Dtsch. Entomol. Z. 71 (1) 2024, 1-15 | DOI 10.3897/dez.71.104130

> PENSUFT.

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BERLIN

Two new genera and four new species of Neotropical Eccritotarsin1

(Heteroptera, Miridae, Bryocorinae)

Eugenia Minghetti!, Sara Itzel Montemayor’, Pablo Matias Dellapé?

1 Division Entomologia, Museo de La Plata, Universidad Nacional de la Plata, CONICET. Paseo del Bosque s/n B1900F WA, La Plata, Buenos Aires,

Argentina

https://zoobank. org/A0C8 103C-B56C-40E9-9C38-1D2617F8A3C4

Corresponding author: Eugenia Minghetti (eugeniaminghetti@fcnym.unlp.edu.ar)

Academic editor: David Rédei # Received 27 March 2023 @ Accepted 20 November 2023 @ Published 8 January 2024

Abstract

In this study two new genera and four new species of Eccritotarsini (Hemiptera: Heteroptera: Miridae: Bryocorinae) are described.

Thomasomiris gen. nov. is established to accommodate 7homasomiris setosus sp. nov. (from Panama) and /gerocoris gen. nov., 1s

described for . ecuatorianus sp. nov. (the type species), £. dimorphus sp. nov. (both from Ecuador) and /. chaparensis sp. nov.

(from Bolivia). Adult habitus and male genitalia photographs are provided for each species. A key to species is also provided and

the affinities of the new genera are discussed.

Key Words

Insecta, Hemiptera, Bolivia, Ecuador, Panama

Introduction

Eccritotarsini is the most diverse among the five tribes
currently recognized in Bryocorinae (Hemiptera: Het-
eroptera: Miridae) (Namyatova et al. 2016), with about
115 genera (Konstantinov and Knyshov 2015; Schuh and
Weirauch 2020) and more than 628 species worldwide
(Ferreira et al. 2015; Konstantinov et al. 2018; Henry and
Menard 2020). The New World has the greatest diversity
with over 70 genera and more than 450 species (Ferreira
et al. 2015; Henry and Menard 2020). Carvalho and Fer-
reira (1995) keyed 66 Neotropical genera and Ferreira
and Henry (2011) provided a key to 18 genera occurring
in Minas Gerais, Brazil.

Neotropical eccritotarsines can be recognized by the
reduced scent gland evaporative area; the large disc-
shaped pulvilli covering nearly the entire inner surface
of the claws, with a comb-like row of long setae on its
outer surfaces (Konstantinov and Knyshov 2015; Namya-
tova et al. 2016; Konstantinov et al. 2018; Schuh and

Weirauch 2020); the bothrium of femoral trichobothria
being deeply recessed and tuberculate (Konstantinov
and Knyshov 2015; Namyatova et al. 2016; Schuh and
Weirauch 2020); the asymmetrical setiform parempodia
(Konstantinov and Knyshov 2015; Konstantinov et al.
2018; Schuh and Weirauch 2020); and the female gen-
italia possessing an asymmetric, complexly sclerotized
vestibulum (Namyatova et al. 2016).

The Eccritotarsini have a great morphological hetero-
geneity, including their general appearance, size, color-
ation, and genitalia (Konstantinov and Knyshov 2015;
Namyatova et al. 2016; Konstantinov et al. 2018). Males
exhibit several projections around the aperture of the
genital capsule (Mu and Liu 2012; Henry and Howard
2016; Konstantinov and Zinovjeva 2016; Konstantin-
ov et al. 2018; Menard and Schwartz 2018; Henry and
Menard 2020), and internally (Menard and Schwartz
2018). The parameres are diverse, with the right param-
ere usually being larger and more complex than the
left one (Konstantinov 2003; Namyatova et al. 2016;

Copyright Eugenia Minghetti et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted
use, distribution, and reproduction in any medium, provided the original author and source are credited.

2

Konstantinov et al. 2018; Menard and Schwartz 2018;
Henry and Menard 2020; Henry 2022). The aedeagus
can be simple, membranous, or complex with sclero-
tizations in different areas (Kerzhner and Konstantinov
1999; Konstantinov 2003; Henry and Howard 2016;
Konstantinov et al. 2018). The ductus seminis is entirely
membranous or with its apical part typically sclerotized
and may be needle-shaped, extends to mouth of the phal-
lotheca in repose, and can be attached to the endosoma
(Kerzhner and Konstantinov 1999; Konstantinov 2003;
Konstantinov et al. 2018; Menard and Schwartz 2018;
Henry and Menard 2020; Henry 2022). According to
Namyatova et al. (2016) Eccritotarsini 1s the sister group
of Bryocorini, but the phylogenetic relationships among
its genera are poorly resolved.

In the present contribution two new genera, the first
one with one new species from Panama and the other one
with three new species from Ecuador and Bolivia, are de-
scribed and illustrated, and their affinities are discussed.
Colour images of the adult male and female and of the
male genitalia, except for one of the species that is known
only from females, are provided, together with a key to
facilitate identification of the species.

Materials and methods

Specimens are deposited in the Museo de La Plata, La
Plata, Argentina (MLP) and the National Museum of
Natural History, Smithsonian Institution, Washington,
DC, USA (USNM).

Label data are cited verbatim. Lines on labels are sep-
arated with ‘/’, contents of different labels are separated
with ‘//’, remarks are given between square brackets ‘[]’.

All measurements are given in millimetres; measure-
ments of the holotype between brackets; identical measure-
ments of the same structure of different specimens are pro-
vided only once. Anterior width of pronotum is defined as
the width across behind margin of collar, and posterior width
of pronotum as the width across humeral angles. Genital
structures were dissected under stereomicroscope, cleared in
85% lactic acid solution for 45 min, washed in distilled wa-
ter and preserved in vials with glycerinol. Terminology for
male genitalia follows Kerzhner and Konstantinov (1999),
Konstantinov (2003) and Menard and Schwartz (2018), ex-
cept for the endosoma which follows Cassis (2008).

Images were captured using a cellphone Galaxy A52s
5G attached to an Olympus stereomicroscope SZX7 for
dorsal habitus, and with a Canon EOS Rebel T7i with a
Professional Grade Raynox DCR 150 DSLR Objective
Tube Lens and an Objective PLAN Achromatic LWD
infinity SX for external morphology and 10X for dissect-
ed genitalia, mounted on a WeMacro’s automatic focus
stacking rail. Multiple focal planes were taken with Hel-
icon Remote software and merged using Helicon Focus
software. Maps were built using QGIS 3.2, localities
were georeferenced with Google Earth Pro.

