New species and records of Zebragryllus Desutter-Grandcolas & Cadena-Castañeda, 2014 (Orthoptera: Gryllidae: Gryllinae) from the Brazilian Amazon rainforest Tavares, Gustavo Costa Oya, Beatriz Harumi Kondo Cadena-Castañeda, Oscar J. Oliveira, Marcus Paulo Alves de Castro-Souza, Rodrigo Antônio European Journal of Taxonomy 2024 2024-04-24 932 82 111 Tavares, Oya & Cadena-Castaneda Tavares, Oya & Cadena-Castaneda 2024 [401,1088,1364,1391] Insecta Gryllidae Zebragryllus Animalia Orthoptera 4 86 Arthropoda species aphonus sp. nov.   urn:lsid:zoobank.org:act: C2212D6F-194D-49C7-BA27-B113E0B51C55  Figs 1–5, 16A–C    Diagnosis Specimens dark ( Figs 1A–B, 4A–B). Tympana reduced or absent ( Fig. 2A–F). All femora entirely ochre, without the ‘zebra’ pattern ( Fig. 2A–C, G–K). Males dorsally with head, pronotum, tegmina, and first abdominal tergite black ( Fig. 1A–E); without stridulatory apparatus ( Fig. 1E). Comparatively, all the other known species of  Zebragryllusare easily distinguished for having stridulatory apparatus on the male tegmina. Male genitalia: pseudepiphallic median lophi elongated, conspicuously produced hind- and upwards, very narrow, and apically acute, ventrally with a sharp keel. Pseudepiphallic lateral lophi acute. Pseudepiphallic paramere surpassing the lateral lophi; in ventral view, slightly arched. Ectophallic fold very thin, not reaching the level of the pseudepiphallic paramere apex ( Fig. 3A–D). Female with the black coloration extending till the second abdominal tergite ( Fig. 4A–D); remaining tergites dark brown; ovipositor straight and almost as long as the hind femora ( Fig. 4A, H–I). Female genitalia: copulatory papilla with posterior margin of dorsal surface widely concave, and the anterior margin with medial process emarginated, flanking the spermatheca duct ( Fig. 4L); ventral surface acute anteriorly ( Fig. 4M); in lateral view, posterior portion taller than anterior one ( Fig. 4N).    Etymology The specific epithet “ aphonus” refers to the absence of a stridulatory apparatus, which makes the cricket incapable of producing sound.    Type material    Holotype BRAZIL• ♂; Pará, Cannã dos Carajás, S11C-0033 cave; 6°23′53.5″ S, 50°23′26.5″ W;  16 Aug. 2022; J.P. Alvesand D. Souzaleg.; except for the right foreleg, all the remaining legs have fallen but are stored in the same specimen vial, one palpus, the subgenital plate, the right tegmen, and the phallic complex were removed but kept in a microvial with the specimen; MPEG. HEX 05050607.    Paratypes BRAZIL• 1 ♀; same collection data as for holotype; except for the left hindleg, all the remaining legs have fallen but are stored in the same specimen vial, the subgenital plate, the seventh abdominal sternite, and the copulatory papilla were removed but kept in a microvial with the specimen; MPEG. HEX 05050608•  1 ♂; same collection data as for preceding; S11C-0066 cave; 6°23′53.5″ S, 50°22′58.1″ W;  9 Oct. 2022; except for the left foreleg and right hindleg, all the remaining legs have fallen but are stored in the same specimen vial; MPEG. HEX 05050609.   Other material   BRAZIL• 1 immature(sex unknown); same locality as for holotype; 6°23′53.5″ S, 50°23′26.5″ W;  15 Feb. 2023; J.P. Alvesand D. Souzaleg.; ISLA107969•  1 immature(sex unknown); same collection data as for preceding; S11C-0110 cave; 6°23′55.5″ S, 50°23′23.2″ W;  15 Feb. 2023; ISLA107970•  1 ♀, immature; same collection data as for preceding; S11C-0110 cave; 6°23′55.5″ S, 50°23′23.2″ W;  16 Aug. 2022; ISLA107971•  1 ♂, immature; same collection data as for preceding; S11C-0121 cave; 6°22′56.8″ S, 50°23′56.0″ W;  4 Aug. 2022; ISLA107972•  1 ♂, immature; same collection data as for preceding; S11C-0037 cave; 6°23′28.2″ S, 50°23′33.7″ W;  12 Aug. 2022; ISLA107973•  1 immature(sex unknown); same collection data as for preceding; S11C-0050 cave; 6°23′58.2″ S, 50°23′8.5″ W;  8 Aug. 2022; ISLA107974•  2 immatures(sex unknown); same collection data as for preceding; S11C-0092 cave; 6°24′7.2″ S, 50°23′18.7″ W;  17 Aug. 2022; ISLA107975 to 107976.    