3 JANUARY 1997 MEMOIR ENTOMOL. SOC. WASH. 18, 1996, pp. 261-272

SPILOMYIA FLOWER OF THE NEW WORLD (DIPTERA: SYRPHIDAE)

E CHRISTIAN THOMPSON

Systematic Entomology Laboratory, ARS, USDA, NHB-168, Smithsonian Institution, Washington, DC. 20560.

Abstract.— wirthi, new species, is described. A key to the species of Spi- lomyia of the New World is given, along with tables of their diagnostic characters and the nomenclatural details for the various names which apply to them. Key Words: key, Neotropical

Biodiversity is now a global concern of two at a time. However, regardless of how high priority. Most nations have signed a they are produced, they are all critical to biodiversity convention that proclaims this the edifice. So, with that knowledge, I add concern (Convention on Biological Diver- another brick to the edifice and our under- sity 1994). So, we now must understand our standing of biodiversity. In doing so, I also biodiversity, for only through understand- honor a great mason, Willis Wagner Wirth: ing can we learn to use biodiversity in a Bill Wirth was an enthusiastic contributor, sustainable manner. The first task is enu- who over the past half century contributed meration, or the listing of all components more to building our knowledge of Diptera of biodiversity, from genes to populations than any of his contemporaries. While Bill (Solbrig 1990), with species being the cen- understood the need for architects, like Wil- tral component. Enumeration requires that li Hennig, he preferred to produce the basic the components have names. Unfortunately, alpha taxonomic work from which great most species have none. To name a species classifications are derived. Without the gen- scientifically requires more than just creat- era and species Bill described, the classifi- ing an epithet. A scientific name is a state- cations created by dipterists would be noth- ment of relationship, a hypothesis of where ing more than barren trees! the organism belongs in a classification. The Spilomyia species found in the New Our biological classification is a great edi- World have never been revised as a whole. fice that includes millions of names in a hi- Williston (1886) published a treatment of erarchical structure that expresses the rela- the North American species he knew; Cur- tionships among the concepts and organ- ran (1951) provided a synopsis; Vockeroth isms represented by those names. All great (1958) revised a species, adding two new edifices are built one brick at a time. So, it ones; Nayar (1968) figured the male geni- is with our classification and, hence, our talia of some species; Nayar and Cole knowledge of biodiversity. The first bricks (1968) redescribed two species under new are new species descriptions. Sometimes names; Maier, Waldbauer and associates these bricks are produced in great batches, (Maier 1982; Maier and Waldbauer 1979 a but more often they are produced one or and b; Waldbauer 1970, 1983; Waldbauer 262 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Sheldon 1971; Waldbauer and Ghent on anepisternum and katepisternum; hind fe- 3 1984; Waldbauer and LaBerge 1985; Wald- mur black on apical /3 S. sayi - Pleuron extensively yellow, only black ven- bauer et al. 1977) provided data on biology, trally on katepisternum and katepimeron and flower visiting, mate-seeking, and mimicry; anteriorly on laterotergite (Fig. 15); hind fe- Thompson (1972) redescribed the mur yellow S. gratiosa and discussed its phylogenetic relationships 8. Abdomen with yellow fasciae narrow, with (as did Hippa (1978)); and Hippa (1986) black areas extensive, not reduced to fascia, with yellow fasciae narrower than black areas included one species in