Systematics of the American Katydids (Orthoptera: Tettigoniidae). Communication 9

2020·1.5 MB·

by  Gorochov, A V

Proceedings of the Zoological Institute RAS
Vol. 324, No. 2, 2020, pp. 187–220
10.31610/trudyzin/2020.324.2.187
УДК 595.729
Systematics of the American Katydids (Orthoptera: Tettigoniidae). Communication 9
A.V. Gorochov
Zoological Institute of the Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Petersburg, Russia;
e-mail: [email protected]
ABSTRACT
The Neotropical genus Anaulacomera Stål, 1873 is tentatively divided into six subgenera: Anaulacomera s. str.;
Anallomes Stål, 1875, stat. nov.; Oecella Kirby, 1890, stat. nov.; Bovicercora subgen. nov.; Cervicercora subgen.
nov. and Munticercora subgen. nov. A new material on the two latter subgenera as well as on Oecella, stat. nov. are
considered with description of 25 new species and 9 new species groups from Peru, Ecuador and Bolivia: Bispinosa
Species Group of Cervicercora subgen. nov. with A. (C.) bispinosa sp. nov. and A. (C.) eubispinosa sp. nov.;
Superapex Species Group of Cervicercora subgen. nov. with A. (C.) superapex sp. nov.; Schunkei Species Group
of Cervicercora subgen. nov. with A. (C.) unispinosa sp. nov. and A. (C.) forceps sp. nov.; Cercalis Species Group
of Cervicercora subgen. nov. with A. (C.) paracercalis sp. nov., A. (C.) spinulata sp. nov., A. (C.) abbreviata sp.
nov., A. (C.) neohirsuta sp. nov., A. (C.) apicalis sp. nov. and A. (C.) denticulata sp. nov.; Daedala Species Group of
Cervicercora subgen. nov. with A. (C.) daedala sp. nov.; Mariposa Species Group of Cervicercora subgen. nov. with
A. (C.) mariposa sp. nov., possibly A. (C.) rectiapex sp. nov. and A. (C.) aenigma sp. nov.; Virgula Species Group of
Cervicercora subgen. nov. with A. (C.) virgula sp. nov., possibly A. (C.) grandiramus sp. nov. and A. (C.) parviramus
sp. nov.; Bellator Species Group of Cervicercora subgen. nov.; Confusa Species Group of Oecella; A. (O.) appendi-
culosa sp. nov.; A. (O.) tuberculosa sp. nov.; A. (O.) lobulosa sp. nov.; A. (O.) redunca sp. nov.; A. (M.) sclerogenitalis
sp. nov.; A. (M.) pseudoepiproctalis sp. nov. and A. (M.) spinolobata sp. nov. All the species from “Grupo Didieri” of
Anaulacomera s. l. (sensu Cadena-Castañeda) are included in the subgenus Deragramma Ebner, 1953 of the genus
Grammadera Brunner-Wattenwyl, 1878: G. (D.) impudica (Piza, 1952), comb. nov. and G. (D.) didieri (Cadena-
Castañeda, 2012), comb. nov.
Key words: America, Anaulacomera, new taxa, Phaneropterinae, Tettigoniidae
Систематика американских кузнечиков (Orthoptera: Tettigoniidae). Сообщение 9
А.В. Горохов
Зоологический институт Российской академии наук, Университетская наб. 1, 199034 Санкт-Петербург,
Россия; e-mail: [email protected]
РЕЗЮМЕ
Неотропический род Anaulacomera Stål, 1873 предварительно подразделен на шесть подродов:
Anaulacomera s. str.; Anallomes Stål, 1875, stat. nov.; Oecella Kirby, 1890, stat. nov.; Bovicercora subgen. nov.;
Cervicercora subgen. nov.; Munticercora subgen. nov. Рассмотрен новый материал по двум последним под-
родам, а также по Oecella, stat. nov.; из Перу, Эквадора и Боливии описаны 25 новых видов и 9 новых
видовых групп: группа Bispinosa (Cervicercora subgen. nov.) с A. (C.) bispinosa sp. nov. и A. (C.) eubispinosa sp.
nov.; группа Superapex (Cervicercora subgen. nov.) с A. (C.) superapex sp. nov.; группа Schunkei (Cervicercora
subgen. nov.) с A. (C.) unispinosa sp. nov. и, возможно, A. (C.) forceps sp. nov.; группа Cercalis (Cervicercora
subgen. nov.) с A. (C.) paracercalis sp. nov., A. (C.) spinulata sp. nov., A. (C.) abbreviata sp. nov., A. (C.) neohirsuta
sp. nov., A. (C.) apicalis sp. nov. и A. (C.) denticulata sp. nov.; группа Daedala (Cervicercora subgen. nov.) с A. (C.)
daedala sp. nov.; группа Mariposa (Cervicercora subgen. nov.) с A. (C.) mariposa sp. nov. и, возможно, A. (C.)
