The American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae): a new generic concept based on a reinforced cathrema in the phallus

Abstract

We describe one new genus (Brachinepticula Stonis & Diškus, gen. nov.) with two new species (B. plurilobata Diškus & Stonis, sp. nov., B. elongata Remeikis & Stonis, sp. nov.), and one species with uncertain taxonomic position (Johanssoniella bina Remeikis & Stonis, sp. nov.). We also provide diagnostic characters and an updated annotated catalogue of the following related genera: newly restored Johanssoniella Koçak, Brachinepticula gen. nov., Enteucha Meyrick (s. str.) and Manoneura Davis. In the Annotated Catalogue, we provide five new combinations and new data on morphology, biology, and distribution of some species, including the first photographic documentation of Manoneura basidactyla Davis discovered in the Amazon rainforest, results of re-examination of the male genitalia of the European Johanssonia acetosae (Stt.), also the first documentation of the male genitalia, host plant and leaf mines of the little known Johanssoniella diplocosma (Meyrick) from the Himalayas. All new taxa treated in the paper are illustrated with drawings and (or) photographs of the adults and genitalia; photographs of the leaf mines of Brachinepticula plurilobata Diškus & Stonis, sp. nov. and Johanssoniella diplocosma (Meyrick) are also provided.

Full text

BIOLOGIJA. 2018. Vol. 64. No. 2. P. 99–128
© Lietuvos mokslų akademija, 2018
e American Brachinepticula gen.nov.
and Manoneura Davis (Nepticulidae):
anew generic concept based on areinforced
cathrema in thephallus
* Corresponding author. Email: stonis@leu.lt
JonasR.Stonis1*,
ArūnasDiškus1,
AndriusRemeikis1,2,
MariaAlmaSolis3
1Lithuanian University
of Educational Sciences
and Baltic-American
Biotaxonomy Institute,
StudentųSt.39,
Vilnius 08106, Lithuania
2Institute of Ecology,
Nature Research Centre,
AkademijosSt.2,
Vilnius 08412, Lithuania
3Systematic Entomology Laboratory,
ARS, USDA, National Museum
of Natural History,
Smithsonian Institution,
Washington, D.C.,
20013-7012, USA
We describe one new genus (Brachinepticula Stonis & Diškus,
gen.nov.) with two new species (B.plurilobata Diškus&Stonis,
sp.nov., B. elongata Remeikis & Stonis, sp.nov.), and one spe-
cies with uncertain taxonomic position (Johanssoniella bina
Remeikis & Stonis, sp.nov.). We also provide diagnostic char-
acters and an updated annotated catalogue of the following re-
lated genera: newly restored Johanssoniella Koçak, Brachinept-
icula gen.nov., Enteucha Meyrick (s.str.) and Manoneura Davis.
In theAnnotated Catalogue, we provide ve new combinations
and new data on morphology, biology, and distribution of some
species, including the rst photographic documentation of
Manoneura basidactyla Davis discovered in theAmazon rainfor-
est, results of re-examination of themale genitalia of theEuro-
pean Johanssonia acetosae (Stt.), also therst documentation of
themale genitalia, host plant and leaf mines of thelittle known
Johanssoniella diplocosma (Meyrick) from theHimalayas. All new
taxa treated in thepaper are illustrated with drawings and (or)
photographs of theadults and genitalia; photographs of theleaf
mines of Brachinepticula plurilobata Diškus&Stonis, sp.nov. and
Johanssoniella diplocosma (Meyrick) are also provided.
Keywords: American fauna, Brachinepticula Stonis & Diškus,
Enteucha Meyrick, Johanssoniella Koçak, leaf mines, Manoneura
Davis, Nepticulidae, new genus, new species
INTRODUCTION
Among leaf-mining pygmy moths (Lepidoptera,
Nepticulidae), there are over a dozen described
(and many undescribed) nepticulid species which
we informally name the“Enteucha-like assemblage”.
ese species are characterized by thelamellar col-
lar (homologous to thecollar of Stigmella Schrank,
Simplimorpha Scoble, and afew other genera), ab-
sence of transtilla in themale genitalia (paralleled in
Glaucolepis Braun and some other taxa), thedistinc-
tive band-like tegumen, extremely weak or totally
undeveloped cornuti, and reduced forewing vena-
tion lacking either vein CuA or Rs4. Some species of
this “assemblage” occur in North America, Europe,
and Asia, however, the majority occur in Central
and South America, and namely they include some

100 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
of themost striking representatives in the Neo-
tropical fauna with remarkable forewing colora-
tion and with distictively modied structures of
thegenitalia. Host-plant preferences are still un-
known for nearly two-thirds of the species, but
species with known biology feed on Polygonace-
ae plants. Phylogenetically this “assemblage” has
been found to be a sister group to Stigmella
Schrank (Puplesis, 1994) or to all remaining Nep-
ticulidae (Doorenweerdetal., 2016).
In recent literature, species from the“Enteu-
cha-like assemblage” were either divided between
two genera (Enteucha Meyrick and Manoneura
Davis; see Davis, 1978, 1979; Puplesis&Robin-
son, 2000; Puplesisetal. 2002b; Diškus&Puple-
sis, 2003; Stonisetal., 2017) or treated as belong-
ing to one genus (Enteucha Meyrick, sensu lato;
see van Nieukerken, 1986a; 1986b, van Nieuker-
kenetal., 2016a).
Recent molecular research (Regieretal., 2013;
Doorenweerdetal., 2016) indicated aheteroge-
neity of this artical assemblage. Despite avery
few shared characters, including the variously
reduced wing venation, morphology of these
species shows a great diversity (Figs. 1–4). In
thecourse of our recent study of the“Enteucha-
like assemblage”, we discovered novel morpho-
logical characters, notably thepresence of vaginal
sclerites in thefemale genitalia, and agreatly de-
veloped pseudoanellus, and astrongly reinforced
cathrema of the phallus in the male genitalia.
eterm “cathrema” was proposed by Schoo-
rletal. (1985) and is characteristic of all Neptic-
ulidae (i.e., an apomorphy of thefamily). It rep-
resents astriate or smooth thickening at thebase
of theductus ejaculatorius and is variously devel-
oped in dierent genera of Nepticulidae. Among
the species of the “Enteucha-like assemblage”,
we can observe cathrema rather weakly devel-
oped (Fig. 3) in Enteucha or extremely weakly
developed, indistinctive or maybe even absent
(Fig.1) in Johanssoniella. However, there are two
putative monophyletic clusters of species, each
of which exhibit strongly reinforced cathrema:
either with unique, very long, lateral processes
in Brachinepticula gen.nov. described here (see
Fig. 2) or in the genus Manoneura, cathrema
strongly extended caudally (Fig.4). It is interest-
ing to note that these latter unusual evolution-
ary trends are paralleled with another morpho-
logical feature of Brachinepticula gen.nov. and
Manoneura–theabsence of cornuti (Figs.2, 4)
so characteristic of thevast majority of Nepticu-
lidae outside of the“Enteucha-like assemblage”.
For researchers studying theAmerican Nep-
ticuloidea, it does not take very long to perceive
that Manoneura Davis represents atruly exotic
genus, themost distinctive and probably most
beautiful and endemic element of theNeotropi-
cal Nepticulidae fauna. Externally, the species
of Manoneura are very distinctive because of
thestrong purple or ochreous golden lustre (see
Figs.93–98) and internally thehighly modied
uncus and gnathos of Manoneura form aunique,
movable, lock-like formation (Figs.80, 83, 89).
Moreover, the phallus has distinctive carinae,
thesublateral process of thevalva is unusually
enlarged, the vinculum has a specic lateral
thickening (apodeme), and thevalva has ador-
sal process (Fig. 4). Additionally, the strongly
developed cathrema makes the genus even
more derived. erecently described Peruvian
Manoneura forcipis Remeikis & Stonis (Stonis
etal., 2017) probably represents the most out-
standing example: thecathrema is protruding
from thephallus (Fig. 81). Similarly, although
not so strongly reinforced, a cathrema is ob-
served in other species of Manoneura (Figs.79,
84) and species of Brachinepticula gen. nov.
(Figs.2, 15, 22). To accommodate themarkedly
apomorphic Manoneura (Fig.4) and newly rec-
ognized Brachinepticula genus (Fig.77) within
the classication of Nepticulidae and the “En-
teucha-like assemblage” in particular, we needed
anew taxonomic concept of four genera (or ve
including themonotypic African Va r iu s Scoble).
Detailed characterizations of all these genera are
provided in Fig.1 and additionally discussed af-
ter each genus in theAnnotated Catalogue be-
low. Now it appears that theearlier synonymi-
zation of Johanssoniella Koҫak (=Johanssonia
Borkowski) with Enteucha by van Nieukerken
(1986a) and followed by Puplesis (1994) and
Diškus&Puplesis (2003) was premature. ege-
neric status of Johanssoniella (Fig.1) is restored,
although it can be characterized only by avery

