Neoperla leroiana Klapálek, 1911

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 : 173-176

82. Neoperla leroiana Klapálek, 1911

( Figs. 471–484 )

Neoperla leroiana Klapálek, 1911 , Wien. Ent. Ztg. 28: 103, 2 text figures of the male (see Figs 477, 478 ). Type locality: Redjaf , Lado [ Republic of South Sudan, Rejaf 4.75, 31.59].

Ochthopetina innexa Navás, 1920: 287 , fig. 1; new synonymy.

Ochthopetina petiolata Navás, 1920: 288 , fig. 2; new synonymy.

Neoperla leroiana , Neoperla sp. 8 — Zwick 1973b: 387, figs. 8, 9, 12,13.

Type material of N. leroiana . O. LeRoi collected the two syntypes in Sudan while he worked for the Museum Koenig, Bonn, to which LeRoi’s personal collection was later also given ( Horn & Kahle 1935). The types are apparently lost. We searched in vain for them at Bonn, but also in Klapálek’s collection in Prague, and the MfNBerlin which holds some old material studied by Klapálek. Our interpretation of the name agrees with the illustration of the male abdominal tip in the original description which we designate as Lectotype ( Figs. 477, 478 ). The ♀ paralectotype (here designated) had no subgenital plate ( Klapálek 1911: 104) and was evidently misassociated; see below under Taxonomy and Geography.

Holotype J of N. innexa , Soudan Égyptien Roseires (Haut Nil Bleu) Ch.Alluaud 1906 [print, white paper] \ Soudan Égyptien Roseires (Haut Nil Bleu) Ch.Alluaud 1907 [print, green paper] \ Ochthopetina innexa J Nav. [in Navás’ hand] Navás S.J. det. [print, green paper] \ Type ( MNHN, pinned, WL 9mm; P.Zwick vidit 1974).

Holotype J of N. petiolata , Soudan Égyptien Roseires (Haut Nil Bleu) Ch.Alluaud 1906 [print, white paper] \ Soudan Égyptien Roseires (Haut Nil Bleu) Ch.Alluaud 1907 [print, green paper] \ Ochthopetina petiolata J Nav. [in Navás’ hand] Navás S.J. det. [print, green paper] \ Type ( MNHN, pinned, WL 10 mm; P.Zwick vidit 1974).

