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Psilota aegeae Vujić, Ståhls et Smit sp. n. ZooBank link: urn:lsid:zoobank.org:act: C15AA253-D7AB-4BE7-BAAE-F7617409E9B6 Figs 9, 10, 11A, C Type material. HOLOTYPE: Greece, ♂, pinned, in FSUNS. Original label: “Agiassos, 21.iv.2007, Sanatorio Site, leg. Pérez-Bañón C. & Vujić A.”. PARATYPES: Greece: 1♂ + 1♀, same locality as holotype, Lesvos, Agiassos, Sanatorio Site, 39°06’00.26”N 26°21’25.45”E, 21.iv.2007, leg. C. Pérez-Bañón, A. Vujić (FSUNS); 1♂ + 2♀, Lesvos, Agiassos, 39°06’00.26”N 26°21’25.45”E, 13.iv.2013, leg. A. Vujić (FSUNS); 1♂, Lesvos, Agiassos, 39°06’00.26”N 26°21’25.45”E, 8.v.2007, leg. G. Ståhls (MZH; collection code http://id.luomus.fi/GJ.2051 ♀, Lesvos, Agiassos, 39°06’00.26”N 26°21’25.45”E, 9.v.2007, leg. G. Ståhls (MZH; DNA voucher, collection code http://id.luomus. fi/GJ.2501 and http://id.luomus.fi/GJ.2052 1♂, Lesvos, Megali Limni, 28.iv.2008, 39°05’49”N 26°19’55”E, leg. G. Ståhls (MZH; http://id.luomus.fi/GJ.2053 1♂, Lesvos, Megali Limni, 39°05’49”N 26°19’55”E, 9.v.2009, leg. G. Ståhls (MZH; http://id.luomus.fi/GJ.2054 1♂, Lesvos, Megali Limni, 39°05’49”N 26°19’55”E, 9.v.2009, leg. G. Ståhls (MZH; http://id.luomus.fi/GJ.2055 1♂, Lesvos, Megali Limni, 1.v.2008, 39°05’49”N 26°19’55”E, leg. G. Ståhls (MZH; DNA voucher http://id.luomus.fi/GJ.2655 1♂, Lesvos, 30.iv.2008, leg. S. Radenković (FSUNS). Diagnosis. Psilota aegeae sp. n. belongs to the P. atra species group, with metafemur in male not swollen (Fig. 9E) and epandrium in male genitalia longer than broad, outer surstyle lobe shorter than inner surstyle lobe (Fig. 11 A–B). Antenna in female implanted in the upper half of the face (Fig. 10A), basoflagellomere about 2 times longer than broad, with clearly brown elongated macula basoventrally (Fig. 10A); abdomen covered with predominantly whitish pile, while the scutellum covered with whitish pile intermixed with the black ones (Fig. 10 C–D). Based on male genitalia characters, Psilota aegeae sp. n. is the most similar to Psilota atra, but differs as follows: in P. aegeae sp. n. epandrium 1.2 times longer than broad, while in P. atra almost 2 times (Fig. 11 A–B), furthermore there are differences in morphological characters of other body parts as well: basoflagellomere in P. aegeae sp. n. longer (about 2 times longer than broad) than in P. atra (1.5 times); cell R on wing in P. aegeae sp. n. is hyaline compared to P. atra where this cell is distinctly yellow; in P. aegeae sp. n. pile on tergum 2 mostly pale-yellow, and much longer on lateral margins of all terga with regard to P. atra (pile on all terga mostly black and much shorter on lateral margin); tergum 2 with less developed transverse grooves in P. aegeae sp. n. which are distinctively developed in P. atra. The female of P. aegeae sp. n. can be separated from all other Psilota species by the combination of the following characters: antennae implanted in the upper half of the face; anepisternum covered with white pile and metafemur not swollen. From the related P. atra, it differs by the color of the pilosity: pale-yellow on frons in P. aegeae sp. n. (in some specimens the pile on frons are pale-yellow in anterior part and black in posterior part), while black in P. atra; mostly pale-yellow on scutum in P. aegeae sp. n., while black in P. atra; white on metathoracic pleuron in P. aegeae sp. n., while mixed black and pale-yellow in P. atra. Molecular data: The clustering of the mtDNA COI barcode sequences of the included Psilota species and specimens is shown in Fig. 12. The uncorrected sequence divergences between Psilota aegeae sp. n. and included Psilota spp. ranged between 3.3–4.4%. Description. MALE. Head (Fig. 9 A–B). Antennae elongated, basoflagellomere about 2 times longer than broad, rounded at the tip, mostly dark-brown, except small light-brown basoventral inner area; face and frons black, mostly shiny, with scarce, indistinct dark-brown/gray microtrichia, covered with black pile, as well as vertex; face with a bare vitta, from the base of antenna to upper margin of mouth; genae and ventral part of mouth edge without pile; eyes densely covered with long yellowish pile; eye contiguity about as 2 times longer than ocellar triangle; ocellar triangle isosceles, covered with very long black pile; occiput very narrow, covered with short black pile. Thorax (Fig. 9C, E). Entirely black, with fine punctuation, covered with black pile somewhat wavy at the top; pile on scutum and scutellum clearly of two different lengths (Fig. 9C); scutellum with a row of long pile-like bristles at the posterior margin; subscutellar fringe composed of short black pile; metathoracic pleuron mostly shiny, predominantly covered with black pile (except whitish ones at grey microtrichose anterior end of anterior anepisternum), the longer pile somewhat wavy at the top; wings hyaline, veins clearly yellow at the base; pterostigma yellow; wing covered with microtrichia, except basal half of the cell R almost bare; calypteres yellowish, with some brown pile at the posterodistal margin of fringe, the rest yellow; halteres yellow-brown; legs entirely black, except apex of femora and base of the tibiae slightly lighter; all legs covered with mixed pale and black pile; metafemur thin, about 4.7 times longer than wide; the apical half of the metafemur ventrally with a shallow groove, formed by two rows of mixed short and long black spines (Fig. 9E); the ventral surface of the tarsi with the short yellow pile. Abdomen (Fig. 9D). Entirely black, predominantly shining; tergum 2 mostly covered with yellowish pile, only the posterior half with some black pile; terga 3 and 4 usually almost entirely covered with black pile, except some yellowish pile on lateral margins of both terga and anterior end of tergum 4 (area with yellowish pile on tergum 4 can be larger); all pile on the abdomen erect; sterna black, shiny, covered with long and erect whitish pile wavy at the top. Male genitalia (Fig. 11A, C). Epandrium compactly built, rectangular, longer than broad; inner surstyle lobe elongated, broad apically; outer surstyle lobe narrow and curved, shorter than inner surstyle lobe; cercus very elongated, rectangular (Fig. 11A); height of narrowest part of epandrium in dorsal view small (Fig. 11C). Superior lobe fused with hypandrium, narrow with tapering tip; aedeagus with a serrate ventral margin and pointed apex (Fig. 11A). FEMALE (Fig. 10). Similar to the male except for the following characters: face, frons and larger part of occiput covered with pale-yellow/whitish pile (Fig. 10 A–B); basoflagellomere with a clear elongated brown to reddish macula basoventrally, developed on both sides of basoflagellomere, inner and outer (Fig. 10A); pile on the eyes much shorter than in males; pile on scutum all pale-yellow, except at the wing basis, postalar callus and on posterior margin of scutellum mixed white and black pile (Fig. 10C); the metathoracic pleuron entirely covered with pale-yellow pile; legs and abdomen predominantly covered with whitish pile (Fig. 10D, E). Etymology. The specific epithet is the genitive case of the Greek name Aegea (to which the Aegean Sea owes its name) and refers to its origin, an island in the Aegean Sea. Distribution. Only known from Lesvos Island (Greece) (Fig. 6). Probably local endemic. Biology. Adults were collected in open areas within Mediterranean pine forest Pinus brutia Tenore (255–355m above sea level). They were visiting flowers of Smirnium perfoliatum L. Flight period is from April to beginning of May., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on pages 114-119, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960

Cheilosia candida Vujić et Radenković sp. n. ZooBank link: urn:lsid:zoobank.org:act: 316154DF-6472-4362-9269-F484233B277B Figs 1A, 1 C–D, 3, 4 Type material. HOLOTYPE. Greece, ♂, pinned, in NBCN. Original label: “Griekenland, Prov. Pindos gebergte, Plaats Kalliroi, 19.v.1988, leg. G. den Hollander ” (det. as Cheilosia longula by G. den Hollander, 1984). Diagnosis. Species related to Cheilosia paralobi, but differs in the following characters: face partly microtrichose on genae, along paraface and between antennae and facial tubercle in C. candida sp. n. (Fig. 1A), while in C. paralobi entire face microtrichose, except facial tubercle (Fig. 1B); basoflagellomere of male partly reddish in C. candida sp. n. (Fig. 1D), while in C. paralobi almost completely black (Fig. 1E) (in some specimens inner side basally with paler macula); frons covered with intermixed black and yellow pile in C. candida sp. n., while in C. paralobi only with yellow pile. Male genitalia: surstylus shorter in C. candida sp. n., about three times longer than broad basally (Fig. 4B), while in C. paralobi very narrow apically and elongated, more than five times longer than broad basally (Fig. 5E). Description. MALE. Head (Figs 1A, 1 C–D). Antennae dark, brown-reddish; basoflagellomere dark-brown with reddish macula basoventrally, ellipsoidal, about 1.3 times as long as broad, and 3.5 longer than pedicel; arista 1.5 times longer than antennae, covered with short pile (Fig. 1D); face wide, 1/2 of the width of head anteriorly, covered with long, pale-yellow pile along paraface in upper 1/3 (Fig. 1A); facial tubercle rounded; genae, face between antennae and facial tubercle, and along paraface covered with grey microtrichia; frons convex, shiny, covered with mixed yellow and black pile; vertex convex and shiny; ocellar triangle equilateral, covered with mixed yellow and black pile; eyes bare; eye contiguity shorter than length of frons (Fig. 1C); occiput narrow, with white-grey microtrichia, covered with yellow pile, with additional black pile dorsally along eye margin. Thorax (Fig. 3). Scutum black, with dark-golden color, laterally slightly microtrichose, covered with long yellow pile, except wing bases covered with black pile and few black ones in notopleural area; central disk shiny, with fine punctuation; scutellum covered with yellow pile (Fig. 3A, C); metathoracic pleuron slightly microtrichose, covered with pale-yellow pile and wavy at apical end; wing membrane brownish medially, with brown veins, completely microtrichose; vein M1 meeting vein R4+5 at acute angle; halters light to dark-brown; calypteres yellowish-gray; all legs dark-brown/black, except light-brown apex of femora, basal 1/3 and apical end of tibiae, and ventral surface of tarsi; legs grey microtrichose and with mixed yellow and black pile, although yellow ones dominate (Fig. 3B). Abdomen (Fig. 3C). Terga dull, covered with mostly yellow pile, except few black ones on pregenital segment; sterna grey microtrichose, covered with yellow pile, except few black ones on sternum 4. Male genitalia (Fig. 4). Surstylus about three times longer than broad basally, with central bulge from lateral view (Fig. 4B); dorsal lobus of gonostylus sickle-shaped, with spine-like dorsal prolongation (marked with arrow on Fig. 4A). FEMALE. Unknown. Etymology. The new species is named according to the Latin adjective ‘ candida ’ meaning bright (not dark), indicating yellow color of body pile and reddish antennae. Distribution. Known only from Greece, from a locality on Pindos Mountains (Fig. 6)., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on pages 107-108, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960

Subgenus Paragus Latreille Diagnosis. Eyes with two dorso-ventral vittae of whitish pile, as on Figs 7 A–B. Scutellum without conspicuous teeth on posterior margin. Only terga 1 and 2 completely fused., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on page 111, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960

Genus Psilota Meigen Diagnosis of the P. atra species group Within the Palaearctic, two well defined species groups can be separated: the P. atra species group, recognized in both sexes by the only slightly microtrichose and therefore clearly shiny face below the antenna, and by bluishblack and shiny thorax and abdomen, while the P. innupta species group is recognized in both sexes by the heavily microtrichose and therefore clearly dull face below the antenna, by bronze-brown color of thorax and abdomen, and by the medially dull terga in males. The P. atra group consists of the following species: Psilota anthracina, Psilota atra (Fallén), Psilota exilistyla Smit et Vujić, Psilota kroshka Mutin, and Psilota nana Smit et Vujić., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on page 114, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960