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Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

Results

Taxonomy

Thomasoniiris gen. nov.
https://zoobank.org/9FF34EA5-DAB6-426D-87B7-D9BD97C ADECS5

Type species. Thomasomiris setosus sp. nov.

Diagnosis. Head, collar, posterior pronotal lobe, scute-
llum, and hemelytra with short, erect setae; eyes small,
stylate, directed anteriorly (Fig. 1A), less than half head
height (Figs 2A, 3A); vertex with a medial longitudinal
depression; antennal segment I about half the length of
vertex width and almost 4 times as short as antennal seg-
ment II; anterior pronotal lobe depressed in lateral view;
cuneal fracture extending to middle of corium; cuneus
wider than long (Fig. 1A); veins angled at the middle of
membrane, slightly surpassing apex of cuneus; and tibiae
wide, with abundant semierect setae.

Description. Macropterous female. Body length to
apex of membrane 5.6—5.7; body length to apex of cu-
neus 4.64.7.

Surface shiny, pilose; posterior pronotal lobe even-
ly punctate. Head with short and erect setae (Fig. 3A);
antennal segment I with recumbent setae on distal half
(Fig. 3A), antennal segments II-IV with short, recumbent
and longer, semierect setae. Collar, posterior pronotal
lobe, scutellum and hemelytra with short, erect setae.

Structure. Head width more than twice its length.
Clypeus flat in lateral view (Fig. 3A), not visible from
above. Frons flat in lateral view. Eyes small, less than
half head height (Figs 2A, 3A), stylate, directed upwards
and anteriorly, interior and posterior margins straight,
interior margin at same level with lateral collar margins
(Fig. 1A). Maxillary plates rectangular, large, long. La-
bium extending to base of abdomen. Antennal segment I
shortest, about half the length of vertex width, narrower
basally; segment II almost 4 times as long as antennal
segment I; segment III thinner than II (Fig. 1A); segment
IV shorter and thinner than III. Thorax: pronotum tra-
peziform, slightly directed downwards. Anterior pronotal
lobe clearly distinguished from posterior pronotal lobe,
depressed in lateral view (Fig. 3A); collar wider than an-
tennal segment I; calli evident, medium-sized, depressed
and separated from each other by a median depression.
Posterior pronotal lobe flat, with shallow lateral depres-
sions before humeral angles, posterior margin convex
(Fig. 1A). Hemelytron flat in lateral view; lateral margin
convex (Fig. 1A); embolium flat, thin and slightly ex-
panded in posterior margin; medial fracture visible, ad-
jacent to R+M vein; cuneus wider than long (Fig. 1A);
veins angled at middle of membrane and directed poste-
riorly behind posterior margin of cuneus. Legs: femora
flat; metafemora slightly curved; tibiae robust; protibiae
flat on distal interior face.

Male. Unknown.

Geographic distribution. Panama (Fig. 11).

Dtsch. Entomol. Z. 71 (1) 2024, 1-15

Figure 1. Zhomasomiris gen. nov. and Egerocoris gen. nov. new species, dorsal habitus and labels. A. 7’ setosus sp. nov., female holo-
type; B. E. dimorphus sp. nov., male holotype; C. £. ecuatorianus sp. nov., male holotype; D. E. chaparensis sp. nov., male holotype.

Etymology. The name of the new genus 1s formed from
the given name of our friend and colleague Dr Thomas J.
Henry, who has published numerous important papers on
Heteroptera, including many on Miridae, combined with
miris in reference to its assignment to the family Miridae.
Gender masculine.

Discussion. The first couplet of the key to the Neotrop-
ical eccritotarsine genera (Carvalho and Ferreira 1995) re-
fers to the development of male hemelytra; we assumed

that males of Thomasomiris have normal hemelytra, with
clavus and corium distinguished and membrane present,
so we followed to couplet 3. The two monotypic genera
identified in the couplet 2, Aztecariella Carvalho, 1951
and Coleopteromiris Carvalho, 1946 show different char-
acters from Thomasomiris, as a pronotum coarsely punc-
tate and females without division of clavus, coritum and
cuneus, and without membrane in Aztecariella, and sessile
eyes and females with coleopteroid hemelytra in Coleop-

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4

teromiris. Following the key, Thomasomiris runs to the
couplet 8 where the genera Hesperolabops Kirkaldy, 1902
and Aztecarina Calvalho, 1974 are identified and are dis-
criminated by male genitalia characters, but unlike 7iom-
asomiris, Hesperolabops shows strongly pedunculated
eyes, a lobate collar, a longer antennal segment I, and a
cuneus longer than width, and Aztecarina shows a strongly
punctate pronotum with its posterior margin sinuated and
antennal segments I and II of similar length. Four Neo-
tropical genera were omitted in the Carvalho and Ferreira’
key, Perissobasis Reuter, 1892, Eurycipitia Reuter, 1905,
Bugabacoris Carvalho & China, 1959 and Pycnoderiella
Henry, 1993, and five genera were described after 1995,
Cubanomiris Hernandez & Stonedahl, 1996 and Agaveo-
coris, Laterospinocoris, Nigrotomocoris and Schaffnero-
coris, described by Henry and Menard 2020 including
some species with neotropical distribution. The medium
size, the stylate eyes, the labium extending at least to the
metacoxae, and the convex lateral margins of hemelytra
are similar to those found in species of Neoneella Costa
Lima, 1942. However, the short, erect setae, the upward
and anteriorly directed eyes located far from the collar,
the antennal segment IT almost 4 times as long as antennal
segment I, the flattened anterior pronotal lobe clearly sepa-
rated from the evenly punctate posterior pronotal lobe with
the posterior margin convex, the short, wide cuneus, and
the concave posterior margin of cell on membrane forming
an acute angle at the middle, distinguish this new genus
from Neoneella and from all other eccritotarsines.

