Gryllacrididae Ingrisch, 2018, sensu lato
Creators: name: Ingrisch, Sigfrid
Publisher: name: Zenodo
Year: 2018
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Orthoptera
Gryllacrididae

description: Key to the genera of Gryllacrididae occurring from South East Asia to New Guinea
Included are all genera so far reported from an area between about India in the West, China in the North and New Guinea in the Southeast. For most genera I studied and illustrated at least one species, and, if possible, the type species. A few genera I only know from their descriptions (see Material and methods).
1. Ventral margins of fore and mid-tibiae with 4 pairs of long spines plus one pair of apical spurs (Figs. 2A, C) and with a pre-apical spine on ventral margins of hind tibiae in addition to the apical spurs (Figs. 1H, 2C); if on fore or on mid tibiae only 3 pairs of long spines then at least with a pre-apical spine on both ventral margins of hind tibiae (Figs. 2C, 2F)................ 2.
- Ventral margins of fore and mid-tibiae with only 3 pairs (rarely only 2 pairs) of long spines plus one pair of apical spurs (Figs. 2B, 2D). Hind tibiae without pre-apical spine in addition to the apical spurs (Fig. 2D)............................. 67.
2. In adult specimens fore and hind wings fully developed, surpassing or at least covering abdomen; if not fully covering abdomen or more strongly shortened, wing venation clearly identifiable.............................................. 3.
- In adult specimens fore and hind wings strongly reduced or absent: apterous, micropterous, or strongly brachypterous species, in which the identification of single veins is not clear or impossible............................................ 54.
3. Fore wings with media in basal area fused with radius, giving the impression that radius has two posterior branches; cubitus anterior simply divided into two branches (Fig. 1B, D) or undivided (Fig. 1C); usually media anterior does not send a connecting branch to cubitus and there is no visible media posterior. Female ovipositor with apical widening and with a longitudinal lateral furrow that ends just before apical widening (Fig. 2E). (If media fused in basal area with radius but ovipositor without apical widening compare Asarcogryllacris.)................................................................ 4.
- Fore wings with media in basal area free, although sometimes running very close to it; in fully winged species, media anterior usually sends a short connecting branch to the anterior branch of cubitus anterior that then divides a second time; thus that MP+CuA consists of three branches, but in species with shorter wings their number can be reduced (Figs. 1 E–G) or fore wings with somewhat elevated parallel veins that do not show a clear branching (Fig. 1A). If media anterior is fused at base with radius then there are two branches between media anterior and the last branch of cubitus anterior (Fig. 2G) and/or the ovipositor has the tip simply rounded or acute. Female ovipositor without apical widening and without longitudinal furrow (Fig. 2F)................................................................................................... 14.
4. Median ocellus very large (Fig. 22). Both, fore and hind wings darkened with veins and veinlets light and bordered by light bands; in species with less darkened wings, the pattern is still distinct (Fig. 21). Veins of fore wings running parallel, substraight and in nearly equal distance to each other, only the basically fused radius/RS/MA system branched, other veins unbranched and straight (Fig. 1C). Male abdomen with projections from ninth abdominal tergite (Fig. 23)... Ocellarnaca Gor.
- Median ocellus often small and in dried specimens often indistinct; if large and distinct then wings with different color pattern. In fore wings, cubitus anterior usually divided into two branches and cubitus posterior and the first anal veins in middle of wing with sinuate course (Figs. 1B, D).................................................................... 5.
5. Medium sized to large and robust species, pronotum relatively wide and short. Hind wings transparent or with dark color pattern; if with dark color pattern then the centers of the cells are dark while bands along the cross veins light (Figs. 4, 5K). Fore wings of uniform color, often green when alive, or orange brown with subtransparent marginal area; some species with indicated or fully expressed dark pattern as on hind wings and/or with shining black maculae (Figs. 4, 9 A–E). Male ninth tergite prolonged and down-curved with median furrow; at tip with a pair of rather small teeth, spines or otherwise formed projections (Fig. 6). Female seventh abdominal sternite with apical process of variable size and shape, covering base of subgenital plate to a variable extend (Figs. 7–8)......................................................... Eugryllacris Karny, 1937
- With relatively less wide pronotum compared to body size. Hind wings transparent or of light basic color (e.g. yellow); if with dark color pattern then with dark bands running alongside of cross veins while centers of cells are transparent or of light basic color............................................................................................... 6.
6. Medium sized to large species. Male eighth abdominal tergite markedly prolonged and ninth tergite strongly modified: either the lateral areas restricted to base and in middle with a large process, or the lateral areas reduced and the ninth tergite partly moved to the underside of the eighth tergite and/or is moveable against it, or lateral areas extended and strongly bent down from central area thus that the ninth tergite appears nearly box or hood shaped and has at apical margin a pair of small tooth- or hook-shaped projections that are in situ completely hidden by the subgenital plate. Female subgenital plate wide at base, then strongly narrowed to the often bi-oval centro-apical area with medial furrow and apical notch; either with a small stylus-like projection in about central area of plate with slightly widened tip or female subgenital plate with a wide extra sclerite from preceding intersegmental membrane or very base of subgenital plate. Gryllacris sensu lato ............................. 7. - Male ninth abdominal tergite of rather normal shape, with a medial furrow or a pair of oval swellings in subapical area and with a pair of small apical appendages. Female subgenital plate without stylus-like projection or wide basal extra sclerite...................................................................................................... 10.
7. Male ninth abdominal tergite with lateral areas extended, strongly curved or bent down from central area laterally and apically thus that the ninth tergite appears nearly box or hood shaped, sometimes with small horns or other marks where the dorsal surface is bent to the apical surface or with projections from apico-lateral areas (Figs. 15B, G, K); apical margin pointing ventroproximad or in some species fully proximad, provided with a pair of small tooth- or hook-shaped projections often difficult to find since in situ they are hidden by the posterior surface of the ninth tergite and the subgenital plate (Figs. 15F, E, H arrows, 18G). Subgenus Gigantogryllacris and fuscifrons group of Gryllacris. ........................................... 8.
- Male ninth abdominal tergite with lateral areas strongly reduced (Figs. 13–14)..................................... 9.
