Systematics and Phylogeny of Leptomorphus Curtis (Diptera: Mycetophilidae) 3549
Borkent, Christopher J.
Wheeler, Terry A.
Zootaxa
2012
2012-11-15
3549
1
117
[356,625,151,177]
Insecta
Mycetophilidae
Leptomorphus
Animalia
Diptera
10
11
Arthropoda
species
undetermined
1. Vein R 4present (i.e. Figs. 64, 65)......................................................................... 2 - Vein R 4absent (i.e. Fig. 57)............................................................................. 8 2. (1) Lateral ocelli no more than their own diameter away from eye margin ( Fig. 79); foretarsomere slightly shorter than foretibia; male unknown ( USA: California, Fig. 148)................................... L. perplexusBorkent, n. sp. - Lateral ocelli more than their own diameter away from eye margin, grouped closely with median ocellus at vertex ( Fig. 78); foretarsomere I longer than foretibia...................................................................... 3 3. (2) Vein R 4joining Rs or junction of Rs and r-m, forming a triangular cell ( Fig. 65); genitalia as in Fig. 112(southeastern South America, Fig. 142)..................................................................... L. neivaiEdwards. - Vein R 4joining Rs beyond junction of Rs and r-m, forming a quadrilateral cell (i.e. Figs. 60, 64) (Nearctic, Palaearctic).... 4 4. (3) Apex of antennae (at least apical 5 flagellomeres) white, base black or dark brown (e.g. Figs. 24–27, 31)............. 5 - Antennal flagellomeres (except, rarely, base of first flagellomere) uniformly coloured (e.g. Figs. 1–7).................. 6 5. (4) Male with large apical spine-like process on midfemur ( Fig. 83); male genitalia with two long sickle-like structures curving dorsally ( Fig. 111); colour varying from completely black to mostly yellow with dark brown markings on thorax and abdomen ( Figs. 24–27) ( Canada, northern and eastern USA, Fig. 144)................................. L. nebulosus(Walker). - Male without large apical spine-like process on midfemur; male genitalia as in Fig. 114without long sickle-like structures; body completely black/ dark brown ( Fig. 31) ( Japanand far eastern Russia, Fig. 141)...... L. panorpiformis(Matsumura). 6. (4) Male genitalia with two long sickle-like structures curving dorsally ( Figs. 108, 117); scutum with setae on entire surface; colour either yellow with brown head, or dark brown/ black (females with tergite 4 red-orange) with yellow coxae and femora ( Figs. 21, 36–37)...................................................................................... 7 - Male genitalia without sickle-like structures ( Fig. 94); scutum with setae restricted to dorsocentral line and margins; thorax varying from dark to light brown, when light brown scutum with two posteriorly converging dark lines; abdomen varying from light to reddish brown, tergites 1–2 often dark brown ( Figs. 4–5) (southern Canada, south in eastern USAto NC, Fig. 147)....................................................................................... L. bifasciatus(Say). 7. (6) Completely yellow with brown head ( Fig. 21) (eastern North America, Fig. 145)........... L. magnificus(Johannsen). - Thorax and abdomen completely dark brown/black in male, female identical except with tergite 4 reddish-brown; coxae and femora yellow ( Fig. 36–37) (central and eastern North America, Fig. 146)................. L. subcaeruleus(Coquillett). 8. (1) Scutum evenly covered with trichia ( Fig. 80) or fine setae; male foretibia with 1–2 dense rows (combs) of short anteroventral bristles (spacing between bristles usually less than bristle length) ( Figs. 85, 89–90).............................. 9 - Scutal setae or trichia (if present) restricted to dorsocentral and lateral rows ( Fig. 81); male foretibia without dense row of anteroventral bristles ( Figs. 87–88)...................................................................... 22 9. (8) Scutellum with covering of yellow or white fine trichia and without bristles; Neotropical or Afrotropical............ 10 - Scutellum without covering of fine trichia, but with brown, small or large, bristles (i.e. with sockets); Nearctic or Palaearctic. ................................................................................................... 17 10. (9) Palp segment 5 with thick covering of fine, white setulae and with or without a few setae ( Fig. 82); vein sc-r present (i.e. Fig. 57); Neotropical.................................................................................. 11 - Palp segment 5 without covering of fine, white setulae, but with a few dark setae; vein sc-r absent ( Fig. 58); genitalia as in Fig. 105(eastern Madagascar, Fig. 134)........................................................ L. grjebineiMatile. 11. (10) Scutellum dark brown (rarely yellow), same colour as scutum and mediotergite............................... 12 - Scutellum light brown, yellow or white, distinctly paler than scutum and mediotergite.............................. 15 12. (11) Fore- and midfemora and tibia yellow ( Figs. 3, 14)...................................................... 13 - Fore- and midfemora with brown or dark brown spots basally extending from 1/3 to ¾ towards apex, fore- and midtibia with apical ½ or entirely brown or dark brown ( Figs. 6, 15)....................................................... 14 13. (12) At least abdominal segments 2–4 mostly yellow ( Fig. 14); metepisternum yellow or white; male genitalia with gonocoxite arising at base of tergite 9 and almost as long as tergite 9, gonocoxite with long thin process, as long as gonocoxite, arising mediobasally and armed with 4 long apical bristles pointed laterally ( Fig. 101) ( Panama, northern South America and Amazonia, Fig. 142)................................................................ L. fasciculatusEdwards. - Abdominal segments 2–4 with some brown or dark brown ( Fig. 3); metepisternum completely or partially brown; male genitalia with gonocoxite arising ¼ of length of tergite 9 towards apex, gonocoxite with scalloped edge apicomedially and hook-like process laterally ( Fig. 92) (southern Brazil, Fig. 142)..................................... L. amorimiBorkent, n. sp. 14. (12) Metepisternum brown; male genitalia yellow or white; tergite 9 rounded basally, tapering to a single, long, point posteriorly that extends beyond cerci ( Fig. 95) ( Costa Rica, Fig. 143)............................. L. brandiaeBorkent, n. sp. - Metepisternum yellow or yellow with brown dorsally; male genitalia partially brown; apex of tergite 9 not extending past cerci; tergite 9 posteriorly with some scalloping on apicolateral corner and with 3 rounded medial processes, with central shorter than laterals ( Fig. 102) ( Peruand Bolivia, Fig. 143)................................... L. femoratusEdwards. 15. (11) Abdomen mostly yellow, brown posteriorly (posterior half of tergite 5 and all of T6–7, Fig. 39); medial wing spot (macula) only around junction of R and M 1+2, not reaching anterior or posterior wing margin as a complete band ( Fig. 77); halter yellow; antennal flagellomere 6 almost square (slightly longer than wide); male genitalia with tergite 9 tapering posteriorly to a point, with short pointed processes laterally at 2/3 of length towards apex; gonocoxite with long projection on medial margin ( Fig. 124); female unknown ( Ecuador, Fig. 143)......................................... L. waodaniBorkent, n. sp. - Abdomen striped brown and yellow ( Figs. 10, 12); medial wing spot (macula) strong and running from anterior to posterior wing margin as a complete band, though fainter posteriorly ( Figs. 50, 52); halter with base of stem yellow, remainder brown; antennal flagellomere 6 1.5–2X longer than wide; male genitalia not as above.................................... 16 16. (15) Male genitalia with tergite 9 tridentate posteriorly (medial process laterally compressed); gonocoxite placed posteriorly on tergite 9, gonostylus small (~1/4 length of gonocoxite) with two short, round, apical projections ( Fig. 100) ( Costa Rica, Fig. 143)........................................................................... L. eberhardiBorkent, n. sp. - Male genitalia with tergite 9 posterior margin rounded laterally and concave medially, lateral process apex with 5 rows of tightly spaced short blunt brown bristles; gonocoxite placed basally on, and subequal in length to, tergite 9; gonostylus ~2/3 length of gonocoxite with two apical processes, medial one most pronounced and tapering to a point, the lateral ½ the length of medial, thin and rod-like ( Fig. 98); female unknown (northern Argentina( Tucumán province), Fig. 142).............................................................................................. L. crassipilusBorkent, n. sp. 17. (9) Abdomen brown or dark brown ( Fig. 33)............................................................... 18 - Abdomen mostly yellow, tergite 7 and often posterior margin of other tergites brown ( Figs. 16–17, 41)................. 19 18. (17). Scutum and mediotergite dark brown; gonocoxite with lateral process of uniform thickness on apical 2/3 ( Fig. 116) (far eastern Palaearctic, Fig. 141).................................................. L. quadrimaculatus(Matsumura). - Scutum with posterolateral corners yellow; mediotergite yellow with central brown spot; gonocoxite with lateral process swollen at apex ( Fig. 118) (north-western Russiaand Finland, Fig. 140)................... L. subforcipatusZaitzev & Ševčík. 