188 A.V. Gorochov
rectiapex sp. nov. и A. (C.) aenigma sp. nov.; группа Virgula (Cervicercora subgen. nov.) с A. (C.) virgula sp. nov. и,
возможно, A. (C.) grandiramus sp. nov. и A. (C.) parviramus sp. nov.; группа Bellator (Cervicercora subgen. nov.);
группа Сonfusa (Oecella); A. (O.) appendiculosa sp. nov.; A. (O.) tuberculosa sp. nov.; A. (O.) lobulosa sp. nov.;
A. (O.) redunca sp. nov.; A. (M.) sclerogenitalis sp. nov.; A. (M.) pseudoepiproctalis sp. nov. и A. (M.) spinolobata sp.
nov. Все виды из “Grupo Didieri”, помещенной Каденой-Кастанедой в род Anaulacomera, включены здесь
в подрод Deragramma Ebner, 1953 рода Grammadera Brunner-Wattenwyl, 1878: G. (D.) impudica (Piza, 1952),
comb. nov. и G. (D.) didieri (Cadena-Castañeda, 2012), comb. nov.
Ключевые слова: Америка, Anaulacomera, новые таксоны, Phaneropterinae, Tettigoniidae
INTRODUCTION (Cadena-Castañeda 2012, 2015; OSF). Thus, this ge-
nus must be divided into several preliminary subgen-
This communication is the ninth one in the se- era, some of which may be distinct genera. However,
ries of publications on the American Tettigoniidae the nominotypical subgenus is not very understanda-
(Gorochov 2012a, 2012b, 2014a, 2014b, 2015, 2016, ble in connection with the above-mentioned problem
2017, 2018). It contains some results of the study of a of its type species. Here, we tentatively ascribed the
new material on the genus Anaulacomera Stål, 1873 name Anaulacomera s. str. to “Grupo Gracilis” sensu
(Phaneropterinae: Phaneropterini: Anaulacomeri- Cadena-Castañeda (2012, 2015), because its repre-
na). The taxonomy of Anaulacomera is not very clear, sentatives have distinct whitish marks on the female
because its type species has been described from a tegminal dorsal field (as in the original description of
single female, but the majority of species included in Anaulacomera type species; Stål 1861).
this genus differ from each other and from some relat-
ed genera in the male abdominal apex only. Moreover, MATERIAL AND METHODS
a few species described in Anaulacomera by the pre-
vious authors belong to some other genera of the same The material studied (including type specimens)
subtribe Anaulacomerina: 1) Separatula Gorochov, is deposited at the Zoological Institute, Russian
2018 [= “Grupo Falcata” of the genus Anaulacomera Academy of Sciences, Saint Petersburg. All the spec-
sensu Cadena-Castañeda (2012, 2015)] with S. falca- imens are dry and pinned. The photographs of their
ta (Giglio-Tos, 1898), S. wilsoni (Cadena-Castañeda, morphological structures were made with a Leica
2015) and S. adunca Gorochov, 2018 (type species); MZ 16 stereomicroscope. Most part of this material
2) Grammadera Brunner-Wattenwyl, 1878 [its sub- was collected in mountain and plain rainforests at
genus Deragramma Ebner, 1953 = “Grupo Didieri” light. But some specimens were collected in these
in Anaulacomera sensu Cadena-Castañeda (2012, forests on leaves of bushes during the night work
2015)] with G. (D.) impudica (Piza, 1952), comb. with a flashlight. This work was carried out within
nov. and G. (D.) didieri (Cadena-Castañeda, 2012), the framework of a large project on the invertebrate
comb. nov. fauna of the Ene and Tambo river basins (Proyecto
The remaining species groups of Anaulacomera de Conservación de la Biodiversidad de la Selva
are also significantly distinguished from each other Amazónica: Identificación taxonómica de la fauna
by the male characters; some of these differences invertebrada en la cuenca del Rio Ene y Rio Tambo)
almost reach generic level. It is surprising that some under the supervision of the Peruvian and Ukrainian
subgenera in the neotropical genus Theia Brun- entomologist Volodymyr Izerskyy (Asociación para
ner-Wattenwyl, 1891 from the same tribe (Gorochov el desarrollo y conservación de los recursos natu-
2015: subtribe Pycnopalpina) are considered by Ca- rales del Perú – ACRENAP, Satipo). This project
dena-Castañeda (2014) and Orthoptera Species File is founded by the National Service of Natural Areas
(OSF) as separate genera; however, these “genera” Protected by the State (Servicio Nacional de Áreas
have differences in their most important morphologi- Naturales Protegidas por el Estado – SERNANP) of
cal characters more or less similar (in general volume) the Environment Ministry of Peru. The online cata-
to those between some groups of Anaulacomera, but logue Orthoptera Species File (Cigliano et al. 2020)
the latter genus is subdivided only into species groups is cited in this paper as OSF.
Systematics of American Tettigoniidae 9 189
SYSTEMATICS – Male cercus different in length, divided into branches
or having rather large specializations (spines, processes
or lobes) (Figs 20, 25, 30, 33, 35, 36, 39, 44, 48, 51–53,
Subfamily Phaneropterinae Burmeister, 1838
59, 63, 67, 71, 73, 78, 82, 83, 88, 89, 93, 95, 99, 100,
Tribe Phaneropterini Burmeister, 1838 104–112, 116, 117, 122, 123, 126, 130, 136, 139–144,
149, 151, 153, 155, 189, 191–195, 197, 198, 200) . . . 2
Subtribe Anaulacomerina Brunner-Wattenwyl,
2. Male epiproct rather small and simple (oval, angular
1878
and so on; Figs 24, 29, 38, 56, 58, 77, 85, 91, 96, 102,
113, 119, 189, 194, 197, 200), or with more or less
Note. In accordance to Gorochov (2018) and elongate posteromedian process (Figs 126, 128, 130,
OSF, this subtribe contains the following genera: 132, 134, 136, 137, 139, 140–144, 146, 147, 149); male
Anaulacomera; Grammadera; Abrodiaeta Brun- cercus elongate (distinctly protruding beyond genital
plate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
ner-Wattenwyl, 1878; Mendesius Piza, 1960;
– Male epiproct rather large (widened) and with more or
Phaneropteroides Piza, 1971; Montealegrezia Cade-
less deep posteromedian notch; male cercus short (not
na-Castañeda, 2012; Tenellulus Cadena-Castañeda,
protruding beyond genital plate), with two short and
2012; Rostellula Gorochov, 2018 and Separatula
rather strong processes (dorsal and ventral ones) . . .
