16 October 1998 PROC.ENTOMOL.SOC.WASH. 100(4), 1998, pp. 636-657

THE GENERA MACROXYEU KIRBY AND MEGAXYELA ASHMEAD (: ) IN NORTH AMERICA

D AVID R. S MITH AND N ATHAN M. SCHIFF

(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, MRC-168, Washing- ton, DC 20560, U.S.A. (e-mail: [email protected]); (NMS) U.S. Department of Agriculture, Forest Service, Southern Hardwoods Laboratory, PO. Box 227, Stoneville, MS 38776, U.S.A. (e-mail: [email protected]).

Abstract.-Five species of Megaxyela, including Megaryela alzkonae, n. sp., and two species of Macroxyela occur in North America. Macroxyela bicolor MacGillivray is a new synonym of Macroxyela ferruginea (Say). The species are keyed, described and illustrated, and biological information is summarized. Hosts include hickory, pecan, and elm. Key Words: Megaxyela, Macroxyela, Xyelidae, Ulmus, Carya, pecan, North America

The large xyelids of the genera Megax- for Xyelidae, he suggested that Megaxyela yela and Macroxyela are among the most and Macroxyela form a natural unit based intriguing groups of because of on wing venation and that Xyelu Dalman, their odd appearance, scarcity, and probable Neoxyela Curran (= Xyela) and Pleroneura primitive status in the Symphyta and Hy- Konow form a separate closely related menoptera. They are apparently two relic group. Smith (1967) proposed a classifica- genera in a family with a long fossil record tion based on larval and adult characters (Rasnitsyn 1969). Megaxyela occurs only in and concluded that Macroxyela and Megax- eastern Asia and eastern North America, yela form the tribe Macroxyelini in the sub- and Macroxyela only in North America. family Macroxyelinae. The other tribe of Larvae of both genera are associated with Macroxyelinae, Xyeleciini, includes Xyele- angiospermous trees, whereas all other cia Ross, a genus found in western North Xyelidae are associated with conifers. They America and Japan. are seldom collected, although Macroxyelu We have accumulated about 600 speci- ferruginea (Say) can be collected in num- mens of the genera Megaxyela and Ma- bers on its host during its flight time. Why croxyelu and are now able to resolve some Megaxyela is so scarce is puzzling. Such questions and present a more complete pic- large (lo-15 mm long) and colorful ture of species composition in North Amer- with long hindlegs (Fig. 3) certainly should ica. We recognize five species of Megaxyela be sighted more frequently by collectors. and two species of Macroxyela. Both genera were revised by Ross Specimen depositories and acronyms (1932), but few specimens were available used are in Acknowledgments. w to him, and he was not able to resolve sex associations or present a complete concept T RIBE M ACROXYELINI of the North American fauna. Although he The Macroxyelini differ from other Xye- did not use subfamily or tribal categories lidae by wing venation (Figs. 4-6): forewing

VOLUME 100. NUMBER 4 637

Figs. I-2. Clypeus and labrum. 1, Megaxyela aviingrata. 2, Macroxyela ferruginea.

with vein Rsl; C free from SC forming costal (Fig. 1); forewing with Sc2 joining SC + Rl ‘/4 and subcostal cells; vein Sc2 distinct; Sc2 to % distance between separation of Rs and stigma (Fig. 4); labial palpus 4-segmented C joining S + Rl apical to separation of Rs; Megaxyela hindwing with two closed submarginal cells Clypeus with small anterior median notch and (1Rs and 2Rs) (wing venation terminology labrum broad, rectangular, and nearly truncate follows Goulet and Huber 1993). Also, the anteriorly (Fig. 2); forewing with Sc2 joining apical filament of the antenna is short, one- SC + RI very slightly apical to juncture of Rs with SC + Rl (Figs. 5, 6); labial palpus 3.seg- quarter to one-third the length of the thud mented Macroxyela segment, and 5-l 1 segmented, and the tarsal claws have a long inner tooth, nearly as long G ENUS MEGAXYELA ASHMEAD as the outer tooth. Megaxyela Ashmead 1898: 214.-Ross K EY TO G ENERA OF M ACROXYELINI 1932: 161-164 (N. Amer. species; syn.: 1. Clypeus with large median triangular tooth and Odontophyes, Paraxyela).-Ross 195 1: 7 labrum rounded with notch on anterior margin (N. Amer. catalog).-Smith 1978: 24 638 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(world catalog).-Smith 1979: 10 (N. with clearly defined serrulae at its apex Amer. catalog).-Shinohara 1992: 783- (Figs. 22-29). 786 (Asian species). Type species: Xyela Known host plants are members of the major Cresson. Orig. desig. Juglandaceae. The hosts for the North Odontophyes Konow 1899: 42. Type spe- American Megaxyela aviingrata (Dyar) cies: Pleroneura aviingrata Dyar. By and M. major (Cresson) are Carya spp. monotypy. and perhaps Juglans spp. The hosts for M. Megaloxyeta Schulz 1906: 88. Emendation. gigantea in eastern Asia are Juglans spp. Paraxyelu MacGillivray 1912: 294. Type and Pterocarya spp. (Shinohara 1992). species: Xyelu tricolor Norton. Orig. desig. The few known aspects of their biology is given under each species. A brief account Clypeus with sharp, median triangular of M. gigantea in Korea by Sato (1941), tooth and labrum rounded with anterior, as given by Shinohara (1992), is note- median notch (Fig. 1); labial palpus 4-seg- worthy in that it contains some unusual mented; forewing with SC long, Sc2 joins habits for sawflies: “This species has a Sc+Rl % to ‘/z distance between the sepa- univoltine life cycle, with adult appear- ration of Rs and stigma. Antenna1 flagellum ance from late April to early May. The 6-l 1 segmented. General habitus of all spe- female lays an egg along the midrib on cies similar to that of M. alisonae, n. sp. the upper surface of the apical part of a (Fig. 3). newly growing leaf. Before oviposition, Few specimens are available for study, the female stands on the under surface of but we have seen enough (about 65) to the leaf, with the head directing toward resolve several problems, including spe- the base of the leaf. Then the female cies identity and association of sexes, and bends the tip of her abdomen to the upper present a better understanding of the surface of the apex of the leaf and ovi- North American fauna. We recognize five posits there, folding by hind legs the apex species. of the leaf in half along the midrib upward The genus occurs only in eastern North and attaches the folded halves with glu- America and eastern Asia. Shinohara tinous material. One egg is laid per leaf. (1992) treated three species from eastern The larval period is from late May to ear- Asia, M. gigantea Mocsky from eastern ly June. The larva always starts feeding Russia (Primorski Krai), Korea, and China; from the apex of the leaf and coils on the M. togashii Shinohara from Japan; and M. midrib at rest. The larva has four molts parki Shinohara from Korea. and overwinters in the soil.” Two species groups are apparent. The

