Neoperla excisa, Klapalek, 1909
publication ID |
https://doi.org/10.11646/zootaxa.5316.1.1 |
publication LSID |
lsid:zoobank.org:pub:BC922E16-2614-4F3D-AD82-87A845DE7E2B |
DOI |
https://doi.org/10.5281/zenodo.16763903 |
persistent identifier |
https://treatment.plazi.org/id/E12C876C-4AEB-FF07-FF4F-FE47FA0C0B60 |
treatment provided by |
Plazi |
scientific name |
Neoperla excisa |
status |
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V. The Neoperla excisa View in CoL and N. sjostedti -operational complex (~ clade J, clade K + L)
Species in this operational complex were from the beginning of our work distinguished from all other species groups because:
1. known males have no unpaired medio-caudal process of T7. In many T7 is instead concave between a pair of paramedian humps,
2. females with S8 with a projecting hard nail or with a sclerite resembling a fingernail occur only in the present assemblage, and
3. egg striae have narrow sulci which lack macroscopic punctures but have micropunctures visible only in transmitted light, at magnifications ≥ 200*. They form a single regular row on each side of a sulcus and separate it from the costa. Many eggs have spiral striae with species-specific direction of spin while most other species have straight or at best oblique striae.
Unfortunately, half of the included taxa are known in only one sex, morphology did not reveal how the species are interrelated.Accordingly, the keys which rely on morphology cover the entire operational complex.Arrangement of texts follows the strong molecular evidence (except for N. multiserrata sp. n. and the exclusion of node I).
Keys to species
Males
1 T7 is flat, a trapezoidal sclerite in the middle has a wide truncate caudal edge which is opposed to and only narrowly separated from the wide, transverse top of a high process on T8 ( Figs. 373 View FIGURES 373–385 , 386 View FIGURES 386–390 )........................................... 2
1' T7 and T8 different, the high process on T8 with narrow top, or hook-like........................................ 3
2 Tips of HT10 bisinuous, the hook-like tips often grasp and hold the T8 process; mediobasal callus parabolic ( Fig. 373 View FIGURES 373–385 ). Penis dorsally with subterminal and terminal spine groups ( Figs. 374–375, 385 View FIGURES 373–385 ). Endophallus dorsally with several rows of spines, ventrally with a single orderly spine row ( Figs. 376, 380 View FIGURES 373–385 )....................... 65 N. heideae and 66 N. larvata n. spp.
2' Tip of HT process not hook-shaped ( Fig. 386 View FIGURES 386–390 ). Penis with only one subterminal spine group. Endophallus dorsally with a wide spine field, ventral face bare ( Fig. 387 View FIGURES 386–390 )...................................................... 67 N. sorella n. sp.
3 T7 with a strongly raised wide caudal process, its hind corners diverge ( Figs. 365, 366 View FIGURES 365–372 , 486 View FIGURES 486–489 ).......................... 4
3' T7 without caudal process, concave between the blunt corners of a caudal bench-like elevation ( Fig. 422 View FIGURES 422–428 ), or between 2 low paramedian humps. T8 and penis variable.................................................................. 5
4 T8 with wide, forward-curved process which fits into the excavation of the process on T7 ( Figs. 365 View FIGURES 365–372 , 388 View FIGURES 386–390 ). The penis is a hard narrow long tube, the long narrow endophallus with fine short armature ( Fig. 368 View FIGURES 365–372 )................. 64 N. excisa Klapálek View in CoL
4‘ Process of T8 narrow, pointed, standing erect behind the bilobed projection on T7. Penis soft and short, endophallus with sharp spreading spines ( Figs. 486–489 View FIGURES 486–489 )............................................................. 83 N. sp. Afr_A
5 Penis tip bent like a knee, only part of the short endophallus with spines, several form a short crest on the ventral side ( Figs. 423–424 View FIGURES 422–428 )........................................................................... 74 N. crenulata n. sp.
5' Penis straight, distally annulated, the endophallus differs...................................................... 6
6 Endophallus with a dorsoventral armature of long groups or rows of spines (e.g., Figs. 440 View FIGURES 439–442 , 445 View FIGURES 443–449 )...................... 7
6‘ Endophallic armature dominated by basolateral rows of individual spines ( Figs. 482, 484–485 View FIGURES 477–485 ), dorsal and ventral faces are bare or have only fine spines............................................................................ 9
7 Penis straight, its annulated tip soft. Spinose part of endophallus not much longer than the tube, the wide basal half dorsally with a wide spine field, ventrally with a row or a narrow band of many small spines, the long and narrow distal section is bare ( Fig. 440 View FIGURES 439–442 ).............................................................................. 77 N. socia n. sp.
