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Erebonyx de Mello, 2021 Type-species: E. catacumbae de Mello, 2021, Published as part of Merlo, Rayanne Lays Sant'Ana, Castro-Souza, Rodrigo Antônio, Bento, Diego De Medeiros & Ferreira, Rodrigo Lopes, 2022, A new troglophilic species of Erebonyx (Orthoptera: Grylloidea: Phalangopsidae) from Brazilian caves, pp. 83-93 in Zootaxa 5222 (1) on page 84, DOI: 10.11646/zootaxa.5222.1.7, http://zenodo.org/record/7456493, {"references":["Mello, F. D. A. D. & Ferreira, R. L. (2021) Erebonyx catacumbae, n. gen. et sp.: a blind, troglobitic cricket from Brazil (Orthoptera, Grylloidea, Phalangopsidae). Zootaxa, 4975 (2), 343 - 356. https: // doi. org / 10.11646 / zootaxa. 4975.2.5"]}

Erebonyx potiguar n. sp. (Figures 1, 2–7, 8–17, 18–22, 23–29, Table 1) Material examined. Holotype ♁, code ISLA 97132, Brazil, Rio Grande do Norte state, Caraúbas municipality, Casa de Homens Cave (5º 34′ 34.5792″ S; 37º 34′ 25.7052″ W), 11.vii.2021, Ferreira R. L. leg. Holotype condition: tegmen, legs I, II and III detached, and maintained in the holotype tube. Paratypes, 4♁♁, 11.vii.2021, (ISLA 97133, ISLA 97134 *, ISLA 97135 *, ISLA 97136 *) (* = nymph stage), 3♀♀, 15.v.2022 (ISLA 97129, ISLA 97130, ISLA 97131), Casa de Homens cave (5º 34′ 34.5792″ S; 37º 34′ 25.7052″ W), Caraúbas municipality, Rio Grande do Norte state, Brazil; 1♁♁, 12.vii.2021, (ISLA 97128), Gruta Capoeira de João Carlos Cave (5º 30′ 56.9946″ S; 37º 31′ 41.4336″ W), Governador Dix-sept Rosado municipality, Rio Grande do Norte state, Brazil. Distribution. Known from two caves: Gruta Capoeira de João Carlos (5º 30′ 56.9946″ S; 37º 31′ 41.4336″ W), Governador Dix-sept Rosado municipality; Casa de Homens Cave (5º 34′ 34.5792″ S; 37º 34′ 25.7052″ W), Caraúbas municipality. Immatures were observed in other caves in the area. Etymology. Specific epithet “potiguar” refers to a person who was born in Rio Grande do Norte state, Brazil. The name is to be treated as a noun in apposition. Diagnosis. E. potiguar can be distinguished from E. catacumbae by the combination of the following characteristics: fore wings slightly developed, symmetrical layout (Figs 11 and 12, 21 and 22); tergites III, IV, V, VI and VII with bristle brushes (T.III–VI, Br., Fig. 14); median lophi developed, apex dilated and curved inward, convex shaped at apex (MPLs, Figs 2–6); paramere well developed and sclerotized, rectangular-lamellar shaped (PsP, Figs 2–6); endophallus with a pair of upper outward projections, narrow and filled, lower part elongated and tapered towards the ectophallic arch, surpassing its margin (End.Sc, Fig. 3); female copulatory papilla well sclerotized, flattened, cone shaped, apex membranous and slightly convex (a, b and c, Fig. 7). Description, male holotype. General Coloration. Body and head light brown, fore wings dark brown (Figs 8–17). Head. similar in color to the body and pronotum, slightly pubescent and with two long bristles between scapes (pers. obs. apparently some were lost after fixation), vertex slightly flattened in lateral view, occiput region vestigially darkened behind the eyes (Figs 8 and 9). Eyes. eyes sub-quadrangular rounded, compound with black ommatidia rounded by slightly margin of depigmented ommatidia, and a superior region more depigmented near the scape insertion (Figs 8 and 9); median and lateral vestigial ocellus present (Fig. 8: circle in red on left head face). Mouthparts. clypeus and labrum whitish brown, mandibles dark brown outlined (Figs 8 and 9); maxillary and labial palps lightly darkened between articulations, with distal region outlined in white (Figs 8 and 9); maxillary palp slightly pubescent, elongated, with five articulations; the first and second palpomeres of same size and shorter than the others; the third and fourth of same sized and bigger than the first two; fifth palpomere is longer than the third and fourth, claviform, dilated in distal portion (Figs 8 and 9); labial palps with three articulations of increasing size, third palpomere claviform (Figs 8 and 9). Antennae. scape slightly pubescent, oval and dilated shaped, with long bristles on interior distal portion (pers. obs. apparently some were lost after fixation); pedicel whitish brown, narrow, cylindrical and slightly compressed on middle region; antennomeres whitish at base and light brown at tip; flagellum light brown (Fig. 9). Thorax. pronotum pubescent, light brown and slightly darkened at the extremities, marked with a vertical median white stripe with darkened regions symmetrically distributed around (Figs 10 and 11); dorsal disc wider than long, lateral lobe rounded, with long bristles at the anterior and posterior margin (pers. obs. apparently some were lost after fixation) (Fig. 10); metanotum non-glandular (Fig. 14). Abdomen. light brown at the proximal part becoming darkened distally; tergites pubescent, light brown (Figs 11 and 14); tergites