Paraphasma marginale Redtenbacher, 1906

Chiquetto-Machado, Pedro I., Morales, Adriana C. & Cancello, Eliana M., 2022, Taxonomic revision of Paraphasma Redtenbacher, 1906 (Phasmatodea, Pseudophasmatidae) based on phallic and external morphology, Zootaxa 5122 (1), pp. 1-80 : 31-36

Paraphasma marginale Redtenbacher, 1906

Figs 20–25 , Table 5 .

Paraphasma marginale Redtenbacher, 1906: 115–116 ; Chopard, 1911: 338 [probable misidentification]; Piza, 1939: 116 [probable misidentification]; Klante, 1960: 90; Weidner, 1966: 231; Hennemann et al., 1995: 438; Brock, 1998a: 41 [lectotype designation]; Zompro, 2000: 95 [designation as type species of Paraphasma ]; Zompro, 2002a: 192; Zompro, 2002b: 6 [repetition of designation as type species of Paraphasma ]; Zompro, 2003: 39; Hennemann & Conle, 2003: 6; Zompro, 2004: 159, figs 9.9, 92a,b; Otte & Brock, 2005: 251; Brock, 2007: 51 ; Delfosse et al., 2019: 212 ; ChiquettoMachado & Cancello, 2021: 4, 23, 26, figs 5, 15C,D, 25, 26. Lectotype: ♂, Paraguay, Puerto 14 de Mayo ( MSNG) . Paralectotypes: 1♂, 2♀, Paraguay, Puerto 14 de Mayo ( MSNG) ; 1♂, 1♀, Paraguay, Puerto 14 de Mayo ( NHMW *) ( Figs 20 , 23 ) ; 2♂, 1♀, Paraguay, San Bernardino ( NHMW) ; 1♀, Paraguay ( NHMW) ; 3♂, Brazil, Goiás, Jataí ( NHMW *) ; 1♂, Brazil, Mato Grosso, Cuiabá ( NHMW) ; 1♂, Brazil, São Paulo ( NHMW) ; 1♀, Brazil ( NHMW) ; 1♂, Brazil ( ZIRAS) ; 1♂, “ Schrerik’s Reise ” ( ZIRAS) ; 2♂, Paraguay ( ZMUH) ; 1♂, 1♀, Brazil, Goiás ( ZMUH) ; 2♂, Brazil, Santos ( ZMUH) ; 4♂, Brazil, Goiás, Jataí ( MNHN) ; 1♂, Brazil, Minas Gerais, Caraça ( MNHN) ; 4♂, Brazil, Rio de Janeiro ( MNHN) ; 2♂, Brazil ( MNHN) ; 1♂, Brazil, Rio de Janeiro ( SMNS) ; 2♂, no locality data ( SMNS) ; number of specimens and localities unknown for the paralectotypes at MNCN and SMTD.

Diagnosis. Similar to P. conspersum , P. indistinctum sp. nov., P. laterale , P. minus and P. sooretama sp. nov., but clearly distinguishable by the phallic organ, with a “large type ” sclerite of the ventral lobe ( Fig. 22 , in green), which is roughly semicircular in dorsal view and has three protuberances: the apical one ( Fig. 22 : ap), which is large and rounded, the left basal one ( Fig. 22 : lp), undivided and strongly protruding, and an inconspicuous right basal protuberance ( Fig. 22 : rp), restricted to a gentle bulge. Although not exclusive of P. marginale , the following features may also be useful for the identification of this species: tegmina relatively long for the genus (at least 2.5x longer than wide) ( Figs 20D , 23D , 25C,D ), with non-spiniform shoulder pads and rounded margins; male cerci with spatulate apex ( Fig. 21A–D ); vomer longer than wide ( Fig. 21E ); male subgenital plate with posterolateral margins forming a pair of approximately triangular expansions with malleable aspect ( Fig. 21 : arrows); female sternite VII with a rounded indentation on the posterior margin; live specimens cryptic-colored ( Fig. 25 ), with body mostly pale brown or pale orange and a pair of pale yellow lateral stripes.

