Neoperla burgeoni Navás, 1926

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 : 89-94

38. Neoperla burgeoni Navás, 1926

( Figs. 205–211 , 213–214 )

Neoperla burgeoni Navás, 1926: 108 .

Neoperla bredoana Navás, 1932: 283 , fig. 82. New synonymy.

Neoperla sp. D — Zwick 1973b: 386, figs. 5–7.

Neoperla bredoana — Zwick 1973b: 394, figs. 19, 28.

6 presumed in N. spaghetti Type material studied: Democratic Republic of the Congo :

Holotype J N. burgeoni : Museum Paris, Congo Belge Central, Prov. de Maniéma, Kindou [2.57°S, 25.55°E, 465 m], L. Burgeon 1917 \ Neoperla Burgeoni Nav. P.Navás S. J. det. [in Navás’ hand] \ Type ( MNHN).

Holotype ♀ N. bredoana : Type \ Musée du Congo Waika (P.or.) [~ 70km N of Kindu, on the Lualaba R.] 18-V-1931 H.J.Bredo \ R. dét. 2088 [print], Neoperla bredoana Nav. P.Navás S.J. det. [in Navás’ hand] ( MRAC).

Additional material studied. Republic of Angola: 1 ♀, Angola, Dundo, Piège lumineux VIII.53 Carvalho; 1♀, Angola Dundo IX.48 Barros Machado (gift P. Brinck, SMNS). Republic of Cameroon: 1♀, mis-associated paralectotype of N. africana Klapálek : N.Kamerun Joh.-Albrechtsĥhe [Kumba, 4°38’N, 9°27’E] V. 96 L. Conradt S. \ africana Klapálek \ COTYPE [print, red paper] \ COTYPUS [print, white paper] \ Neoperla burgeoni Navás, 1926 det.P. Zwick 1980 —falsch zugeordneter Paralectotypus von N. africana Klapálek, 1909 [handwritten, Neoperla sp. N [handwritten on photoprint] (pinned, with plastic slide; NMCZ). 1♀, mis-associated paralectotype of N. camerunensis (End.) : Ochthopetina camerunensis Enderl. Type ♀ [in Enderlein’s hand] det. Dr. Enderlein [print] \ Kamerun Barombi Conradt [Kumba, lake Barombi Mbo, 4°38’56.44”N 9°24’38.62”E] \ Co-Typus [print, yellow paper] \ Mus. Zool. Polonicum Warszawa 12/45 \ Mus. Zool. Polonicum Warszawa Typus n. 1298 Ochthopetina camerunensis Enderlein 1909 Paralectotypus [red label]; ( MIZ; pinned genitalia and egg slide on Zelluloid, on the specimen pin). 1♀, Kamerun L.Conradt S. 1898–1899 (Z19.112); 1♀, N. Kamerun, Johann-Albrechtshöhe L.Conradt S. v.1896 (Z19.114) (both NMCZ). 1♀, Cameroon, at light, Mamfe [5.75N, 9.32E] 4.3.75, slide Z17.011, eggs. 1♁, 1♀: Adamaua [6.89, 12.81] Poli (500m) b. Garua A. Weidholz 1.X.36 ( NHMW). 2♁, 3♀: Cameroun, 52 mi S Garoua [9.32N, 13.39E], Boki R., 330m, 29.IX.1966, E.S.Ross & K.Lorenzen ( CASENT 8413082). 1♀, 2°14'13.54''N, 12°48'17.14''E, Cameroon, S-Region, Sangmelina 18 km E Djoum, Anwam 2km NWbW on logging road 680m asl, 3.Nov.2011, MV-lamp, A.Zwick ( NEOP 215; SMNS). 1♀: Kamerun, Mukonje,28.3.–4.4.38 Buhr S.G.; 1♀: Kamerun, Ekona, 17.4.–20.5.98 Buhr S.G.; Kamerun Stat. Bascho, Bartsch (all MfNB). Central African Republi c, near Bayanga: 3♀, 06-III-2012, nuit, chasse de nuit (18h00-00h30): banc de sable au milieu de la Sangha, M’boki (sud village Likembé, bord Sangha) ( NEOP 211, NEOP 213, NEOP 216; SMNS). Democratic Republic of the Congo: 6♀: Kongo, Odzala Nat. Park [1.19°N, 14.85°E; 560m], 29.9.– 3.3.1997 leg. V.Sinaev ( MfNB); 1♀, Musée du Congo Maniema Lowa [-1.4020°, 25.8462°] 30-VII-1930 Dr. P.Gérard \ Neoperla haugi Nav. P. Navás S. J. det. (green label, MNHNP; egg slide Z16/ 186 in SMNS); 1♀, Musée du Congo Ituri [1.5330, 29.4348]: Kaporata 1-vii-1932 L. Burgeon \ Neoperla africana Klap , \ det. Navás S.J. ( MNHN, pinned slide). 