Asian species and Megaxyela major, M. K EY TO N EARCTIC S PECIES OF MEGAXYELA uviingratu, and M. alisonae from North Legs black or orange brown, usually with hind- America form one group characterized by tarsus white 2 the short sheath with the ventral and dorsal Legs bright orange with hindtibia and tarsus margins curving mesally to an acute apex black or hindtibia black and hindtarsus white 3 (Figs. 7-9) a thick hindbasitarsus (Fig. 15) Entirely black except for white hindtarsal seg- ments 225 or 3-5; sheath about 2.5X longer the lance lacking dorsal teeth, and the lancet than broad (Fig. 8) aviingruta (Dyar) having indistinct serrulae (except M. alison- Usually entirely orange brown, sometimes var. ae) (Figs. 16-19). The other group includes ious black markings on dorsum of head, tho- the North American Megaxyela tricolor and rax, and/or abdomen and on femora and tibiae; M. bicoloripes, which have a long sheath, sheath about 3X longer than broad (Fig. 7) major (Cresson) straight on its ventral margin (Figs. 10, 1 l), Thorax black; sheath longer than hindtarsus, a slender hindbasitarsus (Fig. 14), the lance about 4X longer than broad (Fig. 11) (legs with with distinct teeth at its apex, and the lancet hindtibia and hindtarsus black; hindbasitarsus VOLUME 100, NUMBER 4 639

Fig. 3. Habitus of Megaqela alisonae, length 12 mm.

slender, as in Fig. 14, about 6.5X longer than Megaxyela alisonae Smith and Schiff, broad) tricolor (Norton) new species Thorax black with mesostemum and mesepi- sternum orange; sheath shorter than hindtarsus (Figs. 3, 9, 15, 20, 21) and less than 3X longer than broad (Figs. 9, 10) ...... ___.....__.. 4 Female.-Length, 10.5-12.0 mm. Anten- Hindtarsal segments 2-5 white; hindbasitarsus na black with scape brownish. Head black slender, about 6.5X longer than broad and tar- with clypeus, labrum, mouthparts, and man- sal segments 2-4 much longer than broad (Fig. dible white, tips of mandibles brownish, 14); sheath at apex in lateral view nearly straight below, slightly emarginate on dorsum and apical labial palpal segment blackish. (Fig. 10) hicoloripes (Rohwer) Thorax orange with pronotum, mesonotum, Hindtarsal segments 2-5 black; hindbasitarsus and upper corners of mesepisterum and me- broad, about 3.5X longer than broad and tarsal sepimeron black to purplish metallic; pos- segments 2-4 about as long as broad (Figs. 15); sheath at apex in lateral view emarginate terolateral portion of pronotum sometimes on both dorsal and ventral margins (Fig. 9) dark orange; mesonotal lateral lobe may be ...... alisonae, new species partly orange; tegula white. Abdomen or- 640 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Wings of Megmyla major. ange, terga with narrow black posterior greatly amused by her brother’s (NMS) margins; apical tergum black or about half choice of an entomological career. of 8th and all of 9th black; sterna with pos- Host.-Unknown. terior margins narrowly white; sheath Discussion.-The sheath shape (Figs. 7- black; basal plate orange. Legs orange with 9) and thick hindbasitarsus (Fig. 15) are extreme tip of hindfemur, hindtibia, and similar to Megaxyela major and M. aviin- hindtarsus black. grutu. The largely orange thorax, abdomen, Antenna1 filament 9-segmented and sub- and legs and the serrate dorsal margin of equal in length to scape; antenna1 length the lance and distinct serrulae of the lancet about 2.3X head width. Hindbasitarsus will separate M. alisonae from those spe- thick, about 3.5 X longer than broad. Length cies. of hindbasitarsus about 0.8X length of re- maining tarsal segments combined; length Megaxyela aviingrata (Dyar) of hindtarsus about 0.7X length of hindtib- (Figs. 1, 8, 16, 17) ia. Sheath (Fig. 9) length about 1.1 X length Pleroneura aviingrata Dyar 1898a: 213.- of basal plate and 0.6~ length of hindtibia. Dyar 1899: 127 (larva).-Smith 1978: 26 Lance with serrate dorsal margin (Fig. 20); (as a syn. of M. tricolor).-Smith 1979: serrulae of lancet small, narrowly pointed lO.-Smith 1987: 386 (lectotype). at apices (Fig. 21). Odontophyes aviingrata: Konow 1899: Male.-Unknown 42.-MacGillivray 1906: pl. 23, fig. 32 Holotype.-?, labeled “U.S.A.: Pine- (wings). bush, N.Y.: Albany Co., 42.42.45-75.52.53, Megaxyela aviingrata: Ross 1932: 163.- 21 May 1983, T. McCabe, 100 m.” Depos- Ross 1951: 7.-Burks 1958: 8. ited in the New York State Museum, Al- bany, New York. Female.-Length, 11.0-12.5 mm. Black Paratypes.-NEW YORK: Ithaca, 2 1 with following white: labrum except for May ‘36 (1 0, UCB). OHIO: Franklin Co., black apical margin, labial palpus except 5-22-79 (1 9, OSU). for black apical segment, extreme apices of Distribution.-New York, Ohio. fore- and midtrochanters and most of hind- Etymology.-The specific epithet is in trochanter; extreme apices of fore- and mid- memory of Alison Wyn Schiff who was tibiae; hindtarsal segments 2 or 3 to 5 with VOLUME 100, NUMBER 4 641