his study of female (Figs. 3,9,10) 10 genitalia. Hence, the New World Spilomyia - Abdomen with yellow fasciae very broad, are ready for a monographic treatment, but with black areas reduced to narrow fascia, this is postponed until the more compre- with these black fasciae narrower than yellow hensive taxon, Syrphidae, can be treated as ones (Fig. 2) 9 a whole. Instead, all the essential informa- 9. Propleuron yellow above coxa; discal cell completely microtrichose; anal cell almost tion necessary to know the New World spe- completely microtrichose S. texana cies of Spilomyia is presented in a slightly - Propleuron black (Fig. 12); discal cell bare on nontraditional, but highly compact format. basal % or more; anal cell completely bare Diagnoses of previously described species S. alcimus have been reduced to tables of diagnostic 10. Abdomen with some yellow fasciae discon- tinuous, with basal fasciae on 2nd, 3rd and characters and taxa. Likewise, the critical 4th terga interrupted medially (Figs. 9, 10) 13 nomenclatorial data are summarized in a ta- - Abdomen with yellow fasciae always contin- ble. uous on 2nd tergum, usually on 3rd tergum, frequently on 4th tergum also (Fig. 3) .... 11 KEY TO NEW WORLD SPECIES OF SPILOMYIA 11. Metasternum yellow S. sp. CR-3 - Metasternum black 12 1. Scutellum entirely black (Fig. 20) . . S. obscura 12. Pleuron with yellow areas extending contin- - Scutellum yellow at least on apical rim (Figs. uously from katepisternum to laterotergite 19, 21) 2 S. longicornis 2. 2nd abdominal tergum entirely black, rarely - Pleuron with yellow areas broadly interrupted with apicolateral margin narrowly yellow, but along suture at junction of epimeron and la- never continuous (Fig. 1) S. fusca terotergite S. crandalli - 2nd tergum with at least a narrow continuous 13. Anepimeron and katepimeron black (as in yellow fascia (Figs. 2-11) 3 Fig. 14) S. liturata 3. Scutum dark anterior to scutellum (Fig. 21) - Anepimeron at least partially yellow (Fig. 13) S. ephippium 14 - Scutum with yellow prescutellar macula 14. Hind femur entirely yellow on anterior face; (Figs. 19-20) 4 abdomen with yellow basal and apical fasciae 4. 2nd and 3rd abdominal terga with basal fascia continuous laterally, so lateral margin entirely reduced to triangular mediolateral maculae; yellow; 4th tergum with yellow apical fascia 4th tergum without apicolateral fascia (Fig. 15 11) S. pleuralis - Hind femur black on apical %\ abdomen with - 2nd tergum with basal fascia elongate, contin- yellow fasciae isolated laterally, so lateral uous or narrowly interrupted medially, not re- margin alternating black and yellow; 4th ter- duced to lateral triangles (Figs. 2-10) 5 gum without apical yellow fascia S. wirthi 5. 2nd and 3rd terga with 2 yellow fasciae .... 8 15. Face without tubercle, not produced ventrally, - 2nd tergum with a single basal yellow fascia so ventral margin of head horizontal (see 6 Vockeroth 1958: 285, Fig. 3); pleuron with 6. 4th tergum entirely reddish yellow to yellow yellow on anepimeron extending on katepi- in ground color, yellow pollinose (Fig. 7); meron (as in Fig. 13) S. foxleei foreleg entirely pale orange S. kahli - Face with weak, but distinct tubercle, pro- - 4th tergum dark with a single medial yellow duced ventrally so ventral margin of head an- fascia (Figs. 4, 6); foreleg yellow basally, gular (see Vockeroth 1958: 285, Fig. 4); ka- dark on apex of tibia and on tarsus 7 tepimeron entirely black (as in Fig. 12) .... 16 7. Pleuron extensively black, only small supra- 16. Pleuron with yellow maculae on anepimeron procoxal yellow macula and yellow maculae and laterotergite narrowly separated by thin CONTRIBUTIONS ON DIPTERA 263