Paragus thracusi Radenković, Likov et Vujić sp. n. ZooBank link: urn:lsid:zoobank.org:act: 2DCF31FB-8425-41D2-8725-22FF35B4EE4A Figs 7, 8 Type material. HOLOTYPE: Greece, ♂, pinned, in NBCN. Original label: “Hellas, Thraki, Rodopi, Porto Lagos SE, 1.v.2000, leg. W. van Steenis, E. S. Bakker ”. Diagnosis. Paragus thracusi sp. n. is by habitus most similar to P. oltenicus Stćnescu, but easily distinguishable by the following characters: mesonotum of P. thracusi sp. n. covered with longer erect whitish-yellow pile in contrast to short yellow adpressed pile in P. oltenicus; microtrichose maculae at the end of transverse suture present in P. thracusi sp. n., but lack in P. oltenicus; punctuation of mesonotum scarce in P. thracusi sp. n., but dense in P. oltenicus; femora dark-brown/black basally in P. thracusi sp. n., while orange-yellow in P. oltenicus; tergum 1 black in P. thracusi sp. n., and medially reddish in P.oltenicus; microtrichose fasciae on terga 2–4 distinct in P. thracusi sp. n., but absent or scarce in P. oltenicus; microtrichia on terga completely whitish-yellow, while in P. oltenicus partly black; black transversal maculae on terga 2–4 lack in P. thracusi sp. n., but usually present in P. oltenicus. Based on the male genitalia P. thracusi sp. n. is immediately recognizable by the characters of hypandrium: pointed, hood shaped paramere and cliff shaped lingula. Description. MALE. Head (Figs 7 A–B). Face cream-yellow, with a faint brown macula on facial tubercle; ventral margin of face dark-brown/black, except the light-yellow frontal part; face and frons covered with only whitish pile; antennae long, basoflagellomere elongated and pointed, about 1.5 times as long as scape and pedicel together; arista 0.7 times the length of basoflagellomere; antennae predominantly brown, except orange basal 2/3 of basoflagellomere ventrally and pedicel ventrally; arista light-brown basally and dark-brown apically; eye contiguity 0.75 times the length of ocellar triangle; eyes with two clearly visible vittae of white pile, typical of the nominal subgenus; vertical triangle covered with grayish microtrichia in front of frontal ocellus, covered with light-yellow pile; occiput whitish microtrichose and with light-yellow pile, and with silver pile in lower parts, turning yellowish in upper parts. Thorax. Scutum black with paler anterior and posterior end of postalar callus, shiny, except two longitudinal submedian whitish microtrichose vittae ending at the level of transverse suture, and with two whitish microtrichose maculae at the end of transverse suture; mesonotum with scarce punctuation, and covered with longer erect whitish-yellow pile; metathoracic pleuron whitish microtrichose and pilose, except for the anteroventral part of posterior anepisternum and anepimeron, which are shining black; scutellum black in anterior half, orange-yellow in posterior half, without conspicuous teeth on posterior margin; wings hyaline, covered with microtrichia, except for basal cells, which are nearly entirely bare; halters light-brown, with a yellow knob; legs two colored, covered with whitish-yellow pile; femora dark-brown, black at base (pro- and mesofemora in basal 1/3, and metafemur in basal 2/3, except for tiny area near to the base yellow), and light-yellow apically; tibiae light-yellow in basal half, and dark-yellow in apical half, metatibia with brown submedian ring; tarsi orange-yellow (borders between tarsomeres black), basotarsomere of metatarsus little darker dorsally; claws yellow in basal half and black in the apical half. Abdomen (Fig. 7C). Abdomen predominantly red-brown, covered with whitish-yellow pile and with three pairs of whitish-yellow microtrichose fasciae on terga 2–4; tergum 1 brown-black, more or less fused with tergum 2 that is dark-brown anteriorly to the microtrichose fasciae and laterally, while red-brown posteriorly; terga 3 and 4 predominantly red-brown, except dark-brown longitudinal areas near the lateral margins and on medial parts; tergum 5 red-brown; sterna translucent orange-brown (sternum 1 darker), covered with erect whitish pile; pregenital segment white microtrichose and pilose. Male genitalia (Fig. 8). Hypandrium in lateral view wide (Fig. 8A) with clearly developed lingula cliff shaped (Fig. 8A: l); lateral lobe of aedeagus (= inferior claspers) well developed, almost quadrilateral in lateral view (Fig. 8A: la); paramere (= superior claspers) pointed, hood shaped (Fig. 8 A–B: p); ejaculatory apodeme pendulum like (Fig. 8C); epandrium in lateral view narrow, with narrow, not prominent cercus (Fig. 8E); surstylus (= styles) a little shorter than theca of epandrium (Fig. 8E: sst), curved and tapering in dorsal view (Fig. 8D: sst). FEMALE. Unknown. Etymology. The species is named after the type locality, in the North-Eastern part of Greece that belongs to the Thrace (Greek word: Θράκη), geographical and historical area in southeast Europe. Distribution. Only known from Rhodope Mountains in Greece (Fig. 6)., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on pages 111-114, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960

Subgenus Convocheila Barkalov Diagnosis. Face along paraface in upper 1/3 covered with long whitish or black pile (Figs 1 A–B), long as pile around ocellar triangle; eyes usually pilose (in some species without pile: C. candida Vujić et Radenković sp. n., C. paralobi and C. laticornis); face wide, with facial tubercle nose-like; frons convex; antennal pits separated; basoflagellomere rounded or ellipsoidal, dark-brown/black with reddish markings (except in some specimens of C. paralobi and C. cumanica with entirely black basoflagellomere); pile on scutum and scutellum mixed pale-yellow and black (black pile at notopleural area, at wing bases and on posterior margin of scutellum); pile on metathoracic pleuron long, whitish, and wavy apically; all legs black, except light-brown apical parts of femora, basal 1/3 and apical end of tibiae and ventral surface of tarsi; pile on legs mixed pale-yellow and black; abdomen microtrichose, predominantly covered with long pale pile (except the posterior half of lateral margins of terga 3 and 4 with black pile in some males of C. laticornis and C. hypena (Becker)). Male genitalia: ventral lobus of gonostylus sickle-shaped (Figs 2, 4A). In the Mediterranean area, this subgenus comprises besides the four previously mentioned species in Barkalov (2002), C. paralobi, C. cumanica, C. sulcifrons and C. laticornis, also two additional species: C. hypena and C. candida sp. n., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on pages 105-107, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960, {"references":["Barkalov, A. V. (2002) Subgeneric classification of the genus Cheilosia Meigen, 1822 (Diptera, Syrphidae). Entomological Review, 82, 518 - 531."]}

Psilota anthracina Taxonomic notes. Speight (2020) recognized the problems associated with recognition of Psilota anthracina based on male genitalia features, because of the differences in the figures given by Smit & Zeegers (2005) and Smit & Vujić (2008). In order to resolve this confusion, we studied available European material, including specimens from the Balkan Peninsula, and particularly checked original specimens whose drawings were presented in the two above mentioned publications referred to. After detailed analysis, we concluded that all specimens belong to Psilota anthracina, and that the main reason for the differences in these drawings was a different angle of observation and additionally a slightly aberrant epandrium in the Montenegro specimen figured by Smit & Vujić (2008). Examined material (published). Smit and Vujić (2008): Austria: 1♀, Carinthia, Eisenkappel, 24.iv.1955, (NBCN); Belgium: 1♀, Hautes Fagnes, 7.vi.2004, leg. B.Wakkie (coll. BW); 1♀, Malmedi, Mont Rigi, 29.v.2005, leg. W. Renema (coll. WR); 1♀, Ottenburg, Rodebos, 25.v.2005, leg. F. v. d. Meutter (coll. FvdM); Croatia: 1♀, Zagreb, Sljeme, 1000m, 27.v.1969; 2f, 30.v.1969, leg. Lambeck (NBCN); England: 1♂, Brockenhurst, 7.v.1945, leg. Wainwright (coll. TN); 1♀, Monach. (= Outer Hebrides?) (NBCN); France: 3♀, Col de Jeau, Mosset, 9–15.vi.2003, leg. M. Willemse (coll. MW); 1♂ + 4♀, Ile de France, Foret d Evry, 25.v.1997, leg. A. v. Eck (coll. AvE); Germany: 1♀, Sachsen-Anhalt, 3km NO Aken, 15.v.2002, leg. F. Dziock (coll. FD); Greece: 1♀, Ipiros, Peristeri Mts., 12–1700m, 24–8.v.1994, leg. S. Andersen (coll. CC); Italy: 1♀, Raibl (nowadays Cave del Predil), vii.1897, leg. Bergenstamm (NMW); Montenegro: 1♂ + 1♀, Boka Kotorska, Morinj, 5.v.1991; 4♂ + 1♀, 4.v.1994; 1m, 7.v.1994; 1m +1f, 1–3.v.1997; 1♀, 12.v.1997; 3m +1f, 7.v.1998; 1f, Durmitor, Sušičko jezero-Zlatica, 19.v.2000 (FSUNS); Netherlands: 1♀, Boxtel, 16.v.2005, leg. L. v. d. Leij (coll. LL); 1♂ + 1♀, Ede, Planken Wambuis, Nieuw Reemst, 12.v.2006, leg. J. T. Smit (coll. JTS); 1♀, idem, leg. John T. Smit (coll. JTS); Spain: 1♀, Pyrenees, Valle de Ordesa, 1500m, 23.v.2004, leg. F. Dziock (coll. FD). Examined material (new). The United Kingdom: England: 1♂, Brockenhurst, 7.v.1945, leg. Wainwright (coll. TN); Croatia: 1♀, Zagreb, Sljeme, 1000m, 27.v.1969, leg. Lambeck (NBCN); Czech Republic: 1♀, Podyji NP, Nad Sobesem forest-steppe, MT, 340m, 3.v.–2.vi.2003, leg. Bartak, Kubik (coll. MB); 1♂, Podyji NP, Pod Ledovymi slujemi, wetland near river, 26.iii–13.v.2002, leg. Bartak, Kubik (coll. MB); Montenegro: 1♂, Boka Kotorska, Morinj, 4.v.1994; 1m +1f, 7.v.1994; 1f, 1.–3.v.1997; 1♀, Durmitor, Sušičko jezero-Zlatica, 19.v.2000 (FSUNS)., Published as part of Radenković, Snežana, Likov, Laura, Ståhls, Gunilla, Rojo, Santos, Pérez-Bañón, Celeste, Smit, John, Petanidou, Theodora, Steenis, Wouter Van & Vujić, Ante, 2020, Three new hoverfly species from Greece (Diptera: Syrphidae), pp. 103-124 in Zootaxa 4830 (1) on page 119, DOI: 10.11646/zootaxa.4830.1.4, http://zenodo.org/record/4402960, {"references":["Speight, M. C. D. (2020) Species accounts of European Syrphidae, 2020. Syrph the Net, the database of European Syrphidae (Diptera). Vol. 104. Syrph the Net publications, Dublin, 314 pp.","Smit, J. T. & Zeegers, T. (2005) Overzicht van het Zweefvliegen genus Psilota in Nederland (Diptera: Syrphidae). Nederlandse Faunistische Mededelingen, 22, 113 - 120. [https: // www. researchgate. net / publication / 254911940 _ Overzicht _ van _ het _ zweefvliegengenus _ Psilota _ in _ Nederland _ Diptera _ Syrphidae]","Smit, J. T. & Vujic, A. (2008) The Palaearctic species of the genus Psilota Meigen (Diptera, Syrphidae) with the description of two new species. Studia dipterologica, 14 (2), 345 - 364."]}