Thomasomiiris setosus sp. nov.
https://zoobank.org/8F6E7B92-1FB9-4259-8AC5-B99C4B492E56
Figs 1A, 2A, 3A

Diagnosis. Pale yellow with extensive dark areas (Fig. 1A);
clypeus longer than half of head height; vertex more than
two and a half times as wide as eye width; antenna short-
er than length from apex of clypeus to apex of abdomen;
scutellum with basal depression extended longitudinally;
embolium bent upwards from its anterior end to more than
half its length; cuneal fracture long; and cell in membrane
with the interior margin anteriorly concave.

Description. Macropterous female. Coloration. Head
pale yellow (Fig. 1A). Clypeus apex black (Figs 2A,
3A). Labium pale yellow, segments IN-—I'V darker. Eyes
black. Antennal segment I dark brown, base pale yellow;
segment II dark brown, with faint basal ring yellow; seg-
ments III-IV brown. Thorax: collar and calli pale yellow.
Posterior pronotal lobe dark brown, lateral margins pale
yellow. Scutellum pale yellow, apex dark brown. Pleural
area pale yellow, excepting metaepisternum dark brown
dorsally. Hemelytron: embolium pale yellow; clavus
basal half pale yellow and posterior half dark brown; co-
rium anterior and posterior areas pale yellow, and me-
dially dark brown; external apex of cortum and cuneus
orange, cuneus apex dark brown; membrane dark brown
with dark brown veins. Legs: coxae, trochanters, femora

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Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

except apex and basal three quarters of inner face of
protibiae pale yellow; apex of femora, rest of protibiae,
meso- and metatibiae dark brown; tarsi dark brown with
segments I—II paler. Abdomen dark brown.

Surface. Head with short, erect setae, more abundant
on posterior margin and area adjacent to eyes in dorsal
view (Fig. 1A). Labium with semierect setae. Antennal
segments II-IV with abundant recumbent setae and some
longer semierect setae. Collar, posterior pronotal lobe,
scutellum, and hemelytra with abundant, short, erect, yel-
lowish setae, except those on dark brown areas of hem-
elytra darker. Calli glabrous (Fig. 3A). Veins with very
short erect setae. Pleura and abdomen with longer and
more dispersed erect setae than dorsum. Coxae, trochan-
ters, femora and tarsi with semierect setae. Tibiae with
abundant dark recumbent and semierect setae, thicker
than the ones in the femora.

Structure. Head: vertex almost twice length of anten-
nal segment I; upper margin at eye level (Fig. 2A), with
median longitudinal depression. Labrum not extending
half way on labial segment I (Fig. 3A). Gular area not
visible. Antennal segment II thinner and between 3-4
times as long as antennal segment I. Antennal segment
IH almost twice the length of antennal segment I. Tho-
rax: anterior collar margin concave and posterior margin
convex, posterior sulcus shallow medially (Fig. 1A). Pos-
terior pronotal margin width more than twice its length;
anterior pronotal width less than half posterior pronotal
margin width. Calli not attaining lateral pronotal margins.
Scutellum with triangular basal depression extended lon-
gitudinally. Hemelytron: embolium bent upwards from its
anterior end to more than a half its length. Medial fracture
shorter than one quarter of corium length. Claval commis-
sure longer than scutellum and half of pronotum length to-
gether. Cuneal fracture evident, straight, attaining half of
corium width. Interior margin of cuneus concave. Mem-
brane with interior margin concave anteriorly, and posteri-
or margin straight with middle weakly concave (Fig. 1A).

Measurements (n: 2): Body length to apex of mem-
brane (5.60), 5.70; body length to apex of cuneus 4.60,
(4.70). Head: width 1.28; interocular distance 0.73,
(0.74), and (1.38), 1.42 times head height. Labium: seg-
ment I length 0.80, (0.86); II, 0.91, (0.96); III, (0.32),
0.34; IV, 0.27, (0.29). Antenna: segment I length (0.40),
0.41; If, 1.55; III, (0.75), 0.80; IV, (0.62), 0.66. Pronotum:
length (0.92), 0.95; anterior width 0.95; posterior width
1.92, (1.97). Scutellum: length 0.72, (0.74); width 0.85.
Cuneus: length 1.00; anterior width 1.04. Membrane cell
(2.06), 2.16 times longer than wide.

Geographic distribution. Panama (Fig. 11).

Type material. Holotype 2: Panama: Barro / Collora-
do] Isl., 17 Mar. / 1985, J. E. Loye coll. [9°9'N, 79°51'W]
(USNM). Paratype 2°: Panama: Barro / Col[orado] Isl.,
17 Mar. / 1985, J. E. Loye coll. // Neoneella / sp. / det. T.
J. Henry 1987 (MLP).

Etymology. The specific epithet is the Latin adjec-
tive setosus, -a, -um, meaning ‘bristly’, in allusion to the
abundant short, erect setae of the dorsal surface.

Dtsch. Entomol. Z. 71 (1) 2024, 1-15

Figure 2. Zhomasomiris gen. nov. and Egerocoris gen. nov. new species, head frontal view. A. 7: sefosus sp. nov., female holotype;

B. E. dimorphus sp. nov., male holotype; C. £. ecuatorianus sp. nov., male holotype; D. F. chaparensis sp. nov., male holotype.

Egerocoris gen. nov.
https://zoobank. org/10DAE85 1-82DB-4BD4-9E5 1-2DEEFD77F055

Type species. Egerocoris ecuatorianus sp. nov.

Included species. Egerocoris chaparensis sp. nov.,
E. dimorphus sp. nov., E. ecuatorianus sp. nov.

Diagnosis. Eyes large, stylate (Fig. 1B—D), partly
covering collar in lateral view (Fig. 3B—D); labrum
short, less than half labial segment I length; antennal
segment I slightly narrowing on distal half and wider
than antennal segment II; collar wider than antennal
segment I; embolium flat, bent upwards from its
anterior end to less than a half its length; dorsal wall
of genital capsule sclerotized, left wall shorter than
right wall (Fig. 6); left paramere body triangular, with
apical process initially curved and V-shaped in frontal
view (Fig. 8); and, ductus seminis basally expanded
(Fig. 10A, B).

Description. Macropterous male. Body length to
apex of membrane 4.42-4.66; body length to apex of cu-
neus 3.95—4.66; body width 1.94—2.43.

Surface shiny; posterior pronotal lobe evenly punc-
tate. Head, collar, pronotum and hemelytra with abundant
short, recumbent setae.

