Neoperla juxtadidita, Zwick & Zwick, 2023

Zwick, Peter & Zwick, Andreas, 2023, Revision of the African Neoperla Needham, 1905 (Plecoptera: Perlidae: Perlinae) based on morphological and molecular data, Zootaxa 5316 (1), pp. 1-194 : 18-19

10 SSt forming 1–2 rings, inside with needle-like microtrichia ( Fig. 109 ). Egg slender, spindle-shaped, both poles narrow, no collar, with ~ 20 straight ridges ( Figs. 83–90 )..................... 17 N. didita (Enderlein) and 18 N. juxtadidita n. sp.

10‘ Inside of SSt with dense coat of flat scales ( Fig. 113 ), length varies from a short regularly curved fraction of a ring to long spirals of up to 5 superimposed rings, or very long narrow tubes forming a tangle ( Figs 126 , 167 , 335 )................. 11

11 The entire egg surface is punctate (e.g., Fig. 112 ), with 6–16 straight ridges (e.g., Figs. 100–101 ). Some species with coarse punctation have ornaments on top of ridges (e.g., Fig. 111 )................................. N. africana -group, p. 50

11' Impunctate costae or smooth ridges present on the chorion.................................................... 12

12 Egg striae are composed of impunctate raised costae and low-lying or incised punctate sulci (e.g., Figs. 110 , 214 , 291 , 401 ). Anchor mushroom shaped, a solid stem supports the cap (e.g., Figs. 7 , 31 , 392)................................... 13

12' Egg striation is caused by only raised ridges of variable height ( Figs. 303 , 312 ). Anchor resembling a mushroom but the cap rests on several thin delicate threads ( Figs 302 , 308 ), no solid stem present................ N. orthonema -complex, p. 116

13 Costae and sulci are of similar width, sulci with unordered punctation and freely visible micropyles (e.g., Fig. 257 ), except in

3 In cases of doubt, continue with the alternative couplet 1’ species where costae bear crest of flanges ( Figs. 324–325 )................ N. spio - and N. dubia -complexes, p. 87, p. 130

13' Costae are 2 or more times wider than the sulci which lack macroscopic punctures but have a row of micropunctures on each side (e.g., Figs. 343 , 350 , 360 ). Micropyles are not exposed, in some species they are completely concealed............ 14

14 S8 with a pale anchor-shaped mark, egg without collar....................................................... 15

14' S8 without pattern, caudally not modified. Egg shape variable, with or without collar, striae straight or levogyrous (anticlockwise, spirals in side view running from top left to bottom right). In mounted eggs regular rows of micropunctures are visible at magnifications ≥ 200X. Micropyles variable............... N. excisa and N. sjostedti -operational complex, part, p. 130

Figures

FIGURES 108–113. Neoperla spp., structural details.108, N. africana Klap., 109, N. kalengonis n. sp., 110, N. sjostedti needhami Lestage, all from the stream Kalengo: 108, 109 scales in the SSt. Close-ups of egg chorion: 110, three egg striae, the 2 rows of micropunctures in the sulci diverge around an island with a micropyle; 111, N. africana, 112 N. camerunensis (End.), chorion with 2 micropyle orifices; 113, processes of tergites 7 and 8, and modifications of antecosta 8 (AC8), a combination of inner structures (light grey), and of fine pilosity and sclerites (dark grey) on the surface of tergites; diagrammatical. The SEM microgaphs are courtesy of MPI of Limnology, PL̂n.

FIGURES83–90.Neoperladidita(Enderlein)andN.juxtadiditan.sp.:N.didita:83,spermathecalstalkofthefemalefromMamfe; 84,fragment ofan eggfromthe lectotype;85,eggof thespecimenfrom Mamfe;86,anchor poleofsame,opticalsectionandsurface view combined (diagrammatic, from a photograph). N.juxtadidita: 87, genital situs of holotype female from Belinga; 88, spinules from inside the spermathecal stalk; 89, egg; 90, anchor pole with anchor.Scales are 100µm, 84, 85 to the same, others not to scale. a, anchor; c, collar; r, ridges on egg; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 123–126. Neoperla africana-group, female genital situs and spermathecal stalk: N. africana Klapálek: 123, Ghana, Subri R., vagina and spermathecal coil folded back from S8, for better visibility of anchor spot on S8. N. panafricana n. sp.: 124, Waterberge, Mapote Farm. N. camerunensis (Enderlein): 125, Ethiopia, Ghibe R.; 126, Cameroon, Bamenda, NEOP128. Scales are 100µm. am, anchor mark; ap, angular projection; lf, longitudinal folds; lscl, lateral vaginal sclerite; tf, transverse folds; SSt, spermathecal stalk; V, vagina.