8. Female subgenital plate with a broad and rather large basal sclerite from base or from preceding intersegmental membrane (Figs. 18 E–F)................................................ Gryllacris subgenus Gigantogryllacris Karny, 1937
- Female subgenital plate with a narrow protruding sclerite in midline (Figs. 16 N–O).................................................................................................... fuscifrons group of Gryllacris Serville, 1831
9. Male ninth abdominal tergite with lateral areas absent; ninth tergite hanging down from apical area of eighth tergite and moveable against it, always widening in apical area and with or without a second articulation joint before apical area, or ninth tergite fully hidden under eighth tergite with the prolonged process in situ folded towards body (Figs. 13 A–C, 14A–L). Fore wings often with conspicuous dark cross-veins and narrow dark bands bordering them (Figs. 11 C–D, E–G).................................................................................. Gryllacris subgenus Pardogryllacris Karny, 1937
- Male ninth abdominal tergite with at least short lateral areas along the full hind margin of the eighth tergite (Figs. 13 D–M). Fore wings normally of uniform color with main veins slightly darker than cross veins or with extended dark marks not restricted to the cross-veins (Fig. 10).............................. G. subgenus Gryllacris Serville, 1831 sensu stricto
10. Hind wings often yellow or ochre with dark bands along veinlets or dark color more extended but centre of cells in those areas at least partly with a light spot (Figs. 29A, D, F). Male ninth abdominal tergite rather weakly furrowed in midline, before apical margin split along furrow and provided with a pair of rather stout but short and often black acute teeth pointing ventro-mediad (Figs. 30 A–H). Female subgenital plate wide at base, afterwards with converging margins and split at tip into two short lobes or at least with concave apical margin; seventh abdominal sternite with a projection of varying size (minute to huge) covering middle of base of subgenital plate (Figs. 30 I–J, 31C–E)................................ Xanthogryllacris Karny, 1937
- Hind wings colored or transparent. Male ninth abdominal tergite more strongly furrowed along midline; on both sides of furrow often with an oval swelling; at hind margin with a pair of upright or upcurved projections or oval swelling with a short acute ventral process................................................................................. 11.
11. Fore wings subtransparent with hardly or strongly darkened veins; hind wings with or without narrow dark bands along crossveins, not very conspicuous (Figs. 3A, F). Male ninth abdominal tergite with a pair of stout oval swellings projecting below apical margin and carrying a compressed acute tooth at ventral tip or somewhat proximad of tip, thus not visible from behind (Figs. 3C, H, I). Female subgenital plate semi-oval to triangular with apex angularly excised (Fig. 3D)................................................................................................ Caustogryllacris Karny, 1937
- Male ninth abdominal tergite with a pair of small elongate projections inserted at or between apical swellings and pointing dorsad or dorso-mediad.................................................................................. 12.
12. Fore wings unicolored, often yellowish with nearly transparent anterior area; hind wings semi-transparent without marks (Figs. 25A, F, K). Face pale. Male ninth abdominal tergite with posterior area almost vertically bent down and slightly furrowed; tip bilobate and provided with a pair of compressed, upright, and slightly sinuate projections with acute tip (Figs. 25 B–C, M–O). Female seventh abdominal sternite overlap base of subgenital plate or separated from it by long membranous zone; subgenital plate variable (Figs. 25E, I–J); ovipositor sickle-shaped, apical widening weak but slightly thickened (Figs. 25D, G)...................................................................................... Otidiogryllacris Karny, 1937
- Hind wings often with dark cross-veins or with dark bands along cross-veins. Male and female characters different...... 1 3.
13. Hind wings semitransparent with dark bands along veinlets (Figs. 19A, F, I). Male ninth abdominal tergite at least in apical area furrowed in midline and with a pair of oval or more triangular swellings that terminate at tip or at ventral margin into a pair of projections pointing posteriorly or mediad (Fig. 20). Female seventh sternite roundly prolonged or of rather normal shape followed by a separate sclerite from intersegmental membrane, in both cases covering base of subgenital plate in middle; female subgenital plate with bilobate tip and medial keel or terminating into 2 long acute angular lobes (Fig. 19E)....................................................................................... Lyperogryllacris Karny, 1937
- Fore wings unicolored, with or without slightly darkened veins; hind wings semitransparent and then often with darkened cross-veins (Figs. 26A, F, J) or with dark bands in marginal area (Fig. 28H) or of yellowish to ochre basic color and with distinct dark bands or oval spots along cross-veins (Figs. A–B, J). Face often with black areas and with fastigium frontis laterally bordered by distinct furrows (Figs. 26B, G, 27C). Male ninth abdominal tergite more gradually down-curved, furrowed and in apical area with a pair of oval swellings that carry at ventral tip a styliform, upright, blunt or subacute projection (Figs. 2 6 C–E, M–N, 27E–F). Female seventh abdominal sternite often with apical projection or a transverse fold (Figs. 26H, K, 27G, L, 28 J– K). Female subgenital plate with bilobate or truncate apex (Figs. 26 H–I, K–L, 27G, L, 28K–J)...................................................................................................... Prosopogryllacris Karny, 1937 14. More or less brachypterous species with wings just covering abdomen or shorter. Fore wings stiff with orange, yellow, or black areas or with orange/yellow areas alternating with black areas except for transparent tip (Figs. 71 A–K); main veins strong, prominent, and apart from the two subcosta branches running parallel and equidistant, often without branching or only R or M branching (Fig. 1A).................................................................................. 15.
- Fully winged or brachypterous species, but fore wings always with clear branching (Figs. 1 E–G). If male ninth abdominal tergite of similar shape then usually long winged species with main veins of fore wings less prominent and hind wings often with dark bands along veinlets.............................................................................. 17.
15. Spines on ventral margins of fore and mid tibiae shorter than diameter of tibia (Figs. 92 B–C)........................................................................................ Zalarnaca subgenus Glolarnaca Gorochov, 2008
- Spines on ventral margins of fore and mid tibiae longer than diameter of tibia (Figs. 73A, D). Male ninth abdominal tergite globular, ventro-posterior area furrowed and provided with a pair of oval swellings that terminate at medio-internal end into a stout, elongate process with acute tip (Figs. 72 A–K). Female ovipositor falcate or elongate. Larnaca Walker, 1869 ...... 16.
