Metaphotina Piza, 1964

Ferraz, Bernardo R., Queiroga, Drielly, Souza-Dias, Pedro G. B. & Rivera, Julio, 2025, Taxonomic revision and natural history of Metaphotina Piza, 1964 (Mantodea, Acontistidae), Zootaxa 5646 (3), pp. 351-399 : 355-359

Metaphotina Piza, 1964

( Figs. 2 , 3 )

Metaphotina Piza, 1964:125 (original description; Type species: Metaphotina piracicabensis Piza, 1964 , by original designation). Terra, 1995: 23 (as synonym of Acontista ); Ehrmann, 2002:50 (as synonym of Acontiothespis ); Otte & Spearman, 2005:14, 442 (catalog, as synonym of Acontista ); Roy, 2006: 329 (as synonym of Acontista ); Rivera & Svenson, 2016: 642 (phylogenetics and systematics remarks); Schwarz & Roy, 2019:128 (taxonomic remarks); Rivera & Svenson, 2020:124 (as valid, rescued from synonymy with Acontista , redescription, checklist).

Diagnosis. Small, ranging from 14.18 to 22.6 mm ( Fig. 2 ). Males fully winged, wings largely hyaline but with brown markings of variable size on either the meso- or metathoracic wing. Females brachypterous, wings scale-like and not extending beyond abdominal TG3; metathoracic pair opaque and strongly pigmented in red/orange and black. The ventral phallomere of males with strongly sclerotized sdp that distally splits into two processes: one long, curved, and spiniform, the other broad and flattened.

Redescription. Small body size, ranging from 14.18 to 22.6 mm. Males typically emerald green, occasionally appearing brown or reddish. Females usually emerald green with multiple white markings, sometimes in shades of brown, reddish, yellowish and more rarely fully black or pink with traces of white; or combinations thereof.

Head triangular, wider than long, vertex convex to flattened, not exceeding or only slightly surpassing the imaginary line connecting the top of the compound eyes; compound eyes kidney-shaped and slightly protruding, outer margin evenly curved. Lower frons transversal and divided into three parts: central third elevated towards the base of the scape, upper margin moderately arched toward the central ocellus; flanking thirds rectangular. Antennal flagellum moniliform at its base for about 1/4 of the length, becoming filiform distally; sclerite robust, scape cylindrical, pedicel short and cylindrical, slightly longer than the ocelli.

Prothorax short and stout. Supracoxal dilatation broad and rounded. Prozona and proximal two-thirds of the metazona forming an oval outline, with the prozona exhibiting a well-marked expansion of the lateral margin, forming a ledge; metazona strongly constricted proximally; distal margin of metazona slightly curved upwards and elevated. Prothoracic coxae almost as long as the pronotum; prothoracic femur wide, well-developed, triangular, dorsal margin straight; tsg located between the DS and the first AvS; distance between the first and second PvS half the distance between any subsequent pair; ventral aspect of the prothoracic femur pilose between AvS and PvS; foretibial AvS perpendicular or inclined forward relative to the tibial margin, gradually increasing in size distally; the gap between two AvS about as long as a single AvS; foretibial PvS gradually increasing in size distally, strongly inclined forward and tightly arranged, leaving no space between any contiguous PvS; ventral aspect of the foretibia pilose between AvS and PvS; all femoral and tibial spines colored as the rest of legs, but with dark tips. Spination formula: F=3DS/11–13AvS/5PvS; T=10–12AvS/11–15PvS. Mesothoracic legs shorter than the metathoracic legs; tarsi bearing numerous bristles.

Male mesothoracic wings slightly longer than metathoracic wings. Mesothoracic wings hyaline, unpigmented or spotted, costal field densely reticulated with non-parallel veins, subopaque, sometimes the proximal third markedly greenish, membrane sometimes exhibit grayish tones, giving it a smoky appearance. Metathoracic wings hyaline, unpigmented or spotted, veins sometimes with a reddish tint.

Females brachypterous, with wings not extending beyond the TG3. Mesothoracic wings opaque, same color as the body, white stripes and/or a circular distal spot of varying intensities sometimes present. Metathoracic wings shorter than mesothoracic wings. Discoidal field and proximal portion of the anal field pigmented in red or orange, anal field mostly black with white crossveins and black to white longitudinal veins.

Male abdomen slender, cylindrical, slightly flattened dorsoventrally, all terga pigmented in crimson red; TG10 short and apically rounded; CS9 well-developed and spoon-shaped, longer than TG10 and bearing bristles; short cerci bearing numerous long bristles, styli absent. Female abdomen ovoid, robust, the first two segments reddish, remaining segments the same color as the body; abdomen medially expanded but tapering towards TG7–9; CS7 is broad, obtuse, with a central distal fold and a large number of distal bristles. Cerci shorter than in males, not extending beyond CS7.

