The Nearctic Species of Pandivirilia Irwin and Lyneborg (Diptera: Therevidae: Therevinae)

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1 SYSTEMATICS The Nearctic Species of Pandivirilia Irwin and Lyneborg (Diptera: Therevidae: Therevinae) DONALD W. WEBB 1 AND MARK A. METZ 2 Ann. Entomol. Soc. Am. 96(4): 369Ð402 (2003) ABSTRACT The genus Pandivirilia is widespread across the Nearctic and Palaearctic regions. The Nearctic species are redescribed, including the description of two new species: Pandivirilia constricta Webb and P. rufa Webb. Their phylogenetic position among the Palaearctic species and related higher therevine genera is elucidated by cladistic analysis of male and female morphological characters. A key to the Nearctic species is provided along with maps of their distribution. A neotype is designated for P. albifrons (Say) and a lectotype is designated for Psilocephala limata Coquillett. Also, Dichoglena melampodia (Loew), Ps. conspicua (Walker), and Spiriverpa nitoris (Coquillett) are placed as new combinations within Pandivirilia. P. bussi (James) is placed as a newcombination within Cliorismia, making this genus Holarctic in distribution. Ps. canadensis Cole, Ps. limata Coquillett, and Ps. pollinosa Cole are placed in synonomy under P. conspicua (Walker), P. argentifrons Cole is placed in synonomy under P. melampodia (Loew), and Thereva borealis Cole is placed in synonomy under P. albifrons (Say). KEY WORDS Nearctic revision, Dialineura, Incoxoverpa, Pallicephala IRWIN AND LYNEBORG (1981a) described the genera Pandivirilia, Viriliricta, and Dichoglena, but the placement and relationships among species within these genera have been enigmatic since that time. Pandivirilia (Irwin and Lyneborg 1981a: 212) was characterized as having males with eyes separated by a distance less than the width of the median ocellus, lacking a hypandrium, having elongate setae on the mesonotum, and lacking short, scattered macrosetae over the entire ventral surface of the hindfemur. Females have yellowish brown tibiae and the abdominal tergite four at least partially pruinose. They included the Nearctic species Pandivirilia argentifrons (Cole), P. bussi (James), P. limata (Coquillett), and P. pollinosa (Cole). Viriliricta (Irwin and Lyneborg 1981a: 208) was characterized as having males with eyes separated by a distance equal to or greater than the width of the median ocellus, lacking a hypandrium, having short, sparse setae on the mesonotum, and having many short, scattered black macrosetae on the entire ventral surface of the hindfemur. Females have dark yellowor reddish brown tibiae and abdominal tergite four is glossy or pruinose laterally. They included the Nearctic species Viriliricta canadensis (Cole), V. grandis (Johnson), and V. montivaga (Coquillett). Lyneborg (1986) synonomized Viriliricta with Pandivirilia but gave no justiþcation. Dichoglena (Irwin and Lyneborg 1981a: 210) was characterized as having males with eyes separated by a distance equal 1 Center for Biodiversity, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL Frost Entomological Museum, Department of Entomology, Agricultural Sciences and Industries, The Pennsylvania State University, University Park, PA to or greater than the width of the median ocellus, having a hypandrium, having elongate setae on the mesonotum, and lacking short, scattered macrosetae over the entire ventral surface of the hindfemur. Females were characterized as having black or blackish brown tibiae and having the abdominal tergite four entirely shiny black. They included in this genus the Nearctic species Dichoglena amplifrons (Cole), D. borealis (Cole), D. melampodia (Loew), and D. nigrina (Kröber). In this article we attempt to clarify the characteristics deþning Dichoglena and Pandivirilia; to update the generic placement of species that Irwin and Lyneborg had placed in Pandivirilia, Viriliricta, and Dichoglena; and to test LynenborgÕs hypothesis of the synonymy of Viriliricta under Pandivirilia with minimum length tree parsimony analysis. This article is the seventh of a series (Webb and Irwin 1988, 1991aÐc, 1995, 1999) revising the species of the genera treated in the monograph of the nearctic genera of Therevidae (Irwin and Lyneborg 1981a). The phylogenetic position of the Nearctic species of Pandivirilia relative to the Palaearctic species and related higher therevine genera are elucidated by a cladistic analysis of male and female morphological characters. Keys to the Nearctic species are provided, along with species descriptions or redescriptions and distributions. Materials and Methods The morphological terminology used for the male terminalia was deþned by Lyneborg (1968) and subsequently modiþed by Lyneborg (1972, 1976, 1978) /03/0369Ð0402$04.00/ Entomological Society of America

2 370 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 and Irwin (1977a, 1977b, 1976) deþned that used for the female terminalia. The terminology for body vestiture follows Winterton et al. (1999), and for other morphological features follows McAlpine (1981). Setae described as elongate have a length equal to or greater than the width of the scape; those described as short have a length less than the width of the scape. Macrosetae on the thorax (np notopleural, sa supraalar, pa postalar, dc dorsocentral, sc scutellar) were counted from the left half of the specimen. Anteroventral (av) and posteroventral (pv) macrosetae are presented as counts (0:0:0) for the fore-, middle-, and hindfemora. Generally, each specimen was assigned a unique number, MEI#. This number is pinned with the specimens and is used to associate the ecological and label data with a given specimen, and when referred to herein, consist of three letters, MEI followed by a six-digit number. Some lending institutions pinned their own unique number to their specimens and these were used for those specimens. Specimen data were recorded into a database within the architecture of MANDALA (Kampmeier et al. 1998) and can be viewed and queried via the Web site inhs.uiuc.edu/index.htm. To conserve space and include as much information as possible about each specimen, a format adopted from Irwin (1983) is used in the Specimens Examined section of each species. Stuckenberg and Irwin (1973) deþned many of the terms used therein. In the presentation of repetitive locality data, a semicolon terminates one series of specimens and signals the beginning of the next; thus, data not repeated in the series are the same as those of the preceding series. In the discussion of the cladistic analysis, references to characters and character states are in parenthesis (1:0) with the character preceding the colon and the speciþc character state following the colon. The following institutions were kind enough to