Neoperla pilulifera, 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 : 76-79

30. Neoperla pilulifera n. sp.

( Figs. 169–173 , 181–185 )

Neoperla africana — Zwick 1973b: 382, figs. 1–2; partim; not africana Klapálek, 1909 .

Neoperla sp. 11— Zwick 1973b: 393, figs. 20–21.

Neoeprla sp. 10, sp. 11— Moore 1991: 5, figs. 8–11.

Type material. Holotype J, Central African Republic, SE Bayanga, 25-I-2012, UV: 17h30-2h, Nord M’boki, bras mort de la Sangha, S. Danflous et al.; 2,471 964 N 16,081 238 E ( NEOP087 , SMNS).

Paratypes (all SMNS). 1♁, Central African Republic, Bayanga, 15.10.2008 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 ( NEOP 088; abdomen in microvial on pin, penis on microscope slide IV-3); 1♀, C.A.R., SE Bayanga ACX7, NEOP 076; 1♀, C.A.R., SE Bayanga ACA3[3], 26.02.2012, NEOP 077; 1♀, C.A.R., SE Bayanga, LB-2, 2008, NEOP 079; 1♀, C.A.R., SE Bayanga IV[4], 2008, NEOP 081; 1♀, C.A.R., SE Bayanga AFO[2], 2.47N, 16.08E, 2012, NEOP 082; 1♀, C.A.R., SE Bayanga 148[1], 2012, NEOP 083; 1♀, C.A.R., SE Bayanga 148[2], 2012, NEOP 084; 1♀, C.A.R., SE Bayanga 145[1], NEOP 089; 1♀, C.A.R., SE Bayanga TK, 2.48N, 16.21E, 2012, NEOP 090; 1♀, C.A.R., SE Bayanga KT, 23.10.2008, NEOP 094; 1♀, C.A.R., SE Bayanga KT-6 2nd, NEOP 098; 1♀, C.A.R., SE Bayanga NB2, 22.11.2010, NEOP 100.

