Neoperla transvaalensis (Enderlein, 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 : 21-26

1. Neoperla transvaalensis (Enderlein, 1909)

( Figs. 1–3 , 6–7 )

Ochthopetina transvaalensis Enderlein, 1909c: 402 .

Ochthopetina tenera Navás, 1915: 374 ; syn. nov.

Neoperla transvaalensis — Klapálek 1923b: 136, 140.

Ochthopetina transvaalensis — Barnard 1934: 519 (partim: female, egg (figs.4c,d), not the male (figs. 4a,b)) Neoperla kunenensis Barnard, 1934: 552 , fig. 6; syn. nov.

Neoperla seyrigiana Navás, 1934: 17 , fig. 105; syn. nov.

Neoperla transvaalensis — Zwick 1973a: 504, figs. 76–77; redescription from holotype.

Neoperla kunenensis — Zwick 1973b: 389, figs. 10–11.

Neoperla Species B — Picker 1980: 188 ff.

Neoperla sp. B , sp. C, N. transvaalensis — Moore 1991, figs. 32–34, 43–44, 49–52.

Type material studied.

Neoperla transvaalensis : Republic of South Africa: ♀ Holotype: Ochthopetina transvaalensis Enderlein \ Transvaal, Zoutpansberg [23.15°S, 29.77°E;] ( PAN, No. 1311; the pinned type is in fair condition; genitalia (stained with safranin) and eggs in Eukitt on Zelluloid ) .

Neoperla tenera : Federal Democratic Republic of Ethiopia: J Holotype: Muséum Paris Ethiopie mérid. Haut Aouache Endessa \ Ochthopetina ? tenera ♁ Nav. [in Navás’ hand] \ Typus [red card, handwriting] \ Neoperla transvaalensis (End.) det. P. Zwick 1980 ( MNHN; pinned, in poor shape, wingspan 27mm, everted penis in microvial on specimen pin).

Neoperla seyrigiana : Republic of Kenya: J Holotype: Mery (Kenya) 6.VI.32 \ Muséum Paris Longin Navás legit 19[illegible] \ Neoperla seyrigiana Nav. det Navás S.J. \ Type \ Neoperla transvaalensis (End.) det. P. Zwick 1980 ( MNHN; pinned, in poor shape, WL 13.3mm, everted penis in microvial on specimen pin).

Neoperla kunenensis : Republic of Namibia: J Lectotype, ♀ Paralectotype (here designated to ensure stability of nomenclature): Otjimbumbe Kunene R. Mar. 1923 / K.H.Barnard B.M. 1934-162 \ Ochthopetina kunenensis J \ Neoperla transvaalensis (End.) det. P. Zwick 1980 ( BMNH, J genitalia originally glued on card, transferred to Euparal on Zelluloid); J Paralectotype: Erikson’s Drift Kunene River Mar. 1923 / S.W. Africa Mus. Exped.\ K.H.Barnard B.M. 1934-162 \ Ochthopetina kunenensis J \ Neoperla transvaalensis (End.) det. P. Zwick 1980 ( BMNH, cleared J genitalia originally glued on card, penis missing; transferred to Euparal on Zelluloid). Additional paralectotypes (here designated): 3♀: Otjimbumbe Kunene R. Mar. 1923; 1♀: Erikson’s Drift Kunene River, South West Africa, K.H.B. & R.F.Lawrence, III.1923 (S.Afr.Mus.; pinned, genitalia and eggs in Euparal on Zelluloid slides on the pins; egg slide also PZ).

