Monograph |
Corresponding author: Adrian Ardila-Camacho ( aardilac88@gmail.com ) Corresponding author: Atilano Contreras-Ramos ( acontreras@ib.unam.mx ) Academic editor: Davide Badano
© 2024 Adrian Ardila-Camacho, Renato José Pires Machado, Michael Ohl, Atilano Contreras-Ramos.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ardila-Camacho A, Machado RJP, Ohl M, Contreras-Ramos A (2024) A camouflaged diversity: taxonomic revision of the thorny lacewing subfamily Symphrasinae (Neuroptera, Rhachiberothidae). ZooKeys 1199: 1-409. https://doi.org/10.3897/zookeys.1199.115442
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Species of the thorny lacewing subfamily Symphrasinae (Neuroptera: Rhachiberothidae) are revised. Prior to this work, 42 species were known in the genera Anchieta Navás, 1909, Plega Navás, 1928, and Trichoscelia Westwood, 1852. Herein, the number of species is increased to 60, 23 of which are newly described. Species previously known are redescribed, and their taxonomic status is revised. Keys, diagnoses, and high-resolution images for all species are presented. The distribution range of Anchieta is now known from Costa Rica to southern Brazil with a total of 11 species, of which three are newly described. The genus Plega is known from southwestern United States to southern Brazil and includes 28 species of which 14 are described as new. Moreover, the genus Trichoscelia occurs from central and southern Mexico to Argentina, with a total of 21 species, of which six are herein newly described. A phylogenetic analysis of Symphrasinae based on morphological characters recovered the three symphrasine genera as monophyletic, with Anchieta sister to Plega + Trichoscelia. The three genera are newly diagnosed based on a cladistic framework. Within the genus Anchieta, bee-mimicking species comprise a monophyletic group, while wasp-mimicking species form a laddered sequence to that lineage. Within Plega, three lineages are recovered, the first mostly composed of South and Mesoamerican species, the second with species predominantly from Central America and central and southern Mexico, and a third clade encompassing species mostly from central and northern Mexico and southwestern United States. By contrast, relationships between species of Trichoscelia were poorly resolved because of a simplified and conserved morphology of this group.
Lacewings, Mantispoidea, morphology, New World, systematics
We dedicate this work to the memory of neuropterists who eased the way for today’s taxonomy of Symphrasinae. They have made possible our current understanding on this enigmatic group: Robert G. Beard intended to write a revision of American Mantispidae before his death in 1968, and many type specimens in European museums have determination and synonymy labels from him. Norman D. Penny (1946–2016) revised the Symphrasinae from the Amazon Basin and these publications motivated AAC since an undergraduate (2007–2013). Finally, Lionel A. Stange (1935–2020) engaged in revising the fauna of Symphrasinae from North and Central America; however he did not conclude this project as he dedicated strong efforts to Myrmeleontidae through his long career, yet he generously shared many observations, drawings, and materials with AAC one year before his death.
The superfamily Mantispoidea includes three extant families, i.e., Berothidae, Rhachiberothidae, and Mantispidae, of which the latter two are collectively known as raptorial Mantispoidea or Raptoneuroptera due to the presence of grasping forelegs for prey capture (
In a recent phylogenomic study of Neuropterida, previous hypotheses of the relationships between the Mantispoid subfamilies were challenged, as the Mantispidae was recovered as a polyphyletic group, with Symphrasinae and Rhachiberothinae as sister groups in a clade sister to Berothidae and the remainder Mantispidae subfamilies (
The subfamily Symphrasinae was proposed by
The type genus of this subfamily, i.e., Symphrasis was proposed by
The phylogenetic relationships of the genera in the Symphrasinae are not well understood, and the boundaries of the genera are not clearly outlined (
Besides the contentious background on the understanding of the phylogenetic relationships between the symphrasine genera, the extant diversity of the group has never been previously revised, except for the fauna from Amazonia and the United States (
We examined and identified > 3000 specimens of Symphrasinae deposited in collections and museums of Europe, United States, Mexico, Costa Rica, Colombia, and Brazil (see collections section below). Nearly all the primary types of the subfamily were directly examined and redescribed, and for the majority high resolution photographs were obtained.
Genitalia preparations were made by clearing the last abdominal segments in a hot solution of 10% potassium hydroxide (KOH). Residuals of the alkaline solution, gut contents, and tracheae were washed with a series of washings of distilled water and 80% ethyl alcohol, sometimes with the aid of a tuberculin syringe, a minute steel hook inserted at the tip of a wood stick, and fine forceps with blunt tip. A cut along the pleural membrane allowed extraction of the male genital complex for examination on an excavated microscope slide (
Foreleg morphology of Symphrasinae A–C Anchieta fumosellus (Westwood, 1867) A forefemur, ventral Surface B trochanter, femur, tibia, and tarsus, anterior surface C fore- tarsus and pretarsus, anterior surface D–F Plega dactylota Rehn, 1939 D forefemur, ventral Surface E trochanter, femur, tibia, and tarsus, anterior surface F fore- tarsus and pretarsus, anterior surface. Abbreviations: avr, anteroventral processes row bt, basitarsus cvs, clavate setae et, eutarsus fe, femur fcs, femoral closing surface ftso, foretarsal Stitz organ lcp, lanceolate process p, primary process ptc, pretarsal claw pvr, posteroventral processes row ta, tarsus ti, tibia tp, trochanter process tr, trochanter tbs, tubercle-shaped specialization tsgb, thickened seta with globular base.
Wing venation and genitalic morphology of Plega dactylota Rehn, 1939 as an example for Symphrasinae A forewing B hind wing C male terminalia, lateral D same, ventral E male genitalia, dorsal F same, ventral G same, lateral H female terminalia, ventral I same, lateral J female genitalia, ventral. Abbreviations: h, humeral vein C, Costa scvl, subcostal veinlets 1r-m, first radiomedial crossvein rarp1, first radial anterior cell rarp2, second anterior radial cell pt, pterostigma g.s., gradate series jv, jugal vein jl, jugal lobe 1cua-cup, first intracubital crossvein tVIII, eighth tergite, tIX, ninth tergite cc, callus cercus ect, ectoproct sVII, seventh sternite sVIII, eighth sternite sIX, ninth sternite ml, median lobe of gonocoxites XI dp, digitiform processes bc, bursa copulatrix p.s., proximal section of spermatheca m.s., medial section of spermatheca d.s., distal section of spermatheca fc, fertilization canal.
CAS California Academy of Sciences, San Francisco, USA.
CN Navás collection housed at (MZBS) Museo Zoologia, Barcelona, Spain.
CZMA Coleção Zoológica do Maranhão, Caixias, Maranhão, Brazil.
DZUP Coleção Entomológica Padre Jesus Santiago Moure, Curitiba, Paraná, Brazil.
IAvH Instituto Alexander von Humbolt, Villa de Leyva, Colombia.
MHN-UPN Museo de Historia Natural, Universidad Pedagógica Nacional, Bogotá, Colombia.
MPUJ Museo Javeriano de Ciencias Naturales, Bogotá, Colombia.
NMBS Naturhistorisches Museum, Bern, Switzerland.
For the phylogenetic analysis of Symphrasinae, 10 species of Anchieta, 25 of Plega, and 19 of Trichoscelia studied herein were included. One species of Mantispidae, i.e., Gerstaeckerella irrorata (Erichson, 1839), and one of Rhachiberothinae, i.e., Mucroberotha vesicaria Tjeder, 1968 were selected as outgroups.
The morphological characters used for the phylogenetic analysis were determined by comparing the majority of species of Symphrasinae described herein with the outgroups, plus the detailed comparative study of the raptorial Mantispoidea by
The Bremer absolute (
The final topology, including all characters, states and Bremer values, was edited using Adobe Illustrator CS® software.
Head
Prothorax
Foreleg
Mid- and hind leg
Wings
Forewing
Hind wing
Abdomen
Male genitalia
Female genitalia
The cladistic analysis performed using equal weight and New Technology Search recovered 33 most parsimonious trees with 311 steps in length (L), consistency index (CI) of 62 and retention index (RI) of 86, which generated a strict consensus tree with L = 319, CI = 61, and RI = 86. A weighted analysis using New Technology Searching yielded similar results to the analysis using equal weighting, and thus was not used for further discussion.
In this analysis, the subfamily Symphrasinae was recovered as a monophyletic group with strong support, with Anchieta as sister of Plega + Trichoscelia (Fig.
The genus Anchieta was recovered as monophyletic with A. fasciatellus (Westwood, 1867) sister to the rest of the species (Fig.
The sister-group relationship between the genera Trichoscelia and Plega is supported by synapomorphic venational characters such as a trapezoidal pterostigma of FW (char 42:3) which is dark with pale medial area (char 43:2), a candelabrum-shaped first branch of CuA of HW (char 68:2), and two crossveins in the cubitoanal space of HW (char 72:2) (Fig.
The genus Trichoscelia was rendered monophyletic with strong support, although the relationships within the genus were largely unresolved and recovered in big polytomies (Fig.
The genus Plega was recovered as monophyletic with strong support based on six unequivocal changes such as the collar-like (i.e., ventrally fused) postfurcasterna (char 18:3), the posteroventral row of processes with two primary, spine-shaped processes on proximal ½ of forefemur (char 25:3), the presence of stinger-shaped setae on the integumentary specializations rows of forefemur (char 27:1), male sternite IX pentagonal with rounded posterolateral corners (char 76:3), distal section of the spermatheca slightly wider than proximal and medial sections, lacking diverticulum (char 100:3), and fertilization canal duct sinusoid with proximal, triangular, concave portion (char 102:3) (Fig.
The first clade in this group is composed by P. disrupta Ardila-Camacho & Contreras-Ramos, sp. nov., P. yucatanae Parker & Stange, 1965, P. duckei Penny, 1983, P. paraensis Penny, 1983, P. hagenella (Westwood, 1867), P. bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov., and P. radicaudata Ardila-Camacho & Contreras-Ramos, sp. nov. In this group, the former species is sister of the rest, and the whole clade is supported by a homoplasious character state, i.e., a bursa copulatrix with sclerotized and membranous areas (char 99:2). Within this clade, there is a group conformed by three species, P. hagenella, P. bowlesi, and P. radicaudata, of which the former is sister of the latter two. Synapomorphies supporting this clade are the ventral region of the male ectoproct which is equipped with anterior, flattened lobe anteriorly covered with conical, translucent setae (char 79:5), and the ventral part of the gonocoxites XI medial lobe as a concave covering forming a curved process (char 88:1). A clade composed of P. paraensis and P. duckei was recovered as sister of that group and is supported only by homoplastic features such as a subcylindrical forefemur (char 21:1) and antennal scape as long as wide (char 6:0). These two sister clades are supported by homoplasious character states such as an unmodified supraantennal region (char 10:0) and an arched substigmal cell of FW (char 51:4). Plega yucatanae was found sister of these clades based on two homoplastic features, i.e., the ventromedial, posterior area of the gonocoxites XI median lobe being undeveloped (char 87:0), and the female gonapophyses IX as two tiny sclerites concealed on the inner surface of the base of gonocoxites IX (char 97:1).