Description  Male  COLORATION. Specimens very dark ( Figs 1A–E, 4A–D). Cephalic capsule, pronotum, and tegmina black ( Fig. 1A–E). Abdominal tergites dark brown and posteriorly black ( Figs 1B, 4B), each with midline of black punctuations extending from lateral to the dorsolateral portion. Some whitish spots sometimes present on head dorsum ( Fig. 1B). Eyes black, ocelli whitish. Scapus and pedicel yellowish, flagellomeres brown. Clypeus and labrum dark brown. Mandibles proximally black and mediodistally reddish-brown ( Fig. 1C). Maxillary palpi dark brown, except for whitish ventral areas on mediodistal portion and apex ( Fig. 1G). Two first segments of labial palpi dark brown, and last ochre, with tip and ventrodistal portion whitish. Femora entirely ochre, and tibiae dark brown ( Fig. 2A–C, G–K) – fore tibia with two inner whitish circular areas on most proximal portion ( Fig. 2G). Ventral sclerites of thorax and abdomen ochre, except for brown sternite 8 and male subgenital plate, which are brown ( Fig. 1F, I).  HEAD. In frontal view ( Fig. 1C), semicircular, almost as wide as high, with eyes slightly produced laterally; ocelli circular and almost aligned to eyes dorsal margin – median ocellus slightly lower. Antennal pits located low on frons, near epistomal suture, and almost on same level as eyes’ most ventral border (these last also close to subgenal suture). Clypeus dorsal area notably separated from ventral area by clypeal suture. Ventral area conspicuously constricted laterally, almost half as wide as dorsal area, with median white stripe and transverse dark line on sides. Labrum circular, with pair of incomplete transverse mediolateral sutures. First two segments of maxillary palpi subequal in length; third and fourth as long as first and second together; fifth longer than any other segment, ventrally truncated on mediodistal area ( Fig. 1G). In lateral view ( Fig. 1D), head convex, with frons slightly tumescent; eyes ovoid, higher than wide. In dorsal view ( Fig. 1B), eyes protruding; fastigium subtriangular and apically convex. Scapus wide, almost as wide as half fastigium apex.   Fig. 1.  Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov., habitus. A–F, H–J. Paratype, ♂, (MPEG.HEX 05050609). G. Holotype, ♂ (MPEG.HEX 05050607). A. Lateral view. B. Dorsal view. C. Frons. D. Lateral view of head and thorax. E. Tegmina. F. Sternum. G. Maxillary palpi, outer view. H–J. Terminalia. H. Dorsal view. I. Lateral view. J. Ventral view.   Fig. 2.  Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov.legs, outer views of legs. A. ♂, paratype (MPEG.HEX 05050609), with reduced tympanum. B. ♀, paratype (MPEG.HEX 05050608), very reduced tympanum. C. ♂, holotype (MPEG.HEX 05050607), no tympanum. D. ♂, paratype (MPEG. HEX 05050609), detail of reduced tympanum. E. ♀, paratype (MPEG.HEX 05050608), detail of very reduced tympanum. F. ♂, holotype (MPEG.HEX 05050607), detail of the tibia with no tympanum. G–I. ♂, holotype (MPEG.HEX 05050607). G. Inner view of the fore leg. H. Outer view of the mid leg. I. Inner view of the mid leg. J–L. ♂, paratype (MPEG.HEX 05050609). J. Outer view of the hind leg. K. Inner view of the hind leg. L. First tarsus, in detail.  THORAX. Pronotum wider than long, bearing bristles on margins, with both anterior and posterior dorsal margins slightly concave ( Fig. 1B). Lateral lobes almost squared, with anterior and posterior margins almost straight and ventral margin obliquely straight ( Fig. 1A, D). Prosternum reduced, triangular, and continuous with conjoint cervical sclerites. Mesosternum sub-squared, with posterior margin slightly bilobed. Metasternum hexagonal, with posterior margin slightly incised ( Fig. 1F)  LEGS. All femora covered by fine pubescence and some longer bristles ( Fig. 2A–C, G–K). Fore and mid tibia with two apical spurs on outer side ( Fig. 2A–C, H) and one on inner side ( Fig. 2G, I). Tympana reduced ( Fig. 2A–B, D–E) or absent ( Fig. 2C, F) (different development levels can be found in same specimen). Hind tibia with 6–7 external and 5–6 internal dorsal spurs and three apical spurs on each side ( Fig. 2J–K); externally, dorsal and ventral apical spurs equal in length, mid one twice longer; internally, dorsal and mid spurs same-lengthened, ventral one at least three times smaller ( Fig. 2J–K). Fore and mid basitarsi covered by strong setae on ventral side ( Fig. 2A–C, G–I, L). Hind basitarsi ventrally with setae, dorsally with two rows of spines, and apically with two spurs, outer one smaller ( Fig. 2J–K).   