Structure. Head width more than twice length. Cly-
peus rounded in lateral view, not visible from above. Frons
rounded in lateral view (Fig. 3B—D). Vertex flat (Fig. 2B—
D), wider than head length. Eyes large, more than half head
height, stylate, interior margin straight and at lateral collar
margins level (Fig. 1B—D). Gular area not visible. Labium
extending at least to metacoxae; segment I concave ven-
trally. Antennal segment I slightly narrowing on distal half.
Antennal segment II straight, slender. Antennal segment III
narrowing towards the apex, more slender than segment II.
Antennal segment IV straight. Thorax: collar wider than
antennal segment I; anterior margin concave and posteri-
or margin straight. Pronotum slightly directed downwards.
Calli evident, small to large, separated by a median de-
pression, sometimes attaining to pronotal lateral margins.
Posterior pronotal lobe with shallow lateral depressions be-
fore humeral angles; posterior margin straight (Fig. 1 B—D).
Mesoscutum exposed (Fig. 1B). Scutellum with anterior

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Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

Figure 3. Thomasomiris gen. nov. and Egerocoris gen. nov. new species, head and pronotum lateral view. A. 7’ sefosus sp. nov.,
female holotype; B. £. dimorphus sp. nov., male holotype; C. -. ecuatorianus sp. nov., male holotype; D. £. chaparensis sp. nov.,

male holotype.

depression. Hemelytron flat in lateral view; lateral margin
convex (Fig. 1B—D). Embolium flat, straight to slightly ex-
panded on posterior margin, bent upwards. Medial fracture
visible, shorter than half the corium length. Cuneus longer
than wide (in one species the length sexually dimorphic
as in Fig. 4). Veins angled in middle of membrane before

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posterior margin of cuneus; cell with interior and posterior
margins straight (Fig. 4). Legs: profemora broader at base.
Metafemora slightly curved. Protibiae wider than meso- and
metatibiae. Male genitalia: genital capsule length variable
relative to abdomen length, longer than wide. Right wall
more developed than left wall. Dorsal wall in lateral view

Dtsch. Entomol. Z. 71 (1) 2024, 1-15

straight, well developed laterally, with sclerotized process-
es (Fig. 5B, D, F); posterior margin concave. Ventral wall
in lateral view oblique, more developed than the dorsal wall
(Fig. 6); posterior margin sinuate. Subgenital plate with two
processes, one on the left side and one on the right side
(Fig. 7); each one embedded with corresponding paramere.
Genital opening broad, dorsally directed. Left paramere
(Fig. 8) smaller or larger than right paramere; body broad,
triangular; apical process well developed, basally curved;
expanded or not at apex. Right paramere (Fig. 9) simpler
than left paramere; uniformly wide; slightly to strongly
curved. Aedeagus (Fig. 10) small and simple; phallotheca
membranous on basal half, sclerotized on distal half. Duc-
tus seminis basally expanded, membranous and not folded:
sclerotized behind last curvature; apex extending to middle
of phallotheca; endosoma membranous to sclerotized.

Macropterous female. Body length to apex of mem-
brane 4.28-4.94:; body length to apex of cuneus 3.7-4.51;
body width 1.97—2.09. Lateral margins of hemelytra e1-
ther convex or (in species with sexual dimorphism) paral-
lel with less well developed cuneus.

Geographic distribution. Bolivia, Ecuador (Fig. 11).

Etymology. The name of the genus is formed from the
family name of Joe Eger, who has extensively collected
Heteroptera, including one of the specimens studied here-
in, and published several papers on the group, combined
with the latinized Greek noun coris, meaning “true bug”.
Gender masculine.

Discussion. Egerocoris resembles Nee//a in general
appearance, but in Nee/la the eyes are stylated but not
covering the lateral margins of collar, the postocular re-
gion of head present a patch of setae, the antennal seg-
ment I 1s straight and as wide as II, the subgenital plate
lacks processes and the parameres are simpler.

Egerocoris runs to the couplet 8 in the key to the Neo-
tropical eccritotarsine genera (Carvalho and Ferreira
1995) where the genera Hesperolabops Kirkaldy, 1902
and Aztecarina Calvalho, 1974 are identified. These gen-

Key to species of Egerocoris

era are quite different from Egerocoris, since Hespero-
labops shows strongly pedunculated eyes, a lobate collar,
and a curved spiniform projection in the genital capsule,
and Aztecarina shows a strongly punctate pronotum with
its posterior margin sinuated and antennal segments I
and II of similar length. The character states of the large,
stylate eyes that partly cover the collar in lateral view,
the short labrum, the long labium extending at least to
metacoxae, the slightly narrowing antennal segment I on
the distal half and wider than antennal segment II, the
collar being wider than antennal segment I, the posterior
pronotal lobe evenly punctate with its posterior margin
straight, and the embolium being flat, bent upwards from
its anterior end to less than a half its length, combined
with characters of the male genitalia, including the pres-
ence of sclerotizations in the dorsal wall of the genital
capsule, the processes on the subgenital plate embedding
the parameres, and the triangular left paramere with a
well-developed apical process, allow the recognition of
this genus from the four genera omitted in Carvalho and
Ferreira’s key and the five genera described afterwards
detailed in the Thomasomiris discussion above.

Egerocoris dimorphus sp. nov. shows sexual dimorphism
in the shape and length of cuneus similar to Proneella Car-
valho, 1960, but the hemelytra are slightly more convex,
and the cuneus is wider relative to its length. This species
runs to the couplet 13 in the Carvalho and Ferreira’s key
(1995) where Proneella and Neoneella Costa Lima, 1942
are identified. According to Carvalho (1960), the sexual
dimorphism relates Proneella with Neoneella, and in both
genera the included species also share several characters
from internal and external morphology. Among the three
species of Egerocoris two of them lack sexual dimorphism,
but share several characters from the male genitalia and
from the external morphology allowing us to infer a close
relationship and justifying the inclusion of these species in
the same genus, besides the absence of sexual dimorphism
in E. ecuatorianus sp. nov. and E. chaparensis sp. nov.

il Antennal segment | unicolorous (Fig. 3D). Clypeus length less than or equal to half head height. Labial segment | deeply
concave ventrally. Internal margins of cell separated with hemelytra in repose (Fig. 1D). Right paramere larger than left,
C-shaped in dorsal view (Fig. 9F). Right process of subgenital plate ending in numerous asymmetric teeth (Fig. 7E, F)

irk. Bone ee eR Ae haat Ae ak ae Bel E. chaparensis sp. nov.

