Neoperla sjostedti needhami Lestage, 1921

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 : 164-167

78B. Neoperla sjostedti needhami Lestage, 1921 , new status

( Figs. 450–456 )

Neoperla excisa — Needham 1920 (nec Klapálek, 1909), Bull. Amer. Mus. Nat. Hist. 43: 36, pl. 5, figs. 13, 15, 17.

Neoperla needhami Lestage, 1921 , Ann. Soc. Ent. Belg. 61: 153.

Neoperla bottegoana Navás, 1933: 109 , fig. 102. New synonymy.

Neoperla sp. 33— Zwick 1976a: 386, 387, figs 4, 8.

Neoperla species C — Picker, 1980, only the male.

Type material studied. J Lectotypus N. needhami , Democratic Republic of the Congo, on 2 microscope slides (Coll. Needham, Cornell University): 1: Both pairs of wings: Neoperla excisa Klp ♁ Faradié Congo [pencil, in Needham’s hand; [apparently Faradje, 3.73186, 29.70647]; Neoperla needhami Lestage, 1921 , wings of ♁ Lectotype design. P. Zwick 1980; 2, with two cover slips: A, all 3 pairs of legs; B, head [torn], pronotum, dorsal and ventral sides of abdomen [penis well visible, Figs ##). Label on front of slide: Cornell University [print] \ ♁ Neoperla excisa Klap. Faradie Congo” [pencil, in Needham’s hand]; label on reverse of slide: ♁ Neoperla excisa Klp Faradié Congo [pencil, in Needham’s hand]. Text scratched into the slide: Lectotypus ♁ Neoperla needhami Lestage, 1921 , design. P. Zwick 1980 (= N. bottegoana Navás, 1933 ). The previously unpublished lectotype designation is now validated, to stabilise nomenclature. We have not seen a female which Needham also had, it is an unavailable paralectotype. ♁ Holotype N. bottegoana : [ Federal Democratic Republic of Ethiopia]: ARUSSI GALLA [Arsi, 7.84N, 39.65E] A.Gamale III V.93 V. Bottego Tipo \ Neoperla bottegoana ♁ Nav. (Mus. Genua, labels in Navás’ hand; vidi X.1974).

