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https://doi.org/10.11646/zootaxa.5524.1.1
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by R. Mariño-Pérez: 18 Jun. 2024; published: 21 Oct. 2024
5524
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Copyright © 2024 Magnolia Press Monograph
Revision of the polyphyletic genus Hirrius Bolívar, 1887 (Orthoptera:
Tetrigidae), with descriptions of three new genera and insights into antennal
sensilla morphology
JOSIP SKEJO1,+,*, MARTIN HUSEMANN1,2,+, ROMEO PATANO JR.3,+, JOSEF TUMBRINCK4,
TARAS I. PUSHKAR5,+, MARKO PAVLOVIĆ6, ALMA MOHAGAN3, JAN-HENRIK PAMIN1 &
NIKO KASALO7,+
1
Leibniz-Institut zur Analyse des Biodiversitätswandels - Standort Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
2Staatliches Museum für Naturkunde Karlsruhe, Erbprinzenstraße 13, 76133 Karlsruhe, Germany
3 Center for Biodiversity Research and Extension in Mindanao & Institute of Biological Sciences, Animal Biology Division, Central
Mindanao University, Musuan, Maramag, Bukidnon 8710, Philippines
4 Auf der Hees 1, D-41849 Wassenberg, Germany
5 Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Khmelnytskogo Str. 15, Kyiv, UA-01030, Ukraine;
6 Prva privatna gimnazija Varaždin s pravom javnosti, Franja Supila 22, Varaždin, HR-42000, Croatia
7 Matice hrvatske 11, 80101 Livno, Bosnia and Herzegovina
+ equal contribution
* Corresponding author,
�
skejo.josip@gmail.com
Josip Skejo: https://orcid.org/0000-0002-2554-4499;
�
skejo.josip@gmail.com
Martin Husemann: https://orcid.org/0000-0001-5536-6681;
�
martin.husemann@smnk.de
Romeo Patano Jr.: https://orcid.org/0000-0001-5020-6048;
�
romeonojrpatano@gmail.com
Josef Tumbrinck: https://orcid.org/0000-0002-8955-7934;
�
j.tumbrinck@t-online.de
Taras I. Pushkar: https://orcid.org/0009-0008-5845-8803;
�
taras.i.pushkar@gmail.com
Marko Pavlović: https://orcid.org/0000-0002-2956-6548;
�
marko09.pavlovic@gmail.com
Alma Mohagan: https://orcid.org/0000-0002-8303-5131;
�
almsmohagan@gmail.com
Jan-Henrik Pamin: https://orcid.org/0009-0009-2161-6764;
�
j.pamin@leibniz-lib.de
Niko Kasalo: https://orcid.org/0000-0002-3139-6349;
�
nikokasalo5@gmail.com
SKEJO ET AL.
2 · Zootaxa 5524 (1) © 2024 Magnolia Press
JOSIP SKEJO, MARTIN HUSEMANN, ROMEO PATANO JR., JOSEF TUMBRINCK, TARAS I. PUSHKAR,
MARKO PAVLOVIĆ, ALMA MOHAGAN, JAN-HENRIK PAMIN & NIKO KASALO
Revision of the polyphyletic genus Hirrius Bolívar, 1887 (Orthoptera: Tetrigidae), with descriptions of
three new genera and insights into antennal sensilla morphology
(Zootaxa 5524)
61 pp.; 30 cm.
21 Oct. 2024
ISBN 978-1-77973-181-4 (paperback)
ISBN 978-1-77973-182-1 (Online edition)
FIRST PUBLISHED IN 2024 BY
Magnolia Press
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 3
Table of Contents
Introduction ..............................................................................................4
Materials and methods .....................................................................................5
Museum abbreviations and online databases ......................................................................5
Taxonomy, nomenclature, and terminology .......................................................................5
Measurements .............................................................................................6
Overview of all the localities ..................................................................................6
Results ...................................................................................................8
Taxonomic part ...........................................................................................8
Subfamily Metrodorinae ....................................................................................8
Genus group Melainotettix ..................................................................................8
Genus Hirrius .............................................................................................8
A key to species of Hirrius ..................................................................................10
Catalogue of Hirrius species .................................................................................10
Hirrius punctatus ..........................................................................................10
Hirrius mindanaensis .......................................................................................14
Hirrius ruber sp. nov. ......................................................................................20
Genus Guentheracris gen. nov. ..............................................................................23
Guentheracris scrobiculata comb. nov. ........................................................................24
Tetrigidae: incertae sedis ...................................................................................26
Genus Parahirrius gen. nov. ................................................................................26
A key to species of Parahirrius ...............................................................................27
Catalogue of Parahirrius species ..............................................................................27
Parahirrius parvus sp. nov. ..................................................................................27
Parahirrius amorosus sp. nov. ...............................................................................32
Subfamily Scelimeninae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Tribe Discotettigini ........................................................................................36
Genus Zvierckia gen. nov. ..................................................................................36
A key to species of Zvierckia .................................................................................37
Catalogue of Zvierckia species ...............................................................................38
Zvierckia montana .........................................................................................38
Zvierckia sarasinorum ......................................................................................39
Zvierckia storozhenkoi sp. nov. ...............................................................................42
Widened antennal segments in Zvierckia and Phaesticus .........................................................49
Zvierckia storozhenkoi ......................................................................................49
Phaesticus mellerborgi ......................................................................................51
Discussion ...............................................................................................57
Acknowledgements ........................................................................................59
Literature ...............................................................................................59
Abstract
Discotettiginae were recently synonymized with Scelimeninae, but the polyphyletic genus Hirrius Bolívar, 1887, with
five species endemic to the Philippines and Sulawesi, remained an unsolved issue. Besides similarly widened subapical
antennomeres, head and pronotum, other traits suggest that Hirrius members belong to different subfamilies.
The genus is now split into four genera, with four new species. Hirrius (Tetrigidae: incertae sedis) now includes
three species, one of which is new, all endemic to Mindanao, the Philippines: H. punctatus (Stål 1877), H. mindanaensis
(Günther, 1938), and H. ruber Skejo, Patano et Kasalo sp. nov. A lectotype is designated for H. punctatus, because the
type series was found to contain two different species belonging to two different genera. Three new genera are described.
Genus Parahirrius Skejo, Patano et Kasalo gen. nov. (Tetrigidae: incertae sedis) consists of P. parvus Skejo, Patano et
Kasalo sp. nov. and P. amorosus Skejo, Patano et Kasalo sp. nov., both endemic to Mindanao. Genus Zvierckia Skejo,
Tumbrinck et Pushkar gen. nov. (Scelimeninae: Discotettigini) includes Z. montana (Günther, 1937), comb. nov., Z.
sarasinorum (Günther, 1937), comb nov., and Z. storozhenkoi Skejo et Tumbrinck sp. nov., all endemic to Sulawesi.
Genus Guentheracris Skejo, Tumbrinck et Pushkar gen. nov. (Tetrigidae: “Asian Metrodorinae”) includes G. scrobiculata
(Günther, 1937), comb. nov. from Sulawesi.
Subapical antennomeres of Zvierckia storozhenkoi sp. nov. and Phaesticus mellerborgi (Stål, 1855) were scanned for
the first time and compared to Discotettix belzebuth (Serville, 1838). Zvierckia gen. nov. and Discotettix Costa, 1864, both
Scelimeninae: Discotettigini members, share many features of the antennal morphology, such as many basiconic sensilla
on the antennal margins giving the ridge a saw-like impression. On the other hand, antennae of Phaesticus Uvarov, 1940
SKEJO ET AL.
4 · Zootaxa 5524 (1) © 2024 Magnolia Press
contain many elongated placoid sensilla, which are fully absent in the subapical widened segments of Zvierckia gen. nov.
and Discotettix.
Key words: Scelimeninae, Metrodorinae, taxonomy, new taxa, new combinations, morphology, antennae, coeloconic
field, basiconic sensilla, cuticular plates, elongated placoid sensilla
Introduction
Members of the genus Hirrius Bolívar, 1887 are characterized by the reduction of wings (small size or complete
absence) and by the widened subapical antennal segments (Fig. 1) (Günther 1937, 1938; Skejo 2017). The genus
FIGURE 1. Already the antenna of the former members of the genus Hirrius, currently Hirrius mindanaensis, Guentheracris
scrobiculata comb. nov., Zvierckia sarasinorum comb. nov., Zvierckia montana comb. nov., and Zvierckia storozhenkoi sp.
nov. Evidence the diversity of the Tetrigidae representatives assigned to this genus till this revision.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 5
was originally established to include only Arulenus punctatus Stål, 1877 from Mindanao, but currently counts five
seemingly unrelated species from Mindanao (H. punctatus, H. mindanaensis Günther, 1938) and Sulawesi (H.
montanus Günther, 1937, H. sarasinorum Günther, 1937, H. scrobiculatus Günther, 1937) (Stål 1877; Bolívar 1887;
Günther 1937).
Pygmy grasshoppers’ (Tetrigidae) taxonomy had been based mostly on the presence and the absence of wings,
morphology of pronotum, and morphology of legs (Bolívar 1887; Hancock 1907), but recently also head characters
came into use (Devriese 1999; Tumbrinck 2014) and have proven since to be very important in supergeneric
taxonomy (Muhammad et al. 2018). The presence of the widened antennomeres was thought to be a Discotettiginae
synapomorphy, but was proven to represent only a homoplasy and Discotettiginae were synonymized with
Scelimeninae (Skejo et al. 2022).
Antennae represent the most important sensory organ of insects and were under strong selective pressure because
of biotic and abiotic constraints. With antennae, insects can taste (gustatory sensilla), smell (olfactory sensilla),
move and balance (mechanosensory), sense temperature (sensilla with thermoreceptors) and humidity (sensilla with
hygroreceptors). Five main types of sensilla can be identified by the shape, that being trichoid, basiconic, placoid
and coeloconic sensilla. Furthermore, three types can be distinguished by the presence of pores and these are aporous
(usually for thermoreception and hygroreception), uniporous (gustation chemoreceptors) and multiporous (olfactory
chemoreceptors) (Slifer 1970; McIver 1975; Altner & Prillinger 1980; Zacharuk 1980; Hallberg & Hansson 1999).
Fine structure of the antennae, including information on the type, number and arrangement of antennal sensilla
was never used as a diagnostic trait in Tetrigidae, even though it showed perspective (Bland 1991, Kuřavová et al.
2017).
The lack of wings and the widened subapical antennal segments are found in a variety of unrelated Tetrigidae
(Cigliano et al. 2023), hence, the aim of this study was to review type specimens of all five known Hirrius species, re-
define the genus Hirrius in the light of modern Tetrigidae systematics, to describe new genera and new species from
the Philippines and Sulawesi, and to provide data on the morphology of the widened antennomeres’ morphology in
Zvierckia gen. nov. and Phaesticus Uvarov, 1940.
Materials and methods
Museum abbreviations and online databases are given as follows:
CJT—Collection Josef Tumbrinck, Wassenberg, Germany;
CMU—Central Mindanao University, University Museum, Zoological Section, Mindanao, Philippines;
Flickr—Flickr webpage (https://www.flickr.com/);
iNaturalist—iNaturalist webpage (https://www.inaturalist.org/);
MfN—Museum für Naturkunde, Berlin, Germany;
NHMB—now NMBA, see below;
NHRS—Naturhistoriska Riksmuseet, Stockholm, Sweden;
NMBA—Naturhistorisches Museum Basel, Basel, Switzerland;
OSF—Orthoptera species file database (Cigliano et al. 2023);
SDEI—Das Senckenberg Deutsche Entomologische Institut, Müncheberg, Germany;
SMTD—Staatliche Naturhistorische Sammlung Dresden, Museum für Tierkunde, Dresden, Germany;
ZISP—Zoological Institute, St. Petersburg, Russia
ZMH—Zoologisches Museum Hamburg, Hamburg, Germany;
Taxonomy, nomenclature and terminology. Taxonomy follows the Orthoptera species file database (Cigliano et
al. 2023), while all nomenclatural acts are in accordance with the International Code of the Zoological nomenclature
(ICZN, 1999). Morphological terminology follows Devriese (1991, 1999) and Tumbrinck (2014), while antennal
morphology follows Kuřavová et al. (2017).
SKEJO ET AL.
6 · Zootaxa 5524 (1) © 2024 Magnolia Press
Measurements. Measurements were taken in ImageJ software after calibration with the scale in the each image.
Measurements were taken mostly following Tumbrinck (2014):
Body length—from the tip of the vertex to the apex of the pronotum;
Antenna length—from the base of the scapus to the tip of the apical antennomere;
Eye width—in frontal view, widest compound eye width;
Vertex width—in frontal view, vertex width between the eyes;
Pronotum length—in dorsal view, length between the anterior margin and the tip of the pronotum;
Pronotum width— in dorsal view, between the shoulders;
Tegmen length—for species with tegmina, in lateral view, length between the base and the tip;
Tegmen width—for species with tegmina, in lateral view, width of visible part of tegmen in its widest part;
Fore femur length—from the base to the tip in lateral view;
Fore femur width—widest mid part of the fore femur;
Mid femur length—from the base to the tip in lateral view;
Mid femur width—widest mid part of the mid femur;
Hind femur length—from the base to the tip in lateral view;
Hind femur width—widest part of the hind femur;
Overview of all the localities. List of all the localities mentioned in literature and in our research, with accompanying
coordinates, is given in Table 1. A thorough literature overview was made in order to identify some localities, such
as Tangke Salokko, a mountain supposedly part of Mengokka Mts. in Sulawesi, but which is not mentioned in
any source except for the G. Heinrich expeditions. Unfortunately, for most of the localities from Sulawesi the
coordinates are approximate, as it is usually not known in detail where G. Heinrich or K. F. Sarasin and P. B. Sarasin
collected certain specimens.
TABLE 1. List of localities mentioned in this study with island annotated, coordinates, elevation, date of collection and
collector’s names.
