Arumatia dubia Ghirotto & Crispino & Engelking & Neves & Góis & Chiquetto-Machado 2022, gen. et comb. nov.

Arumatia dubia (Caudell, 1904) gen. et comb. nov. Figs 1–13, 46, 52, 53A–D Bacunculus dubius Caudell, 1904: 186. Bacunculus fragilis Brunner von Wattenwyl, 1907: 336. Syn. nov. Echetlus evoneobertii Zompro & Adis, 2001: 294. Syn. nov. Heteronemia dubia – Zompro 2001: 223. — Otte & Brock 2005: 158. Heteronemia fragilis – Zompro 2001: 223. — Otte & Brock 2005: 158. Echetlus evoneobertii – Zompro 2004b: 138. — Otte & Brock 2005: 132. — Zompro & Domenico 2005: 257. — Zompro et al. 2006: 131 (specimen record for Australia in error, misidentification). Candovia evoneobertii – Brock & Hasenpusch 2007: 7, 70; 2009: 151. — Araujo & Garraffoni 2012: 235. Diagnosis Females From Arumatia fulgens gen. et comb. nov., A. dubia gen. et comb. nov. differs by the shorter subgenital plate not exceeding half the length of the tergum X, the slightly shorter cerci (not reaching the length of terga VIII–X combined) and the thinner body. From A. anyami gen. et sp. nov. it differs by the lack of a black spot on the ventral region of the prothorax, the longer median segment, equal in size to the metanotum (vs ¾ the length of metanotum), the shorter subgenital plate, larger epiproct, posterior margin of anal segment less emarginate, tergum VII having a widened and incurved posterior margin and shorter gonapophyses VIII, and narrower gonoplac. Some mouthparts differ between the two species, because A. dubia has a different microtrichia pattern as a band on the galea and not a circle, has less and stouter setae on the lacinia, a narrower galea and narrower labial palp segments than those of A. anyami. From A. crassicercata gen. et sp. nov. it differs by the lack of a praeopercular organ, the longer median segment, much longer cerci and thinner legs with less prominent keels. Arumatia dubia also has fairly less and stouter setae on the lacinia and shorter microtrichia of the galea than A. crassicercata, and the microtrichia are arranged in a band, not a circle. From A. aramatia gen. et sp. nov. it differs by the shorter, less elongate head, the more elongate galea, the longer median segment, as long as the metanotum, the shorter terga VIII–X, the emarginate posterior margin of tergum X, the slightly shorter subgenital plate, the lack of a praeopercular organ and the shorter gonapophyses VIII. From A. motenata gen. et sp. nov. and A. diamante gen. et sp. nov. it differs by the shorter, less elongate head, the more elongate galea, the shorter microtrichia of the galea, the longer median segment, as long as the metanotum, the presence of stronger spiniform setae on the carinae of the tarsi and on the apex of mid and hind tibiae, the absence of an apical sessile spine on the apex of the ventral, antero- and posteroventral carinae of the tibiae, the slightly longer cerci, the shorter epiproct, the longer subgenital plate, the absence of a praeopercular organ, the narrower gonoplac and the shorter gonapophyses VIII. Eggs The egg of Arumatia dubia gen. et comb. nov. has a smoother capsule than that of A. anyami gen. et sp. nov., and a lumpier and grosser texture (rather than finely punctuated) than that of A. motenata gen. et sp. nov. or A. crassicercata gen. et sp. nov. The egg also has a reticular, netted aspect of the keels of the capitulum rather than radial on the capitulum of the eggs of A. aramatia gen. et sp. nov., A. motenata and A. diamante gen. et sp. nov. as well as a widened posteriormost region of the micropylar plate and dorsal region less convex in lateral view than that of A. crassicercata. Type material PARAGUAY • 1 ♀, holotype of Bacunculus dubius; Sapucay; W.T. Foster leg.; “10-3[?]1/[?]”, “ Bacunculus dubia ♀ type. Caudell; Type No. 8027. U.S.N.M.”; USNM 8027 (examined from photo). SOUTH AMERICA • 1 ♀, holotype of Bacunculus fragilis; “America merid.” [South America]; Bol. leg.; MNMS (examined from photo). BRAZIL • 1 ♀, holotype of Echetlus evoneobertii; São Paulo, Boa Esperança do Sul; 18 Feb. 1998; J.A. Cerignoni leg.; MZUSP 0035 (examined) • 1 ♀, eggs, paratypes of Echetlus evoneobertii; São Paulo, Piracicaba, ESALQ / USP; Sep. 2000; “criado em laboratório em folhas de Eucalyptus ” [raised in lab with Eucalyptus leaves]; MZUSP 0036 (examined) • 1 ♀, paratype of Echetlus evoneobertii; same collection data as for preceding; MZUSP 0037 (examined) • 1 egg, paratype of Echetlus evoneobertii; same collection data as for preceding; ANIC (examined from photo) • 1 ♀, paratype of Echetlus evoneobertii; same collection data as for preceding; ANIC (not examined) • 5 ♀♀, paratypes of Echetlus evoneobertii; same collection data as for preceding; MZUSP (not traced) • 2 ♀♀, paratypes of Echetlus evoneobertii; São Paulo, Boa Esperança do Sul; 18 Feb. 1998; J.A. Cerignoni leg.; ZMUK 457-1, 457-3 (not traced) • 5 eggs, paratypes of Echetlus evoneobertii; same collection data as for preceding; ZMUK 457-7 (not traced). Other material examined BRAZIL – Paraná • 1 ♀; Londrina; 6 Apr. 1998; E.P. Frazão leg.; ESALQENT 000438. – Distrito Federal • 1 ♀; Brasília, urban area near Parque Olhos D’água; 15°44′56.6″ S, 47°53′25.1″ W; 2020; J. Góis and P. Souza leg.; MZUSP. – Mato Grosso do Sul • 2 ♀♀; Chapadão do Sul, margin of Sucuriú River; 19°28′31.3″ S, 52°32′17.7″ W; 30 Nov. 2020; E.B. Crispino and D. Castro-Pereira leg.; in cerradão [deciduous forest]; MZUSP V0534, V0535 • 2 ♀♀, nymphs, eggs; same collection data as for preceding; MZUSP. – Minas Gerais • 3 ♀♀; Paraopeba; 18 Jun. 1988; J.A. Cerignoni leg.; in Eucalyptus sp.; ESALQENT 000439, 000444, 000451 • 1 ♀; Delfinópolis, Pousada Cachoeira Paraíso, near Serra da Canastra; 20°20′35.5″ S, 46°47′14.0″ W; E.W. Engelking leg.; in cerrado [savannah formation]; UNESP • 1 ♀; Araguari, near Emborcação Dam; 18°27′23.0″ S, 48°00′40.7″ W; 20 Jan. 2021; V.M. Ghirotto leg.; in cerradão [deciduous forest]; MZUSP V0547 • eggs; same collection data as for preceding; from female V0547; MZUSP • 1 ♀, first instar nymph; Minas Gerais, Araguari; reared by V.M. Ghirotto 2021 from female V0547; MZUSP V0556 • 1 ♀, second instar nymph; same collection data as for preceding; MZUSP V0557 • 1 ♀; same collection data as for preceding; MZUSP V0676. – São Paulo • 1 ♀; Piracicaba, ESALQ; Jun. 1992; J.A. Cerignoni leg.; host plant Psidium guajava; Echetlus evoneobertii ♀ det. O. Zompro Nov. 2003; ESALQENT 000443 • 1 ♀; Piracicaba; Jun. 2001; S.S. Prado leg.; ESALQENT 000442 • 1 ♀; Piracicaba; 16 May 2000; R.A. Polanczyk leg.; ESALQENT 000436 • 1 ♀; Piracicaba, ESALQ; 18 May 1998; S.R. Magro leg.; ESALQENT 000450 • 1 ♀; Piracicaba, ESALQ, Dept. Entomology; 17 May 1998; V.D.A. dos Reis leg.; ESALQENT 000440 • 3 ♀♀; Limeira; 23 Jan. 2004; P. Milano leg.; ESALQENT 000446, 000447, 000448 • 1 ♀; Limeira; 10 Jun. 1999; “criação particular” [culture stock] P. Milano; ESALQENT 000445 • 1 ♀; Descalvado; 26 Apr. 1968; Renato leg.; ESALQENT 000441 • 1 ♀; “R. Preto” [Ribeirão Preto]; Lordello leg.; “4-51”; ESALQENT 000437 • 1 ♀; “Boa Esperança” [Boa Esperança do Sul]; 21 Apr. 1998; R.B.Q. Silva leg.; ESALQENT 000449 • 8 ♀♀; Ibaté; Jan.–Jul. 2019; reared by V.M. Ghirotto, culture obtained at ESALQ entomology lab; MZUSP 1222, 1224, 1227, V0314, V0315, V0333, V0415, V0503 • 1 ♀, first instar nymph; same collection data as for preceding; MZUSP V0272 • eggs; same collection data as for preceding; MZUSP • 2 ♀♀; same collection data as for preceding; 2021; MZUSP V0542, V0558 • eggs; same collection data as for preceding; MZUSP • 2 ♀♀; Assis, Campus UNESP; 22°39′00.2″ S, 50°26′18.8″ W; Aug. 2018; P.W. Engelking leg.; UNESP WE059 • 2 ♀♀; Echaporã, Trilha da cachoeira Stn Rosa; 22°25′23.8″ S, 50°11′60.0″ W; Sept. 2020; P.W. Engelking and G.A. Nunes leg.; UNESP WE060. Remarks The Australian specimen of Arumatia dubia gen. et comb. nov. reportedly housed at the Zoological Museum of the Christian-Albrechts University of Kiel in Germany (as Candovia evoneobertii) could not be traced (Thies Büscher, pers. comm., 2021) and no description or illustration of this specimen was provided by Zompro et al. (2006). It is very likely that this specimen was misidentified and represents some distinct species native to Australia (Paul Brock, pers. comm., 2021). Redescription Female MEASUREMENTS (in mm, N = 3). Body (without cerci) 69.1–72.4, head 4–4.1, antennae 43.5–48.5, pronotum 2.4–2.6, mesonotum 15.7–16.6, metanotum 4.5, median segment 4.5, abdomen (excluding median segment, with cerci) 37.6–39.4, cercus 4.7–5.0, profemur 15.9–16.4, protibia 16.5–17.2, mesofemur 11.2–11.8, mesotibia 10.8–11.6, metafemur 