101
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
few apomorphies (we expect more will be found
in the future) (also see Discussion in the An-
notated Catalogue). We also describe one new
genus, Brachinepticula Stonis & Diškus, with
two new species (B.plurilobata Diškus&Stonis,
sp.nov., B. elongata Remeikis&Stonis, sp.nov.),
and one species with an uncertain taxonomic
position (Johanssoniella bina Remeikis & Sto-
nis, sp.nov.). We provide an updated Annotated
Catalogue for therelated genera of the“Enteu-
cha-like assemblage”. We also provide the rst
photographic documentation of Manonera
basidactyla discovered in theAmazon rainfor-
est, theresults of re-examination of male genita-
lia of theEuropean Johanssoniella acetosae (Stt.),
and therst documentation of male genitalia,
host plant, and leaf mines of the little known
Johanssoniella diplocosma (Meyrick) from
theHimalayas.
MATERIALS AND METHODS
Descriptions of new taxa are based on material
deposited in the collection of the Zoological
Museum, Natural History Museum of Den-
mark, Copenhagen, Denmark (ZMUC), col-
lected in Argentina (by E.S.Nielsen and Ole
Karsholt, Mision Cientica Danesa) and Ecua-
dor (by Arūnas Diškus and J.R.Stonis). Other
material in the current paper was mostly re-
ceived from theUSNM, and also from other in-
stitutions including theBMNH, ZIN, and LEU.
Methods and protocols for species identi-
cation and description are outlined in Puplesis
(1994), Puplesis & Robinson (2000), Puple-
sis&Diškus (2003), and Stonisetal. (2016).
Preparation of temporary and permanent mi-
cro-mounts of genital structures was undertaken
following themethod by Stonisetal. (2014). Af-
ter maceration of theabdomen in 10% KOH and
subsequent cleaning, abdominal pelts and fe-
male genitalia were stained with Chlorazol Black
(Direct Black 38/Azo Black), and male genitalia
were le unstained. Male genital capsules were
removed from theabdomen and mounted ven-
tral side uppermost. e phallus was severed
from thegenital capsule. Both male and female
genitalia were mounted in Euparal.
Permanent slides were photographed and
studied using aLeica DM2500 microscope and
Leica DFC420 digital camera.
e descriptive terminology of morphologi-
cal structures follows Johanssonetal. (1990) and
Puplesis&Robinson (2000), except for theterm
“aedeagus”, which is referred here as “phallus” and
theterm “cilia”, which is referred here as “fringe”.
Institutional abbreviations used in thetext:
BMNH – e Natural History Museum,
London, United Kingdom;
LEU–Lithuanian University of Educational
Sciences (formerly VPU), Vilnius, Lithuania;
USNM – United States National Museum
of Natural History, Smithsonian Institution,
Washington, D.C., USA;
ZIN – Zoological Institute of the Russian
Academy of Sciences, St.Petersburg, Russia;
ZMUC–Zoological Museum, University of
Copenhagen, Denmark.
DESCRIPTION OF BRACHINEPTICULA
STONIS&DIŠKUS, GEN.N O V.
Type species. Brachinepticula plurilobata Diš-
kus & Stonis, sp.nov. by present designation.
Diagnosis (Figs.1–4). Brachinepticula is rec-
ognised by thepresence of well-developed pseu-
doanellus and distinctive, paired juxta, long lat-
eral processes of cathrema, also lobate valva in
the male genitalia, and the vaginal sclerites in
thefemale genitalia. Wing venation with asepa-
rate vein CuA. As far as we know, blotch-like leaf
mines also make this genus distinctive.
Description. Adult (Figs.5–7). Rather small
nepticulid moths, forewing length 1.8–2.5mm;
wingspan 5.1–5.5mm. Head (Figs.8, 9): palpi
greyish cream to grey brown; maxillary palpus
four-segmented; labial palpus three-segment-
ed; frontal tu beige cream to orangish yellow;
collar distinctive, comprised of lamellar scales;
scape large; male antenna longer than half of
thelength of forewing; agellum (Fig. 10) with
36 segments in female, 43–45 in male. Fore-
wing dark, with or without purple iridenscence,
uniformly coloured or with two fasciae (Fig.5);
fringe grey-brown, without fringe line; under-
side of forewing without spots or androconia.

102 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs.1–4. Diagnostics of Johanssoniella Koҫak, Brachinepticula gen.nov., Enteucha Meyrick (s. str.), and
Manoneura Davis
*–aer van Nieukerken, 1986, modied; **–aer Davis, 1978, modied.

103
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.5–11. Morphology of Brachinepticula gen.nov. 5–Brachinepticula elongata Remeikis&Stonis, sp.nov.,
male adult, dorsal view; 6–same, ventral view; 7–B.plurilobata Diškus&Stonis, sp.nov., male adult, re-
constructed, enhanced; 8–same, descaled head, slide no. AD686; 9–same, slide no. AD687; 10–same,
descaled agellum, slide no. AD686; 11–same, forewing venation, slide no. AD689 (ZMUC)

