Exocnophila hastata ( Burmeister, 1838 ) Ghirotto & Conle & Hennemann & Valero & Cancello, 2024

Ghirotto, Victor Morais, Conle, Oskar, Hennemann, Frank, Valero, Pablo & Cancello, Eliana M., 2024, On the Brazilian Exocnophila stick insects (Insecta: Phasmatodea), with the description of a new species, Zootaxa 5536 (1), pp. 59-98 : 69-80

Exocnophila hastata ( Burmeister, 1838) comb. nov.

( Figs 8–18 )

Bacteria hastata Burmeister, 1838: 567 ; Westwood 1859: 348; Redtenbacher 1908: 423; Otte & Brock 2005: 64; Brock & Büscher 2022: 510.

Heteronemia hastata, Kirby 1904: 348 .

Dyme hastata, Zompro 2005: 265 [without expression of new combination].

>> Lectotype (here designated), ♂ ( MfN): hastata, Burm. * ♂ ♀, Brasil. Sello. v. Lgsdf.; 756 ; Bacunculus hastata ♂ Burmeister, 1838 , LECTOTYPUS det. O. Zompro III.2001 ( Fig. 8–10 ) . Paralectotype (here designated), ♀, not found, most likely lost ( MfN): Sello .

Further material examined

1♀ ( MZUSP 2055), eggs: Brasil. Rio de Janeiro. Trajano de Moraes. Visconde de Imbé, Mata Atlântica, 22°05’23.9”S 42°10’13.6”W, on Mimosoideae. i.2021. VM Ghirotto col. Died 12.v.2021; 3♂ ( MZUSP 2056, 1686, 2632), 3♀ ( MZUSP V 0608, 1998, V 0739), eggs: Brasil. Rio de Janeiro. Magé. Vila Mimosa, próx. Camping da Paz, 22°34’03.7”S 43°10’23.8”W, 300m a.s.l. 29.xi.2021. VM Ghirotto & RP Indicatti col.; 4♀ juvenile ( MZUSP V 0610, V 0759, V 0767, V 0786), 4♀ ( MZUSP V 0611, V 0653, V 0785): Brasil. Rio de Janeiro. Angra dos Reis. Pontal, próx. a Camping Recanto do Pontal, 22°56’28.6”S 44°18’58.6”W, on Bambusoideae . VM Ghirotto & RP Indicatti col. 28.xi.2021; 1♀ ( MZUSP 2631), eggs: Brasil. Rio de Janeiro. Magé. Trilha para Pico da Coruja, Santo Aleixo, 22°30’32.5”S 43°02’10.4”W. 23.vii.2022. VM Ghirotto, EB Crispino, PW Engelking, B Ferraz, EA Ribeiro col.; 1♀ ( MZUSP 0564): Brasil. Rio de Janeiro. Teresópolis. Parque Nacional da Serra dos Órgãos, Núcleo Teresópolis. 22º27’14”S 42º59’32”W. 5–6.xii.2015. P.I.Chiquetto-Machado & A.Z.Ramin col.; 1♂ ( MZUSP 1997): Brasil. Rio de Janeiro. Niterói. Parque da Cidade, 22°55’44.9”S 43°05’10.1”W. 19.vii.2022. P.W. Engelking col.; 2♀ ( MNRJ-ENT 15-11, MNRJ-ENT 15-16), 2♂ ( MNRJ-ENT 15-12, MNRJ-ENT 15-13): Brasil. Rio de Janeiro. Rio das Ostras. REBIO União, 22°25’31”S 42°02’21”W. 21–25.ii.2022. P. Souza Dias et al. col.; 2♀ ( MNRJ-ENT 15- 14, MNRJ-ENT 15-15): Brasil. Rio de Janeiro. Nova Iguaçu. REBIO do Tinguá. 14–16.ii.2020. P. Souza Dias, A.M. Siqueira col.; 1♀ ( MNRJ-ENT 15-44): Brasil. Rio de Janeiro. Teresópolis. 2022; 1♀ ( MZUSP 1218): Brasil. Rio de Janeiro. Rio de Janeiro. Jardim Botânico do Rio de Janeiro, 22°58’10”S 43°13’37”W. 18.i.2019. L. Lanna, E.B. Crispino, P.W. Engelking, S.K. Harumi col.