loan material relevant to this study: D. Azuma, Academy of Natural Sciences of Philadelphia (ANSP); D. A. Grimaldi, American Museum of Natural History (AMNH); M. Cazier, Arizona State University (ASUT); L. G. Bezark Collection, (BEZC); N. Evenhuis, Bernice P. Bishop Museum (BPBM); C. R. Nelson and R. W. Baumann, Brigham Young University (BYUC); N. D. Penny, California Academy of Science (CASC); E. Fisher, California Department of Food and Agriculture, State Collection of Arthropods (CDAE); H. J. Teskey, D. M. Wood, J. M. Cumming, Canadian National Collection (CNCI); C. Young, Carnegie Museum of Natural History (CMNH); J. Keiper, Cleveland Museum of Natural History (CEMU); B. C. Kondratieff, Colorado State University (CSUC); C. T. Maier, Connecticut Agriculture Experiment Station (CAES); L. L. Pechuman, E. R. Hoebeke, Cornell University (CUIC); H. V. Weems, Jr., G. J. Steck, Florida State Collection of Arthropods (FSCA); Illinois Natural History Survey (INHS); Michael E. Irwin Collection, (MEI); D. Blocker, Kansas State University (KSUC); C. T. Maier Collection, (CTMA); B. Brown, Los Angeles County Museum (LACM); F. Stehr, Michigan State University (MSUC); Montana State University, MT Entomological Collection, M. Ivie, (MTEC); D. Furth, S. Shaw, Museum of Comparative Zoology, Harvard University (MCZC); J. C. Chainey, The Natural History Museum (NHM(E)); R. C. Bechtel, Nevada State Department of Agriculture (NVDA); J. K. Barnes, NewYork State Museum (NYSM); R. Blinn, North Carolina State University (NCSU); N. Johnson, Ohio State University (OSUC); W. A. Drew, Oklahoma State University (OSEC); R. L. Westcott, Oregon State Department of Agriculture (ODAC); J. Lattin, Oregon State University (OSUO); L. E. Munstermann, Peabody Museum of Natural History, Yale University (PMNH); K. Valley, Pennsylvania Department of Agriculture (PADA); K. C. Kim, Pennsylvania State University, Frost Entomological Museum (PSUC); A. Provonsha, Purdue University (PURC); R. A. Cannings, Royal British Columbia Museum (RBCM); B. Hubley, G. B. Wiggins, Royal Ontario Museum (ROME); San Diego Natural History Museum (SDMC); J. Macpherson, Southern Illinois University (SIUC); A. Freidberg, Tel Aviv University (TAUI); E. G. Riley, Texas A and M University (TAMU); L. Cloutier, Université de Montréal (QMOR); F. C. Thompson, United States National Museum (USNM); S. G. Cannings, University of British Columbia, Spencer Entomological Museum (SMDV); C. B. Barr, J. Powell, University of California, Berkeley/California Insect Survey Collection (EMEC); S. Heydon, University of California, Davis (UCDC); S. Frommer, University of California, Riverside (UCRC); C. L. Smith, University of Georgia (UGCA); S. A. Marshall, University of Guelph (DEBU); F. W. Merickel, University of Idaho, W. F. Barr Entomological Museum (WFBM); G. W. Byers, University of Kansas, SnowEntomological Museum (SEMC); S. Wood, University of Maine (UMDE); M. F. OÕBrien, University of Michigan, Museum of Zoology (UMMZ); P. J. Clausen, University of Minnesota (UMSP); K. B. Simpson, University of Missouri (UMRM); D. S. Chandler, University of NewHampshire (DENH); C. R. Nelson, University of Texas (UTAC); R. T. Bell, University of Vermont (UVIC); University of Wisconsin Entomological Museum (UWEM); S. R. Shaw, R. Lavigne, University of Wyoming (ESUW); W. J. Hanson, Utah State University (EMUS); W. J. Turner, R. S. Zack, Washington State University, James Entomological Collection (WSUC). The ambiguous relationships among these genera and their placement in the subfamily Therevinae required an extensive taxon sampling. To clarify the status and relationships between Pandivirilia, Viriliricta, and Dichoglena, the putataive ingroup included the nearctic species of the three genera, Þve Palaearctic species of Pandivirilia [P. amurensis Lyneborg, P. caesia Meigen, P. exima Meigen, P. isolata Lyneborg, and P. melaleuca (Loew)], and the Palaearctic species of Dichoglena [D. nigripennis (Ruthe)]. The putative outgroup was estimated to be those species with probable close afþnities to Pandivirilia and Dichoglena and included 13 species from six genera: four species of Cliorismia [C. ardea (Fabricius), C. latiphalangis Na-

3 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 371 gatomi and Lyneborg, C. rustica (Panzer), Cliorismia n. sp.], two species of Dialineura [D. anilis (Linnaeus) and D. gorodkovi (Zaitzev)], Incoxoverpa [I. borealis (Cole)], two species of Pallicephala [P. slossonae (Coquillett) and P. pachyceras (Williston)], two species of Spiriverpa [S. albiceps (Cole) and S. senex (Walker)], and two species of Tabuda [T. planiceps (Loew) and T. varia (Walker)]. The analysis was performed to determine the relationships among the various species of Pandivirilia and the outgroup genera and was rooted from Incoxoverpa, which was recently treated (Webb and Irwin 1998) and determined to be monotypic, but closely associated with the other taxa in the analysis. Character Descriptions Thirty characters were used in this analysis. Most characters are binary, but characters 3, 5, 18, 28 each have three or more states. For species scored without an available female specimen, female characters were coded as missing data?. Head 1. Ommatidia of male compound eyes (0) of equal size, (1) smaller ventrally. 2. Distance between male eyes (0) less than width of median ocellus, (1) equal to or greater than width of median ocellus. 3. Pruinescence on dorsal half of female frons (0) present, brown to black, (1) absent, glossy brown to black cuticle exposed, (2) present, silver. 4. Setae on ventral half of female frons (0) present, (1) absent. 5. Black pruinescent band lateral to antennal sockets, both sexes (0) absent, (1) present, blending into frons, (2) present, isolated from frons. 6. Parafacial setae, male (0) present, (1) absent. Thorax 7. Setae in and around prosternal depression, both sexes (0) present, (1) absent. 8. Setae on anterior half of proepimeron, both sexes (0) present, (1) absent. 9. Setae on posterior half of proepimeron, both sexes (0) present, (1) absent. Wings 10. Wing cell m 3, both sexes (0) widely open at apex, (1) closed, often petiolate. Legs 11. Male forefemur with anteroventral macrosetae (0) absent, (1) present. 12. Forefemur of female with posteroventral macrosetae, (0) absent, (1) present. 13. Midfemur of male with short, scattered macrosetae (0) absent, (1) present. 14. Hindfemur of male with short scattered macrosetae (0) absent, (1) present. Male Terminalia 15. Male epandrium in lateral view(0) oblong, tapered posteriorly, (1) posterior half expanded ventrally (see Figs. 9Ð15). 16. Posteroventral area of male hypoproct (0) ßat, (1) expanded ventrally. 17. Male hypandrium (0) present, (1) absent. 