Trichogramma were used in this study, namely 7. exiguum Pinto and Platner, 7. maltbyi Nagaraja and Nagar- katti, 7. minutum Riley, T. nubilale Ertle and Davis, 7. parkeri Nagakatti, 7. pre- tiosum Riley, and 7. stampae Vincent. He- liothis virescens (F.) eggs killed by exposure to ca. 30 krad of gamma radiation were used as the host in rearing the cultures. All cul- tures were maintained individually in 10- dram plastic vials at 27°C, 70-80% relative humidity. SEM, morphology, behavior, aerodynamic, semiochemical Live wasps were fixed in chilled 3% glu- taraldehyde for 3 h at room temperature and dehydrated through 100% ethanol. They were then critical point dried, mounted on stubs with silver paint and coated with gold/ palladium alloy. The specimens were stud- ied with both Hitachi S-430 and Hitachi HHS-2R scanning electron microscopes at an accelerating voltage of 15 and 20 kV. Specimens freshly killed with CO, can also be mounted directly on stubs with TV tube coat and studied at 15 kV without gold/ palladium coating for up to one hour before they collapse. Behavioral observation was carried out under WILD MSD stereomicro- scope. RESULTS AND DISCUSSION Nine scale-like structures were found in both sexes of all seven species studied. They are located on the inner surface at the distal end of hind tibia (Fig. 1). Six of them form a half-ring around the tarsal socket (Fig. 2). The 7th scale is located about 5 micra above the ring, in line with the tibial spur (sp). The remaining two are on the same line between scales 6 and 7. Scale 9 is about 15 micra VOLUME 92, NUMBER 3 above the ring with 8 about half-way in be- tween. Each scale is socketed. Their dimen- sions are 10-19 micra in length, 2.5—7.6 mi- cra in width and 0.16—-0.5 micra in thickness, with #1 the largest and #9 the smallest in size. Each scale is corrugated on both sides (Fig. 3) and keeled underneath (Fig. 2). No pore can be detected even at 20K magnifi- cation. Each scale is so thin that it can be seen through when two scales are partially overlapped (Fig. 4). The thinness of this structure has caused problems in studying it under SEM, because the tip can readily curve up under the electron beam even at 15 kV with metal-coated specimens. Rosen and DeBach (1976) reported the presence of strigil on fore leg and saltatorial mid-tibial spur in Aphytis chilensis Howard. However, they did not mention any scale- like structure. Despite the size of these scales in relation to the tarsal segment (see Fig. 1), they cannot be clearly detected with the phase contrast microscope in slide prepa- rations even when mounted in glycerine or distilled water. Under both compound and stereo microscopes, only what appear to be setae at the apex of the tibia can be seen to have an arrangement very similar to that of these scale-like structures. Under stereo- microscope, if the position of the leg or the light source is manipulated at various an- gles, a thin membrane can be detected around these “‘setae.”” Therefore, it is ap- parent that these structures are so thin and transparent that they cannot be discerned under light microscopes and only the mid- dle ridges can be seen which have the ap- pearance of setae. Cals and Cals-Usciati (1987) briefly de- scribed these structures in 7. maidis Pin- tureau and Voegele. They also reported the occurrence of “similar” structures on the pretarsus which they called “scraping se- tae.” However, these “‘linear serie (sic) of scraping setae” are not “‘petal-like” as orig- inally reported by Hung (1982). Schmidt and Smith (1987) also found the same struc- ture in 7. minutum in their SEM study. Ac- 549 cording to them, there are eight flattened and grooved modified hairs that form a wing comb at the distal rim of the metatibia; the outer surface of each hair is marked with four to five sculpted ridges and the inner surface is smooth (Schmidt and Smith 1987). However, as pointed out by Hung (1982), there are nine such structures (see Fig. 2). Furthermore, each scale is corrugated on both sides with more than 10 ridges and keeled underneath (Figs. 2, 3). Whether these differences reflect variations not observed previously cannot be confirmed at this stage. The lack of any pores rules out the pos- sibility that they are chemoreceptors. It is possible that the pores may be very small and not visible at the resolution and mag- nifications used in this study. However, the use of higher magnification will certainly be very difficult, if not impossible, because these structures can readily be deformed un- der the electron beam. They could be me- chanoreceptors which are stimulated when the legs touch the abdomen or ovipositor. However, it is hard to understand why such mechanoreceptors need to be shaped like scales. Many species of insects communicate with each other by tapping their antenna or feet against the bodies of other insects. Radia- tion from thin layers of molecules coated in insect bodies can be modulated by this tap- ping (Callahan 1977). The thinness and cor- rugations of these scale-like structures on the hind leg might be used for the imped- ance match for some incoming far IR ra- diation, possibly from some host oscillating molecule (P. S. Callahan, per. comm.). The function of this structure can only be speculated, based on my observation of the behavior of this wasp. Both male and female Trichogramma frequently rub the legs against the abdomen and the wings. It is this particular part of the leg where the scales are located that is used in this abdominal stroking and wing brushing. Although they also rub the legs against each other, only the tarsal segments are involved. It is, therefore, 550 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. The scale-like structure in Trichogramma spp. 1. T. exiguum female, hind tibia (TB) and tarsus (TS) with the scales. 2. 7. stampae female, distal end of hind tibia (pointing upward) showing nine scales (SC) and the spur (SP). 3. 7. maltbyi female, the corrugated surface of the scale. 4. T. nubilale female, two overlapped scales showing the thin membrane. Bars = 5 microns. VOLUME 92, NUMBER 3 conceivable that these scale-like structures are used to transfer some kind of secretion from the abdomen to the wings. This is sup- ported by the oily appearance of the wings. The brushing of wings with abdominal se- cretion might be used to keep the wings from drying and thus increase the aerodynamic function of the wings, or it might be that some unknown behavioral semiochemical is transferred to the wing. ACKNOWLEDGMENTS I am grateful to Norita Chaney for her assistance with SEM work, to David L. Vin- cent for maintaining the Trichogramma cultures, and to Philip S. Callahan, Ronald S. Petralia and John D. Pinto for reviewing the manuscript. LITERATURE CITED Callahan, P.S. 1977. Tapping modulation of the far infrared (17-um region) emission from the cab- bage looper moth pheromone (sex scent). Appl. Optics 16: 3098-3102. Cals, P. and J. Cals-Usciati. 1987. Repartition me- tamerique des structures cuticularies d’un insecte evolue: Trichogramma maidis (Hymenoptera: Chalcidoidea). C. R. Acad. Sci. Paris (Serie 3) 305: 129-133. Hung, A.C. F. 1982. Petal-like structure in Tricho- gramma (Abstract). Les Colloques de PTINRA 9: MIF, Rosen, D. and P. DeBach. 1976. Biosystematic stud- ies on the species of Aphytis (Hymenoptera: Aphe- linidae). Mushi 49: 1-17. Schmidt, J. M. and J. J. B. Smith. 1987. The external sensory morphology of the legs and hairplate sys- tem of female Trichogramma minutum Riley (Hy- menoptera: Trichogrammatidae). Proc. Roy. Soc. Lond. (B) 232: 323-366. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 552-560 ALTITUDINAL PATTERNS IN SPECIES RICHNESS OF NEOTROPICAL TREEHOPPERS (HOMOPTERA: MEMBRACIDAE): THE ROLE OF ANTS K. L. OLMSTEAD! AND T. K. Woop? ‘Department of Entomology, University of Maryland, College Park, Maryland 20742; *Department of Entomology and Applied Ecology, University of Delaware, Newark, Del- aware 19711. Abstract.—Treehoppers are sap-feeding insects that vary widely in degrees of both sociality and ant mutualism. Based on these life histories, treehoppers may be classified as (1) species that are ant mutualists and that aggregate as individuals, (2) species exhibiting parental care that are not ant mutualists, and (3) solitary species that rarely interact with ant mutualists. We predicted the availability of ants should influence the distribution of treehopper species that depend upon ants for protection. Because ant abundance has been shown to decline with increasing altitude in tropical regions, we examined the elevational distribution of treehopper species in Colombia that are obligate ant mutualists and those treehopper species that are not. The proportion of treehopper species that are dependent upon ants for defense declined with increasing altitude. Those species having parental care, that do not rely on ants for defense, were more common at higher elevations. Solitary treehoppers, species that only occasionally interact with ants, did not show a changing relationship with altitude. Thus, mutualistic ants are not only important in the evolution of treehopper life histories but also appear to be important in determining the geographic distribution of treehoppers. Key Words: membracids, mutualism, altitude, Colombia Sociality in treehoppers (Homoptera: ants (Wood 1977, Bristow 1983, Olmstead Membracidae) ranges from solitary species to those with highly developed parental care. Additionally, many species rely on ant mu- tualists for protection from natural enemies (Wood 1984). In return, treehoppers pro- vide ants with a source of carbohydrates, free amino acids and amides, and water in the form of honeydew (Way 1963). Thus, treehopper sociality and ant mutualism may allow alternative defenses against predators (Wood 1982b). For example, aggregations of nymphs and young adults of many species are tended by ants (Wood 1984). In some of these species, parent females also guard eggs and early instars, but their survival is still dependent on the defense provided by 1984). The size of treehopper aggregations and the volume of honeydew produced are important factors contributing to the con- stancy of ant attendance (McEvoy 1979, Fritz 1982, Wood 1982a, Cushman and Whitham 1989). Other treehopper species also aggregate, but do not interact with ant mutualists. In these presocial species (species with paren- tal care) parent females actively guard eggs and nymphs protecting them from preda- tors (Hinton 1976, 1977, Wood 1976, 1982b, Eberhard 1986). The benefit of ag- gregation in these species is the effective guarding of offspring by parent females rath- er than the attraction of ants (Wood 1976). VOLUME 92, NUMBER 3 Alternatively, some treehopper species are solitary throughout their life cycle, are rare, and may incur lower levels of predation simply by virtue of their crypsis. Because a relatively small volume of honeydew is pro- duced by solitary treehoppers, ant-treehop- per mutualisms are relatively uncommon in these species (Wood 1984). Based on their level of sociality and in- teractions with ant mutualists, treehoppers may be classified as (1) species that are ant mutualists and that form aggregations as in- dividuals, (2) species with parental care that are not ant mutualists, or (3) solitary species that rarely interact with ant mutualists. In the tropics, ant mutualisms decline with increasing altitude in myrmecophilous an- imals (Wood 1984) and plants (Bentley 1977a, b, Koptur 1985). This pattern re- flects the decline in ant abundance along an increasing elevational gradient (Janzen 1973, Janzen et al. 1976, Bentley 1977a). The cool air temperatures and high soil moisture of tropical montane regions preclude ants from exploiting these habitats (Bentley 1977a). Koptur (1985) and Bentley (1977b) have shown that nectary plants in areas of low ant activity have alternative defenses against herbivores. Thus, the plasticity of the de- fensive repertoire of these plants (Jnga and Bixa) permits them to grow in areas where ant activity is low. In contrast, the defensive mechanisms of treehoppers are not labile within species. Consequently, treehopper species that rely solely upon ants for defense are relatively undefended in the absence of ants. Furthermore, the increased conspic- uousness of individuals in aggregations formed by ant-dependent species elevates their risk of predation compared to species that do not rely upon ants. Non-attended species reduce their risk of predation in oth- er ways. Specifically, aggregations formed by presocial species are protected by parent females while solitary species are cryptic. Given the effects of altitude on ant abun- dance and the importance of ants to some treehopper species, we predicted a decline 553 with increasing altitude in the number of tropical treehopper species that depend upon ants for protection. We also predicted that species not dependent on ants for defense should be more common at higher altitudes in the tropics where ants are rare. We used treehoppers to test our predictions because they have a wide geographic distribution and they exhibit diverse life history types (Wood 1982b, 1984). We chose to restrict our study to the treehoppers of Colombia because it has an altitudinal range of 5000 m and membracid taxonomists have made extensive collections there. METHODS We obtained locality and altitude records for all treehopper species from the literature (Richter 1940, 1941a, b, 1942a, b, c, 1943, 1945, 1955, Striimpel 1972, 1973, Striim- pel and Striimpel 1975, 1978) and used gaz- etteers and relief maps to determine the al- titude of those localities for which authors did not provide this information. We di- vided the altitudinal gradient into 13 classes of 250 m increments, from sea level to 3000 m and above. We found no records of tree- hoppers collected above 4200 m in Colom- bia. One problem inherent in analyzing col- lection data from published works is the accuracy of locality records. For example, workers may designate the nearest large city as the collection site rather than a more ac- curate locality. Because we used published data, such errors may exist in our data set. We followed Metcalfand Wade (1965) for species synonymies and Deitz (1975, 1983, 1985) for classification at the subfamily and tribal levels. We assumed an equal error rate in species identification across taxa relative to life history type. We used Wood’s (1976, 1977, 1984) studies of membracids as well as those by Eberhard (1986), Ekkens (1972), Fritz (1982), Haviland (1925), and Hinton (1976, 1977) to determine sociality and ant mutualism for 330 (86%) of the 384 tree- hopper species recorded from Colombia. 554 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. The number of treehopper species within each 250 m altitudinal class are given for Colombian treehoppers exhibiting one of three life history types. The number of species for which life history types was not available or could not be inferred is also given (see text for explanation of life history types). No. of Species No. of Ant No. of Presocial with Unknown Dependent Species (No Ant No. of Solitary Life History Altitude Range (in Meters) Midpoint Species Mutualism) Species Types Total 0-249 125 40 1 21 2 64 250-499 375 61 2 40 3 106 500-749 625 72 l 35 7 115 750-999 875 51 2 25 6 84 1000-1249 1125 32 3 14 6 55 1250-1499 1375 32 6 11 5 54 1500-1749 1625 5 7 8 4 24 1750-1999 1875 17 11 8 7 43 2000-2249 2125 10 3 8 1 22 2250-2499 2375 0 3 4 1 8 2500-2749 2625 3 i 4 Y) 23 2750-2999 2875 8 22 14 18 62 3000+ 3125 1 3 7 4 15 Total Number of Species 156 45 129 54 384 When sociality or ant mutualism informa- tion was not available for a particular species, we designated the life history type on the basis of congeners for which these data were available. In the Membracidae, life history patterns are often invariable within tribes, and with few exceptions, are consistent at the generic level (Wood, per- sonal observation). We categorized each species as one of three types (1) species that aggregate as individuals and that are ant mutualists, (2) presocial species that are not ant mutualists, and (3) solitary species. Ap- pendix A is a list of the treehopper genera and includes data on level of sociality and the presence of ant mutualism. To control for unequal sampling effort throughout Colombia, we evaluated species richness as the proportion of species with a particular life history type relative to all species with known life history types that occur at that elevational class. For example, in our data set 21 solitary species occurred between sea-level and 250 m. Because 62 treehopper species occur in Colombia be- tween 0 and 250 m, solitary species repre- sent 33.87% of the treehopper species in this elevational class. We assumed that although some zones may be less well sampled than others, the proportions of species approxi- mate the relative richness of species with different life history types. Proportional val- ues were arcsine transformed prior to anal- ysis. We employed polynomial regression models (SAS Institute 1986) to describe the relationship between altitude and the pro- portion of treehopper species of each life history type. These models are appropriate because there was no reason to assume the relationship between species richness and altitude was linear. This approach also made it possible to describe the form of the re- lationship. We used sequential (Type I) sums of squares to determine the order of the polynomial regression that was appropriate (Freund et al. 1986). Initially, we used fourth degree polynomials and retained terms sig- nificant at P < .05 in the models. We ex- amined the Studentized residuals to deter- mine if our data met the assumptions of the models. VOLUME 92, NUMBER 3 RESULTS AND DISCUSSION We were able to assign life history clas- sification to 330 species of treehoppers in our data set. Of these species, 156 (47.27%) were dependent upon ant mutualists, 45 (13.64%) were presocial species that do not interact with ants, and 129 (39.09%) were solitary. The species richness of treehoppers with each of these three life history types across the altitudinal gradient is given in Table 1. In Colombia, the proportion of treehop- per species that depend upon ants for pro- tection declines with increasing altitude (Figure la). A linear model best described the relationship (y = 61.99 — 0.014x!, R? = .62, P < .01). Because ants are less common at higher elevations (Janzen 1973, Janzen et al. 1976, Bentley 1977a, b, Koptur 1985), treehoppers in these zones may be at a higher risk of predation than those occurring at lower elevations where ants are more abun- dant. Thus, our data supported our hypoth- esis that the altitudinal distribution of tree- hoppers that depend upon ants for protection reflects the availability of ant mutualists. We found a significant positive relation- ship between altitude and the proportion of presocial treehopper species that are not ant mutualists. A linear model best described the relationship (Fig. 1b; y = 2.71 + 0.135x', R2 = .77, P < .01). Because protection of offspring is provided by parent females rath- er than by ant mutualists, these species are less likely to be restricted to areas where ants are abundant. For this reason, species with parental care are overrepresented at higher elevations where ants, and consequently ant- dependent treehoppers, are rare. Because ant-treehopper mutualisms are not common but do occur in solitary tree- hoppers (Wood 1984), these species should not be restricted to areas in which ant mu- tualists are common. Our data supported this hypothesis since the proportion of sol- itary species is nearly equal over the ele- vational gradient (Fig. 1c) and the propor- 555 i) 1000 2000 3000 4000 Proportion of Species (%) 1) 1000 2000 3000 4000 0 1000 2000 3000 4000 Altitude (in meters) Fig. 1. The relationship between altitude and the proportion of Colombian treehopper species of three life history types: 1a) aggregating ant-dependent species, 1b) presocial species that do not interact with ants, and lc) solitary species. Proportional values were arcsine transformed. Solid lines represent significant regres- sions (see text for regression equations and explanation of life history type). tion of solitary treehopper species was not statistically related to elevation in any of the models tested. Solitary species represent on average 36.77 + 7.23% (arcsine trans- formed mean +1 SD) of the membracid species at any elevation. We have focused here upon the relation- 556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ships between ants, altitude, and treehopper life histories. Admittedly, ant mutualism is not the only selective factor varying across the environmental gradient. Differential plant productivity, seasonality, and a num- ber of other factors may also affect the el- evational distribution of phytophagous in- sects (Begon et al. 1986, Descimon 1986). We assume, however, that the differing se- lective pressures resulting from the clinal variation in these factors are evenly im- posed on species with all three life history types. It appears that the decline with in- creasing altitude in the relative richness of treehopper species that depend upon ants for protection is due in large part to the corresponding decline in the abundance of mutualistic ants. ACKNOWLEDGMENTS We thank E. Russek-Cohen, R. Denno, L. Deitz, J. Davidson, L. Hanks, G. Rod- erick, and C. von Dohlen for their helpful comments on earlier drafts of this report. The computer time for this project was pro- vided in full by the Computer Science Cen- ter at the University of Maryland. This re- port is Scientific Article No. A-4783, Contribution No. 7803 of the Maryland Ag- ricultural Experiment Station, Department of Entomology. This report is also pub- lished as miscellaneous paper No. 1223 of the Delaware Agricultural Experiment Sta- tion, Contribution No. 591 of the Depart- ment of Entomology and Applied Ecology and Contribution No. 132 of the Ecology Program, School of Life and Health Sci- ences, University of Delaware, Newark, DE. LITERATURE CITED Begon, M., J. L. Harper, and C. R. Townsend. 1986. Ecology: Individuals, Populations, and Commu- nities. Sinauer Associates Inc., Sunderland, Mas- sachusetts. 876 pp. Bentley, B. L. 1977a. Extrafloral nectaries and pro- tection by pugnacious bodyguards. Ann. Rev. Ecol. Sys. 8: 407-427. 1977b. The protective function of ants vis- iting the extrafloral nectaries of Bixa orellana (Bix- aceae). J. Ecol. 65: 27-38. Bristow, C. M. 1983. Treehoppers transfer parental care to ants: A new benefit of mutualism. Science 220: 532-533. Cushman, J. H. and T. G. Whitham. 1989. Condi- tional mutualism in a membracid-ant association: Temporal, age-specific, and density-dependent ef- fects. Ecology 70: 1040-1047. Deitz, L. L. 1975. Classification of the higher cate- gories of the New World treehoppers (Homoptera: Membracidae). N.C. Agric. Exp. Sta. Tech. Bull. 225, Raleigh, N.C. 1983. Name changes in the Membracidae (Homoptera). Proc. Entomol. Soc. Wash. 85(4): 856-857. 1985. Placement of the genera Abelus Stal and Hemicentrus Melichar in the subfamily Cen- trotinae (Homoptera: Membracidae). Proc. Ento- mol. Soc. Wash. 87(1): 161-170. Descimon, H. 1986. Origins of lepidopteran faunas of the high tropical Andes, pp. 500-532. Jn Vuil- leumier, F. and M. Monasterio, eds., High Alti- tude Tropical Biology. Oxford University Press, New York. Eberhard, W. G. 1986. Possible mutualism between females of the subsocial membracid Polyg/ypta dispar (Homoptera). Oecologia (Berlin) 19: 447- 453. Ekkens, D. 1972. Peruvian treehopper behavior (Ho- moptera: Membracidae). Entomol. News 83: 257- 2d Freund, R. J., R. C. Littell, and P. C. Spector. 1986. SAS system for linear models, 1986 edition. SAS Institute, Cary, N.C. Fritz, R.S. 1982. An ant-treehopper mutualism: Ef- fects of Formica subsericea on the survival of Van- duzea arquata. Ecol. Entomol. 7: 267-276. Haviland, M. D. 1925. The Membracidae of Kar- tabo, Bartica District, British Guiana. Zoologica (New York) 6: 229-290. Hinton, H. E. 1976. Maternal care in the Membraci- dae. Proc. R. Entomol. Soc. Lond. (C) 40: 33. 1977. Subsocial behaviour and biology of some Mexican membracid bugs. Ecol. Entomol. 2: 61-79. Janzen, D.H. 1973. Sweep samples of tropical foliage insects: Effect of seasons, vegetation types, ele- vation, time of day and insularity. Ecology 54: 687-708. Janzen, D. H., M. Ataroff, M. Farinas, S. Reyes, N. Rincon, A. Soler, P. Soriano, and M. Vera. 1976. Changes in the arthropod community along an elevational transect in the Venezuelan Andes. Bio- tropica 8: 193-203. Koptur, S. 1985. Alternative defenses against her- VOLUME 92, NUMBER 3 bivores in /nga (Fabaceae: Mimosoideae) over an elevational gradient. Ecology 66: 1639-1650. McEvoy, P. B. 1979. Advantages and disadvantages of group living in treehoppers (Homoptera: Mem- bracidae). Misc. Publ. Entomol. Soc. Amer. 11: 1-13. Metcalf, Z. P. and V. Wade. 1965. General catalogue of the Homoptera. Membracoidea I and II. N.C. State University, Raleigh, N.C. Olmstead, K. L. 1984. Maternal behavior to establish ant-membracid associations. MS Thesis, Univer- sity of Delaware, Newark, DE. Richter, L. 1940. Catalogo de los Membracidae de Colombia. Rev. Acad. Colombia 3: 462-463. 1941a. Contribuci6n al conocimiento de los Membracidae de Colombia. Caldasia 2: 67-74. . 1941b. Catalogo de los Membracidae de Co- lombia. Rev. Acad. Colombia 4: 83-85. 1942a. Contribucion al conocimiento de los Membracidae de Colombia. II. Caldasia 3: 41-48. 1942b. Contribucion al conocimiento de los Membracidae de Colombia. III. Caldasia 5: 41- 49. . 1942c. Catalogo de los membracidos de Co- lombia. Rev. Acad. Colombia 4: 405-409. 1943. Contribucion al conocimiento de los Membracidae de Colombia. IV. Caldasia 6: 81- 12 1945. Membracidae Colombianae. Rev. Acad. Colombia 6: 339-354. 1955. Entomologia. Membracidae Colom- bianae. Caldasia 6: 269-380. SAS Institute. 1986. SAS system for regression. SAS Institute, Cary, N.C. Striimpel, H. 1972. Die Membraciden-Fauna Ko- lumbiens. |. Die Gattung Notocera Amyot and 557 Serville, 1943. Mitt. Hamburg Zool. Mus. Inst. 69: 33-55. 1973. Die Membraciden-Fauna Kolum- biens. 2. Die Gattung Spongophorus Fairmaire, 1846. Entomol. Mitt. Zool. Mus. Hamburg 4: 327- 350. Strimpel, H. and R. Striimpel. 1975. Die Membra- ciden-Fauna Kolumbiens. 3. Die Gattung Leios- cyta Fowler, 1894. Mitt. Hamburg Zool. Mus. Inst. 72: 177-200. 1978. Die Membraciden-Fauna Kolum- biens. 4. Die Gattung Tritropidia Stal, 1869. Ento- mol. Mitt. Zool. Mus. Hamburg 6: 133-149. Way, M. J. 1963. Mutualism between ants and hon- eydew-producing Homoptera. Ann. Rev. Ento- mol. 8: 307-344. Wood, T. K. 1976. Alarm behavior of brooding fe- male Umbonia crassicornis (Homoptera: Mem- bracidae). Ann. Entomol. Soc. Amer. 69: 340-344. 1977. Role of parent females and attendant ants in the maturation of the treehopper, Entylia bactriana (Homoptera: Membracidae). Sociobiol- ogy 2: 257-272. 1982a. Ant-attended nymphal aggregations in the Enchenopa binotata complex (Homoptera: Membracidae). Ann. Entomol. Soc. Amer. 75: 649- 653. 1982b. Selective factors associated with the evolution of membracid sociality, pp. 175-179. In Breed, M. D., C. D. Michener, and H. E. Evans, eds., The Biology of Social Insects. Westview Press, Boulder, CO. 1984. Life history patterns of tropical mem- bracids (Homoptera: Membracidae). Sociobiology 8: 299-344. 558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Appendix A. A list of the treehopper genera reported in Colombia including the number of species, life history type, ant mutualism, and references. Life histories were categorized as one of three types: aggregative/ ant-dependent, presocial, or solitary. Genus No. of Species Life History Type (Reference)* Ant Mutualism (Reference)* Subfamily Centrotinae Tribe Abelini Ischnocentrus 3 Aggregative (8) Yes (8) Subfamily Membracinae Tribe Aconophorini Aconophora 14 Presocial (3, 5, 8) Yes (8) Guayaquila 1 Presocial (8) No (8) Tribe Hoplophorionini Potnia 2 Presocial (8) No (8) Ochropepla 2 Presocial (8) No (8) Hoplophorion 10 Presocial (8) No (8) Alchisme 12 Presocial (8) No (8) Umbonia 4 Presocial (1, 5, 6, 8) No (8) Tribe Membracini Bolbonota 7 Aggregative (1, 8) Yes (8) Tritropidia 5 Aggregative (3) Yes (9) Erechtia 5 Presocial (8) Yes (8) Tylopelta 3 Aggregative (8) Yes (8) Leioscyta 14 Aggregative (8) Yes (8) Campylenchia 72 Aggregative (8) Yes (8) Enchophyllum 6 Aggregative (8) Yes (8) Enchenopa 8 Agegregative (8) Yes (8) Membracis 28 Aggregative (5, 8) Yes (8) Tribe Hypsoprorini Notocera 11 Aggregative (8) Unknown Philya 4 Solitary (8) No (8) Hypsoprora 8 Unknown Unknown Sphongophorus 15 Solitary (8) Unknown Subfamily Darninae Darnoides 4 Unknown Unknown Hypheodana 1 Unknown Unknown Tribe Cymbomorphini Cymbomorpha 1 Solitary (8) No (8) Tribe Darnini Darnis 3 Solitary (8) No (8) Hebetica 2 Unknown Unknown Stictopelta 2 Solitary (8) No (8) Alcmeone 1 Solitary (8) No (8) Tribe Hyphinoini Bubalopa 2 Unknown Unknown Hyphinoe 2 Solitary (5, 8) No (8) Tomogonia 2 Unknown Unknown Tribe Hemikypthini Proterpia 1 Unknown Unknown Atypa 1 Solitary (8) No (8) Subfamily Smiliinae Tribe Acutalini Acutalis 4 Solitary (8) No (8) VOLUME 92, NUMBER 3 Appendix A. Continued. Genus Euritea Thrasymedes Tribe Micrutalini Micrutalis Tribe Ceresini Antonae Centrogonia Penichrophorus Tlithucia Melusinella Ceresa Stictolobus Vestistilus Cyphonia Poppea Tribe Amastrini Vanduzea Harmonides Tynelia Lallemandia Amastris Tribe Smiluni Telamona Antianthe Tribe Tragopini Horiola Tragopa Stilbophora Chelyoidea Tropidolomia Tribe Polyglyptini Eucatoriana Heranice Adippe Dioclophara Ennya Hille Polyglyptodes Maturnaria Metheisa Polyrhyssa Entylia Polyglypta Aphetea Phormophora Subfamily Stegaspidinae Tribe Stegaspidini Bocydium Stylocentrus Oeda Lycoderes No. of Species tN Ne} Ne $s Ww oNnNnno-— Nw hs Ne wre Nm how h Neue = WN KK DAN PONN ~~) on wN Life History Type (Reference)* Unknown Solitary (8) Solitary (8) Unknown Solitary (9) Solitary (9) Unknown Unknown Solitary (9) Solitary (9) Solitary (8) Solitary (8) Aggregative (8) Aggregative (8) Aggregative (8) Aggregative (3) Unknown Solitary (8) Solitary (9) Presocial (5, 8) Presocial (3, 8) Solitary (8) Aggregative (9) Aggregative (9) Aggregative (9) Unknown Unknown Presocial (8) Unknown Presocial (8) Presocial (3) Presocial (8) Unknown Presocial (8) Unknown Presocial (5, 7, 8) Presocial (la, 5, 8) Presocial (3, 8) Unknown Solitary (8) Solitary (8) Solitary (8) Solitary (8) Ant Mutualism (Reference)* Unknown No (8) Yes (8) Unknown Unknown Unknown Unknown Unknown No (9) No (9) No (8) No (8) Yes (8) Yes (2, 8) Yes (8) Unknown Unknown Yes (1, 8) Unknown Yes (8) Yes (8) Yes (8) Unknown Unknown Yes (9) Unknown Unknown Yes (8) Unknown No (8) Unknown No (8) Unknown Yes (8) Unknown Yes (7, 8) No (8) Yes (1, 8) Unknown No (8) No (8) No (8) No (8) 560 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | Appendix A. Continued. Genus No. of Species Life History Type (Reference)* Ant Mutualism (Reference)* Stegaspis 5 Aggregative (9) Unknown Euwalkeria 1 Unknown Unknown Subfamily Heteronotinae Tribe Heteronotini Nassunia 2 Aggregative (8) Yes (8) Anchistrotus 2 Aggregative (9) Unknown Heteronotus 3 Aggregative (1) Yes (1, 8) Smiliorachis ! Unknown Unknown Rhexia 11 Aggregative (9) Yes Total Number of Genera = 84 Total Number of Species = 384 * la) Eberhard 1986, 1) Ekkens 1972, 2) Fritz 1982, 3) Haviland 1925, 4) Hinton 1976, 5) Hinton 1977, 6) Wood 1975, 7) Wood 1977, 8) Wood 1984, 9) Wood, personal observation. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 561-570 A REVISION OF ZACREMNOPS SHARKEY AND WHARTON (HYMENOPTERA: BRACONIDAE: AGATHIDINAE) MICHAEL J. SHARKEY Biosystematics Research Centre, Agriculture Canada, Research Branch, Central Ex- perimental Farm, Ottawa, Ontario K1A 0C6, Canada. Abstract.—The genus Zacremnops Sharkey and Wharton (Hymenoptera: Braconidae: Agathidinae) is revised. Four species are recognized of which two, Z. ekchuah and Z. coatlicue are new to science. The nominal species Z. oranensis de Fernandez and Z. petiolatus (Szépligeti) are synonymized with Z. chiriquensis (Cameron). A diagnostic key to species is presented and phylogenetic relationships among the species are discussed. Key Words: INTRODUCTION AND HISTORICAL REVIEW The genus Zacremnops was proposed by Sharkey and Wharton (1985) to include two species, Z. petiolatus (Szépligeti) and Z. al- bitarsus (Cresson), previously placed in Megagathis Kriechbaumer. De Fernandez (1987) revised the species of Argentina and Bolivia and described a new species Z. oranensis. Studies of additional material since the publication of these works has in- dicated two new synonyms, two new species and apparently two morphotypes in one previously described species. These findings are presented in this paper. PHYLOGENY The relationships of Zacremnops within the Agathidinae were discussed by Sharkey and Wharton (1985) and nothing will be added to the argumentation presented there. The phylogenetic relationships among the species of Zacremnops are problematical. The species vary in only a few characters and all of these occur in both states in the outgroups, Cremnops and Labagathis. The sister group of Zacremnops, Labagathis, has a color pattern similar to that of Z. chiri- Hymenoptera, Zacremnops, revision quensis and Z. coatlicue. Indeed, this color pattern is widespread throughout the sec- ondary outgroup, Cremnops, and therefore appears to be the plesiomorphic condition within Zacremnops. The predominately black coloration, which is very rare in Za- cremnops and absent in the monotypic sis- ter group, is hypothesized to be a synapo- morphy diagnosing Z. cressoni and Z. ekchuah. Thus the preferred, though weakly supported, hypothesis can be summarized as follows: ((Z. chiriquensis) (Z. coatlicue) (Z. cressoni + Z. ekchuah)) or Z. cressoni and Z. ekchuah are sister species but the phylogenetic placement of the two other species remains uncertain. TEXT CONVENTIONS Abbreviations of depositories follow the conventions of Arnett and Samuelson (1986). AEIC: American Entomological Insti- tute, Gainesville, Florida, U.S.A. AMNH: American Museum of Natural History, New York, New York, USA. 562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ANSP: Academy of Natural Sciences, Philadelphia, Pennsylvania. British Museum of Natural His- tory, London, England. Canadian National Collection of Insects, Ottawa, Ontario, Cana- da. Florida State Collection of Ar- thropods, Gainesville, Florida, U.S.A. Hungarian Natural History Mu- seum, Budapest, Hungary. Fundacion e Instituto Miguel Lillo, Tucuman, Argentina. Museum of Comparative Zoolo- gy, Cambridge, Massachusetts. Muséum National d’Histoire Na- turelle, Paris, France. Department of Entcmology In- sect Collection, Texas A&M Uni- versity, College Station, Texas. Department of Entomology Col- lection, University of Georgia, Athens, Georgia. United States National Museum of Natural History, Washington, DIG UEStA® Bereich Zoologisches Museum, Berlin, German Democratic Re- public. BMNH: CNCI: FSCA: HNHM: IMLA: MCZC: MNHN: TAMU: UGCA: USNM: ZMHB: TAXONOMIC TREATMENT Zacremnops Sharkey and Wharton, 1985. Type-species.—Agathis albitarsis Cres- son (= A. cressoni Cameron, new name for albitarsis Cresson). 1865, p. 63, male, ANSP, type no. 1729.1. Type designation by Shar- key and Wharton (1985). Description. — Males and females. (From Sharkey and Wharton, 1985). Head (Figs. 1, 4). Distance between median ocellus and lateral ocellus much greater than distance between lateral ocelli; median ocellus situ- ated much lower on face than lateral ocelli; frons, between median ocellus and antennal sockets, smooth and shining, carinae lack- ing; occiput slightly excavated for reception of pronotum (Fig. |); malar space long, 0.7- 0.8 eye height; longitudinal carina be- tween antennae weak or absent; anterior tentorial pit about 2 x closer to eye than to mandibular condyle; maxillary palpus 5-segmented, second and last segments sometimes longest, often all segments sub- equal (Fig. 4); labial palpus 4-segmented, basal 3 segments subequal, apical segment may be slightly longer (Fig. 4); clypeus al- most as high as wide, height: width ratio about 0.8. Mesosoma (Figs. |, 2, 5—7).— Anterior and posterior portions of pronotum, medially, separated by deep transverse groove (i.e. subpronopes confluent) (Fig. 1); propleuron without protuberances; sternaulus well de- veloped, usually complete to epicnemial ca- rina (Fig. 7); epicnemial carina approaching pronotum near mid-height of posterior margin of pronotum; at least ventral half of metapleuron areolate-rugose; notauli deep- ly impressed, smooth; scutellum with pos- terior transverse ridge weak or absent; pro- podeum areolate (Fig. 2); propodeal spiracles oval; hind coxal cavities closed; fore tibia lacking spines apically; mid tibia with apical spines but lacking spines admedially; all tar- sal claws bifid with pectination basally (Fig. 5); hind trochanterellus lacking longitudinal carina; hind coxa large, about 2 longer than mid coxa; IRS cell of fore wing quad- rate (Fig. 6); 2RS2 vein (really a spurious vein) absent or present as a stub (Fig. 6); cells 1M and 1R1 of fore wing confluent (Fig. 6); last abscissa of Cu vein of hind wing present and well sclerotized basally; last ab- scissa of Cu vein of hind wing positioned closer to vein A than to vein M+Cu (Fig. 6); 2r-m crossvein (really a spurious vein) of hind wing weakly indicated or completely absent. Metasoma (Fig. 3).— First tergum long and narrow, 2.6—4.2 x longer than apical width; apex of first tergum more than 2 wider than base; all terga mostly smooth; ovipos- itor longer than metasoma but shorter than body length when fully exposed. VOLUME 92, NUMBER 3 563 Figs. 1-5. Color.—Wings infuscated, with several small hyaline patches (Fig. 6), lacking yel- low bands or spots; body mostly to entirely black and usually with some yellow, red or yellowish red color on legs, head, mesosoma or metasoma anteriorly. Body length.— Most specimens are large, from 9 to 13 mm; some specimens of Z. chiriquensis may be as small as 6.5 mm. Key TO ADULTS OF SPECIES OF ZACREMNOPS Ie Hind tarsus yellow ..... ...Z. cressoni 1 inditarsusiblacks 35.20ceeen.c asec cece .05 mm Zacremnops cressoni female: 1, dorsal aspect of head and pronotum; 2, propodeum; 3, dorsal aspect three basal metasomal segments; 4, lateral aspect of head; 5, tarsal claw. 2(1'). Body entirely melanic Z. ekchuah Pd Body partly reddish orange or yellow 3 Z. coatlicue Z. chiriquensis 3(2'). Mesopleuron black Mesopleuron reddish orange Zacremnops chiriquensis (Cameron), NEw CoMBINATION Agathis chiriquensis Cameron, 1887, p. 399. Cremnops petiolatus Szépligeti, 1902, p. 65. Syn. N. Megagathis? petiolata (Szépligeti), 1904, p. 122. Megagathis petiolata: Enderlein, 1920 (1918), p. 167. 564 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1mm Figs. 6-7. Zacremnops petiolatus: Sharkey and Whar- ton, 1985, p. 603. Zacremnops oranensis de Fernandez, 1987, p. 90. New SYNONYMY Diagnosis.—Males and females. Color mostly black, metapleuron and propodeum reddish orange. Zacremnops cressoni female: 6, fore and hind wing; 7, lateral aspect of mesosoma. Description.—Antenna with 39-46 fla- gellomeres; metapleuron rugose reticulate in ventral 73, smooth to partly rugose in dorsal 3; hind femur from weakly rugose punctate to smooth with scattered punc- tures; body length 6.5 to 12.5 mm; body color mostly black; mesopleuron, meta- pleuron and propodeum reddish orange, VOLUME 92, NUMBER 3 some or all of following parts sometimes reddish orange: mouthparts, fore tarsus, base of hind coxa, scutellum, metanotum and basal half of first metasomal segment; rarely mouthparts yellow or reddish orange. Distribution. — Map 1. Restricted to South America with one record from Panama. Probably widespread north of the 30th par- allel except in excessively dry or high areas. Type material.—Agathis chiriquensis (Cameron), Holotype female, PANAMA, David, (BMNH, 3.c.933) (examined). Zac- remnops oranensis de Fernandez, Holotype female. ARGENTINA, Salta, Rio Pescado, Oran, 30.1V.1968, (Porter, IMLA) (exam- ined). Cremnops petiolatus Szepligeti, Lec- totype female, BRAZIL, Amazonas, Ton- antins, (HNHM) (examined). Depositories.—The more than 400 spec- imens that I have identified are in the fol- lowing collections: AEIC, AMNH, BMNH, CNCI, FSCA, HNHM, IMLA, MCZC, USNM, ZMHB. Remarks. — This species is widespread and rather variable morphologically. De Fer- nandez (1987) based the recognition of a new species, Z. oranensis, which I consider to be a junior synonym of Z. chiriquensis, on the presence or absence of two longitu- dinal carinae on the first metasomal tergum. After examining over 400 specimens of this species I have come to the conclusion that the character is variable intraspecifically. The carinae grade from quite strong to com- pletely absent with no correlation with other characters or with geographic distribution. De Fernandez (1987) based her conclusions on ten specimens and therefore did not have the advantage of observing the variation of this character. She also used a character to separate Z. cressoni from (what I consider to be) Z. chiriquensis that does not prove to be valid when many specimens are ex- amined. This is the presence or absence of a complete median longitudinal carina sep- arating the subpronopes. Z. coatlicue is very close to Z. chiriquensis and the two may prove to be conspecific. 565 Though, at present Iam persuaded that they are separate species because of three char- acters: the difference in coloration, with Z. chiriquensis having more reddish-orange coloration; the difference in size, Z. coatli- cue generally being composed of larger spec- imens; and Z. coatlicue having a relatively more robust and wider metapleuron with the dorsal smooth area being substantially larger and mostly lying on a horizontal rath- er than vertical plane. All of these characters are somewhat variable, but taken together they indicate distinct species. Zacremnops coatlicue, NEW SPECIES Etymology.—Named after Coatlicue, a hideous Mixica chthonic goddess, with a thirst for blood. Diagnosis.—Males and females. Color black except metapleuron and propodeum reddish orange. Description. — Males and females. Anten- na with 44-46 flagellomeres; metapleuron rugose reticulate in ventral *4, smooth in dorsal *4; hind femur from weakly rugose punctate to smooth with scattered punc- tures ventrally; body length 11.3 to 15.0 mm.; body color entirely black except meta- pleuron and propodeum reddish orange, rarely (two specimens) the mesopleuron is reddish-black posteriorly. Distribution.—Map 2. Restricted to Mexican and Central American lowlands. Near the Pacific specimens are found in de- ciduous forest as far south as Guanacaste Province in Costa Rica and as far North as Sinaloa State in Mexico. Specimens col- lected near the Caribbean coast are from habitats that once were tropical rainforest, perhaps indicating that the species has a wide range of moisture tolerance. Type material.— Holotype female, COS- TA RICA, Guanacaste, Santa Rosa Park, 22.V.1978 (Janzen, AEIC). Paratypes. —Allotype, male, same data as holotype except date 8.V.1977. COSTA RICA: Guanacaste: 3 females, same data as holotype except dates 20.V.1978, 21.V.1978 566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-9. Zacremnops cressoni female: 8, ventral aspect of punctate hind femur; 9, ventral aspect of rugose hind femur. and 18.VI.1978. MEXICO: Jalisco: 1 fe- ’ male, Puerto Vallarta, 18-23.VII.1961 Zacremnops cressoni (Cameron) (Grant, CNCD). Sinaloa: 1 female, 8 mi. (13 (Figs. 1-9) km)S. Elota, 2. VII.1963, (Parkerand Stange, Agathis albitarsus Cresson, 1865, 4: 63. USNM). | male, Venodio, (Rosche, USNM). (preoccupied by 4A. albitarsus Spinola, Veracruz: 1 female, Tecolutla, 19.VI.1951 1840). (Evans, AEIC). Cremnops albitarsis: Schulz, 1906, p. 137. VOLUME 92, NUMBER 3 567 MAP 1 @ Z. chiriquensis Map. 1. Distribution of Z. chiriquensis. 568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MAP 2 @ Zekchuah @ Z. coatlicue Map 2. Megagathis albitarsis: Enderlein, 1920 (1918), p. 167. Agathis cressoni Cameron, 1887 (new name for A. albitarsus), p. 398. Cremnops cressoni: Ashmead, 1895 (1894), p: 123: Zacremnops cressoni: Sharkey and Whar- ton, 1985, p. 599. Diagnosis.—Males and females. Hind tarsus yellow, remainder of body usually black, often with yellow markings, never with reddish orange markings. Description. — Males and females. Anten- na with 42-49 flagellomeres; metapleuron rugose reticulate in ventral 74, smooth to partly rugose in dorsal 3; hind femur from rugose to smooth with scattered punctures; Distribution of Z. ekchuah @ and Z. coatlicue @. body length 9.2 to 13.0 mm; usually body color entirely black except yellow hind tar- sus, rarely some or all of following parts also yellow: gena, face, mouthparts, propleuron, mesopleuron, pronotum, mesonotum, fore leg, middle leg, trochanter, trochanterellus, femur and tibia of hind leg, basal '4 of meta- soma. Distribution.—Map 3. This is the only species of Zacremnops in the U.S.A., oc- curring in Florida and southern Texas. It is widespread throughout Mexico except in the north-central region, widespread through- out the Greater Antilles, and Central Amer- ica south to northern Colombia and Ven- ezuela. Type material.—Holotype male, CUBA, (ANSP) (examined). VOLUME 92, NUMBER 3 569 MAP 3 @ Z. cressoni Map 3. Distribution of Z. cressoni. Specimens with ventral surface of hind femur smooth with punctures @. Specimens with ventral surface of hind femur rugose to rugose punctate Depositories.— The 440 specimens that I have identified are in the following collec- tions: AEIC, AMNH, BMNH, CNCI, FSCA, MCZC, MNHN, TAMU, UGCA, USNM. Remarks.—There appear to be two dis- tinct morphotypes of Z. cressoni. Speci- mens from the Greater Antilles, Florida and the Yucatan peninsula differ from the re- mainder of the specimens in that the ventral surface of the hind femur is smooth with scattered punctures (Fig. 8) as opposed to rugose (Fig. 9). Members of this population (Greater Antilles etc.) are generally smaller, darker and have fewer flagellar segments. On the basis of these character states, it seems possible that the two populations constitute separate species. Since the known distributions of the two groups are allopat- ric, I take a conservative approach and treat them as one species. Zacremnops ekchuah, NEw SPECIES Etymology.—Named after Ekchuah, Maya god of merchants, who is said to cover himself in black paint. Diagnosis. — Males and females. Body en- tirely black. Description. — Males and females. Anten- na with 45-47 flagellomeres; metapleuron rugose reticulate in ventral 74, smooth to completely rugose in dorsal 3; hind femur from weakly rugose punctate to smooth with scattered punctures ventrally; body length 10.6 to 15.0 mm; body color entirely black. 570 Distribution.—Map 2. Found in semi- tropical and tropical areas from Mexico south to Costa Rica. Type material.—Holotype, female, MEXICO, Colima, 9 mi. (14.4 km) nee. Comala, 17-18.VII.1983, (Kovarik, Har- rison, Schaffer, CNCI type #20268). Paratypes.—Allotype, male, same data as holotype. COSTA RICA: 2 males, Turrial- ba, VIII.1963, (Porter, MCZC). 1 male, Turrialba, 24.V.1944, (Schrader, USNM). GUATEMALA: 1 female, Yepocapa, VIII.1949, (Dalmat, USNM). MEXICO: Chiapas: 20-25 mi. (32-40 km) N. Huixtla, 3000’ (1000 m), 2. VI. and 4.VI.1969, (Tes- key and Peterson, CNCI). Durango: 2 males, Nombre de Dios, VIII.1951, (Evans, AEIC). Guerrero: | male, 2.1 mi. (3.4 km) N. Ca- cahuamilpa, 19.VII.1984, (Carroll, Schaff- ner, Friedlander, TAMU). 4 males, Omil- temi, 8000’ (2700 m), VIII.1904, (Smith, BMNH). 11 females, 4 males, Xucuma- natlan, VII.1904, (Smith, BMNH). Jalisco: (Flint and Ortiz, USNM). | female, Gua- dalupe, 26.VII.1951, (Hurd, USNM). 1 male, Puente Grande, 5000’ (1700 m), 20.VIHI.1954, (Chillcott, CNCI). Michoa- can: | male, 18 mi. (29 km) N.W. Quiroga, 6400’ (2100 m), 22. VIII.1962, (Painter and Painter, AEIC), Morelos: 5 females, Cuer- navaca, VII-VIII.1944 and 1965, (Krauss, USNM). 1 female, Cuernavaca, IX.1923, (Smyth, USNM). | male, 5 mi. (8 km) E. Cuernavaca, 16.VII.1963, (Parker and Stange, USNM). 2 females, Teptoztlan, 20.VIII.1956, (Dreisbach, USNM), 1 fe- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON male, 3 mi. (4.8 km) S. Tepoztlan, 16. VIII.1962, (Painter and Painter, AEIC). Sinaloa: 2 females, 3 males, Santa Lucia, 4000’ (1300 m), 25. VII-4. VIII.1964, (Ma- son and McAlpine, CNCI). Tabasco: | fe- male, | male, Teapa, H-III.1904, (Smith, BMNH). Yucatan: | male, Pisté, V.1957, (Townes, AEIC). ACKNOWLEDGMENTS I thank Gene Bisdee and Greg Esnard for technical assistance and Evert Lindquist and William Mason for reviewing an earlier draft of the text. REFERENCES Arnett, R. H.and G. A. Samuelson. 1986. The Insect and Spider Collections of the World. E. J. Brill/ Flora & Fauna Publications, Gainesville. 220 pp. Cameron, P. 1887. Insecta. Hymenoptera, |. Biol. cent-am. 1: 1-487. Cresson, E. T. 1865. On the Hymenoptera of Cuba. Proc. Entomol. Soc. Phil. 4: 1-200. De Fernandez, C. D. B. 1987. El genero Zacremnops Sharkey y Wharton (Hymenoptera, Braconidae, Agathidinae) en Argentina y Bolivia. Acta Zool. Lilloana 39(1): 89-93. Enderlein, G. 1920 (1918). Zur Kenntnis aussereu- ropadischer Braconiden, Agathidinae. Arch. Na- tugeschichte 84(A)11: 162-213. Sharkey, M. J. and R. A. Wharton. 1985. Redefini- tion of Megagathis, and reassignment of New World species to Zacremnops new genus (Hy- menoptera: Braconidae: Agathidinae). Can. Ento- mol. 117: 599-603. Szépligeti, G. V. 1902. Tropische Cenocoelioniden und Braconiden aus der Sammlung des Ungfar- ischen National-Museums. Termész. Fiizetek 25: 39-84. . 1904. Hymenoptera Fam. Braconidae. Gen- era Insectorum 4(22): 253 pp. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 571-583 A REVISION OF THE NEARCTIC SPECIES OF DICERURA KIEFFER (DIPTERA: CECIDOMYIIDAE) A. BORKENT * Biosystematics Research Centre, Agriculture Canada, Research Branch, Central Ex- perimental Farm, Ottawa, Ontario, K1A 0C6, Canada. Abstract.—This revision recognizes 8 species of Dicerura in the Nearctic Region in- cluding 6 as new. Descriptions are based on the male adults. A key is provided to all species. Key Words: North America, Palaearctic Species in the genus Dicerura Kieffer are relatively distinctive within the Cecido- mylidae. They are medium sized to rela- tively large and males can be recognized as members of the genus as alcohol or pinned specimens through examination of their wings, antennal flagellomere numbers and their modified genitalia. Overall, the Cecidomyiidae are one of the most poorly taxonomically understood groups of Diptera (Vockeroth 1979). This is reflected in the present revision, where only two species of Dicerura have been pre- viously named in North America (one in another genus) and an additional 6 species are recognized on the basis of only 14 spec- imens. The species in the Palaearctic are better known. Mamaev (1960, 1964, 1966, 1968, 1972, 1975) has described a number of species in the USSR. The species occurring in Latvia, which includes many of those more broadly distributed in the Palaearctic region, have been recently revised by Spun- gis (1987). * Current address: 2330—70th St. SE, Salmon Arm, British Columbia, VIE 4M3, Canada. MATERIALS AND METHODS This study was based on the examination of 81 males of Dicerura, housed in either the CNCI or USNM. Requests for addi- tional material resulted in only negative responses. Cecidomyiidae are not well rep- resented in most North American collec- tions! The acronyms used to represent the mu- seums from which material was studied are those provided by Arnett and Samuelson (1986): CNCI—Canadian National Collection of Insects, Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario, KIA 0C6, Canada. NYSM—New York State Museum, Biolog- ical Survey, 3132 Cultural Education Center, Albany, New York, 12230, USA. USNM-— United States National Entomo- logical Collection, Dept. of Entomol- ogy, U.S. National Museum of Natural History, Washington, DC., 20560, USA. The few specimens studied for this revi- sion were collected either by sweeping, with malaise traps or with a light trap. 572 Of the Palaearctic species, I have been able to examine material of only D. iridis and D. rossica. Otherwise, the Nearctic ma- terial described in this paper was compared only to literature descriptions. Adults were preserved in 70% ethanol and were mounted on microscope slides using a method developed by Leo Forster of our centre. Adults had their wings removed and placed in 15% acetic acid. The head and abdomen were dissected from the thorax and all were placed in 10% KOH which was then heated in a hot water bath. When fully cleared these were placed with the wings in the acetic acid. All parts were then placed through successive baths of 100% 2-pro- panol, 2-propanol layered over clove oil, pure clove oil, where the antennae and left legs were further separated from the head and thorax respectively, and finally into Canada Balsam on the slide. The antennae and legs were sometimes removed while the specimen was in the Canada Balsam. Structures were measured using a mi- crometer in a Nikon compound micro- scope. I used the terms provided by Gagné (1981) for discussing various characters. Type specimens in the Canadian Natio- nal Collection are given numbers in a ref- erence text and these numbers are reported here with the description of the type locality of each named species as “CNC No.’ Some locality labels are accompanied by a ‘CD’ number. These refer to detailed notes for a given locality housed in the Diptera Unit of the Biosystematic Research Centre. Because of a lack of phylogenetic reso- lution, the species are arranged alphabeti- cally in the text. Dicerura Kieffer Dicerura Kieffer 1898: 57. Type-species Di- cerura scirpicola Kieffer (by monotypy). Iridomyza Ribsaamen 1899: 67. Type- species Iridomyza kaltenbachii Ribsaa- men (= Dicerura iridis (Kaltenbach)) (by monotypy). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hormosomyia Felt 1919: 220. Type-species Hormosomyia oregonensis Felt (by orig- inal designation). Ulmomyia Mamaev 1960: 1521. Type- species Ul/momyia rossica Mamaev (by original designation). Neosynepidosis Parnell 1971: 313. Type- species Neosynepidosis furcata (Felt) (by original designation). NEw SYNONYM. Diagnosis. — Males: only Holarctic Ceci- domyiidae with the following combination of character states: 14 antennal flagello- meres; first tarsomere of each leg shorter than second; M, present (in some barely discernable); CuA, absent; parameres fused; gonostylus lacking tooth or dense apical brush. Description. — Male: coloration: head, thorax, legs, abdomen light to dark brown, thorax with vittae evident; wings pale to generally infuscated. Head. eye bridge either present or absent (with a gap equal to 2-3 ommatidia); an- tenna with 14 flagellomeres; flagellomeres with well developed node and neck, each with basal encircling circumfila, with one or two extensions distally; palpus with 4-5 seg- ments. Thorax: with dorsal outline of scutum and scutellum in lateral view forming an unin- terrupted curve; dorsocentral, acrostichal, anepisternal setae present; anepimeral, kat- episternal setae present or absent. Wing: with macrotrichia on membrane and veins; Rs lying in similar direction as R,.5; Ry, extending past apex of wing; apex of M, ,, present or absent; apical portion of M; present or absent; CuA, absent; CuA, present. Legs: first tarsomere of each leg with api- cal projection; claws with 24 ventral teeth; empodium about '4-'» length of claw. Abdomen: posterior setae on tergites ar- ranged in continuous transverse row or in two lateral groups. Genitalia: tergite 9 densely pruinose; cerci present on hypoproct; parameres fused, forming an aedeagal guide posterodorsally, VOLUME 92, NUMBER 3 Table 1. Dicerura. List of species now placed in the genus adunca n. sp. Borkent. Ontario, Canada. barbata Mamaev 1966: 226. Ukrainian SSR. carpiensis n. sp. Borkent. Ontario, Canada. cooperi n. sp. Borkent. Newfoundland, Canada. complicata Spungis 1987: 27. Latvia. curva n. sp. Borkent. Ontario, Canada. dentata Spungis 1979: 84. Latvia. elongata n. sp. Borkent. Arizona, USA. foliicola Mamaev 1968: 614. Maritime Territory, USSR. fungicola (Mamaev) 1964: 904 (Ulmomyia). Moscow Region, USSR. furcata (Felt) 1907: 52 (Winnertzia). New York, USA. NEW COMBINATION. furculata Mamaev 1968: 614. Ukrainian SSR. iridis (Kaltenbach) 1874: 717 (Cecidomyia). Germany. kaltenbachi (Riibsaamen) 1899: 67 (Iridomyza). Germany. loba n. sp. Borkent. Ontario, Canada. mixta Spungis 1987: 24. Latvia. oregonensis (Felt) 1919: 220 (Hormosomyia). Oregon, USA. padi Mamaev 1975: 60. Maritime Territory, USSR. rossica (Mamaev) 1960: 1521 (U/momyia). Voronezh region, USSR. scirpicola Kieffer 1898: 57. Europe. scirpi Kieffer 1899: 165 (new name for scirpicola). separata Spungis 1987: 26. Latvia. stipator Mamaev 1972: 113. Maritime Ternitory, USSR. triangularis Mamaev 1966: 227. Ukrainian SSR. unidentata Spungis 1987: 20. Latvia. xvlophila Mamaev 1966: 227. Ukrainian SSR. with posterolateral barbs present or absent; aedeagus elongate, single or bifid posterior- ly; gonocoxite with mediobasal, pruinose lobe; gonostylus lacking tooth, with or with- out developed mediobasal lobes. Distribution and bionomics. —The genus is presently known only from the Holarctic Region. Biological information is available for only some of the Palaearctic species where larvae have been collected from the leaf axils of various plants such as Scirpus sylvaticus, Iris pseudacorus, and Acorus ca- lamus, from rotting wood, from decaying leaves, or from soil (Mamaev 1973, Spungis 1987). 573 Taxonomic discussion.—All the species that are presently included in Dicerura are listed in Table 1. D. indica Grover has been recently transferred to Cryptoneurus by Grover (1981). I consider the genus Neosynepidosis as a synonym of Dicerura because of the fun- damental similarity between the species of the two genera. The characters previously considered to distinguish Neosynepidosis are now known within Dicerura: the presence of circumfila with distal extensions and the presence of an unforked aedeagus. The only other genus in the Dicerurini in which males are reported to lack a tooth on the gonostylus is in the monotypic genus Synepidosis Mamaev (Mamaev 1964). However, Dr. V. Spungis (pers. comm.) has examined the type of S. longiventris Ma- maev (from Voronezh Province, USSR) and reports that the gonostylus does in fact bear a small tooth. Mamaev (1966) reported in a generic di- agnosis that some Dicerura lack teeth on their tarsal claws. Examination of material and literature descriptions show or report that all Dicerura possess at least one tooth on their claws. However, one specimen of D. oregonensis had the teeth broken off one midleg claw, indicating that this condition may be an artifact of age or preparation. Female Dicerura have been previously described only from several Palaearctic species (Kieffer 1899, Mamaev 1960, 1964, 1966, 1968, Panelius 1965, Spungis 1987). Although I collected several female Dicer- ura in the Nearctic, I was unable to confi- dently associate these with males, which ap- pear to be more easily differentiated from one another than are females. I was there- fore unable to include descriptions of iden- tified females in this paper. Furthermore, the morphological variation of these and previously described Palaearctic females, in combination with our generally poor un- derstanding of the females of Cecidomyi- idae, does not allow for a generic diagnosis of the females at this time. Similarly, the 574 larvae of some Palaearctic species have been described (Kaltenbach 1874, Mamaev and Krivosheina 1965, Riibsaamen 1899, Spun- gis 1979, 1987) but they too, for the same reasons as for the females, cannot be diag- nosed generically. Panelius (1965) provided a cladistic anal- ysis of the generic relationships within the Porricondylinae. His major groupings in- dicated that the subfamily is paraphyletic. No synapomorphy was proposed for Dicer- ura and the monophyly of the genus 1s there- fore uncertain. The only character state which may argue for recognition of this clade is the lack of a tooth on the gonostylus of the male. However, outgroup comparisons within the Lestreminae and Cecidomyiinae indicate that both toothed and bare gono- stylil are present. In addition, the bare gono- stylus of Dicerura is not unique within the Porricondylinae (e.g. present in some Win- nertzini, some Porricondyla), indicating that the character is susceptible to homoplasy. Consequently, the genus may not be mono- phyletic. Nevertheless, there is a general similarity of appearance of members of the group and they are presented as a single genus in this paper. KEY Members of the genus Dicerura may be recognized as such in the Holarctic Region using Gagné (1981), with the consideration that Neosynepidosis 1s considered a syn- onym of Dicerura here. The outline of the gonostylus can be markedly affected by differences in position. Care must be taken, therefore, in comparing specimens to the illustrations provided here. Key TO ADULT MALES OF NEARCTIC DICERURA SPECIES 1. Gonostylus with markedly developed medio- basalilobel(Figss3 ANB D946) ie. eect cacae 2) — Gonostylus with no or only slightly developed mediobasal lobe (Figs. 3C, 4A, B, D) ........ 5 2. Gonostylus with portion distal to mediobasal lobes elongate, apex abruptly bent ventrally (Fig. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3D); parameres lacking posterolateral barbs .. 5 eto See Tra aoa S MDa a curva Nn. sp. — Gonostylus with portion distal to mediobasal lobes relatively short and apex, at most, some- what bent medially (Figs. 3A, B, 4C); para- meres with posterolateral, stout barbs 3. Mediobasal lobe pointed distally (Fig. 3A); ae- deagus with a single apex and an associated sclerite; tergite 9 markedly bilobed .. adunca n. sp. — Mediobasal lobe rounded distally (Figs. 3B, 4C); aedeagus biramous apically, lacking associated sclerite; tergite 9 truncated or slightly bilobed ... 4 4. Apex of aedeagus with divergent and separate ends (Fig. 3B); parameres parallel sided pos- teriorly; flagellomere 3 with stem 0.50-0.67 times length of basal node (Fig. 1B) Bef a, Mice Pee tach Rees MOA OE carpiensis N. sp. — Apex of aedeagus with ends closely appressed (Fig. 4C); parameres somewhat bulbous pos- teriorly; flagellomere 3 with stem 0.94 times length of basal node (Fig. 1G) ....... loba n. sp. 5. Aedeagus with single apex (Fig. 4B); paramere lacking posterolateral barbs; flagellomere 3 with stem 0.58—0.86 times length of basal node (Fig. LB): CREA Sa he PAR OS furcata (Felt) — Aedeagus with biramous apex (Figs. 3C, 4A, D); paramere with posterolateral barbs; flag- ellomere 3 with stem at least 1.00 times length of basal node (Fig. 1C, E, H) 6. Tergite 9 truncated posteriorly (Fig. 4A); ae- deagus with ends of biramous apex closely ap- pressed for most of length; gonostylus elongate and of nearly equal diameter; flagellomere 3 with stem 1.06 times length of basal node (Fig. DE) FSi eee eee elongata n. sp. — Tergite 9 rounded or bilobed apically (Fig. 3C, 4D); adeagus with ends of biramous apex di- vergent and distinctly separated; gonostylus ta- pering to apex; flagellomere 3 with stem more than 1.41 times length of basal node (Fig. 1C, 139) snob eee 7. Parameres forming rounded lobe posteriorly, with apex with bilobed nipple (Fig. 3C); gono- stylus thick, squat for most of its length SO sre Lae en oitG eee cooperi n. sp. — Parameres forming triangular lobe posteriorly, with apex with single nipple (Fig. 4D); gono- stylus evenly tapered posteriorly Dicerura adunca Borkent, NEw SPECIES Types.