Adaptive shifts in Neotropical planthoppers: new troglobitic species and the first surface species of Ferricixius Hoch & Ferreira, 2012 (Hemiptera: Fulgoromorpha Cixiidae) from Brazilian caves Author Santos, Júlio César Do Carmo Vaz Centro de Estudos em Biologia Subterrânea, Departamento de Ecologia e Conservação, Instituto de Ciências Naturais, Universidade Federal de Lavras, MG. CEP 37200 - 900, Brazil. & Programa de Pós-graduação em Ecologia Aplicada (PPGECO), Universidade Federal de Lavras. Author Hoch, Hannelore Museum f ¸ r Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Humboldt-University, Invalidenstr. 43, D- 10115 Berlin, Germany. Author Zampaulo, Robson De Almeida 0000-0002-0955-9930 Gerência de Espeleologia e Licenciamento Ambiental - Vale S. A, Nova Lima, Minas Gerais, CEP: 34.006 - 200, Brazil. robsonzampaulo @ gmail. com; https: // orcid. org / 0000 - 0002 - 0955 - 9930 robsonzampaulo@gmail.com Author Simões, Matheus Henrique 0000-0002-0955-9930 Gerência de Espeleologia e Licenciamento Ambiental - Vale S. A, Nova Lima, Minas Gerais, CEP: 34.006 - 200, Brazil. robsonzampaulo @ gmail. com; https: // orcid. org / 0000 - 0002 - 0955 - 9930 robsonzampaulo@gmail.com Author Ferreira, Rodrigo Lopes Centro de Estudos em Biologia Subterrânea, Departamento de Ecologia e Conservação, Instituto de Ciências Naturais, Universidade Federal de Lavras, MG. CEP 37200 - 900, Brazil. & Programa de Pós-graduação em Ecologia Aplicada (PPGECO), Universidade Federal de Lavras. text Zootaxa 2023 2023-08-16 5330 3 375 397 http://dx.doi.org/10.11646/zootaxa.5330.3.3 journal article 10.11646/zootaxa.5330.3.3 1175-5326 8254746 F958F4AD-89F0-4804-96E4-E4E1FB181931 Ferricixius urieli sp. nov. Santos, Hoch & Ferreira ( Figs 1A ; 2A ; 3A–F ; 4A–G ; 5D ; 12A ; 17A, E ) https://zoobank.org/ urn:lsid:zoobank.org:act: EEDFB4AF-109F-48C3-9814-53A6F3816E29 Type material. Holotype : Male , Brazil , Minas Gerais ; Catas Altas municipality, CPC-13 Cave , (UTM 670405W, 7772216S, 23K), 24.viii.2016 , ( Alves, J.P ), ( ISLA 41337 ) . Holotype condition: Dissected, stored in individual vials in 70% ethanol. Additional material. Brazil , Minas Gerais : 1♁ , 2♀♀ , Ouro Preto municipality, Porcos cave ( ISLA 100987 ); 1♁ , Lima Duarte municipality, Casas Cave , (UTM 615466W, 7599900S, 23K), 27.ii.2022 , ( Ferreira RL ) ( ISLA 100988 ) and 1♀♀ , ( ISLA 100989 ); 1♀♀ , Nova Lima municipality, RM-03 cave , (UTM 603947W, 7783375S, 23K), 17.vi.2021 , ( Ferreira R.L. ), ( ISLA 100990 ) . Brazil , São Paulo : 1♀♀ , Iporanga municipality, Bomba Cave (UTM 736984W, 7277175S, 23K), 23.vii.2013 , ( Ferreira RL ), ( ISLA 100991 ) . Description. Coloration (specimen preserved in 70% ethanol): As in figures 1A, 3A–C, generally brilliant orange yellow (67) contrasting with some darker areas on head, mesonotum and abdomen which are Brownish black (65), Deep yellowish brown (75), and Brownish black (65). Tegmina hyaline with Dark yellowish brown (78) veins as in figures 1A and 3D. FIGURE 1. Habitus in dorsal view: Ferricixius urieli sp. nov. (A); Ferricixius michaeli sp. nov. (B); Ferricixius goliathi sp. nov. (C); F erricixius davidi Hoch & Ferreira, 2012 (D). FIGURE 2. Tegmina: Ferricixius urieli sp. nov. (A); Ferricixius michaeli sp. nov. (B); Ferricixius goliathi sp. nov. (C). FIGURE 3. Ferricixius urieli sp. nov. : head (A–C), pedicel with flagellum (C1); tegmen (forewing) (D); hind tibia and tarsomeres (E); hind tarsomeres (F). Body length . Male. 4.6–5.8 mm (n = 3) Female. 5.5–5.7 mm (n=2). Head . Vertex ( Figs 3A, C ): Approx. 1.4 times longer (0.4) than wide (0.2); apical compartment narrow and elongated medially, approx. 