taxonID	type	description	language	source
06AA0E4B88D25FF6968F05131E48747E.taxon	description	Figs 2 E, F, 3 A – C, 4 A – C, 5 A – G, 6 A – D, 7 A – D, 8 A – P, 9 A – F, 10 A – F, 11 A – D, 12 A – D	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
06AA0E4B88D25FF6968F05131E48747E.taxon	description	Description. Adults. Figs 2 E, F, 3 A – C, 4 A – C, 5 A – G, 6 A – D, 7 A – D, 8 A – P.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
06AA0E4B88D25FF6968F05131E48747E.taxon	diagnosis	Diagnosis. Habitus. Small, light-colored and strongly troglomorphic delphacid devoid of compound eyes and ocelli, tegmina strongly reduced in length without recognizable venation, hind wings reduced to narrow pads. Configuration of male and female genitalia as described for the genus. For details of male genitalia see below. Coloration (preserved specimen). As in Figs 3, 4, predominantly deep yellow (85) on head, thorax, abdomen and genitalia, some areas in pale yellow (89) principally legs, head laterally (gena) and spaces between segments, especially of the abdomen. Body length. Male. 1.6 – 1.8 mm (n = 7) (Fig. 3); Female. 1.9 – 2.0 mm (n = 2) (Fig. 4). Configuration, shape and proportions of head, thorax and female terminalia as proposed for the genus. Some figures and measurements are provided here. Head. Vertex (Figs 3 A, 5 A, 6 A): width (0.20) length (0.04). Frons (Figs 3 B, 5 B, 6 B, C): width (0.39) length (0.38). Antennae (Fig. 5 E): Pedicel width (0.09) length (0.13). Thorax. Pronotum (Figs 3 A, C, 5 A, C). Mesonotum (Figs 3 A, C, 5 A, C): width (0.54) length (0.30). Tegmina (forewings) (Fig. 5 F, G): length (0.57). Hind legs (Figs 3 B, C, 5 D): length (0.91). Male genitalia. Pygofer (Fig. 8 A – D, G, H) bilaterally symmetrical; with caudal margin irregularly convex without lateral processes; in lateral view with irregular caudal margin, without lateral processes; dorsal margin slightly convex / irregular; in ventral view, margins in W-shaped with small triangular ventromedial process; in caudal view rounded, armature of diaphragm weakly sclerotized wider than tall; opening to inner chamber rounded, opening for gonostyli ovoid small approx. 3.0 times smaller than opening to inner chamber. Anal segment (Fig. 8 A – D, I – J) with paired lobes slightly asymmetrical, left lobe slightly larger than right one; in caudal view margin between lobes deeply concave; epiproct and paraproct rather small. Gonostyles (Fig. 1 A – C, E, F) short and slender, in lateral view pointing dorsocaudad with tip slightly surpassing laterocaudal margin of pygofer, widest in basal two thirds, with median margins opposed and close to each other, ventral and dorsal margins slightly sinuate; in distal third narrowing, forming a pair of pincers with keyhole-shaped opening between styles. Aedeagus (Fig. 8 K – N) tubular, asymmetrical; shaft with four spines: a rigid spine arising right laterally near base (long, tubular, compressed ventrally, curved and directed towards apex of shaft), 2 rigid spines at about midlength on left and right side (both compressed, triangular, one short, the other slightly longer), and a movable sinuate spine arising ventrally near apex (in repose directed basally). Flagellum moderately long, slightly surpassing midlength of aedeagal shaft, laterally with serrated margin. Female genitalia. As described for the genus. Immatures. Figs 9 A – F, 10 A – F, 11 A – D, 12 A – D. Nymph 5 th instar. Body length. 1.9 mm (n = 3). Habitus. Slightly dorsoventrally flattened, about twice as long as maximally wide, widest at level of forewing pads and 5 th abdominal segment. Colouration. All parts whitish to very pale yellow. Head. Head without compound eyes, ocelli or traces of them; width at posterior margin of vertex distinctly narrower than pronotum at posterior margin (about 0.4: 1). Vertex short and broad, almost pentagonal, apex anteriorly slightly produced and medially notched, at posterior margin about twice as wide as medially long, lateral and anterior margins ridged, medially a very faint longitudinal carina separating vertex in 2 halves; transition to frons broadly rounded. Frons rather wide, about 1.3 times wider than medially high, maximal width at level of antennae; lateral margins strongly convex, paired intermediate carinae very shallowly