Neoperla sjostedti sjostedti Klapálek, 1909

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 : 161-164

78A. Neoperla sjostedti sjostedti Klapálek, 1909

( Figs. 443–449 )

Neoperla sjöstedti Klapálek, 1909a: 218 .

Neoperla sjöstedti — Klapálek, 1910: 55, fig. 1.

Neoperla sjöstedti — Klapálek, 1912: 447.

Neoperla sjöstedti — Klapálek, 1923b: 134, fig. 6.

Type material studied. Republic of Kenya: Holotype ♀: Kilimandjaro, Kibonoto, Kulturzone 1300–1900 m 1905 Yngue Sĵstedt. The specimen is in alcohol, the detached cleared abdominal tip in a microvial inside the main tube (Mus. Stockholm; studied in 1971).

Additional material studied: Republic of Kenya: 3♀: Kenya, Kakamega Rain Forest Reserve [0.29°N, 34.86°E; 1500m], 12.II.1999, 1450m, leg.Deckert/Göllner (MfnB & SMNS, slide Z21.01). 1♁, 2♀: van Someren, Nyeri [0.25°S, 36°57’E] ( BMNH). Republic of South Africa: 1♁: Univ. Pretoria: SE24,30 BU V.1976, E. A. Boomker (Uni. Pretoria, glued on a card, cleared genitalia in glycerin in a pinned microvial). 1♁: SE 24,30 BU V.1976 E. A.Boomker \ Neoperla spec . aff. leroiana Klap., Zwick det. 1982 (Univ. Pretoria, glued on card [originally pinned], genitalia in microvial); 1♁: Natal (penis slide); 1 penis slide, S. Africa, Malta Forest [~ 27.26E, 25.81S] 4/12/76 \ M.Picker det. sp. C ( SMNS, gift of M.Picker). 2♁, RSA/15 E Cape 60–90m - 31.4465S / 29.7372E Mbotyi

Forest Mkozi River, beating 14– 15.11.2013 leg. R. Ruta (slide Ruta ♁, NEOP 309; SMNS). 1♀, Cathedral Peak, Natal area South Africa Febr. 1954 B.Stuckenberg (Slide Z 17-004, SMNS) . 12♀, Cape Peninsula, Cape Point Nature Reserve 10.12.1950 (B&R 79) ; 1♀, Cape prov., George [-34.12, 22.43], 27.2.1951 (B&R 188) ; 1♀, Butterworth , 50 mls N East London, 4.3.1951 (B&R 204) \; 1♀, Cape, Lundean‘s Nek [Lundin‘s Nek] 20mls NW Rhodes 11.3.1951 (B&R 229); ca 50♀, Natal, Royal Natal Ntl Park, The Hostel , 3.4.1951, 3.4.1951 (B&R 259) ; 13♀, Royal Natal Ntl Park Tugela Valley 4.4.1951 (B&R 261) (all Brinck & Rudebeck, MZLU). Republic of Zimbabwe: 1♁. S. Rhodesia Cashel Dist. [-19.56, 32.79], Umvumvumvu R. 18.12.1954 C.N.Smithers ( SMNS) . 1♀, by stream eastern slopes of Vumba Mts [-19.15, 32.73] S. Rhodesia 22.12.58 C.N.Smithers (slide Z17-014) ; 1♀: Salisbury, S. Rhodesia [Harare, -17.87, 31.00]), 23.3.58, C.N.Smithers, at light (slide Z17-016) ; 1♀: N. Rhodesia Livingstone [-17.824, 31.050] 17.5.1951 Brinck & Rudebeck #309 (Slide Z 17-008) ; 1♀: Salisbury Expt. Stn. [Harare -17.87, 31.03] Light Trap 6. March 1957 ( MZLU) . 3♀: Rhodesia, Victoria Falls [-17.83, 25.83], 900m, 11/ 12.X.1966 Ross & A.R.Stephen ( CAS). Republic of Zambia : 1♀, Copperbelt Province, Kitwe Dist.; Kumasamba Lodge near Kafue R. abt. 5km S Kitwe; S12°54’20’’, E 28°14’22.3’’ 18–30 April 2006, G. Svenson ( BYU; slide Z18.74) .

Habitus. WL 10.5–11.5mm in males, 13.5–17.2 mm in females. Body yellowish to light ochre. Ocelli one diameter apart, in a dark 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. 443–446 ). A slightly elevated caudal sclerite occupies the median third of T7. Its corners are rounded, the area between them is concave, saddle-like, smooth, with a low transverse fold (arrow in Fig. 443 ) behind which stand several SB. A sclerite with scattered SB curves down into the intersegmental fold. Median sclerite on T8 band-shaped, with a forward-curved hook with some SB on front and at tip Fig. 444 ). T9 unmodified, with paramedian pilose humps and median furrow. HT10 process variable, short, sinuous, mediobasal callus large, squarish.

