Revision of the African Neoperla Needham, 1905 (Plecoptera: Perlidae: Perlinae) based on morphological and molecular data
Author
Zwick, Peter
0000-0002-6730-0011
pleco-p.zwick@t-online.de
Author
Zwick, Andreas
0000-0002-7532-1752
andreas.zwick@csiro.au
text
Zootaxa
2023
2023-07-17
5316
1
1
194
http://dx.doi.org/10.11646/zootaxa.5316.1.1
journal article
10.11646/zootaxa.5316.1.1
1175-5326
8154005
BC922E16-2614-4F3D-AD82-87A845DE7E2B
52.
Neoperla orthonema
n. sp.
(
Figs. 294–304
)
Type material studied:
Republic of Ghana
: J
Holotype
(
NEOP
171, slide Z17_037_sp_Ax \GH.10), Ghana
Volta region
, Agumatsa waterfalls, Wli, St. 10
19.11.1993
H. Olsvik light trap; NUFU project (
ZMBN
).
3♀
paratypes
, Ghana, same data as
holotype
(slides GH.10\2, GH.11\2,
ZMBN
).
1♀
Paratype
, Ghana
Volta Region
, Wli Falls, Afegame [
7.12°N
,
0.59°E
], St. 12
07.03.1993
J.S.Amakye & J. Kjaerendsen light trap; NUFU-project
ZMBN
Sample no. 1 (
NEOP
169, slide GH.09\1,
ZMNS
);
4♀
paratypes
, Ghana Western region, Ankasa game prod. Reserve [
5.44N
,
2.07W
,
47m
], 10–16.12.93 St. 12 J. Kjaerendsen & T. Andersen light trap; NUFU-project
ZMBN
Ref. No. 3 (GH.01/17,
NEOP
164;
NEOP
168, slides GH.01/6, /9, /16, /17;
SMNS
); 1♁
paratype
(
NEOP
170),
1♀
paratype
(
NEOP
165, slide GH12\1),
1♀
(Slide Z17.073) Ghana
Volta Region
, Wli Falls, Afegame, St. 10
08.03.1993
J.S.Amakye & J. Kjaerendsen light trap; NUFU-project
ZMBN
Sample no. 12; 1♁
paratype
, Ghana
Volta Region
, Wli Falls, Afegame, St. 12
07.03.1993
J.S.Amakye & J. Kjaerendsen light trap; NUFU-project
ZMBN
Sample no. 1); 1♁
paratype
, Ghana
Volta Region
, Wli Falls, Afegame, St. 12
07.03.1993
J.S.Amakye & J. Kjaerendsen light trap; NUFU-project
ZMBN
Sample no. 1 (
NEOP
166, slide 98\9,
SMNS
).
Republic of Cameroon
,
4♁ p
aratypes
: Muyuka [
4.7217N
,
9.6408E
;
305m
], Victoria Div., BRIT. CAMEROONS,VI-25-49 B.Malkin at Light (Slides Z17.021-023;
ZMB
&
SMNS
); 1♁
paratype
,
7 mi
S Ebolowa [
2.92N
,
11.16E
],
580m
,
15.–17-X-1966
Ross & Leech CASENT8413020 (penis slide).
République de Côte d’Ivoire
:
1
J
paratype
,
15 km
N Lakota [
5.86N
,
5.68W
]
23.Oct. 1971
white light trap J.A.Gruwell (
USNM
152) (
Figs 294–299
).
Togolese Republic
,
1♁
paratype
, penis in microvial,
2♀
,
paratypes
(Z16/131,
NEOP
167 and Z16/132,
NEOP
161;
SMNS
) Amoutchou,
28.11.1985
, V. Landa (Z16.132).
Additional material studied:
3♀
,
Ghana
, same data as holotype,
4♀
slides,
Ghana Volta Region
,
Wli Falls
,
Afegame
,
St.
6
11.03.1993
J.S.Amakye
& J.
Kjaerendsen light trap
(slides
Afr.Z
98\09,
Afr.
Z98\14; GH.03/3,
Ghana
♀
14;
SMNS
)
.
Ghana Volta Region
,
Wli Falls
,
Afegame
,
St.
10
11.03.1993
J.S.Amakye
, J.
Kjaerendsen light trap
(
♀
slides:
Ghana
♀
2,
Ghana
♀
4, Z17/73;
Ghana
♀
10;
1♀
(
Slide Z
17.073); 3♁,
4♀
,
Ghana Western region
,
Ankasa
game prod.
Reserve
[
5.44N
,
2.07W
,
47m
], 10–16.12.93
St.
12
J. Kjaerendsen
& T.
Andersen light trap
; NUFUproject
ZMBN
Ref. No.
3 (slides GH1\18–20); 1♁ (penis in mv),
2♀
(
NEOP163
)
:
Ghana Subri R.
[
5.22851
,
- 1.7481
],
05.11.1993
light trap
; NUFU project,
Uni Bergen.
4♁,
1♀
, GHANA,
Western Reg.
,
Ankasa Resource Res.
,
Nkwanta Camp
05°16.566’N
002°18342’
7–8 Jun 2005
K.B.Miller
, colr. (
BYU
, slide Z18.76)
.