convex, well separated from each other, converging towards apex; surface of lateral areas shallowly vaulted, of median area deeply concave; frontoclypeal margin slightly arched and notched. Head with a total of 30 sensory pits, i. e., 12 pits in lateral areas of frons and 3 pits in “ preocular ” area, frontal pits arranged in rows of 6 along intermediate and lateral carinae; variation: in one individual only 5 lateral pits on right side. Sensory pits on head as well as on the other tagmata oval to subcircular, from a cup-like socket at the outer margin arising a seta which is exposed horizontally over the pit without attaining lateral or opposite margins; seta distally knob-like dilated. Post- and anteclypeus shallowly vaulted, smooth, devoid of carinae, post- and anteclypeus separated by a transverse notch. Rostrum distinctly surpassing post-trochanters; tip of rostrum as in adults. Antennae short, scape ring-like; pedicel about 4 times longer than scape, subglobose, distally with about 10 sensory-plaques consisting of a crown of 3 – 4 longer finger-shaped processes arising from a common socle and 4 – 5 short rigid spines at base. 3 rd antennal joint (“ flagellum ”) distinct, knob-like, about 0.25 times shorter than pedicel, surface with 2 – 3 transverse wrinkles, distally continuing in a long arista (“ first projection ” sensu Shi and Yang 1996) and a short peg-like process or arista (“ second projection ” sensu Shi and Yang 1996), both with surface wrinkled; dorsally at distal margin of the 3 rd antennal joint above the bases of the aristae a distinct Bourgoin´s organ consisting of about 20 slender finger-shaped processes forming a distally converging crown. Genae devoid of oblique carina, antennal bases large, subcircular, lamina mandibularis not defined by sutures, distally filiformous. Thorax. Pronotum broad and short, about 5.4 times wider than long in middle line, medially slightly shorter than vertex (0.8 – 0.9: 1); posterior margin straight transverse; medially a membraneous cleft, no ridged carina; dorsolateral carinae oblique, ridged, reaching from anteromedian corner of disc towards lateral posterior margin, dorsal area of disc along carina with 4 sensory pits; lateral lobes at posterior and ventral margin with a total of 4 sensory pits; surface of disc very shallowly concave, nearly plane, surface of lateral lobes slightly convex. Meso- and metanotum medially divided by a longitudinal membraneous furrow, no median carina. Mesonotum approximately 3.8 – 4 times wider than medially long, medially about twice as long as pronotum; posterior margin shallowly concave, at lateral discal carina almost rectangularly bent inwards; disc narrow, lateral discal carina diverging laterocaudad, attaining posterior margin; mesonotum with a total of 16 sensory pits; mediad along discal carina 2 sensory pits, laterad of carina close to posterior margin another sensory pit; forewing pads large, slightly surpassing midlength of hindwing pads; pads sublaterally at posterior margin with a short ridge and a long subcostal carina; forewing pads with a single pit laterad of sublateral ridge, 2 pits at distal margin and 2 pits between subcostal and costal margin. Metanotum about 3.5 times wider than medially long, medially about as long as mesonotum, posterior margin oblique with pads rounded, attaining anterior margin of first abdominal tergite; disc plane, narrow, lateral discal carina straight, slightly diverging caudad, laterad along carina 2 sensory pits. Legs moderately long, slender, normally shaped, without significant dilation or flattening of particular limbs. Pro- and meso-femora at outer distal margin with a short tooth-like protuberance; pro- and mesotarsomeres 2 - jointed with 1 st tarsomere rather short, scale-like; post-tarsomeres 3 - jointed. Post-trochanters medially with a narrow row of 10 parallel ribs forming a cog-wheel structure. Post-femora anteriorly at outer side with a small knob-like protuberance. Post-tibiae about twice as long as post-tarsomeres combined, laterally with 2 small spines, one near base, the other distinctly basad of midlength; distally with 5 coarse spines and a short post-tibial spur. Postbasitarsi only little longer than 2 nd and 3 rd post-tarsomeres together, about 3 times longer than post-tibial