18. Male inner gonocoxal process (0) short, bluntly pointed, (1) narrow, elongate, tapered posteriorly (see Figs. 16Ð22), (2) enlarged posteriorly. 19. Male dorsal apodeme of aedeagus with sides broad, rounded (0) absent, (1) present (see Figs. 38, 40, 42, and 43). 20. Sclerotized area of anterior margin of male dorsal apodeme of aedeagus strongly bifurcate (0) absent, (1) present. 21. Male dorsal apodeme of aedeagus narrow, diamond shaped, (0) absent, (1) present. 22. Male dorsal apodeme of aedeagus narrow, subrectangular, posterior corners obtuse, (0) absent, (1) present (see Figs. 37, 39, and 41). 23. Male dorsal apodeme of aedeagus broad, square (0) absent, (1) present. 24. Male dorsal apodeme of aedeagus reduced, rectangular, posterior corners strongly angulate (0) absent, (1) present. 25. Male distiphallus of aedeagus, lateral view(0) short, broad basally (see Figs. 51Ð57), (1) elongate with a narrow base. 26. Posterior end of male aedeagal ejaculatory apodeme (0) slightly bulbous (see Figs. 37Ð39) and 41), (1) distinctly enlarged (see Figs. 40, 42, and 43). 27. Anterior half of male aedeagal ejaculatory apodeme (0) distinctly enlarged laterally, (1) cylindrical, slightly bulbous (see Figs. 37Ð43). 28. Male sclerotized areas of ejaculatory process (0) split into bilaterally paired discs, (1) crescent shaped (see Figs. 37Ð43), (2) absent. Female Terminalia 29. Setae on medial lobe of female tergite 9 10 (0) present, (1) absent. 30. Posterior fourth of female spermathecal ducts (0) narrow, smooth, (1) broad, crenulate (see Figs. 58Ð63). The character matrix used in the cladistic analysis is given in Table 1. All characters were treated as unordered. Cladistic Analysis The data were analyzed with PAUP* 4.0b10 (Swofford 2002) running a heuristic search with 1,000 random addition sequence replicates and three trees held at each step during stepwise addition. Rearrangements were made by the method of tree bisection and reconnection and the steepest descent option was in effect. The search resulted in 18 equally parsimonius trees (CI 0.425, RI 0.776, RC 0.330) of 80 steps. Characters were successively reweighted by the max-

4 372 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 Fig. 1. (a) The strict consensus cladogram of the 18 most parsimonious unweighted trees (see text for tree statistics). The circles indicate the character state changes under UN- AMBIGUOUS character optimization. Characters are numbered as in the text with hash marks as follows: black forward change with no homoplasy; white forward change with homoplasy or reversal. Genus names in parentheses indicate the previous generic assignment of the species. (b) Bremer support displayed on a strict consensus of the 18 most parsimonius unweighted trees resulting from TBR branch swapping with all characters assigned weight 1.

5 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 373 Figs Male tergite 8, epandrium, cerci, and hypoproct, dorsal view: (2) P. albifrons. (3) P. conspicua. (4) P. constricta. (5) P. grandis. (6) P. melampodia. (7) P. montivaga. (8) P. rufa. Scale 0.1 mm. C, Cercus; Epa, epandrium; Hpt, hypoproct; T 8, tergite 8. imum value of the rescaled consistency indices until tree lengths stabilized after two repetitions. Successive reweighting resulted in three equally parsimonious trees (CI 0.626, RI 0.880, RC 0.550) that were a subset of the original 18 trees of 80 steps. The strict consensus trees resulting from unweighted and successively reweighted analyses were completely congruent in regards to generic monophyly with the latter providing increased resolution among the species within the genera Cliorismia and Pandivirilia. The program Winclada (Nixon 2002) was used to map character changes. Taxonomic changes proposed herein are based on the successively reweighted strict consensus cladogram (Fig. 1a). Decay indices (Bremer 1988, 1994) were calculated using SEPAL (Salisbury 2000) and are shown on the unweighted consensus tree in Fig. 1b The analysis supported the monophyly of Dichoglena based on the synapomorphy of the male dorsal apodeme being reduced, rectangular, and with distinct angular corners (24:1). In addition, this clade is supported by the dorsal half of the female frons being dark reddish brown and glossy (3:1); setae being absent on the ventral half of the female frons (4:1); a black pruinescent band with diffuse margins extending from the base of the antennal sockets to the eyes (5:1); and the median lobe of female tergite 9 10 lacking setae (29:1). The analysis also supported the removal of Dichoglena borealis (Cole) [ P. albifrons (Say)] (sensu Irwin and Lyneborg 1981a: 211) and D. melampodia (Loew) (sensu Irwin and Lyneborg 1981a: 212) from this genus and their placement into the genus Pandivirilia. Dichoglena amplifrons (Cole) (sensu Irwin and Lyneborg 1981a: 211) has been determined to be a junior synonym of D. nigrina. This leaves Dichoglena with only a single Nearctic species, D. nigrina. The analysis supported the placement of P. bussi (sensu Irwin and Lyneborg 1981a: 214) and an undescribed newspecies into the genus Cliorismia based on the synapomorphy of the male inner gonocoxal process being enlarged and clavate apically (18:2). In addition this clade is supported by the absence of a black pruinescent band lateral to the antennal sockets in both sexes (5:0); the wing cell m 3 being closed (10:1); and the posteroventral area of the male hypoproct being expanded ventrally (16:1). The placement of these two Nearctic species within Cliorismia adds a newgenus of therevids to the Nearctic fauna as Cliorismia nowbecomes Holarctic in distribution. The analysis also supports the monophyly of Pandivirilia based on the synapomorphy of the male distiphallus in lateral viewbeing short and having a broad base (25:0). In addition, this clade is supported by the absence of a black pruinescent band lateral to the antennal sockets (5:0) (reversed to state one in P. isolata and to autapomorphic for state two in P. amurensis); prosternum generally lacking setae (7:1); the male epandrium in lateral viewwith posteroventral expansion (15:1); male hypandrium absent (17:1); the male ejaculatory process being narrowand crescent shaped (28:1); and the basal portion of the female spermathecal ducts broad, crenulate (30:1). This clade has strong Bremer support (Fig. 1b) and justiþes

