Arumatia motenata Ghirotto, 2022

Ghirotto, Victor Morais, Crispino, Edgar Blois, Engelking, Phillip Watzke, Neves, Pedro Alvaro Barbosa Aguiar, Góis, Júlia de & Chiquetto-Machado, Pedro Ivo, 2022, Arumatia, a new genus of Diapheromerinae stick insects (Insecta, Phasmatodea) from Brazil, with the description of five new species and a reassessment of species misplaced in Australian genera, European Journal of Taxonomy 827 (827), pp. 1-85 : 53-70

Arumatia motenata Ghirotto gen. et sp. nov.

urn:lsid:zoobank.org:act:935C1661-AF01-473F-A0B3-884C968257DF

Figs 34–46 , 53G–H

Diagnosis

Females

Differs from all other known females of Arumatia Ghirotto gen. nov., except those of A. diamante gen. et sp. nov., by the very elongate and subrectangular head, larger microtrichia on the galea, the presence of an apical sessile spine on the apex of ventral, antero- and posteroventral carinae of the tibiae, the very elongate epiproct, the shorter subgenital plate and the wider gonoplac in lateral view and from all other known species of the genus by the lobed, well-developed praeopercular organ. It also differs from all other species of the genus except A. diamante and A. anyami gen. et sp. nov. by the absence of stronger spiniform setae on the carinae of the tarsi and on the apex of the mid and hind tibiae. It further differs from A. dubia gen. et comb. nov. and A. fulgens gen. et comb. nov. by the shorter median segment in relation to the metanotum, the longer segment VII in relation to VIII, the slightly shorter cerci and the longer gonapophyses VIII. From A. anyami it further differs by the less emarginate posterior margin of tergum X. From A. crassicercata gen. et sp. nov. it further differs by the shorter median segment in relation to the metanotum and the longer cerci. From A. aramatia gen. et sp. nov. it further differs by the shorter median segment in relation to the metanotum, the shorter cerci and the longer gonapophyses VIII. From A. diamante it further differs by the less elongate head, the relatively longer cerci, the shorter epiproct, and paraprocts and the lack of tubercles on tergum VI.

Males

From Arumatia aramatia gen. et sp. nov., the only other species with known male, it differs by the head with parallel to concave sides slightly widening towards the anterior in dorsal view rather than convex and not widening, the absence of stouter spiniform setae at the posterior region of all carinae of the tibiae, the presence of an apical sessile spine on the apex of the ventral, antero- and posteroventral carinae of the tibiae, tergum IX as long as X, the poculum shortened and round in lateral view, tergum X with lateral flat expansions with thorn pads further extending to the ventral area of the lateral expansions.

Eggs

The egg of Arumatia motenata gen. et sp. nov. differs from the eggs of all other Arumatia gen. nov. whose eggs are known by the not sinuous, somewhat radial elevations of the capitulum and from eggs of A. diamante gen. et sp. nov. it further differs by the constricted opercular collar, not enlarged.

Etymology

This species is named after its unusual male-female attaching mechanism consisting of the posterior margin of the tergum X of the male bearing lateral expansions that fold around the well-developed and prominent female praeopercular organ. The name is a Latinized adjective formed by the old Tupi word ‘ motena ’ (= ‘to attach, to fit’) with the Latin suffix ‘-ata’ meaning ‘shaped to attach’. The old Tupi language was spoken by the indigenous Tupi people of Brazil and is now extinct. However, the word survives with the same meaning in the more recently developed Nheengatu language of the TupiGuarani family which is spoken by some peoples of the Amazon.

Material examined

Holotype BRAZIL • ♀; Minas Gerais, Santana do Riacho, side of highway MG-010, next to Sítio Velozias; 19°16’51.7″ S, 43°34′52.6″ W; 1100–1200 m a.s.l.; 31 Oct. 2021; V.M. Ghirotto leg.; Serra do Cipó plateau , in campo rupestre vegetation; MZUSP V0603 .

Paratypes BRAZIL • 5 ♀♀; same collection data as for holotype; MZUSP V0626 , V0659 , V0662 , V0663 , V0675 • 6 ♂♂; same collection data as for holotype; MZUSP V0620 to V0625 • eggs; same collection data as for holotype; MZUSP .

Other material

BRAZIL • 3 ♀♀; same collection data as for holotype; MZUSP V0660 , V0661 , V0664 .