Merodon aureus Fabricius Puparium description. Dimensions and shape. Length: 7.5 mm, width: 3.5 mm, height: 3 mm; sub-cylindrical; anterior end truncate, tapering posteriorly and flattened Ventrally; cream to light brown in colour; rough integument with larVal segmentation persisting as transVerse folds and wrinkles. Cephalopharyngeal skeleton. Robust mandibles with black sclerotised hooks and blunt apex, without accessory teeth, fused to the external mandibular lobes; the mouth hooks separated at the apex by about the same distance as their basal width; the dorsal cornu narrowed towards the apex, shark���s fin shaped, representing almost the whole length of the Ventral cornu; clipeal sclerite and tentorium highly sclerotised and apparently fused to the intermediate sclerites; Ventral cornu elongated and narrow in profile View, with the cibarium located at the base (Fig. 1A). Pupal spiracles. Sclerotised, blackish-brown in colour, cylindrical in shape, gradually tapering and pointed at the end; 0.6 mm long; separated by a distance of 4�� their length; apex with a granular surface; surface of the base smooth or slightly rough; each spiracle bearing numerous irregularly-spaced domed tubercles (Fig. 2A) except for the pointed apex; surface between tubercles smooth or slightly rough; tubercles composed of two layers; 4���7 radially-arranged spiracular openings on each tubercle (Fig. 2B). Prp. Reddish-brown and shiny, sub-elliptical in cross-section, short (Visible from dorsal View) and wider than long; with an annular grooVe at the base and another just before the spiracular plate; surface between the two grooVes ornamented with a network of small rounded protuberances (Fig. 3A); the outline of the spiracular plate irregular with indentations at the locations of the inter-spiracular setae; spiracular plate of prp with four pairs of slightly curVed and conVoluted spiracular openings around two central scars (Fig. 3B); the spiracular openings clearly separated from each other; four pairs of well deVeloped inter-spiracular setae emerging from the edge of the spiracular plate; each seta with at least four main branches, each of which possesses multiple sub-branches from the base to the tip of the seta (Fig. 3C). Examined material. Đerdap National Park, Serbia (44�� 32 ' 38"N, 22��01'08"E), leg. J. PreradoVić: 1 puparium in the ground near bulbs of Ornithogalum umbellatum, V.2013 (P 1 in FSUNS); 1 puparium on the ground close to O. umbellatum plants, Viii. 2015 (P 4 in FSUNS). Biology. Males and females of M. aureus were obserVed flying near and resting on plants of Ornithogalum sp. from the beginning of May to the beginning of June in 2011 -2013 and 2015-2016 (more than 50 obserVations in total). Up to four adults were noted to sometimes rest simultaneously on the same plant leaVes. Females of M. aureus were often obserVed settling for oViposition between the base of an Ornithogalum sp. plant and the surrounding soil, behaVing as if they were laying eggs on the ground, Very close to the plant. HoweVer, we neVer found eggs. The females made intermittent buzzing sounds while maintaining their positions for oVer fiVe minutes. Thereafter, they cleaned themselVes for a few minutes before flying away., Published as part of Preradovi��, Jelena, Andri��, Andrijana, Radenkovi��, Sne��ana, Zori��, Ljiljana ��a��i��, P��rez-Ba����n, Celeste, Campoy, Andr��s & Vuji��, Ante, 2018, Pupal stages of three species of the phytophagous genus Merodon Meigen (Diptera: Syrphidae), pp. 229-242 in Zootaxa 4420 (2) on pages 232-233, DOI: 10.11646/zootaxa.4420.2.5, http://zenodo.org/record/1250819

Merodon avidus (Rossi) Puparium description. Dimensions and shape. Length: 9.7 mm, width: 4.7 mm, height: 4.7 mm (n=2); subcylindrical; anterior end truncate, slightly tapering posteriorly and flattened Ventrally; dark brown in colour; rough integument with larVal segmentation persisting as transVerse folds and wrinkles. Cephalopharyngeal skeleton (n=3). Mandibles with black sclerotised hooks and blunt apex, without accessory teeth, fused to the external mandibular lobes; the mouth hooks separated at the apex by a distance wider than or equal as their basal width; the dorsal cornu tapered towards the apex, shark���s fin shaped, representing two thirds of the whole length of the Ventral cornu; clipeal sclerite and tentorium slightly sclerotised and separated from the highly sclerotised intermediate sclerites; Ventral cornu elongated and narrow in profile View, with the cibarium at the base (Fig. 1B). Pupal spiracles. Sclerotised, blackish-brown in colour, stout, cylindrical in shape, gradually tapering and pointed at the end; length ��� 0.4 mm (length: width ratio of spiracle 1.5:1); separated by a distance of 4�� their length; surface of the spiracle with a polygonal pattern (Fig. 2C); dorsal surface facing toward the lateral margins of the puparium and with the distal one-fifth slightly curVed backwards; about 60% of the dorsal and lateral surface coVered with irregularly-spaced, oVal-shaped tubercles; tubercle composed of two layers; 5���6 radially-arranged spiracular openings on each tubercle. Prp. Blackish-brown, sub-elliptical in cross-section, extremely short (not Visible from dorsal View); spiracular plate of prp with four pairs of sinuous spiracular openings around two central scars (Fig. 3D); four pairs of interspiracular setae emerging from the edge of the spiracular plate. Examined material. Đerdap National Park, Serbia (44�� 32 ' 38"N, 22��01'08"E), leg. J. PreradoVić: 2 puparia in the ground near bulbs of Ornithogalum umbellatum, Viii.2015 and xi.2015 (P2 and P 5 in FSUNS); 1 puparium on the ground close to O. umbellatum plants, Viii.2015 (P 3 in FSUNS); 1 puparium in the ground near bulbs of O. umbellatum, 29.V.2016, ♂ emerged on 8.Vi. 2016 (P 6 in FSUNS)., Published as part of Preradovi��, Jelena, Andri��, Andrijana, Radenkovi��, Sne��ana, Zori��, Ljiljana ��a��i��, P��rez-Ba����n, Celeste, Campoy, Andr��s & Vuji��, Ante, 2018, Pupal stages of three species of the phytophagous genus Merodon Meigen (Diptera: Syrphidae), pp. 229-242 in Zootaxa 4420 (2) on pages 233-234, DOI: 10.11646/zootaxa.4420.2.5, http://zenodo.org/record/1250819

Identification of Merodon puparia BLAST search results reVealed 99% COI sequence identity (corresponding to a difference of one nucleotide) between puparium P2 (640 base pairs, bp) and a sequence of M. avidus deposited in GenBank under accession number FM206517. The query coVerage was 97%. HoweVer, BLAST search did not help in resolVing the species of the P1 puparium. We aligned and compared the P1 pupal sequence (634bp) with sequences resolVed from adult Merodon specimens deposited in the FSUNS sequence database and concluded that the puparium P1 belonged to M. aureus since the P1 sequence was identical to the sequences of nine adult specimens of M. aureus collected in Đerdap National Park (Table 1) and differed by only one mutational step from a tenth adult specimen of M. aureus from the same locality. Genetic identification of the two other hoVerfly puparia found on the ground close to O. umbellatum plants in August 2015, namely P 3 and P 4, was unsuccessful because the sequencing failed to obtain DNA from a hoVerfly species. Instead, we obtained sequences of two hymenopterans. Thus, sequencing analysis reVealed parasitoidism by wasps of the Hymenopteran families Diapriidae and Ichneumonidae. NeVertheless, based on morphological characters, puparium P3 (parasitoidized by Diapriidae) was identified as M. avidus and puparium P4 (parasitoidized by Ichneumonidae) as M. aureus. As a summary, our genetic results and morphological comparisons reVealed that the puparia found in the ground near bulbs of O. umbellatum and on the ground close to these plants in the Đerdap National Park belong to M. aureus (P1 and P4) and to M. avidus (P2, P3 and P5) (Table 1).

Key to the members of the Brachyopa quadrimaculosa species group in the Eastern Mediterranean (following Doczkal & Dziock 2004) 1. Scutellum entirely covered in microtrichia................................. Brachyopa bimaculosa Doczkal & Dziock - Scutellum largely bare of microtrichia, at least on posterior 1 / 2 ��� 1 / 3.............................................. 2 2. Proepimeron with a few pile anteriorly. Scutum with rounded bare spots at the transverse suture (Figs 3 C & D)........... 3 - Proepimeron bare. Scutum with microtrichose transverse suture................................................. 5 3. Lateral sides of scutum uniformly covered with light grey microtrichia....... Brachyopa vernalis van Steenis & van Steenis - Lateral sides of scutum with stripes of dark grey microtrichia... (as in Fig. 3 C)..................................... 4 4. Postalar callus with a very small bare spot (Fig. 3 C); male terminalia illustrated in Kaplan & Thompson (1981: 209, Figs 16��� 18)............................................... Brachyopa quadrimaculosa Thompson in Kaplan & Thompson - Postalar callus with large bare spot (see van Steenis & van Steenis 2004: Fig. 4).................................................................................................. Brachyopa cruriscutum van Steenis & van Steenis 5. Sensory pit present; scutum with bare spots and stripes (Figs 3 A & B); tergites with pale pile; male terminalia in Figs 4���5.................................................................................. Brachyopa minima sp. nov. - Sensory pit absent; scutum completely microtrichose except for the postalar callus; second tergite with black pile posteromedi- ally in female (can be absent in male); (male terminalia illustrated in Vujić (1991: 147, Fig. 4 B)....................................................................................................... Brachyopa insensilis Collin, Published as part of P��rez-Ba����n, Celeste, Radenkovi��, Snezana, Vuji��, Ante, St��hls, Gunilla, Rojo, Santos, Grkovi��, Ana & Petanidou, Theodora, 2016, Brachyopa minima (Diptera: Syrphidae), a new species from Greece with notes on the biodiversity and conservation of the genus Brachyopa Meigen in the Northern Aegean Islands, pp. 217-234 in Zootaxa 4072 (2) on page 223, DOI: 10.11646/zootaxa.4072.2.5, http://zenodo.org/record/270665, {"references":["Doczkal, D. & Dziock, F. (2004) Two new species of Brachyopa Meigen from Germany, with notes on B. grunewaldensis Kassebeer (Diptera, Syrphidae). Volucella, 7, 35 - 59.","Kaplan, M. & Thompson, F. C. (1981) New Syrphidae from Israel (Diptera). Proceedings of the Entomological Society of Washington, 83, 198 - 212."]}

Brachyopa minima Vujić & P��rez-Ba����n sp. nov. Diagnosis. Small species (6���7 mm). Scutum (Figs 3 A, B) covered with grey microtrichia except: pair of small bare spots posteromedian to the postpronotum; a pair of large bare spots anterior to transverse suture and median to notopleural sulcus; spots at the median ends of transverse sulcus covered by brown microtrichia; postalar callus with a bare spot; two stripes bare of microtrichia on posterior part of scutum located laterally. Male terminalia on Figs 4���5; surstylus with well-developed ventral lobe (Figs 4 A, 5 A); dorsal and ventral margins of surstylus straight (Figs 4 A, 5 A); hypandrium wide with complex apical appendages (Figs 5 C, D). Type material. HOLOTYPE: Greece, Lesvos, Karini, 13.04.2011, 1 male (G0322), leg. Vujić (in FSUNS). PARATYPES: Greece, Lesvos, Karini, 13.04.2011, 4 males (G018���G020, G023), 1 female (G0324), 13.04.2013, 1 female (G 2805), leg. Vujić (in FSUNS); 30.04.2008, 1 male (G 3069), leg. Vujić (in FSUNS); Greece, Lesvos, Karini, 26.04.2007, 1 male (G 3068) 1 female (G 3067), leg. P��rez-Ba����n and Vujić (in FSUNS). Greece, Lesvos, Karini, 3.05. 2007, leg. P��rez-Ba����n, 1 female, DNA voucher specimen MZH _Y 544 (in MZH). Etymology. The specific epithet minima means "smallest" and refers to the size of the species, which is smaller than most other members of the genus. Description. MALE. Body length 6���7 mm, wing length 7 mm. Head (Figs 1 A, C; 2 A���C). Length to height ratio of head 0.97. Face moderately protruding (Fig. 1 C). Length to width ratio of clypeus 1.4. Length to width ratio of subcranial cavity 1.6; 1.7 times as wide as the shortest distance from the eye to the subcranial cavity. Orbital strip (Figs 2 A, B) shallow, in narrower part as wide as the maximum width of arista. Pile on all parts of the head capsule pale, except black pile on upper part of occiput near eye margin. Frons and vertex (Fig. 1 A) microtrichose, only narrow area above lunule and around ocelli bare. Length of eye contiguity about 0.4 times the length of frons, shorter than ocellar triangle. Distance between posterior ocellus and eye margin about equal to two ommatidium diameters. Hypostomal bridge grey-reddish, paler anteriorly, entirely microtrichose, except bare hypostomal sulcus. Face (Figs 2 A, B) microtrichose except for a fascia from the tentorial sulcus to the mouth. Length to height ratio of postpedicellus 1���1.15, with small sensory pit (Fig. 2 C). Arista (Fig. 2 C) with very short pile (shorter than 1 / 3 the maximum width of arista), with orange base and darkened toward apex. Thorax. Black except for the postalar callus, which is partly reddish, and the metasternum and probasisternum which are more or less reddish. Scutum (Figs 3 A, B) entirely densely covered in grey microtrichia except for a pair of bare maculae posteromedian to the postpronotum; a pair of bare maculae anterior to transverse suture and median to notopleural sulcus (maculae at the median ends of transverse sulcus covered by brown microtrichia); postalar callus with a bare spot; two stripes bare of microtrichia on posterior part of scutum located laterally. Notopleural sulcus impressed. Thoracic bristles not well differentiated from the pile. Postpronotum pale haired, with few black pile mixed in. Scutum predominantly black haired except for its anterior margin and the notopleuron, which are pale haired, a few pale pile that are present at the supra-alar area, and the postalar callus that is pale haired. Scutellum rather trapezoidal, without a distinct depression, mainly orange, only slightly darkened anteriorly and anterolaterally, with two pairs of strong black marginal bristles; predominantly pale haired mixed with black pile medially; only anterior half covered with microtrichia (Fig. 2 D). Mediotergite microtrichose below subscutellum, except at posterior margin. Proepimeron and anterior flat part of mesanepisternum bare. Posterior surface of mid coxa without pile. Legs orange, 5 th tarsomere of all legs black, tarsomeres 1���4 of metatarsus darkened dorsally. Legs pale pilose except for the usual black bristles on hind femur ventrally, apex of mesotibia ventrally with strong black bristles, the ventral surface of mesotarsus with black bristles. Haltere yellow with light yellow capitulum. Wing without dark spots. Abdomen. Tergites and sternites orange, except first tergite and first sternite with dark grey area laterally, pile all pale. First tergite entirely microtrichose, second tergite extensively covered with microtrichia except for the posterior margin narrowly and large area at the posterolateral corners, which are bare. Sternites entirely densely microtrichose. Male terminalia on Figs 4���5. FEMALE. Differs from the male in the following characters: length to width ratio of frons 1.46���1.54. Frons (Fig. 1 B) entirely microtrichose except shiny anterior area above antennae, with small central microtrichose spot (this spot lacking in B. quadrimaculosa). A narrow pollinose stripe running from the dusted area of the frons downward along the eye margin, widening beside the lunule and merging with the wide pollinose cross band that covers the dorsal part of face (face less covered by microtrichia than in male, without bare spot below each antenna, contrary to B. quadrimaculosa) (Fig. 1 B). Length to height ratio of postpedicellus 1.14���1.15, a little larger than in male, with a minute sensory pit. Tibiae and tarsi darker than in male., Published as part of P��rez-Ba����n, Celeste, Radenkovi��, Snezana, Vuji��, Ante, St��hls, Gunilla, Rojo, Santos, Grkovi��, Ana & Petanidou, Theodora, 2016, Brachyopa minima (Diptera: Syrphidae), a new species from Greece with notes on the biodiversity and conservation of the genus Brachyopa Meigen in the Northern Aegean Islands, pp. 217-234 in Zootaxa 4072 (2) on pages 220-222, DOI: 10.11646/zootaxa.4072.2.5, http://zenodo.org/record/270665

