Thirteen new species of Urucumania (Phasmatodea: Pseudophasmatidae) stick insects from Brazil and Bolivia

Abstract

Stick insects (Phasmatodea) in general are remarkable for presenting striking camouflage-related adaptations. Even so, a considerable number of species are colorful and present other defense mechanisms, such as the New World lineage Pseudophasmatidae. Within this family, Urucumania currently comprises two described species occurring in Brazil, Bolivia and Paraguay, both of which have small scale-like wings. Urucumania is currently placed in Anisomorphini. However, two similar species under Pseudophasma present remarkable similarities with Urucumania species. These two species, Pseudophasma nigrovittatum and Pseudophasma dentata, are the only Pseudophasma presenting scale-like wings. Both inhabit Brazil, are known from a single sex and present a generally similar coloration pattern contrasting to that of Urucumania. Aiming to resolve whether these and other similar specimens represent distinct species, and if they belong to Urucumania rather than to Pseudophasma, we conducted a careful analysis of the external morphology of both sexes, eggs and the male genitalia. We transfer both scale-like winged Pseudophasma to Urucumania, redescribing them based on both sexes showing that the two described species are in fact distinct, and furthermore present 13 new species of Urucumania from Brazil and Bolivia: Urucumania pirulai sp. n., U. varellai sp. n., U. guadanuccii sp. n., U. atilai sp. n., U. intervalica sp. n., U. brasil sp. n., U. tapirape sp. n., U. rasocatarinensis sp. n., U. dilatata sp. n., U. sertaneja sp. n., U. albopunctata sp. n., U. oriomadeirensis sp. n. and U. candanga sp. n.

Introduction

Several insect lineages developed camouflage-related traits. Stick insects (Phasmatodea) are remarkable among them presenting striking adaptations, such as a very elongate body, cryptic coloration and forelegs that when extended conceal the head (Bradler and Buckley 2018). Nevertheless, colorful or very robust forms are not rare. These less cryptic representatives usually present other defense mechanisms such as large spines or the capacity of spraying an irritating substance from the prothoracic gland, an organ typical of Phasmatodea (Bradler and Buckley 2018).

In the Neotropics, one of the world’s hotspots of Phasmatodea diversity, many of the colorful sprayer phasmids belong to the Pseudophasmatidae lineage (Brock et al. 2024). To date 325 species are described for this family (Brock et al. 2024), which is divided into three subfamilies, including Pseudophasmatinae containing around one third of that diversity. This group is further subdivided into the tribe Paraprisopodini Zompro, 2004, containing only a peculiar genus specialized in bark camouflage, the tribe Anisomorphini Redtenbacher, 1906 and the tribe Pseudophasmatini Kirby, 1904. While a proper delimitation of Anisomorphini and Pseudophasmatini is still pending, their type genera are easily distinguishable: Anisomorpha Gray, 1835 presents short and flattened subgenital plates for both sexes and Pseudophasma Kirby, 1896 presents longer, convex subgenital plates for both sexes with the male poculum strongly convex and bearing a lateral finger-like projection (Conle and Hennemann 2002). Species of these groups are able to spray a pungent, minty substance from the prothoracic glands and often display characteristic coloration patterns including lines, stains and contrasting tones. The coloration pattern differs among species and is often used as diagnosis for new species (e.g., Conle et al. 2011).

The Anisomorphini genus Urucumania Zompro, 2004 currently comprises two described species presenting small scale-like wings and a robust body with relatively stout legs, occurring in Brazil, Bolivia and Paraguay. The type species Urucumania urucumana (Giglio-Tos, 1910) was described from females from the Urucum massif in the Pantanal in central Brazil, from where it takes the specific name and later the generic name as given by Zompro (2004). Urucumania borellii (Giglio-Tos, 1897), described from males and females from Paraguay, was also included by Zompro (2004) when stablishing the genus. Prior to the transfer of U. borellii to its current genus, Anisomorpha lurida Redtenbacher, 1906, also from Paraguay, was synonymized under the former by Conle and Hennemann (2002), an act accepted by Zompro (2004).

Two Brazilian species currently placed in Pseudophasma present remarkable similarities with Urucumania, also presenting a robust body with relatively large legs, being the only ones in the genus to bear scale-like wings. These species, Pseudophasma dentata (Stål, 1875) and Pseudophasma nigrovittatum (Piza, 1939), the former known only from the female and the latter from the male, show a similar overall contrasting coloration of black and beige to orange but with slightly differing patterns. Additionally, we observed that specimens from other localities also show slightly differing patterns with similar general colors. We conducted a careful analysis of the external morphology of both sexes, eggs and male genitalia on several specimens in order to resolve whether P. dentata and P. nigrovittatum should be transferred to Urucumania, and if they truly represent distinct species or are conspecific and together with specimens from other localities represent a single species with polymorphism of color pattern.

Therefore, in the present work we propose the new combinations Urucumania dentata comb. n. and Urucumania nigrovittata comb. n., redescribing both species based on type and new material collected near the type localities, and pointing out morphological differences between the two species. We also describe the male of U. dentata comb. n., the female and egg of U. nigrovittata comb. n., and 13 new species of Urucumania: Urucumania pirulai sp. n. (from Parauapebas, Pará, Brazil), Urucumania varellai sp. n. (from Abel Figueiredo, Pará, Brazil), Urucumania guadanuccii sp. n. (from Diamantina, Minas Gerais, Brazil), Urucumania atilai sp. n. (from São Miguel Arcanjo and Iperó, São Paulo, Brazil), Urucumania intervalica sp. n. (from Ribeirão Grande and Tapiraí, São Paulo, Brazil), Urucumania brasil sp. n. (from Loreto, Maranhão, Brazil), Urucumania tapirape sp. n. (from Barra do Tapirapé, Mato Grosso, Brazil), Urucumania rasocatarinensis sp. n. (from Jeremoabo, Bahia, Brazil), Urucumania dilatata sp. n. (from Umburanas and Sento Sé, Bahia, Brazil), Urucumania sertaneja sp. n. (from Piranhas, Alagoas, and Natal, Rio Grande do Norte, Brazil), Urucumania albopunctata sp. n. (from Santa Cruz, Buena Vista, Bolivia), Urucumania oriomadeirensis sp. n. (from Itaituba and Xinguara, Pará, and Itacoatiara, Amazonas, Brazil) and Urucumania candanga sp. n. (from Brasília, Distrito Federal, Brazil).

Material and Methods

The analyzed specimens are housed at the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP) except for the holotype of U. nigrovittata comb. n., housed at Museu de Entomologia Luiz de Queiroz, Piracicaba, Brazil (MELQ), the type material of U. dentata comb. n., housed at Naturhistorisches Museum Wien, Vienna, Austria (NMW), type material of Urucumania albopunctata sp. n., housed at the collection of the Georg August University of Göttingen, Göttingen, Germany (Göttingen), type material of Urucumania candanga sp. n., housed at the collection of the Universidade de Brasília, Brasília, Brazil (UNB) and non-type material of Urucumania sertaneja sp. n., housed at Museum Paraense Emílio Goeldi, Belém, Brazil (MPEG). Supplementary information deduced from specimen labels is given in brackets. Depository institution for each specimen is given in parenthesis.

Biological observations were based on live specimens collected by active searches at night. Dry-preserved eggs were examined when possible, except for the eggs of U. dentata, extracted from within a female abdomen. Specimens and eggs were examined at Universidade Estadual Paulista “Júlio de Mesquita Filho”, Rio Claro campus (UNESP), and at MZUSP under Leica M125 and Leica M205 C stereomicroscopes and measured with a Mitutoyo digital caliper. Photographs of live specimens and the habitus were taken with a Canon EOS Rebel T5 camera equipped with a Canon EF-S 18–55 mm lens; a Canon EOS Rebel SL1 camera equipped with a Canon EF 100 mm F/2.8 Macro USM lens; or with a Canon EOS 70D camera equipped with a Canon 100 mm Macro lens. Pinned, dry preserved specimens were photographed with a Canon EOS Rebel SL1 camera equipped with a Canon EF 100 mm F/2.8 Macro USM lens and a Nikon D5000 camera equipped with a Nikon DX AF-S Nikkor 18-140 mm 1:3 5-56G ED lens. Specimens and structures were photographed under a Leica M205 C stereomicroscope with LAS Core software or with a Zeiss MRc5 camera coupled to a Zeiss SteREO DiscoveryV12 stereomicroscope and focus-stacked in the software Helicon Focus version 6.3.7 Pro. Illustrations were made using Adobe Photoshop CS6. The distribution map was prepared in QGIS 3.28.

Redescriptions are based on type specimens and the additional material cited for each species. Color descriptions are based on live specimens when possible, but the colors change in most dried specimens, especially due to fading. Measurements are given in millimeters, and the body is measured from the apex of the head to the posteriormost portion of the abdomen. In the text, the median segment can be referred to as “tergum I” when descriptions concern more parts of the abdomen. General terminology of the body follows Ghirotto (2021), that of the egg capsule follows Clark Sellick (1997), that of the terminalia follows Bradler (2009), and that of the internal male genitalia follows Chiquetto-Machado and Cancello (2021) and Ghirotto (2021).

Order Phasmatodea Jacobson and Bianchi, 1902

Family Pseudophasmatidae Rehn, 1904

Subfamily Pseudophasmatinae Rehn, 1904

Genus Urucumania Zompro, 2004

(Figs. 1, 2)

Fig. 1

Genital structures of Urucumania. a–d Entire phallic organ in dorsal (a), ventral (b), caudal (c), left side (d), right side (e), and caudal view of the internal sclerite (f). In red, dorsal sclerite; orange, dorsal lobe; purple, ventral lobe; pink, ventral sclerite; yellow, internal sclerite; green, main process of internal sclerite; blue, accessory process of internal sclerite. Scale bars = 1 mm

Fig. 2

Urucumania urucumana. Couple from Serra do Urucum, Mato Grosso, Brazil. a–b Female in dorsal view (a) and female terminalia in lateral view (b). c–f Male in dorsal view (c) and male terminalia in dorsal (d) and lateral (e, f) views. Scale bars = 0.5 mm

Type species: Anisomorpha urucumana Giglio-Tos, 1910:8, by original designation.

Otte and Brock 2005 (citation in catalog); Brock and Büscher 2022 (citation in catalog).

Characteristics. The male genitalia of Urucumania species consist of a dorsal, a ventral and an internal sclerite, and a ventral and a dorsal lobe (Fig. 1). The dorsal sclerite is nominally divided into basal part and apical part, the latter bearing a main and a distal process (Fig. 1a). The ventral sclerite covers the tip of the ventral lobe and has a small narrow bump anteriorly. The internal sclerite is somewhat crown-shaped and is also nominally divided, into a main process bearing four projections, the two centralmost projections and the outer two flank projections, and an accessory process bearing an inner branch and an outer branch which usually bears a distal and a proximal part (Fig. 1f). The dorsal lobe is short and originates atop the dorsal sclerite.

External characteristics of Urucumania are illustrated in reference to the morphology of the type species of the genus, Urucumania urucumana, in Fig. 2, from males and females collected very near the type locality (Urucum massif in Corumbá and Ladário, Mato Grosso do Sul, Brazil, in the same mountain, the Urucum massif, where type specimens are from). Both sexes possess small, scale-like wings. The male has an emarginated posterior margin of the tergum X with small thorn pads pointing inward, near median line, with small teeth, and has a very large, bulgy poculum with large anterior portion further bearing an elongated dextral process originating superiorly from the anterior portion of sternum IX. The cerci of the female are short and cylindrical, and those of the male are longer, usually gently incurved.

Biology. Most Urucumania specimens observed alive are very active, moving a lot upon disturbances such as vibrations or handling. During the day, it is common for them to hide under logs, large leaves or stones, near the ground. They are able to spray a minty, slightly irritating substance from their prothoracic glands, if grabbed or poked. In captivity, specimens fed on Lantana (Verbernaceae), Ligustrum (Oleaceae), Rosa (Rosaceae), Hibiscus (Malvaceae) and Oldenlandia (Rubiaceae). Urucumania rasocatarinensis sp. n. were observed to feed on an undetermined Spermacoceae (Rubiaceae) plant in situ (Fig. 46).

Identification key. We provide an identification key for Urucumania based on adult males and females. We refrain from using eggs in the key due to its higher variation, generalized morphology between similar species and the lack of knowledge of the egg of several species.

• 1. Well-defined pale dorsolongitudinal band, without a dorsolongitudinal black line in the center……........ 2

  - Pale dorsolongitudinal band absent or present but with a black line in the center..................…….. 4

• 2. Vestigial tegmina and alae present. Males with a finger-like process present only on the right side …………………………………………………………………………….. 3

  - Apterous. Males with a finger-like process present on both sides, asymmetric ……………………………………………………………….. U. albopunctata sp. n.

• 3. Abdominal terga posteriorly smooth in lateral view, without ornamentations on males and with only small bumps on the posterior margin of each terga, not spine-shaped. Males with a finger-like process widened towards the middle, forming a silhouette similar to that of a boomerang. Females with a rounded posterior margin of the anal segment ……………………………………………………………………U. candanga sp. n.

  - Presence of a small, dorsomedial, spine-shaped ornamentation near the posterior margin of each abdominal tergum. Males with a simple and slender finger-like process. Females with a truncated, quadrangular posterior margin of the anal segment ……U. intervalica sp. n.

• 4. Without a pale band or dorsolongitudinal black line …………………………… 5

  - With a dorsolongitudinal pale band and black line ... 9

• 5. Beige or yellow band along the lateral and posterior margins of the pro-, meso-, metathorax, as well as abdominal terga I-VIII ………………………. U. brasil sp. n.

  - Nota and terga uniformly colored or with small discontinuous beige marks …….. 6

• 6. Black tegmina and alae, but with yellowish or orangish overall tone due to venation strongly colored in those tones ………………………………………………….. 7

  - Tegmina and alae without marked venation in a different color ……………………. 8

• 7. Body and legs uniformly black, with the base of the femora at most slightly orange. In males, the finger-like process clearly extends beyond the posterior margin of tergum IX …………………………………………………………………….. U. dilatata sp. n.

  - Posterior margin of abdominal terga I-VIII with discontinuous beige marks. Legs orange-brown, only dark at the junction between femora and tibiae. In males, a short finger-like process, not extending beyond the posterior margin of tergum IX ………. U. urucumana

• 8. Overall body color and legs dark brown, almost black. Alae and tegmina of the same color as the body but bordered by a thin beige line ………………… U. sertaneja sp. n.

  - Overall dark body color with dorsal tones of camel brown to dark reddish. Femora reddish at the base, darken progressively towards the apex. Alae and tegmina uniformly colored in beige ……………………………………………………………………….. U. borellii

• 9. Absent dorsolongitudinal light band but thin black line present along the head and thorax ………………………………………………………………………………….. 10

  - Dorsolongitudinal light band and black line present ………………………….. 11

• 10. Thin dorsolongitudinal line along the head, pro- and mesonotum; becomes diffuse from the metanotum. Femora reddish-brown and tibiae black ……. U. oriomadeirensis sp. n.

  - Wide dorsolongitudinal line along the head, nota, and terga. Femora nearly black and tibiae bronze-brown …………………………………………………. U. tapirape sp. n.

• 11. Apex of vestigial tegmina extending beyond the posterior margin of the mesonotum ………………………………………………………………………………….. 12

  - Vestigial tegmina not extending beyond the posterior margin of the mesonotum ….. 13

• 12. Posterior margin of the anal segment in females without medial incision. In females, head slightly wider than pronotum and tergum X not shortened, only 1.2 × wider than long …………………………………………………………………… U. varellai sp. n.

  - Posterior margin of the anal segment in females with slight medial incision. In females, head as wide as pronotum and tergum X short in length, 2.4 × wider than long …………………………………………………………………… U. pirulai sp. n.

• 13. Femora and tibiae uniformly colored in the same general tone as the body …… 14

  - Femora and/or tibiae with two different color tones …………………………… 15

• 14. Wide black dorsolongitudinal line along the head, thorax, and abdominal terga I-VII. Overall body color dark brown. Finger-like process in males slender and not exceeding the posterior margin of tergum IX …………………………… U. rasocatarinensis sp. n.

  - Very thin black dorsolongitudinal line, not extending beyond abdominal tergum III. Overall body color black. Finger-like process in males widened in the center and clearly extending beyond the posterior end of tergum IX ………………. U. guadanuccii sp. n.

• 15. Black femora except at the base, which is reddish-brown. Tibiae reddish-brown, orange or dark brown to blackish ………………………………………………………….. 16

  - Black femora. Tibiae black except at the base, which is yellowish-brown or beige ………………………………………………………………….. U. nigrovittata

• 16. Vestigial tegmina yellow; vestigial alae black and smaller than the tegmina. Tibiae dark brown to blackish. Body sprinkled with conspicuous, very light dots contrasting with background and noticeable from full body view ……………… U. atilai sp. n.

  - Vestigial tegmina yellow; vestigial alae same color as tegmina and smaller. Tibiae reddish brown to orange. Body sprinkled with small light dots that are not noticeable from full body view ………………………………………………………………………….. U. dentata

Urucumania dentata (Stål, 1875) comb. n.