Fig. 3.  Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov., ♂, holotype (MPEG. HEX 05050607), phallic complex. A. Dorsal view. B. Ventral view. C. Axial view. D. Lateral view. Abbreviations: see Material and methods.   Fig. 4.  Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov., paratype, ♀ (MPEG.HEX 05050608). A–B. Habitus. A. Lateral view. B. Dorsal view. C. Dorsal view of head and thorax. D. Tegmina. E. Supra-anal plate. F–G. Subgenital plate. F. Lateral view. G. Ventral view. H–I. Ovipositor. H. Lateral view. I. Dorsal view. J–K. Detail of the ovipositor apex. J. Lateral view. K. Dorsal view. L–M. Copulatory papilla. L. Dorsal view. M. Ventral view. N. Lateral view.  WINGS. Tegmina short, reaching up to the third tergite, without stridulatory apparatus or transverse veins; lateral field with six longitudinal veins ( Fig. 1A–B, D–E). Hind wings absent.   Fig. 5.  Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov., immatures. B. Immature (ISLA 107975). C. Immature (ISLA 107976). D. Immature (ISLA 107969). A. Alive. B–D. Alcoholpreserved.  ABDOMEN. Cylindrical, covered with bristles. Male supra-anal plate triangular, medially incised posteriorly and with transverse suture ( Fig. 1H). Subgenital plate slightly longer than wide, with widely rounded posterior margin ( Fig. 1I–J).  GENITALIA. Pseudepiphallic median lophi conspicuously elongated, notably produced hind- and upwards, very narrow, apically acute ( Fig. 3A–B, D), and ventrally with sharp keel ( Fig. 3C). Pseudepiphallic lateral lophi triangular, apically acute ( Fig. 3D). Pseudepiphallic parameres elongated, surpassing lateral lophi; in ventral view, slightly arched, with rounded apex ( Fig. 3B). Pseudepiphallic apodemes acute, incurved ( Fig. 3A). Ectophallic fold very thin, not reaching level of pseudepiphallic paramere apex ( Fig. 3B). Ectophallic arc very thin and notably curved; in dorsal view, forming ‘W’ with ectophallic apodemes ( Fig. 3A). Rami basally bifid and notably arched along its length, contiguous, and medially incised ( Fig. 3A–B).  Female Similar to males in very dark coloration, with head, pronotum, tegmina, and first two abdominal tergites blackish ( Fig. 4A–B). Tegmina reduced, reaching only first abdominal tergite, with only longitudinal veins ( Fig. 4C–D). Supra-anal plate notably triangular, with transverse suture ( Fig. 4E). Subgenital plate short; in ventral view, wider than long, with posterior margin slightly concave medially ( Fig. 4G); in lateral view, higher than long, with posterior margin truncated ( Fig. 4F). Ovipositor almost straight, with triangular apex. Dorsal valves dorsally sulcated ( Fig. 4H–K).  GENITALIA. Copulatory papilla with posterior margin of dorsal surface widely concave, and anterior margin with medial process emarginated, flanking spermatheca duct ( Fig. 4L); ventral surface acute anteriorly ( Fig. 4M); in lateral view, posterior portion taller than anterior one ( Fig. 4N).  Nymphs In life, bearing same black coloration as head and pronotum of adults, but abdomen and legs gray ( Fig. 5A). When preserved in alcohol, gray body parts become ochre ( Fig. 5B–D).  MEASUREMENTS(mm). ♂, holotype. TL: 11.0; LP: 2.0; WP: 3.0; Tg: 2.5; HF: 6.0; HT: 4.0. ♂, paratype. TL: 9.5; LP: 2.0; WP: 2.5; Tg: 3.0; HF: 7.0; HT: 4.0. ♀, paratype. TL: 9.0; LP: 2.0; WP: 3.0; Tg: 1.5; HF: 7.0; HT: 4.0; Ov: 6.5.    