Penis a caudally narrowing tube, distally soft and annulate. Near midlength, there may ventrally be a transverse fold (arrow in Fig. 458 ) or a rudimentary caecum ( Fig. 445 , black arrow). The everted endophallus is about twice as long as the penis tube ( Figs. 445–446 ), with an armature of small spines in 1–3 rows on both the dorsal and ventral sides. Distally, spinules grow smaller, they disappear near the tip.

Female ( Figs. 447, 449 ). Caudal edge of S8 with an often indistinct short transverse pale subgenital plate. Vagina unmodified, without spines. SSt very long, a wide coil of 3–5 rings, the basal half is bare except for a narrow row of scales on the convex side (Fig, 447). Near midlength the scale row widens gradually until it covers most of the SSt diameter but a short section before the curled transparent spermatheca is bare ( Fig. 449 ).

Egg ( Fig. 448 ). Average size 354*178µm. Slender, club-shaped, operculum wide, with distinct cells. Collar with two rings of cells. There are about 35 straight striae, the narrow sulci with two rows of micropunctures which diverge around the micropyles. The anchor cavity is wide and deep, the mushroom-shaped anchor has a solid stem.

DNA ( Figs. 491–492 , 498). Only a male specimen from the Republic of South Africa was sequenced for the COX1 DNA barcode fragment. This sequence is distinct from and near maximally supported (99.5/100/100) as sister to the remaining sequences in this species complex. The entire species complex has maximum support (100/100/100) as sister to N. socia n. sp..

Notes. The holotype is completely faded, even the eyes lack pigment. Staining with Safranin ( Ludwig & Schmidbauer 1967) was unsuccessful, but a drawing of the female genitalia was nevertheless possible ( Fig. 449 ) but there are no eggs. The combination of a short pale transverse subgenital plate with a very long largely bare SSt is characteristic of N. sjostedti . Females of the two subspecies cannot reliably be separated but the type locality is east of the Rift Valley from where only males with rudimentary penis caecum are known ( Fig. 458 ).

The correct spelling of the name is sjostedti ( ICZN 32.5.2.1). Klapálek had submitted a detailed description in 1908, in the same manuscript he also proposed another new species, N. africana . However, the paper was published only in 1910 (from the journal’s archives: the late P. Brinck, Lund, in a letter). In the meantime, the names had been published ( Klapálek 1909) in a key to known African species of Neoperla , an indication which makes both names available.

Klapálek (1910) associated a pinned male from Ukami with dark tibiae with the female type. He did so with doubts which excludes a type status ( ICZN 72.4.1). This male is N. tansanica n. sp., see above. A female from Daressalam in coll. Klapálek in Prague was also labelled N. sjostedti . However, it was not mentioned in Klapálek’s papers and therefore has also no type status. The specimen is N. sambarua n. sp. (see above—#51). Klapálek’s comparisons with other species and suggested synonymies ( Klapálek 1912, 1923b) are unsubstantiated.

The disposition of spines on the endophallus differs from most other African species. Usually, there are many more spines or longer spine bands dorsally than ventrally, the opposite is the case here. A rather uniform small size of spines is not often observed (e.g. in N. socia n. sp.) The long ventral spine band on the endophallus comprises several rows of small teeth but the male from Zimbabwe has a single row of slightly larger teeth, like N sjostedti needhami .

Picker (1980) illustrated one male of his sp. C “in normal resting position” (Picker’s fig. 23, left hand side). We everted the endophallus of this particular specimen, it is N. sjostedti with a vestigial caecum ( Fig. 445 ). The presumed female of sp. C of Picker (1980) is N. panafricana n. sp.. Picker’s material also included N. burgeoni , as we now noticed.

Neoperla sjostedti sjostedti occurs east of the Great Rift Valley, from Kenya to the eastern coast of South Africa, inland to the Victoria Falls; there is a possible contact zone with N. sjostedti needhami 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 443–449. Neoperla sjostedti sjostedti Klapálek: ♁ from Pretoria: 443, 444, terminal segments in dorsal and lateral views; arrows indicate the low crest in the sclerotised area on T8, tergites, hemitergite 10 and cercus labeled; 445, penis of male from Mbotyi Forest, black arrow points at the rudimentary caecum, open arrows at the location of details; 446, penis, diagammatic, shades are spinose areas. Penes and details of armature to the same scales, respectively. Female from Kenia, Kakamega Rain Forest Reserve: 447, genital situs; 448, egg (not to scale). 449, ♀ Holotype, genital situs. sa, spermatheca; SSt, spermathecal stalk; V, vagina.