186 Florida Entomologist 86(2) June 2003 THE GENUS RHOPALOSYRPHUS (DIPTERA: SYRPHIDAE) HOWARD V. WEEMS, JR.1, F. CHRISTIAN THOMPSON2, GRAHAM ROTHERAY3 AND MARK A. DEYRUP4 '(retired) Florida State Collection of Arthropods, P.O. Box 2309, Hawthorne, FL 32640-2309 'SystematicEntomology Laboratory, USDA, NHB-168 Smithsonian Institution, Washington, DC. 20560 ^Department of Natural History, Royal Museum of Scotland, Chambers Street, Edinburgh, EH1 1JF Scotland 'Archbold Biological Station, P.O. Box 2057, Lake Placid, FL 33862 ABSTRACT The flower fly genus Rhopalosyrphus Giglio-Tos (Diptera: Syrphidae) is revised. The genus is redescribed; a key to species is presented; the phylogenetic relationships of the genus and species are hypothesized; the included species are described; with new species, R. ramu- lorum Weems & Deyrup, described from Florida (type) and Mexico; R. australis Thompson from Brazil and Paraguay (type); and the critical characters are illustrated. Key Words: taxonomy, identification key, neotropics, nearctic RESUMEN El genero de la mosca de la flor del genero, Rhopalosyrphus, (DIPTERA: Syrphidae) es revi- sada y es redescrito; se presenta una clave para las especies; la relacion filogenetica del ge- nero y las especies es formulada; las especies incluidas son descritas; con las nuevas especies, R. ramulorum Weems & Deyrup, descrita de Florida (tipo) y Mexico; R. australis Thompson de Brasil y Paraguay (tipo); y los caracteres criticos son ilustrados. Translation provided by author. Rhopalosyrphus Giglio-Tos is a small group of microdontine flower flies restricted to the New World subtropics and tropics, ranging from south- ern United States to northern Argentina. These flies are rarely collected, only some two dozen specimens are known. The adults mimic eume- nine vespids, such as Zethus, which nest in twigs (Bohart & Stange 1965). The immature stages are known for only one species. These were found in ant nests (Pseudomyrmex) in twigs and grass culms, where the larvae probably prey on ant brood. The genus contains only three species. One wide ranging species, R. guentherii, is found from southwestern United States to northern Argen- tina. The others are more restricted in their ranges; R. ramulorum, presently known from a few specimens from Florida and Mexico, and R. australis from southeastern Brazil, Peru and Par- aguay. The genus is here revised, with complete synonymies, descriptions, and distributional and biological data given for all taxa. Adult terminol- ogy follows Thompson (1999), larval terminology follows Hartley (1961) and Rotheray (1991). GENUS RHOPALOSYRPHUS GIGLIO-TOS Rhopalosyrphus Gilgio-Tos 1891: 3. Type spe- cies, Holmbergia guentherii Lynch Arribalzaga (subsequent monotypy, Giglio-Tos 1892a: 2). Willis- ton 1892: 78 (catalog citation, descriptive note); Giglio-Tos 1892b: 34 [journal (1893:130] (descrip- tion); Aldrich 1905:347 (catalog citation); Kertesz 1910: 360 (catalog citation); Hull 1949: 312, figs, (description, figures of habitus, head, abdomen, hind leg); Capelle 1956 (review, key); Cole & Schlinger 1969: 307 (descriptive notes); Thomp- son et al. 1976: 60 (catalog citation); Vockeroth & Thompson 1987: 729 (key reference). Holmbergia Lynch Arribalzaga 1891: 195. Type species, guentherii Lynch Arribalzaga (monotypy). Synonymy by Giglio-Tos (1892a). Head: face convex, produced anteroventrally, pilose; gena small, linear, pilose; frontal promi- nence absent, antenna inserted above middle of head; frons short, about yA as long as face, as wide as face (?) or slightly narrowed dorsally (8), pi- lose; vertex broad, about 3 times as long as frons, as wide as frons, not swollen, pilose and punctuate; ocellar triangle small, equilateral, well separated from eye margins; occiput broad on dorsal Vr, eye bare, dichoptic in male. Antenna elongate, longer than face; scape and basoflagellomere elongate, at least 4 times as long as pedicel; scape about 6 times as long as broad; arista bare, inserted baso- laterally on mesal surface, about as long as scape. Thorax: longer than broad; postpronotum pi- lose; meso-anepisternum with anterior portion not differentiated, uniformly pilose; meso-katepister- num completely pilose; meso-anepimeron with posterior portion bare; meropleuron with barrette Weems et al.: Genus Rhopalosyrphus (Diptera: Syrphidae) 187 pilose; metasternum developed, pilose (although reduced in some species); scutum punctuate, with appressed