14.2–14.9, metatibia 15.5–16.9. COLOUR (Figs 1–2, 13). Body generally light to dark green, yellow, beige, salmon, orange, brown, grey or black with or without light granulations on body and legs (mostly femora) and with or without irregular stains of different tones (Fig. 13). Some individuals with longitudinal band running along entire body, slightly darker than body to black (Fig. 13A, F). Eyes same colour as rest of body but darker, or reddish brown. Antennae darker ventrally. Apical palp segments and other mouthparts in same colour as body or brownish to beige. HEAD (Figs 3–4). Elongate, smooth, with sparse setae mostly on dorsal region, vertex flat but gently convex at posterior margin to fit underneath the pronotum, frontal convexity developed and frontal suture round (Fig. 3A–D). Eyes small and slightly elongate, approximately 0.2× as long as head. Cervix covering more than half of head, cervical sclerites elongate and weakly sclerotized. Gula somewhat spatulate, bearing setae and covering ca more than half of cervix (Fig. 3D). Subgena narrow with posterior projection ca as high as eye. Submentum narrow and only slightly curved backwards, mentum simple, prementum somewhat wide. Glossa elongate and rounded, paraglossa larger, roundly falcate and almost reaching labrum. Lacinia with three distal teeth, one large medially and two smaller laterally, with mesal edge bearing bundles of ca 12 large setae from base to sclerotized portion (Fig. 4D). Lobe over base of lacinia almost indistinct, showing as small and gentle round bump. Galea elliptical and bearing long setae (Fig. 4F), apically with dense tuft of hairy, large microtrichia forming band along apical edge. Dorsally and posteriad to this patch ca 29 distinct circular granules of same size as base of setae (Fig. 4E). Galealobulus present, small and widely round, basally fused to galea (Fig. 4D, F). Palpal segments cylindrical. Clypeus wide, approximately elliptical, anterior surface with two parasagittal furrows and anterior margin medially with round, wide notch (Fig. 3C). Labrum strongly notched anteromedially, asymmetric with right lobe significantly larger than left one (Fig. 3C). Left mandible with two smooth and straight edges, dorsal one cutting and sharp, and ventral one blunt, mesal surface with convex globose protuberance near ventral cutting edge, and sinuous furrow between both edges (Fig. 4A, C). Right mandible with sharp dorsal cutting edge, straight but interrupted at border with ventral edge by round edge, ventral edge irregular, level and molariform with two larger projections, mesal surface without protuberances (Fig. 4A–B). Both mandibles with dense row of long setae at base of dorsal margin. Antennae filiform, extending approximately until posterior region of third abdominal segment and exceeding forelegs, scapus ca 1.9× as long as wide, basally constricted in dorsal view and slightly compressed dorsoventrally, pedicellus subglobose, large, more than half the length of scapus; composed of 68–72 segments. Antennomeres bear three types of setae (Fig. 3E–F), a very short, small and densely distributed one, a transparent and porrect one, and a black, slightly more elongate, straight one. From 47 th segment, smaller setae becoming sparser, on segments 62–67 smaller type very sparse and almost absent and on segments 67–72 absent. Other two types of setae gradually becoming thinner and more elongate towards last segments. Antennal bump very discrete on dorsal surface of posterior region of 12 th antennomere (Fig. 3F). THORAX (Figs 1–2, 5). Smooth with scattered setae inserted in paler small bumps. Pronotum slightly longer than wide and slightly constricted pre-medially, anteriorly and posteriorly convex in lateral view, transverse sulcus conspicuous and straight, gently curved in lateral edges, longitudinal median sulcus distinct. Paranota curved and ca 3× as long as wide, procoxopleurite apically round (Fig. 5A– B). Probasisternum tapering towards anterior, profurcasternum round (Fig. 5C). Mesothorax 6× as long as prothorax, as wide as prothorax anteriorly and slightly widening towards posterior region. Mesonotum with pair of distinct lateral carinae, mesepisternum lanceolate and regularly widening posteriorly, mesepimeron slightly elongate, pointing towards