Gorochov, 2018. However, Mendesius, Phanerop- . . . . . . . . . . . . subgenus Bovicercora subgen. nov.
teroides and Tenellelus are in need of examination of [Type species: Anaulacomera alfaroi Rehn, 1918.
their taxonomic positions, because structure of their Composition: type species; A. crassicerca Márquez
male tegminal mirror is unclear or unknown. Mayaudón, 1965; possibly A. securifera Brunner-Wat-
tenwyl, 1878 (all these species from “Grupo Alfaroi”);
possibly also A. spinata Brunner-Wattenwyl, 1878
Genus Anaulacomera Stål, 1873
(from “Grupo Spinata”). Etymology: from Bovinae
(mammal subfamily name), cercus (Latinized Greek
Note. Taxonomy of this genus is insufficiently un- morphological term) and Anaulacomera (generic
derstandable, because the structure of its male abdom- name) in connection with some similarity between
inal apex is more diverse than in many other genera of horns of Bovinae and male cerci of this subgenus.]
Phaneropterinae and was studied only for a part of its 3. Male last tergite with unpaired posteromedian lobe
very numerous congeners. Cadena-Castañeda (2012, sometimes having weakly concave posterior edge (Figs
2015) subdivided Anaulacomera s. l. into 17 species 24, 29, 38, 56, 58, 77, 85, 102, 103, 113), without pos-
terior lobe and often slightly concave posteriorly (Figs
groups, but two of them belong to some other genera
91, 96, 119, 128, 132, 134, 137, 141, 142), or fused with
(see above, in the introduction to this communica-
epiproct (146) . . . . . . . . . . . . . . . . . . . . . . . 4
tion). The remaining groups (15) are here reorganized
– Male last tergite with a pair of distinct posterior lobes
into six preliminary subgenera, some of which may be
and rather deep notch between them (Figs 189, 194,
divided into additional subgenera or erected up to
197, 200); male cercus distinctly bi- or trifurcate (Figs
generic level; these subgenera are described below, in 189, 191, 192, 194, 195, 197, 198, 200) . . . . . . . . . .
a subgeneric key for Anaulacomera s. l. . . . . . . . . . . subgenus Munticercora subgen. nov.
1. Male cercus simple, long or moderately long, undivid- [Type species: Anaulacomera poculigera Hebard, 1924.
ed into branches and without any large or moderately Composition: type species (from “Grupo Poculigera”;
large spines, processes or lobes (but with small spe- but A. franciscoi Cadena-Castañeda, 2012 and A.
cializations at or near apex: small spinules and lobules, trispinata Fianco, Faria et Cadena-Castañeda, 2019,
denticles, small hook and so on) . . . . . . . . . . . . . . also included by their authors in this group, lacking
. . . . . . . . . . . . . . subgenus Anaulacomera s. str. a pair of posterior lobes on male last tergite and be-
[Type species: Phaneroptera submaculata Stål, 1861. longing to unclear subgenus or subgenera); A. hebardi
Composition: type species; possibly almost all species Cadena-Castañeda, 2015 (from “Grupo Hebardi”);
of Cadena-Castañeda’s “Grupo Gracilis”, “Grupo A. rosea (Giglio-Tos, 1898) (from “Grupo Rosea”);
Inermis”, “Grupo Subinermis”, “Grupo Uncinata” A. (M.) pseudoepiproctalis sp. nov.; A. (M.) spinolo-
and “Grupo Latifolia” (belonging of type species and bata sp. nov.; A. (M.) sclerogenitalis sp. nov.; possibly
numerous other species to same subgenus problematic A. neofurcifera Otte, 1997 (from “Grupo Furcata”
but possible and based on presence of whitish spots on of Cadena-Castañeda). Etymology: from Muntiacus
female dorsal tegminal fields, because such colouration (mammal generic name) and Bovicercora (subgeneric
especially usual in “Grupo Gracilis” which most widely name) in connection with similarity of male cerci to
distributed in America).] horns of Muntiacus.]
