Neoperla africana Klapálek, 1909

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 : 59-64

23. Neoperla africana Klapálek, 1909

( Figs. 114–123 , 127–130 )

Neoperla africana Klapálek, 1909: 218 .

Neoperla africana — Klapálek, 1910: 56, figs 2–3.

Neoperla africana — Klapálek, 1923b: 133, figs 4–5. not Neoperla africana — Zwick 1973b: 382, figs. 1–2. Neoperla sp. 9 — Zwick 1976: 686-7, figs 3, 7.

Type material studied: Republic of Cameroon: J lectotype, here designated to stabilise the use of the name: N. Kamerun Joh.-Albrechtsĥhe V. 96 L. Conradt S. \ africana Klapálek \ COTYPE [print, red paper] \ COTYPUS [print, white paper] \ det.P. Zwick 1980 Neoperla sp. N [handwritten on photoprint] ( NMCZ; pinned, genitalia in microvial) ( Fig. 19 ).

Paralectotypes: 1♀: africana Klapálek \ N. Kamerun, Johann-Albrechtshöhe L.Conradt S. 26.ii–3.iii.1896 ( NMCZ; slide Z19.115) .

Additional material studied. Republic of Angola: 1♁, M.Picker 10/1/17 Angola Leba Gorge 15°0’32.13’’S 13°11’54.92’’E. Republic of Cameroon: 1♀, mis-associated paralectotype of N.camerunensis (End.) : Ochthopetina camerunensis Enderl. Type ♀ [in Enderlein’s hand] det. Dr. Enderlein [print] \ Kamerun Barombi Conradt \ CoTypus [print, yellow paper] \ Mus. Zool. Polonicum Warszawa 12/45 \ Mus. Zool. Polonicum Warszawa Typus n.1299 Ochthopetina camerunensis Enderlein 1909 Paralectotypus [red label] ( MIZ; wingless torso, genitalia and egg on a plastic slide, on the specimen pin). 1♀, mis-associated paralectotype of N. didita (End.) : Ochthopetina didita Enderl. Type ♀ [in Enderlein’s hand] det. Dr. Enderlein [print] \ Kamerun Barombi Conradt \ Co-Typus [print, yellow paper] \ Mus. Zool. Polonicum Warszawa 12/45 \ Mus. Zool. Polonicum Warszawa Typus n.1303 \ Ochthopetina didita Enderlein 1909 Paralectotypus [red label] ( MIZ; pinned genitalia and egg slide on Zelluloid, on the specimen pin). 2♁, Kamerun Ekona 17.4.–20.5.38 Buhr S. G.; 1♁, 1♀,Kamerun Bumbu 16.4.1912 leg. Dr. Dr. Leo Waibel \abends an Lampe gefangen; 1♀, Kamerun, Bascho, freigeschlagene Urwaldkuppe, 600m, leg. Bartsch; 1♁, 1♀, Kamerun Stat. Bascho, Bartsch; 1♁, 1♀, Kamerun, Soppo [=Sappo, Buea, 4°9’N, 9°14’E], II.1912 leg. v. Rothkirsch (MfNBerlin). 1♀, N.Kamerun, Joh.Albrechtshöhe [Kumba, 4°38’N, 9°27’E], H.Rolle vend. 1904 (Mus. Hamburg). 