Redescription of male. Color ( Figs 20 , 21 , 25A ): Live specimens with cryptic color pattern ( Fig. 25A ), with a pair of pale yellow lateral stripes extending along head, prothorax, mesothorax and costal region of tegmina and hindwings; rest of body pale brown or pale orange with black regions. Head, pro- and mesonotum with brown or orangish dorsomedian line; pro- and mesonotum black, with spots in the same color as dorsomedian line. Legs black or brown; usually darkening towards apex of femora and tibiae.Anal region of hindwing orangish; colorless in dried specimens. Body ventrally brown; subgenital plate partly or entirely black. Head, thorax and legs ( Fig. 20A–D ): As in Paraphasma conspersum . Wings ( Fig. 20A–D ): Tegmina short, not reaching median region of metanotum; in dorsal view 2.5–3.3x longer than wide; posterior and apical margins rounded; shoulder pad as a dull, nonspiniform protuberance; anal region with conspicuous reticulate venation. Hindwing reaching abdominal tergite VII. Abdomen ( Figs 20A–C , 21 ): As in Paraphasma conspersum , except for vomer with narrow basal region ( Fig. 21E ). Phallic organ ( Fig. 22 ): Dorsal sclerite wider than long, roughly rectangular in dorsal view ( Fig. 22 , in red); distal process fairly short and wide, directed to the left, almost perpendicular in relation to longitudinal axis of organ ( Fig. 22 : dp). Dorsal and ventral lobes partially fused on left side. Dorsal lobe ( Fig. 22 : dl) subdivided into main body on the right and a smaller, rounded dorsal pouch on the left ( Fig. 22 : asterisks). Sclerite of the ventral lobe of “large type ” ( Fig. 22 , in green), covering entire inner face of ventral lobe and slightly extending to outer face on right side; roughly semicircular in dorsal view, with rounded posterior margin; apical and left basal protuberances well-developed ( Fig. 22 : ap, lp), the former large and rounded, the latter undivided and strongly protruding; right basal protuberance inconspicuous ( Fig. 22 : rp), restricted to a gentle, rounded bulge. One of base apodemes ( Fig. 22 , in blue) projecting into dorsal lobe as a distinct spatulate expansion.

Redescription of female. Color ( Figs 23 , 25B–D ): As in male, but subgenital plate varying from light brown to predominantly black. Head and thorax ( Fig. 23A–D ): As in male. Legs ( Fig. 23A–D ): As in male, but slightly shorter in relation to body and anterodorsal carina of profemur distinctly raised medially ( Fig. 23D : arrow). Wings ( Fig. 23A–D ): As in male, but hindwing reaching or surpassing abdominal tergite VIII. Abdomen ( Fig. 23A–C,E– G ): As in Paraphasma conspersum .

Description of egg ( Fig. 24 ). External morphology as in Paraphasma conspersum , but capsule with less and shorter bristles. Internal micropylar plate open ( Fig. 24D ); oval and elongate; internal median line distinct ( Fig. 24D : black arrow), more than half as long as plate. Egg in varying shades of brown, often grayish-brown with darker stains. Measurements (mm, n = 8): capsule length, 2.7–3.2; capsule width, 1.6–1.8; capsule height, 1.8–1.9; operculum width, 1.0–1.2; operculum height, 1.2–1.4; micropylar plate length, 0.9–1.2; micropylar plate width, 0.5–0.6.

Distribution ( Fig. 1 : white circles). Paraphasma marginale is widely distributed throughout the Cerrado of Central-West Brazil, also occurring in Cerrado-Amazon transition zones (central and northeastern Mato Grosso; southeastern Rondônia) and Cerrado-Atlantic Forest transition zones (southwestern Minas Gerais). It is also recorded from the Brazilian and Paraguayan Pantanal (there is a record from Cáceres, Mato Grosso, besides the type locality, in northeastern Paraguay close to the borders with Brazil and Bolivia).