1♀ Musée du Congo Stanleyville [Kisangani] 5-VII-1923 A. Collart; 1♁: same except 20-XII-1929; 1♀: same, except IV.26 J. Ghesquière; 1♁, 1♀: Musée du Congo Ituri: Kaporata 1-VII-1932 L. Burgeon \ R. dét. JJ 2969 \ Neoperla africana Klap. P. Navás S.J. det.; 1♁, 1♀: Coll. Mus. Congo Lualaba: Kindu 18-VII-1947 Dr. M. Poll; 1♁, 1♀: Coll. Mus. Congo Uelé: Gangala na Bodio 14–16-IV- 36 L. Lippens; 2♀: Coll. Mus. Congo Ruwenzori W. Mutwanga [29.40E, 0.20N] IV 1937 Lisfranc; 1♀: I.R.S.A.C-Mus. Congo, Bendera terr. Albertville 1000m (à la lumière) N. Leleup X.1958; 1♁: Coll. Mus. Congo Stanleyville Yalemba 8-VIII-1947 Dr. M.Poll \ Neoperla spio Newman det Hynes 1952; 1♁: Musée du Congo Bassin Lukuga IV– VII-1937 de Saeger (all MRAC). 1♀: Lukolela left bank Congo R., 21.7.1930 J.P.Chapin (American Museum, N.Y.; egg slide SMNS). 1♀, B. Congo, 39 km S of Walikale [-1.47, 28.07], 700m, XII-25–57 E.S.Ross & R.E.Leech ( CASENT 8413043); 2♁: B.Congo, 20 mi E of Kibombo [3.93S, 25.93E] ( CASENT 8413080). Federal Democratic Republic of Ethiopia: 1♁, 2♀: Cojeb River [7.059N, 37.366E] 9– 11/4–61 at light S.Chojnacki leg.; 16♁: Ghibe River [8.469N, 37.364E] 13–14/ 5–61, 215 km from Adis Ababa wooden bridge: road: track to The Crown Prince’s Saw Mill, A.Tj[̂nneland] coll. (at light) ( SMNS, gift P.Brinck). slides checked & confirmed 180408. 1♁: Äthiopien, Illubabor Prov., 15km NW Chora [8.98N, 38.75E], 1600m, vi.1973, G. de Rougemont ( SMNS). Republic of Ghana: 1♁: Ghana Western region, Ankasa game prod. Reserve [6.238N, - 2.282W] 10–16.12.93 St.12, J.Kjaerendsen & T.Andersen light trap ( ZMBN, NEOP 203); 1♁, Ghana, Western region, Ankasa game prod. Reserve 6–12. xii.93 St. 7 J. Kjaerendsen & T. Andersen light trap; NUFUproject ZMBN Ref. No. 4 ( SMNS, Slide Z17/026, NEOP 208); Ghana, Brong Ahafo Region, Bui Camp (8°17’N, 2°15’W, 106m), Endr̂dy-Younga, several specimens; Banda-Nkwanta, light trap near yam field, kerosene lamp, 15– 19.7.1965 1♀, 26– 31.7.1965 1♁ ( HNHM). 1♀: Obuasi [6.14N, - 1.70W] Ashanti, Dr. W.M.Graham, on table near lamp12.7.07 ( BMNH). 1♁: Ghana 7miles NW of Kade 16-V-1971 J.A.Gruwell ( USNM). Republic of Guinea: 4♁, 4♀, Niandan [9.17°N, 10.07W] 19.3.87, light trap, evening (Z16/46 & 47, penis in microvial); ibidem, 14.5.85, 3♁ (genitalia in microvial), ~ 12♀ Z16/83; 2♁, Guinea Riv. Milo à Kankan (bassin du Niger) 26.10.1984, (Z16/178); 1♀: Cavally R. at Goéyabolé (Mt. Nimba [~ 7.63°N, 8.42°W]) 22.1.1988 (Z16.92, NEOP 204); 1♁ ( NEOP 214), 1♀ (Z16.70, NEOP 212): Konkouré à Linsan [11.62°N, 12.66°W] 29.01.1987; 5♁ (Z18.70, Z20.83A–C, 20.85) 3♀ (Z16.61), Kolenté à Simbareya [9.89°N, 12.61°W], 3.12.89; 4♀ Guinea, Riv. Rokel à Badala, 11.8.1989 ( NEOP 207; slide Z16.88); 2♀, Guinea, Riv. Léleko affluent Milo près de Kousankoro 11.3.88 (Z16/51; Z16.52: NEOP 205); 1♀, Guinea, PNHN3,10°28‘40‘‘N, -10°26‘42‘‘WFaranan, F. Niger, Somorye 20.06.1996 (Z16.15); 1♀, Guinea: Tominé à Karmafassa, 31.1.1989 (Z16/100) (all J.M.