ISLAND Locality Coordinates Elevation Date Collector
MINDANAO: Mt. Kitanglad, Cinchona Natural Park
Forest Reserve, Kaatuan, Lantapan, Bukidnon
8.10N, 124.93E 1 500 m 7.–8.VII.2019 R. Jr. R Patano
MINDANAO: Mt. Pantaron, Barangay Mandahican,
Sitio Miaray, Cabanglasan, Bukidnon
8.09N, 125.43E 1 004 m 9.–12.II.2020 R. Jr. R Patano
MINDANAO: Mt. Apo, Barangay Mahongkog, Sitio
V, Magpet, North Cotabato
7.1898N, 125.1648E 1 257 m 25.I.2021. R. Jr. R Patano
MINDANAO: Mt. Malimumu (Pantaron Range, Sitio
Nabangkal, Barangay Magkalungay, San Fernando,
Bukidnon)
7.8807N, 125.409E 1 000 m
19.–21.VIII.2020
R. Jr. R Patano
MINDANAO: Mt. Malambo (Barangay Datu Salumay,
Marilog Forest Reserve, Marilog District, Davao)
7.483N, 125.25E 1 175 m 23.–24.II.2018,
26.–27.VII.2019
R. Jr. R Patano
MINDANAO: Mt. Agad-Agad (Barangay Tipanoy,
Sitio Pindugangan, Iligan, Lanao del Norte)
8.209N, 124.271E 305 m 28.-30.XI.2020 R. Jr. R Patano
MINDANAO: Lanao del Norte: Iligan 8.21N, 124.26E 120 m Unknown
(before 1938)
W. Schultze
MINDANAO: Lanao del Norte: Mumungan 8.1164,124.225E 350 m Unknown
(before 1938)
W. Schultze
MINDANAO: Mount Balatukan (Civoleg, Gingoog,
Misamis Oriental)
8.77N, 124.98E 1 300 m 18.–20.I.2023. R. Jr. R Patano
MINDANAO: Mount Natampod (Barangay Namnam,
San Fernando, Bukidnon)
7.86N, 125.42E 900–980 m 24.–25.VII.2022. R. Jr. R Patano
......continued on the next page
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 7
TABLE 1. (Continued)
ISLAND Locality Coordinates Elevation Date Collector
MINDANAO: Eureka, Talangisog, Gingoog City
(Misamis Oriental)
8.6N,125.24E 652 m 20.–21.VII.2023 R. Jr. R Patano
MINDANAO: Bukidnon: Lantapan 8.0017N, 125.0105E 650 m 21.III.2022. obs. Evedreine
Mingo
MINDANAO: Lanao del Norte: Tagoloan
8.12122N,
124.285639E
390 m 20.XI.2021. obs. Mark
Gregory Rule.
MINDANAO: Surigao del Sur: Bislig 8.214N, 126.237E 220 m 16.III.2011. obs. Shirley
Sekarajasingham
MINDANAO: Zamboanga del Norte: Sandayong 7.96N, 122.71E 200 m VIII.2020. unknown
MINDANAO: Zamboanga del Sur: Lapaz
Experimental Forestry
7.046N, 122.0147E 920 m 31.I.2020. iNaturalist user
@anncabras24
SULAWESI: SE Sulawesi: Tangke Salokko Mt.
(Mekongga Mts.)
3.67S, 121.18E 1 500 m 1.–15.I.1932. G. Heinrich
SULAWESI: Mt. Tambusisi (Sulawesi Tengah) 1.656S, 121.373E 1 200 m 3.-13.IV.1980. M. Brendell
SULAWESI: Toli Toli Mts. (Northern Sulawesi) 0.75N, 120.74E 400-800 m 11.XII.1895. Fruhstorfer
SULAWESI: Ile-Ile Mts. (northern Sulawesi) 0.967N, 121.80E 500 m 11.XII.1930. G. Heinrich
SULAWESI: Loka, Piek van Bonthain
(= Lompobattang) (SW Sulawesi)
5.34S, 119.93E >1000 m X.1895. drs. Sarasin
SULAWESI: 20 km NE Pale, ca. 6km W Tawaeli
(C Sulawesi)
0.729S, 119.933E 250 m 02.III.2009 A. Skale
SULAWESI: Palu district, Palolo env. (CW Sulawesi) 1.132S, 120.131E 700 m V.2017. unknown
Imaging of specimens and scanning electron microscopy (SEM) of the subapical antennomeres. Specimens
were photographed by Eileen Ngyuen at the LIB, Hamburg using a custom-made stacking system (DUN Inc.,
California, USA). Images were either taken with a Canon EOS 5DSR with a 65 mm or 100 mm lens and stacked
with Zerene Stacker (PMax algorithm) or with a Canon EOS 6D with a 55 mm or 65 mm lens and stacked with
Helicon Focus 5.3.
Further, we performed SEM analyses for two species with widened antennae: Zvierckia storozhenkoi sp. nov.
belongs to Scelimeninae, Discotettigini, while Phaesticus mellerborgi does not have tribal or subfamily placement.
A male specimen of Zvierckia storozhenkoi sp. nov. (ZMH 831024) originating from Palopo Palu, Sulawesi
(Indonesia) collected on I. 2017, and a female specimen of Phaesticus mellerborgi (ZMH 848501) originating from
Lebang Hara (Borneo) collected on 1.–7. 1. 1925 were investigated. The specimens were both examined with a
tabletop SEM (Hitachi TM4000Plus at the LIB, Hamburg). The pinned specimens were placed inside the chamber
using either styrofoam or dental wax to fix the pin to an aluminum label in a way that the antennal segments
of interest were horizontal. The antennae were observed and photographed using the following settings: 15 kV,
150-9000x magnification, WD 14.1-18.8 mm, LensMode 3. Because both antennae of individual specimens are
symmetrical, only the left antennae were looked at both from ventral and dorsal view. The structures of the antennae
were measured in ImageJ (Bourne 2010). The terminology of the antennal sensillae follows Bland (1991) and
Kuravova et al. (2018).
Structure of the study. The genus Hirrius Bolívar, 1887 is herewith divided into four genera belonging to three
distinct evolutionary lineages: Hirrius Bolívar, 1887 from Mindanao and Guentheracris gen. nov. from Sulawesi
belong to the Melainotettix genus group nov., together with Arulenus Stål, 1877 from Mindanao, Melainotettix
Günther, 1939 and Camelotettix Hancock, 1907 from Papua; Parahirrius gen. nov. is a genus of uncertain placement
(incertae sedis) within Tetrigidae; and finally, Zvierckia gen. nov. belongs to tribe Discotettigini of the subfamily
Scelimeninae. The study provides an annotated diagnosis of Hirrius and descriptions of all other genera; a key to
the species of each genus; a catalogue of included species with annotated diagnoses and/or descriptions (for new
taxa); and finally, a discussion on the steps that need to be undertaken to contextualize the identified groups, as well
as insights into the morphology of widened antennal segments.
SKEJO ET AL.
8 · Zootaxa 5524 (1) © 2024 Magnolia Press
Results
Taxonomic part
Family Tetrigidae Rambur, 1838
A key to genera hitherto assigned to Hirrius
1A) Antennae with widened segments. ........................................................................2
1B) Antennae filiform. (Apterous. Vertex as wide as an eye or narrower. In Mindanao.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
............................................. Parahirrius Skejo, Patano et Kasalo gen. nov. (2 species, both new)
2A) Antennae with smooth margins ..........................................................................3
2B) Antennae with toothed margins. (In Sulawesi) . . . Zvierckia Skejo, Tumbrinck et Pushkar gen. nov. (3 species, of which 1 new)
3A) Pronotum flat. Vertex as wide as an eye. (In Mindanao) . . . . . . . . . . . . . . . . Hirrius Bolívar, 1887 (3 species, of which 1 new)
3B) Pronotum with a hump. Vertex wider than an eye. (In Sulawesi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
............................................... Guentheracris Skejo, Tumbrinck et Pushkar gen. nov. (1 species)
Subfamily Metrodorinae Bolívar, 1887
(Tentative placement, as this subfamily needs comprehensive revision.)
Genus group Melainotettix nov.
Diagnosis. Body usually black. In frontal view, vertex with C-shaped carinae. Dorsal margins of the antennal
grooves positioned at the lower third of the compound eyes. In dorsal view, vertex slightly indrawn. Pronotal discus
wavy, with several depressions. Lateral lobes of the pronotum projected sidewards. Prozonal carinae present or
reduced. Legs smooth and elongated, without tubercles or teeth. Hind femur with a concavity just before the hind
knee.
Composition and distribution. Two genera endemic to Mindanao (Hirrius Bolívar, 1887 and Arulenus Stål,
1877), one monotypic genus endemic to northern Sulawesi (Guentheracris gen. nov., described here) and two
genera endemic to Papua (Camelotettix Hancock, 1907 and Melainotettix Günther, 1939). Possibly, Parahirrius
gen. nov. also belongs to this group.
Genus Hirrius Bolívar, 1887
Vernacular name: Mindanao Blackhoppers (Figs. 1–13)
Historical mentions: under Arulenus Stål 1877: 55–56; As Hirrius Bolívar 1887: 191, 196, 308; Hancock 1907:
7; Kirby 1910: 3; Bolívar 1931: 30; Günther 1937: 176; 1938: 303; Blackith 1992: 90; Yin et al. 1996: 875; Otte
1997: 33; Skejo 2017: 29, 48, 117–121; Patano et al. 2022: 776.
Type species Arulenus punctatus Stål, 1877 (= Hirrius punctatus), by original monotypy.
Composition and distribution. Three species endemic to Mindanao, the Philippines (Fig. 2A, 2B, 2C): Hirrius
punctatus (Stål, 1877), H. mindanaensis Günther, 1938, and Hirrius ruber sp. nov.
Diagnosis. The genus Hirrius shares similarities with genera Guentheracris gen. nov., Parahirrius gen. nov.
and Zvierckia gen. nov., that were assigned to the genus Hirrius hitherto, but can easily be distinguished from these
genera, as well as from other similar Tetrigidae genera, such as Melainotettix and Camelotettix, by the following
set of traits: (1) Antennae smooth, long and with almost all the segments widened (filiform and short in Parahirrius
gen. nov., Melainotettix and Camelotettix; only subapical antennomeres widened in Guentheracris gen. nov.;
antennae with toothed margins and widened subapical antennomeres in Zvierckia gen. nov.); (2) Antennal grooves
inserted between the eyes, in similar fashion as in Guentheracris gen. nov., Melainotettix and Camelotettix (at the
level of the lower margins of the compound eyes or slightly below in Parahirrius gen. nov., while strongly below
the lower margins of the compound eyes in Zvierckia gen. nov.); (3) pronotal discus in lateral view flat with a barely
perceptible hump after prozona (with weak projections/tubercles in Zvierckia gen. nov.; undulated in Guentheracris
gen. nov., Melainotettix and Camelotettix); (4) tegmina not visible (visible in Zvierckia gen. nov., Melainotettix and
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 9
Camelotettix); (5) hindwings visible in some species (visible in Zvierckia gen. nov. and Guentheracris gen. nov.,
while large and easily visible in Melainotettix and Camelotettix, not visible at all in Parahirrius gen. nov.). The
genus Arulenus Stål, 1877 from Mindanao with widened subapical antennal segments can easily be distinguished
from all the mentioned genera by the high spines between the shoulders.
FIGURE 2. Distribution of the species belonging to the genera Hirrius and Parahirrius gen. nov., both endemic to the island
of Mindanao. A) H. punctatus, B) H. mindanaensis, C) H. ruber sp. nov., D) P. parvus sp. nov., E) P. amorosus sp. nov.
SKEJO ET AL.
10 · Zootaxa 5524 (1) © 2024 Magnolia Press
A key to species of Hirrius
1A) Hindwings visible ....................................................................................2
1B) Hindwings not visible. (Pronotum black with characteristic yellow or red lines and dots) . . . . . . . . . . . . . . Hirrius punctatus
2A) Ventral side of the abdomen black. Pronotum black or dark brown with light markings. Prozonal carinae distinct. Hind femora
robust .............................................................................Hirrius mindanaensis
2B) Ventral side of the abdomen red. Pronotum uniformly black. Prozonal carinae indistinct. Hind femora elongated ..........
..................................................................................Hirrius ruber sp. nov.
List of Hirrius species
Hirrius punctatus (Stål, 1877)
Vernacular name: Spotted Mindanao Blackhopper (Figs. 1, 3, 4, 5, 6)
Historical mentions: As Arulenus punctatus Stål 1877: 56; Casto de Elera 1895: 207; As Hirrius punctatus
Bolívar 1887: 309; Hancock 1907: 7; Kirby 1910: 3; Bruner 1915: 241; Günther 1938: 303, 424; Blackith 1992: 90;
Skejo 2017: 118–121; Patano et al. 2022: 776.
FIGURE 3. Lectotype ♀ Hirrius punctatus (Stål, 1877) from NHRS. A) dorsal view, B) left lateral view, C) right lateral view,
D) labels, E) head in frontal view. Photo J. Tumbrinck.
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Lectotypization. An open taxonomic issue occurred, which we and the researchers before us were unaware
of. Arulenus punctatus Stål 1877 type series consists of two syntypes, a female and a male, belonging to two
species in two genera. Because of this problem, here we designate the female specimen from Naturhistoriska
Riksmuseet Stockholm, Sweden catalogue number NRM-ORTH 0012904 as the lectotype of Arulenus punctatus
and single name-bearing type. By this act, we fix the name Hirrius punctatus to the apterous species with widened
antennomeres, pronotum black and ornamented in yellow or red dots and lines, endemic to Mindanao. The male
specimen, cotype of A. punctatus at MfN Berlin, is considered here as paratype of endemic to Mindanao Parahirrius
parvus sp. nov., a small species with short filiform antennae (see below).
Type specimens. 1♀ lectotype ‘Ins. Phillipp.’, ‘Semper’, ‘Arulenus punctatus Stål’, ‘Typus’, ‘NRM-ORTH
0012904) (NHRS); here designated.
Additional records
Museum records:
1♂ ‘231’, ‘O. Philippinen C. Semper Coll 231.’, ‘Hirrius punctatus Stål K. Günther det.’ ‘ZMH 831020’ (ZMH);
FIGURE 4. Hirrius punctatus (Stål, 1877) ♂ from Semper’s collection, i.e., from the same collector and probably the collecting
event as the lectotype ♀ from fig. 3. Deposited in ZMH. A) dorsal view, B) left lateral view, C) labels, D) head in frontal view.
Photo E. Nguyen.
SKEJO ET AL.
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1♀, 1♂ THE PHILIPPINES: Davao: Marilog Forest Reserve 1175 m a. s. l. 23.–24.II.2018.; 26.–27.VII.2019. leg.
et obs. R. Jr. R Patano (CMU);
1♀, 2♂ THE PHILIPPINES: Davao: Mount Hamiguitan, Malambo 1175 m a. s. l. 23.–24.II.2018., 26.–27.VII.2019.
leg. et obs. R. Jr. R Patano (CMU);
2♂ THE PHILIPPINES: Bukidnon: Kitanglad 1500 m a. s. l. 7.–8.VII.2019. leg. et obs. R. Jr. R Patano (CMU);
1♀ THE PHILIPPINES: North Cotabato: Mount Apo 1257 m a. s. l. 25.I.2021. leg. et obs. R. Jr. R Patano (CMU);
2♀♀, 1♂ THE PHILIPPINES: Bukidnon: Pantaron Range about 1000 m a. s. l. 9.–12.II.2020., 19.–21. VIII.2020.
leg. et obs. R. Jr. R Patano (CMU);
1♀ THE PHILIPPINES: Misamis Oriental: Eureka, Talangisog 652 m a. s. l. 20.–21.VII.2023 leg. et obs. R. Jr. R
Patano (CMU);
FIGURE 5. Hirrius punctatus (Stål, 1877), first photograph of a living specimen, ♀ from the Philippines, Mindanao, taken on
16.III.2011. by Shirley Sekarajasingham and reproduced with author’s permission. A) dorsolateral view, B) dorsal view.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 13
Online social media records:
1♀ THE PHILIPPINES: Mindanao: Surigao del Sur: Bislig 16.III.2011. obs. Shirley Sekarajasingham, available
at link https://www.flickr.com/photos/grandma-shirley/6145150892 (and https://www.flickr.com/photos/grandma-
shirley/6144599435);
1♀ THE PHILIPPINES: Mindanao: Lanao del Sur: Kapai (8.12122, 124.285639) iNaturalist observation number
112726760 20.XI.2021. obs. Mark Gregory Rule.