— Holotype, male adult on micro- scope slide, labelled ““Holotype Dicerura VOLUME 92, NUMBER 3 Circumfilum Fig. 1. Third flagellomere of male antenna. Setae and spicules not drawn. Scale = 0.1 mm. A, D. adunca. B, D. carpiensis. C, D. cooperi. D, D. curva. E, D. elon- gata. F, D. furcata. G, D. loba. H, D. oregonensis. adunca Borkent, 6, Ancaster Spring, Valley, Cons. Area, Wentworth Co., Ont., 6-7-VI- 1984, I. M. Smith, CD207, CNC NO. 20343” (CNCI); paratype: | male labelled as for holotype (CNCI). Diagnosis.— Male: only Nearctic species in which the circumfilum has two exten- sions reaching to the distal margin of the flagellomere node; otherwise, this is the only species in which the mediobasal lobe of the gonostylus is short and hooked. Description of male adult. — Head: dorsal eye bridge lacking 1-2 ommatidia medially; antennal length/wing length = 0.90; anten- nal flagellomere 3 with stem 1.00-1.39 length of basal node (Fig. 1A), circumfilum a basal ring with two extensions reaching to BY) distal margin of node; palpus with 5 seg- ments. Wing: length = 2.5 mm; M, ,, absent, M,; present apically; halter with moderately elongate stem (Fig. 2A). Genitalia (Fig. 3A): tergite 9 narrowed posteriorly, bilobed; gonocoxites with slight bilobed posteromedial projection; gono- stylus with anteroventral dense patch of short setae, with short, hooked, densely se- tose mediobasal lobe, gonostylus of similar diameter to rounded apex; paramere form- ing elongate, somewhat parallel sided pro- jection posteriorly, with very rounded apex, posterolateral margins with double row of short, stout barbs; aedeagus undivided at midlength, with single pointed apex, with subapical barbs, associated oval sclerite. Taxonomic discussion.—Aside from the characters noted in the diagnosis, this species is unique in the presence of a separate scler- ite associated with aedeagus. Derivation of specific epithet.— The name adunca (bent inward) refers to the shape of the mediobasal lobe on the gonostylus. Dicerura carpiensis Borkent, NEw SPECIES Types. — Holotype, male adult on micro- scope slide, labelled “Holotype Dicerura carpiensis Borkent, ¢, 3 km E. Carp, Ont., 19-V-3-VI-1983, A. Borkent, CD42, Ma- laise trap, CNC No. 20344” (CNCI); para- types: | male labelled as for holotype; 1 male from km 75-125, Dempster Hwy, Yu- kon Territory (19-VI-1984, S. & J. Peck) (CNCI); 1 male from Springwater Cons. area, nr. Alymer, Ont. (27-VI-11-VII-1984, K. Ferguson) (CNCI); | male from Motts Creek, Atlantic Co., New Jersey (21-V-?, R. J. Gagné). Diagnosis.— Male: only Nearctic species with a gonostylus with an apically rounded mediobasal lobe and with a relatively short stem on the flagellomeres (flagellomere 3 with stem 0.50-0.67 times length of basal node). Description of male adult. — Head: dorsal 576 eye bridge 3-4 ommatidia wide; antennal length/wing length = 0.64—-0.69; antennal flagellomere 3 with stem 0.50-0.67 length of basal node (Fig. 1B), circumfilum a basal ring with a single distal extension; palpus with 5 segments. Wing: length = 2.35-2.81 mm; M, ,, ab- sent, M, present apically; halter with mod- erately elongate stem (Fig. 2B). Genitalia (Fig. 3B): tergite 9 somewhat truncated to rounded posteriorly; gonocox- ites with single, posteromedial concavity with small, medial, posteriorly directed lobe, bordered laterally by slightly developed lobes; gonostylus with anteroventral dense patch of short setae, with well developed, densely setose mediobasal lobe, gonostylus distal to mediobasal lobe gradually taper- ing; paramere forming elongate, somewhat parallel-sided projection posteriorly, with apex bilobed, posterolateral margins with short, stout barbs; aedeagus undivided at midlength, biramous for apical quarter, with ends distinctly separate for their entire length, lacking spicules. Taxonomic discussion.—All specimens were collected with malaise traps except for the male from the Yukon Territory which was collected with a car net (net mounted on a vehicle). Derivation of specific epithet. —The name carpiensis refers to the type locality, near Carp, Ontario, where the author resided for several years. Dicerura cooperi Borkent, NEw SPECIES Type.— Holotype, male adult on micro- scope slide, labelled “Holotype Dicerura cooperi Borkent, 6, CNC No. 20345, 3 km. N. Picadilly, Nfld. [Newfoundland], 25-VI- 83, A. Borkent, CD71, Sweeping grass” (CNCI). Diagnosis.— Male: only Nearctic species with parameres forming rounded lobe pos- teriorly, with its very apex bearing a bilobed nipple and with a thick, squat gonostylus bearing a short mediobasal lobe. Description of male adult. — Head: dorsal PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eye bridge 2 ommatidia wide; antennal length/wing length unknown (terminal flag- ellomeres missing); antennal flagellomere 3 with stem 1.76 length of basal node (Fig. 1C), circumfilum a basal ring with a single distal extension; palpus with 5 segments. Wing: length = 2.77 mm; M,,, absent, M, present apically; halter with elongate stem (Fig. 2C). Genitalia (Fig. 3C): tergite 9 slightly bi- lobed posteriorly; gonocoxites with single, medially convex posteromedial concavity, bordered laterally by slightly developed lobes; gonostylus with anteroventral dense patch of short setae, with short, densely de- tose mediobasal lobe, gonostylus tapering gradually to apex, small apical projection slightly bent medially; paramere forming wide, somewhat rounded projection pos- teriorly, very apex with bilobed nipple, pos- terolateral margins with short, stout barbs; aedeagus undivided at midlength, biramous for apical half with ends distinctly separate and ridged, lacking spicules. Taxonomic discussion.—D. cooperi 1s somewhat similar to the Palaearctic D. den- tata. However, D. dentata differs in having the apex of the parameres entirely rounded while in D. cooperi the parameres have an apical, bilobed nipple. In addition, the ae- deagus of D. dentata 1s straight apically but this may be an artifical difference due to position. Derivation of specific epithet.—The name cooper! is given in recognition of Mr. Bruce E. Cooper. A number of specimens he col- lected were important to this study and re- flect his major contributions to the growth, health and welfare of the Diptera collection housed in the Canadian National Collec- tion. Dicerura curva Borkent, NEW SPECIES Types. — Holotype, male adult on micro- scope slide, labelled ‘“‘Holotype Dicerura curva Borkent, 6, Springwater Cons. area, nr. Alymer, Ont., 27-VI-11-VII-1984, K. Ferguson, CD 253, CNC No. 20346” VOLUME 92, NUMBER 3 II if S77 Cc D G H ———— Fig. 2. Outline of male halter. Scale = 0.5 mm A, D. adunca. B, D. carpiensis. C, D. cooperi. D, D. curva. E, D. elongata. F, D. furcata. G, D. loba. H, D. oregonensis. (CNCI); paratypes: 1 male from 4 km NW Kagawong, Manitoulin Is., Ontario, 1-16- VI-1982, A. Ritchie (CNCI); | male from Cheticamp, Nova Scotia, 6-VI-1984, B. E. Cooper (CNCI). Diagnosis. — Male: only Nearctic species with a gonostylus with a well developed me- diobasal lobe and an apex which is bent ventrally. Description of male adult. — Head: dorsal eye bridge 3-6 ommatidia wide; antennal length/wing length = 0.62-0.67; antennal flagellomere 3 with stem 0.76-0.88 length of basal node (Fig. 1D), circumfilum a basal ring with a single distal extension; palpus with 5 segments. Wing: length = 1.9-2.4 mm; M, ,, absent, M, barely discernable apically; halter with relatively short stem (Fig. 2D). Genitalia (Fig. 3D): tergite 9 somewhat truncated to rounded posteriorly; gonocox- ites with single, posteromedial concavity with small, medial, posteriorly directed lobe, bordered laterally by slightly developed 578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON d t SN -Ppee WA. Fig. 3. Male genitalia in ventral view; apex of tergite 9 drawn below. A, D. adunca. B, D. carpiensis. C, D. cooperi. D, D. curva. VOLUME 92, NUMBER 3 lobes; gonostylus with anteroventral dense patch of short setae, with well developed, densely setose mediobasal lobe, gonostylus distal to mediobasal lobe of equal diameter, with very apex abruptly directed ventrally; paramere forming elongate, somewhat par- allel-sided projection posteriorly, with apex bilobed, posterolateral margins lacking barbs; aedeagus undivided at midlength, biramous for apical half, with ends distinct- ly separate for their entire length, lacking spicules. Taxonomic discussion.—This species is very similar to the Palaearctic D. separata but tergite 9 is not so tapered apically, the parameres have posteriorly directed barbs apically, and the mediobasal lobe of the gonostylus is proportionally larger. If fur- ther collection supports the view that these two are separate species, they are probably sister species, based on the shared unique conformation of the apex of the gonostylus. The two specimens from Ontario were collected with a Malaise trap. Derivation of specific epithet. —The name curva refers to the unusual subapical curve in the gonostylus of the male of this species (shared by D. separata in the Palearctic). Dicerura elongata Borkent, NEw SPECIES Type.— Holotype, male adult on micro- scope slide, labelled ““Holotype Dicerura elongata Borkent, 6, USNM, Portal, Co- chise Co., Ariz., VII-1-67, at light, Saul and Suzy Frommer, Can. Balsam’? (USNM). Diagnosis.— Male: only Nearctic species with a markedly elongate gonostylus, of nearly constant diameter, and lacking a well defined mediobasal lobe. Description of male adult. — Head: dorsal eye bridge 6 ommatidia wide; antennal length/wing length = 0.91; antennal flag- ellomere 3 with stem 1.06 length of basal node (Fig. 1E), circumfilum a basal ring with a single distal extension; palpus with 5 seg- ments. Wing: length = 2.39 mm; M,,, present 579 apically, M, present apically; halter with short stem (Fig. 2E). Genitalia (Fig. 4A): tergite 9 truncate with posterolateral corner with lobe; gonocoxites with posteromedial concavity with medial spiculose lobe, bordered laterally by fleshy lobes; gonostylus with anteroventral dense patch of short setae, lacking mediobasal lobe (patch of spicules present), gonostylus of nearly constant diameter to rounded apex; paramere forming expanded projection pos- teriorly, truncated apically, posterolateral margins with short, stout barbs; aedeagu divided at midlength, biramous apically with ends closely appressed, lacking spicules. Derivation of specific epithet.—The name elongata refers to the distinctive, elongate gonostylus of the male of this species. Dicerura furcata (Felt), NEw CoMBINATION Winnertzia furcata Felt 1907: 148. Holo- type, male adult on microscope slide, not seen (housed in USNM on long term loan from NYSM). Type locality, Nassau, New York. Asynapta furcata: Felt 1908: 420. Neosynepidosis furcata: Parnell 1971: 313. Diagnosis.— Male: only Nearctic species with a gonostylus lacking a mediobasal lobe and with the aedeagus undivided. Description of male adult. — Head: dorsal eye bridge 6-8 ommatidia wide; antennal length/wing length = 0.83-0.86; antennal flagellomere 3 with stem 0.58-0.86 length of basal node (Fig. 1F), circumfilum a basal ring with a single distal extension; palpus with 5 segments. Wing: length = 2.6-3.3 mm; M, ,, present apically, M, present apically; halter with rel- atively short stem (Fig. 2F). Genitalia (Fig. 4B): tergite 9 broad, slight- ly bilobed; gonocoxites with single, evenly curved posteromedial concavity bordered laterally by fleshy lobes, gonostylus with an- teromedial dense patch of short setae, lack- ing mediobasal lobe, gonostylus broadly bent 580 at midlength; paramere forming parallel sided projection posteriorly, posterolateral margins lacking barbs but forming convo- luted thick cuticle; aedeagus divided at mid- length, biramous apically with ends closely appressed, lacking spicules. Taxonomic discussion.—The justifica- tion for considering this species, previously placed in Neosynepidosis, as a member of Dicerura is noted above in the taxonomic discussion of the genus. Material examined.—I have examined 54 males from the following localities: 14 mi. (22 km) E. Dawson, Y.T.; 40 km. E. Daw- son, Y.T.; Kemptville, Ontario; 4 km N. Metcalf, Ontario; 3 km SW Richmond, On- tario; Dunbar Lk., St. Mauricie Provincial Park, Quebec; 19 km N. Grand-Mere, Que- bec. Specimens were collected from May 31 to August 22. Dicerura loba Borkent, NEw SPECIES Types.— Holotype, male adult on micro- scope slide, labelled ““Holotype Dicerura loba Borkent, 4, 3 km E. Carp, Ont., 21-V- 11-VI-1984, A. Borkent, CD251, CNC No. 20347” (CNCI); paratype: 1 male labelled as for holotype (CNCI). Diagnosis.— Male: only Nearctic species with the apex of the aedeagus with ends closely appressed, the parameres somewhat bulbous posteriorly and flagellomere 3 with its stem 0.94 times length of the basal node. Description of male adult. — Head: dorsal eye bridge without ommatidia medially; an- tennal length/wing length = 0.89; antennal flagellomere 3 with stem 0.94 length of basal node (Fig. 1G), circumfilum a basal ring with a single distal extension; palpus with 5 seg- ments. Wing: length = 2.12-2.55 mm; M,,, barely discernable or absent, M, present apically; halter with moderately elongate stem (Fig. 2G). Genitalia (Fig. 4C): tergite 9 slightly bi- lobed; gonocoxites with single, evenly curved posteromedial concavity bordered PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON laterally by moderately developed lobes; gonostylus with anteroventral dense patch of short setae, with well developed, densely setose mediobasal lobe, gonostylus tapering gradually to rounded apex; paramere form- ing wide projection posteriorly, postero- lateral margins with short, stout barbs; ae- deagus undivided at midlength, thicker for apical third, biramous apically with ends closely appressed for basal portion, distinct- ly separate at very apex, lacking spicules. Derivation of specific epithet. —The name loba refers to the shape of the mediobasal lobe on the gonostylus of this species. Dicerura oregonensis (Felt) Hormosomyia oregonensis Felt 1919: 220. Holotype, male adult on microscope slide, labelled ““Type n.g. s.sp. Entomologic Di- vision Hormosomyia oregoniensis (sic) F.R. Cole. 4 Felt. Forest Grove, Ore., C1790, N.Y. State Museum, 13 Mar. 1919” (housed in USNM on long term loan from NYSM). Dicerura oregonensis: Mamaev 1966: 226; Parnell 1971: 308. Diagnosis.— Male: only Nearctic species with a gonostylus which tapers gradually to its apex, with a poorly defined mediobasal lobe. Description of male adult. — Head: dorsal eye bridge 3-8 ommatidia wide; antennal length/wing length = 1.06-1.14; antennal flagellomere 3 with stem 1.41-1.92 length of basal node (Fig. 1H), circumfilum a basal ring with one or two distal extensions; pal- pus with 5 segments. Wing: length = 3.0-5.6 mm; M, ,, present apically, M, present apically; halter with moderately elongate stem (Fig. 2H). Genitalia (Fig. 4D): tergite 9 narrowed posteriorly, slightly bilobed; gonocoxites with single, evenly curved posteromedial concavity bordered laterally by moderately developed lobes; gonostylus with antero- ventral dense patch of short setae, with short to very short, rounded to angular, in some 581 VOLUME 92, NUMBER 3 Fig. 4. Male genitalia in ventral view; apex of tergite 9 drawn below. A, D. elongata. B, D. furcata. C, D. loba. D. D. oregonensis, with variation in gonostylus shape shown. 582 poorly defined, densely setose mediobasal lobe, gonostylus tapering gradually to apex, small apical projection slightly bent medi- ally, medial surface of gonostylus smooth to jagged; paramere forming triangular pro- jection posteriorly, posterolateral margins with short, stout barbs, some with subapical margin also with barbs; aedeagus undivided at midlength, biramous apically with ends diverging, a few spicules present or absent. Taxonomic discussion. —Specimens of D. oregonensis were collected by sweeping, with a malaise trap, or at a light. Of the six specimens examined for this study, some differences were noted in the shape of gonostylus and in the shape and extent of the posterolateral barbs of the par- ameres. These character states should be re- studied once more material becomes avail- able to test the possibility of the presence of more than one species under this name. The specimen from the Yukon Territory had palpal segments 4 and 5 fused on one side but completely separate on the other. Although the type label bears a date of March 13, 1919, Felt (1919) reported the date of collection as October 10, 1918. This later date is more consistent with the other specimens of D. oregonensis which were col- lected late in the season. D. oregonensis is similar to several Pa- laearctic species. However, I consider the following differences to be significant. In D. xylophila the gonostylus is relatively longer, in D. triangularis the gonostylus is relatively shorter and broader, in D. iridis the ends of the biramous aedeagus are divergent from this base, the spicules on the aedeagus are restricted to the very apex, and the medial area of the gonocoxites is a straight line, and in D. rossica the apex of the parameres is rounded. Material examined.—Five specimens, aside from the holotype, were studied: one specimen from each of Falls Church, Vir- ginia (10-IX-1960, W W. Wirth), km 155, Dempster Highway, Yukon Territory (12- VII-1984, D. M. Wood), 11 km E. Griffith, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ontario (9—20-IX-1984, B. E. Cooper) and 2 specimens from Salmon Arm, British Co- lumbia (29-VIII-1988, A. Borkent). Derivation of specific epithet.—The name oregonensis refers to the state from which the holotype was collected. ACKNOWLEDGMENTS I thank Mr. Leo Forster for mounting most of the specimens on microscope slides. His typically excellent preparations were crucial to this study. Mr. Barry Flahey drew most of the male genitalia for this paper and I appreciate his skills in this area. The genitalia of D. ore- gonensis and D. furcata were drawn by Mr. Ralph Idema. Ms. Barbara Bissett provided help with some library work and labelling. Dr. V. Spungis kindly compared my drawings of Nearctic Dicerura to those Pa- laearctic species in his collection and made comments on their similarities or differ- ences. He also kindly reviewed a manuscript of this revision. I also thank Dr. R. J. Gagné for lending material from the USNM (Washington, D.C.). Drs. Yves Bousquet and Lubomir Masner made helpful remarks on an early draft of this paper. Finally I thank Mr. Bruce Cooper (of our Centre) and Mr. Kevin Ferguson (Outdoor Education Center, Alymer, Ontario) for servicing malaise traps which resulted in the capture of some of this material. LITERATURE CITED Arnett, R. H. and G. A. Samuelson 1986. The insect and spider collections of the world. E. J. Brill/Flora and Fauna Pub., Gainesville, Florida, 220 pp. Felt, E. P. 1907. Appendix: New species of Cecido- myiidae, pp. 97-165. Jn his 22nd report of the State Entomologist on injurious and other insects of the State of New York, 1906. N.Y. State Mus. Bull. 110: 39-186. . 1908. Appendix D, pp. 286-422. In his 23rd report of the State Entomologist on injurious and other insects of the State of New York, 1907. N.Y. State Mus. Bull. 124: 5-541. VOLUME 92, NUMBER 3 1919. Five non-gall-making midges (Dip., Cecidomyidae). Entomol. News 30: 219-223. Gagné, R. J. 1981. Cecidomyiidae (chapter) 16, pp. 257-292. In McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 1. Research Branch Ag- riculture Canada Ottawa, Monograph 27, 674 pp. Grover, P. 1981. A catalogue of Indian Gall-midges. Cecidologia Internationale 2: 63-108. Kaltenbach, J. H. 1874. Die Pflanzenfeinde aus der Klasse der Insekten. 848 pp., 401 figs. Stuttgart. Kieffer, J. J. 1898. Synopse des Cécidomyies d’Eu- rope et d’Algerie décrites jusqu’a ce jour. Bull. Soc. Hist. nat. Metz 20: 1-63. . 1899. Ueber Dicerura Kieff. Iridomyza Rbs.). Wien. Entomol. Ztg. 18: 165-169. Mamaev, B. M. 1960. Description of two new genera and one species of gall midges (Itonididae, Dip- tera) developing in rotten wood (in Russian, with English summary). Zool. Zh. 39: 1521-1524. 1964. Gall-midges of the USSR. 6. New species of the tribe Porricondylini (Diptera, Ce- cidomyiidae) (in Russian, with English summary). Entomol. Obozr. 43: 894-913. 1966. New and little known Palaearctic gall midges of the tribe Porricondylini (Diptera, Ce- cidomyiidae) (in Russian, with English summary). Acta Entomol. Bohemoslov. 63: 213-239. 1968. New nematocerous Diptera of the USSR fauna (Diptera, Axymyiidae, Mycetobiidae, Sciaridae, Cecidomyiidae) (in Russian, with En- glish summary). Entomol. Obozr. 47: 605-616. 1972. Review of species and ecological re- lations of insects-decompositors of wood-matter 583 of Ulmus propinqua Loidz. (in Russian). Tr. biol.- pochy. Inst. Vladivostok (N.S.) 7: 106-120. 1973. The biology of Dicerura iridis Kalten- bach—the new species of gall-midges in the fauna of the USSR (in Russian, with English summary). Acta Entomol. Lituanica 2: 177-181. . 1975. Comparative character of entomocom- plexes developing in wood-matter of Padus asia- tica in the south Maritime Territory (in Russian). Tr. biol.-pochv. Inst. Vladivostok (N.S.) 28: 58- 62. Mamaey, B. M. and E. P. Krivosheina. 1965. Larvae of gall midges. Diptera, Cecidomyiidae (in Rus- sian). Akad. Nauk USSR, Moscow. Panelius, S. 1965. A revision of the European gall midges of the subfamily Porricondylinae (Diptera: Itonididae). Acta Zool. Fenn. 113: 1-157. Parnell, J. R. 1971. A revision of the Nearctic Por- ricondylinae (Diptera: Cecidomyiidae) based largely on an examination of the Felt types. Misc. Publ. Entomol. Soc. Am. 7: 275-348. Riibsaamen, E.H. 1899. Ueber Gallmiicken auf Car- ex und Jris. Wien. Entomol. Ztg. 18: 57-76. Spungis, V. V. 1979. Eine neue Art der Gallmicken der Gattung Dicerura Kieffer aus Lettland. Latv. Entomol. 22: 83-87. . 1987. Gall midges of the subtribe Dicerurina (Diptera, Cecidomyiidae) in Latvia (in Russian, with English summary). Latv. Entomol. 30: 15- 42. Vockeroth, J. R. 1979. Mycetophilioidea. pp. 390, 404405. Jn Canada and Its Insect Fauna, H. V. Danks, ed. Mem. Entomol. Soc. Can. 108, 573 pp. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, p. 584 NOTE The type locality of Sciapus pressipes Parent (Diptera: Dolichopodidae) Parent (1929, Annls Soc. sci. Brux. (B) 49: 244-246) described Sciapus pressipes in his account of the non-European material in the von Roder collection, giving the type locality as “‘Caroline.”” He included char- acters to distinguish the species from other american Sciapus, and provided an addi- tional couplet to fit it into Becker’s key to the nearctic and neotropical species of the genus (Becker, 1922, Abh. Zool.-Bot. Ges. Wien. 13(1): 361-363). The species has been included in subsequent keys to North American Sciapus by Robinson (1964, Misc. Publs Ent. Soc. Am. 4: 112-113) and Stey- skal (1966, Proc. Ent. Soc. Wash. 68: 292- 294), and was included in the catalog of North American species by Foote et al. (1965, pp. 485-486 in USDA Agr. Hndbk No. 276). However, these subsequent au- thors all give the distribution as “Carolina,” with no extra localities, nor any indication as to whether North or South Carolina is the appropriate state. This suggests that no further material has been collected. The descriptions of new species in Par- ent’s paper were preceded by a list of lo- calities for both known and previously un- known species (Parent, loc. cit., pp. 169- 173). This gives the origin of his material of S. pressipes as ““Penope (Caroline).” It appears that Penope is an alternative or misspelling of Ponape, and that S. pressipes comes from Ponape in the Caroline Is. It is thus a member of the pacific rather than the nearctic fauna. Parent’s description shows that whereas S. pressipes is unlike any North American species of Sciapus, it is very sim- ilar to S. occultus Parent from the Admiralty Is., particularly in the genitalia and male secondary sexual characters. C. E. Dyte, Department of Pure and Ap- plied Zoology, University of Reading, Whitenights, Reading, England. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 585-586 OBITUARY Anne Ma 1958 Anne Marie Wieber, an entomologist with the Maryland Department of Agriculture and an officer in the Entomological Society of Washington, died suddenly on May 22, 1989. She perished in a helicopter accident near Cumberland, Maryland, while direct- ing spray operations to control gypsy moths. She resided in Wheaton, Maryland, and is survived by her parents Paul R. and Mar- ilyn J. Wieber of Morgantown, West Vir- ginia, and by a sister, Paula, and a brother, Jerome, both residents of Maryland. Anne was born in Lakewood, Ohio, where she spent the early years of her life. She lived in Cary, North Carolina, with her family between 1973 and 1975, and periodically returned to North Carolina to visit and va- cation along the coast. The family moved to Derwood, Maryland, in 1975, where Anne attended Col. Zadok Magruder High School in Rockville, graduating with the class of 1976. She then entered the University of Maryland where she majored in Entomol- ogy, and from which she was awarded the degree of Bachelor of Science in 1980. Anne was employed as an Entomologist by the rie Wiebe -1989 T Forest Pest Management Section of the Maryland Department of Agriculture from 1981 until her death. Anne became actively involved in the problems of entomological research and pest management while an undergraduate em- ployee of the Department of Entomology, maintaining insect colonies used for re- search on insect pheromones, and serving as a scout for ornamental and turf pest man- agement programs. As a scout, she refined skills of communicating with the public that made her an invaluable team member in her later career. During the later part of her undergraduate studies, she was a part time employee of the Vegetable Laboratory at the Beltsville Agricultural Research Center (BARC), Beltsville, Maryland (Agricultural Research Service, United States Depart- ment of Agriculture). Here she supported research on pests of vegetable crops and commercial mushroom production, prepar- ing illustrations of mushroom flies that were used in scientific presentations. Upon graduation, Anne became a mem- ber of an interagency cooperative team 586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON searching for improved methods to manage the gypsy moth under the technical direc- tion of Ralph E. Webb. Although an em- ployee of the Maryland Department of Ag- riculture, she was physically located at BARC, initially in association with the Flo- rist and Nursery Crops Laboratory, and lat- er with the Insect Chemical Ecology Lab- oratory. She participated in a wide variety of projects that involved cooperation with scientists from a number of ARS labora- tories and from the USDA-Forest Service, the Animal Plant Health Inspection Ser- vice, the University of Maryland, and the Illinois Natural History Survey. Anne co- operated in research to control gypsy moth as a pest of homeowners including evalua- tion of homeowner spray products for gypsy moth egg mass control, and ground appli- cation and systemic implantation of insec- ticides for control of gypsy moth on indi- vidual trees. She collaborated with Michael McManus (USDA-FS, Hamden, Connect- icut) to evaluate the use of pheromone traps and burlap banding for gypsy moth moni- toring, and was a key participant in a co- operative effort by the Forest Service, Ih- nois Natural History Survey, and ARS to establish species of gypsy moth adapted mi- crosporidia collected from European gypsy moth populations into North American populations. She led the field research for a detailed study of the hyperparasite complex of the gypsy moth parasite Cotesia melan- oscela in Maryland. Her last research col- laboration involved the design and imple- mentation of a biologically-based gypsy moth suppression program for a county park system. Finally, Anne provided valuable as- sistance each year to the Maryland Gypsy Moth Cooperative Suppression Program, directing a portion of the spray operations. Reflecting her contributions to the project, Anne coauthored 10 published papers, with another in preparation, and her name will appear on several projected manuscripts. Her collaborators in the C. melanoscela study have agreed to publish the results of this research, with Anne to be the senior author. Anne’s interests were not limited to ento- mology. She had an appreciation for all forms of life and her field work provided great satisfaction for the opportunity to ob- serve nature. She enjoyed pets and over the years had kept tarantulas and ferrets as well as more conventional companions. Anne had as many houseplants as practicality and aesthetics would allow in her home and of- fice. She was especially fond of orchids, suc- culents and African violets. During the years she worked on Entomology Road in Belts- ville, the flower beds around her building were replete with snapdragons, impatiens and marigolds. Anne maintained a small plot of roses for several years at Beltsville that were used in research on rose midges and she also helped with and encouraged many other floricultural projects. She was especially helpful to Floyd F. Smith in his last years, accompanying him to his willow plots and cleaning off his sticky boards. Anne is missed not only by her colleagues in Entomology, but by her family and mul- titude of friends. She will be remembered for the enthusiasm and perseverance she displayed in her endeavors, her sense of hu- mor, her love of people. It is a grievous loss that her life and career ended so early. Acknowledgment.—We wish to thank Maureen Gough for her helpful comments. Geoffrey B. White, Vegetable Laboratory, Beltsville Agricultural Research Center, Beltsville, Maryland 20705. Ralph E. Webb, Insect Chemical Ecology Laboratory, Beltsville Agricultural Research Center, Beltsville, Maryland 20705. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 587-589 Book REVIEWS Insect-Plant Interactions, Vol. I. Edited by E. A. Bernays. CRC Press, Boca Raton, Florida, 1989, 164 pp. Three years ago an NSF administrator told me that the study of plant-insect interac- tions—as a viable (and fundable) field ecol- ogy—was dead. The central theories had been hashed and rehashed; once innovative experimental approaches were now passé; and controversies that had gripped much of community and population ecology for two decades generated no further debate. An av- alanche of data has, in fact, appropriately tamed much of the early enthusiasm for models such as reciprocal coevolution and plant apparency. But to conclude that we have exhausted the subject or—heaven for- bid—know all that we need to about the relationships between insect herbivores and their hosts would be scientifically irrespon- sible. Though they risk obscurity under the guise of an uncommonly common title, the reviews assembled by Elizabeth Bernays in Insect-Plant Interactions illustrate the pau- city of our knowledge in several areas of this vast subject. The contributors provide a “no- frills” yet thorough treatment of five areas that are rapidly moving into the main- stream of theoretical and empirical plant- insect research. Bruce Campbell of the Western Regional Research Center, USDA-ARS, begins with a lengthy review (over 570 references) of the importance of intra- and extracellular mi- crobial symbiotes in the nutritional ecology of herbivorous insects. Throughout his dis- cussion he emphasizes how advances in re- combinant DNA techniques have replaced primitive manipulations involving anti- biotics in studies of microbial symbiotes, particularly endosymbiotes. Molecular techniques have led us beyond the notion that all endosymbiotes are yeasts and to- ward a more useful understanding of the systematics of symbiotic rickettsiae, bac- teria, plasmids, and Bochman bodies. The best evidence suggests that the endosym- biotes supplement nutritionally depauper- ate host plant resources by synthesizing amino acids, vitamins, and sterols for insect herbivores. Campbell treats extracellular symbiotes with examples from termites, roaches, beetles and flies. In addition to their role in the nutritional adaptation of insects to plants, Campbell also recognizes the po- tential significance of prokaryotic symbiotes in inducing nongenetic reproductive isola- tion between different populations of her- bivorous insects. Documentation of this phenomenon will have major implications for our understanding of herbivore specia- tion. Michael Crawley of Imperial College (U.K.) presents a stimulating synthesis of studies examining the relative impact of vertebrate vs invertebrate herbivores on the population dynamics of plants. First he rather cleverly compares the attributes of vertebrate and invertebrate herbivores, such as body size, metabolic rate, host specificity and mobility because these most influence plant recruitment and mortality. This dis- cussion establishes the hypothesis that ver- tebrates impose a measurably greater effect on plant communities than do inverte- brates. Crawley then tests this hypothesis by examining what is known about the im- pact of vertebrate and invertebrate herbi- vores on flower, fruit and seed production, seed and seedling predation, and vegetative growth rate. Finally, he compares obser- vational evidence from herbivore out- breaks, introduced herbivores, and studies of overgrazing, with data generated from experimental manipulations such as fencing exclosures and insecticide applications. Though he presents overwhelming support 588 for his hypothesis, Crawley concludes by emphasizing that the distinction he draws in this chapter 1s not between insects and vertebrates, but between small sedentary monophagous herbivores and large mobile polyphagous herbivores. Insects appear to be limited by the abundance of chemically undefended, high quality plants rather than the presence or absence of the plants them- selves. Thus, their impact on plant popu- lation dynamics and community structure is considered to be ecologically insignificant compared to that of vertebrates. Chapter 3 is a collaborative effort by J. Riemer (University of Copenhagen) and J. B. Whittaker (University of Lancaster) to draw together a disjunct literature on the effects of air pollutants on insect herbivores and their host plants. This area 1s truly in its infancy, but Riemer and Whittaker cau- tiously propose that there is a relationship between aerial pollution and changes in in- sect attacks on plants. The majority of their evidence comes from European studies which suggest that, on a regional scale, pop- ulations of insect predators and parasites generally decline, while herbivore popula- tions—particularly those of sap feeders not directly exposed to the pollutants them- selves—increase. The principal hypothesis examined in this chapter is that pollution increases plant susceptibility to herbivore attack by weakening defense systems and/ or improving resources in sap or foliage in terms of insect nutrition. The authors freely discuss inadequacies in the existing pool of data and call for controlled manipulations in future studies of this obviously important but heretofore neglected area. The fourth contribution takes a long over- due look at the extrinsic factors regulating the production of secondary metabolites in plants. This chapter alone makes the vol- ume a worthwhile purchase, for in it Peter Waterman (University of Strathclyde, Scot- land) and Simon Mole (Purdue University) critically examine the essence of nearly all plant-insect interactions: the functions and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON regulation of secondary metabolites. They begin by dismissing the simplistic views of secondary metabolites that dominated plant- insect theory in the 60’s and 70’s. It is not the ‘apparency’ of plants that determines the qualitative and quantitative presence of secondary metabolites in plant tissues, but rather a fundamental shift from metabolic products of the shikimic acid pathway (commonly employed by woody perennials) to products derived from acetate or meva- lonate (typical of more advanced herba- ceous plants). Beyond these phylogenetic re- strictions, the authors stress the importance of light intensity and soil nutrient quality in controlling the production and distribu- tion of plant metabolites. Although it 1s brief, I applaud their discussion of costs related to the production and maintenance of sec- ondary metabolites. Here they make the necessary distinction between energy costs and the cost of substrate depletion, a dis- tinction that has escaped too many ecolo- gists in the past. Of over-riding importance is the chapter’s message that insect herbi- vores are only one of many extrinsic factors influencing the production of secondary metabolites in plants. Evolutionary inter- actions between plants and insects must be interpreted with this in mind. Stephen Welter of the University of Cal- ifornia concludes the volume with a dis- cussion of the consequences of herbivory on gas exchange and thus the physiological pro- cesses of photosynthesis and respiration. His approach is to focus on the impact of var- ious types of herbivory, including that in- flicted by defoliators, mesophyll feeders, gall formers, epidermal feeders, phloem feeders, stem borers, and insects attacking roots. The strength of his chapter lies in its portrayal of our current knowledge of the subject, wanting as it 1s, rather than with patterns or predictions emerging from the data. The effect of herbivores on gas exchange in plants is yet another area about which we know too little to draw reliable conclusions. I recommend this first volume of /nsect- VOLUME 92, NUMBER 3 Plant Interactions to anyone wanting to stay abreast of current research trends in this area of ecology, and I commend the editor for her insight in compiling 5 reviews which should convincingly demonstrate to non- believers that the science of plant-insect in- teractions is alive and well. 589 Douglas W. Tallamy, Dept. of Entomol- ogy & Applied Ecclogy, University of Dela- ware, Newark, Delaware 19717-1303. PROC. ENTOMOL. SOC. WASH. 92(3), 1990, pp. 589-592 Book REVIEW A Guide to the Breeding Habits and Im- mature Stages of Diptera Cyclorrhapha. By P. Ferrar. 1987. Entomograph, Volume 8. E. J. Brill/Scandinavian Science Press, Leiden, Copenhagen. Two parts (not available separately). 907 pp. $180.00. Hard cover. ISBN 90 04 08539 4. This century, 90% of which has already passed, has seen enormous progress in the science of Dipterology. The monumental series “Die Fliegen der Palaerktischen Re- gion,” almost as old as the century and now nearly completed, has set an example for a similar, Nearctic series that has recently been started. Catalogs for all the major zoogeo- graphical regions have either been pub- lished recently or are nearing completion, and several teams of workers have launched a compilation of world catalogs for families. The recently completed “Manual of Nearc- tic Diptera” has set excellent guidelines for faunal treatments elsewhere, and there are countless smaller contributions. However, all these major contributions, plus the over- whelming majority of smaller publications on Diptera, are based on, or deal entirely with adult flies, whereas the immature stages have received relatively little attention. One exception is Hennig’s (1948-1952) series of three books that summarized all descriptive work on immature Diptera available up to the middle of this century. The main reason for this bias stems from the difficulty in- herent in locating, collecting, rearing and identifying immature dipterans. This is also the reason that most of the work published on immatures is descriptive and the entire knowledge on these stages is still very frag- mentary. Ferrar’s guide, in a way compa- rable to Hennig’s books, is a very successful attempt at summarizing the progress made during the second half of the century on a large subgroup of Diptera, the Cyclorrha- pha. Ferrar’s monograph is rather convenient- ly divided into two volumes: a) The text which covers 478 pages in addition to 8 text-figures and numerous tables, which are numbered separately in each chapter; and b) the figures that cover 431 pages including 11 pages presenting a list of the sources. The text is arranged in a simple and efficient way. After the sections entitled “Preface and Acknowledgments” and ‘Abbreviations” (five of the six abbreviations refer to the five stadia of immature stages, and the sixth— ANIC—to the Australian National Insect Collection), there is a short section (Chapter 1) devoted to techniques and dealing with the preparation and examination of eggs, larvae, and puparia. Chapter 2 1s a com- prehensive treatment of anatomical features of all immature stadia, and chapter 3 ana- lyzes, mostly in table form, the topic of breeding and other habits of Diptera Cy- 590 clorrhapha. Chapter 4 is a key to 3rd instar larvae of 74 families, out of the 87 treated in the work. The treatment of individual families, which are presented alphabetical- ly, begins on page 60. The following infor- mation is given for each family: 1. Scope and distribution; 2. Economic impor- tance—often treated under the subtitles “Harmful” and “Beneficial”; 3. Notes on literature; 4. Biology—arranged according to the unique biological and ecological fea- tures of the family; 5. Immature stages— description of all known stages. Literature is sometimes given under generic or specific titles in the Biology section. The text length for a family ranges from a few lines to more than 15 pages (e.g. Muscidae—22, Dro- sophilidae— 19), with an average of just un- der 4 pages. The second volume, with some 5000 fig- ures, is probably the largest assemblage of figures ever published for the Cyclorrhapha as a whole, as well as for many of the in- cluded families. The first 1142 figures are a selection of duplications from the rest of the figures, and are arranged in the following general order: egg; first instar larva: whole body, head skeleton, spiracles; second and third instar larvae (same internal order as in first instar); and pupa. The remaining fig- ures are arranged by family, and then again by stage and body part. Ferrar’s guide is, in my opinion, one of the major contributions to Dipterology this century. However, there are several short- comings and errors, that should be noted. Beginning with the more general ones: Al- though this work is basically a compilation (“A new summary” in Ferrar’s own words) based on over 2000 papers, the author does not explain the procedures used. As a result, there is no feasible way of knowing the ex- tent of the literature review, or which papers were not included. Indeed, many references have been omitted, either unintentionally or deliberately. This deficiency would make it very difficult to trace published descrip- tions of immatures that are not mentioned PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in this work. A second point is the lack of indication regarding the author’s criteria for either re-writing information, or presenting it unmodified. Likewise, there is no indi- cation as to how many of the described im- matures the author has seen or studied him- self. In a compilation of such magnitude it is difficult to correct mistakes of other re- searchers, or even to resolve inconsistencies or contradictions between different sets of data. It is therefore especially important not to create new confusion. One such incon- sistency 1s on page 29 where the author states that ““Myopites (Tephritidae) is unique among known Cyclorrhapha in having only 2 slits per spiracular plate in the 3rd instar,” but on page 26 (Table 2.3) he says that some Phoridae have 2 “slits” and that Nycteri- biidae (page 26) and Streblidae (page 27) have 2 “‘apertures.”’ Likewise, there is a dis- crepancy between the statements on pages 12 and 45 with regards to families for which immature stages are not known. The family Asteiidae is included in the list on page 45 only. Later, however, the egg and puparium of Asteia sabroskyi are redescribed (page 79). Additional examples of copied errors are given below when dealing with the chapter on Tephritidae. Perhaps due to his Austra- lian background, Ferrar refers repeatedly and with partiality to the Australian fauna. He explains in the preface that because of the alphabetical layout of the families, an index would be largely superfluous. I, however, believe that an index to names, and espe- cially to terminology, would have been de- sirable. The positioning of the titles of the tables below rather than above the tables is especially disturbing in tables covering more than one page, where the reader sometimes needs to turn the page in order to read the title. I have tested the key on examples from several well-studied families as well as from poorly known ones, including the unde- scribed larvae ofa species of Canacidae and those of Tethinidae. All except the Tethini- VOLUME 92, NUMBER 3 dae ran smoothly into their families. The Tethinidae dead-ended, which is gratifying for a key that does not include this family (larvae of this family have apparently never been described). In the second volume it is admittedly convenient to have the separate batch of initial 1142 figures at hand, so that a selec- tion of figures from across the Cyclorrhapha can be viewed easily. However, I believe it would be more practical to omit this luxury of 80 duplicate pages in favor of a propor- tionate reduction in the price the book, which is very high. The list of figure sources on page 897 does give the relevant refer- ences. However, is this a legal substitute for copyrights and receiving permission to copy figures? My final comment about the figures in volume 2 is with regards to their quality. The mixed styles of the figures is less than aesthetic, but is understandable and ac- ceptable. The great majority of the figures are reasonable reproductions, and some of them are, surprisingly, better than the orig- inals. However, the quality of certain figures is extremely low, even to the point of being useless (the figures on page 705 and figure 88.31 are just a few examples). More specific comments: ““Techniques” (Chapter 1). This chapter is generally in- structive. The author, however, refers only very briefly to the use of scanning electron microscopy (SEM) for larvae. Although this is a-relatively new approach, it is gaining momentum quickly and deserves more than the 10 short lines given to it in this work. It is perhaps appropriate to add here a quick method for studying larvae that I have found satisfactory in most cases: use (with caution) hot 10% KOH to macerate the larva, clean in water, and mount in a drop of glycerin. T have not taken special efforts to discover errors, but the one on p. 22, where Figure 6.252 was cited was prominent. Such a fig- ure does not exist, and the author probably meant to say Figure 14.6. Two good practices, on the other hand, deserve mentioning: In volume one the au- 591 thor is consistent in giving in parentheses the family to which every mentioned species belongs. In volume 2 the figure legends are located immediately below their respective figures. These two practices, which are not very common elsewhere, facilitate the use of the work. In relating to the individual chapters dealing with the various families, I prefer to restrict myself to the chapter dealing with Tephritidae, a family which I myself have been studying for some time. I have only a few comments: The examples of beneficial species are enlightening, but I would also have added a further example, namely the genus Urophora, which has been the focus of extensive research in recent years. Several species of this genus have been introduced from Europe to North America, some suc- cessfully, to combat knapweed and related weeds there. The first paragraph of column 2 on page 388 deals with the generalization that all Aciurinae whose hosts are known breed in plants of the families Labiatae, Acanthaceae or Verbenaceae [within Tephritidae this as- sociation is practically unique (A.F.)]. Al- though I am still a strong supporter of this statement, this particular paragraph is er- roneous and misleading, partly due to my own earlier mistake. The genus I meant to record in 1979 is Perirhithrum, not Trirhith- romyia. Perirhithrum was recently trans- ferred from the Schistopterinae to the Te- phritinae but, as this genus is now considered to be in the Aciurinae (Freidberg, unpub- lished), it supports the generalization about hosts of this group rather than being an ex- ception. Likewise, Aciurina is a tephritine (and not an aciurine as might be alluded from its name) and therefore is not an ex- ception to the above generalization, despite the fact that its hosts are Compositae. The only known exception to the above gener- alization is Eutreta xanthochaeta (Tephri- tinae), a species that induces the formation of stem galls on Lantana camara (Verben- aceae). 592 The only described immatures of a true Chetostoma are those of C. continuans which are frugivorous on Lonicera spp. (Caprifoli- aceae; Kandybina, 1966: 679). Chetostoma completum (p. 390) is actually a Chaetos- tomella (based on my unpublished obser- vations on specimens from the type series), a genus attacking various Compositae. Fi- nally, some names were misspelled: The correct spelling is Carpomya and not Car- pomyia (p. 389); and heracleii (a species of Euleia) and not heraclei (p. 390). Despite these shortcomings, this chapter makes in- teresting reading and is an important ref- erence in the study of Tephritidae. Iam sure that other researchers will reach the same conclusion with regards to the chapters that deal with families of their own particular interest. In summary, this excellent work is a must PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for any laboratory studying Diptera or im- matures and ecology of insects. Its appear- ance is warmly welcomed. As a result of its presence, the necessity for similar sets for the rest of the Diptera became much more prominent. LITERATURE CITED Freidberg, A. 1979. The Afrotropical species assigned to Terellia R.D. (Diptera: Tephritidae). J. Wash. Acad. Sci. 69: 164-174. Hennig, W. 1948-1952. Die Larvenformen der Dip- teren. 1.-3. Teil. Berlin, Akademie Verlag. Kandybina, M. N. 1966. Contribution to the study of fruit flies (Tephritidae, Diptera) in the far east of the USSR. Ent. Rev., Wash. 45: 383-388 [orig. in Ent. Obozr. 45: 677-687]. Amnon Freidberg, Department of Zool- ogy, The George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Tsrael. PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Gynipid Gals ofithe/Easteras United States) by LewisiH. Weld == = Se GynipidsGalls ofthersouthwestnoyseewis tle nW Cl Css ene ne TBYaK HOY jORENO STR Cay CORN YG | CNS a I rt Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman eee Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. A Short History of the Entomological Society of Washington, by Ashley B. Gurney. Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. S fey Ska lesen ete & eereenain eee eee ee SS ee Pole Soe Lis eek tO Te ee Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. Steyskal . 10. paluis lly; MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939... A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. 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All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) LUYKX, PETER, DAVID A. NICKLE, and BRIAN I. CROTHER.—A morphological, allo- zymic, and karyotypic assessment of the phylogeny of some lower termites (Isoptera: Kalotermitidae) MADDOX, D. M., A. MAYFIELD, and C. E. TURNER.—Host specificity of Chaetorellia australis (Diptera: Tephritidae) for biological control of yellow starthistle (Centaurea solstitialis, Asteraceae) McDANIEL, B. and A. BOE.—A new host record for Eurytomocharis eragrostidis Howard (Chalcidoidea: Eurytomidae) infesting Eragrostis tef in South Dakota NEUNZIG, H. H.—A new species of Dioryctria (Pyralidae: Phycitinae) from Mexico OLMSTEAD, K. L. and T. K. WOOD.—Altitudinal patterns in species richness of Neotropical treehoppers (Homoptera: Membracidae): the role of ants PINTO, JOHN D.—The genus Xiphogramma, its occurrence in North America, and remarks on closely related genera (Hymenoptera: Trichogrammatidae) SHARKEY, MICHAEL J.—A revision of Zacremnops Sharkey and Wharton (Hymenoptera: Braconidae: Agathidinae) SPANGLER, PAUL J.—A new species of water scavenger beetle, Guyanobius simmonsorum, from Brazil (Coleoptera: Hydrophilidae) TOGASHI, ICHIJI.—A new Darjilingia (Symphyta: Tenthredinidae) from Taiwan VANDENBERG, NATALIA J.—First North American records for Harmonia quadripunctata (Pontopiddian) (Coleoptera: Coccinellidae); a lady beetle native to the Palaearctic VEGA, FERNANDO E. and PEDRO BARBOSA.—Gonatopus bartletti Olmi [Hymenoptera: Dryinidae] in México: a previously unreported parasitoid of the corn leafhopper Dalbulus maidis (Delong & Wolcott) and the Mexican corn leafhopper Dalbulus elimatus (Ball) [Homoptera: Cicadellidae] WIRTH, WILLIS W. and FRANCIS E. GILES. —New species and records of predaceous midges from Fiji (Diptera: Ceratopogonidae) NOTES DYTE, C. E.—The type locality of Sciapus pressipes Parent (Diptera: Dolichopodidae) WHITE, GEOFFREY B. and RALPH E. WEBB.—Obituary. Anne Marie Wieber 1958-1989 BOOK REVIEWS FREIDBERG, AMNON.—A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha TALLAMY, DOUGLAS W.—Jnsect-Plant Interactions | VOL. 92 OCTOBER 1990 |. NO. 4 4 Kd (ISSN 0013-8797) PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED QUARTERLY CONTENTS BATRA, S. W. T.—Bionomics of Evylaeus comagenensis (Knerer and Atwood) (Halictidae), a facultatively polygynous, univoltine, boreal halictine bee ............................. 725 DAVIS, DONALD R.—Three new species of Acrolophus from the southeastern United States with remarks on the status of the family Acrolophidae (Lepidoptera: Tineoidea) ........ 694 DAVIS, DONALD R. and JORGE E. PENA — Biology and morphology of the banana moth, Opogona sacchari (Bojer), and its introduction into Florida (Lepidoptera: Tineidae) ..... 593 EISENBERG, R. M. and L.E. HURD—Egg dispersion in two species of praying mantids (Man- TOMEA=MIVian tl Gace heey cee ace ee ere Ae eee A ME Sl ones Aa ee te ala oe 808 EPSTEIN, MARC E. and SCOTT E. MILLER—Systematics of the West Indian moth genus Heuretes Grote and Robinson (Lepidoptera: Limacodidae) ........................... 705 GAGNE, RAYMOND J. and GWENDOLYN L. WARING— The Asphondylia (Cecidomyiidae: Diptera) of creosote bush (Larrea tridentata) in North America ....................... 649 GOEDEN, RICHARD D. and DAVID HEADRICK — Notes on the biology and immature stages of Stenopa affinis Quisenberry (Diptera: Tephritidae) ..........................-2---- 641 GRACE, J. KENNETH— Effect of antioxidants on eastern subterranean termite (Isoptera: Rhi- moter tidae) Orientation tO mUnPAlextTaCt Spies cay caucicstec avec eo acta se eae wer ATS, HALBERT, SUSAN E. and KEITH S. PIKE—An Asian elm aphid (Homoptera: Aphididae) ME WEOVN OLAV AMEDICANS antes a7 ieee Me Tae ee ik ee els Cee ee a code Reno's + a0 cise salah aedees HALSTEAD, JEFFREY A.—Revision of Hockeria Walker in the Nearctic region with descrip- tions of males and five new species (Hymenoptera: Chalcididae) ...................... 619 HENRY, THOMAS J. and DANIEL J. HILBURN—An annotated list of the true bugs (Het- CLODICIA) OMBCLOTUC Ae ee eee ey ee Se ee oe 2s dcyeee 4a ote en Ie bes 675 LINLEY, JOHN R.—Scanning electron microscopy of the eggs of Aedes vexans and Aedes BAILII pLeTAaGUIICIGAl) Petpet eee ee sense hitene cite wy sacs oleae ale. seseys 685 MATHIS, WAYNE N.—A revision of the shore-fly genus Diphuia Cresson (Diptera: Ephy- (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1990 JEFFREY R. ALDRICH, President NORMAN E. WooDLey, Treasurer Davip R. SmitH, President-Elect Gary STECK, Program Chairman RICHARD G. RosBINns, Recording Secretary GEOFFREY B. WHITE, Membership Chairman Ho us B. WILLIAMS, Corresponding Secretary F. CHRISTIAN THOMPSON, Past President JAMES B. STRIBLING, Custodian RoBERT D. Gorpbon, Editor THOMAS J. HENRY, Associate Editor Publications Committee DONALD R. DAvis WAYNE N. MATHIS GEORGE C. STEYSKAL F. CHRISTIAN THOMPSON Honorary President Curtis W. SABROSKY Honorary Members LoulIsE M. RUSSELL ALAN STONE THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $20.00 (U.S. currency) of which $18.00 is for a subscription to the Proceedings of the Entomological Society of Washington for one year. PROCEEDINGS.-— The Proceedings are published quarterly beginning in January by The Entomological Society of Washington, % Department of Entomology, NHB-168, Smithsonian Institution, Washington, D.C. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $50.00 per year, domestic, and $60.00 per year, foreign (U.S. currency), payable in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE P. 318 OF THE APRIL, 1989 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. Editor: Robert D. Gordon, Systematic Entomology Laboratory, ARS, % Department of Entomology, Smith- sonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: T. Henry, Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 16 October 1990 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. ENTOMOL. SOC. WASH 92(4), 1990, pp. 593-618 BIOLOGY AND MORPHOLOGY OF THE BANANA MOTH, OPOGONA SACCHARI (BOJER), AND ITS INTRODUCTION INTO FLORIDA (LEPIDOPTERA: TINEIDAE) DONALD R. DAVIS AND JORGE E. PENA (DRD) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (JEP) Institute of Food and Agricultural Sciences, University of Florida, Homestead, Florida 33031. Abstract. —The general distribution and recent introduction into Florida of the banana moth, Opogona sacchari (Bojer) is reviewed. Currently, the principal damage caused by this species in Florida consists of larval stem boring in certain nursery stock and orna- mental palms in particular. The biology of the species is summarized and all stages of the insect are described, supplemented by numerous illustrations. Key Words: mature stages The banana moth, Opogona sacchari (Bojer), is known from several tropical-sub- tropical humid regions around the world but has yet to be reported from the Indo- Australian region. Originally reported from the Mascarene Islands in the Indian Ocean (Bojer 1856, Walker 1863, Butler 1876), the species was later discovered in Africa (Vari and Kroon 1986) as well as islands near the African continent (Walker 1875, Durrant 1925) and Europe (Ciampolini 1973, D’ Aguilar and Martinez 1982, Mourikis and Vassilaina-Alexopoulon 1983). More re- cently the species has spread to South America (Cintra 1975, Giannoti et al. 1977) and the West Indies (Alam 1984). The ear- liest evidence we have of this species in the New World is represented by adult speci- mens in the Smithsonian Institution (USNM) that were collected in 1970 from Aragua, Venezuela on potato. Specimens received for identification from B. Kumashiro of the Hawaiian Department of Agriculture just prior to publication in- dicate that O. sacchari is now established in Hawaii. The species has been reared from Lepidoptera, Tineidae, Opogona sacchari, banana moth, moth biology, im- rotting coconut tree tops (central whorl of leaves) from Kaneohe, Oahu and from an unidentified palm at Kohala, Hawaii. Within the last few years O. sacchari has become established on various nursery stock in southern Florida, particularly in Dade and Palm Beach Counties (Heppner, Pena, and Glenn 1987). Nursery stock particularly affected in Florida include corn plant or casse (Dracena fragrans (L.) Ker-Gaus, variety massangeana) and bamboo palms (Cha- maedorea sp.) as well as Hawaiian good luck plant (Cordyline terminalis (L.) Kunth) and aralias (Polyscias sp.). Although sugar cane is a major host, O. sacchari has not yet been reported on that plant in the United States. The appearance of this new pest in the United States has necessitated a careful ex- amination of all developmental stages and a report of its biology pertinent to its current significance as a pest of nursery stock in Florida. Opogona sacchari (Bojer) Alucita sacchari Bojer 1856: 21, pl. 5, figs. 1-10. 594 Opogona sacchari (Bojer).— Vinson, 1938: 56 (synonym of Opogona subcervinel- la).—Viette, 1957: 145; 1958: 4.—Ciam- polini, 1973: 221.—Cintra, 1975: 223.— Giannotiet al., 1977: 209.—Declercq and Van Luchene, 1977: 499.—Zimmerman, 1978: 386.—Cintra et al., 1978: 3.—Pi- gatii, 1978: 21.—Pigatiietal., 1979: 61.— Veenenbos, 1981: 235.—D’Aguilar and Martinez, 1982: 28.—Rotundo and Tremlay, 1982: 123.—Suplicy and Sam- paio, 1982: 174.—Bennett and Alam, 1985: 41.—Heppner et al., 1987: 1. Tinea subcervinella Walker, 1863: 477. Opogona subcervinella (Walker). —Wal- singham, 1907: 713; 1919: 259.—Meyr- ick, 1930: 321.—Vinson, 1938: 56.— Viette, 1951: 339; 1957: 145 (synonym of Opogona sacchari); 1958: 4.—Paulian and Viette, 1955: 147.—Box, 1953: 34.— Davis, 1978: 14; 1984: 22.—Vari and Kroon, 1986: 84, 156. Hieroxestis subcervinella (Walker).—Mey- rick, 1910: 375; 1911: 298.—Cockerell, 1923: 247.—Meyrick, 1924: 556.—Dur- rant, 1925: 12.—Oldham, 1928: 147.— Rebel, 1939: 63; 1949: 56.—Jannone, 1966: 24. Gelechia sanctaehelenae Walker, 1875: 192.—Durrant, 1925: 12 (synonym of Hi- eroxestis subcervinella). Euplocamus sanctaehelenae (Walker).— Wollaston, 1879: 417. Hieroxestis sanctaehelenae (Walker).— Durrant, 1923: xvii. Gelechia ligniferella Walker, 1875: 192.— Durrant, 1925: 12 (synonym of Hierox- estis subcervinella). Laverna plumipes Butler, 1876: 409.— Wal- singham, 1907: 713 (synonym of Opo- gona subcervinella), 1919: 259. Adult (Figs. 1, 2).—Length of forewing: 6, 7.3-11.2 mm; 2, 9-12.5 mm. Moderately large, generally dark grayish brown moths with a small, black subapical spot on fore- wing near apex of discal cell and a similar one at basal third on anal fold; male slightly PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON paler in color with faint longitudinal streaks of light brown or buff. Head: Vestiture smooth except for a pair of bilateral tufts of erect, pale brown to cream, piliform scales between antennal bases; scales of frons broad, cream colored; vertex covered by a dense row of broad cream scales that curve over frons; caudal portion of vertex with broad, grayish brown scales; occiput light brown with a scattered arch or median patch of dark fuscous scales. Antenna 103-125 segmented, approxi- mately 0.8 the length of forewing; scape cream dorsally, with scattered fuscous scales ventrally; flagellum uniformly covered with small but moderately broad, cream to buff scales arranged in a single row, completely encircling each flagellomere (Fig. 12); sen- silla chaetica (with spiral grooves) and trich- odea (longitudinal grooves) relatively dense and randomly scattered over each flagello- mere; a few pair of sensilla coeloconica lo- cated near distal margin (Figs. 13, 14). Pil- ifers moderately developed with setae nearly meeting at midline (Fig. 9). Labrum densely covered with microtrichia. Mandible ves- tigial but exceeding pilifer setae in length. Maxillary palpus 5-segmented, elongate, ex- ceeding length of relatively short haustel- lum; dorsal and lateral surfaces densely cov- ered with cream scales; venter naked with dense sensilla; apex of fifth segment with a slender basiconic sensillum and another slightly smaller one at subapex (Figs. 15, 16). Haustellum with a series of shallow plates over basal half; largest plates with a pair of short sensilla basiconica (Fig. 10). Labial palpus upcurved, smoothly covered with cream scales except for a lateral, sub- apical series of 3-4 cream bristles and a ven- tral subapical tuft of 6-8 bristles; third seg- ment with an elongate, narrow sensory pit located just beyond middle (Figs. 17, 18). Thorax: Pronotum light brown with heavy to sparse scattering of fuscous scales and usually with a small to large median patch of fuscous on anterior margin and on tegula. Venter uniformly shiny cream. Forewing VOLUME 92, NUMBER 4 595 Figs. 1-4. Opogona sacchari. 