Taxonomic revision of Paraphasma Redtenbacher, 1906 (Phasmatodea, Pseudophasmatidae) based on phallic and external morphology Author Chiquetto-Machado, Pedro I. 0000-0002-3966-3202 pedroivocm@gmail.com Author Morales, Adriana C. 0000-0002-2108-427X adriana.morales@unesp.br Author Cancello, Eliana M. 0000-0003-3125-6335 ecancell@usp.br text Zootaxa 2022 2022-03-30 5122 1 1 80 http://dx.doi.org/10.11646/zootaxa.5122.1.1 journal article 289470 10.11646/zootaxa.5122.1.1 05b411d8-94fd-4bec-8aa8-4b8b5f7daed8 1175-5326 6399540 EC13A69D-D6FA-4926-AC59-648A5626C9B9 Paraphasma laterale ( Fabricius, 1775 ) Figs 11–19 , Table 4 . Mantis lateralis Fabricius, 1775: 275 ; Fabricius, 1787: 227 ; Gmelin, 1789: 2054 ; Olivier, 1792: 634 ; Fabricius, 1793: 15 ; Lichtenstein, 1796: 78 [in part]; Zimsen, 1964: 615 . Holotype : ♀ , Brazil (NHM 012502929). Phasma lateralis , Fabricius, 1798: 188 ; Latreille, 1807: 87 ; Serville, 1831: 58 [in part]; Peletier de Saint Fargeau & Serville, 1825: 101 [in part]. Phasma laterale , Lichtenstein, 1802: 15 [in part]; De Haan, 1842: 123 [in part]; Gray, 1835: 22 [in part]; Westwood, 1859: 122 [in part]. Phocylides lateralis , Giglio-Tos, 1895: 806 [probable misidentification]. Olcyphides lateralis , Caudell, 1904: 186 [probable misidentification]. Prexaspes lateralis , Kirby, 1904a: 414 ; Brock, 1998b: 34 . Stratocles lateralis , Redtenbacher, 1906: 104 [in part]. Prexaspes ( Prexaspes ) lateralis , Otte & Brock, 2005: 282 ; Brock et al. , 2016: 180 . Paraphasma lateralis , Conle et al. , 2020: 70 , 126. Paraphasma laterale , Chiquetto-Machado & Cancello, 2021: 3–4 , 26–27, figs 3, 15F, 16A,D,G, 18A, 22C, 25, 26. Phasma hopii Gray, 1835: 25 ; Chiquetto-Machado & Cancello, 2021: 26 [= Paraphasma laterale ( Fabricius, 1775 ) ]. Lectotype (here designated): ♂ , Brazil (OUM). Paralectotypes (here designated): 2♀ , Brazil (NHM 012502945, 012502946). Phasma hopei , Westwood, 1859: 117 . Olcyphides hopii , Kirby, 1904a: 410 ; Rehn, 1907: 165 [probable misidentification]; Otte & Brock, 2005: 231 ; Brock et al. , 2016: 176 . Stratocles hopei , Redtenbacher, 1906: 107 . Phasma lineolatum Serville, 1839: 273 ; De Haan, 1842: 123 ; Westwood, 1859: 120 ; Kirby, 1904a: 410 [= Olcyphides hopii ]; Giglio-Tos, 1910: 10 [= Phasma fasciatum Gray, 1835 ]; Otte & Brock, 2005: 231 [= Olcyphides hopii ( Kirby, 1904a ) ]; Chiquetto-Machado & Cancello, 2021: 26 [= Paraphasma laterale ( Fabricius, 1775 ) ]. Unspecified type (s): Brazil [probably lost]. Phocylides lineolatus , Stål, 1875: 97 ; Giglio-Tos, 1910: 10 [= Phasma fasciatum Gray, 1835 ]. Stratocles dentatus Piza, 1937: 7–8 ; Piza, 1946: 155 ; Chiquetto-Machado & Cancello, 2021: 26 [= Paraphasma laterale ( Fabricius, 1775 ) ]. Holotype : ♂ , Brazil , Mato Grosso do Sul , Três Lagoas (ESALQ*) ( Fig. 11 ). Paraphasma dentatum , Zompro & Domenico, 2005: 257 ; Otte & Brock, 2005: 251 . Diagnosis. Similar to P. conspersum , P. indistinctum sp. nov. , P. marginale , P. minus and P. sooretama sp. nov. , but clearly distinguishable by the phallic organ, with a “small-simple type ” sclerite of the ventral lobe ( Fig. 14 , in green), which is roughly oval in dorsal view and has two rounded protuberances, the right one directed anteriorly and left one directed caudally. Although not exclusive of P. laterale , the following features may also be useful for the identification of this species: male cerci with rounded and internally concave apex ( Fig. 12E ) (a feature that is shared only with P. sooretama sp. nov. ); vomer longer than wide ( Fig. 13 ); male subgenital plate with posterolateral margins forming a pair of approximately triangular expansions with malleable aspect ( Fig. 12 : arrows); female sternite VII with a rounded indentation on the posterior margin; live specimens bright-colored ( Figs 18 , 19 ), with a marked contrast between the bright yellow lateral stripes and the mostly shiny black body. Also, contrary to most species of Paraphasma , the females of P. laterale sometimes lack a distinctly raised anterodorsal carina on the profemur. Redescription of male. Color ( Figs 11 , 12 , 18 ): Live specimens ( Fig. 18 ) with a pair of bright yellow lateral stripes extending along head, prothorax, mesothorax and costal region of tegmina and hindwings; rest of body mostly shiny black. Head, pro- and mesonotum with yellow or light brown dorsomedian line; pro- and mesonotum with spots in the same color as dorsomedian line. Legs black or dark brown; femora sometimes with longitudinal light brown stripes.Anal region of hindwing bright yellow. Body ventrally yellowish-brown, but abdominal sternites gradually darkening towards posterior end of body; subgenital plate mostly or entirely black. Dried specimens usually retain general color pattern ( Fig. 11 ), but less bright and with anal region of hindwing becoming colorless; in some specimens entire body becomes brownish. Head, thorax and legs ( Figs 11 , 18A–C ): As in Paraphasma conspersum . Wings ( Figs 11 , 18B ): Tegmina short, not reaching median region of metanotum; in dorsal view 2– 2.6x longer than wide; posterior and apical margins gently rounded; shoulder pad varying from dull protuberance to sharp spine; anal region with conspicuous reticulate venation. Hindwing reaching abdominal tergite VII. Abdomen ( Figs 11A–C , 12 , 13 ): As in Paraphasma conspersum , except for the following: apex of cerci rounded in lateral view and internally concave ( Fig. 12E ); vomer shape varying from a gradual narrowing towards apex to an abrupt constriction between apical and basal regions ( Fig. 13 ). Phallic organ ( Fig. 14 ): Dorsal sclerite wider than long, roughly rectangular in dorsal view, but anterior margin sometimes fairly concave ( Fig. 14 , in red); distal process short and wide, directed to the left, almost perpendicular in relation to longitudinal axis of organ ( Fig. 14 : dp). Dorsal and ventral lobes partially fused on left side. Dorsal lobe ( Fig. 14 : dl) subdivided into main body on the right and a smaller, rounded dorsal pouch on the left ( Fig. 14 : asterisks); inner face with a small sclerotized region ( Fig. 14 , dark gray). Sclerite of the ventral lobe of “small-simple type ” ( Fig. 14 , in green), covering only central region of inner face of ventral lobe; roughly oval in dorsal view, with right region broader than left region; bearing two rounded protuberances, the larger one on the right of sclerite and directed anteriorly, the smaller one on the left of sclerite and directed caudally. One of base apodemes ( Fig. 14 , in blue) projecting into dorsal lobe as a distinct spatulate expansion. FIGURE 11. Paraphasma laterale ( Fabricius, 1775 ) , male, holotype of junior synonym Stratocles dentatus Piza, 1937 (ESALQ). A–C. Habitus, dorsal ( A ), ventral ( B ) and lateral ( C ) views. D. Labels. E. Anterior region of the body, dorsolateral view. Scale bars = 10 mm. FIGURE 12. Paraphasma laterale ( Fabricius, 1775 ) , male terminalia. A–D. Terminalia in dorsal ( A ), lateral ( B ), ventral ( C ) and oblique caudal ( D ) views. E. Cerci, lateral view. 