Figs. 5-6. Wings. 5, Macroxyela ferruginea. 6, M. aenea; arrow points to crossvein in cell Rsl. sometimes extreme apex of hindbasitarus nal length about 1.9X head width. Hind- whitish and basal part of hindtarsal segment basitarsus broad, about 3.5 X or more longer 2 blackish. Apex of mandible brownish. than broad (similar to Fig. 15). Length of Wings hyaline with veins and stigma black. hindbasitarus about 0.7X length of remain- Antenna1 filament 8-9 segmented; anten- ing tarsal segments combined; length of 642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hindtarsus 0.8X length of hindtibia. Sheath Zuns are based on Dyar’s observations of (Fig. 8) about 1.1 X length of basal plate larvae. and 0.6X length of hindtibia. Dorsal margin Biology.-Dyar’s rearings bear his code of lance smooth, without serrations; lancet “3C” (Smith 1987). Dyar (1898a) reared without discernible teeth (Figs. 16, 17). this species and presented some sketchy Male.-Length, 11.0 mm. Color and notes. He stated “curious larva resembling structure as for female. Genitalia similar to the excrement of birds”. . .“on the young that of M. major (Fig. 33). tender leaves of the hickory and butternut Specimens examined.-20 0, 1 8. in May. The larvae disappear by the end of Distribution.-INDIANA: Pine, V-2 l- the month and reappear the following sea- 05. KANSAS: Riley Co. MASSACHU- son. The larva is solitary, if abundant there SETTS: Boston, V-18-15. MISSISSIPPI: may be several on one leaf. In the natural State College, 3-24-32, pecan L834-13; 3- position of rest, curled spirally around a 27-3 1, pecan L834-9; 4-28-3 1, pecan L834- leaf petiole. . . . ” “Enters the earth without 11; 3-3 1, adult L777-22; Oktibbeha Co., ultimate stage. Single brooded, feeding only Starkville, 111-23-1975, in black light trap on immature leaves.” His larval records edge of deciduous woods. MISSOURI: Co- are: Staten Island, N.Y., June 1; Bellport, lumbia, 9-30-56 (male). NEW YORK: Ith- Long Island; Bedford Park, N.Y., May 25; aca, IV-29-13; V-9-36; Bedford Park (Dyar and Fort Lee and Plainfield, N.J., May 16. 1898b). OKLAHOMA: Oklahoma City, V- Discussion.-Only one male (a specimen 4-1964, on pecan (larvae). ONTARIO: collected in Missouri) was available for Guelph, VIII-8-86; Hamilton, V-l 8-80. study. This species closely resembles h4. PENNSYLVANIA: Notch, V-13 & 15-13 major, except for the black coloration and (larvae). Ross (1951) gave the distribution stouter sheath (about 2.5X longer than as eastern, west to Illinois. Ross (1932) stat- broad; Fig. 8). Both A4. uviingrutu and M. major have a similar distribution and both ed that he collected two larvae on hickory have been recorded from pecan in Missis- at Charleston, Ill., May 3 1, 193 1, and his sippi. record may be based on this larval collec- tion. Identification of larvae, however, is Meguxyelu bicoloripes (Rohwer) tenuous. The identity of the two records (Figs. 10, 14, 26-29, 34) from larvae need confirmation. Lectotype.-The female designated by Odontophyes bicoloripes Rohwer 1924: Smith (1987) is in the USNM; it was reared 215. from larvae on hickory from Bedford Park, Meguxyelu bicoloripes: Ross 1932: 163.- New York, by Dyar. Ross 1951: 7.-Smith 1978: 24.-Smith Hosts.-Carya sp. and pecan are the only 1979: 10. definite hosts recorded. The records from Female.-Length, 10-l 1 mm. Antenna pecan are from Mississippi (see Distribu- black. Head black with clypeus and labrum tion). Dyar (1898a) observed larvae which white; apex of mandible and mouthparts he called A4. aviingrutu on hickory and but- brownish. Thorax mostly orange with pro- ternut at other locations, but his rearings notum, mesonotum, metanotum, upper cor- from these collections were not successful. ner of mesepistemum, upper half of mese- Several species of Meguxyela are possible pimeron, and upper comer of metapleuron in the area of his collections, and it is pos- black; tegula white. Abdomen black dor- sible these larvae may be something else. sally and laterally, orange ventrally; basal Ross (1951) recorded “Hicoria,” and plates orange; segments with a narrow Burks (1958) recorded C’uryu sp. and Jug- white posterior margin; sheath black with Zuns cinereu L. Both of their records of Jug- extreme apex dark orange. Legs orange VOLUME 100. NUMBER 4 643

Figs. 7-l 1. Female sheaths. 7, Megaxyelu major. 8, M. aviingrata. 9, M. ulisonae. 10, M. bicoloripes. 11, M. tricolor. with extreme apex of hind femur black on sometimes black at extreme base). Head, outer surface, hindtibia black, hindbasitar- body and legs except tarsi with faint dark sus black with extreme apex white; and purplish tinge. Wings hyaline; veins and hind tarsal segments 2-5 white (segment 2 stigma black. 644 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Antenna1 length about 1.9X head width; Megaxyela major (Cresson) apical filament 9-segmented (in one speci- (Figs. 4, 7, 18, 19, 33) men; others broken) and subequal in length Xyela major Cresson 1880: 34.-Cresson to scape. Hindbasitarsus about 6.5X longer 1916: 5 (lectotype). than broad and length about 0.6X length of Macroxyela major: Kirby 1882: 351. remaining tarsal segments combined; hind- Megaxyela major: Ashmead 1898: 214.- tarsus about 0.7X length of hindtibia. MacGillivray 1906: pl. 23, fig. 32 Sheath (Fig. 10) length about 1.1 X length (wings).-Ross 1932: 162 (syn.: ferru- of basal plate and about 0.8X length of ginea Bridwell).-Ross 195 1: 7.-Smith hindtibia. Lance (Figs. 26; 28) with dorsal 1978: 25 (syn.: langstoni Ross).-Smith serration restricted to distal third; lancet 1979: 10. (Figs. 27, 29) with fine serration on distal Odontophyes ferruginea Bridwell 1906: 94. quarter. Megaxyela langstoni Ross 1936: 13 l.- Male.-Length, 9.5 mm. Coloration as Ross 1951: 9.-Webb 1980: 132. for female; hypandrium black to dark or- ange. Antenna1 filament 9-segmented. Gen- Female.-Length, 9.5-l 1.5 mm. Mostly italia as in Fig. 34. orange to reddish brown with yellowish and Specimens examined.4 9, 1 d. sometimes black markings, usually with Distribution.-MICHIGAN: Washtenaw following more yellowish: eye orbits, or Co., Ann Arbor, IV-21-32. MISSISSIPPI: much of head and mouthparts except for Agricultural College, 4-16-15 (holotype). postocellar area; tegula; fore- and midlegs, MISSOURI: Boone Co., Columbia, 4-26- hindtrochanter; basal third of hindtibia, 73; 4-25-73. PENNSYLVANIA: Spring hindbasitarus and sometimes two or more Br., V-7-1945, H.K. Townes; Ross (1932) following segments; hind margin of abdom- designated an allotype from “Penn.“; we inal segments; and sheath. Following may be blackish: frons; part of cervical sclerites; did not see this specimen. VIRGINIA: Ar- upper corner of mesepisternum; small spots lington, VIII-2 l-5 1. anteriorly on mesoprescutum, lateral por- Holotype.-Female, USNM type no. tions of mesonotal lateral lobes; small lat- 26040, labeled “BarbarinAE., A&M.C, 4/ eral spots on metanotum; hindcoxa except 6/15.” Rohwer (1924) gave the type local- apex; apical half hindfemur and hindtibia; ity as “Agricultural College, Mississippi”; center of basal plates; terga 6-7. Wings hy- The collector is A.E. Barbarin aline to a slightly yellowish tinge; veins Host.-Unknown. brown; forewing stigma black, sometimes Discussion.-Megaxyela bicoloripes and with apical half yellowish. M. tricolor form a group based on the shape Antenna1 filament 6-7 segmented, short- of the sheath, lance and lancet, and slender er than scape; antenna about 2.1 X head hindtibia, as outlined in the introduction to width. Hindtarsus 0.7X length of hindtibia; the genus. The mostly orange legs, orange hindbasitarsus about 0.9 X length of remain- mesosternum and mesepisternum, and ing tarsal segment combined and and about sheath shape distinguish M. bicoloripes 4.2X longer than broad. Sheath (Fig. 7) from M. tricolor, which has the ‘thorax en- about 1.2X length of basal plate and 0.6X tirely black. Furthermore, the sheath is length of hindtibia. Dorsal margin of lance shorter than the hindtarsus and about 3X without teeth; lancet without discernible longer than broad in M. bicoloripes, where- teeth (Figs. 18, 19). as that of M. tricolor is much longer than Male.-Length, 9.0-10.5 mm. Colora- the hindtarsus and about 6.5X longer than tion and variation similar to that of female, broad (compare Figs. 10, 11). sometimes with slightly more extensive VOLUME 100, NUMBER 4 645