Remarks. For the description of Paraphasma marginale, Redtenbacher (1906) examined a large series of specimens deposited in different collections (NHMW, MSNG, ZIRAS, ZMUH, MNHN, SMNS, MNCN, SMTD), from various localities in Brazil and Paraguay, which very likely includes individuals of different species. Brock (1998a) designated as the lectotype a male from northeastern Paraguay (label data: “ Paraguay, Puerto 14 de Mayo, i.1897, leg. G. Boggiani”) housed at MSNG. We examined this specimen only through an image available in Brock et al. (2022), but one male and one female paralectotypes ( Figs 20 , 23 ) with label data identical to those of the lectotype were studied by PICM at NHMW.

The paralectotypes of P. marginale that are housed in institutions other than the NHMW were examined only through photographs or were not examined. Of the NHMW specimens, only two had their phallic organs dissected for analysis, in order to minimize the damage to type material. As a consequence, for many paralectotypes the conspecificity with the lectotype remains uncertain and is unlikely, especially in view of the very diverse collecting localities and some differences of external morphology among the specimens. Paralectotypes from the following localities were not confirmed as conspecific with the lectotype: “Paraguay”; “Paraguay, San Bernardino”; “Brazil”; “Brazil, São Paulo”; “Brazil, Santos”; “Brazil, Rio de Janeiro”; “Brazil, Minas Gerais, Caraça”. No information could be obtained regarding the number of individuals and collecting localities of the paralectotypes housed at MNCN and SMTD.

One male from Argentina (locality: “Pto. Bemberg, Misiones”) deposited at the Museo Argentino de Ciencias Naturales (Bueno Aires, Argentina) was identified as P. marginale by Piza (1939), but examination of the specimen would be necessary to confirm this identification. In fact, considering that the collecting locality is an area of Atlantic Forest in northeastern Argentina (indicated by a star in Fig. 1 ), it is likely that this specimen belongs to Paraphasma laterale . This is the only record of Paraphasma from Argentina found in the literature or in the material examined by us.

Additional material examined. BRAZIL. Goiás: 1♂, Chapada dos Veadeiros, Alto Paraíso de Goiás, Trilha da Cachoeira do Segredo , 14º15’37”S, 47º52’11”W, 31.xii.2016 – 1.i.2017, P. I. Chiquetto-Machado, A. Z. Ramin ( MZUSP 0758 *) ; 2♀, Formosa, Distrito do Bezerra, Fazenda Santo Antônio , -15.30755, -47.19584, 28.i–5.ii.2012, Excursão Disciplina Entomologia de Verão (UnB) . Mato Grosso: 1♀, Barra dos Bugres, R. E. Serra das Araras , 20.i.1986, Marcio Zanuto ( MPEG 19000228 ) ; 1♀, Buriti , 15.ii.69, N. Tangerioni ( DZUP 380005 ) ; 1♀, Cáceres , 17.xi.1984, Buzzi, Mielke, Elias, Casagrande, Proj Polonoroeste ( DZUP 380011 ) ; 1♀, Chapada de Guimarães , 30.i.65, Sebastião Laroca ( DZUP 380004 ) ; 1♀, Chap Guimarães , 25.xi.1983, Exc Dep Zool, Polonoroeste ( DZUP 380008 ) ; 1♀, 1 egg, Chap Guimarães , 3.xii.1983, Exc Dep Zool, Polonoroeste ( DZUP 380009 ) ; 3♀, Chapada dos Guimarães, Fazenda Buriti , 18.xi.1982, Márcio Zanute, W. Overat ( MPEG 19000225 , 19000226 , 19000227 ) ; 1♀, Cuiabá, Aguaçu , 15.xi.1992, Paula Polon ( UFMT) ; 1♀, 1 egg, Diamantino, Intersecç BR163 c o Rio Arinos, 20.x.83, J. Becker ( UEFS 38373 ) ; 1♀, Mun Livramento, 12.xii.92, Airton Reis ( UFMT) ; 2♂, 2♀, Tapirapés, Carvalho ( MNRJ *) . Minas Gerais: 9♀, 40 eggs, Delfinópolis, Camping Claro Casa de Pedra , 20º20’45”S, 46º48’18”W, 28.xii.2016, P. I. Chiquetto-Machado, A. Z. Ramin ( MZUSP 0576 , 0757 , 0772 , 0773 , 0774 , 0775 , 0853 , 0854 , 0855 ) . Rondônia: 6♂, 2♀, Vilhena, Polo Noroeste , 17.xii.1986, C. Elias ( DZUP 379998 , 379999 , 380000 , 380023 *, 380024 , 380034 , 380035 *, 380036 ) .