6 374 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 Figs Male epandrium, lateral view: (9) P. albifrons. (10) P. conspicua. (11) P. constricta. (12) P. grandis. (13) P. melampodia. (14) P. montivaga. (15) P. rufa. (16Ð22) Male gonocoxite, lateral view. (16) Pandivirilia albifrons. (17) P. conspicua. (18) P. constricta. (19) P. grandis. (20) P. melampodia. (21) P. montivaga. (22). P. rufa. Scale 0.1 mm. IGP, inner gonocoxal process; OGP, outer gonocoxal process. LyneborgÕs synonomy of Viriliricta and Pandivirilia and the placement of D. borealis (Cole) [ P. albifrons (Say)] (sensu Irwin and Lyneborg 1981a: 211) and D. melampodia (Loew) (sensu Irwin and Lyneborg 1981a: 212) within Pandivirilia. Taxonomy With the synonomy of the genus Viriliricta by Lyneborg (1986) and the placement of D. melampodia within Pandivirilia, couplets 17Ð19 (Irwin and Lyneborg 1981a: 202) and couplet 18Ð20 (Irwin and Lyneborg 1981b: 519) in the keys to the genera of Nearctic Therevidae need to be changed. The following key is a modiþcation of these couplets with couplets 17Ð18 for Irwin and Lyneborg (1981a) and couplets 18Ð19 in parentheses for Irwin and Lyneborg (1981b). 17 (18). Face with black pruinescent band extending from antennal sockets to margin of eyes; male dorsal apodeme rectangular, posterolateral corners distinctly rightangled; female with median lobe of tergite 9 10 glabrous...dichoglena Face lacking black pruinescent band extending from antennal sockets to margin of eyes; male dorsal apodeme with posterolateral corners obtuse, not distinctly right-angled; female with dark reddish brown setae on median lobe of tergite (19) 18 (19). Parafacial setae present, extending lateral to antennal sockets. Male with hypandrium; with anterior half of ejaculatory apodeme distinctly enlarged; with ejaculatory process triangular; and apex of male distiphallus turned laterally; female spermathecal ducts narrow, smooth...spiriverpa Parafacial setae absent (exception: P. albifrons, which has setae on the anterior half of the proepimeron). Male without

7 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 375 Figs Male gonocoxite, dorsal view: (23) Pandivirilia albifrons. (24) P. conspicua. (25) P. constricta. (26) P. grandis. (27) P. melampodia. (28) P. montivaga. (29) P. rufa. Scale 0.1 mm. GA, gonocoxal apodeme. hypandrium; anterior half of ejaculatory apodeme cylindrical; ejaculatory process narrow, crescent shaped; apex of distiphallus directed posteriorly; female with basal portion of spermathecal ducts broad, rugose... Pandivirilia Pandivirilia Irwin and Lyneborg Pandivirilia Irwin and Lyneborg (1981a: 212, desc.): Irwin and Lyneborg (1981b: 522, key), Lyneborg (1986: 85, key, desc.). Type species: Psilocephala limata Coquillett (1894:99) by original designation (Ir- Table 1. Character matrix for the genera Cliorismia, Dialineura, Dichoglena, Incoxoverpa, Pallicephala, Pandivirilia, Spiriverpa, and Tabuda. Characters are described in the text Pandivirilia albifrons Pandivirilia amurensis 10?? ? ??? Pandivirilia caesia Pandivirilia conspicua Pandivirilia constricta nsp. 10?? ? ?? Pandivirilia exemia Pandivirilia grandis Pandivirilia isolata 00?? ? ?? Pandivirilia melaleuca ?? Pandivirilia melampodia Pandivirlia montivaga Pandivirilia rufa nsp Cliorismia ardea Cliorismia bussi Cliorismia latiphalangis Cliorismia rustica Cliorismia n.sp Dialineura anilis Dialineura gorodkovi Dichoglena nigrina Dichoglena nigripennis Incoxoverpa borealis Pallicephala slossonae Pallicephala pachyceras Spiriverpa albiceps Spiriverpa senex Tabuda planiceps Tabuda varia

8 376 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 win and Lyneborg 1981a: 212) ( Thereva conspicua Walker 1848:223). Type locality: Nova Scotia. Viriliricta Irwin and Lyneborg (1981a: 208, key, desc.): Irwin and Lyneborg (1981b: 522, key), Lyneborg (1986: 85, desig. syn.). Type species Ps. montivaga Coquillett (1893:226) by original designation. Type locality: California, Los Angeles County. Diagnosis of Male and Female. Medium- to largesized stiletto ßies (7Ð17 mm). Head. Ocellar tubercle dark reddish brown, pruinescence gray; setae dark reddish brown. Eye glabrous; male eyes variable from almost contiguous medially to being separated by a distance twice the width of the median ocellus; female dichoptic; median margin sinuate; male ommatidia generally of similar size although in some males they may grade ventrally into a smaller size. Female frons with dorsal two-thirds reddish brown, pruinescence light brownish gray, ventral third with silver pruinescence. Antenna shorter than head; scape cylindrical, equal to or narrower than width of ßagellum, apical macrosetae dark brown; pedicel lobate; ßagellum three-segmented, awlshaped basally then tapered apically to two cylindrical ßagellomeres ending with a minute apical spine. Male frons generally reduced, triangular, female frons broad, lateral margins convergent dorsally, dorsal half generally dark, ventral half with silver pruinescence. Parafacial with silver pruinescence. Maxillary palpus one-segmented, cylindrical to slightly clavate, rounded apically. Setae elongate on scape, short on pedicel; elongate on ocellar tubercle, gena, and maxillary palpus, absent on eyes, and clypeus. Thorax. Postpronotal lobe generally concolorous with thorax. Vittae variable. Setae color variable, elongate on propleuron, anepisternum, scutellum, and laterotergite, and anterior half of katepisternum; absent on meron and anepimeron. Wing. Hyaline to opaque; veins brown, occasionally with brown margin; pterostigma brown; setulae absent; cell m 3 open; discal cell acute basally; cell cup generally closed with short petiole, occasionally closed at wing margin; m-cu/r-m subequal. Legs. Coxae moderately long; apical macrosetae dark brown. Setae present on posterior surface of middlecoxa. Femur with white Þliform setae, becoming lanceolate and appressed on dorsal surface. Abdomen. Equal to width of thorax basally, tapering posteriorly. Male Terminalia. Tergite eight bilobed, posterior margin deeply emarginate (Figs. 2Ð8). Sternite eight (see Figs. 30Ð36) reduced, posterior margin with shallowemargination. Epandrium (Figs. 2Ð8) quadrate, median length subequal to width. Cercus (Figs. 2Ð8) with apex rounded, ending before or slightly beyond hypoproct. Hypoproct (Figs. 2Ð8) generally bilobed, generally ending belowposterior margin of cerci. Hypandrium (see Figs. 23Ð29) absent. Gonocoxite (see Figs. 23Ð36) bulbous, sides generally rounded; in ventral viewseparated medially; gonocoxal apodeme reduced; inner gonocoxal process (see Figs. 16Ð22) attenuated posteriorly, pointed apically, extending well beyond outer gonocoxal process. Ventral lobe (see Figs. 30Ð36) large, generally ßattened laterally. Gonostylus (see Figs. 23Ð29) broad, sinuate, upturned dorsally. Aedeagus (see Figs. 37Ð43) with ejaculatory process narrow, crescent-shaped. Female Terminalia (see Figs. 58Ð63). Tergite eight subrectangular, subequal in length and width; anterior margin projecting anteriorly. Tergite 9 10 fused, with strong acanthophorite macrosetae, median lobe of tergite 9 10 with setae. Cercus triangular, membranous, with numerous Þne, short setae. Sternite eight large, longer than wide; posterior emarginate with