Figs. 1-9. Spilomyia abdominal pattern, dorsal view. 1, S. fusca. 2, S. alcimus. 3, S. longicornis. 4, S. sayi. 5, S. ephippium. 6, S. gratiosa. 7, S. kahli. 8, 5. obscura. 9, S. wirthi. 264 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 10-16. Figs. 10—11, Spilomyia abdominal pattern, dorsal view. 10, S. interrupta. 11, S. pleuralis. Figs. 12—16, Spilomyia thoracic pattern, lateral view. 12, S. alcimus. 13, S. longicornis. 14, S. obscura. 15, S. gratiosa. 16, S. ephippium.

black vitta; male surstylus with apex blunt; and laterotergite broadly separated by a wide female 5th tergum with yellow medial fascia black vitta; male surstylus with apex distinct- expanded laterally and continuous with apical ly notched; female 5th tergum with yellow yellow fascia, so that entire lateral margin is medial fascia isolated from lateral margin and yellow S. interrupta apical fasciae, so that lateral margin black on Pleuron with yellow maculae on anepimeron basal 2A S. cilima CONTRIBUTIONS ON DIPTERA 265

Spilomyia wirthi Thompson, silvery pollinose, yellow and black pilose; NEW SPECIES trochanters black, black pilose, shiny; fore and mid femora black ventrally and dor- Male.—Head: Face yellow except sally, yellow laterally, black pilose; hind broad black medial vitta which is narrowly femur black except yellow anterobasal V separated from antennal pits and black lat- 3 and posterobasal '/4, black pilose; fore tibia erally along juncture with gena, sparsely yellow except black on anteroapical V5 and white pollinose on yellow areas, shiny on 2 posteroapical /3, yellow pilose except black areas, white pilose; gena yellow ex- brown pilose on posteroapical %; mid tibia cept black at juncture with face, shiny an- yellow, yellow pilose; hind tibia yellow ex- teroventrally, white pollinose and pilose cept brownish apical '/4; fore tarsus black dorsoposteriorly; occiput black except yel- except orange apical tarsomere, dark pilose; low ventral V6, silvery pollinose, white pi- mid and hind tarsi orange, pale pilose. lose except black on dorsal V3; frontal Wing: brownish black anteriorly, hyaline triangle black, shiny, bare; frontal lunule posteriorly, with dark color extending pos- black; vertical triangle black, black pilose, teriorly to vein R, Rs and R4+5 and slightly yellow pollinose anterior to anterior ocel- beyond vein R4+5 in apical V3 of cell R and lus; eye contiguity long, about as long as in fork between crossvein r-m and vein scape; antenna black, black pilose; basofla- R4+5, microtrichose except bare alula, cell gellomere with elongate mesial sensory BM, cell R posterior to spurious vein, cell pit; arista yellow becoming white apically. CuP except microtrichose apicomedially, Thorax: Black with yellow maculae; cell CuA, laterally and anterior to vein A2. postpronotum yellow except black medi- Abdomen: Black with narrow yellow ally, black pilose. Mesonotum black pi- fasciae; sterna black except narrow yellow lose, black pollinose except for broad me- apical margins on 1st, 2nd and 3rd, shiny, dial silvery pollinose vitta on anterior % long yellow pilose except with short op- and shiny dorsad to wing, black except pressed black pile intermixed apically on yellow as follows: submedial anterior mac- 4th; 1st tergum black, black pollinose and ula mesiad to postpronotum, notopleuron, pilose; 2nd tergum black except narrow vitta running anteriorly from postalar cal- subbasal and apical yellow fasciae black lus obliquely to transverse suture, and pollinose and pilose; 3rd tergum black ex- oblique vitta anterior to scutellum. Scutel- cept narrow submedial and apical yellow lum black except yellow apical margin, fasciae black pollinose and pilose; 4th ter- black pilose. Pleuron yellow pilose except gum black except for narrow subbasal and black pilose dorsally on anepisternum and subapical yellow fasciae, shiny, black pi- anterior anepimeron, sparsely black polli- lose; male genitalia black, shiny, black pilose. nose except densely silvery pollinose on Holotype male.—COSTA RICA, San katepisternum, black except yellow as fol- Jose, San Gerardo de Dota, 2000-2500 m, lows: proepimeron, large maculae on pos- lambert coordinates 387400 482700, 22-26 terior anepisternum, dorsal katepisternum, February 1992, Curso Tachnidae y Syrphi- posterior anepimeron including dorsal tri- dae, net-collected, INBIOCRI000406886, angular portion, laterotergite, dorsal kate- deposited in Institute Nacional de Biodiv- pimeron and apicolaterally on metaster- ersidad, Santo Domingo. Para types: COS- num. Halter yellow; calypter with dorsal TA RICA, San Jose, San Gerardo de Dota, lobe black, ventral lobe yellow. Legs: fore Finca Zacatales, 2300 m, LN 387000- coxa black laterally, yellow medially, sil- 483250, 26 February 1992, F. C. Thomp- very pollinose, white pilose; mid and hind son, USNM ENT 00022134 (

Figs. 17-21. Figs. 17-18, Spilomyia wing color and microtrichia pattern, dorsal view. 17, S. ephippium. 18, S. wirthi. Figs. 19-21, Spilomyia thoracic pattern, dorsal view. 19, S. longicornis. 20, S. obscura. 21, S. ephip- pium. CONTRIBUTIONS ON DIPTERA 267