Between the previous clade and the rest of the species of the genus, P. drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov., is sister of a large clade composed of two smaller clades and several intermediary species between them. This whole clade is supported by the distal section of the spermatheca which is wider than proximal and medial sections and equipped with blunt diverticulum (char 100:4) as an unequivocal change. Two homoplastic features further support this clade, i.e., distally forked A1 of FW (char 59:3) and male gonocoxites X with anterior apex expanded and dorsally bent (char 83:2). A transitional species between P. drepanicoides and the rest of species of Plega is P. pseudohagenella Ardila-Camacho & Contreras-Ramos, sp. nov. Such a clade is supported by two synapomorphies: a short and blunt process on the ventral part of the gonocoxites XI median lobe (char 88:4) and the presence of tiny posteromedian process of the female gonapophyses VIII with short lateral lobes (char 94:3). Inside this latter clade, there are two sister subclades whose monophyly is supported only by homoplastic features. The first one is composed of P. vangiersbergenae Ardila-Camacho, sp. nov., P. stangei Ardila et al., 2019, P. lachesis Ardila-Camacho & Contreras-Ramos, sp. nov., P. obtusa Ardila-Camacho & Contreras-Ramos, sp. nov., P. oswaldi Ardila-Camacho & Contreras-Ramos, sp. nov., and P. spinosa Ardila et al., 2019 and is supported by the presence of female gonapophyses IX as two tiny sclerites at the base of the gonocoxites IX (char 97:1). Of these species, the first one is sister of the rest, followed by P. stangei as sister of the remaining species. A clade composed of P. lachesis as sister of P. obtusa + (P. oswaldi + P. spinosa) is supported by an unequivocal change, the presence of modified setae on the posterior surface of the male ectoproct (char 80:1). The sister clade of P. lachesis is supported by a digitiform or enlarged ventromedial posterior area of the gonocoxites XI median lobe (char 87:3) as synapomorphic feature, while the trapezoidal male sternite IX (char 76:4), and the male gonocoxites IX lacking digitiform processes are homoplastic features also supporting this group.
The last clade in the phylogeny of Plega is composed by species found in the Mexican pacific coast, Baja California peninsula, Central and Northern Mexico, and Southwestern United States. This group is well defined and includes P. mixteca Ardila et al., 2019, P. dactylota Rehn, 1939, P. fumosa Linsley & MacSwain, 1955, P. flammata Ardila-Camacho & Contreras-Ramos, sp. nov., P. signata (Hagen, 1877), P. insolita Ardila-Camacho & Contreras-Ramos, sp. nov., P. banksi Rehn, 1939, P. megaptera Ardila-Camacho & Contreras-Ramos, sp. nov., and P. sonorae Ardila et al., 2019. Nonetheless, this a clade is supported only by four homoplasious characters states: basal antennal flagellomeres > 3 × as wide as long (char 7:3), the unmodified supraantennal region (char 10:0), the gently curved substigmal cell of FW (char 51:0), and the gonocoxites X only slightly wider on anterior apex (char 83:0). Within this clade, P. mixteca is sister to the rest of the species, followed by P. dactylota as sister of two sister clades which are supported by two homoplastic features. The first one, is the antennal scape as long as wide (char 6:0), and the posteromedial process on the female gonapophyses VIII medial part elongated and Y-shaped (char 94:4). Within this clade, the first subclade is composed by P. fumosa as sister of P. flammata + P. signata. Such a clade is supported by a homoplastic character state, i.e., the ventromedial posterior area of male gonocoxites XI median lobe not developed (char 87:0), while the sister relationship between the latter two species is supported by an enlarged, curved, rugose, ventral process on the male gonocoxites XI (char 88:3) as a synapomorphy. Additionally, the relationship between both species is supported by two homoplastic features, the basal antennal flagellomeres (char 7:2) and the absence of posteromedian process on the female gonapophyses VIII (char 94:0). The second clade is constituted by P. insolita as sister of P. banksi (P. megaptera + P. sonorae). This group is supported by a single synapomorphy, the presence of a ventral, curved process with bilobed apex on the ventral part of the gonocoxites XI median lobe (char 88:3). The latter clade is supported by a homoplastic character state, i.e., a pentagonal to semi-triangular male sternite IX with straight vertexes (char 76:2).
Class Insecta Linnaeus, 1758
Order Neuroptera Linnaeus, 1758
Family Rhachiberothidae Tjeder, 1959
Symphrasini
Anisopterinae
Platymantispinae Rehn 1939: 82.
This subfamily is distinguished from other mantispoid subfamilies by the presence of hypostomal bridge, the undeveloped laminatentorium and straight ocular plane. The procoxa are inserted approximately at mid-length of the prothorax, the pronotum is shield-shaped, and the postfurcasternum is well-developed and plate-shaped. On the foreleg, the femoral closing surface is proximally curved and presents two rows of thickened setae with globular base adjacent to the rows of processes; the foretarsus is four-segmented in both sexes with basitarsus equipped with a patch of clavate seta on anterior surface, a row of prostrate setae on the closing surface, and a long, lanceolate process set with an apical plug-shaped Stitz organ. On the wings, there are more than two trichosors on the posterior wing margin between the apex of two longitudinal veins, the Sc vein is fused to RA at pterostigma level, and two crossveins in the radial space of the forewing are present. Additionally, the CuP of the forewing is proximally bent, approaching or touching the A1. On the hind wing, the C and Sc veins are fused at or slightly beyond the level of R fork, the M vein is tuning fork-shaped, the 1r-m is sigmoid, and the CuP is developed and free. Male genitalia characteristics of Symphrasinae include gonocoxites IX set with apical, digitiform processes, paired gonapophyses X not fused to gonocoxites X, and gonostyli X remarkably long, incurved or forming loops. Moreover, on the female genitalia, the medial part of the gonapophyses VIII is enlarged and elaborated, the gonocoxites IX remarkably elongated forming an ovipositor, and the tergite IX and ectoproct are fused.
Summarized in
1 | Foretrochanter with blunt process on anterior surface (Fig. |
Anchieta Navás 1909 |
– | Foretrochanter without blunt process on anterior surface; HW with first branch of CuA candelabrum-shaped (Fig. |
2 |
2 | Third labial palpomere expanded, with broadly ovoid palpimacula (Fig. |
Trichoscelia Westwood, 1852 |
– | Third labial palpomere narrow, with sulcate or narrowly ovoid palpimacula; forefemur with rows of thickened setae with globular base with some degree of reduction to distal portions (Fig. |
Plega Navás, 1928 |
Anisoptera
Schneider, 1843: 32. Type species: Mantispa notha Erichson, 1839: 170 (now in Anchieta), by monotypy. Junior homonym of Anisoptera Berthold, 1827: 409 (in Orthoptera) and Anisoptera Herrich-Schäffer, 1840: 57, 69 (in Hymenoptera). Replaced by Platymantispa Rehn, 1939 and Anisopterana Strand, 1942. Synonymized with Anchieta by
Anchieta Navás, 1909: 483. Type species: Anchieta nobilis Navás, 1909: 484 (= Anchieta fumosella (Westwood, 1867: 504)), by monotypy.
Platymantispa Rehn, 1939: 82. Name replacement for Anisoptera Schneider, 1843.
Anisopterana
Strand, 1942: 389. Unnecessary name replacement for Anisoptera Schneider, 1843, previously replaced by
Anchieta Bechyné, 1954: 176 (in Coleoptera). Junior homonym of Anchieta Navás, 1909.
Compound eyes are relatively small, generally ½ the interocular distance at toruli level. The pronotum is as long as wide, unlike the other genera it presents only a slight outgrowth on the posterior margin. The forecoxa is slightly expanded at the apex, the trochanter is unique due to the presence of a blunt process on the anterior surface. The forefemur is setose; on the closing surface, the tubercle-shaped processes are noticeably thickened, with a spine-shaped sub-basal process on the posteroventral row. The anteroventral row is more reduced than in Plega, although like this, it presents the basal, primary, spine-shaped process. The foretibia is glabrous. On the hind leg, the tibia is generally markedly expanded. The pterostigma of the forewing is rectangular, and the rarp2 is straight to gently curved; the hind wing is often noticeably shorter and narrower than the forewing, with the gradate series of crossveins absent or reduced to a single crossvein, and the first branch of the CuA is simple or forked. On the abdomen, the tergites of segments III–VII often present posteroventral keeled processes in both sexes. As in Mucroberotha, sternites VIII and IX of the male are fused, the sternite IX forms a wide canal, and the ectoproct presents a posteroventral patch of conical and thickened setae. The gonostyli X are short and recurved, and the gonocoxites XI have a flattened medial lobe. On the spermatheca, the distal section is expanded and sac-like.
Head. Diamond-shaped in frontal view, smooth to rugose, region of vertex domed over compound eyes, paraocular area concave; coronal suture discrete. Antenna moniliform, flagellomeres discoidal on most of the flagellum. Compound eye hemispheric, as wide as 0.5–0.7 of the interocular distance at toruli. Thorax. Pronotum nearly as long as wide, with a groove adjacent to lateral and distal margins; posterior margin with slightly raised, entire surface with abundant, thickened setae arising flush the pronotal surface; postfurcasternum quadrangular, paired. Mesonotum wider than long, with abundant long, thickened setae, metanotum ~ 3 × as wide as long, glabrous. Foreleg. Coxa as long as femur, cylindrical, slightly distally expanded, densely setose. Trochanter semi-triangular, with a blunt process on anterior surface. Femur robust, densely setose; closing surface covered with fine and sinuous trichoid setae; posteroventral row processes fully developed, slightly carinated on distal ½, composed of thickened tubercle-shaped specializations with conical Stitz organs; proximally with a more developed, spine-shaped, sub-basal process; adjacent rows of thickened setae with globular base reduced to distal ¾ of the closing surface to a single apical seta; anteroventral row of processes reduced to the proximal region and apex, composed of tubercle-shaped integumentary specializations; basal primary process present, curved; adjacent row of thickened setae with globular base present in distal 4/5. Tibia nearly as long as femur, glabrous, curved, ventrally keel, with a row of prostrate setae; anterior surface with a patch of clavate setae at apex. Basitarsus with long lanceolate process surpassing distal margin of third tarsomere, equipped with a plug-shaped Stitz organ at apex; basal ½ with a row of prostrate setae on ventral surface, and patch of clavate setae on anterior surface; second tarsomere articulated in basal ½ of basitarsus on anterior surface, longer than third and fourth tarsomeres together; pretarsal claws simple. Mid- and hind leg. Mid-leg with thin to slightly widened tibia; hind leg thin to noticeably expanded, and laterally flattened, oar-shaped or fusiform. Wings. Forewing oval, trichosors present along wing margin, except at base; costal space medially narrow to slightly widened, humeral vein simple or forked, subcostal veinlets simple; pterostigma rectangular; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge with the RA; radial space with two crossveins; rarp2 straight to gently curved; RP base located near separation of M and R, M fork near such separation; 1r-m between RP base and M fork forming a small trapezoidal cell; RP with single gradate series present; CuP basally angled, approaching A1. Hind wing oval, notably smaller and narrower than forewing; costal space narrow and reduced, C and Sc fused at proximal 1/3 of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, narrow to slightly widened distally; radial space widened with single crossvein, straight to sinuous; RP with gradate series absent or reduced to a single crossvein. M forked at or slightly beyond R fork; Cu deeply forked, CuA ending at posterior margin at level of 1ra-rp, distally simple or forked, first branch simple or forked; intracubital vein generally reclined; CuP distally anteriorly curved or bent near posterior wing, pectinate. Cubitoanal space without crossveins. Abdomen. Tergites III and IV, or III–VI, or III-VII with poorly developed to prominent, posteromedial, keeled processes, present in both sexes.