1♀ THE PHILIPPINES: Mindanao: Bukidnon: Lantapan (8.0017, 123.0105) iNaturalist observation number
116808954 21.III.2022. obs. Evedreine Mingo;
Diagnosis. Hirrius punctatus is similar to its congeners H. mindanaensis and H. ruber sp. nov., but can easily
be distinguished from them by the following set of traits: (1) alae not visible under the pronotum (visible in H.
mindanaensis and H. ruber sp. nov.), (2) prozonal carinae indistinct (distinct in H. mindanaensis), (3) pronotum
black and richly ornamented with yellow and red dots and lines (variable in H. mindanaensis, from fully black
to brown with pale markings; black in H. ruber sp. nov.), (4) hind femora slender (robust in H. mindanaensis),
(5) abdominal sternites black (bright red in H. ruber sp. nov.), (6) slightly larger than H. ruber sp. nov. (see
measurements).
Color variation. The species does not have large color variability. Antennae are fully black. Pronotum almost
always has two bright and wide red, yellow, or orange dots between the humeral angles (sometimes fused), and two
smaller and usually paler dots in the middle of the metazona (also sometimes fused). From the level of these dots
to the tip of the pronotum, lateral carinae are usually brightly colored yellow or orange. Fore and mid femora and
tibiae usually have yellow-orange lines on the dorsal margins. Hind femora black with various yellow dots or lines
on the outer surface and dorsal margins.
Measurements. The measurements of the holotype and five additional specimens from the recent fieldwork
are shown in Table 2.
TABLE 2. Measurements of Hirrius punctatus lectotype female, and four additional females and one male from (1)
Mount Malimumu, (2) Cabanglasan within Pantaron range, (3) Marilog Forest Reserve, (4) Mount Apo, North Cotabato,
(5) Eureka, Talangisog and (6) Mount Kitanglad within Cinchona Forest (Mindanao, the Philippines). All measurements
are given in millimeters.
Lectotype
♀
(1) Mt. Malimumu
(Pantaron)
♀
(2) Cabanglasan
(Pantaron)
♀
(3) Marilog
Forest Reserve
♀
(4) Mt. Apo
♀
(5) Eureka
(Talangisog)
♀
(6) Mt. Kitanglad,
(Cinchona)
♂
Body length 12.10 13.10 11.30 12.00 12.15 12.50 11.20
Antenna length 6.30 6.50 8.00 7.20 6.50 7.0 6.50
Eye width 0.75 0.70 0.80 0.80 0.85 0.75 0.70
Vertex width 0.80 0.70 0.70 0.80 0.75 0.75 0.60
Pronotum length 10.80 9.70 9.60 9.40 9.30 9.50 8.20
Pronotum width
(at shoulders) 3.85 3.95 3.90 3.85 4.00 3.9 3.35
Fore femur length 3.50 3.20 4.00 3.60 2.90 3.45 2.70
Fore femur width 0.70 0.75 0.80 0.60 0.60 0.65 0.60
Mid femur length 3.40 2.90 4.10 3.40 3.20 3.50 2.50
Mid femur width 0.60 0.80 0.90 0.60 0.75 0.70 0.75
Hind femur length 6.25 6.00 6.20 5.40 6.00 6.00 5.50
Hind femur width 2.05 2.10 2.50 2.50 2.20 2.40 2.10
Distribution and habitat. The species seems to be common in various types of rainforests of Mindanao (see
‘Additional records’), the Philippines (Fig. 2A), from the lowlands to the mountainous rainforests, where it can be
observed on leaves and bark.
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FIGURE 6. Color variability of Hirrius punctatus (Stål, 1877) from Mindanao. A) male similar in coloration to the lectotype,
B) dark ♂ with many bright markings and epizoic overgrowth, C) dark ♀ with very weak markings, D) ♂ of a very bright
coloration and fused dorsal spots, E) dark ♂ with bright markings, F) dark ♂ with very bright markings. Photo R. Patano Jr.
Hirrius mindanaensis Günther, 1938
Vernacular name: Western Mindanao Blackhopper (Figs 7, 8, 9, 10, 11)
Historical mentions: As Hirrius mindanaensis Günther 1938: 304; Petersen & Gaedike 1970: 166; Blackith 1992:
90; Skejo 2017: 121.
Type specimens. 2♀♀ syntypes and 1♂ syntype with same labels: ‘Mindanao W. Schultze’ ‘Iligan’ ‘Lanao’
‘Syntypus’ ‘Hirrius mindanaensis n. sp. K. Günther det.’ (SDEI) (Günther 1938).
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 15
FIGURE 7. Hirrius mindanaensis Günther, 1938 syntypes from SDEI (A–D ♀ and E–I ♂). A) ♀ left lateral view, B) ♀ dorsal
view, C) ♀ head in frontal view, D) ♀ syntype labels, E) ♂ left lateral view, F) ♂ right lateral view, G) ♂ dorsal view, H) ♂ head
in frontal view, I) ♂ syntype labels. Photo S. Ingrisch.
SKEJO ET AL.
16 · Zootaxa 5524 (1) © 2024 Magnolia Press
Published records: 2♀♀ the Philippines: Mindanao Isl.: Lanao del Norte: Mumungan, Collector: W. Schultze,
det. K. K. Günther (SMTD) (Günther 1938); 1♀ Philippines: Mindanao: Zamboanga del Norte: Sandayong,
VIII.2020. det. 2022 J. Tumbrinck (CJT) (Cigliano et al. 2023).
Additional records.
Museum records:
1♀+1♂ Philippines: Mindanao: Zamboanga del Norte: Sandayong, VIII.2020. det. 2022 J. Tumbrinck (ZMH);
1♀+1♂ Philippines: Mt. Agad-Agad (Barangay Tipanoy, Sitio Pendugangan, Iligan, Lanao del Norte) 305 m a. s. l.
28.-30.XI.2020. leg. R. Patano Jr. (CMU).
Online social media records:
1♂ Philippines: Zamboanga del Sur: Lapaz Experimental Forestry 920 m a. s. l. observed on 31.I.2020. by iNaturalist
user @anncabras24, available at https://www.inaturalist.org/observations/38141140.
FIGURE 8. Hirrius mindanaensis Günther, 1938 ♀ from Zamboanga peninsula. A) head in frontal view, B) dorsal view, C)
right lateral view, D) labels, E) left dorsolateral view. Photo J. Tumbrinck.
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FIGURE 9. Hirrius mindanaensis Günther, 1938 ♀ from Zamboanga peninsula. A) dorsal view, B) left lateral view, C) labels,
D head in frontal view. Photo E. Nguyen.
SKEJO ET AL.
18 · Zootaxa 5524 (1) © 2024 Magnolia Press
FIGURE 10. Hirrius mindanaensis Günther, 1938 ♂ from Zamboanga peninsula. A) dorsal view, B) left lateral view, C) head
in frontal view, D) labels. Photo E. Nguyen.
Diagnosis. Hirrius mindanaensis shares characters with its congeners, but there is a set of characters by which
it can be easily distinguished from H. punctatus and H. ruber sp. nov.: (1) alae are visible under the pronotum in
lateral view (also visible in H. ruber sp. nov., but not in H. punctatus); (2) prozonal carinae are distinct, while
they are literally absent in H. punctatus and H. ruber sp. nov.; (3) hind femora visibly robust (much slenderer in
H. punctatus and H. ruber sp. nov.); (4) abdominal sternites dark, similar to H. punctatus, while H. ruber sp. nov.
has bright red sternites; (5) slightly larger than H. ruber sp. nov. (see measurements); (6) pronotum of variable
coloration, but generally with much paler ornamentation than in H. punctatus and H. ruber sp. nov., which both
have bright red and yellow markings.
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Syntype measurements (female). Body length from tip of head to the tip of pronotum 10.85 mm; eye width
0.70 mm, vertex width 0.95 mm; pronotum length 10.05 mm; pronotum width (at shoulders) 3.40 mm; fore femur
length 2.80 mm; fore femur width 0.70 mm; mid femur length 2.85; mid femur width 0.70; hind femur length 6.10
mm; hind femur width 2.45 mm.
Syntype measurements (male). Body length from tip of head to the tip of pronotum 8.90 mm; eye width 0.65
mm; vertex width 0.80 mm; pronotum length 7.80 mm; pronotum width (at shoulders) 2.80 mm; fore femur length
2.25 mm; fore femur width 0.50 mm; mid femur length 2.35 mm; mid femur width 0.60 mm; hind femur length 4.80
mm; hind femur width 2.10 mm.
Distribution and habitat. Western Mindanao Blackhopper inhabits lowland and mountainous tropical
rainforests of Mindanao, where it can be found from 100 m a. s. l. up to 950 m a. s. l. in the regions around the lake
of Lanao (e.g., Lanao del Norte) and on the Zamboanga Peninsula (in e.g., Zamboanga del Norte and Zamboanga
del Sur) (Fig. 2B).
FIGURE 11. Hirrius mindanaensis Günther, 1938 A–B) living specimen from Iligan, photo R. Patano Jr., C) living ♂ from
Zamboanga del Sur observed by iNaturalist user @anncabras24 on on 31.I.2020., iNaturalist observation 38141140.
SKEJO ET AL.
20 · Zootaxa 5524 (1) © 2024 Magnolia Press
Hirrius ruber Skejo, Patano et Kasalo, sp. nov.
Zoobank ID: urn:lsid:zoobank.org:act:6A108D3E-9F87-4CC2-9C66-C3BFA89F7766
Vernacular name: Red Mindanao Blackhopper (Figs. 12, 13)
Type specimens: 1♀ holotype THE PHILIPPINES: Mindanao Island: Bukidnon: San Fernando: Mount Natampod
905 m a. s. l. 24.VII.2022. leg. Romeo Patano R. Jr. (CMU); 1♂ paratype THE PHILIPPINES: Mindanao Island:
Bukidnon: San Fernando: Mount Natampod 905 m a. s. l. 24.–25.VII.2022. leg. Romeo Patano R. Jr. (CMU).
Type locality. THE PHILIPPINES: Mindanao Island: Bukidnon: San Fernando: Mount Natampod rainforest
905 m a. s. l. (7.86N, 125.42E), Pantaron Range.
Type depository. Holotype and paratype are deposited in the Central Mindanao University (CMU) Orthoptera
Collection, Mindanao, Philippines
Etymology. From Latin ‘ruber, rubra’ meaning ‘red’, referring to bright red abdomen coloration. Species
epitheton is of masculine grammar gender, fitting the genus gender.
FIGURE 12. Hirrius ruber Skejo, Patano et Kasalo sp. nov., living type specimens at the type locality, Mount Natampod
24.VII.2022., before being caught. A) ♀ holotype right lateral view, B) ♂ paratype left lateral view, C) same as previous one, D)
♀ in left dorsoventral view. Photo R. Patano Jr.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 21
Diagnosis. New species shares similarities with its congeners, H. punctatus and H. mindanaensis, but can
easily be distinguished from them by the following set of traits: (1) alae visible under the pronotum (not visible
in H. punctatus), (2) prozonal carinae indistinct (distinct in H. mindanaensis), (3) pronotum uniformly black in
color (richly ornamented in yellow and red dots and lines in H. punctatus, while variable in H. mindanaensis, from
fully black to brown with pale markings), (4) hind femora slender (robust in H. mindanaensis), (5) abdominal
sternites bright red in color (black in H. punctatus and H. mindanaensis), (6) slightly smaller than H. punctatus (see
measurements).
FIGURE 13. Hirrius ruber Skejo, Patano et Kasalo sp. nov., ♀ holotype (A–G) and ♂ paratype (H). A) ♀ holotype in dorsal
view, B) ♀ holotype in ventral view, C) ♀ holotype in right lateral view, D) ♀ holotype head in frontal view, E) ♀ holotype head
and pronotum details in lateral view, F) ♀ holotype head and pronotum details in dorsal view, G) ♀ holotype abdomen apex in
lateral view, H) ♂ paratype abdomen apex in lateral view. Photo R. Patano Jr.
SKEJO ET AL.
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Description. Relatively small species with smooth head, pronotum, and legs. All structures colored uniformly
black except for a pair of bright yellow lines in the occipital area of the head; yellow maxillary palps; a pair of bright
yellow dots on the basal fifth of the hind femora; a pair of narrow bright yellow lines on the margin of paranota; and
abdomen with bright red ventral surface (sternites).
Antennae. Antennae with 15 black antennomeres and with all the antennomeres flattened and widened (except
for the reduced apical segments): 1st massive scapus; 2nd large pedicel; 3rd to 6th basal antennomeres, robust and
shorter than the central segments; 7th and 8th central antennomeres, elongated and widened; 9th antennomere very long
and widened, pennate; 10th the longest and the widest, the most pennate one; 11th segment widened and elongated;
12th to 15th apical segment fused and small.
Head. Eyes in frontal view globose, the dorsal margins elevated above the level of the fastigium. Vertex in
the level of pronotum in lateral view. Frons and vertex forming 60 degrees angle. In frontal view, vertex of almost
the same width as a compound eye. In lateral view, vertex and frons not visible because of the protruding eyes.
In dorsal view, vertex oblique with middle indrawn/excised into a wide V-shape, not projecting before the eyes.
Lateral carinae of the vertex not elevated (above the eyes). In dorsal view, tip of the vertex slightly narrower than
a compound. Medial carina of the vertex present, recognizable in the apical fourth. Fossulae deep. Lateral and
transverse carinae forming V-shape in dorsal view and U-shape in frontal view. Frontal costa short, recognizable
above the lateral ocelli in frontal view. Whole vertex is lowered in the anterior part, so the frontal costa might be
in fact the upper part of the medial carina of the vertex. Frontal costa bifurcation situated in the upper fourth of the
compound eyes height. Frontal costa in lateral view not projected forwards. Dorsal margin of the antennal groove
placed between the compound eyes. Lateral ocelli placed in upper third of the compound eyes height. Maxillary
palps yellow, smooth. Occipital area as wide as a half of a compound eye.
Pronotum. Pronotum covering whole abdomen. Pronotum surface very smooth, finely granulated. Anterior
margin of the pronotum truncated. Median carina continuous, except in the anterior part where it is intercepted
by two sulci. Generally, median carina of pronotum is low and hardly recognizable. Pronotal discus in lateral
view flat, plain. Prozonal carinae almost indistinct, short and convergent. Interhumeral carinae not recognizable.
Humeroapical carina weak. Tegminal sinus absent. Ventral sinus deep. Infrascapular area narrow and with parallel
margins. Lateral area very narrow. Lateral lobes directed sidewards, without projections. Apex of the lateral lobe
narrowly truncated. Pronotal tip acute, reaching hind knees.
Wings. Tegmina absent or reduced and covered by infrascapular area. Hindwings visible, dark with red tips,
shorter than the tip of pronotum for 0.5 mm.
