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October 1986

Taxonomy of the Adults of the Robineau- Desvoidy(Diptera, )

G. C. Steyskal National Museum of Natural History, Washington, DC

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Steyskal, G. C., " of the Adults of the Genus Strauzia Robineau-Desvoidy(Diptera, Tephritidae)" (1986). Insecta Mundi. 529. https://digitalcommons.unl.edu/insectamundi/529

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Vol. 1, no. 3, October 1986 INSECTA MUNDI 101

Taxonomy of the Adults of the Genus Strauzia Robineau-Desvoidy (Diptera, Tephrltidae)

G. C. Steyskal Systematic Entomology Laboratory, IIBIII Agricultural Research Services, USDA c/o National Museum of Natural History--NHB 168 Washington, DC 20560

INTRODUCTION the inadvisability of erecting any of the members of this group to the importance of The recently increased importance of species, though specimens may be selected sunflowers (Hellanthus annuus Limaeus) which present a vivid contrast. It is as a crop has brought with it an increased even almost impossible to characterize interest in the sunflower maggot, a distinct varieties. The most constant long-recognized conmon name for Strauzla character for such differentiation lies longlpemls (Wiedemann). The of apparently in the relative size of the this species bores downward in the stem frontal bristles in the males. In regard and feeds upon the pith of sunflowers. to the characters of the wing pattern Until recently the genus Strauzla was there are not only gradations between considered to include only the seemingly distinct varieties, but even the type-species S. longipennis , with 7 two wings of an individual show varieties (Foote, 1965). It is now known considerable differences. Characters that some of these varieties should be derived from the picture of the thorax are classed as distinct species, differing also very unstable. *' Much more material morphologically and biologically. has been accumulated since Snow*s time, All species of Strauzia whose biology and the use of the terminalia of both is known to any extent bore in the stems sexes has become comnonplace, but his of various of the family statement is still true to a great (Compositae) during the larval stage and extent. More will be said on this matter feed only on the pith. This mode of under S. longipennis. feeding apparently does not harm the The species of Strauzla are a180 very , although it has been stated (Brink, similar, in general appearance as well as 1922) that a reduction in seed production in detailed morphology, to species of the is caused thereby, and that the stems are genera Edela and Wyolefa. The weakened sufficiently for high winds, American leaf miner of parsnip and related frequent in the Middle States where the plants, fratrla (Loew), and its greater amount of sunflowers is grown. close European relative, the celery leaf This comes at the crucial time when the miner E. heraclei (Limaeus) are so seed heads are ripening, causing similar in wing pattern to females of considerable loss by destroying plants Strauzla longlponnls or to either sex of which, if uninfected, may have survived S. perfecta that they may easily be (Caesar, 1924; Westdal and Barrett, 1960; taken for one another. In fact, the Schulze, personal comrmnication). Other seminal receptacles of Euleia fratrla deleterious effects, such as increased were described by Sturtevant (1925-1926) attack by other organisms, have also been as those of S. longfpenn~s (see attributed to the work of the sunflower Steyskal, 1972). Characters to separate maggot. Strauzla from its closest relatives will The genus Strauzia is restricted to be given under the generic heading. North America. The species are very The species now known as GgmOCdrena similar in general appearance, as may be diffusa (Snow) and originally proposed in inferred from the fact that they at one the genus Oedicarena was for a time time were all treated as varieties of one placed in Strauzia. It somswhat species. This was sumnarized in 1894 by resembles Straula longlpennls and ha8 Snow, who stated, "The material at hand, been found to develop in the seed heads of though not large, is sufficient to show sunflowers. INSECTA MUNDI Val. 1, no. 3, October 1986 Genus Strauzia Robinew-Desvoldy setae, as well as its obvious relationships with Euleia and Hyoleja. Strauzia Robineau-Desvoidy, 1830: 718. The following combination of characters Type-species , inermis R.-D. , by will distinguish Strauzia among all designation of Coquillett, 1910: 609 (= North American Tephritidae: Humeral setae longipennis [Wiedemann]); Agassiz and present ; basal cubital cell closed by Loew, 1846: 37 (Strauzia, derived from strongly bent vein, cell elongated at l*Strauss-Durckheim"); Agassiz, 1846: 354 lower apex; dorsocentral setae anterior to (Strauzia, with note on use of halfway point between supra-alar and Straussia in botany) ; Osten Sacken, acrostichal setae and closer to 1858: 77 (Strauzia, as syn. of supra-alars then to transverse furrow of Ortalis) Loew, 1873 243 ; : thorax (Fig. 8); scutellum not inflated, (wStraussia may be preserved, after " posterior upper fronto-orbital setae giving description of varieties of absent (Fig. 20), rarely present as a longipennis in Ttypeta) Osten ; setule; oral margin anterior to genal seta Sacken, 1878: 189 (Straussia, as without well developed setae; presutural subgenus of ); Snow, 1894: 159 (Straussia); Aldrich, 1905: 602 thoracic setae present; antennae extending (straussia); Cresson, 1907: 100 but little below middle of face; head higher than long, rounded in profile (~traussia) Phillips, 1923: 125 ; below; proboscis not geniculate; wing (Straussia, with key to varieties); pattern (Fig. 1-7) pronounced, consisting Neave, 1940: 325 (strauzia valid, with of bands of yellow or brown color; Straussia Agassiz, 1846 emendation postocular setae few, thin, acute, and thereof); Foote, 1965: 676 (Strauzia, black. with list of varieties). Why Robineau-Desvoidy spelled Strauzia HOST RBLATIONSHIPS AND BEHAVIOR with a z may never be certainly known, but he stated that he dedicated the name All species of Strauzia known to any to his great contemporary Hercule Eugine extent biologically are feeders in the Straus-Durckheim ("Je le didie b M. larval stages on the pith of the stems of Straus, anatomisten). Agassiz noted in plants of the family Asteraceae the index to his nomenclator that (Compositae). The known hosts are listed Straussia had been used in botany by De under each species in the sequel, and Candolle. The botanical name, however, include plants of the genera Agera tina, was stated by De Candolle (1830) to have Ambrosia, , Rudbecki a, been named for one nLwr. Strauss," who in Smal lanthus , and Verbesina . Several 1666 published what was thought to be the species of Helianthus are involved, but first reference to coffee as a drink. species preferences and possible species Loew apparently considered Straussia to specificity are not yet clear. be a proper emendation of Strauzia (" . Unfortunately, the host data cited by . . Strauzia, which may be preserved, Wasbauer (1972) are largely suspect after being modified into the more correct because of the poor previous state of fom of Strauzian), and most Strauzia taxonomy. entomological authors followed his usage The adult , at least of S. until Foote (1965). except that Cresson perfecta and S. longipennis, are already in 1907 stated definitely that late-day crepuscular. When the sun is Strauzia has priority. There is no close to the horizon, females may be seen justification under modern rules for on their host plant. I do not know how emendation of strauzia, and even if late they remain active nor whether they there were it could only be to Strausia. are also active in early morning. They oviposit between the upper internodes of GENERIC CHARACTERIZATION the plant by standing crosswise on the stem and inserting their ovipositors into Because no satisfactory subfdly the pith. Larvae bore downward, feeding classification has been proposed for North on the pith, until they reach the crown of American Tephritidae, no effort is made the plant in early autumn. Pupation takes here to place Strauzia in a subfamily. place in the crown or adjacent thereto in However, many would place it in the the soil, perhaps according to the species because of well-extended basal of . All known species are univoltine, cubital (anal) cell, non-inflated emerging early the following spring. scutellum, and acute, black postocular Epigamic behavior has not been Vol. 1, no. 3, October 1986 INSECTA MUNDI