Élouard, SMNS). République de Côte d’Ivoire: Ivory Coast: Cavally R. near village Grabo, 1st quarter 1978, B. Statzner ( SMNS; slides Z16.33,34)); 4♀: Ivory Coast: Cavally R. (blackwater) nr Taȉ [~ 5.87°N, 7.45°W] 02.07.1977 B.Statzner (Z16.36, 37, 49); Ivory Coast, N’Zi-river: 4♀, ca. 10 km N Bocauda 25.01.1977 (Z16.38,42); 3♀, Fetekro upstream from road bridge Bouaké [7.669N, - 5.020W] - Sokoura, 5.5.1977 (slides Z16.39 & 43); 1♀, Ivory Coast, 25 km W of Bouaké, 7.27.10.1971, Gruwell (Z19.92; NEOP 344); 1♁ Côte d’Ivoire Bandama [5.97N, - 5.14W] aux chutes Gauthier, rég. Tiassalé [5.8933N, - 4.8269W], VIII.82 J.H.Hougard & H. Prod’hon ( SMNS; penis slide, no nr.); 3♀, upstream from road bridge Katiola [8.1242N, -5.8092]—Dabakala, near village Tinbé, 4.5.1977; 1♀, Côte d*Ivoire Cavally r. à Lieupleu 3.2.1988, J.M.Élouard, slide Z16/72 (all SMNS). 1♁, 2♀,Coll. Mus. Tervuren Côte d’Ivoire Divo 28-XI-1963 Decelle ( MRAC). Côte d’Ivoire, Danané, Museum Paris, 12.1930 –4.1931 Ch.Alluaud & P.A..Chappuis (1♁ BMNH, Hynes det. N. spio ; 2♁ MNHN); Côte d’Ivoire, Toumodi, Museum Paris, 12.1930 –4.1931 Ch.Alluaud & P.A.Chappuis (2♁ BMNH, Hynes det. N. spio ; 1♁, 7♀ MNHN). 1♁, Ivory Coast, 25 km North of Bouaké, black light trap. 27–30-X-1971 J.A.Gruwell ( USNM). Republic of Liberia, Suakoko, C.C.Blickenstaff: 4♁, 6.x.52; 1♁, 5.i.52; 2♁, 7.xi.52; 2W, 14.xi.52; 1♀, 10.–20.iii.52; 1♀, 27.–30.iii.52; 2♁, Mt. Coffee, Febr. 1897, Mrs Sharp, R.P.Currie ( USNM). 1♁: Liberia, 36 mi S Voinjama [8.42N, 9.75S], 13.8.66, Ross & Lorenzen ( CASENT 8413083). 1♀, Liberia, 22 mi SE Tapeta [6.49N, 8.86W] 17.VIII.1966, Ross & Lorenzen ( CASENT 8413088). 1♀, Liberia, 8 mi NW Belefuanai [7.26N, 9.43W], S fork S.Paul R., 11.8.66, Ross & Lorenzen ( CASENT 8413089). Republic of Malawi: 1♁, Nyasaland Dr. J.E.S.Old 1913-167 ( BMNH; Hynes det. N. spio ). 2♁, Nyasaland: Chikonje near Chiromo [-16.54, 35.14], XII.1957, C.Sweeney ( CASENT 8413081). Republic of Mali, Chute de Ngougny, Bafing, LF 9./ 10.3.1997 1 ♀, 1 ♁ penis naturally erect; 1♁, Mali, riv. Baoulé à Missira, 09.10.1984, J.M.Gibon ( SMNS, Z16/179). Republicof Namibia: 2♁: Kavango, Popa-Falls, 26.2.– 2.3.1992, W.Mey. 1♁: Namibia Exp. ZMB 1992 Kavango: Gelukiie. Kavango-Ufer 18°03'S / 21°08'E 1.III. leg.J. Deckert (all MfNB). Federal Republic of Nigeria: Nigeria, Samaru, BM 1970-604: 1♀, 26.-31.5.70; 1♀, 3.-10.6.70; 2♁, 7.-14.6.70; 4♁, 15.-22.6.70; 6♀, 26.6.-7.7.70; 6♁ (slides Z17.76 & 77, 19.96), 5♀, 7.-14,7.70; 1♁, 8♀, 13.-20.7.70; 5♁, 21♀, 21.-29.7.70; 3♀, 29.7.-6.8.70; 1♁, 2♀, 6.- 23.8.70; 2♀, 24.-30.8.70 (all P.H.Ward, BMNH). 1 pharate ♀, Nigeria, SE State, Obudu Cr., 13.v.1973, J.E.Medler; NIGERIA, ca. 40♀: Shaganu Field Station, west bank of Lake Kainji, A. Bidwell [10.20N, 4.25E] ( BMNH; SMNS, 1 slide whole body mount). 2♀: Nigeria, N.W. State, Mokwa, nr. Cattle Ranch, 14.– 19.8.1970, P.W.Ward ( BMNH); 1♁: Nigeria, 21 mi W Kaduna [10.508 N, 7.417 E], 650m, 11.IX.66, Ross & Lorenzen ( CASENT 8413084). Republic of Senegal, 2♀, Senegal, Kokofata [12.79°N, 9.94°W] 7.11. 88 P. L.soir (Z16/84&85; J.M. Élouard, SMNS). Republic of Sierra Leone, 1♁, F. Rokel à Makaba [8.40652, -11.84601] 08.02.1985, J.M.Élouard ( SMNS, slide Z16.106). 1 ♀: Pampana près de Magburaka [8.67°N, 11.96°W], 5.2.89, Z16.78; 1♀: Ivory Coast Région de Grabo, petite rivière, route Grabo-Tiboto 11.02.1988 (Z16.73) (J.M.