1, Adult male, length of forewing 9.5 mm. 2, Adult female, length of forewing 12.5 mm. 3, Larvae, maximum length 30 mm. 4, Cocoon with pupal exuvium, cocoon length 12.5 mm. light brown in female with an almost equal amount of scattered dark brown scales; male with dark brown scales more concentrated into longitudinal streaks, particularly evi- dent along upper and lower margins of dis- cal cell; a small, dark fuscous subapical spot near apex of cell and a more elongate fus- cous spot along anal fold at basal third in both sexes; male retinaculum a relatively elongate flap from ventral subcostal margin, with apical margin rolled under (Figs. 24, 25). Hindwing light yellowish brown with slender scales in male, more shiny gray and with siightly broader scales in female; male with elongate hair pencil from dorsal base (Figs. 28-30); ultrastructure of pencil sex scales as illustrated (Figs. 31-33); male fren- ulum a single stout bristle (Fig. 26); female with usually five stout frenular bristles (Figs. 27). Foreleg mostly cream; coxa with a small amount of grayish fuscous suffusion at base; dorsal surfaces of tibia and tarsus heavily suffused with grayish fuscous; normal pec- tinated epiphysis present (Figs. 19, 20). Pre- tarsus of all legs normal, with relatively broad unguitractor plate bearing 7-10 ranks of scutes in a single transverse row (Figs. 21-23). Midleg uniformly cream except for slight grayish fuscous suffusion over dorsal surface of tarsus. Hindleg similar to midleg in color except with less grayish fuscous on tarsus and with dense, elongate piliform se- tae from tibia. Abdomen: Terga mostly grayish brown with caudal margins and pleura buff to cream colored; venter of female uniformly cream, 596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. Damage to Dracaena fragrans by Opogona sacchari larvae (see arrows). 5, Healthy stem. 6-8, Typical feeding damage to Dracaena fragrans canes. VOLUME 92, NUMBER 4 597 Figs. 9-14. Opogona sacchari, adult morphology. 9, Labrum, anterior view (100 um). 10, Base of haustellum (38 um). 11, Haustellum, food canal (15 um). 12, Antenna, scale pattern (50 um). 13, Antenna, sensilla (60 um). 14, Sensillum coeloconicum of antenna (4.3 um). (Scale lengths in parenthesis; bar scale for all photographs = Fig. 9.) 598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 15-20. Opogona sacchari, adult morphology. 15, Maxillary palpus, apical (fifth) segment (100 um). 16, Apex of Fig. 15 (23.1 um). 17, Labial palups, apical (third) segment (176 um). 18, Detail of sensory pit in Fig. 17 (see arrow) (25 um). 19, Epiphysis (120 um). 20, Detail of pecten on epiphysis (27 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 15.) VOLUME 92, NUMBER 4 Figs. 21-26. Opogona sacchari, adult morphology. 21, Pretarsus of midleg, lateral view (20 um). 22, Ventral view (50 um). 23, Unguitractor plate of pretarsus (7.5 um). 24, Subcostal retinaculum of male forewing (0.27 mm). 25, Distal-lateral view of Fig. 24 (176 um). 26, Male frenulum (38 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 21.) 600 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i HAWS Wil] tH 4 y ) WWI / BM) Figs. 27-32. Opogona sacchari, wing morphology. 27, Female frenulum (250 um). 28, Hair pencil of male hindwing, dorsal wing (1 mm). 29, Base of male hair pencil (176 um). 30, Raised scale sockets of hair pencil (30 um). 31, Surface ultrastructure of a single hair pencil scale (3.8 wm). 32, Detail of Fig. 31 (1.2 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 27.) VOLUME 92, NUMBER 4 60! 7 aE > SoS 5 ne vg & ; ™~ ais Any id ~~? % ‘ Se os an tok AG: FUT 4 s = Figs. 33-38. Opogona sacchari, adult, egg, and larval morphology. 33, Cross section of hair pencil scale (Fig. 31) (3.8 um). 34, Egg, lateral view (150 um). 35, Micropyle (136 um). 36, Detail of Fig. 35 (75 um). 37, Detail of central disk (Fig. 36) (17.6 um). 38, Head of larva, anterior view (60 mm). (Scale lengths in parentheses; bar scale for all photographs = Fig. 33.) 002 grayish brown in male; both sexes with a lower lateral series of five dark fuscous spots on A3-7. Male genitalia (Figs. 79-82): Uncus deep- ly divided into two large widely separated lobes arising beneath tegumen and bearing numerous stout elongate setae on their me- sal surfaces. Tegumen a relatively broad ring dorsally. Vinculum tapering to a broad, rel- atively truncate saccus. Valva prominently divided into a large, elongate, rounded api- cal lobe and a much smaller, acute cucullar lobe. Aedoeagus a relatively small, straight, slender tube without cornuti; phallobase much larger and greatly inflated. Female genitalia (Figs. 83, 84): Tertiary apophyses present in AlQ. Genital plate moderately divided into a pair of rounded lobes. Corpus bursae with a single large sag- itate signum bearing elongate anterior arms. Egg (Figs. 34-37).— Length, 0.5-0.55 mm; diameter, 0.38 mm; oval in shape, round in cross section; color light yellow at ovipo- sition, gradually becoming yellowish brown prior to eclosion. Surface irregularly pitted. Micropyle consisting of a single, centrally positioned opening with small radiating grooves forming an enlarged, reticulate pat- tern of low ridges over entire end of egg; reticulations mostly 5-6 sided. Larva (Figs. 3, 38-66).— Length of largest larva 30 mm, maximum diameter 3 mm. Body generally white with dark brown plates and pinacula. Head: Maximum width 2.5 mm. Color reddish brown, becoming darker anteriorly and over frons. Chaetotaxy as illustrated. Stemmata rudimentary, consisting of a pair of widely separated, clear lenses, probably corresponding to stemmata | and 5; the most anterior (5) located ventral to antennal socket; the most posterior (1) midway be- tween A3 and S2. Mandible with five cusps. Spinneret long and slender with a minute orifice. Labial palpus 2-segmented, elongate and slender, equalling or slightly exceeding length of spinneret; apical seta one-third the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON length of entire palpus. Apex of mentum with a pair of minute secondary labial setae. Thorax: Pronotum and spiracular plate dark reddish brown, lighter in color along margins; spiracle together with L setae on same plate. Pinacula over entire body rel- atively large, dark brown and very distinct on whitish integument. Meso- and meta- thorax with L2 arising on a separate pinacu- la from L1 and 3. MSD1 and 2 of similar length and reduced. Legs well developed; tarsal claw elongate; basal lobe actually bi- lobed and with a minute conical seta from inner angle (Figs. 51, 52). Abdomen: Whitish in color with brown- ish pinacula. L2 on separate pinacula from spiracle. Prolegs well developed on A3-6 and 10; crochets A3—6 uniordinal, uniserial, and arranged in a complete ellipse com- posed of approximately 43-45 hooks; a scattered band of much smaller, numerous spines encircling apex of planta (Fig. 53); crochets on Al0 with 20-22 hooks and a dense, scattered band of much smaller spines along anterior edge of planta (Figs. 55, 56). Pupa (Figs. 67, 68).—Length, 9-12.8 mm; maximum width, 2—3.5 mm. Color usually light brown ventrally and dark reddish brown dorsally; wing cases becoming darker with maturity; frontal process, cremaster, and adjacent areas extremely dark, often black. Head with frontal process (cocoon cutter) moderately developed, with a broad, triangular apex (Figs. 68, 78); labrum with a pair of lateral setae. Anterolateral margins of mesonotum with a pair of perforated bands composed of minute, raised slit open- ings (Figs. 69-71). Forewings extending to middle of A5. Hindlegs to A6. A simple, anterior row of short, stout, dorsal spines present on segments A4-8; tabulation of spines as follow: A4 = 56-70, AS = 53-72, A6 = 57-71, A7 = 44-50, A8 = 8-18. Last pair (A8) of spiracles on raised, swollen bas- es (Figs. 74-76). Cremaster consists of a large pair of stout hooks arising dorsally from VOLUME 92, NUMBER 4 603 Figs. 39-44. Opogona sacchari, larval morphology. 39, Head, dorsal view (0.6 mm). 40, Labrum, dorsal view (176 um). 41, Head, lateral view (0.43 um). 42, Stemmal area (136 um). 43, Head, ventral view (0.5 mm). 44, Maxilla and labium (150 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 39.) 004 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 45-50. Opogona sacchari, larval morphology. 45, Maxilla (60 um). 46, Apex of maxillary palpus (6 um). 47, Apex of maxilla (5 um). 48, Apex of antenna (30 um). 49, Apex of antenna (27 um). 50, Ventral view of pro- and mesothorax (0.6 mm). (Scale lengths in parentheses; bar scale for all photographs = Fig. 45.) VOLUME 92, NUMBER 4 605 WH Wi eet ma aii) Figs. 51-56. Opogona sacchari, larval morphology. 51, Pretarsal claw of prothorax (50 um). 52, Detail of axial lobes and seta of Fig. 51 (6 wm). 53, Proleg of A3, | anterior, ~ meson (136 wm). 54, Caudal view last abdominal segment, A10 (0.5 mm). 55, Ventral view, A9, 10 (0.5 mm). 56, Anal proleg, A10 (150 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 51.) 606 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 57-64. Opogona sacchari, larval chaetotaxy. 57, Body segments T1-2, Al, 6, 8-9. 58, Head, dorsal view (0.5 mm). 59, Ventral view. 60, Segments A8-10, dorsal view. 61, Head, lateral view. 62, Mandible. 63, Labrum, dorsal view (0.25 mm). 64, Ventral view. (Scale lengths in parentheses.) VOLUME 92, NUMBER 4 607 Figs. 65-70. Opogona sacchari, larval and pupal morphology. 65, Larval segments A8-10, dorsal view (0.6 mm). 66, Lateral view of Fig. 65 (0.6 mm). 67, Pupa, lateral view of head (0.6 mm). 68, Ventral view of Fig. 67 (0.6 mm). 69, Dorsal view of Fig. 67, perforated band (see arrow) (0.6 mm). 70, Detail of dorsal perforated band (136 um). (Scale lengths in parentheses; bar scale for all photographs = Fig. 65.) 608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON HM Figs. 71-76. Opogona sacchari, pupal morphology. 71, Detail of openings in perforated band (see Fig. 69) (15 um). 72, Dorsum of A4—5 (0.75 mm). 73, Dorsal spines of A4 (100 um). 74, Dorsal view of A7—10 (0.5 mm). 75, Caudal view of A8-10 (0.5 mm). 76, Lateral view of A8-10 (0.5 mm). (Scale lengths in parentheses; bar scale for all photographs = Fig. 71.) VOLUME 92, NUMBER 4 Figs. 77, 78. Opogona sacchari, pupa. 77, Ventral view (2 mm). 78, Dorsal view. (Scale lengths in paren- theses.) A10 (Figs. 74-76); a much smaller pair of conical tubercules also present ventrally. Pupation occurs in a cocoon (Fig. 4) of white silk usually covered with dark frass and plant debris, 14-18 mm long, 3-4 mm in diameter. Normally, the cocoon is con- structed somewhere in or near the feeding site. Types.—Syntypes 6, 2, deposition un- known (sacchari); Holotype, BMNH (sub- cervinella); syntypes, 4, 9, BMNH (sanctae- helenae), holotype, sex unstated, BMNH (ligniferella), syntype(s)?, sex and number of specimens unstated, BMNH (p/umipes). Type localities.—Mascarene Islands: Mauritius (sacchari, subcervinella), St. Hel- ena (sanctaehelenae, ligniferella); Mascar- ene Islands: Rodriguez (p/umipes). Hosts.—46 plant hosts reported (see Ta- ble 1). Distribution (maps 1, 2).—A widely rang- ing, partly pantropical species, early re- ported from several circum-African islands (Canary Islands, St. Helena, Mauritius, 609 Rodriquez and the Seychelles), as well as South Africa; frequently intercepted in sev- eral European countries (Belgium, France, Great Britain, Greece, and the Nether- lands). More recently, O. sacchari has been introduced into South America and the West Indies, including Bermuda, and is now es- tablished in nurseries over much of south- ern Florida. Discussion.—By its relatively large size and predominantly brown color, Opogona sacchari 1s easily distinguished from nearly all other members of this large complex. Its affinities are with O. omoscopa (Meyrick) as suggested by their similar male genitalia and shared presence of a dorsal hair pencil in the hindwings of the male. Present evidence suggests that this complex of obviously re- lated species warrants recognition under and resurrection of the currently synonymized genus Hieroxestis. Hopefully this and sim- ilar generic uncertainties can be resolved following a revision of the large cosmopol- itan genus Opogona. The larva of O. sac- chari may be recognized by the presence of two stemmata, separation of the spiracle from the pinaculum bearing L2 on the first eight abdominal segments, by the large number of crochets (A3-6 = 43-45, Al0 = 20-22), and by complete encirclement of the abdominal planta by a band of small, secondary spines. In O. omoscopa, a pan- tropical species recently introduced into California (Davis 1978), only one anterior stemma has been observed, the first eight abdominal spiracles are united with L2 on acommon pinaculum, the crochets are few- er in number, and the abdominal planta A3- 6 have spines only along the anterior mar- gin. The pupae of these two species are also similar but may be distinguished by the raised spiracle on A8 and larger cremaster spines of O. sacchari. A peculiar structure of unknown function in the pupa of this species deserves further comment. It consists of a bilateral pair of perforated bands curving around the an- terolateral margins of the mesonotum (Figs. 610 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 79-84. Adult genitalia. 79, Male, ventral view (0.25 mm). 80, Lateral view. 81, Lateral view of valva. 82, Aedoeagus, lateral view. 83, Female, ventral view (1 mm). 84, Detail of signum in Fig. 83. (Scale lengths in parentheses.) 69-71). The bands are narrow and elongate and under high magnification can be ob- served to comprise raised, slitlike openings. The senior author has not noted this in other species, perhaps because the bands were simply overlooked. Possibly their function is merely to weaken the cuticle, thereby fa- cilitating rupture of the pupal shell during eclosion. The band was observed to have been expanded or further separated in most pupal exuviae examined, although the ma- jor breaks in this region of the pupal shell during ecdysis did not occur through the perforated bands but were located anterior to the pronotum and down the mid-dorsal line of the pro- and mesonotum. VOLUME 92, NUMBER 4 Table 1. Plant hosts of Opogona sacchari. Plant Species Agavaceae Cordyline terminalis (L.) Kunth Dracaena fragrans (L.) Dracaena fragrans (L.) Ker-Gaus, “var. massangeana”’ Dracaena marginata Lam. Dracaena reflexa Lam. Yucca elephantipes Regel Yucca sp. Araceae Colocasia esculenta Schott. Philodendron scandens Lindl. Araliaceae Polyscias fruticosa (L.) Harms Polyscias fruticosa (L.) Harms, “elegans” Asteraceae Dahlia sp. Bromeliaceae Aechmea fasciata (Lindl.) Baker Aechmea fasciata (Lindl.) Baker ‘‘Variegata” Guzmania lingulata var. x magnifica [Hort.] Nidularium tricolor [species name unknown, possibly = Neoregelea ‘Perfecta Tricolor,’ a cultivar] Caricaceae Carica papaya L. Convolvulaceae Ipomoea batatas Lam. Cycadaceae Cycas revoluta Thunberg Dioscoreaceae Dioscorea sp. Gesneriaceae Gloxinia sp. Saintpaulia sp. Iridaceae Gladiolus sp. Leguminosae Albizia julibrissin Durazz. Enterolobium sp. Erythrina variegata L. Liliaceae Sansevieria laurantii Wildem. Sansevieria trifasciata laurentii Wildem. Malvaceae Hibiscus sp. Reference Heppner et al. Declercq and Van Luchene 1977 Heppner et al. Heppner et al. Heppner et al. Heppner et al. Heppner et al. Cintra 1975 Siiss 1974 Heppner et al. Heppner et al. Cintra 1975 Siiss 1974 Siiss 1974 Siiss 1974 Siiss 1974 Viette 1951 1987 1987 1987 1987 1987 1987 1987 1987 USNM (new record) Heppner et al. Heppner et al. Siiss 1974 Siiss 1974 Cintra 1975 Heppner et al. Heppner et al. 1987 1987 1987 1987 FSCA (new record) Siiss 1974 611 Country United States Belgium United States United States United States United States United States Brazil Italy United States United States Brazil Italy Italy Italy Italy Madagascar Peru United States United States Italy Italy Brazil United States United States United States Italy Declercq and Van Luchene 1977 Belgium Moreton 1974 Great Bntain 612 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Plant Species Reference Country Marantaceae Maranta leuconeura massangeana Schum. Siiss 1974 Italy Stromanthe sanguinea Sonder Siiss 1974 Italy Moraceae Ficus elastica (H. A. Siebrecht) Musaceae Musa cavendishii Paxt Musa paradisiaca L. Musa sapientum L. Strelitzia sp. Orchidaceae “Orchids” Palmae Arecastrum sp. Bactris [= Guilielma] gasipaes HBK Chamaedorea elegans Mart. Chamaedorea erumpens H. E. Moore Chamaedorea seifrizii Burret Poaceae “Bamboo” Saccharum officinarum L. Zea mays L. Solanaceae Capsicum sp. Solanum melongena L. var. esculentum Nees Solanum tuberosum L. Verbenaceae Clerodendrum sp. Moreton 1974 Great Britain Oldham 1928 Canary Islands Heppner et al. 1987 United States Oldham 1928 Canary Islands Zandvoort 1972 Netherlands Heppner et al. 1987 United States United States United States United States United States United States Heppner et al. 1987 Heppner et al. 1987 Heppner et al. 1987 Heppner et al. 1987 Heppner et al. 1987 Oldham 1928 Canary Islands Bojer 1856 Mauritius Oldham 1928 Canary Islands Siiss 1974 Italy Siiss 1974 Italy Oldham 1928 Canary Islands Heppner et al. 1987 United States BIOLOGY The larvae of most species of Opogona for which we have information are detritus feeders and rarely feed on living plant tissue. Opogona sacchari thus departs from the norm and can be a serious pest of banana, maize, potato, sweet potato, sugar cane, and certain greenhouse crops (Alam 1984, Dur- rant 1925, Oldham 1928, Siiss 1975, Vee- nenbos 1981). A total of seven instars are indicated, based on head capsule measure- ments of larvae reared on artificial diet (Ta- ble 2). All measurements were based on pooled data for both sexes. The discreteness of the last two instars as interpreted from head capsule measurements presumably would be better had males only or females only been used. The SD values do not over- lap as it is, so likely the conclusion reached as to instar number is correct. More precise data, however, might have shown indica- tion of variation in instar numbers, which seems likely in a generalist feeder of this sort. Oldham (1928) observed that O. sacchari larvae feed on nearly all parts of the banana plant except the roots and leaf blades. Lar- vae accepted leaves as a food in rearings but normally avoided this part of the plant in nature. The most serious damage occurred in the banana inflorescence. Larvae seldom feed exposed but burrow into the substra- 613 VOLUME 92, NUMBER 4 og} —— | | “uojuney © a | zenBypoy * a - aeosedppew ‘seeyokes *& | BOY pe Ant 81 a | { (oineg ows) 7 Weg x Bulepe se10zy oe “DY IIDS DUOSOdG JO 9dUILINIIO payoday “| dew =i Re CO 06 OB! p \.F | En Vis | Sy | | NOILD3PONd HOLWOWIW | | sunianon cope _otpe_ome con E Ses nme! | YOLVNDI NO 3S i | | | | T st — + Se 614 a Map 2. Distribution of Opogona sacchari in Flor- ida (after Heppner et al. 1987). tum. Their presence is usually indicated by the accumulation of frass and other debris entangled in larval silk over the surface of the injury. The larvae are voracious feeders and construct long meandering galleries through the injury site. Reports of larval damage on sugar cane has varied markedly. On Barbados, Alam (1984) noted extensive damage to live sugar cane, exceeding that caused by Diatraea saccharalis (F.). The young larvae feed un- der the leaf sheaths and, as they mature, penetrate the stalks and destroy the cane tissue. Infested stalks are hollowed out and gradually filled with larval frass. As in the case on banana, pupation occurs at the feed- ing site inside the stalks. At the infestation sites studied by J. E. Jones (in litt.), also on Barbados, O. sacchari was most abundant in dead or dying stumps and dead canes, suggesting they moved in following an in- festation by Diatraea. Jones also found that the high incidence of Opogona on green canes was associated with a high infestation PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Larval head capsule width for 7 instar groupings of Opogona sacchari (Bojer). Mean + SD Ratio of Instar N Range (mm) (mm) Increase 1 26 0.16-0.24 0.18 + 0.03 2 20 0.26-0.32 0.31 + 0.02 72 3 27 0.36-0.56 0.49 + 0.01 1.51 4 20 ~=—-0.58-0.88 =—-0.91 + 0.02 1.85 5 20 0.91-1.16 1.02 + 0.04 1.12 6 20 1.20-1.44 1.42 + 0.04 1.39 7 32 1.52-2.56 2.17 + 0.04 1.42 Average Nesy of Diatraea, again suggesting that O. sac- chari invades burrows previously created by Diatraea. Thus, the observations by Jones agree most closely with those originally re- ported by Bojer (1856). OBSERVATIONS IN FLORIDA A laboratory colony of O. sacchari was initiated in 1985 with larvae and pupae col- lected from infested Dracaena canes found at a nursery in Homestead, Florida. Pupae were held individually in 36 ml diet cups until eclosion. Emerging adults were sexed and 17 pairs were placed in 11 cm x 14.5 cm plastic oviposition chambers along with folded filter paper. Adults were provided honey and water. Pairs were held until the female died, males were replaced as needed. The preoviposition period, fecundity and longevity of females and the time required for egg to hatch were determined. First in- stars were gathered from the oviposition chambers and placed on one of 3 artificial diets in an attempt to rear this species in the laboratory. The 3 diets used were: velvet bean caterpillar diet (VBC), sugar cane borer diet, and the elm spanworm diet (Fedde 1974). All rearing and diet tests were con- ducted in a rearing room with a photoperiod of 12:12 (L:D) (0600 to 1800 photophase), temp 24 + 2°C, and RH 65-70%. Larvae were collected at different days from the diet, and preserved in a 70% alcohol for later measurement of head capsule width. Larvae VOLUME 92, NUMBER 4 615 Table 3. Mating activity of moths in rearing chamber. "| 25 4 % Males and Females in Copula 1°23: 4 5°96 7 8 9° 1017 12°13 1415: 16 17 16: 19:20 21 22 23 24 A.M. P.M. Time of Day (Hours) were checked daily for onset of pupation and pupal development time was recorded for 175 pupae. Emerging adults were also placed in 53 x 53 x 50 cm screen cages with Dracaena and Chamaedorea potted plants. Number of eggs and oviposition site on each plant species was inspected daily. RESULTS AND DISCUSSION Based on laboratory observations, devel- opment of an Opogona generation required 50-70 days. More eggs were present on unexpanded leaves and stems than on ex- panded leaves. Eggs are laid singly or in groups up to 328. They are light yellow at Oviposition, turn a dark yellow color ca. 2 days later, and finally yellowish brown prior to eclosion. Eggs hatched in 7.02 + 0.02 days at 24°C in the laboratory. The preovi- position period for newly emerged females in the laboratory was 1.66 + 0.14 days at 25°C. Female longevity (x + SD, n = 20) in the laboratory was 9-17 days with a mean of 11.45 + 0.72 days. Sixty first instar larvae were placed individually on each of the 3 diets tested. Survivorship on the artificial diet was low with the exception of velvet bean caterpillar diet. The highest percentage survivorship, 83%, was on VBC diet. Experiments were conducted to monitor O. sacchari sexual activity throughout the night. Sets (n = 30) of 2—3-day-old virgin males and females were placed in petri dish- es (9 cm in diameter) with water and honey as source of food. Each set was placed in an environmental chamber (LD 12:12, 24 + 2°C, 75-80% RH). The experiment was rep- licated 4 times. Diel activity was monitored hourly from 6:00 pm to 7:00 am. The period of activity occurred between 1:00 am to 4:00 am. No sexual activity was observed before 1:00 am or after 4:00 am. Through this ex- periment it became apparent that the op- 616 timal response period for O. sacchari occurs 3 h before the end of scotophase (darkness) (Table 3). The pupal stadium of larvae placed on VBC diet lasted 12.53 + 0.33 days. The average weight of pupae was 0.043 g. Table 2 lists a 7 instar model which best fits the data according to Dyar’s rule (Dyar 1890). As noted previously, these measurements are based on pooled data for unsexed larvae. Typical damage of O. sacchari on Dra- caena is characterized by removal of the bark and phloem. Cuttings of Dracaena having Opogona larvae show exterior debris and frass (Figs. 7, 8) deposits, and have in- ternal feeding damage on dead and living portions of the cortex, pith, roots and leaves (Fig. 6). Damage is not evident 4-6 weeks after infestation. Typical damage to Cha- maedorea palms can be observed 2-3 weeks after infestation. Each larva feeds at the base and roots of Chamadorea, and frass accu- mulates at the plant base from feeding into roots and petioles. In palms the leaf blades of the growing point become bleached and necrotic. ACKNOWLEDGMENTS The senior author (Davis) is responsible for the systematic and morphological por- tions of this paper and the biological ob- servations in Florida are by the junior au- thor (Pena). We are indebted to Vichai Malikul and Young Sohn of the Department of Ento- mology, Smithsonian Institution, for the line drawings and to Susann Braden and Brian Kahn of the Smithsonian SEM Lab and Vi- tor Kranz of the Smithsonian Photographic Laboratory for photographic assistance. The final draft of the manuscript was prepared by Callie Sullivan. We with to thank Joél Minet and Pierre Viette of the Museum of National d’Histoire Naturelle (Paris), John Heppner of the Florida State Collection of Arthropods (Gainesville, Florida), and Kevin Tuck of the British Museum (Natural PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON History, London) for providing information essential to our report. We also appreciate the comments by two anonymous reviewers of the manuscript. LITERATURE CITED Alam, M. M. 1984. New insect pests of sugarcane in Barbados. Barbados Sugar Technologists’ Asso- ciation, Second Annual Conference, pp. 1-6. Bennett, F. D. and M. M. Alam. 1985. An annotated check-list of the insect and allied terrestrial ar- thropods of Barbados. Caribbean Agricultural Re- search and Development Institute, pp. 1-81. Bojer, W. 1856. Report of the committee on the “cane borer.” Pp. 1-46, 5 pls. H. Plaidean Government Printer, Port Louis, Mauritius. Box, H. E. 1953. List of sugarcane insects. Pp. Il- 101. Commonwealth Institute of Entomology, London. Butler, A.G. 1876. Preliminary notice of new species of Lepidoptera from Rodriguez. Annals and Mag- azine of Natural History, 44th Series 17(101): 407- 409. Ciampolini, M. 1973. Opogona sacchari Bojer dan- nosa a piante ornamentali. Notiziario sulle Mul- lattie delle Piante, Nos. 88-89, pp. 221-225. So- cieta SIPCAM, Milan, Italy. Cintra, A. F. 1975. Opogona sp. nova praga da ba- nanicultura em Sao Paulo. Biologico 41(8): 223- EME Cintra, A. F., P. R. Almeida, I. Myazaki, and H. S. Neves. 1978. Fumigacao de banana com bro- meto de metila para cotrole de lagartas de Opo- gona sacchari (Bojer, 1856). Biologico 44(1): 3- 10. Cockerell, T. D. A. 1923. The Lepidoptera of the Madeira Islands. The Entomologist 56: 247. D’ Aguilar, J. and M. Martinez. 1982. Opogona sac- chari (Bojer) présent daus les cultures sons serres en France (Lep. Tineidae). Bulletin de la Société Entomologique de France 87(1/2): 28-30. Davis, D. R. 1978. The North American moths of the genera Phaeoses, Opogona, and Oinophia, with a discussion of their supergeneric affinities (Lep- idoptera: Tineidae). Smithsonian Contributions to Zoology 282: 1-39, 128 figs. . 1984. Tineidae, pp. 4-5, 19-24. Jn Heppner, J. B., ed., Atlas of Neotropical Lepidoptera, vol 2 (checklist), part 1 (Micropterigoidea to Immo- idea). Dr. W. Junk, the Hague. Declercq, R. and I. Van Luchene. 1977. Voorkomen en bestrijding van de rupsen van Opogona sacchari Boj. in Dracena fragrans en Sansevieria trifasciata ‘Laurentii.. Verbondsnieuws voor de Belgische Sierteelt 21(15): 499, 501, 1 pl. VOLUME 92, NUMBER 4 Durrant, J. H. 1923. A moth from Saint Helena in Britain. Proceedings of the Entomological Society of London, p. xvii. 1925. The banana moth, Hieroxestis subcer- vinella Wik. Entomologist’s Monthly Magazine 61: 12-13. Dyar, H.G. 1890. The number of molts of Lepidop- terous larvea. Psyche 5: 420-422. Fedde, Y. H. 1974. Rearing the elm spanworm on artificial diet: A preliminary report. USDA Forest Service, Ashville, N. C., SE-204, pp. 1-5. Giannoti, O., B. S. Oliveira, T. Toneda, and D. Fell. 1977. Observacoes gerais sobre 0 desenvolvi- mento e comportamento sexual do Lepidoptera Opogona sacchari (Bojer, 1856) em labortorio. Ar- quivos do Instituto Biologico 44(4): 209-212. Heppner, J. B., J. E. Pena, and H. Glenn. 1987. The banana moth, Opogona sacchari (Bojer) (Lepi- doptera: Tineidae), in Florida. Florida Depart- ment of Agriculture and Consumer Services, Di- vision of Plant Industry. Entomology Circular No. 293, pp. 1-4. Jannone, G. 1966. Risultati di alcuni controlli fitosa- nitari su vegetalie prodotti vegetali esteri nel Porto di Genova. Rivista di Agricoltura subtropicale e Tropicale 60(1-3): 5-34. Meyrick, E. 1910. Descriptions of Microlepidoptera from Mauritius and the Chagos Isles. Transactions of the Entomological Society of London, pp. 366- SiMe 1911. Tortricina and Tineina of the Percy Sladen Trust Expedition to the Indian Ocean in 1905. Transactions of the Linnaean Society of London, Zoology, 2nd series 14: 263-307. . 1924. Microlepidoptera of Rodriguez. Trans- actions of the Entomological Society of London, pp. 544-557. . 1930. Microlepidoptera of Mauritius. Trans- actions of the Entomological Society of London 78: 309-323. Moreton, B. D. 1974. New or uncommon plant dis- eases and pests. Plant Pathology 23: 163-164. Mourikis, P. A. aff P. Vassilaina-Alexopoulon. 1983. Data on the biology of the Opogona sacchari (Bojer 1856), a new pest for ornamental plants in Greece. Anales de L’institute Phytopathologique Benaki 13(1): 59-64. Oldham, J. N. 1928. Hieroxestis subcervinella WIk., an enemy of the banana in the Canary Islands. Bulletin of Entomological Research 1 9(pt. 2): 147- 166, pls. 4-5. Paulin, R. and P. Viette. 1955. Essai d’un catalogue biologique des Lepidopteres heteroceres de Tana- narive. Memoirs de L’Institut Scientifique de Madagascar, ser. E 6: 141-281. Pigatti, A. 1978. Selecao de inseticides em labora- 617 torio para o controle da ‘traca’ da banana—Opo- gona sacchari (Bojer, 1856) (= O. subcervinella Walker, 1863)—Lepidoptera: Lyonetiidae. Biol- Ogico 44(1): 21-23. Pigatti, A., P. Almeyda, A. Cinta, and D. Oliveira. 1979. Ensaios com inseticidas aplicados por via liquida para o controle da ‘“‘traca da banana” — Opogona sacchari (Bojer, 1856) (= O. subcervi- nella, Walker, 1863) Lepidoptera: Lyonetudae. Biologico 45(3/4): 61-68. Rebel, H. 1939. Achten Beitrag zur Lepidopteren- fauna der Kanaren. Annalen des Naturhistorisch- en Museums in Wien 49: 43-68. 1949. Anhang. Eine Lepidopteren-Ausbente von Madeira. Die Lepidopteren fauna des Azo- rischen Archipels 8(1): 50-57. Rotundo, G. and E. Tremlay. 1982. Preliminary ob- servations on the sex pheromone of Opogona sac- chari (Bojer). Bollettino del laboratorio di Ento- mologia Agraria Filippo Silvestri 39: 123-132. Suplicy Fo., N. and A. S. Sampaio. 1982. Pragas da Bananeira. Biologico 48(7): 169-182. Siiss, L. 1975. Opogona sacchari (Bojer) (Lepidop- tera, Lyonetiidae), a new pest of ornamental plants in greenhouses. Bollettino di Zoologia di Ento- mologia Agraria dell Universita 12: 1-28. Vari, L. and D. Kroon. 1986. Southern African Lep- idoptera, a series of cross-referenced indices. 198 pp. The Lepidopterists Society of South African and the Transvaal Museum, Pretoria. Veenenbos, J. A. V. 1981. Opogona sacchari, a pest risk from imports of ornamental plants of tropical origin. Bulletin, Organisation Européene et Méd- iterranéenne pour la Protection des Plantes 1 1(3): 235-237. Viette, P. 1951. Quelques Microlepidopteres de Mad- agascar. Memoires de L’Institut Scientifique de Madagascar, série A 5(2): 335-347. . 1957. Lepidopteres (excepte les tordeuses) et les Geometrides. Memoires de L’Institut Scienti- fique de Madagascar, série E 8: 137-226. 1958. Lepidoptéres Tinéides (s.1.) et Pyrales. Societas Scientiarum Fennica. Commentationes Biologicae, XVII. 8 17(8): 1-12. Vinson, J. 1938. Catalogue of the Lepidoptera of the Mascarene Islands. Mauritius Institute Bulletin 1 (pt. 4): 1-69. Walker, F. 1863. List of the specimens of Lepidop- terous insects in the collections of the British Mu- seum 28: 287-561. . 1875. Fam. Gelechidae [sic], p. 192. Jn Mel- liss, J. C., ed., St. Helena: A Physical, Historical, and Topographical Description of the Island, In- cluding its Geology, Fauna, Flora, and Meterol- ogy. xiv + 426 +pp., pls. 1-56. L. Reece & Co., London. 618 Walsingham, Lord (Thomas de Gray). 1907. Micro- lepidoptera. /n Sharp, D., ed., Fauna Hawaiiansis, or the Zoology of the Sandwich (Hawaiian) Isles I(pt. 5): 469-759, pls. 10-22. Cambridge Univer- sity Press. 1919. Madeiran Tineina (Lepidoptera). The Entomologist’s Monthly Magazine, second series 21(46): 257-259. Wollaston, E. 1879. Notes on the Lepidoptera of St. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Helena, with descriptions of new species. Annals and Magazine of Natural History, fifth series 3: 219-233, 329-343, 415441. Zandvoort, R. 1972. Bestrijding van Opogona sub- cervinella bij Strelitzia. Gewasbescherming 3(4): 95-96. Zimmerman, E. C. 1978. Insects of Hawaii. Micro- lepidoptera 9: 1-1876, figs. 1-1355. Honolulu: University of Hawaii Press. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 619-640 REVISION OF HOCKERIA WALKER IN THE NEARCTIC REGION WITH DESCRIPTIONS OF MALES AND FIVE NEW SPECIES (HYMENOPTERA: CHALCIDIDAE) JEFFREY A. HALSTEAD California State University Fresno, Fresno, California 93740. Present address: 110 W. Barstow #112, Fresno, California 93704. Abstract. —The genus Hockeria Walker is revised for the Nearctic region. Nine species are recognized; five new species (hainesi, bicolor, brevipennis, micra, and burksi) are de- scribed. Four previously described species, eriensis (Wallace), rubra (Ashmead), tenui- cornis (Girault), and unipunctatipennis (Girault), are diagnosed and discussed. Males of all species are described and allotypes or plesiotypes designated. Hockeria americensis (Girault) is designated a junior synonym of H. unipunctatipennis. A key to the Nearctic species for both males and females is presented. Characters of females and males are illustrated. Biological and distributional information is summarized for each species. Hockeria eriensis and H. bicolor n.sp. are recorded from the Neotropical region. Key Words: The worldwide genus Hockeria Walker contains about thirty described species. Walker described Hockeria in 1834 with H. bispinosa Fabricius, a European species (synonym of H. bifasciata Walker), as the nominal type-species. Major faunal treat- ments of Hockeria include: Japan (Habu 1960, 1962), Europe (Boucek 1951), USSR (Nikolskaya 1952, 1960), the Near East and India (Husain and Agarwal 1982). Husain and Agarwal (1982) presented a key to the world species of Hockeria but none of the Nearctic species were included. The Nearc- tic Hockeria are revised herein for the first time. No taxonomic keys or comprehensive treatments exist for this fauna. The fauna contains nine species, five of which are new- ly described in this paper. Past literature on Nearctic Hockeria is cataloged in Peck (1963), Burks (1979), and DeSantis (1979). Hockeria are small to moderate sized wasps (2 to 10 mm). Females have slender filiform antennae and are entirely black, red Insecta, Hockeria, Chalcididae, revision, new species, Nearctic or orange, or commonly with a combination of red or orange and black. Males have ro- bust filiform antennae, are usually black with some orange markings, and are more robust than females. The forewing of females is clouded in a specific pattern whereas in males it is usually clear. At present, a generic revision of the American Chalcididae is underway (Bou- cek, pers. comm.); therefore, a generic de- scription of Hockeria is omitted. However, to facilitate the identification of this genus for the Nearctic region the following char- acters are diagnostic: vertex not produced into horns, hindtibiae truncate distally, two apical hindtibial spurs present (Haltichel- linae); marginal vein on anterior margin of forewing, postmarginal and stigmal veins present (Haltichellini); tergite 1 without ca- rinae; posterior margin of scutellum with- out a median tooth; frontal carina weak, not joining in ocellar area to form an arch. Useful species characters for females in- 620 clude: clouded pattern of the forewing; col- or; shape of the abdomen, hindfemora, head, antennae, and ovipositor sheath; sculpture of tergite 1 and mesopleural acetabulum; length of flagellomeres; body length; and body sculpture. Characters for males in- clude: body length and color; T1 sculpture; forewing clouding and color; shape of the scutellum (especially the shape of teeth on the posterior margin or their absence), fla- gellomeres, and propodeal carinae. The taxonomy of the Nearctic Hockeria has been based on females, and only the male of H. eriensis (Wallace) has been de- scribed. Strong sexual dimorphism and di- chromatism makes the male-female asso- ciations difficult. Males usually have a robust body, robust filiform antennae, clear wings, and black body coloration; whereas, females usually have a more slender body, slender filiform antennae, clouded wings, and red- orange and black body coloration. Species exhibiting strong sexual dimorphism and dichromatism include: eriensis, rubra (Ash- mead), tenuicornis (Girault), unipunctati- pennis (Girault), and hainesi n. sp. The species micra n. sp., burksin. sp., bicolor n. sp., and brevipennis n. sp. are less dichro- mic. Males are similar morphologically, which complicates the task of distinguishing them and making the proper female asso- ciation. However, examination of large se- ries has permitted the male-female associ- ation for all species. These males are described and specimens designated as al- lotypes or plesiotypes. The nine Nearctic species will not be clas- sified into species groups at this time. I think a world overview is necessary to determine and designate species groups. However, two species (H. eriensis and H. bicolor n. sp.) form a unique group separate from other Nearctic species in having a narrow head, globose abdomen, and strongly arched scu- tellum. Boucek (1951) also noted short, stout forms and slim forms in the European fau- na. World literature denotes a wide range of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hosts for Hockeria: antlion larvae (Neurop- tera), elasmid and tenthredinid pupae (Hy- menoptera), free-living Strepsiptera, dipter- an pupae, and commonly lepidopteran larvae and pupae (Boucek 1951, Habu 1962, Burks 1979, Narendran and Rao 1987). Hosts have been determined for six of the nine Nearctic species, including three eco- nomically important lepidopterous pests: the Western Grapeleaf Skeletonizer (Harrisina brillians Barnes and McDunnough), the Nantucket Pine Tip Moth (RhAyacionia frus- trana (Comstock)), and the Ponderosa Pine Tip Moth (Rhyacionia zozana (Kearfott)) and a new host record of ascalaphid larvae (Neuroptera). Hockeria is widely distributed through- out the Nearctic region (Fig. 53). The dis- tribution map is based upon specimens ex- amined by the author, and it encompasses most of the literature records. Hockeria are found in a variety of habitats and eleva- tions. In California, some species (e.g. erien- sis and rubra) range from coniferous forests to deserts. Several species range throughout the entire Nearctic region. A few species are known only from the western United States although additional collecting will likely ex- tend their range. No species are restricted to the eastern United States or Mexico and interestingly, all Nearctic species have been collected in California. It is possible that the Nearctic species may also occur in the Pale- arctic or Neotropical regions; however, this awaits further study. Despite their broad range, Hockeria are rarely collected and un- common in collections. Sweeping flowering vegetation or vegetation in general, and us- ing Malaise-type traps and pan-traps are successful collecting techniques. Many Hockeria specimens, representing seven species, were collected from a hydro- electric flume which runs through Foothill Woodland and Chamise Chaparral plant communities 660 m (2200 ft) in Tulare County, California. Large series of unde- scribed, rarely collected, and/or poorly rep- resented species were collected from this VOLUME 92, NUMBER 4 source (Halstead and Haines 1987). With- out these specimens, species variation, male/ female associations, and complete distri- butions would have been difficult to deter- mine. During this study, somewhere be- tween 1000 to 2000 specimens of Hockeria were examined. Collections examined and museum ac- ronyms are as follows: American Museum of Natural History, New York; Bernice P. Bishop Museum, Hawaii; California Acad- emy of Sciences, San Francisco (CAS); Cal- ifornia Department of Food and Agricul- ture, Sacramento (CDFA); California State University, Fresno; California State Uni- versity, Sacramento; Canadian National Collection, Ottawa (CNC); Carnegie Mu- seum of Natural History, Pittsburg, Penn- sylvania (CMNH); Florida Department of Agriculture and Consumer Affairs, Gaines- ville (FDA); Fresno County Department of Agriculture Fresno, California; Illinois Nat- ural History Survey, Champaign; Los An- geles County Museum of Natural History, California (LCM); Mississippi State Uni- versity, Mississippi State; Natural History Museum of San Diego, California; Oregon Department of Agriculture, Salem; Royal Ontario Museum, Toronto (ROM); Texas A&M University, College Station; Tulare County Agricultural Commissioner’s Of- fice, Visalia, California; United States Na- tional Museum of Natural History, Wash- ington D.C. (USNM); University of California, Berkeley; University of Califor- nia, Davis; University of California, Riv- erside (UCR); University of Georgia, Ath- ens (UOG); J. A. Halstead personal collection (JAH); H. A. Hespenheide per- sonal collection, Los Angeles, California (HAH); R. B. Miller personal collection, Project City, California (RBM); R. D. Haines personal collection, Visalia, California (RDH). Abbreviations include: T1 for tergite 1, etc.; OOL (ocular-ocellar line) for the small- est distance between the compound eye and lateral ocelli; OL (ocellar line) for the small- 621 est distance between the anterior ocellus and lateral ocelli; LOD for lateral ocellar di- ameter; AOD for anterior ocellar diameter. All measurements were made in the flattest plane possible. Specimens were examined at 30 to 100 x. A mylar, glare reducing screen was used in lighting specimens. Key TO NEARCTIC SPECIES OF HOCKERIA 1. Females; ovipositor present (Figs. 1-10) ... 2 — Males; ovipositor absent ................. 10 2. Gaster (lateral view) about 1'2* as long as wide, apex rounded (Figs. 8-10) Ene 5 — Gaster (lateral view) 2-3 as long as wide, apex pointed (Figs. 1-7) . ercrais tes 4 3. Hindfemur about 3 x as long as wide, without ventral projections (Fig. 40, rarely as in Fig. 38); Tl punctate dorsally, with coriaceous band posteriorly ............ eriensis (Wallace) — Hindfemur about 2 as long as wide, with an anterior toothlike projection and a rounded posterior projection (Fig. 39); T1 coriaceous PO Aan 5 ore Penner bicolor Halstead n. sp. 4. Tl punctate dorsally; forewing with a single clouded area under marginal vein (Figs. 45- 46), rarely with no or 2 clouded areas; body black — TI polished or slightly coriaceous dorsally: forewing with two clouded areas (Figs. 41-44) or a clear circular area laying within a large clouded area (Fig. 47); body partly or entirely red or orange 5. Apex of ovipositor sheath with dorsi margin evenly rounded (lateral view) (Fig. 6); length about 3.8 mm burksi Halstead n. sp. — Apex of ovipositor sheath with dorsal margin angled (Fig. 5); length about 2.5mm ...... ne pa RINE eet Se BT micra Halstead n. sp. 6. Forewing with a clear circular area containing a dense patch of white setae, enclosed within a brown clouded area (Fig. 47) Rr nice eniee unipunctatipennis (Girault) — Forewing without a clear circular area (Figs. 41-44) . set eA, 7. Length ies ane 3mm Q2. 5: to 2 2.8 ane 7 8 Length greater than 4 mm (4 to 10 mm) 9 8. Apex of ovipositor sheath with dorsal margin evenly rounded (lateral view) (Fig. 3); head, thorax, propodeum, and legs partly black .. nto . hainesi Halstead n. sp. ~ eee ah ovipositor aheath with dorsal margin squared (Fig. 4); head, thorax, propodeum and legsorange) 2b). eevect brevipennis Halstead n. sp. 9. Apex of ovipositor sheath with dorsal margin 622 NS}, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON evenly rounded (Fig. 2); head, thorax, pro- podeum, and legs partly black NES Syst ees SARE OOM DOS tenuicornis (Girault) Apex of ovipositor sheath with dorsal margin angled (squared) (Fig. 1); head, thorax, pro- podeum and legs red to orange rubra (Ashmead) T1 punctate dorsally, posterior margin with a thin coriaceous band T1 coriaceous dorsally or if punctate, punc- tures extending *4 or less the length of T1; posterior margin with a broad coriaceous band. Or, if scutellum is strongly arched (Fig. 51) go to couplet 12 . Wings darkly clouded throughout, commonly with an orangish tint; scape, tegulae, and legs usually orange; head (lateral view) oval and interantennal projection large (Fig. 27) . unipunctatipennis (Girault) Wings clear or with a small clouded spot; body color mostly black; head (lateral view) oblong and/or interantennal projection small (Figs. 2526450) 12 . Scutellum strongly arched dorsally (Fig. 51); forewing with apical 7 clouded and a prom- inent brown spot under marginal vein side iotas ieee h rscans etme hisarilis eriensis (Wallace) Scutellum slightly convex; forewing clear or with a faint clouded spot under marginal vein 13 . Flagellomeres 2 to 2'2x as long as wide; me- sopleural acetabulum with sculpture between strong transverse carinae polished a SAA aR burksi Halstead n. sp. Flagellomeres 3-8 | to 12x as long as wide; mesopleural acetabulum with sculpture be- tween weak transverse carinae punctate.... 14 . Propodeum with a strong longitudinal, sub- median carina; anterior area of mesopleuron punctate and rugose .... /ainesi Halstead n. sp. Propodeum with a oval reticulation of carinae medially; mesopleuron anteriorly smooth and polished, punctate only ventrally .......... micra Halstead n. sp. T1 coriaceous dorsally; tergites without oval macropunctures; body partly or entirely or- ‘ange to red-brown) (jsienrds nein eieroieae 16 Tl punctate dorsally, sometimes punctures shallow and faint, appearing somewhat pol- ished—if so, tergites with oval macropunc- tures (Fig. 52); body black . Scutellum with two, wide, triangular teeth at posterior margin, sculpture coriaceous cen- trally; head and thorax with well defined, moderately deep punctures which are sepa- rated by 2 to 's their diameter, sculpture acic- ulate, polished; body orange to red and black bicolor Halstead n. sp. — Scutellum rounded at posterior margin, sculp- ture matte; head and thorax with shallow, vague punctures which are separated by 6 to Y their diameter, sculpture smooth, matte; body orange-brown . . brevipennis Halstead n. sp. 17. Posterior margin of scutellum with two tri- angular teeth; T1 dorsolaterally with macro- punctures; band of macropunctures on other tergites prominent (Fig. 52) .. rubra (Ashmead) — Posterior margin of scutellum rounded to truncate; T1 dorsolaterally without macro- punctures, macropunctures on other tergites ab- sent or faint and shallow ..... tenuicornis (Girault) Hockeria eriensis (Wallace) Figs. 8, 10, 18, 20, 28, 30, 38, 40, 48, 50, 51, 53 Stomatoceras rubra var. eriensis Wallace, 1942: 31,2 &6. Stomatoceras rubrum eriense Wallace; Peck 1951: 585. Hockeria eriensis (Wallace); Burks, in Stef- fan 1959: 304. Female diagnosis (species).— Length about 5.0 mm. Red or orange with teeth of mandibles, mesosternum, anterior margin of mesoscutum, metanotum laterally, fla- gellomere 4 or 5 to apex, and teeth on ven- tral margin of hindfemur black. Head as in Fig. 30. Antennae (Fig. 20) geniculate. Scutellum (Fig. 51) strongly arched. Forewing (Figs. 48, 50) with one or two clouded areas: at apex of marginal vein and in middle of wing near apex. Hindfemur (Fig. 40) narrow, elongate, without promi- nant ventral projections. Abdomen (Fig. 10) globose, apex blunt. The female of Hockeria eriensis is most similar to H. bicolor n. sp. though is distin- guished by its hindfemur shape. These two species (females) differ from other Nearctic Hockeria by having a narrow head, globose abdomen, and strongly arched scutellum. Variation (2).—Length 2.5 to 5.0 mm. Most specimens are orange, or red with black areas. Wallace (1942) noted “‘dark females in which the head and thorax are almost entirely black, and the abdomen heavily VOLUME 92, NUMBER 4 623 O016 1. BU 15KYoK4800 Fig. 4. Third instar larva of St. affinis. (A) Venter, arrow denotes depressions along segmental lines; (B) lateral grouping of sensilla; (C) detail of middle, wart- like sensillum. 5.7 + 0.15 mm in length and 2.5 + 0.06 mm in widest width. The puparium is trans- lucent, oblong, barrel-shaped, rounded at both ends, and smooth. Prior to eclosion the pharate adult is easily visible through VOLUME 92, NUMBER 4 1SKV k44 @018 100.6U KU X40 a eR Fig. 5. Second instar larva of S?. affinis. (A) Ha- bitus; (B) gnathocephalon; |—median oral lobe; 2— lateral sensory organ; (C) anterior thoracic spiracle. the puparial integument; it was also noted that the wings of the adults were fully pig- mented within the puparia. The anterior end bears the open anterior thoracic spiracles, and the lateral fracture line which extends posteriorly for three segments (Fig. 6A, ar- 647 15KU Xeeuad. a) 8006. Le ou Fig. 6. Puparium of Sv. affinis. (A) Anterior end, arrow denotes lateral fracture line; (B) posterior end, arrow denotes raised posterior spiracles; (C) posterior spiracles. row). The posterior spiracular plates slightly protrude dorsal to the transverse midline (Fig. 6B, arrow); the rimae are elongate-oval; the spiracular slits are ca. 0.05 mm long; and the interspiracular processes are small and lack well-defined blades (Fig. 6C). 648 ACKNOWLEDGMENTS We thank D. W. Ricker for technical as- sistance, including the insect photography in Figs. 1 and 2; A. C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for plant identifications; J. LaSalle, now with the Commonwealth Ag- ricultural Bureaux International Institute of Entomology, London, U.K., for identifying Tetrastichus sp.; and A. L. Norrbom, Sys- tematic Entomology Laboratory, USDA, ARS,% U.S. National Museum, Washing- ton, D.C., for identifying St. affinis. We also are grateful for manuscript reviews by F. L. Blanc, G. Gordh, A. L. Norrbom, and G. J. Steck. LITERATURE CITED Carroll, L. E.and R. A. Wharton. 1989. Morphology of the immature stages of Anastrepha ludens (Dip- tera: Tephritidae). Ann. Entomol. Soc. Am. 82: 201-214. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Chu, I-Wu and R. C. Axtell. 1972. Fine structure of the ventral organ of the house fly larva Musca domestica L. Z. Zellforsch. Mikrosk. Anat. 130: 489-495. Foote, R.H. 1965. Family Tephritidae, pp. 658-678. In Stone, A. et al., eds., A Catalog of the Diptera of America North of Mexico. USDA Agric. Handbk. 276. Foote, R. H. and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect Surv. 7, 115 pp. Headrick, D.and R. D. Goeden. In press. Description of the immature stages of Paracantha gentilis (Diptera: Tephritidae). Ann. Entomol. Soc. Am. Munz, P. A. 1974. A Flora of Southern California. Univ. Calif. Press, Berkeley, Los Angeles, London. 1086 pp. Novak, J. A. and B. A. Foote. 1975. Biology and immature stages of fruit flies: The genus Stenopa. J. Kansas Entomol. Soc. 48: 42-52. Novak, J. A., W. B. Stolzfus, E. J. Allen, and B. A. Foote. 1967. New host records for North Amer- ican fruit flies (Diptera: Tephritidae). Proc. Ento- mol. Soc. Wash. 69: 146-148. Quisenberry, B. F. 1949. Notes and description of North American Tephritidae (Diptera). J. Kansas Entomol. Soc. 22: 81-88. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 649-671 THE ASPHONDYLIA (CECIDOMYIIDAE: DIPTERA) OF CREOSOTE BUSH (LARREA TRIDENTATA) IN NORTH AMERICA RAYMOND J. GAGNE AND GWENDOLYN L. WARING (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560, USA; (GLW) Museum of Northern Arizona, Rt. 4, Box 720, Flagstaff, Arizona 86001, USA. Abstract. —Fifteen species of gall midges of the genus Asphondylia that form complex galls on leaves, stems, or buds of creosote bush are described. Fourteen of the species are new to science, the other is redescribed. One other species that was caught in flight and is similar to the leaf gall makers of Larrea is also redescribed. The Asphondylia spp. on creosote bush appear to be a monophyletic group and are treated as the Asphondylia auripila species group. Key Words: Fifteen distinct kinds of complex galls growing on leaves, stems, and buds of Lar- rea tridentata (Sessé & Mocino ex DC.) Cov. (Zygophyllaceae) were found by G. L. War- ing during the course of an ecological study of this plant. Each type of gall is formed by a different species of the genus Asphondylia, all of them except one new to science. In this paper we describe or redescribe these gall midges and place them in context with one another and with the rest of the genus. The natural history, ecology, and natural enemies of these flies have been or will be treated separately in Waring (1987), Waring and Price (1989a, b), and Waring (in prep- aration). Larrea tridentata, or creosote bush, is a dominant member of southwestern desert plant communities from Texas to California (Mabry et al. 1977, Waring 1986). It is a perennial, evergreen shrub, and one of the most drought-tolerant plants in southwest- ern United States. Larrea is restricted to the New World and is one of many taxa of plants and animals that show a disjunct distribu- Complex galls, Southwestern desert, gall midges tion between the southwestern North American and South American deserts. Larrea tridentata is the only species of Lar- rea in North America, while four others oc- cur in southern South America (Waring 1986). Asphondylia is a large, cosmopolitan ge- nus of 247 described species (Foote 1965, Gagné 1968, Gagné 1973, Gagné in press, Gagné in prep., Harris 1980, Skuhrava 1986). To date, 67 species have been de- scribed from the Nearctic Region (Gagné in prep.). Almost as many more Nearctic species are known but not yet described (Gagné 1989). Gagné (1989) listed the de- scribed and undescribed Nearctic species and their hosts and discussed Asphondylia in general. A thorough generic analysis of the tribe (as a supertribe) to which the genus belongs was done by Mohn (1961). The Nearctic species of Asphondylia have not been revised since Felt (1916), but recent studies were made of a monophyletic group of eight species that occurs on Chenopodi- aceae in California (Hawkins et al. 1986) 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON es , Nica = NIA. — @ “PF m b Lees ) A® uN AS "tut ROR SHG Prd 5 Pars to . an = i? ASS sag ~~ (ee : z ' 9g a Fig. 1. Stem and bud galls of Larrea tridentata formed by Asphondylia spp. Sprig of plant in | x, details of galls in 3 x. la: Stem gall of A. auripila, the detail with outer leaves removed to show the individual cells beneath; b, stem galls of A. foliosa; c, stem gall of A. rosetta; d, apical bud gall of A. apicata; e, node galls of A. bullata; f, flower gall of 4. florea; g, stem galls of A. resinosa, the resin of one of the enlarged pair removed to show detail. and of Asphondylia websteri Felt, an appar- ent generalist known from some Fabaceae and other plants (Gagné and Wuensche 1986, Gagné and Woods 1988). In addition, one of us (RJG) made a survey for this study of certain characters on all known described Nearctic species. Asphondylia adults are between 1-5 mm in length and are relatively robust with cy- lindrical antennae, large eyes, and an almost complete covering of scales. They are gen- erally brown to dark brown, but some species, such as Asphondylia monacha Os- ten Sacken and relatives, have black- and white-banded legs and are otherwise cov- ered with black scales. Females have a rigid, protrusible, needlelike ovipositor (Figs. 7, 8) with which they insert their eggs into liv- ing plant tissue. Larvae are generally white to yellow, have three instars, and always occur singly, either taking up the entire gall or an individual cell in aggregate galls. The last instar is robust and has a spatula (Figs. 35-47), a hard, brown to black dermal struc- VOLUME 92, NUMBER 4 k "Xe { \ 4 Cy ye Sate <> 7 ' “28 RESULTS EY Aedes (Aedimorphus) vexans (Figs. 1-3) Size: dimensions as in Table 1. Color: dark bronze. Overall appearance: shape variable, cur- vature of ventral surface greater than dorsal, greatest diameter somewhat anterior to middle, anterior taper more pronounced, posterior more gradual (Fig. 1). Outer cho- rionic cells uniformly elongate longitudi- nally (in long axis of egg), each more or less completely filled by longitudinally aligned outer chorionic tubercles. Tubercles occa- sionally in a single row, but usually in two rows, at least at widest part of cell. Micro- pylar collar indistinct. Chorion, dorsal, lateral and ventral sur- faces: all surfaces very similar (Fig. 1). Outer chorionic cells longitudinally elongate, 20- 42 um long, 8-12 wm wide (2.5-5 times as long as wide), irregularly polygonal with Fig. 1. Ae. vexans. Entire egg lateral view; dorsal boundaries clearly defined but not very surface at right, anterior end at top. Scale = 100 um. straight (Figs. 1, 2a, b). Cell fields 17-39 um VOLUME 92, NUMBER 4 Fig. 2. Ae. vexans. (a) Outer chorionic cells, ventral surface, middle of egg; (b) detail of outer chorionic cells and tubercles; (c) detail of individual outer chorionic tubercles and outer chorionic reticulum. Scale = 10 um (a, b), = 5 um (c). 687 long, 7-10 um wide, with smooth floors (Fig. 2c). Outer chorionic tubercles 6-15 in num- ber, fewer per cell on dorsal surface than on ventral (lateral surface not counted), but not significantly so (Table 2). Tubercles ar- ranged longitudinally in a single row, or more frequently a double row at widest parts of cell (Fig. 2a, b). Outer edges of tubercles almost always touching outer chorionic re- ticulum, gaps separating tubercles strikingly uniform, ca. 1.5 wm (Fig. 2a, b). Shape of tubercles irregular, roughly po- lygonal, shapes of edges tending to match those of adjacent tubercles (Fig. 2b), largest tubercles ca. 5.5 um in longest dimension, smallest ca 2.4 um, but very small tubercles uncommon. In detailed structure, each tu- bercle consists of a base, often with slightly concave inner edges with sloped, tapered walls rising to a smaller, flat top ornamented with poorly defined bumps and fissures (Fig. 2b, c). Outer edges of tubercle bases usually rounded (Fig. 2c). Outer chorionic reticu- lum low, width 1.2-3.5 um, consisting of a very fine reticulate meshwork with central line of small, bead-like protuberances (Fig. 2b, c), more or less evenly spaced (1.0-3.2 um). Meshwork usually touching and con- tinuing some distance up sides of tubercles (Fig. 2c). Anterior pole and micropyle: outer cho- rionic cells diminish somewhat in length to- wards anterior pole, becoming narrower, with outer chorionic cells reduced to single row (Fig. 3a, b). Tubercles not all separated by uniform gaps (Fig. 3a), many gaps nar- rower and shallow. Many tubercles close to- gether or almost fused, appearing more rounded and less distinct, especially just posterior to micropyle (Fig. 3b). Anterior ring present but not well formed, usually incomplete (Fig. 3c, d), diameter 35—45 um, variable, and of very variable width (0-7 um). Micropylar collar not prominent (Fig. 3a), height 6-10 wm and variable, diameter 20-28 um and not always circular or con- tinuous (Fig. 3c, d), internal diameter 18- 23 um, wall width 2-6 um, very variable, 688 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON —— — * a Fig. 3. Ae. vexans. (a) Anterior pole and micropylar apparatus, lateral surface; (b) anterior pole, lateral surface, chorionic cell detail; (c) top view, anterior pole and micropylar apparatus; (d) top view, detail of micropylar apparatus; (e) posterior pole, lateral surface; (f) posterior pole, ventral surface, outer chorionic cell detail. Scale = 10 um. VOLUME 92, NUMBER 4 689 Table 1. Dimensions of the eggs of two species of Aedes (n = 15). Length (um) Width (um) L/W Ratio Species Mean (+SE) Range Mean (+SE) Range Mean (+SE) Range Ae. vexans 650.9 + 3.3 637.3-665.3 197.6 + 1.8 193.7-203.9 3.29 + 0.03 3.14-3.54 Ae. infirmatus 664.4 + 3.6 637.3-685.7 207.8 + 1.9 201.4-224.3 3.20 + 0.03 3.02-3.40 outer margin irregular. Micropylar disc fair- ly clearly defined, domed (Fig. 3c, d), di- ameter 10-18 um, micropyle indistinctly trilobed, diameter ca. 2.6 um. Posterior pole: outer chorionic cells be- come shorter near posterior pole, gaps be- tween outer chorionic tubercles become more irregular (Fig. 3e), some tubercles fused and all fused in cells immediately at and adjacent to pole (Fig. 3e, f). Aedes (Ochlerotatus) infirmatus (Figs. 4-6) Size: dimensions as in Table 1. Color: satiny black. Overall appearance: shape fusiform, somewhat variable, ventral surface more convex than dorsal, widest point just an- terior to middle, anterior taper more abrupt than posterior, both anterior and posterior dorsal margins straighter than more curved ventral margins (Fig. 4). Outer chorionic cells appearing irregular in outline, bound- aries difficult to distinguish, outer chorionic tubercles clearly visible and many quite large, but not conforming to any easily dis- cernible pattern (Fig. 4). Micropylar collar relatively inconspicuous, conforming to ta- per of egg. Chorion, ventral surface: outer chorionic cells somewhat longitudinally elongate, po- Table 2. of Aedes (n = 15). lygonal, but very variable in form, 20-26 um long, 10-13 um wide, boundaries quite straight (Fig. Sa), corresponding cell field dimensions ca. 17-25 wm and 9-11 um. Outer chorionic tubercles 2-7 in number, fewer than on lateral and dorsal surfaces (Table 2), more or less evenly spaced lon- gitudinally in cell, usually but not always touching outer chorionic reticulum on at least one side (Fig. Sa, d). Largest tubercles 6-11 um long (longest dimension), some as small as 1.2 um, but very small tubercles fairly uncommon. Form of tubercles com- plex and irregular, each consisting of a low, smooth base, often with deeply excavated outline (Fig. 5a, d) and a smaller, domed upper portion, which is somewhat less ir- regular in outline and covered with small rounded bumps (Fig. 5d). Tubercles mostly separated from one another, but occasion- ally joined by narrow bridges (Fig. 5a). Floors of outer chorionic cell fields smooth, but partly covered with a very thin layer of irregular outline usually extending some distance from field edges, but occasionally forming narrow bridges to tubercles or com- pletely across cell (Fig. 5d). Outer chorionic reticulum low, width 0.9-3.2 um, consisting of a fine reticulate meshwork with a line of small (ca. 0.3-0.5 wm diameter), rather un- Numbers of outer chorionic tubercles in outer chorionic cells on different egg surfaces of two species Dorsal Surface Lateral Surface Ventral Surface Species Mean (+SE) Range Mean (+SE) Range Mean (+SE) Range Ae. vexans 827. 