Speleoberotha palomae, Machado & Martins & Aspöck & Tavares & Aspöck, 2022

Machado, Renato Jose Pires, Martins, Caleb Califre, Aspöck, Horst, Tavares, Leon Gustavo De Miranda & Aspöck, Ulrike, 2022, The first cave associated genus of Berothidae (Insecta: Neuroptera), and a new interpretation of the subfamily Cyrenoberothinae, Zoological Journal of the Linnean Society 195, pp. 1422-1444 : 1426-1433

SPELEOBEROTHA PALOMAE SP. NOV.

FIGS 4–7

Zoobank registration: urn:lsid:zoobank.org:act:85F50233-3FC4-4C1D-AB75-42C460A1221C .

Etymology: Named after friend and researcher Dr Paloma H. F. Shimabukuro, who collected most of the type series.

Diagnosis

Slightly smaller than Speleoberotha mineira . Male: sternite 9 simple, without lobes; gonapophyses 10 as two small, hooked structures; complex gonocoxites + gonostyli 10 shorter or about the same length as gonocoxites 11 in lateral view.

Description

Measurements (N = 10): Body length average ( Fig. 4A ), 1.98 mm (variation, 1.7–2.5 mm); forewing length average ( Fig. 4B ), 3.96 mm (variation, 3.7–4.2 mm); hindwing length average ( Fig. 4B ), 3.42 mm (variation, 3.2–3.7 mm).

Head ( Fig. 4C, D ): Pale yellow, with amber marks. Vertex elevated above compound eyes, pale yellow without tubercles, with thin, medium-sized brown setae. Antennae moniliform, scape subrectangular, ~2.5 times as long as wide, pale yellow, bearing long and pale setae; pedicel subrectangular, approximately twice as long as wide, with medium-sized pale setae; flagellum pale brown with 45–49 articles; flagellomeres subquadrangular, bearing two rings of brown, long setae interspersed with small setae; apical flagellomere triangular. Compound eyes subspherical, black. Frons elongated, pale yellow, bearing small, pale setae. Clypeus pale yellow on medial region, amber on lateral margin, entire surface with scattered, fine, long, brown setae. Labrum narrow, amber, trapezoidal, with anterior margin concave; a group of preapical, tapered, amber setae is present, two of them longer and located on the lateral edge. Gena and postgena amber. Mandible triangular, with tapered apical tooth and one preapical and triangular tooth. Maxilla with cardo quadrangular and yellow, stipes rectangular and yellow; galea narrow, longer than stipes, pale amber, tapering at apex, bearing medium-sized amber setae, especially at interior margin; lacinia dark amber base and pale amber apex, apical part narrow, bearing medium-sized amber setae; maxillary palpus five-articulated; articles elongated and amber, with apex pale yellow, bearing dark amber setae; distal palpomere tapered apically. Labium with amber mentum, bearing small amber setae; ligula amber, with tapered triangular yellow apex, bearing long and tapered pale setae; labial palpus three-articulated; articles elongated and amber with apex pale yellow, bearing amber setae; distal palpomere tapered at apex. Thorax ( Fig. 4A, D ): Pronotum subquadrangular, wider than long, with one transverse furrow; median region pale yellow, bearing small amber setae; lateral region dark amber, covered with abundant long and thin setae, with projected bases. Pleural region pale yellow interspersed with blackish marks, especially at anterior and posterior regions. Ventral region of prothorax pale yellow, with long and pale setae. Mesonotum slightly wider than long, bearing long amber setae; almost all surface is pale yellow except for the median line and anterolateral margins, which are blackish. Metanotum slightly smaller than mesonotum, similar to mesonotum in colour and shape, bearing amber setae. Pteropleurae mostly pale yellow, with blackish marks below wing bases; entire surface with long, amber setae.

Legs ( Fig. 4A ): All the legs pale yellow. Foreleg: coxa elongated, subcylindrical; entire surface with long and thin setae; trochanter and femur with long, pale setae; tibia narrow, with abundant long, fine and pale setae; tibial spurs absent; first tarsomere as long as the following three together; last tarsomere slightly shorter and darker than other tarsomeres; the whole surface with thick, long, yellow setae; tarsal claws amber. Midleg with coxa bearing some long setae; trochanter and femur covered with long, pale setae; tibia narrow, with abundant long and fine setae, tibial spurs absent; tarsi similar to foreleg tarsi. Hindleg similar to midleg in colour and shape.

