Neoperla brachyphallus, 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 : 85-87

37. Neoperla brachyphallus n. sp.

( Figs. 201–204 )

Holotype J: Republic of Liberia, Suakoko 10–20 March 1952 light trap C.C.Blickenstaff ( USNM-37 , pinned, abdominal tip in microvial in glycerin, penis in Euparal, on pinned plastic slide).

Additional material studied: 1♁: Republic of Liberia, Mt. Coffee , IV-1897 Mrs. Sharp (slide USNM-146 , genitalia in microvial) .

Habitus. Small, WL 8.3mm, specimen from Mt. Coffee only 7.3mm. Body entirely ochre, except a brown spot connecting ocelli. Wings infuscate.

Male ( Figs. 201–204 ). Antecosta T7 connected to a short transverse sclerite separated by a pale zone from the squarish caudal sclerite with a long, sharply pointed triangular process ( Fig. 201 ). Sclerite on T8 T-shaped, with forward-inclined hump ( Fig. 202 ). Processes hinged in the intersegmental fold, antecosta 8 modified, with short anterior apodemes (resembling Fig 113 ). Antecosta of T9 medially divided in front of a membranous field. Lateral pilose humps with some SB, median furrow without. Caudolateral base of HT10 large, with copious pilosity, mediobasal callus wide, almost square, process short, tip blunt. Sternites finely pilose, caudally with larger setae, mainly on S8.

Penis plump, tubular, narrowing caudally ( Figs. 203–204 ). Stout spines in a subterminal dorsal patch point towards base, ♁ from Mt. Coffee additionally with some lateral subterminal spines. Endophallus short, curved to ventral side, distally tapering rapidly. Dorsal side with many minute spinules near base. Caudally large triangular spines form a wide band tapering near midlength of endophallus. Concave ventral side bare in basal third, caudally with slender erect spines, size decreasing distally and changing to small slender sclerites on the flagellum-like apical section of endophallus. Tip bare, recurrent tube not noticed.

Female. Unknown.

DNA. No data.

Notes. Several species with a hump on T8 have subterminal patches of spines on the penis but none has a comparably short endophallus. The armature conforms with the general pattern of a wide dorsobasal spine band, a bare lateral stripe, and a more distal ventral band of spines. The presumedly conspecific small male ( USNM- 146) agrees in T7–T9, but the process of Ht10 appears more slender and is pointed. Eversion of the endophallus failed, specimen in poor shape, details not recognised.

Etymology. A noun in apposition alluding to the shortness of the endophallus, formed from the Greek stems brachys and phallus.

IV.3. The Neoperla spio -complex (~ clade G)

In males, a pale line of weak sclerotisation runs across the middle of T7 which caudomedially has a pyramid-shaped process, with SB near tip and along the sclerotised underside. Median sclerite on T8 raised, with a hump or a process, the penis is tubular and sclerotised.

In several males, especially in the N. burgeoni -complex where males can raise a median sclerite on T8, the antecosta of T8 has paramedian apodemes projecting into segments 7 and 8, between the apodemes remains a passage for the aorta ( Fig. 113 ). The structure acts like a hinge, longitudinal muscles from antecosta 7 and attached to the apodemes affect both T7 and T8: the ventro-median ridge of the process on T7 is exposed and directed backward, resembling a vertical keel on the caudal edge of T7. Comparable modifications of the antecosta 8 occur also in some other species but are less strongly developed.

Female S8 with unmodified caudal edge, well pigmented females with three brown sclerites forming a triangle with caudal tip (e.g., Fig. 215). In variants of this pattern, sclerites may be large and merge around a pale center resembling an anchor (e.g., Fig. 264 ). In several species the caudal sclerite is weak or absent and only the strong paired anterior sclerites remain (e.g., Fig. 213 ). Anchor patterns on S8 occur also in other species groups (e.g., Figs. 289 , 345 , 357 , 362 ). Females of the N. dubia - and the N. orthonema -complexes differ and have on S8 2 brown marks in a longitudinal row which may be difficult to see in pale specimens ( Figs. 323, 330 ).

The vagina is sometimes with spines laterally from the SSt-attachment, otherwise unmodified. The SSt is coiled and of variable length, its opening into the vagina is narrower than in the transvaalensis -group (e.g., Figs 226 , 236 , 244 ). The narrow concave edge of the coil lacks scales and is membranous. The bare edge widens next to the vagina, the length of the basal section varies, sometimes in contains longitudinal folds and seems to be extensible. The rest of the SSt has a dense cover of brown scales resembling roof tiles or fish scales ( Fig. 108 ).

Eggs have straight striae, sulci are about as wide as the impunctate costae (e.g., Figs. 214 , 339 ), punctation in the sulci without order, micropyles freely visible (e.g., Fig. 257 ). Most species have 25– 30 egg striae, but N. spio has only about a dozen ( Fig. 247 ).