Figs. 22-24. Male genitalia of Spilomyia wirthi. 22, 9th tergum and associated structure, lateral view. 23, Aedeagus, lateral view. 24, 9th sternum and associated structures, lateral view. 268 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

G. Zumbado, INBIOCRI000409035 (c? IN . 1986. Morphology and taxonomic value of Bio), INBIOCRI000409036 (? INBio). the female external genitalia of Syrphidae and some other Diptera by new methodology. An- Chirripo National Park, F. Cementario de la nales Zoologici Fennici 23: 307-320. Maquina, 2100-2500 m, LS 378700-512500, Janzen, D. H. 1994. : Universal and es- M. A. Zumbado, INBIOCRI001305812 (c* sential infrastructure for development and man- INBio). agement of tropical wildland biodiversity, pp. 100-113. In Sandlund, O. T. and Schei, P. J., ACKNOWLEDGMENTS eds., Proceedings of the Norway/UNEP Expert Conference on Biodiversity, Trondheim, Nor- I thank J. R. Vockeroth, Canadian Na- way, NINA, 190 pp. tional Collection, Agriculture Canada, Ot- Maier, C. T. 1982. Larval habitats and mate-seeking tawa (CNC); Manuel Zumbado, Institudo sites of flower flies (Diptera: Syrphidae, Eristal- Nacional de Biodiversidad, Santo Domin- inae). Proceedings of the Entomological Society go de Heredia (INBIO); Philip Perkins, of Washington 84: 603-609. Maier, C. T. and G. P. Waldbauer. 1979a. Dual mate- Museum of Comparative Zoology, Har- seeking strategies in male syrphid flies (Diptera: vard University, Cambridge; Paul Hanson, Syrphidae). Annals of the Entomological Society University of Costa Rica, San Jose (UCR); of America 72: 54-61. for permission to study material in their . 1979b. Diurnal activity patterns of flower care [other collection acronyms used are flies (Diptera: Syrphidae) in an Illinois sand area. Annals of the Entomological Society of America USNM for United States National Museum 72: 237-245. (= the National Museum of Natural His- Nayar, J. L. 1968. The male genitalia of the genus tory), Washington, DC. I also thank Drs. Spilomyia Meigen Tribe (Diptera: Syr- Chris Maier, Connecticut Agricultural Ex- phidae). Journal of the New York Entomological perimental Station, New Haven; Wayne N. Society 76: 168-175. Mathis, Smithsonian Institution, Washing- Nayar, J. L. and F. R. Cole. 1968. Two new species of Spilomyia Meigen (Syrphidae: Diptera). Pan- ton, DC; Manya Stoetzel, Allen L. No- Pacific Entomologist 44: 211-214. rrbom and Richard White, Systematic En- Solbrig, O. 1990. From genes to ecosystems. A re- tomology Laboratory, USDA, Washington, search agenda for biodiversity. Report of IUBS- DC; for their critical reviews of the SCOPE-UNESCO Workshop, Harvard Forest, manuscript. Finally, thanks are due to Lin- Petersham, Massachusetts, USA, June 27-July 1, 1991. Cambridge, IUBS, [ii] + 123 pp. da Lawrence, without whose illustrations Thompson, F. C, J. R. Vockeroth, and Sedman, Y S. this paper would not be complete. 1976. Family Syrphidae. In Papavero, N. ed., A Catalogue of the Diptera of the Americas South LITERATURE CITED of the United States. Departmento de Zoologia, Full references to most of the literature Secretaria da Agricultura. Sao Paulo, Brazil, 195 pp. cited in the text are found in the North and Waldbauer, G. P. 1970. Mimicry of hymenopteran an- tennae by Syrphidae. Psyche 77: 45-49. South American Diptera catalogs (Wirth et . 1983. Flower associations of mimetic Syr- al. (1965) and Thompson et al. (1976)). phidae (Diptera) in northern Michigan. Great Only those references not included in these Lakes Entomologist 16: 79-85. references are given here. Waldbauer, G. P. and A. W. Ghent. 1984. Flower as- sociations and mating behavior or its absence at blossoms by Spilomyia spp. (Diptera, Syrphidae). Great Lakes Entomologist 17: 13-16. Convention on biological diversity. 1994. Conven- Waldbauer, G. P. and W. E. LaBerge. 1985. Phenolog- tion on biological diversity. Text and Annexes. ical relationships of wasps, bumblebees, their Geneva, UNEP/CBD/94/1, 34 pp. mimics and insectivorous birds in northern Mich- Goot, V. S. van der 1964. Fluke's catalogue of Neo- igan. Ecological Entomology 10: 99-110. tropical Syrphidae (Insecta, Diptera), a critical Waldbauer, G. P. and J. K. Sheldon. 1971. Phenolog- study with an appendix on new names in Syr- ical relationships of some aculeata Hymenoptera, phidae. Beaufortia 10: 212-221. their dipteran mimics, and insectivorous birds. Hippa, H. 1978. Classification of Xylotini (Diptera, Evolution 25: 371-382. Syrphidae). Acta Zoologica Fennica 156, 153 pp. Waldbauer, G. P., J. G. Sternberg, and C. T. Maier. CONTRIBUTIONS ON DIPTERA 269