7' Entire penis tube moderately sclerotised. The spiny section of the endophallus is at least twice longer than the penis (e. g., Figs. 445 View FIGURES 443–449 , 452 View FIGURES 450–457 )............................................................................................ 8
8 Penis ventrally with an empty caecum as long as the penis is wide, or longer ( Figs. 451–454 View FIGURES 450–457 )................................................................................................. 78A N. sjostedti needhami Lestage View in CoL
8‘ No ( Fig. 446 View FIGURES 443–449 ) or a rudimentary caecum on the ventral side of penis, or only an oblique inconspicuous fold in its place ( Fig. View FIGURES 443–449 445).................................................................... 78B N. sjostedti sjostedti Klapálek View in CoL View at ENA
9 Top of T8-process wide, rounded or truncate ( Figs. 477, 479, 481 View FIGURES 477–485 ), even slightly concave under certain angles of view. Base of endophallus on each side with a short row of rapidly decreasing sharp teeth, distal part of endophallus consisting of long folded membranes without distinct armature.................................................. 82 N. leroiana Klapálek
9' Contour of T8-process triangular, top more or less pointed (Figs. 432,461), endophallus variable..................... 10
10 Endophallus (observed only by transparency) little longer than the penis, two orderly lateral rows of stout conical teeth of uniform size occupy the basal 2/3 ( Figs. 434–435 View FIGURES 432–438 ), no other armature present...................... 76 N. tangana n. sp.
10' Endophallus much longer than the penis tube, endophallus base on each side with an orderly row of individual large teeth which distally decrease rapidly in size......................................................................... 11
11 Basolateral rows of endophallic teeth continue for a considerable distance, much smaller teeth stand between them ( Figs. 460, View FIGURES 459–465 462).......................................................................... 80 N. arambourgana Navás ’ View in CoL
11’ Penis similar to N. leroiana , (i.e., the basolateral rows of teeth on endophallus are very short, comprising only about 12 sharp teeth ( Fig. 485 View FIGURES 477–485 )), the long distal part of endophallus composed of folded membranes without recognizable armature.......................................................................................... 81 N. dundoana n. sp.
Females
At the base of the SSt is usually a section of variable length that can presumably be much expanded, it has scales only along the convex face while the concave side is membranous, looks void but may have longitudinal folds. This basal section is not considered in statements like "entirely" or "completely coated with scales".
1 Caudal edge of S8 with a long nail (e.g., Figs 369 View FIGURES 365–372 , 378, 382 View FIGURES 373–385 ). SSt long........................................... 2
1' Caudal edge of S8 and SSt variable, possible nails are short.................................................... 4
2 The SSt is of similar width over its entire length ( Fig. 369 View FIGURES 365–372 ). Egg ovoid, without distinct striae ( Figs. 370–372 View FIGURES 365–372 ).............................................................................................. 64 N. excisa Klapálek View in CoL
2' SSt with wide base, the longer distal part narrow ( Figs. 377–378 View FIGURES 373–385 ). Egg drum-shaped, with levogyrous striae ( Fig. 379 View FIGURES 373–385 )..... 3
3 Nail tongue-shaped. SSt forms about 1 ring, entirely coated with scales, entirely of the same structure and directly attached to the curled spermatheca ( Figs. 377–378 View FIGURES 373–385 )..................................................... 65 N. heideae n. sp.
3' Nail conical, its base may be narrow or wide. SSt very long, consisting of an entirely scaly basal half and a very narrow distal part with scattered very small cuticular elements which is connected to the spermatheca ( Figs. 381–383 View FIGURES 373–385 )..................................................................................................... 66 N. larvata n. sp.
4 Caudal edge of S8 triangularly recessed, a short conical nail projects from the notch. Floor of vagina sclerotised, a massive inversely U-shaped crest meets an internal ridge along the antecosta 8. Egg with levogyrous spiral striae ( Figs. 397–399 View FIGURES 397–401 )......................................................................................... 70 N. dianae n. sp.
4' Different, eggs and S8 variable.......................................................................... 5
5 The center of the caudal edge of S8 with a distinct notch...................................................... 6
5' Caudal edge of S8 not notched, variable................................................................... 8
6 The bare notch is shallow but wide, from it a strong sclerite extends forward into the vagina. Egg drum-shaped, striae straight ( Figs. 418–419 View FIGURES 418–421 )....................................................................... 73 N. excavata n. sp.
6' Notch of S8 and egg different............................................................................ 7
7 Notch rounded, strong setae reach over it from in front. Egg slender, striae levogyrous, costae near the punctate operculum with pale swellings ( Figs. 402–404 View FIGURES 402–406 )............................................................ 71 N. gibbosa n. sp.
7' Caudal edge of S8 with two soft points flanking a crescent-shaped sclerite which is shifted a little forward towards the vagina ( Figs. 463–465 View FIGURES 459–465 ). Egg slender, ovoid, with levogyrous striae ( Fig. 466 View FIGURES 466–476 )..................... 80 N. arambourgana Navás View in CoL
8 S8 caudally long, parabolic, with an anchor pattern. SSt long, egg sulci spiral and very narrow........................ 9
8' S8 caudally truncate, not prolonged, SSt and egg variable.................................................... 10
9 Egg oval, the levogyrous striae continue onto parabolic operculum ( Figs. 362–363 View FIGURES 362–364 )................. 63 N. muyukae n. sp.
