Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus Paratischeria gen. nov.

Abstract

We describe a new genus Paratischeria Diškus & Stonis, gen. nov. and present the first observations of Urticaceae-feeding Tischeriidae species in South America and two new Urticaceaefeeding species (Paratischeria fasciata sp. nov. and P. ferruginea Diškus & Stonis, sp. nov. from the Andes of Bolivia and Ecuador). Together with the African P. urticicolella (Ghesquière) (comb. nov.) from Congo, they are attributed to the newly designated ferruginea species group.

Full text

BIOLOGIJA. 2017. Vol. 63. No. 1. P. 1–22
© Lietuvos mokslų akademija, 2017
Urticaceae-feeders from thefamily
Tischeriidae: descriptions of two new species
and new genus Paratischeriagen.nov.
*Corresponding author. E-mail: stonis@leu.lt
JonasR.Stonis1*,
ArūnasDiškus1,
BrigitaPaulavičiūtė2,
AlexK.Monro3
1Lithuanian University of Educational
Sciences and Baltic-American
Biotaxonomy Institute, StudentųSt.39,
Vilnius LT-08106, Lithuania
2Tadas Ivanauskas Zoological Museum,
Laisvės Ave.106,
Kaunas LT-44253, Lithuania
3Royal Botanic Gardens,
Kew, Richmond, Surrey TW9 3AE,
London, United Kingdom
We describe a new genus Paratischeria Diškus&Stonis, gen.
nov. and present the rst observations of Urticaceae-feeding
Tischeriidae species in South America and two new Urticaceae-
feeding species (Paratischeria fasciatasp. nov. and P. ferruginea
Diškus&Stonis, sp. nov. from theAndes of Bolivia and Ecuador).
Together with the African P. urticicolella (Ghesquière) (comb.
nov.) from Congo, they are attributed to thenewly designated fer-
ruginea species group.
Keywords: eAndes, leaf-mines, new genus, new species, Para-
tischeria, Phenax Wedd., South America, Tischeriidae, Urticaceae
INTRODUCTION
e Tischeriidae represents the relatively small
family of Lepidoptera. emost recent detailed
family review was provided by Diškus, Puplesis,
2003. Among themost distinctive diagnostic fea-
tures of the
family
is thelong length of male anten-
nal sensillae (Fig.67); females possess
much
short-
er sensillae, therefore, female
antennae
appear bare
in ventral view. Among other characters that also
have diagnostic importance for family recognition
are thefollowing: (1) frontal tu (Fig.26) project-
ing over triangular (or trapezoid) face smoothly
covered with scales; (2) sensillae trichodea with
strongly recurved
bases (Fig.67
); (3) strongly en-
larged 3rd antennal segment (Fig.67); (4) strongly
narrowed phallus (Figs.8, 33, 45), usually bifurcat-
ed or with spines at apex; (5) dark, short, strongly
thickened, stout peg setae on female ovipositor
(usually visible even without dissection) (Fig.29);
(6) specic rod-like or plate-like projections
of
modied 8th and 9th sternites in female genitalia
collectively referred as
prela (Fig.31)
.
Literature on Tischeriidae is still rather scarce
(for thehistory of research on thefamily inves-
tigation and a literature review, see Puplesis,
Diškus, 2003). Since then several papers have
been published, notably: Landry, Roque-Albelo,
2004; Puplesisetal., 2004; Mey, 2004, 2010; Puple-
sis, Diškus, 2005; Diškus, Stonis, 2006, 2012, 2015;

2Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Stonis, Diškus, 2007, 2008; Lees, Stonis, 2007;
Stonisetal., 2008, 2014, 2016; Hua, Cai, 2009;
van Nieukerken, 2010; Navickaitėetal., 2011;
Diškusetal., 2014; Kobayashietal., 2016.
Tischeriidae are trophically associated with
plants belonging to rosid and asterid I core eu-
dicot angiosperm families Fagaceae (Fagales),
Rosaceae, Rhamnaceae (Rosales), Malvaceae
(Tilioideae, Sterculioideae, Malvales), and As-
teraceae (Asterales). A few represantatives of
Tischeriidae have also been recorded on plants
from the Combretaceae (Myrtales), Euphor-
biaceae (Malpighiales), Ericaceae, Symploca-
ceae (Ericales), Anacardiaceae (Sapindales),
eaceae (Ericales), Hypericaceae (Malpighi-
ales), Betulaceae (Fagales), and Apocynaceae
(Gentianales) (Diškus, Puplesis, 2003a, 2003b;
Kobayashietal., 2016). In temperate regions
the family is strongly associated with plants
from theRosaceae and Fagaceae, in theAmeri-
cas also with Malvaceae and Asteraceae. How-
ever, only a single record of the Urticaceae-
feeding species was previously known from
equatorial Africa (Puplesis, Diškus, 2005), and
no records from theAmericas.
In this paper, we review our recent ndings
in South America together with the previous
record of theUrticaceae-feeding species from
Congo, equatorial Africa (Fig.1). We illustrate
theAfrican species and describe two new Tisch-
eriidae species from theAndes: one trophically
associated with Phenax hirtus, other feeding on
unidentied Urticaceae plant but not Phenax.
Host plant Urticaceae is a medium-sized
family comprising ca 2,500 species and 50 gen-
era of woody shrubs, succulent herbs, trees and
vines that are found on all of theworld’s conti-
nents with theexception of Antarctica. Ashort
characterization of Urticaceae will be provided
in our other paper on theUrticaceae-feeding
Nepticulidae (Stonisetal., submitted).
MATERIALS AND METHODS
Descriptions of new species are based on
the material deposited in the collection of
theZoological Museum, Natural History Mu-
seum of Denmark in Copenhagen, Denmark.
Collecting methods and protocols for spe-
cies identication and description are outlined
in Puplesis, Diškus (2003) and Diškus, Stonis
(2012). Aer maceration of theabdomen in 10%
KOH and subsequent cleaning, male genital
capsules were removed from the abdomen and
mounted ventral side uppermost. e phallus
was removed and mounted in Euparal separately
but on thesame genitalia slide. Abdominal pelts
and female genitalia were stained with Chlorazol
Black (Direct Black 38/Azo Black) (for adetailed
method’s description see Stonisetal., 2014).
Permanent slides were photographed and
studied using aLeica DM2500 microscope and
a Leica DFC420 digital camera. edescriptive
terminology of morphological structures fol-
lows Puplesis, Diškus (2003), except for theterm
“aedeagus” that is referred here as “phallus”, and
theterm “cilia” that is referred here as “fringe”.
For DNA extraction we used specimens
which were stored in 96% ethanol. Total
genomic DNA was extracted from head or tho-
rax using a Nucleospin Tissue Kit (Machery-
Nagel, Düren, Germany) according to thepro-
tocol of themanufacturer.
Amplication of a 577-bp fragment of
themtDNA region was performed using mito-
chondrial cytochrome c oxidase subunit Igene
(COI): LCO1490 (5’-GGT CAA CAA ATC ATA
AAG ATA TTG G-3’) and HCO2198 (5’-TAA
ACT TCA GGG TGA CCA AAA AATCA-3’)
(Folmeretal., 1994; Herbertetal., 2003a,
2003b). PCR were performed in 25µl volumes
including: 2µL DNA, 2µL of each primer (MBI
Fermentas, Lithuania), 0.5µL of Amplitaq
DNA polymerase (5U/µl), 2.5µl 25mM MgCl2,
2.5µL 10X Buer (Fermentas) and 1µL 10mM
dNTP (Fermentas) and water.
PCR were carried out in an Eppendorf Mas-
tercycler gradient, 5331 (Germany). Cycling
parameters were an initial denaturation step
at 94°C for 2min, followed by 94°C for 30s,
50°C for 45s, and 72°C for 1min. is cy-
cle was repeated 35 times, followed by 4 min of
extension at 72°C. PCR products were visual-
ized on 1.5% agarose gel stained with ethidium
bromide and UV light photographs of thegels
with DNA bands were taken using “Herolab”