(Figs. 3, 4, 5, 6, 7 and 8)

Fig. 3

Urucumania dentata comb. n., holotype female from Santa Catarina, Brazil (NWM3517/5317). a Body in dorsal view. b Abdomen in lateral view. c Head in dorsal view. d Head in lateral view. e Terminalia in dorsal view. f Terminalia in ventral view. Scale bars: a 1 mm; b–f 0.5 mm 

Fig. 4

Urucumania dentata comb. n., couple from Ilha de Santa Catarina, Florianópolis, Santa Catarina, Brazil. a, b Female in dorsal (a) and lateral (b) views. c, d Male in dorsal (c) and lateral (d) views. Scale bars = 2 mm

Fig. 5

Urucumania dentata comb. n., couple from Ilha de Santa Catarina, Florianópolis, Santa Catarina, Brazil. a–h Male. i–o Female. a, c, g, i, l Head and pronotum in dorsal (a, i), lateral (c, l) and ventral (g) views. b, n Metathorax in dorsal view. d, e, f, h, j, m, o Terminalia in dorsal (d, j), left side (e, m), right side (h) and ventral (f, o) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 6

Urucumania dentata comb. n., phallic organ of male from Ilha de Santa Catarina, Florianópolis, Santa Catarina, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 7

Urucumania dentata comb. n., egg of female from Ilha de Santa Catarina, Florianópolis, Santa Catarina, Brazil. a Lateral view. b Dorsal view, inlet showing mushroom-like structures. c Anterior or opercular view. Scale bars = 1 mm

Fig. 8

Urucumania dentata comb. n., live females from Santa Catarina, Brazil, in situ. a Adult female from Balneário Camboriú, photo courtesy of Felipe Willian Borges Alves. b Female juvenile from Florianópolis, Ilha de Santa Catarina, photo courtesy of Camila T. Cegoni

Anisomorpha dentata Stål, 1875; Kirby 1904 (citation in catalog); Redtenbacher 1906 (redescription, first description of male, illustration); Brock 1998 (type data); Domínguez and Vera 2014 (citation in checklist).

Neophasma dentata, Conle and Hennemann 2002 (redescription, description of egg); Otte and Brock 2005 (citation in catalog).

Pseudophasma dentata, Araujo and Garraffoni 2012 (citation in checklist); Brock and Büscher 2022 (citation in catalog).

Redescription

Female. Color (Figs. 4a, b, 8). Head and body dark red to dark orange, with a black medial longitudinal line running along entire length of head and thorax, sparsely in median segment, and continuing punctuated along the black posterior projections of terga I–VII and along the black elevation of terga VIII and IX; some carinae and granules with darker irregular stains. Head with further paramedian darker lines; antennae dark red to dark orange. Ventrally irregularly dark red to dark orange with irregular stains. Femora black with dark red to dark orange at basal 1/5, tibiae and tarsi dark red to dark orange. Head (Fig. 5i, l). Large, globose to ovoid, 1.3 × longer than wide, 1.8 × longer than high, about as long as pronotum, with sparse short tubercles and sparse dark setae. Vertex slightly convex, frontal area gently elevated. Frontal suture gently curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with 22–23 fully segmented antennomeres; from 13th onwards, each segment is further subsegmented into 3–5 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump absent. Thorax (Figs. 4a, b, 5i, l, n). Cylindrical in cross section, sparsely granulated, covered in short small black setae, bearing lateral carinae. Pronotum subrectangular; sulci shallow, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior two thirds, and a further row of smaller round granules at anterior three fourths at each lateral margin. Metanotum slightly shorter than median segment. Fore and hindwings diminute, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 4a, b). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae gently elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as following two or three tarsomeres combined. Area apicalis delimited by a slightly deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 4a, b, 5j, k, m, n, o). Sparsely tuberculated, covered in short small black setae. Terga I–IX wider than long, X as long as wide. Terga I–VII bearing conical projections pointing posteriad at posterior margin, each a single curved one medially and one in each lateral carinae, except for tergum I without lateral projections. Terga VIII and IX with a medial slight elevation. Tergum X with medial carina at posterior half, posterior margin somewhat straight and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat triangular, posteriorly widening. Subgenital plate wider at mid length, then tapering to a lanceolate posterior margin, reaching the base of the cerci. Epiproct surpassing tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci short, cylindrical, gently widened basally (Fig. 5j, m, o). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX (Fig. 5k). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin wider and with a wide incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 5k). Gonoplac incurved, gently widened basally, apex round, shorter than gonapophysis IX (Fig. 5k).

Measurements (MZUSP 2612): body (without cerci) 66.1, head 5.7, antennae 40.2, pronotum 5.5, mesonotum 12.2, metanotum 5.2, median segment 4.7, abdomen (excluding median segment, without cerci) 32.8, profemur 15.6, protibia 16.1, mesofemur 14.0, mesotibia 13.3, metafemur 18.8, metatibia 19.5.

Male. Similar to that of the female, except: color (Fig. 4c, d). Black longitudinal line reaching median segment, tergum II and fading in terga III–IV. Head (Fig. 5a, c, f). Narrower, eyes larger, frontal convexity v-shaped but still broad, scapus and pedicel relatively larger, antennae with around 21 antennomeres. Thorax (Figs. 4c, 5a-c). Narrower; weaker and less conical granules at mesonotum, metanotum slightly relatively shorter than median segment. Legs (Fig. 4c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 4c, d, 5b, d, e, g, h). Narrower, tergum II barely wider than long, III–IV as wide as long, V–VII longer than wide, VIII–X wider than long. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 10 irregularly sized teeth. Epiproct hidden dorsally. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and slightly curving towards anterior; bearing a large linear dextral process, slightly upcurving, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci longer, of uniform width, gently incurved. Vomer very large, wide, strongly tapering, terminal hook sharp, small, upwardly curved. Genitalia (Fig. 6). Dorsal sclerite with basal part significantly narrower than apical part; main process with deep right angled cleft near apex and with short round anterior projection; distal process long, narrow, with round slightly widened apex. Internal sclerite with main process with four short projections equally apart; accessory process with short, somewhat conical inner branch and widely round proximal part of outer branch and short, somewhat conical distal part of outer branch. Ventral sclerite compound, asymmetric with somewhat straight margin tapering towards the left side.

Measurements (MZUSP 2614): body (without cerci) 52.2, head 4.3, antennae 46.5, pronotum 3.9, mesonotum 9.0, metanotum 4.0, median segment 3.7, abdomen (excluding median segment, without cerci) 27.3, profemur 16.0, protibia 16.5, mesofemur 12.2, mesotibia 11.8, metafemur 17.3, metatibia 18.7.

Egg (Fig. 7). Dissected from the abdomen of a female. Irregularly dark brownish, micropylar plate with creamish edges. Capsule barrel-shaped, higher than wide, longer than wide in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule covered by net of irregular, moderately thick rugose ridges, which are irregular and not forming ridges on the operculum. Ridges further bearing minute mushroom-like structures also on the operculum (Fig. 7b: inset). Micropylar plate elliptical, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line short, less than half the length of the micropylar plate. Operculum with a small central irregular rugose hump. Measurements (N = 5): length 3.5–3.8, width 2.3–2.6, micropylar plate length 0.9–1.4, operculum maximum diameter 1.8–2.1.

Materials examined

Holotype, f#: ♀ Anisomorpha dentata Stål, 1875: Brazil, Santa Catharina [Santa Catarina state] leg. M. Puls, coll. Brunner von Wattenwyl, number 131 (NMW 131, examined, Fig. 3).

Additional material examined, 2♀ (MZUSP 2612, 2613), 1 ♂ (MZUSP 2614), 2♀ nymphs (MZUSP 2615, 2616), 5 eggs: Brazil, Santa Catarina, Florianópolis, Ilha de Santa Catarina, Morro das Aranhas, 27°28′07.9"S 48°22′51.5"W, 2021; 1♀ (MZUSP 2620): Brazil, Santa Catarina, Florianópolis, Ilha de Santa Catarina, UCAD, 27°31′50.5"S 48°30′39.0"W, 11.ii.2023, G. Gomes and J. Conrado.

Differential diagnosis. Differs from other Urucumania by the larger head in dorsal and lateral views (Fig. 5a, c, i, l), the wider than long median segment, the dark orange coloration and wider aspect (extending to the lateral margin) of the dorsal paramedian bands on the body, the color pattern on the legs with femora black with dark orange in basal portion and dark orange tibiae and tarsi (Fig. 4), and the aspect of the male genitalia (Fig. 6), with a short sclerite of the ventral lobe with straight posterior margin, tapering towards the left side and a dorsal sclerite with thin left process with a widened round apex and right process forming three round margins.

Distribution (Fig. 68). Only known from Santa Catarina Island, Santa Catarina, Brazil, in the Atlantic Forest.

Remarks

Redtenbacher (1906) provided a description of male for P. dentata, but we do not consider this description as valid for the species due to the overall similarity between different species of Urucumania, several of which could match with the description provided, added to the vague locality relatively distant from type locality.

Urucumania nigrovittata (Piza, 1939) comb. n.

(Figs. 9, 10, 11, 12, 13 and 14)

Fig. 9

Urucumania nigrovittata comb. n., holotype male from Salto Grande, São Paulo, Brazil, in dorsal (a) and lateral (b) views. Scale bars = 2 mm

Fig. 10

Urucumania nigrovittata comb. n., couple from Assis (male) and Echaporã (female), São Paulo, Brazil. a, b Female in dorsal (a) and lateral (b) views. c, d Male in dorsal (c) and lateral (d) views. Scale bars = 2 mm

Fig. 11

Urucumania nigrovittata comb. n., couple from Assis (male) and Echaporã (female), São Paulo, Brazil. a–h Male. i–n Female. a, c, e, i, k Head and pronotum in dorsal (a, i), lateral (c, k) and ventral (e) views. b, m Metathorax in dorsal view. d, f–h, j, l, n Terminalia in dorsal (d, j), left side (f, l), right side (h) and ventral (g, n) views. Scale bars = 1 mm

Fig. 12

Urucumania nigrovittata comb. n., phallic organ of male from Assis, São Paulo, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars = 1 mm

Fig. 13

Urucumania nigrovittata comb. n., eggs of female from Echaporã, São Paulo, Brazil. a Lateral view. b Dorsal view. c Anterior or opercular view. d Five eggs in different views. Scale bars = 0.5 mm

Fig. 14

Urucumania nigrovittata comb. n., live individuals from Assis (male) and Echaporã (female), São Paulo, Brazil, in situ. a, b Adult male. d Subadult male. c, e Adult female

Donusa nigrovittata Piza, 1939.

Bacunculus nigrovittata, Piza 1946 (citation list).

Neophasma nigrovittata, Conle and Hennemann 2002 (redescription); Zompro and Domenico 2005 (citation in catalog); Otte and Brock 2005 (citation in catalog).

Pseudophasma dentata, Araujo and Garraffoni 2012 (citation in checklist); Brock and Büscher 2022 (citation in catalog).

Redescription

Female. Color (Figs. 10a, b, 14c, e). Head and body orange to dark orange, with a black medial longitudinal line running along entire length of head and thorax, sparsely in median segment; abdomen with darker irregular stains. Head and body laterally black; antennae dark red to dark orange. Ventrally dark brown, subgenital plate dark beige posteriorly. Femora dark brown, darker apically, tibiae and tarsi dark brown to black, tibiae basally beige. Head (Fig. 11i, k). Large, ovoid, 1.2 × longer than wide, 1.5 × longer than high, about as long as pronotum, with sparse dark conical setae. Vertex slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, slightly more than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with at least 20 fully segmented antennomeres; from 14th onwards, each segment is further subsegmented into 2–4 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump small, discrete, in the same color as the rest of the segment, present on the first subsegment of the 15th segment. Thorax (Figs. 10a, b, 11i, k, m). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, with very few small granules except in the carinae. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior three fifths, and a further row of smaller round granules at anterior two thirds at each lateral margin. Metanotum very slightly shorter than median segment. Fore and hindwings diminute, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 10a, b). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as the following two or three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Fig. 10a, b, 11j, l, m, n). Sparsely tuberculated, covered in short small black setae. Terga I–IV, VIII and IX wider than long, V and X as long as wide, VI and VII longer than wide. Terga I–VIII bearing elliptical projections pointing posteriad at posterior margin, each a single curved one medially and one in each lateral carinae, except for tergum I without lateral projections; medial projection slightly away from margin in VIII. Tergum IX only with the medial projection, smaller than in other terga and slightly away from the margin. Terga III–VI with two paramedian carinae near the medial line, connected to the posterior projection; VII and VIII with two paramedian elevations similar to the carinae of anterior segments. Tergum X with medial carina at posterior half, posterior margin somewhat straight and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat triangular, posteriorly widening. Subgenital plate wider at mid length, then tapering to a lanceolate posterior margin, reaching half the length of the cerci and almost reaching the end of tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci very short, slightly thick, cylindrical (Fig. 11j, l, n). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX. Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin. Gonoplac incurved, gently widened basally, apex round, shorter than gonapophysis IX.

Measurements (MZUSP 1738): body (without cerci) 54.1, head 5.0, antennae at least 32.0, pronotum 4.3, mesonotum 10.0, metanotum 4.1, median segment 4.0, abdomen (excluding median segment, without cerci) 26.7, profemur 14.1, protibia 14.4, mesofemur 11.9, mesotibia 11.9, metafemur 16.6, metatibia 17.0.

Male. Similar to that of the female, except: color (Figs. 10c, d, 14a, b, d). Orange color thinner, present in bands bordered by black, and present in dorsal surface of terga I–VI, gradually narrowing and fading in posterior terga and fading away in VII. Black longitudinal line reaching median segment, and continuing punctuated with the black posterior projections of terga, terga I–VII and with the black elevation of terga VIII and IX. Body and rest of abdomen black. Femora and tibiae black except for orange stains basally on all tibiae, and dark orange apically on tibiae, tarsi dark orange. Head (Fig. 11a, c, f). Narrower, eyes larger, scapus and pedicel larger. Thorax (Figs. 10c, d, 11a, b, c, f). Narrower, mesepisternum bearing sparse round granules. Legs (Fig. 10c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 10c, d, 11b, d, e, g, h). Narrower, tergum II–VII longer than wide, VIII wider than long, IX–X as long as wide. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 8 teeth. Epiproct wide, projecting dorsally from posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and slightly curving towards anterior; bearing a large linear dextral process slightly narrowing towards posterior, slightly upcurving, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci longer, cylindrical, slightly constricted medially to post medially, gently incurved. Vomer very large, wide, strongly tapering, terminal hook small, acute but with blunt dorsoventrally flattened apex, upwardly curved. Genitalia (Fig. 12). Dorsal sclerite with basal part wide, narrower than apical part; main process somewhat elongate, with very short, gentle anterior projection; distal process slightly thickened, apex conical and pointing laterally. Internal sclerite with main process with four short projections, the two medial very close to each other; accessory process with round slightly projected inner branch and widely round proximal part of outer branch and conical, slightly projected distal part of outer branch. Ventral sclerite compound, asymmetric with somewhat round margin more projected to the right.

Measurements (MZUSP 2590): body (without cerci) 45.3, head 3.2, antennae at least 15.2, pronotum 3.3, mesonotum 7.8, metanotum 2.9, median segment 3.1, abdomen (excluding median segment, without cerci) 25.0, profemur 14.3, protibia 14.7, mesofemur 11.2, mesotibia 11.3, metafemur 15.6, metatibia 15.9.

Egg (Fig. 13). Irregularly greyish to brownish, micropylar plate with creamish edges. Capsule barrel-shaped, higher than wide, longer than wide in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule covered by net of irregular, moderately thick rugose ridges, which are irregular and barely forming ridges on the operculum. Ridges further bearing minute mushroom-like structures also on the operculum. Micropylar plate elliptical to ovate, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line moderately thick, more than half the length of the micropylar plate. Operculum with a central irregular rugose hump. Measurements (N = 10): length 2.8–3.5, width 1.8–2.2, micropylar plate length 1.0–1.3, operculum maximum diameter 1.5–1.9.

Materials examined

Holotype, m#: ♂ Donusa nigrovittata Piza, 1939. Holotypus, ♂ Donusa nigrovittata Piza, 1939 [Brazil, São Paulo, Salto Grande] (MELQ: ESALQENT 000384, examined, Fig. 9).

Additional material examined, 1f# (MZUSP 1738), 10 eggs: Brazil, São Paulo, Echaporã, 22°24′42.6"S 50°11′48.2"W. xii.2020. P. W. Engelking & G. Annunciato col.; 1 ♂ (MZUSP 2590): Brazil. São Paulo, Assis, 22°33′37.8"S 50°22′19.8"W. 1.xi.2021. P. W. Engelking & G. Annunciato col.

Differential diagnosis. Differs from remaining Urucumania by the lighter orange bands and the not too narrow nor too thick dorsal paramedian bands not reaching the lateral margins of the body, the color pattern on the legs with femora gradually turning from dark orange or red in basal portion to black at apex, tibiae apically with a light orange stain contrasting with black gradually turning dark orange or red towards apex, and orange to red tarsi (Fig. 10), and the aspect of the male genitalia (Fig. 12). From Urucumania dentata comb. n., it differs by the less robust body (Fig. 10), relatively smaller head (Fig. 11a, i), the longer, thinner and more curved cerci of the male (Fig. 11d, e, g, h), and shorter cerci of the female (Fig. 11j, l, n).

Distribution (Fig. 68). Only known from Salto Grande, Assis and Echaporã, São Paulo, Brazil, in areas of dry forests of the Cerrado.

Urucumania pirulai Ghirotto sp. n.