described, but the enlarged frontal setae As in the key, one species after another of some species are likely used in has been recognized as of specific rank on territory-defending skirmishes between the basis of relatively slight males. The elongated and specially morphological characters, the only ones patterned wings of some males may be of that appear to be stable. Strauzia use either in intimidating males or perfecta is distinct from all other inducing females to copulate or both. species, but there seems to be no way to make satisfactory groupings of the others DISTRIBUTION on external or genitalic structures. The final residue, here designated as S. The genus is restricted to North longipennis (Wiedemann) may still be a America. One or more species should be species complex, but it is certainly one found in every contiguous State of the of taxa less distinct than the others here United States and in the southern part of recognized. When satisfactory criteria all Provinces of Canada. Very few records can be distinguished, it will then be are available from California (Foote and proper to make examinations of the types Blanc, 1963), and none so far from Oregon of the species at present left in the and Washington. synonymy of S. longipennis. The taxon that Loew considered typical of TAXONOMY longipennis has not been encountered in specimens other than those Loew had. Although more than 1500 specimens were Except for S. longipennis, those here examined in this study, the taxonomic recognized seem to be host specific. problems remain little more resolved than Although a considerable amount of when Snow (1894) made the statement quoted rearing has been done, much more must yet in the introduction. However, we now know be done, as well as mating experiments and that, as in most cases of pronounced hybridizing. Along with such biological sexual dimorphism, such characters are work chromatographic and other chemical variable and that specimens may occur techniques may provide better conclusions. wherein the dimorphic characters are virtually undeveloped and males look like MORPHOLOGICAL TERMINOLOGY females. The instability of thoracic markings has been confirmed, but it is now The terminology of general body parts is known that some parts of the thoracic traditional, that of wing venation is as pattern are much more stable than others. in my recent papers (Steyskal, 1979b, The extent of black marking on the 1984), and that of female terminalia as in scutellum and metathorax is much more my papers (Steyskal, 1977, 19798, 1979b, stable than the longitudinal striping on 1984) and Foote (1981). the mesoscutum. Of the pleural markings, The chief point of difference between my a roundish spot above the hindcoxa is most terminology for the wing venation and that stable. The wing pattern, except for male of McAlpine (McAlpine et al., 1981) lies dimorphism, is not subject to much in the designation of the erstwhile "analw variation. cell. Because I cannot consider CUP a The male postabdomen, as in many other true vein but merely the claval furrow, I Tephritidae (Stone, 1942; Steyskal, 1977, have called the cell "basal cubital 1979a; Foote, 1981). is much more cell." I have also used the designation difficult to work with than the female of the hindmost vein of compound veins for postabdomen and also much less the sake of simplicity (R3, R5, etc.). characteristic at the species level. The I furthermore still find the common male organs of Strauzia species are European designations for the crossveins similar throughout the genus and subject of higher Diptera useful: ta, tb, tp). to considerable variation; they have not My opinions on the female postabdomen been used in this study. differ more fundamentally from those of The female postabdomen, although of McAlpine (1981, p. 38), who states that quite uniform structure throughout the "often in the Diptera all the elements of genus, has proven more useful than that of the terminalia, including the cerci, are the male. Microscope slide preparations called the ovipositor (oviposition tube, have been made, which, because of the ovicauda) . . ." I believe that this is flatness of the parts, are very too broad an extension of the term. The 3 comparable. The spermathecae, also useful sections - - "oviscape, eversible in Strauzia, show up well on slides. ovipositor sheath, and 8th segment with f 104 A MUNDI Vol. 1, no. 3, October 1986 cerciw (McAlpine, 1981, fig. 2.109, 110) inclusion of the Tanypezidae in the -- have not generally been termed superfamily , as was somewhat ovipositor as a whole, at least in the tentatively proposed by McAlpine (1977). Tephritidae, but they are sufficiently The ovipositor per se, the last of the modified to be termed terminalia, or, as 3 divisions of the female postabdomen (or by Hennig, postabdomen. In my key to the terminalia), was divided by McAlpine genus (Steyskal, 1977), I (1977) into 'stalk' and 'aculeus' and by termed the f irat section 'ovipositor Munro (1947) was termed 'aculeus' as a sheath, ' the second section whole. It is certainly of somewhat lovipositubus,v and the terminal section complex origin and composition inasmuch as 'ovipositor.' everything from the 8th segment to the The 7th abdominal segment has the 8th telson is included in it. Although the 2 and following segments retracted into it flaps on the ventral side are likely the when the ovipositor is not in use, often 8th sternum, there is no assurance that with none externally evident. It is the 8th tergum is directly above them. therefore quite properly termed a sheath. Even in the Tipulidae the 8th sternum Munro (1947) called it 'oviscape' and projects well posterad of the 8th tergum. remarked that "it has been called the Epiproct , hypoproct, and 9th and 10th ovipositor, the base of the ovipositor, segments in some form or other are all and the ovipositor sheath." I believe parts of the ovipositor. In most that it is entirely proper to revert to Tephritidae, the apical part of the the pre-Munrovian term 'ovipositor ovipositor (beyond the oviduct) retains sheath. ' The terms 'oviscape' and traces of compound origin. In the oviscapt' are both inapt. The former is Tanypezidae, the 2 sclerites imnediately from Latin ovi- 'egg' and scapus basad of the cerci are obviously present. 'stem, stalk, trunk, cylinder, column' Above the paired 'flaps' there are (originally Greek). The latter term is structures under which the egg must pass; from the same Latin ovi- with Greek I have proposed the term 'oviprovector' skaptos 'dug. ' Neither has any for one such structure in the Otitidae reference to the laying of eggs. It is (Steyskal, 1979b). The finer structure of difficult to form 3-part Latin compounds, the ovipositor in the Tephritoidea and but 'ovipositivagina' is possible and a related groups needs much study. Although word of Greek derivation would be the vast majority of female Diptera with lootococoleus'; neither of these hardly the tephritoid type of ovipositor have seems comnendable. well-developed mouthparts and apparently The long tube that makes possible the feed, at least a little, the fate of the telescoping of all the other part8 into caudal end of the alimentary canal (anal the sheath was termed lovipositubus' by me orifice, rectum, rectal papillae, etc.) in order to have a eingle-word tenn; the seems to be unknown. tenn 'inversion membrane* (Munro, 1947) is I am referring to divisions of the quite as apt and probably the first term antenna and tarsus as segments, a good to be applied to this part. I believe English word referring to similar parts of that all parts of it, bearing either nearly anything. I do not believe that minute spicules or rasping teeth of having a special name (scape, pedicel, various sizes, represent the flagellum) for each of the 3 segments of intersegmental membrane between the 7th the simple antenna of the Muscoidea, and 8th segments. It is certainly not a regardless of their morphological sheath in the ordinary meaning of the significance, contributes anything to the word. The retrorse spicules and teeth are general understanding of keys and apparently secondary sclerotization well descriptions. divided into small portions to maintain A configuration in the ovipositor per great flexibility of the tube. Similar se, probably homologous to the structure is seen in the membrane serving oviprovector, is here used for the first for inversion in other families. In the time and termed the signum. It consists Tanypezidae both the 7th and the following of fine, arcuate, usually interrupted segments retract into the 6th segment by 2 cuticular ridges and grooves. It is separate sections of such membrane. This sometimes difficult to see and always incidentally casts some doubt on the requires high magnification. Vol. 1, no. 3, October 1986 INSECTA MUNDI 105 Key to Species of Strauzia Robineau-Dssvoidy