Male genitalia. Sternites VIII and IX fused, with fusion line weakly marked or absent; Sternite IX U-shaped, transversely curved to form a wide canal. Gonocoxites IX short and sinuous, or almost completely reduced, with or without apical processes. Ectoproct ovoid, with a posteroventral patch of conical, thickened setae. Gonocoxites X unpaired, forming a triangular or hourglass-shaped sclerite, concave or canaliculate on ventral surface, anterior portion spatulate, straight or dorsally bent; posterior apex with paired dorsal and lateral processes, articulated to gonostyli X and gonapophyses X, respectively; gonostyli X with thickened, concave base, set with curved lateral processes; the rest of the structure whip-like, short, recurved, with apex posteriorly curved, sometimes forming a loop. Gonapophyses X paired, rod-shaped, straight, thin, forming a V-shaped structure, joined by membranes; posterior apex with surrounding membrane set with minute granules. Gonocoxites XI thin, U-shaped, medial lobe dorsoventrally flattened, weakly sclerotized. Hypandrium internum concave, keeled, with two lateral fins.
Female genitalia. Sternite VII (gonocoxites VII) trapezoidal or rectangular, sometimes as medially joined, lateral trapezoidal plates. Tergite VIII narrower medially than laterally, enclosing the spiracle of the segment VIII. Gonocoxites VIII forming a narrow, concave plate; gonapophyses VIII medial part canal-shaped, with or without a tubular process; lateral part as an enlarged plate, hidden under tergite IX + ectoproct, sometimes dorsally fused to form a covering. Tergite IX + ectoproct triangular, elongated. Gonocoxites IX long, straight and narrow; gonapophyses sometimes present as tiny sclerites located basally on inner surface of gonocoxites IX. Bursa copulatrix funnel-shaped, unsclerotized, short to long. Spermatheca short and irregularly entangled or long and spiral-shaped, proximal section short to long and thin; medial section thicker than proximal section, entangled or coiled; distal section expanded, wider than medial section, sac-shaped; fertilization canal duct long, thin, spiral-shaped; fertilization canal short to elongated, J-shaped, covered with microfilaments.
1. A. apiculasaeva Thouvenot, 2009 (Brazil, French Guiana)
2. A. bellus (Westwood, 1867) (Brazil, Colombia, French Guiana, Suriname)
= A. eurydella (Westwood, 1867), new synonym
3. A. fasciatellus (Westwood, 1867) (Colombia, Panama)
4. A. fumosellus (Westwood, 1867) (Brazil)
= Anchieta nobilis Navás, 1909
5. A. nebulosus Ardila-Camacho & Machado, sp. nov. (Brazil)
6. A. nothus (Erichson, 1830) (Brazil)
7. A. partheniellus (Westwood, 1867) (Brazil, Colombia, Ecuador, Suriname, Venezuela)
8. A. remipes (Gerstaecker, 1888) (Colombia)
9. A. romani (Esben-Petersen, 1817) (Brazil, Peru)
10. A. sophiae Ardila-Camacho & Contreras-Ramos, sp. nov. (French Guiana)
11. A. tinctus Ardila-Camacho & Contreras-Ramos, sp. nov. (Costa Rica)
Summarized in
Genus named in honor of the Jesuit missionary San José de Anchieta, evangelizer in Brazil between 1553 and 1597. Masculine gender as the name was erected after a man by
1 | Hind tibia narrow and unmodified or only slightly widened (Fig. |
2 |
– | Hind tibia noticeably expanded and flattened, fusiform or oar-shaped (Fig. |
4 |
2 | Body bright orange with dark brown bands (Fig. |
Anchieta fasciatellus (Westwood, 1867) |
– | Body either yellow with dark brown stripes or dark brown with few orangish areas; male gonocoxite IX with different number and arrangement of the apical processes | 3 |
3 | Male gonocoxite IX with posterior apex blade-shaped, sometimes slightly lanceolate, or with 2 or 3 small, preapical processes (Fig. |
Anchieta fumosellus (Westwood, 1867) |
– | Male gonocoxite IX filiform, with posterior apex sharply pointed (Fig. |
Anchieta sophiae Ardila-Camacho & Contreras-Ramos, sp. nov. |
4 | Body nearly completely dark brown (Fig. |
5 |
– | Body mostly pale orange, with brown marks (Fig. |
7 |
5 | Male gonocoxite IX reduced, fusiform and helical, ventrally attached to lateral arms of gonocoxites XI (Fig. |
Anchieta tinctus Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Male gonocoxite IX well-developed, filiform or thickened with digitiform processes (Figs |
6 |
6 | Male gonocoxite IX with posterior apex ventrally curved, and set with three short processes (Fig. |
Anchieta nothus (Erichson, 1830) |
– | Male gonocoxite IX filiform, lacking processes (Fig. |
Anchieta nebulosus Ardila-Camacho & Machado, sp. nov. |
7 | Forewing costal space narrow (Fig. |
Anchieta bellus (Westwood, 1867) |
– | Forewing costal space medially expanded (Fig. |
8 |
8 | Male sternite IX with broad posteromedial, quadrangular lobe (Fig. |
Anchieta remipes (Gerstaecker, 1888) |
– | Male sternite IX approximately rhomboid, with broad, rounded, posteromedial canal (Fig. |
9 |
9 | Male gonocoxite IX with posterior apex strongly lateroventrally recurved, set with two apical and four preapical, short processes (Fig. |
Anchieta partheniellus (Westwood, 1867) |
– | Male gonocoxite IX with posterior apex laterally or ventrally curved, set with one or two short, preapical processes | 10 |
10 | Male gonocoxite IX apex laterally curved, set with one or two short preapical processes (Fig. |
Anchieta romani (Esben-Petersen, 1817) |
– | Male gonocoxite IX apex ventrally curved, set with one short, preapical process (Fig. |
Anchieta apiculasaeva Thouvenot, 2009 |
Anchieta apiculasaeva
Thouvenot, 2009: 226. Holotype: male, French Guiana, [Régina] Piste de Kaw (
Holotype. French Guiana • ♂; [Régina] Route de KAW pk 3; 03 Mar. 1981; G. Tavakilian leg.;
Brazil • 1 ♂; abdomen cleared and stored in a microvial;
This species exhibits the same general body color pattern as A. bellus, A. partheniellus, A. remipes, and A. romani. It can be separated from A. bellus due to the expanded sub-basal region of the costal space in the forewing and the expanded and oar-shaped hind tibia. Moreover, the gonocoxite IX is short, sinuous, with posterior apex ventromedially gently curved, with two processes, one apical, and a shorter preapical, sometimes a third, more proximal, and shorter is present which separates this species from A. partheniellus. Meanwhile, the gonocoxites X are thickened, with the anterior ½ expanded, which allows to differentiate A. apiculasaeva from A. romani.
Measurements. Male (n = 1). Forewing length: 9.3 mm; Hind wing length: 5.6 mm. Female (n = 1): Forewing length: 8.5 mm; Hind wing length: 5.3 mm.
Coloration (Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Mato Grosso), French Guiana (Matoury, Régina).
This species was previously known solely from the holotype male collected at Régina, French Guiana and was deposited in the private collection of the author of the species (
This species exhibits the same coloration pattern of other species of the genus mimicking stingless bees of the genus Ptilotrigona Moure, 1951 (
Mantispa (Trichoscelia) bella
Westwood, 1867: 502. Holotype: female, Amazonia (
Mantispa (Trichoscelia) eurydella
Westwood, 1867: 501. Holotype: female, Amazonia (
Anisoptera bella
(Westwood, 1867).
Anisoptera eurydella
(Westwood, 1867).
Trichoscelia bella
(Westwood, 1867).
Trichoscelia eurydella
(Westwood, 1867).
Anchieta eurydella
(Westwood, 1867).
Holotype of Mantispa (Trichoscelia) bella.
Brazil • ♀; Amazonas; 1861; H.W. Bates leg.; Type Neur.: No. 8, Mantispa Trichoscelia bella Westwood, HOPE Dept. Oxford; Type Westwood, Trans. Ent. Soc. 1867, p. 502, Coll. Hope Oxon;
Holotype of Mantispa (Trichoscelia) eurydella.
Brazil • ♀; Amazonas; 1861; H.W. Bates leg.; Type Neur.: No. 4, Mantispa Trichoscelia eurydella Westwood, HOPE Dept. Oxford; Type Westwood, Trans. Ent. Soc. 1867, p. 501, Coll. Hope Oxon;
Brazil – Amazonas • 1 ♂; Mamiraua, varzea; 03°02'54"S, 64°51'02"W; 22 Sep. 1993; I.S. Gorayeb and O.T. Silveira leg.;
Colombia • 1 ♀; 1997; Anchieta eurydella det. M. Ohl, 2006;
Suriname • 1 ♂; Brokopondo; 24 Apr. 1965; G.F. Mees leg.; A. remipes, det. L.A. Stange;
This species is distinguished from its congenerics by the narrow costal space of the forewing and the hind tibia moderately expanded and fusiform. On the male genitalia, the gonocoxite IX is narrow, curved, with posterior apex pointed and set with four short, preapical, spine-shaped processes, located on the ventral surface. The gonocoxites X are anteriorly expanded. On the female genitalia, the gonapophyses VIII medial part is ventrally projected, forming a broad, short, blunt canal.
Measurements. Male (n = 1). Forewing length: 8.0 mm; Hind wing length: 4.9 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Amazonas, Pará), Colombia, French Guiana (Saint-Laurent-du-Maroni) (doubtful record), Suriname (Brokopondo).
This species was described based on a female specimen collected by H. Bates. The only collecting data available on the original labels indicate “Amazons” as in many other types of the subfamily. In the present study, further specimens from Suriname, Colombia and Brazil (Amazonas and Pará) were studied and their genital structures and wing venation redescribed. Based on careful comparisons, A. eurydella is herein proposed as a synonym of A. bellus, as both share the same characteristics in the wing shape, venation and hind tibia shape (i.e., fusiform and narrow). The differences in coloration used by
In the phylogeny of the subfamily, A. bellus was recovered as sister of the rest of the species mimicking stingless bees. Like other closely related species, A. bellus has a mimicry pattern resembling meliponine bees of the genus Ptilotrigona (
Mantispa (Trichoscelia) fasciatella
Anisoptera fasciatella
(Westwood, 1867).
Trichoscelia fasciatella
Westwood, 1867.
Plega fasciatella
(Westwood, 1867).
Anchieta fasciatella
(Westwood, 1867).