Legs. Fore and mid femora elongated, smooth, and finely granulated. Dorsal and ventral carinae of the fore and
mid legs straight, finely granulated. Hind femur slender, about 3 times as long as wide. Five strong transverse ridges
recognizable in the middle outer area. Dorsal carina of the hind femur smooth and straight Antegenicular teeth small
and blunt, genicular teeth long and sharp. Hind tibia with blunt spines. First tarsal segment shorter than third tarsal
puvilli obliquely angular. Proximal pulvillus smallest, middle a bit larger, while the distal largest.
Ovipositor. Ovipositor elongated and with strongly armed valvulae. Upper valvulae wider in lateral view than
the lower ones. Also, upper valvulae have more teeth (>10) than the lower ones (8-9). Cerci conical and hairy.
Holotype measurements (female). Body length from tip of head to the tip of pronotum 11.65 mm; antenna
length 6.10 mm; eye width 0.60 mm; vertex width 0.80 mm; pronotum length 9.90 mm; pronotum width (at
shoulders) 3.40 mm; fore femur length 2.80 mm; fore femur width 0.60 mm; mid femur length 3.00 mm; mid femur
width 0.60 mm; hind femur length 6.00 mm; hind femur width 2.00 mm;
Paratype measurements (male). Body length from tip of head to the tip of pronotum 9.95 mm; antenna length
5.80 mm; eye width 0.70 mm; vertex width 0.60 mm; pronotum length 9.25 mm; pronotum width (at shoulders)
3.20 mm; fore femur length 3.00 mm; fore femur width 0.60 mm; mid femur length 3.00 mm; mid femur width 0.60
mm; hind femur length 5.20 mm; hind femur width 1.65 mm.
Distribution and habitat: So far only known from the mountainous tropical rainforests of Mount Natampod,
Pantaron Range, 905 m a. s. l. on the island of Mindanao, the Philippines (Fig. 2C).
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 23
Genus Guentheracris Skejo, Tumbrinck et Pushkar, gen. nov.
Zoobank ID: urn:lsid:zoobank.org:act:F45A1561-E5EC-4ECB-9BB3-E96D5B1830C9
Vernacular name: Ile-Ile Pygmy Devils (Figs. 1, 14–15, 22)
Historical mentions: Under Hirrius Günther 1937: 177; 1938: 303; Blackith 1992; Under Zvierckia Skejo 2017:
193; unavailable according to Article 13.1 of the Code (ICZN, 1999).
Type species Hirrius scrobiculatus Günther, 1937, here designated.
Etymology. Named after Klaus Günther, a famous German entomologist, zoologist, anthropologist, and
philologist. Günther is one of the fathers of modern tetrigidology, who described three tribes, 43 genera (38 valid
today), and 246 species (226 valid today). The generic name is composed of Guenther and acris, Latinized Greek
word (ἀκρίς) for a grasshopper, and is of feminine gender (because of ἡ ᾰκρῐς, τῆς ᾰκρῐδος).
Composition and distribution. Monotypic, endemic to northern Sulawesi (Ile-Ile Mts.) (Fig. 22D).
Diagnosis. New genus may be distinguished from Hirrius from Mindanao by the wide vertex, by the shape of
the antennae, and by the wavy pronotal discus; from Parahirrius gen. nov. it distinguished by the widened antennal
segments, by the wider vertex, and by the presence of the alae; it can also be easily separated from Zvierckia gen.
nov. from Sulawesi by the higher position of the antennal grooves, different shape of the widened segments of the
antennae (lacking saw-like margins in the new genus). The new genus is assigned to the proposed here Melainotettix
genus group because of the shape of the head, especially in frontal view, which is very similar to Melainotettix and
Camelotettix, and because of the wavy pronotum and elongated legs, also shared among Melainotettix genus group
members. New genus may be easily separated from Melainotettix and Camelotettix, Guentheracris gen. nov. by the
covered tegmina, by the short pronotum, and the very wide vertex.
Description. Genus with smooth, black, and moderately large and slender species of wavy pronotum and with
tegmina fully covered by pronotum.
Antennae. Long, with widened subapical segments. With 15 antennomeres, smooth. 1st massive scapus, 2nd
large pedicel, 3rd to 8th basal segments, elongated. 9th antennomere elongated and widened; 10th antennomere strongly
widened. 11th antennomere small, but widened. Antennomeres 12th to 15th reduced and fused.
Head. Eyes in frontal view elliptical. Vertex in the level of pronotum. In frontal view, vertex almost plain, very
slightly elevated in the middle. Vertex not visible in lateral view. In dorsal view, vertex slightly indrawn. Lateral
carinae of the vertex weakly elevated. Vertex two times as wide as an eye. Anterior margin of the vertex truncated,
without convexity. Medial carina of the vertex visible in the upper third only. Fossulae deep. Lateral and transverse
carinae forming wide C-shape. Frontal costa long, bifurcating just between the compound eyes. Frontal costa weakly
produced in lateral view. Dorsal margins of the antennal grooves situated between the lower third of the compound
eyes. Lateral ocelli situated slightly below the frontal costa bifurcation, in the lower half of the compound eyes
height. Maxillary palps small. Occipital area visible.
Pronotum. Pronotum covering whole abdomen. Pronotum smooth, finely granulated. Median carina of the
pronotum continuous, except in the connection of prozona and metazona, where it is indistinct in dorsal view.
Pronotal discus undulated, wavy, with two large depression, first depression just above the tegmina and second
just before the pronotal apex. Prozonal and interhumeral carinae indistinct. Humeroapical carina very weak. The
only distinct carinae are median and internal lateral one. Tegminal sinus almost nonexistent, ventral sinus deep.
Infrascapular area narrow and decurved. Pronotum without any projections. Lateral lobes projected sidewards, with
truncated apex.
Wings. Tegmina almost fully covered by pronotum. Hindwings visible, almost reaching the tip of the
pronotum.
Legs. Fore and mid femora smooth, with almost straight carinae, and without teeth. Hind femur with 7 transverse
ridges on the outer surface. Dorsal carina of the hind femur finely granulated and with a convexity just before the
antigenicular tooth. Antegenicular and genicular teeth short and sharp. Hind tibia margins finely serrated and with
5 larger spines. First and third tarsal segment almost equal in length. First two tarsal pulvilli smaller than the third,
oblique, smooth and almost rounded; while the third pulvillus longer and more angular.
Note. Günther (1937) already noted that Guentheracris scrobiculata comb. nov. has many differences from Zvierckia
members (Hirrius montanus and Hirrius sarasinorum sensu Günther 1937), but did not recognize new genus.
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24 · Zootaxa 5524 (1) © 2024 Magnolia Press
Guentheracris scrobiculata (Günther, 1937), comb. nov.
Vernacular name: Ile-Ile Pygmy Devil (Figs. 14, 15)
Historical mentions: As Hirrius scrobiculatus Günther 1937: 177; 1938: 303; Blackith 1992; As Zvierckia
scrobiculata Skejo 2017: 193; unavailable according to Article 13.1 of the Code (ICZN, 1999).
Type specimens. 1♀ holotype ‘Celebes Ile Ile 500 m 11.12.1930 G Heinrich leg.’ ‘Hirrius scrobiculatus K.
Günther’ ‘Typus’ (MfN); 1♂ paratype ‘Celebes Ile Ile 500 m 11.12.1930 G Heinrich leg.’ ‘Hirrius scrobiculatus
K. Günther’ ‘Typus’ (SMTD).
IUCN Red List assessment: Vulnerable B1ab (iii,v) (Hochkirch & Skejo 2019).
FIGURE 14. Guentheracris scrobiculata (Günther, 1937) comb. nov. ♀ holotype from Sulawesi, Ile-Ile. A) dorsal view, B) left
lateral view, C) head in frontal view, D) labels. Photo E. Nguyen.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 25
FIGURE 15. Guentheracris scrobiculata (Günther, 1937) comb. nov. ♂ paratype from from Sulawesi, Ile-Ile,. A) left lateral
view, B) dorsal view, C) head in frontal view, D) labels. Photo S. Ingrisch.
Diagnosis. Antennae long, 15-segmented, 10th segment strongly widened, pennate. 9th and 7th antennomeres
with wide pale colored rings. Vertex wide, in females two times wider than an eye, in males 1.65 times wider than
an eye. In frontal view, vertex flat; frontal costa bifurcating between the eyes; antennal grooves inserted between
SKEJO ET AL.
26 · Zootaxa 5524 (1) © 2024 Magnolia Press
the eyes: dorsal margins of the antennal grooves between the lower third of the compound eyes height. In lateral
view, the disc of the pronotum wavy, with two deep depressions: one just behind the shoulders, and the other just
before the apex of the pronotum. Prozonal carinae indistinct. Tegmina almost completely covered by infrascapular
area. Hindwings visible. Femora slender, genicular and antegenicular teeth of the hind femur sharp. General color
black.
Holotype measurements (female). Body length from tip of head to the tip of pronotum 14.75 mm; eye width
0.60 mm, vertex width 1.20 mm; pronotum length 13.60 mm; pronotum width (at shoulders) 3.85 mm; tegmen
length 1.0 mm; tegmen width 0.15 mm; fore femur length 4.10 mm; fore femur width 0.85 mm; mid femur length
4.15 mm; mid femur width 0.80 mm; hind femur length 8.00 mm; hind femur width 2.80 mm.
Paratype measurements (male). Body length from tip of head to the tip of pronotum 11.75 mm; eye width
0.60 mm, vertex width 1.00 mm; pronotum length 11.05 mm; pronotum width (at shoulders) 3.45 mm; tegmen
length 0.70 mm; tegmen width 0.10 mm; fore femur length 3.15 mm; fore femur width 0.60 mm; mid femur length
3.15 mm; mid femur width 0.75 mm; hind femur length 6.45 mm; hind femur width 2.00 mm.
Distribution and habitat. Indonesia, northern Sulawesi, endemic to the mountainous tropical rainforests of
Gunung Ile-Ile Mt. (500 m a.s.l) that is part of the Matinan Mountains.
Notes. This interesting genus and its only species are known only from holotype female and paratype male
collected by Gerd Hermann Heinrich during the expedition to Ile-Ile Mts. in northern Sulawesi in 1930. There have
been no new records during the last 93 years. It is likely that more new species will be found in the future, as well
as that Ile-Ile Pygmy Devil lives in other localities in northern Sulawesi where the rainforest is still present. This
species was described based on two speciments, male and female, recorded as holotype and allotype (Günther,
1937), but without type lables on both speciments. Here we considered female as holotype and male as paratype.
Tetrigidae: Incertae sedis
Genus Parahirrius Skejo, Patano et Kasalo, gen. nov.
Zoobank ID: urn:lsid:zoobank.org:act:64064F4C-D428-4469-BB64-6D5603CACE8B
Vernacular name: Mindanao Pygmy Vividhoppers (Figs. 2, 16–21)
Type species: Parahirrius parvus Skejo, Patano et Kasalo, sp. nov., here designated.
Etymology: The new genus name is a combination of the words para (Ancient Greek παρά), meaning ‘near’
and ‘Hirrius’, the name of the genus Hirrius Bolívar, 1887 under which this new genus and species were hidden for
more than 145 years. The genus name is of masculine gender.
Composition and distribution: Two species, Parahirrius parvus sp. nov. and Parahirrius amorosus sp. nov.
endemic to Mindanao, the Philippines (Fig. 2D, 2E).
Diagnosis. This new genus can be clearly separated from Hirrius by the filiform antennae, the narrow vertex
with V-shaped carinae, and by the low position of the antennal grooves. From Zvierckia gen. nov., Parahirrius
gen. nov. can easily be distinguished by the lack of wings, lack of pronotal projections, very small size, and narrow
vertex with V-shaped carinae. From Guentheracris gen. nov., the new genus can be separated by the lack of wings,
wide and long infrascapular area, narrow vertex, low position of the antennal grooves, and very small size. There
is no clear affinity between this genus and any of the known Tetrigidae genera, but because of the V-shaped carinae
and long fore and mid proximal tarsal segments, this genus may belong to the Xistra genus group, whereas the
pronotum shape would point to an affinity to the Melainotettix genus group. From other members of the Xistra
genus group and of Melainotettix genus group it can easily be separated by the lack of wings.
Diagnosis. Small, smooth, flightless, and dorsoventrally flattened colorful species from Mindanao, the
Philippines.
Antennae. Antennae filiform, with 17 antennomeres.
Head. Head light brown or yellow. Eyes in frontal view globose. Vertex in the level of pronotum in lateral
view. Frons and vertex forming acute angle. In frontal view, vertex visibly narrower than a compound eye. In
lateral view, vertex and frons not visible. In dorsal view, vertex oblique, not projected before the eyes. Lateral
carinae of the vertex slightly elevated above the eyes as small horns. In dorsal view, vertex narrower, slightly wider,
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 27
or as wide as a compound eye. Anterior margin of the vertex truncated. Medial carina of the vertex present and
recognizable throughout its whole length. Fossulae deep, so medial carina of the vertex and lateral carinae seem
to be strongly elevated. Lateral and transverse carinae forming V shape in frontal and dorsal view. Frontal costa
long and recognizable between the eyes in frontal view. However, the frontal costa is in fact short as whole vertex
is depressed in the anterior part, so the strongly recognizable structure similar to frontal costa is in fact median
carina. Frontal costa bifurcation situated between the compound eyes or in the lower third. Frontal costa in lateral
view not projected forwards. Dorsal margin of the antennal groove placed below the lower margin of the compound
eye. Lateral ocelli placed in the lower fourth of the compound eyes height, just below the frontal costa bifurcation.
Maxillary palps yellow, smooth, elongated. Occipital area as wide as half of a compound eye.
Pronotum. Pronotum color species-specific. Pronotum covering whole abdomen. Pronotum smooth, finely
granulated. Anterior margin of the pronotum very weakly projected forwards, almost truncated. Median carina
continuous. Pronotal discus in lateral view straight, plain. Prozonal carinae weak, parallel. Interhumeral carinae
present or absent. Humeroapical carina weak. Tegminal sinus absent. Ventral sinus deep. Infrascapular area widened,
extended to the tip of the pronotum in lateral view, but still narrow and with parallel margins. Lateral area very
narrow. Lateral lobes directed sidewards, without projections. Apex of the lateral lobe truncated. Pronotal tip acute,
reaching hind knees.
Wings. Tegmina absent or reduced and covered by infrascapular area. Hindwings absent or reduced and covered
by infrascapular area.
Legs. Fore and mid legs elongated, smooth. Dorsal and ventral carinae of the fore and mid legs smooth, finely
granulated, slightly undulated and almost straight. Hind femur of species-specific coloration. Dorsal carina of the
hind femur smooth and straight. Antegenicular teeth small and blunt, genicular teeth long and sharp. Hind tibia with
4-5 large spines on the outer and inner margins and with many minute teeth. First tarsal segment slightly shorter than
the third Tarsal pulvilli oblique or obliquely angular, distal being the largest one. Hind tibia uniformly yellow.