An on-going study of the hoverfly fauna of the Northern Aegean Islands (Greece) has revealed the presence of four species of the genus Brachyopa Meigen. During the survey the following species were found: B. bicolor (Fallén), B. quadrimaculosa Thompson in Kaplan & Thompson, B. minima Vujić & Pérez-Bañón sp. nov. and an unidentified species very close to B. pilosa (Collin). Morphological characters and mitochondrial COI barcodes were used to link different life stages of B. minima, and to identify a larval specimen of B. bicolor. In this study adult and larval morphology and habitat preferences for B. minima are described. The description of larval morphology of B. bicolor and Brachyopa sp. aff. pilosa is amended too. An identification key to the adults of the B. quadrimaculosa group sensu Kassebeer (2002) in the Eastern Mediterranean (Greece, Israel and Turkey) is provided. The importance of specific microhabitats for the continued existence of these taxa is discussed. The study was funded by the European Union (European Social Fund – ESF) and Greek national funds through the Operational Program "Education and Lifelong Learning" of the National Strategic Reference Framework (NSRF)—Research Funding Program: THALES—Investing in knowledge society through the European Social Fund (Petanidou et al. 2013). Partial support came from the Ministry of Education, Science and Technological Development of the Republic of Serbia projects No. 173002 and the Provincial Secretariat for Science and Technological Development (project: Genetic diversity of agricultural ecosystems).

Eosalpingogaster umbra Mengual and Thompson, 2011 Comparing the morphology and ultrastructure of the posterior breathing tube (prp = posterior respiratory process) of the available puparia from type material showed unexpectedly that there were two different morphotypes. Three specimens were studied: two males, showing one puparial morphotype, and one female with a completely different morphotype. We consider that the true puparial morphotype of Eosalpingogaster umbra is the one associated to the studied male specimens, as one of the males is not totally emerged from the puparium. We also think that the puparium associated to the female individual of E. umbra does not belong to this species, and it may be the result of erroneous taxonomy or human error. Therefore, this puparium is here described as Eosalpingogaster sp. because its identity is unclear. See a more comprehensive discussion below about the improbability of immature sexual dimorphism in this species. Overall appearance. Length 5–6 mm, maximum width 2.5 mm and height 2.5 mm. Pear-shaped, in profile with dorsal and ventral margins of the posterior end forming an acute angle until the anterior bulge (fourth first abdominal segments); see Fig. 1 C, D. Outline serrated, sub-cylindrical in cross-section, anterior end truncate, tapered posteriorly and flattened ventrally. Integumental vestiture and segmental sensilla persisting except on the ventral surface. Dorsal surface covered with minute blunt-tipped papillae. Colour of empty puparium cream with bright brown spots on most basal papillae of segmental sensilla; a slightly dark brown spot in front of the tubercles of segmental pair of sensilla 1 on 3 rd abdominal segment. The serrated outline is due to great development of basal papillae of segmental sensilla as tubercles on the 2 nd, 3 rd and 4 th abdominal segment (Fig. 2 A). There are three transverse rows of paired tubercles (segmental pairs of sensilla 1, 2, 3 and 5–6). Tubercles of segmental pair of sensilla 1 are less developed on the 2 nd than on the 3 rd and 4 th abdominal segment. Tubercles of segmental sensilla 5–6 (born on the same papilla) are clearly smaller than the others tubercles on the three abdominal segments. On the last abdominal segments (5 th, 6 th and 7 th) the basal papillae of segmental sensilla are dried and collapsed not forming tubercles. Dorsal and dorsolateral segmental pairs of sensilla with apical setae not visible. The last three abdominal segment (5 th, 6 th and 7 th) and anal segment narrowed significantly towards the base of posterior breathing tube. Posterior breathing tube (prp): Short, broader than long. Dark brown in colour, lustrous. Spiracular plate little higher than wide, not differentiated in colour from the rest of prp. Spiracular plate divided by a shallow Vshape median groove. Ecdysial scars displaced anteriorly. Diameter of ecdysial scars is almost the length of the spiracular opening I. Dorsal spur absent (compare with Fig. 3 B-D). Spiracular openings straight, mounted on welldeveloped carinae. Carinae about twice as long as wide, extending down the sides of prp. Spiracular opening II closer to III than to I. Spiracular openings II and III parallel, inserted almost at the same level (Fig. 3 A). Interspiracular ornamentation reduced, interspiracular setae very short, hardly visible by light microscope, mounted inside of small circular nodules, periorificial setae between openings I and II close, with their nodules almost connected. Material examined: VENEZUELA: 1 3 puparium+adult, Distrito Federal, cumbre de Boquerón frente a Bajo Seco, 1700 m., 7.iii. 1970, leg. F. Fernández & C.J. Rosales, “ex. Larva predadora de Coccus viridis sobre naranjo” USNM ENT 00037123 (MIZA); 1 3 puparium+adult (imago inside not totally emerged), Falcón, El Supi, 29.xii. 1971, leg. V. Vargas, “ex. larva pred. Aspidiotus destructor en Cocos nucifera ” USNM ENT 00036800 (USNM). FIGURE 1. Dorsal and lateral view of puparium. A, B, Salpingogaster nigra; C, D, Eosalpingogaster umbra; E, F, Eosalpingogaster sp. Arrows point to the transverse rows of segmental tubercles on 2 nd, 3 rd and 4 th abdominal segment., Published as part of Pérez-Bañón, Celeste, Arcaya, Evelin, Mengual, Ximo & Rojo, Santos, 2013, Preimaginal morphology of the genera Salpingogaster Schiner, 1868 and Eosalpingogaster Hull, 1949 (Diptera: Syrphidae), with its systematic implications, pp. 361-370 in Zootaxa 3599 (4) on pages 363-364, DOI: 10.11646/zootaxa.3599.4.4, http://zenodo.org/record/221611, {"references":["Mengual, X. & Thompson F. C. (2011) Carmine cochineal killers: the flower fly genus Eosalpingogaster Hull (Diptera: Syrphidae) revised. Systematic Entomology, 36, 713 - 731. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.2011.00588. x"]}

Salpingogaster nigra Schiner, 1868 Overall appearance. Length 9–10 mm, maximum width 3.0 mm and height 3.5–4 mm. Pear-shaped, oval in crosssection. Anterior extremely truncated, tapered posteriorly and flattened ventrally (see Fig. 1 A, B). Integumental vestiture and segmental sensilla absent (Fig. 2 B). Colour of empty puparium light brown, with three dorsomedial darker spots and a ventrolateral longitudinal dark stripe. Posterior breathing tube (prp): Short, broader than long. Dark brown in colour, lustrous. Spiracular plate little higher than wide, not differentiated in colour from the rest of prp. Spiracular plate divided by a very shallow median groove. Ecdysial scars displaced towards the median groove and anteriorly. Diameter of ecdysial scars is half the length of spiracular opening I. Dorsal spur absent (compare with Fig. 3 B–D). Spiracular opening I displaced towards the centre of the spiracular plate. Spiracular opening I and II mounted on well-developed carinae, straight, parallel and inserted almost at the same level. Carinae of spiracular opening II about three times longer than broad, extending down the sides of prp. Spiracular opening III sinuous with a horseshoe shape (Fig. 3 B). Interspiracular ornamentation reduced, interspiracular setae very short, hardly visible by light microscope, mounted inside of small circular nodules. Interspiracular setae between openings I and II close, with their nodules clearly connected. Material examined: VENEZUELA: 1 Ƥ puparium, Yaracuy, Vía Yaritagua, Hacienda Santa Cruz, 300 m., 10.x. 2000, “sobre Aeneolamia varia ” leg. R. Paz (MJMO); 1 Ƥ adult, Yaracuy, Yaritagua, 300 m., 30.vii. 2002. yellow sticky trap (MJMO); 1 puparium without imago, Yaracuy, Yaritagua, 300 m., 30.vii. 2002, leg. B. Escalona (MJMO); COSTA RICA: 1 3 puparium+adult, Puntarenas Prov., Monteverde, ext. pasture, 16.vii. 1994, D.C. Peck (MZCR); 1 Ƥ puparium+adult, Puntarenas Prov., Monteverde, ext. pasture, 16-23.vii. 1994, D.C. Peck (MZCR)., Published as part of Pérez-Bañón, Celeste, Arcaya, Evelin, Mengual, Ximo & Rojo, Santos, 2013, Preimaginal morphology of the genera Salpingogaster Schiner, 1868 and Eosalpingogaster Hull, 1949 (Diptera: Syrphidae), with its systematic implications, pp. 361-370 in Zootaxa 3599 (4) on pages 365-367, DOI: 10.11646/zootaxa.3599.4.4, http://zenodo.org/record/221611

Eosalpingogaster sp. Overall appearance. Length 6–7 mm, maximum width 2.5–3 mm and height 2.5–3 mm. (see Fig. 1 E, F). Pearshaped, in profile with dorsal and ventral margins of the posterior end forming an acute angle until the anterior bulge (fourth first abdominal segments). Outline slightly serrated, sub-cylindrical in cross-section, anterior end truncate, tapered posteriorly and flattened ventrally. Integumental vestiture and segmental sensilla persisting except on the ventral surface. Dorsal surface covered with short pointed papillae. Colour of empty puparium dull cream with dark brown spots in front of tubercle bases dorsally and along the lateral sides of puparium. The slightly serrated outline is due to medium development of basal papillae of segmental sensilla, as tubercles, on the 2 nd, 3 rd and 4 th abdominal segment. There are three transverse rows of paired tubercles (segmental pairs of sensilla 1, 2, 3). Tubercles of segmental pair of sensilla 1 are less developed on the 2 nd than on the 3 rd and 4 th abdominal segment (Fig. 2 C). Basal papilla of segmental sensilla 5–6 not developed. On the last abdominal segments (5 th, 6 th and 7 th) the basal papillae of segmental sensilla are dried and collapsed not forming tubercles. Dorsal and dorsolateral segmental pairs of sensilla with apical setae visible. The last three abdominal segment (5 th, 6 th and 7 th) and anal segment narrowed significantly towards the base of posterior breathing tube. FIGURE 2. 2nd and 3 th abdominal segments. A, Eosalpingogaster umbra; B, Salpingogaster nigra; C, Eosalpingogaster sp. Arrows point to the tubercles of the segmental pairs of sensilla 1, 2 and 3 on 2 nd abdominal segment. Posterior breathing tube (prp): Short, broader than long. Dark brown in colour, lustrous. Spiracular plate little higher than wide, not differentiated in colour from the rest of prp. Spiracular plate divided by a shallow Vshape median groove. Ecdysial scars displaced anteriorly. Diameter of ecdysial scars is almost the length of the spiracular opening I. Dorsal spur absent (compare with Fig. 3 B-D). Spiracular openings straight, mounted on welldeveloped carinae. Carinae about twice as long as wide, extending down the sides of prp. Spiracular opening II closer to III than to I. Angle between spiracular openings II and III near 90 ° (Fig. 3 C). Interspiracular ornamentation reduced, interspiracular setae very short, hardly visible by light microscope, mounted inside of small circular nodules. Interspiracular setae between opening I and II close, but with their nodules clearly separated. Material examined: VENEZUELA: 1 puparium incorrectly associated to an adult female specimen of E. umbra, Mérida, Timotes, 2050 m., 12.x. 1971, leg. A. Briceño, “ex. larva pred. de Dactylopius Costa en Opuntia ” USNM ENT 00037122 (MIZA).