1♁, 1♀, British Cameroons, Kumba, 19.I.1949 (at light) B. Malkin (situs & egg slide); 1♁ ( NEOP 116) 4°38'56.44''N, 9°24'38.62''E, Cameroon, SW Region, Kumba, Lake Barombi Mbo runoff, 320m asl, 30 Oct.2011, MV-lamp (A.Zwick; Slide CAM 2011.01); 1 ♁, Kumba, Cameroon, light trap, early Dec.1952, lg. Nicholas (gift H.B.N.Hynes, slide Z16/173) (all SMNS). Libamba 10km E of Makak, filtered black light, J.A.Gruwell: 2♁, 2♀ 21–25-VI-1974 ( USNM _080, USNM _119); 1♁, 1♀ 12–13-V-1974 ( USNM _085, 094); 1♁, 3♀, 7–9-XII-1973 ( USNM _087 USNM _092); 1♁, 2♀, 18–19-VI-1974 ( USNM _088, 100, 117); 1♁, en route Douda Yaounde FrCam W.Afr. 36 \ Van Zwaluvenburg & McGough (slide USNM 27) (all USNM). 1♁, Libamba 10km E of Makak, filtered black light, J.A.Gruwell, 2-III-1974 ( SMNS ex USNM, Slide Z17.71, NEOP 107). 3♁, 4°51'42.89''N, 9°38'21.17'' E, Cameroon, SW Region, Tumbel, Ngusi, Etambo River, 490m asl, 23.Oct.2011, MV-lamp, A.Zwick ( NEOP 102, NEOP 103; SMNS). 1♀, CAMEROUN: 28 mi. S. Ngaoundéré [N 7.32800 E 13.58500], 1250m, 1-X-1966, E.S.Ross & K.Lorenzen ( CASENT 8413108). 1♀, Kamerun L.Conradt S. (Z19.113, NMCZ). Central African Republic, near Bayanga: 1♁, no locality details, no date; 3♁, near Bayanga, 15.– 18.10.2008, 03°03’58.3” N 16°08’59.6”, E 528 m ( NEOP 117, NEOP 118, NEOP 119) ( SMNS). Democratic Republic of the Congo: 1♀, Odzala Nat. Park, 29.9.–.3.1997 [sic], V. Sinaev (MfN slide 12/12, NEOP 104). 9♁ (1♁ Slide Z17.67, NEOP 105), Rep. Congo, P.N. Odzala, Mdanza [00°34’N, 14°39’E] XI.1992, G.Carpaneto, 1♀ (slide Z22/07, SMNS, gift R.Fochetti). 17♁, 2♀, B. Congo, 12 ml. N of Matadi [5.85N, 13.46E] VII-28-1957 E.S.Ross & R.E.Leech ( CASENT 8413026 & 27); 4♁, 12♀, B. Congo, 39 km S of Walikale [1.47S, 28.07E], 700m, XII-25-57 E.S.Ross & R.E.Leech ( CASENT 8413030 & 31); 2♁, 1♀, B. Congo, 23 mi E of Kama [3.52S, 27.12E], 16.8.57, Ross & Leech ( CASENT 8413029). 7♀: Rep. Zaire W. of Lake Kivu, Kalengo stream, 1700–1800m 1972–73 B̂ttger, emergence trap ( SMNS; Z73/3. NEOP 101). 1♁, 1♀, Congo, Kivu area, for. Blambwe, terr. Lubero, 950m, XI.1956, R.P.M.J.Célis (egg slide SMNS). 1♁, Musée du Congo Kitobola [-5.38, 14.52] 1911 Rovere \ R. dét. H 230 \ Cotypus \ Ochthopetina camerunensis ♁ Enderl. \ Neoperla camerunensis End. P. Navás S. J. det. \ Paratypus ♁ camerunensis End. 5 \ Cotypus (slide Z19.41) ( Navás 1912); 1♁, Coll. Mus. Congo Basoko V-1946 P.L.G. Benoit; Congo belge: P.N.U. [Upemba] Kaswabilenga: 1♁, affl. g. Lupiala (1300m) 14–20-xi-1947 Mis. G.F.de Witte, 768a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); 4♁, 2♀, (R. Lufira) (700m) 15-ix–6-xi. 1947 Mis. G.F.de Witte, 768a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); 3♁, (R. Lufira) 5 Navás (1912) does not mention his intended paratype designation