III, IV, V, VI and VII with bristle brushes, tergite III divided into two portions of bristle brush (dorsal view), tergite IV with a higher density of bristles compared than others (pers. obs. apparently some were broken), tergal glands are present among the aforementioned tergites (T.III–VI, Br. and Gl., Fig 13); sternites pubescent, slightly whiter than the tergites (Figs 11 and 13); subgenital plate darkened, pubescent, quadrangular shape, distal and lateral margins with long bristles, distal central region with an indentation rounded (Figs 15 and 16); supra-anal plate darkened, pubescent, trapezoidal shaped, with small lateral projections, distal portion rounded and with long bristles (Fig. 15); cerci whitish, subapical region slightly darkened (Figs 16 and 17). Legs. Leg I: femur whitish with long bristles, tibia with two subequal apical spurs; tympanum absent, internal proximal face of tibia with a slight discoloration; first tarsomere twice bigger than the second and third together, second tarsomere with one quarter of the third tarsomere length (Fig. 18). Leg II: similar to leg I, with long bristles more dispersed than leg I; tibial apical spurs longer than in leg I, tympanum absent (Fig. 19). Leg III: femur developed, proximal region slightly whitish, other regions light brown uniform, slightly pubescent on the upper inner; tibia with four outer and inner subapical spurs (S.S., Fig. 20), two outer and three inner apical spurs (a, b (outer); c, d and e (inner), Fig. 20); first tarsomere developed and serrulated with two apical spurs, the interior is slightly bigger than the external, first tarsomere is larger than the third and second in size, respectively (Fig. 20). Fore wings. slightly developed, symmetrical layout, with the right tegmen overlapping at the level of their first third of the left tegmen, both covering the first three abdominal tergites, each one becoming narrower towards at tip and triangular shaped in dorsal view (Figs 11 and 12, 21 and 22). Right tegmen. mirror undeveloped, oval shaped dilated, with one cross-vein slightly marked, forming two small cells (M) (Fig. 22); harp undeveloped, triangular shaped, with three poorly marked cross-veins, the distal is more developed, cross-veins connecting the stridulatory vein (SV) with the first cubital vein (Cu1) (Fig. 22); median inner marginal region undeveloped with two chordal veins (CVs), diagonal vein (DV) connecting the mirror with the chordal vein more inner, which present a discontinuous branch towards mirror (Fig. 22); stridulatory file vestigial and degenerated, with 51 teeth evident; lateral field with two parallel veins (Fig. 22). Female. Coloration similar to the males, but more darkened in tergites and sternites (Fig. 23); body size bigger than holotype /males (12.957 mm) (Fig. 23); fore wings absent (Fig. 23); supra-anal plate brownish, pubescent and with two proximal and medial lateral regions without bristles, distal margin U-shaped, slightly elongated, with long apical bristles (Fig. 26); subgenital plate brownish, short and U-shaped, distal margin concave at apex (Fig. 24); ovipositor shorter than the cerci, elongated, sword format at apex (5.568 mm) (Figs 27–29). Female genitalia (ISLA 97129). Copulatory papilla well sclerotized, flattened in dorsal and ventral view, elongated and dilated at tip, cone shaped, apex membranous and slightly convex (a, b and c, Fig. 7); with opening sinuous in proximal portion (dorsal view) (a, Fig. 7); slightly inclined ventrally near the tip (lateral view) (b, Fig. 7); proximal portion with a ventral arch-shaped (ventral view) (c, Fig. 7). Observations in Paratypes. Male phallic sclerites (Holotype ISLA 97132, Figs 2–6) Pseudepiphallus: median lophi elongated in dorsal, ventral and lateral view, median invagination extending from its middle portion at to apex, slightly pubescent, apex dilated and curved inward, more developed and with convex apex (not triangular) in lateral view than compared Erebonyx catacumbae (MPLs, Figs 2–6); paramere well developed and sclerotized, rectangular-lamellar shaped than compared Erebonyx catacumbae, proximal region inward facing of sclerite, distal region projecting towards MPLs apex, this regions (distal and proximal) are separated by a constriction in front view (PsP, Figs 2–6); ramus elongated, rounded at apex, slightly projected towards inner the sclerite (R, Figs 2–4 and 6). Ectophallic invagination: apodemes elongated and slightly projecting toward ramus (Ec.Ap, Fig. 3); ectophallic arch developed and horizontal shaped similar to E. catacumbae (Ect.Arc, Fig. 3); ectophallic fold less sclerotized, more elongated than E. catacumbae (Ect.F, Fig. 3). Endophallus: well developed and sclerotized, with a pair of upper outward projections, narrower and more filled than E. catacumbae (End.Sc, Fig. 3); lower part elongated and tapered towards the ectophallic arch, surpassing its margin (Fig. 3).