sclerotized medial penis guide. Sternite nine greatly modiþed, invaginated above sternite eight to form internal sclerotized furca which is closed anteriorly; ventral surface of furca covered with lightly sclerotized membrane attaching it to the penis guide. Sternite 10 membranous, subtriangular with short, thick spines or setae. Internal reproductive organs with short common duct; the two spermathecal ducts separate at posterior junction with medial spermathecal sac, posterior third of spermathecal duct broad, rugose. Immature Stages. Cole (1923: 59, Fig. 173) described the pupa of P. conspicua (as Ps. limata). Seasonal Activity and Distribution. Specimens of Pandivirilia have been collected from 1 March to 17 September. Specimens have been hand collected in a damp sphagnum fen, along the side of a stream, and on Asclepias sp., Encelia sp., Prunus serotina Ehrh., Sarcobatus vermiculatus (Hook.) Torr.; by Malaise traps in the oak-chaparral zone or in a Þeld near a dry oak forest; in a ßume debris trap; and by black lighting. Pandivirilia has been collected in the Nearctic region from Florida to Nova Scotia west to eastern Texas and eastern North Dakota and from California to Alaska east to NewMexico and Alberta. Key to Nearctic Species of Pandivirilia 1. Prosternal setae white, elongate P. melampodia (Loew) (in part) Prosternal setae absent Parafacial setae present...3 Parafacial setae absent Male eye with ommatidia of equal size; maxillary palpus without circular basal constriction P. albifrons (Say) Male eye with ommatidia smaller on ventral third; maxillary palpus with circular basal constriction... P. constricta Webb n. sp. 4. Gena with small patch of short dark reddish brown setae anteriorly P. melampodia (Loew(in part) Gena lacking small patch of short dark reddish brown setae anteriorly Femora dark reddish brown to black...6 Femora yellowto dark yellow Tibiae dark yellow, apex dark reddish brown; male with mesonotal setae elongate; male hy-

9 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 377 poproct with posteroventral area ßat; setae in band across proepimeron P. conspicua (Walker) Tibiae dark reddish brown, concolorous with femora and tarsi; male with mesonotal setae short; male hypoproct with posteroventral area projecting ventrally; setae absent on proepimeron...p. grandis (Johnson) 7. Thorax black, abdomen dark reddish brown to black...p. montivaga (Coquillett) Thorax orangish brown; abdomen orange P. rufa Webb n. sp. Cliorismia bussi (James), new combination bussi James (James and Huckett 1952: 265, desc.), Cole (1965: 351, cat.) (Psilocephala), Irwin and Lyneborg (1981a: 214, listing) (Pandivirilia). This species was placed in Pandivirilia by Irwin and Lyneborg (1981a), but in our phylogenetic analysis it shared a suite of characters which placed it in a clade containing species of the Palaearctic genus Cliorismia. On the basis of this phylogenetic analysis, we are placing P. bussi in combination with Cliorismia. Pandivirilia albifrons (Say), new combination albifrons Say (1829: 156, desc.), (Cole 1923: 102, desc.; 1965: 352, cat.) (Thereva); Irwin and Lyneborg (1981a: 216, listing, comb. change) (Spiriverpa). Type; NT in INHS. borealis Cole (1923: 126, key, desc.; 1965: 353, cat.) (Thereva); Irwin and Lyneborg (1981a: 211, comb. change) (Dichoglena). Type; HT in CUIC. New Synonym. nitoris Coquillett (1894: 224, desc.), Cole (1923: 126, key;1965:354, cat.) (Thereva): Irwin and Lyneborg (1981a: 216, listing, comb. change) (Spiriverpa). Type: HT in USNM. The holotype female of T. nitoris (USNM Type Number 993) was compared with the holotype female of T. borealis (CUIC Type Number 4352) and determined to be conspeciþc based on the presence of white elongate setae on the parafacial and the anterior half of the proepimeron. Irwin and Lyneborg (1981a: 216) placed T. nitoris in the genus Spiriverpa, but males of this species do not have the apex of the distiphallus bent laterally, a synapomorphic characteristic of Spiriverpa. Say (1829: 156) described T. albifrons, which he stated resembled T. frontalis Say, although being much smaller. Although Say did not state that T. albifrons had setae on the face, the authors are assuming that this condition was present on T. albifrons because species of Thereva at the time of SayÕs description had the face covered with setae unless otherwise stated by Say. Say also stated that T. albifrons was black with a blackish vitta, the halteres (poisers) black at tip, and being 7.6 mm [three-tenths of an inch] long. The only species of Thereva currently known from Indiana is T. frontalis which is a large brownish yellow species 12.7 mm long with two yellow vittae, dark halteres, and having the abdominal terga with yellow annulations [bands]. Irwin and Lyneborg (1981a: 216) placed albifrons in Spiriverpa. The only species of Spiriverpa currently known from the midwest is S. senex (Walker) but it has the halter knob pale yellow. Acrosathe has setae on the face, but the only eastern species of this genus is A. bimaculata (Cole) from North Carolina. To clarify the taxonomic status of T. albifrons, a neotype is designated for this species. Upon his death, Thomas SayÕs insect collection was given to the Philadelphia Academy of Natural Sciences (Mallis 1971) and was destroyed by dermestid beetles and the transportation by stagecoach when it was shipped to Dr. T. W. Harris in Cambridge. Say described T. albifrons on the basis of a male specimen from Indiana. Currently, no specimens of this taxon are available from Indiana, so a male specimen from Vermilion County, IL, has been selected for the neotype and is deposited in the collection of the Illinois Natural History Survey. During this reviewof the Nearctic species of Pandivirilia, D. borealis (Cole) (Irwin and Lyneborg 1981a: 216) was noted to have setae on the parafacial. This species has been determined to be conspeciþc with Spiriverpa nitoris (Coquillett) (Irwin and Lyneborg 1981a: 216) which herein is placed in the genus Pandivirilia. This species is widespread in eastern North America. The males have dark brown halter knobs and are 7.6 mm in length. The characteristics of size, halter color, presence of white parafacial setae and distribution provides the best evidence for synonomizing nitoris under SayÕs species albifrons. In the key to the Nearctic genera of Therevidae (Irwin and Lyneborg 1981a) the males of P. albifrons would fall into the genus Spiriverpa based on the white frontal and parafacial setae, even though this species does not possess a hypandrium, nor is the apex of the distiphallus bent laterally. Irwin and Lyneborg (1981a: 216) placed T. nitoris in Spiriverpa. An examination of the holotype female determined that it differs from Spiriverpa in having setae present on the anterior half of the proepimeron and absent on the posterior half; in having posteroventral macrosetae on the forefemur; in having the female tergite four glossy without lateral or posterior pruinescence; and in having the basal fourth of the female spermathecal ducts broad and rugose. Diagnosis. This species is similar to P. constricta in having white elongate setae on the parafacial. It is easily separated from P. constricta by the male ommatidia being of equal size; in lacking a basal circular constriction on the maxillary palpus; the forefemur lacking anteroventral macrosetae; and the distiphallus (see Fig. 51) in lateral viewbroad basally and projecting posteriorly at a 45 angle. Male. Description of Neotype (MEI ). Body length 8.9 mm. Head. Length 1.08 mm. Ocellar tubercle black, pruinescence gray; setae black, elongate. Eyes separated by distance one-tenth width of median ocellus; ommatidia of equal size. Frons with silver pruinescence ventrally, becoming grayish black dorsally; setae