190 A.V. Gorochov
4. Male epiproct rather small and simple (oval or roundly Brunner-Wattenwyl, 1893, A. sulcata Brunner-Wat-
angular; Figs 24, 29, 38, 56, 58, 77, 85, 91, 96, 102, 113, tenwyl, 1878, A. valentinae Cadena-Castañeda, 2012
119); male cercus with three or more processes (branch- (these species from “Grupo Lanceolata”; but all they,
es, rather large spines and lobes; Figs 20, 25, 30, 33, 35, including type species, insufficiently studied and
36, 39, 44, 48, 51–53, 59, 63, 67, 71, 73, 78, 82, 83, 88, separated here from other subgenera only on base of
89, 93, 95, 99, 100, 104–110, 111, 112, 116, 117, 122, characters given for this group by Cadena-Castañeda
123). . . . . . . . subgenus Cervicercora subgen. nov. (2015): “cercos bufurcados antes de la mitad de su
[Type species: Anaulacomera (Cervicercora) bispinosa longitud, generalmente cerca de la base del mismo”, but
sp. nov. Composition: type species; A. festae Giglio- male cerci of A. antillarum and A. valentinae with three
Tos, 1898; A. dama Rehn, 1913; A. cercalis Caudell, or more processes, including spines, and looking more
1918; A. schunkei Caudell, 1918; A. longicercata similar to those of Cervicercora subgen. nov.); possibly
Caudell, 1918, possible secondary homonym of A. lon- also A. intermedia Brunner-Wattenwyl, 1878 (Fig.
gicercata (Brunner-Wattenwyl, 1891), but belonging 155) and A. richteri Cadena-Castañeda, 2012 (from
of latter species to this genus problematic; A. hirsuta “Grupo Furcata” sensu Cadena-Castañeda).]
Hebard, 1926; A. apolinari Hebard, 1926; A. caucana
Hebard, 1926; possibly A. cornucervi Brunner-Wat- Species with unclear generic position previously
tenwyl, 1878, A. biramosa Brunner-Wattenwyl, 1891, included in Anaulacomera s. l.
A. bidentata Chopard, 1918 and A. bellator Rehn,
1920 (all these species from “Grupo Apolinari” sensu
Note. Cadena-Castañeda (2015) indicated 12
Cadena-Castañeda, which divided here into 8 species
species of this genus, which were left by him outside
groups (7 groups new) and including also 17 additional
new species). Etymology: from Cervus (mammal ge- of all species groups of this genus. In OSF, number
neric name) and Bovicercora (subgeneric name) in con- of such species was raised to 16 or 17. These species
nection with similarity of male cerci to Cervus horns.] are poorly described or described only from females,
– Male epiproct diverse; male cercus with only two pro- and their generic position inside Anaulacomerina
cesses (rather large spines or lobes; Figs 126, 130, 136, is also insufficiently understandable. Also a few in-
139–144, 149, 151, 153, 155) . . . . . . . . . . . . . . 5 sufficiently described species, tentatively included
5. Male epiproct with angular projection or specialized
here in the subgenus Anaulacomera, may belong to
process at apex (Figs 126, 128, 130, 132, 134, 136, 137,
different genus or genera. Some other species, includ-
139–144, 146, 147, 149); male cercus long or moderate-
ed by Cadena-Castañeda in his “Grupo Libidinosa”,
ly long (elongate), rather thin and with two processes
“Grupo Horti” and “Grupo Euceraiaformis”, may
(branches) or lobes in distal part (Figs 126, 130, 136,
be members of the genus Separatula; because these
139–144, 149, 151, 153) . . . . . . . . . . . . . . . . . .
. . . . . . . . subgenus Oecella Kirby, 1890, stat. nov. species groups have a large dorsal process in the basal
[Type species: Oecella furcifera Kirby, 1890 part of the male cercus, and this process is treated as
(= Anaulacomera harpago Brunner-Wattenwyl, 1878). homologous in these groups and in Separatula, judg-
Composition: type species; A. furcata Brunner-Wat- ing by the same abbreviation “P1” on some figures of
tenwyl, 1878; A. darwini Scudder, 1893; A. rusa Rehn, Cadena-Castañeda (2015: figs 26, 28–30). Moreover,
1917; A. biloba Brunner-Wattenwyl, 1878; A. lingulata almost all the females from my material cannot be
Piza, 1952; A. confusa Piza, 1975; A. maculifemora
exactly identified to species, species groups, subgen-
(Piza, 1977) (all these species from “Grupo Furcata”
era and sometimes even genera. For this reason, they
sensu Cadena-Castañeda); three new species described
will be considered separately after the descriptions of
here; possibly also A. juanchoi Cadena-Castañeda,
species and species groups.
2012 (from “Grupo Juanchoi”) and one new species
described here.
– Male epiproct small and simple (oval or roundly trian- Bispinosa Species Group of Cervicercora
gular); male cercus short or moderately long (elongate), subgen. nov.
more or less different in shape but always with two
distinct processes (spines, lobes, hooks) in proximal or
Note. This new group differs from all the other
middle parts . . . . . . . . . . . . . . . . . . . . . . . . .
species groups of this subgenus in the male cercus
. . . . . . . . subgenus Anallomes Stål, 1875, stat. nov.