Ovipositor. Ovipositor small and with strongly armed valvulae. Cerci triangular and hairy
Key to species of Parahirrius
1A) Vertex visibly narrower than an eye. Smaller species, body length 8.80–9.00 mm. Pronotum base color brown, paranota and
lateral carinae with bright yellow lines. Females with larger ovipositor, about 0.45 mm long . . . . Parahirrius parvus sp. nov.
1B) Vertex almost as wide as an eye. Larger species, body length 9.20–10.85 mm. Pronotum base color yellowish and with many
dark patches/blotches. Females with tiny ovipositor, about 0.35 mm long .................Parahirrius amorosus sp. nov.
List of Parahirrius species
Parahirrius parvus Skejo, Patano et Kasalo, sp. nov.
Zoobank ID: urn:lsid:zoobank.org:act:981EA1F3-FABA-4AF3-8F79-731D75428362
Vernacular name: Mindanao Tiny Vividhopper (Figs. 16, 17, 18, 19)
Historical mentions: Hitherto ‘hidden’ under one of the syntypes of Arulenus punctatus (mentioned as Arulenus
punctatus by Stål 1877: 56; Casto de Elera 1895: 207; and as Hirrius punctatus (partim, male only) by Bolívar 1887:
309; Hancock 1907: 7; Kirby 1910: 3; Bruner 1915: 241; Günther 1938: 303, 424; Blackith 1992: 90; Skejo 2017:
118–121).
Type specimens: 1♀ holotype THE PHILIPPINES: Mindanao: Misamis Oriental: Gingoog City: Civoleg:
Mount Balatukan 1 300 m a. s. l. 18.–20I.2023. leg. Romeo Jr. R. Patano (CMU); 1 ♂ paratype THE PHILIPPINES:
‘Ins. Phillipp.’ ‘Semper’ ‘Hirrus punctatus cotyp Stål’ (MfN) (paratype of Arulenus punctatus)
Type locality. THE PHILIPPINES: Mindanao: Misamis Oriental: Gingoog City: Civoleg: Mount Balatukan 1
300 m a. s. l.
Type depository. Holotype female is deposited in the Central Mindanao University (CMU) Orthoptera
Collection, Mindanao, Philippines, while paratype male (=paratype of Arulenus punctatus) in the Museum für
Naturkunde, Berlin.
Etymology. Latin parvus, meaning ‘small’. Specific epithet is of masculine gender.
SKEJO ET AL.
28 · Zootaxa 5524 (1) © 2024 Magnolia Press
FIGURE 16. Parahirrius parvus Skejo, Patano et Kasalo sp. nov. ♂ paratype (= paratype of Arulenus punctatus Stål, 1877) A)
dorsal view), B) broken antenna glued on a paper, C) right lateral view, D) head in frontal view, E) labels. Photo E. Nguyen.
Diagnosis. The species is similar to Parahirrius amorosus sp. nov., but can be easily separated from that
species by (1) smaller size, (2) characteristic coloration of pronotum and legs, (3) by larger ovipositor in females,
and (4) by a slightly narrower vertex.
Description. Small, smooth, flightless, and dorsoventrally flattened colorful species.
Antennae. Antennae filiform, short. Antennae with 17 antennomeres; 1st massive scapus; 2nd large pedicel; 3rd
elongated first basal segment, 4th, 5th and 6th basal segments shorter than the third; 7th and 8th central segments,
elongated; 9th central segment strongly elongated, more than 4 times as long as wide; 10th and 11th segments
similar to 9th; 12th and 13th segments shorter than the previous; 14th segment as long as wide; 15th to 17th apical
segment fused and small.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 29
FIGURE 17. Parahirrius parvus Skejo, Patano et Kasalo sp. nov. ♀ holotype, from Mount Balatukan rainforest . A) ventral
view, B) left lateral view, C) dorsal view, D) head and pronotum detail in dorsal view, E) abdominal apex in lateral view, F) head
in frontal view. Photo R. Patano Jr.
Head. Head color light brown, sometimes with darker markings. Eyes in frontal view globose. Vertex in the
level of pronotum in lateral view. Frons and vertex forming 55 degrees angle. In frontal view, vertex much narrower
than a compound eye, in the level of the dorsal margins of the compound eyes. In lateral view, vertex and frons not
visible. In dorsal view, vertex oblique, not projected before the eyes. Lateral carinae of the vertex slightly elevated
above the eyes as small horns. In dorsal view, vertex narrower than a compound eye in males, and slightly wider
than an eye in females. Anterior margin of the vertex truncated. Medial carina of the vertex present, recognizable.
Fossulae deep, so medial carina of the vertex and lateral carinae seem to be strongly elevated. Lateral and transverse
SKEJO ET AL.
30 · Zootaxa 5524 (1) © 2024 Magnolia Press
carinae forming V-shape in frontal and dorsal view. Frontal costa long and recognizable between the eyes in frontal
view. However, the frontal costa is in fact short as whole vertex is depressed in the anterior part, so the strongly
recognizable structure similar to frontal costa is in fact median carina. Frontal costa bifurcation situated just between
the eyes. Frontal costa in lateral view not projected forwards. Dorsal margin of the antennal groove placed below
the lower margin of the compound eye. Lateral ocelli placed in the lower fourth of the compound eyes height, just
below the frontal costa bifurcation. Maxillary palps yellow, smooth, elongated. Occipital area as wide as a half of
a compound eye.
FIGURE 18. Parahirrius parvus sp. nov. living ♀ holotype in natural habitat, from Mount Balatukan rainforest. A) dorsal
view, B) right lateral view. Photo R. Patano Jr.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 31
FIGURE 19. Parahirrius parvus sp. nov. living ♀ holotype in natural habitat standing on mosses (A–F, J–L) or on stem (G–I),
from Mount Balatukan rainforest at 1 300 m. a. s. l. Photo R. Patano Jr.
Pronotum and wings. Whole surface of pronotum characteristically ornamented in brown, reddish-brown,
bright yellow and white tints. Pronotum covering whole abdomen. Pronotum smooth, finely granulated. Anterior
margin of the pronotum very weakly projected forwards, almost truncated. Median carina continuous. Pronotal
discus in lateral view straight, plain. Prozonal carinae almost indistinct, parallel. Interhumeral carinae recognizable,
short and yellow. Humeroapical carina weak, bright yellow in color. Tegminal sinus absent. Ventral sinus deep.
Infrascapular area widened, extended to the tip of the pronotum in lateral view, but still narrow and with parallel
margins. Lateral area very narrow. Lateral carinae bright yellow in color. Lateral lobes directed sideward, without
projections. Apex of the lateral lobe truncated. Pronotal tip acute, reaching hind knees.
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32 · Zootaxa 5524 (1) © 2024 Magnolia Press
Wings. Tegmina absent or reduced and covered by infrascapular area. Hindwings absent or reduced and covered
by infrascapular area.
Legs. Fore and mid legs elongated, smooth, grey-brownish in color. Dorsal and ventral carinae of the fore and
mid legs smooth, finely granulated, slightly undulated and almost straight. Hind femur with reddish-brown outer
ventral area, grey-brownish dorsal area, and central area with a white line. Five transverse ridges elevated from
the smooth outer surface. Dorsal carina of the hind femur smooth and straight Antegenicular teeth small and blunt,
genicular teeth long and sharp. Hind tibia with 4 large spines on the outer and inner margins and many fine and blunt
teeth. First tarsal segment slightly shorter than the third Tarsal puvilli obliquely angular. Proximal pulvillus smallest,
middle larger, distal largest. Hind tibia uniformly yellow.
Ovipositor. Ovipositor very small and with strongly armed valvulae. Cerci triangular and hairy
Holotype measurements (female). Body length from tip of head to the tip of pronotum 8.85 mm; antenna
length 1. 90 mm; eye width 0.60 mm; vertex width 0.55 mm; pronotum length 7.70 mm; pronotum width (at
shoulders) 2.80 mm; fore femur length 2.60 mm; fore femur width 0.45 mm; mid femur length 2.60 mm; mid femur
width 0.50 m; hind femur length 5.50 mm; hind femur width 1.65 mm.
Paratype measurements (male). Body length from tip of head to the tip of pronotum 9.00 mm; antenna length
N/A (antennae broken); eye width 0.55 mm, vertex width 0.50 mm; pronotum length 7.90 mm; pronotum width (at
shoulders) 2.75 mm; fore femur length 2.20 mm; fore femur width 0.50 mm; mid femur length 2.60 mm; mid femur
width 0.60 mm; hind femur length 5.60 mm; hind femur width 1.70 mm.
Distribution and habitat: Endemic to the mountainous tropical rainforests of Mount Balatukan at 1 300 m a.
s. l. on the island of Mindanao, the Philippines (Fig. 2D).
Parahirrius amorosus Skejo, Patano et Kasalo, sp. nov.
Zoobank ID: urn:lsid:zoobank.org:act:7DFBB44D-B65E-4612-85DB-A981105538F1
Vernacular name: Mindanao Yellow Vividhopper (Figs. 20, 21)
Type specimens: 1♀ holotype THE PHILIPPINES: Mindanao: Bukidnon: San Fernando: Mount Natampod 980
m a. s. l. 24.–25.VII.2022. leg. R. Patano (CMU); 1♂ paratype THE PHILIPPINES: Mindanao: Bukidnon: San
Fernando: Pantaron Range: Mount Natampod 980 m a. s. l.. 24.–25.VII.2022. leg. R. Patano (CMU).
Type locality. THE PHILIPPINES: Mindanao: Bukidnon: San Fernando: Pantaron Range: Mount Natampod.
Type depository. Holotype and paratype are deposited in the Central Mindanao University (CMU) Orthoptera
Collection, Mindanao, Philippines.
Etymology. Named after Academician Victor B. Amoroso, a renowned scientist and a famous Philippine
botanist, and our dear friend and collaborator. His excellent mentorship and support to Romeo Patano Jr., author
of this study, led to discoveries of many new species of pygmy grasshoppers from Mindanao. Specific epitheton is
the second Latin declension masculine adjective, derived from Professor Amoroso’s surname, ‘amorosus, amorosa,
amorosum’.
Diagnosis. The species is similar to Parahirrius parvus sp. nov., but can be easily separated from that species
by (1) larger size, (2) characteristic coloration of pronotum and legs, (3) by a very small ovipositor in females, and
(4) by a slightly wider vertex.
Description. Small, smooth, flightless, and dorsoventrally flattened colorful species.
Antennae. Antennae filiform. Antennae with 16 antennomeres; 1st massive scapus, 2nd large pedicel, 3rd to
6th basal segments - about 2-3 times as long as wide; 7th to 9th central segments, about 3-4 times as long as wide,
10th to 11th subapical segments, elongated, about 5 times as long as wide, 12th segment as long as wide, 13th to
16th apical segments, reduced and hard to recognize.
Head. Head color bright yellow, with light brown markings in the frons and clypeal triangle. Eyes in frontal
view globose. Vertex in the level of pronotum in lateral view. Frons and vertex forming 45 degrees angle. In frontal
view, vertex narrower than a compound eye, at the level of the dorsal margins of the compound eyes. In lateral
view, vertex not visible, except for the tip of the frontal costa and medial carina connection. In dorsal view, vertex
oblique, not projected before the eyes. Lateral carinae of the vertex slightly elevated above the eyes, so small horns
are recognizable. In dorsal view, vertex almost as wide as a compound eye Anterior margin of the vertex truncated.
Medial carina of the vertex present, recognizable. Fossulae deep, so medial carina of the vertex and lateral carinae
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 33
seem to be strongly elevated. Lateral and transverse carinae forming V shape in frontal and dorsal view. Frontal
costa long and recognizable between the eyes in frontal view. Frontal costa bifurcation situated in the lower third
of the compound eye height. Frontal costa in lateral view almost not projected forwards at all. Dorsal margin of
the antennal groove placed below the lower margin of the compound eye. Lateral ocelli placed in the lower fourth
of the compound eyes height, just below the frontal costa bifurcation. Maxillary palps yellow, smooth, elongated.
Occipital area as wide as a half of a compound eye.
FIGURE 20. Parahirrius amorosus sp. nov. from Mount Natampod ♀ holotype. (A–G) and ♂ paratype (H) deposited in CMU.
A) right lateral view, B) ventral view, C) dorsal view, D) head in frontal view, E) head and pronotum detail in lateral view, F)
head and pronotum detail in dorsal view, G) ♀ abdominal apex in lateral view, H) ♂ abdominal apex in lateral view.
SKEJO ET AL.
34 · Zootaxa 5524 (1) © 2024 Magnolia Press
Pronotum. Whole surface of the pronotum characteristically ornamented in yellow and black markings.
Pronotum covering whole abdomen. Pronotum smooth, finely granulated. Anterior margin of the pronotum very
weakly projected forwards, almost truncated. Median carina continuous. Pronotal discus in lateral view straight,
plain. Prozonal carinae weak, parallel. Interhumeral carinae absent. Humeroapical carina almost non-existent.
Tegminal sinus absent. Ventral sinus deep. Infrascapular area widened, but still narrow and with parallel margins.
Lateral area very narrow. Lateral lobes directed sidewards, without projections. Apex of the lateral lobe truncated.
Pronotal tip acute, reaching hind knees.
FIGURE 21. Parahirrius amorosus Skejo, Patano et Kasalo sp. nov. living ♀ holotype in dorsolateral view, standing on a leaf
in natural habitat, Mount Natampod. Photo R. Patano Jr.
Wings. Tegmina absent or reduced and covered by infrascapular area. Alae absent or reduced and covered by
infrascapular area.
Legs. Fore and mid legs elongated, smooth, with brownish markings. Dorsal and ventral carinae of the fore and
mid legs smooth, finely granulated, slightly undulated and almost straight. Hind femur basal color dark, with yellow
markings. Four to five transverse ridges elevated from the smooth surface. Dorsal carina of the hind femur smooth
and straight. Antegenicular teeth small and angular, genicular teeth long and sharp. Hind tibia with 4-5 large spines
on the outer and inner margins and with many minute teeth. First tarsal segment slightly shorter than the third. Tarsal
pulvilli oblique, almost rounded. Proximal and middle almost equal in size, the distal one larger than them. Hind
tibia uniformly yellow.
Ovipositor. Ovipositor tiny with strongly armed valvulae. Cerci small and smooth.
Holotype measurements (female). Body length from tip of head to the tip of pronotum 10.85 mm; antenna
length 2.90 mm; compound eye width 0.60 mm; vertex width 0.50 mm; pronotum length 8.50 mm; pronotum width
(at shoulders) 2.45 mm; fore femur length 2.80 mm; fore femur width 0.65 mm; mid femur length 2.90 mm; mid
femur width 0.70 mm; hind femur length 6.05 mm; hind femur width 1.70 mm.
Paratype measurements (male). Body length from tip of head to the tip of pronotum 9.20 mm; antenna length
2.10 mm; compound eye width 0.45 mm; vertex width 0.40 mm; pronotum length 7.60 mm; pronotum width (at
shoulders) 2.30 mm; fore femur length 2.50 mm; fore femur width 0.40 mm; mid femur length 2.70 mm; mid femur
width 0.55 mm; hind femur length 5.35 mm; hind femur width 1.65 mm.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 35
Distribution and habitat. Endemic to the lower montane tropical rainforests of Mount Natampod, 980 m a. s.
l. on the island of Mindanao, the Philippines (Fig. 2E).