16. Female ovipositor falcate (Figs. 73 A–B, D). Male ninth abdominal tergite as before...................................................................................................... Larnaca subgenus Larnaca Walker, 1869
- Female ovipositor elongate. Males unknown.......................... Larnaca subgenus Paralarnaca Gorochov, 2003
17. Fore wings with media in basal area free (usually in long winged species, Fig. 2H) or fused with radius (usually in small or very small species with reduced wing length, Fig. 2G). Males at apical margin of ninth abdominal tergite with series of spines or other rough protuberances that have their counterpart at dorsal margin of modified paraprocts and might work together as a pincers-like tool during mating. Female seventh abdominal sternite usually with a projection from hind margin often with bifid tip and covering base of a simple subgenital plate (Figs. 42G, I, 43 I–J). Female ovipositor elongate and moderately curved or straight and with rounded tip or upcurved and with subacute tip. Asarcogryllacris sensu lato ........................ 18.
- Fore wings with media anterior always free from base but may in the further course temporarily fuse for a short distance with radius and/or radius sector and separate again. Male and female abdominal segments different....................... 20.
18. Small species with reduced venation of wings. Brachypterous; tegmen often with M and CuA undivided, in few specimens CuA simply branched (Figs. 43A, G). Hind tibia on ventral side without pre-apical spine; hind femur rather short and strongly thickened in basal half (Fig. 43H). Ovipositor short, straight, tip obtuse (Fig. 43H). Male paraproctes prolonged and modified on top to form together with the obtuse projections of the ninth tergite a gripping device (Fig. 43 C–E).............................................................................. Asarcogryllacris subgenus Pseudasarca subgen. nov.
- Brachypterous or macropterous species; MP+CuA1 vein often branching from both, MA and CuA, in species with wider fore wings then dividing into MP and CuA1, in others it remains undivided. Hind tibia on ventral side with pre-apical spine; hind femur longer, not strongly thickened in basal half........................................................... 19.
19. Slightly brachypterous species; CuP and first analis strongly sinuate in middle (Figs. 42D, H, L). Ovipositor upcurved, tip subacute (Fig. 42K)......................................... Asarcogryllacris subgenus Pseudolarnaca Gorochov, 2005
- Brachypterous or macropterous species; CuP and first analis only faintly curved in middle, in some species almost straight (Figs. 40A, C–D, G). Ovipositor elongate or of medium length, slightly upcurved or substraight, tip obtuse or subacute (Figs. 41A, E, I, L)............................................. Asarcogryllacris subgenus Asarcogryllacris Karny, 1937
20. "The (mostly four) pairs of moveable thorns at fore and mid tibiae remarkably short" (Karny 1937), shorter than the diameter of the tibia (similar as in Figs. 92 B–C). Fore wings (Fig. 1G) light with darkened veins and veinlets at least in basal area. Rather large species with long wings surpassing midst of stretched hind tibia. Male ninth abdominal tergite with a pair of large spine-like projections at apical margin with spines crossing each other. Female ovipositor elongate..................................................................................................... Psilogryllacris Karny, 1937
- The four pairs of moveable thorns at the fore and mid tibiae of normal length or longer............................ 21.
21. Species with colored wings. Fore wings semi-transparent with black or at least distinctly darkened veins and veinlets or with pale and dark pattern or with dark spots, or fore wings fully red brown or suffused with orange brown, or fully black with light veins. Hind wings transparent with darkened veins and veinlets, with or without dark bands along veinlets; in few species hind wings fully suffused with red or yellow brown color and darker bands along veinlets or hind wings fully suffused with brown or black with light cross veins and slightly lighter spots in cells between veinlets. Often large and rather robust species, fully winged or brachypterous; also medium sized or smaller, but not very slender species. Genera with mostly large species are included in this path, although some of the species may have uncolored fore wings with only the veins darkened........ 2 2.
- Mostly unicolored species. Fore wings yellowish semi-transparent, some species light green when alive; veins and veinlets not or hardly darker than surface. Hind wings transparent colorless with veins and veinlets only slightly darker. Often smaller, slender species with long wings reaching or surpassing mid of hind tibia, but some genera include also brachypterous species................................................................................................... 33.
22. Medium sized brachypterous species. Fore wings with strongly reduced costal field, which forms a projecting rounded-triangular flap at base of anterior margin; without visible media anterior or this is fused with radius sector. Hind femur strongly thickened.......................................................................................... 23.
- Fore wings normally developed, without strong reduction of costal field, with visible media anterior.................. 24.
23. Fore wings strongly sclerotised, dark brown to black; CuA simply dividing into two veins behind basal third of fore wings (Fig. 87A). Hind wings brown with light oval spots along veinlets. Hind femur strongly thickened. Male subgenital plate with medial area strongly prolonged, surpassing tip of styli (Fig. 87D). Monotypic genus........... Siderogryllacris Karny, 1937
- Fore wings semi-transparent with black or dark brown, very thick veins and veinlets; hind wings with same pattern (Figs. 81 A–B, E). Fore wings without separate media, but media arises behind basal area from cubitus anterior and divides behind mid length into MA and MP, which in P. cyanipes fuse again to one vein (Figs 81B, E). Hind femur strongly thickened. Male subgenital plate with medial area prolonged but not reaching tip of styli; male ninth abdominal tergite with a pair of lateral spines, tenth abdominal tergite with a pair of central spines. Female ovipositor medium long, curved with acute tip....................................................................................... Phlebogryllacris Karny, 1937 24. Fore wings and hind wings with dark brown infumation, appearing almost black when closed (Figs. 74 J–K); veinlets and basal area of main veins of light color; hind wings with similar color pattern (Fig. 76K). Female ovipositor rather short, in basal area stout and faintly upcurved, afterwards valves narrow and little down-curved (Figs. 74J, 76M). (One species with three subspecies)......................................................................... Melanogryllacris Karny, 1937
- Fore wings not dark with light veins. Ovipositor not with narrowed valves behind base; straight or upcurved........... 25.
25. General color ivory white (or pale yellowish in preserved specimens) with black ornaments (Figs. 88A, D–E). Medium ocellus very large (Fig. 88G). Male ninth abdominal tergite forming a single unpaired huge process that is moveable against the globular and strongly down-curved eighth tergite (Fig. 89). Female seventh abdominal sternite unmodified; ovipositor long and straight with subacute or subobtuse tip (Figs. 88 H–I).................... Ultragryllacris Gorochov & Dawwrueng, 2015
- Different combination of characters. Male abdominal apex with one or two pairs of appendages or projections of different shapes............................................................................................ 26.
