New Species of Endecous (Grylloidea, Phalangopsidae, Luzarinae) from the Iguaçu National Park (Brazil), Including Bioacoustics, Cytogenetic and Distribution Data

Zootaxa 4237 (3): 454–470 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4237.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:BC7FE3FF-4B54-41FB-B74D-0AAAB784CD3B

New species of Endecous (Grylloidea, Phalangopsidae, Luzarinae) from the Iguaçu National Park (Brazil), including bioacoustics, cytogenetic and distribution data

PEDRO G. B. SOUZA-DIAS1,5, NEUCIR SZINWELSKI2, MARCOS FIANCO2, ELLIOTT CENTENO DE OLIVEIRA3, FRANCISCO DE A. G. DE MELLO4 & EDISON ZEFA3 1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, rua do Matão, travessa 14, n. 101, 05508-900, Cidade Universitária, São Paulo, SP, Brazil 2Centro de Ciências Biológicas e da Saúde, Universidade Estadual do Oeste do Paraná, rua Universitária, n. 2069, 85819-110, Universitário, Cascavel, PR, Brazil 3Programa de Pós-Graduação em Biologia Animal, Departamento de Ecologia, Zoologia, Universidade Federal de Pelotas, rua Capão do Leão s/n, 96010-900, Pelotas, RS, Brazil 4Departamento de Zoologia, Instituto de Biociências de Botucatu, Universidade Estadual Paulista "Júlio de Mesquita Filho" (UNESP). 18618-000, Botucatu, SP, Brazil 5Corresponding author. E-mail: [email protected]

Abstract

In this study we describe two new species of cavicolous-straminicolous crickets for southern Brazil. Endecous chape n. sp. and E. naipi n. sp. are sympatric crickets that inhabit the Atlantic Semideciduous Forest of the Iguaçu National Park and adjacent areas. The descriptions were based on morphological characters, mainly from male genitalia and tegmina of adult males. Furthermore, we describe the chromosomes and the calling song of one of the new species, E. chape n. sp., presenting a discussion about the morphology of the phallic complex of Endecous and a distribution map for the species of the genus. The type-material is deposited in the Museu de Zoologia da Universidade de São Paulo (MZSP), and in the Coleção de Insetos do Departamento de Zoologia (Zoology Department Insect Collection), Universidade Estadual Paulis- ta—UNESP—Botucatu campus.

Key words: Insect, Orthoptera, calling song, chromosomes, taxonomy, cricket

Resumo

Neste estudo descrevemos duas novas espécies de grilos cavícolas-estraminícolas para o sul do Brasil. Endecous chape n. sp. e E. naipi n. sp. são grilos simpátricos habitantes da Floresta Estacional Semidecidual do Parque Nacional do Iguaçu e áreas próximas. As descrições basearam-se em caracteres morfológicos, sobretudo da genitália e das tégminas dos ma- chos. Além disso, descrevemos os cromossomos e o som de chamado de uma das espécies, E. chape n. sp., apresentamos uma discussão sobre a morfologia do complexo fálico de Endecous, e um mapa de distribuição das espécies do gênero. O material-tipo está depositado no Museu de Zoologia da Universidade de São Paulo (MZSP) e na Coleção de Insetos do Departamento de Zoologia da Universidade Estadual Paulista—UNESP—campus Botucatu.

Palavras-chave: Inseto, Orthoptera, som de chamado, cromossomos, taxonomia, grilo

Introduction

Endecous Saussure (1878) is one of the largest Luzarinae genera in South America, with 14 known valid species grouped in three subgenera, occurring through Brazil, Argentina, Bolivia and Uruguay (Gorochov, 2014; Cigliano

454 Accepted by D. Rentz: 14 Dec. 2016; published: 1 Mar. 2017 et al., 2016). The genus is widely distributed in Brazil, where its 10 recognized species are spread throughout the most parts of the country (Souza-Dias et al., 2014; Zefa et al., 2014; Bolfarini & Bichuette, 2015). Species of Endecous present tegmina with a functional stridulatory apparatus, except one species. E. apterus Bolfarini & Souza-Dias, 2014 has lost its tegmina as a result of troglobiomorphosis (Souza-Dias et al. 2014). Until now, only E. alejomesai Zefa, 2010, E. betariensis de Mello & Pellegatti-Franco, 1998, and E. itatibensis Rehn, 1918 had their calling songs recorded. These songs are composed of phrases, and including species-specific song parameters (de Mello & Pellegatti-Franco, 1998; Zefa, 2006; Zefa et. al. 2010). Zefa (2006) observed that populations of E. itatibensis from different regions present intraspecific variations in the carrier frequency and rhythm of the emitted signals. The Phalangopsidae comprises 975 valid species (Cigliano et al., 2016). Studies on cytogenetics are known for only 17 species. The karyotypes vary from 2n=11 to 2n=21, and with the basic X0 sex system (Zefa et al., 2010). Regarding Endecous, six of the 14 described species were karyotyped, varying in number, chromosome morphology and sex system determination: E. onthophagus (Berg, 1891) and E. itatibensis have 2n=19, X0 (Cardoso et al., 1984; Mesa & Garcia-Novo, 1997), E. alejomesai, E. cavernicolus Costa-Lima, 1940, and E. betariensis have 2n=21, X0 (Piza, 1945; Zefa et al. 2010), and E. ubajarensis Zefa, 2014 2n=14, X1X20 (Zefa et al. 2014). Endecous is considered a cavicolous-troglobitic cricket: its species are active at night on leaf litter, while during daytime remain inside caves, rock gullies, crevices, burrows, and general cavities, even in urban areas. Some species are generalists, with cavicolous and straminicolous populations, as E. itatibensis and E. aguassay Mews, 2008 and three species are troglobitic, as E. apterus, E. cavernicolus, and E. peruassuensis Bolfarini, 2015. For more information on the habitat occupation of Endecous see Souza–Dias et al. (2014), Zefa et al. (2014), and Bolfarini & Bichuette (2015), and references therein. Here we describe two new species of Endecous from the Iguaçu National Park (Brazil), and the calling song and chromosomes of one of the new species, with comments on the morphology of the phallic complex of the genus. Additionally, a distribution map of Endecous species is presented.

