Xerosoma nannospinus, Engelking & Ghirotto & Crispino & Büscher & Heleodoro & Neves & Bispo, 2023

Engelking, Phillip Watzke, Ghirotto, Victor Morais, Crispino, Edgar Blois, Büscher, Thies H., Heleodoro, Raphael Aquino, Neves, Pedro Alvaro Barbosa Aguiar & Bispo, Pitágoras da Conceição, 2023, Fig. 23. Saiva formosana Kato, 1929, male genitalia. A in Lanternflies (Hemiptera: Fulgoridae) of Taiwan., Zoological Studies 62 (31), pp. 1-59 : 17-40

Xerosoma canaliculatum Serville 1831 ( Figs. 16–48 )

Xerosoma canaliculatum Serville 1831 (61); Gray 1835 (27); Serville 1838 [1839] (3) pl. 6, Fig. 3 ; Burmeister 1840 (40); Blanchard 1840 (18); Westwood 1859 (104); Stål 1875a (100); Kirby 1904 (416); Redtenbacher 1906 (9); Klante 1969 (3); Zompro 2004b (93); Otte and Brock 2005 (342); Araujo and Garraffoni 2012 (236); Delfosse et al. 2019 (193).

Dinelytron neptuna Kaup 1871 (41); Kaup 1871 (23); Kirby 1904 (414) [ Prexaspes neptunus ]; Redtenbacher 1906 (131) [ Harupuna neptunus ]; Zompro 2001 (138), Fig. 13 [Harupuna neptuna ]; Zompro 2004b (93) [ X. canaliculatum ].

Xerosoma senticosa Stål 1875a (99); Kirby 1904 (416); Redtenbacher 1906 (144), pl. 5:10 [ X. senticosum ]; Brock 1998 (57); Zompro 2004a (112) Otte and Brock 2005 (343); Araujo and Garraffoni 2012 (236). syn. nov.

Examined material: 1ñ (MNHN-EO-PHAS497): “ Brésil, 135” (holotype, examined by photo); 1ò ( MNHN) “ Brasilien ” (examined by illustration); 1ñ ( MNW 239): Brazil, Bahia, “ Xerosoma senticosa, Coll. Br. V. W., Coll. Sommer ” (holotype of Xerosoma senticosum , examined). 1ò ( MNW 7654): Brazil, Bahia, “Coll. Br. V. W., Coll. Sommer” (same coll. and locality as the holotype of Xerosoma senticosa , examined); 1ñ ( SMFD (examined by photo)): “ Brasilien ”; 1 egg ( HLDL (examined by illustration)); 1ò ( MZUSP 799): Brasil, Bahia, Parque Nacional do Pau Brasil, 16°28'24"S, 39°16'20"W, 18–19. XI.2016, P. I. Chiquetto‐Machado, J. P. Constantini, N. C. C. P. Barbosa. 2ò 1ñ ( MZUSP 751, 764, 845): Brasil, Espírito Santo, Reserva Biológica Córrego Grande, 18°14'32"S, 39°40'00"W, 20–21.XI.2016, P.

I. Chiquetto‐Machado, J. P. Constantini, N. C. C. P. Barbosa. 1ò 1ñ ( MZUSP 489, 490): Brasil, Minas Gerais, Parque Nacional do Caparaó, Núcleo Alto Caparaó, 20°25'10"S, 41°50'53"W, 30.xi-1.xii.2015, P. I. Chiquetto‐Machado, A. Z. Ramin, J. P. Constantini. 1ò ( MZUSP 851) Brasil, Rio de Janeiro, Parque Nacional de Itatiaia, 22°25'35"S, 44°37'05"W, 31.x-2.xi.2016, P. I. Chiquetto-Machado. 2ò ( MZUSP 1203, 1204): Brasil, Espírito Santo, Aracruz, Itaparica, 9–11.i.2019, E. B. Crispino, E. Travassos Júnior, P. W. Engelking, I. M. Cunha, S. Harumi. 5ò, 5ñ, eggs, nymphs ( MZUSP): raised in captivity by V. M. Ghirotto, origin: Brasil, Espírito Santo, Aracruz. 1ñ (last instar nymph) ( MZUSP 389): Brasil, Espírito Santo, Reserva Biológica de Sooretama, 19°03'15"S, 40°08'48"W, 24– 26.xi.2014, P. I. C. Machado, T. F. Carrijo, R. G. Santos; 2ò 1ñ ( INPA) Brasil, Espírito Santo, Reserva Vale, R. Heleodoro.

Type condition: ( Fig. 17 ). The type ñ is very well preserved. Ventrally, it is possible to see that the specimen had its abdomen filled with cotton through an incision, which made it impossible to see the preopercular organ. The wings are opened.