Subfamily Scelimeninae Bolívar, 1887
Tribe Discotettigini Hancock, 1907
Genus Zvierckia Skejo, Tumbrinck et Pushkar, gen. nov.
Zoobank ID: urn:lsid:zoobank.org:act:A9011CA9-F3E6-4C34-B7EF-CA259292D9F9
Vernacular name: “Sulawesi Pygmy Devils” (Figs. 1, 22–32)
Historical mentions: Zvierckia Skejo, Tumbrinck, Šapina et Puškar in Skejo, 2017: 185–193; unavailable according
to Article 13.1 of the Code (ICZN, 1999).
Type species: Zvierckia storozhenkoi Skejo et Tumbrinck sp. nov., here designated.
Etymology: Diminutive of the Slavic word for ‘beast’ (zvir, zvier, zver in Slavic languages), owing to the beasty
look of Zvierckia members, but also after our friend Dora Zvjerković, who found this genus interesting. The genus
name is of feminine gender.
Composition and distribution. Endemic to Indonesia, Sulawesi (in older literature Celebes), where three
species live, of them Z. sarasinorum comb. nov. in the northern part, Z. storozhenkoi sp. nov. in the central part,
and Z. montanta comb. nov. in the southeastern mountains (Fig. 22A, 22B, 22C).
Diagnosis. Based on (a) the low position of the frontal costa bifurcation, (b) low position of the antennal
grooves, (c) characteristic vertex shape with horns and deep fossulae, (d) pronotal projections arrangements, (e)
triangular infrascapular area, and (f) toothed femora, we place this new genus to the tribe Discotettigini of the
subfamily Scelimeninae. It is generally very similar to Discotettix, from which it can easily be distinguished by the
complete reduction of the pronotal projection of the discus.
Description. Moderately large robust species (body size up to 15 mm) of widened subapical antennal segments,
with tegmina partly covered by the pronotum and alae shorter than pronotum, with finely granulated surface.
Pronotum and body dark with rich maculations (red, yellow, orange). Fore and mid tibiae stripped.
Antennae. With 14 antennomeres, long, and with strongly widened subapical antennomeres. Rough in surface and
with saw-like margins (= basiconic sensilla). Similar to antennae of Discotettix members, but with one basal antennomere
more (so the widest segment is 9
th
and not 8
th
like in Discotettix). 1
st
massive scapus, barrel-like; 2
nd
large pedicel, 3
rd
to 7
th
basal segments, elongated (3
rd
and 4
th
antennomeres short, about 3 times as long as wide; 5
th
antennomere about
4 times as long as wide; 6
th
antennomere more than 6.5 times as long as wide; 7
th
about 7 times as long as wide). 8
th
,
subapical segment very elongated and slightly widened, 9
th
subapical antennomere is the widest segment, pennate and
about 2.3 times as long as wide in the widest part; 10
th
, apical antennomere pennate, reduced, small, about 3 times as
long as wide; 11
th
to 14
th
antennomeres, apical ones, completely reduced, cylindrical, and small.
Head. Frontal costa, in frontal view, long and bifurcating between the compound eyes; lateral ocelli situated
in the lower third of the compound eyes width; upper margins of the antennal grooves situated just below the
compound eyes lower margins. Antennal groove and scapus considerably wider than a narrow and almost parallel
scutellum. In dorsal view vertex truncated, in frontal view concave because of the slightly elevated lateral carinae
of the vertex. Vertex visibly wider (1.4–1.9 times) than a compound eye in dorsal and frontal view. Fossulae deep.
In dorsal view, medial carina of the vertex present only in the apical third. Compound eyes in dorsal view crescent-
moon shaped, in lateral view semi-globular, and in frontal view globular in shape. Maxillary palps flattened, dark
and usually with yellow markings.
Pronotum. Pronotum flat or weakly undulated, covering whole abdomen and extending to the abdominal apex
and lightly extending behind the hind femora in falling fashion. Anterior margin of the pronotum truncated in
dorsal view. Median carina continuous; prozonal carinae parallel. The apex of the pronotum blunt in dorsal view.
Humeroapical carinae forming with the external lateral carinae an obtuse, rounded angle. Pronotal projections
reduced, very low and barely recognizable in lateral view (minute FM, weakly elevated PM and MM1, as well as
very low MML1s), ML absent. Infrascapular area triangular covering most of the tegmina. Tegminal sinus weak,
much weaker than the ventral sinus. Lateral lobes directed outwards and bear weak VL or without it.
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FIGURE 22. Distribution of the species belonging to the genera Zvierckia gen. nov. and Guentheracris gen. nov. A) Z. montana
(Günther, 1937) comb. nov., B) Z. sarasinorum (Günther, 1937) comb. nov., C) Z. storozhenkoi sp. nov., D) G. scrobiculata
(Günther, 1937) comb. nov.
Legs. Dorsal and ventral margins of fore and mid femora slightly undulated and weakly toothed. Ventral margin
of the mid femur strongly undulated and usually with two to three teeth. Tibiae robust and with undulated margins,
rectangular in cross-section. Hind femora slender, about 2.8 to 3.3 times as long as wide. Six transverse ridges
visible on the external surface of hind femora. Femoral carinae and transverse ridges weakly tuberculated. Dorsal
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margin of the hind femora with finely serrated. Genicular and antegenicular teeth small. Hind tibia toothed and with
several larger spines. Proximal and distal segments of the hind tarsi almost equal in length. Tarsal pulvilli obliquely
angular, proximal two pulvilli half in size of the distal one.
Key to species of Zvierckia
1A) Lateral lobes of the pronotum truncated, without spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1B) Lateral lobes of the pronotum bearing spines. (Vertex 1.55 to 1.65 times wider than an eye. Hind femur 3 to 3.33 times as long
as wide. Face 4 to 4.1 times wider than an eye.) N Sulawesi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. sarasinorum, comb. nov.
2A) Dorsum undulated. Antennae black with yellow rings, especially in the subapical area. Vertex 1.47 times wider than an eye.
Hind femur 2.7 times as long as wide. Face 3.56 times wider than an eye. SE Sulawesi . . . . . . . . . . . Z. montana, comb. nov.
2B) Dorsum flat. Antennae uniformly black. Vertex 1.64 to 1.86 times wider than an eye. Hind femur 2.85 to 3.06 times as long as
wide. Face 4.00 to 4.54 times wider than an eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. storozhenkoi, sp. nov.
FIGURE 23. Comparison of head in frontal view in the members of the genus Zvierckia gen. nov.: Z. montana (Günther, 1937)
comb. nov., Z. sarasinorum (Günther, 1937) comb. nov., and Z. storozhenkoi Skejo et Tumbrinck sp. nov..
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List of Zvierckia species
Zvierckia montana (Günther, 1937), comb. nov.
Vernacular name: “Salokko Pygmy Devil” (Fig. 23, 24)
Historical mentions: As Hirrius montanus Günther 1937: 176; Günther 1938: 303; Blackith & Blackith 1987:
2; Blackith 1992: 90; As Zvierckia montana Skejo, 2017: 191; unavailable according to Article 13.1 of the Code
(ICZN, 1999).
Type specimen. 1♂ holotype ‘SO Celebes Berg Tangke Salokko 1500 m 1-15.1932 G. Heinrich’ ‘Hirrius
montanus n. sp. K. Günther’ ‘Typus’ (MfN).
FIGURE 24. Zvierckia montana (Günther, 1937) comb. nov. ♂ holotype from SO Celebes Berg Tangke Salokko. A) left lateral
view, B) dorsal view, C) labels, D) head in frontal view. Photo E. Nguyen.
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Diagnosis. In lateral view, the disc of the pronotum with undulated median carina which is low in prozona,
then elevated between prozona and metazona, and then weakly undulated at the apical descending portion. Lateral
lobes of the pronotum projected outwards, but with a truncated, blunt apex (without spikes, VL projections). Vertex
1.47 times as wide as an eye in males. Face 3.56 times as wide as an eye in males. Male antennae black with bright
yellow-orange rigs between the basal antennomeres, and with especially wide pale ring on the subapical and apical
antennomeres. General color black, but there are many yellow-orange (or red) dots on the median carina and on the
lateral carinae of the pronotum. Hind tibia with yellow rings.
Holotype measurements (male). Body length from tip of head to the tip of pronotum 13.45 mm; eye width
0.70 mm, vertex width 1.05 mm; pronotum length 12.35 mm; pronotum width (at shoulders) 4.40 mm; tegmen
length 1.25 mm; tegmen width 0.30 mm; fore femur length 3.25 mm; fore femur width 0.75 mm; mid femur length
3.70 mm; mid femur width 0.90 mm; hind femur length 6.65 mm; hind femur width 2.45 mm.
Distribution, habitat, and doubtful records. Indonesia, Southeastern Sulawesi, endemic to the mountainous
tropical rainforests of Tangke Salokko hills at 1500 m a. s. l. in the Mekongga Mountains above Kolaka (approximately
3.67S, 121.18E). Blackith’s (1987) report of Z. montana comb. nov. from Sulawesi Tengah, Mount Tambusisi (1200
m a. s. l.) (approximately 1.656S, 121.373E) 3.-13.4.1980. leg. Brendell (BMNH) needs to be checked as it may
represent another undescribed species.
Note. The only specimen of Z. montanta comb. nov, that for sure represents this species is male holotype,
collected by German entomologist and ornithologist Gerd Hermann Heinrich during the expedition to Mekongga
Mts. in early January 1932.
Zvierckia sarasinorum (Günther, 1937), comb. nov.
Vernacular name: “Sulawesi Spiky Pygmy Devil” (Fig. 23, 25, 26)
Historical mentions: As Hirrius sarasinorum Günther 1937: 176; Blackith 1992: 90; As Zvierckia sarasinorum
Skejo, 2017: 191 (not valid nor published combination as Diploma thesis cannot be regarded as a publication under
ICZN 1999).
Type specimens examined. 1♀ holotype ‘Celebes Ile Ile 500 m 11.12.1930 G Heinrich leg.’ ‘Hirrius
sarasinorum K. Günther’ ‘Typus’ (MfN); 3♂♂ paratypes ‘Celebes Ile Ile 500 m 11.12.1930 G Heinrich leg.’
‘Hirrius sarasinorum K. Günther’ ‘Typus’ (MfN).
Type specimens not examined. 1♀ paratype ‘Nord Celebes Toli Toli, 11.XII.1895., Fruhstorfer leg.’ (Stettin);
1♂ paratype ‘Celebes Ile Ile 500 m 11.12.1930 G Heinrich leg.’ (MfN or SMTD); 1♀paratype ‘Celebes’ ‘Dres.
Sarasin legrnt.’ (NHMB = NMBA).
Annotated diagnosis. In lateral view, the disc of the pronotum almost straight, except for the caudal part in
which pronotum gradually descends. Lateral lobes of the pronotum bearing spines (ventrolateral, VL projections).
Vertex 1.55 times as wide as an eye in males, and 1.64 times in females. Face 4.10 times as wide as an eye in males,
4.08 times in females. Female antennae dark with weak yellow rings between basal antennomeres, male antennae
black with pale yellow rings between the basal antennomeres. General color black, except for the yellow-orange
dorsum of the pronotum and hind femora, and with especially bright median and lateral carinae of the pronotum.
Hind tibia uniformly brown.
Holotype measurements (female). Body length from tip of head to the tip of pronotum 16.85 mm; eye width
0.75 mm, vertex width 1.20 mm; pronotum length 15.75 mm; pronotum width (at shoulders) 4.85 mm; tegmen
length 1.55 mm; tegmen width 0.45 mm; fore femur length 4.10 mm; fore femur width 0.95 mm; mid femur length
3.85 mm; mid femur width 0.90 mm; hind femur length 8.60 mm; hind femur width 2.85 mm.
Paratype measurements (male). Body length from tip of head to the tip of pronotum 15.75 mm; eye width
0.75 mm, vertex width 1.10 mm; pronotum length 14.50 mm; pronotum width (at shoulders) 4.30 mm; tegmen
length 1.65 mm; tegmen width 0.50 mm; fore femur length 4.10 mm; fore femur width 0.90 mm; mid femur length
3.80 mm; mid femur width 0.85 mm; hind femur length 8.00 mm; hind femur width 2.40 mm.
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Distribution, habitat, and questionable records. Indonesia, northern Sulawesi, endemic to the mountainous
tropical rainforests above of Mt. Dako above Tolitoli and of Gunung Ile-Ile Mt. (500 m a.s.l) that is part of the
Matinan Mountains. A record from southwestern Sulawesi (1♂ Loka, Piek van Bonthain X.1895. leg. drs. Sarasin
(NMBA)), not examined by us, and that was not included by Günther (1937) in the type material needs to be checked
as it could belong to another undescribed species. Lompobattang (Lompobatang, Piek van Bantaäng, Piek van
Bonthain, approximately 5.34S, 119.93E) is a mountain in the southwestern-most part of Sulawesi, on to opposite
side from other known localities of Z. sarasinorum comb. nov.
FIGURE 25. Zvierckia sarasinorum (Günther, 1937) comb. nov. ♀ holotype from Ile Ile. A) left lateral view, B) dorsal view,
C) labels, D) head in frontal view. Photo E. Nguyen.
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FIGURE 26. Zvierckia sarasinorum (Günther, 1937) comb. nov. ♂ paratype from Ile Ile. A) left lateral view, B) dorsal view,
C) labels, D) head in frontal view. Photo E. Nguyen.
Note. Günther (1937) doubted whether Z. sarasinorum comb. nov. may represent only a local form of Z.
montana comb. nov., but now when we have also a new species from central Sulawesi before us, it is clear that the
differences between the three species are clear enough to distinguish them as separate species and that Sulawesi
Spiky Pygmy Devil represents a unique species. This species was described based on few speciments, males and
females, but lables ‘holotype, allotype and paratype’ under specimens from Ile Ile absent (Günther, 1937). Here we
considered female from Ile Ile as holotype and males from this locality as paratypes.
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Zvierckia storozhenkoi Skejo et Tumbrinck, sp. nov.
Zoobank ID: urn:lsid:zoobank.org:act:A7D33548-28F0-459A-A3E1-612700420969
Vernacular name: “Palu Pygmy Devil” (Figs. 23, 27, 28, 29, 30, 31)
Historical mentions: As Zvierckia storozhenkoi Skejo, Tumbrinck, Šapina et Puškar in Skejo, 2017: 187-190
(not valid name and authorship nor published genus and species name as Diploma thesis cannot be regarded a
publication under ICZN 1999).
FIGURE 27. Zvierckia storozhenkoi Skejo et Tumbrinck gen. et sp. nov. ♀ holotype (A–E) and ♂ paratype (F-J) from Sulawesi,
Palu region, deposited in CJT. A and F) dorsal view, B and G) right lateral view, C and H) left lateral view, D and I) head in
frontal view, E and J) labels. Photo J. Tumbrinck.