(Figs. 15, 16, 17 and 18)

Fig. 15

Urucumania pirulai sp. n., holotype female (a, b) and paratype male (c, d) from Parauapebas, Pará, Brazil, in dorsal (a, c) and lateral (b, d) views. Scale bars = 2 mm

Fig. 16

Urucumania pirulai sp. n., paratype male (a–h) and holotype female (i–n) from Parauapebas, Pará, Brazil. a, c, i, k Head and pronotum in dorsal (a, i) and lateral (c, k) views. b, m Metathorax in dorsal view. d–h, j, l, n Terminalia in dorsal (d, j), left side (f, l), right side (g), caudal (e) and ventral (h, n) views. Scale bars = 1 mm

Fig. 17

Urucumania pirulai sp. n., phallic organ of paratype male from Parauapebas, Pará, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 18

Urucumania pirulai sp. n., live insects from Parauapebas, Pará, Brazil, in situ. a–c Mating pair, male above. d Male in dorsal view. Photos courtesy of Nereston Camargo (a, d), Lourival Tyski (b), and Welligton da Mota Ferreira (c)

Description

Female. Color (Figs. 15, 16, 17, 18a-c). Head and body dark brown to black, with a black medial longitudinal line running dorsally along entire length of head and thorax, median segment, and terga II–VII; medial line flanked by dark yellow to dark beige bands running along the head, thorax and terga I–VII. Alae with yellow veins and edges, cells black. Head with further paramedian black, thin lines; scapus and pedicel brown, flagellomeres dark yellow. Ventrally irregularly dark brown to black with irregular beige stains. Femora black with short areas of dark yellow to brown basally and apically faint in the profemora; tibiae black with a basal short area of dark yellow to brown, tarsi blackish. Head (Fig. 16i, k). Large, ovoid, 1.3 × longer than wide, 1.8 × longer than high, about as long as pronotum, with sparse dark conical setae. Vertex very slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform, long, with around 28 fully segmented antennomeres; from 17th onwards, each segment is further subsegmented into 2–12 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump small, discrete, in the same color as the rest of the segment, present on the first subsegment of the 19th segment. Thorax (Figs. 15a, b, 16i, k, m). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, with scattered small granules. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal gently curved; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior half, and a further row of smaller round granules at anterior two thirds at each lateral margin. Metanotum slightly shorter than median segment. Fore and hindwings diminute, scale-like, elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 15a, b). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi as long as to slightly longer than following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 15a, b, 16j, l, m, n). Sparsely tuberculated, covered in short small black setae. All terga wider than long. Terga I–VIII bearing round projections pointing posteriad at posterior margin, each a single one medially and smaller ones in each lateral carinae, except for terga I and VIII without lateral projections; medial projection slightly away from margin in VIII. Tergum IX only with the medial projection, smaller than in other terga. Terga III–IX with two elevated paramedian carinae near the medial line in the posterior region, connected to the posterior projection. Tergum X with medial carina at posterior one third, posterior margin slightly emarginated forming short round lobes, and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, posteriorly widening. Subgenital plate wider at mid length, then tapering to a roundly lanceolate posterior margin, reaching the base of the cerci and reaching three fourths of tergum X. Epiproct visible dorsally, surpassing tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci 2.1 × longer than wide, short, slightly thick, cylindrical (Fig. 16j, n). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX. Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin. Gonoplac incurved, narrowing towards posterior, apex acute and slightly upcurving, slightly longer than gonapophysis IX.

Measurements (MZUSP 2575): body (without cerci) 64.2, head 5.5, antennae 43.8, pronotum 5.6, mesonotum 11.4, metanotum 3.7, median segment 4.7, abdomen (excluding median segment, without cerci) 33.3, profemur 16.7, protibia 17.3, mesofemur 13.8, mesotibia 13.2, metafemur 19.2, metatibia 19.8.

Male. Similar to that of the female, except: color (Figs. 15c, d, 18). Head with very thin dorsolateral lines, mostly just black laterally; antennae lighter; tibiae apically also yellowish, tarsi lighter. Head (Fig. 16a, c). Narrower, 1.4 × longer than wide, eyes larger more than hemispherical; antennae with around 25 segments, subsegments starting at 14th segment, scapus and pedicel larger. Thorax (Figs. 15c, d, 16a-c). Narrower, slightly more elongate. Mesonotum bearing on each side a paramedian row of conical granules on anterior two thirds, and a further row of smaller round granules at anterior three fourths at each lateral margin; mesepisternum bearing carinae and sparse row of round granules. Legs (Fig. 15c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 15c, d, 16b, d-h). Narrower, terga I–VII longer than wide, VIII–X wider than long; carinae present in most of the tergum’s length. Posterior margin of tergum X gently emarginated, thorn pads pointing inward, near median line, bearing around 8 teeth. Epiproct wide, not projecting dorsally, under tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin straight, gently widened laterally; bearing a large dextral process with a broad base bearing a round projection ventrally and an upcurving linear projection dorsally; dextral process gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci longer, 2.7–2.8 × longer than wide, cylindrical, very slightly constricted medially to post medially, posterior half slightly narrower, gently incurved (Fig. 16d, f, h). Vomer very large, wide, strongly tapering, terminal hook small, acute but with blunt dorsoventrally flattened apex, upwardly curved, slightly tilted to the right side. Genitalia (Fig. 17). Dorsal sclerite with basal part narrower than apical part; main process elongated and conical with roundly lanceolate apex; distal process long, narrow with a widened base, with round slightly widened apex pointing laterally. Internal sclerite with main process with four short projections, the medial ones slightly closer to each other; accessory process with a round, convex edge and with short, conical inner branch and widely elliptical proximal part of outer branch and protruding, irregular, somewhat conical distal part of outer branch. Ventral sclerite compound, asymmetric, somewhat elongate, with round margin pointing to the right.

Measurements (MZUSP 2576): body (without cerci) 48.7, head 3.3, antennae 55.0, pronotum 3.6, mesonotum 8.7, metanotum 2.3, median segment 4.1, abdomen (excluding median segment, without cerci) 26.6, profemur 17, protibia 17.5, mesofemur 12.1, mesotibia 13.0, metafemur 18.0, metatibia 19.9.

Measurements, variation (N = 1, MZUSP 2577): body (without cerci) 49.9, head 3.7, pronotum 3.6, mesonotum 9.3.

Egg. unknown.

Materials examined

Holotype, f#: Brazil, Pará, Parauapebas, arredores da FLONA do Tapirapé-Aquiri, 5°57′08.8"S 50°31′25.0"W, 15.iii.2021, T. S. Soares col., BP 12 (MZUSP 2575).

Paratypes, 2 m# (MZUSP 2576, 2577): Brazil, Pará, Parauapebas, arredores da FLONA do Tapirapé-Aquiri, 5°57′08.8"S 50°31′25.0"W, 15.iii.2021, T. S. Soares col., BP 12.

Etymology. The species is named after Dr. Paulo Miranda Nascimento, nicknamed and mostly known as Pirula or Pirulla, a Brazilian paleontologist and science communicator. Acting mostly via YouTube videos in his channel ''Canal do Pirulla'', Pirula is regarded by us as one of the most important science communicators of Brazil, especially, but not only, in the area of biological sciences, with proficient treatment of scientific and other matters. Pirula was a strong reference as a scientist for VMG in his early career. This homage also makes sense because despite YouTube system favoring only short videos to be more discoverable, Pirula achieved popularity even with his long videos, unusual among youtubers, coinciding with phasmids being unusually long insects. After the dark times that recently shadowed science in Brazil, popularizing the importance of science is vital.

Differential diagnosis. Differs from remaining Urucumania by the dark-orange color pattern with bright yellow wings and black legs with basal and apical area of femora and basal area of tibiae yellowish (Fig. 15), and the shortened tergum X in both sexes (Fig. 16d, j), and the basally widened right process of sternum IX of males (Fig. 16g). The overall coloration pattern resembles U. varellai sp. n., but the female of U. pirulai sp. n. can be distinguished by the other mentioned characteristics.

Distribution (Fig. 68). Only known from Carajás Mountains in Parauapebas, Pará, Brazil, in the Amazon.

Urucumania varellai Ghirotto & Chiquetto-Machado sp. n.

(Figs. 19, 20)

Fig. 19

Urucumania varellai sp. n., holotype female from Abel Figueiredo, Pará, Brazil, in dorsal (a) and lateral (b) views. Scale bars = 2 mm

Fig. 20

Urucumania varellai sp. n., holotype female from Abel Figueiredo, Pará, Brazil. a, c Head and pronotum in dorsal (a) and lateral (c) views. b Metathorax in dorsal view. d–f Terminalia in dorsal (d), lateral (e), and ventral (f) views. Scale bars = 1 mm

Description

Female. Color (Fig. 19). Known only from the dry preserved holotype. Body mostly dark brown, with light brown punctuations denser on thorax and abdominal terga; dorsally with a black medial longitudinal line, more distinct from head to median segment and progressively fainter towards posterior region of abdomen; dorsal surface of head, abdominal tergum X and posterior half of subgenital plate light brown. Antennae light brown. Fore and hindwings dark brown with reticulate light beige venation. Femora dark brown with basal and apical lighter regions; tibiae and tarsi light brown. Head (Fig. 20a, c). Smooth, somewhat ovoid, with vertex and genae gently convex, about as long as pronotum, covered with short dark setae and with short sparse tubercles, 1.2 × longer than wide, 1.6 × longer than high. Eyes slightly elongate; relatively small, covering less than one quarter of head length. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform, reaching tergum VII, entirely covered with small setae; scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus; with around 22 segments. Almost all antennomeres fully segmented, segments after basal portion long, only the last two segments further divided into 2–3 less conspicuous segments. Thorax (Figs. 19, 20a, b, c). Cylindrical in cross section; covered with short dark setae and with significant quantity of granules, more densely on meso- and metathorax. Prothorax about as long and wide as head. Pronotum with slight constriction on anterior third; anterolateral corners with rounded indentations, outlining openings of paired defensive glands; posterior margin convex; pair of gentle dorsolateral carinae originating posterior to defensive glands and extending until nearly posterior margin. Mesothorax slightly rugose, 2.2 × longer than prothorax; about as wide as prothorax on anterior region and gradually widening towards posterior region. Mesonotum bearing on each side a paramedian row of conical granules on anterior two thirds, and a further row of smaller round granules at anterior three fourths at each lateral margin. Metanotum slightly rugose, about 0.4 × the length of mesothorax and slightly longer than median segment; as wide as the posterior region of mesothorax. Meso- and metanotum with a pair of longitudinal carina extending along each lateral margin; mesepisternum and metepisternum with a longitudinal carina extending along ventral margin. Fore and hindwings very small, scale-like, roughly elliptical, the anterior larger than the posterior. Legs (Fig. 19). Hindlegs the longest; midlegs the shortest. All five carinae elevated and marked by rows of short setae. Profemur curved and compressed basally. Meso- and metafemur slightly thinner basally. Each tibia about as long as corresponding femur and 2.0 × longer than corresponding tarsus. Basitarsi about as long as or slightly longer as following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 19, 20b, d-f). Dorsally slightly rugose, covered with significant quantity of granules; ventrally smooth. Terga I–VII each with a pair of longitudinal carina extending along lateral margin and with a subtriangular projection near posterior margin, pointing posteriorly. Median segment slightly shorter than metanotum and 1.2 × longer than segment II. Abdomen excluding median segment approximately as long as combined length of head, thorax and median segment. Segments II–IV similar in length; segments V–VI slightly longer than II–IV; segment VII slightly shorter than VI. Terga VIII–X slightly shorter and narrower than VII. Tergum VIII with a gentle medial constriction in dorsal view; bearing a small bump near posterior margin. Tergum IX slightly longer than wide, gently narrowing towards posterior region. Tergum X tectiform, in dorsal view about as long as wide; posterior margin approximately straight; posterolateral corners with large rounded indentations outlining cerci. Sternum VII with conspicuous preopercular organ, developed into an elongate, shiny dark protuberance with blunt rounded apex surpassing posterior margin of sternum VII. Subgenital plate lanceolate, almost reaching posterior margin of tergum X; apex somewhat sharp. Epiproct under tergum X, slightly wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci very short, somewhat thick (2.4 × longer than wide), conical, slightly tapering, very slightly upcurved; not surpassing posterior margin of tergum X; apex blunt (Fig. 20d-f).

Measurements (MZUSP 0215): body (without cerci) 60.21, head 5.27, antennae at least 40.96, pronotum 4.22, mesonotum 11.53, metanotum 4.36, median segment 4.6, abdomen (excluding median segment, without cerci) 30.23, profemur 17.26, protibia 11.9 (damaged), mesofemur 13.07, mesotibia 12.88, metafemur 17.67, metatibia 19.72.

Male. unknown.

Egg. unknown.

Materials examined

Holotype, f#: Brazil, Pará, Abel Figueiredo, Juca Marhe Farm, 04º51′45.5″S, 48º32′48.5″W, 28.i–5.ii.2010, manual collection (MZUSP 0215).

Etymology. This Amazonian species is named after Drauzio Varella, a Brazilian oncologist, infectious disease specialist, scientist, educator and best-selling author. Varella was a pioneer in the treatment of AIDS in Brazil and since the 1980s has had an important role in educational campaigns about this syndrome. During the past three decades he has also led a prospective research program on the medicinal potential of Brazilian Amazon plants and has been an active and successful science communicator in the radio, television and Internet.

Differential diagnosis. Females (the only known sex) differ from remaining Urucumania by the dark-orange color pattern with bright yellow wings, beige tibiae and tarsi, femora black with beige at the extremities (Fig. 19), the widened and round head (Fig. 20a), the slightly wider than long tergum X (Fig. 20d, e), and relatively short (around 3 × longer than wide) cerci (Fig. 20f). The overall coloration pattern resembles U. pirulai sp. n., but the female of U. varellai sp. n. can be distinguished by the other mentioned characteristics.

Distribution (Fig. 68). Only known from Abel Figueiredo, Pará, Brazil, in the Amazon.

Urucumania guadanuccii Ghirotto sp. n.

(Figs. 21, 22, 23, 24, 25 and 26)

Fig. 21

Urucumania guadanuccii sp. n., holotype female (a, b) and paratype male (c, d) from Diamantina, Minas Gerais, Brazil, in dorsal (a, c) and lateral (b, d) views. Scale bars = 2 mm

Fig. 22

Urucumania guadanuccii sp. n., paratype male (a–h) and holotype female (i–o) from Diamantina, Minas Gerais, Brazil. a, c, f, i, l Head and pronotum in dorsal (a, i), lateral (c, l) and ventral (f) views. b, n Metathorax in dorsal view. d, e, g, h, j, m, o Terminalia in dorsal (d, j), left side (g, m), right side (h) and ventral (e, o) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 23

Urucumania guadanuccii sp. n., phallic organ of paratype male from Diamantina, Minas Gerais, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 24

Urucumania guadanuccii sp. n., egg of holotype female from Diamantina, Minas Gerais, Brazil. a Lateral view. b Dorsal view. c Anterior or opercular view. Scale bars = 0.5 mm

Fig. 25

Urucumania guadanuccii sp. n., live individuals raised in captivity from female from Diamantina, Minas Gerais, Brazil. a, b Male and female paired in dorsal (a) and lateral (b) views. c, d First instar in ventral (c) and dorsal (d) views. Scale bars: a, b 2 mm; c, d 1 mm

Fig. 26

Urucumania guadanuccii sp. n., holotype female in situ (a) and paratype male in captivity (b) from Diamantina, Minas Gerais, Brazil

Description

Female. Color (Figs. 21a, b, 25a, b, 26a). Head and body black with scattered whitish granules varying in density, denser on the abdomen, legs black with orangish beige stains on the basis of metafemora. Head and body with orange paramedian bands interrupted by a black medial longitudinal line, running along the head, pronotum, mesonotum, metanotum, median segment, tergum II and slightly in tergum III, blurred on the head and in the posterior region of tergum II and almost indistinctly present at tergum III. Each orange band breadth reaches around mid-width of each half of the thorax. Subgenital plate dark orange from pre-medial region to the posterior end, with black blurred longitudinal stripes. Head (Fig. 22i, l). Large, approximately globose, 1.1 × longer than wide, 1.4 × longer than high, slightly shorter than pronotum, sparsely tuberculated. Vertex flat, frontal area gently elevated. Frontal suture gently curved and deep, frontal convexity prominent. Eye somewhat large, round, more than hemispherical. Clypeus wide, convex, with shallow depressions, labrum slightly notched anteromedially, round. Antennae filiform with 14 fully segmented antennomeres; from 15th onwards, each segment is further subsegmented into 3–5 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump absent. Thorax (Figs. 21a, b, 22i, l, n). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, with sparsely small granules. Pronotum subrectangular, tapering towards anterior; sulci marked, longitudinal straight and transversal slightly sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Pronotum with granules mainly near lateral edges. Mesonotum bearing on each side a paramedian row of conical to cylindrical granules on anterior three fifths and some smaller granules posteriorly, and a further row of smaller round granules at anterior three fifths at each lateral margin. Metanotum very slightly shorter than median segment. Metanotum bearing two laterodorsal granules at each side and a medial one posteriorly. Mesepisternum and metepisternum also bearing row of granules. Fore and hindwings diminute, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 21a, b). Hindlegs surpassing abdomen and slightly longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as or slightly longer than the following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 21a, b, 22j, k, m–o). Sparsely tuberculated, with scattered larger granules, covered in short small black setae. Terga I–X wider than long. Terga I–VIII bearing short round projections pointing posteriad, one medially and two paramedial at posterior margin, and one in each lateral carinae except for tergum I without lateral projections. Terga IX–X with scattered paramedial granules. Tergum X with medial carina at posterior half, posterior margin somewhat widely round and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat triangular, posteriorly widening. Subgenital plate wider at mid length, then slightly tapering to a round posterior margin, almost reaching the end of tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci relatively elongate (3.5 × longer than wide), cylindrical, posteriorly slightly tapering (Fig. 22i, o). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX (Fig. 22k). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 22k). Gonoplac incurved, gently widened basally, apex round, as long as gonapophysis IX (Fig. 22k).

Measurements (MZUSP 2592): body (without cerci) 52.97, head 3.94, antennae at least 25.62, pronotum 4.4, mesonotum 9.95, metanotum 2.93, median segment 3.7, abdomen (excluding median segment, without cerci) 28.05, profemur 12.0, protibia 12.66, mesofemur 10.1, mesotibia 10.1, metafemur 13.8, metatibia 14.26.

Measurements, variation (N = 5, MZUSP 1746–1750): body (without cerci) 41.3–45.7, head 3.4–4.1, pronotum 3.7–4.2, mesonotum 8.1–9.0.