3
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
transluminator (Germany). epositive bands
of the expected size range were excised from
thegel. DNA was extracted using theGeneJet
Gel Extraction kit (Fermentas, Lithuania). Am-
plicons were then sequenced in both directions
on an automated sequencer, ABI 3130xl (Ap-
plied Biosystems, USA).
Electropherograms were checked by eye
for poor base calls and sequence quality and
then sequences were edited and aligned using
theClustalW (omsonetal., 1994) algorithm
in MEGA v.7.0.21 (Kumaretal. 2015). Phy-
logenetic relationships among control region
haplotypes were reconstructed using theneigh-
bour-joining (NJ) method (Saitou, Nei, 1987).
NJ analysis was performed using theKimura-2
paratemeter correction model (Kimura, 1980)
by bootstrapping with 1000 replicates.
Partial fragment of Cytochrome oxidase I
(COI) gene were amplied in 21 Tischerii-
dae specimens. Amplied fragment produced
a577-bp sequence which corresponded to COI
gene sequence position 2239–2944.
e partial nucleotide sequences of theTis-
cheriidae species COI gene were added to
the GenBank database. Sequences accession
numbers are presented in Table.
Institutional abbreviations used in the text
of the current paper: MRAC – Museé Roy-
al de l’Afrique Centrale, Tervuren, Belgium;
ZMUC – Zoological Museum, University of
Copenhagen, Denmark.
Table 1. GenBank accession numbers of examined specimens of Tischeriidae
Examined specimens Accession number GenBank
Coptotriche marginea(1) (Haworth, 1828) HM244382
Coptotriche marginea(2) (Haworth, 1828) HM244379
Coptotriche marginea(3) (Haworth, 1828) HM244380
Coptotriche marginea(4) (Haworth, 1828) HM244381
Tischeria ekebladella(1) (Bjerkander, 1795) HM244384
Tischeria ekebladella(2) (Bjerkander, 1795) HM244385
Tischeria ekebladella(3) (Bjerkander, 1795) HM244383
Tischeria dodonea(1) Stainton, 1858 HM244386
Tischeria dodonea(3) Stainton, 1858 HM244387
Astrotischeriasp. 4848 HM244389
“Astrotischeria”sp. 4877, Paratischeriasp. HM244393
“Astrotischeria”sp. 4894, Paratischeria ferruginea
Diškus&Stonis, sp. nov.
HM244395
“Astrotischeria”sp. 4909.1, Paratischeria neotropicana
(Diškus&Stonis, 2015)
HM244391
“Astrotischeria”sp. 4909.2, Paratischeria neotropicana
(Diškus&Stonis, 2015)
HM244392
Astrotischeriasp. 4910 HM244390
“Astrotischeria”sp. 4936, Paratischeria ferruginea
Diškus&Stonis, sp. nov.
HM244394
“Astrotischeria”sp. 4937, Paratischeria neotropicana
(Diškus&Stonis, 2015)
HM244396
“Astrotischeria”sp. 4939, Paratischeriasp. HM244398
Astrotischeriasp. 4944.2 HM244397
Astrotischeriasp. 4957.1 HM244399
Astrotischeriasp. 49572 HM244400

4Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
RESULTS
Taxonomy of theUrticaceae-feeding Tisch-
eriidae
ree currently known Urticaceae-feeding Ti-
scheriidae, including one Afrotropical and
two Neotropical species, together with some
American Malvaceae and Asteraceae-feeding
species, belong to a new genus, Paratischeria
Diškus&Stonis, gen. nov., which is described
below.
Genus Paratisheria Diškus&Stonis, gen. nov.
Type species: Paratischeria ferruginea Diš-
kus & Stonis, sp.nov.
Diagnosis. In the male genitalia, the new
genus diers from Astrotischeria in theundi-
vided, usually narrow valva (divided in Astro-
tischeria), long undivided uncus (usually short
and divided in Astrotischeria), and thestrongly
though variously developed anellus (Figs.48–
53); from
Coptotriche it diers
in theslender
valva, well developed anellus with lateral papil-
lae and setae (Figs.48–53), also in theabsence
of
transtilla, absence of
spines on thediaphrag-
ma, larger vinculum, and dierently shaped
phallus (not tulip-shaped as in Coptotriche);
from Tischeria it diers in theabsence of
juxta,
presence of
strongly developed anellus (absent
in Tischeria), and usually larger vinculum. A
lso
see Figs.73, 74.
Head (Figs.54–67)
.
Scape with apecten. 3rd
antennal segment greatly enlarged.
Wing venation (Figs.68–70). Greatly re-
duced, similar to Astrotischeria (Fig. 71). In
the
forewing,
vein Sc almost reaching middle
of wing; R with ve short radial branches
ter-
minat
ing on costa; radial cell reduced; M with
two short
branches
distally and with base ab-
sent; vein Cu single, very long and straight; 2A
well-developed,
but simple; in hindwing, Sc
very short; R, M and Cu expressed only in dis-
tal half (if
not
entirely reduced), vein Asingle
or represented by two short
veins.
Abdomen (Fig.72). Male with ashort anal
tu
of piliform scales.
Male
genitalia.
Uncus with long lateral
lobes (Fig.38). Socii not thickened, similar to
those in Astrotischeria. Tegumen large, with
strongly sclerotized anterior processes; poste-
riorly not extending into alobed pseuduncus.
Diaphragma smooth or with membranous
wringles but without
spines. Valva relatively
slender, undivided. Transtilla absent. Anel-
lus
always well-developed, slightly thickened
laterally
and
with three or four pairs of papillae
and setae, sometimes additionally with ador-
sal sclerite (pseudotranstilla) (Figs.48–53).
Juxta
absent.
Vinculum moderately large to
very large, anteriorly either broadly rounded
or gradually
narrowed. Phallus
long and dis-
tinctly bifurcate in apical half or apical third
(Figs.33, 45
).
Female
genitalia.
Ovipositor lobes large
(Figs.29, 31); thesecond pair of lobes very small.
Anterior and posterior apophyses supplemented
with three pairs of shorter rod-like and plate-like
sclerites (prela) (Fig.31). Vestibulum membra-
nous or hardened but antrum absent
.
Corpus
bursae slender, as long as anterior apophyses or
longer, usually
without
pectination; signa ab-
sent
. Ductus spermathecae
membranous, slen-
der, with afew to many convolutions.
Biology. Larvae produce bloth-like mines
(Figs. 11–19) in leaves of plants from Malva-
ceae, Urticaceae, and Asteraceae.
Distribution.
Known from theNew World
(the Americas) and equatorial Africa. ere are
many species from Central and South America
which have already been dissected, identied
but still undescribed (Stonisetal., in prep).
Ety mology. ename combines Tischeria
(a name of thetype genus of thefamily) and
Greek prex para (at or to one side of, side by
side, beyond).
e ferruginea species group
Designated here for therst time for thePa-
ratischeria species feeding on Urticaceae and
possessing a dorsal sclerite (pseudotranstilla)
in themale genitalia.
Diagnostics: forewing dark brown to brown
(not sparsely speckled as is usual in other Pa-
ratischeria or Astrotischeria), with spot(s)
(Figs. 5, 26), fascia (Fig.5), or without spots
or fascia, uniform. In male genitalia, phallus
deeply divided in distal half (Figs.7, 8, 32–34,