Material and methods

Taxonomy. Individuals of E. chape n. sp. and E. naipi n. sp. were collected at night on trails of the Iguaçu National Park, at the municipalities of Foz do Iguaçu (25°37'6.16"S, 54°28'56.34"W) and Céu Azul (25°9'20.96"S, 53°50'34.51"W), Paraná State. Individuals of E. chape n. sp. were also collected in the municipality of Concórdia, west of Santa Catarina State (27°15'38.50"S, 51°58'37.55"W). The national collecting permit was issued by the Instituto Chico Mendes de Conservação da Biodiversidade—ICMBio to PGBSD (SISBio 31497-1), and NS (SISBio 13543). The specimens were examined, described and compared using a Leica MZ-9.5 stereomicroscope. They were photographed immersed in 80% ethanol using a Leica MZ-16 stereomicroscope attached with a DFC-420 video camera. The same procedure was used for the male genitalia photographs. The software Leica Application Suite LAS v4.0 was used to take the images, and the software Helicon Focus 5.3 to stack them. Subsequently, the images were edited in a licensed Adobe Photoshop CS6 and GIMP (Gnu Image Manipulating Program) 2.8. The male phallic complexes were removed and treated with an aqueous solution of 10% KOH for 24h to remove muscular tissues, and stored in a small glass vial with 80% ethanol together with the respective specimen. The female copulatory papilla were removed and stored in a small glass vial with 80% ethanol together with the respective specimen. The genitalia terminology follows Desutter (1987), Desutter-Grandcolas (2003) and Souza- Dias (2015). Distribution map. The distribution map was built in the software QGIS 2.10. The biogeographic provinces followed the proposition of Morrone (2014), and the shapefiles were provided by Lowenberg-Neto (2014). The distribution data were obtained from Endecous’ previous studies, labels of studied specimens, field trips, and additional data provided by Dr. Marcio Bolfarini (Bolfarini & Bichuette, 2015). Bioacoustics. The calling songs was registered only for E. chape n. sp. Six males were recorded in laboratory with RR-US300 Panasonic tape recorder, and microphone placed about 10 to 20cm from the calling male. The room temperature was measured after each recording with an INCOTERM L-007/09 thermometer. The recorded songs were analyzed with Avisoft-SASlab Lite software, digitized at sampling frequency of

NEW ENDECOUS FROM THE IGUAÇU NATIONAL PARK Zootaxa 4237 (3) © 2017 Magnolia Press · 455 22050 Hz, resolution 16 bit, fast Fourier transformation (FFT) – Length 256, frame 12.5%, window hamming and overlap 99.1%. We selected the best recordings of the calling song of each individual to analyze the phrase duration (i.e., the elapsed time from the first to the last pulse of a phrase); number of subphrases per phrase; subphrase period (elapsed time from the subphrase’s first pulse to the beginning of subsequent subphrase); number of pulses per subphrase; dominant frequency (highest intensity spectral component); and phrase amplitude. We consider a pulse as a train of sound cycles produced during inward movement of the forewings, and a phrase as a train of pulses. Cytogenetics. Cytological preparations were obtained only from E. chape n. sp., from the caeca of two males and three females, previously injected with a 0.05% aqueous solution of colchicine for 5h. The tissues were dipped in a hypotonic solution of sodium citrate (0.075 M) for 10 min prior to fixation in Carnoy I (3 parts of ethanol: 1 part of acetic acid). The cells were squashed in a drop of 45% acetic acid and stained with 0.5% lacto-acetic orcein. The chromosomes were photographed under a microscope Zeiss Lab. A1, and Axio Cam ERc5s. The chromosome terminology is according to Levan et al. (1964). Abbreviations. Male genitalia. Arc, ectophallic arc; Arm, pseudepiphallic arm; D.Ec.P., dorsal ectophallic projection; Ect.Ap., ectophallic apodeme; Ect.F., ectophallic fold; End.Ap., endophallic apodeme; End.Sc., endophallic sclerite; M.S., membranous sphere; PsP1, pseudepiphallic paramere 1; PsP2, pseudepiphallic paramere 2; V.E c.P., ventral ectophallic projection; Sc.A, sclerite A of the pseudepiphallic arms. General morphology. I, II, III, anterior, median, posterior (leg, tarsomere); DD, LL, dorsal disc, lateral lobe of pronotum; FW, forewing; F, femur; T, tibia. Repositories. MZSP, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil; Coleção de Insetos do Departamento de Zoologia (Zoology Department Insect Collection), Instituto de Biociências, Universidade Estadual Paulista (São Paulo State University) – UNESP – Botucatu campus. Institution. ZIN, Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia. Measurements (mm). AWP, anterior width of pronotum; HW, head width; IOD, intra-ocular distance; LFIII, length of hind femur; FWL, forewing length; PL, pronotum length; PW, pronotum width (at mid-line); LBt-III, length of basitarsus III; LTIII, length of hind tibia; OL, ovipositor length; PWP, posterior width of pronotum; WFIII, width of hind femur; FWW, forewing width (at mid-line).

Results

Endecous (Endecous) chape Souza-Dias & de Mello n. sp. Figures 1, 2 (A, C), 3.