15 Figs. 12-15. 12-13, Female sheaths. 12, Mucroxyela ferruginea. 13, M. aeneu. 14-15, Hindtarsus. 14, Megaxyela hicoloripes. 15, M. alisonae. black marks at same spots and thoracic and IV-8-1963, on pecan (larvae). ARKAN- abdominal dorsum. Genitalia as in Fig. 33. SAS: Sheridan, IV-22-25, on pecan leaves Specimens examined.-32 0 and CT ; (larvae). FLORIDA: Alachua Co., 9 mi. about 15 vials of larvae. NW Gainesville, UF Hort. Unit, 19-111- Distribution.-ALABAMA: Madison 1977; Monticello, IV- lo- 1963, on pecan Co., VI-1962, pecan (larvae); Mobile Co., (larvae). GEORGIA: Byron, 4-17-4-20-8 1, 646 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

IV-21-1977, on pecan (larvae); Thomson, ~011. J.M. Langston, reared from pecan McDuffie Co., IV-26-1960, ex pecan (lar- (Curya pecan A. & G.). Both sexes were vae); Jasper Co., IV-30-1958, on pecan (lar- described, and Ross (1936) also designated vae); Jonesboro, IV-29- 1959, pecan trees an allotype and 9 paratypes. (larvae); Ty Ty, IV-1-1964, on pecan (lar- Hosts.-Carya illinoensis (Wang.) K. vae). IOWA: Story Co., state nursery, April Koch (pecan). An adult was collected on 6, 1978. KANSAS: Baldwin (holotype, 0. “early shrub willow” (Bridwell 1906). ferruginea); Douglas Co., V-1933; Burling- Discussion.-Ross (1936) stated that the ton, on pecan, V-2-25 (larvae). MISSISSIP- larvae recorded by Dyar (1898b: 175) and PI: State College, Apr. 7, l932 and March Yuasa (1923) were M. major. Yuasa’s 24, 1932, reared from pecan, J.M. Langston “Megaqela sp. 1” may also be M. major. (mostly paratypes of M. Zangstoni), 4-27- Megaqela langstoni larvae have a large 31, pecan L838B; 4-7-32; Lucedale, reared dark area on the pronotum and penultimate from pecan, emerged April 20, 1930, and abdominal segment. Yuasa’s sp. 1 has these March 28 and April 2, and April 9, 193 1, areas with a pair of black spots. Also, M. H. Dietrich (some paratypes of A4. Zung- major appears to be a solitary feeder on pe- stoni); Ag. Coll. Miss., 4-10-1915, pecan. can and some other hickories, whereas M. MISSOURI: St. Joseph, V-3-1939. NEW lungstoni is gregarious. The dark areas on YORK: McLean, IV-26- 19 13. OKLAHO- the pronotum and penultimate abdominal MA: Latimer Co., IV-1989; Cleveland Co., segment are variable and cannot be used for IV-8- 193 1; Madil, Marshall Co., IV-28-64, species separation. We cannot comment on on pecan (larvae). PENNSYLVANIA: the solitary vs. gregarious feeding habit. Huntingdon, VI-3-34; Dauphin Co., VI-8- Ross (1936) separated Megaxyela lang- 34, on hickory (larvae). SOUTH CARO- stoni from M. major by the chestnut ab- LINA: Clemson, IV-20-1975; Beaufort Co., dominal dorsum (not mostly black as in M. IV-18-1959, on pecan (larvae). TENNES- major), black area at base of the stigma, and SEE: McNairy Co., V-13-71, on pecan (lar- less yellow at the base of the hindfemur and vae). TEXAS: “Tex.” ; Concan, 4- 14-49; hindtibia. Examination of the specimens at Waco, on leaves of pecan, IV-2 1- 1906 (lar- hand show considerable color variation, and vae); Fredericksburg, IV-2 1- 1960, pecan fo- we believe Ross’ M. langstoni is one of the liage (larvae). VIRGINIA: Miners Hill, on paler color forms of M. major. Ross also hickory, V-14-12 (larvae, identity ?). Re- stated that the sheath is shorter than the cords are for adults unless otherwise indi- hindbasitarsus in M. Zangstoni (longer than cated. in M. major). However, the sheath appears Types.-Xyela major Cresson was de- to be longer than the hindbasitarsus in the scribed from a female and a male from specimens Ross examined. “Texas (Belfrage),” The lectotype, chosen In summary, there seems to be behavior- by Cresson (1916), is at ANSP Type No. al, color and morphological variation within 330. M. major, but, at present, it appears most Odontophyes ferruginea Bridwell was likely that this is intraspecific variation rath- described from one female. The holotype is er than several cryptic species. Additional in the USNM, with type labels and “Bald- specimens, especially from intermediate lo- win, Kansas,” “Bridwell, Apr.,” and “on calities, are needed to elucidate the full ex- early shrub willow.” This is one of the tent of this variation. darker specimens with blackish marks on Dyar (1898b) attempted rearing some lar- the frons, mesonotum, and abdominal terga. vae under his code “7D.” These were on The holotype of Megaxyela langstoni hickory, Fort Lee, N.Y., Bronx Park, N.Y., Ross is at the INHS (Webb 1980). It is from and Bedford Park, N.Y. No adults were ob- State College Mississippi, April 7, 1932, tained, and thus the identity of his speci- VOLUME 100, NUMBER 4 647