Description

Female holotype (MZUSP V0603)

MEASUREMENTS (in mm). Body (without cerci) 100.1, head 6.5, antennae 57.0, pronotum 4.5, mesonotum 26.3, metanotum 10.0, median segment 6.7, abdomen (excluding median segment) 57.1, cercus 6.0, profemur 26.5, protibia 27.0, mesofemur 19.5, mesotibia 20.0, metafemur 24.0, metatibia 26.0.

COLOUR ( Figs 35–37 ). Entirely yellowish beige with irregular stains of different tones.

HEAD ( Fig. 35 ). Very elongate, smooth, with sparse small setae, vertex flat but very gently convex at posterior margin to fit underneath the pronotum. Frontal convexity developed, frontal suture round ( Fig. 35A–B ). Eyes small and slightly elongate, approximately 0.15 × as long as head. Cervix covering slightly more than half head, cervical sclerites developed, gula bearing setae and covering slightly less than half of cervix ( Fig. 35C ). Subgena fairly narrow, with posterior projection ca height of middle of eye. Submentum narrow and curved towards posterior, mentum simple, prementum somewhat wide. Glossa elongate, rounded, paraglossa roundly falcate, with round lateral margins and curved inner margins, reaching middle of labrum. Lacinia with three distal teeth, one large medially and two smaller laterally, all prominent, mesal edge bearing bundles of ca 11 large setae from base to sclerotized portion. Galea bearing long setae, slightly elongate, slightly widened before apical portion, apex round. Inner edge of galea apically with dense round tuft of hairy, long, large microtrichia and a few distinct circular granules dorsally and posteriad to this patch. Galealobulus present, small, round and basally fused to galea. Maxillary palpal segments cylindrical and labial slightly widened. Clypeus wide, subrectangular. Labrum strongly notched anteromedially, delimiting two round lobes, asymmetric with right lobe larger. Antennae filiform, reaching at least tergum IV. Scapus ca 2 × as long as wide, slightly compressed dorsoventrally. Pedicellus subglobose, large, slightly more than half as long as scapus ( Fig. 35A–B ). Antennomeres bear three types of setae as in A. dubia .

THORAX ( Figs 34 , 35A , 36A ). Smooth with scattered small and short black setae. Pronotum longer than wide, slightly constricted pre-medially and slightly wider from transverse sulcus towards posterior margin, fairly flat in lateral view. Transverse sulcus of pronotum conspicuous and gently curved towards

posterior, longitudinal sulcus conspicuous and distinct. Paranota curved, slightly widened medially, longer than wide, procoxopleurite apically round and ventrally straight ( Fig. 35A–B ). Probasisternum strongly tapering towards anterior, profurcasternum round ( Fig. 35C ). Mesothorax widened across all of length except near margins, gently convex in dorsal view (similar to paratype, Fig. 34B ). Mesothorax 7 × as long as prothorax. Mesonotum with pair of lateral carinae, mesepisternum lanceolate and regularly widening posteriorly, mesepimeron slightly elongate, medially gently widened, pointing towards posterior and slightly exceeding end of mesothorax ( Fig. 36A ). Mesocoxopleurite small, elongate, mesofurca Y-shaped. Metathorax and median segment widened across all of length except near margins, gently convex in dorsal view ( Fig. 36A ). Metathorax continuing pair of lateral carinae of mesonotum, metepisternum long, similar to mesepisternum, metepimeron extremely elongate, extending across entire length of median segment, posteriorly pointing and slightly exceeding end of median segment

( Fig. 36A ). Metacoxopleurite discrete, slightly elongate, medially gently widened, metafurca T-shaped. Metanotum 1.4 × as long as median segment ( Fig. 36A ). Median segment continuing pair of thoracic lateral carinae but weaker.

LEGS ( Figs 34 , 36B–F ). Slender. Hindlegs extending beyond epiproct, anterior legs around same length as hindlegs, midlegs shorter. Coxae smooth. Profemur with distinct basal curvature. Femora and tibiae with five carinae with few sparse setae among them ( Fig. 36B–F ). Carinae of profemora and protibiae distinctly keeled, mid and hind femora and tibiae with slightly less prominent keels. Antero- and posteroventral carinae of femora with round apical prominence. Carinae of all femora, tibiae and tarsi bearing row of short stout setae, stouter and spiniform on tibiae. Along all basitarsi and near apex of meso- and metatibiae, ventral, antero- and posteroventral carinae bearing two to three rows of setae. Setae on ventral, antero- and posteroventral carinae of basitarsi stouter and spiniform. Ventral, antero-