Additional material studied. Republic of Angola: 2♀, Angola 21.XI.13, Cuanza Norte Prov. 20km W N’dalantando [-9.30, 14.92], 337m, P. Schuele leg. -18- ( SMNS). 1♀: Angola, prov. Uíge, Finca 7°38’08.6’’S, 14°56’25.8’’E 645m, 24.x.2013, at light, leg. M.Nuss ( SMTD, NEOP 280, slide Z17.49); 1♀, 1♁: Angola, 2 mi E Xandel [-9.39, 17.18], 750m, 4.XII.1966, Ross & Lorenzen ( CASENT 8413064 + 65) (Slide Z18.33a: ♀, Z18.33b: ♁). Republic of Cameroon: J, Kamerun, Lolodorf-Stat. Prof. Ziemann leg. Nr.1600/07 (slide Z19.22); 1♀, Kamerun Pipindi Zenker (Z19.23; both MfNB). 1♁: Kamerun int. Dengi 23.VI.09 Riggenbach S.G.; 1♀: Kamerun int. Benuë unth. Garua 1.VIII.09 Riggenbach S.G.; 1♀: Kamerun Ekona 17.4.–20.5.38 Buhr S. G.; 1♀: Kamerun Ngoko Station Hösemann S.G. 9.IV.02; 1♁: KAMERUN, Ekona [4°40'N, 9°20'E], 17.4.–20.5.38 Buhr S.g. (all MFNB). Libamba 10km E of Makak, light traps, J.A.Gruwell: 4♀, 11–16-XI-1972; 2♀, 1.9.– 10.12.1973; 7♀, 1– 9.3.1974 ( USNM). 1♀, CAMEROUN: 28 mi. S. Ngaoundéré, 1250m, 1-X-1966, E.S.Ross & K.Lorenzen ( CASENT 8413109, Z18.031); N. leroiana Cameroun, Nanga-Eboko [4.683 N, 12.367 E], 620m, 5.X.66, Ross & Lorenzen CASENT 8413062); 1♁, Kamerun Mao Godi [?] 9.vii-09 Riggenbach ( SMNS, ex MRAC). Central African Republic: 1♁ ( NEOP 290), 5♀♀ ( NEOP 283, NEOP 284, NEOP 285, NEOP 286, NEOP 291) near Bayanga, around 2.48N, 16.22E, 382–434m; 3♀: 11-II-2012, 2,480417° N 16,215 528 °E 392 m ( NEOP 282, NEOP 287, NEOP 289); 1♀: 13-II-2012, bord lac, 2,480857°N 16,217943°E 403 m; 1♀, 13-II-2012, UV, bord lac 1 sur bute 2,480 857 N 16,217 943 E 403 m ( NEOP 281); 1♀, 25-I-2012, UV: 17h30-2h, Nord M’boki, bras mort de la Sangha 2,471 964 N 16,081 238 E 382 m ( NEOP 288) (all SMNS) 1♁: Ssanga-Lobaje-Exp, Uaingebiet Bosum Tessmann S.G. ( MfNB). Democratic Republic of the Congo: 2♁, 1♀, P. N. d’Odzala, Mbdanza XI.1992 G. Carpaneto (gift R. Fochetti, SMNS, slides Z16.10 & 22). 2♁, 1♀, Kongo, Odzala Nat. Park [1.19°N, 14.85°E; 560m], 29.9.– 3.3.1997 leg. V.Sinaev ( ZMB). 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) ( MNHN); 1♀, Musée du Congo, Maniema: Lowa 30-VII-1930 Dr. P. Gérard \ Neoperla Haugi Nav. P. Navás S. J. det. \ R. Dét. 2088 ( MRAC). 1♀ Lukulela, left bank Congo R., 1°5’S, [17.20] 21.7.1930 J.P.Chapin (Mus. N.York; slide with eggs also in SMNS). 