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 13. Paraphasma laterale ( Fabricius, 1775 ) , drawings of the vomer of two males, illustrating intraspecific variation. Scale bar = 1 mm. FIGURE 14. Paraphasma laterale ( Fabricius, 1775 ) , 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: b, base; dl, dorsal lobe; dp, distal process of the dorsal sclerite; rbp, right basal pouch; vl, ventral lobe. Colors: light gray, membranous regions; red, dorsal sclerite; green, sclerite of the ventral lobe; blue, base apodemes; dark gray, sclerotized region of the dorsal lobe. * = 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. 3). Redescription of female. Color ( Figs 15 , 19A ): As in male, but subgenital plate light brown or yellowish, lighter than preceding sternites. Head and thorax ( Fig. 15A–C,G ): As in male. Legs ( Fig. 15A–C,G ): As in male, but slightly shorter in relation to body and anterodorsal carina of profemur varying from weakly to distinctly raised medially. Wings ( Figs 15A–C,G , 19A ): As in male, but hindwing reaching or surpassing abdominal tergite VIII. Abdomen ( Fig. 15A–F ). As in Paraphasma conspersum . Description of egg ( Figs 16 , 17 ). External morphology as in Paraphasma conspersum , but capsule and operculum densely covered with uniformly distributed granules, each one originating a fairly long bristle, many of which are lost in preserved eggs. Internal micropylar plate open ( Fig. 16D ); oval and elongate; internal median line distinct ( Fig. 16D : black arrow), more than half as long as plate. Egg in varying shades of brown ( Fig. 17 ), often light brown with sparse darker regions. Measurements (mm, n = 10): capsule length, 2.8–2.9; capsule width, 1.8–2.0; capsule height, 1.9–2.1; operculum width, 1.1–1.2; operculum height, 1.2–1.5; micropylar plate length, 0.7–0.9; micropylar plate width, 0.4–0.5. FIGURE 15. Paraphasma laterale ( Fabricius, 1775 ) , female. A–C. Specimen MZUSP 0584, habitus, dorsal ( A ), ventral ( B ) and lateral ( C ) views. D–F. Terminalia, dorsal ( D ), ventral ( E) and lateral ( F ) views. G. Anterior region of the body, dorsolateral view. Abbreviations: Ce, cerci; PO, praeopercular organ; SP, subgenital plate; T8, T9, T10, tergites VIII, IX and X. Scale bars = 10 mm, unless otherwise indicated. FIGURE 16. Paraphasma laterale ( Fabricius, 1775 ) , 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 17. Paraphasma laterale ( Fabricius, 1775 ) , variation in egg color. Scale bar = 1 mm. Distribution ( Fig. 1 : pink circles). Paraphasma laterale occurs in the Brazilian Atlantic Forest in the states of Santa Catarina , Paraná , São Paulo , Rio de Janeiro and southern Minas Gerais , and is also recorded from a CerradoAtlantic Forest transition zone in eastern Mato Grosso do Sul state ( type locality of the junior synonym Stratocles dentatus ). Remarks. The holotype of P. laterale (locality: Brazil ) belongs to the Banks Collection of the NHM, a historic collection composed of Sir Joseph Banks’ material. This specimen was presumably collected during Captain James Cook’s first circumnavigation of the globe, when Banks had the opportunity to collect biological material in the region of Rio de Janeiro , the only Brazilian city visited in this voyage [for more details