Review of the filter-feeding caddisfly subfamily Macronematinae (Trichoptera: Hydropsychidae) in tropical Southeast Asia Uy, Christine Jewel C. Malicky, Hans Bae, Yeon Jae Raffles Bulletin of Zoology 2018 2018-11-15 66 664 703 628L5 Ulmer 1905 [377,571,1853,1877] Insecta Hydropsychidae Animalia Trichoptera 35 665 Arthropoda subFamily Macronematinae    1. Without mouthparts.................................................................2  – With mouthparts ..................................................................12   2. Forewing with ‘false’ discoidal cell formed by fusion of R4 and R5 after basal furcation ( Fig. 1) ........................................ ................................  Aethaloptera(  Aethaloptera sexpunctata)  – Forewing without ‘false’ discoidal cell.................................3   3. Forewing without discoidal cell; Sc and R1 fused ( Fig. 177); thorax without any brown spot markings................................. .......................................  Oestropsyche(  Oestropsyche vitrina)  – Forewing with discoidal cell; Sc and R1 not fused ( Figs. 193– 210); thorax with or without markings, either on mesoscutum or mesoscutellum.......................................  Polymorphanisus4   4. Without markings on mesoscutum and mesoscutellum ( Fig. 183)..........................................................................................5  – With markings on either mesoscutum or mesoscutellum ( Figs. 184–192) .................................................................................6   5. With two brown spots on forewing, one on median cell and one on discoidal cell ( Fig. 194); male eyes large almost meeting ventrally; segment IX of male genitalia squarish apex dorsally ( Fig. 230) .......................................  Polymorphanisus ocularis  – Without markings on forewing; eyes small, not meeting ventrally; segment IX of male genitalia produced into a spatulate lobe dorsally ( Fig. 226....................  Polymorphanisus astictus   6. Without markings on mesoscutellum; with large dark markings on each side of mesoscutum and metascutum ( Fig. 188)........ ...........................................  Polymorphanisus quadripunctatus  – With markings on mesoscutellum; mesoscutum with or without markings, metascutum without markings ...............................7   7. With only one oval spot at center of mesoscutellum (occupying whole mesoscutellum for female ( Fig. 185) and covering only anterior third for male ( Fig. 184); forewing with two brown spots at the basal portion ( Fig. 194)  Polymorphanisus fuscus  – With paired spots or markings on mesoscutellum; forewing without brown spots ................................................................8   8. Paired spots or markings of mesoscutellum situated near center .......................................................................................9  – Paired spots or markings each situated near edge of mesoscutellum.......................................................................10   9. Each of paired markings with darker area concentrated at anterior and posterior portion as shown ( Fig. 186); segment X of male genitalia divided medially, each half with double lobe ( Fig. 228)................................ olymorphanisus muluensis  – Paired spots small ( Fig. 192); segment X of male genitalia divided medially with each lobe outer portion longer than the inner side ( Fig. 234)...................  Polymorphanisus unipunctus   10. With two large oval spots on mesoscutellum located on each side covering most of the area for female ( Fig. 191) and smaller oval markings with darker areas concentrated on anterior and posterior ends for male ( Fig. 190)............................................ ....................................................  Polymorphanisus scutellatus  – With smaller oval spots on mesoscutellum for both male and female ....................................................................................11   11. Mesoscutellum spots at each side elongated ( Fig. 189); segment X in dorsal view crown like in shape, divided medially with each half having longer inner lobe than the outer ( Fig. 232). .........................................................  Polymorphanisus semperi  – Mesoscutellum spots located at each side circular ( Fig. 187); segment X in dorsal view divided medially with each half bilobed ( Fig. 229).......................  Polymorphanisus nigricornis   12. Forewing without discoidal cell, anal area dilated in males ( Figs. 8, 12, 17, 26) ......................................  Amphipsyche13  – Forewing with discoidal cell, sometimes very small...........19   13. Foretibia without spur ...........................................................14  – Foretibia with spur ................................................................16   14. Male phallotheca with three pairs of endothecal spines ( Figs. 25, 33)....................................................................................15  – Male phallotheca with only 2 pairs of endothecal spines, mid and ventral spines short, dorsal endothecal spine absent ( Figs. 29, 30).................................................  Amphipsyche meridiana   15. Very large species 22–25 mm; with pair of round markings on mesoscutellum ( Fig. 23) ....................  