1977. Phenological relationships of wasps, bum- 1965. Family Syrphidae, pp. 557-625. In Stone, blebees, their mimics, and insectivorous birds in A., et al., eds., Catalog of Diptera of North Amer- an Illinois sand area. Ecology 58: 583-591. ica North of Mexico, United States Department of Wirth, W. W., Y. S. Sedman, and H. V. Weems, Jr. Agriculture, Agriculture Handbook 276, 1696 pp. 270 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Taxa and names of New World Spilomyia with nomenclatural details. Format used: valid species group name given in bold and left justified, followed by the author of the name (in parens if proposed in combination with another genus). The range is given next. The original combinations (genus and species group names) followed by the author of name, year and page of the publication of the name, the sex (sexes) described, the type-locality, with the kind of type, sex of type and type-depository given in parens. Abbreviations used are: AMNH = American Museum of Natural History; ANSP = Academy of Natural Sciences, Philadelphia; BMNH = British Museum (Natural History); CAS = California Academy of Sciences; CNC - Canadian National Collection; HT = holotype; Institute Nacional de Biodiversidad; LT = lectotype; MCZ = Museum of Comparative Zoology; ST = syntype(s); UKaL = University of Kansas, Lawrence; USNM = United States National Museum.

alcimus (Walker). Wisconsin to Newfoundland, south to Mississippi and Florida. Alcimus Walker 1849: 563 c* Unknown (LT

Table 2. Diagnostic characters of new world Spilomyia species.

Head 10.2 Anepimeron, posterior portion: 1. Face: a. entirely black (rarely diffusely pale medially a. entirely yellow; C«jy0; or b. with tubercle dark; b. yellow (brownish red in ephippium) or with c. with narrow dark medial vitta; or large yellow macula. d. with broad dark medial vitta. 10.3 Anepimeron, dorsomedial portion: 2. Gena: a. entirely black; or a. entirely yellow (brownish orange in kahli); b. yellow (brownish red in ephippium) or with b. with black vitta or partially black); or large yellow macula. c. entirely black. 11. Katepimeron: 3. Frontal triangle (6*): a. entirely dark; a. entirely yellow; b. yellow (brownish red in ephippium) dorsally; b. yellow laterally, black medially; or or c. entirely black. c. entirely yellow. 4. Scape: 12. Katatergum: a. short, about as long as basoflagellomere; or a. entirely dark; or b. elongate, at least 1.5 (2.5 ephippium) times b. yellow (brownish red in ephippium) or with as long as basoflagellomere. yellow macula. 5. Scape: 13. Metasternum: a. about as long as broad; or a. entirely dark; b. at least twice (3 times ephippium) as long as b. partially yellow; or broad. c. entirely yellow (brownish red in ephippium). Thorax 14. Metepimeron: 6. Notopleuron: a. entirely dark; or a. entirely dark and concolorous with mesonotal b. yellow at least dorsally. disc; 15. Wing cell BM: b. entirely brownish red (ephippium) to orange a. hyaline; or (kahli), contrasting with color of mesonotal b. dark. disc; or c. with yellow macula. 16. Foreleg: 7. Mesonotum: a. entirely pale; or b. dark apically. a. with yellow prescutellar triangular macula; or b. without prescutellar macula. 17. Hind femur: 8. Scutellum: a. entirely yellow; b. anterior face entirely yellow (orange in kah- a. entirely black; li), elsewhere partially black; b. entirely yellow (orange in kahli, brownish red c. black at least 1/2 or more; or in ephippium); d. entirely dark. c. black with narrow yellow apical margin; or d. black with broad yellow apical margin. Abdomen 9. Proanepimeron: 18. 1st tergum: a. entirely dark (rarely diffusely pale medially a. entirely black (brownish reddish in ephip- (alcimus, kahli. sayi); or pium); b. yellow (brownish red in ephippium) or with b. yellow laterally; or large yellow macula. c. entirely pale. 10. Anepimeron: 19. 2nd tergum: a. entirely black; or a. entirely dark; b. at least partially pale, yellow (brownish red b. with two continuous yellow fasciae; in ephippium). c. with interrupted subbasal yellow fascia and continuous apical yellow fascia; 10.1 Anepimeron, anterior portion: d. with basolateral yellow fascia and broadly in- a. entirely black; or terrupted apical yellow fascia; b. yellow (brownish red in ephippium) or with e. yellow basolaterally or with yellow large yellow macula. basolateral fascia, with no apical fascia; or 272 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Continued.