5
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
Fig. 1. Currently known records of Tischeriidae trophically associated with thegenera of Urticaceae: 1–Pa-
ratischeria urticicolella (Ghesquière, 1940) on Fleurya podocarpa Wedd. from Congo (see Puplesis, Diškus,
2005); 2–P. ferruginea Diškus&Stonis, sp. nov. on Phenax hirtus (Sw.) Wedd. from Pichincha Province,
Ecuador; 3–P.fasciata Diškus&Stonis, sp. nov. on unidentied Urticaceae plant, Coroico, Bolivia;
*–earlier, by Ghesquière, 1940, thehost plant was reported as Laportea podocarpa Wedd., however, thegenus
Laportea has been split and thehost plant is attributed now to Fleurya. Fleurya podocarpa Wedd. is placed as
asynonym of ovalifolia, however, such combination of ovalifolia with Fleurya has not been published.
45); valva moderately slender (but wider than
in many other Paratischeria species) or strongly
widened in basal half (Fig. 36); basal process
of valva long (Figs.6, 39, 44); uncus with two
long (Fig.38) to very long lobes (Fig.46); an-
ellus with a dorsal sclerite (pseudotranstilla)
(Figs. 42–44); vinculum large but smaller
than in most of other Paratischeria, triangular
(Fig.35) or distally broadly rounded (Figs.6,
44).
Currently thegroup comprises three equa-
torial or subequatorial species (two from
theAndes of Ecuador and Bolivia), one from
Africa (Congo); all species are trophically as-
sociated with plants from Urticaceae (Phenax
and Fleurya).
Paratischeria fasciata Diškus&Stonis, sp. nov.
Type material. Holotype: ♂, BOLIVIA:
Nor Yungas Province, Coroico, 16°11’39’’S,
67°43’21’’W, elevation 1880 m, mining larva on
unidentied Urticaceae plant 25.iv.2014, ex pupa
v.2014, A. Diškus, genitalia slide no.AD868♂
(ZMUC). Paratype: 1♂, same label data as holo-
type (ZMUC).
Diagnosis. Externally, thenew species dif-
fers from all other known tischeriids by adis-
tinctive fascia of theforewing. In male genitalia,
aspecic shape of phallus and dorsally devel-
oped anellus distinguish Paratischeria fascia-
tasp. nov. from all other Tischeriidae species;
the Urticaceae host plant, shared with three
other related species, also makes this taxon dis-
tinctive.
Male (Figs. 4, 5). Forewing length 3.0–
3.2mm; wingspan 6.5–6.8mm. Head: face
smoothly scaled, palpi and face grey-brown or
olive brown, very glossy and with some golden
shine and purple iridescence; frontal tu grey-
brown to olive brown, smooth, not erected

6Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Fig. 2. Urticaceae clades engaged by leaf-mining Tischeriidae (simplied phylogeny aer Wuetal., 2013.
Note: this is not an ultrametric tree)
*–for detailed relationships of theclades, see Wuetal., 2013
**–originally, thehost plant was reported as belonging to thegenus Laportea (see Ghesquière, 1940), how-
ever, thegenus Laportea has been split and thehost plant is attributed now to genus Fleurya.
but overlapping over face; antenna slightly
longer than half thelength of forewing, grey-
brown, golden glossy, with long hair-like
sensillae trichodea sticking out. orax and
tegula black-brown with strong golden gloss
and some purple iridescence. Forewing black-
brown with some golden gloss and purple and
blue iridescence; median fascia and basal spot
yellow orange, very distinctive; fringe dark
brown with some golden gloss; underside of
forewing brown-black, without spots except
small irregular scaleless patch at the base.
Hindwing dark brown on upper side, brown-
black on underside, without androconia or
spots except small irregular scaleless patch
at the base on underside; its fringe blackish
brown. Legs grey, glossy. Abdomen black-
brown on upper side, dark grey, glossy on un-
derside; genital segments dark grey-brown to
grey, distally white; anal tus short, blackish
brown.
Female. Unknown.
Male genitalia (Figs.6–10). Capsule about
455μm long. Uncus with two long lateral lobes.
Valva about 350μm long, not divided, without
processes (except thebasal one); transtilla ab-
sent. Anellus thickened laterally and with setae
(Figs.6, 51), dorsally with aspecic plate-like
sclerite (pseudotranstilla) (Figs.6, 43). Vin-
culum rather short, rounded distally. Phallus
(Figs.7, 8) 455μm long, with three pairs of
large carinae.

7
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
Fig. 3. Current records of Urticaceae-feeding Tischeriidae from South America
Bionomics. Host plant belongs to Ur-
ticaceae (species and genus unidentied;
host-plant sample neither preserved nor
documented). Larvae mine in leaves in
April. A leaf-mine similar to those illus-
trated in Figs.11–19). Exit slit on theupper
side of the leaf. According to the “Formula
of Determining of Abundance and Occur-
rence of Leaf-Miners” (see Diškus, Stonis 2012:
52–54), Paratischeria fasciata is arare species:
sparse mining of thenew species was observed
in asingle locality of theBolivian Andes.
Distribution (Figs.1, 3). is species occurs
on theeastern slopes of theBolivian Andes in
tropical montane forests at altitudes ca. 1880m.
Ety mology. e species name is derived
from the Latin fascia/fasciata (a band, fascia/
with fascia) in reference to thedistinctive fascia
on theforewing, so unusual among theTisch-
eriidae.

8Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Figs. 4–10. Paratischeria fasciata Diškus&Stonis, sp. nov.: 4, 5–male adult, holotype; 6, genita-
lia slide no. AD868, capsule with phallus removed; 7, 8–same, phallus; 9–same, uncus and socii;
10–same, basal part of valvae (ZMUC)
Paratischeria ferruginea Diškus&Stonis,
sp.nov.
Type material. Holotype: ♂, ECUA-
DOR: Pichincha Province, 11km NW Alóag,
0°26’45’’S, 78°37’34’’W, elevation 3090 m,
mining larvae on Phenax hirtus (Urticaceae)
20.xi.2007, eld card no.4936, A.Diškus, gen-
italia slide no.AD775♂ (ZMUC). Paratypes:
4 ♂ (including agenitalia and head slide of
one adult in thepupal skin, with no pinned

9
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
Figs. 11–19. Leaf-mines of Paratischeria ferruginea Diškus&Stonis, sp. nov. on Phenax hirtus, Ecuador,
Pichincha Province, 11km NW Alóag, 0°26’45’’S, 78°37’34’’W, elevation 3090m

10 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
specimen preserved), 5 ♀, same label data
as holotype, slide nos. AD839♂ (genitalia),
AD866♂ (genitalia), AD865 (head), AD863
(forewing venation), AD867♂ (genitalia and
head; both from adult in thepupal skin with
no pinned specimen preserved), AD840♀
(genitalia) (ZMUC); 1♂, 1♀, same locality
as holotype, mining larvae on Phenax hirtus
Figs. 20–28. Paratischeria ferruginea Diškus&Stonis, sp. nov.: 20, 22–pupae; 21, 23–25–pupal exuviae;
26–adult male, holotype; 27–adult female, paratype AD840; 28–adult male, paratype AD841 (ZMUC)