26. Male ninth abdominal tergite prolonged but apically not down-curved, instead a little narrowed and terminating into a short convex lobe; somewhat before hind margin on both sides with a small obtuse lamella. Tenth abdominal tergite divided in middle; medial margins bulging with obtuse cone at tip (pointing up in preparation) and below that cone with a stout down-curved spine (Figs. 44 D–I). Male subgenital plate divided nearly from base into one short medial and two long lateral lobes; without styli (Fig. 44H). Fore wings and hind wings with cross-veins bordered on both sides by dark bands (Fig. 44A). Last two spines on ventral margin of hind femur markedly enlarged (Fig. 44C). Female subgenital plate with apex bilobate............................................................................................ Australogryllacris Karny, 1937
- Male abdominal apex with paired appendages or projections from ninth and/or tenth abdominal tergite. Male subgenital plate not divided from base but provided with styli, or divided behind base into two lobes with spined tips. Spines on ventral margin of hind femur of normal size........................................................................... 27.
27. Male abdominal apex with two pairs of projections from ninth and from tenth abdominal tergites. Fore wings surpassing middle of hind tibia, colorless or little yellowish brown along midline with markedly darkened veins and veinlets (Figs. 56–57, 91 A–F)........................................................................................... 28.
- Male abdominal apex with only one pair of apical projections................................................ 29.
28. Male ninth abdominal tergite excised in middle of apical margin giving room for the upright, acute projections of the tenth tergite; from both sides of excision with a projection that is wide at base than narrowed and terminating into a compressed acute apical area of various shape, pointing more or less ventrad (Figs. 56–57), sometimes apicad (Figs. 56 D–E), rarely reduced to a pair of short spines (Fig. 57G). Phallus with an unpaired, in lateral view usually sinuate sclerite enclosed in a membranous vesicula, in some species with more extended sclerotization (Fig. 58). Female seventh abdominal sternite and subgenital plate often for a larger part membranous, at least both separated by a distinct membranous zone. Ovipositor long, substraight or very little upcurved (Fig. 59)........................................ Diaphanogryllacris Karny, 1937
- Male ninth abdominal tergite with long, down-curved, sinuate, compressed projections with recurved, acute tip; remnants of tenth abdominal tergite with even longer, rounded, about upright projections with compressed, stiffened tip (Figs. 91 A–D). Phallus membranous. Female seventh abdominal sternite longer than preceding sternites with widened tip; subgenital plate membranous except for rim (Figs. 91 G–H). Ovipositor elongate, little upcurved with subobtuse tip (Figs. 90C, H)...................................................................................... Woznessenskia Gorochov, 2002
29. Brachypterous species with distinct venation (Figs. 45B, D). Male ninth abdominal tergite divided into two lateral parts that are separated by a broad prolongation of eighth tergite and probably moveable against this prolongation; on both sides of ninth tergite with an oval swelling that terminates into a rather large hook-shaped process (Figs. 45 E–F). Male subgenital plate rather short, bilobate, with robust styli longer than length of subgenital plate............... Brachyntheisogryllacris Karny, 1937
- Male ninth abdominal tergite and subgenital plate of different shape........................................... 30.
30. Head wide and robust; face with mat or subrugose surface; median ocellus rather large. Fore wings semi-transparent with distinct dark cross-veins (Figs. 66A, D, F, H). Male ninth abdominal tergite with a pair of short acute or subacute teeth at apical margin (Figs.67 D–E, G–H). Ovipositor elongate, straight or slightly upcurved; tip simple, subobtuse (Fig. 67A).......................................................................................... Haplogryllacris Karny, 1937
- Head not as wide; face nearly smooth with impressed dots and fine transverse riffles.............................. 31.
31. Small to medium sized species with wings little longer than covering abdomen (Figs. 76 C–E). Fore wings in some species yellowish brown with light margin and dark veins (Fig. 76C), in other species semi-transparent. Male ninth abdominal tergite at ventro-apical margin with a pair of band-shaped appendages clinging to tergite and terminating in middle of margin into downcurved hooks or upcurved sinuate spines (Figs. 77A, D–F, I). Female ovipositor short, straight with obtuse or faintly upcurved tip or very long and straight (Figs. 76B, G, J). Metriogryllacris for subgenera see................................. 44.
- Medium sized to large species. Fore wings often yellowish or light brown with dark veins, or transparent with a black area near base that encircles a light spot, or with extended coloration; hind wings transparent with pale or with black veinlets, often veinlets bordered by dark (black, red or brown) bands on both sides that can be extended to hind wings nearly fully dark red or brown (Figs. 46–47). Male ninth abdominal tergite in apical area furrowed in midline and on both sides with oval swelling that terminates on ventromedial, internal side into a stout acute process of varying size and shape; the spines of both sides often crossing each other when at rest (Figs. 49–50). Female subgenital plate with a stiffened pit at very base and/or with an extended preceding membranous area (Fig. 51 above). Ovipositor elongate, straight or slightly curved; tip simple, subacute or subobtuse (Fig. 51 below). Capnogryllacris Karny, 1937 .................................................... 32.
32. Median ocellus very large (Fig. 52J). Apical spines of male ninth abdominal tergite more gracile, thinner, and curved at tip (Fig. 52I, but variation unknown). Ovipositor elongate but curved; fore wings subtransparent with dark and yellow veins or with conspicuous dark bands along cross-veins in anterior area (Figs. 52 K–L)............................................................................................... Capnogryllacris subgenus Dictyogryllacris Karny, 1937 - Median ocellus of normal size (Fig. 48). Spines of male ninth abdominal tergite robust to very robust and usually tapering towards tip (Figs. 49–50). Female seventh abdominal sternite without process from hind margin (Fig. 51 above). Ovipositor elongate, straight or slightly curved (Fig. 51 below).............. Capnogryllacris subgenus Capnogryllacris Karny, 1937
33. Minute species (length of females with wings 14–15 mm). Tegmen without free media, instead media fused at base with cubitus anterior that divides in a first step into M+CuA1 and CuA2, in a second step the anterior branch then divides into M and CuA1 (Fig. 79M). Female subgenital plate membranous with a large transverse groove in basal area (Fig. 78K); ovipositor strongly upcurved in subbasal area then nearly substraight but curved faintly again before tip; tip subacute (Fig. 78L)........................................................................................ Minigryllacris gen. nov.
- Different combination of characters..................................................................... 34.