Remarks: Xerosoma senticosum (Stål, 1875) is solely represented by the female holotype, deposited in NMW ( Fig. 18 ). However, a male also from NMW from the same locality and collector label, highly likely from the same population as the X. senticosum holotype was also analyzed ( Fig. 19 ). The only observable difference of this material from most Xerosoma canaliculatum is the absence of the black longitudinal line on the head of the female. As this line appeared poorly marked in some X. canaliculatum female specimens, in addition to all other morphological characteristics matching those of X. canaliculatum , we synonymized Xerosoma senticosum (Stål, 1875) syn. nov. under Xerosoma canaliculatum . The etymology of X. senticosum was not given by Stål (1875a), however, the epithet likely comes from the Latin words “ sentis ” and “ cosa ”, which mean thorns and hip, respectively, probably referring to the “lateral projections on the last terga”, as mentioned by Stål (1875a). Based on current knowledge and considering that no voucher material was specified, we disregard Bradley and Galil’s (1977) record of X. canaliculatum from Costa Rica.

Diagnosis: Xerosoma canaliculatum differs from X. michaelis by the fully developed hindwings and the reduced spination on femora I –II. It differs from X. nannospinus sp. nov. by the flattened vertex, larger eyes, stronger spines, longer tegmina, longer wings, lateral terga projections more pronounced, presence of three apical projections on tergum X in ñ and broad projections on tergum X in ò, compressed gonapophysis in ñ, longer cerci in ò, flattened vomer in ò, longer eggs, and male genitalia with smaller dorsal sclerite basal projection, as the more schlerotized apical of dorsal sclerite.

Comments on the etymology: The etymology as given by Serville (1831) comes from the Latin word “ canaliculatum ”, which means small canal (canaliculus), referring to the marked longitudinal black line on the head.

Redescription (based on type and additional material).

Female: Total body size of 57–70 mm, winged ( Figs. 20–24 ). Measurements are given in table 4.

Head: ( Fig. 20 ). Elongate, sub-rectangular in dorsal view, and slightly longer than wide; length and width approximately the same as the prothorax. Vertex slightly convex, with a pronounced longitudinal blackened line (canaliculus), less marked in some few specimens, appearing only as a stain in northern populations or depressed into a sulcus on the posterior half in southern populations. Head with a pronounced pair of carinae basally and 8–10 small granules dorsolaterally. Eyes large, pronounced, slightly elongated, about a third the length of the head. Labial and maxillary palps elongated; when extended, the labial palp slightly exceeds the head, while the maxillary palp entirely exceeds the head. Clypeus convex, with two paramedian deep triangular depressions. Lacinia with three distal teeth; mesal edge bearing a restricted bundle of strong setae. Galea broad, spatulate, bearing rows of setae along the outer lateral and the anterior edge; inner laterally and slightly apical with two dense patches of elongate, stout microtrichia, ventralmost patch long, distributed along the edge of the galea, dorsalmost patch smaller, elliptical; between this patch and the lateral rows of setae on both surfaces, several distinct circular granules around the same diameter of the base of nearby setae. Galealobulus well developed, conical, but only gradually tapering, very elongated, almost as long as the galea. Left mandible with two cutting edges, dorsal undulate and blunt, ventral straight and slightly sharper; right mandible with dorsal cutting edge undulate and blunt, ventral blunter, almost entirely straight, except for a convexity basally; both mandibles with a dense row of large setae at the base of the dorsal margin; mesal surfaces smooth without protuberances. Antennae filiform, reaching tergum III; composed of about 30 antennomeres; covered with fine bristles; scapus compressed dorsoventrally, 1.3× longer than wide; pedicel cylindrical, slightly shorter than scapus; first flagellomere (third antennomere) about 2.2× longer than the pedicel. Antennal bump distinct, present on the 14th antennomere.

Thorax: (Fig. 21 A – C). Pro- and mesothorax subquadrate in cross section, with somewhat flat dorsal and lateral surfaces; ventrally bearing small, short setae. Prothorax about 1.7× longer than wide, slightly convex dorsally, with a central transversal groove; anterolateral margins slightly concave outlining the opening of the prothoracic glands. Pronotum bearing a pair of longitudinal, dorsolateral carinae, running from the gland openings and extending almost to the posterior margin, and bearing a pair of large, anteriorly pointed dorsal spines near the posterior margin, accompanied by smaller spines at their enlarged base. Small granules scattered on the dorsal surface and along the dorsolateral carinae of the pronotum. Profurcasternite slightly wider than long. Mesothorax in dorsal view slightly widening towards the posterior region. Mesonotum 1.8× longer than the pronotum, surface slightly rough, with a pair of longitudinal, dorsolateral carinae extending along the anterior two thirds, with a sparse row of 2 to 3 stout, short spines each. Mesepisternum with longitudinal carinae along the ventral margin; medially bearing large and short, stout rounded granules. Mesobasisterum with a longitudinal median carina. Metathorax and median segment smooth; combined length of both structures approximately 1.4× longer than the mesothorax. Metanotum parallel-sided, slightly longer than the mesonotum and 4–5× longer than the median segment. Metepisternum with a thick longitudinal carina along the ventral margin.

