Neoperla africana-group

IV. The N. africana-group

The N. africana- group includes many African species but also a few Asian ones. The N. africana -group is distinguished by three apomorphies but contradicted by the very strongly supported node C:

1. Caudal process of male T7 three-dimensional (e.g., Figs. 102–103 View FIGURES 102–107 , 169–170 View FIGURES 169–176 ). The anterior half of T7 forms a continuous sclerotised ring in front of a pale transverse pale line or a short weakly sclerotised zone. The soft caudal half of T7 accomodates an approximately triangular sclerite ending in a pyramidal process with SB near its tip and along a ridge on the underside. T8 is flat to slightly convex, no raised hump or process. Penis tube sclerotised, dorsal and ventral faces of endophallus differ and are (except in N. planidorsum ) separated by bare lateral strips. Most spines stand on the convex side, details vary (e.g., Figs. 164 View FIGURES 161–168 , 203 View FIGURES 201–204 , 317 View FIGURES 313–322 , 376 View FIGURES 373–385 , 440 View FIGURES 439–442 ).

2. Females with a flat, band-shaped SSt which is strongly curved or coiled, details vary. If short, the SSt resembles a section cut out of a circle (e.g., Figs. 124 View FIGURES 123–126 , 167 View FIGURES 161–168 , 224 View FIGURES 223–226 ) but in most species it forms a full circle or several superimposed rings which may be drawn out into a long spiral (e.g., Figs. 123, 126 View FIGURES 123–126 ). The winding or spiral spermatheca lies in the center of the coil and is not always visible. Female S8 without structural modifications but often with three brown maculae in triangular arrangement. The pattern is variable, the maculae may be large and merge more or less around a pale center which is often anchor-shaped.

3. Pigmented scales form an almost gap-less coat on the wall of the SSt, only the concave edge is free of them. Scales are obliquely rooted in the cuticle, with tips directed towards the vagina (e.g., Fig. 108 View FIGURES 108–113 ). The basal region near the attachment of the SSt to the vagina seems to be extensible and contains many longitudinal folds. In this basal section scales are restricted to near the convex edge, distally scales occupy the entire width of the coil.

External genitalia alone rarely permit species identifications, usually inner genital structures and eggs must also be examined. The penis is a sclerotised tube, only a small ventrobasal region is soft (e.g., Figs. 117 View FIGURES 114–122 , 145 View FIGURES 143–149 ). The endophallus tends to be longer than the penis tube and is folded up inside. The endophallic armature is not dispersed over the surface, instead bare sides separate a long dorsal area with various kinds of spines from a similar but shorter ventral area. Eggs vary between species complexes.

Keys to species in the N. africana View in CoL -group

Males (several unknown)

1 T8 flat, sometimes generally slightly convex but no hump or process............................................ 2

1' T8 with a distinct hump or a process.................................................... N. spio View in CoL -complex, p. 87

2 Region between penis tip and the endophallus base armed with spines........................................... 2

2' A long bare zone between penis tip and spines on endophallus (e.g., Fig. 164 View FIGURES 161–168 )............... N. pilulifera -complex, p. 70

3 Endophallus with slender hooks on all sides ( Figs. 104–107 View FIGURES 102–107 ).................................. 22 N. erinaceus n. sp.

3' Spines mainly on convex face of endophallus, fewer or none on concave face, laterally bare.......................... 4

4 On each side of the contracted penis tip a few spines form a lateral row. On the everted endophallus these spines stand before a background of flat scales and truncate spines forming a sleeve around the endophallus base and are therefore less easily seen. Beyond the sleeve spines and hooks occur all around the endophallus, more distally most of the ventral side is bare. Large hooks form the top dorsolateral spine row ( Figs. 116–118, 120–122 View FIGURES 114–122 ), other spines are smaller........... 23 N. africana Klapálek View in CoL

4' No lateral spine row at penis tip, no scaly pavement around base of endophallus, no row of large hooks along endophallus side, spines of fairly uniform size (e.g., Figs. 138 View FIGURES 135–142 , 143 View FIGURES 143–149 )............................................................ 5

5 Endophallus much longer than penis tube, convex side basally with one row of single widely separated spines (e.g., Figs. 143, 146 View FIGURES 143–149 )...................................................................... 25 N. camerunensis (Enderlein) View in CoL

5' Endophallus barely longer than penis tube, base of endophallus dorsolaterally with dense spine cover ( Figs. 137–142 View FIGURES 135–142 )...................................................................................... 24 N. panafricana n. sp.

Females (unknown of N. erinaceus )

1 Eggs with about a dozen straight ridges, chorion surface similar to the wide concave interstices....................... 2

1' Eggs with approximately 20 or more proper striae, costae with smooth surface, the narrow sulci with unordered punctation................................................................................... N. spio- complex, p. 87

2 Front wing distally with a transverse clear band between dark shades ( Fig. 96 View FIGURES 96–101 )..................... 20 N. decorata n. sp.

2' Front wing not patterned............................................................................... 3

3 Egg subspherical, not striate (e.g., Figs. 168 View FIGURES 161–168 , 187 View FIGURES 181–192 )..................................... N. pilulifera -complex, p. 70

3' Egg striate........................................................................................... 4

4 Egg with a low cap-like operculum and a visible eclosion line ( Fig. 100 View FIGURES 96–101 )........................ 21 N. cataractae n. sp.

4' No visible eclosion line, operculum different............................................................... 5

5 SSt long and flexible, forming 2 or more (up to ~5) rings ( Fig. 126 View FIGURES 123–126 )................... 25 N. camerunensis (Enderlein)

5' SSt forming a single ring, or even shorter, resembling part of a circle............................................ 6

6 SSt very wide, surrounding a small centre. Wide base of SSt on concave side bare and with prominent longitudinal folds. Convex side with a narrow spine band widening distally abruptly which causes a projecting angle. Wide base and narrow distal section of SSt of similar length ( Fig. 123 View FIGURES 123–126 ). Vagina without lateral sclerite rods.................. 23 N. africana Klapálek

6' SSt narrower and longer, about one ring. Base bare on concave side, spine band on convex side widening gradually, rarely an indistinct angle. The completely scaly distal part of the RC equals 1/3 of total length (e.g., Fig. 124 View FIGURES 123–126 ). Vagina calyx-shaped, with sinuous lateral rods................................................................. 24 N. panafricana n. sp.

IV.1 The N. africana -complex (= clades D, E)

Clade D

The monophyly of this small lineage with only three species is only moderately well supported (73.9/98/0) in the DNA sequence data analyses, which is in line with only COX1 DNA barcode sequences being available ( Figs. 492–493 View FIGURE 492 View FIGURE 493 ).