The puparium of Salpingogaster nigra and Eosalpingogaster umbra are described, and diagnosis and illustrations are provided. The morphology of immature stages of the genera Salpingogaster and Eosalpingogaster are studied and compared for the first time. The results are in agreement with previous molecular analyses that recognise both genera as valid. We found two different puparium morphotypes for E. umbra and discuss the improbability of immature sexual dimorphism in this species. An identification key is also provided for all species with known puparia. This paper was partially funded by grant of Consejo de Desarrollo Científico, Humanístico y Tecnológico (CDCHT) de la UCLA (Universidad Centroccidental “Lisandro Alvarado”) awarded to E. Arcaya PhD project.

Eosalpingogaster sp. Overall appearance. Length 6–7 mm, maximum width 2.5–3 mm and height 2.5–3 mm. (see Fig. 1 E, F). Pearshaped, in profile with dorsal and ventral margins of the posterior end forming an acute angle until the anterior bulge (fourth first abdominal segments). Outline slightly serrated, sub-cylindrical in cross-section, anterior end truncate, tapered posteriorly and flattened ventrally. Integumental vestiture and segmental sensilla persisting except on the ventral surface. Dorsal surface covered with short pointed papillae. Colour of empty puparium dull cream with dark brown spots in front of tubercle bases dorsally and along the lateral sides of puparium. The slightly serrated outline is due to medium development of basal papillae of segmental sensilla, as tubercles, on the 2 nd, 3 rd and 4 th abdominal segment. There are three transverse rows of paired tubercles (segmental pairs of sensilla 1, 2, 3). Tubercles of segmental pair of sensilla 1 are less developed on the 2 nd than on the 3 rd and 4 th abdominal segment (Fig. 2 C). Basal papilla of segmental sensilla 5–6 not developed. On the last abdominal segments (5 th, 6 th and 7 th) the basal papillae of segmental sensilla are dried and collapsed not forming tubercles. Dorsal and dorsolateral segmental pairs of sensilla with apical setae visible. The last three abdominal segment (5 th, 6 th and 7 th) and anal segment narrowed significantly towards the base of posterior breathing tube. FIGURE 2. 2nd and 3 th abdominal segments. A, Eosalpingogaster umbra; B, Salpingogaster nigra; C, Eosalpingogaster sp. Arrows point to the tubercles of the segmental pairs of sensilla 1, 2 and 3 on 2 nd abdominal segment. Posterior breathing tube (prp): Short, broader than long. Dark brown in colour, lustrous. Spiracular plate little higher than wide, not differentiated in colour from the rest of prp. Spiracular plate divided by a shallow Vshape median groove. Ecdysial scars displaced anteriorly. Diameter of ecdysial scars is almost the length of the spiracular opening I. Dorsal spur absent (compare with Fig. 3 B-D). Spiracular openings straight, mounted on welldeveloped carinae. Carinae about twice as long as wide, extending down the sides of prp. Spiracular opening II closer to III than to I. Angle between spiracular openings II and III near 90 ° (Fig. 3 C). Interspiracular ornamentation reduced, interspiracular setae very short, hardly visible by light microscope, mounted inside of small circular nodules. Interspiracular setae between opening I and II close, but with their nodules clearly separated. Material examined: VENEZUELA: 1 puparium incorrectly associated to an adult female specimen of E. umbra, Mérida, Timotes, 2050 m., 12.x. 1971, leg. A. Briceño, “ex. larva pred. de Dactylopius Costa en Opuntia ” USNM ENT 00037122 (MIZA)., Published as part of Pérez-Bañón, Celeste, Arcaya, Evelin, Mengual, Ximo & Rojo, Santos, 2013, Preimaginal morphology of the genera Salpingogaster Schiner, 1868 and Eosalpingogaster Hull, 1949 (Diptera: Syrphidae), with its systematic implications, pp. 361-370 in Zootaxa 3599 (4) on pages 364-365, DOI: 10.11646/zootaxa.3599.4.4, http://zenodo.org/record/221611

Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste, Maier, Chris T. (2009): Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species. Zoological Journal of the Linnean Society 156 (4): 697-749, DOI: 10.1111/j.1096-3642.2008.00503.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00503.x

COPESTYLUM MEXICANUM (MACQUART, 1842) Larva and puparium: Posterior breathing tube shiny below TR. Above TR shiny and faintly punctured. AP about as long as TR is wide and with an incomplete and shallow longitudinal groove from spiracular plates to TR. Pupal spiracles slightly curved from the base and about 2¥ as far apart as pupal spiracle is long. Taxonomic notes: This species may be distinguished from others in the group by the AP which is about as long as TR is wide and the slightly curved pupal spiracles. Material examined: 1 ♂, one puparium MEXICO, Hidalgo, Metztitlán, 18.viii.2005 M. A. M. ex Opuntia sp. (CIBIO)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 720, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM IMITANS (CURRAN, 1926) Puparium: Anterior fold without extra large spicules. Anterior spiracles with three openings. Spicules only between longitudinal folds on the dorsum of the prothorax reaching to middle sensilla i.e. about halfway along folds. Abdominal vestiture of short, upright setae. Anal lobe with spicules. Posterior breathing tube orange, not tapering. AP smooth, mostly shiny and as long as TR is wide. Pupal spiracles dark orange, matt, and with three encircling rings of openings in upper third. Pupal spiracles about 1.5¥ as wide apart as a pupal spiracle is long. Taxonomic notes: Copestylum imitans is recognizable by the dark orange pupal spiracles, nontapering posterior breathing tube and numerous spicules between the longitudinal folds of the prothorax. Copestylum ornatum has similarly dark pupal spiracles but the posterior breathing tube tapers in that species. Material examined (INBio): 2 ♂, 1 ♀, three puparia COSTA RICA, Alajuela, Upala, Dos Rios, Estación San Gerado, 600 m, 13.ix.2000, ex columnar cactus (Cactaceae) D. Briceno; 4 ♀, four puparia COSTA RICA Guanacaste, Santa Rosa, 19.i.2005, ex Opuntia M. Zumbado., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 703, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM LATUM (WIEDEMANN, 1830) Larva and puparium: Anterior fold with a narrow band of spicules. Metathorax with two linear groups of distinct spicules on the anteroventral margin and some spicules on the lateral margins. Dorsal vestiture comprising shorter setae in the middle of transverse bands and with anterodorsal groups of large, broad setae on the mesothorax and metathorax. Abdominal segments 4–6 with second row of crochets inconspicuous and not as developed as on abdominal segments 1–3. Posterior breathing tube red-brown at base, dark brown at apex, below TR tube nodulate, above shiny. AP about a third as high to spiky rim as wide at TR. Pupal spiracles about 2¥ as far apart as a pupal spiracle is long and curved in upper third. Openings coating upper half of one side, those at the apex without a lateral projection. Taxonomic notes: Most similar to C. macrocephalum from which it differs in the broader, shorter posterior breathing tube which above the TR to the spiky rim is only about a third as long as TR is wide. Material examined (CIBIO, NMS): 2 ♂, 1 ♀, two puparia, one larva MEXICO Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 7.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen M. geometrizans; 1 ♂, one puparium MEXICO, Hidalgo, Metztitlán 19.ix.2005., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on pages 718-719, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM MARGINATUM (SAY, 1829) Larva and puparium: Anterior fold with a wide spicule band extending from just above the antennomaxillary organs to the apex. Spicules extending almost to halfway along longitudinal folds of the prothorax, at least to beyond the middle sensillum on mid-dorsal folds. Crochets of metathoracic prolegs forming a w-shaped band. Metathorax with lateral folds bearing broad setae at sensilla 4 and 5. Anal lobe with broad setae. Posterior breathing tube with AP 1.6¥ length of TR, faintly coriacous otherwise shining and not tapering. Pupal spiracles (Figs 51, 53) about 1.3¥ length of AP. Taxonomic notes: Copestylum narginatum is most similar to C. limbipenne Williston, 1887, C. truncatum sp. nov. and C. simile Giglio-Tos, 1892 in having the AP the same length as or longer than the width of the TR. It is distinguished from these species by the numerous spicules on the anterior ends of the longitudinal folds of the dorsum of the prothorax, wide band of spicules on the anterior fold, and a nontapering posterior breathing tube. Material examined: Five puparia, six larvae USA California, Coquillett 4741; two puparia USA Texas, St. Antonio 20.iv.1906 W. Griffiths ex Opuntia (SI); 2 ♂, 5 ♀, seven puparia USA Arizona, Pima Co., about 22 km west of Tucson 21.iii.1979 C. T. M. ex decaying Opuntia engelmannii Salm-Dyck ex Engelmann 1850 (NMS); 2 ♂, 1 ♀, three puparia MEXICO, Oaxaca, Valerio Trujano 14.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying platyclades of Opuntia pilifera F.A.C. Weber 1898 (NMS); 2 ♀, two puparia MEXICO Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 6–10.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying Stenocereus stellatus (Pfeiff.) Riccob. 1909 (NMS); 1 ♂, one puparium MEXICO, Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 6–10.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen Myrtillocactus geometrizans (Mart.) Console 1897 (Cactaceae) (NMS); 14 ♂, 18 ♀, 32 puparia MEXICO Hidalgo, Metztitlán 19.viii–3.ix.2005 M. A. M. (CIBIO). Previous records: Johnston (1921); Bugbee & Reigel (1945); Maier (1982) all ex Opuntia., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 707, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Johnston TH. 1921. Biological control of the prickly-pear pest. Queensland Agriculturalist. 16: 65 - 68.","Bugbee RE, Reigel A. 1945. The Cactus Moth, Melitara dentata (Grote), and its effect on Opuntia macrorrhiza in Western Kansas. American Midland Naturalist 33: 117 - 127.","Maier CT. 1982. Larval habitat and mate-seeking sites of flower flies (Diptera: Syrphidae: Eristalinae). Proceedings of the Entomological Society Washington 84: 603 - 609."]}