Republic of Guinea
:
1♁,
Kousankoro
no.2, 21.10.84 (
SMNS
, gift
J.M.Élouard
; slide 16/95)
.
4♁,
Guinea
, piège lumineuse, sans étiquette (slides Z18.66 & 67, gift
J. M. Élouard
)
.
Republic of Liberia
:
2♁,
2♀
,
Suakoko
[
7.11°N
,
9.59°W
349m
] 31
Jan
52
Blickenstaff
(slides
USNM 8
,
11
,
12
)
;
1♁, same, except
3–9 May 1952
(slide
USNM 41
)
.
Republic of Sierra Leone
:
1♁,
Njala Sierra Leone Afr.
XI-16–XII-4 35 \
Van Zwaluvenburg
&
McGough
(
USNM 13
)
.
Togolese Republic
:
1♀
,
Amoutchou
,
28.11.1985
, V.
Landa
(
SMNS
,
Z16.132
)
.
Habitus.
WL of males 8.2–11.0mm, of females 11.7–13.0mm. Uniformly yellowish ochre, the brown ocellar mark is small. Tip of flagellum faintly infuscate, cercus yellow.
Male
(
Figs. 294–300
). Sternites unmodified. A triangular sclerite on T7 has a narrow caudal process with single SB at tip and a smooth keel along rear face (
Fig. 294
); some SB in the intersegmental fold where the sclerite is articulated with T8. Sclerite on T8 T-shaped, with a distinct hump with numerous SB in front (
Fig. 295
). T9 with a median depression between pilose lateral swellings with many SB. HT10 short, regularly curved in dorsal and lateral views, sharply pointed, mediobasal callus tongue-shaped.
Penis (
Figs. 296–299
) a straight slender tube, distal third membranous except for long lateral sclerites between paired dorsal spine patches and a pair of more distal ventral spine patches (
Fig. 300
). Tips of spines point in opposite directions, spines in the ventrodistal groups point towards the penis base. Base of endophallus dorsomedially (
Fig. 296
: eb) with patch of erect slender sharp spines. There are scattered slender spines on the wide endophallus base. Distally, the endophallus is a narrow tube with three spine rows: two regular rows of conical spines dorsally plus a single regular row of more slender and locally also longer spines ventrally (
Fig. 296
, inset). Differences between rows diminish distally, all continue to the tip of the endophallus.
Female
. As for the group (
Fig. 301
).
Egg
(
Figs. 302–304
). Size 345–380µm long, on average 1.85 times longer than wide. Numerous (approximately 30) straight striae, each consisting of a wide rib associated with a slightly narrower rib (
Fig. 303
). The ribs are separated by a single row of micropunctures. All ribs are impunctate, micropyles are located on top of the narrow ribs (similar to
Fig. 312
, after
Moore 1991
). No collar, the wide ribs end in oval depressions on a thickened ring around the anchor pole, the narrow ribs end between them, slightly more distant from the edge (
Fig. 304
). Anchor cavity shallow. Operculum bluntly rounded, with faint reticulation, low impunctate strips separate finely punctate polygones.
DNA
(
Figs. 491–492
,
496
). A total of
10 specimens
from
Togo
and
Ghana
were sequenced for the COX1 DNA barcode fragment and with the genome-skimming approach, providing limited representation of the geographical distribution of the species. This also includes the male
holotype
from
Ghana
for the COX1 DNA barcode fragment. The monophyly of the species is very strongly supported (99.7/99/9), but its placement as sister to
N. filamentosa
n. sp.
is only poorly supported (43.9/90/0). The poor support for this placement is linked to the inclusion of
NEOP
167, which includes particularly badly fragmented mitochondrial genome assembly data and has the potential for the inadvertent inclusion of cross contaminant fragments, potentially resulting in instability of the placement of this specimen relative to
N. filamentosa
n. sp.
.. Together,
N. orthonema
n. sp.
and
N. filamentosa
n. sp.
are maximally supported (100/100/100) as sister to
N. spironema
n. sp.
. Both sexes are well represented.
FIGURES 294–304.
Neoperla orthonema
n. sp.
: Male paratype from Lakota, Ivory Coast: 294, 295, abdominal tip; 296, everted penis, with detail from middle of endophallus; 297–299, apex of contracted penis, in dorsal, ventral and lateral views (from left); 300, lateral view of penis apex, beginning eversion (♁ from Kousankoro); 301, female genital situs (vagina flipped back to show the anchor pattern on S8) and 302, egg, Ankasa game prod. res., Ghana; 303, view onto edge of chorion fragment showing variable size of costae (Ivory Coast, Lakota); 304, detail of collar with end of striae (Wli Falls, Ghana). Scales are 100µm. Tergites, sternites and hemitergite 10 are labeled. d, dorsal spines; eb, spines at base of endophallus; sa, spermatheca; SSt, spermathecal stalk; V, vagina; vs, ventral spines.
Notes
. In all species of
Neoperla
, micropyles are at the bottom of sulci but in both
N. orthonema
n. sp.
and
N. spironema
n. sp.
they are on top of narrow low ribs (
Fig. 312
), which apparently are secondarily uplifted sulci.
Etymology
. The name is a composite noun in apposition, formed from the Greek stems
ortho
, straight, and
nema
, thread, alluding to the straight egg striae and the threads supporting the cap of the anchor.