10 378 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 white, elongate on ventral half, extending lateral to antennal sockets. Antenna dark brown, pruinescence dense gray on scape and pedicel, 0.62 times length of head; scape 0.38 mm long, 1.2 times width of ßagellum, 2.2 times longer than wide, 3.8 times length of pedicel; pedicel 0.10 mm long; Þrst ßagellomere oval, tapered apically to second ßagellomere, length 0.30, width 0.14, ßagellum 2.9 times longer than wide, 2.4 times length of scape; setae white, elongate on scape, black short on pedicel and dorsobasally on Þrst ßagellomere. Parafacial broad, 0.94 times width of clypeus; covered with whitish gray pruinescence; setae white, elongate. Genal setae white. Maxillary palpus dark reddish brown, pruinescence gray; length 0.52 mm, 6.5 times longer than wide; setae white, elongate. Thorax. np 4 sa 2 pa 1 dc 1, sc 2. Thorax dull black, vittae separated by narrowgray stripe; white elongate on mesonotum, postpronotal lobes propleuron, anepisternum, katepisternum, laterotergite, scutellum, metanepisternum, and anterior half of proepimeron, absent on prosternum. Wing. Length 7.0 mm, 2.0 times longer than wide. Membrane pale gray, with faint trace of brown margin on veins and crossveins; pterostigma brown. Halter dark brown. Cell m 3 open widely. Legs. Coxae dark reddish brown, pruinescence gray; setae white elongate, present on posterior area of middlecoxa; apical macrosetae dark reddish brown 2:2:1; papillate projection present on hindcoxa. Femur dark reddish brown, pruinescence gray; anteroventral macrosetae 1:0:3, posteroventral macrosetae absent; setae white elongate on ventral half, becoming lanceolate, appressed on dorsal half. Tibia and tarsi reddish brown, apex of tibiae darker. Abdomen. Black, tergites 2Ð8 with white pruinescence; dorsal and lateral setae white, elongate. Terminalia from (MEI ). Epandrium (Fig. 2) quadrate, dark brown, posterior margin pale yellow, glossy, glabrous, posterolateral corners attenuate, broadly pointed; in lateral view(fig. 9) dark brown with apical half expanded ventrally to form pale yellowblade. Hypoproct with posteroventral area ßat. Gonocoxite dark brown (Figs. 23 and 30); inner gonocoxal process (Fig. 16) pale yellow, apical setae pale yellow; outer gonocoxal process (Fig. 16) broadly pointed, apical setae dense. Gonostylus (Figs. 23 and 30) broad, sinuate; setae yellowish brown on dorsallly, ventrally and along mediolateral margin. Aedeagus (Fig. 37) with dorsal apodeme quadrate, sides slightly constricted, anterior margin slightly emarginate; ventral apodeme (Fig. 44) expanded anteriorly, one-third width of dorsal apodeme, anterior margin rounded, not reaching anterior margin of dorsal apodeme; distiphallus (Fig. 51) short, broad basally, tapered posteriorly at 45 angle; ejaculatory apodeme (Fig. 37) narrow, posterior third slightly bulbous, anterior twothirds ßattened dorsoventrally, extending slightly beyond anterior margin of dorsal apodeme. Variation. Body measurements and thoracic macrosetal counts given in Table 2. Setae on frons, parafacial, and maxillary palpus pale yellow(mei ); veins and crossveins with brown margin becoming broader and more distinct in southern specimens. Female. Similar to male except as follows. Body measurements and thoracic macrosetal counts given in Table 2. Frons with dorsal half dark brown with brownish pruinescence, ventral half with silver pruinescence; setae black, scattered on dorsal half, mixed with white setae on ventral half which extend into area lateral to antennal sockets. Thorax. Setae on mesonotum short, dark brown and white; grayish white on laterotergite; white and pale brown on postpronotal lobe. Abdomen. Dark reddish brown, subshiny, tergites 1Ð3 and tergite Þve with lateral gray pruinescent triangle, tergite four lacking pruinescence; dorsal setae dark reddish brown, short, appressed, lateral setae white, elongate on pruinescent areas of tergites 1Ð3, dark reddish brown, short on tergites 4Ð5. Terminalia (Fig. 58). Tergite eight with anterior projection truncate. Furca oblong, length 0.62 mm, anterior margin rounded with lateral projections short, posterior margin truncate, lateral margins sinuate with quadrate mediolateral sclerotized plate attaching furca ventrally to penal guide. Variation. Body measurements and thoracic macrosetal counts given in Table 2. Parafacial setae white with scattered reddish brown setae (MEI ). Maxillary palpus dark yellow; body, legs, and abdomen reddish brown (MEI ). Seasonal Activity and Distribution. Adults (N 98) have been collected between 1 March and 29 July. Specimens have been collected in Malaise traps, often in oak savanna, in a Þeld near a dry oak forest, at the border of a Þeld and dry oak forest, and in cultivated cotton. P. albifrons (Fig. 64) ranges from northern Florida north to Maine and west to Texas and Kansas. Specimens Examined Type Material. Type specimens of SayÕs Diptera have been destroyed. The neotype of T. albifrons (MEI ), herein described and designated, was collected in a Malaise trap from Forest Glen Forest Preserve, 8 km SE Westville, Vermilion County, Illinois, 18Ð20-IV-1977, by D. W. Webb, and deposited in the Illinois Natural History Survey collection. This neotype is designated to Þx the concept of T. albifrons and to ensure universal and consistent interpretation of the same. The holotype female of T. nitoris Coquillett (USNM Type Number 993) was collected in Missouri. The holotype female of T. borealis Cole (CUIC Type Number 4352) was collected at the Agricultural College, Michigan on 9 May Other Material CANADA. ONTARIO. Carleton County: Constance Bay, 24-V-1971, K. J. G. Deacon, 1, MEI (DEBU). Lambton County: Pinery Provincial Park, 22Ð23-V-1992, 1, MEI (DEBU), Malaise trap, oak savanna.