Élouard, SMNS). 1♁: Sierra Leone Gongo 11.4.09 A.Pearse; 1♁: Sierra Leone Njala X.1926 E.Hargreaves (both ( BMNH, Hynes det. N. spio ). 1♀: Sierra Leone, Njala, 16.xi.–14.xii.35, Vanzwaluwenburg & McBough ( USNM). Republic of South Africa: 6♀: RSA, Transvaal, Pullen Farm, 23.–24.XI.2004, W.Mey [near Nelspruit, 25.48°S, 30.97°E] ( MfNB). 1♀: South Africa, Natal, Pietermaritzburg, 1-5-79 S. McLaughlin (Z16.001; gift J.Illies); Republic of South Africa: 7♁, 2♀, Prov. Limpopo, Waterberge, Mapote Farm, 22.–24.Feb.2012, 24°07’51.41”S 28°38’40.88”E D.Bartsch ( NEOP 218, NEOP 219, NEOP 220, NEOP 221, NEOP 222, NEOP 223, NEOP 224, NEOP 225, NEOP 226; SMNS). 2♁, Südafrika, Transvaal, Mogol R. at Vaalwater & Dwars R. ca 3mls from Mogol sampling point, 24°20’S / 28°03’E, 1.2.1961, slide Z16.007); 1♀: Grootpens Richtersveld SE 28 17 Ab3 2-XII-1980 Univ. van Pretoria \ ex coll. Holm, Pretoria [gift J. Illies]; 3♀, SOUTH AFRICA Drakensberg Garden Natal, III-22-1968 (gift Paul J. Spangler; Z19/27); 1♀, Natal, Drakensberge, Lotheni, 29.27S, 29.31E, T.Wagner; 1♁: Transvaal, Mogol R. at Vaalwater ca 3km from Mogol, 1.2.1961; 1♁: Transvaal, Dwars R. at Vaalwater ca 3km from Mogol, 1.2.1961, J. Illies (all SMNS). 1♀, M’fongosi [-28.71 30.83], Zululand, W.E.Jones Oct. 1911 (S.Afr.Mus., published as N. transvaalensis by Tillyard, 1931). 1♀, Transvaal 5 km w.Kaapmuiden 26 January 1974 Ashley B. Gurney (pinned slide Z16/185, eggs & vag.; USNM _5). 1♀: Natal, 1872; 1♁, S. Africa, R.E.Turner, Port St. Johns, Pondoland 25.– 31.3.1923 [31.38S, 29.33E] ( BMNH). 1♀, South Africa: Cape Prov.. Ravens Wood 125meters Keiskama River, nr Rt N2 bridge, 5 March 1978 D.&M.Davis, B.Akerbergs ( USNM). Republic of Zimbabwe: 1♀, N. Rhodesia Livingstone 17.5.1951 Brinck & Rudebeck #309 ( MZLU); 3♁, 4♀, Rhodesia Victoria Falls Nat’l Park iv-3-6-1968 Paul Spangler ( SMNS; Z17.031 & 032). 1♀, Rhodesia, Victoria Falls [17.83S, 25.83E], 900m, 11/ 12.X.1966 Ross & A.R.Stephen ( CASENT 8413086). 1♀, S. Rhodesia, 41 mi S of Umtali, 640m, 17.III.58, Ross & Leech ( CASENT 8413087). United Republic of Tanzania: 1♀, D.O.Afrika, Amani, Mkulumusi, Usambara-Gebiet 1.– 9.1.1906 Dr. Chr. Schröder ( MfNB). 1♁: G.E. Africa Vy. Ruaha R. [ca - 7.48S, 36.54E], Dec.18–21 1910 S.A.Neave; 1♁: N.E. Rhodesia, Lr Luangawa R., 4.-13.-9-1910 16-18,000 ft S.A.Neave (both BMNH, Hynes det. N. spio ). 1♁, 1♀: Tanganyika-Terr. Massassi, 450m, 15-23.VI.’36, Zerny ( NHMW). 1♁: Rio Mongola Mai 1928 \ Muséum Paris Zambèze Sâne, J. Surcouf 1929 \ Neoperla africana Klap. P.Navás S. J. det. [in Navas' hand] ( MNHN). Togolese Republic: 1♁ (slide Z17.74), many ♀, Togo, R. Mono, Tététou, 25.11.1985, V.Landa; 16♁, 10♀, Togo, R. Mono, 01.12.1985 (1♀ NEOP 210, slide Z16/143, 1♀ NEOP 347, slide 16/126) (slides Z16.136 & 138, 17.78&79, 19.93–95); 2♀, 25. and 30.11.1985, Kpassi (Mono R.) (Z16.112 & 115) (all V. Landa; SMNS). 1♀, Togo, Bismarckburg ( NMCZ, Z19.107). Republic of Uganda: 18♀: Uganda, Jinja, May 1957, A.Tjönneland ( SMNS, slides 19.88 + 19.89 [ NEOP 217]); 1♀: Banks of Nile near Kakindu 3,400 ft Aug. 24–25 1911, S.A.Neave ( BMNH).