11
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
24.ii.2007, eld card no.4894, Diškus and
Stonis, slide no. AD841♂ (dissected genita-
lia) (ZMUC); 1♂, 1♀, Loja Province, 5km
SW Saraguro, Washapamba Forest Reserve,
3°39’43’’S, 79°16’9’’W, elevation 2940m, min-
ing larvae on Phenaxsp. (probably Ph.hirtus
or closely related species) 25.i.2017, eld card
no.5233, A.Diškus (ZMUC).
Figs. 29–31. Female genitalia of Paratischeria ferruginea Diškus& Stonis, sp. nov., paratype,
genitalia slide no. AD840: 29–apophyses and ovipositor; 30–vestibulum; 31–general view of
genitalia (ZMUC)

12 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Figs. 32–39. Male genitalia of Paratischeria ferruginea Diškus&Stonis, sp. nov.: 32, phallus, holotype,
genitalia slide AD775; 33–same, paratype, genitalia slide no.AD867; 34–same, paratype, genitalia slide
no. AD841; 35–general view, AD866; 36–37–capsule, holotype, AD775; 38–uncus and tegumen, para-
type, AD839; 39–dorsal sclerite of anellus (pseudotranstilla), paratype, AD839 (ZMUC)

13
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
Figs. 40–47. Male genitalia of theferruginea group: 40, 41 – capsule of Paratischeria ferruginea
Diškus&Stonis, sp. nov., paratype, genitalia slide no. AD867; 42, same, genitalia slide no.AD841;
43 – dorsal sclerite of P. fasciata Diškus & Stonis, sp. nov., holotype, genitalia slide no.AD868
(ZMUC); 44–ventral view of capsule of P. urticicolella (Ghesquière, 1940) (comb. nov.), holotype,
genitalia slide no.AD091 (MRAC); 45–same, phallus; 46–same, lateral view of capsule, slide no.
AD091 (MRAC); 47–lateral view of capsule of P. ferruginea Diškus&Stonis, sp. nov., paratype,
genitalia slide no.AD841 (ZMUC)

14 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Figs. 48–53. Anellus of Paratischeria Diškus&Stonis, gen. nov.: 48, 49–P. sp., genitalia slide AD844;
50 – P. neotropicana (Diškus & Stonis, 2015), comb. nov., genitalia slide AD861; 51, 52 – P. fasciata
Diškus&Stonis, sp. nov., genitalia slide AD868; 53–P. ferruginea Diškus&Stonis, sp. nov., genitalia slide
AD867 (ZMUC)
Diagnosis. Externally, thenew species dif-
fers from all other known tischeriids by aunique
pattern of forewing with one distinctive ochre-
beige spot. In male genitalia, aspecic shape of
bilobed valva and dorsally developed anellus dis-
tinguish Paratischeria ferrugineasp. nov. from all
other Tischeriidae species; thehost plant Phenax
hirtus also makes this species distinctive.
Male (Figs. 26, 28). Forewing length 4.3–
5.0mm; wingspan 9.1–10.7mm. Head: face
dark grey-brown with golden gloss and purple
iridescence; palpi and proboscis ochre; frontal
tu grey-brown with golden gloss and little or
strong purple iridescence; collar grey-brown
with golden gloss and sometimes with strong
purple iridescence; antenna signicantly long-
er than half thelength of forewing, grey-brown
with some golden gloss on upper side, paler
on underside, with rather indistinctive hair-
like sensillae trichodea. orax and tegula

15
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
Figs. 54–60. Descaled head of Paratischeria Diškus&Stonis, gen. nov.: 54, 55–P. sp., slide no. AD876♀
[5116]; 56 – P. sp., slide no. AD874♀; 57 – P. neotropicana (Diškus & Stonis, 2015), comb. nov., slide
no.AD861; 58–P. ferruginea Diškus & Stonis, sp. nov., paratype, slide no. AD865; 59, 60, same, paratype,
dissected from adult in pupal skin, slide no.AD867 (ZMUC)

16 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Figs. 61–67. Descaled head of Paratischeria Diškus&Stonis, gen. nov.: 61 – P. sp., slide no. AD874♀
[5109]; 62–P. sp., slide no. AD876♀ [5116]; 63–P.neotropicana (Diškus&Stonis, 2015), comb. nov., slide
no.AD861; 64, 65–P. ferruginea Diškus&Stonis, sp. nov., paratype, slide no.AD865; 66, same, paratype,
slide no.AD867; 67–same, paratype, slide no.AD865 (ZMUC)