(700m) 23–24.IX.1947 Mis. G.F.de Witte, 769a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); 1♁, Kongo, Lukavo (afl. dr. Lufira) 700m, 6.–9.10.47 Mis. G.F. de Witte Nr. 814a; 2♁, (700m) 17-x-1947 Mis. G.F.de Witte, 845a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); 1♁, (700m) 21.x. 1947 Mis. G.F.de Witte, 878a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); 1♁, (r.dr. Lufira) (700m) 22–26-X-1947 Mis. G.F.de Witte, 856a \ Coll. Mus. Congo (ex coll. I.P.N.C.B.); suchen! [5.11°S, 27.15°E] (r.dr. Lupiala) 1300m 13.–27.XI.1947, Mis. G. F. de Witte 963a (all Neoperla spio (Newman) Dr. N.Hynes det., 1951). 1♁, Congo Belge, P. N. G. [Garamba] Miss. H. de Saeger I/a/3 , 8-V-1950 Réc. H. de Saeger 497 \ Neoperla spio Hynes det.1959).1♁, Coll. Mus. Congo Bas-Congo: Lukula 1952 Dr R. Wautier; 1♁, Coll. Mus. Congo Tanganika: Mpala 780m XI-1953 H. Bomans [-6.768, 29.487] (all MRAC). 2♀: Ukaika-Mawambi 1910. Grauer \ Neoperla africana Klp. Klapálek det. [handwritten + print]; 1♀: Mawambi-Ukaika, Grauer, Nov.Dez. 10 \ Neoperla \ africana Klapálek [print]; 1♀: Urwald.Beni Okt. ’10. Grauer \ africana Klapálek [print] ( NHMW). Gabonese Republic: 1♁ (slide 75.1, NEOP106 ) 1♀, Makokou, 227L, Coiffait (slide Z16.158); 2♁, 4♀ with the same data ( SMNS). ♁, GABON: Ngounyé Prov. forest opening of logging road 1°41.924’S 10°44.728’ elev. 147m , 22.ii.2018 light trap S.M.Bybee & Y.M.Pacheco ( USNM; slide Z18.98, NEOP108 ). Republic of Ghana: 2♁ Ghana, Volta Reg. Kute River Menu 11.12.1990 J.S. Amakye light trap ( SMNS, Z17.070 ); 3♁, Ghana Western region, Ankasa game prod. Reserve [5.44N, 2.07W, 47m] 10–16.12.93 St. 12 J. Kjaerendsen & T. Andersen light trap ( ZMBN, Slide Z 17.072, NEOP109 ). 13♁, Ghana , Subri R ., 5.11.1993, light trap; NUFU project, Uni.Bergen [Subri: 6,238N, 2.282W] ( ZMBN, NEOP110 , NEOP111 ; 3♁); 3♀, dto. (MfN; Slide SMNS, body used for DNA: 539_F_529_F_Z17.019). 22♁, Ghana Volta region, Agumatsa waterfalls, Wli , St. 10 19.11.1993 H. Olsvik light trap; NUFU project (Z17.036 NEOP112 ; ZMBN & SMNS). 19♁, 1♀, Ghana Volta Region, Wli Falls, Afegame , St. 6 11.03.1993 J.S.Amakye & J. Kjaerendsen light trap, NUFU-project ZMBN Sample no. 1 (♁ Z17.22, NEOP113 ; ♀ Z17.24, NEOP114 ); 1♀, slide Ghana ♀ 13, 3♁, Ghana Volta region, Agumatsa waterfalls, Wli , St. 10 19.11.1993 H. Olsvik light trap; NUFU project (Slide 17.030, NEOP115 ); 2♀, slides Z98/3, Z98/7; ibidem ~80♁, 1♀ , 8.3.1993 ( ZMBN and SMNS, slide Ghana ♀ 12); 9♁, Ghana Volta Region, Wli Falls , Afegame , St. 12 07.03.1993 J.S.Amakye & J. Kjaerendsen light trap; NUFU-project ZMBN Sample no. 1 ( ZMBN & SMNS). Republic of Guinea: 2♁, no details ( SMNS ex coll. Élouard); 2♁, petit affluent du Milo à Kousankoro no.2, 21.10.84 ( SMNS, gift J.M.Gibon; old slide Mali 5/1 & Z17.069); 1♁, Loffa (aff. Diani) W de Boola [8.19°N, 8.67°W], 8.3.88 (Slide Z 17.068; SMNS, gift J.M.Élouard) ( SMNS). Republic of Kenya: 1♀: Brit. O.Afrika, Nimute-Gondokoro [4.8437, 31.6689] 14 .– 29.7.1911 Dr. Nägele (MfNBerlin). Republic of Liberia: 1♁, Gibi, WMMann \ Smithsonian Firestone Expedition 1940 ( USNM 26 ). Federal Republic of Nigeria: 1♀, SE State, Obudu Cr. [6.67°N, 9.16°E, 222m], 13.4.73, Medler ( PZ). 19♁: NIGERIA: Benin 1-IV-75 JTMedler ( BYU). South Suda n: 1♁, Musée du Congo Soudan : Rejaf 1923 L. Burgeon \ R. d´t. 1647 \ Neoperla haugi Nav. P. Navás S. J. det. ( MRAC). Togolese Republic: 2♁, 1♀ (Z16.130) Amoutchou , 28.11.1985, V. Landa ( SMNS). Republic of Uganda: 1♀, Brit. O. Afr., Uganda , Nimute-Gondokoro ,14 .– 29.7.1911 Dr. Nägele (MfNBerlin; slide Z19.08).