FIGURES 161–168. Neoperla amoena n. sp.. ♁ holotype: 161–163, habitus and abdominal tip (not to scale; NEOP070); 164–166, penis, only 164 to scale; ♀ paratype from Bamako: 167, 168, genital situs and egg. Scales are 100µm. bbs, bare basal stretch; tf, transverse folds; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 334–339. Neoperla spaghetti n. sp., ♀ holotype. 334, S8 with the anterior sclerites and the pale furrow (pf, arrow) between them. For better visibility the vagina (V) with the base of the spermathecal stalk (SSt) was flipped back. The overlong SSt (Fig. 335) broke in the process, a blue line connects the fitting ends of the fragments; 336, side view of an egg; 337, oblique view of an egg operculum; 338, anchor pole view of another egg, count of costae in white script; 339, detail of striae and micropyles, enlarged from 336. Scales are 100µm, 337–339 not to scale. co, costa; mi, micropyle; sa, spermatheca; SSt, spermathecal stalk; su, sulcus, v, vagina.

FIGURES 96–101. Neoperla decorata n. sp. and N. cataractae n. sp., the holotypes. N. decorata: 96, front wing; 97, genital situs, diagrammatic, spermatheca removed; 98, egg. N.cataractae: 99, female genital situs, diagrammatic, spermatheca removed; 100, 101, egg, side view and optical cross section (from a photograph). Scales are 100µm, other figures not to scale. a, anchor; c, collar; mi, micropyle; op, operculum; os, opercular suture; sd, spermathecal duct, SSt, spermathecal stalk, V vagina.

FIGURES 213–214. Neoperla burgeoni Navás: 213, ♀ genital situs, Central African Republic (NEOP213), the inset is the holotype of N. bredoana (stained with Safranin); 214, egg, Ghana 81, Banda-Nkwanta, 18–20.9.1965, lighttrap on savanna (Endr̂dy-Younga: 250). FIGURES 215–216. Neoperla beta n. sp. (Guinea, r. Milo at Kankan, slide Z16.26), female genital situs and egg. In Figs 213 and 215, the vagina was flipped back for better visibilty of S8. Situs and eggs to the same scales, respectively.

FIGURES 289–293. Neoperla usambara n. sp. and N. sambarua n. sp., ♀ ♀. N. usambara: 289, genital situs, vagina and the coiled spermathecal stalk flipped back to expose anchor pattern on S8; 290, egg; 291, detail of striae with a micropyle in a sulcus (full arrow). N. sambarua: 292, genital situs with diagrammatic indication of border between S7/S8; 293, egg, variable thickness of chorion indicated by inner contour on right hand side. Open arrows show divergent spinule bands at entry of SSt into vagina. Scales as indicated. A, anchor; c, collar; co, costa; ls, lateral sclerite; mi, micropyle; mic, micropylar canal; sa, spermatheca; SSt, spermathecal stalk.

FIGURES 397–401. Neoperla dianae n. sp., female: 397, genital situs (Libamba, Cameroon); 398, egg from Kumba, Cameroon, with variable thickness of chorion (reddish-brown), arrow points at abrupt thinning; 399, egg from Walikale, D. R. of the Congo; 400, anchor pole with collar and anchor and 401, detail of striae with 2 micropyles (from Libamba). Scale lines are 100µm, unless differently indicated; a, anchor; ac, anchor cavity; co, costa; cr, curved ridge; ct, crest; mi, micropyle; n, nail; sa, spermatheca; su, sulcus; SSt, spermathecal stalk; V, vagina.

FIGURES 6–10. Neoperla transvaalensis (Enderlein) (6, 7) and N. laticollis Klapálek (8–10): 6, female situs, specimen from Rwanda; 7, egg, semi-diagrammatic, not to scale; 8, female situs; 9, egg, lateral view; 10, enlarged detail of a costa. The scales are 100µm; a, anchor; c, collar; fo, stack of membranous folds; gp, gonopore; hk, hard knob, retractor apodeme; ls, lateral vaginal sclerite; mc, mediobasal callus of HT10, the hemitergite 10. mi, micropyle; op, operculum; sa, spermatheca; sd, spermathecal duct; S9, protruding tip of S9; sa, coiled spermatheca; V, vagina.