11 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 379 Figs (30Ð36): Male sternite 8, gonocoxite ventral view. (30) P. albifrons. (31) P. conspicua. (32) P. constricta. (33) P. grandis. (34) P. melampodia. (35) P. montivaga. (36) P. rufa. (37Ð43): Male aedeagus, dorsal viewwith insert of frontal view of ejaculatory apodeme. (37) P. albifrons. (38) P. conspicua. (39) P. constricta. (40) P. grandis. (41) P. melampodia. (42) P. montivaga. (43) P. rufa. Scale 0.1 mm. Gx, gonocoxite; Gs, gonostylus; S 8, sternite eight; VL, ventral lobe. UNITED STATES. CONNECTICUT. Windham County: 1.8 km W Windham, 31-V-1989, C. T. Maier, 1, MEI , (CAES). FLORIDA. Alachua County: W Gainesville, 1-IV- 1976, W. H. Pierce, 1, MEI , (FSCA), ßight trap; 14.4 km NW Gainesville, University of Florida Horticultural Unit, 1Ð2-III-1977, H. N. Greenbaum and H. V. Weems, Jr., 1, MEI (FSCA); 9Ð13-III-1977, 3, MEI , , (FSCA); 16Ð19-III-1977, 1, MEI (FSCA), ßight trap, 25Ð29-III-1977, 1, MEI (FSCA); 12Ð19-III-1978, 1, MEI (FSCA); 23- III-2-IV-1978, H. N. Greenbaum, 1, MEI (FSCA). GEORGIA. Clarke County: Athens, Whitehall Preserve, 2Ð9-IV-1979, R. H. Turnbow, Jr., 1, MEI (UGCA); Whitehall Forest, 7-V-1984, R. T. Franklin, 1, MEI (UGCA), Malaise trap. ILLINOIS. Mason County: Sand Ridge State Forest, 13-V-1975, C. T. Maier, 1, MEI (INHS), Malaise trap in Þeld near dry oak forest; 15Ð18-V-1975, 2, MEI , (INHS); 19-V-1975, 1, MEI (INHS); 19-V-1976, 1, MEI (INHS), Malaise trap at the border of a Þeld and dry oak forest; 20Ð29-VII-1993, M. L. Hartman, 1, MEI (MEIC). McHenry County: Algonquin, Nason, 1, MEI (INHS). Vermilion County: Forest Glen Forest Preserve, 8 km SE Westville, 18Ð20-IV-1977, D. W. Webb, 1, MEI (INHS), Malaise trap; 22Ð25-IV-1977, D. W. Webb, 1, MEI (INHS), Malaise trap. KANSAS. Riley County: -IV-1921, J. B. Norton, 1, MEI (KSUC), 4-V-1964, E. Marston, 2, MEI , (KSUC), Malaise trap. MAINE. Kennebec County: Augusta, 14-VI-1945, 1, MEI (PMNH). MARYLAND. Prince Georges County: Laurel, 2-VI- 1965, 1, MEI (CNCI), 1, MEI (MEIC), 11-V-1965, 1, MEI (CNCI), Malaise trap.

12 380 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 Figs (44Ð50): Male aedeagus, ventral view. (44) P. albifrons. (45) P. conspicua. (46) P. constricta. (47) P. grandis. (48) P. melampodia. (49) P. montivaga. (50) P. rufa. (51Ð57): Male aedeagus, lateral view. (51) P. albifrons. (52) P. conspicua. (53) P. constricta. (54) P. grandis. (55) P. melampodia. (56) P. montivaga. (57) P. rufa. Scale 0.1 mm. MASSACHUSETTS. Hampshire County: Amhurst, 10-V-1949, Hunter, 1, MEI (UGCA); 17-V- 1968, R. DufÞeld, 1, MEI (UGCA); 1-VI-1968, 1, MEI (UGCA); Ware, 4-VII-1968, Lavallee, 1, MEI (UGCA). Middlesex County: Lincoln, 17-V-1-VI-1982, E. T. Armstrong, 1, MEI (MCZC); Framingham, 12-V-1968, T. Lavallee, 1, MEI (UGCA). MISSISSIPPI. Lafayette County: -V-VI-1945, F. M. Hall, 2, MEI , (CNCI). Pontotoc County: 1.6 km SE Ecru, 23-IV-1980, G. L. Snodgrass, 1, MEI (INHS), in cultivated cotton. MISSOURI. Boone County: Columbia, 18-V-1970, F. D. Parker, 1, MEI (UCDC), Malaise trap. Holt County: Mound City, 21-IV-1968, P. L. Andrews, 2, MEI , (UMRM), 1, MEI (INHS); 28-IV-1968, 1, MEI (UMRM); 1-V- 1968, 3 2, MEI , Ð9, Ð6 (UMRM); 2-V-1968, 2, MEI (INHS) (MEIC); 3-V-1968, 1 3, MEI , Ð69 (UMRM); 29-V-1968, 1, MEI (UMRM); 27- VI-1968, 1, MEI (UMRM). Jackson County: Atherton, 30-IV-1901, 1, MEI (OSUC). St. Charles County: Weldon Springs, 3-V-1968, 1, MEI (UMRM). NEW HAMPSHIRE. Strafford County: Durham, 10- VI-1982, W. J. Morse, 1, MEI (DENH); Madbury, 7-VI-1981, 1, MEI (DENH), Malaise trap. NEW YORK. Albany County: Pine Bush, N W, 100 m, 6-V-1982, T. McCabe, 1, MEI (INHS); 18-V-1982, 1, MEI (INHS); 25-V-1982, 1, MEI (MEIC), 1, MEI (NYSM); 27-V-1982, 1, MEI (NYSM); 31-V- 1982, 1, MEI (NYSM); 6-VI-1983, 2, MEI (NYSM), MEI (NYSM); 12-VI-1983, 2, MEI Ð6 (NYSM); 13-VI-1983, 1, MEI (NYSM); 2, MEI , (NYSM).