Habitus. WL 9.8–14.0mm, mean 11 and 13mm in males and females, respectively. Body and appendages yellow to ochre, except for a brown mark over ocelli. Wings often very dark.

Male ( Figs. 114–122 ). Process of T7 sharply pointed and raised, a few SB at tip and on vertical rear face. Sclerite on T8 Y-shaped, some SB on wide part beneath tip of T7. Sclerite caudally band-like, flat to gently convex. T9 unmodified. HT10 slender, pointed, slightly concave in dorsal, slightly angled and curved down in side view, mediobasal callus with wide rounded end ( Figs. 114, 115 ). Hind tibia not distinctly widened.

Penis ( Figs. 116–122 ) a wide sclerotised tube, straight in dorsal, slightly curved in lateral view ( Figs. 116–117 ), distally covered by minute external asperities. Dorsodistally a membranous notch with small spinules, laterally with a short transverse row of a few spines ( Fig. 118 ). Everted endophallus gently curved ventrad, of uniform diameter (Fig, 120), narrowing only at tip where a recurrent tube begins which by transparency can be followed back to inside the penis tube ( Fig. 122 , rt). On the everted endophallus the transverse apical spine row is less obvious because it then stands before a background of numerous truncate spines and asperities forming a rough dark sleeve around the endophallus base. Beyond the sleeve a ring of spines surrounds the endophallus. More distally the ventral side is bare, dorsolaterally stand widely spaced hooks, the largest form a lateral line. Small spinules surround the tip.

Female ( Fig. 123 , and top row of Figs. 127–130 ). S8 with three brownish marks around a variably distinct pale anchor pattern. Vagina unmodified except for some transverse folds near genital opening. The SSt forms a coil of one, rarely up to 1.5 rings. Basal section very wide, with numerous longitudinal folds on concave side and apparently extensible. At base, spines form a narrow band on convex side of SSt, distally the band widens abruptly producing an angular projection. The completely scaly distal section and the basal section of the coil of similar length. The coil is very firm, it usually keeps its shape during maceration and manipulation.

Egg ( Figs. 127–130 ). On average 381*211µm, 1.8 times longer than wide, with 10–12 straight ridges, micropyles exposed, collar with, operculum without cells. Size and shape are variable ( Figs. 127–130 , lower row).

DNA ( Figs. 492 , 495 ). A total of 19 specimens from Ghana, Cameroon, Gabon, Central African Republic and D. R. Congo were sequenced, representing the core of the geographic spread of this species and providing very strong support (94.5/100/100) for the monophyly of the species as here considered. The sequences are markedly diverging and form six distinct lineages, likely representing a species complex that cannot be resolved with current morphological characters and should be investigated with a much broader range of nuclear markers than just COX1. While the lectotype was not sequenced, the sequenced specimen NEOP116 was collected very close to the type locality, Kumba in Cameroon. Both sexes are represented, although male specimens are most abundant .

Variation. Neoperla africana may be a variable species or a complex in speciation. Males can be recognised by the scaly sleeve around the endophallus base and by the row of large hooks dorsolaterally along the endophallus. Most penes are regularly curved with exposed apicolateral spine combs and a short rough basal sleeve ( Fig. 121 , sl), in some Ghanaan males the base of endophallus is sinuous ( Fig. 122 ).

Females are best recognised by the firmly coiled SSt with wide base, a narrow scaly part of similar length and a projecting angle in the spine cover ( Fig. 123 ). Eggs vary, distinction from relatives not always reliable. Different shapes occur, but each female carries only one kind. Egg length ranges from 345–435µm, the length/width relation from 1.5–2.2. Stout eggs resemble N. transvaalensis (e.g., Fig. 127 ) but have a wide instead of a narrow-contracted collar. Egg surface with irregular punctation, mostly fairly coarse, and the top of ridges may have a low fishbone pattern ( Fig. 111 ). However, there are also eggs with fine punctation and smooth ridges which are indistinguishable from N. camerunensis ( Fig. 112 ).