104 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Forewing venation (Fig.11) with aseparate CuA.
Hindwing lanceolate, without androconia. Male
abdomen with two distinctive anal tus.
Female. Similar to male but tends to be paler.
Flagellum of antenna with about 36 segments.
Male genitalia (Figs. 2, 12–30). Capsule
longer (285–380μm) than wide (140–280μm).
Tegumen band-like (Fig. 23). Uncus wide,
rounded (Fig.24), without caudal protrusion.
Gnathos with one stout caudal process, slender
lateral arms, and with or without central plate
(Figs. 14, 25). Valva lobate, with a basal arc;
transtilla absent; pseudoanellus well developed,
arising from basal processes of valva (Fig.26).
Juxta paired, distally pointed and (or) elaborat-
ed, arising from basal arc (Fig.26). Vinculum
without lateral lobes, widely rounded or trian-
gular. Phallus angular (Figs.28–30) or almost
rounded basally (Figs. 15, 16), 260–310 μm
long, without cornuti on vesica; cathrema with
very long, lateral processes (Figs.2, 15, 28–30).
Female genitalia (Fig. 32). Relatively short,
total length about 490μm. Anterior apophyses
lobe-like, slender only distally, but without dis-
tinctive distal processes; posterior apophyses
slender, rod-like. Genitalia with two large, heavily
chitinized vaginal sclerites (Figs. 31, 33). Cor-
pus bursae partially reduced, weakly sclerotized,
without signum or pectinations. Accessory sac
heavily folded; ductus spermathecae without dis-
tinctive coils. Abdominal tip wide, rounded.
Bionomics (Figs.36–47). Larvae of thetype
species, Brachinepticula plurilobatasp. nov.,
mine in leaves in January. Host plant: we were
unable to determine the genus and species of
theplant, but it belongs to thePolygonaceae fam-
ily. Leaf mine starts as ashort but very slender
gallery; later on it develops to ablotch with frass
accumulated in thecentre of theblotch (Fig.45).
Larva spins its cocoon outside themine. Cocoon
of thetype species oval-shaped, very pale, almost
cream. Pupa illustrated in Figs.41–43.
Distribution (Fig.48). especies of thenew
genus occur on thewestern slopes of theequato-
rial (northern) Andes and on theeastern slopes
of central Andes at altitudes up to 2700m.
Etymology. egenus name is derived from
the Latin brachiata (branched) and Nepticula
(synonymous name for thegenus Stigmella) in
reference to thelobate valva, paired juxta, and
divided pseudoanellus in themale genitalia.
Discussion. From Johanssoniella Koçak
(Fig.1), the new genus Brachinepticula gen. nov.
diers in the unique cathrema with long lat-
eral processes (cathrema indistinctive or stick-
shaped in Johanssoniella), presence of astrongly
developed pseudoanellus, widely rounded un-
cus (usually triangular in Johanssoniella), com-
plex, lobate valva (usually simple, triangular in
Johanssoniella), and thepresence of vaginal scle-
rites in thefemale genitalia.
From Enteucha Meyrick (Fig.3), thenew ge-
nus diers in thecathrema with long lateral pro-
cesses, thepresence of distinct, widely rounded
uncus (reduced or partially reduced in Enteu-
cha), gnathos with astout caudal process (with-
out astout caudal process, usually transverse in
Enteucha), the presence of pseudoanellus and
juxta (absent in Enteucha), less reduced forewing
venation (see Figs.2 and 3), and thepresence of
vaginal sclerites in thefemale genitalia.
From Manoneura Davis (Fig. 4), the new
genus diers in thepresence of pseudoanellus
and juxta, widely rounded uncus, gnathos with
a stout caudal process (in Manoneura, unique
uncus, and gnathos form of alock-shaped struc-
ture), short basal process of valva (very long in
Manoneura), absence of athickened apodeme of
vinculum (present in Manoneura), phallus with-
out carinae (with distinct carinae in Manoneu-
ra), cathrema with long processes (half-tubular,
without lateral process in Manoneura), thepres-
ence of vaginal sclerites in thefemale genitalia,
and the less reduced forewing venation, with
aseparate CuA (see Figs.2 and 4).
Brachinepticula elongata Remeikis&Stonis,
sp.nov.
(Figs.5, 6, 12–17, 48)
Type material. Holotype: ♂, ARGENTINA,
Salta, Rosario de la Frontera, Los Banos,
11.iv.1979, Mision Cientica Danesa (E.S.Niels-
enetal.), genitalia slide no. RA625♂ (ZMUC).
Diagnosis. ecombination of theforewing
with afascia, short juxta, valva with pointed api-
cal process and slender inner lobe, triangular

105
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.12–17. Male genitalia of Brachinepticula elongata Remeikis&Stonis, sp.nov., holo-
type, genitalia slide no. RA625 (ZMUC). 12, 13–capsule with phallus removed, 14–same,
enhanced; 15–17–phallus
vinculum, and unique processes of cathrema
in thephallus distinguishes B.elongata sp. nov.
from all Nepticulidae species, including there-
lated B.plurilobata Diškus&Stonis, sp.nov.
Male (Figs. 5, 6). Forewing length about
1.8mm; wingspan about 5.1mm. Head: palpi
grey-brown; frontal tu very pale, orangish
yellow; collar distinctive, comprised of lamel-
lar scales, grey cream; scape cream; agellum
of antenna brown on upper side, pale brown
on underside; number of segments unknown.
orax and tegula dark grey-brown. Forewing

106 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
dark grey-brown, with two silvery shiny fas-
ciae: ill-dened antemedian and wide postme-
dian; fringe grey-brown; underside of forewing
brown, without spots or androconia. Hindwing
and its fringe very pale brown on upper side
and underside. Legs brown on upper side, pale
brown to brownish cream on underside.
Female. Unknown.
Male genitalia (Figs.12–17). Capsule sig-
nicantly longer (285–290 μm) than wide
(140–150μm). Uncus wide, rounded. Gnathos
with one wide caudal process; central plate
absent; lateral arms long and slender. Valva
125–130 μm long, 50 μm wide, with alarge,
apical process and very slender, inner lobe;
transtilla absent; pseudoanellus present, in-
distinctive. Juxta short, but distinctly paired,
distally pointed. Vinculum very long, triangu-
lar, without lateral lobes. Phallus (Figs.15–17)
310–315 μm long, 50–65 μm wide, without
cornuti on vesica; cathrema with two very long
lateral processes.
Bionomics. Adults y in April. Otherwise
biology unknown.
Distribution (Fig. 48). is species occurs
in Argentina on theeastern slopes of the An-
des, in theJungas biogeographical province, at
an elevation of about 750m.
Etymology. especies name is derived
from the Latin elongatus (meaning distant or
remote from) in reference to theunique, slen-
der male genitalia with greatly extended pro-
cesses of thecathrema.
Brachinepticula plurilobata Diškus&Stonis,
sp.nov.
(Figs.7–11, 18–48)
Type material. Holotype:♂, ECUADOR:
Pichincha Province, 24km NW Alóag, 0°27’07”S,
78°41’00”W, elevation 2720m, mining larvae on
Muehlenbeckiasp., 14.i.2005, eld card no.4814,
leg. A.Diškus, J.R.Stonis, genitalia slide no.
AD680♂ (ZMUC). Paratypes: 4♂, 3♀, only
from mature pupae, label data as holotype, slides
no. AD681♂, AD682♂, AD897♂, AD902♂,
AD683♀, AD684♀, AD686♀ (head), AD687♀
(head), AD689♀ (wing venation), AD901♀
(ZMUC).
Diagnosis. ecombination of theuniform
forewing without fascia, long juxta, strongly
developed pseudoanellus, three-lobed valva,
widely rounded vinculum, and unique pro-
cesses of the cathrema in the phallus distin-
guishes B.plurilobatasp.nov. from all Nepticu-
lidae species, including therelated B.elongata
Remeikis&Sto nis, sp.nov.
Male (Figs. 7–11). Forewing length about
2.5mm; wingspan about 5.5mm. Head: max-
illary palpus greyish cream; frontal tu large,
beige cream; collar distinctive, comprised of la-
mellar scales, grey-brown, glossy; scape cream,
glossy; antenna signicantly longer than half
of thelength of forewing; agellum with 43–45
segments, brown with little green iridescence.
orax, tegula and forewing densely speckled
with dark grey-brown scales with distinctive
purple and blue iridescence. Forewing without
fascia; fringe grey-brown; underside of fore-
wing grey-brown, without spots or androconia.
Forewing venation (Fig. 11) with a separate
vein CuA. Hindwing pale grey-brown with lit-
tle purple iridescence on upper side; its fringe
pale grey-brown with some golden gloss. Legs
brownish cream to brown, with golden gloss;
foreleg darkened with fuscous brown scales on
upper side. Abdomen dark fuscous brown on
upper side and underside; genital plates brown-
ish grey, anal tus distinctive, brown.
Female. Similar to male but tends to be paler.
Flagellum of antenna with about 36 segments.
Male genitalia (Figs.18–30). Capsule longer
(375–380μm) than wide (270–280μm). Uncus
wide, rounded. Gnathos with one stout caudal
process, central plate, and slender arms. Valva
235–240μm long, 75–80μm wide, lobate; tran-
stilla absent; pseudoanellus strongly developed,
arising from basal process of valva. Juxta long,
paired, distally eleborated. Vinculum without
lateral lobes, widely rounded. Phallus with chi-
tinized lateral lobes (Figs. 22, 28–30), angular,
240–260μm long, 80μm wide basally, without
cornuti on vesica; cathrema with very large lat-
eral processes.
Female genitalia (Figs.31–35). Total length
about 490 μm. Anterior apophyses lobe-like,
slender only distally, shorter than posterior

