Table Of ContentZOOSYSTEMATICA ROSSICA, 21(2): 204–233 25 DECEMBER 2012
Review of the Neotropical genus Paraphidnia (Orthoptera:
Tettigoniidae: Phaneropterinae)
Обзор неотропического рода Paraphidnia (Orthoptera:
Tettigoniidae: Phaneropterinae)
OSCAR J. CADENA-CASTAÑEDA & A.V. GOROCHOV*
ОСКАР Х. КАДЕНА-КАСТАНЕДА, А.В. ГОРОХОВ
O.J. Cadena-Castañeda, Universidad Distrital Francisco José de Caldas, Grupo de Investigación en Artrópodos
“Kumangui”, Bogotá, Colombia. E-mail: [email protected]
A.V. Gorochov, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034,
Russia. E-mail: [email protected]. *Corresponding author.
The genus Paraphidnia Giglio-Tos, 1898 belonging to the tribe Dysoniini is divided into two
subgenera: Paraphidnia s. str. and Anaphidna subgen. nov. The nominotypical subgenus con-
tains only P. gallina Giglio-Tos, 1898. Paraphidnia verrucosa (Brunner von Wattenwyl, 1878)
and P. lankesteri Rehn, 1918 are included in the new subgenus together with twelve new
species: P. mexicana sp. nov. (type species of Anaphidna), P. peruana sp. nov., P. svetlanae
sp. nov., P. bezverkhovi sp. nov., P. polestshuki sp. nov, P. gracielae sp. nov., P. quirozi sp. nov.,
P. silvai sp. nov., P. osae sp. nov., P. hernandezi sp. nov., P. rubricorpus sp. nov., and P. rhinoceros
sp. nov. The new subgenus and all the new species are described and illustrated.
Род Paraphidnia Giglio-Tos, 1898, относящийся к трибе Dysoniini, подразделен на два
подрода: Paraphidnia s. str. и Anaphidna subgen. nov. Номинативный подрод содержит
только P. gallina Giglio-Tos, 1898. Paraphidnia verrucosa (Brunner von Wattenwyl, 1878) и
P. lankesteri Rehn, 1918 включены в новый подрод вместе с двенадцатью новыми видами:
P. mexicana sp. nov. (типовой вид Anaphidna), P. peruana sp. nov., P. svetlanae sp. nov., P.
bezverkhovi sp. nov., P. polestshuki sp. nov, P. gracielae sp. nov., P. quirozi sp. nov., P. silvai
sp. nov., P. osae sp. nov., P. hernandezi sp. nov., P. rubricorpus sp. nov. и P. rhinoceros sp. nov.
Новый подрод и все новые виды описаны и проиллюстрированы.
Key words: katydids, taxonomy, Neotropics, Orthoptera, Tettigoniidae, Phaneropterinae,
Dysoniini, Paraphidnia, new taxa
Ключевые слова: кузнечики, таксономия, Неотропики, Orthoptera, Tettigoniidae, Pha-
neropterinae, Dysoniini, Paraphidnia, новые таксоны
INTRODUCTION lankesteri Rehn, 1918 from Costa Rica; P.
verrucosa (Brunner-Wattenwyl, 1878) from
This paper is a continuation of the review
Brazil. The taxonomic history of Paraphid-
of the Neotropical tribe Dysoniini started
nia began after the description of Aphid-
by the first coauthor (Cadena-Castañeda,
nia verrucosa by Brunner von Wattenwyl
2011, 2012) and of the series of papers on
(1878). This species was described from a
taxonomy of American Tettigoniidae pub-
female (its male is still unknown) and in-
lished by the second coauthor (Gorochov,
2012a, 2012b). cluded in the genus Aphidna Stål, 1874 in-
Until now, the genus Paraphidnia has in- correctly mentioned by Brunner von Wat-
cluded only three species: P. gallina Giglio- tenwyl as Aphidnia. Later Giglio-Tos (1898)
Tos, 1898 (type species) from Ecuador; P. proposed the genus Paraphidnia with the
© 2012 Zoological Institute, Russian Academy of Scienсes
O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA 205
only species P. gallina, described from two Most of measurements are given in the usu-
males in the same paper; he also indicated al manner (from the base to the apex), but
the differences of his genus from Stål’s one. some of them are defined as follows: length
Kirby (1906) established that Aphidna was of body, distance from the frons to the ab-
a junior synonym of Dysonia White, 1862 dominal apex excluding ovipositor; length
and transferred A. verrucosa to the genus of body with wings, distance from the frons
Paraphidnia. Finally, Rehn described P. to the apex of hind wings; length of prono-
lankesteri from a single male (Rehn, 1918) tum, maximal distance between the anterior
and prepared a key for the genera of Dyso- and posterior pronotal margins; length of
niini including Paraphidnia (Rehn, 1950: male subgenital plate, distance from its base
Dysoniae). After that, the genus has not to its apex excluding styli; length of oviposi-
attracted the attention of orthopterists in tor, distance from the subgenital plate apex
spite of its being insufficiently studied. to the ovipositor apex.