on the Banks Collection and James Cook’s expeditions, see the website of the NHM (http://www.nhm.ac.uk/our-science/collections/entomologycollections/historical-collections) and of the Captain Cook Society (http://www.captaincooksociety.com)].Therefore, in spite of the vague type locality and the very damaged condition of the holotype [images available in Brock et al. (2022) ], it is possible to assume that it was collected near the city of Rio de Janeiro , corroborating its conspecificity with the other specimens here assigned to P. laterale . This species was described by Fabricius (1775) as Mantis lateralis . It has an intricate taxonomic history and was often confused with Parastratocles xanthomela ( Olivier, 1792 ) , as explained below. Olivier (1792) proposed the binomen Mantis xanthomela for Stoll’s species ‘Spectre Noir de Surinam’ and ‘Spectre Noir de Coromandel’ ( Stoll 1787 : pp. 31–32, figs 36, 37). Later, these same species of Stoll (1787) were misidentified by Lichtenstein (1796) as Mantis lateralis Fabricius, 1775 . After M. lateralis was transferred to Phasma by Fabricius (1798) , several authors repeated the mistake of Lichtenstein (1796) , associating the two species of Stoll (1787) with the name Phasma laterale ( Lichtenstein 1802 ; Peletier de Saint Fargeau & Serville 1825 ; Serville 1831 ; Gray 1835 ; De Haan 1842 ; Westwood 1859 —some of these authors incorrectly attributed the name Phasma laterale to Lichtenstein or Stoll). Furthermore, Mantis xanthomela was mistakenly regarded as a junior synonym of Phasma laterale by Peletier de Saint Fargeau & Serville (1825) , Gray (1835) and Westwood (1859) , and also by Redtenbacher (1906) , who proposed the combination Stratocles lateralis . The transfer of this species to Stratocles was probably a consequence of Redtenbacher (1906) having examined the drawings of Mantis xanthomela Olivier in Stoll (1787) instead of the holotype of Mantis lateralis Fabricius , which clearly belongs to a different species. The name Stratocles lateralis remained valid until Brock (1998b) transferred the species to Prexaspes , reestablishing a combination originally proposed by Kirby (1904a) , Prexaspes lateralis . Mantis xanthomela was finally removed from the synonymy of Prexaspes lateralis by Otte & Brock (2005) , who revalidated the species as Stratocles xanthomela . Recently, both species were placed in new combinations by Conle et al. (2020) : Prexaspes lateralis was transferred to Paraphasma , which was corroborated by the phylogenetic hypothesis of Chiquetto-Machado & Cancello (2021) , and Stratocles xanthomela was transferred to Parastratocles Redtenbacher, 1906 . FIGURE 18. Paraphasma laterale ( Fabricius, 1775 ) , live specimens. A–C. Adult male from São Sebastião (São Paulo, Brazil), lateral habitus ( A ) and anterior region of the body in dorsal ( B ) and lateral ( C ) views. D. Adult male feeding on a leaf of Tabebuia cassinoides in an Atlantic Forest area in Bertioga (São Paulo, Brazil). FIGURE 19. Paraphasma laterale ( Fabricius, 1775 ) , live specimens. A. Adult female from Jardim Alviverde (São Paulo, Brazil), dorsolateral habitus. B. Last-instar male nymph from São Sebastião (São Paulo, Brazil), lateral habitus. Here we solve one last taxonomic problem regarding these species. Phasma perspicillaris Stoll, 1813 was synonymized under Stratocles lateralis by Redtenbacher (1906) , probably because of the similarities between the drawings