34. Hind tibia at dorsal-internal or on both dorsal margins with a large hooked spine (Figs. 36A, D–E). Brachypterous species with wings about covering abdomen; MP arises from MA (Fig. 36A, only 2 specimens seen). Male ninth abdominal tergite with a pair of simple teeth at tip. Female ovipositor elongate, little curved, tip subacute (Fig. 36E)..... Ancistrogera Brunner, 1888
- Hind tibia on both dorsal margins with normal spines....................................................... 35.
35. Hind tibia on dorsal internal margin with enlarged and little curved apical spine (Figs. 34C, F–H). Male abdominal apex with projections from ninth and from tenth abdominal tergites, in few species only from ninth or only from tenth tergite (Fig. 33). Male subgenital plate without moveable styli (Figs. 33C, E, I). Female subgenital plate with extended membranous area (Figs. 34B, E). Ovipositor of medium length, straight, tip subacute (Figs. 34A, D)............... Aancistroger Bei-Bienko, 1957
- Hind tibia with apical spine on dorsal internal margin usually larger than on dorsal external margin but less prolonged and not curved. Male subgenital plate with or without moveable styli................................................. 36.
36. "Male last abdominal tergite globular, strongly descending posteriorly; at bottom end terminating into two projections, that are touching each other close to median line, then semi-circularly projecting backwards and terminating into two strongly sclerotised thorns that embrace with their ends the distal part of the subgenital plate" (after Karny 1937). One slender species with long wings (tegmen 34 mm, hind femur 13.5 mm after Karny 1929a). Solomon Islands. Female unknown............................................................................................. Heterogryllacris Karny, 1937
- Male abdominal apex of different shape.................................................................. 37.
37. Male abdominal apex with appendages only at ninth abdominal tergite. Small to medium sized, fully winged to brachypterous species with wings covering about abdomen. Fore wings with MP branching from MA, less commonly from common stem with CuA.......................................................................................... 38.
- Male abdominal apex with appendages at tenth abdominal tergite or at ninth and at tenth abdominal tergite. Male subgenital plate with or without moveable styli. Small to medium sized, brachypterous to long winged species; wing length varying from about covering abdomen to surpassing midst of stretched hind tibia............................................ 47.
38. Small to medium sized species; often fully winged with fore wings surpassing mid of hind tibia, more rarely slightly brachypterous species; MP of fore wings usually arises from common base with CuA, then divides into two branches, MP and CuA1 (Fig. 82 above, exceptions occur). Male ninth abdominal tergite roughly globular, rather variable between species but always with a pair of projections with hook-like tip from apical margin; in some species outlined by a furrow from tergite with hooks meeting in midline, in others more or less projecting from apical margin or even as (tips of) long projections from lateral margins (Figs. 83 A–B, F, J). Male subgenital plate variable, often with bilobate projection between bases of styli or apical margin simply rounded; styli present and stout, or reduced and minute or completely absent (Figs. 83C, G, K). Female ovipositor in many species elongate, moderately curved with tip subobtuse or subacute; in other species of medium length or even short with obtuse tip (Figs. 82B, E, I)..................................................... Phryganogryllacris Karny, 1937
- Male ninth abdominal tergite and its appendages of different shape. Small to medium sized, brachypterous species with wings covering about abdomen. Fore wings with MP branching from MA, less commonly from common stem with CuA. Male subgenital plate with moveable styli........................................................................ 39.
39. Rather short and little stout species with wings covering about abdomen or little shorter. Male ninth abdominal tergite with medial furrow, more or less swollen on both sides of furrow especially in apical area, with swollen areas terminating at apical margin into cones or spines that cross each other in midline. Female ovipositor elongate and curved or short and sickle-shaped.................................................................................................. 40.
- Male ninth abdominal tergite different. Female ovipositor variable............................................. 42.
40. Male eighth abdominal tergite strongly prolonged and globular; ninth abdominal tergite bent proximad in subbasal area, in lateral view it seems to lie beneath eighth tergite; in ventro-apical view deeply furrowed in midline by a membranous zone and with strong oval swellings on both sides that are prolonged into a setose prolongation with strong teeth at tip pointing obliquely proximad (Figs. 52 D–F). Females at seventh abdominal sternite with a cylindrical prolongation that covers base of subgenital plate or longer................................................................. Celebogryllacris Karny, 1937
- Male eighth abdominal tergite less prolonged, not globular; ninth abdominal tergite with longer unfurrowed anterior area. Female seventh abdominal sternite without apical appendage................................................. 41.
41. Male ninth abdominal tergite with posterior area strongly bent down, in lateral view appearing truncate; apical surface with deep furrow, later split and terminating on both sides of divided area into a horizontal, spine-like projection (Figs. 38 D–G). Male subgenital plate with bilobate projection between bases of styli not reaching tip of styli (Fig. 38F). Female ovipositor elongate, slightly upcurved, tip subacute or subobtuse (Fig. 38I)......................... Aphanogryllacris Karny, 1937
– Male ninth abdominal tergite with dorsal surface gradually sloping, posterior area strongly bent proximad, becoming furrowed and membranous in midline, lateral areas swollen, on both sides of furrow with apical projection of different shape (Figs. 35K, J). Subgenital plate with styli very short, in type species hardly longer than wide (Figs. 35K, J). Female ovipositor elongate, slightly upcurved, tip subacute or subobtuse (Figs. 35B, E, N)............................ Anancistrogera Karny, 1937 42. Fore wings not or very little overlapping when at rest (Figs. 37 B–C). Male ninth abdominal tergite with dorsal surface flat and sloping posteriorly, at both lateral angles of apical margin with a short conical projection, carrying at tip a sickle-shaped spine (Figs. 37 J–K). Female seventh abdominal sternite with apical lobe separated by a furrow from sternite and overlapping base of membranous basal area of subgenital plate (Figs. 37 E–G). Female ovipositor elongate, hardly upcurved, tip subacute or subobtuse.......................................................................... Angustogryllacris gen. nov.
- Fore wings overlapping when at rest..................................................................... 43.
43. Male ninth abdominal tergite globular, at ventro-apical margin with a pair of band-shaped appendages clinging to tergite and terminating in middle in down-curved hooks or in narrow sinuate upcurved projections (Figs. 77A, D–F, I). Female ovipositor straight, short or very long. Metriogryllacris .............................................................. 44.
- Different combination of characters..................................................................... 45.