1♀ Kongo, Katanga, La Kipita [-10.69, 27.48] 8.IV.24 Ch. Seydel N5025; 1♀ Kongo, Katanga, Kipopo (Elisabethville) [Lubumbashi, -11.66, 27.48], 27,8.61 Don R. Maréchal; 1♁, 1♀, Congo Belge. P.N.G[aramba] Miss. H. de Saeger II/fd/17, 14-XII-1951 Rec. H. de Saeger 2910; 1♀, Coll. Mus. Congo Bas-Congo: Tumba 1952 R. F. Anselmus; 1♀, Coll. Mus. Congo Elisabethville II/1938 H. J. Bréde \ R. Det. 5891; 1♀, Musée du Congo Elisabethville Ch. Seydel 1952; 1♀, Coll. Mus. Congo Ubangi: Nouv. Anvers 11-VII-1947 Dr. M. Poll; 1♁, Coll. Mus. Congo Haut-Uele: Paulis X-1947 P.L.G.Benoit; 1♁, Musée du Congo Ubangi: Budjala VII—1937 C. Léontovitch; 1♁, Musée du Congo Kikwit 1920 P. Vanderijst; 1♁: Congo Belge, P. N.G. Miss. H. De Saeger, Pidigala [~ 4.12°N, 29.51°E], 23-IV-1952 H. De Saeger 2358 (Hynes det 1959 N.spio Newm. ; strongly damaged) (all MRAC]. 2♁, 3♀, Loudima [-4.117, 13.067], Station Agriculture, 10.XII.1963, J.Balogh, A. Zicsi collected by lamplight ( HNHM). Federal Democratic Republic of Ethiopia: 1♁ ( NEOP 275), 1♀, 15 km NW Chora Illubabor Prov., 1600m, VI.1973, G. de Rougemont ( SMNS); 1♁: Kaffa, Arguba [7.40N, 36.43E], F. Gogeb, 1420m, 25.x.1973, P.Brignoli (coll. C. Consiglio, Roma). République de Côte d’Ivoire: N’Zi-River, ca. 10 km N Bocauda 09.05.1977 B.Statzner 1♁, 4♀ (Z16/41); 1♀, Côte d’Ivoire N’ze & Cavally à Tai 01.02.1988 (gift Élouard, SMNS). Republic of Equatorial Guinea: 1♁, Span. Guinea, Nkolentangan Xi.07–V.08 G.Tessmann S.g. ( MRAC). Gabonese Republic: 1♀, Gabon Moyen-Ogooué Rég., Najolé, Auberge St. Jean, Ogooué R., 0°11’03’’S, 10°46’00’’S 16.12.1993 C.C. Jongkind ( SMNS, gift J.M.Élouard; NEOP 278). 1♀, Gabun, Belinga [1°5’N, 13°12’E], 155L (H. Coiffait, gift C. Berthélemy; SMNS) Coiffait ( SMNS, gift C. Berthélemy). 3♁: Is. Fernando Po [Bioko], Musola [3.43N, 8.60E], 500– 800m.s.m. L.Fea [Mus.Genova, vidi 8.xi.86; in microvial]. Republic of Ghana: 1♁, Northern region, Banda-Nkwanta (8°22’N, 2°09’W), 5.– 8.9.1965; Ashanti region: 1♁, Kumasi (6°43’N, 1°36’W), 20.10.1967; 1♀, Kwadoso (6°42’N, 1°39’W), 12.5.1959 (leg. Endrödy-Younga, Mus. Budapest). 1♁, Volta region, River Menu at Mensu, 18.11.1993, J. Kjaerendsen light trap ( NEOP 277, abd.-tip in microvial + penis slide GH.08.04); 2♀ Ghana Subri R. 05.11.1993 light trap; NUFU project (Mus. Bergen). 2♁, 1♀, Ghana, 22 mi S. 2♀, Ghana, Dodo Tamli, Asukawkaw R., light trap, J.Schorscher 20.XI. 1985 (Afr.Z85/5), 22.XI.85 (Afr.Z86/2; SMNS). Bolgatanga [N 10.78556 W 0.85139], 200m, 31-viii-1966 E.S.Ross & K.Lorenzen; SMNS]. Republic of Guinea: 1♀, Konkouré à Linsan [11.62°N, 12.66°W] 29.01.1987 (gift Élouard, Z16/153). 1, Afrique Guinée Guachedon 1962 III leg. Ferenczy (slide /19.73, HNHM). Republic of Liberia: 1♀, Liberia, Mt. Coffee, III-1895 Mrs. Sharp; 1♀, same locality, 1-III-1897 R.P.Currie; 1♀ Liberia Suakoko 31 Jan 52 Blickenstaff (all USNM; egg slide SMNS). 