107
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.18–22. Male genitalia of Brachinepticula plurilobata Diškus&Stonis, sp.nov., holotype,
genitalia slide no. AD680 (ZMUC). 18, 19– capsule with phallus removed, 20– same, en-
hanced; 21–phallus; 22–same, enhanced
apophyses. Genitalia with two large vaginal
sclerites (Figs.31, 33, 35). Corpus bursae partial-
ly reduced, weakly sclerotized; signum or pec-
tinations absent. Accessory sac rounded, very
heavily folded; ductus spermathecae chitinized,
sinuous but without distinctive coils. Abdominal
tip very wide, rounded.
Bionomics (Figs. 36–47). Larvae mine in
leaves in January. Host plant: Muehlenbeckia
Meisn., M.tamnifolia (Kunth) Meisn. or most

108 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs.23–30. Male genitalia of Brachinepticula plurilobata Diškus&Stonis, sp.nov. 23–tegumen; 24–uncus;
25–gnathos; 26, 27–pseudoanellus and juxta; 28–30–phallus (ZMUC)

109
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.31–35. Female genitalia of Brachinepticula plurilobata Diškus&Stonis, sp.nov. 31–vaginal sclerites,
slide no. AD901; 32– general view, slide no. AD683; 33– apophyses and vaginal sclerites, slide AD684;
34–ductus spermathecae, slide AD683; 35–vaginal sclerite, slide no. AD684 (ZMUC)

110 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs.36–47. Bionomics of Brachinepticula plurilobata Diškus&Stonis, sp.nov. 36, 38–leaf mines, Ecuador,
24 km NW Alóag, 0˚27’07”S, 78˚41’00”W, 2720 m, eld card no. 4814; 37–host plant, Muehlenbeckia sp.;
39, 40–cocoons; 41–43–pupae; 44, 45–adults; 46, 47–leaf mines, eld card no. 4814

111
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
likely M.tiliifolia Wedd. However, the identity
of thehost plant should be treated with caution
because it is based on photographed plant sam-
ples. Leaf mine starts as ashort, but very slender
gallery; later on it develops to ablotch with black
frass accumulated in the centre of the blotch
(Fig.46). Exit slit on upper side of theleaf. Co-
coon outside themine (in forest litter), brownish
cream to cream; length 2.8–3.3mm, maximal
width 1.5–1.8mm. Pupa illustrated in Figs.41–
43. Adults (Figs.44–45) y in late January–early
Februar y.
Distribution (Fig. 48). is species occurs
on thewestern slopes of theequatorial Andes
of Ecuador (Pichincha Province) at an altitude
of about 2700m.
Etymology. especies name is derived from
the Latin plures (meaning most, many, or sev-
eral) and lobatus (meaning lobed) in reference
to thelobate valva, juxta, and pseudoanellus of
themale genitalia.
ANNOTATED CATALOGUE OF RELATED
GENERA
Recent on-going eldwork in South America, an
area historically poorly surveyed for leaf-mining
nepticulids, has provided additional material for
morphological study, and has broadened our
knowledge about distributions and biological
information. Based on these new observations,
we realized that a new taxonomic concept for
thegenera within the“Enteucha assemblage” was
needed. Also, recent molecular results showed
that its internal classication is still in ux. Anew
taxon from theAndes in South America required
that we re-visit and re-evaluate morphological
characters and their distribution within this as-
semblage. We make note that thereinforced cath-
rema in thephallus, shared by Manoneura and
the new genus, Brachinepticula, occurs in tan-
dem with avesica without cornuti, acondition
not found in other Nepticulidae. Its signicance
Figs.48–50. Distribution of Brachinepticula elongata Remeikis&Stonis, sp.nov. and B.plurilobata Diškus&Sto-
nis, sp.nov. 48–map (courtesy of T. Patterson, USA); 49, 50–habitat of B.plurilobata, 24 km NW Alóag, Ecuador