9' Egg drum shaped, operculum wide, smooth, sharply delimited against the dextrogyrous striae ( Figs. 357–358 View FIGURES 351–361 ).......................................................................................... 62 N. conradti (Enderlein View in CoL )
10 S8 with a pale anchor pattern inside a brown macula........................................................ 11
10' S8 without anchor pattern, variable...................................................................... 14
11 Egg with>35 straight striae, a pale ring at base of operculum. Micropyles on the bottom of very narrow sulci. Anchor with single stem ( Figs. 345, 347, 349 View FIGURES 344–350 ).......................................................... 61 N. lineata View in CoL n. sp.
11' Egg without pale ring, with fewer variable striae and with modified anchors: the anchor cap is supported by several delicate filaments instead of one stem........................................................................... 12
12 Egg striae are straight ( Fig. 302 View FIGURES 294–304 )....................................................... 52 N. orthonema n. sp.
12' Egg striae are dextrogyrous............................................................................ 13
13 The micropyles are located in the narrow deep sulci ( Figs. 306–308 View FIGURES 305–308 ).......................... 53 N. filamentosa n. sp.
13' The micropyles are located on top of narrow supplementary costae ( Fig. 312 View FIGURES 309–312 )................... 54 N. spironema n. sp.
14 S8 caudally with a short subgenital plate, or a projecting nail, or a sclerite resembling a fingernail.................... 15
14' S8 not distinctly modified, the SSt is of uniform width, approximately1.5-2* longer than the vagina................... 22
15 Caudal edge of S8 with short rounded projection that is a little thickened. The ribbon-like SSt is several times longer than vagina and S8 and appears void for almost half the length because there are only few scales along the convex edge. The elongate egg has wide blunt poles and straight striae.............................................. 78 N. sjostedti Klapálek View in CoL View at ENA
15' S8 not as above, the SSt is not nearly as long and has no strikingly long void basal section.......................... 16
16 SSt little longer than the vagina, egg not striate............................................... 68 N. simplex n. sp.
16' SSt 2-3 times than vagina and SSt, the egg is striate......................................................... 17
17 Compared to the vagina the SSt is wide and narrows caudally much. The straight egg costae are drawn out into high crests ( Figs. 441–442 View FIGURES 439–442 )............................................................................... 77 N. socia n. sp.
17' Compared to the vagina the SSt is narrow; it is of uniform width............................................... 18
18 Costae with knobs and little columns in alternating positions.................................. 74 N. crenulata n. sp.
18' The egg costae are unmodified, flat...................................................................... 19
19 The nail on S8 projects and is entire or distally minimally excised ( Figs. 470, 472–473 View FIGURES 466–476 )............................ 20
19' Nail on S8 notched, variable (Figs, 463–465, 468–469)...................................................... 21
20 The slender egg has a distinct collar and a conical operculum, the egg striae are levogyrous spirals...................................................................................................... 82 N. leroiana Klapálek View in CoL
20' The egg resembles the egg of N. sjostedti View in CoL View at ENA ( Fig. 456 View FIGURES 450–457 ) and has straight striae, the SSt is only moderately long and forms ~2 rings, there are no long void basal sections ( Fig. 457 View FIGURES 450–457 )................................ 79 N. sjostedti cf. needhami (by DNA)
21 Caudal edge of S8 curved forward into the excavate nail which is S8 shifted forward; egg plump, the wide operculum with small grooves, striae are straight ( Figs. 467–469 View FIGURES 466–476 )........................................... 81 N. dundoana n. sp.
21' Caudal edge of S9 with a deep median notch in front of which lies a crescent-shaped sclerite. The ovoid egg with narrow operculum and oblique striae ( Figs. 463–466 View FIGURES 459–465 View FIGURES 466–476 )......................................... 80 N. arambourgana Navás View in CoL
22 The eggs have straight striae........................................................................... 23
22' The eggs have levogyrous striae......................................................................... 24
23 Anchor pole of egg with a collar, the costae are asperous, like sand paper ( Figs. 437–438 View FIGURES 432–438 )............ 76 N. tangana n. sp.
23' The egg lacks a collar but has in its place a long projection; the costae are smooth ( Figs. 394–396 View FIGURES 394–396 )......6 9 N. claviger n. sp.
24 SSt almost twice longer than the vagina, wide, basal third with longitudinal folds, only a band of scales along the convex edge. The egg striae include the operculum and have extremely narrow sulci which do not widen around micropyles ( Figs. 429–431 View FIGURES 429–431 )........................................................................ 75 N. massevensis n. sp.
24' S8 unmodified, the SSt is shorter, thin, and basally not modified. The egg sulci are widened around micropyles ( Figs. 340– 343 View FIGURES 340–343 ).............................................................................. 60 N. bareensis n. sp.
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