A Revision of Sepedonea, a Neotropical Genus of Snail-killing Flies (Diptera: Sciomyzidae) AMNON FREIDBERG, LLOYD KNUTSON, and JAY ABERCROMBIE SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 506 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 5 0 6 A Revision of Sepedonea, a Neotropical Genus of Snail-killing Flies (Diptera: Sciomyzidae) Amnon Freidberg, Lloyd Knutson, and Jay Abercrombie SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1991 ABSTRACT Freidberg, Amnon, Lloyd Knutson, and Jay Abercrombie. A Revision of Sepedonea, a Neotropical Genus of Snail-killing Flies (Diptera: Sciomyzidae). Smithsonian Contributions to Zoology, number 506, 48 pages, 143 figures, 1991.?The Neotropical genus Sepedonea (Diptera: Sciomyzidae) is revised. Four of the 12 species (S. incipiens, S. neffi, S. trichotypa, and S. veredae) are described as new. Sepedonea vau Mello and Bredt is placed as a nomen nudum under S. guianica. A key to the adults is included, and the male and female terminalia of all species are described and illustrated. The eggs, first-, second-, and third-instar larvae, and puparia of S. guianica, S. incipiens, S. lagoa, S. lindneri, S. telson, and S. trichotypa are described and figured for the first time. These species, plus 5. guatemalana and S. isthmi, are included in keys to the mature third-instar larvae and puparia. The taxonomy, biology, and geographic distribution of the genus is summarized. The taxonomic placement of the Sepedon group is discussed. The geographical distributions of 11 species are mapped. The larvae of all reared species of Sepedonea are predators of non-operculate snails in various freshwater situations. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montaslrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Freidberg, Amnon. A revision of Sepedonea, a neotropical genus of snail-killing flics (DipteraiSciomyzidae) / Amnon Freidberg, Lloyd Knutson, Jay Abercrombie. p. cm.?(Smithsonian contributions to zoology ; no. 506) Includes bibliographical references (p. ). Supt. of Docs, no.: SI 1.27:506 1. Sepedonea?Classification. I. Knutson, Lloyd V, 1934- II. Abercrombie, Jay. III. Title. IV. Series. QL1.S54 no. 506 [QL537.S365] 591 s-dc20 [595.77F4] 90-10224 @ The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48?1984. Contents Page Introduction 1 Historical Review 1 Methods and Materials 2 Acknowledgments 2 Family SdOMYZIDAE 2 Genus Sepedonea Steyskal 2 Description 2 Adult 3 Egg 6 Larva 6 Puparium 6 Natural History 6 Discussion 6 Key to Adults of Sepedonea 9 Key to Mature Third-instar Larvae of Sepedonea 10 Key to Puparia of Sepedonea 11 Sepedonea barbosai Knutson and Bredt 11 Sepedonea canabravana Knutson and Bredt 13 Sepedonea guatemalana (Steyskal) 13 Sepedonea guianica (Steyskal) 16 Sepedonea incipiens, new species 19 Sepedonea isthmi (Steyskal) 21 Sepedonea lagoa (Steyskal) 23 Sepedonea lindneri (Hendel) 26 Sepedonea neffi, new species 29 Sepedonea telson (Steyskal) 29 Sepedonea trichotypa, new species 33 Sepedonea veredae, new species 36 Literature Cited 38 Figures 94-143 39 ui FIGURE 1.?Sepedonea lindturi, adult male, habitus. A Revision of Sepedonea, a Neotropical Genus of Snail-killing Flies (Diptera: Sciomyzidae) Amnon Freidberg, Lloyd Knutson, and Jay Abercrombie Introduction Among families of higher Diptera, the Sciomyzidae are fairly well established as a phylogenetically unified (Griffiths, 1972) and behaviorally diverse family (Berg and Knutson, 1978; Ferrar, 1987) that has more or less uniquely exploited a food resource?mollusks. Mollusks are not rigorously ex- ploited by practically any other dipterans, or indeed, by any other moderately large group of insects. To understand the predatory or parasitoid behavior of Sciomyzidae on mollusks, it is essential to have a good knowledge of the systematics of the taxa involved. This information enables accurate identifica- tion of the species and an understanding of their phylogenetic relationships. What little is known on the phylogeny of Sciomyzidae was summarized by Griffiths (1972) and Berg and Knutson (1978). In the latter paper, the authors also reviewed the biology and systematics of the family, which now includes about 550 species in 61 genera. Data on the natural history and immature stages are known for about a third of the world's species. Although most sciomyzid larvae are primarily predators or parasitoids of aquatic or terrestrial snails, a few species feed on snail egg-masses, slugs, or finger-nail clams. Some Scio- myzidae are potential agents for the biological control of snails that carry diseases (e.g. fascioliasis and schistosomiasis) (Berg and Knutson, 1978). Amnon Freidberg, Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel-Aviv University, Tel Aviv 69978, Israel. Lloyd Knutson, Biological Control of Weeds Laboratory?Europe, American Embassy?AGRIC, APO New York 09794. Jay Abercrom- bie, 10th Medical Laboratory, Department of the Army, APO New York 09180. With about 125 known species (at least 20 are undescribed), Sepedon Latreille, 1804, and related genera comprise nearly 25 percent of the family's species. The strictly neotropical genera of this group, Sepedonea Steyskal, 1973, and Thecomyia Perty, 1833, have never been revised. Our purpose in this paper is to revise the species of Sepedonea. HISTORICAL REVIEW.?Steyskal (1973) proposed the mod- ern classification of Sepedon and related genera (the Sepedon group). He included the following genera in this group: Sepedon Latreille, Sepedonella Verbeke 1950, Sepedoninus Verbeke 1950, Thecomyia Perty, and his new genera Sepe- donea and Sepedomerus. He further suggested that Sepedomyia Verbeke, 1950, is a subgenus of Sepedon, together with the previously established subgenera Mesosepedon Verbeke, 1950, and Parasepedon Verbeke, 1950. In establishing the Sepedon group, Steyskal did not believe that (p. 143) "this group is sufficiently distinct from more typical Tetanocerini, especially from such genera as Hedria and Dichetophora, to be given the rank of tribe or even subtribe." Earlier authors had classified these genera as a tribe, subfamily, or even family (e.g., Berg and Knutson, 1978). The diagnostic characters shared by all members of the Sepedon group as defined by Steyskal (1973) are as follows: 1. Lunule well exposed 2. Face more or less extended ventrally 3. Ocellar setae weak or absent 4. One pair of scutellar setae (absent in the oriental Sepedon lobifera Hendel) 5. Subalar (vallar) setae present In his study of the Western Hemisphere species of Sepedon, Steyskal (1951:272) noted that "the S. lindneri group [= SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Sepedonea] of Central and South America is unique in having the posterior forceps fused mesally to form a 'posterior process,' but