and posteroventral carinae of tibiae ending as apical spiniform sessile projection parallel to tibiae, pointing towards tarsi ( Fig. 36E–F ). Pro- and metabasitarsi very elongate, significantly longer than respective following tarsomeres combined, mesobasitarsi as long as respective following tarsomeres combined ( Fig. 36B–D ). Basitarsi with hairy setae restricted to apical portion on ventro-lateral patches ( Fig. 36B, D ). Remaining tarsomeres with setae on ventrolateral patches in portions not covered by euplantulae. Tarsomeres I–III with dorsal apical projection. Arolium round and broad, bearing setae

dorsally. Pretarsal claws symmetrical, dorsally and outwardly bearing setae. Euplantulae well developed in all tarsomeres, composed of two symmetrical pads separated by median groove in tarsomeres I–IV, and of single flattened lobed pad in tarsomeres V. Euplantulae 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. 36B–D ).

ABDOMEN ( Figs 34 , 36A , 37 ). External surface as in thorax but bearing slightly more scattered short setae across entire length. Median segment shorter than metanotum, anteriorly with two parasagittal ovoid stains ( Fig. 36A ). Combined length of segments II–X as long as combined length of thorax

and median segment. Terga II–VIII and sterna II–VII bearing lateral carinae near lateral margins. All segments longer than wide. Segment II shorter than III, segment III of same length as VII and shorter than IV, segments IV–VI of same length, tergum VIII significantly shorter than preceding segment and slightly longer than IX and X, tergum X slightly longer than IX. Tergum II widening towards posterior, tergum III widest, slightly wider than II and IV, terga IV and V about same width, tergum VI wider than preceding segment, with lateral margins convex in dorsal view, tergum VII narrower than all preceding segments, slightly wider than VIII, terga VIII–X around same width or very slightly tapering ( Fig. 37A ). Tergum X somewhat conical in lateral view ( Fig. 37B ) and in dorsal view almost parallel-sided but at ¾ the length tapering towards posterior, posterior margin emarginate with round lateral margins ( Fig. 37A ). Epiproct very prominent, elongate, lingulate in dorsal view, dorsoventrally flattened, visible dorsally and laterally ( Fig. 37A–B ). Paraprocts elongate, posteriorly fairly acute, straight, bearing setae on posterior margin, laterally bearing cerci ( Fig. 37B–C ). Cerci extremely elongate, straight, basally narrower fitting paraprocts, pointing to posterior, gradually tapering and slightly shorter than terga IX and X combined ( Fig. 37A–C ), bearing four types of setae. Praeopercular organ very prominent, lobed,

elevated and folded backwards, forming a round dorsoventrally flattened, centrally depressed projection ( Fig. 37B–E ). Subgenital plate roundly lanceolate, short, almost reaching ⅓ of length of tergum X. Subgenital plate bearing two parasagittal carinae beginning on anterior margin and running half length of segment before becoming flatter and more setose, tapering towards posterior and totally covering gonapophyses and gonoplac ( Fig. 37B–C ). Gonapophyses and gonoplac flattened, dorsoventrally for gonapophyses VIII, lateroventrally for IX and laterally for gonoplac ( Fig. 37F–G ). Gonapophyses VIII longer than IX, gonapophysis VIII reaching ca 0.8× length of tergum IX, gonapophysis IX reaching ca 0.6 × length of tergum IX. Gonapophysis VIII linear and gradually tapering, IX conical, tapering towards posterior, with lateral margin concave and inner margin straight ( Fig. 37F–G ). Gonapophyses IX ventrally folded to fit within gonapophyses VIII. Gonangulum distinctly reduced, flat, not lobed. Gonoplac lorate, somewhat wide in lateral view, bearing setae and slightly longer than gonapophyses VIII ( Fig. 37F–G ).

Other females

MEASUREMENTS (in mm, N = 8). Body 92–113.5, head 5.0–5.8, antennae 48.0–58.0, pronotum 3.3–4.1, mesonotum 20.0–25.5, metanotum 7.5–9.9, median segment 4.9–7.3, abdomen (excluding median segment) 44.0–54.7, cercus 4.4–6.0, profemur 21.4–24.7, protibia 20.8–26.0, mesofemur 16.5–19.5, mesotibia 16.3–20.3, metafemur 20.0–23.6, metatibia 20.6–28.7.

VARIATION. Some few specimens may present shorter cerci due to regeneration from breaking in previous instars (e.g., Fig. 45C ). Some specimens may present more elongate gonapophyses IX, longer than gonoplac (e.g., Fig. 45D ).