Additional material studied. 1♀, Koteh [spelling, location?] 20.1.25 Schouteden (Hynes det. 1957: N.spio ; slide Z74/10, gift Hynes, ex MRAC). Republic of Angola: 1♀, 54 mi N. Henrique de Carvalho [Saurimo, -9.6618, 20.3904], 3400 feet, 19-III-1970 E.S.Ross ( CASENT 8413113). Republic of Cameroon: Cameroon, Libamba [3.55°N, 11.10°E] 10km E of Makak J.A.Gruwell: 2♁, 9.III.1974; 1♁, 6-10-1973; 3♁, 9.III.1974; 1♁, 10–11-I-1974; 1♁, 11–16-XI-1972; 1♁, 16-III-1974 (all USNM). Cameroon, Libamba, J.A.Gruwell, 1♁, 12.– 13.5.1974, Slide 18.64 (ex USNM, SMNS; Fotos). 1♀ ( NEOP 093), 8♀, Cameroon 82 km NE of Obala [4°15’N, 11°53’E] 13- II-1973 J.A.Gruwell ( USNM). 1♁: 2♀: Cameroun, Nanga-Eboko [4.683 N, 12.367 E], 620m, 5.X.66, Ross & Lorenzen ( CASENT 8413032). 1♁, Cameroon, at light, Mamfe [5.75°N, 9.32°E], 4.3.75, Brinck & Enckell (Z17.010, MZLU). 1♁, Kamerun Bipindi Zenker [N 3.07824 E 10.41199] \ Neoperla camerunensis End. det. Dr. Enderlein (Slide Z19.07, MfNBerlin). Central African Republic: 2♀, C.A.R., SE Bayanga, 26-II-2012, camp lac 1, 2,480 322 N 16,215 608 E 401 m: ACA-1, NEOP 080; ACA3-2, NEOP 099; 2♀, C.A.R., SE Bazanga: RS-3, NEOP 092; KF, NEOP 095; KT-6, NEOP 096; APO_sana_2, NEOP 097. Democratic Republic of the Congo: 2♀: Congo (Zaire), Kisangani, Yoko [0.52°N, 25.20°E; 400m], 12.2.2008, LF, leg. V.Zolotuhin ( ZMB; slide ZMB _9A: NEOP 075). 2♁, 1♀: B. Congo, 20mi E Kizomba [5.05S, 17.25E], 13.8.57, Ross & Leech ( CASENT 8413033); 1♀: B. Congo, 39 km S of Walikale [1.47S, 28.07E], 700m, XII-25-57 E.S.Ross & R.E.Leech ( CASENT 8413036); 1♀: B.Congo, Luhoho R., Bunyakiri [2.07S, 28.57E], 1100m, 6.IX.57 Ross & Leech ( CASENT 8413035). 1♀, Musée du Congo Ituri [1.83306, 29.49808]: Kaporata 1-vii-1932 L. Burgeon \ Neoperla africana Klap. \ det. Navás S.J.; Neoperla leroiana Klap. det P. Zwick 2017 ( MNHN; pinned genitalia & egg slide). 1♀: Kongo, Kasai: Kamonia [-6.995, 20-963] 31, M.Delpierre; 1♀: Bokuma [- 0.666S, 21.016E], 2.4.41, D.Hulstaert; 1♀: Mongupa [2.366N, 20.7811E] (Lisala) 22.9.22, A.Collart; 1♀: Stanleyville, Yalemba [1.1700N, 23.842E], 8.28., A.Collart; 1♀ ibidem, 8.8.47, M.Poll; 1♀: Bassin Lukuga [-5.734, 26.9511], 4.7.34, deSaeger; 1♀: Bassin Lukuga [-5.734, 26.9511]; 1♀: Haut-Uele, Paulis [Isiro, 2.46N, 27.37E], 20.47, L.G.Benoit; 1♀: Ubangi, Nouvelle Anvers, 9.12.52, P.Basilewsky; 2♁: Coll. Mus. Congo Ubangi: Dongo 6-XII-1931 H.J.Brédo (all MRAC). Gabonese Republic: several ♁, Makokou [0.566202N, 12.867E] 227L Coiffait (ex Coll. Berthélemy, 1 SMNS); Belinga,1♁, 1♀, 21.3.63, 1♁ 17.5.63 (coll. Berthélemy). 1♁, Gabon Moyen-Ogooué Rég., Najolé, Auberge St. Jean, Ogooué R., 0°11’03’’S, 10°46’00’’E 16.12.1993 C.C. Jongkind ( ZMBN). Republic of Ghana: 1♀, Ghana Western region, Ankasa game prod. Reserve [5.2167N - 2.6499E] 10–16.12.93 St. 12 J. Kjaerendsen & T. Andersen light trap; NUFU-project ZMBN Ref. No. 3 ( ZMBN). Ghana, leg, Endrödy-Younga: 3♀, Banda-Nkwanta, N 8.22, W 2.09, 122 m, 1.– 8.9.1965, light trap, on savanna; 1♀, Bobiri Forest Res. [6.37N, - 1.38W], 13.11.1965 ( HNHM & SMNS, genitalia slides). Republic of Guinea: 1♀, Loffa (aff. Diani) W de Boola [8.19°N, 8.67°W], 8.3.88 ( SMNS); 1♀: Guinée Française, Kankan, Aout 1937, L. Berland ( MNHN). ♁: Guinea Milo à Boussoulé 23.10.1984 J.M.Gibon. 2♁, piège lumineuse, sans étiquette ( SMNS, gift Élouard; slide Z16/66; 1♀, Guinea Samou, Grde Chute; PL 215 13.02.1986 (Z16.195). République de Côte d’Ivoire: 1♀: Côte d’Ivoire, Bouaké [7.669N, - 5.020W], 21.8.66, Thys & Verheyen ( MRAC). Republic of Liberia: 1♀, Liberia, 8 mi NW Belefuanai [7.26N, 9.43W], S fork S.Paul R., 11.8.66, Ross & Lorenzen ( CASENT 8413034, slide Z18.019); 1♀, 1♀: Liberia, 36 mi S Voinjama [8.42N, 9.75S], 13.8.66, Ross & Lorenzen (all CAS). ♀: Liberia, Mt. Coffee, IV-1897 R.P. Currie ( USNM _136); 1♁, same locality, III-1897, Mrs Sharp ( USNM _147). Republic of Mali: 1♀: Mali, Bafing bei Tinko, N 13°29' 54'' / W 10° 45' 19'', Lichtfalle: 19:45–21:45 28.08.1991 W. Tobias. Federal Republic of Nigeria: 1♁, Nigeria, Samaru [11.1284N, 7.6329E], P.H.Ward, mercury vapour light trap, 6.– 23.8.1970; 1♀, the same, but 24.–30.8.70 (both Brit. Mus.). 1♀: Ifon Ondo Prov. [6.82, 5.77] VII.1-11.34 Nig. \ CNHM \ Straus-F. Mus. W.Afr.Exped.\ Col. by R. & L. Boulton ( CASENT 8413037). Republic of Senegal: 1♀, Haut-Sénégal, Niger, Fabien Girand, Muséum Paris \ Ochthopetina haugi Nav. Navás S.J. det. ( MNHN).

Habitus. WL of males 8.5–9.5mm, of females 11.0– 19.7mm. A pale light ochre species, the dark ocellar spot sometimes continues along the occipital suture. Tips of antenna and cercus bronish. Femora light, basal 4/5th of tibiae and the tarsi infuscate, male metatibia widened little.