Amphipsyche magna  – Small species 10 – 22 mm; without markings on mesoscutellum ...........................................  Amphipsyche parva   16. Fork II on forewing stalked ( Fig. 12); inferior appendages slender....................................................................................17  – Fork II on forewing not stalked ( Fig. 8); coxopodite very much broader than harpago ( Fig. 9) ............  Amphipsyche bifasciata   17. Male phallus with everted endotheca, connective membrane bearing many small spines ( Figs. 15, 16) ................................ ..............................................................  Amphipsyche exsiliens  – Male phallus without everted endotheca ..............................18   18. Forewing pattern with five dark brown spots and other paler brown markings ( Fig. 17); male phallotheca with large dorsal leaf-like lobes ( Figs. 20, 21).................  Amphipsyche gratiosa  – Forewing with only one dark brown spot in fork I; male phallotheca without dorsal leaf-like lobes ( Figs. 35, 36) ........ ..............................................................  Amphipsyche petiolata   19. R1 on hind wing fused with Sc ( Fig. 172); segment X of male genitalia bilobed with warts and clusters of setae, each lobe elongated ( Figs. 49, 92, 128, 77).................  Macrostemum27  – R1 on hind wing ending on R2+3, joined to Sc with a short cross-vein ( Fig. 240); segment X bilobed, lobes not as longer as  Macrostemumsp.( Figs. 237, 260, 265) ..........................20   20. Species size ranging from 15–24 mm; forewing with base of Rs obsolete, joined to R1 by cross-vein; phallus apex without complicated structures...........................  Trichomacronema21  – Smaller species 10–12 mm; forewing base of Rs entire; phallus with more complicated structures in apex than ....................... ................................................................  Pseudoleptonema23   21. Species 15–22 mm; forewing black with white marking rectangular and V-shape markings ( Figs. 258, 263)............22  – Species very large 22–24 mm; forewing lighter with white markings different with previous ( Fig. 268) (only female known) .............  Trichomacronema vietnamensis new species   22. Phallus apex rounded in lateral view ( Fig. 261)...................... .........................................................  Trichomacronema paniae  – Phallus apex oval and broad in lateral view ( Fig. 266)........... ........................................................  Trichomacronema tamdao   23. Phallus having circular apex with tongue-like projection from base pointing apically ( Fig. 254)..........................................24  – Phallus having circular apex without tongue-like projection .. ...............................................................................................25   24. Body black and may appear dark brown in alcohol; forewing with black and white bands ( Fig. 251)..................................... .........................................  Pseudoleptonema quinquefasciatum  – Body yellowish brown; forewing yellowish brown and black with white bands similar to  P. quinquefasciatumbut with large transparent region subapically ( Fig. 257)................................. ..............................................  Pseudoleptonema tansoongnerni   25. Phallus with curved elongated apex ( Fig. 238); segment X widely separated medially in dorsal view forming a U-shaped opening at the middle ( Fig. 237)....  Pseudoleptonema erawan  – Phallus with rounded apex ( Figs. 243, 248); segment X not as widely separated medially as above ( Figs. 242, 247)..........26   26. Segment X with rounded setal area at tip in dorsal view ( Fig. 241); forewing with white circular and long V-shape white markings, apex evenly curved ( Fig. 240)................................. ......................................................  Pseudoleptonema sinuatum  – Segment X without setal area at tip ( Fig. 246); forewing with a small V-shape marking, apex with anterior half longer than posterior half ( Fig. 245)..................  Pseudoleptonema supalak   27. Forewing half dark brown, half pale ( Fig. 37); head is yellow, unmarked except the silvery mark near eyes...........................  Macrostemum albardanum  – Forewing marking not as above; head usually dark brown to yellowish with dark brown bands.........................................28   28. Forewing without markings; phallus apex squaris.................9  – Forewing with evident dark brown and light markings; phallus not squarish ..........................................................................30    29. Segment Xwithout bifid protrusion at apex ( Fig. 138); phallus with tongue-like protrusion in dorsal view ( Fig. 141); distributed in Sabah, Borneo ..................................  Macrostemum pallipes  – Segment X with bifid protrusion at apex in lateral view ( Fig. 173); phallus without protrusion in dorsal view ( Fig 176); distributed in the Philippines.......................  Macrostemumsp.   