Notes on designation of lectotype. A red label accompanying the lectotype states “ LECTOTYPUS det O. Zompro” but such designation is not valid as it was not published, so it is being designated herein.

Notes on the paralectotype. The female paralectotype was not found in the MfN collection and is very likely lost. This paralectotype is certainly not conspecific nor congeneric with the male as its description states it has a very long, acute subgenital plate, which does not correspond to Exocnophila and even to any closely related genera. Therefore, until now, the female of E. hastata comb. nov. was unknown.

Notes on the locality of the lectotype. A large expedition was organised a few years prior to the description of the species by its collector, von Langsdorf, which started in his own farm in Teresópolis in Rio de Janeiro state in Brazil, so there is a reasonable possibility that the lectotype was collected at or near Teresópolis, where some of the recently collected material comes from.

Diagnosis. Differs from Exocnophila brevitarsata comb. nov. by the more elongate head and tergum VIII, the stockier thorax and the slightly thicker cerci with straight inner margins in males, and the less globose head, longer tergum VIII and undulated margin of tergum X forming three round lobes (almost straight in E. brevitarsata ) in females. Differs from Exocnophila ovifuscum sp. nov. by the thicker cerci in both sexes, by the absence of tubercles in the mesothorax and the less compressed and wider head with a more convex vertex and the roundly undulated margin of tergum X (irregularly and roundly serrate in E. ovifuscum sp. nov.) in females. Further differs from females of Exocnophila cornuta and Exocnophila tuberculata by the less globose head with less prominent vertex and the undulated margin of tergum X forming three round lobes.

Redescription

Male lectotype ( Figs 8–10 ).

Colour ( Fig. 8 ). Irregularly dark brown. Head with a dark longitudinal band bordered by two dark beige bands behind the eye.

Head ( Fig. 9 ). Subrectangular, ca. 1.6–1.7x longer than wide, vertex flat; with irregular granules dorsally mainly in two paramedial rows; very slightly widening towards anterior. Frontal convexity developed, frontal suture deep and angulated. Eyes large, slightly more than hemispherical. Labial palp segments wide, maxillary palp segments slightly wide. Clypeus wide, short, sinuous with ventral margin slightly emarginate. Antennae filiform, reaching tergum VII. Scapus dorsoventrally compressed, large, wide, longer than wider in dorsolateral view. Scapus with a dorsolateral carina separating dorsal and lateral surfaces, internal margin narrower at posterior edge, posteriorly very slightly round and then straight, external margin wide and round through all its length. Pedicel more than half the length of scapus, longer than wide. Antennae with around 38 articles.

Thorax ( Figs 8–10A ). Surface smooth with sparse minute setae. Pronotum longer than wide, slightly constricted premedially, slightly narrower anteriorly; smooth ( Fig. 9A–B ). Probasisternum with longitudinal sulcus, anteriorly narrow, posterolateral margins round; with short stiff setae mainly near lateral edges ( Fig. 9C ). Profurcasternum trapezoid to sudquadrate, covered with short stiff setae ( Fig. 9C ). Mesothorax 7.2x longer than prothorax. Meso and metafurcae in “Y” shape. Mesothorax 1.5x longer than metathorax. Meso and metanotum with faint longitudinal medial carina. Metanotum 7.9x the length of median segment.