COLOUR ( Figs 34 , 44–46A ). Body generally light to mid green, beige, brown, grey or dark, with or without irregular stains of different tones.

Male paratype (MZUSP V0602)

MEASUREMENTS (in mm). Body 57.6, head 2.9, antennae at least 48.8, pronotum 2.3, mesonotum 14.0, metanotum 5.6, median segment 4.1, abdomen (excluding median segment) 28.7, cercus 1.1, profemur 19.2, protibia 21.9, mesofemur 15.3, mesotibia 17.0, metafemur 18.8, metatibia 22.1.

COLOUR ( Figs 38–40 , similar to paratype in Fig. 34C ). Entirely light to mid brownish with irregular stains of different tones and scattered whitish to creamy granules.

HEAD ( Fig. 38 ). Similar to female except: eyes, scapus and pedicel proportionally larger, head widening from posterior margin towards anterior in dorsal view, vertex very gently convex near anterior margin above eyes, frontal suture more acute ( Fig. 38A ). Paraglossa wider, inner margin straight ( Fig. 38E ). Galea thinner, more elongate, slightly constricted before apical portion, galealobulus wider, less prominent ( Fig. 38C ). Labial palpal segments slightly less widened ( Fig. 38F ). Antennae reaching at least tergum VI, scapus ca 1.7× as long as wide ( Fig. 38A ).

THORAX ( Figs 38A–B, F , 39A , similar to paratype in Fig. 34C ). Similar to female except: significantly thinner and more elongate than that of female. Meso-, metathorax and median segment with straight lateral margins in dorsal view, not convex ( Fig. 39A ). Mesothorax ca 6.5 × as long as prothorax. Mesocoxopleurite larger, triangular. Metanotum 1.3× as long as median segment ( Fig. 39A ). Metafurca Y-shaped.

LEGS ( Fig. 39B–G , similar to paratype in Fig. 34C ). Similar to female except: legs thinner than those of female, basitarsi relatively longer ( Fig. 39B, D, G ). All setae generally longer than those of female ( Fig. 39B–F ). Setae over carinae of femora slightly sparser, over carinae of tibiae longer ( Fig. 39B, D, G ). Carinae of femora, tibiae and tarsi with single row of setae except for ventral carinae of basitarsi

with two to three rows. Apical spine in ventral carinae of tibiae less developed, discrete. All basitarsi longer than respective following tarsomeres combined ( Fig. 39B, D, G ). Ventrolateral hairy setae of tarsomeres denser. Euplantulae slightly shorter, on tarsomeres V rougher ( Fig. 39E ).

ABDOMEN ( Figs 39A , 40 , 41A–B , similar to paratype in Fig. 34C ). Thinner than that of female. Lateral carinae absent in median segment. Segments II–VII slightly constricted medially from anterior to posterior regions. Terga II, III and VI about same length, terga IV and V about same length and slightly longer than III, tergum VII shorter than VI, tergum VIII shorter than preceding segment and slightly longer than IX, terga IX and X about same length. Terga II–V about same width, tergum VI slightly narrower than preceding segment, tergum VII as wide as VI but slightly wider posteriorly, VIII wider than VII, slightly widening towards posterior, IX slightly narrower than VIII, tapering towards posterior, X narrower anteriorly, slightly widened medially with maximum width around as wide as maximum width of VIII and IX ( Fig. 40A ). Tergum X presenting lateral flat flexing expansions bent downwards from half length to posterior margin, with somewhat acute ventrolateral edges directed posteriad ( Fig. 40A–G ). Paraprocts totally covered by these expansions. Dorsal posterior margin of tergum X widely v-shaped emarginate ( Fig. 40A, F ). Ventral area of posterior dorsal margin of tergum X strongly sclerotized, black ( Fig. 40C ). Thorn pads composed of ca 28–35 strong, inwardly curved sclerotized conical teeth, variable in size, with very few in sclerotized area on ventral surface of posterior dorsal margin of tergum X, with teeth present also anteroventrally (near apex of vomer) and laterally at ventral surface of posterior margin of lateral expansions ( Fig. 40C ). Tergum VIII and sternum VIII separated, not fused ( Fig. 40B, E ). Sternum VIII partly fused with sternum IX, border between both sterna shown as asymmetric sulcus curving towards posterior on right side ( Fig. 40G ). Sternum IX reduced with anterior portion indistinct; only shown as poculum ( Fig. 40B, E ). Poculum reaching around ¾ length of tergum IX. Due to asymmetry of sulcus, poculum shorter at right side than left side ( Fig. 40B, E ). Poculum fairly short, round in lateral view, about as long as tall or slightly longer than tall at left side, round in ventral view, posterior margin medially with wide emarginate short curvature towards anterior ( Fig. 40G ). Cerci slightly shorter than tergum X, inwardly curved and with round apex ( Fig. 40A–F ). Epiproct small, hidden in dorsal and lateral views ( Fig. 40H ). Vomer elongate, dorsoventrally flattened, terminal hook elongate and strongly sclerotized, symmetrically bifid near posterior margin with round apices gently curving upwards ( Fig. 40C, G–H ).