Type specimens (♀ holotype, 9♂♂ paratypes, 10♀♀ paratypes, 8 nymph paratypes). 1♀ holotype
‘INDONESIA: C Sulawesi: 20 km NE Pale, ca. 6km W Tawaeli, 250 m N 0° 43’ 45’’ E 119° 55’ 95’’, 02.III.2009,
leg. A. Skale (019)’ (CJT); 1♂ paratype ‘INDONESIA: C Sulawesi: 20 km NE Pale, ca. 6km W Tawaeli, 250 m
N 0° 43’ 45’’ E 119° 55’ 95’’, 02.III.2009, leg. A. Skale (019)’ (CJT); 1♀ paratype ‘INDONESIA: CW Sulawesi
Palu district, Palolo env. V.2017. ZMH861001’ (ZMH); 1♂ paratype ‘INDONESIA: CW Sulawesi Palu district,
Palolo env. V.2017. ZMH861002’ (ZMH); 1♀ paratype ‘INDONESIA: SC Sulawesi: Palopo Palu I.2017.
ZMH831022’ (ZMH); 1♀ paratype ‘INDONESIA: SC Sulawesi: Palopo Palu I.2017. ZMH831023’ (ZMH); 1♂
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paratype ‘INDONESIA: SC Sulawesi: Palopo Palu I.2017. ZMH831024’ (ZMH); 3♂♂, 5♀♀, 8 nymph paratypes
‘INDONESIA: Sulawesi Island, province Sulawesi Tengah (=Central Sulawesi province), Lore Lindu National
Park, 45 km SSE of Palu City, env. of village Tomado on Lindu Lake, 1000 m, 13–17.II.2011, coll. A.V. Gorochov’
(ZISP); 3♂♂, 2♀♀ paratypes ‘INDONESIA: Sulawesi Island, province Sulawesi Tengah (=Central Sulawesi
province), Lore Lindu National Park, 75 km SE of Palu City, env. of village Wuasa (near Eastern park edge), 1000
m, 7–12.II.2011, 3 males, 2 females, coll. A.V. Gorochov’ (ZISP).
Type locality: INDONESIA: Sulawesi: C Sulawesi: hills around Palu.
Type specimens’ depository: Holotype female and a paratype male are deposited in the Collection Josef
Tumbrinck (CJT) in Wassenberg, Germany, 3 male paratypes and 3 female paratypes are deposited in the Orthoptera
collection of the Zoological Museum Hamburg, while 6 male, 7 female, and 8 nymph paratypes are deposited in the
Zoological Institute, St. Petersburg.
FIGURE 28. Zvierckia storozhenkoi sp. nov. ♀ paratype ZMH 861001 (A–D) and ♂ paratype ZMH 861002 (E-I) from
Sulawesi, Palu region. A and E) dorsal view, B and F) left lateral view, C and H) head in frontal view, D and I) labels, G)
antennae. Photo E. Nguyen.
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FIGURE 29. Zvierckia storozhenkoi sp. nov. ♀♀ paratypes (ZMH 831022, 831023) and a ♂ paratype (ZMH831024) from
Sulawesi, Palu region. A, H and J) dorsal view; B, F and I) left lateral view; C, G and K) paratype labels; D, E, and L) head in
frontal view. M) locality label. Photo E. Nguyen.
Etymology. Named after our dear friend and colleagues, Sergey Yurievich Storozhenko, a famous Russian
entomologist who has described 12 Tetrigidae genera and 55 species. The specific epithet is a Genitive case noun of
the second Latin declension of Sergey’s Latinized surname, storozhenkous, storozhenkoi.
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FIGURE 30. Zvierckia storozhenkoi sp. nov. ♀ paratype (A–C) and ♂ paratype (D–F) from Sulawesi, Palu region. A and D)
left lateral view; B and E) head in frontal view; C and F) dorsal view. Photo S. Yu. Storozhenko.
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FIGURE 31. Zvierckia storozhenkoi sp. nov. abdominal apex of ♂ paratype and (A, B) ♀ paratype (C–E) from Sulawesi, Palu
region. Photo S. Yu. Storozhenko. A) ♂ details of the male apex in dorsal view, B) ♂ apex in dorsal view, C) ♀ apex in dorsal
view, D) ♀ apex in ventral view, E) ♀ apex in lateral view. Photo S. Yu. Storozhenko.
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Diagnosis. Head traits similar to Z. sarasinorum, while pronotum traits similar to Z. montana. In lateral view,
the disc of the pronotum almost straight, only with weakly undulated median carina. Lateral lobes of the pronotum
projected outwards, but with a truncated, blunt apex (without spikes, VL projections). Vertex 1.64–1.85 times as
wide as an eye in males, 1.76–1.86 times in females. Face 4.03–4.25 times as wide as an eye in males, 4.25–4.54
times in females. Male antennae uniformly black. General color black, but there are many orange-red dots on the
median carina and on the pronotal discus. Hind tibia black.
Description. Moderately large and robust species (14–15 mm long). Body finely granulated. Only brachypronotal,
brachypterous specimens known. Coloration seems to be species specific. General body color black or dark grey.
Pronotum often with reds prozona; median carina usually ornamented in orange and red spots; prozonal discus with
alternating red and dark colored regions. Outer surface of the hind femur black with red markings.
Antennae. Antennae relatively long, reaching half of the pronotum length; uniformly black, but in males’
basal segments often with very lightly yellowish proximal margins. Antennae 14-segmented. 1st scapus, 2nd pedicel,
3rd and 4th (basal) antennomeres short; 5th antennomere slightly more elongated than the previous two; 7th (basal)
antennomere very elongated; 8th antennomere (subapical) elongated and weakly widened, 8–9 times as long as wide;
subapical segment 9th strongly widened and foliaceous; 10th (apical) antennomere much smaller than the previous
ones, slightly widened; apical segments (11th to 14th reduced and cylindrical.
Head. Vertex wider than a compound eye 1.64–1.85 times in males, 1.76–1.86 times in females. Frontal costa,
in frontal view, long, bifurcating just between the compound eyes; in lateral view not visible. Eyes in dorsal view
crescent-moon shaped, in lateral view semi-globular, in frontal view globular. Occipital area very narrow. In frontal
view, scutellum narrow, facial carinae almost parallel, slightly diverging from the level of the antennal grooves and
below. Antennal grooves positioned visibly (0.1–0.25 mm) below the lower margin of the compound eyes. Antennal
groove and scapus each considerably wider than scutellum. Lateral ocelli situated very low, in the lower third of
the compound eye height (females) or parallel with the inferior margin of the compound eyes (males). In lateral
view, vertex in the level of pronotum. In dorsal view, fastigium of vertex truncated, not produced forwards, slightly
concave in dorsal and frontal view. Fossulae present. Median carina of vertex present only in the distant third.
Lateral carinae of the vertex visible in frontal view as strong tuberculated elevations, in dorsal view forming acute
angle. Maxillary palps flattened, black with yellow margins.
Pronotum. In lateral view, the discus of the pronotum almost flat. Pronotum covering whole abdomen and
extends to the abdomen apex and slightly surpasses hind femora; in lateral view descending towards the apex.
Pronotal apex, in dorsal view, blunt. Anterior margin of the pronotum truncated. Median carina continuous, in
lateral view weakly undulate. Prozonal carinae parallel. Interhumeral carinae short and parallel. In lateral view,
a weak depression, giving an impression of the undulation of the median carina of the pronotum, visible on the
discus just behind the shoulders. Humeroapical carinae forming with external lateral carinae obtuse, rounded angle.
Infrascapular area triangular and as wide as a half of the mid femora in the widest part, running from tegminal
sinus to the level of the 4/5 of the hind femora length where it fuses with wide lateral area. Lateral lobes directed
outwards, with truncated apices, not bearing spine, i.e., the ventrolateral (VL) projection. Pronotal projections (tiny
FM, fused low PM and MM1, low MMLs) extremely reduced and hardly visible; humeral ML completely absent.
Tegminal sinus lobe weak, produced less than ventral sinus.
Wings. Tegmina small and only partly visible. Tip of hindwings not reaching the tip of the pronotum in lateral view.
Legs. Fore and mid femora elongated, with undulated margins. Dorsal and ventral margin of the fore femora
slightly undulated and finely toothed. Mid femora with weakly undulated dorsal and ventral margins, both distally
bearing a small tooth. Distal tarsal segments of fore and mid legs very long, black with a wide white stripe in the
middle. Hind femora slender (2.9–2.95 times as long as wide). External median area with small and large tubercles.
Six transverse ridges present, of which two proximal to the knee very week. Genicular and antigenicular teeth small.
Dorsal margin of the hind femora finely serrated. Hind tibiae dark gray, finely serrated and with several larger
spines, and with a narrow pale colored band below the knee. First and third tarsal segments equal in length. Tarsal
pulvilli obtusely angular, first two smaller than the third.
Measurements. See Table 3.
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TABLE 3. Measurements of Zvierckia storozhenkoi sp. nov. holotype (HT) and some paratypes (PT). Body length is
measured from the tip of the head to the end of the pronotum. All measurements are given in millimeters.
HT ♀
CJT
PT ♀ ZMH
831023
PT ♀ ZMH
861001
PT ♀ ZMH
831022
PT ♂
CJT
PT ♂ ZMH
861002
PT ♂ ZMH
831024
Body length 16.60 15.40 14.95 14.95 15.05 14.05 12.95
Eye width 0.65 0.80 0.75 0.70 0.60 0.70 0.70
Vertex width 1.20 1.35 1.30 1.30 1.05 1.15 1.15
Pronotum length 15.30 14.10 13.50 13.85 14.20 12.45 12.00
Pronotum width (at shoulders) 6.05 5.05 4.65 4.80 5.00 4.80 4.25
Tegmen length 1.85 1.40 1.30 1.65 1.30 1.25 1.25
Tegmen width 0.60 0.50 0.50 0.50 0.40 0.40 0.45
Fore femur length 4.70 4.05 4.00 3.85 4.10 3.70 3.55
Fore femur width 1.20 0.95 0.90 0.95 1.15 0.70 0.80
Mid femur length 5.00 4.00 4.20 3.75 4.20 4.00 3.50
Mid femur width 1.00 1.00 0.85 0.90 1.05 1.00 0.70
Hind femur length 9.50 8.55 8.10 8.30 7.90 6.80 7.25
Hind femur width 3.10 3.00 2.65 2.75 2.80 2.30 2.50
FIGURE 32. Living individual of Zvierckia sp. from Sulawesi, photographed by macroid.ru user iris-G62 on 14.II.2011. and
available on the link macroid.ru/showphoto.php?photo=74819.
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Distribution and habitat. Endemic to the mountainous tropical rainforests in the hills around Palu in central
Sulawesi, Indonesia (Fig. 22C).
Morphology of antennae in Zvierckia and Phaesticus
The widest subapical antennomeres of a Zvierckia storozhenkoi sp. nov. male from Palu, Sulawesi and all the
widened subapical antennomeres of a Phaesticus mellerborgi (Stål, 1855) male from Sumatra were scanned and
their microscopic morphology was studied. A comparison of the antennae of Zvierckia gen. nov, Phaesticus, and
Discotettix (after Kuřavová et al. 2017) is presented in Table 4. For Zvierckia gen. nov., and Phaesticus these are
the first data on the microscopic morphology of subapical antennomeres.
TABLE 4. Comparison of the microscopic morphology of the widest antennomeres of Zvierckia storozhenkoi sp. nov.,
Discotettix belzebuth (Serville, 1838) (after Kuřavová et al. 2017), and Phaesticus mellerborgi (Stål, 1855). Abbreviations:
D—dorsal surface, V—ventral surface of the antennomere.
Zvierckia storozhenkoi Discotettix belzebuth Phaesticus mellerborgi
Subfamily and tribus placement Scelimeninae: Discotettiginae Scelimeninae: Discotettiginae Incertae sedis within
Tetrigidae
Sex studied male males and females male
Number of antennomeres 14 13 13 or 14
Widest antennomere 9th 8th 8th or 9th
Side of the antennomere D V D V D V
Protuberances and cuticular plates + + + + + +
Elongated multiporous placoid fields - - - + - +
Multiporous coeloconic fields - + - + - +
Modified basiconic sensilla + + + + + +
Zvierckia storozhenkoi
(Figs. 33–34)
This species has 14-segmented antennae, the widest antennomere being the 9th, 1.45 mm long, 0.70 mm wide at
the widest part, and with a surface on each side of about 0.73 mm2. The 9th antennomere is circular in the cross-
section in the base, and elongated, ovoid, or almost rectangular in the cross-section in the tip. The whole surface of
the antennomere is covered in protuberances and cuticular plates (12 µm x 10 µm) (Fig. 33, Fig. 34). A 6-porous
coeloconic field (covering an area of about 310 µm2, one pore with about 5 µm diameter) was observed only in
the apical antennomere’s ventral surface (Fig. 34). Around the coeloconic fields, there were no protuberances and
cuticular plates, i.e., the surface was smooth. No coeloconic fields were observed in the dorsal part (Fig. 33). Huge
(some with the cone longer than 70 µm) modified basiconic sensilla cover the lateral margins of the antennomere,
giving the margin a saw-like appearance. Much smaller basiconic sensilla (about 15 µm long cone at the base)
are found on the dorsal and ventral surfaces. No multiporous coeloconic fields were observed on the dorsal nor
the ventral surface of this antennomere. The segment looks very similar to the widest antennomeres in related
Discotettix (Kuřavová et al. 2017), except for the absence of the placoid fields, which are present in Discotettix.
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FIGURE 33. Scanning electron microscope (SEM) scan of the dorsal side of Zvierckia storozhenkoi sp. nov. 9th antennomere
(widest), with the details of cuticular plates and basiconic sensilla.
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FIGURE 34. Scanning electron microscope (SEM) scan of the dorsal side of Zvierckia storozhenkoi sp. nov. 9th antennomere
(widest), with the details of coeloconic fields, cuticular plates, and basiconic sensilla.
Phaesticus mellerborgi
(Figs. 35–39)
This species has 13- or 14-segmented antennae. Namely, in this species, as already reported by Zha et al. (2021), the
third antennomere (first flagellar) may be divided into two small segments. Thus, the widest antennomere is the 8th
or the 9th. The widest segment is elongated and ovoid in the cross-section in both base and apex. However, its apical
cross-section has a wider perimeter.
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FIGURE 35. Scanning electron microscope (SEM) scan of the dorsal side of Phaesticus mellerborgi (Stål, 1855) 7th antennomere,
with the details of cuticular plates and basiconic sensilla.
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FIGURE 36. Scanning electron microscope (SEM) scan of the ventral side of Phaesticus mellerborgi (Stål, 1855) 7th
antennomere, with the details of coeloconic fields, cuticular plates, and basiconic sensilla.