Holotype
of Mantispa (Trichoscelia) fasciatella. Colombia • ♀; [Magdalena] Santa Marta; 1866; Stevens leg.; Type Neur.: No. 10, Mantispa Trichoscelia fasciatella Westw., HOPE Dept. Oxford; Holotype ♀, Trichoscelia fasciatella Westwood, 1867, ascertained by R.G. Beard, 1968; Type Westwood, Trans. Ent. Soc. 1867, p. 503, Coll. Hope Oxon;
Colombia – Bolívar • 1 ♂; Mompox; 09°14'N, 74°25'W; 33 m; Dec. 1994;
Panama – Canal Zone • 1 ♂; Albrook Forest Site; 30.40 m; 25–26 Jul. 1968; R. S. Hutton leg.; black light trap;
This species is easily recognized by its general body color pattern with orange and dark brown or blackish brown. The forefemur is completely orange, and the forewing has medial and apical, broad, dark amber bands which easily separate A. fasciatellus from T. latifascia. On the hind wing, this species has the intracubital crossvein subparallel to the longitudinal wing axis. On the male genitalia, the gonocoxite IX is long and sinuous, with posterior apex thickened, laterally curved, and equipped with 8–13 apical processes arranged as a brush. On the female genitalia, the gonapophyses VIII medial part is narrow, short, and boat-shaped, with a blunt, dorsally curved process. The spermatheca is short and simple; on the medial section a transparent, digitiform diverticulum is present, and the distal section is progressively expanded and sac-like. The fertilization canal duct is long, spiral-shaped, and the fertilization canal is short, and reniform.
Measurements. Male (n = 8). Forewing length: 9.63–13.56 mm; Hind wing length: 7.4–10.65 mm. Female (n = 1): Forewing length: 11.43 mm; Hind wing length: 8.75 mm.
Coloration (Fig.
Morphology (Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Colombia (Bolívar, Cesar, Magdalena, Santander), Panama (Canal Zone, Darien, Panamá, Panamá Oeste).
This is a rather striking species of Anchieta previously known from Colombia and Panama (
In the phylogeny of Symphrasinae, this species, which is markedly dissimilar from the rest of the species of the genus, was recovered as sister of all other Anchieta. Although it is similar to T. latifascia, both can be easily separated by observing the generic characters of each genus, particularly the structure of the foreleg and the wing venation. Additionally, there are marked differences in the mimicry pattern of both species. As discussed by
Mantispa (Trichoscelia) fumosella
Westwood, 1867: 504. Holotype: male, Amazonia (
Anisoptera fumosella
(Westwood, 1867).
Trichoscelia fumosella
Westwood, 1867.
Anchieta fumosella
(Westwood, 1867).
Anchieta nobilis
Navás, 1909: 484, male, female. Lectotype, sex not indicated, Brazil (
Trichoscelia nobilis
(Navás, 1909).
Holotype of Mantispa (Trichoscelia) fumosella.
[Brazil] • ♂; “fumosella Westwood, male”, “Platymantispa fumosella (Westwood, 1867)”, “Holotype male, Trichoscelia fumosella Westwood, 1867, R.G. Beard 1968, now belongs in genus Platymantispa Rehn”, “Type Westwood, Trans. Entomol. Soc. 1865, p. 504, Coll. Hope Oxon”, “Holotype male, Genitalia of Trichoscelia fumosella Westwood, 1867, prep. R.G. Beard, 1968”, “over? fumosella Westw.”, “Type Neur: No. 13, Mantispa Trichoscelia fumosella Westw. HOPE Dept. Oxford”; Terminalia cleared and stored in a microvial;
Lectotype of Anchieta nobilis.
Brazil • ♂; Goiás, Jataí; Sep.–Nov. 1897; Anchieta nobilis Navás, Longin Navas det. 1907; Lectotype ♂ designated by R.G. Beard, 1968;
Paralectotype of Anchieta nobilis.
Brazil • ♀; Goiás, Jataí; Anchieta nobilis Navás, Longin Navas det. 1907; Lectoallotype ♂ designated by R.G. Beard, 1968;
Brazil – Bahia • 2 ♂; Encruzilhada; 15°32'25"S, 40°50'12"W; 800 m; 10–12 Dec. 2007; J.A. Rafael, P.C. Grossi, D.R. Parizotto leg.; light trap; Anchieta fumosella (Westwood, 1867), det. R.J.P. Machado;
The body coloration pattern of this species may be yellow with dark brown stripes or nearly completely dark brown. The forewing membrane is pale amber, with darker area on distal ½ of costal field, subcostal space, radial space, area between R+M and CuP, and area adjacent to RP. The hind wing is short and narrowly oval. On the male genitalia, the tergite IX has a posterolateral tuft of long setae, which surpasses the posterior margin of ectoproct. The sternite IX is blunt, sclerotized, with lateral margins concave. The gonocoxite IX is thin, short, and sinuous, with posterior apex blade-shaped, with sharp tip, sometimes slightly lanceolate, or with two or three tiny, preapical processes. On the female genitalia, the gonapophyses VIII form a posteromedially projected, tubular process with blunt to pointed apex.
Measurements. Male (n = 5). Forewing length: 11.0–12.8 mm; Hind wing length: 6.6–8.57 mm. Female (n = 3): Forewing length: 7.7–12.15 mm; Hind wing length: 4.6–7.63 mm.
Coloration (Fig.
Chromatic variation
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Bahía, Distrito Federal, Goiás, Minas Gerais, Rio de Janeiro, Paraná, Santa Catarina, Rio Grande do Sul, São Paulo, Tocantins).
This species is so far known only exclusively from Brazil and has been previously recorded in the states of Bahia, Distrito Federal, Goiás, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina, Tocantins, and Rio Grande do Sul (
Anchieta fumosellus expresses a Batesian mimicry pattern with wasps of the family Vespidae (
In the phylogeny of Symphrasinae, this species was recovered as an intermediary species between the braconid mimicking A. fasciatellus and the clade containing all the bee mimicking species. This evolutionary pattern matches with the phylogeny of Hymenoptera, in which the parasitoid wasps of the family Braconidae are part of a large clade that diverged first in the phylogeny of Hymenoptera, then, within Aculeata the Vespoidea had an early divergence, while the bees or Antophila evolved as a highly specialized group within Apoidea (
This species can be recognized by the pale amber area and the anterior region of the forewing, which resembles the folding of the forewings of vespid wasps, the short and narrow hind wing, and the male genitalia. The male genitalia of this species are similar to that of A. nebulosus as both have a tuft of long setae on the lateral region of the tergite IX, arched goncoxites XI, and narrow and short male gonocoxites IX lacking digitiform processes. By contrast, the female goncoxites+gonapophyses VIII are similar to those of A. fasciatellus, but the spermatheca is similar to that of the species included within the bee-mimicking clade.
Brazil, Espírito Santo: Linhares, Estrada, Fazenda St. Terezinha, 50 m a.s.l., Oct. 2004, P. Grossi leg.
Holotype
male, pinned, with genitalia in a separate microvial. Original label: “Brazil, Espírito Santo, Linhares, Estrada, Fazenda St. Terezinha, 50 m a.s.l., X.2004, P. Grossi leg. (coleção E. & P. Grossi)”;
The specific epithet of this species comes from the Latin nebula meaning misty or clouded, in allusion to the wing color pattern of this species. An adjective in the nominative case.
This species has a general body color pattern very similar to that of A. nothus. The body is mostly dark brown; the forewing is dark amber at base, and there are irregular amber areas on membrane surrounding 1m-cu, subcostal veinlets of distal ½ of costal space, medial and apical region of subcostal field, 1ra-rp, and part of M fork, rarp1, and 2m-cu base. The hind wing has a broad, amber, transverse band on proximal 1/3 of wing. The tergites of the abdominal segments III–VI have posteromedial, keeled processes, which are not enlarged. The male tergite IX has posterolateral tuft of long setae, which surpass the posterior margin of the ectoproct. The sternite IX is quadrangular with posterior margin ventrally curved. The gonocoxite IX is short, filiform, with posterior apex pointed. The surrounding membrane of the posterior apex of the gonapophyses X is sclerotized and forms a bilobed structure set with minute granules. On the female genitalia, the gonapophyses VIII form a cubic structure, with concave sides, and with an anteromedial incision.
Measurements. Male (n = 1). Forewing length: 9.5 mm; Hind wing length: 5.9 mm. Female (n = 1): Forewing length: 13.0 mm; Hind wing length: 7.7 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Bahía, Espírito Santo).
This is a new species described from Bahía and Espírito Santo in Brazil. Naked eye, this species could be identified as Anchieta nothus (Erichson, 1939) as both species have exactly the same general coloration pattern, including the extensive shaded areas on the wings. Nonetheless, the arrangement of the processes on the abdominal terga can be used as a first clue to separate both species. Furthermore, the genital sclerites are completely different, the superficial resemblance between both just represents an additional case of cryptic species, a rather common phenomenon in Symphrasinae. In the phylogeny of the subfamily, this species was recovered as sister of A. nothus within the clade of bee mimicking species of Anchieta. Regarding the mimicry pattern, this species appears to mimic the genus Trigona Jurine, 1807 (
Mantispa notha
Erichson, 1839: 170. Holotype: male, Brazil (
Trichoscelia notha
(Erichson, 1839).
Symphrasis notha
(Erichson, 1839).
Anisoptera notha
(Erichson, 1839).
Anchieta notha
(Erichson, 1839).
Holotype. Brazil • ♂; “Virmum Leg., notha Er.”, “Holotype of Mantispa notha Erichson 1839, R.G. Beard” [red label], “holotypus Nr.” [red label], “123”, “prep. R.G. Beard, 1968, Holotypus Nr”; [Male genitalia and left forefemur cleared];
Brazil • 1♀; Bahía; Hagen, Winthom leg.;
This species has the general body color pattern dark brown. The forewing has pale amber, irregular areas at base, area around the 1r-m, proximal ¼ of wing, distal subcostal veinlets, apical area of subcostal field, and area surrounding 1ra-rp. The hind wing exhibits a broad, transverse, pale amber band on proximal 1/3. The tergites of abdominal segments III–VI have prominent, posteromedial, keeled processes. On the male genitalia, the sternite IX is posteromedially produced into a blunt, elongated lobe. The gonocoxite IX is short, narrow, sinuous, with posterior apex ventrally curved, and set with three short apical processes. On the female genitalia, the sternite VII appears as two lateral, broad, subtrapezoidal plates (gonocoxites VII), which are posteromedially fused through a thin bridge. The gonapophyses VIII medial part forms a cubic structure, with concave sides, and with an anteromedial incision.
Measurements. Male (n = 2). Forewing length: 10.3–10.7 mm; Hind wing length: 5.9–6.4 mm. Female (n = 1): Forewing length: 12.0 mm; Hind wing length: 7.4 mm.
Coloration
(Fig.
Morphology (Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Bahía, Minas Gerais).
Prior to this work, this species was known from Brazil, although information about its distribution within the country is not available (
Regarding the phylogenetic relationships of this species, it was recovered as sister to A. nebulosus in the clade comprising the bee-mimicking species. As mentioned above for A. nebulosus, A. nothus has essentially the same coloration pattern which consists of a dark brown body and wings with extensive amber areas. The hind tibia of both species is markedly expanded and flattened, but in A. nothus a broad, yellow, medial area is present. The morphology and coloration of this species represents a mimicry pattern whose model is probably the meliponine genus Trigona Jurine, 1807 (
Mantispa (Trichoscelia) partheniella
Westwood, 1867: 501. Lectotype male, Paralectotype female, Amazonia (
Trichoscelia partheniella
(Westwood, 1867).