0:7 6-14 * 10.0 + 0.5 7-15 Ae. infirmatus 5: 5)200 spp.), mostly neo- tropical families, have only one West Indian species each. ACKNOWLEDGMENTS Miller’s fieldwork, as well as preparation of the illustrations, was supported by The Conservation Agency, with a grant from the The Falconwood Corporation. L. Laszlo Meszoly and Epstein illustrated the geni- talia. We thank the curators of the collec- tions consulted for use of specimens. R. W. Hodges, D. R. Davis, N. L. Evenhuis, and S. J. Weller reviewed the manuscript. LITERATURE CITED Becker, V. O. and M. E. Epstein. Limacodidae. /n Heppner, J. B., ed., Atlas of Neotropical Lepi- doptera. Checklist: Part II. E. J. Brill. (In press.) Brock, J. P. 1971. A contribution towards an under- standing of the morphology and phylogeny of the Ditrysian Lepidoptera. J. Nat. Hist. 5: 29-102. D’Arcy, W. G. 1967. Annotated checklist of the di- cotyledons of Tortola, Virgin Islands. Rhodora, 69: 385-450. Dyar, H. G. 1897. On the white Eucleidae and the larva of Calybia slossoniae (Packard). J. New York Entomol. Soc. 5: 121-126, pl. V. . 1899. The life-histories of the New York slug caterpillars. —(conclusion). J. New York Entomol. Soc. 7: 234-253, pl. VI-VIII. 1905. A list of American cochlidian moths, with descriptions of new genera and species. Proc. U.S. Natl. Mus. 29: 359-396. 1935. Limacodidae. In Seitz, A., ed., Die Gross-Schmetterlinge der Erde. Stuttgart, Alfred Kernen. Volume 6, pp. 1104-1139. Epstein, M. E. 1988. An overview of slug caterpillar moths (Lepidoptera: Limacodidae) with emphasis on genera in the New World Parasa group. Ph.D. Thesis, Univ. Minnesota. x1 + 149 pp. Fletcher, D. S. and I. W. B. Nye. 1982. Jn Nye, I. W. B., ed., The generic Names of Moths of the World, volume 4. London, British Museum (Natural His- tory). xiv + 192 pp. VOLUME 92, NUMBER 4 Forbes, W. T. M. 1930. Insects of Porto Rico and the Virgin Islands— Heterocera or moths (except- ing the Noctuidae, Geometridae and Pyralidae). Sci. Surv. Porto Rico Virgin Islands 12: 1-171. Grote, A. R. 1882. New Check List of North Amer- ican Moths. New York, New York Entomol. Club. 73 pp. 1888. The classification of the Bombycidae (third paper). Can. Entomol. 20: 181-185. Grote, A. R.and C. T. Robinson. 1868. Descriptions of American Lepidoptera—No. 4. Trans. Amer. Entomol. Soc. 2: 179-206, pl. 2-3. Heatwole, H., R. Levins, and M. D. Byer. 1981. Bio- geography of the Puerto Rican Bank. Atoll Res. Bull. 251: 1-55. Kirby, W. F. 1892. A synoptic catalogue of Lepi- doptera Heterocera. (Moths.) Vol. I. Sphinges and Bombyces. London, Gurney and Jackson. xi + 951 pp. Lazell, J. D., Jr. and L. Jarecki. 1985. Bats of Guana, British Virgin Islands. Amer. Mus. Novitates 2819: 1-7. Martorell, L. F. 1948. A survey of the forest insects 715 of Puerto Rico. J. Agr. Univ. Puerto Rico 29: 69- 608. (“1945”) 1976. Annotated Food Plant Catalog of the Insects of Puerto Rico. Agr. Exp. Sta., Univ. Puer- to Rico. 303 pp. Miller, S. E. Systematics of the Neotropical moth fam- ily Dalceridae (Lepidoptera). Bull. Mus. Comp. Zool. (In press.) Smithe, F. B. 1975. Naturalist’s Color Guide. New York, American Museum of Natural History. Van Eecke, R. 1925. Cochlidionidae (Limacodidae). Lepidopterorum Catalogus (32): 1-79. Wolcott, G. N. 1936. “Insectae Borinquenses”: A revised annotated check-list of the insects of Puer- to Rico. J. Agri. Univ. Puerto Rico 20: 1-627. 1951. The insects of Puerto Rico. J. Agni. Univ. Puerto Rico 32: 1-975. (*1948”) Wood, B. J. 1984. Implementation of integrated pest management in plantation crops, pp. 295-309. In Lee, B. S., W. H. Loke, and K. L. Heong, eds., Integrated Pest Management in Malaysia. Kuala Lumpur. Malay. Plant Protection Soc. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 716-724 A NEW SPECIES AND DISTRIBUTION RECORD FOR THE GENUS CAECULUS DUFOUR (ACARI: CAECULIDAE) FROM SOUTH DAKOTA B. MCDANIEL AND A. BOE Plant Science Department, South Dakota State University Brookings, South Dakota 57007. Abstract. —Caeculus lewisi, a new species of the genus Caeculus, is described and illus- trated. A new distribution record for the genus Caeculus is recorded for South Dakota along and a key to the species of the family Caeculidae is presented. Key Words: During the course of ecological studies in the 1970’s connected with the grassland In- ternational Biome Program, invertebrate fauna of grasslands in northwestern South Dakota was investigated. At that time, a new species of Caeculus was discovered. The dominant vegetation of the study area was comprised of buffalograss [Buchloe dacty- loides (Nutt.) Engelm.], blue grama [Bou- teloua gracilis (H.B.K.) Griffiths], and west- ern wheatgrass (Agropyron smithii Rydb.). Interspersed within the Pierre shale that supports these dominant grasses were out- crops of coarser-textured soil on which species more characteristic of tall-grass prairies were found. The most abundant grasses on these outcrops were big bluestem (Andropogon gerardii Vitman), little blue- stem (Andropogon scoparuis Michx.), In- diangrass [Sorghastrum nutans (L.) Nash], and switchgrass (Panicum virgatum L.). Field observations of this new species of mite suggested it was feeding on phlox (Phlox sp.) plants that were growing on the out- crops. No specimens of this new species were collected in the short-grass dominated areas surrounding the outcrops. However, obser- vations of collected mites studied with a dissecting microscope revealed they were Caeculus, rake-legged mites, distribution, fungus feeding on a fungus associated with the phlox stems. Similar studies of other plants col- lected from the rocky outcrop confirmed that this species of Caeculus feeds on fungal spores. Crossley and Merchant (1971), in laboratory observations of a species of Cae- culus, demonstrated with radioactive tracers that their species of Caeculus also fed on fungus. Mites were collected in the field and transferred to petri dishes in the laboratory. Parts of plants containing fungal spores were placed in the dishes as a source of food. However, all captured individuals died without completing their life cycle. Mc- Daniel (1979) only provided a description of the genus Caeculus. However, before the text was shortened for publication a key to each of the then described species of rake- legged mites from the United States and their distributions were included. That key and distribution data are presented here. Caeculus lewisi, NEw SPECIES (Figs. 1-23) Female (Figs. 1-6) anterior border of propodosoma projecting forward over gnathosoma with six flattened crescent shaped curved setae; two pairs of lateral eyes associated with plate region of propodoso- VOLUME 92, NUMBER 4 Fig. 1. dorsal view. Caeculus lewisi new species holotype female ma; propodosomal plate with six spatulate setae; median dorsal plate (Fig. 1) with nine spatulate setae, plate longer than wide, an- terior and posterior borders subequal; lat- eral median dorsal plate with four spatulate setae each, three aligned in a row vertically, fourth setae paired with third iateral plate seta near posterior border of median dorsal plate; opisthosomal transverse plates fused into single transverse plate across opistho- soma, with six spatulate setae; opisthosoma with 10 spatulate setae scattered near pos- terior region of opisthosoma; all dorsal spat- ulate setae with inner plumose region; body heavily armored, wrinkled, dark brown in color; length 1.5 mm; width between eyes 0.5 mm; propodosomal region wider than opisthosomal region; ventral area below an- terior projection of propodosomal plate contains a forked prominence with two spines (Fig. 2), below this structure 1s a cres- cent-shaped groove between chelicerae; pal- pal tibia with single terminal claw; palpal tarsus with terminal setae; first coxa with four spatulate setae; second and third coxa ach ry { % 7 7 x \ \ } \ ra aS / \ y, } f } 4 | ee I~ ARN) YK NX = Naat ay 4 s } Fig. 2. Caeculus lewisi new species holotype female ventral view. with three spatulate setae; fourth coxa with four spatulate setae; 20 setae in center of venter forming a double row starting from between fourth coxa, extending on either side of genital opening, each row with seven setae; three setae are grouped in a circle one either side of row of seven setae; genital opening with two rows of seven simple se- tae; anal opening with two pairs of simple setae surrounded by three simple setae on each side; opisthosomal region below anal opening with three setae; trochanter I with three large spatulate setae on anterior bor- der and a single smaller setae; inner region with three spatulate setae; setal placement on rest of segments as shown in Fig. 3; setal placement on legs II, III, and IV as shown in Figs. 3-6. Male (Figs. 7-12) anterior border of pro- podosoma similar to female projecting for- ward over gnathosoma with only four flat- 718 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5,6. Caeculus lewisi new species holotype fe- male, 3rd and 4th pair of legs. Figs. 3, 4. Caeculus lewisi new species holotype fe- male, Ist and 2nd pair of legs. tened crescent-shaped, curved setae; two pairs of lateral eyes similar in structure to female; propodosomal plate with six spat- ulate setae; median dorsal plate (Fig. 7) with seven spatulate setae, plate longer than wide similar to female; lateral median dorsal plates with three spatulate setae each; opis- thosomal transverse plate fused into single transverse plate across opisthosoma with five spatulate setae; opisthosoma with 10 spatulate setae scattered near posterior re- gion of opisthosoma; dorsal setae similar in structure to female with inner plumose re- gion; body heavily armored, wrinkled, dark brown in color; length 1.2 mm, width be- tween eyes 0.5 mm; opisthosoma wider than propodosomal region; ventral area below anterior projection of propodosomal plate without forked prominence found on fe- male; two large spine-like setae located on Fig. 7. Caeculus lewisi new species allotype male pronounced swellings between palps and dorsal view. VOLUME 92, NUMBER 4 719 Ue, Wi QGEr Guy 5 Mel CA, ry hi 4 4 7 Ww wm * 4 Hy Figs. 11, 12. Caeculus lewisi new species allotype male 3rd and 4th pair of legs. Fig. 8. Caeculus lewisi new species allotype male ventral view. chelicerae; chelicerae reduced; four spatu- late setae located below chelicera and be- tween palps bases; first coxa with four spat- ulate setae; second and third coxa with three 10 Figs. 9, 10. Caeculus lewisi new species allotype Fig. 13. Caeculus lewisi paratype nymph dorsal male Ist and 2nd pair of legs. view. ty we VAG { RRA Hi) ae Z {i WIEN al) edt \ TAN , gp tie { S\N Ka Wer X\ \\ 4 y)* y) (% if \ LYING SVU Fig. 14. view. Caeculus lewisi paratype nymph ventral spatulate setae; fourth coxa with two spat- ulate setae; six setae in center of venter forming a transverse line below fourth coxa (Fig. 8); genital plates each with six simple setae; ten simple small setae associated with genital opening; anal opening with two pairs of setae surrounded by three simple setae; opisthosomal region below anal opening with three simple setae; trochanter I with three large spatulate setae on anterior bor- der (Fig. 9); setal placement on rest of seg- ments as shown in Fig. 9; setal placement on legs II, III, 1V as shown in Figs. 10-12. Nymph (Figs. 13-18), anterior border of propodosoma projecting forward over gnathosoma with four flattened crescent shaped setae; two pairs of lateral eyes as- sociated with striated dorsal integument, two small setae are located on two finger-like projections of propodosomal plate; propo- dosomal plate with two simple, long, setae PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 16 Figs. 15, 16. Caeculus lewisi paratype nymph Ist and 2nd pair of legs. on anterior portion; propodosomal plate with eight spatulate setae, all aligned in a vertical row on lateral margin; propodoso- mal plate about as wide as long; lateral me- dian dorsal plate each with three spatulate setae, all aligned in a vertical row; median dorsal plate with six spatulate setae; plate longer than wide; opisthosomal transverse plates fused into single transverse plate across opisthosoma, with five spatulate se- tae; opisthosoma with five spatulate setae in a transverse row in posterior region of opisthosoma; ventral palpal tibia as shown in Fig. 13; first coxa with four simple setae and a single spatulate setae; second coxa with one simple seta; third and fourth coxa with two simple setae; center of venter be- tween fused third and fourth coxa with two pairs of simple setae; a transverse row of five simple setae located below fourth coxa; a reduced genital plate with three simple VOLUME 92, NUMBER 4 18 Figs. 17, 18. Caeculus lewisi paratype nymph 3rd and 4th pair of legs. setae, plate surrounded by pair of simple setae on either side (Fig. 14); anal plate with three simple setae, surrounded by three sim- ple setae on either side (Fig. 14); opistho- somal region with a pair of setae; setal place- ment of legs shown in Figs. 15-18. Larva (Figs. 19-23), anterior border of propodosoma projecting forward over gnathosoma; propodosomal plate reaching to anterior border with two spatulate setae; two pairs of lateral eyes; two spatulate setae located below lateral eyes and near posterior border of propodosomal plate; eight spat- ulate setae aligned in two rows of four each running from posterior border of propo- dosomal plate to opisthosomal region; ven- tral palpal tibia as shown in Fig. 20; mouth- parts reduced; apex of anterior region of gnathosoma with two setae; below reduced mouth parts are four simple setae; first coxa with two setae; second coxa without setae; Fig. 19. Caeculus lewisi larva dorsal view. Fig. 20. Caeculus lewisi larva ventral view. 122, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mulaik (1945). Mulaik’s work contains no keys, but his descriptions and figures are ) accurate. < / 23 a! 21 Figs. 21-23. Caeculus lewisi larval lst, 2nd, and 3rd pair of legs. third coxa with a single seta; venter with two pairs of setae, one pair located in pos- terior region of opisthosoma, other pair spatulate, located on margin of dorsal and ventral region of opisthosoma; setal place- ment of legs as shown in Figs. 21-23. Female holotype collected July 16, 1972 in Butte County 10 miles north of Newell, South Dakota near Willow Creek. Allotype male, nymphs, and larvae along with 22 specimens collected at type location. The holotype and allotype along with a nymph and larva will be deposited with the USNM. The following key constructed in 1972 from the literature was used to separate species of the genus Caeculus. It has been used with success to identify members of the family Caeculidae collected from Col- orado, North Dakota, South Dakota, New Mexico and Texas. It is here presented for use by others that do not have access to . Median dorsal hysterosomal plate with 2 pairs olispatulate|setacien ce eee eee ere 2 Median dorsal hysterosomal plate with more than’ 2 ipairsiofisetacieea- seme ari 3 . Leg I with 2 setae on femur with clavate tips, 2 on tibia, and 4 on pretarsus, all located on inner margin; outer margin seta smaller than inner margin setae and curved ............ ee C. clavatus 1905 Banks (California) Leg I with 2 setae on femur without clavate tips, 3 on tibia, and 4 on pretarsus, all located on inner margin; outer margin seta smaller than inner setae and curved .............. a e® ohes C. americanus Banks 1899 (California) . Median dorsal hysterosomal plate with 3 pairs Ofisetae erin ect Poet 4 Median dorsal hystersomal plate with more than’S ‘pairsiofisetae aeaiteiaeie eee 9 . Dorsal median hysterosomal plates I and II forming 2 pairs of small oval plates not con- nected at midline, each plate with 3 setae . . eee C. kerrulius Mulaik 1945 (Texas, Utah) Dorsal median hysterosomal plates I and II fused into single transverse plate across opis- thosomaliregiony ; 25 ere ee eee 5 . First opisthosomal transverse plate with 4 se- TAG ee ioieses Rrage apetenstele elses Sere 6 First opisthosomal transverse plate with 5 se- [ae Mier te ieee cetera pain bic ae Ais. 61.6 Gita 7 . First trochanter with 3 large curved spatulate setae on anterior border; specimens normally with an incrustation of minute sand-like par- ticles over most of body and legs ......... ert doce C. dortheae Mulaik 1945 (Arizona, Nevada, Texas) First trochanter with 2 curved setae on prom- inent tubercles on anterior border, one long (59 microns), club-shaped dorsomedially; without incrustation of particles .......... bo ited Coss eh C. hardyi Mulaik and Allred 1961 (New Mexico, Texas) . Second opisthosomal transverse plate with 6 setae; trochanter I with 2 slightly curved cy- lindrical setae, posterior setae much longer than anterior setae; on anterior margin of fe- mur is a single long seta, one on the patella, 3 on the tibia, and 4 on the pretarsus ..... BES ad MONON C. gretschi Mulaik 1945 (Texas) Second opisthosomal transverse plate with 4 Korte paar ame comttiac on sonsdoorBAnonas 8 . Trochanter I with 2 pairs of setae, posterior seta straight, slender, as long as width of tro- chanter, anterior seta smaller, curved, clavate; VOLUME 92, NUMBER 4 coxa with long slender, straight seta on an- terior border, anterior to seta is a small curved clavate seta; femur and patella each with one anterior seta; tibia with 2 setae, pretarsus with 4 setae .... C. hypopachus Mulaik 1945 (Texas) Trochanter I with a single curved spatulate seta On a prominent tubercle; femur and pa- tella with anterior setae on tubercles; tibia with 2 setae; pretarsus with 3 eA ort Sivesiors GARNI C. calechius Mulaik 1945 (Montana, Texas, Utah) . Propodosomal plate not projecting anteriorly over the gnathosomal tubercles and not cov- ering palps from above .................. 10 Propodosomal plate projects anteriorly over gnathosoma, covering gnathosomal, tubercles from above (if palps are observed it is due to mounting of specimen in a flattened position on the slide, specimens that are not on a slide will clearly show projection of propodosomal plate) . Median dorsal hystersomal plate with 13 se- tae arranged in a 4-4-5 sequence; left lateral metapodosomal plate with 7 setae, 2-3-2; night lateral metapososomal plate with 6 setae, 2- Dal sae C. mexicanus Mulaik and Allred 1961 (Texas) Median dorsal hysterosomal plate with less than 13 setae; left lateral metapodosomal plate with less than 7 setae; right lateral metapo- dosomal plate with less than 4 setae ....... 11 . Dorsal-lateral gnathosomal sensillae much expanded distally forming racket-like organs; posterior area of trochanter I set in slightly from edge Moa oO C. oregonus Mulaik and Allred 1961 (California, Oregon) (Higgins and Mulaik 1961 placed C. oregonus in the genus Procaeculus) Without above characters; left and right me- tapososomal plate with 5 setae; distally, pos- terior seta of trochanter I located on posterior edge ... C. brevis Mulaik 1945 (Arizona, Texas) . Opisthosomal transverse plates I and II form- ing 2 pairs of small oval plates not connected at midline; on transverse plate I are II setae on each oval plate and a single seta near each inner margin of oval plate; transverse plate II with 5 setae on each oval plate C. cremnicolus Enns 1958 (Missouri) Opisthosomal transverse plates I and II fused into single transverse plates across opistho- Somalirepionmeeern ae cette niece selene 13 . Median dorsal hysterosomal plate setae num- IDERUNE VEN Cera royr cet seictapa trees tneie crsversieere 14 Median dorsal hysterosomal plate setae num- Derievenly escice toes silent earauiais i) 14. Median dorsal hysterosomal plate with seven setae; dorsal plate of cephalothorax with two clavate setae near posterior corners; second transverse abdominal plate with 6 setae, each located on prominent humps on lateral mar- gins of this plate, with one clavate seta at mid- line of posterior margin of dorsum C. archeri Mulaik 1945 (Alabama, Tennessee) — Median dorsal hysterosomal plate with 9 se- tae; dorsal plate of cephalothorax with 4 pairs of setae, first pair located at anterior margin, second pair near anterio-lateral margin, third and fourth pair in postero-lateral region of plate: eee ee C. pettiti Nevin 1943 (Virginia) 15. Trochanter I with 2 spatulate setae on anterior and posterior border (4 setae); 10 median dor- sal hysterosomal plate setae C. tipus Mulaik 1945 (Texas, Utah) — Trochanter I with more than 4 spatulate setae on anterior and posterior border 16. Trochanter I with 3 spatulate setae on anterior and posterior borders (6 setae); 10 median dorsal hysterosomal plate setae; dorsal region of pretarsus without setae in midregion; ... .... C. valverdius 1945 Mulaik (Arizona, Texas) — Trochanter I with 3 spatulate setae on anterior and 2 on posterior border (5 setae); eight me- dian dorsal hysterosomal plate setae; dorsal region of pretarsus with setae in midregion C. lewisi n. sp. (South Dakota) ACKNOWLEDGMENTS We are especially grateful to J. K. Lewis, Professor emeritus South Dakota State Uni- versity, after whom we have named this new mite. This research was supported by the South Dakota Agricultural Experiment Sta- tion, SDSU, Brookings, project numbers H-277 and H-388, contribution no. 1716. LITERATURE CITED Banks, N. 1899. An American species of the genus Caeculus. Proc. Entomol. Soc. Wash. 4: 221-222. 1905. Descriptions of some new mites. Proc. Entomol. Soc. Wash. 7: 135-136. Crossley, D. A. and V. Merchant. 1971. Feeding by caeculid mites on fungus demonstrated with ra- dioactive tracers. Ann. Entomol. Soc. Amer. 64(4): 760-762. Enns, W.R. 1958. A new species of rake-legged mite from Missouri (Acarina:Caeculidae). Jour. Kan. Entomol. Soc. 31(2): 107-113. Higgins, H. G. and S. B. Mulaik. 1957. Another Cae- 724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON culus from southwestern United States (Caeculi- | Mulaik, S. B. and D. M. Allred. 1961. New species dae). Texas Jour. Sci. 9: 267-269. and distribution records of the genus Caeculus in McDaniel, B. 1979. How To Know the Mites and North America. Proc. Entomol. Soc. Wash. 56(1): Ticks. Wm. C. Brown Publishers, Dubuque, Iowa. 27-40. 319 pp. Nevin, R. R. 1943. Caeculus pettiti a new species of Mulaik,S. 1945. New mites in the family Caeculidae. mite from Virginia. Ann. Entomol. Soc. Amer. 36: Bull. Univ. Utah 35(17): 1-23. 389-393. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 725-731 BIONOMICS OF EV YLAEUS COMAGENENSIS (KNERER AND ATWOOD) (HALICTIDAE), A FACULTATIVELY POLYGYNOUS, UNIVOLTINE, BOREAL HALICTINE BEE S. W. T. BATRA Beneficial Insects Laboratory, Bldg. 476, United States Department of Agriculture, Beltsville, Maryland 2070S. Abstract. —Evylaeus comagenensis (Knerer and Atwood) (Hymenoptera: Halictidae) is a boreal carinate species with solitary to nearly unique semisocial or quasisocial behavior. In the Adirondacks of northern New York, it was at the southern limit of its range, which may be restricted by the deleterious effects of summer heat and drought on its brood. The brood was in exceptionally shallow subterranean clusters (combs) of delicate earthen cells. Its nest architecture, phenology, associates, and sociobiology are discussed. Key Words: There are about 2000 species of halictine bees worldwide (Sakagami et al. 1982). They include many solitary species, as well as species that share nests in a variety of social arrangements (Michener 1974). The genus Evylaeus consists of small, black, incon- spicuous bees that are closely related to La- sioglossum, and are included in this genus by some taxonomists. Evy/aeus, a primarily Palearctic genus, includes 81 species in the Western Hemisphere (Moure and Hurd 1987). Most research concerning the biol- ogy of Evylaeus has been performed in Eu- rope and Japan; only five North American species have been investigated (Moure and Hurd 1987, Packer et al. 1990a, b). Evylaeus comagenensis (Knerer and At- wood) belongs to the ‘carinate’ group, in which the females have a carina on the pos- terior edge of the propodeum (Svensson et al. 1977), and it is closely related to E. niger (Viereck). Bees in this group often construct their subterranean cells so that they form delicate earthen combs, surrounded by air- filled cavities. Several Evylaeus species be- nesting, quasisocial, semisocial, polylectic gin nesting each spring, when groups of overwintered females cooperatively con- struct each nest. This polygynous condition may be quasisocial if all females perform all duties, including oviposition; or it may be semisocial if there is a division of labor, with some females laying eggs and others performing other duties. In species of Evyla- eus that may facultatively begin nests in spring polygynously, the colonies continue to produce broods in the summer in an eu- social manner. Thus the nest-founding fe- males (foundresses) become egglayers (queens), while their daughters stay in their natal nests as workers, e.g. E. nigripes (Lep.) (Knerer and Plateaux-Quénu 1970) and E. linearis (Schenck) (Knerer 1983). Evylaeus comagenensis 1s unusual because it is uni- voltine, and thus does not have any eusocial phase. The only other univoltine, polygy- nous species is Dialictus problematicus (Bliithgen) from northern Japan (Sakagami et al. 1984). Strictly polygynous (quasisocial or semisocial) colonies of Hymenoptera are rare, and, therefore, E. comagenensis is of 726 unusual interest for comparative investi- gations of the probable evolution of insect societies. HABITAT I discovered nests of E. comagenensis in 1986 during an investigation of Andrena (Scrapteropsis) alleghaniensis Viereck, which shared the nesting site (Batra 1990). Nests were aggregated in an insolated south- easterly facing road cut in the Adirondack Mountains of northern New York (44°1 2’N; 73°55'W), at an elevation of 668 m (2035’). This may be the southernmost extension of the range of this boreal bee, which occurs from north of the Arctic circle at Inuvik, N.W.T. (Sakagami and Toda 1986) to On- tario. According to Knerer and Atwood (1964), it is not found south of 45°N, but Svensson et al. (1977, Fig. 23) illustrate a record from the Appalachian Mountains in Pennsylvania. Packer (pers. comm.) found E. comagenensis to be rare in southern On- tario but very common to the north. The soil in which the bees nested con- sisted of well-drained glacial outwash, a mixture of fine, loose, dustlike sand, peb- bles, and cobbles, without surface vegeta- tion. The surrounding area was covered with mature mixed northern hardwood and soft- wood forest. At the time that E. comage- nensis was active, species of Acer, Prunus, Amelanchier, and Viburnum, as well as for- est-floor herbs, were blooming. According to Knerer and Atwood (1964), this species is polylectic. I found pollen of many species in the provisions and in the digestive tracts of adults, which confirms their report. I carefully excavated nests in May and June, 1987 and 1989. Nest contents were preserved, reared, dissected, and evaluated, and foragers were collected to determine each bee’s social rank. I could not visit the site during the exceptionally hot, dry spring of 1988. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nest ARCHITECTURE, PHENOLOGY, AND ASSOCIATES Each of the 21 complete nests that I ex- amined contained a single brood-cell comb (Fig. 1). Most lacked tumuli and were closed at their entrances. Open nest entrances were irregular, cryptic, unguarded, and about 2.5 mm in diameter. The cluster of brood cells was very shallow, the topmost cells being 1.5-5.0 (X = 3.32) cm below the surface of the eroding sand. The lowest cells were 4.5- 7.5 (X = 6.0) cm deep. Clusters (brood combs) were |.5—4.0 (% = 2.6) cm high and 1.5—3.5 (% = 2.0) cm wide, and were without lateroids. They were supported in the 4-7 mm-wide cavities by several earthen pillars. Nests contained 1-12 (X = 5.5) cells. One nest that had only a single cell lacked a cav- ity, and a nest with two cells had an incip- ient cavity, indicating that the initial cells were made before the surrounding cavity was excavated. Nests that contained only one female had 1-7 (X = 3.6) cells and po- lygynous nests contained 6-12 (X = 9) cells. Somewhat sinuous tunnels, 4.5—7 (* = 5.5) mm in diameter, extended below the cell clusters to depths of 7.0-25.0 (¥ = 17.7) cm. Polygynous nests had 1-3 tunnels, but nests built by solitary females had only one tun- nel. The delicate brood cells were constructed of fine sand particles held in a stiff matrix of Dufour’s gland secretion. The thinnest parts of the walls of some cells were trans- lucent, and only 0.30 mm thick. The outline of each cell was visible on the surface of the brood comb. Cells (N = 22) were horizontal, of the usual halictine structure (Michener 1974), and 9.0-13.4 (x = 11.0) mm in length, and 4.5-5.0 (xX = 4.8) mm in maximum width. The cell entrances were 1.8-3.7 (X = 2.7) mm wide, and they were filled after Oviposition with a loose plug of soil that was 1.0-3.5 (< = 2.0) mm deep. Each arched, white egg was laid on a moist, yellow, roughly oval pollen and nectar mass 4.3-4.5 mm long, 3.0-4.3 mm wide, and VOLUME 92, NUMBER 4 Fig. 1. Three nests of Evy/aeus comagenensis in the sandy bank. The clusters of cells, each surrounded by an airspace, are in the sun-warmed layer near the surface. The bees will deepen the burrows below the clusters for use as hibernacula. 2.5-3.7 mm high. The provisions were lon- gitudinally grooved on top, and were sup- ported below on a wide pedestal as in E. linearis (Knerer 1983), which minimized contact with the cells. Nests were initiated in late May. During the first week of June, eggs, larvae and pupae were present, with medium-sized to large larvae and prepupae predominating. By June 10-11 (both years) oviposition had ceased, ovaries were regressing, and the brood in- cluded larvae and pupae. I removed these and reared them in plastic tissue-culture wells at 25°C. By June 24, all were prepupae to black pupae, and from June 30 to July 2, adults emerged. Adults were kept in a small, 728 screened cage outdoors (in Maryland), pro- vided with various cut flowers and sand for nesting. Males patrolled and attempted to mate at flowers, but did not inseminate fe- males (as determined by dissection). All adults disappeared by July 15. The females burrowed into the sand and exhibited a strong photophobic response when they were removed from their burrows in August. Their burrows were initiated beneath ob- jects on the surface of the sand, extended to the bottom of the cage, and each contained up to three females. Hibernation in nature occurred at the ends of the burrows that extended below the cell clusters. I excavated some nests on October 30, 1987, when soil temperatures were 12°C at the surface, 10°C at 2.5 cm, 8°C at 7.5 cm, 5°C at 15 cm, and 3°C at 20 cm where bees were found. Below that level, the earth was frozen and it was not possible to ex- cavate or locate bees. According to Saka- gami et al. (1984) hibernating Evy/aeus and other halictine bees are capable of super- cooling. The cells of E. comagenensis were con- structed unusually near the soil surface. Such shallow placement maximized heating dur- ing the relatively few sunny days in May and early June (17%, 37%, and 24% of days in 1986, 1987 and 1989). On sunny days, the afternoon soil temperature at 2.5 cm was 10—15° warmer than soil at 15 cm, and 2- 5° warmer than soil at 7.5 cm (3 measure- ments). When the air temperature was 32°, soil at 2.5 cm deep was 40°, and at 7.5 cm deep it was 35°. Packer (1990a, b) found that pebbles or stones just above shallow cell clusters in level soil in Nova Scotia en- hance heating. In 1989, there were fewer nests (3 nests/ m7’) than in 1987 (9.6/m72). Probably the pro- longed heat (many days above 32°) and the June drought of 1988 resulted in high brood mortality. However, the population size of Andrena alleghaniensis at this site appeared normal (this species makes cells at 13-23 cm depth, in cooler, moister soil). Augo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON chlorella striata (Prov.), another halictine that makes shallow combs, similarly suf- fered high pupal mortality during an un- usually hot, dry summer (Packer et al. 1990b). Because E. comagenensis is uni- voltine, it has no opportunity to rapidly re- cover population levels. Other species of Evylaeus that make shallow brood cell clus- ters in spring make additional cells at deeper levels for summer broods, thus, they can repopulate more rapidly. Examples are E. duplex (Dalla Torre) (Sakagami and Haya- shida 1961, 1968, Sakagami et al. 1984), E. affine (Smith) (Sakagami et al. 1982) and E. nigripes (Knerer and Plateaux-Quénu 1970). A close relative, Evylaeus nupricola (Saka- gami), is a solitary, univoltine alpine relict species that also makes shallow cell clusters (Sakagami 1988); its distribution may sim- ilarly be limited by heat or drought. In 1987, 82% of nests also included fe- males and larvae of the scutacarid mite, /m- paripes apicola (Banks) in some of the cells that contained eggs, larvae, and prepupae. In 1989, only 14% of nests included mites. According to Eickwort (1979), these mites feed on the feces of the larvae of various halictid and andrenid bees. Other associates included unidentified nematodes and fungi on feces in cells that contained healthy pre- pupae, and a conopid larva (probably The- cophora occidensis (W\k.), Knerer and At- wood 1967) in the abdomen of a dead female. Species of Sphecodes (Halictidae), Leucophora (Anthomyiidae), and Phrosi- nella aurifacies Downes (Sarcophagidae) followed returning foragers, entered nests, or both, but none were found in cells. Larval rhipiphorids and larval Strepsiptera were on adult bees or in their crops, but none were recovered from cells or brood. SOCIOBIOLOGY One possible effect of the low population density following the 1988 drought was the lack of polygynous nests. All of the nine nests that I examined in 1989 were occupied by a single female; however, in 1987, 64% VOLUME 92, NUMBER 4 of 12 nests were polygynous. Thus it appears that E. comagenensis was only facultatively polygynous. Possibly the solitary foundress- es were the only survivors among sisters that would have nested together following a favorable year. Perhaps when competition for suitable nest sites became less, few fe- males joined already occupied nests. In the strictly solitary species, E. oenotherae (Ste- vens), some females attempt to join other nesting females, but they are rejected (Kner- er and MacKay 1969). Similarly, solitary E. duplex foundresses are aggressive toward strangers, although they later cooperate eu- socially with their own daughters (Sakagami and Hayashida 1961). Table 1. 729 All females of E. comagenensis were in- seminated (N = 56) and male production was high in my study. Thus this bee resem- bled a solitary species (see Packer and Kner- er 1985). In 1987 and 1989, broods were 38% and 75% male, respectively. Plateaux- Quénu (1967, Fig. 1) indicates a 50% sex ratio for this species. Fluctuating sex ratios occur also in E. duplex (Sakagami and Ha- yashida 1961). There was neither clear division of labor nor size difference among females that shared the polygynous nests. I measured head width, ovarian and Dufour’s gland en- largement, front wing nicks, and degree to which the mandibles were worn down by Contents of polygynous nests of E. comagenensis in 1987. Head width units (bee size): | mm = 6.4 units. The ovarian development of each female is indicated thus: A, with 2 or more eggs ready to lay; B, with large ovaries, | egg ready to lay; C, with moderately developed or regressing oocytes; D, with slight enlargement of oocytes; E, with no oocyte enlargement. Nicks in forewings counted (if x, wings badly damaged). Mandibular wear ranges from 1, with unworn, sharp mandibles, to 5, mandibles worn to stubs, lacking a notch. Adults Head Width (in Units), Ovary, Wing Nicks, Mandible Wear Eggs Nest Small Medium Number of Immatures Larvae Pupae Large Prepupae Male Female Date 1. 6 bees 13.5, 1355: 13.0, 13.0, 125; 12.0, 14.0, 13.5, 13.0, 13.0, 13.0, 14.0, 12.0, 11.0, 14.0, 13.0, 12.5, 12.0, 14.0, 113255 13.0, 125; 12.0, A o ENE AaN Nw srs 1 to SE uDEeS NUNN FWNWYNW Ww . 3 bees _ 4. 4 bees B 9N0U UU> BONEN PoRAe paw Ww wn . 2 bees iS) Sd) pK 00" OO Ww t (forager) 6. 3 bees ae) mimo Pas 359 oh 2 3 June | June 4 June 4 i) nn nN June 7 June 7 June 10 730 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON digging, in order to detect possible castes (Table 1). Bees that had eggs ready to lay (N = 12) had mean head widths only 1.02% larger than bees (N = 9) with undeveloped ovaries (bees with partly enlarged oocytes were omitted from these comparisons). In 1987, 29% of all females had large oocytes, but in 1989, 56% of females, all of which were alone in nests, had eggs ready to lay. There was no consistent difference in wear between egg layers and non-egg layers in polygynous nests. The wings of the egg lay- ers had 1-9 (x = 3.3) nicks, indicating prob- able foraging activity, and their mandibular wear was 2-5 (X = 4.0). Non-egg layers had 1-8 (< = 3.8) wing nicks and mandibular wear of 3-5 (X = 3.8). Nests that were occupied by solitary fe- males in 1989 had 1-7 (x = 3.6) cells; in 1987, solitary females made 2-5 (X = 3.3) cells. Polygynous nests with 3 females (1 egg layer) had 3-6 (X = 4.5) cells; a nest with 4 females (1 egg layer) had 10 cells; a nest with 5 females (1 egg layer) had 12 cells; and a nest with 6 females (4 egg layers) had 6 cells. It was difficult to estimate productivity be- cause of the probability that most or all of the females in polygynous nests may be- come capable of oviposition at some time. Oophagy and drifting of females to adjacent nests may also occur. It thus appears that polygynous nests more closely approach the democratic quasisocial arrangement than the semisocial pattern of behavior, in which queens and workers are well differentiated. Packer et al. (1990a) found 1-4 females per nest in Nova Scotia, where there appeared to be some division of labor in polygynous nests. Now that many researchers worldwide have begun to investigate the intricacies and complexities of halictine sociobiology, pre- viously unknown and sometimes unique patterns are revealed, and previously known ones are reclassified. Evylaeus comagenen- sis has pioneered social and architectural systems that are particularly well suited to the efficient exploitation of resources during the cool, rainy, and short boreal flowering season. ACKNOWLEDGMENTS I thank L. Packer for identifying the bees (C.W.T.), for sending me his unpublished manuscripts, and for reviewing the manu- script. G. C. Eickwort kindly identified the mites, N. E. Woodley identified flies, and E. M. Barrows also reviewed the manu- script. LITERATURE CITED Batra, S. W. T. 1990. Bionomics of a vernal solitary bee, Andrena (Scrapteropsis) alleghaniensis Vie- reck in the Adirondacks of New York. J. Kansas Entomol. Soc. (In press.) Eickwort, G. C. 1979. Mites associated with sweat bees (Halictidae). Recent Adv. Acarology 1: 575- 581. Knerer, G. 1983. The biology and social behavior of Evylaeus linearis (Schenck) (Apoidea: Halictinae). Zool. Anz. Jena 211: 177-186. Knerer, G. and C. E. Atwood. 1964. Further notes on the genus Evylaeus Robertson (Hymenoptera: Halictidae). Can. Entomol. 96: 957-962. 1967. Parasitization of social halictine bees in southern Ontario. Proc. Entomol. Soc. Ontario 97: 103-110. Knerer, G. and P. MacKay. 1969. Bionomic notes on the solitary Evylaeus oenotherae (Stevens) (Hy- menoptera: Halictinae), a matinal summer bee visiting cultivated Onagraceae. Can. J. Zool. 47: 289-294. Knerer, G. and C. Plateaux-Quénu. 1970. The life cycle and social level of Evylaeus nigripes (Hy- menoptera: Halictinae), a Mediterranean halictine bee. Can. Entomol. 102: 185-196. Michener, C. D. 1974. The Social Behavior of the Bees. Harvard University Press, Cambridge, Mass. 404 pp. Moure, J. S. and P. D. Hurd, Jr. 1987. An Annotated Catalog of the Halictid Bees of the Western Hemi- sphere (Hymenoptera: Halictidae). Smithsonian Inst. Press, Washington, D.C. 405 pp. Packer, L. and G. Knerer. 1985. Social evolution and its correlates in bees of the subgenus Evylaeus (Hy- menoptera: Halictidae). Behav. Ecol. Sociobiol. 17: 143-149. Packer, L., V. Jessome, C. Lockerbie, and B. Sampson. 1990a. The phenology and social biology of four sweat bees in a marginal environment: Cape Bre- ton Island. Can. J. Zool. (In press.) Packer, L., B. Sampson, C. Lockerbie, and V. Jessome. 1990b. Nest architecture and brood mortality in VOLUME 92, NUMBER 4 four species of sweat bees (Hymenoptera: Halic- tidae) from Cape Breton Island. Can. J. Zool. (In press.) Plateaux-Quénu, C. 1967. Tendances évolutives et degré de socialisation chez les Halictinae (Hym., Apoidea). Ann. Soc. Entomol. Fr. (N.S.) 3: 859- 866. Sakagami, S. F. 1988. Bionomics of the halictine bees in northern Japan. IV. Lasioglossum (Evylaeus) nupricola sp. nov., a climatic relic. Kontya 56(2): 337-353. Sakagami, S. F. and K. Hayashida. 1961. Biology of the primitive social bee, Halictus duplex Dalla Torre III. Activities in spring solitary phase. J. Fac. Sci. Hokkaido Univ. Ser. VI 14: 639-682. 1968. Bionomics and sociology of the sum- mer matrifilial phase in the social halictine bee, Lasioglossum duplex. J. Fac. Sci. Hokkaido Univ. Ser. VI 16: 413-513. 731 Sakagami, S. F., Y. Hirashima, Y. Maeta, and T. Mat- sumura. 1982. Bionomic notes on the social hal- ictine bee, Lasioglossum affine (Hymenoptera, Halictidae). Esakia 19: 161-176. Sakagami, S. F., K. Hoshikawa, and H. Fukuda. 1984. Overwintering ecology of two social halictine bees, Lasioglossum duplex and L. problematicum. Res. Popul. Ecol. 26: 363-378. Sakagami, S. F. and M. J. Toda. 1986. Some arctic and subarctic solitary bees collected at Inuvik and Tuktoyaktuk, N.W.T., Canada (Hymenoptera: Apoidea). Can. Entomol. 118: 395-405. Svensson, B. G., A. W. Ebmer, and S. F. Sakagami. 1977. Lasioglossum (Evylaeus) boreale, a new Halictinae (Hymenoptera: Apoidea) species found in northern Sweden and on Hokkaido, Japan, with notes on its biology. Entomol. Scand. 8: 219-229. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 732-735 A NEW SPECIES OF ASPHONDYLIA (DIPTERA: CECIDOMYIIDAE) ON BORRICHIA (ASTERACEAE) FROM FLORIDA ANTHONY M. Rossi AND DONALD R. STRONG Department of Biological Science, Florida State University, Tallahassee, Florida 32306. Abstract. —A new species, Asphondylia borrichiae, a gall former on the apical growing region of the stems of sea daisy, Borrichia frutescens (L.) DC. (Asteraceae), is described with illustrations. Adults from A. borrichiae were reared from galls collected year-round from the salt marshes along the Gulf coast of Wakulla County, Flonda. Key Words: gall former, sea daisy The sea daisy, Borrichia frutescens (L.) DC. is a fleshy salt-tolerant herb that grows in discrete patches along Florida’s northern Gulf coast (Bell and Taylor 1982, Stiling et al. in prep). The apical growing region of B. frutescens is attacked by an undescribed species of Asphondylia (Diptera: Cecido- mylidae) (Gagné 1989). According to Gagné (1989), the gall is known from North Car- olina, Florida, and Mexico. We are cur- rently studying this species in both the lab- oratory and in the field (Stiling et al. in prep.), and describe the gall midge to provide a name for our further studies. While many species of Asphondylia are known, there is no modern revision with keys for identifi- cation. Further, most species have been de- scribed from adults only, a stage that is re- markably homogeneous in the genus (Gagné, pers. comm.). Most cecidomyiids are mono- or oligophagous (Gagné 1989), and because no species has been described from B. fru- tescens, we regard this gall midge from Bor- richia as a new species. This paper describes the new species with the guidelines used in Hawkins et al. (1986). The figures should allow anyone to identify larvae and pupae of this species from Borrichia and compare it with other species, particularly those on Asteraceae. MATERIALS AND METHODS Mature galls lacking emergence holes were collected, approximately once a month, from May to August, 1989 from seven sites around the Oyster Bay area of Wakulla County, Florida. Galls were taken to the laboratory and placed in 25 dram vials with a moist piece of filter paper. The vials were checked daily and any emerged flies were immediately placed in 70% ETOH. A sub- sample of adults was used in the following species description. In addition, several galls were immediately dissected upon return to the laboratory and third instar larvae and pupae were extracted for illustration. To assess gall characteristics, the maxi- mum width of ten mature green galls with emergence holes was measured at each of the seven study sites during mid-July. On 5 September, 112 additional galls were col- lected from the study sites and dissected to estimate clutch size per gall by counting the number of chambers within the gall. Asphondylia borrichiae Rossi and Strong, NEw SPECIES Adult.— Color: Eyes black, scutum dark brown, abdomen light brown. Antenna: fe- males, mean ratio of scape: pedicel, 2.2:1 (N = 10); mean ratio of first flagellomere : VOLUME 92, NUMBER 4 scape, 2.3:1 (N = 10); males, 2.5:1 (N = 10) and 2.2:1 (N = 10) respectively. First fla- gellomere cylindrical, flagellar segments successively shortened distally. Palpal seg- ments bearing many setae, first segment tiny; second and third segments elongate, 2nd segment wider than 3rd; females, mean ra- tio of Ist, 2nd, and 3rd palpal segments, 1:2: 5.8 (N = 10), males, 1:2:5.4 (N = 10). Wing length: males, range: 2.6-3.6 mm (N = 38), mean = 3.2 + 0.04 (SE) mm; females, range: 2.8-3.7 mm (N = 36), mean = 3.3 + 0.04 (SE) mm. Scutum with dorsocentral row of setae single to double approximately 13 from the anterior margin of scutum. Tarsal claws of similar shape among legs and between sexes and approximately as long as empo- dia. Mean ratio of ovipositor from base to tip of rigid shaft 2.8-times (N = 10) length of sternite 7. Pupa (Figs. 1-3).—Antennal horns curved ventrally; serrated along interior margin. Both upper and lower frontal horns pointed anteriorly. Upper frontal horn bifid; lower frontal horn trifid. Posterior abdominal ter- gal spines on segments 3-8 separated from slightly smaller, anterior row of spines. Larva (third instar) (Figs. 4, 5).—Cream to yellow in color. Spatula as in Fig. 4, quad- ridentate, the two inner teeth shorter than the outer pair. Four lateral papillae present on either side of the spatula. Terminal seg- ment with one pair of dorsal papillae. Type material. — All specimens deposited in National Museum of Natural History, Washington, D.C. Holotype.—Pupa, ex Borrichia frutescens gall, 5 May 1989, St. Mark’s Wildlife Refuge, Wakulla County, Florida, A. M. Rossi. Paratypes: 10 females, 10 males, 5 pupae, 5 larvae, all with same data as holotype. Galls.— These are located on the stems at the apical growing point, usually with one or more swollen pairs of leaves associated with them (Fig. 6). Galled shoots are often prevented from flowering, but flowering is occasionally seen above the galled growing point. Post-emergence galls and associated Figs. 1-3. Asphondylia borrichiae. 1 and 2, Pupal head: 1, ventral; 2, lateral; 3, Pupal abdomen, dorsal. Scale: = 100 um. leaves rot and turn black. After the apical growing region rots, the plant sends out a new shoot from a lower meristem. Live galls were collected year-round from Oyster Bay, although gall density varied greatly through- out the year. Adults were reared every month in northwest Florida (Stiling et al. in prep). The maximum diameter of mature galls averaged 0.9 + 0.02 (SE) cm (N = 70; range: 0.645-1.43). In addition, the mean number 734 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4, 5. Asphondylia borrichiae. 4 and 5, Larval parts: 4, spatula; 5, terminal segment, dorsal. Fig. 6. Gall of Asphondylia borrichiae on its host plant, Borrichia frutescens; note: puparium present at 11 o’clock position. VOLUME 92, NUMBER 4 of chambers per gall was 1.91 + 0.08 (SE) (N = 112; range: 1-5). ACKNOWLEDGMENTS We are grateful to Peter Stiling for assis- tance with the field work and Lawrence Abele for the use of his dissecting microscopes. We thank Sandra Silvers and Kim Riddle for their skill and patience while providing us with the electron micrographs. We es- pecially thank Raymond J. Gagné who gra- ciously reviewed and improved the manu- script. In addition, we thank Larry Bird for his inspiration over the past ten years. This 735 research was supported by National Science Foundations Grant BSR-8703416 to D.R.S. LITERATURE CITED Bell, C. R. and B. J. Taylor. 1982. Florida Wild Flow- ers and Roadside Plants. Laurel Hill Press. Chapel Hill, N.C. 308 pp. Gagné, R. J. 1989. The Plant Feeding Gall Midges of North America. Cornell University Press. Ith- aca, N.Y. xi and 356 pp. and 4 pls. Hawkins, B. A., R. D. Goeden, and R. J. Gagné. 1986. Ecology and taxonomy of the Asphondylia spp. (Diptera: Cecidomyiidae) forming galls on Atri- plex spp. (Chenopodiaceae) in Southern Califor- nia. Entomography 4: 55-107. Stiling, P. D., A. M. Rossi, and D. R. Strong. In prep. Insects reared from galls of Borrichia frutescens in north Florida salt marshes. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 736-738 NOTE Examination of Some Sensory Organs on the Head of Last Instar Larvae of the Lesser Cornstalk Borer, Elasmopalpus lignosellus (Zeller) (Lepidoptera: Pyralidae) The lesser cornstalk borer, E/asmopalpus lignosellus (Zeller) (Lepidoptera: Pyralidae) is an economic pest of several crops in the southeastern U.S., including peanut, sor- ghum, soybeans, and small grains (Funder- burk et al. 1987. J. Entomol. Sci. 22: 159- 168). Outbreak population configurations typically occur in hot, dry weather in pea- nuts grown in sandy soils (Mack et al. 1987. Peanut Science 14: 61-66). Larvae are usu- ally subterranean, and may feed on pegs (gynophores), developing pods, or on plant tissue in the root-hypocotyl region. It is probable that E. lignosellus larvae chemically locate their host plants and ac- ceptable plant parts (Huang and Mack 1989. Environ. Entomol. 18(5): 763-767). It would be useful to document the sensilla in E. lig- nosellus available for host-finding, since it is the sensilla that would be involved in chemoreception. This note documents the antennal and maxillary sensilla on the head of E. lignosellus larvae. Scanning electron micrographs were made Fig. 1A-D. Scanning electron micrographs of E. /ignosellus. A, Ventral view of the head showing the antenna (A), and sensilla styloconica on the maxillary galea (S) and maxillary palpus (P). 92x. B, Antenna with three sensilla basiconica (arrows). 850. C, Lateral (L) and medial (M) sensilla styloconica on the maxillary galea. 4600. D, Maxillary palpus with types (L, longer; S, shorter) of sensilla basiconica on the tip. 2900. VOLUME 92, NUMBER 4 Fig. IE-G. E, Long type of sensilla basiconica on the maxillary palpus. 8500 x. F, Short type of sensilla basiconica on the maxillary palpus. 12,700x. G, A maxillary palpus with only three sensilla basiconica on its tip instead of eight. 2500 x. of the antennae, maxillary palpi, and the sensilla styloconica on the maxillae of E. lignosellus. Larvae were taken from a lab- oratory colony, immobilized with CO,, and killed in ethanol. The heads were severed 737 and successively dehydrated in 70%, 85%, 95%, and 100% ethanol, with 5 min in each concentration. Heads were immersed in hexamethyldisilizane for 5 min and then air- dried at room temperature. Tissue was mounted on stainless steel stubs and sputter coated with gold for microscopy. The ventral view of the head of E. lig- nosellus (Fig. 1A) was similar to those of other lepidopterous larvae (Schoonhoven 1987. In Chapman et al. 1987. Perspectives in chemoreception and behavior. Springer- Verlag, NY), with each antenna possessing three sensilla basiconica (Fig. 1B). Each maxillary galea typically bore two sensillae styloconica which were nonsocket pegs with an apical papilla (Fig. 1C). Eight sensillae basiconica, with five being longer than the others, were observed on the tip of the max- illary palpi (Fig. 1D). The long sensillae basiconica had a smoother surface than the smaller ones and a small peg at their distal end (Fig. 1E), while the shorter ones were blunt with creased surfaces (Fig. 1F). It is interesting to note that not all maxillary palpi bore eight sensillae basiconica (Fig. 1G). The antennal and maxillary sensillae of a few lepidopterous species have been ex- amined, including the tobacco hornworm Manduca sexta (L.) (Hanson and Detheir 1973. J. Insect Physiol. 19: 1019-1034), the silkworm Bombyx mori (L.) (Morita and Yamashita 1961. J. Exp. Biol. 38: 851-861), the spruce budworm Choristoneura fumife- rana (Clemens) (Albert 1980. Can. J. Zool. 58: 842-851), and the darksided cutworm Euxoa messoria (Harris) (Devitt and Smith 1982. Int. J. Insect. Morph. Embryol. 11: 255-270). All of these insects are terrestrial. The number and morphology of the sensil- lae on the antennae and maxillary palpi of E. lignosellus are very similar to those of previously described larvae. The morphol- ogy of the sensillae of E. lignosellus, which is primarily subterranean, do not differ from 4, primarily terrestrial larvae. X. P. Huang and T. P. Mack, (7PM) De- 738 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON partment of Entomology and Alabama Ag- ricultural Experiment Station, 301 Fun- chess Hall, Auburn University, Alabama 36849-5413. Alabama Agricultural Exper- iment Station Journal Series no. 17-89236P; (XPH) Department of Plant Protection, Bei- jing Agricultural University, 100094 Bei- jing, People’s Republic of China. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 739-745 THE TRIBE STRONGYLOGASTERINI (HYMENOPTERA, TENTHREDINIDAE) FROM TAIWAN TIKAHIKO NAITO Entomological Laboratory, Faculty of Agriculture, Kobe University, Rokko, Kobe, 657 Japan. Abstract. —The tribe Strongylogasterini is represented by nine species belonging to three genera in Taiwan. Two new species, Strongylogaster nantouensis and Pseudohemitaxonus taiwanus, and the males of Strongylogaster fulva Naito and Huang and S. formosana (Rohwer) are described and illustrated. Three species, Strongylogaster fulva Naito and Huang, Hemitaxonus nigroorolis (Malaise) and H. alboorolis (Malaise) are newly recorded from Taiwan. Two genera, Canonarea Malaise and Trearea Malaise are new synonyms of the genus Hemitaxonus. The genus Pseudohemitaxonus is a new record in Taiwan. A key is given to separate the genera and species of the tribe from Taiwan. Key Words: Naito (1975) discussed the monophylety of five related genera, Eriocampidea, Hemi- taxonus, Pseudohemitaxonus, Nippono- rhynchus and Adelesta, in the subfamily Se- landriinae. The Strongylogasterini, one of five tribes of the subfamily, consists of these five genera and their two related genera, Strongylogaster and Canonias. The mor- phological characters may be highly vari- able in this tribe (Smith 1969), but the members share the following characters: rather long and slender sawflies exclusively associated with ferns; antenna 9-segmented with flagellar segments gradually reduced in length towards apex; fore wing with vein Rs+M curved towards stigma and anal cell usually with suberect crossvein, if absent, prepectus represented by raised shoulder; hind wing with two closed middle cells; and tarsal claw slender, slightly bent at apex. In Taiwan, this tribe was known by four species belonging to two genera (Takeuchi 1941): Strongylogaster lineata (Christ), S. formosana (Rohwer), S. abdominalis (Ta- keuchi) and Hemitaxonus formosanus Ta- Strongylogaster, Hemitaxonus, Pseudohemitaxonus keuchi. The present study, based on speci- mens collected by some _ Japanese entomologists and specimens deposited in the Taiwan Agricultural Research Institute, revealed the occurrence of nine species in three genera, Strongylogaster, Hemitaxo- nus, and Pseudohemitaxonus. Of these, one species of Strongylogaster and one species of Pseudohemitaxonus are new to science, and another species of Strongylogaster and two species of Hemitaxonus are newly re- corded from Taiwan. Two genera, Canona- reaand Trearea described by Malaise (1947) based on species from Burma, are here re- garded as synonyms of Hemitaxonus. KEY TO GENERA AND SPECIES OF THE STRONGYLOGASTERINI KNOWN FROM TAIWAN 1. Posttergite about as wide as scutellum. Anal cell of hind wing sessile or with short petiole at most as long as greatest breadth of cell. Pro- podeum normal, not excised at center ....... 2 — Posttergite about 1.5 times as wide as scutel- lum. Anal cell of hind wing with long petiole about 1.5* greatest breadth of cell. Propo- 740 tN deum deeply and broadly excised at center . . _.(Pseudohemitaxonus Conde) P. taiwanus n. sp. Prepectus represented by raised shoulder, sep- arated from mesepisternum by furrow. Anal cell of fore wing without crossvein (if present, sawsheath bifid at apex in dorsal view) 5 .(Strongylogaster Dahlbom) 3 Prepectus flat, represented by distinct sclerite, separated from mesepisternum by suture. Anal cell of fore wing with suberect crossvein . (Hemitaxonus Ashmead) 7 Head vith large separate punctures. Frontal area in form of raised platform, not surrounded by distinct raised carina. Tarsal claw with large inner tooth about 2x outer one. Sawsheath with large leaflike scopa ...... ‘ ne ti Head without large punctures. Frontal! area surrounded by sharp raised carina. Tarsal claw simple or with very small inner tooth. Saw- sheath with small apical projection or simple, not divided at apex in dorsal view (Fig. 4) 5 . Propodeum reticulate and dull. Clypeus nar- rowly emarginate in front to depth of about ' its medial length. Head and thorax black in female. Second tergite rufous (sometimes with black mark) in male ... Strongylogaster lineata Christ Propodeum smooth and polished. Clypeus broadly emarginate in front to depth of about Ys its medial length. Head and thorax brownish in female. Second tergite black with apical mar- gin rufous in male : Strongylogaster fulva Naito and Fluane Anal cell of hind wing with petiole. Tarsal claw simple, without inner tooth. Sawsheath divid- ed at apex, with small apical projection. Hind femur entirely yellowish : Anal cell of hind wing sessile. Tarsal claw with small inner tooth. Sawsheath simple, not di- vided at apex in dorsal view (Fig. 4). Basal half of hind femur black . : beets Strongyi logaster? nantouensis Nn. sp. Clypeus white. Abdomen black to dark brown . Strongylogaster formosana (Rohwer) Ghpets rial to dark brown. Abdomen with 2nd to 6th terga rufous . Strongylogaster abdominalls (Takeuchi) Anepineron without membranous area. Anal cell of hind wing sessile. Tarsal claw with small inner tooth. Abdomen black with 3rd to Sth or 6th terga rufous Anepimeron with large membranous area. Anal cell of hind wing with petiole. Tarsal claw sim- ple, without inner tooth. Abdominal terga usu- ally entirely rufous .. Hemitaxonus formosanus Takeuchi Clypeus white. Hind femur yellow. Malar space PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON about as long as diameter of front ocellus ... Hemitaxonus albooralis (Malaise) — Clypeus black. Hind femur blackish with apical half yellow. Malar space about a half of di- ameter oftront ocellusiaae nese eee Genus Strongylogaster Dahlbom This genus is represented by 39 species mainly in the temperate zone of the north- ern hemisphere: 11 species in North Amer- ica, 8 species in Europe and 26 species in eastern Asia (Naito and Huang 1988). The following five species occur in Taiwan, in- cluding a new species and a newly recorded species. Strongylogaster lineata (Christ) Strongylogaster lineata: Enslin, 1914, p. 205; Takeuchi, 1941, p. 243; Zhelochovtsev, 1951, p. 152; Benson, 1968, p. 134; Nai- to, 1980, p. 400. Specimens examined.—2 2 2 6, Tungpu, Nantou Hsien, 10.iv.1965, T. Saigusa; | 4, Meifeng, Nantou Hsiesn, 21.iv.1978, T. Niisato; 2 6, Pilushenmu (2200 m), Hualien Hsien, 29.11.1981, T. Shimomura; | 2, Howangshan, 7.1v.1973, C. C. Lo; 1 9, Ali- shan, Chiayi Hsien, 8.iv.1965, T. Shirozu; 1 6, Alishan, 9.iv.1965, Y. Hirashima. Distribution.— Taiwan, Japan, Korea, Kuriles, Siberia, Caucasus, Turkey, Iran, Europe. Remarks.—The coloration of the abdo- men is significantly variable in both sexes of this widespread species, but it is rather uniform in each sex of specimens from Tai- wan. The male abdomen is rufous except for the black basal two segments, and the female abdomen is black with a narrow yel- low band on the apical margins of the 2nd to 8th terga. Strongylogaster fulva Naito and Huang (Fig. 1) Strongylogaster fulva: Naito and Huang, 1988, p. 41. Male.—Length, 8.5-9.5 mm. Similar to VOLUME 92, NUMBER 4 female in morphological structure but quite different in coloration: Head and thorax black with labrum, labial and maxillary pal- pi and tegula yellowish. Legs light brown; coxae black except for extreme apices; tro- chanters partly brown. Abdomen rufous; propodeum and basal part of 2nd tergite black; 2nd to 5th terga with black mark on lateral sides. Penis valve as in Fig. 1. Specimens examined.—1 6, Kuantao- shan, Miaoli Hsien, 9.iv.1984, C. C. Lo; 1 6, Kuantaoshan, 9.v.1984, C. C. Lo; 4 4, Howangshan, 7.iv.1984, C. C. Lo; 1 2, Sung- kang, Nantou Hsien, 4.v.1984, C. C. Lo. Distribution.— Taiwan, South China. Remarks.—This species is newly record- ed from Taiwan. It was described from fe- males from Tibet and Sichuan, China, by Naito and Huang (1988). The male, which was unknown, is similar to the male of S. lineata but is distinguished from the latter by the broadly emarginate clypeus, the shin- ing thorax and propodeum, and the partly rufous 2nd tergite. A female specimen from Taiwan is paler than those from Tibet or Sichuan. The Taiwanese specimen is light brown except for the black Sth to 9th an- tennal segments and the large black mark on the mesonotal lobe, while the Chinese specimens are dark brown to black with var- ious amounts of orange. Strongylogaster nantouensis New SPECIES (Figs. 2, 4, 5) Female.—Length 9.0 mm. Black; apical margin of labrum, maxillary and labial pal- pi, tegula and apical margin of abdominal segments yellowish. Legs black to dark brown; apices of fore and mid coxae, apical half of hind coxa, trochanters, apices of fore and mid femora, apical third of hind femur, basal third of fore and mid tibiae and basal *%, of hind tibia yellowish. Wings hyaline with apical margins somewhat dark. Head shining, with close, minute punc- tures; frontal area with many longitudinal wrinkles; postocellar area smooth. Labrum 741 flat, gently rounded in front. Clypeus nearly flat, shallowly emarginate in front to depth of about % its medial length. Median fovea represented by small, elongated pit. Lateral fovea open below, with indistinct crest above. Frontal area concave, surrounded by narrow raised carina laterally and by broad, dull carina anteriorly. Interocellar furrow opening out towards front ocellus. Postocel- lar furrow represented by deep line. Vertical furrow deep, convergent toward front ocel- lus. Postocellar area convex; breadth : length = 23:13. Vertex with indistinct convex area at outside of lateral ocellus. Post genal ca- rina developed on lower hind margin. Malar space narrow, about 3x diameter of front ocellus. Antenna about 2.2 breadth of head; relative lengths of segments about 8: 5:24:23:23:18:17:16:14. Thorax smooth and shining; prescutum with minute and close punctures; lateral and posterior margins of scutellum with several large and separate punctures; prepectus represented by nar- row, raised shoulder. Anal cell of fore wing without crossvein. Anal cell of hind wing sessile. Tarsal claw with very small inner tooth. Abdomen (including propodeum) with fine microsculpture. Sawsheath short, not divided at apex in dorsal view, truncate at apex in lateral view (Fig. 4). Saw as in Fig. 5. Male. Length, 7.5—8.0 mm. Similar to fe- male except pronotum with upper angle yel- low and vertical furrows subparallel with each other. Penis valve as in Fig. 2. Holotype.