Wings: (Fig. 21D, E). Tegmina distinctly convex, 2.2× longer than wide, reaching tergum I, projecting dorsolaterally into large shoulder pads slightly pointing forward, venation pattern latticed. Costal area with Sc, R, Rs1, Rs2, and Ma veins reaching apical margin; Mp shorter, directed to the Cu margin. Anal area very reduced, A 1 short, following the Cu margin, spanning the first third of the length of the tegmina. Hindwings fully developed, reaching or exceeding tergum V. Costal area with Sc vein ending in C margin, spanning around 75% the total length of the costal area; veins R 1, Rs1, Rs2, Ma, and MP reaching the apical Cu margin. Anal area tesselate and fan-shaped, with 7 Aa veins and 8 Pa veins gradually decreasing in length.

Legs: ( Fig. 22 ). Femora and tibiae rectangular to trapezoidal in cross section, all carinae pronounced bearing one to three rows of short setae. Profemora with basal curvature and laterally compressed, bearing some small acuminate lobes in the ventral-posterior carinae. Area apicalis well developed, densely covered by setae. Tibiae slightly longer than the corresponding femora and 2.5× longer than the corresponding tarsi. Pro- and metabasitarsus slightly longer than the following two corresponding tarsomeres combined; mesobasitarsus relatively shorter, around the same length of the following two tarsomeres combined. Euplantulae strongly developed at the distal ends of tarsomeres 1–4, hemispherical and bilobed. Tarsomere 5 bearing an elongate accessory euplantula. Euplantulae covered with small conical protrusions (nubs). Praetarsus equipped with a large arolium and two curved claws without pectination.

Abdomen: ( Figs. 23 and 24 ). Approximately 1.2× longer than the combined length of the head, thorax and median segment. Dorsoventrally flattened, mainly towards the posterior margin, ventrally bearing short dark setae. Terga VIII–IX with a smooth medial hump near the posterior margin and X with a short and discrete carina near the posterior margin. Posterior margin of tergum X crenate. Sternum VIII (subgenital plate) barely reaching the median length of IX, slightly longer than wide. Epiprocts small, triangular, hidden under tergum X in dorsal view. Paraprocts wide, both briefly separated from each other near the posterior margin. Cerci clavate, apex conical. Praeopercular organ wide, short, shown as a wide elevation preceded by a slight, thin transversal depression and followed by a glabrous thicker depression continuous with the anterior margin. Gonapophyses VIII wide, conical in ventral view, not totally covered by the subgenital plate; surface similar to that of surrounding body areas and similarly bearing setae; gonapophyses IX fused with gonoplac for most of its length, then short, slightly thin, conical, shiny, bearing few setae apically, hidden from ventral view atop gonapophyses VIII and gonoplac. Gonoplac well developed, dorsoventrally flattened, surface similar to that of gonapophyses VIII, longer than both gonapophyses, apically elliptical. Gonangulum absent.

Variations: Some individuals presented a slightly longer and more acute subgenital plate, especially those from Espírito Santo and Bahia states (northern known range of the species) in comparison with those of São Paulo state in the southern range. The cerci also vary from more to less pointed. The holotype bears a slightly rounder cerci ( Fig. 17D ). The hindwings can be more or less marked with tessellate black stripes and vary slightly in length.

Color: Body mostly stained with irregular shades of brown, complemented by black and white spots and may have green, yellowish or reddish stains. Posterior region of the head, prothorax and mesothorax light brown or yellowish to creamish, lighter than the surrounding areas. Head may have a dorsal longitudinal black line; lateral region of the head, prothorax and mesothorax bearing a dark brown band. Metathorax dorsally black. Abdomen dorsally darker in the first 5 segments covered by the wings and laterally brownish; terga VII–VIII with lateral black spots. Antennae similar in color as the rest of the body, becoming lighter from the 7th antennomere. Costal area of tegmina irregularly brownish with lighter and darker spots and anal region with black spots; ventrally reddish basally. Hindwings with costal area similar to the tegmina and including a red stain basally; anal area with light, translucent cells centrally and near the costal area, area near veins brownish, veins dark brown to blackish. Legs in different shades of brown similar to the body, but with black spots mainly on the anterior femora.

Eggs: ( Figs. 25 , 26 ). As for the genus. Elongater than those of X. nannospinus . Micropylar plate rounded to rectangular or triangular. Capsule with medium size hairy setae, smaller than those of X. nannospinus .

Male: Body size of 53–67 mm, thinner, less rough, and less granulated than the female; fully winged ( Figs. 27–31 ). Measurements are given in table 4.