Anchieta partheniella
(Westwood, 1867).
Lectotype. [Brazil] • 1 ♂; Amazonas; 1861; Bates leg.; “Platymantispa partheniella (Westwood, 1867)”, “Lectotype male, Trichoscelia partheniella Westwood, 1867 by R.G. Beard, 1968”, “partheniella Westwood, male”, “Type Westwood, Trans. Entomol. Soc. 1865, p. 501, Coll. Hope Oxon”, “Type Neur: No. 61/2, Mantispa Trichoscelia partheniella Westwood, HOPE Dept. Oxford”; right foreleg and abdomen cleared and stored in a microvial;
Brazil – Amazonas • 1 ♂; Ipixuna, Río Gregorio, com. Lago Grande; 07°10'11.7"S, 70°49'10.3"W; 18–23 May. 2011; J.A. Rafael, J.T. Cámara, R.F. Silva, A. Somavilla, R. Ale-Rocha leg.; Varredura;
Colombia • 1 ♀; Meta, Cabaña Carrillo, PNN Sierra de la Macarena; 3°21'N, 73°56'W; 460 m, 29 Dec. 2003; W. Villalva leg.; Malaise trap, m-4240; IAvH.
Ecuador • 1 ♀; Napo, near Tena Jatun Sacha; 450 m; G. Beccaloni leg.; BMNH(E) 1241394;
Suriname • 1 ♀; Republiek; 13 Nov. 1961; P.H.v. Doesburg leg.; A. romani, det. Geisjkes 1970;
Venezuela • 1 ♂; Amazonas, T.F., Cerro de la Neblina, Basecamp, near Río Baria; 0°50'N, 66°10'W; 140 m; 18 Feb.1985; P.J. and P.M. Spangler, R. Faitoute, W. Steiner leg.; at black light in rainforest clearing; Anchieta bella, det. O.S. Flint, 1986; USNMENT01541897; USNM. • 1 ♀; same data as for preceding; USNMENT01541896; USNM.
This species has a general body coloration pattern of bright orange with brown areas, which is also present in A. apiculasaeva, A. bellus, A. remipes, and A. romani. The forewing costal space is slightly widened medially. The hind tibia is conspicuously widened and laterally flattened. On the male genitalia, the gonocoxites IX is short, with posterior apex strongly recurved lateroventrally, and set with two apical and four preapical, short processes on outer surface. The gonocoxites X form a hourglass-shaped sclerite, whose anterior ½ is expanded, and dorsally bent. On the female genitalia, the gonapophyses VIII medial part appears as a broad canal, with, ventral, medial depression, and distal margin rounded, and ventrally curved.
Measurements. Male (n = 3). Forewing length: 9.3–10.3 mm; Hind wing length: 6.0–6.56 mm. Female (n = 3): Forewing length: 9.3–9.7 mm; Hind wing length: 5.6–5.9 mm.
Coloration (Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Amazonas, Pará, Rondônia), Colombia (Meta), Ecuador (Napo), Suriname, Venezuela (Amazonas).
Lectotypes (male and female) of this species deposited in the Oxford University Museum have only “Amazons” as collecting data.
In the phylogeny of the subfamily, A. partheniellus was recovered as sister to A. nebulosus + A. nothus. However, this species exhibits the same mimicry pattern as A. apiculasaeva, A. bellus, A. remipes, and A. romani whose model is probably the meliponine genus Ptilotrigona Moure, 1951 (
The distinction between the species exhibiting this mimicry pattern may be difficult as all of them have a rather similar morphology, although A. partheniellus can be separated from other species with the same coloration by the arrangement and number of processes on the male gonocoxites IX, the enlarged and curved gonocoxites X and the complex spermatheca.
Anisoptera remipes
Gerstaecker, 1888: 120. Holotype: male, Colombia, Bogotá (
Trichoscelia remipes
(Gerstaecker, 1888).
Anchieta remipes
(Gerstaecker, 1888).
Holotype. Colombia • ♂; “remipes Gerst.* Columb. Thieme”, “An. remipes Gerst.* Columb. Thieme”, “Zool. Mus. Greifswald II 27487”, “Anchieta remipes det. R. Hall (HNRS) 1988”; Male genitalia cleared and stored in a microvial;
This species is separated from other of the genus with similar body coloration pattern (A. apiculasaeva, A. bellus, A. partheniellus, and A. romani), because the forecoxa has the proximal 2/3 brown and distal 1/3 orange. The forewing costal space is slightly widened medially, and the hind tibia is oar-shaped, notably expanded, and laterally flattened. On the male genitalia, the sternite IX has a broad posteromedial, quadrangular lobe. Like in A. tinctus, the gonocoxite IX of this species are fusiform and reduced, and the gonostyli X dorsally recurved and anteroventrally projecting.
Measurements. Male (n = 1). Forewing length: 8.9 mm; Hind wing length: 5.0 mm.
Coloration
(Fig.
Morphology
(Fig.
Abdomen. Cylindrical, without keeled processes.
Male genitalia
(Fig.
Colombia (Cundinamarca?).
This species is so far known only from the original description by
Regarding the phylogenetic relationships of this species, despite presenting the same mimetic coloration pattern of species like A. bellus, A. apiculasaeva, A. partheniella, and A. romani, this species was recovered as sister of A. tinctus from Costa Rica. Both species share many characters of the wing shape and venation as well as of the male genitalia. Nonetheless, A. remipes can be differentiated from A. tinctus by the body coloration and morphology of the sternite IX, while the overall genitalic morphology markedly differs from other species mimicking stingless bees of the genus Ptilotrigona.
Anisoptera romani
Esben-Petersen, 1917: 14. Holotype: male, Brazil, Rio Autaz (
Trichoscelia romani
(Esben-Petersen, 1917).
(non) Anchieta bella (Westwood, 1867).
Holotype. Brazil • ♂; Amazonas, Río Autaz; Sept.; Roman leg.; “Anisoptera romani, male, det. Esben-Petersen (
Peru • 1 ♀ 2 ♂; San Martín, Tarapoto Urku, nr. Boca Toma; 400 m. a.s.l.; 10 Jul. 2012; C. Rasmussen leg.; nest collected;
This species is remarkably similar to A. apiculasaeva and A. partheniella because of the general color pattern of orange with brown marks. Additional similarities include the slightly widened forewing costal space near the medial region, and the notably expanded and laterally flattened hind tibia. On the male genitalia, the sternite IX is U-shaped with posterior ½ glabrous, medially convex, and laterally concave, forming a canal. The gonocoxite IX is short, sinuous, with posterior apex laterally recurved, and set with 1–3 longitudinally arranged, preapical processes on the outer surface. An important distinction from the aforementioned species is the shape of the gonocoxites X, which form a triangular sclerite, with anterior apex laterally flattened and not expanded. On the female genitalia, the gonapophyses VIII medial part forms an enlarged canal, with ventral depression and apex ventrally curved, and rounded.
Measurements. Male (n = 2). Forewing length: 9.0–9.2 mm; Hind wing length: 5.2–5.4 mm.
Coloration
(Fig.
Morphology (Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Brazil (Amazonas), Peru (San Martín).
This species was synonymized with A. bellus by
Regarding its distribution, the collecting data of the type of A. romani appears as “Rio Autaz, Amazon”, locality that was corrected by
In a recent publication by
Regarding the phylogenetic affinities of this species, it was recovered in a polytomy with A. apiculasaeva and a clade containing A. remipes, A. tinctus, A. partheniellus, A. nebulosus, and A. nothus. Considering the morphology of the male genitalia, this species is closely related with A. apiculasaeva. Both species can be separated by the morphology of the male gonocoxites IX and gonocoxites X.
French Guiana, Camopi: Mont St-Marcel de la Haute, 2°23'9.816"N, 53°1'29.964"W, 13 Dec. 2014, light trap, J. Touroult leg.
Holotype
male, pinned. Original label: “French Guiana, Camopi, Mont St-Marcel de la Haute, 2°23'9.816"N, 53°1'29.964"W, 13 Dec. 2014, light trap, J. Touroult”
This species is separated from the yellow morph of A. fumosellus because the frons is completely yellow, the pronotum has a dark brown posterior, inverted U-shaped mark, and the pteropleura is nearly entirely yellow. Furthermore, the hind tibia is slightly widened. On the male genitalia, the gonocoxite IX is pointed, and lacks processes. On the female genitalia, the gonocoxites + gonapophyses VIII form an enlarged, chamber-shaped structure.
First author dedicates this species after his daughter Lauren Sophia Ardila, born on 29 of December of 2015.
Measurements. Male (n = 2). Forewing length: 11.7–12.3 mm; Hind wing length: 6.8–7.0 mm. Female (n = 1): Forewing 10.5 length: mm; Hind wing length: 6.1 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
French Guiana (Camopi, Maripasoula).
This species is known solely from two provinces of French Guiana. Based on its coloration pattern, wing morphology and shape of the hind tibia this species appears to be sister to A. fumosellus. However, considering the genitalic morphology this species could be sister of the clade of bee- mimicking species of Anchieta. The coloration pattern of A. sophiae sp. nov. is markedly similar to that of the yellow form of A. fumosellus, although its model is likely Agelaia myrmecophila (Ducke, 1905) (Vespidae: Polistinae) based on overall coloration and distribution of these species.
Costa Rica, Heredia: Est. Biol. La Selva, Arboleda, 10°26'N, 84°01'W, 50–150 m, 28 Jan. 1998, R. Vargas C. leg.
Holotype
male, pinned. Original label: “Costa Rica, Heredia, Est. Biol. La Selva, Arboleda, 10°26'N, 84°01'W, 50–150 m, 28 Jan. 1998, R. Vargas C. leg.”, “INBIOCRI002286798”,
• 1 ♂; Sector Cocori; 150 m; Feb. 1993; E. Rojas leg.; L-N 286000, 567500, CRI001309729;
This species has the body nearly uniformly dark brown, the antennal flagellum has seven yellow preapical flagellomeres. The forewing membrane is mostly pale amber, with darker areas on basal ½ and area adjacent to proximal part of pterostigma; there are hyaline, fenestrated, whitish areas on costal and subcostal fields, anterior radial cells, mcu2, mcu3, cuacup1, and second a1a2; wing base, anal region and jugal lobe are dark amber. The hind tibia is conspicuously expanded and laterally flattened, with most of the surface including the apex pale brown to orange. On the male genitalia, the sternite IX is quadrangular, dorsally canaliculate, and curved ventrally. The gonocoxite IX is reduced, appearing as a filiform, thin sclerite, ventrally attached to lateral arms of gonocoxites XI. The gonostyli X are whip-like, dorsally curved and posteroventrally directed, forming an apical loop. The gonapophyses X are posteriorly fused forming a long, blunt process. On the female genitalia, the gonapophyses VIII medial part is composed by quadrangular plates that form a canal, whose ventral surface has a longitudinal depression. The bursa copulatrix is thin, moderately sclerotized, and remarkably long, forming a loop.
The specific epithet of this species is a noun in apposition in allusion to the wing coloration pattern of this species.