- Antennal segment | bicolored, darker distally (Fig. 3B, C). Clypeus length greater than half head height. Labial segment
| slightly concave ventrally. Internal margins of cell overlapping in anterior half with hemelytra in repose (Fig. 1B). Right
paramere smaller than left, “V’—shaped in dorsal view (Fig. 9B, D). Right process of subgenital plate ending in simple

poitted-apex-GFig. FAD). 6... Ge yeclllas ce en averelesee.

2 Antennal segment III yellow (Fig. 1B). Labial segment | shorter than vertex width. Embolium longer than abdomen. Cla-
val commissure shorter than twice scutellum length. Internal margin and length of cuneus sexually dimorphic (males
with concave internal margin and cuneus extending to posterior margin of membrane, without contacting each other
with hemelytra in repose; females with cuneus short, not extending to posterior margin of membrane) (Fig. 4). Female
subgenital plate equal to or shorter than anterior Margin WITH ......... ccc ccc cec cence cece eee eneeeeneeeeneeeeeenes E dimorphus sp. nov.

- Antennal segment III basally yellow and distally darker (Fig. 1C). Labial segment | longer than vertex width. Embolium
shorter than abdomen. Claval commissure longer than twice scutellum length. Internal margin and length of cuneus not
sexually dimorphic, not extending to posterior margin of membrane on both sexes. Female subgenital plate longer than

AM TSMOls Pilato Win WHE LIN, hie ottsaaccaecteenrntecetgetutes ode dence bee +

Leanosaaadecmtinerde Od! pm aeroem ener, aetna F. ecuatorianus Sp. nov.

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8

Egerocoris dimorphus sp. nov.
https://zoobank.org/B438 1 DE7-5D26-4F 1 8-A037-F2B845E097F5
Figs 1B, 2B, 3B, 4, 5A, B, 6A, B, 7A, B, 8A, B, 9A, B, 10A

Diagnosis. Antennal segment I bicolored, darker distally
(Figs 1B, 3B), segment III whitish; femora yellow and
irregularly tinged with orange; level of vertex not attain-
ing dorsal margin of eyes; labium extending to metacox-
ae and segment I slightly concave ventrally; calli large,
not attaining pronotal lateral margins; embolium longer
than abdomen; internal margin of cell straight; sexu-
al dimorphism on internal margin and length of cuneus
(Fig. 4A, B), which is concave in males and reaches pos-
terior margin of membrane without contacting each other
with hemelytra in repose; cell 3 times as long as wide;
genital capsule with a dorsal sclerotized tooth in right
side (Figs 5B, 6A); left process of subgenital plate small-
er than the right, sclerotized and asymmetrically forked
(Fig. 7A, B); left paramere larger than right paramere.

Description. Macropterous male. Coloration. Head
yellow; labium yellow; eyes black. Antennal segment I
yellow, darker distally; II dark brown; III pale yellow;
IV pale yellow. Thorax: collar, pronotum, scutellum
and pleural area yellow. Hemelytron yellow, irregularly
tinged with orange; membrane yellowish; veins yellow.
Legs yellow, femora and tibiae tinged with orange, tarsi
darker distally, claws brown. Abdomen yellow.

Surface. Antennal segment I with recumbent setae and
a few erect setae (Fig. 3B); segment IT with semierect setae.
Pleura and abdomen with longer, more dispersed setae than
dorsum. Coxae, trochanters, femora and tarsi with semi-
erect setae; tibiae with abundant, short, recumbent setae.

Structure. Head: level of vertex not attaining dorsal
margin of eyes (Fig. 2B). Maxillary plates large. Labium
extending to metacoxae and slightly concave ventrally.
Antennal segment II more than 2.5 times as long as anten-
nal segment I. Thorax: posterior pronotal margin 2 times

0.5mm

Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

as wide as pronotum length. Calli large not attaining lat-
eral pronotal margins. Pronotal width across calli half as
wide as posterior margin width. Hemelytron: embolium
straight, bent upwards from its anterior end to less than
a half its length. Claval commissure longer than scute-
llum length and half of pronotal length together. Cune-
us extending to posterior membrane margin, apex not
contacting each other with hemelytra in repose (Fig. 4),
internal margin concave. Male genitalia: Genital capsule
more than one third abdomen length. Dorsal wall on right
apex with one sclerotized tooth, tapering towards apex;
left apex blunt and sclerotized. Left process of subgenital
plate directed dorsally as a sclerotized wall, dorsally ex-
panded, and divided into two short branches, the internal
blunt and the external pointed and extended outside the
genital capsule (Figs 5A, B, 6A, 7A, B). Right process
of subgenital plate longer than left process, tapering to-
wards apex, curved (Fig. 7A, B), with a basal, internal
expansion. Left paramere (Fig. 8A, B) larger than the
right paramere, with a sclerotized and pointed dorsal pro-
jection; apical process with tiny teeth on dorsal margin,
dorsally expanded at apex. Right paramere (Fig. 9A, B)
with body wider than basal and apical processes, curved;
apical process blunt, with a sclerotized, pointed tooth.
Aedeagus (Fig. 10A) phallotheca tapering towards apex;
endosoma membranous.

Measurements (n: 2): Body length to apex of mem-
brane 4.42, (4.66); body length to apex of cuneus 4.42,
(4.66); body width 2.23, (2.43). Head: width 1.19, (1.31);
interocular distance 0.53, (0.56), 1.20 (1.27) times as wide
as head length. Labium: segment I length 0.43, (0.52); I,
0.52, (0.60); III, 0.20; IV, 0.20. Antenna: segment I length
0.48, (0.56); II, 1.36, (1.49); ILL, 0.78, (0.85); TV, 0.84, (ab-
sent). Pronotum: length 0.74, (0.82); anterior width (0.86),
0.88; posterior width 1.49, (1.58). Scutellum: length 0.53,
(0.60); width 0.77. Cuneus: length 1.58, (1.70); anterior
width 0.90, (0.95). Cell 3.00 times as long as wide.