of the types of Phasma perspicillaris (Stoll 1813: fig. 86) and Mantis xanthomela ( Stoll 1787 : figs 36, 37), which was also regarded by the author as a junior synonym of Stratocles lateralis . However, when Otte & Brock (2005) revalidated Mantis xanthomela (as Stratocles xanthomela ), these authors mistakenly left the name Phasma perspicillaris as a junior synonym of Prexaspes lateralis instead of synonymizing it under Stratocles xanthomela . We, therefore, remove Phasma perspicillaris from the synonymy of Paraphasma laterale and place it as a new synonym under Parastratocles xanthomela (see “Further taxonomic changes in Pseudophasmatidae ”). The types of both Parastratocles xanthomela and Phasma perspicillaris are from Suriname ( Stoll 1787 , 1813) and are probably lost. Additional material examined. BRAZIL . Minas Gerais : 1♂ , Poços de Caldas , Morro S Domingos , xi.1965 , J. Becker ( UEFS 38376 *) . Paraná : 1♂ , 1♀ , C Mourão , i.52 ( DZUP 380032 *, 380033 ) ; 1♀ , São Jorge , Copacabana , 15.xi.1975 , Rosado & Mielke ( DZUP 380013 ) ; 1♂ , Tapejara , i.53 ( DZUP 380031 ) . Rio de Janeiro : 2♂ , 1♀ , Barra de Itabapoana , x–xi.1935 , M. Rosa ( MNRJ *) ; 1♂ , 1♀ , Cabo Frio , i.1978 , Amilton Ferreira ( ESALQ *) ; 1♀ , Distrito Federal , Floresta da Tijuca , 25.i.1953 , C. A. Campos Seabra ( MNRJ ) ; 1♂ , Distrito Federal , Jacarepaguá , 10.xi.52 , Newton Santos ( MNRJ *) ; 1♂ , Itaguaí , Serra da Caveira , 600 m , 25.ii.1948 , W. Zikán ( MNRJ ) ; 1♂ , Organ Mts. near Tijuca , 1902–287, S. R. Wagner ( NHM 012502947 ) ; 2♀ , Petrópolis , Pr. Independência , 9.v. 929, Príncipe Paulo Gagarin ( MNRJ ) ; 2♂ , Petrópolis , Le Vallon , Alt. Mosélla , 24.i–23.ii.1956 , D’Albuquerque ( MNRJ ) ; 1♂ , Petrópolis , Taquara , 27.xii.70 , H. S. Lopes ( MNRJ ) ; 1♂ , Petrópolis , Taquara , 25.i.71 , H. S. Lopes ( MNRJ *) ; 2♂ , 2♀ , Serra dos Órgãos , xii.1940 , Parko ( MNRJ ) . Santa Catarina : 1♂ , Nova Teutônia , 4.i.1949 ( MZUSP 0364 *) . São Paulo : 1♂ , Bertioga , Parque Estadual Restinga de Bertioga , Fazenda Içassu , 23º45’06”S , 46º03’09”W , 16.ii.2013 , Biffi, G. & Ament, D. C. ( MZUSP 0536 ) ; 2♂ , 3♀ , Ilha da Vitória , 16–27.iii.1964 , Exp Dep Zool ( ESALQ ; MZUSP 0165 , 0167 , 0168 *, 0169 ) ; 1♂ , 1♀ , Ilha da Vitória , 29.iii–6.iv.1965 , Exp Depto Zool ( MZUSP 0172 , 0173 ) ; 1♂ , Mogi das Cruzes , próximo à Pedreira Itapeti , 23º29’36”S , 46º15’07”W , 5.ii.2016 , V. M. Ghirotto ( MZUSP 0611 *) ; 1♂ , Raiz Serra ( ESALQ 6532 ) ; 1♀ , Rio Paraná , xii.1929 ( ESALQ ) ; 1♂ , Salesópolis , Est Biol Boraceia , 17.ii. 965, E. X. Rabello ( MZUSP 0166 ) ; 1♀ , 1 egg , Salesópolis , Estação Biológica de Boraceia , 23º39’S , 45º53’W , 6–8.ii.2016 , P. I. Chiquetto-Machado , A. Z. Ramin ( MZUSP 0584 ) ; 1♀ , Santos , Praia do Itaguaré , 6.ii.62 , Rabello ( MZUSP 0171 ) ; 1♂ , Santos , Praia do Itaguaré ( MZUSP 0170 ) ; 2♀ , 14 eggs , São Paulo , Jardim Alviverde , 23º45’56”S , 46º45’34”W , ii.2016 , R. Rossi , P. W. Engelking ( MZUSP 0582 , 0583 ) ; 1♂ , São Paulo , Jardim Alviverde , 23º45’58”S , 46º45’35”W , xi.2016 , P. W. Engelking , E. W. Engelking ( MZUSP 0794 *) ; 2♂ , 4♀ , 40 eggs , São Paulo , Jardim Alviverde , 23º45’58”S , 46º45’35”W , 9.i.2017 , P. I. Chiquetto-Machado , A. Z. Ramin ( MZUSP 0760 , 0761 , 0762 , 0763 , 0771 , 0856 ) ; 1♂ , São Sebastião , Centro de Biologia Marinha da Universidade de São Paulo , 23º49’36”S , 45º25’26”W , 12.vii.2017 , E. B. Crispino ( MZUSP 0872 *) .