Type locality. Brazil, Paraná State, Parque Nacional do Iguaçu (Iguaçu National Park). Type material. Holotype male, 7 male paratypes, 10 female paratypes. Holotype: Brasil, Parque Nacional do Iguaçu. Trilha do Poço Preto. 20-30.i.2008. Dias, P.G.B.S. & de Mello, F.A.G. col. (MZSP). Paratypes: 1 male, 5 females, same data as the holotype (MZSP); 1 male, same data as the holotype, with FWs and genitalia removed and kept with the specimen (MZSP); 1 male, same data as the holotype (PSD33) (MZSP); 1 male, same data as the holotype, with right FW removed and fixed in microscope slides (MZSP); 1 male, labeled: Brazil, Parque Nacional do Iguaçu, Poço Preto. 22-28.xii.2010. Dias, P.G.B.S. col. Pitfall (MZSP); 1 male, labeled: “BR [Brazil], SC [Santa Catarina State], Concórdia. 28.xi-01.xii.2011. Dias, P.G.B.S. col.”, (PSD34) (MZSP); 1 female, same data as the holotype (PSD35); 1 female, with the copulatory papilla removed and kept with the specimen, labeled: Brazil, Parque Nacional do Iguaçu, Trilha das Cataratas. 21.xii.2010. Dias, P.G.B.S. (MZSP); 1 female, with the copulatory papilla removed and kept with the specimen, labeled: Brazil, PR [Paraná State], Foz do Iguaçu, Parque Nacional do Iguaçu, Trilha do Poço Preto. P2—adultrap. 02-08.xi.2005. Szinwelski, N. col. (MZSP); 1 female, labeled: Brazil, PR [Paraná State], Foz do Iguaçu, Parque Nacional do Iguaçu, Trilha do Poço Preto. 11.x.2010. Dias, P.G.B.S. col. (MZSP); 1 male, 1 female, labeled: Brazil, Parque Nacional do Iguaçu, Trilha das Cataratas. 21.xii.2010. Dias, P.G.B.S. (Departamento de Zoologia, Instituto de Biociências, UNESP Botucatu). Distribution. Southern Brazil, Atlantic Forest of western Paraná and Santa Catarina States. E. chape n. sp. is reported for the Iguaçu National Park, municipalities of Foz do Iguaçu and Céu Azul (Paraná), and the area of Concórdia municipality (Santa Catarina).

456 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. FIGURE 1. Endecous chape n. sp. General morphology. A, male habitus; B, male head, pronotum and tegmen, dorsal; C, male head, pronotum and tegmen, lateral; D, male head and pronotum, dorsal; E, male subgenital plate; F, male supra anal plate; G, female subgenital plate; H, female supra anal plate; I, male TIII and basitarsus.

458 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. FIGURE 3. Endecous chape n. sp. genitalia. Male genitalia in dorsal (A, B), ventral (C, D), dorsolateral (E), and lateral view (F). Female copulatory papilla in dorsal (G), ventral (H), and lateral view (I). Conventions: Arc—ectophallic arc; Arm— pseudepiphallic arm; D.Ec.P.—dorsal ectophallic projection; Ect.Ap.—ectophallic apodeme; Ect.F.—ectophallic fold; End.Ap.—endophallic apodeme; End.Sc.—endophallic sclerite; M.S.—membranous sphere; PsP1—pseudepiphallic paramere 1; PsP2—pseudepiphallic paramere 2; V.Ec.P.—ventral ectophallic projection; Sc.A—sclerite A of the pseudepiphallic arms. Scale bar: 1mm.

NEW ENDECOUS FROM THE IGUAÇU NATIONAL PARK Zootaxa 4237 (3) © 2017 Magnolia Press · 459 Etymology. Chape is the nickname of Associação Chapecoense de Futebol, a football team from the city of Chapecó, the largest city of the west of Santa Catarina State, which is part of the distribution of this new species. On 29th November 2016, a plane crash killed almost all the entire team, including the players, coaches, staff and journalists. E. chape n. sp. is a tribute to all people that made part of this team’s beautiful history, which was, unfortunately, interrupted by this terrible tragedy. The word “chape” is a noun in apposition. Diagnosis. Within the genus, E. chape n. sp. can be recognized by the following characters: males FWs short, rounded, not trespassing the third abdominal tergite; mirror irregular, with six cells, the distal weak and irregular; harp with three weak veins; lateral field with veins irregular, weak. Median part of the pseudepiphallic sclerite fused; pseudepiphallic arms lateral, hard, straight, apex pointed, curved inwards, with a conspicuous membranous sphere on the apex; sclerite A fused with the pseudepiphallic arm; PsP1 and PsP2 connected, forming a circular structure; PsP2 smaller and less sclerotized than PsP2; dorsal ectophallic projections (D.Ec.P.) long, almost reaching the apex of the pseudepiphallic arms, clearly separated and detached of the phallic complex; endophallic sclerite medial, long, grooved, forming a medio-dorsal crest; endophallic apodeme present, small, paired. Description. General body coloration medium to reddish brown, almost uniform (Figs 1A, B). Head. Dorsum with soft pubescence, medium brown, yellowish in the middle (Fig. 1B). Vertex and fastigium medium brown (Figs 1B, D). Fastigium little wider than long, narrower than scape, slightly narrower toward the apex, with a double row of small bristles, and a spot yellowish brown on apical portion, corresponding to the area of the median ocellus (Figs 1B, D). Ocelli absent. Eyes obovate, prominent (Figs 1B, C). Antennal scape medium brown, outer border yellowish brown. Antenna not annulated; antenomeres medium brown (Figs 1B–D). Frons medium brown speckled with small, light brown spots, and a macula light brown below each antennal fossa. In frontal view, gena medium brown; line diagonal, thin, light brown that goes from gena toward frons. In lateral view, gena medium brown with diagonal band light brown, which surround the external and superior margin of the eye, and weak ascendant anastomosed lines medium brown (Fig. 1C). Clypeus and labrum light brown. Maxillary palpi elongated, thin; joints 1–2 whitish, joints 3–5 elongate, same-sized, medium brown, speckled with very small light brown spots; apical third of joint 5 curved, apex rounded, whitish (Figs 1C, D). Thorax. Pronotum DD wider than long, slightly pubescent, general coloration medium brown; two spots yellowish brown on anterior portion, with reticulated lines medium brown; two blotches large, yellowish brown on median portion, divided by stripe medium brown, which is divided by a line yellowish brown that goes from the anterior to posterior portion of DD; two spots yellowish brown on posterior portion; DD cephalic and caudal margins sub-straight; LL ventro-cephalic angle slightly salient, rounded, ventro-caudal angle almost straight, ventral margin gradually ascendant (Figs. 1C, D). Legs. Legs I and II yellowish to medium brown, not annulated (Fig. 1A). Auditory tympanum present on both sides of TI; internal tympanum oval, small; external rounded. TI and TII with two same-sized ventral apical spurs. FIII light to medium brown, with diagonal medium brown stripes, distal half medium brown. TIII medium brown; subapical spurs 4/4, the first smaller and associated to the apical; serrulation between and above subapical spurs 1 and 2; apical spurs 3/3, more developed on inner face; dorsal and median apical spurs same-sized and the longest on both faces (Fig. 1A, I). Basitarsus I, II and III medium brown; basitarsus III with double row of spines (Fig. 1I). Abdomen. Medium to yellowish brown, pubescent (Fig. 1A). Subgenital plate wider than long; anterior margin sub-straight, posterior margin broad, rounded, with short distal projections (Fig. 1E). Supra anal plate medium brown, with a central macula yellowish brown; anterior margin slightly concave, posterior margin sub-straight, without distal projections (Fig. 1F). Cerci as long as FIII, medium brown. Male. FWs membranous, short, rounded, not trespassing the third abdominal tergite (Figs 1A, B); right FW hard, medium brown (Figs. 1B, 2A, C); left FW membranous, transparent; stridulatory vein present, with ca. 82 - 94 teeth (n=09) (Figs 1B, 2A, C); chord with two elongated cells; mirror irregular, with six cells, the distal weak and irregular (Figs. 1B, 2A, C); harp with three weak veins, one incomplete (Figs. 1B, 2B, D); lateral field with veins irregular, weak (Figs 2A, C). Male genitalia. Pseudepiphallus: median part of the pseudepiphallic sclerite fused (Figs 3A, B); pseudepiphallic arms lateral, hard, straight, apex pointed, curved inwards (Figs 3A–D); presence of a conspicuous membranous sphere on the apex of the pseudepiphallic arms (Figs 3A–E); sclerite A fused with the pseudepiphallic arm, the apical part of the sclerite is visible in ventral view (Figs 3C, D). PsP1 and PsP2 connected, forming a circular structure, visible in superior and lateral views (Figs. 3B, E, F); presence of a membrane between the contact points of PsP1 and PsP2 (Figs. 3B, E); PsP1, hard, visible in ventral and dorsal views (Figs. 3B–F); PsP2 small, less sclerotized than PsP1, curved inwards (Figs 3B, E, F). Ectophallic invagination: ectophallic apodeme short, robust (Figs 3A, B); ectophallic arc below the median part of the pseudepiphallic sclerite (Figs 3A, B); dorsal ectophallic projections long, almost