Head: ( Fig. 27 ). Similar to that of the female, except: with fewer and less pronounced granulations; eyes larger. Ocelli reduced; antennae longer, almost as long as the body; both maxillary and labial palps

D E

Fig. 21. Thorax and wings of ñ Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: dorsal view. B: lateral view. C: ventral view. D: open wings in dorsal view. E: schematic drawing of wing venation. Abbreviations: C, costa vein; Cu cubitus vein; Sc, subcosta vein; R 1, radius 1 vein; Rs, radial sector vein; MA, media anterior vein; Mds, median segment; MP, media posterior vein; Mst, mesonotum; Mtt, metanotum; Pt, pronotum; Tg, tegmen; Wg, wing; 1 A, first anal vein; 1 AA –7 AA, first to seventh anterior anal veins; 1 PA –8 PA, first to fifth posterior anal veins. Scale bars = 1 cm.

more elongated; fewer setae on the mesal surface of the lacinia; galea shorter and thicker basally, with the area bearing the microtrichia projected ( Fig. 28E ); microtrichia of the galea similar, but with shorter patches than those of the female; galealobulus shorter and thicker. Antennal bump on the 13th antennomere.

Thorax: (Fig. 29). Similar to that of the female, except: thinner and more elongated; surface less

A B

granulated; ventral setae thinner, longer, and lighter in color; pair of spines on the pronotum thinner and bearing only few granules basally; profurcasternite longer than wide.

Wings: (Fig. 29). Similar to that of females, except: slightly elongated. Also presenting Rs subdivided into Rs1 and Rs2, reaching the tegmina apical costal area.

Legs: ( Fig. 30 ). Similar to those of the female,

E F G

except: slightly longer and more slender, profemora only gently compressed laterally with a weaker basal curvature. Tibiae slightly longer than the corresponding femora and 2.5× longer than the corresponding tarsi. Pro- and metabasitarsus slightly shorter than the respective following four tarsomeres combined; about the same length as the respective following three

A tarsomeres combined. Euplantulae strongly developed at the distal ends of tarsomeres 1–4, hemispherical and bilobed. Tarsomere 5 bearing an elongated accessory euplantula. Euplantulae covered with small conical protrusions (nubs). Praetarsus equipped with a large arolium and two curved claws without pectination.

Abdomen: ( Fig. 31 ). Dorsally glabrous under

D

wings and bearing keels in the posterior, exposed region, but in comparison to the female ventral setae are thinner and longer, light in color. Similar to that of the female, except: approximately 1.5× longer than the combined length of the head, thorax, and median segment; terga II– VI equal in length, parallel-sided, about 1.8× longer than wide; VII slightly shorter than VI; VII–X keeled, VII slightly longer than wide; VIII–IX slightly wider than long, convex; X slightly wider than long, slightly tapering from the mid-length to the posterior margin. Terga VIII–IX bearing a small medial round hump near the posterior margin pointing backwards. Cerci elongated, slightly flattened laterally from middle to apex. Thorn pads along the posterior ventral margin of tergum X with few short, strong dark thorns. Anterior portion of sternum IX short, not surpassing the posterior end of VIII; poculum (posterior portion of sternum IX) dorsoventrally flattened, posterior margin truncated.

Color: Similar to that of females, except: more frequently brown; fewer individuals greenish; mesonotum with a more contrasting wide, longitudinal beige band than that of the female.

Genitalia (phallic organ): ( Figs. 32–33 ). Dorsal sclerite elongated, bending towards the left; posterior margin from its wide attachment area connected to the dorsal wall of the body and continuing to the left, slightly curved towards the right side ( Fig. 33A ). Both the membrane covering the right side of the wall of the genitalia and the dorsal sclerite surface itself are dorsally covered with circular sclerotized granules. Longitudinal lobe with around three large folding parts ( Fig. 32B ). Great area of the longitudinal lobe in the central and right sides densely covered with minute, spiny conical setae ( Fig. 32L ), conferring a yellow appearance to the membrane in a cleared genitalia ( Fig. 32A–J ). Right side of the lobe basally bearing around a dozen large, straight, long bristles varying in size (similar to those in Fig. 32K ). Basal (left) lobe short and folding within the longitudinal lobe in untreated genitalia and before dissection ( Fig. 32M–P ), bearing around ten large, straight, long bristles (similar to those of the longitudinal lobe) of different sizes at the ventro-lateral right edge ( Fig. 32K ).

Nymphs: ( Fig. 34 ). First instar nymphs with proportionately large head, smooth body with only very few granules and scattered tiny dark bristles; body green with scattered small black spots and reddish eyes. Nymphs throughout ontogeny gain the adults usual brownish color and the rough and granulate surface with acute spines. Females take 6 instars to become adults, while males take 5 instars ( Fig. 35 ). Nymphs exhibiting sexual dimorphism since the first instar, males bearing a shortened sternum IX (poculum) and females with a shortened sternum VIII (subgenital plate) and budded gonapophyses and gonoplac, similar to Canuleius similis Redtenbacher 1906 ( Ghirotto 2021). Gonapophysis IX with the same size as other valves until subadult stage. First instar nymphs with four tarsomeres and inconspicuous euplantulae ( Fig. 36 ). Euplantulae of second instar nymphs larger and with four tarsomeres, but basitarsus showing a faint fissure. Nymphs from the third instar onwards with five tarsomeres. Euplantulae of all instars covered with a nubby microsculpture ( Fig. 36 ).