Male genitalia with ventral phallomere well-developed, convex, rhomboidal in shape; sdp well-sclerotized, bifurcated, with a broad, flattened main projection (sometimes with a small distal hook) and a spiniform, long secondary projection. L4A medially divided by a central sigmoid outline from the top to the sdp; slight sclerotization on the left margin and a large membranous area on the right. The left phallomere square, L4B relatively well sclerotized compared to the rest of the genitalia, divided diagonally by a mostly smooth membranous region with tufts of spines posterior to afa and at loa. The afa well-sclerotized, with a relatively uniform prism-like appearance, the paa is short with a curved apex under the posterior bending of the left phallomere, which can be flattened or concave; loa simple, bearing many setae. The right phallomere with a projecting fda, with tufts of spines similar to left phallomere; R3 straight, elongated, with a globular and flattened apex, pva more developed than the pia, possibly overlapping the latter.

Female genitalia with hexagonal or lozenge CG8, median portion facing sbu well-sclerotized, apex projected, proximally with two projections or tubercles; spb smooth; cxal curved caudally reaching cxvl, the latter only presenting a few inconspicuous bristles at the end; cxdl membranous or strongly sclerotized; gpmo8 short in length but quite pronounced, smooth; gpal8 smooth; gl9 wide, smooth, with sparse bristles.

Oothecae guttiform (broad at the base and tapering distally), cylindrical to slightly flattened, featuring a filiform residual process of varying length. External coating yellowish-white and extensive, surrounding the entire ootheca. Fixed to supporting substrate by its proximal end.

Distribution. Northern Argentina, Bolivia, Brazil (BA, CE, DF, GO, MA, MT, MS, MG, PA, PB, PR, PE, RN, RS, SP, TO), Paraguay, Uruguay.

Taxonomic history

Piza (1964) formulated Metaphotina to classify his new species Metaphotina piracicabensis Piza, 1964 . Terra (1995) subsequently synonymized Metaphotina with Acontista . Based on molecular data and comparative morphology, Rivera & Svenson (2016, 2020) concluded that Metaphotina represented an independent lineage closely related to Raptrix Terra, 1995 and distinct from Acontista . To reflect their proposed phylogenetic classification ( Rivera & Svenson 2016), they restored and taxonomically circumscribed Metaphotina , adding four species ( Rivera & Svenson 2020): M. bimaculata ( Saussure, 1870) , M. brevipennis (Saussure, 1872) , M. rehni ( Giglio-Tos, 1927) , and M. piracicabensis Piza, 1964 . Ferraz et al. (2023) added a fifth species, M. austri Ferraz, Souza-Dias & Rivera, 2023 . A summary of the taxonomic history of each species follows.

M. bimaculata . Saussure (1870) described Acontista bimaculata Saussure, 1870 based on a male holotype from “Brasilia” (= Brazil), which we located at the MNHN ( Fig. 3A ). Consequently, the purported syntype held at the MHNG ( Fig. 3C ), reported in Pfäuti & Hollier (2012) and later portrayed in Rivera & Svenson (2020: Fig. 44D), is not a primary type as they presumed, but a specimen reported much later in Saussure & Zehntner (1894) and collected by von Ihering. Saussure (1871) expanded the geographic range of A. bimaculata by adding Chiquitos ( Bolivia) and Goiás ( Brazil) (see Fig. 3D ) and reported new morphological details to help distinguish it from the other then-known species of Acontista . In the same publication, Saussure (1871) disclosed Goiás as the actual type locality of A. bimaculata (omitted in the original description of 1870). Therefore, the specimen from Goiás reported in Saussure (1871) and later portrayed in Rivera & Svenson (2020: Fig. 44C) ( Fig. 3D ) is not a primary type specimen (despite being labeled as such) but a topotypical one. Subsequent contributions added to our knowledge of the distribution of this species: Rio Grande do Sul ( Brazil) ( Saussure & Zehntner 1894), Asunción ( Paraguay) ( Giglio-Tos 1894), San Lorenzo ( Argentina), and Caiza ( Bolivia) ( Giglio-Tos 1897), as well as Sapucay (= Sapucai) ( Paraguay) ( Caudell 1904; Rehn 1907). Rehn (1907) further identified and described the female of A. bimaculata based on topotypical specimens and remarked on its chromatic variability. Later, Rehn (1916) transferred A. bimaculata to a newly established genus, Acontiothespis Rehn, 1916 and mapped its distribution from southern Bolivia to northern Argentina, with additional records in Misiones ( Rehn 1920). However, the taxonomic journey of A. bimaculata took a confusing turn when Beier (1934) unwarrantedly synonymized it with A. concinna Perty, 1833 , this time under the genus Acontista . Beier’s (1934) nomenclatural act led to considerable confusion regarding the identity of A. bimaculata through the rest of the 20th century, a problem aggravated by another controversy centered around the pertinence of the name Acontiothespis that Rehn (1916) proposed (unnecessarily) as a replacement name for Acontista . For instance, Terra (1995) followed Beier’s (1934) opinion regarding A. bimaculata as a synonym of A. concinna and treated it under Acontista , whereas Ehrmann (2002) considered A. bimaculata as a synonym of yet another species, Acontista aurantiaca Burmeister, 1838 , instead of A. concinna , but under Acontiothespis . Following Roy’s (2004) confirmation of Acontista as the accepted generic name, he conducted a thorough review of all names linked to Acontistinae and their respective type repositories in a follow-up work ( Roy 2006), in which he also correctly rejected the synonymy of A. bimaculata with A. concinna that Beier (1934) had proposed. Rivera & Svenson’s (2016) phylogenetic studies and the concomitant confirmation of A. concinna and A. aurantiaca as valid species, distinct from A. bimaculata ( Rivera & Vergara-Cobian 2017; Schwarz et al. 2020), prompted Rivera & Svenson (2020) to restore Metaphotina Piza, 1964 to accommodate A. bimaculata as Metaphotina bimaculata ( Saussure, 1870) .