Here, twelve new species of Paraphid-
nia are described. All these species are very TAXONOMIC PART
similar to P. lankesteri by the armament of
Order ORTHOPTERA
hind femora and structure of male cerci.
Probably P. verrucosa is also closely related
Family TETTIGONIIDAE
to these species. But P. gallina has very dif-
ferent male cerci; this difference does not Subfamily PHANEROPTERINAE
allow its male to use these cerci (specialised
for the fixation of female during copulation) Tribe DYSONIINI
in the same manner as in the other conge-
Genus Paraphidnia Giglio-Tos, 1898
ners. Thus P. gallina and all the other spe-
cies clearly belong to different supraspecific Type species: Paraphidnia gallina Giglio-
taxa. However, these taxa have some gen- Tos, 1898.
eral characters allowing us to consider them Note. The type species is insufficiently
as two subgenera of the same genus. studied, and it belongs to the monotypical
subgenus Paraphidnia s. str. All the other
MATERIAL AND METHODS congeners strongly differ from P. gallina but
similar to each other. They are here includ-
This research is based on field observa- ed in the new subgenus described below.
tions of the first co-author and on material Some of its diagnostic characters allow us
deposited at the following institutions: Zo- to propose that it may be a separate genus.
ological Institute of the Russian Academy However, these supraspecific taxa have at
of Sciences, St Petersburg, Russia (ZIN); least a few common characters: the head
Museo de Historia Natural de la Universi- surface is somewhat rugose and/or granu-
dad Distrital Francisco José de Caldas, Bo- late, with a rather large but low and apically
gotá, Colombia (MUD); Instituto Nacional rounded tubercle on each gena near the
de Biodiversidad, Santo Domingo de Here- posteroventral corners of epicranium; the
dia, Costa Rica (INBio). Most of the speci- upper rostral tubercle forms a longitudinal
mens studied were collected in tropical crest; this crest is compressed from sides,
forests at light, but some specimens, camou- rather high or with a more or less high pos-
flaged under bryophytes and under lichens terior denticle (Figs 1–12, 69–77, 88–101);
of the family Lobariaceae, were found living the antennal flagellum has distinct sparse
in trees. and not very small inflations with numer-
The photographs of body parts of these ous short setae (Figs 38, 41, 45, 77); the
insects were made with Sony α300 and Can- pronotum has a more or less deep transverse
on 40D cameras as well as with Leica M216. fold in the anterior half, and its hind lobe
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
206 O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA
is rather short and somewhat erected, sepa- middle and hind femora with widened and
rated from the lateral lobes by a pair of dis- lamellar ventral spines; two of them (situ-
tinct posterolateral notches situated above ated most distally on outer side of hind
(not behind) the hind parts of these lobes femur) fused together and forming large
(Figs 13–17, 69–87, 109–117); the tegmi- lobe-like (strongly widened) spine hav-
nal surface is somewhat crumpled, having ing smaller additional lobule on distal edge
the texture more or less adapted for imita- (Figs 28, 32, 36); fore tibia with strongly
tion of tree bark with bryophytes and/or widened tympanal area; middle and hind
lichens; the mirror of left (upper) tegmen tibiae with widened and lamellar spines on
rather weakly distinct (its membrane with both sides of dorsal surface (Figs 19, 21, 23,
irregular veinlets, and its shape angularly 25, 27, 28, 30, 32, 34, 36). Tegmina long,
elongate; Figs 48, 50, 52, 54, 56, 78–87); with two branches on RS; in male, stridula-
the legs have many of the spines widened tory vein of left tegmen rather short (Figs
and lamellar, similar to those of the genus 102–108), and mirror of right (lower) teg-
Dysonia (including the trend for a strong men similar to that of left (upper) tegmen
increase of the distal outer spine of hind in structure but with less distinct irregular
femora); the abdominal tergites have rather veinlets (Figs 49, 51, 53, 55, 57); hind wings
large and more or less inflate tubercles. So, significantly longer than tegmina and with
at present it is reasonable to consider the texture of distal part of costal lobe more or
above-mentioned taxa as two subgenera of less similar to that of tegmina (Figs 38–47).