Male. Similar to that of the female, except: color (Figs. 21c, d, 25a, b, 26b). Basal half of femora dark orange, tarsi dark orange. Head (Fig. 22a, c, f). Narrower, eyes larger, scapus and pedicel larger. Thorax (Figs. 21c, d, 22a-c, f). Narrower, granules of mesonotum and metanotum more prominent, conical. Legs (Fig. 21c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 21c, d, 22b, d, e, g, h). Narrower, tergum II–III, VIII, IX wider than long, IV–VII longer than wide, X as long as wide. Posterior margin of tergum X very gently emarginated, thorn pads pointing inward, near median line, bearing around 10 teeth. Epiproct not exceeding posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin almost straight, slightly curving towards anterior; bearing a large dextral process widened at basal two thirds then somewhat linear and slightly narrowing towards posterior, slightly upcurving, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci long (4.3 × longer than wide), cylindrical, gently incurved (Fig. 22d, g, h). Vomer very large, wide, strongly tapering, terminal hook small, acute, upwardly curved. Genitalia (Fig. 23). Dorsal sclerite with basal part narrower than apical part; main process with deep right angled cleft, slightly tapering towards a round apex, with round projection near apex and with round anterior projection before the cleft; distal process moderately thick, with round slightly widened apex gently bent laterally. Internal sclerite with main process with four short projections equally apart; accessory process with elongate, somewhat cylindrical inner branch and widely round proximal part of outer branch and very short, slightly prominent round distal part of outer branch. Ventral sclerite compound, slightly asymmetric with round margin extending to the left side.

Measurements (MZUSP 2591): body (without cerci) 34.6, head 2.88, antennae 34.82, pronotum 2.81, mesonotum 6.75, metanotum 1.92, median segment 2.48, abdomen (excluding median segment, without cerci) 17.76, profemur 10.73, protibia 11.38, mesofemur 7.91, mesotibia 8.82, metafemur 11.91, metatibia 12.64.

Measurements, variation (N = 2, MZUSP 1744, 1745): body (without cerci) 31.8–32.3, head 2.5–2.8, pronotum 2.4, mesonotum 6.2–6.5.

Egg (Fig. 24). Irregularly dark blueish gray, micropylar plate creamish. Capsule barrel-shaped, higher than wide, longer than wide in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule covered by net of irregular, moderately thick rugose ridges, which are irregular and not forming ridges on the operculum. Ridges further bearing minute mushroom-like structures also on the operculum. Micropylar plate elliptical to ovate, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line moderately thick, slightly less than half the length of the micropylar plate. Operculum with a central irregular rugose hump. Measurements (N = 10): length 2.6–3.0, width 1.7–2.1, micropylar plate length 0.8–1.1, operculum maximum diameter 1.4–1.7.

First instar (Fig. 25c, d). Brownish beige to dark, tibiae and metathorax lighter, sprinkled with small white stains, ventrally with a medial creamish line on head through sternum II. Body setose, head with moderately large eyes, antennae with 9 clear segments, legs with developed keels, cerci cylindrical.

Materials examined

Holotype, 1f#: Brazil, Minas Gerais, Diamantina, campus of Universidade Federal dos Vales do Jequitinhonha e Mucuri—UFVJM, 18°11′39.6"S 43°34′18.5"W. Cerrado de campos rupestres. 25.x.2018. V. M. Ghirotto, R.F. Ferreira and A. Galleti-Lima col. (MZUSP 2592).

Paratypes, 3 m# (MZUSP 2591, 1744, 1745), 5f# (MZUSP 1746–1750), 25 eggs: Brazil, Minas Gerais, Diamantina. Reared in captivity by V. M. Ghirotto from female 2592 [holotype], died 2.iv–1.vii.2019.

Etymology. The species is named after Dr. José Paulo Leite Guadanucci, an expert Brazilian arachnologist and skillful researcher. He was a professor and researcher in UFVJM at Diamantina, the type locality of the new species, before moving to Universidade Estadual ''Júlio de Mesquita Filho'' (UNESP). He advised VMG in his undergraduate course in biology at UNESP, working with mygalomorph spiders. Besides, he supported VMG in his first practices regarding stick insects that eventually led to the foundation of Projeto Phasma—a freestanding phasmid research group of which some of the authors, VMG, PWE, PABAN, EBC and PICM, are part of. The early start in researching phasmids by us and early foundation of the Projeto Phasma were made possible by Dr. Guadanucci's assistance. This comes in the form of our sincerely gratefulness for kindly helping us and supporting research on Phasmatodea in Brazil.

Differential diagnosis. Differs from remaining Urucumania by the dark-orange color pattern with dark legs (Fig. 21), the elongate terga VIII–X in both sexes (Fig. 22e, j), the widened dextral process of sternum IX of males (Fig. 22h), and elongated cerci of the female (Fig. 22j). Females further differ from Urucumania tapirape sp. n., another species with elongated terga VIII–X, by the shorter head (Fig. 22a, i) and terga VIII–X (Fig. 22e, j) and distinct color pattern (Fig. 21).

Distribution (Fig. 68). Only known from Diamantina, Minas Gerais, Brazil, in areas of campos rupestres, a rocky savannic Cerrado in the Espinhaço formation.

Urucumania atilai Ghirotto sp. n.

(Figs. 27, 28, 29, 30 and 31)

Fig. 27

Urucumania atilai sp. n., holotype female (a, b) and paratype male (c, d) from São Miguel do Arcanjo, São Paulo, Brazil, in dorsal (a, c) and lateral (b, d) views. Scale bars = 2 mm

Fig. 28

Urucumania atilai sp. n., paratype male (a–h) and paratype female (i–o) from São Miguel do Arcanjo, São Paulo, Brazil. a, c, g, i, l Head and pronotum in dorsal (a, i), lateral (c, l) and ventral (g) views. b, n Metathorax in dorsal view. d, e, f, h, j, m, o Terminalia in dorsal (d, j), left side (e, m), right side (h) and ventral (f, o) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 29

Urucumania atilai sp. n., phallic organ of paratype male from São Miguel do Arcanjo, São Paulo, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 30

Urucumania atilai sp. n., eggs of paratype female from São Miguel do Arcanjo, São Paulo, Brazil. a Lateral view. b Dorsal view. c Several eggs in lateral and dorsal views. Scale bars = 0.5 mm

Fig. 31

Urucumania atilai sp. n., live females from São Miguel do Arcanjo, São Paulo, Brazil, in situ. Photos courtesy of Thales de Válio

Description

Female. Color (Figs. 27a, b, 31). Head and body dorsally orangish brown to dark orange, abdomen gradually darkening in the last terga, with VIII and IX black, X lighter; laterally blackish. With a black medial longitudinal line running along entire length of head, thorax and median segment. Head darkening towards anterior; antennae yellowish to reddish brown. Ventrally irregularly beige or brown, subgenital plate lighter. Femora black with dark red to dark orange at basal one fourth, tibiae and tarsi dark red to dark orange; in some individuals entire leg slightly lighter, with irregular, small lighter stains. Head (Fig. 28i, l). Large, subquadrate, 1.1–1.2 × longer than wide, 1.9 × longer than high, about as long as pronotum, with sparse dark conical setae, scattered with moderate quantity of whitish short tubercles. Vertex slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, slightly more than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with around 22–23 fully segmented antennomeres; from 18th onwards, some segments are further subsegmented into 2–4 less articulated subsegments of equal width, with the two apicalmost subsegmented into 6–7; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump absent. Thorax (Figs. 27a, b, 28i, l, n). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, scattered with whitish short tubercles. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior two thirds, and a further row of smaller round granules at anterior three fourths at each lateral margin. Metanotum very slightly shorter than median segment. Fore and hindwings diminute, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 27a, b). Hindlegs surpassing abdomen and slightly longer than forelegs, midlegs slightly shorter than forelegs. All five carinae conspicuously elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as the following two to three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 27a, b, 28j, k, m–o). Scattered with few to moderate whitish short tubercles, covered in short small black setae. Terga I–X wider than long, X about as long as wide. Terga I–VII bearing elliptical projections and VIII shorter projections, all those pointing posteriad and at posterior margin; each a single curved one medially and one shorter in each lateral carinae, except for tergum I without lateral projections. Tergum IX only with the medial projection, shorter and less conspicuous than in other. Terga III–VI with two very weak paramedian carinae near the medial line, connected to the posterior projection; VII and VIII with two paramedian elevations similar to the carinae of anterior segments. Tergum X with medial carina at posterior half, posterior margin somewhat straight, very gently curved and gently carinated, slightly widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat triangular, posteriorly widening. Subgenital plate conspicuously convex, much widened medially with lateral margins round, then tapering to a round to gently acute posterior margin, reaching half the length of tergum X or slightly more. Epiproct slightly surpassing tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci short, moderately thick (around 2.2–2.4 × longer than wide), cylindrical (Fig. 28j, m, o). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX (Fig. 28k). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 28k). Gonoplac slightly incurved, gently widened basally, apex round, as long as or slightly longer than gonapophysis IX (Fig. 28k).

Measurements (MZUSP 2589): body (without cerci) 50.9, head 4.2, antennae 35, pronotum 4.63, mesonotum 9.4, metanotum 4.81, median segment 3.81, abdomen (excluding median segment, without cerci) 24.05, profemur 13.46, protibia 12.85, mesofemur 11.06, mesotibia 10.68, metafemur 15.18, metatibia 15.55.

Measurements, variation (N = 4, MZUSP 1743, 2005–2007): body (without cerci) 37.4–50.0, head 4.1–4.3, pronotum 3.5–4.4, mesonotum 7.0–9.5.

Male. Similar to that of the female, except: color (Fig. 27c, d). Dorsally the reddish color is lighter. The lighter coloration of the female subgenital plate is reflected in the poculum of the male. Head (Fig. 28a, c, g). Narrower, eyes larger, scapus and pedicel larger. Thorax (Figs. 27c, d, 28a-c, g). Narrower. Legs (Fig. 27c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 27c, d, 28b, d-f, h). Narrower, tergum I, III, IV, VII as long as wide, II, VIII, IX slightly wider than long, V, VI slightly longer than wide, X as long as wide or slightly wider than long. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 8–12 teeth. Epiproct hidden under tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and slightly curving towards anterior; bearing a large linear dextral process, slightly upcurving, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci long, cylindrical, somewhat thick (3.4 × longer than wide) slightly constricted medially to post medially, gently incurved (Fig. 28d-f, h). Vomer very large, wide, strongly tapering, terminal hook small, acute but with blunt dorsoventrally flattened apex, upwardly curved. Genitalia (Fig. 29). Dorsal sclerite with basal part significantly narrower than apical part; main process somewhat narrow and elongate, with gentle posterior cleft laterally and with short round anterior projection; distal process long, narrow, with round slightly widened apex. Internal sclerite with main process with four short projections equally apart; accessory process with short, round inner branch and widely round proximal part of outer branch and distal part inconspicuous. Ventral sclerite compound, asymmetric with somewhat round margin wider at the middle, slightly sinuous.

Measurements (MZUSP 2516): body (without cerci) 36.1, head 2.7, antennae 29.2, pronotum 2.4, mesonotum 6.2, metanotum 2.7, median segment 2.9, abdomen (excluding median segment, without cerci) 19.2, profemur 9.8, protibia 10.3, mesofemur 8.3, mesotibia 8.5, metafemur 11.7, metatibia 12.2.

Measurements, variation (N = 7, MZUSP 2517, 2518, 2674, 2675, 2557, 2558, 2588): body (without cerci) 32.6–39.4, head 2.6–3.0, pronotum 2.2–2.9, mesonotum 5.1–6.7.

Egg (Fig. 30). Irregularly greyish to brownish, micropylar plate beige to grayish. Capsule barrel-shaped, higher than wide, considerably longer than wide but in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule covered by net of irregular, moderately thick rugose ridges, which are irregular and barely forming ridges on the operculum. Ridges further bearing minute mushroom-like structures also on the operculum. Micropylar plate elliptical to strongly ovate, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line moderately thick, less than about the same length of half the length of the micropylar plate. Operculum with a central irregular rugose hump. Measurements (N = 10): length 2.5–2.7, width 1.4–1.7, micropylar plate length 0.8–1.1, operculum maximum diameter 1.2–1.4.

Materials examined

Holotype, 1 m#: Brazil, São Paulo, São Miguel Arcanjo, Vila Rica, 23°53′11.9"S 47°59′37.6"W. (MZUSP 2516).

Paratypes, 1f# (MZUSP 2589), 5 m# (MZUSP 2517, 2518, 2674, 2675, 2588), 30 eggs: Brasil, São Paulo, São Miguel Arcanjo, Vila Rica, 23°53′11.9"S 47°59′37.6"W; 3f# (MZUSP 2005–2007), 2 m# (MZUSP 2008, 2009), 12 eggs: raised in captivity from female from Brazil, São Paulo, São Miguel Arcanjo, by E. B. Crispino, died vii.2022; 1f#: Brazil, São Paulo, Iperó, área de floresta antropizada, FLONA Ipanema, 10.x.2019, A. Galleti-Lima, J.P.L. Guadanucci and R.P. Indicatti col. (MZUSP 1743); 1f#: Brazil, São Paulo, Iperó, FLONA Ipanema, collected 15.x.2018 as nymph, reared in captivity and matured 01.xi.2019, R.P. Indicatti col. (MZUSP 2040); 3 m (MZUSP 2550, 2557, 2558): raised in captivity from female from Brazil, São Paulo, São Miguel Arcanjo, by V. M. Ghirotto, died xii.2022.

Etymology. The species is named after Dr. Atila Iamarino, a Brazilian microbiologist best known for his significant science communication contribution spanning several fields of knowledge, mostly through the YouTube channels “Nerdologia” and “Atila Iamarino”. He was also a guiding voice to Brazil amidst the recent COVID-19 pandemic, broadcasting trustful, scientific and public health information.

Differential diagnosis. Differs from remaining Urucumania by the lighter, reddish to beigeish-orange coloration and the narrow width aspect of the dorsal paramedian bands on the body (not reaching the lateral margins), by the color pattern on the legs with dark tibiae and femora but femora basally reddish to orangish (Fig. 27), and by the aspect of the male genitalia (Fig. 29). Both sexes are also distinguished by the body and head proportion (Fig. 28a, i), males can be further distinguished by the short, thickened cerci (Fig. 28d, e), and females by the medium sized cerci (Fig. 28j, m, o).

Distribution (Fig. 68). Only known from Iperó and São Miguel Arcanjo, São Paulo, Brazil, in areas of inland Atlantic Forest.

Urucumania intervalica Ghirotto & Crispino sp. n.

(Figs. 32, 33, 34, 35 and 36)

Fig. 32

Urucumania intervalica sp. n., holotype female (a, b) and paratype male (c, d) from Ribeirão Grande and paratype male (e) from Tapiraí, both in São Paulo, Brazil, in dorsal (a, c, e) and lateral (b, d) views. Scale bars = 2 mm

Fig. 33

Urucumania intervalica sp. n., paratype male (a–h) and holotype female (i–o) from Ribeirão Grande, São Paulo, Brazil. a, c, g, i, l Head and pronotum in dorsal (a, i), lateral (c, l) and ventral (g) views. b, n Metathorax in dorsal view. d, e, f, h, j, m, o Terminalia in dorsal (d, j), left side (e, m), right side (h) and ventral (f, o) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 34

Urucumania intervalica sp. n., phallic organ of paratype male from from Ribeirão Grande, São Paulo, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 35

Urucumania intervalica sp. n., eggs of paratype female from Ribeirão Grande, São Paulo, Brazil. a Lateral view. b Dorsal view, inlet showing mushroom-like structures. c Anterior or opercular view. d Several eggs in lateral and dorsal views. Scale bars = 1 mm

Fig. 36

Urucumania intervalica sp. n., live insects from Ribeirão Grande, São Paulo, Brazi, in situ. a Dark-colored female, photo courtesy of Ericson Cernawsky Igual. b Light-colored female, photo courtesy of Fabrice Schmitt. c Dark-colored male, photo courtesy of Luis Felipe Teixeira

Description

Female. Color (Figs. 32a, b, 36a, b). Head and body dark beige, brown or dark orange, usually with a median dorsal black line on the head; laterally usually darker. Head with paramedian dorsolateral black stripes. Coxae dark. Femora dark beige, apically black, irregularly mottled in lighter stains. Tibiae basally beige and gradually darkening to brown or dark beige; tarsi brown or dark beige. Head (Fig. 33i, l). Large, subquadrate, 1.2–1.3 × longer than wide, 1.5 × longer than high, slightly shorter than pronotum, with sparse dark conical setae. Vertex flat to very slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with 22–24 fully segmented antennomeres; from 15th onwards, each segment is further subsegmented into 2–9 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal absent. Thorax (Figs. 32a, b, 33i, l, n). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, with very few small granules except in the carinae, where they are more prominent and symmetrical. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina and two posterior granules; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of 4–5 conical granules on anterior three fifths; a further row of smaller round granules at anterior two thirds at each lateral margin and a faint row of small granules located almost dorsally. Metanotum slightly shorter than median segment bearing an inconspicuous median spike. Fore and hindwings are very small, scale-like; forewing slightly elliptical, robust; hindwing elliptical, scale-like. Meso and metafurca each separated longitudinally, curved medially. Legs (Fig. 32a, b). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as the same size than following two or three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 32a, b, 33j, k, m–o). Sparsely tuberculated, covered in short small black setae. Terga I–IV, VIII and IX wider than long, V and X as long as wide, VI and VII longer than wide. Terga I–VIII bearing elliptical projections pointing posteriorly at posterior margin, each a single curved one medially and one in each lateral carinae, except for tergum I without lateral projections; medial projection slightly away from margin in VIII, wider and less pointed. Tergum IX only with the medial projection, similar to the one in VIII but slightly more conspicuous and positioned at the margin. Terga III–VI with two paramedian carinae near the medial line, connected to the posterior projection; VII and VI with two paramedian elevations similar to the carinae of anterior segments, also present in the anterior half of VII. Tergum X with medial carina at posterior half, posterior margin with three projections facing away from the body, a somewhat inconspicuous median one and two prominent lateromedian pair. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat trapezoid, posteriorly widening, posterior margin with an indentation facing anteriorly. Subgenital plate wider at mid length, then tapering to a lanceolate posterior margin, reaching two thirds the length of the cerci and reaching the end of tergum X. Epiproct under tergum X, wide, somewhat triangular with a medial carina. Paraprocta apart at the very most apical part, inner margins long, beginning at the posterior end of tergum IX. Cerci short, slightly thick, cylindrical, tapering apically (Fig. 33j, m, o). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX (Fig. 33k). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 33k). Gonoplac incurved, gently widened basally, apex round, shorter than gonapophysis IX (Fig. 33k).