44. Male ninth abdominal tergite at ventro-apical margin with a pair of band-shaped appendages clinging to tergite and terminating in middle in down-curved hooks (Figs. 77A, D–F). Female ovipositor short, straight with obtuse or faintly upcurved tip (Figs. 76B, G)................................................. Metriogryllacris subgenus Metriogryllacris Karny, 1937
- Male ninth abdominal tergite at ventro-apical margin with a pair of upcurved sinuate spines (Figs. 77 I). Female ovipositor very long and straight (Fig. 76J)................................ Metriogryllacris subgenus Xiphilarnaca subgen. nov.
45. Male ninth abdominal tergite with one pair of processes posteriorly and one pair of small lobate prominences below them; these prominences directed forward or and partly downward, each bearing a small spine at or near apex; paraproctes of male simple, without any processes; male subgenital plate transverse, with styli. Female subgenital plate (at least in type species) almost triangular, with a transverse fold at base ventrally and with characteristic median vulva between its base and seventh abdominal sternite (from Gorochov 2008). Female ovipositor sickle-shaped, short and strongly upcurved, strongly narrowing from very wide in subbasal area to acute tip.......................................... Dinolarnaca Gorochov, 2008
- Different combination of characters..................................................................... 46.
46. Male ninth abdominal tergite with long and thin down-curved projections with sinuate apical area. Epiproct with a pair of long curved projections from base, each terminating into an acute tooth. Female subgenital plate largely membranous except for marginal areas. Ovipositor short, strongly curved, tip subobtuse......................... Microlarnaca Gorochov, 2004
- Small sized species (12.5–20 mm after Hugel 2009) with fore wings covering about abdomen; MA free throughout or arising free at (very) base but then fused for a distance with R (male) or RS (female of N. polygramma), MP arising from common base with CuA. Male abdominal apex with short cerci hardly longer than styli; subgenital plate with hind margin prolonged, almost reaching tip of styli. Female seventh abdominal sternite with a projection from hind margin (similar to the condition in Eugryllacris from which it however differs by the free MA and the ovipositor without apical widening). Female subgenital plate simple with sub-truncate apical margin. Ovipositor varying, moderately long, little upcurved, in New Guinean species substraight with tip subacute. New Guinea and Pacific Islands.......................... Nannogryllacris * Karny, 1937
* This key is restricted to New Guinean species of the genus; see Hugel (2009) for the distinction of the Pacific species and for differentiation against the Pacific genus Amphibologryllacris. Regarding the New Guinean species, only for N. polygramma the male is known. The type species, Nannogryllacris exigua (Brunner von Wattenwyl, 1888) from New Caledonia, was described from a single specimen without abdomen.
47. Long winged and brachypterous species (Fig. 63). Male subgenital plate considerably prolonged between bases of styli, prolonged area divided into two lobes and largely surpassing tip of styli. Male abdominal tergites nine and ten both with a pair of short spines, acute teeth or other short projections at tip (Fig. 64). Female ovipositor of medium length, substraight or in some species faintly up- or faintly down-curved, tip subacute (Figs. 65A, E, H)................... Furcilarnaca Gorochov, 2004
- Male subgenital plate with central area not projecting beyond tip of styli........................................ 48.
48. Small to medium sized, fully winged species with wings little surpassing hind knees but not fully reaching mid of stretched hind tibia, but also brachypterous and apterous species (Figs. 68A, E, H). Male ninth abdominal tergite without processes; tenth abdominal tergite narrow, with a pair of long projections often folded when at rest, with flexible basal and sclerotised apical areas (Figs. 69 A–F). Ovipositor elongate, longer than hind femur, almost straight; tip subobtuse (Fig. 68D)............................................................................................ Homogryllacris Liu 2007
- Male at abdominal apex with projections from ninth and from tenth tergites..................................... 49.
49. Male ninth abdominal tergite globular with a short unpaired lobe or protrusion in middle of posterior surface, if less expressed then with concave impression of surface; tenth abdominal tergite with pairs of spines, hooks or compressed projections....................................................................................................... 50.
- Male ninth and tenth abdominal tergites both with pairs of projections.......................................... 51.
50. Male ninth abdominal tergite globular with a short lobe in middle of hind margin; tenth abdominal tergite with a pair of upright hook-like projections in middle. Male subgenital plate with apical margin projecting between bases of styli but not reaching tip of styli. Female ovipositor elongate, faintly curved, in middle substraight, at tip upcurved and subobtuse............................................................................................... Dialarnaca Gorochov, 2005
- Medium sized species, fore wings covering about abdomen; MP of fore wings usually arises from a fusion of short branches from MA and CuA that shortly after divides into MP and CuA1, but there is some variation between specimens (Figs. 85 A–E). Male ninth abdominal tergite globular with a rugose obtuse lobe in middle of apical surface, followed on ventral side by a furrow; tenth abdominal tergite with a pair of projections in middle that point mediad and are provided with widened tip that carries on internal side a stiffened black rim, thus that the projections from both sides together form a pincers like tool (Figs. 86 A– E). Female sixth abdominal sternite with a short apical projection; seventh sternite little prolonged, at end somewhat depressed (Figs. 86 G–I). Ovipositor elongate, straight, and rather stout; tip subobtuse (Fig. 85I)............. Siamgryllacris gen. nov. 51. Male ninth abdominal tergite with one pair of approached projections carrying small hooks directed medially touching (but not crossing) one another; tenth tergite narrow transverse, in middle with a pair of digitate processes pressed to each other, directed upwards, and partly laterally overlapped by the projections of ninth tergite, this looks as though the projections of the tenth tergite are framed or encircled by the projections of the ninth tergite (Figs. 93E, L). Male subgenital plate with styli inserted at apical-lateral angles. Female seventh abdominal sternite with a process directed backwards, in some species only indicated; subgenital plate with apical margin more or less emarginated. Genus Zalarnaca ................................... 52.
- Male abdominal apex with projections from tenth tergite not framed or encircled by projections of the ninth tergite...... 53.