Legs ( Fig. 8 ). Hind legs slighty longer than anterior legs;mid legs shortest. Coxae smooth.Profemur considerably longer than mesothorax. Mesofemur around 0.8x the length of profemur. Profemur with distinct basal curvature occupying around one sixth of the segment. Metafemur with a basal swelling occupying around one eightieth to one seventh of the length of the segment. Carinae of femora and tibiae with sparse setae between them and bearing row of porrect spiniform setae larger in the tibiae, longer than those of the female. Carinae of tibiae presenting small short spines at the apex, one in dorsal carinae, four to seven in lateroventral carinae and five to 10 in the ventral carina, with more spines in mid and hind tibiae. Basitarsi short, around the same size than the respective following three tarsomeres combined; ventrally with dense tuft of stiff, porrect setae. Euplantulae well developed and present in all tarsomeres.

Abdomen ( Figs 8 , 10 ). Surface smooth, without granules but with carinae. Median segment well marked by a sulcus, around 0.13x the length of the metanotum ( Fig. 10A ). Terga II–X longer than wide. Terga II–VII with two pairs of paramedial longitudinal carinae near lateral margins; IV–VII with further two pairs of faint carinae near centre; II–VIII with a further short longitudinal carinae near anterior margin. Tergum X with one pair of paramedial carinae and a longitudinal medial carina; posterior margin elevated and widely, roundely emarginate into two somewhat triangular lobes bearing thorn pads at the apexes ( Fig. 10B ). Tergum X very slightly shorter than IX; both shorter than VIII. Tergum VIII widening towards posterior and IX tapering towards posterior; X wider centrally. Sterna with pair of paramedial carinae near lateral margins and IV–VII with a further, faint pair. Poculum well developed with angulated posteroventral tip forming a laterally compressed lobe pointing backwards ( Fig. 10C–D, F ); anterior region of sternum IX short; cerci elongate and dorsoventrally compressed, with straight inner margins, ellipsoid, apex round ( Fig. 10E ), thornpads apart, well developed with around 10 incurved teeth varying in size ( Fig. 10E–F ). Vomer well developed, triangular, terminal hook short, thin, curved upwards.

Measurements (in mm), lectotype. Body (without cerci) 71.6, head 3.3, antennae at least 53.0, pronotum 2.6, mesonotum 20.2, metanotum and median segment 13.9, abdomen (excluding median segment) 31.2, profemur 25.2, protibia 30.5, mesofemur 20.8, mesotibia 26.5, metafemur 26.2, metatibia 35.7.

Variation in males ( Figs 11A–D , 15–17 ). Colour in different shades of brown, olive, gray, with or without whitish irregular small spots. Head slightly narrower ( Fig. 15 ). Thorn pads with 8–11 teeth ( Fig. 17E ). Mesonotum 7x longer than median segment which is 0.14x of mesonotum length.

Measurements (in mm), N = 2. Body (without cerci) 73.7–75.8, head 3.1–3.2, antennae 59.0–63.1, pronotum 2.8, mesonotum 20.4–21.5, metanotum 12.4–13.7, median segment 1.5–1.6, abdomen (excluding median segment) 33.1–33.4, profemur 24.7–25.6, protibia 30.8–31.6, mesofemur 20.0–21.1, mesotibia 24.6–25.4, metafemur 23.2– 24.6, metatibia 34.0–34.2.

Female ( Figs 11E–H , 12–14 ).

Colour ( Figs 11E–H , 12–14 ). Entirely beige, beige orangish, brown, reddish brown, orangish brown, gray, or dark brown, with or without irregular stains, with or without a longitudinal black line in pronotum to the end of the abdomen. Head with a dark longitudinal band bordered by two dark beige bands behind the eye.

Head ( Fig. 12 ). Subrectangular, ca. 1.6x longer than wide, smooth, vertex gently convex; with irregular granules dorsally, the central ones the largest, in pair or irregular ( Fig. 12A–B ); in some individuals the central pair is developed into large, irregular leaf-like lobes. Frontal convexity developed, frontal suture deep and roundly angulated. Eyes hemispherical. Labial palp segments wide, maxillary palp segments slightly wide ( Fig. 12C–D ). Clypeus wide, short, sinuous with ventral margin slightly emarginate.Antennae filiform, reaching tergum II. Scapus dorsoventrally compressed, large, very wide, longer than wider in dorsolateral view. Scapus with a dorsolateral carina separating dorsal and lateral surfaces, internal margin narrower at posterior edge, posteriorly slightly round and then straight, external margin wide and round through all its length ( Fig. 12A–B ). Pedicel more than half the length of scapus, longer than wide. Antennae with 37 articles.