Natural history and biological aspects of Xerosoma canaliculatum

Geographic distribution: Xerosoma canaliculatum is recorded from five Brazilian states: Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro, and São Paulo.

Habitat: Populations of Xerosoma canaliculatum a r e r e c o r d e d f r o m a l t i t u d e s r a n g i n g f r o m 1 4 meters above sea level (Bertioga, São Paulo state) to 1295 meters (Parque Nacional do Caparaó, Minas Gerais state). They inhabit distinct kinds of phytophysiognomies in the Atlantic Forest, such as Restingas Altas (coastal sandbank forests) ( Fig. 37 ), lowland ombrophilous forests, and submontane ombrophilous forests. In Bertioga, specimens were sighted from near the ground resting in the herbaceous stratum to the forest understory at 6 meters above the ground.

Ecdysis: ( Fig. 38 ). Reared individuals remained

D E

Fig. 29. Thorax and wings of ò Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: dorsal view. B: lateral view. C: ventral view. D: open wings in dorsal view. E: schematic drawing of wing venation. Abbreviations: C, costa vein; Cu cubitus vein; Sc, subcosta vein; R 1, radius 1 vein; Rs, radial sector vein; MA, media anterior vein; Mds, median segment vein; MP, media posterior vein; Mst, mesonotum; Mtt, metanotum; Pt, pronotum; Tg, tegmen; Wg, wing; 1 A, first anal vein; 1 AA –7 AA, first to seventh anterior anal veins; 1 PA –8 PA, first to fifth posterior anal veins. Scale bars = 1 cm.

about 14 to 18 days in each instar. The duration of the ecdysis processes were approximately 30 minutes. During this process, it was possible for individuals to recover lost appendages ( Fig. 39 ).

Seasonality and longevity: This species appears to be perennial since individuals in all instars were observed during samplings in the rainy period. The lifespan of individuals in captivity and in a controlled

A B

environment ranged between 6 and 8 months, with around 3 to 4 months in the nymphal stage and 3 to 4 months in the adult stage. Egg incubation time was approximately 4 to 6 months.

Movement: This species is strictly nocturnal, remaining immobile during the day unless disturbed. Some individuals observed feeding or foraging at night were reencountered during daylight resting in situ ( Fig.

E F G

40) more than 2 meters away from the feeding site ( Fig. 40A ). Both in nature and in captivity, they rested during the day in different positions and on several parts of the plants, such as leaves, branches, trunks, dead leaves, or other plant material available ( Figs. 40 , 41 ). The most common resting position was with the first pair of legs extended concealing the head, the second pair usually facing forward, flexed at an angle of 45°, and

A the third pair extended very close to the abdomen. ( Figs. 40 , 44A ). During nocturnal activity most nymphs and adults hanged from the underside of leaves, while some adults, mainly males, were observed on the upper side of leaves, with some males eventually flying away.

Camouflage: Xerosoma are visually very similar to dry branches, exhibiting different patterns of colors to match their surroundings ( Fig. 42A–F ). During the

D

day, the insects tend to hide next to thicker branches or bark pieces when available, while at night individuals forage. Active individuals frequently stop moving when exposed to a flashlight. Specimens also vary in color according to the time of day or brightness of the environment, matching tones with their surroundings and perfecting their crypsis ( Fig. 42G, H ).

Secondary defense mechanisms: Nymphs mostly exhibit thanatosis when slightly disturbed, falling immobile to the ground with legs held close to the body. When intensely disturbed, i.e., grasped, nymphs always expel an irritating substance from their prothoracic glands. The substance is expelled in a microjet and quickly volatilizes ( Fig. 43D ). The smell is minty, sour in taste, and causes mild irritation in the airways when inhaled by a person, resulting in coughing or sneezing. Early-stage nymphs frequently autotomize legs when disturbed, probably to minimize damage from an attack ( Bedford 1978).

Adults bear red spots at the base of their wings, which can be opened as a deimatic display when they are disturbed. Males and females fully extend their wings, lift their bodies, and curve up their abdomen, while swaying from side to side or running away ( Fig. 43 ). The red color of the wing together with the black and white streak of the membranous area likely create an intimidating pattern of color and shape for visual predators. Whilst males were sometimes observed flying away when disturbed, females never tried to fly in response to a disturbance; but, they are likely to dampen a fall if dropped while running away (Video S 1).