1.6 times wider (0.2) than medially long (0.1); apical transverse carina and subapical carina moderately elevated and elongated medially; Irregular caudal border, laterally curved, with a small portion at right angles medially. Frons ( Fig. 3B ): Approx. 1.2 times longer (0.7) than wide (0.6), approx. 1.9 times wider at the height of the antennae than level of the apical carina (0.3). Compound eyes and ocelli ( Figs 3A–C ; 12A ): Well developed; frontoclypeal suture ( Fig. 3B ). Thorax . Pronotum ( Figs 3A, C ): Submedian carinae well developed, in dorsal view slightly concave laterally, evanescent distally; hind margin obtusely angled, slightly concave laterally. Mesonotum ( Fig. 3A ): Tricarinate, lateral carinae well developed and almost straight; median carina well developed proximally, evanescent distally. Tegmina (forewings) ( Fig. 2A , 3D ): Length ( 6.1 mm ); Well developed, cell C3a and C2a approximately the same size, sometimes C3a slightly larger, Cell C1 larger, and C1’ touching the distal margin, vein ScP present Hind legs ( Figs 3E–F ): Hind tibia 2.2 mm long, with 4–5 lateral spines and 6 apical teeth, medially separated (3+3); 1 st tarsomere approx. same size, with 8 apical teeth; 2 nd tarsomere with 8 apical teeth, and three thin setae. FIGURE 4 . Ferricixius urieli sp. nov. , male genitalia: Genital capsule right lateral (A); genital capsule left lateral (B); genital capsule dorsal (C); aedeagus right lateral (D); aedeagus in dorsal view (E); aedeagus left lateral (F); genital capsule ventral (G). Male genitalia . Genital segment ( Figs 4A–C, G ): Ventromedian process triangular elongated, tapering in distal third. Anal segment ( Figs 4A–C ): in dorsal view approx. 2.0 times as long (0.8) as wide (0.4). Genital styles ( Figs 4A–B, G ): as detailed in generic description. Aedeagus ( Figs 4D–F ): Shaft laterally flattened with one large process inserting near the base; weakly concave laterally and curved on the proximal margin, almost straight on the distal margin, and serrated/irregularly dentate on the ventral margin (in distal section). Aedeagal shaft with two subapical spinose processes, one of which is bifid; the bifid process originates right laterally, the longer spine of the bifid process reaches the ventral process when at rest and is twice as long as the shorter spine; bifurcation begins moderately distant from the base of this process; aedeagal shaft with a second spinose process dorsally, moderately short and curved; sometimes shaft with a small process dorso-laterally. Flagellum moderately tubular; process moderately short, approx. 1/3 the size of flagellum. Differential diagnosis. F. urieli sp. nov. can be distinguished from other species of the genus Ferricixius mainly by its epigeomorphic morphology, exhibiting well developed eyes, tegmina, hindwings, presence of a median ocellus and moderately large body size, versus tegmina and hind wings moderately reduced in F. michaeli sp. nov. , and very reduced in F. goliathi sp. nov. and F. davidi . Furthermore, median ocellus absent in F. michaeli sp. nov. , F. goliathi sp. nov. and F. davidi and the eyes are very reduced in F. michaeli sp. nov. , F. goliathi sp. nov. and F. davidi (almost absent in the latter—it is possible to see a small reddish spot in the living specimen, Fig. 16C ). Etymology. The epithet urieli refers to the Archangel Uriel, which is considered to be the angel of wisdom, who shines the light of God’s truth to those who seek information, solutions, or even require wisdom. He is commonly portrayed carrying a book or a scroll, which symbolize wisdom. The name “Uriel” is derived from the Hebrew אוּר̹יא‥ל and means “God is my light.” It is also translated as “Fire of God.” This name was given because F. urieli sp. nov. is the only known epigean species of Ferricixius , thus, living