1 (2). Scutellum, metanotum, and pleural sclerites wholly yellowish, without dark markings; wing (Fig. 3) with hyaline band between crossveins extending anteriorly to costa and posteriorly at least a little beyond Cu; postocellar setae nearly or quite as far apart as distance from one of them to nearest inner vertical seta (Fig. 20). $: Ovipositor sheath (Fig. 9) with broad black apical ring; ovipositor (Fig. 9, 10) with preapical lateral projections; major rasper teeth small, approximately rectangular, with slightly blunt tip; spermathecae teardrop-shaped, preterminal vesicle more than half diameter of terminal vesicle, latter with appressed teeth oblique or even mostly longitudinal; signum absent, present only in New Mexico specimen . d) : Frontal setae usually enlarged, blunt-tipped; wing usually longer than in 8, but with same pattern. Host: Ambrosia trifida L...... S. perfecta Loew Scutellum with lateral corners black; metanotum usually with at least lower black strip, often largely black; pleura unmarked or with black mark above hindcoxae and sometimes also with other dark marking; wing with hyaline band between crossveins not extending posterad of vein Cu, sometimes not reaching costa anteriorly and somtimes in dl variously modified; postocellar setae much closer together (Fig. 21) than distance from one of them to nearest inner vertical seta. $: Ovipositor sheath with narrow black apical ring, sometimes also with dark basal marking; ovipositor (except in S. verbesinae, Fig. 13) with arcuate transition from sides to tip or (S. stoltzfusi, Fig. 14) with slight angulation, oviprovector not develop; major rasper teeth larger, often with extended point or lobe; spermathecae more or less globular, appressed teeth of terminal vesicle latitudinal; signum of ovipositor present. 8 : Frontal setae various; wing various, often with greatly modified pattern.

3 (4). Last tarsal segment blackish, at least in large part; median hyaline band of wing (Fig. 6) extending from costa nearly to vein Cu, ending roundly a little before vein; scutum with short blackish stripe mesad of edge of supra-alar declivity and sometimes with interrupted sublateral stripe; mesopleuron, pteropleuron, sternopleuron, and sometimes hypopleuron above hindcoxa with dark marks; lateral margins of abdominal terga usually darkened. $: Ovipositor sheath (Fig. 18) yellowish or with some basal darkening; ovipositor (Fig. 11) with tips of ventral flaps divergent, rounded; major rasper teeth bilaterally concave to narrow tip; spermathecae with globular terminal vesicle and discoidal preterminal vesicle, about 3/4 of diameter of terminal vesicle. : Frontal setae usually enlarged and blunt-t ipped ; wing somewhat elongated but with pattern as in g. Host: Rudbeckia laciniata L...... S. intermedia Loew

4 (3). Last tarsal segment concolorous with other segments, at most elightly darkened at tip; median hyaline band of wing reaching vein Cu; pleural markings various. : Ovipositor sheath (Fig. 19) largely blackish or yellow with narrow blackish apical band; tips of ventral flaps various; major rasper teeth various; sparmathecae with more or less globular terminal vesicle, sometimes basally truncate, and with preterminal vesicle usually half or less of diameter of terminal vesicle and at least one-third as long as ite own diameter. : Frontal setae and winge various. - -- 106 INSECTA MUNDI Vol. 1, no. 3, October 1986

5 (6). Wings (Fig. 4, 5) with median hyaline band not extending to costa but turned apicad at vein RS or ending; cell Sc (pterostigma) and hyaline spot in apical part of cell br long (pterostigna 3x as long as basal width and hyaline spot attaining level of tip of vein Rl); mesoscutum with well developed pair of sublateral dark stripes interrupted at transverse furrow; large part of mesopleuron with coarse black setae; black spot present on pleuron above hindcoxa. : Ovipositor sheath (Fig. 17) narrowly blackish at tip; ovipositor (Fig. 12a, b) with lateral margins distinctly concave before tip, preapical "shoulders" broadly arcuate and wider then stem; major rasper teeth with distinct, short, apical mucro; spermathecae nearly globular, terminal vesicle with small basal truncation, preteminal vesicle yellow, about 2/3 diameter of terminal vesicle. : Similar to p, frontal setae sometimes slightly enlarged. Host: Helianthus grosseserratus M. Martens .... S. arculata Loew 6 (5). Wing with median hyaline band extending more or less directly to costa; if somewhat constricted, then other characters differing.

