Neoperla panafricana, Zwick & Zwick, 2023

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 : 64-67

24. Neoperla panafricana n. sp.

( Figs. 133–142 )

Type material: Republic of South Africa: Holotype ♁ ( NEOP158 , penis slide), paratype ♀ ( NEOP156 , situs & egg slides: RSA/15 E Cape 60–90m - 31.4465S / 29.7372E Mbotyi Forest , Mkozi River, beating 14– 15.11.2013 ( SMNS, gift R.Ruta). 2♁, p aratypes, RSA, 15.12.95, Legalameetse [24.20, 30.31; 1100m], leg. F.Koch (MfN; NEOP154 , NEOP155 ); 13♁, 5♀, paratypes, Prov. Limpopo, Waterberge, Mapote Farm, 22.–24.Feb.2012, 24°07’51.41”S 28°38’40.88”E D. Bartsch ( NEOP136 , NEOP137 , NEOP138 , NEOP139 , NEOP140 , NEOP141 , NEOP142 , NEOP143 , NEOP144 , NEOP145 , NEOP146 , NEOP147 , NEOP148 , NEOP149 , NEOP150 , NEOP151 , NEOP152 , NEOP153 ; SMNS). Central African Republic: 1♀ paratype ( NEOP129 ): 06-III-2012, nuit, chasse de nuit (18h00-00h30): banc de sable au milieu de la Sangha, M’boki (sud village Likembé, bord Sangha), [ca 2,471 964 N 16,081 238 E]. Republic of Namibia: Kunene R., Epupa Falls [17.00S, 13.25E], 1♁, 1♀ ( NEOP133 ), 20.04.1994, W.Mey. Republic of Senegal, 1♁ paratype ( NEOP131 , Z16.89), 1♀ paratype ( NEOP130 , Z16.159), Touba [14.87°N, 15.88°W] août 1982 (gift J.M.Élouard, SMNS).

Additional material: Central African Republic, near Bayanga: 1♀ ( NEOP 135): 15.10.2008 18h–5h 03°03’58.3” N 16°08’59.6” E 528 m Sur plate-forme à 54 m du sol dans un Ayous ( Triplochiton scleroxylon , Sterculiaceae ) à 50 m du camp. 2♀, near Bayanga, sur plate-forme à 40 m du sol dans un kungu, à 50 m du camp 23.10.2008 18h–6h 03°01’49.5” N 16°08’31.7” E 567 m ( NEOP 134). Republic of Angola: Angola 20.XI.13, Malanje Prov.. 2km N of Socqueco, 994m, P. Schuele leg. -16- ( NEOP 132). République de Côte d’Ivoire: 15 ♁, riv. Férédougouba (bassin Sassandra [4.9433, -6.0850]) 12.08.1982 J.M.Gibon ( SMNS). Republic of Guinea: 4 ♁, Guinea, Light trap, no other collecting details ( SMNS, Slide 18.104, NEOP 335, gift Élouard). Republic of Namibia: Kunene R., Epupa Falls [17.00S, 13.25E]: 4♀, 20.02.1994; 2♀, 21.– 23.2.2008; 2♁, Namibia, Kunene River, Swaartboisdrift, 28.XI.2000 (all leg. W.Mey; MfN & SMNS). 1♁, CSIR Water Research AML218 \ Epupa falls Kaokoveld, S.S. Africa, 26.4.1970, Brown taken at light (gift J.Illies, slide Z16/005); 1♁: Kunbu-[illegible] Kavango 2.1.50 \ leg. F.Gaerdes ( SMNS). Republic of Senegal, 3♀, Touba [14.87°N, 15.88°W] août 1982 (gift J.M.Élouard, SMNS). Republic of South Africa: 3♀: RSA, Transvaal, Pullen Farm, 23.–24.XI.2004, W.Mey [near Nelspruit, 25.48°S, 30.97°E] ( MfNB & SMNS). 1♀ Kwa Zulu-Natal, Mooi R. [-29.21, 30.00, 1383m] (coll. Brinck). 2♁: R.S.Africa: Cape, 8 miles W Humansdorp [-34.04, 24.74], 1.3.1950, Brinck & Rudebeck (slides Z21.12 + 13). Republic of Zimbabwe: 2♀, Glendale [-17.362, 31.062] s.r. 26/12/56 \ C.N.Smithers [print]; 1♁: C. N. Smithers \ Beatrice [-18.27, 30.87] S. R. 26/12/56 ( SMNS). United Republic of Tanzania: Nyassa-See, Langenburg [Lake Malawi, Tukuyu], Fülleborn S. 1♁ 16.VII.-8.VIII.98, 1♀: 20.VIII.- IX.[18]98 ( MfNB).