the same genus Paraphidnia s. l. Last abdominal tergite in male with shallow
and wide posteromedian notch; epiproct of
Subgenus Anaphidna Gorochov et male moderately large, flat, oval in distal
Cadena-Castañeda, subgen. nov. half (but having almost angular apex), and
with width and length almost equal; male
Type species: Paraphidnia (Anaphidna) cerci with rather long and spine-like outer
mexicana sp. nov. lobe (this lobe curved upwards and having
Diagnosis. Lower rostral tubercle low, acute apex), and with rather long and partly
rounded at apex, slightly narrower than lamellar inner lobe (this lobe movable in re-
basal half of scape, and with large median lation to outer lobe and having two projec-
ocellus anteriorly; upper rostral tubercle (= tions: rather long and acute upper one, and
crest) divided into three denticles (ante- shorter lower projection provided with a
rior one low and thin, with narrow median few very small apical hooks; Figs 58, 60, 61,
groove and narrowly rounded apices; mid- 63, 65, 67, 121–128); male subgenital plate
dle one also low and with similar groove, elongate (but not long and not large), with
but distinctly thicker, rounded or obtuse distinct posteromedian notch and thin and
in profile, and having large lateral ocelli on moderately long styles (Figs 59, 62, 64, 66,
lateral sides; posterior denticle higher and 68, 129–135); ovipositor short and strongly
diverse in shape; Figs 1–12, 69–76, 88–101, curved upwards (Figs 118–120).
109–117); first and second proximal seg- Included species. Type species; P. ver-
ments of antennae or only second one with rucosa; P. lankesteri; P. peruana sp. nov.; P.
finger-like process on medial part (Figs svetlanae sp. nov.; P. bezverkhovi sp. nov.;
110, 111). Pronotum with one tubercle on P. polestshuki sp. nov.; P. gracielae sp. nov.;
central part of each lateral lobe and a few P. quirozi sp. nov.; P. silvai sp. nov.; P. osae
tubercles on disc (latter tubercles rather sp. nov.; P. hernandezi sp. nov.; P. rubricor-
strongly varied in height, and some of them pus sp. nov.; P. rhinoceros sp. nov.