GENITALIA ( Figs 41 , 43 , right side). Male genitalia or phallic organ attached ventrally at area corresponding to asymmetric sulcus between sterna VIII and IX ( Fig. 41A–B ) by large muscles connected to anterior portion of dorsal sclerite ( Fig 41C–D ). Phallic organ composed of dorsalmost sclerite (dorsal sclerite), wide lobe with fine granulation originating ventrally from dorsal sclerite (longitudinal lobe), and smaller and shorter lobe basally, slightly ventrally, at right side, the (right) basal lobe. Dorsal sclerite very elongate and penetrating inside body cavity ( Fig. 43 ). Anterior portion penetrating body, wider and medially constricted. Short truncate branch at right side pointing towards posterior, connecting dorsal sclerite to body wall at end of anterior portion ( Fig. 43C–D ). Posterior portion narrower, slender, slightly tapering towards apex and bent to left near apex ( Figs 41D , 43A–D ). Longitudinal lobe wide, anteriorly to left branching in round lobule pointing towards anterior, with densely finely granulated round lobule dorsally before apex, originating near dorsal sclerite. Longitudinal lobe posteriorly with several folding and smaller lobules, with dense fine granulations ( Fig. 41F ), larger and sparser circular granulations, ventrally with weakly sclerotized area near apex and more posteriorly with strong, projecting, small and round sclerite ( Fig. 41C–F ). Basal lobe short and apically bearing claw-like sclerite, strongly sclerotized, with few sparse granules and with wide base ( Fig. 41E ).

Other males

MEASUREMENTS (paratypes, range in mm, N = 6). Body 61–70.1, head 2.7–3.2, antennae 50.0–57.0, pronotum 2.0–2.6, mesonotum 15.8–17.4, metanotum 6.3–7.5, median segment 4.4–5.2, abdomen

(excluding median segment) 29.8–34.2, cercus 0.9–1.2, profemur 20.5–22.7, protibia 22.4–24.8, mesofemur 16.0–17.8, mesotibia 17.9–19.1, metafemur 20.0–23.2, metatibia 22.4–25.2.

VARIATION. Thorn pad at ventral area of posterior dorsal margin of tergum X bearing 2–10 spines.

COLOUR ( Figs 34A, C , 44–45 ). Entirely light to dark brownish, light greenish, grey or dark with irregular stains of different tones and with or without stains of beige, dark red, black with scattered whitish to creamy granules.

Egg ( Fig. 42 )

Measurements (in mm, N = 10): length 2.7–2.8, height 1.9–2.0, width 1.4–1.5. Relatively small,somewhat subrectangular in lateral view, slightly constricted at opercular collar, laterally compressed and ca 1.8 × as long as wide and 1.4× as long as tall ( Fig. 42A–D ). Capsule surface smooth but with finely punctuated appearance ( Fig. 42A–B ). Colour varying in shades of orangish or reddish brown with creamy lighter

band on dorsal surface extending in short posterior lateral patches around polar area. Micropylar plate very elongate, occupying large area of dorsal region, with round margins, almost parallel-sided with posterior portion gently widened ( Fig. 42C–D ). Micropylar plate with lighter round elevation delimiting inner flat region confluent with micropylar cup ( Fig. 42C–D ). Micropylar cup small, rounded and only slightly elevated. Median line short and of same colour and elevation as elevated margin of micropylar plate, almost reaching polar area, sometimes disjunct with micropylar plate ( Fig. 42C–D ). Opercular collar very slightly narrower (constricted) than rest of capsule, smooth, whitish with several minute and delicate long bristles surrounding edge ( Fig. 42E–F ). Operculum elliptical with marginally radial elevations with amber texture, irregular and less elevated at centre with some elevations fused centrally ( Fig. 42A–F ).