460 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. reaching the apex of the pseudepiphallic arms (Fig. 3A), clearly separated and detached of the phallic complex (Figs 3A, F), apex curved inwards; apex of the ventral ectophallic projections pointed, curved inwards (Figs 3C, D); ectophallic fold membranous, with a weak sclerotization visible in dorsal view (Figs 3A, B). Endophallus: endophallic sclerite medial, long, grooved, forming a medio-dorsal crest (Figs 3C, D); endophallic apodeme present, small, paired (Figs 3C, D). Female. Larger than male, general coloration similar. Apterous. Supra anal plate medium brown, similar to male. Subgenital plate medium brown, short, wider than long, anterior margin slightly convex, posterior margin rounded, with a central concavity. Female genitalia: copulatory papilla laterally flattened, apex rounded, basis with a dorsal projection, as in Figs 3G–I. Remarks on paratype. One male paratype had the FWs different from the holotype showing chord with two cells, one elongated and other reduced (Figs 2A, C). Measurements (mm). Males (n=8): HW—3.53 (3.2–3.9); IOD—1.88 (1.5–2.3); PL—2.96 (2.5–3.9); AWP—3.31 (2.6–3.9); PWP—4.57 (4.2–5.1); PW—5.22 (4.5–5.6); FWL—5.66 (5.1–6.2); FWW—4.9 (4.5–5.2); LFIII—16.25 (14–17.5); WFIII—3.9 (3.5–4.1); LTIII—17.2 (15.2–19.2); LBt–III—5.35 (4.1–6). Females (n=11): HW—3.42 (3.2–3.6); IOD—1.8 (1.5–1.9); PL—2.84 (2.2–3.5); AWP—3.16 (2.7–3.4); PWP— 4.57 (4.2–5.1); PW—5.02 (4.9–5.2); LFIII—15.6 (12.2–16.9); WFIII—3.86 (3.5–4.1); LTIII—17 (14.9–18.9); LBt-III—5.25 (4.5–5.9); OL—13.6 (10.1–15.1).

Endecous (Endecous) naipi Souza-Dias n. sp. Figures 2 (B, D), 4, 5.