in view of the diversity of form of the male terminalia within that group and the conformity otherwise with other forms, I have not thought it wise to distinguish the group nomenclatorially." Later, however, he reversed his opinion and described Sepedonea (Steyskal, 1973) and designated Sepedon lindneri Hendel as the type species of the genus. Eight species were recognized in the genus in the catalog of Neotropical Sciomyzidae (Knutson et al., 1976). These, plus four new species described herein, are now included in the genus. METHODS AND MATERIALS.?During the course of this study slightly over 1000 adult specimens from most major North and South American collections were examined, includ- ing the primary types of all but one previously described species. Label data for all correctly identified specimens we examined were recorded, organized according to locality (country, state or province), and are presented under the appropriate species. The descriptive terminology essentially follows that pub- lished in the recent Manual of Nearctic Diptera (MeAlpine, 1981 (adults); Knutson, 1987 (adults and immature stages)). Because of the paucity of characters available for descrip- tions of adults, and because the key to adults contains most of the distinguishing characters, separate diagnoses are not given for each species. ACKNOWLEDGMENTS.?Numerous individuals and institu- tions have contributed to this revision. Without their coopera- tion and assistance, much of the study could not have been completed. We are grateful for their time and thoughtful consideration. We thank the following curators and institutions for lending specimens: BMNH British Museum (Natural History), London, England (Mr. B.H. Cogan, Dr. B.R. Pitkin) CUI Cornell University, Ithaca, New York, USA. (Dr.C.O. Berg) IOC Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (Dr. L.M. Deane, Dr. D.V. Ferreira, Dr. A.P.A. Lunas Dias) IZAM Instituto de Zoologia Agricola, Maracay, Venezuela (Dr. Carlos Julio Rosoles) MNRJ Museu Nacional, Rio de Janeiro, Brazil (Dr. Hugo de Souza Lopes) MZUSP Museu de Zoologia da Universidade de Sao Paulo, S3o Paulo, Brazil (Dr. Nelson Papav- ero) OSU Ohio State University, Columbus, Ohio, USA. (Dr. Charles A. Triplehorn) SMNS Staatliches Museum fur Naturkunde, Lud- wigsberg, Federal Republic of Germany (Dr. B. Herting) USNM Collections of the United States National Museum in the National Museum of Natural History, Smithsonian Institution, Washing- ton, DC, USA. (Dr. W.N. Mathis) We thank WN. Mathis, F.C. Thompson, and R.E. Orth for reviewing the manuscript. The advice of the master scio- myzidologist, G.C. Steyskal, was appreciated throughout the study. We also thank Mary Lou Cooley, Systematic Entomol- ogy Laboratory, IIBIII, for completion of the illustrations prepared by AF, and for preparing the plates. We are also grateful to Linda Heath Lawrence for handling some graphic aspects. Most specimens of Sepedonea available for this study were collected by CO. Berg and his students and coworkers while conducting life-cycle studies in Central and South America. Most of these specimens are in the Department of Entomology, Cornell University. We feel that when systematists conduct a taxonomic study, they should, if at all possible, place specimens, especially paratypes of new species, in various collections in pertinent geographical areas, to serve as reference material. Hence, we appreciate the cooperation of Cornell University for allowing us to place paratypes and other specimens from the Cornell material in several collections. Whenever possible specimens were placed in the following collections in addition to those listed above: Universidade Federal do Parana', Curitiba, Brazil; Universidad Nacional de Colombia, Medellin, Colombia; Universidad de Panama, Panama; Fundaci6n M. Lillo, Tucuman, Argentina. Family SCIOMYZIDAE Sciomyzidae are small to moderately large acalyptrate Diptera, often rather slender, ranging from 1.8 to about 12 mm in length. The antenna is usually porrect, and the pedicel is usually elongate. Two orbital setae are usually present, but sometimes only one. The wings are usually longer than the abdomen and are hyaline, slightly spotted, or heavily patterned. The costa (C) is unbroken and the subcosta (Sc) is complete and free from Rl distally. The abdomen is moderately long and cylindrical. The general color varies from yellowish to shiny black, but often is grayish or brownish. Sciomyzidae are best recognized by the above characters and by the close association of most species with aquatic or otherwise damp and shaded habitats. Adults often rest on emergent vegetation. Genus Sepedonea Steyskal, 1973 Sepedonea Steyskal, 1973:145 [type species: Sepedon lindneri Hendel, by original designation].?Steyskal and Knutson, 1975:276 [generic key].? Knutson et al., 1976:10 [catalog].?Knutson and Bredt, 1976:113 [review].?Knutson and Valley, 1978:198 [review]. DESCRIPTION.?The following is a brief characterization of NUMBER 506 FIGURES 2-6.?Sepedonea spp., various parts: 2, S. lagoa, head in profile; 3-5, left posterior spiracle: 3, S. trichotypa; 4, 5. lindneri; 5, S. guatemalana; 6, S. is thru, wing. the various life stages of Sepedonea (first- and second-instar larvae are not included): Adult: Elongate, generally brown, Sepedon-]ike flies (Fig- ure 1). Head (Figure 2): One (posterior) pair of orbital setae, occasionally anterior orbital setae weakly developed; postocel- lar setae well developed; frons largely dull yellow with slender, shiny, frontal vitta; lacking round black spots on orbit a short distance ventral to antenna; face with dark brown spot on each ventral corner, orbit, face, and gena spotted by silvery microtomentum; antenna with pedicel 3.5-5 times as long as wide, with setulae as long as diameter of pedicel; first flagellomere with distinct, elongate, sensory pit near base; ventral part of head not forming a tube into which the proboscis may be withdrawn; palpus well developed. Thorax: Scutum with brownish gray microtomentum; SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 10 FIGURES 7-13.?Sepedonea spp., legs: 7-9, right midfemur, posterior view: 7, S. guianica; 8, S. telson; 9, 5. isthmi; 10, S. telson, right hindleg, anterior view; 11-13, right hindcoxa, posterior view: 11, S. lindneri; 12, S. trichotypa; 13, S. canabravana. pleura and coxae with silvery microtomentum, dense on pleura; pleura with numerous setulae; posterior spiracle (Figures 3-5) with a varying number of setulae of various lengths; postcoxal bridge divided by membranous area; forefemur with at least 1 outstanding dorsal seta; midfemur with large anterior seta near midlength, usually with a posteroventral row of spines (Figures 7-9) extended to varying degrees from apex toward base; hindfemur (Figure 10) often with dorsal and/or lateral, preapical dark marks (these marks are somewhat variable intraspecifically and the least reliable of the characters used for identification); hindcoxa posteriorly (Figures 11-13) with a varying number of setulae of varying length; wing (Figure 6) without spots, except sometimes with slight infuscations about crossveins; crossvein dm-cu straight, at right angle with penultimate section of vein CuAl. Abdomen (Figures 14-19): with light gray microtomen- NUMBER 506 CERC FIGURES 14-16.?Sepedonea isthmi, male abdomen: 14, lateral view; 15, ventral view; 16, dorsal view, (a = anterior plate of sternum 5 (left-hand half); a.SS = anterior surstylus; CERC = cercus; D = distiphallus; EPAN = epandrium; p = posterior plate of sternum 5 (left-hand half); p.SS = posterior surstyli; S 1-5 = sternum 1 through 5; T 1-5 = tergum 1 through 5.) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY turn; male posterior surstyli fused to form a median structure (Figures 14-15); female sterna 6-8 fused to form a synsternum (Figures 17-18). Egg: Eggs of Sepedonea are remarkably uniform from species to species (Figures 94,96), and are very similar to those described for Sepedon sphegea (Fabricius), S. spinipes (Sco- poli), and Sepedomerus macropus (Walker). Some eggs of S. telson differ from other species in having a dorsal depression anteriorly in the central area between the dorsal longitudinal ridges. It is not present in all individuals, however, so it is unreliable as a distinguishing character. Neff and Berg (1966) found the same structure in eggs of Sepedomerus macropus but not in others. The known eggs of eight species of Sepedonea treated here can be divided roughly into 2 groups on the basis of size, a somewhat arbitrary division that is nonetheless reliable in most instances. One group, that includes S. incipiens, S. lindneri, and 5. trichotypa, has eggs which range from 0.90 to 1.12 mm long. Sepedonea incipiens is the only species in this group with eggs as large as 1.12 mm; only one specimen of 10 examined reached this length. Other species in the first group (S. lindneri and S. trichotypa) have eggs with lengths of 0.90-1.06 mm. The second group, which includes S. guatemalana, S. guianica, S. isthmi, S. lagoa, and S. telson, has eggs ranging from 1.12 to 1.40 mm in length. Sepedonea lagoa is the only species in the second group with eggs as small as 1.12 mm; of 17 individuals examined, 5 were 1.12 mm long and the remainder were considerably longer. Other species in the second group have eggs with lengths of 1.16-1.24 mm (S. guianica and S. telson), 1.26-1.40 mm (S. guatemalana; Neff and Berg, 1966) and 1.28-1.38 mm (5. isthmi; Knutson and Valley, 1978). The micropyle is subterminal in all eight species. Larva: The third-instar larva of 5. guianica, S. incipiens, S. isthmi, S. lagoa, S. lindneri, S. telson, and S. trichotypa has smooth posterior stigmatic tubes that are distinctly scalloped basally (Figure 131). Neff and Berg (1966) reported that in S. guatemalana (as well as in species of Sepedomerus and Sepedon), surfaces of the stigmatic tubes are tuberculate or papillose; hence, the tubes may appear scalloped basally because of these projections. All third-instar larvae of Sepedonea except S. lagoa have a prominent middorsal stripe and dorsolateral V-shaped marks anteriorly. Larvae of Sepedomerus and Sepedon do not have dorsolateral markings. Dorsolateral patch of setulae occur on segments 5-11, except in S. guatemalana, S. guianica, and S. isthmi, which lack them on segment 11. All species of Sepedomerus and some of Sepedon have dorsolateral patches of setulae on segments 5-10. Larvae of Sepedonea, Sepedomerus, and Sepedon all possess 5 pairs of lobes on the posterior spiracular disc (Figures 127-130,135). The most reliable character to separate larvae of Sepedonea from Sepedomerus and Sepedon is the dorsal most accessory tooth of the cephalopharyngeal skeleton. In all 3 genera, the accessory teeth are directed mesally (Neff and Berg, 1966; Knutson and Valley, 1978) (Figures 107, 109-110). However, in Sepedonea the dorsal tooth is larger and darkly sclerotized; the remaining teeth are smaller and only lightly sclerotized. In Sepedomerus and Sepedon the accessory teeth are subequal in size and evenly sclerotized, usually lightly. Puparium: As pointed out by Neff and Berg (1966) for Sepedon, most external features of puparia are more useful for species recognition than for distinguishing the genus from other Tetanocerini. The same is true of Sepedonea. No characters of the puparia themselves are as dependably diagnostic for the genus Sepedonea as the dorsalmost accessory tooth of the third-instar larva, as discussed above. This may be examined by removing the third-instar larval cephalopharyn- geal skeleton from the inner surface of the ventral cephalic cap. All puparia of Sepedonea except 5. lagoa have more or less prominent dorsolateral V-shaped marks anteriorly (Figures 95, 97-100). The puparium of S. lagoa lacks dorsolateral V-shaped marks; it also is considerably different in shape and has prominent ventral lobes on the posterior spiracular disc (Figure 101). NATURAL HISTORY.?Behaviorally, the reared species of Sepedonea, like almost all species of the Sepedon group, are typical predators of non-operculate snails in various freshwater situations (Figures 20-21). There is biological information available on all species except S. neffi and S. veredae. References to the biology and descriptions of immature stages of the reared species are included in the synonymy of the species. The immature stages described in the present paper were all previously described by Abercrombie (1970) in his Ph.D. dissertation. Mello and Bredt (1978) presented informa- tion on the monthly variation in abundance of four species of Sepedonea (5. barbosai, S. canabravana, S. telson, and "5. vau" (= S. guianica (Steyskal)) and Sepedomerus bipuncticeps (Malloch) in Brazil. DISCUSSION.?The monophyly of Sepedonea is established by the following synapomorphies: (1) face with dark brown spot on each ventral corner; (2) forefemur with at least one outstanding dorsal seta; (3) male with posterior surstyli fused to form a median structure; and (4) female sterna six through eight are fused to form a synsternum. Sepedonea is strictly neotropical and appears to be most closely related to Sepedon, the species of which are found in all major zoogeographical regions. Sepedon is primarily distin- guished from Sepedonea as follows: the face lacks spots in the ventral corner; the forefemur lacks an outstanding dorsal setae; the male surstyli are well separated; and the female stema six through eight are separate (in Sepedonea they form a synsternum). No species of Sepedonea has an hydrostatic sperm pump as found in some Afrotropical and Oriental species of Sepedon. The species of Sepedonea are very similar externally, and identifications are most reliably based on the terminalia, especially those of the males. Various external characters and the female terminalia are useful in the identification of certain species, and both external and terminalia characters are used in the key to species. NUMBER 506 T7 18 19 FIGURES 17-19.?Sepedonea veredae, female abdomen: 17, lateral view; 18, ventral view; 19, dorsal view. (CERC = cercus; EP = epiproct; HY = hypoproct; S 2-5 = sternum 2 through 5; SYNST = synstemum; T 1-8 = tergum 1 through 8.) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGURES 20-21.?Habitats that are collecting sites for species of Sepedonea. 20, Rio Preto, 70 km northeast of Brasilia, Goiis, Brazil; November 1974; collecting site for S. canabravana, S. guianica, and S. isthmi. 21, Lago de Pedra, west of Rio Cana Brava, 160 km northeast of Brasilia on Brasflia-Fortaleza Highway (BR-020), Goiis, Brazil; October 1974; collecting site for S. barbosai, S. canabravana, and S. telson. NUMBER 506 Key to Adults of Sepedonea 1. Midfemur posteroventrally with 8-15 spines extended 2/3 distance to base (frequently, the basad spines becoming weak and hardly distinguishable from apical setulae in the same row) [Figure 9]; hindfemur without preapical dark marks; wing usually with crossveins r-m and dm-cu distinctly clouded, especially at junction with vein M [Figure 6]. Male terminalia: posterior margin of sternum 4 smoothly curved, without protuberances; posterior surstylus with median lobe triangular, large; with lateral lobes curved anterad. Female synstemum: posterior margin slightly concave (Panama", Trinidad, Venezuela, Colombia, Bolivia, Brazil) S. isthmi Midfemur posteroventrally usually with less than 8 spines, extended no more than 72 distance to base; other characters variable 2 2. Hindfemur with more or less discrete, usually dark, dorsal preapical mark [Figure 10] 3 Hindfemur without dark, dorsal preapical mark, although this area may be diffusely reddened 5 3. Dorsal mark on hindfemur elongate. Male terminalia: posterior margin of sternum 4 straight, with 1 pair of approximate posterior tubercles; a subequal ventral pair at ends of interior apodome and just anterior to posterior pair, and a larger ventral pair lateral of these; posterior surstyli with median lobe moderately developed and with large lateral lobes moderately curved anterad. Female synsternum: posterior margin distinctly projected in middle, with apical xh distinctly narrowed; in lateral view with apical xh attenuate, pointed, and with posterodorsal shoulder (Brazil). S. telson Dorsal mark on hindfemur rounded; male and female terminalia d i f f e r e n t . . . . 4 4. Setulae near posterior thoracic spiracle numerous, long and black, obscuring spiracular opening similar to Figure 3. Male terminalia: posterior margin of sternum 4 sclerotized, straight, with median projection bearing a pair of tubercles, posterior surstyli with median lobe low and rounded; lateral lobe moderately curved anterad. Female synsternum: apically with 3 subequal tubercles (Brazil and Argentina) S. incipiens, new species Setulae near posterior thoracic spiracle fewer and shorter, not obscuring spiracular opening [Figure 4]. Male terminalia: posterior margin of sternum 4 deeply emarginate, with pair of small, posteromesally directed protuberances; posterior surstyli with median lobe emarginate ventrally, with lateral lobe slightly curved anterad. Female synstemum: posterior margin emarginate, with 2 large lateral lobes (Brazil, Paraguay, Argentina) S. lindneri 5. Hindfemur with lateral preapical marks 6 Hindfemur without preapical marks 8 6. Mesonotum yellowish; hindcoxa posteriorly with row of long setulae, mostly longer than setulae on dorsum of abdomen [Figure 12]. Male terminalia: posterior margin of sternum 4 straight, with short, truncate median lobe that is slightly bifid at apex; posterior surstyli with median lobe short, with lateral lobe strongly curved anterad. Female synsternum: short, about 1.6 times as wide as high, in lateral view with distinct ventral convexity, with dorsal margin receding gradually (forming broad apical point), with more or less distinct shoulder (Brazil and Argentina) . S. trichotypa, new species Mesonotum grayish black; hindcoxa with or without long setulae; male and female terminalia different 7 7. Hindcoxa posteriorly with dense, long setulae [Figure 13]. Male terminalia: posterior margin of sternum 4 more or less straight, with median lobe; posterior surstyli with median lobe triangular. Female synsternum: posterior margin 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY apically with rather inconspicuous median convexity and lateral shoulders; in lateral view ventral surface almost straight; posterior margin with apex and shoulder short and rounded (Brazil) S. canabravana Hindcoxa posteriorly with a few short setulae only as in Figure 11. Male terminalia: posterior margin of sternum 4 gently emarginate, with median lobe; posterior surstyli with median lobe large and lateral lobes short and narrow. Female synsternum: gradually pointed apically, in lateral view with ventral convexity, posterior margin without ridge (Venezuela, Brazil) . . . . S. neffi, new species 8. Hindcoxa posteriorly with row of setulae that are longer than setulae on dorsum of abdomen as in Figure 12; crossvein r-m clouded at junction with vein M; setulae near posterior thoracic spiracle moderately strong. Male terminalia: posterior margin of sternum 4 straight, with median, hooklike process; posterior surstyli without median lobe, lateral lobe acuminate, strongly curved anterad. Female synsternum: posterior margin with broad, straight apex; in lateral view ventral surface gently convex and posterior margin with small ridge (Brazil) S. barbosai Hindcoxa posteriorly with long setulae restricted mesally as in Figure 11; crossvein r-m clouded or not; setulae near posterior thoracic spiracle weak; male and female terminalia different 9 9. Mesonotum usually yellowish; wing uniformly hyaline. Male terminalia: posterior margin of sternum 4 gently concave, without protuberances; posterior surstyli with median lobe undeveloped, with lateral lobe strongly curved anterad. Female synsternum: posterior margin very slightly concave; posterodorsal ridge project- ing beyond posterior margin (Brazil) S. veredae, new species Mesonotum grayish black; wing usually clouded 10 10. Setulae near posterior thoracic spiracle moderately strong as in Figure 5. Male terminalia: posterior margin of sternum 4 deeply emarginate, with pair of posteriorly directed protuberances; posterior surstyli with median lobe rounded. Female synsternum: posterior margin distinctly projecting in middle; in lateral view with ventral surface more or less straight and with distinct posterior ridge (Surinam, Guiana, Venezuela, Colombia, Brazil, Argentina) S. guianica Setulae near posterior thoracic spiracle weak as in Figure 4. Male and female terminalia different 11 11. Wing distinctly clouded, especially around distal half of vein R2+3; crossvein r-m slightly clouded. Male terminalia: posterior margin of sternum 4 concave; posterior surstyli in form of narrow, elongate, anteriorly directed rod. Female synsternum: posterior margin pointed apically; with large light depression centrally; in lateral view with distinct ventral and posterior shoulders (Costa Rica, Surinam, Brazil) S. lagoa Wing not clouded, or clouded only around end of longitudinal veins; crossvein r-m not clouded. Male terminalia: posterior margin of sternum 4 slightly convex with short median, triangular lobe and pair of large mammilliform to lobelike, ventrally directed protuberances; posterior surstyli without median lobe, with lateral lobe long, strongly curved anterad. Female synsternum: posterodorsal margin straight; in lateral view without shoulders, but with posterior ridge projecting slightly beyond posterior margin (Mexico, Guatemala, Nicaragua, Costa Rica) S. guatemalana Key to Mature Third-instar Larvae of Sepedonea 1. Dorsolateral patches of setulae found only on segments 5-10 2 Segments 5-11 each bearing a prominent dorsolateral patch of setulae 4 2. Middorsal stripe incomplete; stigmatic tubes tuberculate or papillose S. guatemalana NUMBER 506 11 Middorsal stripe complete; stigmatic tubes smooth, scalloped basally [Figures 127-130, 135] 3 3. Lateral tubercles on segments 5-10 bearing short setulae and/or a single seta . . . S. isthmi Lateral tubercles on segments 5-10 with no setulae or setae S. guianica 4. Middorsal stripe and dorsolateral V-shaped marks lacking [Figures 106, 107,115, 118, 127, 132] S. lagoa With middorsal stripe and dorsolateral V-shaped marks anteriorly 5 5. Posterior spiracular plates black as in Figures 129 and 130 6 Posterior spiracular plates not black as in Figures 128 and 131 7 6. Epipharyngeal sclerite small, lightly sclerotized as in Figure 117 S. incipiens, new species Epipharyngeal sclerite larger, darkly sclerotized as in Figure 119 . . . . S. telson 7. Hypopharyngeal sclerite wide as in Figure 115; ventral arch as in Figure 112 . . . S. tindneri Hypopharyngeal sclerite long and narrow [Figure 114]; ventral arch as in Figure 111 S. trichotypa, new species Key to Puparia of Sepedonea 1. No dorsolateral marks [Figure 101]; elongate S. lagoa With at least 2 dorsolateral V-shaped marks anteriorly; roughly barrel-shaped. . 2 2. Found in Mexico, ranging southward to Costa Rica 5. guatemalana Found in South America and/or Panamd 3 3. Ground color light, with black markings as in Figures 95 and 97 4 Ground color dark, with light markings as in Figures 98-100 6 4. Short, less than 5 mm long S. tindneri Longer, ranging from 5.2 to 6.0 mm in length 5 5. Labial sclerite gently arcuate, lying loosely on median projection of hypopharyn- geal sclerite as in Figure 115 S. isthmi Labial sclerite strongly arcuate, lying wrapped around median projection of hypopharyngeal sclerite [Figures 97 and 116] S. guianica 6. Anterior segments slightly upturned from longitudinal body axis; small, usually under 4.5 mm [Figure 100] S. incipiens, new species Anterior segments parallel to longitudinal body axis; usually larger, above 4.5 mm [Figures 98-99] 7 7. Two very prominent, light-colored dorsolateral V-shaped marks anteriorly [Figure 99]; hypopharyngeal sclerite long and narrow [Figure 114] S. trichotypa, new species Usually 3 inconspicuous dorsolateral V-shaped marks anteriorly, sometimes almost completely obscured [Figure 98]; hypopharyngeal sclerite wide, as in Figure 115 S. telson Sepedonea barbosai Knutson and Bredt FIGURES 22-27,139 Sepedonea barbosai Knutson and Bredt, 1976:114.?Bredt and Mello, 1978:767 [biology].?Knutson and Valley, 1978:198 [review].?Mello and Bredt, 1978:1459 [phenology]. ADULT.?Head: Lateral facial spot usually small. Thorax: Mesonotum grayish black; postpronotum yel- lowish; setulae near posterior spiracle moderately dense and strong. Legs: Midfemur posteroventrally with 6-8 spines, not extended beyond half distance to base; hindcoxa posteriorly with row of long setulae that are longer than setulae on dorsum of abdomen; hindfemur lacking dark preapical marks. Wing: Brownish with anteroapical margin and crossvein r-m clouded; length 5.5 mm. Abdomen: Male terminalia: Posterior margin of sternum 4 straight, with median, hooklike process (Figur?. 22); anterior plate of sternum 5 strongly concave, without fingerlike projections (Figure 22); distiphallus (Figure 23) strongly 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 22 23 24 25 FIGURES 22-27.?Sepedonea barbosai: 22, male, sterna 3-5; 23, distiphallus, lateral view; 24, posterior surstyli, anterior view; 25, same, lateral view; 26 female, synsternum, ventral view; 27, same, lateral view. NUMBER 506 13 curved, with sinuous posteroventral process; anterior surstylus moderately elongate; posterior surstyli without median lobe (Figure 24), and with lateral lobe acuminate and strongly curved anterad (Figure 25). Female synsternum: In ventral view posterior margin with broad, straight apex (Figure 26); in lateral view ventral surface gently convex and posterior margin with small ridge (Figure 27). TYPE SPECIMENS.?Holotype d": BRAZIL. GoiAs: Cana Brava, 160 km NE Brasilia, 9 January 1974, D. Barbosa, MZUSP. Allotype: same data, but 30 October 1974, [abdomen dissected], MZUSP. Paratypes: BRAZIL. GoiAs: Cana Brava, 160 km N Brasilia, 23 October 1974, Knutson and Bredt, Id"; 30 October 1974, Knutson and Bredt, 4cf, USNM. OTHER SPECIMENS EXAMINED.?None. ADDITIONAL RECORDS FROM LITERATURE.?BRAZIL. Dis- TRITO FEDERAL: Brasilia, Lagoa Paranoa, 5 November 1974, Knutson and Bredt (Knutson and Bredt, 1976). Highway L 2 Norte, Brasilia, 5 November 1974, Mello and Bredt. GoiAS: Lagoa do Piripiri, Formosa, 30 October 1974, Mello and Bredt. Lagoa do Golfe, Formosa, 11 February 1974, Mello and Bredt. Lagoa das Pedras, Formosa, 9 January 1974, January- December 1975, January-February, and June-November 1976, Mello and Bredt (Bredt and Mello, 1978; Mello and Bredt, 1978). IMMATURE STAGES.?Unknown. REMARKS.?The similarity between S. barbosai and S. canabravana in both the male and female terminalia suggests a close relationship between the two species. Both species have a similar distribution, in central Brazil, that is the most restricted of any species of Sepedonea (Figures 139,140). Sepedonea canabravana Knutson and Bredt FIGURES 13,28-33,140 Sepedonea canabravana Knutson and Bredt, 