112 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
remains to be discovered. We could not jus-
tify inclusion of this new taxon into theexisting
genera and therefore described the new genus,
Brachinepticula. We continue to expand our bio-
logical discoveries, specically on thebiology of
two Brachinepticula species, and provide an An-
notated Catalogue to summarize our ndings
in other closely related genera in the“Enteucha
assemblage”. We expect that future biological re-
search will provide more clarity to theknowledge
about, and relationhips among, these beautiful
and biologically interesting moths.
Genus Johanssoniella Koçak, 1981 (Fig.1)
(status restored)
Type species: Nepticula acetosae Stainton,
1854: 303.
Johanssonia Borkowski, 1972: 702–705.
Johanssoniella Koçak, 1981: 99 (replacement
name for Johanssonia Borkowski).
Discussion. etaxonomic status is re-
stored. Any other solution, i.e., placing it with-
in the highly apomorphic Enteucha Meyrick
or other genus, seems impossible. At aglance,
Johanssoniella may seem to lack apomorphies,
but we found the triangular uncus and dis-
tinct triangular valva with long apical process
supports thetaxon. We expect that more apo-
morphies for Johanssoniella may be found
in thefuture. ediagnostics of thegenus is
straightfoward.
From Brachinepticula gen. nov. (Fig. 2),
the genus diers in the indistinctive or stick-
shaped cathrema (with long lateral processes
in Brachinepticula), absent or very weakly de-
veloped juxta and pseudoanellus, triangular
uncus, and theabsence of vaginal sclerites in
thefemale genitalia.
From Enteucha Meyrick (Fig.3), thegenus
diers in thesimple, undivided valva, thepres-
ence of a distinct triangular uncus (fully or
partially reduced in Enteucha), gnathos with
a stout, caudal process, usually rounded or
slightly lobate vinculum (mostly triangular
in Enteucha), and the forewing venation with
apreserved Rs4 (reduced in Enteucha).
From Manoneura Davis (Fig.4), thegenus dif-
fers in thetriangular uncus, gnathos with astout
caudal process, simple, triangular valva, short ba-
sal process of valva, theabsence of thickened ap-
odeme of vinculum, indistinctive or stick-shaped
cathrema, and thephallus without carinae.
Johanssoniella acetosae (Stainton, 1854)
(Figs.51–58)
Nepticula acetosae Stainton, 1854: 303.
Nepticula acetosella Doubleday, 1859: 36.
Nepticula arifoliella Klimesch, 1940: 92.
Stigmella acetosae (Stainton), Beirne, 1945:
200.
Johanssonia acetosae (Stainton), Borkowski,
1972: 702; Scoble, 1983: 15, 16.
Johanssoniella acetosae (Stainton), Koçak,
1981: 99.
Stigmella arifoliella (Klimesch) Hering, 1957:
912.
Enteucha acetosae (Stainton), Emmet, 1976:
266; van Nieukerken, 1986a: 54; Johanssonetal.,
1990:138, 139; Puplesis, 1994: 57; Diškus&Pu-
plesis, 2003: 321; van Nieukerkenetal., 2016a:
103.
Host plants. Rumex acetosaL., R.acetosel-
laL., R.arifolius All (Polygonaceae).
Distribution. Europe (except southern and
northern regions): from Ireland, Sweden and
Estonia to Slovenia and Ukraine.
Discussion. Two distinctly dierent forms
of adults were reported (see Johansson et al.,
1990: 139): asmaller and paler form from West-
ern Europe (“true” J. acetosae), and alarger and
darker form, with distinct purple iridescence,
feeding on Rumex arifolius predominantly
from Germany and alpine localities (previously
described as aseparate species arifoliella). Re-
cently we discovered both forms occurring side
by side in Lithuania, but thelarger and darker
form in thecountry prevails. ere is still aslim
chance that theforms may represent two dier-
ent species; more research is still needed.
In thecourse of our study (see Figs.51–58),
we made some discoveries about themorpho-
logical characters of Johanssoniella acetosae,
specically themale genitalia with variously de-
veloped juxta (Figs.1, 52, 57) and pseudoanel-
lus (Figs.55, 57, 58). elatter was previously
unknown; it is variously expressed from speci-
men to specimen even from the same reared
sample, but visible only in stained slides with

113
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.51–58. Re-examination of Johanssoniella acetosae (Stt), thetype species of thegenus Johanssoniella
Koҫak., the “larger and darker form” (see Discussion on thespecies), collected in Trakai, Lithuania,
54°38’20’’N, 24°57’29’’E, 150 m, on Rumex acetosa L., 18.vi.2000, eld card no. 4635. 51, 52–capsule
with phallus removed, slide no. AD903; 53–same, phallus; 54–phallus, slide no. AD907; 55–pseu-
doanellus, slide no. AD904; 56–valvae, slide no. 907; 57, 58–pseudoanellus, slide no. AD907 (LEU)

114 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
slightly ventrally turned genitalia. e pseu-
doanellus and varied juxta was conrmed for
both forms of J.acetosae collected in Lithuania.
Johanssoniella sp. (undescribed species from
Japan)
Eneucha spec., van Nieukerken, 1986a: 53, 54.
Discussion. emale genitalia of an unde-
scribed species from Japan were illustrated by
van Nieukerken, 1986a: Figs.99–101. is new,
but not yet described species, exhibits astrik-
ing similarity of themale genitalia to those of
theprobably very closely related J. acetosae.
Johanssoniella hilli (Puplesis & Robinson,
2000) comb. nov.
Enteucha hilli Puplesis & Robinson, 2000:
19, 20.
Enteucha hilli Puplesis &Robinson, in Pu-
plesisetal., 2002b: 64; Diškus&Puplesis, 2003:
321; van Nieukerkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Central America: Belize (rain-
forest).
Discussion. Illustrated in Puplesis & Rob-
inson, 2000: Figs. 8, 69, 70. is remarkable,
relatively large (7.4mm in wingspan) species
is characterized by the unicolorous forewing
coarsely speckled with brown scales, distinc-
tive, practically unpaired collar comprised of
wide lamellar scales, unusually large apical
process of valva, narrowed ventral plate of vin-
culum, and thechitinized cathrema in themale
genitalia; conuti in thephallus absent.
Johanssoniella contracolorea (Puplesis &
Robinson, 2000) com. nov.
Enteucha contracolorea Puplesis & Robin-
son, 2000: 20.
Enteucha hilli Puplesis &Robinson, in Pu-
plesisetal., 2002b: 64; Diškus&Puplesis, 2003:
321; van Nieukerkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Central America: Belize (rain-
forest).
Discussion. Illustrated in Puplesis&Robin-
son, 2000: Figs.9, 71–74. is distinct species
is characterized by thedark brown collar, large
and unusually elongated scape, smooth-scaled,
brown forewing with golden gloss and some
blue iridesncence, highly contrasting fringe,
and thepeculiar uncus in a shape of inverted
Y in themale genitalia; cathrema very simple,
cornuti absent.
Johanssoniella diplocosma (Meyrick, 1921)
(Figs.59–70) comb. nov.
Nepticula diplocosma Meyrick, 1921: 410, 411.
Enteucha diplocosma (Meyrick), in Diškus
& Puplesis, 2003: 321; van Nieukerken et al.,
2016a: 103.
Host plant. Persicaria amplexicaulis (D.Don)
Ronse Decraene (syn. Bistorta amplexicaulis
(D.Don) Greene), Polygoncaceae.
Distribution. India: Assam (type locality)
and Uttarakhand (the Himalayas, at an eleva-
tion of about 2200m) (Figs.64, 65).
Discussion. especies was described in
1921 on thebasis of a single male specimen
deposited at the BMNH. Externally it can
be recognized by thepresence of two golden
glossy fasciae on abrown (or purplish brown)
forewing. We compared themale genitalia of
theold specimen (theholotype, slide 25652)
with themale genitalia of two recently reared
male pupae from Uttarakhand, India (ZIN;
see Figs. 59–63); the Himalayan specimens
(mature pupae) appear to be conspecic
with Meyrick’s holotype; it allows us to name
thehost plant (see Figs.66–68) and document
the leaf mines of J.diplocosma (see Figs. 69,
70) for therst time. We also added new dis-
tribution data (Fig.65) and updated the in-
formation of mining (August) and y period
(June and August–early September). ead-
dition is based on thefollowing new material
examined: 2♂ (from mature pupae, no pinned
adults preserved), India, the Himalayas, Ut-
tarakhand, Tehri Garhwal District, Dhanaulti,
30°25’11’’N, 78°15’37’’E, elevation 2200 m,
mining larvae on Persicaria amplexicaulis (Po-
lygonaceae), 18.viii.2010, eld card no.5003,
leg.A.Diškus and A.Navickaitė, genitalia
slide nos. AD490♂, AD491♂ (ZIN) (Note:
thestudied type with thelabel “Shillong, As-
sam, TBF6.19”, genitalia slide no.25652 is de-
posited in theBMNH).
Johanssoniellasp. (undescribed species from
Japan)
Eneucha spec., van Nieukerken, 1986a: 53, 54.