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20

21

Figs. 1621. Female ovipositors. 16, Lance of Megaxyela aviingrata. 17, Lancet of M. aviingrata. 18, Lance of M. major. 19, Lancet of M. major. 20, Lance of M. alisonae. 21, Lancet of M. alisonae. Figs. 16-21 drawn to same scale mens are unknown. Authors, e.g. Ross Megaxyela major belongs to the group of 1932, Smith 1987, recording Dyar’s rear- species with M. aviingrata and M. alisonae ings as M. major may or may not be cor- which have a thick hindbasitarus (Fig. 15) rect. and similar sheath shape (Figs. 7-9). It can 648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON be distinguished from those species by the length 2.1 X head width. Otherwise similar nearly entirely orange-brown coloration, to female. Genitalia as in Fig. 35. but with various degrees of black on the Specimens examined.-1 ? , 4 d . dorsum of the head, thorax, abdomen, fem- Distribution.-CONNECTICUT Storrs, ora, and tibiae. May 1935. ILLINOIS: Algonquin; Urbana, April 12, 1891 (Ross 1932; specimen not Meguxyela tricolor (Norton) seen). KANSAS: holotype, no further data. (Figs. 11, 22-25, 35) MISSOURI: Sapp, V- 10-56. ONTARIO: Xyela tricolor Norton 1862b: 144. Simcoe, 2-V-55. Locality unknown: One Macroxyela tricolor: Kirby 1882: 351. male labeled “16093” possibly from Illi- Paraxyela tricolor: MacGillivray 19 12: nois, at INHS. 294. Holotype.-At ANSP, male, labeled Megaxyela tricolor: Ross 1932: 162.- “KS.,” red label “Type No. 331,” and “tri- Ross 1951: 7.-&&h 1978: 26.-Smith color. ” Norton described the species from 1979: 10. “One specimen from Kansas.” Host.-Unknown. Female.-Length, 10 mm. [Antennae Discussion.-For separation from M. bi- missing.] Head and thorax black with a pur- coloripes, see the preceeding key and dis- plish metallic tinge; clypeus white except cussion of that species. We have seen only for black narrow margins and central spot; one female (from Ontario), and this is as- labrum white with apical half black; man- sociated with males only by the coloration. dible brown to black at base underneath; Previously, M. tricolor was known only mouthparts brown; and tegula white. Legs from several males and M. bicoloripes only orange, spot on outer surface of mid- and from females. Because of the similarity of hindcoxa, spot at base of outer surface of M. tricolor and M. bicoloripes, Ross (1932) hindcoxa, extreme apex on outer surface of thought they may eventually be shown to hindfemur black, all hindtibia and hindtar- be opposite sexes of the same species. Our sus black. Abdomen black with white nar- studies show that there are two separate row posterior margin on each segment, ba- species involved. sal plate of sheath brown to orange; and sheath black. Wings hyaline with veins and Genus Macroxyela Kirby stigma black. Hindbasitarsus slender, about 6.5 X lon- Mucroxyela Kirby 1882: 35 1 .-Ashmead ger than broad and about 6X length of re- 1898: 214 (in key).-Ross 1932: 164 (re- maining tarsal segments together; length of vision; syn.: Protoxyela MacGillivray).- hindtarsus about 0.8X length of hindtibia. Ross 195 1: 7 (N. Amer. catalog).-Smith Sheath (Fig. 11) unusually long and slender, 1978: 23 (world catalog).-Smith 1979: about 1.6X length of basal plate and about 10 (N. Amer. catalog). Type species: Xye- 1.3X length of hindtibia. Lance (Figs. 22, la ferruginea Say. Orig. desig. 24) with dorsal serrations restricted to api- Protoxyela MacGillivray 1912: 295. Type cal third or less; lancet (Figs. 23, 25) with species: XyeZu aenia (!) Norton. Orig. de- distinct teeth only on ventral margin of api- sig. cal quarter. Male.-Length, 9.0-9.5 mm. Coloration Median lobe of clypeus notched medial- mostly same as for female; antenna black; ly, with two small round lobes, on each side clypeus and mandible mostly whitish; hy- of emarginated meson and labrum short, pandrium sometimes dark orange. Apical rectangular, nearly truncate anteriorly (Fig. antenna1 filament 6-8 segmented with 2). Labial palpus 3-segmented. Forewing length subequal to length of scape; antenna1 with SC short, Scl joining SC + Rl very VOLUME 100, NUMBER 4 649

23

24

26

Figs. 22-29. Female ovipositors, 22, Entire lance of M. fr~cobr. 23, Entire lancet of M. tricolor. 24, Apex of lance of ,14. tricnlor. 25, Apex of lancet of M. tricolor. 26, Entire lance of M. bicolori~es. 27, Entire lancet of M. bicoloripes. 28, Apex of lance of M. bicnloripes. 29, Apex of lancet of M. bicoloripes. Figs. 22-29 drawn to same scale.

slightly apical to juncture of Rs with S C + KEY TO S PECIES OF M ACROXYELA Rl (Figs. 5, 6). 1. Sheath slender, about 4X longer than broad Two species are known for this exclu- (Fig. 12); radial region of forewing with 3 cells sively North American genus. A4acroxyela (in 85% of specimens examined) (Fig. 5), if ferruginea is by far the most commonly with 4 cells, then crossvein in cell Rsl located toward distal portion of cell; female mostly collected species. The larva feeds on elm; reddish brown; male black or with abdomen further biological notes are given under that partly to mostly reddish ,ferruginea (Say) species. - Sheath stouter, about 3X longer than broad 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 30-32. Female ovipositors. 30, Entire lance of Macroxyela ,ferru@zea. 3 1, Entire lancet of M. ferru- ginea. 32, Entire lancet of M. aenea. Figs. 30-32 drawn to same scale.

(Fig. 13); radial region of forewing with 4 of sheath; apical margins of abdominal seg- cells, cell Rsl divided by a crossvein at center ments yellowish. Abdomen with metallic of cell (Fig. 6); both sexes concolorous black to bronze aenea (Norton) bronze tinge. Head and thorax finely punctate and sha- Mucroxyelu aeneu (Norton) greened; abdomen more shining with fine (Figs. 6, 13, 32, 37, 38) transverse microsculpture. Cell Rs 1 of fore- wing divided into two cells (Fig. 6). Xq’ela ueneu Norton 1872: 86.-Cresson 1928: 3. Male.-Length, 7.0-8.0 mm. Black, Mucroxyelu aenea: Kirby 1882: 351.- mostly like female, except thorax entirely Ross 1932: 165.-Ross 1951: 7.-Smith black and hypandrium pale orange. Geni- 1978: 23.-Smith 1979: 10. talia as in Fig. 37. Protoxyelu ueneu: MacGillivray 19 12: 296. Specimens examined.-32 0 and 8’. Distribution.-ILLINOIS: Pope Co., Female.-Length, 7.5-9.0 mm. Black Burden Falls, April 20, 1960; Dixon with antenna1 segments 1 and 2 dark or- Springs, IV-3-1948; Camp Point, Adams ange, antenna1 filament brownish, and with Co., IV-23-1960, on elm. INDIANA: Perry following usually orange brown to orange: Co., T4S RlW S36,21 April 1978. GEOR- supraclypeal area, clypeus, labrum, mouth- GIA: Jones Co., l-8 March 1994. LOUI- parts, anterior pronotum or all pronotum, SIANA: Tallulah. MISSISSIPPI: Clarks- mesoscutellum, sometimes mesosternum; dale, 3-10-55. MISSOURI: Webster Grove, legs (except hindtibia and hindtarsus usu- 3-26-21, 3-31-25. NORTH CAROLINA: ally blackish; sometimes venter of abdo- Orange Co., Chapel Hill, 8 Oct. 1963. men; and basal half or only apical margin OKLAHOMA: Latimer Co., in flight trap, VOLUME 100, NUMBER 4 651