Wings ( Fig. 4A, B ); forewing: Broadened, with posterodistal margin convex.Wing venation alternating pale and dark brown, with abundant long and fine setae of the same colour as cuticle, margins beaded; trichosors present along wing margin. Membrane mostly hyaline but with dark markings surrounding the major forks and crossveins. Costal area narrow, with humeral recurrent vein; subcostal veinlets forked. Pterostigma suffused, weakly marked, with about six crossveins. Sc ending after the pterostigma and before the wing apex, not fused with radius anterior (RA). Subcostal area with two single crossveins, one of which is brown and located near the wing base, the other one is located at two-thirds of the wing length, brown and surrounded by a suffused dark mark. RA running parallel to subcostal (Sc). Radial area with one brown crossvein, surrounded by a suffused dark mark. Radius posterior (RP) with three branches forking at wing margin (two branches in a few smaller specimens); all radial forks with suffused dark marks. Gradate series inconspicuous. Basal branch of RP forked apically. Presence of two brown RP–media anterior (MA) crossveins: the basal one is located right after the origin of RP basal branch, and the distal one is located after the fork of the RP basal branch. M forking basally to R forks; each branch of M with a secondary fork. MA and media posterior (MP) ending in four main branches with some ramifications at wing margin; only one medial crossvein is present between MA and MP. One M–CuA and one MP–CuA crossvein are present near wing base. Cu forking near wing base, before M and R forks; CuA simple, ending in two main branches with some ramifications at wing margin, before the mid-length of the wing; CuP simple, ending in some small ramifications at wing margin. CuA– CuP crossveins absent. One basal CuP–A1 crossvein is present near wing base. Anal veins A1, A2 and A3 ending in some small ramifications at wing margin. A1–A2 and A2–A3 crossveins present near wing base.

Hindwing: Broadened, shorter and narrower than forewing, with posterodistal margin convex. Wing venation alternating pale and dark brown, with abundant long and fine setae of the same colour as cuticle; margins beaded; trichosors present along wing margin. Membrane mostly hyaline but with dark markings surrounding some forks and crossveins. Costal area narrow, with ~25 unforked subcostal veinlets. Pterostigma suffused, weakly marked, with approximately six crossveins. Sc ending after the pterostigma and before the wing apex, not fused with RA. Two Sc–RA are present near two-thirds of wing length. RA running parallel to SC. Radial area with one brown crossvein surrounded by a suffused dark mark. RP with three branches ending forked at wing margin; RP forks with small, suffused dark mark. Gradate series inconspicuous. Basal branch of RP straight and forking apically. Presence of one brown RP–MA crossvein, located near MA bifurcation. M forking origin of the basal branch of RP, near one-third of wing length; MA ending in four main branches with some ramifications at wing margin; MP ending in two main branches with small ramifications; only one medial crossvein is present, located before the forks of MA and MP. One MP–Cu crossvein is located near Cu apex. Cu forks inconspicuous, making the difference between CuA and CuP difficult to see. One long and sinuous Cu–A1 crossvein is located near the wing base. A1 bifurcated; A2 and A3 simple.

Abdomen ( Fig. 4A ): Tergites and pleural membrane yellow, with spotted small dark brown marks. Sternites yellow. The entire abdominal surface covered with abundant long, fine, yellowish setae.

Male genitalia ( Figs 5 , 6 ): Tergite 9 wider ventrally and not fused with ectoproct. Ectoproct without callus cerci, distally extending in lateral view; in dorsal view, the distal extensions curving outward. Sternite 9 subtriangular in ventral view and not bearing any lobes, but bearing four large, modified and clavate setae. Gonocoxites 11 and 9 as two simple, unpaired bows, connected basally. Complex gonocoxites + gonostyli 10 long and acute apically, but shorter or of the same length as gonocoxites 11 in lateral view. Gonapophyses 10 as two small ventral hooked structures.

Female genitalia ( Fig. 7 ): Gonocoxites and g o n a p o p h y s e s o f s e g m e n t 8 a b s e n t; t e r g i t e 9 n o t f u s e d w i t h e c t o p r o c t; t e r g i t e 9 v e n t r a l l y elongated; gonocoxites 9 ovoid, hypocaudae lacking; gonapophyses 9 as tiny curved sclerites; spermatheca elongated and coiled, becoming narrower and sclerotized in the medial and final portions; proximal portion opening in a large and membranous bursa copulatrix.

Holotype: BRAZIL: Ceará: Ubajara: Parque Nacional de Ubajara : 03°50′04″S, 40°54′29″W, 24 November–1 December 2017, CDC trap, Shimabukuro P.H.F. & Lopes T.A., DZUP 381919 (male; DZUP).

Paratypes (23 ♂, 22 ♀): S ame data as holotype, DZUP 381920–381923 (ten ♂, nine ♀; DZUP); (two ♂, two ♀; NHMW); (one ♂, one ♀; RPSP) ; same data as holotype but Serra do Ibiapaba , 03°50′40.8″S, 40°54′35″W, 799 m, 23 October 2011, light trap, Silva-Neto, A., Xavier, M. & Lima, E. (eight ♂, nine ♀; UFBA) ; Pernambuco: Triunfo: Riacho do Pinga, Cachoeira do Pinga , 07°52′03″S, 38°07′13″W, 890 m, light pan trap, Cavalcante, A. (two ♂, one ♀; UFBA) .