COPESTYLUM MACROCEPHALUM (GIGLO- TOS, 1892) Larva and puparium: Anterior fold with a narrow band of spicules. Metathorax with two linear groups of distinct spicules on the anteroventral margin and some spicules on the lateral margins. Dorsal vestiture comprising shorter setae in the middle of transverse bands and with anterodorsal groups of large, broad setae on the mesothorax and metathorax. Abdominal segments 1–3 with three rows of crochets, third row inconspicuous and not as developed as on abdominal segments 4–6. Posterior breathing tube red-brown at base, dark brown at apex, below TR tube nodulate, above shiny. AP about half as high to spiky rim as wide at TR (Figs 68, 70). Pupal spiracles wide apart, about 2.5¥ as far apart as a pupal spiracle is long and curved in upper third. Openings coating upper half of one side, those at the apex without a lateral projection. Taxonomic notes: Most similar to C. latum from which it differs in the narrower, longer posterior breathing tube which above the TR to the spiky rim is about half as long as TR is wide. Material examined: 2 ♂, 2 ♀, four puparia MEXICO, Oaxaca, Santiago Dominguillo 14.vii.1999 E. G. H., M. A. M., G. E. R. ex Ferocactus latispinus var. spiralis (Haw) Britton and Rose (NMS); 3 ♂, 3 ♀, six puparia MEXICO, Oaxaca, Valerio Trujano 14.vii.1999 E. G. H., M. A. M., G. E. R. ex Pachycereus hollianus (CIBIO); 1 ♂, 1 ♀, two puparia MEXICO Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 7.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen M.geometrizans (HM); one larva MEXICO Tehuacán, Puebla Zapotitlán, El Salado 8.vii.1999 ex F. latispinus var. spiralis (NMS); 1 ♂, 1 ♀, two puparia MEXICO Tehuacán, Puebla Zapotitlán, Plan de Fierro 10.vii.1999 ex decaying fallen M. geometrizans (CIBIO); one puparium, one larva COSTA RICA Alajuela, Arado 6.xii.1999 Y. Cárdenas, no further data; 8 ♂, 14 ♀, 22 puparia MEXICO, Chiapas, Mazapa de Maderos, 850 m, 22–25.vi.2004 M. A. M. and C. P. B. (CIBIO)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 719, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM POSTICUM (SAY, 1829) Larva and puparium: Anterior fold with a wide band of spicules. Metathorax with two groups of a few indistinct spicules on the anteroventral margin. Dorsal vestiture comprising evenly sized setae except for anterodorsal groups of large, broad setae on the mesothorax and metathorax and smaller setae on middle transverse bands on abdominal segments 1–2. Abdominal segments 4–6 with second row of crochets inconspicuous and not as developed as on abdominal segments 1–3. Posterior breathing tube red-brown at base, dark brown at apex, below TR tube nodulate, above coriaceous. AP about a third as high to spiky rim as wide at TR. Pupal spiracles approximated about as far apart as a pupal spiracle is wide and slightly curved in upper third (Fig. 71). Openings coating upper half of one side, those at the apex without a lateral projection. Taxonomic notes: Very similar in larval characters to C. isabellina from which C. posticum differs in the pupal spiracles being close, about as far apart as a spiracle is wide, and the openings at the apex lacking spiky projections. Material examined (CIBIO, NMS): 3 ♀, three puparia MEXICO, Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 7.vii.1999 E. G. H., M. A. M., G. E. R. ex one each from decaying N. tetetzo, Stenocereus sp., and an unidentified columnar cactus; 11 ♂, 18 ♀, 29 puparia MEXICO, Chiapas, Mazapa de Maderos, 850 m, 23–25.vi.2004 M. A. M. and C. P. B.., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 719, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM TAMAULIPANUM (TOWNSEND, 1898) Larva and puparium: Anterior fold without extra large spicules. Anterior spiracles with three openings. Abdominal vestiture of short, upright setae. Anal lobe without spicules. Posterior breathing tube orange, tapering. AP smooth, shiny and as long as TR is wide. Pupal spiracles orange, matt, and with three encircling rings of openings in upper third. Pupal spiracles as wide apart as a pupal spiracle is long. Taxonomic notes: Copestylum tamaulipanum is very similar to C. alberlena and C. multipunctatum in having a tapering posterior breathing tube and light orange pupal spiracles. It differs from both these species in the AP that is as long as the TR is wide. In the latter two species the AP is shorter than the TR is wide. Material examined: 1 ♀, one puparium USA Hunter No. 1612-72, 30.xi.1908; 1 ♂, one puparium USA Texas, Victoria 14.xii.1908 JD Mitchell ex Opuntia (SI); 1 ♂, one puparium, six larva MEXICO Veracruz, Alvarado vii.1998 M. A. M. and C. P. B. ex decaying platyclades of Opuntia stricta (Haw) Haw 1812 (CIBIO). Previous records: ex O. stricta (Marcos-García & Pérez-Bañón, 2001, 2002), Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 706, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Marcos-Garcia MA, Perez-Banon C. 2001. Immature stages of Copestylum tamaulipanum and Copestylum lentum (Diptera: Syrphidae). European Journal of Entomology 98: 375 - 385.","Marcos-Garcia MA, Perez-Banon C. 2002. Life cycle, adult and immature stages of a new species of Copestylum (Diptera: Syrphidae) from Mexico reared from Cactaceae. Annals of the Entomological Society America 95: 432 - 440."]}

COPESTYLUM VIOLACEUM (SAY, 1829) Larva and puparium: Posterior breathing tube shiny below TR. AP shiny, faintly punctured about as long as TR is wide, and with a shallow depression just below apex not forming a complete longitudinal groove to the TR. AP about as long as TR is wide. Pupal spiracles curved in upper half and about 1.6¥ apart as pupal spiracle is long. Taxonomic notes: This species may be distinguished from others in the group by the slightly curved pupal spiracles and the faintly punctured AP which is about as long as the TR is wide. Material examined: 2 ♀, two puparia MEXICO Hidalgo 26.iii.1924 no other data (SI); 5 ♂, five puparia MEXICO Hidalgo, Metztitlán 19.viii– 3.ix.2005 ex columnar cactus M. A. M. (CIBIO, NMS)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 720, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM SICA (CURRAN, 1953) Larva and puparium (Fig 27): Posterior breathing tube shiny below TR. Above TR shiny and punctured. AP just over half as long, about 0.65¥ as TR is wide. Pupal spiracles curved from the base and about 1.4¥ as far apart as pupal spiracle is long. Taxonomic notes: This species may be distinguished from others in the group by the punctured AP and the AP just over half as long as the TR is wide. Material examined: 1 ♂, 1 ♀, two puparia, one larva ECUADOR Pinchincha, Guayllamaba 29.viii.2000 Elicio Tapir, E. G. H., G. E. R., ex decaying leaves of a flowered and recently dead unidentified agave (Agavaceae) (NMS); 1 ♂, 4 ♀, five puparia COSTA RICA Guanacaste, Rincón de la Vieja 19.i.2005 C. P. B. and M. Zumbado., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 720, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Curran CH. 1953. Notes and Descriptions of Some Mydaidae and Syrphidae (Diptera). American Museum Novitates 1645: 1 - 15."]}

COPESTYLUM FRAUDULENTUM (WILLISTON, 1891) Larva and puparium (prothoracic plate and pupal spiracles missing): Dorsal abdominal vestiture coating the surface but shorter on either side of the transverse folds. Ventral abdominal vestiture dense and of varying length across folds: anterior sides of folds with longer setae than posterior sides. Posterior breathing tube red-brown, surface corrugated and lightly punctured, about half width of puparium. TR deep and distinct. AP bulging, not tapering, about 0.77¥ as long as TR is wide and with spiracular openings of simple s-shaped curves. Taxonomic notes: The larva and puparium are readily distinguished from those of other species in this group by the clearly discernable TR on the posterior breathing tube and the dorsal abdominal vestiture which completely coats the integument. Material examined: 1 ♂, one puparium USA Arizona, Phoenix 12.ii.1939 collector unknown ex cactus (SI). Previous records: Mann (1969) ex Opuntia., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 712, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Mann J. 1969. Cactus-feeding insects and mites. Bulletin of the US National Museum 256: 1 - 158."]}

COPESTYLUM APICIFERUM (TOWNSEND, 1895) Larva and puparium (Fig. 25): Anterior fold with band of spicules rounded not tapered at outer limits. Longest vestiture of dorsum of prothorax shorter than longest spicules on the anterior fold. Posterior breathing tube about as long as wide above poorly defined TR (Fig. 66). AP tapering to half width of TR and 0.9¥ as long as TR is wide. Pupal spiracles separated by 2.1¥ their length and openings not clustered closely together at the apex but spread down the upper third and separated by about 2¥ diameter of an opening (Fig. 67). Taxonomic notes: Copestylum apiciferum is separated from C. quadratum (Williston, 1891) and C. milae sp. nov. by the pattern of distribution formed by spicules at the ends of the band on the anterior fold that are rounded not tapered, AP not as long as TR is wide and the distribution of the openings on the pupal spiracles, which are distributed over the apical third and not clustered at the apex. Copestylum apiciferum is separated from C. hidalgense sp. nov. by the absence of spicules on the anterodorsal margin of the mesothorax. Material examined: Five larvae USA Arizona, Pima Co., Santa Catalina Mountains, Lower Bear Canyon 1.ii.1965 J. Burger ex decaying Carnegiea gigantea (Englem.) Britton and Rose 1908 (= Cereus giganteus Englem.) (SI); one larva USA Arizona, Pima Co., 20.iv.1973 C. T. M. ex decaying C. gigantea (NMS) 1 ♂, 3 ♀, six puparia USA Arizona, Pima Co., 24 km west of Tucson 21.iii.1979 C. T. M. ex rot pocket of Ferocactus sp. (Britton and Rose 1922) (NMS). Previous records: Ryckman & Ames (1953), Santana (1961) both ex C. gigantea; Maier (1982) ex Ferocactus sp., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 715, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Ryckman RE, Ames CT. 1953. Insects reared from cacti in Arizona. The Pan-Pacific Entomologist 60: 326 - 331.","Santana FJ. 1961. The biology of immature Diptera associated with bacterial decay in the giant sagaro cactus (Cereus giganteus Englemann). MS Thesis, University of Arizona.","Maier CT. 1982. Larval habitat and mate-seeking sites of flower flies (Diptera: Syrphidae: Eristalinae). Proceedings of the Entomological Society Washington 84: 603 - 609."]}

COPESTYLUM FORNAX (TOWNSEND, 1895) Puparium (lacking anterior prothoracic plate): Dorsal abdominal vestiture confined to medial areas on the transverse bands. Ventral abdominal vestiture of varying length across folds: anterior sides of folds with longer setae than posterior sides. Posterior breathing tube red-brown, surface corrugated and shiny, about half width of puparium, and not tapering. TR barely discernable. AP bulging at apex and spiracular openings at apex with more curves than a simple s-shape. Pupal spiracles short, about 1.5¥ basal width, straight, shiny red with openings in upper third and separated by about 4.5¥ their length (Figs 48, 50). Taxonomic notes: Copestylum fornax is more similar to C. satur than to C. fraudulentum. It is distinguished from C. fraudulentum by the shallow TR and vestiture confined to medial areas of abdominal transverse folds. It is separated from C. satur by a redbrown, nontapering AP which is the same length as a pupal spiracle, the shorter, 1.5¥ basal width pupal spiracles which are separated by more than 4¥ their length and by the more complex curves of the spiracular openings. Material examined: One puparium USA Arizona, Tucson 7323 23.i.1897 HG Hubbard ex Opuntia; ten larvae USA Arizona, Yuma 25.ii.1940 HT Rainwater ex decaying Opuntia (SI). Previous records: Mann (1969) ex Opuntia., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 712, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Mann J. 1969. Cactus-feeding insects and mites. Bulletin of the US National Museum 256: 1 - 158."]}

COPESTYLUM ISABELLINA (WILLISTON, 1887) Larva and puparium (Fig. 26): Anterior fold with a wide band of spicules. Metathorax with two groups of a few indistinct spicules on the anteroventral margin (Fig. 30). Dorsal vestiture comprising evenly sized setae except for anterodorsal groups of large, broad setae on the mesothorax and metathorax (Fig. 29) and smaller setae on middle transverse bands on abdominal segments 1–2. Abdominal segments 4–6 with second row of crochets inconspicuous and not as developed as on abdominal segments 1–3. Posterior breathing tube red-brown at base, dark brown at apex, below TR tube nodulate, above coriaceous. AP about a third as high to spiky rim as wide at TR (Fig. 69). Pupal spiracles, about as far apart as a pupal spiracle is long and slightly curved in upper third. Openings coating upper half of one side, those at the apex with a short spiky projection (Figs 72, 73). Taxonomic notes: Very similar in larval characters to C. posticum from which C. isabellina differs in the pupal spiracles being about as far apart as a spiracle is long and the openings at the apex having a short spiky projection. Material examined: 15 larvae USA Arizona, Tucson mountains east of Gates Pass, 1.iii.1965 J. F. Burger ex decaying stems of C. giganteus (Cactaceae) (SI); 1 ♀, one puparium USA Arizona, Yuma Co., Kofz mountains 20.i.1969 J. F. Burger ex decaying stems of C. giganteus (SI) 1 ♂, one puparium USA Arizona, Pima Co., 16 km west of Tucson 14.iii.1979 C. T. M. ex decaying stems of C. giganteus (NMS); 2 ♂, 2 ♀ four puparia USA Arizona, Pima Co., 22.5 km west of Tucson 21.iii.1979 C. T. M. ex rot-pocket in C. giganteus (NMS). Previous records: Hubbard & Schwarz (1899) ex C. giganteus; Ryckman & Ames (1953) ex C. giganteus; Maier (1982) ex C. giganteus; Mangan (1984) ex C. giganteus; Myles (1986) ex C. giganteus., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 718, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Hubbard HG, Schwarz EA. 1899. Insect fauna of the giant cactus of Arizona: letters from the Southwest. Psyche Supplement 1: 1 - 14.","Ryckman RE, Ames CT. 1953. Insects reared from cacti in Arizona. The Pan-Pacific Entomologist 60: 326 - 331.","Maier CT. 1982. Larval habitat and mate-seeking sites of flower flies (Diptera: Syrphidae: Eristalinae). Proceedings of the Entomological Society Washington 84: 603 - 609.","Mangan RL. 1984. Reproductive ecology of three cactophilic Diptera (Diptera: Drosophilidae, Neriidae, Syrphidae). Pan- Pacific Entomologist 60: 326 - 331.","Myles TG. 1986. Oviposition and development of Volucella isabellina (Diptera: Syrphidae) on Saguaro cactus, Cereus giganteus. Entomological News 97: 104 - 108."]}