Republic of Cameroon: Libamba [3.55°N, 11.10°E, 646m] 10km East of Makak, 7–9-XII-1973, at black light, J.A.Gruwell ( PZ ex USNM; 1♀ with similar data except - III-1974: T. Moore, 1991, figs. 43, 44) ; 1♀, Cameroon (Littoral) Reg. de Kribi [2.94°N, 9.91°E], D.Gronier 1925, Mai, Juillet, Neoperla africana Klap. det. Navás S.J. ( MNHN) . 1♁, Kamerun, Ekona [4°40’N, 9°20’E], 17.4.–20.5.38 Buhr S.G.; 1♁, N.W. Kamerun, Moliwe [4.08°N, 9.29°E] b. Victoria, Vorwerk Wöhrmannshöhe, Frfr v. Maltzau G. 1.–8.12.07 (both MfNB) . 1♀: Cameroon, Akom [2.7866 N, 11.304], 450m, 18.X.66, Ross & Lorenzen ( CASENT 8413006 ) ; 1♀: Cameroun, Nanga-Eboko [4.683 N, 12.367 E], 620m, 5.X.66, Ross & Lorenzen ( CASENT 8413008 ) ; 2♀: Cameroon, Nkongsamba [4.968 N, 9.935 E], 960m, 2.XI.66 Ross & Lorenzen ( CASENT 8413011 ) . 2♀ CAMEROUN: Bertoua [4.5196, 13.6998], 660m, 4-X-1966, E.S.Ross & K.Lorenzen ( CASENT 8413111 , 8413112 ) . Central African Republic: 1♀, Oubanghi-Chari , Bangui [4.39°N, 18.56°E, 376m] I/III.68 coll. Breuning ( MRAC) . Democratic Republic of the Congo: 1♀, Rep. Congo, P.N. d’Odzala [1.19°N, 14.85°E; 560m], Mbdanza XI.1993 G. Carpaneto ( PZ, gift R.Fochetti) ; 1♀, Kongo, Odzala Nat. Park , 29.9.– 3.3.1997 leg. V.Sinaev ( NEOP023 ; MfNB) . 1♀, Belg. Congo, Rwindi Camp at light, 6.4.1948, Emersen [0.78°S, 29.29°E, 1005m] (Mus. N.Y.) ; 3♀: B. Congo , 39 km S of Walikale [1.47S, 28.07E], 700m, XII-25-57 E.S.Ross & R.E.Leech ( CASENT 8413012 and 8413016) ; 1♀: B. Congo , 20mi E Kizomba [5.05S, 17.25E], 13.8.57, Ross & Leech ( CASENT 8413004 ) ; 1♀, Congo (Zaire), Kisangani , Yoko [0.52°N, 25.20°E; 400m], 12.2.2008, LF, leg. V.Zolotuhin ( NEOP032 ;MfN) . 1♀, Kongo, Kaswabilenga, 17.X.47 Miss. de Witte Nr. 845a ; 1♀, Lusinga [8.93°S, 27.18°E] (1760m) 12.–17.XII.1947 Mission de Witte \ Hynes det. N. spio ; 1♀, Costermansville [Bukavu, -2.49, 28.84] V-1938 J. Ghesquière ; 1♀, Kongo, Katanga [9.28°S, 26.48°E, 814m], Kinda, 1950 M. Dierckx ( PZ) ; 1♀, Musée du Congo Mongbwalu (Kilo) 1937 Mme Scheitz; 1♀, Coll. Mus. Congo W.Kivu Mulungu près Shabunda [-2.98693, 27.55371] 1939 Dr. Hautman ; 1♀, Mus. roy. Afr.Centr. Lualaba: Ruwa (à la lumière) I-1980 Dr. V. Allard ; 3♀, Coll. Mus. Tervuren Oubanghi-Chari: Bangui I/ III.1968 ex coll. Breuning ; 1♀, Coll. Mus. Congo Uelé: Gangala na Bodio 14–15-IV- 36 L. Lippens 139 (all MRAC) . 2♁, 4♀, Loudima [-4.117, 13.067], 10.XII.1963, J.Balogh, A. Zicsi collected by lamplight ( HNHM) . 1♁, 1♀, Kwango [6.7°S, 17.72°E], terr. de Feshi, Bae de la Rivière de Kwenge , light trap, Leleup ; 1♀, Kundelungu [Parc National de K., 10.65°S, 27.83°E], Riv. Lualala 16.10.51 22.10.51, Marlier; 1♁, Kundelungu, Riv. Kalumbulua, 16.10.51, Marlier ; 1♀, Etang de Kanere No. 12, 15.7.1950, G. Marlier Vmtl. MNHN, Paris . Federal Democratic Republic of Ethiopia: 1♀, Bahar-dar, edge of Lake Tana , 1500m, at light 18.10.1973 G. de Rougemont ( PZ) . 