pile; metatibia expanded apically; scutellum with or without small apical calcar, without distinct ventral pile fringe. Wing: brown on anterior 1/3, extensively microtrichious; mar- ginal cell broadly open; stigmatic crossvein present; vein Ml with apical portion straight, joining vein R4+5 perpendicularly; vein M2 present or absent; vein R4+5 with spur. Abdomen: petiolate; 1st segment short; 2nd segment as broad as thorax basally, but con- stricted, cylindrical apically; 3rd segment cylin- drical; 4th and 5th segments forming a compact club; aedeagus bifid. Puparium: elongate with broader ventral than dorsal surface; marginal band of variously-sized setae; dorsal surface flat; ventral surface convex; marginal band notched anteriorly; prothorax and mesothorax hidden beneath metathorax; mandi- ble with serrate ventral margin. Rhopalosyrphus belongs to the subfamily Micro- dontinae and is the sister group of Ceriomicrodon Hull, together these taxa are the sister group of Microdon Meigen, sensu lato. Rhopalosyrphus is defined (synapomorphy) by its 1) abdominal struc- ture and 2) pilose meropleuron. Other diagnostic characters are 3) antenna elongate, longer than face, usually about twice as long; 4) scape and ba- soflagellomere elongate; 5) face produced ven- trally; 6) occiput greatly developed on dorsal 1/3; 7) metasternum developed, not reduced; and 8) metatibia flared apically. The relationship to Micr- odon, sensu lato is unresolved: Rhopalosyrphus shares with the Microdon clade the bifid aedeagus and appears closely related to Ceriomicrodon. Ce- riomicrodon shares characters 3, 4, 5, 6, 7, 8 and its abdominal shape could be considered derived from that of Rhopalosyrphus. The two differ only by the presence of pile on the meropleuron. Microdon aurcinctus, described by Sack (1921: 138) in Rhopalosyrphus, belongs to the Pseudomi- crodon group of Microdon. The species of this group differ from Rhopalosyrphus in the charac- ters listed above and in having the vertex swollen and shiny. Based on puparial characters, the immature stages of Rhopalosyrphus closely resemble Micro- don. They both have the anterior end consisting of the metathorax, with the prothorax and mesotho- rax hidden beneath it, a marginal band of setae which surrounds the puparium except for a notch at the anterior end, sharply-pointed antennomax- illary organs and mandibles with a serrated ven- tral margin. The very distinctive shape of the puparium of Rhopalosyrphus separates it from that of Micro- don: it has a curved ventral surface that is broader than the narrow, fiat dorsal surface. Also, the whole structure is elongate rather than oval in outline. The reverse appears in Microdon, with the dorsal surface being broader and curved and the ventral surface narrower and flat. These dif- ferences in shape suit the larva to life in hollow twigs and grass culms in which its prey, larvae and pupae of the ant, Pseudomyrmex, live. 2. KEY TO SPECIES OF RHOPALOSYRPHUS 3rd tergum short, about 1/3 as long as 2nd; 2nd tergum elongate (Fig. 9); eye with an area of enlarged ran- matidia medially and posterior to antenna australis 3rd tergum elongate, as long as 2nd; 2nd tergum not greatly elongate posteriorly (Fig. 10); eye without en- larged ommatidia 2 Alula completely microtrichose; cell R extensively microtrichose, bare only on basoposterior 1/4 or less; metasternum appearing bare, with pile greatly reduced; face and anepisternum partially black pilose ramulorum Alula bare basomedially; cell R completely bare behind spurious vein; metasternum with long, distinct pile, not reduced; face and anepisternum entirely white pilose guentherii Rhopalosyrphus guentherii Lynch Arribalzaga Figs. 10-13 Holmbergia giintherii Lynch Arribalzaga 1891: 198, Fig. 3 (habitus) Argentina, Buenos Aires (T 2 MACN lost?). Giglio-Tos 1892a: 2 (notes), 1893: 131 [sep. 35], pi. 1, Figs. 10, lOa-b (description, figures of abdomen, wing); Aldrich 1905: 347 (catalog citation); Kertesz 1910: 360 (catalog citation); Fluke 1957: 36 (catalog cita- tion); Capelle 1956: 172, Fig. 2 (description, syn- onymy, key reference, figure of head); Thompson et al. 1976: 60 (catalog citation). Rhopalosyrphus carolae Capelle 1956: 174 A* 6 2 Arizona, Huachuca Mts., Sunnyside Canyon (HT 2 UKaL). Byers et al. 1962: 168 (HT UKaL); Wirth et al. 1965: 599 (catalog citation); Cole & Schlinger 1969: 307 (descr. note, distr. western N.A.); Thompson et al. 1976:60. NEW SYNONYM Wing length: 8.8 mm (-.'