7 (8)- $: ovipositor (Fig. 13) with lateral preapical angulations continuing directly to side margins of ovipositor but with small concavity where joining tapered apex; ventral flaps coarctate apically; signum well developed; major rasper teeth bilaterally concave and with narrow points; spermathecae with subglobular terminal vesicle and small colorless preterminal vesicle; ovipositor sheath yellowish with slight apical darkening; thorax with dark markings only above neck and wings, at lateral corners of scutellum, and in lower and lateral corners of metanotum. dr : Similar to p in thoracic markings and wing characters; upper frontal setae enlarged, usually blunt-tipped. Host: Verbesina occidentalls (L.) Walter ...... S. verbesinae, n. ep.

8 (7). : Ovipositor with lateral preapical arcuation or blunt, very obtuse angulation; otherwise differing.

9 (10). Mesoscutum with blackish markings as follows: nearly complete submesal, interrupted sublateral, and short lateral (above supra-alar declivity) stripes; well developed blackish areas also on mesopleuron, pteropleuron, sternopleuron, and hypopleuron above hindcoxa; last tarsal segment not darkened. @: Ovipositor sheath largely blackish; ovipositor (Fig. 14) with very obtuse, blunt lateral preapical angulations; ventral flaps separately rounded apically; major rasper teeth small, bluntly angulate; signum present; spermathecae with small terminal vesicle (collapsed in type specimen), preterminal vesicle long-ovate, relatively large. 8 : Only available specimen with frontal setae not enlarged, wing pattern as in $. Host unknown ...... S. stoltzfusi, n. sp. 10 (9). Mesonoturn variously marked, but lacking long submesal stripes, lateral stripes lacking; mesopleuron, pteropleuron, and sternopleuron at most with small or faint dark markings. : Ovipositor sheath (Fig. 19 largely yellowish; ovipositor with lateral preapical arcuation; rasper teeth frequently with narrow points; spermathecae with large globular terminal vesicle and small preterminal vesicle (Figs. 15, 16 a-g). dr : Frontal setae often enlarged and blunt-tipped; wing often elongated and with modified pattern (differing from that of 9). - - -- Vol. 1, no. 3, October 1986 INSECTA MUNDI 107

11 (12). Distinct spot present on pleuron above hindcoxa. : Ovipositor (Fig. 15) with unusually blunt tip, serrations very fine and not extending full length of tip; major rasper teeth small. rounded; spermathecae with terminal vesicle strongly flattened at base. : Frontal setae not enlarged; wing aa in 9. Host: Hellanthus giganteus L...... S. glgantei, n. sp. Dark spot above hindcoxae usually lacking or faint. $: Ovipositor (Fig. 16a-g) with more sharply pointed tip; major rasper teeth usually distinctly pointed, often bilaterally concave; spennathecae usually with virtually globular terminal vesicle. dl : wing in specimens with well developed sexual dimorphism as in Fig. 1, 2a-c, elongated and with greatly modified pattern; frontal setae usually enlarged and blunt-tipped. Host: Species of Hellanthus and other genera of Asteraceae ...... S. longipennis (Wiedemann) Strauzla arculata (Loew), new status spot above hindcoxa, and most of metanotum (Fig. 4, 5, 12) (except anteromesally). Wing very similar to that of S. longipe~ls (Fig. 1) but Trypeta longlpennis (Wied.), var. hyaline spot below pterostigma in cell arculata Loew, 1873: 242. br with distal end rounded nearly at Straussia longipe~fsvar. arculata level of end of Rl and without small (Loew) Phillips, 1923: 127, pl. 18, fig. spot in cell 2c; tp virtually 7. straight. Legs wholly yellowish. This species has often been confused Abdomen yellowish, with a few faint with S. longipennis, to which it is slightly darker areas; ovipositor sheath indeed more similar than to other very narrowly blackish at apex; terminalia species. I have examined 370 specimens (Fig. 15) with spermathecae consisting of from New Mexico (Valencia Co., in UTA*), roundish, basally truncate terminal Colorado ("Colorado" in CUI), North Dakota vesicle and preterminal vesicle about 0.6 (Hunter, in USNM), South Dakota as large in diameter; major rasper teeth (Brookings, in USNM), Wyoming (nr. Lander, virtually semicircular; ovipositor 2.0 nm in UKAL, Iowa (many localities. in ISU and long, greatest width 0.52 mn; tip USNM) , and Illinois (McHenry Co ., in USNM; unusually blunt, serrations distinct only "Illinoisn holotype, in MCZ). Many were in approximately apical 3/4 of tip, each reared in Iowa from Hellanthus serration 8-10 w wide; ventral flaps grosseserratus M. Martens by W. Bryan apically rounded; signum faint, apparently Stoltzfus; the only other rearing (if of 2 apically convex groups of shallow indeed it is such) from a definite host is striae and reticulations. of a single specimen from Hunter, North 8. Length of wing 5.0-6.4 w. Similar Dakota "ex H. annuus." to $, even in wing shape and pattern and The unique wing pattern described in the structure of head. key and figured makes S. arculata Host: Helianthus giganteus L. The relatively easy to recognize. record of S. intermedia from this host by Novak et al. (1967) actually refers to Strauzia gigantei, n. sp. (Fig. 15) S. gigantei. The species name is the genitive case of giganteus, meaning 'of This species is most similar to S. gigan teus ' longlpe~is, but differs most appreciably . from all forms of that species in details Holotype , 1 mi. S. Kent, Portage of the female genitalia. County, Ohio, E (merged) 4/13/1966, Biol. Length of wing 5.1 - 5.3 mn. Note No. 6552 (W. B. Stolzfus). with Thorax yellowish with following blackish puparium in capsule, ovipositor and sheath markings: Submesal (only near anterior on microslide. Paratypes (all with same end and sometimes fused anteriorly), locality and collection data as holotype): scutellar (at lateral ends of scutellum), 8 allotype, 8 4/22/1966, note no. 6552; adalar (on supra-alar declivity), wedge with puparium in capsule; 9, E 3/24/1966 above halter pointing downward, roundish note no. 6604, with puparium in capsule; c? E 4/26/1965, note no. 6552; dr , E *see appended list of abbreviations. ;/22/1965; cr" , 5/27/1966; all in USNM. 108 INSEC TA MUNDI Vol. 1, no. 3, October 1986