Figures

FIGURE 1. Distribution map of Paraphasma. Circles indicate records based on material examined; triangles indicate records from the literature (Stoll 1787; Serville 1839; Chopard 1911; Conle et al. 2011; Jourdan et al. 2014); stars indicate records of undetermined Paraphasma species based on Rehn (1906) and Piza (1939) (although these authors identified the specimens, the identifications are doubtful or insufficient).

FIGURE 20. Paraphasma marginale Redtenbacher, 1906, male, paralectotype (NHMW). A–C. Habitus, dorsal (A), lateral (B) and ventral (C) views. D. Anterior region of the body, dorsolateral view. E. Labels. Scale bars = 10 mm.

FIGURE 21. Paraphasma marginale Redtenbacher, 1906, male terminalia. A–D. Terminalia in dorsal (A), lateral (B), ventral (C) and oblique caudal (D) views. E. Drawing of the vomer. Abbreviations: Ce, cerci; S8, sternite VIII; SPa, anterior region of the subgenital plate; SPp, posterior region of the subgenital plate; T8, T9, T10, tergites VIII, IX and X; TP, thorn pad. Arrows = expansion of the posterolateral margin of the subgenital plate. Scale bars = 1 mm.

FIGURE 22. Paraphasma marginale Redtenbacher, 1906, illustrations of the phallic organ. A. Dorsal view. B. Detail of the sclerite of the ventral lobe in dorsal view. C. Caudal view. D. Left view. E. Right view. Abbreviations: ap, apical protuberance of the sclerite of the ventral lobe; b, base; dl, dorsal lobe; dp, distal process of the dorsal sclerite; lp, left basal protuberance of the sclerite of the ventral lobe; rbp, right basal pouch; rp, right basal protuberance of the sclerite of the ventral lobe; vl, ventral lobe. Colors: light gray, membranous regions; red, dorsal sclerite; green, sclerite of the ventral lobe; blue, base apodemes. * = left ramification of the dorsal lobe. In D and E the final portion of the ejaculatory duct is represented by transparency with dotted lines. Images A, B, D and E adapted from Chiquetto-Machado & Cancello (2021: fig. 5).

FIGURE 23. Paraphasma marginale Redtenbacher, 1906, female, paralectotype (NHMW). A–C. Habitus, dorsal (A), lateral (B) and ventral (C) views. D. Anterior region of the body, dorsolateral view. E–G. Terminalia, dorsal (E), ventral (F) and lateral (G) views. H. Labels.Abbreviations: Ce, cerci; PO, praeopercular organ; SP, subgenital plate; T8, T9, T10, tergites VIII, IX and X. Arrow = anterodorsal carina of profemur. Scale bars = 10 mm, unless otherwise indicated.

FIGURE 24. Paraphasma marginale Redtenbacher, 1906, egg. Dorsal (A), lateral (B) and anterodorsal (C) views, and drawing of the internal micropylar plate (D). Abbreviations: Ca, capsule; Op, operculum; MP, micropylar plate. White arrow = median line; black arrow = internal median line.

FIGURE 25. Paraphasma marginale Redtenbacher, 1906, live specimens. A. Adult male from Alto Paraíso de Goiás (Goiás, Brazil), dorsolateral habitus. B–D.Adult female from Delfinópolis (Minas Gerais, Brazil), lateral habitus (B) and anterior region of the body in dorsal (C) and dorsolateral (D) views.