B 0.5mm

Figure 4. /-gerocoris dimorphus sp. nov., dorsal view of cuneus. A. Male holotype; B. Female paratype.

dez.pensoft.net

Dtsch. Entomol. Z. 71 (1) 2024, 1-15

Macropterous female. Similar to male in size, color-
ation, surface, and structure. Lateral margins of hemely-
tra parallel; cuneus less developed, without extending to
posterior membrane margin, 1.5 times as long as anterior
margin width, and internal margin concave with apex trun-
cate; posterior veins margin straight. Body length to apex
of membrane 4.94; body length to apex of cuneus 4.51;
body width 2.02. Head: width 1.23; interocular distance
0.58. Labium: segment I length 0.46; II, 0.62; III, 0.19;
IV, 0.2. Antenna: segment I length 0.52; II, 1.36; II, 0.74;
IV, 0.79. Pronotum: length 0.82; anterior width 0.89; pos-
terior width 1.66. Scutellum: length not measured; width
0.76. Cuneus: length 1.28; anterior width 0.84.

Geographic distribution. Ecuador (Fig. 11).

Etymology. The specific epithet is from the Greek di
(two), and morphe (form), referring to the sexual dimor-
phism in cuneal development.

Type material. Holotype @: Ecuador: Prov. Napo
[Orellana] / vic. Puerto Misahuali, / 1650—1900 ft, 6-19-—
IX— / 1998, J. E. Eger, coll. // 1°2'4.2"S, 77°39'49.2"W/
Mercury vapor & / Ultraviolet lights (USNM). Paratypes:
3, Ecuador: Napo [Orellana] Res. Ethnica / Waorani, 1
km S. Onkone Gare / Camp, Trans. Ent. 21 June 1994
/ 220m 00°39'10"S, 076°26'00"W / T. L. Erwin, et al. //
Insecticidal fogging of mostly bare / green leaves, some
with covering / of lichenous or bryophytic plants in / terre
firme forest At 8 x-trans, / 94 m mark Project MAXUS
Lot 708 (MLP); 9, Ecuador: Napo [Orellana], Tiputini /
Biodiversity Station, 216 m, / 00°37'55"S, 76°08'39"W, /

B =i fe

~~ LPSp

2

9 February 1999 / T.L Erwin et al., collectors // Insecti-
cidal fogging of mostly / bare green leaves, some with /
covering of lichenous or / bryophytic plants; / Lot #2002,
Transect #T—1 (USNM).

Remarks. This species is sexually dimorphic in the
shape and length of cuneus (Fig. 4), and in this respect
it is similar to species of Proneella, which have the lat-
eral margins of the hemelytra slightly convex and the cu-
neus narrower relative to its length. Also, in species of
Proneella the eyes are sessile, the level of vertex attains
the dorsal margin of the eyes, the antennal segment I is
straight, the antennal segment II is widest in the central or
distal area, the calli are small, and the posterior margin of
the cell is straight.

Egerocoris ecuatorianus sp. nov.
https://zoobank. org/EB3EE0D7-40C 1 -4FFO-BAB9-ECE165ED5521
Figs 1C, 2C, 3C, 5C, D, 6C, D, 7C, D, 8C, D, 9C, D, 10B

Diagnosis. Antennal segment I bicolored, lighter basal-
ly (Figs 2C, 3C), and segment III with basal half pale
yellow, distal half darker; femora yellow and irregularly
tinged with orange ; level of vertex not attaining the dor-
sal margin of eyes (Fig. 2C); labium extending to abdom-
inal segment III and segment I slightly concave ventrally;
calli large, attaining lateral pronotal margins; embolium
shorter than abdomen; internal margin of cell straight; in-
ternal margin and length of cuneus not sexually dimorphic

DT
A RPSP

. a «LPSP
0.2mm F a

0.2mm

Figure 5. E-gerocoris gen. nov. new species, left lateral view of genital capsule. A, B. £. dimorphus sp. nov.; C, D. E. ecuatorianus

sp. nov.; E, K. £. chaparensis sp. nov.; A, C, E. Genital capsule not dissected; B, D, F. Genital capsule dissected; DT—dorsal tooth;

LPSP-left process of subgenital plate; RPSP-right process of subgenital plate.

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10 Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

A = rm.

B ram LP

DT
0.2mm E 0.2mm
RP
7 RPSP
0.2mm F 0.2mm

Figure 6. /-gerocoris gen. nov. new species, dorsal and ventral views of genital capsule. A, B. E. dimorphus sp. nov.; C, D. E. ecu-

atorianus sp. nov.; E, F. £. chaparensis sp. nov.; A, C, E. Genital capsule dorsal view; B, D, F. Genital capsule ventral view; DT-

dorsal tooth; RP-right paramere; LP—left paramere; LPSP-left process of subgenital plate; RPSP-—right process of subgenital plate.

(Fig. 1C); genital capsule with a sclerotized broad tooth
on left apex of posterior margin (Figs 5D, 6C); left pro-
cess of the subgenital plate sclerotized, pointed, curved
(Fig. 7C); left paramere same size as right paramere; and
female subgenital plate longer than anterior margin width.

Description. Macropterous male. Coloration. Head
yellow. Clypeus apex reddish. Labrum reddish. Labium
yellow, segments II—IV pale yellow. Eyes silver. Antennal
segment I brown, base yellow; II, brown; III, basal half
pale yellow, distal half darker; IV, brown. Thorax: collar
yellow. Pronotum orange. Mesoscutum yellow. Scutel-
lum yellow, apex orange. Pleural area yellow. Hemely-
tron orange. Membrane and veins yellow. Legs: coxae
and trochanters pale yellow. Femora and tibiae yellow
and irregularly tinged with oranges. Tarsi pale yellow.
Claws brown. Abdomen yellow.

Surface. Antennal segment I with recumbent setae and
a few erect setae (Fig. 3C). Antennal segments I-III with
semierect setae and some erect setae. Antennal segment
IV with semierect setae. Pleural area and abdomen with
longer and more dispersed setae than dorsum. Coxae, tro-
chanters, femora, and tarsi with semierect setae. Tibiae
with abundant recumbent setae.