Type locality. Brazil, Paraná State, Parque Nacional do Iguaçu (Iguaçu National Park). Type material. Holotype, male 1 male paratype. Holotype: BR [Brazil], Paraná, Foz do Iguaçu. Parque das Aves. 03.xii.2010. Dias, P.G.B.S. col. (MZSP). Paratype: BR [Brazil], Paraná, Céu Azul. Parque Nacional do Iguaçu. 10.x.2012. Dias, P.G.B.S. col. (PSD36) (MZSP). Etymology. The name naipi is given in allusion to the myth of the creation of the Iguaçu Falls, as told by the naipi people. According to the myth, Taroba was a warrior that fell in love with Naipi, daughter of the tribal chief. However, Naipi was promised to the snake-god M’Boy. During a ceremony, the couple ran away using a small boat through the Iguaçu river, and was persecuted by M’Boy. The furious god dug the river, creating the waterfalls, in which the couple fell out. Then, M’Boy turned Taroba in a palm tree and Naipi in a stone, at the Iguaçu Falls. Diagnosis. Within the genus, E. naipi n. sp. can be recognized by the following characters: males FWs square, reaching the 4th abdominal tergite; mirror sub-triangular, with four cells; harp with four well-visible veins; lateral field with transversal veins. Phallic complex elongated; pseudepiphallic arms lateral, straight, apex upcurved, without membranous sphere; sclerite A ventral, reaching PsP1, not fused with pseudepiphallic arms; dorsal ectophallic projections (D.Ec.P.) weak, thin, barely visible; endophallic sclerite proximal, sub-triangular, without endophallic apodeme. Description. General body coloration yellowish to medium brown, almost uniform (Fig. 4A). Head. Dorsum with soft pubescence, yellowish brown, with two lines thin, medium brown toward the vertex. Vertex and fastigium medium brown. Fastigium little wider than long, narrower than scape, slightly narrower toward the apex, with a double row of small bristles, and without the spot observed in E. chape sp. n. Ocelli absent. Eyes obovate, prominent. Antennal scape light to medium brown. Antenna not annulated; antenomeres medium brown (Figs 4A, B). Frons light brown, almost uniform, maculae inconspicuous. Gena light brown (Fig. 4B). Clypeus, labrum and mandibles light brown. Maxillary palpi elongated, thin; joints 1–2 whitish, joints 3–5 elongate, medium brown; joint 5 the longest, apical third of joint 5 curved, apex rounded, whitish (Figs 4A, B). Thorax. Pronotum DD wider than long, slightly pubescent, general coloration medium brown (Figs 4A–C); anterior portion with two spots yellowish brown with reticulated lines medium brown (Figs 4B, C); two blotches large, yellowish brown, on median portion, divided by stripe medium brown, which is divided by a line yellowish brown that goes from the anterior to posterior portion of DD (Figs 4B, C); posterior part with two spots yellowish brown (Fig 4C); DD cephalic and caudal margins sub-straight (Figs 4B, C); LL light brown; ventro-cephalic angle slightly salient, rounded, ventro-caudal angle almost straight, ventral margin gradually ascendant (Figs 4A, B). Legs. Legs I and II yellowish to medium brown, not annulated (Fig. 4A). Auditory tympanum present on both sides of TI; internal

NEW ENDECOUS FROM THE IGUAÇU NATIONAL PARK Zootaxa 4237 (3) © 2017 Magnolia Press · 461 tympanum oval, small, barely visible; external rounded. TI with two same-sized ventral apical spurs; TII with two same-sized ventral apical spurs, and one dorsal, internal. FIII light to medium brown, with diagonal medium brown stripes, distal half medium brown (Figs 4A, B). TIII medium brown; subapical spurs 4/4, the first smaller and associated to the apical; serrulation between and above subapical spurs 1 and 2; apical spurs 3/3, more developed on inner face; dorsal apical spurs the largest on both faces (Fig. 4G). Basitarsus I, II and III medium brown; basitarsus III with double row of spines (Fig. 4G). Abdomen. Medium to yellowish brown, pubescent (Fig. 4A). Supra anal plate medium brown, with spots yellowish brown and macula central, yellowish brown (Figs. 4D, E); anterior margin concave, posterior margin rounded, without distal projections (Figs. 4D, E). Subgenital plate wider than long; anterior margin sub-straight, posterior margin broad, rounded, without distal projections (Fig. 4F). Cerci long, light to medium brown (Fig. 4A). Male. FWs membranous, square, reaching the 4th abdominal tergite (Figs 4A–C); right FW hard, medium brown (Figs 2B, 4A–C); left FW membranous, transparent; stridulatory vein present, with ca. 83 - 87 teeth (n=02) (Figs. 2B, 4A, C); mirror sub-triangular, with four cells (Figs 2B, C, 4C); harp with four veins (Figs 2B, C, 4C); lateral field with transversal veins (Figs 2B, D). Male genitalia. Phallic complex elongated (Figs 5A, C, D). Pseudepiphallus: median part of the pseudepiphallic sclerite fused; pseudepiphallic arms lateral, straight, apex upcurved, without membranous sphere (Figs 5A, B); sclerite A ventral, reaching PsP1, not fused with pseudepiphallic arms (Figs 5C–E). PsP1 and PsP2 connected, not forming a circular structure (Fig. 5C); PsP2 curved inwards (Figs 5A, B). Ectophallic invagination: ectophallic apodeme short, but longer than in E. chape n. sp., robust (Fig. 5A); ectophallic arc below the median part of the pseudepiphallic sclerite (Fig. 5A); dorsal ectophallic projections weak, thin, barely visible (Figs 5A, D); ventral ectophallic projections long, apex straight, rounded (Figs 5D, E); ectophallic fold membranous. Endophallus: endophallic sclerite proximal, sub-triangular (Figs 5A, D), without endophallic apodeme. Female. Unknown. Measurements (mm). Males (n=2): HW—3.6; IOD—1.9; PL—3.1 (3.2–3); AWP—3.65 (3.6–3.7); PWP— 5.05 (5–5.1); PW—5.25 (5.1–5.4); FWL—8.8 (8.6–9); FWW—6.55 (6.4–6.7); LFIII—13.8 (13.6–14); WFIII—4 (3.9–4.1); LTIII—14.5 (14–14.1); LBt–III—4.9 (4.8–5).