Remarks

Speleoberotha palomae and Speleoberotha mineira are almost identical eidonomically; body colour and wing venation are basically the same in both species. However, Speleoberotha mineira is slightly larger than Speleoberotha palomae and their male terminalia are different, particularly in the shape of sternite 9, length of the complex gonocoxites + gonostyli 10 and the presence of the gonapophyses 10 in Speleoberotha palomae . Female genitalia of both species are similar, but Speleoberotha palomae has larger gonapophyses 9 and bursa copulatrix than Speleoberotha mineira .

The paired hooked structures located apically in the male terminalia of Speleoberotha palomae are interpreted here as the gonapophyses 10. These structures are clearly located internally and are not related to sternite IX. They seem to be unique in Berothidae , because no other species have something similar to them. In Mantispoidea , the only structures that could somehow be related to these paired structures of Speleoberotha palomae are traditionally called the hypomeres in Mantispidae , which were interpreted as gonapophyses 10 by Ardila-Camacho et al. (2021). In Symphrasinae , the gonapophyses 10 are a pair of elongated rods located near the complex gonocoxites + gonostyli 10, whereas in Mantispinae they are reduced and located near the apex of the complex gonocoxites + gonostyli 10. No other paired structures besides the gonapophyses 10 have been described in the male terminalia of Mantispoidea , and for this reason we tentative call these structures in Speleoberotha palomae the gonapophyses 10. We suggest that these paired structures are a plesiomorphic character retained by this new species, which could be reinforced by the position of Cyrenoberothinae in our phylogeny and by the fact the members of the subfamily are known to retain plesiomorphic characters, such as the tergite IX and ectoproct not fused, the presence of the recurrent humeral in the forewing and the lack of bristles in the complex gonocoxites + gonostyli 10.

Most of the type series of Speleoberotha palomae was collected at the Ubajara National Park, a protected area in the wider Caatinga Biome, the most xeric biome of Brazil. However, the park is located on the Ibiapaba ridge, an elevated area (950 m a.s.l. at the highest point) with higher precipitation, supporting some forested areas with many elements shared with the Atlantic rainforest biome, usually classified as ‘brejo de altitude’ ( Queiroz et al., 2017). Ubajara National Park contains a total of 11 caves, and the specimens presented here were collected close to one of these (P. H. F. Shimabukuro, personal communication), suggesting that the species might live inside caves, as its congeneric species does. The specimens from Pernambuco State were collected at Triunfo, a municipality located at the highest point of the state and also surrounded by Atlantic forest of a ‘brejo de altitude’ type ( Fig. 3C ). However, the location of these collected specimens lacks any known nearby caves and/or grottos, but has a 15 m waterfall and rocky formations, which includes many rock overhangs and similar sheltered sites that this species might prefer.

Figures

Figure 3. Distribution of Cyrenoberothinae (Neuroptera: Berothidae). A, world map showing the distribution of all genera included here in Cyrenoberothinae. B, map of South America showing the records of Speleoberotha. C, maps showing in detail the three Brazilian states where Speleoberotha were collected: Ceará, Minas Gerais and Pernambuco.

Figure 4. Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, habitus, lateral. B, wings. C, head, frontal view. D, head and thorax, dorsal view. Abbreviations: A1, A2, anal veins; CuA, cubitus anterior; CuP, cubitus posterior; MA, media anterior; MP, media posterior; RA, radius anterior; RP, radius posterior; Sc, subcostal; 1r-m, first radial–medial crossvein.

Figure 5. Male genitalia of Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, B, lateral views. C, oblique view. D, E, genital complex, oblique view. Abbreviations: ect, ectoproct; gph10, tenth gonapophyses; gst10, tenth gonostyli; gx9–gx11, ninth–eleventh gonocoxites; S8, S9, eighth and ninth sternites; T8, T9, eighth and ninth tergites.

Figure 6. Male genitalia of Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, B, ventral views. C, caudal view. D, genital complex, ventral view. E, genital complex, caudal view. Abbreviations: ect, ectoproct; gph10, tenth gonapophyses; gst10, tenth gonostyli; gx9–gx11, ninth–eleventh gonocoxites; S9, ninth sternite; T9, ninth tergite.

Figure 7. Female genitalia of Speleoberotha palomae sp. nov. (Berothidae: Cyrenoberothinae). A, B, lateral views. C, D, ventral view. E, spermatheca, lateral view. F, spermatheca, ventral view. Abbreviations: bc, bursa copulatrix; d.s., distal portion of spermatheca; ect, ectoproct; gph9, gonapophyses 9; m.s., median portion of spermatheca; p.s., proximal portion of spermatheca; S7, seventh sternite; sp, spermatheca; T7–T9, seventh–ninth tergites.