V. The Neoperla excisa and N. sjostedti -operational complex (~ clade J, clade K + L)

Species in this operational complex were from the beginning of our work distinguished from all other species groups because:

1. known males have no unpaired medio-caudal process of T7. In many T7 is instead concave between a pair of paramedian humps,

2. females with S8 with a projecting hard nail or with a sclerite resembling a fingernail occur only in the present assemblage, and

3. egg striae have narrow sulci which lack macroscopic punctures but have micropunctures visible only in transmitted light, at magnifications ≥ 200*. They form a single regular row on each side of a sulcus and separate it from the costa. Many eggs have spiral striae with species-specific direction of spin while most other species have straight or at best oblique striae.

Unfortunately, half of the included taxa are known in only one sex, morphology did not reveal how the species are interrelated.Accordingly, the keys which rely on morphology cover the entire operational complex.Arrangement of texts follows the strong molecular evidence (except for N. multiserrata sp. n. and the exclusion of node I).

Keys to species

Males

1 T7 is flat, a trapezoidal sclerite in the middle has a wide truncate caudal edge which is opposed to and only narrowly separated from the wide, transverse top of a high process on T8 (Figs. 373, 386)........................................... 2

1' T7 and T8 different, the high process on T8 with narrow top, or hook-like........................................ 3

2 Tips of HT10 bisinuous, the hook-like tips often grasp and hold the T8 process; mediobasal callus parabolic (Fig. 373). Penis dorsally with subterminal and terminal spine groups (Figs. 374–375, 385). Endophallus dorsally with several rows of spines, ventrally with a single orderly spine row (Figs. 376, 380)....................... 65 N. heideae and 66 N. larvata n. spp.

2' Tip of HT process not hook-shaped (Fig. 386). Penis with only one subterminal spine group. Endophallus dorsally with a wide spine field, ventral face bare (Fig. 387)...................................................... 67 N. sorella n. sp.

3 T7 with a strongly raised wide caudal process, its hind corners diverge (Figs. 365, 366, 486).......................... 4

3' T7 without caudal process, concave between the blunt corners of a caudal bench-like elevation (Fig. 422), or between 2 low paramedian humps. T8 and penis variable.................................................................. 5

4 T8 with wide, forward-curved process which fits into the excavation of the process on T7 (Figs. 365, 388). The penis is a hard narrow long tube, the long narrow endophallus with fine short armature (Fig. 368)................. 64 N. excisa Klapálek

4‘ Process of T8 narrow, pointed, standing erect behind the bilobed projection on T7. Penis soft and short, endophallus with sharp spreading spines (Figs. 486–489)............................................................. 83 N. sp. Afr_A

5 Penis tip bent like a knee, only part of the short endophallus with spines, several form a short crest on the ventral side (Figs. 423–424)........................................................................... 74 N. crenulata n. sp.

5' Penis straight, distally annulated, the endophallus differs...................................................... 6

6 Endophallus with a dorsoventral armature of long groups or rows of spines (e.g., Figs. 440, 445)...................... 7

6‘ Endophallic armature dominated by basolateral rows of individual spines (Figs. 482, 484–485), dorsal and ventral faces are bare or have only fine spines............................................................................ 9

7 Penis straight, its annulated tip soft. Spinose part of endophallus not much longer than the tube, the wide basal half dorsally with a wide spine field, ventrally with a row or a narrow band of many small spines, the long and narrow distal section is bare (Fig. 440).............................................................................. 77 N. socia n. sp.

7' Entire penis tube moderately sclerotised. The spiny section of the endophallus is at least twice longer than the penis (e. g., Figs. 445, 452)............................................................................................ 8

8 Penis ventrally with an empty caecum as long as the penis is wide, or longer (Figs. 451–454)................................................................................................. 78A N. sjostedti needhami Lestage

8‘ No (Fig. 446) or a rudimentary caecum on the ventral side of penis, or only an oblique inconspicuous fold in its place (Fig. 445).................................................................... 78B N. sjostedti sjostedti Klapálek

9 Top of T8-process wide, rounded or truncate (Figs. 477, 479, 481), even slightly concave under certain angles of view. Base of endophallus on each side with a short row of rapidly decreasing sharp teeth, distal part of endophallus consisting of long folded membranes without distinct armature.................................................. 82 N. leroiana Klapálek

9' Contour of T8-process triangular, top more or less pointed (Figs. 432,461), endophallus variable..................... 10

10 Endophallus (observed only by transparency) little longer than the penis, two orderly lateral rows of stout conical teeth of uniform size occupy the basal 2/3 (Figs. 434–435), no other armature present...................... 76 N. tangana n. sp.