Notes nos. 6552 and 6604 refer to the 1966 Tephri tis trimaculata Nacquart , 1843 : specimens as having been removed from the 383 (sep. 2261, pl. 31, fig. 3. host as late larvae on 18 Sept. 1965, Trypeta cornigera Walker, 1849: 1010. placed in regrigerator as puparia on 5 Trypeta cornigera Walker, 1849: 1011. Oct. 1965, removed on 9 Dec. 1965, again Trypeta longipemis (Wied.) Loew, placed in refrigerator on 4 March 1966, 1862: 59, 65, pl. 2, fig. 2, 3; and finally removed on 28 March; the 1966 1873: 238, pl. 10, fig. 2, 3. specimen with note no. 6604 emerged from a Trypeta longipemis (Wied.), var. puparium found in soil about roots of the typica Loew, 1873: 240. hoet on 5 March. Trypeta longipemis (Wied.), var. longitudinalis Loew, 1873: 240. N. SYN. Strauzia intermedia (Loew), new status Trypeta longipennis (Wied.), var. (Fig. 6, 11, 18) confluens Loew, 1873: 241. Trypeta longipemis (Wied.), var. Trypeta longipemis (Vied. ) , var. vittigera Loew, 1873: 241. intermedia Loew, 1873: 241. (Wied.), var. Straussia longipemis var, intermedia Cresson, 1907: 99, pl. 1, fig. 1 (first (Loew) Phillips, 1923: 127, pl. 18, f. 7. combination). This species is relatively easy to Wiedemannvs type series consisted of an recognize by its largely black last tarsal unstated number of specimens of both segment and the other characters cited in sexes, "in Von Winthem's and my the key and shown in the figures. I have collections." Loew (1862: 59) stated that examined 170 specimens of S. intermedia "26. longipennis Wied. will be more from the following localities, as well as accurately described in the sequel. The the holotype in MCZ from an unknown name of it is acertained from the locality: Manitoba ( Aweme and Minette, in inspection of the originals. It is CNC), Ontario (Beachville, Midland, and surprising that Wiedemann does not mention Ottawa, in CNC), South Dakota, Iowa, the thickening of the frontal bristles in Michigan, Illinois, Indiana, Ohio, the male, though the males in his Pennsylvania, Connecticut, New Jersey, collection show it. Perhaps he had Maryland, District of Columbia, Virginia, specimens enough to satisfy himself that and Georgia (Rabun County, in CNC). this peculiarity is not constant." Then Adults of this species appear earlier in "the sequel" (p. 65, pl. 11, fig. 2, 3) than other species; there is a specimen he gives the figures and description upon from Ag. Coll. (Bast Lansing), Michigan in which the traditional concept of the MCZ with the date "3-30-1892" and one fro species is based. Midland, Ontario in CNC dated 2 May 1959. Several specimens from Indiana, Maryland, Loew (1862: 58-60) also synonymized the and New Jersey were taken in the first 2 Robineau-Desvoidy names, the original week of May. The latest were taken during publication of which provides very little the latter half of June. of moment; the Macquart name, based upon a very sketchy description and a wing figure Host: Rudbecki a 1aciniata L. agreeing well with those of Loew; and the Stoltzfus reared S. intermedia from this 2 Walker names. Walker's notoriously plant in central Iowa, where he found 1 to inadequate descriptions are based upon 3 puparia in the crowns of the plant as males; notes on them kindly furnished by early as 28 July, apparently having done C. W. Sabrosky show that they have the little damage. The report of S. longipennis male wing pattern fairly intermedia in He1 Ian thus gi ganteus by well developed. Novak et al. (1967) is erroneous and Of the 4 varieties erected by Loew and refers actually to S. gigantei. still remaining in synonymy, I feel confident of synonymizing only one, var. Strauzia longipemis (Wiedemann) longitudinalis. A male specimen in MCZ (Fig. 1, 2a-c, 16a-g, 21) bearing the labels "Sharon / Loew coll. / var. longitudinalis / Type 13286" was Tephritfs longipennis Wiedemann, 1830: furnished by me in 1971 with a lectotype 483. label. The male wing pattern is so well Strauzia inermts Robineau-Desvoidy, developed in this specimen that it can 1830: 718. only be longipennis. Strauzia armata Robineau-Desvoidy, A female specimen of var. typica Loew 1830: 719. in MCZ bearing the labels "Penn Auxer / Val. 1, no. 3, October 1986 INSEC TA MUNDI 109