posterior and slightly exceeding end of mesothorax. Mesocoxopleurite indistinct, mesofurca Y-shaped. Metathorax continuing pair of lateral carinae of mesonotum, metepisternum long and similar to mesepisternum. Metepimeron extremely elongate, extending across entire length of median segment, posteriorly pointing and slightly exceeding posterior region of median segment. Metacoxopleurite very discrete but elongate, metafurca Y-shaped. Metanotum and median segment of about same length (Fig. 5E). Median segment anteriorly marked by two parasagittal ovoid stains (Fig. 5E), continuing pair of thoracic lateral carinae. Posterior margin of metanotum and anterior margin of median segment slightly widened (Figs 1, 13). LEGS (Figs 1–2, 6). Slender. Hindlegs slightly extending beyond epiproct but not cerci, anterior legs ca as long as hindlegs, midlegs distinctly shorter. Midlegs slightly longer in some individuals. Coxae smooth (Figs 5B–C, 6A). Profemora of around same length as mesothorax, mesofemora about 0.7 × as long as profemora and metafemora slightly shorter than mesothorax. Protibia varying from slightly shorter to longer than profemur, mesotibia ca as long as mesofemur, metatibia slightly longer than metafemur. Profemur with distinct basal curvature (Fig. 6A). Femora and tibiae with five carinae without setae between them (Fig. 6B). Carinae of profemora and protibiae distinctly keeled, mid and hind femora and tibiae weakly keeled. Antero- and posteroventral carinae of femora with apical toothed prominence. Carinae of all femora, tibiae and tarsi bear row of short setae, last two setae of all five carinae of meso- and metatibiae stouter and spiniform (Fig. 6B–F). In tarsi, setae on ventral carinae of probasitarsi longer (but not stouter), while setae on ventral, antero- and posteroventral carinae of meso- and metabasitarsi stouter and spiniform, distributed across entire length except for apex of mesobasitarsi and on basal half to three quarters of metabasitarsi, stronger on ventral carinae (Fig. 6D–F). Some individuals with weaker, not spiniform setae on antero-ventral carinae of metabasitarsi. All basitarsi very elongate, significantly longer than respective following tarsomeres combined (Fig. 6B–E), with hairy setae restricted to apical portion on ventro-lateral patches (Fig. 6C–D). Remaining tarsomeres with setae on ventro-lateral patches in portions not covered by euplantulae. Tarsomeres I–III with discrete dorsal round apical projection. Arolium round and broad, bearing setae dorsally. Pretarsal claws symmetrical with setae dorsally and laterally. Euplantulae well developed in all tarsomeres, composed of two symmetrical pads separated by median groove in tarsomeres I–III, of single pointing domed pad in tarsomeres IV and of single flattened lobed pad in tarsomeres V, present only apically at tarsomeres I–II, covering ca half length of tarsomeres III, covering two thirds length of tarsomeres IV and covering almost entire ventral surface of tarsomeres V (Fig. 6D). ABDOMEN (Figs 1–2, 7–8). External surface as in thorax, bearing few setae across its entire length. Median segment ca as long as metanotum (Fig. 5E). Combined length of segments II–X slightly longer than combined length of head, thorax and median segment. Terga II–VII and sterna II–VIII bearing discrete lateral carinae near lateral margins. All segments longer than wide (Fig. 1). Segment II as long as VII and slightly shorter than III, segments III–VI slightly increasing in size in relation to anterior segment, tergum VIII significantly shorter than preceding segment and slightly longer than IX, tergum X slightly longer than IX. Terga III–VI wider than II, VII and VIII. Terga IX and X slightly narrower than preceding segments. Lateral borders of tergum VII gradually widening and curving downwards and inwards near posterior margin (Fig. 7B–E). Tergum X conical in lateral view (Fig. 7B), in dorsal view just slightly narrower towards apex, posterior margin emarginate with broad round indent showing epiproct (Fig. 7A). Epiproct domed in dorsal view, dorsoventrally flattened, distinctly prominent, exceeding posteriormost margins of tergum X and visible dorsally and laterally (Fig. 7A–B). Paraprocts elongate, posteriorly acute, straight, setose on posterior margin, laterally bearing