with its ventroproximal process widened and flat-
[Type species: Anallomes nodulosa Stål, 1875. Compo-
sition: type species; possibly A. dentata Brunner-Wat- tened as well as having a posteromedial notch (Figs
tenwyl, 1878, A. lanceolata Brunner-Wattenwyl, 1878, 22, 27), with two dorsal spines, without ventral
A. maculata Brunner-Wattenwyl, 1878, A. antillarum spines or lobules between the distal and ventroprox-
Systematics of American Tettigoniidae 9 191
imal processes of this cercus, and without additional narrowly rounded apex which somewhat narrower
(lateral) process near ventroproximal one (Figs 20, than apical part of upper rostral tubercle, and with
25). This group includes two species: A. (C.) bispino- flattened anterior surface; median ocellus distinct,
sa sp. nov. and A. (C.) eubispinosa sp. nov. almost round, located between ventral parts of an-
tennal cavities and slightly not reaching these cavi-
Anaulacomera (Cervicercora) bispinosa sp. nov. ties; lateral ocelli also round but almost twice as large
(Figs 1–4, 20–24, 165) as median ocellus and located laterally on proximal
half of upper rostral tubercle. Pronotum with anteri-
or edge almost truncate, posterior edge of hind lobe
Etymology. This species name originates from
roundly convex, lateral lobes rather high and having
the Latin word “spinosa” (spinous) and the Latin
rounded ventral and posterior edges, and humeral
prefix “bi-“ (two), because the new species has two
notches moderately deep and clearly angular. Wings
dorsal spines on its male cercus.
long (strongly protruding beyond abdominal apex
Type material. Holotype – male, PERU: Junin
and apices of hind femora); tegmina clearly shorter
Department, Satipo Prov., ~25 km SE of Satipo
than hind wings, rather narrow, with almost straight
Town, environs of Rio Venado Vill., 1000–1200 m,
costal and anal edges, with rounded apex, with one
primary/secondary forest, at light, 20–23 October
distinct branch on RA directed distomedially, with
2008, A. Gorochov, M. Berezin, L. Anisyutkin,
E. Tkatsheva, V. Izerskyy. Paratypes: 4 males, same more or less normal RS running from proximal half
data as for holotype; 8 males, PERU, Ucayali De- of tegmen and having two branches in distal half of
partment, Atalaya Prov., ~35 km NWW of Atalaya tegmen, and with stridulatory apparatus as in Figs
Town on Ucayali River, environs of Sapani Vill., 3, 4. Last tergite with large (cup-like) dorsal part
~300 m, primary/secondary forest, at light, 26–31 having rather widely rounded and short posterome-
October 2008, A. Gorochov, M. Berezin, L. Anisyut- dian lobe directed downwards; epiproct small and
kin, E. Tkatsheva, V. Izerskyy. simple (oval); paraprocts also small and rounded
Description. Male (holotype). General appear- (Fig. 24); cercus with two large dorsal spines, long
ance typical of this genus. Body slender, medium-sized distal process and characteristic (widened, flattened
for this genus. Colouration yellowish (yellowish and strongly hooked) ventroproximal process (Figs
green in living specimen) with a pair of small reddish 20–22); genital plate narrowing to almost angularly
marks on dorsum of upper rostral tubercle between notched apex, but this notch with somewhat sinuous
lateral ocelli (and slightly behind them; Figs 1, 2), edges and almost spine-like lobules around it (Fig.
brown eyes and very small marks on subapical and 23); genitalia mainly membranous but with a sem-
apical tarsal segments, some tegminal veins brown- imembranous distal parts of a pair of lobes having
ish rose (two longitudinal veins along lateral edge of rounded apical tubercles (one on each lobe) with
stridulatory apparatus and one vein in basal area of numerous microscopical denticles, and with a pair of
dorsal field), stridulatory teeth and some membranes semisclerotized oblique structures in middle part of
between venation light brown (these membranes lo- genitalia (Fig. 165).
cated along lateral and medial edges of this apparatus Variations. Paratypes (males) very similar to
as well as in above-mentioned basal area and along holotype including colouration and structure of cer-
anal edge of remaining tegminal part), transparent ci, but reddish marks on upper rostral tubercle often
mirror and nearest membranes of stridulatory ap- absent, and anterior and posterior halves of this tu-
paratus as well as majority of membranes in hind bercle sometimes separated from each other by slight
wing, and rose venation of latter wing (but three narrowing.
veins along costal edge and nearest crossveins in this Female unknown.
wings yellowish). Upper rostral tubercle short (not Length (mm). Body 13.0–16.0; body with wings
projected before apex of lower rostral tubercle), with 34.0–36.0; pronotum 3.2–3.5; tegmina 24.0–27.0;
rounded (but not wide) anterior half slightly flat- hind femora 16.0–17.0.
tened apically, with somewhat widened posterior half Comparison. The new species differs from all the
(above lateral ocelli), and with dorsal median groove other species groups of this subgenus in the charac-
running also to hind part of head; lower rostral ters listed in the note on Bispinosa Species Group
tubercle barely not reaching upper one, with rather above.
192 A.V. Gorochov
Anaulacomera (Cervicercora) eubispinosa Superapex Species Group of Cervicercora
sp. nov. subgen. nov.
(Figs 5, 25–29)
Note. This group is new one and characterized
Etymology. This species name originates from the by the following characters of male cercus: its ven-
species name A. bispinosa sp. nov. and the Latinized troproximal process is as in the Bispinosa Species
Greek prefix “eu-” (true), because the new species is Group, i.e. widened, flattened and with a posterome-
most similar and related to A. bispinosa sp. nov. dial notch (Figs 31, 34); dorsal cercal edge is with one
Type material. Holotype – male, PERU: Junin spine; ventral cercal edge is with one spine between
Department, Satipo Prov., ~40 km NE of Satipo the distal and ventroproximal processes; additional
Town, environs of Calabaza Vill., ~2000 m, primary large lateral process near ventroproximal process is
forest, at light, 16–17 October 2008, A. Gorochov, absent (Figs 30, 33).