Remarks   Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov.has quite a unique feature. It is the only known species of the genus in which the male has no stridulatory apparatus. The different development levels of the tympana are also singular to this species. These two characteristics may be adaptations to the caves in which they were found. Still, it is hard to ensure since no phylogenetic study was made and ecological data are lacking. The shape of the pseudepiphallic median lophi is similar to that of  Z. nauta. However, it differs by the presence of conspicuous pseudepiphallic lateral lophi, which are very reduced and almost inconspicuous in  Z. nauta.These two species may be related. We believe this species is facultative to a cave-dwelling way of life or caviculous (sensu Desutter-Grandcolas et al.1998), foraging on forest ground during the nights and passing the day inside the caves or cavities at ground level. We believe in this assumption since adult and immature stages were found inside caves as field work was carried out during the day. This species may also inhabit or take refuge in pores or voids on the soil, characteristic microhabitats of the environment where it was collected. The structure of the landscape is known as Banded Ferruginous Formations, which are iron formations composed of alternated bands of jaspilite and ore bodies rich in iron. In this area, extensive iron ore plateaus are formed, comprising superficial ferrugineous breccia, also known as canga formation. A metallophilic savannah covers the canga formations on the plateaus and mountains, while typical tropical vegetation of the Amazon rainforest covers the slopes. The highly porous canga formations result from the removal of silica and carbonates due to the tropical rains, which causes the dissolution and chemical change found in iron ore, forming several interconnected small spaces in the canga formations – the voids ( Ferreira et al.2018).   Zebragryllus aphonusTavares, Oya & Cadena-Castañeda sp. nov.was collected in seven caves in the municipality of Canaã dos Carajás, located in the east region of Pará State, Brazil( Fig. 16A–C). The caves are coded as S11C-0033, S11C-0037, S11C-0050, S11C-0066, S11C-0092, S11C-0110, S11C-0121. They are situated in the Amazon rainforest and are part of the federal conservation unit “Floresta Nacional de Carajás” (FLONA) ( Carmo & Kamino 2015). They are located within the “Grão-Pará” geological group, known for its iron-ore lithology. This region is popularly known as “Serra dos Carajás”, which comprises vast plateaus separated by depressions in the landscape, dividing it into three regions: Serra Norte, Serra Leste, and Serra Sul ( Carmo & Kamino 2015). The seven caves where the specimens were collected are situated in the “Serra Sul” plateaus region ( Fig. 16). 4849264315 2022-08-16 MPEG J. P. Alves & D. Souza Brazil -6.3981943 S 11 C- 0033 cave 1 -50.390697 Canna dos Carajas 5 87 HEX 05050607 1 1 Para holotype 4849264301 2022-08-16 MPEG J. P. Alves & D. Souza Brazil -6.3981943 BRAZIL 1 -50.390697 Canna dos Carajas 5 87 HEX 05050608 1 1 Para paratype 4849264318 2022-10-09 MPEG J. P. Alves & D. Souza Brazil -6.3981943 S 11 C- 0066 cave 1 -50.382805 Canna dos Carajas 5 87 HEX 05050609 1 1 Para paratype 4849264308 2023-02-15 ISLA J. P. Alves & D. Souza Brazil -6.3981943 BRAZIL 1 -50.390697 Canna dos Carajas 5 87 ISLA107969 1 1 Para 4849264312 2023-02-15 ISLA J. P. Alves & D. Souza Brazil -6.39875 S 11 C- 0110 cave 1 -50.38978 Canna dos Carajas 5 87 ISLA107970 1 1 Para 4849264304 2022-08-16 ISLA J. P. Alves & D. Souza Brazil -6.39875 S 11 C- 0110 cave 1 -50.38978 Canna dos Carajas 5 87 ISLA107971 1 1 Para 4849264313 2022-08-04 ISLA J. P. Alves & D. Souza Brazil -6.3824444 S 11 C- 0121 cave 1 -50.39889 Canna dos Carajas 5 87 ISLA107972 1 1 Para 4849264307 2022-08-12 ISLA J. P. Alves & D. Souza Brazil -6.3911667 S 11 C- 0037 cave 1 -50.392696 Canna dos Carajas 5 87 ISLA107973 1 1 Para 4849264309 2022-08-08 ISLA J. P. Alves & D. Souza Brazil -6.3995 S 11 C- 0050 cave 1 -50.385696 Canna dos Carajas 5 87 ISLA107974 1 1 Para 4849264311 2022-08-17 ISLA J. P. Alves & D. Souza Brazil -6.402 S 11 C- 0092 cave 1 -50.38853 Canna dos Carajas 5 87 ISLA107975, ISLA107976 2 2 Para