1976:114.?Bredt and Mello, 1978:767 [biology].?Knutson and Valley, 1978:198 [review].?Mello and Bredt, 1978:1459 [phenology]. ADULT.?Head: Lateral facial spot rather elongate. Thorax: Mesonotum grayish black, postpronotum yel- lowish; setulae near posterior spiracle dense and strong. Legs: Midfemur posteroventrally with 5-6 spines, not extended beyond half distance to base; hindcoxa (Figure 13) posteroventrally with 2 irregular rows of long setulae that are longer than setulae on dorsum of abdomen; hindfemur with dark lateral preapical marks, posterior mark larger and darker than anterior mark. Wing: Brownish yellow with anteroapical margin, crossveins r-m and dm-cu clouded; length 5-5.5 mm. Abdomen: Male terminalia: Posterior margin of sternum 4 more or less straight, with large, median hooklike lobe (Figure 28); anterior plate of sternum 5 with 2 fingerlike projections (Figure 28); distiphallus (Figure 29) moderately curved, with short posteroventral spur; anterior surstylus small; posterior surstyli with short, triangular, median lobe (Figure 30) and with lateral lobe moderately curved anterad (Figure 31). Female synsternum: In ventral view posterior margin apically with rather inconspicuous median convexity and lateral shoulders (Figure 32); in lateral view ventral surface almost straight; posterior margin with apex and shoulder short and rounded (Figure 33). TYPE SPECIMENS.?Holotype d": BRAZIL. GoiAs: Rio Preto, 70 km NE Brasilia, 21 October 1974, L. Knutson [abdomen dissected], MZUSP. Allotype: BRAZIL. GOIAS: Rio Preto, 30 km NE Brasflia, 7 November 1974, Bredt and Knutson [abdomen dissected], MZUSP. Paratypes: BRAZIL. GOIAS: Rio Preto, 30 km NE Brasilia, 7 November 1974, Bredt and Knutson, Id". Lagoa de Pedra, W Rio Cana Brava, 160 km NE Brasilia, Brasilia-FortalezaHwy. (BR-020), 25 April 1974, D. Barbosa, Id", (This is erroneously reported as "Lagoa Preta" in Knutson and Bredt, 1976), USNM. OTHER SPECIMENS EXAMINED.?None. ADDITIONAL RECORDS FROM LITERATURE.?BRAZIL. GOIAS: Rio Preto, DF 06, Formosa, 15 September 1975, Mello and Bredt. Lagoa das Pedras, Formosa, 25 April 1974, January-December 1975, January-December 1976, Mello and Bredt (Bredt and Mello, 1978; Mello and Bredt, 1978). IMMATURE STAGES.?Unknown. REMARKS.?See remarks under 5. barbosai. Sepedonea guatemalana (Steyskal) FIGURES 5,34-39,140 Sepedon guatemalana Steyskal, 1951:293.?Neff and Berg, 1966:41 [biology and immature stages]. Sepedonea guatemalana.?Steyskal 1973:145 [list].?Knutson et al., 1976:11 [catalog].?Knutson and Valley, 1978:198 [review]. ADULT.?Head: Lateral facial spot small to large. Thorax: Mesonotum grayish black; postpronotum yel- lowish; setulae near posterior spiracle weak and sparse (Figure 5). Legs: Midfemur posteroventrally with 3-6 spines; hind- coxa posteriorly with long setulae restricted mesally, with or without short setulae laterally; hindfemur without dark preapi- cal marks. Wing: Grayish; crossveins r-m and dm-cu not clouded; length 4.5-6 mm. Abdomen: Male terminalia: Posterior margin of sternum 4 slightly convex, with short, median lobe and pair of large, mammilliform, ventrally directed protuberances (Figure 34); anterior plate of sternum 5 without fingerlike processes (Figure 34); distiphallus (Figure 35) slightly curved, with rather long ventral process and hairy epiphallus; anterior surstylus very small or indistinct; posterior surstyli without median lobe (Figure 36), and with lateral lobe long, strongly curved anterad (Figure 37). Female synsternum: In ventral view posterior margin straight (Figure 38), in lateral view with 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 29 28 31 32 33 FIGURES 28-33.?Sepedonea canabravana: 28, male, sterna 3-5; 29, distiphallus, lateral view; 30, posterior surstyli, anterior view; 31, same, lateral view; 32, female, synslernum, ventral view; 33, same, lateral view. NUMBER 506 15 34 36 35 37 38 39 FIGURES 34-39.?Sepedonea guatemalana: 34, male, sterna 3-5; 35, distiphallus, lateral view; 36, posterior surstyli, anterior view; 37, same, lateral view; 38, female, synstemum, ventral view; 39, same, lateral view. 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY shallow ventral depression, and with posterior ridge projecting slightly beyond posteroventral margin (Figure 39). TYPE SPECIMEN.?Holotype d": GUATEMALA. Los Amates, 18-28 February 1905, Ja[me]s. S. Hine, [red label], OSU. OTHER SPECIMENS EXAMINED.?HONDURAS. FRANCISCO MORAZAN: Zamorano, nr. Tegucigalpa, 19, 20 November 1987, L. Knutson, 37c?, I9. MEXICO. CHIAPAS: Las Cruces, 16 July 1958, SB. Neff, 12d", 59 (field collected), 7d \ 4$ (laboratory reared). NICARAGUA. Managua, April 1943, lcT, D. Gilail, USNM. ADDITIONAL RECORDS FROM LITERATURE.?NICARA- GUA. Chinandega, Baker, Id1 (Steyskal, 1951). GUATE- MALA. SANTA ROSA: 2.1 km E Barbarena (1,341 m), 23 July 1958, 2 d \ 1$. COSTA RICA. SAN JOSE: San Antonio- Desamparados (1,160 m), 16 June 1964,4d", 19 (both Neff and Berg, 1966). IMMATURE STAGES.?See Neff and Berg, 1966. REMARKS.?This species is found from Chiapas (Mexico) to Costa Rica (Figure 140) and thus is the northernmost representative of the genus and the only species not occurring in South America. With a lack of accepted phylogeny for the genus, this distribution may either suggest that the species has evolved relatively late or, conversely, that it is an early offshoot of the genus and a sister group to all remaining species. The shape of the posterior surstyli, which are only weakly fused, supports the latter possibility. Neff and Berg (1966) studied the immature stages. Sepedonea guianica (Steyskal) FIGURES 7.40-45,97,110,113,116.126.135,138.141 Septdon guianica Steyskal, 1951:295. Sepedonea guianica.?Steyskal, 1973:145 [list].?Knutson et al., 1976:11 [catalog].?Knutson and Valley, 1978:198 [review]. Sepedonea vau.?Mello and Bredt, 1978:1459 [nomen nudum; phenology]. ADULT.?Head: Lateral facial spot large. Thorax: Mesonotum grayish black; postpronotum brow- nish; setulae near posterior spiracle moderately strong and dense. Legs: Midfemur posteroventrally usually without spines but with setulae only, sometimes setulae near apex of femur distinctly stronger than basal setulae (Figure 7), and in some Brazilian specimens there are 3-6 spines, not extending beyond half distance to base; hindcoxa posteriorly with short setulae, mainly restricted mesally; hindfemur usually without dark preapical marks. Wing: Grayish, with crossveins r-m and dm-cu not clouded, or brownish and clouded anteroapically and over the crossveins; length 5-6 mm. Abdomen: Male terminalia: Posterior margin of sternum 4 deeply emarginate, with pair of posteriorly directed protuber- ances (Figure 40); anterior plate of sternum 5 with wide, indented flange at posteromesal margin (Figure 40); distiphal- lus (Figure 41) sinuous, with posteroventral angle covered by large flat, hairy epiphallus; anterior surstylus indistinct; posterior surstyli with (Figure 42a) or without (Figure 42b) median lobe; median lobe, when present, rounded; posterior surstyli with lateral lobe moderately strongly curved anterad (Figure 43a, b). Female synsternum: In ventral view posterior margin distinctly projecting in middle (Figure 44); in lateral view with ventral surface more or less straight, and with posterior ridge distinctly projecting (Figure 45). TYPE SPECIMEN.?Holotype d": Comoro Is., Maroni, Guyane, Purch. E. Le Moult, B.M. 1933-189, Type H. T. [round label with red margin], [red label], BMNH. This locality refers to an island in the Maroni River (= Marowijne River), which is the border between Surinam and Guiana. The river is about 150 km in length, and the exact location is not known. OTHER SPECIMENS EXAMINED.?COLOMBIA. VALLE DEL CAUCA: Morga, 20 km SE Univ., 22-25 June 1964, CO. Berg, 16d\ 129. 