Measurements (MZUSP 1238): body (without cerci) 60.2, head 5.1, antennae at least 35.0, pronotum 5.5, mesonotum 10.8, metanotum 3.6, median segment 4.5, abdomen (excluding median segment, without cerci) 30.7, profemur 16.3, protibia 16.1, mesofemur 13.5, mesotibia 12.8, metafemur 18.1, metatibia 18.6.

Measurements, variation (N = 3, MZUSP 1237, 1239, 1986): body (without cerci) 54.6–60.7, head 4.4–5.1, pronotum 4.7–5.3, mesonotum 9.8–11.2.

Male. Similar to that of the female, except: color (Figs. 32c-e, 36c). Dorsolateral portion of body blackish, median portion dark beige to dark brown; legs bearing black or dark brown femora, with a faint crimson hue at the base and with irregular faint white or beige stains along the length. Antennae with a distinct orange hue. Tibiae usually beige with a conspicuous lighter patch where it articulates with the femora and irregular dark brown to black stains along the length. Head (Fig. 33a, c, g). Smaller. Antennae similar to that of the female, but from the 14th antennomeres onwards, each segment is further subsegmented into 2–6 less articulated subsegments of equal width. Thorax (Figs. 32c-e, 33a-c, f). Narrower, mesepisternum bearing sparse round granules. Legs (Fig. 32c-e). Setae in carinae longer, distributed in a single row with alternated distribution. Abdomen (Figs. 32c-e, 33d-f, h). Narrower, tergum I–VII longer than wide, VIII wider than long, IX–X as long as wide. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 6 small teeth distributed in a circle partially covered by setae. Epiproct wide, projecting dorsally from posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and slightly curving towards anterior; bearing a large linear dextral process slightly narrowing towards posterior, slightly upcurving in the last third, gently laterally flattened, and a short triangular sinistral process, both originating superiorly from the anterior portion of sternum IX. Cerci longer, cylindrical, slightly expanded basally, gently incurved. Vomer very large, wide, strongly tapering, terminal hook small, acute but with blunt dorsoventrally flattened apex, upwardly curved. Genitalia (Fig. 34). Dorsal sclerite with basal part wider than apical part; main process with sinuous lateral margin and with widely round apex; distal process somewhat long, narrow, with round very slightly widened apex. Internal sclerite with main process with four short projections equally apart; accessory process with short, somewhat conical inner branch and widely round proximal part of outer branch and short, round distal part of outer branch. Ventral sclerite compound, asymmetric with somewhat straight margin tapering towards the left side and then round.

Measurements (MZUSP 1967): body (without cerci) 51.1, head 3.4, antennae 45.5, pronotum 3.7, mesonotum 9.5, metanotum 3.2, median segment 4.1, abdomen (excluding median segment, without cerci) 27.2, profemur 16.3, protibia 17.6, mesofemur 12.5, mesotibia 13.7, metafemur 17.5, metatibia 20.4.

Measurements, variation (N = 2, MZUSP 1342, 3939): body (without cerci) 48.0–55.3, head 3.2–3.3, pronotum 3.4–3.7, mesonotum 9.1–9.8.

Egg (Fig. 35). Irregularly brownish, rarely grayish, micropylar plate beige. Capsule barrel-shaped, higher than wide, longer than wide in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule and operculum covered by net of irregular, significantly thick rugose ridges. Ridges further bearing minute mushroom-like structures also on the operculum. Micropylar plate round, elliptic or ovate, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line very short to short, less than a fourth of the length of the micropylar plate. Operculum with a central irregular rugose ridge. Measurements (N = 10): length 3.3–3.9, width 2.2–2.5, micropylar plate length 0.8–1.2, operculum maximum diameter 1.7–1.9.

Materials examined

Holotype, 1 m#: Brazil, São Paulo, Ribeirão Grande, Intervales, Pousada Onça Pintada, 24°16′03.0"S 48°24′55.5"W. 02.iv.2022, E.B. Crispino col. (MZUSP 1967).

Paratypes, 1f#: Brazil, São Paulo, Ribeirão Grande, Intervales, Pousada Onça Pintada, 24°16′03.0"S 48°24′55.5"W. 02.iv.2022, E.B. Crispino col. (MZUSP 1986); 3f# (MZUSP 1237–1239), 25 eggs: Brazil, São Paulo, Ribeirão Grande, Parque Estadual Intervales, próximo à Pousada Onça Pintada, 24º16′04"S 48º24′53"W. 14–17.iii.2019, E.B. Crispino col.; 1 m#: Brasil, São Paulo, Ribeirão Grande, P.E. Intervales (MZUSP 1342).

Additional material examined, 1 m#: Brazil, São Paulo, Tapiraí, antes da Trilha dos Tucanos, 24°00′29.2"S 47°33′50.2"W. 12.i.2019, V. M. Ghirotto col. (MZUSP 3939).

Etymology. The specific epithet is an adjective referring to the type locality, Parque Estadual Intervales, and the characteristic of the region itself, which is a section of a mountain formation located among valleys, in Portuguese “intervales”.

Differential diagnosis. Differs from other Urucumania by the dark orange, stripeless coloration with light tibiae and tarsi and mottled femora lighter basally and darker apically (Fig. 32), the elongate (around 4 × longer than wide) cerci in both sexes (Fig. 33d, e, j, m), and very elongate, linear right process of sternum IX in males (Fig. 33h).

Distribution (Fig. 68). Only known from Ribeirão Grande and Tapiraí, São Paulo, Brazil, in areas of ombrophilous Atlantic Forest in a section of the Serra do Mar formation.

Urucumania brasil Ghirotto & Soares sp. n.

(Figs. 37, 38)

Fig. 37

Urucumania brasil sp. n., female holotype from Loreto, Maranhão, Brasil, in dorsal (a, b) and lateral (c) views. Scale bars = 5 mm

Fig. 38

Urucumania brasil sp. n., female holotype from Loreto, Maranhão, Brasil. a, c Head and pronotum in dorsal (a) and lateral (c) views. b Metathorax in dorsal view. d–f Terminalia in dorsal (d), lateral (e), and ventral (f) views. Scale bars = 1 mm

Description

Female. Color (Fig. 37). Head and body black dorsally and laterally, except for beige abdomen laterally, and dark yellow to beige ventrally. Antennae annulated in black and yellow. Presenting several yellow stripes, two short dorsally present only at anterior portion and two dorsolaterally on the head; laterally at each side and posteriorly at pronotum, mesonotum, metanotum and terga I–VIII, the posterior stripe is medially interrupted, but only a faded interruption in the mesonotum. Terga II–VIII with two paramedial yellow stripes (becoming more beigish towards posteriormost segments), metanotum and median segment bearing interrupted paramedial yellow stripes, anteriorly and posteriorly in the median segment and anteriorly in the metanotum. Terga IX–X irregularly covered in beige-yellowish, with a medial black stripe stronger in IX. Coxae superiorly black and inferiorly yellowish-beige. Femora black with dark red in basal third, tibiae and tarsi black. Head (Fig. 38a, c). Large, ovoid, 1.2 × longer than wide, 1.8 × longer than high, about as long as pronotum, with sparse brown conical setae, tuberculated. Vertex flat, slightly convex posteriorly, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, slightly less than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with at least 14 fully segmented antennomeres; from 15th onwards, each segment is further subsegmented into 2–4 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, longer than wide, subcylindrical, slightly flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump absent. Thorax (Figs. 37, 38a-c). Cylindrical in cross section, somewhat densely granulated, covered in short small black setae, bearing lateral carinae. Pronotum subrectangular; sulci shallow, longitudinal straight and transversal slightly sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior two thirds, and a further row of smaller round granules at anterior three fourths at each lateral margin. Metanotum very slightly longer than median segment. Fore and hindwings very small, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 37). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae gently elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as or longer than following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 37, 38b, d-f). Somewhat densely tuberculated, covered in short small black setae. Terga I–IX wider than long, X as long as wide. Terga I–IX bearing small, short round projections pointing posteriad at posterior margin, each a single medially and one in each lateral carinae, except for tergum I (median segment) without lateral projections. Tergum X with medial carina at posterior half, posterior margin somewhat straight and gently emarginated, and widening towards lateral edge. Preopercular organ elliptical, rugose, black, convex, posteriorly with a further round rugosity. Subgenital plate wider at mid length, tapering at posterior third to a slightly lanceolate posterior margin, almost reaching posterior margin of tergum X. Epiproct surpassing tergum X, very wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci cylindrical, conical posteriorly tapering to a round apex (Fig. 38d-f). Gonapophysis VIII long and very narrow, tapering towards apex, apically setose, longer than gonapophysis IX. Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin wider and with a wide incurved sclerotization from the base of gonoplac to before the prominence of the inner margin. Gonoplac incurved, gently widened basally, apex round, as long as gonapophysis IX.

Measurements (MZUSP 2572): body (without cerci) 62.5, head 5.2, antennae at least 32.6, pronotum 5.05, mesonotum 10.48, metanotum 4.3, median segment 4, abdomen (excluding median segment, without cerci) 33.47, profemur 16.51, protibia 16.72, mesofemur 13.7, mesotibia 13.13, metafemur 19.4, metatibia 19.3.

Male. unknown.

Egg. unknown.

Materials examined

Holotype, 1f#: Brazil, Maranhão, Loreto, Pedrinhas district, cerrado strictu sensu, 6°57′20.6"S 45°11′16.9"W, 14.xi.2021 W.L. da Silva col. (MZUSP 2572).

Etymology. The specific epithet is a noun in apposition derived from a Portuguese adjective, in reference to the general color of the species as it might resemble a burning piece of carbonized wood, as ember, “brasa” in portuguese and “brasil” when meaning ‘like ember’. Specimens seen but not collected showed an orangish coloration much resembling ember. The epithet is also a reference to Brazil, especially as the country also takes its name, although indirectly, from ember, due to the coloration of the wood of a Brazilian tree species, the pau-brasil.

Differential diagnosis. Females (the only known sex) differ from remaining Urucumania by the striking black and yellow coloration pattern with legs black with femora basally reddish (Fig. 37), proportions of the head (1.2 × longer than wide; Fig. 38a) and terga VIII–X (tergum X as long as wide; Fig. 38d), the very wide epiproct (Fig. 38d), and the thick, slightly conical cerci about 2.7 × longer than wide (Fig. 38d-f).

Distribution (Fig. 68). Only known from Loreto, Maranhão, Brazil, in a savannic area of the Cerrado.

Remarks. Although bearing contrasting patches of black and yellow resembling some aposematic animals, in fact the coloration pattern of the species likely helps it camouflaging amidst clear, pale and darker dry leaves in the soil of its habitat, as seen for the live specimen.

Urucumania tapirape Ghirotto sp. n.

(Figs. 39, 40)

Fig. 39

Urucumania tapirape sp. n., female holotype from Barra do Tapirapé, Mato Grosso, Brazil, in dorsal (a) and lateral (b) views. Scale bars = 2 mm

Fig. 40

Urucumania tapirape sp. n., female holotype from Barra do Tapirapé, Mato Grosso, Brazil. a, c Head and pronotum in dorsal (a) and lateral (c) views. b Metathorax in dorsal view. d–f Terminalia in dorsal (d), lateral (e), and ventral (f) views. Scale bars = 1 mm

Description

Female. Color (Fig. 39). Head and body dorsally beige, laterally and ventrally dark beige, dorsally with a black medial longitudinal line running along the entire length of head, thorax and abdomen, entirely sparsely covered in white irregular small stains. Alae veins light beige, cells black. Femora black with orangish brown base and short stains of orangish brown at the apex, tibiae and tarsi orangish brown. Head (Fig. 40a, c). Large, subrectangular, 1.4 × longer than wide, 2.2 × longer than high, about as long as pronotum, irregularly tuberculated, with sparse setae. Vertex straight, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye small, round, slightly less than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennae broken at pedicellus. Thorax (Figs. 39, 40a-c). Cylindrical in cross section, covered in short small setae, bearing lateral carinae, irregularly tuberculated. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal slightly sinuous; lateral edges marked with a conspicuous tuberculated carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior half, and a further row of smaller round granules at anterior two thirds at each lateral margin. Metanotum very slightly shorter than median segment. Fore and hindwings very small, scale-like, elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 39). Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than foreleg (although the only foreleg left is regenerated). All five carinae elevated and marked by rows of short setae. Femora slightly thinner and curved basally. Basitarsi as long as to longer than following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 39, 40b, d-f). Tuberculated, covered in short small black setae. Median segment and tergum II–III wider than long, IV and VIII about as long as wide, V–VII and IX–X longer than wide. Terga I–VIII bearing elliptical projections pointing posteriad at posterior margin, each a single medially curved on I–VI or small, round on VII–VIII, and one in each lateral carinae, except for tergum I without lateral projections. Terga III–VI with two paramedian carinae near the medial line, connected to the posterior projection; VII and VIII with two paramedian elevations similar to the carinae of anterior segments. Tergum X with medial carina at posterior half, with a seemingly artificial carinae possibly being a fold at anterior half; posterior margin broadly curved, but widening towards lateral edge. Preopercular organ, wrinkled and rugose, dark, convex. Subgenital plate wider at mid length, then tapering to a roundly lanceolate posterior margin, reaching two thirds of tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins anteriorly straight. Cerci 2.8–3.0 × longer than wide, slightly thick, slightly tapering towards posterior, apex round (Fig. 40d-f).

Measurements (MZUSP 0203): body (without cerci) 59.46, head 4.39, antennae —, pronotum 4.71, mesonotum 10.6, metanotum 2.81, median segment 5.29, abdomen (excluding median segment, without cerci) 31.66, profemur 14.52, protibia 14.72, mesofemur 11.50, mesotibia 11.9, metafemur 17.38, metatibia 17.78.

Male. unknown.

Egg. unknown.

Materials examined

Holotype, 1f#: Barra do Tapirapé [Santa Teresinha, Tapirapé], Mato Grosso, xii.1962, B. Malkin col. (MZUSP 0203).

Etymology. The specific epithet is a noun in apposition referring both to the type locality, the mouth of the Tapirapé river, as well as the Tapirapé indigenous people inhabitants of that area. The Tapirapé people were driven out of their original territory from centuries ago by urban and farming expansion and were divided in more than one area, some inhabiting the Tapirapé area in Mato Grosso for centuries until today. Their craft often display patterns of beige and black similar to those of the new species.

Differential diagnosis. Females (the only known sex) differ from remaining female Urucumania by the beige color pattern with orangish tibiae and tarsi and femora black with orangish base (Fig. 39), and by the elongate head (Fig. 40a) and terga VIII–X (Fig. 40d, e).

Distribution (Fig. 68). Only known from Santa Teresinha, Mato Grosso, Brazil, in a savannic area of the Cerrado.

Remarks. The locality is given as “Barra do Tapirapé” which means “Tapirapé’s [river] mouth bar’’, so the locality most probably refers to the mouth of the Tapirapé flowing into the Araguaia river, in Santa Teresinha municipality—Mato Grosso state.

Urucumania rasocatarinesis Ghirotto sp. n.

(Figs. 41, 42, 43, 44, 45 and 46)

Fig. 41

Urucumania rasocatarinensis sp. n., female paratype from Jeremoabo, Bahia, Brazil, in dorsal (a) and lateral (b) views. Scale bars = 5 mm

Fig. 42

Urucumania rasocatarinensis sp. n., male paratype from Jeremoabo, Bahia, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 5 mm

Fig. 43

Urucumania rasocatarinensis sp. n., paratype male (a–h) and holotype female (i–o) from Jeremoabo, Bahia, Brazil. a, c, g, i, k Head and pronotum in dorsal (a, i), lateral (c, k) and ventral (g) views. b, m Metathorax in dorsal view. d, e, f, h, j, k, n Terminalia in dorsal (d, j), left side (e, l), right side (h) and ventral (f, n) views. Scale bars = 1 mm

Fig. 44

Urucumania rasocatarinensis sp. n., phallic organ of paratype male from Jeremoabo, Bahia, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 45

Urucumania rasocatarinensis sp. n., eggs of paratype female from Jeremoabo, Bahia, Brazil. a Lateral view. b Dorsal view. c Two operculi in anterior view. d Several eggs in lateral and dorsal views. Scale bars = 1 mm

Fig. 46

Urucumania rasocatarinensis sp. n., live insects in situ. a Adult female. b Adult male. c Juvenile female. d Mating pair, male above female

Description

Female. Color (Figs. 41, 46a, d). Body, antennae and legs entirely brown to moderately dark brown, body dorsally with a thick medial black longitudinal line running along the head, thorax and then gradually fading from sterna II to VII, flanked by an orange band dimming laterally, very thin in the anterior half of pronotum, fading to brown in the anterior portion of head and orangish light brown on the abdomen. Head with lateraldorsal black lines behind the eye. Alae dark brown on superior half and beige on inferior half. Subgenital plate slightly lighter on posterior two thirds. Head (Fig. 43i, k). Large, subquadrate to ovoid, 1.3 × longer than wide, 1.3 × longer than high, slightly longer than pronotum, with sparse stiff setae. Vertex flat, frontal area gently elevated. Frontal suture slightly curved and deep, frontal convexity prominent. Eye medium sized, round, less than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with around 23 fully segmented antennomeres; from 16th onwards, each segment is further subsegmented into 2–8 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump apparently present, small, discrete, in the same color as the rest of the segment and poorly elevated, present on the 14th segment and the first subsegment of the 16th segment. Thorax (Figs. 41, 43i, k, m). Robust. Cylindrical in cross section, covered in short small stiff setae, bearing lateral carinae, with few diminute granules except in the carinae. Pronotum subrectangular, slightly longer than wide, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum bearing on each side a paramedian row of conical granules on anterior half on an elevation, and a further sparse row of smaller, short round granules at anterior two thirds at each lateral margin on a prominent keel. Metanotum very slightly longer than median segment. Fore and hindwings diminute, scale-like; both round, hind slightly elliptical. Meso and metafurca shortened, each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 41). Legs moderately thickened. Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae slightly elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as the following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 41, 43j, l-n). Robust. Sparsely tuberculated, covered in short small black setae. All terga wider than long, tergum X only slightly. Terga II–VII with lateral margins keeled. Terga I–IX dorsally bearing very short wide humps near posterior margin, developed in a round short projection at terga VI–IX; terga II–VIII further bearing a conical projection at the end of each lateral carinae. Terga VII–IX with a medial elevation accompanying the posterior projection. Tergum X with medial carina at posterior half, posterior margin straight to slightly emarginated, and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, posteriorly slightly widening. Subgenital plate wider at mid length, then tapering to a slightly lanceolate posterior margin, reaching the base of the cerci and reaching two thirds of tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci short, slightly thick, 2.1 × longer than wide, cylindrical (Fig. 43j, l, n). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, as long as gonapophysis IX. Gonapophysis IX triangular basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin. Gonoplac incurved, gently widened basally, apex round, as long as gonapophysis IX.