19. (17) Wing with apical spot very pale and restricted to apical ¼ of cell r1; medial spot absent ( Fig. 56); male genitalia with sternite 9 bearing a medial invagination for posterior 2/3 and with 2 dark, apical points ( Fig. 104) ( New Mexico, Arizonaand Northern Mexico, Fig. 148).......................................................... L. furcatusBorkent, n. sp. - Wing with dark apical and medial spots present ( Figs. 55, 76); male genitalia with sternite 9 two dimensional (no ventral points or invaginations, Figs. 103, 123)......................................................................... 20 20. (19) Halter yellow; male genitalia with gonocoxite shorter than tergite 9; sternite 9 not reaching lateral margins of tergite 9 ( Fig. 123) (Europe, Fig. 140)............................................................... L. walkeriCurtis. - Halter with stem yellow and knob brown; male genitalia with gonocoxite at least 1.5X length of tergite 9; sternite 9 wider than long, reaching lateral margins of tergite 9 ( Figs. 103, 121).................................................... 21 21. (20) Gonocoxite straight for ¾ of length and then angularly bent medially and swollen beyond bend ( Fig. 121) ( Azerbaijan, Fig. 140)................................................................... L. talyshensisZaitzev & Ševčík. - Gonocoxite smoothly curved along entire length, not swollen at apex ( Fig. 103) (Europe, Fig. 140)... L. forcipatusLandrock. 22. (8) Laterotergite and anepisternum brown; anepimeron at least partially brown ( Figs. 7, 40); Orientalor Palaearctic....... 23 - Laterotergite, anepisternum and anepimeron yellow (i.e. Figs. 8, 20); Afrotropical or Nearctic........................ 28 23. (22) Katepisternum brown; metepisternum at least partially brown or light brown; wing without apical spot ( Figs. 48, 73, 74); segments 3–5 of abdomen noticeably swollen relative to other segments ( Figs. 35, 40).............................. 24 - Katepisternum and metepisternum yellow; wing with pale brown apical wing spot reaching to wing tip ( Figs. 44, 67, 75); abdominal segments all relatively the same thickness ( Figs. 2, 30, 38)........................................... 26 24. (23) Scutum dark brown............................................................................... 25 - Scutum dark brown with small yellow spots on posterolateral corners; genitalia with tergite 9 bearing two long lateral processes posteriorly, so that tergite 9 is ~2X as long as wide ( Fig. 120); female unknown ( Sulawesi, Fig. 137)............................................................................................... L. tabatiusBorkent, n. sp. 25. (24) Metepisternum brown; antennal flagellomere 1 with base yellow, remainder brown; vein sc-r joining Rwithin its own length of origin of Rs; known only from female holotype( Fig. 7) ( northern Borneo, Fig. 137).......... L. chaseniEdwards - Metepisternum brown anteriorly, yellow posteriorly; antennal flagellomere 1 yellow; vein sc-r joining R at ~2X its own length from origin of Rs; genitalia wider than long ( Fig. 119); female unknown ( Philippines, Fig. 137).. L. tagbanuaBorkent, n. sp. 26. (23) Scutellum yellow; pale medial wing spot absent ( Figs. 67, 75); genitalia with gonostylus bearing two apical projections ( Fig. 122)........................................................................................... 27 - Scutellum dark brown; pale medial wing spot present ( Fig. 44); genitalia with gonostylus a single blunt taper ( Fig. 93); female unknown ( Japan, Fig. 141)............................................................... L. babaiSasakawa. 27. (26) Vein sc-r present (though faint in one specimen); genitalia as in Fig. 122( Peninsular Malaysia, Fig. 137)........................................................................................ L. titiwangsensisBorkent, n. sp. - Vein sc-r absent or present; genitalia presumably different from above (male unknown) (eastern Indiaand Nepal, Fig. 137)........................................................................................ L. ornatusBrunetti. 28. (22) Scutellum brown or dark brown..................................................................... 29 - Scutellum yellow..................................................................................... 