3♁, 2♀, Ghibe River 13-14/5-61 215 km from Adis Ababa wooden bridge: road: track to The Crown Prince’s Saw Mill) A.Tj. coll (at light); 3 ♁, 2♀, Cojeb River 9-11/4-61 at light S.Chojnacki leg. ; 2♀, Awash River nr. Hot Springs 1/5-61 (at light) S. Chojnacki leg. ; 2♀, Oma River 29/4-61 at light S.Chojnacki leg. (gift P. Brinck, SMNS) ; 1♀, Abessinien 1938 Prov. Wallega [Welega, 8.57°N, 35.73°E] F.B.Neuhaus \ Jubdo Wallega 1900m IV–V ( ZSM) . 1♁, 2♀: Kaffa, Assandabà , F. Piccole Ghibie, 1700m, 22.x.1973 ; 1♀: Kaffa, Bonga [7.27N, 36.25E], 1800m, 27.x.1973, C.Consiglio; 1 ♁, 1♀: Gado, Dodola [6.97N, 39.18E], guado del F. Uebi Scebeli, 2500m, 6.xi.1973, A.Vigna (all CCR) . Guder River: 1♁ Site 1, 17/09/2016, 8°57’56.79N, 37°45’15.42E, 1,987.2 m, W. Graf; 2♁ Site 3, 24/09/2016, 8°58’10.44N, 37°45’32.24E, 1,980.5 m (W.Graf, NEOP022 , PZ) . 1♀, Abessinien 1938 Prov. Wallega F.B.Neuhaus \ Jubdo Uollega 1900m IV–V (no body, only pinned slide with genitalia and some macerated eggs, ZSM) . Gabonese Republic: 4♀, Gabon Moyen-Ogooué Rég., Najolé , Auberge St. Jean , Ogooué R., 0°11’03’’S, 10°46’00’’S 16.12.1993 C.C. Jongkind ( ZMBN) . ~40 ♁, several ♀, Gabun, Makokou [0.56°N, 12.86°E, 484m], 227L & 229L ( Fig. 8G , coll. Berthélemy, PZ) ; 4♀, ibidem, 21– 24.3.1971, J. Mateu (egg slide PZ) ; 2♀, Makokou , light trap, J. Mateu leg. March 14–24, 1971 [ NEOP029 ] (Z16.110; PZ) . Republic of Ghana: 2♁, 5♀, Ghana Subri R. 05.11.1993 light trap, NUFU project, ( ZMBN, PZ) ; 8♀, Ghana Western region, Ankasa game prod. Reserve 10–16.12.93 J. Kjaerendsen & T. Andersen light trap; NUFU-project ZMBN Ref. No. 3 [ NEOP027 ] ; 2♁ (P), 5♀, Ghana Volta Region, Wli Falls, Afegame , St. 10 08.03.1993 J.S.Amakye & J. Kjaerendsen light trap; NUFU-project ZMBN Sample no. 1; 2♁ [ NEOP028 ] , 5♀, Ghana Volta region, Agumatsa waterfalls, Wli , St. 10 19.11.1993 J. Kjaerendsen light trap; Ref. No. 2 ; 3♀, Ghana Volta Region, Wli Falls, Afegame , St. 3 10.03.1993 J.S.Amakye & J. Kjaerendsen light trap; NUFU-project ZMBN Sample no. 1; same locality , 2♀, 17.11.1993 J. Kjaerendsen light trap; 1♀, Ghana, Dodo Tamli , Asukawkaw R., [~ 7.70N, 0.52E] 22.XI.85, light trap, J.Schorscher ; 2♀, Ghana, Eastern Region, Boti Falls, River Pawmp ???? 19.11.1991, J.S.Amakye, light trap (all PZ) ; 1♀: Ghana: Andem-Volta lake 29./ 30.9.1966 Thys v.d. Andenaerde ( MRAC) . GHANA, Banda-Nkwanta , 1.9.– 23.10.1965, light trap, savannah [8°22’N, 2°09’W] (Endr ̂dy-Younga 71, 81 90), 15♁ ( HNHM) . Republic of Guinea: 2♀, Guinea: Niandan, Bambaya 25.10.1984, light trap, [slide geprüft 19.8.16] J.M.Gibon. 1 ♁, 2♀, Guinea, Loffa (aff. Diani ) W de Boola [8.19°N, 8.67°W], 8.3.88; 1♁, Guinea , Konkouré à Linsan [11.62°N, 12.66°W] 29.01.1987; 1♁, Guinea, Kousankoro no. 2 21.10.1984 (penis slide); 1♁, Guinea, Riv. Léleko affluent Milo près de Kousankoro 11.3.88 (all PZ, gift J.M.Elouard) . 1♀: Guinée franc. Mamou Mlles Homburger & Morrel, 1924 juin-juillet saison humide \ Navás det.: africana Klap. ( MNHNP; MP3 , egg slide PZ). 1♁, Guinea, riv. Koumba à Gaoval 29.1.1989 [ NEOP030 ; penis slide] ; 1♀, Guinea, Kousankoro no.2, 21.10.84 [ NEOP024 ] ; 1♁, Guinea, riv. Koulountou, rte. Youkoukou-Koundara [12.49°N, 13.31°W], 28.1.1989 [ NEOP026 ; PZ] . République de Côte d’Ivoire: 2♀, 25 km N Bouaké 27.