Food plants: In situ, most specimens of Xerosoma canaliculatum were found in epiphytic Araceae plants, which are common in several Atlantic Forest areas, especially in the trunks of trees, in several distinct locations. Furthermore, whole generations could be bred with only Araceae as their food plant. In captivity, they fed well on several Araceae plant species (i.e., Monstera adansonii , Monstera deliciosa , Xanthosoma violaceum , Epipremnum aureum , Thaumatophyllum bipinnatifidum , Philodendron spp. , and others), but also accepted other plants such as the exotic Jalapa mirabilis and the Neotropical Bougainvillea glabra , both Nyctaginaceae , as well as the native Ludwigia sp. ( Onagraceae ) and Comelina spp. (Comelinaceae). To feed on the leaves, the insects chew from the outside to the inside along the leaf margin, in semicircles ( Fig. 34 ). They usually feed one to three times each night.

Mating behavior: While mating pairs were commonly observed in the rearing cage, only one courtship could be observed and filmed. When the male perceived the female, it moved back-and-forth palpating its partner with its antennae, palps, and abdomen. Then it twisted its abdomen to touch the female ventrally. These movements were alternate and were repeated a few times. The entire process took about one hour. The female eventually refused the male, pushing it away with its mid and hind legs (Video S 2).

When the male is accepted by the female, it uses its thorn pads to attach to the female’s preopercular organ. After fixation, the male extroverts its phallic organ which penetrates the female. This process always occurs at night, when individuals are active, but can extend into the day, with the mating pair remaining immobile ( Fig. 45 ). Males remain on the back of females throughout the entire process, which lasts about eight to ten hours.

In situ, adult males were observed flying, or about to fly, hanging on random plants, shaking their heads and antennae, probably trying to find a female.

Oviposition: The eggs of X. canaliculatum are glued to the substrate, close to food plants, and individually or most frequently in batches of 4– 14 eggs organized in two alternating rows ( Fig. 46 ). In captivity, each observed ñ laid around 100– 200 eggs. The female holds the eggs with the gonapophysis and presses them against the substrate with the cerci for some seconds, making slight lateral movements of its abdomen. A batch of seven eggs was glued in about 20 minutes. In Bertioga, an egg batch was spotted among roots of epiphytic plants in the bark of a tree bearing several Araceae food plants, where young nymphs were previously observed the night before ( Fig. 47 ).

Ecological relations: One ñ nymph in Bertioga at night was parasitized by a fly ( Forcipomyia sp. : Ceratopogonidae : Diptera ) attached to its antenna ( Fig. 48 ).

Figures

Fig. 16. Mounted ñ and ò Xerosoma canaliculatum specimens from Bertioga, São Paulo, Brazil. A: dorsal view of ñ. B: dorsal view of ò. C: lateral view of ñ. D: lateral view of ò. Scale bars = 1 cm.

Fig. 17. ñ Xerosoma canaliculatum holotype. A: labels. B: lateral view. C: dorsal view D: end of abdomen in ventral view. E: ventral view. Scale bars: B, C, E = 1 cm; D = 5 mm.

Fig. 18. Holotype ñ of the synonym Xerosoma senticosum (= X. canaliculatum) (NMW). A: lateral view. B: labels. C: dorsal view. D: end of abdomen in ventral view. E: ventral view. Scale bars: A, C, E = 1 cm; D = 5 mm. Photographs provided by Natural History Museum Vienna, NOaS Image Collection / H. Bruckner; published with permission.

Fig. 19. Male X. canaliculatum from the same collection and locality as the holotype ñ of the synonym Xerosoma senticosum (= X. canaliculatum). Photographs taken at the Natural History Museum Vienna. A: lateral view. B: dorsal view C: labels. D: Head, pro- and mesothorax in lateral view. E: end of abdomen in lateral, dorsal and ventral views. Scale bars:A, C, E = 5 mm; D = 2 mm.

Fig. 20. Head and prothorax of ñ Xerosoma canaliculatum from Bertioga, São Paulo, Brazil.A: 13th and 14th antennomeres. B: head and prothorax in lateral view. C: dorsal view. D: ventral view. E: head in anterior view. F: schematic drawing of mouth parts in ventral view. Abbreviations: Ant, antennae; Bp, Bump; Cn, canaliculus; Cly, clypeus; Cd, cardo; Eye, compound eye; Glo, glossa; Lab, labrum; Me, mentum; Pd, pedicellus; Pgl, paraglossa; Plb, labial palpus; Pme, prementum; Pmx, maxillary palpus; Prn, pronotum; Ptg, prothoracic gland; Sc, scapus; St, stipes. Scale bars: A = 5 mm; B = 1 cm.

Fig. 22. Left legs of ñ Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: front leg in dorsal view. B: front leg in ventral view. C: mid leg in dorsal view. D: mid leg in ventral view. E: hind leg in dorsal view. F: hind leg in ventral view. G: front tarsus in lateral view. Abbreviations: Ap, area apicalis; Ar, arolium; Cl, claw; Eu, euplantula; Fe, femur; Ta1–Ta5, tarsomere; Tb, tibia; Ts, tarsus. Scale bars:A–F = 1 cm; G = 5 mm.