Additional material studied. Republic of Angola: 2♀: Dundo [7°22’S, 20°49’E], Lunda prov., Angola 21.IX.1949 B.Malkin; 1♀: ANG. 230. Dundo, Piège lumineux 5.XII.47 (M) (slide Z18.03); 1♀: ANG. 3126.2 Dundo XI.48 (M) (slide Z17.80); 1♀: Angola, Dundo, Piège lumineux VIII.53 Luna de Carvalho (slides Z17.81; [♀ ist needhami!] 1♀: ANG. 3158.7. Dundo, piège lumineux 30./31. XII. 53 (L) (Z19.6); 1♀: Angola: 54 mi N. Henrique de Carvalho [Saurimo, -9.6618, 20.3904], 3400 feet, 19-III-1970 E.S.Ross ( CASENT 8413114). 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 308, Slide Z17.45). Republic of Cameroon: 3♁, 1♀: Cameroon, at light, Mamfe [5.75, 9.32] 4.3.75 ( SMNS, penis slide Z17.009, ♀ situs Z17.012). 1♁: Süd-Kamerun, Molundu [1.93, 15.20] XII.09, v.d.Marwitz; 1♁: Kamerun: Ngoko- Station 4.5.02 Hösemann S.G.; 1♀, same, except 9.4. [19]02; 1♀, Kamerun, Soppo [=Sappo, Buea, 4°9'N, 9°14'E], II.1912 leg. v. Rothkirsch (all MfNB). 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, A.Zwick ( SMNS: slide CAM_2011-34, NEOP 310; CAM_2011-43, NEOP 321; CAM_2011-43, NEOP 322). 1♁: Cameroun, Nanga-Eboko [4.683 N, 12.367 E], 620m, 5.X.66, Ross & Lorenzen ( CASENT 8413062; penis with large caecum in a microvial). Central African Republic: 1♀, Bayanga, 06-III-2012, nuit, chasse de nuit (18h00-00h30): banc de sable au milieu de la Sangha, M’boki (sud village Likembé, bord Sangha; Z16.08, NEOP 324); 3♁, 1♀: Rép.Centrafricaine, Bambari [5.73, 20.68] III.1964 G.Pierrard ( MRAC, 1♁ SMNS). Democratic Republic of the Congo: 1♀: Kongo, Odzala Nat. Park [1.19, 14.85], 29.9.– 3.3.1997 leg. V.Sinaev ( ZMB). 2♀, B. Congo, 39 km S of Walikale [-1.47, 28.07], 700m, XII-25-57 E.S.Ross & R.E.Leech; 1♀ B.Congo, Irangi, Luhoho R., 900m, 10.IX.57 Ross & Leech ( CAS). 2♀: O.Afrika, Ruwenzori [0.385, -29.872], Westseite, Febr. 1908, leg. Schultz ( MfNB). 1♁, Musée du Congo Haut-Uele [3.53, 28.31]: Moto IV-V-1923 L. Burgeon \ Neoperla camerunensis End. P. Navás S. J. det.; 1♁, Congo belge: P.N.A. Ishango [-0.136, 29.602] 9-XII-1933 Mission H. Damas: 462 \ N. Hynes det. 1959 Neoperla spio (Newman) [hdschr.]; Kongo, Lukavo (afl. dr. Lufira) 700m, 6.–9.10.47 Mis. G.F. de Witte Nr. 814A; 1♀ Kongo, Gandalika IV.1959 J. Decelle ( MRAC). 1♀: Urwald Beni Okt.10 Grauer \ Neoperla \ africana Klapálek [print] ( NHMW). Federal Democratic Republic of Ethiopia: 2♁, 1♀: 15km NW of Chora [8.98, 38.75], Illubabor Prov., 1600m, vii.1973, G.de Rougemont ( SMNS; slides z74/4, 16/103). 2♁ (penis in mv), 15♀: Cojeb river 9-11/4- 61 at light S.Chojnacki leg.; 1♀, ETHIOPIA: Valaga, Awash River (125km by road from A.A.) 18/6-61 at light S.Chojnacki leg.; 3♀: ETHIOPIA: Ghibe River (260 km) (at light) 6/5-61 S. Chojnacki leg.; 3♁,> 50♀: ETHIOPIA: 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) (gift P. Brinck). 