—Female, Nanshanchi, Nan- tou Hsien, 21.111.1979, A. Shinohara (De- posited in the Entomological Laboratory of Kobe University). Paratypes.—1 6, same locality and date to the holotype; | 4, Nanshanchi, 23.11.1977, T. Naito; 1 4, Meifeng, Nantou Hsien, 8- 11.v.1984, K. C. Chou and C. C. Pan; 1 4; ‘*Hokuko, Kaminoshimaonsen,” Miaoli Hsien, 11.iv.1967, T. Shirozu. Distribution. — Taiwan. Remarks. —This new species is similar to S. soriculatipes Cresson from North Amer- 742 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Fig. 1, Strongylogaster fulva, penis valve. Figs. 2, 4, 5, Strongylogaster nantouensis. 2, Penis valve. 4, Sawsheath in dorsal view (left) and in lateral view (right). 5, Saw. Fig. 3, Strongylogaster formosana, penis valve. Figs. 6, 7, Pseudohemitaxonus taiwanus. 6, Sawsheath in dorsal view (left) and in lateral view (right). 7, Saw. Scale: 0.2 mm in Figs. 1-4, 6; 0.1 mm in Figs. 5, 7. VOLUME 92, NUMBER 4 ica, but the latter species differs from S. nantouensis by the yellowish clypeus, yel- lowish hind femur, dull frontal carina, and broad sawsheath in lateral view. Strongylogaster formosana (Rohwer) (Fig. 3) Thrinax formosana Rohwer, 1916, p. 100; Takeuchi, 1941, p. 235. Male.—Length, 6.5 mm. Similar to fe- male except for entirely black abdomen and anal cell of hind wing with petiole a little longer than breadth of the cell. Penis valve as in Fig. 3. Specimens examined.—1 4, Shitonshan, Miaoli Hsien, 15.11.1980, T. Tanabe; | 4, Meifeng, Nantou Hsien, 28.v.1975, S. Ima- saka; 1 6, Tsuifeng (2300 m), Nantou Hsien, 23.11.1979, A. Shinohara; 1 ¢, Alishan, Chiayi Hsien, 9.v.1922, S. Mori. Distribution. — Taiwan. Remarks.—Only two females had been recorded from Taiwan. The male was pre- viously unknown. Strongylogaster abdominalis (Takeuchi) Thrinax abdominalis Takeuchi, 1928, p. 42; Takeuchi, 1941, p. 235. Specimens examined.—1 ? 2 6, Pianan- anbu-Shikikun, Taipei Hsien, 20.vii.1932, T. Esaki; 1 6, Tapingshan, Taipei Hsien, 22.vii.1932, T. Esaki; 1 6, Sungkan, Nantou Hsien, 16.11.1977, A. Shinohara; | 2, Tsui- feng, Nantou Shien, 17.11.1977, A. Shino- hara; 1 2 1 4, Alishan (2300 m), Chiayi Hsien, 9.1v.1965, T. Saigusa; 3 4, Alishan (2400 m), 5-9.viii.1981, L. Y. Chou and S. C. Lin; 1 4, Alishan, 17—20.viii.1982, K. C. Chou and C. C. Pan. Distribution. — Taiwan. Genus Hemitaxonus Ashmead Trearea Malaise, 1947, p. 35. New syn- onym. Type species: T7rearea compressi- cornis Malaise. Monotypic. Canonarea Malaise, 1947, p. 38. New syn- onym. Type species: Canonarea alboora- lis Malaise. Original designation. 743 Malaise (1947) described Trearea and Canonarea as new genera of the Selandri- inae and commented that these two genera together with another related genus, Cano- nias Konow, compose a distinct and iso- lated group in the subfamily. No comments were given on the relationships to other groups. Canonarea, however, is quite iden- tical to Hemitaxonus. Characters separating Trearea from Canonarea are the absence of the Ist cubital crossvein in the fore wing and the compressed antenna, but these are considered to be variations within Hemi- taxonus. The first two genera, therefore, should be included in Hemitaxonus as new synonyms. This genus was represented by 16 species. Four species were recorded from North America (Smith 1969) and 12 species from East Asia, of which only one spreads to Eu- rope (Naito 1971). Three species described by Malaise (1947) as new species of Cano- narea or Trearea are newly included in this genus. Other than the species below, Hemi- taxonus compressicornis (Malaise), de- scribed in 7rearea, is a new combination. The following three species occur in Tai- wan, the last two representing the first rec- ords of these species there. Hemitaxonus formosanus Takeuchi Hemitaxonus formosanus Takeuchi, 1928, p. 43; Takeuchi, 1941, p. 247. Specimens examined.—1 2°, Tapingshan, Taipei Hsien, 24.vii.1932, T. Esaki; 1 9, Pahsienshan, Taichung Hsien, 30.viui.1929, K. Takeuchi; 1 @ 5 4, Sungkang, Nantou Hsien, 18.vii.1972, T. Naito; 1 4, Tsuifeng, Nantou Hsien, 18.11.1977, A. Shinohara; | 4, Tsuifeng (2300 m), 23.111.1979, A. Shi- nohara. Distribution. — Taiwan. Hemitaxonus albooralis (Malaise), New CoMBINATION Canonarea albooralis Malaise, 1947, p. 38. Specimens examined.—1 2, Meifeng- 744 Sungkang, Nantou Hsien, 4.v.1978, A. Shi- nohara; 2 2, Meifeng (2150 m), 8-11.v.1984, K. C. Chou and C. C. Pan. Distribution.— Taiwan, Burma. Remarks.—Three females from Taiwan are identical with the paratype from Burma in structure and coloration. This is the first record of this species from Taiwan and also outside the type locality, Burma. Hemitaxonus nigrooralis (Malaise), NEw CoMBINATION Canonarea nigrooralis Malaise, 1947, p. 39. Specimens examined.—1 2, Lushan (1000 m), Nantou Hsien, 27—31.v.1980, K. S. Lin and L. Y. Chou; | 2, Wusho (1159 m), Nan- tou Hsien, 6-1 1.v.1981, K. S. Lin and S. C. Lin; 2 2, Tunpu (1200 m), Nantou Hsien, 23-27.vi1.1984, K. C. Chou and C. H. Yang; 1 2, Lienhwachi, Nantou Hsien, 26.111.1984, GCXEe2Lo: Distribution.—Taiwan, Burma, Hima- laya. Remarks.—This is the first record of this species from Taiwan. I also examined a fe- male specimen from Hymalaya labeled “Panjab, Himalaya, Khajjiar, 23.vi.1965, Tikav leg.”’ These are the first occurrences outside the type locality, Burma. Genus Pseudohemitaxonus Conde Three species have been described, one from Europe and two from Japan (Naito 1969). The following species represents the first record of this genus from Taiwan. Pseudohemitaxonus taiwanus New SPECIES (Figs. 6, 7) Female.—Length, 5.3 mm. Head and thorax black; clypeus, labrum, labial and maxillary palpi, scape, pedicel (with dark band on anterior half), pronotum, tegula and legs (except for hind tarsus infuscate) yel- lowish. Abdomen dark brown; 2nd to 4th terga pale brown and with yellowish lon- gitudinal line in middle; 2nd to 6th terga PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with apical margins narrowly and lateral margins broadly yellowish. Head shining, feebly shagreened, with small punctures and sparse pubescence. La- brum nearly flat, rounded in front. Clypeus gently convex, shallowly emarginate in front to depth of about 4 its medial length. Me- dian fovea indistinct and lateral fovea punc- tiform. Frontal area represented by feebly raised platform, not clearly defined. Inter- ocellar and postocellar furrows absent. Ver- tical furrow represented by shallow line, somewhat convergent toward apex. Post- ocellar area nearly flat; breadth: length = 15:7. Malar space about %x diameter of front ocellus. Postgenal carina developed on lateral side. Antenna filiform, about 2.1 x breadth of head; relative lengths of seg- ments about 6:5:24:24:20:15:15:12:14. Thorax shining and smooth; mesonotal middle lobe and lateral lobes with small and close punctures; pronotum, scutellum and mesepisternum with sparse and fine punc- tures. Posttergite large, about 1.5 times as wide as scutellum. Prepectus defined by dis- tinct suture. Anal cell of fore wing with sub- erect crossvein; Ist cubital crossvein absent; vein Rs+M strongly curved toward stigma. Anal cell of hind wing with petiole about 1.5 greatest breadth of cell. Inner front tibial spur slender, slightly bifid at apex. Tarsal claw with very small inner tooth. Ab- domen feebly reticulate throughout. Pro- podeum deeply and broadly excised. Saw- sheath slender, not bifid at apex (Fig. 6). Saw as in Fig. 7. Male. —unknown. Holotype.—Female, Meifeng, 2150 m, Nantou Hsien, 19-21.iv.1983, K. C. Chou and S. P. Huang (Deposited in the Taiwan Agricultural Research Institute). Distribution. — Taiwan. Remarks.—This new species resembles P. dryopteridis Naito from Japan but is eas- ily distinguished from the latter by the fron- tal area which is not surrounded by a carina, the clearly defined prepectus, and the very VOLUME 92, NUMBER 4 small but distinct inner tooth on the tarsal claw. ACKNOWLEDGMENTS I express my sincere thanks to Dr. D. R. Smith, Systematic Entomology Laboratory, U.S.D.A., Washington, D.C., for critical reading of the manuscript. I also express appreciation to the following people who have allowed study of valuable specimens in their private or institute collections: Dr. L. Y. Chou, Taiwan Agricultural Research Institute; Dr. A. Shinohara, Department of Zoology, National Science Museum, Tokyo; Mr. H. Kumamoto, Hirakata City, Osaka; and Prof. T. Saigusa, Kyushu University, Fukuoka. LITERATURE CITED Benson, R. B. 1968. Hymenoptera from Turkey. Symphyta. Brit. Mus. (Nat. Hist.) Ent. Bull. 22: 109-207. Enslin, E. 1913-1914. Die Tenthredinoidea Mitte- leuropas. Beit. Deut. Ent. Zeit., Beihefte, pp. 99- 309. Malaise, R. 1947. The Tenthredinoidea of south- eastern Asia. The Emphytus-Athlophorus group. Ark. Zool. 39A: 1-39. 745 Naito, T. 1969. The genus Pseudohemitaxonus of Japan, with descriptions of two new species. Kon- tyu, Tokyo 37: 403408. 1971. A revision of the genus Hemitaxonus in the old world, I (Hymenoptera, Tenthredini- dae). Konty, Tokyo 39: 19-28. 1975. Phylogeny and distribution of five re- lated genera, Eriocampidea, Hemitaxonus, Pseu- dohemitaxonus, Nipponorhynchus and Adelesta, referred to the Selandriinae (Hymenoptera, Ten- thredinidae). Kontya, Tokyo 43: 330-342. . 1980. Studies on the Japanese Sawflies of the Genus Strongylogaster Dahlbom (Hymenoptera, Tenthredinidae). Kontyi, Tokyo 48: 390-401. Naito, T. and F. S. Huang. 1988. The sawfly genus Strongylogaster in Tibet and Sichuan, China (Hy- menoptera, Tenthredinidae). Sci. Rept. Fac. Agr. Kobe Univ. 18: 41-48. Rohwer, S. A. 1916. H. Sauter’s Formosa-Ausbeute. Chalastogastra (Hymenoptera). Suppl. Ent. 5: 81- 113: Smith, D.R. 1969. Nearctic sawflies II. Selandriinae: Adults. U.S. Dep. Agr. Tech. Bull., 1938, 48 pp. Takeuchi, K. 1928. New sawflies from Formosa—II. Formosa Nat. Hist. Soc. Trans. 18: 38-45. 1941. A systematic study of the suborder Symphyta (Hymenoptera) of the Japanese empire (IV). Tenthredo 3: 230-274. Zhelochovtsev, A. N. 1951. [Survey of the Palaearc- tic sawflies of the subfamily Selandriinae (Hym., Tenthr.)]. Moscow Gosud. Univ. Zool. Muz. Sborn. Trudy 7: 123-152. [In Russian.] PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 746-756 A REVISION OF THE SHORE-FLY GENUS DIPHUIA CRESSON (DIPTERA: EPHYDRIDAE) Wayne N. MATHIS Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560. Abstract. — Diphuia Cresson, a New World genus of shore flies, is revised and found to be close phylogenetically (the sister group) to the lineage giving rise to Allotrichoma Becker, including Pseudohecamede Hendel. Although four species (D. anomala Cresson, D. nasalis Wirth, D. nitida Sturtevant and Wheeler, and D. zatwarnickii, new species (Jamaica) are recognized, the second and third are very similar and may prove to be conspecific when adequate material of D. nitida is available. Characters of the male postabdomen and terminalia are described and illustrated. Key Words: While conducting field work on several cays within the Stann Creek District of Be- lize, I found a tiny, black-colored shore fly that is associated with mangrove peat. The peat, which is exposed at low tide, is shaded during most of the day beneath the dense canopy of well-developed fringe red man- grove (Rhizophora mangle L.). The speci- mens did not occur on peat that is the sub- strate for scrub or dwarf red mangrove and where little or no shade is provided. Deter- mining the identity of this species, which is less than two mm in length, has led to this revision of Diphuia Cresson, the genus to which the species has been assigned. In ad- dition to determining the identity of the specimens from Belize, the other specific purposes of this revision are to provide the first illustrations of the male terminalia and to determine the phylogenetic position of Diphuia, which Cresson (1944) suggested was related to Allotrichoma Becker in the tribe Atissini. Shore flies of the genus Diphuia are anom- alies among atissines in being mostly black, lacking dense vestiture of gray to brown mi- Ephydridae, shore flies, Diphuia, revision, systematics crotomentum, and having a distinctively marked face that is black with silvery white, microtomentose lines. The facial markings of microtomentum are similar to specimens of Discocerina nitida Cresson (tribe Disco- cerinini) and several genera of the tribe Gymnomyzini. The superficial resemblance of this genus to discocerines or gymnomy- zines prompted Cresson to formulate the generic name Diphuia, which is a Latin transliteration of Greek words meaning double nature. Although similar to genera in other tribes, the genus is related most closely to Allotrichoma, as Cresson con- cluded in the original description, and the similarities noted are apparently the result of convergences. When Cresson (1944) originally proposed Diphuia he included only the type species, D. anomala Cresson, which was described in the same paper. Two years later, Cresson again treated Diphuia and its type species in his synopsis of Neotropical Psilopinae (= Gymnomyzinae). The genus then remained unstudied for nearly a decade, which is not surprising in view of their diminutive size, VOLUME 92, NUMBER 4 restricted distribution, and rarity in collec- tions. Sturtevant and Wheeler (1954) wrote the concluding part for Cresson’s synopses of Nearctic shore flies, following the latter’s death, and described D. nitida from a single specimen that was collected near New York City. Two years afterwards, Wirth (1956) reviewed the shore flies of the Bahamas and described a third species, D. nasalis. Aside from catalogs of the Nearctic and Neotrop- ical Regions (Wirth 1965, 1968, respective- ly), no further work has been published on Diphuia. Nothing is known of the immature stages, no key 1s available, and the structures of the male terminalia have not been in- vestigated, described, or illustrated. Methods.—The terminology and meth- ods used in this study were explained pre- viously (Mathis 1986a, b). Because of the small size of specimens, study and illustra- tion of the male terminalia required the use of a compound microscope. To better as- sure effective communication about struc- tures of the male terminalia, I have adopted the terminology of other workers in Ephyd- ridae (see references in Mathis 1986b). Us- age of these terms, however, should not be taken as an endorsement of them from a theoretical or morphological view over al- ternatives that have been proposed (Grif- fiths 1972, McAlpine 1981). Rather, I am deferring to tradition until the morpholog- ical issues are better resolved. Two venational ratios are used common- ly in the descriptions and are defined here for the convenience of the user (ratios are averages of three specimens). 1. Costal vein ratio: the straight line dis- tance between the apices of R,,, and R,,;/ distance between the apices of R, and Rj, ;. 2. M vein ratio: the straight line distance along M between crossveins (dm-cu and r-m)/distance apicad of crossvein dm-cu. Most of the specimens used in this study, a total of 225, are housed in the National Museum of Natural History (USNM), Smithsonian Institution. Prior to my tenure at the Smithsonian, W. W. Wirth had ac- 747 cumulated several specimens of what ap- peared to be the same tiny fly. His collec- tions from Jamaica and Dominica are especially noteworthy in that regard. I also examined collections of the Academy of Natural Sciences of Philadelphia (ANSP), the American Museum of Natural History (AMNH), and the University of Texas (UTA). Diphuia Cresson Diphuia Cresson, 1944: 4. Type species: Di- phuia anomala Cresson, 1944, by original designation; 1946: 138, 140 [note, key].— Sturtevant and Wheeler, 1954: 248 [notes].—Wirth, 1956: 4 [discussion of species]; 1968: 5 [Neotropical catalog]. Diagnosis.— Mostly black, subshiny to shiny, microtomentum usually sparse; small shore flies, length 1.35 to 1.80 mm. Head: Wider than high; face width-to- head width ratio 0.28; frons black, mostly unicolorous, lacking distinctively colored ocellar triangle; frons wider than long, fron- tal length-to-width ratio 0.58; frontal ves- titure variable; ocellar seta well developed, inserted slightly in front of alignment of an- terior ocellus and at about the same distance apart as between posterior ocelli; pseudo- postocellar setae usually well developed, length subequal to ocellar setae, proclinate, slightly divergent; | reclinate and 1| procli- nate fronto-orbital seta present, reclinate seta inserted slightly anteromediad of proclinate seta; both inner and outer vertical setae present; ocelli arranged to form isosceles tri- angle, with distance between posterior ocelli larger than between anterior ocellus and ei- ther posterior ocellus. Antenna exerted; pedicel with well-developed, proclinate, dorsal seta; aristal length subequal to an- tennal length and bearing 4—5 dorsal rays, with basal 3 rays longer than apical 1-2, the latter subequal. Eye apparently bare of mi- crosetulae (using a stereomicroscope). Face black in both sexes and with silvery white, microtomentose antennal grooves and with 748 2 lines, sometimes irregular, paralleling par- afacials, these and similarly invested and colored ventral margin (microtomentum sometimes interrupted at middle) form a facial triangle that has a small microtomen- tose area below facial prominence; face not carinate between antennal bases but slight- ly, conically protrudent at middle (best seen in lateral view); ventral facial margin shal- lowly emarginate; face bearing 2 facial setae, the dorsal seta very slightly larger, both in- serted near parafacials; parafacials densely microtomentose, silvery white; clypeus very sparsely microtomentose, black; palpus blackish brown to black; mouthparts not geniculate, labella shorter than medipro- boscis. Thorax: Generally black, vestiture of microtomentum variable with species, al- though generally sparse; pleural areas lack- ing stripes of distinctly colored microto- mentum. Chaetotaxy with mesonotal setae poorly developed except for those at pos- terior margin; mesonotal setulae numerous and not arranged in well-defined setal tracks; prescutellar acrostichal setae much larger than other acrostichal setulae and more widely set apart; only | dorsocentral seta, inserted posteriad; intra-alar setulae irreg- ularly seriated; presutural seta well devel- oped, length subequal to notopleural setae; 2 scutellar setae and scutellar disc with sparse, scattered setulae; postpronotal seta 1; postalar seta 1; notopleural setae 2, in- sertion of posterior seta elevated dorsally above anterior one; anepisternal setae 2, in- serted along posterior margin; katepisternal seta well developed, conspicuous. Wing: membrane mostly hyaline to very slightly milky white; veins behind costa pale, usu- ally yellowish to yellowish brown; vein R,,, extended well beyond level of crossvein dm- cu, 2nd costal section at least 1' times lon- ger than 3rd section; alular marginal setulae short, less than '2 alular height. Legs: fem- ora black; tibiae dark basally, concolorous with femora, apices yellowish. Abdomen: Fifth segment of male well PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sclerotized, elongate, not normally visible from a dorsal view, usually retracted within 4th segment; 5th tergum and sternum of male united anteriorly to form a complete annulus. Male terminalia as follows: cercus rod shaped, bearing 2-3 conspicuously lon- ger setae at ventral margin; surstylus well developed, well sclerotized, and conspicu- ous, length as long as cercus. Distribution.—New World. Temperate to tropical zones, in North America along the east coast (New York south to Florida) and the Caribbean to Colombia and Eucador in South America. Phylogenetic relationships.—Diphuia is related to a group of taxa (Allotrichoma Becker, Eremotrichoma Soika, Pseudohe- camede Hendel, and Hecamede Haliday) within the tribe Atissini that is character- ized by having very sparse or lacking mi- crosetulae on the compound eyes; a coni- cally prominent face (degree of development varies) that is emarginate ventrally and with the clypeus exposed in the emargination; oral opening and clypeus narrow; area sur- rounding crossvein dm-cu not infuscate; the apex of the wing broadly rounded, not pointed at the apex of vein R,,.; fifth ter- gum of male retracted within the enlarged fourth, usually not visible; and cerci with elongate setulae at ventral margin. Diphuia appears to be the sister group to the lineage giving rise to taxa closely related to Allotri- choma sensu lato, including Pseudohe- camede. This relationship is evidenced by the characters noted previously, especially the retracted fifth tergum of the male, which is moderately elongate, almost tubular. Al- though related and similar to this group, Diphuia may be distinguished as follows (characters indicated by an asterisk are aut- apomorphies that corroborate the monophy- ly of Diphuia): *coloration very dark, usu- ally black; *microtomentum of head and thorax generally sparse, giving a subshiny to faintly dull appearance; facial coloration of male and females similar, lacking sexual dimorphism; face, although slightly pro- VOLUME 92, NUMBER 4 trudent medially (best seen in profile), not acutely pointed in lateral view; *face with silvery microtomentose markings, antennal grooves, 2 vertical lines, ventral margin, an area below the facial prominence, and par- afacials; presutural and prescutellar setae well developed; *pleural region lacking a stripe or stripes; 5th segment of male well sclerotized and its tergum moderately en- longate; *Sth tergum and sternum of male united anteriorly to form a complete an- nulus; and male genitalia with distinct, well- sclerotized, elongate surstyli. The place- ment of Diphuia as the sister group to the lineage of Al/otrichoma sensu lato follows Cresson’s original assessment. The evi- dence supporting this relationship is not strong, however, and Diphuia could be re- lated to Hecamede. Two species groups are evident within D- phuia, each comprising two species: D. anomala Cresson and D. zatwarnickii, a new species that is described below; and D. ni- tidaand D. nasalis. For species in the former group, I have found that characters of the male terminalia only are adequate to dis- tinguish between the species. In the second group, the degree of microtomentum on the frons may be a distinguishing character, al- though that character is questionable (see ““Remarks” under D. nitida). Discussion.— Two shore-fly species (Dis- cocerina quadripectinata (Becker) and A/- lotrichoma argentipraetextum Lamb) that are now or perhaps should be assigned to Allotrichoma and related genera and that are dark colored are not closely related to Di- phuia (Zatwarnicki, in litt.). Nothing is known about the immature stages or natural history of any of the species included in Diphuia. Key TO SPECIES OF DIPHUIA 1. Mesofrons bare of microtomentum, shiny (New VOLK) fen Crean D. nitida Sturtevant and Wheeler — Mesofrons mostly densely microtomentose, at most with small shiny area immediately before anterior ocellus (southeastern USA and Neo- ELODICS) eee eres eee chip heceners Sere at 2 749 2. Mesonotum thinly invested with microtomen- tum, subshiny; anepisternum with anteroven- tral 3-2 bare, shiny black, otherwise with thin investment of whitish gray microtomentum dex Pra eae area tthe rates ctions D. nasalis Wirth — Mesonotum moderately densely microtomen- tose, golden brown; anepisternum almost en- tirely invested with whitish gray microtomen- tum ... Pe es rere ochre 3 3. Surstyli long and narrow, length subequal to that of cercus (Fig. 3); gonite with pointed pos- teroventrally; aedeagus only moderately curved apically see tate ... D. anomala Cresson — Sturstyli moderately short and robust (Fig. 18), length shorter than cercus; gonite with poster- oventral portion broadly bifurcate; aedeagus more curved apically, point oriented anteriorly C. zatwarnickii, new species Diphuia anomala Cresson Figs. 1-7 Diphuia anomala Cresson, 1944: 4; 1946: 138 [review].— Wirth, 1968: 5 [Neotrop- ical catalog]. Description.—Small shore flies, length 1.60 to 1.80 mm. Head: Frons moderately invested with brownish microtomentum, microtomen- tum sparse or lacking on 2 small areas la- terad of posterior ocelli and 2 spots along the anterior margin. Thorax: Mesonotum densely invested with brownish to golden brown microto- mentum, especially medially, along poste- rior portion of scutum and scutellum; an- episternum with fine investment of whitish microtomentum. Wing with costal vein ra- tio 0.50; M vein ratio 0.41. Abdomen: Sth tergum (Figs. 1, 2) almost as high as long, anterior margin in dorsal view with deep, broadly V-shaped emargi- nation (Fig. 2), posterior margin with sparse setae; 5th sternum clearly divided into 2 broad sternites that are connected only an- teroventrally (Fig. 1). Male terminalia (Figs. 3-7) as follows: epandrium bulbous, shiny, in lateral view almost as wide as high (Fig. 3); surstylus long, narrow, parallel sided, width and length subequal to that of cercus, apex angulate, pointed anteriorly, and bear- 750 ing a few setulae (Fig. 3); gonite broad bas- ally, with posteriorly extended process sheathing aedeagus, posterior apex of gonite curved anteroventrally (Figs. 4, 5); aedeagal apodeme triangular in lateral view (Figs. 5, 7), narrowly produced dorsally; aedeagus in lateral view broad, thumblike, produced posteroventrad to a ventral point, in dorsal view becoming wider apically, apex broadly rounded (Figs. 5, 7); hypandrium in ventral view longer than wide, anterior margin with a small, anterior process (Figs. 5, 6). Type material.—The holotype male is labeled ‘‘Monte Lirio[,] PANAMAJ,] RCShannon{[,] IV.6.23 [6 Apr 1923]/4/ TYPE DIPHUIA ANOMALA ¢ E.T. Cres- son, Jr. [red, species name and ¢ handwrit- ten]; TypeNo 70450 USNM [red, number handwritten].’"> The holotype is point mounted, is in good condition (the right first flagellomere is missing), and is deposited in the USNM. The allotype and several para- types are also deposited in the USNM. Other specimens examined.—COLOM- BIA. Rio Raposo (light trap), Jan 1964, V. H. Lee (1 6; USNM). ECUADOR. Los Rios Province. Guare, Aug 1955, J. R. Levi-Cas- tillo (6 6, USNM). Manabi Province. Ca- marones, 9 Sep 1955, J. R. Levi-Castillo (1 6; USNM); Estero Balsa, 9 Sep 1955, J. R. Levi-Castillo (13 6, 3 2; USNM); La Palma, Aug 1955, J. R. Levi-Castillo (1 6; USNM). EL SALVADOR. Laguna de Zapotitan, Dec 1953, W. B. Heed (1 2; UTA). PANAMA. Canal Zone: Balboa, Feb 1958, M. R. Wheeler (1 6; UTA), Monte Lirio, 6 Apr 1923, R. C. Shannon (29 4, 19 2; ANSP, USNM); Pedro Miguel, 10 Apr 1923, R. C. Shannon (4 4, 1 2; USNM). Panama: Da- rién: Sabanas, 20 Apr 1923, R. C. Shannon (4 8, 22; USNM). Panama City, 5 Apr 1923, R. C. Shannon (2 ?; ANSP, USNM). Distribution.—Colombia, Ecuador, El Salvador, and Panama. Remarks. —This is the type species of Di- phuia. It is very similar externally to D. zat- warnickii and can be distinguished only by reference to structures of the male termin- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON alia (see ““Remarks” under D. zatwarnickii and couplet 3 of the key). From D. nasalis and D. nitida it may be distinguished by the following characters: frons and mesonotum invested moderately densely with brownish to golden brown microtomentum; anepi- sternum invested with fine, grayish to whit- ish microtomentum, anteroventral portion not bare, shiny; second costal section long, costal vein ratio 0.50; and several characters of the male terminalia (see description and figures). Diphuia nasalis Wirth Figs. 8-15 Diphuia nasalis Wirth, 1956: 3; 1968: 5 [Neotropical catalog]. Description.—Small shore flies, length 1235; to le7S)mm: Head: Frons sparsely invested with fine brownish microtomentum, microtomen- tum becoming sparser or lacking on 2 small areas laterad of posterior ocelli, 2 spots along the anterior margin, and sometimes a small spot in front of the anterior ocellus. Thorax: Mesonotum sparsely invested with fine brownish to golden brown micro- tomentum, mostly subshiny to shiny; anepi- sternum with anteroventral 3 to '2 bare of microtomentum, shiny, posterodorsal por- tion invested with fine, grayish microto- mentum. Wing with costal vein ratio 0.58; M vein ratio 0.40. Abdomen: 5th tergum with anterior mar- gin essentially straight, at most very shal- lowly arched anteriorly (Fig. 9); 5th sternum undivided, as a narrow band connected dor- sally with anteroventral portion of Sth ter- gum (Fig. 8). Male terminalia (Figs. 10-15) as follows: epandrium narrow in lateral view, much higher than wide (Fig. 10); surstylus as long as cercus but almost twice its width, broadly rounded apically (Fig. 10); gonite in lateral view parallelogram-shaped, pos- terior angles produced into pointed pro- cesses, posteroventral process sinuate (Figs. 11, 12); aedeagal apodeme rounded anter- VOLUME 92, NUMBER 4 Figs. 1-7. oventrally (Figs. 12, 13); aedeagus acutely pointed apically, in dorsal or ventral view bifurcate apically (Figs. 12-14); hypan- drium in ventral view wider than long, an- terior margin shallowly arched anteriorly (Figs. 12, 14, 15). Type material.—The holotype female is labeled “Long Island[,] Deadman’s Cay[,] March 11, 1953/Van Voast—A.M.N.H. Bahama Isls. Exped Coll. E. B. Hayden/é/ SHOLOTY PE Diphuia nasalis W. W. Wirth [red, gender and species name handwrit- ten].”” The holotype is point mounted, is in 71 Diphuia anomala. 1, 5th tergum and sternum, lateral view. 2, 5th tergum, dorsal view. 3, Male terminalia (epandrium, cercus, surstyli), lateral view. 4, Gonite, lateral view. 5, Internal male terminalia (gonite, hypandrium, aedeagal apodeme, aedeagus), lateral view. 6, Hypandrium and aedeagal apodeme, ventral view. 7, Aedeagus and aedeagal apodeme, lateral view. Scale bar = 0.1 mm. good condition (tip of right wing folded back on itself), and is deposited in the AMNH. Although the holotype was listed as a male (Wirth 1956: 4) and the specimen is so marked, it is a female. Other specimens examined.— BAHA- MAS. Crooked Island, Landrail Point, 5 Mar 1953, E. B. Hayden, L. Giovannoli (1 2; AMNH); Exuma Cays, Staniard Bay, 13 Jan 1953, E. B. Hayden (1 2; AMNH); Long Island, Deadman’s Cay, 11 Mar 1953, E. B. Hayden (2 6, AMNH, USNM). BELIZE. Stann Creek District: Bread and Butter Cay, 752 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-15. terminalia (epandrium, cercus, surstyli), lateral view. 11, Gonite, lateral view. 12, Internal male terminalia (gonite, hypandrium, aedeagal apodeme, aedeagus), lateral view. 13, Aedeagus and aedeagal apodeme, lateral view. 14, Hypandrium, aedeagal apodeme, and aedeagus, ventral view. 15, Hypandrium, lateral view. Scale bar = 0.1 mm. Diphuia nasalis. 8, 5th tergum and sternum, lateral view. 9, 5th tergum, dorsal view. 10, Male 25 Mar 1988, W. N. Mathis (5 46, 1 2; Cays(West Bay), 22 Mar 1988, W. N. Math- USNM); Glover’s Reef (Long Cay, Middle is (1 6, 1 2; USNM); Wee Wee Cay, 24-25 Cay, Northeast Cay, Southwest Cay), 26- Mar 1988, 21 Jul 1989, W. N. Mathis (5 4; 28 Jul 1989, W. N. Mathis (29 6, 12 2; USNM). Six Belize, 1959, N. L. H. Krauss USNM); Man of War Cay, 8-15 Nov 1987, (1 6; USNM). BERMUDA. Hamilton Par- W.N. & D. Mathis (7 6, 4 2; USNM); Twin _ ish. Shelly Bay, 20 Nov 1987, D. J. Hilburn, VOLUME 92, NUMBER 4 i538 Figs. 16-22. Diphuia zatwarnickii. 16, Sth tergum and sternum, lateral view. 17, 5th tergum, dorsal view. 18, Male terminalia (epandrium, cercus, surstyli), lateral view. 19, Internal male terminalia (gonite, hypandrium, aedeagal apodeme, aedeagus), lateral view. 20, Gonite, lateral view. 21, Aedeagus and aedeagal apodeme, lateral view. 22, Hypandrium, ventral view. Scale bar = 0.1 mm. N. E. Woodley (2 2; USNM). JAMAICA. Runaway Bay (bay shore), 16-28 Feb 1969, Falmouth (bay shore), | Mar 1969, W. W. W. W. Wirth (3 4; USNM). UNITED Wirth (1 6; USNM); Milk River Bath(man- STATES. Florida. Monroe Co., Bahia Hon- groves), 11 Mar 1970, T. Farr, W.W. Wirth da Key (seashore), 11 Apr 1970, W. W. (5.4, 1 9; USNM); Negril Beach (rocky shore), Wirth (4 4, 3 ¢; USNM); Big Pine Key, 11 12 Mar 1970, W. W. Wirth (1 6, USNM); Apr-30 Dec 1954, 1970, H. V. Weems (1 754 6, 1 2; USNM); Long Key, 23 Jun 1953, M. R. Wheeler (1 6; UTA); Saddlebunch Keys, 29 Dec 1953, H. V. Weems (1 4, 1 2; USNM). North Carolina. Onalow Co., Ashe Island, 11 Aug 1975, J. C. Dukes (1 2; USNM). Distribution. — Bahamas, Belize, Bermu- da, Jamaica, and USA (Florida, North Ca- rolina). Natural history.—The vast majority of specimens from the Belizean cays were col- lected by sweeping just above mangrove peat that is well shaded most of the day. A few specimens, apparently feeding, were col- lected on flowers. The association with mangrove peat must be opportunistic, as the species occurs in areas where mangrove does not now exist. Remarks.—This species is distinguished from congeners, especially D. anomala and D. zatwarnickii, by the sparsely microto- mentose mesofrons (although not shiny as in D. nitida); the subshiny mesonotum that is very thinly invested with fine microto- mentum; the shiny anteroventral one-third to one-half of the anepisternum; and several characters of the male terminalia (see de- scription and figures). Diphuia nitida Sturtevant and Wheeler Diphuia nitida Sturtevant and Wheeler, 1954: 248.—Wirth, 1965: 737 [Nearctic catalog]. Description.—Small shore flies, length 1.60 mm. Head: Frons, except for fronto-orbits, bare, shiny, especially mesofrons and fron- tal triangle; fronto-orbits invested with brown microtomentum; frontal triangle chestnut brown, mesofrons otherwise black. Thorax: Mesonotum sparsely microto- mentose, subshiny to shiny, black; anepi- sternum mostly shiny, posterodorsal angle with some grayish to whitish microtomen- tum. Type material.— The holotype ° is labeled ““Dougl[t]s[t]on[,] Lfong]. I[sland]., N[ew]. Y{ork].[,] Au[gust]. 17, [19]52/HOLO- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON TYPE Diphuia nitida Stvt & Whlr [pink]/ TYPE 6695 [dark pink; number handwrit- ten].”’ The holotype is point mounted, is in poor condition (the antennae and several setae are missing and the left side of the body and wings are covered partially with glue), and is deposited in the ANSP (6695). Sturtevant and Wheeler stated that this specimen is a male, but is clearly a female. Distribution.— USA. New York: Long Is- land, Douglaston (just within the city limits of New York City). Remarks.—This species is very similar to, and may be conspecific with, D. nasalis. Resolution of this question will depend on collection and study of additional material, especially males, from the type locality or a locality nearby. At present, the only known specimen of this “species” is the female ho- lotype, which, as noted earlier, is in poor condition, making it impossible to ascertain its identity with certainty. I recognize the holotype as being different and possibly rep- resenting a separate species mostly because the few external features that are discern- able, especially those of the head, are not within the variation among specimens of D. nasalis that I have studied. The shinier frons of the holotype appears to be unique. Fur- thermore, the distance between New York, which is the type locality of this species, and the nearest locality where D. nasalis is known to occur (North Carolina) is several hun- dreds of miles. As these populations are somewhat disjunct and are apparently dif- ferent, I am provisionally recognizing them as representing separate species. If the pop- ulations prove to be conspecific, D. nitida is the senior synonym. Diphuia zatwarnickiit, NEw SPECIES Figs. 16-22 Description.—Small shore flies, length 1.40 to 1.90 mm. Head: Frons moderately invested with brownish microtomentum, microtomen- tum sparse or lacking on 2 small areas la- VOLUME 92, NUMBER 4 terad of posterior ocelli and 2 spots along the anterior margin. Thorax: Mesonotum densely invested with brownish to golden brown microto- mentum, especially medially, along poste- rior portion of scutum and scutellum; an- episternum with fine investment of whitish microtomentum. Wing with costal vein ra- tio 0.52; M vein ratio 0.42. Abdomen: 5th tergum (Figs. 16, 17) about as high as long, anterior margin in dorsal view with deep, broadly V-shaped emargi- nation (Fig. 17), posterior margin with sparse setae; 5th sternum clearly divided into 2 broad sternites that are connected only an- teroventrally (Fig. 16). Male terminalia (Figs. 18-22) as follows: epandnum bul- bous, shiny, in lateral view almost as wide as high (Fig. 18); cercus cylindrical; sursty- lus moderately long and narrow, parallel sided, width subequal to that of cercus but length shorter, apex angulate, pointed an- teriorly, and bearing a few setulae (Fig. 18); gonite broad basally, with posteriorly ex- tended process sheathing aedeagus, poste- rior apex of gonite bifurcate (Figs. 19, 20); aedeagal apodeme triangular in lateral view (Figs. 19, 21), narrowly produced dorsally; aedeagus in lateral view broad, thumblike, curved posteroventrad to an anteroventral point, in dorsal view becoming wider api- cally, apex broadly rounded (Figs. 19, 21); hypandrium in ventral view longer than wide, anterior margin with a small, anterior process (Fig. 22). Type material.— The holotype male is la- beled “JAMAICA 5mi.E.Negril 13March 1970 W. W. Wirth fresh marsh.” The al- lotype female and three other paratypes (2 6, 1 2; USNM) bear the same label data as the holotype. Other paratypes are from: DOMINICA. Cabrit Swamp, 22-25 Mar 1965, W. W. Wirth (6 6, 2 2; USNM); Woodford Hill, 27 Feb 1965, W. W. Wirth (2 6; USNM). JAMAICA. Kingston, Fresh River, 24 Feb 1969, W. W. Wirth (8 4, 5 2; USNM); Milk River Bath, 11 Mar 1970, T. Farr, W. W. Wirth (7 4, 4 2; USNM); Rio 755 Bueno, 21 Feb 1969, W. W. Wirth (1 2; USNM); Savanna La Mar, 13 Mar 1970, W. W. Wirth (2 6; USNM). The holotype is double mounted (minute nadel in polyporus block), is in excellent condition, and is de- posited in the Smithsonian Institution (USNM). Distribution.— West Indies: Dominica and Jamaica. Etymology.—This species is named for Tadeusz Zatwarnicki, who first brought this species to my attention and who has con- tributed significantly to the study of shore flies. Remarks.—This species is distinguished from D. nasalis and D. nitida by the sparsely microtomentose mesofrons; the subshiny mesonotum that is very thinly invested with fine microtomentum; the shiny anteroven- tral one-third to one-half of the anepister- num; and several characters of the male ter- minalia. This species is distinguished from D. anomala by characters of the male ter- minalia: especially the shorter, more robust surstyli; the gonite that is broadly bifurcate posteroventrally; and the more apically curved aedeagus (see description and fig- ures). ACKNOWLEDGMENTS I am grateful for the assistance in the field from Candy Feller, Holly Williams, and Dianne Mathis. For the opportunity to ex- amine specimens housed in their collec- tions, I thank David Grimaldi and Julian Stark (AMNH) and Donald Azuma (ANSP). For critically reviewing a draft of this paper, I thank Allen Norrbom, Norman Woodley, Willis Wirth, and Tadeusz Zatwarnicki. The illustrations were skillfully inked by Elaine R. S. Hodges. Funding for this research project, espe- cially the field work in Belize, was provided by the Caribbean Coral Reef Ecosystems (CCRE), Smithsonian Institution. This is contribution number 282 of the CCRE pro- ject, which is partially supported by a grant from the Exxon Corporation. 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Cresson, E. T., Jr. 1944. Descriptions of new genera and species of the dipterous family Ephydridae. Paper XIV. Notulae Nat. Acad. Nat. Sci. Phila- delphia. 135: 1-9. 1946. A systematic annotated arrangement of the genera and species of the Neotropical Ephydridae (Diptera). 1. The subfamily Psilopi- nae. Trans. Amer. Entomol. Soc. 71: 129-163. Griffiths, G. C. D. 1972. The Phylogenetic Classifi- cation of Diptera Cyclorrhapha with Special Ref- erence to the Structure of the Male Postabdomen. Series Entomologica 8: 340 pp. W. Junk, Hague. Mathis, W. N. 1986a. A revision of the subgenus Eremotrichoma Soika of the shore fly genus A/- lotrichoma Becker (Diptera: Ephydridae). Israel J. Entomol. 19: 127-139. 1986b. Studies of Psilopinae (Diptera: Ephydridae), I: A revision of the shore fly genus Placopsidella Kertész. Smithson. Contrib. Zool. 430: 1-30. McAlpine, J. F. 1981. Morphology and terminolo- gy—Adults, pp. 9-63. In McAlpine, J. F., et al., eds., Manual of Nearctic Diptera. Ottawa. [Vol- ume | is Monograph 27 of Research Branch Agric. Canada.] Runyan, J. T. and D. L. Deonier. 1979. A compar- ative study of Pseudohecamede and Allotrichoma (Diptera: Ephydridae), pp. 123-127. Jn Deonier, D. L., ed., First Symposium on the Systematics and Ecology of Ephydridae (Diptera). Oxford, Ohio: North Amer. Bentho. Soc. Sturtevant, A. H.and M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera). Trans. Amer. Entomol. Soc. 79: 151-257. Wirth, W. W. 1956. The Ephydridae (Diptera) of the Bahama Islands. Amer. Mus. Novitates 1817: 1- 20. 1965. Family Ephydridae, pp. 734-759. In Stone, A., et al., eds., A Catalog of the Diptera of America North of Mexico. Agri. Hankbk. 276. 1968. 77. Family Ephydridae, pp. 1-43. Jn Papavero, N., ed., A Catalogue of the Diptera of the Americas South of the United States. Dept. Zool. Sec. Agri. Sao Paulo. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 757-759 NOTE Dispersal of the Southern Green Stink Bug, Nezara viridula (L.) (Heteroptera: Pentatomidae), by Hurricane Hugo Nezara viridula (L.) (Heteroptera: Pen- tatomidae), called the southern green stink bug in the U.S., is native to the Ethiopian Region (Jones 1988, Ann. Entomol. Soc. Am. 81: 262-273; Hokkanen 1986, Ann. Entomol. Fenn. 52: 28-31), but now is a worldwide pest ofa multitude of crops (Todd 1989, Annu. Rev. Entomol. 34: 273-292). Range expansion of N. viridula is ongoing although its distribution has long been cos- mopolitan, including tropical areas of Asia, northern Africa, Europe, and the Americas by the 18th century (Hokkanen 1986). Most recently, the southern green stink bug has invaded the Sacramento Valley in Califor- nia (Hoffmann, Wilson, and Zalom 1987, Calif. Agric. 41: 4-6). On October 4, 1989, an adult N. viridula female was collected by sweep net in a soy- bean plo: at the South Farm, Agricultural Research Center, Beltsville, Maryland. This locale is over 600 km north of the normal range for the southern green stink bug in the eastern U.S. (Jones and Sullivan 1981, En- viron. Entomol. 10: 409-414); **. . . records for more northern states, like Ohio, New York, and Virginia, probably are adventi- tious occurrences” (Froeschner, /n Henry and Froeschner 1988, Catalog of the Het- eroptera, or True Bugs, of Canada and the Continental United States, E. J. Brill: 588). Extensive sweeping of the same and adja- cent soybean plots during the following two weeks yielded no additional specimens of N. viridula, nor were any southern green stink bugs collected in soybean during this period from weekly sampling in Wicomico Mention of commercial products does not constitute an endorsement by the U.S. Department of Agricul- ture. County, on the eastern peninsula of Mary- land (T. C. Elden, pers. comm.). Several adults and late instars of Euschistus spp. stink bugs were captured at the South Farm, and fifth instars and adults of the green stink bug, Acrosternum hilare (Say), were excep- tionally common on the eastern shore of Maryland. The errant Nezara female was housed in the laboratory insectary (16:8 h L:D, 28°C, 65% RH) and proceeded to ovi- posit fertile egg masses on October 7, 13, 19, 23, 30, and November 13. Approxi- mately half of the eggs in the sixth mass were infertile and the female died within a week of the last oviposition date. Shortly before midnight, September 21, a hurricane with winds in excess of 215 km/h made landfall near Charleston, South Ca- rolina, and moved rapidly to the northwest (Fig. 1 insert; Weekly Climate Bulletin No. 89/38, U.S. Department of Commerce). Though the eye of the storm (designated ‘“*Hugo”’) passed east of Maryland through West Virginia, this powerful hurricane en- compassed the entire eastern portion of the continent after landfall (Fig. 1; Satellite Data Service Division, U.S. Department of Com- merce). Thus, the most likely explanation for the appearance of N. viridula in Mary- land is that the insect was swept northward from South Carolina by the intense coun- terclockwise, uplifting winds of the hurri- cane and deposited some 13-14 h later in Maryland. This scenario 1s supported by the knowledge that the insect is capable of sus- tained flight for at least 12 h (Kester and Smith 1984, Entomol. Exp. Appl. 35: 75- 81) and, in fact, has often been collected more than 150 km from land without ab- normally strong winds (ref. in Hokkanen 1986; Baust, Benton, and Aumann 1981, 758 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. | the day before and after landfall. Bull. Entomol. Soc. Am. 27: 23-25; Sparks, Jackson, Carpenter, and Muller 1986, Ann. Entomol. Soc. Am. 79: 132-139). Dissection of five 4. hilare females col- lected October 5 in Wicomico County, Maryland, revealed that these bugs were committed to diapause (large fat body with little ovarian development); a similar con- dition might be expected for the Nezara fe- male in question had she developed in Maryland. Moreover, aeration of 18 F, N. viridula males (ca. 10 days old) produced a pheromone extract that was indistinguish- able by gas chromatography from phero- mone of N. viridula males from the south- eastern U.S. (Aldrich, Lusby, Marron, = September 22 diminishes to tropical storm o: September 21 Charleston, SC Photograph: Satellite view of Hurricane Hugo at ca. 13:30, September 22, 1989. Insert: Path of Hugo Nicolaou, Hoffmann, and Wilson 1989, Na- turwissenschaften 76: 173-175). This record of a fertile N. viridula female about 750 km north of its probable point of origin near Charleston, South Carolina, provokes the question: Will the southern green stink bug, and other insects of similar distribution in the southeastern U.S., be en- countered farther north next season than usual? In describing the long-range dispersal of certain aquatic Heteroptera via hurri- canes, Herring (1958, Pan-Pac. Ent. 34: 174, 175) emphasized that “Hurricanes are not rare phenomena but occur with amazing frequency in the tropics and provide a dy- namic means of distributing organisms.” As VOLUME 92, NUMBER 4 such, hurricanes should be included among Wellington’s “exploitable kinds of weather” available to insects for the evolution of dis- persive adaptations (1983, Bull. Entomol. Soc. Am. 29: 24-29). I thank Dr. Owen Thompson, Depart- ment of Meteorology, University of Mary- land, College Park, for helpful discussions about hurricanes and for showing me the satellite video of the landfall of hurricane Hugo. I am especially grateful to Thomas C. Elden, USDA-ARS, Germplasm Quality 759 and Enhancement Laboratory, Beltsville, for informing me of stink bug infestations in soybean and helping to collect bugs. Tom Elden also reviewed the manuscript, as did Kent D. Elsey and Thomas J. Henry, for which I am grateful. Jeffrey R. Aldrich, USDA-ARS, Insect Chemical Ecology Laboratory, Agricultural Research Center-East, B-467, Beltsville, Maryland 20705. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 760-764 A NEW SPECIES OF STENONEMA (EPHEMEROPTERA: HEPTAGENIIDAE) FROM NORTH CAROLINA W. P. MCCAFFERTY Department of Entomology, Purdue University, West Lafayette, Indiana 47907. Abstract. —The new species, Stenonema lenati McCafferty, is described and figured from larval specimens taken from unpolluted reaches of Piedmont rivers in North Carolina. The new species has a distinctive dorsal color pattern and is easily keyed from other Stenonema larvae. It apparently belongs to Cluster IIIB of the subgenus Maccaffertium and is perhaps most closely related to Stenonema mexicanum. It and some other Sten- onema species demonstrate highly restricted geographic and ecological distributions. Key Words: Stenonema larvae that were taken in larg- er rivers of the North Carolina inner coastal plain but that could not be definitively iden- tified to any known species using Bednarik and McCafferty (1979) were first found and brought to my attention in 1985. Subse- quent examination of this material revealed that these populations represent a new spe- cies of Stenonema, the second of which to have been discovered since the extensive revision of the genus by Bednarik and McCafferty (1979). Its discovery, much like that of Stenonema bednariki McCafferty in Kentucky and Missouri (McCafferty 1981), was precipitated by aquatic biologists con- ducting water quality studies and then being able to recognize enigmatic populations through the use of comprehensive taxo- nomic literature. The new species is based on the larval stage alone since adults have yet to be reared. The species-level morphological characters in Stenonema, however, are primarily and sometimes entirely based on the larval stage. Whereas larvae tend to have specific char- acterization, perhaps related to adaptations to specific running water habitats and ecol- mayflies, Heptageniidae, Stenonema, new species ogy, adults have been less prone to mor- phological evolution, possibly due to the relatively young geological age of the group as hypothesized by Bednarik and Mc- Cafferty (1979). The new species belongs to the subgenus Maccaffertium. | name it in honor of David Lenat in recognition of his collecting of the new material. Stenonema lenati McCafferty New SPECIES Fig. 1 Larva (in alcohol).—Mature length ex- cluding caudal filaments: 11.0 mm (male) to 14.0 (female). Head: Vertex (Fig. 1) generally medium to dark brown with fine, pale speckling and with small pale area anterior to median ocellus, pair of pale areas extending between compound eyes and lateral ocelli, large pale areas anterolateral and lateral to compound eyes, and 3 pale areas along posterior mar- gin between compound eyes, sometimes be- coming continuous; narrow brown band sometimes extending laterad of each com- pound eye, dividing anterolateral pale areas from lateral pale areas; lateral and posterior VOLUME 92, NUMBER 4 Fig. 1. Stenonema lenati, larva. pale areas obliterated to large degree in ma- ture males by well-developed compound eyes. Scapes of antennae light; pedicels and bases of flagella dark brown; remainder of 761 flagella light. Mandibles each with 8-10 teeth on inner margin of outer incisor. Maxillae each with 1-3 spinelike setae on crown of galealaciniae (usually 2, with pectination 762 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON poorly developed on subapical spinelike seta and hardly developed on terminal seta), and with 25-35 hairlike setae on galealacinial crown, and 19-26 setae in submedial row of galealaciniae. Thorax: Nota (Fig. 1) generally medium brown; pronotum with pale lateral and an- terolateral marginal areas and pair of dis- tinctive sublateral, pale spots at anterior margin. Legs (Fig. 1) generally pale with brown banding dorsally; all tarsi with single, brown band at midlength; fore- and mid- tibiae with single, broad bands at approxi- mately midlength, most prominent on fore- tibiae; hindtibiae without bands; all femora with pair of broad bands, sometimes ap- pearing diffuse or coalescing with each oth- er, more basal bands of mid- and hindfem- ora often broken and appearing more as 2 large markings. Dorsal armature of forefe- mora consisting of small, mostly blunt, spinelike setae scattered over apical 1 to 4 of surface; posterior margin of forefemora with short, spinelike setae and long, hairlike setae; anterior margin without distinct rows of either spinelike or hairlike setae. Fore- tarsal claws adenticulate. Hindfemora not conspicuously broader than fore- and mid- femora. Abdomen: Dorsally (Fig. 1) pale with con- spicuous dark brown and light brown pat- terns: terga 1-5 with variable markings but often with triangulate anteromedial mark, pair of submedian small spots, and some sublateral markings, sometimes diffuse. Some individuals, however, with terga | and 2 generally light; tergum 3 almost complete- ly dark; tergum 4 with large, dark, sublateral markings; and tergum 5 with pair of small, dark, submedian spots and posterior, dark brown transverse band. Terga 6-10 more consistently patterned among individuals, varying mostly in degree and extent of pig- mentation: terga 6 almost entirely brown to dark brown; terga 7-9 with conspicuous, dark, transverse bands at posterior margin, often dark, medial, longitudinal stripe with shape somewhat variable, darkened lateral areas under gills, and pair of small, sub- median spots; tergum 10 with dark pig- mentation covering anywhere from poste- rior half to virtually entire tergum. Rows of well-developed spinules along posterior borders of all terga. Lateral patches of light, hairlike setae well developed on terga 3-7. Ventrally, sterna 1-6 pale, without mark- ings; sterna 7 and 8 entirely pale with no markings to somewhat ferruginous with small, dark, anteromedial, triangulate markings (anterior markings usually seen beneath overlapping edge of preceding ster- num) and narrow, diffuse shadings or mark- ings; sternum 9 of females ranging from pale to ferruginous with markings ranging from having no anterolateral markings and pair of sublateral, dark spots at posterior margin to having anterolateral markings and well- developed dark band following rather horseshoe shaped distal margin, darkest at sublateral areas corresponding to spotted areas of lighter individuals; sternum 9 of males instead usually with pair of diffuse to well-defined spots sublaterally and subter- minally and diffuse, subterminal, transverse band (tips of developing genital forceps also shaded with ferruginous in mature individ- uals). Posterolateral projections not devel- oped on segments 1-5; projections on seg- ment 8 slightly larger than those of segment 7 and much larger than those of segment 10. Gills 1-6 truncate apically; gills 7 un- tracheated. Caudal filaments uniformly fer- ruginous, with short, spinelike setae and long, hairlike setae well developed on each segment. Holotype.—Mature male larva, North Carolina: Stanley Co.: Rocky River at Nor- wood, March, 1985, D. R. Lenat, deposited in the Purdue Entomological Research Col- lection (PERC), West Lafayette, IN. Paratypes. — Five mature female and four mature male larvae (some mouthparts slidemounted), same data as holotype, all in PERC except one mature male larva at United State National Museum, Washing- ton, D.C. VOLUME 92, NUMBER 4 Additional material examined.—Ten lar- vae, North Carolina: Chatham Co., Haw River, May, 1985, D. R. Lenat, in PERC; nine larvae, North Carolina: Moore/Ran- dolph Cos., Deep River, August, 1985, D. R. Lenat, in PERC. Discussion.—The dorsal color pattern of abdominal segments 6-10 (Fig. 1) found in S. lenati is unlike that of any other Steno- nema. Many species have a darkened ter- gum 6, e.g. S. pulchellum, S. mexicanum, and S. exiguum: however, their following terga differ considerably from those of S. lenati. The pattern of sternum 9 of either males or females are also distinctive but are subject to considerable individual variabil- ity. Sternum 9 of S. femoratum may have a darkened, horseshoe-shaped, posterior margin similar to that of the female larva of S. /enati, but the remainder of the ventral abdominal pattern is very different. The pattern of the vertex and pronotum together can also be used to diagnose this species, particularly when taken in combination with other characters. By using the larval key of Bednarik and McCafferty (1979) and ignoring the color patterns mentioned above, one would tend to key S. /enati to couplet 18 and S. integ- rum [= S. mexicanum (see McCafferty 1984)]. This is due to the large number of hairlike setae but only one or two spinelike setae found on the crown of the galealacinia. Couplet 18 should be modified into a triplet as follows to accommodate identification of S. lenati: 18. Abdominal terga 7, 8, and 9 together with a distinct V-shaped pale area; galealacinial crown with 2-3 spinelike setae .. S. mexicanum — Abdominal terga 7, 8, and 9 each pale with darkened lateral areas, posterior borders, variable median stripe, and pair of subme- dian, small spots; galealacinial crown with |- 3 spinelike setae (usually 2 and with only | conspicuously pectinate) ............. S. lenati — Abdominal terga 7, 8, and 9 with various pat- terns but not exactly as either of above pat- terns; galealacinial crown with 3-7 spinelike setae (if only 3, then terga 8 and 9 mostly dark) 763 It is difficult to determine the exact re- lationships of S. /enati. It is definitely a member of the subgenus Maccaffertium, sharing truncated gills 1-6 and untracheated gills 7 with other members of that subgenus. The new species probably belongs to species Cluster III (Bednarik and McCafferty 1979) because of the loss of posterolateral projec- tions on anterior segments of the larval ab- domen. This proposition, however, is con- tingent on male adult penes and eye separation characters proving to be consis- tent with others in this phyletic position. This appears likely because, based on its larval characteristics, the species would fur- thermore belong to Cluster IIIB, a group which retains considerable hairlike setae on the galealacinial crown. Cluster IIB also in- cludes S. modestum, S. smithae, and S. mexicanum. The reduction of spinelike setae on the galealacinial crown of S. mexicanum and S. lenati suggests a possible sister relationship of these two species. Based on published distributions, the geographically restricted S. lenati apparently does not overlap with the relatively very widespread S. mexica- num (see Bednarik and McCafferty 1979 under S. integrum). This suspected allopa- try is supported by collecting records of D. R. Lenat (pers. comm.), who has not col- lected the two species together in North Ca- rolina. Stenonema lenati has been taken only in coastal plains, whereas records of S. mex- icanum in North Carolina are from other regions of the state. Stenonema lenati larvae appear to be lim- ited to unpolluted sections of Piedmont riv- ers near the fall line. They have been found in slower currents and have been taken co- habiting with the following other heptagen- iid mayflies: Stenonema exiguum, S. fe- moratum (rarely), S. modestum, S. terminatum, Stenacron interpunctatum, S. pallidum, Heptagenia marginalis, and Leu- crocuta aphrodite. With the discovery of S. /enati, there are now four species of Stenonema known to 764 have highly restricted distributions and what appear to be highly restricted ecological re- quirements. The other three are S. bedna- riki, S. carlsoni, and S. sinclairi. All of these are clean-water species (Lewis 1974, 1979, McCafferty 1981) and may be in jeopardy of survival because of narrow ecological tol- erances. Of the four, only S. /enati is found in large rivers, the others occurring in small- er streams. Collections of S. /enati larvae were made in March, May, and August. Only the March collections included mature individuals with darkened wingpads. May and August sam- ples contained predominantly middle instar larvae and no mature individuals. While such data suggest an early spring emergence and possibly suggest another mid-summer emergence, data are too limited to draw def- inite conclusions about voltinism and other aspects of phenology at this time. ACKNOWLEDGMENTS I thank D. R. Lenat of the North Carolina Department of Natural Resources and Community Development, Division of En- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vironmental Management, Raleigh, for pro- viding study material of the new species. I also thank D. W. Bloodgood and A. V. Pro- vonsha, Purdue University, for their assist- ance. This paper has been published as Pur- due University Agricultural Experiment Station Journal No. 12348. LITERATURE CITED Bednarik, A. F. and W. P. McCafferty. 1979. Biosys- tematic revision of the genus Stenonema (Ephem- eroptera: Heptageniidae). Can. Bull. Fish. Aquat. Sci. 201: vi + 73 pp. Lewis, P. A. 1974. Three new Stenonema species from eastern North America (Heptageniidae: Ephemeroptera). Proc. Entomol. Soc. Wash. 76: 347-355. 1979. A new species of the mayfly genus Stenonema Traver from eastern United States (Ephemeroptera: Heptagentidae). Proc. Entomol. Soc. Wash. 81: 321-325. McCafferty, W. P. 1981. A distinctive new species of Stenonema (Ephemeroptera: Heptageniidae) from Kentucky and Missouri. Proc. Entomol. Soc. Wash. 83: 512-515. 1984. The relationship between North and Middle American Stenonema (Ephemeroptera: Heptageniidae). Gr. Lakes Entomol. 17: 125-128. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 765-770 DESCRIPTIONS OF THE LARVA AND PUPA OF FLAVOHELODES THORACICA (GUERIN-MENEVILLE) WITH NOTES ON APHYTOTELMA ASSOCIATION (COLEOPTERA: SCIRTIDAE) JAMES B. STRIBLING AND DANIEL K. YOUNG (JBS) Department of Biology, 37th and O St., NW, Georgetown University, Washington, D.C. 20057; (DKY) Department of Entomology, 1630 Linden Drive, University of Wis- consin, Madison, Wisconsin 53706. Current address: (JBS) EA Engineering, Science, & Technology, 15 Loveton Circle, Sparks, Maryland 21152. Abstract. — Descriptions of the larva and pupa of Flavohelodes thoracica (Guérin-Méne- ville) are given and based on specimens collected and reared from a phytotelma (water- filled treehole) in a Quercus sp. These descriptions are presented in order to facilitate the recognition of immature F. thoracica and to document the occurrence of this species in phytotelmata. Morphological comparisons are made with F. /flavicollis (Kiesenwetter). The status of phytotelmata as a habitat association with the genus is briefly discussed. Key Words: Scirtidae larvae inhabit a wide variety of restricted lentic habitats including phyto- telmata (e.g. water-filled treeholes, tank- forming Bromeliaceae); shallow, leafy groundpools; and leaf packs in nearly still waters of stream sidepools. Larvae are rel- atively easily collected from these habitats, occasionally in large numbers. In spite of this and the relative ease of rearing, life stages or species/habitat associations have been published for few species of Scirtidae. Among those treated in detail are Priono- cyphon discoideus Say (Osten-Sacken 1862, Snow 1958a, b), P. serricornis Miller (Ben- ick 1924, Rohnert 1951, Horion 1955, Kitching 1969, 1971, 1983), P. niger Kitch- ing and Allsopp (1987), Scirtes championi Picado (1913), and Flavohelodes flavicollis (Kiesenwetter) (Klausnitzer 1980). Klausnitzer (1974) recognized the Elodes flavicollis species group as distinct from the remaining species of the genus. Later, he (Klausnitzer 1980) elevated this species immature stages, phytotelmata, Scirtidae, Flavohelodes group to generic status, naming it Flavo- helodes. This action was partially based on larval mouthpart and antennal modifica- tions which he interpreted as adaptations to phytotelmata. Klausnitzer (1980) also re- ported the larval stages of F. flavicollis (Kie- senwetter) as having been recently discov- ered inhabiting water-filled treeholes in Europe. This represents the sole source of published microhabitat data for the genus until now. In this paper, we add another with the description of the larval and pupal stages of Flavohelodes thoracica (Guérin- Méneville) which were collected and reared from a Quercus phytotelma near Great Falls, Montgomery Co., Maryland. METHODS Collection and rearing. — Loose leaves and the leaf pack at the bottom of the treehole were removed by hand and long forceps and sorted in a white porcelain pan. Some larvae were immediately placed into vials contain- 766 Rigs: 15) 2: Flavohelodes thoracica (Guérin-Méne- ville), 1. larva, dorsal habitus, dotted lines on prono- tum indicate patterns of melanization, the median, lon- gitudinal vitta and the anterolateral arms being lighter; 2. pupa, dorsal habitus. ing 70% ethanol for preservation. The re- mainder were segregated into plastic contain- ers with leaf litter and water from the phytotelma, and returned to the lab. Water was removed by siphoning with rubber lab- oratory tubing or commercial basting syringe. Larvae to be reared were placed into 4— 6 dram glass, stoppered vials. Each vial was partially filled with treehole water and a few leaf fragments from the orginal treehole. Personal observations have demonstrated that mature larvae crawl out of the water to pupate, attaching the abdominal apex to the leaf surface. Since attachment to the glass walls of a vial has not been observed, it may be essential to orient the leaf fragments so that portions of them protrude well above the level of the water. Dissections and illustrations. —Dissec- tions were performed in 70% ethanol. Dis- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs; 3514: Flavohelodes thoracica (Guérin-Méne- ville), 3. larva, abdominal terga 8-9, dorsal view; 4. pupa, apical abdominal segments, dorsal view. sected structures were cleared in an aqueous KOH solution, rinsed in distilled water, and observed in glycerine. The dissected struc- tures were stored in genitalic microvials with a drop of glycerine; these microvials were than placed in glass alcohol vials along with the remainder of the specimen. Habitus illustrations were drawn with the aid of a camera lucida attached to a Wild MSA dissecting microscope. Mouthpart il- lustrations were prepared using a micropro- jector. Actual preparation of dissections for illustration were with the K-Y procedure as described by Young and Stribling (1990). In the following description, terms for mouthpart and abdominal characters of the larva were translated directly from or mod- ified after Hannappel and Paulus (1987). Some terms in the pupal description were adopted from Rozen (1963). Voucher specimens are deposited in the personal collections of the authors (DYCC, Madison, Wisconsin and JBSC, Wood- bridge, Virginia). Flavohelodes thoracica (Guerin-Meneville) Description. — Mature larva (Figs. 1, 3, S— 12). Body size.