Thorax ( Figs 11E–H , 12–13 ). Surface smooth with very few sparse round short granules and presenting irregular carinae. Pronotum longer than wide, slightly constricted premedially, slightly narrower anteriorly; with paramedial granules larger at posterior half ( Fig. 12A–B ). Probasisternum with longitudinal sulcus, anteriorly narrow, posterolateral margins round; with short stiff setae mainly near lateral edges ( Fig. 12C–D ). Profurcasternum round with an anterior trapezoidal region densely covered with short stiff setae ( Fig. 12C ). Mesothorax 5–5.2x longer than prothorax; gradually widening posteriorly. Mesonotum, metanotum, mesosternum and metasternum with distinct pair of lateral carinae ( Figs 11E–H , 13 A–B ). Meso and metafurcae in ‘’Y’’ shape, metafurca wider and shorter than mesofurca. Mesothorax 1.8x longer than metathorax. Metathorax with irregular sinuous paramedial carinae ( Fig. 13A–B ). Metanotum 4.8–5.5x the length of median segment.

Legs ( Figs 11E–H , 13C–H ). Anterior legs only slightly shorter than hind legs; mid legs shorter than both. Hind legs considerably exceeding end of the abdomen. Coxae smooth. Profemur slightly longer than mesothorax. Mesofemur around 0.8x the length of profemur. Profemur with distinct basal curvature occupying slightly more than one third of the segment ( Fig. 11E ). Metafemur with a basal swelling occupying slightly less than one sixth of the length of the segment ( Fig. 11E ). Carinae of femora and tibiae with sparse setae between them and bearing row of short porrect spiniform setae larger in the tibiae ( Fig. 13C–H ). Carinae of tibiae presenting small short spines at the apex, one to two in dorsal carinae, three to eight in lateroventral carinae and nine to 12 in the ventral carina, with more spines in mid and hind tibiae ( Fig. 13F–H ). Basitarsi short, pro and metabasitarsi around the same size and mesobasitarsi slightly shorter than the respective following three tarsomeres combined; ventrally with dense tuft of spiniform, thick setae ( Fig. 13C–E ). Euplantulae well developed and present in all tarsomeres ( Fig. 13C–E ).

Abdomen ( Figs 11E–H , 13A–B , 14 ). Surface slightly rugose, with granules and with several carinae ( Figs 13A–B , 14A–F ). Median segment well marked by a sulcus, around 0.2x the length of the metanotum, bearing two sinuous pairs of paramedial carinae ( Fig. 13A ). Terga II–VIII and X longer than wide, IX slightly shorter than wide to as long as wide. Terga II–IX with four to five pairs of paramedial longitudinal carinae, X with two pairs and a single medial longitudinal carina. Tergum X with posterior margin undulate forming three round lobes, a medial one slightly shorter to as the same size as the others, and two aside ( Fig. 14A–B ); laterally upcurving, shorter not covering the cerci from lateral view ( Fig. 14C–D ). Sterna with more setae than terga, bearing two pairs of paramedial carinae. Praeopercular organ developed, swollen, rugose, slightly sclerotised ( Fig. 14E ). Subgenital plate convex, with two strong paramedial carinae in posterior three fourths, with round apex, reaching half the length of tergum X or slightly more ( Fig. 14C–F ). Cerci very short, centrally widened, apex round or with a wide angulation, ventrally concave, not surpassing the posterior margin of tergum X ( Fig. 14E–F ). Paraprocts divided posteriorly, each with round posterior margin ( Fig. 14E ). Epiproct wide. Gonapophyses and gonoplac covered by the subgenital plate, all elongate and gradually tapering to a somewhat pointy apex; gonapophyses VIII the longest, gonoplac slightly shorter, upcurving, gonapophyses IX shorter than both ( Fig. 14G ).