10' Endophallus much longer than the penis tube, endophallus base on each side with an orderly row of individual large teeth which distally decrease rapidly in size......................................................................... 11

11 Basolateral rows of endophallic teeth continue for a considerable distance, much smaller teeth stand between them (Figs. 460, 462).......................................................................... 80 N. arambourgana Navás ’

11’ Penis similar to N. leroiana, (i.e., the basolateral rows of teeth on endophallus are very short, comprising only about 12 sharp teeth (Fig. 485)), the long distal part of endophallus composed of folded membranes without recognizable armature.......................................................................................... 81 N. dundoana n. sp.

Females

At the base of the SSt is usually a section of variable length that can presumably be much expanded, it has scales only along the convex face while the concave side is membranous, looks void but may have longitudinal folds. This basal section is not considered in statements like "entirely" or "completely coated with scales".

1 Caudal edge of S8 with a long nail (e.g., Figs 369, 378, 382). SSt long........................................... 2

1' Caudal edge of S8 and SSt variable, possible nails are short.................................................... 4

2 The SSt is of similar width over its entire length (Fig. 369). Egg ovoid, without distinct striae (Figs. 370–372).............................................................................................. 64 N. excisa Klapálek

2' SSt with wide base, the longer distal part narrow (Figs. 377–378). Egg drum-shaped, with levogyrous striae (Fig. 379)..... 3

3 Nail tongue-shaped. SSt forms about 1 ring, entirely coated with scales, entirely of the same structure and directly attached to the curled spermatheca (Figs. 377–378)..................................................... 65 N. heideae n. sp.

3' Nail conical, its base may be narrow or wide. SSt very long, consisting of an entirely scaly basal half and a very narrow distal part with scattered very small cuticular elements which is connected to the spermatheca (Figs. 381–383)..................................................................................................... 66 N. larvata n. sp.

4 Caudal edge of S8 triangularly recessed, a short conical nail projects from the notch. Floor of vagina sclerotised, a massive inversely U-shaped crest meets an internal ridge along the antecosta 8. Egg with levogyrous spiral striae (Figs. 397–399)......................................................................................... 70 N. dianae n. sp.

4' Different, eggs and S8 variable.......................................................................... 5

5 The center of the caudal edge of S8 with a distinct notch...................................................... 6

5' Caudal edge of S8 not notched, variable................................................................... 8

6 The bare notch is shallow but wide, from it a strong sclerite extends forward into the vagina. Egg drum-shaped, striae straight (Figs. 418–419)....................................................................... 73 N. excavata n. sp.

6' Notch of S8 and egg different............................................................................ 7

7 Notch rounded, strong setae reach over it from in front. Egg slender, striae levogyrous, costae near the punctate operculum with pale swellings (Figs. 402–404)............................................................ 71 N. gibbosa n. sp.

7' Caudal edge of S8 with two soft points flanking a crescent-shaped sclerite which is shifted a little forward towards the vagina (Figs. 463–465). Egg slender, ovoid, with levogyrous striae (Fig. 466)..................... 80 N. arambourgana Navás

8 S8 caudally long, parabolic, with an anchor pattern. SSt long, egg sulci spiral and very narrow........................ 9

8' S8 caudally truncate, not prolonged, SSt and egg variable.................................................... 10

9 Egg oval, the levogyrous striae continue onto parabolic operculum (Figs. 362–363)................. 63 N. muyukae n. sp.