Cavernas sempre despertaram a curiosidade dos cientistas em função dos troglóbios, organismos exclusivamente subterrâneos que frequentemente apresentam distribuição restrita e elevados níveis de endemismo. Eventos paleoclimáticos têm sido apontados como os principais responsáveis pela origem de espécies troglóbias, e o oeste da formação Jandaíra, nordeste do Brasil, alia a ocorrência de extensas áreas cársticas, enorme concentração de cavernas e aquíferos subterrâneos em uma região atualmente semiárida. Essa região é reconhecida pela riqueza e concentração de espécies troglóbias, algumas das quais (isópodes, anfípodes e hemípteros) possuem distribuição relativamente ampla em áreas geológica e hidrologicamente distintas. Por meio de análises filogenéticas e filogeográficas, objetivamos avaliar os padrões de variação genética desses táxons, investigar possível diversidade críptica e delimitar Unidades Taxonômicas Operacionais (UTO´s), bem como indicar os fatores que possivelmente influenciaram sua diversificação. Além disso, objetivamos avaliar o estado de conservação das linhagens e identificar áreas prioritárias para sua conservação. Com exceção de Cirolanidae sp.1 (Isopoda), os demais grupos apresentaram extensa diversidade de linhagens crípticas e, além da identificação de três UTO´s para Cirolanidae sp.2 e cinco para Potiberaba (Amphipoda), Kinnapotiguara troglobia (Hemiptera) provavelmente é um complexo com sete espécies crípticas, com linhagens restritas às microbacias (táxons aquáticos) e lajedos (Kinnapotiguara). Somente uma pequena parcela das linhagens encontra-se em áreas protegidas, enquanto a maioria ocorre em áreas expostas a ameaças antrópicas e podem ser consideradas ameaçadas de extinção: Cirolanidae sp.1 foi categorizada como Vulnerável, enquanto Cirolanidae sp.2 possui uma linhagem Menos Preocupante (LC) e duas Criticamente Ameaçadas (CR). Para Potiberaba, há duas linhagens LC, uma Em Perigo (EN, P. porakuara) e duas CR, e, para Kinnapotiguara, há quatro linhagens EN e três CR (incluindo K. troglobia). Foram identificadas duas áreas prioritárias para conservação dessas linhagens, com destaque para a região da microbacia do riacho do Abreu e lajedo do Rosário, em Felipe Guerra/RN, que abriga metade (oito, sendo seis endêmicas) das linhagens identificadas. O lajedo do Rosário, que pode ser considerado um sistema de cavernas e abriga 24 espécies troglóbias, é um hotspot de biodiversidade subterrânea. Por fim, apresentamos a cartilha “Vida nas CaveRNas”, elaborada como uma estratégia educacional com o objetivo de despertar o interesse sobre as cavernas, seus habitantes e sua conservação. Caves have historically attracted attention of scientists due to troglobites, exclusively subterranean organisms that often have restricted distribution and high levels of endemism. Paleoclimatic events have been pointed out as the main responsible for the origin of troglobitic species, and the west of the Jandaíra formation, northeast of Brazil, combines the occurrence of extensive karst areas, huge concentration of caves and subterranean aquifers in a region that is currently semi-arid. This region is recognized due to the richness and concentration of troglobitic species, some of which (isopods, amphipods and planthoppers) have a relatively wide distribution in geologically and hydrologically distinct areas. Through phylogenetic and phylogeographic analyzes, we aim to assess the patterns of genetic variation of these taxa, investigate possible cryptic diversity and delimit operational taxonomic units (OTU’s), as well as indicate the factors that possibly influenced their diversification. In addition, we aim to assess the conservation status of the lineages and identify priority areas for their conservation. With the exception of Cirolanidae sp.1 (Isopoda), the other taxa showed extensive diversity of cryptic lineages and, in addition to the identification of three OTU’s for Cirolanidae sp.2 and five for Potiberaba (Amphipoda), Kinnapotiguara troglobia (Hemiptera) probably is a complex with seven cryptic species, with lineages restricted to microbasins (aquatic taxa) and outcrops (Kinnapotiguara). Only a small portion of the lineages are found in protected areas, while the majority occur in areas exposed to anthropic threats and can be considered endangered: Cirolanidae sp.1 was categorized as Vulnerable, while Cirolanidae sp.2 has one Least Concern (LC) and two Critically Endangered (CR) lineages. For Potiberaba, there are two LC, one Endangered (EN, P. porakuara) and two CR lineages, and, for Kinnapotiguara, there are four EN and three CR (including K. troglobia) lineages. Two priority areas were identified for the conservation of these lineages, with emphasis on the region of Abreu stream microbasin and Rosário outcrop, in Felipe Guerra municipality, which harbor half (eight, six of which are endemic) of the identified lineages. The Rosário outcrop, which can be considered a cave system and harbor 24 troglobitic species, is a new hotspot of subterranean biodiversity. Finally, we present the booklet “Vida nas CaveRNas”, designed as an educational strategy with the aim of arousing interest in caves, their inhabitants and their conservation.