7th antennomere (Fig. 35, Fig. 36) is a bit narrower than the 8th, 0.93 m long and 0.47 mm wide at the widest part
and with an area on each side of about 0.35 mm2. It is the second widest antennal segment in this species. Dorsal
side fully covered in cuticular plates (each about 12 µm x 8 µm). Ventral side (Fig. 36) with 9-porous coeloconic
field covering area of about 575 µm2 and with about 5 µm of the pore diameter. The coeloconic field similar in shape
to that of the 9th segment, with two upper pores and 7 lower ones. No elongated placoid fields are situated on this
antennomere and no basiconic sensilla on the antennal surface on any side. Small basiconic sensilla, with the base
cone up to 30 µm long are however situated on the margins.
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8th antennomere (Fig. 37, Fig. 38) is the widest one. It is 0.84 mm long, 0.52 mm wide and with area of one side
of about 0.35 m3. Its dorsal part is fully covered in triangular cuticular plates (12 µm x 9 µm). Ventral part (Fig. 38)
has a 4-porous coeloconic field covering an area of about 365 µm2, each pore with diameter of about 5 µm2. The
coeloconic field is situated in the apical part of the antennomere and has a single elongated placoid field (26 µm
long, 10 µm wide) next to it. The area around these fields is smooth and does not bear cuticular plates. No modified
basiconic sensilla were found on the antennal surface, but many small basiconic sensilla are present on the edge.
FIGURE 37. Scanning electron microscope (SEM) scan of the dorsal side of Phaesticus mellerborgi (Stål, 1855) 8th, i.e., the
widest antennomere, with the details of cuticular plates and basiconic sensilla.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 55
FIGURE 38. Scanning electron microscope (SEM) scan of the ventral side of Phaesticus mellerborgi (Stål, 1855) 8th, i.e., the
widest antennomere, with the details of one elongated placoid field, coeloconic fields, cuticular plates, and basiconic sensilla.
The 9th antennomere (Fig. 39) is 0.65 mm long, 0.28 mm wide at the widest part, and with an area of one side
of about 0.15 mm2 It is very similar in overall morphology to the previous two antennomeres, and looks on the
first sight as a minute version of any of them. It is also covered in cuticular plates (12 µm x 8 µm). However, upon
closer inspection it has a quite different set of traits. Its apical ventral side has, besides a 9-porous coeloconic field
(covering about 630 µm2 and with each of the pores with diameter from 3.5 µm to 5.5 µm), 11 large elongated
placoid fields (each about 30 µm long, and about 8 µm wide). The area around the coeloconic and placoid fields is
smooth. On each margin, there are small basiconic sensilla.
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56 · Zootaxa 5524 (1) © 2024 Magnolia Press
FIGURE 39. Scanning electron microscope (SEM) scan of the ventral side of Phaesticus mellerborgi (Stål, 1855) 9th
antennomere with the details of elongated placoid fields, coeloconic fields, and cuticular plates.
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 57
Discussion
Many new genera, but one still with a very uncertain placement
A reviewed system is presented for all the species hitherto assigned to the genus Hirrius. This is an old genus from
the standpoint of nomenclature and the one on which a lot of work has been done (Stål 1877, Bolívar 1887, Günther
1937, 1938). The last revision of the genus took place 85 years ago (Günther 1937, 1938) and in that revision,
Günther already noted that the genus had been composed of several likely unrelated groups. After our revision, we
confirm Günther’s doubts as we break up Hirrius into 4 genera, altogether including 9 species.
We have assigned Hirrius and Guentheracris gen. nov. to the Melainotettix genus group, defined here, based on
its similarity to Melainotettix and Camelotettiix from Papua, Guentheracris gen. nov. from Sulawesi, and Arulenus
from Mindanao. We have assigned Zvierckia gen. nov. to Scelimeninae and Discotettigini based on the similarity
to Discotettix. The genus Parahirrius gen. nov. is still a ‘lonely wolf’, a genus of uncertain placement within
Tetrigidae. This new genus shares some similarities with the members of the tribe Ophiotettigini Tumbrinck et
Skejo, 2017, namely the flat and largely smooth pronotum, projected lateral lobes, the first and third hind tarsal
segments of equal length, and mostly filiform antennae. However, Ophiotettigini members have highly derived head
morphology, with the narrow vertex, elongated head, and the higher placement of facial features. All of this is not
present in Parahirrius gen. nov. and thus makes a complete comparison and a confident conclusion impossible. This
potential relationship should be considered while gathering data for future molecular studies. Because of the many
unique traits Parahirrius gen. nov. exhibit we refrained from assigning this genus to the Melainotettix genus group
or to any other subfamily, tribe, or genus group within Tetrigidae.
In conclusion, we have described three new genera in order to sort out Hirrius, but for Parahirrius gen. nov. we
still have many questions to answer in the future. In future studies, we will try to obtain a series of fresh material,
especially of Parahirrius Skejo, Patano, Tan et Kasalo gen. nov., which would be suitable for DNA sequencing,
but also a series of all other genera from SE Asia for comparison. Namely, the lack of DNA sequences is significant
among Tetrigidae in this part of the world, so we regard it necessary to incorporate molecular methods in the
future.
A new genus and species hidden under the name “Hirrius punctatus” for more than 140 years
Problems with Parahirrius gen. nov. are not novel. The genus was problematic even before it was described. But
how is that possible? The Mindanao Tiny Vividhopper (Parahirrius parvus sp. nov.) is an excellent example of the
complexity of Tetrigidae taxonomy and the negligence some species and groups were exposed to. The exploration
of the Mindanao Tiny Vividhopper dates back to the designation of the syntype of Hirrius punctatus by Stål in 1877,
exactly 146 years ago or even at its discovery.
The potential oblivion of a species waiting to be discovered in another’s species syntype series was ended by
Filipino scientists and the coauthors of this study who stumbled upon a population of a previously unrecognized
species in Mount Balatukan at 1 300 m above sea level. It soon became clear that the members of this unique
population corresponded to a single male specimen that had been stored in Berlin for a century and a half before
its true identity came to light. The locality where the syntype of H. punctatus was caught was not known, so Mt.
Balatukan is the only locality from which Parahirrius parvus sp. nov. is known.
During the realization that one of the individuals within the syntypes of Hirrius punctatus belonged to another,
yet undescribed, genus and species, a dilemma arose regarding which species should bear the name of Hirrius
punctatus, and which should be considered a novel genus and species. This decision on this issue was relatively
straightforward, as Stål’s description clearly portrayed the presence of the widened antennal segments and the bright
spots on the pronotum, characteristics that undoubtedly set it apart from here described Parahirrius parvus sp. nov.,
a species of filiform antennae and without spots on the pronotal discus. This species testifies to the ever-evolving
nature of taxonomic classification in sheds of modern methods and detailed analyses.
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58 · Zootaxa 5524 (1) © 2024 Magnolia Press
Widened antennal segments in Zvierckia, Phaesticus and Discotettix
Microscopic morphology of antennae is well understood in many insect orders, e.g., Hymenoptera (Ren et al.
2023), Lepidoptera (Guo et al. 2022), Coleoptera (Haddad et al. 2023), including Orthoptera (Bland 1989, Ochieng
et al. 1998; Nakano et al. 2023). The data are unfortunately almost completely missing on the antennal sensilla in
pygmy grasshoppers, with only two papers (Bland 1991; Kuřavová et al. 2017) that dealt with that topic till today.
Bland (1991) examined mouthparts and antennal sensilla in 8 Nearctic Tetrigidae species, the Tetriginae Neotettix
femoratus, N. cristatus, Paratettix cucullatus, Tetrix arenosa, T. ornata, and T subulata; and the Batrachideinae
Paxilla obesa, and Tettigidea lateralis. Kuřavová et al. (2017) studied the antennal morphology of Bornean
Discotettix belzebuth (Scelimeninae: Discotettigini). In this study, we provide the first data on the antennal sensilla
on the subapical antennomeres of Zvierckia storozhenkoi sp. nov. (Scelimeninae: Discotettigini) and Phaesticus
mellerborgi (uncertain subfamily placement within Tetrigidae). Taking into account how scarce the data on the
microscopic antennal morphology of Tetrigidae are, especially on SE Asian taxa, our study certainly represents a
huge contribution to the field.
First insights into the diversity of the antennal sensilla type, number, and abundance in pygmy grasshoppers
(Bland 1991, Kuřavová et al. 2017, this study) show promising results for future research. Corticolous Discotettigini,
phytophylous Phaesticus and ground dwelling Tetriginae, and Batrachideinae (Bland 1991) show significant
differences in the sensilla type, number, and distribution on the antennomeres. Differences were reported also
between the sexes (Bland 1991, Kuřavová et al. 2017). Systematic comparison of several species from the same
genera should determine whether the number and arrangement of the sensilla are species-specific traits or do they
have a higher phylogenetic signal. From the comparison of Discotettix and Zvierckia gen. nov. we could hypothesize
that the microstructure of the antennae might prove valuable in higher taxonomy. Widened antennomeres are
not common among Tetrigidae (Cigliano et al. 2023) and filiform shape seems to be ancestral for the family.
Widened antennomeres are present in taxonomically and geographically unrelated groups, such as Lophotettiginae
in Americas (all species), Tripetalocerinae and Clinophestini Storozhenko, 2013 in SE Asia (all species), some
members of Discotettigini, Ophiotettigini, Metrodorini, Malagasy Metrodorinae, in Asian Phaesticus, and in some
species Mazarredia Bolívar, 1887 (Cigliano et al. 2023). The function of the widened Tetrigidae antennomeres
is still completely unknown (Kuřavová et al. 2017) and we will probably find out in future that the function is
not analogous among the different groups. For example, in Papuan phytophylous and colorful genus Ophiotettix
widened antennomeres may be significant in courtship, while we do not expect the same function in corticolous
cryptic Discotettigini members, in which it is already hypothesized that widened antennae may provide selective
advantage to the cryptic appearance (Kuřavová et al. 2017).
The genera Zvierckia gen. nov. and Discotettix are both members of the tribe Discotettigini and this is also evident
in many shared features related to the antennal morphology. Namely, Zvierckia gen. nov., and Discotettix are the
only known genera that have hugely enlarged basiconic sensilla on the antennal margins. On the other hand, unlike
D. belzebuth (Serville, 1838) (Kuřavová et al. 2017), Zvierckia storozhenkoi sp. nov. lacks elongated multiporous
placoid fields on the widest subapical antennomere. In D. belzebuth (Serville, 1838) elongated multiporous placoid
fields always come next to the multiporous coeloconic fields.
The antennae of Phaesticus mellerborgi are much smoother than those of Zvierckia gen. nov. There are many
basiconic sensilla on the P. mellerborgi antenna margins, but those are not as enlarged and elevated as in Discotettix
and Zvierckia gen. nov. Both Zvierckia storozhenkoi sp. nov. and P. mellerborgi lack coeloconic and placoid fields
on the dorsal surfaces of the antennae. Interestingly, the same was observed in Discotettix belzebuth (Kuřavová et
al. 2017) and in North American Tetriginae and Batrachideinae (Bland 1991), so it could represent a more general
pattern. Interestingly, in North American Tetriginae no basiconic sensilla type II were observed, but they were
reported in Batrachideinae (Bland 1991) and Scelimeninae (Kuřavová et al. 2017). Many of the huge marginal
basiconic sensilla in Zvierckia storozhenkoi sp. nov. represent this very type of sensilla. On the other hand, we
suspect that the basiconic sensilla in P. mellerborgi represent basiconic sensilla type I, which are common in
Tetriginae and Batrachideinae (Bland 1991).
REVISION OF POLYPHYLETIC HIRRIUS Zootaxa 5524 (1) © 2024 Magnolia Press · 59
Acknowledgements
Many people are to be thanked for their help, discussions, photographs, specimens, and literature that altogether
resulted in a comprehensive revision of what was hitherto Hirrius. Thanks to all the amateur photographers who
photographed pygmy grasshoppers, uploaded them to iNaturalist, or permitted us to use their photographs, among
them Shirley Sekarajasingham, Evedreine Mingo, Mark Gregory Rule, Ann Cabras (https://www.inaturalist.org/
people/anncabras24). Thanks to Sigfrid Ingrisch for photographing the specimens during the DORSA project, as
these photographs are valuable and much appreciated even today. We thank Eileen Nguyen (ZMH) for imaging the
types of Zvierckia montana and other specimens. We also would like to thank Birgit Jaenicke (MfN) for preparing
loans of type specimens. This study was part of Josip Skejo’s Master Thesis “Taxonomic revision of the pygmy
devils (Tetrigidae: Discotettiginae) with online social media as a new tool for discovering hidden diversity”
defended in 2017—so thanks to Damjan Franjević and Axel Hochkirch for valuable comments during mentoring
of the thesis—as well as a part of Josef Tumbrinck’s and Josip Skejo’s project funded by OSF ‘Digitalization of
Tetrigoidea types in European Musea’. Ivan Šapina considered the genus Hirrius for his Master’s thesis and we
thank him for this work. Many thanks to Ming Kai Tan, Hendrik Devriese and Sergey Yu. Storozhenko for all
the help, discussions, and literature. Special thanks to Sergey Yu. Storozhenko for preparing nice photographs of
Zvierckia storozhenkoi. The authors would like to extend their warmest thanks to the following funding agencies
in supporting biodiversity research programs in Mindanao; (1) Commission on Higher Education, as the funding
agency of our research in Marilog District, as part of DARE TO (Discovery-Applied Research and Extension for
Trans/Interdisciplinary Opportunities) program entitled “Saving Terrestrial Biodiversity: Inventory, Assessment,
Conservation and Capability Building in Marilog Forest Reserve, Southern Mindanao, Philippines”, (2) National
Research Council of the Philippines for funding our research in Mt. Agad-Agad entitled “Biodiversity, Inventory,
Assessment, and Conservation for Ecotourism Development in Mt. Agad-Agad, Lanao del Norte”, (3) Department
of Science and Technology Grants-in-Aid (DOST-GIA) program entitled “Biodiversity in Selected Mountain
Ecosystems of Mindanao for Conservation and Sustainable Development”, and the (4) Department of Environment
and Natural Resources for the funding the research on “Long Term Biodiversity Monitoring of the two-hectare
permanent plot and BAMS plots of Mt. Kitanglad Range Natural Park – ASEAN Heritage Park, Bukidnon,
Philippines” as well as for the issuance of Gratuitous permits in compliance to the with Republic Act No. 9147
for the collection of the specimens. Romeo Patano Jr. is grateful to the DOST–Science Education Institute for
the scholarship under the Science and Technology Regional Alliance of Universities for National Development or
STRAND program for the support of his Ph.D. degree program. In addition, we would like also to thank the different
local government units, local stakeholders, local researchers, indigenous people organizations (the Mamacila Tribal
Community of the Mount Balatukan, the MAMATRIPCEDI (Matigsalug-Manobo Tribal People Council of Elders
Davao Inc.) of the Marilog District and the Manobo-Tigwahanon indigenous people of the Barangay Namnam,
San Fernando Bukidnon). Lastly, the researchers and research assistants of CEBREM and to the administration of
Central Mindanao University, and the leadership of Dr. Rolito G. Eballe, University President, are also recognized
by the authors for their important contributions to the implementation and support of the study.
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