30. Species yellowish; with dark brown band markings in the middle of the forewing running through the entire width ( Figs. 90, 126)..................................................................................31  – Species either dark brown or light brown, or if species is yellowish, without forewing markings as above..................32   31. Dark brown stripe along middle of the width of forewing somewhat straight, anterior band thicker and narrowing posteriorly until the arculus and continuing with a dot like dark brown spot ( Fig. 90) .................  Macrostemum fastosum  – Dark brown stripe along middle of the width of forewing curving anteriorly ( Fig. 126) .................  Macrostemum midas   32. Forewing with orange, white and black markings ( Fig. 54); segment X with each lobes longer as compared to the male genitalia of other  Macrostemumspecies( Fig. 56); species endemic in the Philippines...............  Macrostemum boettcheri  – Forewing not as above; segment X not as long as above; species can also be found in other geographic area.........................33   33. Species pale yellowish; forewing with dark brown spots ...34  – Species dark brown ...............................................................38   34. Forewing with three dark brown bands: one near base not reaching costa, one at middle formed by four connected spots, and one broad band almost covering apex ( Fig. 165) ............. .......................................................  Macrostemum trifasciatum  – Forewing with dark brown spots instead of bands ( Figs. 42, 149, 142–144)........................................................................35   35. Forewing with three dark brown spots or less; segment IX of male genitalia usually broad ................................................36  – Forewing with five dark brown spots or more ( Fig. 149); segment IX of male genitalia thin and annular ( Fig. 151) ...... .............................................  Macrostemum quinquepunctatum   36. Forewing with two or no dark brown spot ( Figs. 142–144); phallus in lateral view with longer posterior portion of the apex ( Fig. 147) making it having an opening anteriorly ......... ..........................................................  Macrostemum punctatum  – Forewing with three dark brown spots; phallus in lateral view if with posterior portion, making it just like a protrusion and not an opening.......................................................................37   37. Forewing with three dark brown spots located on the radial sector, cubital area near base, and anal area almost near the posterior edge of forewing ( Fig. 42) ...  Macrostemum bacham  – Forewing with three dark brown spots located on crossvein behind median cell, in front of anal vein, and on vein from median to cubitus near base.........  Macrostemum tripunctatum   38. Endemic to the Philippines...................................................39  – Distributed in other area .......................................................40   39. Forewing darker apically and lighter basally, with two curved light bands divided in the middle, with lighter bands running toward the length of forewing occupying three-fourths of the length, and four lighter streak bands near the apex ( Fig. 59). ............................................................  Macrostemum caliptera  – Forewing dark brown, with several silvery white marks as a subapical streak, two coastal spots tending to form V-shape, two spots near the base, and two streaks located toward the posterior portion of the forewing ( Fig. 52) .............................. ................................................................  Macrostemum bellum   40. Phallus apex divided into two lobes ( Figs. 68, 117, 163) ...41  – Phallus not divided into two lobes .......................................43   41. Phallus apex in lateral view opening widely forming a U-shape ( Figs. 68, 163), in ventral view, apex round; segment X medially separated and joined at the center forming a V-shape area in dorsal view ( Figs. 67, 162) ...................................................42  – Phallus opening V-shaped and not as deep as wide ( Fig. 117), in ventral view, apex heart-shaped ( Fig. 118); segment X medially separated with lobes directed downwards ( Figs. 114, 116)..........................................................  Macrostemum hestia   42. Phallus opening broad with each lobe thick ( Fig. 163); forewing with white markings where one is C-shape occupying almost half of anterior area ( Fig. 160).......  Macrostemum tonkinensis  – Phallus opening narrow, twice as deep as wide ( Fig. 68); forewing with white markings as shown ( Fig. 65).................. .............................................................  Macrostemum dairiana   43. Phallus with spine-like structure at the tip of apex ( Figs. 73, 105)........................................................................................44  – Phallus without spine-like structure at the tip of apex.......45   44. Segment X with apical lobes separated by parallel-sided gap narrower than width of either lobe ( Fig. 72); spine-like structure at the tip of phallus seems not pointed in dorsal view ( Fig. 74), but obviously pointed in lateral view ( Fig. 73) ....................... ..................................................................  Macrostemum dione  – Segment X with apical lobes separated by U-shaped gap ( Figs. 103, 104); spine-like structure at the tip of the phallus pointed in dorsal view ( Fig. 106) ..............  