111-1986, IV-1987, 111-1987, 111-1989; Still- Xyela infuscata Harris 1835: 583 (nomen water, 1948; Seminole Co., Wewoka, III- m&m).-Rohwer 1920: 515. 29-1966. TEXAS: types, no further data, as Xyela infuscata Norton 1862a: 224. well as several other specimens labeled Macroxyela infuscata: Kirby 1882: 351. “Tex.“; Harrison Co., 3 mi. E. Kamack, on Macroxyela obsoleta MacGillivray 19 12: pine, 6-III- 1972. WISCONSIN: Dane Co., 294.-Frison 1927: 234. April 14, 1915. Ross (1932) recorded Stat- Macroxyela bicolor MacGillivray 19 12: en Island, N.Y., a locality we were unable 294.-Frison 1927: 234.-Ross 1932: to verify. 166.-Ross 1951: 7 (syn.: var. harti Lectotype.-At ANSP, male, labeled Ross).-Smith 1978: 23.-Smith 1979: “Tex.” and red label “Type No. 332 Xyela 10. New synonymy. aenea Norton.” Norton described the spe- Macroxyela distincta MacGillivray 1912: cies from “Texas. Five specimens.” The 295.-Frison 1927: 234. one labeled as type is the lectotype as des- Macroxyela bicolor var. harti Ross 1932: ignated by Cresson (1928). 166. Host.-Unknown. Discussion.-The division of cell Rsl of Female.-Length, 7.5-9.0 mm. Mostly the forewing is more consistent than in M. reddish brown; clypeus, labrum, and mouthparts sometimes more yellowish, me- ferruginea. Of the 32 specimens examined, only 2 had this crossvein absent in one of sostemum usually black; in darker forms, the forewings and none in both forewings. antenna1 flagellum, head between eyes, and ocellar area blackish; mesonotal lateral Other distinguishing features of A4. aenea are the shorter, stouter female sheath (com- lobes, sometimes most or all of thorax ven- pare Figs. 12, 13), and the concolorous trally and laterally, basal 3 to 5 abdominal black to bronze coloration in both sexes segments, hindcoxa, hindtibia and apex of sheath sometimes black, with various com- (usually reddish brown in M. ferruginea). binations. Abdominal segments usually Nothing is known of the habits of A4. aenea. Records indicate a more southerly distri- with yellowish apical margins, especially noticeable on venter. Wings hyaline to very bution than for A4. ferruginea whose north- ernmost record is Madison, Wisconsin, al- lightly, uniformly infuscated, or rarely though there is substantial overlap between darkly infuscated. the species. Head and thorax finely shagreened; ab- domen more shining, with fine transverse microsculpture. Apical antenna1 filament 5- Macroxyela ferruginea (Say) (Figs. 2, 5, 12, 30, 31, 36, 38) 6 segmented, slightly shorter than scape. Forewing with cell Rsl not divided in most Xyela ferruginea Say 1824: 310. specimens. Macroxyela ferruginea: Kirby 1882: 351, Male.-Length, 7.0-8.0 mm. Mostly pl. 14, fig. I.-Young 1899: 41 (larva, black with following reddish-brown to or- habits; syn: infuscata Norton).-Mac- ange: most of clypeus, labrum and mouth- Gillivray 1906: pl. 23, fig. 33 (wings).- parts; most of legs beyond coxae (inner sur- Garlick 1923: 256-257 (adult feeding faces of fore and mid tibiae, sometimes tar- habits).-Yuasa 1923: 41’ (larva).-Ross si, and sometimes apical portion of hind- 1932: 165 (syn.: obsoleta MacGillivray, coxa paler); apical 3 or 5 abdominal distincta MacGillivray).-Smith et al. segments on venter and sometimes lateral- 1943: 382 (Kansas).-Ross 1951: 7.- ly; genitalia. Some forms with almost en- Maxwell 1955: 17 (internal larval anat- tirely orange abdomen. Genitalia as in Fig. omy).-Smith 1967: 379-380 (larva).- 36. Smith 1978: 23.-Smith 1979: 10. Specimens examined.462 9 and c?. 652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 100, NUMBER 4 653

. =Macroxyela ferrugin A dfacroxyela aenea

Fig. 38. Distribution of Macroxyela. Circles = M. ferruginea. Triangles = M. aenea.

Distribution.-Specimens examined manuscript list, was collected April 25, from the following states and provinces, 1827. (Fig. 38): Arkansas, Connecticut, Idaho Macroxyela bicolor MacGillivray was (Moscow), Illinois, Iowa, Kansas, Maine, described from a male, “Columbus, Ohio, Maryland, Massachusetts, Michigan, Min- Professor J.S. Hine, collector.” It is at nesota, Mississippi, Missouri, Montana INHS (Webb 1980), labeled “947” and (Lost Horse, Ravalli Co., 7-2-49), Nebras- “Columbus, Ohio J.S. Hine.” ka, New Hampshire, New York, North Car- Macroxyela obsoleta MacGillivray was olina, Ohio, Oklahoma, Ontario, Pennsyl- described from a female labeled “Ithaca, vania, TexasVirginia, West Virginia, Wis- N.Y., J.O. Martin, collector.” It is at INHS consin. (Webb 1980), labeled “Ithaca, N.Y., Apr. Types.-The type of Xyela ferruginea ‘97.” Say is lost. Say described the female and Macroxyela distincta MacGillivray was stated “Inhabits Arkansa.” The description described from a female and a male from agrees with the concept of this species. “Ithaca, N.Y, J.O. Martin, collector.” The Norton described Xyela infuscatu from a lectotype female, designated by Frison male, “One specimen in Harris’s Coll. (1927) is at INHS (Webb 1980), labeled Mass.” Rohwer (1920) stated the type is “Ithaca, N.Y., April 3, ‘97.” no. 186 in the Harris Collection (MCZ), is Macroxyela bicolor var. harti Ross, de- in good condition, and, according to Harris’ scribed from a male is at INHS (Webb