spur; postbasitarsi distally with 4, 2 nd post-tarsomeres with 3 stout spines in a transverse row, 2 nd post-tarsomeres relatively short. Pretarsi with aroliae very large, pillow-like with several longitudinal ribs, lateral claws short. Post-tibial spur subconical, terete, with a few irregularly ordered bristles. Abdomen. Tergites mediodorsally incised, distal tergites with sharply ridged crest, no wax-plates on tergites; tergites I – III devoid of sensory pits; tergite IV with a single sublateral pit, extralateral area devoid of pit; tergite V with a sublateral and a lateral pit, tergites VI – VIII with a subdorsal, a sublateral and a lateral pit, extralateral areas V – VIII anteriorly with a single pit; an intermediate or sublateral carina of tergites missing, only on tergite VIII a short sublateral ridge; 9 th abdominal segment with a single dorsal and 2 ventrocaudal sensory pits; in the late 5 th instar males the outline of the pygofer and lateral anal combs, and in the females the developing of paired gonapophyses of the ovipositor are visible. In the individuals examined the dicision for a particular sex appears difficult as there are on one hand clearly recognizable paired structures which may be interpreted as gonapophyses, on the other hand mediolateral protuberances which may be interpreted as anal combs. Stigmata are present in the intersegmental fold between extralateral parts of tergites and sternites, mostly located anteriorly. Nymph 4 th instar. Body length. 1.25 mm (n = 2). Habitus, coloration and equipment with sensory pits as in 5 th instar nymphs; rostrum relatively longer, attaining anterior margin of 5 th sternite; pedicel with surface smooth, distally as in 5 th instar with about 10 sensory-plaques; 3 rd antennal joints relatively larger, with 2 distinct transverse wrinkles with serrate margin; Bourgoin´s organ large, consisting of about 20 long finger-shaped processes arising from a common plate. In mesonotum forewing pads shorter, attaining anterior margin of metanotal hindwing pads, posterior margin shallowly concave. Legs in shape, proportions and spine-configuration of hindlegs as in 5 th instar, cog-wheel structure of post-trochanters with about 10 parallel ribs. Anlagen of male and female genitalia as in 5 th instar. Nymph 3 rd instar. Body length. 0.98 mm (n = 1). Habitus, coloration, proportions, number and arrangement of sensory pits as in 5 th and 4 th instar nymphs; rostrum long as in 4 th instar; pedicel 3.4 times longer than scape (in 4 th and 5 th instar about 4 times longer), distally with about 6 sensory-plaques (10 in 4 th and 5 th instar); 3 rd antennal joint relatively large with surface finely sculptured, Bourgoin´s organ distinct, shaped as in 4 th and 5 th instar. Legs with cog-wheel structure of post-trochanters as in 5 th and 4 th instar with 10 parallel ribs; all tarsi 2 segmented, in second post-tarsomeres limitation of later separation marked by 2 small spines on ventral side; lateral margins of post-tibiae without spines, or spines vestigial; post-tibial spur rather small, developed as rigid cone. Anlage for genital structures faint.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
06AA0E4B88D25FF6968F05131E48747E.taxon	etymology	Etymology. The species epithet is a noun in apposition treated as indeclinable. It is derived from the 1994 movie « Star Trek: Generations », in which Nexus refers to a place of bliss that connects different points in space-time, acting as a link between distinct realities. The gender is masculine.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
06AA0E4B88D25FF6968F05131E48747E.taxon	distribution	Distribution. The species newly described herein was collected in a cave located in the Serra do Ramalho, a vast karstic province situated in southwestern Bahia, Brazil. This region lies within the São Francisco Carbonate Supergroup, particularly the Bambuí Group, and is renowned for its thick limestone deposits and complex subterranean networks (Auler et al. 2001). The karst extends over roughly 12,000 km ², framed by the Carinhanha and Corrente rivers, which are important tributaries of the São Francisco River, forming one of the largest continuous carbonate outcrops in eastern South America. The lithostratigraphy of Serra do Ramalho is dominated by the Sete