Measurements (MZUSP 2579): body (without cerci) 52.83, head 5.45, antennae 35.22, pronotum 5, mesonotum 8.3, metanotum 3.98, median segment 3.75, abdomen (excluding median segment, without cerci) 26.35, profemur 14.2, protibia 14.77, mesofemur 12.06, mesotibia 12.04, metafemur 16.03, metatibia 17.5.

Measurements, variation (N = 2, MZUSP 2580, 2582): body (without cerci) 46.5–50.0, head 4.4–4.6, pronotum 4.7–5.3, mesonotum 7.6–8.3.

Male. Similar to that of the female, except: color (Figs. 42, 46b, d). Generally darker, dorsal stripes flanking the dark medial line are more beigish. Head (Fig. 43i, k). 1.1 × longer than wide, narrower, eyes larger, scapus and pedicel larger. Thorax (Figs. 42, 43a-c, g). Slightly narrower, mesepisternum bearing sparse round granules. Legs (Fig. 42). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 42, 43b, d-f, h). Slightly narrower, tergum X longer than wide. Posterior margin of tergum X straight to very slightly emarginated, thorn pads pointing inward, near median line, bearing around 12 teeth. Epiproct wide, not projecting dorsally. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin overall round and slightly curving towards anterior; bearing a linear dextral process slightly narrowing towards posterior, slightly upcurving, gently laterally flattened at basal half, originating superiorly from the anterior portion of sternum IX. Cerci longer, 3.5 × longer than wide, cylindrical, very slightly constricted medially to post medially, gently incurved (Fig. 43d, e, h). Vomer very large, wide, strongly tapering, terminal hook small, acute but with slightly blunt apex, slightly upwardly curved. Genitalia (Fig. 44). Dorsal sclerite with basal part narrower than apical part; main process somewhat rectangular, wide, with protruding round posterior apex, with slightly sinuous, convex lateral edges; distal process short, somewhat thick, with round apex directed laterally. Internal sclerite with main process with four short projections; accessory process with protruding bent conical inner branch and widely round parts of outer branch. Ventral sclerite compound, asymmetric with somewhat round margin, thicker at the middle and slightly pointing right.

Measurements (MZUSP 2578): body (without cerci) 39.4, head 3.2, antennae 34.5, pronotum 3.3, mesonotum 6.4, metanotum 3.2, median segment 2.8, abdomen (excluding median segment, without cerci) 20.5, profemur 12.2, protibia 12.6, mesofemur 9.8, mesotibia 9.8, metafemur 13.2, metatibia 14.5.

Measurements, variation (N = 1, MZUSP 2581): body (without cerci) 39.2, head 3.2, pronotum 3.5, mesonotum 6.3.

Egg (Fig. 45). Irregularly creamish, beige or brownish, micropylar plate beige to creamish. Capsule ovoid, slightly barrel-shaped, higher than wide, longer than wide in variable ratios, oval in cross section; polar area somewhat flat to round. Operculum perpendicular, oval and flat. Capsule covered by net of irregular rugose ridges, which are irregular and forming numerous ridges on the operculum. Ridges further bearing very minute mushroom-like structures also on the operculum. Micropylar plate oblong to elliptical, or slightly ovate, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose round hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line moderately thick, more than half the length of the micropylar plate. Measurements (N = 15): length 2.8–3.6, width 1.8–2.4, micropylar plate length 0.9–1.3, operculum maximum diameter 1.3–1.9.

Materials examined

Holotype, 1 m#: Brazil. Bahia. Jeremoabo. Caatinga densa diversa em plantas, borda do Raso da Catarina. 10°02′09.0"S, 38°24′10.6"W. 20.iii.2023. V.M. Ghirotto, P.W. Engelking, E.W. Engelking col. (MZUSP 2578).

Paratypes, 3f# (MZUSP 2579, 2580, 2582), 1 m# (MZUSP 2581), 30 eggs: Brazil. Bahia. Jeremoabo. Caatinga densa diversa em plantas, borda do Raso da Catarina. 10°02′09.0"S, 38°24′10.6"W. 20.iii.2023. V.M. Ghirotto, P.W. Engelking, E.W. Engelking col.

Etymology. The specific epithet is an adjective derived from the type locality, Raso da Catarina, a plateau in the Caatinga biome. The Raso da Catarina region appears to be a very important area for the Caatinga bearing endemic species such as the endangered Lear's indigo macaw.

Differential diagnosis. Differs from remaining Urucumania by the dark orange and brown color pattern with brown legs, and by the very robust body (Figs. 41, 42). The robust body resembles Urucumania dilatata sp. n. and Urucumania sertaneja sp. n., from which it differs by the orange banded color pattern, less developed posterior projections of terga and the smaller wings (Figs. 41, 42).

Distribution (Fig. 68). Only known from Jeremoabo, Bahia, Brazil, in the Raso da Catarina plateau in the xeric Caatinga.

Urucumania dilatata Ghirotto sp. n.

(Figs. 47, 48, 49, 50, 51 and 52)

Fig. 47

Urucumania dilatata sp. n., holotype female from Umburanas, Bahia, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 5 mm

Fig. 48

Urucumania dilatata sp. n., paratype male from Sento Sé, Bahia, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 5 mm

Fig. 49

Urucumania dilatata sp. n., holotype female from Umburanas, Bahia, Brazil. a, c, e Head and pronotum in dorsal (a), lateral (c) and ventral (e) views. g Metathorax in dorsal view. b, d, f, h Terminalia in dorsal (b), lateral (d) and ventral (f) views. h Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 50

Urucumania dilatata sp. n., paratype male from Sento Sé, Bahia, Brazil. a, c Head and pronotum in dorsal (a) and lateral (c) views. e Metathorax in dorsal view. b, d, f, g, h Terminalia in dorsal (b), right side (d), left side (f) and ventral (g, h) views. Scale bars = 1 mm

Fig. 51

Urucumania dilatata sp. n., phallic organ of paratype male from Sento Sé, Bahia, Brazil. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f Internal sclerite. Scale bars: a–e 1 mm; f 0.5 mm

Fig. 52

Urucumania dilatata sp. n., paratypes from Sento Sé, Bahia, Brazil, in situ, mating, male above (a, b) and holotype from Umburanas, Bahia, in situ, when sighted, walking on a cactus (c, d). Photos courtesy of Willianilson Pessoa da Silva (a, b) and Márcio Perez Bolfarini (c, d)

Description

Female. Color (Figs. 47, 52). Entirely black; tip of subgenital plate, base of femora and tarsi very dark, blackish red; head with dorsolateral blackish dark red bands; antennae dark red and ringed in creamish from apical half. Allae bright yellow except for the blackish or dark yellow cells. Head (Fig. 49a, c, e). Large, slightly rugose, considerably wider posteriorly and gradually tapering towards anterior, around 1.0–1.1 × longer than wide, 1.1 × longer than high, about as long as pronotum, with dark conical setae. Vertex slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with at least 26–28 fully segmented antennomeres; from 16–18th onwards, each segment is further subsegmented into 2–8 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, slightly flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump small, discrete, not prominent, in the same color as the rest of the segment, present on the first subsegment of the 16–17th segment. Thorax (Figs. 47, 49a, c, e, g). Very robust, widened, rugose. Subrectangular in cross section, covered in short small black setae, bearing lateral strong carinae, with short and round to conical granules, including conical ones on the carinae. Pronotum somewhat trapezoidal, tapering towards anterior; sulci marked, longitudinal straight and transversal approximately in “V” shape; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum slightly wider than long, lateral margins round, bearing on each side a paramedian row of conical granules around anterior half on an elevation, and a further row of smaller round to conical granules at anterior two thirds at each lateral margin on a prominent keel. Metanotum slightly shorter than median segment. Fore and hindwings round, prominent but scale-like; forewing reaching half the length of metanotum, hindwing almost reaching posterior margin of median segment. Meso and metafurca each separated longitudinally, mesofurca curved, metafurca somewhat straight, anteriorly deeper. Legs (Fig. 47). Robust, thick. Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as the following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 47, 49b, d, f, g, h). Widened, rugose. Sparsely tuberculated, covered in short small black setae. All terga wider than long. Terga II–VII with lateral margins keeled. Terga I–VIII bearing elliptical projections pointing posteriad at posterior margin, each a single curved one medially and one in each lateral carinae, except for tergum I without lateral projections; additionally, two pairs of paramedial projections on terga I–VI, gradually becoming smaller. Terga VII–IX only with the medial projection, shorter than in other terga. Terga III–VII with two paramedian carinae near the medial line, connected to the posterior projection and with two paramedian carinae each near one of the lateral margins; VIII and IX with two paramedian central elevations similar to the carinae of anterior segments. Tergum X with medial elevation or wide carina at posterior third, posterior margin slightly curved to very slightly emarginated, widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black to dark brown, convex. Subgenital plate wider at mid length, tapering to a rounded to somewhat lanceolate posterior margin, reaching the end of tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically slightly apart, inner margins almost straight. Cerci short, 2.3–2.6 × longer than wide, thick, cylindrical (Fig. 49b, d, f). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, longer than gonapophysis IX (Fig. 49h). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 49h). Gonoplac incurved, widened basally, apex round, longer than gonapophysis IX (Fig. 49h).

Measurements (MZUSP 2587): body (without cerci) 57.7, head 5.5, antennae 38.9, pronotum 5.8, mesonotum 9.1, metanotum 3.6, median segment 4.4, abdomen (excluding median segment, without cerci) 29.3, profemur 14.3, protibia 14.0, mesofemur 11.6, mesotibia 11.3, metafemur 15.4, metatibia 16.8.

Measurements, variation (N = 1, MZUSP 2574): body (without cerci) 54.2, head 5.1, pronotum 5.2, mesonotum 8.0.

Male. Similar to that of the female, except: color (Figs. 48, 52). Dorsally with blackish dark red tint in paramedial bands, dark red on the head more conspicuous. Head (Fig. 50a, c). Narrower, with more homogeneous width, eyes larger, slightly more than hemispherical, antennae larger. Thorax (Figs. 48, 50a, c, e). Slightly narrower, more elongate, granules rounder. Forewings relatively shorter. Legs (Fig. 48). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 48, 50b, d, e–h). Slightly narrower, tergum X bearing a longitudinal carina at posterior half, posterior margin of tergum X very gently emarginated, widening towards lateral edges, thorn pads small, near median line, bearing around 4 teeth. Epiproct moderately wide, projecting dorsally from posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and round, gently curving towards anterior; bearing a large linear dextral process moderately narrowing towards posterior, slightly upcurving, gently laterally flattened and a short triangular sinistral process, both originating superiorly from the anterior portion of sternum IX. Cerci longer, around 2.6 × longer than wide, cylindrical, straight (Fig. 50b, g). Vomer very large, very wide, strongly tapering, terminal hook very small, acute, slightly upwardly curved (Fig. 50h). Genitalia (Fig. 51). Dorsal sclerite with basal part narrower than apical part; main process somewhat rectangular, elongate, with protruding conical posterior apex, with slightly sinuous lateral edges; distal process short, with round slightly pointed apex directed laterally. Internal sclerite with main process with four short projections; accessory process with protruding conical inner branch and widely round parts of outer branch. Ventral sclerite compound, asymmetric with somewhat round margin, thicker at the middle and slightly pointing right.

Measurements (MZUSP 2573): body (without cerci) 44.1, head 3.9, antennae 37.0, pronotum 3.7, mesonotum 6.4, metanotum 2.6, median segment 3.3, abdomen (excluding median segment, without cerci) 24.2, profemur 11.3, protibia 11.9, mesofemur 8.9, mesotibia 9.4, metafemur 11.3, metatibia 13.7.

Egg. unknown.

Materials examined

Holotype, f#: Brazil, Bahia, Umburanas, na serra, 10º41′47.0″S, 41º11′16.6″W, 27.v.2021, M. Bolfarini col. (MZUSP 2587).

Paratypes, 1 m# (MZUSP 2573), f# (MZUSP 2574): Brazil, Bahia, Sento Sé. Minas do Mimoso, 10º19′32.5″S, 41º33′22.2″W. 11.v.2022, W. Pessoa col.

Etymology. The specific epithet is an adjective from the Latin word "dilatatus" meaning thick, enlarged, referring to the very widened body, especially the thorax, characteristic of the new species.

Differential diagnosis. Differs from remaining Urucumania by the very dark color pattern with bright yellow wings and dark legs, and by the very robust body with laterally projecting terga (Figs. 47, 48). The very robust body resembles Urucumania rasocatarinensis sp. n. and Urucumania sertaneja sp. n. from which it differs by the relatively more widened thorax and larger wings (Figs. 47, 48).

Distribution (Fig. 68). Only known from Umburanas and Sento Sé, Bahia, Brazil, in very dry areas of xeric Caatinga.

Urucumania sertaneja Ghirotto & Alves sp. n.

(Figs. 53, 54, 55, 56 and 57)

Fig. 53

Urucumania sertaneja sp. n., paratype female from Piranhas, Alagoas, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 2 mm

Fig. 54

Urucumania sertaneja sp. n., holotype male from Piranhas, Alagoas, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 2 mm

Fig. 55

Urucumania sertaneja sp. n., paratype male (a–h) and holotype female (i–o) from Piranhas, Alagoas, Brazil. a, c, g, i Head and pronotum in dorsal (a, g) and lateral (c, i) views. b, l Metathorax in dorsal view. d, e, f, h, j, k, m Terminalia in dorsal (d, j), left side (e, k), right side (h) and ventral (f, m) views. Scale bars = 1 mm

Fig. 56

Urucumania sertaneja sp. n., female from Natal, Rio Grande do Norte, Brazil, in dorsal (a), lateral (b) and ventral (c) views. Scale bars = 2 mm

Fig. 57

Urucumania sertaneja sp. n., female from Natal, Rio Grande do Norte, Brazil. a, c, e Head and pronotum in dorsal (a), lateral (c) and ventral (e) views. f Metathorax in dorsal view. b, d, g Terminalia in dorsal (b), lateral (d) and ventral (g) views. Scale bars = 1 mm

Description

Female. Color (Fig. 53). Entirely dark, with faint tints of dark orange dorsally; head with dorsolateral dark orange bands; antennae black and ringed in creamish from apical half. Allae black with creamish edges. Head (Fig. 55g, i). Large, slightly rugose, considerably wider posteriorly and gradually tapering towards anterior, as long as wide, 1.2 × longer than high, slightly shorter than pronotum, with dark conical setae. Vertex slightly convex, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with at least 14–15 fully segmented antennomeres; from 12th onwards, each segment is further subsegmented into 2–4 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, slightly flattened dorsoventrally only at basal portion; pedicellus subcylindrical, almost as long as scapus. Antennal bump small, discrete, not prominent, in the same color as the rest of the segment, present on the first subsegment of the 12th segment. Thorax (Figs. 53, 55g, i, m). Very robust, widened, rugose, densely granulated. Subrectangular in cross section, covered in short small black setae, bearing lateral strong carinae, with short and round to conical, pointed granules, including conical ones on the carinae. Pronotum somewhat trapezoidal, tapering towards anterior; sulci marked, longitudinal straight and transversal approximately in “U” shape; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum slightly longer than wide, lateral margins gently round, bearing on each side a paramedian row of conical granules around anterior half on a slight elevation, and a further row of smaller round to conical granules at anterior two thirds at each lateral margin on a keel. Metanotum slightly longer than median segment. Fore and hindwings round, slightly elongated, scale-like; forewing reaching anterior edge metanotum, hindwing almost reaching the middle of median segment. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 53). Robust, thick. Hindlegs surpassing abdomen and longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Probasitarsi slightly longer than following three tarsomeres combined, meso- and metabasitarsi about as long as the following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 53, 55j-n). Widened, rugose. Sparsely tuberculated, covered in short small black setae. All terga wider than long. Terga II–IX with lateral margins keeled. Terga I–IX bearing elliptical projections pointing posteriad at posterior margin, each a single curved one medially and one in each lateral carinae, except for tergum I without lateral projections; additionally, two pairs of paramedial projections on terga I–VI. Terga III–VII with two paramedian carinae near the medial line, connected to the posterior projection and with two paramedian carinae each near one of the lateral margins. Tergum X with medial elevation or wide carina at posterior third, posterior margin slightly curved to very slightly emarginated, widening towards lateral edge. Preopercular organ as long as wide, wrinkled and rugose, black, convex. Subgenital plate wider at mid length, tapering to a rounded to somewhat lanceolate posterior margin, almost reaching the end of tergum X. Epiproct exceeding tergum X, wide, round, with a medial carina. Paraprocta apically slightly apart, inner margins almost straight. Cerci short, 2.3–2.5 × longer than wide, cylindrical (Fig. 55k, n). Gonapophysis VIII long and narrow, tapering towards apex, apically setose. Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin. Gonoplac incurved, widened basally, apex round.