Habitus. Moderate to large size, WL 11.7–17.7mm, mean 12.5mm in males, 15.3mm in females. Similar to other members of the complex, body light ochre.

Male ( Figs. 135–142 ). Similar to N. africana , but process of T7 shorter, tip less sharp. HT10 short, curved little, moderately pointed, mediobasal callus tongue-shaped ( Figs. 135, 136 ). Penis slightly sinuous, apex curving ventrad and truncate, the dorsodistal notch with small spinules is U-shaped and well delimited. The everted endophallus about as long as the penis ( Figs. 137, 139 ), a regularly curved tapering tube, base wide. Convex face and side with many strong hooks, laterobasal hooks and those near tip smaller and stouter, concave ventral side of endophallus bare (e.g., Figs. 138, 141 ), or with occasional single spines ( Fig. 140 ).

Female ( Figs. 124 , 133–134 ). S8 structurally unmodified, with three brown spots or with a pale center in a heart-shaped brown macula. The vagina is supported by sclerotised lateral folds: near the gonopore the folds are straight, parallel and stand close together, in front they curve outward and surround a wider area, the ensemble resembles a racket or a calyx ( Fig. 124 ). SSt short and forming only a ring fragment, the modified basal section with longitudinal folds and scales only on the convex side is long. The scale cover widens caudally, often appears a projecting angle (ap in Fig. 134 ).

Egg ( Figs. 133–134 ). Slender ovoid, size varies, 370–445 * 190–240µm, on average 410*218µm and 1.9 times longer than wide. Nine to 11 straight low ridges blend into the base of the operculum. The operculum is a blunt cone or almost rounded. The well-developed collar bears cells. Chorion punctures are as fine as in N. camerunensis .

DNA ( Figs. 491–492 , 495 ).A total of 30 specimens from Senegal, the Central African Republic, Namibia, Angola and the Republic of South Africa were sequenced, representing most of the geographic spread of this species and providing very strong support (98.2/100/97) for a species complex that also includes N. camerunensis (Enderlein) . While sequence variation is distinct and multiple lineages are indicated, specimens from different localities within the Republic of South Africa differ little in their DNA sequences and cluster with similarly looking specimens from Namibia, Angola, and Senegal, but also N. camerunensis (Enderlein) . Both sexes are well represented.

This species and N. camerunensis (Enderlein) form a complex that cannot be resolved with current molecular characters and should be investigated further with a much broader range of nuclear markers than just the mitochondrial COX1. The male holotype from the strikingly homogeneous South African population was sequenced with the genome-skimming approach and obtained 11,001bp of mitochondrial, protein-coding genes. This sequence is clearly different from the sequences of specimens collected in Cameroon, and the disagreement between molecular and morphological data is unlikely to affect the validity of both species but will probably require the description of further species in future.

Notes and variation. The characteristically shaped vaginal sclerites are an apomorphy for N. panafricana . The range is disjunct and there is no evidence of contact between the northwestern and the southern populations. Neoperla panafricana is the only South African representative of the africana -complex, Picker’s figures of eggs suggest his N. spec. C ( Picker 1980, his figs. 13, 16) is N. panafricana . The species is uniform across the huge, divided range, only the 2♁ from the extreme SW (Humansdorp) differ in that there are only very few tiny spinules in the dorsodistal notch of the penis tube.

Etymology. The Latin adjective panafricana refers to the wide distribution of the species over much of the Ethiopian Region.

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 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 131–134. Neoperla spp., female situs in the top row, an egg of the same specimen in the bottom row: N. camerunensis (Enderlein):131, the lectotype (situs after Zwick 1973a); 132, ♀ from Cameroon, Libamba. N. panafricana n. sp.: 133, Namibia, Kunene R. (NEOP133); 134, Senegal, Touba (NEOP130). All to similar scales. ap, angular process; lf, longitudinal folds; od, oviduct; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 135–142. Neoperla panafricana n. sp., male abdominal tip and penis with everted endophallus: 135–137, holotype from Mkozi R. NEOP158; 138, Guinea (no details); 139, Namibia, Kunene R.; 140, R. S. Afr., Legalameetse, NEOP154; 141, Ivory Coast, Férédougouba; 142, Senegal, Touba, NEOP131. The scale is 200µm, all penes to similar scales.

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.