Structure. Head: level of vertex not attaining dorsal
margin of eyes (Fig. 2C). Maxillary plates medium-sized,
wide. Labium extending to abdominal segment III, seg-
ment I extending to procoxae and slightly concave ven-
trally. Antennal segment II more than 2 times as long as

dez.pensoft.net

antennal segment I. Thorax: posterior pronotal margin 2
times as wide as pronotum length. Calli attaining lateral
pronotal margins. Pronotal width across calli more than
half of posterior margin width. Hemelytron: embolium
slightly wider at posterior margin, bent upwards from its
anterior end to less than a half its length. Internal mar-
gin of cuneus concave (Fig. 1C). Male genitalia: genital
capsule less than one quarter abdomen length, almost 2
times as wide as long. Right wall straight. Left wall con-
vex on anterior half and concave on posterior half. Dorsal
wall at left apex with evident concavity, adjacent mar-
gin sclerotized and bent upwards as a broad tooth (Figs
5C, D, 6C). Left process of subgenital plate smaller than
right process, flat and broad internally, curved at point-
ed apex (Fig. 7D). Right process of subgenital plate lon-
ger than left process, broad basally and tapered towards
apex, curved, with a left basal expansion (Fig. 7C, D).
Left paramere (Fig. 8C, D) almost the same size as right
paramere, with a sclerotized and pointed dorsal projec-
tion; ventral margin concave; apical process tapering to-
wards the sclerotized apex. Right paramere (Fig. 9C, D)
with body wider than basal and apical processes, curved;
apical process narrower apically, curved, apex rounded,
flat and bent, appearing as an internal, flat, broad tooth.
Aedeagus (Fig. 10B) phallotheca sclerotized on distal
half dorsally and at middle as a stripe. Ductus seminis
sclerotized at the same level as phallotheca dorsally; en-
dosoma membranous, base sclerotized.

Dtsch. Entomol. Z. 71 (1) 2024, 1-15 11

0.2mm 0.2mm

J
0.2mm D 0.2mm F 0.2mm 7

Figure 7. Egerocoris gen. nov. new species, subgenital plate. A, B. E. dimorphus sp. nov.; C, D. FE. ecuatorianus sp. nov.; E,
F. E. chaparensis sp. nov.; A. Subgenital plate right posterolateral view; C, E. Subgenital plate right lateral view; B, D, F. Subgenital
plate posterior view; DT—dorsal tooth; LPSP-left process of subgenital plate; RPSP-right process of subgenital plate.

A 0.15mm C 0.15mm E 0.15mm

B 0.15mm D 0.15mm F 0.15mm

Figure 8. Egerocoris gen. nov. new species, left paramere. A, B. . dimorphus sp. nov.; C, D. E. ecuatorianus sp. nov.; E, F. E. chapa-
rensis sp. nov.; A, C, E. Interior view; B, D, F. Posterior view.

dez.pensoft.net

12

A =a) Ae

B “in|

Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

0.15mm E

of
0.15mm F 0.15mm

Figure 9. Egerocoris gen. nov. new species, right paramere. A, B. £. dimorphus sp. nov.; C, D. E. ecuatorianus sp. nov.;

E, F. E. chaparensis sp. nov.; A, C, E. Interior view; B, D, F. Dorsal view.

Measurenments: Body length to apex of membrane
4.61; body length to apex of cuneus 4.3; body width 2.18.
Head: width 1.13; interocular distance 0.54, 1.17 times as
wide as head length and 1.08 times as wide as antennal
segment length. Labium: segment I length 0.56; II, 0.66;
IIT, 0.23; IV, 0.22. Antenna: segment I length 0.5; I, 1.26;
II, 0.77; IV, 0.67. Pronotum: length 0.71; anterior width
0.80; posterior width 1.42. Scutellum: length 0.48; width
0.72. Claval commissure more than 2 times as long as
scutellum. Cuneus: length 1.18; anterior width 0.67. Cell
2.55 times as long as wide.

Macropterous female. Similar to males in size, col-
oration, surface, and structure, with medial fracture
longer than half of corium. Body length to apex of
membrane 4.84; body length to apex of cuneus 4.37;
body width 2.02. Head: width 1.16; interocular dis-
tance 0.54; vertex 1.22 times as wide as head length
and 1.14 times as wide as antennal segment I length.
Labium: segment I length 0.58; II, 0.68; HI, 0.22; IV,
0.22. Antenna: segment I length 0.50; H, 1.18; III,
0.80; IV, 0.78. Pronotum: length 0.74; anterior width
0.84; posterior width 1.50. Scutellum: length 0.47;
width 0.73. Cuneus: length 1.03; anterior width 0.68.
Cell 2.63 times as long as wide.

Geographic distribution. Ecuador (Fig. 11).

Etymology. The specific epithet is a latinized adjective
created from the Spanish adjective ecuatoriano, -a, mean-
ing “related to the Ecuador’, in reference to the country
Ecuador, where the specimens were collected.

dez.pensoft.net

Type material. Holotype 3: Ecuador: Napo [Orella-
na]: Tiputini / Biodiversity Station, 216 m, / 0°37'55"S,
76°08'39"W, / 7 February 2002, / T. J. Henry & P. S. F.
Ferreira (USNM). Paratype 9°: Ecuador: Napo [Orella-
na]: Tiputini / Biodiversity Sta., Tower 1, / 0°37'55"S,
76°08'39"W, / 216 m, 3 February 2002, / T. J. Henry & P.
S. F. Ferreira / Ex. Mercury vapor light (USNM).

Egerocoris chaparensis sp. nov.
https://zoobank.org/7664DC A6-2FCC-4A6C-9C82-8BD4BD8C999B
Figs 1D, 2D, 3D, 5E, F, 6E, F, 7E, F, 8E, F, 9E, F, 10C

Diagnosis. Antennal segment I yellow (Fig. 3D), segment
Il with basal half pale yellow, distal half darker; femora
yellow; level of vertex attaining dorsal margin of eyes (Fig.
2D); labium extending at least to metacoxae and segment
I deeply concave ventrally and narrowing distally; calli
small, not attaining lateral pronotal margins; embolium
longer than abdomen; internal margin of cell convex; with-
out sexual dimorphism on internal margin and length of
cuneus (Fig. 1D); dorsal wall of genital capsule well devel-
oped, with two tiny medial and sclerotized teeth (Figs SF,
7E, F); left process of subgenital plate C-shaped in dorsal
view, divided into two broad branches; right process end-
ing in many asymmetrical teeth (Fig. 7E, F); left paramere
smaller than the right paramere; right paramere curved,
with apex expanded dorsoventrally (Fig. 9E); and aedea-
gus less than or as long as the genital capsule.