The analyses of DNA sequence data support strongly (all-NT: 65.7/100/90) to very strongly (mt-NT) the monophyly of the N. spio -complex + N. orthonema- complex (clade G, including clade H; see Figs. 491–492 , 496 ).

Keys to species in the N. spio -complex

The spio -group was originally defined by morphology. The key covers all originally included species, regardless of whether membership in the group was subsequently confirmed by DNA (clades G + H) or not.

Males (unknown of N. gordius , N. spaghetti , N. sassandrae , N. benti )

1 T8 with a longitudinal sclerite that can be raised. In lateral view the raised sclerite resembles a kidney. A fine pale suture surrounds the button-like top of the T7 process in front. HT10 long, slender, regularly arched ( Figs. 205–208 )............ 2

1' T8 without movable sclerite or process, remainder variable. Species with narrow tube-like endophallus belong here....... 4

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 496. Partial (clades G, H) codon-model maximum likelihood phylogram based on 356 mitochondrial genome and COX1 DNA barcode sequences (all-NT). See Fig. 493 for common details.

FIGURES 201–204. Neoperla brachyphallus n. sp., ♁ holotype: 201, 202, terminal tergites in dorsal and lateral views, semidiagrammatic, tergites and hemitergite 10 are labelled; 203, penis with everted endophallus; 204, distal part of endophallus, enlarged. Only 203 to scale.

FIGURES 108–113. Neoperla spp., structural details.108, N. africana Klap., 109, N. kalengonis n. sp., 110, N. sjostedti needhami Lestage, all from the stream Kalengo: 108, 109 scales in the SSt. Close-ups of egg chorion: 110, three egg striae, the 2 rows of micropunctures in the sulci diverge around an island with a micropyle; 111, N. africana, 112 N. camerunensis (End.), chorion with 2 micropyle orifices; 113, processes of tergites 7 and 8, and modifications of antecosta 8 (AC8), a combination of inner structures (light grey), and of fine pilosity and sclerites (dark grey) on the surface of tergites; diagrammatical. The SEM microgaphs are courtesy of MPI of Limnology, PL̂n.

FIGURES 258–266. Neoperla funiculata n. sp.: ♁ holotype 258, 259, ♁ abdominal tip, dorsal and lateral; not to scale; 260, apex of penis with incompletely everted endophallus; 261, enlarged apex of endophallus; note that it contains 2 more rows of un-everted spines. Paratypes from Kalengo stream, D. R. Congo: 262, 263, contracted penis in lateral and dorsal views; 264, ♀ genital situs, vagina flipped back to expose pattern on S8; 265, SEM-micrograph of egg (courtesy of MPI for Limnology, Pl̂n); 266, detail of chorion. Unless differently indicated, scale lines are 100µm, other figures not to scale. FIGURES 267–271. Neoperla biserrata n. sp., male holotype. 267, 268, cleared abdominal tip in oblique dorsal and lateral views, not to scale; 269, 270, contracted penis in dorsal, lateral, and ventral views, scale line is 500µm; 271, detail of endophallus near base. Tergites, HT10 and cercus are labeled. co, costa; mi, micropyle; sa, spermatheca; SSt, spermathecal stalk; su, sulcus; V, vagina.

FIGURES 213–214. Neoperla burgeoni Navás: 213, ♀ genital situs, Central African Republic (NEOP213), the inset is the holotype of N. bredoana (stained with Safranin); 214, egg, Ghana 81, Banda-Nkwanta, 18–20.9.1965, lighttrap on savanna (Endr̂dy-Younga: 250). FIGURES 215–216. Neoperla beta n. sp. (Guinea, r. Milo at Kankan, slide Z16.26), female genital situs and egg. In Figs 213 and 215, the vagina was flipped back for better visibilty of S8. Situs and eggs to the same scales, respectively.

FIGURES 289–293. Neoperla usambara n. sp. and N. sambarua n. sp., ♀ ♀. N. usambara: 289, genital situs, vagina and the coiled spermathecal stalk flipped back to expose anchor pattern on S8; 290, egg; 291, detail of striae with a micropyle in a sulcus (full arrow). N. sambarua: 292, genital situs with diagrammatic indication of border between S7/S8; 293, egg, variable thickness of chorion indicated by inner contour on right hand side. Open arrows show divergent spinule bands at entry of SSt into vagina. Scales as indicated. A, anchor; c, collar; co, costa; ls, lateral sclerite; mi, micropyle; mic, micropylar canal; sa, spermatheca; SSt, spermathecal stalk.