FIGURES 26–32.Neoperla kalengonisn. sp.: 26, artificiallyevertedpenis ofa paratype;27, spines on endophallus of same, scale is 50µm; 28, paratype ♀, genital situs; 29, 30, eggs of paratypes, 30 courtesy of MPI of Limnology; 31, egg from South Sudan; 32, anchorpoleandcollarofeggfromLwiroR.Scalesare100µmunlessdifferentlyindicated,30nottoscale,29and31tothesamescale. sa, spermatheca; SSt, spermathecal stalk.

FIGURES 294–304. Neoperla orthonema n. sp.: Male paratype from Lakota, Ivory Coast: 294, 295, abdominal tip; 296, everted penis, with detail from middle of endophallus; 297–299, apex of contracted penis, in dorsal, ventral and lateral views (from left); 300, lateral view of penis apex, beginning eversion (♁ from Kousankoro); 301, female genital situs (vagina flipped back to show the anchor pattern on S8) and 302, egg, Ankasa game prod. res., Ghana; 303, view onto edge of chorion fragment showing variable size of costae (Ivory Coast, Lakota); 304, detail of collar with end of striae (Wli Falls, Ghana). Scales are 100µm. Tergites, sternites and hemitergite 10 are labeled. d, dorsal spines; eb, spines at base of endophallus; sa, spermatheca; SSt, spermathecal stalk; V, vagina; vs, ventral spines.

FIGURES 309–312. Neoperla spironema n. sp.: 309, 310, holotype; 311, a paratype; both from Ankasa Game. Res; 312, SEMdetail of chorion with the 2 kinds of costae and a micropyle (Ivory Coast, USNM; from Moore, 1991). Scales are 100µm, or as indicated. mi, micropyle; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 305–308. Neoperla filamentosa, n. sp., holotype. 305, genital situs; 306, egg of HT; 307, detail of striae, arrows indicate micropyles. 308, egg of doubtful ♀ from Touba, Senegal. Scales are 100µm, eggs to same, details not to scale. Sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 250–257. Neoperla bella n. sp., 250, ♁ abdominal tip, dorsal view; 251, abdominal tip of ♁ holotype with naturally fully everted penis; 252, contracted penis; 253, penis tip and everted endophallus, 254, distal end of endophallus; 255, ♀ situs, vagina flipped back to expose pattern on S8; 256, slide mounted egg; 257, detail of chorion structure, arrow indicates a micropyle.All from Kalengo stream, D.R.Congo. Not all figures to scale, scales are 100µm, except as indicated in 257. Tergites, HT10, cercus and S8 are labeled. ea, external asperities on penis tube; es, endophallic spines; SSt, spermathecal stalk; V, vagina; vd, vas deferens, visible through the delicate integument.

FIGURES 323–331. Neoperla dubia-complex, femalesand eggs.N.dubia Klapálek:323, genital situs; 324, eggwith moderately raised costae (Ghana, Agumatsa waterfalls); 325–327, egg with flanged costae (D. R. Congo, Odzala N. P.). Neoperla proxima n. sp.: 328, genital situs (D. R.Congo, N of Matadi), 329, egg (Cameroon, Sangmelina). Neoperla vicina n. sp. (D.R. Congo): 330, female situs, 331,egg, arrow points at end of raised part of costa (paratype, Walikale, Z2101). Scales are 100µm, similar structures shown to the same scale. Top view of egg (326) based on combined halves of a photograph. Sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 340–343. Neoperla bareensis n. sp., ♀ holotype: 340, genital situs, stained with safranin, base of SSt twisted and deformed during handling; 341, eggs; 342, opercular egg pole; 5, detail of striae, with a micropyle. Scales are 100µm or as indicated; a, anchor; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 344–350. Neoperla lineata n. sp., female holotype: 344, fore body; 345, genital situs, vagina and SSt were flipped back to expose the anchor pattern on S8; 346, entry of SSt into vagina, arrow points at fine spicules; 347, egg; 348, count of costae on one quadrant of an egg cross-section (from a photograph); 349, detail of pale ring between striae and operculum; 350, micropyle (arrow) in the narrow sulcus between 2 costae, scale is 10µm; the other scales are 100µm. S8, sternite 8; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 351–361. Neoperla conradti (Enderlein): 351–354, male from Baré, abdominal tip and contracted penis; 355–356, male from Matadi, everted penis and detail near midlength of endophallus; 357, 358, ♀ genital situs and anchor pole of egg (from Bangem, Cameroon); the arrow points at spine patches in the vagina; 359, SEM, egg from holotype (courtesy of MPI for Limnology). Figs 353–355 to the same scale. Tergites, HT10 and cercus labeled; a, anchor; ac, anchor cavity; Mi = pores between costae above a concealed micropyle.