— Length 6-10 mm, most spec- imens 7-9 mm; maximum width (usually in thoracic area) 1-2 mm. Head.— Antennal scape ca. % length ped- VOLUME 92, NUMBER 4 767 Figs. 5-8. Flavohelodes thoracica (Guérin-Méneville), larva, 5. basal antennal segments, dorsal view; 6. left maxilla, ventral view; 7. metathoracic leg; 8. mesothoracic tarsus and tarsungulus. icel (Fig. 5), 37-43 flagellomeres (Fig. 1), scape with larger and approx. 2 as many setae on medial surface as on lateral aspect; pedicel with 10-12 small, unevenly scat- tered setae, apically with 4-6 small setae, and a single, very large, stout seta mesally on apex (Fig. 5); 3 non-melanized stemmata either side of cranium in a dorsal, anterior- posterior row; anterior stemma separated from posterior pair by melanized band; row of large subocular setae; clypeolabrum (epi- pharynx) (Fig. 10) with ventral lobes not protruding anteriorly past margin of clypeo- labral forewall, 6 teeth anterad of 5 sub- dental sensilla; mandibular prostheca (Figs. 11, 12) of 2 main sections, proximal sclerite with row of setae, and large, distal com- pound macroseta with main body socketed to the mandible and 2-3 distal subdivisions; mola cleft proximally; maxilla (Fig. 6) with 3 palpomeres, 5 galeal comb setae visible in ventral view; lacinial setal area of short, dense pile of mesally directed setae; hypo- pharynx (Fig. 9) with two apically notched hypopharyngeal teeth, setigerous sclerite (Fig. 9a) with two long setae and four cam- 768 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-12. Flavohelodes thoracica (Guérin-Méneville), larva, 9. hypopharynx, dorsal view; a—setigerous sclerite, b—grip apparatus, c—comb plate, d—central compress; 10. clypeolabrum (epipharynx), ventral view; 11. left mandible, mola, and prostheca, ventral view; 12. left mandible, ventral view. paniform sensilla, usually arranged in row (specimen figured has one lateral sensillum apically offset); one pair of medial campan- iform sensilla proximad to setigerous scler- ite; comb plate (Fig. 9c) with oval setal patch bearing | 8-20 teeth; grip apparatus (Fig. 9b) covered anterolaterally with short, fine setae, posteriorly covered with thicker, more dis- tinct setae (much longer on mesal margin); setal cluster near the anterior margin be- tween setigerous sclerite and grip apparatus; central compress (Fig. 9d) wing-shaped, completely covered with tiny tubercles; la- bium broad, each palpus of 2 palpomeres. Thorax (Fig. 1).—Mediodorsal length ra- tio (pro-: meso: metathoracic nota) 2:1:1; paler bands of melanization extend from an- terolateral pronotal angles to midpoint of posterior pronotal margin, meso- and me- tathoracic nota somewhat paler at middorsal line; notal setation sparse except at margins, setae usually of increasing length from an- terior to posterior angles; nota punctate, punctations evenly distributed, each with single, short fine seta; prothoracic legs (Figs. 7, 8) slightly shorter than meso- and meta- thoracic legs, other features invariant; coxae (Fig. 7) transverse (some appear conical if larva has inflated in preservation), sparsely setose, most setae dispersed on ventral and anterior surfaces; femur with two longitu- dinal, parallel rows of strong setae on outer VOLUME 92, NUMBER 4 surface, continuing around tibial insertion; tibia with stout setae on lateral and mesal surfaces, setation sparse anteriorly and pos- teriorly; tarsungulus (Fig. 8) with two strong, subapical setae; meso- and metathoracic sterna with intercoxal patches of long setae. Abdomen (Fig. 1).—Eight visible seg- ments, gradually decreasing in circumfer- ence and increasing in mediodorsal length posteriorly; punctation and setation similar to thoracic nota; tergum eight (Fig. 3) emar- ginate and slightly concave apically, ninth tergum as in Fig. 3. Pupa (Figs. 2, 4).—Body length 5.5 mm; head smooth; dorsally covered with short, fine setae concolorous with cuticle; prono- tum with two anterior and two posterior spines (“‘tubercles”’ of Rozen [1963]); elytra long, approx. ' body length, projecting pos- tero-laterally,; nine abdominal segments, segment nine with short, bifurcate, terminal projection, ninth tergum (Fig. 4) with blind posterior pit, bounded dorsally by sclerite with 2 acute, lateral apices, and ventrally with 2 small spines. Material examined (and _ repositories): MARYLAND, Montg. Co., near Great Falls, 02 June 1987, J. B. Stribling & W. E. Steiner/reared from larva in water from tree hole (14 larvae, 2 exuviae, | pupa, 3 adults; DYCC); (same data)/in water from tree hole (13 larvae; 7 DYCC, 6 JBSC). Comparisons with Flavohelodes flavicollis (Kiesenwetter) Comparisons of F. thoracica are based on descriptions in Hannappel and Paulus (1987). Some structures are unavailable for comparison because they have not been de- scribed in prior publications and the lack of specimens of F. flavicollis. Those which are common to our studies are discussed here. Hypopharynx (Fig. 9).— Near the anterior margin between the grip apparatus and the setigerous sclerite there exists a cluster of setae which is lacking in F. flavicollis. A further difference is the existence of a row of heavier spines or setae on the anterior 769 surface of the grip apparatus in F. flavicollis that is lacking in F. thoracica. Clypeolabrum (Fig. 10).—In F. flavicollis, the ventral lobes are protruding past the an- terior margin, there are 6 subdental sensilla, the setation of the anterior lobes 1s more uniform and stouter, the pair of medioan- terior setae is short, and the lateral setation is abundant. In contrast, in F. thoracica, the ventral lobes do not protrude past the an- terior margin, there are 5 subdental sensilla, the setation of the anterior lobe is less uni- form and less stout, the pair of medioanterior setae 1s longer, and lateral setation is lacking. Mandible (Figs. 11, 12).—The main spe- cies differences exist in the form of the pros- theca. In F. flavicollis this structure is not setose and does not have an associated com- pound macroseta as is found in F. thoracica. Maxillary palpus (Fig. 6).—Setal abun- dance and distribution are similar in the two species. In palpomere | of F. thoracica se- tation is more proximal than that of F. fla- vicollis which is more evenly and longitu- dinally distributed. Status of the Flavohelodes/phytotelmata association In his paper elevating the flavicollis spe- cies group to generic status, Klausnitzer (1980) hypothesized phytotelmata use as an ecological synapomorphy. Here, we have documented hardwood phytotelmata as a habitat for larval development of F. thor- acica in support of this hypothesis. Al- though this study confirms phytotelmata use, we still know little of the degree of habitat specificity for this species. For example, are phytotelmata necessary for F. thoracica de- velopment or is it only one of the many potential breeding sites? Frank and O’Meara (1985) found habitat segregation by oviposition site preference in bromeliad-breeding Wyeomyia spp. (Dip- tera: Culicidae) and attributed this to both microhabitat and macrohabitat differences. Preference of Tillandsia utriculata Linnaeus over Catopsis berteroniana (Schultes) Mez 770 ex de Candolle (Bromeliaceae) phytotel- mata for oviposition was interpreted as a microhabitat difference. In another experi- ment reported in the same paper, brome- liads in shaded habitat were found to be preferred for oviposition over those in non- shaded locations. The shaded vs. non-shad- ed habitats were interpreted as macrohabi- tat differences. In order to categorize our F. thoracica- phytotelma association as microhabitat or macrohabitat, in the sense of Frank and O’Meara (1985), as well as further testing of Klausnitzer’s (1980) hypothesis, more must be known about the ecological distribution of this species. Further collection and rearing of scirtid larvae from hardwood phytotel- mata and other potential breeding sites, with documentation of habitat specifics, will al- low these questions to be answered. ACKNOWLEDGMENT We are grateful to W. E. Steiner (De- partment of Entomology, Smithsonian In- stitution, Washington, D.C.) for accompa- niment in the field and for his suggestions; to P. J. Spangler (Entomology, Smithsonian Institution) and D. R. Whitehead (USDA— Systematic Entomology Laboratory, Wash- ington, D.C.) for providing space and equip- ment; and, to M. Rodon-Naviera, P. Thaler, and E. Barrows (Georgetown University) for comments and suggestions on earlier drafts of the manuscript. We thank several anon- ymous reviewers for well-taken suggestions. LITERATURE CITED Benick, L. 1924. Zur Biologie der Kaferfamilie Helo- didae. Mitt. Geog. Gesell. Naturhist. Mus. Liibeck (2)29: 45-47. Frank, J. H. and G. F. O’Meara. 1985. Influence of micro- and macrohabitat on distribution of some bromeliad-inhabiting mosquitoes. Entomol. Exp. Appl. 37: 169-174. Hannappel, U. and H. F. Paulus. 1987. Arbeiten zu PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON einem phylogenetischen System der Helodidae (Coleoptera)— Feinstrukturuntersuchungen an eu- ropdischen Larven. Zool. Beitr. N. F. 31: 77-150. Horion, A. 1955. Faunistik der Mitteleuropaischen Kafer. Band 4. Eigenverlag, Miinchen. Kitching, R. L. 1969. A preliminary note on the fauna of water-filled tree-holes. Entomologist 102: 7-9. . 1971. Anecological study of water-filled tree- holes and their position in the woodland ecosys- tem. Ecology 40: 281-302. 1983. Community structure in water-filled treeholes in Europe and Australia—comparisons and speculations. /n Frank, J. H. and L. P. Louni- bos, eds., Phytotelmata: Terrestrial Plants as Hosts for Aquatic Insect Communities. Plexus Publ., Inc., Medford, N.J. 293 pp. Kitching, R. L.and P.G. Allsop. 1987. Prionocyphon niger sp. n. (Coleoptera: Scirtidae) from water- filled treeholes in Australia. J. Aust. Entomol. Soc. 26: 73-79. Klausnitzer, B. 1974. Anwendung der Phylogene- tischen Systematik innerhalb von Gattungen, dar- gestellt am Beispiel der Gattung Helodes Latreille 1796 (Coleoptera, Helodidae). Zool. Jb. Syst. 101: 479-559. 1980. Bemerkungen zur Kenntnis der He- lodidae Bulgariens mit Beschreibung einer neuen Art der Gattung Helodes Latreille. Entomol. Blatt. 76(1): 58-64. Osten-Sacken, R. 1862. Descriptions of some larvae of North American Coleoptera. Proc. Entomol. Soc. Phil. 1: 105-130. Picado, C. 1913. Les Bromeliacées epiphytes consi- derées comme milieu biologique (1). Bull. scien- tifiques France et Belg. 47: 215-360. Rohnert, U. 1951. Wassererfullte Baumhohlen und ihre Besiedlung. Ein Beitrag zur Fauna dendrolim- netica. Arch. Hydrobiol. 44: 472-516. Rozen, J. F. 1963. Preliminary systematic study of the pupae of Nitidulidae (Coleoptera). Amer. Mus. Novit. 2124: 1-13. Snow, W. E. 1958a. Stratification of arthropods in a wet stump cavity. Ecol. 39: 83-88. 1958b. Vertical and horizontal stratification of insect communities in wet tree cavities. Proc. 10th Int. Congr. Entomol. 2: 697. Young, D. K. and J. B. Stribling. 1990. Systematics of the North American Cyphon collaris species complex with the description of a new species (Co- leoptera: Scirtidae). Proc. Entomol. Soc. Wash. 92: 194-204. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 771-772 NOTE Andrena macra Mitchell (Hymenoptera: Andrenidae) Overwinter and Delay Spring Emergence in Virginia Andrena macra Mitchell are known from West Virginia and Maryland southward to Florida, and westward to Texas and Okla- homa (LaBerge 1985. Trans. Amer. Ento- mol. Soc. 112(3): 191-248). The only bio- logical investigation of this species was done by Sivik (1954. Entomol. News 65(10): 253- 256), where he witnessed the emergence of adults, and determined the distribution of nests in a site near East Raleigh, North Ca- rolina. The purpose of this note is to present some unusual observations on the life cycle of this solitary, soil-nesting bee. The nesting site observed in this study was adjacent to a graveled off-road on the United States Marine Corps Reservation in Quantico, Virginia. Andrena macra nests were excavated so that the overwintering brood of this bee could be observed. Ex- cavations were conducted on the following days of each month: 15, 17, 19, 31 March 1984; 7, 12, 26 April 1984; and 2, 10, 11 May 1984. Brood were removed from the nests with a microspatula and placed, in- dividually, inside 4-dram glass shell vials. The numbers of live A. macra found on daily excavations beneath the surface of this site during Winter and Spring 1984 are list- ed in Table 1. A total of 59 overwintering prepupae were kept in the laboratory at room temperature (24-27 degrees Celsius), inside the shell vials, to see ifand when they would complete their development to the mature adult stage. Eleven were reared successfully (Table 2). Prepupae required 78 + 8.4 days (mean and standard error) to metamor- phose into pupae. These pupae required 21 + 1.3 days to metamorphose into mature adults. Andrena macra overwintered in the pre- pupal and adult stages of development. Spe- cies of Andrena were previously known to overwinter only as adults (Linsley 1958. Hilgardia 27(19): 543-599). Some of the overwintering 4. macra also delayed spring emergence in 1984. The evidence in favor of delayed emergence is as follows: (1) pre- pupae were found inside nest brood cells on the same days that adults had emerged and were present on the surface of the nesting site, and (2) prepupae that were found in March and May completed their develop- ment to the mature adult stage near the end, or after the end, of the Spring 1984 nesting season (see Table 2). All adult nesting ac- tivities had ceased by mid-June 1984. Delayed emergence has been reported for several species of Andrena distributed in the western United States. Andrena mojavensis Linsley and MacSwain and Andrena om- ninigra clarkiae Linsley and MacSwain de- layed spring emergence in California and these spring seasons were unfavorable for host flower blooming (Linsley et al. 1964. Univ. Calif. Publ. Entomol. 33(2): 59-98; MacSwain et al. 1973. Univ. Calif. Publ. Entomol. 70: 1-80). Thorp (1979. Ann. Mo. Bot. Gard. 66: 788-812) indicated that a species of Andrena delayed emergence dur- Table 1. The numbers of live Andrena macra found in nest brood cells during daily excavations in 1984. Adults Prepupae Date Found 2 15 15 March 2 17 March 19 March — 31 March 7 April 12 April 26 April 2 May 10 May 11 May — nN oO Co Ne P= wWwnwoneo —-lruanl 772 Table 2. Results of the rearing experiment in 1984. Sex of Reared Date Date at Mature Andrena Found *P=P Adult Stage macra 15 March 40 4 June F 15 March 49 6 June M 15 March 51 13 June M 15 March 51 18 June M 15 March 70 8 July M 15 March 83 15 July M 15 March 83 17 July FE 31 March 126 30 August Ip 2 May 95 27 August M 2 May 98 29 August M 10 May 109 23 September 12 * P-P = number of days required for prepupae to metamorphose into pupae. F = female; M = male. ing the time that a drought occurred in Cal- ifornia. Delaying spring emergence in the eastern United States may insure survival for 4. macra especially when they nest in dense PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aggregations. Female 4. macra of large nest- ing sites may have to rely on limited sources of pollen and nectar to provision their nest brood cells with, each spring season. Emer- gence of the individuals of a population in two years, instead of one, will prevent the depletion of floral resources. Dr. W. E. LaBerge (Illinois Natural His- tory Survey) identified the 4. macra. This research was a portion of a M.S. thesis sub- mitted to the graduate faculty (Department of Zoology) at Howard University. Partial support was made available by a Minority Biomedical Research grant (RR-08016) awarded to Drs. R. M. Duffield and J. Wheeler from the Division of Research Re- sources, National Institutes of Health. Eric W. Riddick, Department of Zoology, Howard University, Washington, D.C. 20059. Present address: Department of En- tomological Sciences, University of Califor- nia—Berkeley, Berkeley, California 94720. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 773-777 EFFECT OF ANTIOXIDANTS ON EASTERN SUBTERRANEAN TERMITE (ISOPTERA: RHINOTERMITIDAE) ORIENTATION TO FUNGAL EXTRACT J. KENNETH GRACE Faculty of Forestry, University of Toronto, Toronto, Ontario MSS 1A1, Canada. Cur- rent address: Department of Entomology, 3050 Maile Way, Room 310, University of Hawaii, Honolulu, Hawaii 96822. Abstract. —Dichloromethane and cyclohexane extracts of wood decayed by Gloeophyl- lum trabeum induce trail-following, arrestment, and/or aggregation of Reticulitermes fla- vipes in behavioral bioassays. In trail-following assays with R. flavipes workers, addition of the antioxidant BHA completely suppressed termite response to the fungal extracts. Addition of the antioxidant BHT did not eliminate termite responses to the extracts, but concentration-dependent repellency was noted in orientation (preference) assays with individual termites and groups of workers. As measured by R. flavipes behavioral response, addition of BHT at the concentrations tested did not increase the longevity of the active semiochemicals in G. trabeum extracts. Key Words: icals Wood decayed by the fungus G/oeophyl- lum trabeum (Pers. ex Fr.) Murr. (Basidi- omycetes: Polyporaceae) contains the compound (Z,Z,E£)-3,6,8-dodecatrien-1-ol (Matsumura et al. 1969) and other uniden- tified chemicals (Watanabe and Casida 1963, Ritter and Coenen-Saraber 1969) that affect the orientation behavior of subterra- nean termites (Isoptera: Rhinotermitidae). Solvent extracts of wood decayed by G. tra- beum and other decay fungi (Grace and Wil- cox 1988) elicit both trail-following and ag- gregation in Reticulitermes species (Esenther etal. 1961, Allen et al. 1964, Grace 1989b). Currently, subterranean termites are ex- cluded from buildings by the injection of large quantities of insecticides into the sur- rounding soil. An alternative approach is the development of toxic baits employing decayed wood to contaminate foraging ter- mites and eradicate the colony through Reticulitermes, termite behavior, Gloeophyllum, decay fungus, semiochem- trophallaxis and grooming’ behavior (Esenther and Beal 1979). Natural or syn- thetic chemical termite “‘attractants’’ would offer more flexibility than decayed wood in developing such baits, and toxic analogues of dodecatrienol have been investigated by Carvalho and Prestwich (1984). In addition, subterranean termites are able to follow a chemical gradient (Clement et al. 1988, Grace et al. 1988), and compounds aggre- gating foragers might prove useful in en- hancing the efficacy of pesticides applied to the soil for termite control. The study reported here was undertaken to determine whether addition of the anti- oxidants BHA and BHT to solvent extracts of wood decayed by G. trabeum could in- crease the longevity of the compounds in- ducting a positive orientation response in the eastern subterranean termite, Reticuli- termes flavipes (Kollar). These antioxidants 774 Table 1. Mean (+SE) distance traveled by Reticuli- termes flavipes workers on artificial trails drawn with solvent (controls) and with dichloromethane extracts of Gloeophyllum trabeum decayed red pine containing the antioxidants BHT and BHA. Trails were air-dried 15 minutes, and each mean represents 25 individual assays. Means followed by the same letter are not sig- nificantly different at the 0.05 level (ANOVA, REGW multiple F test). Treatment Mean Distance (mm) G. trabeum 35 + 8b G. t. + BHT (1 mg/ml) 48 + 9ab G. t. + BHT (10 mg/ml) 66 + 12a G. t. + BHA (1 mg/ml) 2ieE alc G. t. + BHA (10 mg/ml) 1+ Ic Dichloromethane Control 3) Ic D.C. + BHT (1 mg/ml) 0 + Oc D.C. + BHT (10 mg/ml) 2 =EN¢ D.C. + BHA (1 mg/ml) leEalc D.C. + BHA (10 mg/ml) 4 + 3c have been found to protect some semio- chemicals with internal conjugated dienes from both oxidation and isomerization (Ideses and Shani 1988, Shani and Klug 1980) and thereby enhance their field life in pest management applications. METHODS Termites. — Foraging eastern subterrane- an termites, R. flavipes, were collected from corrugated paper rolled within short lengths of plastic pipe buried just below the soil Table 2. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON surface at a site in the city of Scarborough, Ontario (Grace 1989a). Prior to their use in bioassays, termites were maintained on cor- rugated paper and filter paper in plastic box- es in an unlighted incubator (27 + 0.5°C, 90 + 5% RH). Fungus extracts.—Red pine, Pinus re- sinosa Ait., decayed for 6-8 weeks after in- oculation with G. trabeum was provided by E. E. Doyle and K. Seifert, Forintek Canada Corp., Ottawa, Ontario. Ten grams of de- cayed wood, ground in a Wiley mill to pass a 40-mesh screen, were shaken in 100 ml dichloromethane or cyclohexane for 15 minutes at room temperature (24°C), and gravity-filtered through Whatman No. | fil- ter paper to yield approximately 70 ml of filtrate. The antioxidants 3(2)-tert-butyl-4- hydroxyanoisole (BHA) and 2,6-di-tert-bu- tyl-4-methylphenol (butylated hydroxytol- uene, BHT) were purchased from Sigma Chemical Co., St. Louis, MO. Trail-following assay.—Three bioassays were used to test the effects of G. trabeum extracts containing either 1, 10, or 100 mg/ ml BHA or BHT on R. flavipes orientation behavior. In the trail-following assay, de- scribed by Grace and Wilcox (1988), a straight 20-cm line was drawn on tracing paper with 4 ul of solution, applied by a microliter syringe. This artificial trail was Mean (+SE) distance traveled by Reticulitermes flavipes workers on artificial trails drawn with dichloromethane and cyclohexane extracts of Gloeophyllum trabeum decayed red pine containing BHT. Trails were air-dried for different time intervals, and each mean represents 25 individual assays. Treatment means with the same solvent in each column followed by the same letter are not significantly different at the 0.05 level (¢ test, or ANOVA and REGW multiple F test). Trail Aeration Time (minutes) Solvent Treatment 15 45 GHG! G. trabeum 47 == lay (30'-£76a GE BAT 9 45<£ 10a 30)-e18a (1 mg/ml) G. t. + BHT 32+ 10a 48+ 8a (10 mg/ml) GHG G. trabeum 18 + 5a G. t. + BHT 22) ea (1 mg/ml) 15 19 60 75 105 120 Whesstys als ceisyy 3a 23) idan see 4a 3a Sich sbe ew lsieesa OF-Es3a + 3b 11 + 3a 59a) 6+ 1b VOLUME 92, NUMBER 4 Us} Table 3. Mean (+SE) number of Reticulitermes flavipes workers in contact with paper disks treated with dichloromethane extracts of decayed red pine containing BHT, during successive five-minute intervals. The positions of 50 workers, tested individually in separate petri dishes, were recorded every 30 seconds, with each mean representing 10 successive 30-second observations. Means within each column followed by the same letter are not significantly different at the 0.05 level (ANOVA, REGW multiple F test). Number of Termites on Treated Papers (Individual Assays) Treatment 0-5 min G. trabeum 28 + la G. t. + BHT (1 mg/ml) 12+ 1b G. t. + BHT (10 mg/ml) 23) Enh G. t. + BHT (100 mg/ml) 7a Dichloromethane Control + Id air-dried from 15 to 135 minutes, and an R. flavipes worker (pseudergate older than the third instar as determined by size) placed at one end. The forward distance traveled on the trail in 30 seconds by the worker was recorded, and the distance traveled by 25 workers on 25 such trails per treatment compared by f test, or analysis of variance (ANOVA) and the Ryan-Einot-Gabriel- Welsch (REGW) multiple F test (SAS In- stitute 1987). Onientation assays.—In addition to the trail-following assays, orientation assays us- ing both individual workers and groups of ten workers were designed after those de- scribed by Grace (1989b) and Grace et al. (1989). In both individual and group assays, 100 ul of G. trabeum extract was applied by pipette to a 23 mm Whatman No. 3 filter paper disk. This disk was paired with a sol- vent-treated disk in a 5-cm diameter glass 5.5-10 min 10.5-15 min 15.5-20 min 17+ la 14+ la 2s 12-5 be Tile 7 += Ub 14 + labe 10 + lb 1 = 1b 14 + lab 10 + lb 9+ 1b Lil + Je 12 = lab 2=E al petri disk, and aired 15 minutes to evapo- rate the solvent. Either an individual ter- mite worker or a group of ten workers was then placed in the dish, and their positions recorded every 30 seconds for 20 minutes. In the individual assays, 50 workers were tested independently in separate petri dish- es. The number of individuals in each treat- ment in contact with an extract-treated pa- per at each 30-second observation were compared for each five-minute interval (n = 10 observations per five-minute interval) by ANOVA and the REGW multiple F test (SAS Institute 1987). A series of control as- says was included in which one of two equivalent solvent-treated disks was arbi- trarily designated the “‘treatment”’ disk. In the group orientation assays, 20 groups of ten workers were evaluated with each treatment. The 30-second observations were pooled over each five-minute interval (n = Table 4. Mean (+SE) number of Reticulitermes flavipes workers in group assays in contact with paper disks treated with dichloromethane extracts of decayed red pine containing BHT, during successive five-minute intervals. The positions of 20 groups of 10 workers were recorded every 30 seconds and pooled for analysis, with each mean representing 200 observations. Means within each column followed by the same letter are not significantly different at the 0.05 level (ANOVA, REGW multiple F test). Number of Termites on Treated Papers (Group Assays) Treatment 0-5 min 5.5-10 min 10.5-15 min 15.5-20 min G. trabeum 3.1 Ola 3.4 + 0.2a 2.6 + 0.2a 2.4+ 0.la G. t. + BHT (1 mg/ml) 3.5 =;0.1lb 2.8 + 0.1b 1.8 + 0.1b 1.3: = Ob G. t. + BHT (10 mg/ml) 535) (SE (05) Lo) 3.2 += 0:la 2.0 + 0.1b 123. Of1b G. t. + BHT (100 mg/ml) 201d Ie: Bee 0)4) Ke. 1:0'+ O:lc 0.9 + 0.1c Dichloromethane Control 24+ 0-1c 1.8 + O.lc ESE ONIc 1.3+0.1b 776 Table 5. Mean (+SE) number of Reticulitermes fla- vipes workers in contact with paper disks treated with a cyclohexane extract of decayed red pine or an extract containing | mg/ml BHT, during successive 5-minute intervals. Treated papers were aired either 15 or 60 minutes before the assay. The positions of 50 workers, tested individually in separate petri dishes, were re- corded every 30 seconds, with each mean representing 10 successive 30-second observations. Treatment pairs (same time interval) within each column followed by an asterisk are significantly different at the 0.05 level (¢ test). Paper Aeration Time Time Interval Treatment 15 min 60 min 0.5—5 min G. trabeum Xan At” CNarah| G. t. + BHT 20) eee Ola 5.5-10 min G. trabeum Liesl 5.1 G. t. + BHT Wests ils;ee il 10.5-15 min G. trabeum Ses 4 G. t. + BHT Seen OES 15.5-20 min G. trabeum 15 I+ I+ * o G. t. + BHT 6 200 observations per five-minute interval), and the mean numbers in contact with the extract-treated papers compared by ANO- VA and the REGW multiple F test (SAS Institute 1987). Both individual and group orientation as- says, in which the test papers were air-dried 15 minutes, were performed with the di- chloromethane G. trabeum extracts, con- taining either 0, 1, 10, or 100 mg/ml BHT. Cyclohexane G. trabeum extracts contain- ing 0 or | mg/ml BHT were compared in individual assays in which the test papers were aired either 15 or 60 minutes. This latter procedure was adopted to preclude termite behavioral habituation to the test extracts (Grace 1989b) concealing differ- ences between them. RESULTS AND DISCUSSION Neither BHA nor BHT alone elicited trail- following by R. flavipes, and addition of BHA to the dichloromethane extract of wood decayed by G. trabeum completely suppressed trail-following (Table 1). Thus, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BHA was not included in the individual and group orientation assays. With short aera- tion periods, addition of BHT did not affect the trail-following activity of the fungus ex- tract (Table 2). Neither were any consistent effects on trail-following activity apparent with longer aeration periods. Addition of BHT did not arrest the observed decrease in extract trail-following activity with trail aeration time. In the individual (Table 3) and group (Ta- ble 4) orientation assays, the BHT fortified extracts exhibited less activity than the dichloromethane G. trabeum extract, and decreased in activity over time. In several instances, the activity of the BHT extracts, particularly that containing 100 mg/ml BHT, fell below that of the solvent control, suggesting concentration-dependent repel- lency. Results of the individual orientation as- says with the cyclohexane extract (Table 5) were comparable to those obtained in the dichloromethane solvent system. Whether the test papers were aired 15 or 60 minutes, addition of | mg/ml BHT either made no difference in comparison to the activity of the stock G. trabeum extract, or resulted in a significant decrease in positive responses to the extract. At the concentrations tested here, addi- tion of BHA to G. trabeum extract resulted in a complete loss of behavioral activity to- wards R. flavipes, while BHT slightly de- creased activity. Neither antioxidant ap- peared to provide protection to the fungal semiochemicals nor extend their longevity. However, the concentrations of behavior- ally active compounds in the G. trabeum extracts were not determined, and very low concentrations of semiochemicals can elicit activity. More encouraging results might therefore be obtained with lower concen- trations of antioxidants. This study indi- cates that BHT, which demonstrated a con- centration-dependent repellency rather than complete suppression of activity, is the more VOLUME 92, NUMBER 4 promising additive for use in such investi- gations. Although studies with isomers and ana- logs of the identified dodecatrienol (Pres- twich et al. 1984 and included citations) have demonstrated that the (Z, Z)-3,6-alkadien- 1-ol functionality is more important than the conjugated 6,8-diene in eliciting trail- following, 6,8-dodecadien-1-ol also shows activity with Reticulitermes (J. K. Grace and M. Kim, unpublished results). Chemical protection of the identified semiochemical in G. trabeum, as well as other unidentified biologically active compounds in the sol- vent extracts, thus deserves further study as an approach to extending the field life of these potentially useful extracts. ACKNOWLEDGMENTS I am grateful to A. Abdallay, J. Inah, and H. Jakimowicz for performing bioassays and help in data tabulation, and E. E. Doyle and K. Seifert (Forintek Canada Corp., Eastern Division, Ottawa, Ontario) for providing the decayed wood. This is part ofa study funded by the Ontario Ministry of the Environ- ment, Ontario Ministry of Housing, Canada Mortgage and Housing Corporation, On- tario Real Estate Association Foundation, Toronto Real Estate Board, and the munici- palities of Toronto, Scarborough, North York, Hamilton, Guelph, Etobicoke, East York, Leamington, Oakville, and Kincar- dine. LITERATURE CITED Allen, T. C., R. V. Smythe, and H. C. Coppel. 1964. Response of twenty-one termite species to aqueous extracts of wood invaded by the fungus Lenzites trabea Pers. ex Fr. J. Econ. Entomol. 57: 1009- 1011. Carvalho, J. F. and G. D. Prestwich. 1984. Synthesis of -triated and -fluorinated analogues of the trail pheromone of subterranean termites. J. Org. Chem. 49: 1251-1258. Clement, J. L., M. Lemaire, P. Nagnan, P. Escoubas, A. G. Bagneres, and C. Joulie. 1988. Chemical ecology of European termites of the genus Reticuli- Vil termes. Allomones, pheromones and kairomones. Sociobiology 14: 165-174. Esenther, G. R., T. C. Allen, J. E. Casida, and R. D. Shenefelt. 1961. Termite attractant from fungus- infected wood. Science 134: 50. Esenther, G. R. and R. H. Beal. 1979. Termite con- trol: Decayed wood bait. Sociobiology 4: 215-222. Grace, J. K. 1989a. A modified trap technique for monitoring Reticulitermes subterranean termite populations (Isoptera: Rhinotermitidae). Pan-Pac. Entomol. 65: 381-384. 1989b. Habituation in termite orientation response to fungal semiochemicals. Sociobiology 16: 175-182. Grace, J. K. and W. W. Wilcox. 1988. Isolation and trail-following bioassay of a decay fungus associ- ated with Reticulitermes hesperus Banks (Isoptera: Rhinotermitidae). Pan-Pac. Entomol. 64: 243-249. Grace, J. K., D. L. Wood, and G. W. Frankie. 1988. Trail-following behavior of Reticulitermes hespe- rus Banks (Isoptera: Rhinotermitidae). J. Chem. Ecol. 14: 653-667. 1989. Behavior and survival of Reticuliter- mes hesperus Banks (Isoptera: Rhinotermitidae) on selected sawdusts and wood extracts. J. Chem. Ecol. 15: 129-139. Ideses, R. and A. Shani. 1988. Chemical protection of pheromones containing an internal conjugated diene system from isomerization and oxidation. J. Chem. Ecol. 14: 1657-1669. Matsumura, F., A. Tai, and H.C. Coppel. 1969. Ter- mite trail-following substance, isolation and pu- rification from Reticulitermes virginicus and fun- gus infected wood. J. Econ. Entomol. 62: 599-603. Prestwich, G. D., W.-S. Eng, E. Deaton, and D. Wich- ern. 1984. Structure-activity relationships among aromatic analogs of trail-following pheromone of subterranean termites. J. Chem. Ecol. 10: 1201- 1217. Ritter, F. J. and C. M. A. Coenen-Saraber. 1969. Food attractants and a pheromone as trail-following substance for the saintonge termite: Multiplicity of the trail-following substances in L. trabea in- fected wood. Entomol. Exp. & Appl. 12: 611-622. SAS Institute Inc. 1987. SAS/STAT Guide for Per- sonal Computers, Version 6 Edition. SAS Institute Inc., Cary, N.C. Shani, A. and J. Klug. 1980. Sex pheromone of Egyp- tian cotton leafworm (Spodoptera littoralis): Its chemical transformations under field conditions. J. Chem. Ecol. 6: 875-881. Watanabe, T. and J. E. Casida. 1963. Response of Reticulitermes flavipes to fractions from fungus in- fected wood and synthetic chemicals. J. Econ. En- tomol. 56: 300-307. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 778-792 REDESCRIPTION AND IMMATURE STAGES OF FICIOMYIA PERARTICULATA (DIPTERA: CECIDOMYIIDAE), A GALL MIDGE INHABITING SYCONIA OF FICUS CITRIFOLIA J. C. ROSKAM AND H. NADEL (JCR) University of Leiden, Dept. of Population Biology, Schelpenkade 14a, 2313 ZT Leiden, Netherlands; (HN) University of Florida, Tropical Research and Education Cen- ter, 18905 S. W. 280th Street, Homestead, FL 33031. Abstract. —Felt described Ficiomyia perarticulata and F. birdi from syconia of Ficus aurea Nutt. and F. citrifolia Mill., respectively, collected in southern Florida. We put Ficiomyia birdi Felt into synonymy because qualitative, quantitative and principal com- ponent analyses revealed no differences between Felt’s type series of F. perarticulata and F. birdi. Felt’s host plant record for F. perarticulata is probably wrong. After extensive sampling we conclude that the host-plant association of Ficiomyia, at least in Florida, is restricted to Ficus citrifolia. We redescribe the adult stages of Ficiomyia perarticulata and describe the immature stages for the first time. Adult males have considerably fewer antennal segments than females, which is otherwise unknown in gall midges. Other pe- culiarities include the development of a spatula in the second larval instar and the de- velopment of sexually dimorphic galls. Females cause longer, stalked galls and males sessile galls. Key Words: Ficiomyia, Ficus, gall midge, immature stage, sexual dimorphism, emer- gence, principal component analysis (PCA), host-plant association, bio- geography One of the most striking examples of mu- tualism among phytophagous insects and their host plants is the association between figs and agaonid fig wasps. This association resulted in patterns of radiation that display remarkable co-evolved traits between the insects and figs. Fig wasps, which pollinate fig flowers and, in turn, use ovaries as larval development sites, are not the only group of insects that inhabit fig fruit. Other Hy- menoptera, as well as gall midges, inhabit syconial galls (Docters van Leeuwen-Reijn- vaan and Docters van Leeuwen 1926, Wil- liams 1928, Barnes 1948, Condit 1969, Mani 1973). Felt (1922, 1934) erected the genus Ficiomyia for midges inducing pocket-like swellings within syconia of figs native to Florida, and described two species, F. per- articulata on Ficus aurea Nutt. and F. birdi on Ficus citrifolia Mill. (= F. laevigata Vahl). Both species are characterized by a large number of antennal segments, a reduced number of palpal segments, and peculiar male genitalia. Felt (1925) placed Ficiomyia in the tribe Dasyneuriariae: Gagné, in McAlpine et al. (1981) placed it, and all other Dasyneuriariae, in the tribe Oligotro- phini. Like Ficus, Ficiomyia may extend into Florida from the Neotropics. Close relatives of the midge are unknown. Because Felt described both species from a few poor and incomplete specimens, he VOLUME 92, NUMBER 4 drew some erroneous conclusions concern- ing species differences and probably report- ed an incorrect host plant association. These errors became apparent after we analysed Felt’s type series and a large sample of fresh specimens reared from authoritatively iden- tified host plants. In this paper we rede- scribe F. perarticulata Felt and place F. birdi into synonymy. Extensive sampling of the purported host plant species native to Flor- ida, and a comparison of both of Felt’s type series with the midges we reared, show that his material, attributed to two species, is conspecific. We also described the imma- ture stages of Ficiomyia, which, like the adults, possess some remarkable traits. Lastly, we described some aspects of the midge’s biology, including emergence be- havior and periodicity, mating, and the sex ratio. METHODS We studied newly reared material and compared it with Felt’s type series. Samples of syconia in various stages of development were dissected to determine the presence and abundance of midge galls. Immature stages were macerated in warm 80% lactic acid. First instar larvae were mounted in polyvinyl-lactophenol. Larvae of later in- stars and pupae were mounted in Euparal. Fruits of identified fig species were isolated in vials to rear adult gall midges. Adults were stored in 70% alcohol and eventually mounted in Euparal using the method out- lined in Roskam (1977). Adult terminology follows McAlpine et al. (1981); for larval terminology, see Méhn (1955), and for pu- pal terminology M6éhn (1961). All mea- surements were taken from slide-mounted material. The statistical methods are ex- plained in the paragraph dealing with the comparison of new material with Felt’s type series. Behavioral observations were made in several locations in Dade and Monroe Counties, Florida. Sex ratio and periodicity of midge emergence from galls were studied 779 in excised fruits which were isolated in plas- tic vials with mesh caps and ambient con- ditions. Ficiomyia perarticulata Felt Ficiomyia perarticulata Felt 1922: 5. Ficiomyia birdi Felt 1934: 132. New Syn- onym. For quantitative characters, see Tables | and 2. Male.— Head: Eyes very large, holoptic, about 11 facets long at vertex; facets hex- agonal, closely abutting one another (Fig. 1). Occiput diamond-shaped, largely cov- ered with setae (Fig. 2). Antenna with 29- 31 stalked flagellomeres, the fused first and second flagellomeres counted separately; node with a basal whorl of short, rigid setae and long, bent setae on horseshoe-shaped sockets scattered over its anterosubdistal surface, circumfila appressed, forming one complete basal whorl and a partial, anter- odistally situated whorl, the two connected at the medial and lateral surfaces of the node (Fig. 6). Antennal plate and clypeus covered with many setae; labrum sclerotized; la- bium heart-shaped; labella hemispherical in lateral view, the distal half covered with rig- id setae. Palpus 3-segmented, basal segment partly to completely fused with second, third segment variable in length; basal segment with two to six laterally-situated, long setae (Fig. 3). Thorax: Scutum with four longitudinal rows of setae interspersed with scales. Scu- tellum with scattered setae and scales. An- episternum with three groups of anterodor- sally, anteroventrally and centrally situated setae and scales. Anepimeron with a central group of setae and scales. Legs densely cov- ered with scales; claws toothed, teeth usu- ally bifid, first tarsomere of all legs with pointed asetulose lateroventral projection (Fig. 5); empodia as long as or slightly longer than claws and twice as long as pulvilli (Fig. 4). Wing densely covered with scales, hya- line if scales absent, maculate if scales pres- 780 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Quantitative comparison of newly collected material with Felt’s type series, males. All measurements are in um, bold-printed values indicate ranges for Felt’s series which exceed the ranges of the newly collected material. *, characters used for PCA. perarticulata new birdi Felt perart. Felt Character N Mean cv % Range N Range N Range *he head 11 584 1.7 568-598 3 568-637 1 617 he antennal plate 11 198 3.6 184-208 3 162-184 l 184 nr flagellomeres 8 30 2.8 29-31 1 30 0 = le antenna 8 2879 5.0 2684-3063 l 2716 0 — *le 10 flagellomeres 11 1011 3.3 947-1074 2 917-1074 2 995-1074 le node 5. flagm. 11 68 7.0 63-77 2 68-70 2 63-75 wi node 5. flagm. 11 60 3.3 56-63 2 58-61 2 56-61 le stalk 5. flagm. 11 38 933 31441 2 39-39 Z 41-41 le seta node 5. fm. 8 255 10.0 223-300 3 290-339 2 227-307 le 3. palp segment 8 39 16.3 29-46 0 0 =- *le metafemur 11 1473 3.0 1421-1547 1 1532 2 1437-1468 le metatibia 11 1438 537) 1295-1547 1 1437 2 1421-1500 le 2. metatarsus 11 1172 32 1105-1232 1 1232 2 1137-1263 le 5. metatarsus 11 229 4.8 205-237 0 = 1 221 wi wing 11 1061 3.6 979-1103 3 1074-1232 1 884 le vein R1 11 939 4.4 884-995 3 1026-1184 1 1105 *le vein R5 11 2367 2.6 2290-2495 3 2605-2842 1 2416 le basal branch Cu 11 687 4.9 647-742 3 647-774 0 - le proximal br. Cu 11 1451 3.4 1374-1547 3 1515-1768 0 = le distal br. Cu 11 812 4.8 758-900 3 979-1011 0 - le gonocoxite 11 242 3.5 227-254 2 247-252 2 237-249 le lobe gonocoxite 11 128 Ue 104-138 2 116-121 0 - wi gonocoxite 11 81 10.3 68-92 2 68-73 0 — *le gonostylus 11 111 1.8 109-114 2 99-121 1 116 incisision cercus 11 96 12.9 80-121 2 97-102 a) 92-94 incis hypopr 11 90 121 70-106 2 77-80 D 85-90 ent (fresh material) due to patches of broad, pigmented scales among patches of narrow, hyaline ones; R5 almost straight, declining at the very end towards wing tip and ter- minating slightly anterior to it, Cu forked, its branches straight (Fig. 8). Abdomen: oblong cylindrical, yellowish and densely covered with broad, dark brown scales. Tergites 2-6 rectangular, with sim- ple, uninterrupted, posterior rows of setae, lateral setae lacking, two trichoid sensilla on anterior margins, sclerotized parts covered with scales, tergite 4 about 2.5 times as wide as long, tergite 7 narrower, with double row of posterior setae, tergite 8 not sclerotized, indicated by some posterolaterally situated setae, trichoid sensilla present; pleura thick- ly covered with scales; sternites 2-7 rect- angular, wider than long, setae in double rows On posterior margins, scattered along lateral margins and on anterolateral parts, a pair of closely placed trichoid sensilla on medioanterior margin, sclerotized parts with scattered scales, sclerite 8 narrower and without trichoid sensilla. Cerci rounded posteriorly, with rigid setae on posterior parts of both surfaces, arrangement of mi- crotrichia in transverse, elongate patches anterodorsally, gradually changing to a scat- tered arrangement posteriorly, ventral sur- face with microtrichia in transverse rows; hypoproct variable, oblong, parallel-sided, and deeply emarginate, with microtrichia scattered on its dorsal surface and in trans- verse rows on the ventral surface, each lobe with two to three apical setae; gonocoxite oblong, narrowed at mid length, with a con- spicuous apicoventral lobe, setae mainly on VOLUME 92, NUMBER 4 Table 2. explanation, see Table 1. 781 Quantitative comparison of newly collected material with Felt’s type series, females. For further perarticulata new birdi Felt perart. Felt Character N Mean cv % Range N Range N Range *he head 11 584 3.0 559-608 2 549-568 4 568-608 he antennal plate 10 193 6.5 184-201 1 169 3 186-203 nr flagellomeres 9 38 2.2 37-39 (0) _ 2 39-39 le antenna 9 2482 5.8 2290-2700 0 - 1 2921 *le 10 flagellomeres 11 723 4.1 679-774 1 790 1 821 le node 5. flagm. 11 56 Sui) 51-63 1 58 1 53 wi node 5. flagm. 11 57 4.8 53-61 1 58 1 63 le stalk 5. flagm. 11 23 10.9 19-24 1 27 1 24 le seta node 5. fm. 11 106 2122 82-145 1 148 1 85 le 3. palp segment 10 41 27.0 24-58 0 1 34 *le metafemur 11 1503 5.4 1405-1642 2 1547-1579 3 1579-1705 le metatibia 11 1457 6.9 1342-1611 2 1547-1705 3 1611-1721 le 2. metatarsus 8 SS 8.4 963-1263 1 1421 0 = le 5. metatarsus 7 229 5.0 221-237 0 — 0) = wi wing 11 1192 3.8 1137-1263 2 1326-1405 2 1168-1326 le vein R1 11 1041 4.1 995-1105 2 1137-1231 2 1137-1200 *le vein RS 11 2626 Shi) 2526-2779 2 3047-3363 2 2684-3174 le basal branch Cu 11 748 5.3 695-821 1 916 2 679-868 le proximal br. Cu 11 1652 4.9 1547-1753 2 2053-2116 2 1721-2100 le distal br. Cu 11 895 4.3 837-947 2 1105-1105 2 995-1058 le cercus 10 177 4.7 165-189 (0) — 2 143-155 *he cercus 11 49 6.3 44-53 1 51 3 44-48 le hypoproct 11 28 1g 22-34 1 29 3 29-39 distal part of dorsal surface and scattered over lateral and ventral surfaces; gonostylus cylindrical, blunt at apex, with curved, stout, apical tooth and with numerous setae, mainly on its lateral surfaces; mediobasal lobe about half as long as gonocercus, stout, blunt, with transverse, narrowly oblong patches of microtrichia; aedeagus slender, parallel sided, and distinctly longer than the mediobasal lobes (Fig. 10). Female (characters not mentioned similar to those of the male). — Head: Antenna with 37-39 stalked flagellomeres; node with only basal whorl of short setae and complete, laterally and medially interconnected cir- cumfila (Fig. 7). Abdomen: ovoid, deep orange-red; sper- mathecae conspicuously pigmented. Ovi- positor telescoped, abdominal segment 9 about 1.5 times as long as segment 8, with setae scattered all over its surface, the mi- crotrichia in groups, forming a reticulate pattern, one pair of trichoid sensilla on an- terolateral margin; segment 10 slightly long- er than segment 9, with microtrichia in dense transverse rows; cercus oblong, with setae scattered over whole surface, the longer ones mainly distal, microtrichia densely scat- tered anteriorly, in more or less reticulate pattern posteriorly; hypoproct short, ob- tuse-triangular, with microtrichia in trans- verse rows (Fig. 9). Pupa.— Color turning from creamy-white to yellowish in male and pink-orange in fe- male. Apical and lateral spines on head, as well as lateral facial papillae, which are usu- ally present in gall midge pupae, absent and indicated by blunt projections. One pair of trichoid sensilla on apical projections (Fig. 14). Thoracic horns absent, prothoracic spi- racle protruding from its respective surface. Abdomen with five pairs of protruding spi- racles, on segments 2-6, the pair of stigmata on segment 7 vestigial (Fig. 13). On seg- 782 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10 Figs. 1-10. F. perarticulata, adult structures. 1, Male head, frontal. 2, Same, distal. 3, Mouthparts. 4, Detail of male hind fifth tarsomere. 5, Detail of distal part of male hind first tarsomere. 6, Male fifth flagellomere. 7, Female, same. 8, Male wing. 9, Ovipositor. 10, Male postabdomen, dorsal. 8, x25. 1-2, x 100. 9-10, x 180. 6— 7, x 240. 4-5, x 290. VOLUME 92, NUMBER 4 Figs. 11-18. 783 ca 17 F. perarticulata, structures of immature stages. | 1, Male pupa, distal view of ultimate abdominal segments. 12, Female pupa, same. 13, Male pupa, stigma and skin structures on sixth abdominal segment, lateroventral. 14, Male pupa, head and thorax, lateral. 15, Third instar larva, antenna. 16, Same, stigma on first thoracic segment. 17, Same, head, supernumerary segment and first thoracic segment with spatula, ventral. 18, Second instar larva, detail of spatula. 14, x65. 13, x 100. 11-12, 17, x 150. 18, x 450. 16, «725. 15, x 1450. ments 2-6, three pairs of dorsal trichoid sensilla and one pair of pleural sensilla, the sublateral pair of dorsals closely situated to the intermediate pair. On segment 7, one pair of dorsals and one pair of laterals. Dor- sal and lateral surfaces with pointed setulae, ventral surface glabrous. Male pupa with two large posterior convexities in which the gonocoxites develop; female pupae with these cavities vestigial (Figs. 11-12). Third (final) instar.—Body obconic, with broad thorax and upwardly curved, gradu- ally narrowing abdomen, creamy-white; segmentation as usual for gall midge larvae and consisting of head (h), supernumerary segment (ss), three thoracic (t1—3), nine ab- 784 H ss tip 2" tSe=alava2 20) 2d 2d) 2d) 2d S SS 3c Pp PP p p ip. ip ip “pv py PA\~ 972) 975! Sos, )S Vav. av PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a7 a8 as d Ses PPS pv pv av av Fig. 19. F. perarticulata, diagram of pattern of papillae and tracheal system of third instar larva. al-8, abdominal segments. as, anal segment. av, anterior ventral papilla. c, collar papilla. d, dorsal papilla. h, head. ip, interior pleural papilla. |, lateral papilla. p, pleural papilla. py, posterior ventral papilla. S, stigma. s, sternal papilla. ss, supernumerary segment. t, terminal papilla. tl—3, thoracic segments. dominal (al—9), of which the final one is the anal segment (as). Body length 1850-2520 um, width 850-1410 um, head capsule width 123-157 um, width between antennae 53- 70 wm. Head capsule flat, weakly sclero- tized, antennae truncate with sclerotized, plumose appendages (Fig. 15); thoracic sur- face glabrous, anteroventral surface of a2— a7 with about 25 transverse spinule-rows, a8 with such rows on its posteroventral sur- face. Spatula transverse, bilobed, the lobes acute; stalk not differentiated. Spatula height 94-126 wm, width 236-314 um, height of lobes 34-46 um, distance between tips 48- 85 um (Fig. 17). Respiration peripneustic, with spiracles on tl and al—a8; spiracles with oblong sclerotized outgrowths (Fig. 16). Papillae, unless indicated otherwise, with- Out setae; pattern of papillae; three pairs of lateral collar papillae, the papillae of two of these pairs abutting (Fig. 17); thoracic seg- ments with one pair of sternal papillae each, the pair on tl abutting and confined by the spatula lobes (Fig. 17), two pairs of abutting laterals with short setae, one pair of interior pleurals, two pairs of dorsals; segments al— a7 with one pair of anterior ventral papillae, one pair of posterior ventrals, one pair of laterals and two pairs of dorsals; a8 with dorsal papillae missing; anal segment with usually three pairs of terminal papillae, anal papillae absent (Fig. 19). N = 10. First instar. — Body ellipsoid, transparent. Body length 588-735 um, width 225-254 um. Head capsule width 33-41 um, flat- tened, with truncate antennae. Body surface glabrous with patches of minute spinule- rows on dorsal and ventral surfaces. Spatula absent. Respiration apneustic with a pair of vestigial spiracles on abdominal segment 8. All papillae without setae except the pleural pair on abdominal segment 8. Pattern of papillae basically as in third instar; the pair of pleural papillae on abdominal segment 8 very distinct and with short setae. N = 4. Second instar.— Body ellipsoid, creamy- white. Body length and width not defined because of the poor condition of the ma- terial. Head capsule width 87 um, head cap- sule convex, with truncate antennae. Body surfaces as in first instar. Spatula present, bilobed, stalk not differentiated, width 17 um, between lobes 7 um (Fig. 18). Respi- ration peripneustic, with spiracles situated as in third instar, width of first thoracic spi- racle 10 um. Pattern of papillae as in third instar. N = 1. Gall. — Barnes (1948) described the gall as an enlarged seed capsule which becomes ab- normally lengthened. However, because the young gall may bear one to three flowers on its exterior, the gall must be considered a pocket-shaped outgrowth of the receptacle (Fig. 20). The associated flowers are grad- ually reduced as the gall matures. One larva is present per gall. The first instar is oriented VOLUME 92, NUMBER 4 with its head towards the syconial cavity and is completely enclosed by the gall tissue; the head is traceable by a pair of dark eye- spots (Fig. 21). A gall chamber becomes dis- tinct when the larval spatula develops. At the late pupal stage a conspicuous ‘crown,’ with a central window-pit, grows from the top of the gall on the fruit surface. Often, two or three galls are concentrated around one window-pit. One to several crowns oc- cur per galled fruit. Two shapes are distinguished in mature galls (Fig. 22): shorter, sessile galls with the gall chamber situated close to the window- pit, and longer, stalked galls with a distinct constriction between gall chamber and win- dow-pit. Males developed exclusively in the sessile galls, and females in the stalked galls. This was determined by dissection of pupae from galls (N = 15 per gall type), and dis- section of galls after emergence of adults (N = 30 per gall type). Such a dimorphism has also been reported by Coutin and Riom (1967) for Mikiola faqi Hartig on beech in Europe. Types. —Felt did not designate holotypes from among his type series. Lectotypes are therefore designated here by one of us (JCR). For F. perarticulata slide ‘‘a,” male, with left hand label ““Lectotype, design. by J. C. Roskam 1989”; for F. birdi the slide marked with “type” by Felt, male, remounted by R. J. Gagné, again with left hand label “Lec- totype, design. by J. C. Roskam 1989.” All specimens of both Felt series belong to the New York State Museum at Albany and are now on indefinite loan to the Systematic Entomology Laboratory in Washington, D.C. The specimens we mounted have been deposited in U.S. National Museum in Washington. COMPARISON OF NEw MATERIAL WITH FELT’s TYPE SERIES The type series of F. perarticulata consists of 10 slides with specimens or fragments mounted in Canada balsam. Felt (1922) de- scribed the specimens as ““somewhat broken 785 in transit and as a consequence, the descrip- tions ... are not complete in certain de- tails.” All slides are labeled ‘“‘Ficiomyia per- articulata Felt on Ficus aurea Miami, Fla. Feb. 9 °22 Type a32128.”’ On two of these slides, labeled by one of us (JCR) with “male a” and ‘“‘male b,” a male is mounted, both with incomplete antennae and shriveled heads and legs. Five slides, labeled with “female a’’—‘‘female e,”’ bear incomplete fe- males. All this material was cleared before mounting. A slide labeled ‘“‘f’ contains two wings of different sizes, apparently taken from specimens before clearing; a slide la- beled “‘g” contains a complete female fla- gellum in shriveled condition. Finally, a slide marked ‘th” contains a mixture of incom- plete male and female flagella, legs, and the thorax and gaster of a female fig wasp, Pe- goscapus sp. (species identification not pos- sible without the head). The type series of F. birdi Felt consists of three slides with a male on each and two slides with a female on each. One male and one female are marked “‘type”; all slides are labeled “Fi- ciomyia birdi Felt Florida 1933.” The male that we designate as lectotype has been cleared and remounted by R. J. Gagné, Washington, D.C. (R. J. Gagné, in litt.). Felt’s (1934) statement about the number of antennal segments is based on material of the male marked ‘“‘b” by JCR. All ma- terial, specimen “‘b”’ excepted, has incom- plete antennae and legs and shriveled palpi. The lobed gonocoxites and the high num- ber of stalked antennal segments in both sexes, characters on which the genus Ficio- myia was erected, are distinct in the ma- terial of both type series. The statements “palpi probably uniarticulate” (Felt 1922) or “palp consisting of one, slender, rather long segment” (Felt 1934), which are used to distinguish Ficiomyia (Felt 1925, Gagné in McAlpine et al. 1981) are not correct. These must be replaced by “palpi usually 3-segmented, with the first and second seg- ments partly to completely fused.” Examination ofnew material revealed that 786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 20-22. Longitudinal sections of infested F. citrifolia syconia. 20, Gall with reduced flowers on its exterior (ga). 21, Gall with first instar gall midge larva. 22, Syconium with two stalked (female) and one sessile (male) gall. 22, x8. 20-21, x12. VOLUME 92, NUMBER 4 the species differences presented by Felt (1934) for F. perarticulata and F. birdi are not valid. The difference in body color is apparently due to different clearing of the material. The fuscous markings on the wings reported by Felt (1934) of F. birdi (caused by a thick layer of scales) are absent in the material of F. perarticulata, but this is prob- ably an artifact: the scales are easily dis- lodged in alcohol-stored material. The most striking difference reported by Felt, how- ever, concerns the number of antennal seg- ments. He apparently used a complete fe- male antenna for his description of F. perarticulata, whereas he used the antenna of a male in his description of F. birdi. This difference in the number of antennal seg- ments, though, is not due to a species dif- ference but to sexual dimorphism. This is the first record of considerably fewer flag- ellomeres in males than females in Ceci- domyiidae. Male cecidomyiids generally have as many or a few more flagellomeres than females. Ficiomyia is, therefore, re- markable among gall midges (for a review, see Mamaev 1968). Finally, we doubt the identification of the host plant of F. perar- ticulata. For a discussion, see the section on host plant associations. Because we found no qualitative char- acters to differentiate between the two species, we sought differences by using quantitative analysis. The quantitative traits are listed in Tables 1-2. Although both of Felt’s series comprise considerably less ma- terial than the material we reared, his ma- terial is more variable. His midges are gen- erally larger, but in eight cases from both series some characters exceeded both the lower and upper boundaries of the ranges set by our material. Therefore, although our material should be conspecific with one of Felt’s series, the difference between the Felt material and our midges is larger than the difference between Felt’s series. Based on this analysis, we again find no evidence to support the view that F. perarticulata and F. birdi are different species. 787 With principal component analysis (PCA) it is possible to analyse several characters simultaneously; the technique therefore al- lows a more accurate judgment on the status of F. perarticulata and F. birdi. For a de- scription of the technique and an outline of its possibilities, see Pimentel (1979). Among its attributes, PCA results in a graphic rep- resentation of specimens in a coordinate system with axes that show zero intercor- relations and to which the original charac- ters (by measured values of the data matrix) have contributed proportionally to their variation. Put otherwise, the original axes, representing the characters, are rotated while the original relationships among the data points, representing the specimens, are maintained. If F. perarticulata and F. birdi are differ- ent species, they should take different po- sitions in a PCA hyperspace. The members of one of such supposed species should oc- cupy closer mutual positions than members of a different species. The data matrix con- sists of values measured for the specimens we reared. Because the technique does not allow missing values, and missing values are frequent in Felt’s type series, we represented the type series by mean values (Tables 1, 2). We are aware of the flaws of this decision but, given the poor condition of the mate- rial, it was our best option. Therefore, apart from the matrix of 11 specimens per sex we include two ‘type representatives,’ one for F. perarticulata and one for F. birdi, again per sex. The data matrix is standardized (means zero, standard deviations 1) because the characters used are of a different size order (Tables 1, 2). The values for Felt’s material have been standardized using the mean val- ues and standard deviations from the char- acters as listed in Tables | and 2. All char- acters were tested for normality. A PCA technique constrains the number of char- acters which can be used relative to the number of specimens, and, in our case ne- cessitates a low ‘within species variation.’ 