–30.Oct. 1971 black light trap J.A. Gruwell ( USNM) . 1♀, Côte d’Ivoire, Cavally à Lieupleu 03.02.1988 (ex coll. Élouard) ; 1♀: Ivory Coast , 6mi N Toulépleu, 250m, 18.VIII.1966, E.S.Ross & K.Lorenzen ( CASENT 8413009 ) . Republic of Kenya: Kaimosi For. [0.13°N, 34.85°E; 1681m], Kakamega, April 1950, E. Pinhey ( USNM) . 1♀ Kenya: Narok District, Maasai Mara National Park, Mara River , 27–30 January 1990, Robert C. Drewes collector ( CASENT 8413110 ) . Kingdom of Lesotho: 1♀, S. Afr. Basutoland , Quthing [30.40°S, 27.71°E] 12–13.III.1951 ( Brinck & Rudebeck 1955: no. 232) . Republic of Liberia: 1 J, Liberia, Suakoku, 31.1.52 Blickenstaff ; 1 J, same, but 10–20.3.52 ( USNM, pinned) . Republic of Mali: chute de Ngougny, Bafing , lf 9./ 10.3.1997 1 ♀, W. Tobias ( SMNS) . Republic of Namibia: Kunene, Epupa Falls [17.00S, 13.25E]: 8 ♁, 1♀, 20.2.2008; 4♁, 5♀, 21.– 23.2.2008 ( NEOP033 , NEOP034 , NEOP035 ) ; 9♁, 1♀, 3.– 4.9.2012; 1♀, 15.4.2013 (all W. Mey, MfNB) ; 1♁, 1♀. C.S.I.R Water Research AML218 \ Epupa falls Kaokoveld, S.W. Africa 26.4.1970 [illegible] Brown taken at light (gift J. Illies, PZ) . Federal Republic of Nigeria: 1♁, Nigeria, Samaru [11.17°N, 7.63°E], P.H.Ward 3.–10.70 ; 2♀, S. Nigeria , U. C. Ibadan [7.37°N, 3.95°E; 203m], H.T. Sutton 26.X.1957 (all BMNH) . 2♀, Nigeria, N.W. State, Mokwa [9.29°N, 5.05°E], I. A.R., Mile 4, 17.–19.8.70, light trap, P.H. Ward ( BMNH; PZ, slides 16/45 & 18/99) . 1♀: Nigeria , 21 mi W Kaduna [10.508 N, 7.417 E], 650m, 11.IX.66, Ross & Lorenzen ( CASENT 8413010 ) . Republic of Malawi: 1♀, Tanganjika, Njassa See [Lake Malawi], Mango , 600m, 10.12.1952, Lindemann + Pavlitzki ( ZSM) . 1♀, TANGANYIKA: 52 mi. NE Iringa [-7.7854, 35.6855], 575m, XI-15-57 ( CASENT 8413016 ; slide Z17.60) . 1♀, Tanganjika, Njassa-See [Lake Malawi], Mango 600m, 10.12.1952 leg. Lindemann und Pavlitzki (pinned, with pinned plastic slide; ZSM) . Republic of South Africa: 1♁, 3♀, South Africa: Natal Lindeque spruit. 1050m 14km SE Bergville [28.93S, 29.36E] 19 February 1979 D&M.Davis, B. Akerbergs ; 3♀, South Africa: Transkei Border Station, Great Kei River & Rt #N2 [~ 32.51S, 27.98E], 5 March 1978 D.&M.Davis, B.Akerbergs ( USNM) ; 3♁, 3♀, Prov. Limpopo, Waterberge, Mapote Farm , 22.–24.Feb.2012, 24°07’51.41”S 28°38’40.88”E D.Bartsch ( NEOP036 , NEOP037 , NEOP038 , NEOP039 , NEOP040 , NEOP041 ; SMNS) ; 1♁, Drakensbergen Natal J.C. Faure 4.IV.42 (S.Afr.Mus., cleared abdomen and everted penis in microvial on pin) ; 1♀, Natal, Drakensberge, Lotheni , 29.27S, 29.31E, T.Wagner ( Fig. 8H , SMNS) ; 1♁, South Africa, Natal, Pietermaritzburg area , Entomology Class N.U. J.Truran 13 May 1976 (gift ILLIES, PZ) ; 1 J, South Africa : Cape Prov. Groebal R, Schoemanspoort ca. 18km N of Oudtshoorn ca. 700m, 17 March 1978, D. & M.Davis, B. Akerbergs (Depository ??; from photos provided by W. Li) ; 1♀, Upington [28.40°S, 21.24°E, 835m], Cape, Fath. Sollier Mar. 1919 ( Barnard 1934, S. Afr. Mus.); 1 ♁, 15♀ (most spent), Kwazulu-Natal, Ukhalamba-Drakensberg NP, Giant’s Castle, River Forest Trail , 23-III-2005, G.Svenson, K. Miller, S. Bybee ( BYU, Slide Z18.71) . 