Measurements (in mm), N = 4. Body (without cerci) 60.9–73.0, head 3.7–4.5, antennae 31.0–40.1, pronotum 3.0–3.4, mesonotum 15.7–19.0, metanotum 7.9–9.9, median segment 1.5–1.9, abdomen (excluding median segment) 29.1–34.4, profemur 16.3–19.2, protibia 18.1–22.0, mesofemur 13.4–15.6, mesotibia 13.8–16.8, metafemur 16.2– 19.8, metatibia 18.8–23.7.

Egg ( Fig. 18 ). Laterally flattened ( Fig. 18E ), oval in lateral view and ellipsoid in dorsal view, opercular collar constricted on the dorsal surface from the end of the micropylar plate to the operculum ( Fig. 18A–D, I ); 1.2–1.3x longer than tall and 1.8–1.9x longer than wide. Capsule surface densely granulated with minute wide rounded prominences. Micropylar plate ellipsoid, elongated around 1.5–2.2x longer than wide, with round, parallel or slightly constricted lateral edges ( Fig. 18E–G ). Micropylar plate internally flat and smooth. Edge of the micropylar plate raised, prominent and divided into a rougher inner part with a foamy appearance, interrupted by the micropyle, and a narrower outer part, slightly rough, bordering the micropyle, with a smooth inner margin and a rougher outer margin with texture similar to the inner part; these two parts are interrupted by a narrow depression ( Fig. 18E–G ). Micropyle oval, with a rounded projection between the inner edges posteriorly bordered by a rounded depression. Median line very short, truncated, thick and continuous with the outer edge of the external part of the micropylar plate ( Fig. 18E–G ). Operculum round, with a raised and convex ring with a surface similar to the rest of the capsule in the outer half of the diameter and with a round, smooth central depression ( Fig. 18H ). Entirely orange to orangish brown in colour with sparse irregular black dots, micropylar plate surrounded by a creamish rim very short at anterior and posterior edges; with little extension in the anterior and posterior parts; micropylar plaque internally orange darkening towards polar area, inner part of the border ice-coloured and outer part of the border beige to yellowish.

Distribution ( Fig. 31 ). Known from the Atlantic Forest of Rio de Janeiro, recorded for Parque Nacional da Serra dos Órgãos (municipalities of Magé and Teresópolis) and Angra dos Reis, Nova Iguaçu, Rio das Ostras, Rio de Janeiro and Trajano de Moraes in Rio de Janeiro state, Brazil.

Biology. Individuals were only found at 0–180 cm above the ground, but mostly near the ground up to 40 cm. Specimens present a still behaviour, never running away, even when disturbed. If disturbed and even during handling, specimens assume camouflage position and remain motionless. A few specimens may take gradual, interrupted steps upon constant handling. Specimens may also spew stomach content or release a light minty scent from their prothoracic glands. In nature, specimens were observed eating a variety of plants, including undetermined Bambusoideae ( Poaceae ), Asteraceae and Malvaceae plants, and undetermined Mimosoidea ( Fabaceae ) trees very similar to representatives of the Anadenanthera genus. In captivity, specimens ate garden roses ( Rosa spp. , Rosaceae ), Lantana camara ( Verbenaceae ), Hibiscus spp. , Malvaviscus arboreus , Callianthe picta ( Malvaceae ), Anadenanthera sp. , Calliandra spp. , Piptadenia sp. (Mimosoideae: Fabaceae ), Psidium guajava , Eugenia uniflora ( Myrtaceae ), and Passiflora suberosa ( Passifloraceae ).

Figures

FIGURE 8. Exocnophila hastata comb. nov., male body and habitus, lectotype. A, habitus in dorsal view; B, body in dorsal view; C, habitus in lateral view. Scale bars: 2 mm.