Fig. 23. Posterior region of the abdomen of ñ Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: dorsal view. B: lateral view. C: ventral view. D: 7th sternum in lateral view. E: schematic drawing in ventral view. Abbreviations: Ce, cerci; Gnp, gonaplac; Gp8, 8th gonapophyses; Gp9, 9th gonapophyses; Pre, preopercular organ; Pp, paraproct; Sp, subgenital plate; S7, 7th sternum; T7–T10, terga. Scale bars: A–C = 1 cm; D, E = 5 mm.

Fig. 24. Detail of genital structures of ñ Xerosoma canaliculatum from Aracruz, Espírito Santo, Brazil. A: detail of gonapophyses, gonoplac and subgenital plate in ventral view. B: detail of gonapophyses and gonoplac in dorsal view. C: praeopercular organ in ventral view. D: praeopercular organ in lateral view. Scale bars = 5 mm.

Fig. 25. Egg of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: dorsal view. B: lateral view. C: ventral view. D: anterior view showing the operculum. E: micropylar plate in dorsal view. F: operculum in dorsal view. G: schematic drawing of the micropylar plate. Abbreviations: Op, operculum; Mp, micropylar plate. Scale bars: A–D = 2 mm; F = 2 mm; E, G = 2 mm.

Fig. 26. Egg of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil under scanning electron micrographs (SEM). A: lateral view. B: dorsal view. C: ventral view. D: operculum in lateral view. E: micropylar cup in lateral view. F: micropylar plate in dorsal view. G: detail of capsule surface. H: operculum rim. Scale bars: A, B, C = 500 µm; D, E, F, G, H = 100 µm.

Fig. 27. Head and prothorax of ò Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: 12th and 13th antennomeres. B: head and prothorax in lateral view. C: dorsal view. D: ventral view. E: head in anterior view. F: schematic drawing of the mouth parts in ventral view. Abbreviations: Ant, antennae; Bp, Bump; Cn, canaliculus; Cly, clypeus; Cd, cardo; Eye, compound eye; Glo, glossa; Lab, labrum; Me, mentum; Oc, oceli; Pd, pedicellus; Pgl, paraglossa; Plb, labial palpus; Pme, prementum; Pmx, maxillary palpus; Prn, pronotum; Ptg, protoraxic gland; Sc, scapus; St, stipes. Scale bars = 1 cm.

Fig. 28. Mouth parts of Xerosoma canaliculatum from Aracruz, Espírito Santo, Brazil. A, B: maxilla of ñ in ventral view, right maxilla dissected in B. C, D: mandibles of ñ opened to show internal view. E: internal (dorsal) view of maxilla of ò. F: two left side maxillae of two ñ, right one in internal view and left one in external (ventral) view. G, H: detail of the apex of the galea apex, left maxilla of ñ, antero-internal view (G) and anterointernal view at right side and dorsal view at left side (H). Scale bars:A–F = 5 mm; G–H = not to scale.

Fig. 30. Left legs of ò Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: front leg in dorsal view. B: front leg in ventral view. C: mid leg in dorsal view. D: mid leg in ventral view. E: hind leg in dorsal view. F: hind leg in ventral view. G: front tarsus in lateral view. Abbreviations: Ap, area apicalis; Ar, arolium; Cl, claw; Eu, euplantula; Fe, femur; Ta1—Ta5, tarsomere; Tb, tibia; Ts, tarsus. Scale bars: A–F = 1 cm; G = 5 mm.

Fig. 31. Posterior region of the abdomen of ò Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: dorsal view. B: lateral view. C: ventral view. D: detail of vomer and paraprocts in ventral view. E: schematic drawing in ventral view. Abbreviations: Ce, cerci; Po, poculum; Pp, paraproct; S6–S9, sternum; Tp; thorn pads; T7–T10, terga; Vo, vomer; Wg, wing. Scale bars: A, B, C, E = 1 cm; D = 5 mm.

Fig. 32. Male genitalia of Xerosoma canaliculatum from Aracruz, Espírito Santo, Brazil. A–L: genitalia treated with KOH. M–P: untreated genitalia attached to the body. B, D, F, H, J: same images as respective previous. N, P: same image as respective above, all artificially colored for structure visualization: red, dorsal sclerite; purplish blue, longitudinal lobe; green, basal lobe. A–B: lateral view from left side. C–D: ventral view. E–F: posterior view. G–H: dorsal view I–J: lateral view from right side. K: detail of setae of the basal lobe. L: detail of minute setae of the longitudinal lobe. M–N: genitalia of specimen preserved in alcohol attached to the body in ventral view. O–P: genitalia of recently dead specimen attached to the body in ventral view. Scale bars = 5 mm.

Fig. 33. Male genitalia of Xerosoma canaliculatum from Reserva Vale, Espírito Santo, Brazil treated with KOH and expanded. A: latero-dorsal view. B: lateral view. C: ventral view. D: dorsal view.Abbreviations: Ds, dorsal sclerite, Ll, longitudinal lobe, Bl, basal lobe. Scale bar = 5 mm.