COPESTYLUM SATUR (OSTEN SACKEN, 1877) Larva and puparium (Fig. 20): Dorsal abdominal vestiture confined to medial areas on the transverse bands. Ventral abdominal vestiture of varying length with anterior ends of folds with longer setae than posterior ends. Posterior breathing tube yellowbrown, surface corrugated and shiny, about half width of puparium. TR barely discernable. AP bulging at apex, tapering slightly and 0.75¥ as long as a pupal spiracle. Spiracular openings at apex with simple s-shaped curves. Pupal spiracles long, about 3¥ as long as basal width, slightly outcurved, shiny red with openings in upper third and separated by 1.4¥ their length (Fig. 49). Taxonomic notes: Copestylum satur is more similar to C. fornax (Townsend, 1895) than to C. fraudulentum (Williston, 1891). It is distinguished from C. fraudulentum by the shallow TR and vestiture confined to medial areas of abdominal transverse folds and from C. fornax by the yellow-brown, slightly tapering posterior breathing tube and the longer pupal spiracles which are about 3¥ basal length and which are separated by less than 2¥ their length. Material examined: 3 ♂, 1 ♀, four puparia, two larvae USA Arizona, Yuma 25.ii.1940 HT Rainwater ex decaying Opuntia (SI). Previous records: Mann (1969) ex Opuntia., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on pages 711-712, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Mann J. 1969. Cactus-feeding insects and mites. Bulletin of the US National Museum 256: 1 - 158."]}

COPESTYLUM SMITHAE (THOMPSON, 1965) Puparium: (prothoracic plate missing), length 4 mm, anterior fold without extra large spicules. Abdominal vestiture of short, upright setae. Anal lobe with indistinct broad setae. Anteriorly directed spicules little sclerotized and indistinct or possibly absent on the ventral surface of the abdomen. Posterior breathing tube orange, not tapering. AP smooth and shiny and about 0.8¥ as long as TR is wide. Pupal spiracles orange, matt, shining apically, and with openings in upper two thirds. Pupal spiracles about 3¥ as wide apart as a pupal spiracle is long. Taxonomic notes: The adult of C. smithae is similar to adults of the Pictum group in general form but has unmarked wings and thick, scale-like setae on the eyes that set this group apart (F. C. Thompson, unpubl. data). The larva is distinct from other species described here by its small size and pupal spiracles separated by about 3¥ their length. The absence of anteriorly directed spicules on the ventral surface of the abdomen indicates that this is not a member of the Pictum group. It was not possible to determine the presence of anterior spiracles, another character indicating membership of the Pictum group, because of the absence of the prothoracic plate from the specimen. Material examined: 1 ♀, one puparium BOLIVIA, Santa Cruz, Jardín Botánico, 17–20.ix.2004 ex Opuntia cactus G. E. R., E. G. H. (NMS)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 706, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM QUADRATUM (WILLISTON, 1891) Larva and puparium: Anterior fold with band of spicules narrowing towards the lateral ends. Longest vestiture of dorsum of prothorax longer than longest spicules on the anterior fold. Posterior breathing tube with AP about as long as wide above inconspicuous TR. AP tapering to about two-thirds of width of TR and 1.1¥ as long as TR is wide. Pupal spiracles about the same length as the AP and separated by about 1.3¥ their length with openings surrounding the apex. Taxonomic notes: Copestylum quadratum is most similar to C. milae from which it differs by having the openings of the pupal spiracles surrounding the apex rather than being on the apex and by having the AP slightly longer than the TR is wide. Copestylum quadratum is separated from C. hidalgense by the absence of spicules on the anterodorsal margin of the mesothorax. Material examined (INBio): 1 ♂, one puparium COSTA RICA San José, Santa Cruz, Central 5.ix.1999 Y. Cárdenas ex decaying cactus; 1 ♂, one puparium COSTA RICA Alajuela, Arado 4.xii.1999 Y. Cárdenas ex decaying cactus; 2 ♂, 1 ♀, three puparia, four larvae COSTA RICA Guanacaste, La Cruz, Santa Elena, Est. Junquilla 29.x.1999 Y. Cárdenas and D. Briceño ex decaying Stenocereus aragonii (F.A.C. Weber) Buxb. 1961 (Cactaceae)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 717, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM ORNATUM (WILLISTON, 1891) Puparium: (prothoracic plate missing), largest species of the group, puparium length 9 mm. Anterior fold without extra large spicules. Thorax with lateral fold bearing sensilla 4 of mesothorax wider than band immediately below bearing sensilla 5. Abdominal vestiture of short, upright setae. Anal lobe without spicules. Posterior breathing tube orange, tapering. AP punctured and shiny, and as long as TR is wide. Pupal spiracles dark orange, matt, shining apically, and with openings in upper third. Pupal spiracles 1.1¥ as wide apart as a pupal spiracle is long. Taxonomic notes: Copestylum ornatum is similar to those species of the Pictum group having a tapering posterior breathing tube. It can be separated from these species by its large size (puparium length 9 mm), upper thoracic lateral fold being wider than the fold immediately below and having dark orange pupal spiracles. In having dark orange pupal spiracles it is similar to C. imitans but may be separated from that species by its tapering posterior breathing tube. Material examined (INBio): 1 ♂, one puparium COSTA RICA Alajuela, La Garita, Fca. Periodistas 14.i.1994 ex Hylocereus costaricensis (F.A.C. Weber) Britton and Rose 1909 (Cactaceae); 1 ♂, 1 ♀, two puparia COSTA RICA Heredia, Santo Domingo, INBioparque 15.vii.2000 ex decaying columnar cactus E. G. H., G. E. R.., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on pages 704-705, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM COCHABAMBA ROTHERAY, MARCOS & HANCOCK SP. NOV. Adult: Male holotype: face with lateral margins inflated and with slight lateral tubercles near the eye margins below the antennae. Face indented slightly from just above margin of mouth so lower part of face appears as a narrow projection (view from in front). Face and genae yellow and yellow haired except for some black hairs on the central tubercle and on the lower lateral margins of the face. Face without vittae. Tubercle large and prominent and elongate oval in side view. Mid point of tubercle appearing below the mid point between the antennae and the mouth Fig. 5). Frons yellow with black hairs. Antennae yellow with plumose arista and basoflagellomere slightly constricted in the middle and 2.5¥ as long as broad, rounded apically. Ocellar triangle dark yellow and black haired, suture about same length as ocellar triangle. Eyes with yellow hairs reaching lower margins, slightly longer on top of head. Ground colour of mesonotum black except for vague yellowish lateral and posterior margins and transverse suture also yellow marked. Postpronotum well developed, forming a rounded yellow lobe as wide as long with long, yellow and black hairs and two to three black bristles on posterolateral margin. In profile view mesonotum has mixed length pale and black hairs but when viewed from behind, mesonotum has an alternating pattern of hairy, gold and black stripes. Lateral margins with mixed long pale hairs and black bristles. Posterior margin with a row of eight+, thin, prescutellar black bristles, these bristles stronger towards the medial line. Scutellum dark yellow with mixed length black hairs except for pale haired margins and posterior margin with six very strong, socketed, black bristles. Pleurae black except for yellow mark on dorsal margin of posterior anepisternum and densely coated in long yellowish hairs and a row of six+ black bristles on the posterior anepisternum. Wing membrane hyaline, without microtrichia. Legs black with yellowish tibiae and completely black haired. All sternites brown, shiny with long pale hairs. Tergites 1–3 dark yellow, tergite 4 black all coated in short and posteriorly reclined black hairs. Hairs on tergite 4 longer than on other abdominal tergites. Genitalia: in profile view, surstylus longer than basally wide, rounded and slightly tapered at the apex with dark bristles on the inner and anterior sides. Cerci flat tipped. In profile view, epandrium longer than basally broad. Hypandrium indented ventrobasally with below, a rounded ridge across the ventral margin. Hypandrium smoothly punctured and covered with fine pile. Superior lobe dorsally rounded and with a hook shaped apex (Figs 10, 11). Female: Similar to holotype male; width of vertex about 2/5 width of head. Length: Body 13.0– 13.1 mm; wing 12.5 mm (N = 2). Puparium: Anterodorsal margin of mesothorax with enlarged setae and spicules. Dorsal vestiture of abdomen spread across folds and not strongly confined to medial transverse bands. Crochets translucent with red tips. Posterior breathing tube dark brown, short, about 1/10 as wide as puparium, AP only 0.6¥ as long as TR is wide, coriaceous not tapering and slightly bulging ventrally with midventral groove ending before halfway point to TR. Pupal spiracles about 1.6¥ as long as AP and 1.2¥ as wide apart as a pupal spiracle is long. Material examined: Holotype: male with puparium BOLIVIA, Cochabamba, ix.2004, ex decaying Opuntia. Known only from this locality, collected by G. E. R. and E. G. H. (NMS). Paratype: 1 ♀ with puparia, same data as holotype (HM). Etymology: The name refers to ‘Cochabamba’, the Bolivian locality where the larva of the type specimen was collected. Taxonomic notes: In the Scutellata group key of Fluke (1951), the adult of C. cochabamba keys out to C. rectum (Wulp, 1882). Fluke (1951) provided a description of a species that he determined as C. rectum but questioned whether his determination was correct on the basis of discrepancies between his specimens and the description of recta given by Wulp. Copestylum cochabamba differs from both C. recta Wulp and C. rectum sensu Fluke (1951) in the following combination of characters: a flattened not a well rounded facial tubercle, black not pale haired facial margins, pale not black hairs on the abdominal sternites, and in the male genitalia by a rectangular not squareshaped epandrium with surstyli that are taller rather than shorter than basally long. The puparium of C. cochabamba is distinguished from other early stages of the Scutellatum group by the dark brown posterior breathing tube which does not taper, AP about half as long as TR is wide and the pupal spiracles which are about as long as TR is wide., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on pages 713-714, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Fluke CT. 1951. Syrphid flies related to Volucella scutellata Macquart. American Museum Novitates 1503: 1 - 83."]}

COPESTYLUM SIMILE GIGLO- TOS, 1892 Larva and puparium: Anterior fold with a wide spicule band extending from just above the antennomaxillary organs to the apex. Spicules confined to the extreme ends of the longitudinal folds of the prothorax, only one or two further along between the folds. Crochets of metathoracic prolegs forming a w-shaped band. Metathorax with lateral folds bearing broad setae at sensilla 4 and 5. Anal lobe with spicules. Posterior breathing tube with AP 1.4¥ length of TR, matt and shiny in parts and tapering. Pupal spiracles about 0.78¥ length of AP. Taxonomic notes: Copestylum simile is most similar to other members of the Marginatum group having an AP longer than the width of the TR. It differs from these species in having a wide band of spicules on the anterior fold, bands of broad setae on the anterior margins of the transverse folds on the dorsum of the abdomen, and in having a tapered posterior breathing tube. Material examined (CIBIO, NMS): 2 ♀, two puparia MEXICO, Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 6–10.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen M. geometrizans (Cactaceae); 1 ♂, 2 ♀, three puparia MEXICO, Tehuacán, Puebla Zapotitlán, Salinas, Plan de Fierro 10.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen unidentified cactus; 1 ♂, one puparium MEXICO, Tehuacán, Puebla Zapotitlán, Salinas, Plan de Fierro 10.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying fallen M. geometrizans., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 709, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM HAMBLETONI (FLUKE, 1951) Puparium: Anterodorsal margin of mesothorax with enlarged setae, not spicules. Dorsal vestiture of abdomen spread across folds and not strongly confined to medial transverse bands although shorter in front and behind. Crochets translucent with dark red tips. Posterior breathing tube dark brown, long almost as long as puparium is wide with AP about 0.8¥ as long as TR is wide, coriaceous and faintly punctate and tapering sharply to about 0.5¥ of width of TR (Fig. 61). Pupal spiracles short, about 0.7¥ as long as AP and about 1.4¥ as wide apart as a pupal spiracle is long (Figs 56, 57). Material examined: 2 ♂, 3 ♀, five puparia (CIBIO), 1 ♀ and puparium (NMS), PERU, Trujillo, Cerro Campana, 17.i.2005, ex decaying columnar Cactaceae, E. melanostele. Known only from Cerro Campana, Trujillo, Peru, collected by M. A. M. (CIBIO). Taxonomic notes: The puparium of C. hambletoni is distinguished from other Scutellatum group species considered here by the long posterior breathing tube with its sharply tapering AP and the crochets on the prolegs, which have dark red tips., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 714, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Fluke CT. 1951. Syrphid flies related to Volucella scutellata Macquart. American Museum Novitates 1503: 1 - 83."]}