1♀, Kaffa, tra Gimma e Cossa, Torr. Aueddu del Didezza e pozza, ~ 1650m, 24.X.1973, P. Brignoli leg.; 1♀, Ethiopia, Sidamo, Billa Motel, 18.xi.1973, A. Vigna leg. (both coll. Consiglio, Rome). Republic of Ghana: 3♁, 5♀: Ghana Western region, Ankasa game prod. Reserve 10–16.12.93 St. 12 J. Kjaerendsen & T. Andersen light trap; NUFU-project ZMBN Ref. No. 3; 3♀: Volta Region, Wli Falls, Afegame, St. 6, 8.3.93, Mus. Bergen; 21♁, 18♀: same locality, 11.03.93 (penis slides GH09/03, NEOP 314; GH09/04, NEOP 312; ♀ slide 98/15); 1♁, 2♀ ( NEOP 313): Ghana Volta region, Agumatsa waterfalls, Wli, St. 10 19.11.1993 H. Olsvik light trap; 3♀: Ghana Volta region, River Menu at Mensu 18.11.1993 J. Kjaerendsen light trap (all Mus. Bergen & SMNS). 3♁: Ghana, Volta Reg., Kute River [7.37, 0.61], Menu, 11.12.1990, J.S.Amakyé light trap ( SMNS); 4♁, 1♀: Ghana, Dodo Tamli, Asukawkaw R. 20–22.XI.85, light trap, J.Schorscher ( SMNS; slides Z86/1 & 7). Republic of Guinea: 2♀, Kolenté à Simbareya [9.89, -12.61], 3.2.89 ( NEOP 317, slide 16/60 & NEOP 315, Z16.62); 1♀, Milo à Kankan (Bassin du Niger) 26.10.1984 J.M.Gibon (slide 16.26) (all SMNS, gift J. M. Élouard). République de Côte d’Ivoire: 3♀: Ivory Coast: Cavally R. (blackwater) nr Taȉ [~5.87, -7.45] 02.07.1977 B.Statzner ( SMNS, Slide 86.16). 2♀: Ivory Coast Région de Grabo, petite rivière, route Grabo-Tiboto 11.02.1988 ( SMNS, ex coll. Élouard); 1♀: Côte d’Ivoire riv. Férédougouba (bassin Sassandra) 12.08.1982 J.M.Gibon ( SMNS, slide Z86/10). Republic of Mali: 1 pharate ♁: Tienfala [Tyenfala 12.66, -7.76] 24.06.1985 \ élevage #23 (gift J.M. Élouard, SMNS). Federal Republic of Nigeria: 2♁, Oban 7-iv-1975 J.T.Medler ( BYU).! ♀; Nigeria, Ibadan, 8.8.23 [ BMNH, London; egg slide SMNS]. Republic of Rwanda 1pharate ♀, S.W. Ruanda, Rukarara-Bach ca. 1800m, Exp. Herzog Ad. Fr. z.Mecklenburg 20.8.08 (egg slide Z19.16; ZMB). Republic of Senegal: 6♀: Gambie à Samédouta, 26.01.89 ( SMNS ex coll.Élouard, NEOP 316, slide 16/74; SMNS). Republic of Sierra Leone: 3♁ (Z16/79, NEOP 318), 1 ♀, Pampana près de Magburaka [8.67, -11.96], 5.2.89 ( SMNS); 12♁ (Z17/019, NEOP 319), 1♀ ( NEOP 320, Z16/151), Sierra Leone Seli á Bumbuna [8.91, -11.73] 10.02.1989. 1♀: Kaba à Gotamba, Kilims Nat. Pk. [9.716, -12.026] (Z16/103) 4.2.1988 ( SMNS, gift Élouard). South Sudan: 1 pharate ♀: Sudan, Wau [7.6868, 27.9853], Busseri Causeway, 14-ii-83 #4 j. Schorscher ( SMNS; slide DNA 511, NEOP 323); 1 pharate ♁, Sudan on rocks Ngohalima 24.iv.81 285 Schorscher ( SMNS; Z21.27). United Republic of Tanzania: D. O. Afrika Langenburg [Tukuyu, -9.26, 33.65, ~ 1500m] Fülleborn: 1♁, 3.11.1899; 1♁, 15. Nov. 99 (Z19.17); 1♀, 9.–19.VIII.98 Fülleborn S.; D.O. Afrika, Amani [-5.10, 38.63], 1♀, II.1907, Prof. Vosseler; 1♀, Amani, II.12., Dr. Chr. Schröder (all MFNB, in alcohol). Togolese Republic: 1♀: Togo, Amu Oblo [7.38, 0.86], 23.11.1985, V. Landa ( SMNS: slide 16/145). 1♀: Togo, Bismarckburg [8.1770, 0.6874], R.Büttner S. 20.vii–20.ix.1890 ( MfNB; egg slide also in SMNS). 1♁, 1♀, TOGO N 08°31.151’ E 000° 36.169’ 16 June 2005 KB Miller ( BYU; slide Z18.79). 1♀: Togo, Amoutchou, 28.11.1985, V. Landa ( NEOP 311, MfNB)