Notes on types and nomenclature. A “vorläufiger Bericht”, i.e., a preliminary report on exotic Plecoptera ( Klapálek 1909) contained the previously unpublished names of six African Neoperla species in a key, which fulfills the requirements of ICZN, Article 12.1 (= the names are available). However, the names N. africana and N. sjostedti were presented in a format suggesting both had already been described. Indeed, Klapálek had previously submitted their descriptions but the intended original descriptions were late ( Klapálek 1910). Klapálek (1923b) quoted both papers as published in 1909, adding “Id.” (Latin idem —the same); the specimen lists do agree. Claassen (1940) and Illies (1966) also gave 1909 for both papers. However, documents of the Swedish Academy of Sciences show that the detailed descriptions in the report on Sjoestedt’s expedition were published only in 1910 (the late P. Brinck, in a letter).

Klapálek (1910, 1923b) had several specimens, most from the Zoological Museum in Berlin one of which is the lectotype in Klapálek's collection. Neoperla africana cannot be recognised from the descriptions and illustrations (Klapálek’s figs. 3–4, 1923b). Klapálek did not recognise his own species: of 8 ♀ in the Vienna Museum which he identified as N. africana only four are N. africana , the other are N. arambourgana Navás , N. heideae n. sp., and N. sp., cf. dundoana n. sp. A lectotype is designated to stabilise the use of the name N. africana .

Today, no specimen in the Berlin museum has Klapálek’s characteristic labels ( Fig. 119 ). P. Zwick (1973b) believed to have found the male in alcohol from D. O. Afrika, Langenburg (leg. Dr. Fülleborn, 15.III.–23.IV.99: Klapálek 1923b) and talked of a lectotype but without the necessary details, his attempt to designate a lectotype was invalid (ICZN Article 74). We have now not retrieved the specimen in question. Instead, we found in Berlin two males from Langenburg, Dr Fülleborn, 3. and 15. November 1899, respectively, which are N. sjostedti needhami . Fülleborn collected additional species at or near Langenburg (today Tukuyu) which Klapálek probably never saw: N.sjostedti sjostedti , N. panafricana , and N. bareensis .

Figures

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 495. Partial (clade E) codon-model maximum likelihood phylogram based on 356 mitochondrial genome and COX1 DNA barcode sequences (all-NT). See Fig. 493 for common details.

FIGURES 114–122. Neoperla africana Klapálek: ♁ from Makokou, Gaboon: 114, 115, abdominal tip, dorsal and lateral view; 116, 117 contracted penis, dorsal and lateral; 118, partly everted endophallus, lateral. 119–121, the lectotype ♁: 119, Klapálek’s labels; 120, penis, endophallus artificially everted; 121, base of same, white arrow points at the transverse short spine group. 122, male from Ghana, Volta region, Wli Falls, black arrow points at the recurrent tube inside the penis. Scales are 100µm, others not to scale. ll, lateral line; n, membranous notch; rt, recurrent tube; sl, sleeve of dense scale-like spinules; so, soft ventrobasal region of penis tube; sr, short spine row.

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 127–130. Neoperla africana, female genital situs with focus on spermathecal stalk (SSt; not to scale) in top row, and an egg from the same specimen in the bottom row. 127, D. R.Congo, Odzala Nat. Park (NEOP104); 128, D. R. Congo, Odzala Nat. Park, Mbdanza (NEOP105); 129, D.R.Congo, Kalengo stream (NEOP101); 130, D. R. Congo, Kivu area, Blambwe. All eggs to similar scales, the scales are 100µm long.

FIGURES 18–25. eggs of the N. lujana-complex: 18, the N. lujana Navás holotype; 19, from Bayanga (CAR, NEOP019); 20, from Odzala N. P. (NEOP018); 21, top view of egg (D. R. Congo, N of Bukavu) (after Moore 1991). N. costata n. sp.: 22, optical cross section; 23, lateral view of an egg, with enlarged profile of a ridge (Liberia, near Tapeta) to the left hand side; 24, egg from Adamaua. N. serrula n. sp.: 25, egg, an enlarged profile of a ridge (Holotype, NEOP010, Ghana, Agumatsa falls, slide 20.12) to the right hand side. Scales are 100µm, unless differently indicated. Details of costae are to the same scale.

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.