Fig. 34. Live nymphs of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: first instar in lateral view. B: first instar in dorsal view. C: third instar of ò. D: fifth instar of ñ. Scale bars = 1 cm.

Fig. 35. Ventral view of the posterior region of the abdomen of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil showing development in all instars for both sexes (left ñ, right ò). A: first instar. B: second instar. C: third instar. D: fourth instar. E: fifth instar. F: sixth instar. Image not to scale.

Fig. 36. Tarsal morphology of different stages. A: First instar nymph. B: Second instar nymph. C: Third instar nymph. D: Male. E: Female. F: Euplantula 4 of second instar nymph. G: Attachment microstructure on the euplantulae of nymphs and adults. H: Euplantula 4 of the adult female. Abbreviations: ar, arolium; cl, claw; eu 1-5, euplantulae 1 to 5; fs, fissure; L1-3, first to third instar nymphs; ta1-5, tarsomeres 1 to 5. Scale bars: A–E = 500 µm; F = 50 µm; G: 10 µm; H = 150 µm.

Fig. 37. Habitat of X. canaliculatum. A: Parque Nacional de Itatiaia, Rio de Janeiro, Brazil B: Reserva Natural SESC Bertioga, São Paulo, Brazil. Images not to scale.

Fig. 38. Xerosoma canaliculatum from Bertioga, São Paulo, Brazil, nymph under ecdysis process from the second to the third instar. A: nymph in second instar, right before molting. B: nymph under ecdysis process (molting). C: freshly molted third instar nymph. Scale bars = 1 cm.

Fig. 39. Leg regeneration of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: second instar nymph with lost right leg III. B: third instar nymph starting leg regeneration. C: fourth instar nymph with semi-regenerated leg. D, E: ò adult (D) with fully regenerated leg with four tarsi instead of five (E). Images not to scale.

Fig. 40. Xerosoma canaliculatum nymphs from Bertioga, São Paulo, Brazil resting in situ. A: nymph resting on an Araceae food plant stalk. B: nymph resting on an Araceae food plant petiole. C: nymph resting on dead bromeliad leaves. Images not to scale.

Fig. 41. Xerosoma canaliculatum from Bertioga, São Paulo, Brazil, adults resting on branches and bark. A: two ñ on a branch. B: ñ on tree bark. Images not to scale.

Fig. 42. Adults of Xerosoma canaliculatum from Bertioga, São Paulo (A, E, G, H), from Aracruz, Espírito Santo (F), from Jardim Botânico, Rio de Janeiro (C), and Parque Nacional de Itatiaia, Rio de Janeiro (B, D), in different color patterns. A, B: distinct males. C–F: distinct females. G, H: same female during daylight (G) and during the night (H). Images not to scale.

Fig. 43. Xerosoma canaliculatum from Bertioga, São Paulo, Brazil showing deimatic display (A–C) and irritating volatile chemical compound expelled by the prothoracic gland (D). A: ñ. B: ò. C: detail of ñ wing base. C: ñ grabbed by a researcher, arrow showing the chemical compound. Images not to scale.

Fig. 44. Feeding habits of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A: nymph resting on an eaten Epipremnum aureum (Araceae) leaf. B: ò adult feeding on a Philodendron (Araceae) leaf. Image not to scale.

Fig. 45. Copulation process of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil (photographed in captivity). A: mating pair in lateral view, smaller male atop the female. B: contact detail in lateral view. C: contact detail in ventro-lateral view. D: contact detail in posterior view. Image not to scale.

Fig. 46. Oviposition process and egg batch of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil. A, B: female gluing eggs on tree bark. C: glued freshly laid eggs. Image not to scale.

Fig. 47. Xerosoma canaliculatum eggs found in situ in Bertioga, São Paulo, Brazil. A: Base of a tree in the Atlantic Forest covered with X. canaliculatum food plants (Araceae epiphytes), arrow showing location of eggs. B: eggs glued to the tree bark and epiphyte root. Image not to scale.

Fig. 48. Parasite fly (Forcipomya sp.) feeding on the antenna of a fourth instar female nymph of Xerosoma canaliculatum from Bertioga, São Paulo, Brazil.

Fig. 3. Males of Xerosoma species, anterior region of the body in dorsal and lateral views.A: X. canaliculatum in dorsal view. B: paratype (1442) of X. nannospinus sp. nov. in dorsal view. C: X. canaliculatum in lateral view, black arrow indicating the strongly pronotal spines. D: paratype (1442) of X. nannospinus sp. nov. in lateral view, black arrow indicating the reduced pronotal spines. Scale bar = 1 cm.

Fig. 13. Wings of ò Xerosoma nannospinus sp. nov. paratype MZUSP 1441. A: dorsal view with wings opened. B: dorsal view of tegmina closed. Scale bars: A = 1 cm; B = 3 mm.