Distribution ( Fig. 52 )

Known only from the type locality, in the Serra do Cipó District, Santana do Riacho, Minas Gerais, Brazil. Serra do Cipó is part of the Serra do Espinhaço (Espinhaço mountain range), which is a wide and long mountain range in Brazil typically dominated by campos rupestres (“rupestrian grasslands”), a grassland to savannic rocky formation ( Fig. 53G–H ). All individuals of the species were found in campos rupestres vegetation near a road running through the Serra do Cipó ( Fig. 53G–H ). Certainly, the species also occurs in a nearby protected area continuous with the type locality, also in the Serra do Cipó formation, the Parque Nacional da Serra do Cipó, although it has not yet been recorded for the Park.

Biology

At the type locality at Serra do Cipó specimens were found in the rainy season at several stages from early nymphs to mating adult pairs. In two nights of searching, more than 20 specimens were seen over a few kilometres. Specimens were observed feeding on native locust berries ( Byrsonima Rich ex. Kunth, Malpighiaceae ) ( Fig. 44 ) and on guava trees alongside the road (native to Brazil but not to that particular environment; it is widely dispersed in urbanized areas). In captivity, they also fed on Brazilian cherry. Similar to Arumatia dubia gen. et comb. nov. both nymphs and adults are agitated when handled but in a somewhat lessened manner. Reared specimens were not observed to hide resting along a branch, but usually with mid and hindlegs keeping the body elevated. Eggs laid by adult females since collecting took 90–150 days to hatch but most of 10 controlled eggs hatched within 105 days. The first nymphs to hatch were adult males or pre-subadult females when totalling 60–73 days of development and females became adults at around 90–105 days of development.

In one of the mating attempts observed under captivity, the male repeatedly touched the female with his antennae while climbing on her, but the pair parted ways shortly after. Another two mating attempts resulted in a quick tapping of the antennae on the female while climbing on her and either only the male or the pair started to frenetically move their legs. The male quickly jumped on top of the female facing

the same direction with all tarsi touching her thorax. Then, while the female was shaking, the male positioned his abdomen beneath the abdomen of the female by the left side, tapped with the apex of his abdomen and then significantly bent his terga VIII to X anteriorly and slid the apex of his abdomen up and down the ventral surface of the abdomen of the female until the posterior margin of his tergum X attached to the female’s praeopercular organ ( Fig. 45A ). Then, the male pushed his abdomen backwards so the poculum contacted the female’s subgenital plate and copulation started ( Fig. 45A–B ). At this moment, the male’s abdomen curved upwards so the ventral side of the apex of his abdomen contacted the ventro-lateral area of the tip of the female’s abdomen ( Fig. 45B ). The poculum faced the right side of the female’s subgenital plate and attached that way. Later on, the male was seen to hang attached to the female only by the abdomen and eventually mounted on the top of the female again but with the abdomen now curving to the right side so that his abdomen curved downwards instead of upwards ( Fig. 45C ). They remained in copulation for around 50 hours. Males seen already in copulation with females were always positioned by the right side ( Figs 34A , 44 , 45 ).

Males attach to the female in three ways: by grasping the female’s subgenital plate with his curved cerci ( Fig. 45D–E ), by inserting his vomer underneath the praeopercular organ (hidden from view in the pictures) and by folding the lateral expansions of its tergum X, grasping the female praeopercular organ ( Fig. 45D–E ).

Figures

Fig. 34. Live habitus of Arumatia motenata Ghirotto gen. et sp. nov. from Serra do Cipó, Minas Gerais, Brazil. A. Mating pair. B. Adult ♀, holotype (MZUSP V0603). C. Adult ♂. Scale bars: 1 mm.

Fig. 35. Arumatia motenata Ghirotto gen. et sp. nov., holotype, ♀ (MZUSP V0603), head and prothorax morphology. A. Head and prothorax in dorsal view. B. Head and prothorax in lateral view. C. Head and prothorax in ventral view. Abbreviations: Ant = antennomere; BsI-II =pro- and mesothoracic basisterna; Cly =clypeus; Csc =cervical sclerite; Cx =coxa; Cxp =coxopleurite; Eye=compound eye; Fc =frontal convexity; Fs=prothoracic furcasternite; Gu =gula; Lb =labrum; Md=mandible; Msn =mesonotum; Par =paranota; Pd =pedicellus; Plb =labial palpus; Pmx=maxillary palpus; Prn =pronotum; Sc =scapus; Sg = subgena; Sme =submentum. Scale bars: 1 mm.