32 29. (28) Halter with most of stem and knob dark brown; gonostylus with two large lobes ( Figs. 99, 106, 113), Afrotropical; females not currently distinguishable beyond this point............................................................. 30 - Halter mostly yellow with only tip of knob brown; gonostylus with single large, hook-tipped lobe ( Fig. 107) Nearctic ( Fig. 149)............................................................................... L. hyalinusCoquillett. 30. (29) Genitalia with tergite 9 gradually tapering on apical quarter, apex bearing medial spur and often a secondary small bump or point, also with strong, ventrally-produced, thin ridge running across ventral surface at approximately 2/3 the distance towards apex, creating a semicircular dorsal margin when viewed caudally ( Fig. 113) (central-western Africa, Fig. 136).......................................................................................... L. obscurusMatile. - Genitalia with tergite 9 ending in more or less blunt apex laterally and with or without small spine-like projection on medial corner (e.g. Figs. 99, 106).............................................................................. 31 31. (30) Genitalia with posterolateral projection of tergite 9 either square or with slightly acute angle on medial corner; gonostylus with shortest lobe rounded ( Fig. 106); female unknown ( Gabonand Central African Republic, Fig. 135)... L. gracilisMatile. - Genitalia with posterolateral projection of tergite 9 bearing distinct thin point on medial corner; gonostylus with shortest lobe gradually tapering to thick point ( Fig. 99) (central western Africa, Fig. 136)........................ L. crosskeyiMatile. 32. (28) Abdomen mostly yellow with at most brown bands posteriorly on tergites 3–6, tergite 7 brown or yellow ( Figs. 22, 23, 34) ................................................................................................... 33 - Abdomen yellow with varying amounts of brown on tergites 3–5, tergite 6 brown, tergite 7 yellow ( Figs. 1, 8, 9), females not currently distinguishable beyond this point................................................................ 35 33. (32) Scutum mostly yellow with prealar brown spots (sometimes united as a band) or brown Y-shaped medial marking; abdomen with tergite 7 yellow ( Figs. 22, 34)................................................................... 34 - Scutum mostly dark brown with yellow restricted to anterior and lateral margins; abdomen with tergite 7 brown ( Fig. 23); genitalia with tergite 9 bearing ventrally-directed fold along most of posterior margin and small point on apicomedial corner ( Fig. 109); female unknown ( South Africa, Fig. 134)......................................... L. mandelaiBorkent, n. sp. 34. (33) Scutum with brown Y-shaped medial marking and prealar brown spots ( Fig. 81); genitalia with tergite 9 ending in square lobe bearing small medial point; sternite 9 more or less oval shaped ( Fig. 115) ( Tanzania, Fig. 134).................................................................................................... L. stigmatusBorkent, n. sp. - Scutum with prealar brown spots or band only ( Fig. 22); genitalia with lateral lobes of tergal evagination gradually tapering to a medial point; sternite 9 a posteriorly directed triangle ( Fig. 110) ( Guineaand Nigeria, Fig. 136)........ L. medleriMatile. 35. (32) Tergite 9 with basal 2/3 of lateral margins parallel; ventrally directed process at base of posterior lobe of tergite 9 forked into two points at apex; apex of posterior lobe blunt and slightly bulbous ( Fig. 91) (central and western Africa, Fig. 135)........................................................................................... L. aliciaeMatile. - Tergite 9 with basal 2/3 of lateral margins bulbous tapering posteriorly; ventrally directed process at base of posterior lobe of tergite 9 a single point. ( Figs. 96, 97)..................................................................... 36 36. (35) Concavity of posterior margin of tergite 9 longer in diameter than width of sternite 9; posterior lobes of tergite 9 tapering to blunt points ( Fig. 97) ( Ivory Coastand Ghana, Fig. 135)...................................... L. couturieriMatile. - Concavity of posterior margin of tergite 9 smaller in diameter than width of sternite 9; posterior lobes of tergite 9 tapering first to create a lateral corner and then tapering to a point on medial margin ( Fig. 96) ( Republic of the Congoand Democratic Republic of the Congo, Fig. 135)......................................................... L. carnevaleiMatile.
R
northern Borneo
11
12
1
holotype
Malaysia
Peninsular
11
12
1