Habitus. WL 8.8–14.0mm in males, 10.0– 17.2mm in females; females from the Drakensberg Garden are the largest. Fully pigmented specimens, especially males, are dorsally dark yellow to ochre-coloured with dark grey wings. Distal third of femora, tarsi and tibiae dark, tip of tibiae pale. Antenna blackish, cercus light.

Male ( Figs. 205–210 ). Hind tibiae flat and wide. T7 sclerite square, corners rounded ( Fig. 205 ). A fine white line (fs in Fig. 207 ) separates the sclerite from its short raised caudal process with a cap-like tip. The cap is the end of a keel on the ventral side of the T7-process. T8 has in front a wide sclerite caudally from which it is membranous. The sclerite has a caudal extension which at rest is concealed in the membranous center of T8 but can be raised as a flat kidney-shaped process ( Fig. 207 ) by contraction of longitudinal muscles attaching to extensions of antecosta 8 ( Fig. 113 ) which leave a passage for the aorta. T9 is squeesed in between T8 and HT10, it has lateral pilose humps and short median furrow. HT10 with long, slender, regularly curved process, mediobasal callus tongue-shaped ( Figs. 205, 208 ). Stout caudal setae on S7, setae on S8 longer and thinner.

Penis tube well sclerotised, straight. The wide apex ventrally and dorsally with pale areas between which long slender lateral sclerites remain. There are many spinules in the dorsal membranous area ( Figs. 209–210 ). The everted endophallus is much wider than the penis tube for a stretch about as long as the tube. Distally it grows narrow and resembles a rostrum or an elephant's trunk. A narrow bare recurrent tube extends over the entire length, from the tip to the basal penis opening. The endophallic armature begins some distance from the penis apex and consists of unordered sharp spines, the dorsobasal ones are large, ventral side nearly bare. Distal spines smaller, those on the trunk-like portion are delicate.