TABLE 1. Calling song parameters of Endecous chape n. sp. Spp.—specimens; PD—phrase duration; SP/P—number of subphrase per phrase; PPS—number of pulses per subphrase; SP—subphrase period; SPD—subphrase duration; SFI— subphrase interval; DF—dominant frequency (kHz); T—temperature oC. Spp. PD SP/P PPS SP SPD SFI DF T EFM02 2.23± 0.17 11.64±1.15 5.50±1.93 0.19±0.04 0.12±0.04 0.08±0.02 4.92±0.12 27ºC (2–2.59) (10–14) (3–12) (0.13–0.40) (0.06–0.26) (0.05–0.16) (4.74–5.42) n=11) n=11) n=127 n=116 n=127 n=116 n=127 EFM04 1.80±0.20 11.43±1.54 3.99±0.10 0. 16±0.03 0.07±0.04 0.10±0.02 3.73±0.05 27ºC (1.55–2.34) (9–14) (1–7) (0.11–0.28) (0.02–0.5) (0.06–0.21) (3.47–3.78) n=14 n=14 n=146 n=133 n=144 n=133 n=146 ENIM03 1.40 5 5.2±0.4 0.31±0. 03 0.10±0.01 0.21±0.02 4.32±0.04 23ºC n=1 n=1 (5–6) (0.28–0.35) (0.09–0.11) (0.19–0.24) (4.3–4.4) n=5 n=4 n=5 n=4 n=5 ENIM04 2.41 8 3.67±0.47 0.29±0.03 0.08±0.01 0.21±0.02 4±0.2 23ºC n=1 n=1 (3–4) (0.25–0.35) (0.06–0.09) (0.19–0.26) (3.8–4.3) n=9 n=8 n=9 n=8 n=8 ENIM09 2.35 9 4.67±0.82 0.28±0.04 0.11±0.02 0.18±0.02 3.85±0.07 23ºC n=1 n=1 (3–6) (0.21–0.34) (0.06–0.14) (0.15–0.22) (3.8–4) n=9 n=8 n=9 n=8 n=8 ENIM013 2.85 12 4.5±1.38 0.25±0.07 0.10±0.03 0.15±0.03 3.86±0.07 23ºC n=1 n=1 (1–6) (0.15–0.38) (0.01–0.14) (0.11–0.24) (3.8–4) n=12 n=11 n=12 n=11 n=11 Mean 2.02±0.34 11.10±1.90 4.63±1.61 0.19±0.05 0.09±0.04 0.10±0.04 4.22±0.57 N=6 (1.40–2.85) (5–14) (1–12) (0.11–0.41) (0.01–0.50) (0.05–0.26) (3.47–5.42) n=29 n=29 n=324 n=295 n=322 n=295 n=321

462 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. FIGURE 4. Endecous naipi n. sp. General morphology. A, male habitus; B, male head, pronotum and FW, lateral; C, male head, pronotum and FW, dorsal; D, male supra anal plate; E, male terminalia, showing the phallic complex; F, male subgenital plate; G, male TIII and basitarsus.

NEW ENDECOUS FROM THE IGUAÇU NATIONAL PARK Zootaxa 4237 (3) © 2017 Magnolia Press · 463 FIGURE 5. Endecous naipi n. sp. Male genitalia in dorsal (A), dorsolateral (B), lateral (C), and ventral view (D, E). Conventions: Arc—ectophallic arc; Arm—pseudepiphallic arm; D.Ec.P.—dorsal ectophallic projection; Ect.Ap.—ectophallic apodeme; Ect.F.—ectophallic fold; End.Ap.—endophallic apodeme; End.Sc.—endophallic sclerite; M.S.—membranous sphere; PsP1—pseudepiphallic paramere 1; PsP2—pseudepiphallic paramere 2; V.Ec.P.—ventral ectophallic projection; Sc.A—sclerite A of the pseudepiphallic arms. Scale bar: 1mm.

464 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. Calling song of Endecous chape n. sp. (Fig. 6, Tab. 1). Phrase duration 2.02±0.34 (1.40–2.85, n=29); number of subphrases per phrase 11.10±1.90 (5–14, n=29); number of pulses per subphrase 4.63±1.61 (1–12, n=324); subphrase period 0.19±0.05 (0.11–0.41, n=295); subphrase duration 0.09±0.04 (0.01–0.5, n=322); subphrase interval 0.10±0.04 (0.05–0.26, n=295); dominant frequency 4.22±0.57 (3.47–5.42, n=321); phrase amplitude may gradually increase or remain constant (Figs 6B–D). Note: Males emit phrases sporadically along the night. Therefore, we registered two individuals (specimen EFM02 and EFM04, Tab. 01) along one uninterrupted hour in laboratory condition at 27oC, and they produced interval between phrases of 320.2s±230.03 (91–806, n=10) and 263.84s±193.64 (2.7–671, n=14), respectively. Chromosomes of Endecous chape n. sp. This species is 2n=19, X0♂, XX♀, with three pairs of metacentrics (pair 1, centromeric index (ci) = 44.8; pair 4, ci = 42.9; pair 5, ci = 46.2), four pairs of submetacentrics (pair 2, ci = 31.6; pair 3, ci = 28.6; pair 6, ci = 30.8; pair 9, ci = 33.3), two pairs of acrocentrics (pair 7, ci = 9.1; pair 8, ci = 9,1); the X chromosome is metacentric, ci = 41.6 (Fig. 7).

FIGURE 6. Calling song of Endecous chape n. sp. (A) Sonogram of a phrase with 11 subphrases; (B, C and D) Oscillogram of three phrases showing amplitude variation.

FIGURE 7. Karyotype of Endecous chape n. sp. with 2n=19, X0.

Discussion

Phallic complex of Endecous. The new species were compared with the types of Endecous (Pedroecous) apterus, E. (Endecous) aguassay, E. (Endecous) alejomesai, and E. (Endecous) betariensis from MZSP; topotypes of E. (Notendecous) onthophagus and E. (Endecous) peruassuensis, provided by Dr. Edison Zefa and Dr. Márcio Bolfarini; and individuals of E. (Endecous) arachnopsis, E. (Notendecous) lizeri and E. (Notendecous) onthophagus, studied by the senior author at the ZIN. Thus, the comparisons made encompass all the subgenera of Endecous, and reach almost all of its taxonomical diversity. The main characters used to define and delimitate the species described here are the male forewings and genitalia, which comprises the main character sets used in the taxonomy of Endecous (Zefa et al., 2010; Gorochov, 2014; Souza-Dias et al., 2014; Zefa et al., 2014; Bolfarini & Bichuette, 2015).