52. Rather elongate species with long spines on fore and mid tibiae (Fig. 92H). "Ovipositor well developed, with narrowly rounded apex slightly deflexed upwards." (Gorochov 2005).............. Zalarnaca subgenus Zalarnaca Gorochov, 2005
- Rather small species of variegated coloration. Head wide, fastigium frontis about 2.1–2.3 times as wide as scapus (Figs. 92C, 93I). Legs rather short; the four pairs of moveable spines on fore and mid tibiae short (longest 0.5–0.8 mm, Fig. 92B); hind femur stout (2.7–3.1 times as long as wide); hind tibia slightly arcuate, with sparse, very small dorsal denticles (Fig. 92B). Ovipositor rather short, upcurved (but not sickle-shaped), tip subacute or subobtuse (Fig. 92F)......................................................................................... Zalarnaca subgenus Glolarnaca Gorochov, 2008
53. Brachypterous, wings covering about abdomen. Male ninth abdominal tergite globular with one pair of small, (immovable?) hooks pointing downwards; tenth abdominal tergite "narrow, divided into 2 transverse and typically curved lateral sclerites each bearing in median part long and heavily sclerotized spine directed upwards; male genitalia membranous" (Gorochov 2005)......................................................................... Idiolarnaca Gorochov, 2005
- Small to medium sized, fully winged to little brachypterous species; MP undivided, arising from MA and CuA (in type species after Fig. 2 in Karny 1937 and in N. dravida after Fig. 1 in Karny 1929c) or reaching about middle of stretched hind tibia. Male abdominal apex with projections from ninth and tenth tergites (in N. barkudensis after drawing in Chopard 1924) or with a pair of tubercular extensions on hind margin of ninth abdominal tergite and a pair of hook-shaped projections with widened base from tenth abdominal tergite (in N. succinea, Fig. 70 N–P). (Insufficiently defined heterogenous genus, in need of revision)............................................................................... Niphetogryllacris Karny, 1937
54. Strongly brachypterous to micropterous species. Fore wings reaching at least the first abdominal segments but are distinctly shorter than half the length of abdomen; often still overlapping in midline or almost so............................ 55.
- Micropterous or apterous species. Fore wings reduced to lateral scales with or without venation, usually not surpassing hind margin of metanotum or completely missing.............................................................. 57.
55. Fore wings reaching tip of second abdominal segment (Fig. 80E, one long winged species also exist). Male ninth abdominal tergite little narrowing towards down-curved apex; at apical margin with a pair of long, little curved, acute spines from lateral angles (Fig. 80I). Male subgenital plate with bilobate apex; styli inserted at rounded apico-lateral angles (Fig. 80K). Female ovipositor wide at base, afterwards strongly upcurved, tip subacute (Fig. 80G)............... Neolarnaca Gorochov, 2004
- Male abdominal apex with projections from tenth abdominal tergite or ninth abdominal tergite replaced along midline by a long projection of eighth tergite........................................................................ 56.
56. Fore wings micropterous, lateral and reaching only first abdominal tergite, in other species reaching mid length of abdomen (species with longer wings also exist). Male tenth abdominal tergite with a pair of long, compressed, apically acute projections surpassing beyond tip of ninth tergite (Figs. 70 A–B, I, K, M). Subgenital plate at apex bilobate, styli rather small, sitting at tips of apical lobes. Female subgenital plate with bilobate apical margin (Figs. 70C, J).............. Homogryllacris Liu, 2007
- Males with a pair of projections only from ninth abdominal tergite. Fore wings of male almost reaching middle of abdomen, in female shorter (Figs. 45B, D). Male ninth abdominal tergite divided into two lateral parts that are separated by a broad prolongation of eighth tergite and probably moveable against this prolongation; on both sides of ninth tergite with an oval swelling that terminates into a rather large hook-shaped process (Figs. 45 E–F). Male subgenital plate rather short, bilobate, with robust styli longer than length of subgenital plate (Fig. 45E)............................. Brachyntheisogryllacris Karny, 1937
57. Micropterous; fore wings surpassing hind margin of metanotum............................................... 58.
- Squamipterous, remnants of fore wings hardly surpassing hind margin of mesonotum, or apterous‚ without remnants of wings.................................................................................................. 59.
58. Micropterous, fore wings roughly semicircular, little surpassing hind margin of metanotum. Moveable spines on fore and mid tibiae rather short, not longer than tibia wide. Ovipositor almost as long as hind femur, little curved, more strongly curved in subbasal area, tip subobtuse........................................................ Celeboneanias Karny, 1937
- Micropterous or squamipterous. Female ovipositor short and straight with wide obtuse tip with dorsal margin faintly upcurved. Male ninth abdominal tergite globular with a pair of curved acute spines / hooks at ventro-apical margin. Male subgenital plate with apical margin little projecting between bases of styli and with broadly sub-truncate tip. Female seventh sternite with a pair of pits; subgenital plate with ventral projections. (for identification of species see Kim, Jeong & Shim (2014) and Gorochov 2002b).............................................................. Metriogryllacris Karny, 1937 sensu stricto
59. Squamipterous species, fore wings often lanceolate, hardly surpassing hind margin of mesonotum. The four pairs of moveable spines on fore and mid tibiae of normal size............................................................... 60.
- Apterous, without traces of wings....................................................................... 62.
60. Fore wings reduced to small lateral scales (Figs. 74 F–G); type species sub-apterous (Fig. 74I). Male ninth abdominal tergite deep and long excised from ventral hind margin in which a pair of upright projections from tenth tergite finds room (type species, Fig. 75K) or ninth tergite restricted to dorsal area leaving the posterior surface open in which a pair sclerotised plates (of tenth tergite?) give rise to large curved spines (fruhstorferi group, Fig. 75F). Male subgenital plate wider than long with apical margin concave or with bilobate apical margin (Figs. 75B, F, J). Female (of type species unknown) with seventh sternite unmodified; subgenital plate simply rounded with fine concavity at tip. Ovipositor short, distinctly upcurved, tip subobtuse (fruhstorferi group Fig. 75E) or long and straight (M. harmandi Fig. 75P, M. henry). (The genus is a heterogenous assemblage of nearly apterous (type species), apterous (M. henry), and squamipterous (fruhstorferi group) species and in need of revision)................................................................................ Melaneremus Karny, 1937 - Different combination of characters..................................................................... 61.
61. Male ninth abdominal tergite globular, ventro-posterior area furrowed and provided with a pair of oval swellings that terminate at medio-internal end into a stout, elongate process with acute tip (Figs. 72 I–J). Female ovipositor moderately short and strongly curved or elongate, tip unmodified (Fig. 73G). Mostly species with wings covering abdomen but with strongly brachypterous (with fore wings covering less than half of abdomen but still overlapping in midline), sub-micropterous (with fore wings still touching each other on back), or sub-apterous (fore wing reduced to a tiny lateral scale) (Figs. 71 L–M, O, 73G). As in the fully winged species of the genus, species with strongly shortened wings have fore wings stiffened with parallel veins............................................................ genus Larnaca Walker, 1869 (for subgenera see 16.).