M. Berezin, L. Anisyutkin, E. Tkatsheva, V. Izer- Composition: A. (C.) superapex sp. nov.; possibly
skyy. Paratype – male, same data as for holotype. A. (C.) dama Rehn, 1913.
Description. Male (holotype). Size, colouration
and structure of body similar to those of holotype of Anaulacomera (Cervicercora) superapex sp. nov.
A. (C.) bispinosa sp. nov. but with following differ- (Figs 6, 30–34, 166)
ences: upper rostral tubercle without reddish marks
(as in some paratypes of A. bispinosa sp. nov.); each Etymology. This species name originates from
tegmen with two distinct branches on RA; its dorsal the Latin word “apex” (top, apex) with the Latin pre-
field with white stridulatory vein having stridulatory fix “super-” (over-, super-), because the male cercus of
teeth brown, white chord near previous vein, partly the new species has a very long apical part.
whitish and partly brown venation in basal area, and Type material. Holotype – male, PERU: Junin
light brown space behind mirror (this space yellow- Department, Satipo Prov., ~25 km SE of Satipo
ish in A. bispinosa sp. nov.; compare Figs 3, 4 and Town, environs of Rio Venado Vill., 1000–1200 m,
5); all veins of hind wing yellowish; epiproct larger primary/secondary forest, at light, 20–23 October
(wider; Fig. 29); cercus with distinctly shorter dor- 2008, A. Gorochov, M. Berezin, L. Anisyutkin,
sal spines and longer distal process as well as longer E. Tkatsheva, V. Izerskyy. Paratypes: 8 males, same
and narrower ventroproximal process having wider data as for holotype; 2 males, same province, Rio
distomedial notch (Figs 25–27); genital plate with Tambo Distr., 6 km N of Pichiguia Vill., protected
rounded posteromedian notch (Fig. 28). area “Reserva Comunal Ashaninka”, “11.358244°S,
Variations. Second male with apical part of distal 74.0320473°W”, ~500 m, primary forest, at light, 14–
cercal process slightly more curved medially, and 23 November 2017, A. Gorochov, G. Irisov; 2 males,
with posteromedian notch of genital plate almost PERU, Ucayali Department, Atalaya Prov., ~35 km
roundly angular (but edges of this notch not sinuate). NWW of Atalaya Town on Ucayali River, environs
Female unknown. of Sapani Vill., ~300 m, primary/secondary forest, at
Length (mm). Body 16.0–17.0; body with wings light, 26–31 October 2008, A. Gorochov, M. Bere-
36.0–38.0; pronotum 3.5–3.6; tegmina 27.0–28.0; zin, L. Anisyutkin, E. Tkatsheva, V. Izerskyy.
hind femora 16.0–16.5. Description. Male (holotype). Size, colouration
Comparison. The new species differs from its and structure of body similar to those of A. (C.) bispi-
most similar and related species A. (C.) bispinosa nosa sp. nov. and A. (C.) eubispinosa sp. nov. How-
sp. nov. in the male epiproct somewhat larger, dorsal ever, this male with following characteristic features:
spines of male cercus shorter, distal process of this upper rostral tubercle with slight rose marks on dor-
cercus different in shape (compare Figs 20 and 25), sal part; tegmina with darkened parts (brown strid-
ventroproximal process of this cercus narrower and ulatory vein, most part of basal area, and medial half
longer as well as with wider distomedial notch, and of stridulatory apparatus including chord and space
posteromedian notch of male genital plate more behind mirror) only in region of stridulatory appara-
rounded and having non-sinuous edges (see Figs 22, tus and with whitish marks only in right tegmen (on
23 and 27, 28). stridulatory vein and near it; Fig. 6); hind wing with
Systematics of American Tettigoniidae 9 193
Figs 1–9. Anaulacomera (Cervicercora subgen. nov.), males: 1–4 – A. (C.) bispinosa sp. nov.; 5 – A. (C.) eubispinosa sp. nov.; 6 – A. (C.)
superapex sp. nov.; 7 – A. (C.) unispinosa sp. nov.; 8 – A. (C.) forceps sp. nov.; 9 – A. (C.) paracercalis sp. nov. Head in front (1) and from
above (2); stridulatory apparatus of left (3) and right (4) tegmina as well as of both tegmina (5, 6) from above; this stridulatory apparatus
and head with pronotum from above (7–9).
majority of longitudinal veins rose, but such veins with ventral spine between this and ventroproximal
along costal edge and all crossveins yellowish to processes well developed and having widened basal
whitish; upper rostral tubercle with slight narrowing part (Fig. 30), and with latter (ventroproximal) pro-
between anterior and posterior halves; last tergite cess as in Fig. 31; genital plate with apical part as in
and epiproct almost as in A. (C.) eubispinosa sp. nov.; Fig. 32; genitalia with semimembranous distal parts
cercus with one dorsal spine, with very long distal of a pair of lobes somewhat smaller than in Bispina
process having dorsoapical part denticulate (Fig. 30), Species Group and lacking any distinct tubercles and
194 A.V. Gorochov
Figs 10–19. Anaulacomera (Cervicercora subgen. nov.), males: 10 – A. (C.) spinulata sp. nov.; 11 – A. (C.) abbreviata sp. nov.; 12 – A. (C.)
neohirsuta sp. nov.; 13 – A. (C.) apicalis sp. nov.; 14 – A. (C.) denticulata sp. nov.; 15 – A. (C.) daedala sp. nov.; 16 – A. (C.) mariposa sp.
nov.; 17 – A. (C.) rectiapex sp. nov.; 18 – A. (C.) virgula sp. nov.; 19 – A. (C.) aenigma sp. nov. Head, pronotum and stridulatory apparatus
of both tegmina from above (10, 16, 19); stridulatory apparatus of both tegmina from above (11–15, 17, 18).