Measurements. Male (n = 4). Forewing length: 10.42–11.85 mm; Hind wing length: 5.81–7.33 mm. Female (n = 1): Forewing length: 10.20 mm; Hind wing length: 6.63 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Costa Rica (Heredia, Limón).
This species is known only from two provinces of Costa Rica. Regarding its phylogenetic relationships, this species was recovered sister to A. remipes as both share many characteristics on the wings and male genitalia. Specifically, the genital sclerites in this pair of species is highly modified, with reduced male gonocoxites IX and fused gonapophyses X at the posterior apex. Besides the coloration pattern, this species can be separated from A. remipes by the morphology of the male sternite IX. This species probably mimics Melipona fuliginosa Lepeletier, 1836, a species of stingless bee with similar coloration and size found in Costa Rica.
Plega Navas, 1928: 326. Type species: Symphrasis signata Hagen, by original designation.
This genus differs from the other Symphrasinae by having a slightly to strongly domed region of the vertex; compound eyes generally are as wide as ½ of the interocular distance at toruli level. The pronotum is typically as long as wide with anterior, medial, and posterior regions unevenly raised, and covered with pedicellate thickened setae; the postfurcasternum is generally ventrally fused, collar-like (unfused in P. insolita). On the forefemur, the closing surface presents both rows of processes fully developed in one species, while the rest has the anteroventral row reduced to proximal ½ and apical portion; the anteroventral row of processes generally presents the basal, primary, curved process; the posteroventral row has two primary processes on the proximal ½ of the femur length, and the proximal part of the row has two spine-shaped processes that may be secondary or tertiary; the distal region of the row is slightly keeled and composed mostly of tubercle-shaped specializations; between the cuticular processes, there are stinger-shaped setae. The rows of thickened setae with globular base adjacent to each row of processes are modified, with the posteroventral row reduced to the apical portion, while the anteroventral row is at least present on the distal ½ of the femur. The fore basitarsus has the lanceolate process is generally long, reaching the middle of the fourth tarsomere, although in one species it is short, reaching the middle of the third tarsomere. The distal section of the spermatheca generally has an apical diverticulum, and the proximal part of the fertilization canal duct is triangular and concave.
Head. Diamond-shaped or quadrangular in frontal view, rugose, region of vertex domed over compound eyes, paraocular area concave; coronal suture generally distinct. Antenna moniliform to filiform, basal flagellomeres from discoidal to as long as wide in frontal view. Compound eye hemispheric, as wide as ½ of interocular distance at toruli level. Thorax. Pronotum either as long as wide or slightly longer than wide, a groove adjacent to lateral and distal margins is generally present; anterior margin, and medial and posterior regions are generally unevenly raised, covered with long, thickened, generally pedicellate setae; postfurcasternum generally unpaired, and ventrally fused. Mesonotum wider than long, with long, thickened, pedicellate setae, metanotum generally 2.5× as wide as long, glabrous. Foreleg. Coxa slightly shorter than femur, cylindrical, with pedicellate setae or not. Trochanter trapezoidal, generally with protuberance on anterior surface. Femur narrow to robust, closing surface covered generally with microtrichia; posteroventral row processes fully developed, slightly carinated on distal ½ or ⅓, composed of tubercle-shaped specializations and stinger-shaped setae; two primary, spine-shaped specializations are present on proximal ½; secondary or tertiary processes are frequent between the primary ones, and on proximal portion of the row; adjacent row of thickened setae with globular base typically reduced to distal ¼ of femur length; anteroventral row generally reduced to proximal ½ and apex, fully-developed in one species, composed of tubercle-shaped integumentary specializations, and stinger-shaped setae; basal, primary, spine-shaped process is generally present, curved (absent in P. insolita); adjacent row of thickened setae with globular base present on distal ½–⅘. Tibia nearly as long as femur, setose, curved or straight, ventrally keeled, with a row of prostrate setae; anterior surface with a patch of clavate setae at apex. Basitarsus generally with long lanceolate process surpassing distal margin of third tarsomere (reaching to the middle of the third tarsomere in P. insolita), equipped with a plug-shaped Stitz organ at apex; basal ½ with a row of prostrate setae on ventral surface, and patch of clavate setae on anterior surface; second tarsomere articulated on anterior surface of basal ½ of basitarsus, longer than third and fourth tarsomeres together; pretarsal claws simple. Mid- and hind leg. Unmodified, basitarsus typically 3.5–4× as long as wide. Wings. Forewing oval, trichosors present along wing margin, except at base; costal space medially slightly widened, humeral vein generally branched, subcostal veinlets generally simple, sometimes with some forked; pterostigma elongated and narrow; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; radial space with two crossveins; rarp2 curved; RP separated or close to separation of M and R, M fork near this separation; 1r-m between RP base and M fork forming a trapezoidal cell; RP with single series of gradate crossveins; CuP basally angled, approaching A1. Hind wing oval, smaller and narrower than forewing; costal space narrow and reduced, C and Sc fused at proximal ¼ of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, curved, narrow; radial space with single crossvein, generally straight; RP with single gradate series. M forked beyond R fork; Cu deeply forked, CuA distally branched, first branch generally candelabrum-shaped; intracubital vein subparallel to longitudinal wing axis; CuP distally anteriorly bent near posterior wing margin, pectinate. Cubitoanal space generally with two crossveins. Abdomen. unmodified.
Male genitalia. Sternites VIII and IX not fused; sternite IX pentagonal or trapezoidal, posteromedial lobe dorsally canaliculated, generally triangular. Gonocoxites IX generally long, thin to thickened, with or without apical processes. Ectoproct generally ovoid or trapezoidal, sometimes with pedicellate setae, sometimes developed as prominent spine-shaped structures; ventral surface typically sclerotized, with anterior rounded, flattened lobe that is continuous with ventromedial, curved sulcus. Gonocoxites X unpaired, forming a ventrally canaliculated, bar-shaped sclerite, anterior apex generally slightly expanded, straight or dorsally bent; posterior apex with paired dorsal and lateroventral, preapical processes, articulated to gonostyli X and gonapophyses X, respectively; gonostyli X with thickened base, set with curved, lateral processes; the rest of the structure from short and recurved to long and coiled, forming one or two internal loops, before protruding from abdomen. Gonapophyses X paired, straight, thin, forming a V-shaped structure; gonapophyses joined by membrane that forms a prominent covering over gonostyli X base, which may be medially sclerotized or not. Gonocoxites XI U-shaped, medial lobe elaborated, with two differentiated parts, a dorsal arch and lobe and ventral convex area, continuous with a process; between these parts a less sclerotized is present; lateral arms of gonocoxites XI straight or sinuous, with anterior apex bent or expanded. Hypandrium internum triangular, keeled, with two lateral fins.
Female genitalia. Sternite VII (gonocoxites VII) generally trapezoidal, in a species (P. drepanicoides) as two lateral, medially joined, trapezoidal plates. Tergite VIII narrower medially than laterally, enclosing the spiracle of the segment VIII. Gonocoxites VIII forming a narrow plate; gonapophyses VIII medial part keel-shaped, with posteromedial bilobed or bifid process; lateral part as a trapezoidal or oval plate, hidden under tergite IX + ectoproct. Tergite IX + ectoproct triangular, ovoid or D-shaped. Gonocoxites IX long, straight and narrow; gonapophyses IX sometimes present as tiny sclerites, located basally, on inner surface of gonocoxites IX. Bursa copulatrix variable in shape, with sclerotized portion or not, short to long, sometimes with an enlarged chamber-shaped part. Spermatheca short and simple to remarkably long and entangled; proximal section short to long and thin, generally forming several coils; medial section thin, entangled or forming a long spiral; distal section moderately expanded, wider than proximal and medial sections, generally forming a blunt diverticulum at apex; fertilization canal duct short, proximal part triangular and concave; fertilization canal generally elongated, pod-shaped, covered with microfilaments.
1. P. acevedoi sp. nov. (Mexico)
2. P. banksi Rehn, 1939 (Mexico, USA)
3. P. bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov. (Ecuador)
4. P. dactylota Rehn, 1939 (Mexico, USA)
= P. fratercula Rehn, 1939, new synonym
5. P. disrupta Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
6. P. drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
7. P. duckei Penny, 1982 (Brazil)
8. P. flammata Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
9. P. fumosa Linsley & MacSwain, 1955 (Mexico)
10. P. hagenella (Westwood, 1867) (Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Mexico, Nicaragua, Panama, Trinidad and Tobago, Venezuela)
= P. beardi Penny, 1982, new synonym
= P. melitomae Linsley & MacSwain, 1955, new synonym
= Mantispa cognatella Westwood, 1867
11. P. insolita Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
12. P. lachesis Ardila-Camacho & Contreras-Ramos, sp. nov. (Guatemala)
13. P. longicornis Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
14. P. megaptera Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
15. P. mixteca Ardila et al., 2019 (Guatemala, Mexico)
16. P. obtusa Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico, USA)
17. P. oswaldi Ardila-Camacho & Contreras-Ramos, sp. nov. (Mexico)
18. P. paraensis Penny, 1982 (Brazil)
19. P. pseudohagenella Ardila-Camacho & Contreras-Ramos, sp. nov. (Costa Rica, Guatemala)
20. P. radicaudata Ardila-Camacho & Contreras-Ramos, sp. nov. (Bolivia, Peru)
21. P. signata (Hagen, 1877) (Mexico, USA)
22. P. sonorae Ardila et al., 2019 (Mexico, USA)
23. P. spinosa Ardila et al., 2019 (Mexico)
24. P. stangei Ardila et al., 2019 (Mexico)
25. P. vangiersbergenae Ardila-Camacho, sp. nov. (Guatemala, Mexico)
26. P. variegata Navás, 1928 (Mexico), nomen dubium
27. P. yucatanae Parker & Stange, 1965 (Guatemala, Honduras, Mexico)
28. P. zikani Navás, 1936 (Brazil)
Summarized in
Uncertain, and not specified in the original description of the genus by
1 | Forefemur remarkably elongated and narrow in lateral view (Fig. |
2 |
– | Forefemur narrow and short or moderate- to strongly robust in lateral view (Fig. |
5 |
2 | Forefemur closing surface with anteroventral row of integumentary specializations fully developed (Fig. |
Plega radicaudata Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Forefemur closing surface with anteroventral row of integumentary specializations reduced to proximal region and apex | 3 |
3 | Antennal flagellum brown with eight pale, preapical flagellomeres (Fig. |
Plega zikani Navás, 1936 |
– | Antennal flagellum uniformly brown | 4 |
4 | Antennal flagellum with basal flagellomeres as wide as long (Fig. |
Plega paraensis Penny, 1982 |
– | Antennal flagellum with basal flagellomeres 3× as wide as long and subconical (Fig. |