sometimes almost indistinct; Figs 13–17, Comparison. The new subgenus dif-
69–76, 78–84, 86, 87, 109–117). Fore coxa fers from the nominotypical subgenus
with short (almost tubercle-like) spine; in the posterior denticle of upper rostral
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O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA 207
crest much thinner or smaller (spine-like lum missing) having sparse small yellowish
or lobule-like, but not shaped as high and spots, and with numerous dark brown setae
rather long keel) and the other denticles on inflations of flagellum; mouthparts with
of this crest low (for comparison see Figs reddish tinge on clypeus and upper half of
1–12, 69–76, 88–101, 109–117 and Fig. labrum as well as reddish marble pattern on
77), presence of distinct pronotal tubercles mandibles and palpi; pronotum with light
(vs. pronotum without any tubercles; for brown disc and narrow upper part of lateral
comparison see Figs 13–17, 69–76, 78–84, lobes (lateral part of this light brown area
86, 87, 109–117 and Figs 77, 85), fusion of with reddish tinge, anterior part of this area
two distal outer spines of hind femora, and with a few small lighter and darker spots
male cerci divided into two lobes almost along anterior edge of disc, and hind part of
equal in length and movable in relation to this area mainly reddish brown); legs with
each other (in Paraphidnia s. str., these cerci numerous light brown areas on fore and
are undivided, with the distal part long and middle femora and on proximal half of hind
medially curved, and with the proximal part femora, with numerous reddish spots on dis-
having a small spine-like medial process im- tal half of hind femora and on all tibiae, with
movable in relation to the rest of cercus). small dark brown spots on inner side of fore
Etymology. This name is derived from femora (only one spot not far from femoral
the Greek prefix “ana-“ (un-, in-) and the base) and on outer side of middle femora
former genus Aphidna. (one spot near femoral apex, and 3–4 spots
on ventral outer keel), with almost blackish
Paraphidnia (Anaphidna) mexicana marks on outer and dorsal surfaces of proxi-
Gorochov, sp. nov. mal half of hind femora, with almost reddish
(Figs 1, 2, 13, 18, 19, 28, 29, 38, 39, 48, 49, brown tarsi having last segment light brown
58, 59) and with yellowish spot near base, and with
darkened thin spines as well as distal parts of
Holotype. Male; Mexico, Chiapas, north- apical spurs and of lamellar widened spines
eastern part of this state, Ocosingo Distr., on all legs; tegmina transparent with most
Selva Lacandona between Montes Azules veins and veinlets yellowish, with some vein-
Biosphere Reserve and Bonampak Natural lets of proximal and distal tegminal parts
Monument (near border with Guatemala), reddish brown, with some veins and veinlets
environs of Lacanja-Chansayab Vill., pri- between these tegminal parts almost dark
mary forest, at light, 20–27 May 2007, coll. brown, and with apical area whitish (this
M. Berezin, E. Tkatsheva (ZIN). area with small darkened marks; Fig. 38);
Description. Male (holotype). Coloura- hind wings transparent but with apical area
tion yellowish with following numerous of costal lobe almost whitish as well as with
marks: epicranium light brown with reddish subapical one brownish and having darker
tinge, whitish ocelli, a few small dark brown (brown) spot (Fig. 39); abdomen with small
spots on face (one under median ocellus, a brown and dark brown spots on ventrolat-
pair of spots near eyes and ventrolateral part eral parts of most tergites, on median part of
of antennal cavities, a pair of spots near lat- 3 distal tergites, and at base of epiproct (api-
eral corners of clypeus, and a pair of spots ces of cercal lobes also darkened).
on middle part of clypeus), and more or less Median ocellus almost as wide as long;
yellowish genae; antennae with light brown apex of anterior denticle of upper rostral
proximal part having reddish tinge on two crest narrow and practically not bifurcate
proximal segments and numerous small red- (Fig. 2); middle denticle of this crest widely
dish spots on flagellum (this flagellum also rounded in profile; posterior denticle of this
with two small yellowish spots), with red- crest high, spine-like, and with apical part
dish brown middle part (distal part of flagel- rather thin and directed upwards; Fig. 1);
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
208 O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA
Figs 1–12. Paraphidnia, male: 1, 2 – P. mexicana sp. nov.; 3 – P. lankesteri, holotype; 4 – P. verrucosa,
holotype; 5, 6 – P. peruana sp. nov.; 7, 8 – P. svetlanae sp. nov.; 9, 10 – P. bezverkhovi sp. nov.; 11,
12 – P. polestshuki sp. nov. Denticles of upper rostral crest from side (1, 3), and from side and slightly
above (5, 7, 9, 11); anterior denticle of upper rostral crest from above and slightly in front (2, 6, 8, 10,
12); posterior denticle of this crest from side (4). [3, after Rehn (1918); 4, after Eades et al. (2012)].