1♀, S.Africa, R.E.Turner, B.M. 1923-189 Umtata [Mthata, 31.59°S, 28.78°E], Transkei, 18.II.–18.III.1923 ( BMNH) . 1♀ S.Afr., Bas., Quthing, 12.–13.iii.1951 (Brinck 232). 2♁, N.Transvaal, the Downs, Malta Forest [~ 27.26E, 25.81S] 3/12/75, M.Picker; 1♁, Tonquani gorge [~ 27.86E, 25.76S] 21/12/75, M.Picker; 1♁, Magaliesberg [~ 27.86E, 25.76S] Bartlett’s Stream, 26/11/78 on riverside vegetation ; 1♀, Injasuti Drakensberg Wazni River 1463 m 29°07.102S 29°03.467E 15/3/2002 coll. D. Stevens (all PZ, gift M.Picker) . Republic of Rwanda: 1♁, 2♀, Ruanda, Mahembe, Terr. Nyanza [2.35°N, 29.75°E], 1400m, 13.–15.1.53, Basilewski ; 1♀, Ruanda, Nyabikenke , 1700m, terr. Nyanza, 12.1.53, Basilewski ; 1♀, Ruanda, Rukara, s. lac Mohasi , 1400m, 3.–5.2.53, Basilewski; 4 ♁, 7♀, Ruanda, affl. Ruzizi, Bugarama [2.71°S, 29.01°E] 21.7.1949 Marlier ( MRAC) (all MRAC) . 1♀, RWANDA: Western Prov., Rusizi District, Nvungwe N.P., Cyamodongo Forest , S 02.54546°, E 028.98478, 2041m, 21.iv.2014, Gen. coll. S.M.Bybee lab ( BYU, NEOP025 ; slide Z18.95) . 1♀, Urundi, riv. Ruzizi vii-1934 (Dr. van Sanghem) ( PZ, ex MRAC) . Republic of Senegal: 2♁, f. Gambie à Mako [12.31°N, 12.35W], 26.01.89, ex coll. Élouard; 1♁, Senegal, Kokofata [12.79°N, 9.94°W] 7.11 . 88 P. L.soir, ex coll.Élouard. Republic of Sierra Leone: 1♀, seli à bumbuna [9.05°n, 11.75°w] 10.02.1989 (pz, gift j.m. Élouard) ; 1♀, Sierra Leone, Northern Province, Bumbuna [9.05°N, 11.75°W] 19.11.1984 W. Rossi ( PZ, gift R.Fochetti) . Republic of Zimbabwe: 1♁, 2♀, [ Shainowe ] R. S. R. 12/9/56 \ C.N.Smithers ; 1♀: C. N. Smithers \ Shawanoe [Shainowe] R. S. R. 17/6/56 (coll. Brinck) ; 1♀, Domboshawa [17.62°S, 31.14°E, 1511m] S.Rhodesia 7/4/57 C.N.Smithers (coll. Brinck). S. Rhodesia Cashel Dist. [-19.56, 32.79], Umvumvumvu R. 18.12.1954 C.N.Smithers (penis in microvial, SMNS) . Republic of Sudan: 1 pharate ♀: sub. No. 3662 Nubische Wüste [20.86, 33.79] Hartmann (MfNBerlin, egg slides Z19.09, Z18.106). United Republic of Tanzania: 2 ♁, 2♀, From light trap, Sigi R. [-5.0318, 38.9071], Amani, Tanganyika 5/6 I 63 ( USNM 144 ) . 1♀, 25. Dec. 1987. Tanzania: Kimboza Forest , near Morogoro: lowland rain forest (Slide Z19.72; HNHM) . Togolese Republic: ♀, Togo, Amu Oblo [région plateaux, 7.30°N, 0.90°E], 23.11.1985 ; 4♀, Togo, Sio à Kati [Région Plateaux, 6.88°n, 0.85°e], 24.11.1985; 10 ♁, 2♀, Togo, Amoutchou , 28.11.1985 (Z16.119) ; 6♀, Togo, Kpassi ( Mono ), 30.11.1985 ; 3♀, Togo, Mono , 01.12.1985 ; 1♀, Togo, Tchamba 2.12.1985 (slide Z16.122) (all gift V.Landa, SMNS) . Republic of Uganda: 2♀, Jinja Bridge [0.42°N, 33.20°E, 1143m], R. Nile, Uganda 3.8.1949 coll. R.Lowe ; 1♀, Uganda : Jinja, 5.5.1954 N.E.Hickin B.M. 1958-25 ; 1♀, same locality & collector, 7.5.1954 ( BMNH) ; 2♁, 20♀, ibidem, leg. Tjönneland ( PZ, gift H.B.N. Hynes) ; 2♀, Uganda, Bussu Busoga [~ 0.42°N, 33.20°E] 1909 Dott. E.Banyon (Mus. Genova) .