- Male eighth abdominal tergite prolonged, together with ninth tergite globular (Fig. 78B); ninth abdominal tergite lying on posterior-ventral side of eighth tergite, with a pair of transverse oval swellings before apical margin, little excised in middle and at ventral margin with a pair of short acute teeth somewhat away from midline (Fig. 79B). Male subgenital plate wider than long with apical margin convex; styli about as long as subgenital plate. Female subgenital plate of type species simply rounded (Fig. 79B). Ovipositor elongate, moderately curved up (Fig. 78A). Females of species currently included in Neanias have long or short, straight or upcurved ovipositors. Squamipterous. In the type species fore wings lanceolate, not reaching tip of metanotum, some longitudinal veins visible..................................................... Neanias Brunner, 1888
62. Apterous (there are also fully winged species in the genus). Male abdominal tergites eight and nine prolonged but without projections. Tenth abdominal tergite with a pair of long or very long tube-like projections crossing each other behind base, for the greatest part membranous and flexible, in apical area sclerotised, with acute tip (similar as in Figs. 69B, F). Female ovipositor elongate, substraight, tip little upcurved, subacute (Fig. 68D)............................... Homogryllacris Liu, 2007
- Male abdominal apex without flexible projection from tenth tergite............................................ 63.
63. Hind tibia on dorso-internal margin with only one large spine, in some species plus one small spine; on external margin with normal spines. Hind femur rather stout. Male ninth abdominal tergite with curved acute projections from apico-lateral margins. Male subgenital plate little bi-lobate projecting between bases of short styli. Female subgenital plate rather short, apex widely sub-truncate. Ovipositor elongate but not very long, tip subobtuse........................ Nippancistroger Griffini, 1913
- Hind tibia with several normal spines on dorso-internal margin............................................... 64.
64. Male subgenital plate with styli. Female ovipositor long, straight or slightly curved............................... 65.
- Male subgenital plate in posterior area divided into two narrowing lobes but without distinct styli. Ovipositor short, upcurved, tip obtuse.......................................................................................... 66.
65. Medium large species. Apterous. Face rather wide and little rugulose (Figs. 61A, C, G, L). Male ninth abdominal tergite at apical margin with a pair of rather short stout acute teeth in middle (Figs. 61B, K, M, 62 E–F). Male subgenital plate wider than long, with short styli, apical margin between styli little angular-concave (Figs. 61B, E, M, 62I). Legs normal. Female seventh abdominal sternite often with a pit before hind margin (Figs. 61D, F, N, 62K); subgenital plate often elongate with rounded margin (type species) or triangular with tip rounded with short lateral lobes. Ovipositor long, slightly curved, tip subacute (Figs. 60 E–F, H–J, 62L)............................................................... Eremus Brunner, 1888
- Medium large species (Figs. 78 E–G). Apterous. Female seventh abdominal sternite depressed before hind margin and then prolonged into a rounded-triangular process; subgenital plate about semicircular with depressed surface and bulging margin, which forms at both sides of base a droopy clubbed process (Figs. 79 I–K)........................ Monseremus gen. nov.
66. Hind tibia straight. Small species. Apterous. Face rather wide. Male ninth abdominal tergite furrowed in midline, then divided into a pair of long and down-curved and often at tip recurved apical lobes. Male subgenital plate in posterior area divided into two narrowing lobes but without distinct styli. Female subgenital plate simple. Ovipositor short, upcurved, tip obtuse..................................................................... Apotrechus Brunner, 1888 (Chinese species only)
- Hind tibia in males distinctly curved, in females little curved (Figs. 39 A–C). Small species. Apterous. Face not peculiar wide (Fig. 39G). Male ninth abdominal tergite furrowed in midline, then divided into a pair of long and down-curved and often at tip recurved apical lobes (Figs. 39 E–F). Male subgenital plate in posterior area divided into two narrowing lobes but without distinct styli. Female subgenital plate simple. Ovipositor short, upcurved, tip obtuse (Fig. 39D). Hind femur on ventro-external margin in males with (one or two) greatly enlarged spines in apical area (sometimes also with one enlarged spine on ventrointernal margin), in females less enlarged (Fig. 39D).................................. Apterolarnaca Gorochov, 2004
67. Ventral margins of fore and mid-tibiae with only two pairs of short spines. Male unknown (monotypic genus)........................................................................................... Glenogryllacris Karny, 1930
- Ventral margins of fore and mid-tibiae with three pairs of long spines plus one pair of apical spurs (Fig. 2B). Male subgenital plate with a medial prolongation with bifid tip............................................................. 68.
68. Apterous. Fore and mid tibiae with only three pairs of rather short moveable spines on ventral margins plus one pair of apical spines (Fig. 98C). Hind femur rather stout, about 2.5x longer than wide, with narrow apical area very short (Fig. 98B). Hind tibia without pre-apical spine on ventral margin. Ovipositor long and straight (except at base), tip subacute. Seventh abdominal sclerite with median carina, with a small sclerite in following intersegmental membrane; subgenital plate membranous with rounded apical margin. (Fig. 98 E–G). Oriental Region....................................... Paraneanias gen. nov.
- Fully winged or micropterous. Papuan Region............................................................. 69.
69. Male ninth abdominal tergite in dorsal view only little longer than preceding tergites; apical margin with down-curved restiform projection with bifid tip in middle and with very long little upcurved spine-like projections from ventro-lateral angles. Remnants of tenth tergite terminating into long, twisted, acute projections (Figs. 99 C–F). Female unknown................................................................................................ Plexigryllacris gen. nov.
- Male ninth abdominal tergite globular. With a pair of short projections from below ninth tergite..................... 70. 70. Fully winged species (Fig. 94)................................................... Papuogryllacris Griffini, 1909f - Micropterous species; tegmina reduced to short lateral lobes (Fig. 97)...................... Papuoneanias Karny, 1929a
description: Published as part of Ingrisch, Sigfrid, 2018, New taxa and records of Gryllacrididae (Orthoptera, Stenopelmatoidea) from South East Asia and New Guinea with a key to the genera, pp. 1-278 in Zootaxa 4510 (1) on pages 16-24, DOI: 10.11646/zootaxa.4510.1.1, http://zenodo.org/record/10072806