Female ( Fig. 213 ). Anterior half of S8 with a trapezoidal dark brown mark which is incised in front and truncate caudally. The caudal half of S8 is completely unpigmented ( Fig. 213 ). Vagina with slender base and rounded front, unmodified. SSt coiled, of uniform width and coated with slender scales, length variable, about 1.5–10 times as long as the vagina.

Egg ( Fig. 214 ). Approximately 370*205µm, ovoid, surface often very shining. Collar fused, integrated into general egg contour, only the walls of 2 rings of cells project a little. Operculum parabolic, no visible eclosion line, surface with fine punctures without order, smooth ( Fig. 214 ). Anchor mushroom-shaped, inserted in a deep cavity. About 25 straight striae, the bare costae raised, punctures in sulci fine, unordered, micropyles exposed.

DNA ( Figs. 491–492 , 496 ). A total of 25 specimens from Guinea, Ghana, Togo, Cameroon, the Central African Republic, Uganda, and the Republic of South Africa were sequenced, representing a large part of the distribution of this very widespread species. The monophyly of the species is maximally supported if N. sassandrae sp. n. is included (100/100/100), but the cluster is partly heterogeneous, with at least three distinct lineages that cannot be accounted for by just genetic variation across a huge geographical distribution. A cluster of (near) identical sequences for specimens from Guinea, Cameroon, and Central African Republic might appear as cross-contaminants as specimens from the same countries are placed in different lineages, but collecting localities within each country differ and specimens were sequenced in different laboratories over several years. Future research is likely to split this species complex further. The species is near maximally supported (97/100/100) as sister to N. beta n. sp.. Both sexes are well represented.

Notes. The type locality of N. burgeoni on the Lualaba R. is only approximately 70km downstream from the type locality of N. bredoana . The two nominal species are conspecific.

The most obvious distinctive character of females is the contrast between the trapezoid brown sclerite in the anterior half of S8 and the almost white caudal half of the sternite. The egg of the typical morph is unmistakable, no other egg has strong abruptly ending striae and a completely smooth operculum. The most obvious distinctive character of females is the contrast between the transverse brown sclerite in the anterior half of S8 and the unpigmented almost white caudal half of the sternite.

There is a single female from the Comoro Islands 7, located about halfway between the African continent and Madagascar where no Neoperla occur. The specimen differs from N. burgeoni only by some slender spines in the vagina next to the SSt-attachment.