788 We therefore chose a limited number (5) of characters showing a low coefficient of vari- ation (Tables 1, 2). We selected these char- acters from different body regions to avoid combinations of characters with high inter- correlations (e.g. different parts of one leg). The PCA was done for males and females separately. Results. — Males (Fig. 23). The type series of F. perarticulata (P) and F. birdi (B) are excentrally situated: both represent large midges. The F. perarticulata representation (P) is nearest to the centroid, even closer than specimen marked 1. The latter speci- men, like all others (a—-k), supposedly be- longs to F. birdi on the basis of host plant association. The F. birdi representation (B) is situated farthest from the centroid, but, on the other hand, is so closely situated to specimen i that we cannot assign it to a separate species. For females (Fig. 23) the result is even clearer: here both F. birdi and F. perarticulata representations are excen- tric, but mutually very close. Therefore we must conclude again that a// material, i.e., Felt’s type series and our material, is con- specific, and F. birdi must be put into syn- onymy for reason of priority. DISTRIBUTION AND Host PLANT ASSOCIATION Ficus aurea, the Florida strangler fig, and F. citrifolia, the shortleaf fig, are native to Florida. The Florida strangler fig is more abundant and occurs also on the Bahama Islands. The shortleaf fig is restricted in Florida to hammocks in the southernmost part and the Florida Keys, and occurs also in the Bahamas and the West Indies (Elias 1987). Although we examined many sam- ples of F. aurea, we never found Ficiomyia or its parasitoid, Physothorax bidentulus Burks, in this fig species (there is one record of P. bidentulus from F. aurea, in Burks (1969), but it is from the same collection from which F. perarticulata was described). D. McKey, botanist at the University of Florida at Miami has also never found the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON x 0.75 MALES Na = S o a : sere F 0.50 Ya 8 4 Ep TES VA et ° e / re & 0.25 / 0.00 +3 le metatemur h -0.25} is ] -0.50 | Ts, Can -0.75}¢4 = °o "3 -1.00 . s ‘ a , -050 -025 0.00 025 0.50 0.75 1.00 1.25 —— PC axis! : 42.2% n 08 FEMALES Ab Re é e #) es é mn O06 2 2 / 3 / o. 0.4} oP ie / bh 9 0.2 —. le vein RS oo} P = 5 he cercus f ’ b xX — -0.2 . SS ys \ x heag ~ -o.4t \S, 2% a” =O sf —1.2 -0.8 -0.4 0.0 O4 0.8 ue 16 — PC axis I: 57.4% Fig. 23. Principal component analysis of adult characters. The vectors indicate amount and direction in which the separate characters attribute to the posi- tion of the specimens. a—k, Newly reared specimens. B, Representation of F. birdi. P, Same, F. perarcticu- /ata. For further explanation, see text. conspicuous Ficiomyia galls in syconia of F. aurea (pers. comm.). The record of Fi- ciomyia on F. aurea, therefore, was prob- ably due to a misidentification of the host plant. The host plant of F. perarticulata has also been erroneously listed as F. carica in Gagné (1989). We examined the following samples: Fi- cus aurea, Dade Co., Miami, 3-22-89, leg. H. Nadel and J. C. Roskam, 147 syconia, VOLUME 92, NUMBER 4 diameter 5-8 mm, two kinds of ovaries, small and large, containing different species of fig wasps, the species in the larger ovaries containing Physothorax; Dade Co., Miami, 3-22-89, leg. H. Nadel and J. C. Roskam, 50 syconia, diameter 5-8 mm, two sizes of fig wasp-containing ovaries, the larger con- taining Physothorax; Dade Co., Miami, 3-22-89, leg. H. Nadel and J. C. Roskam, 50 syconia, diameter 5—6 mm, ovipositing agaonids, no size differences among ovaries; Lee Co., Sanibel Island, 3-25-89, leg. H. Nadel and J. C. Roskam, 100 syconia, di- ameter 5-9 mm, small and large ovaries, one Physothorax russelli Crawford. Monroe Co., No Name Key, 9-17-88, leg. H. Nadel, 15 syconia, diameter 7-8 mm, some with many fig wasps, no midge galls. Monroe Co., Key Largo, 9-12-88, leg. H. Nadel, 46 sy- conia, diameter 5-8 mm, some with rem- nants of founding fig wasp females, others with almost emerged fig wasps, no midge galls. Ficus citrifolia, Dade Co., Miami, 6-24- 88, leg. H. Nadel and M. Matthews, 49 sy- conia, diameter 7-10 mm, with fig wasp remnants; 31 syconia with 80 galls, second and third instar midge larvae, midge pupae and emerged galls, third instar parasitoid larvae and parasitoid pupae; Dade Co., Homestead, 3-13-89, leg. C. Campbell, 200 syconia, diameter 7-11 mm, 20 of these sy- conia with one to five Ficiomyia galls each. Midge larvae parasitized by Physothorax bi- dentulus Burks, pupae by an unidentified chalcidoid. Furthermore we reared species of Pegoscapus (Agaonidae), Colyostichus, Idarnes (Torymidae), and an eurytomid, probably a Syceurytoma. The latter form was also dissected from galls. Heavy mor- tality of wasps was caused by larvae of a staphylinid beetle and a few ant species; Dade Co., Miami, 3-15-89, leg. H. Nadel and J. C. Roskam, 200 syconia, diameter 5-8 mm, 30 galls, induced by Ficiomyia, larvae in second and third instar, parasit- oids absent; Monroe Co., No Name Key, 8-7-88, leg. H. Nadel, 30 syconia, diameter 789 45+4+ + +—+—_+_+_+_+_+_+_+_+_+_+_+_+—++++ ++ tr 40+ + Co moles | 357 He Ea females | . 30+ + o 3 2st | E 20+ f 2 © 454 10+ 5+ 0 np | 12 14 #16 16 20 22 00 O02 04 06 O08 10 12 time of day Fig. 24. Distribution of emergence of adult F. per- articulata from syconia of Ficus citrifolia in southern Florida. Data were combined from four days of hourly observation during April and May 1989. 8 mm, with remnants of fig wasps and agaonid larvae, no galls; Monroe Co., No Name Key, 9-17-88, leg. H. Nadel, 17 sy- conia, diameter 8-10 mm, with fig wasp remnants, many galls with midge pupae and emerged adults; Monroe Co., Key Largo, 9-12-88, leg. H. Nadel, 22 syconia, diameter 4-5 mm, too young for ovipositing fig wasps; five syconia, each with one gall, one syco- nium with three galls, all with first instar midge larvae; Monroe Co., Key Largo, 3-28- 89, leg. H. Nadel and J. C. Roskam, 30 syconia, galls absent, two kinds of ovaries containing agaonid and torymid wasps. ADULT EMERGENCE FROM GALLS The formation of the crown-like ridge around the opening of the gall begins about two days before eclosion of F. perarticulata. The fig skin splits, exposing the whitish tis- sue of the syconial wall, and curls outward radially to form a crown around the exit area. The tissue within the base of the gall becomes moist and vicid. The pupa pushes its way head first to the gall exterior and comes to rest with about three-quarters of its length protruding from the fig surface. The adult ecloses after a few minutes, leav- ing the pupal skin partly embedded in the crown, and rests hanging from the side or bottom of the fig by its front and middle legs. The wings are fully expanded within 790 five minutes and are capable of sustained flight within an hour after eclosion. Daily emergence within a population is periodic, with a minor peak in late after- noon and a major peak during the night, and with males tending to emerge earlier than females during the major peak (Fig. 24). Within a single fig, emergence of all adults usually spans a few days, and may span even weeks, as suggested by our ob- servations of second instar larvae in figs with already emerged galls. Emergence occurs throughout the year in Florida, as F. citrifolia trees fruit asynchron- ously in relation to each other and thus af- ford year-round development by the gall midges. This scenario probably prevails over the entire range of the midge. MATING Mating probably occurs on or around the tree from which adults emerged. Females extrude their ovipositors directly after eclo- sion, apparently to emit pheromones. On three evenings around sunset we observed males, solitary and in small groups, flying in zigzag motion towards and around tips of branches. Males also frequently landed on fruits and leaves. We observed only one mating pair, which was on a fig at 17:45 on 6 December. The female, however, was cap- tured by an ant either before or during cop- ulation. The male continued to hang, head- down, by its terminalia for five minutes before disengaging from the female. SEX RATIO Both sexes may inhabit one fig. The sex- ual dimorphism in gall shape allowed us to determine midge sex ratio before mortality due to parasitism. In one sample of 13 fruits with 134 galls, 69 galls were male (sessile) and 65 were female (stalked), which is es- sentially a 1:1 ratio; however, in a sample of 50 fruits from a different location, 61 were male and 24 were female, which is highly male-biased (Chi-square = 16.11; P < .005). The sex ratio of emerged adults PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON was studied in four other locations. Collec- tion | yielded 32 adult males and 35 fe- males; collection 2 yielded 32 males and 20 females; collection 3 yielded 70 males and 46 females; and collection 4 yielded 88 males and 81 females. The sex ratio in collection 3 deviated significantly from 1:1 (Chi-square = 4.97; .025 < P < .050). Apparently, sex ratio of F. perarticulata varies from equality to male-biased. SYCONIAL GALL MIDGES IN OTHER GEOGRAPHIC AREAS If close relatives of Ficiomyia exist, we would expect them in syconia of figs occur- ring elsewhere. Docters van Leeuwen- Reijvaan and Docters van Leeuwen (1926) listed 28 kinds of galls from Malaysian figs on leaves, stems, aerial roots, and three kinds in syconia. Samples of these fruit galls, dried and in alcohol, have been deposited in the Rijksherbarium at Leiden, the Netherlands. None of the syconia of this material bear galls of the type induced by Ficiomyia. We dissected one mature larva from a syconi- um. This larva differs in many respects from Ficiomyia larva and probably belongs to the tribe Cecidomyiini. Barnes, in Williams (1928), described two species, one cecido- myiine and the other an asphondyliine, from fruits of figs native to the Philippines. These species, together with a cecidomyiine oc- curring in India, have also been listed in Barnes (1948). Although seven species of figs occur in Japan (Ohwi 1965), no ceci- domyiids have been reported from them (Yukawa 1971). Finally, Mani (1973) dis- tinguished eight kinds of leaf galls, but none is induced by Oligotrophini. Two females, undoubtedly belonging to Ficiomyia, have been collected in a light trap in Dominica (West Indies, Clarke Hall, 1-10 February and 1-10 March 1965, leg. W. W. Wirth). The sole complete flagellum yields a count of only 33 flagellomeres and the hypoproct is slightly longer (41 um) in one specimen. All remaining measurements are within the ranges given for F. perarticu- VOLUME 92, NUMBER 4 lata. Because of the low number of flagello- meres this material might belong to a new species. More material and host data are required, though, to draw a conclusion and to provide a formal description. Possible Ficiomyia parasitoids, in the ge- nus Physothorax, have been described from Brazil, reared from big, stalked galls in the syconia of F. doliaria Mart. (Mayr 1885, 1906, Miller 1886). We cannot conclude, however, whether this indicates the pres- ence of Ficiomyia in Brazil, since P. russelli Crawford and P. pallidus Ashmead also emerge from large, stalked galls in F. aurea. These galls are not induced by Ficiomyia, but probably by torymid sp. The genus Phy- sothorax is not known outside the New World (Bouéek et al. 1981). Hence, al- though our data are incomplete, there is no evidence that Ficilomyia or its parasitoid, Physothorax, occurs outside the New World. Syceurytoma has been described from Af- rican material (Bouéek et al. 1981), but again there is no evidence that any of its species are associated with Ficiomyia. ACKNOWLEDGMENTS We thank M. Zandee, Institute of Theo- retical Biology, University of Leiden for his statistical help. E. Grissell, Systematic Ento- mology Laboratory checked the identifica- tions of the parasitoids. H. Heyn is grate- fully acknowledged for drawing some of the figures. For his critical and helpful reviews of this manuscript and for the loan of spec- imens in his care we thank R. J. Gagné, Systematic Entomology Laboratory, Wash- ington, D.C. LITERATURE CITED Barnes, H. F. 1948. Gall Midges of Economic Im- portance. III. Fruit. Crosby Lockwood & Son, London. 184 pp. Boucek, Z., A. Watsham, and J.T. Wiebes. 1981. The fig wasp fauna of the receptacles of Ficus thonnin- 791 gil (Hymenoptera, Chalcidoidea). Tijdschr. Ent. 124: 149-233. Burks, B. D. 1969. Redefinitions of two genera of chalcidoids from figs, with new Florida species (Hymenoptera). Flonda Ent. 52: 115-122. Condit, I. J. 1969. Ficus: The Exotic Species. Univ. Calif. Div. Agri. Sci., Berkeley. 363 pp. Coutin, R. and J. Riom. 1967. Dimorphisme des galles provoquées par Mikiola faqi Hartig. (Dipt. Cécid.) sur Fagus silvatica L. Androcécidie et gyn- océcidie. C. R. Acad. Sc. Paris 265 Ser. D: 975— 978. Docters van Leeuwen-Reijnvaan, J. and W. M. Docters van Leeuwen. 1926. The Zoocecidia of the Neth- erlands East Indies. De Unie, Batavia. 601 pp. Elias, T. S. 1987. The Complete Trees of North America. Gramercy, New York. 948 pp. Felt, E. P. 1922. A new and remarkable fig midge. Florida Ent. 6: 5-6. ——.. 1925. Key to gall midges (A resumé of studies 1-7, Itonididae). Bull. New York St. Mus. 257: 135-186. ——. 1934. Anew gall midge on fig. Ent. News 45: 131-133. Gagne, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ith- aca and London. 356 pp. Mamaey, B. M. 1968. Evolution of the Gall Forming Insects— Gall Midges. British Library Lending Di- vision, Boston Spa, Wetherby, translated by A. Crozy (1975). 317 pp. Mani, M. S. 1973. Plant Galls of India. Macmillan, Madras. 354 pp. Mayr, G., 1885. Feigeninsecten. Verh. zool.-bot. Ges. Wien 35: 147-251. Mayr, G. 1906. Neue Feigen-Insekten. Wien. Ent. Zeit. 25: 153-187. McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood. 1981. Manual of Nearctic Diptera. Vol. I. Research Branch Agriculture Canada, Monograph No. 27. 674 pp. Mohn, E. 1955. Beitrage zur Systematik der Larven der Itonididae (= Cecidomyiidae, Diptera). Zoo- logica, Stuttgart 105: 1-247. 1961. Gallmiicken aus El Salvador (Diptera, Itonididae). 4. Zur Phylogenie der Asphondyliidi der neotropischen und holarktischen Region. Senckenb. Biol. 42: 131-330. Miiller, F. 1886. Zur Kenntnis der Feigenwespen. Ent. Nachr. 12: 193-199. Ohwi, J. 1965. Flora of Japan. Smithsonian Inst., Washington, D.C. (English ed.). 1067 pp. Pimentel, R. A. 1979. Morphometrics. The Multi- variate Analysis of Biological Data. Kendall-Hunt, Dubuque, Iowa. 276 pp. Roskam, J.C. 1977. Biosystematics of insects living 792 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in female birch catkins. I. Gall midges of the genus species). Bull. Exp. Sta. Hawaiian Sugar PI. Ass. Semudobia Kieffer (Diptera, Cecidomyiidae). No. 19. Honolulu, Hawaii. Tijdschr. Ent. 120: 153-197. Yukawa, J. 1971. A revision of the Japanese gall Williams, F. X. 1928. Studies in tropical wasps— midges (Diptera: Cecidomyiidae). Mem. Fac. Agr. Their hosts and associates (with description of new Kagoshima Univ. 8: 1-203. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 793-801 TWO NEW SPECIES OF METAJAPYX (DIPLURA: JAPYGIDAE) FROM TENNESSEE MarK A. MUEGGE AND ERNEST C. BERNARD (MAM) Department of Entomology and Plant Pathology, University of Tennessee, Knoxville, Tennessee 37901-1071. Current address: Department of Entomology, Loui- siana State University, 402 Life Sciences Bldg., Baton Rouge, Louisiana 70803; (ECB) Department of Entomology and Plant Pathology, University of Tennessee, Knoxville, Tennessee 37901-1071. Abstract.—Two new species of Japygidae from eastern Tennessee are described and illustrated. Metajapyx heterocercus n. sp. exhibits sexual dimorphism of the cerci and chaetotaxy of the tenth abdominal tergum, and Metajapyx magnifimbriatus n. sp. exhibits only 30 antennal segments. Both sexual dimorphism and the reduced number of antennal segments are new characters of the genus in the United States. Key Words: Daiplura, Japygidae, Metajapyx, taxonomy The genus Metajapyx Silvestri is repre- sented in North America by eight eastern species (Smith and Bolton 1964). Recently, Rathman et al. (1988) illustrated a japygid from eastern Washington State as a Meta- Japyx sp.; some generic criteria for the genus Metajapyx were not reported for those spec- imens and therefore, further study is re- quired to determine their generic place- ment. Although two species of Metajapyx are quite widespread and frequently encoun- tered (M. subterraneus [Packard] and M. steevesi Smith and Bolton), the other six species are locally distributed and rarely en- countered. During intensive collection of Metajapyx spp. in the eastern United States for the purpose of revisionary studies we found two new Metajapyx species from eastern Tennessee. The objective of this pa- per is to describe and illustrate these new species. MATERIALS AND METHODS All specimens were cleared and stained in Essig’s Aphid Fluid (Wilky 1962) and double stain (1 part 5% aqueous lignin pink and 5% aqueous acid fuchsin to 10 parts of Essig’s Aphid Fluid), then mounted in poly- vinyl alcohol-lactophenol to further clear and expand the specimens, and dried in an oven (60°C) for two days. Measurements and illustrations were made with an ocular micrometer and a Nikon phase-contrast mi- croscope equipped with a drawing tube or a Unitron stereo microscope. Body length measurements were made from the basal articulation of the antennae to the apex of the cerci. Measurements are presented as a mean followed in parentheses by the range. Because the distinctions among certain taxonomic characters used to describe Ja- pygidae are sometimes vague it may be helpful to define the more salient of these. The following definitions are those of Smith (1962): M = larger macrosetae of the body set in reinforced setal sockets, so as to move in a plane parallel to the body; m = medi- um-sized sub-macrosetae usually set in sim- ple setal sockets; Microsetae = minute setae visible only under high magnification, al- ways set in simple setal sockets; Friction 794 setae = a type of microseta with large sock- ets that occur in groups where the body in- tegument folds or moves upon itself; Calcar setae = two setae at the ventral apex of a tibia which may be thicker or more robust, but no longer than other tibial setae; Apot- ome = the anterior sclerite of an abdominal sternum. For a more complete treatise of japygid terminology one should refer to Smith (1962), Pagés (1952), and Steinmann and Zombori (1984). Another potentially important taxonom- ic character of japygids is cercal chaetotaxy. This character has been either ignored or used only sparingly in japygid taxonomy, yet it appears to be useful at the generic and specific levels. In this paper we propose a new scheme to describe the cercal chaeto- taxy of the genus Metajapyx. The cercal se- tae are arranged into four general categories: the lateral setae (L), the dorsal setae (D), the ventral setae (V), and the postdental margin setae (P). Within each group, setae are num- bered from proximal to apical, and given relative size descriptions (long, short or minute). In this paper cercal chaetotaxy is used to help distinguish closely related spe- cies. Metajapyx heterocercus Muegge and Bernard New SPECIES (Figs. la—f, 2a—d; Table 1) Females.—Body length 12.1 (9.3-14.5) mm. Entire body and head yellowish-white, except for median sclerotized area at apex of epicranial and postoccipital sutures, and tergites VI through X, which are progres- sively more sclerotized posteriorly. Cerci heavily sclerotized. Antennae: Thirty-two segmented and heavily setose, setae arranged in two irreg- ular whorls, the basal whorl alternating as simple and hooked setae on segments 7 to 31 (Fig. la); terminal segment with 6 plac- oid sensilla arranged with 2 in basal whorl, 4 in apical whorl; lateral proliferation of setae occurring on segments 15 to 19; tri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON chobothria present on segments 4-6 in a 3,5,5 pattern and nearly as long as longest seta on segment. Head: Numerous M, m, and microsetae; admental plate with 35+35 (29-42) m + microsetae; prementum with 27+27 (30- 43) m + microsetae, most restricted to an- terior half of sclerite; postmentum 1+1M, 1+I1m, and 19+19 (17-20) scattered mi- crosetae; submentum 1+1M, 4+4m, and 10+10 (8-12) scattered microsetae. Labial palp conical with one or two apical sensory cones and 11 +11 (10-12) setae, the longest slightly longer than palpus. Terminal seg- ment of maxillary palpus with 22 (20-27) setae, the longest as long as the segment; galea with 3 (3-4) external setae in a row, thumb of galea sclerotized, with 3 long sen- sory cones and 9 (6-11) short sensory cones; lacinia falciform and heavily sclerotized, all five laminae pectinate (Table 1), with a small basal spur between laminae III and IV. Thorax: Pronotum 5+5M, 5+5m, and scattered microsetae; prescutum | + 1M and posteromedian and posterolateral groups of friction setae; mesonotum 6+6M, 7+7m, and scattered microsetae (Fig. 1b); prescu- tum 1+1M:; posteromedian and _ lateral margin with friction setae; metanotum 5+5M, 5+5m, and scattered microsetae. Legs: Pro-, meso-, and metacoxae each with a row of friction setae near apex of segment; dorsal base of trochanter with a circular group of friction setae; dorsal apex of femur with a row of 4 long setae; ventral apex of tibia with 2 calcar setae, one much longer than the other; tarsus with two ven- tral rows of 4-6 large setae becoming more robust distally. Additional setae on all seg- ments scattered, most restricted to distal 74 of segment. Empodium minute on protar- sus and becoming progressively larger on meso- and metatarsi, metatarsal empodium subequal to pretarsus. Abdomen: Prescutum 1+1M and scat- tered friction setae; scutum 1+ 1M and scat- tered friction setae. Tergite II 3+3M, 2+2m, and scattered microsetae; anteromedian pair 795 VOLUME 92, NUMBER 4 ij m3, a L oe ec ee 7 a oOo - Py € i) . Fig. la-f. Metajapyx heterocercus. a, 14th antennal segment, L = 0.25 mm. b, Mesothorax, L = 1.0 mm. c, Tergites VI-X, L = 1.0 mm. d, Ist abdomenal sternum, L = 0.25 mm. e, Tergite VI of male, L = 1.0 mm. f, Cerci of male, L = 1.0 mm. Terms: AM, anteromedian pair of setae; M, macroseta; AS, antecedent setae; GS, glandular setae; SS, sensory setae. 796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ( =~] \ vi3 Fig. 2a-d. Chaetotaxy of female Metajapyx heterocercus cerci. a, Right dorsal view. b, Right ventral view. c, Left dorsal view. d, Left ventral view. L = 0.5 mm. Terms: D, dorsal setae; L, lateral setae; P, postdental margin stae; V, ventral setae. of large setae progressively smaller on ter- gites III-VI and absent on tergite VII. Ter- gites III-V 5+5M, 2+2m, and scattered microsetae; tergite VI with M1 usually ab- sent and posterolateral margins rounded, tergite VII with M1 always absent and pos- terolateral angles projected to the rear as blunt points (Fig. lc). Tergite VIII 2+2M VOLUME 92, NUMBER 4 Table 1. Mean number and range of laminal teeth in Metajapyx heterocercus and Metajapyx magnifim- briatus. Mean (Range) Lamina M. heterocercus M. magnifimbriatus I 13 (11-14) 10 (10-11) il 26 (24-29) 23 (21-24) Ill 16 (14-18) 10 (9-11) IV 17 (16-19) 14 (13-15) Vv 14 (13-15) 13 (12-13) and a few scattered microsetae; tergite IX 1+1M anda few scattered microsetae; plur- ae meeting in mid-ventral line, 2+2M and a few scattered microsetae. Tergite X with macrosetae A and B long, C absent, D min- ute or absent (Fig. 1c). Carinae distinct and nearly parallel; pygidium prominent and rounded. Sternite I apotome 3+ 3m alternating with microsetae; sternite I 12+12M in four ir- regular transverse rows, scattered microse- tae, and a group of two large posterolateral setae; antecedent setae 36+36 (29-39) and simple, in two irregular transverse rows, be- coming sparse mesally; lateral subcoxal or- gans protruding from sternite, with 18+ 18 (12-23) glandular setae broad basally and gradually tapering apically, nearly as long as antecedent setae; 27 +27 (21-38) hairlike sensory setae set 1n large setal sockets, about half as long as glandular setae (Fig. 1d); me- dian glandular organ protruding, with 11 (8-14) contiguous disculi. Sternites II to VII 16+16M in transverse rows, 2+2m, and scattered microsetae; sternite VIII 7+7M and scattered microsetae. Genitalia: Papillary area with groups of 11+11 (9-14) sensory pegs; anterior lobes with 4+4 (3-5) apical sensory pegs; poste- rior lobes with 4+4 (3-4) apical sensory pegs. Ventral carinae of segment X distinct with 10+10M and scattered microsetae. Cerci: Right arm predental margin uni- serrate with 4 (4—5) denticles, median tooth large and slightly rounded, postdental mar- gin crenate; dorsal (D) surface with 7 setae, 797 right lateral margin (L) with 11 setae, post- dental margin (P) with 3 minute setae, and ventral (V) surface with 13 setae: D1 long, D2 short, D3-5 long, D6 short, and D7 long, L1-5 long, L6-1 1 alternating short with long; V 1-2 short, V3-5 long, V6 short, V7—9 long, V10 short, V11-13 alternating long with short (Fig. 2a, b). Left arm predental margin biserrate with 9 (9-10) denticles in dorsal row and 9 (8-11) denticles in ventral row; median tooth large and slightly rounded, postdental margin crenate. Dorsal surface with the same number of setae as the right arm and L and P setal patterns same as right arm; D1-7 alternating long and short, V1 minute, V2 medium, V3-5 long, V6 short, V7-9 long, V10-13 alternating short and long (Fig. 2c, d). Males.—Resembling females except ter- gite V with M1 sometimes absent, tergite VI with M1 usually absent and posterolat- eral margins usually projected back into rounded points (Fig. le), tergite X with A and C absent, B long, D minute or absent. Sternite I antecedent setae more numerous; slightly fewer glandular and sensory setae. Genital papillae conical, with 2+2 (1-3) apical sensory pegs and numerous short se- tae mesally and long setae distally; genital opening with numerous short marginal se- tae becoming progressively longer distally. Cerci: Right arm predental margin uni- serrate with four denticles, median tooth large, rounded, occasionally projected pos- teriorly, postdental margin crenate, slightly falcate (Fig. If). Chaetotaxy similar to fe- male except L1 and V1 absent, and a short extra dorsal seta (D4’). Left arm similar to female except median tooth minute or ab- sent, L1 absent, L6 long, L7 short; D4 min- ute, V1 and V6-8 absent. Type material.—Holotype male and al- lotype female: Tennessee, Knox County, Cherokee Trail, 3.2 km south of the Uni- versity of Tennessee (elevation ca. 650 me- ters), 24-X-1988; M. A. Muegge. Paratypes (4 males, 4 females): Same data as holotype and allotype. Habitat.— All specimens were 798 found 5-10 cm deep in moist gravelly clay soil or under rocks in a beech-maple forest. The holotype, allotype and two paratypes (1 male, | female) are deposited in the Unit- ed States National Museum (USNM). The other paratypes (3 males, 3 females) are de- posited in the Apterygote Section of the University of Tennessee Entomology Mu- seum, Knoxville, TN. Etymology.—Greek hetero (‘different’), Greek cercus (‘‘tail’’), which referes to the sexually dimorphic dif- ferences in the cerci. Diagnosis.— Within the North American Metajapyx spp., female M. heterocercus ap- pear to be most closely related to M. stee- vesi. Females of both species are nearly identical in form and chaetotaxy; however, M. heterocercus possess only simple ante- cedent setae, while /. steevesi usually pos- sess at least a few fimbriate setae. Addi- tionally, M. heterocercus differs by the chaetotaxy of tergite X (seta C absent and B bisects tergite X at 74 its length rather than % its length as in M. steevesi), and both cerci lack dorsal setae D1’ and usually D7’. Male M. heterocercus are taxonomically similar to M. folsomi Silvestri, yet M. heterocercus males are distinguished by possessing only a few simple antecedent setae that are gen- erally uniform in size, tergite VI with the anteromedian pair of setae always present, and the median tooth of the left cercus min- ute or absent. Metajapyx magnifimbriatus Muegge and Bernard NEw SPECIES (Figs. 3a—f, 4a-e; Table 1) Females.— Body length 9.2 (7.4-10.9) mm; all aspects of this species are similar to M. heterocercus unless otherwise noted. Head: 30 antennal segments, terminal segment with 2 whorls each with 4 placoid sensilla; posterolateral proliferation of setae occurring on segments 13-17. Labial palpus with 10+10 setae; maxillary palpus with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18+18 (15-21) setae on terminal segment; all laminae of lacinia pectinate (Table 1). Thorax: Mesonotum with 5+5M, 6+6m, and scattered microsetae (Fig. 3a). Abdomen: Tergites VI and VII with M1 and AM always present, posterolateral mar- gin of VI rectangular, VII projected back into sharp points, the latter occasionally fal- cate; tergite X setae A, B and C large, D minute or absent (Fig. 3b). Sternite I with 13+13M (12-14) in three transverse rows and scattered microsetae; antecedent setae 30+30 (29-32); subcoxal organs with 7+7 (4-10) glandular and 18+18 (15-23) sen- sory setae (Fig. 3c, d); median glandular or- gan protruding and with 11 (9-13) contig- uous disculi; sternites H—VII with 17+17M in four transverse rows. Genital papillae with 8+8 (5-10) sensory pegs; anterior and pos- terior lobes with 3+3 sensory pegs. Cerci: Right arm predental margin biser- rate with 1:2, 2:2 or 2:3 denticles; median tooth medium and sharply pointed; post- dental margin smooth to slightly crenulate and strongly falcate; chaetotaxy similar to M. heterocercus n. sp. except extra L4’ short, D1’ and D3’ long, and V7’ long (Fig. 4a, b). Left arm predental margin biserrate with 5 (4-5) dorsal toothlets and 7 (6-7) ventral denticles; median tooth small and sharply pointed; postdental margin smooth to slightly crenulate and sharply falcate; chae- totaxy similar to M. heterocercus n. sp. ex- cept extra L4’ short, D1’ long, D3’ long, and V7’ long (Fig. 4c, d). Males.— Body length 9.4 (9.3-9.4) mm. Similar to female except: tergite VI with M1 occasionally absent and posterolateral mar- gins usually projected back into sharp points; tergite VII with M1 always absent (Fig. 3e); sternite I with numerous fimbriate antece- dent setae restricted to posterior third of sclerite becoming sparse mesally, and a few simple antecedent setae in an irregular transverse row anterior of subcoxal organs becoming sparse mesally (Fig. 3f). Genital papillae conical, with 2+2 apical sensory VOLUME 92, NUMBER 4 799 Fig. 3a-f. Metajapyx magnifimbriatus. a, Mesothorax, L = 0.5 mm. b, Tergites VI-X, L = 1.0 mm. ¢, Ist abdominal sternite of female, L = 0.25 mm. d, Sensory and glanular setae of lateral subcoxal organ, L = 0.1 mm. e, Tergites VI-VII of male, L = 1.0 mm. f, Ist abdominal sternite of male, L = 0.25 mm. Terms: AM, anteromedian pair of setae; M, macroseta; AS, antecedent setae; GS, glandular setae; SS, sensory setae. pegs and numerous short setae mesally and long setae distally; genital opening with nu- merous minute marginal setae becoming longer distally. Cerci: Right arm with predental margin biserrate with 1:2 or 1:3 denticles, chaeto- taxy similar to female except L1 and L4’ absent, D2 and D4 minute, V1 and V2 ab- sent, V3 and V5 medium, V7 and V7’ ab- sent; left arm predental margin biserrate with 800 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4a-e. Chaetotaxy of male and female Metajapyx magnifimbriatus. a, Right dorsal view, L = 0.5 mm. b, Right ventral view, L = 0.5 mm. c, Left dorsal view, L = 0.5 mm. d, Left ventral view, L = 0.5 mm. e, Dorsal view of male cerci, L = 1.0 mm. Terms: D, dorsal setae; L, lateral setae; P, postdental margin setae; V, ventral setae. VOLUME 92, NUMBER 4 5 (5-6) dorsal toothlets and 5 (4-6) ventral toothlets, median tooth minute and sharply pointed; chaetotaxy similar to female except L1 absent, D1 short, V1—2 absent, V3 me- dium, V5 short, and V6-7 absent (Fig. 4e). Type material.—Holotype male and al- lotype female: Tennessee, Knox County, Cherokee Trail, 3.2 km south of the Uni- versity of Tennessee (elevation ca. 650 me- ters), 24-X-1988; M. A. Muegge. Paratypes (2 males, 3 females): Same data as holotype and allotype. Habitat.— All specimens were found 15-30 cm deep in moist clay soil in a beech-maple forest. The holotype, allo- type and two paratypes (1 male, | female) are deposited in the United States National Museum (USNM). The remaining para- types (1 male, 2 females) are deposited in the Apterygote Section of the University of Tennessee Entomology Museum, Knox- ville, TN. Etymology.—Latin magni (“great”), Latin fimbriatus (“fringed”), which refers to the numerous fringed antecedent setae of the male. Diagnosis.— Metajapyx magnifimbriatus most closely resembles M. multidens (Cook) 801 and M. propinguus (Silvestri), but may be easily distinguished from the latter two spe- cies by the following characters: 30 antennal segments, anteromedian pair of setae al- ways present on tergites VI and VII, seta M1 always present on tergites VI and VII of female (M1 absent on VII and sometimes on VI for male), median tooth of cerci small and sharply pointed, and postdental mar- gins of cerci smooth. LITERATURE CITED Pages, J. 1952. Diploures Japygidés de Nouvelle-Zé- lande. Rec. Canterbury Mus. 6: 149-162. Rathman, R. J., R. D. Akre, and J. F. Brunner. 1988. External morphology of a species of Metajapyx (Diplura: Japygidae) from Washington. Pan-Pa- cific Entomol. 64: 185-192. Smith, L. M. 1962. Japygidae of South America. 3: Japygidae of Chile. Proc. Biol. Soc. Washington. 75: 273-292. Smith, L. M. and C. L. Bolton. 1964. Japygidae of North America 9. The genus Metajapyx. J. Kansas Entomol. Soc. 37: 126-138. Steinmann, H. and L. Zombori. 1984. A Morpho- logical Atlas of Insect Larvae. Akademiai Kiado, Budapest, Hungary. 403 pp. Wilky, R. F. 1962. A simplified technique for clear- ing, staining and permanently mounting small ar- thropods. Ann. Entomol. Soc. Amer. 44: 606. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 802-807 SEASONAL ABUNDANCE AND HABITS OF THE BOXELDER BUG, BOISEA TRIVITTATA (SAY), IN AN URBAN ENVIRONMENT KAREN M. YODER AND WILLIAM H. ROBINSON Department of Entomology, Urban Pest Control Research Center, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061-0319. Abstract.— Boisea trivittata (Say) (Family Rhopalidae), the boxelder bug, is a common household pest in the United States. Large aggregations of adults and nymphs of this plant bug can form on the primary host tree, Acer negundo L., and other maples, or on buildings in urban environments. Overwintering boxelder bugs often seek harborage in protected areas around houses. The pest status of this insect is based primarily on the presence of large aggregations in the spring and the fall, and movement indoors where their excrement can stain fabric. Information on the biology and influence of substrate temperature on its activity was obtained from studies of laboratory and field populations in urban environ- ments in Virginia. Adult boxelder bugs were present twice during the calendar year 1988. Overwintering adults were active from April to June, and first generation adults were present from August to October. In laboratory colonies, the number of days between mating and egg laying was 1-5, the mean number of eggs per mass was 9.5 (laboratory) and 9.9 (field), and the mean incubation period was 10 days. The range of survival was 1-22 days for males, and 1-20 days for females. Adult and nymphal boxelder bugs responded to temperature differences by aggregating on the warmest substrates available. Thoracic temperatures of adult boxelder bugs were significantly higher than ambient air, and the differences between substrate and thoracic temperatures ranged from 3.4 to 7.0°C. Key Words: biology, mating, feeding, thoracic temperatures Boisea trivittata (Say), the boxelder bug, is a scentless plant bug in the family Rho- palidae. It isa minor pest of some fruit crops, and a widely distributed, common house- hold pest in the United States (Wheeler 1982). Boxelder bugs are usually found near their primary host, the boxelder tree, Acer negundo L. However, feeding is not restrict- ed to this species of maple. B. trivittata has been found feeding on other trees, including A. saccharinum (L.), Quercus spp., and Ai- lanthus altissima (Mill.) (McDaniel 1933, Smith and Shepherd 1937, Wheeler 1982). Large aggregations of adults and nymphs can form on houses during spring and fall and cause concern among homeowners. Overwintering boxelder bugs move into houses through open windows and doors, and cracks and crevices around windows, doors, and eaves in the fall. When indoors, adults can stain fabric with their excrement (Davis and Joos 1982). Taub (1970) re- ported severe asthmatic reactions during fall migrations of boxelder bugs into homes. The pest status of B. trivittata was reported by Swenk (1929), Ascerno (1981), and Pink- ston (1988). Early studies focused on the geographic distribution, taxonomy, morphology, and development of B. trivittata (Howard 1898, Smith and Shepherd 1937, Knowlton 1944, Tinker 1952). The objectives of the research Se ee VOLUME 92, NUMBER 4 reported here were to elucidate seasonal abundance, life cycle data, microhabitat temperature preferences, and boxelder bug activity in response to temperature in an urban environment. MATERIALS AND METHODS Seasonal abundance.— Field populations of boxelder bugs were sampled from April to December, 1988, at three houses, three apartment buildings, and one utility build- ing in Blacksburg, Virginia. These struc- tures were inspected three times per month for boxelder bugs. The base of the founda- tion walls on the south and west exposures were partitioned into one meter squares with spray paint. The wall, ground, and vegeta- tion located within sample squares were ex- amined, and all adult and nymphal boxelder bugs within the square were recorded. The total number of insects in each square were summed over all squares per site. At one apartment building (site F) three boxelder trees located approximately 10 m from the building were sampled for adults and nymphs with a plastic container and hand brush. The opening of the container contained a funnel and was held at the base of the tree. Boxelder bugs were lightly brushed from the bark into the container. After sampling, the containers were re- turned to the laboratory and the bugs were sexed, counted, and later returned to the trees and released. The sex ratio of overwintering adults was determined from boxelder bug populations at four building sites. Four samples were taken each week for a five-week period from September and December, 1988. Samples of soil and leaf litter were removed from the southwest exposure of the building and taken to the laboratory where boxelder bugs were removed, sexed and counted. Biological information. — Laboratory col- onies were initiated with adults obtained from field populations in Blacksburg, Vir- ginia in spring, 1988. Rearing methods were similar to those reported by Smith and 803 Shepherd (1937). Thirty-eight females and 30 males were maintained in the laboratory for life history studies. Pairs were main- tained in 3.8 liter glass jars and provided with fresh boxelder leaves and water. Water bottles and leaf material were replaced dai- ly. Mortality was recorded daily and dead individuals were removed and replaced. Egg masses were transferred from jars to petri dishes (100 x 15 mm). After hatch, first instar nymphs were transferred to plastic cups (9.5 x 5.5 x 8 cm) and supplied with small boxelder leaves. Cheesecloth was used to retain nymphs and adults in the contain- ers. Rearing conditions were 20-—22°C, 58- 62% RH, and 12:12 h photoperiod. Temperature selection.— Visual obser- vations were conducted on the activity of adults and nymphs in response to sunlight. The movement of second-generation box- elder bugs on the trunks of three boxelder trees was recorded several times during the day in September. Temperatures from substrates on three houses (A, B, C) in Blacksburg were record- ed from October 21 to November 15. The substrates included white-painted, alumi- num siding, concrete, brick, and colored canvas. The houses were selected on the ba- sis of existing infestations of boxelder bugs. An insulated thermistor probe attached to a micrologger (Campbell CR21 Microlog- ger) was used to measure temperatures in the microhabitats sampled. The styrofoam- insulated (2 layers) thermistor probe was attached to the substrates for 6 min. Am- bient air temperature was measured by plac- ing the probe in the air and recording for 15 min. The micrologger was programmed to record maximum, minimum, and aver- age temperatures every 3 min. Records were maintained for the time, Julian day, and microhabitat where the probe recorded temperature. Data were loaded onto a cas- sette recorder and transferred to an IBM mainframe computer program for analysis. Internal thoracic temperatures of 156 adult boxelder bugs at sites A, B, and C were 804 taken with a 0.03 cm chrome-alumel (Ome- ga) thermocouple insulated with Tygon tub- ing. The thorax was held with rubber gloves and forceps, and the thermocouple inserted posteroventrally between the second and third coxae. Because of the large size of the thermocouple, a time-constant of fifteen seconds was used to record temperatures. The specific microhabitat where the insects were found was also recorded. Data were analyzed using the SAS-GLM procedures (SAS Institute Inc. 1985); the Student-Newman-Keuls Range test was used to separate means. RESULTS AND DISCUSSION Seasonal abundance.— Adult boxelder bugs are present twice during the calendar year. The surviving overwintering adults are active in the spring. The first generation adults produced that year are present in late summer and fall, and form the overwinter- ing population. Wollerman (1965) reported two generations of the boxelder bug in the South and one in the North. Smith and Shepherd (1937) reported two generations of boxelder bugs in Kansas. Overwintering generation.— Adults were found primarily in leaf litter, on bark around A. negundo and A. saccharinum trees, and in mulch on southern exposures of resi- dences. During April, some adults observed at overwintering sites became active. Mat- ing activity was observed from March 11 to June 15 on the outside walls of resi- dences, at the base of A. negundo and A. saccharinum trees, and on low vegetation around residences. Females laid eggs predominantly on the surface of buildings in April and May. Egg deposition sites appeared to shift to low veg- etation from June to August. First-genera- tion eggs were first observed on April 14, and nymphs appeared during the second week of May. Egg deposition by first-gen- eration adults ceased the last week in June. The mean number of eggs per mass counted PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in the field was 9.9 (n = 123.0; SD = 4.3; range = 2-21). The mean percentage egg hatch was 84.4%. The number of nymphs increased during the last week of June, and numbers peaked during the first week of July. This increase was followed by a low number of nymphs observed from the end of July through August. The number of adult box- elder bugs at all sites remained low until mid-June. There was a gradual increase in the number of adults observed from mid- June to the end of July. Adults were observed mating and feeding on female boxelder trees in mid-July. No mating was observed on low vegetation dur- ing July or August. A general decrease in the adult population was observed during August. Tinker (1951) reported adults feed- ing and mating on female boxelder trees in August and September. The results from this study indicate that during late summer first- generation adults move from buildings to host trees and ground sites. First generation.—An increase in the number of first-generation nymphs oc- curred during late August to September (Fig. 2). A decline in the nymphal population oc- curred in October, and only small numbers of nymphs were observed from October through December. A decreased number of nymphs was followed by an increase in the number of adults observed on the surfaces of buildings in mid-October. Movement of adults to buildings at all of the sites occurred at this time. Tinker (1951) reported an abundance of adults in August and Septem- ber on female boxelder trees in Minnesota, which coincided with the development and maturity of the A. negundo ovules. Smith and Shepherd (1937) reported that boxelder bugs sought shelter in early October, and movement out of hibernation began in late March in Kansas. Time of dispersal to search for feeding or overwintering sites may vary with geographic location (Wollerman 1965). The sex ratio of second-generation adults was approximately 1:1. The ratio reported here contradicts the ratio (2:1, female : male) VOLUME 92, NUMBER 4 reported by Smith and Shepherd (1937). The method of sampling adults could account for the disparity in the ratio reported by Smith and Shepherd (1937) and that re- ported here. Life cycle Laboratory colony.— Mating did not oc- cur immediately after adults were paired in rearing jars. Mating behavior was not ob- served until several days after males and females were placed in the jars. When two males were placed with one female there was Often aggression between the males when both attempted to copulate with the female. Several observations were made of males that mounted females on the posterodorsal side and extended their beak between the elytra of the female. Of the 38 females ob- served, 16 did not lay eggs. The number of days between mating and egg laying ranged from | to 5 (n = 22). In the laboratory the mean number of eggs per mass was 9.5 (n = 33, SD = 7.7). Smith and Shepherd (1937) reported an av- erage of 10 eggs per mass. The mean incu- bation period calculated from all eggs for all batches was 10 days. The incubation pe- riod was 12 and 13 days for eggs from fe- males that laid 2 and 3 batches, respectively. Smith and Shepherd (1937) reported a range between |1 and 19 days and a mean of 13 days for boxelder bug oviposition period. Egg batches were laid in various size groups. Newly laid eggs were light orange, but turned dark red before hatching. Forty-one percent of the females laid no eggs, 43.6% laid 1 batch, 10.3% laid 2 batches, and 5.1% laid 3 egg batches. The mean survival of adults in the lab- oratory was 7.6 days (n = 37; SE = 5.6) for the females and 8.8 days for males (n = 29; SE = 6.27). The range of survival was 1-22 days for males and 1-20 days for females. The average number of days death occurred after eggs were laid was 4.3 days. 805 Microhabitat temperature selection Substrates.— Observations of second- generation adults and nymphs on three trees indicated that they oriented primarily to- ward the side of the trees that received the most sunlight. In the morning a large num- ber of bugs gathered on the eastern-exposed portion of the trunk, and in late afternoon more boxelder bugs were seen on the west- ern exposure. At approximately 9:00 am, an increase in adult and nymphal activity was observed, including movement to the ground at the base of the trees. They may have responded to a change in the incident angle of the sun and temperature. Tinker (1951) reported a preference for sunny ex- posures of A. negundo, but aggregations of bugs dispersed when the area became shad- ed. The mean temperatures recorded on southern-exposed substrates of three houses (A-C) are shown in Table 1. The number of boxelder bugs was greatest on substrates that had higher temperatures. At house A the mean temperature (26.5°C) recorded on the white-painted aluminum siding was not significantly different (P > 0.05) from the mean temperature (24.3°C) recorded at the concrete foundation. Although there was only a 2.2°C difference in temperatures re- corded from the two substrates, there were 113 boxelder bugs on the siding, and 23 boxelder bugs on the concrete foundation. At house B a mean temperature of 25.2°C was recorded on a window shutter, and this temperature was significantly different (P < 0.001) from the 17.9°C recorded behind a rain gutter on that house. Apparently, box- elder bugs at this site responded to the 7.3°C temperature difference by aggregating on the warmer substrate (240 on the window shut- ter vs. 8 at the rain gutter). Of the approx- imately 500 boxelder bugs collected at house C, 56 boxelder bugs were recorded on the cement foundation, which had a tempera- ture of 17.1°C. However, 240 boxelder bugs were collected from a piece of blue canvas 806 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Microhabitat mean temperatures for boxelder bugs at three sites. Site Microhabitat Temp (C)* No. Bugs A white aluminum siding 26.5a 113 white alum. siding (east) 15.4b 10 white alum. siding (north) 10.9¢ 0 white alum. siding (west) 27.4a 27 concrete foundation 24.3a 23 B window shutter 25.2a 240 brick 24.7a 140 rain gutter 17.9b 8 & blue canvas 24.9a 240 white aluminum siding 20.2b 121 concrete 17.1b 56 * Range of 2-5 separate temperature recordings. Means in each column followed by the same letter are not significantly different (Student-Newman-Keuls multiple range test; P > 0.05). (24.9°C) on the house. Tinker (1952) re- ported that as little as a 1.0°C to 3.3°C dif- ference between substrates resulted in box- elder bug aggregations on surfaces with the higher temperature. Insect.— Boxelder bug thoracic tempera- tures were significantly higher than the am- bient temperature; the differences ranged from 3.2 to 3.5°C at three locations (P < 0.001) (Table 2). The substrate surface tem- peratures were always significantly greater than the ambient and thoracic temperatures in the four locations sampled (P < 0.001). Differences between substrates and thoracic temperatures ranged between 3.4 and 7.0. The thoracic temperatures may more close- ly match the temperature of the air imme- diately above the substrate. Solar radiation heats the boundary layer of air above sun- lighted substrates. The temperature differ- ences between substrates, boundary layer air, and ambient may be several degrees (Heath and Wilkin 1970). Heath and Wilkin (1970) reported the thoracic temperature of a de- sert cicada to be closer to the temperature of the boundary layer air than to that of the substrate or ambient air. CONCLUSIONS The boxelder bug is a unique pest in the urban environment. Unlike a year-round indoor pest, such as the German cockroach, the boxelder bug 1s a pest only during spring and fall, but affects urban residents both indoors and outdoors. The apparent pref- erence of the adults and nymphs for warm or heat-retentive surfaces, and the large populations twice each year make this insect an important urban pest. Its pest status is based on the numbers found aggregating in Table 2. Mean comparisons of thoracic, substrate and ambient air temperatures of boxelder bugs at four locations. Temperatures (n) and Locations Subject A B c D Substrate 28.7a (38) 28.5a (6) 20.3a (11) 20.8a (8) Thorax 24.9b (8) 21.5b (27) 16.9b (42) 19.8a (17) Ambient air 21.4c (14) 21.9b (20) 13.7c¢ (10) 16.3b (17) Means in each column followed by the same letter are not significantly different (Student-Newman-Keuls multiple range test; P > 0.05). VOLUME 92, NUMBER 4 and around houses, and its pest importance is indicated by the amount of money urban residents spend on control (Yoder and Rob- inson 1988). Nonchemical control strategies for this insect are limited. Smith and Shepherd (1937) reported that no parasites emerged from 452 eggs they observed during rearing. Smith and Shepherd (1937) reported para- sitic flagellates in the intestinal tract of B. trivittata, but did not appear to cause mor- tality. Removal of the host tree may be ef- fective (Davis and Joos 1982), and sealing crevices in structures may prevent boxelder bugs from moving indoors. However, Smith and Shepherd (1937) and Wheeler (1982) reported that B. trivittata can feed on a va- riety of ornamental trees and plants. Chem- ical control of this insect presents several problems. Because this insect is a problem to homeowners primarily during spring and fall, chemical controls are usually initiated after large numbers of bugs aggregate around houses. Photodegradation and volatiliza- tion of commonly used insecticides can oc- cur because they are applied to sun-exposed and heat-retaining surfaces. These surfaces are attractive to the insects, but limit insec- ticide effectiveness. It would be more effec- tive to time and direct chemical applica- tions to the eggs and early instars of the first generation. ACKNOWLEDGMENTS We thank D. Mullins and R. Fell, VPI&SU Entomology Department, for their suggestions and help with the temperature studies. A. G. Wheeler, Jr., Pennsylvania Department of Agriculture, reviewed and improved the manuscript. 807 LITERATURE CITED Ascerno, M. 198i. Diagnostic clinics: More than a public service? Bull. Entomol. Soc. Am. 27: 97- 101. Davis, C. S. and L. Joos. 1982. Controlling western boxelder bug. Leaflet— University Calif. Cooper- ative Extension Service. 2 pp. Heath, J. and P. Wilkin. 1970. Temperature re- sponses of the desert cicada, Diceroprocta apache. Physiol. Zool. 43: 145-154. Howard, L. O. 1898. The box-elder plant bug. U.S. Dep. Agric. Div. Entomol. Bull. 28: 1-3. Knowlton, G. F. 1944. Boxelder bug observations. J. Econ. Entomol. 37: 443. McDaniel, E. J. 1933. The boxelder bug as a house- hold pest. Quart. Bull. Mich. Agric. Exp. Stn. 15: 226-227. Pinkston, K. 1988. Results of an industry survey. Oklahoma Pest Control Association Annual Con- ference. Coop. Ext. Serv. 6 pp. SAS institute Inc. 1985. SAS User’s Guide; Statistics, Version 5 Edition. SAS Inst. Inc., Cary, NC. 956 pp. Smith, R. C. and B. L. Shepherd. 1937. The life his- tory and control of the boxelder bug in Kansas. Trans. Kans. Acad. Sci. 40: 143-159. Swenk, M. H. 1929. Boxelder bug (Leptocoris trivit- tatus Say). Insect Pest Surv. Bull. 9(3): 87. Taub, S. J. 1970. Severe asthma attacks caused by gnats and box elder bugs. Eye, Ear, Nose and Throat Month. 49: 475. Tinker, M. E. 1951. The seasonal behavior and ecol- ogy of the boxelder bug, Leptocoris trivittatus Say. Master of Science thesis. University of Minnesota. 36 pp. 1952. The seasonal behavior and ecology of the boxelder bug Leptocoris trivittata in Minne- sota. Ecology 33: 407-414. Wheeler, A. G., Jr. 1982. Bed bugs and other bugs, pp. 319-351. Jn Mallis, A., ed., Handbook of Pest Control. 6th ed. Franzak & Foster Co., Cleveland, Ohio. Wollerman, E.H. 1965. The boxelder bug [Leptocoris trivittatus, Acer negundo]. U.S. For. Serv. Forest Pest Leaflet. 6 pp. Yoder, K. and W. Robinson. 1988. Do you know how to identify and control boxelder bugs? Pest Control Technology 16(5): 54, 58. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 808-810 EGG DISPERSION IN TWO SPECIES OF PRAYING MANTIDS (MANTODEA: MANTIDAE) R. M. EISENBERG AND L. E. HurRD Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware 19716. Abstract. — Tenodera sinensis (Saussure) and Mantis religiosa (Linnaeus) are univoltine generalist predators which produce eggs at the end of the growing season. Oothecae of both species exhibit a markedly contagious dispersion in old fields in northern Delaware. In view of the large number of eggs contained in each ootheca, the propensity for syn- chronous egg hatch for each species, and severe food limitation during emergence early in the spring, such clumping is surprising since it places newly-hatched nymphs at a greater risk of cannibalism and competition from their cohort than if they were more uniformly distributed in space. Possible explanations for such clumping are discussed. Key Words: ators The praying mantids Tenodera sinensis (Saussure) and Mantis religiosa (Linnaeus) are generalist predators which commonly occur in old fields in northern Delaware. Both species produce oothecae which may contain several hundred eggs each, and al- though eggs of 7. sinensis generally hatch before those of M. religiosa, there is con- siderable intraspecific synchrony (Hurd and Eisenberg 1988a). Egg hatch occurs early in the spring, when there is normally a short- age of suitable prey (Hurd and Eisenberg 1988b, Rathet and Hurd 1983). We have shown that early instar nymphs of 7. si- nensis show a strong tendency to disperse (Hurd and Eisenberg 1984). We also know that first stadia of both species will resort to cannibalism when alternative food 1s ab- sent (Hurd and Eisenberg 1984, unpub- lished laboratory observations). Given the above information, it would seem reason- able to expect that for both of these species, oothecae should not be clumped. Females which deposit an egg mass close to another Mantodea, Mantidae, Tenodera sinensis, Mantis regligiosa, dispersion, pred- egg mass would seem to be placing their young at a disadvantage in terms of increas- ing the potential for both intraspecific com- petition and cannibalism (Hurd 1988). We therefore decided to determine the spatial dispersion of oothecae of these two species in local old fields. MATERIALS AND METHODS Three different old fields in the vicinity of the University of Delaware campus in Newark, Delaware, were used for the col- lection of data. One was a late successional goldenrod field (field #1); the other two were sites AG (field #2) and CHRY (field #3) referred to by Hurd and Eisenberg (1988a, b). All three had been examined for several years and were known to harbor persistent populations of mantids. The portion of each field which consti- tuted mantid habitat was staked out. Areas thereby delineated were searched for oothe- cae and the location of each was marked with a 1.0 m wooden dowel. After searching VOLUME 92, NUMBER 4 Table 1. Results of oothecae censuses and nearest neighbor analyses for three fields. Ts = Tenodera si- nensis, Mr = Mantis religiosa. R = ratio of observed mean distance to nearest neighbor to expected mean distance to nearest neighbor. P values are based on values of C (Clark and Evans 1954). Number Site ° Area Oothe- P. Site (m’) Species cae R Value Field #1 1200 Ts 119 0.76 <.01 Field #2 700 ~=Ts 59 0.48 <.01 Field #3 1000 Mr 101 0.76 <.01 was completed the distance of each ootheca to two reference stakes was measured to the nearest cm using a pair of measuring tapes. In the laboratory this information was used to locate each ootheca on a scale map of each area, and dispersion was determined by nearest neighbor measurements (Clark and Evans 1954). RESULTS AND DISCUSSION While we encountered both species of mantids in each of our collections, only a single species was numerous enough in each field to permit dispersion analysis. In fields #1 and #2 the dominant mantid was 7. si- nensis, and oothecae were mainly located on dead, upright plant stems from 0.3 to 1.0 m above the ground. In field #3 the domi- nant mantid was M. religiosa, the oothecae of which were mainly found in the dense, overlapping grasses which comprised ground cover in these fields. In all three fields the nearest neighbor analysis shows a highly sig- nificant departure from random expectation (Table 1). An R value of 1.0 is expected if the observed pattern is random. Our R val- ues of 0.48 to 0.76 indicate that the distri- butions of oothecae of both M. religiosa and T. sinensis are very contagious; both of these mantid species show a strong tendency to deposit egg masses in close proximity to other conspecific oothecae. The average field-collected ootheca of 7. sinensis weighs approximately 1.9 g and re- leases 240 nymphs (Eisenberg and Hurd 809 1977). Mean weight of M. religiosa oothecae collected from our study fields was 1.07 g, emergence ranging from 30 to 370 nymphs with a mean of 156. Both species display considerable synchrony of emergence in the field. Thus, in addition to the high local density produced by the hatching of a single ootheca, the close proximity of additional oothecae can result in even higher densities. In field #1, 29.4% of oothecae were within the same | m? area as another ootheca and 48.7% were within the same 2 m? area. In field #2, 61.0% of oothecae were within the same | m? area as another ootheca and 71.2% were within the same 2 m? area. For field #3 the values were 19.8% and 30.7% respectively. Thus at emergence time, local densities of nymphs easily could reach or exceed 300 to 400 nymphs per m?. How can we explain the contagious na- ture of the oothecae pattern? During the late summer and fall of each year, 7. sinensis often can be found on inflorescences of late- flowering plants such as goldenrods and as- ters, which attract prey in the form of flow- er-foraging insects including pollinators. This represents an important source of nu- trition for females while they are undergoing oogenesis; females so positioned produce more eggs than those which are on plants not in flower (Hurd 1989). Females gener- ally do not move around once they mature (Bartley 1982), so that a female’s position on a specific plant (in flower or not) may be a matter of chance rather than choice. How- ever, the clonal nature of these flowering plants produces clumps of the most nutri- tionally rewarding oviposition sites, which could in turn explain contagion among oo- thecae. An alternative explanation, that fe- males oviposit more than once, 1s less likely because normally there is not sufficient time between first oviposition and killing frost for this species to generate a second ootheca in our geographical region in the face of rap- idly decreasing food levels at the end of the growing season (Eisenberg, Hurd and Bar- tley 1981). 810 Mantis religiosa exhibits a very different set of behaviors in regard to its foraging activities. This species tends to forage closer to the ground than 7. sinensis (Rathet and Hurd 1983), and thus is less likely to be found on the taller flowering plants. While this species also will deposit its oothecae on upright stems, apparently it prefers grasses located much closer to the ground. These grasses do not have flowers to attract sup- plemental food, and constitute a much denser, less patchy vegetational layer than goldenrod. We do not know if this species faces the same degree of food limitation as T. sinensis at the end of the growing season, so that it is possible that multiple oviposi- tion is responsible for the contagious dis- tribution of M. religiosa oothecae. How- ever, this still leaves open the question as to why a female would crowd her own off- spring, an apparently maladaptive trait. ACKNOWLEDGMENTS This work was supported by NSF grant BSR 8506181. This is contribution #137 from the Ecology Program, University of Delaware. LITERATURE CITED Bartley, J. A. 1982. Movement patterns of adult male and female mantids (7enodera sinensis Saussure). Environmental Entomology 11: 1108-1111. Clark, P. J.and F.C. Evans. 1954. Distance to nearest PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON neighbor as a measure of spatial relationships in populations. Ecology 35: 445-453. Eisenberg, R. M.and L. E. Hurd. 1977. An ecological study of the emergence characteristics for egg cases of the Chinese mantis (Tenodera aridifolia sinensis Saussure). American Midland Naturalist 97: 478- 482. Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981. Ecological consequences of food limitation for adult mantids Tenodera sinensis Saussure). American Midland Nat. 106: 209-218. Hurd, L. E. 1988. Consequences of divergent egg phenology to predation and coexistence in two sympatric, congeneric mantids (Orthoptera: Man- tidae). Oecologia 76: 547-550. 1989. The importance of late season flowers to the fitness of an insect predator, Tenodera si- nensis Saussure (Orthoptera: Mantidae), in an old field community. Entomologist (U.K.) 108: 223- 228. Hurd, L. E.and R. M. Eisenberg. 1984. Experimental density manipulations of the predator Tenodera sinensis (Orthoptera: Mantidae) in an old-field community. I. Mortality, development and dis- persal of juvenile mantids. Journal Animal Ecol- ogy 53: 269-281. 1988a. A mid-summer comparison of sizes and growth rates among nymphs of three sym- patric mantids (Mantodea: Mantidae) in two old- field habitats. Proceedings Entomological Society Washington 91: 51-54. 1988b. Temporal distribution of hatching times in three sympatric mantids (Mantodea: Mantidae) with implications for niche separation and coexistence. Proceedings Entomological So- ciety Washington 91: 55-58. Rathet, I. H. and L. E. Hurd. 1983. Ecological rela- tionships among three co-occurring mantids. Te- nodera sinensis (Saussure), 7. angustipennis (S.), and Mantis religiosa. American Midland Natu- ralist 110: 240-248. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 811 NOTE First Distributional Records of Cimexopsis nyctalis List (Hemiptera: Cimicidae) in Connecticut Cimexopsis nyctalis List 1s restricted to the eastern United States where it inhabits nests of the chimney swift, Chaetura pelag- ica (Linnaeus) (Usinger 1966. Monograph of Cimicidae (Hemiptera—Heteroptera), Vol. 7. Thomas Say Foundation. Entomological Society of America, Baltimore, Maryland. 