115
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.59–63. Male genitalia of Johanssoniella diplocosma (Meyrick), theHimalaya, Uttarakhand, Dhanaulti,
30°25’11’’N, 78°15’37’’E, 2200 m. 59–capsule, freshly made slide AD490; 60–same, dry slide; 61–same,
phallus; 62–capsule, slide AD491; 63–capsule with phallus, slide AD491 (ZIN)

116 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs. 64–70. Distribution and bionomics of Johanssoniella diplocosma (Meyrick).
64–habitat, theHimalayas, Uttarakhand, Dhanaulti, 30°25’11’’N, 78°15’37’’E, 2200 m;
distribution map; 66–68–host plant, Persicaria amplexicaulis; 69, 70–leaf mines
Discussion. e male genitalia of an unde-
scribed species from Japan were illustrated by
van Nieukerken, 1986a: Figs.96–98. is new,
but not yet described species, exhibits strik-
ing similarity of themale genitalia to those of
theprobably very closely related diplocosma.
Johanssoniella acuta (Puplesis & Diškus,
2002) comb. nov.
Enteucha acuta Puplesis & Diškus, 2002a:
21, 22.
Enteucha acuta Puplesis&Diškus, in Puple-
siset al., 2002a: 21–22, Figs. 4, 26–28; 2002b:

117
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
64; Diškus& Puplesis, 2003: 321; van Nieuke-
rkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Ecuador (Amazon rainforest).
Discussion. Illustrated in Puplesis et al.
2002a: Figs. 4, 26–28. is distinctive spe-
cies from theAmazon Basin markedly diers
from other species in theslender, curved val-
va and large, plate-like pseudoanellus; phallus
probably with alarge horn-like cathrema, not
alarge cornutus as it was reported earlier (Pu-
plesisetal., 2002a); however, the morphology
of thephallus was not re-examined by us. Ex-
ternally, J.acuta is characterized by thesilvery
shiny fascia on thedark colour of theforewing.
Species with provisional (uncertain) place-
ment in Johanssoniella
Johanssoniella bina Remeikis & Stonis,
sp.nov. (Figs.71–76)
Type material. Holotype: ♂, Argentina,
Neuquen, Piedro del Aguila, 23.xii.1978, Mision
Cientica Danesa (leg.E.S.Nielsenetal.), geni-
talia slide no.RA523♂ (ZMUC). Paratype: 1♂,
Neuquen, Rio Limay, Arroyito, 17.xi.1978, Mi-
sion Cientica Danesa, (leg.E.S.Nielsenetal.),
genitalia slide no.RA522♂ (ZMUC).
Diagnosis. e combination of a sparsely
speckled forewing, inverted V-shaped gnathos,
trapezoid vinculum and alarge, and distinctive
cathrema in the male genitalia distinguishes
E.bina sp.nov. from all currently known nept-
iculid species, including all Johanssoniella (also
see Discussion).
Male (Figs. 71, 72). Forewing length 2.1–
2.7 mm; wingspan 4.6–5.9mm. Head: palpi
golden cream to brownish-cream, distally grey-
brown on upper side; frontal tu brown on
vertex, cream on frons; collar comprised of pili-
form scales, cream; scape cream; antenna half
thelength of forewing; agellum with about 45
segments, pale brownish-cream on upper side
and underside. orax, tegula and forewing
cream with some golden gloss, sparsely speck-
led with pale brown and brown scales; fringe
cream; underside of forewing brownish-yellow,
without spots or androconia. Hindwing cream
on upper side and underside, without spots or
androconia; fringe cream. Legs golden cream to
brownish-cream; foreleg darkened distally with
brown scales on upper side. Abdomen grey
cream on upper side, grey on underside; anal
tus short, cream; genital plates grey.
Female. Unknown.
Male genitalia (Figs.73–76). Capsule about
350 μm long. Uncus widely rounded, with
avery short caudal process (Fig.73). Gnathos
inverted V-shaped, with widely rounded caudal
element (Fig.73). Valva with alarge apical pro-
cess. Transtilla absent. Vinculum wide, without
lateral lobes. Phallus (Figs.74–76) 370μm long,
130–140μm wide, with alarge, binary cathre-
ma and numerous indistinctive, spine-like cor-
nuti (Fig.76).
Bionomics. Host plant unknown. Adults y
in November and December.
Distribution. is species occurs in thesouth-
ern Andes (Argentina: Neuquen).
Etymology. especies name is derived from
Latin binus (two at atime, binary) in reference to
thebinary cathrema in themale genitalia.
Discussion. etaxonomic position of
J. bina is uncertain and, therefore, provisional.
is new species is placed in Johanssoniella
because of the general similarity of the male
genital capsule to Johanssoniella. However, this
species possesses an odd, binary cathrema in
thephallus and, in contrast to other Johansso-
niella or related taxa, acollar comprised of pili-
form (not lamellar) scales. In thefuture, when
wing venation or molecular data becomes
available, thetaxonomic position of bina may
be rened.
Johanssoniella terricula (Puplesis&Robin-
son, 2000) comb. nov.
Enteucha terricula Puplesis & Robinson,
2000: 20, 21.
Enteucha terricula Puplesis& Robinson, in
Puplesis et al., 2002b: 64; Diškus& Puplesis,
2003: 321; van Nieukerkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Central Andes (Peru: Depart-
ment Puno).
Discussion. eplacement of J.terricula in
Johanssoniella is very uncertain and, therefore,
only provisional. ere are only two characters

118 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs. 71–76. Johanssoniella bina Remeikis & Stonis, sp. nov., Argentina, Neuquen.
71–fragment of male adult, paratype; 72–male adult, holotype; 73–capsule with phal-
lus removed, holotype, slide no. RA523; 74– phallus, holotype, slide no. RA523; 75,
76–same, paratype, slide RA522 (ZMUC)
of themale genitalia which J. terricula shares
with other Johanssoniella: thetriangular uncus
and indistinctive cathrema. is new species
is placed in Johanssoniella mostly because of
thelack of apomorphies characteristic of other
described genera.
Genus Brachinepticula Stonis & Diškus,
gen.nov. (described above) (Fig.2)
Type species Brachinepticula plurilobata
Diškus&Stonis, sp.nov.
Discussion. Along with the unusually re-
inforced cathrema or the vaginal sclerites in