Habitus. Most specimens large (WL 12.0– 19.8mm) and uniformly pale ochre, except for dark ocellar rings. A few specimens from the Ethiopian Highlands have darker anterolateral swellings of meso- and metanotum, a pale rhombus on the scutum remains between them. Wing membrane a little turbid, the Angolan female ( NEOP 031) has glass clear wings with fine dark veins.

Male. As for the group. The HT10 process is a straight spike, the mediobasal callus is plump, almost square ( Fig. 1 ). Caudal edge of S8 with a short row of stout spines.

Penis large, plump, soft, about twice as long as wide. Base dorsally with a weak triangular sclerite, apex flat ( Fig. 2 ). Everted endophallus ( Fig. 3 ) plump, nearly straight, apex wide, conical. Triangular sharply pointed teeth with tips directed basad dispersed on about the distal half of endophallus, except on a bare dorsal strip (bs in Fig. 3 ). The largest spines form 2 inconspicuous ventrolateral bands that extend far basad.

Female ( Fig. 6 ). Caudal edge of S8 slightly convex. The vagina has lateral sclerite rods, a large stack of folds (fo) rests on it, the basal folds are the widest. SSt originating in front of the folds, flat, initially directed forward, then caudad. Cuticle on convex side of SSt thick, next to it a narrow strip with microscopical plicae (magnification 200X, the bare zone in Fig. 6 ). Most of stalk with many longitudinal folds ending near blunt tip. Spermatheca an elongate tangle.