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA 209
Figs 13–17. Paraphidnia, male pronotum from side: 13 – P. mexicana sp. nov.; 14 – P. peruana
sp. nov.; 15 – P. svetlanae sp. nov.; 16 – P. bezverkhovi sp. nov.; 17 – P. polestshuki sp. nov.
each of two proximal antennal segments and low inner ventral keel; hind femur with
with medial process. Pronotal disc with a six inarticulate outer ventral spines (these
pair of moderately high tubercles near ante- spines as on middle femur but longer, and
rior edge, a pair of somewhat higher (spine- most distal spine strongly widened; Fig.
like) tubercles on middle part, and a pair 28), 1–2 inner ventral spines similar to
of thick rounded but not high tubercles on outer ones but not large and situated only
lateroproximal parts of hind lobe; outer sur- in distal part of femur, and two pairs of api-
face of lateral lobes clearly granulated along cal spinules (upper spinules distinctly lon-
ventral edge (Fig. 13). Fore femur with ger and slightly protruding behind femoral
shagreen inner surface, three short and in- apex; Fig. 29); fore tibia with a pair of small
articulate inner ventral spines in distal half, articulate dorsal spinules near distal edges
two inner apical spinules, and rather high of tympana (Fig. 18), six pairs of short and
outer ventral keel; middle femur with three thin ventral spines (including a pair of api-
short and inarticulate outer ventral spines cal spurs), one outer dorsoapical spur, very
(these spines widened and lamellar; distal high outer dorsal keel, and very low inner
one rather large), two outer apical spinules, one having three very low and rounded
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
210 O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA
Figs 18–27. Paraphidnia, male: 18, 19 – P. mexicana sp. nov.; 20, 21 – P. peruana sp. nov.; 22, 23 –
P. svetlanae sp. nov.; 24, 25 – P. bezverkhovi sp. nov.; 26, 27 – P. polestshuki sp. nov. Dorsal surface
of fore tibia near distal part of tympana (18, 20, 22, 24, 26); dorsal surface of middle tibia (19, 21,
23, 25, 27).
(weakly distinct) tubercles; middle tibia ner dorsoapical spur, seven pairs of thin and
with three outer and 4–5 inner dorsal in- articulate ventral spines (including a pair of
articulate spines similar to ventral spines apical spurs), and rather narrow distal part
of hind femur (inner spines distinctly nar- of tibia (Fig. 19); hind tibia with numerous
rower than outer ones), one thin and inar- moderately widened and inarticulate dor-
ticulate inner dorsal subapical spine, one in- sal spines, rather sparse thin and articulate
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O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA 211
ventral spines, six apical spurs, and rather reddish tinge, without dark spots on face,
narrow distal part (Fig. 28). Tegmina with and practically without yellowish spots on
general part of RS slightly longer than its antennae (distal antennal part missing),
longest branch (Fig. 38); stridulatory appa- but reddish marble pattern on palpi present;
ratus and distal part of costal lobe of hind pronotum without reddish tinge and with
wings as in Figs 39, 48, 49; stridulatory vein only hind tubercles darkened; legs with less
of left tegmen with 27 teeth. Cerci with distinct light brown spots (but reddish ar-
lower projection of inner lobe wide and eas on inner side of fore femora and on tibiae
having five very small apical hooks (Fig. and tarsi more or less present), with brown
58); subgenital plate with almost angular spot on middle femora near apex, and with
posteromedian notch (Fig. 59). dark brown band along dorsal surface of
Female unknown. hind femora (this band interrupted by three
Length in mm. Body 15; body with wings very light spots); tegmina with distinct
44; pronotum: median length 2.8, maximal greenish tinge, with reddish brown veinlets
length 3.6; tegmina 32; hind femora 13. only in basal part, and with less transparent
Comparison. The new species is most distal half (this half having oblique band
similar to P. lankesteri, however it differs near middle of tegmen almost whitish, more
from the latter in the posterior denticle of distal part marble greenish grey, and apical
upper rostral crest almost straight (not with part almost completely whitish; Fig. 41);
the apical part hooked) and inner lobe of hind wings with a few small dark spots near
male cerci with the lower projection clearly apical part of costal lobe (Fig. 40); abdo-
wider (for comparison see Figs 1, 3, 58, 60). men less spotted: with somewhat darkened
From P. verrucosa, the new species is distin- dorsal areas near base, with small dark spots
guished by the posterior denticle of upper on lateral parts of posterior abdominal half
rostral crest much longer (for comparison and on median part of 10th tergite as well as
see Figs 1, 4, 76), scape with a distinct fin- at base of epiproct, and with darkened api-
ger-like medial process, and pronotal disc ces of cercal lobes. Structure of head simi-
with distinctly higher tubercles. lar to that of P. mexicana, however middle
Etymology. The species name is a Latin denticle of upper rostral crest more angular
variant of the word Mexican (from Mexico). (angularly rounded), and posterior denticle
of this crest much lower and with 2 small
apical projections in profile (anterior one
Paraphidnia (Anaphidna) peruana Goro-
thinner and directed almost upwards; Fig.