.-5.-*Jfiw* '? ? :? ?"?? v^\fes?>SK*??? Figs. 1-8. Puparium of Rhopalosyrphus ramulorum. 1-3, habitus, 1, dorsal, 2, lateral, 3, anterior. 4, anterior spi- racular process, lateral. 5, 6, posterior respiratory process. 5, dorsal view and surrounding papillae, anterior end uppermost; 6, lateral view, anterior end to the left. 7-8. Papillae from marginal band. Weems et al.: Genus Rhopalosyrphus (Diptera: Syrphidae) 191 Figs. 9-14. Features of Rhopalosyrphus. 9, abdomen, australis, dorsal; 10. Habitus guentherii, dorsal; 11, hind leg, guentherii, lateral; 12. head, guentherii, lateral; 13. abdomen, guentherii, lateral; 14. cephalopharyngeal skel- eton, ramulorum, lateral. Figures 10-13 from Hull (1949). gregated creating impression of vague pair of vittae running along dorsal surface (Fig. 1). Pos- terior respiratory process (Fig. 6): 0.3 mm long, 0.2 mm high, oval, nodulate with mid-dorsal pro- jection, surrounded by papillae, with 4 pairs of in- terspiracular setae and 3 pairs of spiracular openings (Fig. 5). Distribution: USA (Florida) south to Mexico (Chiapas). Holotype S: USA: Florida, Highlands Co., Lake Placid, Archbold Biological Station, Trail 1 SSo, 22-V-1985, Malaise Trap, M. Deyrup, depos- ited in the National Museum of Natural History (USNM), Washington. Paratypes: USA. FLOR- IDA, same locality as holotype, 5-V-1986, reared from nest of Pseudomyrmex simplex in twig of Carya floridana, M. Deyrup (1 9 USNM);..., L. L. Lampert, Jr. & H. W. Weems, Jr., 8-IV-1978 (2 6 USNM ENT 0003891-2 FSCA);..., ll-IV-1978 (1 5 USNM ENT 0003893 FSCA);..., 18-111-1975, H. W. Weems, Jr. (1 6 USNM ENT 0003894 FSCA);..., 17-IX-1979, T. A. Webber & H. W. Weems, Jr. (1 9 USNM ENT 0003895 FSCA); F. E. Lohrer, H. W. Weems, Jr., 11-15-IV-1980 (1 9 USNM ENT 0003896 FSCA),..., 21-22-IV-1980 (1 6 USNM ENT 0003897 FSCA);..., 14-15-V-1980 (1 6 USNM ENT 0003898 FSCA);..., 18-20-V- 1980 (1 6 USNM ENT 0003899 FSCA); ..., High- lands Hammock State Park, 3-IV-1965, H. Weems, Jr. (1 9 USNM ENT 0003877 FSCA);..., 27-111-1966 (1 9 USNM ENT 0003878 FSCA); Collier County, SR94, 1.8 miles south of US 41, 25-11-1992, M. Deyrup & B. Ferster, reared from nest of Pseudomyrmex ejectus in culm of Cladium jamaicense (ABS); Liberty Co., Torreya State Park, 14-V-1964, H. Weems, Jr. (1 9 USNM ENT 0003875 FSCA);..., 30-IV-5-V-1973, C. R. Artaud & H. Weems, Jr., Malaise trap, (1 S, 4 9 9 USNM ENT 0003879-83 FSCA, USNM); Alachua Co., Gainesville, Seville Heights, L. A. Stange, Black- light, 2-VII-1980 (2 9 9 USNM ENT 0003886-7 FSCA);..., l-VII-1980 (1 9 USNM ENT 0003888 FSCA);..., 5-VII-1980 (1 6 USNM ENT 0003889 FSCA);..., 30-VII-1979 (1 9 USNM ENT 0003890 FSCA); Dade Co., Ross & Castello Ham- mock, 30-111-1963, C. F. Zeiger (1 6 USNM ENT 0003884 FSCA);..., Fuch's Hammock, near Homestead, 27-29-VII-1978, Terhune S. Dickel & H. Weems, Jr. (1 9 USNM ENT 0003885 FSCA); Chekika State Recreation Area, 10-XI-1982, FD Fee (1 8 Fee). MEXICO. Morelos, 3 miles N Alpuyeka, 3400', 5 June 1959, HE Evans, 14-V- 192 Florida Entomologist 86(2) June 2003 1959, Biol. Note 601 (1 6 USNM ENT 0003876 FSCA); Chiapas, 28 miles west Cintalpa, 9-IV- 1962, F. D. Parker (1 3 USNM). Another broken 6 specimen is in the Canadian National Collec- tion and is labelled "Letitia, Colombia? (or Flor- ida)." According to Vockeroth (pers. comm.) this specimen was found in a Malaise trap which had been used both in the Florida Keys and Colombia. Rhopalosyrphus ramulorum is similar to guen- therii, but is smaller, narrower, not as robust, and has much more extensive black pile on face, scutum and anepisternum. The metasternal pile is greatly reduced and closely appressed, so the metasternum appears bare at low magnifications. Although many specimens of R. ramulorum have been collected, two of these are reared spec- imens and provide most of our insights into the natural history of the genus Rhopalosyrphus. When alive, the reared adults, like many other syrphids, bore a strong resemblance to stinging Hymenoptera. The wasp-like features of elongate antennae, narrow abdominal "petiole," pale bands and spots, and dark wings are enhanced by the wasp-like habit of holding the wings out from the body. The wings are also partially folded, so that they appear long and narrow. The general impres- sion is of a very small individual of the twig-nest- ing eumenid genus Zethus. One specimen was found in a nest of the ant Pseudomyrmex simplex in a small twig (hence the species