cerci and not concealing them from ventral or lateral view (Fig. 7C–E). Cerci extremely elongate, straight and basally narrower, fitting paraprocts, pointing to posterior, gradually tapering, slightly longer than terga IX, X and epiproct combined (Fig. 7). Cerci bearing four types of setae (as in nymphal stages, see Fig. 11F–G), three very similar to those of antennae, densely covered in very short, small setae similar to those of antennae, sparser transparent setae inclined towards posterior and black and slightly more elongate straight setae and with inner and ventral surfaces also bearing several sensory hairs significantly thinner than other setae. Praeopercular organ absent (see praeopercular region in Fig. 7C–E). Subgenital plate roundly lanceolate to slightly more acute, reaching medial region of tergum X and bearing two parasagittal carinae beginning at anterior margin and running half length of segment before becoming flatter and more setose, gradually tapering towards posterior and totally covering gonapophyses and gonoplac (Fig. 7C– E). Gonapophyses and gonoplac flattened, dorsoventrally for gonapophyses VIII, lateroventrally for IX and laterally for gonoplac (Figs 7E, 8). Gonapophyses VIII and IX short and of similar length, reaching ca ¾ of length of tergum IX; VIII linear to lorate, blunt to tapering towards posterior, IX lorate to conical, tapering towards posterior (Fig. 8). Gonapophyses IX ventrally folded to fit within gonapophyses VIII. Gonangulum distinctly reduced, flat and not lobed (Fig. 8C). Gonoplac significantly elongate, widened basally, subsequently linear to lorate, bearing setae, exceeding both gonapophyses and reaching ca anterior margin of tergum X (Fig. 8). Distinct triangular sclerotization pointing towards posterior and slightly towards ventral present at dorsal face of gonapophyses, separating them from ventral wall of segment IX. Inside oviduct around median region of segment VIII with triangular and slightly sclerotized structure dorsoventrally flattened and pointing towards posterior. Egg (Fig. 12) Measurements in mm (N = 10): length 2.5, height 1.7–1.8, width 1.4–1.5. Relatively small, capsule ovoid, constricted at opercular collar, laterally compressed and ca 1.8× as long as wide and 1.4× as long as tall (Fig. 12A–D). Capsule surface slightly roughly textured with lumpy or net appearance. Colour varying in shades of orangish or reddish brown, rarely greyish, with lighter band on dorsal, anterior and posterior regions, surrounding operculum and polar region (Fig. 12A–D). Micropylar plate very elongate, occupying large area of dorsal region and varying around 0,6–0,8 × as long as capsule and 2,8–5,7× as long as wide, elliptical with round margins and almost parallel-sided with posterior portion gently but noticeably widened (Fig. 12C–D, G). Micropylar plate with lighter elevated outer circle delimiting darker inner flat region confluent with micropylar cup (Fig. 12C–D). Internal micropylar plate closed, accompanying external micropylar plate, interior surface smooth. Micropylar cup small, rounded, only slightly elevated, dark and anteriorly merged to widened polar edge of micropylar plate. Median line short and of same colour and elevation as elevated margin of micropylar plate, almost reaching polar area (Fig. 12C–D). Opercular collar narrower (constricted) than rest of capsule, smooth and lighter, with minute and delicate bristles surrounding edge (Fig. 12F). Operculum elliptical with irregular reticular and sinuous elevation, with amber texture composing non-stalked capitulum (Fig. 12A–F). Very few eggs present flat, smooth and amber operculum (Fig. 12G–H). Nymphs (Figs 2B, 9–11) FIRST- INSTAR NYMPH (Figs 1, 9–10, 11A–B). Entirely green with black blurred longitudinal lateral line running along head behind eyes, further crossing prothorax and extending to anterior portion of mesothorax. Dorsally green but lighter than lateral and ventral areas delimited by lateral black line. Body with sparse setae and surface with smooth and flat scaly texture. Head and eyes large. Head globose, just slightly elongate (Fig. 9). Gula not sclerotized, visually absent, cervix more than half length of head, slightly wider than that of adults. Subgena similar in shape to that of adults. Labrum notched and asy