1♀, Région de Gabo [6° 53’ N, 10° 11’ W], petite rivière, route Gabo-Tiboto 11.02.1988 ( SMNS, gift Élouard). Federal Republic of Nigeria: 1♁, mis-associated paralectotype of N. excisa (here designated): Abutshi, R. Niger 5.1908 [print, date hand-written] \ Neoperla \ excisa Klapàlek \ Neoperla excisa ♁ Klapálek, 1909 Paralectotypus design. P. Zwick 1988; falsch zugeordneter Syntyp! \ Neoperla leroiana Klapálek, 1911 det. P. Zwick 1988 [cleared abdomen in microvial, penis missing] ( NMCZ). 2♁, NIGERIA: Benin 1-IV-75 JTMedler \ Neoperla leroiana Klapalek det. 1985 B. Stark; 3♀, same locality & date are morph 8A ( BYU + SMNS, slides Z18.72 & 73 FOTO). 2♀, Nigeria, SE State, Obudu Cr., 13.v.1973, J.E.Medler; slides Z16/167 ( NEW 2016-261, für NGS) & Z16/168. 3♀, NIGERIA: Benin 1-IV-75 JTMedler (remains of 2 abdomina and a few residual eggs) Slide 18.72 ( BYU). 3♀, Nigeria, 21 mi W Kaduna [10.508 N, 7.417 E], 650m, 11.IX.66, Ross & Lorenzen ( CAS); 1♀, Nigeria, NW State, Mokwa, I.A.R. Mile 4 17– 19.8.1970 HG-Lamp (gift P.H.Ward; Z16/164). 1♀, AFRICA, Nigeria Umuahia, 10-IV-1975 JTMedler (Morph 8A, BYU). 4♁ Ikom [6°5’N, 8°37’E], Ogoja Prov., 6–7.I.1949 at light B.Malkin (Brinck XI 08A; penis slide Z18.09). Republic of South Africa: 1♀, Transvaal Lydenburg [Mashishing; - 25.08S, 30.45E] P. Kroeger (S. Afr. Museum, Barnard det. transvaalensis ). Republic of Sierra Leone: 1♀, Sierra Leone Kaba à Gotamba, Kilims Nat. Pk. 04.02.1988 (gift Élouard, Z16/154). 7♀, Makaba [8.40652, -11.84601] 08.02.1985; 1♁, Pampana près de Magburaka [8.67°N, 11.96°W], 5.2.89, ex coll. Élouard ( SMNS, slide Z16.81, NEOP 276). 1♁, Sierra Leone, Séli à Bumbuna [8.91N, 11.73W] (Z16/101) (all gift J. M. Élouard, SMNS). Republic of Zimbabwe: 1♁ [penis slide], 2♀, N. Rhodesia Livingstone [Harare, -17.824, 31.050] 16.05.51 ( SMNS). 1♀, Victoria Falls Zimbabwe Jan.2015 (gift M.Picker, Slide MP6, NEOP 279, SMNS): 1♀, Rhodesia, Victoria Falls Nat.Pk., IV.3–6.1968 (slide (Z17.015, gift P. Spangler, SMNS). Republic of South Sudan: 1 pharate ♀: Sudan, 266 Schorscher, Numatinna [SW of Wau, 7.6868, 27.9853], genitalia & eggs on slide ( SMNS). 1♁, 1♀, Musée du Congo Sudan: Nbongalla L. Burgeon \ Neoperla Haugi Nav. P. Navás S.J. det. \ R. det. 1647 \ R. Det. 5891; 1♁, Musée du Congo Sudan: Nbongalla L. Burgeon; 1♁, Musée du Congo Soudan: Rejaf 1923 L. Burgeon (all MRAC). Togolese Republic: 8♀, Togo, Amu Oblo [7.38N. 0.86E], 23.11.1985, V. Landa; 2♁, 25♀, Togo, Sio à Kati, 24.xi.1985 (AfrZ86/22&23, Z19/); 17♁ [in extra vial],> 100♀,Togo, Tététou [7.00N, 1.50E] (Mono R.), 25.11.1985, V. Landa; ca. 15♀, Togo, R. Mono, Ngonlutu, 29.11.1985 V. Landa; ca. 30♀, Togo, Kpassi (Mono R.), 30.11.1985, V. Landa (Z16/113); 2♁, 22♀, Togo, R. Mono, 01.12.1985, V. Landa (Z16/117, ♁ & ♀ Z.16.125) (all V. Landa; SMNS).