Loew coll. / Type 13290" was furnished by tuberosus in Bethesda, Maryland. Most of me in 1983 with a lectotype label. the males of this population had wings Additional specimens (2 dr , 1 (2) are nearly as extreme in pattern dimorphism as also in MCZ bearing the same original in Fig. 2c, which is of a specimen from labels. It is quite doubtful that var. northern Ohio. Of the female genitalia, typica Loew is S. longipemis Fig. 16a was made from a specimen of the longipennis (Wied.). A microslide of Bethesda population on He1 1anthus the abdomen of the lectotype has been tuberosus, Fig. 16b from H. annuus in returned to MCZ with the remainder of the Ames, Iowa, and Fig. 16c from H. specimen. Fig. 16g in the present work hirsutus in Ames, Iowa. Two further was made from that preparation. The series, represented by Fig. 16d from ovipositor somewhat resembles that of S. uvedalia (L. ) in Blkton, arculata (Fig. 12), but the wing pattern Virginia and Fig. 16e from Ageratina is much like that of a longipennis with altissima (L.) A. H. King and H. Robinson rather expanded dark pattern. It is in Wadsworth, Ohio are somewhat more possible that when more material and some doubtful, although they seem to be well biological data on this entity is within any kind of morphological variation available, it may prove to be a distinct of the Helianthus-feeding forms. The species. No dark mark is present above mentioned series are deposited in USNM. the hindcoxa. Strauzia longipennis may be expected Loew*s var. confl uens , as he to occur throughout the North American originally stated, is represented by a range of the genus Helianthus or even single c? "Connecticut; Mr. Norton." I beyond, and therefore farther west than examined in 1971 a specimen in MCZ which any other species of the genus. I have had only the labels (plain white circle) / seen specimens from the western States of Loew coll. / var. confluens / type 13284. Arizona, Colorado, New Mexico, and Utah. It is a small specimen; Loew said "it is Because of possible recognition of one of the smallest specimens in my distinct species within this complex, collection.** I would interpret it as a detailed description of the range is not small, heavily colored specimen of S. given. longipennis with poorly developed dimorphic male characteristics of head Strauzia perfecta (Loew), n. comb. bristles and wing pattern. The two arms (Fig. 3, 9, 10, 20) of the F-mark in the apical part of the wing are so broad that, as Loew stated, Trypeta longipennis (Uied.), var. they fuse for a short distance in their perfecta Loew, 1873: 239, pl. x. fig. middle. The fl surstyli, as I noted in 2. 1971, are similar to those of other S. Straussia longipennis var. perfecta longipennis. (Loew) Phillips, 1923: 126. LoewVs var. vittigera was described No other species of Strauzia is as from "a male and a female from Nebraska obviously distinct and of specific rank as (Dr.Heyden)"; they were examined by me in S. perfecta. It varies but slightly and 1983 and furnished with a lectotype label may be easily recognized by the lack of on the pin with the female specimen. Both dark markings in the lateral corners of specimens in MCZ, have labels "Neb. / Loew the scutellum and in the 9 (Pig. 9) by coll. / Type # 13287." The lectotype is the broad apical black ring on the rather small, wing 4.4 nm long, and ovipositor sheath. This latter character somewhat shrunken. A slide preparation of was first noted by Snow (1894: 169) but the lectotype (basis of Fig. 16f) was made not recognized as specific. The and returned to MCZ. I see no reason for definitely angulate preapical corners of considering it as other than a rather the ovipositor are also characteristic; cotmuonly found variation in Midwestern the only other species of Strauzia with populations, viz., occasional specimens such angulations is s. verbesinae, but with well developed dark sublateral that species is quite different otherwise stripes. and even the angulations are somewhat The more typical wing patterns and different (cf. key and Fig. 10 and 13). female genitalic structures are shown in It should be noted that Loew used the same Fig. 1 (q) and 2a-c (d)of wings and figure of the male wing for 10nglpeIInI~ Fig. 16a-c of 9 genitalia. Of the and the var. perfecta. wings, Fig. 2a and b are of specimens from I have examined the holotype (in MCZ a single population on Helianthus with locality data) and about 660 110 INSEC TA MUNDI Vol. 1, no. 3, October 1986 specimens from the following localities: ovipositor 1.8 ma long, greatest width 0.4 Ontario (Point Pelee, in CNC), New Mexico ma, tip virtually equilaterally triangular (Pecos, in USNM), Colorado ("Colorado", in beyond rather suddenly slightly projecting USNM; Castle Rock, in ISu; Fort Collins, shoulders, side of ovipositor basad in MCZ) , South Dakota (Brooking8 , in USNM; thereof nearly straight; ventral flaps Jefferson, in ISu), Nebraska (Valentine apically coarctate and ending near (before and Holdrege, in ISU; Overton, in CNC); or beyond) level of shoulders; signum Kansas (Douglas County, in UKaL; Lawrence, present, consisting of semicircular ridges in CNC), Iowa, Illinois, Michigm and striae; lateral serrations of tip 28 - (southern: Saginaw, Berrien, Kalarnazoo, 35, extending nearly to shoulders. St. Joseph, Branch, Ingham, Wayne, and . Similar to ?. Wing length 4.37 - Monroe Counties), Ohio, Kentucky, 5.18 (average 4.76) nm, pattern similar to Tennessee, New York (Ithaca), Connecticut, that of $, but tip sometimes a little Massachusetts, New Jersey, Pennsylvania, more tapering; upper fronto-orbital setae West Virginia, Maryland, Virginia. enlarged and blunt or uncate. Dates of capture of adults extend from Host: Verbesina occi dentalis (L. ) the last week of May to about mid-July, Walter, reared by W. B. Stoltzfua. with a few later stragglers. The species name is that of the genus of Host: S. perfecta has been reared only its host in the genitive case. from Ambrosia trifida L. (first reported Holotype: 5 mi R Harrisonburg, by Novak et al. (1972). In a vacant lot Rockingham County, Virginia, R(merged) in Bethesda, Md., in which numerous plants 10.1V.1967, Biol. Note 6701 (puparia of both A. trifida and Helianthus removed from stalks of host on tuberosus were growing, no Strauzia 25.111.1967), abdomen on microslide, specimens were taken on the "wrong" plant puparium in microvial (W. B. Stoltzfus). - S. perfecta was always on A. trifida Paratypes: 4 9 and 2 (including and S. longipenni s always on H. allotype dr ), asme data as holotype; 1 tuberosus . $, 3 07, same locality, ll.VI.1967; 1 $ , same locality, with puparium, R 25-11-1968: 2 , same locality, with Strauzia verbesinae, n. sp. puparium, R 8.111.1968; 4 d , same (Fig. 13) locality, 21.V.1968 (all collected and/or reared by W. B. Stoltzfus) ; 1 g, 9 d , This species is evidently fairly closely Claytor Lake State Park, Pulaski County, related to S. longipe~fs, but it Virginia, 5.VI.1971 (Wm. H. Robinson); 1 b differs most decidedly in the obviously , Peteraburg, Prince George County, angulate preapical corners (shoulders) of Virignia, 1.VI.1917 (R. C. Shannon); 1 the ovipositor (Fig. 13), the very smell, , Lexington, Kentucky, 23-25. V. 1971, spherical preterminal vesicle of the Malaise trap (P.H. Freytag, G. Leppert); spermathecae, and the lack of specialized all In USNM. wing pattern in the male. The only other species of Strauzia with angulate Strauzia stoltzfusl, n. sp. (Fig. 14) preapical corners on the ovipositor is S. perfecta (Loew), q. v. There is much more extensive darkening . Length of wing 4.64 - 5.50 on the body of this species than in any (average 4.95) mn. Thorax yellowish with other in the genus, but the last segment only following brown to black markings : of the tarsi is not darkened, as they are pair of submesal spots above neck, in S. intermedia, which is probably the sometimes fused mesally; spot above wing next most darkened species. Characters of base; basilateral scutellar spots; lower the female terminalis bring S. and lateral borders of metanotum. Wing stoltzfusi into relationship with S. virtually as in S. longipemis. Legs gigantel, a very pale species, especially wholly yellowish. by the rather blunt tip of the ovipositor. Abdomen yellowish, often with 9. Length of wing 4.75 mn. Head ill-defined brownish patches; ovipositor yellowish with brown area on occiput at sheath narrowly blackish at tip; each side of neck, narrow brown stripe terminalla as in Fig. 13; spermathecae along upper part of dorsal occipital with globular terminal vesicle and very sutures; and black ocellar triangle. small preterminal vesicle oblate spherical Thorax yellowish with following dark brown in shape and of yellowish color; major to black markings: narrowly separated pair rasper teeth mucronate, concave laterally; of continuous submesal black stripes from Vol. 1, no. 3, October 1986 INSEC:TA MUNDI 111