It is estimated that the Brazilian karst areas sum about 200.000 km2. The caves, one of the main components of karst, are important windows into the biological studies on hypogean environments. In Rio Grande do Norte are known 563 caves, and 476 of them are in the municipalities of Baraúna, Felipe Guerra, Governador Dix-Sept Rosado, Apodi and Mossoró, the Western region of the State. However, like in the rest of the country, the cave fauna of the State is still poorly understood. This study used data from invertebrates harvested in 47 caves and aimed to analyze the effect of environmental change between the dry and rainy seasons in the communities of cave invertebrates, characterize these communities and evaluate the relationships between morphological and biotic variables of the caves and surroundings, and to define priority areas for conservation of cave environments of the study area from biotic parameters. Strong effects were found in the community structure of cave invertebrates due environmental changes between seasons, with values of total richness, abundance, diversity and ecological complexity significantly higher in the rainy season. It was possible to assess how the morphology of the cave and the external environment variables affect the biotic system, so that the variety of resources, forest cover in the vicinity, the area of the cave and its entrance were the variables that best explained the structure communities of cave invertebrates in the region. High values of total richness of invertebrates (36,62 ± 14,04 spp / cave) and troglomorphic species (61 species, mean 1,77 ± 2,34 spp / cave) were found and, given the biological relevance in the context of the area national and the imminent anthropogenic pressures existing, we defined four priority areas for actions aiming cave biodiversity conservation in the region. Estima-se que as áreas cársticas brasileiras perfaçam cerca de 200 mil km2. As cavernas, um dos principais componentes do relevo cárstico, são uma importante janela para a realização de estudos biológicos sobre os ambientes hipógeos. No Rio Grande do Norte são conhecidas 563 cavidades, sendo que 476 delas encontram-se nos municípios de Baraúna, Felipe Guerra, Governador Dix-Sept Rosado, Apodi e Mossoró, região Oeste do Estado. No entanto, a exemplo do restante do país, a fauna cavernícola do RN ainda é pobremente conhecida. O presente estudo utilizou dados de coletas de invertebrados realizadas em 47 cavernas e teve como objetivos analisar o efeito das alterações ambientais entre as estações seca e chuvosa nas comunidades de invertebrados cavernícolas, caracterizar essas comunidades e avaliar as relações entre variáveis bióticas e morfológicas das cavernas e do entorno, bem como definir áreas prioritárias para a conservação de ambientes cavernícolas da área de estudo a partir de parâmetros bióticos. Foram encontrados fortes efeitos sobre a estrutura das comunidades de invertebrados cavernícolas em função das mudanças ambientais entre as estações, com valores de riqueza total, abundância, diversidade e complexidade ecológica significativamente maiores na estação chuvosa. Foi possível avaliar como a morfologia da caverna e variáveis do ambiente externo influenciam as variáveis bióticas do sistema, de forma que a variedade de recursos, a cobertura florestal no entorno, a área da caverna e de suas entradas foram as variáveis que melhor explicaram a estrutura das comunidades de invertebrados cavernícolas da região. Foram encontrados elevados valores de riqueza total de invertebrados (36,62 ± 14,04 spp/caverna) e de espécies troglóbias (61 espécies, média de 1,77 ± 2,34 spp/caverna) e, diante da relevância biológica da área no contexto nacional e das iminentes pressões antrópicas existentes, foram definidas quatro áreas prioritárias para ações de conservação da biodiversidade cavernícola regional.