6.5 km SE Cali (Navarro), 11 June 1969, Karl R. Valley, Id" (field collected), Id", 19 (laboratory reared). 14 June 1969, Karl R. Valley, 2d", 29,1 puparium. 5 km SE Cali, near Navarro, 11 June 1969, Karl R. Valley, 19. VENE- ZUELA. COJEDES: L. Taguanes, near Tinaquillo, 13 April 1972, L. Knutson, Id". BRAZIL. MINAS GERAIS: Lagoa Santa, near Belo Horizonte, 18 January 1977, L. Knutson, l id", 69. 17 km N Belo Horizonte, 18-23 July 1964, CO. Berg, 49 (field collected), 1 puparium (laboratory reared). Jockey Club, 23 August 1966, CO. Berg, Id1, 15 September 1966, CO. Berg, 29. Serra Verde, 30 km E Belo Horizonte, 22 August 1974, A. Bredt, Id", 10 October 1974, A. Bredt, Id". PARANA: Praia do Leste, 4 May 1967, Berg and Abercrombie, 3d1, 17 May 1967, Berg and Abercrombie, Id", 1 puparium. 4,17 May 1967, Berg and Abercrombie, 3d" (field collected), 4d", IO9 (laboratory reared). 61 km S Curitiba, Rio Varzea, 16 May 1967, Berg and Abercrombie, 2d". SAo PAULO: Sao Vicente, Parque Bitaru, 29 May 1967, Berg and Abercrombie, Id", 19 (field collected), 8 eggs (laboratory reared). Rio Claro, 14 January 1977, L. Knutson, Id". S3o Jose" do Rio Preto, 27 July 1966, N. Papavero, 9d". GoiAs: Rio Preto, 70 km NE Brasilia, 7 November 1974, Bredt and Knutson, Id". ESPIRTTO SANTO: Baixo Guandu, October 1970, P C Elias, Id". ARGENTINA. TUCUMAN: Monteros, 7 February 1967, Berg and Abercrom- bie, Id", (all USNM). ADDITIONAL RECORDS FROM LITERATURE.?BRAZIL. PA- RANA: Campina Grande, 2 August 1975, Mello and Bredt. DISTRTTO FEDERAL: Nucleo Bandeirantes, 11 November 74, Mello and Bredt. Highway L 2 Norte, Brasilia, 9 January 74, Mello and Bredt. Riberao Extrema, DF 21, 6 February 1974, Mello and Bredt. MINAS GERAIS: Hip6dromo Serra Verde, Santa Luzia, 24 August 1974, Mello and Bredt. GoiAs: Rio Preto, DF 06, Formosa, 19 June 1974, Mello and Bredt. Lagoa das Pedras, Formosa, 9 January 1974, January-May, July- October, December 1975, January-February, April, June, July, October-December 1976, Mello and Bredt (all from Mello and Bredt, 1978). IMMATURE STAGES.?Egg: White. Length 1.20-1.24 mm (average = 1.24); greatest width 0.32-0.44 mm (average = NUMBER 506 17 44 FIGURES 40-45.?Sepedonea guianica: 40, male, sterna 3-5; 41, distiphallus, lateral view; 42, posterior surstyli, anterior view, two variations; 43, same, lateral view, two variations; 44, female, synstemum, ventral view; 45, same, lateral view. 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 0.40). Larger than S. incipiens, but otherwise very similar in shape, ridging, reticulation, and subglobular ends. (Based on 9 specimens: Praia do Leste, Paran?, Brazil). First-instar Larva: White. Integument transparent. Length 1.2-2.9 mm (average = 1.9); greatest width 0.3-0.7 mm (average = 0.4). Cephalopharyngeal skeleton 0.28-0.31 mm long; mandible 0.05-0.06 mm long, with 3 component parts; very similar to that of other Sepedonea. Indentation index 29-38. Segment 1 bilobed anteriorly, each lobe bearing a sensory papilla. Segments 2-4 each with 1 seta dorsolaterally, 1 laterally, and 1 ventrally, all quite small. Segments 5-10 each with dorsolateral patch of setulae; segment 11 with single seta dorsolaterally; segments 5-11 each with lateral tubercle group arrangement as in other Sepedonea; none with setae; main ventral tubercle group with 4 tubercles in transverse row, each bearing short, stout setae. Posterior spiracular disc with 5 pairs of lobes: ventral pair subconical, quite wide basally; ventro- lateral pair two-segmented, basal section truncate, distal section digitiform; lateral, dorsolateral, and dorsal lobes low, rounded, each bearing 1 long seta distally; annulation of ventral lobes and distal section of ventrolateral lobes mostly obscured by very long setulae; center of disc glabrous. Two stigmatic tubes, each bearing a spiracular plate with a B-shaped spiracular slit and 4 transparent, irregularly branched float setulae, larger in proportion to size of disc than in second- or third-instar larvae. Anal proleg small, inconspicuous, hookless. (Based on 15 specimens: 4 from Praia do Leste, Parana1, and 11 from Parque Bitaru, Sao Vicente, Sao Paulo, both in Brazil). Second-instar Larva (Figure 126): Light brown. Integu- ment diaphanous. Length 2.8-5.3 mm (average = 3.9); greatest width 0.6-1.1 mm (average = 0.8). Cephalopharyngeal skeleton 0.49-0.52 mm long, with paired mandibles 0.09-0.10 mm in length, each with 3 mesally directed accessory teeth; very similar to others in Sepedonea. Ventral arch connected posterolaterally on each side to mandibles. Parastomal bars of epipharyngeal sclerite connected to fused hypopharyngeal- pharyngeal sclerite; the latter light to dark brown with small clear areas near dorsal margin of dorsal cornu; ventral window indistinct in light ventral cornu. Indentation index 31-35. Segment 1 as in first-instar larva. Seta arrangement of segments 2-4 as in first-instar larva. Segments 5-10 each with dorsolateral patch of setulae; segment 11 with single seta dorsolaterally. Segments 5-11 each with lateral tubercle group of 3 contiguous tubercles in a vertical row, the middle 1 slightly anterior to other 2; dorsal and ventral tubercles of lateral tubercle group of segment 11 with single seta each and middle tubercle with group of small, stout setae; main ventral tubercle group of 4 tubercles in a transverse row. Posterior spiracular disc with 5 pairs of lobes, very similar to that of S. telson; ventral pair subconical; ventrolateral pair two-segmented; lateral pair low, rounded; dorsolateral pair even less conspicu- ous; dorsal pair somewhat larger, broadly rounded. Two stigmatic tubes arising from center of disc, each bearing a spiracular plate (Figure 126) with a stigmatic scar, 3 rather small spiracular slits, and 4 large, prominent, irregularly