In Luzarinae, the development of pseudepiphallic arms are considered a putative synapomorphy and are the most conspicuous part of the male genitalia, with distinct morphological patterns and complexity among the subfamily (Souza-Dias & Desutter-Grandcolas, 2014). Thus, the pseudepiphallic arms are a good source of information for morphological datasets and phylogenetic studies. Nevertheless, the evolution and morphology of the male genitalia in crickets is still discussed, with several terminologies proposed (e.g. Chopard, 1961; Randell, 1964; Alexander & Otte, 1967; Desutter-Grandcolas, 1987, 1990; Desutter-Grandcolas, 2003; Gorochov 1995, 2002, 2013). Regarding Luzarinae, Desutter-Grandcolas (1992, 1993a, b) identified three sclerites on the pseudepiphallic arms, namely sclerites A, B and C. In genera as Eidmanacris, Lerneca, Luzarida, Luzaridella and Strinatia, it is possible to identify some projections or sclerites associated to the pseudepiphallic arms, which correspond to the sclerites originally described by Desutter-Grandcolas (op. cit.). In these genera, a ventrolateral sclerotized projection is present, adjacent to the pseudepiphallic arms (or connected to it), which fits in the definition of the

466 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. sclerite A: “more or less clearly disconnected from epiphallic arm by sclerotization discontinuity, and separating it from epiphallic parameres” (Desutter-Grandcolas, 1993a). Therefore, in this study we consider this projection as the sclerite A of the pseudepiphallic arm. The sclerite A is present in Endecous, and its shape and position is variable within the genus—this structure was recognized in previous studies (Gorochov, 2014; Zefa et al., 2014). In E. chape n. sp., the sclerite is fused to the pseudepiphallic arm, while in E. aguassay, E. alejomesai, E. apterus, E. betariensis, E. peruassuensis, and E. naipi n. sp. the sclerite is separated from the pseudepiphallic arms at the basis. The pseudepiphallic parameres are also very informative structures for taxonomic and phylogenetic studies (Souza-Dias et al., 2015a, b). In Endecous, as in several Luzarinae genera, these structures are highly variable, providing good characters for genera and species delimitation. In E. chape n. sp. the pseudepiphallic parameres are medial, and the PsP1 and PsP2 are fused, forming a circular structure. The PsP1 is sclerotized, curved inwards, large and connected to the small PsP2—the connection is visible in dorsal and superior views. In E. naipi n. sp. the PsP1 is smaller, connected to the distal, parallel, slightly incurved PsP2—the whole structure is not circular as in E. chape n. sp.. This pattern is observed in other Endecous species, as E. aguassay, E. apterus, E. arachnopsis, E. betariensis, E. onthophagus, and E. peruassuensis. In several Endecous species, the pseudepiphallic sclerite is modified, mainly its median part. In E. naipi n. sp. and E. chape n. sp. this portion is fused, while in other species as E. aguassay, E. alejomesai, and E. peruassuensis, this structure is disconnected at the median line. The ectophallic invagination comprises the intermediate portion of the crickets’ phallic complex. In the Phalangopsidae, this structure is composed of a pair of proximal, ventral apodemes (ectophallic apodemes), connected by a median bridge (ectophallic arc), and two pairs of distal projections: a pair of dorsal ectophallic projections, and a pair of ventral ectophallic projections—the latter is connected to the PsP1. The ectophallic dorsal projections are highly modified among Luzarinae, and are reduced or absent in several genera. In other taxa, as Eidmanacris and Endecous, the structure is present, but varying in size, sclerotization and length. In Endecous, the ectophallic dorsal projection is developed and visible in several species, as E. aguassay, E. apterus, E. arachnopsis, E. betariensis, E. onthophagus, E. peruassuensis, E. chape n. sp., and E. naipi n. sp. However, in E. chape n. sp., the ectophallic dorsal projection is totally separated from the phallic complex (Figs 3E, F)—in E. onthophagus, the ectophallic dorsal projection is also separated from the genitalia, but connected to it by a membrane. In the species cited above, the ectophallic dorsal projection, although well visible, is not separated from the genitalia. The inner portion of the phallic complex comprises the endophallus. Its structures were used by Gorochov (2014) to propose the subgenera of Endecous. According to the author, the subgenus Notendecous comprises species with rachis (=ectophallic fold) very large, significantly protruding behind epiphallic posterolateral arms (=pseudepiphallic arms), lamellar, and almost completely membranous, and formula (=endophallic sclerite) very long. The author grouped two species in this subgenus: E. (Notendecous) lizeri and E. (Notendecous) onthophagus. The subgenus Pedroecous comprises only the species E. (Pedroecous) apterus. The subgenus Endecous comprises species with rachis not very large, not protruding behind epiphallic posterolateral arms, more or less finger-like but not lamellar, and partly membranous or semi-sclerotized, and formula moderately long or short. All remaining species are grouped in this subgenus, including E. chape n. sp. and E. naipi n. sp. The endophallus in Endecous is also informative and a good source of morphological characters. In E. chape n. sp., the endophallic sclerite is medial, long, grooved, forming a medio-dorsal crest, and with small, paired endophallic apodemes—this pattern is more or less similar to E. arachnopsis and E. betariensis. In E. naipi n. sp., the endophallic sclerite is proximal, small, triangular, and without endophallic apodemes—this pattern is observed in E. aguassay, E. apterus and E. peruassuensis. For that reason, all the layers of the male genitalia (pseudepiphallus, ectophallic invagination and endophallus) and related structures in Endecous, as well as in Luzarinae, are very informative for taxonomic and systematic studies. A comprehensive comparative and morphological study of male genitalia in Phalangopsidae, mainly Luzarinae, is much needed in order to clarify the genital morphology of the family and identify homologies. Distribution, habit and habitat data. In the Neotropics, Endecous occurs in eight biogeographical provinces, according to Morrone’s regionalisation (2014), and in Brazil, Endecous’ species are found in almost all biomes, excepting the Amazon (Fig. 6). Endecous chape n. sp. and E. naipi n sp. are the 4th and 5th species of cricket described for the Iguaçu National Park, or adjacent areas. The other species described are Adelosgryllus rubricephalus Mesa & Zefa, 2004, Taroba