Habitus. WL ♁ 8–12 mm, ♀ 9–13 mm. Yellowish, with dark ocellar spot. Wings slightly infuscate. Pronotum greyish, femora, tibiae and tarsi partly or entirely infuscate. Flagellum distally brownish, cercus light.

Male ( Figs. 477–484 ). T7 caudally with a pair of low humps between which the tergite is shallowly concave, slightly sclerotised, with some setae and SB ( Fig. 479 ). A narrow median sclerite continues from this area over the exposed rear face of T7 into the intersegmental fold where it is connected to the T8 antecosta. T8 bears a high forward-inclined erect process with wide top. T9 with pilose lateral humps, the little sclerite at the front of the median furrow is very small. The tip of HT10 is of moderate size and very variable, the obliquely truncate or tongue-shaped hemitergal callus is dorsally a little concave ( Figs. 476–477, 479–480 ).

The penis ( Figs. 481–484 ) is a straight thick tube that narrows caudally to a distal soft nipple. The spines on the endophallus stand in lateral rows, the 10–12 basal spines are sharp spikes, distally spines grow successively smaller, near the base the space between the rows is bare.

Female ( Figs. 472–475 ). A short brown nail of variable shape projects from the caudal edge of S8. It has anterolateral crests and is connected to a sclerite in the floor of vagina. The vagina is unmodified, no spines near the insertion of the SSt which is a narrow tube of variable length. It is of uniform width and coated by triangular scales, the incompletely scaly basal stretch is short.

Egg ( Fig. 476 ). Size 290–365µm long, an elongate ovoid 1.6–2.1 times longer than wide. Numerous levogyrous striae form cells on the collar. On the operculum striae form a few punctate cells with low walls. Costae are rather narrow but still much wider than the very narrow sulci, the two rows of micropunctures are hard to detect and resemble microscopic serrations along the edge of the costa. Sulci widen where micropyles are. The anchor cavity is bowl-shaped, the anchor is mushroom-shaped.

DNA ( Figs. 491–492 , 497).A total of 17 specimens from Sierra Leone, Ghana, Gabon, Central African Republic, Ethiopia, Angola, and Zimbabwe were sequenced for the COX1 DNA barcode fragment and with the genome-skimming approach, representing a large part of the very wide geographic spread of this species. The monophyly of the species is near maximally supported (99.5/100/100), despite noticeable sequence divergences between species. It is placed with maximum support (100/100/100) as sister to N. arambourgana Navás + N. dundoana n. sp.. Both sexes are well represented.

Variation. The top of the process on male T8 is mostly plump and rounded but is sometimes truncate and may even seem to be slightly concave when some lateral SB on front of the process are visible from above. Regional morphs occur among females: most females from Ethiopia, Sudan, Cameroon, Ghana, and westward to Guinea have elongate nails, the edge is rounded, truncate or shallowly notched, and the SSt varies but tends to be long ( Figs. 472–473 ). Nails barely project and the SSt is long in females from the Central African Republic ( Fig. 474 ). Females from Zimbabwe and the Republic of South Africa have nails that are wider than long, distally concave or widely notched; the SSt is shorter ( Fig. 475 ). Most eggs are slender, the egg contour is regular and continuously curved but stout inflated eggs with an angle at the base of the collar also occur.

Eggs of females from Nigeria ( Benin, Obudu Creek, and Umuahia) lack a collar and have a wide anchor pole surrounded by a rim and a short central nipple for anchor attachment ( Fig. 476 ).

Notes. Penes of N. leroiana and N. dundoana are very similar, the apex of the process on T8 differs but varies between individuals. Recognition as distinct species is supported by DNA. Neoperla leroiana has an extremely wide distribution from Sierra Leone and Ethiopia south to South Africa. Types are not available but N. leroiana is the only member of the N. arambourgana- complex known from near the type locality in South Uganda.

VI. Species that are presently not named.

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.

FIGURES 466–476. females in the Neoperla leroiana cluster, eggs and diagrammatic genital situs to similar sizes, from photographs: N. arambourgana Navás: 466, egg of ♀ from Kama (D. R. Congo; NEOP272); 467, ♀ from Popokabaka (D. R. Congo, NEOP271). N. dundoana n. sp.: 468, 469, egg and genital situs of HT (Dundo, Z16.180, NEOP268). Neoperla leroiana Klapálek: 470, egg of ♀ from Ngonlutu, Togo; ♀ genital situs: 471, Xandel, Angola; 472, Odzala N. P.; 473, Guinea, Konkouré à Linsan; 474, Central Afr. Rep., Bayanga (NEOP281); 475, Victoria Falls (NEOP279); 476, SEM of anchor pole of female from Nigeria (courtesy MPI of Limnology, Pl̂n).

FIGURES 477–485. Male Neoperla leroiana and N. dundoana: N. leroiana Klapálek: 477, 478, abdominal tip of lectotype in dorsal and lateral views (modified from Klapálek 1911); 479, 480, the same of a Ghanaan specimen (modified from Zwick 1973a); 481, tip of T8-process of another Ghanaan specimen; 482, penis with largely everted endophallus, ♁ from Togo, Tététou; 483, intentionally broken penis of another ♁ from Tététou showing details of endophallus, with enlarged detail; 484, endophallus base of ♁ from Ghana, R. Menu (NEOP277). 485, N. dundoana, endophallus base of ♁ from Dundo (NEOP267).