Dtsch. Entomol. Z. 71 (1) 2024, 1-15

0.15mm B

13

BE

0.15mm C

0.15mm

Figure 10. Egerocoris gen. nov. new species, aedeagus right lateral view. A. FE. dimorphus sp. nov.; B. E. ecuatorianus sp. nov.;

C. E. chaparensis sp. nov.; Ae-—aedeagus; BE—basal expansion.

Figure 11. Distributional map of the species included in the genera 7homasomiris gen. nov. and Egerocoris gen. nov. Blue dot rep-

resents occurrence of 7homasomiris setosus sp. nov.; yellow dots represent occurrences of Egerocoris dimorphus sp. nov.; orange
dot represents ocurrence of E. ecuatorianus sp. nov.; in grey the province Chapare (Bolivia) where /. chaparensis sp. nov. occurs.

Description. Macropterous male. Coloration. Head
yellow. Labium yellow. Eyes black with silver spots. An-
tennal segment I yellow; II, dark brown; III, basal half
whitish, more than distal half brown. Thorax: collar, pro-
notum, scutellum and pleural area yellow. Hemelytron,
legs and abdomen yellow, tarsi whitish.

Surface. Antennal segment J-II with recumbent se-
tae, and a few semierect setae, more abundant on seg-
ment II. Antennal segments HI with semierect setae and
a few erect setae. Pleural area and abdomen with longer

and more disperse setae than dorsum. Coxae, trochanters,
femora, and tarsi with semierect setae. Tibiae with abun-
dant recumbent setae.

Structure. Head: level of vertex attaining dorsal mar-
gin of eyes (Fig. 2D). Maxillary plates long and thin.
Labium just beyond metacoxae; segment I, deeply con-
cave ventrally and narrowing distally; I, thin and long;
IH-IV, wider and short. Antennal segment II more than
2 times as long as antennal segment I. Thorax: posteri-
or pronotal margin 2 times as wide as pronotum length.

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14

Calli small, not attaining lateral pronotal margins. Pro-
notal width across calli more than half of posterior mar-
gin width. Hemelytron: embolium straight, bent upwards
from its anterior end to more than anterior third; longer
than abdomen. Claval commissure longer than scutellum
and half pronotum length combined. Internal margin of
cuneus concave (Fig. 1D). Male genitalia: genital capsule
more than one third abdomen length. Dorsal wall with
two tiny, medial, sclerotized teeth directed upwards; the
left tooth larger than the right (Figs SE, F, 7E, F). Subgen-
ital plate with two large processes at the level of paramere
insertions. Left process, shorter than right process, flat,
broad basally and forked. Right process longer, thin-
ner, extremely broad basally with small external tooth,
C-shaped, apex with several asymmetrical teeth (Fig.
7E, F). Left paramere (Fig. 8E, F) smaller and broader
than right paramere, with a blunt dorsal projection, api-
cal process with an internal, triangular, tooth, apex flat,
thin and sclerotized. Right paramere (Fig. 9E, F) curved;
body wide basally, narrowed distally; apical process ex-
panded dorsoventrally, dorsal expansion blunt, with ven-
tral expansion pointed. Aedeagus (Fig. 10C) phallotheca
sclerotized on left side, with a dorsal stripe and a ventral
oblique stripe sclerotized; endosoma sclerotized.

Measurements: Body length to apex of membrane
4.46; body length to apex of cuneus 3.95; body width
1.94. Head: width 1.08; interocular distance 0.56, 1.22
times as wide as head length and 1.47 times as wide as an-
tennal segment I length. Labium: segment I length 0.52;
II, 0.62; III, 0.17; IV, 0.18. Antenna: segment I length
0.38; II, 0.91; III, not measured; IV, absent. Pronotum:
length 0.71; anterior width 0.80; posterior width 1.46.
Scutellum: length 0.50; width 0.74. Cuneus: length 0.98;
anterior width 0.76. Cell 2.78 times as long as wide.

Macropterous female. Similar to males in size, color-
ation, surface, and structure. Antennal segment II basally
yellow, distally dark brown; IV, brown.

Structure. Antennal segment II more than 2.5 times
as long as antennal segment I. Thorax: posterior pronotal
margin more than twice as wide as pronotum length. Pro-
notal width across calli less than half of posterior margin
width. Hemelytron: embolium bent upwards from its an-
terior end to a half or more than a half its length. Medial
fracture attaining half corium length. Claval commissure
equal to or longer than scutellum and half pronotal length
combined. Cuneus not bent downwards.

Measurements (n: 4): Body length to apex of mem-
brane 4.28-4.79; body length to apex of cuneus 3.7-4.23;
body width 1.97—2.09. Head: width 1.07—1.10; interocu-
lar distance 0.54—0.59, 1.20—1.34 times as wide as head
length and 1.50—1.64 times as wide as antennal segment
I length. Labium: segment I length 0.48—0.54; II, 0.49-
0.65; III, 0.18—0.19; IV, 0.18. Antenna: segment I length
0.36—-0.37; II, 0.92-1.0; III, 0.44—0.46; IV, 0.58—0.62.
Pronotum: length 0.74—0.80; anterior width 0.77—0.82;
posterior width 1.54—1.58. Scutellum: length 0.49-0.55;
width 0.72—-0.77. Cuneus: length 0.98—-1.09; anterior
width 0.74—0.78. Cell 2.34—2.8 times as long as wide.

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Eugenia Minghetti et al.: New genera and species of Neotropical Bryocorinae

Geographic distribution. Bolivia (Fig. 11).

Etymology. The specific epithet refers to Chapare Prov-
ince, Bolivia, where the specimens were collected. Adjective.

Type material. Holotype 3: Sajta, XI-93 / Chapare,
Bolivia (USNM). Paratypes: 42, same data as the holo-
type (2 USNM, 2 MLP).

Acknowledgements

We thank Thomas Henry (Systematic Entomology Labo-
ratory, ARS, USDA, (U.S. National Museum of Natural
History (USNM), Washington, DC) for kindly reviewing
the manuscript and his hospitality and support during our
visit to the USNM collection, which was also made pos-
sible thanks to a Smithsonian Short-Term Visitor Grant.
This work was funded by the Consejo Nacional de Inves-
tigaciones Cientificas y Técnicas (CONICET).

References

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Henry TJ (1993) A striking new genus and species of bryocorine
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with new and revised synonymies, lectotype designations, and de-
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