17
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
dark grey-brown to black-brown, with golden
gloss and strong purple iridescence. Forewing
dark grey-brown with some purple and blue
iridescence; median spot along costal margin,
ochre beige (ferruginous), usually large and
very distinctive, occasionally small and indis-
tinctive; fringe grey-brown with some golden
gloss (particularly prominent at costa), dis-
tinctly ochre at thevery apex of forewing; un-
derside of forewing dark grey-brown, without
spots except small, irregular scaleless patch
at thebase. Hindwing glossy, grey-brown to
olive grey on theupper side, grey-brown on
underside, without androconia or spots except
very small irregular scaleless patch at thebase
on underside; fringe grey-brown. Legs glossy,
grey-brown, except femur of middlegs which
distinctly ochre beige (ferruginous); tibia of
hindlegs with alarge tu of ochre yellow pili-
form scales. Abdomen dark brown with some
golden gloss on theupper side, bright yellow
ochre to orange yellow and glossy on under-
side; genital segments dark grey-brown; anal
tus indistinctive.
Female (Fig. 27). Forewing length 4.0–
4.5mm; wingspan 8.7–9.7mm. Antenna
shorter, sensillae trichodea indistinctive, 8–12
distal segments greyish white. Legs dark grey
on theupper side, mostly ochre beige on un-
derside. Abdomen dark brown on the upper
side, bright yellow ochre yellow and glossy on
underside. Otherwise as in male.
Male genitalia (Figs.32–42, 47, 53). Capsule
about 420–470μm long, 305μm wide. Uncus
with two long lateral lobes. Valva about 320–
375μm long, slender in distal half, with wide
lobe basally; transtilla absent. Anellus slightly
thickened and with setae laterally (Fig. 40),
dorsally with aspecic plate-like sclerite (pseu-
dotranstilla) (Figs.41, 42, 47). Vinculum trian-
gular. Phallus (Figs.32–34) 500–570 μm long,
distally 140–150 μm wide, distinctly bilobed;
each lobe with atiny spine.
Female genitalia (Figs.29–31). Ovipositor
clothed in short, stout and darker, modied se-
tae which we refer to as ‘peg setae’ (Fig.29). Sec-
ond pair of lobes, lateral and anterior to theovi-
positor lobes, are three times smaller, bearing
Figs. 68–72. Forewing venation and abdominal pelt: 68, forewing venation, Paratischeria ferruginea
Diškus&Stonis, sp. nov., slide no. AD863; 69–same, P. neotropicana (Diškus&Stonis, 2015), comb. nov.,
slide no.AD862; 70–same, slide no. AD861; 71–same, Astrotischeriasp., slide no.AD864; 72–fragment
of abdominal pelt, lateral view, P. ferruginea Diškus &Stonis, sp. nov., slide AD841 (ZMUC)

18 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Figs. 73, 74. Molecular phylogeny of Tischeriidae: 73–neighbour-joining tree for specimens of Tischerii-
dae with Elachista alpinella (GU248257.1) as outgroup; bootstrap values are shown above; 74–same, other
presentation