9' Egg drum shaped, operculum wide, smooth, sharply delimited against the dextrogyrous striae (Figs. 357–358).......................................................................................... 62 N. conradti (Enderlein)

10 S8 with a pale anchor pattern inside a brown macula........................................................ 11

10' S8 without anchor pattern, variable...................................................................... 14

11 Egg with>35 straight striae, a pale ring at base of operculum. Micropyles on the bottom of very narrow sulci. Anchor with single stem (Figs. 345, 347, 349).......................................................... 61 N. lineata n. sp.

11' Egg without pale ring, with fewer variable striae and with modified anchors: the anchor cap is supported by several delicate filaments instead of one stem........................................................................... 12

12 Egg striae are straight (Fig. 302)....................................................... 52 N. orthonema n. sp.

12' Egg striae are dextrogyrous............................................................................ 13

13 The micropyles are located in the narrow deep sulci (Figs. 306–308).......................... 53 N. filamentosa n. sp.

13' The micropyles are located on top of narrow supplementary costae (Fig. 312)................... 54 N. spironema n. sp.

14 S8 caudally with a short subgenital plate, or a projecting nail, or a sclerite resembling a fingernail.................... 15

14' S8 not distinctly modified, the SSt is of uniform width, approximately1.5-2* longer than the vagina................... 22

15 Caudal edge of S8 with short rounded projection that is a little thickened. The ribbon-like SSt is several times longer than vagina and S8 and appears void for almost half the length because there are only few scales along the convex edge. The elongate egg has wide blunt poles and straight striae.............................................. 78 N. sjostedti Klapálek

15' S8 not as above, the SSt is not nearly as long and has no strikingly long void basal section.......................... 16

16 SSt little longer than the vagina, egg not striate............................................... 68 N. simplex n. sp.

16' SSt 2-3 times than vagina and SSt, the egg is striate......................................................... 17

17 Compared to the vagina the SSt is wide and narrows caudally much. The straight egg costae are drawn out into high crests (Figs. 441–442)............................................................................... 77 N. socia n. sp.

17' Compared to the vagina the SSt is narrow; it is of uniform width............................................... 18

18 Costae with knobs and little columns in alternating positions.................................. 74 N. crenulata n. sp.

18' The egg costae are unmodified, flat...................................................................... 19

19 The nail on S8 projects and is entire or distally minimally excised (Figs. 470, 472–473)............................ 20

19' Nail on S8 notched, variable (Figs, 463–465, 468–469)...................................................... 21

20 The slender egg has a distinct collar and a conical operculum, the egg striae are levogyrous spirals...................................................................................................... 82 N. leroiana Klapálek

20' The egg resembles the egg of N. sjostedti (Fig. 456) and has straight striae, the SSt is only moderately long and forms ~2 rings, there are no long void basal sections (Fig. 457)................................ 79 N. sjostedti cf. needhami (by DNA)

21 Caudal edge of S8 curved forward into the excavate nail which is S8 shifted forward; egg plump, the wide operculum with small grooves, striae are straight (Figs. 467–469)........................................... 81 N. dundoana n. sp.

21' Caudal edge of S9 with a deep median notch in front of which lies a crescent-shaped sclerite. The ovoid egg with narrow operculum and oblique striae (Figs. 463–466)......................................... 80 N. arambourgana Navás

22 The eggs have straight striae........................................................................... 23

22' The eggs have levogyrous striae......................................................................... 24

23 Anchor pole of egg with a collar, the costae are asperous, like sand paper (Figs. 437–438)............ 76 N. tangana n. sp.

23' The egg lacks a collar but has in its place a long projection; the costae are smooth (Figs. 394–396)......6 9 N. claviger n. sp.

24 SSt almost twice longer than the vagina, wide, basal third with longitudinal folds, only a band of scales along the convex edge. The egg striae include the operculum and have extremely narrow sulci which do not widen around micropyles (Figs. 429–431)........................................................................ 75 N. massevensis n. sp.

24' S8 unmodified, the SSt is shorter, thin, and basally not modified. The egg sulci are widened around micropyles (Figs. 340– 343).............................................................................. 60 N. bareensis n. sp.