Systematics of American Tettigoniidae 9 195
denticles, and with a pair of oblique semimembranous Town, environs of Calabaza Vill., ~2000 m, primary
plates in middle part of genitalia (Fig. 166). forest, at light, 16–17 October 2008, A. Gorochov,
Variations. Other males sometimes with darker M. Berezin, L. Anisyutkin, E. Tkatsheva, V. Izerskyy.
(almost dark brown) or lighter (light brown) dark- Paratypes: 2 males, same data as for holotype; 1 male,
ened parts of dorsal tegminal field; their cerci also same province, ~25 km SE of Satipo Town, environs of
varied from barely shorter to somewhat longer, i.e. Rio Venado Vill., 1000–1200 m, primary/secondary
distal cercal process 2.5–6.5 times as long as ventral forest, at light, 20–23 October 2008, A. Gorochov, M.
cercal spine between this process and ventroproximal Berezin, L. Anisyutkin, E. Tka tsheva, V. Izerskyy; 3
cercal process; these ventral spine and ventroproxi- males, same province, Rio Tambo Distr., 6 km N of
mal process sometimes as in Figs 33, 34. Pichiguia Vill., protected area “Reserva Comunal
Female unknown. Ashaninka”, “11.358244°S, 74.0320473°W”, ~500
Length (mm). Body 13.0–16.5; body with wings m, primary forest, at light, 14–23 November 2017, A.
32.0–35.0; pronotum 3.2–3.4; tegmina 24.0–26.0; Gorochov, G. Irisov.
hind femora 14.5–16.0. Description. Male (holotype). General appear-
Comparison. The new species differs from A. (C.) ance very similar to that of holotype of A. (C.) supera-
dama in the male distal cercal process clearly longer pex sp. nov. but with following differences: apical part
and directed partly downwards (not partly upwards) of upper rostral tubercle barely wider and almost sem-
as well as in the male dorsal cercal spine directed iglobular, separated from rest of this tubercle by dis-
strictly upwards (vs. this spine directed upwards/ tinct narrowing (this narrowing somewhat more dis-
backwards and almost parallel to distal process; tinct than in all previous species described here); this
Fig. 51). From all the other species of Cervicercora tubercle with rose loop-like line on dorsum (Fig. 7);
subgen. nov., the new species is distinguished by the longitudinal vein in basal part of dorsal tegminal field
characters listed above, in the note about Superapex yellowish (not rose; Fig. 7); epiproct barely narrower
Species Group. (but not smaller; Fig. 38); cercus with dorsal spine
having its basal part somewhat separated from rest
Schunkei Species Group of Cervicercora of cercus by short and almost vertical posterolateral
subgen. nov. groove, with distal process shorter but rather large as
well as having small and almost angular dorsal pro-
Note. This new group is distinguished from all jection near dorsal spine (this distal process widened
the other species groups of this subgenus by the fol- under latter projection, narrowing to apex, somewhat
lowing characters of male cercus: its ventroproximal arcuately curved and having almost completely den-
process is as in Bispinosa and Superapex Species ticulate dorsal edge; Fig. 39), with two ventral lobules
Groups (Figs 37, 41, 45, 49); dorsal cercal edge is between distal and ventroproximal processes (larger
with one spine; ventral cercal edge is with two spines and almost horizontal ventrodistal lobule, and smaller
or lobules between the distal and ventroproximal and hooked as well as partly denticulate ventroproxi-
processes; additional large lateral process near ven- mal lobule; Figs 39, 40, 42), and with ventroproximal
troproximal one is absent (Figs 35, 36, 39, 44, 48). process having posteromedial (subapical) lobe slightly
Composition: A. (C.) schunkei Caudell, 1918 and less projected backwards (Fig. 41); genital plate with
A. (C.) forceps sp. nov.; possibly A. (C.) festae Giglio- posteromedian notch as in Fig. 43. Genitalia almost
Tos, 1898 and A. (C.) unispinosa sp. nov. as in A. (C.) superapex sp. nov.
Variations. Darkened parts of tegmina sometimes
Anaulacomera (Cervicercora) unispinosa sp. nov. slightly darker or barely lighter; genital plate with
(Figs 7, 38–43) apical part also somewhat varied: from that as in Fig.
43 to almost that as in Fig. 32.
Etymology. This species name originates from Female unknown.
the Latin word “spinosa” (spinous) and the Latin Length (mm). Body 11.0–14.0; body with wings
prefix “uni-“ (one), because the new species has one 28.0–32.0; pronotum 2.9–3.1; tegmina 2.2–2.3; hind
dorsal spine on its male cercus. femur 14.0–15.0.