13 July 2003 WEBB AND METZ: THE NEARCTIC SPECIES OF Pandivirilia IRWIN AND LYNEBORG 381 Table 2. the mean) Morphometric variation in the species of Pandivirilia (range [millimeters] and ratios for each measurement are followed by P. albifrons P. conspicua P. constricta P. grandis P. melampodia P. montivaga P. rufa Male (n 10) (n 10) (n 5) (n 4) (n 10) (n 10) (n 1) Body length (excluding 8.2Ð10.4, Ð12.4, Ð9.6, Ð13.9, Ð10.7, Ð11.5, antenna) Wing length 6.7Ð8.2, Ð9.4, Ð8.0, Ð11.5, Ð7.8, Ð9.8, Wing length/width 2.6Ð3.1, Ð3.7, Ð3.5, Ð3.2, Ð3.7, Ð3.5, Head length 0.98Ð1.24, Ð1.38, Ð1.20, Ð1.44, Ð1.16, Ð1.4, Dist. betw. eyes/width med. ocellu 0.2Ð0.5, Ð0.30, Ð0.25, Ð1.2, Ð2.3, Ð1.3, Antenna/head length 0.68Ð0.83, Ð0.79, Ð0.79, Ð0.80, Ð0.99, Ð0.98, Scape length 0.26Ð0.32, Ð0.34, Ð0.36, Ð0.36, Ð0.34, Ð0.36, Scape length/width 1.6Ð2.3, Ð2.00, Ð2.3, Ð2.0, Ð2.4, Ð2.3, Scape length/pedicel length 2.6Ð3.3, Ð3.8, Ð3.5, Ð3.6, Ð3.8, Ð4.5, Scape width/ßagellum width 0.8Ð1.1, Ð1.1, Ð1.0, Ð1.1, Ð0.9, Ð1.3, Pedicel length 0.08Ð0.10, Ð0.12, Ð0.11, Ð0.11, Ð1.12, Ð0.11, Pedicel length/width 0.5Ð0.8, Ð0.83, Ð0.8, Ð0.6, Ð0.9, Ð0.7, Flagellum length 0.38Ð0.46, Ð0.61, Ð0.53, Ð0.63, Ð0.53, Ð0.68, Flagellum length/width 2.2Ð3.1, Ð3.1, Ð3.8, Ð0.8, Ð3.3, Ð4.3, Flagellum length/scape length 1.3Ð1.7, Ð2.2, Ð1.8, Ð2.1, Ð2.0, Ð1.9, Maxillary palpus length 0.54Ð0.67, Ð0.72, Ð0.66, Ð0.67, Ð0.64, Ð0.70, Maxillary palpus length/width 5.4Ð9.0, Ð12.0, Ð8.3, Ð6.9, Ð7.5, Ð8.8, Notopleural macrosetae 3Ð4, 4 2Ð7, 4 3Ð4, 4 4Ð5, 4 3Ð4, 4 3Ð4, 4 7 Supraalar macrosetae 2Ð3, 2 1Ð3, 2 2 2Ð3, 2Ð3 2 2Ð4, 2 2 Postalar macrosetae Dorsocentral macrosetae 1 1Ð4, 2 1Ð2, 2 1-2, 2 0Ð2, 1 0Ð2, 1 1 Scutellar macrosetae Female (n 10) (n 10) (n 10) (n 10) (n 10) (n 3) Body length (excluding antenna) 7.5Ð11.5, Ð12.7, Ð16.6, Ð12.7, Ð11.9, Ð16.8, 15.1 Wing length 5.8Ð9.3, Ð9.4, Ð12.0, Ð9.4, Ð10.0, Ð11.7, 11.0 Wing length/width 2.9Ð3.4, Ð3.4, Ð3.3, Ð3.4, Ð5.3, Ð3.3, 3.4 Head length 1.0Ð1.3, Ð1.54, Ð1.50, Ð1.42, Ð1.4, Ð1.36, 1.20 Antenna/head length 0.7Ð0.8, Ð0.8, Ð0.9, Ð0.95, Ð0.97, Ð0.98, 0.91 Scape length 0.26Ð0.40, Ð0.34, Ð0.40, Ð0.42, Ð0.44, Ð0.44, 0.33 Scape length/width 1.6Ð2.3, Ð2.1, Ð2.4, Ð2.5, Ð2.4, Ð2.0, 1.7 Scape length/pedicel length 2.9Ð4.5, Ð3.3, Ð4.0, Ð4.0, Ð3.7, Ð4.4, 3.1 Scape width/ßagellum width 0.8Ð1.1, Ð1.0, Ð1.1, Ð0.90, Ð1.1, Ð1.3, 1.1 Pedicel length 0.07Ð0.12, Ð0.12, Ð0.12, Ð0.12, Ð0.12, Ð0.12, 1.11 Pedicel length/width 0.5Ð0.7, Ð0.8, Ð0.7, Ð0.8, Ð0.8, Ð0.7, 0.7 Flagellum length 0.38Ð0.54, Ð0.61, Ð0.66, Ð0.64, Ð0.82, Flagellum length/width 2.1Ð3.2, Ð2.9, Ð3.3, Ð3.1, Ð4.6, Ð3.8, 3.4 Flagellum length/scape length 1.1Ð1.7, Ð2.1, Ð1.8, Ð1.6, Ð2.9, Ð2.7, 2.0 Maxillary palpus length 0.50Ð0.68, Ð0.70, Ð0.76, Ð0.86, Ð0.70, Ð0.82, 0.71 Maxillary palpus length/width 5.4Ð8.8, Ð10.5, Ð7.0, Ð6.7, Ð7.5, Ð6.8, 6.3 Notopleural macrosetae 1Ð5, 3 3Ð6, 3 3Ð6, 4 3Ð6, 3Ð4 3Ð4, 3Ð4 3Ð4, 4 Supraalar macrosetae 2Ð3, 2 2Ð3, 2 2Ð4, 2 1Ð2, 2 2Ð3, 2 2 Postalar macrosetae 1 1Ð2, Dorsocentral macrosetae 1Ð2, 1 1Ð2, 2 1Ð2, 2 0Ð2, 1 0Ð2, 1Ð2 1Ð2, 2 Scutellar macrosetae Ð2, 1 2 OHIO. Jefferson County: Salem Township, 19-V- 1965, J. Flenniken, 1, MEI (OSUC). Nassau County: Oyster Bay, 10-VI-1961, D. W. S. Sutherland, 1, MEI (DENH). OKLAHOMA. Bryan County: Bennington, 28-IV- 1983, 1, MEI (INHS). McClain County: Payne, near Lake Carl Blackwell, 21-IV-1979, 1, MEI (OSEC); Payne, west of Lake Carl Blackwell, 20-IV-1979, 1, MEI (OSEC); Payne, south side of Lake Carl Blackwell, 20-IV-1979, 1, MEI (OSEC). RHODE ISLAND. Washington County: Kingston, 22-V-1970, T. Lavallee, 1, MEI (UGCA). SOUTH CAROLINA. Beaufort County: Parris Island, 2Ð9-IV-1981, J. F. Reinert, 2, MEI , (FSCA). TEXAS. Brazos County: College Station, 16-III-9- IV-1988, R. A. Wharton, 1, MEI (TAMU). VIRGINIA. Arlington County: Falls Church, 18-V- 1917, C. T. Greene, 1, MEI (USNM); Glen Carlyn, 82 m, 4-V-1919, C. T. Greene, 1, MEI (INHS). Louisa County: Louisa, 19-IV-1964, 1, MEI (FSCA). Norfolk County: Dismal Swamp, 25- IV-1971, 1, MEI (INHS). WISCONSIN. Burnett County: 3.2 km W Grantsburg, 5Ð12-V-1199, M. Sabourin, 2, MEI Ð7 (MEIC), Malaise trap; 19Ð25-V-1999, 4, MEI , , , (MEIC); 26-V-1-VI-1999, 4, MEI , , , (MEIC); 2Ð10- VI-199, 3, MEI Ð2, (MEIC). Pandivirilia conspicua (Walker), new combination conspicua Walker (1848: 223, desc.) (Thereva); Cole (1923: 74, key, desc.; 1965:351, cat.), Irwin and Lyneborg (1981a: 227, listing) (Psilocephala). Type: LT in NHM(E).