NEW ENDECOUS FROM THE IGUAÇU NATIONAL PARK Zootaxa 4237 (3) © 2017 Magnolia Press · 467 elephantina de Mello & Souza-Dias, 2010, and Brazitrypa paranaensis (de Mello & Souza-Dias, 2010) (Mesa & Zefa, 2004; Dias & de Mello, 2010a, b). The Iguaçu National Park is located at the west of Paraná State, on the border with Argentina. That area, together with Santa Catarina State (Brazil), and part of the provinces of Alto Paraná (Paraguay) and Misiones (Argentina), is covered by the Brazilian Atlantic Forest. There, two phytophysiognomies of this forest are found, the Atlantic Semideciduous Forest, occupying lowlands (150 to 700m), and the Ombrophylus Mist forest (Araucaria forest), in altitudes from 750m on northern and northeastern of the Iguaçu National Park (Joly et al., 1999; Curcio et al., 2007; Ortiz, 2010). The altitude on that area varies from <200 to 800 meters and the main soil type, the red latosol, is a result of extensive volcanic flows, occurred between late Jurassic and early Cretaceous (145–120 mya) (Salamuni et al., 2002). There are no records of caves in that area, and the nearest karst areas, the Ribeira Valley (São Paulo State) and Bodoquena (Mato Grosso do Sul State) are at least 800km far. Thus, there are no troglobitic species of crickets in southern Brazil—the three known troglobitic species from Brazil, Endecous apterus, E. cavernicolus and E. peruassuensis are recorded from caves in Bahia and Minas Gerais States (Bambui karst area), respectively (Souza- Dias et al., 2014; Bolfarini & Bichuette, 2015). Therefore, regarding the use of habitat, the species E. naipi n. sp. and E. chape n. sp. are cavicolous- straminicolous. Individuals were observed during the night occupying the leaf litter and small natural cavities, as gullies, crevices and burrows. The species E. chape n. sp. seems to have a large population, and its individuals are relatively easy to find—this species is observed since the first field studies on the Iguaçu National Park area, made by Alejo Mesa and Francisco de A. G. de Mello in 1985. Collections made since 2006 resulted in the capture of several individuals of E. chape n. sp., while in all these years, only two males of E. naipi n. sp. were found. Probably, this species has small populations or the individuals are secretive and not easy to find, although they are larger than the individuals of E. chape n. sp. The females of Endecous species are very similar, and the females of E. naipi n. sp. are unknown—the type material of females of E. chape n. sp. is composed of females that were observed mating with conspecific males. A molecular analysis of Luzarinae crickets are being carried out, and is expected that the results will improve the delimitation of these species. Bioacustics. The calling songs of Endecous itatibensis and E. alejomesai are similar, with phrases composed of pairs of pulses, but diverging in dominant frequency with 3.2 kHz and 4.4 kHz, respectively. On the other hand, E. betariensis and E. chape n. sp. show phrases without pair of pulses, instead group of pulses, here denominated as “subphrases”. The number of pulses per subphrase is different between both species, but it is not possible to compare accurately other song parameters since data published by de Mello & Pellegatti-Franco (1998) is not sufficient. Furthermore, both species have calling songs with the different dominant frequency being about 3 kHz in E. betariensis, and ranging from 3.7 to 4.9 kHz in E. chape n. sp. Due to the difficulties to sampling E. naipi n. sp., the calling song of this species was not recorded. The calling song of E. chape n. sp. has peculiarities that are not common in other species of Grylloidea whose calling song carries rhythmical repetitions of the same phrases (Alexander, 1957; Leroy, 1969). Here we found that the calling of E. chape n. sp. varies not only at phrases rhythm emission, as in physical and temporal parameters of each phrase. It will be necessary to analyze large samples of individuals to understand the range of the calling song variation, as well as parameters used for females to found males. For now, all detected variation in the calling song parameters of the six individuals here described must be used to recognize the new species. Cytogenetics. The karyotype of E. chape n. sp. presents the same chromosome number as E. onthophagus and E. itatibensis, and chromosome morphology similar to E. onthophagus with the same proportion of metacentric, submetacentric and acrocentric chromosomes. However, they differ in metacentric (E. chape n. sp. pairs 4 and 5; E. onthophagus and E. itatibensis pairs 7 and 8) and acrocentric (E. chape n. sp. pairs 7 and 8; E. onthophagus pair 4 and 5, and E. itatibensis pair 5). The karyotypes of E. onthophagus and E. itatibensis differ from each other in the pair 4, which is submetacentric in E. itatibensis. In these species, there are lower acrocentric chromosomes than E. betariensis, E. alejomesai, and E. cavernicolus, with 2n=21, X0, and karyotypes including 5, 6 e 7 pairs of acrocentric chromosomes, respectively. In Orthoptera, the most derived karyotypes present larger amounts of metacentric chromosomes, which result of centric fusion between acrocentric bivalents, with consequent reduction of diploid number (White 1951; White, 1973). In this perspective, the basic chromosome number in Endecous is 2n=21, X0, highlighting E. ubajarensis karyotype, with 2n=14, X1X20, including only meta and submetacentric chromosomes. Additionally, E. ubajarensis

468 · Zootaxa 4237 (3) © 2017 Magnolia Press SOUZA-DIAS ET AL. presents the most derived karyotype to date, with reduction in chromosome number involving one or more fusion events or translocation X/autosome that originated the sex system mechanism determination X1X20 (Zefa et al. 2010; Zefa et al. 2014).

Acknowledgments

The authors would like to thank the administration of the Iguaçu National Park for the support during the field work, especially Apolonio N. Rodrigues, Marina X. Silva, Pedro Fogaça and Raphael Xavier; Dr. Márcio P. Bolfarini for the loan of Endecous peruassuensis specimens; and Dr. Andrej Gorochov, for the support during PhD internship in the Zoological Institute of the Russian Academy of Sciences, in Saint Petersburg (ZIN). Financial support received from CAPES (Sandwich Doctorate Scholarship, Proc. 5568-13-7 and PNPD 20131527).

References

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