Egg ( Fig. 7 ). As for the group, length 340–415, mean 368µm, elongate, anchor pole truncate, collar constricted, narrow and short with slightly flanged rim, no cells. Most eggs with 11–12, occasionally up to 16 straight ridges which all end at the same distance from the punctate operculum; no suture. Chorion densely punctate, rough, between the ridges punctures are arranged in groups of transverse lines. Ornaments on top of ridges are small and variable.

DNA ( Figs. 491–493 ). A total of 20 specimens from Ethiopia, Rwanda, D. R. Congo, Gabon, Ghana, Guinea, Angola, Namibia, and the Republic of South Africa were sequenced for the COX1 DNA barcode fragment and with the genome-skimming approach, representing the large geographic spread of this species and providing maximum support (100/100/100) for the monophyly of the species. The species is placed as sister to a group comprising N. spectabilis n. sp., N. kalengonis n. sp., and N. lujana Navás , but without strong statistical support (33.6/56/4). Both sexes are well represented.

Notes. Neoperla tenera and N. seyrigiana are synonyms of N. transvaalensis . Navás relied on unsuitable habitus characters and often compared his material with misidentified specimens.

Neoperla kunenensis: Barnard (1934) redescribed N. transvaalensis from subjectively associated specimens, his figure shows an egg of N. transvaalensis , probably from the Kunene River. The presumed N. transvaalensis male (from Krantzkop, Barnard’s figs. 4a, b) was a different species ( N. burgeoni , see below) which Barnard compared with N. leroiana Klapálek. A female from Lydenburg (today: Mashishing) in the South African Museum which Barnard identified as N. transvaalensis is in fact N. leroiana . His error made Barnard believe males of the presumedly new N. kunenensis differed much from N. transvaalensis . The large females from the Kunene River are paralectotypes of N. kunenensis that were said to resemble N. transvaalensis “in all characters”. Barnard’s specimens (in the South African Museum) were in a small box labelled “ Ochthopetina kunenensis Barnard ” but there were no name labels on individual specimen pins.

Distribution and variation. N. transvaalensis is very widespread, populations vary. Because endophallus and SSt are membranous and soft, it was not possible to quantify differences. Length/width proportion of eggs varies from 1.6–2.1 within and between populations but biometry of selected egg samples from the entire range revealed no geographical pattern. The holotype and some Ethiopian females have the longest, moderately slender eggs. The shortest and stoutest eggs were observed from the Odzala National Park, in Sierra Leone, and in Rwanda. Eggs from Rwanda and Urundi have wide, very short collars. Size of punctures, roughness of chorion, and accordingly the development of ornaments on ridges vary, one of three N. transvaalensis females from Walikale (in CAS) bears spheres which are very much smaller than spheres in N. laticollis .

Several other species in the N. transvaalensis complex may have similar ornaments.

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 493. Partial (outgroup, clades A, B, D) codon-model maximum likelihood phylogram based on 356 mitochondrial genome and COX1 DNA barcode sequences (all-NT). Statistical support values above branches or near nodes are SH-aLRT and UFBoot for the codon-model analyses, followed by NT123 UFBoot. 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. See Fig. 492 for an overview. HT = holotype; PT = paratype; PCR = Polymerase Chaine Reaction; WGS = Whole Genome Shotgun.

FIGURES 1–5. Neoperla transvaalensis (Enderlein) (1–3) and N. laticollis Klapálek (4, 5): 1, male tergites 7–10, dorsal and lateral views; 2, penis, dorsoventral; 3, artificially everted penis, the retractor apodemes (sclerotised knob) are on the dorsal side; 4, male tergites 7–10; 5, artificially everted penis. The scales are 100µm, other figures not to scale. bs, bare strip; ce, cercus; gp, gonopore; vlb, ventrolateral spine bands.

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.