neck to 2/3 distance from transverse relinquishing work on the Tephritidae furrow to scutellum; broad presutural part magnanimously turned over to the narrowly disjunct from narrowed Systematic Entomology Laboratory, USDA, postsutural part of pair of sublateral the exceedingly important results of his black stripes; narrow pair of subalar work on the genus; and John T. Schulz, who black stripes and pair of roundish black was very encouraging in the early stages adalar blotches; small brown mark just of the problem. And far from least, I am lateral to each anterior notopleural seta; grateful to Linda Heath for making true pair of black basilateral marks on art out of my pencil sketches. scutellum; each side of thorax with dorsal The reviews of the manuscript by W. border of propleuron and its posterior Bryan Stoltzfus, Edward W. Baker, and edge alongside spiracle blackish; broad Robert V. Peterson are also gratefully border nearly all around mesopleuron; acknowledged. lower 2/3 of sternopleuron; entire hypopleuron and sclerotic connections with metanotum; entire metanotum and all except ABSTRACT narrow dorsal edge of postscutellurn. Legs wholly yellowish. Wing as in S. Of the 7 presently accepted varieties of longipennis, markings rather deep-colored. the single species of the genus, Strauzia Abdomen with ill-defined but broad and longipennis (Wiedemann), 4 are given conspicuous brown crossbands on tergites; species status: S. 1ongi penni s ovipositor sheath nearly wholly blackish; Wiedemsnn) (type), . arculata (Loew) , terminalia (Fig. 14) with terminal vesicle S. intermedia (Loew), and 5. perfecta of spermathecae collapsed in only (Loew). Three new species, all from the available specimen, preterminal vesicle United States, are proposed: S. large, longitudinally oval, light brown; gigantei, S. st01 tzfusi, and S. major teeth of rasper minute, verbesinae. A key to species is given round-tipped, short, obtusely triangular; and the female terminalia of all species ovipositor 2.0 mn long, greatest width and other details of sow, of them are 0.55 w; ovipositor tip rather blunt, given. Known host plants are also cited. serrations very fine and shallow, about 40 The genus is considered as one of unusual on each side, becoming evanescent about difficulty and one that still requires 2/3 distance from tip to shoulder; extensive work on the biology of its shoulders very obtusely angulate; ventral members before meaningful type examination flaps extending to shoulder level; signum can lead to better solution of their faint, in single mesa1 row of apically systematics. concave ridges and striae. dr. Similar to . Length of wing 4.55 w. Head with frontal setae well Literature Cited developed but neither enlarged nor blunt-tipped. Agassiz, L. 1846. Nomenclatoris Host: Unknown. zoologici index universalis. Soloduri I take pleasure in dedicating this (=Solothurn). viii, 393 p. species to W. Bryan Stoltzfus (genitive Agassiz, L., and H. Loew. 1846. Nomina case, stoltzfusi) in token of his very systematics generum dipterorum. In: considerable and significant work on Agassiz, L., Nomenclator zoologicus, Pt. Strauzia species. 4. 42 p. Holotype $ and 1 dl paratype Aldrich, J. M. 1905. A catalogue of (allotype): St. Anthony Park (Ramsey North American Diptera. Smithsn. Inst., County, St. Paul, Minnesota, no date Smithsn. Misc. Collect. 46 (2 = pub. (Lugger), 57, abdomen and 1 wing of each 1444): 1-680. specimen on microslides; in USNM. Brink, J. E. 1923. The sunflower maggot (Straussia longipennis Wied.). 53rd Ann. ACKNOWLEDGMENTS Report Entomol. Soc. Ontario 1922: 72-74. Caesar, L. 1924. The sunflower maggot - I am most grateful to the many who have generously assisted in the preparation of a pest of sunflowers. Farmers Advocate this paper, and three persons should be 59: 990. Coquillett, D. W. 1910. The type-species especially acknowledged, viz., Richard H. of the North American genera of Foote, who has given constant help and Diptera. U.S. Natl. Mus. Proc. 37: encouragement; W. Bryan Stoltzfus, who in 499-647. 112 INSEC TA MUNDI Vol. 1, no. 3, October 1986