M. brevipennis . Saussure (1872a) described Acontista brevipennis Saussure, 1872 based on a female holotype from “ Brazil ”. Saussure (1872a) distinguished his new species by its brachypterous condition. The type specimen is presumed lost, but an illustration of it ( Saussure 1872b: Fig. 23 ) was added to the original description (reproduced here in Fig. 3B ). Subsequent contributions significantly expanded the known range of A. brevipennis , encompassing San Francisco ( Paraguay) and Resistencia ( Argentina) ( Giglio-Tos 1894), as well as San Lorenzo ( Argentina) and Aguayrenda and Caiza ( Bolivia) ( Giglio-Tos 1897). However, Giglio-Tos (1897) speculated that only the female specimens he examined could be confidently attributed to A. brevipennis , whereas some males, previously identified as A. bimaculata by himself ( Giglio-Tos 1894), actually represented Acontista vitrea Saussure and Zehntner, 1894 , a species primarily found in Central America (currently regarded as a synonym of Acontista cordillerae Saussure, 1869 ). In other words, Giglio-Tos (1897) believed that the species then referred to as A. vitrea represented the male of A. brevipennis . If Giglio-Tos hadn’t been misled by the marked sexual dimorphism in this lineage and the similarities between its males and those of A. vitrea , he might have suspected that A. bimaculata and A. brevipennis actually were names given to different sexes of the same species. Caudell (1904) extended the range of A. brevipennis to include Sapucay (= Sapucai, Paraguay). Rehn (1916) described the presumed male of A. brevipennis , but under Acontiothespis , and mapped its distribution to encompass Paraguay and southern Brazil, noting its presence also in northeastern Brazil (Caatinga biome); however, the latter were eventually demonstrated to belong to an undescribed species, not to A. brevipennis ( Ferraz et al. 2023) . Giglio-Tos (1927) listed A. brevipennis under Acontista and provided descriptions of both sexes, this time without addressing his previous assertions ( Giglio-Tos 1894, 1897) of synonymy with A. vitrea . Heitzmann-Fontenelle (1968) offered a detailed morphological description of A. brevipennis (under Acontiothespis ), and proposed A. bimaculata as its synonym, marking the first instance in the historical literature of both names being attributed to the same species. However, Heitzmann-Fontenelle (1968) erroneously accorded priority to the name A. brevipennis over A. bimaculata , as per the Principle of Priority outlined in the ICZN (1999), despite the latter being described two years earlier. Terra (1995) and Otte & Spearman (2005) listed A. brevipennis under Acontista , summarizing its distribution as Argentina, Bolivia, Paraguay, and Brazil, while Ehrmann (2002) placed it in Acontiothespis . Finally, after recovering Metaphotina from synonymy with Acontista , Rivera & Svenson (2020) settled the name of this species as Metaphotina brevipennis (Saussure, 1872) . Trillo et al. (2021) added new records of M. brevipennis from Uruguay; however, the species reported is actually M. bimaculata .