chov, sp. nov.
5). Pronotal disc with anterior tubercles
(Figs 5, 6, 14, 20, 21, 30, 31, 40, 41, 50, 51,
almost as in P. mexicana, middle tubercles
61, 62)
distinctly lower than anterior ones (and
Holotype. Male; Peru, Junin Department, than middle tubercles of P. mexicana), and
Satipo Prov., ~35 km NE of Satipo Town, en- posterior tubercles clearly larger (higher)
virons of Mariposa Vill., ~1200 m, disturbed than all previous tubercles (and than poste-
primary forest, at light, 17–18 Oct. 2008,
rior tubercles of P. mexicana); outer surface
coll. A. Gorochov, M. Berezin, L. Anisyutkin,
of lateral pronotal lobes clearly granulated
E. Tkatsheva, V. Izersky (ZIN).
along ventral edge (Fig. 14). Legs also simi-
Paratypes. Two males, same data as for holo-
lar to those of P. mexicana in structure, but
type; 1 male, same province, ~40 km NE of Sati-
po Town, environs of Calabaza Vill., 11°26.7´S, with 3–4 inner ventral spines of fore femur,
74°46.4´W, 1492 m, at light, 22 Dec. 2010, coll. 5–6 outer ventral spines of hind femur (1–3
V. Sinyaev, S. Sinyaeva, V. Izersky (ZIN). proximal spines much smaller than oth-
Description. Male (holotype). Coloura- ers), 2–3 inner ventral spines of hind femur,
tion similar to that of P. mexicana but with 1–2 very small (low) lobes between latter
following differences: head almost without spines, 1–2 pairs of apical spinules of hind
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
212 O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA
Figs 28–37. Paraphidnia, male: 28, 29 – P. mexicana sp. nov.; 30, 31 – P. peruana sp. nov.; 32, 33 – P.
svetlanae sp. nov.; 34, 35 – P. bezverkhovi sp. nov.; 36, 37 – P. polestshuki sp. nov. Dorsal surface of
hind tibia (28, 30, 32, 34, 36); apical part of hind femur (29, 31, 33, 35, 37).
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(2): 204–233
O.J. CADENA-CASTAÑEDA & A.V. GOROCHOV. REVIEW OF NEOTROPICAL PARAPHIDNIA 213
Figs 38–47. Paraphidnia, male: 38, 39 – P. mexicana sp. nov.; 40, 41 – P. peruana sp. nov.; 42, 43 – P.
svetlanae sp. nov.; 44, 45 – P. bezverkhovi sp. nov.; 46, 47 – P. polestshuki sp. nov. Right (38) and left
(41, 43, 45, 47) tegmina; distal part of costal lobe of hind wing (39, 40, 42, 44, 46).
femur short (lower spinules very small, ab- ventral spines of fore tibia, 3–4 outer and
sent in one of legs; upper spinules short, not 7–8 inner dorsal spines of middle tibia (only
protruding behind femoral apex; Fig. 31), some of inner spines clearly smaller than
dorsal surface of fore tibia near distal edges outer ones; Fig. 21), and outer dorsal spines
of tympana having a pair of very small an- of hind tibia more widened than inner ones
gular tubercles (outer tubercle almost in- (Fig. 30). Tegmina with general part of RS
distinct; Fig. 20), four outer and 5–6 inner distinctly shorter than its longest branch
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