Variation. Even if the endophallus is incompletely everted N. burgeoni can usually be recognised by the combination of the dorsodistal spine patch at the penis tip, the size difference between spines on the dorsal and ventral side of the endophallus, and also by the fine spines on the trunk-like section of the endophallus which are visible by transparency (sp on Figs. 209–211 ).

In several males from Garoua in western Cameroon and from Togo the penis tip is entirely weakly sclerotised, instead of having slender lateral sclerite strips between dorsal and ventral membranous areas. However, the dorsodistal patch of small spines is present. The endophallus is not much wider than the tube and is largely covered with uniform spines, except the distal ones smaller. The rostrum-like distal part may be longer than the rest, it lacks spines ( Fig. 211 ).

In females the length of spermathecal stalk varies from hardly twice as long to about 10 times longer than the vagina and scales in the SSt are sometimes rather pale. Shape of eggs varies little, probably only with degree of hydratisation. For example, inflated ovoid eggs show a step in the contour at the base of the projecting collar. In eggs from Guinea, Cameroon, the Central African Republic, and Uganda, the costae near midlength bulge a little over the sulci, which in vertical view of the chorion suggests parallel stripes. In a female from Pietermaritzburg costae are about twice as wide as the sulci but not in other South African specimens. In the type specimen of N. bredoana and others with contrastingly pigmented S8, the operculum is completely smooth and finely punctate. However, a wide spectrum of different operculum structures occurs, from a barely perceptible delicate network to deep punctate grooves resembling N. beta . We noticed no correspondence between morphology, geographical distribution, or topology of the DNA-tree.

7 FRANCE ÔUTRE MER: 1♀, Mayotte Comoro Is. 5.8.1911 Leigh \ Neoperla sp. F det. R.E. Dewalt 2018 ( ANSP; pinned, dust covered, some fragments on a card, genitalia and eggs on microscope slide Z18.100)

Figures

FIGURE 491. Codon-model maximum likelihood phylogram based on 83 largely complete mitochondrial genome sequences only (mt-NT). Statistical support values (SH-aLRT, UFBoot) from this and other analyses summarised by squares as explained in the figure. Terminal labels include information on the species, type status, gender, originating country, unique sequence identifier (NEOP###), sequencing approach and total basepairs of 13 mitochondrial, protein-coding genes. Clades of particular interest highlighted with colours that correspond to the clades in other trees. HT = holotype; PT = paratype; PCR = Polymerase Chaine Reaction; WGS = Whole Genome Shotgun.

FIGURE 492. Simplified overview of codon-model maximum likelihood phylogram based on 356 mitochondrial genome and COX1 DNA barcode sequences (all-NT). Terminal labels include information on the species only. Clades of particular interest highlighted with colours that correspond to the clades in other trees. The full tree is shown in Figs 493-498.

FIGURE 496. Partial (clades G, H) codon-model maximum likelihood phylogram based on 356 mitochondrial genome and COX1 DNA barcode sequences (all-NT). See Fig. 493 for common details.

FIGURES 205–208. Neoperla burgeoni Navás, ♁: 205, 206, abdominal tip in resting position, dorsal and lateral view; 207, same, lateral, process on T8 raised, tergites and hemitergite 10 labeled; 208, medial view of detached right-hand HT10. Only 208 to scale, 205–207 to similar scales. at9, antecosta 9; fs, fine suture; mc, mediobasal callus on HT10; p7, p8, processes on tergites 7 and 8; p10, front part of hemitergite 10, its so-called process; ps9, paramedian swelling on T9.

FIGURES 209–212. Neoperla burgeoni Navás, everted penes: 209, ♁ from Ghana, modified from Zwick (1973), endophallus fully everted; 210, ♁ from Guinea, Simbareya, endophallus incompletely everted; 211, ♁ from Cameroon, Garua, the slender penis (base not shown, light safranin stain) with long, bare distal part of endophallus. FIGURE 212. Neoperla beta n. sp., holotype ♁ (Senegal, Gambia r. at Samédouta), everted penis (NEOP200). Figures 209–212 to similar scales.

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 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.