Cresson, E. T., Jr. 1907. Some North Smithsn. Inst., Smithsn. Misc. Collect. American Diptera from the Southwest, 16 (2= pub. 270): 1-276. Paper 11. Trans. Am. Entomol. Soc. 33: Phillips, V. T. 1923. A revision of the 99-108, pl. 1. Trypetidae of northeastern America. De Candolle, A. P. 1830. Prodromus sys- J.N.Y. Entomol. Soc. 31: 119-155, pls. teraatia naturalis regni vegetabilis. 18-19. Pars Quarta. Paris. 683 p. Robineau-Desvoidy, J. B. 1830. Essai sur Foote, R. H. 1965. Family Tephritidae les Myodaires. Inst. France, C1. Sci. (Trypetidae, Trupaneidae). In Stone, Math. Phys., Acad. Roy. Sci., Mem. (ser. A., et al., A catalog of the Diptera of 2) 2: 1-813. America north of Mexico. U.S. Dep. Snow, W. A. 1894. Descriptions of North Agric., Agric. Res. Serv., Agric. American Trypetidae, with notes. Kans. Handbook No. 276: 658-678. Genus Univ. Quart. 2: 159-274, pls. 6-7. Strauzia, p. 676. Steyskal, G. C. 1977. Pictorial key to ------1981. The genus species of the genus Anastrepha Loew south of the United States (Diptera; Tephritidae). Entomol. Soc. (Diptera: Tephritidae). U.S. Dep. Wash., Spec. Publ., unnumbered. 35 p. Agric. Techn. Bull. 1607. iv, 75 p. ------1979a. Biological, anatomical, Foote, R. H., and F. L. Blanc. 1963. and distributional notes on the genus The fruit flies or Tephritidae of Callopfstromyia Hendel (Diptera: California. Bull. Calif. Insect Surv. Otitidae). Proc. Entomol. Soc. Wash. 7: 1-117. 81: 450-455- Loew, H. 1862. Monographs of the Diptera ------1979b. Taxonomic studies on of North America. Part I. Smithsn. fruit flies of the genus Inst., Smithan. Misc. Collect. 6 (I= (Diptera, Tephritidae). Entomol. Soc. pub. 141): 1-221, pls. 1-11. Wash., Spec. Publ., unnumbered. 61 p. ------1864. Monographs of the Diptera ------1984. A synoptic revision of of North America. Part 11. Smithsn. the genus Aciurina Curran, 1932 Inst., Smithsn. Misc. Collect. 6 (2= (Diptera, Tephritidae). Proc. Entomol. pub. 171): 1-360. Pls. 111-VII. Soc. Wash. (in press). Stone, A. 1942. The fruit flies of the McAlpine, J. F. 1977. A revised classi- genus Anastrepha. U.S. Dep. Agric. fication of the Piophilidae, including *Neottiophilidae8 and 'Thyreophoridae* Misc. Publ. 439. 112 p. Sturtevant, A. H. 1925-1926. The seminal (Diptera: ). Mem. Entomol. SOC. Can. 103. (vi), 66 p. receptacles and accessory glands of the Diptera, with special reference to the McAlpine, J. F., et al., eds. 1981. Acalypterae. J.N.Y. Entomol. Soc. 33: Manual of Nearctic Diptera. Volume 1. 195-215; 34: 1-21, pls. 1-111. Res. Branch Agric., Can., Monogr. 27. Walker, F. 1849. List of the specimens vi, 674 p. of dipterous in the collection Macquart , J. 1843. ~iptkresexotiques of the British Museum. Part IV. nouveaux et peu connus. Mem. Soc. Sci. London. p. 688-1172. Agr. Lille 1842: 162-460, pls. I-XXXVI Wasbauer, M. S. 1972. An annotated host (sep. vol. 2, pt. 3: 304). catalog of the fruit flies of America Munro, H. K. 1947. African Trypetidae north of Mexico (Diptera: Tephritidae). (Diptera). Mem. Entomol. Soc. So. Afr. Ocass. Pap. Bur. Entomol. Calif. Dep. 1. (viii) , 284 p., 16 unnumbered plates (fig. 1-321). Agric. 16: 1-172. Neave, S. A. 1940. Nomenclator Zoolo- Westdal, P. H., and C. F. Barrett. 1960. gicus. Vol. iv, Q-Z and Supplement. Life-history and habits of the sunflower 2001. Soc. London, London. p. 1-758. maggot, Strauzia longipemis (Wied.) Novak, J. A., et al. 1967. New host (Diptera: Trypetidae) , in Man1 toba. records for North American fruit flies Can. Entomol. 92: 481-488. (Diptera: Tephritidae). Proc. Entomol. Wiedemann, C. R. W. 1830. Soc. Wash. 69: 146-148. ~ussereuropaische zweifliigelige Inaekten Osten Sacken, C. R. 1858. Catalogue of Vol. 2. Hamn. xii, 684 p., pls. 7-lob. the described Diptera of North America. Smithsn. Inat., Smithsn. Misc. Collect. 2 (1= pub. 102): vii-XX, 1-92. ------1878. Catalogue of the described Diptera of North America. Ed. 2. Vol. 1, no. 3, October 1986 INSECTA MUNDI 113

-- ABBREVIATIONS OF NAMBS OF DEPOSITORIES EXPLANATION OF FIGURES

CNC - Canadian National Collection, The names of the species appear with Ottawa each figure. Figs. 1-7 - wings; 8 - CUI - Cornell University, Ithaca, New scheme of thoracic chaetotaxy and pattern; York 9 - ovipositor sheath and ovipositor; ISU - Iowa State University, Ames 10-16 (left to right) spermatheca MCZ - Museum of Comparative Zoology, (appressed spinules shown only in fig. 10; Harvard University, Cambridge, they are directed equatorially in a11 Massachusetts others)., outline of largest rasper UKaL - University of Kansas, Lawrence spicule, ovipositor; 17-19 - ovipositor USNM - United States National Museum of sheath in profile; 20-21 - median dorsal Natural History, Washington, view of head, showing setae. D.C. UTA - University of Texas, Austin.

1 longipennis

2a longipennis

2b longipennis

2c longipennis 0 \\ ---+/ 114 INSECTA MUNDI Vol. 1, no. 3, October 1986

3 perfecta 9

4 arculata 9

5 arculata d

6 intermedia d

7 stoltzfusi 9 Vol. 1, no. 3, October 1986 INSECTA MUNDI 115

8

anterior intra-alar Dorsal Thoracic Chaetotaxy and Pattern notopleural orsocentral Primary pattern: submesal (uauaYy only anterkrly), rublateral. and scutelkr (exc. In 8. porfocta) Secondary pattern: supra-alar and extended rubmoral Not rhown: adakr scutellar

9a perfecta dorsal

ventral

9c perfecta

9b perfecta

10 perfecta signum \

11 intermedia

I 12a arculata 116 INSECTA MUNDI Vol. 1, no. 3, October 1986

------~ ~~ ~

12b arculata

13 verbesinae

(

14 stoltzfusi .

C .\.--, ------.- - .------.-.- 15 gigantei

.- ___ -- .._ ,I,/?-lc~:X.'~rqi:;++. r

16b longipennis

( - .-.-=::= -

16c longipennis

-----.__.___

16d longipennis L-

( < 16e longipennis Vol. 1, no. 3, October 1986 INSECTA MUNDI 117

16f longipennis

---- ..-.--

16g longipennis

17 arculata

20 perfecta

18 intermedia

2 1 longipennis 19 longipennis