M. rehni . Giglio-Tos (1927) formulated Acontista rehni Giglio-Tos, 1927 , based on a female specimen ( Fig. 3G ) from “Sapucay” (= Sapucai, Paraguay) that Rehn (1907) had identified as Acontista bimaculata ; Giglio-Tos (1927) provided no justification for this action. Beier (1934) and subsequent catalogs and checklists ( Terra 1995; Ehrmann 2002; Otte & Spearman 2005; Roy 2006; Agudelo et al. 2007), continued to list A. rehni either under Acontista or Acontiothespis . No further taxonomic developments on A. rehni occurred until Rivera & Svenson (2020) reinstated Metaphotina , relocating it as Metaphotina rehni ( Giglio-Tos, 1927) .

M. piracicabensis . Piza (1964) described Metaphotina piracicabensis Piza, 1964 both as a new genus and species based on a male specimen ( Fig. 3E ) collected in São Paulo, Brazil (ESALQ), but erroneously classifying it among the Photinainae ( Photinaidae ). Terra (1995) transferred M. piracicabensis to Acontista after establishing Metaphotina as its synonym. Besides appearing in subsequent catalogs and checklists ( Ehrmann 2002; Otte & Spearman 2005; Agudelo et al. 2007), the taxonomic status of M. piracicabensis remained unchallenged. Following phylogenetic and morphological studies, Rivera & Svenson (2016, 2020) recognized the distinctiveness of Metaphotina , rescuing A. piracicabensis ( Piza, 1964) from synonymy under Acontista and returning it to its original placement as Metaphotina piracicabensis Piza, 1964 . Additionally, Rivera & Svenson (2020) reported a female specimen ( Fig. 3F ), labeled as “ allotype ”, accompanying the holotype; however, this specimen, collected a decade after the holotype, is evidently not part of the type series and thus cannot be considered a primary type.

M. austri . Ferraz et al. (2023) described both sexes of Metaphotina austri Ferraz, Souza-Dias & Rivera, 2023 , from northeastern Brazil, with the reported type locality in Aiuaba, Ceará, in addition to multiple sites in Bahia, Maranhão, Paraíba, Pernambuco, and Rio Grande do Norte states. Ferraz et al. (2023) attributed specimens from northeastern Brazil that Rehn (1916) had reported as Aconthiothespis brevipennis to their new species.

Figures

FIGURE 2. Live specimens of Metaphotina bimaculata (A–F) and Metaphotina austri (G–J) retrieved from iNaturalist platform, license CC-BY-NC. Images may have been cropped and/or rotated for better observation. (A) Observation 163494503, by goncrisdi [GonzaloDias],Argentina, 06/May/2023.(B) Bernardo Ferraz, Brazil (MG).(C) Observation 199319301, by jcsullivan [John Sullivan], Bolívia, Jan/2024. (D) Observation 223539397, by lucas_fornero [Lucas Fornero], Argentina, 11/May/2024. (E) Bernardo Ferraz, Brazil (MG). (F) Observation 137464405, by entomofobia [Juan Zara], Argentina, 16/Apr/2022. (G) Observation 167907017, by fredericosonntag [Frederico Acaz Sonntag], Brazil (PB), 17/Jun/2023. (H) Observation 147265948, by offvinn, Brazil (CE) 15/Jan/2023. (I) Observation 137450242, by fredericosonntag [Frederico Acaz Sonntag], Brazil (PB), 26/Jan/2000. (J) Observation 62552485, by alenilson [Alenilson], Brazil (AL), 19/Aug/2020.

FIGURE 3. Type material of Metaphotina spp. (A) True holotype of A. bimaculata described by Saussure (1870), deposited at MNHN. (B) Illustration of the holotype of A. brevipennis Saussure, 1872 and a reference to it from the precedent page: “Fig. 20. Acontista brevipennis, Sauss. Femelle de grandeur naturelle.” (C) A. bimaculata cited in Saussure & Zehntner (1894), mislabeled as “syntypus” (Pfäuti & Hollier 2012), deposited at MHNG. (D) A. bimaculata specimen cited in Saussure (1871) and mislabeled as “Type”, deposited at MNHN (adapted from Rivera & Svenson 2020). (E) Holotype of M. piracicabensis, deposited at ESALQ. (F) M. piracicabensis labeled (incorrectly) as “allotype”, deposited at ESALQ. (G) Holotype of A. rehni, deposited at ANSP. Scale: 10 mm.

FIGURE 23. Metaphotina spp. habitus, dorsal view. (A) Metaphotina bimaculata, male. (B) Metaphotina bimaculata, female. (C) Metaphotina austri, male holotype. (D) Metaphotina austri, female allotype. (E) Metaphotina akaraje sp. nov., male holotype. (F) Metaphotina akaraje sp. nov., female allotype. Scale: 10 mm.