t

Figs. 33-37. Male genitalia, dorsal view, right half of genital capsule, and lateral view of penis valve with dorsal side to left. 33, Megaxyelu major. 34, M. bicoloripes. 35, M. tricolor. 36, Macroxyela,ferruginea. 37, M. aenea. 654 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1980), labeled “Havana, Illinois, April bud % up from the base and eat only the 1896 (CA. Hart).” central part, working down and enlarging Hosts.-Ulmus americana L. (Young the hole as necessary. They left the bud an 1899, Garlick 1923); on elm (Yuasa 1923). empty shell and so badly injured that no We collected adults from Ulmus carpinifol- further growth of the bud was possible. ia Gleditsch at the University of Virginia About 200 specimens were netted, the sex Blandy Experimental Farm and State Ar- ratio being 5 females to 1 male. Many of boretum of Virginia, Clarke Co.; very few these voucher specimens are in the collec- were on American elm, U. americana, and tion of the Snow Entomological Museum, some exotic elms in the same vicinity in the University of Kansas. Arboretum. Specimens collected in Malaise Smith et al. (1943) reported that this is a traps were in a nearby woodlot where slip- common in Kansas and occurs only pery elm, Ulmus rubra Mtihl., is present, in early spring in April and May. but we cannot confirm this as a host. Most NMS collected adults and observed them collection records from label data are from in abundance on three elms at the Beltsville elm or Ulmus sp. Agricultural Research Center, Prince Biology.-The description of the larva George’s Co., Maryland. Hundreds of in- and biological notes by Young (1899) are dividuals were netted while they flew from the first published on M. ferruginea. His the ground to the branches of the trees classical observation of the active pupa above. However, diligent searching of the phenomenon has not been reported since. ground over several weeks failed to find According to Young “Larvae sit curled “active pupae” as described by Young around the young leaves or terminal buds (1899). After continual observations, it be- with the front part of their body free. They came easy to distinguish M. ferruginea feed by eating a line across the bunch of from other insects because of their slow leaves, thus truncating them all. A single flight, shiny wings, and because M. ferru- brood each year, the adults appearing in ginea would land on the twigs of the elm April. A few days before the appearance of trees while other insects would hover near the adult the larva which has lain in the the branches, but seldom land. The elm larval state in its cocoon until this time trees where quite large (15 m) and only the changes to an active pupa, which bursts its lowest branches were attainable for collect- cocoon and comes forth without shedding ing even with an extended net handle (3 m). its skin and exists then as an active pupa at Nonetheless, examination of the upper the surface of the ground from twenty-four branches of the trees with binoculars re- to forty-eight hours. It then sheds its skin vealed thousands of adults on two trees in and becomes the true imago. Larvae enter 1994. the ground about June 1st.” He describes Specimens were netted during the fol- the cocoon as a light shell of sand held to- lowing periods: 8-14 April 1993, 31 gether by a few strands of silk. March-5 April 1994, and 16 March-15 Garlick (1923) reported on adult season- April 1995. During these periods, NMS ality and feeding habits at Lawrence, Kan- looked for individuals at least once a day. sas. Adults were abundant in spring, March Macroxyela ferruginea only flew when it 30 to April 24, especially the week of April was warm and sunny. On cold or windy 10-17. The first specimen was found on days, even in the middle of the flight peri- March 30, and the last on April 24. They od, no individuals were caught. Males fed on buds of American elm and to a lesser emerged before females in 1995, the first extent buds of pear and plum. The buds are male (followed by many others) was caught grown in length, but leaves hardly begun to on 15 March and the first female not until show. Adults gnaw a hole in the side of the 29 March. The sex ratio was notably biased VOLUMElOO.NUMBER4 655 toward females after 1 April. Collections in brown and are strongly violaceous at the 1993 and 1994 were not started prior to base. This is similar to the specimens with catching the first female specimens so pre- darkly infuscated wings as cited above. We sumably collections started in the middle of regard these as color variants; thus, their the flight period. Probably Garlick (1923) synonymy above. collected late in the flight period after males had flown. Sex ratio among (NMS) speci- A CKNOWLEDGMENTS mens (not necessarily a uniform sampling) We appreciate the loan of specimens is 124 m to 30 f (but at least 30 females from the following: D. Azuma, Academy of were ground up for allozyme studies and a Natural Sciences, Philadelphia, PA large number for sterol analysis). Rare in- (ANSP); R.W. Brooks, University of Kan- dividual males with red abdomens (former- sas, Lawrence; I? J. Clausen, University of ly M. bicolor) were collected occasionally Minnesota, St. Paul; S. Cover and G. Cha- in the middle of the flight period (first part varria, Museum of Comparative Zoology, of April), and, since there were no associ- Harvard University, Cambridge, MA ated females, we presume they are merely (MCZ); E. Diller, Zoologische Staatssam- a sexually dimorphic color form of M. fer- mlung, Mtinchen, Germany; H. Goulet, Ag- ruginea. riculture Canada, Ottawa; W.J. Hanson, Discussion.-Cell Rsl of the forewing is Utah State University, Logan; E.R. Hoe- not divided in most specimens examined. beke, Cornell University, Ithaca, NY; M.A. Of 461 specimens examined, 5 specimens Ivie, Montana State University, Bozeman; had this cell divided in both forewings and N. E Johnson, The Ohio State University, 26 specimens had this cell divided in one Columbus (OSU); B. Kondratieff and H.E. of the two forewings. In most instances the Evans, Colorado State University, Fort Col- division of this cell is not as uniform as in lins; B. Kopper, Kansas State University, M. aenea. The cell may be smaller, or the Manhattan; S.J. Krauth, University of Wis- crossvein dividing the cell may be partially consin, Madison; R.E. Lewis, Iowa State atrophied; the crossvein is not centrally lo- University, Ames; T. McCabe, New York cated as in M. aenea but toward the apex State Museum, Albany; K. Methven, Illi- of the cell. For other characteristics, see the nois Natural History Survey, Champaign key to species and discussion under M. (INHS); J.C. Morse and B. Robinson, aenea. Clemson University, SC; V.L. Moseley, About five specimens examined had rath- Louisiana State University, Baton Rouge; er darkly infuscated wings. They were from M.E O’Brien, University of Michigan, Ann Missouri, Ohio, and Illinois. About ten Arbor; PI? Parrillo, Field Museum of Nat- males examined had a red abdomen. Some ural History, Chicago, IL; E.G. Riley, Texas of these were collected in the same situation A&M University, College Station; T.L. and at the same trees as the typically-col- Schiefer, Mississippi State University, MS; ored black specimens, and we found no R.W. Sites, University of Missouri, Colum- structural differences between the two. We bia; M.E. Smethurst, American Museum of examined red abdomen specimens from Natural History, New York, NY; R.R. Maryland, Missouri, Illinois, Mississippi, Snelling, Natural History Museum of Los Ohio, and Pennsylvania. NMS collected Angeles County, Los Angeles, CA; E W. several red abdomen forms when making Stehr, Michigan State University, East Lan- large collections in Maryland. This red sing; K. Stephan, Lattimer Co., OK; W. W. form was described as M. bicolor by Middlekauff, University of California, MacGillivray (1912). Macroxyela bicolor Berkeley (UCB); D. Ubick, California var. harti Ross (1932) is also this red form, Academy of Sciences, San Francisco; D. but the wings are infuscated with deep Wahl, American Entomological Institute, 656 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Gainesville, FL; J.B. Whitfield, University Dyar, H. G. 1899. Larvae of Xyelidae. The Canadian of Arkansas, Fayetteville; and J.R. Wiley, Entomologist 3 1: 127. Frison, T H. 1927. A list of the types in the Florida State Collection of , collection of the Illinois State Natural History Gainesville, FL. Other specimens are in the Survey and the University of Illinois. Bulletin of National Museum of Natural History, the Illinois Natural History Survey 16: 137-309. Smithsonian Institution, Washington, DC Garlick, W. G. 1923. Notes on the feeding habits of (USNM). an adult sawfly. The Canadian Entomologist 55: We thank Michael Bowers for allowing 256-257. Goulet, H. and .I. T. Huber. 1993. Hymenoptera of the us to collect at the University of Virginia World: An Identification Guide to Families. Re- Blandy Experimental Farm and State Ar- search Branch, Agriculture Canada Publication boretum of Virginia, Clarke Co. Figures of 1894/E. 668 pp. the lances and lancets are by Linda Law- Harris, T W. 1835. Chapter VIII. Insects, pp. 553-602. rence, staff artist, Systematic Entomology In Hitchcock, E., Report on the Geology, Miner- alogy, Botany, and Zoology of Massachusetts. Laboratory. The habitus in Fig. 1 was Amherst, Massachusetts. drawn by John Heraty, University of Cali- Kirby, W. E 1882. List of Hymenoptera with Descrip- fornia, Riverside. We thank Terry Nuhn and tions and Figures of the Typical Specimens in the Cathy Anderson, Museum Technicians, British Museum, Vol. 1. and Siri- Systematic Entomology Laboratory, for cidae. London. 450 pp., 16 plates. Konow, E W. 1899. Ueber einige neue Chalastogastra. taking the photos and SEM’s and helping Wiener Entomologische Zeitung 18: 41-46. with the manuscript. We thank the follow- MacGillivray, A. D. 1906. A study of the wings of the ing for reviewing the manuscript: Hemi , a superfamily of Hymenoptera. Goulet, Agriculture Canada, Ottawa; W. W. Proceedings of the United States National Muse- Middlekauff, University of California, um 29: 569-654. MacGillivray, A. D. 1912. New genera and species of Berkeley. Xyelidae and Lydidae. The Canadian Entomolo- gist 44: 394-299. L ITERATURE C ITED Maxwell, D. E. 1955. The comparative internal larval anatomy of sawflies (Hymenoptera: Tenthredini- Ashmead, W. H. 1898. Table of genera of Xyelidae, dae). The Canadian Entomologist 87, Supplement pp. 214. In Dyar, H. G. Psyche 8: 214. 1, 132 pp. Bridwell, J. C. 1906. A second species of the hyme- Norton, E. 1862a. Descriptions of several of Harris’s nopterous genus Odontoph?jes Konow (Xyelinae). named Tenthredinidae. Proceedings of the Boston Entomological News 17: 94. Society of Natural History 8: 223-224. Burks, B. D. 1958. Symphyta, pp. 8817. In Krombein, Norton, E. 1862b. Notice of several new species of K. V., ed., Hymenoptera of America North of Mexico, Synoptic Catalog, First Supplement. Tenthredinidae. Proceedings of the Entomological United States Department of Agriculture, Agri- Society of Philadelphia 1: 143-144. culture Monograph 2. Norton, E. 1872. Notes on North American Tenthre- Cresson, E. T 1880. Descriptions of new North Amer- dinidae, with descriptions of new species. Trans- ican Hymenoptera in the collection of the Amer- actions of the American Entomological Society 4: ican Entomological Society. Transactions of the 77-86. American Entomological Society 8: l-52. Rasnitsyn, A. P 1969. Origin and Evolution of Lower Cresson, E. T. 1916. The Cresson Types of Hymenop- Hymenoptera. Trudy Paleontologischeskogo Insti- tera. Memoirs of the American Entomological So- tuta, Vol. 123, 195 pp. ciety I, 141 pp. Rohwer, S. A. 1920. Notes on the Harris Collection of Cresson, E. T 1928. The types of Hymenoptera in the sawflies, and the species described by Harris. Academy of Natural Sciences Other ‘Than Those Journal of the Washington Academy of Sciences of Ezra T Cresson. Memoirs of the American En- 10: 508-518. tomological Society 5, 90 pp. Rohwer, S. A. 1924. A new sawfly of the family Xye- Dyar, H. G. 1898a. Description of an unusual saw-fly lidae (Hymenoptera). Proceedings of the Ento- larva belonging to the Xyelinae. Psyche 8: 212- mological Society of Washington 26: 215. 214. Ross, H. H. 1932. The Hymenopterous family Xyeli- Dyar, H. G. 1898b. Notes on some sawfly larvae es- dae in North America. Annals of the Entomolog- pecially the Xyelidae. The Canadian Entomologist ical Society of America 25: 153-167, 2 plates. 30: 1733176. Ross, H. H. 1936. A new pecan sawfly, Meguqelu VOLUME 100, NUMBER 4 657