19
Urticaceae-feeders from the family Tischeriidae: descriptions of two new species and new genus...
long slender setae. Anterior and posterior apo-
physes very long and slender (Fig.29). Addi-
tionally there are three rod-like and plate-like
projections (of possibly modied 8th and 9th
sternites) collectively referred to as theprela de-
scribed by Braun (1972) (Figs.30, 31). Ductus
bursae considerably narrower than corpus bur-
sae, without spines but with a hardened plate
(Fig. 30). Corpus bursae membranous, rela-
tively very small (500μm long, 200μm wide)
(Fig. 31), without spines or signum. Ductus
spermathaecae with many large coils (Fig.31).
Bionomics (Figs.11–25). Host plant: Phenax
hirtus (Sw.) Wedd. (Urticaceae) (Fig.11). Larvae
mine in late January–February and October–
November. Leaf-mine as a blotch (at the early
stage triangular or distally widely rounded, later
very irregular), without frass. Larva pale green-
ish grey, with pale grey-brown intestine. Pupa-
tion inside of the leaf-mine without cocoon;
pupa brown (Figs.20–25). Exit slit on theupper
side of the leaf. Adults known from February,
March, and December. According to the“For-
mula of Determining of Abundance and Occur-
rence of Leaf-miners” (see Diškus&Stonis 2012:
52–54), P. ferruginea is not arare species: limited
in distribution, but abundant mining in thelo-
cality where it has been recorded.
Distribution (Figs.1, 3). is species oc-
curs in theEcuadorian Andes in tropical mon-
tane and cloud forests at altitudes about 2940–
3100m.
Ety mology. especies name is derived
from the Latin ferruginea (brown, yellowish
brown) in reference to the distinctive yellow
beige spot of the forewing and same ferrugi-
nous abdomen on underside.
DISCUSSION
Within thefamily Tischeriidae, analysis of mor-
phological characters indicates at least four main
lineages of ageneric rank (Stonisetal., in prep.).
During our molecular studies, partial frag-
ment of the COI gene were amplied in 21
specimens of Tischeriidae resulting in a577-bp
fragment. e sequences have 147 polymor-
phic nucleotides. 145 nucleotides were found to
be parsimony-informative characters. All four
lineages indicated by morphological studies
were well supported by our molecular data; see
thecladogram (NJ tree) in Figs.73, 74. eNJ
tree of Tischeriidae (Fig.74) has two big clus-
ters: I and II. ecluster I includes species from
thegenus Tischeria: European T.dodonaea and
T.ekebladella. Among these two species, 24 pol-
ymorphic nucleotides were detected.
Cluster II is formed of two groups: therst
group (A) is represented by Coptotriche (Eu-
ropean Coptotriche marginea) and thesecond
group (B) by thespecies of South American As-
trotischeria and Paratischeria. eCOI gene se-
quences of investigated specimens from cluster
B indicated 52 polymorphic and 525 conserva-
tive nucleotides.
ere is adierence between thecladogram
published by Diškus, Puplesis (2003b) and our
NJ tree (Fig.74). Previously two clades, Tisch-
eria and Astrotischeria, were shown as sibling
(Diškus, Puplesis, 2003); in our NJ tree, thesib-
ling group is Coptotriche and Astrotischeria
+ Paratischeria; other major cluster presents
thegenus Tischeria.
Our study has identied several distinct
groups which are supported by the COI gene
region (it also has shown that this particu-
lar gene is suitable for the reconstruction of
the Tischeriidae phylogeny). However, more
molecular data and further studies are neces-
sary to conrm thedetected cladas in our pre-
liminary cladogram.
e conducted specic molecular research
provides atheoretical and practical framework
for afurther use of DNA molecular markers in
taxonomy and phylogenetic studies of Tisch-
eriidae.
ACKNOWLEDGEMENTS
For helpful and frequent discussions on various
host plants (not only those which are listed in
thecurrent paper) we thank Nixon Cumbicus
Torres (Universidad Técnica Particular de Loja,
Loja, Ecuador), Arvind Singh (Banaras Hindu
University, Varanasi, India); Narayanan Nair
Mohanan (Jawaharlal Nehru Tropical Botanic

20 Jonas R. Stonis, Arūnas Diškus, Brigita Paulavičiūtė, Alex K. Monro
Garden and Research Institute, India), José Luis
Fernández-Alonso (Universidad de Salamanca,
Spain), eodor C.H.Cole (Universität Heidel-
berg, Germany), Maximilian Weigend (Univer-
sity of Bonn, Germany), and Franz Starlinger
(Federal Research and Training Centre for For-
ests, Natural Hazards and Landscape, Vienna,
Austria).
Further we would like to thank Andrius Re-
meikis (Nature Research Centre, Lithuania) for
providing us with photographs of genitalia and
adults of Tischeriidae.
Received 3 January 2017
Accepted 30 March 2017
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JonasRimantasStonis, ArūnasDiškus,
BrigitaPaulavičiūtė, AlexK.Monro
URTICACEAE AUGALUS MINUOJANTYS
TISCHERIIDAE: DVI NAUJOS MOKSLUI
RŪŠYS IR NAUJA PARATISCHERIA GENTIS
Santrauka
Šiame straipsnyje pirmą kartą publikuojami duo-
menys apie Urticaceae šeimos augalus minuo-
jančius Tischeriidae Pietų Amerikoje. Aprašoma
nauja Paratischeria gentis ir dvi naujos mokslui
rūšys: Paratischeria fasciata (aptikta Bolivijoje) ir
P. ferruginea (aptikta Ekvadore). Šios dvi rūšys kar-
tu su pusiaujo Afrikoje paplitusia P. urticicolella
(Ghesquière) (comb. nov.) pirmą kartą išskiriamos
į atskirą Paratischeria genties rūšių grupę, kuriai
būdingas pakitęs patino anellus ir dilgėlinių augalų
(Phenax, Fleurya) minavimas.
Raktažodžiai: Andai, nauja gentis, nauja rū-
šis, Paratischeria, Pietų Amerika, Phenax Wedd.,
Tischeriidae, Urticaceae