Measurements (MZUSP 3869): body (without cerci) 50.5, head 4.8, antennae 36.8, pronotum 4.6, mesonotum 8.2, metanotum 3.0, median segment 3.8, abdomen (excluding median segment, without cerci) 26.1, profemur 12.6, protibia 13.2, mesofemur 9.9, mesotibia 10.8, metafemur 13.2, metatibia 15.3.

Measurements, variation (N = 1, MZUSP 3870): body (without cerci) 53.2, head 4.9, pronotum 5.0, mesonotum 8.5.

Male. Similar to that of the female, except: color (Fig. 54). Dorsal faint dark orange tints spread in two paramedian wide bands; orange dorsolateral head bands brighter; creamish edge of allae thicker. Head (Fig. 55a, c). Narrower, with more homogeneous width, eyes larger, antennae larger and longer with around 20 fully segmented antennomeres. Thorax (Figs. 54, 55a-c). Slightly narrower, more elongate, granules rounder. Wings relatively larger. Legs (Fig. 54). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 54, 55b, d-f, h). Slightly narrower, tergum X bearing a longitudinal carina, posterior margin of tergum X very gently emarginated, widening towards lateral edges, thorn pads small, near median line, bearing around 4 teeth. Epiproct moderately wide, projecting dorsally from posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently widened and round, gently curving towards anterior; bearing a large linear dextral process moderately narrowing towards posterior, slightly upcurving, gently laterally flattened and a short triangular sinistral process, both originating superiorly from the anterior portion of sternum IX. Cerci longer, around 2.6 × longer than wide, cylindrical, straight (Fig. 55d, f, h). Vomer very large, very wide, strongly tapering, terminal hook very small, acute, slightly upwardly curved.

Measurements (MZUSP 3871): body (without cerci) 44.0, head 3.8, antennae 40.4, pronotum 3.8, mesonotum 6.5, metanotum 2.9, median segment 3.0, abdomen (excluding median segment, without cerci) 24.0, profemur 11.6, protibia 11.8, mesofemur 8.5, mesotibia 9.4, metafemur 11.7, metatibia 13.6.

Measurements, variation (N = 1, MZUSP 3872): body (without cerci) 44.7, head 3.8, pronotum 3.8, mesonotum 6.5.

Egg. unknown.

Materials examined

Holotype, m#: Brazil, Alagoas, Piranhas, near city and near the margins of São Francisco river, 9°37′33.6"S, 37°45′25.2"W, 29.10.2023, T.J.S. Alves col. (MZUSP 3871).

Paratypes, m# (MZUSP 3872), 2f# (MZUSP 3869, 3870): Brazil, Alagoas, Piranhas, near city and near the margins of São Francisco river, 9°37′33.6"S, 37°45′25.2"W, 29.10.2023, T.J.S. Alves col.

Additional material examined, 2f#: Brazil, Rio Grande do Norte, Natal, Sanvalle, Parque da Cidade. 23.x.2009. M. L. S. Lopes col. (MPEG).

Etymology. The specific epithet is an adjective from Portuguese meaning inhabitant of the “sertão”, a Brazilian semi-arid region bearing the Caatinga biome. The new species was found very near the São Francisco river, which has a high significance for the “sertão” being the most important water source in that dry area.

Differential diagnosis. Differs from other Urucumania by the very darker color pattern with faint dark orange to reddish dorsal stripes, and by the very robust body with laterally projecting terga (Figs. 53, 54). The very robust body resembles Urucumania dilatata sp. n., from which it differs by the smaller alae and less widened thorax (Figs. 53, 54) and Urucumania rasocatarinensis sp. n., from which it differs by the distinct color, the slightly larger allae, the larger thoracic spines, shorter tergum X of the female and more robust thorax and abdomen of the male (Figs. 53, 54, 55).

Distribution (Fig. 68). The type material is known from Piranhas, Alagoas, Brazil, in very dry areas of the xeric Caatinga. The females from Rio Grande do Norte could be revealed to represent a distinct species (see Remarks below).

Remarks. Two females from Natal, Rio Grande do Norte state (Figs. 56, 57) only differ from female type specimens from Piranhas by their very short cerci (Fig. 57b, d, g). Males from that population would be helpful to confirm its identity.

Urucumania albopunctata Ghirotto sp. n.

(Figs. 58, 59, 60 and 61)

Fig. 58

Urucumania albopunctata sp. n., holotype male from Santa Cruz, Buena Vista, Bolivia, in dorsal (a, b) and lateral (c) views. Scale bars = 2 mm

Fig. 59

Urucumania albopunctata sp. n., holotype male from Santa Cruz, Buena Vista, Bolivia. a, c, e Head and pronotum in dorsal (a), lateral (c) and ventral (e) views. b Metathorax in dorsal view. d, f, g, h, i Terminalia in dorsal (d), right side (h), left side (i) and ventral (g, f) views. Scale bars = 1 mm

Fig. 60

Urucumania albopunctata sp. n., phallic organ of holotype male from Santa Cruz, Buena Vista, Bolivia. a Dorsal view. b Ventral view. c Caudal view. d Right side view. e Left side view. f latero-posterior view. g Internal sclerite. Scale bars = 1 mm

Fig. 61

Urucumania albopunctata sp. n., live insects in situ from different municipalities in Santa Cruz department, Bolivia. a Adult male from Yapacani, photo courtesy of Kozue Kawakami. b Mating pair from Buena Vista, photo courtesy of Ramon and Suzanne Vargas

Description

Male. Color (Figs. 58, 61). Head, antennae, body and legs in a black to dark brown background, head dorsally creamy, body with a somewhat thick white median line running from the pronotum to tergum II, terga III to VII dorsally creamy, lighter at median line accompanying the preceding median white line. Femora blackish with a short white stain dorsally at the apex, tibiae dark brown with a basal white stain dorsally and apex darker, tarsi blackish. Coxae dorsally white, mesocoxopleurite, posterior portion of meso and metaepisternum, metacoxopleurite and metaepimeron white. Terga II to IV with a white stain at the apex of each lateroposterior, gradually less marked in posterior terga; sterna II to VI with a round white stain at each lateroposterior edge. Ventrally dark brown in the thorax to blackish on the abdomen. Head (Fig. 59a, c, e). Large, elliptical, 1.4 × longer than wide, 1.3 × longer than high, about as long as pronotum, with sparse round granules and short setae. Vertex flat, frontal area gently elevated. Frontal suture curved and deep, frontal convexity prominent. Eye medium sized, round, slightly more than hemispherical. Clypeus wide, convex, with shallow depressions, labrum notched anteromedially, round. Antennae filiform with at least 21 fully segmented antennomeres; from 14th onwards, each segment is further subsegmented into 2–5 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, slightly shorter than scapus. Antennal bump apparently absent, but first subsegment of the 15th segment with a very slight widening basally. Thorax (Figs. 58, 59a-c, e). Cylindrical in cross section, dorsally and laterally covered in granules more or less arranged in rows and very sparse short small black setae, bearing lateral carinae with granules near the edge of the mesothoracic carinae, ventrally smooth. Pronotum with round, short, not pronounced granules, mesonotum and metanotum with spine-like, pronounced granules, more pronounced on the mesonotum and generally rounder, less acute on the metanotum. Metanotum with a paramedian pair of larger, pronounced round granules at the posterior margin. Pronotum subrectangular slightly tapering towards anterior; sulci shallow, longitudinal straight and transversal curved towards posterior; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Metanotum very slightly shorter than median segment. Completely apterous. Meso and metepisternum with three granules, on mesepisternum very close to each other and located near posterior margin, on metepisternum separated and spread across its length. Mesofurca barely separated and metafurca separated longitudinally, both curved, anteriorly deeper, and both with a prominent round convexity anteriorly. Legs (Fig. 58). Hindlegs surpassing abdomen and slightly longer than forelegs, midlegs shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi about as long as following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 58, 59b, d, f-i). Sparsely granulated dorsally, granules stronger in more anterior terga, covered in short small black setae sparsely dorsally and denser with thicker setae ventrally. Terga I–VIII each bearing a medial curved elliptical projection pointing posteriad, slightly away from the posterior margin; projection on tergum I with further two round granules anteriorly; projection on tergum VII and VIII weaker than in preceding segments. Terga IX and X with two medial round elevations in the same position to the projections of anterior segments. Tergum X with a medial carina and two paramedian carinae at posterior half, the medial reaching the posterior margin and the paramedians not reaching posterior margin. Posterior margin somewhat roundly gently carinated, widening towards lateral edge. Paraprocta apically apart, inner margins short. Tergum II–VII longer than wide, VIII–IX wider than long, X as long as wide. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 4 teeth. Epiproct wide, dorsally hidden from view. Sternum IX voluminous, with very large anterior portion and very large poculum, posterior margin gently widened and slightly curving towards anterior; bearing a large linear sinistral process slightly narrowing towards posterior, gently laterally flattened, originating superiorly from the anterior portion of sternum IX and a similar dextral process but much smaller. Cerci long, cylindrical, slightly constricted medially to post medially, gently incurved (Fig. 59d, f, h, i). Vomer very large, wide, strongly tapering, terminal hook originating from the left, small, acute but with blunt dorsoventrally flattened apex, upwardly curved (Fig. 59g). Genitalia (Fig. 60). Dorsal sclerite with basal part significantly narrower than apical part; main process located to the left, with almost straight margin gently round at edges, apex blunt and round; distal process located to the right, short, apex round, slightly bent towards anterior. Internal sclerite with main process with two short projections; accessory process with long, incurved, cylindrical, slightly narrowing inner branch and wide, triangular outer branch. Ventral sclerite simple, long and extending from the base of the genitalia, narrow, located and bent to the left side.

Variation. Thorn pads with 4–6 teeth; poculum with posterior margin slightly emarginated or round; epiproct projecting or not beyond tergum X.

Measurements, range in mm (Göttingen WS375): body (without cerci) 45.8, head 3.6, antennae at least 41.0, pronotum 3.2, mesonotum 8.0, metanotum 3.0, median segment 3.2, abdomen (excluding median segment, without cerci) 24.8, profemur 14.7, protibia 15.2, mesofemur 11.6, mesotibia 12.9, metafemur 15.9, metatibia 17.9.

Measurements, range in mm (Göttingen WS374): body (without cerci) 49.8, head 3.7, antennae at least 41.0, pronotum 3.4, mesonotum 9.0, metanotum 3.5, median segment 3.4, abdomen (excluding median segment, without cerci) 26.8, profemur 15.2, protibia 16.6, mesofemur 12.1, mesotibia 13.4, metafemur 17.4, metatibia 19.6.

Female. see Remarks and Fig. 61b.

Egg. unknown.

Materials examined

Holotype, 1 m#: Bolivia, Santa Cruz, Buena Vista, border of Buena Vista N.P., MVL, 17º41.721 S, 63º33.766 W, 10.i.2004, Svenson, Cameron, Bybee col. (Göttingen WS375).

Paratypes, 1 m#: Bolivia, Depto. Santa Cruz, Potrerillos del Guenda Res., elev. 400 m, MVL, 17º40.262 S, 63º27.445 W, 16–12.i.2005, J. McHugh lab exped. (Göttingen WS374).

Etymology. The specific epithet is an adjective derived from the Latin words “albus” meaning white and “punctatus” meaning punctuated, referring to the lateral white dots present on the new species.

Differential diagnosis. Males (the only known sex) differs from remaining males of Urucumania by the dark color pattern with white medial line and white lateral dots at the abdomen (Fig. 58), absence of wings (Fig. 59b), and presence of two lateral processes, on each side, of sternum IX (Fig. 59h, i). A female photographed alive but not collected also seems to bear a light-colored medial line and lateral dots and also be totally wingless (Fig. 61b).

Distribution (Fig. 68). Only known from Buena Vista, Santa Cruz, Bolivia, in areas of humid and mountainous forests with Amazonian influences.

Remarks. Although we lack a female voucher to describe, a living mating pair (Fig. 61b) of Urucumania albopuncatata sp. n. was photographed by Ramon and Suzanne Vargas, found by their son Rio Vargas, in Buena Vista, Santa Cruz, Bolivia, around 25 km from the holotype and paratype localities. The female has the same overall color pattern as the male, including the white dots laterally on the abdomen, as well as some characteristics that are shared by both sexes in other species of the genus, such as the small curved projection at the posterior margin of terga and the prominent granules on thorax (Fig. 61b). Furthermore, the female appears to be totally wingless like the male.

Urucumania oriomadeirensis Conle, Ghirotto, Henneman & Valero sp. n.

(Figs. 62, 63)

Fig. 62

Urucumania oriomadeirensis sp. n., female (a–b) and male (c) paratypes from Itacoatiara, Amazonas, and male paratype (d–e) from Itaituba, Pará, Brazil, in dorsal (a, c, d) and lateral (b, e) views. Scale bars = 5 mm

Fig. 63

Urucumania oriomadeirensis sp. n., male (a, d–h) and female (j, m, n) paratype from Itacoatiara, Amazonas, and male (b, c) and female (i, l, k, o, p) paratypes from Itaituba, Pará, Brazil. a, c, f, i, l, m Head and pronotum in dorsal (a, i), lateral (c, l, m) and ventral (f) views. b, o Metathorax in dorsal view. d, e, g, h, j, n, p Terminalia in dorsal (d, j), left side (g), right side (h, n) and ventral (e, p) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Description

Female. Color (Fig. 62a, b). Known only from aged dry preserved specimens. Head and body dorsally mid to dark orange, ventrally dark orangish brown, laterally head, pronotum and anterior two thirds of mesonotum black, last three to five terga darker, dark brown, and the rest of body orangish brown. Head, pronotum and mesonotum with more vivid orange interrupted by a thin median longitudinal black line that also runs faintly through metanotum. Posterior third to half of the lateral margins of mesonotum, lateral margins of metanotum, and posterior two thirds of the lateral margins of median segment, with a thin beige to light orange band, including the alae. Ventral region of thorax, lateral posterior third of mesonotum, and entire metanotum and abdomen with several light orange circular very fine granules in a dark orange to brown background; light granules denser dorsally. Antennae basally light orange then brown to black. Coxae dark brown, femora dark orange except for black apical two thirds on profemora and black apical one third on meso and metafemora; tibiae and tarsi dark brown to black. Head (Fig. 63 i, m). Large, ovoid to subrectangular, 1.3 × longer than wide, 1.6 × longer than high, sparsely tuberculated, covered in short brown setae, about as long as pronotum. Vertex slightly convex, frontal area gently elevated. Frontal suture gently curved and deep, frontal convexity prominent. Eye medium sized, round, hemispherical. Clypeus wide, convex, with shallow depressions and strongly notched anteromedially, labrum notched anteromedially, round. Antennae filiform with around 21–25 fully segmented antennomeres; from 15–17th onwards, each segment is further subsegmented into 4–9 less articulated subsegments of equal width, more conspicuous and numerous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, as long as scapus. Third antennomere as long as or slightly longer than scapus and pedicel combined. Antennal bump absent. Thorax (Figs. 62a, b, 63i, l, o). Subquadrate in cross section, densely granulated, covered in short small brownish setae, bearing lateral carinae. Pronotum with sparser granules, rectangular; sulci shallow, longitudinal straight and transversal sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Mesonotum laterodorsally bearing two rows of conical granules on anterior three fifths, and a further row of very sparse and small round granules at each lateral margin. Metanotum slightly shorter than median segment. Fore and hindwings very small, scale-like, elongated; forewing bud elliptical, hindwing bud triangular with acute apex. Mesofurca X shaped, metafurca each separated longitudinally, both furca curved, anteriorly deeper. Legs (Fig. 62a, b). Hindlegs surpassing abdomen and slightly longer than forelegs, midlegs slightly shorter than forelegs. All five carinae gently elevated and marked by rows of short dark setae. Femora slightly widened apically. Mesobasitarsi about the same size than following three tarsomeres combined, pro- and metabasitarsi slightly longer than following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 62a, b, 63j, k, n-p). Densely granulated, covered in short small black setae. Terga I and IV–VII longer than wide, II slightly wider than long, III and VIII–X about as long as wide. Terga III–VII bearing medial projections at posterior margin, in some specimens also on VIII; projections on IV–VI slightly conical, pointing posteriad, also the one on VII when a further projection is present on VIII; on III and VII or VIII round, smaller. Terga VII–VIII further bearing conical projections on posterior margin of lateral carinae. Terga VIII and IX with a medial slight elevation. Tergum X with medial carina at posterior half, eventually on all length of the tergum, posterior margin gently carinated, trilobed with two paramedian round short lobes and a medial short, slightly pointed lobe, slightly widening towards lateral edge. Preopercular organ longer than wide, rugose, black, convex, anteriorly widening. Subgenital plate wider at mid length, then tapering to a lanceolate posterior margin, reaching from the base of the cerci to the posterior margin of tergum X. Epiproct not surpassing tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci short, conical, 2.5–2.6 × longer than wide, not or barely projecting from the posterior margin of tergum X (Fig. 63j, n). Gonapophysis VIII long and narrow, tapering towards apex, apically setose, apex thin; significantly longer than gonapophysis IX (Fig. 63k). Gonapophysis IX wide, triangular; basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin wider and with a wide incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 63k). Gonoplac incurved, gently widened basally, apex round, slightly shorter than gonapophysis IX (Fig. 63k).