Habitus. WL 10.5–11.5mm in males, 13.5–17.2 mm in females.Body yellowish to light ochre, a dark interocellar spot, forehead in front of M-line often darker than rest. Appendages light, flagellum distally light brown. Legs light, knees and tibiae mostly slightly infuscate.

Male ( Figs. 450–454 ). Indistinguishable from N. sjostedti sjostedti by habitus and external male structures. Penis a caudally narrowing tube, distally soft and annulate. A large empty caecum (* in the figures, e.g., Fig. 451 ) with elastic surface projects ventrally near midlength. The everted endophallus is much longer than the penis tube and often strongly curved ( Fig. 454 ). Annulate penis tip and endophallus base with dorsolateral spinules which merge to a short dorsal band but only the spinule band on the ventral side is well developed. Ventral side of endophallus bare at base, distally follow triangular spines in a single row which continues to the tip.

Female and egg ( Figs. 455–456 ). Similar to the nominal subspecies, eggs indistinguishable. The brownish nail (n in Fig. 455 ) on the caudal edge of S8 has no internal crests and is often very pale and less distinct than in the figured specimen.

Variation. The length of the caecum varies but it is longer than penis width at the point where the caecum originates ( Figs. 451–454 ). The everted endophallus may curve in various directions and is always very long, regardless of caecum size. Possible differences in shape of the mediobasal callus of HT10 were not studied in detail. It is large and truncate in the type of N. bottegoana , yet about tongue-shaped in the N. needhami type. In addition, Ethiopian females have a projecting nail, a brown spot in front of it on S8 that makes the nail look longer than it actually is ( Fig. 455 ).

DNA ( Figs.491–492 , 498).A total of 16 specimens from Senegal, Guinea, Sierra Leone, Ghana, Togo, Cameroon, Central African Republic, South Sudan, and Angola were sequenced for the COX1 DNA barcode fragment and with the genome-skimming approach. Including an additional eight distinct female specimens from Cameroon (see below) and excluding the female specimen from Angola (NEOP308), the monophyly of this cluster is very strongly supported (99.3/100/100). Including the genetically distinct female specimen from Angola (NEOP308), statistical support for the monophyly of this group is not significant (47.4/85/73). This indicates that contrary to the best topology, the single representative of N. sjostedti sjostedi (NEOP309) might be closer to the other specimens than NEOP308 from Angola, which probably represents a distinct subspecies. As females of the named subspecies of S. sjostedti cannot be reliably distinguished, this female specimen from Angola is not formally described as a new subspecies based on DNA sequence differences alone.A similar situation exists for eight additional female specimens from Cameroon (see below), which are also not described as a new subspecies. More specimens, additional data, and careful examination would be needed to resolve this cryptic species complex. For now, we include all specimens under N. sjostedti needhami to ensure monophyly of the subspecies of N. sjostedti .

Notes. Neoperla needhami Lestage, 1921 was not replacing some other name ( Illies 1966: “ nov. nom. ”) but was a previously misidentified new species. The subspecies of N. sjostedti are allopatric, N. sjostedti needhami occurs north of the nominal subspecies in a wide east-west range from Ethiopia to the Senegal but also far into the eastern Congo area, with a possible contact zone with the nominal subspecies near Tukuyu ( Fig. 458 ).

Figures

FIGURE 458. Distribution map of N. sjostedti sjostedti and N sjostedti needhami. The type localities of N sjostedti, N. bottegoana and N. needhami are identified by the initials of the names. Other specimen records concern only males, the arrow indicates the possible contact zone between the 2 subspecies.

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 450–457. Neoperla sjostedti needhami Lestage and N. sp. aff. sjostedti. N. s. needhami: 450, 451, abdominal tip and contracted penis of the N. needhami lectotype; 452, ♁ from Pampana, Sierra Leone; 453, ♁ from Tukuyu, Tansania; 454, ♁ from Illubabaor, Ethiopia. Scales are 0.5mm, except for the details in 451; asterisks identify the caecum of penis. 455, 456, ♀ genital situs and an egg, Ethiopia, NW of Chora. N. sp. aff. N. sjostedti (by DNA) from Mentchum River, Cameroon: 457, ♀ genital situs. Scale is 100µm. Scales are 100µm; n, nail; sa, spermatheca; SSt, spermathecal stalk; V, vagina.