epithet "ramulorum" the epithet to be treated as a noun in the genitive case) of Carya floridana in long unburned Florida scrub habitat. The adult emerged from its pupa the day after the twig was opened. A second specimen was in a nest of P. ejectus in a culm ofCladiumjamaicen.se; this adult also emerged a day after the nest was opened. The nests of these two species of Pseudomyrmex are kept clean and free of debris and fungi, and it is probable that the fly larvae are not scavengers, but predators feeding on ant brood. This would fit well with the known larval habits of the closely re- lated genus Microdon, (Duffleld 1981; Garnett et al. 1985). This is apparently the only known exam- ple of a predatory inquiline attacking members of the large neotropical ant genus Pseudomyrmex, though there must be others, especially among the Eucharitidae. Pseudomyrmex species are less sus- ceptible to inquilines than most ants because the nests are in plant cavities with access by only one or a few small, well-guarded holes, and the nests themselves are bare, with minimal edible detritus and no hiding places for inquilines. The holes used by Pseudomyrmex simplex and P. ejectus are much too small to permit the adult fly to escape, and it seems probable that emergence from the puparium is delayed until the nest has been broken open. Since the term "strategy" has been used extensively in discussing Microdon (Dufneld 1981), the problem of adult egress is a major flaw in the strategy of R. ramulorum. It may be, however, that there are ways an ovipositing fe- male can increase the likelihood that her offspring will be freed. Small, dead, exposed twigs are more likely to get broken off than larger twigs in the in- terior of the tree crown. Culms on the edge of a tus- sock of sedge or grass are more likely to get broken off than culms in the interior. If there is some spe- cial site selection by the female, this may explain why only two pupae were found in a 10-year study of Florida ants, a study that involved opening hun- dreds of colonies of Pseudomyrmex. This brings up the topic of apparent rarity of Rhopalosyrphus species, especially in Florida. At the Archbold Biological Station, two Townes traps running continuously for 3 years captured only one specimen. If the adults spend their time in the tops of trees or in extensive open marshes, this would explain why so few specimens appear in Malaise traps, which are usually set-up in un- derstory fryways. If R. ramulorum is actually de- pendent on chance events to release the adults, actual populations would need to be quite high for the sexes to meet, even if there were a mechanism for adult aggregation, and even if some synchro- nous emergence were provided by wind storms. Whatever the actual abundance of Rhopalosyr- phus species, the rarity of specimens in collec- tions suggests that there could be additional undiscovered species, especially in the neotropics, where the fauna of arboreal ants is large. ACKNOWLEDGMENTS This was a cooperative project: the larval taxonomy was done by Rotheray; nomenclature, literature review and adult taxonomy by Thompson; and biology by Dey- rup. After the manuscript was submitted for publication and during the review process, it was discovered that Weems was about to also describe the Florida species. Hence, Weems was added to this manuscript as a co-au- thor and his material was incorporated into it. We thank Drs. Paul Arnaud, Jr., California Academy of Sciences, San Francisco (CAS); George Byers, Snow Entomological Museum, University of Kansas (UKaL), Lawrence; Peter Cranston, University of California, Davis (UCD); Brian Pitkin, Entomology, The Natural History Museum (for- merly the British Museum (Natural History)), London (BMNH); Mauro Daccordi, Museuo Regionale di Storia Naturale, Torino (UTOR); John Gelhaus and Douglas Azuma, Academy of Natural Sciences, Philadelphia; David A. Grimaldi, the American Museum of Natural His- tory, New York (AMNH); Adriana Oliva, Museo Argentine de Ciencias Naturales Bernardino Rivadavia, Buenos Aires (MACN); Nelson Papavero, Museum of Zoology, Universidade de Sao Paulo, Sao Paulo (MZSP); J. R. Voc- keroth, the Canadian National Collection, Ottawa (CNC); Manuel Zumbado, Institute Nacional de Biodiversidad, Santo Domingo (INBIO), for permission to study material in their care. Suzanne Mangan assembled the plates, etc. We also thank Chris T. 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