Macrostemum fenestratum   45. Phallus with protrusion almost in the middle of the apex in lateral view ( Figs. 63, 86).....................................................46  – Phallus without protrusion, or if with protrusion at apex, concentrated on posterior area..............................................47   46. Phallus with curved protrusion very obvious in ventral view ( Fig. 89); forewing black in fresh specimens with white markings, three large across costa, other marking originating from arculus running upwards across the width of forewing ( Fig. 85) ........ ............................................................  Macrostemum eleanora  – Phallus without protrusion in dorsal view; forewing with three white markings, two across costa and one originating from arculus ( Fig. 60) ...............................  Macrostemum centrotum   47. Phallus strongly convex subapicoventrally with apicoventral protrusion 90° from ventral margin basal of convexity in lateral view ( Figs. 78, 50) ................................................................48  – Phallus rounded to an angle less than 90° in lateral view ..50   48. Phallus in dorsal view with apex oval narrowing in the body and with broad base ( Fig. 51); segment X in lateral view narrowing toward apex ( Fig. 48); forewing pattern dark brown in alcohol with white bands as shown in Fig. 47...................................... .......................................................  Macrostemum bellerophon  – Phallus in dorsal view with rounded apex; segment X in lateral view narrowing toward the apex but with broader tips with presence of warts ( Fig. 76); forewing pattern with three white bands running across the length of the forewing as  M. bellerophon,but no white markings running towards the length of the forewing length, or if there is very tiny white streak ( Figs. 75, 170) .......................................................................49   49. Phallus in dorsal view with raised curve protrusion at apex ( Fig. 79); forewing pattern with two white bands in anterior and one white band in posterior and one small streak band running toward forewing length ( Fig. 75), but in some individuals, the small white streak is not evident .............................................. ...................................................  Macrostemum distinguendum  – Phallus in dorsal view rounded; forewing pattern with four white bands running across the length of the forewing, two anterior and two posterior ( Fig. 170) ....  Macrostemum zenon   50. Phallus apex with ventral area much more protruded than anterior ( Fig. 125); segment X lobes separated only half of length of segment X by narrow V-shaped notch ( Fig. 124) ... ..............................................................  Macrostemum luteipes  – Phallus apex not as above, if with protrusion, not as much protruded as above; segment X usually widely separated or if not, the lobes are not as attached as above..........................51   51. Phallus with truncate apex, in dorsal view as broad basally as apically, slightly narrower in middle ( Fig. 165); forewing with variation ( Figs. 131, 132).................  Macrostemum opulentum  – Phallus rounded; forewing pattern mostly black or brown with fewer pale areas that are not interconnected........................52   52. Forewing with white markings as shown in Fig. 53, most prominent large triangular marking at the middle part............ .....................................................  Macrostemum bifenestratum  – Forewing pattern with various white markings, but none dominating middle of wing...................................................53   53. Phallus apex with small tongue like projection in lateral view ( Figs. 83, 110) .......................................................................54  – Phallus without small projection at apex .............................55   54. Phallus in dorsal view with apex as wide as base and twice as wide as middle ( Fig. 84); forewing with four white markings, two triangular across costa, one linear running parallel to the length of forewing, and one circular near basal area ( Fig. 80); segment X divided mesally by narrow V-shaped pad, with each side consisting of two lobes, one is smaller and located at anterior of major lobe ( Fig. 82).........  Macrostemum dohrni  – Phallus in dorsal view with apical portion twice as wide as basal portion ( Fig. 111); forewing with more than four white markings ( Fig. 107); segment X bilobed and each lobe widely separated in dorsal view ( Fig. 109.....  Macrostemum floridum   55. Phallus with dorsal part of apex more protruded ( Fig. 158) in lateral view, protrusion very much obvious in dorsal view ( Fig. 159); segment X with each lobes widely separated forming a U-shape space ( Fig. 157); forewing with variation ( Fig. 154, 155)............................................................  Macrostemum seba  – Phallus with dorsal part of apex flat, ventral part a little longer in lateral view ( Fig. 122); segment X with each lobe flat at apex ( Fig. 121); forewing pattern as shown in Fig. 119......... .......................................................  Macrostemum indistinctum Segment X Sabah 36 700 1 Sabah