585 pp.). This insect is known to occur in 16 states (Lee 1955. Bull. Brooklyn Ento- mol. Soc. 50: 51—52.), (Usinger, ibid.). Three unreported records of C. nyctalis from Connecticut are: New Haven County, Branford, 30 July 1942 (1 specimen); Litch- field County, Thomaston, | August 1942 (3 specimens from a chimney); and Tolland County, Manchester, 22 August 1988 (1 specimen). The specimens from Branford and Thomaston, housed in the Connecticut Agricultural Experiment Station insect col- lection, were previously identified incor- rectly as Oeciacus vicarius Horvath. The specimen from Manchester was collected inside a home where birds were nesting in the chimney during 1987 and 1988. The person who collected C. nyctalis in Man- chester reported that she was bitten by sim- ilar insects. C. nyctalis, whose only recorded host is C. pelagica, is not known to bite humans. However, it is not uncommon for other species of bird bugs to do so on oc- casion. For example, Harwood and James (1979. Entomology in Human and Animal Health. Macmillan Publishing Co., Inc., New York) reported that the swallow bug, O. vicarius Horvath, and the Mexican chicken bug, Haematosiphon inodorus (Duges), bite humans infrequently. All spec- imens are deposited in the insect collection of the Connecticut Agricultural Experiment Station. No specimens were found in the insect collections of the University of Con- necticut (Storrs, CT) and Peabody Museum, Yale University (New Haven, CT). I thank Edmond Marrotte, of the Uni- versity of Connecticut Cooperative Exten- sion Service, for sending me the specimen from Manchester; Carl W. Schaefer from the University of Connecticut and Ray- mond J. Pupedis from the Peabody Muse- um, Yale University for checking their re- spective collections; and T. J. Henry, Systematic Entomology Laboratory, PSI, USDA, for confirming my identification. Kenneth A. Welch, Department of En- tomology, The Connecticut Agricultural Ex- periment Station, P.O. Box 1106, New Ha- ven, Connecticut 06504-1106. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 812 NOTE Liliacina diversipes (Kirby) (Hymenoptera: Tenthredinidae), a Sawfly Genus and Species New to the United States Dr. Henri Goulet, Biosystematics Re- search Centre, Agriculture Canada, Ottawa, discovered two specimens of a neotropical- like sawfly collected by Malaise trap in Ala- chua Co., Florida. These specimens are here identified as Liliacina diversipes (Kirby) (Se- landriinae), a species native to Mexico and Central America. The species is apparently established in Florida and undoubtedly the result of an accidental introduction. Both the genus and species are new to the United States. The host is not known, though most nearctic Selandriinae feed on various ferns and sedges. Liliacina was described by Malaise (1942, Ent. Tidskr. 63: 94, 99-100; type species— Liliacina carinifrons Malaise), includes about 8 species, and occurs from Mexico to southeastern Brazil and northern Argenti- na. It is distinguished from other North American Selandriinae genera (revision by Smith, D. R. 1969, U.S. Dept. Agr., Tech. Bull. 1398, 48 pp.) by the clypeus shallowly, semicircularly emarginate; tarsal claw with long inner tooth but without basal lobe; epicnemium (previously termed prepectus) elongate, on same level as mesepisternum and separated from mesepisternum by su- ture; genal carina absent; anal crossvein of forewing absent; width of malar space about half diameter of an ocellus; and antennal pedicel slightly longer than wide. Liliacina diversipes (Kirby 1882, List Hym. Brit. Mus., vol. 1, pp. 189-190, pl. 8, fig. 22; described as Selandria diversipes: transferred to Liliacina by Malaise 1942) was described from “Mexico, Orizaba.” I have seen specimens from Mexico (Jalisco, Veracruz), Guatemala, El Salvador, Costa Rica, Panama, and Colombia. The species is 8.0-9.0 mm in length, and coloration dis- tinguishes it from all other North American Selandriinae: Head and antenna black with scape and pedicel brownish and clypeus, la- brum, and base of mandible white; apex of mandible reddish; palpi blackish though basal segments and labium whitish. Thorax whitish (pale areas probably lilac-colored when alive) with cervical sclerites, meso- sternum (except medial stripe), lower half of mesepimeron, most of mesoprescutum, most of mesoscutellum, and metathorax (except upper portion of metapleuron) black; mesonotal lateral lobes whitish to orange. Abdomen black, narrow posterior margin of segments whitish. Legs white with ex- treme base of hindcoxa, apical half of fore- and midtibiae, apical third of hindtibia, and all tarsi black. Wings blackish; veins and stigma black. The pale areas of the thorax are bright lilac in some neotropical speci- mens examined, but this turns to a sordid whitish in some preserved specimens, es- pecially those collected in alcohol. The collection data are as follows: USA: Fl., Alachua Co., Gainesville, AEI [Amer- ican Entomological Institute], B.R.C. [Bios- ystematics Research Centre] Hym. Team. The two females were collected on 3-17- VII-1987 and 10-31-VIII-1987. David R. Smith, Systematic Entomology Laboratory, PSI, ARS, U.S. Department of Agriculture, % National Museum of Natural History, NHB 168, Washington, D.C. 20560. er a PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 813 NOTE Coptotermes crassus Ping Preoccupied by C. crassus Snyder Renamed C. pingi (Isoptera: Rhinotermitidae) In the course of compiling a list of species of the Isoptera of the world, an unusual tax- onomic mistake was discovered. A newly described termite from China of the genus Coptotermes was given the specific name crassus (Ping, Z. 1985. Eight new species of the genus Coptotermes and Reticulitermes from Guangdong Province, China. Ento- motaxonomia 7(4): 317-328 (pp. 319-320, 327, fig. 3A-E)). Unfortunately, the name Coptotermes crassus 1s already occupied by a Neotropical species (Snyder 1922. Proc. U.S. Natl. Mus. 61: 1-32 (pp. 21-22, fig. 6)). Thus according to Article 60(a) of ICZN (Ride et al. 1985. Internatl. Code Zool. No- men. 3rd ed., Univ. Cal. Press, Berkeley, 338 pp.) Coptotermes crassus Ping is a ju- nior homonym and must be rejected. Since Ping placed crassus directly in the genus Coptotermes, 1.e. this combination did not arise from revision, C. crassus Ping 1985 is known as an objectively invalid primary ju- nior homonym. Barring the possible avail- ability of other names, a replacement name (nomen novum) with its own author and date needs to be designated (Article 60(c)). There is no reason to suspect that C. crassus Snyder and C. crassus Ping are synony- mous—neither on the basis of morphology nor on the basis of geographic distribution (Araujo 1977. Catalogo dos Isoptera do Nova Mundo. Academia Brasileira de Cien- cias, Rio de Janeiro). The only possible available name for a Coptotermes from that area is hongkonensis Oshima (Oshima 1914. Annot. Zool. Jap. 8: 553-585 (pp. 559-560, pl. IX, fig. 5)) which was sunken into syn- onymy with Coptotermes formosanus Shiraki. However, the ranges of all the measurements given by Oshima for hong- konensis are outside of those given for cras- sus Ping. Other details of the descriptions and illustrations also do not match. The right soldier mandible of crassus Ping is strongly curved toward the tip while it is relatively straight, ““saber-shaped,” in the photograph of hongkonensis. The soldier gula is strongly contracted in the middle in crassus Ping while only “‘slightly contracted at middle” in hongkonensis. The labrum reaches well beyond the middle of the soldier mandibles in crassus Ping while Oshima states that in hongkonensis it is “scarcely reaching the middle of the mandibles.”’ Therefore, since C. crassus Ping cannot be referred to C. hongkonensis, and since there are no other available names from that area I propose the new name: Coptotermes pingi Myles Nomen Novum = Coptotermes crassus Ping 1985. It is appropriate to rename this species after Dr. Ping Zhengming in recognition of his substantial contributions to termite tax- onomy in China. Timothy G. Myles, Research Associate, Department of Entomology, University of Arizona, Tucson, Arizona 85721. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 814-815 Book REVIEW Bird Blow Flies (Protocalliphora) in North America (Diptera: Calliphoridae) with Notes on the Palearctic Species. C. W. Sa- brosky, G. F. Bennett, and T. L. Whit- worth. Smithsonian Institution Press, Washington, D.C., and London [x and] 312 pp., paperbound. 1989. $16.95. Or- der from: Smithsonian Institution Press, Dept. 900, Blue Ridge Summit, PA 17294 (add $2.25 for postage and handling). Maggots of bird blow flies are obligatory blood feeders that live exclusively at the expense of nestling birds. Adult flies are not commonly seen or collected in flight. They are fairly large, 9-12 mm long, and most commonly dark metallic blue, although the females of some species are bronze or cop- per. Protocalliphora is a Holarctic genus; 26 species, 15 of them new to science, are now known to occur in North America, and the ranges of two species extend into Europe. This work is a fine, comprehensive sys- tematic study of the genus Protocalliphora and its Nearctic species. It is based on Sa- brosky’s long-term taxonomic studies, Ben- nett’s and Whitworth’s individual field in- vestigations, and Bennett’s anatomical studies of the immature stages, all collated here by Sabrosky. The book is divided into two parts. The first, 43 pages long, is a gen- eral exposition of the taxonomy, geograph- ical distribution, life history, and ecology, with tips on study and control. The re- mainder of the book, titled the taxonomic section, covers generic taxonomy, anatomy of all stages supported by illustrations for adults and a glossary for immature stages, keys to species, and individual treatment of all species. The descriptions are followed by a list of the avian hosts of Nearctic Proto- calliphora, annotated references, an index to bird hosts and one to Protocalliphora and miscellaneous names, plates of illustrations and distribution maps. The first part, at least, should fascinate anyone with passing interest in bird life, and the whole book, even the “‘legal”’ aspects of taxonomy, is engagingly written. The au- thors point out that these insects are barely or not at all mentioned in bird books or textbooks on parasitology, even though the maggots are common and numerous in bird nests. Publication of this book should make researchers much more aware that these in- sects exist. The species treatments are helpfully bro- ken down into short, topical subsections. The bibliography is annotated, a measure of the care taken with the book. I like the combined key to males, females, and pu- paria, which the authors show is the best way to determine Protocalliphora species. Anyone doing serious work with these flies will have both sexes and all stages available. Additional keys to each sex alone and to third instars are here also, but are less useful: the few easy-to-separate species are keyed first, but as the key goes along differences become finer so that a person will be less sure of what is in hand. The authors wring their hands a little too much for my taste about what they consider to be drawbacks of the study: that it is not definitive (preface), that the distribution of many species is spottily known and that few areas have been studied thoroughly (p. 3), that any general statements should be qual- ified ‘‘as far as known”’ (p. 16), that the en- tire life history has not been followed for any species (p. 20), and that speculation on phylogeny seems premature (p. 37). These are all true statements that could have been rephrased in a positive manner, and de- servedly so for all the painstaking study and time that the authors have invested in this project. This book will still be the author- itative work on Protocalliphora for a long time to come. Non-entomologists who need this book and have no mental picture of a calliphorid may be put off by the lack of illustrations VOLUME 92, NUMBER 4 of an entire adult and larva. An illustration of a larva to show the characteristic fringe of long setae around the head segment would also have served to distinguish Protocalliph- ora from other maggots likely to be found in bird nests. Reference is made to a draw- ing of anterior parts of a larva in another work, but that book may not be readily available to all. Further, books for general readers shouldn’t have a lengthy quote in a foreign language (p. 9) or use dipterists’ jar- gon or technical terms (eclose, myiasis, pu- parium) without a glossary. Authors’ lapses are few and minor, among which are the lack of an authorial reference for a future publication on details of life history and bi- ology (top p. 24) and five misspelled place names. A publisher’s note on the verso of the title page advises that, ‘For reasons of speed and economy, this book is prepared from cam- era-ready copy prepared electronically by the authors, who assume full responsibility for the contents and form.” The claim not- withstanding, I cannot fault the authors for 815 the Press’s lack of oversight on this book. It resulted in weak major headings, the awk- ward blank bottom third of p. 56, the need of additional lines and leaders in some key couplets, a clumsy insert in the key on p. 86 (couplet 9a), the indexes appearing be- fore the end of the book, and the fact that there are two indexes (one for bird names, one to other names) where one index would have served better. The indexes inconve- niently precede the 33 pages of plates and maps, which themselves are not indexed. Further, the figures on pages 276-298 are reduced too much and are less effective for it; they could have been shown twice as large if they had been reconfigured. It is under- standable that a press would want to keep costs low, but it must still attract authors and stand behind its product. Raymond J. Gagné, Systematic Ento- mology Laboratory, PSI, Agricultural Re- search Service, USDA, % USNM, NHB 168, Washington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 816-817 Book REVIEW Insect Flight. 1989. G. J. Goldsworthy and C. H. Wheeler (eds.) C.R.C. Press, Inc., Boca Raton, Florida. Few insect behaviors rival the signifi- cance of flight. Considering all the inver- tebrates, insects are the only ones to explore and succeed in using flight as a mechanism to locate new breeding sites, find food, es- cape predators, avoid unfavorable environ- mental conditions, and search for potential mates. /nsect Flight is a literary tribute to this singular behavior that has culminated in the success of insects. This recent look at flight is an attempt to provide its readers with an up-to-date synthesis of the diverse topics that encompass this most important insect adaptation. The editors Graham Goldsworthy and Colin Wheeler should be complimented for providing such an im- pressive group of contributing authors (i.e. 18 in all). The coverage is broad and ranges in scope from the mechanisms and aero- dynamics of flight (Chapt. 1) to the impor- tance of understanding flight in the context of developing better control strategies (Chapt. 15). The chapters are well written, up-to-date, provide excellent illustrations and contain few editorial mistakes. Overall, this book is for specialists who are interested in specific aspects of insect flight. The au- thors selected for this overwhelming task were well chosen and are leaders in their respective areas. Some of the topics pre- sented have been covered by the same au- thors elsewhere; however, they all have up- dated the information and presented new ideas. I am taking the privilege to list all the chapters presented and the respective au- thors. The reasons for doing this are two- fold: cost and specialization. Because of the exorbitant price (i.e. $195 U.S., $225 out- side U.S.) for the 371 pages, I strongly rec- ommend that individuals interested in just one or a few chapters contact a wealthy friend for a loaner or, for most of us (i.e. either poor or not having wealthy friends), have the library order it. The only glaring weak- ness of the book, despite its cost, is the last two chapters. The authors of Chapt. 14 make the following statement on p. 322: “Our ob- jective in this chapter is to describe the pres- ent state of insect control, considering its failures as well as its successes, because it 1s only by taking this ‘warts and all’ approach that we can realistically set the scene for an analysis of future requirements.” The au- thors completed this objective, even if it is disjointed from the rest of the book. The final decision to make sure that all the chap- ters of a book somehow relate to the theme of the book lies with the editors and here they failed. Not only did they fail to rec- ognize that this chapter, as written, did not fit well into this volume, but they missed an excellent opportunity to stress the im- portance of flight to current and future con- trol strategies. I am sure the editors initially recognized the importance of including a chapter that would integrate these two top- ics, otherwise they wouldn’t have included it in the volume. They could have discussed such topics as dispersion, flight capabilities of different pests, founder effects on resis- tance management strategies, passive flight of certain vectors (i.e. aphids) of plant dis- eases and their dependence on prevailing winds, and effect(s) parasite loads may have on flight in medically important species. Chapt. 15, by the same authors as Chapt. 14, again met the stated objective of the authors but failed to hit the target. Its pres- ence, however, in a treatise that is mainly biased towards basic research should serve an important function. Chapt. 14 provides the novice with an excellent overview of future control strategies. Maybe, and just maybe, individuals interested in the more basic aspects of flight biology will read it. If they do, and I strongly recommend that they do, they should attempt to relate how their VOLUME 92, NUMBER 4 area of interest in insect flight relates to im- proving control strategies. I am sure that this type of synthesis is what the editors wanted to promote when they decided to include these last two chapters. This reviewer recommends you treat this book as a buffet: only take those dishes that interest you. Following is the menu: Chapt. 1. Mechanics and Aerodynamics of Flight — W. Nachtigall, pp. 1-29; Chapt. 2. Structure and Function in Flight Muscle—D. J. Aid- ley, pp. 31-49; Chapt. 3. Development of the Flight Motor Pattern— Wolfram Kutsch, pp. 51-73; Chapt. 4. Sense Organs and the Control of Flight— Bernhard M6hl, pp. 75- 97; Chapt. 5. The Evolution and Signifi- cance of Migratory Flight— Hugh Dingle, pp. 99-114; Chapt. 6. Genes, Environment, and Insect Flight—A. G. Gatehouse, pp. 115- 138; Chapt. 7. Hormonal Control of Flight — Mary Ann Rankin, pp. 139-163; Chapt. 8. Swarm Flight Behavior in Flies and Lo- 817 custs— Richard John Cooter, pp. 165-203; Chapt. 9. Orientation and Foraging in Hon- eybees— Fred C. Dyer and Thomas D. See- ley, pp. 205-230; Chapt. 10. Pheromones and Flight Behavior—T. C. Baker, pp. 231- 255; Chapt. 11. Oxygen Consumption Dur- ing Flight—Timothy M. Casey, pp. 257- 272; Chapt. 12. Mobilization and Transport of Fuels to the Flight Muscles—Colin H. Wheeler, pp. 273-303; Chapt. 13. Utiliza- tion of Fuels by the Flight Muscles—D. J. Candy, pp. 305-319; Chapt. 14. Problems in the Control of Flying Insect Pests—D. P. Giles and A. R. Jutsum, pp. 321-336; Chapt. 15. Prospects for Better Control Strategies — A. R. Jutsum and D. P. Giles, pp. 337-371. Reviewed by John G. Stoffolano, Jr., De- partment of Entomology, Fernald Hall, University of Massachusetts, Amherst, Mas- sachusetts 01003. PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 818-824 SociETY MEETINGS 954th Regular Meeting—January 4, 1990 The 954th Regular Meeting of the Ento- mological Society of Washington was called to order by President Jeffrey R. Aldrich in the Naturalist Center, National Museum of Natural History, at 8 p.m. on 4 January 1990. Twenty-two members, two guests and a joker were present. Minutes of the De- cember meeting were read and approved. Membership Chairman G. B. White read the names of the following applicants for membership: James M. Hill, Natural Re- sources Division, Maryland-National Cap- ital Park and Planning Commission, Chevy Chase; Roberta (“Bobbe’’) Krueger, Hay- market, Virginia; Timothy George Myles, Department of Entomology, University of Arizona, Tucson; and David H. Young, Waltham, Massachusetts. President Aldrich reviewed the results of the Executive Committee meeting held ear- lier in the day. The Committee has decided to hold our Regular Meeting for May in the Visitors Center (or “Log Lodge”) at the Beltsville Agricultural Research Center. Tentative plans for this event include talks by a number of entomologists belonging to the Agricultural Research Service. The Ex- ecutive Committee also noted that our an- nual donation of $750 to the American As- sociation for Zoological Nomenclature is now a routine feature of ESW budgets (Proc. Entomol. Soc. Wash. 91: 651, 653, October 1989) but that the nation’s largest ento- mological organization, the Entomological Society of America, has refused to support the AAZN, ostensibly because all such con- tributions must be approved by the entire ESA membership. However, it is well known that there is pressure within ESA to bring this matter to a vote, and Dr. Aldrich sug- gests that our Society can encourage a fa- vorable outcome by publishing a letter of challenge in either the ESA Newsletter or Bulletin. Doug Sutherland circulated a notebook containing letters from the American and North Carolina entomological societies supporting the selection of the monarch butterfly, Danaus plexippus (Linnaeus), as our national insect. R. G. Robbins thanked J. M. Kingsolver for providing the Smithsonian’s Acarology Unit with a recent listing of 139 references on Lyme disease produced through the Quick Bibliography Series of the National Agricultural Library (NAL-BIBL. QB 89- 87). All bibliographies in this series are de- rived from searches of the AGRICOLA da- tabase and can be obtained by writing to the NAL, Public Services Division, Room 111, Beltsville, Maryland 2070S. The speaker for the evening was our Past President, F. Christian Thompson, System- atic Entomology Laboratory, U.S. Depart- ment of Agriculture, whose talk was entitled “Will Systematics Survive the 21st Centu- ry?” Dr. Thompson is pessimistic, in part because entomology, like other branches of natural science, has suffered a certain loss of identity in recent decades; in part also because systematists are often their own worst enemies, lending credence to the view that they are just esoteric academics; but largely because taxonomists across the board are so intransigently conservative, refusing to adapt to, let alone take advantage of, technological advances that could automate their workplaces, save them countless thousands of hours, and even elevate their profession in the eyes of the nonsystematic scientific community. Given the immensity of the systematist’s task —that of generating and disseminating biosystematic informa- tion—and the presumably permanent pau- city of funds, space and personnel, there is no question that automation is the key to survival. But in any such scheme, system- atists are the bottleneck, so their modus op- erandi must be to divert some of the ava- lanche of data from themselves while VOLUME 92, NUMBER 4 increasing the rate of data flow. In a word: computerize! Presently, a relational data- base model for biosystematic information is being developed by Dr. Thompson and several colleagues. This and other expert systems will use names of taxa, characters, attributes, and certain comments (e.g. ge- ography, season) to generate biosystematic information with unsurpassed speed and flexibility. Over time, the volume of this information can be further condensed via CD ROM technology. Mignon Davis thanked Rose Ella and Ted Spilman for providing this evening’s treats. Visitors were introduced and the meeting was adjourned at 9:20 p.m. Richard G. Robbins, Recording Secretary 955th Regular Meeting—February 1, 1990 The 955th Regular Meeting of the Ento- mological Society of Washington was called to order by President Jeffrey R. Aldrich in the Naturalist Center, National Museum of Natural History, at 8:08 p.m. on | February 1990. Fifteen members and 15 guests were present. Minutes of the January meeting were read and approved. Recording Secretary R. G. Robbins read the names of the following applicants for membership: Ralph P. Eckerlin, Division of Natural Sciences, Northern Virginia Community College, Annandale; Terry A. Wheeler, Department of Environmental Bi- ology, University of Guelph, Ontario, Can- ada; and Jing Zhai, Department of Ento- mology, Virginia Polytechnic Institute and State University, Blacksburg. Mignon Davis circulated a sign-up sheet for volunteers to bring refreshments to our meetings. President Aldrich recounted a recent meeting with Jean Boek, President of the Washington Academy of Sciences, who dis- closed that he is being impeached! Because our Society is a member of the Academy, Dr. Aldrich has decided not to hobnob with that organization’s mutineers, who alleg- 819 edly are headquartered at George Washing- ton University. President Aldrich also an- nounced that he had met with a realty specialist at the Beltsville Agricultural Re- search Center, who informed him that as of this March we will be required to pay a yearly fee of approximately $4.30 per square foot for the storage of back issues of the Proceedings and other Society publications on reservation grounds. Since the shelf space required for our present backlog is estimat- ed to be no less than 200 square feet, we will soon be facing an insufferable annual assessment of almost $900. A discount sale is probably inevitable, but Dr. Aldrich ap- pealed to the membership for additional means of drastically reducing this literary logjam. W. E. Bickley displayed a winsome new book for insect enthusiasts, Ninety-nine Gnats, Nits, and Nibblers, by May R. Ber- enbaum, 1989, University of Illinois Press, Urbana and Chicago, xxi + 254 + 9 pages, illustrated, $29.95/cloth (ISBN 0-252- 01571-1), $9.95/paper (ISBN 0-252- 06027X). R. P. Eckerlin exhibited two specimens of the squirrel flea Orchopeas howardi (Ba- ker) (Siphonaptera: Ceratophyllidae) that had been found on a child. Radioecological studies of the population dynamics of this flea have been conducted in Virginia, where O. howardi has been implicated as a poten- tial vector of sylvan epidemic typhus (Rick- ettsia prowazekii), although experimental attempts to induce transmission by biting have been unsuccessful (Sonenshine et al. 1978, Amer. J. Trop. Med. Hyg. 27: 339- 349: Bozeman et al. 1981, Amer. J. Trop. Med. Hyg. 30: 253-263). Edd Barrows circulated two breathtaking guides to the entomological exhibits at the Staatliches Naturhistorisches Museum in Braunschweig, Federal Republic of Ger- many, Insekten: Begleitheft zum Insekten- saal, by Jiirgen Hevers (ISBN 3-925538-00- 3), and Insekten und Spinnen aus Edelstahl Plastiken, by Hans Jéhne and Jiirgen He- 820 vers (ISBN 3-7682-1393-5). Though lavish- ly illustrated and printed on glossy paper, both guides can only hint at the elegance of the treasures on view in this small museum, which encompasses Taxonomie, Morpholo- gie, Schutztrachten zum Uberleben (cam- ouflage), staatenbildende Insekten (social insects), and Insekten im Wald und der Wasseroberflache (forest and aquatic in- sects). At the time of Dr. Barrows’ visit (Au- gust 1989), the museum also contained a living colony of Honigbienen, all hard at work in the best German tradition. Ed Saugstad exhibited one end of a towel rack that had been made from an Asian hardwood but, apparently unbeknownst to the manufacturer, rendered hollow by the borings of cerambycid larvae. The speaker for the evening was M. Alma Solis, Systematic Entomology Laboratory, U.S. Department of Agriculture, whose talk was entitled “Contributions of Annette Braun, an Early 20th Century Microlepi- dopterist.”” One of the pioneer students of North American leaf-mining Lepidoptera (a vast assemblage of some 16 families com- prising over 335 species), Annette Frances Braun is chiefly remembered for her large monographic works. Her Ph.D. dissertation at the University of Cincinnati in 1912 fo- cused on the evolution of color pattern in the gracilariid genus Lithocolletis Hiibner, but in this and subsequent papers the careful reader will discern seminal concepts of ho- mology and outgroup comparison that pres- aged modern phylogenetic methods. In later life, Miss Braun produced revisions of the North American Elachistidae and Tischer- iidae, as well as the lyonetiid genus Buc- culatrix Zeller; all were published as Mem- oirs of the American Entomological Society. Together with her sister Lucy (a famous bot- anist), she was an early advocate of habitat conservation. A long-overdue tribute to this kind, sedulous scientist will be published by Dr. Solis in the summer 1990 issue of Amer- ican Entomologist (formerly the Bulletin of the Entomological Society of America). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Our numerous visitors were introduced and the meeting was adjourned at 8:55 p.m. Refreshments followed. Richard G. Robbins, Recording Secretary 956th Regular Meeting— March 1, 1990 The 956th Regular Meeting of the Ento- mological Society of Washington was called to order by President Jeffrey R. Aldrich in the Naturalist Center, National Museum of Natural History, at 8 p.m. on 1 March 1990. Fifteen members and 10 guests were pres- ent. Minutes of the February meeting were read and approved. Membership Chairman G. B. White read the names of the following applicants for membership: William A. Bruce, Stephen Hight, and Walter S. Sheppard, all U.S. De- partment of Agriculture, Agricultural Re- search Service, Beneficial Insects Labora- tory, Beltsville, Maryland; Ernie May, Snow Entomological Museum, University of Kansas, Lawrence; Richard L. Orr, College Park, Maryland; Michael P. Parrella, De- partment of Entomology, University of Cal- ifornia, Davis; and Kurt L. Schmude, De- partment of Entomology, University of Wisconsin, Madison. Chairman White also announced the establishment of the Anne M. Wieber Memorial Fund, in honor of our late Custodian. Begun in the closing months of 1989 and spearheaded by Gaye L. Wil- liams, Maryland Department of Agricul- ture, the Fund currently contains about $400, all of which will be used to purchase books on insects for the Naturalist Center, National Museum of Natural History. Cer- tainly, this is a cause worthy of espousal by the entire ESW membership, which greatly benefited from Anne’s selfless dedication. Checks in any amount should be made pay- able to the Naturalist Center and sent to either Chairman White or Ms. Williams. President Aldrich again solicited ideas for reducing the Society’s backlog of unsold Proceedings and other publications. A new storage facility has been located, but it will VOLUME 92, NUMBER 4 accommodate only about half our present holdings. Dr. Aldrich suggested that authors of Memoirs and miscellaneous publications could do the Society a service by personally taking responsibility for surplus stock. James M. Hill, Ecologist, Maryland-Na- tional Capital Park and Planning Commis- sion, exhibited a bag of belostomatids that he had purchased in the frozen food section of an Oriental supermarket in downtown Silver Spring. An aficionado of Thai cook- ery, Mr. Hill observed that these rapacious aquatic bugs are standard fare in Southeast Asia and invited those present to try some samples. A male specimen examined by P. J. Spangler, Department of Entomology, Smithsonian Institution, was assigned to the genus Lethocerus, which includes some of the world’s largest Hemiptera. Several spec- imens bore the calyptostases of hydrach- nidian larvae (Acari: Acariformes: Actine- dida), which commonly parasitize aquatic and semiaquatic insects. President Aldrich recounted his recent trip to Brazil in search of stink bugs. Among his finds were the predaceous pentatomid Stir- etrus erythrocephalus (Lepeletier and Ser- ville) and its curiously similar prey, the chrysomelid beetle Phaedon sp. near con- finis Klug. Striking similarity between pred- ator and prey suggests aggressive mimicry but other interpretations are possible. Spec- imens of both bug and beetle were exhibited to the membership. R. G. Robbins displayed a new dictionary for entomogermanophiles: Worterbuch fiir Veterindrmedizin und Biowissenschaften, compiled by Roy Mack, published in 1988 by Paul Parey, Berlin and Hamburg, 321 pages, $35.00/paper, ISBN 3-489-50516-6. Though not intended as a replacement for the classic De Vries dictionary, which is out of print and will not be republished, this work largely fills the gap by focusing on technical terms in the fields of anatomy, microbiology, physiology, parasitology, pa- thology, pharmacology, toxicology, and sys- tematics. Besides the usual German-English 821 and English-German sections, the dictio- nary contains an appendix of Latin anatom- ical and medical terms together with their German and English equivalents. Two ad- ditional appendices provide the German and English common names of animals (e.g. Bombyx mori, De: Seidenraupe, En: silk- worm) and plants (e.g. Papaver somniferum, De: Schlaf-Mohn, En: opium poppy). In the United States, orders for this invaluable ref- erence should be sent to Paul Parey Scien- tific Publishers, 35-37 West 38th Street, No. 3W, New York, NY 10018. The speaker for the evening was Susan J. Weller, Smithsonian Fellow, Department of Entomology, whose talk was entitled “Why are Notodontid Genitalia so Variable—Sex- ual Selection?” Males of Neotropical no- todontids belonging to the tribe Nystaleini possess a number of novel courtship struc- tures, the most spectacular of which is a more or less elaborate modification of the external genitalia: the lower portion of the valve, the sacculus, contains glandular ma- terial, and androconial scales are attached to its pleated, membranous base. In the rel- atively large Nystalea clade, which includes nearly 200 species, all males possess these valve scent organs as well as secondary scent organs on either the legs or abdomen. By contrast, the Dasylophia clade comprises just 55 species, only one genus possesses an ab- dominal courtship structure, and the valve scent organs are reduced or lost. Such dif- ferences are consistent with the theory of sexual selection via female choice. In the Nystalea clade, chemical communication may allow for rapid divergence of signals driven by female choice, thereby promoting speciation. However, in the Dasylophia clade, mechanical communication through genitalic contact is the only means available for females to assess different males. Either female choice does not occur in the Dasy- lophia clade or morphological structures cannot evolve as quickly as chemical sig- nals. Dr. Weller illustrated her fluent pre- sentation with slides of some striking no- 822 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON todontid larvae, including the lobster moth, Stauropus fagi Linnaeus, and a spider mim- ic, Cnethodonta grisescens Staudinger. She also exhibited the eminently perusable text Sexual Selection and Animal Genitalia, by William G. Eberhard, Harvard University Press, cloth edition 1985, 288 pages, $26.00, ISBN 0-674-80283-7; paper edition 1988, 256 pages, $14.95, ISBN 0-674-80284-5. President Aldrich thanked Bill Bickley and Edd Barrows for supplying this evening’s genteel repast of Girl Scout cookies and soft drinks. Visitors were introduced and the meeting was adjourned at 9 p.m. Richard G. Robbins, Recording Secretary 957th Regular Meeting—April 5, 1990 The 957th Regular Meeting of the Ento- mological Society of Washington was called to order by President Jeffrey R. Aldrich in the Naturalist Center, National Museum of Natural History, at 8 p.m. on 5 April 1990. Sixteen members and nine guests were pres- ent. Minutes of the March meeting were read and approved. President Aldrich read the name of to- night’s guest speaker (vide infra), who has applied for membership. President-Elect David R. Smith an- nounced that the annual joint banquet of the Entomological Society of Washington, Pest Science Society of Washington, and Maryland Entomological Society will be held on the evening of 7 June in the Associates Court, National Museum of Natural His- tory. This year’s Master of Ceremonies will be Douglas W. S. Sutherland, Entomologist, U.S. Environmental Protection Agency. Our after-dinner speaker will be David A. Nick- le, Research Entomologist, Systematic En- tomology Laboratory, U.S. Department of Agriculture, who will regale us with an au- dio-visual presentation entitled “Hide and Sing in the Rainforest: the Katydids of the Peruvian Amazon.” President Aldrich reminded the mem- bership that our Regular Meeting for May will be held in the National Visitors Center (or “Log Lodge’”’) at the Beltsville Agricul- tural Research Center. He also announced that on 11 April the Center for Agricultural Biotechnology and the Department of En- tomology, University of Maryland, College Park, will co-sponsor a special seminar en- titled ““Mating Systems in Ghost Moths (Hepialidae) and other Primitive Lepidop- tera: Role Reversal, Sex Scaling, and Con- tingency Analyses,” presented by David L. Wagner, Department of Ecology and Evo- lutionary Biology, University of Connecti- cut, Storrs. Dr. Wagner’s data suggest that basal lineages of Lepidoptera and Trichop- tera possessed a female-released, long-range sex attractant and that the unusual male- based calling systems found in the Hepiali- dae represent one or more special deriva- tions. The morphological diversity and tax- onomic distribution of male scent scales (androconia) have been examined in several clades and appear to be the most rapidly evolving adult structures. Explanations of this evolutionary lability will be explored, with emphasis on the use of statistical cor- relation analyses, which treat species as in- dependent data points. At the request of Bill Bickley, Rich Rob- bins read an announcement from Towson State University, which will host the 51st annual meeting of the Association of South- eastern Biologists (ASB), 18-21 April, in Baltimore. Societies meeting in conjunction with the ASB include Beta Beta Beta; the Botanical Society of America, Southeastern Section; the Ecological Society of America, Southeastern and Washington, D.C., Chap- ters; the Society of Wetlands Scientists, Northeastern and South Atlantic Chapters; and the Southern Appalachian Botanical Club. Field trips are planned to such nearby attractions as Fort McHenry, the McCor- mick Spice Company, and the National Aquarium in Baltimore, as well as to the Beltsville Agricultural Research Center, the Patuxent Wildlife Research Center, Soldiers Delight Serpentine Area, the Smithsonian VOLUME 92, NUMBER 4 Environmental Research Center on the Rhode River south of Annapolis, and Long- wood Gardens in southeastern Pennsylva- nia. Registration, paper and poster presen- tations, exhibits, and general sessions will be held at the Lord Baltimore Radisson Pla- za, about 10 miles south of the Towson State campus. ASB membership is not required for registration. D. R. Smith exhibited a lavish new pub- lication on what are probably the world’s most ubiquitous insects, The Ants, by Bert HGlldobler and Edward O. Wilson, 1990, published by the Belknap Press of Harvard University Press, Cambridge, Massachu- setts, xiv + 732 pages, ISBN 0-674-04075- 9, $65.00/cloth (alkaline paper). A large page format, numerous full-color paintings, and an illustrated key to the approximately 300 recognized ant genera make this authori- tative but engrossing volume a “must buy” for any entomologist, regardless of special- ization. T. J. Spilman displayed an attractive new paperback, Nymphs of the Sahelian Grass- hoppers: An Illustrated Guide, by G. B. Po- pov, 1989, published by the Overseas De- velopment Natural Resources Institute, Chatham, Kent, United Kingdom, with partial funding from the U.S. Agency for International Development, v + 158 pages, ISBN 0-85954-264-5. Though small enough to slip into a pocket, this book succeeds in illustrating with color drawings and pho- tographs one or more instars of some 78 species of “hoppers” from the vast sub-Sa- haran semidesert known as the Sahel. As- sociated symbols provide instant summa- ries of life cycles, food and habitat preferences, maximum number of genera- tions per year, number of instars by sex, gregariousness, infraspecific variation and, of course, economic importance. Common names appear in both English and French, a reflection of this region’s equally colorful colonial history. President Aldrich projected slides of the spined soldier bug, Podisus maculiventris 823 (Say) (Hemiptera: Pentatomidae), an efh- cient springtime predator of lepidopterous larvae. He also distributed samples of this species’ aggregative pheromone, formulated into small, gumball-like spheres for shotgun distribution. The speaker for the evening was Timothy P. Friedlander, Center for Agricultural Bio- technology, University of Maryland, whose talk was entitled ““Moths and Molecules: Reconstructing a Phylogeny from Nucleic Acid Sequence Data.” Molecular system- atics is a useful tool in taxonomic situations where there are many species but few mor- phological characters. While at Louisiana State University, Dr. Friedlander attempted to construct a phylogeny of the superfami- lies of higher Lepidoptera, infraorder Di- trysia, by comparing ribosomal RNA (rRNA) sequence data among 35 represen- tative species. Taxonomic characters from rRNA sequences were shown to be various kinds of nucleotide base substitutions, in- cluding transitions, transversions, and in- sertion/deletion events. Sequence charac- ters were evaluated for homology and independence, after alignment, by looking at secondary structure, and relative fre- quency classes were assessed. The software package Phylogenetic Analysis Using Par- simony (PAUP) was chosen for evolution- ary hypothesis construction. Characters of the less frequent character classes were an- alyzed, these being more appropriate to such inference. Taxa were chosen with the goal of estimating the hypothetical ancestral character states for each independent su- perfamily; this method reduces both the number of taxa in a computer run and the amount of homoplasy—without reducing the number of characters. Dr. Friedlander’s molecular analysis of the Ditrysia yielded some unexpected results: butterflies did not group with other macrolepidopterans, while lasiocampids grouped strongly with cossids and castniids rather than with other bom- bycoids. Edd Barrows projected a series of slides 824 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from last year’s annual banquet, including and horseradish dip.” Visitors were intro- one of the irrepressible Doug Sutherland duced and the meeting was adjourned at 9 sporting a plexippian bow tie. p.m. Davis thanked his daughter Marisa : : : Don ay tha ed Sue ugh a Richard G. Robbins, Recording Secretary for providing this evening’s “onion bread PROCEEDINGS of the PROC. ENTOMOL. SOC. WASH. 92(4), 1990, pp. 825-831 ENTOMOLOGICAL SOCIETY President President-Elect Recording Secretary Corresponding Secretary Treasurer Program Chairman Membership Chairman Custodian Editor OF WASHINGTON Volume 92 OFFICERS FOR THE YEAR 1990 Jeffrey R. Aldrich David R. Smith Richard G. Robbins Hollis B. Williams Norman E. Woodley Gary Stech Geoffrey B. White James B. Stribling Robert D. Gordon Past-President Published by The Society WASHINGTON, D.C. 1990 TABLE OF CONTENTS, VOLUME 92 ARTICLES ADLER, PETER H.—Two new species of black flies (Diptera: Simuliidae) from North America BAKER, GERALD T. and SONNY B. RAMASWAMY. —Tarsal and ovipositor sensilla of Heliothis virescens and H. subflexa (Lepidoptera: Noctuidae) . Een aes BAO, NONGGANG and WILLIAM H. ROBINSON. — Morphology and mating ‘conheuration of genitalia of the oriental cockroach, Blatta orientalis L. (Blattodea: Blattidae) ...... BARBER, K. N.—See HERATY, J. M. BARBOSA, PEDRO—See VEGA, FERNANDO E. BARR, C. B.—See MORSE, J. C. F. Christian Thompson 826 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BATRA, S. W. T.—Bionomics of Evylaeus comagenensis (Knerer and Atwood) (Halictidae), a facultatively polygynous, univoltine, boreal Halictine bee ..................-5.....20.. BECKER, V.O.—See HODGES, R. W. BERNARD, ERNEST C.—See MUEGGE, MARK A. BOE, A.—See McDANIEL, B. BORKENT, A.—A revision of the Nearctic species of Dicerura Kieffer (Diptera: Cecidomyiidae) BOTTORFF, R. L., 8S. W. SZCYTKO, and A. W. KNIGHT.—Descriptions of a new spe- cies and three incompletely known species of western Nearctic /soperla (Plecoptera: Per- |korc b(s 1) ee oh ee Oi So Se SR a BO on ih A DG anidiGn ocean Omlann BOWLES, D. E.— See MATHIS, M. L. BROE, R.—See SANDS, D. P. A. BROWN, B. V.—Triphleba vitrinervis (Malloch), an unrecognized species of Crinophleba Borg- meier(Dipteras Phoridae). .¢-- hase ero er eee fee a alb a fener steiaeiehalans Srexeuete ets CARROLL, L. E.—See STECK, G. J. CELEDONIO-HURTADO, H.—See STECK, G. J. CHADEE, DAVE D.—See LINLEY, JOHN R. COOPER, KENNETH W.-— Likely causes and explanation of probable atavism in somatically mosaic fly from a wild population (Diptera, Asilidae, Nannocyrtopogon minutus) ........ CROTHER, BRIAN I.—See LUYKX, PETER DAVIDSON, R. L. and H. J. LEE, Jr.—Distribution and habitat of Dyschirius campicola Lindroth (Coleoptera: Carabidae) with new state records for Ohio and Illinois, first records Eastiot the Mississippl RIVED: Yrco.sckcpeqece cccccie-ate snoiel amar ates ol evarstsueeenaees Glens eerie eet eee DAVIS, D. R.—Neotropical Microlepidoptera XXIII. First report of the family Eriocottidae from the New World, with descriptions of new taxa ...........22...0000-2- ec cee eee enes DAVIS, D. R.—First record of a bagworm moth from Hawaii: description and introduction of Brachycyttarus griseus De Joannis (Lepidoptera: Psychidae) ......................5-- DAVIS, DONALD R.—Three new species of Acrolophus from the southeastern United States with remarks on the status of the family Acrolophidae (Lepidoptera: Tineoidea) ......... DAVIS, DONALD R. and JORGE E. PENA—Biology and morphology of the banana moth, Opogona sacchari (Bojer), and its introduction into Florida (Lepidoptera: Tineidae) ...... DIETRICH, C. H. and S. H. MCKAMEY.—Three new idiocerine leafhoppers (Homoptera: Cicadellidae) from Guyana with notes on ant-mutualism and subsociality ............... DUFFIELD, R. M. and C. H. NELSON.—Seasonal emergence patterns and diversity of Ple- coptera on Big Hunting Creek, Maryland, with a checklist of the stoneflies of Maryland DUNCAN, R. W.—See GAGNE, R. J. EASTOP V. F. and D. J. VOEGTLIN.—Taxonomic notes on some North American aphids EISENBERG, R. M. and L. E. HURD.—Egg dispersion in two species of praying mantids (MantodeaMantidae)) 0. fortes ee seicie reer o ee aoe tesco ete teket reaps es eer poe Tare ELBERSON, LESLIE—See KIRIAC, ION EPSTEIN, MARC E. and SCOTT E. MILLER.—Systematics of the West Indian moth genus Heuretes Grote and Robinson (Lepidoptera: Limacodidae) .........................-.. FREIDBERG, A. and W. N. MATHIS.—A new species of Craspedoxantha and a revised phylogeny, for the/genus;(Diptera Tephritidae)) 22-2 nye aieniaie ie iorsiototereiee ste eke FREYTAG, PAUL H.—A new genus Nancyana and nine new species with a review of the related genus Rhogosana (Homoptera: Cicadellidae) ...................000000 0 ee eee GAGNE, R. J. and R. W. DUNCAN.—A new species of Cecidomyiidae (Diptera) damaging shoot tips of yellow cypress, Chamaecyparis nootkatensis, and a new genus for two gall midgesonl @upressaCeaep cee seer ee ek = een ee ee etelen tee rete er keer GAGNE, RAYMOND J. and GWENDOLYN L. WARING.—The Asphondylia (Decidomyi- idae: Diptera) of creosote bush (Larrea tridentata) in North America .................-- GILES, FRANCIS E.—See WIRTH, WILLIS W. 571 286 271 373 VOLUME 92, NUMBER 4 827 GOEDEN, RICHARD D. and DAVID HEADRICK.—Notes on the biology and immature stages of Stenopa affinis Quisenberry (Diptera: Tephridae) .................00.202000000. 641 GOEDEN, RICHARD D.—See HEADRICK, DAVID GRACE, J. KENNETH.—Effect of antioxidants on eastern subterranean termite (Isoptera: Rhinotermitidae) orientation to fungal extract... 22... eee 773 GRISSELL, E. E. and M. E. SCHAUFF.—A synopsis of the seed-feeding genus Bephratelloides (GhaleidoideastEunytomidae) i ecac ce cc eyecare eccicrectstce ates ce stn es enn eee eee 77 GROGAN, W. L., Jr. and W. W. WIRTH. as new species of the: minute predaceous maidee genus Nannohelea from Sri Lanka (Diptera: Ceratopogonidae) ..................... ... 347 GROGAN, W. L., Jr.—See SPINELLI, G. R. GRUBER, FRANCIS—See KIRIAC, ION GUILLEN-AGUILAR, J.—See STECK, G. J. HALBERT, SUSAN E. and KEITH S. PIKE—An Asian elm aphid (Homoptera: Aphididae) New toi NOntHVAMEFICa sa ace cre eee sae ee cere Sas eater ars cle Cn. cara ee es 2 HALBERT, SUSAN—See KIRIAC, ION HALSTEAD, J. A.—Review of Haltichella Spinola in the Nearctic Region (Hymenoptera: Ghalcidid ae) ira ae era a ere eee ere ere eee Me reac a 153 HALSTEAD, JEFFREY A.— Revision of Hockeria Walker in the Nearctic region “with de- scriptions of males and five new species (Hymenoptera: Chalcididae) ................... 619 HEADRICK, DAVID and RICHARD D. GOEDEN.— Resource utilization by larvae of Para- cantha gentilis (Diptera: Tephritidae) in capitula of Cirsium californicum and C. proteanum (Asteraceae) in‘southern California .:....2.....22-.405c--ssseeee-- ee : Cmnaccces, eoul2 HEADRICK, DAVID—See GOEDEN, RICHARD D. HENRY, THOMAS J. and DANIEL J. HILBURN.—An annotated list of the true bugs (Het- ELOptera) Ot Bena sa Sap eget ts eee eee eee ee Genes Mens scare), es sik cvadepean digs atte lela 675 HERATY, J. M. and K. N. BARBER.—Biology of Obeza floridana (Ashmead) and Pseudo- chalcura gibbosa (Provancher) (Hymenoptera: Eucharitidae) ........................-.. 248 HESPENHEIDE, HENRY A.—New species of Buprestidae (Coleoptera) fom the Doniici IREPUbLICA Pee ee metre meet Soe serene freee ne Syed ere aken oer Cicadellidae] (= cmemsemecr rere ee ieee eestor rer tote eeveeniate reketere tare renecratst 461 VOEGTLIN, D. J.—See EASTOP, V. F. WALTZ, R. D.—Baetis jesmondensis McDunnough, a New Junior Synonym of Baetis tricau- datus' Dodds (EphemeropterasBactidae) anv isy irate) slover-lclotclatolateieie cleteretatetaiee erate 138 WARING, GWENDOLYN L.—See GAGNE, RAYMOND J. WEAVER, J. S., I1].—Two new synonyms in Rhyacophilidae (Trichoptera) ............... 358 WEAVER, J. S., II and J. L. SYKORA.—A new synonym in Hydroptilidae (Trichoptera) .. 360 WELCH, JUDITH L.—See KONDRATIEFF, BORIS C. WHITFIELD, J. B.—Phylogenetic review of the Stiropius group of genera (Hymenoptera: Braconidae, Rogadinae) with description of a new neotropical genus ..........-.....-...55 36 WILLIAMS, M. L.—See SHEFFER, B. J. WIRTH, W. W.—The biting midges of Aldabra Atoll, Indian Ocean (Diptera: Ceratopogonidae) 230 WIRTH, WILLIS W. and FRANCIS E. GILES.—New species and records of predaceous midges; from) Fiji (Diptera: Geratopogonidae)) 2. 75 etre siete aeleh eters fiedtets era elersetretedt os 444 WIRTH, W. W.—See GROGAN, W. L., Jr. WOOD, T. K.—See OLMSTEAD, K. L. YODER, KAREN M. and WILLIAM H. ROBINSON.—Seasonal abundance and habits of the boxelder bug, Boisea trivittata (Say), in an urban environment ...................--- 802 YOUNG, D. K. and J. B. STRIBLING.—Systematics of the North American Cyphon collaris species complex with the description of a new species (Coleoptera: Scirtidae) ............ 194 YOUNG, DANIEL K.—See STRIBLING, JAMES B. NOTES ALDRICH, JEFFREY R.—Dispersal of the southern green stink bug, Nezara viridula (L.) (Heteroptera: Pentatomidae); by hurricane Hugo’ 2222-322. cae els seo este einen stele elles Test) DYTE, .C. E.—The. type. locality of Sciapus pressipes Parent (Diptera: .Dolichopodidae) 584 HUANG, X. P. and T. P. MACK.—Examination of some sensory organs on the head of last instar larvae of the lesser cornstalk borer, E/asmopalpus lignosellus (Zeller) (Lepidoptera: lagriicht) hes ceoopcuospaddao Sra chor ob hoch nmuoconnmods bhobboodds Ap daooONCOdosOaNn aC 736 MACK, T. P.—See HUANG, X. P. MYLES, TIMOTHY G.— Coptotermes crassus Ping preoccupied by C. crassus Snyder renamed G./pingi Tsoptera: Rhinotermitidae)) << cies sate sels sole eraveelate eyevel letede ele lately Yel l= fel 1-Yoree tated 813 RIDDICK, ERIC W.—Andrena macra Mitchell (Hymenoptera: Andrenidae) overwinter and delay/spring:emergence int VaAnginiagerys ere ters tere teeter lot vale eted niet dete te eked okletve fare eete tee ret 771 SMITH, DAVID R.—Liliacina diversipes (Kirby) (Hymenoptera: Tenthredinidae), a sawfly genus.and’speciesinew toi the Wnited'States! = 222 2.1 ni cee yaciele eters -m eleteelaleioie terme tore 812 WELCH, KENNETH A.—First distributional records of Cimexopsis nyctalis list (Hemiptera: Gimicidae)iin’ Connecticut). seen rec renee cease ieee ose ei ere ei crete ateieer het ie 811 VOLUME 92, NUMBER 4 831 BOOK REVIEWS BOLDT, E.—The Plant-Feeding Gall Midges of North America ..................... scsi SOD CHANDLER, L. D.—Cotton Insect Pests and Their Management .................... we) 16 CHANDLER, L. D.—Crop and Plant Protection: The Practical Foundation .......... er its) ELZINGA, R. J:-—Jnsect Spiracillar SyStOMs. «oi ic ccc cae e es cece asec eee seseeens 362 FREIDBERG, AMNON.—4 Guide to the Breeding Habits and Immature States of Diptera Gyclorrhaphaees eons ee ee ee PPE E A rteRaton ARTE RAEI ae 589 FUNK, V. A.—Novel Aspects of Insect-Plant Titerantions A eo ta ee nee 171 GAGNE, RAYMOND J.—Bird Blow Flies (Protocalliphora) in North America (Diptera: Cal- liphoridae) with Notes on the Palearctic Species 0.2.00... ccc ccc cee ee 814 JACOBSON, M.—Cyanide Compounds in Biology ..... 0.000000 ooo oc ce eee 1:72 LACEY, L. A.—Combating Resistance Xenobiotics: Biological and Chemical Approaches ... 169 LAMP, W. E.—Plant Stress—Insect Interactions ..... AE Shick cre ROT 361 STEYSKAL, G. C.—The Torre-Bueno Glossary of Eniomalosy ee Sic gute Barware LOLS STOEEOLANOMIOHIN Gy, Jr-—Insect Flight 2.5.2 neue ss vecsss acess coms seesccenvas 816 TALLAMY, DOUGLAS W.—/nsect-Plant Interactions ............ Eesd Awad eee Oa OBITUARIES WHITE, GEOFFREY B. and RALPH E. WEBB.—Obituary. Anne Marie Wieber 1958-1989 585 MISCELLANEOUS INSTRUCTIONS TO AUTHORS ..... Se ee ar eee ee OF ee ecenent arene ate ... 166 SOCIETY MEETINGS AND REPORTS TO OFFICERS FOR 1989 .... saree ove, oO) SOGIETIVZMEE MING Sita ne oti aracave ciate Griesoiyeieiars ane ehehaveucteiaia gee Se. oyb.a0.4 dod andi edie @ Sites 818 vy ee i ee. > PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS @ynipidiGallsiof theEasterm United’ States; by Lewis)/H. Weld 22 2 ee GynipidiGallsiofithe Southwest. by MeewiSjh) Weld ee en Soe ee eee Bothspaperston Gypipidigalls ese See a ee ee 2 ee Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. Gea OTe ec ee ey Oe ier nr, ee ee A ees one cana Nee eS Le eh A Short History of the Entomological Society of Washington, by Ashley B. Gurney Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. SS LSS kc a are ae ce See Ce ee A 1 a A I Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. Steys kal Seem ese te arsine See A eee eh ese ee ee See ee ee Lee MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939... No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. CGY jo) oe SY eae Se ee ee ee ee Se ee 2 eS ee ee ee No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. 2S SAD a O AD es ke nt Ne Te A ee ee ee es eee ES ee 4 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952000. No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. SSNPS a OEP a ON Ie ae ee SO doe, Pe ee ee Ok PER eee SRG N EP Been No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Makahasieg23 Oippiyel 969 et be Laka ter te, SEA a ee Ree Ye A So Ee No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. NOFA waste eB Ee a IY Dae oh al Pe Ee Sl A ee Ae ee eS aS No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera- topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 oceans No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 DD Se OSilvests eee: se See eS ee ee reels Oe eS eS No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982... No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. NCO YS3 hoes S ee A e, i RRe e P A ee c ie ica ee ae No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. GREDD sO OA ye see ee eee eT ee pee hae MOT Cee Se ee) Ee Dae ae 2 ae eS No. 13. .An Identification Manual for the North American Genera of the Family Braconidae (Hy- menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 0. 1.50 2.00 15.00 11.00 12.00 10.00 11.00 18.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume to non-members and $13.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560. CONTENTS (Continued from front cover) McCAFFERTY, W. P.—A new species of Stenonema (Ephemeroptera: Heptageniidae) from North Carolina McDANIEL, B. and A. BOE—A new species and distribution record for the genus Caeculus Dufour (Acari: Caeculidae) from South Dakota MUEGGE, MARK A. and ERNEST C. BERNARD—Two new species of Metajapyx (Diplura: Japygidae) from Tennessee NAITO, TIKAHIKO-—The tribe Strongylogasterini (Hymenoptera: Tenthredinidae) from Tai- wan ROSKAM J. C. and H. NADEL—Redescription and immature stages of Ficiomyia perarticulata (Diptera: Cecidomyiidae), a gall midge inhabiting syconia of Ficus citrifolia ............ ROSSI, ANTHONY M. and DONALD R. STRONG—A new species of Asphondylia (Diptera: Cecidomyiidae) on Borrichia (Asteraceae) from Florida STRIBLING, JAMES B. and DANIEL K. YOUNG—Descriptions of the larva and pupa of Flavohelodes thoracica (Guérin-Méneville) with notes on aphytotelma association (Coleop- tera: Scirtidae) YODER, KAREN M. and WILLIAM H. ROBINSON—Seasonal abundance and habits of the boxelder bug, Boisea trivittata (Say), in an urban environment NOTES ALDRICH, JEFFREY R.—Dispersal of the southern green stink bug, Nezara viridula (L.) Het- eroptera: Pentatomidae); by hurnicane Hugo: 22.5 9. edccasucem ee oe econ oan HUANG, X. P. and T. P. MACK—Examination of some sensory organs on the head of last instar larvae of the lesser cornstalk borer, E/asmopalpus lignosellus (Zeller) (Lepidoptera: Pyralidae).". 04 aw Ske cid tec ee AS Me. Se A Sete oh Ae cet ia Seater eee ree MYLES, TIMOTHY G.—Coptotermes crassus Ping preoccupied by C. crassus Snyder renamed C: pingt (soptera: Rhinotermintidae), 4/5... 5 6 ie os. = Pela neice aces a seg alone hs ore peices eee RIDDICK, ERIC W.—Andrena macra Mitchell (Hymenoptera: Andrenidae) overwinter and delay;spring emergence an) Virpinia. 3 ior. cicero > te age ie vte= ala ot =) har perma SMITH, DAVID R.— Liliacina diversipes (Kirby) (Hymenoptera: Tenthredinidae), a sawfly genus andi species HEwto, them mited States Re fast oes = yale} ale yt Rigo » Sole Cota eee eee WELCH, KENNETH A.—First distributional records of Cimexopsis nyctalis List (Hemiptera: Gimicidae)iin\Gonnecticuts. 225 5.. nls lge «2-0-0 ba 8 a one Ac ictaya ee ates ei eet cater nicNe Cer eee BOOK REVIEWS GAGNE, RAYMOND J.—Bird Blow Flies (Protocalliphora) in North America (Diptera: Calli- phoridae) with Notes on the Palearctic Speci€S ........ 0000000 c cece ccc cece eee neneees STOFFOLANOS JOHN GIR: =Insect Flight: 1.427 20 ke wee etd = acts ak ee ee eerie) SOCIETY: MEETINGS) faved acre s.<5 os eects om Hele ike See teeta aes ieeiele eeice Ret eerre TABLE. OPIGONDEIND Ss o5 fei 5 sie dpoeis le cmcics ecu Mees Woe coanoge ey TeV SIe (alist te eer cet ene eee eae cree 760 716 793 79 778 732 765 802 ie 736 813 771 812 NS [2 3] > YY J > | a reas a = | See \S \ a> rs) ae “Ge 2 xm £ = % yay a 2 insist al ‘fl m ONDE Z unis mt ITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3S1IYVYUEIT LIBRARIES Zz Re 7) z 77) z eee 2) R . =| ; A Pe te S YS, =) 1h fg 5 z UY fy ro) &: ”. = 2 NRO 2 Zs D bY D WS & < 0 GOAG =e (e} Wid x= \. fe) = z “yp eS zYZY = NS z = . > = > = ee 7) i Fe ” 7-2 7) ee INOSHLINS S3IYVYNSIT LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLII ee ”n = S = Hf S ier Aa B Se wee - @ WY Ss oc = x. YY YA = Ls = XY cc =| oc Ge, S cc S = iS) Se re) = 3 SS eel 2 a} aes = Zz. ES IITHSONIAN INSTITUTION | NOILNLILSNI NVINOSHLIWS (S31 YVaal LIBRARIES SMITHSONI. ° mz z = ro) ro - ro) Y w —_— 5 —_ ow 4 — - = dy 2 5 Ng 5 5 5 gy = = SS = & E bi LY, : b SS 2 Bo o z sk at Oo z (INOSHLINS S3SIYVYEIT LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLI! SMITHSONIAN wtt f % j Li, , Ye NVINOSHLINS S3IYVUaI wn ~~ ae w oS = < = < I x = re) Se AS é z. NS E Ae. = 2 . ee 3 AITHSONIAN INSTITUTION NOILALILSNI_NVINOSHLINS | S3 1YVYE!IT LIBRARIES SMITHSONI. = no = o s, = a = ec = z A = eS = = < = a a a =| es re) = (e) —s ro) a. Fs aj ee wall —_ Sy INOSHLINS S3IYVYUEIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLII : z S Zz iz z l= f fe) = S) wo ° = NS Ley be) = me] iE D wy =) x = = > S >: S = Be) = son = eS) = = _ = ape a z a z a AITHSONIAN INSTITUTION NOMIMLIESNI NVINOSHLINS | S34 | uveai tie! BRARI ES ,SMITHSONI = 2. = = = ya ee 2S 5 2 ‘2 NS 5 D9 Xs Zc re) ce ORENNG Be 2S 2 2 g ZR 8 fs ; $ = ad E Sete = . > G = > = bet y= ”o a z ” ae ” Sees JVINOSHLINS S3IYVYUSIT LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLI Jp) = > = Sache . Ww = Ww ; : - Ww = = Ho Toad See 4, 4H oe — ee 5 “PUY 5 2 5 e = = Uyty Ss e 5 = 3 eo 3 = 3 Sey =a = 2 ay zZ S MITHSONIAN INSTITUTION _ NOILALILSNI_NWINOSHLIWS 24 lYvudgd 7 tl BRARI ESS MHSOM! 2 E > We 5 2 = Y x = ay NKAY EF 2 = YY p = D = Ys a = a : wo = wo ; z Ww z VINOSHLINS SSIYVYAIT_ LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLI ww Yj Yr HSONIAN AHSONIAN ISHLINS HSONIAN TPO oa VASA ISHLIWS ‘G OSHLIWS an Ttfh 4 ns = wl - na Vs WS: n > -ixZ# MG > Eee > MU 2 > 2 EG a \ Aiea rs) SS = Es) ae = Yfy = Os of 6 prs b ae m +2) ti m Qn OC “ m 12) m ra) = wo = wo Sas wo ITUTION NOILALILSNI NVINOSHLINS S31YVYE!IT LIBRARIES SMITHSONIAN INSTITUTION N Re wo z no Zz oe ” z : n x a = “Ss = af = \N \ = = > = . 2F = > ha aed 77) Zz 75) - 2 an 2 y4v¥uYgia LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI L ) = es re - “a ri a ul L418 ul Ss ws & a ta ye 3 x =t = = _(\7 £4 4 = t c iY i = = = ps c = a YO > S = = = fe) £2 fe) = 5 2 re) 2 =) z = a asl 2 ITUTION NOILNLILSNI NVINOSHLINS S3IYVYEIT LIBRARIES SMITHSONIAN INSTITUTION N = a amie: : a a 2 e av \ = a ra es = > =) 5 - S = 2 SSS E = ra = = eB | = 2 NS Ie a = z = [¢p) nS © w z . 2 p z R z 4VYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S3ZIYVYEIT L' fer ITUTION NOILALILSNI_ NVINOSHLINS S3ZIYVYEIT LIBRARIES SMITHSONIAN MG NVINOSHLIWS NVINOSHLIWS NVINOSHLIWS SMITHSONIAN SMITHSONIAN SMITHSONIAN INSTITUTION N LIBRARIES SMITHSONIAN NOILNLILSNI at 4 LIB RAR eS) LIBRARIES NOILNLILSNI NOILALILSNI LIBRARIES YWYdIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYVYEIT L c z : c : c Ee a = a o — mee) J w st Pl = a = 2 E s > es > = > res > 2 Ee = E = e e (ep) D 2 Oo z 4 . Z D ITUTION NOILONLILSNI NVINOSHLINS S3I1YVYSEIT LIBRARIES SMITHSONIAN INSTITUTION N a ov z o z oe oO a) ee, wo re = < = cS &e = x =< ®. = RS = z = “2 WS 3S z Ko 5 * a 5 (eo) as (e) SS Sie x as oO RSs SY s¥e BS Hw Za Ry 2 / = > = . FS = . > ee a z a Ae a . 5 yvudit LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IuYvYuRaIT L ZS es = = i) = = WwW = WwW 2 Ta Ww = g = ih jp 3 = 2 WX = g a = t/ > E E > QAre zi EY ane = ” m yy 2 m os m g = 7) 3 = wo = wo = yVYUdIT LIBRAKIES SMITHSONIAN INSTITUTION NOILOLILSNI NVINOSHLINS S3IYVESIT_L <0 wOr> S 4) fe x! SONIAN SHLIWS SONIAN SHLIWS SONIAN SHLIWS SONIAN iy py. re nn eaane ONIAN INSTITUTION sIES ILIA 88 00908 0730