Fig. 36. Arumatia motenata Ghirotto gen. et sp. nov., holotype, ♀ (MZUSP V0603), thorax and leg morphology. A. Metathorax and median segment in dorsal view, arrows indicate the length of each segment. B. Left fore tarsus, ventro-prolateral view. C. Left mid tarsus, ventral view. D. Right hind tarsus, ventral view. E. Detail of right mid tibia and tarsus (mirrored), prolateral view. F. Detail of right hind tibia and tarsus, ventral view, arrow showing apical sessile spine. Abbreviations: Ar =arolium; Cda =dorsal anterior carina; Cl =claw; Cva =ventral anterior carina; Cve= ventral carina; Cvp; ventral posterior carina; Epm =epimeron; Eps =episternum; FTa = fore tarsomere; HTa =hind tarsomere; Mds =median segment; Msn =mesonotum; MTa=mid tarsomere; Mtn =metanotum; Tg =tergum; Tib= tibia. Scale bars: A =1 mm; B–F =0.5 mm.

Fig. 37. Arumatia motenata Ghirotto gen. et sp. nov., holotype, ♀ (MZUSP V0603), terminalia morphology. A. Dorsal view. B. Lateral view. C. Ventral view. D–E. Detail of praeopercular organ, ventral (D) and lateral view (E). F–G. Detail of external genitalia, ventral (F) and lateral view (G). Abbreviations: Ce= cercus; Ep=epiproct; Gap =gonapophysis; Gpl=gonoplac; Pre =praeopercular organ; Prp= paraproct; Sp= subgenital plate; St =sternum; Tg=tergum. Scale bars: A–C =1 mm; D–G =0.5 mm.

Fig. 38. Arumatia motenata Ghirotto gen. et sp. nov., paratype, ♂ (MZUSP V0602), head and prothorax morphology. A. Head and prothorax in dorsal view.B. Head and prothorax in lateral view.C. Left maxilla in dorsal (internal) view. D. Detail of labrum, anterior view. E. Detail of paraglossa and glossa, anteroventral view. F. Head and prothorax in ventral view. Abbreviations: Ant=antennomere; BsI-II =pro- and mesothoracic basisterna; Cly = clypeus; Csc =cervical sclerite; Eye= compound eye; Fc =frontal convexity; Fs=prothoracic furcasternite; Ga =galea; Galo= galealobulus; Gl =glossa; Gra =granules; Gu =gula; Lac =lacinia; Lase =lacinial setae; Lb=labrum; Md= mandible; Msn= mesonotum; Mtr =microtrichia; Pd =pedicellus; Pgl =paraglossa; Plb =labial palpus; Pmx= maxillary palpus; Prn =pronotum; Sc = scapus; Sg =subgena; Sme =submentum. Scale bars: A–B, D–F = 0.5 mm; C= 0.2 mm.

Fig. 39. Arumatia motenata Ghirotto gen. et sp. nov., paratype, ♂ (MZUSP V0602), thorax and leg morphology.A. Metathoraxand mediansegmentindorsalview, arrowsindicatethelength ofeachsegment. B. Left fore tarsus, ventral view. C. Detail of left fore tibia and tarsus. D. Right mid tarsus, retrolateral view. E. Detail of right mid tarsus, ventral view. F. Detail of right mid tibia and tarsus, ventral view, arrow showing apical sessile spine. G. Right hind tarsus, retrolateral view. Abbreviations: Cva =ventral anterior carina; Cve =ventral carina; Cvp; ventral posterior carina; Epm =epimeron; Eps =episternum; FTa =fore tarsomere; HTa =hind tarsomere; Mds= median segment; Msn =mesonotum; MTa =mid tarsomere; Mtn =metanotum; Tg=tergum; Tib=tibia. Scale bars: 0.5 mm.

Fig. 40. Arumatia motenata Ghirotto gen. et sp. nov., paratype, ♂ (MZUSP V0602), terminalia morphology. A. Dorsal view. B. Left side lateral view. C. Posterior view, detail of tergum X. D. Left side latero-dorsal view. E. Right side lateral view. F. Posterodorsal view, detail of tergum X. G. Ventral view. H. Detail of vomer, ventral view. Abbreviations: Ce =cercus; Ep= epiproct; Pha =phallic organ; Po=poculum; Prp= paraproct; St = sternum; Tg=tergum; Tp=thorn pads; Vo =vomer. Scale bars: 0.5 mm.