Measurements (MZ1130): body (without cerci) 88.2, head 6.3, antennae at least 60.0, pronotum 6.8, mesonotum 15.4, metanotum 5.5, median segment 7.1, abdomen (excluding median segment, without cerci) 47.1, profemur 27.0, protibia 27.2, mesofemur 20.3, mesotibia 20.1, metafemur 28.4, metatibia 30.1.

Measurements, variation (N = 20, all paratypes): body (without cerci) 68.7–86.2, head 5.3–6.4, pronotum 5.2–7.0, mesonotum 12.2–15.6.

Male. Similar to that of the female (Fig. 62c-e), except: head (Fig. 63a, c, f). Narrower, eyes much larger, scapus and pedicel larger, third antennomere longer than pedicel and scapus combined. Thorax (Figs. 62c-e, 63a-c, f). narrower. Legs (Fig. 62c-e). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 62c-e, 63b, d, e, f, g, h). Terga I–VII and IX longer than wide, VIII and X wider than long to as long as wide. Terga VI–VIII further bearing conical projections on posterior margin of lateral carinae. Posterior margin of tergum X emarginated, thorn pads pointing inward, near median line, bearing around 12 wide teeth of similar size. Epiproct hidden dorsally. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin gently broadly emarginated; bearing a large dextral process with anterior half widened, with ventral margin round and prominent and posterior half linear, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci longer, of more uniform width only slightly conical, 2.7–3 × longer than wide, apex very slightly curved (Fig. 63g, h). Vomer very large, wide, strongly tapering, terminal hook sharp, small, upwardly curved.

Measurements (MZUSP 0184): body (without cerci) 60.0, head 3.8, antennae 67.8, pronotum 4.0, mesonotum 10.4, metanotum 4.0, median segment 3.9, abdomen (excluding median segment, without cerci) 33.9, profemur 21.6, protibia 23.6, mesofemur 16.8, mesotibia 18.4, metafemur 22.0, metatibia 25.3.

Measurements, variation (N = 31, all paratypes): body (without cerci) 50.9–59.8, head 3.4–4.3, pronotum 3.8–4.5, mesonotum 9.4–10.7.

Egg. unknown.

Materials examined

Holotype, m#: Brazil, Pará, Itaituba, Rio Tapajós. iv.1961. Dirings col. (MZUSP 0184).

Paratypes, 6 m# (MZUSP 1128–1129, 2261, 2263–2265), 5f# (MZUSP 0186, 1130–1131, 2266, 2267): Brazil, Amazonas, Itacoatiara. iv.1964. [Dirings col.]; 1 m# (MZUSP 2262): Brazil, Amazonas, Itacoatiara. [Dirings col.]; 1f#: Brazil, Pará, Xinguara, Rio Vermelho, iii.1964 [Dirings col.] (MZUSP 1424); 1 m#: Brazil, Pará, Santarém, iv.1961 [Dirings col.] (MZUSP 1425); 1f# (MZUSP 1428), 2 m# (MZUSP 1127, 2257): Brazil, Pará, Itaituba. [Dirings col.]; 1 m#: Brazil, Pará, Itaituba, Camargo, Rio Tapajós. [Dirings col.] (MZUSP 2260); 1 m#: Brazil, Pará, Itaituba, Rio Tapajós, i.1961 [Dirings col.] (MZUSP 2253); 4 m# (MZUSP 0187, 0190, 1134, 2244): Brazil, Pará, Itaituba, Rio Tapajós, Santarenzinho. iv.1961. Dirings col.; 4f# (MZUSP 2236–2239), 1 m# (MZUSP 2243): Brazil, Pará, [Itaituba], Santarenzinho. v.1961. Dirings col.; 1f#: Brazil, Pará, Itaituba, ii.1964 [Dirings col.] (MZUSP 2259); 1 m#: Brazil, Pará, Itaituba, Rio Tapajós, iv.1963 [Dirings col.] (MZUSP 0185); 1f#: Brazil, Pará, Itaituba, Santarenzinho, iv.1963 [Dirings col.] (MZUSP 1133); 2f# (MZUSP 2240, 2281), 2 m# (MZUSP 2241, 2242): Brazil, Pará, Itaituba, Santarenzinho, iii.1961 [Dirings col.]; 1f#: Brazil, Pará, Itaituba, Santarenzinho, viii.1961 [Dirings col.] (MZUSP 2235); 1 m#: Brazil, Pará, [Itaituba], Santarenzinho, iii.1964 [Dirings col.] (MZUSP 2245); 1 m# (MZUSP 1124), 1f# (MZUSP 1123): Brazil, Pará, Itaituba, Camargo, Rio Tapajós. iv.1963. [Dirings col.]; 1 m#: Brazil, Pará, Itaituba, Rio Tapajós, iv.1964 [Dirings col.] (MZUSP 2258); 4 m#: Brazil, Pará, Itaituba, vi.1964 [Dirings col.] (MZUSP 1125, 2246–2248); 4f# (MZUSP 1126, 2249, 2250, 2256), 6 m# (MZUSP 0188, 0189, 2251, 2252, 2254, 2255): Brazil, Pará, Itaituba, Rio Tapajós, iii.1964 [Dirings col.].

Etymology. The specific epithet is an adjective and refers to the known distribution of the new species in the Amazon, occurring east of the Madeira river and south of the Amazon river. The biogeography of the Amazon is often divided into areas roughly corresponding to interfluves, mainly delimited by the Amazonas, Madeira and Negro rivers as already recognized by earlier biogeographers (Wallace 1854). The new species is unable to fly and thus could be more liable to be isolated by large rivers.

Differential diagnosis. Differs from remaining Urucumania by the dorsally orange, laterally black and posteriorly darker color pattern (Fig. 62), the fine granulation on the body surface (Fig. 63), the third antennomere more elongate, as long as pedicel and scapus in females and longer than pedicel and scapus in male, the slightly elongated alae (Fig. 63b, o), the terga I–II without elliptical dorsal projections (Fig. 62b, e), the female conical cerci (Fig. 63p), and the basally widened right process of sternum IX of males (Fig. 63h).

Distribution (Fig. 68). Only known from Itaituba and Xinguara, Pará and Itacoatiara, Amazonas, Brazil, in the Amazon.

Urucumania candanga Ghirotto & Sisnando sp. n.

(Figs. 64, 65, 66 and 67)

Fig. 64

Urucumania candanga sp. n., paratype females from Brasília, Distrito Federal, Brazil, in dorsal (a, b) and lateral (c) views. Scale bars = 2 mm

Fig. 65

Urucumania candanga sp. n., holotype male from Brasília, Distrito Federal, Brazil, in dorsal (a,) and lateral (b) views. Scale bars = 2 mm

Fig. 66

Urucumania candanga sp. n., holotype male (a–e, g–h) and paratype females (f, i–o) from Brasília, Distrito Federal, Brazil. a, c, f, i, l Head and pronotum in dorsal (a, f, i) and lateral (c, l) views. b, n Metathorax in dorsal view. d, e, g, h, j, m, o Terminalia in dorsal (d, j), left side (g, m), right side (h) and ventral (e, o) views. k Gonapophyses in lateral view. Scale bars = 1 mm

Fig. 67

Urucumania candanga sp. n., egg of paratype female from Brasília, Distrito Federal, Brazil. a Lateral view. b Dorsal view. c Anterior or opercular view. Scale bars = 0.5 mm

Description

Female. Color (Fig. 64). Head and body with a dorsal beigeish orange band and light to dark brown on the remaining dorsal areas and on all lateral and ventral areas, with several scattered whitish granules varying in density, denser on the abdomen but scarce on terga VIII–IX and almost absent on X; ventrally slightly lighter and with much scarcer whitish granules. Head and body with an orange beigeish wide medial band running along the head, pronotum, mesonotum, metanotum and terga I–VIII and sometimes also on IX–X, slightly duller on head and VIII–X. Dorsally bordering the medial band the body is slightly darker than near lateral margins, and terga I–VIII display at the posterior margin orange beigeish round stains almost touching the band. Legs dorsally and laterally dark, dark brownish red basally in femora and almost all length of tibiae expect apex, rest of femora, apex of tibiae and all tarsi blackish; ventrally lighter, with irregular creamish stains.

Head (Fig. 66f, i, l). Large, approximately subrectangular, around 1.3 × longer than wide, 1.4 × longer than high, slightly longer than pronotum, sparsely tuberculated. Vertex flat, frontal area gently elevated. Frontal suture gently curved and deep, frontal convexity prominent. Eye somewhat medium sized, round, less than hemispherical. Clypeus wide, convex, with shallow depressions, labrum slightly notched anteromedially, round. Antennae filiform with 24–25 fully segmented antennomeres; from 16–17th onwards, each segment is further subsegmented into 2–7 less articulated subsegments of equal width, more conspicuous apically; bearing small setae on all surface. Scapus smooth, slightly longer than wide, subcylindrical, flattened dorsoventrally only at basal portion; pedicellus subcylindrical, slightly shorter than scapus. Antennal bump small and discrete, on the 18th segment. Thorax (Figs. 64, 66f, i, l, n). Cylindrical in cross section, covered in short small black setae, bearing lateral carinae, with several small granules. Pronotum subrectangular, slightly tapering towards anterior; sulci marked, longitudinal straight and transversal slightly sinuous; lateral edges marked with a conspicuous smooth carina; prothoracic gland openings large, delimited by elevated carinae, with a deep sulcus posteriorly. Pronotum with granules almost absent from central region. Mesonotum bearing on each side a paramedian row of conical to cylindrical granules on anterior three fifths and some smaller granules posteriorly, and a further row of smaller round granules at anterior three fifths at each lateral margin. Metanotum slightly shorter to slightly longer than median segment. Metanotum bearing two laterodorsal granules at each side. Mesepisternum and metepisternum also bearing row of granules. Fore and hindwings diminute, discrete, scale-like; forewing round, hindwing elliptical. Meso and metafurca each separated longitudinally, curved, anteriorly deeper. Legs (Fig. 64). Hindlegs surpassing abdomen and slightly longer than forelegs, midlegs slightly shorter than forelegs. All five carinae elevated and marked by rows of short setae. Femora slightly thinner basally. Basitarsi slightly longer than the following three tarsomeres combined. Area apicalis delimited by a somewhat deep sulcus and bearing tuft of setae. Euplantulae well developed in all tarsomeres. Abdomen (Figs. 64, 66j, k, m–o). Tuberculated, with scattered larger granules, covered in short small black setae. Terga I–X wider than long. Terga I–VIII bearing short round projections pointing posteriad, one medially and two paramedial at posterior margin, and one in each lateral carinae except for tergum I without lateral projections; IX with a short round projection medially at posterior margin, shorter than previous. Terga IX–X with scattered paramedial granules. Tergum X with medial discrete carina at posterior half, posterior margin somewhat widely round and gently carinated, but widening towards lateral edge. Preopercular organ longer than wide, wrinkled and rugose, black, convex, somewhat triangular, posteriorly widening. Subgenital plate wider at mid length, then slightly tapering to a round to slightly roundly lanceolate posterior margin, reaching half the length tergum X. Epiproct under tergum X, wide, round, with a medial carina. Paraprocta apically apart, inner margins short. Cerci relatively elongate (2.6–2.7 × longer than wide), cylindrical, posteriorly slightly tapering (Fig. 66j, o). Gonapophysis VIII long, somewhat triangular, tapering towards apex, slightly widened basally, apically setose, about the same length as gonapophysis IX (Fig. 66k). Gonapophysis IX basally and subapically setose, ventral margin with a narrow smooth keel outwardly curved along its edge; inner margin soft, ending in a posterior narrow prominence protruding distinctly; outer margin slightly wider and with an incurved sclerotization from the base of gonoplac to before the prominence of the inner margin (Fig. 66k). Gonoplac incurved, gently widened basally, apex round, longer than both gonapophyses (Fig. 66k).

Measurements (paratype MZUSP): body (without cerci) 50.7, head 4.0, antennae 28.5, pronotum 3.8, mesonotum 9.4, metanotum 2.8, median segment 3.5, abdomen (excluding median segment, without cerci) 27.2, profemur 11.4, protibia 10.5, mesofemur 9.4, mesotibia 8.8, metafemur 12.9, metatibia 12.8.

Measurements, variation (N = 1, paratype UNB 104237): body (without cerci) 46.9, head 3.8, pronotum 3.0, mesonotum 8.3.

Male. Similar to that of the female, except: color (Fig. 65). Orange band slightly brighter and almost fading on tergum VIII; rest of body darker, dark brown to blackish, whitish granules scarcer, sparse; round stains on terga absent; legs ventrally the same as dorsally. Head (Fig. 66a, c). Narrower, eyes larger, scapus and pedicel larger, antennae with around 22 segments. Thorax (Figs. 65c, d, 66a-c). Narrower, granules of mesonotum and metanotum wider, conical. Legs (Fig. 65c, d). Setae in carinae longer, distributed in fewer rows; basitarsi relatively longer. Abdomen (Figs. 66c, d, b, d, e, g, h). Narrower, terga I, IV–VI longer than wide, II–III, VII, IX as long as wide, VIII and X wider than long. Posterior margin of tergum X gently emarginated, thorn pads pointing inward, near median line, bearing around 10 teeth. Epiproct not exceeding posterior margin of tergum X. Sternum IX voluminous, with large anterior portion and very large poculum, posterior margin almost straight, slightly curving towards anterior; bearing a large dextral process widened at mid length then slightly narrowing towards posterior, slightly upcurving, gently laterally flattened, originating superiorly from the anterior portion of sternum IX. Cerci moderately long (3.2 × longer than wide), cylindrical, very slightly inwardly curved (Fig. 66d, g, h). Vomer very large, wide, strongly tapering, terminal hook small, acute, upwardly curved.

Measurements (holotype): body (without cerci) 34.4, head 2.6, antennae 28.2, pronotum 2.3, mesonotum 6.0, metanotum 2.2, median segment 2.6, abdomen (excluding median segment, without cerci) 18.7, profemur 9.4, protibia 9.4, mesofemur 7.3, mesotibia 7.3, metafemur 10.3, metatibia 10.2.

Egg (Fig. 67). Irregularly dark blueish gray, micropylar plate creamish. Capsule barrel-shaped, higher than wide, longer than wide in variable ratios, slightly wider anteriorly, oval in cross section; polar area somewhat flat. Operculum perpendicular, oval and flat. Capsule covered by net of irregular, moderately thick rugose ridges, forming a circular ridge on the operculum. Ridges further present minute mushroom-like structures also on the operculum. Micropylar plate slightly elongate, elliptical, rugose, with elevated edges; centrally and posteriorly bearing an elongate rugose hump widened anteriorly, connected to the micropylar cup. Micropylar cup distinct. Median line moderately thick, slightly less than half the length of the micropylar plate. Capsule presenting a short furrow below the median line reaching the polar area. Measurements (N = 10): length 2.7–3.0, width 1.5–1.7, micropylar plate length 1.0–1.3, operculum maximum diameter 1.3–1.6.

Materials examined

Holotype, 1 m#: Brazil, Distrito Federal, Brasília, ARIE Capetinga, Cerrado campo sujo, 15°56′53.7"S 47°57′33.6"W, 24.x.2023, P. Sisnando, rede de varredura. (MZUSP).

Paratypes, 1f#, 20 eggs: Brazil, Distrito Federal, Brasília, ARIE Capetinga, Cerrado campo sujo, 15°56′53.7"S 47°57′33.6"W, 24.x.2023, P. Sisnando, rede de varredura (MZUSP); 1f#: Brazil, Distrito Federal, Brasília, ARIE Capetinga, Cerrado savânico, 15°57′27.0"S 47°55′55.2"W, 24.i.2024, P. Sisnando (UNB 104237).

Etymology. The specific epithet is an adjective from Portuguese meaning inhabitant of Brasília, the type locality.

Differential diagnosis. Differs from remaining Urucumania by the dark-orange color pattern with dark legs (Figs. 64, 65), the elongate terga VIII–X in both sexes (Fig. 66d, g, j, m), the widened dextral process of sternum IX of males (Fig. 66h), and elongated cerci of the female (Fig. 66j). Females further differ from Urucumania tapirape sp. n., another species with elongated terga VIII–X, by the shorter head (Fig. 66a, f, i) and terga VIII–X (Fig. 66j, m) and distinct color pattern (Fig. 64).

Distribution (Fig. 68). Only known from Brasília, Distrito Federal, Brazil, in savannic areas of the Cerrado.

Discussion

With the 13 new species described here and the additional two species transferred from Pseudophasma, Urucumania now contains 17 species, representing an increase of 850% in the number of species. The high diversity revealed in this research highlights the potential for discovering new species of stick insects, especially in poorly sampled areas, as well as the need of revision for several groups of stick-insects. Urucumania is a more diverse genus than previously thought, now known to be broadly distributed in Brazil, occurring in dry to humid, open to forested habitats. Moreover, nine out of the 13 new species derive from recently collected material, mostly from active efforts of Projeto Phasma to sample phasmids specifically, also counting on the help of third parties. This shows that even few resources dedicated to sampling can result in the discovery of high amounts of new phasmid species, greatly contributing to reducing the taxonomic impediment (Engel et al. 2021). Preliminary observation by the authors further suggest many more undescribed species of Urucumania which would be discovered with increased sampling effort; for example, note the wide gaps in the known records for Urucumania in Fig. 68. Some unsampled populations from the South of Brazil show much similarity to one another, as seen for example in pictures taken by citizens of unsampled material. Studies focusing on similar Urucumania representatives would be a welcoming effort on future studies on the genus, especially if including in other analyses such as the use of molecular work. The genus Pseudophasma still includes several heterogeneous species and is in need of revision (see also Chiquetto-Machado 2018 and Robertson et al. 2018), with the expectation that more of its species will be transferred to other genera.

Fig. 68

Distribution map for Urucumania in South America. Brazil is further divided in States, which are shown with a thinner line