Plega duckei Penny, 1982 |
5 | Posterior surface of forefemur pale with numerous brown dots (Fig. |
6 |
– | Posterior surface of forefemur pale with brown areas, which may or may not be interconnected (Fig. |
17 |
6 | Pale medial area of forewing pterostigma with anterior notches (Fig. |
7 |
– | Pale medial area of pterostigma with different pattern, or if notched, additional smaller pale areas are present (Fig. |
8 |
7 | Posterior apex of male gonocoxite IX equipped with long and thin preapical process on inner surface, and 2–4 shorter, apical processes (Fig. |
Plega hagenella (Westwood, 1867) |
– | Posterior apex of male gonocoxite IX equipped with an elongate preapical process on inner surface and seven or eight short processes on outer surface, arranged in a single plane (Fig. |
Plega bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov. |
8 | Supraantennal region unmodified, without lateral protuberant areas (Fig. |
Plega lachesis Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Supraantennal region with lateral protuberances covered with reclined setae (Fig. |
9 |
9 | Male gonocoxites IX lacking digitiform processes (Fig. |
10 |
– | Male gonocoxites IX se with 3–12 digitiform processes | 15 |
10 | Antennal flagellum completely brown (Fig. |
11 |
– | Antennal flagellum with 3–5 pale, preapical flagellomeres (Fig. |
12 |
11 | Pale area of pterostigmata enlarged, cream colored (Fig. |
Plega obtusa Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Pale area of pterostigmata small (Fig. |
Plega disrupta Ardila-Camacho & Contreras-Ramos, sp. nov. |
12 | Forewing hyaline (Fig. |
Plega oswaldi Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Forewing with area adjacent to crossveins and area between apical twigging amber (Fig. |
13 |
13 | Area adjacent to frontal sutures strongly sunken (Fig. |
Plega spinosa Ardila et al., 2019 |
– | Area adjacent to frontal sutures slightly sunken (Fig. |
14 |
14 | Antennal scape 2× as long as wide at middle (Fig. |
Plega acevedoi Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Antennal scape 3.5× as long as wide at middle (Fig. |
Plega longicornis Ardila-Camacho & Contreras-Ramos, sp. nov. |
15 | Forefemur remarkably robust (Fig. |
Plega pseudohagenella Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Forefemur not remarkably robust (Fig. |
16 |
16 | Head with vertexal region patterned with narrow pale and brown stripes (Fig. |
Plega stangei Ardila et al. 2019 |
– | Head with vertexal region with lateral broad brown stripes (Fig. |
Plega vangiersbergenae Ardila-Camacho, sp. nov. |
17 | Basal antennal flagellomeres as wide as long (Fig. |
18 |
– | Basal antennal flagellomeres > 2× as wide as long (Fig. |
19 |
18 | Antennal flagellum entirely brown (Fig. |
Plega drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Antennal flagellum with 3–5 pale preapical flagellomeres (Fig. |
Plega yucatanae Parker & Stange, 1965 |
19 | Forecoxa with fine and long setae on anterior and posterior surfaces not arising from protuberant bases; forefemur noticeably robust (Fig. |
20 |
– | Forecoxa with thickened setae arising from protuberant bases; forefemur not noticeably robust (Fig. |
21 |
20 | Body mostly dark reddish brown (Fig. |
Plega fumosa Linsley & MacSwain, 1955 |
– | Body with a mixture of dark brown and yellow (Fig. |
Plega mixteca Ardila et al. 2019 |
21 | Anteroventral row of integumentary specializations lacking curved, basal, primary process (Fig. |
Plega insolita Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Anteroventral row of integumentary specializations with curved, basal, primary process (Fig. |
22 |
22 | Basal antennal flagellomeres > 3× as wide as long (i.e., discoidal) (Fig. |
23 |
– | Basal antennal flagellomeres 2× as wide as long (Fig. |
24 |
23 | Body color pattern pale and dark reddish brown (Fig. |
Plega banksi Rehn, 1939 |
– | Body color pattern pale and dark brown (Fig. |
Plega dactylota Rehn, 1939 |
24 | Region of the vertex often with a well-defined, arrow-shaped, pale area surrounding the coronal suture (Fig. |
25 |
– | Region of the vertex with pale area surrounding the coronal suture not arrow-shaped (Fig. |
26 |
25 | Forewing distinctively enlarged (Fig. |
Plega megaptera Ardila-Camacho & Contreras-Ramos, sp. nov. |
– | Forewing not distinctively enlarged (Fig. |
Plega sonorae Ardila et al., 2019 |
26 | Forefemur anterior surface completely dark brown (Fig. |
Plega signata (Hagen, 1877) |
– | Forefemur anterior surface brown with pale base (Fig. |
Plega flammata Ardila-Camacho & Contreras-Ramos, sp. nov. |
Mexico, Morelos: Balneareo de los Manantiales, 18°28.059'N, 99°9.315'W, 2253 m, 20.IX.2018, C. Rodríguez Leg., selva baja caducifolia.
Holotype
male, pinned. Original label: “Mexico, Morelos, Balneareo de los Manantiales, 18°28.059'N, 99°9.315'W, 2253 m, 20 Sep. 2018, C. Rodríguez Leg., selva baja caducifolia;
Mexico – Morelos • 1 ♂; Tlaquiltenango, Huautla; 18°23'08.4"N, 99°03'04"W, 1023 m, 28–31 Jan. 2009; V.H. Toledo leg.; selva baja caducifolia; light trap;
This species is closely related to P. longicornis and P. spinosa as the area adjacent to frontal sutures is sunken, and the supra-antennal is raised laterally and set with abundant, fine, reclined setae. Plega acevedoi can be separated from them by the antennal scape 2× as long as wide at middle with dorsal surface set with stout setae near the distal margin in males. This species exhibits the femoral posterior surface pale with brown dots, while the anterior surface is dark brown, except basal ¼ pale. On the male genitalia, the gonocoxites IX lack digitiform processes as in P. longicornis, P. oswaldi, and P. spinosa, and the ectoproct is equipped with spine-shaped setal bases on the posteroventral surface; the gonostyli X are posteriorly recurved at mid length before protruding from abdomen, with an abruptly narrowed and pointed apex. The ventral part of the gonocoxites XI medial lobe has a prominent, rounded, transversal ridge and ventrally it forms a pointed, curved process. Moreover, in the female genitalia, the gonocoxites + gonapophyses VIII form two lateral quadrangular plates with a tubular, medial process whose apex is shallowly bifid; the spermatheca is short and simple, with undifferentiated proximal and medial sections; the proximal-most part and the apex form a blunt diverticulum each.
This species is dedicated to Fernando Acevedo Ramos, Spanish entomologist and expert in Myrmeleontidae, who supported the first author in many ways. This allowed the development of much of this revision during its first stages.
Measurements. Male (n = 6). Forewing length: 6.6–14.9 mm; Hind wing length: 5.3–12.0 mm. Female (n = 5). Forewing length: 11.2–14.3 mm; Hind wing length: 9.4–11.4 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Mexico (Morelos, Puebla, Veracruz).
This species is closely related to P. spinosa and P. longicornis based on the sexual dimorphism of the antennae, morphology of the supra-antennal area, coloration of the forefemur, the overall morphology of the male terminalia and genitalia, plus the female genitalia. Furthermore, its distribution in central Mexico (Morelos, Puebla and Veracruz) supports its phylogenetic affinities. This new species, together the aforementioned ones form a complex of species together P. oswaldi and P. obtusa.
Plega banksi
Rehn, 1939: 248. Holotype: male, USA, Arizona (
Paratypes. USA • 1 ♂; Arizona, Oracle 14 m E.; 27 Jul. 1924; E.P.V. Van Duzee leg.; “Plega banksi Rehn, Paratype”;
Mexico – Sinaloa • 2 ♀; 16 mi S Guamuchil; 20 May. 1962; F.D. Parker & L.A. Stange leg.; Plega fratercula Rehn det. L. Stange, 1991;
USA – Arizona • 1 ♂; Madera Canyon; 24 Jul. 1976; B.K. Dozier leg.;
Distinguished from other species in the genus by the dark-reddish brown body coloration pattern. The vertexal region has a pale arrow-shaped mark around the coronal suture. The basal antennal flagellomeres are discoidal. The mesonotum often has a wide, pale, medial band. The forewing generally exhibits a zigzagged amber mark connecting MA and MP forks, terminal part of MA and 2r-m and 1ma-mp. The area between CuP and posterior wing margin is often amber. The sternite IX is pentagonal, it is posteromedially elongated, blunt, surpassing posterior margin of ectoproct. The gonocoxite IX is short, thickened, and sinusoid, with posterior apex laterally curved, and equipped with 4–7 processes of different lengths. The dorsal part of the median lobe of gonocoxites XI is rounded and anteriorly curved lobe; the ventral part is convex, and continuous with a ventral, curved, prominent hook-shaped process whose apex is covered with microspinulae. The gonapophyses VIII of female have a chamber-shaped medial part, posteromedially is set with an elongated Y-shaped process. The female gonocoxites IX are nearly as long as the whole abdomen. The bursa copulatrix is short, with the proximal part strongly expanded, ovoid, and sclerotized, continuous with a membranous and striated part, which is abruptly narrowed.
Measurements. Male (n = 10). Forewing length: 7.8–14.6 mm; Hind wing length: 6.0–11.5 mm. Female (n = 7): Forewing length: 7.3–14.5 mm; Hind wing length: 5.6–11.7 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Mexico (Sinaloa, Sonora), USA (Arizona).
This species appears to have a more restricted distribution than the closely related P. dactylota, being distributed in northern Mexico (Sinaloa and Sonora) and Arizona. Specimens of Sinaloa are slightly different to those from Arizona, particularly in the overall morphology of male genitalia, although there are not enough differences to consider the specimens from the former location as a different species. Such difference is interpreted as part of intraspecific variation, probably related to size or geographic distribution. Records from the Mexican state of Chihuahua presented by
Specimens of this species may be difficult to separate from P. dactylota, particularly when only females are available. However, the general dark reddish brown body coloration pattern with a cream medial band on the mesonotum, the strongly marked forewing and the longer ovipositor are characters that allow to make a relatively reliable distinction between both.
Ecuador, [Guayas]: Guayaquil, Rosemberg leg.
Holotype
male, pinned, with genitalia in a separate vial. Original label: “Ecuador, Guayaquil, Rosemberg coll., Acq. 1903”;
This species presents the antennal flagellomeres as long as wide at proximal region of flagellum; three preapical flagellomeres are pale. The wings are broadly oval, and the forefemur is narrow with brown dots on the posterior surface. On the male genitalia, the sternite IX is pentagonal in ventral view, with rounded posterolateral lobes. The gonocoxite IX is long, gently sigmoid, with posterior apex straight, thickened, set with elongate preapical process on inner surface and seven or eight short processes on outer surface, arranged in a single plane. Additionally, the ventral part of the gonocoxites XI forms covering with two lateral concavities; such covering is ventrally produced as a straight, trapezoidal process.
This species is named after David E. Bowles, a great person, aquatic entomologist and neuropterologist, who has steadily helped AAC in their studies in Neotropical Neuropterida.
Measurements. Male (n = 1). Forewing length: 10.4 mm; Hind wing length: 8.3 mm.
Coloration
(Fig.
Morphology (Fig.
Male genitalia
(Fig.
Ecuador (Guayas).
This species is solely from its type locality in Guayas, Ecuador. It is remarkably similar to P. hagenella in the general coloration pattern and overall morphology, although the shape of the male gonocoxites IX and the gonostyli X rapidly differentiate both species. Another important difference is the shape of the ventral part of the median lobe of the gonocoxites XI.