Lagoas and Lagoa do Jacaré formations, which are composed mainly of calcilutites, calcarenites, calcirudites, and dolomitic facies (Ferreira et al. 2023). These rock assemblages have given rise to an impressive array of geomorphological features, including extensive cave systems, dolines, sinkholes, and subterranean drainage networks. Further details on the geological framework and hydrogeological dynamics of the region are available in Ferreira et al. (2023). Climatically, the area is classified as tropical savanna (Aw, Köppen system). The local regime is characterized by a pronounced dry season extending from March to October, followed by a shorter wet season. Annual precipitation averages approximately 640 mm, while mean temperatures hover near 26 ° C (Alvares et al. 2013; INMET 2018). This seasonal variability strongly influences the ecological processes of the karst, shaping both the surface vegetation and the subterranean communities. Ecological associations, habitat characteristics, and potential threats concerning the new species are discussed in the section “ Ecological Remarks ” below.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
9B87E015D41254A0A82B5D3B7ECC1691.taxon	type_taxon	Type species. Spelaeodelphax nexus Santos & Asche, sp. nov.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
9B87E015D41254A0A82B5D3B7ECC1691.taxon	diagnosis	Diagnosis. Small delphacid, habitus strongly troglomorphic: compound eyes and ocelli totally absent, tegmina strongly reduced, wings vestigial, without sexual dimorphism in the degree of troglomorphy. Females slightly larger in body size than males. Spelaeodelphax gen. nov. differs from all other delphacid genera by the unique combination of the following characters: vertex wide and short, median carina present; frons wider than long, median carina presents in Y-shape apically, lateral carinae not elevated. Hind tibia with lateral spines, five apical teeth, calcar small, not serrated, not reaching the midlength of basitarsus. Male terminalia, pygofer rounded in caudal view; anal segment short, semicylindrical, without process; gonostyli very short, in ventral view pincer-shaped on last third distal, lateral process sub-triangular basally; aedeagus tubular asymmetrical, shaft of the aedeagus elongated, with flattened and tubular spines, flagellum tubular developed, without spine processes.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
9B87E015D41254A0A82B5D3B7ECC1691.taxon	description	Description. Structure. Body length 1.6 – 2.0 mm (male and females). Head in dorsal view distinctly narrower than pronotum at posterior margin (0.6 – 0.7: 1); compound eyes and ocelli totally absent. Vertex short, at posterior margin about 4 times wider than medially long, in dorsal view, apical transverse carina evanescent, slightly concave, incised medially; caudal border moderately concave, median carina evanescent; transition to frons smoothly rounded. Frons about 1.3 times wider than high, widest at level of antennal bases; no median carinae recognizable, surface smooth, shallowly convex, lateral carinae not elevated. Frontoclypeal suture notched, shallowly arched and slightly convex towards postclypeus. Post- and anteclypeus with median carina absent, postclypeus swollen medially. Rostrum distinctly surpassing post-trochanters; distal labial segment furnished with several long bristles and a crown of numerous (14 + 2) short conical or peg-like, on ventral side one sensilla on each side located slightly basad; stylets elongate, filiformous, mandibular stylets distally split, maxillary stylets distally with a row of 4 well-spaced teeth. Antennae with scape short, ring-like; pedicel subglobose, about 3.2 times longer than scape, 1.3 – 1.4 times longer than wide, distally with about 10 sensory-plaques and several bristles; 3 rd antennal joint relatively short, Bourgoin´s organ small but recognizable, long arista filiformous, second arista short arising laterally at base. Genae devoid of oblique carina; lamina mandibularis defined by sutures. Thorax bearing pronotum tricarinate with 2 distinct depressions laterally, at posterior margin about 5 times wider than medially long, posterior margin shallowly concave; lateral carinae strongly divergent, sharply ridged, median carina weakly developed, evanescent