x. with a yellow basolateral triangular macula d. entirely pale; or and narrow yellow apical fascia. x. with a yellow medial and subapical triangular maculae. 20. 3rd tergum: a. entirely dark; 22. 5th (?) tergum: b. with two (subbasal and apical) continuous a. entirely dark; yellow fasciae; b. with two (subbasal and apical) continuous c. with interrupted subbasal yellow fascia and yellow fasciae; continuous apical yellow fascia; c. yellow except black basomedially (or basally d. with only narrow medial discontinuous yel- in obscura); or low fascia; d. entirely pale, e. with a narrow apical yellow fascia; 23. 6* terminalia f. yellow basolaterally, with narrow yellow apical fascia; or a. entirely black; x. with a yellow medial triangular macula and b. partially yellow; or narrow yellow apical fascia. c. entirely yellow. 21. 4th tergum: 24. Abdominal margins from 2nd to 4th (d*) or 5 th (9) terga: a. entirely dark; b. with two (subbasal and apical) continuous a. entirely pale; or yellow fasciae; b. alternating dark and pale. c. with interrupted subbasal yellow fascia and continuous apical yellow fascia;

Table 3. Distribution of diagnostic character states among New World Spilomyia species.

Species l 2 3 4 5 6 7 8 9 10 10.1 10.2 10.3 n 12 13 14 15 16 17 18 19 20 21 22 23 24 alcimus b b a a a a a b a a a a a a b a b a b b a b b a b C a citima c b b b b c a d b b a b a a b a b a b b b c c c b C a CR-3 c b b b b c a d b b a b b b b c b a b b b b b b d C a crandalli c b b b b c a d b b a b b b b a b a b b b b b b d c a epitippium c b c b b b b b b b b b b b c c a b b d a e e a a a b foxleei c a b b b c a d b b a b b b b a b a b b b c c c b c a fusca b a b a a a a c b a a a a c b c b a b a c e b b c c b gratiosa c b b a a c a d b b b b b c b c b a b a c e b b c c b interrupta c b b b b c a d b b a b a a b a b a b b b c c c b c a kahli a a a b b be a b a a a a a a b a b b a b b e f d d c a liturata c b b b b a b d b a a a a a b a b a b b b c c c b c a longicornis c b b a b c a d b b a b b b b a b a b b b b b(c)b(c) d c a obscura a c a b b a a a b a a a a a a a b a b b a d b b c a by pleuralis c b b b b c a d b b b b b b b c b a b b b X X X a a b sayi c c b b b a a c a a a a a a a a a a b c a e d c a a b texana b b a a a a a b b a a a a a b a b a b b a b b a b c a wirthi d b c a a c a d b b a b b b b b b a b c a c c c ? a b