Plega dactylota
Rehn, 1982: 423. Holotype: male, USA, Arizona (
Plega dactylota lipanica
Rehn, 1982: 423. Holotype: male, USA, Texas (
Plega fratercula
Rehn, 1939: 247. Holotype: male, USA, Arizona (
Holotype of Plega fratercula
USA • ♂; Arizona, Capitain Mt.; 8 Aug. 1933; R. Anderson leg.;
Paratype of Plega fratercula.
USA • 1 ♀; Arizona, Capitan Mt.; 24 Jul. 1933; R. Anderson leg.; “Allotype Plega fratercula Rehn”;
Paratypes of Plega dactylota.
USA • 1 ♂; Arizona, Huachuca Mtn. Carr. Cn.; 06 Aug. 1924; J.O. Martin leg.; “Plega dactylota dactylota Rehn, Paratype”;
Mexico • 1 ♀; Chihuahua, 2 mi W. Matachic; 30 Jul. 1984; D. Thomas leg.; TAMU-ENTOX0285142;
USA – Arizona • 1 ♂ 1 ♀; Cochise Co., S.W. Research Sta., nr. Portal; 1982; L.L. Lampert leg.; black light trap;
This species is distinguished by the long antennae with discoidal basal flagellomeres. On the pronotum, often lateral pale bands are present. The forefemur posterior surface is either nearly completely brown or pale with two pairs of brown areas near the medial region. On the male genitalia, the gonocoxite IX is sinusoidal, with posterior apex equipped with 4–15 closely adpressed, twisted, digitiform processes. The median lobe of the gonocoxites XI has the dorsal part as an expanded arch with a less sclerotized medial area from which a ventrally incised process protrudes; the ventral part consists of a ventral, curved trapezoidal process. On the female genitalia, the gonocoxites + gonapophyses VIII medial part is keel-shaped, formed by to oval, medially joined plates; such a keel forms a posteromedial curved, bilobed process. The bursa copulatrix is short, conical, and moderately sclerotized. The spermatheca is thin, with the proximal section forming several coils; the medial section is a long spiral; the distal section is slightly expanded forming a D-shaped diverticulum.
Measurements. Male (n = 8). Forewing length: 7.3–12.84 mm; Hind wing length: 5.4–9.78 mm. Female (n = 12): Forewing length: 10.7–19.5 mm; Hind wing length: 8.7–15.3 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Mexico (Chihuahua, Coahuila, Guerrero, Jalisco, Morelos, Nayarit, Nuevo León, Puebla, Querétaro, Sinaloa, Sonora, Tamaulipas), USA (Arizona, Colorado?, New Mexico, Texas, Utah).
This species is widely distributed in Mexico and the United States. The records from Coahuila and Sinaloa were extracted from
This species is highly variable in coloration, size, and morphology.
Mexico, Oaxaca: Distrito de Ixtlán, Ixtlán de Juárez, Universidad de la Sierra Juárez, 17°18'55"N, 96°28'57.8"W, 1956 m, 20.X.2017, J.A. Casasola leg., “sobre pared de habitación”.
Holotype
male, pinned. Original label: “Mexico, Oaxaca, Distrito de Ixtlán, Ixtlán de Juárez, Universidad de la Sierra Juárez, 17°18'55"N, 96°28'57.8"W, 1956 m, 20.X.2017, J.A. Casasola, “sobre pared de habitación”,
Mexico • 1 ♀; Oaxaca, Distrito de Ixtlán, Ixtlán de Juárez, Universidad de la Sierra Juárez; 17°18'55"N, 96°28'57.8"W; 1956 m; 16 Jan. 2017; J.A. Casasola leg.; “sobre pared de habitación”;
This species is distinguished because of the presence of a brown flagellum with single preapical paler flagellomere, and the supra-antennal region slightly raised and covered with reclined setae. Furthermore, the antennal flagellomeres are as long as wide on the proximal part of the flagellum. The area adjacent to the posterior and wing margin presents intermittent amber infuscations, yet a pale area at wing tips is remarkable. The forefemur posterior surface is pale with brown dots, whereas the anterior surface is nearly completely brown. On the male genitalia, the sternite IX is trapezoidal with lateral indentation, and the gonocoxite IX is filiform, thin, and sinusoid, with the posterior apex pointed and devoid of processes. The gonocoxite XI medial lobe has a posteroventral U-shaped projection, with lateral rounded fins. The female genitalia have the gonapophyses VIII as two medially joined plates forming a tubular structure; posteromedially two lateral, digitiform lobes are present. The bursa copulatrix is long, with area adjacent to genital pore strongly sclerotized; the spermatheca has the distal section slightly expanded, wider than medial and proximal sections, but abruptly narrowed at apex.
Measurements. Male (n = 1). Forewing length: 12 mm; Hind wing length: 9.5 mm. Female (n = 1): Forewing length: 12 mm; Hind wing length: 9.6 mm.
Coloration
(Fig.
Morphology (Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
The name of this species comes from the Latin disrumpere which means disrupt(ed), alluding to the discontinuous or disruptive color pattern of the wings of this species. An adjective in the nominative case.
Mexico (Oaxaca).
This species is known only from the type locality in Oaxaca, Mexico. This is an anomalous species which was recovered in a clade with P. yucatanae, P. hagenella, and other South American species. The genitalia of both sexes of this species are remarkable, as the male has the gonocoxites XI similar to those of Trichoscelia, and the gonocoxites IX are devoid of processes. The female spermatheca is markedly different to other species of the genus, the gonocoxites + gonapophyses VIII have certain similarities with other South America species, for instance P. zikani.
Mexico, Jalisco: Sierra Manatlán, Puerto Los Mazos, 1500 m, 3–7 Aug. 1992, Luz Hg, Encinar, D. Curoe leg.
Holotype
male, pinned. Original label: “Mexico, Jalisco, Sierra Manatlán, Puerto Los Mazos, 1500 m, 3–7 Aug. 1992, D. Curoe leg., Luz Hg, Encinar”,
Mexico • 1 ?; Jalisco, Sierra Manatlán, Puerto Los Mazos; 1500 m; 3–7 Aug. 1992; D. Curoe leg.; Luz Hg, Encinar;
This species presents the antennal flagellum completely brown, and the basal flagellomeres are as wide as long. The supra-antennal region is raised and the area adjacent to frontal sutures is sunken; both areas are covered with reclined setae. The forefemur has the posterior surface pale with discontinuous brown spots on proximal and distal ½; the anterior surface is mainly dark brown with small pale areas. The forewing has the membrane on apex of intracubital area strongly marked. On the male genitalia, the sternite IX is U-shaped; the gonocoxite IX is short, strongly laterally arched, set with three digitiform, apical processes, subequal in shape and size, two of which are dorsally situated; the surface of such processes has scattered, minute spinules. The gonostyli X are anteriorly recurved, forming a loop before protruding from the abdomen. On the female terminalia, the sternum VII appears as two medially joined, trapezoidal plates (gonocoxites VII), with fusion line discernible. The gonapophyses VIII medial part consist of two medially joined, trapezoidal plates forming a keel, which is set with posteromedial bilobed process. The proximal section of the spermatheca is spiral-shaped, thin, with intermittent swellings.
The specific epithet of this species is a composite of the mantispid genus name Drepanicus and –oides, used to refer to a species that resembles other species. This name alludes to the superficial resemblance of this new species with some South American species of Drepanicinae.
Measurements. Male (n = 4). Forewing length: 11.9–15.4 mm; Hind wing length: 9.9–13.2 mm.
Coloration
(Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Mexico (Chihuahua, Jalisco).
This species is known only from the Mexican states of Chihuahua and Jalisco, being probably endemic to the country. This is an anomalous species recovered as intermediary between a clade containing species from South and Mesoamerica and the rest of the species of Plega. In the male genitalia, the morphology of the male gonocoxites XI is similar to that of Trichoscelia. Furthermore, this species is the unique species of Symphrasinae that has the female gonocoxites VII as paired lateral plates as expressed by several species of Anchieta and Rhachiberothinae.
Plega duckei
Penny, 1982a: 424. Holotype: male, Brazil, Amazonas (
Holotype. Brazil • 1 ♂; Amazonas, Reserva Ducke; 11 Oct. 1977; J.R. Arias leg.;
Brazil • 1 ♀; Amazonas, Manaus; 22 Oct. 1991; G.A.R. Melo leg.; DZUP. • 1 ♀; same data as for preceding; 17 Aug. 1991; DZUP.
This species has most of the vertexal region, frons, and clypeus brown. The antennal flagellum has subconical flagellomeres, those of proximal ¼ are 3× as wide as long, and the palpimacula is grooved. The forefemur is remarkably narrow. On the male genitalia, the sternite IX is U-shaped; the gonocoxite IX is short, thin, and straight, with posterior apex set with three or four short processes. The gonostyli X are ribbon-shaped, short, and recurved at middle before protruding from the abdomen.
Measurements. Male (n = 1). Forewing length: 8.5 mm; Hind wing length: 6.2 mm.
Coloration
(Fig.
Morphology (Fig.
Male genitalia
(Fig.
Brazil (Amazonas).
This species was described from Amazonas, Brazil and is known solely from its type locality and adjacent areas (
Mexico, Baja California Sur: San Venancio, 08 Oct. 1941; Rosa & Bohart leg.
Holotype
male, pinned. Original label: “Mexico, Baja California Sur, San Venancio, 08 Oct. 1941, Rosa & Bohart leg.”,
Mexico – Baja California • 1 ♂; San Lorenzo [Island]; N. Banks leg.;
This species has the antennal flagellum with basal flagellomeres 2× as long as wide. The posterior surface of the forefemur has three brown weakly to well defined spots on the medial area; the anterior surface is dark brown, except the base pale. On the male genitalia, the gonocoxite IX is long, thin, and sigmoid, with posterior apex thickened, laterally curved, and set with 8–13 processes arranged in a group. The ventral part of the median lobe of the gonocoxites XI forms a prominent, curved process with trapezoidal apex, whose surface is rugose and has microspinulae. On the female genitalia, the medial part of the gonapophyses VIII is boat-shaped, lacking processes.
The specific name of this species comes from the Latin flammare, which means flame. This alludes to the flame-shaped apex of the male gonocoxite IX of this species.
Measurements. Male (n = 7). Forewing length: 11.3–14.6 mm; Hind wing length: 8.8–11.3 mm. Female (n = 2): Forewing length: 13.2–14.2 mm; Hind wing length: 10.4–11.3 mm.
Coloration (Fig.
Morphology
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
Mexico (Baja California, Baja California Sur).
This species is restricted to the Peninsula of Baja California (Baja California and Baja California Sur) in Mexico. It was recovered within the clade of species of Central Mexico and the Nearctic as sister to Plega signata to which it is noticeably similar. The anterior surface of forefemur with pale base and the numerous short processes on the male gonocoxites IX arranged as a flame easily separate P. flammata from P. signata. The females may be difficult to separate as there are no clear differences on the genitalia, however they can be separated based on the general color pattern.
Plega fumosa
Linsley & MacSwain, 1955: 16. Holotype: male, Mexico, Michoacán (
Holotype. Mexico • ♂; Michoacán, 11 mi E. Apatzingán; 20 Aug. 1954; E.G. Linsley, J.W. MacSwain, R.F. Smith leg.; Type No. 7202;