Neoperla lujana Navás, 1931

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 : 28-29

3. Neoperla lujana Navás, 1931

( Figs. 13–14 , 16 , 18–21 )

Neoperla lujana Navás, 1931: 135 , fig. 71.

Type material studied. Democratic Republic of the Congo: ♀ holotype: Holotype \ Musée du Congo Kivu: Buserengenye (Rutshuru [1.32°S, 29.42°E, 2273m]) IX-1929 Ed.Luja \ R. dét. 1943 \ Neoperla lujana Nav. P.Navás S.J. det. (pinned, wingspan ~ 30mm) [ MRAC].

Additional material studied. Republic of Cameroon: 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 ( NEOP014 , NEOP015 ) ; 2♁, 6°8’26.25’’N, 10°6’3-68’’E, Cameroon, NW Reg., Bamenda, Mentchum River contributor 15 km NNW Bafut, 630m asl, 9 Nov.2011, MV-lamp ( NEOP016 , NEOP017 , SMNS, PZ) . Central African Republic: 1♁, Bayanga _HS, ( NEOP020 , slideZ19.120) au dessus d’une chute d’eau (ruisseau Songo 1) à proximité du camp 10.10.2008 18h30-3h 03°06’14.6” N 16°08’52.1” E 493 m Camp de transit 2 ; 3♀, Bayanga , near a Bayous-tree ( Triplochiton scleroxylon ) at 3.10N, 3.15E ( Fig.14B ; depository) (♀ IT, NEOP019 , ♀ JL, NEOP021 ) . Democratic Republic of the Congo: 1♀, Belg. Congo , Rwindi Camp ,at light, 6.5.28, Emerson (Mus. New York; FOTO) . 1♀, Kongo , N Lac Kivu, Rwankwi XII1957 J.V.Leroy ( PZ ex MRAC) ; 1♀, Kongo , Kindu, Kitundu, 19.9.1914, L. Charliers ( MRAC) ; 1♀, No. 4 Riv. Nyabarongo, Bunyakiri, 28.3.50, G.Marlier; 1♀, Terr. Kabare, Riv. Ulindi 20.5.49, G.Marlier; 2♁, 1♀, Bendera , terr. Albertville, Katanga 950m X.1958, G.Marlier [1♁ PZ] ; 1♀, Tshibati , Riv. Lwiro , gite, 8.3.59, G.Marlier; 1♁, Riv. Ulindi , Zimbe [reading?] aval, 20.8.50, G.Marlier; 1♁, Riv. Zizi , Bilmba, 24.8.50, G. Marlier ( MNHN) . 1♀, Kongo , Odzala Nat. Park [1.19°N, 14.85°E; 560m], 29.9.– 3.3.1997 leg. V.Sinaev ( Fig. 10 , MfN \ NEOP018 ) . 2♁, B. Congo , 12 ml. N of Matadi [5.85N, 13.46E] VII-28-1957 E.S.Ross & R.E.Leech ( CASENT 8413073 ) [1 in PZ]; 1♀, B.Congo: Lwiro River , 47km N.of Bukavu [-2.49, 28.84], 1950m, XII-15-57 ( CASENT 8413074 ) ; 1♁, B. Congo , 50km S of Tshela [5.01S, 12.96E], 26.7.57, Ross & Leech ( CASENT 8413075 ; SEM-micrograph 55 of Moore, 1991) . Republic of Equatorial Guinea: 1♀, is. Fernando Poo [= Bioko], Bahia de S. Carlos, XII.1901, 500–600m, L.Fea (Mus. Genua) . 1♀, Fernado Poo Spanisch Guinea [Bioko, 3.43°N, 8.60°E] 12.9.1939 \ Prof. Eidmann legit 41. \ coll. Eidmann ( SMNS, slide Z18/101) . République de Côte d’Ivoire: 2♁, 3♀, Ivory Coast , stream on Mt. Tonkui [7.37°N, 7.63°W, 511m], near Man, B. Statzner ( PZ; slides Z86.18, Z20.07) . Republic of Uganda: UGANDA: 8-IV-62, T.R. Williams ( USNM; from SEM-micrograph 31 of Moore, 1991, as sp. B) .

Habitus. Large, WL of ♁ 13.5–14.3mm, of ♀ 15.0– 17.3mm. Rs with 2–4 free branches. Pale species, no pattern, narrow ocellar rings separate, appendages pale.

Male ( Figs. 13–14 ). Tergites as for the complex. Process of HT10 short, slender, with blunt tip. A row of stout spines along caudal edge of S8.

Penis ( Figs. 13–14 ) plump, soft, straight, penis tube in line with everted endophallus. Two large patches of small triangular spines at the endophallus base are separate from distal spines, large areas in between are bare. Spines in the distal area are large, slender, and stand obliquely on an oval base.

Female ( Fig. 16 ). S8 without pattern, caudal edge straight to weakly bilobed (female from Mt. Tonkui). Vagina with widely separate lateral sclerites, the wide caudal ends asperous (hard to see in juveniles) and located in front of caudal segment edge. SSt about as long as vagina, strongly curved, basal half with many longitudinal folds parallel to front edge. Basal half to 2/3 of SSt coated with delicate pale rhomboid scales, apical portion and a narrow strip along the concave side only with folds and wrinkles.

Egg ( Figs. 18–21 ). Length 435–500, average 477µm. The length/width proportion varies from 1.79–2.45, because of variable length of the collar. About 11 rounded ridges, sometimes with spherical ornaments along the top. Collar distally widened, with rudimentary cells, rim flared. Operculum punctate, shallowly domed, in profile a low triangle. Between ridges punctures are arranged in transverse lines. Anchor mushroom-shaped.

DNA ( Figs. 491–493 ). A total of eight specimens from Cameroon, Central African Republic and D. R. Congo were sequenced, representing the core of the geographic spread of this species and provided very strong support (98/100/100) for the monophyly of the species. The specimens from the Central African Republic and D. R. Congo form a distinct, very strongly supported (99.5/100/100) subgroup. Both sexes are well represented. The species is well-supported (46.7/98/99) as sister to N. kalengonis n. sp . .

Variation and notes. Despite variation in chorion sculpture and length of collar, the widespread N. lujana is best identified by the egg in combination with the short curved SSt with longitudinal folds and some pale scales. The holotype (Kivu area, Fig. 18 ) and the female from Odzala N. P. ( Fig. 20 ) have the longest, while those from Bayanga ( Fig. 19 ), Bioko, and Mt. Tonkui much shorter collars. In the last female, punctures on the operculum are coarse and deep and stand in polygonal patterns reminiscent of follicle cell impressions. Normally, punctation of the operculum is unordered.

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 11–15. Males in the N. lujana-complex. N. costata n. sp.: 11, male tergites 7–10, dorsal view; 12, artificially everted penis. N. lujana Navás, artificially everted penes: 13, from Cameroon, Etambo R.; 14, from Bayanga (CAR). 15, N. serrula paratype from Ghana (NEOP009).

FIGURES 16–17. Female genital situs: 16, N. lujana Navás, not to scale; 17, N. costata n. sp. (Liberia, near Tapeta). co, common oviduct; scl, sclerite in vagina; sd, spermathecal duct; SSt, spermathecal stalk; V, vagina.

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 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.