119
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
the female genitalia, the presence of pseu-
doanellus or bid juxta seem to be very strong
apomorphies (Fig.2). However, our morpho-
logical studies of theEuropean Johanssoniella
acetosae unexpectedly revealed thepresence of
apseudoanellus and juxta, though very weakly
developed, in thegenus Johanssoniella as well.
It may indicate apossible relationship between
Brachinepticula with Johanssoniella. Neverthe-
less, Brachinepticula is a very distinctive and
highly apomorphic genus (see the legend of
Fig. 2). Diagnostics of the taxon is provided
above, in thedescription of Brachinepticula.
Brachinepticula elongata Remeikis & Sto-
nis, sp.nov. (described above) (Figs.5, 6, 12–17)
Host plant. Unknown.
Distribution. Argentina (the Jungas bioge-
ographical province) (Fig.48).
Brachinepticula plurilobata Diškus&Ston-
is, sp.nov. (described above) (Figs.7–11, 18–50)
Host plant. Belongs to the Polygonace-
ae family; probably Muehlenbeckia Meisn.:
M. tamnifolia (Kunth) Meisn., or, most likely,
M.tiliifolia Wedd., but theidentication needs
to be conrmed (Figs.36–38, 46, 47).
Distribution. Ecuador (Pichincha Province)
(Figs.48–50).
Genus Enteucha Meyrick, 1915 (Fig.3)
Enteucha Meyrick, 1915: 241.
Type species: Enteucha cyanochlora Mey-
rick, 1915: 241.
Artaversala Davis, 1978: 219, 221.
Type species: Artaversala gilvafascia Davis,
1978: 221–223.
Discussion. From Johanssoniella Koçak
(Fig.1), thegenus diers in thedivided valva,
reduced or partially reduced uncus (distinct,
triangular in Johanssoniella), usually transverse
gnathos without astout, caudal process, trian-
gular vinculum (rounded or slightly lobate in
Johanssoniella), and theforewing venation with
reduced Rs4 (preserved in Enteucha).
From Brachinepticula gen. nov. (Fig. 2),
thegenus diers in thecathrema without large
lateral processes, divided, two-folded valva,
reduced or partially reduced uncus (distinct,
rounded in Brachinepticula), usually trans-
verse gnathos without astout caudal process,
absence of pseudoanellus and juxta, reduced
forewing venation (see Figs. 2 and 3), and
the absence of vaginal sclerites in the female
genitalia.
From Manoneura Davis (Fig.4), thegenus
diers in thesimple cathrema (extended, half-
tubular in Manoneura), usually transverse gna-
thos, reduced or partially reduced uncus (un-
cus with gnathos form alock-shaped structure
in Manoneura), absence of thickened lateral
apodeme, triangular vinculum, phallus without
carinae (latter strongly developed in Manoneu-
ra), and thedierently reduced forewing vena-
tion (see Figs.3 and 4).
Enteucha cyanochlora Meyrick, 1915
Enteucha cyanochlora Meyrick, 1915: 241.
Enteucha cyanochlora Meyrick, in Davis, 1985:
142–145; van Nieukerken, 1986: 52, Figs.87–89;
Puplesis et al., 2002b: 64; Diškus & Puplesis,
2003: 321; van Nieukerkenetal., 2016: 103.
Host plant. Unknown.
Distribution. South America: Guyana (Bar-
tica).
Enteucha gilvafascia (Davis, 1978)
Artaversala gilvafascia Davis, 1978: 221–223.
Enteucha gilvafascia (Davis), in van Nieu-
kerken, 1986a: 52, Figs.90–92; Puplesisetal.,
2002b: 64; Diškus & Puplesis, 2003: 321; van
Nieukerkenetal., 2016a: 103.
Host plant. Coccoloba uvifera (L.) L., Po-
lygonaceae.
Distribution. USA: Florida (Davis, 1978;
1984) and Cuba (Núñez Aguila& Barro Caña-
mero, 2012).
Enteucha snaddoni Puplesis & Robinson,
2000
Enteucha snaddoni Puplesis & Robinson,
2000: 21, 22.
Enteucha snaddoni Puplesis&Robinson, in
Puplesis et al., 2002b: 64; Diškus& Puplesis,
2003: 321; van Nieukerkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Central America: Belize (rain-
forest).
Discussion. especies is illustrated in Pu-
plesis & Robinson, 2002: Figs.10, 75–79. is
species from the Belize rainforest diers from
other species by the partially reduced, almost

120 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
membranous, but lobe-like uncus and by
thecombination of alarge triangular vinculum,
very long and inwardly bent apical process of
thevalva, and adistinctly transverse gnathos.
Genus Manoneura Davis, 1979 (Figs. 4,
77–101)
Manoneura Davis, 1979: 276. Replacement
name for Oligoneura Davis, nec Bigot, 1878.
Type species: Oligoneura basidactyla Davis,
1978: 218, 219.
Oligoneura Davis, 1978: 217, 218.
Type species: Oligoneura basidactyla Davis,
1978: 218, 219.
Discussion. From Johanssoniella Koçak
(Fig. 1), the genus diers in the unique un-
cus and gnathos in the form of lock-shaped
Fig.77. Dierentiation of Manoneura Davis and Brachinepticula gen.nov.

121
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.78–84. Manoneura Davis, male genitalia. 78–M. basidactyla (Davis), slide no. Diškus002 USNM,
capsule with phallus, Dominica; 79, same, slide no. 29120 BMNH, phallus, Belize; 80–M. forcipis Re-
meikis&Stonis, slide no. RA552 ZMUC, uncus and gnathos, Peru; 81, same, cathrema and carinae;
82–M. trinaria Puplesis&Robinson, slide no. Diškus003 USNM, capsule with phallus removed, Ven-
ezuela; 83–same, uncus and gnathos; 84–same, phallus

122 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Figs. 85–89. Documentation of male genitalia of Manoneura basidactyla (Davis) from
theAmazon rainforest, Ecuador, Napo Region. 85–capsule with phallus removed, slide no.
AD327; 86–lateral apodemes of vinculum, slide no. AD905; 87–valva, lateral view, slide no.
AD905; 88–phallus, slide no. AD327 ; 89–uncus and gnathos, slide no. AD905 (ZMUC)
structure (uncus wide, triangular in Johanssoniel-
la, gnathos with a stout caudal process), ver y long
basal process of valva (short in Johanssoniella),
the presence of a thickened apodeme of vinculum
(absent in Johanssoniella), valva with a dor-
sal process (absent in Johanssoniella), phallus
with distinct carinae (absent in Johanssoniella),
greatly extended, half-tubular cathrema, and the
greatly reduced forewing venation (see Figs.1
and 4).

123
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Figs.90–92. Documentation of female genitalia of Manoneura basidactyla (Davis) from
the Amazon rainforest, Ecuador, Napo Region, genitalia slide no. AD326 (ZMUC).
90–general view; 91, 92–apophyses
From Brachinepticula gen. nov. (Fig. 2),
thegenus diers in theabsence of pseudoanel-
lus and juxta, theunique uncus and gnathos in
the form of amoveable lock-shaped structure
(uncus wide, rounded in Brachinepticula, gna-
thos with astout caudal process),very long ba-
sal process of valva (short in Brachinepticula),
thepresence of athickened apodeme of vincu-
lum (absent in Brachinepticula), phallus with
distinct carinae (absent in Brachinepticula),
half-tubular cathrema, and thegreatly reduced
forewing venation (see Figs.2 and 4).
From Enteucha Meyrick (Fig.3), thegenus dif-
fers in theunique half-tubular cathrema, unique

124 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
uncus and gnathos in the form of lock-shaped
structure (uncus reduced or partially reduced
in Enteucha), presence of athickened lateral ap-
odeme (absent in Enteucha), very wide vinculum
(triangular in Enteucha), phallus with distinct
carinae (absent in Enteucha), and thedierently
reduced forewing venation (see Figs.3 and 4).
Manoneura basidactyla (Davis, 1978)
(Figs.78, 79, 85–92, 97–101)
Oligoneura basidactyla Davis, 1978: 218, 219.
Manoneura basidactyla (Davis), in Puple-
sis& Robinson, 2000: 22–23: Figs. 12, 83–85,
207; Puplesisetal., 2002b: 61, 62, Figs.6, 9–14;
Diškus&Puplesis, 2003: 321.
Figs.93–98. Adults of Manoneura Davis. 93, 94–M. trinaria Puplesis&Robinson, Ven-
ezuela (USNM); 95, 96–M. forcipis Remeikis&Stonis, Peru (ZMUC); 97, 98–M. basi-
dactyla (Davis), Dominica (USNM)