COPESTYLUM DISPAR (MACQUART, 1846) Puparium: Posterior breathing tube, shiny and faintly nodulate below TR. Above TR shiny to coriaceous. AP 0.75¥ as long as TR is wide, faintly punctate. Pupal spiracles sharply bent over at halfway point and separated by 1.2¥ the length of a pupal spiracle. Taxonomic notes: This species may be distinguished from others in the group by the sharply curved pupal spiracles and the faintly punctured AP which is about 0.75¥ as long as TR is wide. Material examined (CIBIO, NMS except where stated): 2 ♂, 3 ♀, five puparia MEXICO Oaxaca, Valerio Trujano 14.vii.1999 E. G. H., M. A. M., G. E. R. ex P. hollianus; 1 ♀, one puparium MEXICO Oaxaca, Valerio Trujano 15.vii.1999 E. G. H., M. A. M., G. E. R. ex F. latispinus; 2 ♀, two puparia COSTA RICA Puntarenas, Buenos Aires, Sabanas Esperanza, 3–9.ix.2000 BHB ex decaying leaves of Furcraea cabuya Trel. (Agavaceae) (INBio); 1 ♂, 4 ♀, five puparia MEXICO Chiapas, Mazapa de Maderos, 850 m, 22–25.vi.2004 ex columnar cactus M. A. M. and C. P. B.., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 720, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM PACHECOI (CURRAN, 1939) Puparium: Anterior fold without extra large spicules. Anterior spiracles with three openings. Anterodorsal margin of metathorax with some spicules. Abdominal vestiture of long, fine setae. Anal lobe with spicules. Posterior breathing tube orange, not tapering. AP smooth and shiny and about 1.2¥ as long as TR is wide. Pupal spiracles orange, matt, shining apically, and with three encircling rings of openings in upper half. Pupal spiracles about 1.5¥ as wide apart as a pupal spiracle is long. Taxonomic notes: Copestylum pachecoi is similar to those Pictum group species with a nontapering posterior breathing tube. It differs from these species in having spicules and broad setae on the anterodorsal margin of the metathorax. Material examined: 7 ♀, seven puparia MEXICO Chiapas, Mazapa de Madero, 850 m, 23–25.vi.2004, ex Opuntia cactus, M. A. M. and C. P. B. (CIBIO)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 705, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM COCKERELLI (CURRAN, 1927) Puparium: Anterodorsal margin of mesothorax with enlarged setae (conspicuously longer than surrounding vestiture, not spicules. Dorsal vestiture of abdomen confined to medial areas of transverse folds. Crochets entirely black. Posterior breathing tube dark brown, short, less than half as wide as puparium, AP short, only about 0.4¥ as long as TR is wide, lightly punctate, tapering but bulging out horizontally (Fig. 60). Pupal spiracles as long as AP and about 2.5¥ as wide apart as a pupal spiracle is long. Material examined: 1 ♂, 3 ♀, four puparia PERU, Trujillo, Cerro Campana, 17.i.2005, ex decaying columnar Cactaceae, Espostoa melanostele (Vaupel) Bullock 1938, collected by M. A. M. (CIBIO); 1 ♀, one puparium same data as previous specimens (NMS). Taxonomic notes: The puparium of C. cockerelli is distinguished from other Scutellatum group larvae considered here by the dorsal vestiture of the abdomen being confined to medial areas of the transverse folds, the entirely black crochets, and the short, horizontally bulging AP., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 714, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM TETETZOI MARCOS, HANCOCK & ROTHERAY SP. NOV. Adult: Male holotype: face with lateral margins inflated and sharply tapered from just above margin of mouth (view from in front). Face yellow with a narrow (about as wide as antennal bases are apart) black, mid-dorsal vitta extending from the base of the antennae to the margin of the mouth and face yellow haired. Tubercle slight and below mid point between antennae and mouth. Frons black with black hairs. Lateral vittae black reaching mouth and genae black and pale haired (Fig. 3). Antennae dark yellow and arista densely pilose with short, incurved hairs and basoflagellomere dorsally not excavated and more than 3¥ as long as broad. Ocellar triangle black with long black hairs and suture same length as ocellar triangle. Eyes black haired, hairs longer at top of head and reaching lower margin. Eye hairs denser and darker in the middle of the eye forming a vertical, black stripe. Ground colour of mesonotum black except for uniformly pale yellow lateral margins and a medial pair of triangular shaped yellow markings on the posterior margin. Mesonotum with mostly long hairs with a few shorter ones on the posterior margin. Longer hairs about 2¥ as long as shorter hairs. Hairs white except for some black hairs on the lateral margins and short, black hairs on the posterior margin. Mesonotum with a pair of dusted stripes dorsocentrally ending before the transverse suture. Scutellum yellow with short black hairs and long pale hairs. Pleurae black, only pale on the dorsal part of posterior anepisternum and with long white hairs on posterior anepisternum, anepimeron, and katepisternum. Wings hyaline, without markings or microtrichia. Legs black haired and blackish with knees and basal two thirds of tibiae yellow. Sternites 1, 2, and 3 yellow and pale haired, fuscous medially, remaining sternites black and black haired except anterior corners of sternite 4 with a pair of yellow spots. Tergites black except tergite 2 with yellow triangular spots tapering almost to a point towards the medial line and reaching the side margins. Tergite 3 with a pair of semicircular spots in anterior two thirds and not reaching side margins. Tergite 4 with a pair of yellow bars on the anterior margin and not reaching the side margins. Tergite 1 pale haired except for posterior lateral margins. Tergite 2 pale haired except black haired on posterior third and medial line. Tergite 3 black haired except for pale hairs on anterolateral margins. Male genitalia: epandrium subrectangular in shape; surstylus square in shape, truncate at apex; hypandrium not inclined backwards, straight in lateral view; superior lobe with a hook-like, sharply tapered apex (Figs 14, 15). Female: Similar to holotype male except: frons with one medial and two lateral grooves; width of vertex about one fourth width of head. Length: Body 8.0–9.0 mm; wing 8.0– 9.5 mm (N = 8). Larva and puparium (Fig. 21): Anterior fold with a wide spicule band extending from just above the antennomaxillary organs to the apex. Spicules confined to the extreme ends of the longitudinal folds of the prothorax, only one or two further along between the folds. Crochets of metathoracic prolegs forming a w-shaped band. Metathorax with lateral folds bearing broad setae at sensilla 4 and 5. Anal lobe with spicules. Posterior breathing tube with AP 0.8¥ length of TR, with light punctures and not tapered. Pupal spiracles about 0.85¥ length of AP. Material examined: holotype: male with puparium MEXICO, Puebla, Tehuacán, Jardín Botánico, 1590 m (18°19′58″N, 97°27′28″W), 7.vii.1999, ex live decaying Neobuxbaumia tetetzo (F.A.C. Weber) Backeberg 1938 (Cactaceae), collected by M. A. M. (CIBIO). Paratypes: 2 ♂ and 3 ♀ with puparia, same data as holotype (CIBIO); 1 ♂, MEXICO, Puebla, San Juan Raya, 9.vii.1999, collected by M. A. M. (CIBIO), ex decaying Cactaceae; 1 ♀, MEXICO, Oaxaca, Santiago Dominguillo, 14.vii.1999, collected by M. A. M. (CIBIO), ex decaying Cactaceae; 1 ♀ MEXICO, Puebla, Zapotitlán Salinas, Jardín Botánico, 10.vii.1999, ex live decaying (Cactaceae), collected by E. G. H. (HM); 1 ♀, one puparium MEXICO, Oaxaca, Santiago Dominguillo 14.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying stem of Ferocactus latispinus var. spiralis (Karw. ex Pfeiller) Taylor 1984; 3 ♂, 4 ♀, seven puparia, one larva MEXICO Tehuacán, Puebla Zapotitlán, Salinas, Jardín Botánico 7.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying stems of fallen N. tetetzo; 1 ♂, one puparium MEXICO, Tehuacán, Puebla, Zapotitlán, San Juan Raya (El Salado) 9.vii.1999 E. G. H., M. A. M., G. E. R. ex decaying stem of fallen Neobuxbaumia mezcalaensis (Bravo) Backeb 1941 (Cactaceae) (NMS). Etymology: The name tetetzoi refers to the cactus species Neobuxbaumia tetetzo from which the holotype was reared. Taxonomic notes: The adult of this species can be easily separated from other Marginatum group adults by the male genitalia, black frons, and wings hyaline without brown spots. The larva and puparium of C. tetetzoi is similar to C. bequaerti and C. lentum in having an AP which is shorter than the width of the TR. The nontapering punctured AP separate C. tetetzoi from C. lentum and the absence of anterior spiracles separates it from C. bequaerti., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on pages 709-710, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474

COPESTYLUM LENTUM (WILLISTON, 1887) Larva and puparium: Anterior fold with a wide spicule band extending from just above the antennomaxillary organs to the apex. Spicules confined to the extreme ends of the longitudinal folds of the prothorax, only a few further along between the folds. Anterodorsal margin of metathorax with broad setae and spicules. Crochets of metathoracic prolegs forming a w-shaped band. Metathorax with lateral folds bearing spicules at sensilla 4 and 5. Anal lobe with broad setae. Posterior breathing tube with AP 0.9¥ length of TR, smooth, shiny and not tapered. Pupal spiracles about 1.1¥ length of AP. Taxonomic notes: Copestylum lentum is similar to other members of the Marginatum group with an AP shorter than the width of the TR. The absence of anterior spiracles, mixed long and short abdominal vestiture, smooth not punctured AP, and spicules on the anterodorsal margin of the metathorax separate this species from others in this species group. Material examined: 1 ♂, one puparium MEXICO Chapingo 15.iv.1964 CG Martell ex ‘nopal’; 1 ♀, one puparium no data (SI); 1 ♂, one puparium, two larvae MEXICO Veracruz, Guadalupe Victoria 26.vii.1998 ex Opuntia M. A. M. (CIBIO). Previous records: ex Opuntia (Marcos-García & Pérez-Bañón, 2001)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 708, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Marcos-Garcia MA, Perez-Banon C. 2001. Immature stages of Copestylum tamaulipanum and Copestylum lentum (Diptera: Syrphidae). European Journal of Entomology 98: 375 - 385."]}

COPESTYLUM CURRANI (FLUKE, 1951) Puparium: Anterodorsal margin of mesothorax with enlarged setae, not spicules. Dorsal vestiture of abdomen spread across folds and not strongly confined to medial transverse bands. Crochets translucent with red tips. Posterior breathing tube dark brown, about 1/10 as wide as puparium, AP about 0.67¥ as long as TR is wide, lightly punctate and slightly tapering. Pupal spiracles about as long as AP and about 1.8¥ as wide apart as a pupal spiracle is long. Taxonomic notes: The puparium of C. currani is distinguished from other Scutellatum group species by the slightly tapering posterior breathing tube which is dark brown and with the AP just over half as long as TR is wide. Material examined: 3 ♂, 3 ♀, six puparia, one larva ECUADOR Pinchincha, Guayllamaba 29.viii.2000 E. G. H., G. E. R. and Elicio Tapia ex decaying platyclades of Opuntia and a small columnar cactus (Cactaceae) (NMS, PUCE)., Published as part of Rotheray, Graham E., Marcos-García, Maria-Angeles, Hancock, Geoff, Pérez-Bañón, Celeste & Maier, Chris T., 2009, Neotropical Copestylum (Diptera, Syrphidae) breeding in Agavaceae and Cactaceae including seven new species, pp. 697-749 in Zoological Journal of the Linnean Society 156 (4) on page 714, DOI: 10.1111/j.1096-3642.2008.00503.x, http://zenodo.org/record/5443474, {"references":["Fluke CT. 1951. Syrphid flies related to Volucella scutellata Macquart. American Museum Novitates 1503: 1 - 83."]}