distally. Mesonotum about 1.7 – 1.8 times wider than medially long, surface smoothly rounded, seemingly tricarinate, carinae rather weak, almost obsolete, faintly visible with particular light exposure only, lateral carinae diverging caudad, median carina only in anterior disc only. Tegmina coriaceous, surface smooth devoid of bristles, strongly reduced, attaining anterior margin of 2 nd abdominal tergites; about 1.4 – 1.5 times longer than wide, widest at midlength, distal margin broadly rounded, almost subtruncate; venation patterns evanescent, almost absent, ScP + R anteriorly recognizable. Hindwings reduced to small and narrow elongate pads. Legs moderately long, slender, normally shaped, devoid of obvious flattening or dilation of particular limbs; between post-trochanters and - femora on ventral side a short, densely scale-sculptured, nail-like structure (see corresponding scanning photography); such a structure, reminding of the skin of a pangolin, is present in adult Delphacidae and most other Fulgoromorpha, but is rarely referred to. Post-tibia about twice as long as post-tarsomeres combined (2.1: 1), about 6 times longer than post-tibial spur; laterally with 2 spines, one near base, the other shortly basad of midlength; distally with 5 spines, increasingly longer towards outside. Postbasitarsus about 1.6 – 1.7 times longer than 2 nd and 3 rd post-tarsomers combined, about 2.5 times longer than post-tibial spur; distally with 4 equally sized stout spines in a transverse row; 2 nd post-tarsomeres distally with equally small spines. Post-tibial spur (calcar) subulate, terete, distally acuminate, furnished with a few irregular bristles. Pretarsi with relatively small claws, arolium pillow-like with numerous horizontal ridges. Abdomen bearing abdominal tergites 3 – 6 chevron-shaped. Male genitalia. Pygofer moderately long, in lateral view trapezoidal, in caudal view subcircular, dorsally rather narrow, laterocaudal margins convex, ventrocaudal margin concave, medially with a short triangular process; diaphragm weakly sclerotized without a prominent armature or process; foramen for gonotyli small, flat ovate. Anal segment relatively large, hood-like, in ventrocaudal view bilobate; epiproct and paraproct rather small. Aedeagus with shaft tubular, straight, from right basal part arising a long rigid spinose process which is curved toward shaft and left side; shortly distad of midlength of shaft on left and right side a rigid spinose process pointing almost rectangularly laterad; from apex of shaft on ventral side arising a slender, slightly sinuate and apparently mobile process which is curved ventrobasad with tip pointing right laterad; flagellum tubular, devoid of spinose processes, quartercircularly curved over dorsal side, reaching basad to nearly midlength of shaft, phallotreme apically exposed to dorsal side. Gonostyli short, slightly surpassing dorsal margin of diaphragm, in basal 2 thirds medially close to each other in a nearly straight line, inner margin subapically quarter circularly excavate, distally forming a pair of pincers with distal margin truncate. Female genitalia. Ovipositor short and robust, ensiform, slightly surpassing tip of tergite IX and anal segment, basal part nearly at midlength of abdomen, gently curved dorsad; tergite IX ventrally broad, convex, ventromedially with a deep furrow to receive ovipositor, attaining or slightly surpassing midlength of anal segment; anal segment short, dorsocaudally semicircularly excavate, ventrocaudal margin nearly straight transverse with laterocaudally corners rounded; paraproct very short.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
9B87E015D41254A0A82B5D3B7ECC1691.taxon	etymology	Etymology. The genus name is derived from the Greek word for cave (σπήλαιο). The gender is masculine.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
9B87E015D41254A0A82B5D3B7ECC1691.taxon	distribution	Distribution. Brazil, Bahia state, Carinhanha municipality.	en	Santos, Júlio C. C. V., Asche, Manfred, Hoch, Hannelore, Ferreira, Rodrigo L. (2026): First record of a troglobitic Delphacidae in the Neotropics: Spelaeodelphax nexus Santos & Asche gen. nov. et sp. nov. from Brazil (Hemiptera, Fulgoromorpha). Subterranean Biology 55: 1-25, DOI: 10.3897/subtbiol.55.175661