branched, semitransparent float setulae each with glandular pore at base. Anal proleg scarcely larger than a main ventral tubercle when viewed laterally; without hooks. (Based on 12 specimens: Praia do Leste, Parana\ Brazil). Third-instar Larva (Figures 110, 113, 116, 135, 138): Length 8.2-9.3 mm (average = 8.7); greatest width 1.8-2.3 mm (average = 2.0). Light to dark brown, with dark brown middorsal stripe and dorsolateral V-shaped marks prominent anteriorly; laterally, a less prominent, broken, irregular lateral stripe; quite similar to S. trichotypa. Integu- ment opaque. Cephalopharyngeal skeleton length 0.74-0.82 mm, with paired mandibles (Figure 110), each with 4-6 large, sharply pointed accessory teeth, directed mesally; hook strongly decurved; entire sclerite 0.08-0.16 mm long; ventral arch (Figure 113) with 20-25 denticles directed anteriorly; connected posterolaterally with both mouth-hooks; hypophar- yngeal and labial sclerites (Figure 116) uniquely shaped among Sepedonea; former free from tentoropharyngeal sclerite; epipharyngeal sclerite very similar to that of S. telson and 5. trichotypa, with parastomal bars fused to tentoropharyngeal sclerite; latter light to dark brown, lacking small clear areas in dorsal cornu and with ventral window very indistinct. Indentation index 31-33. Segment 1 as in first-instar larva. Seta arrangement of segments 2-4 as in first-instar larva. Segment 2 with 2 anterior spiracles (Figure 138), each 0.13-0.14 mm long and bearing 6 or 7 papillae distally. Segments 5-10 each with dorsolateral patch of setulae with short setulae; segment 11 with single dorsolateral seta; lateral tubercle group on segment 11 with dorsal and ventral ones bearing single seta each; other lateral tubercles setaless; segments 5-11 each with main ventral tubercle group of 4 tubercles in a transverse row, followed posteriorly by 2 rows of much smaller intrasegmental tubercles arranged in transverse rows. Segment 8 with posterior spiracular disc (Figure 135) of 5 lobes: ventral pair quite wide at base, tapered sharply distally; ventrolateral pair basically similar to those of other species of Sepedonea; lateral pair rather acute; dorsolateral pair very low and inconspicuous; dorsal pair each with tuft of setulae distally, lobes broadly rounded. Entire disc moderately setulose, with glabrous center, with 2 separate stigmatic tubes, each scalloped basally, and each with a stigmatic scar, 3 spiracular slits, and 4 irregularly branched float setulae on a spiracular plate much lighter in color than 5. incipiens and S. telson. Anal proleg larger than in other species of Sepedonea but still relatively inconspicuous and hookless. (Based on 8 specimens: Praia do Leste, Parana^ Brazil). Puparium (Figure 97): Light brown with dark brown stripes along middorsal line and dorsolaterally on anterior segments, strongest on segments 6 and 7. Integument opaque. Length 5.2-5.7 mm (average = 5.4); greatest width 2.2-2.3 mm (average = 2.2). Barrel-shaped, with ends of cephalic caps projecting anteriorly slightly dorsal to the longitudinal body axis and slightly upturned. Anterior spiracles protruding from NUMBER 506 19 anterolateral corners of dorsal cephalic cap. Lateral tubercle group of segments 5-11 visible as strongly contrasting dark areas; main tubercles of ventral tubercle group persisting as 4 less noticeable light spots in a transverse row. Dorsolateral patches of setulae of segments 5-10 with setulae appearing appressed to puparium surface. Posterior end sharply upturned, stigmatic tubes forming an angle of 100-110 degrees with longitudinal body axis. Lobes of posterior spiracular disc shrunken. Anal plate invaginated; anal proleg lacking. Quite similar to S. telson and S. trichotypa, but much lighter, color pattern being dark-on-light rather than light-on-dark as in 5. telson and S. trichotypa. (Based on 5 specimens; Praia do Leste, Parana", Brazil). REMARKS.?This species, which occurs widely from north- ern to central South America, is especially well represented in eastern Brazil, but is practically lacking in the entire Amazon basin (Figure 141). Most specimens from Brazil differ from the specimens originating farther in the north, including the holotype, in having the midfemur distinctly spinose, in usually having dark lateral preapical marks on the hindfemur, in the darker and clouded wing, and in the posterior surstyli, which lack a median lobe. However, the overall similarity between the two groups of specimens, the male and female terminalia in particular, suggests that these differences merely represent intraspecific variation. Sepedonea incipiens, new species FIGURES 46-51,94, 100.104, 108-109,117,123,129. 134.139 ADULT.?Head: Lateral facial spot moderately large. Thorax: Mesonotum usually grayish black; postpronotum brownish yellow; setulae near posterior spiracle very strong and dense. Legs: Midfemur posteroventrally with 3-5 spines, not extended beyond half distance to base; hindcoxa posteriorly with short setulae, restricted to mesal area; hindfemur with dark posteroventral and anteroventral preapical marks and with dark or light dorsal preapical marks. Wing: Yellowish, with anteroapical margin and crossveins r-m and dm-cu more or less distinctly clouded; length 4-5 mm. Abdomen: Male terminalia: Posterior margin of sternum 4 straight, with median projection bearing a pair of tubercles (Figure 46); anterior plate of sternum 5 with triangular, anterolaterally directed flange (Figure 46); distiphallus (Figure 47) slightly curved, with short, slightly curved posteroventral spur; anterior surstylus indistinct; posterior surstyli with median lobe low and rounded (Figure 48) and with lateral lobe moderately curved anterad (Figure 49). Female synster- num: In ventral view posterior margin with 3 subequal tubercles (Figure 50); in lateral view ventral surface straight, posterior margin with apex and shoulders short and rounded (Figure 51). TYPE SPECIMENS.?Holotype d": ARGENTINA. BUENOS AIRES: 28 km SW Buenos Aires, 15-16 April 1967, J. Abercrombie, USNM. [Label: J. Abercrombie field trip #6729]. Allotype: BRAZIL. GUANABARA: InsL Oswaldo Cruz, 6 June 1967, Berg and Souza-Lopes. [Label: J. Abercrombie Biol. Note #67118]. Paratypesr. ARGENTINA. BUENOS AIRES: 28 km SW Buenos Aires, 15-16 April 1967, J. Abercrombie, Id", USNM. BRAZIL. GUANABARA: InsL Oswaldo Cruz, 6 June 1967, Berg and Souza-Lopes, Id", 19 (field collected), 14d", 149 (laboratory reared), USNM; 6 April 1967, Berg and Souza- Lopes, 29, USNM; 7 April 1967, Berg and Souza-Lopes, lcT, USNM. SAo PAULO: S3o Paulo, InsL de Botanica, July 1964, Berg and Papavero, Id", MZUSP. 15 July 1964, N. Papavero, lcf, MZUSP. September 1964, N. Papavero, lcf, MZUSP; Sao Paulo, InsL Bot. Serv. Agric, 11-13 July 1964, Berg and Papavero, \