in the light. Remarks. The individuals listed in other material examined were collected at different dates and locations and show the following small morphological variations from the holotype . The specimen collected in Ouro Preto-MG ( ISLA 100987) exhibits some variation compared to the holotype : tegmina with C3a cell slightly larger than C2a; anal segment with paraproct slightly smaller than the holotype ; ventral process of the shaft serrated distally, but with an almost straight margin and many teeth; body color Strong orange yellow (68) contrasting with some areas with light orange yellow (70), deep yellowish brown (75) and dark yellowish brown (78), tegmina hyaline with dark yellowish brown veins (78). The specimen collected in Lima Duarte-MG ( ISLA 100988) has the shortest body length of all the individuals collected ( 4.6mm ) and also exhibits some variations in comparison to the holotype : tegmina with C3a cell slightly larger than C2a, as in the specimen from Ouro Preto-MG; anal segment with paraproct slightly smaller than the holotype ; ventral process of the shaft distally serrated, with a moderately concave margin and few teeth, apically with two slightly larger teeth in close proximity to each other; body color as in the collected specimen from Ouro Preto-MG but slightly more Deep yellowish brown (75) and Dark yellowish brown (78), in the darker regions. The female associated with the specimen from Lima Duarte-MG exhibits a distinct color pattern when compared to the females from other locations: it is much darker, mostly Brownish black (65) contrasting in some areas with Strong yellowish brown (74), in addition it exhibits two highly pigmented spots on the medial/distal region between veins A1 and A3 of the hindwings. FIGURE 5. Habitat Ferricixius urieli sp. nov. : Casas cave, Limas Duarte municipality- MG (A–C); habitus Ferricixius urieli sp. nov. (D). Ecology. Most specimens of F. urieli sp. nov. were sampled by consultancy companies, which unfortunately did not provide any specific information on host plants or habitat. One of the authors however, collected the specimens from Casas cave (Lima Duarte municipality-MG). This cave is inserted in quartzite rocks and presents around 600 meters of horizontal projection. Ferricixius urieli presents traits of external morphology like epigean species, thus, would be expected to occur in external habitats or, if living in caves, inhabiting areas close to the entrance ( Fig. 5A ), both the male and female were found in a deep cave chamber, in an aphotic area ( Fig. 5B–C ). This chamber consists of a collapsed chamber, trespassed by a small autogenic stream. The male was found resting on a rock ( Fig. 5D ), while the female was attracted by the headlamps. Some root mats from the epigean vegetation were observed in some areas. However, immatures of F. urieli sp. nov. we’re not observed. It should be noted that although nymphs were not observed in root mats that were accessible to the collectors, there were other roots in areas inaccessible to people on the cave walls and ceiling that could not be examined. This cave is located within the limits of a State protected area (Parque Estadual do Ibitipoca), and entering this cave is only allowed for research. Other caves in the area were sampled, but no specimens were found. It is noteworthy that specimens of F. urieli sp. nov. were also found in caves from other lithologies, e.g. limestone and iron ore, located quite distant from each other (in straight line, the most distant caves where the species occurs are around 700 km away). This species likely also occurs in external habitats, but considering its frequent association to caves, it might constitute a troglophilic species.