Male ( Figs. 169–173 ). Tergites similar to N. amoena but process on T7 sharply pointed ( Figs. 169, 170 ). Penis thin, approximately 3mm long when endophallus is everted. Tube curved dorsad, sides near midlength with a membranous window with external baso-lateral spines (bl) ( Fig. 172 ). Membranous zone between tube and endophallus with small groups of disto-lateral spines (dl) plus a larger disto-dorsal (dd) spine group ( Figs 171– 173 ). Everted endophallus as narrow as penis tube, curved ventrad, diameter decreasing distally. Basal fourth bare, then follows a dorsal armature of scattered slender large hooks. Distal hooks are more slender, smaller, and closer together, the long narrow end is bare. Ventral side only with few small spines.

Female ( Fig. 181 ). S 8 in some exemplars with an indistinct anchor-like pattern, caudal edge unmodified.About 10 transverse folds across base of vagina, widest and most distinct close to genital opening. Vagina calyx-shaped, unmodified. The SSt forms a complete ring with long bare inner edge, remainder completely covered with scales. About distal third of coil narrow, spermatheca curled.

Egg ( Figs. 182–184 ). Ovoid ( Fig. 182 ), but some females with eggs whose operculum is narrower than the anchor pole. Average size 354*250µm, including the 25µm long collar. Body of the egg (excl. collar) on average 1.32 times longer than wide. Collar wide, surrounded by dense row of punctures, with deep smooth cells ( Fig. 183 ).

DNA ( Figs. 491–492 , 494). A total of 22 specimens from Cameroon, the Central African Republic, and the D. R. Congo were sequenced, representing only a small part of the wide distribution of this species. The sequences include the male holotype from the Central African Republic, for which only a COX1 DNA barcode fragment was obtained, and 21 female specimens, many of which are paratypes. The sequences form a tight cluster, and the monophyly of the species is maximally supported (100/100/100). The species is very strongly supported (98.2/99/99) as sister to N. aethiopica n. sp . .

Notes. Some females are very large and also much larger than males in the same sample. Length of collar varies, probably regionally, from wide and short with a single ring of cells (e.g., Fig. 183 , fig. 8 of Moore 1991) to longer telescoping collars with several rings of cells of stepwise decreasing diameter ( Fig. 185 , fig. 10 of Moore 1991).

Distinction of females from N. occulta n. sp. is difficult. The spermathecal coil seems to be shorter and stouter than in N. occulta , attempts to quantify differences were unsuccessful. Eggs of N. pilulifera n. sp. seems to be a little larger, eggs of N. occulta tend to have parallel sides, like a short cylinder. The N. pilulifera n. sp. collar width corresponds to approximately17% of egg width, more than of N. occulta n. sp..

Enderlein mistook a male of N. pilulifera n. sp. in the Naturkundemuseum Berlin for N. camerunensis ,which he had named from females many years before. The true male N. camerunensis are described here for the first time.

Etymology. The name (Latin, a pill carrier) alludes to the many smooth eggs found in gravid females.

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 169–176. ♁♁ in the Neoperla pilulifera-complex. N. pilulifera n. sp.: 169, 170, abdominal tip, dorsal and lateral, semi-diagrammatic, tergites and hemitergite 10 labelled; 171, contracted, 172, everted penis; 173, distal half of penis tube and base of endophallus (holotype ♁): all from Bayanga. N. crustata n. sp.: 174, 175, ♁ from Mali, Niger at Bamako; by transparency of the slide mount the basal end of the distodorsal spine group seems to overlap with the far distally displaced basolateral spine group (encircled); 176, holotype from Guinea (NEOP073). 174 and 178 to the same, other figures not to scale. bbs, bare basal stretch; bl, basolateral, dd, distodorsal, dl, distolateral spines.

FIGURES 181–192. ♀♀ of the Neoperla pilulifera-complex. N. pilulifera n. sp.: 181, genital situs, ♀ from Ghana, Ankasa Reservation; 182, egg of same, arrows show micropyles; 183, 184, collar and micropyle of ♀ from Gabun, Belinga; 185, “teleskoping” collar of ♀ from Schouteden, Koteh / Belinga. N. occulta n. sp.: 186, 187, genital situs and egg, Central African Republic, Bayanga. N. nichollsi n. sp., 188–192: 188, 189, genital situs and egg (Liberia, 8 mi NW Belefuanai, S fork S.Paul R., 11.8.66, Ross & Lorenzen diagrammatic, from photographs) (NEOP057); 190–192 are anchor pole regions of an intact and of 2 teratological eggs, all from the same ♀. In 186–188 ♀ situs and eggs to the same scales, respectively. The scale in 181 is 100µm, others figures as indicated, or not to scale. a, anchor; od, oviduct; rd, receptacular duct; sa, spermatheca; SSt, spermathecal stalk.