Type material. Holotype – male, PERU: Junin Comparison. The new species is very similar to
Department, Satipo Prov., ~40 km NE of Satipo A. (C.) festae in the shape of its male cerci. However,
196 A.V. Gorochov
male syntype of A. (C.) festae, judging by its pho- distal process (this process spine-like and without
tographs in OSF, has the male cercal distal process denticulation), somewhat longer (than in A. unispi-
with a clearly concave dorsal edge (i.e. this edge is nosa sp. nov.) ventrodistal lobule having rounded
with a rather large dorsoproximal projection, but in and setose apical part, moderately low widening near
A. unispinosa sp. nov., this edge completely convex base of this lobule, small ventral spinule located be-
and denticulate from small dorsoproximal projection fore this base but also near it (this spinule probably
to apical part; compare Figs 35, 36 and 39); the latter homologous to ventroproximal lobule of A. unispino-
syntype has also the distal process of the left male sa sp. nov.), moderately high keel along ventromedial
cercus distinctly shorter than that of the right male cercal surface having roundly angular apical projec-
cercus (such asymmetry may be a result of strong tion, and characteristic (with very large apical lobe)
medial curvature of the apical part of this process ventroproximal process (Figs 44–46); genital plate
in the left cercus, or this part might be deformed with very narrow and rather deep posteromedian
or damaged; Figs 35, 36), but the above-mentioned notch (Fig. 47); genitalia almost as in A. (C.) super-
process even in the right cercus is clearly smaller apex sp. nov. and practically indistinguishable from
(shorter) than in A. (C.) unispinosa sp. nov. (see Figs those of A. (C.) schunkei (Fig. 167).
35 and 39). Differences from all the other species of Variations. Some males almost without rose
this subgenus are given in the note about Schunkei marks on dorsum of upper rostral tubercle; ventrodis-
Species Group. tal lobule of cercus sometimes directed more medial-
ly or partly upwards (this lobule probably movable in
Anaulacomera (Cervicercora) forceps sp. nov. relation to rest of cercus).
(Figs 8, 44–47) Female unknown.
Length (mm). Body 12.0–14.0; body with wings
Etymology. This species name is the Latin word 31.0–33.0; pronotum 3.0–3.2; tegmina 23.0–24.5;
“forceps” (ticks, tongs) in connection with the struc- hind femora 13.0–14.0.
ture of the male cercal distal part. Comparison. The new species is most similar
Type material. Holotype – male, PERU: Junin to A. (C.) schunkei but distinguished from it by the
Department, Satipo Prov., ~25 km SE of Satipo distal process of male cercus longer and slightly nar-
Town, environs of Rio Venado Vill., 1000–1200 m, rower, subdistal part of this cercus (near the base of
primary/secondary forest, at light, 20–23 October its movable ventrodistal lobule) lower, longitudinal
2008, A. Gorochov, M. Berezin, L. Anisyutkin, keel of this cercus located along its ventromedial
E. Tkatsheva, V. Izerskyy. Paratypes: 3 males, same surface and having distinct apical projection (vs. this
data as for holotype. keel located along medial cercal surface and with-
Description. Male (holotype). Size, coloration out apical projection, i.e. almost gradually reducing
and structure of body almost as in A. (C.) unispinosa towards apex of this keel), apical (lateral) lobe of
sp. nov. but with following characteristic features: ventroproximal process of this cercus larger, and
upper rostral tubercle with reddish rose lateral ocelli subapical (medial) lobe of this process less projected
(Fig. 8); tegmina with darkened parts light brown backwards (compare Figs 44, 45 and 48, 49). From
and with rest of venation yellowish (Fig. 8); cercus A. (C.) unispinosa sp. nov. and A. (C.) festae as well
with smaller dorsal spine, clearly shorter and thinner as from all the other species of this subgenus, the new
Figs 20–51. Anaulacomera (Cervicercora subgen. nov.), males: 20–24 – A. (C.) bispinosa sp. nov.; 25–29 – A. (C.) eubispinosa sp. nov.;
30–34 – A. (C.) superapex sp. nov.; 35–37 – A. (C.) festae Giglio-Tos, syntype; 38–43 – A. (C.) unispinosa sp. nov.; 44–47 – A. (C.)
forceps sp. nov.; 48–50 – A. (C.) schunkei Caudell; 51 – A. (C.) dama Rehn. Left cercus from side (20, 25, 30, 33, 36, 39, 44, 48, 51); its
apical part, posterolateral view (21, 26); ventroproximal cercal process, posterodorsal view (22, 27, 31, 34, 37, 41, 45, 49); genital plate
(23, 28, 47) and its distal part (32, 43) from below; posterior part of last tergite and epiproct with cercal bases (24, 38) or without these
bases (29); right cercus without apices of ventral lobules and of posteroventral process, inner view (35); distal half of smaller ventral
lobule of left cercus, ventrolateral view (40); middle portion (42) and distal half (46, 50) of right cercus from below. [35–37 – after
photographs from OSF; 51 – after Rehn (1913), modified.]. Abbreviations: a, apical lobe of ventroproximal process; d, distal process of
cercus; ds, dorsal spine or spines of cercus; gp, genital plate; k, longitudinal keel on ventromedial or medial surface of cercus; s, subapical
lobe of ventroproximal process; v, vd, ventral lobules of cercus (ventroproximal and ventrodistal lobules, respectively); vp, ventroproxi-
mal process of cercus.