Fig. 41. Arumatia motenata Ghirotto gen. et sp. nov., ♂♂, genital morphology. A–D. Paratype (MZUSP V0620). E–F. Paratype (MZUSP V0621). A–B. Terminalia, ventral view. A. Poculum pushed anteriorly showing the phallic organ. B. Phallic organ tilted, arrows showing attachment area corresponding to the sulcus between sterna VIII and IX. C–D. Dissected untreated phallic organ, ventral (C) and dorsal view (D), grey line highlights area of attachment to the body. E. Detail of phallic organ treated with KOH, ventral view.F. Detail of granulations on the longitudinal lobe of phallic organ treated with KOH, lateral view. Abbreviations: BL =basal lobe; DS = dorsal sclerite; LL=longitudinal lobe; Pha =phallic organ; Po =poculum; Tg=tergum. Scale bars: A–E=0.5 mm; F =0.1 mm.

Fig. 42. Arumatia motenata Ghirotto gen. et sp. nov., egg morphology. A–B. Lateral view. C–D. Dorsal view. E–F. Detail of operculum, anterior view, arrowheads pointing to bristles on opercular collar (E). Abbreviations: Cap =capitulum; Mcup =micropylar cup; Mip =micropylar plate; MLi =median line; Opc = operculum. Scale bars: 0.5 mm.

Fig. 43. Male genitalia of Arumatia aramatia Ghirotto gen. et sp. nov. and Arumatia motenata Ghirotto gen. et sp. nov., treated with KOH.A, C, E, G. Treated genitalia of A. arumatia (left side) andA. motenata (right side) in different views. B, D, F, H. Same pictures as respective preceding, structures coloured for visualisation: red =dorsal sclerite; blue=longitudinal lobe; green =basal lobe; yellow= internal sclerotization; white line = area of attachment to the body; inset (F) =detail of internal sclerotization, internal view. All pictures to the same scale. Scale bar: 0.5 mm.

Fig. 44. Arumatia motenata Ghirotto gen. et sp. nov., live habitus in situ at Serra do Cipó, Minas Gerais, Brazil. Adult pair, at night, on locustberry plant (Byrsonima sp.). Note female feeding on the plant and male hanging attached to the female by the terminalia. Image not to scale.

Fig. 45. Arumatia motenata Ghirotto gen. et sp. nov., sexual behaviour and mating attachmentmechanism, from Serra do Cipó, Minas Gerais, Brazil. A–C. Sequential copulatory behaviour. D–E. Attachment of male and female structures. A. Male (dark) on top of female (green) extends his terga VIII-X anteriorly and touches the ventral surface of the abdomen of the female trying to attach. B. Male attached to the female by the left side and inserting the phallic organ; note abdomen of male curved upwards. C. Male and female in copulation, male hanging from femal; note abdomen of male extended.D–E. Male attached to the praeopercular organ of the female with the posterior margin of its tergum X (arrow), cerci pressing the subgenital plate of the female, lateral (D) and ventral view (E). Images not to scale.

Fig. 46. Comparison of live adult females. A. Arumatia motenata Ghirotto gen. et sp. nov. B. Arumatia dubia gen. et comb. nov. Scale bar: 10 mm.

Fig. 53. Habitats of Arumatia Ghirotto gen. nov. in the Cerrado of Brazil. A–D. Habitat of Arumatia dubia gen. et comb. nov., seasonal forest at Assis, São Paulo (A), seasonal forest at Echaporã, São Paulo (B), savannah formation at Delfinópolis, Minas Gerais (C–D). E. Type locality of Arumatia crassicercata Ghirotto, Crispino & Engelking gen. et sp. nov., gallery forest at Chapada dos Veadeiros plateau, Alto Paraíso de Goiás, Goiás. F. Type locality of Arumatia aramatia Ghirotto gen. et sp. nov., savannah formation in Luzimangues district, Porto Nacional, Tocantins (photo courtesy of Dante Pavan). G–H. Type locality of Arumatia motenata Ghirotto gen. et sp. nov., rocky savannah formations (campos rupestres) at Serra do Cipó district, Santana do Riacho, Minas Gerais. I. Type locality of Arumatia diamante Ghirotto gen. et sp. nov., dense rocky savannah formations (campos rupestres) at Catolés, Abaíra, Bahia (photo courtesy of Pedro H. Martins).

Fig. 52. Distribution map of species of Arumatia Ghirotto gen. nov. in central South America. Brazil is further divided into states which are shown in a thinner line. Note that grey dotted circles represent photographic records of Arumatia dubia gen. et comb. nov. from the iNaturalist (iNat) online platform, the black shield represents an unidentified nymph of Arumatia gen. nov. from MZUSP collection and the grey dotted shields represent photographic records of a potentially new species from iNaturalist.