FIGURE 9. Exocnophila hastata comb. nov., head and thorax, lectotype.A, dorsal view; B, lateral view; C, ventral view. Scale bars: 2 mm.

FIGURE 10. Exocnophila hastata comb. nov., male thorax and abdomen, lectotype.A, metathorax in dorsal view; B, terminalia in dorsal view; C, terminalia in lateral view; D, terminalia in lateral view after warming in hot water to spread cercus; E, tergum X and cercus in ventral view; F, terminalia in ventral view. Scale bars: 2 mm.

FIGURE 11. Exocnophila hastata comb. nov. from Rio de Janeiro, Brazil, male V0613 (A–D) and female MZ0564 (E–H) body and habitus. A, E, habitus in dorsal view; B, F, body in dorsal view; C, G, body in lateral view; D, H, body in ventral view. Scale bars: 2 mm.

FIGURE 12. Exocnophila hastata comb. nov., head and thorax, females from Rio de Janeiro, MZV0677 (A–C), MZ0564 (D). A, dorsal view; B, lateral view; C–D, ventral view. Scale bars: 1 mm.

FIGURE 13. Exocnophila hastata comb. nov., thorax and abdomen, females from Rio de Janeiro, MZV0677 (A, C–H), MZ0564 (B). A–B, metathorax in dorsal view; C, F, right protarsus, ventral view, F showing detail of tibia apex; D, G, right mesotarsus, ventral view, G showing detail of tibia apex; E, H, right metatarsus, ventral view, H showing detail of tibia apex. Scale bars: A–E = 1 mm; F–H = 0.5 mm.

FIGURE 14. Exocnophila hastata comb. nov., abdomen, females from Rio de Janeiro, MZV0677 (A, C, E, G), MZ0564 (B, D, F). A, abdomen in dorsal view; B, terminalia in dorsal view; C–D, terminalia in lateral view; E–F, terminalia in ventral view; G, valves in lateral view with subgenital plate spread out. Scale bars: 1 mm.

FIGURE 15. Exocnophila hastata comb. nov., head and thorax, male V0608 from Rio de Janeiro, Brazil.A, C, dorsal view; B, ventral view; D, lateral view. Scale bars: 1 mm.

FIGURE 16. Exocnophila hastata comb. nov., thorax and abdomen, male V0608 from Rio de Janeiro, Brazil. A, median segment, dorsal view; B, metathorax, dorsal view; C, protarsi, ventral view; D, right mesotarsus, ventral view, E, apex of tibia of left leg, ventral view; F, apex of tibia of right leg; G, right metatarsus, ventral view. Scale bars: A–D, F–G = 1 mm; E = 0.5 mm.

FIGURE 17. Exocnophila hastata comb. nov., terminalia, male V0608 from Rio de Janeiro, Brazil. A, dorsal view; B, lateral view; C, ventral view; D, cerci and vomer in ventral view; E, thorn pads in ventral view. Scale bars: 1 mm.

FIGURE 18. Exocnophila hastata comb. nov., eggs from different females from Rio de Janeiro, Brazil.A, E, H, I, from Magé, Rio de Janeiro (MZV0677); B, F, from Trajano de Moraes, Rio de Janeiro (MZV0549); C, D, G, from Angra dos Reis, Rio de Janeiro (MZV0611). A–D, different eggs in lateral view; E–G, different eggs in dorsal view; H, egg in anterior view showing operculum; I, several eggs in the same scale in lateral view. Scale bars: A–G, I = 1 mm; H = 0.5 mm.

FIGURE 31. Map of the known distribution for Exocnophila species, in southeast Brazil. Precise localities are known only for Rio de Janeiro state; the inlet map shows Brazilian states with records for the genus highlighted in red, map colours indicating biomes (dark green: Amazon, green: Atlantic Forest; orange: Cerrado; beige: Caatinga).