lungstoni n. sp. (Hymenoptera: Xyelidae). Ento- Karatavitidae, Gigasiricidae, Sepulcidae, Pseudo- mological News 47: 131-132. siricidae, , Siricidae, , Ross, H. H. 1951. Xyelidae, pp. 5-7. In Muesebeck, Paroryssidae, Xylotomidae, , Per- C. E W. et al., eds., Hymenoptera of America gidae). In van der Vecht, J. and R. D. Shenefeld, North of Mexico, Synoptic Catalog. United States eds., Hymenopterorum Catalogus, pars 14, 193 . Department of Agriculture, Agricultural Mono- PP. graph 2. Smith, D. R. 1979. Symphyta, pp. 3-137. In Krom- Sate, K. 1941. Ein dendro-entomologischer Beitrag. bein, K. V. et al., eds., Catalog of Hymenoptera in American North of Mexico, Vol. 1. Smithson- Bulletin of Agriculture and Forestry, College Sui- ian Institution Press, Washington D.C. gen 6, 235 pp. [In Japanese.] Smith, D. R. 1987. The sawfly work of H. Cl. Dyar Say, T 1824. Appendix. Part I.-Natural History. 1. (Hymenoptera: Symphyta). Transactions of the Zoology. E. Class Insecta, pp. 268-378. In Keat- American Entomological Society 112: 369-396. ing, W. H., Major Long’s Second Expedition, Vol. Smith, R. C., E. G. Kelly, G. A. Dean, H. R. Bryson, 2, 459 pp. pls. 6615. Philadelphia. and R. L. Parker. 1943. Common Insects of Kan- Schulz, W. A. 1906. Spolia Hymenopterologica. Pa- sas. Report of the Kansas State Board of Agricul- derborn, 355 pp. ture, June, 1943, Topeka, Kansas, 440 pp. Shinohara, A. 1992. The sawfly genus Megaqelu (Hy- Webb, D. W. 1980. Primary insect types in the Illinois menoptera, Xyelidae) in East Asia. Japanese Jour- Natural History Survey Collection, exclusive of nal of Entomology 60: 783-796. the Collembola and Thysanoptera. Illinois Natural Smith, D. R. 1967. A review of the larvae of Xyelidae, History Survey Bulletin 32: 55-191. with notes on the family classification (Hymenop- Young, C. 1899. Descriptions of sawfly larvae. The tera). Annals of the Entomological Society of Canadian Entomologist 3 1: 4 l-43. America 60: 376-384. Yuasa, H. 1923. A classification of the larvae of the Smith, D. R. 1978. Suborder Symphyta (Xyelidae, Par- Tenthredinidae. Illinois Biological Monographs 7, archexyelidae, Parapamphiliidae, Xyelydidae, 172 pp.