FIGURES 344–350. Neoperla lineata n. sp., female holotype: 344, fore body; 345, genital situs, vagina and SSt were flipped back to expose the anchor pattern on S8; 346, entry of SSt into vagina, arrow points at fine spicules; 347, egg; 348, count of costae on one quadrant of an egg cross-section (from a photograph); 349, detail of pale ring between striae and operculum; 350, micropyle (arrow) in the narrow sulcus between 2 costae, scale is 10µm; the other scales are 100µm. S8, sternite 8; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 351–361. Neoperla conradti (Enderlein): 351–354, male from Baré, abdominal tip and contracted penis; 355–356, male from Matadi, everted penis and detail near midlength of endophallus; 357, 358, ♀ genital situs and anchor pole of egg (from Bangem, Cameroon); the arrow points at spine patches in the vagina; 359, SEM, egg from holotype (courtesy of MPI for Limnology). Figs 353–355 to the same scale. Tergites, HT10 and cercus labeled; a, anchor; ac, anchor cavity; Mi = pores between costae above a concealed micropyle.

FIGURES 362–364. Neoperla muyukae n. sp. ♀ holotype: 362, genital situs, vagina and spermathecal coil flipped back to expose S8; 363, 364, egg and detail of chorion structure. mi, micropyle; od, oviduct; sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 323–331. Neoperla dubia-complex, femalesand eggs.N.dubia Klapálek:323, genital situs; 324, eggwith moderately raised costae (Ghana, Agumatsa waterfalls); 325–327, egg with flanged costae (D. R. Congo, Odzala N. P.). Neoperla proxima n. sp.: 328, genital situs (D. R.Congo, N of Matadi), 329, egg (Cameroon, Sangmelina). Neoperla vicina n. sp. (D.R. Congo): 330, female situs, 331,egg, arrow points at end of raised part of costa (paratype, Walikale, Z2101). Scales are 100µm, similar structures shown to the same scale. Top view of egg (326) based on combined halves of a photograph. Sa, spermatheca; SSt, spermathecal stalk; V, vagina.

FIGURES 223–226. Neoperla bipolaris n. sp.: 223, 224, egg, SEM micrograph (from Moore, 1991); 225–226, egg and genital situs of holotype. FIGURES 227–230. Neoperla schuelei n. sp.: 227, female genital situs, paratype; 228–230, egg from holotype; 229, enlarged detail of chorion; 230, oblique view of anchor pole showing the relation between striae and cells on collar. Scale lines are 100µm, unless differently indicated. Arrows indicate micropyles.

FIGURES 231–237. Neoperla nigricauda Navás: 231–234, male fore body, abdominal tip and penis (length 1.18mm), specimens from Cameroon, Etambo r.; 235, everted endophallus of pale ♁ from Kumba. 236, ♀ genital situs, arrow indicates the large spine patch in the vagina); 237, egg (Etambo r.).

FIGURES 244–249. Neoperla spio (Newman), ♀ and egg from Ghana, Subri R.: 244–245, genital situs and detail of vaginal spines; 246, detail of spines in spermathecal stalk (SSt); 247, egg, 248 detail of chorion, arrow indicates micropyle; 249, optical cross section of egg (from a photograph). Scales are 100µm, unless differently indicated. co,costa; sa, spermatheca; sp, spicules; SSt, spermathecal stalk; su, sulcus; V, vagina.

FIGURES 334–339. Neoperla spaghetti n. sp., ♀ holotype. 334, S8 with the anterior sclerites and the pale furrow (pf, arrow) between them. For better visibility the vagina (V) with the base of the spermathecal stalk (SSt) was flipped back. The overlong SSt (Fig. 335) broke in the process, a blue line connects the fitting ends of the fragments; 336, side view of an egg; 337, oblique view of an egg operculum; 338, anchor pole view of another egg, count of costae in white script; 339, detail of striae and micropyles, enlarged from 336. Scales are 100µm, 337–339 not to scale. co, costa; mi, micropyle; sa, spermatheca; SSt, spermathecal stalk; su, sulcus, v, vagina.

FIGURES 250–257. Neoperla bella n. sp., 250, ♁ abdominal tip, dorsal view; 251, abdominal tip of ♁ holotype with naturally fully everted penis; 252, contracted penis; 253, penis tip and everted endophallus, 254, distal end of endophallus; 255, ♀ situs, vagina flipped back to expose pattern on S8; 256, slide mounted egg; 257, detail of chorion structure, arrow indicates a micropyle.All from Kalengo stream, D.R.Congo. Not all figures to scale, scales are 100µm, except as indicated in 257. Tergites, HT10, cercus and S8 are labeled. ea, external asperities on penis tube; es, endophallic spines; SSt, spermathecal stalk; V, vagina; vd, vas deferens, visible through the delicate integument.

FIGURES 205–208. Neoperla burgeoni Navás, ♁: 205, 206, abdominal tip in resting position, dorsal and lateral view; 207, same, lateral, process on T8 raised, tergites and hemitergite 10 labeled; 208, medial view of detached right-hand HT10. Only 208 to scale, 205–207 to similar scales. at9, antecosta 9; fs, fine suture; mc, mediobasal callus on HT10; p7, p8, processes on tergites 7 and 8; p10, front part of hemitergite 10, its so-called process; ps9, paramedian swelling on T9.