125
e American Brachinepticula gen. nov. and Manoneura Davis (Nepticulidae)...
Enteucha basidactyla (Davis), in van Nieu-
kerkenetal., 2016a: 103.
Host plant. Coccoloba uvifera (L.) L., Po-
lygonaceae.
Distribution. USA (Florida, Davis, 1978),
Cuba (Núñez Aguila&Barro Cañamero, 2012),
Dominica (Puplesis&Robinson, 2000), Belize
(rainforest, Puplesis & Robinson, 2000), Ec-
uador (Amazon Basin, Puplesisetal., 2002b),
French Guiana (D.C.Lees, pers. comm.).
Discussion. Manoneura basidactyla pos-
sesses avery strong purple iridescence (lustre)
on theforewing (Figs.97, 98). is remarkable
species was re-desribed and illustrated in Pu-
plesis & Robinson, 2000: 22–23; here we add
only thedata lacking in theformer re-descrip-
tion: abdomen black to fuscous with yellowish
bronzy lustre on upper side, blackish grey on
underside; genital plates blackish grey to black,
not contrasting with themain color of theab-
domen, anal tus grey-black, short. Here we
also provide therst photographic documenta-
tion of thespecimens collected in theAmazon
Basin (Figs.85–92, 97, 98) rst reported by Pu-
plesisetal. 2002b; these specimens dier very
little from specimens from other localities (see
Distribution).
Manoneura trinaria Puplesis & Robinson,
2000 (Figs.82–84 93, 94, 99)
Manoneura trinaria Puplesis & Robinson,
2000: 23.
Manoneura trinaria Puplesis & Robinson,
in Puplesisetal., 2002b: 64; Diškus&Puplesis,
2003: 322.
Enteucha trinaria (Puplesis&Robinson), in
van Nieukerkenetal., 2016a: 103.
Host plant. Unknown.
Distribution. Venezuela (Puplesis&Robin-
son, 2000).
Discussion. Manoneura trinaria possesses
very strong purple iridescence on theforewing
(Figs. 93, 94). Originally the species was de-
scribed and illustrated in Puplesis&Robinson,
2000: 23; Figs.13, 86, 87. In thecurrent paper
we provide therst photographic documenta-
tion of theadult (Figs.93, 94) and male geni-
talia of M. trinaria (Fig. 85–92, 97, 98); note
theuncus turned caudally in Fig.78.
Figs.99–101. Distribution of Manoneura (Davis). 99–distribution map of thecurrently known species
(courtesy of T. Patterson, USA); 100, 101, habitat of M. basidactyla (Davis), theAmazon rainforest, Napo
Region, Ecuador

126 Jonas R. Stonis, Arūnas Diškus, Andrius Remeikis, Maria Alma Solis
Manoneura forcipis Remeikis & Stonis,
2017 (Figs.80, 81, 95, 96, 99)
Manoneura forcipis Remeikis & Stonis, in
Stonisetal., 2017: 58.
Host plant. Unknown.
Distribution. Central Andes (Peru: Apuri-
mac Department) (Stonisetal., 2017).
Discussion. is distinctive, highly apo-
morphic species was recently described and
illustrated by Stonis et al., 2017: Figs. 2, 9, 33,
95, 96 (note theunusually extended cathrema,
illustrated in Fig.81 of this paper).
A genus not discussed here, but possibly
related to Enteucha Meyrick.
Genus Va r i us Scoble, 1983
Varius Scoble, 1983: 14.
Type species: Stigmella ochnicola Vári, 1955:
336, 337.
Discussion. egenus is characterized by
less reduced forewing venation (see Scoble,
1983: Fig. 12), deeply divided valva, elongated
cathrema, and theabsence of cornuti in themale
genitalia. Female genitalia without vaginal scle-
rites. Single known species is trophically associ-
ated with Ochnaceae.
Varius ochnicolus (Vári, 1955)
Stigmella ochnicola Vári, 1955: 336, 337.
Varius ochnicolus (Vári), in van Nieuker-
kenetal., 2016a: 103.
Host plant. Ochna pulchra Hook., Ochnace-
ae (Malpighiales).
Distribution. Republic of South Africa:
Gauteng (Pretoria).
ACKNOWLEDGEMENTS
We are indebted to Ole Karsholt and the late
Professor NielsP. Kristensen (ZMUC) for
theinitial stimulus to start theNeotropical pro-
ject as well as for generous support during its
course and theloan of the Neotropical mate-
rial. For helpful and frequent discussions on
various host plants, we thank Dr.Arvind Singh
(Banaras Hindu University, Varanasi, India),
Dr.José Luis Fernández-Alonso (Universidad
de Salamanca, Spain), Dr.Franz Starlinger
(Federal Research and Training Centre for For-
ests, Natural Hazards and Landscape, Vienna,
Austria), Dr.Asok Ghosh (University of Burd-
wan, Barddhamān, India), and Dr.Nixon Cum-
bicus Torres (Universidad Técnica Particular de
Loja, Ecuador). We are very grateful to Lauri
Kaila (Finnish Museum of Natural History,
University of Helsinki, Finland) for helpful dis-
cussions. is study was supported by theRe-
search Foundation of theLithuanian University
of Educational Sciences.
Mention of trade names or commercial
products in this publication is solely for thepur-
pose of providing specic information and does
not imply recommendation or endorsement by
theUSDA. eUSDA is an equal opportunity
provider and employer.
Received 2 November 2017
Accepted 12 December 2017
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JonasRimantasStonis, ArūnasDiškus,
Andrius Remeikis, MariaAlmaSolis
AMERIKINĖMS BRACHINEPTICULA
GEN.NOV. IR MANONEURA DAVIS GENTIMS
PRIKLAUSANČIOS RŪŠYS SU NEĮPRASTAIS
PATINO LYTINIAIS LATAKAIS CATHREMA:
NAUJA GENČIŲ KONCEPCIJA
Santrauka
Straipsnyje aprašoma nauja mokslui Brachinepticula
gentis ir trys naujos rūšys: Brachinepticula plurilo-
bata sp. nov., B. elongatasp.nov. ir Johanssoniella
bina sp. nov. Pateikiamos diagnostinės schemos ir
nauja taksonominė koncepcija, kuria remiantis pir-
mą kartą pripažįstamos keturios giminingos gen-
tys: Johanssoniella Koçak, Brachinepticulagen.nov.,
Enteucha Meyrick ir Manoneura Davis. Genčių ir
rūšių kataloge pateiktos naujos taksonominės pa-
vadinimų kombinacijos ir nauji duomenys apie
kai kurių rūšių morfologiją, biologiją bei papliti-
mą. Pirmą kartą dokumentuojama Amazonijoje
surinkta Manoneura basidactyla (Davis), taip pat
Johanssoniella diplocosma (Meyrick) medžiaga iš
Himalajų. Pateikiami nauji tyrimų duomenys apie
iki šiol nežinomas europinės Johanssonia acetosae
(Stt.) patino genitalinių struktūrų morfologijos ypa-
tybes. Straipsnis itin gausiai iliustruotas lapų pažei-
dimų (minų) ir morfologinių struktūrų nuotrauko-
mis, piešiniais ir schemomis.
Raktažodžiai: Amerikos fauna, Brachinepticula
Stonis & Diškus, Enteucha Meyrick, Johanssoniella
Koçak, lapų minos, Manoneura Davis, nauja gentis,
naujos rūšys, Nepticulidae